THE

J

OF

THE LINNEAN SOCIETY

■SjSflS^i**-^.-

BOTANY.

VOL. XV. G

EDITED BT DR. JAMES MURIE, F.D.S., F.G.S., F.Z.S., ETC

LONDON:

SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON HOUSE;

AND BY

LONGMANS, GREEN, READER, AND DYER,

AND

WILLIAMS AND NORGATE.

1877.

*

Dates of Publication of the several Numbers included in this Volume.

No. 81, pp. 1-40, was published October 11, 1875

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86,

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40-90,

83, 90-159,

84, 159-252,

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85, 253-303,

363-422,

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87, 423-480, 88, 481-548,

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March 3, 1876 May 11,

July 1 1 , September 14,

October 23,

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>>

December 15, February 28, 1877.

PRINTED BY TAYLOR AND FRANCIS,

RFI> r

COURT, FLEET STREET.

LIST OF PAPERS.

Archer, William, Esq., F.R.S., M.R.I.A.

Note on the Freshwater Algae collected by H. N. Moseley, M. A., in Kerguelen's Land. (Communicated by J. D. Hooker, M.D., Pres. Roy. Soc.) 445

Baker, John Gilbert, Esq., F.L.S.

On the Polynesian Ferns of the < Challenger' Expedition 104

Revision of the Genera and Species of Anthericese and Erio- spermeae 253

On a Collection of Ferns made by Mr. William Pool in the inte- rior of Madagascar 4U

Balfour, Isaac Bayley, D.Sc, F.L.S.

Extract of a Letter from I. B. Balfour, Esq., Botanist to the Expedition to Rodriguez to observe the Transit of Venus ; addressed to, and communicated by, Dr. Hooker 24

On a new Genus of Turneraceas from Rodriguez 159

Bentiiam, George, Esq., F.R.S., F.L.S., Vice-Pres. Linn. Soc. Notes on the Gamopetalous Orders belonging to the Cainpanu-

laceous and Oleaceous Groups

On the Distribution of the Monocotyledonous Orders into Pri- mary Groups, more especially in reference to the Australian Flora, with notes on some points of Terminology. (Plates VI1.-IX.) #

-

Berkeley, The Rev. M. J., M. A., F.L.S.

Enumeration of Fungi collected during the Expedition of H.M.S. ' Challenger.' (Second Notice.) 48

Report on the Fungi collected in Kerguelen's Land by the Rev. A. E. Eaton during the stay of the Transit-of- Venus Expedi- tion of 1874-75 221

1

490

f

IV

Page

Berkeley, The Rev. M. J., and C. E. Broome.

Supplement to the Enumeration of Fungi of Ceylon. (Plate II.) 82

Berkeley, The Rev. M. J., and M. C. Cooke.

The Fungi of Brazil, including those collected by J. W. H.

Trail, Esq., M.A., in 1874

363

Broome, C. E., and The Rev. M. J. Berkeley.

Supplement to the Enumeration of Fungi of Ceylon. (Plate II.) 82

113

116

Clarke, C. B., Esq., F.L.S.

On Edgctria, a new Genus of Cucurbitacese. (With wood- cuts.)

Botanic Notes from Darjeeling to Tonglo. (With wood- cuts.)

Crombie, The Rev. James M., F.L.S. &c.

Lichenes Capenses : An Enumeration of the Lichens collected at the Cape of Good Hope by the Rev. A. Eaton during the Venus-transit Expedition in 1874 165

Lichenes Terrse Kergueleni : An Enumeration of the Lichens collected in Kerguelen's Land by the Rev. A. E. Eaton during the Venus-transit Expedition in 1874-75 180

On the Lichens collected by Professor R. O. Cunningham in the Falkland Islands, Fuegia, Patagonia, and the Island of Chiloe, during the Voyage of TI.M.S. < Nassau/ 1887-69 222

Lichens collected by W. Pool, Esq., in Madagascar 940

Lichenes Insulse Rodriguesii. An Enumeration of the Lichens collected by Dr. I. B. Balfour in the Island of Rodriguez during the Venus-transit Expedition, 1874 431

*

Contributions to the Botany of II.M.S. i Challenger':

, Numbers XXI. to XXXVII inclusive. Consult respectively Archer, XXXIV., p. 445 ; Baker, XXXI., p. 104; Ber- keley, XXVI., p. 48; Dickie, XXI.-XXV., pp. 40-47, XXXIII., p. 235, XXXV., p. 446, XXXVII., p. 486 ; Mitten, XXIX., p. 59; Moseley, XXVIL, p. 53, XXX., p. 73, XXXVL, p. 481 ; O'Meara, XXVIIL, p. 55 ; Reichen- bach, XXXII. , p. 112.

Cooke, M. C, and The Rev. ML J. Berkeley.

The Fungi of Brazil, including those collected by J. W. H. Trail, Esq., M.A., in 1874 368

Page

Dar*win, Francis, Esq., M.B., F.L.S.

