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Sir W. JARDINE, Bart.— P. J. SELBY, Esq.,


DAVID DON, Esq., Prof. Bot. King’s Coll. Lond.,










Oinnes res creatse sunt divinae sapientiae et potentiae testes, divitiae felicitatis humanae : exharum usu honitas Creatoris ; ex pulchritudine sapientia Domini; ex teconomia in conservatione, proportione, renovatione^ potentia majestatis elucet. Earum itaque indagatio ab hominibus sibi relictis semper aestimata ; a vere eruditis et sapientibus semper exculta ; male doctis et barbaris semper inimica fuit.” Linn.








XLIV. On Gloionema paradoxum. By the Rev. M. J. Berkeley, M.A., F.L.S 449

XLV, Supplement to descriptions of Exotic Fungi in Annals of Nat. Hist.,’ vol. iii. pp. 322 and 375. By the Rev. M, J; Berkeley,

M.A., F.L.S,


XLVI. A List of Plants collected by Charles Fellows, Esq., during his Tour in Lycia and Caria ; with descriptions of the New Species.

By David Don, Esq., Prof. Bot. King’s College 45 4

XLVI I. Report of the Results of Researches in Physiological Botany made in the year 1839. By the late F. J. Meyen, M.D., Professor of Botany in the University of Berlin 460

XLVIII. Descriptions of new or little known Arachnida. By Mr. Adam White, Assistant in the Zoological Department of the British Museum 471

XLIX. Additions to the Fauna of Ireland. By Wm. Thompson,

Esq., Vice-Pres. Natural History Society of Belfast 477

L. Description of two new Genera of Irish Zoophytes. By Arthur Hill Hassall, Esq., Corresponding Member of the Natural History Society of Dublin 483

LI. Notes on Birds. By T. C. Eyton, Esq., F.L.S 486^

LII. A Catalogue of Fossil Fish in the Collections of the Earl of Enniskillen, F.G.S., &c., and Sir Philip Grey Egerton, Bart., F.R.S., &c 487

LI 1 1. Information respecting Zoological and Botanical Travellers :

Mr. Forbes and Mr. Thompson. Mr. Schomburgk. Mr. W. S. Mac- Leay 520

New BooTcs : Natural History as a Branch of General Education,

by Robert Patterson, Esq 498

Proceedings of the Zoological Society ; Geological Society 503—519

New Genus of Mexican Glirine Mammalia ; On a new European Ge- nus of Freshwater Fish, by F. Heckel ; Forbes’s Starfishes, Echi- nus Uvidus', Speaking Canary Bird; Birds of Kent; Emberiza Hortulana; Diluvial {^Glacial}'] Scratches on Rocks in America;

Mr. Shuckard on the Proceedings of the Entomological Society; Obituary C. S. Rafinesque; Meteorological Observations and Table 521—528


Proceedings of the Royal Society ; Linnaean Society ; Entomological

Society; Zoological Society 529 578

Index 579

Irish Zoophytes.


Plate I. British Actiniadse.

{Development of Stomata in Hyacinthus orientalis.

Fig. 1, 2, 3. Cyanea coccinea ; (from Tenby.)

Fig. 4, 5. Helix conoidea; (fossil from Essex.)

III. Horny Sponge from Australia.

IV. Nest of Myrapetra scutellaris, a South American Wasp.

V. Horns of various species of Sheep.

VI. Flustra carnosa * ; Hermia glandulosa ; Sertularia Margarita; Sertularia pumila.

VII. Thuiaria articulata.

VIII. Plumularia frutescens ; Valkeria imbricata; Langenella repens ? Valkeria (new species) ; Hippothoa lanceolata.

IX. Cellepora bimucronata ; Lepralia ciliata, appensa, pedilostoma, insignis, cylindrica, punctata, linearis.

X. Tubulipora lobulata ; Alcyonidium hirsutum ; Echinochorium clavigerum.

XL Metamorphoses of the Spur-shaped Nectaries of Aquilegia vul- garis.

XII. Urari Plant, Strychnos toxifera, Schomb. r Fruit of the Urari Plant.

* \ Gloionema paradoxum ; fig. a. (1 8).

* In Plate VI. this figure is by mistake given as Coryne squamata. It is, in reality, a representation of Flustra carnosa, Johnston (see p. 369), of which it is a correct figure, with the exception of the number of the tenta- cula, which should have been 30.

Erratum, p. 3/3, line 33, for Coryne squamata read Flustra carnosa.




