# The Journal of Indian Botany/Volume 2/June 1921/The Indian Species of Eriocaulon

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THE INDIAN SPECIES OF ERIOCAULON *
By P. F. Fyson, M.A., F.L.S.,
Presidency College, Madras.
The investigation of the Indian Species of Eriocaulon on which the following account is written arose out of the difficulty found in determining some of my own collections in South India, which led me to examine the collections in the herbaria of the Royal Botanic Gardens, Calcutta, and at the Agricultural Station, Coimbatore (Madras), and subsequently in those of the Peradiniya Gardens, Ceylon, the Forest College, Dehra Dun, and the Agricultural College, Poona, for types not found in the two former.
From an examination of the sheets in the Calcutta herbarium it was soon clear that other collectors besides myself have found the identification of species difficult. As far as India is concerned there are, if we exclude local Floras which have largely followed the F.B.I., only two works in which descriptions of the species are given : Hooker's Flora of British India, Vol. VI, (1894) and Ruhland's monograph in Engler's Das Palanzenreich (1903). The former of these is naturally now incomplete, being without species which have been founded since its date. The latter is not available to the ordinary botanist, and even if it were is, since it contains all the species of the world, too cumbersome for the collector. It is therefore thought that a revision of the Indian species accompanied by illustrations would perhaps be welcomed by collectors of this interesting but difficult genus.
It may not be out of place in this connection to note that the identification of a species from its published description alone is nearly always fraught with some, often with very grave risk of error; and that only by reference to the actual type sheet can certainty be attained. None of the species of this genus were, as far as I know, founded in this country, so that the actual specimens from which they were described are not here but in Europe. We have however in this country duplicates of many of the type sheets, and though the possibility of ​error in the duplicates is not excluded, it is not unreasonable to ac- cept them as identical with tha types. Kurz made, certainly, on a sheet in the Calcutta herbarium a note to the effect that the label must be wrong and that most of them seem to be wrong, but the latter part of the remark appears to me unduly pessimistic for with most of the sheets there is strong internal evidence of their correctness. There are however some species which I have not been able to trace, especially those founded by Ruhland, the types of which are per- sumably in Berlin. While therefore there are several forms which appear to be new species, they may not be so in reality. My new species are therefore advanced tentatively and with the object of avoiding the confusion which would inevitably follow a mistaken identification.
The illustrations given herein are mainly from photographs of her- barium sheets, and wherever possible, of the actual types cr co-types of the species. It is hoped that this will make identification easier and more confident.
In the descriptive portion I shall as a rule give references to the Flora of British India and Ruhland's monograph only, and shall omit all synonyms, since the ordinary collector does not need them. But for reference in herbaria I shall give in Appendix II the identi- fication marks and previous namings of all the sheets seen by me (practically all those in India and Ceylon) under the true species-name, as believed by me, and those interested will find the synonyms there. The usual enumeration of the collectors and their marks, in connec- tion with the distribution of each species, is (with this appendix) unnecessary and has been omitted.
Following the name and the references, I refer (in the usual place) to the type sheet or its duplicate whenever I have seen this. If the type has not been seen no reference is made as a rule to any sheet. My authority for the name will be found in the appendix.
I tender my most grateful thanks to Lit. -Col. Gage, I.M.S., at the Royal Botanic Gardens, Calcutta ; to Rao Bahadur K. Rangachariar of Coimbatore ; to Dr. Burns and Mr. R. K. Bhide of the Agricultural College, Poona ; to Mr. R. S. Hole of the Forest College, Dehra Dun, and to the Dr. Petch of Ceylon for very kindly lending me all the collections under their charge for examination ; and to Col. Gage in addition for lending me reference books. I am the more indebted to them since on account of military duties my work was so often and lengthily interrupted that I had to keep the sheets far longer than I had expected, the work being begun in the autumn of 1917. To Mr. Streenivasiah, my herbarium-keeper, I am also gratefully indebt- ed for much assistance, and he prepared;, nearly all the photographs, ​
Introduction.