On the Glandular Bodies on Acacia sphcerocephala and Cecropia peltata serving as food for Ants. With an Appendix on the Nectar-Elands of the Common Brake Fern, Pteris aquilina.

(Plate VI.)

398

Dickie, Professor George, M.A., M.D., F.L.S.

Notes on Alffffl from the Island of Mangaia, South Pacific

30

Alo-re collected bv Mr. Moselev at Simon's Bay, Cape of Good Hope 40 (N.B. Title of this correct on wrapper, hut at p. 40, XXI. of Contributions to the Botany of II.M.S. « Challenger,' con- tains part title of Nos. XXII. to XXV. inclusive.)

Algse collected by Mr. Moseley at Seal Island 41

Algse collected by Mr. Moseley at Marion Island in 40 fathoms 42 Marine Algse collected by Mr. Moseley at the Island of Kerguelen 43 Al°-ae collected by Mr. Moseley at Heard Island, 250 miles S. of

A*7

Kerguelen

Notes on Alffffl found at Kerguelen's Land by the Key. A. E.

Eaton

193

Algse chiefly Polynesian : being Contributions to the Botany of H.M.S. < Challenger ' 235

Notes on Alga collected by II. N. Moseley, M.A., of H.M.S. « Challenger,' chiefly obtained in Torres Straits, Coasts of Japan, and Juan Fernandez 446

Supplemental Notes on Algse collected by H. N. Moseley, M.A.,

of II.M.S. ' Challenger,' from various localities 486

Dyer, W. T. Thtselton, Esq.,M.A.,B.Sc, F.L.S., Assist. Director

Roy. Gardens, Kew.

On the Plant yielding Latakia Tobacco (vide Corrigenda et

246

25

Addenda, Lattakia)

On the Genus Jloodia, with a diagnosis of a new Species. (PI. V.

and one woodcut.)

Gammie, J., Esq.

Extract of a Letter from Mr. J. Gammie to Dr. Hooker [On spadix

of Ariscema speciosum]

Gilbert, J. H, Ph.D., F.R.S., F.C.S.

Note on the Occurrence of " Fairy-Rings " 17

Horne, John, Esq., F.L.S.

Extract of a Letter addressed to Dr. Hooker by John Home, E«q F L S., Subdirector of the Botanic Gardens, dated Mau- ritius 12 November, 1874. [On the Botany of the Seychelles.] 27

VI

Pag#

King, G., Esq., M.B., F.L.S., Superintendent of Botanic Gardens,

Calcutta. Note on a Sport in Paritium ti'icuspe, G. Don 101

Kirk, John, M.D., F.L.S. (H.B.M. Consul, Zanzibar.)

Identification of the Modern Copal Tree ( Trachylobium Home- mannianum) with that which yielded the Copal or Aninii, now found in the earth on the East Coast of Africa, often

where no copal-yielding trees now exist 234

Note on specimens of Hibiscus allied to //. rosa-sinensis, L., col- lected in E. Tropical Africa, with remarks by Prof. Oliver,

(Woodcuts.) 478

Masters, Maxwell T., M.D., F.R.S., F.L.S.

Remarks on the a Superposed " Arrangement of the Flower . . 456

Mittkn, William, Esq., A.L.S.

The Musci and Hepaticre collected by H. N. Moseley, M.A., Naturalist to H.M.S. < Challenger ' 59

A List of the Musci and Hepaticse collected in Kerguelen's Land bv the Rev. A. E. Eaton, M. A 193

Mooiie, S. Le Marchant, Esq., F.L.S.

Occurrence of Staminal Pistillody in an Acanthad. (Plates

in. &iv.)

86

Moseley, H. N., Esq., M.A., late Naturalist H.M.S. ' Challenger f Expedition.

Further Notes on the Plants of Kerguelen, with some remarks

on the Insects. (In a letter addressed to Dr. Hooker, Pres.R.S.) 53 Notes on Plants collected and observed at the Admiralty Islands,

March 3 to 10, 1875 73

Notes on the Flora of Marion Island. (Communicated by Prof. * Oliver, F.R.S., F.L.S.) 481

Oliver, Prof. Daniel, F.R.S., F.L.S., Keeper of the Herbarium, Roval Gardens, Kew.

List of Plants collected in New Guinea by Dr. A. B. Meyer, sent to Kew December 1874 29

Enumeration of Plants collected by V. Lovett Cameron, Lieut. R.N., in the region about Lake Tanganyika 90

Note on a Collection of North-Celebes Plants made by Mr. Riedel of Gorontalo 97

VII

Pa«re

O'Mkara, The Rev. E., M.A.

On the Diatoraaceous Gatherings made at Kerguelen's Land by H. N. Moseley, M.A., H.M.S. ' Challenger.' (Communicated by Dr. Hooker, Pres.R.S., V.P.L.S.) (Plate I.) 55

Potts, T. H., Esq., F.L.S.

