Darwin spent over a year intensely studying orchids, conducting experiments and dissecting flowers. He examined both British and foreign orchid species. Darwin was fascinated by the complex adaptations between orchids and their insect pollinators. He saw the orchid's structures as evidence of multiple modifications through natural selection. Darwin published his findings in 1862 in his book On the Various Contrivances by which British and Foreign Orchids are Fertilised by Insects.

1. Darwin’s Sexy Orchids:
Case Study On The Origin
of Species by Means of
Natural Selection
Harvey Brenneise, Associate Dean for Research
Services
University of Southern Mississippi (Hattiesburg) &
Research Associate,
Rancho Santa Ana Botanic Garden (CA)

2. Charles Robert Darwin (1809-82)
 Childhood and youth
 Education
 Voyage of the Beagle
 Marriage
 Move to the country
 Experiments

3. Life (cont.)
 Publications
 Scientific friends
 Publication of Origin of Species
 Publication of Orchids
 Life after Orchids

4. Darwin as a child

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

6. University of Edinburgh (1825-27)

7. Cambridge (1828-31)

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

9. Darwin as a young man

10. Orchis morio L. (1753)

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

12. Capt. Robert FitzRoy (1805-65)

15. Darwin’s room on board

16. Voyage of the Beagle, 1831-36

17. Orchidaceae taxa collected:
by date of collection
 Epidendrum difforme Jacq. [1760] (Brazil,
May/June 1832)
 Codonorchis lessonii Lindl. [1840] (Tierra del
Fuego, Jan./Feb. 1833)
 Chloraea gaudichaudii Brongn. [1834]
(Argentina, Dec. 1833 or Jan. 1834)
 Chloraea magellanica Hook. f. [1846] (Chile,
Jan. 30, 1834) Isolectotype
 Bipinnula fimbriata I.M. Johnst. [1929] (Chile,
Aug. 1834)
 Epidendrum spicatum Hook. f. [1847]
(Galápagos, Oct. 1835) Holotype [Endemic]

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

20. Epidendrum difforme (May/June 1832)

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

30. Darwin’s field notes

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

39. Jan. 30, 1834

44. Bipinnula fimbriata (Aug. 1834)

48. Orchidaceae in the Galápagos
Wiggins and Porter, Flora of the Galápagos Islands (1971)
 Cranichis schlimii Rchb. f. (1876)
 Epidendrum spicatum Hook. f. (1847) Endemic
 Erythrodes weberiana Garay (1970) Endemic
(Isabella)
 Govenia utriculata (Sw.) Lindl. (1839)
 Habenaria alata Hook. (1826)
 Habenaria monorrhiza (Sw.) Rchb. f. (1885)
 Ionopsis utriculariodes (Sw) Lindl. (1826)
 Liparis nervosa (Thunb.) Lindl. (1830)
 Ponthieva maculata Lindl. (1845)
 Presscottia oligantha (Sw.) Lindl. 1840
 Tropidia polystachya (Sw.) Ames (1908)

49. Epidendrum spicatum (Oct. 1835)

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

54. What next?
 First love, Fanny Owens, had become
engaged while on the voyage.
 What to do? The practical thing.

55. Reasons to get married
 Children (if it please God)
 Constant companion (& friend in old age)
who will feel interested in one, object to be
beloved & played with, better than a dog
anyhow
 Home, & someone to take care of house
 Charms of music & female chit-chat
 These things good for one's health
 Forced to visit & receive relations but
terrible loss of time. —

56. Reasons not to stay single
 My God, it is intolerable to think of spending
ones whole life, like a neuter bee, working,
working, & nothing after all. — No, no won't do.
 Imagine living all one's day solitarily in smoky
dirty London House
 Only picture to yourself a nice soft wife on a sofa
with good fire, & books & music perhaps —
Compare this vision with the dingy reality of Grt.
Marlbro' St.
 No children, (no second life), no one to care for
one in old age.— What is the use of working 'in'
without sympathy from near & dear friends—who
are near & dear friends to the old, except
relatives

57. Reasons not to get married
 Freedom to go where one liked — choice of
Society & little of it
 Conversation of clever men at clubs
 Not forced to visit relatives, & to bend in every
trifle
 to have the expense & anxiety of children —
perhaps quarelling
 Loss of time. — cannot read in the Evenings
 Fatness & idleness — Anxiety & responsibility
 Less money for books &c — if many children
forced to gain one's bread. — (But then it is very
bad for ones health to work too much)
 Perhaps my wife wont like London; then the
sentence is banishment & degradation into
indolent, idle fool —