I. Considerations respecting Spur-shaped Nectaries, and those of tlie Aquilegia vulgaris in particular. By M. Ch. Morren, Professor in ordinary at the University of Liege, Member of the Royal Academy of

Brussels, &c. (With a Plate.) 1

II. On the Fungi of the Neighbourhood of Bristol. By Mr. H. O. Stephens 17

III. Description of Four Bats taken in Cuba. By Dr. Grundlach 19

IV. Horae Zoologicae. By Sir W. Jardine, Bart. :

Remarks on the Structure and Habits of Le'pidosiren annec- tens 21

V. Commentary on Mr. G. R. Gray’s Genera of Birds,’ 1S40. By

H. E. Strickland, Esq., M.A., F.G.S., &c 26

VI. Mr. Shuckard on his falsely alleged participation in Mr. Swain-

son’s views of Natural Arrangement 41

VII. Excerpta Botanica, or abridged Extracts translated from the

Foreign Journals, illustrative of, or connected with, the Botany of Great Britain. By W. A. Leighton, Esq., B.A., F.B.S.E., &c.

On the Anther of Chara vulgaris and Chara hispida, and the Animalcules contained in it. By M. Gustavus Thuret... 44 VIII. Notes on Saxifraga umhrosa: By Charles C. Babington, M.A., F.L.S., F.G.S., &c. ; and by the Reviewer of Baines’s ‘York-

shire Flora’ 47

IX. Notes on Birds. By T. C. Eyton, Esq., F.L.S. : Menura Lyra 48

New Books : An Introduction to the modern Classification of Insects, founded on the Natural Habits and corresponding Organization of the dilferent Families, by J. O. Westwood, F.L.S. , &c. Otia Hispanica, Auctore P. B. Webb, No. 2 53 58

Proceedings of the Zoological Society ; Geological Society ; Microsco- pical Society 58 74

Absorption of Liquid Solutions by the Sap-vessels of Plants ; Congres Scientifique de France ; Mr. Thompson on Eels killed by the late Frost ; Obituary Francis Bauer, Esq. ; Mr. Paget on Halichcerus Gryphus ; Meteorological Observations and Table 74 80





X. Contributions to British Actinology. By Edward Forbes,

M.W.S., For. Sec. B.S., &c. (With a Plate.) 81

Xr. Description of some new Species and four new Genera of Rep- tiles from Western Australia, discovered by John Gould, Esq. By J.

E. Gray, Esq., F.R.S., &c. 86

XII. Description of some new Species of Madeiran Fishes, with ad- ditional information relating to those already described. By the Rev.

R. T. Lowe, M.A 92

XIII. On the Species of Stickleback (Gasterostens, Linn.) found in Ireland. By Wm. Thompson, Vice-Pres. Nat. Hist. Society of Belfast 95

XIV. Notice of Plants and Animals found in the Sulphureous

Waters of Harrowgate and Askern, Yorkshire. By E. Lankester, M.D., F.L.S., &c 105

XV. Remarks upon the Recent and Fossil Cycadese. By J. Morris,

Esq no

XVI. Carabideous Insects collected by Charles Darwin, Esq., during the Voyage of Her Majesty’s Ship Beagle. By G. R. Waterhouse,

Esq 120

XVII. Observations on a Keratose Sponge from Australia. By J.

S. Bowerbank, Esq, F.G.S. (With a Plate.) 129

XVIII. Notices of European Herbaria, particularly those most in- teresting to the North American Botanist 132

New Books : Monographic des Libellulidees d’Europe, par Edm. De

Selys Longchamps, Membre de plusieurs Societes savantes 141

Proceedings of the Entomological Society ; Zoological Society ; Royal Botanical Society of Edinburgh; Wernerian Natural History So- ciety of Edinburgh 143 159

Mr. Gray’s Genera of Birds Birds of Kent ; Meteorological Observa- tions and Table 159 160


XIX. On the Alteration which the Atmosphere undergoes during

the Development of Heat in the Spadix of Colocasia odora. By Pro- fessors G. Vrolik and W. H. De Vriese 161

XX. Note on the Occurrence of the Genus Diphya on the Coast of

Ireland. By G. C. Hyndman, Esq., Member of the Natural History Society of Belfast 164

XXL Report of the Results of Researches in Physiological Botany made in the year 1839. By the late F. J. Meyen, M.D., Professor of Botany in the University of Berlin 166

XXII. Notes on Birds. By T. C. Eyton, Esq., F.L.S 177

XXIII. Notices of European Herbaria, particularly those most in- teresting to the North American Botanist 179



XXIV. Excerpta Botanica, or abridged Extracts translated from the Foreign Jounials, illustrative of, or connected with, the Botany of Great Britain. By W. A. Leighton, Esq., B.A., F.B.S.E., &c.