Habits of Filices observed about the Malvern Hills, near the Gorge of the Rakaia, Canterbury, New Zealand 423

Reichenbach, Prof. H. G. . On some Orchidacere collected by Mr. Moseley, of the l Chal- lenger ' Expedition, in the Admiralty Islands, Ternate, and Cape York, one of which forms the type of a new section of the Genus Dendrobium. (Communicated by Prof. Oliver, F.R.S., F.L.S.) 112

Reinsch, Paul Frederick, Esq.

Species ac Genera nova Algarum aquae dulcis quae sunt inventa in Speciminibus in Expeditione Vener. transit, hieme 1874-75 in Insula Kerguelensi a clar. Eaton collectis. (Communicated by Dr. Hooker, C.B., Pres.R.S., F.L.S.) 205

Sorby, H. C, Esq., F.R.S., F.L.S., (fee, Pres. Roy. Micros. Soc.

On the Characteristic Colouring-matters of the Red Groups of Algae. (With woodcuts.) 34

Trimen Henry, Esq., M.B., F.L.S.

Note on Boe'ea Commersoniij R. Br 163

Vines, S. H., Esq., B.A., Fellow of Christ's College, Cambridge. On the Digestive Ferment of Nepenthes. (Communicated by W. T. Thiselton Dyer, M.A., F.L.S., Assist. Director Roy. Gardens, Kew.) 427

EXPLANATION OP THE PLATES.

Plate

I. Diatoms. Illustrations of new Marine and Freshwater Species from Kerguelen's Land as determined by the Rev. E. O'Meara.

II. Fungi of Ceylon. Alwisia bombarda and Actlniceps Thivaitesi, and

details of same figured as referred to in the Rev. M. J. Berkeley and Mr. C. E. Broome's paper.

III. | Whiteieldia lateritia. Various Views of Normal and Abnormal & I Flower of, to illustrate Mr. S. Le Marchant Moore's paper on Sta- I"V. I minal Pistillody.

V. IIoodia and Decabelone. Gynostemium of Species of these Genera,

viewed from above and in section, to illustrate Mr. W. T. T. Dyer's paper on the Genus Hoodia.

VI. Glandular Bodies and Nectar-glands of Acacia sphcerocephala,

Cecropia peltata, and Pteris aquilina, illustrating Natural Appear- ance and Histology of these Organs according to Mr. Francis Darwin's researches thereon.

VII. Xtris and Eriocatjlon. Diagrammatic Illustrations of these Genera,

showing the Homology of Perianth-segments &c, in illustration of Mr. Bentham's paper.

VIII. Cyperace^:. Diagrams to represent the Homology of Parts of same in

various Genera and Species as determined by Mr. Bentham.

IX. Gramine^:. Diagrams illustrating the Homology of their Parts in seve- ral Grenera, according to Mr. G. Bentham's vi

COKKIGENDA ET ADDENDA.

XXII

XXV

wrapper, viz. "Algae collected by H. N. Moseley, Esq., M.A., at Simon's Bay, Cape of Good Hope." 56, 4 from top, /or " Herd Island " read Heard Island.

163, Mr. Trimen's paper was read in 1876, not 1875.

167, 16 from bottom, "F. subcornuta" should be C. subcornuta.

170, 12 from top, " F. minor w should be P. minor.

224, 11 from top, "F. costata" should be C. costata.

224, 10 from bottom, in Lamarkii " c has been omitted.

235, 12 from bottom, add XXXIII., after " Contributions to the

Botany of H.M.S. ' Challenger/ "

238, 3 from top. The asterisk (*) attached to Sargassum ilici-

folium refers to the two lower lines of p. 239, which ought to have formed a footnote on p. 238.

246, 4 from top. The author of the paper states that Mr. Eldridge

and Prof. Post, his correspondents and authorities for certain statements, gave the spelling Lattakia (probably

Latakia

in this country.

445, 17 from top. After "Communicated by" delete " Sir".

452, 6 from bottom, for " micranthum " read micracanthum.

453, 12 from top, the first " u" has dropped out in Scutellum.

453, 19 from bottom " Climacosphaua " should be Climaco-

sphenia ; and " moniligeea " should be moniligera.

454, 2 from top " Cespitalis" should be C.espitalus. 454, 13 from bottom,/or "inserta" read incerta.

454, 7 from bottom, the specific name "coNCIONA,, should be

CONCINNA.

THE JOURNAL

Off

THE LINNEAN SOCIETY.

Notes on the Gamopetalous Orders belonging to the Campanu- laceous and Oleaceous Groups, By George Ben¥ham, F.R.S.

[Read March 4, 1875.]

