THE DIFFERENT FORMS OF FLOWERS ON PLANTS OF THE SAME SPECIES, DARWIN

THE DIFFERENT FORMS OF FLOWERS ON PLANTS OF THE SAME SPECIES By CHAKLES DAEWIN, M.A., F.E.S WITH ILLUSTRATIONS D NEW YORK: APPLETON AND COMPANY, 549 AND 551 BROADWAY 1877 TO PEOFESSOR ASA GRAY £^ts ©'olunu is gtbuatfir BY THE AUTHOR AS A SMALL TRIBUTE OF EESPECT AND AFFECTION 4:58(H)2 Digitized by the Internet Archive in 2010 with funding from University of Illinois Urbana-Champaign http://www.archive.org/details/differentformsof01darw — CONTENTS Introduction Page 1-13 CHAPTEE I Heterostyled Dimorphic Plants : Primulace^ — Primula veris or the Cowslip Dift'erences iu structure between the two forms Their degrees of fertility when legitimately and ille- — gitimately united — P elatior, vulgaris, Sinensis, auricula, &c Summary on the fertility of the heterostyled species of Primula Homostyled species of Primula —Hottonia palustris — Androsacc 14-54 Vitalliana CHAPTER 11 Hybrid Primulas The Oxlip a hybrid naturally produced between Primula veris and vulgaris two —The diflerences in pareiit-specics — Effects structure and function between the of crossing long-styled and short- styled Oxlijis with one another and with the two forms of both — Character of the offspring from Oxiips in a state of nature — Primula and elatior shown to be a distinct species — Hybrids between other heterostyled species of Primula — Sui)plementary note on sponartificially parent-si^jcies self-fertilised cross-fertilised taneously produced hybrids iti the genus Verbascum 55-80 " — CONTENTS VI CHAPTER III Hetebostyled Dimorphic Plants Liuum — continued grandiflorum, long-st}led form utterly sterile with own-form jx)llen —Linura perenne, torsion of —Homostyled furm alcne nalis, singular difference the pistils in the long-styled in Linum — Pulmonaria officibetween the English Pulmonaria angustifolia shown species of self-fertility and German long-styled plants — to be a distinct si^ecies, long-styled form completely self-sterile — Various —Mitchella repens, Houstonia— Faramea, remarkable Polygonum fagopyrum RubiacefE other heterostyled genera fertility of the flowers in pairs difference in the pollen-grains two forms; torsion of the stamens in the short-styled form alone; development not as yet perfect The heterostyled structure in the several Rubiaceous genera not due to descent in of the — common Page 81-136 CHAPTER IV Heterostyled Trimorphic Plants — — — Lythrum salicaria Description of the three forms Their power and complex manner of fertilising one another Eighteen different — Mid-styled form eminently feminine in nature likewise trimorphic — L thymifolia dimor— Lythrum phic — L hyssopifolia homostyled — Nesaja verticillata trimorphic — Lagerstroemia, nature doubtful— Oxalis, trimorphic species of — Valdiviana— Eegnelli, the illegitimate unions quite barren— speciosa — sensitiva—Homostyled species of Oxalis unions possible Gra^fteri — Pontederia, the one monocotyledonous genus known to include 137-187 heterostyled species CHAPTER V Illegitimate Offspring of Heterostyled Plants Illegitimate ofifspring from all three forms of Their dwarfed stature and sterility, Lythrum salicaria some utterly barren, some — CONTENTS fertile — Oxalis, Vll transmission of form to the legitimate and il- legitimate seedlings— Primula Sinensis, illegitimate