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111THE WATERLILIES
CARNEGIE INSTITUTION OF WASHINGTON. WATERLILIES, PLATE 1.NYMPH/EA GIGANTEA VIOLACEA.
THE WATERLILIESA MONOGRAPH OF THE GENUSNYMPHAEABYHENRY S. CONARDSenior Harrison Fellow in BotanyUniversityof PennsylvaniaPublished byThe Carnegie Institution of Washington1905
miciCARNEGIE INSTITUTION OF WASHINGTONPublication No. 4*ZU Barb Qgafttmor* (JptietTHE FRIEDENWALD COMPANYKAI.T1MORI. MD.. U.S. A.
?^PREFACE.This work is the outcome of four years' study on the HarrisonFoundation in the University of Pennsylvania.It would have beendifficult or impossible without the generous support given by the Provostand the Board of Trustees. In the University Botanic Garden overthirty species and varieties of waterlily were cultivated, and severalhybrids originated. These and other collections of living plants furnishedexcellent material for the present review of the group.Nothing like a complete synopsis of the Waterlilies has hithertobeen put before the English-speaking world. DeCandolle made a specialstudy of the genus Nymphaea for his Prodromus, and both Lehmannand Planchon gave much attention to the genus from 1850 to 1853.Beginning in 1855 and continuing until 1888, a number of exhaustivetreatises by Robert Caspary went far toward perfecting our knowledgeof these plants. Indeed, had the learned doctor of Konigsbergassembled his vast knowledge into one connected whole, the presentwork would be needed chiefly as a translation. Since his day, however,of water-great strides have been made in the culture and hybridizationlilies, with the result that they now appear in almost every collectionof plants. Already by crossing and recrossing and the use of horticulturalnames, serious confusion exists concerningthe identities of theparent species. It has therefore seemed important to bring togetherthe knowledge of the genus in all of its botanical relations and in itsbearings on human life and history. In the present attempt there maybe many omissions, in spite of the utmost efforts in collating the material,but the paper may serve to show how incomplete is our knowledge ofany one group of plants, and it is hoped that it may make the way ofthe future investigator easier.In the adoption of the generic name Nymphaea, the writer regretsthat he is at variance with a number of American botanists for whomhe has the utmost respect;but a glance at the synonymy of Nymphaea,in
ivPreface.page 125, shows the leading facts. It seemed patent to the writer thatall of the general, and several special, arguments against a change ofname are particularly cogentin the present example. The new code,however, proposed by a committee of the Botanical Club of the AmericanAssociation for the Advancement of Science, seems to sanction the nomenclaturehere chosen. 1 In the use of specific names, authorities have beenomitted wherever possible, since the names adopted in Chapter V arefollowed. Citations of literature are based on the bibliography on pages243 to 263.Throughout the time occupied by the preparation of this work, I havebeen deeply indebted to Professor John M. Macfarlane, to whom I desireto express my gratitude.To Mr. William Tricker, formerly of Riverton, N. J.,and through himto the Henry A. Dreer Co., of Philadelphia,I am under obligationfor the privilege of studying their magnificent collection of waterlilies.Acknowledgment is also due to Dr. J.W. Harshberger, of the Universityof Pennsylvania, and Dr. J. N. Rose, of the National Herbarium,for suggestions and herbarium material ;to Mr. A. J. Mela, of Helsingfors,Finland ;J. J. Soar, of Little River, Fla.; Wm. Gollan, Saharanpur, India ;Wm. Fawcett, of Kingston, Jamaica R. M.; Gray, gardener to O. Ames,North Easton, Mass. ;Gopal Chandradatta, Calcutta, India ;Sr. Albertothe directorLofgren, director of the Botanical Garden of Sao Paulo, Brazil ;of the Royal Botanic Garden of Tokyo, Japan to the Gray Herbarium,;Boston Natural History Society, Massachusetts Horticultural Society,Columbia University;to the Academy of Natural Sciences of Philadelphiafor library facilities; and to John B. Lieberg, Joseph H. Painter, Chas. C.Abbott, Prof. Conway MacMillan, Albert J. Edmunds, E. D. Sturtevant,and C. G. Pringle for various courtesies. Mr. Frederick V. Coville andMr. J.C. Bay, of Washington, D. C, have also given helpful aid andcriticism while the matter was going through the press.In order to give the greatest possible accuracy to the taxonomicportion of the monograph, I was enabled by the trustees of the CarnegieInstitution of Washington to examine important type material in severalEuropean herbaria, viz, in Kew, British Museum, Linnaean Society,Museum d' Histoire Naturelle, the herbaria of De Candolle, Delessert, andBarbey-Boissier, and the herbaria of Munich, Berlin, and Copenhagen. Theresults of this trip were very valuable, thanks to the courteous assistance'Bulletin Torr. Bot. Club, 31: 249-290. May, 1904.
Preface.vof those in charge of the various collections. The principal changesresulting in the manuscript were in the addition of a few species.That the book appearsin its present form is also due to the CarnegieInstitution, on account of their acceptance of the manuscript for publication.Were it not for this privilege, much of the illustrative material must havebeen omitted. These many obligations I have felt keenly, and havetried in a measure to meet by conscientious care in the execution ofevery detail.University of Pennsylvania, HENRY o. CONARD,Philadelphia, December i, 1904. Senior Harrison Fellow.
VICONTENTS.PrefacePAGEiii-vDescription of Plates ix-xiList of Text Figuresxii-xiiiI. History 3Oriental Literature 3Pre-Linnaean Literature 10II. Structure 27The Root 28The Stem 37The Leaf 54
xDESCRIPTION OF PLATES.Plate i. Frontispiece. Nytnphaea gigantea violacea. From a painting made by Mrs. E. Rowan,in the Cape York Peninsula, Australia.Plate 2. Nytnphaea odorata. 1-7, development of leaf and stipules; 1, 2, front and back views ofthe same rudiment; 3, older; 4, 5, front and side views of one rudiment; 6, 7, frcntand back views; 8, rhizome with roots and leaves trimmed off, natural size; 9, transversesection of rhizome, natural size. A. Koronski del.Plate 3. Nymphaea gigantea. Under side of leaf and details of flower. 1, anterior sepal ; 2, outermoststamen ; 3, inner stamens ; 4, median petal ; 5, vertical section of ovary ; 6, outermostpetal. Photograph from plant grown at University of Pennsylvania.Plate 4. Nymphaea elegans. Flower and under side of leaf. From plant grown at University ofPennsylvania stock of the Dreer Co. A. Koronski del.;Plate 5. Nymphaea ampla, from Caspary, 1878, pi. 28. 1, flower from hb. St. Petersburg, coll. B.Jaeger, No. 233, Hayti, with 149 stamens, 24 carpels ; 2, leaf from hb. Geo. Engelmann,coll. Wright, No. 1857, Cuba; 3, flower, seen from the side; 4, rays of stigma andcarpellary styles; 5, stamens (2-5 from the same specimen) ; 6, two carpels of a flowerin hb. St. Petersburg, coll. Karwinski, 1841, in Colipa River, Mexico.Plate 6.Nymphaea Havo-virens (N. gracilis Hort, not Zucc, 1832). Flower and under side of leaf,Photo, from plant grown at University of Pennsylvania, from stock obtained from theDreer Co.Plate 7. Nymphaeastellata versicolor. Under surface of leaf and details of flower, a, sepal ;b-e, petals of successive whorls; f, stamens; g, closed flower. From plant grown bythe Dreer Co. (Mr. W. Tricker), from seed sent by W. Gollan, Saharanpur, India.Plate 8.Nymphaea caerulea.Pennsylvania.Flower and under side of leaf.From plant grown at University ofPlate 9. Nymphaea micrantho. 1, under side of leaf; 2, upper side with young plant developing;3 caudex, with a branch ; 4, scars of leaves and of a flower, from surface of a caudex,enlarged ; 5, cross section of caudex ; 6, longitudinal section of a caudex ; 7, superficialtangential section through leaf cushions of Fig. 4; /, base of a petiole; f, base of apeduncle; 8, section a little deeper than 7; 9, same as 8, a little deeper. From a drawingin color by R. Caspary, now in hb. Berlin, by kind permission of the Directors.Plate 10. Nymphaea capensis. Under side of leaf. From plant grown at University of Pennsylvania;stock from Dreer Co.Plate ii. Nymphaea capensis zanzibariensis. Flower and leaf. Plant grown at University ofPennsylvania; stock of Dreer Co. A Koronski del.
xDescription of Plates.Plate ia. Nymphaea tulfurea. I, s, leaves viewed from the upper surface; 2, under side of leaf;3, a closed flower; 4, fully open flower; 6, rhirome. From a co-type in hb. BritishMuseum, by courtesy of the Curators. P. Highly del.Plate 13. Nymphaea mexicana. Flower and leaf, natural size. From plants grown at Universityof Pennsylvania, from roots collected in Little River, Fla., by Prof. J. M. Macfarlane.A. Koronski del.Plate 14. Nymphaea tetrogona. Flower and under side of leaf, natural size. From plants grownby the Dreer Co.Plate 15. Nymphaea alba rubra, 1-21 ;N. Candida, 22. Nymphaea alba rubra.I, partly openedflower; 2, fully open flower on its last day; 3-8, petals; 9-12, stamens of successivewhorls from outside inward, viewed from front, side and back; 13, ovary stripped ofsepals, petals and stamens, from the side; 14 stigma and styles from above; 15, sectionof ovary ; 16, pollen, from pole and from side ; 17-19, fruits ; 20, petal ; 31, stigmaticlobes. Nymphaea Candida. 22, flower cut in half. From drawings in color by R. Caspary,now in hb. Berlin, by kind permission of the Directors.Plate 16. Nymphaea lotus. Leaf, flower, transverse and vertical section of fruit. Photographedby P. Highley from a specimen coll. Schweinfurth, Nov., 1887, at Damietta,Egypt. >n hb. British Museum, by courtesy of the Curators.Plate 17. Nymphaea pubescens. Flower and under side of leaf. From a plant grown by the DreerCo.(Mr. Wm. Tricker), from seed sent by Mr. W. Gollan, Saharanpur, India.Plate 18. Nymphaea rubra. 1, flower of second day, between 9 and 9.30 a. m. ; 2, flower at sametime of third day; 3, flower at same time of first day; 4, flower of fourth day, between8.45 and 9 a.m.; 5, sepal from outer and inner sides; 6, petals of first (at left)and second whorls; 7, petal of innermost whorl; 8-10, stamens from outer and innersides; 11, 12, dry pollen; 13-15, pollen germinating in sugar solution; 16, 17, pollen,examined in citron-oil, 18, 19, in water, 20, in dilute sulphuric acid; 21, germinatingpollen; 22, section of carpellary style; 23-26, hairs of ovary, 24 treated with potash;27, pistil seen from above ; 28, ovary of Fig. 3, with stamens ; 29, ovary on second day.From color drawings by R. Caspary, in hb. Berlin, by kind permission of the Directors.Plate 19. Nymphaea amazonum. 1, closed flower after its first night; 2, fully open flower,second night between 4.30 and 4.45 a. m.; 3, the same flower on the first night between4.30 and s a. m. ; 4, sepal, inner side ; 5, petal of outermost whorl, from without;6, 7, 8,9 petals of the next four whorls; 10-13, stamens of successive whorls;14, is, side and section views of ovary after the second night; 16, ovary after thefirst night; 17, transverse section of ovary; 18, 19, pollen, dry and in water; 20,epidermis of ovary. From color drawings by R. Caspary, in hb. Berlin, by kindpermission of the Directors.Plate 20. Nymphaea rudgeana. 1, flower on its first night, 10.15 P- rn. ' 2, the same between9.20 and 9.40 p. m. ; 3, 4, ripe fruits ; 5, closed flower after first night ; 6, flower with3 outer whorls of floral leaves removed; 7, sepal from within; 8, petal of outermostwhorl; 9, petal of second whorl; 10, petal of fourth whorl; 11, stamen of outermostwhorl of four; 12, stamen of second whorl of four; 13, stamen of third whorl ofseven or eight; 14, 15, inner and innermost stamens; 16-19, views of ovary; 20, 21,22, pollen in dilute sulphuric acid, dry, and in water; 23, ripe fruit with floral leavesremoved; 24, transverse section of ovary; 25-27, sections of carpellary style at apex,just below the club-shaped apex, and at base ; 28, papilla; of stigma. From colordrawings by R. Caspary, in hb. Berlin, by kind permission of the Directors.
Description of Plates.XIPlate 21. Nymphaea blanda, N. blanda fenzliana. N. tenerinervia, N. stenaspidota. From Caspary,1878, pi. 36. 1-7, N. tenerinervia, from a specimen coll. Martius at Joazeiro, Provinceof Bahia, Brazil. 1, flower; 2, lower surface of leaf; 3, apex of outer stamen; 4, ofinnermost stamen ; 5, carpellary style ; 6, upper surface of leaf, near the point of insertionof the petiole, showing series of raised dot 1 ; and short raised lines; the arrowpoints to the insertion of the petiole ; 7, outer surface of sepal showing short crowdedraised lines ; 8, outer surface of sepal of N. stenaspidota with raised dots between therather long raised lines; 9-12, N. blanda fenzliana, specimen from San Juan tie Nicaragua,in hb. Vienna. 9, young fruit; 10, 11, outer and median stamens, from hb. Bentham;12, leaf; 13-16, N. blanda. 13, flower viewed from side, coll. Parker in BritishGuiana; 14, leaf of the same; 15, part of upper surface of same leaf, showing raphid-likelines and raised dots; 16, leaf of Meyer's type, coll. Dr. Rodscheid in the EssequiboRiver (from hb. Grisebach).Plate 22. Nymphaea lasiophylla and N. rudgeana, from Caspary, 1878, pi. 34. 1-7, N. lasiophylla.I, 2, flower from face and back; 3, leaf, showing lower side, with distant lobes; 4, partof a larger leaf seen from above, lobes touching ; 5, 6, outermost and innermost stamens;7, part of lower surface of leaf marked with raised dots and branching lines; 8-19,N. rudgeana. 8, leaf of a plant from Jamaica, cult, in Berlin Botanical Garden; 9, 10,II, stipules of three leaves of a Jamaica plant, cult, in Konigsberg Botanical Garden;12, fruit, raised at Konigsberg ; 13, another fruit, raised at Konigsberg, with persistentsepals and petals; 14, Konigsberg fruit in median section; 15, small flower, coll. Poeppig,No. 3033, 1832, in the Island of Colares of the Amazon, in hb. Berlin; 16, sepal ofa flower, coll. Sello at Victoria, Brazil; 17, 18, stamens of this flower; 19, apices ofmedian and innermost stamens showing a narrow rim of the connective, but no appendage.Plate 23. Nymphaea jamesoniana, from Caspary, 1878, pi. 32. 1, 2, flower from front and back, inhb. Boissier; s, sepal; 3, 4, 5, innermost, outer and outermost stamens; 6, 7, leaves,from hb. St. Petersburg and hb. Lehm; 8, part of leaf from beneath, showing forkedbranching lines and raphid-like idioblasts.Plate 24. Nymphaea stenaspidota and N. gardncriana, from Caspary, 1878, pi. 33. 1-3, AT stenaspidota.1, whole plant from type of hb. Lehm ; s, sepal ; 2, flower ; 3, part of leaf, uppersurface, showing raised dots, the arrow points toward point of insertion of petiole;4-n, N. gardneriana. 4, flower, from hb. Vienna; s, sepal; 5, 6, 7, upper part ofoutermost, median and innermost stamens; 8, 9, 10, leaves; 11, upper surface of leafshowing series of raised dots, with raphid-like lines interspersed, the arrow as in Fig. 3.Plate 25. Nymphaea oxypetala, from Caspary, 1878, pi. 31. I, 2, flower, from hb. Boissier, fromface and back ; 3, 4, outermost and innermost stamens ; 5, three carpellary styles, withsurface of stigma and axile process; 6, 7, leaves from hb. Lehm. and St. Petersburg;8, globose idioblasts of inner face of anther at the point of Fig. 3.Plate 26. Nymphaea Pennsylvania = N. caerulea X zanzibariensis. From the original plant grownat University of Pennsylvania.A. Koronski del.Plate 27. Nymphaea capensis X zanzibariensis. Flower and under side of leaf. From plants raisedat University of Pennsylvania.Plate 28. Nymphaea elegans X zanzibariensis. Flower and under side of leaf. From plants raisedat University of Pennsylvania.Plate 29. Nymphaea Havo-vircns X zanzibariensis. Flower and under side of leaf. From plantsraised and bred at University of Pennsylvania.Plate 30. Nymphaea omarana, hybrid. Stock from Dreer Co. A. Koronski del.
LIST OF TEXT FIGURES.Tail-piece. Guest at a feast, with lotus (after Wilkinson) iFigure I. Nefer Turn (after Wilkinson) 62. Egyptian lotus (designs after Wilkinson) 73. Pleasure boat in a pond of lotuses (after Wilkinson) 7"4. Faba Aegyptia." Facsimile from Matthiolus 15"5. Nymphaea alba." Facsimile from Lobelius 156. Longitudinal section of root tip of N. caerulea 297. True and false epidermis of roots 308. Root of N. odorata, transverse section 329. Contractile root of N. Aavo-virens, transverse section 3310. Vascular bundles of the roots of N. odorata 361 1.Development erf air-canals 3712. Diaphragms in the air-canals of roots 3813. Rhizome of N. tuberosa 3914. Tubers 4015. Perennating body of N. mexicana 42i6j Rhizome of N. odorata; transverse section of internode 4317. Microscopic section of rhizome of N. odorata 4418. Caudex of N. caerulea; transverse section 4619. Cork formation at base of root scar 4720. Details of leaf scar 4821. Transverse section of caudex of N. flava SO22. Course of vascular bundles at node of Nymphaea flava 5123. Vascular system of stolon of N. rubra? at a node 5224. Stipules 5525. Collenchyma in petioles 5726. Sections of petioles and peduncles 5927. Types of vascular bundle 6328. Leaf of N. ftava 6529. Epidermis of leaf, N. tetragona 6730. Leaf of N. rubra, vertical section 6831. Arrangement of vascular bundles in veins of leaf 7132. Development of mucilage hair of N. alba 7233. Hairs and hair-bases 7334. First leaves from germinating tubers 7535. Transverse sections of petioles of water-leaves 7636. Floral diagram for Nymphaea 8037. Details of anther and stigma 8438. Pollen 8539. Diagram of ovary (Brachyceras) in longitudinal section 8640. Diagram of ovary in transverse section 8641. The aril 9142. Sections of outer cell-layer of testa (after Weberbauer) 9243. Distribution of vascular bundles from the chalaza (after Chifflot) 93xiiPAGE
List of Text Figures.xiiiFigure 44. Longitudinal section of seed 9445. Development of flower 9946. Ovule of N. odorata from a half-grown bud 10247. Embryo sac of N. odorata 10248. Embryology 10549. Seedlings of a hybrid Nymphaea of the Lotos group 10850. Seedling leaves no51. First broad (submerged) leaf of N. lotus seedling 11252. A strong-growing waterlily in shallow water 1 1953. Nymphaea elegans, details of flower 13254. Nymphaea Aavo-virens, details of flower 13855. Nymphaea caerulea, details of flower and fruit 14256. Nymphaea heudelotii, flower and leaves 14857. Nymphaea heudelotii nana, leaf and flowers 149leaf and flower 15058. Nymphaea ovalifolia,59. Nymphaea calliantha, leaf and flower 15160. Nymphaea copensis, details of flower 15461. Nymphaea capensis zanzibariensis, details of flower 15862. Nymphaea stuhlmannii, leaf and flower 16263. Nymphaea mexicana, details of flower and fruit 16464. Nymphaea mexicana, seedling leaves 165details of flower 16865. Nymphaea tetragona,66. Nymphaea Candida, leaf 17367. Nymphaea odorata, details of flower and leaf 18168. Nymphaea odorata minor, details of flower 18469. Nymphaea odorata gigantea, details of flower 18770. Nymphaea tuberosa, details of flower 18971. Nymphaea tuberosa, tubers and water-leaves 19072. Nymphaea tuberosa, details of leaf 19173. Floral organs of the Lotos group 19374. Early leaves of the Lotos group, from germinating tuber 19575. Nymphaea zenkeri, flower and leaf 19776. Nymphaea dumasii, leaf (after Schenck) 21677. Nymphaea pulcherrima,PAOBdetails of flower 22478. Nymphaea Pennsylvania, details of flower 22579. Nymphaea capensis X zanzibariensis, details of flower 22680. Nymphaea elegans X zanzibariensis, details of flower 22681. Nymphaea " Wm. Stone," details of flower 22782. Nymphaea " Mrs. C. W. Ward',' details of flower 228
."THE WATERLILIESBYHENRY S.CONARD
3CHAPTER I.HISTORICAL.ORIENTAL LITERATURE.Man was probably impressed from the first with the grace and beautyof the waterlilies ;for their distribution is worldwide. There is evidenceenough that the seeds and roots (tubers) of various species have beenused for food by many savage races, as in Australia, Madagascar, WestAfrica and Central America ;but naturallyit remained for more civilizedpeoples to rise to anything like an aesthetic interest. A Chinese writer ofthe eleventh century a. d., Chou Tun-I, is thus quoted in Giles's " Gemsof Chinese Literature,"Since the opening: days of the T'angr dynasty [600 a. d.],it has been fashionableto admire the paeony ;but my favorite is the waterlily. How stainless it risesfrom its slimy bed ! Howmodestly it reposes on the clear pool an emblem ofpurity and truth !Symmetrically perfect, its subtle perfume is wafted far and wide ;while there it rests in spotless state, something to be regarded reverently from adistance, and not to be profaned by familiar approach.In my opinion, the chrysanthemum is the flower of retirement and culture ;thepaeony, the flower of rank and wealth ;the waterlily, the Lady Virtue sans pareille.Alas ! few have loved the chrysanthemum since T'ao Yuan-ming ; and nonenow love the waterlily like myself ;whereas the paeony is a general favorite with allmankind.The waterlily of China is the littleNymphaea tetragona, which,though pure white and of pleasant scent, never attains to much morethan a pigmy size ;and its odor is not nearly so rich as that of ourown N. odorata. It is interesting also to note that our author is expressingonly a personal opinion, one which is not at all general, andwhich indeed may be regarded as heretical.India is much richer ingorgeous varieties of waterlily than anyother portion of the globe. By day the blue N. stellata exhibitsseveral shades of color, even running off by degrees into pink in partsof Bengal. At night and early in the morning the great white N. lotusand the deep crimson N. rubra expand their flowers 6 to 10 inches in3
4 The Waterulies.diameter, and the displayis enriched by every shade of color connectingthe one form with the other. That these have found place in the richliterature of the Hindus is quite certain. But some confusion has arisenon account of the presence and importance of Nelumbo nucifera, thepink-tipped sacred lotus of India. Nelumbo is unquestionably the sacredplant of the Buddhists, the flower upon which Buddha sits, the padmaof the Hindu prayer. The only questionis whether the few statementsassigning a religious value to Nymphaea are to be entirely disregarded,or whether plants of this genus did really receive a minor degree of" " "worship. Lotus-eyed," lotus-hearted and the figure of water on thelotus leaf, all so frequent in Sanskrit literature, certainly refer toNelumbo. 1 But when we read in the Mahabharata (Vana Parva, ch.307, 19) of a hero "possessed of a complexion bright as the coppercoloredlotus leaves," we think at once of N. rubra or its varieties. Anexamination of the Sanskrit, however, 2 reveals that the root padma occurshere (padmatdmradalojjvalam), showing that the plant in the poet's mindwas Nelumbo, and we may translate " "lotus-petals in place of " lotusleaves,"improving the simile. Again in Adi Parva (ch. I, 86)we read"As the Full-Moon with its mild light opens the buds of the waterlily,so does this Purana with the light of Sruti expandthe human intellect."This waterlily isprobably N. lotus or N. rubra, as they are nightbloomers. Kalidasa makes a similar reference in Sakoontala," And nowWhile the round Moon withdraws his looming discBeneath the western sky, the full-blown flowerOf the night-loving: lotus sheds her leavesIn sorrow for his loss, bequeathing naughtBut the sweet memory of her lovelinessTo my bereaved sight:" (Williams, 1856, p. 92.)There is some poetic license here ;for Nelumbo does not open atnight, but does shed its petals when full-blown, while Nymphaea lotusopens at night, but retreats under water after three nights, taking itspetals with it ! It seems, however, that N. lotus is the plant referredto; for Williams adds in a note (p. 14) that "the moon is often calledthe ' lover or lord of the lotuses '."lCf. Dhammapada, ch. iv, 58, 9; ch. xxvi, 401; Miiller 1881. Mahabharata, Adi Parva,ch. 122, 29 ; ch. 199, 11 ;Vana parva, ch. 5, 16 ; ch. 231, 13 ;ch. 307, 23 ; Dutt 1895.1Kindly made for me by my learned friend Albert J. Edmunds of Philadelphia.
jHistorical.Nymphaea stellata is alluded to in the same poem (p. 15) thus"thoughtlessly attemptsTo cleave in twain the hard acacia's stemWith the soft edge of a blue lotus leaf."Graceful and appreciative as are these literary references, and manysuchmight be added, they cannot be regarded as having any religioussignificance. The Sanskrit words utpala and kantala for the blue lotus(N stellata), kumuda the white night lotus (N. lotus) and pundarika whitelotus flower, so far as we know, have no religious associations (Macdowell,Elwell, Lanman). Williams (p. 5, notes) states very pertinently that thelotus " is as favorite a subject of allusion and comparison with Hindupoets as the rose is with Persian."On the other hand, it is stated in an early number of the " BotanicalMagazine" (1805) that when a Hindu once entered the study of SirWilliam Jones where a flower of Nymphaea lotus chanced to lie for examination,he " "made prostrations before it as though this were an act ofpiety. This reverence is further said to " pervade Hindostan, Tibet andNepal." Tratinnick (1822) considers the same species to be the sacredlotus of India, without giving evidence. Pickering (1879) records thatN. stellata is said to be " distinctly figured in the cave-temples of Adjunta,"and that he himself saw itfigured in Braminical cave-temples elsewherein India. Nymphaea mbra appears as the " autumn lotus " in Dhammapada,285, according to Pickering, and he thinks that this species may bethe padma of the Hitopadesa and Vetala panchavinsati Graham I is;quotedas having observed it around Bombay, " in tanks, particularly in theneighborhood of caves or temples." But to the writer, none of theserecords present any certain evidence of the religious importance ofNymphaeas in India. The account in the " Botanical Magazine " is themost suggestive, but isby no means conclusive, and the use of the flowersas decorative designs for temples indicates no more than an appreciationof their beauty. The presence of the plants themselves " especiallynear temples,"if the observation is trustworthy, may be explained bothby the beauty of the flowers and the food-value of seeds and tubers.In the absence, therefore, of any direct evidence that waterlilies areregarded as sacred in India, I believe we have no sufficient reason forattributing this regard to them.
The Waterlilies.EGYPTIAN ARCHEOLOGY.In Egypt the relations of the waterlilies to the social and religiouslife were more extended. But after an impartial examination of the factsavailable, am I strongly of opinion that they received no worship oreven reverence here. The teachings of archaeological research werebrought together long ago by Duppa (1813-1816), and to some extentby Tratinnick (1822), and more recently by Dr. Pleyte (1875), whereasthe critical observations of Schweinfurth from 1882 to 1886 have addedmany interesting facts, and placed many points beyond the reach of doubt.In spite of a complete unanimity among scholars, considerable confusionexists in the popular mind as to the identity of theso-called Sacred Lotusof Egypt. In America, at least, Nelumbo nuciferaiscommonly styledSacred or Egyptian Lotus. But Pickering,Fio. 1.Nefer Turn (after Wilkinson).Pleyte, Joret and Schweinfurth from the botanicalside, and Wilkinson, at least, amongarchaeologists, unite in the opinion that Nelumbois never found on the ancient monuments,and that it was not known inEgyptbefore the advent of the Persians. Not untilthe Roman period did it find a place in Egyptianart ;it does become more or less prominent atthis time. But this comes within the limit ofrecorded history and is discussed in anotherplace.We must understand, therefore, that the " lotus " of the palmy dayswhen Egypt was truly Egyptian was a waterlily native to the Nile Valleyand Delta. Two species are figured on the monuments and tombs,N. lotus, the white night-lotus, and N. caerulea, the blue diurnal one ;of these the latter occurs very much more frequently. Pleytefound thewhite lotus only on a single tomb, belonging to the XII dynasty, about2500 B.C.; 1 this example, however, was plainly distinguishable from theallied species. Schweinfurth (1884 c) found petals of the white lotusalong with those of the blue in the funeral wreaths of Ramses II andAmenhotep but he never observed I, any carvings or pictures of thisplant, though he says Unger found such at Beni-Hassan. The lotusborne on the head of Nefer Turn in one of his manifestations (Wilkinson,1883, 2: 180; Wiedemann, 1897, p. 138) is long and obtuse and suggests'Petrie's dates are followed throughout.
Historical. 7N. lottts rather than N. caerulea, but it is too much conventionalized tooffer any conclusive argument (Fig. i, 6).Since the leaves of the blue species are entire and its petals acute,whereas the white one has sharply dentate leaves and broad petalsrounded at the apex, the difference is usually evident even in very cruderepresentations (Fig. 2).We recognize the blue one easily in manifoldapplications. It occurs, according to Schweinfurth (1883 b),on all theancient monuments of Egypt. The use of its tuberous rhizome for food,is said to have been given to the people by Menes, or perhaps even byIsis (Pickering Diodorus). At convivial meetings a flower of the blue;lotus was presented to each of the guests, and in feasts for the dead, thefeasters delighted themselves with the color and odor of this lotus(Buckley). In a carving figured by Wilkinson, the guests are distinwFig. 2.Egyptian lotus (designsafter Wilkinson).Fig. 3.Pleasure boat in a pond of lotuses (afterWilkinson).guished from the hosts and servants partly by their lotuses (see page 1,underneath title).In most of these picturings the flowers are considerablyconventionalized, but Schweinfurth saw them in the temple of Ramses IIat Abydos, and on coffins of the Ptolemaic period, distinctly painted blue.One of Champollion's figures shows a lotus with green sepals, blue outerpetals and red inner petals, all of the parts being tipped with black the red;and black must have been inserted for love of variety! An interestingTheban picture shows a pleasure boat, being towed round a pond by threeslaves ;the water isrepresented by the characteristic wavy lines, and ismade more vivid by a liberal sprinkling of lotuses, both leaf and flower,over the water-surface 1 (Fig. 3).Just as the blue lotus appears in figures of social life and inrecreation, it occurs as a favorite flower in religious observances. It isfigured among offerings to the gods in the IV dynasty (Pickering ;1This lotus has also been found depicted on the pavement of the palace of Sardanapalus, buthas evidently been copied by the Assyrian artists from Egyptian sources (Bonavia, 1894).
8 The Waterlilies.Pleyte), and stands in front of Osiris at the judgment of the dead(Buckley). "When the Egyptians approached the place of divineworship," Diodorus tells us, " they held the flower of the ' 'agrostis intheir hand, indicating that man had proceeded from a well-watered ormarshy land, and that he required a moist rather than a dry aliment "(Wilkinson, /. 428). This " "agrostis Wilkinson considers to be anothername for the lotus (1.c.J: 350). The god Nefer Turn "was figured asa man crowned with an upspringing lotus flower (Fig. a 1), symbol of theresurrection, and of his power to grant continuous life in the world tocome" (Wiedemann, p. 138). In one manifestation this is distinctly theblue lotus (Fig. 1, a). Ahi, an infant deityis "sometimes represented assitting on the lotus flower," the vase from which the plant grows representinga lake of water (Wilkinson j: 132-3). "Osiris, too, swam on alotus leaf, and Harpocrates was cradled in one " (Whewell). Hapi, or theNile, a god widely known and honored, " was sometimes resolved into theNile of Upper and the Nile of Lower Egypt, the lotus being consideredemblematic of the former, and the papyrus of the latter ;a design consistingof both plants tied together formed a favorite subject for thedecoration of the royal throne, as typifying the king's rule over Upperand Lower Egypt" (Wiedemann, p. 145-6).The use of flowers in funeral decorations seems to have been veryprominent in the XIX to XXI dynasties.The custom was to lay wreathsand semicircles of flowers on the breast of the enwrapped corpse untilsarcophagus was quite packed with these floral tributes. Flowers ofNymphaea caerulea on petioles 18 to 20 inches long were fastenedbetween the bands encircling the mummies of Ramses II and the priestNisboni, scattered singly all over them (Schweinfurth 1883 b).Breastwreathsconsisting mostly of petals and sepals of the same plant,sometimes also with petals of N. lotus, were found in the coffins ofRamses II, Amenhotep Ahmes I, I(1580 b. c), the priest Nisboni, theprincess Nzi-Khonsu, and the mummy of the case marked kent, from theXX and XXI dynasties, found at Deir-el-Bahari (Thebes) in July 1881.These are probably the " Egyptian wreaths " of Pliny and Plutarch, the"lotus garlands" of Athenseus. Most of these plant-remainsthedate fromnearly 2000 b. c, but those of Ramses II were renewed about 1100 b. c. ;for inmoving these sarcophagi at that time into their secret resting-placein the Valley of the Tombs of the Kings, to avoid the marauding hordesfrom the desert, that of Ramses was accidentally broken a new coffin;and new floral decorations were therefore supplied. The wreaths of
Amenhotep are perhaps older than the Trojan war !Historical. 9(Schweinfurth,1883b). In every case the preservation of the vegetable relics was as perfectas in newly-dried herbarium specimens they were easily soaked out in;water, and Schweinfurth was able not only to identify them beyond doubtand to find out the manner in which the wreaths were fastened together,but also to prepare and mount examples for the Boolak Museum, wherehis success is attested by eleven cases of plants. Specimens were alsosent to the British and several continental museums. A wreath of lotuspetals from the coffin of Ramses II was figured in Nature, in 1883(Schweinfurth, 1883 b). Such a wreath consists of foliage leaves ofMimusops schimperi folded in two or in four and stitched together with fibersfrom the date-palm in such a manner as to clamp and hold the lotus petalswithout piercing them the whole was then; strung on strips of leaves ofthe date-palm. The mummy marked rent had small whole flowers ofN. caerulea woven in the wreaths by means of papyrus threads. Sometimes,as in the case of the princess Nzi-Khonsu, the whole body of thelaid inmummy was covered several layers deep with these garlandsconcentric semicircles from the chin downward. It is interesting to notein passing, that Schweinfurth was not able to detect the slightest differencebetween these ancient plant remains and the species now growinginEgypt. Either these species are not suffering transmutations, or thechanges are infinitely slow. Of real importance, however, is the conclusionthat since the species are unchanged, the climate also of Egyptmust be very nearly the same as it was four thousand years ago. If thisbe true, the absence of Nelumbo and Papyrus from modern Egypt cannotbe attributed to meteorological changes, but we are strengthenedin theconclusion that these plants only existed there in ancient times throughartificial, i. e. human, agencies, namely by cultivation.Several additional references to the Egyptian lotus are unquestionablymeant for a species of Nymphaea, but whether the blue or the white,the writer cannot now say. Bes, probably a god adopted from the negrotribes, was, at the end of the New Kingdom (500b. c ?) " fused with . . .and endowed with all the attributes of the young sun-god, and representedlike him as sitting upon a lotus flower" (Wiedemann, p. 167). Plutarchsays the Egyptians " characterize the sun as though it sprang every dayfresh out of the lotus plant" (Wilkinson, 3 : 132-3), and Proclus pretendsthat the lotus was particularly typical of the sun, which itappeared tohonor by the expansion and contraction of its petals (1. c, p. 350).Wilkinson indicates that this lotus isalways N. lotus L. (p. 132-3), but
ioThe Waterlilies.in this he is certainly mistaken for;the one referred to by Proclus isevidently diurnal, and therefore must be N. caerulea. The reader mayalso be warned against the statement of Prichard that " Nymphaea lotusand N. Nelumbo [= Nelutnbo nucifera\ appear plentifully on the sculpturesof Thebes." The unanimous voice of later observers sufficientlynegativesSo itthis.appears that the lotus was enjoyed in many ways, but especiallyas an ornament, a thing of beauty.It was the emblem of the Nile God,for it was the evident product of the river. It was offered to Osiris asany treasured object is consigned to the gods.It was laid upon thebodies of the dead, but other plants could take its place at seasons ofthe year when the lotus was out of bloom. Thus we see no decidedevidence that it was a sacred flower, or an object of worship.It wasthe " favorite flower in the hands of the Egyptians" (Wilkinson, j: 132-3)as the rose or others might be in the hands of any modern people.Itwas the symbol of Nefer Turn and the resurrection just as Lilium longi-with us and is used for decoration atflorum has a similar significanceEaster.pre-linn,ean literature.The facts of the foregoing account are drawn from the researchesof scholars during the past hundred years, and have had no influenceupon the growth of botanical science. The beginnings of our presentknowledge of the waterlilies are to be found in early Greek and Romanliterature, from which a growth can be continuously traced. It is true,however, that classic writers became acquainted with one or two Africanspecies of Nymphaea by traveling in Egypt and noting the customs ofthe people and the plants which surrounded them. For example,Herodotus (Bk. 2, 92) tells us that when the Nile " is full and hasmade the plains like a sea, great numbers of lilies {xpivov),which theEgyptians call lotus {Xmro^j, spring up in the water : these they gatherand dryin the sun, then having pounded the middle of the lotus, whichresembles a poppy, they make bread of it and bake it. The root alsoof this lotus is fit for food, and is tolerably sweet, and is round, and ofthe size of an apple." This evidently refers to a Nymphaea, probablyto N. lotus L. ;for the white-flowered form would be more likely to appealto the writer as a " "lily than the blue N. caerulea. The descriptionapplies equally well to either species ; perhaps, like most other earlywriters, Herodotus did not distinguish between the two, or did not notice
Historical.i ithat there were two varieties. He goes on to mention other " lilies,like roses," with fruit " like a wasp's nest," and edible seeds " of thesize of an olive stone " this is; evidently Nelumbo nucifera. He alsouses the word "lotus" in book iv, par. 177, to designate a tree whosefruit is the food of the Lotophagi (Lotus-eaters) and their neighborsthe Machyles, tribes of Northern Africa. This, the Forgetful Lotus, isa species of Zizyphus.Theophrastus (Bk. 9, ch. 13),in a chapter dealing with various plantsand roots, speaks of some which are " sour, others bitter, others sweet.""The sweet [one] called nymphaea (wfiyaia) is accustomed to grow inpools and marshes, as in the Orchomenian country, and Marathon, andthe island of Crete. The Boeotians call it madonia, and eat the fruit.It bears large leaves upon {supra) the water ;[these] ground and placedon wounds are said to stop blood. It is useful also as a drink forintestinal disorders." This has been generally accepted,I think correctly,to refer to Nymphaea alba, the common white waterlily of Europe.Theophrastus also describes Nelumbo, under the name of xua/xo^.A brief mention of the "lotus " of Egypt, evidently Nymphaea, occursin the Historical Library of Diodorus Siculus, written about the beginningof our era. He says (Bk. I, ch. 3)itgrows in great plenty in the delta ofthe Nile, and is used for bread. He too speaks of Nelumbo, but underthe name" "of xtftmpwv. Celsus gives lotus in a list of therapeuticagents for " cooling the body." The word may not belong to the originalmanuscript, but may have been added in later copies. It is considered tostand for the Egyptian lotus as above.Dioscorides, in his Materia Medica, devotes a chapter (112) in bookiv to " the Egyptian lotus (kwTbt; 6 'at AquTcry j-eww^svoc). which growsin plains flooded by the water " of the "Nile. The stem resembles thatof xua/ioi; [Nelumbo], having a white flower like a lily (xpivov) which,they say, opens at sunrise, and closes at sunset, and the head isaltogether hidden under water, and again emerges at sunrise. The headlike millet,is like a very large poppyin it is seed; (xo/otoc;grand)which the Egyptians dry and beat into bread. It has a root like a quince,which is also eaten both raw and cooked ;when cooked it resembles theyolk of an egg." The identity of this with our Nymphaealotus L. canscarcely be doubted. We find a difficulty in that N lotus is a nightbloomer,and therefore could not possibly be thought by those familiarwith it to sink under water at night. But N caerulea grows with N lotusin Egypt, and is a day-bloomer, closing into a very insignificant bud by
j 2The Waterlilies.evening ; its flowers, however, are not white, but pale blue. We think,however, that N. lotus is the one meant by Dioscorides, as its tubers arelarger than those of N. caerulea, it fruits more freely, and the fruits whenripening are devoid of all the floral envelopes except the bases of the outerstamens, exposing a large poppy-like ovary. The fruit of N. caerulea issurrounded by its four sepals and four alternating sepaloid petals, so thatits ovary, though equally poppy-like, remains hidden. The storyof itsbehavior at night is a traveler's tale, enlarged upon, as we shall see, byPliny, and copied many times afterward. Three other kinds of " lotus "are given in Dioscorides one is the tree of Herodotus' : Lotophagi the;other two are leguminous plants near to or part of our present genusLotus. Nymphaea {wptpaia) appears in book m, chapter 138. Hesays it grows in marshes and standing water has leaves like;xifiwpwv[Nelumbo], but smaller and more oblong, several from the same root,some floating and some submerged flower; white, like a lily (xpivov),with a bulb in (xpoxwdei;)the middle. " But when done blooming, therecomes a round black body of the shape of an apple or a poppy-head,which contains copious, black, crowded seed, of a glutinous taste. Stemsmooth, slender, black, like the stem of xtfiibptov.Root (/!>ia) [Rhizome]black, rough, like a club {p'apaloidrjz) gatheredin autumn." Dried and;taken with wine, it is recommended for dysentery and bowel complaints ;it is also powerfully anaphrodisiac. 1 For all these uses the seeds are alsogood.It is called Nymphaea because " it loves a watery place " it is;found plentifully {ttoMjj)in Elis in the river Anigros, and in Aliartia ofBceotia. This description all together is quite recognizable for Nymphaeaalba; the black ovary is the only discrepancy. There is, continuesDioscorides, another nymphaea [which is called nymphona {yufupCova),and its flower nouphar (voixp />)] with leaves like the former, but the root[rhizome] white and rough, flowers yellow, brilliant, resembling a rose.This can be no other than a species of Nuphar. He describes further, inbook n, two kinds of xiaptoz,the Greek and the ;Egyptian the formerthe fruit of this (whichappears to be a kind of bean, the latter Nelumbo;is used for food) he calls xiftwptovor xtfiioztov,and the root, which is likethat of a cane, isalso eaten both raw and cooked, and is called xoXoxaala.Pliny in several places in his Natural History, uses the word "lotos"for four different plants, the herb lotos, the Egyptian lotos, a shrub lotos(perhaps the pomegranate)and the tree lotos. The fruit of the second of1" Eadem contra veneris insomnia bibitur, quae adimit :quin et aliquot diebus continuo pota,genitale flaccidum et enerve reddit." Edit. Sprengel, /.' 479.
Historical. 13these (Bk. 22, chap. 21) he calls "lotometra" and states that theEgyptian bakers knead the flour of -its seed with milk or water to makebread. He adds, to give the quaint English of Philemon Holland's freetranslation (p. 125)."There is not any bread in the world (by report)more wholesome and lighter than this, so long as it is hot ; but being oncecold, it is harder of digestion and becommeth weightie and ponderous."In book xiii (ch. 17) the " Egyptian lotus " (a Nymphaea)is described inalmost the same words as those of Dioscorides :its occurrence and habitat,itspoppy-like head, and seed resemblingmillet. He tells us that theEgyptians gather the heads, lay them in heaps until they putrefy, thenwash the seeds clean, dry and grind them. Pliny repeats the story of theclosing of the flower at sunset, stating only that it is covered with leavesuntil sunrise ;also that the flower is white. But though he is morereserved in statement here than is Dioscorides, he goes farther in tellingof the behavior of the " Egyptian lotus" in the Euphrates (chap. 18).There, he says, the flower isplunged beneath the water at evening, sodeep that a man cannot reach any part of it with his hand;but aftermidnight it rises slowly, appearing above water again at sunrise. Hetells us in addition that its root is covered with a black rind, and isgoodto eat, especially when boiled or roasted, and that it is excellent forfattening hogs. Nelumbo is mentioned by Pliny in two places under thenames " colocasia," "cyamos" and Egyptian bean {/aba). The descriptionof Nymphaea in Bk. 25, ch. 7, reads much like that of Dioscorides. Afew new features are introduced. The name and the plant,it is said,originated from a nymph who died of jealousy through love of Hercules ;hence the plantis called by some Heraclion but ;by others it is calledrhipsalos, for its club-like rhizome {radix). Itgrowsin water and hasbroad, floating leaves. The other features of the plant, its names, localities,and uses, and the yellow Thessalian variety are exactly as given byDioscorides, and the identities of all these are subject to the same doubtsand assurances as those given in discussing that writer.At that remarkable banquet imagined by Athenaeus, the conversationturned on Alexandria, whereupon one of the feasters tells about the" " lotus which grows in the marshes near that city in summer "; it bearsflowers of two colors, one like that of the rose, and it is the garlandswoven of the flowers of this color which are properly called garlandsof Antinous ;but the other kind is called the lotus garland, being of abluish {xuaUav) color" (Bk. 15, ch. 21). The firstplant is probablyNelumbo, the second Nymphaea caerulea Sav. ;wehave here the first
14 The Waterlilies.mention of Nelumbo under the name of lotus, and the earliest Latinor Greek record of a blue Egyptian lotus. Nelumbo is frequently mentionedelsewhere by Athenaeus, once again under the name of lotus,but more often as xtfttbpiov, xuapoz atyunuoz, or xolozaoiu (Bk. 3, ch. 2).Hesays, at variance with all other writers, that the Egyptians call this plantlotus, but quotes the other names from Nicander, Theophrastus, DiphilustheSiphnian, and Phylarchus.Galenus adds nothing to the facts and theories of his predecessorsconcerning these plants. He recognizes three kinds of lotus (Bk. 7,ch. 11), the herb, the tree and the Egyptian lotus (Nymphaea), andone with black rhizomes and onetwo kinds of Nymphaea (Bk. 7, ch. 13),with white these ; correspond with Dioscorides' descriptions of Nymphaeaalba and Nuphar. He simply mentions Nelumbo, under the name of/aba Aegyptia (Bk. 1, ch. 20).From this time onward we find nothingof note for our scienceuntil it revived with the Revival of Learning. William Turner in 1548gives this quaint note in his catalogue of *' The Names of Herbes :"Nymphea is also named in greeke nymphaia & madonais & is called inenglish water Roses, & some with the Potecaries cal it nenufar. The duch mecal thys herbe Seeblumen. Boeth the kyndes of water Roses growe in standyngwaters."Mr. Britten (cf. Turner, 1 548, p. 1 30) has identified " boeth the kyndes "lutea Sm.as Nymphaea alba L. and NupharThree years later Fuchsius and Hieronymus Bock (Tragus)inGermany publishedtheir botanical classics. The former has a rudewoodcut of the white waterlily with the Latin name Nymphaea Candidaand the Greek Nu/ifaia huxy.On the next page occurs "Nymphaealutea" with a figure which identifies it with Nuphar.Tragus goes into the subject much more extensively and showsThere are two kinds of " Seeblumen,"evidences of personal observation.the white and the yellow, of which the former is commoner. He comparesthe flower of this to a beautiful double {gefult) lilyor rose, with eachpetal shaped like one's thumb ;and in the midst of the flower is agolden yellow sun (ein schone goltgale Sonneri),and it is odorless. Hedescribes the shape of the bud, and itsgreen "outer leaves." Thebroad, round, disc-like foliage leaves, he says, are stiff and leathery,and grow on smooth hollow {lucken) stems. His figures are quite crudebut recognizable, the " Weiss Seebluomen " as Nymphaea alba, the other"
Historical. *5as Nuphar. A longlist of common names isgiven Wasser : Gilgen,Haarwurz, Haarstrang, Nenuphar in apothecary books, Clavus Veneris,Digitus Veneris, Alga palustris, Papaver palustre. The account of themedicinal virtues of these plants does not materially differ from thosepreviously given.Matthiolus, in his Commentary on Dioscorides (1554), transcribesthe chapter on Nymphaea from that writer with numerous slight verbalFig. 4."Faba Aegyptla.'facsimile from Mftttliiotas.Fig. 5." Nymphaea alba.'Facsimile from Lobelias.changes. He understands " Nymphaea " to signify our N. alba, andgives " Nymphaea altera," also called " blephara," for Nuphar. Thetwo are figured side by side in rude but quite recognizable cuts ;butthese may have been inserted by C. Bauhin, whose notes were addedto the original in the edition which I have seen. On the identity ofthe Egyptian Nymphaea and Nelumbo, Matthiolus was hopelessly atsea ;for Nelumbo had been so long extinct inEgypt that nothingcould be found there to correspond with the descriptions given by theclassic writers.So, after much weighing of evidence, our author invented
1 6 The Waterlilies.a unique figure (Fig. 4) to show what this confused plant must be afigurewhich formed the basis of a heated discussion for the next threequarters of a century. We can only refer to this in passing, sinceNelumbo, and Arum, and Colocasia are the genera most nearly concerned.Lobelius' " Observations of Plants "(1576) figures two " Nymphaeas,"alba and lutea, and gives their medicinal virtues at length. The cuts arerude, but original and easily recognized (Fig. 5) as the speciesstillbearing those specific names. In the " Adversaria Nova," Nymphaealutea isgiven before alba, with most of the synonyms used by Tragus andin addition Nenuphar officinarum, plompen (Dutch), Escudetes del Rioand Higos del Rio (Spanish), Water lilie, blanc d'eau and Lis d'estang.From this time forward, waterlily is a term consistently used as asynonym for Nymphaea to the present day, except that in Devonshire,Eng., Iris pseud-acorus has shared the same common name (Friend, 1882).Nymphaea alba is not directly given any common name by Lobelius, butit is evident that some of those given above apply to it ;hesaysit issometimes called "major" to distinguish it from "N. alba minor," bywhich some small white flower, probably Limnanthemum, is designated.A " Nymphaea lutea minor Septentrionalium " is also figuredit; may be"our Heteranthera. Nymphaea," therefore, would seem to be definableas an aquatic plant with evident flowers and floating leaves ;the idearepresented by the word is acquiring a more distributed content. " LotusAegyptia " {Nymphaea lotus L.) is discussed on pages 385 and 386, butLobelius is unable to interpret the descriptions given by the older writers,and regards the whole thing as a fabrication. He seems to have no ideaof such a plant as Nelumbo.Lyte's Herball (1578) {fide Britten and Holland, 1879) gives a singlename for the white and yellow waterlilies, viz., nenuphar. This name waswith this view.also corrupted into " ninnyvers."In 1589, Petrus Bellonus published observations from Greece, Asia,Egypt, &c. he found; nothing inEgypt to correspond with the olddescriptions of Nelumbo, but found the word " colocasia " used for theroot of an aroid which was used for food, as was already known byMatthiolus. So he concluded that Colocasia, Lotus, and Faba Aegyptiawere synonyms, and made vain endeavors to interpret Herodotus, Theophrastus,Dioscorides and Pliny in harmonyA great beam of light was shed on the subject, however, by Alpinus(1592), who made a special study of the Egyptian lotus. Finding no signof Nelumbo, but only nymphaeas and aroids in Egypt,he considered the
Historical. 17ancient writers wrong in their synonymy;colocasia, he said, was an aroid." Lotus Aegyptia " he believed to be the white flower which, from itsresemblance to the European waterlily, he had no hesitation in calling a" larger nymphaea";in fact he regardsit as practically identical with thespecies so familiar about Venice. For both have the poppy-like head, andseed like millet, and both, he said, retreat under water at sundown toemerge again in the early morning, " as is observed by everyonein thecommon nymphaea"{quae in communi nymphaea ab omnibus animadvertitur).Indeed he was surprised that Pliny and others thought thisbehavior strangein the Egyptian lotus. Both flowers and leaves aresubmerged at night, according to Alpinus, the flowers emerging first, atsunrise, and a little later all the leaves appear. Of course this does notreally occur; but the flower of Nymphaea alba closes into a tight budduring the afternoon and is covered by the dull green sepals, to openagain with the light of the next ;day the disappearance of the leaves,however, must have been purely owing to darkness, and their reappearanceafter the flower must have been due to their darker color. Alpinusstated that the white Egyptian lotus was called by the Arabs " nuphar,"and by the Hebrews " Arais el Nil." After the publication of the workjust discussed, Alpinus received numerous tubers of the Egyptian lotusfrom a Venetian doctor in Cairo ;they were of the size of a walnut, ovate,with fibrous roots, blackish outside, yellowish within ;they were said to beeaten freely by the Egyptians, both raw and cooked the sender also said;that the plant resembled the lesser nymphaea (JV. alba L.) exceptflowers"The which, if it be true, this is the lotus niloticus ofwere blue.that tHfeAthenaeus." Alpinus figures it with entire leaves, insuring its identitywith N. caertilea Sav. At a still later date (1735 b), the same writersummed up the knowledge of the Egyptian lotus (" Lotus Aegyptia ") ina " dissertation " on the subject. As synonyms are given Lotus niliacum,Nymphaea, and Nuphar Aegyptium. The flower and its stalk are calledArais el Nil, the leaf and petiole Bis el Nil, the root (tuber) Biarum.Four plates are devoted to it ;one shows the whole plant, with its dentateleaves, ovoid tuber and the fruit devoid of floral leaves ;another showsthe calyx and fruit on a larger scale ;the third shows two fully openflowers, and the fourth gives an enlarged view of a leaf and a tuber. Theflower is described as having the four outer leaves green outside andwhite within, each with five to seven or more longitudinal nerves, spreadingwide open when in bloom. Within these are about 12 smaller leaves[petals], pure white ;then 25 to 40 long acute leaves [outer stamens],3
1 8 The Waterlilies.white below and yellow above then a;fourth series of slender yellowleaves [stamens] and a fifth series of almost capillary members [innermoststamens], followed by a row of narrow incurved bodies [carpellary styles].At the middle is a small, hard globule, and a head containing the seed.The fruit is round like that of Nymphaea alba, with a green cortex, andseed like that of cabbage. The peduncles are hollow, and of the size ofone's finger.The leaves are crenate (the crucial point of the description),of a dark green color, and so numerous as to cover the whole surface ofthe water ;they die away in November, so that Alpinus considered theplant annual, in spite of the fact that it comes up again at the nextflooding of the Nile. The root, he says, is ovate, of the size of a hen'segg, blackish outside, yellowish inside, and tastes at once sweetish andastringent. This excellent description of Nymphaea lotus deserves permanentrecognition.Itrepresents a masterly attempt to bring order out ofthe chaos which then existed, and that by the true scientific method ofobservation. Had subsequent descriptions been as well written, many ofour difficulties in nomenclature would not have arisen.We may mention here, following the chronological order, that PhilemonHolland, in 1601, translates the word nymphaea used by Pliny as"Nemphar"; this name for the waterlily must have been more or lesscommon, therefore, inEngland at that time.Clusius in his earlier works (1576 and 1601) continues the fruitlessdiscussion of the relations of Lotus, Faba Aegyptia and Colocasia.Concerning Lotus Aegyptia, however, he quotes at length from thevaluable text of Alpinus; but, ignoring the latter's excellent illustrations,he copies the crude figure of the white waterlily from Matthiolus (infact, the same block was evidently used in the printing), and introducesconfusion by labeling it " Nymphaea alba. Lotus Aegyptia Alpini."Beside itappears Matthiolus' cut of Nuphar, marked " Nymphaea luteamajor." In 1605, however, he raised the cloud which obscured theEgyptian bean of classic writers by figuring and describingNelumbo, brought from Java by an intelligent sea-captain.Dodonaeus (1583)a fruit ofrecognizes only two species of Nymphaea, namelyalba and lutea. His figures of these are almost if not quite identicalwith those given by Lobelius, and his description and synonymy beara striking resemblance to the text of Hieronymus Bock. The whitespecies (candidd), he says, has great, broad and nearly round, smooth,floating leaves on long, terete, smooth, porous petioles ; flowers solitary,on similar stalks, composed of many oblong and acuminate leaves, in
Historical. 19the midst of which are many yellow stamens. After the flower, a roundhead isproduced, in which are very many black, shining seeds, likemillet. The rhizomes are thick, nodular, black outside, white and spongywithin, fastened in the mud by many fibrous roots. In addition to thesynonyms given by Lobelius, Dodonaeus tells us that Nymphaea isNenuphar of the shops, and that Apuleius gives the Latin names " materHerculania," " alga palustris," " clavum Veneris,"" digitum Veneris," andthe Gallic name Baditin. To these Dodonaeus adds Wassermahnenand Haar-strang of the Germans. The medicinal powers of the herbare discussed and the subjectis then dismissed without any referenceto the Egyptian species.Besler's magnificent folio (161 3)illustrates in color three kinds of:Nymphaea N. alba major, N. alba minor, and N. lutea, known now asNymphaea alba, N. Candida, and Nuphar lutea respectively.The descriptionof " N. alba major " isgiven with singular accuracy in many details :leaves broad, smooth, thick, green, cleft to the petiole, etc. radix ;spongywithin, and marked with blackish scars (foliorum vestigiis),etc. In thefigure, the characteristic round root-scars are very plain.The material is now in hand for a satisfactory summing up ofthe knowledge of botany as determined by the progressive increaseof accuracy and the widening horizon of science up to this time ;andthe summing up was duly made by Caspar Bauhin. In the Pinax (1623)he divides Nymphaea into two "genera," one with white flowers and,black rhizome, the other with yellow flowers and white rhizome ;the *latter, called " Nymphaea lutea "The former,"includes Nuphar and some other types.Nymphaea alba," is divided into four species. The firstof these, with itssynonymy, is :" Nymphaea alba majorNenuphar album, Brunfelsius.Nymphaea alba, Brunf. Matth. Ang. Dodon. Cord. hist. Lon. Ad. Lobel.Taber. Thai. Ger. Clus. hist.Nymphaea Candida, Trag. Fuch. Tur. Caes.Nymphaea major alba, Lugd. Cast. Eyst.Lotus Aegyptia, Alpini."The last synonym is unquestionably wrongly placed. The secondspecies of Bauhin is " Nymphaea alba minor," being " Nymphaea parva,Matth." Whatever the plant may be, it is not a Nymphaea in the presentsense ;and the same may be said of the third and fourth species of Bauhin.The consideration reduces for our present purposes to a single species,in which, however, two, and possibly three, quite distinct plants are
20 The Waterlilies.included, namely, the European species N. alba, and perhaps N. Candida,and the Egyptian white lotus, N. lotus L. The blue Egyptian lotus,IV. caerulea Sav., mentioned by Athenaeus and Alpinus, is quiteA unique volume is Gerard's " Herball or Generall Historie ofleft out.Plantes " published in 1636. Of his five "Water Lillies" only the firstwould now be placed in the ;genus Nymphaea this is " Nymphaea alba,White Water Lillie," givenin the index of the work, however, as " N. albamajor." The second speciesis"Nymphaea lutea," our Nuphar; theOthers have no relationship with the Nymphaeaceae. The figures ofthe first, second, fourth and fifth species were apparently printed fromthe same blocks as were those of Matthiolus, and the text is but anabbreviated translation of the same writer. The Egyptian lotus is notmentioned at allby Gerard.Vesling, editing Alpinus in 1640, recognized the difference between theEuropean and Egyptian white waterlilies and discussed their relationships.He tells us also that the white lotus flower is woven into triumphal crowns.In the same year Parkinson's fascinating " Theatrum " appeared ; hisarchaic English is most entertaining. He gives a burning paragraph onthe " Egyptian bean " question, having full knowledge of that plant(which we know as Nelumbo), and arraigning Matthiolus for his " false "and imaginary figure of it (Fig. 4).At no point does he refer to thisplant as " lotus." In chapter 29 he gives " Nymphaea, the Water Lilly,"with seven species:1. N. alba major vulgaris, The great common white Water Lilly.2. N. alba major Aegyptiaca sive Lotus Aegyptia, The great white Water Lillyof Egypt.3. N. alba minor [Not a Nymphaea as now understood].4. N. alba minima quae et Morsus ranae vocatur.5. N. lutea major [Nuphar].6. N. minima lutea \ rxT . XT , , ,-\,_ , ,,. > LMot Nymphaea as now understood J.7. N. lutea llore minore )The first is of course our N. alba L. ;it is illustrated by a copy ofLobel's figure (Fig. 5).The second is our N. lotus L., and is accompaniedby Alpinus' figure of the whole plant. Referring to the restingperiod of this plant in winter, Parkinson suggests perhaps in criticism ofAlpinus that the plant has been therefore " called annual, but I wouldcall itrestible."A compilation of former observations with valuable comments, wasmade by Bodaeus a Stapel in the 1644 (Amsterdam) edition of Theophras-
Historical. 21tus. He considered the word Nenuphar a corruption of the older Greeknouphar. But his description of the European waterlilyis the valuablefeature. The leaves, he says, are on long, terete, glabrous, porouspeduncles, are approximately round, firm, almost coriaceous, for the mostpart floating on the water ;upper surface smooth, distinctly veinedbeneath. Those which lie concealed below the water are thinner andsofter. Flowers solitary,on stems like the petioles, made of many floralleaves, so that sometimes a single flower iscomposed of twenty-five tothirty or even forty leaves, each of the shape of one's thumb or a leaf ofthe greater Sedum. From the middle of the flower many yellow stamensproject.Bud oblong,its outer leaves [sepals] purplish green. Flowersscentless, shining like the sun. Here, curiously enough, reappears theat sunrise.myth of its retreat below the water at evening and emergenceThe account continues : After the flower, there isproduced a head like apoppy or round apple, with black shining seed, larger than millet. Therhizome {radix)is about as thick as one's arm, knotty, black outside, whiteand spongy within, odorless, tasteless, beset with and fastened in themud by fibrous roots. Bodaeus calls this plant " Nymphaea alba major."Two other varieties of " N. alba" and four of " N. lutea " are next described,but they are all outside our present genus Nymphaea and neednot be considered here. At another place Nymphaealotus is fully treatedunder the name of " Lotus Aegyptia," givingall of Alpinus' figures.Bodaeus thinks this plant should not be classed with Nymphaea on accountof its bulbous root. He thus differs in opinion from Alpinus, and especiallyfrom C. Bauhin.Piso (1648) gives us the earliest account of an American Nymphaea,in a Brazilian species. " Among those plants," he says, " which arecommon to Europe and the western world is Nymphaea, called by theBrazilians Aguape, by the Portuguese Galvaon, which is noticed everywherefloating on the surface of pools and still waters. The leaves are similarto our Nymphaea, with a great network of veins beneath." The flowerhas a pleasant odor, and consists of four green sepals and about twelvenarrowly oblong, acute, white petals. Its medicinal virtues are alsodetailed. Just what species is here referred to cannot be positivelyidentified, nor does the crude figure accompanying assist. But from thecharacter and number of the petals, and the fact that we know of but onewhite day-blooming waterlily in Brazil, we have but little hesitation inreferring it, as did Caspary (1878), to N. ampla DC.Bontius, whose accounts of the East Indies were bound in with Piso's
22 The Waterlilies.of the West Indies in 1658, devotes considerable space to " NymphaeaIndica flore purpureo"; the figure and description relate very clearly toNelumbo and may be omitted here.Another new feature is introduced by Flacourt (1661) in his " Historyof the Great Island of Madagascar." He found there a blue waterlily,which was considered a delicious food. It is called by the natives " Tantamou."Instead of being anaphrodisiac like the European species, thisone provokes to excesses. The white waterlily, says Flacourt, is called inthis country " Laze-Laze," and the yellow flowered one " Talifouc." Theroots of all three are used for food. The identities of all of these are inthe highest degree uncertain. The blue isprobably N capensis or N.madagascariensis. The white may be N. lotus, or a mere variety of theblue. The yellow one may be a Nuphar at any rate, no yellow Nymphaeais known at present from that island.;Returning to the European flora, we may gather a few points fromJ. Bauhin's " Histoire des Plantes " (1670). Curiously enough he quotesthe name Nymphaea alba minor, with the synonym N parva Matth. fromC. Bauhin, as signifying the European white waterlily, and adds anumber of -the common names given by Lobelius. The real identity ofthe plantis rendered certain by a crude picture. The peculiar mixing ofthe names remains unexplained.Chabraeus (1677) records five species of Nymphaea,viz.: N albavulgaris, N lutea, N minor lutea Lobelii, N. minor lutea flore fimbriato,N. alba minor. The last four would not now be classed in the genusNymphaea at all. The figure of N. alba vulgaris, is an enlarged andvery slightly modified copy from Fuchsius.John Ray's "Catalogue of Plants of England" (1670) mentionsfiveNymphaeas, of which only the first, "Nymphaea alba',' belongs inthe genus as now constituted. The only information given in additionto the name is the words " In fluviis.Florem habet plenum!' The" Methodus Plantarum" of the same author (1682) is especially interestingfor its freedom from slavery to the classic writers. The twenty-fifthgenus contains "Anomalous herbs and those of uncertain position";the first section of these is headed "Anomalae Aquaticae," and the firstof these isNymphaea, plants with round to oblong, floating leaves,differingin their flowers. Three varieties are mentioned, of which" N. alba vulgaris"is the only true waterlily, the common white oneof Europe. The " Historia Plantarum" (1688) follows the classificationof the " Methodus " in general, but describes the European white water-
Historical. 23lily as N alba simply. In many points the description is only a repetitionof earlier writers ;the flowers are said to be odorless, with the outerleaves greenish as in Ornithogalum (ut Ornithogali).A " NymphaeaBrasiliensis," known to the natives as Aguape, and described in nearlythe same words as used by Piso, isgiven here, on the authority ofMarggrav's observations, and classed as belongingto N alba. A fifthspecies is added, N alba major Aegyptiaca. The specific descriptionof this refers to the " crenate leaves " which identifyit with N lotus L.,and the statement is added, " flores . . .semper supra superficiem aquae."The object of this addition comes out when in the succeeding paragraphthe stories of Theophrastus, Pliny, and Alpinus are given regarding theretirement of the flower far under water at night,and the writer boldlyconcludes, " Nobis certe incredibilia videntur."Jacobus Breynius (1680) is one of the few among the early botanistswho considered new and foreign plants. His second Prodromus gives"Nymphaea flore coeruleo odoratissimo, Cap. bonae spei nobis," and;"Nymphaea flore suave purpurascente Japonica nobis." The former;is doubtless N. capensis Thunb., the latter Nelumbo.The magnificent "Flora Malabarica" of Van Rheede (1692) containsfigures of a number of water plants. In vol. 11, plates 26, 27, 28 and 29represent plants with floating leaves, all under the general Malabar nameof Ambel. The first, called simply Ambel, isNymphaea pnibescens; the second,Cit-Ambel is the source of Willdenow's Nymphaea stellata, a bluefloweredwaterlily. The other two, Nedel-Ambel and Tsjeroa-Ambel areLimnanthemums. All are quite fairly recognizable from the engravings,at least as to their generic position. It is interestingto note that thenatives of the Malabar coast classed Nymphaea and Limnanthemumunder the same type or genus, just as did their European contemporaries.The text accompanying the plates refers chieflyto the medicinalproperties of the plants.In 1696 Commelin published Latin names and synonymy for thespecies illustrated by Van Rheede. To Ambel he gave the title " Nymphaeaindica, flore candido, folio in ambitu serrato," although he regarded itas identical with the Egyptian white lotus according to Parkinson, Alpinus,and Vesling. Cit-Ambel was dubbed "Nymphaea malabarica minor, folioserrato" and was a new species to European botany.Perhaps the most magnificent and exhaustive botanical work of thepre-Linnaean period is the " Phytographia " of Plukenet (1691-96), withits exquisite engravings and its several companion volumes of text. We
24 The Waterulies.cannot tell exactly what this writer understood by the term Nymphaea;for of the four varieties figured in the Phytographia, one appears to beTrientalis, another is plainly Marsilia, a third is Nelumbo, and the fourthLimnanthemum. The genusis still more comprehensivein the " Amaltheum" (1705), including in addition to those above mentioned our Nymphaeaalba and N lotus as well as Nuphar, Hydrocharis, Podophyllum, andsome others whose brief descriptions are unrecognizable. Several ofthese had been much more scientifically disposed of bywith whose writings Plukenet was fully conversant. In the " Mantissa "earlier authors( 1 700) Brasenia and Pistia are added as species of Nymphaea, additionswhich one can easily understand. Nymphaea capensis is also given underthe name and on the authority of Breynius as quoted above, but it isconfused with Nelumbo. Finally an American terrestrial plant classed byprevious writers as a Brassica, is here classed as a species of Nymphaea.The white Nymphaea or Laze-Laze of Flacourt is referred by Plukenet inthe "Amaltheum" to his"Nymphaea Indiae" (Nelumbo). Plukenetbrings in no new information for our subject.In 1 707 Sloane described a white flowered Nymphaea with the edgesof the leaves "deeply cut," from the "Fresh River" in Jamaica; he saysits leaves agree with the description of the white lotus of India, and hencehe speaks of it as " N Indira flore candido folio in ambitu serraio" ofCommelin. The identity must have been mistaken, since none of theOld World species are native in the new. Caspary has rightly identifiedthis with N ampla DC. In discussing the names, Sloan suggests thatthe Indian and Egyptian white lotus seem to him to differ very little ;English botanists now rank them as one species. He also states, quotingauthority, that his Jamaica waterlily was carried to the Indies by way ofmerchandise. Perhaps this is an attempt to explain its supposed identitywith the Egyptian plant.By all odds the best pre-Linnaean classification of plants,as is wellknown, was that of Tournefort (1700). The waterlilies are placed in hisSixth Class, " herbs and suffruticose plants with rosaceous flowers."Nymphaea is the last (eleventh) genus in the fourth section of this class,while Nelumbo isseparated from it as the first genusin section five. Inthis respect Tournefort came nearer our presentideas than Linnaeus, whomade Nelumbo a species of Nymphaea. The genus Nymphaeais definedin the " Institutiones " as having a circle of many petals, and a pistil risingfrom the midst which forms a globose or conical multilocular fruit filledwith many oblong seeds. The floral leaves and the fruit are putrescent, and
Historical. 25the genus is" easily recognized " by the broad sub-rotund floating leaves." Nenufar " is given as a generic synonym. Four species are named :1. " Nymphaea alba, major C. B. Pin.," our N. alba.2."Nymphaea Americana, foliis circinatis, maximis, acute crenatis,subtus purjmreis, florealboPlum."3. "Nymphaea Americana, foliis circinatis, minoribus, obtuse crenatis,subtus viridibus, flore albo Plum."4. " Nymphaea lutea, major? our Nuphar luteum.In vol. 2, p. 137-8, the flowers, petals, ovary, fruit and seed of Nymphaeaalba and Nuphar are figured,each. We are not prepared to settle the identitywith also a cross-section of the fruit ofof Plumier's Americanspecies ; it seems likely that No. 2may be referred to N. ampla DC, andNo. 3 to N rudgeana Mey.;Caspary so placed them (1878).We cannotexplain the absence of N. lotus from the list, unless it was meant to beincluded under the species N alba major, as classed by C. Bauhin, who isquoted as authority for this name.Finally Ludwig's (1737) arrangement of the genera may be mentioned.He separated Nymphaea from Nelumbo, as did Tournefort, and describedthe former genus as having five sepals and many small petals this is;doubtless our genus Nuphar. He made an additional genus " Leuconymphaea" which has many petaloid stamens, and covers our present genusNymphaea, as was recognized by Kuntze (1891) and MacMillan (1892).The early plant anatomists seem to have neglected Nymphaeaentirely. Grew and Malpighi make no mention of the genus. Vaillantrefers to the "Grandin his essay on the structure of flowers (17 17),Nenufar-blanc " as illustrating the condition where the stamens arisefrom the sides (reins) of the ovary. Older writers, as we have noted,referred occasionally to the "spongy" rhizomes and the "porous" petiolesand peduncles. The Linnaean and post-Linnsean literature will be citedthroughout the course of the present paper.