It is now more than thirty years since the volumes of the ' Pro- dromus * containing the above orders were published ; and during that period the great additions made to our knowledge, especially of tropical vegetation, have in many instances required consider- able modifications of their tribual as well as generic arrangement. I have endeavoured to carry this out in preparing them for the forthcoming part of our ' Genera Plantarum ;' but there is much of detail respecting the reasons which have induced me sometimes to depart from the principles adopted by former monographists, which the limits of that work prevent me from there entering into, and which I should therefore be desirous of placing on re* cord in the ' Journal of the Linnean Society/

1. Campanulaceae and their allies.

The orders constituting this generally recognized and well- defined group are usually reckoned as seven Stylidiese, Goodeno- vieae, Brunoniaceae, Lobeliaceae, Cyphiaceae, Campanulaceae, and Sphenocleaceae. They would appear to me to be more naturally and clearly defined if reduced to three Stylidieae, Goodenovieae and Campanulaceae ; for the exceptional characters upon which Brunonia, Cyp}iiay and Sphenoclea have been isolated do not

than

ac

LINN. JOURN, BOTANY, VOL. XV. B

2

MB. G. BEJSTHAM OK THE

count proposed to separate ; and the union of Lobeliaceae with Campanulacese, often proposed and rarely distinctly objected to, appears to me to be a most natural one, as I shall presently endeavour to show.

With regard to Stylidiese, having so recently worked them up for the Australian Flora, I have now no remarks to make with regard to the genera Stylidium and Levenhoolcia, almost exclu- sively Australian, and constituting the great bulk of the order.

I would only observe, as to the two small southern genera {Fors- tera and Fhyllachne), that they were very well distinguished by Swartz in Schrader's Journal for 1799, and in the first volume of Konig and Sims's * Annals of Botany.' Willdenow and, after him, Endlicher and De Candolle united them under the name of Forstera. F. Mueller in a recent number of his c Fragmenta ' adopts the same view, but proposes to substitute the name of Phyllachne for the generally received one of Forstera. In the * Flora Austra- liensis ' I had only to deal with a true Forstera, which I entered under that name. Dr. Hooker, for his series of Southern Floras, had to examine them both ; but not having before him the fruit of the typical Phyllachne, he presumed that his predecessors were right in treating it as the same as that of Forstera, and main- tained the two only as sections of one genus ; but having received from New Zealand a new species with the habit of his section

with

For

Hera, he therefore proposed it as a new genus under the name of Holophyllum. It is now, however, shown, both by spe- cimens and by Hombron and Jaquinot's figures (under Forstera), that the original Phyllachne has not only the habit and inflores- cence, but also the indehiscent fruit of Holophyllum, and forms

with it a generic group which Swartz was fully justified in distin- guishing from Forstera.

Goodenovie m, as the order was originally named by its founder, Robert Brown, or Goodeniacece, as they are styled by the modern advocates of monotony in the names of orders, are, as well as StylideaB, almost exclusively Australian, and are worked up in de- tail in my Flora. On that occasion, having at my disposal a large proportion of original specimens described by De Vriese, I was fortunately enabled to clear up much of the extraordinary confu- sion exhibited in that botanist's monograph. I also there gave my reasons for following Brown in including JBrunonia in the order, which, notwithstanding the anomalies of this monotypic genus, is

CAMPANULACEOUS AND OLEAOEOUfl ORDERS. 3

so definitely characterizad by the indusium as to have no ambi- guity in its circumscription. The homology of this indusium has been variously interpreted. R. Brown, in the then state of our acquaintance with the organs of other plants that might be com- pared with it, put the following queries (Trans. Linn. Soc. xii. 134) : "Is this remarkable covering of the stigma merely a pro- cess of the apex of the style ? or is it a part of distinct origin, though intimately cohering with the pistillum ? On the latter supposition, may it not be considered as analogous to the glan- dular disk surrounding or covering the ovarium in other families ? And in adopting the hypothesis I had formerly advanced (Trans. Linn. Soc. x. 159) respecting the nature of this disk in certain families, namely that it is composed of a series of modified sta- mina, has not the part in question a considerable resemblance in apparent origin and division to the stamina of the nearly related family Stylidiaceae ? To render this supposition somewhat less paradoxical, let the comparison be made, especially between the indusium oiBrunonia and the imperfect antherae in the female flowers of Forstera" All further observation, however, has tended rather to oppose than to confirm this supposition. Brown's idea of the conformity of the disk, scales, and stamens as alluded to in the passage above quoted, appears to have been founded, not so much on his own observation, as on Labillardiere's having given the name of sterile stamens to the hypogynous glands or scales of CenarrJienes, which glands, however, in fact give no indication of any structural or morphological analogy to stamens. It is very rarely indeed that the disk has been shown to be a rudimentary whorl of organs, whether a reduced remnant of extinct organs or a first development of new ones. It is, on the contrary, now generally admitted that whether peripetalous, as in the case of the ring of calycine glands or scales in many Apocynese and Aselepiadeae, or peristaminal, as in some Polypeta- lous orders or genera, or, as is more frequently the case, truly pe- rigynous, hypogynous, or epigynous, the disk is a mere outgrowth of the receptacle or summit of the floral pedicels, promoted pro- bably in many instances by the attractions its secretions afford to insects and then a true nectary, possibly developed in others as a store of nutriment during the early stages of the young embryo, but seldom, if ever, independently organic. If on some occasions it shows a ring of lobes alternating with those of the androecium or other immediately external whorl, that is merely due to the

b2

4 MR. G. BENTIIA.M ON THE

pressure of that external whorl during its growth in the bud- That the indusium cannot well be a second abortive whorl of sta- mens is further proved by the consideration that that whorl of organs is always uniseriate in the whole Campanulaceous group of orders, and that that single series exists in a perfect state in Goodenovieae in its usual position at the base of, or slightly adnate to the corolla ; nor can we readily compare it with a disk, as that excrescence, when epigynous, is, I believe, always at the base, not at the summit of the style.