On the Development of the Reproductive Organs of the Mis- seltoe {Viscum album, Linn,). By M. Decaisne 185

XXV. On the Origin of some of the Lower Forms of Vegetation.

By Mr. Henry Oxley Stephens 190

XXVI. An Amended List of the Species of the Genus Ovis. By

Edward Blyth, Esq 195

XXVII. Addenda to the Flora of Norfolk. By Mr. S. P. Woodward 201 XXVIII. On the Formation of the Stomata. By Hugo Mohl.

(With a Plate.) 206

New Books: The Natural History of South Devon, by J. C. Bellamy,

Esq. Memorie della Reale Accademia delle Scienze di To- rino 209 211

Proceedings of the Microscopical Society ; Botanical Societ}' of Edin- burgh ; Linnaean Society ; Geological Society ; Zoological So- ciety 211—234

Zoological Observations at Tenby, by Dr. Davis (with a Plate) ; Mr. Jennings on Eels killed by Frost; On the Occurrence oi Anemone ranunculoides, by the Rev. W. Hincks ; On the Irish localities for Dia7ithiis plumarius, by W. T. Alexander, Esq. ; Suicidal Powers of Luidia ; Meteorological Observations and Table 234 240


XXIX. On the Anatomy of Nautilus. By M. Valenciennes 241

XXX. Remarks on Red and Green Snow. By the late Prof. Meyen 245

XXXI. An Amended List of the Species of the Genus Ovis. By

Edward Blyth, Esq 248

XXXII. Notes on some of the smaller British Mammalia, including the Description of a New Species of Arvicola, found in Scotland. By

the Rev. Leonard Jenyns, M.A., F.L.S,, &c 261

XXXI II. Supplement to a Catalogue of Irish Zoophytes. By Arthur Hill Hassall, Esq. Read before the Natural History So- ciety of Dublin, November 6th, 1840 276

XXXIV. List of Phanerogamous Plants, together with the Crypto- gamic Orders Filices, Equisetacece, and Lycopodiacece, observed in the

Shetland Islands. By Thomas Edmondston, Jun., Esq 287

XXXV. On the Composition of Chalk Rocks and Chalk Marl by in- visible Organic Bodies : from the Observations of Dr. Ehrenberg. By

Thomas Weaver, Esq., F.R.S., F.G.S., M.R.I.A., &c. &c 296

XXXVI. Description of a South American Wasp which collects Honey. By Mr. Adam White, M.E.S., Assistant in the Zoological Department of the British Museum. (With a Plate.) 315




XXXVII. Information respecting Zoological and Botanical Tra- vellers : Mr. Schomburgk. Letter from Messrs. Forbes and Thomp- son, with Captain Graves, in tbe Archipelago 348

New Books : The Principles of Botany, by W. Hughes Willshire, M.D. Arcana Entomologica, or Illustrations of new, rare, and interesting Exotic Insects, by J. O. Westwood, F.L.S., &c. ; Na- turhistorisk Tidskrift : edited by Henrik Krbyer, Copenhagen. Tijdschrift voor Natuurlijke Geschiedenis en Physiologie, by Pro- fessors Van der Hoeven and De Vriese. Part VI. Leyden . 322 326

Proceedings of the Geological Society; Zoological Society; Werne-

rian Society; Royal Society of Edinburgh 326 348

Mr. Yarrell on Motacilla alba of Linnaeus; Mr. Johnson on Chcp.tura

rujicollis ; Meteorological Observations and Table 350 352


XXXVIII. On the Existence of Branchiae in the young C cb cilice \ and on a Modification and Extension of the Branchial Classification of

the Amphibia. By John Hogg, Esq., M.A., F.R.S., &c 353

XXXIX. Supplement to a Catalogue of Irish Zoophytes. By Arthur Hill Hassall, Esq. Read before the Natural History So- ciety of Dublin, November 6th, 1840. (With Plates) 363

XL. On the Composition of Chalk Rocks and Chalk Marl by invi- sible Organic Bodies : from the Observations of Dr. Ehrenberg. By Thomas Weaver, Esq., F.R.S., F.G.S., M.R.I.A., &c. &c 374

XLI. Report of the Results of Researches in Physiological Botany

made in the year 1839. By the late F. J. Meyen, M.D., Professor of

Botany in the University of Berlin 399

XLII. On the Urari, the Arrow Poison of the Indians of Guiana ; with a description of the Plant from which it is extracted. By Robert H. Schomburgk, Esq 407

XLIII. A List of the Fossil Shells found in a Fluvio-Marine Deposit

at Clacton in Essex. By John Brown, Esq., F.G.S 427

New Books : On the Relation between the Holy Scriptures and some parts of Geological Science, by J. Pye Smith, D.D. The Cer- tainties of Geology, by W. Sidney Gibson, F.G.S. Linnaea, ein Journal fiir die Botanik, &c. leones Fungorum hucusque cogni- torum 429 434

Proceedings of the Entomological Society ; Linnaean Society ... 434 444

Third Meeting of the Men of Science of Italy ; Dr. Lush on the Madi, or Chili Oil-seed, Madia saliva ; Diurnea Novembris, or Novem- ber Dagger ; Meteorological Observations and Table 445 418




No. 41. MARCH 1841.