CHAPTER II.STRUCTURE.The habits of waterlilies are too well known to need more than a briefstatement here. These plants are found in the shallows of slow streams orstill water all around the world. The long rhizomes of the Castalia groupcreep along in or upon the mud bottom of ponds or in a tangle of vegetationoverlying this, in a depth of water from a few centimeters to as muchas 5 or 6 meters, branching here and there, and dying off behind as theyadvance by apical growth. By far the greater number, however, have anerect, tuberous stem or caudex, which never acquires any considerablelength, but stands with its growing apex about on a level with the mud inwhich its lower portionis buried. From the stem in every case a greatmany stout, fibrous roots pass downward and anchor the plant in place,while the leaves and flowers rise upward. The former lie flat upon thewater surface, the upper side being dry and exposed to the air. Undersome conditions the petioles may be strong enough to hold up the leavesand make them totally aerial. All of the species, at certain periods ofgrowth (germination and beginning of each annual growing season),produce small, thin leaves which lead a submerged existence. The flowersof about half of the species open at the surface of the water and seem tofloat upon it, those of the other half are raised on strong peduncles somedistance above the water level. Only in one or two species (N. rudgeana,N. amazonuni) is cleistogamy known to occur. The flowers in such casesnever reach the air, or only do so for a few hours, after self-pollination hasbeen effected. After anthesis the fertile flowers are drawn down into thewater by movements of the peduncle, and here the seeds ripen. Bymeans of a buoyant aril, the ripe seeds from the irregularly bursting fruitsrise to the water-surface and are floated far and wide.These peculiar conditions of life have impressed themselves stronglyupon the forms and functions of the waterlilies. Manifold adaptationshave arisen in every part of the plant. We shall discuss these in orderin the succeeding pages, taking upfirst the gross morphology, then thefiner anatomy of root, stem, leaf, and flower, and finally the physiology ofthe genus.27
28 The Waterulies.THE ROOT.The primary root inNymphaeaceae is of short duration ;in Nelumbo,indeed, it never becomes more than an embryonic rudiment. The entireroot-system, therefore, of mature plants of this familyis adventitious.The following description relates to conditions in mature Nymphaeas, thecourse of affairs at germination and immediately afterward being reservedfor a later chapter. On digging out a well-grown rhizome of a waterlily,we find it beset by a great number of long, terete, white, spongy roots.In large tropical species (Al. caerulea, lotus, flavo-virens, zanzibariensis,capensis, and hybrids) these may be a centimeter or more in diameter, andthey are nearly as large in strong specimens of N. alba; in N. odoratathey measure usually 0.3 to 0.6 cm. in diameter. Each root is decidedlycontracted at its point of junction with the stem, but reaches its greatestdiameter about 5 cm. out, or less ;thence ittapers almost imperceptiblyto itsapex 30 to 60 or 100 cm. away.In species with a short caudex (Apocarpiae ; Lotos, Hydrocallis)these roots may be so numerous and close together as to be deformed atthe base into prismatic shapes in Eu-castalia the internodes are so;elongated as to cause the roots to appear much more scattered. For theroots all arise from the bases of the petioles, or (in Eu-castalia) from anupraised cushion-like portion of the rhizome upon which also the petiole isinserted (Fig. 20). In N. flavo-virens and its hybrids a thick irregularexcrescence remains after the decay of the petiole, projecting from thetuber a centimeter or more, on the lower side of which the roots areborne, to the number of 5 to 13. In N. odorata 3 to 5 or 6 roots occupya triangular area on the cushion behind each petiole the ;apex of thetriangle is farthest from the petiole and is occupied by the first formedroot ;this one is of medium size. Next to it is a second and much largerroot, and the base of the triangular area is covered with a few muchsmaller ones. Some of the roots of young plants, and of adult specimensof N. flavo-virens, are transversely wrinkled in their upper parts, showinga contractile power. No co-ordinate branches are given off by the rootsas a rule, but they are all clothed with a great number of slender rootlets,and the larger of these may again be beset with capillary branchlets this;arrangement may compensate for the total absence of root-hairs throughoutthe adult plants. A root-capispresent on the tip of every root androotlet, often showing the long, thimble-like shape so frequent in waterplantsit may be black or brown in ;color, but is frequently colorless orwhitish on the smallest rootlets.
Structure. 29The root-capis said by Henslow (1892, p. 507-8) and Van Tieghem(1898) to resemble in structure that of monocotyls rather than dicotyls.In N. flavo-virens and N. caerulea, however, the tip of the root agreesexactly with the fourth type of structure given by De Bary (1884, p. 12),Fio. 6.~ Longitudinal section of root-tip of IV. caerulea. a, air-canul ; s, spacebetween root and upper portion of cap. From a photomicrograph.i. e., there is a transverse layer of initial cells from which all of the tissuesare developed (Fig. 6).togrow only for a very short period,Caspary and Nicolai assert that the cap continuesafter which no farther addition to itis made, though it is persistent, and not deciduous as inNuphar (De Bary1884, p. 413). In a root of N. flavooirsns from a plant just breakingfrom a strong tuber, the capis short-conical and rounded at apex, with
3QThe Waterlilies.long thimble-like prolongation around the root. Its outer surface isbounded by a distinct, smooth layer of cells, which are much larger thanthe underlying ones (cf. Fig. 6).At the apex these outer cells arecolumnar in shape and arrangement and are removed from the region ofinitial cells by 12 or 13 elongated elements (columella) they become;nearly cubical opposite the initial region, where the capthick, and elongatedin the upper free portion of the cap,is five to six cellswhere for somedistance this layer alone is present.The relative size of the root-cap in this species is shown by thefollowing measurements. The cap extends 0.035 mm. beyond the initialcells, and 1.33 mm. above this around the root; its diameter oppositethe initials is 0.51 mm., the same as the diameter of the root aboveFlo. 7. True and false epidermis of roots. N.marliacca chromatella. (a) from very young, (b)older, (d) mature root In transverse section; (c)surface view of (b), t , epidermis, h, hypodermallayer, c, cortex.the cap. A somewhat larger root-tipof N. caerulea, (Fig. 6), taken from ayoung root of a mature plant, shows astructure exactly similar to that of N.flavo-virensin one specimen, however,;the apical portion of the cap had beenworn away in irregular fashion so thatthe primitive outer layer of large clearcells is visible only on the sides. Allof the cells of the cap are arranged inlongitudinal rows. The peripheral portionof the capis continuous with theupper, sheathing part, and iscomposedof narrower cells than those below thebody of the root. The root-meristem and root-cap are formed by atransverse row of cells as described above.A regular epidermis of columnar cells is found near the apex of theroots (Fig. 7 (a) e)the outer walls of these cells are; cutinized, but theirThey increase somewhat in size as the rootbut their outer ends become rounded and somewhat drawnpartition walls are quite thin.enlarges,apart (Fig. 7 (b) e).In most of the species examined (N. flava, tuberosa,odorata minor, marliacea-chromatella, lotus) they become almost cubical inshape (i. e., in cross section ;they are elongatedin the direction of theaxis of the root), then are rupturedin their outer walls, the radial wallsremaining as a black, irregular fringe on the surface of the root (Fig. 7 (d)e), giving it a powdery appearance. In N. elegans X zanzibariensis, theepidermal cells become greatly flattened and epithelioid before rupturing.
Structure. 31In all cases, however, the epidermis persists on the narrower portion ofthe root within about 2.5 cm. of the rhizome. Its loss is the direct resultof the great expansion of the root, caused by the enlarging and maturingof the air-canals.Meanwhile the subjacent outermost layer of cortex has developed inan especiai manner (see Fig. 7 h). At a very early stage this layer iscomposed of columnar cells closely resembling the epidermis at the sameperiod. But as its cells become broader and more nearly cubical, thewalls become collenchymatously thickened, as also do those of the nextone or two layers below (N. lotus, tuberosa, marliacea-chromatella). Theouter walls of the hypodermal cells unite to form a fairly even surface,but the inner walls meet at various angles, owing to the varying depthsof the cells. Thus an even hypodermal coveringis formed in place of thelost epidermis.The cortex may be divided into five layers :(1) the single hypodermallayer ; (2) exo-cortex, a layer of from one to three or four tiers of closelypacked cells without intercellular spaces ; (3) medio-cortex, a tissuecomposed chiefly of large air-canals and constitutingbulk of the root ;nine-tenths of the(4) endo-cortex, consisting of one or two layersof cellsclosely set together, lying adjacent to (5) the endodermis (Figs. 8, 9).The first of these layers has already been described. The exo-cortexis continuous, between the air-canals, with the medio-cortex. In N.odorata (Fig. 8) there are between the outermost air-canals and thehypodermis two series of cortical cells with thin walls and in size of lumenequal to or larger than the hypodermalcells. N. tuberosa and N.marliacea-chromatella have usually three series of cells in the exo-cortex ;they are rounded, with collenchymatous walls, and larger by half than thehypodermal cells. The exo-cortex in N. lotus consists of a single rowof cells (four or five near the base of a mature root),in N. /lava of one ortwo rows. In N. elegans x zanzibariensis there are two or three series ofcells which are thin-walled and three or four times as large in crosssectionas the hypodermal cells ; in longitudinal section, however, thehypodermal cells are much the longer, being about 0.025 mm. long; theexo-cortical cells are only 0.0175 mm. in length, and the cells of the mediocortexabout 0.0076 mm. N. flavo-virens, at least near the proximal endof the root, has five or six cell-layers in the exo-cortex.The outstanding characteristic of the medio-cortex is the presence ofThese are elliptic or rounded or more or less hexagonallarge air-canals.in cross-section, and run parallel throughout the length of the root,
32 The Waterlilies.Fia. 8. Root of N. odorata, transverse section. From photomicrograph; e, remnants of epidermis ,* h, false epidermis.beginning in the meristem immediately back of the initial layer of cellsFig. 6 a). They appear to be without communication of any kind withone another, i. e., the cellsbounding them fittogetherwithout transverse intercellularspaces. A singleperforation 0.0025 mm. indiameter has been observedin the side wallof a canalin a root of N. tuber osa, butthis must for the presentbe considered exceptional.At more or less frequentintervals (0.127 mm. to3.0 mm. for N. tuberosa,1.2 mm. to 5 mm. for N.elegans X zanzibariensis)diaphragms lie directly orobliquely across the canals.A diaphragmis a very thinplate of epithelioid cells(0.0076 mm. to 0.013 mm.thick for N. tuberosd) withsinuous margins the protuberancesof the margins;of the cells meet oneanother, leaving spaces betweenthe sinuses (Fig.12 /). In jV. lotus thecells are more even in outline,with large, round, intercellularspaces. Thusthe passage of gases isleftpractically unimpeded.The walls of the canals area single cell-layer thick, andbear stellate cells (idioblasts)here and there. In A^. tuberosa these occur about 0.32 cm. aparton the average along each canal ;in N. elegans X zanzibariensis they are
AStructure. 33two or three times as frequent, but somewhat more slender, and, thoughof nearly the same actual size, are much smaller in proportionto the size ofthe canals ;they are few in the root of N. flavo-virens, numerous in N.flavaand N. odorata, very plentiful and of massive build in N. marliaceachromatella.Transverse sections show that they occur only in angleswhere three partition walls meet, andtheir arms project into the three adjoiningcanals (Fig. 8).The canals vary in actual and relativesize in the different species, but arealways largest at the middle of themedio-cortex and smaller toward exoandendo-cortex. N. odorata (Fig. 8)has six or seven concentric rings ofcanals, i. e., that number can be countedalong any radius of the root ;they areall quite large and distinct. N. tuberosahas seven to ten rows, very irregularlyplaced the middle and outer ones are;approximately circular in outline, thosenear the endo-cortex are greatly elongatedradially, and the innermost andoutermost ones are scarcely more thanlarge intercellular spaces. N.flava andN. lotus have six or seven rows ofrounded and quite regular canals in themain part of the root, becoming irregularin N. lotus near the stem. N. marliacea-Fio. 9./m/Contractile root of N. flavo-virens, trano-CkrOtnatella has nine rOWS, arranged verse section ; en, endodermis ; ft, false epidermis ;i \T i L .i j i t, latex or tannin cell.much as in N. tuberosa; the radialelongation of the inner canals causes the partition walls in this region tolook like radial bead-like strings of cells. N. elegans X zanzibariensis haseight or nine rows of large canals, each of a rounded-hexagonal shape ;N.flavo-virens shows a similar structure in its nutritive roots.The cells of the partition walls are nearly squarein side view(longitudinal section). The angles of the canals are occupied mostly byrounded-hexagonal cells ;three alternate sides of these face the canals,the other three are joined to the neighboring partition cells. Betweentwo angle-cells is usually a single square or rectangular cell ;but often4
34 The Waterlilies.this divides into two smaller ones. Occasionally the angleis formed notby a single cell but by three partition cells which meet at a point and areconnected by beveled ends. In N. flavo-virens, N. tuberosa, and to someextent in N. marliacea-chromatella, all of these cells are greatly roundedout and turgescent, but their areas of union with one another (commonwall) are narrow and bounded by slightly curved lines in transverse;sections these lines give the appearance of small tubular and branchedcells lying between larger rounded cells ;longitudinal sections reveal thetrue state of affairs. The contents of the medio-cortex cells may be verysmall indeed, as in N. flavo-virens and N. elegans X zanzibariensis, or theremay be starch in the inner parts, as in N. tuberosa and N. marliaceachromatella,or the whole medio-cortex may be loaded with starch, as inN.flava. N. marliacea-chromatella is intermediate in this respect betweentuberosa and flava. The grains of starch are rounded and simple orelongate and double.The endo-cortex is an ill-defined layer, most distinct in N. flavaand N. elegans X zanzibariensis ; in the former it consists of one or twocell-layers,in the latter of but one it is;composed of small cubicalcells, and has rather large intercellular spaces, both between its owncells and between these and the endodermis on the one side and themedio-cortex on the other ;it should not, perhaps, be separatedin thedescription from the medio-cortex.Bounding the cortical tissues on the inner side, an endodermis withthickened radial walls may always be recognized. Its circular outlineisinterrupted by occasional slight sinuosities it is most evident in;N- lotus, odorata (Fig. 8)and elegans X zanzibariensis, least so in N. flavovirensand tuberosa. Its cells contain starchName.in N. flava and marliacea-chromatella.A typical radial bundle system passesalong the middle of the root. Its outermostcell-layer constitutes the pericambium.Against this the xylem and phloem patchesabut ;their number in different species isshown in the accompanying table.In the center of the bundle an areaof pith remains ;this may extend overone-third of the diameter of the bundle inN. odorata and lotus, one-fourth in N.flava, tuberosa, and marliacea-chromatella,or one-sixth in N. elegans X zanzibariensis. In a general way this
Structure. 35shows a greater amount of xylem tissue in species with a greater extent ofleaf surface. The first-formed and smallest xylem tracheae are in contactwith the pericambium one such trachea occurs in each xylem plate, and;from it a single row of tracheae continues radially inward. These are indirect contact with one another in N. tuberosa, elegans-zanzibariensis, flavovirensX wnzibariensis ,and sometimes in N. odorata ; parenchyma cellsmay be interposed between them in N. marliacea x chromatella, lotus, andodorata ; in N. flava the radial lines are quite irregular and difficult tofollow, the tracheae having much the appearance of being scattered throughoutthe parenchyma. Only spiral and spiro-reticulate tracheae have beenobserved (A 7 ",tuberosa, elegans X zanzibariensis). Large phloem patcheswith sieve-tubes and companion cells, but no fibers, lie between the xylemplates they are rounded on the inner side and do not extend so far;toward the center of the bundle as the xylem plates that is, the innermosttracheae project more deeply into the pith than do the inner ; boundariesof the phloem masses. All of the cells of the bundle appear polygonaland nearly isodiametric in transverse section, but even the parenchyma ismuch elongatedin the direction of the axis of the root.Contractile roots differ from the more numerous nutritive roots inthat the xylem is less in quantity in the former, and the air-canals ofthe cortex are much smaller in proportion to the cells bounding them.In a root of an adult plant of N. flavo-virens (Fig. 9) taken up inOctober, the false epidermis and exo-cortex are apparently normal.The outer part of the medio-cortex is collapsed wherever a transversewrinkle occurs, but it consists for the most part of comparatively densetissue, in which the intercellular spaces are not larger than the surroundingcells. The vascular system is cut off from the cortex by adistinct endodermis. Each of the 5 to 9 xylem rays consists of asingle line of tracheae occupying about one-third of the diameter ofthe vascular cylinder. The pith in the center of the cylinder is sharplymarked off from the surrounding tissues by an endodermis with verydistinct radial thickenings. This occurrence of an intrafascicular endodermisseems very anomalous in a flowering plant, but was constantlyfound in contractile roots of adults of N. flavo-virens.In non-contractileroots of this species the xylem rays are often two cells wide, theyextend farther toward the center of the vascular cylinder, and thereis no inner endodermis. In none of the roots of other species wasan inner endodermis found.As stated before, a radial vascular bundle system traverses the entire
36 The Waterlilies.length of each adventitious root. Branches are given off from it at rightangles and pass straight out through the cortex to the side rootlets.Where the root joins the stem, the root-bundle passes in and bendssharply backward (away from the stem apex) and runs more or lessobliquely into the plexus of bundles of the stem. In N. odorata the bundlesof all of the roots from a single leaf-base unite in the peripheral region ofthe stem into one strong trunk (Fig. 10) the ;appearance of a pith in themidst of this trunk in N. alba candidissima gives rise to the " root-bearingIn N. lotus each root-bundlestele" of Gwynne-Vaughan (1896 b).passes in separately and breaks up into a number of branches beforejoining the plexusFio. 10. Vascular bundles ofthe roots of N. odorata. Longitudinalsection of leaf-base.of the stem. It is notable that there is no directconnection between the bundles of any leaf andthe roots which spring from its base, close as theirproximity is.The innumerable small rootlets which coverthe main roots seem to repeat entirely the structureof the latter. The vascular system (in N.elegans x zanzibariensis) is hexarch, with uniseriatepericambium and a tiny central pith ; endodermisand endo-cortex are uniseriate and distinct ;themeso-cortex has only one or two series of verysmall and irregular air-canals ;hypodermis andepidermis occur as above described.The rudiments of roots are laid down belowthe bases of the petioles at a very early period.In a strong rhizome of N. odorata a leaf bud lessthan 0.3 cm. long, including the petiole, had four subtending roots alreadylaid down in the tissues of the cushion just below it ;the posterior two ofthese had the root-cap well defined and had destroyed some of theoverlying cortical cells of the stem. After this time, however, theirdevelopment is very slow ;at the base of a petiole over 8 cm. long theroot apices had just penetrated the epidermis, but were still concealed inthe short pubescence of the leaf-cushion, No pericambial tissue has yetbeen distinguished in Nymphaeaceous stems for the source of adventitiousroots.Reference has been made to cell-multiplication in the apex of growingroots, and to the early origin of the air-canals. These canals are at firstsurrounded by six cells. Between every two of these another cell is cutoff, giving one at each angle of the canal, and one on each side (Fig.1 1b).
Structure. 37The angle-cells are, in this stage, triangular and nearly equilateral(Fig. 1 1 a) ; the lateral cells are narrow and brick-shaped, with their endsabutting on the canals. It is largely by the greatextension of the lateralcells that the air-canals and the roots themselves increase in diameter ;for the shortest side of the young cell becomes much the longest inmaturity ; meanwhilethe angle-cells have expanded also, their threepoints becoming first rounded, then flattened, to give the cell its hexagonalshape (cf. Fig. 8 and n).A diaphragm of an air-canal may be traced back (Fig. 12) to a singlelateral cell near the tip of the root ;this extends out into the lumen of thecanal, by distension of its wall, as a rounded papilla. The papilla is cutoff, and soon divides longitudinally;each half then divides transversely,and again longitudinally.Thus by growth a plate of cells is formed, extendingacross thecanal. As thesecells enlarge, intercellularspaces areformed betweenthem until the maturediaphragm isproduced. Stellatecells develop fromthe angle-cells in/&fc/ fa,Jthe USUal Fl - n.way, viz.,Development of air-canals. Transverse section of root^tip of N. jlavofire/i*.From,photomicrograph, a, earliest stage ; b, division of one angle-cell ;by extension OI the e, cell divisions nearly completed; e, epidermis; h, hypodermal layer; (a) is11 1 farther from apex than (b), but from the same root.wall at severalpoints, with subsequent thickening and deposition of crystals. This isexactly as described for Nuphar lutea by Trecul (1845).Lateral rootlets take their origin while the root tissues are still veryyoung, even within the root cap. They spring from the pericambial layeropposite each xylem ray, and make their exit in the usual way, but notuntil a considerable time after their inception;they do not occur in amature state near to the root apex.THE STEM.Externally the stems of Nymphaeas offer some striking differences,which are at times characteristic of subgenera or even of species.Members of the Eu-castalia group have always a stout, spongy, horizontalrhizome which decaysin its older parts as itelongates by apical growth.(b)
3 The Waterlilies.In JV. tuberosa (var. parva Abbot) and JV. odorata mhtor this may be only5 to 10 cm. long and 1.2 to 2 cm. in diameter, but in the typical forms ofthese speciesit becomes 30, 60 or even 100 cm. long and 4 cm. indiameter, while in the larger forms of JV. alba and Candida itmay reach adiameter of 10 cm. In general shape these rhizomes are terete, thoughthe largest one I have seen (JV. alba candidissima) was somewhat flatteneddorso-ventrally.In the American species they are of a dull whitish color,though this is nearly obscured by fine black hairs ;in the EuropeanFid. 13. Diaphragms in the air-canals of roots ; a, d, e, early stages from transverse ; b, c, from longitudinalmicrotome sections of the root of N. flavo-virens ; /, mature diaphragm with beadlike intercellular spaces fromthe root of ZV. elegana X zanzibarlemi*.species the color is brownish or nearly black. All of the species of thisgroup except JV. tuberosa branch rather infrequently in their stems, andthe branches are stout and attached to the main shoot by a broad base.In JV. tuberosa, however, the rhizome (Fig. 13) is often crowded with short,tuber-like shoots from 1.2 to 8 cm. long and 1.2 to 2 cm. in diameter,attached to the parent axis by a short and very slender isthmus, sometimesonly 0.3 cm. in diameter. In external appearance these do not differ fromthe main rhizome, exceptin size and a darker color. One such tuber oftenhas others, one, two, three or four, of nearly equal size, growing out from itssides and base, so that bunches of tubers are formed. Being light enough
Structure. 39to float on water and easily detached from the parent plant and from oneanother by reason of the brittleness of the tissues of the isthmus, thesetubers serve to distribute the plant widely. Waterlily leaves (" lily-pads ")are eaten by large herbivorous animals (deer, &c), which wade about inthe shallow water ;these must detach numberless tubers and leave themfloating about, as a person does when wading among them. Doubtlessalso wading birds, turtles, bull-frogs and many kinds of fish accomplish thesame end. Special mention of the structure of these tubers will be madelater.Each of the Nymphaeas except the group just mentioned possessesa short, thick, erect caudex. Where this remains throughout the yearFio. 13. Rhizome of N. tuberoaa. Natural size.in a wet and more or less vegetative condition, it dies off in the lowerpart as it elongates above, about keeping pacein its elongation withthe amount of sedimentation taking place around it. Such conditionshabitually surround the Xanthanthae and Chamaenymphaeae, and thesecannot otherwise survive. Species of the Hydrocallis group often havea like perennial growth, as also sometimes have the Lotos and apocarpousgroups in cultivation. But the habit of all of the truly tropical members(Lotos, Hydrocallis, Apocarpiae) is to be dried off completely at oneseason of the year. In this case the large mature blooming plantsdie and rot away. N. flavo-virens alone, with some of its hybrids, isable to withdraw from vigorous flowering growth into a dry, restingall of their nourish-tuber. Young plants, however, readily store awayment in the caudex, which then forms a tuber from the size of a pea tothat of a hen's egg (or much larger in N. flavo-virens). Such resting
40 The Waterlilies.tubers are usually of an ellipsoid shape, or spheroidal in N. gigantea andelegans (Fig. 14). They are of a brownish black color, smooth androunded in the lower half, but more or less acute above, becausethe apical regionisprotected by a dense upward growth of long fine hairsintermingled with dried and shriveled bases of petioles. These tuberslive through the dry season without injury, and begin growth again ina characteristic manner on the return of sufficient moisture and heat.Mature stems of the types under discussion are, like the tubers, of anellipsoid shape, and have grown in our Botanic Garden to a length ofFio. H. Tubers :a, N. amazonum ; b, b', N. gigantea, from two sides 90 degrees apart ; c, e', N. eapensis, twospecimens; d, a", N. elegans, two specimens; e, N. tanzibariensis rosea; f, N. lotus, seedling; g. If. lotus,lateral tuber from mature plant. All natural size.20 cm., and a diameter of 10 cm. in N. rubra var., 12.7 by 7.5 cm. inN. dentata, caerulea and zanzibariensis ; I have seen a resting tuber ofN. Mrs. C. W. Ward (N. flavo-virens X zanzibariensis rosea) in the DreerNurseries which measured 21 cm. by 10 cm. All of these are of abrownish black color. With us N. flava has not exceeded 7.6 cm. longby 1.6 cm. broad, and has a yellowish color, but Mrs. Treat (1877) recordsthe length of an old caudex in the St. Johns River, Florida as 60 cm., anddiameter 10 cm. N. tetragona stem is rather slender, 7 to 12 cm.long by 2.5 cm. in diameter, and is of a blackish color ;N. fennicaissimilar in appearance but stouter, 10 cm. long by 3.75 cm. in diameter.No branching of the stem, that is, no development of vegetativebuds, occurs in N. tetragona, and none inany of the apocarpous species
Structure. 41( neglecting some hybrids), except in breaking out from the resting tuber.N.fennica may rarely fork, as one of our specimens shows. The Lotosand Hydrocallis species, however, give rise more or less freely to budsfrom the lower part of the stem, and these develop into young plantsentangled among the roots and leaves of the parent. On the approachof winter or the dry season, these lateral shoots harden off into restingtubers for the perennation of the plant. Tubers thus produced (inN, lotus) are very irregular in shape (Fig. 14 g), appearing knotty andgnarly, but they still show in their lower half a smooth surface, and onthe upper parts a dense growth of long protective hairs intermingledwith remains of dead petioles. A remarkable development of lateralshoots occurs in N. mexicana. Long stolons originate from the lower partof the stem, even in very young plants, and run horizontally just beneaththe surface of the mud to form a new plant at a distance of 15 to 60 cm.from the parent. No sooner has the young member established a fewroots and leaves than it in turn sends out runners, and so on. Threeor more such runners may be found attached to a single plant. Theirminuter details will be described in another place. In autumn the endsof these runners develop a peculiar form of perennating stem (Fig. 15);it consists of an axis 1 to 1.6 cm. long and 0.48 cm. in diameter lyinghorizontally or obliquely in the soil on;itsupper surface is a continuousrow of 3 to 6 small leaf buds, while from the under surface depends abunch of thick, swollen, starch-laden roots 1 to 3.5 cm. long and 0.3 to0.6 cm. in diameter ;these lie crowded over each other somewhat inrows like bananas on a bunch ;at the tip of each a flattened and distendedroot-cap remains.A last type of stem exhibited by Nymphaea is that produced fromFrom the center of the upperresting tubers when they resume growth.end of the tuber, in apocarpous species, a short, smooth, white stem(stolon), 0.16 to 0.3 cm. in diameter, rises a centimeter or so, and at itssummit a tuft of leaves and roots is formed ;thismay now be broken offand willgrow into a perfect plant, while the tuber will repeat the process,ifstrong enough, as many as 8 or 10 times (N. flavo-virens and hybrids).In Lotos species a much greater area of the tuber, in fact nearlyall of itsupper surface, develops a very large number of tiny shoots, crowded allover it. One, two, three or four of these start forward at nearly the sametime and each develops a long internode 1.2 to 8 or 10 cm. in length asmooth, slender, white or reddish, often much curved stolon 0.16 to 0.3 cm.in diameter bearing a simple lanceolate or subulate bract 0.6 to 2.5 cm.
42 The Waterlilies.long at its summit. This is followed by a second internode, much shorter,0.3 to 1.3 cm. long, and this bears the tuft of leaves and roots which arethe beginning of a new plant. If these shoots are detached, other shootsdevelop, and so on almost indefinitely, even from a tuber no bigger thanthe end of one's thumb.Phyllotaxy in the Nymphaeas presents some peculiarities. As mightbe expected, forms with elongated rhizomes (Eu-castalia) have the simplerorders of leaf arrangement, and where dorso-ventrality appears leaf scarsare only half as numerous on the lower as on the upper side. In N.tuberosa and odorata, the phyllotaxy seems to be as low as f or f, andthe internodes are 1.3 cm. or more long. N. alba shows a much higherorder, probably /r or \\ (Raciborski 1894). But the erect tuberousstems are so densely crowded withleaf attachments as to leave onlynarrow, sinuous lines of stem-surfacevisible ;Ithe order of the spiralshave not determined.One can speak of a definiteanthotaxy also in the genus Nymphaea,since a plant once in flowerproduces its bloom continuouslyuntil it dies. In N. gigantea theFig.peduncular attachments form 15. Perennating body of N. mexieana. Seenseparateand distinct secondary spiralsalternating with two parallel spirals of leaves (Caspary, 1865). In otherspecies the flowers occur in the place of leaves, as members of theleaf-spirals. In every case the peduncles arise entirely independentlyof any leaf axil and there is no visible trace of a bract. The relationof leaves and flowers in the primary leaf-spirals is shown in the followingseries taken from Raciborski (1894). The numbers represent leaves,the letter f stands for a flower. A rhizome of A^. alba taken up inautumn gave 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, f, 12, 13, f, 14, 15, f, 16,17,from nearly opposite sides. Natural size.28, 29, f, 30, 31, f,f, 18, 19, f, 20, 21, f, 22, 23, 24, 25, 26, 27,3 2 > 33- f . 34. 35. 36, 37. 38, 39, 40, 41. 42, 43. f.48, 49, f, 50, 51,f, 52, 53, f, 54, 55, f, 56, 57, f,44, 45,58, 59,f, 46, 47, f,f, 60, 61, f,62, 6$, f, 64, &c. ;the flowers up to 45 have already bloomed, the otherswould come out the next season. In N. zanzibariensis, a very floriferousspecies, leaf and flower alternate regularly after the plant once begins tobloom. "N. stellata" gave 1, 2, f, 3, 4, 5, f, 6, 7, 8, f, 9, 10, 11, f,
Structure. 4312, 13, 14, f, 15, 16, 17, f, 18, 19, 20, f, 2i, 22, 23, f, 24, 25, 26, f, &c;the primary spiral seemed to be sometimes xr sometimes higher. N.rubra in the Munich garden (which Raciborski considered was moreprobably a hybrid) showed 1, 2, f, 3, 4, 5, 6, f, 7, 8, f, 9, 10, 11, 12, f,13, 14, 15, 16, f, 17, 18, f, 19, 20, 2i, 22, f, 23, 24, 25, 26, f, 27, 28, f,2 9> 30, 31, 32, f, 33, 34, 35, 36,f, 37, 38, f, 39, 40, 41, 42, f, 43, 44,45, 46, f, 47, 48, f, 49, 50, 51, 52, f, 53, 54, 55, 56, f, 57, 58, f, 59, 60,61, 62, f, &c.'The structure of the stem inNymphaeaceae has long been a puzzle,and for the most partstill remains so. A transverse section of anyvegetative stem of Nymphaea (Fig. 16) shows to the unaided eye threewell-marked regions, viz., (1) an outer wall of dense cellular tissue, about0.16 cm. thick, consisting of epidermis and exo-cortex, (2) a medio-cortexfull of large lacunae or air-spaces, (3) a dense central core traversed inevery direction by vascular bundles. The secondlayer is traversed by numerous trabecular of densertissue connecting the exo-cortex with the centralcore. The following description applies only tonormal vegetative stems, not totubers.stolons or restingMicroscopically examined, a well-marked epidermisisalways present. In N. lotus and odorata..... ,. 111 /T^.Fig. 16. Rhizome of iV. odor-(rig. 17) this is an irregular layer as to the level a ta; transverse section of inofthe cells on the outer and inner sides, and as5^253to their size, though they are approximately cubicalin shape. The walls are equally thin on all sides.avy a are SCUlar bUnafe s Na tura i 8 urIn N. flava the cells are much shallower than long or broad (about2:1) and form a more even surface. With the exception of a veryminute area at the growing point of the stem, its whole surface iscovered with hairs. Where these firstappear outside the naked area,every epidermal cell bears one. At a short distance back of this point,however, many of the epidermal cells undergo an oblique division, cuttingoff a wedge-shaped segment with its apex at the outer surface on oneside of the cell, and a base nearly equal to half the base of the mothercell ; the apex of the wedge widens out, and soon the new cell is cubical.A single cell thus comes to intervene between two formerly contiguous1Raciborski's TV. flava, having no runners, but an ordinary branching rhizome as in otherNymphaeas, was almost certainly N. odorata sulfurea (N. odorata X flava) and not the true N .flava. His A'', slellala was probably N. caerulea Sav.
44 The Waterlilies.Specie*.
Structure. 45and here andsuch cells, with occasionally a small intercellular space,there a typical stellate cellmany times as large as the cortical cells.In N. lotus the exo-cortex is23 or 24 cells thick ;each cell is ellipsoidal,the radial axis (with regard to the stem) being shortest, and the horizontaltangential axis longest. The walls appear thickened, and probably arecolloid. There are no starch grains, no intercellular spaces and nostellatecells.The relative amount of lacunar cortex isvery different in the differenttypes of stem. In the Castalia group it is quite prominent (Fig. 16), inthe others rather insignificant the; following table shows the relation infive types examined. " Central core " includes endo-cortex and all within it.Species.
46 The Waterlilies.The endo-cortex contains some starch.The ground tissue of the centralcore of the stem consists of spherical cells with large intercellular spaces,as in N. alba candidissima, and is densely filled with starch, even in alarge vegetative stem taken up in autumn and otherwise destined only todecay. In N. flava (Fig. 21) the endo-cortex and ground-tissue areconstituted as in N. alba-candidissima, but the cortex cells seem morecrowded together than the central tissue of the core, and more denselyladen with large starch grains.Stellate cells are occasional in the centralcore, and very plentiful in the cortex of this species.The trabecular which cross the medio cortex (Fig. 16) are composedfor the most part of tissue similar to the endo-cortex, but at the outer endsFig. 18.Caudex of N. caerulea; transverse section.Heavy lines represent vascular tissue; e,exo-cortex ; /, peduncle ; m, medio-cortex ; n, endocortex; p, petiole ; r, root ; 8, stipule. Natural size.they gradually take on the characteristicsof the exo-cortex. The numberof trabecular in a cross-section is about5 in N. flava, 7 or 8 in N. odorata,8 in N. tuberosa, 14 to 15 in a verylarge rhizome of N. alba candidissima,15 or 1 6 in a large caudex of N. lotus.On stripping off the exo-cortex ofa stem o{ N. odorata, I found eightbands of trabecular tissue runninglongitudinally throughout the length ofthe portion studied, connected by transverseor oblique cross-trabeculae atintervals of a half-inch or more.Thepetioles and peduncles are alwayssituated over a longitudinal trabecula,butin which the vascular bundles travel to join the central vascular plexus ;these leaf and flower insertions stood in no discoverable relation to thetransverse and oblique trabecular ;in two cases carefully dissected therewere no cross-trabeculae near the insertion of the leaf. In N. flava slendertrabeculae divide the medio-cortex into fusiform areas, each lying withitslonger axis horizontal and tangential to the stem a leaf or flower;or stolon stands over each such area, and a stout vertical trabeculacrossing the short diameter of the area bears the central leaf traceto the vascular cylinder of the stem, while the lateral traces travelinward near the points of the fusiform area, surrounded by a thin layer ofdense cortex. Caspary (1857 b) states that such a division of the spongycortex into areas is common to Victoria, Euryale and Nymphaea.
Structure. 47In waterlilies which are adapted to enduring a dry season, theresting tubers differ markedly from the vegetative caudex. In the caseof young seedling plants, which alone form resting tubers in mostapocarpous species, the tuber acquires a smooth, corky bark on itsintrusion oflower parts. The leaf and root scars are obliterated bycortical cells, which absorb the spiral thickenings of the xylem (Caspary,1859) and give rise to several layers of typical cork. Although I havenever found these in section to have the regularity of arrangementshown in some trees with a well-developed cork cambium (Ginkgo),I have often with my hands peeled off the bark from a living tuber,and found it to separate as smoothly as the bark of a willow twig inApril. The interior tissues are dense and hard with their load of starch,and the vascular system remains intact except in its peripheral parts.The irregular lateral tubers produced upon the sides of large plantsof the Lotos group seem to possessvegetative caudex and to have much more cortex,much less vascular tissue than theheavily laden with starch. The lacunar cortex is whollyabsent from resting tubers, probably because, as in N.flavo-virens, the cork layer is formed inside the lacunartissue, and all of this region dries or decays away.T t- .. , r^, ,...Fio. 19. Cork formalnh.u-castaha and Chamaenymphaea, subenzation tion at base of root soar,takes place only over the scars of*petioles, pedunclesand roots, and on the boundary of the living and dead tissues at the olddecaying end of the stem. The root scars are healed over by a curvedsurface, concave on the outer side, in the middle of which the vascularbundle projects slightly. The cortical cells along this concave surface (whichlies four or five cells farther in than the proximal ends of the air-canals of theroot) undergo one to three parallel divisions (Fig. 19) to form the cork layer,and not until some time afterward do the adjoining root tissues decay away.I do not know how the vascular bundle of the root is closed. In N. albacandidissima, where the above facts were observed, the old rhizome,decaying at its posterior end, seems to have itsepidermis shriveledand more or less disintegrated and its exo-cortex suberized in its cellwalls,but usually without cell-division. When this layer peels off,carrying with it the leaf scars, a rounded corky invagination or pouchis seen (Fig. 20 b) at the base of each main air-canal of the petiolesand peduncles. The largest pouches extend inward a quarter of aninch, and point slightly backward;they are hollow, and wide open tothe exterior, and are usuallyfull of mud. The whole surface of the
48 The Waterlilies.scar is suberized in a manner identical with that of the root. In onecase (Fig. 20 c)the spiro-reticulate elements of a bundle in a leaf scarwere closed by large thyloses from the neighboring parenchyma. Sincebut 3 to 5, usually 3, layers of cork are formed, the protection isonlytemporary ; and when the water breaks through into the underlyingtissues, a thin corky layer is thrown across the cortex in front of it,sometimes simply by modification of the cell-walls without division oftheir contents. Soon the water breaks through this, and again a partitionis thrown across farther forward. So in the older parts of the stem,the plant keeps up a continually losing conflict with death, resultingin a gradually advancing decay of the tissues.LX(c)a>)TVS*f*JLIn the outer, corkycells of the scars, and toa less extent in the innerones, crystalsare found.Some of these are irregularand rounded in outline, butmost are beautifully rectangular,prismaticor tabu-Fio. 20. (a) Old leaf cushion on rhizome of N. alba candldUsima, Jar. The Tegular CrVStalsshowing: base of petiole with air-canals, insertion of stipule, , andiroot scars, r; (b) transverse section of (a) through the two largest may be transparent, homOair-canals; (
Structure. 49space is filled with longitudinal air-canals separated by partitions of asingle cell-layer in thickness. Throughout the length of the stolon threeto five vascular strands pass along, each surrounded by endodermis withthe characteristic thickenings on the radial walls. Gwynne-Vaughan(1896, a, b) called these strands "steles," and considered the stolonspolystelic. He states that the narrow isthmus connecting the tubersof N. tuberosa with the parent axis are similarly polystelic. I am notprepared to express an opinion upon the stelar nature of these strands,especially since the present idea of a " stele " seems to be of doubtfulvalue, and, to say the least, needs more accurate definition. It will sufficeto describe the appearance of the vascular strands in transverse section,and to mention their constituents. In N. flava a strand has at center alarge, regular and thickly cuticularized air-canal, with two or three(usually three) projecting vascular lobes. Each lobe has one to three(sometimes none) spiral xylem cells next to the cells bounding the aircanal;the rest of the area consists of phloem, with large sieve-tubesand small companioncells. The vascular strands of N. rubra stolonshave the large air-canal with only one or two vascular lobes.In N. tuberosa the very short stolon resembles the rhizome in itsepidermis and superficial hairs, and the large, starch-laden cells of theinterior ;these latter cells include very large intercellular spaces.There are about three vascular strands, bounded by endodermis. Atthe middle of each, or near one side, is a large group of spiral elements,corresponding in size and position to the air-canal in N. flava. A moderateamount of phloem lies about this, either concentrated in fourphloem lobes, or half surrounding the xylem and bearing two or threelesser lobes, or completely surrounding the xylem and scarcely lobed.Two or three of these conditions are seen in a single section.The vascular system of nymphaeaceous stems has always beendescribed as consisting of scattered bundles (astelic), resembling monocotyledonousstructure. In most cases this is about all that can be said(cf. Figs. 16, 18). Trecul (1845) studied the stem of Nuphar for a yearwith considerable success, but was far from being satisfied with his results.Nor do his researches shed much light upon the vascular structure ofNymphaea. Although we can not yet unravel the complicated networkof tracheae and phloem strands, which seem to run in every imaginabledirection and position throughout the inner tissues of the stem, somepoints are brought out by an examination of the stolons and young plantsof N. rubra, and the general and nodal structure of N. flava. This latter5
5oThe Wateri.ilies.species has an additional interest as representing probably the mostprimitivemember of the syncarpous series.In a transverse section of the caudex of N. flava (Fig. 21),one findsa pretty well marked cortex, a broad vascularring with one to three leafgaps,and a well defined pith which is continuous with the cortex throughthe leaf-gaps. The vascular ring, however, presents a most peculiarFig. 21. Transverse section of caudex of JV. flava. b, base of leaf with bundles andair-canals ; e, exocortex ; g, leaf gap ; I, point of branching of leaf trace ; m, mediocortex; n, endocortex ; p, pith (medulla); r, root trace ; t, central leaf trace.structure. Projecting slightly into the pith from the inside of the ring areseen seven or eight vascular bundles, t,of varying size, cut transversely,each with itsphloem against the pith, and its xylem (of equal or greateramount) next to the vascular ring ; these are the central leaf-traces.The greater part of the "ring" consists of horizontal or oblique elongatedelements, mostly of phloem, but with some spiral tracheae ;these arederived from root-traces and lateral leaf-traces. In this ring andsometimes on the outer side of it, next the cortex, are additional bundles
Structure.5icut transversely, with xylem inside and phloem next the cortex ;theiroriginor end I was unable to learn. On the surface of the stem ateach node or point of union of a petiole with the caudex, there is inN. flava a prominent conical projection, on the upper face of whichthe petioleis attached, and on the lower face the roots. Within thetissues of this projectionthe bundles from the attached roots uniteinto a single trace, and this passes horizontally throughthe cortex tothe lower edge of the leaf-gap. A few cells join the vascular cylinderof the stem directly, but most of the trace is contained in two largelateral forks, one right and one left, which start almost horizontallyinto the vascular cylinder and could not be followed. The petiolarFio. 22.-Course of vascular bundles at node of Nymphaeaflam. i) base of petiole ; (b) base of peduncle g leafgap; rr, roots; . vascular cylinder of stem, semi-diagrammatic; (c) diagrammatic projection of (a); p, petiole.bundles are nine in number, three in a median vertical plane, and sixsmaller lateral ones. Of the first three, one is anterior, one median,and one posterior. The others are right and left antero- and posterolateral,with one directly right and one left of the centre of the petiole.On entering the stem, the posterior and median bundles fuse, andare shortly joined by the anterior, the whole forming a large centralleaf-trace. The right and left antero-lateral bundles are joined at aboutthe same level as the above, by the directly right and left bundles, andthe two fused bundles then join the central leaf-trace. This trace passeshorizontally in through the cortex and comes in contact with the roottrace,then passes over the brink of the leaf-gap and turns sharply downwardnext to the pith, giving the appearance noted in the cross-section.About midway on its course through the cortex, the central leaf-trace
52 The Waterlilies.gives off a strong branch to either side. Each branch passes about 45degrees around the stem and then runs obliquely into the vascular cylinder.Meanwhile each has been joined near the middle of its course bya postero-lateral bundle from the petiole, which, on firstentering thestem, bent sharply away from the other bundles and ran along horizontallyin the exo-cortex to the corner of the leaf-area before turning inward.The whole is made reasonably clear by a glance at Fig. 22, (a), (c).A very similar arrangement of bundles was found at the node ofa stolon of N. rubra. It is usual in the Lotos group to find a slenderleaf a short distance below the young plant near theapex of the stolons. This is the only place in anyNymphaea, I believe, where a leaf can be found atsufficient distance from others to admit of simplicityin the arrangement of the bundles. At some distancebelow the leaf in question there are four bundles(stelesof Gwynne-Vaughan) passing longitudinallyin the stolon. Nearer the leaf one of these forks,making five bundles, nearly equidistant from oneanother (Fig. 23, upper end). Opposite the leafbase,the bundle on that side of the stolon dividesFig. 28. Vascular system ofstolon of N. rubra ? at a node ;b, e, lateral leaf traces ;
Structure. 53can only be traced so far as to show that all of the four stolon-bundlesare sooner or later joined by leaf-traces, and therefore that all of thebundles of the stem are common.Concerning the origin and distribution of the bundles in the stolon ofN.flava, we can only say that at the distal end of the stolon the vascularstrands branch and anastomose, in fusing with leaf and root traces, toform the vascular cylinder or plexus of the new caudex. The strands ofthe stolon originate from the four sides of a ramular gapin the vascularsystem of the parent plant similar to a leaf-gap, and the stolons occur inthe place of leaves in the leaf-spirals of the surface of the stem. Theabove evidently refers to a stolon with four vascular strands, which seemsto be the normal number. I have examined one specimen with fivestrands, one central and four peripheral. In this the central strand fusedwith the lowest peripheral one before entering the parent stem, so thatboth passed in as a single strand over the brink of the ramular gapexactly like a central leaf-trace.It would seem from these accounts that the bundles of the stolon arescarcely to be called steles, and that the behavior of the leaf-traces ismore in line with dicotyledonous than with monocotyledonous structures.Prof. Nageli remarked in 1857 (Caspary, 1857, b) that "of the fivevascular bundles which enter each leaf in N. alba, the two lateral pairsin the interior of the stem go to the sides of the stem, but the middleone sends a branch inward to form a central strand, and only in thislatter point does N. alba differ in structure from the characteristics ofa dicotyledonous stem." 1The relations of the bundles of the flower stalk in joining the stemhave been most accurately worked out in N. flava. Seventeen longitudinalbundles were found in the peduncle, viz, a central bundle four; largeones 90 degrees apart around the periphery, of which one is anterior,one posterior, one right and one left ;alternating with these are foursmaller bundles, and eight tiny ones alternate with the last two sets. Thelast eight were not traced into the stem, but they doubtless unite withtheir larger neighbors at the base of the peduncle.Great variation existsin the details of the fusion and anastomosis of the remaining bundles,Fig. 22, (b). However, it seems constant for the four oblique bundles to1 "dass von den 5 Gefassbundeln, die bei N. alba in jedes Blatt eintreten, die beidenseitlichen Paare im Innern des Stammes nach dessen Rande zugingen, das mittlere jedoch einenZweig nach Innen entsende, urn einen centralen Strang zu bilden, und nur in diesem letzterenPunkte weiche der Bau von N. alba von dem Charakter des Dicotyledonenstammes ab." p. 791-2.
54 The Waterlilies.fuse with one or both of the adjacent large bundles, leaving,central one, five strong vascular trunks. Shortlywith thethe central and posteriortrunks unite, leaving four, and then the anterior and right trunks unite,leaving but three traces. The left trunk may now join the fused right andanterior, or these may remain separate and pass to opposite sides of thegap in the vascular cylinder of the stem which is prepared to receivethem. This gap is identical in size and shape with those for the leavesand stolons ;since a peduncle is a form of stem, this should probably becalled a ramular gap. The fused posterior and central trace passes inover the lower edge of the gap and turns downward after the manner ofa central leaf-trace, with itsphloem uppermost (innermost after turningdownward). Some analogy between the bundles of peduncle and petiolewould be desirable, but opinions on this would as yet be premature.THE LEAF.The three types of waterlily leaf mentioned on a previous pagehave been designated by Arcangeli (1890, b) hydrophylls (idrofilli), aerophylls{aerofilli) and aerohydrophylls {aeridrofilli) ; the last includes allof those commonly seen and familiarly known as lily-pads. It will bemore convenient, however, to speak in plain English of submergedor water leaves, aerial leaves and floating leaves. The aerial type isaltogether exceptional. It occurs in N. odorata, alba, tuberosa, mexicanaand marliacea-chromatella when the plants are much crowded, or inN. odorata minor where the rhizome is bedded indeep peat or sphagnumfrom which the surface water entirely dries awayin summer. In the lastcase, specimens from Atco, N. J.have shown a few stomata on the underside of the leaf near the apex, and such leaves are always firmer andtougher in texture than floating leaves ;but other than this no differencehas been noted, and no further mention of them will be made.Floating leaves are present in all Nymphaeas, excepting N. alba,when growing in deep, swift streams (which is a rare occurrence), andperhaps in N. amazonum submersa (Sagot, 1881) and N. oxypetala. Amature flowering plant will have at one time from 4 of these in N.tetragona to 6 or 8 in N. odorata or 10 to 15 in the large tropical species.They are produced in rapid succession throughout the growing season,and after functioning for a few weeks, turn yellow and decay.At the base of the petiole occur stipules of characteristic shapes(Fig. 24). In Castalia the two " stipules are inserted on the edge of thedilated pedestal of the leaf-stalk, and below the articulation of the latter ;
Structure. 55and are connate in one plate-like piece in front of the petiole. They aretherefore intrapetiolar"(Lubbock, 1894). The insertion is usually a littleoblique in Eu-castalia, the curved line of attachment extending fartherback on one side of the petiole than on the other. The fused stipularplate persists for some time after the decay of the leaf. It is about 2.5cm. long, more or less, and lies closely appressed to the rhizome ;ingeneral outline it is broadly oval, with emarginate apex. A shallowfurrow exists along the line of fusion, into which the petiole fits, and fromthe ridges on either side of this the tissues taper out to an extremely thinmargin. The stipular plate of N. flavais erect, lanceolate, 2.5 cm. ormore in length, thin, rounded at apex, and not persistent. In Brachycerasthestipules are fused at base, with distinctand more or less long-attenuateapices. The other subgenera have distinct,slightly unequal stipules, consistingeach of a long (2.5 to 4 cm.), narrow(0.3 cm.) wing, adnate to the side of thepetiole, with free acuminate apex. Inall cases the stipules are whitish or semitransparent,and soft in texture, andcovered on both sides, but especiallyon the back, with deciduous mucilagehairs.Interspersed with these on thebacks of the stipules in N. odorata and ** **-. x ; Mr.Fio. U. Stipules.jte, ; . w. otvata.lotus are short fibrous hairs, one to three cells longin the shaft in the formerspecies, much longer, but still few celled, in the latter. Both kinds of hairrest on similar bases of disc-like cells, as will be described shortly.Ihave observed no vascular supply to the adult stipules of N. lotus, butN. odorata has a principal vein running longitudinally in each ridge of thestipular plate, with about nine smaller ones parallel and nearer themargin. N. flava has also a midrib in each and ridge, may have oneparallelvein outside of this. No others have been examined. Thecellular structure isvery simple. The epidermis of the two sides iscontinuous around the margins of the stipules, verysurface, but irregular and angular next to the parenchyma;even on its outerit consistsof more or less cubical cells in N. odorata and lotus, but in N. flavathe cells are wider than deep and longer (in the direction of the lengthof the stipules) than wide. The interior is filled with nearly uniform,thin-walled cells, roundish or oval, with evident intercellular spaces in
56 The Waterlilies.N. odorata and lotus, polygonal and with extremely minute intercellulars inN. flava.In thickness this parenchyma reaches about eleven cells nextto the petiole in A^. lotus, tapering to one cell at the margin N. flava;has about 9 cells in thickness at the ridges;N. odorata has 16 cells atthe ridges, and throughoutits tissues are many very thin-walled, twopointedor variously forked idioblasts. Numerous small starch grainswere observed in the parenchyma of a young stipule of N. lotus. Inthis species also the stipules are much more prominent when young thanon mature leaves, lending weight to Lubbock's (1894) statement that theyare throughout the genus of especial use in protecting the young leavesand flowers.The petioles of Nymphaeas necessarily vary in length according tothe depth of water in which the plant grows, being always long enoughto allow the leaf some freedom in floating about ; the rangeis thereforefrom a few centimeters to 5 or 6 meters, and that in the same species(N. alba,odorata).In diameter the smallest are those of N. tetragona,0.3 cm. through, and the largest are found in N. lotus, 1.9 cm. through.They vary very little from a cylindrical shape, being, near the ends,slightly flattened on the upper side. Of a uniform green in most species,many of the cells contain a blue-purple pigment in the blue-floweringspecies, or red-purple in the Lotos group N. odorata has; usually deepcrimson-brown petioles, and in N. tuberosa they are quite characteristicallymarked with longitudinal brown stripes in the upper part. Alwayspubescent when young, the hairs persist only at the base in most species,but are permanent in the majority of the Lotos group, and in occasionalspecimens of N. odorata ; Caspary (1865) had such from New York Stateon which he founded his variety villosa, and we have cultivated such aplant, sent from Florida by Mr. Soar of Little River, in the BotanicGarden of the University of Pennsylvania. A plentiful growth of shortfilamentous algae on the petioles and under surfaces of leaves has beenmistaken for pubescence by some writers.Surface sections of the petiole reveal an epidermis composed of foursidedcells arranged in fairly regular longitudinal rows, mostly with thelongest dimension lengthwise of the petiole the cross-walls are often;oblique. At intervals of 1 or 2 to 1 5 or 20 cells, the rows are interruptedby circular, thick-walled cells, the bases of the hairs mentioned above. Inthe Castalia group, hair-bases seem to be most plentiful in N. odorata,though in all species theydiffer in number at different levels on thepetioles. In transverse section, the epidermal cells vary in shape from
slightly wider than deep to slightly deeperlatter shape occurs near each end of the petiole,Structure. 57than wide. In N. iuberosa thewhile near the middle thecells are approximately squarein section. The bases of the hairs, Fig.25, (a), (b) are now seen to consist of a short epidermalcell whose innerend is about flush with its neighbors, surmounted by one or two shallow,disc-like oells which come up level with the outer surface. Two disc-likecells are constant in N. rubra and zanzibariensis X and occasional in N.tuberosa and amazonum, only one being found elsewhere.The disc-likecells are very shallow, one-third to one-fourth the depth of the epidermisin N. odorata and its varieties, but in other species they occupy about halfroOOOfiOo&qb1 0,(a,)Fio. 25.--Collenchyma in petioles,(a) N. sturtevantii ; (b) N. flava ; b, basal cells of a hair.the depth of the epidermis. A thick cuticle isalways present, and wasseen in N. tetragona to be distinctly lamellate.Beneath the epidermis there is a single subepidermal layer, distinguishablein all but N. tetragona by its thin walls, and in N. rubra bythe larger size and somewhat columnar arrangement of the cells. Underthis layer is a zone of collenchyma (Fig. 25),whose cells remain of smallor medium size but are followed internally by much larger fundamentalcells in N. mexicana, flava, rubra, amazonum, and the middle partsof the petiole of N, zanzibariensis X but which become, gradually largerinward and pass more or less insensibly into the fundamental tissue inN. odorata, tuberosa, tetragona and the lower part of petiole of N. zanzibariensisX . This zone is about 3 cells wide in N. flava (a small petiole),4 in N. tetragona, mexicana, amazonum, 5 or 6 in the upper and lowerends of N. tuberosa petiole and the middle of N. zanzibariensis X 6 or, 7in the middle of N. tuberosa, and 8 in N. odorata, rubra, and the two ends
58 The Waterlilies.of N. zanzibariensis X .\x\ N. tetragona the subepidermal layer isso in its innerdecidedly colloid, and in other speciesitmay be slightlywalls. These colloid tissues are of the type usually described in textbooksfor collenchyma the thickenings are massed in the angles where;three or more cells meet, sometimes from their great amount seeming toextend along the narrower sides. The appearanceis often that of a groupof round dull bodies (the cell-cavities) in a highly refractive but homogeneousmedium in less ;pronounced cases the likeness is more that of anetwork with large knots. In Miiller's (1890) valuable classification ofcollenchyma, Nymphaea is cited as an example of his first class, " Cornercollenchyma" (Eckencollenchyni), and as having the colloid matter sometimesconvex and sometimes concave to the cell-lumen ;the latter is mostcommon, but the former may be seen in N. rubra. The greatest amountof thickening occurs about the middle of the colloid zone.The remaining tissue of the petiole, designated already as fundamentaltissue, consists of larger or smaller rounded, thin-walled cells, withlarge intercellular spacesit is traversed ; longitudinally by numerouslysigenous air-canals and by vascular bundles. In N. odorata these cellsare of fairlyuniform size, but in N. rubra the cells bounding the air-canals,and those between the canals, are much smaller than those between thecanal-region and the collenchyma. Many of the largeare bi-nucleate, and in N. flava and N. zanzibariensisfundamental cellsX contain starch.For many years the air-canals of waterlily petioles and peduncleshave attracted the attention of botanists on account of their large size andconstant and characteristic arrangement. Thus Mackintosh (1876),Parlatore (1881) and Masters (1902, a, b),as well as Planchon, Casparyand others have written of them. Two principal types may be distinguishedin petioles of mature plants; (1) that of Eu-castalia, with fournearly equal, large canals placed in a square near the middle of thepetiole, two anterior and two posterior, and (2) that of Lotos, Hydrocallisand Xanthantha, with two greatly predominant canals and two lesser onesat each end of these (anterior and posterior). Other species, as shown inthe diagrams (Fig. 26), approach one or other type more or less, or showintermediate conditions. Starting from such an arrangement as is shownin N. tetragona (19) or gigantea (n), we can easily imagine specializationleading in two directions to the well-marked types above mentioned.Each canal is bounded by an even layer of small cells, with walls slightlyconvex toward the canal, and is lined with a thick cuticular coating, whosefunction, if the researches of Barthelemy (1874) are of isweight, to aid in
Structure. 59Fio. 28.Sections of Petioles and Peduncles.No.1
60 The Waterlilies.the exchange of gases between the air in the canals and the surroundingtissues. For, following Goebel (1893), it seems generally accepted thatthe air-spaces of aquatic plants are for the purpose of internal breathing ;and the researches of Lechartier (1867) as well as those cited by Goebel(1. c.) are in favor of this view. Such an internal cuticle can hardly befor protectionouter surfaces of plants, thoughitmight be simply a chemical product ofin the same sense as this function is attributed to it on thethe contact of air upon an otherwise unmodified livingcell-wall. Similararguments apply to the cuticular coatings on the surfaces of mesophyllcellswhere in contact with air, as in the air-chambers connected with thestomata in most plants. Russow (1884) and Mellinck (1886) regardedthese coatings as intercellular protoplasm; but Kny (1900), working overthe same ground very recently, considers the presence of living extracellularprotoplasm in the large air-canals of water plants improbable ;the writer favors this view for Nymphaea. Kohl (1889) found on thewalls of the air-canals of N. alba small free crystals of calcium oxalate.Between two adjacent air-canals there is often but a single layerof cells (e. g., N.fiava, tetragona, and small leaves of rubra, amazonum,etc.); usually, however, the partitions are 2 to 6 or even 12 cells thick.The thickness varies slightly in different parts of the same petiole, sincethe canals taper a little at each end. The largest ones, at least, open atthe proximal end into the lacunar mediocortex of the stem by pores inthe end-wall ;at the distal end they connect similarly with a mass ofsimilar lacunar tissue which again makes communication with the aircanalsof the veins of the leaf. At each of these points of opening thewalls of the canals and the lacunar tissues are loaded with stellate idioblasts; indeed, the lacunar tissue in N. flava, odorata and tuberosa seemsto be little else than a tangled mass of such cells, with their arms interlacingin every direction. Diaphragms do not occur in the air-canals ofthe petioles and peduncles.It will be well here to dispose of some matters relatingto the internalhairs of Nymphaea. We have adopted for them the Sachsian term" idioblast" as used by Weiss (1878). 1 Their common features are thegreat induration of the walls, and the presenceof numberless small'The earlier terms "pachycyst " of Caspary (1865), " pneumatocyst " of Planchon (1850, e)and " Knorpelzelle " of Hanstein, have never come into general use. Solereder's (1898)" Sclerenchymzelle"is a good term, but a little too broad. Ischirch (1889) has proposed theterm " astrosclereide," but Arcangeli (1890, a) rightly objects to this because so many forms arefound in addition to the stellate-cell; he therefore proposes " cladosclereide."
Structure.6imore or less cubical or rhombohedral crystals of calcium oxalate embeddedin the wall. Schenck (1884) states that the completed crystals are formedon the inside of the primary walls of the cell, followed in the usual wayby deposits of layers of secondary thickening. Kohl (1889) and Arcangeli(1890, a) support this view, in opposition to Molisch (1882) who thoughtthe crystals were only partly embedded in the wall, and to Van Tieghem(Traite de Bot., 12 ed., p. 644-5) wno considers them to be simply incrustationsand not crystals at all. That they are truly embedded is shownby the fact that a protuberancestill remains after the salt is dissolved outby acid. Kohl (1. c.) gives a copious list of plants in which such " tertiary"calcium oxalate is found. Idioblasts are derived from ordinaryyoung parenchyma cells ;these become distended on one or more sidesinto the air-canals or smaller intercellular spaces. In a narrow canal,the further extension of the cell takes place in two directions, resultingin a long, double-pointed (bipolar) idioblast, attached somewhere on oneside. This typeisvery plentiful all through the petioles of the Eucastaliaspecies a row of such idioblasts occurs in these; petioles nextto or near the epidermis;a similar row was seen in N. zanzibariensisand in a hybrid of this near the leaf, but in no other species were theyobserved. If a cell bears two such elongated outgrowths,itsshaperesembles a long narrow H. But where the future idioblast lies next to abroad air-canal, itusually puts out several arms in different directions,giving the well-known stellate form all ;possible shapes from the simpletwo-pointed fiber to a star with 8 or more arms may be found. Thesehave been frequently observed and figured heretofore. They vary innumber at different levels in the petiole. Thus, in N. hiberosa, near therhizome the marginal ring of bipolar idioblasts numbered about 18, nearthe middle of the same petiole 100, near the leaf 60 ;in N. odorata therewere none at the rhizome, 60 to 70 midway, and 100 near the leaf; N.zanzibariensis x had about 100 near the leaf. A comparison of the numberof idioblasts of all kinds in different species was made by countingthose visible in a single field of the microscope (Leitz, oc. 2, obj. 3) atthe center of the petiole (between the four main air-canals);the resultsare shown in the subjoined table. The varieties of N. odorata, viz, minor,rosea, and A^. exquisita Hort., are like the type in this respect. N. flavaand mexicana agree in having no bipolar idioblasts, but many small, shortarmed,stellate cells in the canals.In addition to idioblasts, there are found rather plentifully (one every6 to 15 cm.) in the air-canals of petioles (and peduncles) of Apocarpiae,
62 The Waterlilies.and less frequently in Lotos, irregular cell-masses projecting from thewalls ;these appear as whitish specks to the unaided eye, and are foundto consist of numerous oval, thin-walled, turgid cells with one end freein the canal and the other attached. They develop from a single wallcell, much after the manner of the diaphragms of the canals of roots.Similar outgrowths were found by Mellinck (1886) filling up the air-
Structure. 63side of the canals equidistant from each other and from the former two.According to the size of the petiole there may be an additional, stillsmaller bundle in the middle of each space between the six just located ;Fio. 27. Types of vascular bundle; (a) simple bundle;(b) small bundle with air-canal from petiole of N. luberota;(c) double bundle from peduncle of N. zanzibariensis ; (d)double bundle with two canals, petiole of N. dentata. a, aircanal; 8, sieve tube ; sp, spiral trachasa ; t, tannin or latex cell.and stillanother series of twelve very slenderones may alternate with all of the precedingones.Where four main air-canals are present,there isalways a large bundle in the middleof the petiole where the four partition wallsmeet the; peripheral bundles are arranged onthe same principle as before, though usuallythere is but one large lateral bundle on eachside ;thus the number of peripheral bundleswill be a multiple of four instead of six.In structure the vascular bundles ofthe petiole (and peduncle) of Nymphaeashow that extreme reduction of xylem which usually characterizes waterplants. In the absence of any distinguishable endodermis, the bundlesare sheathed by ordinary, large, fundamental cells occasional;intercellularspaces occur between these and the bundle-cells, but this isnot common the outer side of the;sheathing-cells, however, joins with the
64 The Waterlilies.surroundingfundamental tissue with the looseness common to that tissue.Three types of bundle may be distinguished :(1) The smallest bundles, Fig. 27, (a), consist of a strand of phloem,with perhaps five or six sieve-tubes and a number of companion-cells, andon the inner (central) side there may or may not be one or two spiralelements.(2) Medium-sized bundles, Fig. 27, (b), like the former but largeras to the amount of phloem, and accompanied on the inner side by asmall but extremely regular and round air-canal.(3) Still larger bundles, Fig. 27, (c), differing from (2) in having aphloem strand on the inner side of the air-canal as well as on the outerside ;the inner strand is usually smaller than the outer.Sometimes two air-canals intervene between the phloem strands,Fig- 2 7> ("0- while in the center of a large petiole (or peduncle) theremay be three phloem strands about a single air-canal. The four orsix large peripheral bundles of the petiole usually have two phloemstrands and one or two canals. These air-canals are quite small,scarcely visible to the naked eye, and are bounded by a very regularrow of clear cells, whose walls next to the lumen of the canal areconvex and strongly cuticularized. In very young petioles remnantsof spiral elements may be seen in these canals, showing that they bearthe same relation to the bundle as the well-known canals in the xylemof monocotyls, Zea mays, etc. Where the petiole joins the leaf severalspirals are found at the sides of the canals, and these rapidly increase innumber as we approach the collar until the canal iswholly obliterated,and there is, instead, a mass of spiral elements nearly or quite as large incross-section as the phloemstrands. The neat wall of the canal disappearspari passu with the encroachment of the spirals (cf. Caspary,1858, p. 382, 385 ; 1888, p. 2 ; Solereder, 1898).The lamina of the waterlily leaf (Fig. 28) varies in size from 5 to 7cm. in diameter in N. tetragona and odorata minor to 40 cm. or 60 cm. inthe large tropical forms. It is cleft nearly to the center where the petioleis attached, and has been termed fissi-cordate. Nearly orbicular in mostspecies, it is somewhat oval in N. caerulea, and decidedly so in N. elegans,pubescens, and tetragona. The marginis entire in Eu-castalia and Chamaenymphaea,slightly wavy at base in Xanthantha in;Hydrocallis andBrachyceras it varies from entire in N. amazonum and elegans to deeplysinuate in N. rudgeana, ampla, flavo-virens, capensis, and zanzibariensis ;in N. gigantea and the Lotos group the leaves are sinuate-dentate, with
Structure. 65wide rounded sinuses and sharp, almost acuminate teeth. The apexisusually rounded, but somewhat emarginatein N. caeruleaand sometimesin N. odorata, especially the variety gigantea. In the sinuate-marginedApocarpiae, the tendency is for the apex to lie in a sinus, while inLotos the midrib is continued out into one of the serrations. At basethe two lobes may overlap or just touch or divaricate in the sameplant or species, so that this featureis of verylittlesystematic value.Near the petiole the lobes may befused by their edges for a distanceof 2.5 to 5 cm. in Lotos, 1.3 cm. insome Apocarpiae, or almost nonein Castalia ;this causes the leaf tobe more or less peltate, and thefused portionI have called the pelta(Fig. 28, p). At the apex of thelobes they tend to develop distinctpoints marking off the borders ofthe sinus (the space between thelobes) from the rest of the marginof the leaf; this point is acuminateand sharp in N. gigantea, capensis,and the Lotos group, obtuse but stillproduced and evident in Castalia,altogether absent and the lobesbroadly rounded in N. amazonum.The thickness of the leaves ofmost species gives them a leatheryFio. 28. Leaf of JV. flava, showing upper surfaceon right, lower surface on left half ; a, angle of lobe ;I, lobe; p, pelta; r, principal area; 8, sinus; t,astomatic area ; 1, 2, 3, 4, 5, primary veins.feeling, but they are rather brittleand easily torn. Mostly lying flat upon the water, we have exceptionsin the crisped margins of JV. jlavo-virens, capensis, zanzibariensis andthe Lotos group. N. odorata gigantea often has the margin of thedistal half of the leaf turned up at right angles to the water-surfaceto the height of 2 or 3 cm., somewhat after the manner of Victoria.A rich chlorophyll-greenis the usual color of the upper surface ofthe leaves. In N. flava, mexicana, tetragona, rubra-rosea, amazonum andzanzibariensis, however, there is more or less of brownish-red blotching,and in N. rubra the whole leaf is of a bronzy-red hue. The undersurface of the leaf is quite different. Dull green in N. tuberosa, it is
66 The Waterlilies.reddish to dark crimson-brown in N. odorata, alba, tetragona and flava,dull brownish in the Lotos group, and blue-purple in N. zanzibariensisand gigantea. Dark brownish or blackish blotches and specks markthe under surface in N. flava, caerulea, ampla, elegans and amazonum.The leaves are rolled inward from the sides (involute) in venation.They come up through the water in a more or less erect position, andonly on reaching the surface does the lamina bend over at right anglesi to the petiole (Frank, 1872).The venation, as inferred above, is in general palmate, but there is adistinct midrib with pinnate branching. The lowest pair of veins turnbackward into the lobes, and others radiate out at various it isangles ;sometimes difficult to decide whether the most anterior of these aredistinct or branches of the midrib. Five {N. flava, tetragona) to twelve{N. zanzibariensis) veins on either side of the midrib originate thusdirectly from the base of the leaf, and were called by Caspary (1865)primary veins (Fig. 28, 1-5). They give off small branches along theircourse, but sooner or later fork into two nearly equal parts. A strongcross-vein soon connects the forks of adjacent veins, enclosing a distinctarea between each pair of veins. The area whose long axis stands mostnearly at right angles to the midrib was called by Caspary (1. c.) theprincipal area (Fig. 28, r),and its length in relation to the radius of theleaf was used as a standard measurement in all of his taxonomic work.The vein below it is the principal vein (Fig. 28, 3). Beyond theseprimary areas there is a series of similar meshes, becoming rapidlysmaller out to the margin of the leaf. Here the outer boundaries of theultimate meshes make an irregular, submarginal vein in all but the Lotosgroupin these, three or four veinlets run together to a point in the apex;of each tooth of the leaf-margin. Great variety exists in regard to theprominence of the veins on the under side of the leaves. In Castalia theveins are easily seen, but they are marked by very slight and gradualelevations of the leaf surface. In N. amazonum (Hydrocallis) the veinsare prominent beneath as narrow, low, but quite abruptly raised lines.Among the Apocarpiae, N. gigantea has the veins quite abruptly raisedabove the leaf surface, and standing out as a prominent reticulum ;N.zanzibariensis is similar save that the ultimate veinlets at the leaf margindo not project, and so we pass through N. flavo-virens to A^. caerulea, inwhich the primary veins alone are markedly prominent, and the tertiarybranches are raised very little if any above the leaf surface. The Lotosgroup, however, has an extremely prominent network of veins. The
Structure. 67midrib near the base forms a ridge a centimeter high and 0.3 to 0.5 cm.sides and rounded summit. The primary veinswide, with vertical, parallelare but little lower ;the ultimate branches also, especially where they runout into the serrations, stand out like little cords. Thus by a system ofunderlying rafters the lamina is rendered firm and stiff. In the largeleaves of Euryale and the gigantic foliage of Victoria this method ofstrengthening isIn describing leaf-structure we may speak of (1) the upper epider-(2) the palisade layer, (3) spongy parenchyma, (4) idioblasts, (5) vein-mis,carried to a remarkable degree of development.structure,(6)lower epidermis, and (7) external hairs.Fig. 29. Epidermis of leaf, N. tetragona: (a), lower epidermis with bases ofhairs ; (b), upper epidermis with average stomata ; (c), upper epidermis with verylarge stoma from margin of astomatic area.The upper epidermisisperfectlysmooth and even and somewhatcuticularized on its outer surface. The cells are square or polygonal andfrom 0.0127 to 0.0178 mm. across (in eight species and varieties). Theiroutlines are quite wavyin N. flava, and mexicana, less so in N. tetragonaand alba candidissima, and nearly straight in N. amazonum, odorata, tuberosaand zanzibariensis x . The cells vary in depth, but are about 0.01 mm.deep in N. rubra and odorata, 0.01 to 0.013 mm. in N. tuberosa, 0.013 inN. amazonum, 0.01 to 0.0178 mm. in N. tetragona, and 0.0178 to 0.02 mm.in N. zanzibariensis X . Stomata are present only on the upper surfacesof the floating leaves. They are usually small and round, Fig. 29, (b),and separated from one another by two to three or four epidermal cells ;their openings are comparatively large. The guard-cells stand on a level
68 The Waterlilies.with the surrounding epidermis. The size is quite constant in the eightspecies studied, namely about 0.0176 mm. in diameter ;those of N. tetragona,however, are slightly oval and 0.02 to 0.025 mm. wide by 0.025 to0.028 mm. in length. The average number of stomata per square centimeteris, in round numbers, 340 for N. tetragona and odorata, 390 for N. tuberosa,500 for N. zanzibariensis X ,530 for N. mexicana, 720 for N. amazonum,960 for N. alba candidissima, 1,000 for N. flava. Immediatelyover the insertion of the petiole there is an area totally devoid of stomata,Fig. 28, (t)it is of a rounded; shape with a slender prolongation overFio. 80. Leaf of N. rubra, vertical section ; o, air chamber ; Id, idioblast ;p, palisade layer ; 8, stoma ; up, spongy parenchyma ; v, a small rein.each primary vein, and varies in size from 1.2 cm. to 3.8 cm. across.Here the epidermal cells are slightly larger than elsewhere. At the edgeof this area the stomata are at first far apart and extremely large, Fig.29, [c) ; in N. tetragonaI found them 0.05 to 0.052 mm. long by 0.038 to0.045 mm -wide, in iV. alba candidissima 0.045 mm - l n l by 0.038 mm.wide, in N. odorata 0.043 to 0.045 mm n - lg by 0.038 to 0.043 mm -wide, inN. zanzibariensis X 0.038 to 0.05 mm. long by 0.031 mm. wide, in N. amazonum0.031 to 0.038 mm. long by 0.028 to 0.033 mm. wide. From theseexceptionally large dimensions the stomata rapidly become smaller atgreater distances from the petiole, reaching the averagesize within thelength of 0.5 to 0.7 cm. A border of about 0.16 cm. in width around themargin of the leaf is also devoid of stomata, but there is no increase insize of those nearest the margin.