An analogy between the indusiums of Goodenovieae and the ring of collecting-hairs observable in many Lobelieae has often been distinctly brought forward or tacitly assumed even very lately, as by Sonder in the ' Flora Capensis ' with regard to Cyphia, or by Asa Gray in his note on Nemacladus in our Journal (xiv. 28); but it was ably refuted by Brown when pointing out the impor- tant characters separating Goodenovieae from Lobelieae. These collecting-hairs are indeed in a ring below the stigma in many Lobelieae and in some Campanuleae ; but in some of the former and most of the latter they are differently arranged, either coTer- ing the whole of the upper moiety of the stjde, or in longitudinal rows, or more or less unilateral, &c. ; and in all cases they are en- tirely epidermal productions, whilst the indusium is a development of the substance itself of the style,

Lindley may have come much nearer the truth when he says (Veg. Kingd. 694) that it " is to be regarded as nothing more than a remarkable exaggeration of the rim which surrounds the stigmatic surface of Heathworts (Ericaceae), and of the plates which cover the style of Cranesbills and Balsams (Geraniaceae and Balsamineae) . It is, in fact, the free upper extremity of the carpellary leaves, distinct from that prolongation of the placenta which is named style and stigma." I cannot quite concur with the latter part of this proposition ; for I should confine the term placenta to that substance which more or less lines the inner walls of the ovary-cells or protrudes from them and bears the ovules, and regard the whole style as the prolonged apex of the carpellary leaves, entire or variously lobed, or enlarged at the extremity and bearing the stigmatic surface either on the summit, or within, or on the margins of the lobes, or below them on the outside. And if, with Lindley, we may compare the indusium with the ring sur- rounding the stigma in Ericaceae, we may perhaps still better com- pare it with the dilatation at the end of the style commonly called

CAMPANULACEOUS AND OLEACEOUS ORDERS.

the stigma, in Apocyneae and Asclepiadeae, with this difference, that in the latter two orders the dilatation bears the stigmatic surface or spots on its outside, whilst the terminal bifid apiculus, some- times forming a large beak into which the dilatation tapers, some- times reduced to very small dimensions in the centre of the con- cave disk-like dilatation, is usually, if not always, inert, whilst in Groodenoviesc the concave dilatation or indusium is inert, and the small bifid apiculus in its centre is stigmatic. The same differ- ence in the position of the stigmatic surface is observable in He- liotropieae as compared with other Boragineae : that the indusium is thus in fact the concave apex of the style may be further con- firmed by the case of Leschenaultia, where it is more or less di- stinctly two-lobed, and the stigmatic surface, instead of being sti- pitate in the base of the cavity, lines the inside of one at least of the lobes, thus forming an approach to the ordinary bilamellate stigmas of a large number of Gamopetalae.

In the Campanulacejs we have, as has often been proposed, re- united as tribes or suborders the two great groups of Lobelieae and Campanuleae. Together they form a determinate wholesur rounded on every side by a broad gap without any intermediate or ambi- guous form. Their nearest neighbours are the above-mentioned Stylidieae and Goodenovieae ; but besides the minor characters brought forward by Brown, it may be enough to observe that there is in Campanulaceae no indication either of the singular consolida- tion of the filaments and style of the former, or of the indusium of the latter order. Between the tribes, however, there is no such definite line of distinction. At first sight, indeed, the irre- gular flowers and alternate anthers of Lobelieae would seem to form together a good ordinal character to separate them from the regular flowers and free anthers of Campanuleae; but though constant in the great majority of cases, there are various except tions with" different combinations. There are species of Isotoma and Lobelia where the obliquity of the corolla is very slight ; and, on the other hand, in Leptocodon and several species of Campanula and Phyteuma it is very perceptible. At the most, indeed, the irregularity of Lobelieae is but little more than obliquity. There is no tendency to the bilabiate aestivation or to the didynamy of the Personate orders ; the equally valvate aestivation and the iso- mery of the stamens are constant, both in Lobelieae and Campa- nuleae ; the structure of the pistil, fruit, and seed is the same in both tribes as to all essentials, and exhibits similar variations in both as to minor points.

