398 MR. F. TOWNBEND ON AN ERXTHRBA

On an Erythrea new to England, from the Isle of Wight and South Coast. By FREDERICK TOVNSEND, M.A., F.L.S.

[Read December2,1880.]

(PLATE XV.)

IN the ‘ Journal of Botany ’ for October 1879 I drew attention t o an Erythrea which 1 had lately found on the downs of Freshwater in the Isle of Wight, and to which at that time I was unable t o give a name, though inclined to think i t might be Erythrcea capi- tata, Willd. I hesitated-to refer i t to that species, because my plant differed in some respects from the longer description given by Rcemer and Schultes in their ‘ Systeina Vegetabilium,’ and I was unacquainted with Willdenow’s plant either in the living or dried state.

On first gathering the Erythraea in the island, I was much struck with its habitual dwarf character, in which it approached a dwarf and capitate variety of Erythrcea Centaurium, which also grew abundantly in the same locality. I soon found it easy to distinguish the former by its narrower leaves, by the shortness of its corolla-tube, and, above all, by its almost free stamens, the filaments of which, without exception, I found to be attached only a t the base of the corolla-tube, and to be otherwise perfectly free within it (compare fig. 3 b and fig. 6). This latter character seemed to have been hitherto unobserved in any species of the genus Erythrcea; and indeed it would take the plant out of the genus, as described by Grisebach in his monograph of the order Gentianaces, given in the ‘ Prodromiis ’ of De Candolle, where he says “Stamina 5-4, corollm tubo superiore inserta ” (DC. Prod. ix. p. 57).

It has been suggested to me both by Dr. Syme andDr. Ascher- son that the comparatively free stamens might represent a ten- dency to, or “ the last remains of,” dimorphism, like Menyanthes and Limnanthemuin in the same order, or like Primula &c. ; and if this were the only character by which the plants could be distin- guished, or that a similar tendency could be traced in other spe- cies of the genus, I should myself be disposed to give some weight to the suggestion ; I have, however, carefully searched for other examples in the same genus, and have examined numerous speci- mens, especially of the following species, E. Centauritrm, E. pub- chella, and E, littoralis (E. linariflia, Pers.), in all of which I

FROM THE ISLE OF WIGHT AND SOUTH COAST. 399

have invariably found the filaments attached at or near the throat of the corolla, and of comparatively equal length, whilst, as already observed, I do not know an instance in which the filaments are ao attached in the Freshwater plant

Through the kindness of Dr. Ascherson, I have lately been able to examine an authentic specimen of Erythrma capitata, Willd., from the Berlin Herbarium. In this specimen I find the fila- ments are likewise attached only a t the base of the corolla-tube ; and having asked Dr. Ascherson to examine specimens with espe- cial regard to the insertion of the filaments, he informs me that the Berlin specimens agree in this curious character and also in general habit. I have not been able to consult other specimens myself, and cannot find that any of true E. capitata, Willd., exist except in the Berlin Herbarium.

The earliest record of E. capitata is that given by Chamisso in his ‘Adnotationes quaedam ad Floram Berolinensem, auc- tore Adalbert de Chamisso ’ (1815), 8 ~ 0 ~ 1 3 pp. These ‘Adnota- tiones ’ are found in an appendix to a third edition of a cata- logue published anonymously, the author of which is F. Walter (chief gardener of Frau von Friedland and Count Itxenplik), who died in 1865. The title of the catalogue is ‘ Verzeichniss der auf den friedlandischen Gutern cultivirten GtewPchse, nebst einem Beitrage zur Flora der Mittelmark,’ and the first edition waa edited by Willdenow. The third edition appeared in 1806. Chamisso’s notice of E. capitata runs thua :-“ Erythrma capitata, foliis elliptico-lanceolatis obovatisque tri- et quinque nervibus ses- ailibus, floribus capitatis bracteatis.- Willd.” “ Species notabilis, inedita, ex herb. celeberrimi viri. Planta aemel sed copiose a phptopola adlata rursus haud reperta est. Latet locus natalis ”*