The eriocaulace^: are a very distinct family of flowering plants which in one respect occupies among the Monocotyledons a position analagous to that of the compositae among the Dicotyledons The flowers are very small and aggregated into a head, which is enclosed at first and subsequently backed by a involucre of outer bracts. Unlike the compositae the flowers are unisexual, and are typical- ly complete in every other respect, that is they possess complete whorls of sepals and petals, and either two of stamens or one of carpels. The genus Eriocaulon with its three sepals, three petals, and three carpels in the female flower ; and 3 sepals usually united into a spathe-like calyx, a trumpet-shaped 3-lobed corolla and stamens in the male, comes nearest to the monocotyledonous type : but we find two-merous flowers in some species ; and in some, otherwise trime- rous, two sepals only, or two or fewer petals, obviously by reduction.
The genus was founded by Linnaeus in 1742, and subsequently placed by Kunth along with one or two related genera as a tribe of the order RESTIACE.E. Martius, reviewing this tribe in 1835, raised it to the rank of a distinct order, the Eriocaulonaceae, a name afterwards changed by Eichard to ERIOCAULACEAE. Koerniche wrote a mono- graph with very full descriptions and several new species in 1856.* Steudel gave short descriptions of all the known species in his Syn. PI. Cyperacearum in 1858 and other authors, notably Sir J. D. Hooker, have founded species in " Floras " of Ceylon, India, Tropical Africa and Brazil. In 1903 there appeared in Engler's Das Pflanzen- reich a monograph by Euhlaud of all the species known in the family, with several new ones founded by him. The number of species des- cribed was 420, of which 200 belonged to Paepalanthus found only in America, and 193 to Eriocaulon. The latter genus is distributed all over the warmer parts of the world, being found in America, Africa, Asia and Australia, and even in Europe as far west as Ireland. But although there are in India some fifty species, occurring over tho whole of South India as for North as Mount Aboo and the Central Provinces, and along the Himalayas from tho eastern end to Dehra Dun, there are no collections from the United Provinces or the Punjab. In India therefore excluding the Himalayas, the genus is confined to the tropics.
Most species of Eriocaulon grow exclusively in wet places, a few only fully submerged, and only one or two I believe in ground always dry enough to be firm. It has long been recognised that the conditions of water and marsh are much more uniform the world over
is a sameness about the vegetative parts which we do not find in dry land plants. Thus in Eriocaulon the stem is nearly always very short, little more than a flat disc, with the leaves all radical and narrow, and the flower-heads are carried well above the ground or the water- level on tall scapes. There are of course differences ; submerged species have linear leaves, and some have elongate stems ; some land species are hairy, most glabrous ; in some species there is only one, in most there are numerous scapes : but except for these the differences are mostly small and difficult to define. This sameness in the vegeta- tive parts is accompanied by a surprising amount of difference in the floral, on which therefore the separation of the species is of necessity largely based. But since the parts are always very small and need a good lens for their study the species are difficult to diagnose, and in most cases it is impossible to do so with the naked eye without con- siderable practice. Collecting therefore, for any one not thoroughly conversant with the species, is like pulling things out of a bran-tub, one cannot tell at once what one has got ; and the determination of the species afterwards, unless the descriptions are very clear and to the point, is difficult in the extreme. This is borne out by an exami- nation of the material in the Indian Herbaria. Thus six collections made by Meebold in Mysore and Coorg, from September 1897 to November 1898, of what is certainly one species, were named by him, E. sexangulare three times, E. truncation twice and E. trilobum once. Yet these three species should never be confused, and the plants col- lected did not as a matter of fact belong to any one of them. The name E. sexangulare I find wrongly given on 15 collectings in the Herbarium 'of the "Royal Botanic Gardens, Calcutta, belonging to seven species quite distinct from the true E. sexangulare of Linnaeus ; and E. luzit- laefolium to 13 collectings belonging to six species, four of them being among the seven just referred to ; and these are not exceptional cases ; the third column of the Appendix will show that 2 species have each been given names of 7 other species, 2 more names of 6 other species, 7 of 3 or more other species.
One result of this is that species have bean given a wider distri- bution than they are entitled to, and this has of course led many a collector to suppose that he may have a species which really does not exist in his area. Especially is this the case with E. luzulaefolium Linn, which has been reported as all over South India, from Khasia to Bombay, Madras, Malabar and Ceylon. But, unless the sheets which bear Wallich's number in the Calcutta Herbarium do so wrongly, the species is confined to Nepal, Assam, Bengal and Upper Burma. ​
The Arrangement of the Species.