Structure. 69The palisade layer of waterlily leaves isvery deep, and composed ofparallel strings of cylindrical cells. Between these are long, narrow airchambers,opening through the stomata above, and connecting with thelacunae of the spongy mesophyll below. In N. tetragona the palisade layeris about 0.09 mm. deep, in N. rubra (Fig. 30) o. 1to o. 14 mm. in depth, beingin each case one-half the thickness of the whole mesophyll, and consistingof two to three series of long, narrow cells. In N. odorata the palisadeoccupies a little more than one-third of the thickness of the mesophylland consists of five to seven series of cells with an aggregate depth of0.14 to 0.15 mm. In N. tuberosa there are four or five series of cellswith a depth of about 0.16 mm., or about one-half of the mesophyll. N.zanzibariensisY.has five series of cells, making about 0.14 mm., or a littleless than half the mesophyll;finally, N. amazonum has four series of cells,altogether about o. 14 mm. deep, or half the depth of the mesophyll. Wherethe palisade layeris more than two cells thick, the uppermost ones areshorter than the rest, often but little longer than wide. Ameling (1893,p. 189) gives 0.0166 mm. as the average diameter of a cell of the palisadeparenchyma of N. alba, and points out the interesting fact that this isexactly the same in a leaf 230 mm. in diameter and in one only 95 mm.across. In another table (p. 198) a leaf 190 mm. across had the palisadecells 0.019 mm. in diameter, and this iscompared with a like measurementof 0.0197 mm. in a leaf of Victoria regia 900 mm. across. The closeagreement in size of cells is striking. The lower ends of the strings ofpalisade cells rest against a very loose, irregular layer ofthe mesophyll cells,upper members of the spongy parenchyma (Fig. 30). They are large,thin-walled and turgid, and of the most varied forms ;they connect withtrabeculae or rods and plates of soft cells which pass vertically from thislayer to the lower epidermis. Great air-chambers lying between thetrabeculae occupy five-sixths of the space of the spongy layer. Boundingand supporting the lower epidermis in N. odorata, tuberosa, fiava,marliacea chromatella, and zanzibariensis X. is a 1 to 3 seriate layer ofsmaller, fairly regular, and close-fitting parenchyma cells. This layerbe mayabsent in N. rubra, amazonum, and tetragcma, the air-spaces of themesophyll being bounded below only by the epidermis. Immediately overthe insertion of the petiole the mesophyll cells are nearly cubical, withwalls evenly colloid-thickened, and without intercellular spaces.A very striking feature of the mesophyllis the presence of innumerableidioblasts.In the spongy layer these are usually of the multipolar orstellate form.They project from every side into the air-spaces, and
joThe Waterulies.assume the most grotesque shapes one; might truly say that of thethousands one sees, no two are alike. Since most of the arms end freelyinopen cavities, as do those in the air-canals of the roots, petioles, andpeduncles, it is impossible to assign to them any function of strengtheningthe tissues as Arcangeli (1890, a) suggested, although bipolar and H-shaped idioblasts in leaves may have that use in a measure. Arcangeli'sadditional opinion that they serve as depositories for waste calciumoxalate seems to be a good one, although Kohl (1889) reverses the caseand considers that in all such cases the salt is a by-product of thedeposition of thickening carbohydrates on the cell-wall he; imagines thatthe glucose or dextrose is transported as a calcium compound. Thereare, however, great numbers of idioblasts in the leaf, of still another type,which seem to have a definite supporting office. These are cells of greatsize, with one end spread out in three or four lobes against the inner sideof the upper epidermis from this broad base a stout shaft runs down;between the strings of palisade cells (Fig. 30, id). Reaching the spongyparenchyma, it may break up at once into several radiating arms, or maycontinue straight on down to the lower epidermis and attach itself thereby a narrow base all ;imaginable intermediate conditions may be found.The walls of such idioblasts are extremely heavily thickened. In driedspecimens these make prominences on the upper surface of the leaves,and since their number differs according to the species, they were usedby Caspary (pachycysts, 1865) for systematic distinctions.The veins which ramify copiously throughout the leaf present severalinteresting features. The midrib is marked by a line of modified parenchyma.In the palisade region this line is three to five cells wide. The cellsare nearly cubical and very poor in chlorophyll the upper ones have;thickened walls. These grade off insensibly into a plate of larger, closefittingparenchyma cells which runs through the leaf to the lower epidermis.There is a broad arc of corner-collenchyma just within the epidermis onthe lower side of the rib. Embedded in the plate of parenchyma are thevascular bundles, and alongits sides are two {N. odoratd) to several{N. rubra, zanzibariensisX.) air-canals. Each vascular bundle is surroundedby a sheath of parenchyma cells much smaller than those of theneighboring tissue. Although a well-developed xylem of spiral tracheaeand companioncells ispresent, it is less in amount than the phloem.Near the base of the midrib several types and positions of bundle are metwith (Fig. 31). In N. odorata there is shortly below the epidermis a tinybundle, and just below this a much larger one, both with xylem above and
Structure.7iphloem below. Under this is a double bundle with xylemin the middleand phloem above and below, then another double bundle, placed transversely,and finally a simple bundle with xylem uppermost. The lowestbundle is underlaid by a transverse band of thin yellowish collenchyma.Near the apex of the same midrib the lowest bundle with its band ofcollenhymastill persists above it is an inverted simple bundle, and;above this again, near the base of the palisade layer, is a simple bundlewith normal orientation. Lateral bundles seem to arise along the midribfrom the median double bundles, and as they go farther from the rib theyswing round so as to present a normal orientation of xylem and phloem.At a point nearer the petiole than the section first described, the place ofthe transverse double bundle is taken by an irregular vascular mass,1ift(c)f*J$Fio. 31. Arrangement of vascular bundles in veins of leaf: (a) section near the base; (b) near apexof midrib, N. odorata; (c), (d) from iV. eapemi* X; e, upper epidermis; p, parenchyma; pal, palisadelayer ; pe, lower epidermis and parenchyma ; ph, phloem ; *j>, spongy parenchyma ; x, xylem.consisting of one or two xylem areas, each surrounded by two to fourphloem lobes each ;phloem lobe represents a bundle which emanates fromthe vascular collar, several of which fuse to form the transverse doublebundle of the rib.The primary veins are traversed by a lower large simple bundle withphloem below and xylem above, and a similar smaller bundle above this.In all of the lesser veins the orientation of bundle tissues is strictly normal.The lower curve of the vein also is lined within with corner-collenchyma,but there is no interruption in the palisade layer above;the bundle has aclose sheath of small cells, and a wider investment of parenchyma. In allbut the finest ramifications the veins lie nearer to the lower surface of theleaf and are made fast there by the continuity of their surroundingparenchyma with that which lines the lower epidermis ; for there is such alining layer near the veins, even in N. rubra. The smallest veins lie justbeneath the palisade layer (Fig. 30, v),where the exchange of crude andelaborated materials takes place.I have not observed any free endingsof veins;anastomosis seems to be universal.
72 The Waterlilies.The lower epidermis (Fig. 29, a) has been frequently referred to inprevious pages.It is a perfectly continuous layer, 0.01 to 0.018 mm.thick, quite smooth on the outer surface, and consisting for the most partof polygonal cells, many of which are about twice as long as broad (0.018 X0.036 mm.). In these is located the pigment which colors the under sideof the leaf. More or less plentifully interspersed with them are thickenedcircular cells (suberized, according to Schilling, 1894) representing thebases of hairs. Frequently the other epidermal cells are arrangedaround these in a radial manner. This caused Barthelemy (1874) tospeak of them as pseudostomata, and he thought he could see in thecenter of some of them a small perforation. His opinion lacks confirmation.The circular cells are about 0.02 mm.Via. 32. Development of mucilage hair of N.alba; 1-5, successive stages. (After Schilling.)acrossin JV. flava, mexicana, amazonum,odorata, 0. rosea, and tuberosa ;0.02 to0.028 mm. in N. zanzibariensis X and alba~~~*f^yf candidissima, and 0.025 to 0.029 mm. inj( || ~~^/f^^-^^ A^ tetragona. Their relative number isshown by the following figures, representingthe number visible in a single fieldof my microscope (Leitz, oc. 2, obj. 7) :TV. flava 32, mexicana 19-20, amazonum 52, tetragona 21, odorata 36,o. rosea 34, tuberosa 28, rubra 46. In the last-named species permanenthairs stand on many of these basal cells. Hofmeister in 1868 mentionedN. alba incompany with Fagus, Salix, Quercus, &c, as having hairy leavesonlyin the bud.From what has already been said concerningthe occurrence of basalcells of hairs in the epidermis of the petioles and lower leaf-surface,it willbe seen that hair structures always occur on these parts at some time intheir history the main ribs of the leaves and their;upper surface are alonetotally devoid of them. On a leaf rudiment of N. flava a trifle overo. 1 6 cm. long, some of these hairs are already complete, but others continueto develop, so that on a leaf three times as long all stages mayfound. A young epidermalcell swells outward beyond its neighbors andstill beis cut in half by a transverse wall ;the outer cell continues to swell, and issoon divided by a wall which isstrongly concave outward, making a termi-follow the first innal spherical cell (Fig. 32).A second concave wall maythe apical cell. Now, according to Schilling (1894), who investigated N.alba and other aquatics, the cellulose walls of this cell become mucilagin-and the outer cell breaks down into a mass ofized, the cuticle is ruptured,
mucilage. Cylindrical and clavate mucilageStructure. 73hairs are common in N. alba(Schilling, 1. c), and appressed finger-like ones occur in N. rubra. Theyare shed as soon as the tissues of the leaf begin to elongate rapidly andbecome mature, being of use only to keep the developing organs embeddedin a coat of slime. This phenomenon was noted by Irmisch in1859. Subsequent to the development of mucilage hairs, or along withand among them, longer hairs are formed in most Nymphaeas. Theearly stages of these are exactly as just described ;the apical cell, however,instead of remaining spherical, elongates and may undergo divisionFio. 33. Hairs and hair bases :(a) on petiole of N. odorata ; (o) on petiole of iV. rubra, the shaft being:shed ; (c) on petiole of iV. tetragona, showing- also the collenchyma Immediately below the epidermis ;(d), (e), from rhizome of N. tuberosa.into a number of cylindrical elements as in Eu-castalia. The terminalcell is bluntly rounded in Eu-castalia or acute in Lotos. These longerhairs are persistent all over in members of the Lotos group, but only atthe bases of the petioles in Eu-castalia, and as a specially developed ringof long hairs at the summit of the petiole in N. amazonum. In Lotos theshaft of the hair is unicellular. Its base may be verymuch wider than thebasal epidermal cells in the petiole of N. rubra and sturtevantii, or but littlewider, as on the under side of the leaves of N. rubra. The hairs of thering on N. amazonum consist of 1 2 to 15 cells, and reach a length of 0.48cm. Stahl (1888) considered the mucilage and other hairs as intended forprotection against animals (snails, &c), and Goebel (1893) suggested thatthey served to prevent contact of the young organs with one another inthe bud but; Schilling (1894) regards them rather as useful in preventing
74 The Waterlilies.the access of water to the embryonic tissues. The first idea could applyto the hairs on mature leaves as well as to the younger.It would be uselessto assert that either purpose isunimportant.In a few cases the leaves of Nymphaea have been viviparous.Planchon(1852, c) records this for "N. stellata" (probably N. micranthd) asgrown by Van Houtte. The plant came from English cultivators, withwhom it had never shown this tendency. The new plant springs from theupper surface of the leaf just over the insertion of the petiole. "Alreadyvisible," says Planchon, "though very small on the young leaf, the axis buta tubercle with very rudimentary leaves, [the bud] remains almost stationaryon the adult and functional leaf so long as this retains its connectionwith the living plant.If the leaf is detached or artificially by decay of thepetiole, it puts out roots, develops its leaves rapidly, and seems to hastento live an independentlife." 'recorded.Further details of the process have not beenIt was observed in JV. micrantha Guill. & Perr. and given thusin the description of the species:" Folia . . . inter lobos saepius bulbifera,bulbillis sphaericis pubescentibus." Hooker (1850) also records the factin a few words in describing JV. micrantha in the Botanical Magazine,where the leaf figured is viviparous, and Paxton (1853, a) mentions it in his" Flower Garden." Lehmann (1853, a) went so far as to establish a " TribeIII Bulbophyllon" for his JV. vivipara, JV. guineensis Thonn., and JV. micranthaGuill. & Perr. We have not considered these as specifically distinct.Water-leaves are produced in all seedling Nymphaeas to the numberleaves. Of these moreof 2 (JV. flava) to 15 or 16 before any floatingwill be said in another chapter. In species which have dry resting tubersthe first leaves produced on resuming growth are also entirely submerged.JV. flavo-virens alone, with itshuge tubers, is at times an exceptionto thisrule ;small tubers follow the rule. The perennating bodies of JV. flava(doubtless also of JV. mexicand) begin the growth of each bud with a fewwater-leaves, and the other Castalia species produce them either in latefall or early spring, or both. These members also lose their floatingleaves and produce only water-leaves in unfavorable conditions 2 of culture,Deja visibles, mais tres-petits sur la jeune feuille, pourvus d'un axe en forme de tubercule, mais'avec des feuilles encore tres-rudimentaires, sur la feuille adulte et pleine de vie, ils restent presquestationnaires tant que la feuille tient a son pied nourricier. Qu'on detache artificiellement cet organeou qu'il s'isole spontanement par la decomposition de son petiole, aussi tot le bourgeon poussedes racines, developpe rapidement ses feuilles et semble se hater de vivre d'une vie independante.(Flore des Serres, 8: 285.)'Bonnier (1800) found in several lakes of the Alps shoots of A^. alba without floating leaves,but with an unusually large number of submerged leaves, "as is often seen in streams with rapidcurrent."
Structure. 75while the tuberous species in like circumstances lose all of their foliageand form tubers, from which they come out againfirst with water-leaves.A^r^Fia. 84. First leaves from germinating tubers: 1, first, 2, second submerged leaf; 3, first, 4, thirdfloating leaf of If. gigmitea ; 5, second floating, 6, first, 7, second submerged leaf of N. flavo-virens ; 8, first,8, second, 10, third submerged leaf of If. eaerulea ; 11, 12, first submerged leaf from brood body ofN. mexicana and If. Jlava, respectively ; 13, second floating, 14, second, 15, first submerged leaf ofIf. elegans ; 16, first, 17, second, IS. third submerged. 19, first floating leaf of If. zamibariemiis rosea ; 20, 21,first and second submerged leaves of N. "Mrs. C. W. Word" (N. fiavo-virens X zanzibarienaU). Allnatural size.Wachter's (1897) observations on N. stellata, dentata, rubra, thermalis,and alba showed, by cutting off the roots or leaves from growing plants,that production of water-leaves could be induced by enfeebled nutrition,
76 The Waterlilies.especially by lack of elaborated materials in the absence of mature foliage ;cutting off the roots had a much less marked effect. I have noted also inN. rubra that young shoots acquire floating leaves much sooner if leftattached to the parent tuber than if early broken off and planted out. Itseems quite certain, therefore, that the occurrence of water-leaves in falland springin Castalia is directly due to the cutting off of the leaves byfrost, and the suspension of vitality in very cold water ; and, except inthe large and powerful tubers of N. flavo-virens, the youngshoots of otherspecies must gather strength in themselves before they are able to producefloating leaves. In other words, a leaf-rudiment will, up to a certain^3Fig. 85. Transverse sections of petioles of water-leaves; a. N. rubra, from tuber;b, N. ortorata, from rhizome in spring ; e, N. tuberosa, from tuber.age, develop into a submerged or a floating leaf according to nutritiveconditions.Such submerged leaves as are now under discussion assume a moreor less deltoid shape, varying to hastate (Lotos), cordate, or nearly orbicular(Fig. 34). Specific shapes, so far as known, are given in the taxonomicsection. The petioles are usually 2.5 to 7 or 8 cm. long, thoughsometimes longer, and rather stout in proportion. Structurally they showbut slight differences from the petioles of floating leaves. The thicklycutinized epidermal cells contain many bright-green chloroplasts in theupper part of the petiole, where it is not covered by mud. The fundamentaltissue, too, is rich in chlorophyll throughout. Its outer two to fourlayers are more or less colloid in the angles, but only slightly so at most ;here also chloroplasts are most plentiful. The arrangement of air-canalsdiffers from that of petioles of floating leaves, but in the larger waterleavesthis character approaches nearer and nearer to the typical arrangementfor the species. The tendencyin petioles of water-leaves is towardthe condition with two main canals (Fig. 35) and a great many smaller onesaround and between these, with partitions but one cell thick. Thisapproaches the adult structure of N. flava and tetragona, and points alsoto a probable primitive state in which, like Nuphar and the roots of
Structure. 77Nymphaea and Nuphar, the fundamental tissue of the petioles wastraversed by innumerable small and equal canals. Specialization hasenlarged some of these at the expense of others in Nymphaea, Euryale,and Victoria. The distribution of vascular bundles in petioles of waterleavesfollows the plan laid down for floating leaves. Stellate cells occurin the air-canals, but no thyll-like bodieshave been found.The lamina is of a reddish color in Lotos and Eu-castalia, but is plaingreen in N. tetragona, fennica, and amazonum, and Apocarpiae.Its textureisextremely thin and fragile in Lotos, much firmer, thoughstillvery thin, inCastalia, Hydrocallis, and Apocarpiae. The upper epidermisiscomposedof rather larger, somewhat sinuous cells, and possesses a few stomata.Arcangeli noted stomata on water-leaves of N. alba in 1890 (b),but Brand(1894) failed to find them, and considered Arcangeli's specimens to bearrested floating leaves. Wachter (1897) corroborated Arcangeli'sobservation by examining leaves in chloral hydrate, and extended thework to N. rubra, dentata, stellata, and thermalis, in all of which a fewstomata were found on the uppersurface of the leaves. The lowerepidermis is like that of floating leaves. Great reduction has taken place,however, in the mesophyll. Arcangeli (1. c.) found the palisade layer inN. alba reduced to two strata of globular or ovoid cells, and Wachterfound a single layer of crescent-shaped cells representing the palisade inN. rubra and thermalis. The spongy parenchyma is greatly reduced inquantity, and consists of loose cells with large intercellular spaces.Idioblasts are present, but few in number, and the vascular network issimple. The ordinary involute vernation is found in the water-leaves.They are, therefore, to be considered as reduced floating leaves, and noindication is offered of the wholly aerial habits which must have belongedto the distant ancestors of the Nymphaeaceae. Even the large aerialleaves of Nelumbo, from having stomata still restricted to the uppersurface, must be considered as derived from floating leaves, and in noclose relation to those of the probable terrestrial ancestors of the order.THE FLOWER, FRUIT,AND SEED.Few genera of plants offer such variety in size and color of flowers asis found among the waterlilies. The primitive yellow, but of a rich hue,ispreserved in Xanthantha and the newly discovered N. sulfurea andstuhlmanni; white prevails in Castalia, varying to pink in N. fennica,odorata rosea, and alba rubra. All shades from pure white (N. dentata) todeep crimson (N. rubra) occur in the Lotos group, and from white
78 The Waterlilies.(A 7 ",ampla) to pink (N. versicolor) and royal blue (N. zanzibariensis)in Apocarpiae. In Hydrocallis a creamy white prevails. Orange tintshave been obtained in hybrids of N. flava and alba rubra (JV. aurora,seignoreti).In size, N. tetragona, fennica, and odorata minor stand at oneend of the line, with flowers 3.5 to 5 cm. across, and at the other end areA^. rubra, dentata, zanzibariensis, and giganlea, whose flowers measure30 cm. or more in diameter. There is, of course, great variation withina single species, according to the food supply and other conditions. Othergeneral characters will be brought out in the following description of theflower and flower-stalk.So close is the resemblance between petiole and peduncleinNymphaeathat we shall greatly abbreviate the present section by referringback to the previous one. The peduncles are always terete, and in colorand presence or absence of pubescence are like the petioles of the samespecies (no brown stripes occur on the peduncles of N. tuberosa). InEu-castalia the likeness extends to the length, diameter, and number ofair-canals of the two organs. In Chamaenymphaea the likeness includeslength and diameter. The same is true of Xanthantha when in deepwater ;in shallow water, the peduncles are stouter. These species are alsopeculiar in that the upper part of the flower-stalk is somewhat fourfurrowedwith shallow rounded grooves. In all other species theto raise thepeduncles are 1 to 2.5 cm. in diameter, and stiffenoughmassive flowers 5 to 30 or 40 cm. above the water. The more slenderpeduncles of N. amazonum (Hydrocallis) are yet quite stiff and reach toor verylittle above the water surface. In exceptionally shallow water thesame may be true for all of the Castalia group.The epidermis presents nothing characteristic. It bears a large numberof hair bases. Beneath it there is a single layer of thin-walled cells inN. dentata, rubra rosea, devoniensis, amoldiana, and zanzibariensis. Theinner wall of this layer is somewhat collenchymatous in N. zanzibariensis X ,marliacea chromatella, alba candidissima, and gladstoniana. Following thislayer in the above-named forms, or includingit in N. rubra, lotus, slurlevantii,caerulea, odorata, and tuberosa, there are 3 to 1 1layers of collen-tissue is alsochyma as in the petiole. The character of the remaininglike that of the petioles. In Apocarpiae the cells contain a good deal ofstarch, but not so in Lotos and Eu-castalia. Four main air-canalstraverse the peduncles of Castalia, but six isthe standard number for theremaining;groups these numbers are subject to an occasional variationof one more or less, but much oftener of one more. The canals are equal
Structure. 79and arranged symmetrically about the centre of the peduncle (Fig. 26).Outside of each of these are two rather prominent but much smallercanals, and in N. zanzibariensis each of these is followed by two morequite evident canals. Many smaller scattered ones are always present.Idioblasts are extremely scarce in the Lotos group, frequent in Apocarpiae,plentiful in Castalia fibrous; (bipolar) idioblasts occur throughoutthe peduncle in Eu-castalia and around the periphery of the upper part inBrachyceras. Thyll-like masses of soft cells are frequent on the sides ofthe larger air-canals in Apocarpiae, occasional in Lotos, and not found inCastalia.Vascular bundles are distributed on the same principle as in thepetiole (Fig. 26). In the centre is a large double or triple bundle. Oppositeeach of the 4 or 6 main partitions there is a large double bundle, oftenwith two well-formed air-canals in Apocarpiae and Castalia, but with thesecond canal a mere fissure without definite wall in Lotos. Midwaybetween these bundles are smaller ones, and so on in several series,according to the size of the peduncle. Examples are shown in detail inthe diagrams. The constituents of the bundles are exactly as describedfor petioles.Just beneath the flower the peduncle widens out to form a receptaclefor the insertion of the floral organs. This widening extends over about1.3 cm. in Lotos, making an inverted frustum of a cone with a slope ofabout 45. In Apocarpiae, Hydrocallis, and Eu-castalia the transition ismore sudden, almost like the limb of a salver-form corolla. The resultingreceptacle is usually round, but inChamaenymphaea its upper margin issquare, with distinct and prominent angles. In Xanthantha there are fourrounded prominences on the sides of the receptacle, with distinct groovesbetween the ;grooves are continued down a short distance on the petiole,but they swell again quickly above, so that the receptacle is nearly roundat the line of insertion of the sepals. The color of the receptacleis usuallylike the peduncle, but paler. Within the upper part of this regionthe vascular bundles of the peduncle unite into an extremely complexhorizontal network, from which bundles are separated out for the floralorgans. The complexity of this anastomosis is too great to admit of anydefinition or unraveling as yet.The flower proper has four sepals, of which one is anterior and outer.The two lateral sepals are covered on one edge by the anterior one, andon their posterior sides they overlap the posterior sepal. At times threeor five sepals may occur. I have seen the smaller number in N. rubra
80 The Waterlilies.and the larger in N. elegans, and Eichler (1878, 2 :183-186) records bothnumbers for N. alba. A trimerous flower of N. atnpla was the type ofGaudichaud's N. trisepala.Hexamerous and heptamerous flowers areknown. When five sepals occur, two stand in place of one in theposterior position when only three are present, the posterior one is;lacking (Payer, 1857 Eichler, 1. c).;The number of petals varies from 7 (A 7 ",tetragona) to 33 (Al. capensis).The 4 outermost are largest and alternate with the sepals over each;interval between these, i. e., opposite each sepal, is a pairof smallerpetals, making a whorl of 8. This seems to be followed in Brachycerasand Lotos by another alternating whorl of 8. Of this row usually only 4or 5 are present in N. caerulea, while an additional row is found more orless complete in N. capensis and zanzibariensis. InHydrocallis the petals are all regularly arranged inalternating whorls of 4. Here, also, the outermoststamens follow in one or two whorls of 4 ; then comeone or two whorls of 8, the first of which has twomembers over each interspace between the preceding 4;within this it isimpossible to make out any order.fio. .-Fiorai diagram The stamens of the Lotos group follow on in more orjegs definite whorls of 8, and the same may frequentlyforNymphaea.be seen in Brachyceras. In old flowers of N. caerulea, long after inflorescence,when the organs are spread apart by the growth of theripening ovary, I have several times found the stamens arrangedin 16vertical lines, with mostly 4 stamens in each line. The whorled arrangementis not distinct in Castalia, and Eichler (1. c.) states for A^. albathat there are 4 petals alternate with the sepals, and 4 alternate withthese, i. e., over the sepals. These 8 are the beginning points foroblique lines of successively smaller petals, rising right and left withequal inclination. Each series has, on the average, four members. Theinclination of the spirals is such that each successive petal stands overthe space between two neighbors, so that in all 16 vertical rows of 2petals each are formed. The arrangement of the stamens was not madeout. The inner petals and stamens are inserted at various levels up thesides of the ovary (receptacle). There is a narrow, naked zone betweenthe petals and stamens in Lotos and Anecphya, but in other genera thetwo sets of organs are in continuous series, exceptin some forms of N.Candida and tetragona. The carpels are alwaysin a circle in the middleof the flower, abutting against a central prolongation of the floral axis.
Structure.8iWe will now take up some details relating to the individual floralorgans. The line of insertion of the sepalsisusually smooth androunded but in N. Candida the;bases of the sepals are slightly produceddownward, and in N. fennica the sepals make a sharp, clean-cut anglewith the receptacle. The general shape of the sepalsis ovate. Theyare always greenish and chlorophylloid outside, and soft and petaloidinside the ;margins which are covered in the bud are wholly petaloid.Specific details of shape, size, color, and texture are givenin the taxonomicportion of this paper. The epidermis of the outside of the sepals hasa very smooth surface, and consists of irregularly polygonal cells in surfaceview, nearly square in section. Bases of mucilage hairs are frequentin N. tuberosa and plentiful in N. odorata. In the latter species, also,stomata are present. They begin at or just below the middle of the sepaland are more plentiful toward the apex and are a little larger than thoseof the leaf (0.015 by 0.015 mm.). The cells of the inner epidermis arerounded and prominent on the exposed side, of approximately rectangularshape, and shallower than those of the back of the sepal. Here alsorounded cells like the bases of mucilage hairs are frequent, but no stomata.The parenchyma of the sepalis traversed longitudinally by vascularbundles and in the lower parts by air-canals. Inside the outer epidermisthere are in N. zanzibariensis x about 8 tiers of close-fitting, nearly sphericalcells containing chlorophyll they become gradually larger from without;inward. Then follow 1 or 2 flattened cells and a vascular bundle, or,between the bundles, 2 to 4 flattened cells and an air-canal. About 5roundish cells intervene between the air-canals and the inner epidermis.At the apex of the sepals the structure differs from this only in theabsence of air-canals. Just within the epidermis on each side two or threelayers of cells are feebly collenchymatous, but most noticeably so on the innerround orside of the sepal. The air-canals in this hybrid are rather large,oval, and arrangedin a single row between the middle of the parenchymaand the inner epidermis. The partitions between them are two orthree to several cells thick. The vascular bundles have a well-developedprotoxylem, and at some distance from this, in the larger ones, a smallair-canal. There is a row of larger bundles about the middle of theparenchyma and a row of smaller ones between the air-canals and theinner epidermis, with still others between the canals. Many stellateidioblasts project into the air-canals, but none was found elsewhere inthese sepals. Thyll-like ingrowths also occur. One of these consisted ofbut two dense clavate cells, another had a multicellular base surmounted7
82 The Waterlilies.by a crown of ovate cells with dense granular contents. Intermediateforms were seen.In N. odorata the sepal is quite thick at the base, but very thin above.In the basal region there are about 1 2 tiers of cells between the outerepidermis and the air-canals, of which the first 4 or 5 are slightlycollenchymatous. They are all rounded in outline and become graduallylarger from without inward. Among them are very many bipolar idioblastslying lengthwise of the sepal. The air-canals are very numerous andirregular in shape and size. They are separated by partitions only onecell thick, and occupy a zone about as thick as the denser parenchymajust described. Great numbers of idioblasts of irregularly stellate formsoccur here. Between this zone and the inner epidermis there is a zone ofabout 6 layers of parenchyma. A single irregular row of vascular bundlestraverses the partitions between the air-canals. They have well-developedprotoxylem and, in all the larger ones, an air-canal. The same structureis continued in the apex of the sepal, but with fewer tiers of cells andmuch smaller air-passages. Nearlyall of the parenchymacells containchloroplasts, especially the two or three outer layers, where the cells arecrowded together and polygonal in outline.A section near the middle of the sepal of N. tuberosa shows a stillmore leaf-like arrangement. Within the outer epidermis are two to fourlayers of round cells with large intercellular spaces and containing manychloroplasts. Then comes a zone 3 to 5 times wider than the first, madeup of branching lines of cells and large intercellular spaces. Throughthis zone the vascular bundles travel, accompanied by one or two layersof close-fitting parenchyma. Phloem and xylem are well represented, butno air-canal was seen. The orientation of the bundles is normal in allparts of the flower. Stellate idioblasts are plentiful in the air-chambers.The inner epidermisis backed by a single continuous layer of roundedcells. Near the apex of the sepal the air-chambers are quite small, theintervening partitions are multicellular, and idioblasts much fewer. Thetotal thickness of the sepal, however, may be greater than at the middle.Most of the general features of the petals have been already mentioned,and specific details are sufficiently covered in the chapter on taxonomy.The outer four petals are usually of about the size and shape of thesepals. They are thicker at base than elsewhere, and are traversed by anumber of longitudinal veins which divide up into a fine network in theupper half or two-thirds of the petal. There may be some longitudinalair-canals in the basal parts. No distinction of blade and claw occurs, but
Structure. 83the oval to lanceolate lamina tapers gently to a comparatively broad insertion.Where the petals are brightly colored, the lower parts are usuallypaler, and often yellowish, even in blue flowers. The margins of the petals,as of the sepals, are always entire and are of softer texture than the sepals.The inner ones are shorter and narrower than the outer.Th^ structure of the basal and sepaloid portions of the outer petals ofN. odorata is exactly like that of the sepals, even to the presence of largestomata on the back. The softer parts of these petals, and all of theinner ones, are much thinner. The epidermis of both surfaces is like theinner epidermis of the sepals, and has numerous hair-bases.Parallel andanastomosing veins traverse the parenchyma, accompaniedin the middleline by idioblasts of varied shapes, but without air-canals.The stamens are quite characteristic in the different groups ofwaterlilies. In Castalia, especially N. alba, all gradations are found, frominner petals with a callous yellow tip or with one or two tiny anther cells,to stamens with petaloid base and a small but perfect anther, and so on tothe innermost stamens, where the anther is large and thick and broaderthan the filament. N. amazonum shows similar but more rapid gradations;the innermost filaments are filiform and the anthers narrowly linear. InN. gigantea, however, all of the filaments are slender, and anthers narrow.Lotos also shows no gradation from petal to stamen, but the outerfilaments are as much as half an inch wide at base ;their insertion is wideand somewhat decurrent at the sides. The anther is broad and ligulate,with blunt, round apex. The inner stamens are shorter and narrower,ligulate in shape, and not decurrent. In all of the foregoing groups theconnective is not continued above the anther cells more than as a limitingborder to the anther, with the exception of the outermost stamens ofN' gigantea, which bear a tiny subulate In tip. Brachyceras, however,there is a prominent prolongation of the connective at the apex of a largenumber of the outer stamens. This is least in N. elegans (0.08 cm.), smallin N. flavo-virens, but may reach nearly 1.3 cm. in length in N. zanzibariensis.In this group, also, transitional forms between petal and stamenare sometimes seen. The outer filaments are ovate, often shorter thanthe lanceolate or linear anthers these ;grade by slow degrees into theinnermost stamens, which are short and slender and without appendage.In all species the outer stamens are colored like the petals, excepting theyellow anther cells in Castalia and Apocarpiae and the reddish-brownanthers of some Lotos species. The innermost stamens are wholly yellowin all but N. rubra, where the reddish-brown color persists.
84 The Waterlilies.The structure of the filament resembles that of the inner petals.The epidermis bears hair-bases, the parenchyma is made of soft, roundedcells, and air-spaces are present. There are one to three or morelongitudinal vascular bundles. The mature anther is covered by a singlelayer of square cells, a littledeeper than wide, with thickened inner andlateral walls ;the outer wall is unthickened. The anther cells areparallel and introrse, with longitudinal dehiscence (Fig. 37, a, b). In Lotosand Brachyceras the division between the two loculi of each cell isFia. 8?. a, b, trausverse sections of anther of N. flava nearbase and near apex, respectively ; c, vertical section of stigma,showing papillae, N. tuberosa. All after Chifflot.quite evident after dehiscence,as the pollen dust remainingafter most has been shed liesin two pairs of grooves onthe inner face of the anther.ChifRot (1902) has recentlymade a minute study of sporophyllstructure in Nymphaeaceae,including nine species ofNymphaea. He finds slightdifferences in the venation ofthe stamens. He sums up hiswork (whichI have not hadtime to verify)in a classificationbased on the distribution of idioblasts ("sclerites ") and stomata inthe stamen, and of idioblasts in the carpels (1. c, p. 280). We quotethisin translation. The names are correct and mostly well chosen, unlessin the case of "N. blanda." The material of this came from the KewHerbarium, where N. rudgeana occurs under the name of N. blanda, andis much more plentiful.Syncarpiae :Idioblasts rare in the filament, always present in the anther a.Idioblasts absent from filament, anther, and ovary (Xanthantha)N. flava.Idioblasts present in the filament and outer part of ovary (Castalia) c .a. Stomata present on upper epidermis of filament (Lotos). Idioblasts absent frommesophyll of ovary, present in carpellary styles A^. Ihcrmalis .a. Stomata wholly absent from the stamen (Hydrocallis) b.b. Idioblasts of but one kind, present in filament and anther "N. blanda."b. Idioblasts of two kinds, localized in the anther N. oxypetala.c. Idioblasts absent from the anther, but numerous in the carpellary rays d.c. Idioblasts present in anther, rare in carpellary ray N. tuberosa.d. Stomata absent from whole stamen N. alba.d. Stomata present at base of loculi on both faces of filament. N. pygmaea [ = N. tetragonal.
Structure. 85Apocarpiae :Idioblasts rare in the filament, always present in the anther. Stomata absent from allthe surface of the filament (Brachyceras). Idioblasts absent from all partsof the mesophyll of the ovary and carpellary stylesIdioblasts absent from filament and anther (Anecphya)TV. caerulea.N. gigantea.The dry microspores (pollen grains) of N. alba are oval and deeplyfolded ion one side (Kerner, 1895, fig. 218); in Brachyceras (Fig. 38, b-e)they are spheroid, with greatly flattened poles, or, in N. capensis, lenticular,and the edges may be folded under from three sides. Fischer (1890)speaks of them as folded in (eingefaltet) in N. dentata, lotus, rubra, andalba, the infolded half being separated from the other by an equatorial line.Along this line the exine separates in germination. The exine on theinfolded half is thinner and smoother than on the other half, and hasparallel to the equatorial line a ring of similar thickness and sculpture tothe outer half (Fischer,1.c).In water the grains become spherical in aAFio. 88. Pollen: a, young grain of S. flavo-virtns ;b e, N. caerulea; b, justmatured ; c, nearly dry ; d, e, as shed from the anther, seen from opposite sides.few minutes, and along the equatorial line in N. capensis, caerulea, zanzibariensis,and hybrids the two halves of the exine separate slightly;theyseem to be too small to cover the whole swollen grain. In pollen of N.odorata mounted in glycerine some jelly, grains have the exine caps stilltouching at one edge, but wide apart at the other, while some are equidistantall round, and some are unbroken. Fine rods, points, and granulescover the exine in Eu-castalia, and one-half of it in N. tetragona and flava(Caspary, 1865 ; 1888). It is smooth inBrachyceras, and, according toCaspary (11. a), in Anecphya, Lotos, and Hydrocallis. Fischer (1890),however, found itgranulate (kornig-walzig) in N. rubra and dentata, andsmooth in N. lotus, and I have found it smooth in N. mexicana. Ingermination Caspary frequently refers to an operculum, and in one place(1888) to the "smaller half" of the exine but the two; parts which I havenoticed in Brachyceras and N. odorata are practically equal, and Fischerspeaks simply of two pieces (zwei Stilcke) in N. alba, and elsewhere ofthe two " halves " of the exine. The term " operculum " is thereforemisleading.The pistil iscomposed of 5 to 35 carpels, the number being extremelyvariable for any one species. They are placed radially round a central
86 The Waterlilies.prolongation of the floral axis and are immersed in the excavatedreceptacle with about half of the ventral suture exposed above and theother half fused with the central axis. The back of each carpelis more orless prolonged above the receptacle (except in N. gigantea) into a stylarprocess. We shall speak of this as a carpellary style (Fig. 39, c).Theexposed portions of the ventral sutures are connivent or united into abroad, round disc, cup, or funnel, the whole surface of which is stigmatic.At the center of this the floral axis projects as a process (Fig. 39, a) whichis ovoid or nearly spherical in Eu-castalia, narrowly ovate in N. tetragona,linear and rounded in Xanthantha, linear and acute in Hydrocallis, broadlyconical and obtuse in Lotos and Apocarpiae. It is of a dull white color.Fig. 39. Fig. 40.Fig. 38. Diagram of ovary (Brachyceras) in longitudinal section : a, axile process ; c, carpellary style ;n, air-oanals ; r, receptacle : a, stigma tic papllhe ; t, torus, with bases ot floral leaves.Fig. 40. Diagram of ovary in transverse section :a, syncarpous ; b, apocarpous.The stigmatic surface (Fig. 39, is s), usually yellow and densely papillose.The papillae completely cover the upper surface of the small rounded dorsalprojections of the carpels of N. gigantea, and in other species they extendfor a short distance up on the inner face of the carpellary styles. Thestyles themselves are, as indicated, wantingin N. gigantea. In Brachycerasthey are short, stiff, orange-colored bodies, grading from mere pointsin N. elegans to erect processes in N. flavo-virens or stout, incurved bodies0.6 cm. long in N. zanzibariensis. In Xanthantha and Chamaenymphaeathey are more or less ligulate, and stand out horizontally in the basal part.In Eu-castalia and Lotos they are 0.6 to 1.3 cm. long, ligulate and flexible,and in Hydrocallis of about the same length, slender and terete below, witha large clavate end.In the structure of the ovary Caspary (1865) was first to recognize twofundamentally different types. In most of the old group Cyanea of DeCandolle and Planchon he found the walls between the cells of the ovary
Structure. 87are composed of two distinct lamellae, which easily fallapart along theirsmooth surfaces of contact. In all other species there is no such divisionin the walls, but the tissues in each partition are perfectly continuous.The firstgroup was called on this account Lytopleura [Xuzo
88 The Waterlilies.division between the carpels lies in the middle of each partition andextends from a point in the wall of the ovary a little farther in than theback of the ovary cells to a point in the central axis distinctly farther inthan the inner edge of the cells. On each side of this line is a smoothface of square or columnar epidermal cells ;the two sides are in contact.This epidermisis backed by two layers of closely packed rectangularcells ;then comes a line of open spongy tissue, then a single layer ofparenchyma supporting the epidermis which lines the cells of the ovary.This epidermisiscomposed of short columnar cells of highly glandularnature ;each one bulges out into the cavity with a round, turgid end.They doubtless secrete the mucilage which fills the ovary. The axis ofthe ovary is composed of spherical cells with large intercellular spaces inN. zanzibariensis X in Eu-castalia ;(A^ alba candidissima, odorata, (uberosa)the spaces are so large that the cells are drawn out into several broadarms. Many stellate idioblasts were present in N. odorata. In the upperin which it termi-parts the tissues of the axis become solid. The processnates is firm and composed of thin-walled parenchyma the cells are often;slightly elongated and arranged in rows bending from the center out to theperiphery. A regular epidermis of small cubical cells covers the process.The stigmais marked with a number of narrow radiating furrows, onebetween every two carpels and one along the ventral suture of each.Indeed,the ventral sutures are scarcely closed and are easily torn apart.The stigmatic papillae (Fig. 37, c)are crowded as close as they can standand consist of rows of 2 to 8 short cylindrical cells. The terminal one isrounded and contains large, oily-looking bodies. In Hydrocallis the cellsseparate from one another and the stigma is covered with a moist,powdery mass of oily-looking cells, some of which remain attached toeach other in strings.The carpellary styles have numerous mucilage hairs on the epidermisand are made up internally of spherical cells loosely laid together. On theclavate apex of the styles of N. amazonum, the epidermis has a slightlycorrugated cuticle, and the cells contain a pink-purple pigment.The vascular system of the ovaiy presents certain regular features.From the complicated plexus in the base of the receptacle the immediateconnections have not been worked out, but sooner or later there isseparated a ring of bundles which ascend with numerous branchings andanastomoses in and just outside of the lacunar tissue of the ovary wall,and a ring which ascend in the central axis near the edges of the cells ofthe ovary. In N. odorata another bundle passes up the centre of the axis
Structure. 89to terminate in the axile process. From the bundles of the outer ringbranches are given off to the petals and stamens on the one side and tothe walls and partitions of the ovary and ultimatelyto the ovules on theother. Finally, a bundle extends into each carpellary style. In Syncarpiaethe bundles of the inner ring, like those of the outer, are irregularlyplaced with regard to the carpels, and they seem to unite among themselvesby oblique branches but in ;Apocarpiae there are one or twobundles in the inner edge of each carpel, and these are separated fromneighboring ones by the divisions between the carpels. The innerbundles branch copiously into the partitions between the cells of the ovaryin order to connect with the ovules. The bundles are so oriented as tobring xylem next to the ovary cells and phloem away from these, that is,normally with regard to the carpel itself. Consequently the bundles ofthe outer ring have xylem on the axial side, but the inner ring has phloemtoward the axis. In the partitions the bundles nearest one cell have theirxylem next to that cell since those on the other side of the; partition aresides turned toward onesimilarly arranged, the two sets have their phloemanother. The central axile bundle of N. odorata has one or two spiraltracheae with phloem all round, but especially thick on two opposite sides.Some peculiar vascular bundles occur in the ovary wall. The largestbundles of this type have an outer mass of phloem, then a band of xylem;this isfollowed by a band of small-celled parenchyma, and this by anotherxylem band equal to the former. These are comparable with the smallerveins of the leaves where two bundles stand one above the other ;in theovary the phloem of the upper bundle is suppressed and the interveningparenchyma reduced.The fruit of a waterlily is technically a berry. The outer wall of theovary remains firm, but its inner tissues become soft and mealy,i. e., thecells easily separate from one another. The ovary-cells are filled with agreater or lesser number of hard spheroidor ovoid seeds embedded inmucilage, and each surrounded with an aril. Dehiscence is irregular the;fruit simply bursts open by the growth of the arils and swelling of themucilage. In N. flava, however, there is a decided tendency for the carpelsto separate, still containing the seeds. The ripening of the fruittakes place under water, during which time various changes occur in thefloral leaves. In Castalia these completely rot away, leaving only smallscars. In Lotos about half an inch of the bases of the outer stamensremains attached to the fruit.In N. amazovum the sepals persist and theother parts of the flower may be only partly decayed. In Brachyceras
90 The Waterlilies.the sepals and outer four petals remain and become stiff and turgid anddevelop a good deal of additional chlorophyll. The fruits of N. tetragonaAnd/lava are small (2 to 2.5 cm. long), and the persistent carpellary stylesform only an insignificant crown around the top. In Eu-castalia thestigma grows very little if any after anthesis, but is about half as wide asthe diameter of the spheroid fruit. But in Brachyceras and Lotos, wherethe fruits become 8 to 12 cm. across, the stigma keeps pace with thegrowth of the rest of the pistil,so that the fruit has much the same shapeas the ovary of the flower.The seeds of waterlilies offer marked characteristics which are oftenof great systematic value.They were made an object of special study byWeberbauer (1894), from whose valuable paper many of the followingdetails, especially regarding N. alba, are taken. Some important species,however, e. g.,N. tetragona and flava, did not come under his notice.The largest seeds are those of Xanthantha and Anecphya, then N. tetragona,followed by N. tuberosa, Candida, alba, and odorata ; those of Lotosand Brachyceras are considerably smaller than the preceding; the smallestare found in Hydrocallis. They vary in number inversely as the size.N. flava may have only seven perfect seeds to a fruit, or one to three percarpel ; N.tuberosa has sixty to seventy, while N. lotus, rubra, caerulea,capensis, zanzibariensis, and amazonum have thousands. They are nearlyglobular in N. flava, flavo-virens, elegans, and amazo?ium, and ellipsoidinthe rest. The micropylar end isalways slightly pointed, and in Eu-castaliaone side.and Chamaenymphaea the raphe forms a gentle swelling alongIn Castalia the color of the seed is usually greenish black, but brown is thecolor in N. tuberosa. Dark brownish-olive tints are found in allregularothers that I have seen, except N. amazonum, which is reddish brown,covered with silvery hairs.The micropyle and hilum lie near together at one end of the seed.The character of the surface, whether smooth or ridged or hairy, can bestbe discussed in describing the microscopic details of structure.The aril which is attached to the ripe seed is a bell-shaped outgrowthof the funiculus. It is whitish and translucent, mucilaginous, and holdsbetween its two coats tiny air bubbles. It is much shorter than the seed inN. tuberosa, a littlelonger than the seed in N. odorata (Fig. 41, a), lotus,caerulea, capensis, zanzibariensis, and amazonum, but in these the seed isvisible at the open end of the aril. In N. alba (Weberbauer) and gladstonianathe seed iscompletely inclosed and hidden by the aril, while thearil of N. flavais a large, wrinkled bag, big enough,if stretched out, to
Structure.9icover two seeds with ease. Its cells (Fig. 41, c-e) are tubular andstretched out in the direction of itsgrowth, thin-walled, and somewhatpointed. In N.flava they are about three times longer and wider than inN. odorata and caerulea. The inner layer of cells adheres closely to theseed, and its cells are much smaller than those of the outer layer. Thewalls are supplied with fine pores (Weberbauer). The aril serves, by itsmucilage and inclosed air, to float the seed for a short time ;itdrops theseed in the course of a few hours (JV. tuberosd) or a day or so {N. caerulea),partly by being bursted and torn through absorption of water, partly bydecay (Fig. 41, b).The seed properisprotected by a firm shell consisting of two parts,the outer a single layer of thick-walled, close-fitting cells, the inner aFig. 41. The aril. N. odorata.- a, diagrammatic section of aril and seed ; b, portion of arilafter the seed is dropped ; c, cell of the aril where the outer wall bends In, In continuity withthe inner wall of the aril ; d, cell of inner wall ; e, cell of outer wall.few layers of thick-walled, but loosely placed cells. The latter layerisof the same thickness as the former ;its cells are much flattened tangentiallyand contain large tangential air-spaces ; the walls are lignified, withthe exception of a thin layer next to the lumen. The cells of the outerlayer are also strongly compressed tangentially (except in N. flava). Onaccount of the great induration, which is thickest on the outer wall, theyhave but a small lumen (Fig. 42). The walls are penetrated by porecanals.They are composed of lignin, excepting a thin layer of cellulosenext to the lumen and an outer thin cuticula which projects inward alongthe boundaries of the cells. In the cells there are (in N. alba) bluish-greengranules which give color to the seed. In surface view the outer cells arearranged in longitudinal rows, and, except at the micropylar end, they havevery sinuous walls. Over the raphe the cells are elongatedin the directionof the rows, but elsewhere at right angles to the rows. At the micropylarend of the seed the cells of the hard (outer) layer are nearly squareand in rows, but interrupted over the hilum, which lies (in N. alba) about
9 2 The Waterlilies.three cells from the micropyle. At the hilum the thick-walled cells of theinner seed coat are rounder and more closely packed than elsewhere.Based on the examination of the seeds of about twenty species ofNymphaea, Weberbauer distinguished four types of testa, as follows:(i) Surface bare, smooth, and shining: N. alba, Candida, tuberosa, odorata.(2) Bare, with longitudinal ridges;adjacent edges of rows of cells arealternately raised up or level ;the ridges may be interrupted atmiddle of seed {N. rubra (Fig. 42, a), pubescens).Fio. 42. Sections of outer cell layer of testa: a, N. rubra; b, N.fiavovirens(" N. gracilte" Weberb.) ; c, N. stellata. After Weberbauer.(3) The seed bears longitudinal rows of hairs in the position of the ridgesof the second type;the rows may be similarly interrupted (A 7 ,blanda,amazonum, gigantea, caerulea, stellata (Fig. 42, c), capensis, ampla,madagascariensis); N. lotus,hairy and show transitions to the second type.thermalis, and dentata are only slightly(4) Both longitudinal and transverse cell walls have short papillar outgrowths(N. Jlavo-irirens, Fig. 42, b).Differences occur in the degree of flattening of the outer cells andthickening of the walls, as also in the number of hairs and character ofthe surface under them. In N. tfiermalis the amount of thickeningis lessthan in other Lotos species, and in N. gigantea the outer walls of thecells are scarcely thicker than the lateral and inner walls. In N. rubra.the cells are but littlecompressed. N. amazonum has a dull, roughenedsurface beneath the hairs, but in "A^. blanda " (probably N. rudgeana) thisisvery smooth and shiny and the hairs are bent at the tip.N. ampla
Structure. 93and madagascariensis show an inclination to the formation of ridges as inthe second type, with a reduction of the hairs.The outer coat of N. flava is peculiar in several points. It is coveredwith long, slender, appressed hairs just visible to the naked eye,springing from all sides of the cells. It thus combines the charactersof the third and fourth types of Weberbauer. The cells are extremelywavy superficially, and so irregular in shape as to render the longitudinalrows obscure. They differ from those of all other species investigated inbeing nearly twice as deep as wide. A great amount of induration of theouter walls leaves the lumen as a small pear-shaped space at the innerend of the cell. This is a striking approach towardthe narrow columnar structure of the hard layer ofVictoria and other genera of Nymphaeaceae.I findthat N. tetragona belongs plainly to the first type of jseed coat according to Weberbauer, and N. elegans tothe fourth type.The inner seed coat ispressed to a thin layer,u '8'rib" tion fFlB -except ~ at the tip r of the nucellus. The walls are rvascular bundles from thethickened on the inner side, but as it approaches the ebaiaza, in seed coat of n.. . il ,,alba.After Chlfflot.micropyle the thickening extends up the side walls,until at the micropyle only the outer walls are unthickened. With thisthickening there is an increase in the height and sinuosity of the sidewalls. The micropyleis closed by slightlythickened rounded cells anda layer of thinner cells overlying the inner coat (cf. Fig. 44). A vascularbundle passes along the raphe, and spreadsin the seed coat irregularly(Fig. 43), as figured by Chifflot (1902).Nine-tenths of the space of the seed is occupied by perisperm(nucellar tissue). This consists of large, thin-walled polyhedral cells (Fig.44, p), densely packed with starch. The outermost cells of the nucellusare shriveled and compressed together into a thin membrane, which isthickest between the embryo and the seed coat. The outer food-storingcells are small and somewhat flattened, but in most of the nucellar tissuethey are elongated and disposed somewhat in lines {Leitungsbahnen ofWeberbauer) running obliquely upward toward an air-space beneath theembryo. The central axis of the seed is filled with looser, thin, elongatedstarch-laden cells. The nuclei of the cells, where distinct, are pressedover against one wall. The starch is in simple or compound granules,and these, in N. gladstoniana and odorata, are aggregated into sphericalmasses. The grains are very small, and the spherical masses are not
94 The Waterlilies.larger than single grains of some plants. Each spheruleis surroundedby a pellicle of protoplasm, and a network of the same substanceramifies among the constituent granules. This type of aggregationexists in a less perfect condition in N. jiava, but was not seen elsewhere.In N. odorata stained sections of the perisperm show a network ofthicker plasma strands between the starchspherules near the middle of each cell.In the micropylar end of the seed aFig. 44. Longitudinal section of seed :c, cotyledon ; e, endosperm ; g, tegmcn ;m, micropyle ; p, perisperm ; I, testa.smalllenticular or globose space (Fig. 44) isoccupied by the endosperm and embryo.The former is a single layer of cubical cells(e) inclosing the latter. The endosperm cellsare thin-walled, but Weberbauer was able tofind the walls covered with pits so that thethicker portions formed a fine network. Theyhave moderate nuclei, and contain a richsupply of proteids and oil, but no starch.Over the radicle the endosperm is very thinand poorin contents. The embryo has twoequal, fleshy, hemispherical cotyledons, excavatedon the inner side to receive theplumule. Between their symmetrical basesis the tiny caulicle, and a slight prominenceat the end of this is the radicle. Foodstoringtissue of isodiametric cells and elongated conductingcells aredistinguishable ; an axial strand of the latter is found in the caulicle,with a branch into each cotyledon. No starch is found in the embryo,but it is richly supplied in all its parts with oil and protein. In acrushed embryo of N. flava the oil globules run together in considerabledrops which stain easily with alcannin. The plumule consists of tworounded papillae, the leaf rudiments, placed transversely to the cotyledons.One of these is about four times as large in volume as the other.
CHAPTER III.DEVELOPMENT.ORGANOGENY AND EMBRYOLOGY.If it were not for the long hairs which clothe the apex of the stem ofNymphaea, there could hardly be a more favorable object for the study ofthe development of the organs of a mature plant. We cut off the broadtip of a rhizome of N. alba, odorata, or tuberosa i cm. back and pinit down to the wax bottom of a dissecting pan. Then with the aid ofa dissecting microscope the larger leaves and their stipule's are cut away.The hairs may be pulled out with fine forceps. This is the most difficultpart of the task. Finally one discovers that the stem apex is slightly basinshaped.At the center is a very minute elevation, the growing point, andaround it the rudiments of leaves and flowers are arranged in spiral order.The figures of leaf development given by Payer (1857) for N. alba areequally applicable to N. odorata, and would need but little alteration forN. rubra. I have examined the last two by dissection, and have studiedN. caerulea, odorata, and flava brieflyin serial microtome section. Theyoungest rudiments at the stem apex are mere rounded papillae, withoutindication of their future destiny. They gradually elongate and on reachinga length of 0.08 cm. the foliar or floral character becomes evident.The leaf rudiment (cf.PI. II, 1-7) takes on a more and more conical form,while the floral rudiment remains broad and rounded and develops theanterior sepal. A leaf of N. rubra var., 0.16 cm. long, is about three-quartersas wide at base, and tapers to a rounded apex on the inner;face abroad triangular depression is sharply marked off by two lateral ridgeswhich are continuous over the apex and curve inward a little below, endingabout one-fifth of the length of the rudiment above its base. Below thisis the petiole the ;upper partis the lamina. The rudiment becomeslonger and more acute, and the lateral ridges become slightly rolledinward until in a leaf 0.3 cm. long the former triangular depression hasbecome a narrow groove, rounded above. At the base of the petiole thereis now on either side a narrow rounded auricle, three times as long as
96 The Wateri.iues.broad, placed so that one end touches the rhizome and the other is a veryshort distance above. These are the stipules. In a leaf 0.64 cm. long thelamina is still more than twice the length of the petiole ; the inrolled sidesnow meet in the median line and the veins are prominent. The lobes ofthe leaf extend down along the front of the petiole and are slightly divaricate.The petioleis about as broad as long and somewhat contracted ateach end. The stipules are slightly larger and more nearly vertical. Thecoating of hairs on petiole, stipules, and lamina is well developed, and theswollen collar at the base of the leaf isbecoming evident. Subsequentdevelopment is, to outward appearances, chiefly a matter of size, exceptwith regard to the stipules.On a leaf of N. rubra whose petioleisalready more than 5cm. long these form lunate auricles, with nearly verticalline of attachment to the petiole,and a lobe extending upward a shortdistance above the line of insertion. Such a stipule is about one-third aswide as the diameter of the petiole and twice as long as wide. After thisit decreases in proportionate width, while greatly elongating to reach themature form.less than 0.16 cm. in length,In N. odorata (PI. II)and N. alba the young leaf rudiment,is more rounded and less acute than in N.rubra and the stipules are quite different. About the time the flatteningof the inner side of the rudiment to form the lamina occurs, two elongatedridges appear, one on either side of the base of the leaf, extending alongtoward each other just in front of the rudimentary petiole. The adjacentends soon meet in the median line and the beginning of the stipular plateof the Castalia group is evident. Beingshorter in the median line thanat the sides, its outline is obcordate. As the whole ridge grows higher,itovertops its leaf at the length of about 0.3 cm., and on either side of themiddle line an S-shaped fold occurs, which throws the median part of thestipules away from the leaf, or, to state it otherwise, forms a groove intowhich the leaf fits. A great thickening along these folds makes twostrong keels on the inner side of the stipule. Later this broad plateelongates to the typical adult form. A leaf rudiment oi N. odorata 0.3 cm.long had downward pointing hairs covering the back of the lamina and thepetiole, but on the next younger leaf these were present only on the lamina.The formation of the principal air-canals takes place at an extremelyearly period. The firstbeginning of separation of the cells was seen inN. caerulea in a rudiment 0.5 mm. long. The canals were seen with thenaked eyein a leaf rudiment of N. flava about 1 mm. long;the boundingwalls were composed of columnar cells. At 1.6 mm. longall of the canalsare indicated. In the smaller leaf the fusion of the three main bundles
CARNEGIE INSTITUTION OF WASHINGTON. WATERLILIES, PLATE II.A. KORONSKI DEL.NYMPHAEA ODORATA.1-7, development of leaf and stipules; 1, 2, front and back views of the same rudiment; 3, older;4, 5, front and side views of one rudiment; 6, 7, front and back views; 8, rhizome with roots andleaves trimmed off, natural size; 9, transverse section of rhizome, natural size.
Development. 97from the petiole to a single trace and the separation of this again into thecentral and two lateral traces was seen. The bundles were chiefly composedof procambial cells, though some spirals were presentin the threebranches within the inner fork. The lamina of the older leaf mentionedhad elongated cells along the lines of the veins, but the parenchymawas almost totally undifferentiated and was without intercellular spaces.A leaf of-iV. tuberosa richardsoni whose lamina and petiole are each 1.3 cm.long has the air-canals well developed along the midrib. Between theveins the mesophyll consists of five layers of cubical cells arranged in verticallines from the upper to the lower epidermis. The layer next to eachepidermis is continuous, but between the vertical rows in the three middlelayers long intercellular spaces are present. Idioblasts are already presentnear the midrib. A leaf of N. odorata with lamina 2.5 cm. long andpetiole 3.2 cm. has the air-chambers of the mesophyll well marked off byplates of cells ;the palisade consists of a single layer of dense cubicalcells, underlaid by two or three layers of flat cells elongated parallel tothe surface of the leaf ; the mucilage hairs are largeand functional. Whenthe lamina is 8 cm. long the palisadestill consists of a single cell-layer ;the big idioblasts have not yet appeared. Stoma-mother-cells are presentin full number and nearly mature size ; some, indeed, have just divided toform the guard cells. They are round, granular, and highly refractive.The maturation of the stomata occurs only a day or two before the leaffinally expands. Caspary (1854; 1858), in announcing first the apicalgrowth of leaves, showed that this apical growth continues longer inNymphaea than in other plants, and that the margin of the leaf in allNymphaeaceae is the latest part to complete its development.The tissues of the stem begin to differentiate within 0.25 mm. of thepoint of apical growth. No pleromeis at any time distinguishable. Thelacunar cortex acquires intercellular spaces while all of the tissues are stillmeristematic, and the early establishment of desmogen strands, adventitiousroots, and external hairs has already been referred to.Concerning the development of the flower, we are able to corroboratethe work of Payer (1857) and Goebel (1886) on N. alba by observationson N. caerulea and lotus, and to add some details. The discussionof important morphological problems will be deferred whenever possibleuntil the observed facts are set forth. The earliest rudiment of a flowerthat I have seen was a smooth, thimble-shaped upgrowth at the stem apexin N. caerulea. On such a papilla, according to Payer, there appears inN. alba the rudiment of the anterior sepal then the two lateral ; sepals
98 The Waterlilies.arise on the sides, and later the posterior sepal. Hence comes theirpeculiar aestivation. Four papillae next appear alternate with the sepals.They arise simultaneously, grow rapidly, and are always larger than theinner floral organs, which originate later. They are at the base of a hemisphericalreceptacle whose surface, naked hitherto, becomes covered littleby little from base to summit with a number of new papillae. These arearranged in ten vertical rows, one abutting on each petal, one on eachlateral sepal, and two on the posterior and anterior sepals. The lowestones soon flatten out and become petals the ;uppermost become typicalstamens ;between these two sets one finds all gradations from stamen topetal. The stamen rudiments, however, do not cover the whole receptacle.A bare space remains at the summit, around the edges of which arow of papillae appears. These (the carpels) become horse-shoe shaped,and their inner edges join, making a sinuous line. The sinuosities extendsomewhat toward the center, and a thick process of the receptacle growsup to fill the central space, closing the ovary-cells. Now the sepals andthe four outer petals remain stationary, as it were, while the rest of thereceptacle elongates, carrying with it the carpels, stamens, and innerpetals. Thus the ovary comes to be inferior with regard to the stamensand inner petals while the outer petals and the sepals have an inferiorinsertion. There is no fusion of the floral leaves outside the carpels withthe ovary or receptacle, as Koehne {fide Goebel) believed.According to this account from Payer, the ovary would bear thecarpels at its summit, and its lower parts would be of axial origin.Goebel(1. c.) noted this difficulty and showed that before the carpels are mappedout there is a hollowing of the receptacle as is common in inferior ovaries.From the edge of the axial excavation, but reaching to the bottom of it,arise the carpel-rudiments. They are never free from the receptacle onthe outer side. The whole wall of the ovary-cells, however, is of carpellaryorigin, and no part is formed from the vegetative apex of the flower stalk.Our own observations on N. caerulea and lotus support Goebel's view.In N. lotus the time relations of the origin of the outer floral organsdiffer slightly from that given by Payer. In the apex of a large floweringplant we found three rudimentary flowers of successive ages and sizes, inall of which only the anterior and lateral sepal-rudiments were present(Fig. 45, a, b).This shows that these organs originate much earlier thanthose farther in. The anterior sepal was much larger than the others, andwas inserted considerably farther down. In such a rudiment one couldhardly speak of a peduncle, for this sepal seemed almost sessile on the
Development. 99parent stem. The next older flower noted (Fig. 45, had c) a broad papillarepresenting the posterior sepal, a rounded one for the right antero-lateralpetal and a partly formed papilla for the left antero-lateral petal. Theright petal was about half as large as the posterior sepal, and this wasabout half as large as a lateral sepal, and this again half as large as theanterior sepal. The next flower had the two anterior petals about equallydeveloped. The remaining organs (except the carpels) originate oneby one ; two postero-lateral petals come next, then four petals opposite thesepals then four opposite the first petals. The exact order of these was not;determined (Fig. 45, d).In early stages the bud exceeds the peduncle inlength, but after the total length of the two reaches 2.5 cm. the pedunclebegins to exceed the bud. Previous to this stage the anterior and lateralFio. 45. Development of flower, N. lotut var.; a, end view; b, side view ofvery early stage ; c, with rudi ments o f sepals and two anterior petals ; d, sepals,two whorls of petals, and one member of the next whorl laid down.sepals alone cover the bud ; later they are pushed apart and the posteriorsepal is exposed. The carpels are rather late to develop. In a bud 1 cm.the back with thelong they resemble little folded leaves, fused half way uphollowed receptacle, but open along the ventral suture, and, even in N.free from each other.The above facts throw light on several questions, and firstupon thelotus, whollykind of inflorescence found in Nymphaea.It will be remembered that theflowers stand in place of leaves on the stem, but without any subtendingbract. Payer says the case is very simple : Eachflower arises in the axilbe atof a leaf, but on account of crowding in the bud, the flower comes tosome distance from its leaf, and itrequires some experience ("il fautquelque habitude ") to find the leaf to which a given flower belongs.Schumann (1894) maintains the same view and cites the case of Victoria,in which there are parallel spirals of leaves and flowers on the stem, andat the apex each leaf has a flower to its left as they originate. Raciborski(1894) opposes this with the theory that the bract is entirely suppressed inNymphaea and Victoria. This better accords with the exact location of theflowers in the leaf spirals;and inNuphar there is at times, if not always, arudimentary bract subtending the young flowers.Eichler (1875), however,
iooThe Waterlilies.on the authority of Caspary and Braun (in personal letters), reluctantlyaccepts the belief that the flowers are axillary, but the bract and bracteolesare carried up on the summit of the peduncle as anterior and lateralsepals. The argument for this has been based largely upon the uncertainground of teratology. Planchon (1850, e) figured a flower of N. caerulea" whose peduncle showed accidentally [at the lower end] three spatulatebracts, with petioles wavy here and there on the margins as the sepals areat base." * Caspary {fide Eichler) found in the same species a flowerwith the anterior sepal at the base of the peduncle as a ribbon-shapedbract, the lateral sepals also pushed down, and the posterior sepal in itsproper place. This was followed by four sepals in a whorl, placed diagonallylike the second whorl of a normal flower. Caspary found similarcases in JV. alba, gigantea, rubra, and hybrids of M. caerulea and capensis.My own observations on the development of the flower of N. lotus fullysubstantiate Caspary's view, and, I may say, completely turned me froman opponent to an adherent of his theory. The low insertion of therudiment of the outer sepal and itsvery early appearance,followed at onceby the two lateral sepals, are convincing. This also explains the peculiarrelations of the sepals in the bud ;on any other theory, except a blankdisplacement, the anterior sepal should be, like the posterior, covered bythe lateral sepals.It remains, however, to explain the presence of a single posterior sepaland the oblique position of the next four floral leaves. Caspary and Braunlooked upon the single sepal as an " Erganzungsblatt. " But if the floraldiagram of Nuphar as given by Baillon (j : 82) can be depended on, inwhich the outermost of the five sepals is posterior and the petals " areinserted along a spiral," we have the most perfect starting point for thedevelopment of the flower of Nymphaea. The outermost sepal of Nupharhomologizes with the posterior sepal of Nymphaea, and a very slightshifting of the remaining four sepals away from the median anterior linegives them the oblique position of the four outer petals of Nymphaea.The strongly sepaloid character of these petals, especially in Brachyceras,indicates such an origin. As to the inner petals, from the acycly ofNuphar, Nymphaea shows a continually increasing tendency to cyclicsymmetry. In N. lotus the rudiments of all of the floral leaves (exceptingthe three outer sepals) arise in series, one after another, but Payer'saccount ol N. alba places the four outer petals in a whorl, and the inner*dont le pedoncule offre accidentellement trois bractees spathulees, a petiole ondulec_a-et-la sur les bords, comme les sepales le sont a leur base.
Development.ioipetals and the stamens in alternating whorls of five ;probably this plantshould be worked over anew. In the mature flower the cyclic arrangementis least evident in Castalia and but little more so in Lotos ; but, asstated, in old flowers of Brachyceras, and open flowers of Hydrocallisitseems to prevail in all of the parts (cf. Fig. 36). Probably cyclic arrangementisalways to be considered as derived from acyclic, and Nymphaeaseems to be in the transition stage. For such flowers the term hemicyclicor spirocyclic (Engler, 1898)is best. Itgoes beyond the truth to placeNymphaeaceae among acyclic orders as Goebel did (1887, p. 412).While stillmaintaining that the terms sepal and petal as commonlyused for waterlilies are physiologically correct, the morphological value ofthe petals must be considered. We have shown that the anterior sepalrepresents the bract, the lateral sepals the bracteoles, and the inner sepalwith the four outer petals the primitive sepals. For the last homology Iam indebted to a suggestion from my instructor, Dr. John W. Harshberger.In development no distinction can be made between the rudimentsof the remaining petals and the stamens, while in N. alba the fourouter petals do have a distinct and simultaneous origin. Payer, therefore,pointed out, and rightly so, that the inner petals are really modifiedstamens, and the waterlilies are habitually double flowers in the samesense as roses, almonds, etc., are doubled in cultivation. This at onceexplains and issupported by the extremely variable number of petals andthe occurrence of transitional forms between petal and stamen. It seemsprobable, however, that in last analysis all petals are modified stamens,and in Lotos and Anecphya, where a naked space occurs between theinsertion of stamens and petals, and intermediate forms rarelyif everoccur, the distinction between the two classes of organs is being rapidlydrawn, and the homologyis more distant than in Castalia. The fixationof definitive petal and stamen characters upon the floral rudiments istaking place before our eyes in the genus Nymphaea.It is striking, buton second thought only natural, to find this kind of specialization going onalongside of the change from spiral to cyclic symmetry. The two phenomenaare but manifestations of the same law of adaptation and differentiationin form and function.After the stamen rudiment isestablished itgrows considerably beforethe archesporium appears. It has been claimed that the anther cells areat first extrorse and are pushed around to the inner side by unevengrowth of the connective. In N. lotus, at least, this is not the case. Twopairs of archesporial lines are differentiated in the substance of the anther,
102 The Waterlilies.each consisting of a single straight row of cells.They are removed fouror five cells from the inner epidermis and seven to twelve cells from theouter epidermis, and between each pair of lines eight cells intervene. Thecells immediately surrounding the archesporia take on a more or lessconcentric arrangement, and the development goes on in the usual wayfor flowering plants. In N. caerulea the outer stamens develop slightlyinadvance of the inner ones ;where the former are 1.3 cm. long and thespore-mother-cells are already separate from one another and becomingrounded off, the mother-cells of the inner stamens form a continuous tissue.The ovules originate after the ovary has attained a considerable size,say 0.5 cm. in diameter in N. caerulea. At first a tiny rounded columnFig. 4. Ovule of N. odo- Fio. 47. Embryo sac of N. odorata; longitudinal section of ovule at time ofrata, from a half-grown bud, flowering. From a photomicrograph,showing archesporium.From a photomicrograph.projects into the enlarging ovarycell. This swells at the end into aknob, an integument of two cell-layers grows up around the knob, andthen a second integument outside the first. When the inner integumenthas reached the height of the nucellus, the outer one is about half as high(cf. Fig. 46).Meanwhile the knob has turned over at right angles to thesupporting column in its change toward anatropy. A large archesporial cellis now visible at the apex of the nucellus, covered by a layer of cells. Thearchesporial cell presents five times as large a surface in longitudinal sectionas the surrounding cells its nucleus is as; large as an entire cell ofthe nucellus, is rather poor in contents, and has a single dense nucleolus.Its further development has been described for N. odorata by Cook (1902).According to his account, a tapetal cell is cut off at the outer end of thearchesporial cell (cf. Fig. 46), and from the former several tapetal cellsarise by a variable series of divisions. Three potential megaspores arenext cut off from the mother-cell. These rapidly degenerate, leaving a
Development. 103single large megaspore. This cell elongates to form the embryo-sac.Returning to the stage in which only the archesporial cell is found, we notean elongation of cells in the center of the funiculus which foreshadows thevascular bundle. This was seen in an ovary of N. odorata 0.6 cm. indiameter ;the ovules were just visible to the naked eye. In an antherfrom the same flower the microspores are formed, but have not receivedthe chj -acteristic markings on the exine ;the tapetum appears as a layerof disintegrated matter around the walls of the anther cell ;the thickeningsare not yet laid down in the anther wall. After this the ovule rapidlygrows to its mature form. The outer integument becomes three cellsthick and loses itsprotoplasm, while the inner integument consists of twolayers, of which the inner alone remains richly protoplasmic. The nucellusconsists of many cells arrangedin radial rows the outer ones are;tangentially flattened, the central ones rounded. The rapheisprominentand is traversed by a slender vascular bundle with a spiral trachaea on theside next to the nucellus. The funiculus is covered at base with a glandularepidermis like that of the ovary, but the glandular cells end at a sharpline near the seed, beyond which the epidermis is plain and smooth.Between the epidermis and the vascular bundle the round parenchymacells enclose large intercellular spaces. The embryo-sac (Fig. 47)isprominent as a clear area below the micropyle, and is shapedlike a flaskwith the neck extending down into the axis of the seed. The ratio of thediameter of the body and neck of the flask is as 5 to 1 in N. odorata andzanzibariensis, but in N. lotus the whole ovule islonger and narrower, andthe embryo-sac is lanceolate in longitudinal section, with rounded ends.A single layer of columnar nucellar cells covers its outer end.A day or two before the flower opens the egg apparatus is complete.The embryo-sac is slightly narrowed at the micropylar end, where it iscovered, as before, with one layer of columnar cells. In this upper partof the sac three similar cells are crowded, two above and one below. Therest of the sac is lined already by a few cells around the sides, visibleonly in my preparations as compressed nuclei. Hofmeister (1858) doubtlessreferred to the same thing in Nuphar, " whose embryo-sac," he says,"even before fecundation shows clearly a cellular wall" (Zellstoffhaut).A pair of nuclei indistinctly seen in the extreme end of the narrow innerpart of the sac in N. odorata doubtless represent the antipodal cells; a singlecell with very large nucleolus occupies the same position in N. zanzibariensis.The outer integument now consists of two (or near the raphe three) layers
104 The Waterlilies.of slightly flattened cells, poor in contents. The inner integument hasboth of its layers very thin, and the outer one nearly empty.Dahmen (1892) criticallyexamined the funiculus of the ovules ofon theNymphaea. Since the so-called aril originates as a swellingfuniculus before fecundation occurs, he considers that it is not an arilin the usual sense, restricting the use of the term to outgrowths whichoccur when the seed is ripening. The distinction, however, isextremelyunimportant, especially as the aril of Nymphaea does not reach maturityuntil the seed is ripe, and most of its growth occurs after the seedhas attained its full size. In N. lotus Dahmen noted two swollen zoneson the funiculus near the ovule, but only one in N. rosea [quid?) andzanzibariensis. The epidermis of the funiculus has slightly thickenedwalls ;within it is a thin-walled parenchyma and a vascular bundle withphloem and a little xylem. 'The bundle has no connection with the aril.Small round starch grains are found temporarily in the parenchyma,epidermis, and phloem. Intercellular spaces occur onlyin the arillarswellings according to Dahmen, but I find them plentifully elsewhere inN. odorata. The tissues are all filled with mucilage, which is only precipitatedby strong alcohol. The protoplasm is dull yellowish. Asparaginoccurs in the funiculi, but no sugar was found.Unfortunately we cannot accurately trace all of these tissues to theirmature state, but the changes are not important in any part except theembryo-sac. The early stages were worked out by Hofmeister (1858) inNuphar luteum and advena, which differ from Nymphaeain that the narrowinner part of the embryo-sac is much longer, and extends after fecundationto the chalaza ;an axial tube remains open even in the mature seed(Weberbauer, 1894).In Nuphar there are three cells at the top of theembryo-sac, two upper synergidae and an ovum below. About the timeof fecundation a transverse wall divides the narrow part of the embryo sacfrom the upper wide part, and the latter is soon filled by two or threeendosperm cells. The fertilized ovum now divides transversely, and asuspensor of one to five cells is formed above the embryo. The observationsof Mr. Cook (1902) on N. odorata seem to be corroborated up tothe first cell-wall laid down in the embryo-sac. His views on the developmentof the embryo, however, which he considers to be monocotyledonousand without suspensor, are totally at variance with my own preparations.I cannot imagine that his interpretations are correct. This portion of Mr.Cook's paper is based chiefly on Nuphar advena, in which Caspary'sdrawings (inHb. Berlin) seem also to indicate the absence of a suspensor.