offspring in some degree dwarfed and infertile Sinensis, auricula, farinosa, variety, illegitimate and — Equal-styled varieties of P —P vulgaris, red-flowered elatior seedlings sterile — P veris, illegitimate plants raised during several successive generations, their dwarfed stature and sterility —Equal-styled remarks —Close parallelism between illegitimate and Page 188-243 fertilisation hybridism CHAPTER —Trans—Concluding varieties of P veris mission of form by Pulmonaria and Polygonum VI Concluding Eemarks on Heteeostyled Plants The essential character of heterostyled plants — Summary of the between legitimately and illegitimately Diameter of the pollen-grains, size of anthers fertilised plants and structure of stigma in the different forms Affinities of the genera which include heterostyled sj^ecies Nature of the advantages derived from heterostylism Tlie means by which Transmission of form Equalplants became heterostyled Final remarks 244-277 styled varieties of heterostyled plants differences in fertility — — — — — — — CHAPTEE VII Polygamous, Diojcious, and Gyno-dicecious Plants Tlie conversion in various ways of hermaphrodite into dioecious —Kubiaceaj —Heterostyled plants rendered Verbenaceaj— Polygamous and sub-dioecious plants— Euonymus — Fragaria— The two sub-forms of both sexes of Ehamnus and Epigaia — Ilex — Gyno-dioecious plants — Thymus, difference of the hermaphrodite and female individuals — Satnreia —Manner which the two forms i)robably originated—Scabiosa the of the and other gyno-dicecious plants— Difference dioecious plants in fertility in in corolla in the forms of plants size polygamous, dioecious, and gyno-dioccious 278-309 — vm CONTENTS CHAPTER VIII Cleistogamic Floweks General character of cleistoganiic flowers— List of tlie genera producing such flowers, and their distribution in the vegetable —Viola, description of the cleistogamic flowers in the their compared with that of the perfect three forms of flowers— Oxalis acetosella— O gamic flowers— Vandellia—Ononis — Impatiens—Drosera— Miscellaneous observations on various other cleistogamic plants Anemophilous producing cleistogamic flowers—Leersia, perfect flowers rarely developed —Summary and concluding cleistogamic flowers — The remarks on the origin conseries several species fertility ; sensitiva, cleisto- species of ciiicf which may be drawn from the observations in this Pao;e 310-345 volume clusions Index 34C-352 THE DIFFERENT FORMS OF FLOWERS ON PLANTS OF THE SAME SPECIES INTEODUCTION The namely the difby certain kinds of plants, either on the same stock or on distinct stocks, ought to have been treated by a professed botanist, to which distinction lean lay no claim As far as subject of the present volume, ferently formed flowers normally produced the sexual relations of flowers are concerned, Linnseus long ago divided them into hermaphrodite, monoecious, dioecious, and polygamous species This fundamental distinction, with the aid of several subdivisions in each of the four classes, will serve classification is artificial, my purpose ; but the and the groups often pass into one another The hermaphrodite sub-groups, namely, class contains heterostyled two interesting and cleistogamic but there are several other less important subdivisions, presently to be given, in which flowers differing