JOURNAL . D ILE LINNEAN SOCIETY., BOTANY. . VOL, XLIV, LONDON: SOLD AT THE SOCIETY'S APARTMENTS, “BURLINGTON HOUSE, ` PICCADILLY, AV, : AND bY LONGMANS, GREEN, AND CO., ; EA WILLIAMS AND NORGATE, 1917-1920. a Dates of Publication of the several Numbers included in this Volume. No. 295, pp. 1-105, published November 27, 1917. 2511996: 107-172, » . May 22, 1918. SE o OPS, ES September 12, 1918. » 298, ., 993.802, „ — May 16, 1919. , 999, , 363-482, . — July 31, 1919. » 900, , 483-530 is March 18, 1920. (Index, Title, ete.) : PRINTED BY (c ER AND FRANCIS, RED LION COURT, FLEET STREET. momm CPI LIST OF PAPERS. Page ARBER, E. A. NgwzrL, M.A., Sc.D., F.L.S., and F. W. LawrrELD, B.A. On the External Morphology of the Stems of Calamites, with a Revision of the British Species of Calamophloios and Dictyocalamites of Upper Carboniferous: Age. (Plates 28-25) 5555... meer 507 Bower, Prof. Freperick Orpen, M.A., Se.D., F.R.S., F.L.S. Hooker Lec- ture: On the Natural Classification of Plants, as exemplified in the Brisror, B. Murier, M.Sc. Ona Malay Form of Chlorococewm humicola (Nüg.), Rabenh. (Communicated by Prof. G. S. Wxsr, F.L.S.) (Plates17&18.) 473 Gares, Rrarxanp RuaGLEs, Ph.D., F.L.S. A Systematic Study of the North American Melanthacee from the Genetic Standpoint. (With Map (Plate 5), and 1 Text-figure.) ............ MERCURIO Jeu E 131 Gurry, Henry Bnovanaw, M.B., F.R.S., F.L.S. Plant- Distribution from the Standpoint of an Idealist M ALL ees tueri PE 439 Hasrinas, SOMERVILLE, M.S., see PAursox, R. Herpman, Prof. WinLiaw. Assort, D.Sc., F.R.S., F.L.S. Sports RUNIANA.— ILI. The Distribution of certain Diatoms and Copepoda, throughout the year, in the Irish Sea. (With 21 Text-figures.)......0.0.0000.000000. 173 Lacarra, CHARLES CARMICHAEL, F.L.S. Two Critical Plants of the Greek —— A Revision of some Critical Species of Hchiwm, as exemplified in the Linnean and other Herbaria ; with a Description of Echium judeum, a new Species from-Palestitigmne si o o euer 363 LawriELD, F. W., see ARBER, E. A. NEWELL. iv MacLzop, Prof. Jutrus. Quantitative Description of Ten British Species of the Genus Mnium. (Communicated by Prof. F. E. Werss, D.Sc., F.L.S.) (Aaa 9) Theocr DI duse cH aM M COUTE Paunso, Rogert, F.L.S., and SowEnviLLE HasriNGs, M.S. The Relation between the Alga and Fungus of a Lichen, (Plates 21 & 22.) ......... PucsuEv, HERBERT WiLLIAM, B.A. A Revision of the Genera Fumaria and Rupicapnos. (Communicated by A. B. Reypux, D.Sc., F.R.S., Sec. L.S.) (ID bic OI ve od p PNE cC RENDLE, ALFRED BARTON, M.A., D.Sc., F.R.S., Sec.L.S., see Puasuey, H. W. SanwoN, Cuarues Epaar, F.L.S. A Hybrid Stachys. (With one Text- EE ii UE EU E a dgrnsra rear Scorr, DukixrIELD Henry, M.A., LL.D., E.R.S., F.L.S. The Heterangiums of the British Coal Measures. (Plates 1-4.) 0... inl nie Notes on Calamopitys, Unger. (Plates 6-8, and 1 Text-figure.) ......... Srorzs, Marre CARMICHAEL, D.Sc., Ph.D., F.L.8. — Jennettites Scottit, sp. nov., a European Petrifaction with Foliage. (Plates 19 & 20, and 4 Text- uiseplty ssp rp DM nee TI» Wess, Prof. Freperick Ernest, D.Sc., F.R.S., F.L., see MacrEop, Prof. J. West, George Sreruey, M.A., D.Sc., F.L.S., see Biusror, B. M. Page 233 357 EXPLANATION OF THE PLATES Heterangium shorense, and H. tiliwoides. Heterangium Lomaxii. Heterangium shorense, and H. tilieoides. Heterangium tilieoides, H. Lomazcii, and H. minimum. Distribution of North American Melanthacem. (Map.) Calamopitys americana. Calamopitys annularis. ' Calamopitys Saturni, C. fasicularis, and C. Beinertiana. Flowers of Fumaria and Rupicapnos. Fumaria Ballii, sp. nov. Fumaria berberica, sp. nov., and F. montana, J. A. Schmidt. Fumaria coccinea, sp. nov. Fumaria australis, sp. nov. Rupicapnos muricaria, Pomel, and X. pretermissa, sp. nov. Liupicapnos longipes, Pomel, and R. sublevis, sp. nov. Rupicapnos surcocapnoides, Pomel. 17,18. Chlor humicol 19, 20. Bennettites Scotti. Cladonia digitata, and C. pycidata. Cladonia digitata. 23-25. Calamophloios and Dictyocalamites. ERRATA. [Norr.—Slight errors have been corrected in the Index.] Page 142, line 29, for Juss. (/.e. D. Juss.) read Moehring. 147 , 13 , pumillia read pumila. 168 ,, 5 from bottom, for Wats. read S. Wats. 373 „ 9, for macanthum read macranthum. i : 981 , 15 , specimems read specimens. j 383 , 24 , L.suppl , L. fil. Suppl. 898 ,, 19., spicat » Spicat ; a 408 , 37 , Antlantica read Atlantica. 432 „ 7 , ramosum read ramosius. 1 j i | THE JOURNAL OF THE LINNEAN SOCIETY. (BOTANY. Quantitative Description of Ten British Species of the Genus Mniwn. By Professor Junius MacLEop, University of Ghent. (Communicated by Prof. F. E. Weiss, D.Se., F.L.S.) (9 Text-figures.) [Read 1st June, 1916.] INTRODUCTION. WHEN we give a specific description of an animal or a plant, we use terms, and an extensive terminology has been created for the description of the characters of living beings. : The leaves of a certain species A are, for instance, described as oblong, elongate, with numerous marginal teeth and a short. petiole. "Those of species B are oval, short, with distant marginal teeth and a longer petiole. So long as we compare the two descriptions in a book the difference between A and B is distinct, but when we look at the objects we begin to hesitate. The words are vague: what is, for instance, the difference between a short and a longer petiole ? Moreover, we are very often puzzled by the variation of the characters. We may find in A leaves which are really oblong and other leaves which are rather oval, and in B we may find some leaves which are oblong or oboval and petioles of very different lengths. We are told, for instance, that the pronotum of a certain species of LINN. JOURN.—BOTANY, VOL. XLIV. B 2 : PROF. JULIUS MACLEOD ON TEN Carabus is longer than broad, and that this property is quite characteristie of the species under consideration. Measuring the length and the breadth of a dozen specimens, we find that the pronotum is always distinctly broader than long! The author of the description has been deceived by an optieal illusion, the system of curved lines which limit the pronotum producing a false appearance. Similar examples are to be found in thousands. Our method of describing animal and vegetable species has made little progress since the days of Lamarck and De Candolle. It is true our descriptions are more complete than in former times, attention being paid to characters which were unknown or neglected a century ago, but the way in which the characters are described has been little improved. The defects of the method are more seriously felt in proportion as the number of the known species becomes greater. The result of this state of things is that it is often very difficult—in fact, impossible—to identify a specimen, even by means of the best books. Again and again species already known to science are described afresh under new names, because it was impossible to recognize them in the previously pub- lished descriptions. Notwithstanding all this, the work accomplished by systematic botany and zoology is a very valuable one; tens of thousands of species have been exactly diseriminated, for the most part by a sort of instinct, which enables the experienced systematist to recognize the species by their facies. The masters of descriptive science possess this instinct to an astonishing degree. The difficulty begins when a specific description is to be made. Is it possible to adopt a better descriptive method, whieh would render the valuable knowledge collected by the systematists more exact, and make the inventory of living nature more useful? Is it possible to describe and to identify an animal or a vegetable species by means of numbers representing the values of the specific characters ? * Such a quantitative method is used for the description of inorganic objects. When we want to describe a specific mineral or a chemical substance (for instance, quartz, sea-salt, copper, sulphate, water, etc.), we measure a certain number of its properties, such as density, indea of refraction, coefficient of expansion, angles of the crystalline form, etc. The measurements are made according to certain conventions with reference to temperature, pressure, and other conditions of observation. The description consists of figures (so-called constants). * In deference to customary usage, I have used throughout the present communication the word “character ” in speaking of the various features the measurements of which give us specific differences. I should have preferred the term “property” as emphasizing the analogy between the specifie characters of living organisms and the various properties of inorganic objects. I hope to develop this point of view more fully in a subsequent publication. BRITISH SPECIES OF THE GENUS MNIUM. i 3 I have tried to apply a similar method to the deseription of living species by measuring 38 characters of about 90 species and 20 varieties of the genus Carabus. For each property I have determined the minimal, median, and maximal value in each species. The figures, set in order in tables, enabled me to describe and to identify the species and varieties more accurately than by the usual method of description. Unfortunately, the war prevented me from finishing and publishing my work. I have tried to carry out similar work with plants, in the Cryptogamic Laboratory of the University of Manchester. On the suggestion. of Prof. Lang, I took mosses of the genus Mnium. I have measured fourteen different characters of 10 British species of this genus (altogether more than 30,000 measurements have been made). Four other species of Mniwn belong to the British flora, but they are very rare in these islands and could not be measured for want of material. I hope to publish later a more extensive work on the theoretical part of this quantitative method. In the present paper I confine myself to a quantitative description of the ten studied species, giving only the theoretical indications which are necessary to make the given descriptions com- prehensible. I read a short abstract of my work at the Meeting of the British Asso- ciation for the Advancement of Science, in Manchester, on September 7th, 1915*. I may perhaps be allowed to reply here in a few words to some remarks which were made on that occasion. The first criticism was to the effect that I was using a rather complicated method for the identification ot species which are easily distinguished by every student of British mosses. I give the deseription of British species of Mnium as a first example of the application of the quantitative method, because I wanted to begin with well-known, easily recognizable species, in order to put the method to the test and to establish a starting-point for further application. . Secondly, it was said that the use of the method is too laborious to allow of an extensive application. It is true that the quantitative description of a certain number of species is a long and rather tedious task, but this is not a sufficient reason to cause us to be discouraged. When the descriptions and the tables of identification are made (once for all), the measurements which are needed for the identification of a given specimen of Mniuin may be made in less than an hour, and for the use of the tables no more than about ten minutes are needed. Let us remember that the exact measurement of one single physical constant is also a delicate and sometimes a very long task. * “On the Expression by Measurement of Specific Characters, with Special Reference to Mosses,” Report of the Meeting of the British Association for the Advancement of Science, held at Manchester in September 1915. B2 Lei art pee. ed ED aa MONUI M x4 rog i Pag caus 4 PROF. JULIUS MACLEOD ON TEN Despite this fact, innumerable constants have been determined by physicists and chemists. In a modern Dictionary of Chemistry we find hundreds of thousands of numbers which have been calculated during more than a century. Many of them (for instance, the atomic weight of the elements) were at the beginning only approwimate and have been corrected later on. It is certainly possible to carry out a similar work with animals and plants. My address seems to have made on some persons the impression that I am complicating things which are rather simple. In realitv, the method enables us to kace and to surmount difficulties which actually exist and are ordinarily overlooked. When we are told, for instance, that in Mnium rostratum the cells of the leaves are larger than in Mniwm serratum, this information is on the whole correct. But we may find specimens of rostratum the cells of which are really smaller than those of certain specimens of serratum! Such facts bring about hesitation and doubt. When we say that the breadth of the cells at the place of the yreatest breadth of the leaf, in the longest leaf of a fertile stem of M. serratum, varies between 16 and 26 p, and that the same character varies in rostratum between 22 and 36 p, we have replaced, it is true, a simple notion by a complieated one, but the given information is complete. The student is no longer disconcerted by the disagreement between the descriptive text and the faets he is observing. The material used for obtaining the measurements that follow was for the most part obtained from the Barker collection of British Bryophytes. This herbarium was given to the University of. Manchester by the late Thomas Barker, Professor of Mathematies, who was the founder of the Chair of Cryptogamie Botany in the University of Manchester. A few specimens were taken from the Manchester Herbarium. A number of rare specimens were kindly given to me by Mr. H. N. Dixon, M.A., F.L.S., to whom I offer here my best thanks. To Prof. W. H. Lang, in whose laboratory the present work was carried out, and to Prof. F. E. Weiss I am deeply indebted, as they made this work possible for me, and I wish to express to them my gratitude. I beg Prof. G. Unwin, who kindly helped me with reference to the language, to accept my most sincere thanks. "T BRITISH SPECIES OF THE GENUS MNIUM. 5 PARTE GRADATION. Variation of a Given Character along a Given A«is.— Variation within the Limits of One Specimen. $1. DrrricuLTIES.— When we try to apply the quantitative method to the description of an animal or a vegetable species, we meet two serious difficulties. The FIRST DIFFICULTY depends on the individual variation. The great majority of the characters of animals and plants are variable within the limits of each species, and the differences between several individuals of the same species are often great. When we measure, for instance, the length and the breadth of the pronotum (prothoraz) and the elytra of a dozen Beetles (Coleoptera) between which there is no specific difference, we see that they are all different from each other in the four characters under consideration ; and one who makes for the first time such a series of measurements is astonished by the importance of the observed differ- ences, although the measured specimens seemed to be, at first sight, almost identical. The figures seem to vary in a most capricious way, and to be useless for the description of a species or the identification of a specimen. In the description of living beings, quantitative data have been used only in those rather exceptional cases in which a character is not (or only very slightly) variable (examples: number of teeth of the peristome of the Mosses, number of stamens and styles in many flowers, etc.). We shall see further (Part II.) how it is possible to use even such VARIABLE figures for description and identification. The SECOND DIFFICULTY occurs especially in the Vegetable Kingdom and in polypiform animals; also to a certain degree in other animals. It is the result of a peculiar sort of variation which is quite distinct from the individual variation (or variation properly so-called), and may be called GRADATION. Example: Let us suppose that we want to compare the length of the leaves 1n two species of plants, A and B. Enormous differences are almost always observed between the leaves of each specimen. In one fertile stem of Mnium orthorrhynchum, with 53 leaves, the shortest leaf had a length of 046 mm., the length of the longest leaf was 3:91 mm. In a similar stem of M. undulatum, with 51 leaves, the length of the longest leaf was 8:97 mm., that of the shortest leaf 1:06 mm.* What answer can be given * In a branch of Ulmus montana with seven leaves the length of the longest leaf (petiole +limb) was 15:5 cm., that of the shortest leaf 6 cm. I ARR eee. ee WE WIN qe v NUNT e OMNEM. SS ST CNET aa LIII 6 PROF. JULIUS MACLEOD ON TEN when we are asked what is the length of the leaves of a specimen of Mnium orthorrhynchum or Mn. undulatum? To find the answer, we want to establish the conditions under which the measurements are to be made in order to obtain comparable figures. In other words, when we have measured a leaf / belonging to a specimen of the species A, we want to find in a specimen of the species B a leaf which is comparable with l. § 2. Grapation.—The way in which it is possible to solve this problem may be indicated by the study of an example. We take the Mosses of the genus Mnium. All the species belong to the ordinary type of the Acrocarpie Mosses. In a complete specimen we find fertile stems (which bear a fruit) and, according to the species, erect and horizontal sterile branches, male branches, ete. The leaves of all those branches are more or less different from each other in all the specimens of a given species. We agree, therefore, to limit our measurements to certain definite and restricted features, in the same way as we do when we measure the properties of inorganic objects. We select the fertile stem, and exclude all the other parts of the plant. The fertile stems may be considered as comparable in all the specimens and all the species. When we measure, for instance, the length of all the successive leaves of such a stem from the base to the summit, we see that the character length increases up to a maximum and then diminishes (see Table I.). TABLE I. Mnium punctatum, Hedw. : a fertile stem with 10 leaves ; length of the successive leaves in mm. Leaves: 1. 2: B: 4. 5. 6. rA 8. 9. 10. Length .... 0°97 1:31 QAUM ID. OUI. 0OUA TIS - :12- 689 246 Here we have a first simple example of gradation. GRADATION IS THE VARIATION OF A GIVEN PROPERTY ALONG A GIVEN AXIS. In this case the axis of gradation or line of gradation is the longitudinal axis of the stem which bears the measured leaves. We know that those leaves have been developed successively in acropetal order ; therefore the whole system of ten leaves represents a period of growth, and at the same time a period of gradation of the character under consideration. By means of the figures in Table I. we may construct the Gradation Curve of one specimen (specimen curve), in which the axis is represented by a horizonial line, the length of the successive leaves being represented by equidistant vertical ordinates (fig. 1). A specimen curve is merely a graphic representation of facts, without any calculation. BRITISH SPECIES OF THE GENUS MNIUM. 7 -7 mm J -6 Q x} T 2 i 2°53 ^ — > 8 8 6 Fie. 1.—Mnium punctatum. Gradation Curve of the length of the leaves of one fertile stem, o, Base of the stem; v, Summit of the stem; 1, 2, .. . 10, successive leaves. Scale in mm. See the figures in Table I. ” At first sight a gradation curve * recalls by its ascending-descending form a variation curve properly so-called ; it is, however, something quite different. In the ordinary (individual) variation, the differences between the individuals depend on cHaxcE. A variation curve properly so-called represents the distribution of tl e observed values of a given property among * Percy Groom, * Longitudinal Symmetry in Phanerogamia,” Phil. Trans. Roy. Soc. London, ser. B, vol. cc. (1908) pp. 57-115, with numérous figures. Percy Groom measured the length of the successive internodes of the stem in a number of Phaneroganis ( Rhinanthus, Chenopodiacer, Boraginacec, Solanacer, Lysimachia, Scrophu- laria, Ampelopsis, etc.). In his paper one finds numerous internode-curves which represent the variation of the length of the internodes along the axis, in the same way as our fig. 1 represents the variation of the length of the leaves. By the study of what I call Gradation, Percy Groom was brought to interesting morphological conclusions—among others, “ that the alternate-leaved Chenopodiacee have a phyllotaxis transitional between alternate and opposite, and that opposite phyllotaxis is primitive in this family.” (Loe. cit. p. 65.) From this example one sees that the study of gradation may render good service in a field of research quite different from the subject of the present paper on Mosses. B 8 PROF. JULIUS MACLEOD ON TEN a certain number of comparable individuals. The form of such a curve is governed by the laws of chance (Quetelet, Galton). The Gradation (variation along an axis) does not depend on chance, but on a certain cause (force or factor) which varies regularly all along the axis according to a certain law which is quite different from the laws of chance. In other words, in our example (fig. 1), the length y' of a given leaf 7 depends on (is a function of) the distance «' between the base of the leaf | and the origin o of the axis*. Chance has only an influence in producing slight irregularities of the curve, which are easily discerned when the curve is drawn. In fig. 1 such irregularities are less conspicuous than in the majority of the gradation curves (fig. 2). $3. ImpoRTANCE OF THE LONGEST LEAF.—In the fertile stem of all the species of Mnium the gradation curve of the length of the leaves is in a certain degree variable, but on the whole similar to the example represented in fig. l. The longest leaf of a given fertile stem (corresponding to the summit of the curve) is therefore comparable with the longest leaf of any fertile stem of any species of Mnium whatever (see $ 12, Remark II.). $4. SIMULTANEOUS GRADATION OF SEVERAL CHARACTERS ALONG THE SAME AXIS.— When we measure a second charaeter (for instance, the breadth of the successive leaves of a fertile stem) we obtain a second gradation curve.— Example: The gradation of the character length and the character breadth of the leaves of a fertile stem of Mnium serratum ; see Table II. and fig. 2. Tipun Lb Mnium serratum, Schrad. : a fertile stem with 27 leaves ; length and breadth of the suecessive leaves in mm. Leaves: 15 2. 9. 4. 5. 6. pa 8. 9. ength 5. ET 2:00.86. 2°06 2:15 2:28 3242 2:44 9:44 Breadth .... a m f TOG CIO sol. DPI. 118. 115 Leaves: 10. BE 12: 13. 14. 15. 16. 17. 18. 264 2836 302 295 3819 309 3824 3:40 Length ...... OYA 2 Breadth... 1:36 1:27 1:35 1:34 1:20 i 140. 1:52 1: Or Leaves: 189. 90. 2]. 22. 93. 24. 25. 26. 27. Length ...... 29:99. 350. 905. | 9:84, 468. 497. AV 402 .- 3°22 ibrendihie dece «1244. AD. 11:40. 1272, 1-24. 1:07, 080. 0:55 Here the two characters are independent of each other with reference to their gradation: the maximal length corresponds to leaf 25, the maximal breadth to leaf 17. * The relation between the value y and the distance x may be expressed by y=fv. The nature of f is hitherto unknown. — — — — e — o BRITISH SPECIES OF THE GENUS. MNIUM. 9 I have studied the gradation of twelve characters of the leaves in ten species : the figures are given in the description of the species, Part IV. The general conclusion is that in each species each character has its own gradation, different from all others. (There is, however, often a certain analogy between certain gradation curves. See Types of gradation, § 11.) nel HU Ex 5 10 a. 90 Fie. 2.—Mnium serratum. Gradation Curves of the length and the breadth of the leaves of one fertile stem. o, Base of the stem; x, Summit of the stem; 1,...5,... 27, Successive leaves; B, Breadth; Z, Length. See the figures in Table II.—In these curves the fortuitous irregularities produced by chance are more conspicuous than in fo ale $4«a. PERIOD or Grapation.—So long as I was acquainted only with the gradation of the length, I thought that there was a relation between gradation and the so-called grand period of growth, the development of the longest leaf synchronising with the period of greatest velocity of growth. But the faet that each character has its own independent gradation and that there is no coincidence between the highest values of several characters of the same stem, makes it very difficult to admit such a relation. The suggestion that a connection really exists between the variation of a character along an axis and the variation of the velocity of growth along the same axis, can only be accepted when we admit that each character has its own grand period coinciding with the summit of its gradation curve. According to this view, we should no longer speak of one grand period, but of as many grand periods as there are characters. Percy Groom, discussing the question of a possible connection between the grand period of growth and the normal curve of a main axis of a herb (gradation of the length of the internodes) has pointed out that : “What Sachs describes as the grand period of growth of a cell involves merely a kinetic conception dealing exclu- sively with the velocity or rate of growth of a cell. The proposition in question . . does not define or treat of the duration of growth or the ultimate dimensions attained at insu dci iia: VE Pp ior ae ee ee ea a eee Pe ee UIT 10 PROF, JULIUS MACLEOD ON TEN by acell.... The length attained by an internode, on the other hand, depends upon several factors: the original length of the internode, the mean velocity and duration of its growth." (Phil. Trans. Roy. Soc. ser. B, vol. cc. (1908) p. 96.) The question is a very complicated one (see Percy Groom, l.c. pp. 96-101). We do not need to discuss it further in the present paper. In any case, we may consider all the leaves which belong to one gradation curve as representing one continuous (uninterrupted) period of growth (period of gradation), abstaining from any reference to a possible connection between the form of the gradation curve and the variation of the velocity of growth during that period. $5. LEADING CHARACTER.—When we want to study the gradation of several characters along the same axis, at first each character must be studied separately. But, in order to obtain a general view of the whole, we may take one of the characters as a standard or leading character, all the others being referred to the standard. The choice of the leading character is, in each peculiar case, arbitrary. As a rule, we may take a conspicuous character, the gradation of which is distinct. I take in this paper the length of the leaves, the curve of which has ordinarily a prominent summit. §5a. We may represent the gradation of a given property in four different ways, viz., by means of 1°. A curve of one specimen (specimen curve): this is merely an empiric representation of facts. See $$ 2 & 4. 2°, An interval curve of one specimen (specimen interval curve). See $ 6. 3°. A mean interval curve of several specimens. See $ 7. 4°, The curves 1^, 2°, and 3° may be brought into the form of percentage curves. See $ 8. $6. INTERVAL CURVE OF ONE SPECIMEN.—In this paper I limit myself to that part of the stem which extends from the lowest leaf to the longest one. As the number of leaves is very variable, even within the limits of one species * (which makes comparisons diffieult), I divide this part of the stem into 10 equal intervals : having measured the value of a given character of all the leaves, I calculate the mean value of the character in each interval. The figures of each interval thus become comparable with the figures of the same interval in all the specimens and species. Moreover, as the leaves are brought together into groups, the small irregularities produced by chance disappear to some extent, especially when the leaves are numerous. Example : I take the length of the leaves of the stem of Mnium serratum, the figures of which are given in Table II. (fig. 2). From the lowest leaf to the longest one, the number of leaves is 25. divide this part of the * Examples.—Fertile stem of Mnium punctatum, 7-19 leaves; fertile stem of M. hornum, 46-64 leaves. BRITISH SPECIES OF THE GENUS MNIUM. bL stem first into 1000 grades. The leaf 1 (lowest leaf) is placed at 0°, the leaf 2 at 429 *, .... the leaf 6 at 209°, .... the leaf 25 (the longest one) at 1000°. I divide now the part of the stem under consideration into ten intervals, viz., interval 1 from 0° to 99°, interval 2 from 100° to 199°, .... interval 10 from 900° to 1000°. Table III. gives the number of leaves in each interval. TABLE III. A fertile stem of Mnium serratum, Schrad. : number of leaves in the ten intervals. Total number =25. Compare Table II. Intervals: 1. 9. 3. 2 5. 6. 7. 8. 9. 10. Number of leaves.. 8 2 8 2 9 3 2 3 9 8 I calculate now the mean value of the property length in each interval, using the figures given in Table II. See Table IV. TABLE IV. Interval curve of one fertile stem of Mnium serratum. Mean length of the leaves T in 10 intervals, in mm. Intervals: 1. 9. ioc 4. 5. 6. the 8. 9. 10. Meandonsth. 2.0176 2:10 . 288emibeeee oO. 58:05. 8373 Bal 377. 4:50 As the curve in Table IV. represents one specimen (fertile stem), I call it a specimen interval curve. § 7. MEAN INTERVAL CURVE.—We may, in a similar way, construct the mean curve of a given character of a certain number of stems which were developed under the same conditions of existence. The safest way is to take stems belonging to the same patch. Example : I have measured the lengths of all the leaves (114 in number) of 11 fertile stems of Mnium punctatum collected by Prof. Barker (Llanberis to Bangor, Wales). Table V. gives the number of leaves of each of them in each interval (see the method of calculation in $ 6). It must be observed that in several stems the number of leaves is smaller than 10 ; even in such cases the place of each leaf is calculated by means of the general method used in $ 9. * 1000: (25—1) = 417. T Leaf 1 was damaged and could not be measured (see Table IL.). Therefore only two leaves are taken into account in interval 1. ; E2 PROF. JULIUS MACLEOD ON TEN TABLE V. Mnium punctatum, Hedw. : distribution of the leaves of 11 fertile stems among 10 intervals. Intervals : T 9. nA 4. b. 6. y? 8. 0,7 IQ NAI... DEI: 4.1 4 3» 3 SN ee Eg 04 PEEL Y LESS E T$ à 13204 151 0 d» a o E Ec gu. Bop 31.1 4 1. 3 LB v EUST. £1 NR 9.9 cw £c cup Ea i1 9 14 1292 T 154 UNS EST. I i fu 359 Sos DICE E Tj ]1 1 LI ub d e a. ETC. 1 Pee a x oto r j 1 | d | a abs. E X 1 ‘een rey Me Tett sich | B xu 1 mH 8 19 1 ANLOP VG. ce Dividing in each interval the total length of all the leaves by the number of leaves, we find the mean length in each interval: this gives the mean interval curve of the whole group. See Table VI. TABLE VI. Mean interval eurve of the length of the leaves of 11 fertile stems of Mnium punctatum. Intervals: 1. 2. 3. 4. 5. 6. T. 8. 9. 10. Meanlength,in mm... 2:10 256 353 363 459 498 573 017 699 7:19 Id. reduced in °/, of the length in | 29 36 49 50 64 69 80 86 97 100 interval 10 ...... $8. PERCENTAGE OURVES.— Comparison between an interval curve of one specimen and a mean interval curve. A comparison between specimen curves of two or several stems * is only suitable when the number of leaves is the same (see $6); as this number is very variable, equality rarely occurs. Therefore we must have recourse to interval curves ($$ 6 & 7). Interval curves (specimen as well as mean curves) in which absolute values are given are all comparable with each other, because they have all the same number of ordinates. In such curves, however, the absolute values of the characters under consideration are of secondary importance ; the relative values of the successive ordinates, indicating the variation along the acis, are much more important (see also $ 14). Therefore we are allowed to bring the * Belonging to the same species or to difterent species. BRITISH SPECIES QF THE GENUS MNIUM. 13 interval curves to such a form that the absolute values are eliminated, only the relative values (proportions) being retained. This method enables us to reduce all the curves to the same scale; this makes the comparisons much easier. In all the interval curves we suppose the mean value of the 10th interval — 100, and calculate the value of the other intervals in hundreds of the 10th one. We call such eurves percentage interval eurves or percentage curves (°/) curves). Example: We want to compare a mean interval curve of a group of specimens with the curve of one of the specimens. In Table VII. we give the mean curve of the length of the leaves * of eight stems of Mniwm hornum (belonging to the same patch) and the individual curve of one of those stems taken at random, with the corresponding percentage curves. TABLE VII. Mnium hornum, Linn.—Length of the leaves : mean curve of eight stems and one individual eurve, with the corresponding percentage curves. Intervals: — 1. 2. 5: 0M Ub. 6. 7. 8. Oe 105 8 stems: length, in mm. .. 200 251 290 3:28 375 423 465 5:204 601 679 Percentage ...... 80 820014100 MS 55 - 02-69 - 77 89 100 l stem: length, in mm. .. 2°05 253 281 339 413 445 494 564 643 7:16 Percentage ...... 29 0OMENOON Qn e 58 02. GB “990; - 100 Comparing the percentage curves, we see that the individual eurve does not deviate much from the mean curve. This is an ordinary rule : in the great majority of the cases the gradation of a given character in a stem follows more or less approximately the mean curve of the group of stems to which it belongs. Therefore we may admit that a mean curve is not the result of an arbitrary method of computation, but that it has a real significance. A mean curve represents the most probable (approximately the most frequent) gradation in a group of stems. The differences between the specimen curves are very likely produced by chance—in the same way as the ordinary individual variation. $9. PERCENTAGE CURVES (continued).— Comparison between two species with reference to the gradation of the same property. Let us compare, for instance, the mean percentage curve in Table VII. CM. hornum) with the similar curve in Table VI. (M. punctatum). The difference between both curves is obvious—too great to be explained by chance. In the intervals 7, 8, 9 the percentage values are much higher in punctatum than in hornum : the result is that the summit (interval 10) is distinctly more prominent in hornum. (See also Table IX.) * 340 leaves distributed among 10 intervals. 14 PROF. JULIUS MACLEOD ON TEN $ 10. PERCENTAGE CURVES (continued).— Comparison between several cha- racters of the same species. Table VIII. enables us to compare the gradation of four characters of the leaves of M. hornum by means of their mean percentage curves (eight specimens belonging to the same patch). The absolute values are given in the descriptive table of M. hornum. TABLE VIII. Mnium hornum, Linn.—Mean percentage curves of the gradation of four characters. Hight stems (340 leaves). See fig. 3. Intervals: 1. 2. 8. 4. 5. 6. T. 8. 9. 10. IDSIOUR d uus B BER AB dB. DD 09 00. 7/7 89 100 Toys vcl c DD MEM H0 2584 09. 4^ — 94. 107. 111^:120^ 191 —TO0 Number of cells at the place of the greatest | 80 5:05... 70 00- 108 114. 1190- 194 :194 . 100 DISSE S ose ck. ] Breadth of those cells...... 67 73 78; 385 ..-80 94 94 94 100 100 ie E VA N/K Ty f D 0 A E L 22 X wae 8 7.93 35 19 Fro, 9.— The curves represent the figures in Table VIII. Z, length of the leaves; Bi, breadth of the leaves; Ne, number of cells; Be, breadth of the cells. We see here (fig. 3) more examples of the independence of the gradation curves; the curves L, Bc, and Ne are, indeed, quite different. The curve B? is interesting. The character Bl (breadth of the leaves) depends on the whole on two factors: the number of cells (in the transverse direction) and BRITISH SPECIES OF THE GENUS MNIUM. 15 their breadth. In the intervals 1-6 the curves Ve (number) and Be (breadth of the cells) are both ascending; the increase of Blin those intervals depends on the two mentioned factors. In the intervals 6-9, the increase of Be being less rapid, the influence of this factor on the increase of £7 becomes smaller and this increase depends almost exclusively on Ve. In the intervals 9-10 Be is invariable ; therefore the variation (negative increase) of B? depends merely on the variation of Ne. The relations between B? and the factors Be and We ave thus variable all along the axis. Moreover, those relations are not the same in all the species. If we try to study those relations by taking a number of leaves at random, we may be brought to contradictory results, and general conclusions drawn from such fragmentary observations may be rather illusory. It is merely by the study of the gradation curves in a sufficient number of species that we may hope to discover general rules governing the relations between the dimensions of an organ, the number of its constituent cells, and their dimensions *. We see from this example that the knowledge of the gradation may have a certain importance for the solution of certain physiological, and morphological problems—apart from its significance for the study of species and their variation. $11. Types or GnApATION.—The gradation curves of twelve characters of the leaves of the fertile stem in ten species of Mnium may be brought roughly under five types :— Type I. : The character under consideration increases from the lowest leaf to the longest one, and decreases beyond this. This is, of course, the case with the length of the leaves (leading property) in all the species, and also in some other cases (examples: figs. 1 & 2; L and Bein fig. 35 Mn. rostratum and subglobosum in Table LX.). Tyrer II.: The character, incréasing from the lowest leaf, reaches its maximum (summit of the curve) below the longest leaf, the position of the maximum being very variable. See, for instance Ve and Bl in fig. 3. More examples are given in Table IX.; in this table the summit of the curve of the same character (breadth of the leaves at their base) has a different position according to the species. Two species have their summit in interval 10: these belong thus to Type I. The eight other species (hornum is doubtful) belong to Type lI., the summit being situated below the longest leaf. A comparison between the curves in Table IX. is the occasion of some interesting remarks. We see, for instance, that in two species (rostratum, subglobosum) the longest leaves have the broadest base, but in four other species (afine, cuspi- datum, undulatum, punctatum) the longest leaves have the narrowest base. In subglobosum the shortest leaves (intervals 1, 2) are much narrower at their base than the longer ones (intervals 9, 10) ; in punctatwm we see rather the inverse. This is interesting, because these two species resemble each other in many other characters. * See Percy Groom, Phil. Trans. Roy. Soc. London, ser. B, vol. cc. (1908) pp. 96-98. g } 16 PROF. JULIUS MACLEOD ON TEN TABLE IX. Mean percentage curves of the breadth of the leaves at their base in ten species of Mnium. The absolute values (in mm.) are given in the descriptive tables of the species. Hach species is represented by one patch. Intervals: 1. 2; 3. 4. 5. 6. ae 8. 9. 10. mme a aan 117 4128 — 18D 131: 124 .127 .119 104. 97. .100 cuspidatum ........ E073 51012774189 188 $5132 25197. Loe. 116. 08: 100) onum. iei 59 67 73 81 87 90 92 96: 101. 100 orthorrhynchum .... T$ 84 92 96 104 | 192 .194 118 105 100 undulatum ........ NOS 10 028 — 145^ 149^ /185 184^ 199 — 114 100 NOTA est 65 70 68 73 70 71 77 78 90 100 BODIORHA Sites 40s) ts DO oona a 122%, 122 :-118 : 108 108-- 100; 100 punctatum.......... 105 11]. 04 -.118: 114-109. 112.110 4-106. 100 subglobosum ........ 72 83 94 94 89 95 88 92 98 100 TEDQUM TTL EN RE 91 97 Bo 108: 108- 014 . 115.110 — 100^ 300 Type III. : In some curves the increase is rapid in the first intervals ; when a certain value is reached it remains constant or almost constant in the next intervals, the curve assuming the form of a horizontal line—LZwamples : The number of cells at the place of the greatest breadth of the leaves in M. undulatum (almost constant in intervals 4-10) and also M. spinosum (intervals 4-8). See Table X., in which the curve of a third species is given for comparison. TABLE X. Mean interval curves of the number of cells of the leaves at the place of the greatest breadth in three species of Mnium. Absolute values. Intervals: 1. 9. 3. 4. 5. 6. T: 8. 9. 10. OD Pen] HORS RU ra 63 71 76 91 92 89 91 90 89 89 PRED ORS AS aS 40 59 84 91 91 90 88 88 87 89 orthorrhynchum .... 28 82 37 40 46 58 60 60 52 42 Tyrs IV.: In some specimens and in some species a character has the value 0 (does not exist) in the first intervals.— Example : In the great majority of the specimens of Mnium hornum the teeth at the margin of the leaves and on the vein (at the back of the leaf) do not exist jn the lowest leaves. See Table XI. TABLE Al. Muium hornum.—Gradation of the number of teeth (1) at the margin of the leaves and (2) on the nerve. One stem ; interval curves. Intervals: 1. 3. 4, 5. 6. ve 8. 9. 10. 2 Teeth: margin...... 0 0 il 14 32 42 50 53 59 65 nerve dou or 0 0 0 0:2 d 3 5 7 7 10 » BRITISH SPECIES OF THE GENUS MNIUM. T Tyre V.: Sometimes a character reaches its maximum in the leaves which are above the longest leaf.— Example : In Mn. punctatum the nerve is short and never reaches the summit of the leaf in the three first intervals ; in intervals 4—10 it grows longer, more and more approaching the summit and often (not always) reaching this. In the second leaf above the longest leaf, and further upwards, the nerve reaches the summit in all the specimens I have observed. Remark : The classification of the characters in five groups with reference to their gradation is an artificial one, as all possible transitions between the types are found to exist. A common feature of all the gradation curves is their ascending-descending form. All the observed differences may be brought under two principles :—1°, the variation of the position of the summit ; 2°, the variation of the rapidity of increase (or decrease) along the axis. In other words, the differences between the curves are quantitative *. § 12.—From the examples given in the preceding paragraphs, we may conclude that the gradation of a particular series of measurements is a character in itself, independent of the absolute value of the feature under consideration. The gradation of a given character is more or less variable among the specimens of the same species (see § 8) and often very different from one species to another (see Table IX.). The gradation curve may be modified by the conditions of existence—we know at least one good example of such a modification T. The principle of gradation rules greatly in the morphology of plants, and also in a certain degree the structure of animals (see $ 23, note *, p. 33). A full discussion of this subject would go far beyond the limits of this paper. Therefore 1 want to close this part with five final remarks. Remark I.: The knowledge of the gradation of a certain number of characters enables us to make clear the variation of the leaves of the same stem.— Examples : In fig. 2 a given leaf (for instance, 25) is quite different from another given leaf (for instance, 17) with reference to the measurements of length and breadth. As each character has its own independent curve (see figs. 2 & 3), and as much diversity exists among the curves, the result is a practically unlimited number of combinations 1, the characters of each leaf depending on its position $. This form of variation seems to be of the same kind as the individual variation, but it is something quite different (see 82). * We may suppose that in the relation j—f (see note, $ 2, p. 8) certain variable factors intervene, the knowledge of which would perhaps enable us to explain the observed differences. + The gradation of the number of marginal florets in Chrysanthemum carinatum, Schousb. i In Table IL the figures in each vertical column give the combination of the cor- responding leat. $ This is also the case with branches, buds, flowers, fruits, trichomes, hydathodes, etc. LINN. JOURN.-—BOTANY, VOL. XLIV. C 1 f UN ie darm E ing es oy ag 18 PROF. JULIUS MACLEOD ON TEN Individual variation depends on chance, but the variation of the twenty-seven leaves in fig. 2 is independent of it *. The two forms of variability are often confounded, and the fruit of many laborious researches has been spoiled by that confusion. Let us suppose that we want to study the variation of the length of the leaves of the fertile stem of M. hornum (or any other species). If we collect and measure a certain number of leaves taken at random from a certain number of stems, and if we construct a variation curve by means of the collected figures, we bring together material which is not homogeneous. The variation depends here on two causes—chance and gradation,—the effects of which are not governed by the same laws. If we start from the idea that variation, in this and all similar cases, is merely governed by the law of chance, the calculation of a mean value and other more complicated computations will give artificial results. As this question is important, I may be allowed to give one more example. In Table XII. one finds the mean interval curves of four characters of the leaves of Mnium orthorrhynehum. ‘TABLE X Ll Mnium orthorrhynchum, Bruch, Schimp. & Guemb.— Mean interval curves of four characters of the leaves of the fertile stem. Intervals: 1. 2. 3. 4, 5. 6. i 8. 9. 10, Length (mm............. 076 0:99 193 141 159 2:06 234 291 319 396 Breadth (mm) .......... 0:36 042 050 056 0-65 084 093 0:93 083 061 PON. B et i3 jj HH 14 M 14 1 Marginal teeth (number) .. 1 2 5 7 dO 145 . 21:960 96 920 Although only four characters are taken into account in this example, the enormous differences existing between the leaves of the ten intervals are obvious (compare, for instance, the combination of figures in the vertical column 3 with the combination in column 8, etc.). The knowledge of the gradation curves is our leading clue among this disconcerting variation, which does not depend on chance. Remark II.: The principle of gradation enables us to answer the question, which leaf of a given specimen is comparable with a given leaf of another specimen : the leaves which are situafed at the same degree of the gradation axist are comparable.— Ezample: In a stem with 15 leaves (from the lowest leaf to the longest one) the 8th leaf (numbered from below) is at 500° (see the method of computation in $6) ; in a stem with 33 leaves the 17th leaf is also at 500°, and is thus comparable with the 8th leaf in the first stem. In the same way, the longest leaves of all the stems are comparable, as they are all at 1000? (see $ 3). * We leave out of account the fortuitous irregularities of the curves (see $ 21. T In other words, the leaves which have the same value « (see § 2). BRITISH SPECIES OF THE GENUS MNIUM. "m Unfortunately, apart from the longest leaf, this exact method is only applicable in exceptional cases, In a stem with 38 leaves, for instance, we do not find any leaf at 500°, and there is thus no leaf strictly comparable with the two former ones. A similar difficulty very often occurs. Therefore, if we want to study the individual variation and to construct a variation curve properly so-called, we must content ourselves with collecting and measuring the leaves which belong to the same interval in each stem. In this way the variation produced by gradation is eliminated, and we may consider the obtained variation curve as being governed only by the laws of chance. The easiest method is to limit ourselves to the 10th interval. This method is not quite accurate, because the leaves which belong to the same interval are not exactly at the same degree ; but it is approximate enough for our purpose, which is the description of species and the identification of specimens, If we wanted to be more accurate for any other purpose, we should take in each stem the longest leaf only. Remark III.: It is obvious that certain relations exist between the Characters of a given leaf and those of all other leaves.— Ewample : In Table XII. the leaves in intervals 5 and 10 have almost exactly the same breadth. Starting from this equality, we find that In interval 10: Length —3:96 mm.; breadth of the cells 213 u; number of marginal teeth =29. In interval 5: Length —1:59 mm.; breadth of the cells —13 u; number of marginal teeth —10. The proportions interval 5: interval 10 are Breadth: a o tal 0°65:0°64 = 102:100 Length <.. ieee. ee 1:59:3°96 = 40:100 Breadth of cells; js 18:18 = 100:100 Number of teeth 25 1020; = MF T00 Numerous comparisons of the same kind are possible : the results seem to be capricious to such a degree that one would think that any attempt to dis- cover any rule whatever would be vain. All similar relations between the leaves of a given stem depend, however, on the gradation of the characters under consideration. If we want to study the correlation between the leaves of several stems of the same species, we must compare leaves which belong to the same interval (in preference the 10th) or the longest leaf in all the stems, in order to eliminate the influence of gradation. The same method enables us to study the correlation between the characters of distinct species. Tables such as Tables IL., VIIL, XI., XII., and the specific tables B which are given in Part IV., are, in a certain sense, correlation tables. Without taking into account the principle of gradation, it is hardly possible to make o2 20 PROF. JULIUS MACLEOD ON TEN clear correlation in the Vegetable Kingdom. The limits of this paper do not allow a further discussion of this subject. Remark IV.: Gradation may produce, within the limits of one specimen, a segregation (disjunetion) of characters, which recalls the segregation occurring in Mendelian hybrids.— Example: Adopting the well-known presence-absence principle with reference to the visible properties, we consider three properties of the leaves of Mnium hornum :— (a) In some leaves the vein reaches the summit of the leaf: we call this character S. In other leaves of the same stem it does not reach the summit (absence of the charaeter) : we call this s. (b) Some leaves have marginal teeth: we call this character B. The absence of teeth, which is observed in other leaves of the same stem, is called b. (c) The presence of teeth on the nerve at the back of the leaf is called N. In some leaves of the same stem those teeth are absent; their absence is called n. Table XIII. gives the gradation of the three properties in one stem taken at random, only with reference to presence-absence. (See also Table XT.) TABLE XIII. Mnium hornum, Linn.—Presence-absence table of three characters in the leaves of one stem. Interval table. Intervals: 1. 2. 3. - B. 5. 6. T: 8. 9. 10. BERI ILU E 8 8 8 E E E 8 S S 1072 PENES "m b b B B B B B B B B NEM S A n n N N N N N N N Let us suppose now that a species SBN (in all the leaves: midrib reaching the summit, teeth at the margin, teeth on the nerve) is crossed with a species sbn (nerve ceasing below the summit ; margin and nerve without teeth). If the transmission of the properties happens regularly according to the Law of Mendel, the plants F, (second hybrid generation) are of eight different kinds, viz. : SBN, SBn, Sdn, sbn. SON, sBn, sBN, sbN, Of those eight combinations, four occur among the leaves of the stem in Table XIII., viz. : InantényalB:s1-2 [15 opo hors son. PE aAa OA ig. Pesala fs sDn. : e N C ad e COUP DAS sBN a A 135995... UMS d SBN, BRITISH SPECIES OF THE GENUS MNIUM. 21 It may be pointed out that it is impossible to find the explanation of those facts in the ordinary hypotheses of the Mendelian theory—such as the prineiple of segregation of the zygotes, the dominant-regressive principle, the hypothesis of the hereditary factors, and the presence-absence theory with reference to these factors,—because all the leaves in Table XIII. belong to one stem. Here the base of the explanation is found in a quite different principle—gradation. This calls for reflection *. Kemark V.: We have seen (§ 12) that the gradation of a given character is a character in itself. It would be interesting to observe the transmission of a gradation curve in hybridization. We see, for instance, in Table IX. (p. 16) that in JM. rostratum the gradation curve of the breadth of the leaves at their base has its summit in interval 10 ; the summit of the corresponding curve of M. spinosum is found in interval 3. What would be the form of the curve in the hybrids between the two species? Hybridization of Mosses seems to be impracticable ; but it is certainly possible to find among the Phanerogams two species different from each other by the form of the gradation eurve of a given character and suitable for experiment. PARE- LE INDIVIDUAL VARIATION. The Use of Figures for the Description of Species and the Identification of Specimens. $13. INDIVIDUAL VARIATION.—Let us suppose that we have measured a given character of the longest leaf of a certain number of specimens (fertile stems) belonging to a given species of Mniwn—for instance, the length of the longest leaf of eight stems of M. spinosum. In this example the figures were (in mm.) : 655 668 798 758 764 764 748 — 810 As all the measured leaves are exactly comparable (see § 12, Remark II.), the influence of gradation is eliminated. Here the variation is governed by the laws of chance ; it is individual variation. What answer will be given when we are asked which value is characteristic for the species ? Here we meet the first difficulty mentioned in $ 1. $14. MEAN VALUE.—According to the classic method, we might calculate the mean value (in the above example, 7°38 mm.) and take this as a cha- racteristic one. But the significance of a mean value, and especially its * See on the general importance of gradation, $ 12, p. 17, second paragraph. 22 PROF. JULIUS MACLEOD ON TEN biological significance, is something VERY VARIABLE and therefore rather vague. When we want to construct, for instance, a mean interval curve of a certain character of a group of specimens (see § 7), we need to find the relative value of the ten ordinates. As the variation of the character within the limits of a given interval depends (almost entirely) on chance, the mean value in an interval a is approximately the most probable (the most frequent) value of the character in the middle of a. Taking the mean values of the ten intervals, we eliminate as much as possible the influence of chance; the differences between the ten figures depend (almost entirely) on the influence of gradation. The construction of a mean interval curve (in which the ten mean values are used) is thus justified ; such a curve gives us, by the relative values of its ordinates, an adequate representation of the variation of the character along the axis. But the absolute values of the mean values (ordinates) are of secondary importance, because those absolute values vary from one patch to another. (See the construction of percentage interval curves, in which the absolute values are eliminated, § 8.) We see from this example that, when we want to use mean values, their real significance ought to be exactly defined in each peculiar case. What is, for instance, the significance of the mean length 7:38 mm., calculated by means of the eight figures in §13? As the variation of these eight figures depends merely on chance, the calculation of a mean value is justified from a mathematical standpoint. But its significance is limited to the patch to which the eight stems belong; similar values may be useful ‘for the comparison of several patches with each other *, but they are not characteristic for the species. It is, in fact, IMPOSSIBLE to establish the mean value of a variable character of a species. In the above example (M. spinosum, § 13) the mean length of the leaves was calculated by means of specimens belonging to one patch. But when we collect several patches of the same species in different localities, each patch gives a different mean value ; when we bring these values together and try to calculate in that way the mean value of the species, the result is, of course, an ARTIFICIAL one and will be modified again and again by each new observation. Moreover, even if we succeeded in establishing a specific mean value f, this would give us little help, if any help at all, for the identification of a given specimen.— Example : We have found for the length of the longest leaf of Mnium spinosum the value 7°38; in a similar way we found for Maium cuspidatum the value 4:63 mm. Let us suppose that those values were really * For instance, with reference to the influence of different conditions of existence. + For instance, by cultivating the species under certain conditions of existence defined by convention—a method which is practicable only in exceptional cases. BRITISH SPECIES OF THE GENUS MNIUM. 23 specific. As the character under consideration is variable in each species, it is impossible to say to which of the two belongs a stem, the longest leaf of which has a length intermediate between the two figures ; and if the individual figure of the unknown stem is, for instance, 4 mm. or 8 mm., we can give no more than a guess at its identity. $15. MaxiMAL AND MINIMAL VALUES.—In a series of measurements representing the individual variation of a given property of a species, the MAXIMAL value and (with certain restrictions) the minimal value have a more definite BIOLOGICAL significance than the mean value. Moreover, it is possible to find these two limits, once for all, by a sufficient number of Observations. "THEREFORE THE MAXIMAL AND MINIMAL VALUES ARE THE CHARACTERISTIO CONSTANTS OF THE SPECIES. Adopting this view, I deviate from the classical principle of the mean value introduced in biometrical science by Quetelet (1846). This principle, borrowed from the theory of errors, is quite justified from a mathematical standpoint, and may do good service in certain cases (see $14); but from a biological point of view a mean value is very often deceptive. There are serious physiological reasons for considering the minimal and maximal limits as constants and for ascribing to them a definite biological and mathematical significance. I hope to expound this subject more completely in another publication *. $16. PRACTICAL USE OF THE LIMIT VALUES.—In the present paper I want to content myself by pointing out the practical importance of the minimal and maximal values for the description of species and the identification of specimens. We take, as first example, the length of the leaves of the tenth interval of the fertile stem of two species, Mnium cuspidatum and M. hornum. The measurement of a certain number of specimens, belonging to patches collected in several localities, has given the following limits :— cuspidatum. iTia MESE 373 mm. 7'44 mm. hornum :.......- ERU. DIT 80817 In this case the variation is transgressive ; in other words, the two series (variation eurves) overlap each other. Let us suppose that we want to identify a specimen a, and that we are hesitating between the two mentioned species. The length of the longest leaf of w is 5:33 mm. ; this figure being between the minimum of hornum and the maximum of cuspidatum, it is impossible to make a choice. As the examples of transgressive variability are very numerous, a similar difficulty * I beg the reader of the present paper to accept as a postulate that the maximal valne and (with certain restrictions) the minimal value are specific constants. Edu o See ee a i eee Oe 24 PROF, JULIUS MACLEOD ON TEN occurs again and again, and it seems to be insuperable. There are, however, three methods to surmount it. The FIRST METHOD consists in trying a second character of the same specimen æ : for instance, the breadth of the longest leaf, which is in our example 1:93 mm. On the other hand, the limits of the breadth are in the two species : E Oumtdatum- Dove ee 1:29 mm. 3:26 mm. LIST s 1127 ABUSE IRSE E ets 004 5.35 1:446. Although the two specific curves are again overlapping each other, hesitation is no longer possible : æ is cuspidatum. In the above example, two measurements allow us to identify 2 *. According to my experience with Carabus and Mnium, this happens more frequently than one would think. It is, of course, often necessary to measure more than two characters of a specimen, especially when we want to make a choice between numerous species. The SECOND METHOD is applicable when we have at our disposal two or more specimens which belong with certainty to the same species, This is the ordinary case with Mosses, as several stems of the same species are to be found in one patch f.— Example: In a patch w, the breadth of the longest leaf of four fertile stems was measured ; the figures were 1:19 mm. 1:15 mm, 0:05 mm. 0:67 mm. Comparing the specific limits given above, we see that the first figure (1:19) is doubtful ; it points to hornum, but the difference between 1:19 and the minimum of cuspidatum is too small to give a decision. The second figure (1:15) is also doubtful, but the two figures together make it probable that the patch æ is hornum. The third and the fourth figures (0°95 and 0°67) are decisive in favour of hornum ; the four stems (and the whole patch 4) belong thus to this species. The principle of the second method is that, in a series of specimens which certainly belong to the same species, the identification of al/ the specimens is possible if one of them gives a decisive figure—even if the figures of all others are not decisive. The THIRD METHOD is applicable if one specimen affords opportunity of measuring a given character two or several times. In the above example of application of the first method we limited ourselves to the measurements of the longest leaf of the fertile stem æ which we wanted to identify ; but we may go further, and take all the leaves of the tenth interval of z ($ 12, Remark II.). The number of those leaves depends, of course, on the total number of leaves of the stem, as one sees in Table XIII. A. * If we had taken at first the breadth, one figure would have been sufficient. T It is, of course, necessary to ascertain that the patch is homogeneous. wee Ree ee ea. m s —""- * TA eu urit s BRITISH SPECIES OF THE GENUS MNIUM. 25 Application of the third method is possible when the total number of leaves is greater than ten. TanLE XIII. A. Number of leaves in the tenth interval. (See the method of calculation in § 6.) Total number of leaves (from the lowest leaf Number of leaves to the longest one). in the tenth interval. Eoss than Ll... -. T IU MEER HK 1 NA Ce. Bar oi UNT. 2 eee DEUS n MUN eet ar 3 PI MGS aE ge 4 41-50 .....57.. ee ; 5 etc etc. Example : The breadth of the leaves in the tenth interval is variable— In Mnium orthorrhynchum between 0:30 and 1:03 mm. In Mnium serratum » Doon 10r ays We want to discover to which of the two species belongs a stem s. The number of leaves is 19; in the tenth interval we find thus two leaves. The figures of those leaves are : Breadth : longest. leai m Oy urs 1:02 mm. B leaf below the longest one........ 120. The figure of the longest leaf is dubious, but the second figure (1:26) is decisive : « belongs to the species serratum*. $17. PRACTICAL USE OF THE LIMIT VALUES (continued).—I give in Part V. a certain number of tables of identification, in which are found the lowest and the highest value of the fourteen measured characters for the ten species of Mnium I have studied. The figures were obtained for each species by the measurement of specimens from as many localities as possible. Let us try to identify a specimen « belonging to one of the ten mentioned species. * It is possible to apply the third method to the identification of animals—for instance, of Beetles. A considerable difference may exist between the right and the left side ot one specimen with reference to certain characters; it may happen that a certain ligure is dubious at one side, the corresponding figure at the other side of the same specimen being decisive. 26 PROF. JULIUS MACLEOD ON TEN First mernop : Identification of one fertile stem z. The figures are : gimpa i eei (0) length ches. ees eere 4:66 mm. 5 ncc doctob cor TE TER ER 17855 5 nuo DORAIN at ble DARE . ii eee hn nnn v82: a » (d) breadth of the cells at the em of the ErentesbDISAQth. 63. 0s cae c's red nn 19 y. 5 » (e) number of cells at the place of the giviitat IGA Ue ARO a EO ag oret 88 p » (f) breadth of the border at the place of the mprearestipneaath .. 0 5 0. ic dete 4 3 37 p. $5 » (g) number of cells of the border at the place of the greatest breadth ............... 6 ^ » (A) number of marginal teeth .............. 4l 5 » (č) number of double teeth at the margin .... 0 5 » (J) teeth on the nerve at the back of the Hos WEE c EN CO CI A v CES DES 0 the nerve reaches 5 ni Ce) leneih/of themerve *- 5... srl the summit of the leaf. 5; » (4) tooth at the summit of the leaf.......... exists. Eum On A A Und ceo e rex rere 24 KE ogean sd p OMNE Co RR QOEM 1 The order in which the tables may be used is arbitrary. We begin with Table XXV. (length) : the minimum table excludes undulatum and probably affine and hornum ; the maximum table does not exclude any species. Table XXVI. (breadth): the minimum table excludes subglobosum, punc- tatum, and probably affine; the maximum table excludes orthorrhynchum, probably serratum, and perhaps hornum. Table XXVII. (breadth of the leaves at their base) : no more species are excluded. Table XXVIII. (breadth of the cells): the minimum table excludes sub- globosum and punctatum (both already excluded), and brings confirmation of the exclusion of afine; no more species are excluded by the maximum table. Table XXIX. (number of cells) : no species are excluded by the minimum table; the maximum table brings confirmation of the exclusion of serratum, excludes orthorrhynchum (already excluded), and perhaps hornum. Table XXX. (breadth of the border): the minimum table excludes rostratum, spinosum, and also punctatum and subglobosum (both already excluded) ; no species are excluded by the maximum table. Hitherto eight species have been excluded and a ninth species (hornum) has been excluded three times, although not with certainty. It may be remarked that we have been prudent in excluding species, allowing a wide * On the measurement of the character k, see $ 35. BRITISH SPECIES OF THE GENUS MNIUM. 21 margin of error, because the REAL specific limits are probably not yet reached in this FIRST ATTEMPT of construction of minimum and maximum tables *. We may conclude that the stem æ is probably cuspidatum, perhaps hornum. We take now Tabie XXXI. (number of cells of the border) : no more species excluded. Table XXXII. (number of marginal teeth) : no more species excluded. Table XXXIII. (number of double teeth at the margin) : the exclusion of hornum is confirmed by the minimum table. The stem æ belongs thus to the species Mnium cuspidatum. One may perhaps think that we aseribe too high a significance to the figures when we conclude, from rather small differences between the figures of specimen w and the figures in the tables, that æ does not belong to such or such a species. One must realize, once for all, that each figure in the tables is a Limir deduced from a series of measurements, and has thus a significance quite different from that of a figure given by one specimen taken at random. We may, moreover, identify specimen æ a second time, taking the tables in the reverse order and beginning with Table XXXVIII. (number of fruits): the minimum table excludes perhaps spinosum and undulatum ; no species are excluded by the maximum table. Table XX XVII. (number of leaves): the minimum table excludes hornum ; no species are excluded with certainty by the maximum table. Table XXXVI. (tooth at the summit of the longest leaf) exeludes sub- globosum. Table XXXV. (length of the nerve) excludes no more species. Table XXXIV. (teeth on the nerve at the back of the leaf): the minimum table excludes hornum (already excluded) and probably spinosum. No species are excluded by the maximum table. Table XXXIII. (number of double teeth at the leaf-margin) excludes serratum and orthorrhynchum, brings confirmation of the exclusion of spinosum and excludes kornum (already excluded). No species are excluded by the maximum table. Table XX XII. (number of marginal teeth) : the minimum table confirms ° the exclusion of undulatum. The maximum table excludes punctatum, and also subglobosum and serratum (both previously excluded). Table XX XI. (number of cells of the border) does not exclude more species. * The figures in the tables are approximate enough to demonstrate the use of the method, but they want to be verified and corrected by more measurements. It may be remembered that many constants, used by physicists and chemists, were at the beginning only approximate. They have been corrected later on—often at the cost of long labour. eee we ee TRES T 28 PKOF. JULIUS MACLEOD ON TEN Table XXX. (breadth of the border): the minimum table excludes rostratum, perhaps affine (and also subglobosum, spinosum, and punctatum previously excluded), Eight species are already excluded. The stem æ is thus probaby cuspi- datum, perhaps affine. Table XXIX. (number of cells) does not give an answer. Table XXVIII. (breadth of the cells): here the minimum table is decisive, affine is excluded. The stem æ belongs thus to the species Mn. cuspidatum : the first identi- fication is confirmed (continued $ 19). $18. THE QUANTITATIVE DESCRIPTION OF ONE INDIVIDUAL (BERTIL- LONAGE): its value for descriptive botany and zoology.—The description of one individual of Mnium cuspidatum given in § 17* may be compared with the well-known bertillonage, which is the quantitative description of a person. A bertillonage is a combination of figures obtained by the measurement of a certain number of characters of a person n. As a given combination practically never occurs a second time, it is characteristic of the person n ; it enables us to differentiate n from any other human individual whatever, and thus to identify it. In a similar way the quantitative description of an animal or a vegetable specimen enables us:—1°, to discover the species to which it belongs (by means of maximum and minimum tables) ; 2°, to differentiate it from any other living individual of the same species or of any species whatever. Ithinkit would be very useful to add to the description of a new species the quantitative description of a few or at least of one specimen T. By means of the exact information given in such a description we would be enabled to identify later on the described species, even if the type is lost. In innumerable cases it is possible to make a sufficient number of measure- ments without disturbing the specimens. I succeeded in measuring 38 characters of thousands of specimens of Carabus mounted on needles (more than 250,000 measurements) practically without spoiling them, except in the early days when I had not yet had enough practice in the work i. In the case of the Mosses and other minute objects (small insects, Crustacea, flowers, etc.) one is compelled to dissect the specimens, but in case of need * To this description the gradation curves of the characters a-/ might have been added, but this was useless for our purpose. T Measuring as many characters as possible. f If one wants to measure precious specimens (specimens of certain species of Carabus have a value of 10 shillings each and even more) it is advisable to practise at first by measuring similar, but valueless objects. "BWEARWISST- . BRITISH SPECIES OF THE GENUS MNIUM. 29 it is possible to keep the measured parts in permanent microscopical preparations, glass tubes, etc. $19. (Continued from § 17.) PRACTICAL USE OF THE LIMIT VALUES.— Now we want to make a trial of the SECOND METHOD of identification by means of limit tables (see $ 16). We take a fertile stem a belonging toa patch æ. We measure merely three characters. The figures are :— Length of the longest leaf ................% . 615mm Breadth ,, i A aae A CIG es; E i m p Sat its basó, "os e 075 We begin with Table XXV. (length): the minimum table excludes no species with certainty, the maximum table excludes serratum and perhaps orthorrhynchum. Table XXVI. (breadth): the minimum table excludes affine, punctatum, subglobosum, and perhaps rostratum and undulatum ; the maximum table does not exclude any species. Table XXVII. (breadth at the base): the minimum table excludes perhaps undulatum ; the maximum table excludes perhaps orthorrhynchum. By means of three measurements of stem @ we were enabled to exclude four species :—serratum, affine, punctatum, subglobosum, and perhaps three more species: rostratum, orthorrhynchum, and undulatum. We now take a second stem b of the same patch æ ; the figures are :— ength ofthe longest louie, M ese 6:92 mm. Breadth ., N Re yee S esr QT T. b "t Py een see Or; Table XXV. (length): no species are excluded by the minimum table, the maximum table excludes serratum (previously excluded) and confirms the exclusion of orthorrhynchum. Table XXVI. (breadth): the minimum table excludes cuspidatum, spinosum, afine (already excluded), punctatum (id. id.), subglobosum (id. id.), and confirms the exclusion of rostratum and undulatum. Nine species being excluded, it becomes useless to go further: the patch æ belongs to the species Mnium hornum *. Ewample of the use of limit tables according to the THIRD METHOD of identification: we take one stem æ with 43 leaves and measure four properties of the 5 leaves which belong to the 10th interval (see Table XIII. A). The figures are given in Table XIV. * It may be remarked that in this example it was possible to identify the patch v by measuring merely three properties of two leaves belonging to two stems, without taking into account any of the characters used in the classical books for the identification of Mnium hornum. ERA IND T. 30 PROF. JULIUS MACLEOD ON TEN TABLE XIV. | Mnium hornum, Linn.—One stem with 43 leaves. | : Characters, Particulars of the leaves in the 10th interval. A. 3 E. Breadth, leaves .......... s... 181 124 188 087 115 - E # base leaves ,.:....... 071 0:75 076 0:66 0:75 | ‘ Marginal-teeth .......... 1:1... 58 52 58 46 60 [ AW (D9 4 SE TRECE RET ER Sess yd 9 9 6 T Table XXVI. (breadth), minimum : by means of the lowest figure of æ in Table XIV. (0°87) we exclude spinosum, undulatum, rostratum, affine, punc- tatum, and subglobosum. Using the maximum table, we exclude orthorrhyn- chum by means of the highest figure of æ (1:38). Table XX XII. (marginal teeth), minimum: by means of the lowest value of æ (46) we exclude undulatum, previously excluded. The maximum table enables us to exclude by means of the highest value of æ (60) : punctatum, subglobosum, and orthorrhynchum (previously excluded), and also serratum. The stem æ thus belongs to hornum or cuspidatum. Table XXXIV. (teeth on the nerve): cuspidatum is excluded by the maximum table. The stem w thus belongs to the species Mnium hornum. $20. REMARKS ON THE USE OF NUMBERS FOR THE IDENTIFICATION OF SPECIMENS.—For the identification of specimens we have three methods at our disposal, each of them allowing verification of the result given by the others. Each of the three methods affords a rather wide range of possibilities. Applying the first method (with one leaf of one stem, see $ 17) we may very often use the limit-tables in two ways, starting from the first or from the last table: this allows verification (see the example in $ 17). Moreover, it is possible to measure a greater number of properties, for instance (taking the Acrocarpic Mosses as an example) : the length of the so-called fruit-stalk, some properties of the teeth of the peristome, the dimensions of the ripe spores. A section of a fertile stem (exactly in the transverse direction) at the insertion of the longest leaf would furnish some more measurable properties. Of course, the practical value of the method will increase in proportion as the number of the measured properties becomes greater *, Using the SECOND METHOD (see $ 19) we may take more than two speci- mens and, in case of need, verify the result as many times as the material at our disposal. is afforded by * I have measured 38 characters of Carabus. Tn this paper I limit myself to 14 characters of Mnium, because I want to avoid too long a delay in the publication of a first paper on the results of the work with which I have been occupied for many years. CARNE E SERT e Wet rw qp TA METER E TS MS EINE LE ETER BRITISH SPECIES OF THE GENUS MNIUM. 2L The second and the third method will be useful, especially in those cases in which it is impossible to measure a great number of characters (for instance, in many Fungi, Algæ, and lower animals). Finally, the three methods may be used at the same time. For more than a century innumerable species have been described in several instances under different names, because the available descriptions did not allow an exact identification. The result has been an enormous waste of time and a prejudicial disorder in the synonymy. We are still going on in the same way, and it is to be feared that in the future more labour will be wasted and more disorder will arise. ‘Would it be too much to hope that the state of things will be improved by the exact measurement of characters and the construction of limit-tables ? PART: III. THE MEASUREMENT OF THE CHARACTERS OF THE LEAVES IN THE GENUS Mxriua. $21. EXACT MEASUREMENTS. POSSIBLE ERROR.—The exact expression by measurement of the characters of a living being is only possible if each character is strictly determined. When we want to measure, for instance, the length of the leaves of a certain plant, we must know exactly which leaves are to be measured, and we must determine the significance which we ascribe to the word length and even to the word leaf. There is something elusive in the variation of animals and plants. This variation is so great that one might be tempted to content himself with more or less approximate measurements, This would be a grievous mistake, by which the use of the quantitative method might be seriously endangered. As soon as we use the quantitative method we discover how many of our notions are rather superficial; and, as we proceed, we become more and more aware of the necessity of reaching the highest possible degree of exactitude. Errors are unavoidable, but even the limits of error ought to be as much as possible determined. In order to obtain an estimate of the possible errors, I measure a given character p of an object several times ; after a few days I repeat the same measurement with the same object several times. The difference between the highest and the lowest figure divided by 2 is the possible (positive or negative) error with reference to the character p. All the figures given iu this paper were verified at least by a second measurement. 32 PROF. JULIUS MACLEOD ON TEN It may be recommended to the biologist who wants to apply the quanti- tative method to exercise himself, by measuring some exactly known properties of inorganic objects *. In the present Part I give the definition of all the characters I have measured, the methods of measurement, and the possible error of the figures obtained. N.B.—In all the calculations the decimal 0°5 has been taken as 1. (Example : 31:5 is brought into account as 38.) 22, FERTILE stem.—I call fertile stem a stem which bears a ripe or almost ripe fruit, or at least a fruit sufficiently advanced in its development to make it certain that the leaves, even those which are iuserted near the summit, are adult. Here there is no difficulty. $33. DEFINITION OF THE WORD “ LEAF." NUMBER OF LEAVES.— The lowest phyllomes of a fertile stem often differ in their facies from the true leaves which are inserted higher up: therefore they are sometimes called scales (for instance, in M. hornum). Asa gradual transition exists between scales and leaves, it is impossible to find a strict limit between both: therefore I call them all leaves T. On the other hand, near the summit of the stem, we find almost always some phyllomes which constitute a sort of perianth. Sometimes there is a distinct breach of continuity in the gradation between the upper leaf and the first phyllome of the perianth, the latter being, for instance, much smaller than the former : in such cases, no doubt exists about the limit between the true leaves and the perianth. But ordinarily the gradation between both is continuous, and we must come to an agreement as to where the limit is to be made. Proceeding towards the summit (and neglecting the lowest leaves) we find: 1°, a certain number of leaves which belong (according to the species) to the forms represented in fig. 4, 7, 2, 3 (in 7 the breadth increases from the base b to D, and decreases further to the summit of the leaf; in 2 the breadth increases from b to B'and is constant between D' and D : in 3 the breadth is constant between 6 and B); 2°, approaching the summit we very often find the form 4 (narrowed between b and B); 5°, near the summit we find ordinarily phyllomes, the breadth of which is decreasing continuously from b to the summit without constriction. I give the name of leaves to the * For instance: the angles of a crystal, the density of a substance, the boiling-point of a liquid, ete. + The differences and also the transition between scales and leaves find their accurate expression in the successive figures of the gradation curves (for instance, with reference to length, breadth, number of teeth, dimensions of the cells, etc.). BRITISH SPECIES OF THE GENUS MNIUM. oo forms 7, 2, 3, 4, excluding form 5, which I consider as belonging to the perianth *. B B B B A eli JL shl] T 2 3 4 5 Fia. 4.