I. Considerations respecting Spur-shaped Nectaries, and those of the Aquilegia vulgaris in particular. By M. Cii. Mor- REN, Professor in ordinary at the University of Liege, Member of the Royal Academy of Brussels, &c.*

[With a Plate.]

The Columbine, that pretty ranunculaceous flower of our woods, deserves attention, as well on account of its structure, curious as it is, and, we venture to add, but little known, as from the historical recollections which it brings to mind. To say nothing here of the medicinal virtues which Dioscorides attributed to his Isopyron or to his Phasiolon, a plant which Fabius Columna, Clusius, Dodonseus and many other learned botanists suppose to be no other than the Columbine itself ; and not to mention Adrian Junius, who also quotes it as a medical plant ; or Francois Rapard, a celebrated physician of Bruges, who addressed to Clusius a letter upon its uses in difficult labours ; ought we not to remark that its singular nec- taries, compared by some to the beak and talons of an eagle, by others to the graceful neck of the pigeon, by some to rams’-horns, and by others to capuchins’ hoods, had so gained the attention of the painters of the middle ages, that it be- came one of the favourite flowers, placed in great profusion in the illuminations of missals and manuscripts of the time ? The ^ ancoiles^ or the ^ ancolyes^ were there intermixed with the leaves, flowers, or fruit of the strawberry of of the campanula ; and Memling was most particularly attached to it. When Dodoens wrote his CruptJt-BlWk,^’ the name Aquileia or Aquilina was still a novelty introduced, he says, by the latest phytographers of his own time. The name Aquilegia

* Translated from the original communicated by the Author.

Ann, ^ Mag. N. Hist. Vol. vii. B


M. Morren on the Spur-shaped Nectaries

which he gives it calls to mind the comparison already men- tioned, of the beak and talons of an eagle; but since that time that of Columbine prevailed in England and in Holland, where they were fonder of likening the spurred petal of this flower to the stately neck of a pigeon. When one of these flowers is turned upside-down, says an English author * * * §, we might fancy we saw a group of young eagles, or, if we like better, a nest of pigeons. It is evident that these spur-shaped necta- ries had considerable influence on the mind of Dodoens, since on their account he places his Akeleyen {^ancolies/ columbines,) between the Cypripedium Calceolus, a monocotyledoiious plant, and his Grant or Antirrhinum majus, an alliance which would not at all square with our present ideas of classi- fication by families. By a singular chance, the Cypripedium, which in our mythological language we call the Venus^s foot, was at that period of religious struggles called Our Lady^s shoe {Calceolus Marm), and the Columbine w^as named in its turn the Virginias glove : thus we see that shoes and gloves shook hands in our Lady’s toilet.

Chief ornament of the gardens of the middle ages, and even of those of the sixteenth century, the Aquilegia, carefully and almost universally cultivated, produced those varieties which modern botanists have pointed out in this species. Joost van Ravelingen, the commentator of Dodoens, and L’Obelt, men- tion the varieties in colour and those of form to be met with in their time : blue, red, flesh-coloured, blue and white, white and variegated. The garden of a gentleman of Leyden, Jean van Hogelande, produced an Aquilegia pleno fore roseo, v hich Clusius described and illustrated. The same botanist had also recorded a variety truly monstrous {Aquilegia degener), in that the petals, being reduced to their primitive type of leaves, had remained green ; only Van Ravelingen did not think that such monstrosities were worth the pains of describing. Now- a-days we should be eager to do it, and not without reason ; for the organography of such a flower is very interesting to know, in relation to the subject which vvill occupy us further on.

The varieties of structure known under the name of Aqui- legia vulgaris corniculata, in which Biria J and DeCandolle § discovered that the cornets are deviations of the anther,

* Burnett’s Outlines of Botany, p. 840.

f Generally written Lobel ; but the true name of the author of the Stir- pium Histor'ia was Matthias De L’Obel, as appears from his letters and the portrait engraved during his life-time.

t Biria. Monographie des Renonculacees. Montpellier, 1811.