6 MR. G. BENTHAM ON THE

Nemacladm

w » »/X XT * 7 •*—- *- ^^Jf*1"*

the two former from western extratropical America (south and north), the latter from South Africa, which have very decidedly the irregular corolla of Lobelieae with the free anthers of Cainpanulea?. We have for technical convenience placed them together in a small third tribe intermediate between the two others ; but, strictly speaking, they may not perhaps form a truly natural group. In general character indicative of natural affinity, each one of the three may prove to be more nearly allied to some one belonging to one of the other tribes than to either of the two now associated with it ; but yet the actually inserting them in those tribes would have interfered too much with a clear methodical exposition of the order to be admissible at present, besides that it is by no means certain that they have not a common connexion.

An elaborate and careful monograph of the tribe Campanulas was the first botanical publication of Alphonse De CandoUe, and gave him at once a distinguished place amongst the followers of systematic botany, although that has never been his special branch of the science. He subsequently edited the whole group for the 1 Prodromus ; ' and he may always be relied upon for accuracy of detail as far as his own observations went. If we have found it necessary to remodel some of his genera consisting chiefly or en- tirely of extra-European species, or to lower the systematic grade he had assigned to some of his groups, this is owing either to the additional lights thrown on the subject by subsequent discoveries, or, in the case of several Lobelieae, to the too great reliance placed by him on the preceding observations and conclusions of Presl and Don ( George Don, assisted, I believe, in many respects by his brother David), many of which are very loose and hasty.

The Lobelieae, however, are exceedingly difficult to divide into definite genera. The most dissimilar groups often run into each other by almost insensible gradations. I have endeavoured to keep up all genera which appeared to be founded on more appre- ciable characters, such as the consistency and dehiscence of the fruit, the placentation of the ovary, the dorsal or ventral fissure of the corolla, the attachment of the stamens, &c. But I have felt obliged to leave the genus Lobelia itself a very large one, however heteromorphous it may appear at first sight. The St.-Helena Trimeris, the large tropical species which I have referred to the section Bhynchopetalwn, the Central- American section Homochilus, the large scarlet North-Ameriean Euhbeli<t. and the small Mm

CAMPANULACEOUS AND OLEACEOTJS ORDERS. 7

South- African and other species, all retained as Lobelia by De Can- dolle and others, appear to me to differ more from each other in habit and character, than some of them do from the American Tupce, or from the small South- African groups generically sepa- rated by De Candolle after Don and Presl, chiefly from the degree of irregularity or fissure of the corolla, which it is often impossible to define in words. Amongst the latter, however, I was at first disposed to maintain the genus Parastranthus, founded on two species with yellow flowers quite sessile, without the reversion which takes place generally in the tribe by the torsion of the pedicel ; but to these two Sonder has added, as a third species, the Lobelia leptocarpa, DC, with shortly pedicellate blue flowers, in which, as far as I can judge from dried specimens, the re- version takes place in some flowers and not in others, thus quite invalidating the generic character. There are also, I be- lieve, other cases of partial or variable reversion.

Lobelia Bergiana, Cham., under Presl's generic name Gramma- totheca, had been associated by De Candolle with Clintonia, Dougl. (now Bowningia), in a distinct tribe, with the remark- able character of a unilocular capsule with two parietal pla- centae, but opening in three valves. . To this tribe he gave the name of Clintonieae, altered to Grammatothecese by A. Gray, who (Journ. Linn. Soc. xiv. 29) remarks on it as a second in- stance of Lobeliaceous genera related to each other found in the two distant localities California and South Africa. But all this is founded on the hasty erroneous observations of Presl, never since verified, as I pointed out (Fl. Austral, iv. 128). The only character the two plants have in common is the long narrow ovary and capsule, which occurs here and there in single species of other genera. . L. ( Grammatotheca) Bergiana, accurately figured in Delessert, Ic. Sel. v. t. 6, has the true fruit of a Lobelia, completely two-celled and opening at the apex in two short locu- licidal valves between the calyx-lobes ; and the only way I can account for Presl's singular error is from his having observed old empty capsules, which in their decay have split up longitudinally below the calyx-lobes, the walls of the cells separating from the dissepiments, which latter he must have mistaken for a third placentiferous valve. In Bowningia the capsule remains closed at the apex, but really opens below the calyx-lobes in lateral slits for the emission of the seeds. As the walls of the cap- sule are made up of the pericarp and the adnate calyx-tube

8 MR. G. BENTHAM ON THE

composed of five sepals, each with its midrib, the lateral dehis- cence would naturally take place in the thinner spaces between those nerves or midribs, as is clearly shown in Campanula, where the thin part of the pericarp between the dissepiments coincides with the thin part of the calyx-tube between the nerves or ribs. In Downingia, however, the ovary, two-celled at an early stage, becomes one-celled by the drying up of the exceedingly thin dis- sepiment, and the two placentae remain attached to the inner walls of the cavity, each one opposite the junction of two sepals. "When the capsule at maturity has to split open between the se- pals, four of them are held together in pairs by the adnate pla- centae, and between these pairs there is on one side one slit, and on the other side the intervening odd sepal induces two slits, one on each side of it ; and we have as the natural result three valves, two of them placentiferous and the third naked, as described. This is the only instance among Lobeliese proper of the lateral dehiscence, which is also exemplified, but apparently in a single slit, in the Chilian Cypliocarpus among Cyphiese, a genus otherwise allied in many respects to Dow?iingia, and in a few genera of Campanuleae, although there rather in short valves than in long slits, excepting, indeed, in Githopsis, an outlying member of Campanulese, as Downingia is of Lobeliese, and connected with the latter in some measure in this respect as in native country.