—1-4, leaves; 5, perianth ; b, base. See text. In a fertile stem several leaves near the base are usually destroyed. There- fore the number of leaves given in Table XXXVII. (page 56) includes merely the leaves which are still in existence when the fruit is ripe or half ripe. This error is unavoidable ; it does not prevent the specific figures from being comparable with each other. The lowest leaves being often concealed among the hairs with which the lowest part of the stem is clothed, they are to be searched for under the simple microscope, the hairs being carefully removed by means of needles. As this is a rather delicate task, owing to the small dimensions of those leaves, it may happen that some of them are overlooked. The figures given in Table XXXVII. are therefore a little below the reality. I estimate the possible error at 3 for the species with smaller figures (punctatum, ete.) and at 6 for the species with very numerous leaves (undulatum, ete.). Some fertile stems show distinctly two successive periods of gradation f : * It may be allowed to make a momentary excursion into a by-path. The breadth of a moss-phyllome varies along an axis of gradation, the direction of which is given by the nerve. The margin of the phyllome is, on each side of it, a curve of the gradation of the breadth drawn by nature itself. Comparing the phyllomes in fig. 4, we see that in leaf Z the summit B of the curve is situated between the base and the summit of the leaf; in the leaves 2-3 the breadth is invariable from 4 to B (or B' to B), and decreases further towards the summit; in phyllome 5 the summit of the curve corresponds to the base ; in leaf 4 we find two summits b and D. As the development of a phyllome is basipetal, its successive parts from the summit to the base have been developed successively, in a similar way as the leaves along the stem (although in reverse order). We may thus say that in the phyllomes 1, 2,3, and 5 one period of gradation exists, and in leaf 4 two periods are observed. The variation of the gradation curves in fig. 4 recalls the variation of the curves in Table IX. (page 16). See $ 12, second paragraph. T This occurs rather frequently in certain species: for instance, Mnium subglobosum and punctutum. It is the ordinary rule in Cinclidium stygium, Sw. LINN. JOURN.—BUTANY, VOL. XLIV. D jl 34 PROF. JULIUS MACLEOD ON TEN the first period, including the leaves of the lower part of the stem, is suddenly interrupted at a certain level, and followed by a second period which begins with very small leaves and goes on regularly to the summit. The limit between both periods is distinctly indicated by a breach of continuity in the gradation of the length of the successive leaves. When two periods exist, only the ssconp PERIOD (which ends at the summit of the stem) is taken into account. $24. PREPARATION OF THE LEAVES.—Herbarium specimens and alcohol material, having been placed in hot water for a few minutes, may be used as well as fresh objects. All the leaves are separated from the stem, proceeding from the base towards the summit, under the simple microscope, by means of a scalpel. The leaves are placed in their order of succession (their back upward) on slides, in a very small quantity of water (just enough to prevent desiccation). They are then covered, the underside of each cover-glass being moistened with diluted glycerine. If a leaf is brought at once in (even diluted) glycerine, it may curl up before it is possible to place the cover-glass over it. $25. LENGTH OF THE LEAF.—At the base of a leaf separated from the stem one finds easily a more or less irregular curved line e, corresponding to the upper limit of the surface of insertion of the leaf. I take as inferior limit of the leaf the point 7, which is in the centre of the surface limited by the curve ¢ and the transversal straight line bb, the latter being the span of the arch c. Following the nerve, I measure under the mieroscope the length of the leaf from the extreme summit to ¿ As it is impossible to use a high power according to the dimen- sions of the object (I used obj. 3 Leitz), and as the position of 7 is not strictly determin- able, a positive or negative error of 30 u (0:03 mm.) is possible. Fic. 5.— Lower part of a leaf separated from the stem. See text. n, nerve. $26. BREADTH OF THE LEAF. PLACE OF THE GREATEST BREADTH.—In almost all the leaves the place of the greatest breadth is easily found, the forms represented in fig. 4, 7 and 4 (page 33), being the commonest. In some leaves the greatest breadth extends over a certain length, the margins being parallel (fig. 4, 2, 3) : here we agree to take the upper limit of the parallel portion (indicated by B in fig. 4, 2, 3) as the place of the greatest breadth. Among the lowest leaves a form similar to fig. 4, 5, sometimes BRITISH SPECIES OF THE GENUS MNIUM. 35 occurs : here the base is the broadest part of the leaf. I have considered the breadth of such leaves (which are rare) as unmeasurable. Sometimes the differentiated border of the leaf is recurved (this happens often in M. hornum). Here I measure first the visible breadth b (with- out taking the recurved parts into account); then the recurved parts are measured separately and their breadth is added to b. Iestimate the possible + or — error in the measurement of the breadth at 20 u (0°02 mm.). § 27. BREADTH AT THE BASE.—This property is measured in the direction of a transverse line passing through the point / mentioned in $ 25 (see fig. 5, p.34). If the margin is recurved, I use the method described in § 26. As the margins at the base of the leaf are often oblique, a small error in the estimation of the position of i may bring about a sensible error in the measurement of the character under eonsideration. Therefore the possible positive or negative error is proportionally great: I estimate it at 30 u (0°03 mm.). $38. NUMBER OF CELLS AT THE PLACE OF THE GREATEST BREADTH.—The place of the greatest breadth being determined (see $ 26), I count the cells in the transversal direction, separately in the right and the left half of the leaf, I exclude:—1°, the nerve; 2°, the elongated cells of the differentiated border, except the interior one which is taken into account*. In each half of the leaf I count the cells four times (proceeding twice from the nerve to the margin and twice in the reverse sense) and take the mean value of the four figures. Ordinarily these figures differ from each other by one or two units. If the differences are greater f, I go on, counting two or four times more. The sum of the mean values of the two halves is the figure of the leaf. I estimate the possible positive or negative error at two cells. This estimation is rather too high :when the cells are not very numerous (see for instance, the minimal values of serratum and orthorrhynchum in Table XXIX.). $29. BREADTH OF THE CELLS AT THE PLACE OF THE GREATEST BREADTH.— As the form and the size of the cells are very variable, even in one leaf, it is practically impossible to obtain useful information about their dimensions by measuring them separately. I have tried to surmount this difficulty by determining the mean dimension of the cells in the transverse direction (in other words, their average breadth) at the place of the greatest breadth of the leaf. * If the differentiated border consists of one cell (a single row of cells) this is excluded. + If the figures are, for instance, 22, —23, —25, —26 (a rather unsatisfactory result). D2 36 PROF. JULIUS MACLEOD ON TEN Having subtracted the breadth of the nerve and the breadth of the two. differentiated borders (see § 31) from the total breadth of the leaf, I divide the rest by the number of cells (see § 28) : the quotient is the mean breadth of the cells. It is very difficult to measure exactly the breadth of the nerve, its lateral limits being rather indistinct. The error, being divided by the number of cells, can be neglected. I estimate the possible positive or negative error in the value of the mean breadth at 0*5 u. If the number of cells is small (for instance, 25) the error may reach 1 w (maximum) *. The method here described gives very regular results, and is applicable (with the necessary changes) to the measurement of cells, fibres, ete., in numerous objects. § 30. NUMBER OF CELLS OF THE BORDER AT THE PLACE OF THE GREATEST BREADTH.—In the 10 species of Mniwm which I have studied (and in many other Mosses) we find along the margin of the leaves one or several rows of cells which constitute the so-called border. The border-cells are differ- entiated from the neighbouring cells of the leaf by two properties :— 1°, they are distinctly elongated, often several times longer than broad, whilst the neighbouring cells are about isodiametrical ; 2°, their cell-walls are thickened. These two characters, however, seem to be to a certain degree independent of each other, the differentiation being sometimes limited to one of the two. We find, for instance (often in the lowest leaves), border-cells which are distinctly elongated, their walls being hardly or not thickened, and other border-cells which are hardly longer than broad, but the cell-walls of which are thickened. I consider as belonging to the border the cells which are differentiated at least by one of both properties. At the inner limit of the border we often find cells (fig. 6, e) which are elongated but not thickened. Such cells are taken into account in counting the cells of the border. As the number of border-cells is variable along the margin, I number the cells three times in each leaf at each side: 1°, at the place of the greatest breadth ; 2°, about 50 to 100 u (according to the dimension of the leaf) above this place ; 3°, the same below the mentioned place. I take the highest of the three figures. The two maxima (right and left border) are * I want to call attention once more to the usefulness of strict conventions. Instead of saying, for instance, “in Mnium sp. the diameter of the cells of the leaves is about 20 4,” I say, “in Mnium sp. at the place of the greatest breadth of the longest leaf of a fertile stem the mean breadth of the cells varies between 17 and 24 p,” and I add the possible error, One might say that the disconcerting variation of the character under consideration has been caught and confined between two constants. BRITISH SPECIES OF THE GENUS MNIUM. 37 added: their sum is the figure of the leaf. Places immediately below a tooth are excluded (fig. 6, n). Example: In fig. 6, I number 3 cells at a, 4 cells at b, 4 cells at c (in- cluding ¢), 2 cells at d. I take the maximum: 4 cells. There is no difficulty if the border contains one layer of cells (fig. 6). If the border-cells are placed in two (or more ? ) superposed layers, the cells are numbered as they are seen in the optical section: here an error of one cell is possible, and this may occur in the same sense at both sides of the leaf. I estimate the possible positive or negative error of the figure of the leaf at 2 cells (1 cell if the cells are not numerous) *. $31. BREADTH OF THE BORDER.— This character also is very variable along the margin. I measure the breadth at three places (same method as for the number of cells, § 30) and take the mean value of the three measurements. The sum of the two mean values (right and left border of the leaf) is the figure of the leaf. I always exclude the in- terior border-cell, but I include the exterior wall of this cell in the measure- ment.— Example (fig. 6): Measuring Fia. 6.—Border of a leaf—B, place of the breadth of the border at a, I take the greatest breadth of the leaf. the two exterior cells, including the t, tooth ; a, b, c, d, e, n, see § 30. wall between the 2nd and the 3rd cell ; measuring at B I take three cells, excluding e and including the exterior wallof e; measuring at d I take the exterior cell and the interior wall of this. I calculate the mean value of the measurements a, b, c, and add this to the mean value of the other side of the leaf. I have used obj. 6 Leitz. Iestimate the possible positive or negative error of the figure of a leaf at 5 p. * If the border is recurved, it is, of course, impossible to count the border-cells without making sections 38 PROF. JULIUS MACLEOD ON TEN § 32. NUMBER OF MARGINAL TEETH.—No explanation is needed if all the teeth are simple. If certain teeth are in groups of 2 (even of 3) each group is brought into account as 1. ‘Teeth which are situated below the base of the leaf (see 7 in fig. 5, page 34) are excluded *. It is sometimes difficult to say whether a given marginal cell ought to be considered as a tooth or not. I consider as a tooth each marginal cell, the distal extremity of which projects beyond the margin by at least one-half of its breadth (see fig. 7). Fre. 7.—Rudimentary teeth. m, margin; a and o are considered as teeth ; c is not taken into account as a tooth. The figure of a leaf is the total number of teeth at the right and the left margin. I estimate the possible positive or negative error at 1 tooth, if all the teeth are simple. If double or triple teeth exist, the error may reach 3. § 33. NUMBER OF DOUBLE (OR TRIPLE) MARGINAL TEETH.—In several species double or even triple teeth exist. These are counted separately. Each group (of 2 or 3 teeth) is taken into account as 1 unit. Figure of the leaf noted as in $32. Possible positive or negative error, 2. § 34. TEETH ON THE NERVE AT THE BACK OF THE L&AF.—(Careful obser- vation is needed for the counting of these teeth. A group of 2 teeth (situated at the same level—this rarely occurs) js brought into account as l unit, Error: the observed number may be too small by 1 unit; a positive error is almost impossible. $35. LENGTH OF THE MIDRIB (reaching the summit or not).—The length of the nerve is very variable. A measurement of this length is difficult, because * Such teeth occur in certain decurrent leaves: for instance, in some leaves of Mnium undulatum. BRITISH SPECIES OF THE GENUS MNIUM. 39 it is often practically impossible to determine the distal limit of the nerve. Therefore I admit three degrees: 1*, the midrib is short and does not reach the summit (fig. 8, 7) ; 2°, there is a trace of relation between the extremity of the nerve and the summit (fig. 8, 2) ; 3°, the nerve undoubtedly reaches the summit (fig. 8, 3). There is no hesitation possible about 1° and 3°. In the 2nd form (inter- mediate between 1? and 3°) it is sometimes difficult to come to a decision. I bring under 1° (nerve ceasing below the summit; abbreviation —) a leaf in which there is at least one ordinary (isodiametric) cell above the extremity of the nerve (in fig. 8, 7, there are two such cells). (vL a Fic. 8.—Relation between the nerve and the summit of the leaf. n, nerve. See text. About the summit of the leaf the two differentiated borders meet each other, and we find a greater or smaller number of differentiated cells. If the nerve reaches those cells, I bring the leaf under 3° (abbreviation +. Fig. 8, 3: the nerve often reaches further than in this figure). If the extremity of the nerve is connected with those differentiated cells by one or more distinetly elongated cells, which constitute a sort of bridge among the ordinary isodiametric cells, I bring the leaf under 2° (fig. 8, 2. Trace of relation; abbreviation +). In some eases the decision is rather arbitrary, because the structure near the summit is sometimes indistinct. $36. ToorH AT THE SUMMIT OF THE LEAF.—No explanation is needed about this character. (Tooth existing, abbreviation + ; tooth not existing, abbreviation —.) § 37. NUMBER or Fruits.—See the explanation in Table XX XVIII. 40 PROF, JULIUS MACLEOD ON TEN PART IV; DESCRIPTIVE TABLES OF TEN SPECIES OF THE Genus Mxrua. § 38.—In the present part I give the quantitative description of ten British species of the genus Mnium*. For two species (orthorrhynchum, spinosum) which are rarely found in fruit in these Islands, it has been impossible to obtain fertile British specimens ; therefore I examined specimens from the continent, which were kindly given to me by Mr. H. N. Dixon, F.L.S. The description of each species includes two Tables, viz. : 1°. A Table A, in which I give the minimum and maximum values of twelve characters of the leaves in the 10th interval (see § 6; $16, Table XIII. A) of the fertile stem, and of two more characters (total number of leaves of the fertile stem and number of fruits). The limits are obtained by the measure- ment of specimens from as many localities as possible. Many of the given limits are merely approximate and are very likely to be corrected by further measurements. I hope, however, that corrections will be rather unimportant, except perhaps for three rare species (affine, orthorrhynchum, spinosum), material of which could not be obtained in sufficient quantity. As they are, the Tables may give an adequate idea of the method. 2". A Table B, in whieh I give the gradation curves of the twelve characters of the leaves. For each species the figures in the ten intervals were obtained from specimens of one patch, the curves being mean interval curves (see $ T). Therefore the figures have not the significance of exact specific figures. Gradation is, indeed, in a certain degree, variable from one patch to another. A complete study of this variation would go beyond the limits of this paper. As was remarked in $8, the relative values of the ordinates of a gradation curve are more important than the absolute values. I prefer, however, to give the absolute values, by means of which the relative values (percentage curves) may be easily calculated (§ 8)—whilst it is impossible to caleulate the absolute values by means of percentage curves. On the definition and the measurement of the characters, see Part III. ABBREVIATIONS : 1°, Length of the midrib (see § 35) : (a) Midrib short (ceasing below the summit of the leaf) : —. (6) Nerve longer (a trace of connection between nerve and summit): +. (c) Nerve as long as possible (reaching the summit of the leaf): +. 2°. Tooth at the summit of the leaf : (a) Tooth present: +. (b) Tooth absent : —. * The same method is applicable to other Acrocarpic Mosses. BRITISH SPECIES OF THE GENUS MNIUM. 41 Taste XV. Description of Mnium affine, Bland. A.— Specific limits of variation. [Cheshire, 3 stems; e Vogeso superiore, 2 stems.] As the number of specimens is so small, the figures in this Table are merely given for com- parison with further measurements. According to Messrs. H. N. Dixon and H. G. Jameson (* British Mosses, 2nd ed. p. 379), this species varies very greatly. There were no noticeable differences between the specimens from the two mentioned localities. 1. Length of the leaves: 544-824 mm. . Breadth of the leaves: 2:20-4:-08 mm. 2» a » at their base: 0°76-1:16 mm. m » cells: 30-43 y. . Number of cells: 56-93. . Breadth of the border: 47-92 y. . Number of cells of the border: 5-10. w marginal teeth : 42-106. » double marginal teeth : 0. OMNAHBR AR wb pia E a teeth on the nerve at the back of the leaves: 0. E [um . Length of the nerve: ? (indistinct). = bo . Tooth at the summit of the leaves: always existing. . Number of leaves: 24-29, 53 fruits: 1-4. = m P Co B.—Gradation of the characters. [One patch: Tatton Park, Cheshire, 3 stems.] Mean interval curves. Intervals: 1. 2: 3. 4, 5. 6. T. 8. 9. 10. Length leaves (mm.)...... 2:02 3:390 3°88 8:80 414 387 447 444 551 6038 Breadth leaves (mm.) .... 1:007 2-61] 2:90 2-58 2:64 2:45 2:01 2:58 303 2:93 25: "base mm: a. 2 ee 1:04 1:12 1:24 136 (1:10 118 1:06 0:02. 0:86 0:89 3 CELIAC) 3 ee ds 84. 3805855 1894. 932 31 23925 $30 3b, 88 Number. cella? occ... ose 5. 4070 0 a d 70. 70 3D yer *72 Breadth border (y) ...... 60: Gor 0S7 76. 78 000 789 4 72*. 0T ias border |... E cU, 9 9 9 0. 8b 18908 85. 52- TI Marginal teeth .......... 5 995 056 08. GL- .65 . 05 ? 78 62 Double marginalteeth.... 0 0 0 0 0 0 0 0 0 0 llipsnpinerve iaoe oc-+ eere 0 0 0 0 0 0 0 0 0 0 Iancih'nerve...........- F — E -— — — — — — — Tooth summit ........ eos cto FA T c^ c o4 t4 — 42 PROF. JULIUS MACLEOD ON TEN TABLE XVI. Description of Mnium cuspidatum, Hedw. A.— Specific limits of variation. [Great Britain : Borrowdale, Harleyford (Bucks), Windermere.— Europe: Helsingfors.— Canada : Ottawa.] . Length of the leaves: 39:73-7:44 mm. . Breadth of the leaves: 1:13-3:26 mm. » » F at their base: 0:62-1:26 mm. ^ » cells: 17-22 p. . Number of cells: 56-146. . Breadth of the border: 35-89 y. . Number of cells of the border: 5-9. » marginal teeth : 18-77. NOAA PP wo Oo ped UD ,» double marginal teeth: 0. 10. m teeth on the nerve at the back of the leaves: 0. 11. Length of the nerve: the nerve always reaches the summit of the leaf. 12. Tooth at the summit of the leaves: always existing. 3. Number of leaves: 11-28. 14. M fruits: 1-2. B.— Gradation of the characters. [One patch: Borrowdale, 10 stems. | Mean interval curves. Intervals: 1. 2; EA 4. 5. 6. Ni 8. 9. 10. Length leaves (mm.)...... 1:31 185 227 266 9202 3:20 861 8:86 421 441 Breadth leaves (mm.) .... 1:08 1L:44 1:65 179 1:80 L85 1:91 L88 1:80 1:66 » —base (mm.) ...... 081 0:96 100 1:01 100 096 095 0:88 080 0:76 S AMEND Sii. wo nm « 9 3. 9 —90..I9 i9 Penn Der celle i. muere 58 79 81 81 85 86 88 88 86 80 Breadth border (p) ...... 0122 8.70.7 07 70-80. EB. (40%, 02... 6S UBI DordeR ve iere « 00 9:4: 100. 10:3) 10:3: 1031. |9:8...05..80..84. 77 Maremal teeth... ss. ve 8 15 2b 82 36 96 4] 41 4l 40 38 Double marginal teeth .... 0 0 0 0 0 0 0 0 0 0 Teeth nerve ........ ee 0 0 0 0 0 0 0... 0 0 0 ngih nerve 9... 9o eei — t- +- t- +— +t + + + + Tooth summit... veces es zE T + T — ES + + + + BRITISH SPECIES OF THE GENUS MNIUM. 43 TasLE XVII. Description of Mnium hornum, Linn. A.—Specific limits of variation. [6 localities in Great Britain. | l. Length of the leaves: 5:17-8:06 mm, 2. Breadth of the leaves: 0:64-1:46 mm. 3. ie is 5 at their base: 0°55-0°98 mm. 4. 5 » cells: 14-20 pg. 5. Number of cells: 34-73. 6. Breadth of the border of the leaves (often recurved): ?. 7. Number of cells of the border (often recurved, see § 30): ?. 8. 2 marginal teeth: 46-84. 9. = double (and triple) marginal teeth : 34-61. 10. Teeth on the nerve at the back of the leaf: 4-9. Length of the nerve: nerve reaching the summit (4-), ceasing below the summit = [zr ( —), or intermediate form (+). 12. Tooth at the summit of the leaf: always existing. 13. Number of leaves: 46-64 (perhaps more, as many as 80 ?—See § 23). 14. » fruits: always 1. bo B.—Gradation of the characters. [One patch: Dungeon Ghyll, Lake District, 8 stems. ] Mean interval curves. Intervals : i 2. 3. 4. 5. 6. a 8. 9. 10. Length leaves (mm.)...... 200 200a 900 aes 9/70. 428) 405 524 6:01 6779 Breadth leaves (mm.) .... 0:53 0°63 074 0:991 111 1:26 1832 1:41 143 118 o bsimm.)-.....-. 0:50 O2 OG (i68. 0:74 0708:0777 OBI 085 084 EB NN... p. NEU C de. 1005 03:7 4. des "18 INIIDONGBlNS si eee reni 85 gp D bo: 00 oam oao TA — 72. OS Breadth border .......... p COL DOTAN, use rris F Ri E : T i ipf TE e. a Marginal teeth .......... 1:5 3 10 23 35 42 49 55 DD. Of Double marginal teeth .. . O1 L3 4 WS: 98. 808 8D AI 48. 48 lleoth;nerye. ...de eser 0 0 EDAM q0 94: mU 04 76 9 length nerye*........... — +- - — — t- +- +- +- +- Bob minmit......:-..-- T ete CL dubio qub c d dE * In the intervals 1-8 all the leaves were —, except: 1°, in each of the intervals 2, 7, 8, one leaf +; 2°, in interval 6, one leaf +. 44 PROF, JULIUS MACLEOD ON TEN Taste XVIII. Description of Mnium orthorrhynchum, Bruch, Schimp. & Guemb. A.—Specific limits of variation. [ Boulder, Engstligen Falls, Adelboden (Switzerland).—H. N. Dixon dedit. | 1. Length of the leaves: 3:00-5:41 mm. 2. Breadth of the leaves: 0:30-1:03 mm. 3. » T » at their base: 0:31-0:56 mm. 4. d. » cells: 12-15 y. 5. Number of cells: 21-62. 6. Breadth of the border: 23-45 w. 7. Number of cells of the border: 3-7. 8. j marginal teeth: 22-38. 9. m double marginal teeth: 12-31, 10. m teeth on the nerve at the back of the leaves; 0-11. 11. Length of the nerve: in 48 leaves I found the nerve reaching the summit; in 2 leaves merely a trace of connection between nerve and summit. 12, Tooth at the summit of the leaves: always existing. 13. Number of leaves: 25-61. 14. j fruits: always 1. B.—Gradation of the characters. [One patch : Adelboden, 8 stems.] Mean interval curves. Intervals: 1. 9. 3. 4. 5. 6. Ë 8. 9. 10. Length leaves (mm.)...... 0776 099 1:23 141 1:59 2:00 2:84 2:91 319 3:96 Breadth leaves (mm.) .... (0:36 0:42 0:50 0:56 0:65 0°84 0:93 0°93 083 0:64 5 Dasa (W) o) s. 0:381 034 036 038 0'41 048 049 0'47 042 0'40 3j COS (ies cas. TL T 12 12 13 14 14 14 14 13 Number cells ........ c aM wm yz p M 5» wo 60 b2 o Breadth border (u)........ 24 27 28 28 28 31 33 34 36 33 NPOLIMIDOLGOD T ree 5 6 6 6 6 6 6 6 6 5 Marginal teeth .......... 1 2 5 (i 10 15 21 26 26 29 Double marginal teeth .... 0 04 18 3 5 WwW 15 18 19 20 Teeth nerve...... EAA s 0 0 0 0-702059 1 2 3 4 een pth Nerves. ou. ss. .« t— t- +— +- t- +t += +- +t + Tooth summit...... Iw Ue E b LGB + + +4 BRITISH SPECIES OF THE GENUS MNIUM. Taste XIX. Description of Mnium punctatum, Linn. À.— Specific limits of variation. [Seven localities in Great Britain.—Island Griisen (Sweden).] l. Length of the leaves: 4:35-9:21 mm. 2. Breadth of the leaves: 2:71-6:48 mm. 3. be jt ji at their base: 0:44—0:82 mm. 4. 2 » cells: 36-65 u. 5. Number of cells: 61-111. 6. Breadth of the border: 82-195 p. 7. Number of cells of the border: 8-11. 8. T marginal teeth : 0. 9. 2 double marginal teeth: 0. 10. E teeth on the nerve at the back of the leaves: 0. 45 11. Length of the nerve: the nerve reaches the summit in the great majority of the cases, It always reaches the summit in the second leaf above the longest one and higher upwards. 12. Tooth at the summit of the leaves: a tooth is almost always existing. It always exists in the second leaf above the longest one and higher upwards. 18. Number of leaves: 7-19. 14. 5 fruits: 1-2. B.—Gradation of the characters. [One patch : Llanberis to Bangor, 11 stems.] Intervals: 1. 2. 8. 4, 5. 6. yj^ Length leaves (mm.)...... 2:10 2:56 8:58 3863 459 498 578 Breadth leaves (mm.) .... 1°68 198 298 317 335 359 3277 Bo base (mm.) ...... 0:58 0:63 0:64 0:64 0:64 0:62 0-64 a; cs GE UT) REN 90- 20 0000 46 40 47 A5 NHMDOPGOIB. T. eel. 38 38 52 65 70 TE 80 Breadth border (y). ....... SS ADD IIIS 0107. 194 :190 190 Cells border ©. 2. os. se ek: 02 QNT so ess 93 90 p5 Marginal teeth 55.42... 0 0 0 0 0 0 0 Double marginal teeth .... 0 0 0 0 0 0 0 BREST nerves... eec ete 0 0 0 0 0 0 0 I[ienpth nerve... ......... — = — Lo nl ec ll/'ooth summit .......:..- tT— +- +— +— + te dum +— 2 t ACE, TO" TIT NER P ; E 1 E : 46 PROF. JULIUS MACLEOD ON TEN TABLE XX. Description of. Mnium rostratum, Schrad. A.—Specific limits of variation. [Great Britain: Alston to High Force, Dunkeld, Llangadock, Whaley Bridge. ] . . Length of the leaves: 4:35-8:32 mm. 2. Breadth of the leaves: 1:80-3:19 mun. ” ” 1 2 3. ” ” » 4 5 at their base: 0:73-1:24 mm. 5. Number of cells: 62-97. cells: 23-34 u. 6. Breadth of the border: 69-125 y. 7. Number of cells of the border: 6-12. 8. 5 9 T double marginal teeth: 0. Oa 11. Length of the nerve: nerve always reaching the summit (one leaf doubtful). 19. Tooth at the summit of the leaves: always existing. 13. Number of leaves: 12-21. 14. » fruits: 1-4. B.—Gradation of the characters. teeth on the nerve at the back of the leaves: 0. [One patch: Alston to High Force, 9 stems.] Intervals : Length leaves (mm.) ...... Breadth leaves (mm.) 2.1 -Dase (m). 2:21 quoa EMISIT S Number cells ....... Vis Breadth border (u). ....... O DOTAS. esie rene Marginal teeth .......... Double marginal teeth .... RIBOD EDEL VG. LAN Le ces i sve ono NERVE 5 opos Ces vie OON BAITIT a o errors l. 223 132 2. 2:95 1-80 075 29 55 89 9 0 0 TE JJ 3. 30i D 0-73 4. 4:07 2-04 0:78 5. 4:04 2:18 0775 3l 59 88 8. 5:59 2:40 0:84 30 73 90 9 marginal teeth : ? (teeth rather numerous, but partly obsolete). BRITISH SPECIES OF THE GENUS MNIUM. ; 4T TABLE XXI. Deseription of Mnium serratum, Schrad. A.— Specific limits of variation. (Great Britain: Ben Lawers, Ingleton, High Force, South Wales, Gilsland Alston, Craven.— Canada. | . Length of the leaves: 2:-49-4:71 mm. . Breadth of the leaves: 0:59-1:31 mm. E " » at their base: 0:88-0:71 mm. E 5 » cells: 17-25 gp. . Number of cells: 98-57. . Breadth of the border: 36-101 p. . Number of cells of the border; 5-9. 5 marginal teeth: 11-29. OPNP PP a double marginal teeth: 6-21. 10, i teeth on the nerve at the back of the leaves: 0. ll. Length of the nerve: nerve always reaching the summit in Great Britain. (Canada: ZONE) p 12. Tooth at the summit of the leaves : always existing. 13. Number of leaves: 16-36. (Great Britain: 21-36. Canada: 16-24.) 14. » fruits: 1. ; : . B.—Gradation of the characters. [One pateh : Ben Lawers, 9 stems.— Barker leg., July 1870.] Intervals: ^ 1. | 20 E b 16 c8 470. 10. Length leaves (mm.)...... 144 1°70 1:96 215 2:87 92:62:9-79 3:09 3-48 408 Breadth leaves (mm.) .... 0°63 078 087 101 1:09 119 126 1:29 1-29 113 Ec pase (im. ...... 047 0:50 051 053 0:56 059 059 0:57 0:52 052 come EY (79 oa. s ess l/ eee S0 2].* 99 10h 08.95 Ves P. INTIBDEROOISS e eere neret 37 40 4l ae AG AT, 50 50 50 48 am Breadth border (g) ........ . 54° Sees Ole SO . 08 690 1709 901 76. 78 ISTIS DOLdBE sik. wes cscs 85 8 9 D 88 Shee INB 75. TO Marginal teeth .......... 0- 0 E One T6 5 84 12 T7 18 16 Double marginal teeth .... 0 0 0 05.08. Stow age, 12" '12 1l MEPRh CTV 2.0005 ees n 0 0 0 0 0 0 0 Wength nerve .........-++ t- t— F- F- +- +E tt +- + ME Tooth EINIDES 2.25508 + + + uen p HE + FiF g SMS t e i oE JM ee EM : 1 5 i. vit: zr on 2^ DP E EEE E Pre te Ee, a ee DR oa rm n 48 PROF. JULIUS MACLEOD ON TEN TABLE XXII. Description of Mnium spinosum, Schwaeg. A.—Specific limits of variation. [Switzerland : Engstligen Falls, Adelboden ; H. N. Dixon dedit.—Styria : in sylvis prope St. Nicolai, in vallo Solk.— Very rare in the British Islands.] » ” ” $ 0 - O0 Ov RD WY » mn e » "m mw 2 nm ” . Length of the leaves: 5:01-8:10 mm. . Breadth of the leaves: 1:55-2:84 mm. f j at their base: 0:68-1*06 mm, cells: 21-31 p. » . Number of cells: 65-96. . Breadth of the border: 62-92 y. . Number of cells of the border: 10-13. marginal teeth : 43-72. double marginal teeth : 53-47. teeth on the nerve at the back of the leaves: 2-12. . Length of the nerve: nerve always reaching the summit. . Tooth at the summit of the leaves: always existing. . Number of leaves: 24-33. fruits: 2-6. B.—Gradation of the characters. [One pateh : Adelboden, 8 stems.] Intervals: 1. 9; 3. 4. 5. 6. Ts 8 9. 10. Length leaves (mm.) .. Breadth leaves (mm.) 5. base:(mm.).... » cells (p) Number cells Breadth border (u)... . Calla Dorder. se: eee Marginal teeth Double marginal teeth .... Higeth:nerve... es. Length nerve ........ Tooth summit. Ls» 6:60 nios Tog 280 455 087. 6:91 616 6088 6:93 7:26 O75 1:41 2:42 2:80 286 2:81 276 274 266 2°52 m. Oel 095 114. 10 110 102 0°95 0902 089. 090 PU 15 22 27 29 29 29 29 29 28 28 nic ae 90 59 R4 91 91 90 88 88 87 82 iia o OF a 75 16 73 TE 75 73 74 76 Hio. SLE DE ll 1l IU 1 i 11 11 ll Oe ic eee 18 35 44 47 52 55 57 63 63 0 2 12 24 2m 80 $4 88 40 . 42 re ie 0 001 1 Lp 245 OI 40 48 76 n +-+ + Rock cb ok o4 c4 c Be Fo + + + + + + + + + DET WT BRITISH SPECIES OF THE GENUS MNIUM. 49 Taste XXIII. Description of Mnium subglobosum, Bruch, Schimp. & Guemb. A.— Specific limits of variation. [Great Britain: Benson Knot (Kendal), Whaley Bridge, Hathersage, Derbyshire.] . Length of the leaves: 3:62-7:33 mm. : Breadth of the leaves: 2:95-5:50 mm. at their base: 0:58-0:98 mm, d » ” ” A » cells: 38-54 y. . Number of cells: 71-100. . Breadth of the border: 53-85 p. . Number of cells of the border: 4-6. jd marginal teeth: 0. OM ASMP ow double marginal teeth: 0. teeth on the nerve at the back of the leaves: 0. ” m" e ” . Length of the nerve: nerve never reaching the summit. (Very rarely a trace of T -— connection between nerve and summit in the third leaf above the longest one. Nerve sometimes reaching the summit in the phyllomes of the perianth.) 12. Tooth at the summit of the leaves: never existing. (A trace of a tooth in the above- mentioned exceptionalleaf. The phyllomes of the perianth sometimes with a tooth.) 13. Number of leaves: 7-i4. 14. a fruits: 1-2. B.—Gradation of the characters. [One patch: 11 stems with about 70 leaves in good condition ; many leaves were spoiled and not measurable, especially in the intervals 1-5. Benson Knot, Kendal.] Intervals: 1. 2. 3. 4. 5. 6. ts 8. 9. 10. Length leaves (mm.)...... 1:34 2:04 2°58 3:08 2:06 3°65 413 4°77 6:00 579 Breadth leaves (mm.) .... 1:07 1:59 1:99 2°56 2°31 2:86 3:28 8:80 3870 441 » base (mm.) ...... 0-56 0:05 073 073 0-69 074 068 071 077 078 COMED) Eiee 31 36 38 42 3s 41 41 45 44 47 Mmmm ber Cols Ging. cs. ss 20. Wl 500. 50 .00 20 7B 75> 9] Breadth border (u)........ 59 606 65 08. 65. 05 2027 | Bl 63 65 BULbonder $3.60 4 4 4 5 5 5 5 5 5 5 Marginal teeth .......... 0 0 0 0 0 0 0 0 0 0 Double marginal teeth .... 0 0 0 0 0 0 0 0 0 0 ENDE. .oeeecnnennn 0 0 0 0 0 0 0 0 0 0 length nerve ........ se AS - E imr - -= lg mit. se ooo eia oo — — — — — — — — _ LINN. JOURN.— BOTANY, VOL. XLIV, E 50 PROF. JULIUS MACLEOD ON TEN Taste XXIV. Description of Mnium undulatum, Linn. A.— Specific limits of variation. [Great Britain: Manchester ; Bolton Abbey. | 1. Length of the leaves: 6:903-15:58 mm. 2. Breadth of the leaves: 1:71-2:35 mm. 3. » ^ 5 at their base: 1:02-1:47 mm. 4. » » cells: 17-28 y. 5. Number of cells: 76-103. 6. Breadth of the border: 40-97 p. 7. Number of cells of the border: 6-11. 8 marginal teeth : 84-134. 9 double marginal teeth : 0. ” ” 10, P teeth on the nerve at the back of the leaves: 0. 11. Length of the nerve: nerve always reaching the summit. 12. Tooth at the summit of the leaves: always existing. 13. Number of leaves: 33-58 (or more ?). 14. D fruits: 2-8 (the fertile stem is sometimes ramified, the branches being fertile; the fruits of such lateral branches are not taken into account). B.—Gradation of the characters. [Two patches: Manchester, 7 stems with about 210 leaves, the leaves above the longest leaf being excluded. ] Intervals: 1. 2. 3. 4. 5. 6. i 8. Oy 16 Length leaves (mm.) ...... 159 235 3:72 540 627 678 823 854 894 10:15 Breadth leaves (mm.) .... 1°55 165 189 225 2:20 212 222 218 212 209 base. (MM) «e. ses 1:32), -1:89.°. 102 1:82...1-80 1/70 1:69 1:54 148 1:26 5 Gell (A) obesse vs 22 22 22 22 21 21 21 21 21 21 INOMIDEY COMA Ls 2 st sas 850. 71. 16. Ole 98. 4 89...) 00a 89. . 80 Breadth border (#)........ 81 82 84 85 8T 86 .. M5 88 488 78 Welle border 2. xs... 100195: 122 11a 19. | IL... 14 LE Al 9 Marginal teeth .......... 29 48 66 87 96. 102 109. 115 .116 119 Double marginal teeth ..,, O 0 0 0 0 0 0 0 0 0 APELI DOTO. Wi... 3s SFO 0 0 0 0 0 0 0 0 0 Mongth (3 42) 2 BR SNR T— +- +t +24 +t +t +4 +t + E TOt SUING sissies. vi + + + + + + + + E + BRITISH SPECIES OF THE GENUS MNIUM. 51 PART V. TABLES OF IDENTIFICATION. § 39.—In the present part I give tables of identification for the ten species of Mnium which are described in Part IV. — The British Flora includes four species (lycopodioides, Schwaeg., riparium, Mitt., stellare, Reich., cinclidioides, Blytt) which are not mentioned in the tables ; these are rare (or very rarely or never found in fruit) in the British Islands. In each table the minimum and maximum limits of the character under consideration are given separately. In the descriptive tables (Part IV.) the figures are given as they were observed. As the limit-values are hitherto merely approximate, they are MODIFIED in the tables of identification in the following way :— I'. The minimum figures are diminished by about 5 per cent. of the observed value, and for three species (affine, orthorrhynchum, spinosum, see $ 38, p. 40) by about 10 per cent, Z'. The maximum figures are augmented by about 5 per cent. of the observed value, and for the three mentioned species by about 10 per cent. By these modifications (which are arbitrary) the limits between which variation may occur are widened by about one-tenth (or one-fifth) in order to diminish the possibility of mistakes in identifying a given specimen. Despite this, it is advisable to be prudent in the use of the tables and to allow a wide range of error, as we have done in § 17 (see p. 26), in which the THREE METHODS of using the tables are illustrated by examples. ABBREVIATIONS : see $ 38, p. 40. TABLE XXV. (see § 25). Length of the leaves in the 10th interval (mm.). Minimum. Maximum. CURATO: ESPERE 2°36 RCNP OUUM ...: ELS 4:95 orthorrhynchum ........ 2:70 orthorrhynchum ........ 5:94 SUO IQUOSQIN seo RE 3:44 subglobosum | .......... 7T'10 QUADIdQTU ee esci ss 3°55 cuspidatum |. esses. 7T'8l Unda woes occ es 4:12 BOW o ee bw 8:46 piegata Sye iei die. eins 4:12 DEA aaran ri C. OR LU 8:73 PROUN e i oea ae 4:50 EORUM a e C oo B RORIS 8:91 HET t E S ER 4:90 Ce Rees RR 9:17 OH LS NCC 4:90 BELAUN Lies enin 9:67 undulatum ........ eee, 6:60 undulatum eee 16:36 52 PROF. JULIUS MACLEOD ON TEN TaBLE XXVI. (see $ 26). Breadth of the leaves in the 10th interval (mm.). Minimum. orthorrhynchum ........ 0:27 BOETHII oe eer ee 0156. ZU hU dose TREAT 0:61 eusmidatum .... ee 1:07 WRMLOMETE ie E e 3's 0 + 1:40 UOU nei 1:62 POO i Las ARCEM Ly e gna uds Ie E aE 1:98 punctatum. foie a fon OMG subglobosum .... 64... 2:80 | Maximum. orthorrhynchum ........ serratum does PRA ME TA hornum undulatum ......... vs. POMEM i seas A ow TOS Blum ori. ovv Y cusptddtum- e see ee affine subglobosum punctatum TaBLE XXVII. (see § 27). Breadth of the leaves at their base in the 10th interval (nim.). Minimum, orthorrhynchum ........ 0:28 BS eis eco ° 0:36 punciatum ....ceeseees 042 WOPRTSS ends port. 0:52 subglobosum — .......... 0:55 euspidatum 5... vs 0:59 SPANOBUIE S E AE 0-61 E OI: outed: oats: 0.0 e pnr 0:68 Job gupo 211 E TC PECTUS 0:69 undulatum ooo. 0:96 Maximum. orthorrhynchum ........ hs mE PETERE IT YE punctatum ......eee ees DORRI Silas aaa re ES ED subglobosum |—........ su. EPIO S edic alae a affine rostratum CUSAAAEUM . i.. eee se undwlatum ............ TasgLE XXVIII. (see § 29). Breadth of the cells at the place of the greatest breadth of the leaves in the 10th interval (4). Minimum. orthorrhynchum |... ..... TI IURI LOSS E ER CER E 13 0177 117/71] 77, EQUES e's» 16 LLLI LUCUS S PPP TENE 16 Qr Ua n Lv au CREER LL AIL C1 CORO 8 ear eee 18 FOE Deere tee 22 QUE eRe ss ue 27 punctatum oo cccccceceses 94 subglobosum |... veru. 36 Maximum. orthorrhynchum hornum cuspidatum |... TE HI p 7, an ase OTP is a 5,0 rostratum spinosum affine subglobosum punctatum ...... TT . BRITISH SPECIES OF THE GENUS MNIUM. ; 58 TABLE XXIX. (see § 28). Number of cells at the place of the greatest breadth of the leaves in the 10th interval. Minimum. | Maximum. orthorrhynchum ...... D 19 serratum: na sus 60 serratum ow re es 27 orthorrhynchum | ........ 68 Topnum o ETE 32 hornum soe a ente "i ARES Uer ELM 50 affine iesea EN r 102 CURPHUTTUmu n. nats eh + 6 ote 53 rosratum EE. 102 DURUM S Ee cer 58 | Spinosum, creme he iis 106 R A E iie 59 undulatum : 6500s are 108 n e te vam 59 subglobosum ...... Seea L2 RISD LOUORUM oiiaao 67 punctatum sad deksesa T INQUIT. eerte 72 duamidatum | LESE. 154 TABLE XXX. (see $31). Breadth of tlie border at the place of the greatest breadth of the leaves in the 10th interval (gu). Minimum. Maximum. TODO erudi R orthorrhynchum.......... 50 OPtAOrrhiynohum o e... AE suboloboswmn. o 31:5:90 CUSmOGNMAE <- o e nier n B5 Olptdatum ieaie ee eae 94 EURO one rene Riri OE ne se. . sen “aR eS Oe 102 RIO cc a vs eie e s ae morum: o e is ats sch 102 Cdi SO ID. 42 undwatum heres, les, 102 SUDQLODOSUM ioes nereo ee 50 maium EE Loma 106 Gul UCUGOSN MONIS 56 OBL GUI. ). EO a 132 EUG ine a oa 65 punctatum ..... E Tus 205 unciam E rene 77 OPA r.c sara |. shal dy TaBLE XXXI. (see § 30). Number of cells of the border at the place of the greatest breadth of the leaves in the 10th interval. Minimum. Maximum. ROPNE. o c ae eh A ? | subglobosun Es 7 orthorrhynchum |....... ss. 2 | orthorrhynchum — ........ 8 subglobosum . 6. es v. a 3 | cuspidatum s osm dela ds ssc 10 affine ARGO eae edo T 4 | CIAL) o M rer. 10 cuspidatum ......... ee 4 | nc c MM d. ll BOUT ATU. 21s o E 4 | punctatum ... sse 12 torgum LE 5 | Damn CORRI ree 12 hatum 5 sS 5 PORT GUT. eese es ANO IBIR CHITI. ciis cei il aang T i PROSITE eee ae:e = 14 DEnonun 2:10. 5.5 E 9 | hornum ........ nc Í x 2 : PROF. JULLUS MACLEOD ON TEN TABLE XXXII. (see § 32). Number of marginal teeth of the leaves in the 10th interval. Minimum. HUQ Uie etn 0 subglobosun i 0 TONPFQUIEE een n? S8 ratum. vss. MERC DA UM: 10 CURD IORI renes 17 orthorrhynchum .......... 20 qna. vv. DUE ES 38 pub] C M 39 BONUM has ere 45 ORONIA sss 80 Maximum, punctatum we ce ese 0 GUD QLODOBUM SIR Pao 0 POFAM
E 89
P R eata 117
tfhdulütum. ir i aro eia 141
roBirddum $ S HU. us n?
Taste XXXIII. (see § 33).
Number of double (and triple) marginal teeth of the leaves
Minimum.
Ws 1 e o. 0
DUSAGATUm VERDE ees 0
tandi i ae eoe sissy 0
WOSU GUI. ee re sc ess 0
SUOGLODOBINTE 10303. 0
UndidNm el les 0
aOR C cuu. 5
orthorrhynchum —.... LL... 11
Lydi ni dte nc CORE .90
ADU Ty ooh ec shee O
in the 10th interval.
TaBLE XXXIV. (see § 34).
Number of teeth on the nerve at the back of the leaves
in the 10th interval.
Minimum,
GIG Mares oor co ino 0
OHSPIGOURDI 5. osse de 0
orthorrhynchum o... 0
OV Lo d RS 0
EINE Verre 0
OUI aq d M MEE 0
BUDUIOUONUTT eens. oc Near 0
ODES iss E ra 0
polano 16 000 GO HONO 1
TOFU etos EDD Va 3
Maximum.
EE d.i mir. 0
DUSDITDUHTAEA edes 0
pundtatum dy 6e: 0
TÜUBLPRDUP. Ugo... 0
Subploboku 3. eve. 0
UTOR. lli. 0
naana I A Ute a ev 22
orthorrhynchum vv ceca 34
EOT a E dios dri. 51
Wn I) VADO, 64
Maximum.
DUIS E I is 45 TREAT A 0
cuspidatum © s.es., 0
punctatum ........... ls. 0
LEEUUM 4007 0 oi 0
E c e Ea, 0
SUDOIDDDFU esu. 0
UNGURU Jo ee er 0
orthorrhynchum ....... LL. 12
BOVIOSUTE a ans 13
HORE Gao kee. ss 20
them exactly. See Table XX., A.
x In Mnium rostratum marginal teeth always exist, but it was impossible to count
CIE TM - AGS i i a a e
BRITISH SPECIES OF THE GENUS MNIUM. 55
TaBLE XXXV. (see § 35).
Length of the nerve in the leaves of the 10th interval. (Nerve
reaching the summit, +; a trace of connection between nerve
and summit, +; nerve ceasing below the summit, — .)
Minimal | Maximal
length of the nerve. | length of the nerve.
VGL ERE AR MUR P aubglobosum. i Sii... —
iranta i Ee A 5 cuspidatum Sail... oc. +
MFT Tp o o Ee RRS — Aornim Le vex +
E ens ello rne — orthorrhynchum l...a.. +
RAC HOR: oe — | Ennoiatum . aod ms +
orthorrhynchum ........ EE | Zostratum.- ae a +
EROR D a o nr | ERs S25 Mee ly wate +
CUSDIGTE S iso's tis» eh + | BAAD LITE. 0 e oin +
Gn rp PRESE S ha + nuam 23e. +
LSOTLITI a E 5 cus. ie ES l pU 3... a tn R
TABLE XXXVI. (see § 36).
Tooth at the summit of the leaves in the 10th interval.
A tooth at the summit of the leaves in the 10th interval exists always in
all the species, except in two of them, viz. :—
1°. Mn. punctatum. The tooth exists ordinarily, or there is a trace of
a tooth. The absence of a tooth is rather rare. The first leaf
above the longest one is variable in the same way. In the second
leaf above the longest one and further upwards I always found
a tooth.
2°, Mn. subglobosum. . Never a terminal tooth in the 10th interval, nor
in the two leaves above the longest one. In a few cases a trace of
a tooth exists in the third leaf above the longest one. A tooth may
oceur at the summit of the phyllomes of the perianth.
See, with reference to this character, the specific tables of gradation
(Tables XV.-XXIV., B).
56 PROF. JULIUS MACLEOD ON TEN
TaBLE XXXVIT. (see § 23).
Number of leaves of the fertile stem.
In this table a rather great possible error is to be admitted. The figures
indicate the number of leaves which still remain in a stem which bears a ripe
or almost ripe fruit. In such stems the lowest leaves have almost always
disappeared. (See $ 23.)
Minimum. Maximum.
CO es rss 6 subglobosum |............ 15
subglobosum. =... 8... ee es 6 | PUNAM eroi c cese 20
QIBDUIAUUTE his 5 hs. se eire 10 | haa a E e Ee 22
PORVOO a 1l | euspidatum o cece s 30
anaia Ce 00 cree i a's sra 15 DUME LUE a i mene >. 30
a a ee A E P 290 | NOSU A oes cmi E
orthorrhynchum | ......... 25 | MIETEN E te ek ef co 38
OE VA E 25 | undulatum ......... suus. 61
undulatum |... ees. 31 | orthorrhynchum | ......... 67
BOMUN E I TRA INST sss 46 | OTM anra E 67 (or
| more).
Tastre XXXVIII. (see $37).
Number of fruits at the summit of the fertile stem (see § 37).
In several species this property is very variable. The figures in this table
are to be considered as a rather incomplete information. Besides the fully-
developed fruits, abortive fruits (length 2 mm. or longer) which are easily
distinguished from sterile archegonia are sometimes found; such abortive
fruits have always been counted and brought into account. In all the
species, stems with one fruit will likely be found. Fruits which sometimes
occur at the summit of lateral branches (Mn undulatum and perhaps other
species) are not taken into account.
Minimum. | Maximum.
Zn c Um ELE T A l | ey cr VAPE D RR A MEO l
iy Th UIC): S e Lr cen l | orthorrhynehum sss... |
[OCA SSO MAR l CTO E in Apka l
orthorrhynchum .......... 1 QUEDIQQPAMT Ivo voces 2
DURO UU erro Y BUncauum. oso 2
E Aa a TREE A ER T 1 subglobosum .............. 2
MAQUI S T 1 WL e ect rers 4
$ubglobONIm S eres X u a AT 4
SANOMM ae Ge Pein es io. 2 SARUM e stb s+.» 6
undulatum ........ ues hoe wndglatim eee io enn 8
BRITISH SPECIES OF THE GENUS MNIUM.
2
1:
Fro. 9.— Marginal teeth (see p. 58).
1,2,8: Mnium rostratum.
4,5: Mnium affine.—4. Part of the margin of a leaf,
with l-cellular, 2-cellular, and 3-cellular teeth ;
5. A 4-cellular tooth (rare).
LINN. JOURN.—BOTANY, VOL. XLIV.
57
TEN TT IR
x
58 ON TEN BRITISH SPECIES OF THE GENUS MNIUM.
TABLE XXXIX.
Number of cells of the marginal teeth. (With reference to
M. affine and rostratum.)
In order to test the practical value of the method, I have tried to identify
several specimens of each species by means of the Tables XXV.-X XXVIII.
I always succeeded, except with some specimens of M. affine and rostratum.
These species cannot be confounded when the classic characters of the fruit
and the inflorescence * are taken into account. But when we limit ourselves
to the characters of the leaves, there is frequently some difficulty in the
determination.
Using merely the figures of the longest leaf of one fertile stem (first
method, see $17), some specimens of both species could be determined.
But the figures of other specimens were not decisive ; after all the tables had
been used, I was still hesitating between affine and rostratum. Therefore
I have recourse to a supplementary character.
In both species, the longest leaves (10th interval) of the fertile stem are
adorned with numerous marginal teeth, which are always simple (never in
pairs). In M. rostratum, about the place of the greatest breadth of the
longest leaves, the salient part of each tooth always consists of one cell
(fig. 9: 7, 2, 3). In M. affine, at the mentioned place, the salient part of
the teeth consists of 1, 2, 3, or even 4 cells. Among those teeth, one always
finds some teeth the salient part of which consists of more than one cell
(Bg. 9: 4, 5).
All this may be shown in the form of the following table :—
Minimum. | Maximum.
a o 1 cell. | rostratum 1 cell.
i os ean an LLL 1 cell. | n E Na ae ene ng 4 cells,
* M. affine : lid of the capsule conical, apiculate ; inflorescence dioicous.
M. rostratum: lid of the capsule with a long, straight, or curved rostrate beak ;
inflorescence synoicous,
See Drxon and Jameson, * British Mosses, second edition, 1904, pp. 379, 381.
THE HETERANGIUMS OF THE BRITISH COAL MEASURES. 59
The Heterangiums of the British Coal Measures.
By D. H. Scorr, M.A., LL.D., F.R.S., FES!
(PLATES 1-4.)
[Read 19th April, 1917.]
llzrERANGIUM is a genus of Carboniferous plants based on petrified speci-
mens with the anatomical structure preserved ; it is now classed among the
Pteridosperms. The characters of the genus are briefly as follows :—
Stem monostelie; vascular cylinder a protostele, the primary xylem, inter-
spersed with conjunctive parenchyma, extending to the centre and consisting
throughout of elongated tracheides ; peripheral strands of stele and leaf-trace
bundles mesarch,
Secondary wood and bast formed. Tracheides (apart from the protoxylem-
region) with multiseriate bordered pits.
Leaf-trace bundles one or more for each leaf,
Outer cortex consisting of alternating radial bands of sclerenchyma and
parenchyma.
Inner cortex, or pericycle, or both, usually with plates of sclereides.
Leaves (when known) large, compound, and of the Sphenopteris type.
This definition, while correct so far as it goes, will be found to need some
emendation when the results of the present investigation are taken into
consideration (see p. 99).
The genus Heterangium was founded by Corda in 1845, on some fragments
from the ** sphzero-siderites ” (coal-balls) of Radnitz in Bohemia, probably of
Middle Coal-Measure age (Corda, 1845). Corda’s specimens showed little
more than portions of the wood ; somewhat better examples of the species,
H. paradoxum, have recently been obtained by Kubart from the same
material (Kubart, 1911).
In 1873 Williamson described in detail a fossil plant from the Pettycur
deposits, near Burntisland, of Lower Carboniferous age; he referred his
specimens to Corda’s genus, under the name Heterangium Grievii (William-
son, 1873). This has now come to be the best-known species of the genus, as
it is also the most ancient *. Williamson’s reference of his plant to Corda's
genus was criticised by Renault (1879, p. 277), but has been confirmed by
subsequent writers (Solms-Laubach, 1887; Kubart, 1911), and H. Grievii is
now for practical purposes the type-species of Heterangiwum, though tech-
nically 77. paradoxum, Corda, claims this position.
Mucb later, in 1887, Williamson deseribed a species of Heterangium from
the Halifax Beds (Lower Coal Measures). This was H. tiliwoides, a form
* See also Williamson & Scott, 1895.
F 2
os me
60 DR. D. H. SCOTT ON THE HETERANGIUMS
remarkable for its perfect preservation, especially as regards the phloem
(Williamson, 1887; Williamson & Scott, 1895).
These two species, H. Grievii of Lower and H. tiliwoides of Upper Car-
boniferous age, were the only British species recognised by Williamson in
his published works.
In 1890, however, he described some specimens from Dulesgate (Lower
Coal Measures) under the name H. Grievii, and thus made this species range
from the Lower to the Upper Carboniferous (Williamson, 1890). Williamson
at first thought that some, at least, of these Dulesgate specimens were a
different species, and gave them in MS. the name 77. Lomazii, but in his
published account all are included under the name H. Grievii. The difficulty
of identifying plants of such different horizons does not seem te have troubled
Williamson at all. In the joint paper by Williamson and the present writer
the same nomenclature was unfortunately adopted, though some distinctions
between the Dulesgate and Burntisland forms were pointed out (Williamson
& Scott, 1895, pp. 750 & 753). At the same time a Dulesgate specimen,
differing somewhat from the others, was given the provisional name of
H. cylindricum, though its claims to specific distinction were regarded as
doubtful (Z. e. p. 765).
In my ‘Studies in Fossil Botany’ (1st ed. 1900) it was recognised that
Williamson’s Dulesgate specimens of “ Heterangium Grievii” probably repre-
sented a separate species (Z. c. p. 345) ; in a second edition (1909, p. 410)
Williamson's MS. name, HM. Lomawii, was definitely adopted for them. No
full or critical account of the Dulesgate Heterangiums has, however, yet
been given, and it is one object of the present communication to supply this
omission.
The stimulus to undertake a revision of British Coal-Measure Heterangiums
was in part due to the discovery, in 1912, by Mr. Lomax, of a fine specimen
of Heterangium from the coal-balls of Shore, Littleborough. This plant
presents some remarkable features, and has aided in the interpretation of
other forms ; it will be taken first in the description.
The important discoveries of Dr. Kubart (1908; 1911; 1914), who has
investigated a number of forms of Heterangium and Lyginopteris* from
Upper Silesia, also render a survey of the British Coal-Measure species
desirable. A consideration of Dr. Kubart's results (as yet only published in
a preliminary form) will be postponed to the end of the paper; at the
same time the species of Permo-Carboniferous age, previously discovered
by Renault, will be dealt with.
Some interesting points of comparison with the Lower Carboniferous
genus Rhetinangium, discovered by Dr. Gordon (1912), also arise.
* Potonié’s name is adopted in preference to the familiar Lyginodendron, Gourlie, for
reasons fully set forth in Seward's * Fossil Plants,’ vol. iii. pp. 96-38 (1917).
TRAY M wae Y EE T TEE E Y. N
OF THE BRITISH COAL MEASURES. 61
HETERANGIUM SHORENSE, sp. nov.
The one specimen on whieh this species is based was discovered by
Mr. Lomax and his son on August 12, 1912, in material from Shore, Little-
borough. The fragment was 1°5 inches (3:75 cm.) long, and a series of
15 transverse sections was cut (now numbered 2781-2795 in my collection).
It is a fine stem, 17-18 mm. in diameter, the wood of the stele, though
almost without secondary thickening, reaching 7 mm. in diameter (Pl. 1.
Phot. 1; Pl. 3. fig. 1). The general outline of the stem is approximately
cireular, with more or less marked prominences where a leaf-base is met with.
The cortex is for the most part well preserved; it is 3-4 mm. thick. A
considerable part of the pericycle is also preserved, often in contact with
the inner edge of the cortex. The phloem has almost wholly perished ; the
gap between pericycle and wood was no doubt caused partly by shrinkage
and partly by actual destruction of tissue.
Stele.
We will at first confine our attention to the wood, which in three of the
sections is almost or quite complete (Pl. 1. Phot. 1; P1.3. fig. 1). The great
feature of the primary xylem (and little else is present) is the grouping of
the tracheides in a large number of definite packets, separated by a narrow
reticulum of conjunctive parenchyma. ‘The tracheal packets commonly
contain from 6-15 elements each, and range from about 400 to 650 w in
diameter. In the middle part of the wood they are fairly isodiametric, while
towards the periphery they become lengthened in the radial direction. "These
peripheral groups are very distinct, as is usual in the Coal-Measure species.
The deep clefts between them, seen in some places, are merely due to
shrinkage. The surface of the wood is somewhat ribbed, the peripheral
groups being convex (fig. 1).
The parenchymatous reticulum between the tracheal packets appears
narrower than it should be, owing to a partial collapse of the cells.
The tracheides of the metaxylem are generally of large size; there is,
of course, much variety, but we may take 200 pu as an approximate mean
diameter. "Towards the outside, i. e. in the neighbourhood of the protoxylem
of the peripheral strands, the tracheides become rapidly smailer, down to 25 u
or 20 p. The preservation is such that it is not, as a rule, easy to determine
the position of the actual protoxylem. Where this can be fixed it is found
near the outside, separated by from two to four small elements from the outer
edge of the wood (P1.3. fig. 5). The question may still arise whether all these
centrifugal xylem-elements were really primary, for a certain amount of
secondary growth had taken place—at least locally. On one side of the stele
there are a few irregular layers of radially arranged tracheides—at most five
in a radial row (fig. 6). These secondary elements are little larger than the
smallest primary tracheides—about 25-35 u in diameter. On other parts of
¢
62 DR. D, H. SCOTT ON THE HETERANGIUMS
the circumference no secondary xylem can be distinguished. Thus secondary
growth had just started, the newly formed elements being, as usual, small,
corresponding rather to the centrifugal primary tracheides on which they
abut, than to the wide elements of the metaxylem.
The primary strueture of the peripheral strands was thus, as we should
expect, mesarch, but with the centrifugal portion of the primary xylem very
small in amount (fig. 5).
Deferring the important subject of the leaf-traces, we come next to the
pericycle, for the phloem is practically lost.
The inner margin of the pericyclie zone is more or less crenulated, its
concavities corresponding to the convexities of the xylem cylinder (fig. 7).
Hence they must originally have fitted on to each other, with the inter-
vention of the phloem only. Now they are almost everywhere separated by
a wide gap, containing some Stigmarian rootlets. In places some thickness
of thin-walled tissue is preserved ; its inner portion may include some remains
of the phloem (fig. 6). In other places the pericycle is represented almost
wholly by sclerotic tissue, most of the elements having apparently very thick
walls, like those of the sclerotic cell-plates in the cortex. In the pericycle,
however, the sclerotie tissue is less continuous, a certain number of thin-
walled elements being interspersed. In these more resistant parts of the
pericycle the crenulations are specially well marked, the sclereides extending
into the projections between the hollows (Pl. 8. fig. 7). The pericyclic
sclereides are unusually abundant in Zeterangium shorense, forming exten-
sive tracts, but they occur also in most of our other Coal-Measure forms.
Cortex.
The cortex, as in other Heterangiums, may be divided into inner cortex
and hypoderma—the former, of course, much wider than the latter. The
inner zone consists of a matrix of thin-walled parenchyma, in which great
sclerotic plates are imbedded (Pl. 3. figs. 1 & 7). These plates sometimes
border on the pericycle, and elsewhere may almost reach the hypoderma.
As a rule they consist, as seen in transverse section, of hexagonal cells with
their lumina more or less completely blocked with a brown substance, which
may be altered cell-wall. In other cases the sclereides are flattened and
ranged in regular rows, suggesting the activity of a special meristem. These
appearances are like what we find in other forms, in longitudinal sections
through the sclerotic plates, and are no doubt seen owing to displacement of
certain of these plates.
The great development of the cortical sclerotie masses is characteristic of,
though not peculiar to, the Shore Zeterangium. The thin-walled cortical
tissue between the plates often consists of large cells tangentially dilated ; in
other places the cells are hexagonal and small, resembling those of the
sclerotie plates in form and size.
TU C PT ie h
OF THE BRITISH COAL MEASURES. 63
In the hypoderma the radial bands of sclerenchyma are well marked, and
sometimes anastomose across the intervening tracts of cellular tissue, as in
Lyginopteris. On the prominent leaf-base the proportion of sclerenchyma to
parenchyma is greater than elsewhere (Pl. 8. fig. 2). The leaf-base is delimited
on either side by a barrier of sclerenchyma, resembling that of the hypoderma
on which it abuts (figs. 1, 2). The whole structure of the cortex is typical
of a Heterangium and shows the various tissues in a well-developed form.
The elements commonly described as “secretory sacs” are, however,
obscure. There are a certain number of cells, especially in the cortex, with
dark contents, but, owing to the state of preservation and the absence of any
longitudinal sections, it has not been possible to distinguish with certainty
between differentiated sacs and disorganised cells.
The Leaf-traces.
We now come to the most characteristic point in our plant—the arrange-
ment and course of the leaf-traces. Several traces are present in each
transverse section, but no one section shows them all perfectly. That repre-
sented in fig. 1 may serve to give an idea of their distribution. Four leaf-
traces are shown in the section.
At the top of the figure is the large leaf-base, in which three bundles are
preserved (Trace A). The full number, however, is four, as shown in fig. 2,
which represents the same leaf-base several sections higher up the stem. All
four are likewise shown in a section eut lower down than that represented in
fig. 1, so the presence of three only in the latter is merely due to accidents
of preservation.
The first fact, then, which we learn as to the leaf-traces is that no less than
four bundles entered the base of the leaf.
In the petiole itself still further division took place, as shown in fig. 3,
which represents the section of a detached petiole clearly belonging to the
same plant. Here each of the four original bundles has divided into two, the
division being more advanced in the middle than in the lateral pairs. This
plurality of bundles is a new feature for the genus Heterangium and a marked
difference from the Lower Carboniferous species, M. Grievii, where only a
single bundle enters the petiole.
How, then, do the four bundles entering the leaf-base originate? Turning
to fig. 1, we see in the “ S.E.” of the figure, in the inner cortex, a pair of
bundles close together, and above them another, ill-preserved, pair. In other
sections this second pair is quite well shown ; the space between the two
pairs is about 1*6 mm. There is thus a strong presumption that the two pairs
of strands together constitute a single leaf-trace (Trace Bj. This is con-
firmed on observing the bundles on the left of the figure (Trace C). There
are two on the inner edge of the cortex, and each is evidently in the act of
division. The lower of the two is shown on a larger scale in fig. 6. The
64 DR. D. H. SCOTT ON THE HETERANGIUMS
two strands are about 1:2 mm. apart. The whole is evidently one leaf-trace,
the two constituent bundles just dividing to form the four seen in traces
further out. Lastly, just above the stele in fig. 1, we see two strands
(Traee D) lying loose in the empty space between wood and perieycle. This
tells us little, but in other sections the same pair is found in connection with
the stele. This is shown in fig. 4, from the 3rd section below that from
which fig. 1 was drawn. Here the two bundles are just detaching themselves
from the wood of the stele, with which they are still in continuity. (See
also Pl. 1. Phot. 1, which shows the same trace in an adjacent section.)
We see, then, that in H. shorense two strands left the stele, that they
divided into four in passing through the inner cortex, entered the leaf-base
as a 4-bundle trace, and underwent further division in the petiole. The
middle bundles, in fact, begin to show signs of division before the leaf-base
separates from the stem.
Although the available piece of stem was one and a half inches long, the
changes in this length are but slight, and the comparison of the successive
sections has only shown a fraction of the course of each leaf-trace. Thus, in
the case of the four bundles entering a leaf-base (Trace A, fig. 1), the latter
becomes decidedly more prominent as we follow the series upwards and more
cut off from the cortex by selerenehyma. The bundles, however, are four in
number all through, and, beyond moving outwards and showing some signs
of further division, undergo little change.
Trace B, which is somewhat difficult to follow owing to defects of preser-
vation, is a 4-bundle trace from the bottom of the series ; following the trace
upwards, the two pairs separate widely from each other, and those of each
pair also diverge, but the stage shown in trace A seems not to be reached
within the series.
Trace C first appears as a 2-bundle trace, each bundle showing signs of
division. The division becomes complete as we follow the trace upwards,
but that is about the only change.
The trace D, which is seen starting from the stele in the lower sections,
cannot be followed continuously owing to damage. At the top of the series,
however, what appears to be the same trace appears in the pericycle, each of
the two bundles showing signs of division.
Piecing together the evidence from the various traces, the whole course
becomes clear, as explained above—namely, a double trace leaving the stele
and further dividing into four bundles before entering the leaf-base.
A word may be added as to the free petiole (fig. 2). It runs practically
through the whole series, with little or no change. The transverse section
measures about 12x 5:5 mm. It is concave on one side, convex on the other,
the former having no doubt faced the stem.
TON CNET rS Ts
P TTE j
OF THE BRITISH COAL MEASURES. 65
The structure is the same as that of the leaf-base attached to the stem,
excepting only the further subdivision of the bundles. In tbe two lateral
bundles the xylem is divided, but still in connection. In the middle bundles
the division is complete, the ground-tissue already separating the daughter-
bundles of each pair (fig. 3). The structure of each bundle appears to be the
same as in the xylem-strands of the stem, ¿. e. mesarch with little centrifugal
wood. There is no appreciable change in the arrangement of the bundles
throaghout the series. The ground-tissue of the petiole, like that of the leaf-
base, consists of large-celled parenchyma, in which plates of sclerotic cells,
and also isolated elements of the same kind, are embedded.
The hypoderma is much thicker on the convex than on the concave side.
On the former it is composed chiefly of thick-walled elements, with narrow
radial bands of parenchyma at intervals resembling the hypoderma of the
leaf-base. The thin layer of hypoderma on the concave side of the petiole
seems to consist entirely of sclerenchyma, with smaller cells and thicker walls
than that on the opposite face. Towards one end of the specimen (that
corresponding to the lower end of the stem-fragment) the hypoderma on
the concave side dies out. This is probably merely a matter of preservation,
but it is possible that in this part the petiole or leaf-base was originally
attached to its stem.
The characters of the new species (for as such it must be reckoned in the
absence of evidence to connect it with any other form) may be summed up
as follows :—
Heterangium shorense, sp. nov.
Stem large, reaching a diameter of 18 mm. in the one specimen known.
Primary wood arranged in small definite packets, separated by a reticulum
of parenchyma.
Secondary wood sinall-celled (little developed in the specimen).
Peripheral xylem-strands and leaf-trace bundles mesarch, with little
centrifugal wood.
Selerotic tissue highly developed, both in the pericycle and inner cortex.
Leaf-trace consisting of two bundles where it leaves the stele, dividing into
four on entering the cortex, and further dividing into eight in the leaf-base
and petiole. The traces of at least four leaves present in each transverse
section of the stem. Leaf-traces traversing the stem for a long distance
before passing out.
Locality. Shore, Littleborough, Lancashire: Lower Coal Measures.
Found by Messrs. James and Joseph Lomax, August 1912.
MOS Vu. ee RAE S I E. LEE, gn
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zT QVE UON SIS ee Ca M n q TX MARATA ETTR
DR. D. H. SCOTT ON THE HETERANGIUMS
HETERANGIUM TILLEOIDES, Will.
The anatomy of the stem of this well-known species from the Halifax Hard
Bed has been fully described (Williamson, 1887; Williamson & Scott, 1895)
and need not be recapitulated here. The species is remarkable for the great
development of the phloem, sometimes equalling that of the secondary wood,
and for the characteristic form of the main phloem-rays, dilated outwards
like those of the Lime Tree. The specimens are among the most beautifully
preserved of petrified plants, so far at least as the stelar tissues are
concerned.
Only three points demand our attention—the course of the leaf-traces, the
adventitious roots, and the specific distinction from other members of the
genus.
The Leaf-traces.
No satisfactory account of the course of the leaf-traces has ever been given.
Williamson, in his original description of the species, mentions and figures
“ascending vascular bundles grouped in pairs” (Williamson, 1887, p. 290),
and further speaks of “the twin bundles going off to what, both in this plant
and the allied Aeterangium Grievii, I presume to have been petiolate leaves iil
(L. e. p. 293).
In our joint memoir (Williamson & Scott, 1895, p. 763) it was recognised
that the twin bundles with secondary wood just leaving the stele, already
figured by Williamson in 1887, represent a double leaf-trace. Thus the
trace was shown to start as a pair of distinct bundles. On the other hand,
the conjecture that “ the twin bundles may have completely fused on entering
the leat” has proved to be incorrect. The reverse is the case.
Beyond the fact that the leaf-traces were double, nothing definite was, in
fact, established as to this part of the anatomy of Æ. tiliwoides.
Thanks to the kind help of Prof. F. W. Oliver and Prof. F. E. Weiss in
lending slides from their collections, I have been able to examine most, if
not all, of the sections of this species in this country. Good as the sections
are for anatomical details, the material is not particularly favourable for
following the course of theleaf-traces. This is owing partly to the absence
of continuous series of sufficient length and partly to the fact that in this
species the cortex is usually the worst-preserved region of the stem, so that
the leaí-traces are imperfectly shown in the outer part of their course. Still,
it has been possible to determine the essential points.
The sections of Æ. tilimoides which have come under observation appear to
belong to four distinet stems, reckoning only those which clearly belong
to this species. Three of these are represented in the Williamson Collection,
and sections from them are described and figured in his original memoir of
1887. These may be regarded as types. The fourth specimen is represented
in my own collection and that of the Manchester Museum. A synopsis of
OF THE BRITISH COAL MEASURES. 67
all the sections known to me is given in a footnote *. The four stems will
be shortly referred to as
Specimen 1 (Williamson, slide 1301 ete.).
Specimen 2 (Williamson, slide 1302 ete.).
Specimen 3 (Williamson, slide 1620 eto.).
Specimen 4 (Scott, slide 1323 ete.).
Specimen 3 is the only one which affords a fairly long series of trans-
verse sections. I have succeeded in determining the order of the 10 sections
(from various collections) and find it to be as follows, from below upwards :—
Q. 60; R. 658; Q. 59; N. 12; W. 1620; W. 1624; W. 1623; W. 1625;
Q. 58; R. 634.
Two leaf-traces are shown ; the divergence between them would be con-
sistent with a 2 phyllotaxis. At the bottom of the series the inner trace (T. 1)
is scarcely beginning to leave the stele; it consists of two distinct strands
which, however, are here almost in contact laterally (P1. 8. fig. 8, from the
4th section). As they pass out through the pericycle they separate rapidly;
and gradually lose their own secondary tissues, while those of the stele close
in behind them. At the top of the series (the ninth section, Q. 58, is the last
which shows this trace complete) the bundles are 2 mm. apart and are in the
middle of the pericycle, in which they cause two marked projections. They
show, however, no sign as yet of further division.
At the bottom of the series the outer trace (T. 2) is about as advanced as
the inner trace was in the uppermost section. In the 2nd section the two
* [n the following list the letter W. indieates the Williamson Collection in the British
Museum (Natural History); the letters Q. & R. the collections of the Manchester Museum,
Q. being the Cash, R. the Hick Collection; S. indicates my own collection; and N. that of
the Botanical Department of University College, London (Heterangium slides).
` Specimen 1.—2 transverse sections, W. 1301; R. 632.
Specimen 2.—6 transverse and 7 longitudinal sections. Transverse: W. 1302, 1303, 1619;
Q. 55, R. 654; S. 237. Longitudinal: W. 1304, 1621, 1622, 1627, 1628; Q. 56,
R. 631 (?). All these sections appear to be from the same stem, a point not
recognised in the joint memoir (Williamson and Scott, 1895, p. 778).
Specimen 3.—10 transverse sections: W. 1620, 1623, 1624, 1625; Q. 58, 59, 60; R. 634,
658; N. 12.
Specimen 4.—9 transverse and 2 longitudinal sections. Transverse: R. 633; S. 1323.
Longitudinal: S. 238, 1324. ;
The order of the transverse sections, so far as it could be determined, is given in
the text.
The above list is limited to stems from Halifax. There is a Dulesgate specimen (slides
S. 625 & R. 639) which bears some resemblance to H. tiliæoides, but seems to be really
referable to H. Lomavii (see pp. 77, 79, 82).
A small stem from Halifax, collected by the late Mr. Spencer (sections S. 239, S. 1322;
N.2 a, N. 2 b) may be a young specimen of the same species.
ase A SEN
68 DR. D. H. SCOTT ON THE HETERANGIUMS
bundles are quite in the outer part of the pericycle, each occupying a very
marked protrusion of this zone. They are about 1°3 mm. apart, and each
bundle is tangentially widened and slightły concave on the outer surface—
indications of approaching division. The bundles are shown in fig. 8 from
the 4th section (N. 12) where the pericyclic bulges are very prominent, and
the preparations for division of the bundles evident. Two sections higher up
the bundles have left the pericycle altogether, and are now in the ill-preserved
cortex. The further changes consist chiefly in the progress of division in
each bundle. The trace is shown in fig. 9 from the 9th section (Q. 58),
where each bundle is distinetly double *.
This series shows, then, that the trace leaves the stele as a pair of bundles,
which become widely separated as they pass outwards, and that each of these
divides into two in the cortex. But the complete division of the two bundles
into four is not shown.
This series, however, is of additional interest from the fact that three
of the sections contain a detached petiole (Pl. 4. fig. 10). The preservation
is imperfect but the structure is fairly shown. On the outside is a rather
narrow hypoderma of the * Sparganum" type, with the sclerenchyma exceed-
ing the parenchymatous bands in extent. The large-celled ground-tissue
contains two sclerotic patches, in one of which the cells are ranged in rows.
There are also some scattered elements with contents, which resemble the
so-called secretory sacs of the stem.
The petiole measures about 6 x 2:5 mm., and has one side flat, the other
convex. The important point is that it contains four distinct bundles (shown
in all the sections). They have been displaced by the intrusion of Stigmarian
rootlets, but appear to have been in two pairs, those of one pair further apart
than those of the other. The xylem of the bundles consists of a mass of
large tracheides, with a band of smaller elements on one side—exactly like
the strands in the petiole of H. shorense (cf. fig. 3). The organ is obviously
the petiole of a //eterangium, and there is no reason to doubt that it belongs
to the stem with which it is associated. Its dimensions relative to the stem
are about in proportion to those in the Shore plant (H. shorense, stem
18 mm., petiole 12 x 5'5 mm. ; H. tilieoides, specimen 3, stem, if complete,
about 10 mm., petiole 6 x 2:5 mm.).
This specimen, then, establishes the strongest presumption that the petiole
of FI. tiliwoides contained four distinct vascular bundles, confirming the
evidence for a quadruple leaf-trace derived from the stem.
To complete the proof it is clearly desirable to observe the structure
of a leaf-base still in connection with the stem. This is shown, though
* One of the double bundles is figured by Williamson (1887, pl. 22. fig. 7) from W. 1623,
the 7th section.
OF THE BRITISH COAL MEASURES. 69
imperfectly, in a section (S. 237) of specimen 2 * (PI. 1. Phot. 3). This is a
large stem, quite 15 mm. in diameter when complete. Though the internal
tissues are so admirably preserved, the cortex, as usual, is in a fragmentary
state.
A pair of bundles is very beautifully shown just separating from the stele
(Pl. 1. Phot. 3, T. 1; cf. Phot. 5). Far out in the pericycle is another pair
(T. 2) very widely separated, the interval between the strands being nearly
4 mm. These bundles show some signs of division. What chiefly interests
us is the presence of an evident, though badly preserved leaf-base, pro-
jecting far out from the stem and beginning to be marked off at the sides
by internal bands of sclerenchyma (Phots. 3 & 4). The whole is much
damaged, but in the middle part the continuity of the tissues is preserved.
The tangential width of the leaf-base is about 7:5 mm. Towards one side
of the leaf-base and in its inner part is a distinct pair of bundles ; the two
strands are about 200 p apart and separated by ground-tissue. In a corre-
sponding position on the opposite side are the remains of another pair ; one
of these strands is fairly preserved, the other partly destroyed (Phot. 4, v.b.).
The leaf-base, so far as preserved, shows the usual Heterangium structure,
with a rather narrow hypoderma and large plates of sclereides in the ground-
tissue. This specimen proves that four bundles, in two pairs, entered the
leaf-base, and thus confirms the evidence of the detached petiole associated
with specimen 3.
It may be mentioned that in the section W. 1302, which appears to have been
cut just below S. 237, two pairs of bundles are shown in the cortex which are
probably the same as those which pass out into the leaf-base just described.
Both pairs are shown in Williamson's fig. 1 (1887) (one pair is marked v’, u’;
the other, not specially lettered, lies on and below the line p). The bundles
shown in Williamson's fig. 8, on a larger scale, appear to be the former pair.
The main points in the course of the leaf-traces are now clear, but it may
be well to summarise the evidence from the remaining sections.
The transverse sections of specimen 2 are not in a series, but appear to run
in pairsf. The sections R. 654 and W. 1303 come next each other, the
former being below the latter. In R. 654 two traces are shown ; the inner
is the pair just leaving the stele, shown in Phot. 5. The phase is almost the
same as that of the corresponding pair in S. 237 (cf. Phot. 3) ; a betier
example could not be found. The two bundles of the trace are quite distinct
as regards their primary xylem, but close together ; they are separated from
* This is the magnificent specimen from sections of which most of the figures of
H. tilieoides in Williamson's memoir of 1887 were drawn.
S. 237 was cut for me by Mr. F. Chapman from the original block, next to W. 1302
shown in Williamson's fig. 1 (1887).
+ W. 1302 and S. 237 have already been dealt with.
Pee Sid
70 DR. D. H. SCOTT ON THE HETERANGIUMS
the metaxylem of the stele by a few layers of conjunctive parenchyma, in
which some tangential divisions have taken place. The secondary wood
proper to the double leaf-trace is still continuous, but the ares corresponding
to the two bundles are already evident. Most of the phloem is perfectly
preserved ; it about equals the secondary xylem in thickness.
In the outer part of the pericycle there is a second trace; the two strands
are wide apart (about 4 mm.) and each is preparing to divide. In the next
section (W. 1303) the inner trace has moved out into the pericycle ; the two
strands are now diverging, each has its own are of secondary wood and bast.
These bundles were admirably figured by Williamson (fig. 5, 1887) at a time
when their leaf-trace nature had not yet been recognised. The outer trace
is here in the cortex, but only fragments are preserved.
The two sections W. 1619 and Q. 55 are similar and must have been eut
near together, but I cannot exactly correlate them. In W. 1619 no leaf-
trace is starting from the stele—the appendage shown is no doubt an adven-
titious root (figured by Williamson, fig. 13, 1887). "Two double bundles are
seen in the cortex, just beyond the pericycle. They are about 6 mm. apart,
but may have been displaced ; they probably belong to the same trace.