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

59. When? Jan. 29, 1839
Then how should I manage all my business if I were
obliged to go every day walking with one’s my wife. —
Eheu!! I never should know French, — or see the
Continent — or go to America, or go up in a Balloon, or
take solitary trip in Wales — poor slave. — you will be
worse than a negro — And then horrid poverty, (without
one's wife was better than an angel & had money) —
Never mind my boy — Cheer up — One cannot live
this solitary life, with groggy old age, friendless & cold, &
childless staring one in ones face, already beginning to
wrinkle. — Never mind, trust to chance —keep a sharp
look out — There is many a happy slave —

60. Emma Wedgwood Darwin
(1808-96)

61. Emma a paragon of virtue
 Loyal
 Cared for him when he was sick
 Read to him at night
 Fundamentalist religious views
 Little sense of humor
 Angelic!

63. Move to Kent (Down House)--1842

68. Darwin’s scientific “buddies”

69. Joseph Dalton Hooker (1817-1911)

71. Darwin, Hooker, Charles Lyell

72. Asa Gray (1810-88)

75. Alfred Russel Wallace (1823-1913)

78. Experiments (“little triumphs”)
 “Are you going to beat Dr. Hooker?”
 Experiment Book (1856-)
 Weed Garden (experimental theorist)
 Gardeners’ Chronicle
 “I am like a gambler, & love a wild experiment.”
 Field studies and practical scientific
investigations both an amusement and part of
his research program

79. Books--to Origin
 The zoology of the voyage of H.M.S.
Beagle (1838-43)
 The narrative of the voyages of H.M.
Ships Adventure and Beagle [Journal of
Researches] (3 vols.) (1839, 1845, 1860,
1880, 1890, 1905)
 Geology of the voyage of the Beagle,
(1842-46,1876-76, 1889-90)

80. Books—to Origin (cont.)
 A monograph of the sub-class Cirripedia,
with figures of all the species (2 vols.)
(1851-54)
 A monograph on the fossil Lepadidae, or,
pedunculated cirripedes of Great Britain .
(2 vols.) (1851-54)
 On the origin of species (1859, 1860 [1st
American], 1860, 1861, 1866, 1869, 1872,
1876)

83. What next?
 Variation among domestic plants and
animals?
 Drosera (sundew)?
 Dimorphism among primulas?

84. Carl Linnaeus (Carl von Linné) in 1746 had
portrayed a flower as a marriage bed of 9
gentlemen and 1 lady.

85. Christian Konrad Sprengel (1750-1816), Das
entdeckte Geheimnis der Natur im Bau und in der
Befruchtung der Blumen [floral ecology]

86. Orchids!
 Summer 1860: “Orchis Bank”
 Appeal to Gardeners’ Chronicle and replies
 1861: All rest of year Orchid Book (diary)
 To Hooker: "I am intensely interested on subject,
just as at a game of chess.” In September, he
"dissected with the greatest interest.”
 "The contrivances for insect fertilisation in
Orchids are multiform & truly wonderful &
beautiful.”

87. Darwin’s periodical articles on orchids
prior to publication of Orchids
 “Fertilisation of British orchids by insect agency.”
Gardeners' Chronicle no. 23 (9 June 1860): 528
and no. 6 (9 Feb. 1861): 122.
 “Orchids, Fertilization of.” Gardeners' Chronicle
no. 37 (14 Sept. 1861): 831.
 “On the three remarkable sexual forms of
Catasetum tridentatum, an orchid in the
possession of the Linnean Society.” [Read 3
April] Proceedings of the Linnean Society of
London. Botany 6: 151-157. 1862

88. Wrote to Asa Gray prior to publication: “It
really seems to me incredibly monstrous to look
at an orchid as created as we now see it. Every
part reveals modification on modification.”
Publication was delayed because of illness,
but Darwin looked at it as “a hobby-horse, which
has given me great pleasure to ride.”