Development. 105The earliest fertilized ovule of Nymphaea that I have seen is ofN. lotus, and has already reached nearlyits mature size. The integumentsare considerably hardened, and the perispermis loaded with starch.A broad space at the micropylar end of the seed is occupied by the endosperm,the cells of which are very thin and watery (Fig. 48, a).A distinctlayer of perisperm still bounds the endosperm on the side next to theseed c-^at. From a point near the micropyle a row of three cells projectsinto the endosperm, representing the embryo. An older seed of the samespecies shows the endosperm occupying an approximately spherical space,somewhat pointed above and flattened below ;its cells are large andFio. 48. Embryology : (a), (b) successive stages, N. lotus; (c) sphericalembryo with suspensor, N. odorata ; (d) early differentiation of cotyledonsand plumule, N. caerulea.distinctly nucleate, but very thin walled and poor in contents. Theembryo (Fig. 48, b) consists of a line of three suspensor cells and aterminal knob of probably eight cells (four in longitudinal section). Theuppermost suspensor cell is hexagonal, the others square (in section).After this the endosperm enlarges very little. As the embryo encroachesupon it, the outer cells become denser and more regular until they reachthe mature condition, and the inner ones are crushed to a thin layer ofremnants between the permanent outer layer and the cotyledons. Theembryo proper (cf. Conard, 1902) becomes accurately spherical as itenlarges, until it consists of some hundreds of cells (Fig. 48, c).Then thelower end becomes flattened by reason of the outgrowth of cells on eachside of the lower pole. In section the embryo now appears triangularand almost equilateral. The two lobes increase symmetrically and formthe two equal cotyledons. At first these spread out nearly horizontally(Fig. 48, d), while in the middle between them a slight cone-like papilla
io6The Waterlilies.represents the plumule, and a similar prominence opposite (next to thesuspensor)is the radicle. Subsequently the cotyledons spread round thewalls of the endosperm and fill up all of the limited space available, whilebetween them the two leaf-rudiments of the plumule are differentiated.GERMINATION AND EARLY GROWTH.Corresponding with the habits of life of the various species of Nymphaeaare certain requirements for the proper growth of seedlings. Seedsof the Eu-castalia group quickly lose the power of germination if dried in;nature only those which are, by chance freshets, stranded hopelessly abovethe normal water level ever become dry. N. tetragona can withstanddrought fairly well. Seeds from Kashmir communicated by Mr. Wm. Gollan,of Saharanpur, India, germinated quite well, but of several sendings receivedin the early spring of successive years from Japan, only one seed has eversprouted. The former were sent to us directly after they were gathered,while the latter have doubtless been keptin a seed-house since theprevious summer. Seeds of the Lotos and Brachyceras group can withstanda great deal of drought they are accustomed to it but ; ; they arekept much more successfully, as Caspary (1877) recommended, packed inand air dried. Under these conditions a considerable amount ofclaymoisture is retained, even when the mass appears to be perfectly dry.N. zanzibariensis was introduced into Europe from Zanzibar by seedspacked in chalk ;and of a box of seeds of N. dentata, mingled with soil, receivedlast summer (1901) from Jamaica, nearly every one sown germinated.In all of the Castalia group the seeds should be keptin bottles of water.It was thus, also, that Victoria regia was successfully carried to Englandafter a long series of costly failures.For germination the seeds of waterlilies need to be submerged in 5to 30 cm. of water. Those of the Castalia group prefer a temperature of15 to 18 R., but the tropical species do best at 23 R. (Siber, 1883).Avery irregular length of time is required. Of a quantity of N. elegans Xzanzibariensis seeds sown as soon as collected, a large number came upin two weeks after more than a;year some earth from this pot chanced tobe placed in a warmer tank, and numerous new and vigorous seedlingsappeared. In the Dreer Gardens at Riverton, N. I J.,have seen floweringplants of N, caerulea which came up spontaneously out of doors wheretwo years previously that species had been grown the seeds had lain;dormant over all one summer. In this case Mr. Tricker informed me thatthe bottom of the pond had been dug over each spring. Probably the
Development. 107seeds were buried in the first digging and brought to the surface in thesecond. In midsummer N. zansibariensis seeds will germinate quicklyafter maturing, and the bags of seed must be gathered as soon as the fruitbursts ;but in the summer of 1900 numerous seedlings of this plantsprang up in the pond of the Botanic Garden (University of Pennsylvania)and blossomed before frost ;they must have come from seed ripened inthe previous autumn. Mr. Waters (1886) gives an interesting account ofthe germination of N. odorata. Seeds gathered in the fall of 1 883 weresown at once in an aquarium keptin a living room with north light. InMarch, 1884, many were germinating. In June, 1885, the vessel was placedwhere it received direct skylight with sun for an hour at noon a half;dozen new plants appeared in August. In the fallthey were returned tothe living room, and about Christmas another seed had germinated.From February to April, 1886, four others sprouted these had therefore;lain dormant for about two and a half years, although they were constantlyin conditions which would permit germination.The general features of a seedling Nymphaea were noted by Tittmannin 1821. Treviranus, Caspary, Klebs, Goebel, and others have sinceremarked upon or figured such seedlings. The first visible evidence oflife in the seed is the loosening of an operculum at the apex (Fig. 49, 8, 6).This is a conical body bearing the micropyle and hilum its;edge is raggedby reason of the sinuous margins of the cells of the testa. The operculumispushed aside by the protruding embryo and its wrappings, and lies onthese at one side near the edge of the opening from which it was torn.The seed coat is also rent with short longitudinal fissures, dividing the lipof the aperture first formed into numerous acute teeth (Fig. 49, 8).Asthe embryo emerges it is covered by a white sheath produced by theswelling and straightening out of the overlying " rumpled up " portion ofthe inner seed coat this ; protects it from the sharp and ragged edges ofthe testa. The cotyledons remain permanently within the seed and showno changesin the inner parts exceptin the emptying of the cells. It is bythe elongation of the bases of the cotyledons into a kind of petioles thatthe radicle and plumule are passively carried out of the seed. These cotyledonarypetioles also curve geotropically downward until the caulicle isa vertical position, and then they cease to grow (Fig. 49, 3, 4).Through all this time the plumule and radicle have remained dormant,but now both start into activity, the former very rapidly. The hypocotyldoes not elongate at all. The epicotyl and first leaf shoot upward as aslender subulate body (Fig. 49, 3, 4).The elongation of the epicotylin
io8The Waterlilies.Fig. 49. Seedlings of a hybrid Nymphaca of the J,otos group. 1, tuber formation at summit of epicotylwell advanced. 2, the first broad leaf expanded, the primary root, with root hairs, mature. 3, the first (filiform)leaf nearly mature, and rhizoids springing from cotyledon. 4, a slightly older plant, from a seed which wasburied more deeply in earth ; the epicotyl is much elongated. 6, first secondary root. 6, small ripening tuber ;epicotyl rotted off. 7, beginning of second secondary root and third broad leaf (4th leaf). 8, soon after germination; the tegmen (dotted) extends beyond the hard seed coat, bearing the detached operculum. 9, plant letof same aye as 3, with cotyledons broken off from their petioles, a, first (filiform) leaf ; o, operculum.
Development. 109depends upon the depth to which the seeds are buried it reaches ;up justto the surface of the soil. If they are uncovered at the bottom of thewater, the epicotyl is scarcely visible (Fig. 49,it is3) about as long as;the width of the cotyledonary petioles. In other circumstances this internodemay be 2.5 cm. long (2, 4, 5).The stimulus which effects thisdifference is light.Seeds of N. gladstoniana and flava grown in glassbottles of water in the dark sent out very long epicotyls reaching farabove the layer of seeds in the bottom of the bottle. The first leafproduced by the seedling is a slender awl-shaped object, like a spear ofgrass (Tricker, 1897) ;it looks like a prolongation of the epicotyl (Fig. 49, (a);rudiment of thethe boundary between the two is marked by a tinynext leaf. While these changes are going on in the plumule, a large tuftof root-hairs is formed on a broad prominence at the base of each cotyledonarypetiole. These hairs serve to anchor the plantlet to the substratumand doubtless to absorb some food during the inactivity of the radicle(Goebel, 1893). After the subulate leaf has nearly reached maturity, andthe second leaf-rudiment is swelling, the radicle begins to elongate (Fig.49, 4).Hitherto it has been a tiny papilla between the bases of the cotyledons,but it soon grows out into a primary root 5 or 8 centimeters long (5).The second leaf of the plantlet arises opposite the first and developsa petiole and an ovate to linear lamina (2, 5).At its base a single adventitiousroot comes out and descends into the earth ;itonly becomesevident after the leaf is nearly mature (5). Subsequent leaves are broaderand broader in the lamina, becoming deltoid in Lotos and cordate tonearly orbicular in Brachyceras (cf. Fig. 50). Several of them remainon short petioles and are typically aquatic, but finally small floatingleaves are sent up. Each one bears one or more adventitious roots atthe base. The petioles of all the earlier leaves are winged and claspinguntil later.at the base ;no stipules are developedThe plantlet being thus pretty well established, the primary rootceases to function and soon dies away. As the nutriment is baled outfrom the seed, the central portions of the cotyledons are first emptied,then their outer parts, and then the endosperm. Finally the solution ofthe perisperm begins at the end next to the embryo and works steadilyback until the supply is exhausted. The nutriment furnished to the youngplant, therefore, is at first highly proteinaceous, and at the last whollycarbohydrate. The cotyledons remain of the same size as before germination,or very nearly so. When the seed has been emptied of its food, theepicotyl shrivels and dies as the primary root did before it (Fig. 49, 1, 6).
noThe Waterulies.The internodes above the epicotyl are not developed, but the stemgradually thickens and enlarges into a miniature tuber. In nearlyall of thegenus the plant is now essentially mature it needs; only to enlarge. Butin Eu-castalia a marked change takes place. The internodes after a timeFio. 60. Seedling leaves :(a), N. amazonum ; a, first leaf ; b, second (first broad) leaf ; c, third leaf ;e, epicotyl ; p, primary root ; si, first secondary root ; (b). If. catrulea ; 1. 2, 8, 4, first and following broadleaves ; (e). first floating leaf, X. odorata giyantea ; (d), If. lotus var., 1, 2, , 4, first and following leaves, thelast very young ; (e), N. zanzibariensis ; 1, 2, 8, first and following broad leaves ; If), If. tetroyoiia, 1, 2, 8, firstand following broad submerged leaves ; 1 fi, first floating leaf. Twice natural size.regularly, but slightly, elongate, and the stem suddenly becomes plagiotropicthus a ; tiny horizontal rhizome isproduced. From this pointonward maturityissimply a matter of size.In structure the characteristic features of the order are evident at avery tender age. While the primary root is still a mere papilla,the rootcapconsists of a single layer of cells, and behind it are three to four close
Development.i i ilayers of meristem cells ; immediately back of these, large intercellularspaces abound. The epidermis of the primary and early adventitiousroots differs from all mature rootage in having a copious supply of roothairs(Fig. 49, 2, 5).It will be recalled that root-hairs are generally saidto be wholly lacking inNymphaea, Nuphar, Euryale, and Victoria. As wehave observed them in N. dentata, lotus, caerulea, zanzibariensis, amazonum,elegans, odorata, capensis, and hybrids, they are doubtless universal in thegenus. On one occasion also they were found on early roots of a plantspringing from a tuber of N. caerulea. Victoria lacks them even on theprimary root. The epidermal cells of the early roots (in N. dentata) arelong and rectangular in surface view, and about as deep as wide where;from cubical cells. No shedding ofroot-hairs occur, however, they springepidermis occurs in these roots. The root-cap is long and thimble-shaped.Inside the epidermis there are in the primary root about six layers ofthin-walled cortex cells and a small (diarch ?)vascular bundle. Theadventitious roots are like this, but successively larger. In the earliest ofthese the diarch bundle is quite plain and is surrounded by a distinctendodermis;the two xylem patches are continuous across the middle ofthe bundle. The appearance of the central vascular strand of the hypocotyland lower part of the epicotyl is exactly the same as this, but it isdescribed here as " two opposite bundles with their xylems confluent "(cf. Gwynne-Vaughan, 1897). A short process of procambial cells extendstoward the base of each cotyledon, but no vascular tissue extends intothem. They are composed of elongated thin-walled cells, and have arather narrow insertion. The bundles of the stem are turned with theirsides toward the cotyledons,i. e., a line connecting the bases of the cotyledonswould pass through the confluent xylem and between the phloemmasses. Higher up in the epicotyl the two bundles become more distinct,and each, instead of being semicircular in section, is nearly round the;xylems are still in contact, though torn asunder to leave an irregular aircanalin the middle. The first two leaves are placed opposite the phloemsof the two bundles and at their point of insertion the vascular systemspreads out toward them on each side. In close succession the bundlesof the first adventitious root, the acicular leaf, the first broad leaf, and thesecond root and broad leaf join the widened portion of the system, withanastomoses across from bundle to bundle. Each of the above-namedorgans has a single bundle. No central strands exist above this, but thebundles spread out through the now fleshy stem on their way to thevarious organs. The communication from vessel to vessel which easily
ii2The Waterlilies.occurs in ordinary plants by reason of their proximity is made possible inthese thick stems with widely separated bundles by the profusionof crossbranches.Gwynne-Vaughan (1897) noticed in the center of the youngstem of N. zanzibariensis a cylinder of meristem tissue suggesting a stele,in which later desmogen strands were differentiated. He evidently consideredthis as furnishing a kind of transition from the "monostelic"epicotyl to the " astelic " stem.The epidermis of all parts of the plant above the cotyledons, exceptingthe upper surface of the leaves, issupplied with mucilage hairs.In N. Jlava these are so numerous on the acicular leaf as to make a plainlyvisible coating of jelly all over it, as is seen on petioles of Brasenia. Thehair-bases remain on the full-grownparts as in mature plants. The innertissues are very loose throughout.In the epicotyl the intercellularspaces are often as large as therounded cortex cells. The acicularleaf is similarly constructed. Thefirst broad leaf of N. dentata hasFio. 61. First broad (submerged) leaf of N. lotus seed- r 11 j1 1 1 . ,lin?: a, stoma In upper epidermis; b, portion of tip oftOUr Well-aevelOpeQ air-CanalS in tUeleaf, showing distribution of stomata by dots.petiole ;the second has six canals,which are at first of almost equal size. The lamina of these leaves isextremely thin and fragile.Stomata are found scattered over the uppersurface in N. lotus (Fig. 51), caerulea, elegans, amazonum, zanzibariensis, andcapensis, but I failed to discover any in N. odorata or rubra ; probably theobservation is at fault, for they are at best rather difficult to see. The bestsuccess was had with material fixed in chrom-acetic acid and mounted inbalsam;probably Wachter's (1897) chloral hydrate method would proveequally effective. The palisade-layer of these leaves is reduced to a singlelayer of hemispherical cells against the upper epidermis. Between theseand the lower epidermis there are branching partitions two cells high,enclosing large air-spaces (Constantin, 1886). The venation isvery simple.Between the bases of the second and third leaves a few long protectivehairs appear as on the adult stem-apex. The young stem is at firstdevoid of lacunar tissue ;its fundamental parenchymaconsists of roundedcells in which starch is rapidly deposited. From this time on the plantisin a position to go into a state of rest on any emergency.
CHAPTER IV.PHYSIOLOGY.Special observations on the physiology of waterlilies are not numerous.That such plants have the usual "senses of direction " geotropismand heliotropism goes without saying.The primary and adventitiousroots are all alike highly sensitive to gravitation at first, but the latterand run obliquelysoon become more or less passive downward, while thesecondary and tertiary roots ramify throughout the soil from its surfaceto a depth of 30 cm. or more. The peculiar type of root spoken of on aprevious page as " contractile " exhibits a striking physiological adaptation.Such roots have been found only on young plants of the Lotos andapocarpous groups, excepting N. inflavo-virens, which adults also havecontractile roots. In all of the plants mentioned, however, resting tubersare produced at the close of the season. The object evidentlyis to drawthese tubers well down into the earth for protection against drought andherbivorous animals.For the adult plants, with the one exception named,perish when the flowering season is over, and consequently are not suppliedwith any special contractile organs. N. flavo-virens y^zanzibariensisbears a few contractile roots on the lower part of the tuber of adult plants.This hybrid usually has persistent tubers as in N. flavo-virens;its contractileroots, however, show none of the peculiar structure which characterizesthose of N. flavo-virens. As a rule it is the lowest (oldest) root fromeach of the older leaf-bases which is contractile ;in N. lotus one or twoupper roots occasionally contract. This lowest root is strongly geotropic,while the others are rather feebly so. It also persists in a living conditionafter all of its companion roots have completely rotted away. After contractionit becomes fusiform in shape. The greater density of the cortexin contractile roots indicates that this is the layer where contraction takesplace, probably, as Rimbach (1902) has shown for liliaceous plants, byshortening and widening of the cells. Thus we see a strong differentiation,apparently greatest in N. flavo-virens, into nutritive and mechanical roots.In stems the great density of tissue in all but the Castalia group, andespecially in resting tubers, has already been shown. This is distinctlyrelated to the storage of starch which takes place so largely in such plants.IX93
ii4The Waterlilies.Iknow of no experiments upon the geotropismof stems outside ofthe Eu-castalia group. Among these plagiotropism is very pronounced.A stem of N. odorata 3 cm. in diameter and 10 cm. long planted uprightin a pot in early spring had turned its apex to a horizontal position bymid-summer.Young petioles and peduncles of different species turn through 90in the course of twenty-four hours ifplaced in a horizontal position. Thistendency is, of course, well developed in all species. The leaf-stalks andflower-stalks, however, are often so slender that their positions are mostlypassive, except at the ends. The buds in Castalia, and the leaves in allspecies, by their buoyancy make their wayunless opposed by other forces. We have alreadycoming up through the water has its midrib in a straightto the surface of the waternoted that the leaf inline with thepetiole, and on reaching the air the petiole bends over at the top nearlyat right angles to the lamina (Frank, 1872). That this is not a purelypassive condition is shown by the stiffness and turgidity of the organs.Also, when plants are moved close together or placed in small tanks,where the upper parts of the petioles are thrown out of their formerdirections, they bend the lamina down until it is more or less submerged.In the course of twenty-four hours, however, the leaf isagain lying flatupon the water surface. A corresponding bending has taken place in theupper part of the petiole. This bending may be due simply to readjustmentsof a mechanical kind in the elastic collenchyma of the petiole, or itvital action. In the latter case it is relatedmay be the result of a distinctlyto the phenomenanext to be described.It has been noted that the " length of the petioles and pedunclesdepends upon the depth of the water in which the plant is growing."Not onlyis this true for plants in different localities, but a given plant,if moved from shallow into deeper water, where all of its leaves are submergeda few centimeters, will be found on the following day with allits leaves floating in the normal manner. Plants moved from deep toshallow water need no adjustment exceptin the angle between the petioleand lamina ;but leaves produced afterward have petioles only long enoughfor the new conditions. Now, it is certain that throughout the life of a leafits petiole is continually growing longer, even though the water levelremains constant. This is necessaryin order to make room for the newleaves which are rising in succession from the center of the plant. Oneneed only look at a large plant of any tropical Nymphaea as grown ingardensto be convinced of this. We see an area ten or twelve feet in
Physiology. i i 5diameter covered with the foliage from a single well-grown specimen. Ifthe gardenershave not been too careful we see that the outer border ofthis ring is occupied by yellow and decaying leaves on very long petioles.Within these comes a zone of large green and shining healthy leaves, thensmaller tender ones, and, at the center, one or two are only partly unrolledfrom the bud. The rate of elongation of a petiole of N. rubra detachedfrom th parent plant was 2.5 cm. per day. This, with perhaps a certainamount of stretching of the collenchyma and softer tissues, may accountfor the rise of leaves on plants moved from shallow to deeper water, butit does not reach the case of adjustment from deepFrank (1872)to shallow water.investigated the causes of this phenomenon, giving especialattention to Hydrocharis morsus-ranae, and concluded that gravitation(as expressed by water pressure) and light, though not the causes, areused by the plant as signs by which to measure how much it has yet toaccomplish before a growing organ has reached its most advantageousposition (" Gravitation und Licht sind nicht die Erreger jener Wachsthumsformen,sondern die Pflanze bedient sich ihrer nur als Merkmale, an denensie abmisst, wieviel sie noch zu leisten hat, bis das durch Wachsthum zurichtende Glied seine vortheilhafteste Lage erreicht hat." p. 85). Theexpression is an excellent one, even if the " signs " are insufficient. Foris it not so with all stimuli and allorganisms? No external stimulus is initself a final cause of any physiological action that; is, the irritability ofprotoplasm is the indispensable condition of all vital functions. Karsten(1888), experimenting on Hydrocharis and Ranunculus sceleratus, showedthe insufficiency of Frank's explanation, and proved that it is the oxygen ofthe air which inhibits the growth of the petioles as soon as the leaf-blade"reaches the surface of the water (dass esder Sauerstoff der Atmosphereist, welche bei den Schwimmblattern jene constatirte Hemmung imWachsthum ihrer Stiele bewirkt, sobald sie die Wasseroberfiache erreichen."p. 577). In Nymphaea, however, the inhibiting influence of theair stillpermits the slow elongation of the petioles by which the olderleaves are moved farther and farther away from the center of growth.Once spread out upon the water surface, the leaves are subjected tothe stress of currents in air and water. Jahn (1896) pointedout fourrequirements for such foliage: (1) lightness and firmness of lamina; (2)the greatest possible amount of surface ;(3) insertion of petiole at centerof leaf ;(4) the petiole must make a large angle with the lamina. Thefirst is attained by the great amount of air space, and in Lotos and Apocarpiaeby the girder-like veins. But in strong winds the leaves are often
n6The Waterlilies.raised up at one edge or blown completely wrong side up. The toughnessof the lamina prevents its tearing. Breadth of leaf surface is carried toits ultimate extent in the circular leaves. The larger the leaves the lesseasily they will be overturned, and a further need will be appreciatedwhen it is noted that waterlilies must have all of their foliage in oneplane ; a distribution in spaceof three dimensions is not available to them.That the rounded outline has been acquired by a backward extensionthe basal lobes of the leaf is due to the third condition. A round laminawith marginal attachment of the petiole is quite a possible thing. But ifsuch a leaf were to exert any tension upon its anchorage, the effect wouldbe to submerge the basal portion. The central attachment obviates thisdifficulty and peltationis a further advance toward the same end.;Various other adaptations of the plant in irritable response to its peculiarenvironment have been referred to in previous pages. A number ofinteresting activities of the floral parts will be described shortly.The nutritive relations of waterlilies have not been specially workedout. They require a large amount of nitrogenous food, as every cultivatorcan testify, but preferit in the form of nitrates rather than ammonia.Excess of calcium, as in limestone waters, is not beneficial. Doubtlessthese substances are absorbed with water by the roots directly,ofsince noroot-hairs are present. And the importance of the roots is shown by thegreat reduction in vigor of plants when moved from one pot to another,Wachter (1897) has shown, how-though they soon regain their strength.ever, that complete removal of the roots is less injurious than removal of theleaves. Water is also absorbed elsewhere than at the roots. A leaf ofN. rubra 35 cm. across, severed from the parent plant and left floating inthe water, with the end of the petiole tied to a stick above the water level,not only remained apparently healthy for three days, but grew 1.3 cm. indiameter, and its petiole elongated 6.3 cm. Whether or not this leaf wastranspiring moisture through the stomata I cannot say ; but that a largeamount of transpiration occurs from waterlily leaves was shown by a smalltank inmy father's garden. This was a water-tight iron vessel four feetsquare, and the surface was almost completely covered with leaves sothat very little free evaporation could occur. After every clear summerday it was necessary to add one to three buckets of water to keep thelevel constant. So reduced is the xylem in the petioles that one questionshow and where any sap can ascend. Some have thought the smallair-canals in the vascular bundles are filled with liquid, but we find noevidence of this. But, in a leaf placed with the end of the petiole in
Physiology. 117watery eosin, the phloem portions of the bundles were stained as thoughthe coloring matter had traveled up in them. The whole subject of thecourse of water in aquatic plants needs investigation.Respiration is to be looked upon as causing some of the most characteristicstructures of water plants, namely the air-canals. In August, 1867,Lechartier experimented on N. alba as it grew naturally in the rivers.He found that on cutting off a leaf just below the water surface a flow ofgas takes place from the air-canals of the petiole. This flow continuesthroughout the day, ceases at dark (7 p. m.), and begins again the nextmorning (8.30 a. It m.). may continue even against a pressure of 26 cm.of water, and after all of the leaves have been cut off the plant. In oneinstance 220 cc. was collected from a single petiole in an afternoon, and inanother 262 cc. The total amount of gas from a single plant was 1,028 cc.in fifty-three hours. In these plantsall of the leaves were submergedduring the experiments. A plant with several floating leaves was nextused. A petiole was cut off near the leaf, and when held 1 cm. below thewater surface no gas came out ;when raised in a closed inverted tube ofwater 10 cm. above the surface of the river, the flow amounted to 600 cc.in fifteen minutes. This gas was tested in portions of 60 cc. collectedsuccessively, and at three times of day, as follows :The first, fifth, and tenth portions of air contained
u8The Waterlilies.therefore believed the flow to be due to a higher pressure in the plant thanoutside. He agreed with Lechartier in believing that there is a movementof gasesin the air-canals of Nymphaeaceae toward the leaf, to makeexit through the stomata, and concluded that stomata in generalare forthe exit rather than for the entrance of gases. Goebel's (1893) experimentseems to differ from these. On a dull autumn day petioles ofN. rubra and stellata (?)were cut off under mercury near the water surfaceand held as nearly as possible erect. The smaller intercellular spacesbecame in every case injected with mercury up to the lamina, even inpieces 60 cm. long.This would indicate a considerable negative pressure.Recognizing the positive pressure as noticed by Sachs (though withoutmention of Lechartier or Barthelemy), Goebel concludes that when theamount of assimilation is slight there is a rarefaction of air in the canalscaused by the continuance of respiration but when assimilation is active,;part of the oxygen is given off into the intercellular spaces, and reachesthe air-canals and finally extends to the rhizome and root. This view atonce harmonizes the observed phenomena and explains the presence ofthe air-canals. It is the more plausible when we consider the peaty soilin which our waterlilies grow and the general absence of free oxygen insubmerged muddy material.Since the behavior of the flowers has for its object the successfulreproduction of the species, and since there is in this connection a verydistinct series of movements and activities, the whole relation of theseparts has been reserved to this point, although there would be good reasonto take them up under the head of irritability.The rise of the flower budfrom the rhizome to the surface of the water takes place in the same manneras that of the leaf, and the peduncle must be regulated in length bythe same stimuli that regulate the petiole ; it is even capable, in Castalia,of a slight elongationif the water level rises after the flower first comesinto bloom. A peculiar adaptation occurs, on the other hand, in plants ofany species, when grown in very shallow water, say 10 to 15 cm. Thisoften occurs in our gardens, where we have the tropical kinds as near tothe surface as possible in order to keep them warm. Now, the flower budis firstpushed out from the apex of the plant in a very immature state,and it is to finish itsdevelopment as it is rising through the water. Invery shallow water, therefore, it reaches the surface when it is by no meansready to be exposed to the air. The situation is recognized at once bythe plant, and the peduncle bends to one side and becomes S-shaped
Physiology. 119(Fig. 52) thus the bud maintains its erect ;position and iskept submerged.Then in due time, the bud having matured and the peduncle havingreached a suitable length, the latter straightens up and lifts the former intothe proper position for opening. In N. flavo-virens,where I have noticedthis most plainly, the flower stands about a foot above the water.The opening of the flower occurs at a particular time of day for eachspecies of Nymphaea. Indeed, we have attemptedin the accompanyingFig. 62.A strong-growing waterlily in shallow water.list * to imitate Linnaeus's floral clock in this genus alone. After remainingopen a few minutes (N. amazonum) or hours, the flower closes at acertain time, and this isrepeated on two (N. flavd) or three {N. odorata,caerulea, etc.) to six or seven (N. gigantea) successive days. This is thefoundation on which was based the statement of Pliny and others that theflowers retreat under water at night.The process of opening and closingusually occupies nearly an hour. On its first day the flower always opens* Floral clock for the latitude of Philadelphia {408- 4 a. m. N. amazonum (new flower) opens.t- 8 N. amazonum (old flower) opens wide.6-6 N. amazonum closes; N. marliacea carneaopens.6-7 N. alba candidissima and odorata minoropen.N. gracilis and elegansXzanzibariensisopen.7- 8 N. elegans, capensis, and caeruleaXzanzlbariensisopen.N. caerulea, marliacea chromatella, andtuberosa open.8-9 N. marliacea chromatella (new flower)opens.N. capensisXzanzlbarlenBis opens.8-10 N. zanzibariensisX opens.10-11 N. mezicana opens.N. rubra and rubra rosea close.11-12 m. N. tetragona and zanzibariensis open.N. omarana closes.N.).13- 1 p. m. N. odorata, tuberosa, caerulea, and elegansclose.1-22-3N. alba candidissima closes.N. elegansXzanzibariensis and marliaceachromatella close.8-4, N. mexicana closes.N. capensisx zanzibariensis closes.4-6) N. tetragona, gracilis, and Pennsylvania6-6close.N. capensis, zanzibariensis, and zanzibariensisXclose.6-77- 8 N. dentata and omarana open.N. amazonum (old flower) opens sepals.8- 9 N. rubra, rubra rosea, and devonlensis9-10)10-lH11-12open.N. rudgeana opens and closes (Casp.).
120 The Waterlilies.about an hour later than on subsequent days, closes earlier, and spreadsbut half as wide. An exception to this must be made for N. amazonum,whose very peculiar habits are detailed in the description of the species.N. rudgeana has also shown a very short nocturnal period of opening.Observations on other Hydrocallis species would undoubtedly yieldinteresting results. The characteristics of each species, so far as known,are givenin the taxonomic chapter.The stimulus to these actions is to be attributed, in diurnal species atleast, wholly to light.If plucked and taken into a room they rapidly close,even though the normal time has not come. On the following morning,however, they reopen even in a very dimly lighted room. Subsequentmovements of the cut flower soon become irregular, and it finally ceases tomove. This is"probably due to the death of the protoplasmin the sensitiveor motor centers, though the flower may remain open without wilting forsome days. That light is the normal stimulus is also shown by the earlieropening of the flowers in more northern latitudes. Thus Kerner (1895)a. m.gives 7 a. m. as the time for opening of N. alba at Upsala, and 8 to 9at Innsbruck, 13 farther south. Heat seems to have no effect upon theEu-castalia group in ordinary atmospheric quantities for the time of;opening and closing remains the same on days and nights when the mercuryis continually above 8o F. and on cool, cloudy days when the temperaturedoes not rise above 65 F. In tropical species as grown out ofdoors in our climate heat does have a marked effect. One eveningin1900, when the mercury registered about 95 F., N. zanzibariensis remainedopen nearly an hour later than usual and in; autumn, when the nighttemperature reaches 40 F. and the plants are looking the worse for wear,the diurnal tropical species become very irregular and open but a littleway. The conclusion to be drawn is that the plant itself must be in acertain condition of health or tonicity to enable its parts to perform theirfunctions. The effect of heat is essentially systemic rather than local. InLotos it seems likely that heat may have a more distinct influence uponthe flowers, but it is still obscure. They do not close until the heat of thesun ismaking itself felt, but on exceedingly hot nights they open toperfection, and on much cooler days they close at the usual time. Butwhen the cold of autumn injures the plants, they sometimes remain opentwo or three days and nights continuously. On one occasion at Riverton,N. I J.,saw about sixty large flowers of this type cut in the morning andplaced in jars of water in a shaded glass house ;they closed very slowlyand incompletely. We must conclude, therefore, that lightisagain the
Physiology. 121principal stimulus, causing in these closure, in other species opening, ofthe flower. The difference in effect is paralleled in positive and negativeheliotropism and positive and negative geotropism. The influence ofdifferent colors of light has never been tried. Chemical stimuli were usedby Tassi (1884). He found that flowers of N. alba and other plants wereunable to perform their movements under the influence of anaesthetics,and in -some plants the color changed.The large and showy flowers of waterlilies are inmany ways designedfor the visits of insects. The colors are mostly conspicuous and attractive.N. caerulea iscomparatively insignificant and N. rubra makes verylittle;showing at night, but in the early morning it is magnificent. N. rudgeana,according to Caspary (1878), is often cleistogamous. The possibility ofsuch a thingissuggested by its regular occurrence in Euryale ; andN. amazonum, with its odd habits, may well be considered as approachingthe same condition. Certain it is that close pollination occurs regularly inthis species.In cold weather, too, the flowers of N. amazonum seem to befertilized without ever rising to the water surface. The occurrence ofsimilar irregular cleistogamyin N. rudgeana may have been due tounnatural conditions. Observations are totally lacking on Hydrocallisspecies in their native haunts.In Castalia and Apocarpiae the flowers have a sweet ethereal odor.This is richest and most delicious in N. odorata, faint in N. tetragona,in N. alba.scarcely perceptible and only present on the first day of openingThe Lotos and Hydrocallis groups have verylittle odor ;in the latter it ispleasant and ethereal, but in the former it is dull and rather offensive.The flowers are always pistillate on the first day of opening, and thestigmatic cup is filled nearly to overflowing with a large quantity of slightlysweetened water. This is excreted by the stigma itself and by the axileprocess. The stamens at this time stand erect around the stigma, leavinga narrow vertical pit in the center of the flower, bounded all over the sidesby the still closed anthers of stamens of different lengths. All the stamensare stiff in Lotos, but in Castalia the innermost filaments are very slenderand flexible, scarcely able to hold up the heavy anthers. The carpellarystylesalso stand erect. Inflowers of the second and later days the stigmaisquite dry and the papillae shriveled (in N. odorata and tuberosa ;Robertson, 1889), thoughin N. alba, according to Schulz (1890) stillreceptive.The carpellary styles, when long enough, have bent inward over thestigma until they lie horizontally or point downward, and the innerstamens meet in the middle so as more or less to hide the pistil.The
122 The Waterlilies.stamens now dehisce. Where the flower isopen only for two days, asin Xanthantha, all of the stamens dehisce on the second day, but the outerones precede the inner by a few hours. In the rest of Castalia the innerstamens dehisce on the second day of opening they do not completely;cover the entrance to the stigma from above, but they hang over farenough to drop their pollen upon it, and sometimes close fertilizationoccurs. In Brachyceras, as a rule, the entrance to the pistil is totallyobstructed on the second day by the inner stamens, and the outer ones aredehiscing; subsequently the inner stamens bend outward and dehisceuntil, on the last day, the passage to the stigma is wide open again. Inspite of this, N. caerulea and stellata are said by Caspary (1877) to becapable of self-pollination, the latter while still in the bud. The stiffstamens of the- Lotos group dehisce simultaneously on the second day,and undergo no considerable change of position.The exact manner of effecting cross-pollination and the insectsconcerned are unknown for the Lotos group, but the white and light-pinkvarieties are extremely fertile when grown out of doors in the vicinity ofPhiladelphia. The agency of insects in Hydrocallisis likewise undetermined.N. alba is, in the opinion of Delpino, fertilized by scarabaeid beetles (Cetoniaeand Glaphyridae); he regards Nymphaea and Victoria as especially adaptedfor beetles. Kirchner, in his "Flora von Stuttgart," page 275, mentions"flower-beetles" as visitors {fide Miiller, 1883, p. 93; Schulz, 1890), andPicciola found Donacia (a chrysomelid beetle) abundantly. But Schulzfound N. alba perfectly fertile with its own pollen, and considered thatflies and beetles which visit italwaysthecause close fertilization. N. tuberosais visited by small mining bees (Andrenidae), flower-flies, and beetles(Robertson, 1889), and N. odorata by Halictus and others. In our ownexperience, mining bees and flower-flies are the only insects found. Theyare drowned in large numbers in the stigmatic fluid, and are found in theclosed flowers after anthesis. In a single bloom of N. capensis X zanzibariensisI once counted thirty-two little bees ! Bacon (1874) noticed deadinsects in N. odorata and supposed that they were caught by the closing ofthe flower ;Delpino attributed their death to the heavy odor of the flower,and Planchon to the accumulation of carbon dioxide in the floral cup {fideRobertson). But Robertson isundoubtedly correct in considering itsimply a case of drowning. The course of events is about thus : Theinsects which visit the flowers are in search of the copious pollen;no truenectar is secreted. The earliest flowers to open each day are the olderones, in which pollen is plentiful and the stigma is dry. The insects fly
Physiology. 123from flower to flower in safety gathering their loads. When an hour laterthe new flowers open with their receptive stigmas, the insects are welldusted over. Coming to the narrow opening of the new flower, they rushin with confident haste; but reaching the slender inner stamens, these benddown under the weight, and the visitor isdropped into the pool of liquid inthe stigmatic basin. As he struggles to get out by crawling up the styles,he must again lay hold of the treacherous inner stamens, and be again letdown into the water. Thus pollenis washed off the bee into the fluid, andthe need of the plantis met it is a matter of; strength and endurance onthe part of the visitor whether or not he gets out alive. When thestigmatic fluid is absorbed, the pollen grains settle down among thepapillae and are in a position to germinate. In some species a considerableamount of heat is evolved in the newly opened flower, but neverin anything like the quantity found in Victoria regia.On the last day of opening the flower begins to sink back into thewater again. In Hydrocallisit takes but 2 to 5 hours for the closed flowerto become totally submergedin N. odorata it ;requires 6 to 1 2 hours, andthe flower remains nearly wide open. In Lotos and Brachyceras also theflower often sinks while stillopen, but on its last day it has already moveddown near to the water surface, and is not totally submerged until 1 2 to 20hours later. The submergence of the flower is due to active, even powerful,movements of the peduncle, and occurs most pronouncedlyin fertilizedflowers. Infertile flowers scarcely get below the surface of the water, andthen quickly disintegrate sometimes the ;peduncle decays before the floralparts, showing how greatis the transmitted effect of fertilization. Underthe most favorable circumstances only about half of the ovules proceedwith development to form mature seeds, but the presence of 10 or 12 fertileovules in each cell of such fecund species as N. lotus or caerulea is sufficientto stimulate the movements of the peduncle.In Lotos the pedunclesimply bends over and carries the flower down, and the ripening fruit lieson its side. In N. amazonum there is a double curve, so that the ripeningfruit is upright. But in other species more or less of spiral coiling accompaniesthe bending. The stout peduncles of Brachyceras descend in thesame manner as N. amazonum, but there is in addition a strong curving ofthe lower half of the peduncle to the right or left. The result is that wefind the fruits near the bottom of the water, 15 to 35 cm. away from thecrown of the plant and on the opposite side from the point of insertion ofthe fruit-stalk. N. flava in shallow water (15 cm.) makes the same kindof a bend ;as the stalk is more slender, the bends are sharper and the
124 The Waterlilies.fruit ispressed close against the mud. N. tetragona resembles N. flavain these respects. In deep water (i to 5 meters) N. flava, odorata, alba,and tuberosa carry the spiral turning to its highest development. Froma few centimeters above the rhizome to within a few centimeters ofthe flower, the ispeduncle coiled into a close helix from 2 to 8 centimetersin diameter, and with 2 to 8 or 10 turns ;thus the fruit is drawn downwithin about half a meter of the bottom of the water, and often quiteinto the mass of algae and other vegetation overlying the mud. Justsuch an adaptation has given the specific name to Vallisneria spiralis.Besides removing the fruit from the drying influence of the air, thepurpose of these movements is doubtless twofold : First, to hide thefruit from the many aerial, aquatic, and amphibious animals which mightfeed upon it, for the majority of such creatures get their food at thesurface rather than in the depths of the water ; secondly, to reduce thedanger of the fruits being broken off, as would easily occur with the long,slender peduncles of Castalia.
CHAPTER Y.*TAXONOMY.'Genus NYMPHAEA (Linn.) J. E. Smith.Generic diagnosis Sepals 4 (rarely 3 or 5), inferior. Petals many(12 to 40), multiseriate, the lowest alternate with the sepals. Stamensmany (20 to 700), multiseriate, inserted above the petals on the upraisedtorus which surrounds the carpels;outer filaments broader, often petaloid;innermost slender or filiform ;anthers introrse, bilocular. Carpels many(8 to 35), sunk in the cup-shaped, fleshy receptacle, fused dorsally with it,and produced above it into shorter or longer processes, the carpellary styles(except in Sec. Anecphya); margins of carpels fused centrally with the floralaxis, which extends above them as a short axile process. Stigma broad,concave, radiate. Ovules very many, anatropous, pendulous from the sidesof the ovary cells. 'Fruit a spongy berry, ripening under water and burstingirregularly. Seeds with a floating sac-like aril, open at the apex;endosperm very small ;perisperm copious. Embryo small, straight.Aquatic herbs with perennial elongated or tuberous rhizome, rootingin mud at the bottom of ponds, streams, etc. ;leaves floating, ovate toorbicular, fissi-cordate, 5 to 60 cm. in diameter flowers; solitary, showy,floating or raised above the water on stiff scapes, 2.5 to 30 cm. in diameter,white, blue, red, or yellow, in all shades.Founded by Smith on N. alba L.Nymphaca, Tournefort 1700, in part. Linnaeus 1753, in part.Nymphaea, J. E. Smith 1809. DeCandolle 1821 b, 1824. Planchon 1853 b. Lehmann 1853 a. Caspary1855, 1865, etc. Bentham & Hooker 1862. "Gray, etc. Not Salisbury 1806 a.Castalia, Salisbury 18060. Greene 1887 a and b, 1888. Britten 1888 a. Lawson 1889. Britton &Brown 1897.Leuconymphaca, Ludwig 1737. Kuntze 1891. MacMillan 1892.Nymphea, Rafinesque 1830.1This chapter, together with most of the bibliography at the end of Chapter VIII, was submittedto the University of Pennsylvania in partial fulfillment of the requirements for the degreeof Doctor of Philosophy, May 1, 1901. At that time I had placed the members of the Lotosgroup in a single species, N. lotus. The separation of these into three distinct species, the fusion ofN. Hava and N. mexicana, the addition of some African species, and the substitution of " N. fiavovirens" for " "N. gracilis are the only material changes made since that time. A few citations andmeasurements have been added in several species.125
126 The Waterlilies.In order to present a tabular view of species and subgenera thefollowing synopsis and key are inserted. The genus Nymphaea dividesreadily into two main divisions, which are subdivided into five subgenera.Group I. Nymphaeae apocarpiae. Subgenus (i) Anecphya. (2) Brachyceras.Group II. Nymphaeae syncarpiae. Subgenus (3) Castalia. (4) Lotos. (5) Hydrocallis.Analytic Key.Carpels free at sides, i. e., walls between ovary cells double (Fig. 40) {Nymphaeaeapocarpiae) 1Carpels fused at sides, i. e., walls between ovary cells single (Fig. 40) {Nymphaeaesyncarpiae 131 .Carpellary styles wanting ;stamens unappendaged (Anecphya) N. gigantea1. Carpellary styles present, thick and fleshy ;stamens appendaged (Brachyceras) 22. Flowers yellow 32. Flowers never yellow 43. Leaves 4 to 6 cm. across ;veins inconspicuous N. sulfurea3. Leaves 20 to 25 cm. across ;veins very prominent N. siuhlmannii4. Leaves entire or nearly so 54. Leaves dentate or crenate 105. Leaves narrowly elliptic (twice as long as broad) ,blotched N, ovalifolia5. Leaves ovate or orbicular 66. American species ;flowers pale violet N. elegans6. African species 77. Mature leaves bulbilliferous N. micrantha7. Not bulbilliferous 88. Leaves deep purple beneath A^. calliantha8. Leaves green or purplish-green beneath , spotted with black 99. Petals s to 10 ;stamens 11 to 16 N heudelotii9. Petals 14 to 20 ;stamens 50 to 75 N. caerulea10. Flowers white 1110. Flowers blue or pink 1211. Petals narrow, acute ;leaves pure green beneath N. flavo-virens11. Petals elliptic, obtuse ;leaves black-spotted beneath N. ampla12. Petals 20 to 30 ;stamens 150 to 275 N. capensis12. Petals 1 1 to 14 ;stamens 33 to 55 A'', stellata13. Sepals prominently veined ; styles linear ;leaves spinose-dentate (Lotos) 1413. Sepals obscurely veined ;outer filaments petaloid 1714. Leaves 10 to 15 cm. long, thin and membranous N. zenkeri14. Leaves 20 to 25 cm. in diameter, stiff ;venation very prominent 1515. Flowers red ;leaves bronzy N. rubra15. Flowers white 1616. Leaf orbicular, smooth or finely puberulent beneath N. lotus16. Leaf ovate, densely pubescent beneath N. pubescens17. Styles clavate ;flowers nocturnal (Hydrocallis) 1817. Styles ligulate ; flowers diurnal (Castalia) 2618. Sepals and petals long acuminate A^. oxypetala18. Sepals and petals acute or obtuse 1919. Leaves sinuate N. rudgeana19. Leaves entire 2020. Petiole bearing a ring of long hairs at topA'', amazonum20. No ring of hairs at top of petiole 21
Taxonomy Nymphaeae apocarpiae. 12721. Styles 12 to 16 mm. long A^. gardneriana21. Styles 4 to 10 mm. long 2222. Sepals with black markings 2322. Sepals marked with dark crimson lines 2422. Sepals marked with fuscous striae 2523. Spots few, large ;leaves large, broadly peltate N. lasiophylla23. Spots few, linear ;leaves small, narrowly peltate N. tenerinervia24. Leaves green above and below N. blanda24. Leaves marked beneath with dark purple forked and branching lines. . . . A^. jamesoniana25. Leaves very narrowly peltate, small N. stenaspidota25. Leaves larger and more peltate N. gibertii26. Rhizome erect 2726. Rhizome horizontal ;flowers white or pink 2927. Flowers yellow (Xanthantha) N. tnexicana27. Flowers white, small (Chamaenymphaea) 2828. Leaves plain green above A'', fennica28. Leaves more or less blotched with brown A^. tetragona29. Leaves crowded on the rhizome ; European or Asiatic 3029. Leaves scattered on the rhizome; American 3130. Line of attachment of sepals to receptacle sharply angular A^. Candida30. Line of attachment of sepals rounded A^. alba31 . Petals spatulate ;rhizome tuberiferous N. tuberosa31. Petals elliptic ;rhizome not tuberiferous N. odorataGroup I. NYMPHAEAE APOCARPIAE (= Lytopleura Casp. 1865, 1878, 1888).Carpels free from one another at the sides, fused along part of the suture with theaxis of the flower, and dorsally with the perigynous torus. Tropical species withdiurnal flowers ranging in color from blue through pink to white, raised 7 to 30 cm.above the water, on stiff peduncles. Venation of sepals inconspicuous. Outermoststamens first to dehisce, innermost last. Principal air-canals in the peduncle 6 (5 to 7),in the petiole 2; idioblasts very few; multicellular ingrowths rather frequent in theair-canals. Rhizomes erect, tuberous, drying off in the resting season. Plants glabrousthroughout, except on the apical portion of the tuber and the base of the petiolesand peduncles.Subgenus 1. ANECPHYA Casp. 1865, 1888.Carpellary styles absent. Stamens, very many, with narrow filaments and short,curved anthers, without appendage, or simply mucronate, inserted very densely at thesummit of the torus around the stigmatic disc, the outermost being at some distancefrom the insertion of the innermost petals. Seeds larger than in the next section.Flowers appearing subtended each by two leaves, i. e., there are on the rhizome obliqueseries of flower bases alternating with two series of leaves. One species, in Australiaand New Guinea.Sec. Cyanea, Planchon 18526, 18536 (in part).Sec. Appendiculatae, trib. Cyananthos, Lehmann 1853a (in part).Sec. Inappendiculatae, trib. Castalia, Lehm. 1853a (in part).
128 The Waterlilies.Nymphaea gigantea Hook. (Plate III.)Diagnosis.See under subgenus Anecphya.Nymphaea gigantea, W. J. Hooker 18520, Ad. original specimen in hb. Kew. Planchon 18516,1852 d, 1853 6. Rev. Hortic. 18526. Lehmann 1853 a. Paxton 1853 d. F. Muller 1861,1862. Bentham & Muller 1863. Caspary 1865. Garden 1883 b. G. & F. 1893 a, with goodhalf-tone. Tricker 1897. Moenkemyer 1897. G. C. 1900. Conard 1901a. Not seen:Lemaire 1853; Duchartre 1858; A. G. 1894, 1897.Castalia stellaris, Salisbury 18066 (in note to tab. 14, referring to specimen collected by Banks atEndeavor River).Castalia gigantea, Britten 1888 a. Lawson 1889.Victoria Fitsroyana, Hort. (fide Hooker, 1. c).Description. Flower 6 to 30 cm. in diameter (12.5 to 19 cm. as cultivated aboutPhiladelphia; 30 cm. in the type specimen, dried), opening for 7 days from 9 a. m. to7 p. m. (Casp. 1865), inodorous. Bud ovoid, rounded at apex. Peduncle terete,rather slender (1 to 2.5 cm. in diameter), tapering upward, smooth, light green, rising10 to 35 cm. (18 inches, Robinson 1865) above the water; air-canals as in groupApocarpiae (figured by Hooker 18520, as petiole). Receptacle short, small, darkgreen. Sepals 4, oval or elliptic, rounded at the apex, breadth : length = 1 :2.5 to 3(average) ; persistent in fruit. Anterior (outermost) sepal pure green without, witha few blackish lines and dots; margins blue-purple above, shading to carmine-purpleat point of attachment. Posterior (innermost) sepal with a broad green area in middleof back; a wide border (3 to 5 mm.) around apex and sides, narrowing to thebase of the sepal, covered by the lateral sepals in the bud, of a dark royal-purple colorin apical part, shading to sky blue at sides and carmine-purple at base. Lateral sepalswith one side covered, and similar in coloring to posterior sepal. Five strong veinsand several smaller ones visible at base of each sepal. Inside of sepals sky blue,shading to royal purple at apex, dark blue on lateral margins, and carmine purple below.Petals 18 to 51, longer than the sepals and not entirely covered by these afterthe first day of opening; outermost petals obovate, obtuse, narrowed at base, deeplyconcave, not at all sepaloid, outside sky blue, shading to royal purple at apex and base,similar in color, but paler, within. Intermediate petal obovate to spatulate, roundedat apex, tapering at base, colored like outer petals, but paler throughout; innermostpetal oblanceolate, rounded at apex, tapering to the base, pale whitish blue, tingedpure blue at apex and base. All of the petals thin and fragile, finely 1 to 7 nerved,satiny and crumpled like chiffon, fading in color on later days of opening. Stamens367 to 745, incurved at summit through an arc of 45 to 90 ,half as longas thepetals, separated from these by a wide interval and inserted densely on the upperpart of the ovary. Anthers bright yellow, about 75 outer ones bearing a short, thin,acute apical appendage (or all unappendaged, Casp. 1865). Filaments paler yellow;10 or 12 outermost ones broader than their anthers; all the rest are nearly or quite"thread-like and longer than their anthers. Outermost anthers first to open, innermostlast. Pollen smooth" (Casp. 1865). Carpels 12 to 20, quite distinct from oneanother and rather easily separated from the receptacle, stigmatic over all their upperfree surfaces ; carpellary styles wanting. Ovules large, few in number. Seed " large,
CARNEGIE INSTITUTION OF WASHINGTON WATERLIUES, PLATE 3titNYMPH/EA GIGANTEA.i. Anterior sepal. 2, Outermost stamen. 3, Inner stamens, 1. Median petal. 5. Vortical -eriion of ovary. 6, Outermost petalHFIIOfYPE CO., B03TON.
Taxonomy Nymphaea gigantae. 129dark colored, ellipsoid, apiculate, usually covered with interrupted longitudinal linesof hairs " (Casp. 1865). (See table of measurements of flowers, below.)Submerged leaves from sprouting tuber (Fig. 34) 2 or 3, entire, green above, reddishbeneath. First leaf triangular-hastate, with rounded apex and angles, and nearlystraight sides; sinus, very broad and shallow. Second leaf ovate-sagittate, broadlyrounded at apex ; sinus broadly triangular, lobes narrowly rounded. First floating leafovate, with deep open sinus and rounded apex and lobes; margin slightly irregularwavy.Second and third leaves similar, with narrower sinus and margins distinctlysinuate-dentate on outer side of lobes; first two dull green beneath, third suffusedwith purplish blue, and with a few blackish spots.Leaf of mature plant very narrowly peltate, orbicular-ovate or elliptic, 60 cm. orless in diameter, breadth :length = 1 :1.4 ; margin flat or slightly wavy, sharply dentate;teeth short, acute (or obtuse, Casp. 1865 and Mrs. Rowan's paintings), 2.5 cm.apart and 0.6 cm. long ; upper surface green and veiny, minutely elevate-punctate undersurface glabrous, brownish pink when young, becoming Tyrian purple when full;grown. Veins prominently reticulated on under side of leaf; primary nerves 9 or 10on each side; length of principal area: length of radius of leaf = 1: 1.33. Depth ofsinus =0.37 of length of leaf; margins entire, touching or slightly overlapping within,then curving out, becoming 7.5 to 15 cm. apart at periphery of leaf; angles producedinto a fine subulate tooth. Petiole smooth, terete, 60 to 150 cm. long, with 2 largeair-canals, 2 smaller ones before and behind these, and a surrounding ring of 10 stillsmaller ones ;2 double bundles anteriorly and posteriorly, with a peripheral ring of 1 1bundles. Stipules absent, but petiole winged at base, the wings gradually vanishingabove.Rhizome (tuber) ovoid; small ones (Fig. 14) nearly spherical, 1.6 to 1.75 cm. indiameter, contracted above, the vegetative bud springing abruptly from the roundedsummit; larger tubers with prominent leaf-bases. Each flower subtended by twoleaves, the peduncular scars on the rhizome forming an oblique series alternating withtwo series of leaf-scars (Casp. 1865).Measurements of Flower from University of Pennsylvania Botanic Garden, 1900 (in cm).Sepals.
'30The Waterlilies.Measurements of Leaves (incm.).Length ofMidrib.Lengthfrom Apexto Angle.Depth ofSfntiB.Width ofLeaf.Width ofSinus.First submerged*..Second submerged*First floating*Second floating*...Third floating*MatureMature242.232.552.873-22.873-245-14.838.2460-50.81.61.92.23%2.53-230.640i-91.91. 121.421-3* From a small tuber.Geographic Distribution. Australia and New Guinea. "Lakes and marshesthroughout tropical Australia" (Benth. & Mull. 1863). "At river mouths in S."Australia," Schomburgk 1875. Moreton Bay, Queensland; Clarence River, N. S.W.," Benth & Mull., 1. c.Papua, Mull. " 1875. New Guinea, coll. Zippel Endeavor;River, coll. Banks," Casp. 1865. Rockhampton, E. Australia, coll. A. Dietrich, in hb.Berlin. King's Creek, Queensland, coll. Miiller, in hb. Munich.Type collected by Bidwill, No. 39, at Wide Bay, Queensland, in hbb. Kew, Berlin.Forma alba with white flowers.Forma rosea with pink flowers.Benth. & Mull., 1. c.Benth. & Mull., 1. c.Notes. Introduced into England in 1852, and fruited by Van Houtte (Ghent)in 1855. Flowered in open air, in heated water, at Glasnevin (Dublin) in 1865. Frequentin cultivation in America, flowering freely out of doors all summer. Tubers aresaid to germinate better in deep water, i. e., about 60 cm. ;but this is not necessaryifgoodbottom heat is furnished.Var. violacea (Lehm.) new comb. (Plate I, Frontispiece.)Smaller than the type. Flower 12 to 15 cm. in diameter. Bud conical-ovoid,acute. Sepals coriaceous, ovate to oblong, tapering above with nearly straight sidesto the obtuse apex ; outer surface dark green more or less densely marked with interruptedheavier and lighter black lines; inner surface pale. Petals many, rich violet,slightly shorter than the sepals ; outer ones larger, obovate oblong, tapering above.Stamens very many, much shorter than the corolla; appendage obsolete. Carpelsabout 10.Leaves subovate, repand, light green above, deep purple beneath ; petiole to apex9 to 11 cm.; width 11.5 to 1 2.7 cm.; lobes 6.4 cm. long, obtuse; primary nerves 10to 12 ;veins numerous, reticulate, prominent beneath, green. Sinus margins curved,open at periphery.Rhizome cylindrical, the size of one's little finger. (Details mostlyfrom Lehmann.)Nymphaea violacea, Lehmann 1853 a, Ad. original specimens coll. John Macgillivray, Oct. io, 1848,Cape York, Voyage of the Rattlesnake, Botany, 410, in hb. Kew.N. Banksii, Cunn., fid. original specimen, coll. A. Cunningham, June, 1819, Endeavor River, N. E.Australia, in hb. Kew.
Taxonomy Nymphaea elegans. 131N. serrata, Muller, fid. original specimen, coll. F.. Miiller, July 15, 1855, Arnheim's Land, Australia,in hb. Kew.N. repanda, Muller, fid. original specimen, coll. F. Muller, July 16, 1856, Arnheim's Land; alsoHead of Sturt's Creek, Australia; both in hb. Kew.N. versicolor, Muller, fid. original specimen, coll. F. Muller, July 15, 1856, Arnheim's Land, Australia,in hb. Kew. Not Roxb. 1809.N. stellata, Muller 1861, fid. original specimens, sent by F. Muller from Rockingham Bay, Queensland,in hb. Munich; from Escape Cliffs, N. Australia, coll. W. Halls, in hb. Kew; fromBa r ron River, Queensland, coll. W. Sayer, in hb. Paris ;also in hb. British Museum. NotWilld. 1797.N. Brownii, F. M. Bailey, 1899, fid. original specimen in hb. British Museum.Geographic Distribution. Queensland, Arnheim's Land, Australia.Notes. Caspary considered this merely a small form of N. gigantea; but thepaintings of both types made by Mrs. Rowan in the native habitat, which I saw at thePhiladelphia Museum, April 16,1901, show the marked peculiarities cited above, whichcertainly entitle the plant to varietal rank. Mrs. Rowan considers it a very distincttype, and says it is restricted to the Cape York Peninsula, a region practically uninhabitedby white people. We are especially fortunate in having opportunity to reproduceMrs. Rowan's painting from the original, which is now the property of theUniversity of Pennsylvania. Examination of the European collections gave fullyconclusive evidence on the rank of this form. (Cf. also its synonymy.)Subgenus 2. BRACHYCERAS Casp. 1865, 1878, 1888.Carpellary styles present, short, stiff, and fleshy. Stamens numerous, followingthe petals in order on the torus directly or without much interval ;exterior antherslong, more or less strongly appendaged with a process of the connective, the correspondingfilaments being flat and more or less expanded. Seed smaller. Twelve speciesof world-wide distribution in the tropics.Cyanea and part of Lotos DC. 1821 b.Cyanea (mostly) Planchon 1852 b, 1853 b.Section I. Appendiculatae Lehmann 1853a (in part).Nymphaea elegans Hook. (Plate IV; Fig. 53.)Leaves entire or slightly wavy at base, narrowly peltate, broadly ovate to ovateorbicular;under surface dark purple; about 18 cm. long. Flower 7 to 13 cm. in diameter,pale violet, open from 8 a. m. to 1p. m. Buds ovate. Sepals marked with blacklines and dots. Petals 12 to 20, ovate, obtuse. Stamens about 100, stout, yellow;appendage minute; filaments narrow.Nymphaea elegans, Hooker 1851, fid. original specimen, cultivated at Kew, from Texas, in hb.Kew. Revue Horticole 1851 b. Paxton 1853 b. Sterns 1888. Tricker 1897. Conard 1901 a.Castalia elegans, Greene 1888. Lawson 1889. Rose 1895.Nymphaea Mexicana, Gray 1850. Not Zucc. 1832.Description. Flower 7 to 13 cm. in diameter, open widely on three successivedays from 8 a. m. to 1 p. m. ;odor sweet, moderately strong. Bud ovate, obtuse orrounded at apex. Peduncle terete, slender, stiff, rising 12 to 18 cm. above the water,
132The Waterlilies.reddish brown, with 5 or 6 main air-canals, each with two very small canals outside.Receptacle very narrow, sloping. Sepals 4 (or sometimes 5) lanceolate, breadth :length= 1 :34 to 3.6, obtuse or sub-acute outer surface dark; green, marked with finebrownish-black lines and dots, especially near the margins 9 longitudinal veins visible;by looking through the sepal at a light; inner surface dull greenish-white. Petals 13to 24, lanceolate, tapering to base and apex, breadth : length= 1 :3.7 in outermostpetals, crowded on lower part of torus. Outer surface of outermost petal sepaloidFig. 58. Nymphaea elegant; a, sepal ; b, petal of outermost whorl ; e, petal of second whorl ;d, petal of Innermost whorl ; e, outer stamen ; /, successively smaller stamens ; g, vertical sectionof ovary. Natural size.cFloral Organs.a
CARNEGIE INSTITUTION OF WASHINGTON. WATERLILIES, PLATE 4.ANYMPH/EA ELEGANS.
Taxonomy Nymphaea elegans. 133from the width of the anther at topof filament to about one-half as wide at base.All of the anthers longer than the filaments. (See measurements at end of description.)Ovary nearly hemispherical, with a naked ring on the sides between petals andstamens. Carpels 15 to 25; styles yellow, erect, minute, 0.8 mm. long, a littlebroader than high, acute, the tip projecting as a very small point beyond and abovethe broadly rounded and centrally grooved stigmatic ray. Stigma more or less curvedfunnel-shape, with large central depression, in which the slender, conical, acute axileprocess stands free. Diameter of stigma 0.8 cm. ; depth of depression 0.3 cm. ; heightof axile process 0.2 cm. Fruit rounded, much depressed, cheese-shaped, 3.2 cm. indiameter by 2.2 cm. high, to 3.5 cm. in diameter by 1.9 cm. high, of a pale greenish color,subtended by the persistent sepals, contracted above to the low crown of hard, inwardlydirectedstyles; stigma basin-shaped, about 1.6 cm. in diameter, black, radially striatesulcate;axile process broadly conical and obtuse.inSeeds nearly globose, about 0.13 cm.diameter, mucronate at hilum by persistence of a portion of the funiculus, dull olivegreen, smooth, with rather strong raphe.Aril about as long as the seed, and closelyinvesting it.Germination may occur immediately on ripening of seed, or after a period ofdrought. First leaf of seedling filiform, 1.6 to 2.8 cm. long. Second leaf ovate, narrowedabove, obtuse, 1.12 cm. long by 0.48 cm. wide, rounded at base; petiole 0.8 cm. long.Third leaf like second, but with slightly cordate base. Fourth leaf broader and morecordate at base.Fifth leaf broadly ovate, cordate, with rounded apex and basal lobes.First submerged leaf from small tuber (Fig. 34) narrowly triangular,acute atapex ; base truncate with rounded angles. Second leaf typically cordate, acute. Firstthree floating leaves entire, ovate, broadest about midway of length, with rather widesinus; apex broadly rounded, angles rounded; first two leaves green on both sides,third green above, purplish beneath. Fourth floating leaf speckled above with smallbrownish blotches, under surface brownish red with a number of fine darker specks ;margin wavy on outside of lobes.Measurements of Leaves from Small Tuber(in cm.).Length ofMidrib.LengthfromApex toAngle.Depth ofSinus.Width ofLeaf.Width ofSinus.First submerged . .Second submergedSecond floating. . .Fourth floating. . .1.62-52.61-552.94-55-7O0-451.92-550.51.93-04.151.01.2Leaf of mature plant broadly oval, or at times nearly orbicular ; breadth :length =1: 1.3; usual length 15 to 18 cm., peltate by 0.6 to 2.0 cm., of rather thin texture,entire in upper half, very gently sinuate-dentate in basal half; apex rounded. Uppersurface dark green,with small blackish-purple markings when young; under surfacebright purplish-red with frequent blue-purple spots, greenish along the midline. Veinsprominent out to the sixth grade; principal nerves 6 to 8 on each side; length of principalarea : radius of leaf = 1 :1.76. Sinus open, depth:length of leaf = 1 : 3
134 The Waterlilies.(approx.), margins straight or slightly concave, converging toward the periphery;angles about 3.2 cm. apart, tapering, acute. Petiole slender, light brown, attached tothe lamina by a slender " collar " ; principal air-canals 4, two larger and two smaller ;stellate cells numerous on walls of larger canals.Rhizome (tuber) erect, ovoid, black, thickly covered with leaf scars, about 7.5 cm.long by 3.5 cm. thick, ripening early in the fall. No stolons formed in vegetativeperiod. Small tubers (Fig. 14), 1.2 to 1.35 cm. in diameter, spherical-ovoid, smoothin lower half, tapering above to the scaly-woolly vegetative end.Measurements in centimeters.Stamens OF Flower A.*
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 5)3-NYMPH/EA AMPLA.i. Flower. 2, Leaf. 3,Flout r si-en from side, 4, Rays ofsti^iiKi and carpellary styles. ;, Stamens.6, Rays ofstiynifl from another (lower.MELIOTYPE CO., BOSTON.
Taxonomy Nymphaea ampla. 135N. Lotus, Linn. 1762 (the American plant). Aublet 1775. Willdenow 1797. Loudon 1855.N. Candolleana, Lehmann 1853 a, as to the leaf (fid. original specimen from hb. Lehmann now inhb. Berlin). Flower is of N. rudgeana.N. foliis circinnatis maximis, Burmann 1759.N. ampla var. Plumieri, Planchon 1853 b. Grisebach 1857. Caspary 1878.N. trisepala Gaudich. ? fid. Eichler 1878.Description. Flowers white, 9 to 13 cm. across. Sepals 4, coriaceous, narrowlyoblong, breadth: length =1:2.3 to 5-8 (average of 51 sepals from 43 flowers =1 :3.6, C-.sp. 1878), obtuse, acute or acuminate, outer surface green, marked with shortblackish lines. Petals 7 to 21, the outermost tinged yellowish-green. Stamens 30 to190, outermost much longer than innermost ;anthers appendiculate. Carpels 14 to 23 ;styles short-conical, gradually narrowed or abruptly apiculate-acuminate, acute, subacute,or obtuse, breadth :length =1:1.5 on the average, =1:1 to 2.5 in extremes;axile process rounded, breadth :length = 1 :0.09 on the average,= 1 :0.3 to 2.0 in extremes;stigma extending out on styles in short, rounded or rarely acute rays. Seedsub-globose-elliptic, about 1 mm. long, with longitudinal rows of hairs. Leaves large,narrowly peltate (3 to 31 mm.; average 10 mm.), sub-orbicular, 15 to 40 cm. indiameter, sinuate or nearly entire, green above, with small black spots ;under surfacered-purple; veins prominent, principal nerves 5 to 14 (average 10) on each side ofarea : radius of leaf =1:2.leaf ; length of principalGeographic Distribution. Tropical and sub-tropicalAmerica, from 26 N. in Texas, through the Antilles,to 8 S. in Brazil; Fort Clark and Spofford, Texas(Plank 1896). Lampasas, Mexico. Neuvo Leon; Lakesbetween Ocuiltzapotlan and Famulte de las Sabanas; Lagunasde Macultepec (very large specimen, coll. J. N.Rovirosa, 1889, in hb. Acad. Nat. Sci. Phila., with petals9.5 cm. long and leaf 56 cm. long by 48 cm. wide).Mexico, coll. Houston, No. 173 1, in hb. British Museum.Springs and streams near Monterey, Mexico (Pringle, Plant. Mex., 1889, No. 2581).La Paila, New Granada. St. Domingo island. Vera Cruz, Hazienda de Mamulique,Hazienda de Tamatoc, Laguna Verde, Mexico (Casp. 1878). Aguada del Labach, Yucatan,coll. Schottmuller, No. 528, in hb. British Museum ;also Gaumer, No. 428, in hb.Boissier. Cuba; St. Miragoane; Alligator Pond, Jamaica; Rio Ulna, Honduras;Belize, Honduras, coll. J. Robertson, Dec. 27, 1889, No. 20, in hb. British Museum.Hog Island, Bahamas, coll. Eggers, No. 4081. Martinique, coll. Plee, in hb. Paris.Guatemala (Smith 1 891, 1893) fid. original specimen in hb. Kew. Encruzilhada, nearPernambuco, Brazil, coll. H. Schenck, Herb. Brazil, No. 4152, in hb. Berlin, and coll.Gardner, No. 915, near Alinda, in hb. Kew. Demarara, British Guiana, coll. Parker,in hb. Kew. Trinidad, coll. Fendler, No. 207, in hb. Kew. St. Vincent, south end, coll.Smith, No. 1708, in hb. Kew. Prov. Quito, Peru, coll.Hartweg, No. 1592, in hb.Boissier.Notes. In the arrangement of this species we have followed Caspary as closelyas possible; he examined the type material in nearly all of the European collections.DeCandolle's type specimen consists of a leaf of this species with a flower of N.rudgeana Mey.Sepal Dimensions.