Q. 55 shows two leaf-traces. The inner is in the perieycle—one strand has
been displaced. These bundles have almost lost their ares of secondary
wood, and are therefore at a more advanced phase than those deseribed in
W. 1303. The other trace is represented by a double bundle in the cortex
the eompanion pair is no doubt lost, for much of the cortex is destroyed,
Of specimen 1, I only know of two sections, one in the Williamson and
one in the Manchester Collection (W. 1301 and R. 632). Both are trans-
verse and cut close together, R. 632 being just above W. 1301. The specimen
is exceptional as regards preservation, for the cortex is fairly preserved,
while the pericycle is almost destroyed. W. 1301 shows three leaf-traces
and a leaf-base. The innermost (Trace 1) consists of a pair of strands,
separated by some conjunctive tissue, about to depart from the stele. A pair
of widely diverging bundles (Trace 2) is seen in the pericycle, and a damaged
one (Trace 3) in the cortex. The leaf-base or petiole in connection with the
stem was described in the joint memoir by Williamson and myself (1895,
p. 763). Only one bundle is shown, but, as the section of the petiole is
longitudinal, this proves nothing as to the total number present. The
conjecture as to fusion of bundles was certainly wrong, as shown by the
evidence from specimens already described (p. 68).
The other section, R. 632 (Phot. 2), shows the same leaf-traves. Trace 1
has searcely moved. The two strands, though only just separating from the
wood of the stele, are about 370 p apart, showing clearly that the bundles are
distinct at their origin. Of trace 2 only one bundle is preserved ; it lies on
the border of pericycle and cortex, and is dividing. Trace 3 is in the inner
OF THE BRITISH COAL MEASURES. 71
cortex ; all four bundles are shown, forming two pairs, about 2:5 mm. apart.
The strands of each pair are quite separate, with about 200 p of cortical
tissue between them. The section is important, as proving the complete
division of the leaf-trace into four while in the cortex ; unfortunately one of
the pairs of bundles is damaged (see Phot. 2). The attached leaf-base is
still shown, but no bundle is visible. The three traces and leaf-base mark
the position of four successive leaves: their arrangement is in agreement
with a 2 phyllotaxis.
There are two transverse sections of specimen 4 (S. 1323 and R. 633) ;
they were cut near together. It is a beautiful specimen of a young stem,
showing all the stelar tissues perfectly (Pl. 1. Phot. 8, & Pl. 4. fig. 12), but,
owing to the loss of the cortex, is notof much value for the course of the leaf-
traces. In the not quite complete sections two traces are shown, both in the
pericycle. In the lower section (S. 1323) the inner trace (T. 1) lies in
the inner part of the wide pericycle, and consists of two bundles nearly 1 mm.
apart. They have no secondary wood at this level, but it is not much
developed on the stele itself. Behind them the stelar wood is beginning
to close up in the form of small tracheides, which for the most part run
horizontally. The outer trace (T. 2), at about 2 divergence from the former,
lies in the extreme outside of the pericycle, which forms a great protrusion
round each of the two bundles; they are over 2 mm. apart (Phot. 8). In
the upper section (R. 633) the strands of the inner trace are about 1:5 mm.
apart, but there is otherwise little change.
Taking the whole of the evidence from the four specimens, we arrive
at the following conclusions regarding the course of the leaf-traces in
H. tiliwoides.
Two distinct strands start from the stele to form a leaf-trace. Each has
at first its own secondary tissues, which it loses in traversing the pericycle.
The strands diverge rapidly, and before the pericycle is passed each begins
to divide. The division is completed in the cortex, and four bundles enter
the leaf-base and petiole. It will be noticed that the facts are essentially
the same as in H. shorense, except that in H. tiliwoides no further subdivision
of the four bundles in the petiole has been observed. This may merely be
correlated with the smaller size of the specimen. The phyllotaxis appears
to be 2.
Adventitious Roots.
The evidence as to the roots of Heterangium is meagre*. It is therefore
of interest to find that in H. tiliwoides the structure of the root can be deter-
mined with a near approach to certainty, a point not hitherto noticed. The
best evidence is provided by specimen 8. At the bottom of the series
(Phot. 7; section Q. 60) the stem bears an appendage (7.) embedded in the
* See Williamson & Scott, 1895, pp. 764, 765; Scott, 1909, p. 410.
72 DR. D. H. SCOTT ON THE HETERANGIUMS
cortex, which from its structure is evidently an adventitious root. In the
2nd section a similar root (Pl. 1. Phot. 6, 7.2.) is shown still connected with
the wood. Immediately to the outside is a free root cut transversely (7.).
Tt is tetrarch, with a considerable amount of secondary wood, and most of the
cortex is preserved. This root reappears in the next two sections, but its con-
nection with the stem is not shown. It agrees, however, as nearly as one can
expect, with the root-base attached to the stem, and there is little doubt that
the free root also belongs to the latter. The drawing (Pl. 4. fig. 11) is taken
from the 4th section (N. 12). Here also the root seems to be tetrarch, the
four protoxylem-groups corresponding to four principal rays. The secondary
wood is well developed. Where it is cut somewhat obliquely the tracheides
show the multiseriate pits on their radial walls as in the stem. The cortex
contains a certain number of cells with dark or brown contents. So far as
can be observed, the structure of the root is of the * Aaloaylon” type. The
characteristic double peripheral layer is not shown, but may have been lost.
The connection with the stem of H. tilivoides, though not strictly proved, is,
as we have seen, beyond measurable doubt.
A very clear root-base in connection with another specimen was figured
by Williamson (1887, pl. 23. fig. 12). The appendage shown in his fig. 13
was no doubt of the same nature.
Specific characters.
Nothing is more difficult than to fix the value of anatomical characters in
discriminating species, especially in the case of petrified specimens, where
practically no other characters are available,
Fleterangium tiliwoides is most probably distinct from the other forms
described in this paper, but an anatomical diagnosis is not easy to draw up.
A few characteristic points may be recalled,
The primary wood has the tracheides grouped in packets, not unlike those
of H. shorense, but on the whole less regular (Phots. 2, 3, 6, & 7).
The pits on the tracheides both of the metaxylem and secondary wood
commonly have a marked spiral arrangement, often with definite spiral bands -
between the rows of pits (Williamson & Scott, 1895, p. 764).
The great development of the phloem and the presence of large dilated
phloem-rays are perhaps the chief characters, but are of course only recog-
nizable in the more advanced specimens *. The phloem may equal or exceed
the secondary wood in thickness. Thus, in specimen 4 (a young stem) the
secondary wood is about 160 w thick and the phloem about 200 p at the
same place; at another point they are just equal (245 u). In an older
stem (specimen 2) the secondary wood ranges from about 570 u to 900 p
in thickness, the phloem from about 650 p to 740 p.
* For example, they do not help us in determining whether ZZ. shorense (a young specimen)
is really distinct.
OF THE BRITISH COAL MEASURES. 73
The pericycle is very wide, 820 u to 900 u, in the young specimen 4
(Phot. 7), and 1:1 to 2'2 mm. in the more mature specimen 2 (Phot. 3).
The cells are very frequently found in a state of division, especially in the
inner layers, so no doubt this zone grew with the growth of the stem. The
pericyclic selereides are chiefly found in the inner portion ; their extent
varies much in the different specimens.
The fact that the pericycle is usually better preserved than the cortex
(almost wholly lost, for example, in specimen 4, otherwise so perfectly
preserved) might be thought to indicate some structural peculiarity. But
specimen 1, as we have seen, is an exception, and, where the cortex is found;
it has the typical Heterangium structure, with plates of sclereides in the
ground-tissue and a “ Sparganum ? hypoderma, but not very strongly
developed.
The course of the leaf-traces has already been dealt with; while, differing
conspicuously from the monodesmie type represented by the Lower Car-
boniferous M. Grievii, it agrees essentially with that of other Coal-Measure
forms. The rapidity with which the leaf-traces were given off varies with
different specimens. Thus, in specimen 3 there are never more than two
traces shown in the transverse section, while in specimen 1 there are as many
as four, if we include the leaf-base attached to the stem. It may be worth
noting that in /7. tilivoides the two strands of the trace, where they leave the
stelar wood, are near together, though distinet; in some other Coal-Measure
forms they are much wider apart to start with ; this, however, affords no
certain specific distinction.
It is not perfectly certain that the stem described as H. shorense may not
be a young specimen of H. tiliwoides, but there are several small distinctions,
such as the more regular grouping of the metaxylem tracheides in H. shorense,
the apparently narrower pericycle with sclereides at the periphery, and the
division of the leaf-trace into eight in the petiole. As the localities are
different, H. tilivoides coming from the Halifax Beds and H. shorense from
Dulesgate, there is no presumption that they are identical, and it is better to
keep the two distinct. The following diagnosis may be given :—
Fleterangium tiliwoides, Will.
Stem from about 10 mm. to about 16 mm. in diameter.
Primary wood arranged in more or less definite packets, separated by
parenchyma.
Tracheides often with a spiral band between the pits.
Secondary phloem much developed, often equalling or exceeding the
secondary wood in thickness. Phloem-rays much dilated.
Peripheral xylem-strands and leaf-trace bundles mesarch, with little
centrifugal wood. — ,
Pericycle broad. Sclerotic tissue present both in perieyele and cortex.
LINN. JOURN.—BOTANY, VOL. XLIV. G
pt A
74 DR. D. H. SCOTT ON THE HETERANGIUMS
Leaf-trace consisting of two bundles where it leaves the stele, dividing
into four in the cortex. Four bundles in the petiole. Number of traces in
the transverse section from two to four. Traces traversing the stem for a
long distance before passing out.
Locality. Halifax Hard Bed, Lower Coal Measures.
Hereraneium Lomaxin, Will. in MS.
Williamson, in his 17th Memoir (1890, p. 96), states, under the heading
Heterangium Grievii, that Mr. Lomax had found specimens of the same plant
from Dulesgate in Lancashire. He proceeds: * But along with these he
brought from the same locality a series of sections of what at first seemed to
be a different species of Heterangium. But long and careful comparison of
all the examples of that genus in my cabinet convinced me that the supposed
new forms were merely the H. Grievii in a younger state of growth.”
The specimens figured and shortly described in the memoir referred to
are those in the “younger state of growth” (Williamson, l. e. pls. 14, 15.
figs. 14-18), whieh Williamson at first regarded as distinct. He did not
deseribe the other, more advanced form, which he had always considered as
simply H. Grierü. In the joint memoir (Williamson & Scott, 1895) the
Dulesgate forms are only referred to incidentally, but both the younger and
more advanced conditions are illustrated for special points (l. c. pl. 26. fig. 21;
pl. 27. figs. 28, 29). It was no doubt to the younger and more peculiar
form that Williamson's unpublished MS. name of H. Lomaati was intended
to apply. In the joint memoir we also described and figured “a Heterangium
of uncertain species," from Dulesgate, and provisionally named it H. cylin-
dricum (l. c. p. 164, pl. 28. figs. 30, 31).
There are therefore three forms of the Dulesgate Heterangiums which
have already been recorded.
1. The young stems with little or no secondary thickening. This form
will be called « (Pl. 4. figs. 16, 17).
2. The older stems, with well-marked secondary tissues, resembling
H. Grievii, now to be called the 8 form (Pl. 2. Phots. 9, 11, 12, 13).
3. The so-called //eterangium cylindricum™ (Pl. 2. Phot. 14).
After careful comparison of all available sections I have come to the con-
clusion that all the above, including one or two minor variants, belong to one
species (of course, quite distinct from the Lower Carboniferous H. G'rievii),
to which the name //. Lomazii is collectively applied. The only Dulesgate
* All the specimens of these three forms appear to have come from one coal-ball. This is
Mr. Lomax's recollection, and it is confirmed by the uniform character of the matrix in which
the stems are imbedded. The single specimen of Heterangium minimum (see below) came
from a different block. It is remarkable that no new specimens of H. Lom«xii have been
met with since the original discovery by Mr. Lomax about 1890. The species must have
been very rare or the conditions for its preservation very unfavourable.
OF THE BRITISH COAL MEASURES. 75
Heterangium which it is proposed to keep distinct is the little stem described
in a subsequent section of the paper under the name /7. minimum (p. 90).
The reasons for the conclusions arrived at will appear in the course of the
description. I take the 8 form first, because the best specimens belong to
it, and it has never been properly described. Its relation to the æ form is
not very obvious, though I think Williamson was right in uniting them.
H. Lomazii, Form £.
The specimens of this form are associated with those of the « form and of
** H. eylindricum." All appear to have come originally from the same block.
I have enumerated in the footnote * seven specimens of the 8 form, though
more than one may possibly belong to the same stem, as in the case of 5 and 6.
The preservation of the outer tissues is seldom so complete as to render
any very accurate measurements of the stem practicable. Roughly speaking,
the dimensions of the different specimens range from 17x10 mm. to
7x4 mm., or, allowing for compression and distortion, the diameter may be
said to vary from about 13 mm. to about 5 or 6 mm.
The general form of the stem, to judge from the least-distorted specimens,
seems to have been nearly cylindrical. The leaves appear to have been, as a
rule, borne on the stem at long intervals, for many of the sections show
the trace of only one leaf, and unfortunately no section happens to pass
through an evident leaf-base in connection with the stem. The best-preserved
specimen is No. 5 (Pl. 2. Phot. 13) ; from the considerable development of the
phloem, this stem at first sight recalls H. tilivoides, but from its structure
must be referred to H. Lomawii.
The Stele.
The more or less compressed primary wood varies in the different specimens
from 8 x 2 mm. to 3 x 1 mm.—say, from 4 or 5 mm. to about 2 mm. in diameter
if restored to its presumably cylindrical natural form. Its size is by no
means always proportional to that of the stem, owing to large differences
in the development of the secondary tissues.
The arrangement of the metaxylem tracheides is not very regular (PI. 2.
Phots. 9, 12), the groups being less definite in form and size than those of
* For reference letters see above, p. 67.
Specimen 1.—7 transverse sections : W. 1915, 1915 A, 1915 B, 1915 C; Q. 6, Q. 11; S. 447.
Specimen 2.—8 transverse sections: W., 1915 H ; Q. 64; S. 445.
Specimen 3.—' transverse sections: W. 1304 A, 1915 P; R. 621; S. 444, 446 ; N. 7, 19.
The longitudinal sections W. 1915 G and 1915 R probably belong to this specimen; possibly
R. 638 also.
Specimen 4.—9 transverse sections: W. 1915 R; R. 645.
Specimen 5.—2 trausverse sections: R. 639; S. 625.
Specimen 6.—3 transverse sections : W. 1915 D, 1915 E, and probably N. 3.
Specimen 1.—9 incomplete transverse sections : IBI0b7; N, 8.
Where necessary, the order of the sections in a series is given in the text.
G2
1
4
x
‘
|
;
—OGUURNRSTYT IO WT i ee "WP TT WT T
T6 DR. D. H. SCOTT ON THE HETERANGIUMS
HĒ. shorense or even H. tiliwoides. Sometimes (as in specimen 5, Phor. 13)
the amount of xylem-parenchyma is considerable. The peripheral xylem-
groups are very distinct, the radial bands of parenchyma between them being
continuous with the principal rays of the secondary wood (Pl. 2. Phots. 9, 12;
Pl.4.fig.13). In this point the present species agrees with H. tiliwou/es and
H. shorense *, and differs from the Lower Carboniferous M. Grievii, in which
the peripheral strands are less isolated and the secondary wood not divided
up by principal rays.
Another feature of H. Lomaxii, which it shares with other Coal-Measure
forms, is the position of the protoxylem, which lies very near the outside of
the primary xylem, with only one or two centrifugal elements before the
secondary wood is reached. "Thus the peripheral xylem-strands, though still
mesarch, are approaching an exarch structure (see Phot. 10). This, again,
is a marked difference from H. Grievii, where the centrifugal part of the
primary wood is considerably more developed (see Williamson & Scott,
1895, pl. 26. figs. 24, 25 t). The position of the protoxylem in 77. Lomari is
well shown in longitudinal section in PI. 4, fig. 14, where the laxly coiled spiral
element, which is no doubt the true protoxylem, is sueceeded to the outside
by a single, more closely wound spiral tracheide, beyond which the wood
appears to be secondary. On the inside the pitted elements are reached at
once. Often, however, there is a little parenchyma on the inner side of the
protoxylem. It appears that here, as in Lyyinopteris and Heterangium
Grievii, the centrifugal part of the xylem was the first to be formed after the
protoxylem.
Elements with dark brown contents are very frequent in the xylem-paren-
chyma, the rays, and throughout the stele. Whether they can be properly
called “secretory sacs” is doubtful. They do not show, as a rule, any special
elongation as compared with the associated cells. They are best shown in
the rays, where they run in a radial direction ; here their contents are some-
times in the form of a mass of granules, simulating altered starch-grains,
but very variable in size. The * secretory " elements were mentioned in the
joint memoir as characteristic of the Dulesgate specimens 1.
The secondary wood varies greatly in amount, not only in the different
specimens, but in different parts of the same specimen : it sometimes attains
a very considerable development. Thus, in specimen 5, the thickness of the
secondary wood ranges from 0°61 to 1:64 mm. (P1. 2. Phot. 13). In specimen 1
the range is even greater, varying in one transverse section (S. 447 ; Phot. 9)
from 0:3 to 2:58 mm. The excessive thickness at certain places is correlated
with the presence of adventitious roots, as was already remarked in the joint
* In the latter, the specimen is too young to show the relation to the principal rays.
+ This distinction was noted iu the joint memoir, l. c. p. 750.
1 Williamson & Scott, 1895, pp. 748, 792.
OF THE BRITISH COAL MEASURES. T
memoir of 1895*. In other specimens the secondary growth has made little
progress. Thus in a section (W. 1915 P) referred to specimen 3, the thick-
ness ranges from *08 to *3 mm. only, and in specimen 2 it is about the same
(see Pl. 4. fig. 13). These relatively voung specimens are of interest for com-
parison with the unthickened stems included under our form «æ (see page 85).
As is usual in //eterangium, the first-formed elements of the secondary
wood are considerably smaller than the primary metaxylem tracheides ;
further out, if the growth is continued, they generally become larger,
though seldom reaching the size of the largest primary tracheides (see Pl. 2.
Phots. 9 and 13). Local tangential bands of exceptionally narrow tracheides
are frequent.
The structure of the secondary wood is like that of other species of
Heterangium. The tracheides have the usual multiseriate bordered pits on
their radial walls. Where, however, the elements are very narrow, the
number of rows may be reduced to two or even, locally, to one. Where the
pits are well preserved, we see that the border has a narrow inclined slit.
In places the pits show traces of a spiral arrangement, but the definite
spiral bands noticed in H. tiliwoides have not been observed here.
As I have seen no good tangential section it is difficult to say anything
definite about the structure of the tangential walls. Certainly the pits are
very few here compared with those on the radial walls, but there is evidence
for the occurrence of scattered tangential pits f.
The principal rays—those corresponding to the intervals between the
primary xylem-strands—are of considerable width, amounting to 9 or 10
cells in the larger specimens. The secondary rays vary much in width and
in some cases appear to be uniseriate. The principal rays are occasionally
divided up by intercalated tracheides, at some distance out, and, conversely,
new rays appear as the wood increases in thickness. The ray-cells are thin-
walled and much elongated in the radial direction.
'The phloem is fairly preserved in one case only, specimen 5 (see Phot. 13).
It here attains a maximum thickness of about 0*8 mm. I have not found such
marked dilation of the phloem-rays as characterizes H. tiliwoides. The
secondary phloem consists for the most part of fairly large, radially arranged
elements. A small-celled irregular tissue sometimes preserved on the outer
border is no doubt the primary phloem. Ii the absence of satisfactory
longitudinal sections no details of the phloem-structure can be given.
The pericycle forms a broad zone, and is often better preserved than the
cortex, as in H. tiliwoides (Phots. 9, 12, 13). Its normal width, apart from
bulges where bundles or roots pass out, is about 0'6 mm. in specimens 1
and 6, where it is best shown.
* Williamson & Scott, 1895, p. 757.
+ Cf. Williamson & Scott, 1895, p. 752.
78 DR. D. H. SCOTT ON THE HETERANGIUMS
Sclerotic groups are frequent in the perieycle: in most stems they are
quite conspicuous ; in specimens 5 and 6 they are not so easy to find, but
this seems to be only because their cell-walls are less thickened. The
sclereides are short elements, but (to judge from the longitudinal section
R. 638) appear to form vertical strands of considerable length. The pericyclic
parenchyma consists of short cells, many of which have the dark contents
above referred to.
On the outer border of the perieyele tangential divisions have sometimes
taken place, giving rise to a tissue resembling an internal periderm, such as
we find in Lyginopteris *. This is best shown in specimen 1 (e. g., S. 447).
The cortex is well preserved in certain cases, though often the parenchy-
matous part of the inner zone has perished, with the result that the
Sparganum hypoderma is sometimes stripped off. When the parenchyma is
preserved it consists of large, short, thin-walled cells. Here, as in other
species, the sclerotic plates of the inner cortex are a conspicuous feature.
Their transverse diameter may be as much as 1°4 mm. In the descriptions
previously given they are confused with the corresponding structures in the
true H. Grievii t ; in reality those of H. Lomazii are quite distinct. They
here form massive blocks, rather than plates, reaching a thickness of about
15 cells and 0*6 mm. In the middle part the cells are flat and thick-walled,
while towards the upper and lower edges they become longer, with thinner
walls. All the cells are regularly ranged in vertical rows (see Pl. 4. fig. 15).
Corresponding to their bulky form, the sclerotic masses are rather widely
spaced. In H. Grievii the plates are much thinner and seldom more than
four cells or about 0'2 mm. in thickness ; they are also more numerous. The
contrast is striking if one compares fig. 15 in this paper with Williamson’s
figure of H. Grievii (Williamson, 1873, pl. 29. fig. 32). H. tiliwoides is
intermediate in these respects (Williamson, 1887, pl. 22. fig. 11). As regards
the sclereides themselves, the apparent thickness of their cell-walls varies
much. In some cases it is probable that the walls were not really as thin
as they appear, the thickening layers originally present having become altered
and obscured.
The parenchyma of the inner cortex consists of rather large, short or
slightly elongated, thin-walled cells. The Sparganum hypoderma 1, or outer
cortex, varies a good deal in the different specimens. As a rule, we find the
usual alternating radial bands of sclerenchyma and parenchyma ; the former
are commonly broad and not much elongated radially, and thus quite
different from those of H. Grievit §. The bands sometimes anastomose. In
* Williamson & Scott, 1895, p. 718, pl. 21. fig. 1; pl. 22. figs. 5-7.
T Asin Williamson & Scott, l. c. p. 753.
t The name“hypoderma,” though commonly used, is not strictly correct, for the Sparganum
zone does not border immediately on the epidermis.
$ Cf. Williamson, 1873, pl. 28. fig. 30; pl. 29. fig. 35.
OF THE BRITISH COAL MEASURES. 19
ihe younger stems the parenchymatous strips are narrow; as the stem
increased in thickness they widened out somewhat (Phot. 9). In one stem
(specimen 5; P1.2. Phot. 13) the Sparganum structure is absent ; all the tissue
of the outer cortex, so far as preserved, is thin-walled, and there is little sign
of any differentiation. The stem is in an advanced stage of growth, so this
peculiarity cannot be explained by youth; possibly this specimen may have
been a rhizeme or underground portion of the stem.
Beyond the Sparganum zone, where it is normally developed, we find two
or three layers of cells larger than the sclerenchymatous fibres; they may
have either thin or thick walls—the latter especially where they abut on
sclerenchyma. In longitudinal section they are found to be elongated, but
differ from the fibres in having transverse end-walls ; their contents are often
carbonaceous.
Lastly, we come to the epidermis, consisting of rather small and thick-
walled cells, often remarkably well preserved. I have not, however, succeeded
in detecting any stomata.
We have now completed the description of the stem-structure, apart from
the leaf-traces. At every point there are differences from the Lower
Carboniferous H. Grievii, and when once the comparison is made no con-
fusion between the species is possible. The relations of Æ. Lomaaii to the
contemporary H. tiliwoides are much closer.
The Leaj-traces.
The distribution of the vascular bundles supplying the leaves is perhaps
the most important point in the anatomy, and has not previously been
investigated in H. Loma. The different specimens of the 8 form show
some variations in this respect. We will begin with specimen 1, of which
there is a series of 7 transverse sections; the order from below upwards
appears to be as follows :
B 447: 0.6; W.19150; Q 315 W. 1915; W.1915 B; W. 1915 A.
There seems to be a gap between W. 1915 and W. 1915 D, but I believe
that the same leaf-trace is present throughout the series.
In the lowest section (8.447; P1.2. Phot. 9) there are two large bundles in the
pericycle, 2:5 mm. apart *; each bundle causes a marked bulge in the pericycle ;
on the inner side only one or two layers of cells invervene between the bundle
and the ill-preserved phloem of the stele. No other leaf-trace is present,
and the two strands are both in the same phase ; it therefore seems clear that
both must belong to one and the same leaf-trace, for if they were two separate
traces one would be further out than the other, and additional leaf-traces
would be shown in other parts of the periphery. It might, indeed, be imagined
that we have to do with a dorsiventral axis, the leaf-traces being given off
from one side only as, e. g., in Helminthostachys ; in that case it is very unlikely
* The clear interval is measured from xylem to xylem.
EE ie, ee E, -..
de al
80 DR. D. H. SCOTT ON THE HETERANGIUMS
that leaves and roots would be borne on the same side of the stem, and, as we
shall see presently, adventitious roots come off in our specimen between the
two leaf-trace bundles.
It thus appears that the leaf-trace was a double one, with the two strands
very far apart.
The bundles are unusually large, the xylem of each measuring at least
1*4 by 0-8 mim. ; each has a coneavity on the outer side and is intersected
radially by a band of narrow elements (parenchyma ?), suggesting an
approaching division, which, however, is never carried out within the series.
In this section the wood is much thickest on the side opposite the bundles ;
this is no doubt due to a previous trace, with its attendant roots, having been
given off in that position at a lower level. The inequality diminishes as we
proceed upwards, and is finally reversed.
In the next two sections (Q. 6 and W. 1915 C) there is no change worthy of
note in the position of the bundles. In the 4th section (Q. 11) one of the
bundles has been displaced ; the natural interval had increased somewhat,
and the strands are also appreciably further out. In the 5th section (W. 1915),
while the strands are still no more than 2*7 mm. apart, they have now
advanced to a considerable distance beyond the phloem, represented by ten
or twelve layers of cells. At this level the secondary thiekening is almost
equal on the two sides of the stele *.
In the 6th section (W. 1915 B) there is a great change. The same two
bundles can be recognized, but they have moved much further apart—the
actual interval is 5 mm., and there does not seem to have been much displace-
ment. One bundle is cut obliquely, and lies considerably further out than
the other. The pericycle around the bundles is immensely enlarged (about
3 mm. thick). Midway between the two strands an adventitious root passes
out horizontally, and can be traced for some distance.
In the last section of the series (W. 1915 A) the bundles are still further
separated (6:5 mm. clear interval). One is still much further out than the
other, about 1:7 mm. as compared with 0*7 mm. from the stele, but not much
importance should be attached to this difference, as the section is here
somewhat oblique and the tissues in places damaged. Only the more remote
bundle shows signs of division. Both strands are here also imbedded in a
great mass of apparently pericyclic tissue (see Pl. 2. Phot. 11). A fragment of
Sparganum hypoderma is present on the outside, but no other cortical
tissues appear. The whole mass might be taken for a leaf-base, but the
bundles are not far out enough for this, and one can only regard the excessive
development on this side of the stem as due to the presence of adventitious
roots. In this section, as in the last, a root (different from the previous one)
* It is in this section that the young root, previously figured, occurs ( Williamson & Scott,
1895, pl. 27. fig. 29). It lies on the side opposite the leaf-trace.
OF THE BRITISH COAL MEASURES. 81
passes out between the two leaf-trace strands; in this case its connection
with the wood of the stem is clearly shown. The wood is here more developed
on the leaf-trace side than on the other, but is very irregular and many of
the tracheides are cut longitudinally.
Tt is evident, from the last two sections, that a row of adventitious roots
was given off between the two bundles of the leaf-trace. This agrees with
the evidence from a longitudinal section (W. 1915 G) previously figured and
described *. Here three roots are seen in a vertical series; just as in the
transverse sections the pericycle “has undergone an enormous thickening,
with tangential cell-divisions, around the bases of the roots." The anomalous
development of secondary wood in the same region was also mentioned
and compared to the “réseau radicifére” of Van Tieghem. Ina parallel
longitudinal section t of the same portion of stem, the roots are missed, but a
leaf-trace bundle is shown in the corresponding position. This was doubtless
one of the two bundles flanking the row of roots, as shown in the transverse
sections. The longitudinal sections show clearly that the local hypertrophy
of the pericycle had nothing to do with a leaf-base, for the leaf-trace strand
shown is still running almost vertically and not bending out.
For the reasons already given, I have no doubt that the two bundles
shown in the transverse series belong to the same trace; their wide
separation may be due to the presence of the roots between them, and it is
possible that they may have again converged before entering the base of
the leaf.
Throughout the series no other trace-bundles are given off. The seven
sections, considering the date at which they were cut, would represent a
length of quite an inch, so we may infer that the leaves were widely spaced
with long internodes.
Specimen 2 (P1. 4. fig.13; S. 445), a smaller stem (about 9 x 8 mm.), likewise
shows two bundles only in the transverse section; they are both at one
end of the compressed stele, and.the interval between them is 16mm. They
lie just outside the wood of the stele. No other trace is given off in the
three sections of this specimen 1, all of which show the same two bundles.
Between them there are two marked protrusions of the secondary wood,
which appear to be composed of short pitted tracheides. These are evidently
connected with the insertion of adventitious roots, which are clearly shown
in one of the sections (Q. 64). Two of the roots are seen side by side; there
may have been a double row, as was possibly the case in the previous specimen
also. A huge mass of ill-preserved tissue encloses the roots and bundles.
* Williamson & Scott, 1895, p. 757, pl. 27. fig. 28.
T W. 1915 R.
t The order from below upwards seems to be: S. 445, Q. 64, W. 1915 H, but as the two
latter are very oblique it is difficult to determine.
82 DR. D. H. SCOTT ON THE HETERANGIUMS
In this specimen the arrangement was evidently the same, in essentials, as
in specimen 1, the leaf-trace being double, with widely separated strands
and roots between them.
Specimen 3 is a little different. In one section (S. 446) the bundles of the
one pair shown are only about 0*8 mm. apart ; the preservation of the outer
tissues is, however, so bad that one eannot be certain that the interval is
natural. In a better-preserved section, probably of the same stem (W. 1915 P),
two bundles are shown well out in the pericycle and 1:6 mm. apart ; each
shows signs of division. There is no certain evidence for a second trace in
any of the sections. [t is evident that this specimen really agrees with the
previous two ; if the strands of a trace are sometimes nearer together, this
may well be due to the absence of adventitious roots between them.
Specimen 4 is of the same type. A pair of bundles lies in the pericycle,
1:64 mm. apart (in R. 645). Between them a root passing out horizontally
is beautifully shown. The stem is a comparatively young one ; the secondary
wood is considerably thicker on the side where the trace and root are, but
there is not yet any excessive hypertrophy such as one finds in older stems.
Specimen 5, already referred to for the good preservation of the phloem
and the absence of Sparganum structure, is peculiar in the character of the
leaf-traces. In the section photographed (S. 625; Pl. 2. Phot. 13) two traces
are shown, each consisting of two bundles. The inner trace (T. 1) is passing
out through the secondary wood; in this position each strand has its own
secondary arc. The primary xylem-groups are about 1:6 mm. apart, and are
unusually small, about 300 u and 240 u respectively in tangential diameter,
about half the diameter of those in the smaller stems previously considered.
The primary tracheides are also small—only the outer part of the stelar
strands appears to have contributed to the trace-bundles. This trace is of
interest, for it is the first one we have met with in the act of leaving the
wood, and it shows that the bundles were well separated at their first origin
from the stele.
The outer trace (T. 2, almost opposite the former) is in the pericycle. It
consists of two small strands (about 220 u in diameter) about 1 mm. apart.
They have no secondary wood. In the next section (R. 639), which appears
to have been eut about 7 or 8 mm. lower down the stem *, the inner trace has
disappeared, having completely fused with the stele. The outer trace is still
in the pericycle, the bundles here are larger (about 320 p) and only about
0:75 mm. apart. They are fully as large as those of the inner trace; so we
must assume that these two bundles represent an entire leaf-trace and not
the product of any subdivision. The distance between the strands of a pair
is evidently a very inconstant feature. In this specimen several adventitious
* This is roughly calculated from the dimensions of a 7rigonocarpus seen in transverse
section in both slides.
OF THE BRITISH COAL MEASURES. 83
roots are present; their position, unlike that of the roots in previous
specimens, shows no relation to the leaf-traces.
I believe that specimen 6 is part of the same stem, though the state of
preservation is different, the phloem having collapsed. The surrounding
objects are the same, and the structure of the stem agrees essentially, especially
in the absence of any well-marked Sparganum hypoderma. The two sections
W.1915 D & E each show two double leaf-traces too near together for any
probable phyllotaxis, but possibly displaced. They are rather small bundles,
300—410 w in diameter. The main point is that here again two double traces
are seen in the same transverse section.
A third section (N.3) is photographed (Pl. 2. Phot. 12). Itiscomplete, which
the two Williamson sections are not, and probably belongs to the same stem,
though there is no absolute proof. Two double leaf-traces are shown, at a
divergence which may be roughly estimated at 2. One trace (T. 1) is in the
outer part of the pericyele, the other (T. 2) may have just entered the cortex.
The bundles of each trace are near together * and their relation to each
other is obvious. In size they are somewhat larger than in the previous
sections (410—490 y).
The stem just described is a variant on the £ type first considered, though
the distinctions are not important; the peculiarities are that two traces are
usually seen in the same transverse section, that the two bundles of a leaf-
trace are usually near together, that the bundles are, on the average, small,
and their tracheides also small. It may be added that no regular relation
appears to obtain in this case between the roots and the leaf-traces. Con-
sidering the poor development of the mechanical tissues it is not improbable
that the stem was of the nature of a rhizome.
The A specimens generally establish the fact that the leaf-trace was
eonstantly double from its starting-point in the stele outwards, the two
bundles being often remarkably far apart. We have so far no conclusive
evidence as to their further subdivision. It may be mentioned that in this
species the trace-bundles carry no secondary wood out with them when they
enter the pericycle.
Further considerations may be postponed till we have compared the other
forms of stem in the species.
H. Lomaxii, Form cylindricum f.
In the joint memoir with Williamson this form was provisionally treated
as distinct, for it was obviously different from Æ. Grievii, with which the
other Dulesgate specimens were at that time confused, while it was not
* Those of the outer trace are evidently displaced, and appear closer together than in
nature.
T Williamson & Scott, 1895, p. 764, pl. 28. figs. 30, 31.
84 DR. D. H. SCOTT ON THE HETERANGIUMS
identical with H. tilivoides. Now that H. Lomaaii is recognized as a distinct
species it becomes evident that the “ cylindricum” specimen is merely a
form of it, a fact which Mr. Lomax realized many years ago.
The specimen happens to be of considerable interest. There are six trans-
verse sections (only two of which are good), two longitudinal, and one
oblique *. The eylindrical form of the stem is preserved with little distortion,
while all the other Dulesgate specimens are more or less compressed. The
outer cortex has a cnrious carbonized appearance, and may have undergone
some change which made it more resistant than in other cases.
The best transverse section is one in the University College collection, and
is shown in Pl. 2. Phot. 14. The Williamson section previously figured
(Williamson & Scott, 1895, pl. 28. fig. 30) is rather less complete, though the
cylindrical form is more perfect. The stem is 6°75 x 5 mm. in diameter.
The primary wood is well preserved, and agrees in every respect with that
of the specimens of //. Lomawii already described. A few layers of small-
celled secondary wood had been added in places. Scarcely any of the phloem
is preserved, and in the pericycle only the sclerotic nests persisi ; their
presence, once regarded as distinetive, is of course common to the Dulesgate
stems generally. The cortex is quite well preserved—much better than is
usual in the 8 specimens. The masses of sclereides, with radiating cells
round them, are conspicuous and have assumed a black colour, The
longitudinal seetions show that they form solid blocks as in other Dulesgate
stems. The Sparganum hypoderma has the structure typical for the younger
stems of H. Lomazii (see above, p. 78).
The point of real interest lies in the leaf-traces, of which two are shown
` in the section photographed. The inner pair (T. 1) is only just beginning to
separate from the stele, in the outline of which the two strands (each about
700 w in tangential diameter) form gentle protrusions, cut off at the back by
conjunctive tissue. They are quite near together, at an interval of about
120 p only. This is the earliest stage in the separation of a leat-trace from
the stele which we have yet met with, and it shows again that the two
bundles were distinct from their origin.
The outer trace (T. 2), at a divergence of perhaps ? from the former, is in the
cortex and consists of two double bundles, about 1150 p apart. The division
of each of the original strands into two is quite evident, and shows that here,
as in H. shorense and H. tiliwoides also, the trace ultimately became a 4-stranded
one. A protrusion of the cortex in the region of the leaf-trace indicates the
beginning of the base of the leaf.
In the section previously figured (W. 1304 A), which appears to have been
* The six transverse sections, in their probable order from below upwards, are: N.19;
W. 1304 A ; R. 621; S. 444; S. 446; N. 7.
The longitudinal sections are W. 1304 B and R. 644; the oblique section is W. 1915 P.
When the joint memoir was written, only the two sections W. 1304 A & B were examined.
OF THE BRITISH COAL MEASURES. 85
cut a little distance above the University College section, the outer trace has
disappeared, having no doubt passed out into the petiole. The double trace
shown (l. e. pl. 28. fig. 30, lt.) in the disorganized pericycle corresponds in
position * to that which in the former section was still in connection with the
wood. At the level shown, the two bundles have not yet begun to divide
into four. No other trace is present in the Williamson section.
The other transverse sections afford no new data, but the Williamson
longitudinal section (W. 1304 B) is of considerable importance, as it shows a
leaf-base in connection with the stem, a point not noticed at the time of the
previous deseription. The preservation of the leaf-base is not very good, but
the main features of the structure are clear. The petiole is attached obliquely
to the stem, which it appears to exceed in width; the cortical tissues are
perfectly continuous throughout. A layer of sclerenchyma, seen in tangential
section, forms a partial barrier between stem and leaf-base. The latter has
a Sparganum hypoderma and sclerotic masses like those of the stem. The
important point is that it contains two double bundles or bundle-pairs. They
are widely separated, but in each pair the division into two has only gone
about as far as in the transverse section figured (Pl. 2. Phot. 14). It appears
then, that the leaf-trace passed into the base of the leaf as two bundle-pairs,
and not as four independent bundles, thus resembling H. tiliwoides rather
than H. shorense.
The “eylindricum” specimen thus throws a good deal of light on the
course of the leaf-traces. It does not differ in any essential respect from the
other examples of Æ. Lomawii described, though it does not exactly agree
with any of them; the trace-bundles are fairly large, but do not separate
very widely from eaeh other in passing through the stem; this is probably
because in this case no roots arose between the strands. The leaves were at
somewhat long intervals, for in most of the transverse sections only one trace
is met with. The * eylindricum" stem is evidently a specimen of the rather
variable Dulesgate species, /7. Loma«ii.
H. Lomazii, Form a.
This form was shortly described and admirably illustrated by Williamson
in his 17th Memoir (1890, p. 96, pls. 14, 15. figs. 14-18). — A section of it
was also figured in our joint paper (Williamson & Scott, 1895, p. 753, pl. 26.
fig. 21) and shows a branch in connection with the stem ; I believe this is
still the only recorded case of branching in the genus Zeterangium.
The specimens are all more or less crushed, and present a rather singular
appearance, different from that of most Heterangiums. This is chiefly due
to the faet that the cortex, owing to the great development of the sclerotic
* As the two slides are reversed, it is impossible to figure them with corresponding
orientation.
$86 DR. D. H. SCOTT ON THE HETERANGIUMS
masses, is less crushed than the stele. The specimens are closely associated
with those of the 8 form, sometimes appearing in the same sections *.
The stems vary in size. Williamson's type-stem (specimen 1 ; W. 1915 H)
measures about 14:5 x 5*5 mm., with wood (badly crushed) about 6 x 2:5 mm. in
diameter. This is only exceeded in size by one of the 8 stems. Specimen 2
is about the same size, with a smaller branch. Specimen 24 is a little
smaller than the first two ; while the obscure specimen 1 a, of which only the
stele is preserved, must have been fairly minute, the wood only measuring
about 2:5 x 1:5 mm.
So far as the erushed state of the tissues allows one to judge, the structure
of the wood did not differ from that of the specimens already described,
unless it be that in the æ stems the centrifugal wood of the bundles is some-
times rather more developed. With one partial exception, to be mentioned
presently, no secondary wood has been detected.
The pericycle contains sclerotic groups as in other specimens. The cortex
is the best-preserved part of the stem, and presents a rather striking appearance
owing to its strong mechanical construction, the amount of sclerotic tissue
exceeding anything observed in the other forms (see Pl. 4. figs. 16 & 17,
and compare with fig. 13 and Pl. 2. Phots. 9, 12, 13, 14). The longitudinal
sections confirm this (see Williamson’s figures, 1890, pl. 15. figs. 17 & 18,
and compare fig. 15 here). The sclerotic nests are both large, often exceeding
1:6 mm. in diameter, and numerous. The outer cells are elongated radially
to the mass. The hypoderma is well developed, with a preponderance of the
fibrous tissue, as is usual in young stems. The epidermis is well preserved
in places.
Owing to the crushing of the stele the specimens are unfavourable for
following the course of the leaf-traces, for portions of the wood are often
mechanically severed from the main mass and may simulate outgoing strands,
while genuine leaf-traces may be crushed in and become unrecognizable.
In Williamson’s figured specimen (l. c. fig. 14) the double strand at a may
probably be a leaf-trace, but there is no proof. On the other hand, the two
double bundles, a little higher up on the left, certainly constitute a trace.
They are best shown in a Manchester section (Q. 10) cut just above Williamson's.
This is altogether a good section, with the wood rather less crushed than
usual; the leaf-trace is particularly clear (Phot. 15). Whether it is in
* The known specimens of the a form are as follows: Specimen 1. W. 1915 IT; Q. 10,
R. 643, R. 657 (transverse); W. 1915 F (longitudinal) from the same stem. la. A smaller
stem appearing in the four transverse sections, as above. The larger stem, 1, may be taken
as the type (Williamson, 1890, figs. 14-16, 18). Specimen 2. W. 1885 H*, 1885 HH,
1915 M*, 1915 N*, 19150; R. 642*; S, 236% (transverse). 2«. A smaller stem occurring
in the sections marked * above, Specimen 3, W. 1915 E (transverse); W. 1915 I, 1915 K,
1915 L (longitudinal) probably from the same stem. The order of the sections will be
mentioned in the text when necessary.
OF THE BRITISH COAL MEASURES. 87
the pericycle or cortex is hard to say, the two tissues not being at all sharply
delimited here—at any rate, it is separated from the disorganized phloem of
the stele by several layers of more or less sclerotic parenchyma. The two
double bundles of the trace are about 120 u apart; they are fairly large
strands, measuring 615 w and 820 p respectively in tangential diameter, and
each is manifestly double, for the xylem is lohed and divided radially down
the middle by a band of collapsed cells. This section, then, proves that the
leaf-trace consisted of two bundles, each of which prepared to divide before
it had got far on its outward course. The adventitious root figured by
Williamson shows no obvious relation to any leaf-trace.
In specimen 2 the order of the sections from below upwards proves to be :
W.1885 HH, W. 1915 N, W. 1885 H, W. 1915 M, W. 1915 0, S. 236, R. 642.
So long a series seems promising, but the stem is in many sections incom-
plete and the crushed condition perplexing. It is doubtful whether the two
“ leaf-traces ” marked on our fig. 21, of 1895, were really of that nature;
the only undoubted traee is near the broken end of the section. In the
present fig. 16 (Pl. 4), from this specimen, the double strand v.b. may be a leaf-
trace. So far as the evidence goes, it confirms that from the type-specimen.
In specimen 2 a, which appears in five sections of the same series, a double
leaf-trace can be followed through the whole thickness of the cortex. [n
the section figured (fig. 17) the bundles are far out and may have been entering
the leaf-base ; the tissue, however, is damaged and one of the two strands
(v.b.!) lies in the débris. The signs of division in each bundle are less marked
here than in the other cases mentioned.
So far as the available data show, the behaviour of the leaf-trace in the a
form of stem seems to have been much the same as in the *cylindricum
form, already described, the trace consisting of two large strands which
pass out without separating very widely, and do not complete their subdivision
till after they have entered the leaf-base ; in both cases more than one trace
may be met with in the transverse section,
A word may be added about the branching, already described and figured
in the joint memoir *. The branch appears in three successive sections of
specimen 2. In the lowest section the continuity between branch and stem
is more complete than in that figured (the middle one); the stele of the branch
is smaller, and its zone of secondary wood thicker. At this level there is a
little secondary wood on the main stele also—a small arc 620 u broad and
about 6 cells thick on the side towards the branch. It dies out higher up the
stem. In the third section (W. 1885 H; above that figured) the branch is
free from the stem, but still shows no leaf-traces of its own ; it is, however,
not quite complete. At this level, the secondary wood of the branch has all
* Williamson & Scott, 1895, p. 753, pl. 26. fig. 21.
T W.1885 HH, 1915 N, 1885 H.
TURCUCETMOEN GUME wa, a "NEN Sn SO eee ke ue i
88 DR. D. H. SCOTT ON THE HETERANGIUMS
but died out, and the structure is that of an ordinary stem of tbe a form.
The branch is very similar to the smaller stem (24) appearing in the same
section, and it is quite likely that the latter may represent another branch of
the same main axis. It is evident that secondary growth started at the base
of the branch, where it also slightly affected the parent stem.
As regards the nature of the æ form of stem, there is no doubt that it has a
distinetive anatomical habit, and we cannot wonder that Williamson at first
inclined to make it a separate species. Even allowing for the better
preservation of the cortex in the younger stems, the difference can hardly
be reduced to one of age; I have seen no B stems that can naturally be
interpreted as the older stages of æ stems. The youngest 8 specimens, which
ought to come nearest to the « form, do not possess the powerfully constructed
cortex which characterizes the latter.
Not much stress can be laid on the fact that an «æ specimen is the only one
that shows branching. ‘The specimens after all are few, and we must
remember that it was a long time before branching stems of Lyginopteris
oldhamia were found, common as the plant is. In ZJeterangium generally,
branching must have been extremely rare ; whether it was soin the Dulesgate
species we cannot tell.
The various forms of stem described under H. Lomaaii all occur in a single
coal-ball ; it appears that none of them have been found in any other block.
It seems almost infinitely improbable that two or more distinct species of the
same genus, each of the extremest rarity, should happen to be associated in
one unique nodule; the presumption is overpoweringly strong that all the
specimens belong to one and the same species. The æ form is the most
distinet in appearance, but I do not doubt. that Williamson's later view was
essentially correct and that it represents a younger stage of growth of one
of the other Dulesgate forms of stem, though not perhaps of a form exsetly
identical with any of those which we possess in a more advanced state.
Whether the differences among the specimens of H. Lomawiti are due to
variability in homologous stems, or to the occurrence of multiform stems in
the same plant, cannot be decided. The latter seems more probable, but we
have no proof, for in the only case where a stem has been observed to branch
the branch merely repeats the characters of the parent axis.
Petioles.
The evidence we have collected so far only proves that in H. Lomazii
two bundles, each showing signs of division, entered the leaf-base ; we do
not yet know how many distinct strands were present in the petiole. The
material contains a few specimens of petioles, though none are well preserved.
As they are associated with various forms of H. Lomaaii stem, it is impossible
to tell to what particular form they belonged ; from their size, they were pro-
bably borne on the larger stems. Three of these petioles are tolerably clear—a
section of one of them is shown in Pl. 4. fig. 18. In other sections of the same
CMT ONERE NAR Sl
OF THE BRITISH COAL MEASURES. 89
petiole the sclerotic nests appear, and the dense hypoderma is also that of a
fleterangium*. The tissues are much permeated by roots and rootlets, but
the vascular bundles belonging to the petiole are easily recognized in all the
sections. There are four large bundles, two widely separated in the section
figured, and two close together. The two more median bundles are clearly
dividing; where the structure is well enough preserved, the bundle is evidently
mesarch with little centrifugal wood, and is thus completely of the Heterangium
Lomacii type. This petiole is much compressed, and measures where complete
about 6x1 mm. There is no doubt that it belongs to the Heterangium,
and it thus proves that the bundles completed their division into four after
entering the petiole, and even initiated further subdivision.
Another larger and much crushed petiole t (75x1 mm. in diameter)
shows, in the best section, four bundles, two of which are dividing. A third
specimen f, of about the same size (6 x 1:5 mm.), contains four large bundles,
quite separate, but not obviously dividing further. All these show the
Heterangium characters unmistakably. It is a pity that we have no petiole
in continuity with the stem, but the evidence is enough to show that in
H. Lomawii, as in H. shorense and H. tiliwoides, the petiole was polydesmic,
the minimum number of bundles being four.
It is evident that the three species of Heterangium which have now been
described were nearly allied; they agree among themselves in having
polydesmie petioles, and are the only species recorded in which this
character has been observed, though it may be found to extend to others.
The forms now grouped under H. Lomasii appear, for reasons already
given, to constitute a single species, distinct though not very different, from
H. shorense and H. tilieoides. The specific characters may be summed up as
follows :—
Heterangium Lomacii, Will. in litt.
Stem from about 5 or 6 mm. to about 198 mm. in diameter.
Primary wood consisting of somewhat irregular groups of tracheides, with
parenchyma between the groups.
Tracheides without any spiral bands between the pits.
Peripheral xylem-strands and leaf-trace bundles inesarch with little
centrifugal wood.
Secondary phloem sometimes well developed, but phloem-rays not much
dilated.
Pericycle generally of moderate breadth, but much enlarged where leaf-
traces or roots pass out.
* This petiole occurs in five sections of the specimen 2 (æ form) series, namely
W. 1915 N, 1885 H, 19150; S. 236 and R. 642.
T Occurring in the sections W. 1915, 1915 B, 1915 A.
t In the University College slide, N. 3.
LINN. JOURN.—BOTANY, VOL. XLIV. H
CT SC TORTE NN
ee may COE Pye Vor, EIRE UU T ER
" i
90 DR. D. H. SCOTT ON THE HETERANGIUMS
Sclerotic tissue present both in pericycle and cortex ; the sclerotic nests
in the cortex forming massive blocks, sometimes (in stems of the « form)
developed to a remarkable extent.
Leaf-trace consisting of two bundles where it leaves the stele, the
constituent bundles often very widely separated in the cortex, when
adventitious roots pass out between them. Trace-bundles without secondary
wood of their own. The double trace beginning to divide into four as it
enters the leaf-base. Four bundles, sometimes further subdividing, in the
petiole.
Number of traces in the transverse section, one only in some specimens,
two or perhaps more in others. Traces traversing the stem for a long
distance before passing out.
Stem observed to branch, the secondary growth in this case starting at the
base of the branch.
Details of cortical structure and of arrangement of leaf-traces varying in
different forms of stem.
Locality. Dulesgate, Lancashire: Lower Coal Measures.
All the specimens, apparently from a single nodule, discovered by
Mr. James Lomax about 1890.
It may be mentioned that the present re-investigation has thrown no new
light on the structure of the root in Heterangium Lomax. That question
remains now as it was in 1895 *.
HETERANGIUM MINIMUM, sp. nov.
A single specimen of this small stem occurs in a block from Dulesgate,
of which sections were received from Mr. James Lomax in February, 1903 f.
Though the locality is the same as that of H. Lomazii, the material is quite
different, and contains different plants. There is therefore no presumption
of specitic identity between this form and those referred to H. Lomaxii. The
characters indicate that it is distinct.
The dimensions of the stem, about 2:9 mm. by 1:7 mm. in its somewhat
crushed condition, are unusually small for a Heterangium. The best section is
shown in Pl. 4. fig. 19. The wood, including a few secondary layers, measures
about l-6x 1 mm. The primary wood consists mainly of tracheides, with
comparatively little conjunctive parenchyma. The peripheral xylem-strands
are not sharply defined, and the principal rays of the secondary wood by no
means conspicuous. In these respects the specimen recalls H. Grievit rather
than the Coal-Measure species. The tracheides of the metaxylem are decidedly
large for the size of the stem, often exceeding 150 p in diameter. The
structure of the peripheral strands is mesarch, with one or two layers of
centrifugal primary wood. The secondary wood, which reaches a maximum
* Williamson & Scott, 1895, pp. 758, 764. The evidence there brought forward relates
entirely to the Dulesgate species, H. Lomarti
t The sections are S. 1801, 1803, 1804, 1805, and 1813, all transverse.
2 ‘OF THE BRITISH COAL MEASURES. 91
4
thickness of about seven elements, is small-celled, as usual, the tracheides
seldom exceeding 36 u in diameter. The phloem is destroyed ; the pericycle
tolerably preserved in places. No sclerotic nests have been detected either
in the pericycle or the cortex—one would not expect a strong mechanical
construction in so smalla stem. The cortex is parenchymatous, with a rather
feeble Sparganum zone on the outside.
In the section figured a leaf-trace is seen in the’ pericycle. — It is a single
strand, though somewhat lobed ; other sections show that the trace divided
into two strands further out on its course, but they have not been seen to
diverge. In one section (S. 1805) a second trace is seen, just separating
from the wood, as a single strand, with two protoxylem groups.
It thus appears that /Jeterangium minimum differs from all other forms
described in this paper in having a leaf-trace consisting of a single bundle
where it starts from the stele. In this point it agrees with M. Grievii, but
differs in the fact that the trace divided into two in the cortex. Of course,
it cannot be strictly proved that /7. minimum may not have been a minute
twig of some other species ; there is, however, nothing to indicate that this
was the case, and on present evidence the plant must be regarded as distinct.
The specific characters may run as follows :—
Heterangium minimum, sp. nov.
Stem minute, under 3 mm. in diameter. !
Primary wood consisting chiefly of tracheides with a few parenchymatous
bands. Peripheral xylem-strands not well defined ; mesarch, with a layer or
two of centrifugal wood. i
Sclerotic tissue apparently absent from both pericycle and cortex. Leaf-
trace a single bundle where it leaves the stele, dividing into two in the
cortex. :
Locatity. Dulesgate, Lancashire : Lower Coal Measures.
Found by Mr. James Lomax in 1903,
A New SUBGENUS.
If for the moment we leave /Jeterangium minimum out of account, we find
that the other three British Coal-Measure species of the genus, while differing
only in trivial points, agree in the following characters: the distinetness of
the peripheral xylem-strands, and consequently of the principal medullary
rays; the tendency towards exarchy in the primary xylem-strands, the
centrifugal primary wood being little developed ; the compound leaf-traces,
double on starting from the stele, quadruple or more on entering the petiole.
The first of these characters may reasonably be considered as an advance
on the structure of the H. Grievii type, in which the peripheral strands of
the stele are not sharply delimited. Their greater distinctness is a departure.
from the pure protostelic structure, and indicates a progressive downward
differentiation of the leaf-trace system.
H2
Ett
92 DR. D. H. SCOTT ON THE HETERANGIUMS
On the other hand, we cannot be certain that the exarch tendency of the
primary xylem-strands marks an advance, for we find both types of structure
in species of similar age, as will be shown below. In the Coal-Measure
forms, the centrifugal xylem, though little developed, is sharply differentiated
and obviously primary ; indeed, from the nature of the traeheides (spiral or
scalariform) it would appear that the centrifugal part of the strand was the
first to be formed after the protoxylem, with which it is in direct connection,
as in H. Grievii and Lyginopteris, a point on which Chodat has laid stress *
(Chodat, 1908, p. 13).
In the double leaf-trace and polydesmie petiole, the Coal-Measure species
undoubtedly show a more complex organization than has hitherto been
described in /Jeterangium, or even in Lyginopteris, for we have here to do
with a trace which is double at its origin, not as in Lyginopteris with one
that merely divides as it passes out. The further subdivision of the trace,
giving rise to four or more bundles in the petiole, gives a most distinct
character to the species in question ; the creation of a new genus for them
might be suggested. There are, however, objections to this course, and I
prefer to be content with proposing a provisional subgenus, which may be
named PorvANGIUM f, and may best be characterized simply by the leaf-trace
being double on starting from the stele, and dividing into four or more
strands in the petiole.
The proposed new subgenus will, of course, include the three species
Hl. shorense, H. tilimoides, and H. Lomasi, with the probable addition of
others. Æ. minimum, on the other hand, will fall under the other subgenus,
which may be called Ev-HETERANCIUM, characterized by the single leaf-trace
given off from the stele remaining single or dividing into two strands only
on its passage outwards. Ihave not thought it desirable to introduce the
other characters (distinctness of xylem-strands and tendency to exarchy )
into the diagnosis of the provisional subgenus, as we have no reason to
assume that they are consistently correlated with the polydesmie condition.
A rapid comparative survey of the other species of the genus and of some
related groups will serve to make the position clear.
CoMPARATIVE CONSIDERATIONS.
Corda’s original species /Jeterangiwm paradoxum 1 was founded on fragmen-
tary material, which practically shows nothing beyond the structure of the.
metaxylem. Kubart (1911, p. 9, Taf. i. fig. 3), who examined smaller
* Prof. Chodat uses the position of the protoxylem as an argument against the homology
of the bundle in Lyginopteride with that in Cycadaceæ, a point with which we are not
concerned in the present paper.
+ Intended to suggest a polydesmie Heterangium.
{ First published in 1845. The reference is to the edition of 1867: ‘ Flora der Vorwelt,’
p. 22, pl. xvi.
OF THE BRITISH COAL MEASURES. S
specimens from the same locality, figures a complete stele, but no details are
given, and beyond confirming the general agreement with other species
referred to the genus, there is nothing to be said. The plant is apparently
of Middle Coal-Measure age.
Williamson’s Lower Carboniferous species H. Grievii is too well known
to need any discussion here. It may, however, be convenient shortly to
recapitulate the principal characters :
Peripheral xylem-strands not sharply delimited.
Primary centrifugal wood of the bundles well developed.
‘Secretory sacs” scarce or absent.
Sclerotic plates (in the form of thin dises) present in the cortex, but not in
the pericycle.
Leaf-trace single and a single bundle in the petiole.
Leaf-bases large and decurrent.
Foliage probably that of Sphenopteris elegans.
This species is the type of the subgenus Hu-heterangium. I am not aware
that the structure of any other Lower Carboniferous species has been
described *.
In the meantime, Renault (1869, p. 177) had described two species, under
the names Lycopodium punctatum, Ren., and Lycopodium Itenaultii, Brongn.,
which have proved to belong to the genus Heterangium. These fossils came
from the Upper Coal Measures of Autun. The reference to Lycopodium
involved a misinterpretation of the structure of the specimens T, and it is
more profitable to refer to Renault’s later description (Renault, 1896, p. 253),
written after he had recognized the true nature of the plants. The only
figures are those in the earlier paper. Both species are represented by small
stems, 5-5-6 mm. in diameter. //. punctatum has a certain amount of
secondary wood and bast, which are absent in H. Renaultii; it is possible, us
Renault pointed out, that the latter may merely be a younger condition of
the former (Renault, 1896, p. 256). 1 examined the original preparations
at Paris in 1905, but at that time my attention was not directed to the points
which now seem most critical. H. punctatum, especially, bears a close
resemblance to our H. tilicoides; it has, for example, large medullary rays,
which are dilated in ihe phloem-zone (Renault, 1869, pl. 12. fig. 3; pl. 13.
fig. 4). The leat-traces are said by Renault (1896, p. 254) to pass out
opposite the rays, a statement which is scarcely confirmed by the figure (1869,
pl. 12. fig. 1). Renault gives no definite information as to the position of
ihe protoxylem, merely stating that the attenuated peripheral extremities
of the vascular groups are occupied by rayed tracheides and spiral elements
(trachées).
* Prof. T. Johnson's species, H. Aibernicum, is based on external characters only (Johnson,
1912).
1 E. Williamson & Scott, 1896, p. 771.
«XX
WENMTITUMERSAT S, he RARIOR SE TERR ERR RE RM ONE eA US P PRESSE RET MA
Yr
94 DR. D. H. SCOTT ON THE HETERANGIUMS
In describing H. Renaultii, he makes the important remark that “on one
side of the woody cylinder we notice two bundles which are separating from
the axis, and in the parenchymatous part of the cortex two others which
have already quitted it for a certain time” (1896, p. 255). It thus appears
that this species gave off its leaf-trace bundles in pairs. He also observed
>in the bundles. I have little doubt that both
forms may be referred to the subgenus Polyangium, but a re-investigation
is required.
Another Autun species, H. Duchartre’, Ren., was originally described under
the name of Porozylon Duchartre’ (Renault, 1879, p. 276, pl. 14. figs. 4-8) ;
it was excluded from that genus in 1886 (Bertrand & Renault, 1886, p. 245),
and appears as a /Jeterangium in the ‘Flore d'Autun et d'Épinac? (Renault,
1896, p. 251, pl. 65. figs. 1, 2). The stem (6:5 mm. in diameter) has very
much the characters of H. tiliwoides, but little beyond the wood is preserved.
'The prineipal rays are conspieuous, dividing up the secondary zone into
15 sectors (Renault, 1896, pl. 65. figs. 1, 2) *. Here, again, the groups of
spiral and scalariform tracheides are described as lying at the periphery of
the primary wood. Two bundles, widely separated, but on the same side of
the stem, are seen in the cortex ; there can be little doubt but that they both
belonged to the same leaf-trace. It appears, then, that this form also may be
included under the provisional subgenus Polyangium. Renault calls atten-
indications of “dichotomy '
tion (1896, p. 254) to the similarity between M. Duchartrei and H. punctatum ;
it is possible that all the three forms just described may eventually be reduced
to one species.
On the other hand, a fourth Autun form, Æ. bibractense, Ren. (Renault,
1896, p. 252, pl. 65. figs. 3-6), seems to be quite distinct. It is remarkable
for the small size of the primary wood (1-1:5 mm. diameter) and the great
thickness of the secondary tissues f. Parts of the phloem and the cork are
preserved. "There are large prineipal rays as in the preceding species.
Nothing is known of the leaf-traces. No other Heterangium has been found
in such an advanced state of growth.
It will be noticed that all the Autun specimens, so far as their structure is
known, agree better with the British Cosl-Measure forms of the Polyangium
type than with the Lower Carboniferous species //. Grievü. his is natural,
considering their age, which is considerably later even than that of our Coal-
Measure forms.
We have now briefly to consider the very interesting series of species of
Heterangium recently described by Dr. Kubart from the Ostrau beds of
Upper Silesia (Kubart, 1908, 1911, 1914). At present only preliminary
accounts have been published, so the data for a full comparison with our own
* The figures of the 1896 memoir appear to be much more exact than those of 1879.
+ Renault gives the thickness of the wood as 55 mm. I found it to reach nearly a
centimetre in places.
OF THE BRITISH COAL MEASURES. 95
species are not yet available *. The Ostrau specimens, which are admirably
preserved, are very interesting from their age, which is somewhat earlier
than that of our Lower Coal Measures, corresponding, according to Dr.
Kubart, to the upper part of our Millstone Grit. The great point which his
investigation has brought out is the existence of transitional forms between
the genera Heterangium and Lyginopteris, confirming the close affinity
between them, as maintained by Williamson and the present writer. Here,
however, our point of view is somewhat different ; we are interested rather
in the comparison between Dr. Kubart’s species and the British Coal-
Measure Heterangiums.
Five species of Heterangium from Ostrau are described—all appear to be
new. The simplest structure is shown by H. Sturii, where there is practi-
cally no differentiation in the stele between metaxylem and primary bundles
(Kubart, 1914, p. 3, Taf. 2. fig. 3) ; the latter only become delimited when
they begin to pass out as leaf-traces. The protoxylem, however, is nearly
exarch. In the figure, five large leaf-traces are shown leaving the stele : all
are single and no sign of division is to be observed ; evidently then, each
leaf received a single bundle, and H. Sturii is a typical representative of the
subgenus Eu-heterangium. It will be noticed that in this case a tendency to
exarchy co-exists with an undifferentiated stele, contrary to what we found
in the British Coal-Measure species.
H. alatum $ has more or less well-defined peripheral primary bundles; their
structure is typically mesarch, with well-developed centrifugal xylem. The
leaf-traces are numerous (I counted 10 in the transverse section) and one
bundle passed out into each leaf. A number of small winged petioles
surrounded the stem, and are sometimes found still in connection with it.
The secondary wood, when formed, is described as a perfectly continuous ring.
H. alatum is thus an obvious member of the Eu-heterangium subgenus—the
numerous small petioles give it, however, a very different character from
H. Grievit.
H. polystichum is a somewhat similar species {, with fairly distinct
peripheral xylem-strands, which, like the leaf-trace bundles, are “ centrally
mesarch,” i. e. with the protoxylem deeply imbedded. A single bundle leaves
the stele for each trace ; the traces are very numerous, 13 appearing in the
transverse section. The trace, on entering the leaf-base, may possibly
have divided into two. The species evidently falls under the subgenus
Eu-heterangium as defined above. — It is distinguished from H. alatum chiefly
by a difference in the petioles.
* Through Dr. Kubart's kindness, I have had an opportunity of examining sections of
several of his species.
+ Kubart, 1914, p. 2, Taf. 2. figs. 1, 2.
+ Kubart, 1911, p. 11, Taf. 1. fig. 4; 1914, p. 4.
96 DR. D. H. SCOTT ON THE HETERANGIUMS
H. Schusteri (Kubart, 1908, fig. 3; 1911, p. 11; 1914, p. 3) is inter-
mediate, as regards the stelar structure, between H. Sturt? and H. alatum ;
the peripheral xylem-strands are only slightly differentiated. Nothing is said
about the position of the protoxylem ; from the figure it appears to lie rather
far out, approaching //. Sturi? in this respect. The figure shows three leaf-
traces very clearly, one just starting as a single strand, another, in the
pericycle, somewhat constricted, and a third, also in the pericycle, just divided
into two. The plant thus falls under /u-heterangium, resembling our
H. minimum in the division of the originally single trace.
LI. Andrei (Kubart, 1914, p. 4, Taf. 2. fig. 4) is the most interesting of
the five species, as it shows a transition towards Lyginopteris. The
transverse section of the large stem shows quite the anatomical habit of a
Lyginopteris ; the primary tracheides, however, extend to the centre of the
pith, where they are scattered and intermixed with much parenchyma. The
peripheral strands of the xylem are quite distinct and few in number, as in
Lyginopteris; the leaf-trace starts from the stele as a single bundle, which
divides into two, much as in Lyginopteris oldhamia. In sections lent me
by Dr. Kubart, which I have no doubt were of the species subsequently
named by him H. Andrei, I found in connection with the stem a leaf-base
containing four bundles, as well as detached petioles with the same
structure *. It appears, then, that Æ. Andrei unites the characters of our two
subgenera, a single trace leaving the stele and ultimately dividing into four
in the leaf-base. This fact, if confirmed, would be a fatal objection to the
establishment of a new genus for the polydesmic forms, but our provisional
subgenera may still serve a useful purpose. Possibly Dr. Kubart may
eventually suggest a third subgenus for the Lyginopteroid species of
Heterangium. We are not here concerned with Dr. Kubart’s new species of
Lyginopteris, but may mention that L. heterangioides (Kubart, 1914, p. 4,
Taf. 2. fig. 5) is a typical member of that genus, except for the constant
presence of a few tracheides about the middle of the pith, an evident vestige
of the metaxylem of Heterangium, already becoming reduced in M. Andrei.
A species from the coal-field of the Ruhr (Westphalia), named //. inter-
medium by Kubart (1914, p. 6), is described as forming a link between
Heterangium Andrei and Lyginopteris heterangioides : we have, in fact, as
the discoverer points out, a continuous series leading gradually from the
protostele typical of /7eterangium to the siphonostele typical of Lyginopteris.
To return to our more immediate subject: the result of our survey is to
show that the older species of /Jeterangium tend, on the whole, to belong to
the simpler Hu-heterangium type, while the later species for the most part
show the more complex structure which characterizes the proposed subgenus
Polyangium.
* I noticed long, stalked glands on both stem and petiole—a very Lyginopteris-like
feature,
OF THE BRITISH COAL MEASURES. 9T
A few words may now be added about other genera which present points
of comparison with Heterangium. The nearest of these is doubtless Rhetin-
angium, founded by Dr. Gordon on the species R. Arberi, which he discovered
in the Pettyeur beds of the Caleiferous Sandstone Series (Gordon, 1912).
The plant is thus of Lower Carboniferous age and contemporary with Meter-
angium Grievit. The structure is protostelic, and the general appearance
of the transverse section suggests a rather large stem of Heterangium.
The new genus, however, differs from Heterangium in important characters :
there are no sclerotic plates in the cortex or perieycle; the primary xylem-
strands are exarch, not mesarch, and the leaf-traee is a complex corrugated
body with numerous external protoxylem groups. Minor peculiarities are
the great development of secretory sacs and cells (also present in some species
of Heterangium) and the enormous enlargement of the leal-base, which
almost equals the whole stem in thickness. The Sparganum, or more strictly
Dictyoaylon, outer cortex is very finely developed, as in Medullosa. The
exarch character of the xylem-strands is important; we have no evidence
that true exarchy occurs in Heterangium, though, as we have seen, there are
considerable variations in the degree of mesarehy. On the other hand, the
structure of the metaxylem seems to me to be altogether that of a Heterangium,
of the type, like H. shorense or H. tiliwoides, where the tracheides are in
definite packets, separated by a reticulum of parenchyma. The secondary
wood is of the ordinary Heterangium kind, with somewhat large rays opposite
the spaces between the peripheral xylem-strands.
But the most interesting point for our comparison is the leaf-trace, a
remarkable and unique structure. It embraces several of the peripheral
xylem-strands, all passing out together and all in connection laterally with
each other. The connection being on the adaxial side, the leaf-trace consists
of a number of fused U-shaped bodies, concave outwards. The protoxylem-
groups, on the extreme outside, are numerous—six or more in number.
The massive trace may remind us for a moment of a meristele of Suteliffia
(Scott, 1906), but is really quite different, for it never breaks up or divides in
any way, but passes out unaltered into the petiole. It is a complex but
never a multiple leaf-trace. — Zhetinangium thus also differs fundamentally
from our polydesmie Heterangiums, where the trace from its origin consists
of two distinct and simple bundles, which divide further on their outward
course to form more bundles of like nature. Rhetinangium is a most inter-
esting parallel development, doubly interesting from its antiquity, bui it has,
in my opinion, no special affinity with the polydesmic species of Heterangium.
Prof. Seward’s genus Megulowylon (Seward, 1899) is more remote from
Heterangium. The large stem agrees with MRhetinangium in the exarch
strueture and in the presence of several protoxylem-groups on the abaxial
side of the leaf-trace, but is peculiar in the character of the metaxylem,
which consists for the most part of short wide tracheides, apparently adapted
RTT TN
a es teh ol ell "TW Tm
98 DR. D. H. SCOTT ON THE HETERANGIUMS
for water-storage. As only the wood is known, we have no information as to
the outward course of the leaf-traces. On the existing evidence, there is
nothing to connect the genus with the polydesmie species of Heterangium,
though Megalowylon appears to represent, in a different direction, an advance
on the original protostelic type of structure.
Polydesmy is common enough among plants grouped under Pteridosperms
or Cycadofilices, but the only families w hich seem near enough to Heterangium
to deserve mention here are the Medullose:e and the Calamopityez.
A simple Medullosean stem, such as that of MJedullosa anglica, has been
described as essentially a polystelic /eterangium. Each stele, in fact,
repeats the characters of the Heterangivm monostele ; the position of the
protoxylem-groups is mesarch, but approaching exarchy, as in our polydesmic
Heterangiums. Several distinct bundles are given off from the stele to form
the trace of a single leaf and they divide up freely on their passage outwards
(Scott, 1899, p. 194; 1914 ; de Fraine, 1914). In all these points there is
a clear analogy with the plants which form the subject of this paper, the
polydesmy, of course, going much further in Medullosa than in any species
of Heterangium.
Suteliffia, though not strictly polystelic, otherwise differs from Heterangium
more widely than does a Medullosa of the anglica type. The stele is exarch,
the emission of leaf-traces takes place through the intermediation of bulky
meristeles, and the trace-bundles are concentric throughout (Scott, 1906 ;
de Fraine, 1912). The polydesmie Heterangiums no doubt show some slight
approach towards certain characters of the Medullosew, but not enough to
justify one in supposing that they are on the line of descent of the latter
family. —
A comparison with the Calamopityez is interesting. These are monostelic
plants whieh until recently have been assumed to have had a true pith,
surrounded by a ring of primary mesarch xylem-strands, the stem-strueture
thus presenting a close analogy with that of Lyginopteris. In a joint paper
on Kentucky fossil plants, of Lower Carboniferous age, Prof. Jeffrey and I
have, however, shown that the structure in the species Calamopitys americana
was essentially protostelic, the “mixed pith” containing a varying proportion
of tracheides among the parenchyma (Scott & Jeffrey, 1914, pp. 318, 326).
There is reason to suspect that the same may be true of the Thuringian
species, C. annularis (Unger). This brings the species in question nearer to
the Heterangium type of structure. The petiole (Aalymma) has long been
known to be polydesmic ; our work on C. americana showed that the leaf-
trace, single at its origin, almost immediately divides into two strands *
further subdividing in the cortex. In C. Saturni, Unger, previously
* Since the paper by Prof. Jeffrey and myself was published, I have obtained even more
convincing evidence of the early division of the trace, .
OF THE BRITISH COAL MEASURES. 99
investigated by Count Solms-Laubach *, the first division takes place further
out. These plants thus present interesting analogies with the polydesmie
species of Heterangium, though never, of course, any exact agreement. I
think this is really the most fertile comparison we can make, and that the
polydesmic petioles of our Coal-Measure Heterangiums may well be described
as showing an approach to the Aalymma (Calamopitys) structuref. Even
here there can be no question of filiation. The species of Calamopitys are
probably older than any known ZZeterangium, and certainly much older than
any which have been found to have polydesmic petioles.
While, then, Heterangium is directly connected with Lyginopteris through
the intermediate species discovered by Dr. Kubart, we have no evidence that
the polydesmie forms indicate a transition to any other genus. They present
interesting analogies with the Medulloseze and Calamopityez, and show that
there was more in common between these groups and the Lyginopteride»
than has hitherto been supposed. But we have no sufficient grounds for
assuming any direct evolutionary connection.
; SUMMARY.
The characters of the various species described have been given above
(pp. 65, 73, 89, 91). The more general results may be summed up as
follows :—
l. The three British Coal-Measure species, Heterangium shorense, H.
tilieoides, and H. Lomavii, agree in their more important characters and
only differ in details. The fourth species, /7. minimum, is of a different type.
2. The three species first named may conveniently be grouped in a pro-
visional subgenus, Polyangium, characterized by the leaf-trace, starting from
the stele as two distinct bundles, which further divide on their outward
course, the number of bundles in the petiole ranging from four to eight.
9. In addition to the polydesmie structure, these three species further agree
in the sharp delimitation of the primary peripheral xylem-strands, with the
consequent differentiation of large interfascicular principal rays, and in the
approach to exarchy in the peripheral strands and leaí-trace bundles, the
primary centrifugal xylem being comparatively little developed.
4. It is probable that, besides H. shorense, H. tilieoides, and H. Lomaazii,
the four species described by Renault, from the Upper Coal Measures of
* Solms-Laubach, 1896, p. 67.
+ There is no reason to believe that the protostelic structure of the stem existed in
Calamopitys Saturni, Unger, or in the two species (C. fascicularis, Scott, and C. Beinertiana
(Goeppert)) separated by Dr. Zalessky, under the name Eristophyton (Zalessky, 1911, p. 27).
I am not sure that I agree with Dr. Zalessky in his proposed division of the genus,
C. Saturni seems to have in some respects more in common with my C. fascicularis than with
C. annularis or C. americana. The whole question is discussed in a forthcoming paper on
the genus.
eee ee Pt
100 DR, D. H. SCOTT ON THE HETERANGIUMS
Autun (H. punctatum *, H. Renaultii*, H. Duchartrei *, and H. bibractense)
are referable to the subgenus Polyangium.
9. The Lower Carboniferous species /7. Grievii may be taken as the type
of another subgenus, u-heterangium, characterized by the leaf-trace con-
sisting of a single bundle only, where it leaves the stele, this bundle either
remaining undivided on its outward course or dividing into two strands only.
One Dulesgate species, H. minimum, falls under this subgenus, as do also
four of the Ostrau species described by Dr. Kubart, namely H. alatum,
+. polystichum, H. Schusteri, and H. Sturii. The peripheral xylem-strands
are, on the whole, less distinct in Hu-heterangium than in Polyangium, but the
degree of their differentiation varies in the different species. The xylem-
strands may either be “centrally mesarch” or show a tendency towards
exarchy. A third subgenus may be required for the Lyginopteroid species,
such as M. Andrei, discovered by Dr. Kubart.