As far as the Indian species are concerned, if we except the local floras founded on the F.B.I, there are, as stated ahove, only two modern works giving descriptions, the F.B.I, and Euhland's mono- graph of the whole family. Previous to this excellent descriptions were given hy Steudel (Syn. PI. Cyperacearum 1858) and Koerniche (Linnsea XXVII, 1854, pp. 577-692).
The species are arranged in these two works on entirely different plans. Hooker after separating the purely aquatic and submerged forms, divided the remainder according to the external appearance of the heads and the presence or absence of hairs on the receptacle. Ruhland on the other hand arranged the species according to the num- ber of parts in the flower, placing in his first section, which though he does not so identify it I take to be Naysmithia Huds., those with 2 parts to each whorl, in his second section those with 3 parts, and in his third those with 3 parts in the staminal and carpellary whorls but with fewer sepals or petals ; these sections being further divided for convenience into the eastern or old-world species and the western or new-world. He then took the nature of the stem whether disciform or elongated, with such characters of the flower as white or black anthers, crested or plain sepals, for the lesser divisions. The difference in the two systems is very great. Two plants classed by Hooker as dimerous and trimerous varieties of the same species, E. sexangulare, appear in Euhlands monograph, as also in Steudel's Syn. PI. Cyp., in different main divisions of the old world species, and in the former s list as numbers 25 and 186 respectively. The plant named by Trimen E. atratum Koerniche var. major was raised by Hooker to the rank of a species, E. caulescens, and placed next to E. robustum Steud. of the Nilgiris, from which it hardly differs except in having a tall and branched stem ; whereas Ruhland separated the two by no fewer than twelve Indian species, and placed E. robustum next to E. quinqucangulare Linn., which in Hooker's arrangement is separated from it by almost the whole of the Indian forms, one being No. 4 and the other No. 35 out of 43.
It would probably be correct to state that except in his main divisions Ruhland in fact did not attempt to arrange the species in phylogenetic groups, but only to provide a general clavis for aid in their identification. Hooker made tentative groupings, but apart from the separation of aquatic from terrestrial species made no definite sections.
Before attempting to classify the species of a genus it is clearly necessary to determine what characters if any are liable to vary with age or with the conditions of the environment, and further ​to estimate if possible the relative importance from the phylogenetic stand- point of the more stable characters.
Stem and Leaf.
As said above since practically all the Eriocaulons grow either in swampy ground or submerged in water the stem and leaves of any one species vary but little, even in size. At the same time all sub- merged forms are for a like reason so alike among themselves, and the swamp forms also among themselves, that such differences as exist are of little use in separating the species. The difference between the usual disc-like stem, and an elongate branching one, which Ruhland following Koerniche used almost at the fore-front of his clavis of old-world species, though at first sight it may seem very definite, is not always a hard and fast line, and is in any case proba- bly bound up with the robustness of the species and the nature of its habitat. As instance the Nilgiri E. robust am and the Ceylon E. caulescens referred to just above. But I have found the former with root-stock over on inch in length, and poorer specimens with leaves narrow enough to be indistinguishable from those of the Ceylon plant. In Ruhland's list one is No. 74, the other No. 120.
The Head and its Involucre.
As regards the heads Koerniche used the difference of hairy and glabrous involucre, as also did Hooker and Euhland ; and without doubt this character is of sectional value. But I find the hairiness varies and "may even be absent from a plant undoubtedly for other reasons allied to hairy species. This character must not therefore bo used two rigidly, as in Ruhland's wide separation on this account of E. Brownianum Mart, from E. nilagirense Steud. (Nos. 93 & 117 respectively.) The specimens in Herb. Calc- show plainly that the type sheet of the former is of a not fully undeveloped plant and that the absence of hairs is here accidental.