136 The Waterlilies.The true species (var. Plumieri Planch., etc.) is of powerful growth, but hasnot yet been introduced into cultivation. We have discarded the varietal name in thisand in other species, allowing the specific name alone to stand for the form originallydescribed.Var. speciosa(Mart. & Zucc.) Casp.Of medium size; stamens 42 to129; leaves less coriaceous, margins more or lessrepand-sinuate or dentate, with obtuse teeth ;nerves slightly prominent beneath ;lobesdivergent, overlapping or approximate. Tuber sub-globose.N. speciosa Martius & Zuccarini 1832, fid. original specimen, coll. Martius, in aquis S. Christoph.,Prov. Rio de Janeiro, Brazil, in hb. Munich, in sched. No. 28.JV. ampla, Hooker 1849.N. ampla P Hookeri, Planchon 1853 b.N. ampla Y Salzmanni, Planchon 18536 (flower).N. tropaeolifolia, Lehmann 1853 a (flower), fid. original specimen in hb. Berlin. Leaf is of N.rudgeana.N. nervosa, Lehmann 18530, from hb. Steudel, fid. Casp. and original specimen in hb. Berlin.N. ampla var. speciosa, Caspary 1878.N. Leiboldiana, Lehmann 1853 a, as to the American plant in hb. Vienna, fid. Caspary 1878. Specimens" ex herb. Lehm." in hbb. Kew and Berlin are from Africa, and appear to be N.copensis.N. gracilis, Zuccarini 1832 a, b, fid. original specimens (2 sheets) coll. Karwinski, Aug. 1827, "exlacu Mexicano," in hb. Munich. Coll. Pringle, Plant. Mex. 1891, State of Jalisco, No. 3891.N. undulata, Lehmann 18530, fid. original specimens, coll. Galeotti, No. 4846 (4840 in Kew), Nov.,1840, " lacs pres Oaxaca, Mexico," ex hb. Lehmann in hbb. Berlin, Kew. Hemsley 1888.Geographic Distribution. Tropical America from 19 N. to 23 S. Valleyof Mexico (Karwinski) ; Oaxaca, Mexico (Galeotti) ; Guadalajara (Pringle). Caguas,Porto Rico, coll. P. Sintenis, No. 2577, Sept. 5, 1885. St. Domingo, coll. Poiteau,1802, in hb. Delessert. Jamaica; Nevis; St. Croix; Lebanon in Antigua; Maracaibo;La Paila in New Granada; Merida; Demerara in British Guiana; Kaw River inFrench Guiana; Pernambuco, Santa Cruz, St. Christophe, Para, Rio de Janeiro, CaboFrio, Capocabana, and Bahia in Brazil (Caspary 1878).Notes. This is the form cultivated in Europe as N. ampla,Jamaica by Macfadyen in 1847 or 1848, and to Caspary from Caracas byin 1869.Specimens from St.sent to Kew fromDr. ErnstChristophe and Maracaibo have the lobes of the leaf widelydivergent; those cultivated in Europe from Jamaica stock have the lobes touching;nearly all the others in collections have an open but very narrow sinus.The type specimens of N. ampla y Salzmanni Planch, and N. tropaeolifolia Lehm.consist each of a flower of this variety with a leaf of N. rudgeana Mey. (Casp. 1878).Var. pulchella (DC.) Casp.Flowers of medium to small size stamens ; 30 to 50 ; marginor slightly repand-sinuate ;veins slender, scarcely prominent.of leaves sub-entireN. pulchella, DC. 1821 b, fid. original specimens, without data, in hb. DeCandolle, but marked coll.Pavon, at Guayaquil (DC. ms.) in hbb. Paris and British Museum. Planchon 1853 b.Lehmann 18530.N. lineata, St. Hilaire 1833, fid. Caspary 1878, and original specimen in hb. Paris.N. ampla var. pulchella, Caspary 1878.
Taxonomy Nymphaea flavo-virens. 137Geographic Distribution. South America, from 2 to 23 S. Guaniva, PortoRico, coll. Sintenis, No. 3662, det. Caspary, in hbb. Berlin, Kew, Munich. Guayaquil,Peru; Province of Rio de Janeiro (between Sitio de Paulista and Sitio do Porier),Brazil (Caspary 1878).Nymphaea flavo-virens Lehmann. (Plate VI; Fig. 54.)Leaves sub-orbicular, 30 to 45 cm. in diameter, narrowly peltate, deeply andirregularly sinuate or nearly entire, plain green on both sides; angle of lobes rounded.Flowers white, 10 to 15 cm. in diameter; sepals pure green, or with 2 or 3 scatteredshort black lines near the margins ; petals 16 to 20, narrow, acuminate; stamens about60, deep yellow ; outermost filaments short, broad, petaloid ; anthers with a yellowishwhiteappendage.Nymphaea flavo-virens, Lehmann 1852 b, 1854, fid. specimens from the Hamburg Botanic Garden,1864, in hb. Kew, and from hb. Lehmann in hb. Berlin. Hooker 1901, fid. original specimenin hb. Kew, cult, in Kew Garden.N. gracilis Hort. Watson 1887? G. & F. 18946. Tricker 1897. Conard 1901 a. Not Zuccarini 1832 a.N. mexicana Hort, not Zuccarini 1832, fid. Hooker,1. c.Description. Flowers 10 to 15 cm. in diameter, open from early morning to 5or 6 p. m., sweet scented, resembling Convallaria majalis. Peduncle terete, slender,glabrous, tapering upward, rising 20 to 30 cm. above water, with 6 small main aircanalsand 12 secondary ones, all with thick partitions between. Sepals 4, lanceolate,tapering to an acute apex, 8.3 cm. long by 2 to 2.4 cm. wide, green and obscurelynerved outside, sometimes with a few scattered short black lines ;inner surface white.Petals 18 or 19, of firm texture, outermost opening nearly horizontally, about as longas the sepals, narrowly lanceolate, acute, breadth :length = 1 : 5.6 to 6.1 (=1.27 cm. :7.8 cm. ;and 1.4 cm. : 8.3 cm.). Outermost petal greenish-white outside, with 2 or 3 reddishlineolse. Inner petals smaller, pure white. Stamens about 60 (56 to 61), yellow;outer ones longer, with short, broad, petaloid filaments, and appendages of mediumlength, stout, light yellow in color. Carpels 12 to 15 ; styles short, stout, fleshy, orangeyellow,becoming elongated and incurved in fruit. Fruit nearly globose, 2.2 cm. indiameter by 2.87 cm. high, crowned by the small circle of styles (1.9 cm. in diameterat base, outside measurement), and slightly depressed, basin-like stigmatic disc; axileprocess broadly conical with rounded apex, about twice as wide as high. Cells ofberry oval, about 1.9 cm. high by 0.95 cm. wide radially. Seed nearly spherical, withinsignificantraphe, about 0.2 cm. in diameter.Floral Organs.
'38 The Waterlilies.Submerged leaves from small tuber 2 or 3; first one ovate, acute, or oftenertriangular-hastate with acute apex and angles ;second leaf much larger, angles slightlyrounded, lobes long and narrow, widely diverging, thin and crisped or even twisted.First floating leaf ovate, abruptly acuminate, broadest opposite the insertion of thepetiole, slightly crenulate; sinus deep, margins straight or slightly curved, slightlydivergent or touching or overlapping, angles narrowly rounded; upper surface greenwith a few brown blotches, especially near the margin ; under surface green, blotchedwith brown.Leaf of mature plant narrowly peltate (1.2 to 1.9 cm.), orbicular-ovate, 30 to45 cm. in diameter, deeply and irregularly sinuate-dentate, with obtuse or rounded teetha- iVia. 54. Symphaea flavo-viretu : a, sepal ; b, petal of outermost whorl ; c, petal of secondwhorl ; d, petal of third whorl ; e, outer stamen ; /, inner stamens ; g, vertical section of ovary.Natural size.1.2 cm. long and 2 cm. apart, rarely nearly entire; pea-green above, paler below; veinsprominent beneath, primary nerves 7 to 9 on each side; length of principal area:radius of leaf =s i: 1.6 to 1.75. Sinus, depth: length of leaf = 1 :2.5 to 2.8; marginsmore or less convex, divergent or overlapping; angles rounded. Petiole terete, 1.2 cm.in diameter, 60 to 150 cm. long.Rhizome (tuber) erect, stout, 3.8 cm. long by 2.9 cm. thick to 21 cm. long by10 cm. thick (!), brownish-black, very rough with projecting leaf cushions, each cappedby the dried base of the petiole and marked on the under side with root scars; upper
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 6NYMPH/EA FLAVOVIRENS.HEUOTYPE CO., BOSTON.
Taxonomy Nymphaea flavo-virens. 139quarter of tuber clothed with long light brown hairs, above which the stout soft leafrudimentsmay project visibly even in the dried state. Phyllotaxy of a high order, butthe leaf scars are 1.2 to 1.9 cm. apart, and by their dry projecting habit give thetuber somewhat the appearance of a rough pine-cone.Measurements of Leaves from Small Tuber, inCentimeters.Leaf.
140 The Waterlilies.Nymphaea stellata Willdenow.Leaves elliptic-orbicular, rather broadly peltate ; margin irregularly sinuate; lobesobtuse; upper surface green; pink or blue-violet beneath. Flower 5 to 12 cm. across,pale blue (varying to pink or white), open from 8 a. m. to 2 p. m. Buds ovate;sepals with minute blackish dots. Petals II to 14, dull white at base ;stamens 33 to 54 ;appendage blue, anthers and filaments pale yellowish.Nymphaea stellata, Willdenow 1797. Andrews tab. 330. Aiton 181 1. DeCandolle 1821b, 1824.Wight & Arnott 1834. Planchon 1853 6. Rev. Hortic, 1854 a, 1855 b, col. plate. Caspary1865 in part ; 1877. Conard 1901 a.Castalia stellaris, Salisbury 1806 a.N. malabarica, Potret, in Lamarck 1802, fid. original specimen coll. Sonnerat, Isle de France, fromhb. Lamarck, in hb.Paris.N. stellata var. parvifiora, Hooker & Thomson, 1855.Description. Flowers small, 5 to 13 cm. across, open on three successive daysfrom 8 a. m. to 2. p. m., nearly odorless. Bud oblong, ovate, rounded at apex.Peduncle terete, slender, rising well above the water-surface. Sepals 4, oblong, subovate-triangular,breadth : length = 1 : 2.8 to 3.5, margins nearly straight toward theapex; outer surface light green, uniform or marked with minute dark purplish lines;inner surface bluish-white. Petals 10 to 16, lanceolate, acute, shorter than the sepals,pale blue above, yellowish-white at base. Stamens 33 to 54, with small blue appendage;anthers and filaments pale yellow. Carpels 10 to 17; styles short-triangular,usually apiculate, obtuse; breadth : length = 1 : 1 to 1.5; stigmatic ray terminating onthe style in a short obtuse triangle, breadth :length = 1 :0.5 to 0.7. Stigma slightly concave.Seed elliptic globose, diameter : length = 0.8 : 0.9 mm. ;=0.8 : 1.0 mm., gray.Leaf of adult plant rather broadly peltate ;breadth of pelta:depth of sinus = 1 :6 to 10; outline sub-orbicular-elliptic, 12 to 15 cm. in diameter, irregularly sinuaterepand,or subentire, rounded or slightly retuse at apex; upper surface bright green,sometimes faintly brownish blotched; under surface purplish to deep blue-violet, withgreen veins. Primary veins 9 to 11 on each side of leaf; length of principal area:radius of leaf =1:1.4 to 1.6. Sinus deep, usually open; margins nearly straight;angles slightly produced, acute or obtuse. Rhizome ovoid.Geographic Distribution. Southern and southeastern Asia and neighboring islands,Java, Borneo, and the Philippines. Malabar, Tanjore in India (Hainan, coll.C. M. & A. Henry, No. 8382).Var. cyanea (Roxburgh) Hooker & Thomson.Flowers of medium size, blue, with faint odor or none.Nymphaea cyanea, Roxburgh 1824. Wight & Arnott 1834.N. stellata /?, Sims 1819. Planchon 18536.Var. versicolor (Roxburgh) Hooker & Thomson. (Plate VII.)Flowers of medium size, pink ;stamens numerous. Leaves sinuate, pink beneath,marked above and below with pustules 1 to 2 mm. in diameter.Nymphaea versicolor, Roxburgh, in Sims 1809. Roxburgh 1814, 1824. Planchon 18536.N. punctata, Edgeworth 1845 = N. Edgeworthii Lehmann 18526, fid. original specimen, coll. M. P.Edgeworth, 1844, Sadhoura, India, in hb. Kew. Not N. punctata Kar. & Kir. 1841 ?
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 7iT oNYMPH/tA STELLATA VERSICOLOR. .>,- )' f;Sepal. whorls, , Stamensof dosed flower. >,- Closed Hover,HEUOTYPE CO., BOSTON.
Taxonomy Nymphaea caerulea. 141N. rhodantha, Lehmann 18530, fid. specimen coll. dimming, No. 702, "in insulis Philippin.," fromhb. Lehm. marked in the handwriting of Lehmann, in hb. Berlin.N. Hookeriana, Lehmann 1853 a, fid. original specimen marked in handwriting of Lehmann, coll.Hooker & Thomson, Dec. 19, 1850, at Noapolly, Bengal, from hb. Lehm. in hbb. Berlin, Kew.N. bella, Lehmann 1853 a, as to the flower ;leaf is of N. pubescens, fid. original specimen from hb.Lehm. in hb. Berlin.Notes. Hooker & Thomson record, and the herbaria bear them out, that nosharp line could be drawn between the varieties of this species as it occurs in India,but they were certainly wrong in including the African plants. The three extremeIndian forms are themselves fairly recognizable. The margin of the leaf may be almostentire. The pustules so evident on the leaf of our N. versicolor cultivated fromseeds sent by Mr. Gollan of Saharanpur, India, may be present at times in the othervarieties. Our plant also had a great number of bulbils on the caudex, as shown inseveral herbarium specimens both of blue and pink forms.Willdenow's description was based solely on the figure and text of Van Rheede'sFlora Malabarica 1 1 :53, tab. 27, and was very brief " N. foliis : integerrimis, lobisdivaricatis, acutis, calyce tetraphyllo petalis acutis longiore." Andrews figured the plantin color and later Sims gave a rather larger form than that of Andrews. Planchon figuresa plant evidently identical with that of Andrews and Van Rheede, and Caspary (1877)places its description clearly in parallel columns with the allied forms N. caerulea,capensis, and zanzibariensis. Of these it comes much nearest to N. caerulea, but isplainly distinct. I have been unable to find it in American gardens.The flowers of this species, according to Caspary (1877), are regularly self-pollinated,often while still in the bud ; it refused to cross with N. caerulea under Caspary'smanagement.Rose-colored and white varieties of N. stellata sent from Canara to Caspary by themissionary Zeigler in 1869 were long cultivated in the botanic garden at Konigsberg.Nymphaea caerulea Savigny. (Plate VIII; Fig. 55.)Leaves entire or slightly wavy at base, orbicular or ovate-orbicular, narrowlypeltate, becoming 30 to 40 cm. in diameter; green above; under surface green withsmall dark purple spots, purplish all round near margin. Flowers 7 to 15 cm. across,open from 7.30 a. m. to 12 m. Buds conical. Sepals thickly marked with black linesand dots. Petals 14 to 20, lanceolate, light blue above, lower half dull white. Stamens50 to 73; appendage long (0.5 cm. on outermost stamens), slender, pale blue.Nymphaea caerulea, Savigny 1802, fid. original specimens coll. Delile, 1798, near Cairo. Ventenat1803. Delile 1813. DeCandolle 1821 b. Planchon 1851 a, 1853 b. Caspary 1877. Wildeman& Dur. 1900. Conard 1901 a.N. stellata, Caspary 1865 in part, 1873. Boissier 1867, 1888. Oliver 1868; and of many authors andcollectors. Not Willdenow 1797.N. maculate, Schum. & Thonn. 1829, fid. original specimens from Copenhagen, kindly loaned bythe curator of the herbarium. Welwitsch ms. in hb. Paris./V. poecila, Lehmann 1853 a, fid. original specimens from hb. Lehmann seen in hb. Berlin.N. scutifolia, Tricker 1897. Not Castalia scutifolia, Salisbury 1806 a.N. discolor, Lehmann 1853 a = JV. caerulea, Kotschy MS. (see note).Castalia stellaris, Salisbury 18066 (in part).C. caerulea, Tratinnick 1822.
142 The Waterlilies.Description. Flowers 7 to 17 cm. across, light blue, with a faint and characteristicsweet odor, open on 3 days from 7.30 a. m. to 12 m. (4 days, 8 a. m. to 2 p. m.Casp. 1877). Bud distinctly conical, with straight sides, obtuse, abruptly contractedbelow to the receptacle, 7 cm. long by 2.5 cm. in diameter at base. Receptacle nearlythree times as wide as peduncle, spreading out at about 45 from the vertical, palepurplish green. Peduncle terete, 0.6 to 0.8 cm. in diameter, 18 to 38 cm. long, mostslender about 1.25 cm. below flower, smooth, dull brownish-green; main air-canals 6,surrounded by a circle of 12 smaller ones. Sepals 4, broadly lanceolate, breadth :length= 1 : 2.8 to 4.7, the sides nearly straight in upper two-thirds, somewhat convergingbelow to the broad base; apex rounded. Outer surface dark green, thickly fleckedwith purplish-black dots and lines most dense near margins of sepal and midway oflength ; the green ground-color assumes a yellowish tint near base of sepal. SevenueFig. SS.Nymphaea eaerulta: a, dosed flower; b, sepal; e, d, e, outer, median, and inner petals;/, h, stamens ; (/, carpellary style viewed en face ; (, vertical section of ovary ; k, vertical section of fruit.Reduced about one-half.longitudinal veins can be seen by looking through the sepal at the light. Inner surfacedull white, tinged blue on midline near apex, greenish and veiny and semi-translucentin lowest three-quarters of length. Petals 12 to 20, of moderately firm texture, openingabout 30 above horizontal, about as long as the sepals. They stand in 3 series:first, a whorl of 4, alternate with the sepals ; second, a whorl of 8, one on each side ofeach outer petal ; third, a whorl of 8 alternate with the second, but usually imperfect,consisting of only 2 to 5 petals on one side of the flower, rendering it so far unsymmetrical.Outermost petal narrowly elliptic-lanceolate, obtuse, with rather broad insertion(0.8 cm. in Flower No. 3, see table of measurements), not at all concave; outersurface colored on lower half and on the midline dark green with black spots exactlylike the sepals ;one-third of width at margin on each side, above, colored pale blue ;inner surface white on lower half, becoming translucent and 7-veined at base, shadingabove to a pale sky-blue at apex and on upper margins. Petals of second whorlshorter than those of the first, rhombic-lanceolate, narrowed at apex and base, 5-veined
CARNEGIE INSTITUTION OF WASHINGTON. WATERLILIES, PLATE 8.If^'fi~"jt *# .> .NYMPH/EA CCERULEA.
Taxonomy Nymphaea caerulea. 143below, white on the lower third, pale blue above, pure sky-blue at apex, the blue andwhite shading together insensibly, the latter predominating in two-thirds of length oninner surface of petal. Petal of third (innermost) whorl linear-lanceolate, obtuse, blueat apex; outer surface with white margins in lower three-fourths of length,the bluecontinuing half way down the midline ;on inner surface, white prevails on lower fiveseventhsof length. Stamens 50 to "J2sometimes appearing inserted without order,sometimes indistinctly spiral, often in about 16 vertical ranks of 3 to 5 stamens each.Outerm.~,t stamens with short and rather broad filaments and long acuminate antherwith two long, nearly parallel anther cells and proportionately long, acute, 3-angled appendage,one angle pointing inward ;blue at tip and half-way down, yellowish-whitebelow. Median stamen has short, broadly elliptic filament, very long parallel anthercells, and short appendage; anther and filament bright yellow, only the appendageblue. Innermost stamen bright yellowall over, short, stout ;anther broader and longerthan filament; no appendage. Ovary nearly hemispherical (e. g., 1.9 cm. in diameterby 0.95 cm. high ; 0.4 cm. by 0.24 cm.). Carpels 14 to 21 ; styles short (0.3 cm. long by0.08 cm. in diameter), fleshy, nearly erect, subacute, concave inward, all alike, pureyellow, papillose up to about 0.08 cm. from apex, the papillose area ending roundly.Stigma nearly flat, bending upward on the styles, yellow. Axile process rising abruptlyfrom the stigma, broader than high (height: breadth = 1 :1.5;= 0.24 cm. :0.38 cm.) and somewhat conical in shape, with obtuse tip, of a whitish color. Fruitlarge, round (4.5 to 6.4 cm. in diameter by 2.5 to 3.8 cm. high), truncate above, withdeep radiating fissures between the carpels, flattened or even excavated beneath aroundthe peduncle, of a pale green color, becoming translucent and brownish, crowned withthe hard, slightly enlarged styles, and surrounded by the sepals and outer 4 petals, all ofwhich are dark green and spotted as in the flower. The peduncle makes a large, rudespiral turn, holding the fruit still erect but with its base nearly or quite resting on theearth. Seed ellipsoidal, acuminate at the hilum, 0.17 cm. long by 0.12 cm. in diameter,dull olive brown ;surface marked with about 14 interrupted longitudinal lines of minutehairs ; raphe evident, not prominent. Aril longer than the seed.Germination may take placeimmediately on ripening, or the seeds will withstanddrying over winter. First leaf of seedling filiform, 1.6 to 1.9 cm. long. Second leafbroadly elliptic on a petiole about 2.5 cm. long; blade about 1.27 cm. long by 0.6 cm.wide. Third leaf broader and shorter. Fourth leaf deltoid-ovate with rounded anglesand apex, and truncate base; lamina 1.6 cm. long by 1.27 cm. wide at base; later leavesfinally deltoid-cordate, with rounded apex and angles, and rather deep sinus. Firstfloating leaves nearly orbicular, 2.5 to 3.8 cm. in diameter, entire ;sinus deep withstraight and touching margins ; under surface marked with irregular blackish spots.First leaves from tuber submerged, 2 or 3 on each shoot; the first one may havea minutely denticulate margin, 1 cm. long by 0.3 cm. wide ; or, more commonly, the laminais entire, broadly deltoid, 3.2 cm. long by 3.2 cm. wide at base, with roundedangles, and petiole 2.5 to 3.8 cm. long. A third one may be produced, of oval shape,fissi cordate, with deep open sinus and rounded apex, 3.8 cm. long by 2.9 cm. wide.First floating leaves from tuber like those of seedling, but larger ; perfectly entire.Leaf of mature plant very narrowly peltate (1:14 to 16, Casp. 1877), ovateorbicularto orbicular, 30 to 40 cm. in diameter (33 cm. long by 30.6 cm. wide), soft,
144The Waterlilies.thin and quickly withering, slightly wavy or sinuate in the basal half, apex slightlyemarginate, dark green and obscurely veiny above, color darker over insertion of petiole.Under surface paler green, with numerous small dark purple blotches, which arelarger near the midrib than at the periphery margins purplish, shading from dark red-;purple at the extreme edge to green, the transition taking place inthe course of abouti cm. near sinus angles, in about 2.5 cm. at apex of leaf. Veins prominent out to thefifth grade ; primary nerves 6 to 10 on each side of leaf. Petiole attached by quite astrong " collar." Length of principal area: radius of leaf = 1:1.3 to 1.7. Sinus usuallyclosed; depth: length of leaf = 1 : 2.7 (= 13.5 cm. :3s. 7 cm.); margins doublycurved, convex and overlapping above, separating about 2.5 cm. from periphery of leafand becoming parallel ; angles subacute, slightly produced, about 0.6 cm. apart. Petioleterete, or flattened near the leaf, about 0.6 cm. in diameter, dull brownish-green ;with2 larger upper air-canals, and 2 smaller ones below, and a ring of 12 still smaller onesoutside. Stipules smooth, fused at base, short, distant above, acuminate, very acute;1.9 to 2.2 cm. long, 1.3 cm. wide at base; free portion 1.1 to 1.6 cm. long, 0.5 cm.wide ; tips 0.9 to 1.2 cm. apart.Rhizome (tuber) erect, thick, ovoid, 2.5 to 7.5 cm. long by 2 to 6.4 cm. in diameter,blackish, apex protected by leaf-bases and long fine hairs. Phyllotaxy of a veryhigh order, the leaf-bases completely covering the larger rhizomes, leaving only about0.3 cm. around each one. Roots springing from the bases of the petioles. Offsetsformed only when the tuber breaks out from the dried state; 2 or 3 shoots may comefrom one tuber; the stolon is terminal, very short, the point of origin of new leavesremaining within the coating of long hairs.
Taxonomy Nymphaea caerulea. 145
146The Waterlilies.the close resemblance of N. caerulea and N. stellata, and partly in the fact that theseand N. capensis were introduced into cultivation near the same time, and the descriptivecharacter of the name caerulea made itequally applicable to all. The geographical proximityof this and N. capensis may have added to the confusion. Indeed, examinationof herbarium material shows that N. caerulea is connected by intermediate forms withN. micrantha on the west and N. capensis on the south. We have not been able tosecure living material of N. micrantha, but the capensis type is very easily distinguishedfrom the present species. Their hybrid (N. pulcherrima Tricker) is entirelysterile. Other related forms are discussed in their proper places. Many botanists haveincluded all the blue waterlilies of Africa and Asia under the name of N. stellata, asdid Hooker & Thorns. (1855) ; Caspary, ten years later, separated off N. capensis, andin 1877 gave exact descriptions of four species in parallel columns; he stated that hisold arrangement (1865) needed complete revision, but unfortunately this work was leftundone.The Kotschy collection from Nubia has long been a stumbling block to studentsof Nymphaea. They are marked No. 166 N. caerulea Sav. and No. 167 N. ampla"DC. Kotschyi Iter Nubicum. In stagnibus pluvialibus ad radices eurontas montisCordofani, Arasch Cool, Jan. 5, 1841, d. Oct. 12, 1839." The first is represented in hbb.Kew, Paris, Boiss., Deless., and Berlin. It was marked by Caspary "N. capensisThunb." The Boissier specimen has narrower petals and fewer stamens than thetype of capensis; the Berlin specimensdiffer between themselves one flower hasspotted sepals, and the leaf gently sinuate; another has leaves typical of N. caeruleabut no spots on the sepal. No. 167 was marked "N. caerulea Sav." by Caspary in hb.Boiss., with the addition of " var. albida " in hb. Paris. The leaf is gently sinuate inthe Kew, Deiessert, and one Paris specimen. It is also represented in hbb. Berlin andBoissier. The leaf may or may not have black spots and some purple coloration beneath.The flower resembles that of N. caerulea, but was evidently white, and in the Kewspecimen, at least, the outer stamen is provided with an unusually long, lanceolate appendage.The two plants seem strikingly intermediate between N. capensis and N.caerulea. No. 167is the type of Lehmann's N. nubica.Self-fertilization occurs regularly in this species.Reference to the importance of N. caerulea as the Blue Lotus, the lotus par excellenceof Egypt, has been made in Chapter I.Nymphaea micrantha Guillemin & Perrottet.(Plate IX.)Leaves small, round-cordate, peltate, entire, petiolate ; petioles very long, slender ;lobes divaricate, acuminate ;under surface reddish, marked with violet-black dots ; uppersurface glabrous, pale green, in mature plants bulbiferous at the summit of the petiole ;stigmatic rays 15.N. micrantha, Guillemin & Perrottet 1830. Hooker 1850. Paxton 18530.N. caerulea, Guillemin & Perrottet 1830. Not Savigny 1802.N. caerulea var. albida, Guillemin & Perrottet 1830 = N. micrantha, Hook.N. rufescens, Guillemin & Perrottet 1830.N. stellata var. bulbillifera, Planchon 1852 c. Not N. stellata, Willdenow 1797.N. vivipara, Lehmann 1852 a, b, 1853 a, 1854.N. guineensis, Schumacher & Thonning 1829, fid. Chifflot 1902.
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 94VNYMPH/EA MICRANTHA.i , Under aide of leaf; 2.Upper side with young plant developing, t, Caudex, with a branch.|,Scar* nf leave* and of a flower, fromaarface of a :caudex, enlarged 5, Cross section of caudex ; 6, Longitudinal section of a caudex. 7. Superficial ftagential section throughm* of fig. 4; /,. base t ;ipetiole;'. base f .1peduncle; B, Section a tittle deeper than 7. o, tame ;t^ S, a little deeper.HELIOTYPE CO., BOSTON
Taxonomy Nymphaea heudelotii. 147Description. Sepals 5, narrow, lanceolate, acuminate, with violet dots. Petals10, 1.3 to 5 cm. long, a little shorter than the sepals, lanceolate, narrow, blue to white.Stamens numerous; filaments whitish, narrow at base; anthers of various lengths,twice as long as the filaments; appendage short, conic-terete. Ovary round. Seedsvery numerous, very small, blackish. Leaves 7.6 cm. long, 5.1 cm. wide (in the type).Rhizomes clustered, with soft tawny wool around the bases of the petioles ;roots verylong.Geographic Distribution. West Coast of Africa. In swamps of the peninsulaof Cape Verde, near n'Batal ;in the Galam region ;in the kingdom of Cayor, etc.Flowers almost all the year.Types. Coll. Perrottet, 1830, in hbb. Delessert, Kew, DeCandolle. Also coll. E.Holub, Eastern Bamangwato territory, Bechuanaland.Notes. The most striking difference between this and N. caerulea Sav. lies inthe viviparous habit. Chifnot (1902) states, and Planchon's description indicates, thatthis character does not appear until the second year of the plant's growth. The firstyear it has nearly white flowers, and corresponds to N. guineensis Sch. & Th. ;thesecond year bulbils appear, but the flowers are small and pale, and the plant correspondsto N. micrantha Guill. & Perr. Only in the third year, according to ChifBot,does the plant acquire its definite morphological characters.We do not find either in the specimens or descriptions of Guillemin & Perrottetany important differences between the several viviparous forms they enumerate. N.rufescens is a larger plant; sepal 7.6 cm. long, petals 18 to 20, stamens very numerouswith petaloid filament; carpels 18 to 20; leaves 12 to 25 cm. in diameter, more or lessspotted beneath, primary veins 12 to 20, prominent; coll. Perrottet, 1830, Senegal, seenin hbb. Delessert, Boissier, DeCandolle ;also coll. Roger, Dec, 1823, Senegambia, inhb. Kew (with the note " Les graines servent de nourriture aux negres ") ;also coll.Buchholz, Eliva Sonanga, Cape Lopez, West Africa, in hb. Berlin. N. caerulea Guill.& Perr. has " leaves strongly peltate, impunctate, red beneath, lobes long, free, acute " ;types coll. Perrottet, 183 1, Senegal, in hbb. Delessert, Kew; also coll. Leprieur, Sept.,1821, " in paludosis prope Ngalete, regio Cago, Senegambia," in hb. Delessert; alsocoll. C. Barter, Baikie's Niger Expedition, No. 7064, 1858, at Nupe, in hb. Paris. N.caerulea var. albida Guill. & Perr. has larger flowers than the type, with narrow lanceolatepetals, nearly white ; types coll. Perrottet, Walo, Senegambia, in hb. Delessert also;coll. Leprieur in same district in hb. Delessert. This is apparently identical with N.micrantha Hook. Caspary's " N. micrantha Hook." (notes and herbarium, now in hb.Berlin) has an orbicular leaf with wide sinus and obtuse lobes, with or without blackspots beneath; petals ovate, white, 3.8 to 4.5 cm. long. This plant was much used byCaspary in hybridization, giving well-marked intermediates with the closelycaerulea Sav. (seen in hbb. Kew, British Museum, Berlin).allied iV.Nymphaea heudelotii Planchon. (Fig. 56.)Flowers small, 2.8 to 5.1 cm. in diameter. Sepals lance-ovate, with black spotsoutside. Petals 5 to 10, lanceolate, acute, bluish-white. Stamens 11 to 16; anthersmuch longer than the filaments. Carpels 4 to 10. Leaves almost or quite entire, ovate
i4The Waterlilies.or orbicular, cleft nearly to the petiole, 2.5 to 8.2 cm. long, reddish beneath, withblack specks. Seed smooth.N. Heudelotii, Planchon 1853 b, according to the original specimen.N. pseudo-pygmaca, Lehmann 1853 a, according to the original specimen in hb. Kew, which consistsof a submerged leaf, with flower and fruit, much dilapidated, coll. Heudelot andcommunicated by hb. Paris.Description. Planchon gives the :following Dwarf, ;glabrous floating leavescordate-suborbiculate, very narrowly peltate, with the sinus narrow and nearly closedVia. S6.Nymphaea heudelotii (a), (b), leaves ; (c), flower ; 1, 2, 8, 4, sepals. From a drawing of thetypes in the notes of B. Caspary, now In hb. Berlin, by kind permission of the Directors. Natural size.(in the membranous submerged leaves wide open) ; margin slightly repand under surfacepurplish, violet spotted. Flowers small ; sepals ovate at base, becoming rather long;attenuate, with violet lines outside ; petals 5 to 8, bluish, appendiculate, anthers five tosix times longer than the rather broader filaments; rays of the stigma 8 to 10, short,triangular, acute. Fruit globose, shorter than the calyx; seed elliptic-globose, smooth(not costate).Geographic Distribution. Senegambia, in shallow flowing streams ;FoutaDhiallon.Types.Coll. Heudelot, No. 844, in 1837, seen in hbb. Delessert, Berlin, and Kew.In hb. Kew I saw another specimen consisting of a half-closed flower and two frag-
Taxonomy Nymphaea heudelotii. 149mentary leaves, coll. Sierra Leone Boundary Commission, Dec. 22, 1891, in "smallstreams, Luseniga, Samu." In hb. Berlin is a specimen, coll. Alex. V. Mechow'sExped., No. 480, Jan., 1880, at Malange, West Africa, marked N. stellata; also coll.Pogge, No. 559, Jan. " 2, auf der Ueberschwemmung ausgesetzten Stellen am Lulua,""and No. 558, Nov. 31, 1881, Wassertumpel in der Campine am Lulua"; also. coll.Schweinfurth, No. 2329, Oct. 30, 1869," Land der Djue: Gr. Scriba Ghattes in derSumpfsteppe im Wasser; Bl.reinweiss."Var. na^a new var. (Fig. 57.)Sepals 4, lanceolate, obtuse, 1.2 to 2 cm. long by about 0.32 cm. wide, green outside,with small and distinct or large and diffuse black spots; nearly white within.Petals 7 to 10, lanceolate, acute, shorter than the sepals, 1.5 to 1.7 cm. long, 4 mm.wide, white. Stamens 11 to 14, yellow, with very short appendage, all nearly alikeand about half as long as the petals, apparently in two series. Stigma of 5 to 6broad carpels, with short, broad styles. Fruit nearly globose, surrounded by all of thefloral parts. Seed gray-green, ellipsoid, 1.3 mm. long, 0.88 mm. in diameter.Fig. 57. Nymphaea heudelotii nana: a, open flower, 1, 2, 8, 4, sepals ; b, under side of leaf;c, flower pressed on one side showing outside of sepals;
150The Waterlilies.Notes.On comparing Mr. Baum's specimens with the originals of N. guineensisSchumacher & Thonning, kindly sent to me at Berlin by the directors of the Museumof Copenhagen, Professor Gilg fully agreed that his former identification of the twowas mistaken. Whether the Baum specimens should have specific rank must remainat present a matter of opinion they are the most divergent forms yet known of a;series of African waterlilies leading by slight gradations up to N. caerulea Sav. Noone could doubt that the two extremesare specifically distinct. The Mechowspecimen, No. 480 cited above, differsfrom Baum's No. 655 in having largerflowers (5 cm. in diameter), sepals 1.7to 2.8 cm. long, stamens two-thirds aslong as the petals, leaves 3.8 to 4.5 cm.across, with big blotches of brownabove ; length of principal area length:of radius = 2:3. Pogge's No. 559 hasthe flower like Baum's 655, the leaf2 cm. across, blotched brown above,dull dark green beneath. Schweinfurth'sNo. 2329has blotched leaves5 cm. across, flower 5 cm. in diameter,stamens 25, a little more than half aslong as the petals. Pogge's 558 hasleaves 5.7 to 7.6 cm. across, and sepal3.2 cm. long (poor specimen). Nextto this come Planchon's types of N.heudelotii. On the other hand, JV.micrantha Guill. & Perr. seems to differfrom JV.in being viviparous.heudelotii Planch, chieflyFio. 58. Nymphaea ovalifolia : leaf and flower. Underside of leaf on right, showing veins. Upper side of leaf onleft, with blotches shaded In on lower half ; folded over atmargin. One-half natural size. From a tracing of theoriginal specimen of lltihin. in hb. Berlin.Nymphaea ovalifolia new sp. (Fig. 58.)Leaf narrowly elliptic, 25 cm.long by 14.7 cm. wide, with largeirregular brown blotches above sinus;10 cm. deep, sides nearly parallel, lobesacute, pelta 0.3 cm. wide. Primaryveins about 6; principal area 2.5 cm.wide by 5.7 cm. long. Under surfaceplain green.Flowers deep blue ("himmelblau "), closed in dull weather. Sepals and petalsacuminate, without dark spots; sepal 4 cm. long by 1.3 cm. wide.Distribution and Types. Coll. C. Bohm, No. 92, March, 1882, Deutsche Expeditionnach Ost-Afrika, in the Wala River, Ugally, in hb. Berlin. With this may beclassed a specimen, coll. F. Stuhlmann, No. 481, July 27, 1890, Expedition Emin Pasha,at Palla Bay, marked " N. stellata var. alba," in hb. Berlin.
Taxonomy Nymphaea calliantha. 151Note. Although a single imperfect specimen affords but a feeble foundation fora species, the peculiarities of this one require us to give it separate consideration. Aname therefore seems necessary.Via. 59. Nymphaea calliantha : Flower and under side of leaf, reduced three-sevenths. Outer stamen, enlargedone-seventh. Drawn by J . Pohl, from Baum's No. 395 in hb. Berlin.Flowers 10 toNymphaea calliantha Conard. (Fig. 59.)15 cm. in diameter, pink, violet purple, or light blue; sepals greenoutside, with black spots near the margin, lanceolate. Leaf entire, ovate-orbicular, cleftnearly to the petiole, purplish to deep royal purple beneath.N. caerulea, Gilg 1903. Not Savigny 1802.N. calliantha, Conard 1903.Description. Sepals 4, lanceolate, 7 cm. long by 1.7 cm. wide, acute, outer surfacedull green with a few black spots near the margin, covered margins tintedinner surface bright rose pink on upper 1.5 to 2 cm., shading to white below. Petalsred;
!c 2 The Waterlilies.17, lanceolate; outer one 1.7 cm. wide, sepaloid on back, itsmargins petaloid; whitishbelow, shading pink in upper two-thirds, to decided rose pink in uppermost 1 cm.;inner surface white in lower three-fourths, becoming rose pink at apex, and on marginsfor 1 to 2 cm. below apex ;inner petals white. Stamens about 100, yellow ;outermoststamen has filament 4 mm. wide, 8 mm. long, anther 2.5 mm. wide at base, 19mm. long, appendage 6 mm. long,1 mm. wide at base, subacute. Stigma? Fruit?Leaf ovate-orbicular, entire, 13 to 28 cm. long by 9 to 23 cm. wide, pelta 0.5 cm., greenabove. Under surface purplish on margin, shading to green in the space of 3 to 4 cm.,spotless. Primary veins 8 on each side, length of principal = area radius : j.y cm. :10.2 cm.Geographic Distribution. Central and Southwest Africa.Types. Coll. H. Baum, Reise nach Siid-West Afrika, Kunene-Zambesi Expedition,No. 395, Nov. 9, 1899, " am Kubango unterh. der Kueio, 1 120 m. ii. M." Nativename " Amavu." In hbb. Delessert, Berlin, British Museum. Gilg (1903) adds, evidentlyquoting Baum, " Bliithen weissblau. * * * Blatter sehr diinn." The pink colorseen in the herbaria may therefore be due to changes in drying, as some other specimensin this collection had undergone the same change. Professor Gilg also considersHopfner's No. 36 from Mossamedes (in hb. Berlin) to be identical with Baum's395. A specimen in hb. Kew, coll. F. D. & E. J. Lugard, Sept., 1896, Botletle River,Ngamiland, Tropical Africa, appears to be the same, as also one, coll. W. P. Johnson,Lake Nyassa, in hb. Kew. Baum's No. 208, Oct. 4, 1899," an ruhigen Stellen im Okachitandabei Kassinga, 1290 m. u M., am Uferrand und in Einbuchtungen des Flusses,auch in der Mundung und in Pfannen hinter Kassinga " (Gilg 1903), has smaller leafand flower, but more numerous floral organs. Leaf 16 cm. long by 13.5 cm. wide ;undersurface deep red-purple. Petals 20 to 26, stamens 115 or more. In hbb. Berlin, Delessert,British Museum. No. 396 of Baum, from the same locality as 395, is smaller thanthe latter, but doubtless of the same species. Flowers rose color ; petals 1 1 to 14, 4 to4.5 cm. long ; stamens more than 50. Leaves very dark purple beneath. In hbb. Delessert,Berlin. Professor Gilg (1903) considers Fischer's No. 4 (in hb. Berlin) from theAfrican lake region between Magu and Kagehi to be identical with the foregoing. Weplace here also a specimen in hb. Kew, coll. A. Carson, 1891, at Karonga, Nyassa, havingflowers 10 cm. across, deep blue ; petals elliptic-lanceolate, stamens stout ;leaf oval,14 cm. long by 10 cm. wide, lobes acute, overlapping, margins wavy in lower half, undersurface dark red-purple.Var. tenuis Conard.Leaves nearly orbicular, 7.5 to 10 cm. long by 7 to 8.6 cm. wide, green above, dulldark purple beneath ; primary veins 6 ; length of principal area : radius = 3.2 cm. :3.8cm.; sinus lobes slightly diverging or overlapping, obtuse, margins nearly straight.Flower 9 to 13 cm. across; petals and sepals very narrow (e. g., sepal 5.75 cm. byI cm.; petals 5.4 cm. by 0.8 cm., 5.1 cm. by 0.85 cm.). Sepals 4, green outside, withnumerous black spots; margin red-purple; inside white.Petals 14 to 17, 3.5 to 5.4 cm.long, white or pale blue, rounded at apex. Stamens 33 or more, slender, yellow, theoutermost with filament 3 mm. wide and I cm. long, anther 7 mm. long, appendage 7mm. long ; innermost stamen with filament 3 mm. long and 0.8 mm. wide, anther 3 mm.long and 1.3 mm. wide, appendage 1.25 mm. long. Carpels 10.
Taxonomy Nymphaea capensis. 153N. caerulea, Gilg 1903. Not Savigny 1802.N. calliantha var. tenuis, Conard 1903.Distribution and Types. Coll. H. Baum, No. 771, March 10, 1900, "an ruhigenStellen in der Kuito-Niederung zwischen Kutue und Sobi, 1200 m. u. M." in hbb.Delessert, Berlin. Professor Gilg unites with this form Pogge's No. 560 from the" " " "Baschilangebiet and Buchner's No. 88 from the Malangebiet I ; prefer to classNo. 88 with N. heudelotii, but I have no notes on Pogge's No. 560. Both are in hb.BerlirNotes. This species is distinct enough, we believe, from the nearest relative,N. caerulea. The color of the under side of the leaf, and the peculiar long narrow areabetween the two primary veins nearest the sinus, are easily noted characteristics. Itshould prove a valuable acquisitionfor the water-garden.Nymphaea capensis Thunberg. (Plate X; Fig. 60.)Leaves large, nearly orbicular, deeply sinuate; angles produced, acuminate ; greenon both sides. Flowers bright blue, large, open from 7 a. m. to 4 p. m. Budsovate. Sepals pure green, spotless. Petals 20 to 30, elliptic. Stamens about 150,appendaged.Nymphaea capensis, Thunberg 1800, fid. original specimens, coll. Burmann, No. 491, from hb. Thunb.in hbb. Delessert, DeCandolle. Harvey 1838. Lehmann 1853 a. Caspary 1865, 1877. Conard1901 a.N. caerulea, Andrews, tab. 197. Kerner 1795. Dryander 1801, fid. original specimens in hb. BritishMuseum. Aiton 181 1. Spach 1846. Tricker 1897, and others. Not Savigny 1802.Castalia scutifolia, Salisbury 18060 and b.N. scutifolia, DeCandolle 1821 b, 1824, fid. original specimen from hb. Thunb. in hb. DeCandolle.Planchon 1850 c, 1853 b. Paxton 1853 c.N. cyanea Hort. fid. Paxton, 1. c.N. Berneriana, Planchon 1852 b; 1853 a, b; fid. original specimens, coll. Bernier, No. 158, 1835," Nord de Madagascar," in hbb. Delessert, Paris.N. Emirnensis, Planchon 1852 b; 1853 a, b; fid. original specimens coll. Bojer, " in ubisque paludosisin provinc. Emirna, insula Madagascar," in hbb. Paris, DeCandolle.N. Leiboldiana, Lehmann 1853 a, as to the African plant, fid. specimens from hb. Lehmann in hbb.Kew, Berlin.N. Petersiana, Klotsch 1862, fid. original specimen, coll. Peters, Feb. 8, 1848, Mozambique, in hb.Berlin.N. stellata var. macrantha, Gilg MS., fid. specimens coll. F. Wilms, Flora Africae Australis, No. 10,Dec, 1883, Distr. Middelberg, Transvaal, in hb. Berlin, and No. 13, July, 1884, Distr. Lydenburg," zwischen Spitzkop und dem Komatiflusse," in hbb. Kew, British Museum, Berlin.Not. N. stellata, Willdenow 1797.N. stellata of Cape collectors and (in part) of many botanists. Not Willdenow 1797.Description. Flower 15 to 20 cm. across, opening on 4 successive days from7 a. m. to 4 p. m. (5 days, 7 a. m. to 1 p. m. ; 5 a. m. to 3 p. m. Caspary 1877), sweetscented. Bud narrowly ovate, pure green. Peduncle terete, 60 cm. long, rising 22 cm.above water; green; 0.8 to 1.3 cm. in diameter, main air canals 6, surrounded by 12smaller ones, and these by 24 very small ones. Receptacle yellowish. Sepals 4, narrowlywedge-ovate, breadth : length= 1 :2.4 to 4.3, tapering above to a rounded apex,opening to about 30 above horizontal ;thick and fleshy in texture ;outer surface deep
154The Waterlilies.green toward apex, paler and yellowish in lower third, and on margins ; that part ofmargin which is covered in the bud is white with a bluish tinge ; inner surface greenishwhite,tinged with blue above, pale green in lower quarter ; obscurely many nerved (18to 19) at base. Persistent in fruit. Petals 21 to 38 (15 to 27, Casp. 1877)of ratherthin texture, shorter than the sepals, all plane (i. e., not concave). Outermost petalelliptic-lanceolate, tapering at base, obtuse at apex outer surface rich sky-blue above,;lower third greenish in midline, with nearly white margins; 5-veined; inner surfacerich sky-blue above, shading to nearly white in lower third, and then to yellowish at thebase.Inner petals successively shorter and narrower, colored like the outermost except-Fio. 60. Nymphaea capensti: o, sepal; 6, outermost petal, hooded at apex; c, median petal;d, innermost petal ; e, /, outer and inner stamens ; g, vertical section of ovary. Natural size.ing the green color on the back. Inner 3 series of petals acute ;innermost sometimesnarrowly long-acuminate, simulating a stamen. Stamens 120 to 225 (82 to 243, Casp.1877), the outermost about half as long as the petals. Anthers all longer than the filaments; appendages obtusely triangular, long on outer stamens, becoming obsolete onlyon the innermost. Outer filaments broadly elliptic, much wider (3 times) than anther;inner-median filaments about as wide as the anthers; innermost filaments narrowerthan the anthers. Appendages always blue, like the petals ; filaments yellow throughout;backs of the outer anthers (connectives) shading from blue above throughdeep blue-purple to yellow at base. Inner stamens yellow throughout. Just outsideof the ring of carpellary styles there often appear short filiform yellow processes,which evidently represent stamens reduced by crowding and lack of nourishment.Ovary about 2 cm. in diameter. Carpels 24 to 31 (12 to 47, Casp. 1877) ;styles stout, oblong to linear-lanceolate (breadth: length = 1 : 2 to 5.5), fleshy, pure
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 10NYMPH/EA CAPENSIS.HELIOTYPE CO., BOSTON.
Taxonomy Nymphaea capensis. 155yellow, rounded at apex, 0.64 cm. long on back, 0.16 cm. wide at base. Stigmaticrays extending up two-thirds of the length of the styles, and ending at an acuteangle but with rounded apex. Axile process conical, white, translucent. Fruitdepressed spheroidal, 5.7 cm. in diameter by 3.6 cm. high, crowned with the comparativelylong (0.64 cm.) obtuse incurved styles; stigma concave. Seed ellipsoidal tonearly globose, 0.16 cm. by o.l cm. (average) (0.15 by o.n; 0.16 by 0.09; 0.17 by0.11), acuminate at hilum, dull dark olive-brown, with about 18 interrupted longitudinallines of rather long hairs, giving the seeds a rough appearance to the naked eye, andmaking" i:hem roll about irregularly.Germination immediate or after drying off. First leaf of seedling filiform, 1.75to 2 cm. long. Second leaf elliptic, I.I cm. long by 0.46 cm. wide, rounded at bothends. Third leaf broadly elliptic, 1.1 cm. long by 0.64 cm. wide, rounded at apex,abruptly rounded at base. Late submerged leaf nearly orbicular, entire, deeply cordate,broadly rounded at apex, lobes rounded, diverging 30 to 40 ; length of leaf 2.4cm., width 2.2 cm., depth of sinus 1 cm., petiole stout, about 1.3 cm. long by 0.08 cm.thick.Germinating tuber produces 3 or 4 submerged leaves. First leaf broadly triangularwith rounded angles and truncate base ; second leaf ovate-cordate, apex abruptlyshort-acuminate, sides parallel in lower half, sinus about 8o open, with straight marginsand rounded angles ; third leaf cordate-ovate, with rounded apex and lobes, sinusdeep and wide, open about 45 ; fourth leaf similar to third, but with acute angles andmargin a little wavy on outside of lobes. First floating leaf coriaceous, orbicularovate,narrowly peltate (0.25 cm.), margin slightly wavy in lower half, excavated onouter side of angles of lobes; apex rounded; sinus deep, narrow, margins convex,touching above, angles obtuse or subacute.Leaf of mature plant narrowly peltate (1.3 to 1.6 cm.), orbicular or orbicularovate,25 to 40 cm. in diameter, thin and soft in texture, doubly or trebly sinuate dentate,somewhat crisped, teeth mostly rounded, sometimes angular and obtuse, depressionsbetween teeth 0.3 to 1.3 cm. deep; apex of leaf rounded. Upper surface brightgreen, depressed-veiny, somewhat brown-spotted when young; astomatic area large,paler in color, umbonate over the insertion of the petiole. Stomata very many andsmall on most of the lamina, larger and scattered at margins of astomatic area. Undersurface pea-green, veins all prominent, 10 to 12 on each side; length of principal area:radius of leaf =1:1.1 to 2.0. Sinus, depth: length of leaf = 1 : 1 to 1.16, marginsoverlapping above, diverging toward periphery, angles produced, acuminate, awntipped.Petiole terete, dark green, about 1 cm. in diameter, 60 to 150 cm. long, with4 main air-canals, 2 larger and 2 smaller, and 10 small ones surrounding these ;transversesection near the leaf shows several large thickened fibers in the parenchymaand around the periphery ;stellate cells very few, with short, thick rays. Both of thesetypes of cell diminish in prominence further down the petiole. Base of petiole finelypuberulent. Stipules partly adnate.Rhizome (tuber) ovoid; small ones distinctly egg-shaped, smooth in lower twothirdsof length, tapering above to the bud, 2.5 cm. long by 1.5 to 1.6 in diameter;older ones clothed with long, dark silky hairs and thickly beset with leaf-bases; producingno offsets during active growth. Roots spring from the bases of the petioles.
156The Waterlilies.Geographic Distribution. South and East Africa; Madagascar; ComorroIslands. Zeekoevley, Promontory of the Cape of Good Hope, and Zwartkops river atUitenhage, Thunberg 1800. Port Natal, Pretoria, L'oldstrom ; Hopefield, Delagoa Bay,Albany, Semliki Valley; Zanzibar, Island of Nossi-be; Island of Johanna, Comorro.Tananarivo ;Province of Emirna, Madagascar.Flower.
Taxonomy Nymphaea capensis. 157capensis; after examining the types, beside much other material from South Africaand Madagascar in the European herbaria, we can see no basis for a separation ofthese forms.N. capensis grows easily and flowers and fruits freely. Caspary (1877) considereditincapable of self-pollination, but it produced much seed in the Botanic Gardenof the University of Pennsylvania (1900), though it had usually but one flower openat a time.Introduced at Kew by F. Masson about 1792, carried from the Cape in H. M. S." Gorgon," Captain Parker.Var. madagascariensis (DC.) new comb." Leaves narrowly peltate, 5.1 to 6.3 cm. in diameter, obtusely sinuate, impunctate,glabrous, lobes divaricate. Flowers blue. Sepals 4, green outside, oblong, subobtuse,2.1 cm. long. Petals 8, oblong, subacute, 4 inner ones smaller. Stamens 8 to 12,shorter than the petals. Carpels 8 to 12. Rhizome small, sub-spherical." (DeCandolle,1821 b, 1824). A small form of N. capensis.Nymphaea madagascariensis, DeCandolle 1821 b, 1824. Planchon 1853 b. Not Caspary 1880.Geographic Distribution.Madagascar.Types. Coll. Brugiere, Madagascar, 1801, in hb. DeCandolle. Other specimenscoll. Bernier, No. 159,"etangs a Ling-raton," in hb. Paris; coll. Perville, No.677, Feb., 1841, Ambongo, Madagascar, in hb. Paris; coll. Perrottet, 1820, Madagascar.in hb. Paris ;coll. Chapelin, Madagascar, in hb. Paris ;coll. Boivin, 1847-52, Madagascar,in hb. Paris.Notes. Caspary (1880) regarded DeCandolle's type as a dwarf, of which thelarger specimens of Perrottet, Goudot, and Rutenberg were more perfect forms.recognize, with Caspary, that our knowledge of Mascarene Nymphaeas is extremelydeficient, but it has seemed best to include all the larger ones in N. capensis, and torecognize these smaller ones simply as dwarfs of that species. This view is supportedby the known capacity of the species, and especially of the var. zanzibariensis, todevelop small flowers when in limited quarters. A study of living plants can alonesettlethe question.WeVar. zanzibariensis (Casp.) new comb. 1 (Plate XI; Fig. 61.)Leaves large, somewhat peltate, nearly orbicular, closely and irregularly sinuatedentate;sinus closed, angles not produced; under surface more or less suffused violet.Flower large, open from 1 1 a. m. to 5 p. m. Sepals green, spotless, margins purple,inner surface deep purplish-blue. Petals 18 to 24, oblong, obtuse, deep blue (lightblue or rosy in varieties). Stamens very many (136 to 242), appendages and back ofouter anthers deep blue, shading to carmine-purple (in the type).Nymphaea zanzibariensis, Caspary 1877, fid. original specimens from hb. Casp. in hbb. Berlin, Kew,Munich ;1882. Andre 1882. Moenkemyer 1897. Tricker 1897. Conard 1901 a.N. stellata var. zanzibariensis, Hooker 1885. Rev. Hortic. 1897. Not N. stellata, Willdenow 1797.1Referred to in the text, for the sake of brevity, as " N. zanzibariensis."
158The Waterlilies.Description. Flowers 15 to 25 cm. across (25 to 250 mm., Casp. 1877), openingon 3 to 5 successive days from 1 1 a. m. to 5 p. m. ; highly fragrant number of;floral organs 175 to 300, average about 220. Bud rather broad, oblong-ovate, obtuse orrounded at apex. Peduncle stout, rising 10 to 20 cm. above the water, with 6 mainair-canals. Receptacle rather long, swollen, with sloping sides. Sepals 4, oblongovate,breadth :length= 1 :2.3 to 3.0, obtuse, opening about 15 above horizontal ;marginsgently curved toward the apex outer surface dark green, often shaded with reddish-brownin upper three-quarters uncovered margins deep carmine-brown, covered;;margins shading from carmine-brown to deep violet inner surface;deep purplish-blueabove, shading to yellowish-green at base. Innermost sepal narrowest, with broad(12 mm.) covered margins. Outer surface of sepals with 7 principal veins (5 in smallFio. 61. Nymphaea capensis zaiizibariensU : Sepals, petals, and stamens of successiveseries, slightly reduced.flowers) alternate with these are 8 more slender nerves. Inner;surface shows theveins as darker lines, there being 32 or more at base of each sepal. Petals 16 to 24,long obovate, the outer obtuse, innermost acute, all rich deep blue above, with obscuredarker veins, shading through carmine-purple to whitish at the point of attachment tothe torus. The outer 4 stand in a whorl alternate with the sepals ; all, or only the outer2, carmine-brown on the back just above the base. Second whorl consists of 8 smallerpetals, alternate by twos with the sepals. A third whorl of 8 still smaller petals follows,alternating with the second, but often only the anterior and lateral ones are developed.A few petals of a fourth whorl, alternate with the third, may be present, evenwhen the third is not fully developed. All of the petals, except those of the fourthwhorl, have 5 principal nerves, visible on the outer side. Stamens 126 to 242 accordingto size of flower, slightly shorter than the petals; appendage and inner side ofouter anthers deep blue, back of anthers deep carmine-purple, giving way to light yellowon the long-elliptic filament. Inner filaments inverted wedge-shape. In very large
CARNEGIE INSTITUTION OF WASHINGTON.WATERLILIES, PLATE 11.I**VNYMPH/EA CAPENSIS ZANZIBARIENSIS.
Taxonomy Nymphaea capensis. 159flowers the carmine-purple on the back of the outermost anthers extends down to themiddle of the filament ;the inner side of the filament is also suffused with blue above.Pollen yellow to the naked eye, colorless under the microscope with transmitted light ;grains flattened-spherical, smooth, with large nucleus. Appendage of outermost stamenslanceolate, about 7 times as long as broad, with a swelling on inner side whichgrows to a keel on median stamens, and on innermost stamens the appendage is thickerthan wide. Anthers bent inward, making an angle with the filament. Carpels 15 to31 styk^ triangular, breadth length =1 ; : :2.5 to 2.8, tapering above, acute, straight anderect on first day of opening, afterward bending gradually inward until in fruit theyare hook-shaped. Stigma yellow, its rays short, triangular. Fruit yellowish, depressedspherical,56 mm. high by 67 mm. in diameter, covered by the enlarged, thick andleathery sepals, containing several thousand seeds. Seed dull brown, elliptic, ratherlong, with papilke at one end; diameter :length = 0.75 mm. : 1.2 mm.; =0.8 mm.:1.45 mm., with 15 to 24 interrupted rows of short hairs, the rows at one end oftenalternate with those at the other.Germination immediate, or after drying. First leaf of seedling filiform. Secondleaf broadly ovate, sub-acute, rather abrupt at base, 1.3 cm. long by 0.8 cm. wide.Third leaf orbicular-ovate, truncate near its junction with the petiole, I.I cm. longby 1.0 cm. wide. Fourth leaf round-reniform, with shallow, rounded sinus androunded lobes, 1.6 cm. long by 1.4 cm. wide.Late submerged leaf orbicular, fissi-cor-Measurements.date, 3.8 cm. long by 3.5 cm. wide, entire,rounded at apex; sinus 1.9 cm. deep, marginsconvex, divergent, angles rounded.First floating leaves orbicular, 2 to 2.5 cm.in diameter, entire; sinus deep, closed, itsmargins straight and touching, angles meetingand completing the circular outline ofthe leaf; upper surface more or less thicklyblotched with irregular reddish-brownspots under surface pale green, with a distinctviolet tinge all around near the mar-;gins. The later leaves have less and less ofbrown spotting above, and gradually acquirethe characteristic margin and color beneathas the plant approaches maturity.Sepals.Submerged leaves from sprouting tuber 2 or 3. First leaf deltoid, with straightsides and curved base ; apex acute. Second leaf broadly orbicular-cordate, with broad,open, rounded sinus and narrowly rounded angles. Third leaf similar to second,larger, with narrower, more acute sinus. First floating leaf orbicular to orbicular-ovate,entire, blotched more or less densely with red-brown above, under surface red on agreen ground.form.)Source of Data.Caspary, 1882..Riverton flowers \(This paragraph describes the growth from a small tuber of the roseaLeaf of mature plant 15 to 60 cm. in diameter, circular or slightly elliptic, narrowlypeltate, length of pelta: depth of sinus = 1:9 to 17;lLength.cm.8.28.48.48.46.0Petals.6.85Riverton flowers < ' 5-76-5Width.cm.3-73-83-73-02-52.41-31-3texture rather firm and
i6oThe Waterlilies.leathery margin coarsely and irregularly sinuate-dentate, the teeth mostly angular and;obtuse, the larger sinuses as much as 1.3 cm. deep. Upper surface green, faintly brownblotchedwhen young ;under surface suffused violet or even dark purple veins ;prominent,primary nerves 10 to 12 on each side ; length of principal area : radius of leaf= 1 : 1.1 to 1.7. Sinus, depth: length of leaf =1:2 to 2.9, margins convex, more orless widely overlapping, or oftener touching and both curled upward, forming a raisedkeel 1.3 to 2.5 cm. high in place of the sinus; angles forming an acute tooth in serieswith the marginal serration of the leaf, but lying within the circle described by theleaf-margin. Stipules fused at base, long-lanceolate, tapering, acuminate.Rhizome ovate or cylindric-ovate, rather stout, protected by long hairs at summit,smooth below when dry ; producing no offsets during vegetative season.Measurements of Leaves.Leaf.Length.Breadth.Length ofLotirs.Length ofIYlta.Length ofprincipalArea.Numberof Veins.MatureMatureMatureCaspary, 1882Maturecm.36.331237-36034-4cm.333334-460cm.15-314-712.7cm.1.61.61-31.9cm.9.611. 111. 110II10Leaf (small tuber of var. rosea).Length ofMidrib.Lengthfrom Apexto Angle.Width.Depth.Sinus.Width.First submerged .Second submergedThird submerged .First floating ....First floatingcm.LIS1-51-752.01.6scm.1.02-453-252.03-0cm.0.7S2.22.92.72-5cm.0.81-41-71.4cm.o1-51.80-51.0Geographic Distribution.The island and town of Zanzibar, in permanent rainpools(Caspary 1877).Notes. Seeds of this species were received by Caspary from J. M. Hildebrandtin the middle' of April, 1874, packed in chalk. Planted in water at 22 R., they germinatedfreely in about two months, and in 1875 seven strong plants came into flower;these were the types on which the species was established. In herbaria, however, it isalmost indistinguishable from N. capensis, and they form fertile crosses ;we have thereforereduced this one to varietal rank. It hybridizes readily with all the members ofthe Brachyceras group.Seeds developed late in the season will withstand the winters of Philadelphia andvicinity, germinate spontaneously, and form blooming plants the following summer;but the flowers are produced too late to ripen seed for the succeeding season. Thesame is true of N. caerulea, whose seeds have lain two years before developing plants.Planted in small pots, N. zanzibariensis will produce perfect flowers of any desired sizedown to 2.5 to 3 cm. in diameter.
CARNEGIE INSTITUTION OF WASHINGTON. WATERLILIES, PLATE 12.NYMPH^EA SULFUREA.
Taxonomy Nymphaea stuhlmannii. 161When raised from seed, itsports freely into the two following forms, withnumerous intermediates :Form azurea Hort, with light blue flowers, sepals lacking thepurple shadings form; rosea, with pink or reddish flowers, sepals green, under side ofleaves shaded reddish. This form appeared first with Herr L. Graebener (1887) in1884 in the Botanic Garden of Carlsruhe, although the first published description wasby W. Siber in 1887. Graebener claims to have obtained it at will by crossing N. zanzibariensiswith N. dentata, though many of the hybrid seedlings came blue.Casparydid not, nor do we, accept this theory of its origin. The experiment needs to be successfullyrepeated, but we have not had opportunity to try it. [Cf. also Revue Horticole,1890a, and Rev. de l'horticult. beige et etrangere, 1890, May 1 (not seen).]Nymphaea sulfurea Gilg.(Plate XII.)"Rhizome stout, conical.Petioles long [38 to 46 cm.] and slender; leaves deeplycordate, i. e. narrowly triangular excised ; outline broadly ovate or suborbicular, entire ;subcoriaceous, glabrous, reddish above, [deep] red beneath; primary veins 6, slightlyprominent, outer veins sub-inconspicuous or invisible. Flowers 4.5 to 7 cm. in diameter,deep sulphur color, sweet scented; sepals lanceolate, acute, purple or purplish onboth sides, with many slightly prominent parallel nerves; petals equaling the sepals,but often a little narrower, shorter and more acute ;stamens 40 to 50, linear ;antherselongate, connective evidently elongate and dilated; carpels 12 to 14." Leaves 4.5 to 5.5 cm. long and about as wide, the sinus being usually about 1 to1.5 cm. deep. Sepals 2 to 3 cm. long, 1 to 1.5 cm. wide. The deep sulphur yellowpetals are 2 to 2.8 cm. long, 0.7 to 1.2 cm. wide. Anthers bright yellow." (Gilg, 1903.)Rhizome 6.4 to 12 or 15 cm. long, about 2.5 cm. across, only one-third of whichis flesh, the rest being a dense layer of black hairs leaf bases; many, prominent ;therhizome was evidently erect but not tuberous. In the specimen at Kew the sepalsare copper-red outside, greenish-yellow inside. Petals 13, cadmium-yellow.N. sulfurea, Gilg 1903.Distribution and Types. Coll. H. Baum, No. 657, Jan. 17, 1900, Kunene-Sambesi-Expedition, " am linken Longa Ufer bei der Imbala, Minnesera, in flachensumpfigen Graben," altitude 1,250 m., in hbb. Berlin, Kew, Delessert, Munich, BritishMuseum. Also No. 325, October, 1899, " in der Nahe des Kuebe, in einem Bache,welche in einem Sumpfe entspringt und in den Kuebe mundet, nicht im Kuebe selbst,"altitude 1,150 m., in hbb. Delessert, Berlin; this number has larger leaves and flowersand broader petals than the former, with the leaves plain reddish-brown above ;in No.657 the leaves are blotched all over the upper surface with large reddish-brown blotches.No. 325 is mentioned first in Gilg's text, and appears, from the color of the leaves, tobe especially designated in the descriptiontranslation) ;(our first two paragraphs are a nearly literalin case of separation of the forms, No. 325 would have to stand as the type.Nymphaea stuhlmannii (Schw.) Gilg. (Fig. 62.)Sepals yellowish green, petals bright sulphur yellow, stamens orange yellow, withsulphur yellow anther. Stigma orange. Receptacle brown.N. lotus var. stuhlmannii, Schweinfurth in Engler 1895.N. stuhlmannii, Gilg 1903.13
162 The Waterlilies.Description. Flowers 10 to 15 cm. across, sweet scented (" duftet sehr aromatisch"). Sepals 4, obovate, broadly rounded above, with numerous longitudinal veins,8.3 cm. long by 2.5 cm. wide. Petals about 22, broadly obovate, with one to severallongitudinal veins. Stamens about 125, stout and thick, all appendaged, the innermostvery shortly so. Carpels about 23 ; stigma shallow, flat. Leaf orbicular-ovate, entire,21 cm. wide by 25.5 cm. long; sinus margins nearly straight, lobes diverging, obtuse;green on both sides ;veins prominent ; primary veins 7 ; principal area 7 cm. long.Via. 88. Nymphaea ttuhlmannii: Flower, under side of leaf, and outer stamen. Drawn by J. Pohl, from thetype in hb. Berlin, by kind permission of the Directors. Reduced one-half ; stamen natural size.Distribution and Type. Coll. Stuhlmann, No. 410, July 16, 1890, at " Uniamweni,Gunda mkali, bei Bibisande," Africa, altitude 1,200 m.,in hb. Berlin.Notes. A single sheet of this plant exists in the Berlin collection it has one; leaf,two separate open halves of a large flower, and one small wrinkled flower enough toshow that a splendid garden plant is yet to be introduced.
Taxonomy Nymphaea mexicana. 163GroupII. NYMPHAEAE SYNCARPIAE (Symphytopleura, Casp. 1865, 1878, 1888).Carpels completely fused with one another at the sides attached also to the axis;of the flower and to the torus as in Group I. Stamens not Flowersappendiculate.white, rose, purple or yellow, never blue nocturnal or diurnal.;Subgenus 3. CASTALIA DC. 1821.Flowers diurnal, usually floating. Sepals not evidently veined. Stamens insertedin series wHh the petals and grading into them in size and shape; innermost stamenswith narrow filaments. Styles linear, more or less ligulate. Leaves entire or sinuateor crenulate, never sharply dentate. Rhizomes not protected against drought, butlying dormant in cold weather. Plant glabrous except on the rhizome and bases ofpetioles and peduncles. Seed smooth. Seven species, found in all of the North TemperateZone, except the Pacific Slope of North America.Castalia and part of Lotos, Planchon 1852 b, 1853 b.Sec. II. Inappendiculatae, trib. III. Castalia (in part), Lehmann 1853 a.Castalia and Xanthantha, Caspary 1888.A. XANTHANTHA Caspary 1888.Flowers yellow, opening near noon and closing in the late afternoon. Outermoststamens first to dehisce. Pollen smooth. Seed very large. Floating leaves sinuate,more or less blotched with reddish brown. Principal air-canals in peduncle 4, mpetiole 2; idioblasts in course of petioles and peduncles stellate only. Rhizomeserect short, giving rise to numerous long stolons or runners which propagate the plantvery widely during summer, but produce in autumn on geotropicstolons characteristicbrood-bodies, consisting of a row of buds and a cluster of short fleshy roots (resemblingthose of Limnanthemum). One species,native in Florida, Southern Texas, andMexico.Nymphaea mexicana Zuccarini. (Plate XIII; Figs. 63, 64.)Flowers 6 to 13 cm. in diameter, bright yellow; sepals and petals elliptic, ovateto lanceolate, acute or rather obtuse. Receptacle with four swollen ridges Leavesfloating or aerial, orbicular to ovate, narrowly peltate, nearly entire or denticulate-wavy,10 to 18 cm. in diameter, with the petioleinserted much nearer the lobes than the apex ;upper surface blotched with brown, at least when young; lower surface deep purpleor purplish-green (green in aerial leaves), with small blackish spots.Nymphaea mexicana, Zuccarini 1832a and b, fid. original specimens ,., .oil. K^winsk^Sept"aus dem See bei Mexico," in hb. Munich. Planchon 1853 *. Hemsley 1888 Pringle1890. Gerard 1890 b; 189:. Plank 1896? Tricker 1897. Conard 1901 a. Not
1 64The Waterulies.N. planchonii, Caspary MS. (flower) in hb. Berlin. This specimen bears several marks :(i) coll."Poeppig, No. 3033, 1832, Colufer Amazon, Nymphaea hepatica"; (2) "No. 10 Nymphaealobtusiloba striken out] tussilagifolia Lehm." in Lehmann's handwriting; (3) "Nymphaea(flore partim rubro, partim flavo). In lacu chalcensi prope Yotla (sit. Mexico),14 Mayi, G. Andrieux, Plant. Mex. exsicc. No. 541" The leaf is of N. rudgeana, theflower is of N. mexicana. No. 541 of Andrieux in hb. Paris is N. mexicana.Description. Flowers floating, or raised 5 to 12 cm. above the water when inshallow places, 7 to10 cm. in diameter, open from 11 a. m. to 4 p. m., with faint odorof a tea-rose or odorless. Peduncle terete, 3 to 6 mm. in diameter, 16 to 150 cm. long.Receptacle four-sided,the corners broadly rounded and prominent, the sides slightlygrooved two of the corner-ridges are larger than the alternate two. Sepals 4, elliptic;to lance-ovate, obtuse, length width = 4.5 cm. : :1.3 cm. ;outer surface green with palebrownish margins; seven longitudinal veins visible by looking through the sepal at aa b AFio. 68. N ympltaea mexicana : a, sepal ; b-e, successive petals ; /, 0, stamens ; h, section of ovary ; fc, sectionof fruit, a-h from Mexican (?) plants, cult, at Riverton, N. J. ; k, from Florida plant, in the Botanic Garden, Universityof Pennsylvania. Six-sevenths natural size.strong light; inner surface yellow; opening to about 15 above horizontal. Petals12 to 23, outermost of the size and shape of the sepals or somewhat narrower, obtuseor acute, slightly purplish without. Inner ones narrowly obovate, rounded at apex.Stamens about 50, stout, golden yellow, grading inward from petaloid to typical stamens.Innermost stamens short, anther about as long as or shorter than its filament andof the same width. Back of anthers convex, face deeply grooved between the anthercells. Pollen smooth. Carpels 7 to 10; styles short, fleshy, with a deep furrow onthe inner face. Stigma deeply curved-funnel-shape, the papillose rays ending roundlyon the base of the styles ;each ray with a short longitudinal median fissure (the suture).Axile process small, rounded, in the bottom of the narrow central pit of the stigma.Fruit spherical in Texan specimens, ovoid in those from Florida, 2.5 cm. in diameter,crowned with the small stigma 0.6 cm. in diameter. Seeds globose, very large, 0.48cm. in diameter, very finely appressed-hairy, dull greenish-black.Submerged leaves from brood-body 1 to 3, narrowly hastate, acuminate. Firstfloating leaves ovate to orbicular-ovate, much blotched above with reddish-brown ;sinusslightly open, or overlapping margin entire in apical portion, sinuate below, ;especiallyat angles under surface ;green, tinged with reddish-brown, with a few blackish specks.
CARNEGIE INSTITUTION OF WASHINGTON. WATERLILIES, PLATE 13.\(ANYMPH/EA MEXICANA.