§ DeCandolle. Organographie, tom. i. p. 496.

of the Aquilegia vulgaris. 3

and those which bear the name of Aquilegia vulgaris ecal- carata or stellata, in which^ according to these authors, the petals proceed from modified staminal filaments without an- thers— these varieties had been already described by Clusius, Dodoens and L’Obel*. Moreover, these authors, besides the simple, semi-double and double varieties of these two principal forms (true sub-varieties which we still possess), also mention Columbines with inverted flowers [Aquileia flora inverso variegato). We might suppose, from the Dodonaean context, that it was hereby understood that the flowers, in- stead of being pendent, were upright nift 011 htt-

0cljiUen ban be anbet ban bat be lUoemen abevecljt^

0taen.” But we know that in this variety, the bases of the spurs being twisted, the spur itself has an upward direction.

We cannot but take an interest in observing the pleasures of the horticulturists of those times. Now-a-days these Co- lumbines are treated with disregard, and dismissed as fit only for the garden of the cottager or village Cure, or, at most, are only permitted to grow" in the shade of some forgotten nook of our villas; but let us not say too much : fashion, which revives all antiquated things, may some day assert its claim upon these Columbines of the Castels. Already in the Botanic Gar- den at Brussels, we have seen pretty borders entirely filled w ith this plant of the middle ages.

The Columbine is really a ver}^ interesting flow^er, on ac- count of its nectaries ; and their genesis not being know n, at least so far as we are aware, we have taken them as the prin- cipal subject of our researches : our object has been to study them comparatively in the different varieties of the common species, and in some other species which w e had at hand ; se- condly, to observe the monstrous structures ; thirdly, to take them at their different degrees of evolution, in order to esta- blish their true genesis ; and, lastly, to examine their histo- logy, so as to ascertain how in their forms so varied the in- terior tissues were affected.

So long as the laws of metamorphoses had not acquired the right of citizenship in the domains of science, calcariform nectaries had to be considered as special pieces, born ad hoc, and being such by their proper nature, without an anterior nature, without a type from which they were derived. Although Linnaeus had said, ^‘Si nectarium a pet alis distinctum, com- muniter ludit'f he also said, distincta esse nectaria a corolla constat eocemplis : Aconitum, Aquilegia f &c.f. They were, then, the floral pieces whose secretion of honey determined

* Dodoens. Cruydt-Boek, 1644, p. 274. f Linnsei Philosophia Botanica : Fructificatio i. 110.

B 2

4 M. Morren on the Sjmr-shaped Nectaries

their character^ and that character raised them to the rank of organs sui generis, not proceeding from any other : they were, because they were.

They were, however, not nectaries, because by their nature they were stamens : here is that truth which science had not then become possessed of.

But when, at the end of the last century, Goethe, following the example of Wolff, established his celebrated theory of the metamorphosis of plants, the nectaries at once lost their auto- chthonous nature ; they were no longer aboriginal organs. On the contrary, in this new theory the nectaries became essen- tially organs of transition, mere forms of anterior organs ; they were, in short, intermediary organs of passage between the pe- tals and the stamens^. In the spirit of this philosophic me- thod, it was necessary to understand, that in order for the petal to become a stamen, in an ascending metamorphosis, it must previously pass through the form of a nectary. More- over, Goethe, who took precisely the Columbine as the exam- ple of one of the most remarkable and most striking trans- formations, considered, as he says, the cuculliform nectaries of this flower as a derivation from the petalsf. We shall see, on the contrary, that the progress of nature is a descending metamor- phosis ; that is to say, that the nectary is, in its genesis, a sta- men, and subsidiarily, that a stamen being developed as such, it may afterwards turn into a nectary.

The theory of Goethe had made too little impression in France to admit of the supposition, that in 1815 Mirbel set out from it when he regarded the nectaries of the Columbine, as well as all organs of the same kind, as anomalous forms of the parts of the perianthium. The spur-shaped cornets of the Aquilegia were also, in his eyes, forms of petals ; but the ano- maly attacking all the petals at once, the flower remained re- gular J. It was one of the successive alterations of types, and in the Columbine particularly this alteration was created in order to become an organ of secretion. A glandular lamina existed for this purpose at the bottom of the cornet-shaped petals §. The petal was the type.

This lamina we have never found ; and in the Aquilegia glan- dulosa, the Aquilegia atrata, &c., we have seen that there only exist one, or two, or three cornets without the regularity of the flower being perverted, as is the case in the Nasturtium,

* Goethe. CEuvres d’Histoire Naturelle. Edition de Martins et Turpin. Paris, 1837, p. 22G.

+ Ibid, p. 228, chap. 56.