In geographical distribution the Campanulaceae may be reckoned, for the most part, amongst the herbaceous races of extratropical origin, and in this respect analogous to a large portion of the Compositae. There is nothing also to oppose the idea of their early development having been contemporaneous with that of the Compositae, although their subsequent progress has been so much more limited. But I confess myself quite unable to see any grounds for supposing, with Delpino, that Lobeliese are the parents of Compositae. Moreover, if the two orders had a com- mon parent, it must have been a very remote one with a long- passed extinction of all the races which had formed the inter- mediate stages. The intervening gap which now separates them is too wide and deep. In the important point of the pistillary structure no genus or species of the tribe Lobeliese or of the whole order of Campanulaceae shows any approach to that which is so uniform in Compositae ; nor indeed does any of the whole group of Campanulaceous orders, unless it be some slight indica- tion in one or two species of Goodenovieae, the furthest removed

CAMPANULACEOUS AJND OLEACEOUS ORDEBS. 9

from Lobelia. Turning it over again and again, I can discover

no plausible foundation for Delpino's genealogical Table from

Lobelia to Artemisia.

The present distribution of Campanulaceae seems to indicate a

southern origin for Lobelieas and a northern one for Campanulese. The southern extratropical or mountain-range of herbaceous LobeliesB is extensive and varied. They are represented by iden- tical genera, sections, or even species in South Africa and Aus- tralia {Lobelia Bergiana, L. anceps, and allies), or in Antarctic America, South-east Australia, and New Zealand (Pratia), and have established several minor more or less endemic groups gra- dually diverging from the common types in South Africa, Aus- tralia, and Extratropical America. From thence Lobeliese appear to have spread in several distinct directions into and beyond the tropics, without any transverse northern connexion between the several lines, which may be traced as follows :

First, and most abundantly, along the western mountain- ranges of America, where they have developed into the shrubby genera Siphocampylus, Centropogon, and Burmeisteria, of which nearly two hundred species are already known, all remaining en- demic in the tropical or subtropical mountain-regions, with the single exception of the Centropogon surinamensis, which has gene- rally spread over tropical America. Secondly into tall herba- ceous true Lobelice with a wider general range, having formed, however, special groups of a more local character, such as the Tupce of Chili, the thapsoid Brazilian species, the Tylomia of the West Indies, the Homochili of Mexico, the JEulobelice of extra- tropical North America. Thirdly, into the Pratioid genera Hyp- sela, Lysipoma, and BkizocepJialuni, which have remained almost entirely confined within the South-American, Andine, or extra- tropical regions, the JPratia hederacea alone, the exact counterpart of the Himalayan P. begonicefolia, having extended further into South Brazil. Fourthly, the southern Hemipogon group has main- tained the typical characters in a few species thinly scattered over the range, chiefly in Mexico, but has also developed into endemic groups passing gradually into the Mexican Heterotopia, more ab- ruptly into the north-western Downingia. The southern Lau- rentia is also very closely represented by the Mexican L. ratno- sissima and by the Californian Porterella, and the southern Iso- toma by the West-Indian Hippobroma.

In the Old World the northern dispersion of the Lobelieae ap-

10 . ME. G. BENTHAM ON THE

pears to have been chiefly in the extreme east over the Indo- Australian region with much less of modification of their original southern character than in America. The Asiatic Lobeliese have remained within the limits of the two widely spread genera Pratia and Lobelia, slightly modified as they enter or pass the tropics, in a manner corresponding to that observed in Brazil, but with- out any tendency to the marked endemic groups of Western America. Thus, in JPratia, the Himalayan P. begonicefolia is, as above-mentioned, a close representative of the Brazilian P. Jiede- racea. The P. montana, from the Archipelago and eastern pro- vinces of India (Spirema, Hook. f. et Thorns.), is an endemic mo- dification, but scarcely generic, diverging in habit, but much less in character than the New-Zealand Colensoa. The genus Lobelia itself has preserved much of the southern type in some species both of Hemipogon and Holopogon, but has also, by somewhat gradual modifications, developed in considerable variety the tall herbaceous JRhynchopetalum forms, sometimes nearly counterparts of the Brazilian thapsoid species, and ranging across the Asiatic continent and to the islands of the China seas.

There has also been some extension of the tribe northwards along the extreme west of the Old World : one or two represen- tatives of South- African types (Laurentia Michelii, DC, and L. tenellciy DC.) are now very rare in the Mediterranean region ; another (L. urens) extends along Western France to Britain, but has been unable to penetrate eastward. From this L. urens in the extreme west of Europe to the L. sessilifolia in the extreme east of Asia, a continuous expanse of land nearly half encircling the globe, there is not to the north of the great Alpine Caucasian and Himalayan line a single trace of the tribe.