in various ways from one another are proplants; duced by the same Some species plants were described by me in a scries of papers read before the * " Ou the Two Forms or Di- morphic Condition in the Species several years ago, Linnean Society,* of Primula, and on their reuiarkable Sexual Relations." ' Journal : INTRODUCTION the individuals of which exist under two or three the length of their pistils and stamens and in other respects They were called by me dimorphic and trimorphic, but have since been forms, differing in named by Hildebrand, heterostyled.* As I many still unpublished observations with respect these plants, it has seemed to me advisable to re- better have to publish my former papers in a connected and cor- rected form, together with the new matter It will shown that these heterostyled plants are adapted be for so that the two or three forms, hermaphrodites, are related to one another almost like the males and-females of ordinary I will also give a full abstract of unisexual animals reciprocal fertilisation though all ; are such observations as have been published since the appearance of my papers but only those cases will be noticed, with respect to which the evidence seems fairly Some plants- have been supposed to be satisfactory heterostyled merely from their pistils and stamens varying greatly in length, and I have been myself more than once thus deceived With some species the ; of the Proceedings of the Linaean Society,' vol vi 1862, p 77 " On tlie Existence of Two Forms, and on tlieir Reciprocal Scxual Relation, in several Species of the Genus Linum." Ibid vol vii " 18G3, p 69 On tlic Sexual Tliree Relations of the Forms of 2/ //^/trwjn saZtcar/a.' Ibid vol viii 1864, p 169 " Oil theCliaracter and Hybriillike Nature of the Ofl'spring from the Illegitimate Unions of Dinwr- phic and Trimorphic Plants." Ibid vol X 1868, p 393 " On the Specific Differences between Primula veris, Brit Fl (var officinalis, Linn.), P vulgar!.^, Brit Fl (var acaulis, Linn.), and Jacq and on tlio Hybrid Nature of the Common Oxllp With Supplementary Romarks on Naturally Produced Hy- P elatior, ; brids in the Genus Vorl)abcum." Ibid vol x 18G8, p 437 * The term "heterostyled "does not express all the difterences beforms; but tliis is a in rnany cases the term has been adopted by tween failure As tlie common writers in various countries, I am unwilling to change it for that of heterogone or heterogonmis, though this has been proposed by so high an authority as Prof Asa Gray see the 'American Naturalist,' Jan 1877, p 42 ; ( 346 ) INDEX ACANTHACE^ Bentham, Mr., on tho cleistogamic flowers of Ononis, 326 A Acanthacex, 313 Acer campestre, 12, 308 Acloxa, jJSgiphila elata, 123 mollis, 123 ohdurata, 124, 286 Alefuld, Dr., on Linum, 100 Alisma natans, 311 Amphicarpxa, 327 Amsinckia spectabilis, 110; bility in varia- length of stamens and ; 2G1, 266 pistil, Boraginew, 101 Boreau on eowslip and primrose, 57 Borreria, 127 Bouche' on Pavonia, 313; effect of temperature and light on corolla, 342 Bouvardia leiantha, 135 Braun on Dracocephalum, 299 Breitenbach, \V., on Primula elatior, 34, 272 Bromfield, Dr., on jirimrose and ct)wslip, 57 Primula elatior, 73 Specularia perfoliuta, 330 Brown, Eobert, on sexual changes, 282 Buckwheat, the common, 111 Anchusa arvensis, 111 Androsace vitalliana, 53 