6. The polydesmie species of Heterangium (Polyangium) show an interesting
analogy with Medulloseæ and with the protostelic Calamopityee. There is,
however, no sufficient evidence of any direct phylogenetie relation to either
group. With Rhetinangium there is no special affinity, for, while this genus
appears to have much in common with /eterangium, its leaf-trace is complex
and not multiple, thus presenting only a remote analogy with that of the
polydesmie Heterangiums. Megaloaylon is altogether more remote.
The generic characters given on p. 59 of this paper can now be made more
precise as regards the nature of the leaf-trace in different subdivisions of the
genus as stated in the Summary above, under the headings 2 and 5. 1t also
appears that in such species as /7. alatum, Kubart, and H. polystichum, Kubart,
the leaves were small and probably very different from the foliage of Spheno-
pteris elegans, attributed with good reason to H. G'rievii.
I am much indebted to my friends Prof. F. W. Oliver, F.R.S., and Prof.
F. E. Weiss, F.R.S., for the loan of slides in the University College, London,
and the Manchester Museum Collections. Without the opportunity of con-
sulting these preparations, in addition to those in the Williamson Collection
and in my own, it would have been impossible to deal at all adequately with
the subject.
The majority of the sections described were eut by Mr. Lomax, to whom
the discovery of //. shorense and of the Dulesgate species is due.
The photographie illustrations are the work of Mr. W. Tams, while the
drawings were made by Mr. G. T. Gwilliam, to both of whom my best thanks
are due for their valuable aid.
* These three may perhaps be forms of the same species.
T I
OF THE BRITISH COAL MEASURES. 101
LrrERATURE CITED.
Bertrand & RENAULT, 1886.—C. E. Bertrand & B. Renault, * Recherches
sur les Poroxylons." Archives Bot. du Nord de la France, ii.
pp. 243-389.
CnopaAr, 1908.—R. Chodat, * Les Ptéropsides des Temps Paléozoiques.”
Archives des Sciences Physiques et Naturelles, 4me Période, t. xxvi.
pp. 1-44. Geneva.
Corpa, 1845.—A. J. Corda, Flora Protogea—Beitrige zur Flora der
Vorwelt. Edition quoted, 1867, Berlin.
De Franne, 1912.—Ethel de Fraine, “On the Structure and Affinities of
Sutcliffia, in the Light of a newly discovered Specimen.” Ann. of
Bot. vol. xxvi. pp. 1031-1066.
De Frame, 1914.—* On Medullosa centrojilis, a new Species of Medullosa
from the Lower Coal Measures.” Ann. of Bot. vol. xxviii. pp. 252-
264.
Gorpon, 1912.—W. T. Gordon, * On Rhetinangium Arberi, a new Genus of
Cycadofilices from the Calciferous Sandstone Series.” Trans. Roy.
Soc. Edinburgh, vol. xlviii. part rv. pp. 813-825.
Jouxson, 1912.—T. Johnson, * Heterangium hibernicum, sp. nov., a Seed-
bearing /eterangium from County Cork." Scientific Proc. of Royal
Dublin Soc. vol. xiii. (n. s.) pp. 247-252.
Kunanr, 1908.—B. Kubart, ** Pflanzenversteinerungen enthaltende Knollen
aus dem Ostrau-Karwiner Kohlenbecken." Sitzungsber. d. K. Akad.
d. Wiss. in Wien, Math.-naturw. Klasse, Bd. cxvii. pp. 1-6 (57a
518).
Kunanr, 1911.— * Corda's Sphaerosiderite aus dem Steinkohlenbecken
Radnitz-Braz in Böhmen.” Jbid. Bd. cxx. pp. 1-14 (1035-1048).
Kupart, 1914.—* Über die Cycadofilicineen Heterangium und Lyginodendron
aus dem Ostrauer Kohlenbecken." Oesterreichische Bot. Zeitschrift,
Jahrg. 1914, pp- 1-11 (8-19).
RENAULT, 1869.—B. Renault, “ Étude de quelques végétaux silicifiés des
Environs d'Autun." Ann. des Sci. Nat. (Bot.) sér. 5, t. xii. pp. 161-
190.
RENAULT, 1879.—* Structure comparée de quelques Tiges de la Flore car-
bonifére.” Nouvelles Archives du Muséum, 2e sér. ii. pp. 213-348.
HENAULT, 1896.—* Bassin Houiller et Permien d'Autun et d'Épinae."
Flore Fossile, ii. (with Atlas, 1893). Paris.
Scorr, 1899.—D. H. Scott, “On the Structure and Affinities of Fossil Plants
from the Paleozoic Rocks. III. On Medullosa anglica, a new Repre-
sentative of the Cycadofilices.” Phil. Trans. Roy. Soc. B, vol. 191.
pp. 81-126.
102 DR. D. H. SCOTT ON THE HETERANGIUMS
Scorr, 1900.— Studies in Fossil Botany (1st edition). London.
Scorr, 1906.—* On Sutelifia insignis, a new Type of Medullosez from the
Lower Coal Measures." Trans. Linn. Soc. London, 2nd ser., Bot.
vol. vii. pp. 45-68.
Scorr, 1909.—Studies in Fossil Botany (2nd edition), vol. ii. London.
Scorr, 1914.— * On Medullosa pusilla.” Proc. Roy. Soc. B, vol. 87.
pp. 221-228.
Scorr & JEFFREY, 1914.—D. H. Scott & E. C. Jeffrey, “On Fossil Plants,
showing Structure, from the Base of the Waverley Shale of
Kentucky." Phil. Trans. Roy. Soc. B, vol. 205. pp. 315-373.
SEWARD, 1899.—A. C. Seward, * Notes on the Binney Collection of Coal
Measure Plants, Part IT. Megalozylon, gen. nov.” Proc. Cambridge
Phil. Soc. vol. x. part 11. pp. 158-174.
SoLws-LaAuBACH, 1887.—H. Graf zu Solms-Laubach, Einleitung in die
Paliophytologie. Leipzig.
Sonws-LaunAcH, 1896.—‘‘ Ueber die seinerzeit von Unger beschriebenen
strukturbietenden Pflanzenreste, ete.” Abhandl. d. K. Preuss. geol.
Landesanstalt, Neue Folge, Heft 23, pp. 1-100.
WiLLIAMSON, 1873.—W. C. Williamson, * On the Organisation of the Fossil
Plants of the Coal Measures, Part IV. Dictyoaylon, Lyginodendron,
and /Teterangium.’”’ Phil. Trans. Roy. Soc. 1873, pp. 377—408.
Wiuiamson, 1887.— bid. Part XIII. Heterangium tiliwoides and Kaloaylon
Hookeri. Ibid. 1887, B, pp. 289-304.
WiurAmson, 1890.—J/bid. Part XVII. — bid. 1890, D, pp. 89-106.
WinLIAMSON & Scorr, 1895.—W. C. Williamson and D. H. Scott, * Further
Observations on the Organisation of the Fossil Plants of the Coal
Measures. Part III. Lyginodendron and Heterangium.” Phil. Trans.
Roye Soe. vol. 186. (1895) B, pp. 703-779.
ZALESSKY, 1911.—M. D. Zalessky, * Étude sur Anatomie du Dadozylon
Tehihatehejfi," Mém. du Comité Géologique, Nouv. Sér. Livr. 68,
pp. 18-29. St. Petersburg.
EXPLANATION OF THE PLATES.
PLATES 1 & 2.
From Photographs by Mr. W. Tams. They require to be examined with a
lens in most cases.
PLATE 1,
Phot. 1. Heterangium shorense. General transverse section of stem. Cf, Plate 3. fig. 1.
A. Three bundles in the leaf-base (one is missing).
D. Double leaf-trace still attached to the stelar wood.
X about 5. Slide S. 2793. `
OF THE BRITISH COAL MEASURES. 103
Photographs 2-8. Heterangium tilicoides.
Phot. 2. General transverse section, from specimen 1. Note the well-preserved cortex.
T. 1. Double leaf-trace, in connection with stelar wood.
T. 2. Next outer trace; only one bundle shown.
T. 3. One pair of bundles of outermost trace. T. 3!. The other pair, damaged.
X about 6. Slide, Manchester, R. 632.
Phot. 3. General transverse section, from specimen 2.
T. 1. Double trace, just leaving stele.
T. 2. Bundles of a second trace, far out in pericycle.
Lb., outer limit of large leaf-base in connection with stem.
X about 6. Slide S. 237.
Phot. 4. Leaf-base from Phot. 3, enlarged.
v.b., one pair of bundles in the leaf-base ; v.5.', the other pair, damaged; C, outer cortex
of leaf-base; se., sclerenchyma delimiting leaf-base laterally; p.c., portions of
pericycle of stem.
X about 11. Slide S. 237.
Phot. 5. Part of another transverse section of specimen 2, showing a leaf-trace, consisting
of two large bundles, starting from the stele.
£., primary; x., secondary xylem of stele; ph.*, secondary phloem of leaf-trace ;
p.c., pericycle.
X about 40. Slide, Manchester, R. 654.
Phot. 6, General transverse section from specimen 3.
r.x., root-base, connected with the xylem of the stem; 7., free root.
T. 2. Leaf-trace in pericycle.
X about 6. Slide, Manchester, R. 658.
Phot. 7. From the section next below the last.
r., root, imbedded in the cortex of the stem.
X about & Slide, Manchester, Q. 60.
Phot. 8. Genera] transverse section from specimen 4. Note that everything beyond the
pericycle is lost.
T. 1. Inner trace, of two bundles, just leaving the stele. :
T. 2. Outer trace, of two widely separated bundles, passing out from the pericycle.
Cf. Pl. 4. fig. 19.
X about 6. Slide S. 1323.
PLATE 2.
Photographs 9-15. Heterangium Lomaxii.
Phot. 9, Form 8. General transverse section from specimen 1. Lowest section.
v.b., the two widely separated bundles of the leaf-trace.
x 6}. Slide S. 447.
Phot. 10. Peripheral xylem-strands from Phot. 9, enlarged.
px., protoxylem of one of the strands; x., primary; 4.7, secondary xylem.
X about 40. Slide S. 447.
Phot. 11. Form B. General transverse section from specimen 1, Uppermost section.
v.b., the two bundles of the leaf-trace, here much more widely separated than iv
Phot. 9; r., adventitious root passing out between them.
x about 8, Slide W. 1915 A.
E
^
:
104 DR. D. H. SCOTT ON THE HETERANGIUMS
Phot. 12. Form 8. General transverse section from specimen 6.
T. 1. Double leaf-trace in the outer part of the pericycle.
T. 2. Another double trace, just entering the cortex.
x about 18. Slide, Univ. Coll., N. 3.
Phot. 18. Form 8. General transverse section from specimen 5.
ph?, secondary phloem; Ay., thin-walled hypoderma.
T. 1. Inner leaf-trace, of two separate bundles, just leaving the stelar wood.
T. 2. Outer trace, two minute bundles in pericycle.
x about 10. Slide S. 625.
Phot. 14. Form * eylindricum." | General transverse section.
T, 1. Inner leaf-trace, consisting of two strands, just starting from the stelar wood.
T, 9. Outer trace of two double bundles in the cortex.
x about 10. Slide, Univ. Coll., N.
Phot. 15. Form æ. Part of a transverse section from specimen 1, showing a leaf-trace,
consisting of two double bundles, v.b. a, xylem of stele.
x about 35. Slide, Manchester, Q. 10.
PLATES 3 & 4.
From drawings by Mr. G. T. Gwilliam.
PLATE 3.
Figs. 1-7. Heterengium shorense.
Fig. 1. Transverse section of stem, showing stele with perieyele, and cortex with sclerotic
masses and Sparganum hypoderma. |
A., A., leaf-base with three bundles preserved (cf. fig. 2); D., B., next inner trace, with
four bundles in pairs; C., C., trace further in, consisting of two double bundles
(cf. fig. 6) ; D., innermost trace of two bundles.
x 7. Slide S. 2791.
Fig. 2. Trausverse section of leaf-base attached to stem. The four bunda (v.b.) in the
leaf-base are evident.
sc., sc., sclerenchyma delimiting leaf-base at the sides ; s.p., sclerotic plate.
x6. Slide S. 2787. :
Fig. 3. Detached petiole, containing eight bundles in four pairs.
s p., sclerotic plate.
x6. Slide S. 2787.
ig. 4. Leaf-trace consisting of two bundles (v.b.), starting from the stele.
zi
a
st., stelar wood.
x 96. Slide S. 2794.
Fig. 5. Part of primary xylem, showing three peripheral strands.
px., protoxylem of one of the strands.
x96. Slide S. 2792.
Fig. 6. Part of stele and half a leaf-trace.
x2, secondary xylem beginning to form on stele; pe., pericycle; v.b., double bundle,
. forming one-half of the trace C., C., shown in fig. 1.
x 46. Slide S. 2791.
Fig. 7. Part of pericycle and inner cortex.
pe., pericycle, with numerous sclereides ; ¢., inner cortex with a large sclerotic plate.
x 33. Slide S. 2791.
Scott.
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OF THE BRITISH COAL MEASURES. 105
Figs. 8-12. Heterangium tilieoides.
Fig. 8. Transverse section of stem from specimen 3, showing two leaf-traces.
v.b.', inner trace consisting of two bundles etaitiha. from stele; »0.2, outer trace,
consisting of two bundles in the pericycle, each SS signs of division ;
pc., pericycle ; e., portions of cortex.
x 9. Slide, Univ. Coll., N. 12.
Fig. 9. Part of stem from the same series, higher.up, showing the outer trace (v.2.), now in
the cortex, and consisting of two double bundles.
X., primary, x.?, secondary xylem of stele ; ph., phloem; pe., pericycle; c., cortex.
x 12. Slide, Manchester, Q. 58.
PLATE 4,
Fig. 10. Detached petiole, containing four distinct bundles. s.p., sclerotic plates.
x 18. Slide W. 1694,
Fig. 11. Transverse section of adventitious root, tetrarch or possibly pentarch, with secondary
wood ; outer cortex probably missing,
x 30. Slide, Univ. Coll., N. 12.
ES 12. Part of young stem, from specimen 4, showing a double leaf-trace.
v., primary, v., secondary xylem of stele; ph., phloem; pe., pericycle in which the
two a of the leaf-trace are embedded. Cf. Pl. 1. Phot. 8.
x 18. Slide S. 1323.
Figs. 13-18. Heterangium Lomavii.
Fig. 13. Form 8. ‘Transverse section of stem, specimen 2, showing stele with a small
amount of secondary wood, pericyele, and cortex.
V.b., v.b., the two bundles of the leaf-trace; the two protrusions of the wood between
them are connected with the bases of adventitious roots.
x 15. Slide S. 445.
Fig. 14. Form 8. Part of the xylem in radial section, passing through a peripheral strand.
Ppt., protoxylem ; æ., primary, .”, secondary xylem-elements.
x 183. Slide W. 1915. R.
Fig. 15. Ferm 8. Part of the cortex in radial section, showing the Sparganum hypoderma
on the outside, three sclerotic masses in the inner cortex, and pericycle below.
x 20. Slide, Manchester, R. 638.
Fig. 16, Form g, specimen 2. Transverse section of stem, showing the crushed stele, with
a possible double leaf-trace (v.b.), pericycle, Sd cortex; in the latter the large
and numerous sclerotic masses are conspicuous.
x 10. Slide S. 936.
Fig. 17. Form g, specimen 2a. Transverse section of a smaller stem. A double leaf-trace
is passing out.
v.b., one of the two strands in the outer cortex ; v./.!, the other strand displaced.
x 15. Slide S. 2306.
Fig. 18. Detached petiole, transverse.
v.b., the four bundles of the petiole ; r., an intrusive root of the “ Kaloxylon” type.
x 90. Slide S. 236.
Hed minimum.
Fig. 19. Transverse section of stem.
v.b., leaf-trace beginning to divide.
x32, Slide S. 1813.
LINN. JOURN.—BOTANY, VOL. XLIV. T
ee Re oak
k;
:
NP TTE
:
PROF. BOWER: HOOKER LECTURE, I9I7. 107
HOOKER LECTURE:
THURSDAY, ?ru JUNE, 1917.
By Proressor F. O. BOWER, F.R.S., F.L.S.
Havine for over thirty years occupied the Chair of Botany which Sir William
Hooker rendered famous by two decades of strenuous work,—in the University
of Glasgow where Sir Joseph Hooker graduated,—it seems apposite for me to
take as the subject of the Hooker Memorial Lecture the Natural Classification
of Plants. For no greater Systematists than the two Hookers existed during
their own period. And I may be pardoned if I draw my illustrations largely
from the group of the Filicales, since Sir William Hooker was the greatest
of descriptive Pteridologists.
The life-work of the Hookers, father and son, bridged a critical period in
the progress of Natural Classification, A vital difference of view separated
their outlook on living organisms. Sir William was the very embodiment
of the older régime, which laboured under the dogma of fixity of species.
Sir Joseph was the first botanical exponent of the belief that species are
mutable. Between them came Charles Darwin, who created that complete
change of outlook which followed on the publication of the ‘ Origin of
Species’ in 1859. The effect upon Natural Classification, brought about
by this revolution of thought, is the theme I shall endeavour to discuss.
It will not be necessary to refer in detail to the earlier history of Natural
Classification. We need do no more than take passing note of archaic
efforts. The recognition and naming of plants must have been initiated
by primitive man, for plant-names figure in all languages. More specific
designation and delineation were introduced as a consequence of the appli-
cation of plants to practical uses as foods, or medicines, or in the arts. The
mediæval Herbalists depicted many such forms ; but they treated them rather
as isolated phenomena, illustrating the ability of the Creator, than as having
any nearer causal relation. Their similarities led to their grouping into
families ; but the very likeness which the members of these families showed
was regarded as exhibiting ingenious variation upon a divine plan of con-
struction. And so, amid admiration tinged with reverence, there grew up
Systems of Classification, chiefly founded upon external similarity. Such
early schemes are described as “ artificial,” for they were based upon few
qualities, and those arbitrarily selected. Convenience was the motor impulse
LINN. JOURN.—BOTANY, VOL. XLIV, K
108 PROF. BOWER :
of their construction. The System of Linnæus was the culmination of such
efforts. But later the method became more “natural,” being based, like
that of De Jussieu, upon a wider area of diagnostic features. This change
led up to the period, about the opening of the 19th Century, when Sir William
Hooker began his life's work. He joined with vigour in the recognition,
delineation, and description of new forms, and the classification of them in
their natural affinities. Under his hands the Glasgow Garden became one
of the most notable in Europe. The port of Glasgow traded with every
quarter of the globe. This gave unusual facilities, and Sir William used
them to the full. The result was that amazing output of descriptive work
which marked his tenure of the Glasgow Chair, and it was continued at Kew
till the end of his life in 1865.
The five volumes of the ‘Species Filicum, condensed and erystallized
later into the ‘Synopsis Filicum,’ constituted his magnum opus. It may
be taken as a typical example of the best systematic works of the immediate |
Pre-Darwinian Period. It gives an insight into the method of its author,
and in some degree reflects his outlook. The work is analytie rather than
synthetie, though flashes of synthetic inspiration are scattered through its
pages. It describes, from personal observation and in meticulous detail,
a vast number of forms. The specific diagnoses and records of distribution
are a veritable mine of exact statement, which later systematists have freely
worked. But the author did not originate any new system. He adopted and
modified that of Presl's* Tentamen.' The chief merit of the book will always
rest upon what its very title conveys—that it gives a conspectus of all the
then known species of Ferns. The limits of the genera in which those species
were grouped gave rise to differences of opinion with other writers. Presl
had published his ‘Tentamen’ in 1836, somewhat earlier than Sir William
Hooker's beautifully illustrated work on the ‘Genera Filicum. Comparison
of the two books shows that there is a wide discrepancy in the number
of their genera. Where Presl recognized 132 genera, Hooker retained
only 75. This at once indicates a salient feature of his method. He
merged many genera, ranking them as sub-genera under more comprehensive
headings. One reason for this was his mistrust of anatomical data, of which
Presl made great use. But at least one other reason influenced him,—that
of convenience in diagnosis. An example will illustrate this point. In
placing Plagiogyria under Lomaria he objects to the use of its oblique
annulus as a diagnostic character, because of its requiring microscopic
examination, and the inconvenience “in a work whose main object is to
assist the tyro in the verification of genera and species.” Natural habit,
he remarks, is often a safer guide than minute microscopic characters.
And further, he states that “those Botanists who have showed themselves
peculiarly addicted to multiplying genera have not always taken Nature as
their guide, nor succeeded in eliciting a simple and tangible arrangement,”
HOOKER LECTURE, 1917. 109
It thus appears that convenience of application was at times admitted by
Sir William Hooker as a factor in his method.
Within the genus he grouped the species in a manner best illustrated in
large genera, such as Cyathea and Alsophila. The species are arranged
according to the complexity of branching of their leaves, from those which
are entire or simply pinnate, to those with more complex branchings. In
many genera the venation is used in diagnosis and grouping. A further
segregation was made according to geographical distribution. These and
many other characters are used ; but it appears that they are called in -
for purposes of diagnosis rather than those of synthesis. The object of the
author may have been in wish and intention to follow the dictates of
Nature. But along with this went in practice the laudable desire to make
the determination of species easy.
That the arrangement shown in the ‘Synopsis Filicum’ resulted in
something approaching to a natural grouping of the Filicales is undeniable.
Anyone, in the light of present knowledge, can criticise it in detail. But to
denounce it as thoroughly artificial (“ durchaus künstlich "), as Professor
von Goebel has done, is grievously to underestimate the merits of a great
systematic work *.
Nevertheless, it must be confessed that the arrangement of the Sub-
Orders of the Filicales in the ‘Synopsis’ was in some measure according to
tradition, or to chance. Circumstance rather than conviction appears to
have dictated it. The placing of the Gleicheniaces first coincides with the
arrangement in the ‘Tentamen’ of Presl. The sequence of other families
corresponds frequently, though not in detail, to that of the Prag Professor.
The relegation of the Osmundacez, Schizeeaceze, Marattiacee, and Ophio-
glossaceæ to the end of the book is probably a mere consequence of those
families having been omitted from the ‘Species Filicum,’ and they appear in
the ‘Synopsis’ as an addition (see Preface). Thus the arrangement of the
Sub-Orders seems to have been for Sir William a relatively trivial matter,
and it is certainly not according to Nature. Dut the intra-generic and
generie groupings are often those which will take permanent place as
knowledge increases, though this is certainly not the case invariably. We
conclude, then, that the permanent importance of the work centres in the
specifie diagnoses, rather than in the limitation or arrangement of the genera,
.or larger groups. In this respect it is characteristic of those systematic
works which were produced under the belief in the Constancy of Species.
The outlook of the Pre-Darwinian Systematist must have been highly
unsatisfactory to any intelligent man. On the one hand, he found the
* This verdict, expressed in ‘ Flora, 1896, p. 75, has been rebutted in detail elsewhere.
* Studies in the Morphology of Spore-producing Members," Phil. Trans. vol. 192 (1899),
p. 131.
K2
110 PROF. BOWER :
deeply ingrained belief inithe Constancy of Species. This doctrine, intro-
duced originally by Linnæus as a summation of his experience, was for a
century accepted by his followers as established truth. But, on the other
hand, there was a growing sense of the kinship of living organisms.
“Natural Affinity ” was instinctively recognized as a consequence of close
comparison. The instinct translated itself into methods of grouping together
such forms as have prominent features in common into genera and families.
Such relationship and consequent grouping was exemplified in all divisions
of the Vegetable Kingdom. If this was merely a reflection of the plan of
separate Creation of Constant Species, well might Elias Fries remark that
there was ** quoddam supernaturale” in the Natural System.
The fact is that the doctrines of independent creation, and of constancy
uf species are incompatible with the idea of affinity in the ordinary sense of
the word. This was doubtless in the minds of the Systematists of the early
19th Century. But many of them, like Sir William Hooker, were content to
expend their labour upon the recognition and record of those * affinities,”
without raising the inconvenient question of causality. Such difficulties
vanished with the collapse of the doctrine of Constancy of Species, and
the establishment of the theory of Evolution through Mutability. The
results already achieved in Natural Classification then found a new inter-
pretation. Affinity was held to mean some degree of relationship by
Descent. A Natural Classification, if correctly constructed, would then
be understood to visualise what remains of the evidences of Genetic History.
In fact, Classification and Phylesis should coincide.
The question may, however, be asked by those concerned especially in
Systematic Work, whether convenience is to be wholly ignored in the
systematic arrangements in use at the present day. It has been seen that
Sir William Hooker still admitted that convenience of application should
modify the method he used in the ‘Species Filicum.’ Therein he, as a
Pre-Darwinian, continued the practice which underlay, in more or less
degree, all the antecedent Artificial Systems. A very real object before
the early writers was readiness of identification of species believed to be
distinct, though in some way related. An insistent question for us now,
in the light of Evolutionary Belief, is whether or not convenience is
still to be allowed to modify the systematic exposition of phyletie con-
clusions, so as to render identification practically easy? The answer
should be a resolute negative. This is, indeed, the only answer for those
who clearly see in Systematic Arrangement a reflection of Genetic History.
But still some degree of convenience can be gained without the violation
of affinities demonstrated by more minute analysis. When such demon-
stration has been given, it should still be possible to construct analytical
keys, which would serve for ready systematic identification, without
involving all of the exacting observations of the laboratory. But in writing
HOOKER LEOTURE, 1917. 111
detailed Systematic Works the sole endeavour must be to arrange the material
so as to indicate phylesis.
It seems easy at the present day to grant this in theory, but it is difficult
indeed to carry it out consistently in practice. For it involves the whole
problem of Natural Relationships, which should be based upon the sum
of all knowledge relating to the organisms classified. It is then little
matter for surprise that the change of outlook, necessarily following on
the acceptance of Evolutionary Theory, reacted very slowly upon the
Classifications current during the second half of the 19th Century. We
may take as an example the Magnum Opus of Bentham and Hooker, the
‘Genera Plantarum,’ undoubtedly the greatest systematic work of the
period. The object of the book was to formulate, on the basis of personal
observation, the generic diagnoses of Flowering Plants. The grouping
of those Genera into Orders, and the arrangement of the Orders among
themselves were left substantially as in the systematic works current at
the time: for instance, in the * Prodromus of De Candolle. The old Sub-
Classes were retained. The Monochlamydez were kept apart as a separate
Sub-Class, and the Gymnosperms were spliced in between the Dicotyledons
and Monocotyledons. It does not follow, however, that this would be
the considered opinion of the author of the Monograph on Welwitschia.
It seems obvious that in a work, the value of which consisted in the
definition of genera, the further problem of the grouping of those genera
was not undertaken. The current groupings were retained so as not to
raise questions apart from the immediate task. However deeply Sir Joseph
Hooker’s mind was imbued with evolutionary belief, such views were not
obtruded in the text of the joint authors. Nevertheless, those who study
the last sections of the Ordinal Diagnoses of the ‘Genera Plantarum’ will
find indications of affinity far in advance of the Classification which the
authors had retained. But since the general scheme of arrangement
followed that in vogue in Pre-Evolutionary days, the ‘Genera Plantarum’
may be classed as technically belonging to that period. We thus see
that the effect of the Dogma of Constancy of Species, as reflected in
Systematic books, lasted long after it had ceased to hold the field. The
reason for this is to be found in the impossibility of remodelling the
broader lines of classification until time had familiarized the new aspect
of old facts ; and it was necessary to wait till the multitude of new facts
essential for full argument should have been acquired.
It might thus seem to the superficial observer as though Natural Classi-
fication had benefited little by the advent of Evolution. The charge
of ultra-conservatism, or even of inconsistency, might ignorantly be laid
against great men of science, such as Bentham and Hooker; though the
only possible ground for such a charge might actually be their unwillingness
to force a change prematurely. This is probably the chief reason why,
112 PROF. BOWER :
for long after the publication of the ‘Origin of Species, the Natural
Classification of Plants was still concerned rather with the distal twigs
than with the main branches of the Evolutionary Tree. Even now, the
question whether there was really a single trunk is unsettled, not only
for the Vegetable Kingdom at large, but also for its chief Divisions. Such
questions as the genetic relations of the Bryophytes and Pteridophytes : or
of the Gymnosperms and Angiosperms: of the inter-relations of Mosses
and Liverworts, or of the several phyla of Pteridophytes, are still matters
of speculation rather than of demonstration. Nevertheless, the assembly of
the members of each into genera, and groups of genera, is already well
advanced. The sequence of steps is naturally backwards from species and
genera to groups of genera, and orders: and thence to groups of orders.
But the argument proceeds from the distal to the proximal with ever
increasing uncertainty. Natural History is in fact like any other history—
a cult which is liable to lose its precision as it extends backwards into the
mists of the past.
In 1909 the sixth edition of Engler's ‘Syllabus of the Families of
Plants’ was published. It contains on its opening pages a statement of
the “Principles of Systematic Arrangement." This, though not by any
means exhaustive, may yet be taken as an Index of the point of view
entertained exactly fifty years after the appearance of the ‘Origin of
Species.’ It is true that it relates almost entirely to Flowering Plants,
as is natural since Professor Engler has been chiefly engaged upon them.
Using freely the observational results acquired by such predecessors as
Braun, Eichler, Bentham, and Hooker, he has proposed extensive re-
arrangements of the Families of Flowering Plants, in accordance with
the phyletie views set forth in his Introduction to the Syllabus. Apart
from those statements which have their special reference to Flowering
Plants, he enunciates among his Principles some which are of general
application. For instance, the facts of Ontogeny are held to be of the
first importance, as reflecting the probable course of Descent. But it is
pointed out that the facts of progress of individual organs from a phyletically
early state to the advanced are not usually available. It is recognized that
a character of value for comparison in one circle of affinity may be valueless
in another. On the other hand, those characters which remain constant in a
phylum are of special value in its comparative treatment. Insistently the
question will arise whether parts relatively simple in character are actually
primitive, or the result of reduction. The value of anatomical data is freely
acknowledged ; but those characters which are not in direct relation to
external conditions are to be estimated more highly than those referable
to adaptation. Combinations of progression in characters independent of
one another are recognized as specially important. Finally, the facts
HOOKER LECTURE, 1917. 113
of Paleontology are held as trustworthy, so far as they go. These are
among the principles which may be observed in the quest of a Natural,
that is essentially a Phyletie Classification.
Those engaged in such enquiry will assent to the soundness of Pro-
fessor Engler's Principles, which are indeed substantially those which were
already in general use. But the feeling most prominent in the mind, after
reading his statement of them, will be the hopelessness of the full realization
of the quest under present conditions, in the case of Flowering Plants. The
weakness of the evidence is specially marked in respect of the two lines
which should be most prominent: viz. Anatomy, and the Palsontologica)
record. Vascular Anatomy in Phanerogams has been robbed of its chief
phyletie interest, as Dr. D. H. Scott has indieated, by the disappearance
of the old centripetal wood, and the substitution of the new centrifugal
wood, which is secondary botl in the individual and in the race. Only
vestiges of the old wood remain in some few Gymnosperms. In the
Angiosperms it is absent. Accordingly the study of vascular anatomy
in them relates to relatively recent developments. The ancient structural
documents have been irretrievably lost.
Similarly, in the fossil record the documents are wanting for the phylesis
of Flowering Plants. The outburst of the Angiosperms in the Mesozoic
Period has left, so far as we yet know, little trace in the form of fossils with
structure. Moreover, such impressions as are found relate commonly to
leaves and stems, while flowers and fruits are wanting. The attention com-
manded by the record of Cretovarium by Dr. Marie Stopes (Ann. of Bot. xxiv.
p. 679) depended greatly on the rarity of the occurrence of such remains,
for it harmonized readily enough with general anticipation. Three other
circumstances affect the solution of the phyletie problem of the Angiosperms
adversely. The first is the completeness of differentiation of the vegetative
and propagative regions, whieh have diverged under adaptive modification
each along its own line. The second is the directness of adaptation of their
vegetative system to the environment, which obliterates the archaic, and
tends rather to present the recent features. The third is the relatively dead
level which these plants have reached in the details of their propagative
process. The depressing uniformity of the pollen-sac and of the ovule
in Angiosperms leaves little scope for comparative treatment in organs
which might otherwise have been full of hope for the morphologist. :
Before any general success can be expected in resolving the phylesis
of Flowering Plants, so as to establish their main lines of descent, new
evidence will be necessary. There will have to be an extension of the
eriteria of comparison. Already there are signs of this in the more accurate
comparison of details in the ovule, introduced by Van Tieghem. An
intensified search will also have to be made after specimens showing
114 PROF. BOWER :
structural detail from the Mesozoie Hocks. For it was at the period
when these were laid down that the immediate ancestors of the Angiosperms
flourished.
Methods of enquiry such as these applied to Flowering Plants are equally
applicable to other divisions of the Vegetable Kingdom. The cogency of
the results will vary with the variety and consecutiveness of the evidence.
The criteria of comparison will naturally be different. For instance, in the
Fungi dependence must be placed on the propagative, rather than the
vegetative system, while paleontological evidence is virtually absent. In
the Bryophytes, as also in the Algze, the vegetative system gives greater help
than in the Fungi: but again paleontological evidence is lacking. Recent
observations, however, point to the existence of sporogonium-like bodies even
from the Lower Devonian: thus indicating, on a basis of direct evidence,
a higher degree of antiquity than had previously been contemplated.
There remain the Pteridophyta. In them, and particularly in the Filicales,
evidence from various sources converges, so as to form a wide basis for their
phyletic arrangement. The results may indeed be held as more effective
here than in any other large group of Plants. We know from the fossil
record that organisms rightly ranked as Ferns date back far into the
Paleozoic Period. Such types, with modifications, may be consecutively
followed through successive horizons to the Present Day. Comparison shows
that some, though relatively few of the Ferns now living, correspond to the
archaic types of the Paleozoic ; while those of successively later horizons are
represented more and more freely in our present Flora. Finally, the great
mass of our living Ferns show characters which stamp them as distinctly
modern. Old though the Filical type undoubtedly is, we conclude from
inner comparison, as well as from the Fossil Record, that the fullness of its
development is that which we see in the majority of the living Ferns of the
Present Day. It is not a type which has stood still, but one which has con-
sistently advanced: and the advance has been as definite as that of the
Flowering Plants, but along quite different lines. Thus the Filicales offer
a singularly instructive field for the application of a phyletic method, so as
to elicit a really Natural Classification.
The basis upon which conclusions as to the evolutionary sequence of such
a group as the Filicales are to be arrived at, is at root that of the Natural
System of Classification, as commonly practised. It depends upon the
recognition not of one character or of two, arbitrarily selected as suitable
for ready use : but of as many characters as possible, which shall collectively
serve as criteria for comparison. In respect of each of these, variation will
be found as we pass from type to type. Such variations must be seriated,
and it will usually appear that they fall between two extremes. The question
then arises of the relative age of these extreme types. They may, of course,
have resulted from divergence from some middle type, and the enquirer
HOOKER LECTURE, I9I7. j 115
should always be prepared for this. But usually they bear the relation of
one extreme being relatively primitive, and the other relatively advanced.
Such a conclusion, based on comparison, should whenever possible be
checked by reference to the Fossil Record. Thus with a high degree of
certainty that which is archaic may be distinguished from the more modern
type, in respect of the character in question.
A good illustration of the effect of the Paleontological check is found in
the Filicales in respect of the type of sporangium. Von Goebel, in 1881
(Bot. Zeit. p. 717), distinguished the more massive types of sporangium seen
in the Ophioglossacez and Marattiacee as Eusporangiate ; while the more
delicate types characteristic of the Polypodiacew were styled Lepto-
sporangiate. Both are Filical types. Do they stand to one another as
relatively primitive, and relatively advanced ? If so, which was the earlier ?
I had written in 1889 (Ann. of Bot. iii. p. 305) a paper on * The Com-
parative Examination of the Meristems of Ferns as a phylogenetic study.”
All their meristems were shown to exhibit a parallelism with the sporangia
in point of complexity. So that the difference between the Eusporangiate
and the Leptosporangiate is really a difference of organization of the whole
plant. Influenced by the general opinion of the time,—itself based on
the assumed affinity of the Hymenophyllacez to the Mosses,—I then held
the simpler Leptosporangiate type of organization to be the more primitive.
But here came in the value of the Palzontological check. Stimulated by a
paper of Professor Campbell, as cogent as it is brief (Bot. Gaz. vol. xv.
Jan. 1890), the question was re-examined in the light of the fossil evidence.
The virtual absence of Leptosporangiate Ferns from the Paleozoic, and the
prevalence of the Eusporangiates at that period led to the inversion of
the series (Ann. of Bot. vol. v. 1891, p. 109)—a position now generally
accepted. Sporangial structure, which is an index of a bulky organization
and complex segmentation of all the parts in certain types, and of a less
bulky and complex construction in others, may accordingly be taken as a
criterion. As exemplified by Fern sporangia, the more bulky Eusporangiate
type is the more primitive, the less bulky Leptosporangiate type is relatively
advanced.
How far this will serve as a real index of their general organization is
shown by the comparison between the sporangia and antheridia of the same
plants, first instituted by von Goebel. It is found that where the sporangia
are large and thick-stalked, or even sunken as in Ophioglossum, the antheridia
are relatively large and are sunken too ; while in the Leptosporangiates, with
their smaller sporangia and long stalks, the antheridia are also stalked and
relatively small. The comparison even extends, though not with numerical
accuracy, to the numbers respectively of spores and of sperms. Where the
number of spores per sporangium is large, as in the Eusporangiate Ferns,
the spermatozoids are also very numerous in each antheridium: in the
116 PROF. BOWER :
Leptosporan giates, where the spore-numbers are relatively small, the number
of sperms is smaller also. Such facts give confidence in the value of this
. criterion, indicating that the sporangium is a real index of relative com-
plexity of organization, which is thus seen to extend to the gametophyte
generation,
In similar ways other criteria have been established. Each has been
checked by comparison, and as far as possible according to the fossil
evidence. The most important may be tabulated as follows for the Filicales,
i.-xiii. relating to the sporophyte, xiv.-xvii. to the gametophyte :—
i. External form, the upright radial shoot being relatively primitive,
the prone derivative, but with frequent reversions to the upright.
ii. Cellular segmentation, the more complex being primitive, and the
less complex derivative.
iii. Dermal appendages, simple hairs being primitive, and scales de-
rivative.
iv. Stelar structure, the protostele being primitive, and successively
the medullated protostele, the Lindsaya-type, the solenostele, and
the dictyostele being derivative. The progression may in certain
cases be followed in the ontogeny.
v. Leaf-trace, the coherent being primitive, and the divided trace
derivative. This may also be followed in the ontogeny.
vi. Venation, the open venation being primitive, and the reticulate
derivative. With this goes the “webbing” of the leaf to form
large laminar areas.
vii. Soral position, the marginal being the prevalent primitive type, and
the superficial frequently, or perhaps always derivative.
viii. Soral construction, the simple simultaneous sorus being primitive,
the gradate a frequent middle condition, and the mixed sorus
derivative.
ix. Indusial protections, none being present in primitive types: various
in form in intermediate types : but again indusia are absent in the
most advanced.
X. Sporangial structure, eusporangiate being primitive, and lepto-
sporangiate derivative.
xi. Mechanism of dehiscence: the annulus is indefinite in structure, with
median dehiscence in primitive types: oblique and continuous,
with lateral dehiscence in intermediate types: vertical and inter-
rupted, with lateral dehiscence in advanced types.
xii. Spore-output : large (i.e., typical numbers 128, 256, 512, 1024, &e.)
in primitive types: small (e.g., 64, 48, 32, 24, 16, 8, &c.) in
derivative types.
xii. Character of spore: the form is not a reliable character, except in
ERRATA IN No. 295.
Heterangiums of the British Coal Measures :—
Page 75. Foot-note, line 4, at end, for 19 read 13.
5» 8. be Bd GE AME for N. 19 read N. 13.
„ 104. Line 13, at end, after N, add 13.
SSN a oo ae, csi T TEES
HOOKER LECTURE, 1917. 117
near circles of affinity. The perispore is absent in most primitive
types ; but it is present in many derivative types.
xiv. Form of prothallus : relatively massive in primitive types, relatively
delicate in derivative types. But the character is unreliable.
xv. Sexual organs: sunken in primitive types, projecting in derivative
types.
xvi. Number of spermatocytes in each antheridium : large in primitive
types, smaller in derivative types.
xvii. Embryology : with suspensor present in a few primitive types, but
mostly without. The embryo relatively massive with axis vertical
in primitive types, more delicate with axis prone in derivative
ty pes.
These are the most important criteria for the phyletie seriation of the
Filicales. Many others are possible, and those stated raise many side-issues.
Their value individually is undeniable; but it is vastly enhanced by the fact
that the progression which each criterion shows in its own individual feature
runs parallel, as a rule, with progressions in the features of other criteria.
For example, Eusporangiate Ferns have a relatively complex cellular seg-
mentation, hairs as dermal appendages, usually an open venation, a simple
type of sorus, without indusium, massive sporangia, deeply-seated sexual
organs, and an upright embryo, sometimes with a suspensor. The more
advanced Leptosporangiate Ferns have a relatively simple cellular con-
stitution, scales as dermal appendages, frequently a reticulate venation,
gradate or mixed sorus, often indusiate, small sporangia, exposed sexual
organs, and a prone embryo with no suspensor. Such parallelisms of
progression in a plurality of criteria may also be traced more or less
clearly, and with a high degree of constancy, within nearer circles of
affinity.
On the other hand, occasional exceptions occur. Thus Cheiropleuria,
which is Leptosporangiate, has a protostelie stem with dermal hairs; but
its leaf shows an advanced type of reticulation, and an Acrostichoid sorus.
The axis in this case has retained its archaic characters, while the leaf has
advanced. But the converse is seen in the Marattiaceæ; for in them the
typical eusporangiate sorus is retained while the vascular system of the
stock is broken up into a complex plexus of reticulated strands. Here
the sporophyll remains archaic, while the axis has structurally advanced. A
somewhat parallel case is seen in Ophtoglossum (Cheiroglossa) palmatum,
where also the stock is swollen and sappy, and the stele is disrupted into
numerous strands. Such exceptional cases do not negative the method ; but
they serve to remind us that it is Organic Nature, with all its resources of
adaptation, that is being dealt with, not a mere mechanism designed after
fixed rules.
we e RS
118 PROF. BOWER :
This is neither the time, nor the place to describe in detail the results
whieh have followed from this more searching application of the Natural
Method of Classification to the Filicales. It must suffice to say that in
certain features they coincide with the results of the older Systematists,
which were based largely on external features. This is a high testimony to
the acuteness of their perception of affinity. But it applies rather to the
genera and species than to the larger groups. We have seen how assumed
affinities or mere circumstance appear to have determined the arrange-
ment of these in the ‘Synopsis Filicum.’ There is little evidence of any
better method in the arrangement in Engler’s ‘ Natiirlichen Pflanzen-
familien’? The Hymenophyllacese are there placed first, and then in
succession the Cyatheacee, Polypodiaces, Parkeriacew, Matoniaces,
Gleieheniaces, Schizæaceæ, and Osmundacee ; while the Hydropterides
are spliced in between these and the Marattiaceæ and Ophioglossacex. It
is difficult at first sight to trace any method in such a disposition. It may,
however, be seen to coincide in the leading features with the views of Prantl.
For he regarded the Hymenophyllacee as the source of the Lepto-
sporangiates and the Schizæaceæ as the source of the Eusporangiates,
these two main phyla being distinct. The arrangement is better in Christ's
* Farnkrüuter'; for the Eusporangiates, though placed last, are at least
in near relation to other Simplices. But it would be difficult to justify
phyletically the juxtaposition of the Hymenophyllacez, still placed first
of all, with the Acrosticheze, which immediately follow them. The plain
fact is, that up to the end of the 19th Century there was little attempt
at a definite method in the disposition of the main groups of Ferns in
the Systematic Works. The larger groups were still treated as though
they represented types isolated from one another in their Descent. It is
true that Prantl (Arb. Kónigl. Bot. Garten zu Breslau, 1892) represented
by a graphic figure his conception of the phyletic relations of the main
groups. But that figure shows that he believed his Osmundales (which
included the Eusporangiates, together with the Schizwacex, Gleicheniacez,
and Osmundacee) to be phyletically distinct from his Pteridales (which
ineluded all other Leptosporangiates). The one he traces from the
Schizæaceæ ; the other from the Hymenophyllaces, which his figure
suggests as having originated from some common but unknown ancestor.
This view is an improvement on the haphazard methods that preceded it.
But it breaks the continuity of those lines of. descent which are now rapidly
assuming clearer definition.
Two channels of recent investigation have materially helped towards that
clearer presentment : viz., the pursuit of vascular anatomy in Ferns, living
and fossil ; and the study of those types which had suftered vicissitudes of
classification, as shown by the richness of their synonymy. To the former
the main contributions have been those of Russow, Poirault, the Bertrands
father and son, Gordon, Boodle, Tansley, Kidston, Jeffrey, Gwynne-
OO Gn, Se a CNN.
HOOKER LECTURE, 1917. 119
Vaughan, and Lang. These have filled in many of the links in the chain
of evidence. They tend to demonstrate a consistent progression in stelar
structure of the primitive Ferns, fossil and living. It is found to accord
generally with the characters of their fructification, and with stratigraphical
sequence. It is true that on purely anatomical grounds, and without due
reference to the phyletie position of the Ferns compared, or the cogency
of their stratigraphical sequence, objection had been taken by the Harvard
School to certain well-founded conclusions. But this serves mainly as
a warning against reasoning based, as theirs has been, upon a single line
of evidence, pursued without due consideration of others.
The second method of investigation is by the enquiry into those Ferns
which bear many synonyms, and have obviously been difficult of classi-
fication under less searching methods. They now stand revealed as missing
links in the phyletic chain. Their being such explains the difficulty they
had presented to the early Systematists. As examples, Lophosoria, Metarya,
and Cheiropleuria may be quoted. Each of them had been merged by
Sir William Hooker, in consequence of his aversion to anatomical evidence,
into other large comprehensive genera. But they have since been re-
established as substantive genera. Lophosoria links the Gleicheniaces with
the Cyatheæ ; Metaxya helps to connect the Gleicheniacee with Elapho-
glossum ; while Cheiropleuria indicates how, from a type like the Dipterids,
a number of later forms may have sprung—and in particular that strange
genus, Platycerium. Another most important connecting type is Lowsoma,
and probably also the little-known Costa-Rican Fern, Loxsomopsis, which
indicate a transition from the Schizæaceæ to the Dicksonioid Ferns. But
we need not here attempt to exhaust the list of such connecting types. It
is merely intended to indicate how their recognition serves to link up those
sequences which form the lower branches of a phyletic system of the
Filicales.
An intensive study of large genera is another line which has been pursued
with success. For instance, the examination of many species of Blechnum
has disclosed how that genus, springing from some Matteuccioid source,
formed its characteristic fusion-sorus. This then spread over the leaf-
surface to form the Acrostichoid developments of Brainea and Stenochlena :
or it broke up into short lengths, as in Woodwardia and Doodya ; or, if
these were displaced by surface-growth, the result appears in the cha-
racteristic features of Scolopendrium and Asplenium. These are quoted
as instances of the methods now being pursued in the phylesis of Ferns,
and of the results which have followed.
A consequence of the more firm establishment of a number of distinct
phyletic lines has been to show more distinctly than before the falseness of
certain old-established genera, practically convenient as these may have been.
Polypodium, Acrostichum, and Gymnogramme, in the old comprehensive
sense, are all doomed to dissolution, They are not genera in the phyletic
$
;
!
1
120 PROF. BOWER :
sense, but states or conditions, which appear to have been arrived at along
a plurality of lines of evolution. This had long ago been recognized in
the case of Phegopteris, which is Polypodioid merely by abortion of the
Dryopterid indusium: it is quite distinct racially from — Eupolypodium,
Niphobolus, or Phymatodes, in which there is reason to believe that no
indusium ever existed. It thus seems probable that the Polypodioid state
may have arrived along a plurality of distinct lines. The same is the
ease with the Acrostichoid state. All that now remains of the genus
Acrostichum, according to Christensen’s Index, are A. aureum and prea-
stantissimum. And these are now seen to be derived from Pteris. Other
Acrostichoid Ferns have arrived from some five other phyletie sources.
Though the matter is not yet worked out in detail, it may be confidently
asserted that the large congeries of Ferns styled Gymnogramme had also a
plural phyletic origin, some of the species being originally destitute of
indusium ; others, such as G. Pozoi, being destitute of indusium by abortion.
The species named is clearly a non-indusiate Asplenium, as is also the genus
Aspleniopsis Mett.
New attempts to group the Filicales phyletically have thus resulted in the
recognition of a number of lines, divergent, parallel, or convergent, all
starting from Eusporangiate sources. These were characterized by having a
simple sorus, consisting of few sporangia, or even of a single one. There is
reason to believe that the position of that sorus was in the first instance
marginal. That is its position in the Botryopterids, the Ophioglossaces, the
Schizæaceæ, and Osmunda, all very primitive types. In others, as in the
Marattiaceæ, the Gleicheniacew, and Todea, the position is superficial.
But in several lines of descent, notably among the derivatives from the
Schizæaceæ, a transition from the marginal to the superficial by gradual
steps can be traced. And it seems probable that such a transition is
accountable for the origin of a superficial position in the other types
also. But in them it was carried out at an earlier period in Descent.
We may designate the first collectively as the * Marginales," and the
latter as the ** Superficiales?” This may be recognized as a broad phyletie
distinction, dividing the later Filicales into two, for the most part easily
distinguished sequences. But our working hypothesis will be that this
distinction does not mark an absolute difference: it is only a difference
of degree in respect of the time of the transition of the sorus from the
margin to the surface. But since the soral position is as a rule constant
in the species or the individual, the distinction is reliable as a basis for
phyletie segregation.
Each of the two main phyla thus distinguished includes a number of
subsidiary phyla. Of the Superficiales the Marattiaceze probably ended
blind, except for their relation to the Cycadales. The Gleicheniacem
probably led on the one hand to the Cyatheoid and Nephrodioid Ferns,
and finally to their Polypodioid and Achrostichoid derivatives, such as
HOOKER LECTURE, 1917. 121
the Oak and Beech Ferns, or Polybotrya osmundacea. A side branch
from Matteuccia led to Blechnum, with Scolopendrium and Asplenium as
further derivatives : also to Acrostichoid types, such as Stenochlaena and
Brainea. A second related sequence starting from Matonia, led through
Dipterid types to the Acrostichoid state of Cheiropleuria, Gymnopteris,
and Platycerium. A third line is indicated by Metaxya and Syngramme,
leading to the Acrostichoid genus Elaphoglossum.
On the other hand, from the Schizæaceæ, which are the most important
central stock of the living Marginales, we may trace the Dicksonioid-
Davallioid Series, culminating in Polypodioid forms, such as P. punctatum.
A side branch indicated by Lindsaya, Pæsia, and Pteris, culminates in
Acrostichum aureum ; while a collateral line , probably leads from Mohria
and Cheilanthes to such types as Hemionitis, and to the fully Acrostichoid
state of Z'rismeria.
From such seriations the fact of parallel development, or Homoplasy,
as it has been well designated in one of the earliest papers of Lankester,
emerges clearly. It is seen not in one case, nor yet in a single feature,
but in many. A few examples may be quoted. The dendroid habit is
characteristic of the Cyatheacez and the Dicksonies, two families which
were merged by earlier writers, chiefly on the ground of habit. Both have
sprung from a creeping ancestry. But the Cyatheacem have superficial
sori and chaffy scales; the Dicksonieæ have marginal sori and dermal hairs.
They represent distinct phyletie lines, the former being of Gleichenioid
origin, the latter of Schizewoid descent. Yet they appear so similar in
habit that detailed examination is necessary to distinguish them. They
exemplify a parallel or homoplastie origin of the dendroid state. Again,
dermal appendages illustrate in many distinct sequences the progression
from simple hairs which are primitive, to scales which are advanced. The
anatomical progression from protostely, to solenostely and dietyostely, as
also to polycycly, is illustrated in a plurality of lines phyletically distinct.
Soral characters show parallel progressions in many ways. The origin of
the gradate and mixed conditions of the sori: the progressive reduction
of the spore-output: the swinging of the oblique annulus to the vertical
position: the change from median to lateral dehiscence: the protection
of the sorus by those heterogeneous growths called indusia, of which there
are some half-dozen distinct types: the loss of such indusia, giving the
“ Polypodioid ” state: the spread of the sori to give the “ Acrostichoid ”
condition :—these and many other progressive changes can be shown to
have originated along a plurality of phyletic lines ; and they have resulted
in some cases in so high a degree of similarity that the segregation of the
forms showing them according to descent is difficult, however certain it may
actually be.
Thus the faet becomes clear as we proceed along the lines of phyletic
progress demonstrated by the comparative study of the Filicales, that
d.
122 PROF. BOWER :
parallel development, and even convergence of characters are common
phenomena. They may be traced in respect of almost all of the characters,
as having been carried out in sequences that must have been phyletically
distinct. It is possible, and with some degree of probability, to correlate
certain of these changes with the external cireumstances. For instance, in
the Paleozoic Period forest shade was either absent or imperfect. The
robust constitution of the Eusporangiate Ferns suited such conditions well
enough. The great outburst of the more delicate Leptosporangiate Ferns
took place in the Mesozoic Period, and doubtless the advent then of broad-
leaved trees, with their more effective shade, permitted plants like these, of
less robust habit, to flourish. At the same time many of them, through their
self-protection by dermal scales and indusial growths, had become more
independent. A curious biological feature becomes, however, evident-in their
further advance. For along many separate but parallel lines their indusial
coverings have become aborted in the species of the present day, and the
sori fully exposed again. It is as though the organisms had themselves
become in some way more resistant to exposure. For such present-day plants
often occupy exposed stations. This is the case in our own Polypodium
alpestre, properly called by Newman Pseudathyrium ; also in the Oak Fern
and Beech Fern. All of these may be found on hill stations, though they
have exposed sori. This condition is due to the abortion of indusia, in the
first case of the type of Athyrium, in the latter of Dryopteris.
It is not only parallelism of development, however, but even convergence
which is frequently seen. A good example is provided by the genera
Cystopteris and Acrophorus, which have been notoriously difficult to place.
They are ranked sometimes with the Davallioid, sometimes with the
Nephrodioid Ferns, two quite distinct phyla. So far as their venation
or their sori go, they might belong to either. But their anatomy, and
the presence of the characteristic chaffy scales, proves them to be of
Nephrodioid affinity. Another case is that of Doryopteris, long ranked
with Pteris, with which its fusion-sorus is in close agreement. But it
was separated by Prantl on more general grounds, and ranked, probably
correctly, with Pellea and the Cheilanthinz. Other examples are the
Davallioid genera Nephrolepis and Oleandra, which curiously mimic the
Nephrodioid sorus, though of quite a different origin from it. Thus con-
vergence of characters is a recurrent feature in the Filicales. There is,
indeed, no group of highly organized plants which shows more frequently,
and indeed consistently, evidence of parallel or convergent progressions in
distinct phyletic lines ; while the parallelisms and convergences involve
a great variety of characters, both vegetative and propagative. I commend
such facts to evolutionary theorists as a pressing problem for them.
The question will be, what eauses have been at work to produce such
results ? They are usually set down to the selection of favourable diver-
gences from type out of inheritable variations, or mutations, produced
VP TUNIS TT TNTCHNPS TT NEN
ME.
HOOKER LECTURE, 1917. 123 y
at random. But the prevalence of parallel development and convergence
suggests that they are not produced at random. Possibly such progressions
may be directed by some internal or physiological necessity. They raise,
however, in my mind very forcibly the question whether or not these
changes are promoted, or actually determined in their direction, or their
number, or their quality, in some way by the external conditions. I am
not aware of any facts which would raise this beyond the level of reasonable
suggestion, or probability. It is far from being proved; but it is equally far
from being disproved. It is in fact an open question. Until the contrary
is proved it would, in my opinion, be wiser to entertain as a working hvpo-
thesis some such view as that suggested than positively to deny it. The
impress of external circumstance cannot properly be ruled out in the genesis
of inheritable characters simply because up to the present time no definite
case of inheritance of observable characters acquired in the individual life-
time has been demonstrated. Already evidence is available from the side of
Zoology which, though it may not yet amount to demonstration, makes the
negation of inheritance of acquired characters perilous. A single positive
observation may at a stroke upset the whole negative position. Moreover,
the prevalence of parallel and convergent characters has made that position
suspect to many of those who pursue Morphology, whether of the animal or
of the vegetable kingdom. More especially I have found that this doubt is
entertained by those who have lived in the atmosphere of experiment and
observation found in large Botanic Gardens.
Plants would seem to be particularly favourable subjects for observation
in testing this question. The early segregation of the germ-cells in the
animal body was a fact which weighed greatly with Weismann in his
negation of the inheritance of acquired characters. But in Plants that
early segregation does not take place. In them the tissues, undifferentiated
as somatic and germ-cells, are for long exposed to the conditions under
which vegetation is carried out, before the germ-cells are specialized. When
this circumstance is given its full weight, such results of comparison as we
have been discussing seem to me more intelligible if related causally in some
way with the external conditions under which evolution proceeds, than if that
be denied.
There are various ways of approaching the problem, such as the com-
parative, the experimental, and the line of Mendelian analysis. Those who
approach the facts by the last should keep an observant eye also on the
methods and the conclusions of those who use a different method of enquiry.
More particularly they should watch with sympathetic interest those who
pursue the morphological avenue of approach to final truth: and especially
when these observers are by pursuit of that avenue led to conclusions
divergent from their own. Many Morphologists have found themselves
unable to accept that general application of the theory of “inhibiting
LINN. JOURN.—BOTANY, VOL. XLIV, d
124 PROF. BOWER: HOOKER LECTURE, I917.
elements ” put forward, as accounting for progressive evolution, by Bateson
in his Presidential address to the British Association in Australia. To me
the process of Mendelian Segregation appears to be nothing more than a
distributing agency. It constructs nothing: nor does it originate new
morphological characters. We have to look elsewhere than to Mendelian
Segregation for the origin of those structural innovations upon which
progressive Evolution depends. The chief interest of Evolution lies in
this initiation of new structural characters, rather than in the distributing
agency to which they are subject. The obvious relation of their cumulative
result to the environment, which we recognize as adaptation, carries with it
homoplasy, or even convergent development in lines phyletically distinct.
This is seen in high degree in the Filicales. It suggests, as the most
probable interpretation of the facts, that some causal connection exists
between the environment and those innovations which are the mark of
structural progress.
It has been impossible in this Lecture to do more than to give a slight
sketch of the methods recently employed in elucidating the phylesis of the
Filicales. Any more explicit statement of the results arrived at must be
reserved for some other occasion. Meanwhile such results have been! used
as an illustration of the modern method of Natural Classification. I hope
what has been said may suffice to show that real progress is being made.
Such measure of success in natural grouping as has been achieved among
the Filicales may serve to stimulate like efforts directed to other groups of
Plants. But it is useless to expect success to be as ready to hand in
them as in the Filicales. These plants present an exceptionally favourable
opportunity, owing to the nature of the evidence available. The Angio-
sperms offer a still wider field ; but it suffers from deficiencies of evidence,
which have already been explained. Moreover, the variety and extent
of their special adaptations complicate the problem, and often tend to
disguise the natural affinities. Still, it is not a hopeless quest to do the
same for them. But what is urgently needed for success amid the multi-
plicity of their lines of progress is a widening of the bases of comparison.
There must be a recognition of new criteria. A revision of the relative
values of the old criteria will also be necessary. These innovations, together
with the more free discovery of fossils with structure from the Mesozoic
Rocks, would provide the lines along which opinion may be expected to
develop. It is with this hope that I have used the Filicales as an object-
lesson in this Address. For the method which is being successfully applied
to them may be confidently expected, sooner or later, to unravel even the
tangled skein of Angiospermie Descent. Eventually it may lead to some
more satisfactory, and assured arrangement of them according to Descent
than has yet been achieved.
TWO CRITICAL PLANTS OF THE GREEK FLORA. 125
Two Critieal Plants of the Greek Flora.
By C. C. Lacarra, F.L.S.
{Read 3rd May, 1917.]
(1) Tuymus Sisrnorpn, Benth. — T. lanceolatus, Sibth. § Sm. non Desf.
=T. heterotrichus, Griseb.
The above synonymy was proposed by Celakovsky in 1882, but as neither
he, nor any other author who quotes these names, except Bentham, ever
saw the unique specimen in herb. Sibthorp at Oxford, which lies before me
as I write, it may be useful to confirm Celakovsky’s opinion with the aid of
that specimen, on which alone Bentham, Gen. & Sp. Lab. p. 345 (April 1834),
based his Thymus Sibthorpii.
There is nothing with the specimen to indicate where it was collected by
Sibthorp. The ticket on the sheet only bears the words “ Thymus lanceolatus
Desfont. Atlant. t. 128” in the handwriting of Smith, who in Fl. Gr. Prodr.
p. 419 (1806) merely quotes “ T. lanceolatus Desf. In montosis Gracie.”
Bentham at first rightly distinguished this Thymus from T. lanceolatus, Dest.
of Algeria as being * foliis glabris, rigidis, nec ut in T. lanceolato molliter
villosis." There are indeed stronger reasons for the separation, the chief of
which I summarise as follows :—7T. Sibthorpii differt a T. lanceolato foliis
floralibus non dilatatis, spicastro graciliori, minus compacto, calyce iriplo
minore, omnino alienis, et corolla multo minore.
The Algerian specimens on which I rely for these distinctions are:
(1) Jamin, Pl. Alger. 1851, no. 201; (2) Bové, no. 254; (3) Munby from
Sidi Den Abbas, all of which can be seen in Herb. Kew.
In 1844 Grisebach, Spic. Fl. Rum. ii. p. 116, described very fully a plant
from Mount Athos to which he gave the name of T. heterotrichus, attempting
to distinguish it from T. Sibthorpii only because the latter has “folia latiora
glabra et corollam vix calycem excedentem." Itis, nevertheless, undoubtedly
identical with 7. Sibthorpw. Unfortunately, when Bentham returned to the
thymes in DeCandolle’s *Prodromus, he confused under the name of
T. lanceolatus, Desf. three totally distinct plants, viz.: the true 7. lanceo-
latus of Algeria, a Persian species 7. lancifolius, Celak., and his own
T. Sibthorpii, but at the same time he separated 7. heterotrichus, Griseb.,
although with the remark “ An idem ae T. lanceolatus?” ; see DU. Prodr.
xii. pp. 199 and 203 (1843). It is to be remembered that Grisebach never
saw Sibthorp’s specimen, nor had Bentham seen Grisebach’s.
Boissier, * Flora Orientalis, iv. p. 555 (1879), wrongly identified both
T. Sibthorpii and T. heterotrichus with T. Serpyllum var. Chaubardi (T. Chau-
bardi, Boiss. & Heldr. pl. exsice. 1851, no. 269 — 7. glabratus, Bory & Chaub.
126 MR. ©. C. LACAITA ON TWO
non Hoffmg. & Link = T. angustifolius var. Chaubardi, Boiss. & Heldr. Diagn.
ser. 2, iv p. 6 (1859) 2 T. Serpyllum var. Chaubardi, Boiss. Fl. Or. 1. c.),
which is the common representative of the Chamadrys forms of T. Serpyllum
all over Greece, and also occurs in profusion in the mountains of South-
Western Italy. But in a MS. note posthumously published in Fl. Or. Suppl.
p. 359 (1888) Boissier, in deference to Celakovsky s opinion, separates
T. Chaubardi, * T. Chamedryi nimis affinis,” from. T. heterotrichus, Griseb.,
identified with T. lanceolatus, Sm. and T. Sibthorpii, Benth., which is “ab
omnibus T. Serpylli formis plane diversus." For in the meanwhile
Celakovsky, the most clear-sighted observer in this genus and one not
disposed to overrate lesser variations, had put the matter right in Flora, xl.
(1882) pp. 146 and 172, on the strength of specimens collected on Mount
Athos by Janka and by Pichler, identifying T. heterotrichus with T. Sibthorpit,
whilst clearly distinguishing T. Chaubardi. He does not appear to have
seen Grisebach’s type, but the precise locality ensures Janka’s and Pichler's
specimens being identical with it. On the other hand, he had certainly not
seen Sibthorp’s specimen for which no precise locality is assigned, but it is
known that Sibthorp visited Mount Athos. His herbarium contains many
plants from that mountain. I have compared his type with Pichler’s Athos
plant, which I have examined in Herb. Kew and in an example generously
lent for inspection by the conservator of Herb. Boissier, and feel quite
certain of their identity. Pichler’s plant was collected in August 1873.
The tickets of the two Kew examples read (a) * Macedonia, in sylvis Montis
Athos, Aug. 1873,” (b) ** Macedonien, Auf Pergen (se. Bergen) am M. Athos
bei Charies. Aug. 1873,” and that of Herb. Boiss. “Th. Sibthorpii, no. 183,
Auf dem nordlichem Abhange des Monte Santo.” Janka’s specimens I have
not seen, but there is a plant in Herb. Boiss. from Constantinople, leg.
Coumary, under the name of 7. montanus, Waldst. & Kit., which is cer-
tainly T. Sibthorpit, though the leaves are rather broader and more obtuse.
Velenovsky, Fl. Bulg. p. 469 (1891) and in his Nachtrag of 1903, p. 15,
describes 7. heterotrichus, Griseb. at some length, asserting that it is plenti-
ful in Macedonia on the Konjovo Planina and Osgovska Planina, at Krapee
and above Rilo Selo; “planta in M. Atho a cl. Janka lecta eadem est ac
nostra.” In the Nachtrag he says “in Bulgaria valde frequens " and creates
a subspecies cinerascens, * planta eximia sed transit in typum,” for which he
quotes five Bulgarian localities. But he does not allude to the identity with
T. Sibthorpii, although his Flora was published nearly ten years after
Gelakovsky’s work. It is also surprising that Haláesy in his‘ Conspectus
Flore Grace,’ ii. p. 563 (1902) should have repeated the erroneous
identification of 7. Sibthorpii with T. Chaubardi ; on p. 965 he alludes to
T. heterotrichus, Griseb. as “species ab autoribus varie interpretata, mihi
ignota."
I have not seen specimens of T. heterotrichus from the habitats enumerated
CRITICAL PLANTS OF THE GREEK FLORA. 127
by Velenovsky, or from the Greek localities cited by Formanek in Ver-
handl. Brünn, 1896, p. 67 and 1897, p. 50, but several plants collected by
Haussknecht in Western Greece in 1885 (specimens in hb. Kew) and
distributed as T. Sibthorpii var. grandiflorus and var. subalpinus are certainly
not forms of T. Sibthorpii, perhaps not even of T. Chaubardi. All that
concerns the present argument is the identity of the Athos plant on which
Grisebach's species rests with Sibthorp's specimen. That identity being
firmly established cannot be shaken by possibly incorrect determinations of
plants from other regions.
It only remains to consider possible objections to this identification on the
ground of discrepancies in the descriptions or superficial differences in the
specimens. I will deal with those that occur to me.
(1) Pichler's specimens were gathered in August and are in full bloom ;
Sibthorp’s has almost ceased flowering. Consequently the lower leaves have
fallen, only those of tbe upper part of the stem remaining, and the plant
shows what Gelakovsky, before he knew Pichler’s specimens, described from
Janka’s “gathered long after flowering with dried-up racemes and fallen
bracts,” viz., “ in the leaf axils of the sterile stems as well as in the middle
of the flowering ones there are formed characteristic short sterile branches
with, at the end of the branch, compact bunches of leaves that are half as
long as the supporting leaves but of the same Ee " [n Pichler’s specimens
thas are not yet developed.
(2) Another consequence of late gathering is that the corollas appear
even smaller than when gathered by Pichler in full bloom. Bentham says
“corolla vix calycem excedens "—Grisebach, “corolla calyce sesquilongior ”
(his main reason for separating his species from Bentham's), but Velenovsky»
“corolla vix exserta.” In fact calyx and corolla together are very small—
half the size that those of 7. Chaubardi usually attain. As far as I can
measure on herbarium specimens the calyx, including teeth, is only 3-4 mm.
long and the corolla, though clearly “ exserta,” is less than “ sesquilongior.”
(3) The leaves both in Sibthorp’s and Pichler’s plants are flat, “plana” as
described by Grisebach, not “ margine revoluto” as stated by Velenovsky.
Pichler’s measure 20 by 6 mm.; Sibthorp’s 15 by 5mm. The middle and
upper stem-leaves are narrowly elliptic *, 7. e. tapering almost equally at
both ends, but the lower leaves are obovate or obovato-spathulate.
Grisebach calls them “ lineari-lanceolata versus basin attenuata" ; Velenovsky
says “ovato-oblongis, oblongisve et lineari-oblongis," which will cover a
multitude of sins, including those of Coumary’s specimen which are about
* This botanical use of the word elliptic is of course mathematically incorrect. The
figure bounded by two intersecting menisci is not an ellipse. Bentham describes these
leaves as *oblongo-lanceolata " which is certainly incorrect. An oblong has not curved
sides and a lance is broader towards the base than towards the point.
wo PA AN
128 MR. C. C. LACAITA ON TWO
.13 by ? mm. The only real discrepancy is in the * margine revolutis ” of
Velenovsky, and he was not deseribing Athos specimens.
(4) Bentham says * suffrutex semipedalis," but of the pieces on Sibthorp’s
sheet two measure just 8 inches (20:5 cm.) ; Pichler’s and Coumary's plants
10-22 inches without the base of the stem. Grisebach places T. heterotrichus
among his *frutieuli erecti vel ascendentes, rhizomate descendente," dis-
tinguished from * suffrutices, ramis primariis demum decumbentibus, flori-
feris adscendentibus." Indeed, the whole plant is obviously a much taller
and more erect plant than T. Chaubardi or any of our forms of T. ovatus
(T. Chamedrys).
(5) The rather stiffly hairy calyx is set, as in so many Mediterranean
thymes, with beautiful ruby glands, some of which also stud the outside of
the corolla. Hitherto no one seents to have noticed that a certain number
of similar glands are found on the upper part of the stem itself. These are
conspicuous in Pichler’s and Coumary's plants, but are very difficult to see
on Sibthorp’s. However, I have detected a few.
The Thymus to which this species seems more nearly related by its
remarkably small calyx and corolla is T. montanus, Waldst. & Kit., as figured
in Pl. Rar. Hung. t. 71, and represented Fl. Exsicc. Austr. Hung. no. 2142
and by the Slavonian specimens from Mt. Veliky Gradac, distributed by Dr.
S. Kupeok. his, however, has the leaves of. T. ovatus (=T. Chamadrys),
glabrous stems and ealyces hardly, if at all, glandular. But it would .
require a complete study of the thymes of the Balkan peninsula to establish
the precise affinities and the limits of distribution of Thymus Sibthorpit.
(2) CREPIS RUTILANS, sp. nov., e Sect. Barkhausia, Radix verticalis, nescio
an biennis seu annua. Caulis solitarius, erectus, 85-65 cm. altus, villis albis,
mollibus, tenuibus et pilis robustioribus rubro-purpureis, basi dilatato in-
sidentibus, preeditus. Folia pilosa, mollia, inferiora in exemplari (A)
simplicia, leviter repando-dentata, dentibus retrorsis, in (B) runcinato-
pinnatopartita segmento terminali maximo ; caulina media amplexicaulia,
oblonga, superiora e basi sagittata lanceolata. Pedunculi etiam ante
anthesin erecti, sub anthesi breves (1-2:5 cm.), in fructu elongati (1:5-3 em.),
pilis rubro-purpureis aliisque pallidis brevioribus, nonnullis glanduliferis,
crebre vestiti. Capitula mediocria, eis C. setose eequalia, illis. C. fatide
dimidio minora. Involueri phylla externa subadpressa, pilis longis rubro-
purpureis barbata, interiora sparse cano-tomentosa, dorso etiam in fructu vix
indurato. Achenia tenuia, omnia in rostrum tenuissimum, ipsis vix brevius,
abeuntia, pappo niveo involücrum vix superante.
Differt ab omnibus C. fætidæ varietatibus caule non a medio ramoso,
panicula corymbosa, alabastris erectis, capitulis minoribus, acheniis cum
pappo duplo brevioribus.
"i
T t
PR CHN
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CRITICAL PLANTS OF THE GREEK FLORA. $5199
A Crepidi setoso, cui structura affinior, indumento molli pilisque rubro-
purpureis.
Ceeteris Crepidibus in Grecia aut Italia notis omnino absimilis, insigni
charactere, pilis rutilis, statim dignoscitur.
The above account is based on the only two specimens of whose existence
Iamaware: (A) A plant in seed in John Ball’s herbarium, now at Kew,
left by him without any name but now bearing the words “ Crepis (Bark-
hausia) " in the hand of Mr. J. G. Baker. Dall's label reads “ Iter orientale
1877. Ex Insula Corcyra in maritimis. Mense Junio. J. Ball.” (B) A
plant, hardly yet in full bloom, in the herbarium of Mr. G. C. Druce,
collected by him at Monrepos, Corfu, in May 1914. ^
Itis with some hesitation that I have ventured to propose a new species
based on only two specimens, but I am encouraged to do so in the hope that
botanists who visit Corfu will look out for this remarkable plant and make
plentiful gatherings of it.
March 18th, 1917.
A GENETIC STUDY OF NORTH AMERICAN MELANTHACEJE. X341
A Systematic Study of the North American Melantbacere
from the Genetic Standpoint. By R. Ruaaius Gares, Ph.D., F.L.S.
(With Map (Plate 5) and 1 text-figure.)
[ Read 15th March, 1917.]
INTRODUCTION.
THE purpose of the present paper is a little different from that of any
previous publication, so far as the writer is aware. It aims to furnish the
applieation to a particular group of plants of a definite point of view
regarding the genesis of species. It may be said that every systematist has
a point of. view in his work, and this is doubtless true, the point of view
having been furnished largely by the general Darwinian background, which
assumes the gradual and infinitesimal differentiation of species.
The point of view of the present work is essentially different, in that it
assumes the variations which differentiate species not to have been universally
continuous and infinitesimal, but to have been definite and often discontinuous.
To the writer, one of the most important advances since Darwin has been in
the discovery of the definiteness * of variation. This knowledge is based
upon experimental results obtained largely through the work of De Vries,
Bateson, Baur, Nilsson-Ehle, and many others during the last fifteen years.
The writer has endeavoured to analyze this definiteness in the case of the
(Enothera mutations, and has reached the conclusion + that many different
though definite types of change are involved, each giving rise to a different
and characteristic mutation. The essential conceptions of variation thus
developed through long experience and experiment by the writer with the
(Enotheras are here applied to the specific and generic differentiation of a
group of Liliales which has not been made the subject of experiment. In
short, the method is to apply the mutation conceptions to the systematic
treatment of this Liliaceous group in order to determine in how far the
method is valid and useful. It cannot be expected that final conclusions will
be reached in the absence of experimental analysis, anl yet I believe the
method is sufficiently illuminating to justify its adoption and application to
other groups. On the other hand, it must obviously be used with caution,
and in such a way that speculation does not too far outrun verification by
means of cytological studies and breeding experiments. If, however, the
* By definiteness of variation I mean not necessarily orthogenetie variation, but marked
and clearly defined variations which may occur in any direction or in many directions
simultaneously. :
+ ‘The Mutation Factor in Evolution, p. 853. Macmillans, London, 1915.
LINN, JOURN.—BOTANY, VOL. XLIV. M
T1352 DR. R. R. GATES: A STUDY OF NORTH AMERICAN
- general conceptions of definiteness and discontinuity in variation are sound,
then the principles that arise from these conceptions should serve as a touch-
stone when applied to the study of the origin and relationships of living
species. The present work is published as an example of what may be
accomplished in this direction. It will doubtless be criticized from many
points of view, but it is hoped that as a first step it will not be entirely
without justification.
This work is the outcome of a study which began with several pairs of
species *, taken at random as regards their distribution, differences, and
probable manner of origin. The group Melanthaceze was selected on account
of its compactness and because it is far removed from the Onagracem. The
conceptions here applied to these families will doubiless apply equally well
to many others. That variations are often marked and definite, who will
now deny? It is with the hope of convincing systematic workers that these
conceptions are not only useful but necessary, if we are to reach an under-
standing of specific differences, that this publication is undertaken. The
definite conceptions of germinal variation or mutation, which are founded
on careful analytical, experimental, and cytological work, need to be incor-
porated into the biological thinking of the time, and this can only be done
by applying the results of experiment to groups where experiment has not
taken or cannot take place.