The original MS. ?f Willdenow’s short diagnosis printed by Chamisso aeems to be lost ; the plant is also wanting in Willde- now’s herbarium, as the specimens preaerved at Berlin belong to the “ Herbarium Generale,” which, together with a small number of plants, probably found after the death of Willdenow, had not been placed in hia herbarium. Dr. Ascherson thinks it probable that the label accompanying the specimens was written by Schlechtendal pater, who took care of bis friend

* I am indebted to Dr. Ascherson for this extract from Chamisso’s ‘Adnota-

(P. 9). i

tiones,’ no copy of which could I find in the English libraries. &INN. J0URN.-BOTANY, VOL. XVIII. 26)

400 YB. F. TOWNSEND ON AIV ERYTHRaA

Willdenow’s herbarium, and sent to Schultes mary observations and notes on ‘Reliquiae Willdenowianae.’ Rcemer and Schultes are therefore hardly correct in writing “ Herb. Willd.,” as they do in their notice ofthe plant in their ‘ Syst. Veg.’ Dr. Ascherson also thinks it nearly certain that Schlechtendal pater was the author of the longer description given by Rcemer and Schultes, which is as follows :-“ Caulis 2 poll., simplex, basi dense tectus foliis obovatis lanceolatis. . . foliis binis ternisve cincto. Bracteae lineares acutae, floribus intermixtae. Calycis laciniae acutae longitu- dine tubi. Corolla Centaurei sed laciniae acutiores, angustiores. Caulis saepe in superiore parte, immo ad ipsum capitulum, ramum unum alterumve emittit nudiusculum capitulo terminatum u t planta prolifera videatur.”--R.# 8., 8yst. Peg. vol. iv.p.786 (1819).

From want of’ access to specimens since Willdenow’s time, subsequent authors have unfortunately confounded his E. capitafa with a capitate variety of E. Centazcrivm. In no instance, in any of the extracts I am about to give, are distinctive characters given by which either Willdenow’s plant or that from Freshwater (both of which I refer to one and the same species) can be distinguished from a very different plant, viz. a capitate variety of E. Centaw ~iurn. It is certain that some of the extracts refer only to the latter variety.

“3. Centaurium . . . . variat p. capitafa corymbo etiam post an- thesin compacto, nec elongato : E. capifata, R. et S. Syst. Veg. iv. 163,D. fl. 2, p. 132. In Westph. u. Brandenb.”-Koch, Synop. ed. 2, p. 566. “E. Centaurium . . . . y capitata caule abbreviato, corymbo capitulo-contracto : E. capifata, Cham.”-Bluf 4 Pingephzcth, ed. 2, vol. i. p. 391%. “E. Centaurium . . . . aut caule abbreviato floribus dense capitatis : E. capitata, Charniss.”- Reich. F1. exc. p. 433. “ Specimina suecica [of E. Centalcrium] ad illam formam, qua? E. capitata, Rcem. & Schult. iv. p. 108, pertinent.”-Pries, Novitiae, ed. 2, p. 72. “Erythma Centaurizcm, p. capitata (R. & S.) faseiculata, Selm.”: “ F1. nornbreuses re’unies en corymbe serrk, compacte mhme apres la fleuraison. Coteaux maritimes : Granville, Cherbourg, Carteret.”-~r~bzsson, F1. de Normandie, ed. 4, p. 201. “E. Centazcrii forma abbreviata, 3. latgolia subsirnilis, at pecorum dentibus orta.”-,Grisebach, Gen-

it The author quotes G. L. E. Schmidt’s work entitled ‘ De Erythraea Disser- tatio inauguralis,’ in which E. capitata, Cham., is given a8 one of nine named varieties of E. Centaurium. Five of the varieties axe dgured, but E. cap’tata is not one of the number.