Anthers, size of, in ditierent forms, 252 contabescent, 283 Arachis, 312 Arnebia hispidissima, 111 Ascherson, Dr., on Salvia cleistogaina, 313, 340 Juncus biifonius, 332 Leersia orijzoides, 335 ; ; ; Asclepias, 337 Ash, the common, 11 Asperula scoparia, 285 Axell on Primula stricta, 50 Caltha pahistris, 13 Campanula colorata, 330 Cardamine amara, 307 Caspary, Prof., on llhamnus cathar- 294 313 Chamissoa, 292 Cinchona viicrantha, 134 Cleistogamic flowers, 310; list of genera, 312; on tlieir origin, 343 Cnicus acaulis, 307 palustris, 307 pulleu-grains Coccocypgclum, 133 of, 250 Coprosma, 285 Cordia, 117 pistil of, 253 ticus, Cnttleya, B Biil)ington, Prof., on Primula elalior, 72 Stellaria graminea, 313 Baillon, emission of the tubes from ; 337 on ray-florets, A W., on Impatiens fulva, l>ollen-graius, I3elhf)mnie, M., JioniK^tt, 327 tlie ; flowers fertilised whilst in bud state, 342 Bentliam, Mr., on the differentiation of the sexes, 11 ; ; difference in size in the sexes of the same species, 307- Corolla, 309 ; ; INDEX 347 COBYDALIS 146 Corijdalis, Corylus avt'Uayia, 10 Cowslip, the common, 14 ; shortnnd long-styled, 19-22, 56-71 Cratojeylon formosum, 123 Crocker, C W., on Plantago lanceolata, Epigxa repens, 297 Eranthemum amhigutim, 329 Erioplwrum anguxti/olinm, 307 Erytliroxylum, 121 pollen-grains ; Euphrasia 306 Cryptosfachtjs, 313, 332 of, 250 Euomjmus Europxus, 287-293 officinalis, Euryale, 311 Cuphea purpurea, 168 F Faramca, 128 pollen-grains of, 129 Fitzgerald, Mr., on Thelymitra, 313 Forsythia suspensa, 117; stamens, ; Darwin, Charles, on reproductive organs under cultivation, 7; intercrossed plants, 30 prepotency of pollen, G2 insects fertilising flowers, 79 Cejihnlanthera granEpklendron and Catdiflora, 9S tleya, 313; number of poUcngruins, 338 W., on Pulmonaria angustifolia, 105, 107 Datura arhorea, 251 Delpino, plants fertilised by the wind, 10; on the walnut, 10; Fohjgomicex, 114; pollen-grains, 250 Thymus serpyllum, 299 ; ; ; ; 252 viridissima, 117 Fragaria Chiloensis, 293 elatior, 293 vesca, 293 Virginiana, 293 Fruxiaus excelsior, • , ; or cleistogamic flowers, 311, 337 ; Viola odurata, 317 ch'sed Dianthus bitrhatus, G Galium cruciatum, 2S6 Gartner on the sterility of unions between distinct species, 29 Primida vulgaris and veris, 58, 59 hybrid Verhascums, 76, 77, 80 jirepotency of pollen, 241 variation in the sexual powers of plants, 267 contabesceut anthers, 193, 283 ; ; 30 ; Dickie, Dr., on Erioyhoruiii angu»tifoli'im, 307 Dictamnus fraxinclla, 146 JJiudia, 135 Dioecious and sub-ditecious plants, 287 Discospermum, 286 Duubleday, H., on Frimula elatior, 73 Dracocephalum Moldavicum, 299 Drosera Anglica, 329, 342 rotundifoUa, 328 Duval-Jouve, M., on Cryptonlachys, 313; Leersia oryzoides, 333, 334 Dyer, ThLselton, on Salvia Ilorminum, Cratoxylon formosum, 123 ; E Echium vulgare, Epidendron, 313 HI, 305, 307 ; Gentianex, 115 Geraniacae, 169 Geranium sylvaticum., 308 Gesneria penduUna, 261 Gilia aggregata, 118 coronopifolia, 119 micrantha, 119 —— • nudiraulis, pulchella, 118 Gillibert on Menyanthes, 311 Gloriosa Lilj% tlie, 146 Godron on iiybrid Primulas, 55 Gray, Prof Asa, proposes the term heterogone or heterogonous, ; on Linum, 101 ; Leucosmia Bur1 neitiana and acuminata, 114 ; Forsythia su^penm, 117; Gilia G coronopifolia, pulchella, 118; ;; ; INDEX 348 GYNO-DIOSCIOCS 