Characters which give very distinctive appearance to the head and would certainly appear at first sight of at least specific value are afforded, by the form and length of the involucral bracts. Thus they are horizontal and very obtuse and slightly turned up at the end in E. sexangulare, E. luzulaefolium, E. truncatum and E. Thioaitesii ; they are acute and ultimately reflexed in E quinquangulare and E. trilobum. In E. xeranthcmum, E. roscum, E. martianum Wall, and forms of E. Dianae collected in Coorg, they are very much longer than the head, projecting beyond the general margin like the rays of a sun-flower. (PI. fig. 12 & 13.) Koerniche placed great value on this. But a comparison of a large number of collections made on the
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pect, every intermediate form being found from large conical heads with short reflexed bracts to small flat heads with long bracts. It is clear therefore that, at least with this species, the relative length of the involucral bracts is of varietal rank only. But if this be so then perhaps it is also with other species ; and the peculiar Burmese plant E. Martianum Wall, should be considered a variety of E. quin- guangulare, not as either a distinct species or a proliferous of state {cp. Hooker (1) p. 582.) A similar but less pronounced lengthening of the involucral tracts appears in two other species, in other respects widely separated, E. gracile Mart, and E. cuspidatum Dalz. At the same time extreme variation of this kind should probably be consi- dered specific— e. g. in E. xeranthenum Mart. -and E. roseum, as also of course when accompanied by differences in the flowers {E. Ediuardii). But it is clear that the form and length of the bracts is not in itself of ' sectional ' ranks as Koerniche supposed.
The colour of the involucral bracts is some guide, as in the separa- tion of what I take to be E. atratnm Koern. from E. subcaulescens Hook, f., but it is an uncertain one, the black colour fading often out of these bracts with age. On the other hand the scarious, usually straw- coloured, floral bracts of some species are easily distinguished from the more common black with white hairs. The difference is definite, with no merging, and I take it to be of ' sectional ' value. Black bracts are usually hairy and scarious bracts glabrous, but there are exceptions.
Hairs on the receptacle are a constant feature of most species, but are partly or altogether absent from others ; and Hooker made use of this in the F.B.I. This difference accompanies others of ' sectional ' rank, but may occur inside the section and is I believe of only secon- dary importance. Another character to which Hooker gave some im- portance is the length of the pedicel (stipes.) I am not able to follow him in this. The length appears to me to depend on the age of the individual flower, and to be therefore of no importance.
The Flower , its Petals and Stamens.
It is in the flower itself that the greatest differences are found. In the male the chief distinguishing characters are white or yellow instead of black anthers, and one corolla-lobe being so much longer than the rest or to protrude beyond the floral bracts and cover them. The first gives an absolute line, not in any way bridged over by an occasional species with olive-green colour. Euhland stated (lc. p. 16) that the colour is constant in the species, but used this difference more than once in his clavis as if of only minor importance. Hooker ​did not notice it at all. Steudel and Koerniche were both aware of this difference, but neither paid any particular attention to it. Apart from the improbability of so very definite a change as the loss or acquirement of the black colour happening more than once, yellow anthers are so distinctive that I have no hesitation in making a ' section ' of the species which possess it. The protruding male petal is also a very distinctive character and one naturally used in any scheme of classification. It appears to be a very constant character, and nearly all the species which show it do so quite definitely. But as might be supposed indications of the habit are not wanting in other species, some of which may therefore be regarded as on the line of development. The stamens do not vary in number. They are always six, except in the one dimerous- flowered species.
In the female flower the ovary is invariably 3-Iobed (except of course in dimerous flowered species) — there is no reduction. The petals are 3 or 2, and only slight differences occur between those of dif- ferent species ; except in one direction. Asa rule they are oblanceolate, with thick terminal hairs and slender, longer, lateral ones ; but in some species there is a brush of slender filaments or hairs, which might be regarded as due either to the longitudinal splitting of the petal into many parts or to a narrowing of the petal accompanied by an increase in the number and length of the basal hairs. This latter change may well have come more than once, in different groups, I therefore do not use it as a 'sectional' characteristic.
The Sepals.
The sepals shows the most interesting variation. The simplest probably primitive, form is boat-shaped, black in colour and with a few hairs along the mid-rib or keel. In one Himalayan species the sepals are connected into a calyx similar to that of the male flower. The petals of this species differ from others in having the gland terminal and in being clawed. Ruhland has a group of Chino-Japanese species with these character, I therefore found my section CONNATO-SEPALAE to include them. In all other species the sepals are free. A development of the boat-shaped sepal is the formation of an enlargement (a thick- ening, wing, or crest) along the keel. In some species this takes the form merely of a thickening (E. Thoniasi PI. I. fig. 7) in others of a narrow wing or crest which may be lobed, or pectinate (figs. 5 & 6.) The depth of the thickening or crest appears to vary in the same species but there is usually no doubt about the crest when it is present. I therefore make a ' section ' of those species which possess this en- largement, whatever its precise form may be. Here again it may be that E. sexangulare, E. cuspidatum and E. Thomasi, are not derived
1697—19 ​from the same proximate stock as E. Elenorae, E. Margaretae, and E. minutum, through all show this character. The question could be decided only after examination of extra-Indian species.