On the other hand, it soon becomes clear that, while many variations have
been sharp and definite, this is by no means universally the case. The con-
tinuous and indefinite variations which sometimes lead from species to species
are the difficult ones to analyze. It may be admitted that their evolutionary
significance is at present insufficiently analyzed and practically unknown,
despite the current assumption that * fluctuations” are not inherited and are
therefore not of evolutionary value. The present paper, while pointing out
frequent cases of such “ continuous variation," is equally concerned with the
cases of discontinuity. Many of the latter are obviously due to gaps caused
by extinction, but many more are undoubtedly the result of discontinuous or
definite variations. Is it, for instance, probable that a verticillate arrange-
ment of the leaves, as in certain species of Polygonatum, in the related family
Convallariaceze, arose gradually from an arrangement opposite, in pairs?
And, again, is it not more reasonable to assume that the “knee” in the
peduncle of Streptopus arose as a definite and marked variation, than that it
was slowly and gradually evolved? It appears that in many cases differences
which are now used as generic characters originated by a marked and definite
step. The argument that extinction accounts for their present distinctness is
certainly often excluded.
Moreover, the mere fact that related genera showing marked differences
* “On Pairs of Species,” Bot. Gazette, lxi. 1916, pp. 177 212, figs. 12.
MEMO a oin a aia
MELANTHACEÆ FROM THE GENETIC STANDPOINT. 133
in structural characteristies often co-exist side by side * indicates that these
differences were probably not of selective value, but they arose through what
Darwin called “spontaneous variation,’ and have been perpetuated by
heredity. It is thus believed that, while natural selection accounts for the
gradual development of adaptations through the accumulation of larger or
smaller variations, the mutation theory accounts for the origin of innumerable
generic and specific differences which were never of selective value to their
possessor. In the years when natural selection was the only accepted factor
of evolution, it always requiredalarge measure of credulity to believe that every
character of every organism had been at some time of particular value to its
possessor. The mutation theory renders unnecessary such an assumption,
and thus relieves natural selection of the necessity of explaining too much.
With these and similar conceptions in mind, the North American Melan-
thaceze have been studied systematically. The result is incomplete, because
some of these genera are still insufficiently known, and I have not had an
unlimited time to devote to the subject. It is hoped that the work will be
most useful for its suggestiveness. We know that many variations are
marked steps, and that such variations are inherited, while we as yet know
very little of the inheritance of continuous variations, and what is known is
largely negative in character. If systematists are to reflect the knowledge
of their time in the construction of their species, then obviously they need
to pay at least as great attention to discontinuous as to continuous variability,
our knowledge of the former being firmly based on experiment.
The analyses of specific and generic differences attempted in this paper
may appear speculative, but at least they are no more so than they would
be if they assumed variation to be continuous. They, moreover, have the
definite advantage of being based on analogy—sometimes direct analogy—
with known experimental results. It is hoped that the explanations here
suggested of various specific and generic relationships will lead to the further
study of many of these species both eytologically and experimentally, for this
is the only way in which the questions here propounded can be definitely
solved. It would be an attractive problem to make a comparative study of the
chromosomes throughout the group, and it is hoped that botanists will avail
themselves of this opportunity whenever they can obtain the necessary material.
In the systematic treatment of these groups it has been necessary to
describe several new species and varieties and to make several new com-
binations, though an effort has been made to circumscribe their number.
It is perhaps doubtful whether the segregation from Zigadenus of the genera
T'oxicoscordion, Anticlea, and particularly Oceanorus should be recognized,
though I have here given them the benefit of the doubt. Tt is easily
* Cf. Gates, R. R., “The Mutation Theory and the species-concept," in Amer. Nat.,
Oct. 1917.
; M 2
134 DR. R. R. GATES: A STUDY OF NORTH AMERICAN
possible to carry the process of subdivision of existing genera too far, and
by so doing to destroy the usefulness of the genus and impair its definiteness
as well, It seems certain that no new genera should ever be introduced in
the groups here under consideration, The indiscriminate creation of new
TRIANTHA
TOFIELDIA
METANARTHECIUM
Juncoioes
yx
JUNCUS P dba ELLA
XEROPHYLLUM
STENANTHIUM
AMIANTHIUM
ToxicOSCORDION
CHAMAELIRIUM
HELONIOPSIS SCHOENOC
| -— fe AULON
E HELONIAS
y, HEMIPHYLACUS
OocoNTOSTOMUM
MELANTHIUM
Oceanorus
HASTINGSIA
SCHOENOLIRION
ZIGADENUS
CHLOROGALUM
ORIGINAL ANCESTORS
OF MELANTHACEAE
Phylogeny of Melanthace:e.
generic names obscures relationships, and there is nothing to be said in its
favour.
In the citation of specimens in this paper it is to be assumed that all
specimens are from the Herbarium of the Missouri Botanical Garden, unless
otherwise specified. Iam greatly indebted to Dr. Geo. T. Moore, Director
of the Missouri Botanical Garden, for placing the facilities of the institution
MELANTHACEJE. FROM THE GENETIC STANDPOINT. 1535
at my disposal, and to Dr. J. M. Greenman for frequent consultations in
connection with the systematic work. Mr. C. H. Thompson has kindly
taken a number of photographs. At the University of California, through
the kindness of Dr. W. A. Setchell and Dr. H. M. Hall, I was able to
study the large collections of Californian Liliacez, and I am also indebted
to Dr. Setchell for a fine series of photographs of herbarium specimens.
1. TOFIELDIA, Huds.
1. TorrELDIA PALUSTRIS, Huds. Fl. Angl. ed. 2, 157 (1778); Delar. in
Redouté, Lil. v. t. 256 (1809) *.
T. pusilla, Willd. in Mag. Gesells. Naturf. Fr. ii. (1808) 27.
T. borealis, Wahlenb. Fl. Lapp. (1812) 89.
Narthecium boreale, Wahlenb. in Nov. Act. Holm. xxvi. (1803) 24.
N. alpinum, Michx. Fl. Bor. Am. i. (1803) 209.
N. pusillum, Michx. l. c.
Anthericum calyculatum, 3, Linn, Fl. Suec. ed. 2 (1755), 108.
Greenland and Labrador to Alaska, south to Quebec, Lake Superior, and
the Canadian Rockies. Also in Europe and Northern Asia.
2. 'l'OFIELDIA CALYCULATA, Wahlenb. Veg. Helv. (1813) 68.
Anthericum calyculatum, Linn. Sp. Pl. (1753) 311; Fl. Dan. (1766) t. 36 ; Linn, Fl.
Lapp. (1792) 106, t. 10. f. 3; Smith, Engl. Bot. (1799) t. 536.
Scheuchzeria Pseud- Asphodelus, Scop. Fl. Carn, i. (1772) 268.
Narthecium calyculatum, All. Fl. Pedem. ii, (1785) 165 ; Lam. Encyc. iv. (1823) t. 268.
Helonias borealis, Willd. Sp. Pl. ii. (1799) 274.
Hebelia allemanica, C. C. Gmel. Fl. Bad. ii. (1806) 118, t. 1.
H. collina, C. C. Gmel. l. c.
Europe.
3. TorrELDIA STENOPETALA, Smith, in Trans. Linn. Soc. xii. (1817) 243,
t. 8. f. 1.
This species was founded on specimens collected by Kalm in North
America, three of which are in the Linnean Herbarium. It has not been
recorded since, but it seems probable that the species is distinct and that
T. calyculata is exclusively European. Smith also described T. alpina as a
species of the mountains of southern Europe, which is distinguished from
T. palustris by being much larger in all its parts. At the same time he
united T. calyculata with T. palustris. Ascherson and Graebner f, on the
other hand, unite T. alpina, Smith, with T. calyculata (Linn.), Wahl., and
recognize also 7. palustris, Huds. The relationships of these four forms
can only be determined by comparison of European and American spe-
cimens. Sir James Smith states that T. stenopetala is nearest T. alpina, from
* This apparently represents T. alpina, Sm., which is larger than T. palustris, Huds.
+ Mitteleur. Fl. iii. (1905) 5.
136 DR. R. R. GATES : A STUDY OF NORTH AMERICAN
which it differs in having an inflorescence which forms a dense obtuse
cluster, 13" long; bracts lanceolate, often exceeding the pedicels; calyx
very broad and shallow, usually 3-cleft, petals eadiiels and acute, anthers
pointed, ovary ovato-lanceolate, styles twice as long as in 7. alpina. It
seems probable that further study will demonstrate the distinctness of this
North American species, and perhaps also of the European T. alpina.
4. TorrELDIA coccinea, Richardson, in Frankl. 1st Journ. (1823) 736;
Hook. & Arn. Bot. Beechey’s Voy. (1841) 130, t. 29 bis.
T. borealis, Cham. in Linnea, vi. (1831) 584.
Arctic America.
4a. TOFIELDIA COCCINEA, var. MAJOR, Hook., Fl. Bor. Am. ii. (1840) 179.
Mackenzie River (Richardson).
According to Hooker, this is easily distinguished (1) by its more flaccid
leaves and leafy scape, (2) by its sessile m deeply tinged with red,
(3) by its large bracts which form a deep involucre beneath the flower,
(4) by the singularly deflexed dark purple fruit. It appears quite worthy
of specific rank. The same form is recorded by Hooker from Siberia in
Herb. Pallas. i
Another variety, represented by two specimens collected by Richardson
on the Mackenzie River, is mentioned by Hooker as having longer pedicels
than the type of T. coccinea.
5. TOFIELDIA GLABRA, Nutt. Gen. Amer. i. (1818) 235; A. Gray, in Ann.
Lyc. N.Y. iv. (1837) 136.
T. glaberrima, MacBride, in Ell. Sketch, i. (1821) 424.
North Carolina and South Carolina.
The total number of deseribed species is about 32, over half of which are
contined to Asia. The genus bears many resemblances to Juncus, and it
may be supposed that the ancestors of Tofieldia gave rise to the Juncaceæ.
This seems to the writer to be more probable than to read the series in the
opposite direction, The genus Tofieldia contains more than 15 species in
the north temperate zone and 3 species (7. falcata, Pers., T. Moritziana,
H. Schultze, and T. Sehomburgkiana, Oliver) in the Andes of South
America, A number of species have been described from China and Japan
in recent years.
T. borealis, Wahlenb., of Lapland, Tyrol, Labrador, ete., is apparently
inseparable from 7. palustris, Huds., unless it be in the presence or absence
of pedicels to the flowers. T. coccinea differs from | T. borealis in having
more flaccid leaves of a duller colour and flowers sessile. T. glabra differs
markedly from the other, more boreal, species of Tofieldia in America. It
MELANTHACE/ FROM THE GENETIC STANDPOINT. Lon
occurs in North and South Carolina, and is tall rather than dwarfed, forming
in this respect a transition to the genus Triantha. But it has neither the
arrangement of the flowers in threes nor the scabrous pubescence on
the stem characteristic of Triantha.
2. TRIANTHA (Nutt.), Baker.
l. TRIANTHA RACEMosA (Walt.), Small, Fl. S.E. U.S. (1903) 249.
Melanthium racemosum, Walt. Fl. Car. (1788) 126.
Narthecium pubens, Michx. Fl. Bor. Am. i. (1803) 209.
N. scabrum, Rafin. in Desv, Journ. de Bot. iv. (1814) 273.
Tofieldia pubescens, Delar, in Redouté, Lil. vi. (1812) t. 324.
T. pubens, Hook. in Bot. Mag. (1841) t. 3859.
T. racemosa, Britton, Sterns, & Pogg, Prelim. Cat. N.Y. Plants (1888) 55; Ann. Rep.
N.J. State Mus. 1910 (1911) p. 337, pl. 33. f. 1.
In swamps, southern New Jersey to Florida and Louisiana.
2. TRIANTHA GLUTINOSA (Michx.), Baker, in Journ. Linn. Soc., Bot. xvii.
(1879) 490.
Narthecium glutinosum, Michx. Fl. Bor. Am. i. (1803) 210.
Tofieldia glutinosa, Pers, Syn. i. (1805) 399; Smith, in Trans. Linn. Soc. xii. (1818) 246,
t. 8. f. 2; Hook. Fl. Bor. Am. ii. (1840) 179, t. 191.
In bogs, Newfoundland to Alaska, Minnesota, Maine, in the southern
Alleghanies to N. Carolina and west to Ohio, Wyoming, and Oregon.
3. T 'RIANTHA INTERMEDIA (Rydb.), comb. nov.
Tofieldia glutinosa, Hook. Fl. Bor. Am. ii. (1838) 179, partim; S. Wats. Bot. Calif. ii.
(1880) 184.
T. intermedia, Rydb. in Bull. Torr. Bot. Club, xxvii. (1900) 528.
In bogs, Alaska to Saskatchewan, Montana, Wyoming, and Tulare Co.,
California.
4./ TRIANTHA OCCIDENTALIS (S. Wats.), comb. nov.
Tofieldia occidentalis, S. Wats. in Proc. Am. Acad. xiv. (1879) 283.
N. California (Mendocino Co.) to Washington and British Columbia, in
mountains.
5. TRIANTHA Japonica (Miquel) Baker, in Journ. Linn. Soc., Bot. xvii.
(1879) 490.
Tofieldia japonica, Miquel, Ann. Mus. Lugd.-Bat. iii. (1867) 201.
Japan, in swamps.
The genus Triantha, which contains the above five species, is properly
separated from Tojieldia on account of (1) the scabrous pubescence, (2) the
flowers in threes, and (3) the caudate seeds. The two western species,
T. occidentalis and T. intermedia, should be included in this genus, since
they have these three characters.
In this connection the case of Tofeldia glabra is of very much interest.
NET RS ASTE ONERE NR PEENE ET
138 DR. R. R. GATES : A STUDY OF NORTH AMERICAN
As already remarked, it agrees in habit with the species of Triantha,
particularly T. racemosa, but differs sharply in two characters: (1) the
absence of pubescence from the stem, (2) the flowers not arranged in threes.
Its distribution is restricted, and it seems reasonable to conjecture that it
may have originated from 7. racemosa by two mutations. At any rate,
it appears to differ from that species by what may very well be two unit
characters, one of them a “loss” character. Certain specimens of T. race-
mosa from Carolina depart from the type in having the flowers for the most
part not in threes but arranged as in 7. glabra. Some of these may perhaps
be hybrids between Tojieldia glabra and Triantha racemosa, or possibly
mutations from the latter. Breeding experiments with those two forms
could be profitably inaugurated, and a study of the forms occurring in North
and South Carolina should also be made.
Triantha racemosa, which is more southern and much less widespread
than T. glutinosa, differs from the latter mainly in (1) pubescence more
scabrous, (2) perianth becoming rigid, (3) capsule firm, dark-coloured, long-
beakel. Of the two western species, 7. intermedia is not very sharply
marked. It is distinguished from T. glutinosa by (1) having a short, more
or less globose raceme, not elongated as in the eastern species ; (2) having
longer pedicels and oblong subequal perianth-segments (4 mm. long).
T. occidentalis is somewhat more restricted in range than T. intermedia, from
which it is distinguished by its narrower, less connate bractlets, longer
pedicels, narrower sepals, larger capsules (8 mm. long), and longer ascending
beaks. "These are all quantitative differences, not sharply marked.
The third known species of Triantha, T. japonica, Baker, I have not seen.
It is perhaps related to 7. intermedia.
The three main differences between the closely related Tofieldia and
Triantha—namely, (1) the arrangement of the flowers in clusters of three
instead of singly on the stem, (2) the appearance of a characteristic rough
pubescence, (3) the caudate seeds—are such as might have resulted from
three mutations. But, if so, where are the forms exhibiting only a single one
of these changes, and why have only the forms containing all three been
preserved? There is perhaps no utility in any of these characters, but it
would appear that only forms having all three of these features, or lacking
them all, have survived. Yet to suppose the passage from Tojieldia to
e , à : ;
Triantha in a single step requires a much larger change than we know from
experience. These are but a few of the questions raised by a consideration
of the two genera.
3. PLEEA, Michaux.
PLEEA TENUIFOLIA, Michx. Fl. Bor. Am. i. (1803) 248, t. 25; Delar. in
Redonté, Lil. v. (1809) t. 248; Sims, in Bot. Mag. (1818) t. 1956.
In pine-land swamps, S. Carolina to Florida.
ee ELS ee o is ea
MELANTHACE FROM THE GENETIC STANDPOINT. — 139
This very interesting monotypic genus differs from 7riantha and Tofieldia
chiefly in the following points:—
Triantha (Nutt.), Baker. Pleea, Michx.
Bracts short. Bracts spathe-like.
Flowers subtended by 3 connate Bractlets wanting.
bractlets.
Stamens 6. Stamens 9-12.
Anthers rounded, erect. Anthers elongated, versatile.
Pleea is evidently isolated, and a considerable amount of extinction must
have occurred between it and its nearest relatives, Tofieldia, Triantha, and
Narthecium. Regarding the comparison with Triantha, it is more likely
that the bractlets subtending the flowers were suddenly lost by a negative
mutation than that they were gradually reduced until they finally dis-
appeared. The increase in size and change in shape of the bracts may
have occurred independently, as also the addition of one or two whorls of
stamens. The change from the short, subglobose, erect, terminal antlers of
Tofieldia to the long, narrow, versatile anthers of Pleea probably involved
several different germinal changes. Some species of Luzula have elongated
erect anthers, so the versatile character probably appeared through an
independent change in a related ancestral line.
`
4. NARTHECIUM, Moehring.
. NARTHECIUM OSSIFRAGUM, Huds. Fl. Angl. 145 ; Sm. in Engl. Bot. viii.
(1799) t. 535.
N. anthericoides, Hoppe, in Mert. & Koch, Deutschl. F1. ii. (1826), 559.
N. palustre, Bub. Fl. Pyren. iv. (1901) 169 *.
Anthericum ossifragum, Linn, Sp. Pl. (1758) 446; Fl. Dan. (1761) t. 42.
Abama ossifraga, DC. in Lam, Fl. Fr. ed. 2, iii. 171; DC. in Redouté, Lil. iv. (1808)
t. 218.
Europe.
—
. NARTHECIUM AMERICANUM, Ker, in Bot. Mag. (1812) t. 1505.
N. ossifragum, var. americanum, A. Gray, Man. ed. 5 (1867) 536. :
Abama americana, Morong, in Mem. Torr. Bot. Club, v. (1894) 109; Ann. Rep. NJ.
State Mus. 1910 (1911) 338, pl. 33. ff. 2-3; Bartonia, iv. (1911) 1, t. 1.
Pine barren swamps, southern New Jersey, and Delaware. `
This species is extremely local in its distribution. It was discovered by
Pursh at Quaker Bridge, New Jersey, about 1805. He sent plants to Kew,
from which the figure was made for the ‘Botanical Magazine. Witmer
Stone t gives an account of its present distribution. After the lapse of a
n2
* A full list of figures is given here.
+ Abama americana (Ker), Morong, in * Bartonia, iv. (1911) 1-5.
140 i DR. R. R. GATES: A STUDY OF NORTH AMERICAN
century since its original discovery, N. americanum is now (with one
exception) known only from fifteen localities, included in an area about
20 by 30 miles. It was always supposed to be restricted to the heart of the
New Jersey Pine Barrens, but specimens in Herb. Phila. Acad. Sci. were
collected by Mr. A. Common near Lewes, Delaware, on Aug. 1, 1895.
The plant is more restricted now than formerly, because it is killed by
cranberry culture.
3. NARTHECIUM CALIFORNICUM, Baker, in Journ. Linn. Soc., Bot. xv. (1876)
351.
N. ossifragum, var. occidentale, A. Gray, in Boland. Pl. Calif. (1870) 31.
Abama californica, Heller, Cat. N. Amer. Pl. (1898).
A. occidentalis, Heller, in Muhlenbergia, i. (1904) 47.
California, Mendocino Co., and northwards into Oregon, and in the
Sierras to the Yosemite National Park. The type came from Del Norte Co.,
California.
4. NARTHECIUM AsIATICUM, Maxim. in Bull. Acad. Pétersb. xi. (1867) 438.
Japan.
This genus is particularly interesting on account of its discontinuous
distribution, no one of the four species coming in contact with any of the
others. And yet, though widely separated geographically, the species are
closely similar. W. asiaticum appears to represent the oldest species, and is
found in Japan. N. ossifragum occurs throughout Europe and in Asia
Minor. N. americanum is confined to a narrow area about New Jersey,
where it is rare; and N. californicum appears chiefly in Northern California.
These two species are both fairly close to JN. ossifragum, of which they
have been regarded as varieties, but they appear to be worthy of specific
recognition.
The differences between the species are as follows :—In N. ossifragum
the leaves are about 7:5-15 em. long and 3-6 mm. wide, 4-6-nerved; the
stamens one-third shorter than the perianth-segments, which are narrowly
linear, 6-7 mm. long, considerably exceeding the stamens; capsule 10 mm.
long, and persistent style one-half longer than the perianth ; raceme of few
flowers, 2:2-5 cm. long ; plant usually 15-25 em. high. In N. americanum
the leaves are narrower (2-3 mm. wide), the stamens scarcely shorter
than the perianth, the capsule and style twice as long as the perianth.
A sheet of N. ossifragum in Herb. Univ. Calif. contains four specimens
from Alingsås, Sweden ,L. Linguist, July 1878, all of which are larger
and stouter. The leaves are 4-10 mm. wide, 5-7-nerved, plants 25-30 cm.
high, perianth-segments 6-8 mm. long.
The following differences are pointed out in the * Botanical Magazine’ :—
MELANTHACEA FROM THE GENETIC STANDPOINT. 141
N. americanum differs from N. ossifragum in (1) much shorter pubescence,
(2) whole plant smaller, (3) leaf and stem somewhat yellower, (4) one of
the bracts not above the middle of the pedicel but near its base, (5) corolla
much paler yellow, (6) anthers yellow instead of vermilion. According to
Pursh they differ only in (4) the position of the bracts. Comparison of
herbarium specimens confirms all the above differences except (1) and (6).
N. americanum is decidedly less stout and more yellowish, while the
perianth is paler yellow. The difference in the position of the bracts is
apparently a constant feature, but there appears to be no difference in
pubescence or in the colour of the anthers *.
N. californicum is intermediate between N. americanum and N. ossifragum
in width of leaves, its raceme is rather lax below, and the stamens about
half the length of the perianth. The filaments are also white-woolly to the
top, unlike the other three species, in which the summit of the filaments is
naked. In N. californicum the base of the filaments is frequently naked.
N. asiaticum I have not seen, but it apparently differs from N. ossifragum in
having wider leaves (2-3 lines) with conspicuous veins and a longer raceme.
Examination of a number of specimens shows the following differences
between N. americanum and N. californicum :—
N. americanum. N. californicum.
Leaves usually about 2 mm. wide, 7-9- Leaves about 3-4 mm. wide, 5-7- or 8-
nerved. nerved.
Raceme dense, 2:5-8 cm. long. Raceme lax, 7-12 em. long.
Perianth-segments narrowly linear, 5-6 mm. ^ Perianth-segments oblong-linear, 8-10 mm.
long, slightly exceeding the stamens. long, considerably exceeding the stamens,
Plants from the type-locality have orange-
coloured perianth-segments in fruit.
Capsules 10-11 mm. long. Capsules 13-15 mm. long.
Plant 30-40 em. high. Plant 33-60 em. high.
Hence N. californicum is somewhat larger, though not markedly so, in all
its parts. The number of leaves in the nerves is often the same, but they
are farther apart in N. californicum.
The following specimen in the Herbarium of the California Academy of
Science is much reduced, and is apparently like the eastern species:—
Desolation Valley, Lake Tahoe Region, Cal., Louise Hutchinson, Aug. 1909.
The plant is 2 dm. high, the leaves about 1 mm. wide, 3-4-nerved, 4-5 cm.
long, the raceme 45 cm. long, perianth-segments 5-6 mm. long, about
1mm. wide. This raises the question as to whether the differences between
N. americanum and N. californicum may not be environmentally induced.
* It is possible, however, that in N. ossifragum there may be a dimorphism in the colour
of the anthers similar to the condition found by C. H. Danforth in Tiarella cordifolia
(* Rhodora, xiii. (1911) 192-3).
ET EN,
142 DR. R. R. GATES : A STUDY OF NORTH AMERICAN
It would be very instructive to transplant N. americanum to California and
vice versá, and observe the results in successive generations.
A segregate from N. ossifragum has been described by Celakovsky * from
Corsica as JV. Reverchoni. Ut differs chiefly in having squamous bracts at
the base of the stem, larger flowers and longer erect pedicels, and in the
hairs of the filaments, which inerease in length from below upwards.
A remarkable peculiarity of the genus Narthecium is the dense pile of
white woolly hairs on the filaments. This is probably a feature which is of
no service to the plant, but which persists throughout the genus because
of inheritance, and it very likely originated through a mutation; which may
have been one of several mutations which gave rise to the generic characters
of Narthecium. The specific differences as above outlined are chiefly of a
quantitative nature, although the species are so widely separated in space.
The variations which gave rise to these species have evidently been small,
and many of them might very well be considered of quantitative nature.
It appears that these are exactly the type of variations which Darwin con-
templated in his theory of natural selection. Whether these small differences
are actually of survival value may perhaps be doubted. The question could
be settled without difficulty in this case by transplanting each species into
the range of the others and determining whether it succeeds or fails in com-
petition with another species in its native habitat. "While these morphological
differences are small, it may well be that they are correlated with invisible
physiological changes which are important in the economy of the species.
The genus Narthecium was formerly placed in the family Juncacew. The
differences between it and its nearest relatives. 7riantha and Xerophyllum,
are, unlike the specific differences within the genus, sharp and distinctive,
though T'fieldia and Narthecium are remarkably alike in habit. The latter
is compared with Triantha + in the following table :—
Triantha (Nutt.), Baker. Narthecium, Juss.
No rootstock. A creeping rootstock.
Flowers mostly in threes in a centrifugal Flowers in a terminal raceme, greenish
panicle, white or greenish. yellow.
Perianth-segments for the most part not Perianth-segments obscurely 3-5-nerved.
nerved.
Filaments slender. Filaments subulate, woolly.
Anthers rounded, erect. Anthers linear-oblong, erect, introrse.
* Oest. Bot. Zeitschr. xxxvii. (1887) 154,
T Lophiola aurea, Ker, representing another monotypic North-American genus, is found
in pine barrens from New Jersey to Florida. It is usually placed in the Hemodoracew,
and is rather remote from Narthecium in many features, but suggests a resemblance in
having tufts of wool at the base of the perianth-segments on which the stamens are
inserted. The inflorescence and upper part of the stem are, however, also clothed with
soft matted wool, as in Lachnunthes tinctoria, El.
MELANTHACE/E FROM THE GENETIC STANDPOINT. 143
The two monotypic genera, Nietneria and Petrosavia, are placed by .
Engler* next to Narthecium, though the affinities of the latter are doubtful
and the former has been placed in the Hæmodoraceæ. Nietneria corymbosa,
Klotzch & Schomb., occurs in the mountains of British Guiana. Petrosavia
stellaris, Beccari, is a root-parasite without leaves, found in Borneo. They
are only remotely related to the present group of genera. E
The genas Metanarthecium of Maximowiez, containing two species in
Japan, also resembles Narthecium in certain respects.
5. XEROPHYLLUM, Michaux.
. XEROPHYLLUM ASPHODELOIDES, Nutt.
X. asphodeloides, Nutt. Gen. i. (1818) 235.
X. setifolium, Michx. Fl. Bor, Am. i. (1803) 211; Ann. Rep. N.J. State Mus. 1910,
p. 940, pl. 35 (1911).
Helonias asphodeloides, Linn. Sp. Pl. ed. 2 (1762) 485; Curt. Bot. Mag. (1804) t. 748;
Lodd. Bot. Cab. iv. (1819) t. 394.
Dry pine barrens, southern New Jersey to eastern Tennessee and Florida.
[1
9. XEROPHYLLUM TENAX, Nutt.
X. tenax, Nutt. Gen. i. (1818) 255,
X. setifolium, Lindl. Bot. Reg. (1833) t. 1613, not Michx.
Helonias tenax, Pursh, Fl. Am. Sept, i. (1814) 243, t. 9.
On high lands near the Rocky Mountains, British Columbia to Montana, and
California (Monterey Co.). Abundant, covering hundreds of acres in some
localities.
3. XEROPHYLLUM Dovarasi, S. Wats. in Proc. Am. Acad. xiv. (1879) 284.
? X. setifolium, var., A. Gray, in Proc. Amer. Acad. viii. (1872) 405.
Oregon, Columbia River (Hall), Montana, Idaho.
The genus Xerophyllum is remarkably distinct from its nearest relatives,
Narthecium on the one hand and 77elonias on the other. The three species
are closely in agreement with each other, the differences being entirely
quantitative. Like so many Pacific coast species, X. fena is much larger
than its eastern congener, X. asphodeloides. Lindley, who studied plants
grown from seed collected by Douglas in the north-west, was “unable to
detect the slightest mark by which they may be separated,” though his
figure shows a wider-leaved plant than X. asphodeloides. X. tenax is generally
much larger and stouter, with broader leaves (4-6 mm.), the leaves of
X. asphodeloides being very narrow (1-2 mm.). The flowers of X. tenaz are
also somewhat larger, and the stamens exceed the perianth-segmenís. The
seeds also are said to differ slightly in shape. It is possible that X. tenax
is a tetraploid species, though this seems improbable.
* Nat. Pflanzenfam. T. ii. Abt. v., Liliacem, p. 20 (1887).
144 DR. R. R. GATES : A STUDY OF NORTH AMERICAN
According to Jepson *, X. tenax at Mt. Tamalpais, Calif., appears to bloom
only once in seven years; and at Howell Mountain, Napa Co., it is said to
fruit only once in five years. Plants in Sonoma Co. show “an irregular
perianth ; the (apparently) upper perianth-segment is keeled, the two
adjoining ones auricled or strongly oblique on the upper side at base;
leaves somewhat revolute-concave as if channelled.”
The following specimens of X. tenaz have narrow leaves (2 mm. or more)
as in X. asphodeloides, but flowers somewhat larger (6-8 mm. long) as in the
true X. tenax :—
S. Fork Mountain, Humboldt Co., Cal., Chesnut and Drew, July 21, 1888,
Univ. Cal. Herb. 13722. On plains near Mendocino City, Cal, 4784,
H. N. Bolander, May 8, 1866, U. C. Herb. 4083 and 4084. Mt. Tamalpais,
Marin Co., Cal., Alice Eastwood, June 16, 1901, U. C. Herb. 167934. Mt.
Eddy, Siskiyou Co., Cal., 3872, Dr. E. B. Copeland, Sept. 6, 1903, U. C.
Herb. 142151. Mt. Tamalpais, Cal, Alice Eastwood, May 1, 1898, U. C.
Herb. 142057.
It appears that narrow-leaved forms are most frequent in some localities,
and there are plants with leaves of intermediate width as well, as in specimens
from Yamhill Co., Oregon.
X. Douglasii is an obscure and little-known species. It may be questioned
whether it is distinguishable from the eastern species, with which it evidently
agrees in most particulars, It is said to have shorter pedicels than X. tenax
(12-30 mm., instead of 3-5 em ), smaller flowers, very short styles (1", instead
of 2" long), a cordate-ovate, 6-valved capsule, and shorter and broader seeds.
None of the western specimens I have seen corresponds with all these require-
ments. Intermediates between X. Douglasii and X. tenaz probably occur.
The latter species is exceedingly abundant in Idaho, where it often covers
hundreds of acres f.
S. Watson f speaks of X. Douglasii as follows :—“ Found in the mountains
from the Columbia to Montana, is a similar species [to X. tenax] with a
narrower raceme of smaller flowers ; stamens included ; styles a line long ;
capsule cordate-ovate, 2 lines long, the very abruptly acute cells usually
separating and then dehiscing loculicidally ; seeds shorter and broader.”
The distribution of the genus is interesting, the single species of the south-
eastern United States being widely sundered from X. tenax of the Rocky
Mountain region. It is perhaps worth pointing out that the main difference
between X. asphodeloides and X. tenax is similar to that between Stenanthium
gramineum and S. robustum, namely in general size and leaf-width. In the
latter ease, however, both species occupy much the same area, though
* Jepson, W. L., Fl. W. Mid. Calif. p. 124 (1901).
T According to Piper, Fl. State Washington, Contrib. U.S. Nat. Herb. xi. (1906) 197.
|. Bot. Calif. ii, (1880) 186.
MELANTHACEJE FROM THE GENETIC STANDPOINT. 145
SS. robustum is somewhat more northern and more restricted. Hence if it
originated from S. gramineum through a mutation, its origin must have been
relatively recent ; while the origin of Nerophyllum tena, if it was derived
from X. asphodeloides, is more likely to have occurred before their present
It is not necessary to assume that intermediates
Indeed,
wide separation in space.
occurred and have since become extinct in the intervening region.
I know of no faets which would justify this assumption.
The main differences between NXerophyllum and Narthecium may be set
forth as follows :—
Narthecium.
Leaves linear, those of the stem short and
distant.
Flowers small, greenish-yellow.
Pedicels with a short bract at base, and
usually bearing a small bractlet.
Perianth-segments obscurely 5-5-nerved.
Filaments woolly, anthers introrse.
Style very short or none, stigma slightly
Xerophyllum.
Leaves narrowly linear, rough-margined,
the upper ones shorter than the lower,
plants larger.
Flowers very numerous, medium-size,
white.
Pedicels with a long bract at base, but
without a bractlet.
Perianth-segments usually 5-7-nerved.
Filaments glabrous, anthers extrorsely
dehiscent.
Styles 3, filiform, reflexed or recurved.
3-lobed.
Capsule oblong.
Seeds many, linear, tailed at each end.
Capsule ovoid, 3-grooved.
Seeds 5, oblong, not appendaged or only
minutely so.
Though Narthecium and Xerophyllum resemble each other in foliage and
habit, yet the differences in inflorescence and flower-strueture are numerous
and necessitate the assumption of a considerable amount of extinction between
these forms. It is useless to hazard a guess as to the number of mutations
involved in the passage from one genus to the other, though there must have
been several. Of course, one genus did not give rise to the other, but
Nerophyllum probably came through a series of definite variations from an
ancestor of Narthecium in which the variation producing woolly filaments
had not yet occurred.
6. HELONIAS, Linn.
HELONIAS BULLATA, Linn.
Helonias bullata, Linn. Sp. Pl. (1753) 342, and Amen. Acad. iii. (1756) 12, t. 1. f 1
Mill. Ie. (1758) 181, t. 272; Bot. Mag. (1804) t. 747; Lam. Illustr. (1823) t. 268 ;
Lodd. Bot. Cab. (1824) t. 961; DO. in Redouté, Lil. i. (1805) t. 13; Ann. Rep. N.J.
State Mus. (1910), p. 540, pl. 36 (1911) ; Bartonia, iii. 1 (1910) t. 1.
Helonias latifolia, Michx. Fl. Bor. Am. i. (1803) 212.
Veratrum americanum, Mill. Gard. Dict. ed. 8 (1768) n. 4.
In bogs, southern New York and northern New Jersey to Virginia, and in
the mountains of North Carolina. Rare and local. In New York it is known
$^
146 DR. R. R. GATES: A STUDY OF NORTH AMERICAN
from only one locality, near Rossville, Staten Island *. Its northern limit is
probably Morris Co., N.J., and it is most abundant in southern New Jersey.
The plant was originally discovered by Kalm near Philadelphia, probably on
April 26, 1749, at Pennsneck, N.J. It was formerly supposed to have
-occurred in eastern Pennsylvania, but this record was probably a mistake.
The range of the species was only recently extended to North Carolina f,
where it was found to be common in sphagnum swamps of the Pink Bed
Valley, Transylvania Co., in thickets of Kalmia, Alnus, and Viburnum.
It was found in full bloom April 29, 1909, and was distributed as No. 4117.
Miller speaks of the plant as follows: Veratrum racemo simplicissimo,
corollis patentibus, staminibus longioribus. ‘This is titled by Mr. John
Bartram, who discovered the plant growing naturally in North America,
Veratrum sempervirens.” Miller received the plant from Peter Collinson,
later a specimen and drawing from Mr. John Bartram, Jr., and he afterwards
obtained more plants from Dr. Bensel, of German Town, in Philadelphia,
* who found it growing plentifully in shady moist places."
This monotypic genus of eastern North America is evidently disappearing,
and on account of its isolation must be regarded as the sole survivor of a
group of forms. Its nearest relative is the Asiatic genus Heloniopsis,
A. Gray, containing three species in Japan and one in Formosa. This
is another instance of the remarkable similarity between the floras of
eastern North America and Japan. According to Gray, Heloniopsis pauciflora
combines the extrorse anthers of Colchicaces with the loculicidal capsule,
entire style, and capitate stigma of the Liliacese, and a mass of little seeds
tailed at one or both ends as in Juncus and Narthecium, in addition to
its resemblances to //elonias. It is like a /Telonias with a single slender
style, few flowers, depressed-capitate stigma, and seed appendaged only
at the hilum.
The genera Xerophyllum and Helonias may be compared as follows :—
Xerophyllum.
Leaves narrowly linear, rough-margined.
Flowers very numerous, medium size,
white, in a large, dense, terminal raceme.
Perianth-segments oblong or ovate, 5-7-
nerved.
Stamensshorter than theperianth-segments.
Filaments subulate, anthers oblong.
Capsule ovoid, 3-grooved.
Seeds 5, not appendaged or only minutely
80.
Helonias.
Basal leaves oblanceolate, persistent.
Flowers rather large, purple, in a terminal
raceme, on a hollow, bracted scape.
Perianth-segments spatulate.
Stamens longer than the perianth-segments.
Filaments filiform, anthers ovate, blue.
Capsule obovoid, deeply 3-lobed.
Seeds numerous, white-appendaged at each
end.
The contrasts in texture and habit of these plants are much greater than
appears in a catalogue of their main taxonomic differences.
* The following facts are taken from Stewardson Brown, in ‘ Bartonia,’ iii. (1910) 1-6.
+ House, H. D., in * Mublenbergia,’ vi. (1910) 73.
MELANTHACEJE FROM THE GENETIC STANDPOINT. 147
7. CHAMAELIRIUM, Willd.
This is another monotypic genus of eastern North America, but is some-
what less isolated than the preceding.
JHAMJELIRIUM LUTEUM, A. Gray.
C. luteum, A. Gray, Man. (1848) 503.
C. carolinianum, Willd. in Ges. Nat. Fr. Berl. Mag. ii. (1808) 19.
C. obovale, Small, in Torreya, i. (1901) 108.
Veratrum luteum, Linn. Sp. Pl. (1753) 1044; Ameen. Acad. iii. (1756) 12, t. 1. f. 2.
V. flavum, Herb. ex Schult. f. Syst. Veg. vii. (1830) 1566.
Melanthium dioicum, Walt. Fl. Car. (1788) 126.
M. luteum, Willd. in Ges. Nat. Fr. Berl. Mag. ii. (1808) 22.
M. densum, Lam. Encyc. iv. (1823) 26, t. 269, f. 2.
Helonias pumilia, Jacq. Icon. Rar. (1786-1793) 2, t. 453.
H. lutea, Ker-Gawl. in Bot. Mag. (1807) t. 1062.
H. dioica, Pursh, Fl. Am. Sept. i. (1814) 243.
Ophiostachys virginica, Delile, in Redouté, Lil. viii. (1816) t. 464.
Diclinotrys, Raf. Neogen. iii. (1825).
This species is much more widespread than the preceding ones, occurring
from Ontario, Michigan, and Massachusetts, south to Florida and west to
Nebraska and Arkansas.
C. obovale, described by Small from New York, New Jersey, and the
mountains of West Virginia to North Carolina and Alabama, is said to differ
from C. luteum in having capsules obovoid or oblong-obovoid (not oblong or
ovoid-oblong), 12-14 mm. (not 7-10 mm.).long, pedicels stout, club-shaped,
about as long as the capsule (not 1-5 mm. long). The stem is also said to
be zigzag, the leaves various, and the flowers larger. I have seen specimens
showing some of these capsule characters, but they appear to be too vague and
evanescent to serve for specific distinction.
Redouté’s figure differs somewhat in foliage from that in the Botanical
Magazine. The latter has elliptical-lanceolate leaves, pointed, with 3 or 4
main parallel nerves ; the former broader, ovate-elliptical to obovate leaves,
obtuse, petioled and pinnately veined.
The following comparison shows the differences between Helonias and
Chamelirium :—
Chamelirium, Willd.
Flowers dicecious, small, white.
Scape slender, bearing small lanceolate
leaves, flowers in a long bractless spike-
like raceme.
Helonias, Linn.
Flowers perfect, rather large, purple.
Scape stout, hollow, sparsely bracted,
flowers in a bractless raceme.
Anthers blue.
Ovary ovoid, 3-grooved.
Capsule obovoid, deeply 3-lobed, the lobes
divergent.
LINN. JOURN.—BOTANY, VOL, XLIV,
Anthers white.
Ovary oblong.
Capsule oblong or obovoid, slightly 3-lobed,
N
]48 , DR. R. R. GATES : A STUDY OF NORTH AMERICAN
Seeds numerous, linear, white-appendaged Seeds 6-12 in each cavity, linear-oblong,
at each end. broadly winged at both ends, narrowly
winged at the sides.
Leaves lanceolate, the lowest spatulate, Leaves oblong-spatulate or oblanceolate,
tapering into a petiole. evergreen.
It is difficult to estimate the value of the differences between these two
genera, or the number of steps which would be necessary to bridge the gap
between them. But I believe we are more likely to be right in thinking of
the differences as having come about through definite variations than in
pieturing the gradual accumulation of infinitesimal differences.
8. AMIANTHIUM, A. Gray.
1, AMIANTHIUM MUSCJETOXICUM, A. Gray.
A. muscetoxicum, A. Gray, in Ann. Lye. N.Y. iv. (1837) 122.
Melanthium muscetoxicum, Walt. Fl. Car. (1788) 195.
M. letum, Soland. in Ait. Hort. Kew. i. (1789) 488.
M. myoctonum, J. F. Gmel. Syst. Veg. i. (1796) 587.
Helonias leta, Ker, in Bot. Mag. (1805) t. 803; Lodd. Bot. Cab. (1894) t. 998.
H. erythrosperma, Michx, Fl. Bor. Am. i. (1803) 212.
Amiantanthus muscetoxicum, Kunth, Enum. Plant. iv. (1843) 180.
Zigadenus muscetozicus, Regel, Gartenfl, xxxii. (1883) 164, t. 1121. f. 1.
Chrosperma muscetoxicum, Kuntze, Rev. Gen. Pl. (1891) 708.
Long Island and eastern Pennsylvania to Florida, Tennessee, Missouri, and
Arkansas.
2. AMIANTHIUM ANGUSTIFOLIUM, A. Gray.
A. angustifolium, &. Gray, in Ann. Lyc. N.Y. iv. (1837) 194.
Anthericum subtrigynum, Jacq. Ic. Rar. ii. (1793) t. 419.
Helonias angustifolia, Michx. Fl. Bor. Am. i. (1803) 212,
H. leta, B. minor, Ker-Gawl. in Bot. Mag. (1813) t. 1540.
Zygadenus angustifolius, S. Wats. in Proc. Am. Acad. xiv. (1879) 980.
Tracyanthus angustifolius, Small, Fl. SE. Un. St. (1903) 251.
Melanthium phalangioides? Lam, Encyc. iv. (1823) t. 269. f, 4.
North Carolina to Florida, in moist pine woods.
3." AMIANTHIUM TEXANUM, comb. nov.
Tracyanthus angustifolius, var. tevanus, Bush, Rep. Mo. Bot. Gard. xvii. ( 1906) 119.
T. texanus, Small, Fl. S.E. Un. St. 2nd ed. (1913) p. 1829.
In sandy swamps, eastern Texas.
The genus Amianthium of Gray (Chrosperma, Rafin.) contains three
species: A. muscetoxicum, extending from Long Island to Penn., Florida, and
Arkansas ; A. angustifolium of narrower distribution, from N. Carolina to
Florida ; and A. texanum in Texas. The latter two species have been placed
in a separate genus, Tracyanthus, by Small, but it would be difficult to find
MELANTHACE/E FROM THE GENETIC STANDPOINT. 149
a reason for creating a new genus in this case. A. angustifolium is smaller,
and it differs from A. muscetoxicum in its narrower leaves and smaller
flowers, in addition to certain minor differences, as in the shape of the sepals
and relative length of the stamens. But intermediates in certain localities
occupied by both species make it very difficult to draw any definite line
between them. Indeed, the two forms are by no means always easily
separated. There is a possibility of tetraploidy here. Amianthium,
Xerophyllum, and Stenanthium are all examples of essentially bitypic genera,
in which the two species differ chiefly in having broad or narrow leaves.
This type of dimorphism is a common one in the plant kingdom, and probably
has some special explanation which is at present obscure. Schanocaulon
dubium and S. Drummondii form a similar pair (see p. 150). A. texanum
differs from A. angustifolium in its larger size, broader leaves, large
compound panicles, and decidedly yellowish flowers, the perianth-segments
being oval, 4-5 mm. long. It resembles Stenanthium, but the panicle is
much smaller and less branched and the perianth-segments not narrow and
acuminate. The differences between Chamelirium and Amianthium will
appear in the following comparative table :—
Chamelirium.
Bitter, tuberous rootstock,
Basal leaves short, spatulate, tapering into
a petiole; stem-leaves lanceolate.
Flowers small, dicecious, in a long, narrow,
bractless, spike-like raceme.
Perianth-segments linear-spatulate, 1l-
nerved, withering-persistent.
Anthers subglobose.
Ovary oblong or obovate, somewhat
grooved.
Styles short, stigmatic along the inner side.
Capsules slightly 3-lobed, ellipsoid.
Seeds 6-12 in each cavity, linear-oblong,
broadly winged at both ends, narrowly
winged at the sides.
Amianthium.
Ovoid-oblong coated bulb.
Basal leaves numerous, long, linear, and
blunt; stem-leaves few and short.
Flowers perfect, in a dense terminal raceme.
Perianth-segments obtuse, persistent.
Anthers reniform, stamens inserted on the
base of the sepals.
Ovary ovoid, with 3 divergent lobes.
Styles subulate.
Capsules 3-celled, dehiscent above the
middle.
Seeds 1-2 in each cavity, ovoid, reddish
brown.
The diccism of Chamelirium and the bulb of Amianthium separate
these genera much more widely than their other morphological features,
which show many resemblances. Diccism has been attained independently
a number of times, but we do not know whether it is usually accomplished
by one step from hermaphroditism, or whether the passage is more gradual
through moncecism and polygamodicecism *. The bulbous condition has also
* The converse change, from diccism to hermaphroditism, seems to have occurred much
earlier in the evolution of the flower,
N2
E r TTS
ee
M Ws
"m
150 DR. R. R. GATES: A STUDY OF NORTH AMERICAN
no doubt been independently derived many times, as we see in Amianthium,
Anticlea, and Allium within this group. It is more difficult to visualize the
passage from a rootstock to a bulb than in the above case of dicecism. It is
very difficult to believe that the passage is ever abrupt, and in certain plants,
such as Amianthium, the condition appears to be somewhat intermediate, the
bulbous condition not having been fully arrived at.
Amianthium is perhaps more closely related to Schanocaulon, whose main
distinguishing features are as follows :—
Scheenocaulon, Amianthium.
Fibrous-coated bulbs. Bulb-like rootstocks.
Leaves basal, narrowly linear. Leaves linear, blunt-pointed, those of the
stem much fewer and shorter.
Flowers in slender spikes, or in spike-like Flowers larger, in cylindrical racemes with
racemes with short pedicels. long pedicels.
Perianth green, persistent, sepals and petals Perianth white, sepals and petals broad.
narrow. j
Seeds slender, nearly terete. Seeds thick, reddish brown.
9. SCHCENOCAULON, A. Gray.
The genus contains about twelve species, of which all but two are confined
to Mexico and will not be considered here. These two are as follows :—
1. Scu&NocAULON DUBIUM (Michx.) Small.
S. dubium, Small, Fl. SE. Un. St. (1905) 250.
S. gracile, A. Gray, in Ann. Lye. N.Y. iv. (1837) 127.
Helonias dubia, Michx. Fl. Bor, Am. i. (1803) 213.
In dry pine lands, Georgia and Florida.
2. Scnawocauton DRUMMONDII, A. Gray; Hook. & Arn. Dot. Beechey
Voy. (1841) 388.
S. texanum, Scheele, in Linnæa, xxv. (1852) 262.
On prairies, Texas and Northern Mexico.
These two species, apparently discontinuous in their distribution, form an
interesting pair :—
S. dubium: compact portion of spike 5 mm. thick ; sepals oblong; capsules sessile.
S. Drummondii: compact portion of raceme 10 mm. thick; sepals linear; capsules
pedicelled.
S. Drummondii is stouter in every part, having larger flowers and wider
leaves. ©. dubium has apparently been derived from it by reduction. It is
not a dwarf in the ordinary sense, but more like a miniature. The
chromosome relations of these species would be worth looking into,
UND CUXM€CUK-
MELANTHACEJÉ FROM THE GENETIC STANDPOINT. 151
10. STENANTHIUM, Kunth.
This genus as now recognized contains only two species.
1. STENANTHIUM GRAMINEUM (Ker), Morong.
S. gramineum, Morong, in Mem. Torr. Club, v. (1894) 110.
S. angustifolium, Kunth, Enum. iv. (1843) 190.
Helonias graminea, Ker, in Bot. Mag. (1813) t. 1599.
Veratrum angustifolium, Pursh, Fl. Am. Sept. i. 242, ii. 747 (1814).
Xerophyllum gramineum, Nutt. Gen. i. (1818) 235.
Virginia to Kentucky, Missouri, Alabama, and Florida.
bo
. STENANTHIUM ROBUSTUM, S. Wats., in Proc. Am. Acad. xiv. (1879) 278.
Pennsylvania and Ohio to South Carolina, Tennessee, and Missouri.
These two species have been united by some writers, but S. robustum is no
doubt worthy of independent recognition. It is larger and stouter, as the
name indicates, and may be a tetraploid mutant or cell-giant from
S. gramineum. In S. robustum the capsule is erect, in S. gramineum reflexed.
The former has leaves 4-6 mm. wide; the latter, leaves 6-20 mm. wide.
The capsules of S. robustum are also larger (10 mm. long, instead of 8 mm.)
and the perianth is said to be usually green, while in S. gramineum it
is usually white.
The two species occupy nearly the same territory, S. robustum extending a
little further north and not so far south.
improbably hybrids.
should be made.
Intermediate specimens are not
A careful anatomical comparison of these species
The genus Stenanthium is closely related to Amianthium, which it resembles
in foliage and with which it agrees in having bulb-like rootstocks. There
are, however, conspicuous differences in inflorescence and flower-structure,
as shown in the following comparison :—
Amianthium, Gray.
Leaves linear, blunt-pointed, stem-leaves
few and short.
Flowers perfect, white, in a dense terminal
cylindrical raceme.
Perianth-segments broad, obtuse.
Stamens as long as or longer than the
perianth-segments.
Capsule 3-celled, dehiscent above the
middle, the divergent lobes tipped by the
subulate styles.
Stenanthium, Kunth.
Leaves linear, narrower, blunt - pointed,
basal leaves less numerous.
Flowers polygamous, white or greenish, in
an ample terminal panicle.
Perianth-segments narrowly lanceolate,
acuminate, spreading.
Stamens shorter than the perianth-segments.
Capsule ovoid-oblong, 3-lobed, dehiscent to
base, the lobes with short, slightly
divergent beaks.
The paniculate inflorescence and narrow acuminate petals of Stenanthium
are its most striking features.
The change in the petals might represent a
single variation, but it is difficult to say whether one or two germinal changes
could transform the racemose inflorescence of Amianthium into the loose
Log DR. R. R. GATES: A STUDY OF NORTH AMERICAN
panicle of Stenanthium. It may suffice to point out that the traditional
conception of gradual transition is based upon suppositions which require
proof as much as do those of sharp steps. If crossing experiments between
these two genera could be instituted, it might throw light on the nature of
all these differences.
11. STENANTHELLA, Rydb.
The genus Stenanthella was segregated from Stenanthium by Rydberg, to
contain the two species S. sachalinensis (F. Schmidt), Rydb., and S. occidentalis.
To these should be added S. frigidum from Mexico, which stands much nearer
to Stenanthella than to Stenanthium. The North-American species are then
as follows :—
1. STENANTHELLA OCCIDENTALIS (A. Gray), Rydb.
S. occidentalis, Rydb. in Bull. Torr. Bot. Club, xxvii. (1900) 531.
Stenanthium occidentale, A. Gray, in Proc. Amer. Acad. viii. (1872) 405.
Alberta and British Columbia to Oregon and Montana, and south to Trinity
Co., California.
2. STENANTHELLA FRIGIDA (Schlecht. et Cham.), comb. nov.
Veratrum frigidum, Schlecht. et Chamisso, in Linnea, vi. (1831) 46.
Zigadenus frigidus, D. Don, in Edinb. N. Phil. Journ. (1832) 933.
Stenanthium frigidum, Kunth, Enum, Plant. iv. (1843) 189.
Mexico.
In S. frigida the ovary is partly inferior, but this alone is not sufticient
reason for separating it from ©. occidentalis, which it otherwise clearly
resembles. It differs from the latter chiefly in its somewhat larger size,
larger flowers, dark purple perianth, and shorter and less pointed capsules.
The genus Stenanthella, which is markedly distinct from Stenanthium,
differs mainly in the following characters :—
Stenanthium. Stenanthella,
Inflorescence a large panicle of many sessile Inflorescence a raceme of few flowers on
flowers. long pedicels,
Flowers polygamous, small, white or Flowers perfect, much larger, greenish,
greenish, open, not campanulate. brownish, or purplish, campanulate.
Tips of perianth-segments not reflexed. Tips of perianth-segments reflexed and
finally involute.
12. ZIGADENUS, Michaux.
1. ZIGADENUS GLABERRIMUS, Michx. Fl. Bor. Am. i. (1803) 214, t. 22 ; Del.
in Redouté, Lil. viii. t. 461 (1816) ; Rydb. in Bull. Torr. Bot. Club,
xxx. (1903) 280, pl. 12. f. 1.
Helonias glaberrima, Link, Enum, Hort. Berol. i. (1821) 348.
In swamps, Virginia to Florida, and west to Mississippi, usually near the
coast,
MELANTHACEA FROM THE GENETIC STANDPOINT. 153
Specimens: Bolling, Butler Co., Alabama, John D. Smith, 1884. Ocean
Springs, Miss., J. Skehan, 1895. Long Beach, Miss., J. F. Joor, 1891.
2.
ZIGADENUS BRACTEATUS, Roem. & Schult. Syst. vii. (1830) 1559.
Helonias bracteata, Sims, Bot. Mag. (1815) t. 1703; Lodd. Bot. Cab. (1828) t. 1380.
This has usually been referred to Z. glaberrimus, but it appears to be
a different species, though I have seen no specimens. It is stated in Lodd.
Bot. Cab. to be native of Carolina and Georgia, whence it was introduced
into cultivation in 1802. The species differs from Z. glaberrimus in its very
much larger and broader bracts and shorter pedicels. In Z. glaberrimus
the bracts scarcely equal the pedicels in length, while in Z. bracteatus they
exceed the flowers. The inflorescence is also perhaps more compact and the
petals apparently broader. This species should be looked for in association
with Z. glaberrimus, with which it has apparently been confused. The figure
in the Bot. Mag. is evidently from a very young specimen in which the
inflorescence has not yet elongated, but, nevertheless, the size and shape of
the bracts seem to separate it from Z. glaberrimus.
_
13. ANTICLEA, Kunth.
. ANTICLEA SIBIRICA (Linn.), Kunth.
Melanthium sibiricum, Linn. Sp. Pl. (1758) 339.
Zigadenus sibiricus, Kunth, Enum. Pl. iv. (1843) 192; Endl. Gen. 135; A. Gray, in Ann.
Lyc. N.Y. iv. (1837) 112.
Anticlea sibirica, Kunth, Enum. Pl. iv. (1843) 191 ; Rydb. in Bull. Torr. Bot. Club, xxx.
(1903) 280, pl. 13. f. 4.
Siberia.
. ANTIOLEA JAPONICA (Makino), comb. nov.
Zigadenus japonicus, Makino, in Bot. Mag. Tokyo, xvii. (1903) 162.
Japan.
Said to resemble A. elegans, but has smaller flowers, narrower not glaucous
leaves, stem shorter, and pedicels shorter. It apparently differs from
A. sibirica.
8.
ANTICLEA ELEGANS (Pursh), Rydb.
Ziyadenus elegans, Pursh, Fl. Am. Sept. i. (1814) 241.
Helonias glaberrima, Ker, in Bot. Mag. (1814) t. 1680.
? Asphodelo affinis floridana, ramoso eaule, floribus Ornithogali obsoletis, Pluk. Amalth.
(1705) p. 40, t. 572. f. 3.
Anticlea elegans, Rydb. in Bull. Torr. Bot. Club, xxx. (1903) 273, pl. 13. f. 3.
Zigadenus dilatatus, Greene, Pl. Baker. i. (1901) 51. Colorado.
Zigadenus alpinus, Blankin. in Mont. Agr. Coll. Sci. Studies, Bot. i. (1905) 44.
Anticlea alpina, Heller, in Muhlenbergia, vi. (1910) 12. Montana.
Saskatchewan to Alaska, south to Colorado, Nevada, and Minnesota.
154 DR. R, R. GATES: A STUDY OF NORTH AMERICAN
4. ANTICLEA CHLOKANTHA (Richards), Rydb.
A, glauca, Kunth, Enum, PI. iv. (1843) 192.
A. chlorantha, Rydb. in Bull. Torr. Bot. Club, xxx. (1903) 273.
Zigadenus chloranthus, Richards, in Frankl. Ist Journ. (1821) 736.
Z. glaucus, Nutt. in Journ. Phila. Acad. vii. (1834) 56; Lindl. in Bot. Reg. xxiv. (1838)
t. 67.
Z. canadensis, Hort. ex Baker, in Journ. Linn, Soc., Bot. xvii. (1879) 482.
Z. chloranthus, B. major, Hook. & Arn. Bot. Beechey Voy. (1841) 402.
Z. glaberrimus, Hook. & Arn. Bot. Beechey Voy. (1841) 160.
Melanthium glaucum, Nutt. Gen. i, (1818) 232.
New Brunswick to Minnesota and south to Vermont.
e
. ANTICLEA COLORADENSIS, Rydb.
A. coloradensis, Rydb. in Bull. Torr. Bot, Club, xxx. (1903) 273. i
Zigadenus coloradensis, Rydb, in Bull. Torr. Bot. Club, xxvii. (1900) 534.
Z. elegans, var. coloradensis, M. E. Jones, in Bull, Univ. Mont., Biol. Ser. xv. (1910) 22.
Colorado.
Specimens : Red Creek, Colo., N. L. T. Nelson, July 18, 1908 (2 sheets).
Near Empire, Colo, /7. N. Patterson, 298, July 28, Sept. 6, 1892.
Jarbridge, Head of Jack and Swamp Creeks, 2018, Nelson and Macbride,
July 11, 1912. These specimens are by no means uniform, but they represent
some at least of the supposed peculiarities of A. coloradensis.
According to Rydberg, this species differs from A. elegans in its “smaller
flowers, greener foliage, long and narrow bracts equalling or exceeding the
pedicels, and a brownish or purplish tint of the inflorescence, bracts, and
flowers.” These differences are, however, by no means sharp in all cases,
and intermediates occur in size of flowers and colour and length of bracts.
The leaves vary in width from 3-15 mm. The perianth-segments are said to
be 7-8 mm. long in Z. elegans, 5-6 mm. in Z. coloradensis ; 7-13-nerved in
the former, 3-7-nerved in the latter.
The toxic properties of several species of Zigadenus are discussed in a
recent Bulletin *. It was found that Anticlea elegans, Toaicoscordion venenosum
and T. paniculatum were constantly poisonous to sheep, while A. coloradensis,
though it contained small quantities of alkaloid, was not very toxic. This
perhaps indicates the specific distinctness of the last species, for the properties
of each species were found to be generally constant. But 7. venenosum from
one locality was less toxic than when collected elsewhere.
6. ANTICLEA VAGINATA, Rydb. in Bull. Torr. Bot. Club, xxxix. (1912) 108.
Utah.
Differs from other species in its habit of growing in big clumps, and in its
numerous loose sheaths at the base of the stem. In the perianth it resembles
* Marsh, C., Dwight,'Clawson, A. B., and Marsh, H., 1915. *Zygadenus, or Death Camas."
Bull. No. 125, U.S. Dept. Agr. pp. 46, pls. 6, including 3 figs. of * Z. venenosus."
MELANTHACEJE FROM THE GENETIC STANDPOINT. 155
A. coloradensis, and in their veins A. porrifolia. The perianth-segments are
smaller than in the former and broader than in the latter. A. vaginata
resembles A. porrifolia also in its branched inflorescence, but has shorter
pedicels and broader leaves.
7. ANTICLEA LONGA (Greene), Heller.
A. longa, Heller, in Muhlenbergia, vi. (1910) 12.
Zigadenus longus, Greene, Pittonia, iv. (1901) 240.
Oregon.
8. ANTICLEA PORRIFOLIA (Greene), Rydb.
A. porrifolia, Rydb. in Bull. Torr. Bot. Club, xxx. (1903) 273.
Zigadenus porrifolius, Greene, in Bull. Torr. Bot. Club, viii. (1881) 125.
New Mexico.
Pale and soft, garlic-like, somewhat glaucous leaves; bulb oblong with
white outer coats; raceme simple or few-branched below; bracts ovate-
lanceolate, green and glaucous ; flowers small, nodding; perianth adnate,
persistent, segments greenish, broad, none unguiculate; gland elongated,
truncate, or obcordate.
9. "ANTICLEA MOHINORENSIS (Greenm.), comb. nov.
Zigad hinorensis, Greenm. in Proc. Amer. Acad. xxxix. (1903) 71.
g
Chihuahua, Mexico.
Nearest to A. elegans, from which it differs in its somewhat larger flowers,
more conspicuous and longer bracts of the inflorescence, and in the nature of
the glands, which are obeordate or Y-shaped, erose-margined.
10. ANTICLEA VOLCANICA (Benth.), Baker.
A, volcanica, Baker, in Journ. Linn. Soc., Bot. xvii. (1879) 482.
Zigadenus volcanicus, Benth. Pl. Hartweg. (1842) 96.
Guatemala.
ll. ANTICLEA VIRESCENS (H. B. K.), Rydb.
A, virescens, Rydb. in Bull. Torr. Bot. Club, xxx. (1903) 273.
Helonias virescens, Kunth, Nov. Gen. Voy. Humb. et Bonpl. i. (1815) 267.
Veratrum virescens, Mart. et Gal. Enum, Pl. Mex. 10; Kunth, Enum, Pl. iv. (1843) 698.
Melanthium virescens, Willd. Herb, No. 7076 (ex Kunth, 7. c.).
Anticlea mexicana, Kunth, Enum. Pl. iv. (1843) 193.
Zigadenus mexicanus, Hemsl. Biol. Centr.-Am., Bot. iii. (1885) 382.
Zigadenus volcanicus, S. Wats. in Proc. Am. Acad. xviii. (1883) 165.
Mexico.
A
12. ANTICLEA GRACILENTA (Greene), comb, nov.
Zigadenus gracilentus, Greene, Pittonia, iv. (1901) 241.
Mexico.
OTTIMA TERN Ee Sl) RT NISUS TT E S
156 DR. R. R. GATES: A STUDY OF NORTH AMERICAN
Said to be nearly related to A. elegans, but distinguished by the oblanceolate
leaves and the dicecious flowers.
14. TOXICOSCORDION, Rydberg.
1. TOXICOSCORDION VENENOSUM (S Wats.), Rydb.
T. venenosum, Rydb. in Bull. Torr. Bot. Club, xxx. (1903) 272.
Zigad. venenosus, S. Wats. in Proc. Am. Acad. xiv. (1879) 279.
Z. Nuttallii, S. Wats. partim, in Bot. King’s Exped. v. (1871) 343.
British Columbia to California. Not east of Wyoming. The type may
be considered to come from Monterey Co., Calif. Watson's description
included plants from the Coast Range and also from the Sierra Nevada.
These belong to distinct species, and the Monterey Co. locality is selected
merely because mentioned first. hat the Sierran species is distinct
is proven by specimens of the flowers shown me by Dr. H. M. Hall. The
flowers of the latter are much smaller, and the stamens exserted beyond the
petals.
2. TOXICOSCORDION INTERMEDIUM, Rydb.
T. intermedium, Rydb. in Bull. Torr. Bot. Club, xxx. (1903) 272, pl. 13. f. 2.
T. gramineum, Rydb. 7. c. xxx. (1903) 272.
Zigadenus intermedius, Rydb. l. c. xxvii. (1900) 535.
Z. gramineus, Rydb. l. c.
Z. venenosus, var. ambiguus, M. E. Jones, in Contrib. West. Bot. xii. (1908) 77.
Montana, Idaho, Wyoming, and Utah.
Rydberg now considers (l. c. xxxix. 109, 1912) that T. gramineum cannot
be separated from 7. intermedium, being a dry hill state of the same with
smaller flowers and narrower leaves. It is doubtful if this species can be
separated from T. venenosum.
3. TOXICOSCORDION ACUTUM, Rydb.
T. acutum, Rydb. in Bull. Torr. Bot. Club, xxx. (1903) 272.
Zigadenus acutus, Rydb. l c. xxvii. (1900) 536.
South Dakota. A
This is said by Rydberg to be easily distinguished from T. venenosum by
*the acute short-clawed petals and sepals, which are both cuneate at
the base." The species is apparently not well marked.
4. "l'OXICOSCORDION SALINUM (A. Nelson), comb. nov.
Zigadenus salinus, A, Nelson, in Coult. Bot. Gazette, liv. (1912) 406.
Idaho.
This species is near to 7. venenosum and T. intermedium, but has globose
bulbs and small glands. Tt lives in alkali-bog lands.
MELANTHACE FROM THE GENETIC STANDPOINT. 15
5. TOXICOSCORDION ARENICOLA, Heller, in Muhlenbergia, ii. (1906) 182.
Inyo Co., California, Washoe Co., Nevada. ;
This is a segregate from T. venenosum. The bulbs are broadly ovoid,
clothed with dark brown outer coats. The leaves about half the length
of stem. Inflorescence at first short and dense, becoming 1-2 dm. long ;
pedicels very slender, spreading and ascending ; bracts membranous,
acuminate ; perianth white, segments faintly 3-toothed at the apex and very
short-clawed, with a small, yellow, orbicular, fimbriate gland just above
the claw.
6. Toxicoscorpion NurrTALLI (A. Gray), Rydb.
T. Nuttall, Rydb. in Bull. Torr. Bot. Club, xxx. (1903) 272.
Amianthium Nuttallii, var. a, A. Gray, in Ann. Lyc. N.Y. iv. (1837) 123.
Amiantanthus Nuttallii, Kunth, Enum, Pl. iv. (1843) 181.
Anticlea Nuttallii, Torr. Pacif. R. R. Rep. iv. (1856) 144.
Zigadenus Nuttallit, A. Gray, Man. ed. 2 (1857) 476; Gartenfl. xxxii. (1883) 163
t 1121 172.
Helonias angustifolia, Nutt. in Trans, Am. Phil. Soc. v. (1837) 154.
Colorado, Kansas, and Arkansas.
Bulb with blackish skin, white within, sepals variable, gland small,
roundish, without distinct margin.
1. Toxicoscorpion Fremontit (Torr.), Rydb.
T. Fremontii, Rydb. in Bull. Torr. Bot. Club, xxx. (1903) 273.
Zigadenus speciosus, Dougl. ex Hook. Fl. Bor. Am. ii. (1840) 177.
Z. glaberrimus, Hook. & Arn. Bot. Beechey Voy. (1843) 160.
Z. Douglasit, Torr, Pacif. R. R. Rep. vii. (1856) 20,
Z. Fremontüi, Torr. ex S. Wats., Bot. King's Exped. v. (1871) 343.
Z. glaucus, Nutt. ex Baker, in Journ. Linn. Soc., Bot. xvii. (1879) 480.
Anticlea Fremontii, Torr. Pacif. R. R. Rep. iv. (1856) 144.
California.
Sepals not clawed, 5-7-nerved, gland irregular-margined above ; stamens
half as long as the perianth.
v,
ToxicosconprioN Fremont (Torr.), Rydb., var. MINOR (Hook. & Arn.), comb.
nov.
" Zigadenus chloranthus, a, minor, Hook. & Arn. Bot. Beechey Voy. (1841) 402.
Z. commutatus, Schult, Syst. Veg. vii. (1830) 1561.
Z. speciosus, var. minor, Greene, Man. Bay Region, (1894) 315.
Z. Fremontii, var. minor, Hook. & Arn, fide Jepson, Fl. W. Mid. Calif. (1901) 129.
Smaller in every part. This is an early dwarf form, 4 or 5 inches high,
with few flowers, in open wet ground near the coast (Jepson, Fl. 122).
8. TOXICOSCORDION ExALTATUM (Eastw.), Heller.
T. exaltatum, Heller, in Muhlenbergia, vi. (1910) 83.
Zigadenus exaltatus, Wastw. in Coult. Bot. Gazette, xli. (1906) 283.
158 DR. R. R. GATES: A STUDY OF NORTH AMERICAN
California.
This is said "by the author to be the largest species of Zigadenus and to be
related to 7. paniculatum in habit and size of bulb, but the flower parts are
of different shape.
9. ToxICOSCORDION BREVIBRACTEATUS (M. E. Jones), comb. nov.
Zigadenus Fremontii, var. brevibracteatus, M. E. Jones, Contrib. West. Bot. xii. (1908) 78.
Z. brevibracteatus, Hall, in Univ. of Cal. Publ., Botany, vi. (1915) 165.
Victor, Calif., and along the southern borders of the Mohave Desert.
Differs in having very narrow leaves, small flowers, ample compound
racemes, stout horizontal pedicels 25-40 mm. long, and ovate-lanceolate
bracts. "This is a desert species.
10. ToxrcoscoRDION PANICULATUM (Nutt.), Rydb.
T. paniculatum, Rydb. in Bull. Torr. Bot. Club, xxx. (1903) 272.
Helonias paniculata, Nutt. in Journ. Philad. Acad. vii. (1834) 57.
Amianthium Nuttallii, var, B, Gray, in Ann, Lyc. N.Y. iv. (1887) 123.
Zigadenus iculatus, S. Wats. in Bot. King’s Exped. v. (1871) 343.
£
Z. Nutta, var. paniculatus, Baker, in Journ. Linn. Soc., Bot. xvii, (1879) 481.
Montana and Washington to New Mexico and California.
Distinguished, according to Rydberg, by its “stout habit, generally
branched inflorescence, and rhombic-ovate, acute, and almost clawless petals
and sepals.”
11. Toxicoscorpion rALcATUM, Rydb.
T. falcatum, Rydb. Bull. Torr. Bot. Club, xxx. (1903) 272.
Zigadenus falcatus, Rydb. l. c. xxvii. (1900) 536.
Colorado.
The following specimens in Herb. Missouri Bot. Gard. are referred to this
species : Canon City, Colo., 24, T. S. Brandegee, May, 1871. Fort Collins,
Colo., C. S. Crandall, May 17, 1894 (cotype). Denver, Colo., E. C. Smith,
May 21, 1891. Near Fort Collins, C. F. Baker, May 24, 1896.
Said to differ from 7. paniculatum in the distinctly clawed petals, which
are subcordate at base. “These characters, together with the more distinct
glands and the slightly adnate filaments, distinguish it from T. Nuttalli.”
12. ToxtcoscorDION MICRANTHUM (Hastw.), Heller.
T. micranthum, Heller, in Muhlenbergia, vi. (1910) 85.
Zigadenus micranthus, Easvw. in Bull. Torr. Bot. Club, xxx. (1903) 483.
Mendocino Co., California.
It is very doubttul if this differs from the true 7. venenosum.
13. TOXICOSCORDION TEXENSE, Rydb. ex Small, Fl. S.E. Un. St. (1903)
252, 1328.
Texas.
MELANTHACEA FROM THE GENETIC STANDPOINT. 159
Raceme many-flowered ; perianth yellow, segments clawed, subcordate,
about 5 mm. long; glands half-orbicular, the upper edge free.
15. OCEANORUS, Small.
OCEANORUS LEIMANTHOIDES (A. Gray), Smail.
Amianthium leimanthoides, A. Gray, in Ann. Lyc. N.Y. iv. (1837) 125.
Amiantanthus leimanthoides, Kunth, Enum. Pl. iv. (1848) 183.
Zigadenus leimanthoides, A. Gray, Man. ed. 2 (1857) p. 476.
Helonias graminea, Ell. Herb. (ex Kunth, /. c.).
Oceanorus leimanthoides, Small, Fl. S.E. Un. St. (1903) 252, 1328.
Long Island to Georgia and Tennessee.
The old genus Zigadenus has been broken up into four closely related
genera. These are Anticlea, Kunth, Enum. iv. (1843) 191; Towicoscordion,
Rydberg, in Bull. Torr. Bot. Club, xxx. (1903) 272, and Oceanorus Small, FI.
S.E. Un. St. (1903) 252, leaving Zigadenus itself with two species, only one
of which is known in the wild condition. Z. glaberrimus comes close to
Melanthium, so close that very good reasons might be adduced for including
it in that genus. The differences between these genera are as follows :—
Plant witha rootstock; perianth-segments each with two glands. Zigadenus.
Plant with a bulb-like rootstock ; perianth-segments each with a single
thick gland. Oceanorus.
Plant with a bulb; perianth-segments each with a single gland.
Plant poisonous, ovary wholly superior; gland obovate or semi-orbicular.
"Toxicoscordion.
Ovary partly inferior ; gland obcordate or notched. Anticlea.
The habit, with a rootstock or bulb; and the number and shape of the
glands at the base of the perianth-segments, are thus the main distinguishing
features. It is reasonable to suppose that the gland differences at any rate
may have originated by single steps. Anticlea and Toxicoscordion, the former
having about 10 species in North America and two in northern Asia, and the
latter with about 12 species in western N. America, might be regarded as
the oldest genera. But, so far as I am aware, the bulbous condition, which
has arisen independently in many families, has always been derived from an
ordinary stem or a rootstock, and not vice versá. Hence we must regard the
ancestor of Zigadenus glaberrimus as representing the original stem, from
which in one direction Z. glaberrimus was derived, perhaps through a
mutation in which the basal glands became divided into two, while in
Z. (Oceanorus) leimanthoides the bulb-like development began, to be completed
in the ancestors of Toxicoscordion and Anticlea. In the latter, meantime, the
glands became obcordate or more or less bifurcated, and the ovary partly
inferior.
It is necessary to assume an extinct ancestor having a rootstock and
a single undivided gland, Aside from this, there is no evidence of
160 DR. R. R. GATES: A STUDY OF NORTH AMERICAN
any extinction having taken place between these genera. There is, for
example, no gap between the species of Tozicoscordion and those of Anticlea.
In the absence of such a gap, the separation of these two genera is by
no means a sharp one, and indeed is only justifiable on grounds of convenience.
Since Z. glaberrimus is confined to the south-eastern States, while Towicoscordion
and Anticlea cover a much wider area, it may perhaps be assumed, considering
also the other relationships of Zigadenus, that its origin took place relatively
recently. On the other hand, it is possible that it may be a survivor of a
group similar to the present Anticlea.
The distinctions between species in Anticlea and Toxicoscordion are founded
chiefly upon such features as the shape of the perianth-glands, the size and
shape of sepals and petals, whether clawed or clawless, the width of leaves,
and the nature of the raceme. These differences are for the most part not
sharply marked presence and absence characters, but tend to be quantitative
and continuous. This makes the discrimination of species difficult. The
two genera are, in fact, in rather a chaotic condition, owing partly to the
nature of the specific differences, partly to their variability, and partly to the
description of a number of new species on a rather slender basis and from
insufficient material to determine the range and manner of variation of the
various forms.
In the preceding pages I have merely assembled the species, some of which
will very likely be reduced to synonymy, but the genus can only be adequately
treated after some botanist has made a careful study in the field.
We have already concluded that the present Zigadenus represents an
ancestral condition from which Oceanorus, Anticlea, and Toxicoscordion were
derived. On the other hand, Zigadenus has clearly led to Melanthium, and
Melanthium to Veratrum through the loss of the glands on the perianth-
segments. Hence we may conclude that Zigadenus is the oldest genus in
this section of the Melanthacez. Presumably it and Stenanthium are
descended from a common ancestor, Zigadenus having developed the pairs of
glands on the perianth-segments and also the claws on these segments, How
these two features developed, we have at present no means of knowing, but
there has probably been considerable extinction between the present genera
Zigadenus and Stenanthium.