119; I'Moz snlmlata, 119; Mitlieterostyleil chella repens, 125 Coprosma, 285 Euplants, 24-1 ottymus, 287 Bliamnus lanceoEpigxa repens, hiius, 295, 29G 297 Ilex opaca, 298 PUintago media, 31 17; Oxybaphus and Nyrtaginia, 313; Impatlensfulva, 328; Leersia, 334 cleistogamic flowers, 342 Gyno-dioecious plants, 298 ; ; ; ; ; ; ; ; on Nepeta glechoma, 301 Hautbois Strawberry, the, 293 Hedyotis, 133 Henslow, Rev Prof., on hybrid Primxdx, 61 Henslow, Kev G., on flowers selffertilised during the winter, 343 Herbert, Dr., on hybrid Primulx, becoming Uordeum, 333 plants l)hrodite sexual, 283 ; uni- Homostyled species of Primula, 49 Hooker, Dr., on Campanula, 330 Hordeum, 332 Hottunia inflata, 53, 313 relative fertility, palustris, 50 52; anthers of, 252; papilla} on stigma, 254 Houstouia ccerulea, 132, 254 Hoy a carnosa, 331 Hybrid Primulas, 55-71 Hydrangea, 6, HypericincK, 123 Hyssojms officinalis, 299 ; Hai't, Mr., 61 Heterostyled plants, illegitimate offspring of, 188-243; essential character of, 244 sunnnary of the ; differences of fertility between legrtimately and illegitimately fertilised plants, 246; diameter of pollen-grains, 249 ; size of anthers, struetm-e of stigma, 252; list of genera, 255 ; advantages derived from Heterostylism, 258 means by which plants became hetero; 260 transmission of form, equal-styled varieties, 272 final remarks, 275 dimorphic i)lanta, 14-54, 81- styled, 268 ; ; 136 Hex aqnifuUum, 297 opaca, 298 Illegitimate offspring of heterostyled plants, 188; Lythrum salicaria, dwarfed stature and sterility, 192 Oxalis, transmission of form to seedlings, 212 Primula Sinensis, in some degree dwarfed, 215; ; 218-223; 224 ti'ansmission of form and colour, 225 seedlings, 227; P veris, 228; dwarfed stature and sterility, 229-234; equal-styled varieties, parallelism between 234, 238 illegitimate fertilisation and hybridism, 242 Hlecebrum, 311 Impatiens, pollen-grains of, 338 halsamina, 328 fulva, 327 noli-me-tangere, 328 equal-styled varieties, Primula vulgaris, ; ; ; trimor])luc plants, 137-187 Jlihhcus, pollen-grains, 338 Hildebrand, Prof., introduces the word " heterostyled," on the ray-florets of the ComponUe, 5, Primula Sinensi.", 38, 40-43, 192, 217 Linum grandijlorum, 86, 87 L perenne, 92 Pulmonaria officinalis, 101-103, 107, 239; P azurea, 110; Polygonum fagopijrum 111; Oxalis, 169, 171-174, 178, 182, 211-213, 322; henna; ; ; J Juglans regia, 10 Juncus bujfonius, 332, 343 Jussieu, A de, on Malpighiacew, 331 E ; Kerner, Prof., on ray-florets, Atiricula, 43 hybrid forms of Prtmuia, on use of hairs within 55, 73 ; ; ; ;; M9 INDEX MENYANTHE8 128; size of corolla in male flowers, 309 use of glauds as a protectiiin to flowers, HlU Kiik Dr on Moiwchoria vatjiualis, 331 Kuoxia, 135 Koch on Primula longijlora, 50 Kraschenviihou'ia, 312 Kulm Dr., on cleistogamic flowers, 3, 310, 311; list of plants produciji^ diftVrently formed seeds, heterostyled plants, 244 Vantlie corolla, ; , ; dtllia ; nummular iJulia, 324; sessiflora, V Linum Austriacum, 97 catharticum, 100 corymhi/erum, 100 flavum, 81, 98 stamens, 252 ; grand ifliirum, 81 various experiments, 87-89, 96 pistils and stamens, 253, 254 sterile with its own-form pollen, 264, 266 •, ; ; Leicisii, 101 perenne, styles, 95 ; torsion of the 90 long-styled form, 97 ; stigma, 247 100 trigynum, 100 salsoloides, 325 