Taxonomy Nymphaea mexicana. 165Leaves of mature plant floating, or aerial and 7 to 12 cm. above the water,broadly ovate to orbicular, evidently peltate, entire, or more commonly sinuate or crenulate-wavyat base, becoming entire or slightly emarginate at apex ; green above, withbrown mottlings, at least on the floating leaves ;under surface deep purplish-red, oftenbecoming greenish, marked with small blackish dots. Primary veins scarcely visible,5 to 10 on each = :side, depressed from the second fork outward. Sinus short, depthlength of leaf 1 2.6 to 2.8 : ; margins nearly straight, slightly separated or overlappingangles slightly or evidently produced, obtuse or somewhat acute, made prominentby a deep concavity of the margin of the leaf on outer side of lobe. The aerial;leaves may become concave above by the overlapping of the sinus margins. Petiolewith two main air-canals placed transversely, and two smaller ones placed anteriorlyand posteriorly; stellate cells small,numerous, short armed. Stipulesfused into a single entire lanceolateorgan, thin, membranaceous andtranslucent; apex rounded; 3.3 cm.long by 0.8 cm. wide, standing erectand clasping the younger leaves;smooth, and uniform in thickness.Rhizome erect (1.9 cm. longby 1.4 cm. thick), becoming asmuch as 30 cm. long and 5 to 6 cm.thick (Treat 1877), densely coveredbetween the leaf bases withlong, light-colored hairs, and on thesides marked with large, projecting,spirally arranged leaf cushions,which are about 0.64 cm. apartfrom center to center. From theupper part of the rhizome nakedstolons 0.5 to 1.0 cm. in diameter areFig. 64 Nymphaea mexicam 1 a, first broad leaf , b, acicularleaf of seedling ; c, d, e, submerged leaves of seedlings ; /, firstfloating leaf of seedling. Florida stock. Natural size.given off ; these pass along under the mud in which the plant grows for a distance of 15or 20 cm. to 1 meter, when the terminal bud turns upward and forms a new plant ; in afew weeks the new plant again sends out runners, and the connection with the parentstem dies away. In autumn the stolons become geotropic and bury their tips 20 cm.or more in the mud, and there form brood-bodies. A brood-body consists of an axis2.5 cm. long, with a row of 3 to 6 or 7 upward pointing buds on one side, and acluster of about a dozen or more fleshy roots 1.3 to 3.8 cm. long and 0.4 to 0.48 cm.in diameter hanging down like a " hand " of bananas on the other side. In spring, theuppermost bud sends up a naked stolon at whose summit leaves spring up as notedabove; roots spring from the bases of these leaves, and soon an independent plantisformed. If the first bud is broken off, the second sprouts, and so on ; probably the secondand lower buds would sprout sooner or later in any case, so long as the food sup-was not used up.ply in the fleshy roots
1 66 The Waterulies.Measurements of Floral Parts in centimeters.BIPILB.
Taxonomy Nymphaea tetragona. 167Geographic Distribution. Florida, Texas, and Mexico. "Lagoons of thelower Rio Grande southward to the Valley of Mexico, and westward to the valley ofthe Lerma, State of Jalisco" (Pringle 1890). Lake Xochimilcho, Valley of Mexico,coll. J. W. Harshberger. Brownsville, Texas, coll. Pringle, Flora of Texas, 1888, No.1956. In Florida : St. Johns River, from Jacksonville southward for 40 miles (Treat,1877, fid. original specimens) ; Curtis, North American Plants, No. 101 a and SecondDistribution No. 4815. Miami River; Prairie Creek (both in hb. Acad. Nat. Sci.Phila.). Little River, coll. Dr. J. M. Macfarlane, cult, in Botanic Garden, Univ. ofPenns) 'r-vania. Reservoir Pond, Palm Beach, coll. Mr. Thomas. Also coll. Rugel, " adfl.Alachua, Florida or." July, 1848, in hb. British Museum.Notes. Zuccarini's description of N. mexicana ascribes white flowers to it,whereas the leaf agrees with that given above. Examination of the types leaves nodoubt as to the identity of the plants now so familiar in cultivation ; the white colormust have been purely a guess, as Professor Radlkofer suggested to me. The driedspecimens are brown like the average herbarium specimens of the European waterlily.Our conclusion to unite the Florida and the Mexican plants comes only after prolongedconsideration and comparison, and it is to be regretted that the very suggestivename N. fiava has to be abandoned. It has seemed to us that the leaves of the Mexicanplant are thicker and rounder, with less of purple coloration beneath. The floralleaves also are a little broader than those of the Florida plants. A fruit on Pringle'sNo. 1956 in hb. Acad. Nat. Sci. Phila. seems to have been nearly spherical, while thefruits of the Florida plants in our garden were ovoid. We have not been able tosecure living plants direct from any Mexican locality. The Prairie Creek specimens,collected by Garber in 1878, have leaves more nearly orbicular than those from otherFlorida localities ;this isprobably because they grew in shallow water. We see in cultivationlike variations from this cause.The name N. fiava Leitner rested on the floral decorations and name on Audubon'splate of the great white swan; the leaves accompanying the yellow flowers areevidently those of a Nuphar. Excepting Rugel's unrecognized specimen, the species wasknown only by that plate until Dr. Edw. Palmer in 1874 sent a dried specimen to Harvard,with a statement of itsyellow color. Mrs. Treat, however, in 1876, again discoveredthe plant, and made it known by sending ample material both fresh and driedto Harvard ;from there it was sent to Kew, where it flowered first in July, 1882.B. CHAMAENYMPHAEA Planchon 1853ft.Flowers white or rosy, opening near noon and closing in the late afternoon.Innermost stamens first to dehisce. Pollen aculeate or tuberculate. Seed of mediumsize. Leaves entire, obovate to oval. Principal air-canals in peduncle 4, in petiole 2;idioblasts stellate only. Rhizome erect, short, entirely without offsets (at least in N.tetragona) .Native of the North Temperate Zone, especially in China and Japan.Nymphaea tetragona Georgi. (Plate XIV; Fig. 65.)Flower small, 2.5 to 5 cm. across, white, with a small number of floral parts ;receptacledistinctly tetragonal. Leaves ovate, small (2.5 to 12 cm. long), green abovewith brown blotches when young, under surface dull red; sinus open, lobes acute.Petiole with 2 main air-canals. Rhizome erect, covered with projecting leaf-scars.
1 68 The Waterlilies.Nymphaea Utragono, Georgi 1775, Ad. original specimen from hb. Pallas, in hb. British Museum;:8co. Caspary 1865. Robinson 1896. Conard 1901 a.N. tetrogonanthos, Pallas MS., in Sims 1813, Ad. specimen cited above.N. pygmaea, Alton 181 1. Sims 1813. Smith 1819. DeCandolle 182 1 b, 1824. Sprengel 1825. Don1831. Ledebour 1842. Turczaninow 1842. Planchon 1852 b; 18530 and b. Lehmann1853 a. Hooker & Thomson 1855. Loudon 1855. Hooker 1872. Garden 1883 b. Tricker1897. Moenkemyer 1897.N. foliis cordatis integerrimis and N. alba minor, Gmelin 1769.N. pygmaea alba, Hort, Gerard 1890 a?Castalia pygmaea, Salisbury 1806 b.C. Leibergi, Morong 1888, Ad. original specimens. G. & F. 1888.C. odorata var. minor, Macoun 1891 (in part) Ad. specimen coll. J. M. Macoun, July 17, 1886, No. 94from hb. Geol. and Nat Hist. Survey, Canada, in hb. British Museum. Not N. odoratavar. minor Sims 1814.C. Utragona, Lawson 1889. Britton & Brown 1897.Fia. 66. Nymphaea tetraaona : a, section of ovary ; b, section of receptacle ; c, outer, d, inner sepal ; e,outer, /, median, 0, innermost petal ; h, stamens. Natural size.Description. Flower floating, 3.3 to 7.6 cm. across, open on 3 or 4 days fromnoon until 5 p. m., sweet-scented (or odorless, Britton & Brown 1897). Peduncleslender, 0.3 to 0.4 cm. in diameter, terete,10 to 30 cm. long, with 4 main air-canals.Receptacle enlarging abruptly from the scape, distinctly tetragonal, with rounded angles,green. Sepals 4, ovate to oblong-ovate or broadly lanceolate, breadth :length =1:1.16 to 3.61, obtuse or slightly acute; opening horizontally or less, putrescent infruit; green and veinless outside, white within; when dry, marked outside withshort elevated lines. Petals 8 to 17, rather thin, pure white (or faintly stripedwith purple lines, Morong 1888), about as long as the sepals, oblong, obovate orbroadly lanceolate, or oblong-elliptic, apex obtuse or somewhat acute, narrowedtoward the base, grading into the stamens. Stamens about 40, with golden yellowanthers, shorter than the petals (stamens 12 to 16 in the American plant, in 3 or4 rows, running up the ovary about half way, Morong 1888) ; outer stamens shorterthan median ones; outermost filaments elliptic or obovate to lanceolate-oblong, verybroad, narrowed toward the base (inserted up to the summit of the ovary, Casp. 1865 ;Turcz. 1842) ;outermost anthers shorter than median and inner, cells diverging below,sometimes only one cell developed. Median stamen with filament as in outer, but narrower;cells of anther parallel. Innermost stamens very short, filaments slender, cells
'CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES. PLATE 14NYMPH/CA TETRAGONA.HEUOTVPE CO., BOSTON.
Taxonomy Nymphaea tetragona. 169of anthers as in median stamens. Pollen yellow, finely tuberculate, operculum smooth ;diameter 0.03 to 0.04 mm., average 0.034 mm. Carpels 6 to 8 (4 to 12, as hithertodescribed) styles thin, subcoriaceous, ovate or ;oblong-elliptic, short and blunt, breadth:length = 1 :1.25 to 3, concave; stigmatic area extending on their bases as an upraisedcushion, ending obtusely. Stigma little or not at all depressed. Fruit spherical, naked,green, crowned with the small stigma. Seed large, 0.3 to 0.2 cm. long by 0.16 to0.17 cm. in diameter, dark brown, smooth.Seeds which germinate in late summer form small plants, and at approach ofwinter, I*se their leaves and retire into a small tuber. These germinate, producingfirst a series of submerged leaves, then small floating leaves as follows :Submergedleaves broadly "obovate, rounded at apex, thin, pale green; sinus deep, margins nearlystraight near the petiole and parallel or diverging or slightly overlapping, curved outwardbelow to the broadly rounded lobes. First floating leaf obovate-oblong, roundedat apex; sinus overlapping, lobes narrow, rounded at angles, suggesting rabbit-ears;green above with a brown line around margin; under surface greenish-red, denselymarked with fine bright red marblings.Leaf of mature plant cleft to the petiole, more or less coriaceous, broadly oblong,ovate or elliptic, 5 to 10 cm. long by 3.3 to 7.6 cm. wide, entire, rounded at apex(rarely emarginate, Casp. 1865), dark green above, more or less mottled (at least whenyoung) with reddish-brown under surface ; green to dull red ; primary veins 5 or 6 oneach side of leaf, sunk in the tissues; length of principal area: radius of leaf = 1 :1.5to 1.9 (average 1: 1.3). Sinus depth: length of leaf = 1 : 1.8 to 2.2 (average 1:2),lobes unequal, margins nearly straight or slightly convex, touching, or diverging and1.9 to 3.8 cm. apart at periphery, angles more or less produced, rounded to sub-acuteor rarely acuminate. Astomatic area very small; stomata larger and not so crowdedas in Eu-castalia, scattered toward the point of insertion of thepetiole ;cells of upper epidermis large. Petiole smooth, terete,very slender, 0.24 to 0.4 cm. in diameter, 10 to 30 cm. long,with 4 main air-canals, 2 larger and 2 smaller, with 4 stillsmaller ones anteriorly and posteriorly, and one on each side;2 double bundles are present (1 anterior and 1 posterior) with3 or 4 smaller ones on either side; stellate cells projecting intothe 6 larger air-canals, rays very slender, long, densely coveredwith small crystals, quite brittle. Stipules oblong-ovate orelongate-triangular, acute, not at all keeled.Rhizome erect, 7 to 15 cm. long, blackish, covered with long hairs (1.3 cm. long) ;apex covered with leaf-bases and stipules.Leaves fromEntirely without branches or offsets.Small Seedling-Tuber (in centimeters).Measurements of Sepals(after Caspary, 1865).Breadth.
!7oThe Waterlilies.Geographic Distribution. Eastern Europe, Asia and North America, south toAustralia. Russia (Baikal region), Georgi 1775. Khasia Hills, and marshes at Nonkreem,alt. 5,600 ft., India, Hook. & Thorn. 1855, fid. specimens coll. Hooker, Duthie,Stewart, etc. N. Queensland, Australia, Midler 1887? Siberia, Gmelin 1769. Commonin China and Japan,fid. numerous specimens. Granite Station, Kootenai County,Idaho, U. S. A., coll. Leiberg, Heller and the writer. Misinaibi River and along SevernRiver, Canada (Ontario), Macoun 1891.Notes. Caspary (1865), from whom the measurements in the text were mostlytaken, divides the species thus:I. Lata: Sepals and petals shorter and broader, outermost petals slightly exceedingthe sepals ; depth of sinus less than half the length of the leaf. Leaves not spotted.Pollen smaller. Found in eastern and northern Asia.II.Angusta: Sepals and petals longer and narrower; depth of sinus equalto orexceeding half the length of the leaf; pollen larger. This varies into (1) orientalis,outermost petals a little shorter than the sepals ;lobes of leaves long-produced, rarelyacuminate ;leaves not spotted. China and Japan. (2) indica, outermost petals slightlyexceeding the sepals; lobes of leaves long-produced, tapering and acuminate; leavesusually brown-spotted above. Northern India (Nonkreem). Georgi's specimens areof the broader sepaled type. Those with narrower petals and sepals may deserve varietalrank as var. angusta Casp.The American plant, from itssmall number of stamens and their insertion belowthe summit of the torus, may deserve the rank of a variety.N. tetragona was introduced into England from China in 1805 by the East Indiain our climate.Company, in the ship " Winchelsea," Capt. Campbell. It ishardyBeing of small size it is suitable for aquaria.The seeds are used for food in Japan, and the leaf buds are also eaten, dressedwith vinegar (Rein 1886).Nymphaea acutiloba, DC. 1821, is of uncertain identity, but as it came fromChina it isprobably N. tetragona Georgi.Nymphaea fennicaMela.Flower cup-shaped, small, 3.8 to 6.9 cm. across, white or rosy ; receptacle large,square, sloping upward, with a keel running from the peduncle to each corner; sepalsforming a sharp angleat their line of insertion. Petiole with 2 main air-canals.Nymphaea fennica, Mela 1897, fid. specimens coll. A. J. Mela, in hb. Univ. of Penna. G. C. 1899.Description (taken from Mela's paper). Flower cup-shaped, small. Bud shortovate, square at base, constricted just above the base, then rounded and rapidly drawnin to the acute apex. Receptacle characteristic, very broad ;breadth of receptacle:length of sepals= 6 mm.: 14 mm. (small flower) ;= 9 mm. :19 mm. (medium) ;= 12of the budmm. :25 mm. (large flower). Looking at the receptacle with the apexturned away, the surface slopes gently, roof-like, from the top of the peduncle, with alow, narrow keel running to each angle. Edges of receptacle not projecting downwardas in N. Candida, but plane beneath, making a sharp angle on the side. In fruit thereceptacle is a little wider than the berry and calyx, these being completely hidden
Taxonomy Nymphaea fennica. 171when the fruit is viewed from beneath. Sepals elliptic-ovate,a little contracted belowthe apex, often very broad; length : breadth = 3:2 (18 to 19 mm.: 12 to 13 mm.) ;calyx somewhat contracted just above its origin, so that the edges of the receptacle,which are sharp-angled outwardly, appear to project. Petals 10 to 15, very concave,and crowded, with apex directed upward so that the corolla iscup-shaped; lanceolateor narrowly-lanceolate. Stamens about 50. Anthers very short and broad (3 mm. longby 1 mm. wide) ; filaments petaloid, with broad, elliptic, deep yellow blade and stronglycontracted, whitish claw ;blade often with 1 to 3 longitudinal folds, with 1 or 2 transversefolds between blade and claw. Stigma 5 to 9 rayed, with yellow, often violettipped,styles. Stigmatic surface usually (not always) deep violet, with generally 3-pointed rays. Fruit broad conical-ovate, broadest at base, but not so broad as thereceptacle which forms a square disc, whose edges (and especially the corners) extendnoticeably beyond the circumference of the fruit; the calyx persists as a 4-sidedpyramid surrounding the fruit.Leaf small, 7 to 12 cm. long by 5 to 9 cm. wide, elliptic-cordate lobes;acute orobtuse, margins overlapping or approaching or more or less wide apart (3 cm.).Primary veins of the lobes usually divergent, even when the lobes overlap, thoughthey are sometimes parallel or even converging. Petiole slightly flattened, with 2 largeair-canals.Rhizome erect, 5 to 10 cm. long, about 2.5 cm. in diameter, conical at apex, cov-One of our specimens has two equal branches.Geographic Distribution. Middle and East Finland, mostly in small forestered with long black hairs.ponds, sometimes in running water, from the shallows to a depth of 1 or 2 meters,often accompanied by large and small forms of N. Candida. Found also in SouthernFinland. (Cf. specimens coll. Ldnnbohm, Aug., 1898, No. 3601, " locus classicus," inseveral herbaria.)Notes. The receptacle and air-canals of petiole place this species in near relationOne isto N. tetragona; in several other respectsitapproaches N. Candida semiaperta.inclined to suggest that N. tetragona of China shades off through the Siberian formsinto N. fennica, this then runs into N. Candida, and itagain into N. alba of Germanyand England. A careful review of these forms, in the living state by cultivation andhybridization, as also in histological details, is highly desirable.The seeds of N. fennica have not as yet been described.Numerous stations for this species throughout Finland are given in Mela's paper.Red-flowered specimens occur "im See Keskimmainen im Kirchspiel Lapinlahti," andinPieni Musto and Jalajarvi.Through the kindness of Mr. Mela, we have a fine series of specimens of N. fennica;of several rhizomes sent to us alive, only one survives, and it has never bloomed.C. EUCASTALIA Planchon 1853 b.Flowers white or rosy to deep carmine, opening in early morning, closing fromnoon to mid-afternoon. Innermost stamens first to dehisce. Pollen aculeate or tuberculate.Seed of medium size. Leaves entire, nearly orbicular, of solid color above.Principal air-canals in peduncle 4, in petiole 4; thickened bipolar idioblasts more orless numerous in the tissues of leaf and flower stalks, this type of cell being absent
iy 2The Waterulies.from other syncarpous members of the genus.Rhizome horizontal, long and creeping,more or less branched. North temperate zone, south to Algeria, Cashmir, and BritishGuiana.Nymphaea Candida Presl. (Plate XV; Fig. 66.)Leaves floating, crowded on the rhizome, phyllotaxy about 13 on 34; lowest pairof veins on under side of leaf (veins of the lobes) curved toward one another.Flower odorless, white, 6.4 to 7.6 cm. across. Line of insertion of the sepals angular,projecting downward. All of the filaments broader than the anthers. Ovary muchcontracted and destitute of stamens immediately below the stigmas. Fruit ovoid, seedlarge.Nympkaeo Candida, Presl 1822, Ad. specimens coll. Topitz, Flora exsiccata austro-hungarica, No.1282, "in stagno Egersee ad pagum Sonnberg prope urbem Gratzen (locus classicus)."Reichenbach 1845. Planchon 1853 b. Schuchardt 1853. Celakovsky 1867. Caspary 1879.Grisebach 1882. Gerard 1892. Tricker 1897. Moenkemyer 1897. Conard 1901a.N. alba, Linnaeus 1753; 1762 (in part), and other European authors.N. alba oocarpa, Caspary 1855 = N. alba oligostigma Casp. 1863 ; 1865.N. biradiata, Sommerauer 1833, Ad. specimens coll. Sommerauer, in Tricbnersee in Styria in hbb./inI.Paris, Boissier, and coll. Wettstein, Flora exsiccata austro-hungarica, No. 1281. Coll. StatzerDorfler's Herb. Norm., No. 3502; coll. Keller in Magnier, Fl. select, exsicc No. 3453!coll. F. G. Strobl, " in aquis stagnantibus : Triebner vel Gaishornsee vallis Paltinae (locusclassicus)." Reichenbach 1845. Fries 1846. Planchon 1853ft. Schuchardt 1853.N. semiaperta, Klinggraeff 1848, Ad. F. Schulz, Herb. Norm. nov. ser. Cent. 3, No. 207, andspecimen coll. Caspary, Aug. 9, 1859, near Konigsberg, in hb. Boissier. Schuchardt 1853.N. neglecta, Hausleutner 1850, Ad. specimen coll. R. Fritz, July, 1870, in a pond near Rybnick,Ober-Schlesien, in hb. Berlin.N. Kosteletzkyi, Palliardi MS., Ad. Lehmann 1852 b, and specimens sent by Palliardi to Lehmann,from hb. Lehmann in hbb. Kew, Berlin. Schuchardt 1853.N. pauciradiato, Bunge, in Meyer & Bunge 1829 = N. alba Patrin, Ad. Meyer & Bunge,1. c ;Ad.specimens coll. Ledebour, 1845, in the Altai region, in hbb. Paris, Munich.N. intermedia, Weiker, in Reichenbach 1842.N. Basniniana, Turczaninow 1842, Ad. specimens sent by Turczaninow from hb. St. Petersburg, inhbb. Kew, Boissier, DeCandolle = N. bashiniana, Steudel 1841.N. Cachemiriana, Cambessedes in Jacquemont 1844, Ad. original specimensin hb. Paris = N. albavar. Kashmiriona, Hook. & Thorns. 1855. Hooker 1872.N. punctata, Kar. & Kir. 1842, Ad. Casp. 1879, from Semenow 1854, and specimens coll. Kar. &Kir. " in Songoriae lacubus prope fluvium Lapsa," in hbb. Berlin, Paris, DeCandolle.AT. alba, L. var. minor DC, Ad. specimen coll. Fischer, 1819, in hb. DeCandolle. Besler 1613, cf.Flora 1833. Schuchardt 1853. =AT. minor Reichenbach 1842; 1845. =N. permixta,Boreau 1859.N. Milletii, Boreau 1859, Ad. specimen coll. Ed. Longeron, June 12, 1861, Maine et Loire (locusclassicus?) in hb. British Museum.N. Wenzelii, Maack in litt, Ad. plate from Ruprecht 1859, in hb. British Museum.Description. Flower floating, 6 to 13 cm. in diameter. Sepals 4 (or 5) oblongor ovate-oblong (3.8 cm. long by 1.3 cm. wide), narrowed at apex, acute or obtuse,about7-veined, green outside, white within.Petals 12 to 20, white, oblong to oblong-lanceolate,obtuse or somewhat acute, 7-nerved, outer ones nearly as long as the sepals.Stamens 32 to 70, shorter than in N. alba, grading inward from the petals, orangeyellow;filaments lanceolate (outer) to linear-lanceolate (inner), long acuminate, nevernarrower than the anthers, usually inserted only on the sides, not on the summit, of
Taxonomy Nymphaea Candida. 173the ovary. Pollen granulate with smooth operculum, larger than in N. alba. Ovaryovate or roundish, usually contracted and destitute of stamen insertions below thestigma. Carpels 5 to 14; styles linear, flattened, breadth :length = 1 :1.5 to 2.5 ; apexrounded or somewhat acute. Stigma concave, yellow, red or violet, the rays terminatingusually in 3 teeth (a central and two lateral) on the base of each style. Axile processslender. Fruit ovoid or spherical ;in the former case, narrowed below the stigma,which stands as it were on a short thick column surmounting the berry; greenish orreddish; subtended by the persistent sepals; diameter 1.9 to 4.4 cm.in N. albit, ellipsoid, brownish, 0.3 cm. long aril ; longer than the seed.Leaf cleft at base to thepetiole, sub-orbicular to oval, 10 or12 to 30 cm. long and 25 cm. wide,entire. Sinus, depth length of leaf= : 1 :2.5 lobes ; nearly equal, innermargins curved, overlapping a littleabove, and spreading out towardperiphery of leaf (or straight, andparallel or touching) angles more;or less acute. Veins on under surfaceof leaf prominent, the lowestpair (i. e., those runninginto thelobes) curved, and, if produced,would cross, inclosing an oval area.Petiole terete, smooth.Rhizome stout, 5 cm. or morein diameter, horizontal, with stronglateral roots, but often without sidebranches; apex surrounded by petioles,peduncles and stipules. Phyllotaxyabout 13 on 34 with veryshort internodes ;the surface of therhizome may almost be said to consistof large leaf-cushions.Geographic Distribution.Northern and arctic Europe andAsia, south to the mountains of Germanyand the Himalayas. Krumme-See, near Grubno, in West Prus-Seed larger thanFig. 66. Nymphaea Candida : Under side of leaf of specimenin ho. British Museum, from the classic locality (Flora exsiccataaustro-hungarica. No. 1882), by courtesy of the Directors. P.Highley, del. Reduced one-quarter.sia (Caspary 1884). Canton St. Gallen, in Switzerland (Gremli 1882). Lojo inPoland (Lindberg 1881). Herjedalen in Sweden (Behm 1887). Far north in Finland(Saelan 1879). Egersee, near Gratzen, in Austria (Kerner 1886). Koniggratz, inBohemia (Hansgirg 1880). Veszto, in Hungary (Borbas 1881). Ascends 500 metersin southern Bavaria (Schenck 1885). East slopes of Urals (Cler 1873). Triebnersee,Palatinathal, Austria (Kerner 1886). Bohemia, in ponds and ditches (Presl1822). Austro-Hungary (Leopoldskron, coll. Stohl), Flora exsiccata austro-hungarica,
174The Waterlilies.No. 1283. Franzensbrunn :3 Nomenteiche, coll. Caspary, Aug. 25, 1855, in hb. Kew.Lakes of the Vosges near Sembach, Bavaria, coll. Koch, July 20, 1859. Hammerteichnear Falkenberg, Ober-Schlesien, fid. specimen in hb. Kew. Teplitz in Bohemia, coll.Winkler, 1853, in hbb. Boissier, DeCandolle. Near Stockholm, Sweden, coll. Ljogruen,1845, in hbb. Munich, DeCandolle. Lithuania, coll. E. Rudominowna, in hb.Kew. Neusattel near Ellnbogen, coll. Caspary & Ortman, 1855, in hb. Kew. Schwarzseenear Kitzbuhel, Tyrol, coll. Traunsteiner, No. 1608, marked " N. alba," in hbb.Boissier, Paris. Lakes of the Baikal region, Siberia, coll. Fischer, 1836, in hb. Paris.Cashmere Lake, coll. J. E. Winterbottom, May 13, 1847, and cou Falconer, both-marked " N. alba" in hb. Kew.Notes. The species varies widely as indicated above in the description, and asmay be inferred from the lengthy synonymy. Presl's distinctions were not widelyknown, or were lost sight of by most botanists. He separated N. Candida from N. albalargely on account of the absence of stamens from the summit of the ovary. Thesame name had been used for a European white waterlily by Tragus, Fuchs, Caesalpinusand others. N. pauciradiata was characterized byits small flowers and few carpels(8 to 9). N. biradiata was noticed on account of the blood-red star on thestigma. N. basniniana was separated from N. alba on account of having oval leavesinstead of round, and its small number (9) of carpels. N. cachcmiriana has also only7 to 8 carpels the; original description is quite complete. N. semiaperta has as its outstandingtrait the fact that even at noonday the flowers remain only half open. N.neglecta was marked off from alba by its small oval fruit, few rayed stigma, and theconverging course of the lowest pair of veins of the leaf; from Candida in havingpetals longer than the calyx, and ovary only one-third bare, instead of two-thirds asin Presl's Candida; from semiaperta by smallness of ovary and color of stigma. It isinteresting to notice how various observers have been impressed with different featuresof the plant as they appear more or less exaggerated in certain localities.Probably the best disposition of the case is that of Caspary in1879, when he firstgranted specific recognition to this group of forms. His paper of 1855 placed theseplants in variety oocarpa of N. alba, with the following divisions:N. alba var. oocarpa.E. angustata styles longer and narrower; KonigswerthF. semiaperta styles shorter; flower opening one half.in Bohemia.a. erythrocarpa fruit reddish (=N. semiaperta and biradiata)b. chlorocarpa fruit greenish (= N. neglecta)G. aperta:a. erythrocarpa (=N. kosteletzkyi and cubogermen, from Franzensbrunn)(=N, Candida Presl from Pardubitz in Bohemia)b. chlorocarpa (=N. Candida Ortm. from Ellnbogen, Bohemia)(=N. Candida Presl from Pardubitz, Bohemia)The name oocarpa was changed by the author to oligostigma in 1863, and finally,on discovering that crosses between N. alba oocarpa and N. alba sphaerocarpa weretrue hybrids, with imperfect sexual apparatus, the two were made distinct species, andthe following disposition of the one in hand was given, with a few, mostly Swedish,localities.
Taxonomy Nymphaea alba. 175N. Candida Presl.Var. I. oocarpa; fruit ovoid, higher than broad.1. aperta; sepals and petals opening to form an obtuse angle.A. xantho stigma; stigma yellow. Nerike; Fagertarn; E. Prussia; Konigsberg; afew lakes in West Prussia.B. erythrostigma; stigma more or less dark crimson, especially near apices of stigmaticrays. Sodermanland and Norbotten.a. erythrocarpa. Norbotten.b. chlorocarpa. Norbotten.2. .iimiaperta; sepals and petals forming an acute angle when open;fl. campanulate. Prussia,east of the Weichsel, and Bohemia.Erythrocarpa and chlorocarpa, N. biradiata and N. semiapcrta.Var. II. sphaeroides; fruit spheroidal, broader than tall.1. aperta; sepals and petals opening to form an obtuse angle.A. erythrocarpa. Norbotten.B. chlorocarpa. Norbotten.2. semiapcrta; sepals and petals forming an acute angle when open;fl.campanulate.A. erythrocarpa. Norbotten; Prussia.B. chlorocarpa. Prussia.N. alba X Candida occurs in East Prussia at Neuhausen near Konigsberg, nearGoldap, and elsewhere (Caspary 1879). Sernander (1894) claims to have found ared variety of N. Candida in Fagertarn, Sweden.Nymphaea nitida, Sims 181 1 a, with entire, suborbicular leaves, with obtuse lobes;flowers white, cup-shaped, may belong here. Its habitat is unknown, and it has notrecently been positively identified anywhere. The figure of the Botanical Magazinewas " drawn at Mr. Vere's garden " in Kensington-Gore, Mr. Anderson, gardener. Aspecimen exists in hb. Caspary (now Berlin), which was presented by Prof. M. T.Masters in 1854 from the botanic garden of Oxford. Here it was received in 1840from Kent, whither it came directly from Anderson. This specimen appears to belongto N. Candida as here defined.Nymphaea alba (L.)Presl.Leaves nearly orbicular, floating, red when very young, 10 to 30 cm. in diameter,crowded on the rhizome, phyllotaxy about 13 on 34 lowest;pair of veins straight anddiverging. Flower odorless, pure white (in the type), 10 to 12 cm. across, opening fully.Line of insertion of sepals rounded, not prominent. Stamens covering the ovary tothe summit. Filaments of inmost stamens not wider than the anthers. Axile processspherical. Fruit more or less spherical, seed small.Nymphaea alba, Linnaeus 1753, fid. original specimen in hb. Linnaeus, without data; 1762 (in part).Willdenow 1797 (in part). Aiton 1789; 1811 (in part). Smith 1800; 1809 (in part), andall writers previous to 1822, and many since.N. alba, Presl 1822. Reichenbach 1845. Hentze 1848 a; 1852 a, b. Schuchardt 1853. Caspary 1879.De Halacsy 1900. Conard 1901 a.N. alba melocarpa, Caspary 1855 = AT. alba polystigma Casp. 1863.N. Aesopii, Orphanides MS. in Boissier 1888; fid. specimens coll. T. G. Orphanides, Flora Graecaexsiccata, No. 1051 (hb. Orphanid., No. 264) Feb., 1862, "ad lacu Baldovo supra VodenaMacedoniae," in hbb. Paris, Boissier.N. Dioscoridis, Heldr. MS., in Boissier 1888; fid. specimens coll. Heldreich, Herbar. Flor. Hellenic,May 23, 1878, at " Aetolia : in lacu Lysimachion," in hb. Boissier.
176The Waterlilies.N. venusta, Hentze 18486; 1853 a, fid. specimens coll. Hentze from hb. Lehmann,Berlin.in hbb. Kew,N. rotundifolia, Hentze 1848 b; 1852 a, fid. specimens coll. Hentze, from hbb. Lehmann and Caspary,in hbb. Kew, BerliaN. erythrocarpa, Hentze 1852 6 = AT. alba Hentze 18480; 18520, fid. specimen coll. Hentze, fromhb. Lehmann, in hb. Berlin.N. parvifiora, Hentze 1848 b; 18520, fid. specimens coll. Hentze, from hb. Lehmann, in hbb. Kew,Berlin.N. spUndens, Hentze 18480; 18520, fid. specimens coll. Hentze, from hb. Lehmann,Berlin.in hbb. Kew,N. urceolata, Hentze 18486; 1852 a, fid. specimen from hb. Lehmann, in hb. Berlin.N. alba var. intermedia Gave 1892; fid. original specimen coll. P. Gave, June 30, 1891, St. Andre,Haute Savoie.Castalta speciosa, Salisbury, 1806 a. Britten 18880.C. alba, Woodville & Wood 1806. Link 183 1. Greene 1888. Lawson 1889.Description. Flower floating, 7 to 15 cm. in diameter, open on 4 days from 7a. m. to 4 p. m. (7 a. m. Upsala; 8 a. m. Innsbruck, Kerner 1895), (5 p. m. Schulz1890), faintly sweet-scented on first day of opening, odorless afterward. Peduncleterete. Receptacle rounded along the line of insertion of the sepals. Sepals 4 (rarely 3or 5, Eichler 1875) oblong or ovate, obtuse or somewhat acute, smooth, opening outhorizontally or more, putrescent in fruit, green mostly shaded with reddish-brown outside,whitish or pale reddish within, stiff, obscurely 7-nerved.Petals 12 to 24, of ratherfirm texture, outermost opening horizontally, grading inward in size, shape and positionto the stamens. Outermost petals about as long as the calyx, lanceolate to oblong, orovate, obtuse, white, often striped with green, gray, or reddish on the back Stamens 64to 100 or more, covering the torus to its summit, innermost first to ripen. Outer ones 15to 17 mm. long, innermost 4 to 5 mm. long. Anthers sulphur-yellow to orange-yellow,sickel shaped with the convexity inward, inserted at an angle with the filament, theangle being slight in the outer stamens, reaching 90 in the innermost; lines of dehiscencefacing inward and downward ;outermost anthers 10 to 12 mm. long, innermostabout 3 mm. Filaments curved, convex side outward, bringing the anthers oerthe middle of the flower ;outer filaments white, petaloid : innermost filament fifitnnn,Pollen aculeate, operculum smooth; 0,038 mm. inyellow, narrower than the anther.diameter. Ovary slightly contracted below the stigma, covered with stamens to thesummit. Carpels 8 to 24 (mean = 12 to 20) ; styles fleshy, triangular-ovate, roundedat apex, more or less concave and 1 to 5 sukate on inner side, subphur yeDow, oftentinted orange.Stigmatic area scarcely produced above to the bases of die styles, endingacutely. Stigma more or less deeply depressed, basin or funnel shape, sulphuryellow.Axile process conical, acute, yellowish-white. Fruit globose or depressedglobose,crowned with the styles, which are bent sharply inward. Inner wall ofcells whitish, spongy tissues ; green or blood-red. Rqe fnrit yellowish-greendevoid of floral leaves.-^Staf 3 mm, long by 2 mm. in diameter , *lKpni^i odark olive green, smooth ; raphe evident. Aril in closing the seed, open at end
Taxonomy Nymphaea alba. 177Size of Leaves.Length.
178The Wateri.ilies.panicd by leaves of the usual size; it was collected at two stations near Lich. All ofthese were supported by numbers of specimens in all stages of flower and fruit.In Greece, N. aesopii of Macedonia was separated off by Orphanides on accountof itshuge size, and N. dioscoridis from Lake Lysimachia in Aetolia for its smallness.Caspary began his work on waterlilies by reviewing this species. He rightlymaintained that the smallness or largeness of the plant is not a safe guide to classification,but that there were other characteristics of a more profound and constant natureon which divisions could be founded. Accordingly in 1855 he gave the followingarrangement :N. alba.I. melocarpa.A. dcpressa.II.a. chlorocarpa N. venusta Htze.= JV. rolundifolia Htze.b. erythrocarpo = N. erythrocarpa Htze.B. circumvallata.C. sphaerocarpa.a. chlorocarpa.a. ftava ==N. parviHora Htze.= N. alba Htze.P. splendens= N. splendens Htze.b. erythrocarpa.D. urceolata = N. urceolata Htze.Oocarpa (see under N. Candida Presl).In 1863 Caspary changed the main headings, substitutingI.Polystigma and II.Oligostigma in the above table. Finally in 1879, as referred to under N. Candida, heraised the two varieties to specific rank under the names of N. alba (L.) Presl andN. Candida Presl, dividing the former species and giving a few localities, thus :N. alba.I. sphaerocarpa. Fruit subglobose.1. platystigma. Stigma more than half the diameter of the fruit (commonest form).A. chlorocarpa. Fruit green (widely distributed).a. Aava. Stamens sulphur color.b. splendens. Stamens orange.B. erythrocarpa. Fruit more or less suffused crimson.a. vulgaris. Petals white, or outer ones pale rosy.b. rosea. Petals and stamens deep crimson == N. alba and rosea Hartm. Handb.Skand. Fl., 1864, p. 76.2. engystigma. Stigma, measured from the backs of the processes, equal to or less thanhalf the diameter of the fruit (rare) ;West Prussia, near Danzig.II. dcpressa. Fruit depressed-globose ; alt. : lat. = 2:3 to 5:6.1. chlorocarpa here and there in Germany.2. erythrocarpa here and there in Germany.III. urceolata. Stigma deeply infundibuliforin in fruit, sunk in to middle of fruit in Germany,very rare. Chlorocarpa and erythrocarpa.IV. oviformis. Fruit ovoid ; upper one-fourth to one-third of height destitute of filaments insome lakes of West Prussia.
:--; e r~ "." :. : r -:- :--:. "E-_ . E5 "_i _ E ':NYWHvEA ALBA RUBRA,A (M7 E--.-T1 rf?
Taxonomy Nymphaea odorata. 179In all of these varieties, both larger and smaller forms occur. This division hasbeen adopted by some botanists, but is decidedly too cumbrous for practical use. Wehave therefore inserted the most important variety and the only one requiring specialnotice, namely, the pink Swedish waterlily, under its usual name, Nymphaea alba vanrubra.Var. rubra, Lonnroth. (Plate XV.)Outermost petals rosy, intermediate intensely rosy, innermost deep carmine-red ;anthers and stigma yellow; filaments and styles orange to deep red-brown.depth and purity of color, deepening on second and third days of flowering.Variable inNymphaea alba var. rubra, Lonnroth 1856. Liebman & Lange 1874. Garden 1879. Hooker 1884,fid. original specimen from Kew Garden, in hb. Kew. Andre 1884.N. alba var. rosea, Hartman. Trimen 1872. Masters 1878. Hemsley 1879. Tricker 1897. Conard1901a.N. alba sphaerocarpa platystigma erythrocarpa rosea, Caspary 1879.N. alba sphaerocarpa rubra, Caspary 187 1.N. alba van, Fries, 1858.N. alba var. purpurea, Fries MS. in Herb. Norm., fid. specimen, coll. T. G. Gjobel, " Nerica, in lacuFagertarn, par. Hammar," in hb. Kew.JV. sphaerocarpa, Carriere 1878. Tricker 1897.JV. sphaerocarpa var. rubra, Duchartre 1877.JV. Caspary, Carriere 1879 & (JV. Caspari Carriere 1880; JV. Casparyi Carriere 18Description. Flowers 7 to 15 cm. in diameter, opening pale pink, changing torose-pink or even deep red, often with a bluish cast; the color appears as linings ona white or pink ground. Petals narrower than in the type, obtuse or rounded.Rhizomes strong, branching as in the type.Geographic Distribution. Lake Fayer (Fagertarn) in the Parish of Hammar,Nerike, Sweden, collected by B. E. Kjellmark in July, 1856.Notes. Introduced by Caspary in 1871 and later by Froebel of Zurich, and soldin England in 1878 at 5 per root. Requires cool weather and water, and is difficult ofcultivation, at least in this country. Flowered first in America by Mr. Hovey of Boston,and later by Wm. Tricker at Dongan Hills, N. J. It 'was received by Caspary in1863, and in the Botanic Garden of Konigsberg was found to produce highly fertilecrosses with the type (JV. alba sphaerocarpa) ;a close relationship was thus proven.E. Fries, in a letter to the " Botanische Zeitung," published in 1858 (p. 73),refers to this as a Nymphaea with rosy-purple flowers, giving to its native pond inNerike, the. name of " Fagersjo," " the beautiful lake." He considered itonly a formof JV.alba.Carriere writes in 1888 of a JV. casparyi alba, as a new seedling from JV. casparyi;the description is peculiar; but if it refers to a white form, it can be no less than areturn to the typical color.Nymphaea odorata Ait. (Fig. 67.)Rhizome horizontal, with few stout persistent branches. Phyllotaxy about 2 on5, with long internodes (2 to 5 cm.). Petioles even greenish-purple. Leaves orbicular,usually purplish beneath, coriaceous; sinus open or closed. Flower very sweetI.-SP*
180 The Waterliues.scented, opening early (6 a. m.) sepals usually purplish-green or purple (pure green;in var. gigantea) Seed of medium size..Nymphaea odorata, Aiton 1789, fid. original specimen in hb. British Museum, from Kew Garden(excluding Siberian plant); 1811. Willdenow 1797; 1816. Sims 1805b. Andrews tab. 297.Pursh 1814. Nuttall 1818. Smith 1819. DeCandolle 1821 *; 1824. Rafinesque 1830. Planchon1853 b. Loudon 1855. Caspary 185s ; 1865. Meehan 1880. Wood 1880. Gray 1889. Gray,etc., 1895. Tricker 1897. Conard 1901 a.N. alba, Linnaeus 1753 (the North American plant). Walter 1788. Michaux 1803, M. specimenfrom hb. Michx., in hb. Paris.N. alba canadensis, Graham in Jameson Edinb. rev. str. 1. journ., vol. 8, p. 386, 1826 (not seen).N. alba /lore plena odorata, Gronovius 1762.Castalia pudica, Salisbury 1806 a. Britten 1888 a.C. odorata, Woodville & Wood 1806. Greene 1888. Branner & Coville 1888. Lawson 1889. Britton1889 a. Britton, etc., 1894. Britton & Brown 1897.Description. Flowers usually floating, 7 to 12 or 15 cm. across, opening on 3or 4 successive days from 6 a. m. to 12 m., with a strong, sweet, characteristic odor.Buds of two forms, the one slender, conical, rounded at apex, the other ovoid.Peduncle terete, greenish-purple, usually slender and weak (0.3 to 0.5 cm. in diameter) ;in the case of strong crowded plants in shallow water (30 cm. or less), the pedunclesmay be rigid and bear the flowers 15 cm. above the water (Sugar Hollow Pond, Danbury,Conn.). In fruit the peduncle coils into a close spiral (or loose if peduncle isvery stout), bent to right or left, or both ways in one peduncle; coils 5 to 9, about 2.5cm. in diameter. Air-canals as in Sec. Castalia. Receptacle short, round, scarcely evident,horizontal. Sepals 4, ovate to lance-ovate, rounded at apex, narrowed at base,putrescent after flowering. Outside green, more or less shaded with reddish-brown orred, covered margins nearly white; veinless; inside greenish-white. Petals 23 to 32(average 27), smaller than the sepals; outer ones ovate to elliptic-lanceolate, slightlyconcave, apex obtuse or rounded, narrowed at base, pure white, spreading out as muchas 1 5 below the horizontal. Median petal slightly obovate, obtuse at apex, wedgeshapedat base. Inner petal broadly ob-lanceolate, obtuse, sides curved and tapering atbase. Outermost staminode ob-lanceolate, acute, with one short anther cell. Stamens55 to 106 (average 79) inserted in direct series with the petals; innermost ripen first,outermost last. Filaments of all dimensions from staminodes to filiform threads, theinnermost being more slender than their anthers, which are bent inward at rightangles. Anthers curved, concave outward, placed obliquely so that the apex standsover the point of attachment or nearer the center of the flower; anthers largest inintermediate stamens. Pollen grains spherical, thickly beset with longer and shorterrods and lumps, alike from all the stamens, rupturing in germination by a large circularcap which isrough like the other parts of the grain. Carpels 13 to 25 (average17), with short linear styles, 0.64 cm. long by 0.16 cm. wide, which are erect on thefirst day of opening, later strongly incurved over the stigmatic disc. The latter isbasin-shaped, shallow, but of varying depth. Stigmatic surface continued out on thestyles one-half of their length or less, then narrowed suddenly and finally endingacuminately. Axile process nearly spherical, pointed at top, much narrowed at pointof attachment. Ovules many, small. Seed ovoid, of medium size, 0.23 cm. long by0.16 cm. in diameter, dark olive color, smooth, shining; aril one-fourth longer than theseed.
Taxonomy Nymphaea odorata. 181The seeds of N. odorata may germinate immediately after escaping from thefruit, or may remain dormant in cold water until the following spring; the first isprobably the case with fruits ripened before September I, the second with thoseFio. &1.Nymphaea odryrata: Under side of leaf on left; upper surface on right; a, sepal; b-d, outer,median, and inner petals; e, staminode; /.outer, median, and inner stamens; g, section of ovary at close offlowering period ; 8, astomatic area. Natural size of leaf 20.9 cm. X 28.6 cm. ; all reduced proportionately.ripened later. In both cases probably the first floating leaves are thrown up in thesummer after the seed was formed. I found great numbers of seedlings (var. minor)in a pond south of Marlton, N. J., August 30, 1900, with only submerged leaves. Thefirst leaf is filiform, about 1.6 cm. long. The second is elliptic-oblong. Intermediate
1 82 The Waterlilies.leaves are ovate, cordate, rounded at apex, with rounded basal lobes and sinus. Latesubmerged leaves are sagittate-cordate, with deep narrow sinus; are stiff in texture,bright emerald green, and stand erect or semi-erect, 1.12 to 1.42 cm. long by 0.8 to 1.0cm. wide.Leaf of mature plant nearly orbicular, fissi-cordate, never peltate, entire orslightly emarginate at apex, 12 to 25 cm. in diameter, smooth and dark green above,veins not evident; under side pustulate or spongy in appearance, smooth or slightlypubescent (?), varying in color from dark dull purplish-red to green tinged withred in the lateral regions. Veins evident but not prominent beneath, the wholelamina being thickened about them. Primary veins 8 to 9 on each side of leaf.Length of principal area: length of radius of leaf =1:1.12 to 1.2 (=7.6 cm. : 8.6cm.; =6.4:7.6 cm.). Depth of sinus less than one-half the length of the leaf( 1 : 2.5, average) margins straight or gently curved, touching, or spreading as much;as 30 ; angles obtuse, not at all produced. Petiole terete, slender, smooth or slightlypubescent (always pubescent or hirsute when young, and at base in maturity), 30 to180 cm. long, reddish-green to dark purplish-red, with 4 main air-canals and 2 to 4smaller ones. Idioblasts very numerous, about 150 to 200 in central quadrangle onecentimeter below leaf; many bipolar idioblasts in the smaller intercellular spaces, anda ring of thickened fibers around periphery. Stipules fused into a broadly wedgeshaped,thin, white and semi-transparent scale, closely appressed to the rhizome, withtwo-cleft apex and slightly decurrent, oblique base; outer surface pubescent, smoothnext to the rhizome; length 1.3 to 2 cm., width 1.2 to 14. cm.; bearing a shallowrounded furrow next to the petiole, with 2 longitudinal sub-median keels, 0.3 cm. apartbeneath.Rhizome stout, 2.25 to 3.2 cm. thick and 30 to 100 cm. long, pale in color, coveredwith a dense, short, black pubescence. Hairs of two types, one larger and stouter, ofcomparatively short, iso-diametric cells, the others much more numerous, slender,composed of few elongated cells ;all with 2 or 3 discoid basal cells. Interior dull whitewith a pinkish tinge. Apex rounded, clothed with stipules and long fine hairs. Branchesstout, few in number, their attachment 1.3 to 2 cm. in diameter.Geographic Distribution. Common in the Eastern United States; north toNova Scotia and Manitoba, south to Virginia and Arkansas.Notes. Introduced into Europe (England) in 1786, by Wm. Hamilton. In1801 it was " far less common " there than N. caerulea, and was considered a tenderplant; was well known in Europe in 1850; was cultivated in McMahon's gardenin Philadelphia in 1830 (Harshberger 1899), and appearsin American trade-lists in1883 and probably earlier.Flower from June to October ; fruit July to October. The roots were found tobe uninjured by drying away of the water in the fall at State College, Pa. (Buckhout1891) ;the plant was also seen blooming well in the absence of water at Baury'sLay, Buzzards Bay, R. I.(Sturtevant 1889). The plants which I have regarded astypes grow at Bristol, Pa., in a mill-pond.
Taxonomy Nymphaea odorata. 183Number of Floral Organs.Organs.
1 84 The Waterlilies.Rhizome 1.3 to 1.9 cm. inlong by 1.3 cm. wide, bifid, otherwise like those of the type.diameter, densely covered with long black hairs ; yellowish in cross-section.Measurements.Diameter of flowers 8.9 cm. ; 8.3 cm. ; 7.9 cm. ; 6.7 cm. ; 5.7 cm.Carpellary styles 0.4 cm. long, 0.08 cm. wide, continuation of stigma 0.13 cm. long.Geographic Distribution. Shallow water in cold bogs and sandy soil, occurringwith the type, but chiefly on the Atlantic Slope. The commonest form in New Jersey.FiO. 68. Nymphaea odorata minor: Upper row of figures from one flower of the narrowform ; 8, sepal ; p 1, 2, 3, 4, successive petals ; 8t 1-7, successive stamens ; c, carpellarystyle, with stigmatic surface dotted ; 82, b2, sepal and bud of the broader form ; 1, bl,sepal and bud of the narrow form. */ 2, staminode ; 2 at 2, outer stamen ;
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 16"VNYMPH/EA LOTUS.Collected by Schweinfatth at Damiuttn, Egypt.HEUOTYPE CO., BOSTON.
Taxonomy Nympmaea odorata. 185and had stomata on the back near the apical region. Such leaves were 5 to 10 cm.in diameter, mostly or entirely green beneath. The petioles were green or purplishgreenand had two large upper air-canals and two smaller lower ones, with no othersvisible to the unaided eye.The thickened idioblasts were very numerous, probably givingthe necessary stiffness to the petiole.
1 86 The Waterlilies.Plants from Raquette Lake, N. Y., differ a little from the New Jersey form ;theleaves tend more to an ovate shape, with overlapping sinus and slightly produced lobes ;in 30 to 180 cm. of water. The flowers have a slightly larger number of organs.Introduced into England by Pursh as N. odorata rosea in 1812.Var. rosea Pursh.Flowers of medium size, 8 to 10 cm. across, deep pink, or red; when closed,after once blooming, the sepals remain spread out horizontal on the water. Leaves12 cm. in diameter, deep crimson beneath, dark purplish-green above; primary veinsabout 7 on each side; length of principal area: radius of leaf = i: 1.2 to 1.26; sinusopen, angles somewhat produced.Nymphaea odorata var. rosea, Pursh 1814, fid. specimen coll. Pursh 1808, in Bass and WadingRivers, in hb. Kew. Garden 1883 b. Tricker 1897. Conard 1901 a.N. odorata minor fioribus roseis, Hooker 1883, marked N. odorata var. rubra on the typeKew; raised in Kew Garden from plants sent by A. Gray.N. rosea, Rafinesque 1830. Donn 1831.N. sanguinea, Donn 1831 ?Castalia odorata forma rosea, Britton 1889 a; 1890. Millspaugh 1893.C. odorata rosea, Britton, etc., 1894. Britton & Brown 1897.N. odorata rubra, Rev. Hortic. 1881 b. Wittmack 111. Gartenztg., 1882, p. 367.in hb.Geographic Distribution. Cape Cod; Barnstable, Mass. (Gray 1889); Lancaster,Mass. Cape May County, N. J. ; Woodstown, Salem County, N. J. ;Manchester,Ocean County, N. J. (Britton 18890). Buffalo, Putnam County, W. Va.(Millspaugh 1893). Rare and local.Notes. Millspaugh gives the following remarkable account of the Buffalo station" : On a plat of low bottom land near Buffalo, Putnam County, W. Va., the ploughturns up a large number of small tubers each season that the soil is cultivated. Theseplanted in tubs produce, much to the astonishment of the neighborhood, beautiful deeppinkwaterlilies. How long this bottom has been drained is not known, but the evidenceadduced by the fact above stated of the existence of a pond here, certainly over acentury ago, is very interesting, as is also the light thrown uponthe distribution ofthis pretty form of waterlily."The plants in cultivation are said to have sprung from a single original specimen.Var. gigantea Tricker. (Fig. 69.)Distinguished by large size in all its parts.Flowers less odorous than in the type;sepals mostly green. Leaves turned up around apical part of margin.N. odorata var. gigantea, Tricker 1897. Conard 1901 a.N. reniformis, Walter 1788? DeCandolle 1821 b. Donn 1831. Not Gray 1889.Castalia reniformis, Nash 1895, fid. original specimens.C. tuberosa, Lloyd & Tracy 1901 ? MacCarthy 1885 ?N. odorata var. chlorhisa, Rafinesque 1830?N. spiralis, Rafinesque 1830 (excluding synonyms).N. Maximiliani, Lehmann 18530, as to the flower; leaf is of N. tenerinervia ; fid. specimen from hb.Lehmann in hb. Berlin.N. Parkeriana, Lehmann 1853 a; 1854. Watson 1894. Conard 1901 a. Fid. specimens coll. Parkerin British Guiana, from hb. Lehmann in hbb. Kew, Berlin.AT. Pringlei, Rose MS., coll. Pringle, Plantae Mexicans: 1896, No. 6464, from Lake Xochimilcho= A r .tussilagifolia, Pringle 1897?
Taxonomy Nymphaea odorata. 187Description. Flowers floating, 10 to 15 cm. in diameter, closed before 2 p. m. ;odor dull or absent. Peduncles terete, rather stout, 8 to 10 mm. in diameter, in fruitshowing about 3 spiral turns 3.9 to 5 cm. in diameter. Sepals narrowly ovate, roundedat apex, breadth : = length 1 :2.4; outer surface green. Petals 24 to 31 (average of 13flowers,27.6), ovate or obovate, broadly rounded at apex, tapering at base, all alike inshape, pure white, outermost petals very faintly tinged with purple on the back near thebase ;shorter than the sepals ;breadth : length = 1 :2.9. Stamens 69 to 120 (averageof 9 flowers, 90.5). Carpels 15 to 23 (average of 9 flowers, 17.7) ; styles long, flat,thin, bending strongly inward after anthesis ; papillose ray ending bluntly far below thetip of the style. Fruit depressed globose, reddish around the bases of the styles whennearly ripe. Seeds as in the type, dark olive brown, smooth, 0.224 cm. long by 0.155cm. in diameter, inclosed in the aril.Mature leaves 40 cm.in diameter (15 to 60 cm.,Nash 1895), apical marginand part of sides turned upfor 2 to 5 cm., somewhat asin Victoria; upper surfacedark green; green beneath,with only a very faint tingeof purplish near the marginof some leaves (bright red,Tricker 1897). Veins, especiallythe midrib, prominent,9 on each side; principalarea : radius of leaf =1 : 1.2. Sinus open, marginscurved; angles obtuse,verylittle or not at allproduced. Petioles stout, plain green.Rhizomes stout, clothed with short dark hairs; yellowish-white inside; leavesscattered, about 2.5 cm. apart. Branches few and stoutly attached.Geographic Distribution. Southeastern United States ;Delaware to Floridaand Louisiana; Cuba; Mexico; British Guiana. E. g. C. Wright, Plantae Cubenses,No. 1856, coll. at Nueva Felipina, Cuba. R. M. Harper, No. 1212, Cane Water Pond,Decatur County, Ga. Nash, PI. Cent Penins. Florida, No. 1153, vicinity of Eustis.Curtiss, No. 6672, Lake Como, Fla.Notes. The plants which I have taken as types of this variety came from theDelaware and Chesapeake Canal at the railroad bridge, 2 miles west of St. George'sCity, Del. Here it grows in water from 60 to 120 cm. deep; in Lake Ella, Florida,itgrows from the shallows of the shore out to a depth of 3 to 5 meters (Nash 1895).If Walter's N. reniformis can be identified with any known plant, this must bethe one; but there is nothing but its geographical range by which to verify it, asno type exists in the Walter herbarium in hb. British Museum. His description isas follows :Fia. 09. Nymphaea odorata gigantea: , sepal; pi, p2, outer andmedian petals ; st 1, 2, 3, outer, median, and inner stamens. Natural size." N. reniformis, foliis reniformibus, corollis polypetalis,loculis mono-
1 88 The Waterlilies.spermis."If we accept the last two words as binding, the name cannot apply to anyspecies of the present genus Nymphaea, but rather to Nelumbo, as was understood byRafinesque (1830). It seems best, therefore, as Caspary already advised (1865), todiscard the name entirely. Gray was certainly wrong in applyingit to N. tuberosa,a species which does not exist in the region covered by Walter's Flora. In choosinga name, therefore, we have discarded the suggestion of Mr. Nash, and acceptedthe established and expressive name which the plant bears in gardens. Britton &Brown (1897) are certainly right in pronouncing this plant distinct from N. tuberosaPaine, as is shown by the absence of tubers and the small size of the seeds. Inthese respects, however, as well as in the great number of idioblasts in the petioles, andthe behavior in germination and thereafter, it is in agreement with N. odorata. Aninteresting parallelism exists in the European N. alba, of which gigantic varietiesoccur in the warm and sunny climate of Greece.
CARNEGIE INSTITUTION OF WASHINGTON WATERULIES, PLATE 17NYMPH/EA PU8ESCENS.MELIOTYPE CO., BOSTON
Taxonomy Nymphaea tuberosa. 189Nymphaea tuberosa Paine. (Figs. 70, 71, 72.)Rhizome horizontal, with many, often compound, slenderlyattached tuber-likebranches. Phyllotaxy 2 on 5, with long internodes. Petioles green, with longitudinalbrown stripes above. Leaves orbicular, pure green beneath; sinus usually narrow,angles slightly produced. Flowers scentless or nearly so, opening about 8 a. m. ;sepals pure green; petals obovate to oblanceolate, pure dazzling white, often deflcxed.Seed large.Nymphc -t tuberosa, Paine 1865, fid. specimen coll. Paine on S. shore of Lake Ontario, from hb. A.Gray, in hb. Kew. Hooker 1881. Garden 1882; 1807. Gray, Manual ed. 5. Caspary 1888.Sargent 1888. Tricker 1897. Conard 1901 a.N. reniformis, Gray 1889. Not Walter 1788, etc.N. maculata, Rafinesque 1830.N. alba, Nuttall 1818. Not Linn., etc.N. blanda, Hort. (in some American gardens). Tricker 1897. Not Meyer 1818.N. tuberosa maxima, Hort. Conard 1901 a.N. tuberosa var. parva, Abbott 1888.Leuconymphaea reniformis, MacMillan 1892.Castalia reniformis, Branner & Coville 1888.C. tuberosa, Greene 1888. Britton 1889 a. Lawson 1889. Hitchcock 1890. Britton & Brown 1897.Fia. 70. Nymphaea tuberosa: s, sepal; pi, p2, p3, successive petals; 8t 1-4, successivestamens. Natural size.Description. Flower 10 to 23 cm. across, floating, or when crowded and inshallow water raisecT 10 to 15 cm. above the surface. Open 3 or 4 days (2 or 3,Sargent 1888) from 8 a. m. to 1 p. m. (2 or 3 p. m., Sargent, 1. c.) odorless; (or with afaint odor of apples, Gray 1889). Peduncle terete, stout, 0.5 to 0.9 cm. in diameter,green, forming in fruit, in the lower portion, about 3 close spirals 6 cm. in diameter, theupper portion only slightly bent ; 30 cm. to 2 meters long ; air-canals as in N. odorata.Sepals 4, often becoming reflexed to the peduncle in flower, putrescent ; pure greenoutside, rounded at apex. Petals all pure white, obovate or almost spatulate, concave,rounded at apex, wedge-shaped at base, outer ones reflexed in flower, gradually smallerinward but retaining the spatulate or obovate shape. Stamens as in N. odorata.Carpels about 14; axile process globular. Fruit depressed globose, bare of all floralparts, 3.2 cm. in diameter by 2.2 cm. high. Seeds few (157 in a fruit), large, darkolive-brown, smooth, with very prominent raphe ; 0.44 cm. long by 0.28 cm. in diameter.Aril about as long as the seed, or shorter, not stipitate.
190The Waterlilies.The germinating tuber produces at first 3 or 4 tliin and fragile submerged leaves.The first is broadly deltoid, rounded or emarginate at apex, truncate at base, 2.9 to3.8 cm. wide at base by 1.6 to 1.9 cm. long on a slender petiole 7 to 12 cm. long. Thesecond is similar to the first or may be much larger, somewhat drawn in at the petiole,approaching a hastate shape, 6.4 cm. wide by 3.8 cm. long. Third and fourthleaves hastate, rounded at apex, 5.4 cm. wide at base by 6.4 cm. long, the sinus 1.9cm. deep. The first floating leaf is orbicular-ovate, entire, with rounded apex, open,straight narrow sinus with lobes produced and acuminate, 5.7 to 7.6 cm. in diameter.By degrees the mature foliage is attained.Leaf of mature plant floating, or sometimes borne 2 to 7 cm. above the water,cleft to the petiole, less leathery in texture than in N. odorata, orbicular, 12 to 38 cm.Fig. 71. Nymphaea tuberota: (a) group of tubers; 1, S, 3, first, second, and third submerged leaves fromsprouting tuber. Natural size.across, entire, or slightly curved outward at apex, smooth, bright greenand somewhatveiny above (purplish and shiny when very young), darker in color over thepoint of insertion of the petiole. Under surface mostly smooth, pitted toward thecenter (sometimes pubescent, Britton & Brown, 1. c), always green at maturity,slightly purplish toward margins when young; veins rather prominent, 7 to 13 on eachside; length of principal area: radius of leaf =1:1.6. Sinus depth: length of leaf =1 :2.3, open or closed, margins gently curved, touching at proximal end, 3.2 to 6.4 cm.wide at periphery of leaf, angles short-acuminate, abrupt on outer side, obtuse atpoint. Petiole pubescent when very young, smooth at maturity, terete or a little flattenedventrally, with 1 or 2 longitudinal ridges, 30 cm. to 2 m. long, 0.6 to 0.9 cm.in diameter; 4 main air-canals surrounded by 12 smaller ones; idioblasts in verysmall number, about 40 in central quadrangle 1 cm. below leaf. Color green, with12 to 24 longitudinal brown stripes due to pigment dissolved in subepidermal tissue,running from near the leaf downward 1 or 2 dm. or even the whole length of thepetiole ;about one-half of the stripes are broader and more prominent than the rest.
Taxonomy Nymphaea tuberosa. 191Stipules fused, whitish, appressed, broadly triangular, slightly emarginate, with aswelling near the base, 1.4 cm. long by 1.4 cm. wide.Rhizome horizontal, stout, 2.5 to 5 cm. in diameter, 10 cm. to 90 cm. long; colorpale, almost obscured by fine dark hairs ; apex protected by stipules and long fine hairs.Phyllotaxy 2 on 5, becoming less evident on older parts; leaves 2 to 2.5 cm. apart.Roots 8 or more from one leafbase.Branches numerous, 2.5to 7 or 10 cm. apart ; originatingwith strong connection (1.12 cm.in diameter) with parent rhizome,but the attachment laterbecomes very slender (0.3 to 0.8cm.) so that the pieces easilybreak off, forming the so-calledtubers. Mature tubers 2.2 to 6or 8 cm. long by 1.3 to 1.9 cm.thick, occurring singly or ingroups of 2 to 5 from an originallysingle shoot. They areloaded with starch, and show theusual rhizome structure; theyspring from any point withoutany visible order on the parentstem; the connecting isthmusshows (in transverse section) 3vascular bundles with centralxylem, embedded in a mass oflarge-celled, loose, starch-ladenparenchyma.Geographic Distribution.North and west of the AppalachianMountains in NorthAmerica : Lake Champlain, westthrough the Great Lakes toMinnesota, south to Arkansas.Very rare on the Atlantic Slope.Lake Hopatcong, New Jersey,fid. specimens cultivated atRiverton.Fio. 72. Nymphaea tuberosa. Under side of leaf on right,upper side on left ; 8, automatic area ; o, petiole, showing stripes ;b, stipule. Natural size of leaf 24.5 cm. long; stipule reducedproportionately. Petiole enlarged.Poeatquissing Creek,Mercer County, N. J. (Abbott 1888, and our own collections). Meadville, Pa. (Britton& Brown 1897). Raquette River, New York (Prentiss in Bull. Torr. Bot. C\., 10: 43-45). Ontario, Canada (Britton 1889 b). Grosse Isle, mouth of Detroit River (Campbell,Bull. Torr. Bot. CI., 13: 93-94, 1886). Southeastern Minnesota (Wheeler 1900). Ames,Iowa (Hitchcock 1890). Swamps near Little Rock, Ark. (Branner & Coville 1888).Notes. A smaller variety from Lake Hopatcong, N. J., is known in gardens
192 The Waterlii.ies.as N. odorata maxima (N. tuberosa maxima (Hort.) Conard 1901 a) ;it has leaves 7 to15 cm. in diameter; margin of leaf a little drawn in at the sinus; lobes overlapping;angles acuminate. Petiole pubescent, with a few faint longitudinal stripes. Rhizomesomewhat tuberiferous. Flower medium to large, cup-shaped, with the fragrance ofN. odorata; sepals pure green outside, petals pure white like the type; rarely producesseed. This may be a natural hybrid with N. odorata, since that species is commonin New Jersey, and Dr. Abbott has reported a diminutive but otherwise typical formof N. tuberosa under the name of N. t. var. parva from near Trenton, N. J.A pink odorless waterlily found at Painesville, Ohio, may be a varietyof this(Beardslee 1877).Nytnphaea tuberosa is a very strong grower ; when one wades among the plants,great numbers of the tubers are detached, and float to the surface of the water; thusthe plants are distributed and will in short time take complete possession of a smallpond. Gray (1889) speaks of the tubers as " spontaneously detaching "; we have seenno way for this to .occur; it seems more likely that they are naturally broken off by wadinganimals (deer, etc.) which feed on the leaves of Nymphaeas, as also by frogs,turtles, and even by fish.At the margin of the astomatic area on the leaves, the stomata are large, oval orround, and scattered, immersed in shallow pits; those of the rest of the lamina aresmall, round and crowded. Stomata are absent from a narrow marginleaf and along the sinus.all round theFlower, June to September. Fruit, July to October.According to Robertson (1889) the flowers are visited by Hymenoptera: Andrenidae:(1) Agapostemon radiatus Say ?; (2) A. nigricans F. 9; (3, 4) Halictusspp. ?; (5) H. pectoralis Sm. 9; (6) H. occidentalis Cress. ?; (7) H. coriaceus Sm. 2;(8) Prosopis sp. ?. Diptera: Syrphidae: (9) Helophilus divisus Lw. (10) H. latifronsLw. Bombyliidae:( 1 1 ) Sparnopolius fulvus Wied. Coleoptera Rhipiphoridae : :(12) Rhipiphorus limbatus F.Subgenus 4. LOTOS, DeCandolle 1821 (in part).Flowers nocturnal, raised on stout peduncles 10 to 30 cm. above the water.Sepals with 7 to 13 conspicuous and prominent veins. Stamens inserted at some distanceabove the petals, all flat and more or less ligulate, rounded at apex, dehiscingsimultaneously. Pollen smooth. Styles linear, ligulate. Leaves sinuate-dentate withacute sub-spinous teeth. Petioles, peduncles, and under side of leaves more or lesspubescent. Main air-canals in peduncles 6 (5 to 7), in petioles 2; idioblasts and multicellularingrowths absent. Rhizome tuberous, erect, ovoid, drying off in the restingseason. Seed rather small, with interrupted longitudinal lines of short hairs. Fourspecies native to the tropics of the Old World, from Senegambia to the PhilippineIslands, extending north to Hungary (two isolated localities) and south to Madagascar.Lotos, Planchon 18536. Caspary 1865; 1888.Sec. II. Inappendiculatae trib. I Lotos (in part) and trib. II Chamaelotos (in part), Lehmann18530.
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Taxonomy Lotos. 193Notes. The following species differ chiefly in color of leaf and flower, in shapeof leaf, and in production of seed. It is stated in Hooker & Thomson (1855) thatallgradations in color occur in India from the white to the deep-red forms; theytherefore united them into one species, N. lotus L., with three varieties : a. Lotus; leavesorbicular or reniform, densely pubescent beneath ; lobes divergent or approximate ;flowers ample, red, rosy or white. /9. cordifolia; leaves cordate-ovate, densely pubescentbeneath ;lobes divergent flowers;of medium size, white or carneous. y. pubescens;leave, puberulent or pubescent beneath ;flowers smaller, white, rosy or red.These characters are wholly unreliable in cultivation, whereas the color relations comepretty true to seed, and are usually reproduced precisely from the tuber. Caspary's(1865) division of the species into var. lata with sepals ovate to ovate-oblong, petalswhite or rosy, filaments yellow; var. angusta, with sepals mostly oblong-lanceolate,petals white, inner filaments with a purple spot; var. oblonga, with sepals mostlyoblong, rarely oblong-lanceolate,petals white, rosy or purple,is also unsatisfactory.There seems to be no constantmorphological differenceon which to base a classification.In 1888, however,Caspary speaks of N. lotus,pubescens, and rubra as distinctspecies. This opinionhave been founded onmaythe relations of hybrids, ofwhich many are in cultivation.But the evidence from thissource is not conclusive. Forexample, the hybrid N. omaranahas bronzy green leavesand large flowers with petalspinkish-red with a nearly white streak up the middle, and the floral leaves openout horizontally ; it shows plainly the blending of its N. lotus and N. rubra parentage ;it bears seed, however, much more freely than N. rubra, but less freelythan N. lotus.Its exact parentage is stated to be N. lotus X N. sturtevantii; the latter is a seedlingfrom N. devoniensis, raised without crossing, according to Mr. Sturtevant, but itgives evidence of N. lotus parentage. N. devoniensis was at first considered to be ahybrid of N. lotus and N. rubra, but it shows no evidence of white parentage, andis probably not of hybrid origin, and Hooker & Thomson claim to have found itcommon in India ; itvery rarely produces seed, however, a character which it seemsto have inherited in exaggerated form from its red parent. Whatever else may beinferred from these facts, certain it is that crossingof these forms does not inducesterility, but rather that fertility increases the nearer the mixed progeny approaches tothe highly fertile N. lotus. From what we have seen, however, there seems sufficientreason to consider N. lotus, pubescens, and rubra quite distinct.15Fig. 73. Floral organs of the Lotos group. Outer, median and innerstamens; carpellary style; vertical section of ovary. Natural size.From a white flowered garden form.