J Mirbel. Elemens de Physiologie, vol. i. p. 269.

§ Goethe. CEuvres d’Histoire Naturelle. Edition de Martins.


of the Aquilegia vulgaris.

Tropceolum, or the Lark^s-spur. The great German poet’s notions had not indeed at first all possible success in this country. Willdenow always asserted that the spur {calcar) was more an organ intended to preserve the nectar than to prepare it, and that it was furthermore a sacciform elongation of the corolline corona* * * §. The first of these facts is evidently erroneous. The second was also admitted by Jacquin.

Sprengel, when opposing Vaillant, who had also himself declared that the nectary was always a production of the corolla, placed the spurs of the Columbine in his class of Nec- tarothecce, and characterized by the presence of the secreting gland at the bottom of the cornet. Moreover, it never occurred to his mind to investigate the anterior nature of this appara- tus in the Passiflorece, in the Aconites, and a multitude of other plants ; he sees only peculiar little machines^ more or less ornamented : machinulce peculiares eleganter co-

lor alee

DeCandolle, in 1819, adopts this view of the subject; but the spur, according to him, is of a very different nature, an elongation, one while of the calyx, one while of the corolla, one while of the perigonium ; but the stamens are still ex- cluded from the floral organs which may produce this nec- tary J. However, a year before, the celebrated botanist of Ge- neva had positively declared that, in Aquilegiae corniculatce, without regard to species, the supplementary spurs arose from a modification of the anthers which lengthened downwards ; moreover, he recognises the origin of the stellated varieties from the abortion of the anthers, and from the hypertrophy of the filaments ; and lastly, that the scales which are situated between the carpels and the stamens are stamens without anthers, and with dilated and membranous filaments§. Biria had made known the former facts |1. In 1827 these ideas were again brought forward in the Organographie vegetate^. They are, undeniably, the most accordant to the real state of things.

Among the most recent authors we may mention Kurr, who places the spurs of the Columbine with his nectarostig- mata, A very curious remark of this accurate writer is, that the greenish glands which secrete the nectar at the bottom of

* Willdenow. Grundriss der Krauterkunde, cap. 86-88. (Terminologie.)

t Linnaei Phil. Bot. edit. Sprengel (notes). Fructificatio ilO.

I DeCandolle. 7'lieorie elementaire, p. 406, § 395.

§ DeCandolle. Systema Regni Vegetabilis, vol. i. p. 333.

II Biria. Histoirenaturelle etmedicale desRenoncules, 1 fasc. Montpellier^ 1811.

^ DeCandolle. Organographie, vol. i. pp. 484 496. ,


M, Morren on the Spur-shaped Nectaries

the spurred cornet, do not begin to afford this sugared liquid until precisely when the first anther blows. The secretion lasts only as long as the stamens are capable of performing their functions, and at the end of three or four days the flower leaves off this ejection of fluid and of pollen, and drops the organs which produced both the one and the other*. This curious remark is quite correct; we have verified it. From this we might be led to suppose that the secretion of the nectar, which is here so intimately connected with the functions of the stamens, becomes necessary to the action of the sexes ; but from ten unblown flowers, where there had been neither dehiscence of the anthers, nor secretion of nectar by the spurs, Kurr cut away those organs : the further de- velopment took place without any difference, and these flowers bore as many and as large fruits as they ordinarily do ; the seeds germinated as usual f. This experiment gives great support to those who consider the nectar as being only a true excretion, comparable to our urine, and which is of no use, at least in the great majority of cases, in the process of fecun- dation, as was generally supposed. Kurr, however, does not give his opinion as to the proper nature of the spurs.

Lindley, in his new edition of the ^ Introduction to Botany,^ (1839) no longer gives (to the great regret of the friends of deep scientific research) the interesting and useful part on mor- phology; but this judicious author, in his edition of 1832 J, had published some very curious details upon the Aquilegia vulgaris, The petals of this planV’ says he, consist of a long, sessile, purple horn or bag, with a spreading margin, while the stamens consist of a slender filament, bearing a small, oblong, 2- celled, yellow anther. In single and regularly- formed flowers, nothing can be more unlike than the petals and stamens; but in double flowers the transition is complete. The petals which first begin to change, provide themselves with slender ungues : the next contract their margin, and acquire a still longer unguis : in the next the purple margin disappears entirely; two yellow lobes like the cells of the anther take its place, and the horn, diminished in size, no longer proceeds from the base, as in the genuine petal, but from the apex of the now filiform unguis : in the last transi- tion the lobes of the anther are more fully formed, and the horn is almost contracted within the dimensions of the con- nective, retaining, however,- its purple colour : the next stage

* Kiut. Untersucliungen iiber die Bedeutung, &c. Stutgard, 1833.

t Ibid, p. 128.