16. MELANTHIUM, Clayt., ex Linn.
1. MELANTHIUM VIRGINICUM, Linn. Sp. Pl. (1753) 339; Lamk. Encyc. iv.
(1823) 24, t, 269. f. 1,
M. letum, Kinn, ex Kunth, Enum. Pl, iv. (1843) 195.
M. biglandulosum, Bertol. in Mem. Acc. Sc. Bologn. ii. (1850) 316.
Asphodelus elatior Floridanus gramineis foliis, floribus parvis et herbaceo-pallescentibus,
Pluk, Amalth. (1705) 40, t. 434. f. 8,
a
MELANTHACEJE FROM THE GENETIC STANDPOINT. 161
Leimanthium virginicum, Willd. Sp. Pl. ii. (1799) p. 266; A. Gray, in Ann. Lyc. N.Y.
iv. (1837) 115, partim.
Helonias virginica, Sims, Bot. Mag. (1806) t. 985.
Veratrum virginicum, Ait. Hort. Kew. ed. 2, v. (1813) p. 426.
Zigadenus virginicus, Kunth, Enum. Pl. iv. (1843) 195 partim.
In meadows and swamps, Rhode Island to Minnesota, and south to Florida
and Texas.
2. MELANTHIUM HYBRIDUM, Walt. Fl. Carol. (1788) 125; Pursh, Fl. Am.
Sept. i. (1814) 241 ; Ell. Sketch, i. (1821) 418.
M. racemosum, Michx. Fl. Bor. Am. ii. (1803) 251.
M. latifolium, Desrouss. in Lam. Encyc. iv. (1797) 25.
Leimanthium hybridum, Schult. Syst. Veg. vii. (1830) 1550; A. Gray, in Ann, Lyc. N.Y.
iv. (1837) 115.
Zigadenus hybridus, Kunth, Enum, Pl. iv. (1843) 196.
In dry woods and on hills, Connecticut to Pennsylvania, and South
Carolina.
3. MEgLANTHIUM MONOICUM, Walt. Fl. Carol. (1788) 125; Pursh, Fl. Am.
Sept. i. (1814) 241 ; Ell. Sketch, i. (1821) 418.
M. polygamum, Desrouss, in Lam. Encyc. iv. (1797) 95.
M. dispersum, Small, in Bull. Torr. Bot. Club, xxv. (1898) 606.
Leimanthium monoicum, Schult. Syst. Veg. vii. (1830) 1550; A. Gray, in Ann. Lyc. N.Y.
iv. (1837) 116 partim.
In woods. This species appears to be quite restricted in distribution.
Small described his M. dispersum from Walton Co., in the north of Florida,
and the following specimen is in Herb. Missouri Bot. Gard.: Sphagnous
bog about 2 miles east of Folkston, Charlton Co., Georgia, 1506, Roland M.
Harper, Aug. 12, 1902. Pursh gives as the distribution, “ on the mountains
of Virginia and Carolina."
This genus of three species is confined for the most part to south-eastern
North America, though M. virginicum extends farther north and west.
M. monoicum occurs on the borders of Florida and Georgia, and probably
also in Virginia and Carolina. The main specific differences are as follows :—
M. virginicum. M. hybridum.
Leaves linear, acuminate, 4’—12” wide. Leaves oblanceolate, acute, 6’-2' wide.
Flowers 6"-10" broad, greenish yellow. Flowers 6"-8" broad, greenish white.
Perianth-segments oblong, obtuse, flat, Perianth-segments orbicular or ovate,
entire, blade at least twice as long as undulate or crisped, blade longer than
the claw *. the narrow claw.
M. monoicum, Walt., differs from M. virginicum in having (1) longer and
narrower leaves, (2) a very broad instead of a narrow paniele, (3) fewer
flowers on stouter pedicels, (4) perianth-segments thicker and with shorter
claws. The stamens are attached near the top of the claw.
* The perianth-segments are also usually more pubescent on the outside,
pom Pm ES
162 DR. R. R. GATES: A STUDY OF NORTH AMERICAN
Melanthium is closely related to Zigadenus, which it resembles in having
the perianth-segments clawed, with two glands at the base of the blade, but
the glands in Melanthiwm tend to be confluent. The two genera also
resemble each other in foliage and in inflorescence, but Zigadenus is glabrous
while Melanthium is characteristically seabrous-pubescent. As regards the
stamens, they are free from the perianth-segments and about equalling them
in length in Zigadenus, while in Melanthium they are shorter than the
segments to which they are adnate. The seeds of the former are oblong or
linear, while those of the latter are flat and broadly winged. The capsules
also show characteristic differences. Thus in Melanthium they are truncate
at the top, while in Zigadenus they are more gradually narrowed and pulled
out, as it were, at the points where the stigmas protrude. Another difference
is found in the flowers, which are moncecious in Melanthium, hermaphrodite
in Zigadenus.
The differences between the three species of Melanthium are typical specific
differences, and there is no direct evidence as to how they came about. The
genus is closely related to Zigadenus on the one hand, and through
V. parviflorum to Veratrum on the other hand. The scabrous pubescence
of Melanthium may be supposed to have originated suddenly in a Zigadenus-
like ancestor. ‘The differences in relative stamen-length and in the adnation
of the stamens to the petals are no greater than in the species of, e. g.,
Trillium. Hence it is unnecessary to suppose any greater amount of
extinction between these three genera than occurs between species of the
sume genus.
The resemblances and differences between Melanthium and Veratrum are
pointed out in the following table. The two genera agree in certain features,
such as the pubescence and the shape of ilie seeds, in which they are con-
trasted with Zigadenus. On the other hand, Melanthium and Zigadenus are
closely similar in the glands on the perianth, which in Veratrum have been
almost completely lost :—
Melanthium.
Stem and inflorescence scabrous-pubescent.
Leaves narrow, sheathing.
Flowers greenish, white or cream-colored ;
moneecious or polygamous; on slender
pedicels; in large terminal panicles.
Perianth-segments clawed, with 2 more or
less confluent elands at base, mostly free
from the ovary.
Stamens shorter than the perianth-segments
and adnate to them.
Seeds very flat, broadly winged.
Veratrum.
Rootstocks poisonous.
Stem and inflorescence pubescent.
Leaves broad, clasping, strongly veined.
Flowers greenish, yellowish, or purple,
rather large; moncecious or polygamous ;
on short stout pedicels ; in large terminal
panicles.
Perianth-segments not clawed, glandless or
nearly so, sometimes adnate to the base
of the ovary.
Stamens shorter than the perianth-segments
and free from them.
Seeds very flat, broadly winged.
TS Ue CAE
Eu Res a ii. d MER T
MELANTHACEA FROM THE GENETIC STANDPOINT. 163
We may now consider the synonymy of the three species of Melanthium,
together with that of Veratrum parviflorum, Michx., which has often been
included in the same genus. It will be seen tist M. monoicum, Walter,
replaces M. dispersum, Small, and M. hybridum, Walt., takes the place of
M. latifolium, Desrouss. The reasons for these changes are as follows: Walter,
in the * Flora Caroliniana,’ recognizes the three known species of Melanthium
under a special section of his genus Melanthium which is thus described :
* Petalis unguiculatis imprimis albis demum obscuro-rubris seminibus semi-
ovatis," The first species is M. virginicum, Linn., the second he describes as
M. hybridum, and the third as M. monoicum. Let us consider M. monoicum
first. It is described as follows: “ Petalis planis, maculis 2 luteis ; floribus
inferioribus masculis majoribus, paniculis lateralibus; superioribus foemineis
racemo terminali.” Pursh describes the species as follows: ** M. panicula
inferne mascula, superne feminea racemosa, petalis oblongis planis brevi-
unguiculatis bimaculatis, stylis germine duplo brevioribus. Walt. Fl. Car.
195." He adds that the flowers are smaller than in M. virginicum, and not
so apt to change colour. Elliott merely repeats the description of Pursh.
There can be no doubt concerning the identity of this species with the
M. dispersum of Small. Particularly the oblong flat petals with two glands
and a short claw, and the flowers smaller than in M. virginicum, serves to
identify the species; also the broad panicle and the fact that the upper
flowers are mostly pistillate. This identification clears up a long-standing
difficulty with Walter's M. monoieum. Assurance is made doubly sure by the
detailed description of Leimanthium monoicum, Schult., in which measurements
and numerous other details are given.
When M. monoicum, Walt., has been identified, the remaining synonymy
falls easily into place. The identity of M. Aybridum, Walt., is not quite so
conclusive as that of M. monoicum, but that it is the same as M. latifolium,
Desrouss., there can be very little doubt. Britton * has referred it there
with a question mark, which I think must be removed now that Walter's
M. monoicum is recognized.
The description of M. hybridum by Walter is in these words: “ petalis
plicato-undulatis immaculatis, floribus masculis et fceminis mixtis." The
word *immaeulatis" has been the source of trouble, for the other two
features, (1) undulate petals and (2) staminate and pistillate flowers inter-
mingled, are peculiarities of this species, while glands are usually present on
the perianth-segments as in other species of Melanthium. The description of
Pursh, however, leaves no possible doubt as to the plant intended, for he
says under M. hybridum : * M. panicula superne racemosa feminea, petalis
subrotundis plieato-undulatis vix maculatis, extus hirsutis." He gives as
* Britton & Brown, Ill. Fl, i. (1896) 407.
LINN, JOURN.—BOTANY, VOL. XLIV. Oo
TOTS Ce ee TS ON TIT ENERO
164 DR. R. R. GATES: A STUDY OF NORTH AMERICAN
synonyms M. latifolium, Lam. Encycl. iv. p. 25, and M. racemosum, Michx.
Fl. Amer. i. p. 251. He further states that the flowers are “small, pale
white, and do not change." That Michaux's M. racemosum is the same plant
is shown by the character of the petals in his description: * panicula recemoso-
oblonga, confertiuscula ; laciniis calycinis orbiculatis.”
Elliott, under the name M. hybridum, Walt., also clearly describes this
species, his diagnosis being as follows: ^ M. panicula pubescente, racemosa ;
petalis orbiculatis, plicatis, longe unguiculatis ; glandulis coalitis.’ He
further says, * glands forming an emarginate circle, at the summit of the
claw, with a furrow along the centre,” which exactly describes the usual
condition in this species.
In the absence of glands, Nuttall concluded that M. hybridum, Walt., must
be the plant now known as Veratrum parviflorum, Michx., which Nuttall
clearly describes under the name M. hybridum. But Asa Gray, in his
description of M. hybridum, Walt., under the name Leimanthium hybridum,
mentions that the glands are sometimes obsolete. It is evident, then, that
Walter founded his description on plants of this character. Gray’s
full description is as follows : “ Foliis lineari-lanceolatis, elongatis ; perianthii
foliolis anguste unguiculatis, lamina rhomboideo-suborbiculata, margine
undulata; glandulis conniventibus (quandoque obsoletis); unguibus convoluto-
canalieulatis, infra medium staminiferus." He evidently gave the species
careful study, for he deseribes * three forms of Leimanthium hybridum
(M. hybridum, Walt.) as follows :—
* a gracilis ; panicula sparsiflora, ramis plerisque simplicibus.
“ B robustior; foliis inferioribus plantagineis ; ramis imis paniculæ
compositis.
* y elata; ramis panicul: plerisque compositis, multifloris.”
A study should be made of these forms, which differ in the branching of
the panicles and in the foliage.
The Schultes’? description of L. hybridum adds further details. ‘There
remains, then, no reasonable ground for doubt that Pursh, Elliott, and
Asa Gray were right in their designation of M. Aybridum, Walt., which must
therefore be known under that name.
Gray was less fortunate in his characterization of Leimanthium monoicum.
His description is that of Veratrum parviflorum, while his synonymy refers
in part to M. monoicum. Kunth also made the mistake of describing
Veratrum parviflorum under the name Zigadenus monacus. Gray also
confused Melanthium virginicum, Linn., with M. hybridum, Walt., under the
name Leimanthium virginicum, while Kunth following him confused the same
forms under the name Zigadenus virginicus.
* A. Gray, in Ann. Lyc. New York iv. (1837) 116.
TOM
MELANTHACEH FROM THE GENETIC STANDPOINT. 165
17. VERATRUM, Tourn.; Linn.
l. VERATRUM PARVIFLORUM has been considered with the genus Melanthium
because its synonymy has been entangled with that genus, standing as
it does in some respects intermediate between these two genera. We
may now consider the remaining species of Veratrum.
2. VeRATRUM VIRIDE, Ait. Hort. Kew. iii, (1789) 422; Bigel. Amer. Med.
Bot. ii. (1818) 121, t. 33.
V. album, Michx. Fl. Bor. Am. ii. (1803) 249, non Linn.
Helonias viridis, Ker, in Bot. Mag. (1808) t. 1096.
In swamps and wet woods, New Brunswick and Quebec to Ontario, south to
Georgia and Tennessee, west to Minnesota.
3. Veratrum Wooprr, Robbins, in Wood, Classbook, ed. 2, 557 (1855).
Dry woods and hills, S. Indiana to Missouri, Illinois, and Iowa.
Specimens: Hancock Co., Ill, S. B. Mead, 1842. (resembles V. inter-
medium). Desmoines River opp. Portland, Iowa Terr., Chas. A. Geyer, 1841.
4. VERATRUM INTERMEDIUM, Chapm. Flora S. Un. St. (1860) 489.
Middle Georgia to middle Florida.
Specimens : Gadsden Co., Fla., Chapman, 1837. Foot of Stone Mountain,
De Kalb Co., Ga., Eggert, 1897 (resembles V. Woodii).
5, VERATRUM CALIFoRNIcUM, Durand, in Journ. Phila. Acad. iii. (1855)
103; Piper, Fl. Washington, Contrib, U.S. Nat. Herb. xi. (1906) 196,
pl. 20.
V. album, S. Wats. Bot. King’s Exped. 344.
V. speciosum, Rydb. in Bull. Torr. Bot. Club, xxvii. (1900) 531.
Washington State to California, Colorado, and Montana.
6. VERATRUM FIMBRIATUM, A. Gray, in Proc. Amer. Acad. vii. (1868) 391.
Calif. (Mendocino Co.) along the coast.
7. VERATRUM CAUDATUM, Heller, in Bull. Torr. Bot. Club, xxvi. (1899) 588.
Western Washington.
8. Veratrum Jonesu, Heller, in Muhlenbergia, i. (1905) 124. Idaho.
9. VERATRUM TENUIPETALUM, Heller, l. c. i. (1904) 39. Colorado.
10. Veratrum Esconscnorrzit (Schult. f.), A. Gray.
V. Eschscholtzit, A. Gray, in Ann. Lyc. New York, iv. (1837) 119.
V. Lobelianum, B. Eschscholzianum, Schult. f. Syst. vii. (1830) 1555.
V. parviflorum, Bongard. in Mém. Acad. St. Pétersb. ser. 6, ii. (1831) 166.
Alaska and British Columbia.
Specimen: Rodman Bay, Alaska, 56, Mrs. K. Stephens, Aug. 15, 1907,
Univ. Cal. Herb. 127687.
166 DR. R. R. GATES : A STUDY OF NORTH AMERICAN
The poisonous genus Veratrum contains some 17 species in the north
temperate zone. In northern Asia are found V. album, Linn., and V. nigrum,
Linn., and several other species. Of the North American species V. parvi-
lorum, V. viride, V. intermedium, and V. Woodii may be ranked as eastern
species. V. parviflorum, Michx., is frequently included in Melanthium, but
in the absence of glands on the perianth-segments and in view of its close
relationship to V. intermedium and V. Woodii there is no doubt that it is
properly excluded from Melanthium.
In distribution, V. viride is spread most widely, extending west to
Minnesota in wet habitats. V. parviflorum occurs in drier localities on
mountains from Virginia to South Carolina, V. intermedium in Georgia and
Florida, while V. Woodii occurs farther west, in Indiana, Illinois, Missouri,
and Iowa. Itmust be said, however, that V. intermedium and V. Woodit are
particularly difficult to separate, and this difficulty is enhanced by finding
from Georgia certain specimens (Foot of Stone Mountain, De Kalb Co.,
Georgia, H. Eggert, July 23, 1897) verging on V. Woodii and from Illinois
a specimen (Hancock Co., Ill, S. B. Mead, July, 1842) practically
inseparable from J). intermedium. The differences between the species
in their typical condition are as follows :—
V. Woodii : Perianth-segments broad, purple; pedicels typically very
short, 2" or less.
V. intermedium : Perianth-segments narrower, dark brown within ;
pedicels 3"—4" long.
On the other hand, V. intermedium comes nearest V. parviflorum, from
which it is clearly separated, however, as follows :—
V. intermedium : Stamens nearly as long as perianth-segment, ovary
pubescent.
V. parviflorum : Stamens less than half as long as the perianth-segment,
ovary smooth.
In the two features of V. intermedium above mentioned it agrees with
V. Woodü. V. viride is much stouter, with broad elliptical leaves, in which
it agrees with the western V. californicum.
The differences between V. viride and V. Woodii are shown in the
following table :—
V. viride. V. Woodit.
Stem stout, very leafy to the top. Stem slender, sparingly leafy.
Leaves broadly oval, sheath-clasping. Leaves oblanceolate, only the lowest
sheathing.
Panicle pyramidal, Panicle very narrow.
Perianth yellowish green, segments ciliate- Perianth greenish purple, segments entire
serrulate, pubescent. glabrous.
Ovary glabrous. Ovary tomentose.
Flowers 15-25 mm. broad. Flowers 19-16 mm. broad.
UU TNNT ae EE
MELANTHACEJ FROM THE GENETIC STANDPOINT. 167
The seven western species are as yet imperfectly known. V. fimbriatum, A.
Gray, the most striking species, having fimbriate perianth-segments with
two glands at the base and an obtuse, obovate, few-seeded capsule, is found
only along the coast of California in Mendocino Co. We may suppose that
the fimbriate character at any rate originated through a mutation. Plants
belonging to V. californicum, Durand, and V. speciosum, Rydb., have broad
perianth-segments, acute with a green V-shaped basal mark, or obtuse and
without this mark, but the difference does not appear to be constant.
V. caudatum, Heller, and V. tenuipetalum, Heller, have narrow segments,
2-3 mm. wide in the former and 1 mm. wide in the latter. The chief peculiarity
of V. caudatum is the greatly elongated central rhachis of the inflorescence.
It may be doubted whether these two species are well founded. Heller, in
Muhlenbergia, i. (1905) 120, has formed a key of the seven western species
he recognizes, but neither the characters nor the distribution of these species
are as yet sufticiently well known to warrant further discussion. They all
appear to have been derived from the viride type, perhaps through
V. californicum.
18. SCHOINOLIRION, Torr.
1. ScHaNoLIRION cRocEUM (Michx.), A. Gray.
S. croceum, A. Gray, in Amer. Nat. x. (1876) 427.
Phalangium croceum, Michx. Fl. Bor. Am. i. (1803) 196; Nutt. Gen. i. (1818) 220.
Anthericum croceum, Schult. Syst. Veg. vii. (1829) 476.
A. Nuttallianum, Schult. l. c. (1829) 477.
Oxytria crocea, Raf. Fl. Tellur. ii. (1836) 26.
Ornithogalum croceum, Kunth, Enum. Pl. iv. (1843) 371.
O. Nuttallianum, Kunth, l. c. (1843) 372.
Georgia to Florida and Louisiana, in sandy soil.
bo
. ScH@NOLIRION TEXANUM (Scheele), A. Gray.
S. texanum, A. Gray, in Amer. Nat. x. (1876) 427.
S. Michauxii, Torr. Bot. Mex. Bound. (1859) 220, partim.
Ornithogalum texanum, Scheele, in Linnea, xxiii. (1850) 146.
Oxytria texana, Pollard, in Bull. Torr. Bot. Club, xxiv. (1897) 407.
Texas and Louisiana, on prairies.
3." SCH@NOLIRION ALBIFLORUM (Rafin.), comb. nov.
S. Michauzii, Chapm. Fl. S. States (1860) 483.
S. Elliottii, Feay, ex A. Gray, in Amer. Nat. x. (1876) 427.
Ornithogalum croceum, Elliott, Sketch, i. (1821) 397, non Michx.
Amblostima albiflora, Ratin. Fl. Tellur. ii. (1836) 26.
Anthericum croceum, Baker, in Journ. Linn. Soc., Bot. xv. (1876) 297, non Schult.
Oxytria albiflora, Pollard, in Bull. Torr. Bot. Club, xxiv. (1897) 406.
Southern Georgia and Florida, in low pine lands.
The genus Schwnolirion, being one of the genera conservanda of the Vienna
Congress, is retained. Otherwise it would be superseded by Rafinesque's
PCM era? ELEM Oe A 11 a COT TT TL EL EE MIA e RS WEIT
CRAT T SU ai aaa da NT TATEM SM ME MM S UC TV
168 DR. R. R. GATES: A STUDY OF NORTH AMERICAN
name Owytria. There are only three species, S. eroceum and S. albiflorum
occupying much the same limited range in the south-eastern States, while
S. texanum is farther west but overlaps |S. eroceum in Louisiana. The
differences between these species may be regarded as typical specific
differences, the three species forming a rather compact group. ©. eroceum
differs from the other two species in having (1) bright yellow instead of
white or greenish-white flowers, and (2) obtuse instead of acute or acuminate
bracts. The other two species are differentiated as follows * :—
S. texanum.
Leaves narrowly linear, 1-4 dm. long.
Scapes as loug as the leaves, simple below
the inflorescence.
Pedicels 8-14 mm. long.
Perianth greenish white, 1 em. broad.
Segments oblong, obtuse, 3-nerved.
Seeds 2 mm. long.
S. albiflorum.
Leaves narrowly linear, 4-6 dm. long.
Scapes longer than the leaves, paniculately
branched above.
Pedicels 10-25 mm. long.
Perianth white, 10-12 mm. broad.
Segments broadly oblong or oval, obtuse,
mostly 5-nerved.,
Seeds 4 mm. long.
S. albiflorum is thus a sturdier species with longer branching scapes,
longer pedicels, somewhat larger flowers, and larger seeds, These differences
suggest very much the contrasts between GZnothera Lamarckiana and
CE. gigas, and this is another pair of species which may be suspected
of tetraploidy or cell-gigantism. It is to be hoped that someone will make
a cytological study of these three species.
Anthericum Nuttallianum, Schult., is supposed to differ from S. eroceum
chiefly in having the basal portion of the raceme interrupted, with the pedicels
about an inch apart. I have seen specimens having this feature. The form
should be carefully studied in the field or from a large suite of specimens
to determine whether it represents a distinct species. Its segregation by
Schultes is probably to be attributed to the error of Michaux in stating the
inflorescence of his Phalangium croceum to be pyramidal.
In his S. Michauwii, Torrey really combined the three species of Schanolirion
into one.
19. HASTINGSIA, S. Wats.
The western genus /astingsia is a segregate from Schenolirion, to which it
is not, however, closely related. It contains two species as follows :—
1. Hasriyesta ALBA (Durand), Wats.
H. alba, S. Wats. in Proc. Amer. Acad. xiv. (1879) 242.
Schenolirion album, Durand, in Journ. Phila. Acad, ii. 111. (1855) 108.
N. California and S. Oregon.
* These characters are taken mostly from Small; Fl, S.E. Un. St. (1903) 268.
"E
bue urbc PR ee
MELANTHACEJE FROM THE GENETIC STANDPOINT. 169
2. Hastinesta BRAOTEOSA, S. Wats. in Proc. Amer. Acad. xx. (1885) 377.
S. Oregon, in marshes near Waldo.
Specimen: Kerbyville, Ore., /7owell, 1884.
H. bracteosa is said to differ from H. alba in its longer leaves, more
` conspicuous bracts, flowers twice larger, and shorter stamens. "The last two
characters are the only constant ones in the specimens I have seen. The two
species are contrasted by Howell * as follows :—
H. alba.
Leaves 4-12 mm. wide, attenuate above.
Raceme densely many-flowered.
Bracts narrow acuminate.
Pedicels 2-4 mm, long, shorter than the
bracts.
Perianth -segments about 6 mm. long,
oblong, obtuse, white or tinged with
green or pink.
Stamens about equalling the perianth.
H. bracteosa.
Leaves 2-8 mm. wide, acuminate above.
Raceme loosely many-flowered.
Bracts filiform-subulate from a broad base.
Pedicels ascending, about 4 mm. long.
Perianth-segments 8-10 mm. leng, lanceo-
late, acuminate, dull white.
Stamens half as long as perianth-segments.
This pair of species also requires a careful comparative examination.
H. bracteosa is apparently very limited in range and may perhaps be a
mutation of recent origin.
The generic characters are very well marked. The nearest relatives
of Hastingsia are Schenolirion and Chlorogalum. The differences from the
former genus may be tabulated as follows :—
Schanolirton.
Seapose herbs with bulb-like rootstocks.
Leaves basal, often numerous.
Leaves very narrow, flat, rather rigid.
Flowers in loose, simple or sparingly
branched racemes, white or yellow.
Perianth-segments ^ withering- persistent,
not scarious, 3-5-nerved, exceeding the
stamens.
Anthers versatile, introrsely dehiscing.
Style short.
Pedicels slender, exceeding the flowers and
bracts.
Capsule depressed globose, deeply lobed.
Hastingsia,
Perennial herbs with scape-like sparingly
branched stems from coated bulbs.
Leaves basal, rather numerous.
Leaves narrow, flat, more fleshy.
Flowers in densely many-flowered sparingly
branched racemes, white or greenish,
Perianth-segments lax and scarious, per-
sistent, closely 3-nerved (appearing
l-nerved), equalling the stamens.
Anthers versatile.
Style short.
Pedicels stout, much shorter than the
flowers and bracts.
Capsule oblong-ovate, not deeply lobed,
very shortly stipitate.
* Howell, T., in Fl. N.W. Amer. (1902) 647,
170 DR. R. R. GATES : A STUDY OF NORTH AMERICAN
Since Schenolirion, Hastingsia, and Chlorogalum form a group of three
allied genera we may ‘compare the last two as follows :—
Hastingsia.
Bulbs white-coated.
Leaves narrower.
Perianth-segments lax and scarious, closely
3-nerved.,
Style short.
Flowers white or greenish, in densely
many - flowered sparingly panicled
racemes,
Anthers versatile.
Chlorogalum.
Bulbs with membranous or densely fibrous
coats.
Leaves with undulate margins.
Perianth-segments at length twisted over
the ovary, distinctly 3-nerved.
Style long, deciduous,
Flowers white, pinkish or purplish, in loose
paniculate racemes,
Anthers versatile.
Of these three genera, Schwnolirion occupies the Southern States from
Florida to Texas, while Chlorogalum occurs in the greater part of California,
and //astingsia is found in Northern California and adjacent Oregon. Itis
difficult to trace the steps by which these genera have become separated.
Hastingsia appears to be more closely related to Schanolirion despite its
western position. In such cases, as in those of bifurcated genera with
eastern and western species, it seems necessary to assume that the ancestors
originally occupied a more northerly position and that the eastern and
western forms became separated as they were driven southward by the ice.
In the central plains area, plants of this group failed to find suitable habitats,
at least after the plains took on their present aspect, and so only eastern and
western forms, variously modified, have persisted to the present time.
Another monotypic genus, represented by Hemiphylacus latifolius, S. Wats.*,
in Northern Mexico, belongs with this group. Among its peculiarities the
most interesting are (1) the tuberous roots, (2) the filaments adnate to the
perianth-segments for half their length, (3) only the inner and shorter ones
bearing anthers, which are versatile, rounded, introrse, and (4) the scarious
l-nerved perianth-segments.
20. CHLOROGALUM, Kunth.
l. CHLOROGALUM POMERIDIANUM (Ker), Kunth.
C. pomeridianum, Kunth, Enum. iv. (1843) 682.
C. divaricatum, Kunth, l. c.; Torr. Bot. Mex. Bound, (1859) 218, t. 60.
Scilla pomeridiana, DC. in Redouté, Lil. (1816) t. 421.
Anthericum pomeridianum, Ker, in Bot. Reg. (1821) t. 564.
Phalangium pomeridianum, D. Don, in Sweet, Hort. Brit. ed. 1, 416 (1827).
Ornithogalum divaricatum, Lindl. Bot. Reg. xxviii. (1842) t. 98.
Laothoe pomeridiana, Rafin. Fl. Tellur. iii. (1886) 53.
L. divaricata, Greene, Leaflets Bot. Obs. i. (1904) 91.
California, from Mendocino Co. southward to San Bernardino Co.
* [n Proc. Amer. Acad. xviii. (1888) 164.
MELANTHACEJE FROM THE GENETIC STANDPOINT. igl
2. CHLOROGALUM PARVIFLORUM, S. Wats. in Proc. Amer. Acad. xiv. (1879)
243.
Laothoe parviflora, Greene, Leaflets Bot. Obs. i. (1904) 91.
S. California, San Diego Co.
3. CHLOROGALUM ANGUSTIFOLIUM, Kellogg, in Proc. Calif. Acad. ii. (1863)
105, t. 30.
Laothoe angustifolia, Greene, Leaflets Bot. Obs. i. (1904) 91.
Middle California, Sacramento Valley, and Mendocino Co.
4. CHLOROGALUM PURPURECM, T. S. Brandegee, in Zoe, iv. (1893) 159.
Laothoe purpurea, Greene, Leaflets Bot. Obs. i. (1904) 91.
California, Coast Range, Monterey Co. southward.
The genus CAlorogalum contains four species and is practically confined
to California. The following key, compiled by Mr. C. H. Thompson, shows
the nature of the specific differences :—
A. Bulb large, thiekly coated with coarse brown fibres; perianth-segments narrowly
ligulate, spreading widely from the base in the open flower. C. pomeridianum
(Ker), Kunth.
B. Bulbs smaller, with membranaceous coat; perianth-segments spreading from above
the base.
a, Perianth-segments oblong-ovate; pedicels as long as or longer than the perianth ;
flowers purplish blue. C. purpureum, T. S. Brandegee.
b. Perianth-segments oblong-lanceolate ; pedicels very short.
a. Flowers pinkish ; ovary broad and obtuse. C. parviflorum, S. Wats.
B. Flowers white with yellowish-green lines; ovary oblong-ovate, acutish above,
short stipitate. C. angustifolium, Kellogg.
In C. pomeridianum the branches are spreading, the stems dark brown, and
the leaves remarkably undulate. C. angustifolium differs in having narrow
linear-lanceolate leaves, slightly undulate, and a bulb which is ovo-conie
rather than long-ovate.
C. pomeridianum and C. angustifolium apparently have much the same
range in central California, the latter occurring chiefly farther inland in the
Sacramento Valley, while the former extends farther northward into Oregon.
The other species are more local in distribution and probably more recent in
origin, as they are in discovery. C. parviflorum occurs in the extreme south,
in San Diego Co., and C. purpureum in the Coast range of Monterey Co.
This would be an interesting genus for experiments in cultivation, but
until such are instituted it is superfluous to comment further on the specific
differences exhibited, except to say that there is an unusual range of colour
variation for so small a genus.
21. ODONTOSTOMUM, Torr.
A monotypie genus of western North America.
OpoxrosroMUM Harrweeu, Torr. in Pacif. R. R. Rep. iv. (1856) 150, t. 24;
Baker, in Journ, Linn. Soc., Bot. xi. (1870) 436.
LINN. JOURN.—BOTANY, VOL. XLIV. y
172 A GENETIC STUDY OF NORTH AMERICAN MELANTHACE.E.
In California, from Shasta Co. to Napa and Amador Co., chiefly in the
Sacramento Valley. Said to be rare.
Specimens: Near Napa, Calif., A. W. Robinson, 1892. Redding, Shasta
Co., California, A. A. Heller, May 26, 1905.
Odontostomum, while most nearly related to the group of Chlorogalez, is
rather isolated, especially in flower-structure, though the foliage and habit
clearly resemble that group. Of the three genera it is nearest Chlorogalum.
The flowers are small and yellowish and the base of the stem forms a fibrous-
coated corm. The deciduous perianth of Odontostomum in the form of a
cylindrical 12-nerved tube, bearing in salver-form and finally reflexed the
5-nerved segments, is a new feature in this group. Our experience of
mutations is as yet too limited to permit an understanding of the way
in which the development of this organ took place. The perianth-segments
are obtuse and the three outer ones cucullate at the tip.
Another striking peculiarity is in the stamens. There are 6 stamens with
subglobose anthers at the summit of short filaments, and alternating with
them are 6 staminodia, the whole forming a ring at the top of the perianth-
tube*. In this feature Odontostomum bears some resemblance to the
Mexican Hemiphylacus, though in the latter there are only 3 stamens and
3 (longer) staminodia. Evidently several definite variations occurred to
produce the Odontostomum flower, but what their sequence has been it is
impossible to say. In connection with the stamens there has evidently been
a loss mutation in Hemiphylacus (see p. 170), while there must have been a
positive mutation in Odontostomum.
The fact of parallel mutations introduces an important principle into
conceptions of phylogeny. Thus Chlorogalum and its relatives have versatile
anthers, but it is not necessary to assume that this feature has been handed
down from a common ancestor of Chlorogalumand Lilium. It is much more
probable that the versatile condition has been developed independently in the
Chlorogaleæ and the Lilieze through parallel mutations.
KEY TO MAP, PLATE 5.
GENERA OF Melanthacee.
. Tofieldia. 2. Triantha. 3. Pleea. 4. Narthecium. 5, Xerophyllum. 6, Helonias.
7. Chamwlirium. — 8. Amianthium., 9. Scheenocaulon. — 10. Stenanthium.
11. Stenanthella. 12. Zigadenus. 13. Anticlea, 14. Toxicoscordion. 15. Oceanorus,
16. Melanthium. 17. Veratrum. 18. Schcnolirion, 19. Hastingsia. 20. Chloro-
galum, 21. Odontostomum.
* According to Jepson [in Fl. W. Mid. Calif. (1901) 114 * the stamen opposite the lower
outer segment stands alone and faces the remaining 5, which approximate each other by
their filaments on the upper side of the flower.”
GATES. JOURN. LINN. SOC., Bor. VoL. XLIV. PL. 5.
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DISTRIBUTION OF NORTH
AMERICAN MELANTHACEZE.
Ads co fige Piodecnd ide Mice ts al Lagu; or i
DISTRIBUTION OF DIATOMS AND COPEPODA IN THE IRISH SEA. 173
Spora RuNIANA.—IIT. The Distribution of certain Diatoms and Copepoda,
throughout the year, in the Irish Sea. By W. A. Herpmay, F.R.S.,
F.L.S., Professor of Zoology in the University of Liverpool.
(With 21 Text-figures.)
[Read Ist November, 1917.)
Part of the work * of the yacht ‘Runa’ for some years previous to 1914
consisted in taking periedic samples of the marine plankton at various
localities around Port Erin, at the south end of the Isle of Man, during the
two most interesting times in the annual cycle—viz., spring (March-April)
and autumn (July-September). During the remaining months, when the
yacht was not in commission, plankton gatherings in Port Erin bay were
taken with great regularity at the rate of six in the week, three at a time on
two occasions per week, two of the three hauls being horizontal and the
third vertical. This systematic plankton survey has been continued for fully
10 years (1907-1917 inclusive), and over 5000 + samples have been collected
and examined. ‘The general results of this intensive study of the plankton of
a central area of the Irish Sea have been given in a series of reports $ drawn
up in collaboration with Mr. Andrew Scott, A.L.S., and others, and published
by the Lancashire and Western Sea-Fisheries Committee; but the material
and statistics collected still contain much information which has not yet
been made use of. It is proposed in the present communication § to deal
with the records of the occurrence throughout the year in our district of a
few of the most abundant of the Diatoms and the Copepoda which make up
the bulk of the phytoplankton and of the zooplankton respectively at those
periods of the year when they are most abundant. At the time of the
spring maximum (usually in April or May) a small silk tow-net hauled for
about 15 minutes through about half-a-mile of the surface water of the Irish
Sea will usually catch some millions of individual Diatoms (up to a couple of
hundred millions || on occasions), constituting probably, on the average, some
999,999 out of each million of organisms in the gathering €. This is almost
* For Parts I. and IT. of *Spolia Runiana" see Journ. Linn. Soc., Zool. xxxii. p. 163
(1913), and p. 269 (1914).
T More precisely 5116, to the end of 1916,
t Trans. Biol. Soc. Liverpool, xxii. (1908) to xxxi. (1917).
§ I wish to acknowledge, with thanks, the help I have received in the preparation of
these plankton records from Mr. Andrew Scott, A.L.S., and from my secretary, Miss
H. M. Lewis, B.A. Mr. Scott took for me the excellent photo-micrographs of the plankton
from which most of the illustrations have been reproduced.
|| Estimated by counting measured samples.
«| The average of a number of cases where smaller, but still very large, hauls of Diatoms
were taken is—Diatoms=about 99 per cent. of the total organisms present.
LINN. JOURN.—BOTANY, VOL. XLIV, Q
174 PROF. W. A. HERDMAN ON THE DISTRIBUTION
a pure sample of Diatoms—a “ monotonic phytoplankton.” Similarly, when
the zooplankton is at its height in late summer (usually September) the same
net may contain almost a pure gathering of Copepoda numbering some tens
to hundreds of thousands of individuals (up to 214,000), and making up
perhaps 999 out of every thousand organisms presenta “monotonic
zooplankton." But we may still regard the gathering as a zooplankton if
over 50 per cent. of the organisms are animals—on account of their greater
bulk.
Moreover, these very abundant Diatoms and Copepoda belong in each ease
to very few species, so that one can select about half-a-dozen species of
Copepoda which constitute by far the greater part of the summer zooplankton,
and about the same number of Diatoms whieh similarly make up the bulk
of the spring phytoplankton. These few species, belonging to these two
very widely separated groups, thus come to be the most significant organisms
~
Fra. 1.—“ Hensen,” “ Nansen,” * Funnel” and other plankton nets
drying after use on the yacht.
in relation to the annual metabolic cycle of our seas and the food-supply
from our coastal fisheries, Consequently it is of both scientific and economic
importance to obtain such data as seem possible from our long series of
observations, extending over a decade, as to the occurrence of these dominant
factors in the plankton. No doubt there are exceptional years with unusual
occurrences which will have a disturbing effect, but the ten or eleven years’
results ought to give us an average of some value.
We have endeavoured, in our work from the yacht, as the result of many
experiments, to make use of a standard net in a constant manner so that the
hauls should be approximately comparable. Our two horizontal tow-nets of
fine-meshed miller’s silk (No. 20 and No. 9) measure 35 em. (about 14 inches)
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 175
in diameter at the mouth, and are in our weekly gatherings throughout the
year hauled through a definite course in the open water of Port Erin bay.
The two together (* coarse? and “ fine” nets) constitute a “standard haul."
The gatherings taken outside the bay during the periods of the vernal and
autumnal plankton maxima were made with the same nets as in the bay, at
fixed “stations” respectively three and five (and on a few occasions ten)
miles from land *.
In addition to these standard hauls many others were taken with special
nets, such as the closing * Petersen-Hensen " and “Nansen,” the “ shear-
net” and “funnel-nets,” usually at depths of 5 to 30 fathoms, and on occasions
down to 60 fathoms out in the deep central valley between the Isle of Man
QC b
40 A
N
/ N
35 V. wm. y
i
25 }
^
E
Sets o ipee
E
N
\
i
i5 \
\
/ bel p
\
JAN Fes Mar Arc May June Juw Aucr Sep Ocr Nov. Dec
Fra. 2.—Typical Irish Sea plankton curves for the years 1913 (dotted line)
and 1914 (whole line).
and Ireland. Figure 1 gives a general view of the various nets used on one
of our plankton cruises. Temperatures, surface and deeper, weather observa-
tions and water-samples were also taken systematically f.
The plankton gatherings when taken are at once treated with formol to
kill and fix the organisms and prevent further changes. They are then
placed (at the Port Erin Biological Station) in graduated cylinders, and
after some hours, when the material has fallen to the bottom and the super-
jacent fluid is clear, the quantity of plankton in cubic centimetres is
recorded. The fixed plankton is then concentrated and preserved in
* Determined by cross-bearings, and by distance run calculated from the engines,
T For further particulars as to methods, see the earlier reports published by the Lanca-
shire and Western Sea-Fisheries Committee.
Q2
176 . PROF. W. A. HERDMAN ON THE DISTRIBUTION
5 per cent. formol, and is re-measured it may be weeks later when
submitted to detailed examination. It is this last more accurate measure-
ment that has been made use of for calculations and curves. The
first estimation of the volume is only taken in case some accident happens
later ; but we have been very fortunate in that respect: we have only lost
about 10 gatherings in the 10 years out of over 5000 samples. Figure 2
shows typical plankton curves for the years 1913 (dotted line) and 1914,
taken from the recorded monthly averages of the plankton hauls. As an
example of the height to which individual hauls may rise in spring, I may
quote from our records of this year that on April 18th, 1917, the standard haul
measured 165 c.c, and on April 19th 150 c.c., and consisted mainly of
Chetoceras.
Our confidence that these samples are adequate and representative receives
support from the fact that the same organisms are recorded in much the
same quantities year after year, and that practically no new forms turn up.
Mr. Andrew Scott, A.L.S., who has made a detailed microscopic examination
of all the material for the purpose of determining the species, has met with
none new to science. Some rare species previously unknown in British
seas, such as the Norwegian Copepod Mierocalanus pusillus, Sars, which
appeared first in our deeper nets in the summer of 1907, and the Indo-Pacific
Diatom Biddulphia sinensis, Grev., have occurred ; but throughout the series
of over 5000 gatherings, extending over nearly every week of 10 years,
no species actually new to science has been determined from the macro-
plankton. The various new Copepoda which have been described from time
to time from our work at Port Erin have all been bottom-living forms
obtained by dredging. This is, so far, a satisfactory result of our work, as
it seems to indicate that probably all the pelagic species of Copepoda in our
sea are now known. It is not the rare species that are of most interest.
They may have an interest of their own—morphological or distributional—
but for my present purpose it is the common species that are of most import-
ance, those species which by their abundance in nature play their part in
providing fish-food for man or in affecting the public health either by
keeping the sea clean or by causing plagues.
For the purpose then of arriving at some conclusion as to the distribution
throughout the year of these really significant organisms, I have picked out
from our records the following six species of Copepoda as being undoubtedly
the most abundant and economically the most important representatives of
that section of the plankton :— Oithona helgolandica (= similis), Pseudo-
calanus elongatus, Acartia elausi, Temora longicornis, Paracalanus parvus, and
Calanus jinmarchicus. These are all cases of genera where there is only one
species in our seas (e. g., Calanus) or only one common species (e. g., Oithona),
so that we are dealing with half-a-dozen very distinct forms, and there can
be no doubt as to what is in question even if the genus only is referred to,
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 147
When we turn to the Diatoms the case is rather different. There several
of the more prominent genera are represented by a number of common
species, and moreover some of the species are closely related, and variable,
so that doubts may arise as to the exact identifications, and authorities may
differ as to the relative proportions in which certain species or varieties of,
say, Chetoceras or Biddulphia are present in the plankton. Under these
circumstances I have considered it will best serve my purpose, which is a
general and economic rather than a detailed speciographic one, if I deal with
generic titles only, grouping together for example all the species of
Chetoceras that may occur under that one name. I have chosen the following
genera as being the most important representatives of the Diatoms in our
plankton :—Biddulphia, Chetoceras, Coscinodiscus, Rhizosolenia, Thalassio-
sira, Guinardia, and Lauderia. In some of these genera (e.g., (ruinardia
and Lauderia) as in the case of the Copepoda there is only one possible
species in question, in Biddulphia it is in most cases only the species B. mobi-
liensis, but in others (e. g., Chetoceras, Coscinodiscus, and Rhizosolenia) there
are usually several allied species occurring together in profusion in any large
gathering of the genus.
I may add that our commonest species in the Irish Sea off Port Erin are
not necessarily those that are most abundant in other seas of North-West
Europe. For example, in the Baltic near Kiel, according to Lohmann, the
most abundant Diatom is Skeletonema costatum, a comparatively rare form in
our plankton, and George Murray similarly found that to be the commonest
form he had met with in a plankton survey of some of the more sheltered
lochs of the West of Scotland. It is, however, one of the minuter forms
which readily escapes notice, and may to a considerable extent pass through
the meshes of the net.
Then again, in July 1911, in Upper Loch Torridon, on the West Coast of
Scotland, I got a haul of 334,000,000 Nitzschia delicatissima, which is rare
with us in the Irish Sea, but is apparently more abundant at Plymouth.
I think it probable, however, that our Port Erin results will be found to hold
good for the more open sea-water of high salinity * around the British Isles. A
valuable paper which appeared recently on the Plankton of Plymouth Sound,
by Dr. Marie V. Lebour t, while dealing mainly with the more minute
Protozoa and Protophyta which escape the tow-net and can only be obtained
by centrifuging samples of water, gave also some records of the occurrence
of some of the larger forms which enables a comparison to be made between
the plankton conditions in the English Channel and in the northern part of
the Irish Sea.
* The salinity off Port Erin averages about 34:2 per mille. Its range for April, May, and
June in the year when we took the most complete series of observations is from 34:02 to
34:4 per mille.
+ Journ. Mar. Biol, Assoc. vol. xi. 2 (1917), p. 138.
178 PROF. W. A. HERDMAN ON THE DISTRIBUTION
There are certain differenees in detail. For example, the total Diatom
curve at Plymouth has three maxima or crests, in April, August, and October.
At Port Erin the curve has only two erests, a much greater maximum in
spring and a variable and smaller one in autumn, while Diatoms are usually
wholly absent in August.
On the other hand, there is a general agreement in regard to the distribution
throughout the year of many of the more abundant organisms. For example,
amongst Diatoms Coscinodiscus is a winter and early spring form, Biddulphia
flourishes throughout the winter from November to April or May, Rhizosolenia
is a summer form having its maximum in June, while Chetoceras and
Lauderia have two maxima, the one in spring and the other in autumn, in the
English Channel and the Irish Sea alike. Amongst Copepoda there seems
to be a general agreement along with a certain amount of difference in detail
which will be referred to below when discussing the species.
I may recall that in November 1910 I read a paper before this Society *
in which I made a comparison between the summer (July) plankton on the
West Coast of Scotland and that of the Irish Sea, showing that in some of
the deep fjord-like highland sea-lochs green-coloured phytoplankton can be
obtained even in the height of summer, while a zooplankton may be found
living in abundance a few miles away. This, of course, would be impossible
in the Irish Sea, where a zooplankton and a phytoplankton do not occur
simultaneously.
DIATOMS.
The seven generic forms I have selected for consideration taken together
make up nearly the whole of the Diatom plankton of the year. No other
genus occurs in anything like such profusion as these. In April, for example,
when the Diatoms are usually at their climax, all the remaining genera
(at most 10 or 12) taken together make up only about one-thousandth, or less,
of the whole. Moreover, these common Diatoms often attain their greatest
profusion successively, not. simultaneously, so that single genera, or it may
be single species of a genus, make up on occasions the bulk of the phyto-
plankton. For example, in May 1916 the month's average haul of Diatoms
was 7,171,789, while the average for the genus Chwtoceras taken alone was
6,947,333, leaving only 224,456 as the average of all the rest of the Diatoms.
On the last two individual hauls, taken on May 25th and 29th, the actual
numbers were as follows :—
Chetoceras alone... ie 24,094,500 de 19,461,600
C. sociale alone bis A 23,936,000 5s 19,396,000
All other Diatoms together... 166,300 ne 228,900
So that on these occasions, and they are examples of many, one species makes
up nearly the whole of the plankton.
* Journ, Linn, Soc., Zool. xxxii. (1913) p. 23.
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 119
The maximum on the Diatom curve ranges from March to May. In 1907
it was in March, in 1908 in May, and in 1909 in April. In some years the
Diatom maximum may be divided into two parts, an earlier due mainly to
Chetoceras and Thalassiosira, and a later in June due to Rhizosolenia and
Guinardia. A common order of succession for the species which contribute
most largely to the Diatom maxima is-—Biddulphia mobiliensis and
Coscinodiscus radiatus in early April, Chetoceras debile in late April,
Chetoceras sociale in May, Chetoceras teres and Rhizosolenia Shrubsolei in
early June, and Rhizosolenia (several species) and Guinardia in later June.
The autumn Diatom maximum is constituted mainly in the Irish Sea by
Chetoceras subtile and Rhizosolenia semispina. Certain species of most of the
genera we are dealing with are commonly regarded as “oceanic” in the
sense that they are characteristic of the open sea, although they may
occasionally be carried in shore and so form a part ef the coastal plankton ;
while other species are “neritic,” having their origin aad their home in
coastal waters and not being found normally in the open ocean. For
example, Chetoceras densum, C. boreale,- Coscinodiscus radiatus, and Rhizoso-
lenia. semispina are supposed to be oceanic; while Biddulphia mobiliensis,
Chetoceras subtile (and other species), Coscinodiscus concinnus, Rhizosolenia
setigera, and Lauderia borealis are probably neritie. Other species of these
genera are of doubtful position in this classification, or it may be are
* panthalassic " or equally at home in both regions.
BIDDULPHIA.
This is a winter and early spring group of Diatoms, generally appearing
in September or October, becoming more abundant in November and
reaching a maximum in March or April. It dies off during May, and is
practically absent in June, July, and August.
The species, or forms, that occur in our plankton are Biddulphia mobiliensis
(? B. regia) and B. sinensis (fig. 3). This is one of the cases where there is
some difference of opinion as to the validity of species. Whether B. regia
is only a form of B. mobiliensis, and what is the exact relation of B. sinensis
to the others, has been and may still be a matter of discussion. Specimens
can be picked out that seem distinct and characteristic, but others occur in
nature that are intermediate and possibly abnormal *.
B. sinensis is an exotic, oceanic form which, according to Ostenfeld, made
its appearance at the mouth of the Elbe in 1893, and spread during successive
years in several directions. It appeared suddenly in our plankton gatherings
at Port Erin in November 1909, and has been present in abundance each
year since. Ostenfeld, in 1908, when tracing its spread in the North Sea,
predicted that it would soon be found in the English Channel. Miss Lebour,
who has recently examined the old plankton gatherings at the Plymouth
** See forms figured in Trans. Biol. Soc. Liverpool, xxvii. (1913) p. 210.
180 PROF. W. A. HERDMAN ON THE DISTRIBUTION
Laboratory, finds that as a matter of fact this form did appear in abundance
in the collections of October 1909, within a month of the time when
according to our records it reached Port Erin. Whether or not this species
was brought accidentally by a ship from the far East, there is no doubt that
it was not present in our Irish Sea plankton gatherings previous to 1909,
but has been abundant since that year, and has completely adopted the
habits of its English relations—appearing with B. mobiliensis in late autumn,
Me
Fig. 3.—Photo-micrograph of a plankton preparation showing (a) Biddulphia
mobiliensis, forma “ regia,” and (b) B. sinensis.
persisting during the winter, reaching a maximum in spring, and dying out
before summer. | Biddulphia is generally the first Diatom to show a marked
increase in early spring, and is responsible for the moderate rise in the curve
which takes place in February or March.
Our largest records of Biddulphia are as follows :—
From three hundred thousand to over six hundred thousand per haul on
several occasions between middle of March and middle of April in 1910 and
1911, and towards end of April and beginning of May, 1916.
From three hundred thousand to over seven hundred and fifty thousand on
ten occasions between November 10th and 27th, 1911, 1914, and 1915.
The highest monthly averages in the early spring months occur as
follows :—
February ... in 1907 and 1912.
March — ... in 1910, 1911, 1914, and 1915.
April x in 1908, 1909, 1913, and 1916.
A second, usually slighter, maximum occurs in November, when the
numbers are higher than in December and January ; but on three occasions
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 181
(1911, 1914, and 1915) in our ten years the November monthly average is
the highest in its year. Moreover, two of these November records, viz.,
341,231 in 1911 and 406,100 in 1914, are the highest in the whole series.
So Biddulphia may be regarded as characteristic of the late autumn
(November) as well as the early spring plankton.
COSCINODISCUS.
The more abundant species that our records deal with are Coscinodiscus
concinnus, C. Grani, and C. radiatus. They are mainly winter and spring
forms, the maximum of the genus occurring always in our experience in
either March or April. Coscinodiscus (fig. 4) agrees very closely with
Biddulphia in its distribution throughout the year, beginning in late autumn,
maintaining its position throughout the winter, increasing in January or
Fr. 4.—Photo-micrograph of an early spring phytoplankton
consisting mainly of Coscinodiscus and Biddulphia.
February, and then more rapidly in March and April, and dying away before
the height of the summer; but it sometimes continues to be present in small
quantities further into the summer months than is the case with Biddulphia.
The presence of Coscinodiscus in quantity in the spring plankton is easily
seen through a glass jar with the unaided eye, as the little rounded drum-
like cells give a characteristic granular appearance to the gathering.
Our largest records of Coscinodiscus are :—
From four to five hundred thousand per haul on several occasions
between middle of March and middle of April, 1911, 1912, 1913, 1915,
and 1916.
About nine hundred thousand on several occasions at end of April 1914.
From one million to close on five millions on two occasions in middle of
April 1915.
So Coscinodiscus may outrival its companion form Biddulphia at the time
of the vernal maximum, but does not attain to such high numbers in late
182
PROF. W. 4. HERDMAN ON THE DISTRIBUTION
autumn.
The October and November monthly averages never, in our series
of years, come anywhere near the averages for March and April.
in the Irish Sea than in the English Channel, judging from the Plymouth
records.
Both Biddulphia and Coscinodiscus seem to be slightly later in their maxima
In the years 1907 to 1912, inclusive, the highest monthly averages (March
and April) for Coscinodiscus run in general about 100,000. In 1913 and
1916 they are elose on 200,000, in 1914 over 300,000, and in 1915 there is a
sudden jump, in April, to over 840,000. The other months of that year do not
show unusually high numbers
CH. ETOCERAS.
The chief species of this genus in our Irish Sea plankton are Chetoceras
boreale, C. contortum, C. crioplilum, C. debile, C. decipiens (fig. 6), C. densum,
April
8 millions
Oheetoce y=
Rhizosole ma
| million
1 Chaetoceras
1 \ Sept:
V M +
" \ M Rh ixosoleni a
—; Nm atu duci iir ioia io
Jan. Feb. Mar. Apr. May June July
]
Aug. Sept. Oct. Nov. Dec.
Fia. 5.— Curves of the more important Diatoms of ihe April and June maxima.
C. diadema, C. sociale, and C. teres. Of these, C. boreale, C. criophilum, and
C. decipiens are Arctic oceanic forms, C. densum is a temperate occanic or
Eid. hurt a Z2 LASXE os oos ut E
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 183
Atlantie species, and all the rest may be classified as temperate neritic.
Some of these (C. eriophilum, C. debile, C. decipiens, C. sociale, and C. teres) |
are spring forms with a maximum in April or May, while C. boreale and
C. densum are autumn species having their maxima in September or
Oetober. Consequently the genus is well represented throughout a con-
siderable part of the year, and the numbers are very high in April and May,
and sometimes also in September and October (see curve, fig. 5).
A few of our highest records for the genus, giving the nearest million
in each case, are :—151 millions on May 4th, 1914, 95 millions on April 29th,
1912, 68 millions on May 16th, 1911, 49 millions on April 22nd, 1910, and
44 millions on May 19th, 1911. The highest record we have for the autumn
species is 30 millions on September 26th, 1912. On May 16th, 1911,
C. debile contributed 30 millions and C. sociale 12 millions to the total in the
haul ; and on May 4th, 1914, C. debile gave 148 out of the 151 millions
Fic. 6.— Chetoceras decipiens, showing the active winter growth.
From a photo-micrograph by A. Scott.
present. If we examine the records of the separate species for the year 1914
as an example, we find that C. contortum has an average of 62,700 per haul in
May, C. debile an average of 867,878 in April and 18,972,800 in May (the
record), C. decipiens an average of 821,811 in April and 321,050 in May,
C. sociale an average of 1,229,500 in May, C. teres an average of 577,867 in
April ; while of the autumn species C. boreale has an average of 53,200 in
September and 54,644 in October, and C. densum has 151,120 in September
and 100,624 in October. C. debile and C. decipiens were also very abundant
that October.
The highest monthly averages for Chetoceras fall as follows during our
ten years :—
March <.< in 1907.
April ... "dn 1909, 1910; and 1912;
May ... in 1908, 1911, 1913, 1914, 1915, and 1916.
184 PROF. W. A. HERDMAN ON THE DISTRIBUTION
The highest average is nearly 23 millions, in May 1911. None of the
September and October averages run as high as those in spring, and only two
reach millions, viz., 3,956,047 in October 1911, and 7,702,658 in September
1912. The years 1911 and 1912 had high numbers of Chetoceras throughout
many of the months*. There are no months in the ten years when Chetoceras
was totally unrepresented ; but July and August show the lowest averages—
the lowest of all being only six individual cells in August 1907.
LAUDERIA.
We have only the one species, Lauderia borealis, Gran (fig. 7), in our
records. It is a late spring or early summer form, occurring generally from
March or April to June or July, with a later, smaller, occurrence in autumn.
It is sometimes present in large quantities, e. g., 20,064,000 on April 22nd,
1910; 12} millions on April 29th, 1912; 3,600,000 on May 4th, 1914.
The maximum is towards the end of April or beginning of May, when
Lauderia helps, along with Chetoceras, to form the main crest on the vernal
Diatom curve (see fig. 5).
Fic. 7.—Photo-micrograph showing a chain of Lauderia borealis.
THALASSIOSIRA.
The only species of this genus that are of any importance in our records
are T. gravida, Cleve, and T. Nordenskioldi, Cleve (fig. 8). Apparently
T. gravida is the only one common at Plymouth, but 7. Nordenskioldi, along
with Chetoceras contortum and C. debile, helped to constitute the vernal
maximum at Port Erin in 1907, and has been still more abundant on several
occasions since. T. Nordenskioldi is in the main a neritic, arctic or
Scandinavian species, and probably its occasional occurrences in quantity
are to be regarded as invasions of some arctic water and northern plankton
into our British seas. In April 1917, it was abundant at Port Erin along
with Chetoceras teres, C. debile, aud C. decipiens.
All our high records (over a million per haul) for Thalassiosira lie between
late in April and late in May, and the two highest are six millions on April
29th, 1912, and six and a half millions on May 16th, 1913. Other high
* The largest hauls of Diatoms as a whole, all species taken together, were in May of
1912 and 1913 (see Table on p. 188).
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 185
records have occurred during these same months, in 1910, 1911, 1912, 1913,
1914, 1915, and 1916.
The highest monthly averages for Thalassiosira are 721,000 in April
1912, 898,000 in May 1913, and 976,000 in May 1915. The genus is
totally absent from our records throughout the ten years in December,
Fre. 8,— Photo-micrograph of a phytoplankton consisting mainly
of Thalassiosira Nordenskioldi.
January, and February, and again in July and August, and is very poorly
represented in several other months. In fact, it is only really abundant in
April and May and not always then. In this distribution over the months
of the deeade this genus forms a marked contrast with Chetoceras, which is
so constantly present.
RHIZOSOLENTA.
The species that occur most abundantly in our records are Rhizosolenia
semispina, R. Shrubsolei, Ft. Stolterfothi and, less abundantly, R. setigera and
R.alata (rarely). The two last named are temperate Atlantic forms, and
R. semispina is a typically arctic oceanic species.
They are all summer or autumn species, the genus being often quite
unrepresented in certain of the winter months. It usually begins to appear
in the nets in February and reaches its maximum in June. The numbers
are sometimes enormous, giving the water in a glass jar a characteristic silky
or fibrous appearance and causing the June crest that is usually present as a
distinct elevation on the Diatom curve (see fig. 5, p. 182).
The most abundant species in the Irish Sea is Phizosolenia Shrubsolei—at
Plymouth it is 7. Stolterfothi ; but R. semispina (fig. 9) is sometimes present
in abundance at Port Erin in autumn, causing a second crest or maximum in
September or October. In September 1907, thirteen millions and sixteen
millions of this species were taken in two hauls of the surface-net at Station
III. (three miles off Port Erin), while the following year, at the same time
and locality, it was almost absent,
|
186 PROF. W. A. HERDMAN ON THE DISTRIBUTION
We have had much larger numbers, of R. Shrubsolei, in June; and amongst
our largest records of the genus taken as a whole are :—184 millions on
May 30th, 173 millions on June 3rd, 59 millions on June 8th, 43 millions on
June 13th, all in 1912, and 17 millions on June 21st, 1915.
Fra. 9.—Photo-mierograph of a phytoplankton consisting mainly
of Rhizosolenia semispina.
The highest monthly average is in June in all of our years except 1907 when
it was in May, and 1913 and 1916 when it was in July. The greatest average
recorded is over 40 millions in June 1912.
The autumn records are very irregular and sometimes show no rise
whatever. Even when present it is slight compared with the June maximum
(see fig. 5, p. 182). The highest monthly average in autumn is 117,122, in
September 1912.
GUINARDIA.
Only the one species, Guinardia flaccida, occurs in our records
(tig. 10). It is a summer form occurring mainly between April and July,
with the maximum almost invariably in June. Our largest records are
22,800,000 on June 3rd, 1912; 18,000,000 on May 30th, 1912; and
8,773,000 on June 11th, 1910. We have several records of over 7,000,000
early in June, one of over two millions on July 8th, 1913, and one of over
a million as late as July 16th, 1916.
Guinardia, when present, makes up along with Rhizosolenia the June
elevation, which is sometimes so marked towards the end of the vernal
Diatom maximum. This last summer (1917) it was unusually late, showing
a maximum of five millions on 23rd July, along with one million of
Rhizosolenia.
The highest monthly averages are all in June with the exception of 1916,
when it was in July. The greatest recorded average is close on six millions
for June 1912, The lesser, secondary, increase in autumn has its highest
ipt re. pm
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 187
point in September—the greatest recorded monthly average being 10,967 for
September 1912. It is interesting to notice that according to Pavillard
Guinardia flaccida is abundant in the Gulf of Lyons as late as October.
Fro, 10.—Guinardia flaccida, from a photo-mierograph by A. Scott.
TABLE
showing the monthly averages of the selected Diatoms throughout a year,
| |
1912. | Biddulphia. Mation — Rhizosolenia, ees Guinardia. | Lauderia,
|Jan..... 24,920 7,842! 9,877 | 22 0 957 189.
Feb. .. 306,885 10,201 10,034 12 0 105 0|
Mar. .. 21,176 8,977,292 95446 595 95 162 387 |
Apr ..| 21,869 18,365,750 100,019 ^ — 36,464 721,338 3,158 1,099478 -
May ..| 1,072 | 2,844,861| 14,586 29,008,744 41,083 | 2,789,438 | 170,878
June .. 0 67,543 0 | 40,888,771 0 | 5,919,429 0
Taly. 0 39,527 56 1,228 0 158 0
Aug. .. 1 694,961 0 13 0 0 0
Sept. ..| 4,181 | 7,702058. 1,077 | 117,122 | 3,978 10,907 | 89944
De....| 6897 214,421) 11,914 820 57 2176 99
Nov. ..| 95714 | 9476 5,486 0 0 21 0
Dec. ..| 8,059 1,106! 1,578 0 0 141 2
The above table shows very clearly how these important genera reach
their maxima at different times, and how, for example, Chetoceras in spring
(April) gives place to Rhizosolenia and Guinardia in early summer
(May and June). Curves can be drawn mentally from a glance at
the columns of figures which will demonstrate the waxing and waning of
the several types.
vcr ee eee:
188 PROF. W. A. HERDMAN ON THE DISTRIBUTION
The folowing table showing the largest hauls of total Diatoms and
of total Copepoda in each of the ten years brings out well how greatly the
Diatoms outnumber the Copepoda—in some cases in the proportion of about
a thousand to one.
Diatoms. Copepoda.
a | EIUS
| Largest haul. | ^ Date. Largest haul. | Date.
1907 ....| 698,950 | April 6th 28,900 | Aug. 17th |
1908 ....| 5,746,300 May 28th 68,015 | Sept. 14th
1909/52: 10,358,300 April 22nd 71,010 | Oct. 18th
1910 .... 70,128,400 | April 22nd 119,650 | Sept. 19th
1911 ....| 69,982,500 | May 16th 948045 | July 18th
1919 ....| 202,993,600 | May 30th 223,789* | May 20th
1913 ....| 205,814,700 | May 16th 118600 | July 21st
1914 ....| 155,288,000 ^ May 4th 217,571 | Nov. 9th
1015.29 18,895,500 June 15th 117,940 | Aug. 2nd
1018 2 -| 24,260,800 May 25th 118,524 | Sept. 25th |
wW o 64,339,250 | May 3rd | 147,706 | July 19th |
The relatively high records for Diatoms in 1912 and 1913 are due in the
former case to the usual June maximum of Rhizosolenta and Guinardia, and
in the latter case to a quite exceptional invasion of Asterionella japonica—
the only occasion in our records when this genus has been abundant in the
Irish Sea. We regard it as quite a rare form here. It is exceedingly
abundant to the south of Iceland (Ove Paulsen) and also in the Gulf of Lyons
(Pavillard).
COPEPODA.
The following table shows the total numbers recorded of our six important
species of Copepoda in the years 1909 and 1910 :—
1910. 1909.
Oithona helgolandica — ... 872,678 465,066
Pseudocalanus elongatus 308,926 309,973
Acartia clausi os 340,631 63,373
Paracalanus parvus — ... 217,633 54,120
Temora longicornis P 147,043 62,059
Calanus finmarchicus ... 15,418 21,412
* An exceptionally large haul at such an early date—due apparently to several moderately
large swarms of different Copepoda occurring together, viz., Calanus, Pseudocalanus, Temora,
and Acartia,
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 189
The actual detailed numbers are of no importance except as indications
of the relative abundance of the species. It is clear that Oithona and
Pseudocalanus (in 1909) far outnumber the others. Acartia shows a notable
increase in 1910.
The Copepoda as a whole are a summer and autumn group, all the crests
of their annual curves being found between May and October. Figure 11
shows curves of occurrence of the five most abundant of the above-named
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species for the year 1912, which may be regarded as typical. Curves of
Calanus are shown separately in figure 12. The remaining one of our six
selected species, Calanus finmarchicus, although much the largest individually
and probably one of the most important from the fisheries point of view, only
occasionally occurs in very large quantities in the Irish Sea, and its total
numbers in a year are much below those of the other species, as the following
table shows.
LINN. JOURN.—BOTANY, VOL. XLIV.
R
PROF. W. A. HERDMAN ON THE DISTRIBUTION
190
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OF DIATOMS AND COPEPODA IN THE IRISH SEA. 191
CALANUS.
Calanus finmarchicus (Gunner) * is present throughout the year in the
Irish Sea, and may be taken in small quantities at almost any time and in
larger numbers on different occasions in different years. In 1907 it was
fairly abundant (a few thousands in a haul) in April, and again at the end of
July and the beginning of August, and in October. In 1908, on the other
hand, the thousands per haul were not reached until later in the year, and
the numbers remained relatively high from September to December (2,850
on December 23rd). In 1909, in addition to occasional thousands in May,
EA Calanus
Jew "Fas. ' Mer ' Apr * Moy dane eum "Rag "Bap C Oct “Nov ^ Dec.
Fia. 12.—Curves of Calanus for three years (1907-9) at Port Erin, showing
successive waves of invasion, 2 or 8 in each summer,
June, July, and October, we have evidence of an unusually large swarm that
entered the bay on July 17th and 19th, when hauls estimated at 20,000 each
were obtained. The average for these three years shows maxima in July
and October (see fig. 12).
In 1910 there were no very large hauls, and the thousands were only
reached at the end of July and the beginning of August—perhaps the most
usual time for swarms to appear in the bay.
In 1911 the numbers in general were low, but two of the customary large
swarms appeared in summer, one on July 4th and the other on July 18th.
In 1912 the numbers were higher again, and thousands occurred on
various dates from April to October. The very high record of 50,720 was
obtained on May 17th, unusually early for such a large swarm.
* [ use this older specific name as I am inclined to agree with Wolfenden, Esterly and
others that the characters used in the attempt to separate “ finmarchicus” and “helgolandicus”
as species are too slight and inconstant to be relied upon.
n2
192 PROF. W. A. HERDMAN ON THE DISTRIBUTION
In 1913 the numbers were lower, and the maximum was 4000 on
July 28th.
In 1914 the maximum was 7320 on August 6th.
In 1915 again the numbers were low, the highest being 3232 on July 1st.
In 1916 a swarm appeared in the bay on July 11th, when over 12,000
were taken in a haul. A few days before a similar haul gave only 10
specimens, and a few days after 200. The average per haul at Port Erin
over the ten years is only 266.
On the whole this record agrees well with that at Plymouth, which is
* Common on and off from the end of April to the beginning of November,
generally present in small numbers at other times” (Miss Lebour).
Fic. 13.— Calanus finmarchicus, from a photo-micrograph by A. Scott.
Calanus finmarchicus (fig. 13), as its specific name suggests, is a northern
or Scandinavian form with a wide distribution through the colder waters of
the North Atlantic. According to Ove Paulsen its home and centre of dis-
tribution lies te the south of Iceland.
In Loch Fyne on the west coast of Scotland, off Skate Island, Calanus
appears to be present throughout the year in great quantities, in deep water,
at or near the bottom, along with Jucheta norvegica and Nyctiphanes
norvegica. For example, in July 1907, off Skate Island in 104 fathoms, we
eaught 13,000 Calanus in one vertieal haul, and on another occasion off
East Loch Tarbert, in 76 fathoms, we got 10,000. We have also obtained
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 193
from the yacht at various localities in the deep lochs on the west coast of
Scotland large hauls of Calanus by means of vertical hauls when no
specimens were obtained at the surface. In fact this Copepod seems to be
permanently present in the deeper waters of these lochs.
Tn addition to these deep water specimens of Calanus we find on occasions,
on the west coast of Scotland, large swarms on the surface, which may be
the result of invasions from the ocean, and there is a considerable amount of
evidence that such swarms are the cause of local mackerel fisheries.
Some years ago (1909) Dr. E. J. Allen and Mr. G. E. Bullen published *
some interesting work, from the Plymouth Marine Laboratory, demonstrating
the connection between mackerel and Calanus and sunshine in the English
Channel; and Farranf states that in the spring fishery: on the West of
Ireland the food of the mackerel is mainly composed of Calanus. In the
summer of 1913 we had an experience at Tobermory, in Mull, which I shall
quote from two letters written from the yacht at the time, and published in
* Nature."
* S.Y. ‘Runa,’ Tobermory, July 12th, 1913.
“ On arriving in this bay last night we found that the local boats had been
catching abundance of mackerel close to. We bought some for supper (good
fish for a halfpenny each), and on dissection found that the stomachs of all
of them were crammed full of fresh-looking Calanus (the individual Copepods
being for the most part distinct and perfect), along with a few immature
Nyctiphanes and larval Decapods. Professor Newstead and my daughter
then noticed, while fishing over the side of the yacht, about 8 p.m., that the
gulls in the bay were feeding in groups around patches of agitated water
evidently caused by shoals of fish. On rowing out to these we saw distinctly
the mackerel, large and small, darting about in great numbers in the clear
water, and we also noticed every here and there on the smooth surface of the
water—it was a beautifully calm evening—innumerable small whirls or
circular marks which, looking closely, I found to be caused by large Copepoda
close to the surface.
“ About twenty years ago I senta note to ‘ Nature,’ from the yacht * Argo,’
in regard to large Copepoda (I think it was Anomalocera on that occasion, and
the locality was further north, off Skye), splashing on the surface so as to
give the appearance of fine rain; and this present occurrence at once
reminded me of the former occasion, but here the Copepod was Calanus
inmarchicus of large size and in extraordinary abundance. They could be
clearly seen with the eye on leaning over the side of the boat, a small glass
collecting jar dipped at random into the water brought out twenty to thirty
specimens at each dip, and a coarse grit-gauze tow-net of about 30 em. in
diameter caught about 20 cubic centimetres of the Copepoda in five minutes.
* Journ. Mar. Biol. Assoc. vol. viii. (1909) pp. 394-406.
T Conseil Internat. Bull. Trimestr. 1902-8, Planktonique, p. 89.
TURN
194 PROF. W. A. HERDMAN ON THE DISTRIBUTION
The mackerel were obviously darting about, occasionally leaping to the
surface (which gave the gulls their opportunity) where the whirls, caused by
the Copepoda, were thickest, and an examination of the stomach-contents of
the fish on the yacht afterwards, showed us that the amount in one mackerel
was about the same quantity as that caught by the tow-net in five minutes.
Professor Newstead and I have made a count of 8 c.c. of the tow-net
gathering, and estimate that it contains about 2400 specimens of Calanus.
This would give about 6000 Copepods in the stomach of an average mackerel,
or in a five-minutes’ haul of the tow-net, on this occasion.
Fie. 14.—Photograph of exceptionally large hauls (about 1000 c.c. in a
jar) of Calanus taken from the yacht ‘Runa’ in 1913 on the west coast
of Scotland. The largest haul was estimated to contain at least
half a million individuals,
* [t may be added that these mackerel were evidently not being nourished
y were y g
in accordance with the views of Piitter, and were clearly able to fill their
, y.
stomachs from the plankton around them.”.
The following note, written some weeks later, records the conclusion of the
g , E
matter, so far as that summer’s observations went :—
“S.Y. ‘Runa,’ off Island of Eigg, August 12th, 1913.
* On getting back to Tobermory on Saturday, we found the plankton to be
g 8g y JE l
in marked contrast to its condition four weeks ago. The vast swarm of
I2]
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 195
Calanids has gone, and there are now no signs of mackerel feeding in the
bay. In faet, the change has been noticeable for some days in the seas
outside, and we have not been getting lately the large plankton catches
that were usual in the latter half of July. On July 14th a haul of the
large surface tow-net, in the open sea off Ardnamurchan, gave such a huge
catch of Calanus (about 1000 c.c.) that we promptly took a second similar
haul, and had it cooked as a sort of potted ‘shrimp’ confection for tea
(sampled by ten persons, including the crew, who were much interested to
try this new edible *fish?) ; while on August 11th a haul of the same
net, taken at the same spot, gave only a small catch of some 15 c.c.,
containing very few Calanids, along with the usual scanty summer
zooplankton.” . . ..
The importance of Calanus as a food for migratory pelagic fishes such
as the mackerel cannot be doubted (see fig. 14).
PsSEUDOCALANUS.
Pseudocalanus elongatus (Boeck) is widely distributed in the North Atlantic
and is present in the Irish Sea throughout the year (fig. 15). It reaches its
lowest level in January and February, and has its maximum in late summer
and autumn (June to October in our records, generally September or October).
Fra. 15.— Pseudocalanus elongatus, from a photo-micrograph by A. Scott.
This Copepod comes next after Oithona as the second most abundant
species in the Irish Sea, its average per haul over the ten years being 4583.
Some of the most important records are :—65,200 on May 20th, 1912 ;
91,960 on October 21st, 1912 ; 68,120 on June 24th, 1913 ; 60,600 on May
5th, 1916 ; 59,600 on September 25th, 1916 ; 58,200 on October 31st, 1912;
54,350 on July 31st, 1911; and many other records between 35,000 and
50,000 in these same months.
196 PROF. W. A. HERDMAN ON THE DISTRIBUTION
PARACALANUS.
Paracalanus parvus (Claus) is a southern form (fig. 16) which extends to
the tropics and has a wide distribution. It is present in quantity only during
autumn and winter in the Irish Sea, being practically absent between
February and July, and having its maximum in September or October. The
largest hauls are in autumn, for example, 59,460 on September 7th ; 138,300
on September 10th; 53,500 on September 15th, all in 1914; 61,930 on
August 24th, 1911; 49,800 on September 8th, 1910; 49,390 on September
Tth, 1912 ; 44,060 on October 24th, 1912 ; while other large hauls in the
same months range from 25,000 to 40,000.
Fie. 16.— Paracalanus parvus, from a photo-micrograph by A. Scott.
This is a species that has varied considerably in quantity from year to
year, its average per haul ranging from 325 in 1907 to 5863 in 1914. In
the present year, 1917, it was especially abundant at Port Erin in September,
along with Oithona.
OITHONA.
Oithona helgolandica (=O. similis), Claus, is the most generally abundant
Copepod throughout the year in the Irish Sea ; but the months when it is
taken in greatest numbers are June to November (inclusive) with the
maximum generally in July. There may, however, be a second maximum
later, in October or November.
The following may be quoted as exceptionally large hauls :—The highest,
225,450, is on July 18th, 1911, and the next highest, 199,300, is on
November 9th, 1914 ; then we have 93,580 on July 28th, 1913, 87,530 on
September 26th, 1912, and 83,550 on June 14th, 1910. We have twenty
other records of over 44,000 each scattered over the months June to
November. This is the Copepod which has the highest average number per
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 197
haul over the ten years, viz., 8171 (see table, p. 190). Although this is a
small species (fig. 17), still from its very great abundance it must be of
considerable economie importance as a food-matter in the sea.
Fra. 17.— Oithona helgolandica, from a photo-micrograph by A. Scott,
ACARTIA.