194 The Waterlilies.Nymphaea lotus (Linnaeus) Willdenow. (Plate XVI.)Flowers usually large, white or tinged pink on backs of outer petals; stamensyellow. Fruits numerous, depressed-spherical, truncate above, yellowish-green. Seedsbrown, shining, with very fine meridional ridges.Nymphaea lotus, Linnaeus 1753 (excluding Indian and American plants), fid. original specimen inhb. Linnaeus, now at Linn. Soc. London. Hooker & Thomson 1855 (in part). Miquel1859 (in part). Caspary 1865 (in part). Boissier 1867 (in part).N. lotus, Willdenow 1797. Savigny 1802. Waldstein & Kitaibel 1802. Andrews tab. 391. Sims18050. Aiton 1811. Delile 1813. Sprengel 1817. DeCandolle 182 1 b; 1824. Guillemin &Perrottet 1830. Planchon 1852 b. Lehmann 1853 a. Simkovics 1883. Caspary 1888. Tricker1897. Moenkemyer 1897. Conard 1901 a.N. lotus a Aegyptia, Planchon 1853 b.N. aegyptiaca, Opiz Naturalientausch (fid. Steudel 1841 ; Casp. 1865.)N. thermalis, DeCandolle 1821 b; 1824, fid. specimen coll. Kitaibel at Grosswardein, 1815, in hb.DeCandolle." Reichenbach 1845. Lehmann 1853 a. Planchon 1851 c; 1853 b.N. dentata, Schumacher & Thonning 1829, fid. specimens "ded. Schum." from hb. Lehmann in hb.Berlin; no types of this exist at Copenhagen, fid. Prof. E. Warming. Walpers 1842.Hooker 1846. Planchon 18500. Rev. Horticole 1851 a. Loudon 1855. Tricker 1897.N. Ortgiesiana, Planchon 1852 e, fid. specimen sent by J. E. Planchon Feb. 4, 1853, in hb. Kew.Castalia mystica, Salisbury 18066.C. lotus, Woodville & Wood 1806. Tratinnick 1822.Description. Flowers 15 to 25 cm. across, open on 4 successive nights from7.30 p. m. to 11 a. m., the sepals and petals all standing out horizontally, and stamensall erect; nearly odorless. Buds ovate, obtuse. Peduncle stout, pubescent, 0.6 to 2.0cm. in diameter, with usually 6 main air-canals, surrounded by 12 smaller ones; nospiral bending in fruit. Receptacle conical, about twice as wide as upper part ofpeduncle. Sepals 4, broadly ovate (10.8 cm. long by 5.1 cm. wide), rounded atapex, pure green on back, white at base, with 10 to 16 creamy white, prominent veins ;covered margins and inside white. Petals 19 or 20, oval, broadly rounded at apex,tapering slightly at base, longer than the sepals; texture rather thick and firm; 4outermost petals green along middle of back, otherwise white, or tinged with pink orpurplish; inner petals shorter and narrower, never stamenodial. Stamens about halfas long as the petals, yellow ;outer ones with broad, persistent filament having a lunate,semi-decurrent insertion on the torus ;inner filaments slightly wider than the anthers,with or without a dark purplish-brown spot on upper part ;outer anthers shorter thantheir filaments ;inner anthers as long as, or slightly longer than the filaments. Ovaryfinely pubescent. Carpels about 30 ; styles yellow, tinged purplish-brown on the back,becoming incurved over the stigma in fruit ; stigma funnel-shaped ; axile process short,cylindric, rounded. Fruit large, 6 to 9 cm. in diameter, with copious seeds; stigmafunnel-shaped; calyx and enlarged outer filaments persistent. Seed 1.36 mm. by 1.07mm. to 1.44 mm. by 1.09 mm., with interrupted longitudinal lines of hairs ;aril inclosingthe seed.Germination may occur immediately after ripening of the seed, or better after aperiod of drought. First leaf filiform, 1.6 to 1.9 cm. long. Second leaf linear tolinear-ovate, rounded at apex, tapering at base, 1.9 to 2.2 cm. long by 0.48 cm. wide;petiole slender, about 1.3 cm. long. Third leaf narrowly ovate with abruptly tapering
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Taxonomy Nymphaea lotus. 195base, 1.6 cm. long by 0.64 cm. wide. Fourth leaf ovate, 1.6 cm. long by 0.95 cm.wide, broadest and nearly truncate at base, the sides nearly parallel below, curvingin above the middle to the rounded apex. Fifth leaf cordate-ovate, with rounded apexand lobes, 1.68 cm. long by 1.04 cm. wide, sinus 0.16 cm. deep (cf. Lubbock 1892).The germinating tuber gives rise, all over its apical region, to a large numberof slender stolons about 2.5 cm. long and 0.08 to 0.16 cm. in diameter, with a lanceolate,acute, translucent bract 0.6 to 1.2 cm. long on the basal portion, and an apicalcenter of vegetation, from which about three submerged leaves are first formed, varyinggreatly in size according to the supply of nourishment. The first leaf may havea petiole 3.8 cm. long, bearing a filiform midrib 1.3 cm. long with a minute protuberanceon either side at base representing the lamina ;or the lamina may be expandedas a linear terminal lobe with a shorter linear lobe on either side of the roundedt /*Fio. 74. Early leaves of the Lotos group, from (Terminating tuber of N. devoniemia Hort.: Is, laminaand stipules of first submerged leaf ; 2s, lamina, !, stipules, of second submerged leaf ; S/, 3/,second and third floating leaves. Laminse natural size. Stipules enlarged.base. The second submerged leaf is deltoid, with rounded angles and broad, shallow,rounded sinus. The third leaf is broadly deltoid, with deeper sinus. All three appearat first of a reddish color, later becoming green their texture is very soft and thin,;and margins entire. The first floating leaves vary much in size according to foodsupply,but are entire, oval, deeply cordate, with rounded apex and lobes ; length :breadth = 1.5: 1(about) ; gradually leaves of a more circular outline are produced,and when practically round the dentation appears, at first very faintly, slowly reachingthe adult form in the older leaves.Leaf of mature plant distinctly peltate (2 to 5 cm.), orbicular, 20 to 50 cm. indiameter, subspinose-dentate and more or less wavy at margin, the teeth being sharpand brown-tipped, situated on the crests of the vertical undulations of the margin,1.3 to 5.0 cm. apart, with curved sinuses. Texture very firm and stiff by reason ofthe large prominent veins of the under surface; primary veins about 9 on either side
ig8The Watermmes.sinuate-dentate ;thin and membranaceous ; younger ones glabrous,adult or sub-adultleaves glabrous above, densely pilose beneath with very short hairs; veins slightlyprominent above, manifestly so beneath; venation loose, conspicuous; flowers 5 to 7cm. in diameter, ' white ' ; sepals 4, ovate-lanceolate, acute, whitish within, light greenoutside; petals 7 to 8, smaller than the sepals, and becoming decidedly smaller andnarrower inward ;stamens 35 to 45, broadly linear ;anthers rather short, connectivescarcely produced; carpels several." Leaf 10 to 15 cm. long, 9 to 13 cm. wide; the sinus is about 2 to 4 cm. wideat base of lobes. The sepals are 2.5 to 3 cm. long, 7 to 10 mm. wide ;the petals decreasein size inward down to 2 cm. long and 4 mm. wide. The outermost stamens are about2 cm. long and 3 mm. wide at base, anthers 7 mm. long by 2 mm. wide." (E. Gilg.)N. Zenkeri, Gilg, in herbaria.Geographic Distribution." Cameroon :Bipindi, in slowly flowing brooks andin streams of the Lokundje at Bequanohe. Coll. Zenker, No. 2130. Flower in July."(E. Gilg.)Notes. Finding this species labeled by Professor Gilg in several European herbaria,but there being as yet no published description, we were favored bythe Berlinprofessor with the above description from his own pen, based on the specimens inthe Royal Herbarium. We have gladly inserted it therefore, translated verbatimfrom the Latin and German in which it was communicated.Nymphaea pubescens Willdenow.(Plate XVII.)Flowers small, white; stamens yellow. Leaves small, ovate, dark green. Fruitsnumerous, nearly spherical, contracted above, dark green. Seed spherical-ovoid,brownish-olive, shining, nearly smooth.Nymphaea pubescens, Willdenow 1797. Aiton 1811. Smith 1819. Wight & Arnott 1834. Planchon1853 b. Lehmann 1853 a. Caspary 1888.N. lotus, Linnaeus 1753 (in part), fid. specimen in hb. Hermann, now British Museum. Roxburgh1814; 1824. Hooker & Thomson 1855 (in part). Miquel 1859 (in part). Caspary 1865 (inpart). Kurz Veg. eil. Bangk. (fid. Caspary 1865). Boissier 1867 (in part). Blanco 1878?N. Coteka, Roxburgh 1806= Castalia edulis, Salisbury 18066.N. esculenta, Roxburgh 1814; 1824.N. edulis, DeCandolle 1821 b.N. sagittata, Edgeworth 1845, fid. specimen coll. M. P. Edgeworth, 1844, in N. W. India, in hb.Kew.N. semisterilis, Lehmann 1853 a (fid. Hooker & Thomson 1855, and specimen in hb. Kew).N. bella, Lehmann 1853 o as to the leaf, fid. specimen from hb. Lehmann in hb. Berlin. Flower is ofN. stellata Willd.N. Nouchati, Burman 1768, fid. specimen coll. Burman in India, in hb. Delessert = N. NaccheliJ. F. GmeL Syst 1591, fid. Hooker & Jackson 1893.Castalia sacra, Salisbury 1806 b.Description. Flowers small to medium size. Sepals 7 to 9 cm. long, obtuse;green outside, with white veins. Petals pure white outer ones; broadly obovate, width :length = 1 : 2.6. Outer stamens not lunate-decurrent at insertion, separated from thepetals by a wide gap. Fruit about 4 cm. in diameter. Seed 1.9 mm. long by 1.52 mm.in diameter.
CARNEGIE INSTITUTION OF WASHINGTON. WATERLILIES, PLATE 20.NYMPH/EARUDGEANA.1, 2, Views of the flower. 3, 4,- Pruits. 5, Bud. 6, Same as 5, with outer floral organs removed. 7-10, Petals. 11-15, stamens.16-19. Views and details of gyniecium. 20-22. Pollen. 23, Fruit. 24, Cross section of ovary. 25-27, Sections of stvle. 28, Verticalsection of stigma, showing parrfllte.
Taxonomy Nymphaea rubra. 199Leaves 25 cm. long or less ;breadth : length = 1 : 1.28. Sinus open or closed ;margin mostly curved. Upper surface of leaf dark green, under surface dull purplish-green,more or less pubescent. Length of pelta length of leaf = : 1 : 12.Tuber large, ovoid. (Other details as in N. lotus.)Geographic Distribution. British India to the Philippine Islands; south toJava and Australia.Notes. Of most of the plants classed above as synonyms descriptionsare notavailable. N. sagittate was separated on account of its open sinus and entire, sagittateleaves. ,these were doubtless from a young plant. It was collected at Ambala,province of Sirhind, India. N. semisterilis is said to have the leaves obtusely dentateor crenate. No one except the founders of these two species has ever considered themdistinct. The names edulis and esculenta were given in reference to the use of thetubers and seeds in India as food, as is the case with N. lotus inAfrica.TV. pubescens is said to have been introduced into England from India in 1803.Willdenow took as his type the plate entitled " Ambel " in Van Rheede's Flora Malabarica.The description and diagnosis given above are based on a plant grown by Mr.Tricker in the Dreer gardens at Riverton, N. J., in 1902, from seed sent by Mr.Gollan of Saharanpur, India, corroborated by comparison with herbarium material.Nymphaea rubra Roxburgh. (Plate XVIII.)Flowers deep purplish-red; stamens cinnabar-red, becoming brownish. Leaveslarge, dark bronzy red, becoming greenish in age. Fruits rare. Seed nearly spherical,dull brown, with fine, prominent, meridional ridges.Nymphaea rubra, Roxburgh 1806; 1814; 1824; fid. color drawings in hb. Kew. Andrews tab. 503.Sims 1810. Aiton 1811. Smith 1819. DeCandolle 18216; 1824. Wight & Arnott 1834.Wight 1850. Planchon 1850 b and d; 1852 a. Lehmann 1853 a. Lindley 1853 a.N. lotus, Hooker & Thomson 1855 (in part). Miquel 1859 (in part). Caspary 1865 (in part). Boissier1867 (in part). Not Linn. 1753.N. rubra var. rosea, Sims 181 1 b.N. rosea, Sweet 1827.N. Devoniensis, Hooker 18526, and Lindley 18536, fid. Hooker & Thorns. 1855. Garden 18830.Castalia magnifica, Salisbury 1806 b.C. lotus fiore rubro, Tratinnick 1822.Description. Flowers 15 to 25 cm. across, deep purplish-red, open from 8 p. m.to 11 a. m. Sepals dull purplish-red, about 7 nerved, never opening more than ioabove horizontal. Petals 12 to 20, narrowly oval, breadth :length = 1:3.15, roundedat apex, standing at various angles in the open flower from 45 to the level of thesepals. Stamens about 55, cinnabar-red, becoming brownish. Fruits very few, small.Seeds 1.85 mm. long by 1.6 mm. in diameter.Leaves large, 25 to 45 cm. across, dark reddish-brown, bronzy, becoming greenishin age; under surface pubescent. (Other details as in N. lotus.)Geographic Distribution. British India; Bengal.Notes. In gardens is distinguished N. rubra rosea, with leaves bronzy greenblotched with brown ;flowers large, magenta to dull red, open until 10.30 a. m. ; petalsnarrow, pointed; stamen tips orange-brown.
2ooThe Waterulies.We are doubtful if the true N. rubra exists in this country. Seed communicatedby Mr. Chandradatta of Calcutta has germinated but produced no floweringplants as yet. It was introduced into England in 1803.N. devoniensis at Lyon House, England, in 1885 produced a flower whosepeduncle arose from the axil of a stamen of a previous flower (Barber 1889), andDr. Kirk, according to Masters (1869), observed a similar abnormality in N. lotus inEast Africa. The sepals, petals, stamens and even carpels of these species may becomeleaf-like (Masters 1869).We have to admit that our arrangement of the Lotos group is far from beingsatisfactory or final. Important results would certainly follow a study of the plantsof various parts of Africa, Asia and the outlying islands. This can only be done effectivelyon living plants, either in cultivation or in their native haunts. The severalforms in gardens have proven useless on account of uncertainty of origin and probableadmixture of other stocks by crossing.Subgenus 5. HYDROCALLIS, Planchon 1852 b; 1853 o.Flowers nocturnal, floating. Sepals not evidently veined. Petals in whorls of 4,alternate with the sepals and with each other. Stamens inserted in series with thepetals, the outermost 4 or 8 more or less petaloid and in whorls like the petals anthers;dehiscing about simultaneously. Styles slender, cylindric, with enlarged club-shapedtips. Seeds small or minute, with numerous long hairs. Leaves entire, or dentatesinuatewith obtuse teeth. Rhizome erect, tuberous, drying off in the resting season.Ten known species, natives of the tropics of the Western Hemisphere.Hydrocallis, Caspary 1878; 1888.Castalia (in part), DC. 1821 b.Tribe Leucanthos (in part), Chamaclotus (in part), Lotos (in part) and Castalia (in part), Lehmann1853 a.Nymphaea amazonum, Martius & Zuccarini.(Plate XIX.)Number of floral parts 132 to 376. Sepals ovate-oblong, more or less acuminate.Petals 16 to 20, rows all about equidistant. Stamens 93 to 297. Pollen 0.0142to 0.0174 mm. in diameter. Carpels 19 to 40. Fruit with funnel-shaped stigmatic disc,the funnel surface curved ; sepals and outermost row of petals persistent. Seeds 10,000to 27,000, very small, 0.386 mm. wide by 0.706 mm. long to 0.566 mm. by 0.973 mm.,dull gray. Mature leaf entire, lobes subacuminate or rounded. Top of petiole bearinga ring of long hairs. Tuber producing stolons throughout its vegetative period.Flowers imperfectly opening two nights, becoming fully open suddenly the secondnight between 4 and 5 o'clock a. m. Bud ovate-oblong, acuminate.Nymphaea amazonum, Martius & Zuccarini 1832, fid. specimens coll. Martius, No. 3313, near Para,Brazil, in hbb. Munich, Berlin. Planchon 1853 b. Lehmann 1853 a. Hooker 1854. Caspary1878. Sagot 1881. Conard 1901 a.N. albo-viridis, St. Hilaire 1833, Ad. specimen coll. St. Hilaire, Prov. Rio de Janeiro, Brazil, in hb.Paris.JV. blanda, Macfadyen 1837. Hort. Glasnevin (.fid. Hooker, 1. a). Not Meyer 1818.JV. blanda /? Amazonum, Planchon 1856.JV. Rudgcana /? Amazonum, Grisebach 1864. Not N. rudgeana Mey. 1818.
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 21*/> I**\10 II.-^V. \NYMPH/tA BLANDA, N. BLANDA FENZLIANA, N. TENERINERVIA, N. STENASPIQOTA.A", hn'riv ,, hi. i, i- lower, 2. Lower nr&et of leaf. $, Ap-x ofouter stamen. 4, Apex of Innermosl stamen, 5, Carpellary style.6, Upper surface of k-.it'. 7, Ontei surface of sepal..v.ft IruH. m, ji, Outer and median stamens* 12. Leaf.X. hian flu.13, Flower rlewed from side. 14, j^eaf. i( Partof upper surface of snore Leaf, 16, Flower.HEUOTYPE CO., BOSTON.
Taxonomy Nymphaea amazonum. 201AT.N. integrifolia, Salzmann MS., Ad. specimens coll. Salzmann, 1830, at Bahia.N. foetida, Gardner MS., fid. specimen coll. Gardner, No. 2477, Aug., 1839, in the Li.ke of Paranagua,in hb. Berlin.nocturna, March MS. (Ad. Hooker, 1. c),Description. Flower floating, 10 to 12 cm. across, opening two nights. Thefirst night, the bud stands with its base about 1 cm. or more immersed in the water,and begins to open about 3 o'clock a. m. ; itspreads gradually until 4.15 a. m., whenthe sepals stand out at about 45 and the petals are loosened, leaving an aperture 1.3cm. wiJ-3 into the center of the flower. At 4.45 a. m. itbegins to close, being completelyclosed about 6 a. m. The second night the bud has risen till its base is justout of the water, and begins to loosen at the apex at 6.30 p. m. the; sepals spreadout rapidly to a horizontal position, followed by the outermost series of petals, whichreach nearly the horizontal position; the second series also open slightly from thetip of the bud (about 1 cm.) ;this phase is completed by 8 p. m., a tight creamywhiteacute bud being left in the center of the flower. The flower remains thus untilabout 3.30 a. m., when the parts begin to open out more fully, the process being completedabout 4.45 a. m. At this time the sepals and outermost petalsare horizontal inposition, the inner petals standing at all angles up to about 40 above the horizontal;the outermost stamens lie against the innermost petals; from this position thestamens stand at all angles up to the innermost which bend over the stigma. All ofthe stamens have now dehisced, the flower is warmer than the surrounding air, andhas a very faint sweetish odor with a suggestion of peaches. Closure of the flowerbegins about 5 a. m., and is led by the stamens and petals; these, with exception of1 or 2 outermost petals, are united into a tight bud at 5.30 a. m., while the sepals areonly io to 15 above the horizontal. By 6 o'clock complete closure is effected, andthe flower is retreating below the water, where it becomes completely covered beforethe middle of the succeeding afternoon, i. e., in eight hours or less. From 7 to 7.30p. m. and from 4 to 4.30 a. m. the movements of the petals and sepals on the secondnight is very rapid and may easily be seen; the apex of the organ rotates about itsbase, moving about 1.3 cm. in three minutes; the aperture at the apex of the budtherefore enlarges at twice this rate. In one observation the inmost petals were closedover the stamens at 4 a. m. at ; 4.06 their apices stood 5 cm. !apart The closure ofthe flower after its full maturity is almost as rapid. Bud 5.4 to 7.3 cm. long by 2.7 to3.8 cm. in diameter, conical-ovate, broad at base, acuminate above, more or less obtuse,abruptly contracted below to the petiole. Peduncle 40 to 50 cm. long, 0.6 to 0.8 cm.in diameter, terete, brownish-green, lighter at base, smooth, with 3 minute central aircanals,6 to 7 main canals and 12 to 14 outer canals, separated by extremely thin partitions.Receptacle round, pure dark brown, nearly horizontal, distinctly boundedfrom the sepals at the margins. Sepals 4, fleshy, ovate, rounded at apex, carinate onback in upper part, the ridge vanishing about two-thirds of the length from the tip.Outer surface dark green above, paler below, the lower quarter being greenish-white;whole surface marked with strong purplish-black lines and dots, the lines becomingInner surface apple green above, shadingbright red on the lower quarter of the sepal.to greenish-white at base, with a few dull brownish spots and streaks on margins inupper half. Innermost (posterior) sepal deeply hooded at apex, i. e., the fleshy tip is
202 The Waterlilies.excavated, forming a scoop-like depression about 0.3 cm. deep, which receives the tipof the bud and covers it above and on three sides; outermost (anterior) and rightsepal slightly hooded, left sepal very slightly hooded. Margins of innermost sepalcovered in the bud, about 0.16 cm. wide, whitish and semi-transparent. All veinless,persistent in fruit. Petals 16 to 20, of a yellowish-white color, rather thick and leathery,a little shorter than the sepals, deeply concave and slightly hooded at apex;when cut off from the receptacle (of a closed flower), the basal portion curls inwardstrongly, showing the tension to which it has been subjected in closure. Outer petalbroadly elliptic, obtuse, obscurely n-nerved; second and third series similar; fourthseries similar but broader in proportion at middle and base, and more rounded atapex; fifth series becoming acute at apex. Stamens 136 to 188, yellowish-white, withdirty yellow anthers, unappendaged. The lowest series of 8 have broad filaments witha fleshy midrib and parallel anther cells; filaments of second series about one-thirdas wide as the first, the inner ones becoming more slender and placed without visibleorder; innermost filaments narrower than their anthers. Carpels 25 to 35, styles clavate,about 1.3 cm. long, the enlarged tip 0.5 cm. long by nearly 0.3 cm. indiameter,pure red around the end, the extreme tip whitish ; they stand alternately in two positionsone is bent directly inward so that the tip touches the stigmatic disc two-thirdsof the way to its center; the one on either side of this is coiled spirally so that theapex lies beside and against its base, there being but one turn of the coil. Stigmaticdisc curved, scarcely depressed at center ; papillose area continued 0.3 cm. out on thestyles, where it ends obtusely. The stigmatic papillae break up at the flowering timeinto a moist powdery mass of cells lying singly or in chains, of a golden yellow color.Axile process 0.25 cm. high by 0.08 cm. in diameter, appearing as a slender acute rodprojecting half of its length from a narrow pit at the center of the flower. Ovaryspheroidal, 1.6 to 2.4 cm. in diameter, 1.12 to 1.42 cm. high. Ovules very numerous,very small. Fruit sub-spherical, with sepals and outer petals standing loosely roundit, about 3.9 cm. in diameter. Seeds very small and very numerous, 0.1 cm. long by0.074 cm. in diameter, reddish-brown in color, with a silvery appearance due to manyscattered long white hairs; raphe scarcely evident.Germination takes place as soon as seeds are shed. First leaf filiform, short(0.95 cm. long). Second leaf spatulate, rounded at apex, tapering at base, 0.8 cm.long by 0.32 cm. wide. Third leaf elliptic ovate.Leaf of mature plant orbicular-ovate, breadth : = length 1 : 1.2 to 1.18 (17.2 cm.by 14.3 cm), rounded at apex, peltate by about 0.3 cm., thin and soft in texture;margin entire, somewhat wavy up and down; upper surface green, yellowish towardthe center, with faint brownish spots. Under surface glabrous, brownish-red, withnumerous small spots of a blackish color. Veins all prominent, 7 to 9 on each side.Length of principal area: radius of leaf =1:1.4 to 1.47 (2.4 cm. :3.5 cm.). Sinusopen or closed, depth: length of leaf =1:2.3 to 2.6 (7 cm.: 17.2 cm.), margins approachingor overlapping, nearly straight above, becoming strongly curved outwardtoward periphery, forming two broad, round, more or less widely separated lobes.Astomatic area small ; stomata becoming larger and scattered near it, but stoppingrather suddenly. Petiole glabrous, except a ring of long simple hairs at its junctionwith the leaf; 45 to 60 cm. long, 0.9 cm. in diameter; with two transversely placed
Taxonomy Nymphaea amazonum. 203main air-canals, two smaller ones anteriorly and posteriorly, and two stillsmaller onesbeyond these ;one lateral canal on either side of main pair, and numerous smaller onesabout; main partitions 2 cells thick; one double bundle anteriorly and one posteriorlyand two equidistant from these and from each other on either side; numerous smallerbundles. Idioblasts numerous, all stellate, rather short armed.Rhizome (tuber) erect or semi-erect, thick, about 7 cm. long by 3.8 cm. indiameter, brownish-black, apex protected by leaf-bases and long hairs. Phyllotaxy ofa high order, the leaf-bases being very close together. Branches several (3 to 5 or6), 7 v 10 cm. long on short stolon-like stalks, produced throughout the season ofvegetation, and sending up leaves and flowers among those of the parent plant.Geographic Distribution. In still water ; lakes, ponds and ditches. TropicalAmerica between 18 N., 23 S., 38 W., and 78 W., in Jamaica, Guiana, Surinam,Brazil (Para, Bahia, Rio Janeiro). New Granada. Laguna del Corcho, near Ocuiltzapotlan,Mexico.
204 The Wateri.ilies.Measurements of Petals (incentimeters).Series.Lenvrth.Flowbr 1.Outermost.Second .Third.Fourth ,Fifth .5-865-35-15-35-24 954-954.84-74-34.6
MELIOTYPE CO., BOSTON.CARNEGIE INSTITUTION OF WASHINGTONWATERLILIES, PLATE 221 o^.V.NYMPH/EA LASIOPHYLLA AND N. RUDGEANA'i,2. Flower from face and back..5, Leaf, showing lower side. 4, Part of larger leaf from above, lohes touching.5,6, Outermost arid innermost stamens. 7, ("art of lower surface of leaf.ruthit/tint.plant from Jamaica. 9. 10. 11, Stipules of three leaves of the |amaica plant. 12, Kruit, raised at Konitfsbcrfy.13, Another fruit raised at Kdnlgsberg with scp:i]s and petals removed. 14, Konijrsnertf fruit in median section. 15, Small tlower,island fit Cotara. i'\ s< paj offl Bower rrora Brazil. 17. is. Stamens of same Ho-.ver. to. Apices of median and innermost stamens./
Taxonomy Nymphaea rudgeana. 205sistent. Seeds 1,000 to 8,000, short-elliptic, 1.15 mm. in diameter by 1.6 mm. long,to 1.3 mm. by 2.2 mm., shining, greenish-gray. Mature leaf sinuate-dentate. Petiolenaked at point of insertion. Tuber producing stolons only when first germinatingfrom the dry state. Flowers imperfectly open two or three nights, rarely entirelyopen before midnight. Buds ovate-oblong.Nymphaea Rudgeana, G. F. W. Meyer 1818. Grisebach 1864. Caspary 1878. Garden 18836.
2o6The Waterlilies.dentate with unequal, distant teeth ; primary veins 4 ;under side dull greenish, suffusedwith red-brown at apex upper surface bright green with a narrow red border ; ;marked above and beneath with radiating blue-black spots. Mature leaf of adult plant40 cm. long, elliptic to orbicular, fissi-cordate, more or less coriaceous ; margins coarselyand unequally sinuate-dentate, with obtuse teeth becoming obsolete at apex of leaf;lobes touching or distant ; margins of sinus irregularly sinuate-dentate. Primary veins= 7 to 11 ; length of principal area: length of radius 1 : 1.07 to 1.5. Upper surfacedull greenish, more or less suffused with red-brown, especially toward the margin;both surfaces marked with a greater or less number of blackish spots. Petiole terete,without ring of hairs at top; main air-canals 4, the two upper ones larger. Roots6 to 7 from each leaf-base.The caudex in the native country vegetates throughout the year and becomes verylong and sub-fusiform. Young plants cultivated in central Europe cease growing inwinter, forming ellipsoid tubers of the sire of a hen's egg or larger. Stolons fromtuber cylindric, with 1 to 4 triangular scale-leaves at base ;later leaves as in seedlingplant.Geographic Distribution. In still and slow-flowing waters; tropical Americabetween 18 N. and 22 S., 38 W. and 78 W. Jamaica; Martinique; Guiana; Surinam;Brazil (Caspary, 1. c). Porto Rico (Espinosa 1881). Type collected by Dr.Rodschied in the Essequibo River, British Guiana, in herb. Grisebach (not seen).Examples. Coll. Mosen, No. 3339, Prov. S. Paulo, Brazil; coll. Jenman, coastlands of British Guiana, Nov., 1889, No. 5748 ; coll. Poeppig, No. 2930 and 3033 ;coll.Burchell, No. 5556. Most of these are marked " N. blanda" in hbb. Kew, BritishMuseum, Delessert, &c.Notes. The above description is taken from Caspary's account in Flora Brasiliensis.He had the plant under cultivation, introduced by Dr. Moore from SpanishTown, Jamaica, in 1854. We have only seen young plants, in the garden of OakesAmes, North Easton, Mass., R. M. Grey, gardener. One of these specimens was sentto us by the kindness of Mr. Grey, but, the summer being cool, it did not flower.The seeds of this species are used for food by American natives.Nymphaea blanda G. F. W. Meyer. (Plate XXI.)Sepals densely marked with very slender crimson-brown lines, oblong-ovate,rather obtuse, sub-acuminate. Styles linear-clavate, about 8 mm. long. Peduncle coveredthroughout (or just below the flower) with septate hairs (glabrous in the var.).Leaves small, membranaceous, entire, cordate-cleft, suborbicular or orbicular-elliptic,apex very broadly rounded; lobes slightly produced, sub-acuminate and sub-hastate;pale green, very narrowly peltate (scarcely 1 mm.). Petiole villous at top, or hereand there, or throughout, or (in the var.) glabrous.Nymphaea blanda, G. F. W. Meyer 1818. Schomburgk 1848. Seeman 1852. Dalton 1853. Caspary1878. Hemsley 1888. Conard 1901 a.N. Rudgeana P Amazonum, Grisebach 1857. Not N. rudgeana Meyer 1818.N. Fenzliana, Lehmann 1853 a and b; 1854.N. glandulifera, Rodschied in Meyer 1818.. .
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 23 ,.(9IPNYMPH/EA JAMESONIANA.1. 1, Flower fr.. in front ml bsck ; t, pal. 3, 4, 5. Innermost, outer, and outermost stamen*. 6, 7. Leave*. 8, Part ofleaf from beneath.MELIOTYPE CO., BOSTON.
Taxonomy Nymphaea blanda. 207Description. Flower 81 to 95 mm. across. Sepals 35 to 44 mm. long by 13 to20.5 mm. wide, widest below the middle. Petals 16; outer ones 38 mm. long by 15mm. wide, oblong, ovate; outside sepaloid along the middle, margins and insideyellowish. Inner petals gradually smaller; innermost oblong-lanceolate, acute.Stamens about 65, outer 4 largest, 25 mm. long, with filaments petaloid, longer thanand twice as wide as anthers; inner stamens gradually smaller, the filament becomingshort, and narrower than the anther, innermost 14 mm. long; all the stamens areyellowish. Carpels 26 ; styles 9 to 1 1 mm. long, incurved and purplish-brown at apex.Peduncle clothed with septate hairs.Leaf small, entire, membranaceous, elliptic-orbicular, narrowly peltate, greenabove and below ;lobes distant, sub-hastate, sub-acute ; margins of sinus gently curved ;primary veins 6 on each side. Petiole clothed with long septate hairs in the upperpart or throughout.Tuber oblong or ovate, small, covered with brownish hairs.Geographic Distribution. Mouth of Essequibo River, Guiana, in ponds andstreams, collected by Rodschied about 1814. Swamps near city of Panama, coll. Seeman.Guatemala; Jamaica (Hemsley, 1. c).Notes. The description above is taken from Caspary (la), who studied Meyer'stype collected by Rodschied in hb. Grisebach, and Parker's collection in hb. Hooker &Lindley.Var. fenzliana (Lehmann) Caspary. (Plate XXI.)Peduncles and petioles glabrous.N. Fenzliana Lehmann, 1. c.N. blanda var. Fenzliana Caspary, 1. c, lid. types in hbb. Kew, Berlin.Description. Flower 96 mm. across. Sepals 40 to 42 mm. long by 12 to 16mm. wide, oblong, ovate, sub-acuminate, pale green outside, very densely marked withslender red-brown lines 3 to 4 mm. long; inside yellowish. Petals 16, pale yellowish;outermost 4 petals 38 to 40 mm. long by 13 to 15 mm. wide, acute; innermost shorterand narrower, sub-lanceolate. Outer stamens lanceolate, with anthers 8 mm. long,scarcely appendiculate. Carpels 23; styles about 8 mm. long,apex.1 mm. wide at clavateLeaf floating, coriaceo-membranaceous, entire, sub-orbicular, fissi-cordate, greenon both sides ; margins of sinus curved ;lobes slightly produced, sub-acuminate ; primaryveins 5 to 6 on each side, not prominent.Rhizome small, ovate-globose, 20 mm. long.Geographic Distribution. Collected in Guatemala near San Juan de Nicaraguain 1841.Notes.from which Caspary's description was drawn.Known only from the types in hb. Vienna and hb. Bentham (now Kew),Nymphaea lasiophylla Martius & Zuccarini. (Plate XXII.)Sepals with a few large black spots, ovate-oblong, sub-acute, sub-acuminate. Styles6 to 7 mm. long. Leaves large, entire, cordate to orbicular, cleft at base, broadly peltate(i. e., 8 to 16 mm.) ;lobes rather obtuse. Flowers 9.3 to 11.8 cm. in diameter.Petals 16. Stamens above 42. Carpels 15 to 20.
208 The Wateruues.Nymphaea lasiophylla, Martius & Zuccarini 1832. Planchon 1852 b (name only) ; 1853 b (name andlocality only). Caspary 1878.Description. Sepals 48 to 50 mm. long by 18 mm. wide. Petals white, oblong,sub-elliptic ;outermost 20 mm. wide by 50 mm. long ;innermost 4 lanceolate, 8 mm.wide by 42 mm. long;all acute. Stamens unequal; outermost petaloid, lanceolate;middle and inner ones linear, with anthers wider than the filaments ;outermost slightlyappendiculate. Styles 6 to 7 mm. long, 1 mm. wide at apex ; stigma yellowish ; axileprocess minute. Leaf coriaceous, sub-orbicular, entire; under side reddish, withbrown-purple lines; lobes distant or touching, more or less produced, obtuse; sinusmargins curved or nearly straight; primary veins 7 to 11. Petioles terete, smooth,reddish. (Pubescence of under side of leaf mentioned by Martius & Zuccarini is alayer of algae. Caspary, 1. c.)Geographic Distribution. Collected by Martius, No. 2377, near Joazeiro inLagodisso, province of Bahia, Brazil (9 40' S., 42 10' W.) in March and April. Otherwiseunknown. Type seen in lib. Munich.Nymphaea gardneriana Planchon.(Plate XXIV.)Sepals densely marked with long slender fuscous lines, oblong-ovate, distinctlyacute, acuminate. Petals 16 to 20, outermost oblong-ovate, innermost oblong-lanceolate.Stamens 46 to 62, outer filaments petaloid, inner filiform and short. Carpels 15to 18; rays of stigma rounded at apex; styles linear-clavate, very long (12 to 16 mm.),apex 1 to 1.25 mm. in diameter, dark purple; axile process minute. Leaf floating,6.5 cm. long, slightly coriaceous or sub-mem branaceous, peltate (pelta 5 to 8 mm. wide),ovate or orbicular-hastate, rarely orbicular, apex rounded or sub-retuse; sinus open,broadly triangular ; lobes acute or obtuse ; margin entire, often crisped ; primary nerves5 on each side (rarely 4) ; glabrous, irregularly marked above and beneath with darkviolet spots.Nymphaea Gardneriana, Planchon 1852 b; 1853 a and b; fid. specimens coll. Gardner, No. 2476, about1840, Paranagua, Brazil. Caspary 1878.JV*.fragrans, Gardner MS. in hb. Bentham, now Kew.N. Passiflora, Lehmann 18530 and b; 1854; fid. specimen marked in Lehmann's handwriting in hb.Kew.Description. Flower 92 to 115 mm. across. Sepals 11 to 19 mm. wide by 40to 55 mm. long ;outside nerveless, greenish, with long slender fuscous lines. Petalsacute, usually sub-acuminate; innermost oblong-lanceolate; outermost reddish-yellow,innermost reddish-brown (in dried specimens). Stamens reddish-brown; outermostpetaloid,hairs.innermost slender.Leaf small; veins slightly immersed beneath.Rhizome small, 16 mm. in diameter, sub-globose, covered with long gray-brownGeographic Distribution. Paranagua, Province Piauhy, Brazil (io 45' S.,484o'W.) ; Plain of Mbatolio, Paraguay, coll. Balansa, No. 521.
MEUOl'rPE CO., BOSTONCARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 24NYMPH/EA STENASPIDOTA AND N. GARDNERIANA.'>i, Whole plant ; , sepal. 2, Flower. 3, Partofleaf; upper surface.rt of outermost median and innermost stamens. 8, 9, 10, Leaves. 11, Upper surface of leaf.
Taxonomy Nymphaea stenaspidota. 209Nymphaea jamesoniana Planchon.(Plate XXIII.)Flower 87 to 107 mm. in diameter. Sepals green, densely marked from base toapex with long narrow dark crimson lines, 3 to 7 mm. long by 0.1 to 0.05 mm. wide;oblong-ovate, rather obtuse. Petals 16. Stamens 52 to 65. Carpels about 30; styles6 to 10 mm. long, clavate, purplish-brown at apex. Leaves sub-coriaceous, ovatecordate,cleft at base, sub-sagittate or elliptic, more or less peltate ; margins entire lobes;obtuse or acute ;under surface marked with dark purple forked and branching lines.Nymphaea Jamesoniana, Planchon 1852 b; 1853 a and b; fid. original specimen in hb. Delessert.Caspary 1878. Hemsley 1888.N. sagittariaefolia, Lehmann 1853 a and b; 1854 ;fid. specimen marked in Lehmann's handwritingin hb. Kew.Description. 5V/>a/.y 13 to 21 mm. wide by 40 to 46 mm. long. Petals 16,oblong those of the outer three whorls obtuse, of the inner whorl acute ; ;all brownishyellow(in dried specimens), sub-membranaceous. Stamens like petals in color andtexture ;outermost with an acute triangular appendage, innermost obtuse ;outermostfilaments four times as wide as anthers, innermost filaments and anthers about equalin width.Leaf thin and membranaceous; lobes approximate or distant; primary veins 6 oneach side, very slender, slightly immersed at base, elsewhere level ; principal area halfas long as radius of leaf.Geographic Distribution. Savannah of Guayaquil (2ii'25"S., 8o2'io" W.)(Caspary 1878). Ecuador (Hemsley 1888). Ditches at Hatillo, Afiasco, Porto Rico,coll. P. Sintenis, No. 5688, Dec. 7, 1886.Notes. Fruit, seed, tuber and habits unknown.Nymphaea stenaspidota Caspary. (Plates XXI, XXIV.)Sepals densely marked with long slender fuscous striae, oblong-ovate, somewhatacute, never acuminate. Petals 20 to 24, short-oblong, obtuse. Stamens 60 to 67, resemblingthose of N. gardneriana, but with narrower anthers. Carpels 18, clavate, 4to 5 mm. long. Leaves ovate-triangular, sagittate, small, rounded at apex, entire,scarcely peltate (1 mm.) ; sinus margins straight.Nymphaea stenaspidota, Caspary 1878, fid. specimens coll. Gardner, No. 2476, and 3568, in hbb.Berlin, Kew, British Museum.Description. Flower 91 to 100 mm. in diameter. Sepals 18 to 19 mm. wideby 46 mm. long. Inner petals bearing abortive anthers, very short, elongate-ovate orlanceolate. Stamens like those of N. gardneriana, but narrower; outermost anthers1.5 mm., innermost 1 mm. wide.Lobes of leaf acute, produced ;sinus large, triangular;otherwise as in N. gardneriana.Tuber ovate or sub-globose, small, 15 mm. in diameter, covered above with verylong brownish hairs.Geographic Distribution.(i2i5'S., 57 W.), collected by Gardner in 1841.16In a small lake at Conceicao, province Goyaz, Brazil
2ioThe Waterliues.Nymphaea tenerinervia Caspary. (Plate XXI.)Sepals with a few scattered linear black marks, ovate-oblong, shortly acuminate,sub-acute. Petals 16 (?), sub-membranaceous, outer ones oblong, innermost sub-lanceolate,acute. Stamens about 40, three- fourths as long as the petals;outer filamentspetaloid, median filaments about as wide as the anthers, innermost filaments very shortand slender. Carpels numerous; styles short (5 mm. long), clavate, 1 mm. in diameterabove, incurved. Leaves ovate-cordate, entire, sub-acute, narrowly peltate, slightlycoriaceous ;lower surface usually with small, black, oblong, radially placed spots ;lobesdistant, obtuse, inner margin convex. Top of peduncle and petiole, and thence downwardon petiole, clothed with slender multiseptate hairs.Nymphaea tenerinervia, Caspary 1878, plate 36, and MS. in bbb. Munich, Berlin, etc.N. tenuinervia, Caspary,1. c. in text.N. pulchella, Lehmar.n MS. in hb. Konigsberg. Not DeCandolle 1821 b.N. Maximiliani, Lehmann 1853 a, as to the leaf ;flower is of N. odorata gigantea, q. v.Description. Flower 86 to 89 mm. in diameter, or larger. Sepals 15 to 20mm. wide by 41 to 51 mm. long, subcoriaceous, sub-translucent, shining. Innermostfilaments narrower than the anthers.Veins of leaf immersed beneath, the midrib aloneprominent ;lobes slightly produced.Geographic Distribution. On St. Francisco River, province of Bahia, Brazil(40 W., 9 50' S.), near Joazeiro, collected by Martius, April, 1819.Nymphaea oxypetala Planchon. (Plate XXV.)Sepals without striae, lanceolate, oblong below, long caudate-acuminate, apextwisted or hooked. Petals 20 to 30, shorter than the sepals, lanceolate, acuminate,very acute. Stamens 84 to 106 or more, outermost lanceolate with filaments 4 timeswider than anthers, inner ones linear, innermost with filament about as wide as anther.Carpels 20 to 28, styles very long, linear-clavate, 24 to 26 mm. long. Submergedleaves alone known ; thin, membranaceous, sub-lunate-reniform, sub-hastate or obovate,cleft at base, not peltate; lobes very long; margin subrepand, slightly crisped; primarynerves 3 to 7 on each side; petiole slender.Nymphaea oxypetala, Planchon 18526; 18530 and b. Caspary 1878.N. Raja, Lehmann 18530 and b; 1854 (name only).Description. Flower 123 to 160 mm. across. Peduncle slender. Sepals11 to17 mm. wide by 61 to 74 mm. long. Outer stamens short-appendiculate (1.5 mm.),acute ;inner ones obtuse ;all the anthers longer than the filaments.Geographic Distribution. Guayaquil, Ecuador (2n'25" S., 7959'io" W.),coll. Jameson, March, 1845, No. 334. Types seen in hbb. Delessert, Boissier, BritishMuseum, Berlin.Nymphaea gibertii(Morong), new comb.Leaves oval, entire, rounded at apex lobes ;moderately spreading, obtuse under;surface covered with short branching fuscous lines. Flower white, inodorous, 6 to 7cm. in diameter. Sepals oblong-ovate, 3 to 3^ cm. long, somewhat acute, with numerousfuscous lines on outer surface. Petals in 3 series [12], a little shorter than the
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 25 /i )NYMPH/EA OXYPETALA.i. a. Flower, j, 4, Outermost and innermost stamens. 5, Three carpellary styles. 6, 7. Leares s. Globose idioblaHEUOTYPE CO., BOSTON.
Taxonomy Nymphaea gibertii. 211sepals, marked with faint purplish lines, the outer ones sepaloid on the back, withbroad white margins. Stamens in 4 series ;outermost with anther cells 6 to 7 mm.long. Carpels about 18.Castalia Gibertii, Morong 1893, in Morong & Britton.Leuconymphaea Gibertii, Morong MS. on co-type in hb. Acad. Nat. Sci. Phila.Geographic Distribution. Paraguay, collected on the Pilcomayo River aboveAsuncion by Gibert in 1858 (in hb. Kew), and by Dr. Thos. Morong 1888-90,in hbb. New York, etc. Flower, May and June.Notes. The imperfect published description of this species does not give certainevidence as to which section of the genus it belongs in ;but from the lines on thesepals, the seriate arrangement of petals and stamens, and the locality in which it wasfound, we supposeit belongs here.Names Not Elsewhere Entered.Nymphaea albo-rosea, Teijsm., ex Regel., Ind. Sent. Hort. Petrop. (1866) 101, fid. Index Kewensis.Quid?N. advena, Soland. = Nuphar.N. arifolia, Salisb. = Nuphar.N. Cahlara, Donn, Hort. Cantabr. ed. vii, 159 = N. stellata Willd., fid. Index Kewensis.N. ceramica, Rump., Amb. 6: 173, pi. 72 = Menyanthes indica Willd., fid. Roxburgh 1824.N. crenata, Amadeo, Bot. & Veg. Materia Medica of Porto Rico (Ph. J., 3 ser., vol. 18. London1888, p. 761 .). Quid?N. crcnulala, Raf., in Med. Repos. N. York, 5:354 (1808) name only; fid. Index Kewensis. Quid?N. exumbonata, Rupr., in Bull. Phys. math. Ac. Petersb. 7^:219 (1854) = N. alba, fid. Index Kew.N. hastata, Michx. = Nuphar.N. hirta, Kurz ex Teijsm. & Binn., in Tijds. Nedl. Indie 27: 38 (1864) ; fid. Ind. Kewensis. Quid?N. Kalmiana, Sims = Nuphar.N. longifolia, Michx. = Nuphar.'N. lutea, Linn., Thunb., Walt. = Nuphar spp.N. macrophylla, Small = Nuphar.N. microphylla, Pers.= Nuphar.N. minima, Reichb. = Nuphar.N. nelumbo, Linn. = Nelumbo nucifera, Gaertn.N. odorata, Willd., Hort. Berol. t. 39 = A r .nitida, Sims, fid. Index Kewensis.N. officinalis, Gater., PI. Montaub. 99 = N. alba, fid. Index Kewensis.N. orbiculata, Harper = Nuphar.N. pentapetala, Walt. = Nelumbo lutea Willd.N. polysepala, Greene = Nuphar.N. pumila Hoffm. = Nuphar.N. punctata, Kar. & Kir., in Bull. Soc. Nat. Mosc. (1841), 376, fid. Index Kewensis. Quid?N. Reichardiana, Hoffm., Beitr. z. Kenntniss von Central-ostafrika. Inaug. Dissert. Jena, 1889 (notseen).Quid?N. rubrodisca = Nuphar.N. rufescens, Willd. ex. Casp., in Journ. Sc. Math. Lisboa n. 16 (1873), reinipr. 8, name only, fid.Index Kewensis. Quid?N. sagittata, Vers. Nuphar.N. sagittifolia, Walt. = Nuphar.N. suaveolens, Dura., in Bull. Soc. Bot. Belg. 3 7 : ( 1864) = N. alba, fid. Index Kewensis.N. sumatrana, Miq. Fl. Ind. Bat. Suppl. 387, fid. Index Kewensis. Quid?N. umbilicalis, Salisb. = Nuphar.N. Victoria, Schomb. = Victoria.
CHAPTER VI.DISTRIBUTION.On account of their aquatic habits, requiring still water, mostly fresh,and enduring comparativelylittle sedimentation, the areas in which waterliliesoccur are at times scattered. They are found here and therealmost all over the world. New Zealand and the Pacific Slope of NorthAmerica have no representatives of the genus Nymphaea. The Castaliagroup is wholly restricted to the northern hemisphere, and Brachyceras tothe tropics and subtropical regions, but both are found all round the globe.Lotos ismostly tropical, and is restricted to the Eastern Hemisphere,where itranges from Sierra Leone and Senegambia to the Philippines,while Hydrocallisis native to the tropics of the New World. N. giganteaoccurs only in tropical Australia and New Guinea.Nymphaea is, generally speaking, a genus of the low lands. TheAtlantic coastal plain of North America is the home of N. odorata, andN. flavais found in brackish water in the Little River, Florida. N. oxypetalawas collected at Guayaquil, Ecuador. In the West Indies, the Philippines,and Australia they occur along the coastal swamps, and at rivermouths. The delta of the Nile is a favorite haunt of N. lotus and caertdea.On the other hand, both of these species are reported from central Africa,and similar ones from the interior of India. N. tetragona and alba ascend1500 meters in Kashmir, and our own N. odorata ascends 600 metersin the eastern United States.The distribution of single speciesis often very wide, and the variationscorrespondingly great, insomuch that the limitation of species is, inour present state of knowledge, more than usually arbitrary. Carefulstudies of plants from many localities under cultivation and hybridizationare needed to place this branch of our subject on a better footing.Another uncertain feature is the range of the Hydrocallis species. Theirnative countries are all so little known botanically that most of them mustbe left out of account in these generalizations. Outside of the Hydrocallisgroup, probably the most restricted species is N. mexicana, knownonly from Florida, Texas and Mexico. N. tetragona, on the other hand,213
214 The Waterlii.ies.is, as now constituted, the most widespread species;it extends fromSiberia through Kashmir and China to Australia (?),and is known fromGranite, Idaho, and Ontario Province, Canada, in North America. N. lotusis also very widespread, but further investigationis needed to determinethe relation of the various forms. In N. thermalis of the warm springs ofHungary we have its most northern representative. And its existencehere is a problem of extreme interest. The Grosswardein locality wasfirst recorded by Waldstein and Kitaibel (1802). Plants were removedfrom there by Paul Kitaibel in 1800 and planted in the warm springsof Lucasbad at Budapesth. In the former place the temperature of thewater is33.75 to 41. 25 C, and the water is quite pure; at the latterstation the water isonly 26 C, and ischarged with sulphur and othermineral matters. Nevertheless the plant grew equally well in both places,and flowered freely from May to October. The Lucasbad station was,I believe, destroyed by building operations about 1890 (Schilbersky, 1889).Kitaibel, Boissier, Neilreich, Kerner, Engler, and Simonkai (1890) consideredN. thermalis as identical with A^. lotus of Egypt, but Simkovics(1883) agrees with DeCandolle in pronouncing it a distinct species. Ifit could be shown, as some have believed, that the plant was originallyintroduced here by man, perhaps by the Turks, the question of identitywould be settled. But Simonkai considers this theory impossible, sincethe plant was of no material use; and Staub (1891)' agrees that it isalmost certainly a relic of a pre-glacial vegetation, when the whole ofEurope enjoyed a comparatively warm climate. When the plants of thisperiod were driven back toward the equator by colder conditions, this oneremained in the favorable temperature of the thermal springs. Alongwith it has continued a snail, Melanopsis parreyssi Muhlf., very near of kinto M. costata Fer. of Egypt. According to Staub, the genus Melanopsiswas common inEurope in the Drift period and he considers that the;warm springs of Grosswardein are older than the Drift. It is also notablethat there were in recent geological times throughout Europe waterliliesresembling the Lotos group (Schenck, 1888 ; Staub, 1. c.) which may havebeen the ancestors alike of the Egyptian and the Hungarian plants. Thereare the best of reasons, therefore, for considering N. thermalis as trulynative in itspresent habitat. And if we can believe that its specific characterswere demarcated as early as the Drift period, and have not changed1 It must be noted that Staub's paper, as it appears in Engler's Jahrbuch, is full of the mostegregious errors concerning the relationships and distribution of living species of Nymphaea; wehope they are misprints!
Distribution. 2 15since, we most follow those botanists who make this name a synonym ofN. lotus L.The questions of the place and time of origin of the genus Nymphaeaitself are obscure, although several fossils of this and allied genera areknown. All of these have been found in Europe. They establish theexistence of distinctly Nymphaeaceous types as early as the lower cretaceous.Indeed, at that time the three tribes Nelumboneae, Cabombeae,and Nymphaeinae were already well marked. Saporta (1894 b) recordsBraseniopsis venulosa (a form very near to Brasenia peltata), two speciesof Nelumbium and a Nymphaeineous plant from Valle-de-Sobreiros,Portugal. The last-mentioned plant was represented by rhizomes foundin a schistose marl. The leaf cushions were distinct and slightly prominentthe; petiolar scar showed the traces of many irregularly distributedair-canals ;below this were one to four root-scars of irregular size andarrangement. Alternating with the leaf bases are much smaller peduncularscars, with air-canals as above these ;occupy the place of leaves in thephyllotactic spires. We do not know whether this stem was erect orhorizontal, but the plant can most readily be considered as a generalizedform, from which the tetrasepalous Nymphaeinae, and perhaps Nuphar also,have been derived. Another remarkable fossil from the same beds appearsto be a fruit with the stigma and general shape of N. alba but wholly!superiorIf this interpretation be correct, it is a most interesting find.In the tertiary, the Nymphaeineous stock reached a very high development.The genus Anoectomeria Sap. (1865 a, b),of which we maytake A. brongniartii (Casp.) Sap. as the type, had flowers and fruitsresembling those of N. alba, but twice as large. It had an outer row ofthree floral leaves, and within this a second alternating row of three ;therest of the floral leaves were spirally arranged.It was probably apocarpous; at any rate, the fruit broke up readily into elliptic or lunatepieces, though Saporta says these correspond with the bases of the petals.The rhizome was cylindrical and horizontal, and marked with large rhomboidalleaf-cushions. These have the curved scar of a broad stipular platein front as in Castalia and Victoria.The petiole scar shows two large aircanalsand two smaller ones above these. The root scars are arrangedas in Eucastalia. The leaf was large (28 to 40 cm. long), entire, fissicordate,not peltate. There were 15 to 18 primary veins, which branchand dichotomize to form a very fine network;they form no primary areas,but run pretty distinctly to the margin of the leaf. Thus we see in therhizome, stipules, leaf-margin and shape of fruit a marked resemblance to
2l6The Waterlii.ies.Eucastalia, and in air-canals and venation to Lotos, Nelumbo or Victoria ;but the trimerous symmetry has not come down to us in any living form.Anoectomeria brongniartii (Casp.) Sap. (1865 a, b) (= Nymphaea ArethusaeBrongn., 1822, not of Weber, 1850 ;= Nymphaeites Brongniartii andVia. 76. Nymphaea dumasii Sap. (after Schenk, 1888).Arethusae Casp., 1856) flourished in Eocene times, being plentifulin thelittoral chalk marls of the Marseilles basin. A. media Sap. (1894 c)occursin the Aquitanian (Miocene), and A. nana Sap. in the tertiary of Alsace(Mieg, etc., 1890), although Schenk (1888) speaks of the genus as restrictedto the Oligocene. Nymphaea polyrkiza Sap. (1865 a) (= N eocenica Sap.,1 86 1, in part) from the Aquitanianof St. Jean de Garguier,resembles Anoectomeria in thetrimerous flower, and should onthat account be removed fromthe genus Nymphaea;it alsohas a large number of air-canalsin the petiole.In other ways,however, it is near to N. gypsorumSap. (1865 a).On some other speciesstatements are conflicting. N.calophylla was at first (Saporta,1861) said to have denticulateleaves, but in 1894 (b) the samewriter describes the leaves asentire. The venation of this, asof most other tertiary Nymphaeas, was similar to that of Anoectomeria,and very suggestive of the Lotos ;group but they had the rhizome andstipules of Eucastalia. Saporta considers that their type has becomeentirely extinct. In this list are included also N. gypsorum Sap., N amelianaSap., A^. nalini Sap., N rousseti Sap., N and N. dumasii N Sap. (Fig. 76).near of kin to tetragona, and cordata Sap.Nymphaea minuta Sap. isto the smaller forms of N alba, while N latior Sap. resembles the largeN alba of Greece. Wessel and Weber (1855) compare their N lignitica(= Nymphaeites lignitica Casp.) from the lignites of Rott in the lowerRhine region with N. alba, with regard to the shape of the leaf. N.parvula Sap. and charpentieri Heer {Nymphaeites charpentieri Casp.;=e Nelumbium nymphaeoides Ettingsh.) had double flowers like those ofN. alba (Saporta, 1879). Nymphaeites microrhizus Sap.is a small form
Distribution. 2 17with two air-canals in the petiole and one root per leaf ;the figure givesita certain resemblance to N. mexicana. Nymphaeites palaeopygmaeus Sap.had also two main air-canals and three to five roots, but Saporta consideredit near to Anoectomeria. Nymphaeites weberi Casp. (= NymphaeaArethusae Weber, 1850, not of Brongn., 1822 =; Carpolites granulatusWeber) and Nymphaeites ludwigii Casp. (1856) probably are akin to theNymphaea calophylla group. Nymphaea dubia Wat. was shown by Bureauand of Nymphaea blandusiae Ung., doliolum(1888) to be really a Nuphar;Ludw., doris Heer, and Nymphaeites saxonica Fried, we have learnednothing but the names.During the warmer portions of the Tertiary the genus Nymphaeaextended far into the arctic zone. N arctica Heer (1869) of Spitzbergenhas left quite complete remains. Its rhizome and leaf suggest N. alba ;there were four main canals in the petiole and anumber of evident onessurrounding these. The sepals were elliptic, with slender longitudinalveins. There were about 12 carpels, and the fruit was much enlargedbelow the narrow (6 mm. wide) depressed stigma. It certainly belongs tothe Castalia group. Nymphaeites thulensis Heer was found in the samelocality, but poorly preserved. In Grinnell Land also Captain Feilden ofthe English North Polar Expedition found N arctica (Heer, 1878 a, c)and an allied fragment named Nymphaeites tener (Heer, 1878 b).From such forms and such localities doubtless all of the Castalias ofEurope and America are alike derived. The genus Nymphaea probablyoriginated in western Europe and spread over the Arctic continent. Veryearly, thoughfossil evidence does not indicate when, the syncarpous andapocarpous groups were separated. Of the latter, N. gigantea, from itsabsence of styles and restricted distribution in the ancient Australianflora, is most primitive. The lowest Brachyceras types are the shortstyledand pale-flowered American forms A^. elegans and flavo-virens, fromwhich we ascend throughA^. caerulea and stellata to zanzibariensis, the mostgorgeously colored, largest-flowered, longest-styled species of the group,and also the farthest removed from the original center of distribution.Among Syncarpiae, the Castalia group is most primitive, and of thesewe have already mentioned A^. mexicana as the lowest, with N. tetragonaas a close second. In Europe the Lotos section was segregated off fromthese at a very early date, probably through forms somewhat similar toN calophylla and gypsorum. N. lotus is nearest to the parent stock, andN. rubra most removed. N. thermalis remains to testify to the minimumage of the type. A^. Candida has kept near home and has probably
218 The Waterliues.changed less than its fellow Castalias, whereas N. odorata by its rich odor,and N. tuberosa by its rhizome show marked specialization as comparedwith any other species of Nymphaea. Finally, in the American tropics,Hydrocallis has arisen from the Castalia stock. This groupis too littleknown to assign relationships among the species, but it is probable thatN.OIQAHTCAN.ZANZIBAR1ENSI8N. RUBRAN.CAPENSISN. STELLATAN 7.FIAVQ- V1BEN8
CHAPTER VII.HYBRIDS AND GARDEN VARIETIES.Beginning with Nymphaea devoniensis in 185 1,number of forms of waterlily has become known to cultivators.a continually increasingThe originof these was in some cases accidental, in others due to artificialhybridization.In the first case there may have been cross-pollination by insects,or variation or mutation all equally problematical to the systematist anduncertain for the general student. In the second case, careless or ignorantmethods leave room for much doubt as to parentage, and very oftenthe facts are withheld or distorted for trade purposes. Only in very fewcases, therefore, is our knowledge of the origin of the various gardenvarieties at all reliable or useful.For example, N. devoniensis, which appears on plate 4665 of thePaxton in Gardeners'Botanical Magazine, and was first described byChronicle for July 10, 1852, is said in the former article to have been producedby crossing N. rubra with N. lotus a red species with a white.The seeds were obtained in autumn of 1850 at Chatsworth, England, andthe plantis said to have flowered from April to October, 1851. But thefrom N. rubra are more robustpoints of difference of the so-called hybridgrowth, more floriferous habit, and a very slightly lighter color. In otherwords, there is almost no trace of the influence of N. lotus. Since Hookerand Thomson stated that just such a plantis common and native in India,its hybrid origin is generally disbelieved. It is probably a self-fertilizedseedling of N. rubra. As to the color of leaves and flowers, we had inthe summer of 1901 three flowering plants of A^. rubra (?) raised from oneand the same tuber. One was grownin a half-barrel of earth in a tank1.4 meters square, incompany with N. marliacea-chromatella, N. tetragona,and the water-poppy the flower was of a ;light magenta red, and the leavesbrownish green. Another was in a half-barrel in a pond with plenty ofleaf-room, and had deep red-brown leaves and pure magenta flowers. Thethird was in a tub of earth 1.6 meters in diameter and 60 cm. deep, withplenty of leaf-room ;it resembled the second, but had larger leaves andflowers, and the latter stilldeeper in color. There is, therefore, greatvariation, according to conditions of growth.219
220 The Waterliues.On the other hand, N. ortgiesiano-rubra, exhibited at Chiswick inMay,1852, by M. van Houtte, and described by Planchon (1852 ad), is in manyrespects intermediate between the two parent types. The seed wasobtained in the summer of 1850 by M. Ortgies in Van Houtte's aquaria,by removing the stamens from a flower of N. rubra and dusting the stigmawith pollen of "A 7 ",ortgiesiana." The comparison of hybrid and parentsis well shown in Planchon's parallel column descriptions. The hybrid hasbright rose-pink flowers, but they open wide as in N. ortgiesiana. Thestyles are yellow, washed with red in N. ortgiesiana, reddish orangein thehybrid, dark brownish red in N. rubra. The color of the stamens andupper surface of the leaves isfairly intermediate, but the under surface ofthe leaf and the character of the stigma are as in N. rubra. The robusttemperament of N. ortgiesiana is reproduced in the hybrid, but it is morerloriferous than either parent, probably because no strengthis used upinthe production of seed. In short, N. ortgiesiano-rubrais a well-markedhybrid. Several other hybridizations are recorded by Planchon (1. c.) asproductive of seed in 1852, but no further results reached the public.In 1853 M. Bouche, Inspector of the Royal Botanic Garden of Berlin,obtained another hybrid, named by Planchon (1854) N. boucheana. It isto allappearance intermediate between N. lotus and N. ortgiesiano-rubra,but is of the same parentage as the latter. A slightly different form ofN. lotus, viz., N. dentata, was used as pollen parent. The pale pink petalsbordered with pure pink, the yellow stamens, and the bright green leavesshow a preponderating influence of N. lotus. The sterility of N. boucheanaattests its hybrid origin.N. kewensis was next to appear. It was published in Gardeners'Chronicle in 1887 (p. 366), and in Botanical Magazine in April, 1888(tab. 6988). The cross was made at Kew in 1885 by Mr. Watson, withN. lotus as seed parent and pollen of N. devoniensis. The hybrid has alarge number of broad petals of an even rosy pink all over the;stamensare orange colored the leaves are ;pure dark green above. In recentyears, crossing and re-crossing has been carried on in this group, especiallyin America, until every shade of color may be had between the whiteand red types differences;are also noted in shape of petal, size of flower,and color of leaf.They are propagated readily and quite accurately fromthe tubers, and names have been given to every distinguishable form.N. ortgiesiano-rubra and boucheana are not found under those names, butvery similar plants are cultivated. The name kewensis is still retained,though the original plant "died without issue"; the present stock was
CARNEGIE INSTITUTION OF WASHINGTON. WATERLILIES, PLATE 26.NYMPH/EA PENNSYLVANIA.
Hybrids and Garden Varieties. 221obtained independentlyin America. Hybrids of the Lotos group areknown inGermany under an entirely distinct set of names from thoseused in this country.In 1885-1890 M. Latour-Marliac, of Temple-sur-Lot, France, drew theattention of flower-lovers everywhere by his magnificent hybrids in theCastalia group the hardy waterlilies. Probably no one else has done somuch for waterlily culture. His earliest achievements won universaladmiration but these have been followed;by annual additions of everincreasing splendor. At present, however, the possibilities in that directionseem to be about exhausted. So great was Marliac's lead in thiswork, and so carefully did he guard the secret of his success, for tradereasons, that only within the last three years have any comparable resultsbeen attained, and this only by using Marliac stock for breeding. Theparentage of the Marliac hybrids is a matter of speculation. In severalcases, however, one or both parents can be indicated with considerablecertainty but which was male and which female ; will, I fear, never beknown. The first members of this series were the yellows, N. marliaceachromatella,N. odorata sulfurea and N. tetragona helvola. All are derivedfrom N. mexicana. They are altogether sterile. Of some of the pinksA^. odorata rosea is one parent, and of those with red tints at the center ofthe flower A^ alba rubra is one source.Within the Brachyceras group, hybridization has been carried onalmost exclusively in America. Here Mr. Tricker is the acknowledgedleader, so far as the public goodis concerned. His A^. pulcherrima,derived on one side from A^. caerulea, has been for several years ingeneral cultivation. The new hybrids of N. jlavo-virens with pollen of A7 !zanzibariensis sent out under the rather unwieldy names Mrs. C. W. Wardand Wm. Stone, are easily the finest tender day-blooming hybrids yetavailable. Both have the general habit oi N. jlavo-virens,and in the tuberthey follow this parent exactly ; but the color of the flower and leaf isbeautifully intermediate. N. Pennsylvania and A7 ".elegansX,raised in 1901 at the University of Pennsylvaniaseeds of the previous summer, are fairly intermediate inzanzibariensis,Botanic Garden fromforms, coloring ofall parts, and number of floral organs between their respective parents.succeeded betweenIn spite of many attempts, no cross has yetmembers of different subgenera of Nymphaea as grouped in this paper.We realize that this is in opposition to M. Marliac's published statementsin which he claims A^. rubra as the source of the red color in his hardy redwaterlilies. But so far as naked-eye examination goes (and we acknowl-
222 The Waterlii.ies.edge that is weak argument) the kind of color shown by the hybrids istotally different from that of N. rubra, and no trace of the latter species isto be found in any part of the plant. We are also forced to take issueconcerning the supposed hybrids between Lotos and Hydrocallis l raisedby Mr. Ames at North Easton, Mass. (Ames, 1900; Grey, 1900).1 havethe greatest confidence in the accuracy of Mr. Ames, and of his ableand the latter assures me that the seed-flowers weregardener, Mr. Grey,carefully castrated and kept netted from insects. But the so-called hybridsshow absolutely no trace of the influence of a Hydrocallis parent, and it iseasier to believe that parthenogenesis or asexual budding from the endospermor perisperm has occurred in the highly fertile N. lotus flower thanto imagine a hybrid parentage with one parent obliterated. The casedemands, and might well repay, a rigid investigation. Similar crosses ofdistantly related species have been repeatedly claimed, and by suchauthorities as Planchon, Tricker, and Sturtevant, but they have either failedto reach the public or have shown no trace of the pollen parent;sometimes,indeed, there is strong evidence of the pollenof some relatedspecies.Natural hybrids,i. e., those produced without human intervention, areof frequent occurrence whenever kindred species are in the same pond.Caspary and others have recorded hybrids of N. alba and Candida fromEurope, and intermediate forms between N. odorata rosea and N. tuberosaspring up in our gardens N. odorata caroliniana and N. 0. luciana are of;such origin.N. flavo-virens always becomes mixed with N. zanzibariensisif the two are near each other.From the fragmentary nature of our knowledge it is impossible todraw any general conclusions on the laws of heredity in waterlilies. Sofar as we can see, some characteristics are regularly blended in the hybridoffspring, while others are carried over in toto, or even exaggerated, fromone or other parent. Thus, the spotting of the calyx so characteristic ofN. caerulea is not a whit diminished in N. pulcherrima and A^. Pennsylvania,while in both the color of petal and size, shape and number of allof the floral parts are decidedly intermediate.In N. Wm. Stone, as statedabove, the caudex is exactly like that of the seed parent, N. flavo-virens,in size, shape, character of leaf-bases, hardiness, and keeping qualities. InBrachyceras all known hybrids are sterile, except N. capensisK zanzibariensisraised in our botanic garden, which is highly fertile, and of which a new1Mr. Ames' N. ampla, which I saw in 1901, is N. rudgeana Mey., collected by Mr. Grey inCuba. The so-called crosses may be " false-hybrids."
Hybrids and Garden Varieties. 223generation of hybrids is coming on. All of the hardy yellow hybrids areentirely sterile, and crosses of N. tetragona with Eucastalia types are alsosterile ; as examples of these N. laydekeri rosea and N. pygniaea albaMarliac may be mentioned. Even N. alba x Candida is greatly enfeebledin seed-producing capacity. But N. alba and N. a. rubra, though muchmore distinct than the last two to the casual observer, are perfectly fertiletogether and have fertile progeny. N. odorata and tuberosa hybrids arenot self-fertile, but can be crossed with pure species. In the Lotos group,some forms are sterile, others fertile. Planchon found N. ortgiesianorubrato be impotent as to the ovules, but capable of fertilizing N. lotus orN. rubra. In those now in cultivation, N. devoniensis is nearly sterile, butN. omarana, which closely resembles iV. ortgiesiano-rubra, often producesseed. In general, the nearer the hybrid approachesin color to the highlyfertile N. lotus, the more productiveit is. Several seedlings of N. omaranahave been raised, of which N. Geo. Huster deserves special mention.Its flowers are of an extremely deep dark red, more so than N. rubraitself, as cultivated here. If the original seed was fertilized by pollen of thesame flower, which it isimpossible to know, we would have a case of"separation of characters" according to Mendel's theory; but even ifN. rubra was the pollen parent, the determining elements of the color ofthe flower in this seedling seem to have lost all trace of the influence of awhite ancestor.We have expressed a belief in the occurrence of variations undercultivation without mixture of blood.Probably N. odorata exquisitais sucha variant from N. odorata rosea, as also N. froebelii from N. alba rubra,N. devoniensis from A^. rubra, and N. gladstoniana from A^. alba ; N. gladstonianais quite fertile, and has seeds and arils like A^. alba. WhetherA^. alba candidissima belongsin this class or among the hybridsis uncertain.It is absolutely sterile, and may be a cross of N. Candida with thelarge N. alba of Greece. Quite comparable with these varietal changesin natural forms are some suddenly appearing peculiarities of hybrids.For example, N. robinsoni can be easily recognized by a peculiar crimpednotch and fold about the middle of each side of the sinus of the leaf;N. gloriosa is ;uniformly pentamerous and one of a set of plants ofN. gladstoniana Xmexicana raised by Mr. Tricker habitually twists thepetiole so as to turn the apex of the leaf instead of the sinus toward thecenter of the plant. N. laydekeri rosea proliferais a fasciated strain,which stools out around the crown to a remarkable degree, whereas thetype makes no lateral buds at all.
224 The Waterliues.Without going farther into isolated details, enough has been said toshow that a wide and interesting field liesopen to the student incrossing and inverse crossing of waterlilies. It is unfortunate that Caspary'sextensive investigations in this line have not been formulated, buthis notes, exquisite color sketches, and large collection of pressed specimensare preservedin the Royal Herbarium of Berlin. Among them aresuch crosses as N. capensis X caerulea ; N. capensis X (capensis X caerulea);N. caerulea x [caerulea X {capensis X caerulea)] N. caerulea X; \caerulea X-[caerulea x (capensis X caerulea);N. flavo-virens X [capensisX (caerulea Xmicrantha)] .] \Fio. 77. Numphaea pulcherrima Tricker. Sepal, three successive petals; threesuccessive stamens, vertical section of ovary. Two-thirds natural size.Duplicate specimens of many of these hybrids were distributed toseveral other herbaria. Time did not permit us to make a study of these.We hope they will be worked over by some scholar, and the interestingresults made known. We have, for the convenience of students and cultivators,brought together, in the following pages, some account of the hybridsand garden varieties that have come more directly under our notice.
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 27& kNYMPH/EA CAPENSIS X ZANZIBARIENSIS.MELIOTVPE CO., BOSTON.
Hybrids and Garden Varieties. 225Section I.Subgenus II.Nymphaeae apocarpiae.BRACHYCERAS.i. N. stellata X caerulea. Mentioned by Caspary (1877) as a sterile hybrid.2. N. caerulea X capensis (Focke 1881). Raised by Caspary in 1862. Itresembles N. caerulea very closely and is generally sterile, though Caspary succeededin fertilizing it with pollen of other individuals of the same stock and generation, andobtained a second generation of hybrids. These resembled those of the first generation,except one individual, which produced styles as in N. capensis, and petals of adeeper color than those of either parent. The small violet-black spots on the calyxof N. caerulea are more plentiful in the hybrids. N. pulcherrima (Tricker 1897)gives every evidence of having this parentage ;the spotting of calyx and under sideFio. 78. Ifympfiaea Pennsylvania. Sepal, petals, stamens, and section of ovary. Two-thirds natural size.of leaf,the purple margin of the leaf beneath, and the conical buds are reproduced asin N. caerulea. The sinuate leaves with acuminate lobes are like those of N. capensis.The color of petals and number of floral leaves are fairly intermediate. Sepals 4,petals 20 to 22, stamens 137 to 140, carpels 24. Sterile. (Fig. yy.)3. (N. caerulea X capensis) second generation X caerulea (Caspary 1862).Spots of calyx confluent into large areas.4. N. caerulea X (caerulea X capensis). Raised by Caspary (1862).5. N. caerulea X zanzibariensis (=A7. Pennsylvania Conard 1901 b) (PlateXXVI; Fig. 78). Flowers light blue, open from 8 a. m. to 5 p. m., that is, combiningthe earliest and latest limits of the two parents. Spotting of calyx and lower surface ofleaves as in N. caerulea. Shape of bud, shape, number and color of floral organs, and17
226 The Waterlilies.outline of leaves strikingly intermediate. The seedlings are like those of N. caeruleauntil the first floating leaf; this is spotted with brown above and suffused bluishbeneath as in N. zanzibariensis. Sterile. Raised at the Botanic Garden, UniversityFig. IV. Nymphaea capemi* X zanzlbarlen*U. Sepal, four petals, stamlnode, stamens, vertical section ofovary. Two-thirds natural size.Fia. 80. Nymphaea elegant x zanzibarUruis. Sepal, three petals, stamens, vertical section ofovary. Two-thirds natural size.of Pennsylvania, from seed obtained in 1900; flowered 1901. Sepals 4, petals 23,stamens 140, carpels 26.6. N. capensis X zanzibariensis (Plate XXVII; Fig. 79). Flowers light blue.Floral parts very numerous. Sepals broad, resembling those of N. zanzibariensis.
i".;>rt
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 28NYMPH/EA ELEGANS X ZANZIBARIENSIS.HELIOTYPE CO., BOSTON.