90. British orchids examined by Darwin (2nd ed.)
[15 genera by Darwin’s count]
illustrated in bold
 Aceras anthropophora, A. longibracteata [France]
 Caladenia dimorpha
 Cephalanthera ensifolia, C. grandiflora
 (left out Cypripedium)
 Epipactus latifolia , E., microphylla, E. palustris , E.
purpurata, E. rubiginosa, E. viridiflora
 Epipogium gmelini
 Goodyera discolor, G. pubescens, G. repens
 Gymnadenia albida, G. conopsea, G. odoratissima, G.
tridentata
 Habenaria bifolia, H. chlorantha
 Herminium monorchis
 Listera cordata, L. ovata
 Malaxis paludosa

91. More “British” orchids
 Neotinea intacta [Italy]
 Neottia nidus-avis
 Nigritella angustifolia [alpine]
 Ophrys apifera, O. arachnites , O. aranifera, O.
muscifera , O. scolopas
 Orchis fusca, O. hircina, O. latifolia, O. maculata, O.
mascula , O. morio, O. pyramidalis , O. ustulata
 Peristylus viridis
 Platanthera chlorantha, P. dilatata, P. flava, P. hookeri,
P. hyperborea
 Ptorostylis longifolia [Australia]
 Pogonia ophioglossoides
 Serapias cordigera [France]
 Spiranthes australis, S. autumnalis , S. cernua, S.
gracilis

92. British specimens (1st ed.) donated by:
 Mr. Bateman, Goodyera discolor (foreign)
 Dr. Battersby (Torquay), Spiranthesis autumnalis
 F. Bond (South Kent) Hadena dentina, H. plusia; Orchis
fusca, moths with pollinia attached
 Rev. G. Gordon (Elgin) Goodyera repens
 Mr. Malden (South Kent) Orchis fusca
 A. G. More (Bembridge, Isle of Wight), Epipactis
palustris, Ophrys apifera (sent report of field work),
Spiranthesis autumnalis
 G. Chichester Oxenden (Broome Park, South Kent)
Neottia nidus-avis, Ophyris aranifera, Orchis ustulata
 Mr. Wallis (Hartfield, Sussex) Malaxis paludosa
 Professor Westwood (bees with pollinia attached)

93. Orchis mascula L. (1755)

96. O. Mascula pollinium

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

98. Bilder ur Nordens Flora (1905)

102. Ophrys apifera Huds. (1762)

104. Flora von Deutchland, Osterreich under der Schweiz (1885)

107. Atlas des plantes de France (1891)

112. “Foreign” orchids examined by Darwin
2nd ed. Illustrated in bold
 Acropera loddigesii, A. luteola
 Aerides cornutum, A. odorata, A. virens
 Angraecum distichum, A. eburneum, A.
sesquipedale
 Bulbophyllum barbigerum, B. cocinum, B.
cupreum, B. rhizophorae
 Bonatea speciosa
 Brassia
 Calanthe dominii, C. masuca, C. veratrifolia, C.
vestita
 Catasetum callosum, C. luridum, C. mentosum,
C. planicips, C. saccatum, C. tabulare, C.
tridentatum

113. More “foreign” orchids
 Cattleya crispa
 Chysis
 Coelogyne cristata
 Coryanthes fieldingii, C. macrantha, C.
speciosa , C. triloba
 Cycnoches egertonianum, C. ventricosum
 Cymbidium giganteum
 Cypripedium acaule, C. barbatum, C.
calceolus, C. candidum, C. pubescens, C.
purpuratum
 Dendrobium bigbbum, D. cretaceum, D.
chrysanthemum , D. formosum, D. speciosum
 Disa cornuta, D. grandiflora, D. macrantha

114. And more!
 Epidendrum cochleatum, E. floribundum, E.
glaucum
 Eulophia viridis
 Evelyna carivata
 Galeandra funkii
 Gongora atro-purpurea, G. maculata, G.
truncata
 Laelia cinnabarina
 Leptotes
 Lycaste skinnerii
 Masdevallia fenestrata

115. More!
 Maxillaria ornithorhyncha
 Microstylus rhedii
 Miltonia clowesii
 Monachanthus viridis
 Mormodes ignea, M. luxata
 Myanthus barbatus
 Ornithocephalus
 Phaius grandifolius
 Phalaenopsis amabilis, P. grandiflora
 Pleurothallis ligulata, P. prolifera

116. That’s all, folks!
 Rodriguezia secunda, R. suaveolens
 Sarcanthus parishii, S. teretifolius
 Selenipedium palmifolium
 Sobralia macrantha
 Stanhopea devoniensis, S. oculata
 Stelis racemiflora
 Thelmitra carnea, T. longiflora
 Vanilla aromatica
 Zygopetalum mackai

117. “Foreign” specimens donated for 1st ed. by: [the
kindness of many friends and strangers], 43
“exotic” genera “well dispersed through the
subfamilies of the vast Orchidean series”
 Joseph Hooker, “has never become weary of
sending me specimens from the Royal Gardens
at Kew”
 James Veitch, jun., “many beautiful Orchids”
 R. Parker, “extremely valuable series of forms”
 Lady Dorothy Nevill, “most kindly placed her
magnificent collection of Orchids at my disposal”