These three types of sepal : the united, the boat-shaped and the crested, are quite distinct ; the only specific difference being that in the latter two the three sepals may not be equal in size, and one may be flat instead of boat-shaped, or without the crest. But in the species which show these exceptions individual variation does occur Thus in E, Elenorae the relative sizes of the 3 crested sepals varies, two may be of a size, one smaller, or the three all unequal, or one the smallest without a crest. These variations are shown in plants otherwise indistinguishable. They are therefore not of specific value. I am doubtful indeed whether E. Elenorae where the sepals are un- equal should really be separated from E. Margaretae (were they are all alike), but the species are slightly different in appearance, and I have found no variation in plants with the 3 sepals equal (E. Margaretae). So also in those with two sepals boat-shaped, one not. The odd sepal in E. Dianae may be lanceolate and as long as the other two, or short- er and bristle-like, or so slender that it is difficult to see.
We have in both these lines apparently a reduction in the size of one sepal till is nearly disappears, and if this be so a species with only two sepals may be derived from one with three. An instance where this has actually and unmistakably occurred even inside the species is afforded by E. Xeranthemum Mart. Hooker in F.B.I, gives the female sepals as 2. I find plants on the Himalayas have 2 sepals, but some at least on the Malabar coast have 3. Ruhland says that the sepals are 3, unequal. No one seeing the plants would wish to make several species of them. The same occurs in E. truncatum where the sepals may be 3, but are usually 2 only, and in the same plant I have found 3 equal, 2 and 1 smaller and 2 only ; and also in E. Thioaitcsii Hook. f.
We are thus faced by a set of conditions which must be unique among flowering plants. No other case is known to me of a reduction in the relative number of sepals and petals within the species of genus, though the number may be indefinite — e.g. in species of Banwncalus and Jasminum. The stamens certainly show a reduction in some genera (e.g. Cassia and Bauhinia), and occasionally between the genera of family (e.g. Caryophyllaceae, Acanthacae), but even this is not common. Characters in fact which in all other phanerogamic families are so constant as to be of the first importance in determining fami- lies and cohorts here vary even within the species, and so are of no use at all as guides to the phylogeny. ​
The Sections or Main Groups.
The following deviations from what might be called the normal or primitive conditions seem to be the most constant — hairy in- volucre, one male petal enlarged, scarious floral bracts, crested female sepals, united female sepals, and white anthers ; while the number of sepals or petals, 'in either sex, their relative sizes and shapes, the length of the involucral bracts and of the pedicels are not neces- sarily even of specific rank.
If this supposition be correct, and the evidence is I think too strong to doubt it, the Indian species fall naturally into eight groups distinguished by the above ' sectional ' characteristics, with a ninth composed of species or forms which grow wholly submerged in water and have elongated stems thickly covered for several inches with long capillary leaves. This group I name SETACEvE, from the chief and Linnaean species; not, following Hooker, Aquaticae, because there are other species with just as good a claim to that title. A character which like this is purely adaptative is not as a rule considered of much value in determining relationships, but the five forms which share this in common are so alike among themselves and so differ- ent in appearance from any other species that they must go together in any systematic scheme. The possibility is not excluded that they have sprung severally from the other distinct sections of the genus, but the fact that the male sepals are more or less free instead of being united into a calyx split at the back as with most of the other species, may indicate that they separated off early from the primitive stock. Of my other eight sections the ANISO- PETAL^ with one male petal much enlarged, and the HIRSUTiE with hairy involucre, correspond in part to groups in the F.B.I, of species with these characters but not so named. The LEUCAN- THERiE with white or pale yellow anthers, the CRISTAIO-SEPA- L^ with crested female sepals and the CONNATO-SEPALiE with the female sepals connate, found places in Ruhland's arrangement ; but, except the last of which there is only one species in India, were broken up in the different claves and included several species which I exclude. The SGARIOSiE with scarious floral bracts is I believe anew grouping; and I place together as SIMPLICES all species which do not show one or other of the above mentioned characters.