usitatisK/inum, 100 Lipostoma, 134 Lagerstrcemia Indica, 1G7 parvijiora, lUS ' reninx, 1G8 Lathyrus nissoUa, 326, 342 Leooq, H., on the common maple, 12; cowslips and primroses, 57; 72 Idnum AusLythrum hyssopi98 gvnofolia, 1G6; llhamnus, 2d6 diceeiuus jjlants, 209 Scubioaa gticcisa, 305 Viola odorata, 317 Leersia oryzoides, 333-335 pollengrains of, 338 Leggett, Mr., Poutederia cordata, Primula elatior, Iriacum, ; ; ; ; ; Lysimachia vulgaris, 4, 342 Grxfferi, 165 Lythrum hyssopi/olia, 166 salicaria, 116, 137; power of mutual fertilisation between the three forms, 149-157; summary of results, 157-165; illegitimato offspring from the tliree forms, 191-203 concluding remarks on, 203-211; mid-styled form, 241, seeds, 248 257, 258, 280 ; ; thym/folia, 165 ; 187 Legitimate unions, summary on the fertility of the two, compared with that of the two illegitimate in Primula, 46-49 fertility of, compared with illegitimate, 246 Lcighton, Kev W A., on the cowslip and primrose, 56; Verhascum virgatum, 78 Leoiitodon, pollen-grains, 338 Li' idosiphon, 119 Lt'ucosmia acuminata, 114 Burnettiana, 14 stigma, 253 Lily, the Gloriosa, 146 Liiiinaidhemum Indicum, 116; pollen-grains, 250 aiithers, 252 lAnaria spuria, 325 Lindley on Fragaria elatior, 293 Linnajus on Primula veris, vulgaris, and elatior, 56 Linum anguslifolium 100 ; ; ; M M(dpighiacex, 331 Manettia hiculor, 135 Maple, the common, 12 Marshall, W., on Prirrmla elatior, 73 Phmtago lanceolata, 306 blasters Dr Maxwell, on cleistogamic flowers, Maximowicz on Krascheninihowia, 312 Meehan, Mr., on Mitchella, 285 Epigsea repens, 297 Melissa clinipodium, 299 officinalis, 299 Mello, Correa de, on Amrlii>','M'l; Voandzeia 327 Mentha aquatica, 209 hirsuta, 298 vulgaris, 298 Mcnyauthes, 311 trIfoUuta, 115 ; ;; 350 ; INDEX P0LEM0NIACE2B Blichalet on Oxnlis acetosella, 321 Linaria spuria, 325 Milchella, 285 repens, 125 Mohl, H von, on the common cowslip, 14 size of corolla in the eexcd of Ihe same species, 307, 308; TrifoUum and Arachis, 312 clcifitoL^amic flowers, 314, 342 (Knilin aretoscUa, 321 Impatiens noli-me-tangere, 320 ; Specularia perfuliata, 330 ; ; ; Ohleidandia, 132 Oleacese, Ononis columnx, 325 minutissima, 326, 341 parviflora, 326 vulgare, 298 ; ; pollen-grains, 338 Monochoria vaginalis, 330 Mulberry, the, 10 Miiller, 1)., on Viola canina, 314 Miiller, Fritz, on pollen of the Villarsia, 116; Faramea, 128-130; Fosoqueria fra/jrans, 131 Nesxa, ; 167 Oxalis, 180, 181 Pontederia, 183-185; Oxalis Fegnelli, 212; Chamis^oa, 292 Jliilier, H., on the frequency of ; by insects to the Umhelliferx and Composita', on dichogamy, 10 on Antlmpliora and FomLylius sucking the cowslip, 22 Primula elatior, 32 P villosa, visits ; ; ; Origanum Oxalis acetosella, 181, 182 pistil of, 2iJl ; cleistogamic flowers, 321 Mollia Icpidota, 168 speciosa, 168 Monnier, M., on Viola, 318 ; 117 Oliver, Prof., on ovules of Primula veris 17 ; Viola, 318 ; Campanula Cdhirata, 330 ; 49; Hottonia palustris, 51 table of relative fertility of, 52, 53 PolyLinuin catlmrticum, 100 gonum fagopyrum, 113; Lythrum mlicaria, 145 on the origin of lietero.stylism, 263; on the Lahintx 299, 304 Thymus serpyUum, 300; Scahiosa arvensis, 305 Plantago laneeolata, 306; size of corolla in the two sexes of the same Impatiens balsaspecies, 308; 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