FROM THE ISLE O F WIGHT AND SOUTH UOAST. 401

tian. p. 140. “E. Centaurium . . . . var. y. E. capitata, Cham. : Der Stengel ist vom Grunde an starker gefliigelt-kantig als an der Stammart ; die Doldentraube bleibt auch bei der Fruchtreife sehr gedrungen, i d e m die Aestchen sich kaum verlangern ; der Kelch ist bei der geoffneten Bliithe meist urn die Halfte kurzer a19 die Bohre, und bei der Fruchtreife fast um das Dreifache kurzer als diese. Die Blume wie bei der Stammart . . . . die Abart y. im Pastoreiholze bei Liibke in Westphalen von Dr. Weihe gesammelt und mitgetheilt, auch baben wir sie von Dr. Wickstrom aus Schweden als E. Celztauriwm erha1ten.”-Xertens 4 KO&, in Rohl. Deutscb. Fl., ii. p. 233. “E. Centaurium, Pers. . . 0. E. capituta, Rcem. Plante courte, ramassee, & corymbe toujours compacte; feuil. rad. nombreuses, grandes, arrondies, obtuses, A 5-7 nervures. Rochers, pelouses sablonneuses, graviers de la region maritime, surtout de la cbte du nard."-Lloyd, F1. de l’ouest de la Fr., ed. 3, p. 204. “Erythrffia Centauriunz, L., var. capitata, Rcem. et Sch. On FreshwaterDownand in thewarren atAlum Bay, plentifully’’ *.-A. G. %ore, Supp. to the F1. Vectensis in Journ. of Bot. for 1871. [No characters are given.] Mr. A. G. More is the only English botanist who has applied the name “ capitafa” to English specimens of Erythrffia ; but evidence is wanting to show how far Mr. Nore distinguished the cupitute variety of E. Centa- r ium from the plant which is the principal subject of this paper ; for both grow abundantly in the localitiee named by Mr. More.

Mr. James Lloyd has kindly sent me specimens of his (‘E. Centaurium, p. E. cupitata, Rcem.,” from the Loire Infe- rieure ; and I have no hesitation in referring them to the capitate variety of E. Centaurium, and in considering them quite distinct from the Isle of Wight Erythrffia with free filaments. Mr. Lloyd also sent me a specimen which had been given to him as E. Cen- tairium, p. sufruticosa, Griseb. (Chironia sufruticosa, Salz.). I believe, with Mr. Lloyd, that this specimen must be referred to his E. capitata from the Loire Infhrieure, which, as I have just observed, is a capitate variety of E. Centaurium.

Since Willdenow’s time botanists have been unsuccessful in * Mr. More adds :-“ This is, I believe, the plant given in ‘ Flora Vectensis’

as E. liftoralis, which I have not succeeded in finding in the Isle of Wight.” The stations given by Dr. Bromfield in ‘Fl. Vect.’ are “Alum Bay, between Grove’s Hotel and the sea ; Headon Hill, within twenty yards of Mr. Ward’s cottage ; sea-banks near Compton.” Dr. Snooke, in his F1. Vectiana,’ records ‘ 1 Chironia littoralis,” “ Dwarf Tuf‘ted Centaury,” from “ sea-banks near Comp- ton” (Snooke,‘Fl.Vect.’ p. 12, 1823). Compton is east of and near Freshwater

2 a 2

402 XR. F. TOWNSEND ON AN ERYTHRBA

refinding his B. capitata, though they have frequently sought for it; it appears, therefore, that it is extinct. With the exception of this and the Freshwater plant and two specimens of a similar one from the neighbourhood of Newhaven in Sussex, I know of no instance of an Erythraa with similar characters. That these characters mark a distinct species I feel perfectly satisfied; and the description given in this paper will, I think, satisfy other botanists that I am correct in my conclusious. I n accordance with the generally received rules of nomenclature, it would seem that the English plant must receive the specific name " capitata $' for I believe both it and the German plant agree sufficiently to constitute them two varieties of one and the same species. Which is the type ? The Berlin plant has been only once found, and has not been refound for more than half a century, and is pro- bably extinct. The English plant has been found in two dis- tinct localities ; and in one of these a t least it exists in abundance at the present time: it is also probable that it has a still more extended range. Unfortunately the name " capitata '' has been variously applied, and by existing botanists is now invariably given to a variety of another species. To give the simple name " capitata " to the Isle of Wight plant would therefore, in all probability, either consign it to oblivion, or to the liability of its being confounded with the variety of E. Centaurium. I have therefore given varietal names to both forms.

Itremainsnow to give the specific character and description,and, finally, to point out how the species differs from its neighbours.