Geographical Distribution.
The geographical distribution of the species and groups presents many interesting problems of variation and relationship. As regards the extra-Indian species, the material at my disposal does ​not allow me to say very much ; but certain general conclusions may be drawn from an examination of the very full and careful descriptions of the flowers given by Kuhland in his monograph. It might be supposed at the outset that since the plants grow in water and marshy ground their seeds would be carried on the feet of migrating birds, and that this, coupled with the universally accepted similarity of the conditions of water and marsh tbe world over, would result in a very wide distribution of most of the species. Some certainly are scattered widely, but the majority seem to be confined to comparatively small areas. This question is of course bound up with that of the limits of the species : thus E. Sieboldianum Sieb ot Zucc, aj understood by Hooker in the F.B.I., occurs all over S. E. Asia from Bombay to Japan and N. Australia ; but Euhland separating from it several smaller species gives to them a much narrower distribution, though he retains almost as wide a one for E. Sieboldianum itself. Of the groups which I have proposed in this account the SETACE^j group has one representative, E. bifistulosum Van Huerck, in West Africa and probably others elsewhere. The jglMPLICES being all those with no special modification of the floral parts are no doubt primitive and world-wide. The HIRSUTE and ANISOPETAL^I are spread over S. Eastern Asia from Cochin to China, probably on the mountains of the warmer parts, and the latter seem to have a second centre of distribution in British Guiana. The CRISTATO- SEPAL^ also seem to have a centre in tropical South America, reaching from Mexico to Brazil. But the CONNATO-SEPALAE, which have in India only one representative on the Himalayas, belong almost entirely to China and Japan. Of the LEUCANTHERAE one species, E. Sieboldianum Sieb. et Zucc, is widely spread over tropical S. E. Asia, Malaya and Australia, but the others seem confined to India. E. Sie- boldianum is probably ."the most widely distributed of all the species, and E. Brownianum Mart, with its varieties (or related species of Euh- land) covers almost as wide an area.
Inside India there appear to be on the plains and lower hills no species at all north of a line from Mt. Aboo to Dacca, and not many northwards on the Himalayas, though there are one or two in Kash- mir. They occur all over South India. The hirsutae belong almost entirely to the mountains above 3,000 ft. of Burma, Bengal, S. India and Ceylon, but extend far southwards to Singapore. The ANI- SOPETALAE are developed chiefly in Ceylon, with one species in Ben- gal, one in the Central Provinces and the Deccan and another on the Niligiris ; but not curiously enough collected hitherto on the Palnis which are nearer Ceylon and floristically show closer affinities. Of the CRISTATO-SEPAL2E the smaller species belong to the Western ​
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Map 1. Showing the distribution in India of certain species of the section Simplices. ​Mountains from Mt. Aboo to Coorg, but do not go further north, east or south.
The group collinum-quinquangulare-trilobum-Dianae, has collec- tively the widest distribution, and shows very interesting develop- ments and cross-relationships. E. quinguangulare belongs to the plains of Ceylon, S. India and the Deccan, extending only rarely to the Western side in Canara. On the mountains to the south it is re- placed by E. collinum Hook. f. and to the north in Bengal by E. tri- lobum Ham,, both of which hardly differs from it except in the much darker head and better developed sepals, but are really quite easily distinguished. E. quinguangulare and E. trilobum have their counter- part on the wastern side in E. Dianae Sp. Nov. which in its widely differing varieties is similar to both, but differs in the reduction of one sepal to generally only a bristle and in the involucral bracts being usually longer. E. collinum has likewise in Ceylon one sepal smaller than the others, but the forms are not otherwise distinguishable. Both E. quinquangulare and E. Dianae show lengthening of the invo- ucral bracts, though the former only in Burma ; and E. xeranthemum with sometimes 3 female sepals sometimes 2, might be derived from either. It is in fact as if the two species E. quinquangulare and E. trilobum, and perhaps also E. collinum, were originally one, and developed from it as varieties and later on species, in — (l) the plains of S. India : and L. Bengal, (2) Upper Bengal, and (3) the mountains of S. India : and further as if these have independantly suffered a reduction in one sepal of the female flower ; the first two in travelling westwards across the Ghats to the sea, the third in crossing over to Ceylon. Another change was a lengthening of the involucral bracts, which seems to have proceeded independently in both E. quinquangulare and the derivative E. Dianae, as it has done also in other species.