118. Donors (cont.)
 Mr. Rucker (West Hill, Wandsworth), “sent me
repeatedly large spikes of Catasetum, a
Mormodes of extreme value to me, and some
Dendrobiums”
 Mr. Bateman, “a number of interesting forms,
including the wonderful Angraecum
sesquipedale”
 Mr. Turnbull (Down), “free use of his hot-houses”
and “giving me some interesting orchids, and his
gardener, Mr Horwood, for his aid in some of my
observations”
 Dr. Lindley, “fresh and dried specimens”

119. Lady Dorothy Nevill (1826-
1913)

120. Relationship with Darwin
 Liked to put signed portraits of famous
scientists on her walls
 Flirtatious correspondence “filled with
double entendre, describing the orchids'
body parts and the methods by which
these flowers fertilize each other."

121. Wisdom from Lady Dorothy Nevill:
“The real art of conversation is
not only to say the right thing at
the right place, but to leave
unsaid the wrong thing at the
tempting moment."

122. John Lindley (1799-1865)

123. Jas. Bateman (1811-97)

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

125. Catasetum saccatum Lindl. (1840)

126. Darwin called Catasetum “the most
remarkable of all Orchids", and showed
how in these flowers "as throughout
nature, pre-existing structures and
capacities [had been] utilised for new
purposes".
Catasetum tridendatum showed its
“truly marvelous” mechanism, by which it
shot out a pollinium at any insect touching
a part of the flower with “sticky gland
always foremost.”

127. Darwin imitated the action of an
insect touching the flower’s “antenna”
using a whalebone spring.
"I touched the antennæ of C.
callosum whilst holding the flower at
about a yard's distance from the
window, and the pollinium hit the pane
of glass, and adhered to the smooth
vertical surface by its adhesive disc.”

129. Sertum Orchidaceum (1838)

130. Catasetum experiments
 Fall onto a table from a height of 2-3 in.
 Cut off with a crash with scissors
 Deep pricks of the column and stigmatic
chamber
 A blow hard enough to knock off the anther (an
accident)
 Press hard on pedicel and rostellum
 Nothing works except “violence” to the antennae
(not including stream of air, cold water or human
hair)

132. Curtis’’ v. 131 (1905) as C. christyanum

136. Catasetum macrocarpum [=tridentatum ]:
3 species in 3 genera or one taxon?
Robert Hermann Schomburgk’s
problem: 3 genera on the same plant!
Lindley’s response: “Such cases shake
to the foundation all our ideas of the
stability of genera and species.”

137. Robert Hermann Schomburgk

138. Catasetum macrocarpum Rich. ex Kunth (1822)
IPNI
 Monachanthus viridis Lindl. (1832)
=Catasetum trifidum Hook. (1833)
 Monachanthus viridis Schomb. (1831)
=Catasetum barbatum Lindl. (1844)
 Monachanthus viridis Lindl. (1836 publ.
1835) =C. macrocarpum FEMALE
 C. tridentatum Hook. (1823) = C.
Macrocarpum MALE
 C. tridentatum, var. viridiflorum (1834)
FEMALE?
 Myanthus barbatus Lindl. (1836 publ. 1835)
=C. barbatum BISEXUAL?

139. Catasetum macrocarpum Rich. ex Kunth (1822)

140. Curtis v. 52 (1825) as C. tridentatum

141. Curtis v. 61 (1834) as C. tridentatum

143. Edwards’ v. 21 (1836) as Monachanthus viridis

145. Angraecum sesquipedale Thou. (1822)

146. Thouars. Histoire particulière des plantes orchidées
recueillies sur les trois îles australes d'Afrique (Paris,
1822)

147. Orchid Album

149. Box arrived from Bateman on 25 January
1862
To Hooker: “I have just received such a Box
full from Mr Bateman with the Astounding
Angraecum sesquipedalian with a nectary a
foot long—Good Heavens what insect can
suck it”?