L. Dortmanni, an aquatic species of North-western Europe, may, like the JEriocaulon and others, have come over from North Ame- rica. With regard to the Lobelice of the tthynchopetalum group in tropical Africa we have not sufficient data to form any opinion as to their origin. They are perhaps more likely to be slightly modified Asiatic species than endemic developments of the more distant South-African forms.

Lobeliese must also have found their way very early to the distant islands of the Atlantic and Pacific ; for they have there generated special endemic forms with the usual insular cha- racter of a ligneous development of races more generally her- baceous. Thus they have produced the Trimeris (Lobelia sect.)

CAMPANULACEOUS AND OLEACEOUS OBDEBS. 11

of St. Helena, the Sclerotheca of the Society Islands, and the five genera constituting the Clermontia group of the Sandwich Islands.

In regard to geographical distribution, the CyphiesB may be in- cluded in Lobeliese, to which they are otherwise more nearly allied than to the Campanulese. Of the three genera, the principal one, CypMa, is entirely South African, one species only having reached Abyssinia; the monotypic Cyphianthus, from Chili, smAJVemacladus, from North-west America, have followed the course of other races of southern origin, travelling northwards along the western regions of America.

We

World

beliese had been generally southern. The union of the two in

one very natural and well-defined order would indicate a common

origin ; but what that parent race was, or what was its primitive

home, is, I believe, with our present data, beyond the reach of

conjecture. All we might venture to imagine appears to be :

That the primitive race flourished very early in some region in connexion with Africa.

That the Lobeliese were first developed at a time when the geological or ether conditions afforded some general means of southern communication between South Africa and Australia, between Australia, New Zealand, and Antarctic America, between South Africa and extratropical South America.

That when the Campanulese were differentiated, South Africa was already isolated from the rest of the southern hemisphere.

That this branch of the descendants of the primitive race having then become established in the African region, both to the north and to the south of the hot zone, developed freely, ex- tended widely, and became much varied in the wide area opened to it in the north, but remained much more restricted in num- bers and variations in the limited space allowed it in the south.

That it spread very sparingly over the tropics, with but little variation, and was still more restricted in crossing the tropics southward at a distance from the primitive home.

Such are the principal conjectures which suggest themselves on the consideration of the present distribution of the tribe of which the following is a general sketch.

We may perhaps be justified in taking as the earliest forms in

12 MR. G. BEHTHAM OJST THE

which Campanulese were developed those genera in which the cap- sular dehiscence is normal in terminal valves, especially Wahlen- bergia. This is now the most widely spread genus, being almost cosmopolitan. It is especially abundant in South Africa, where it passes gradually into the closely allied genera Lightfootia and Microcodon. It does not scorn the tropics, where it has generated the two or three species of the closely allied Cephalo- stigma. In the north, besides one almost cosmopolitan species^ it takes the rather distinct forms of the W. hederacea in Western Europe and the several Edrianthi of the East Mediterranean region. It has spread over Asia, passing into the Chino- Japanese JPlatgcodon ; and one Indo- Australian form has extended south- wards to New Zealand, where is also a peculiar endemic species, more nearly connected, however, with the Asiatic than with any other forms. It has also found its way, with other early races, to some of the distant islands, characteristic shrubby forms being found in Saint Helena and in Juan Fernandez. In America it is only to be met with in the tropics, but very sparingly, and not in any peculiar endemic form. The European Jasione, though a very distinct type, may be considered also as one of the Wahlenbergia group. And diverging from it, but in a different direction and in a greater degree, are the three Himalayan genera Leptocodon, Codonopsis, and Cyananthus. Although the latter genus, on account of its trimerous ovary being superior or nearly so, had been originally referred to Polemoniacese, I cannot see any real affinity with that order. The three genera are closely allied to each other, and appear to me to be the result of a differentiation, diverging from the Wahlenbergia group, which has not gone further, and not the remains of the extinct races

which had intervened between Campanulacese and any other order.

Four small genera of Campanuleae, with indehiscent fruits bac- cate or nearly so, are exclusively northern, though not so extra- tropical as most of the others. Canarina, a single Canary-island species, is nearly allied to some of the five East- Asiatic species of Campanumcea ; and these, again, are connected in some measure with the Wahlenbergia group through the above-mentioned Codo- nopsis. But the Himalayan Peracarpa, a single species, and JPentaphragma, from the Malayan archipelago, with three species, are quite isolated, showing no connexion with each other or with any other genus, except the general one with the tribe. The

CAMPANULACEOTJS AND OLEACEOUS ORDERS. 13

intermediate races which might have shown their special deri- vation are lost to us.

The Campanulese in which the capsular valves remain closed and consolidated into a coriaceous or hardened, flat or convex, apex, are numerous, and distributed into a number of more or less distinct genera both in the northern and