The Mendelian would doubtless find in crossing a sufficient ex- planation of these double relationships, but it remains that the species as here defined occupy distinct areas and are found together if at all only on the borders of contiguous fields.
Reference to published Works.
F.B.I.— Flora of British India, by J. D. Hooker. Volume ; VI, (1894)
pages quoted in Arabic numerals : a species number given
thus, No. 3. Ruhl, — Die Eriocaulaceae by W. Ruhland in Engler's Das Pflanzen-
reich (1903). The serial number of the species alone is
quoted. ​ PI. Ceylon. — Handbook to the flora of Ceylon by Trimen and Hooker Vol. V (1900).
Cooke Fl. Bomb.— The Flora of Bombay by Cooke, Volume and page in Roman and arabic numerals respectively.
Fyson Fl. N. & P. H. T.— The Flora of the Nilgiri and Pulney Hill tops by P. F. Fyson. (Madras 1915—21). 3 Vols.
Koern. Linn. — Koerniche in Linnaea Vol. XXVII (1854), pp. 577-592.
Steud. Cyp. — ■Sfceudel in Syn. Plantarum Cyperacearum (1858).
References to Herbaria.
Herb. Bombay. — Herbarium of the Agricultural College, Poona, now at Ganeishkind.
Herb. Calcutta. — Herbarium of the Royal Botanic Gardens, Sibpur, Calcutta.
Herb. Ceylon. — Herbarium of the Royal Botanic Gardens, Peradiniya.
Herb. Dehra Dun. — Herbarium of the Forest College and Research In- stitute, Dehra Dun.
Herb Madras. — Herbarium of the Agricultural College and Research Institute, Coimbatore.
Herb. Presidency College, Madras. — Herbarium of the Presidency College, Madras.
Herb. Sedgwick. — Herbarium of Messrs L. J. Sedgwick, I.C.S. and T. R. Bell, now St. Xavier's Coll., Bombay.
Herb. St. Xavier. — Herbarium of St. Xavier's College, Bombay.
Herb. Talbot. — 'Herbarium of the late W. A. Talbot now at Ganesh- kind with Herb. Bombay.
Terms used in descriptions.
Pale* — of the involucre or the floral bracts, — an absence of black, usually resulting in the bracts being straw-coloured when dry, but sometimes light-brown, sometimes white. When fresh they are in some, perhaps in all cases, scarious and translucent.
* The word pale may appear hardly suitable as a descriptive term, but I know no other that fits the care so well. For the bracts so termed are characterised not by the presence of a light coloured pigment, so that they cannot truly be called white or yellow, but by the absence of the more usual black ; and being thin are translucent, but when dry straw-coloured. ​ Normal — of the flower — Sepals in the male united into a calyx split at the back ; in the female 3, equal and similar. Corolla of male trumpet-shaped with 3 lobes ; of female 3 oblanceo- late free petals. Stamens 6, anthers black. Ovary 3 celled.
Eriocaulon Linn.
Linn. Gen. Ed. II 35 (1743).
Scapigerous herbs, aquatic or on marshy, rarely dry ground. Stem usually very short and disui-form, but in some species elongate and branched. Leaves narrow. Scape slender, with 4 to 7 ribs, twisted usually, and eaclosed at the base in a sheath with oblique mouth. Flowers minute, eachj in the axil of a bract, in involucrate heads ; unisexual, nearly always monoecious ; perianth inferior. Male flower : — Sepals 2 or 3, usually, but not always, connate into a calyx split on the ad-axial side. Corolla mono-petalous, funnel-shaped ; lobes 3, small, usually ciliate and each with a large black gland. Stamens 6, attached to the corolla ; anthers black or pale yellow. Female Flower : — Sepals 2 or 3 flat, boat-shaped or crested, equal or unequal. Petals 3 to free, linear, oblanceolate or spathulate, ciliate, each with a black gland near the upper margin. Ovary three-celled ; style with 3 short branches. Carpels in fruit globose opening down the back to let out the one seed. Seeds translucent yellowish or brown, often with darker markings. Embryo minute, outside the horny endosperm.