150. Experiments
 Bristles and needles
 Cylinder to which the pollinia attached
themselves
 Conjectured that there was a moth with a
long proboscis that could get the nectar
from the bottom of the nectary

151. “The astonishing length of the nectary may
have been acquired by successive modifications.
As certain moths of Madagascar became larger
through natural selection . . . those individual
plants of the Angræcum which had the longest
nectaries . . . and which, consequently, compelled
the moths to insert their proboscises up to the very
base, would be fertilised.
“These plants would yield most seed, and
the seedlings would generally inherit longer
nectaries; and so it would be in successive
generations of the plant and moth. Thus it would
appear that there has been a race in gaining
length between the nectary of the Angræcum and

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

154. John Day scrapbook 039_452_632

155. John Day scrapbook 040-632-44

156. Xanthopan [Macrosila] morgani Walker (1856)
subsp. praedicta Rothschild & Jordan (1903)

160. Reception to Orchids
 Slow sales with general public
 Botanists generally favorable (dialog began that
resulted in expanded 2nd ed.)
 To Lyell: “Entomologists are enough to keep
[evolution] back for half a century.”
 George Campbell, Duke of Argyll, The reign of
law
 Wallace, Creation by law [measured Macrosila
[Xanthopan] morgani in the British Museum
(from South Central Africa) and found the
proboscis to be 7 ½ inches long.]

161. Present entomologists ask:
“What, then, pollinates A.
longicalcar, with a nectary 10
cm. longer than A.
sesquipedale?”

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

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

166. Fritz Müller

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

168. "In my examination of Orchids, hardly any
fact has so much struck me as the endless
diversity of structure ,—the prodigality of
resources,—for gaining the very same end ,
namely, the fertilisation of one flower by the pollen
of another.”
I “found the study of orchids eminently useful
in showing me how nearly all parts of the
flower are coadapted for fertilisation by
insects, & therefore the result of n.
selection ,—even most trifling details of structure.”

169. Books—from Orchids on
 On the various contrivances by which
British and foreign orchids are fertilised by
insects (1862, 1877, 1882).
 On the movements and habits of climbing
plants (1865, 1875, 1876, 1882).
 The variation of animals and plants under
domestication (2 vols.) (1868, 1878).
 The descent of man, and selection in
relation to sex (1871, 1874, 1882).
 The expression of the emotions in man
and animals (1872, 1873, 1890).
 Insectivorous plants (1875, 1888).

170. Books—from Orchids on (cont.)
 The effects of cross and self fertilisation in the
vegetable kingdom (1876, 1877, 1878).
– “A complement to the 'Fertilisation of Orchids,' because it shows
how important are the results of cross-fertilisation which are
ensured by the mechanisms described in that book.”
 The different forms of flowers on plants of the
same species (1877, 1884).
 The power of movement in plants (1880, 1881).
 The formation of vegetable mould, through the
action of worms (1881, 1882).
 The life and letters of Charles Darwin, including
an autobiographical chapter (1887).

171. Articles on orchids post publication
of 1st ed. of Orchids
 “Fertilisation of Orchids.” Journal of Horticulture
(31 March 1863): 237.
 “Appearance of a plant in a singular place.”
Gardeners' Chronicle no. 33 (15 Aug. 1863): 773
[Epipactis latifolia].
 “Fertilisation of Cypripediums.” Gardeners'
Chronicle no. 14 (6 April 1867): 350.
 “Notes on the fertilization of orchids. Annals and
Magazine of Natural History (Ser. 4) 4 (Sept.
1869): 141-159.

172. Darwin’s life after Orchids

173. Darwin’s contributions to botany
 Understanding of and ability to
demonstrate that the flower is a product of
evolution
 Complex ecological relationships resulted
in the coevolution of orchids and insects
 Pollination research and reproductive
ecology (floral ecology)

174. Greenhouse extension project
(1862-63)
 Asked Hooker for plants: "I long to stock it,
just like a school-boy.”
 Sent his butler with a cart to Kew to pick
up 160 plants
 Apologized for depleting the “national
collection”
 To Hooker: "You cannot imagine what
pleasure your plants give me ... Henrietta
& I go & gloat over them."

185. Select Bibliography
 Allan, Mae. Darwin and his flowers: the key to natural
selection (New York: Taplinger, 1977)
 The complete works of Charles Darwin online (
http://darwin-online.org.uk/)
 Darwin Correspondence Project (
http://www.darwinproject.ac.uk/)
 Darwin’s garden: an evolutionary adventure (New York:
New York Botanical Garden, 2008).
 Pollan, Michael. “Love & lies {orchids}.” National
Geographic 216:3 (Sept. 2009), p. 100-121.
 Porter, Duncan M. (various journal articles)
 The works of Charles Darwin (New York: New York
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 Yam, Tim Wing, et al. “’The orchids have been a
splendid sport’—an alternative look at Charles Darwin’s
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186. Questions and comments
 Harvey.Brenneise@gmail.com