Acartia clausi, Giesbr., is a summer species widely distributed over the
North Atlantic, and present in greatest abundance in the Irish Sea from May
to October (inclusive), and having a maximum generally in June, with
another slighter rise in autumn, September or October.
Fre. 18.—Acartia clausi, from a photo-micrograph by A. Scott.
The following are some of our highest records :—59,490 on June 27th,
1912 ; 59,360 on August 24th, 1911 ; 52,200 on May 20th, 1912 : 50,100
on June 2nd, 1913 ; 44,000 on September 29th, 1913 ; and 41,950 on June
17th, 1919. The average of this species per haul throughout the ten years
198 PROF. W. A. HERDMAN ON THE DISTRIBUTION
is 2571. Swarms of this oceanic species (fig. 18) in summer may possibly be
connected with an inflow of Atlantie water ; bnt probably also large numbers
appearing locally in enclosed areas may be the result of reproduction and
development in situ.
In some cases Acartia has been found in unusual abundance quite close to
the shore, and even in water of low salinity.
Towards the end of September, 1917, we emptied for cleaning purposes
the large open-air fish-ponds at the Port Erin Biological Station, and when
most of the water had been pumped out we noticed that what remained was
swarming with small Copepoda. Every bucket and jarful that one removed
for inspection was found to be densely crowded—as densely, in fact, as an
average jar of plankton representing the contents of a standard haul. Some
buckets of these Copepoda were used for feeding the lobster larvæ and
some of the smaller fish and other animals in the Aquarium, and a small
sample that was preserved was found to bea practically pure gathering of
Acartia clausi. One specimen of Centropages was present in the tube amongst
many thousands of Acartia, when Mr. Scott took the photo-micrograph shown
in fig. 18.
TEMORA.
Temora longicornis (Müll.) is a very local species, which may be found on
occasions swarming in great abundance in small areas of the sea, generally
close to land. It is a summer species and its maximum is generally in June
Fia. 19.— Temora longicornis from the stomach of a Mackerel.
From a photo-micrograph by A. Scott.
or July, but was in May in 1912, when the large hauls of 50,400 and 83,400
were taken on the 17th and 20th respectively. Other large hauls of this
TOC PUNCTUM á
a e a e a aiin EI
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 199
species 1n our records are :—45,530 on July 11th, 1916 ; 43,800 on June 11th,
1913 ; and 34,665 on July 26th, 1910. The average for this species per
haul throughout the ten years is 1234.
Temora (fig. 19) is characteristic of coastal as opposed to Atlantic water,
and is the only “ neritic” form in our series. The other five are all usually
classed as '*oceanic," but as I have pointed out above most of them are
really present throughout the year in the Irish Sea.
Temora longicornis is on occasions one of the most abundant of our
'opepoda in the Irish Sea and must be of considerable importance as a food
for fish and especially for the herring in summer. In the latter part of July
and the first half of August, 1917, the shoals of herring to the west of the
Isle of Man came unusually close to land and even penetrated into bays and
creeks ; and during this time they were feeding mainly, if not wholly, on
Fic. 20.— Temora longicornis from the “ red patches.” From a
photo-micrograph by A. Scott.
Temora. Late in July this Copepod was so abundant that its presence
caused large patches of a red colour on the surface of the sea off Port Erin aud
around the Calf Island. These red patches were noticed by the fishermen,
and were spoken of amongst them as being “ fish-food” or “spawn.” A
large jarful from such a red patch, obtained by one of the fishermen, was
brought to the laboratory and found to be swarming with small Copepoda,
which on examination proved to be almost wholly Temora longicornis (fig. 20).
About one-fourth part of the contents of the jar was preserved, and on being
counted later on was found by Mr. Andrew Scott to amount to 50 c.c.
of Copepoda, consisting of 33,340 Temora and 2 Calanus. Mr. Scott
estimated the oil present in 9 c.c. of the dried Temora at 2°47 per cent. of
the weight, which was 0:925 gramme.
During iliis same time the men were catching herring in quantity unusually
close inshove in the neighbourhood of the red patches, and on examining, in
200 PROF. W. A. HERDMAN ON THE DISTRIBUTION
the laboratory at the Diological Station, the stomach-contents of a number of
these herrings, I found in every case that the stomach contained a mass of
red material which was obviously, under the microscope, the broken-down
remains of Copepoda. A few Crab zoea were recognisable, but the bulk of
the material consisted undoubtedly of the Copepoda. Mr. Scott examined
5 c.c. of the stomach-contents for me, and found that it contained 975 easily
recognisable specimens of Temora. A photograph (fig. 21), which Mr. Scott
Fra. 21.— Temora remains from the stomachs of the Herring.
From a photo-micrograph by A. Scott.
has made from one of the microscopic preparations, shows appendages that
undoubtedly belong to this Copepod, while here and there in the stomachs
complete specimens of Temora are to be seen. It is not possible to doubt
that during these weeks, at the height of the summer herring fishery in the
Irish Sea, the fish were feeding mainly upon this species of Copepod.
We recorded a similar occurrence off the Lancashire coast a few years ago,
when in July 1913, at the time of an abundant mackerel fishery off Walney
Island, the stomachs of some of the fish were found to be full either of Temora
alone or of Temora mixed with Jsias and a few other Copepoda (see fig. 19,
p. 198). A few herrings from the Port Erin fishery of July 1916 were
found by Mr. Scott to be feeding mainly on Calanus.
CONCLUDING REMARKS.
Many food-fishes are known to feed upon Copepod plankton during at
least some portion of their life. The Loch -Fyne herrings are frequently
at the time of a fishery found to have their stomachs filled with HKucheta or
Calanus. Mackerel, in the English Channel and to the S. W. of Ireland and
elsewhere, have been recorded as feeding on Calanus. It has been shown in
this paper that in Hebridean Seas the mackerel and in the Irish Sea herrings,
TY 2a, eee eo
OF DIATOMS AND COPEPODA IN THE IRISH SEA. 201
at a time when they are present in great abundance, are feeding on some
prevalent form of Copepod, such as Calanus or Temora. Other similar cases
could no doubt be quoted and are known to marine biologists.
Then, as to demersal fish—young plaice, after their metamorphosis, feed
chiefly on the smaller Copepoda, while in younger stages the post-larval
plaice feeds upon Diatoms. We have noticed at the Port Erin Biological
Station the post-larval plaice with its stomach showing of a golden brown
colour from the Diatoms with which it was filled, and we have watched in a
shallow pond the metamorphosed young plaice darting backwards and for-
wards pursuing, catching, and devouring the individual Copepoda. It is
known that these Copepoda in their turn feed in part at any rate on Diatoms,
so our two main constituents of the plankton are undoubtedly concerned
in the nourishment of either young or adult fishes useful to man.
The association of shoals of fish with abundance of plankton is the
result of the fact that, in order to get an adequate quantity of planktonic
food, the fish must seek out and capture the Copepoda. In other words, the
fish must go where the plankton is abundant and must in its movements
follow the movements of the shoals of Copepoda. It is the very poverty
of the plankton in some sea-areas, insisted on by Piitter, Lohmann and
others, which makes it necessary for plankton-eating fish to move about
in search of more abundant supplies.
Consequently it is of importance to show, as we now can, that in our
coastal seas at least, where the fisheries we are interested in take place, the
plankton is not uniformly distributed. Many of the Copepoda occur very
definitely in local swarms, and various localities and depths are characterised
at the different seasons by particular assemblages of plankton. It is
therefore reasonable to believe, in view of the facts given above as to the
association of fish and plankton, that these variations in the distribution
must have a marked effect upon the presence and abundance of at least
such migratory fish as herring and mackerel, and also of the shoals of post-
larval young of many of our other food-tishes.
No less than three masses of sea-water of different origin and character
may enter or affect the British seas in varying quantity, viz.:—(1) Arctic
water such as normally surrounds Iceland and the east of Greenland, and
may extend further southwards and eastwards towards Norway, the Faroes,
and Shetlands; (2) Atlantic (“ Gulf-stream Drift”) water which impinges
on the western shores of Ireland, and may flood the English Channel and
extend round the Shetlands or down into the North Sea ; and (3) “ Coastal”
water such as flows out from the Baltic and, mixed with the other waters,
bathes the coasts of N.W. Europe generally, and to a large extent surrounds
the British Islands.
The Irish Sea may be regarded as primarily an area of coastal water,
which is, however, liable to be periodically invaded to a greater or less extent
202 PROF. W. A. HERDMAN ON THE DISTRIBUTION
by bodies of warmer and salter Atlantic water (re-inforced possibly by por-
tions of a deeper outflowing Mediterranean current) carrying in oceanic
plankton, and more rarely perhaps by Norwegian or Arctic water causing
an invasion of northern organisms. The variations which we find in different
years in the nature and amount of the plankton at the same localities no
doubt depend to some extent upon the volume and period of such southern or
northern invasions ; but they may depend also upon other faetors, such as
the weather (temperature, sunshine, rainfall, wind, ete.) at the time, and
previously.
Of the six Copepoda discussed above only one— Temora—is a neritie form ;
the others are all usually regarded as oceanie, that is as having their true
home and centre of distribution somewhere to the north, west, or seuth in
the open Atlantic.
The following list gives an approximate indication of what is supposed
to be the source of these five oceanie Copepoda :—
Calarus.—N. Atlantic, about Iceland (** Boreal oceanic ”).
Paracalanus.— Southern, temperate and tropical Atlantic.
Pseudocalanus.—N. Atlantic (* Boreal oceanic”).
Oithona.—N. Atlantic (“ Boreal oceanic ”).
Acartia.—N. Atlantic (* Northern styli-plankton ").
Some no doubt live on during the year in the Irish Sea, but these indi-
genous populations are probably reinforced by waves of immigration from
outside.
In the case of our Diatoms some of the species of a genus may be neritic
and others oceanic, as is shown in the following list, where (N.) stands for
neritic and (O.) for oceanic, and a (?) indicates that the evidence seems to
me conflicting or inconclusive *:—
Biddulphia mobiliensis (N.), B. sinensis (O.).
Coscinodiscus radiatus (O. ?), C. concinnus (N.), C. Grani (N.).
Lauderia borealis (N.).
Chætoceras boreale (O.), C. eriophilum (O.), C. decipiens (O ?), C. densum
(0.), C. eontortum (N.), C. debile (N.), C. diadema (N.), C. sociale (N.),
C. subtile (N.), C. teres (N.).
1 halassiosira gravida (N.), T. Nordenskioldi (N.).
Rhizosolenia alata (O.), R. semispina (O.), R. setigera (N.), It. Shrubsolei
(N. ?). R. Stolterfothi (N. ?).
Guinardia flaccida (N. ?).
It is remarkable how small a number of forms make up the bulk of the
macro-plankton throughout the year. These half-dozen kinds of Diatoms
and half-dozen Copepoda are the all-important organisms upon which our
* The matter was discussed more fully some years ago in a paper by Herdman and
Riddell in Trans. Biol. Soc. Liverpool, xxv. (1911) p.. 178,
iE! DIATOMS AND COPEPODA IN THE IRISH SEA. 203
fate depends so far as concerns food from the sea. That shoals of migratory
fish such as herrings and mackerel are attracted in summer and autumn by
dense swarms of certain Cop^poda such as Calanus and Temora can no
longer be doubted ; and there is abundance of evidence that earlier in the
year the young staves of other food-fishes, such as plaice, are nourished first
by Diatoms and when larger by the smaller Copepoda. These conclusions,
however, do not finish the matter. There remains in the sea the much more
minute Protozoa and Protophyta of the Nannoplankton, which to a large
extent escape. through the meshes of our plankton nets and which are doubt-
less of great importance as the food of Copepoda and other large organisms
which in their turn nourish fish.
There are several other interesting lines of further investigation which
open up as collaterals from plankton investigation. One of these is the
origin of the great vernal phytoplankton maximum. In the spring there
is an awakening of the plant-life of the sea comparable to the growing of the
grass and the budding of the trees on land. This cannot be due to any rise
in temperature, as the sea at the time that the Diatoms start their active
growth and reproduction is at its coldest. We have series of observations
extending over more than ten years showing that the water of Port Erin bay
is generally of lower temperature in March than in December or January.
Of the various causes for the rise in the Diatom curve in these cold waters
of early spring I have for some years * regarded with greatest favour the
view suggested first by Sir John Murray, viz., the increase in sunlight at
that time of year. In that ease it would be a photosynthetie phenomenon—
the increased solar light energy enabling the Diatoms to obtain from their
environment by photosynthesis the materials required for their growth and
reproduction. The view that the spring increase in plankton is due to changes
in the alkalinity of the water does not in my opinion conflict with the photo-
synthesis theory but is supplementary to it. The position in regard to the
relation between variations in alkalinity and in the plankton, in our district,
is as follows :—
The sea around the Isle of Man is a good deal more alkaline in spring
(say April) than it is in summer (say July); and during the years 1912-14
Professor Benjamin Moore, by examining samples of sea-water periodically
at the Port Erin Biological Station, was able to show t that the alkalinity,
which gets low in summer, increases somewhat in autumn, and then decreases
rapidly, to disappear during the winter ; and then once more, after several
months of a minimum, begins to come into evidence again in March and
rapidly rises to its maximum in April or May. This periodic change in
alkalinity will be seen to correspond roughly with the changes in the living
* See ^ Intensive Study, &c." Part IIT. p. 260, 1910, Trans, Biol, Soc. Liverpool, xxiv,
t Trans. Biol. Soc, Liverpool, xxix. (1915) p. 233,
204 DISTRIBUTION OF DIATOMS AND COPEPODA IN THE IRISH SEA.
microscopic contents of the sea represented by the phytoplankton annual
curve, and the connection between the two will be seen when we realise that
the alkalinity of the sea is due to the relative absence of carbon dioxide. In
early spring, then, the developing myriads of Diatoms in their metabolic
processes gradually use up the store of CO, accumulated during the winter
and so increase the alkalinity of the water, till the maximum of alkalinity,
due to the reduction in amount of carbon dioxide, corresponds with the crest
of the phytoplankton curve in, say, April. Prof. Moore has calculated that
the annual turn-over in the form of carbon which is used up or converted
from the inorganic into an organic form probably amounts to something of
the order of 20,000 or 30,000 tons of carbon per cubic mile of sea-water in
the Irish Sea; and this probably means a production each season of about
two tons of dry organic matter, corresponding to at least ten tons of moist
vegetation, per acre—which shows that we are still very far from getting
from our seas anything like the amount of possible food-matters that are
produced annually.
Testing the alkalinity of the sea-water may therefore be said to be merely
ascertaining and measuring the results of the photosynthetic activity of the
great phytoplankton rise in spring due to the daily increase of sunlight.
Other possible causes, more or less related to the above, have been suggested
—such as Brandts hypothesis that the fluctuations in the phytoplankton
depend upon the accumulation, and then the exhaustion, of necessary
inorganic food-matters in the water, such as nitrogen or phosphorus com-
pounds or silica ; and the view of Nathansohn, Gran and others that vertical
currents, carrying up food-matters from the deeper water, have a powerful
effect upon the seasonal development of surface plankton. These may be
contributory causes or may be effective locally, or on occasions; but it seems
probable that a widespread phenomenon of enormous amount such as the
vernal increase of phytoplankton must depend upon an equally widespread
and powerfully-acting cause such as the rapid increase in the amount of
solar light energy which marks the lengthening days of the year in early
spring.
| OP ea
DR. D. H. SCOTT: NOTES ON CALAMOPITYS. 205
Notes on Calamopitys, Unger. By D. H. Scorr, F.R.S., F.L.S.
(PrATES 6-8 and one Text-figure.)
(Read 15th November, 1917.]
CONTENTS.
Page
INTRODUOTION NA E be vss 'sls edes nnn eral dE DERI 205
I. Tux Lrar-rRACE OF Calamopitys americana... eee eee 206
II THE Genus (Clemons. 3 rero ire LA LE OERTOAN 209
Calamopitys annularis ............ E E Lo occa N 210
Calamopitus OR o's. hoes nnn eraran EID 215
Calamopit/s amentoata, o.. ooo eee so QURE oe 218
Calamopitus fasciatlaris o. 5. occ o OE 218
Calamoptiys vBeweortiana, .. «s.aicicirioiszigesssisicnn a se CEE o 219
SYNOPSIG si...» re ue EID PICT Gore's. oc s/o» DEDE 221
Affinities -of the Specionlle 00... S ... 222
Affinities of the Genus Lese a a 226
du e A o as. ss o's oes | OS 00 0 15 S ou 229
LITERATURE OITED ....... QE ets. : soc oe CUERO 230
EXPLANATION OF THE PLATESEE reo os eee e 231
INTRODUCTION.
Five species of Calamopitys, of Lower Carboniferous or possibly in some
cases of Upper Devonian age, have been described : two, C. Saturni, Unger
and C. annularis (Unger), from Thuringia (Unger, 1856 ; Solms, 1896) ;
one, C. Beinertiana (Goepp.) from Silesia and from Scotland (Solms, 1893 ;
Scott, 1902) ; one, C. fascicularis, Scott, from Scotland and England (Scott,
1902) ; and one, C. americana, Scott & Jeffrey, from the State of Kentucky
(Scott & Jeffrey, 1914).
The two species C. Beinertiana and C. fascicularis, which I was responsible
for referring to Calamopitys, have been placed by Dr. Zalessky in a separate
genus, Fristophyton (Zalessky, 1911). The object of the present Notes is
firstly to put on record some fresh evidence as to the course of the leaf-trace
in C. americana, and secondly to discuss the relations of the various species,
with special reference to Dr. Zalessky's proposed division of the genus.
It will be remembered that Calamopitys is a genus of Cycadofilices *, with
a stem-structure comparable to that of Lyginopteris, for the pith is sur-
rounded bya ring of mesarch primary strands of xylem. Their structure,
however, is usually more centrally mesarch than in Lyginopteris. The
leaf-trace is a single strand at its origin, but divides further out, in all cases
* Used here in preference to “Pteridosperms” because there is no evidence whatever as
to the fructification.
LINN. JOURN.—BOTANY, VOL. XLIV. S
T MONEDAS X x UTOR
WOO WS
206 DR. D. H. SCOTT :
where its course has been followed. Secondary wood is present, often
reaching a considerable thickness. The secondary and metaxylem tracheides
have multiseriate bordered pits.
In three species, C. Saturni, C. annularis, and C. americana, the leaf-bases
are present and have the structure of Aalymma, containing a number of
bundles derived from the subdivision of a single leaf-trace.
In C. americana, and probably in C. annularis, the pith is a * mixed” one,
containing tracheides, and the structure therefore protostelie (Scott &
Jeffrey, 1914, p. 318). The differences between the species will be discussed
in the second part of the paper.
I. Tug Lear-rrace OF Calamopitys americana.
Tn the paper, by Prof. Jeffrey and myself, on Fossil Plants from Kentucky,
we showed that in C. americana the leaf-trace, starting as a single bundle,
divides into two as it passes through the zone of secondary wood, thus
entering the cortex as two distinct strands (Scott & Jeffrey, 1914, p. 321).
Certain difficulties, however, remained. In one case the original bundle was
found to divide twice, and there was some doubt as to the nature of the first
division, though it seemed probable that it served to separate the trace as a
whole from a reparatory strand which remained in the stele, while the second
division represented the duplication of the trace itself (l.e. p. 323). Neither
had we, at that time, any section clearly showing the two bundles of the
trace where they pass out of the secondary wood.
I have since had a new series of sections cut for me by Mr. Hemingway,
which shows the whole process of division of the trace quite clearly, as well
as its relation to the reparatory strand. The new evidence only confirms our
previous conclusions, but it makes the case clear, where it was obscure before,
and therefore seems worth bringing forward.
The fragment from which the new series was cut was about an inch long,
and 20 transverse sections were obtained from it. It forms a portion of the
stem marked F, one of the larger specimens, the whole stem measuring about
40 mm., with a pith about 13 mm. in diameter. The uppermost section of
the series lies just below a section previously figured (l. e., Pl. 29. Phot. 22
and PI, 30. Phot. 28), so that the origin of the two bundles there shown can
be followed. In the original section only one bundle was complete, the other
fragmentary ; in the new series both are complete, from their first separation
onwards. A selection of the slides, illustrating the most important changes,
has been photographed (Pl. 6. Phots. 1-10). The series is followed from
below upwards.
In the lowest section (Phot. 1) the trace in question is still at the margin
of the “pith”; it has only just begun to move out wards, but its nature, as an
outgoing trace, is indicated by the fan-lihe arrangement of the rows of
Mu d s cda uM A c ecd
WEN S
NOTES ON CALAMOPITYS. 207
secondary tracheides radiating out from it, and starting from its flanks as
well as from the abaxial side. The protoxylem is disorganized but its
position is evident (Phot. 2); it lies deep in the interior of the xylem-
strand, nearer the outer than the inner edge. It will be noticed that the
elements lying on the inner side of the protoxylem are decidedly larger than
those towards the exterior. On the inner side the xylem-strand is not sharply
delimited, for the pith was *mixed," and also there was some connection with
a reparatory strand, better shown in a later section.
The next section shows little change. In section 3 (Phot. 3) the outward
progress is still very slight. The protoxylem of the outgoing strand appears
to have divided into two, so far as the imperfect preservation of this part
permits one to judge (Phot. 4). The most interesting point in this section,
however, is that the connection of the leaf-trace with a strand remaining in
the stele is clearly shown. This strand lies on the inner side of the xylem of
the trace and somewhat to one side (Phot. 4). Its protoxylem is evident
(though a wide crack passes through it). The two strands are clearly in the
act of separating. The structure is almost identical with that shown, from
another specimen, in Pl. 35. fig. 4 of the Kentucky paper (Scott & Jeffrey,
1914). The new evidence completely confirms the interpretation there
suggested, that the division in question “separated the trace as a whole from
a reparatory strand which did not immediately pass out” (p. 325).
Two sections higher up (Phot. 5) the connection between the reparatory
strand and the leaf-trace is almost severed. Though the xylem-strand of
the trace is damaged, the two distinct protoxylem-groups are now evident,
and there are signs of a division of the centripetal portion of the xylem.
In the following four sections the outward movement of the leaf-trace is
very slow, but the gradual division of the xylem into two becomes more
marked (see Phot. 6, from the 9th section). So far, however, it is only the
inner part of the strand that is affected, while the outer, centrifugal, portion
is still continuous. Even in the former, the separation appears wider than
it actually was, owing to decay of some of the elements.
In the succeeding sections the fan of secondary wood corresponding to the
trace broadens out, the lateral rows making a wider curve. At the same
time the primary xylem is completing its division. In Phot. 7, from
section 13, the two distinct strands are evident, though perhaps still slightly
connected on the abaxial side. The radial series of the secondary wood here
begin to show an arrangement corresponding to the doubling of the primary
strand, the original fan tending to resolve itself into two fans, as shown by
the slight opposite curvature of the adjacent median rows (Phot. 7).
The two strands of the leaf-trace now rapidly separate, and the outward
movement also becomes more marked. In the 15th section (Phot. 8) the
strands are separated by more than their own width and their protoxylem
centres are about 2:5 mm. apart. At this level a median band of secondary
82
208 DR. D. H. SCOTT :
wood abuts on the tissue separating the two strands and appears to belong
to neither : ultimately, however, this also is divided between them.
Two sections further up (section 17, Phot. 9) the two strands are com-
pletely organized as separate bundles, each with its own are of secondary
wood ; the rows of the wood spring from both flanks of either strand, and
are beginning to extend further inwards. The protoxylem-groups are here
over 3 mm. apart, the clear distance between the two primary strands being
nearly 2°5 mm. A wide ray is appearing between the two fans of secondary
wood.
In the next two sections rows of secondary wood appear between the two
bundles of the trace, and in the bundles themselves the secondary growth
begins to extend to the inner, adaxial side. Lastly, in the uppermost section
of the series (section 20, Phot. 10) there is a considerable tract of secondary
wood between the two strands extending far to the inside. It is also closing
in directly behind the right-hand (upper) bundle. In this strand the
secondary wood proper to the strand now clearly extends all round, the rows
on the adaxial side still being short. In the left-hand (lower) strand the
adaxial wood is less evident, but this may be merely a matter of preservation.
The right-hand bundle lies beyond the limits of the adjacent secondary wood,
while that on the left appears to be less far out.
In each bundle the primary xylem is radially elongated, an appearance
no doubt due in some degree to the oblique course of the outgoing trace.
The protoxylem, here as in the original single strand, lies somewhat nearer the
outer than the inner edge: it may be beginning to divide, but this is doubtful.
The section last, described comes next below that already figured in the
Kentucky paper (Scott & Jeffrey, 1914, Pl. 29. Phot. 22 ; Pl. 30. Phot. 28).
This section is reversed as compared with those of the new series; the
complete bundle shown in the former corresponds to the left-hand (lower)
strand in the photographs now given. The chief changes are that the
secondary wood of the stele has at the higher level completely closed in
behind this strand and the strand itself has well-developed wood all round it,
as deseribed in the Kentucky paper (p. 324). The compauion strand (the
right-hand (upper) one in the new series, left-hand in the 1914 figure) is
only represented by a fragment in the old section.
The series just described demonstrates, with perfect clearness, the division
of the leaf-trace into two equivalent strands as it passes out through the
secondary wood. The latter, however, is a late formation, and a better way
of expressing the facts is to say that the trace divided before reaching the
pericycle : i.e., while still within the zone of thickening. In this it differs
from the leaf-trace of C. Saturni, which, according to the detailed observa-
tions of Solms-Laubach, passed beyond the woody zone as a single bundle,
and only divided into two in the cortex (Solms, 1896, pp. 67, 69; Taf. 4.
figs. 1-3). The difference, however, is not very great. Solms-Laubach
7A EPEMERUEREUR T C TES
NOTES ON CALAMOPITYS. ` 209
states (p. 68) that the primary strand of the leaf-trace where it separates
from the wood has two protoxylem-groups, placed near the ends of the trans-
verse section ; his fig. 8 (Taf. 4) shows a leaf-trace still in the woody zone
of the stele and showing two most distinct protoxylem-groups; the
reparatory strand lies behind, and the whole appearance is remarkably like
some of our own sections (Phots. 4—6). The distinction is simply that in
C. americana the division of the trace is completed somewhat earlier in its
course than in C. Saturni. Inthe latter species, as in ours, the leaf-trace
had its own secondary wood, where it first became separate, but it is described
as limited to the outer side (p. 68).
It need hardly be pointed out that the evidence from the new series of
sections finally disposes of the suggestion that the outgoing bundles with
secondary growth might be the steles of branches. A single strand from
the peripheral ring passes out and divides, exactly in the manner of a leaf-
trace and in a way perfectly comparable to that already observed in the
undoubted leaf-traces of Calamopitys Saturni. '
II. Tug Genus Calamopitys.
. A comparative survey of the species which have been included in this
genus is required, now that our knowledge has been extended by the
discovery of a new form, Calamopitys americana, while a division of the
genus has been proposed by Dr. Zalessky, on grounds which demand
consideration.
The genus Calamopitys was established by Unger in 1856 on the species
C. Saturni from Saalfeld in Thuringia. Our knowledge of the true structure
of the type species is entirely due to Count Sol ime- Lanbi (1896), who added
a second species, C. annularis, which Unger had described as a Stigmaria.
These two species have hitherto been regarded as very closely allied, perhaps
not even truly distinct. I find, however, considerable differences between
them ; C. annularis very closely approaches the Kentucky species C. ameri-
cana, while C. Saturni is in some respects more like C. fascicularis, one of
the two species separated by Zalessky under the name Zristophyton.
C. annularis appears never to have been figured since the time of Unger,
who quite misunderstood its structure. I happen to have in my hands
several sections of this species, as well as two of C. Saturni, all lent to me
by the late Count Solms-Laubach some years ago, for comparison with
C. americana. I have taken advantage of this fortunate circumstance to
re-investigate the structure, so far as my limited material allows—it is not
likely to be added to in the near future. For C. annularis the available
sections are fairly adequate, and I have thought it well to give some figures
of this imperfectly known species as well as one or two EE EET illustrations
of the more familiar C. Saturni.
NU
210 DR. D. H. SCOTT :
I propose to begin with C. annularis, which stands most in need of further
elucidation, though I may say at once that my observations in almost every
respect confirm the brief account. given by Solms-Laubach, so far as it goes.
CALAMOPITYS ANNULARIS (Unger), Solms.
The available sections of this species are from three specimens (besides a
doubtful one), two of which (Berlin, 97 & 98) are described by Solms-
Laubach (1896, p. 75) as among the most indubitable examples *.
Specimen 97 shows the general structure best. The two transverse
sections are cut quite close together, and show no important change of
structure. Section 97 appears to come just above 40a.
The diameter of the stele, measured to the outside of the secondary
wood, is about 20 mm.; that of the pith, including the primary wood,
6-7 mm. The phloem is badly preserved, but a large piece of the cortex, :
or leat-base, quite 1 em. thick, is attached. This shows the Kalymma
structure, already described by Solms-Laubach (P1. 7, Phot. 11).
The primary xylem appears to be continuous or nearly so. The preserva-
tion is far from perfect, but the tracheides can generally be recognized by
their thiekened and. pitted walls; the pith is almost every where surrounded
by a band, several layers thick, of such elements; there may have been
occasional small gaps filled by parenchyma. The primary xylem-strands
are not very well defined ; judging from the frequency of protoxylem-
like groups of small elements, they must have been very numerous—
perhaps as many as 20 (see Phots. 12-14). As a rule, the strands are
eccentrically mesarch, the protoxylem lying decidedly nearer the outer
than the inner edge; in these cases there is usually a marked difference
between the centripetal and centrifugal portion of the xylem, the former
consisting of larger tracheides than the latter (see Phot. 13). In both
* The sections examined are as follows :—
1, Specimen, Berlin, 97,
Two almost identical transverse sections, labelled: “97 Stigmaria annularis. Saalfeld.
Richter. (Berl. Ldes. Anst. 97-1)." “40a. Stigm. annul. Culm Saalfeld. Coll. Solms 413."
One radial section, labelled: “97R. Calamop. ann. Berlin Landesanst, Coll. Solms. n. 413.”
2. Specimen, Berlin, 98.
One transverse section only, labelled : “98 Stigmaria annularis. Saalfeld Richter, Berlin
Landesanst. Coll. Solms, n. 411.”
3. Specimen, Halle, 761.
One transverse section, labelled; ** 761. Cal. annul, 9. Mus. Hal. Coll. Solms. n. 420.”
4. Specimen, Berlin, 49.
One radial section labelled: “49R. Stigm. ann. Culm Saalfeld. Berl. Ldesanst. Coll.
Solus. n. 421."
I am not sure that this isolated section is rightly named, and have taken no account of it
in the text.
NOTES ON CALAMOPITYS. 211
respects the xylem-strands of C. annularis agree with those of C. americana
(see Phots. 2 & 4; also Scott & Jeffrey, 1914, Pl. 35. figs. 1, 2, 4; Pl. 36.
fig. 9). In other strands the structure is less eccentric, and there is
correspondingly less differentiation between the inner and outer portions
of the xylem (Phot. 14). The xylem-strands which are in closest contact
with the secondary wood have the most eccentric structure; conversely,
those which are more nearly centrally mesarch stand a little further back,
and even, in some cases, appear to be separated from the secondary wood by
a layer or two of parenchyma.
It was mentioned in the paper by Prof. Jeffrey and myself (l. c. p. 326)
that there is evidence for the occurrence of medullary tracheides in
C. annularis, as in C. americana, I have carefully looked into this question.
In transverse sections one can distinguish at a few places elements with.
thicker walls than the ordinary parenchyma, and resembling the tracheides
of the xylem-ring. But the preservation is not such as to make these
indications at all convincing. The longitudinal section of specimen 97
passes through the middle of the pith as shown by the dimensions. Several
tubes are seen embedded in the medullary parenchyma; they are of very
various diameters, ranging from about 100 to about 360 u —the larger are
probably distorted (Phot. 15). The smaller tubes agree very well in shape
and dimensions with those in the pith of C. americana (Scott & Jeffrey,
1914, Pl. 34. Phot. 51; Pl. 36. fig. 8). These tubes appear to have a wall
of their own (Phot. 16), but pits have not been observed. There is a strong
presumption that these elements are medullary tracheides, but better evidence
is required. — Similar tubes occur in the pith of the only other specimen
under the name C. annularis of which I have a radial section. This will be
described below.
As regards the secondary wood there is not much to be said. The
medullary rays are multiseriate, but not dilated outwards (Phot. 12). In
radial section they are muriform, with narrow cells. The secondary
tracheides are much smaller than the large elements of the primary xylem—
about 60 p in the former to 100 w in the latter. The radial section shows
the pits in 3-6 series; the arrangement in horizontal or slightly oblique
ranks is fairly regular ; longitudinally, the pits may either be alternate or
in irregular vertical series. Their form is commonly somewhat elliptical
or at least elongated in the transverse direction. In all these respects
the structure agrees with that of C. Saturni and C. americana,
The sections at my disposal give little information as to the mode of exit
of the leaf-traces. In specimen 97 there is at one point a disturbance in the
regularity of the wood, no doubt connected with a leaf-trace (see Phots. 11
and 12). Two segments of the secondary wood are somewhat prominent,
with a convex outer edge; between them the tissue is badly preserved, but
there was evidently a median wedge of wood, not extending so far out us its
212 DR. D. H. SCOTT :
two neighbours (Phot. 12). There is a primary xylem-strand corresponding to
each of the three secondary wedges; the median strand (not well preserved)
begins further out from the pith than the other two and presumably alone
represents the leaf-trace (Phot. 13, /.£.). "The structure is comparable to
that shown in C. americana : see, for example, Pl. 6, Phot. 5. The secondary
wedge corresponding to the trace is however narrower in C. annularis.
One of the adjacent strands lies nearly behind the leaf-trace, and may well
be the reparatory strand from which it has separated (Pl. 7, Phot. 13).
Unfortunately there are no sections available to show the outward progress
Text-figure 1.
Calamopitys annularis (Unger), Solms.
Transverse section of leaf-trace strand, shown at /.t. in Phot. 18. <°, outer edge of secondary
wood of stele. The strand shows the ill-preserved primary xylem in the middle, surrounded
by a broad secondary zone. X about 18. Drawn by Mr. G. T. Gwilliam, Section 98,
Berlin. (Coll. Solms 411).
of the trace; the next phase shown (in another specimen) is beyond the
wood altogether (Phot. 18).
This is in specimen 98. The stele, measured to the outside of the wood,
is 28 x 21 mm. in diameter and the pith (including primary xylem) about
8-9 mm. On one side a portion of cortical tissue is attached, which probably
formed part of a leaf-base.
The pith itself is destroyed ; the primary wood is imperfect, but enough
remains to show that it formed a very nearly continuous zone. Where the
NOTES ON CALAMOPITYS. 213
structure of the individual strands can be made out it is very eccentrically
mesarch, with the centrifugal part of the xylem much reduced as compared
with the centripetal.
The leaf-trace above referred to lies outside the wood, probably in the
pericycle, to one side of the fragment of leaf-base (Phot. 18, Text-figure 1).
It is a large strand, measuring about 4:5 x 2:6 mm. in diameter, the longer
axis being tangential to the stem. The greater part is composed of secondary
wood, which is well developed all round the strand, and is thickest on the
flanks. Very little of the primary xylem at the centre is preserved. A
similar strand is mentioned in this specimen by Solms-Laubach (1896, p. 74),
who, however, describes the secondary growth as feeble and limited to the
outer side, forming an arc. In our case, the secondary wood forms a wide
zone, and seems to be about equally developed on the outer and inner face of
the strand (Text-fig. 1). The structure is obviously comparable to that of
a leaf-trace bundle of C. americana, in the corresponding position (cf. Phot. 10).
On one side of the strand there is an irregular extension of the wood, seen
partly in oblique section. I am doubtful as to its nature; it is possible that
it may represent the other bundle of the trace, the two strands having been
erushed together; this part of the tissue has a macerated appearance, and is
less well preserved than the stele and leaf-base. There is another bundle,
on the opposite side of the fragment of leaf-base, which appears to correspond
more nearly with Solms-Laubach's description, but the preservation is bad.
In the cortex or leaf-base itself two complete bundles are preserved, as
deseribed by Solms-Laubach (p. 75). Each is elongated, approximately in
the radial plane *, and has three internal protoxylem-groups, the latter lying
towards that side of the strand which faces its neighbour. There is, of
course, no secondary thickening at this level. The structure is identical with
that of the bundle figured by Solms-Laubach from C. Saturni (l.c. Taf. 4.
fig. 11) and also agrees with that of the bundles in the leaf-base and petiole
of C. americana (Scott & Jeffrey, 1914 ; Pl. 27. Phots. 4, 5 ; Pl. 36. fig. 9).
A small part of the Sparganum hypoderma is preserved. The whole
structure is that of a Kalymma, as Solms-Laubach has shown. From the form
and arrangement of the bundles it seems evident that the fragment formed
part of a leaf-base, rather than of the mere cortex of the stem.
The same probably holds good for the fragment attached to specimen 97,
which shows essentially the same structure (Phot. 11). Here the bundles
have from two to four protoxylem-groups ; there appear to be three in the
one figured (Phot. 17). Two of the bundles are giving off smaller branches,
with a single protoxylem.
As regards the structure of the leaf-base, the three species, C. annularis,
C. Saturni, and C. americana, seem to be practically identical.
* The fragment is obviously displaced (see Phot. 18).
214 DR. D. H. SCOTT :
The Halle specimen under the name C. annularis (Solms, 420) is remark-
able for two features—the narrowness of the medullary rays and the
smallness of the pith. The specimen (Phot. 19) includes the whole of the
latter and a large portion of the wood, but probably not its full thickness,
The present radius is about 14 mm.
The pith, including the primary xylem, only measures about 2:5 x L7 mm.
(Phots. 19, 20). No stems of C. annularis with a small pith like this are
mentioned by Solms-Laubach, so presumably this specimen only came under
observation after his paper was published. The question arises, is it rightly
named Calamopitys annularis? The primary xylem forms an irregular zone
in which some of the strands are fairly well defined, but appear to be con-
nected by intermediate tracheides. At some places the xylem forms a
continuous band, while elsewhere there are only scattered tracheides between
the primary strands; this difference may simply be due to unequal pre-
servation, It is extremely difficult to make out any protoxylem in the
primary strands; where elements smaller than the rest occur, they lie
towards the outer edge of the strand (Phot. 20).
The pith is badly preserved ; a certain number of largish elements, with
somewhat thick walls and clear lumina, persist and suggest tracheides
(Phor. 20). The radial section confirms this suggestion. This section
extends right across the pith, which contains many long tubes, agreeing in
appearance and size with the tracheides of the primary wood. In the case
of a stem with so small a pith one cannot be certain that the section, though
apparently radial, may not be sufficiently tangential to the pith to pass
through the inner part of the xylem-ring, However, it is probable that some
of the more scattered tubes really represent medullary tracheides.
The secondary wood is remarkable, for it nowhere shows the wide
medullary rays characteristic of the species. Most of the rays seen in trans-
verse section are uniseriate; about half as many are biseriate ; larger
numbers (locally four rows of cells were seen in a single case) are extremely
rare. There is no tangential section of the specimen, but one may con-
jecture that the ray in this form commonly hada middle portion two rows
wide, with a uniseriate extension above and below. The whole appearance
of the wood is very different from that of the typical C. annularis
(cf. Phots. 12, 18). The rays are also very narrow in actual dimensions, the
single ray-cell being commonly about 12 w wide.
The secondary tracheides are small; a rough average giving about 36 u
for the diameter. Consequently the radial section shows few rows of pits on
the wall; usually they are in 1-3 rows, rarely 4. Otherwise they appear
to agree with the pits of the first specimen.
The Halle stem is obviously different in certain points from the typical
form of the species. It is certainly not C. Saturni, for, as will be seen
presently, a specimen of that plant of about the same dimensions as regards
NOTES ON CALAMOPITYs. 215
the pith, shows totally different characters. I think it most probable that
the Halle specimen was rightly named by Solms-Laubach and that it repre-
sents an advanced stage of a small branch, with the structure somewhat
modified as compared with that of the main stems. In spite of the
peculiarities mentioned, the general character of the tissues suggests Calamo-
pitys annularis and no other plant.
The special features which require to be emphasized in the species
C. annularis are: 1. The continuous or almost continuous zone of primary
xylem (already pointed out by Solms-Laubach).
2. The eccentrically mesarch structure of the xylem-strands, with a
corresponding reduction of the centrifugal portion.
3. The probable presence of tracheides embedded in the pith.
4. The secondary thickening all round the leaf-trace strand on its exit
from the wood.
In all these points C. annularis agrees with C. americana.
The division of the leaf-trace into two has not been observed in C. annularis;
there can be no doubt that it takes place, but we do not know in what part
of the course of the trace the division oceurs. The narrow medullary rays of
the Halle specimen, assuming, as seems to be justified, that it is rightly
referred to C. annularis, are interesting, for comparison with other species.
CALAMOPITYS SATURNI, Unger.
This species is well known, from the full and admirable account given in
1896 by Solms-Laubach. It has also been examined by Dr. Zalessky (1911).
Solms-Laubach describes the primary wood in this species as “an irregular
tracheal zone, perhaps interrupted here and there, which swells out in places
into expanded nests, projecting into the pith-parenchyma” (1896, p. 65).
In C. annularis it seemed to him that the primary wood was * more strongly
developed, forming a quite or almost closed ring" (l.c. p. 74). Judging
from the one transverse section of C. Saturni now in my hands, I am inclined
to think that the distinction between the two species is more marked than
Solms-Laubach realized. This section (Pl. 8. Phots. 21, 22) is from the
specimen Berlin, 76, and is among those described by Solms-Laubach (l.c.
p. 71), but not figured *.
Two excellent photographs from another section of the specimen are,
however, given by Dr. Zalessky (1911, Pl. 3. figs. 1, 2). The specimen
(stele only) was 1 em. in diameter ; the section here figured is incomplete,
* I have one transverse and one radial section. The former is labelled: “ Cal. Sat. Q.
Berl. 76. Culm Saalfeld. Berlin Landes Anst. Coll. Solms 422."
The radial section is apparently not from the same specimen, for the inscription is: “29K,
Cal. Sat. ? r. Culm Saalfeld. Berlin Ldesanst. Coll. Solms n. 423.” The? makes the value
of this section doubtful, though it was sent to me by Count Solms as an example of the species
C. Saturni and a tangential section of no. 29 is figured by Dr. Zalessky under that name.
216 DR. D. H. SCOTT :
but shows the whole of the pith and primary wood; the preservation is
remarkably good (Phots. 21 & 22). The pith is quite small, about 1°8 x
1-4 mm. in diameter, including the primary wood.
Around the pith there are six distinct primary xylem-strands ; in addition
there is a seventh. strand entering the wood, and lying immediately outside
one of the circum-medullary strands (Phot. 22). The latter stands rather
further back in tbe pith than usual, but at this level is scarcely, if at all,
separate from the outgoing strand. The pair presumably represents a leaf-
trace with its reparatory strand.
The first point that strikes one about the: primary xylem-strands, as
compared with those of C. annularis, is that they are almost circular in
section and centrally mesarch ; the protoxylem is in the middle of the strand ;
the centrifugal portion is just as well developed as the centripetal, and has
equally large elements (Phot. 22). This holds good without any qualification
for five out of the seven strands; only in the strand entering the wood, and
in a less degree in one other, also partly embedded in it, is there any
reduction of the centrifugal primary xylem. In these two cases it may be to
some extent merged in the secondary layers, as we found in the case of
Archwopitys Eastmanii (Scott & Jeffrey, 1914, p. 347; Pl. 38. fig. 17).
The predominance of centrally mesarch structure is a striking contrast to
C. annularis and C. americana, and a striking point of agreement with
C. fascicularis and C. Beinertiana, so far as the large xylem-strands of the
latter two species are concerned,
Another point about the xylem-strands in this specimen of C. Saturni is
that they are quite well defined and to all appearance perfectly separate
from one another. In no case (except of course that of the leaf-trace
and its reparatory strand) have I found any sign of continuity between
the bundles. The perfect preservation allows the tracheides with their
somewhat thick and distinctly pitted walls to stand out quite clearly from
the thin-walled parenchymatous cells* of the pith. In every case the
xylem-strand forms a definite, more or less circular group of tracheides, while
the whole of the tissue lying between the strands, whether the intervals
be broad or narrow, is parenchymatous and identical with that of the pith
(see Phot. 22; also Zalessky, 1911, Pl. 3. figs. 1, 2). The bundles are, in
fact, just as distinct as in Lyginopteris or Porowylon ; there is no sign of a
continuous xylem-ring.
The whole of the pith appears to consist of thin-walled parenchyma.
Every cell is preserved and I find no indication of medullary tracheides,
except in one doubtful case, in the outer part of the pith. If present at
all, the medullary tracheides must here have been in the last stage of
reduction.
* The wall (or middle lamella?) appears as a clear thin line with irregular deposits of a
dark substance on either side.
NOTES ON CALAMOPITYS. 217
Dr. Zalessky gives photographie figures of another section of the same
specimen (I. c. Pl. 3. figs. 1, 2). ‘This was evidently cut near the one I have,
for all the bundles can be identified (cf. Phot. 22). Both Dr. Zalessky
(l.c. p. 25) and Solms-Laubach (l.e. p. 72) mention in this case a single
bundle embedded in the pith, at a little distance from the secondary wood.
From the former author's photograph (l. c. fig. 2) it is clear that this
bundle is the reparatory strand of the leaf-trace. The leaf-trace itself is here
more embedded in the secondary wood and does not seem to have been
noticed. Presumably the section photographed by Dr. Zalessky was cut
just above ours, at a point where the separation of the leaf-trace from the
reparatory strand was more complete.
In the section figured in the present paper there is another bundle
(Phot. 22, x), which appears to be embedded in the pith ; unfortunately the
secondary wood has been cut away at this point, but about three layers of
cells remaining on the outer side of the strand are clearly parenchymatous.
The strand in question is identical with the left-hand one of the two shown
in Dr. Zalessky’s fig. 2. In his section it appears to be almost in contact
with the secondary wood. It does not seem to be a reparatory strand, for
no leaf-trace is visible.
In the circular form and centrally mesarch structure of most of the xylem-
strands, as well as in their complete isolation from one another, C. Saturni,
to judge from this specimen, differs sharply from C. annularis and C. americana.
There is no reason to doubt that the specimen described is typical of the
species in these respects, thongh in other sections, which I examined in 1901,
the exact limits of the xylem-strands were more difficult to trace, owing to
the nature of the preservation.
Another probable difference is in the occurrence of medullary tracheides,
which are a characteristic feature of C. americana and appear to be frequent
in C. annularis, while there is little or no evidence for their presence in
C. Saturni. This, if established, would be an important distinctive character,
but as regards the two Thuringian species the data are inadequate for a
final decision. One small point remains to be mentioned : in our specimen
of C. Saturni and in some others the medullary rays are often dilated
outwards, as noticed by Dr. Zalessky (l. e. p. 26), while this is not the
case in C. annularis or C. americana. This character is obviously of only
specific value, at most.
The question of the course of the leaf-trace has been discussed above
(p. 208), and the distinction in this respect between C. Saturni and
C. americana pointed out; the data are wanting for a comparison with
C. annularis. On the other hand, the structure of the leaf-trace on its
exit from the secondary wood is very much the same in the last-named
species and C. americana.
ur
218 DR. D. H. SCOTT :
On the whole of the characters available for comparison, it seems clear
that C. Saturni is very distinct from C. annularis and C. americana, while
the two latter species show a remarkably close agreement with one another.
ÜALAMOPITYS AMERICANA, Scott & Jeffrey.
A short summary of the chief characters of this species is all that need be
given here,
Primary xylem-strands eccentrically mesarch, with the centrifugal portion
smaller and smaller-celled than the centripetal. Strands united laterally to
form an almost closed xylem-ring.
Pith “ mixed,” containing a varying proportion of medullary tracheides.
Leaf-trace dividing into two as it passes through the secondary wood.
Each leaf-trace bundle, where it leaves the wood, surrounded by a zone of
secondary thickening.
Secondary wood with high, multiseriate medullary rays, not enlarged
outwards, "Tracheides with several series of alternating bordered pits on the
radial walls,
Leaf-base with the structure of Kalymma, containing a number of mesarch
bundles, each with from 2 to 5 protoxylem-groups.
The details are fully described in the paper by Prof. Jeffrey and myself
(1914), supplemented by the more complete account of the course of the leaf-
trace given in the first part of the present communication.
CALAMOPITYS FASCICULARIS, Scott.
Lristophyton fasciculare, Zalessky.
This species was shortly described, under the name Araucarioxylon
Jasciculare, in 1899 (Scott, 1899, p. 615) and a full illustrated account was
given three years later, when I placed it in the genus Calamopitys (Scott,
1902, p. 332). No new specimens have come to light since then and there is
little to add to the published deseription. "The species is mentioned and a
figure given by Dr. Zalessky (1911, p. 22 ; Pl. 3. fig. 7), who contests its
affinity with Calamopitys and transfers the plant to his new genus Evristophyton.
A brief recapitulation of the principal features may be given here, reserving
the discussion of the systematic position to the conclusion of the paper.
The pith is small (2-3 mm. in diameter) and of very uniform parenchy-
matous structure; it contains no sclerotic nests. A few of the cells have
dark contents and may possibly have been secretory in function. There are
no medullary tracheides.
The pith is surrounded by a ring of about 8 or 9 primary xylem-
strands, of mesarch structure (Scott, 1902; Pl, 3. fig. 1). They are of
very unequal size. Those which are about to enter the secondary wood
as leaf-traces are very large, from 0:8 to 1 mm. in diameter. Those
eut lower down in their course are much smaller, diminishing rapidly as
NOTES ON CALAMOPITYS. 219
they pass down in the pith, to a diameter of 0°25 mm. and less. The
smaller strands are as a rule embedded in the pith, a few layers of paren-
chyma separating them from the secondary wood (l.e. Pl. 3. fig. 2). The
large, outgoing strands are centrally mesarch, like those of C. Saturni
(Le. Pl. 1. Phot. 2); the smaller strands, forming their downward
continuation, become eccentrically mesarch with the protoxylem towards the
inner side, and may even approach an endarch structure.
The course of the leaf-traces has been followed in successive transverse
sections ; the phyllotaxis is 2/5, with short internodes; each leaf-trace, as it
passes down in the pith, eventually unites with a reparatory strand on its
kathodic side (l. c., Diagrams 1-4).
In the outgoing trace, as it enters the secondary wood, the protoxylem
divides into two groups, which may become widely separated (Phot. 23).
The division of the trace as a whole has not been observed, and probably
took place too far out to be shown in the incomplete specimens alone
available.
The secondary wood has essentially a Cordaitean structure, with medullary
rays one cell or locally two cells in width, and of no great height (Scott, l: c.
Pl. 4. fig. 6). The tracheides have from two to four rows of bordered pits on
the radial wall ; the pits are arranged in more regular vertical rows than in
the three previous species ; in the form of the pits there is little difference,
for in C. fascicularis, though often isodiametrie, they are about as frequently
elongated in the transverse direction. The pits of adjacent rows alternate
regularly and are in close contact with one another, so that their outline is
more or less hexagonal (l.c. Pl. 3. figs. 4, 5).
The inner layers of the secondary wood are peculiar, for they consist
largely of short, wide tracheides with numerous irregular rows of pits
(l.c. Pl. 3. fig. 3). In this region the medullary rays are also dilated
and distorted.
It is evident that this species differs in several respects from any of those
previously considered. The significance of these differences will be discussed
after the next and last species has been dealt with.
CarAMOPITYS DEINERTIANA (Goeppert), Scott.
Eristophyton Beinertianum (Goeppert), Zalessky.
This is the old Araucarites Beinertianus of Goeppert *.
The primary wood was first described in 1902 (Scott, 1902, p. 341), and
the characters then observed led me to place the species in the genus
Calamopitys, side by side with C. fascicularis.
The figures given in the paper of 1902 have since been supplemented by five
excellent photographs from Solms-Laubach's Falkenberg specimen, published
* For synonyms and references, see Scott, 1902, p. 344 footnote.
220 DR. D. H. SCOTT :
by Dr. Zalessky (1911, Pl. 3. figs. 3-62). This author associates the species
with C. fascieularis in his genus Eristophyton. Two specimens have come
under observation, Solms- Laubach's stem from Silesia and the British one
collected by Dr. Kidston at Norham Bridge on the Tweed. For the purposes
of the present paper I have also examined sections, kindly lent to me by
Dr. Kidston and Dr. Gordon, of a specimen received by the former from
Mr. Dunlop, and also derived from the Norham Bridge locality. This
fragment agrees so closely in dimensions, appearance, and state of preser-
vation with the original Tweed specimen, that I think it must bea part of
the same stem. The new sections have afforded some fresh data for the
study of the leaf-traces.
Except for a fragment of bark, the specimens investigated include only the
pith and wood. The characters of the species are briefly as follows :—
The pith is large, 13-15 mm. in diameter in the British specimens and
about 8 mm. in that from Falkenberg. A striking feature of the pith is the
presence of sclerotic nests, with radiating cell-rows around them (Phot. 27),
recalling the similar structures in Lyginopteris. This character is entirely
absent in C. fascicularis and in all the other species referred to the genus.
There are no medullary tracheides. Around the pith and in contact with the
secondary wood, numerous primary xylem-strands, sometimes confluent with
one another, are disposed. These strands show the same variation in size as
those of C. fascicularis, the maximum diameter being attained as the strand
begins to pass out into the secondary wood. The xylem-strands, owing to the
large size of the pith, appear less important than in the preceding species, but
their absolute dimensions are not much less, the outgoing strands reaching
a diameter of 0'8 mm. In this part of their course they are centrally
mesarch ; lower down, as the strand diminishes in size, the centripetal portion
becomes reduced, and an endarch structure is assumed. Strands in three
different phases are shown, from the Falkenberg specimen, in Phots. 24-26.
The first is a large strand just entering the wood, with centrally mesarch
structure *. Phot. 25 shows a bundle lower down in its course, reduced in
size, but still mesarch, while in Phot. 26 we see a strand at a still lower level,
very small, and distinctly endarch in structure. These changes are quite
parallel to those in the xylem-strands of C. fascicularis, but in C. Beinertiana
they go further, the centripetal xylem dying out altogether.
. The new sections of the British specimen show the leaf-traces at various
points in their passage through the secondary wood. "l'hree such traces may
appear in the same transverse section (Phot. 27), so the internodes must
have been short, for the trace, after leaving the pith, curves rapidly outwards,
so as to be cut very obliquely in transverse sections of the wood. I have not
* There is a group of small elements suggesting a second protoxylem in an exarch
position. I have not seen this elsewhere and cannot offer any explanation. This strand
and the endarch one have already been figured by Zalessky (1911, Pl. 3. Phots. 4, 5, 6, 6a).
Nub 21. s MM E EC.
NOTES ON CALAMOPITYS. 221
been able to determine the phyllotaxis. In the section figured (Phot. 27)
two traces in almost the same phase (about 2 mm. from the pith) are nearly
opposite each other; the arrangement, however, was certainly not decussate
or distiehous, as shown by the position of other traces. Probably it was a
fairly complex spiral.
I have seen no certain case of the división of the protoxylem in the out-
going leaf-trace. The strand shown in Phot. 28, cut almost transversely in
a tangential section of the wood, appears to have a single median protoxylem.
The secondary wood, as in C. fascicularis, has a Cordaitean character, with
the medullary rays seldom more than one cell in width (Scott, 1902, Pl. 4.
fig. 12). The pits of the secondary tracheides are most often in two rows,
sometimes in a single row ; they are often scattered, and even when in
contaet, do not always show an hexagonal form. Those of the primary
elements are in numerous irregular rows (l: c. Pl. 4. fig. 11).
The species C. Beinertiana clearly has much in common with C. fascicularis,
though the large size and peculiar structure of the pith give it a very
different anatomical habit. Doth species have advanced further in a
Gymnospermous direction than the three first described, and, of the two,
C. Beinertiana shows the greater progress.
SYNOPSIS.
A concise synopsis of the chief characters of the five species may be of
service for reference.
1. Xylem-strands of fairly uniform size, all mesarch.
Rays usually multiseriate.
Petiole with Kalyima structure.
A. Xylem-strands usually eecentrically mesarch, with protoxylem
outwards, connected to form a nearly continuous zone. Medul-
lary tracheides present *.
C. americana (from Kentucky).
C. annularist (from Central Germany).
B. Xylem-strands usually centrally mesarch ; quite separate from one
another. Medullary tracheides probably absent. C. Saturni.
2. Xylem-strands of very unequal size; large and centrally mesarch in
the upper part of their course, becoming small and nearly or quite
endarch lower down.
Rays usually uniseriate.
(Petiole unknown.)
* Proved for C. americana; highly probable for C. annularis.
+ It would be premature to give distinctive characters at present. In the meantime the
localities prevent any confusion.
LINN, JOURN,— BOTANY, VOL, XLIV, T
222 DR. D. H. SCOTT:
a. C. fascicularis. Pith small, with no sclerotic nests.
Smaller xylem-strands embedded in the pith, with their centripetal
xylem much reduced.
B. C. Beinertiana. Pith large, with conspicuous sclerotic nests.
All xylem-strands in contact with secondary wood. Centripetal
xylem dying out in the smaller strands, which thus become endarch.
The course of the leaf-trace has not been included in the synopsis as it is
only adequately known in the two species C. americana and C. Saturni; in
the former the first division of the trace takes place in the zone of thickening ;
in the latter not till the trace has passed beyond this zone.
Affinities of the Species.
We have now to consider the relations of the species among themselves
and especially the question of the division of the genus, proposed by
Dr. Zalessky. As mentioned above, this author separates the two species,
C. fascicularis and C. Beinertiana, under the generic name LEristophyton,
leaving in the original genus C. Saturni and C. annularis ; to these the new
species C. americana must now be added, as its close affinity with C. annularis
is beyond dispute.
Dr. Zalessky, indeed, goes beyond generic separation, and while admitting
a certain analogy and even phylogenetic relation between Calamopitys and
Zristophyton * (Zalessky, 1911, p. 27) he points out that the former shows
clear affinity to Lyginopteride, with some approach to Medulloseze also, but
that no such proximity to Lyginopteris t or, in general, to the Cycadofilices
is found in the species referred to Zristophyton (l.c. p. 24). The question at
issue is thus an important one and not merely a matter of the definition of
genera.
In support of his contention Dr. Zalessky brings forward a number of
arguments of very unequal value ; while some are of undoubted weight and
interest, others are trivial or based on a misconception of the facts.
The author goes so far as to state that except for the mesarch xylem-strands
everything in the two groups which can be compared appears to differ
(10. ps 29):
To clear the ground it may be well to deal first with the less important
arguments, reserving for later consideration the more weighty matters on
which I am to a certain extent in agreement with Dr. Zalessky.
He points out that the xylem-strands are rarely embedded in the pith
in Calamopitys Saturni, while all the smaller strands are so embedded in
* For the sake of clearness I provisionally adopt. Dr. Zalessky's nomenclature in discussing
his position.
+ Lyginopteris, Potonié, is the equivalent.of Lyginodendron as used by Williamsou and
formerly by myself,
ae eC s
ii:
CETEROS
NOTES ON CALAMOPITYS. 223
Eristophyton fasciculare. In the other species of LHristophyton, however,
E. Beinertianum, embedding is at least as rare as in the species left in
Calamopitys. We may cite the small endarch strand shown in Phot. 26,
evidently cut low down in its course, but still in contact with the secondary
wood. This character varies in both groups, and affords no criterion between
them. That it sometimes occurs in Calamopitys proper is a point of interest,
but not one on which I should lay much stress. Its recurrence in Pitys may
indicate that it is rather a primitive character. At any rate it does not
constitute a difference between Calamopitys and Eristophyton.
A second argument is based on the structure of the pith. In Calamopitys,
as the author truly says, the groups of sclerenchymatous cells with dark
contents, so conspicuous in Hristophyton Beinertianum, are completely absent.
But they are likewise absent in the pith of ristophyton fasciculare ;
the possible secretory sacs in the latter plant bear no resemblance to the
sclerotic nests of Æ. Beinertianum ; the presence of these structures in the
pith is a specific and not a generic character, and again affords no distinction
between Dr. Zalessky’s two genera.
In one other instance this author has attributed generic value to a character
which proves to be, at most, only specific. He mentions that in C. Saturni
some of the medullary rays are gradually enlarged outwards, while this is
not the case in the Eristophyton species (l.c. p. 26). But, as we have seen,
it is not the case in Calamopitys annularis either, and is therefore of no
value as a distinctive character between Calamopitys and Eristophyton.
The argument from supposed dimensions has been dealt with on a previous
occasion (Scott, 1912, p. 1027). Dr. Zalessky estimates that in his Callizylon
Trifilievi, a plant which he regards as allied to Eristophyton, the stem when
complete may have attained a diameter of a metre. He thinks it improbable
that the stems of Cycadofilices, with their fern-like foliage, should have
reached such dimensions. The argument is at best a very indirect one, but
Iam quite willing to grant that Eristophyton Beinertianum, at any rate, is
likely to have had a thick stem. If this were so, the fact would not afford
the slightest presumption against the plant belonging to the Cycadofilices.
Besides the case of Medullosa stellata, in which a decorticated stem measured
48 x 45 cm. in diameter (Weber & Sterzel, 1896, p. 25), we may cite that
very primitive member of the Cycadofilices Protopitys Buchiana, a stem of
which, though doubtless incomplete, was also almost a foot and a half thick
(Solms-Laubach, 1893, p. 198). The question of dimensions is, in fact, quite
irrelevant to the issue.
We now come to Dr. Zalessky's more serious grounds for the separation of
Eristophyton from Calamopitys. The chief of these relate to the structure of
the secondary wood. Our author lays great stress on the difference in the
medullary rays, high and multiseriate in Calamopitys, low and usually
uniseriate in Eristophyton. The great difference, in typical cases, is strikingly
224 DR. D. H. SCOTT :
shown by Dr. Zalessky's comparative figures of the tangential sections of
C. Saturni and J£. Beinertianum (loc. cit. Pl. 4. figs. 6, 7). There is no
doubt that this is, in most specimens, a real and important distinction ; the
secondary wood of one of the /ristophyton species, considered by itself,
would at once be assigned to Cordaiteæ, while that of Calamopitys
proper would suggest one of the Cycadophyta or Cycadofilices, At the
same time, if the Halle specimen above described (p. 214) was rightly referred
by Solms-Laubach to C. annularis, the distinction is inconstant, for in that
specimen (a small branch) the rays are about as narrow as in an Zristophyton.
Neither are small and narrow rays by any means unknown among undoubted
Cycadofilices, Thus in Protopitys Buchiana, they are low and usually
uniseriate, at most 2-3-seriate in the middle (Solms-Laubach, 1893,
p. 200, Taf. 6. fig. 4); the same is the case in the polystelie Cladoaylon Kidstoni
(Solms-Laubach, 1910, Taf. 3. fig. 13) and other species (Solms-Laubach,
1896, p. 55, Taf. 3. fig. 3). In Pitys the rays seem to be regularly
multiseriate, and this is here accepted as a generic character; their
width, however, is extremely variable and in Callivylon, which appears to be
nearly allied, they are usually uniseriate (Zalessky, 1911, p. 28).
Another point in the structure of the secondary wood on which Dr. Zalessky
lays some stress, is the character of the pitting (l. e. p. 26, footnote). He
finds that in Lristophyton the pits are in regular vertical series, while the
rows are irregular in Calamopitys Saturni ; in the former they are regularly
hexagonal in form, while in the latter they are elliptical to hexagonal. "The
fissures, he says, are oblique or horizontal in Hristophyton, horizontal in
Calamopitys. I have already, in describing the species, called attention to
the difference in the pitting. As Dr. Zalessky says, the pits of Calamopitys
are intermediate between those of Dadoaylon (Cordaiter) and Lyginopteris,
while in /ristophyton they are typically Cordaitean. The difference is a real,
if rather a fine one, and is undoubtedly a point in which the wood of
Eristophyton approaches that of the true Gymnosperms more nearly than is
the case in Calamopitys proper.
In fact there is no doubt that the species placed by Dr. Zalessky in
Mristophyton have made, compared with the typical Calamopitys, a decided
advance in the Gymnospermous direction, as shown, not only in the general
structure of the secondary wood, but also in certain features of the primary
organization. :
These latter points are not much emphasized by Dr. Zalessky, but seem to
me to be the most important of all. While in the typical Calamopitys species
all the primary xylem-strands are more or less uniform in size and similar in
structure, in the species referred to Kristophyton the strands, as they are
followed downwards in the pith, dwindle rapidly in size and tend to assume
an endarch rather than a mesarch structure. In Æ. fasciculare there is
a near approach to endarchy in the lower part of the strands, while in
JE. Beinertianum this structure is completely attained. I have never observed
NOTES ON CALAMOPITYS. 295
any tendency to endarch structure in the xylem-strands of Calmopitys proper;
where there is any departure from central mesarchy it is in the exarch
direction. This seems to me an important difference; it marks a distinct
step, in the case of the ristophyton species, towards the ultimate disappearance
of the centripetal xylem, which finds its realization in the Coniferous type.
of stem. If the species in question are to be placed in a separate genus, T
should be inclined to rest the distinction rather on the primary characters
just diseussed, than on the difference in the structure of the secondary wood.
The latter hardly affords a constant criterion, unless the case of the Halle
specimen, referred to Calamopitys annularis, with its narrow medullary rays,
can be explained away. I have no objection to generic separation, though
I do not, at the moment, adopt it myself. In my first paper I pointed out
that the reference of the two species C. fascicularis and C. Beinertiana to
Calamopitys must be regarded as provisional until the structure of their
cortex and leaf-bases was known (Scott, 1902, pp. 345, 361). ‘This, unfor-
tunately, is not yet the case, and the question is thus still open. If these
plants should turn out to have Kalymma structure in the leaf-base they
should, I think, remain in Calamopitys ; otherwise their separation will
undoubtedly be necessary.
But, on the broader question of the near affinity between the two groups,
I think the evidence is very strong. So far as the larger (i.e. the outgoing)
xylem-strands are concerned the structure of C. Beinertiana and C. fas-
cicularis is identical with that of C. Saturni (compare for example Phot. 22
with Phots. 24 & 25, or with Scott, 1902, Pl. 1. fig. 2). I have seen no
xylem-strands of this type, large, circular and centrally mesarch, in any other
group of plants. The agreement with C. annularis and C. americana is
less close, for in these species the xylem-strands are as a rule somewhat
eccentrically mesareh with the protoxylem nearer the outer side. In
fact the species form a kind of series, as shown in the synopsis on p. 221 ;
C. annularis and C. americana retain much of the protostelic structure,
with an almost continuous primary xylem-ring and medullary tracheides *.
The xylem-strands are usually eccentrically mesarch. In C. Saturni, so
far as I have observed, there is no continuous primary ring, the xylem-
strands being isolated; medullary tracheides are either quite absent or
extremely reduced; the strands are centrally mesarch. In these respects
C. fascicularis agrees with C. Saturni, but there is the important difference
that in the former the xylem-strands, in the lower part of their course, tend
towards an endarch structure and diminish in size, while the secondary
wood assumes a more Cordaitean character. Lastly, in C. Beinertiana the
change has gone further, for while the outgoing xylem-strands are identical
with those of C. fascicularis and C. Saturni, they attain, as they pass
downwards in the pith, a completely endarch structure, the centripetal
xylem here dying out.
* Certainly in C. americana, probably in C. annularis.
226 DR. D. H. SCOTT :
So far as the primary structure is concerned, the relationship throughout
the series appears perfectly clear. It would of course be desirable to put it
on a broader basis, in particular as regards the course of the leaf-traces, but
data are almost lacking in the case of the ** /Zristophyton" species. All we
know is that in C. fascicularis, at any rate, the protoxylem of the leaf-trace
divided into two in passing through the zone of thickening, just as in
C. Saturni, an indication of the subsequent division of the trace itself.
But corticated specimens are needed before the affinities can be fully
cleared up.
In the meantime, on the evidence already available, it seems to me clear
that the species placed by Dr. Zalessky in Zristophyton are more nearly allied
to the Calamopitys of Unger than to any other known plants, and that the
affinity is closest between C. fascicularis and C. Saturni, while C. annularis
and americana show a more primitive type of structure ; C. Beinertiana, on
the other hand, is the most advanced of all the five species.
Affinities of the Genus.
Assuming that the five species which I have included under Calamopitys
form a natural series of nearly allied plants, we may now consider the
affinities of the group to other families. We have to compare our plants on
the one hand with the Cycadofilices and on the other with the Paleozoic
Gymnosperms.
Among the Cycadofilices * the nearest affinity would appear to be with the
Lyginopterideæ. The comparison has hitherto been made with Lyginopteris,
which, in its ring of mesarch xylem-strands and in the first division of
the leaf-trace, presents considerable analogies with Calamopitys Saturni.
The discovery that certain species of Calamopitys (C. americana and probably
C. annularis) were protostelic suggests a relation to Heterangium, while the
recent observation that several species of //eterangium contained a number of
vascular bundles in the petiole (Scott, 1917) strengthens the analogy, and
tends to remove the most obvious discrepancy between the structure of
Calamopitys and that of Lyginopteridem. At the same time it is clear that
no direct filiation of the two groups is admissible; Calamopitys is at least as
ancient as the Heterangiums, and much more ancient than the polydesmic
species: there is at no point any evidence of a transition from one group to
the other; they form parallel series, which, however, may well have had a
common origin. The Medullosex are more remote, recalling Calamopitys
only in their highly polydesmic petioles ; they show, however, a certain
relation to Lyginopteridee through //eterangium, and all three groups may
probably have sprung from a common source.
* [ use this name in a wide sense, to cover all plants which appear to be intermediate
between Ferns and Cycadophyta whether their fructilication is known or not.
NOTES ON CALAMOPITYS. 227
No other Cycadofilices seem to approach Calamopitys at all nearly. There
is a faint analogy in one point between the * Mristophyton” species of
Calamopitys and Protopitys ; in the latter the leaf-trace strands at the edge
of the pith have a mesarch structure, with the protoxylem very near the inner
surface ; traced downwards there seems to be a transition to actual endarchy ;
there is thus some analogy with the lower part of the xylem-strands in
C. Beinertiana and C. fascicularis, but in all other respects the structure is
totally different (Solms-Laubach, 1893). With Stenomyelon, which has
exarch or nearly exarch xylem-strands, there is nothing in common beyond
the polydesmy of the petiole (Kidston & Gwynne-Vaughan, 1912).
It has been pointed out that the Calamopitys series appears to lead in the
direction of Cordaitales, as shown, not only in the structure of the secondary
wood in the “ Eristophyton” species, but in the tendency to a dying out of
the centripetal xylem, shown in the lower part of the course of the leaf-traces
in those species. We may now enquire how far this advance went, and what
was the particular direction which it took.
I am inclined to think that the advance towards a Cordaitalean type of
organization did not after all go very far. I am chiefly influenced by the
great size of the primary xylem-strand which constitutes the leaf-trace in
C. fascicularis and Beinertiana, especially the former*. There are few plants
even among the Cycadofilices in which the primary wood of the outgoing
leaf-trace bears so large a proportion to the size of the stele, asin C. fascicularisf.
It is almost a Filicinean character, and suggests that the leaf which such
a trace supplied must have been large and very probably fern-like. This
is certainly a strong argument for C. fascicularis, at all events, having still
been one of the Cycadofilices. The case is less striking in C. Beinertiana,
where the stele is so much larger in proportion, but these two species are
admittedly nearly allied, and what is true of the one must in essentials be
true of the other, though in C. Beinertiana the leaves may probably have
been smaller in proportion to the stem.
For these reasons I should regard the whole series of species here
included under Calamopitys as having belonged to Cycadofilices, in spite
of the anatomical advance shown by C. fascicularis and Beinertiana. We
may still ask whether these species show an approach to any special family
among the Cordaitales, as at present known to us.
* Scott, 1902, Pl. 3. fig. 1.
T There is a physiological difficulty involved in the large size of the outgoing leaf-trace.
As it dwindled rapidly in the downward direction, how was an adequate water-supply
maintained ? Possibly the solution is to be found, in the case of C. fascicularis, in
the presence of the short wide tracheides of the inner layers of the secondary wood (Scott,
1902, p. 339; Pl. 3. fig. 3). These layers are in immediate contact with the outgoing leaf-
traces, and appear to be adapted to the storage of water. They may have accumulated a
supply on which the leaf-trace was able to draw. The storage-layers are the innermost of
the secondary wood, and may well have been differentiated at a time when the bundles
supplying the leaves were still actively functional.
7”
228 DR. D. H. SCOTT :
There are no doubt some points in common with the Pityeæ, notably
the centrally mesareh primary xylem-strands. In the genus Pitys most of
the strands are embedded in the pith, as is also the case to a lesser extent
in C. fascicularis and more rarely in C. Saturni. On the other hand
Archeopitys, where the xylem-strands are distributed all over the pith +,
goes much beyond anything that we find in Calamopitys, for there is no
homology between those independent xylem-strands and the medullary
metaxylem-tracheides occurring in the protostelic species of Calamopitys
(Scott & Jeffrey, 1914, pp. 345, 365). The secondary wood of the Pityeæ
usually has broad medullary rays; it is only in Callizylon (the most
ancient genus of the family) that uniseriate rays prevail. The xylem-
strands of Pityeæ are on a small scale compared with those of Calamopitys ;
the presumption is that the leaves which they supplied were likewise
relatively small, as Dr. Gordon has now proved (Seward, Fossil PI. iii.).
The two families are roughly contemporary; there are interesting
analogies between them, but they can hardly be on the same line of descent.
The Poroxylese are a much later group (Permo-carboniferous). The
stem-structure recalls that of Lyginopteris, but the primary xylem-strands
are exarch, and the two strands of the double leaf-trace pass down at the
margin of the pith through several internodes before they unite. The
centripetal xylem dies out below, a point of resemblance to Calamopitys
Beinertiana, The secondary wood is of the Lyginopteris type. The
leaves were large and simple, showing a general similarity to those of
Cordaitez. There is little to connect the family with Calamopitys, though
a certain affinity, through Lyginopterides, may be presumed.
The structure of the secondary wood in the more advanced species of
Calamopitys (Eristophyton) suggests a relation to the family Cordaitem ; we
now know that certain members of this family retained the centripetal
wood of the xylem-strands in the stem (Scott & Maslen, 1910; Maslen,
1911; Scott, 1912, 1918). In Mesowylon no primary centrifugal wood can be
distinguished ; the strands are exarch, as in Poroaylon. The leaf-trace here
also is double, the two strands uniting on entering the pith or lower down,
according to the species. Parapitys Spenceri, which I should also place in
Cordaitez, is interesting, because here the primary xylem-strands, though
very small, are distinctly mesarch (Scott, 1902, p. 358). No doubt all
the Cordaitez are far removed from Calamopitys, but it is not impossible
that they muy represent an advanced stage of the same, or at least a
similar, line of descent.
Zalessky's two genera Canowylon (1911*) and Mesopitys (1911), both
probably of Permian age, in which there are well-marked endarch strands
of primary xylem, may also have some affinity with Calamopitys ; as the
author points out, this is especially probable in the case of Mesopitys, in
T Dr. Gordon has found a similar arrangement in Pitys, Seward, 1917, p. 288,
NOTES ON CALAMOPITYS. 229
which he traces a certain resemblance to Calamopitys (Kristophyton)
Beinertiana. The two genera in question seem to me to come nearest to
the family Cordaitez, but to belong to a somewhat different branch from
that which includes Mesorylon and Cordaites.
The conclusion then, to which our comparisons appear to lead, is that while
the genus Calamopitys, in the wide sense, belongs as a whole to the group
Cycadofilices, in the neighbourhood of the Lyginopterides, its more
advanced species, forming Dr. Zalessky's genus Hristophyton, show some
approach in certain characters towards the class Cordaitales, and especially
the family Cordaitez.
SUMMARY.
In the first part of the paper additional evidence is brought forward as to
the course of the leaf-trace in Calamopitys americana. The trace, after
separating from the reparatory strand, divides into two in traversing the
zone of secondary wood. It thus differs from the trace of C. Saturni, in
which division is only completed beyond the zone of thickening.
The second part of the paper is devoted to a re-examination of the five
species.
In C. annularis the zone of primary xylem is almost if not quite continuous.
The primary xylem-strands are, as a rule, eccentrically mesarch, the
protoxylem lying nearer the outer side of the strand. Tubular elements,
which are probably medullary tracheides, are present in the pith, indicating
a protostelic structure.
Each leaf-trace strand, on its exit from the wood, is surrounded by its own
zone of secondary xylem.
In all these points C. annularis agrees with C. americana.