Species about 190 in the tropics and rarely in temperate regions.
The above diagnosis is for the normal 3-merous flower. In one Indian species, the flowers may be dimerous with 4 stamens or a 2-celled ovary. But in the majority of species there are 6 stamens or a 3-celIed ovary, even though the sepals or petals may be reduced to 2 or 0.
Scheme of Sections.
[Note the female flower may have no petals in groups II, V, VI and VIII.]
A. Anthers black.
I. Setace^j. — Stems to over a foot in length. Leaves 3-6 in. Head 1/8—1/5 in.
than an inch long. Male sepals usually but not always united
into a calyx split on the side. i Female sepals free, boat-shaped or flat, except in VII. 1 Floral bracts usually black with white hairs on the back.
Receptacle villous except in II a. ​ II. Simplices.— Involucre glabrous. All floral parts in 3's. and
equal or one female sepal smaller or absent. (a) Receptacle ... glabrous. {!>) Receptacel ... hairy.
III. HIRSUTE— As II b, but involucral bracts hairy, or also the
scapes and leaves.
IV. Anisopetalje. — As II b, but one male petal much enlarged and
projecting so or to hide the floral bracts. I ! Floral bracts scabrid or puberous, pale : receptacle villous. V. SCARIOSAE. — Female sepals 3 or 2 narrow. Male flowers normal. VI. Cristatosepalae. — Female sepals strongly boat-shaped and crested on the keel or at least enlarged. Floral bracts pube- rous in some. Male flowers normal.
t I Female sepals connate.
VII. Connatosepalae. — Female petals clawed. Male flowers nor-
mal. One species only on Himalayas, remainder in China, etc. B. Anthers white.
VIII. Leucantherae. — Involucre black or pale. Floral bracts
usually dark 'with white hairs, but also light and glabrous.
General Key to the Groups.
I Plants entirely submerged, leaves linear or ribbon-like ... ... ... b«
Plants of wet ground, leaves more or less lanceolate ... •-- ... d.
(Stem leafy for several inches; heads 1/6 in. b.j or less ... ... ... ••• Setace^e.
vStem 0, or if longer and branched, under 3 in. c.
Anthers black, Ls. 1—2 in. (Khasia) E. gregatum or E. barba-
caprae. Anthers white or yellow stem ; ls. 2 in. or
more ... ... ... ••• LEUCANTHER.E.
c.i
, (Anthers white or yellow ... ... D°-
' ^Anthers black or greenish .t, ••• e »
1697-20 ​ (Floral bracts at least the lower, hidden by the •J projecting male petals ... ... ANISOPETAL^:.
e * j Petals enclosed, or not projecting conspi- 1 ciously beyond the bracts ... ... f.
/Heads white or grey by the hairs on the
otherwise dark floral bracts ... ... g.
Floral bracts greenish or brownish, puberous, very closely imbri- cated. B. Sexangulare, E. cuspidatum, or E. Thomasi.
Floral bracts black, glabrous ; female sepals
connate (Himalayas) ... ... E. ALPESTRE.
Floral bracts glabrous, yellowish or scarious. h.
a -i
[Whole plant hairy, or heads over 1/2 in. diam.
depressed ... I Plant glabrous or nearly so, heads globose or I ovoid
Hirsute. Simplices.
(Female sepals boatshaped, two at least with h.-j a wing or crest ... ... ... Cristato-Sepal^!.
' Female sepals boatshaped or flat ... ... SCARIOSJE.
(To be Continued)
Explanation of the Figures— (See p. 159.)
qninquangulare L.
do. trilobum Ham. caulesce?is Steud. Eleanor® sp. nov. minutum Hook Thomasi sp. nov. alpestre Hook. f. horsley-kundce sp. nov. Sieboldianum Sieb et Zucc.
do. do.
Diana sp. nov. roseum sp. nov.
Fig.
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Male flower of
E.
Fig.
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Female f
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Male
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Female
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Do.
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Male
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Do.
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Female
do.
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Head of
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Do.
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