ERYTHRBA CAPITATA, Willd.-Caule ( f-3 poll.) plerumque simplici et solitario erecto subangulato, foliis rosulatis ovatis ovato-oblongis obovatisve spathulatis obtusis 3-5-nervatis, foliis caulinis paucis connatis angustioribus, floribus subfasciculatis in capitulo dense congestis sessilibus numerosis cum bracteis obtusis intermixtie, bracteis exterioribus flores squantibus vel superantibus, calyce corolls tubo squali, filamentis in imo tubi corolls insertis, stylo obliquo, quadrante parte ovarii sub anthesi exserto, capsula calycem excedente. 1 vel 9 . 0 G" I n pascuis apricis.

Caulis sirpe in superiore parte ramum nudiusculum unum alter- umve capitulo terminatum emittet.

* The only other notice I can find of any species of Bythrcm being biennial irr that given by Brotero in a long description of his Erythrcea chloodes, where he pays, ," radice &XUnIB, aut non rare biennis " (Brot. FI. Lus. i. p. 277).

FROM THE ISLE OF WIGHT AND SOUTH COAST. 403

Var. a. WILLDENOWIBNIL.-E. capitata, Willd. Bracteis acutis, laciniis calycis lanceolato-subulatis, laciniis corolla elliptico-lan- ceolatis acutis. Latet locus natalis.

Tar. p. S~H~ROCEPHaLA.-BraCteiS subobtusis, laciniis calycis lanceolatis, laciniis corolla ovalibus obtusis. On the downs of Freshwater, Isle of Wight, and Newhaven, Sussex.

Root fibrous, rigid, tapering, slightly branched, pale yellowish white. Stem usually simple and solitary, from & to 3 in. in height, erect, subangular, the angles formed by the decurrent margins of the leaves, and from the central nerve on the underside of each leaf, glabrous. Root-leaves numerous, forming a rosette, sessile, hori- zontal, flat or canaliculate, often curved, ovate or ovate-oblong, more or less spathulate, obtuse, entire, with three principal nerves reaching the apex of the leaf and forming three elevated ridges on the underside, the central nerve often forming an apiculus, glabrous, or with a few long hairs when young, darkish green, paler beneath j the broader leaves five-nerved at the base, but the exterior nerves not reaching the apex of the leaf. Stem- leaves few and in pairs, subconnate, narrower and often longer than the others. Plowers numerous, sessile, in a subfasciculate compact terminal head, formed of many bracteated imperfect cymes, the branches of which are suppressed, and the cymose character almost lost. From the outer bracts long-stalked secondary heads of flowers, which exceed the primary heads, are frequently produced, giving the appearance of the plant being pro- liferous. Bracts numerous, the outer leaf-like, often wider at the base, two or more of them equalling in length the head of flowers ; inner bracts gradually decreasing in size, linear ; larger ones obtuse-apiculate, smaller ones acute. Calyx about equalling the corolla-tube, cylindrical, with minutely serrulate angles, five- lobed ; lobes somewhat unequal, one-nerved, much longer than the tube, triangular-lanceolate, with narrow membranous borders, the border broadening below and sometimes minutely denticulate. Corolla-tube pale greenish yellow, hardly lengthening after flowering, equalling or shorter than the corolla-segments, which are oval, obtuse, concave, of a deep pink without, of a paler colour within. Under the microscope the pink colour on the back of the segmentsis seen to be produced by the pigment enclosed within very minute elongated cells. Stamens 5 , shorter than the corolla-segments ; filaments thread-lilie, flattish, attached only a t

July, Aug. Description of Elzglish form (@.-Annual or biennial.