% Lindley. Introduction to BotiUiy (1832), p. 515. [Ed. 1835, p. 53G.]


of the Aquilegia vulgaris,

is the perfect stamen. No further evidence/’ says our author,

can, I think, be required of the formation of stamens out of petals.”

We see that Lindley had here followed the impulse given by Goethe, and that he looked upon the cuculliform petals (Richard) as proceeding towards the formation of the stamina by an ascending metamorphosis. At present the spur is no longer in his view anything but a modified petal* * * §. A dis- covery which we cannot dispute with him, since the germ of it appears in his words, is that the horn of the Columbine is really a lengthened connective, a thing which we shall also establish by direct proofs hereafter.

Although G. W. Bischoff, Professor of Botany at Heidel- berg, does not give this morphological genesis of the spur in the Aquilegia, still this author helps to lead us to believe that this is really the means which nature employs, in what he has remarked respecting the metamorphosis of the nectar - bearing horns of the Hellehorus fcetidus into normal stamensf. Link sees nothing in the spur but a continuation of the petal, characterized by the presence, at the end of its cavity, of a cellular gland, but of which the cellules have walls thicker than ordinary, a thing which we take the liberty of not admitting^. After M. Vogel of Bonn had sent me his elegant memoir on the development of the parts of the flow^er in the Leguminosae§, the study of the formation of calcariform or cuculliform nectaries, according to the glosso- logy of Richard II, became still more interesting. Indeed, Schleiden and Vogel having proved, by their labours, that it is not merely in idea, as a mental abstraction, that we are to see in the floral organs nothing but the axis of the plant and its leaves, but that this axis and its green leaves are really and substantially found, placed regularly in the very small buds, we thought that the investigation of the genesis of the nectaries in the Columbine could not be without scientific interest. DeCandolle came to consider these horn-shaped nectaries as anthers, by comparison; Lindley came to the same conclusion by the observation of teratological cases ; it was become therefore curious to test these view^s a priori and a posteriori by organogenic proofs : and this is what we have proposed to ourselves.

* Lindley. Introduction to Botany (1839), p. 169.

f Gottlob Wilhelm Bischoff. Lehrbuch der Botanik, vol. i. p. 404. (1833.)

J Link. Elementa Philosophise Botanicse, vol. ii. p. 130.

§ Schleiden und Vogel. Beitrage zur Entwickelungsgeschichte der Blu- mentheile bei den Leguminosen. (Act. Nat. Curios, vol. xix. p. 1.)

II Richard. Nouv. Elemens de Botanique, 1838, p. 333.

S M. Morren on the Spur-shaped Nectaries

Let us see, first, what takes place in a flower of Acpxilegia vulgaris calcarata.

1. Metamorphosis of the stamen into a spur-shaped nectary.

The stamen of the Columbine has a thread-shaped filament slender, flexible and yellow, and a two-celled anther with pa- rallel cells, slightly swelled, opening with a slit and united by a narrow connective, the whole yellow. The connective is even hardly visible (Plate XL fig. 1).

In many flowers, we find along the spire which leads in- sensibly from the andrcEceum to the corolla, stamens which turn into nectaries. To bring this about, the filament en- larges at its base; the connective is, at the opposite pole, the other organ which becomes modified, and it is even the most active of all in this transformation. It enlarges above, by separating the two loculi of the anther, and it grows to a point. This is not slow in becoming bifurcate, so that the connective is soon bilobate. (Figs. 2 and 7*)

To this modification, which up to this period does not at- tack the regularity of the organ, two ways of transformation succeed. In the one, one of the loculi of the anther disap- pears, in the other it remains visible with its fellow. The first of these modifications might induce a belief that the spur is a sac formed by one half of an anther or by a loculus, but this genesis is but a deceitful appearance. The second way of transformation proves that the spur is a sac-shaped con- nective, and that the two lobes of its limb represent the two loculi of the anther originally united by this same connective.

If such a spur-shaped nectary can be obtained, as from its nectar-secreting gland is truly a nectary, and that in it the two lobes of its limb exist as an elongation of the two an- ther-cells, still visible enough to attest their presence, it is clear that this second way of transformation should be ad- mitted. Now this is precisely what experience confirms. In the Columbine w e find this form, not so frequently as the first, it is true ; but it is found, and that is all that is neces- sary. This case we have delineated (Plate VII. fig. 6). On a stamen thus modified, besides a w ell-formed filament (c li), we find the tw^o loculi of the anther, still bearing pollen, but which open more widely [g b), separated by a small connec- tival eminence (c). Each of the swellings which represent the anther-cells produces an elongation in form of a thick margin (e), which, reaching from the inside to the outside, goes to form the circumference of the two lobes of the cornet (d g), separated by a slit {f). Each lobe corresponds to a cell, and originates from it ; it is only indeed that same cell length-


of the Aquilegia vulgaris.

ened. Lastly, i represents the belly of the cornet, and at k we find the gland which, for its part, secretes whilst the pollen no longer issues from the loculi, and little by little its struc- ture is annihilated.