Hybrids and Garden Varieties. 227Leaves dull purplish violet beneath, less brilliant than in N. zanzibariensis.Self fertile.The types are very much alike, but the hybrid is fairly intermediate. Raised in theBotanic Garden of the University of Pennsylvania from seed obtained in 1900; flowered1901. Sepals 4, petals 32, stamens 225, carpels 32. N. stellata eastoniensis (Gardenand Forest p." 475) appears to belong here, the cross having been effected by insects.It is described as having steel-blue flowers, with less of the stellate form, andleaves " more deeply toothed " than the type. It was raised by C. Blomberg, gardenerto O. Ames, North Easton, Mass., from seed obtained in 1895 from " N. stellata"(doubtless N. capensis) ;flowered 1896.Fio. 81. Nymphaea " Wm. Stone." Sepal, petals, stamens, section of ovary.Two-thirds natural size.7. N. elegans X zanzibariensis (Plate XXVIII; Fig. 80). Flowers light bluewith a violet tint, much bluer than N. elegans. Bud, sepals, and petals shaped as inA/, elegans, but much larger. Length of styles and appendages of stamens intermediate.Leaves colored as in N. zanzibariensis, but intermediate in shape. Sterile. Raised in theBotanic Garden, University of Pennsylvania, from seed obtained in 1900; flowered1901. Sepals 4, petals 31, stamens 150.8. N. zanzibariensis X ? (= N. zanzibariensis X capensis, Conard1901 a). Leaves as in N. zanzibariensis, but without brown spots above. Flowers richblue, open from 9 a. m. to 4 or 5 p. m. ; petals and sepals narrower than in N. zanzibariensis.Petals 15 to 20, stamens 60 to 100. Very floriferous, but sterile. Cultivatedat Horticultural Hall, Fairmount Park, Philadelphia, and formerly at University ofPennsylvania. Said to have been originally obtained from Mr. Tricker.9. N. flavo-virens X zanzibariensis (Plate XXIX; Fig. 81) (=N. Greyae,Hort. ;N. Wr.i. Stone Tricker; N. astraea R. M. Grey; N. gracilis purpurea, N. g.
228 The Waterlii.ies.asurea Sturtevant). Sepals, petals and leaves shaped as in N. flavo-virens, but the lasttwo intermediate in color. Stamens and carpels intermediate. Tuber in all respectslike N. Havo-virens. Petals 20, stamens 70, carpels 21. Sterile. First obtained by Ben . jGrey of Maiden, Mass., and named in his honor.Fio. 82. Nymphaea "Mrs. C. W. Ward." Sepal, petals, stamens, section of ovary.Two-thirds natural size.10. N. flavo-virens X zanzibariensis rosea (= N. mauvii Henshaw ;N. Mrs.C. W. Ward Tricker ;N. astraea rosea R. M. Grey ;N. gracilis rosea, N. g. r. perfecta,N. g. rubra Sturtevant). Like No. 9, but with flowers pink instead of blue. Sterile.First obtained by Samuel Henshaw, New Brighton, N. Y., about 1892. There areslight differences in the quality of the pink color as grown by different persons; itusually contains a shade of blue, but Mr. Sturtevant has nearly a pure pink (perfecta) ;his rubra variety is rather deeper in color than any others we have seen. (Fig. 82.)
CARNEGIE INSTITUTION OF WASHINGTON WATERLILIES, PLATE 29NYMPH^EA FLAVOVIRENS X ZANZIBARIENSIS.MELIOTVPE CO., BOSTON.
Hybrids and Garden Varieties. 229Section II.Nymphaeae syncarpiae.Subgenus III.CASTALIA.Chamaenymphaea X Xanthantha.11. N. tetragona X mexicana (=A7. pygmaea helvola Marliac; N. tetragonahelvola Conard 1901a; Froebel Cat. 1901). In shape and size of all the parts likeN. tetragona, with the color in leaves and flowers of N. mexicana. The caudex bearsmany lateral buds, but no runners. Sterile. Produced by Marliac about 1890; introducedinto America about 1892.Chamaenymphaea X Eu-castalia.12. N. tetragona X alba? (= N. pygmaea alba Marl., in part). ResemblesN. tetragona in shape and size of flowers, but the leaves are broader and more rounded.The caudex bears no side shoots. Sterile. Sent out by Marliac as N. pygmaea alba in1893 ; but in other places he uses this name apparently as synonymous with N. tetragona.13. N. tetragona X alba rubra? (= N. Laydekeri rosea Marliac; N. LaydekeriHort.).Closely resembles N. tetragona, except in color of flowers; these are ofa delicate pink color on opening, changing to rose and deep carmine-rose on succeedingdays; this character is drawn from N. alba rubra. Sterile and without sideshoots. Introduced by Marliac about 1891. N. laydekeri rosea prolifera Tricker is afasciated form of the above, producing a great many lateral buds. Introduced byHenry A. Dreer Co., 1901. N. laydekeri lilacea Andre (1895 a, col. plate) (= N.liliacea Gard. and For. 1895, p. 277; N. lilacina 1. c. 1897, p. 316; N. Laydekeri liliaceaTricker in Dreer Cat. 1902) has flowers of a soft rosy lilac, shaded bright carmine, withyellow stamens; otherwise as above. Raised by Marliac; introduced into the UnitedStates in 1895.Eu-castalia X Xanthantha.14. N. odorata X mexicana (=A7. odorata sulfurea Marliac; N. sulfureaHort.). The flowers are intermediate in size, shape of parts, and color betweenthe two parents; the leaf combines the copious brown blotching of N. flava with thedeep red under surface of N. odorata. Rhizome slender, nodose, yellow. Sterile.Obtained by Marliac about 1888, and exhibited by Lagrange at the Trocadero in 1889.Published with colored plate in Revue Horticole 1890 b. About 1899 a larger formwas sent out by Marliac under the name of N. sulfurea grandiflora.15. N. odorata rosea X mexicana? (=A7. arc-en-ciel Marliac 1901). Knownto us only by catalogue description, but apparently of this "parentage. Leaves variegatedwith several lively and distinct tints. * * * The sweet scented flowers are of asalmon-white color, with sepals splashed with rose color."
230The Waterulies.16. N. alba X mexicana? (= N. tmrtiacea chromatello Marliac; A/, chromatelloHort. ;N. tuberosa flavesccns of .Kew) Leaves orbicular, much blotched withrise as much as 20 cm. above thebrown, 8 to 20 cm. across; when crowded they maywater, and are not blotched. Flowers bright yellow ; petals numerous, broad, concave.Very strong grower, but sterile. Said by Marliac to have the parentage cited above,but believed by some to be N. tuberosa X mexicana. Structurallyit bears someresemblance to N. tuberosa, and sometimes the brown stripes of the petioles of thisspecies appear on the hybrid the question cannot yet be decided. Raised by Marliac;about 1888.17. N. alba rubra X mexicana? Several of the Marliac hybrids undoubtedlyhave this parentage, but most distinctly N. robinsoni (N. robinsoniana Hort.), seignoreti,aurora, and fulva (= N. laydekeri fulva Tricker 1897). The leaves are muchmottled with reddish-brown as in N. mexicana. The outer petals are more or lessyellow, shading through rose to deep red on the inner ones; the stamens are redorange. Thus we have a beautiful blending of the colors of the parents. A7. robinsoniis peculiar in having a crimped notch about the middle of each margin of the sinus.There are also differences in the details of the coloring of the flowers. They werefirst published in Revue Horticole in 1895, N. robinsoni and seignoreti with a coloredplate, p. 258, and the other two on p. 568. In the former article JV. andreana andlucida (= N. laydekeri lucida Tricker 1897) are also mentioned, and in the latterplace is N. laydekeri fulgens with a colored plate. All are productions of Marliac.AT. andreana appears on a colored plate in the same journal in 1896, pp. 352-353. Thelast three also agree in having the leaves mottled with brown, but the yelloworcolor isnot nearly so evident in the flower ; they may represent a second cross. N. chrysantha(Marliac 1901) is described in almost the same words as N. aurora, and N. arethusa(Henry A. Dreer Co. 1902) is spoken of as " an improved Laydekeri fulgens."Eu-castalia.(To N. alba we refer the following garden forms A7. alba maxima; N. :a. plenissima;N. gladstoniana, raised by Richardson of Lordstown, Ohio; N. casparyi albaCarriere, Rev. Hort. 1888, p. 300. To N. alba rubra belongs N. froebelii. To N.odorata rosea we refer N. odorata rubra; N. rosacea; N. exquisita. To N. odoratagigantea we refer N. gracillima alba Marliac Cat. 1901 ; N. caroliniana nivea MarliacCat 1901 ;N. eburnea Andre, Rev. Hortic. 1895, p. 568. To N. odorata minor belongN. odorata pumila and N. union. To N. tuberosa we refer N. tuberosa richardsoni;N. t.plena.)18. N. alba X Candida? (= N. alba candidissima Hort.; N. candidissimaHort). A large, white, free-flowering variety ; very robust. Sterile.19. N. alba X odorata? (= JV. marliacea albida Marliac). The flower ispurewhite, stamens yellow, sepals flushed pink ; fragrant. Leaves large, deep red beneath.On account of the color of the sepals, and a tendency to pink on the stamens, thepollen parent may have been N. 0. rosea, and the plant, in that case, belongs withN. m. cornea and rosea (No. 20), which it resembles in every respect except color.Sterile. Introduced by Marliac about 1889.
Hybrids and Garden Varieties. 23120. N. alba X odorata rosea? (N. marliacea carnea; N. m. rosea; N. Wm.Doogue). Differs from No. 19 only in color of flower, which is from light pearlypink to deep rose; all are strong, luxuriant growers but sterile. Under the first twonames it was introduced by Marliac about 1889; Wm. Doogue was obtained by HenryA. Dreer Co. ten years later.21. N. alba X odorata rosea? (N. colossea Marliac 1901). Flowers and leavesvery large petals flesh colored. Blooms from early spring until frost. Sweet scented.;It may be a second hybrid, N. alba candidissima X odorata rosea.22. N. alba rubra X A large number of hybrids and variants havebeen introduced recently possessing the characteristic red shading of the Swedishwaterlily, and without distinct indications of other parentage. Perhaps N. Candidahas been used (as the flowers are often cup-shaped), or they may be second or thirdor multiple crosses ;the latter ishighly likely. Some of the types are fairly distinct,even to an amateur, but most of them can be recognized only by the professional. InN. marliacea Hammea, N. m. ignea, and N. gloriosa there is a faint blotching of theleaf which can come only from N. mexicana or N. tetragona. We have not noted anysuch character in N. ellisiana,sanguinea, marliacea rubra punctata, laydekeri purpurea,Wm. Falconer, James Gurney, or James Brydon (= N. brydonacea elegans).23. N. odorata rosea X tuberosa? (= N. caroliniana, c.perfecta, c. rosea; N.tuberosa superba; N. odorata caroliniana, 0. carolinensis, o. superba, o. luciana, 0.suavissima). This hybrid is a much stronger grower than N. odorata rosea, butotherwise resembles it. The cross easily occurs, but was first cultivated by Dr. Bahnsenof Salem, North Carolina, about 1890.24. N. tuberosa X odorata rosea ? (= N. tuberosa rosea; N. brakeleyi rosea).Flowers pink. The plant has nearly the habit of N. tuberosa, but the rhizome is lesstuberiferous. N. tuberosa rubra (Sturtevant 1901)is said to be N. brakeleyi rosea XN. rubra; but from the descriptionit is more probably (TV. tuberosa X odorata rosea)X alba rubra. The " rosy red " flowers and " bright red stamens " indicate this. Theplant was not flowering when I saw it in Mr. Sturtevant's garden (June, 1902) ; theleaves were small, entire, and petioles dark red.Subgenus IV.LOTOS.25. N. lotus X rubra. Successive crosses between these species have been veryoften made, and every possible gradation between them has been obtained. They differin color and size of leaf and flower and relative breadth of petal and degree of openingof the flower; the hybrids vary also in texture of leaf and in distribution of color onthe petals. In N. sturtevantii the leaves are crimped at margin far more than ineither parent and the color of N. Geo. Huster is darker than any strain we have seen;of N. rubra. According to Focke (1881) N. rubra with N. lotus or ortgiesiana givesthe same product whichever is used as pollen parent, and the same is true of N. lotusand dentata. But in N. rubra and dentata the hybrid progeny resembles the femaleparent in respect to relative length and breadth of early leaves and floral leaves.
232 The Watermues.With regard to color, the forms now in cultivation, with originator,date of origin, and reputed parentage, may be arrangedin series, thus :1. N. dentata magnified (N. omaraua X dentata) Bisset 1902.2. N. dentata grandifiora Sturtevant 1901. White.3. N. eastonensis (N. omarana X smithiana) Ames 1900.4. N. jubilee, Dreer 1899. Delicate pinkish white. N. Queen Elizabeth (N. devoniensis X ortgiesiana)1855 (fid. Focke).5. N. smithiana, Tricker 1893. N. indica Isis (cf. Moenkemeyer 1897). N. laelia (N. colombianaX smithiana), and N. I. colorans (ditto) Ames 1900.6. N. delicatissimo, Tricker 1894. Light pink.7. N. deaniana, Tricker 1894. N. boucheana (N. rubra X dentata) Bouche 1853 (cf. Planchon1854). N. indica Spira (cf. Moenkemeyer 1897).8. N. kewensis (N. dentata X devoniensis). Raised at Kew, 1885 (cf. Botanical Magazine, tab.6988). Pink to red.9. N. sturtevantii (AT. devoniensis seedling) Sturtevant 1884. N. indica Hofgartner Graebener(cf. Moenkemeyer 1897). N. rubicunda (N. sturtevantii X lotus) Ames 1900.10. N. omarana, Plate XXX (N. dentata X sturtevantii), Bisset 1894. Magenta. N. ortgiesianorubra(JV. ortgiesiana X rubra) Ortgies 1852 (cf. Planchon 18510a). N. mariae lagrangei(N. lotus X dentata) Lagrange (cf. Andre 1899). N. niobe (N. ampla X rubra) Ames1900 (cf. page 222 above).11. N. diana (AT. sturtevantii X ampla) Grey 1900 (cf. above, p. 222). N. indica Brahma (FroebelCat. 1901).12. N. devoniensis (N. rubra X lotus) at Chatsworth, Eng., 1851. N. arnoldiana Oliver, about 1899.13. N. Columbiana, Tricker 1894. N. diana grandifiora (AT. sturtevantii X ampla) Ames 1000 (cf.above, p. 222).14. N. George Huster (N. omarana seedling) Dreer 1899. Dark red.15. N. Frank Trelease, Gurney 1900.Focke (1881) mentions, without further comment than is here given,a considerable list of crosses involving species of the Lotos group. Weappend the list, referring back to p. 221 for our views on those cases wheretwo subgenera are involved.N. Queen Elizabeth X dentata gave red and rose-colored flowers.N. Queen Elizabeth X caerulea gave violet flowers.N. rubra X ortgiesiana = N. ortgiesiano-rubra.N. ortgiesiana X rubra = N. ortgiesiano-rubra.N. rubra X lotus = N. lotus X rubra.N. lotus X dentata = N. dentata X lotus.N. alba X ortgiesiano-rubra gave seedlings which never matured. A reported cross of N. devoniensisand Victoria regia, Focke says ismistaken.Planchon (1. c.)mentions the following attempts at hybridization, but wecan find no further record of the plants:N. thermalts X rubra; young plants obtained.N. thermalis X scutifolia; young plants obtained.N. alba X ortgiesiana; young plants obtained.N. dentata X ortgiesiano-rubra; seedlings obtained.N. alba X ortgiesiano-rubra; three seedlings obtained.
CARNEGIE INSTITUTION OF WASHINGTON.WATERLILIES, PLATE 30.NYMPH>EA OMARANA.
Hybrids and Garden Varieties. 233Unsuccessful attempts were made with N. scutifolia X dentata andthe reverse ;N. scutifolia X rubra and reverse ;N. odorata x rubra, andN. ortgiesiana x caerulea. It is notable that all of these failures involvedtwo subgenera.It would be interesting, could we trace any definite and constantrelations among hybrids. A diagram of the pedigree of some of thebest known varieties may aid in this. We select N. omarana and the" "rose aitd red flowered hybrids of N. Queen Elizabeth and dentata.In each couple we place the pollen parent above.N. dentataN. dentataN. dentataN. dentataN. lotusN. lotusN. lotusN. rubraN. dentataN. dentataN. lotusN. devoniensisN. dentataN. sturtevantiiN. omaranaAccording to this a single infusion of JV. rubra has sufficed toproduce a rich magenta color in the fourth generation. But in Mr.Bisset's new N. dentata magnifica (JV. omarana x dentata), the tide iscompletely turned and the flower is pure white. The next instance,however, is like that of N. omarana, viz. :N. dentataN. dentataN. dentataN. dentata}, dentataN. dentata1J[N. dentata'Rose and red flowers.iV. ortgiesiana 1 ., . .N. N.ortpesuma ortgiesiana } ).V. lotusN. rubra},}N. Queen Elisabethdevoniensis JIs red, then, a dominant color in the Mendellian sense, and is ittherefore primitive? Discussion would at present be wholly speculativeand had best be omitted. A wide field for investigation is opened up.But none of our garden stock can be warranted pure enough to starton. We need fresh importations of the natural species, and a carefulcomparison of a large number of sister plants. The custom of gardenersis to discard all seedlings whose foliage does not give indications of thedesired results ;consequently only certain types have been preservedfor us.
Tl; :CHAPTER VIII.CULTURE AND USES.use of waterlilies for food and ornament among the ancient Egyptianshas already been referred to. This still continues to some extent inEgypt, but, on the West Coast of Africa, Nymphaea seeds constitute animportant article of food. Travelers describe seeing the native womenand children coming in at evening to the villages laden with the ripeningfruits. These are laid in heaps until all of the soft parts decay. Theseeds are then easily washed out clean, and are dried and stored away.They contain so much starch, oil, and proteid that they should form a verynutritious diet. When dry the seed coat is brittle and the kernel slightlyshrunken, so that the two are easily separated. The kernels may beground into a kind of flour or may be boiled whole. The resting tubers,in the dry season, are almost solid starch, and are eaten boiled or roastedlike potatoes. The species used in West Africa are N. caerulea andlotus. In Madagascar, N. lotus and capensis are similarly used, and theseeds of N. gigantea are eaten by the Australians. In Central Americaalso waterlily seeds are used for food. Victoria cruziana is known as" water corn " {Mais del aqua) in Paraguay on account of its large farinaceousseeds.In the days of empirical medicine and the " doctrine of signatures"the medicinal virtues of Nymphaea were highly overrated. Its leaveswere said to be antipyretic, and a decoction of the dried leaves andrhizomes was supposed to be powerfully anaphrodisiac. Althoughit stillholds a place in the United States Dispensatory, and is mentioned byTavera (1901) in his medical botany of the Philippines, it is probably neverused. The whole plantisvery astringent and bitter, on account of thepresence of tannin and gallic acid (cf. Griining, 1881 ; Niederstadt, 1883 ;Fridolin, 1884). Whatever value it has is due to these constituents.But it is as ornamental plants that Nymphaeas have achieved theirpresent wide-spread fame, especially inEurope and North America.English colonies in India and Australia are also giving them considerableattention, and a beautifully arranged pond has long been a feature of theBotanic Garden of Buitenzorg in Java. Their culture began in the large235
236 The Waterliues.estates and botanic gardens of England with the introduction of N. odoratain 1786. From that time until 181 2 several species were brought toEnglish gardens. Then there was a period of inactivity until the appearanceof Victoria about 1840 aroused great interest in aquatics. From1850 to i860 this wave reached its maximum, when Planchon, Lehmann,and Hooker described several species, and Caspary beganhis immortalwork on the genus. There followed a lull until near 1880. ProbablyN. zanzibariensis served to bring its congeners before the public again.Another wave of waterlily culture set in, which probably has not yetreached its maximum. So magnificent are the results recently achieved,and of so lasting a nature, that this branch of gardening is to be oflikelyimportance as long as flowers are grown for the beautifying of public orprivate grounds.For cultural purposes we may divide the genus into two groups thehardy and the tender varieties. In the first group Castalia stands alone.All others are tender in our climate. In the Botanical Magazine andFlore des Serres brief cultural directions are given for each species described;these are almost wholly intended for tender varieties when grownunder glass. But since the extended culture of these plants can only becarried on where the climate will admit of their being for three months atleast out of doors, we shall speak only of the latter condition. Out-doorculture of tropical forms is of very recent date. So recently as 1891 HerrBorsig's water garden in Berlin was noted as of especial interest. Hewarmed the pond with waste hot water from his factory. Mr. Vilmorindescribed the place in Revue Horticole, and the article was reproduced intranslation in the London Garden, and in Garden and Forest (1891, p. 375).In Garden and Forest for 1888 (pp. 241-242) cultivation of waterlilies inshallow ponds is urged as both possible and desirable in the eastern UnitedStates. In the garden trade the hardy species were slower than thetender ones in getting recognition. Gardeners' Chronicle in 1848 informsa correspondent concerning N. alba that " the roots of these things maysometimes be had in the nurseries, but they are not usually kept." In1849, however, it is stated that N. rubra-, caerulea, stellata, and ampla may"be had from any of the large nurserymen." Vick's Catalogue for 1883mentions only A^. odorata. At present sixty-four varieties are entered inthe catalogue of one of the leading American dealers.The care of them isvery simple. The pond or tank may be only alarge bucket or a half-barrel ;in such accommodations N. tetragona, N. t.helvola, N. marliacea chromatella, or even N. caerulea, will get along and
Culture and Uses. 237bloom;the first two will do quite well. But the best thingis a brick andcement pool 60 cm. deep and of as large a size as one can get room for ;after one season's trial the tank will seem too small, at any rate.A singlecourse of brick will be sufficient for a small tank, but a double course isbetter. Mr. Arnold, of Sharon Hill, gets beautiful results with a poolwhere the earth was simply scooped out and an even coat of cement laiddirectly on the ground. In any case it is best to protect the edges of thepond in winter by a coat of litter if a weak construction is used this is;absolutely necessary, as a leak caused by frost would let the water outand the resting rhizomes would be fatally frozen. A natural pond,if notfrequented by muskrats, is excellent for hardy varieties, but it would berare to find one north of Washington warm enough for the tender kinds.JV. alba candidissima, odorata gigantea, and colossea will do well in water asdeep as one or two meters. Select a place for the tank where it will getfull sunlight all day. Waterlilies are usually but little shaded in theirnative haunts, and the tropical species need all the light possible whenin our northern climes. It is desirable to have the water-level constant,and to change the water just enough to keep it fresh.For planting the rhizomes it isonly necessary with the strongerhardy varieties in natural waters to tie a stone close to the rhizome andthrow it in where the depthwill be suitable. If one would be more careful,the rhizomes may be planted in weak splint baskets of earth and putneeds more room the basket will be fall-in place. By the time the planting to pieces, or at least, will be easily broken, and the roots will have fullliberty.Such planting may be done at almost any time of year, but preferablyin fall or spring. Sometimes on account of weakness, sometimesbecause of excessive dominance, it is desirable to keep the plants withinprescribed bounds in that case ;permanent pots, boxes, or tubs must beprovided. This isnecessary in artificial ponds. Nothingless than a halfbarrelis really worth while. A tub 100 to 130 cm. in diameter and 45 to50 cm. deep gives the best results for all of the larger species, but I haveseen N. elegans and gigantea bloomingin tubs only 40 cm. across, andN. zanzibariensis will produce pretty little flowers in a "six-inch" pot!Square board boxes of any size may be made to suit circumstances.The character of the soil for these pots or tubs is of prime importance;it must be veryrich. Various prescriptions for its preparationhave been given. R. Miiller (1885) recommends a mixture of three partssoil, two parts compost, and two parts peat for hardy Nymphaeas. We findnothing equal to well-rotted cow manure for fertilizer, and Mr. Tricker's
238 The Wateruues.(1897) preparation of good garden soil or sod compostwith one-third toone-half of manure gives the finest possible results. Swamp muck is notat all to be recommended. If well-rotted cow manure can not be had,bone dust may be substituted in the proportion of "a six-inch potfull to abarrow load of soil " (Tricker, 1. c). Even in such strong materials a goodplant of any of the larger tender forms will completely exhaust the nutrimentin a half-barrel of soil before the end of the season ;hence the desirabilityof larger tubs for the best results. The plants should be placedwith the crown on a level with the surface of the soil. This should becovered with 1 to 3 cm. of clean sand to keep the manure down and toprevent the activities of certain aquatic worms which often carry up muchfine matter to the surface of the tub and tend to bury the plant. For thehardy varieties the tub must be placed deep enoughin the water to besafe from actually freezing. Tender varieties need to be within 15 to 25 cm.of the water surface in order to take advantage of the greater warmthof the upper strata. The area of water surface required per plant dependsupon the species and the size of the tub it is planted in. I have seenN. rubra, N. marliacea chromatella, and N. tetragona in a tank 120 cm.square, with the water-poppy flourishing in whatever space remained ;andall four species bloomed freely all summer. Each of the Nymphaeas hadone flower every day, and sometimes two, and the water-poppy sometimeshad a dozen flowers at once ! But they were badly crowded. A^ tetragonawill be satisfied with a space 60 cm. square most of the hardy varie-;ties require a space 100 to 120 cm. square for a single shoot; and thetender varieties will cover a space 3 to 5 meters across.Once placed as above described, the plants need no attentionthroughout the season except to keep up the water supply and trim offdead leaves and flowers. They should bloom at the rate of a new flowerevery two or three days, or perhaps one every day. As each flower laststwo to five days, the display is always fine. Of course the varieties differin production of flowers, and the situation and other circumstances havetheir effects.Hardy species should not be moved for two or three years atleast, but the tender ones must be wintered with considerable care. Largeblooming plants may be lifted with a ball of earth 30 to 60 cm. in diameter,potted, and kept in a warm tank with plenty of light. Theywill thenbloom all winter, and may be planted out again.It is best, however, tolet old plants of the tender day-blooming species die. The nightbloomersshould be taken up in late October or early November, pottedin sand or planted under the bench of the conservatory, and allowed
Culture and Uses. 239slowly to dry off. In about a month they will almost certainly producestrong tubers, which should be kept in dry sand but not allowed to becomeabsolutely desiccated. In March or April tubers of all of the tender varietiesshould be planted in sand in a warm tank. As each stolon gets itsfirst floating leaf it should be carefully detached from the tuber and pottedoff, and the tuber replanted for the development of additional stolons.With ter-der day-blooming species (apocarpous) it is important to obtaintwo or more plants in the spring. One of these should be kept in a6-inch or 8-inch pot, and dried off about mid-summer toform a tuber forthe next season, as the flowering plants usually die.Hardy varieties arepropagated from the rhizomes, by cutting and planting out in spring.Propagation by seed is also easy, but is not possible for hybrids. Theripening fruits are covered with muslin bags the mouth of the bagis tied;fast around the peduncle, and the bag should also be tied to a stick forsafe keeping. After the fruit has burst and its parts decayed, the bag istaken in and the seeds washed from the debris. Should the pods burstunexpectedly and the seeds be found floating on the pond, they may bedipped up with a fine wire sieve tied on a pole. Seeds of the Castaliagroup must be kept in water in a cool place until ready for sowing.Other seed may be dried and kept in the usual way.It is advisable tosow Castalia seeds as soon as ripe; their germination is at best irregular,and it is unusual to get flowering plants the first year. The tender varietiesmay be sown in flat pans, barely covered with sand, and placed inwarm tanks in February. The young plants should be pricked off intosmall pots and kept moving rapidly;they will produce plantsof thelargest size the first season. Planting out in the pond must be delayeduntil alldanger of frost, or even of a cold rain storm, is over. The first ofJune often proves too early in this latitude. N. gigantea is especiallytender. The labeling of seeds and plants in water can only be successfullydone on metal labels. Thin copper or brass on which one writeswith a stylus, or with asphalt varnish, makes the only satisfactory anddurable label.Some hints on hybridizing have already been given, but itmay beworth while to recapitulate. No haphazard work pays.It is better toget no seeds than to go to the pains of raising a batch of supposed hybridsand find they are all pure species. The flower to be used for seed shouldbe covered with a fine net than (finer mosquito bar) before it begins to;open do it on the previous day. As soon as it allopens (or before) ofthe stamens must be plucked out. For the greatest accuracy the pollen
240 The Wateruues.flower should also be kept netted before and after it;opensavoid any mixing of pollen by insects. Now, on the day the seed-flowerthis is tofirst opens, the net is removed for a moment while, with a brush, pollenfrom the chosen male parentis stirred around in the stigmatic fluid. Thenet is replaced, and in a week or so the result will be indicated by aswelling of the ovary ifsuccessful, or by decay of the flower and peduncleif no fertilization was effected. Some consider that the pollen should beapplied at the close of the first period of opening of the seed-flower. Ihave succeeded very well in the Brachyceras group at that time or earlier,but have no successful experience in any other group.Several enemies attack waterlilies and work great havoc. In theseed beds, algae may grow up and choke the little plants ; the use of lessmanure and more sand in the pots is a preventive measure, and weakBordeaux mixture will kill the algae. After the first transplanting, adamping-off fungus sometimes carries off numbers of seedlings the leaves;become transparent in ever-increasing areas until they are all gone. Thismust be prevented by care in pricking off; if the plants are not wounded,and are not weakened by the shock of moving, they will outstrip the fungus.Snails, too, sometimes devour the leaves, or cut them off and leavethem floating on the tank ;pick out the offenders and throw them away.The first floating leaves are often seriously weakened by swarms ofaphides. A strong spray from hose or syringe will wash them off, and afew sun-fish or paradise fish will effectually prevent their return to theleaves.As to enemies of mature plants,I can do no better than to quoteMr. Tricker's summary from the Cyclopedia of American Horticulture(1901, p. 1 104):Aphides are sometimes troublesome. The best remedy is their natural enemy,the " lady bugs " or " lady birds." A colony of these voracious insects makes shortwork of the aphides, as do also the lace-winged An flies. insect of recent acquaintancewith Nymphaeas is a leaf-miner, the larva of a small fly, which cuts channelsthrough the leaf in all directions. Sometimes only a few of these are in evidence, atother times the leaves are fairly alive with them. The trouble is easily detected. Themarks suggest Japanese writing or the efforts of youthful artists. A simple and effectiveremedy is kerosene emulsion, applied with a fine spray at evening after the flowersare closed. Another troublesome insect has its home in Florida, and has comeNorth to spend the summer in a favored clime. This is a leaf-cutter, Hydrocampaproprialis. The larva cuts out pieces of the leaf and hides between two pieces, whichmake a kind of tent. In this tent the larva moves about. At first it moves slowly,but as it nears maturity the larva becomes ravenous and then eats the surface of the
Culture and Uses. 241leaves near the center, and cuts off much larger pieces of the leaf for camping-outpurposes. The best remedy for this pest is a lamp trap for the mature insect. Frogsand dragon-flies will catch numbers of them.Nymphaeas are also subject to a fungous disease, a leaf spot which is easily discernedafter a spell of warm humid weather. After such a spell of weather, followedby bright sunshine, the leaves are scorched and crumpled up, and as a result, the plantis sadly crippled by being denuded of its foliage; new leaves are weak and smaller,and so too are the flowers, if indeed there are any. This disease must be checked atonce or thr plants will be severely set back, if not ruined. The only remedy isBordeaux mixture, or any of the various mixtures with sulphate of copper as the basis.Use a fine spray, and dilute the mixture to half the strength recommended for mostplants. It is best to spray twice with a weak solution rather than to spray once withtoo strong a solution and damage the foliage.Finally, the ravages of muskrats eating the rhizomes of hardy speciesin winter must be mentioned. Three fine collections, inmy small acquaintance,have been seriously injured or destroyed in this way. Traps anda bounty on hides are two practical aids to the gardener. But here, as inso many cases all along life's journey, " "eternal vigilance is the price ofsuccess. And the success, once it is attained, more than repays the laborof love which itcosts.18
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INDEX.Note. Synonyms and horticultural names are in italics. Asterisk (*) after a number indicates a figure. Numbers inheavy-faced type refer to the taxonomic description of a species or group ; such descriptions always treat in order (1) diagnosis,(2) synonymy, (3) detailed description of flower, leaf and stem, (4) geographic distribution, (5) notes of miscellaneousinformation.PAGEAerial leaves 64Aestivation 98Agrostis 8AM 8Air-canals, function of 60, 118gases in 117-118in leaf 70peduncles 59*, 78-79petals 82petioles 58, 59*, 60, 62, 76development of 96of seedling 112roots 31-33, 37'sepals 81-82Alplnus 16, 17, 20, 21, 23Ambel 23Ameling on cell dimensions 69Amenhotep 6, 8, 9Ames, O 222Analytic Key 126-127Anecphya 127flower 80symmetry of 101phylogeny 218pollen 85seed 90stamens 85Anoectomeria 215, 216, 217fruit 87phylogeny 218brongnlartli 215, 216media 216nana 216Anther, development of 101-102Anthotaxy 42, 43Aphides 240Apical growth of roots 36-37Appendiculatae 127, 131Apocarplae (gee Nymphaeae apocarplae) 127carpels 85-89flower 78odor of 121leaf 65, 66, 77girder-like veins 115ovary 86*peduncle 02, 78, 79petiole 61collar of 62phylogeny 217, 21820PAGEApocarpiae, propagation 239root 28stamens 83, 85stem 39Apogeotropism of peduncles 114petioles 114Arcangeli on idioblasts 60, 61, 70leaves 54stomata of water-leaves 77Archesporlum 101-102*Aril 90-91*development of 104Arum 16Astomatlc area 68Athenseus 13-14, 17Axlle process 80, 88Bacon on dead Insects in flowersBaillon on floral diagram of NupharBarthglemyon air-canals of petioles 58movement of gases 117respiration 117Bauhln, C 15, 19, 22,Hauhin, JBeardsleeBellonus, PBesBeslerBlue lotus 5, 7, 8,Bock, Hieronymus 14Bodaeus a StapelBoissler on N. thermallsBonavlaBontiusBoolak MuseumBotanical MagazineBrachycerasdistributionflowersubmergence of, after anthesls ....germinationhand-pollinationhybrids 221, 222, 225leafovarypedunclepetals sepaloldphylogeny 217,2651221007260118118252219216919146, 182021472195131213801231062402286486*79100218
266 The Waterlilies.PAGEBracbyceras, pistil 86, 89, 80pollen 85pollination 122seedling 109stamens 83, 84, 85symmetry of flower 101Bract 90, 100, 101Bracteole 100, 101Brand on stomata of water leaves 77Brasenia 24mucilage on 112Brasenia peltata 215Brasenlopsls venulosa 215Braun on inflorescence 100Breynins, J 23, 24Britten 14and Holland 16Buckbout 182Buckley 7Cabombeae, geologic history 215phylogeny 218pistil 87Calcium oxalate in idloblasts 61Carpels 80, 84-90Carpellary style 86*, 88, 90Carpolites granulatua 217Caspary 21, 24, 25, 64, 66, 70on development of leaf 97embryo of Nuphar 104hybrids 224inflorescence 100leaf-traces 53ovary 86, 87pachycysU 60phyllotaxy 42pollenJ . 85pollination 122root-cap 29seedlings 107stem structure 46tubers 47vitality of seeds 106Castalia 163, 215adjustment of peduncles to water level 118buoyancy of leaves and buds 114culture 236distribution 213flower 77, 80odor of 121symmetry of 101fruit 89protection of 124geologic history 217, 218germination 106hybrids 221, 229-231leaf 65, 66ovary 84peduncle 78, 79petiole 56phylogeny 218pollination 121, 122seed 90vitality of 106, 239stamens 83, 84rumCastalia, stipules 54development of 96water leaves 74, 76, 77Castalia 125, 127, 200alba 176ampla 134rm nihn 141nlitli* 198, 199elegans 131flava 163gibertii 211Hi !/
Index. 267TAGEContractile roots 113structure of 33*, 35Cook on development of megaspore 102embryo 104Cork formation In stem 47'-48Cortex of roots 31-34stems 44-45Cotyledons In germination 107-109, 108*seed 94*Cultivation, care of plants 238-239extent of 235history of .:. 235-236planting 237soil 237-238tank for 236-238Cyananthos 127Cyanea 86, 87, 127, 131Dahmen on funiculus 104Date-palmDeBary on root cap 29DeCandolle on Cyanea 86N. thermalU 214Delpino on pollination122Dhammapada4-5Diaphragms in air-canals of roots 37, 38*Diodorus Siculus 7"8. HDioscorides 11-18, 15, 16Diseases 240-241Dissection of stem apex 05Distribution 213-214, 217-218Dodonaeus 18-19Double flowers 101DuppaEgyptian lotus ..!, 6, 7, 9, 11, 12, 13, 14, 16, 17, 18, 20Eichler 80. 99-100Elwell5Embryo 94*. 104-107, 105*Embryo sac 102 *> 1 3Endosperm **. 105-106absorption of 1 9Endodermis34 * 35Enemies ****\Engler on V. thermalis 214symmetry of flower 101107, 109, 111EplcotylEpidermis of root 3' 31I 74Erythrocarpa'l 75, I 78Erythrostlgma1J;*Eucastalla "M. 215. 216floral movements 120fruit901 6germinationhybrids 223, 230-231leaf 64, 73, 7758petioles42phyllotaxy218phylogeny86, 88pistil85pollenroots28seedseedlingS4stipulesEucastalla X Xanthantha 229, 230HPAGEEuryale, cleistogamy of 121leaf 67, 77phylogeny 218root-hairs absent Illstem structure 46Faba Aegyptia 15, 16, 18, 20Fagus leaf 72False hybrids 222Fischer on pollen 85Flacourt 22Floral clock 119diagram 80', 100, 101Flower 27, 77-80adjustment to water level 118anesthesia of 121development of 97-99, 101duration of 119-120In relation to pollination 121-123odor of 121opening and closing of 119-121submergence after anthesis 123-124symmetry of 100, 101Focke on hybrids 232-233Food uses 3, 7, 10-12, 17, 22, 170, 196, 235Forgetful lotus 11Frank on position of lamina 114-115Friend 16Fruit 87, 89-90protection of 123-124ripening of 27Fuchsius 14Funeral wreaths 8, 9Funiculus, development of 103structure and contents 104Galenus 14Gaudlchaud 80Geologic history 214-217Geotroplsm of roots 113stems 114Gerard 20Germination 106-110, 108*Giles 3Gilg, E., description of N. zenkerl 197-198Goebel on development of flower 97, 98function of air-canals 60hairs '3respirationMlseedlings 107, 109symmetry of flower 1011Gollan, Wm 6Grew, N 25Grey, K. M 222Grosswardein214Gwynne-Vaughan on steles 36, 49, 52structure of seedling 111-1122 2 l3HabitsJ.Hairs on leaf lz ~' s87ovary'"peduncle56-57, 73*petioleseed 90, 92-93stem73 *
268 The Waterlilies.turnHairs on stem apex4 3-44of seedling 112stipules 06Hansteln *0Hapl 8Hardiness 236Harahberger 101, 182Heat evolved In flowers 123Hellotroplsm113Henslow on root-cap 29Hercules 13Heredity 222, 233Herodotus 10, 16Hofmelster 72on embryo sac of Nuphar 103, 104Holland, Philemon 13, 18Hooker on vlvipary of N. mlcrantha 74Hooker and Thomson 219, 141, 193Hybridizing 239-240Hybrids, history of 219-221limitations In production of 221, 222natural 222sterility of 222, 223200Hydrocalllsdistribution 213flower 78, 80odor of 121submergence after anthesls 123symmetry of 101hybrids222leaf 64, 66, 77peduncle 78petiole, air-canals of 68phylogeny218pistil 86. 88pollen 85pollination 122receptacle 79roots 28seeds 00stamens 84stem 39, 41Hydrocampa proprlalls, leaf-cutter 240-241Hydrocharls24morsus-ranae 115Hypocotyl 107, 111Idloblasts, crystals In 61In carpels 84, 85, 87, 88leaf 69, 70peduncle 79petals 83petiole 60-62roots 32, 33development of 37sepals 81, 82stamens 84, 85stem 46stipules 56water-leaves 77Inappendiculatae 127, 163, 192Inflorescence 99, 100Insects attracted to flowers 121effecting pollination 122-123found dead in flowers 122-123Iris pseudacorus 16l'AOBIrmlsch 73Iscblrch 60Jahn on causes of shape of leaf 115Jones, Sir Wm 6JoretKalldasa 4Karsten on adjustment to water level 115Kerner on opening and closing of N. alba 120pollen 85N. thermal** 214Key to species 126-127Kltalbel, P 214Klebs on seedlings 107Kny on cuticle of air-canals COKoehne on fusion of floral leaves with ovary 98Kohl 60, 61, 70Kotschy collection from Nubia 146Kuntze 25Labeling 239Lanman 5Leaf 27, 64-67, 65*, 74, 77adjustment to water level 114-115air-canals In 70buoyancy of 114development of 95-97hairs on 72-73kinds of 54of germinating tuber 75*, 190*. 195*seedling 107-110* , 108*structure of 112structure of 67*, 68*-72, 76, 77, 115, 116vascular system of 70, 71*venation 66, 67, 115Leaf-cutter 240-241Leaf-gap 51, 53-54Leaf-scar 47, 48*Leaf-spot 241Leaf-trace 51-54Leaves as food for animals 39Lechartler on function of air-canals 60experiments on respiration 117, 118Lehmann on vlvipary 74Leptopleura. See Lytopleura.Lcucanthos 200Leuconymphaea 25, 125gibertU 211reniformis 189Lilium longlflorum 10Llmnanthemum 23, 24Lobelius 15*-16, 18, 20, 22Lotophagl 11Lotos 192, 193*, 216carpels 84, 86, 87, 193*distribution 213, 217flower 77, 80, 193*movements of 120, 121odor of 121submergence after anthesis 123symmetry of 101fruit 89, 90geologic history 214, 217germination 106, 195*hybrids 221-223, 231-232
Index. 269FAGSLotos leaf 6t-66, 76, 77, 195*girder-like veins 115hairs on 73notes 193*, 200peduncle 78, 79idioblasts In 62petiole, air-canals of 58, 62idioblasts in 62thyll-Iike bodies in 62vascular system of 62phytogeny 217, 218pollen 85pollination 121, 122roots 28seed 92seedlings 108*, 109species, discussion of 193stamens 83, 84, 193*stem 39, 41stolons 41, 52*-53Lotoa 131, 163, 200Lotus 0-13, 16, 18, 146Lotus aegyptia 17, 18, 19, 21niliacum 17Lotus, blue 5, 7, 8, 146Egyptian 1*, 6, 7*, 9, 11-14, 16-18, 20genus of Leguminosae 12sacred 4, 6Lotus-eaters 11Lubbock on stipules 54-56Ludwig 25Lyte 16liytopleura 87,%127Macdowell 5Mackintosh on air-canals of petioles 58MacMillan 25Mahabharata 4Malpighl 25Marllac 221Marsilia 24Masters on air-canals of petioles 58Matthlolus 15-16, 18, 20Medicinal uses 235Megaspore, development of 102Melanopsis costata 214parreyssi 214Melllnck 60, 62Mendel's theory 223, 233Menyanthes Indlca 211Meyer, test for cork 48Microspores 85*Mleg, etc 216Millspaugh 186Mimusops schimperi 9Mollsch on crystals 61Movements of floral parts 119-121Mucilage hairs on seedlings 112Miiller, C, on collenchyma 58Mtiller, H., on pollination 122Miiller, H., on soil 237Mummy 8, 9Muskrats 237, 241Nageli on leaf-traces 53Nash 1895 187, 188PAGENatural hybrids 222Nefer TQm6*, 8, 10Neilreich on N. thertnalis 214Nelumblum, geologic history 215Nelumblum nymphaeoides 216Nelumbo 188, 216early notices of 4, 9, 11, 12, 13, 14, 15,16, 18, 20, 21, 23, 24, 25leaves 77primary root 28Nelumbo lutea 211nucifcra 4, 6, 10, 11, 211Nelumboneae, geologic history 215phylogeny 218pistil 87Nemphar 18Nenufar 14, 16, 19, 21, 25Nicolai on root-cap 29Nodal structure, N. flava 51*, 53-54Nuphar 49, 211, 215, 217bract subtending flower 99early notices of 12, 14, 15, 17, 18, 20, 22, 24embryo 104embryo sac 103, 104floral diagram 100fruit 87petiole structure 76, 77phylogeny 218root-cap 29root-hairsIllNuphar advena, embryo 104embryo-sac 104Nuphar aegypHum 17Nuphar lutea, crystals in idioblasts 37early notices of 14, 19, 25embryo sac 104Nutrition 116Nymphaea 9, 10, 11, 12, 13, 14, 15, 17,18, 20, 21, 22, 23, 24, 25, 125floral diagram 80*medicinal value 11, 12, 19, 21, 22, 235Nymphaea abbreviata 145acutiloba 170advena 211aesopU 175, 178Nymphaea alba ..172, 174, 175-179, 180, 189, 215, 216, 217carpels 84development of flower 97, 98, 101leaf 95, 96dissection of stem apex 95distribution 171, 177, 213early notices of ...11, 12, 14, 15*, 16, 17,18, 19, 20, 21, 22, 23, 24, 25floral movements, time of 120under anesthetics . . 121flower 80malformed 100submergence of, after anthesls . . 124symmetry of 100, 101hairs 72*hybrids 222, 223, 232introduction to cultivation 236leaf 64, 66, 69, 72, 73, 75, 77Nageli on leaf-traces 53notes on 177-179, 188odor of flower 121
270The Waterulies.Nymphaea alba MMNymphaea alba, petiole 56, 60phyllotaxy 42phytogeny 218pollen 85pollination 121, 122relation to N. Candida 171, 174, 178respiration 117roots 28seed 80, 91, 92, 93*stamens 83, 84stem 38synonymy 211172thyll-llke bodies In petiole 62variants 230canadensis 180candidissima 223, 230culture 237floral movements 119leaf 67, 08, 72scars 47*-48*peduncle 78pistil 88root structure 36stem 38structure 45, 46cork formation .... 47X odorata rosea 231depreaaa 178flore pleno odorata 180intermedia 176Kashmiriana 172maxima 230melocarpa 175, 178minor 168, 172oligoatigma 172, 174oocarpa 172, 174, 178oviformia 178pleniaaima 230polyatigma 175purpurea 179roaea 178, 179Nympbaea alba rubra (Plate XV) 179flower 77hybrids 78, 221, 223, 229, 231variants 223, 230X mexlcana 230Nymphaea alia aphaerocarpa 174, 178platyatigma erythrocarparoaea 178, 179rubra 179urceolata 178Nymphaea alba X Candida 175, 223, 230X mexlcana 230X odorata 230rosea 231Nymphaea albo-roaea 211albo-viridia 200Nymphaea amazonum (Plate XIX) 200-204clelstogamy in 27, 121flower, duration of 119, 120submergence of 123fruit 89leaf 64-69, 72, 73, 77peduncle 78petiole 57, 59, 60NymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaFAQBamazonum, phytogeny 218pistil 88seedling 110*root hairs on 110*, 111stomata on leaves of . . 112seeds 90, 92stamens 83tuber 40*amazonum goudotlana 204amazonum 205aubmeraa 34, 204ameliana 210americana 25ampla (Plate V) 134-136, 205, 222culture 236early notices of 21, 24, 25flower 78, 80leaf 64, 66notes 135, 136, 139phylogeny 218seed 92ampla 136, 145, 146plumieri 135, 136ampla pulchella 136-137speclosa 136ampla hookeri 136rudgeana 205y aalzmanni 136andreana 230arc-en-ciel 229arctica 217arethuaa 230arethuaae 216, 217arifolia 211arnoldiana 232peduncle 78aatraea 139, 227roaea 228aurora 78, 230bankaii 130baahiniana 172baaniniana 172, 174bella 141, 198berneriana 153, 156, 157btradiata 172, 174blanda (Plate XXI) 206-207phylogeny 218seed 92stamens 84blanda 189, 200, 205, 206blanda fenzliana (Plate XXI) 207blanda amazonum 200, 205bianduslae 217boucheana 220, 232braailienata 23brakeleyi roaea 231brownii 131brydonacea elegana 231cachemiriana 172, 174caerulea (Plate VIII) 141-146, 142, 217anther 102carpels 85, 142*culture 236distribution 145, 213early notices of 6-13, 17, 20embryo 105*I
Index. 271NymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaPAGEcaerulea, flower 80, 142*color of 121development of 97, 98duration of 119submergence of 123malformed 100fruit 142*germination 106hybrids ...221, 222, 224, 225, 226, 233leaf 64-66, 75*development of 95notes 141, 145, 146, 147150, 156, 160, 182ovule, development of 102peduncle 59*, 78petiole 59*development of 96phyllotaxy 43phylogeny 218pollen 85*pollination 122root 28root-cap 29*-30seed 91-92seedling 110*root-hairs onIllstomata of 112stamens 142*stem structure 44, 46*tuber 40used for food 235caerulea 146, 147, 151, 153albida 146, 147caerulea albiflora 145caerulea X, petiole 59*X capensls 225malformed flower .... 100X zanzibariensls (Plate XXVI) 225-226time of floralmovements.... 119cahlara 211calliantha 151*-153tenuis 152-153calophylla 216, 218Candida (Plate XV) 172-175, 173*compared with N. fennlca 170, 171distribution 171, 173-174early notices of 14, 18, 19, 20flower 80hybrids 222, 223, 231leaf 173*notes 174-175, 178phylogeny 217, 218seed 90, 92sepals 81stem 38varieties 174-175Candida oocarpa 175semiaperta 171, 175sphaeroidea 175Candida X alba 175candidissima 230candolleana 135, 205capensls (Plate X)..136, 141, 140, 163-157, 154*early notices of 22-24flower 80, 154*PAGHNymphaea capensls, flower, movements of 119hybrids 224-225leaf 64, 65notes 156-157, 160phylogeny 218pollen 85roots 28seed 90, 92seedling, root-hairs of Illstomata on 112tuber 40*used as food 235Nymphaea capensls madagascarlensis 157zanzibariensls (Plate XI), 167-161, 158*forma azurea 161rosea 161(See N. zanzibariensit.)Nymphaea capensls x ,leaf 71*X zanzibarienaia (Plate XXVII) 222. .223, 226*, 227pollination 122time of floralmovements ... 119Nymphaea caroliniana 231nivea 230perfecta 231rosea 231caspari 179caspary 179casparyi 179Plba 179, 230ceramica 211Nymphaea charpentieri 216Nymphaea chromatella 230chrysantha 230coerulea s N. caerulea.colossea 231, 237Columbiana 232Nymphaea cordata 216Nymphaea coteka 198crenata 211crenulata 211cubogermen 174cyanea 140, 153deaniana 232delicatissima 232dentata 194, 196flower 77, 78movements of 119hybrids 161, 220, 231-233leaf 75, 77peduncle 59*, 78petiole, vascular system of 63*pollen 85seed 92vitality of 106seedling, leaf-structure 112root-hairs of Illtuber 40dentata grandiflora232magniflca232devoniensis 193, 195*, 199, 200floral movements 119history of 219, 220, 223, 232peduncle78(liana 232
272The Wateruues.PAGENymphaea dlana grandiflora 232dlosoortdl* 175, 178discolor 141Nymphaea dollolum 217dorlg 217dubla 217dumasil 216*Nymphaea eastonensis 232eburnea 280edgeicorthil 140cdulis 198, 199Nymphaea elegans (Plate IV) 131-184, 132*, 217culture 237flower 80, 132*movements of 119hybrids 227leaf 64, 66, 75*phylogeny 218pistil 86, 132*seed 90, 93seedling, root-hairs of Illstomata on 112stamens 83, 132*tuber 40*Nymphaea elegans X zanzibariensis (Plate XXVIII) ... 221226*, 227floral movements... 119germination 106root epidermis .... 30structure, 31-36, 38*Nymphaea ellisiana 231emirnensls 153, 156, 157eocenlca 216erythrocarpa 176, 177, 178rsculcnta 198, 199exquislta 61, 230exumbonata 211Nymphaea fennlca 170-171flower 77, 78leaf 77phylogeny 218sepals 81stem 40, 41Nymphaea fenzllana 206, 207flava ( = N. mexlcana) 163, 164*distribution 213flower, duration of 119submergence after anthesls 123, 124fruit 87, 89, 90, 164*hybrids 78, 229leaf 65*-69, 72, 75*. 76development of 95nodal structure 51*, 53-54peduncle 59*perennatlng body 74-75petiole 57*-61, 59*development of 96-97phyllotaxy 43pollen 85root structure 30, 81, 33-35seed 90, 91, 93, 94seedling 109, 165*mucilage hairs on 112stamen 84*. 164*stem 40hairs on apex of 43, 44rinNymphaea flora, stem, structure, 43, 45, 46, 49, 50', 53, 54stipules 55*. 50stolons 48-49, 53Nymphaea flavo-vlrens (Plate VI) 137-189, 138*, 217floral movements 119hybrids 221, 222, 224, 228leaf 64-66, 74-76, 75*peduncle, adjustment to waterlevel 119*petals 138*phylogeny 218pistil 86, 138*pollen 85*root 28contractile 33*, 35, 113structure .31, 33, 34, 37*, 38*root-cap 29-30seed 90, 92*stamens 83, 138*stem 39stolons 41tuber formation 47Nymphaea flavo-vlrens x zanzibariensis (Plate XXIX),227-228root, contrac-** tile 113root, structure 33Nymphaea flavo-vlrens X zanzibariensis rosea 228leaves . . 75*Nymphaea foetida 201folUs ctrcinnatls maximis 135cordatis integerrimis 168fragrans 208Frank Trelease 232froebelii 223, 230fulva 230Nymphaea gardnerlana (Plate XXIV) 208Nymphaea George Huster 223, 231, 232Nymphaea gtbertli 210-211gigantea (Plate III) 128-131culture 237, 239distribution 130, 213flower 78^duration of 110malformed 100leaf 64-66, 75*petiole 58, 59*phyllotaxy 42phylogeny 217, 218pistil 86seed 02stamens 83, 85tuber 40*used as food 235Nymphaea gigantea forma alba ISOrosea 180Nymphaea gigantea vlolacea 130-131Nymphaea gladstoniana 223, 230peduncle 78seed 00, 93seedling 109X mexlcana 223t/landulitera 200gloriosa 223, 231goudotiana 204gracilis 136, 137, 139
Index. 273NymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaNymphaeaPAGEgracilis, petiole ( N. flavo-virons) 59*seed ( = N. flavo-virens) 92*azurea 139, 227-228purpurea 139, 227rosea 228perfects 139, 228rubra 139, 228gracillima alba 230greyae 139, 227uuiancnais. (See N. guineensis.)guineensis ... v 74, 146, 147, 149, 150gypsorum 216, 217hastata 211headelotii 147-150, 148*, 153nana 149-160hcpat ivu 164hirta 211hookeriana 141indica 22, 23, 24brahma 232Hofgartner Qraebener 232isis 232spira 232integrifolia 201intermedia 172James Brydon 231James Qumey 231jamesonlana (Plate XXIII) 209jubilee 232kalmiana 211kewensis 232history of 220-221kosteletzkyi 172, 174laelia 232colorans 232lasiophylla (Plate XXII) 207-208latlor 216laydekeri 229fulgens ,230tulva 230lilacea 229liliacea 229luclda 230purpurea 231rosea 223, 229prolifera 223, 229leiboldiana 136, 153llgnltlca 216Ulacina 229liliacea 229lineata 136longifoUa 211lotus (Plate XVI) 194-196, 199distribution 196, 213-215early notices of ...3-12, 16, 18, 20, 21-25embryo 105*embryo-sac 103flower, development of 97-100, 99*submergence after anthesis . . . 123symmetry of 100transformed to tubers 196funiculus 104hybrids.. 193, 219, 220, 222, 223, 231, 232notes 193, 196, 197, 199, 200peduncle 78petiole 56, 59*PAGENymphaea lotus, phylogeny 218pollen 85root 28contractile 113hairsinstructure 30-34, 36seed 90, 92seedling leaves 110*, 112*stamen, development of 101-102stem structure 43-46stipules 55, 50stolon 48, 49tuber 40*used as food 196, 235varieties 193, 196, 197Nymphaea lotus 135, 198, 199, 205angusta 193lata 193monstrosa 243oblonga 193stuhlmannii 161o in iii ill in 194o lotus 193H cordlfolia 193y pubescens 193Nymphaea lotus X rubra 231-232X sturtevantii 193Nymphaea lucida 230lutea 14, 16, 18-22, 25, 163, 167, 211macrophylla 211maculata 141, 189madagascariensis 157early notices of 22seed .92, 93malabarica 23, 140mariae lagrangei 232marliacea albiia 230carnea 231chromatclla 219, 221, 230culture 236, 238floral movements ... 119idii ib lasts In petiole.. 62leaf 54, 69peduncle 78root structure ...30*, 31,33, 34flammea 231ignea 231rosea 231rubra punctata 231mauvii 139, 228maximilianl 180, 210Nymphaea mexicana (Plate XIII) ..163-167, 164*, 205, 217distribution 167, 213floral movements 119hybrids 221, 229, 230, 231leaf 54, 65, 67, 68, 72, 75*perennatlng body 41, 42*, 74, 75petiole 57, 59*, 61phylogeny 218pollen 85stolon 41(See also N. flava.)Nymphaea mexicana 131, 137, 139Nymphaea nilorantha (Plate IX) 148-147hybrids 22421
274 \ The Waterlilies.PAOBNymphaea mlcrantba, vlvlpary 74Nymphaea mierophylla 211milletU 172minima 211minor 172lutea 22Nymphaea mtnuta 210Nymphaea Urt. O. W. Ward 130, 221, 228*leaf 75*tuber 40naccheli 198Nymphaea nnliui 216Nymphaea ncglecta 172, 174nelumbo I 211nervosa t 130nlobe 232nitida 175, 211nocturna 201nouchali 198nubica '. . 146obtusiloba 164Nymphaea odorata 3, 179-182, 181*aril 91*. 104distribution 182, 213embryo 104-105*embryo-sac 102*, 103flower, duration of 119odor of 121submergence after antbesls,123, 124germination 107hybrids 223, 229, 233Introduction to cultivation 236leaf 54, 65-73, 71*. 181*development of (Plate II) ..95, 96megaspore 102notes 182, 192, 197ovule 102*peduncle 59*, 62, 78petals 83, 181*petiole 56-58, 60-62, 73*, 76*phyllotaxy 42phytogeny 218pistil 87-89, 181*pollen 85pollination 121, 122root 28hairsIllorigin of 36structure 31-36, 32*seed 90-94sepals 81, 82, 181*stem (Plate II) 38dissection of apex 95-97plaglotroplsm of 114structure 43*-46, 44*stipules (Plate II) 65*, 56Nymphaea odorata 211carolinensis 231caroliniana 222, 231chlorhiza 186emquisita 223Nymphaea odorata glgantea 186-188, 187*, 210culture 237flower 187*leaf 65FAQBNymphaea odorata gtgantea, leaf of seedling 110*petiole 59*phytogeny 218variants 230Nymphaea odorata glabra 188luciana 222, 231maxima 192Nymphaea odorata minor 183-186, 184*flower 78, 184*movements of 119leaf 54, 64petiole 01pistil 87, 184*root epidermis 30stem 38variants 230Nymphaea odorata minor floribus roaeU 186pumila 230Nymphaea odorata rosea 186flower 77hybrids 221, 222, 230leaf 72petiole 61variants 223, 230Nymphaea odorata rosea X mexlcana 229X tuberosa 231Nymphaea odorata rubra 186, 230suavissima 231eulfureo 221, 229petiole 59*phyllotaxy 43superba 231villosa 56, 188vUlossistima 188Nymphaea odorata x mexlcana 229Nymphaea officinalis 211omarana (Plate XXX) 193, 232fertility 223floral movements 119pedigree 233orbiculata 211ortgiesiana 194hybrids 220, 231, 232, 233oi lyicsiano rubra 232fertility 223history 220Nymphaea ovallfolla 160*-161oxypetala (Plate XXV) 210distribution 210, 213leaves 54phylogeny 218stamens 84Nymphaea parkcriana 186parvifldra 176, 177, 178parva 19, 22Nymphaea parvula 216Nymphaea passiflora 208pauciradiata 172, 174Pennsylvania (Plate XXVI) ..221, 222, 225*-226floral movements 110pentapetala 211pcrmixta 172prterslana 153planchonil 164, 205poceila14JNymphaea polyrhlza 216
Index.275PAGENymphaea polysepala 211pringlei 186pseudopygmaea 148Nymphaea pubescens (Plate XVII) 141, 198-199early notices of 23leaf 64notes 193, 197, 199seed 92Nymphaea pulchella 136, 210pulcherrima 146, 221, 222, 224*, 225pumila-211punctata 140, 172, 211pygmaea 168ovary 84stamens 84pygmaea alba 168, 223, 229helvola 229Queen Elizabeth 232hybrids 232-233pedigree 233Queen Elizabeth x caerulea 232X dentata 232reichardiana 211reniformis 186-189repanda 131rhodantha 141robinsoni 223, 230robinsoniana 230*rosacea 23(Jrosea 186, 199funiculus 104rotundifolia 176-178Nymphaea roussetl 216Nymphaea rubicunda 232Nymphaea rubra (Plate XVIII) 199color In relation to Insects 121culture 238, 238flower .77-79malformed 100movements of 119hybrids 193, 219-221, 223, 231, 232In Hindu literature 3-5leaf 63, 67, 68*-71, 73, 75-77absorption of water by 116development of . ..95, 96notes 193, 199, 200peduncle 78petiole 57, 58, 60, 73*, 76*growth of ./. 115phyllotaxy 43phylogeny 217, 218pollen r 85respiration 118seed 90, 92*stamens 83stolon 48, 49, 52Nymphaea rubra rosea 199floral movements 119leaf 65peduncle 78Nymphaea rubra X ortgiesiana 232rudgeana (Plate XX) 25, 135,136, 164, 200, 204-206, 222clelstogamy 27, 121floral movements 119, 120leaf 64PAGESNymphaea rudgeana, petiole 59*phylogeny 218seed 92stamens 84Nymphaea rudgeana p amazonum 200, 206rufescens 140, 147, 211sagittariaefolia 209sagittata 198, 199, 211sagittifolia 211sanguinea 186, 231scutifolia 141, 153, 156hybrids 233sei^norert 78, 230semiaperta 172, 174semisterilis 198, 199serrata 131sinuata 205smithiana 232speciosa 136sphaerocarpa 179rubra 179spiralis 186splendens 176-178Nymphaea stellata 140-141, 193, 217culture 236early notices of 3, 5, 23leaf 75, 77notes 141, 146phyllotaxy 42, 43phylogeny 218pollination 122respiration 118seed 92*synonymy 140, 211vivipary 74Nymphaea stellata 131, 141, 153bulbillifera 146Nymphaea stellata cyanea 140Nymphaea stellata eastoniensis 227macrantha 153parviftora 140Nymphaea stellata versicolor (Plate VII) 140-141Nymphaea stellata zanzibariensis 157stellata /S 140Nymphaea stellata X caerulea 225stenaspidota (Plate XXI, XXIV) 209stuhlmannii 161-162*flower 77, 162*leaf 162*Nymphaea sturtevantii 231-232leaf, hairs on 73peduncle 78petiole 57*suaveolens 211Nymphaea sulfurea (Plate XII) 161flower 77Nymphaea sulfurea 229grandiflora 229sumatrana 211Nymphaea tenerinervia (Plate XXI) 186, 210Nymphaea tenuinervia 210Nymphaea tctragona (Plate XIV) 3, 167-171,168*, 216, 217, 219culture 236-238distribution 170, 213, 214flower 78, 80, 168*
The Waterulies.MMNymphaea tetragona, flower, movements of 119odor of 121submergence after anthesls, 124fruit 90germination 106hybrids 223, 229, 231leaf 54, 64-66, 77of seedling 110*structure of 73, 76. .67*-69, 72,ovary > 84peduncle 69*petiole 56-60, 59*, 73*phylogeny ., 218pistil 86pollen i 85stamens 84, 168*stem 40used as food .\ . . 170varieties 170Nymphaea tetragona angusta j 170helvola 221, 229, 236, 237lata 170Nymphaea tetragona X alba 229X alba rubra 229X mexlcana 229Xymphaea tetragonanthos 168thermalis 194, 196, 197carpels 84distribution 214leaf 75, 77pbylogeny 214, 217seed 92stamens 84X rubra 232X scutttoUa 232trisepala 80, 135tropaeolifolla 136, 205Nymphaea tuberosa 189*-192carpels 84, 87, 88flower, movements of 119submergence of 124hybrids 222, 223, 230, 231leaf .54, 65, 67-69. 72, 76', 190*, 191*peduncle 59*, 78ldloblasts In 62petiole 56, 57, 59*-61ldloblasts In 62vascular system of 63*phyllotaxy 42phylogeny 218pollination 121, 122, 192root structure 30-35seed 90, 92sepals 81, 82, 189*stamens 84, 189*stem 38, 39*apex, dissection of 95hairs on 73*structure 44-46stigma 84*, 189*stolon structure 49stomata 192tubers 38, 190*, 192variants 230Nymphaea tuberosa flavescens 230maxima 189, 192PAGENymphaea tuberosa porta 189, 192stem 38plena 230richardtoni 230development of leaf . . 97rosea 231rubra 231superba 231Nymphaea tuberosa X odorata rosea 231Nymphaea tussilagifolia 186umbilicalls 211undulata 136union 230urceolata 176-178venusta 176-178versicolor 131, 140flower 78victoria 211violacea 130vivipara 74, 146wenzelU 172William Doogue 231William Falconer 231TFHam Stone (Plate XXIX) 139, 221, 222,227*228Nymphaea zanzibariensis 157, 217, 236culture 237embryo sac 103flower 77, 78, 80, 158*movements of ....119, 120funiculus 104germination 107hybrids 221, 222, 226, 227Introduction Into Europe. . . 10664-68structure 69, 70, 72\leaf> notes 141peduncle 78, 79ldloblasts In 62petiole 59*, 61Mi. .Wast s in 62vascular structure of . 63*phyllotaxy 42phylogeny 218pistil 86, 158*pollen 85root 28hairsIllseed 90>seedling leaves 110*structure 112stamens 83, 158*tuber 40*Nymphaea zanzibariensis rosea, leaves 75*Nymphaea zanzibariensisnzil. xtuber 40*227floral movements 119ovary 87, 88peduncle 78petiole 57, 58, 61sepals 81stipules 55*X capensls 227Nymphaea zenkerl197M98Nymphaeaceae, flower, symmetry of 101leaf, development of 97
PAGENymphaeaceae, roots, adventitious 28primary 28Nymphaeae apocarpiae (see Apocarpiae) 127hybrids 225-228syncarpiae (see Syncarpiae) 163hybrids 229-232Nymphaeinae, geological history 215phylogeny 218Nymphaeites arethusae 216brongniarttt 216charpentieri 216Nymphaeites lignitlea 216ludwigii 217mlcrorhlzus 216, 217palaeopygmaeus 217polyrhizus 218saxonica 217tener 217thulensis 217weberi 217Nymphea 125Odors of flowers 121Opening and closing of flowers ..11-13, 17, 21, 23, 119-121Ornlthogalurn 23Osiris 8Ovary 84-88, 86*development of 98-99morphology of 86-87, 98Ovules8Vdevelopment of 102*-104Pachycyst 70Padma 4, 5Papyrus 8, 9Parkinson^0, 23Parlatore on air-canals of petiole 68Paxton on vivlpary 74Payer 80on development of flower 97-99, 101leaff . . . 95Peduncle 78, 79air-canals of 59*, 78, 79apogeotropism of J . . . . 114development of t . . . . 99Idioblasts in 7PIso'.21Plstia 24Pistil. (See Carpels.)Plaglotroplsni of stems 114Planchon on Cyanea 86dead Insects In flowers 122hybrids 220, 223, 232malformed flower 100pneumatocyst 60viviparous leaves 74Planting 237-239Pleyte 6, 8Pliny 12, 13, 16-18, 23on disappearance of flowers at night 119Plukenet 23Plutarch 9Pneumatocyst 60Podophyllum 24Pollen 85*Pollination, arrangements for 121-123artificial 240Pond for culture 237, 238Prlchard 10Primary root 28, 108*, 109-111, 110*Principal area 66vein 66Proclus 9, 10Propagation 239Proterogyny 121Pseudostomata 72Quercus leaf 72Raciborski on inflorescence 99phyllotaxy 42-43Eadlkofer, suggestion on N. mexicana 167Ramses II 6-9Ranunculus sceleratus 115Ray, John 22Receptacle 79, 86*morphology of 98Respiration 117-118RImbach on contractile roots 113Robertson 188, 192on pollination 121, 122Rose 5Root 27-28air-canals in 31-33development of 37*, 38*apical growth 36-37structure 29*-30branching of 28cap 28^structure29-30contractile 28, 33*, 35, 113cortex of 31-34, 32*epidermis of 30*-31geotropism of 113hairs 109, 111idioblasts in 32-33development of 37structure 31-30, 32*Roots of seedlings 109
278 The Waterlii.ies.FAOIHoots, origin of 28, 36number of 28vascular system ofSB-SB*Rowan, Mrs > 131RUB80W on cuticle of air-canals 60Sagot ....64Sakoontala 4Sallx leaf 72Sapor ta > 216, 216Schenck ** 210on crystals 61N. thermalia >. 214Schllbersky 214Schilling 72, 73Schulz on pollination of N. alba 121, 122Schumann on Inflorescence 99Schwelnfurth ?\ . 6-9Sedum : 1Seed 90-94*, 92*. 93*as food 3, 10-12, 233collection of 239distribution of 27germination of 100-107, 109-110keeping of 239vitality of 106-107Seedlings 107-110*, 108*structure of 110-112Sepals 79, 81, 82, 89, 90air-canals In 81, 82development of 97-99morphology of 100-101Shedding of epidermis of root 30-31Siber on conditions for germination 106Simkovica on A', thermalia 214Simonkal on N. thermalia 214Sloane 24Soar, J. J 66Soil for culture 237, 238Solereder 60, 64Specific names, use of 136Stahl on mucilage hairs 73Stamens 83-85, 84*development of 98, 101, 102morphology of 101Starch in peduncles 78petioles 68roots 34seed 93, 94stems 46, 113stipules 56Staub on N. thermalia 214Steles 36, 49, 52, 53Stem 27, 37-42, 39*apex, dissection of 95development of tissues In 97geotroplsm of 114hairs on 73*starch stored In 113structure 43*-48, 44*, 46*of seedling 112vascular system of . . .49-54, 50*Stigma 84*, 88Stipules 54-55*development of 96structure of 55, 56PAGEStolons of Lotos group 41, 48, 49, 52*, 53N. flavo 48, 49, 63N. flavo-vlrens 41N. mexlcana 41N. tuberosa 49Stomata 64, 67, bo, 77, 192development of 97on back of leaf 185seedling leaves 112*sepals 81stamens 84, 85Sturtevant 182Submergence of flower after anthesls 123-124Suspensor 104, 105*Symmetry of flower 100, 101Bymphytopleara 87Syncarplae (aee Nymphaeae syncarplae) 163ovary 86*phylogeny 217, 218pistil 86, 87, 89stamens 84Tank for culture 236-2^8TassI on anesthesia of flowers 121Tavera 235Tetrasepaleae, phylogeny 218Theophrastus 11, 16, 20, 23Thyll-llke bodies 61, 02, 77, 79in sepals 81Thylosis 48*Tlttmann on seedling 107Tournefort 24, 25Tragus 14, 16Transpiration 116-117Tratlnnlck 5, 6Treat, Mrs. Mary, on N. /?aeo 40Trecuf 37, 49Treviranus on seedling 107Tricker,' Wm 221on germination 106, 107culture 237, 238, 240, 241Tricntalls 24Trisepaleae, phylogeny 218Tubers 40*, 47, 190*as food 3, 7, 10, 17, 22, 235drawn down by contractile roots 113early leaves from 75*of seedling 108*, 110Turner, Wm 14Unger .... 1 oVaillant 25Vallisnerla spiralis 124Van Houtte 74Van Rheede 23Van Tleghem on crystals 61root cap 29Varietal names, use of 13cVascular system of leaf 70, 71*node 51*, 53, 54peduncle 53, 54, 59*, 63, 79petiole 59*, 62-64, 63*, 76-77root 35-36*seedling ill, 112stem 49-54, 50*
fPAOBVenation of leaves 66, 67, 115Vernation of leaves 66, 07Vesling 20, 23Victoria 211, 215, 216heat evolved in flower 123inflorescence 99introduction of 236leaf 65, 67, 77phytogeny 218pollination 122root hairs absent in Illseed 93stem structure 46Victoria cruziana as food 235Victoria fltzroyana 128Victoria regla 232introduction of seeds into England .... 100leaf 69Vitality of seeds 106, 107Vivipary 74Wachter 75on effect of removing roots and leaves 116on stomata of water-leaves 77, 112Waldsteln and Kitaibel 214
AQKN97C7cop. 2Conard, Henry ShoemakerThe waterliliesBotany
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