404 ME. F. TOWNSEND ON AN ERYTHEBA

the base of the tube of the corolla, perfectly free above. Anthers yellow, oval, spirally twisted when empty. Ovary cylindrical, about & of its upper portion exserted beyond the corolla-mouth at the time of flowering. Style oblique, short, thread-like, cylin- drical. Stigma greenish yellow, expanding into two roundish flattish lobes. Capsule cylindrical (obtuse?), tipped with the style, imperfectly two-celled, about one third longer than the calyx, i ts lower two thirds invested with the enlarged marcescent corolla- tube. Seeds small, numerous, irregular in shape, subspheroid, angular, minutely reticulated, dark brown.-In England the plant grows among the short herbage of open chalk-downs and near the sea. It flowers in July and August, and is in fruit in September. It is abundant on the downs of Freshwater towards the east and west. There are two specimens in Mr. Joseph Woods’s herbarium, from ‘‘ near the signal-post a t Newhaven, Sept. 9, 1836 ; ’’ they are named E. lataflia by Mr. Woods. The only other instance of the probable occurrence of E. cupitafa var. spLrerocephala as an English plant has been furnished me by [the late] Mr. R. A. Pryor, who wrote that, in the Oxford Herbarium, he had examined some specimens, in which both he and Prof. Lawson believed the stamens to be inserted a t the base of the corolla-tube. These specimens were, I believe, from Somersetshire.

It may be presumed that the duration of the plant is both annual and biennial ; that it is biennial is proved by the following facts. Last season Mr. F. Stratton, a t rnT request, kindly col- lected seeds from Freshwater and sent them to Mr. H. C. Watson, who in September last informed me that he had sown them in pots, that they had germinated and formed young plants, but that they had not thrown up any flowering-stem. On hearing this it occurred to me that the plant might be biennial ; and early in October last I paid a visit to Freshwater in the hopes of finding seedlings from nature’s last year’s sowing. I was soon rewarded by the sight of numerous healthy young plants ir, sitzc, a dozen of which I carefully took up and potted. I have sent three of these to Kew, and three to the Botanic gardens of Cambridge. There is every appearance of these young plants having germi- nated many months since; the roots are strong, the leaves are well developed in the form of rosettes, but there is.no sign of any flowering-stem. I could find no young plants of E. Centaurim or E. pzclcltelb, both which species grow abundantly in the same place where I gathered E. cupitata vnr. spkomcephala. Without

F Tomsend dsl LINN SOC JOURN BOT Vor XVIII PL 15

Blam lith I zit ch imp

1-10 ERYTHRrEA CAPITATA,EUd YCW SP~XROCEPHALA,TOWT~S 11-12 E CENTA.lJRIUM,PePS

F R O N THE ISLE OF WIGHT AND SOUTH COAST. 405

further experiment, I would not assume that the latter is the only one of our native species of Erythrma which is biennial. Should it be ascertained that E. capitata var. sphmrocephala doas not stand alone in this peculiarity, there remain other characters by which it may be distinguished ; should the proof lie the other way, and it be found that E. capitata var. spharocephala is the only English biennial species, the character is a very marked one.

I would conclude this paper by pointing out that E. capitata var. sphmocephala may be distinguished from E. Centarizcm, var. capitafa auct., by its narrower leaves, longer and subobtuse bracts, the outer ones equalling or exceeding the flower-heads, by its densely capitate, subfasciculate sessile flowers, its shorter corolla-tube, which does not exceed the calyx, by its filaments attached only at the base of the corolla-tube, by its shorter capsule, only three fourths of which are invested by the mar- cesceiit corolla-tube, &c.

From E. littoralis, Fr., E. capitata var. spherocephalu may be distinguished by its broader and distinctly 3-nerved leaves, by its more humble growth, by its short corolla-tube, its filaments attached at the base of the corolla-tube, by its oblique style, its capsule not covered by the marcescent corolla-tube, by its roundish compact heads of flowers, &c.

DESURIPTION OF PLATE XV. Figs. 1 and 2. Erythraa capitata, Willd., var. sphmocephah, nob.

3. Another plant of the same in fruit, with one central and two lateral

4 and 5. The flower, magnified. 6. Corolla of the same, laid open to show the insertion of the filaments. 7. A small secondary flower-head in fruit. 8. Capsule of E. capitata, Tar. sphmocephala, magnified, four lobes of the

marcescent corolla having been removed, and one left, to show the capsule exceeds the corolla-tube.

main stems.

9 and 10. The stem-leaves, slightly enlarged in fig. 10. 11. Erythrea Cmtauriurn : the corolla laid open, to show the insertion of

the filaments ; one of these has been bent down. 12. A capsule of E. Cmtauriurn : one lobe of the marcescent corolla left

in position (the four other lobes having been removed), to show that the capsule and the corolla-tube are about equal.

Enlarged.