This case of metamorphosis not only proves, as we said above, that the two lobes of the cornet of the Columbine are derivations from the cells of the anther, but it puts out of doubt that the tube of the cornet is the lengthened connective. A circumstance which we must not lose sight of in this phi- losophic study of a metamorphosed flower, is that the nectar- bearing gland, an organ of emission, and which rids the flower of its excess of carbon, is found at the opposite pole to the pollen-bearing loculi, other organs of emission whieh also excrete from the individual, but in this case for the pre- servation of the species, a substance eminently charged with carbon. At the two poles then the same function exists, but the one does not begin till the other ceases ; that is to say, the nectarial gland does not exist or become developed until the pollen apparatus wastes away and becomes obsolete. This subject certainly merits a reflection ; even should I be aeeused of seeing, in Botany, more than my own eyes can see, and especially should I be accused of allowing to myself, in a science of observation, some stretch of imagination. For my part, I could never comprehend how inquiry into the truths of nature should put aside the understanding, and reduce it to a state of inaetion which would render it useless. Be- hind and above facts I always conceive something superior and anterior; for facts are effects, and it is to the know- ledge of causes that w^e ought to endeavour to come. Now here, in the particular problem which occupies us, I see a verification of the law of organic compensation and a realiza- tion of the unity of composition. Thus, the nectaries are one with the stamens, the stamens one with the leaves, the leaves ONE with themselves, as autochthonous organs. So much for the law of unity. Moreover, the gland is at the end of the nectary, because, by its nature a stamen, the pollen is at the other end', there is a change in the product, but not a change of nature, and by the side of this law of polarity there is that of compensation ; for, in proportion as the anther-cell closes to render the pollen abortive, there is a development of the gland which begins to secrete the nectar; the evolution of the gland brings on the atrophy of the cell, but, fundamentally, there always remains an apparatus of emission.

Let us return to the Columbine : we have said that one way of transformation, and it is the most common one, would lead to a belief that the nectar-bearing sac may be in some instances the representative of a loculus of the anther. From

10 M. Morren on the Spur-shaped Nectaries

a slight examination we should in fact conclude so. As a proof of this, see the states delineated figures 3, 4, 5 and 6. We often see a stamen, with a filament dilated at its base, take two horns above (c d), whilst one loculus of the anther, inflated, no longer yields pollen ; and the other, being atro- phized to such a degree as no longer to appear except as a yellow gland [b), seems to have produced a rounded sac (e). This sac, the commencement of the cornet, should we not suppose it to be a modified anther-cell? and yet we have just seen that the tendency of the cells is to produce the lobes of the limb of the cornet, and not its tube. There is a mistake, indeed, as to the true signification of this en- largement, which is nothing but the middle of the connective itself. The connective extends itself outwards, and its hy- pertrophy brings with it the atrophy of the cells or of one cell of the anther ; it signifies little whence substance comes to it, so that it only come. This is why the production of the spur does not always cause the whole anther to be meta- morphosed all at once.

The better-formed cornets, and which even possess all the essential parts, expanded limb, apex with two lobes and a slit, dilated faux, lengthened tube and terminal gland ; these cornets, I say, sometimes still exhibit a trace of their old and primi- tive nature in the anther-cell, hardly visible, but distinguished by its yellow colour, whilst all the rest is white and blue, and, above all, distinguished by the grains of pollen that it still encloses in its bosom (fig. 4).

The conditions (figs. 5 and 6) are tendencies towards a re- gularized form of well-constituted nectaries. The condition (fig. 6) is that found in the common Aquilegice. Nothing here would lead to the supposition of an antherine nature, had not this strange metamorphosis been followed step by step.

It is evident, that all these cornets being hollow, and de- veloped one above the other in several spirals (fig. 16), all like- wise enter one into another (fig. 8), but it is inexact to say that then the glands no longer secrete. This is a mistake : the secretion continues, and, indeed, the tubes never com- pletely close those into which they have entered.

Let us now examine in what manner the cornets are gene- rated in a flower of Aquilegia taken at its first periods of de- velopment.

2. Organogeny of the spur-shaped nectaries.

To ascertain this