In the Halle specimen, a small branch referred by Solms-Laubach to
C. annularis, the medullary rays are narrow, resembling those of C. fascicularis
and C. Beinertiana.
In C. Saturni the primary xylem-strands are found to be completely
isolated ; there is no continuous xylem-ring.
The xylem-strands are, as a rule, centrally mesarch, with the centrifugal
and centripetal portions equally developed. There is little or no indication of
medullary tracheides.
The three species, C. americana, C. annularis, and C. Saturni, all had
Kalymma petioles of very similar structure.
In C. fascicularis the leaf-trace strands at their exit from the small pith
are of large size, centrally mesarch and perfectly similar to those of
C. Saturni. Lower down in their course they diminish in size and the
centripetal portion of the xylem becomes much reduced. The protoxylem of
the leaf-trace divides into two in passing through the zone of secondary wood.
The wood has narrow medullary rays and is generally of a Cordaitean
type. Its inner layers consist of short, wide tracheides, suggesting a water-
storing function.
LINN. JOURN.—BOTANY, VOL. XLIV. U
TWEOCO (^ 17. 7 ee ee re ES ho en ee NE M eT
230 DR. D. H. SCOTT:
In C. Beinertiana the pith is large and contains sclerotic nests. The
xylem-strands are similar to those of the preceding species, except that in
the lower part of their course they become actually endarch. Division of
the protoxylem has not been observed. The wood has a Cordaitean character.
A synopsis of the five species is given on p. 221.
The affinities of the species are then considered, with reference to
Dr. Zalessky's proposed separation of C. fascicularis and C. Beinertiana under
the generic name Hristophyton ; the conclusion is reached that, while generic
separation may ultimately be justified, all the five species form a natural
series, in which €. Saturni occupies, in certain respects, an intermediate
position, between C. annularis and C. americana on the one hand, and
C. fascicularis and C. Beinertiana on the other.
Lastly, the affinities of the genus are discussed. While the whole
Calamopitys series should remain in Cycadofilices, the nearest affinity being
with the Lyginopteridez, through //eterangium, the species C. fascicularis
and C. Beinertiana show some advance towards the structure of Cordaitales
and especially of the family Cordaitec.
LITERATURE CITED.
Krpston & GwvyNNE-VavGHaw, 1912.—R. Kidston & D. T. Gwynne-Vaughan : On the
Carboniferous Flora of Berwickshire—Part I. Stenomyelon tuedianum, Kidston. Trans.
Roy. Soc. Edinburgh, vol. xlviii. Part ii. p. 263.
Masten, 1911.— A. J. Maslen: The Structure of Mesoxylon Sutcliffi. Ann. of Bot.
vol. xxv. p. 381.
Scorr, 1899.—D. H, Scott: On the Primary Wood of certain Araucarioxylons. Ann. of
Bot. vol. xiii. p. 615.
Scorr, 1902.—D. H. Scott: On the Primary Structure of certain Palæozoic Stems with the
Daudoxylon type of Wood. Trans. Roy. Soc. Edinburgh, vol. xl. Part. ii, p. 331.
Scorr, 19012.—D. H. Scott: The Structure of Mesoxylon Lomaaii and M. poroxyloides.
Ann, of Bot. vol. xxvi. p. 1011.
Scorr, 1917.—D. H. Scott: The Heterangiums of the British Coal Measures. Journal
Linn. Soc. London, Botany, vol. xliv. p. 59.
Scorr, 1918.—D. H. Scott: The Structure of Mesoxylon multirame. Ann. of Bot.
vol. xxxii. p. 437.
Scorr & Jrrrrey, 1914.—D. H. Scott & E. Jeffrey: On Fossil Plants, showing
cum. from the Base of the Waverley dd of iiti Phil. Trans. Roy. Soc.
B. vol. 205, p. 315. ;
Scorr & Paid 1910.—D. H. Scott & A. J. Maslen: On Mesoxylon, anew Genus of
Cordaitales—Preliminary Note. Aun. of Bot. vol. xxiv. p. 236.
SEWARD, 1917.— A. C. Seward : Fossil Plants, vol. iii. Cambridge.
SoLms-LAUBACH, 1896.—H. Graf zu Solms-Laubach : Ueber die in den Kalksteinen des
Kulm von Glützisch-Falkenberg in Schlesien enthaltenen structur-bietenden Pflanzen-
reste. IL Abhandlung. Botanische Zeitung, Band li. p. 197:
Sorms-LauBacu, 1898.—H. Graf zu Solms-Laubach: Ueber die seinerzeit von Unger
beschriebenen struetur-bietenden Pflanzenreste des Unter-culm von Saalfeld in
Thüringen. Abhandl.d. K. Preussischen Geologischen Landesanstalt, Neue Folge, Part 93.
SorMs-LavBacH, 1910.—H. Graf zu Solms-Laubach : Ueber die in den Kalketeinen í des Kulm,
atc: . IV; Völkelia refracta, Stelovylon Ludwigii. Zeitschrift für Botanik, 2 Jahrgang,
p. 529
NOTES ON CALAMOPITYS. 231
Unerr, 1856.—R. Richter & F. Unger: Beitrag zur Paläontologie des Thüringer Waldes;
2ter Theil, von F. Unger, Schiefer und Sandstein-flora. Denkschriften der K. Akad.
d. Wiss. Wien, Band xi. p. 139.
WEBER & STERZEL, 1896.—O. Weber & J. T. Sterzel: Beiträge zur Kenntnis der
Medullosew. Ber, d, naturwiss. Gesellschaft zu Chemnitz, Bd. xiii.
Zauussky, 1911.—M. D. Zalessky: Etude sur l'Anatomie du Dadoxylon Tchihatcheffi.
Mém. du Comité Géologique, Nouvelle Série. Livraison 68. Petrograd.
ZALESSKY, 1911*.-—M. D. Zalessky : Études paléobotaniques. lre Partie: Note préliminaire
sur le Cenoxylon Scotti, nov. gen et sp., p. 13.
EXPLANATION OF THE PLATES.
All the figures are photographic and frequently need to be examined with a lens.
PLATE 6.
Calamopitys americana.
(From photographs by Mr. W. Tams.)
Series, from below upwards, to show division of the leaf-trace. All the
figures (except 2 and 4) are x about 7.
Fig. 1. Part of stele, with much secondary wood. The single trace (/.t.) is at the margin
of the pith. Section 1 (S. 2944).
2. Primary xylem-strand of leaf-trace with adjacent secondary wood from same
Section. pæ., the partly disorganized protoxylem of the strand. x 37.
3. Two sections higher up. Zt., leaf-trace ; r.,reparatory strand. Section 3 (S. 2946).
4. Leaf-trace, reparatory strand and adjacent tissues from same section, more
magnified. px., protoxylem of leaf-trace ; pz.., that of reparatory strand, through
which a crack passes. x 18.
5. Two sections above the last. The leaf-trace (/.¢.) shows signs of division, but is
still slightly connected with the reparatory strand, r. Section 5 (S. 2948).
6. Four sections above the last. The leaf-trace (/.t.) is evidently dividing iuto two,
Section 9 (S. 9952).
7. Four sections above the last, The leaf-trace has now divided into two (lt. Lt.).
The reparatory strand (».) is still seen behind the trace. Section 13 (S. 2956),
8. Two sections above the last. The two strands of the leaf-trace (lt, Lt.) are now
widely separated, with secondary wood between them, and the whole has moved
considerably further out. Section 15 (S. 2958).
9. Two sections above the last. The two leaf-trace strands (L4, L/.) have moved
further apart and each has its own separate fan of secondary wood. The cortex
is well shown on the left. Section 17 (S. 2960).
10. Three sections above the last. The two leaf-trace strands (44, Lt.) are now
passing beyond the wood of the stele. Each is almost or quite surrounded by its
own zone of secondary wood, Section 20 (3. 2963).
PLATE 7.
Calamopitys annularis.
From photographs (except 14 & 17) by Mr. W. Tams. 14 & 17 by Mr. L. A. Boodle, F.L.S.
Fig. 11. General section of stele, with part of cortex and leaf-base, containing three
bundles (v.^.) attached on the left. x 921. Section 40a (Coll. Solms 413).
12. Stele, from an adjacent section of the same specimen. lt., probable position of
a leaf-trace. x 3. Section 97, Berlin.
15. Part of primary and secondary wood, from the same section as Phot. 11.
lt., probable primary xylem-strand of a leaf-trace. A reparatory strand lies just
behind. pz., protoxylem of one of the eccentrically mesarch xylem-strands of the
ee eee le a ee ee, ee ee ee NNNM S CNN a ee a, a a a ar
"nj
DR. D. H. SCOTT : NOTES ON CALAMOPITYS.
stele. The primary wood is nearly continuous. x about 25. Section 40« (Coll.
Solms 413).
Fig. 14. Double xylem-strand from the same section as Phot, 12. pæ., pæ., the two proto-
bo
jos
22.
23.
24.
26.
27.
28.
xylem-groups of the strand, which appears to be separated a little from the
secondary wood above. x about 30. Section 97, Berlin.
. Radial section through the pith of the same specimen, The longitudinal axis is
horizontal. ¢.¢.¢.', tubes, probably medullary tracheides. x about 16, Section
97R, Berlin (Coll. Solms 413).
. Shows one of the tubes (¢.') in the pith, more magnified. X 38.
. A vascular bundle and surrounding tissue from the leaf-base of the same specimen.
There appear to be three protoxylem-groups (pz.). X 35. Section 97, Berlin.
. General transverse section of another specimen, with a portion of leaf-base,
containing two bundles (v.d.), on the left. Zt., leaf-trace strand shown in detail in
Text-fig., p. 212. x 1$. Section 98, Berlin (Coll. Solms 411).
. General transverse section of the Halle specimen, showing the dense wood, and,
on the right, the small pith (p.). x 3$. Section 761, Halle (Coll. Solms 420).
. Pith, with primary and part of secondary wood from the same section, more
magnified. The primary wood is almost continuous and there appear to be
medullary tracheides. p.., a protoxylem-group ; m.r., one of the uniseriate rays of
the secondary wood. xX about 25,
PLATE 8.
Calamopitys Saturni.
(From photographs by Mr. W. Tams.)
. General transverse section, showing the small pith and primary wood, the broad
secondary. wood with medullary rays dilated outwards, and the remains of the
phloem (ph.). X 18. Section 76, Berlin (Coll. Solms 422).
Pith and primary wood more magnified. The six distinct. xylem-strands round the
pith and the leaf-trace are seen. pz.Lt, protoxylem of leaf-trace ; pz. proto-
xylem of reparatory strand behind it. 2.,a xylem-strand which appears to be
somewhat embedded in the pith. x about 50,
Calamopitys fascicularis.
Leaf-trace in a tangential section of the wood, showing the two widely separated
protoxylem-groups, pz, pr. X 46. (Mr. W. Tams.) Section S. 2474.
Calamopitys Beinertiana.
Part of transverse section, showing a large mesarch xylem-strand leaving the
pith and entering the secondary wood. pz., protoxylem of strand. x 93, From
a section of Count Solms-Laubach's Falkenberg specimen.
. From the same section, showing a xylem-strand lower down in its course, smaller,
but still mesarch, px., its protoxylem. x about 65.
Another xylem-strand, cut still lower down, very small and endarch. pz.,its
protoxylem. X 65.
The above photographs, 24-26, by Mr. L. A. Boodle, F.L.S.
General transverse section of a Norham Bridge specimen. Numerous sclerotic
nests in the large pith. /..-/.t., the three leaf-traces passing through the wood.
X about 23. From a section (No. 1) lent by Dr. Gordon.
Leaf-trace in a tangential section of the wood, from the same specimen. "There
appears to be one median protoxylem. x about 48. From a section (No. 4) lent
by Dr. Kidston, F.R.S.
Phots. 27 & 28 by Mr. W. Tams.
Huth coll.
Tams phot
CALAMOPIT[S AMERICANA.
Tams & Boodle phot. i E Huth coll
CALAMOPITYS ANNULARIS.
Scott.
42 95
lo pt 3g!
(CR!
$3 (M
21, 28.CALAMO
Jovan. Linn Soc.Bor. Von XLIV. Pr.
Huth coll
ESSIGUDAHIS, 24-28. C. BEINERTIANA.
*
ON THE GENERA FUMARIA AND RUPICAPNOS. 233
A Revision of the Genera Fumaria and Rupicapnos. By HERBERT
Wituiam Puastey, B.A. (Communicated by A. D. RENDLE, D.Sc,
(Lond.), F.R.S., Sec. L.S.)
(PrATES 9-16.)
[Read 3rd May, 1917.]
INTRODUCTION.
Ix the course of writing “The Genus Fumaria in Britain,’
Supplement I to the ‘Journal of Botany’ for 1912, it was found necessary to
examine not only those Foreign Fumitories that were most closely connected
with the British forms, but in a general way all the annual species of the
genus of which material could be obtained ; and remarks on the
features of various non-British species are embodied in that work
reviews of the generic subdivisions.
During the winter of 1912-13 several Foreign collections were received
for examination, notably the fine sets of Mr. Charles Bailey and of Ziirich
University, the latter kindly sent by Professor Schinz ; and with this
material for reference (numbering over 2000 sheets), in addition to the
herbaria at the British Museum and at Kew, where some important types
are preserved, an opportunity presented itself to effect an appreciable advance
in the study cf the genus. About the same time valuable supplies of living
plants were obtained from the Riviera and the neighbourhood of Gibraltar
through the kindness of Mr. H. Stuart Thompson, Major A. H. Wolley-Dod,
and Mr. E. G. Baker, and an opportunity was take
summer to visit the Doissier Herbarium at Geneva
types there.
! published as
salient
in the
n during the succeeding
and examine the important
The result of this work is the present paper, the completion of which has
been seriously delayed owing to the pressure of other duties consequent upon
the war. An attempt is now made to offer a general criticism of all the
species of the genus, which it is thought may be useful for reference üs a
Supplement to the Latin edition of Olof Hammar's * Monographia Generis
Fumariarum " (Nov. Act. Reg. Soe. Scient. Upsal. ser. 3. ii. fasc. post. p. 258),
published at Upsala in 1857, taken in conjunction with ©. Haussknecht’s
* Beitrag zur Kentniss der Arten von Fumaria, sect. Splierocapnos," in * Flora,’
lvi. (new series xxxi.) (1873), the writer’s above-mentioned ‘Genus Fumaria
in Britain,’ and certain other works wherein the Fumarie are dealt with in
more or less detail. Hammars earlier Monograph (* En Monografi öfver
Slägtet Fumaria”), which was published at Lund in 1854 and considerably
amended in his later work, is referred to only in a few special cases.
LINN. JOURN,—BOTANY, VOL. XLIV, X
PTT
234 MR. H, W. PUGSLEY : A REVISION OF THE
CLASSIFICATION OF THE GENUS FUMARIA.
In this Revision X. spicata, Linn. and its allies are omitted as they seem
best referred to a separate genus Platycapnos, Bernhardi in ‘Linnea,’ viii.
471 (1833) ; and Discoeapnos, Cham. & Schlecht. in * Linnea,’ i. 569 (1826),
is similarly excluded.
The genus Fumaria, thus restricted, has been commonly regarded as
comprising two sections, viz. :—(1) Petrocapnos, Cosson & Durieu in Bull.
Soc. Bot. France, ii. 305 (1855), et Cosson, Comp. Fl. Atlant. ii. 80 (1883-
1887) ; (=Rupicapnos, Pomel, Mat. Fl. Atlant. 16 (1860), et Nouv. Mat. Fl.
Atlant. i. 240 (1874), as a genus) ; in which the species are usually perennial,
nearly stemless, with leaves mostly radical, a subcorymbose inflorescence, and
strongly tubercular-rugose fruits: and (2) Spherocapnos, DC. Syst. Nat.
Veg. ii. 131 (1821) ; Cosson, Comp. Fl. Atlant. ii. 83 (1883-1887) ; consisting
of the annual species, with elongate stems, cauline leaves, racemose flowers,
and generally less tubercular fruits.
It was at first intended that this paper should deal only with the second
of these sections, as indicated in the notice read before the Society on the
3rd May, 1917; but subsequently, on the expressed wish of the Council, an
extension was decided upon in order to complete a review of the whole genus.
An examination of the species of Petrocapnos, however, revealed that in
addition to the points of distinction noted by Cosson and Pomel, there are
other important floral and fruiting characters by which they may be separated
as a group from the annual Fumarie, and it is consequently felt essential to
‘follow Pomel in treating them as a distinct genus. An account of these
plants follows that of the true Fumitories.
With this view the genus Fumaria becomes synonymous with the section
Sphwrocapnos, DC., and the classification followed is that already adopted for
the British species, coinciding practically with that of Haussknecht’s Revision
in ‘Flora, lvi. (1873), with an emendation of the sequence of the subdivisions.
It is only after considerable hesitation that this system has been preferred to
that of the more finished Monograph of Hammar, and it is readily admitted
that Haussknecht’s two primary divisions of Latisecter and Angustisecte are
unsatisfactory until their definitions are enlarged by the addition of floral
characters and they become converted into Grandiflora and Parviflora
respectively. A third classification, relying on the curvature of the fruiting
pedicel or its absence as a primary basis of segregation, which was adopted
by Boissier in the ‘Flora Orientalis’ owing to the ready obliteration of the
corolla-characters in exsiccata, seems unreliable owing to the inconstancy
of the recurved pedicel in the species in which it is commonly seen, and
unnatural in its association of plants sometimes widely divergent in other
characters.
It will be remembered that Hammar was led to write his Monograph from
GENERA FUMARIA AND RUPICAPNOS, 235
seeing Fries’ fine series of cultivated Fumitories at Upsala, and asa large
proportion of his descriptions are taken from plants subsequently grown at
Lund, he was evidently well acquainted with many of the living forms and
his choice of the corolla-characters as the basis of generic sections cannot be
lightly set aside. But, on the whole, the annual Fumariv seem to fall
naturally into two rather than three large groups. So far as the small-
flowered species are concerned there is no real diserepaney between Hammar's
arrangement and that of Haussknecht, for the Officinales and the Angustisectie
comprise practically the same species, one author relying on the floral
features and the other on those of the foliage with which they are commonly
associated. But while Hammar's sectional definitions of his Officinales, and
also of his Agrariw, fairly cover all the species placed under them (although
in the former the margins of the upper petal are not often truly patent), it is
not so with his third section, Capreolate, which show marked variations in
the corolla. As instances of this, the spreading margins of the lower petal
in F. Bastardii, Bor. (F. media B. confusa, Hamm.) may be cited, or the
normally obtuse upper petal of F. Gussonei, Boiss. (Pl. 9, figs. 1-3). And
when more recently discovered plants are included, a complete series of
transitional forms may be traced through this section, connecting F. capreo-
lata, Linn., to which Hammar's sectional diagnosis of the corolla correctly
applies, with the species of the section Agrarir. At the same time, neither
the Agrariw nor the Capreolate can be said to be similarly connected with
the Officinales.
It may thus be argued, if the floral characters alone are considered, that
the plants placed by Hammar in his section Capreolate differ from the
Agrarie in that the development of the margins of their outer petals is more
or less arrested. In some other important features, such as size of flower and
character of foliage, most of the Capreolate approximate to the Agrarian
species and are plainly different from the Officinales or Parviflore. The
fruiting characters cannot be similarly contrasted, for while a certain degree
of uniformity prevails among the fruits of the Agrariw, which are usually
large and coarsely rugose, and also to a less extent among the Offcinales,
where they are very rarely large, coarsely rugose, or yet quite smooth, in
the Capreolate of Hammar this organ is eminently variable, being in some
species quite small and perfectly smooth, in others larger than in some
Agrarian species or distinctly rugose. As the variations of the fruit in
Hammar’s Capreolate, therefore, give little indication of any group-affinity,
it seems safest to rely chiefly on the foliage and the flowers, and to regard
the section as related to the Agrariv rather than a distinct group equal
in rank to these and the Officinales or Parviflore.
The contention for two natural groups—G@randijlora or Latisectw and
Parviflora or Angustisecte—is further supported by the geographical
distribution of the species. The Agrarie, especially F. agraria and
x»
236 MR. H. W. PUGSLEY : A REVISION OF THE
F. rupestris, bear a superficial resemblance in flower and fruit to the
perennial F. africana, Lamk., of the section Petrocapnos, which Hammar,
who probably had very scanty material before him, included among them,
Like F. africana they are lovers of warmth, and some of them grow in native
habitats on rocks in North Africa, whence they appear to have spread to
cultivated land. Most of them flourish in North Africa, and while they
extend round nearly the whole Mediterranean littoral, they are absent at
any great altitude or distance from the sea. Away from the Mediterranean
F., agraria alone spreads through Portugal, where rigorous winters are
unknown, while F. occidentalis curiously occurs as an isolated outlier of the
group in West Cornwall, where the mildness of the winters, considering the
latitude, is proverbial.
The other large-flowered Fumitories, Hammar’s Capreolate, are likewise
most abundant round the Mediterranean, especially towards the west, whence
they extend over the islands of the North Atlantic. They also range through
France and Britain, but are rare or absent east and north of a line drawn
southwards along the Rhine and across the Eastern Alps and Balkans to the
Black Sea. Their distribution thus agrees generally with that of the Agrarie,
but is less restricted to the south, the Capreolate forms being evidently less
impatient of cold and able to maintain themselves about as far northwards
as the isotherm of 32° F. for the month of January.
The large-flowered Fumitories, regarded as a whole, may thus be considered
as primarily plants of the Mediterranean region, extending northwards in
Western Europe, but practically absent from Central and Eastern. Europe,
and from Asia except in the vicinity of the Mediterranean. It is evident
that they cannot exist where the winter climate is severe.
On the other hand, the Parviffore or Oficinales of Hammar have a
different and much wider distribution and are very much hardier. While
they inhabit almost the whole region occupied by the Grandijlore, they occur
in the greatest abundance and variety in Eastern. Europe, and in Western
and Central Asia, where they extend as far as India and Mongolia. In some
of their habitats they maintain an existence under elimatie conditions very
severe for annuals. F. Schleicheri and F. Vaillantii are found in the Altai
region, and both of these species grow at an altitude of nearly 2000 m. in
the Swiss Alps. Four species, F. parviflora, F. Vaillantii, F. Schrammii, and
F. officinalis, have a remarkably wide distribution, ranging from the Atlantic
far into Asia; but most of the local species are Asiatic, although some rare
outliers are found in the high mountains of Africa. It will thus be seen
that the small-flowered Fumitories are planis of almost the whole Palearctic
region outside the Arctic Circle and excluding China, with a tendency to
predominate towards the east and with isolated species on certain African
mountains.
GENERA FUMARIA AND RUPICAPNOS. 251
From these considerations, morphological and geographical, it has been
decided to follow Haussknecht in dividing the genus into two Sections,
Latisecte or Grandiffora, and. Angustisecte or Parviffora, based mainly on a
combination of foliar and floral characters, as has been done in “ Fumaria in
Britain."
The Section Grandiflora, representing the Agrarie and Capreolatw of
Hammar, is readily subdivided into three subsections, as is done by Hauss-
knecht, by eliminating, as Murales, from the well-marked Agrarian and
Capreolate species the more dubious and intermediate plants, some of which
present distinct individual features, while one or two others show affinities
with certain of the Parviflore.
Of the Section Parviflora (Oficinales of Hammar) the most strongly
marked species are F. parviflora, Lam. and its allies, which were distinguished
in “ Fumaria in Britain? as Microsepale from their very small sepals. Among
the other species, F. officinalis with some allied forms seems to constitute a
natural subsection (Uficinales) owing to the peculiarly shaped and broad
fruits, as well as the larger flowers with less minute sepals. The remaining
species—all well-marked by relatively large sepals—fall into a homogeneous
subsection Latisepale. In these plants the form of the corolla is less
characteristic than in the other Parviflore, and some of them show afiinities
with a few species of the Murales.
With regard to the generic sequence of these subdivisions, the most natural
arrangement appears to be a commencement with the subsection Agraria,
followed by the Capreolate and the Murales respectively. As the Murales
show affinities with the Latisepale these subsections are placed in juxta-
position, and after them the Offcinales, with the Microsepalæ, comprising the
most extreme small-flowered forms, completing the genus.
SPECIES OF FUMARIA AND THEIR DISTRIBUTION.
The number of species of Fumaria recognized in this Revision is 46—
27 Grandiflore and 19 Parviflore, Hammar's number of species in his final
monograph is 24, including F. Heldreichit but without F. corymbosa ; and
Haussknecht gives 35, if F. microstachys and F. microcarpa are included,
Twenty of Hammar's species are retained, three of the remainder, F. affinis,
F. vagans, and F. Gussonei, being united with F. media B. confusa as one
species F. Bastardii, and F. Heldreichii is reduced to a variety of F. Thuretii.
Of Haussknechvs species F. gaditana and F. Pikermiana are made varieties
of F. sepium and F. Thuretii respectively, and F. malacitana is identified
with F. macrosepala ; the remaining 32 retain their specific rank excepting
F. Borwi, which is reduced to a subspecies of F. muralis.
The 15 additional species are variously accounted for. F. Gaillardotii is
segregated from the allied X. major, F. apiculata from F. muralis, and the
Dalmatian F. Petteri from F. Thuretii. Haussknecht’s varieties F. capreola'a
238 MR. H. W. PUGSLEY : A REVISION OF THE
B. intermedia, F. Vaillantii y. indica, and 8. Sehranunii are raised to specific
rank as F. dubia, F. indica, and F. Schrammii respectively. The Canary
Islands plant referred to F. montana, Schmidt, is separated as a distinct
species P. coccinea. F. bicolor, F. Martini, F. purpurea, and F. occidentalis,
described since the publication of Haussknecht’s work, are admitted as new
species. Two Moroccan plants, ranked by John Ball as F. agraria var.
atlantica and subspecies tenuisecta, are described as new species F. berberica
and F. Ballü. F. micrantha var. Parlatoriana, Boiss. is included as
F. bracteosa ; and F. australis is distinguished for the first time.
The continental distribution of these 46 species may be summarized as
follows :—
Of the Grandijlore 21 occur in Europe and 7 are peculiar to that continent.
2 L| hd
Les, Africa ,, 6 a i:
Dios: ASIA 5230 i i
Of the Parviflore 9 , ^ Europe ,, 2 $ "
9 » Africa ” 4 ” ”
13 Asia Du i =
” ,
Of individual countries, the richest in Europe is Spain, where 17 species
oecur—11 Grandiflore and 6 Parvijlore: France has 12 species, 6 of
each section ; Britain 10, 6 Grandiflore and 4 Parvijlore ; Italy 11,
7 Grandiflore and 4 Parviflore ; Balkan Peninsula 16, 9 Grandiflore and
7 Parviflore; Germany 9, 2 Grandifore and 7 Parvijlore; Russia 9,
1 Grandiflora (introduced ?) and 8 Parvifloræ. In Africa, 14 species
(10 Grandiflore and 4 Parviflore) grow in Algeria, and 8 (3 Grandiflore and
5 Parviflore) in Egypt. The number for Asiatic Turkey is 16, viz.:
5 Grandijlore and 11 Parvijlore ; and for Persia 5, all Parviflore. These
figures indicate the predominance of the Grandijlore in the Mediterranean
region and Western Europe, and of the Parrifore towards the East and
North, and in Asia, as pointed out on page 236.
While six species, F. capreolata, F. micrantha, F. officinalis, F. Vaillantii,
F. Schrammii, and F. parviflora, are remarkable for their wide distribution,
a considerable number of species are distinctly local. Some of these are
endemic in insular habitats, such as F. purpurea and F. occidentalis in Britain,
F. coccinea in the Canaries, and F. montana of the Cape Verde Islands.
F. bicolor, of the islands of the Western Mediterranean, is almost a similar
example, althongh it occurs also in Algeria. F. Ballii, F. dubia, F. berberica,
and F. Munbyi all seem to be rare North African plants : F. amarysia and
F. Petteri are confined to Greece and Dalmatia respectively : and among the
Parviflore, F. Janke appears local in Hungary, F. Boissieri in Mesopotamia,
F. microstachys in Egypt, and F. australis in East Africa. Three other scarce
species, F. rupestris, F. macrosepala, and F. sepium, are notable as only
occurring locally on both sides of the strait of Gibraltar : and F. apiculata
and F. Reuteri seem sparingly distributed in the Iberian Peninsula.
GENERA FUMARIA AND RUPICAPNOS. 239
It is probable that in the Old World the range of the more widely spread
species has been extended through human agency, as has obviously happened
in America, where F. agraria and F. capreolata are extensively naturalized
in the southern continent and F. parviflora in Mexico. The remarkable
distribution of F. muralis from Western Europe round Africa to the East
Indies and New Zealand is clearly due to the same cause,
NATURE or HABITATS oF THE FUMARIX.
In north-western Europe, where the conditions of an open association and
free soil such as the annual Fumitories require are now rarely to be met
with in natural habitats owing to the spread of modern civilization, the
plants of this genus are very generally found as weeds of cultivation : and
owing to the moist summers that usually prevail, they may commonly be seen
from spring to autumn according to the tillage of the ground, for the
frequent rains afford continuously a sufficient degree of moisture for the
germination of their hard-coated seeds. In these regions they show
considerable variation in different situations and as the season progresses,
and the diverse forms that they assume have been carefully diagnosed by
Haussknecht in the case of the small-flowered species with which he was
familiar in Germany.
In the Mediterranean region, however, as well as in Asia, where the
summers are normally hot, and dry rather than wet, it is only after the
winter rains that the seeds are able to germinate, and consequently Fumariæe
are generally flowers of spring or early summer only and are liable to less
vegetative change.
In these southern and eastern countries it is evident from the notes of
collectors that they are by no means confined to disturbed or cultivated
ground, for the conditions of environment which they need frequently exist
in nature on an extensive scale. The following records of apparently natural
habitats have been noted :—Among the G'randiflore, F. rupestris from rock
fissures and calcareous rocks ; F. atlantica from sbady rocks ; F, flabellata
from grassy hills and maritime pastures ; F. dubia from ** mountains " near
Algiers; F. macrosepala from shady rocks, rock-slopes and among Chamerops ;
F. coccinea from moist rocks and mountain wood-edges; F. bicolor from
bushy places and slopes near the sea ; and F. sepium var. gaditana from stony
slopes. Of the Parviflore, F. montana from mountain rocks ; F. Aralikii on
rocks in the Crimea (Pallas) ; F. officinalis var. minor on rocks and in stony
places in the Crimea (Pallas & Bieberstein); F. Vaillantii from sandy hills
of the desert in Songaria; F. asepala in stony, uncultivated places ;
F. parviflora in the desert near Bagdad ; F. parviflora var. latisecta in shady
places of Mt. Sinai ; and F. parviflora var. persica on maritime sands of the
Caspian.
ee
ks
240 MR. H. W. PUGSLEY : A REVISION OF THE
From this list it may not unreasonably be assumed that clearly native
localities for a large proportion of the known species will someday be traced,
although in some countries, such as Britain, where very great natural changes
have taken. place, this may no longer be possible, the original stations for
the endemie species being now lost and the plants compelled to maintain
themselves entirely on ground periodically disturbed by cultivation.
The prevalence of the Grandiflorw in the Old Red Sandstone districts of
Great Britain, and of F. Vaillantii and F. parviflora on the Chalk was pointed
out in “Fumaria in Britain," but it has not been found possible to determine
the geological formations favoured by different species outside the British
Islands.
Tae Genus Ztvprcarxos.
Of the plants treated by Cosson as a section Petrocapnos of Fumaria and
by Pomel as a separate genus Rupicapnos, the earliest known is the large-
flowered species discovered in Algeria by Desfontaines and described by
Lamarck in 1788 as Fumaria africana. This fumitory was placed by
De Candolle in Syst. Nat. Veg. ii. p. 132, with Fumaria spicata, L., to form
a section Platyeapnos of Fumaria, characterized by compressed. fruits—an
arrangement probably due to Desfontaines’ description of the silicule as
" comprimée " rather than to any actual knowledge of the plant.
In 1855 the section Petrocapnos was established by Cosson & Durieu in
Bull. Soe. Bot. Franee, ii. p. 305, with four species, one of these being
F. africana. The sectional diagnosis states that the group consists of rock-
plants, mostly nearly stemless perennials, with long-petioled leaves exceeding
the subcorymbose racemes of flowers, which are borne on very long pedicels,
and with compressed, apiculate fruits. The resemblance of these plants to
the genus Sarcocapnos, both in habit and in the flowers of one species, is duly
noted.
Hammar, at the time of writing his Monograph of Fumaria, seems to
have been unaware of this publication of the section Petrocapnos, ind
included the one species known to him, F. africana, in his section Agrarie.
A more adequate account of these plants, though embracing only the four
species already described, appeared in 1860 in a small pamphlet entitled
^ Matériaux pour la Flore Atlantique,” by A. Pomel. In this work the genus
Rupicapnos is established, and in addition to the cha racters of Petrocapnos
noted by Cosson & Durieu, the prevalent gibbosity at the base of the lower
petal is pointed out, as well as the curious elongation and reflexing of. the
fruiting pedicels whereby the seeds are carried down to the elefts of the rocks
in which the plants grow. The wholly adherent endocarp of the fruit, also,
is contrasted with the apically separated and depressed endocarp of Fumaria,
and it is demonstrated that while the amphitropous seed of this latter genus
shows a bowl-shaped hollow: on the upper side, into which the depressed
GENERA FUMARIA AND RUPICAPNOS. 241
endocarp of the fruit fits, in Rupicapnos the upper part of the seed is simply
rounded without any hollow. The absence of the strongly marked fold in
the albumen below the micropyle, as seen in Fumaria, is also noted.
In a second work fourteen years later (Nouv. Mat. Fl. Atlant. i. p. 240,
and ii. p. 379) a further account of these plants is given by Pomel, and fifteen
species are described, seemingly from material collected by himself, except
in the ease of two species which are adopted from Cosson & Durieu. It is
clear from this work that Pomel possessed a considerable detailed knowledge
of the group, and he has left some accurate deseriptions; but unfortunately
he makes no reference to the various exsiccata that had already been
distributed by other botanists.
The later works of Cosson and of other writers have not maintained Pomel's
separate genus Ztupicapnos despite the distinctive features ascribed to it by
its author. The demonstrated differences of the fruit and seed, however,
seem of the first importance, and not only does Rupicapnos lack the depressed
endocarp and hollowed seed of Fumaria but—what is not so clearly stated by
Pomel—the clefts in the mesocarp which give rise to the “apical pits." On
the other hand, the definition of the fruit of Rupicapnos as compressed, in
contrast with the subglobose fruit of Fumaria, cannot be accepted though
endorsed both by Pomel and Cosson. Except in the case of some of the
annual species, the compression of the fruit in Rupicapnos is not appreciably
greater than what usually obtains in Fumaria.
In addition to the fruiting differences it may be observed that the flowers
of Rupicapnos seem to simulate those of Sarcocapnos rather than those of
Fumaria in that the margins of the two outer petals, which are obsolete or
nearly so in the bud stage in Fumaria, are considerably developed at that
period and generally conceal entirely the inner petals. There appears further
to be an essential difference in the structure of the upper petal, for while in
Fumaria the margins or wings (often strongly reflexed upwards) of the
apical gibbosity are truly marginal only for a short distance back from the
apex of the petal, and further towards the base become extra-marginal and
are prolonged in a blunt, lateral ridge, in Rupicapnos this prolongation is
entirely wanting, the whole petal being more dorsally compressed and the
margins themselves never strongly reflexed over the keel.
The presence of a gibbosity or rudimentary spur at the base of the lower
petal renders this group of plants unique in the family Fumariacew in the
possession of an unequally two-spurred corolla; and it is notable that the
second spur is apparently never developed in the very small-flowered species
and not invariably so in those in which it commonly occurs.
The consideration of these aggregate points of distinction renders it
impossible to ignore the claims of Rupicapnos for recognition as a separate
genus ; and as there is a total absence of any connecting links between it and
the annual species of Fumaria, it has been decided to follow Pomel rather
242 MR. H. W. PUGSLEY : A REVISION OF THE
than Cosson, and to retain Jéupicapnos as a genus. Its natural position in the
family Fumariacer is between the genera Sarcocapnos and Fumaria, to both
of which it approximates in certain features though possessed of other
characters found in neither of them.
CLASSIFICATION OF THE GENUS RUPICAPNOS.
The genus Aupicapnos is a small group of plants of restricted geographical
range. So far as can be judged from the sparse herbarium material
available for examination, including, however, the important collection in
Herb. Mus. Paris, kindly lent by M. Lecomte, it embraces under four well-
marked sections a number of critical forms whose limitations are not always
readily defined, just as happens with the better known and more widely dis-
tributed genus Fumaria. Pomel,in his second account of the genus, divides
his species into three subgroups, viz.: perennials with large flowers, perennials
with small flowers, and small-flowered annuals. These form natural sub-
divisions, unconnected, so far as is known, by intermediate forms, and have been
adopted accordingly. The large-flowered species, with curved, purple-tipped
corollas never very broadly winged, somewhat recall Fumaria agraria in
their floral features (Pl. 9, figs. 4, 5). Their flowers are much more showy
than in the other species of the genus, and the beauty of F. africana is
commented on in Lamarck’s original description. It is proposed to place
these plants in a section. Callianthos.
Pomel’s second group, the small-flowered perennials, resembles the
preceding section in habit and in fruit, albeit usually smaller in all its parts.
Its flowers, however, are widely different, not only in size but in colour and
form, and are quite unlike those of the Fumarie (Pl. 9, figs. 6, 7). Their
colouring is uniformly whitish, with a greenish or yellowish suffusion about the
apex, and the apical purple colouring of the inner petals is practically confined
to their interior. The two outer petals are considerably dilated towards the
apex, with spreading margins, and the inner petals are apically winged, some-
times very broadly so. These wings of the inner petals spread horizontally
between the patent margins or wings of the two outer petals in such a manner
as to give to the flowers a triple winged aspect, and the name Tripteryæ
therefore suggests itself for the section.
In this second section X. sarcogapnoides, described by Cosson, is included
by Pomel, with a note that the actual plant was unknown to him. An
examination of Üosson's type in Herb. Mus. Paris shows that its flowers are
quite unlike those of the other members of the section, the outer petals being
conspicuously dilated towards the apex and much longer than the narrowly
winged, purple-tipped inner petals. The corolla indeed resembles that of a
Sarcocapnos in miniature, as stated by Cosson, and it seems necessary to place
this plant in a separate section, for which the name Sarcocapnoides is
proposed.
GENERA FUMARIA AND RUPICAPNOS. 943
Pomel's last section, comprising the annual species, is readily distinguish-
able by its suberect, shortly branched habit and its few-flowered racemes.
Its fruits are often more compressed than those of the other sections, with a
thinner and peculiarly muricate pericarp. It is proposed to name this section
Muricaria after one of the most strongly marked species.
The first-named of these sections, Callianthos, readily lends itself to a
subdivision into two subsections, some of the species, like R. africana,
showing more or less narrow leaf-segments and whitish flowers, while others
have distinctly broader lobes and pale purple flowers. These two subsections
are termed Africane and Pomeliane respectively.
It will be observed that the sequence of sections adopted in the following
account of these plants places Callianthos last in the series. This order
is followed on the assumption that in an account of the family /umariacee
the genus Rupicapnos follows Sarcocapnos and precedes Fumaria.
SPECIES OF Rvproapyos AND THEIR DISTRIBUTION.
In this Revision all of Pomel’s fifteen species are retained (subject to two
changes of name) although none of his types have been seen and synonyms
are cited by him only in four cases. Herbarium material under different
names, however, has been identified with six of his remaining species with
some confidence, and with a strong degree of probability in the case of a
seventh. The other four species have been adopted out of deference to
Pomels judgment and as the diagnoses themselves show an inherent
probability of the plants being really distinct. In addition, five fresh species
are described for the first time, four of these being unknown to Pomel and
the fifth not distinguished by him. The total number of species of Rupicapnos
thus becomes 20. Of these, eight belong to the large-flowered group
Callianthos, five being Africanw and three Pomelianw ; five are species of the
section Tripterye ; two of Sarcocapnoides ; and there are five known annual
species or Muricarie.
The geographical distribution of the genus, which as a whole is confined to
North Africa from Tunis to Morocco, with one species extending to
Andalusia, is of some interest. The species of the section Callianthos
prevail towards the west and are not known in the Sahara districts. They
occur on mountain rocks in many places in the Algerian province of Oran
north of the plain of Schott, and extend eastward at least as far as Milianah,
in the province of Algiers. In ihe west they have been collected at Tetuan,
near Tangiers, and will presumably be found in intermediate stations in the
eastern half of Morocco when that country becomes more accessible. One
species reaches Europe, inhabiting the Andalusian provinces of Malaga and
Cadiz.
The range of the section Tripteryæ, which is almost exclusively Algerian,
lies to the east, and is nearly contiguous with though apparently not over-
244 MR. H. W. PUGSLEY : A REVISION OF THE
lapping that of the section Callianthos. Species of Tripteryx ave found in the
district of Jebel Amour, in the south-east of the province of Oran ; also near
the Saharan border of the province of Algiers, as at Laghouat ; and more
extensively and generally in the province of Constantine, whence they extend
across the Tunisian frontier in the neighbourhood of Kef,
Of the two species of the section Sarcocapnoides, of which little is yet
known, one was collected near Batna, in the central part of the province of
Constantine near the Aures Mountains, and the other at Bou-Taleb, a locality
in the Algerian montane region that has not been accurately located by the
writer.
Three of the annual species (Muricariv) occur near Biskra, in the south of
the province of Constantine, and they apparently grow there in company.
Plants of this group have also been reported from Beni-Souik, near the Aures
Mountains, and from Elkantara, both localities slightly to the northward.
R. muricaria grows in the Saharan region south of the province of Algiers,
as at Metlili and Ghardaia; and X. delicatula apparently to the west of this
region, in the country south-east of the province of Oran.
METHOD or THE REVISION.
It has already been stated that the following account of the Fumitories,
which includes all the known species with their well-marked varieties, and
has been extended to embrace the plants referred to the separate genus
Rupicapnos, is primarily intended for use as a supplement to the Latin
edition of Olof Hammar's Monograph. A complete synonymy is therefore
not printed, and it is to be understood that Hammar’s synonyms are accepted
unless, as happens in a few instances, they are definitely corrected.
A reference is given to the original description of each species and variety,
and if it exists, to an adequate Latin diagnosis. If this is not to be found, a
fresh diagnosis in that language, with italicised contrasting characters alter
the pattern adopted in “ Fumaria in Britain,” is furnished. A number of
Hammar’s diagnoses, which seem satisfactory except perhaps in minor
details, have been accepted, subject sometimes to some supplementary notes ;
but where the Monographer's descriptions are not considered satisfactory or
where the constitution of his species has been altered, fresh Latin diagnoses
have been written. The descriptions of the British species and varieties
dealt with in “Fumaria in Britain” are cited, but, as a rule, have not been
repeated. References are also given to other works (often to Haussknecht’s
revision in * Flora’) which throw light on various species and their distribution.
The known hybrid-forms are mentioned, but as these are usually barren,
evanescent annuals that do not perpetuate themselves, no new names for their
designation have been introduced. The terms “ey parte” and * pro parte”
are applied to synonyms in the usual sense, į. e, when the synonym includes
more than the recognized species and less than that species respectively.
GENERA FUMARIA AND RUPICAPNOS. 245
Figures are cited in all cases where they are thought good enough to be
helpful in identifying the plants portrayed, but it must be acknowledged
that in few genera are the existing plates so generally inadequate, and the
number of really good drawings of Fumariœ is extremely small. Some of
the icones, such as Hammar’s and most of those in the ‘Journal of Botany,’
show little more than detached flowers and fruits. The general poorness of
the figures is no doubt due to the laboriously detailed work involved in
depicting correctly the elaborately dissected foliage and the racemes of
varying fantastic flowers of these plants.
Exsiccata are usually quoted throughout both genera except in the case
of some British species where this has already been done in “ Fumaria
in Britain,” and in certain British varieties of which no authentic material
has yet been incorporated in any public herbaria. When the herbarium
. where the exsiecata has been seen is not mentioned, the plant will frequently
be found either at the British Museum or at Kew. In some instances the
herbarium is named owing to mixtures in sets having been noticed.
The measurements in the following descriptions are taken from dried
specimens unless the contrary is stated. The length of the corolla is that of
the finest flowers, and is reckoned from the apex of the upper petal to the
end of its spur; the form and measurements of the fruit refer to that organ
in the dry condition as seen in profile with its greatest breadth in view. In
life, the flower is slightly larger than when dry, as is also the fruit, which is
then likewise appreciably longer in some species owing to the presence of a
distinet, fleshy neck which disappears by shrinkage as the fruit becomes dry.
The margins surrounding the green keel of the upper petal are uniformly
referred to as wings in the genus Fumaria, and those, almost invariably
smaller, similarly placed on the lower petal are simply termed margins. The
wings are said to exceed the keel when they are more or less reflexed upwards
and sufficiently developed to hide the keel when the flower is viewed laterally.
Fruits are stated to be mucronate or mueronulate when the keel is produced
at the apex into a short point bearing the deciduous style; they are described
as apiculate when there is no projection of the keel and the style itself
becomes detached more or less above its actual base, thus leaving a small
apiculus of varying length.
It may be well to recall that although the size of the flowers is not stated
in Hammar’s monograph, a line depicting their actual length, and from
which the size of the sepals can also be judged, is shown under each of his
figures.
The sign! implies that the plant mentioned has been seen either in life or
in herbaria. Species and varieties admitted, of which no material has been
seen, are marked “non vidi " (n. v.) and shown in square brackets fF
246 MR. H. W. PUGSLEY : A REVISION OF THE
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Vercuin, L. “x Fumaria Burnati” in Rouy's Revue de Dotauique Systématique et de
Géographie Botanique, vol. ii. Asnières. 1904.
Vistani, R, DE. Flora Dalmatica. Vol. iii. Lipsie. 1852.
— Flora Dalmatica. Supplementum, vol. i. Venetiis. 1872.
Viviant, D. Flore Corsieze Specierum Novarum Diagnosis. Genum. 1824,
Waaener, D. Pharmaceutisch-medicinisehe-Botanik, Wien, 1828-1829.
Wienr, R. Illustrations of Indian Botany. Vol. i. Madras. 1840,
LINN. JOURN.—BOTANY, VOL. XLIV. Y
950 MR. H. W. PUGSLEY : A REVISION OF THE
Wionr, R., & G. A. W. AnNorr. Prodromus Flore Peninsule Indize-Orientalis, Vol. i.
London. 1834.
WinLkoww, H. M. Prodromus Flore Hispanice. Supplementum. Stuttgartiw. 1895.
WinLkoww, H. M. & J. LaxGE. Prodromus Flore Mispaniem. Vol. iii. Stuttgartire.
1880.
Wrrnerina, W. An Arrangement of British Plants. Ed.3. London. 1796.
WoopviLLe, W. Medical Botany. London. 1790-1794.
FUMARIA.
Fumaria, Linn, Species Plantarum, ed. 1, 699 (1753), et Gen. Plant. ed. 5,
314 (1754), ex parte; Gaertner, De Fructibus, ii. 162 (1791), ex parte ;
Bernhardi in Linnzea, viii. 471 (1833); O. Hammar, Mon. Gen. Fumariarum, 2
(1857) [Nov. Act. Reg. Soc. Scient. Upsal. ser. 3, ii. fasc. post. 258 (1858) ],
excl. F, corymbosa ; Fumaria sect. Sphewrocapnos, DC. Syst. Nat. Veg. ii. 131
(1821); Willkomm & Lange, Fl. Hisp. iii. 878 (1880) ; Cosson, Comp. FI.
Atlant. ii. 83 (1883-7).
Plante anne caulibus elongatis ramosis diffusis vel scandentibus. Folia
caulina, primordia longe petiolata, reliqua breviter petiolata vel subsessilia,
foliolis plerisque longe petiolatis et segmentis secundariis vulgo petiolatis
rarius decurrentibus irregulariter 2—4-pinnatisecta ; petioli ssepe cirrhosi.
Inflorescentia pedicellis etiam fructiferis omnibus brevibus nee elongatis
racemosa. Bracteæ pedicellos fructiferos sæpe subæquantes nunquam
multototies breviores. Corolla unicalearata ; petalorum exteriorum margines
apicem versus in statu juvenili parum explicati vel obsoleti ; petalum superius
fere semi-cylindricum, calearatum, superne gibbum apicem attingentem
format qui alas basin versus /n jugum rotundatum laterale productas sæpe
atropurpureas habet ; petalum inferius angustum, plus minusve canaliculatum,
haud. calearatum, inferne gibbum apicem attingentem marginibus patentibus
vel erectis circumdatum efficit; petalorum interiorum nervus medius apice
modice alatus est. Fructus monospermus, indehiscens, nuciformis, muticus
rarius mucronatus vel apiculatus, apice per duas rimas que mesocarpium
perforant bifoveolatus est ; exoearpium interdum lieve vel subleve interdum
tubereulis in sicco videndis rugosum ; endocarpium sub stylo a mesocarpio
disjunctum, in hemisphæræ formå depressum ; semen superne in crateris formd
exeavatum, rugd longa infra mieropylam notatum.
Annual plants, often climbing, with long stems and cauline leaves.
Inflorescence racemose with short pedicels. Lower petal never spurred.
Fruit, when dry, with apical pits.
Although Hammar, in his monograph, gives a generic diagnosis of some
length, a few criticisms are necessary, as well as some notes on the import-
ance of the variations of the different organs in considering specific
characters. In the first place, it is to be remembered that Fumitories, like
some other weeds of cultivation, are very susceptible, especially in the
GENERA FUMARIA AND RUPICAPNOS. 2951
vegotative organs, to the influences of environment. This is often seen in
the general habit, which, while perhaps normally markedly characteristic, i«
readily modified and sometimes quite obscured under abnormal conditions.
The foliage of many species shows distinct inherent differences, but
throughout the genus it is exceedingly responsive to surrounding influences,
always tending to become dwarfer and more compact, with narrower, thicker
and more channelled segments under exposure, and larger and laxer in form,
with more ample and flatter segments, in shade or in damp situations. In
some species the lowest pair of leaflets is always shortly petioled, and in a
few others the leaves are notable for long-inucronate or aristate segments,
Both among the Grandiflore and the Parviflore, however, there are several
species in which the leaves are not readily distinguishable. Hammar defines
the leaves as bipinnatisect in the annual species, but in most of them they
are irregularly 3- or occasionally even 4-pinnatisect in normal individuals.
The relative length of the raceme and its peduncle (i.e. the part of the
rachis below the lowest flower) is usually more or less constant in the different
species, but it should be recollected that the two or three earliest racemes of
a plant frequently show longer peduncles than any that follow them, and the
later ones sometimes become less floriferous.
The length of the pedicels, as seen in fruit, and their direction or curva-
ture, are generally characteristic, but in the case of species where this organ
is areuate-recurved in normal plants, it commonly becomes straight and
divaricate in shade.
The form of the bracts and their length as compared with the fruiting
pedicels are fairly constant and important as affording specific characters,
It generally happens, however, that the two or three lowest bracts in a
raceme are somewhat longer than those above them; and occasionally plants
of various species will be met with in which the pedicels throughout are
abnormally short and the bracts relatively more than usually long.
The sepals are valuable for diagnostic characters, for in form, size, and
marginal dentation they usually show little variation in the same species,
except that in shade an elongate condition sometimes obtains in which the
margin may be either more or less cut than usual.
The features of the corolla are of the first importance, but they
are often obscured and have proved the greatest source of confusion
in the genus. This confusion is largely due to the prevalence, especially
in the section Grandiflora, of a tendency under unfavourable environ-
ment to produce cleistogamous flowers with more or less depauperate
corollas (Pl. 9, figs. 8-10). Such flowers always tend to be pale or
whitish in colour, and are sometimes reduced to less than half their
normal size; their outer petals remain coherent till the corolla falls, and
the margins of these petals are often quite undeveloped. ^ Possessing
no nectary, they are regularly self-fertilized, the sexual organs remaining
y2
252 MR. H. W. PUGSLEY : A REVISION OF THE
tightly enclosed within the connate petals. As in this rudimentary
condition the characteristic features of the flower are more or less lost, it is
desirable in the consideration of species to have regard to perfect flowers,
with the corolla fully coloured and winged, and provided with a nectary.
In these perfect flowers the lower petal is generally deflexed and free, so that
the two outer petals are apically distant, in nearly every species. Hammar’s |
definition that this is so only among the Agrariæ, while in the Capreolate and
the Officinales the lower petal cuheres to the others until separated at the base
by the swelling fruit, cannot be accepted, for apically distant outer petals are
frequently to be seen in good flowers in all three classes, as may be inferred
from the observations of Jordan and other authors; and in all of them the
lower petal eventually becomes detached at the base as the fruit develops.
In some of the Agraria, however, the lower petal seems more caducous than
in most other species.
A feature of the upper petal, not noticed by Hammar, but apparently
constant throughout the genus, is that while the apical wings are formed by
the margins of the petal at the very apex and for the greater part of their
length, towards the base of the petal they leave the margin and are finally
produced in a blunt lateral ridge. The tube of the upper petal may be
regarded as laterally or dorsally compressed in proportion as this lateral ridge
is distant or close to the edge of the petal.
When good flowers are present in exsiccata, a difficulty sometimes arises
owing to the form of the corolla being obliterated in the process of drying, so
that its dorsal or lateral compression, the shape of its spur, and the direction
of the margins or wings surrounding the apical keels of the outer petals are
not readily seen. In the living plant these differences are often very marked,
the acute, laterally compressed corolla, with large, curved spur and narrow,
strongly reflexed wings to the upper petal, seen in such a species as
F. capreolata, being widely divergent from the obtuse, dorsally compressed
corolla, with smaller, straighter spur and broad, erect-spreading wings, of
F. agraria. In adequate exsiccata the direction of the wings and the form
of the spur can usually be fairly judged after some experience, but pressed
flowers are often misleading respecting the compression of the corolla-tube.
A feature of the corolla that does not appear to have been sufficiently
appreciated is the constancy of its colour and of the dark marking about its
apex in almost every species throughout the genus. So far as is known,
there is no Fumaria (with the possible exception of F. parviflora, one variety
of which (indicoides) seems to possess essentially pink corollas) in which the
colour of fully developed flowers may be either pink or white, although pink
flowers may become whitish when depauperate or shade-grown, and white
ones eventually entirely suffused with pink or red after fertilization. But
the essential colour, which is best seen in the later bud stage or as the flower
opens, seems to be invariably constant.
GENERA FUMARÍA AND RUPICAPNOS. 253
The dark red or purplish markings about the apex of the corolla vary
greatly in different species, being occasionally quite absent, sometimes
confined to the inner petals and sometimes spreading thence, either partially
or entirely, over the wings of the upper one. This marking is uniform in
every species (in perfect flowers) excepting F. Bastardii and a few of the
Parviflore. Attention is drawn to these variations under the diagnoses of
the generic subsections, and they may generally be clearly seen in well
dried exsiecata. A well-known peculiarity of this purplish colouring is its
tendency to disperse towards the base of the corolla after fertilization and
as the flower fades.
The characters of the fruit are of course among the most important, and
those which are visible in the dry state are usually recognized without
difficulty. A slight emendation of Hammar’s diagnosis of this organ is,
however, necessary, for although the apical separation of the endocarp is
alluded to, no mention is made of its depression in a hemispherical form
fitting the hollowed upper side of the amphitropous seed. In a few species
that are widely distributed, like F. capreolata, F. muralis and F. parviflora,
the shape of the fruit varies considerably, but otherwise there is a general
uniformity in each species. The fleshy neck-like base, particularly conspicuous
in the subsection Capreolate, disappears by shrinkage as the fruit dries, and
the apical profile is liable to change in some species at maturity or in the dry
state owing in some cases to the subsidence of a small mucronulus, or,
in others, where this is larger and more beak-like, to its splitting and thus
becoming retuse or emarginate. The rugosity referred to in the deseriptions,
and also the apical pits, are only seen when the fruit is dry, and the former
is sometimes ill-marked even then in fruits that have not reached maturity.
A monstrous condition of the fruit, in which it becomes swollen to more than
twice its natural size, occasionally oceurs. This has been noticed especially
in exsiceata of F. muralis from Madeira and in Spanish specimens of
F. parviflora.
From the regularity with whieh the more or less cleistogamous flowers in
this genus develop fruit, it may be concluded that self-fertilization regularly
takes place and that Fumitories are not dependent on insect visitors for their
pollination. In the more northern regions of their range, such as Dritain,
insect visitors are apparently rare; but the flowers of F. Bastardii and
F. muralis have been observed to attract the humble-bee. As might be
expected under these conditions, the occurrence of hybrids is not. common,
and such as have been observed are almost invariably isolated individual
plants that are entirely barren and evanescent.
354 -MR. He W. PUGSLEY : A REVISION OF THE
CONSPECTUS OF SPECIES.
Section I. GRANDIFLORA.
Leaf-segments flat and relatively broad, from broadly oval to oblong or lanceolate (except
in F. Balli, F. Thuretii (partim), and F. Reuteri). Normal flowers large, at least 9 mm.
long, exceeding those of F. officinalis, with the wings of the upper petal reflexed upwards
and the lower petal not spathulate (subspathulate in F. macrocarpa and obscurely so in
F. Reuteri),
Subsection I. AGRARIÆ.
Peduncles rarely long; pedicels usually stout. Sepals small or, more rarely, fairly large.
Lower petal with spreading, more or less broad margins. Fruit usually large, coarsely
rugose when dry.
Seres EU-A GRARIÆ,
Fruit distinctly keeled (except in F. alli), shortly beaked, mucronate or mucronulate.
* Flowers white or rosy-white.
1. F. agraria, Sepals small and narrow; inner petals only tipped with purple; fruit
with subemarginate beak.
2, F, atlantica. Sepals small; inner petals only tipped with purple; wings of outer
petals very broad; fruit mucronate.
3. F. occidentalis. Sepals fairly large; inner petals purple-tipped and wings of upper one
similarly blotched; fruit with subemarginate beak; peduncles rather
‘long.
** Flowers pink.
! Wings of upper petal not clearly purple; inner ones purple-tipped.
4. F. rupestris. Sepals long, lanceolate; outer petals with narrow margins; fruit
shortly beaked, rather large or of moderate size.
5. F. Ballii. Sepals very small, oblong; outer petals with broader margins; fruit
strongly mucronate, of moderate size; leaf-segments small and
narrow.
!! Wings of upper petal clearly purple like the tip of the inner ones.
6. F. major. Sepals small; upper petal obtuse with broad wings; fruit subrotund-
obovate, mucronulate.
. F. Gaiüllardotii, Sepals larger; corolla smaller and paler, with upper petal subacute ;
fruit subrotund-quadrate, submucronulate.
-1
Series ORIENTALES.
Fruit obscurely keeled, quite obtuse.
* Fruiting pedicels areuate-recurved.
8. F. flabellata. Flowers white; sepals fairly large; tip of inner petals and wings of
upper one purple; fruit of moderate size; peduncles long.
** Fruiting pedicels straight.
9. F. judaica. Flowers white; sepals small, oblong-lanceolate; inner petals only
shortly tipped with purple; fruit large.
10. F. amarysia. Flowers pink ; sepals small, ovate-lanceolate ; inner petals only obscurely
tipped with purple; fruit smaller.
11. F. macrocarpa. Flowers pinkish, entirely without purple markings; sepals small, nearly
‘ linear; fruit very large, almost globular.
GENERA FUMARIA AND RUPICAPNOS. 255
Subsection II. CAPREOLATAS.
Peduncles long ; pedicels usually stout with long bracts. Sepals large or of moderate
size, Corolla generally narrow with acute upper petal (tip of inner petals and wings of
upper one purple, except in F. dubia and F. coccinea), and lower petal with erect and narrow
margins. Fruit small to large, usually, when fresh, with a distinct fleshy neck narrower than
the dilated tip of the pedicel, and smooth to rugose when dry.
Series Ev - CAPREOLA TE.
Fruiting pedicels arcuate-recuryed or patent. Fruit smooth or nearly so, not large.
12. F. capreolata. Sepals usually large ; corolla narrow, white (often flushed with pink);
fruit obtuse or truncate.
13. F. dubia. Sepals smaller; corolla narrow, pink ; fruit subacute.
14. F. purpurea. Sepals usually large and blunt; corolla broader and more broadly
winged, pink; fruit broader and truncate.
Series MACROSEPALJE.
Fruiting pedicels suberect. Fruit rugose.
15. F.macrosepala. Sepals very large; corolla rather narrow, white, dorsally flushed with
red; fruit large, mucronulate.
16. F, berberica. Sepals large; corolla broader and more broadly winged, white ; fruit
rather large, mucronulate.
17. F. coccinea. Sepals of moderate size; corolla very narrow, almost wingless, red ;
fruit rather small, obtuse or subacute.
Subsection II. MURALES.
Peduncles short or of moderate length, long only in F. bicolor and F. sepium. Pedicels
usually less thickened above than in Subsections I and II (except in F. Thuretii). Flowers
pink, except in F. bicolor and F. sepium; tip of inner petals purple. Sepals small to fairly
large, Lower petal with narrow, erect or spreading margins, Fruit small or of moderate
size, smooth to rugose when dry.
Series SUB- A GRARI Æ.
Sepals not large. Margins of lower petal spreading. Fruit rugose.
18. F. bicolor. Sepals oval, very small; corolla white, becoming pink; inner petals
only tipped with purple; fruit small, densely rugose; racemes few-
flowered on slender peduncles.
19. F. Bastardi. Sepals oval, less small, serrate; wings of upper petal purpie or not;
fruit larger, less densely rugose; racemes many-flowered on shorter
peduncles.
Series EU-MURALES.
Sepals mostly of moderate size. Upper petal with purple wings, lower one with erect or
more rarely spreading margins. Fruit smooth or rugulose.
* Margins of lower petal spreading; flowers large in many-Howered racemes.
20. F. Munbyi. Sepals ovate, dentate; fruit very small, nearly smooth.
21. F. Martini. Sepals oval, subentire ; fruit of moderate size, nearly smooth.
** Margins of lower petal usually erect; flowers in shorter racemes
(except in F. muralis subsp. neglecta).
22. F. sepium. Sepals ovate or oval, subentire, fairly large; corolla rosy-white, large;
fruit smooth, very obtuse.
256 MR. H. W. PUGSLEY : A REVISION OF THE
23. F. muralis. Sepals ovate, dentate below (or in subsp. neglecta subentire); corolla
of moderate size or sometimes large in subsp. Boræi; fruit smooth
or nearly so, very obtuse or more rarely subacute,
24. F.apiculata. Sepals narrow, ovate-lanceolate, subentire; corolla of moderate size;
fruit smooth, shortly apiculate ; leaf-segments narrow and peduncles
very short.
25. F. Petteri. Sepals sub-rhombic, rather large, toothed mostly about the middle ;
corolla of moderate size; fruit rugulose, subacute-elongate; racemes
subsessile.
Series Sup-LATISEPAL.
Sepals fairly large. Wings of upper petal purple; margins of lower one very narrow,
erect or more rarely spreading with obscurely dilated apex. Fruit very small, rugulose.
Leaf-segments narrower than in the preceding species, and flowers smaller in subsessile
racemes.
26. F. Thuretii. Sepals ovate, obscurely repand-dentate; fruit ovate, subacute; fruiting
pedicels usually arcuate-recurved.
27. F. Reuteri. Sepals broader, subentire; corolla paler pink, with longer spur; fruit
narrower, stipitate; fruiting pedicels patent; leaf-segments smaller.
Srorron IT. PARVIFLORA.
Leaf-segments flat or channelled, relatively narrow, from oblong or lanceolate to linear or
setaceous (except in F. montana). Peduncles usually short. Flowers not large, not
exceeding 9 mm. in length, very rarely larger than those of F. officinalis and generally
smaller, with the wings of the upper petal less reflexed upwards than in Section I and
occasionally even patent; and the lower petal more or less spathulate.
Subsection IV. LATISEPALZE.
Bracts longer than the normally stout fruiting pedicels (except in F, rostellata). Sepals
large, at least one-third as long as the corolla and fairly broad. Fruit very small to rather
large, subrotund, neariy smooth to rugose when dry.
* Flowers white, without purple tip; peduncles rather long.
28. F. montana. Sepals ovate-lanceolate; fruit rather large, rugose; leaf-segments
broad.
** Flowers pink, with tip of inner petals and wings of upper one purple ;
peduncles short.
! Fruiting pedicels arcuate-recurved.
29. F. Kralikii. Sepals oval; fruit very small, nearly smooth; leaf-segments narrow.
!! Fruiting pedicels normally erect-spreading.
30. F. micrantha, Sepals nearly orbicular, more rarely ovate; fruit of moderate size, finely
rugose; leaf-segments narrow,
81. F. bracteosa, Sepals ovate; fruit of moderate size, rugose; leaf-segments setaceous
and flowers minute.
82. F. rostellata. Sepals broadly oval; outer petals rostellate or at least acute; fruit
rather small, rugulose ; leaf-segments rather narrow.
Subsection V. OFFICINALES.
Peduncles more or less short. Bracts shorter than the fruiting pedicels (except in
F. microstachys). Flowers pink ; sepals usually fairly narrow and not more than one-third
as long as the corolla; tip of inner petals and wings of upper one more or less purple. Fruit
of moderate size or very large, broad, truncate or retuse, rugose or rugulose when dry.
GENERA FUMARIA AND RUPICAPNOS. 251
-* Racemes usually long, many-flowered ; pedicels not short; fruit of
; moderate size.
33. F. officinalis, — Racemes 10-40 fld.; sepals ovate or ovate-lanceolate; spur of upper
petal rather short; fruit at most very shortly apiculate.
34. F. cilicica. Racemes 30-80 fld. ; sepals lanceolate; spur of upper petal longer and
more curved; fruit shortly but conspicuously apiculate.
35. F. Boissieri. Racemes very dense, with longer peduncles; sepals lanceolate ; spur as
in F. cilicica ; fruit with longer apiculus.
** Racemes short, few-flowered ; pedicels short; fruit very large.
36, F. microstachys. Sepals ovate; flowers smaller than in preceding species; fruit without
apiculus.
Subsection VI. MICROSEPAL.E.
Peduncles short or obsolescent (except in F. australis). Fruiting pedicels stout or slender,
with variable bracts. Sepals very small or even obsolete, rarely one-fourth aslong as the
corolla and always narrower than its tube. ‘Tip of inner petals purple except in some forms
of F. asepala and F. parviflora. Fruit small or of moderate size, mostly subrotund, very
rarely broad, rugose or rugulose when dry.
Series AM BIGU.X.
Flowers pink, 5-7 mm. long; sepals 1'5-2 mm. long; wings of upper petal reflexed
upwards, except in F. australis.
87. F. abyssinica. — Bracts fully equalling the short fruiting pedicels; wings of upper petal
obscurely purple; fruit subacute or obtuse, apiculate.
38. F. australis. Bracts broad, much exceeding the short pedicels: wings of upper petal
obsolescent, deflexed ; fruit subacute, apiculate ; racemes dense, long-
peduncled.
39. F. Janke. Braets rather shorter than the longer pedicels; wings of upper petal
purple; fruit acuminate.
Series EU-MICROSEPAL Æ.
Flowers pink or white, rarely exceeding 6 mm. tong ; sepals minute (‘5-15 mm. long) or
absent; wings of upper petal variable.
* Flowers pink, with wings of upper petal reflexed upwards.
40. F. Schleicheri. Bracts less than half as long as the long, slender pedicels ; flowers deep
rose, with wings of upper petal purple; fruit rather small, apiculate ;
plant slender.
41. F. microcarpa. Byacts more than half as long as the short pedicels; flowers very small,
lighter pink, with wings of upper petal purple; fruit very small,
apieulate; plant dwarf.
42. F. indica. Bracts about as long as the pedicels or longer; flowers purplish pink,
with wings of upper petal at most obscurely purple; fruit of moderate
size, subtruncate, rarely apiculate; plant robust with very decom-
pound foliage.
** Flowers pink, with erect-spreading or patent (rarely clearly purple)
wings to the broad upper petal.
43. F. Vaillantii. Upper petal emarginate with ascending spur; fruit of moderate size or
rather small, obtuse, rugose; plant usually rather slender with linear-
oblong leaf-segments and shortly peduncled racemes.
44, F. Schrammii. Flowers smaller and paler, with emarginate upper petal and deflexed
spur; fruit smaller, apieulate, rugulose; plant usually slender with
narrower leaf-segments and mostly subsessile racemes.
258 MR. Hl, W. PUGSLEY : A REVISION OF THE
* ** Flowers white (sometimes flushed with pink), with erect-spreading,
patent or deflexed wings to the broad upper petal.
45. F. usepala. Sepals usually wanting; outer petals both emarginate, with no purple
marking; fruit apiculate, rugulose; plant slender, with flat, Jinear-
oblong leaf-segments.
46. F. parviflora. Sepals usually present; outer petals not emarginate, the wings of the
upper usually with a purple blotch; fruit often subacute, rugose ;
plant robust, usually with channelled, linear leaf-segments.
Sscrio I. GRANDIFLORA.
Grandiflore, Pugsley, Fumaria in Britain, 5 (1912). Capreolatr et
Agrariw, Hammar, Mon. 24 & 37 (1857) ; Latisectr, Haussknecht in Flora,
lvi. (new series xxxi.), 513 (1873).
Foliorum laciniæ planæ, late ovales usque ad anguste oblongæ vel
lanceolate (in F. Ballii, F. Thuretii (partim) et F. Reuteri angustiores),
sepius mucronate, Flores normales quam in Æ. officinali, L. majores, plus
9 mm. longi ; petalum superius apice superne gibbum efficit, cujus margines,
sicut alæ, valde sursum reflexi (raro erecto-patentes) sunt; petalum inferius
apice inferne gibbum format, qui margines erectos vel patentes nunquam
apice abrupte dilatatos habet. Petala interiora angusta apicem versus plus
minusve sursum curvata.
SunsEcrIO l. AGRARIÆ.
Agrariæ, Haussk. in Flora, lvi. 550(1873); Pugsley, Fum. in Brit. 42 (1912);
Hamm. Mon. 37 (1857), ut sectio.
Pedunculi breves aut longi pedicellis plerumque apice multo incrassatis
præditi. Bracteæ longitudinis variabilis, sæpe tenuiter acuminatæ. Sepala
plus minusve dentata, raro corollæ tertiâ parte longiora vel ejus tubo latiora.
Petala exteriora obtusa vel subacuta ; inferius marginibus patentibus latius-
culis qui superne nonnunquam paulum dilatati apicem attingunt. Fructûs
sæpissime quam in aliis subsectionibus majores, nonnunquam maximi, semper
siccitate plane tuberculato-rugosi.
In this subsection the varied development of the purple colouring about
the apex of the corolla is characteristic of the different species. F. major,
F. Gaillardotii, and F. flabellata have the tips of the inner petals and the
wings of the upper one entirely purple. — /. occidentalis is similarly marked,
except that the purple wings show clear, white margins; in F. rupestris
the colouring of these wings is obscure and inconstant. In F. agraria,
F. atlantica, F. Balli, F. judaica, and F. amarysia the inner petals only are
purple-tipped, although in the first-named species the wings of the upper
petal often become suffused with a dull rose-colour. In X. judaica this dark
tip is distinctly contracted towards the apex, and in X. amarysia it becomes
both short and obscure. Finally, the purple colour is entirely absent in
F. macrocarpa.
GENERA FUMARIA AND RUPICAPNOS, 259
With the exception of F. flabellata, the plants of this subsection generally
show somewhat narrower leaf-sezments than the Capreolatw and the series
Eu-Murales of the Murales.
* Series Eu-Agraria.
Fructis plane carinati (F. Ballii excepta), breviter rostrati, mucronati
vel mucronulati.
1, FUMARIA AGRARIA, Lagasca, Elenchus Pl. Hort. Matrit. et Gen. Spec.
Pl. Nov. 21 (1816) ; Hamm. Mon. 38, excl. var. major (1857) ; Haussk.
in Flora, lvi. 555 (1873) ; Willkomm & Lange, Fl. Hisp. iii. 881 (1880).
(Pl. 9, fig. 1.)
Icon. Hamm. l.c. tab. iv.
Egsice. Porta & Rigo, Iter II Hispanicum, 1890, No. 228, Cartagena !
Bourgeau, Pl. d'Espagne, 1849, No. 21, Seville! Bourgeau, Pl. d’Esp. et
de Portugal, 1853, No. 1756, Algarve! Balansa, Pl. d'Algérie, 1852, No.
628, Mostaganem !
B. mauritanica, Haussk. l. c. 555.
Lesice. Bourgeau, Pl. d'Algérie, 1856, Tlemcen, in Hb. Boissier, ut
F. agraria!
Bracteis pedicellos breves (circa 3 mm. longos) subeequantibus, sepalis
angustis (5 mm. longis, 1 mm. latis), lanceolatis, integriusculis, longe
acuminatis a typo differt.
Wolley-Dod, Fl. Calpensis, No. 240, from Algeciras, seems to approach
this variety.
y. chilensis, Parlatore, Mon. Fum. 73 (1844) ; Haussk. l. e. 556.
This variety, which seems scarcely separable from the specific type
except for its smaller and more markedly beaked fruits, occurs occa-
sionally in Spain, as may be expected, as well as in Chili and Peru.
Judging from herbarium material, it is by no means the prevalent form
of the species in South America.
There is a Portuguese example of F. agraria in the Sloane Collection in
Herb. Mus. Brit. (H.S. 326/1549) labelled “ F. latiore folio spica laxiore e
Lisbon.”
Lagasca’s diagnosis of this species is meagre and unsatisfactory, but
Jordan, in dealing with F. major, Bad. (Pugillus, p. 6), describes an authentic
specimen in De Candolle's Herbarium which leaves no doubt respecting its
identity. Itis tobe regretted that this fine fumitory has apparently never
been adequately figured.
F. agraria is almost unique among the Grandifloræ in the form of its upper
petal which, while much dorsally compressed and very broad about the apex
with erect spreading wings, is remarkably narrowed towards the base and
terminates in a relatively small and laterally compressed spur. Another
notable feature, only seen in the fresh state, is the yellow colour which
pervades the base of the corolla almost till maturity. This peculiarity is
260 MR. H. W. PUGSLEY : A REVISION Of THE
correctly noted in Hammar’s diagnosis as *lutescentia," and the relatively
dull tints characterizing the flowers of this species are probably due to this
yellow suffusion of the rosy-white corolla.
A further distinctive feature of F. agraria is the polished surface of the
dried fruits and their extremely shallow apical pits.
Of other described forms of this plant Haussknecht’s var. algerica, as
represented in Herb. Kew, belongs to F. major, Bad., as might be expected
from the remarks respecting the sepals.
An authentie specimen, also at Kew, of var. elata, Ball in Journ. Linn. Soc.
xvi. 315 (1878), which the author sug
ggests may be identical with F. rupestris
var. laza, Boiss., has the aspect of a distinct, small-flowered variety of
F. agraria, but as its fruit is now missing, further confirmation is desirable
before this can be established.
F. agraria subsp. Merinoi, Pau ap. Merino, Fl. Galicia, Suppl. iii. 508 (1909),
seems from its description to differ essentially from the type only by its
linear-lanceolate sepals; and hence it is perhaps a form of Haussknecht’s
variety mauritanica.
Material of F. agraria has been observed from Spain, Portugal, Morocco,
Algeria, Tunis, and Sicily, as well as from South America, where it seems to
be extensively naturalized.
2. FUMARIA ATLANTICA, Cosson & Durieu ap. Haussk. in Flora, lvi. 558 (1873).
F. agraria y. atlantica, Cosson, Comp. Fl. Atlant. ii. 90 (1883-1887),
non Dall.
Fwsice. Bourgeau, Tizi Tsennent, Alger, 1856, et cult., in Hb. Boissier !
E. G. Paris, Iter Boreali-Africanum, 1869, No. 205, Sidi Mecid, in cultis,
Constantina, ut F. agraria! Cosson, Djebel Zaghouan, Tunis, 1883, in
Hb. Kew !
Fumaria annua, robusta, foliis irregulariter. 2-3-pinnatisectis, foliolis in
lobos oblongos acutos vel mucronatos fissis. Racemi lawiusculi, sub-20-flori,
pedunculos mediocres vie superantes. Bractee lineari-oblongw, cuspidate,
pedicellos 5-6 mm. longos erecto-patentes paululum incrassatos subaquantes.
Sepala, 3—4 mm. longa, 15-2 mm. lata, ovato-lanceolata, acuminata, parce
irregulariter serrato-dentata, albida, nervo dorsali angusto rubieundo, coroll»
tubo'angustiora. Corolla maxima, 14-16 mm. longa, roseo-albida ; petalo
superiore obtuso, lato, alis albidis latissimis brevibus patenti-reflexis apicem
attingentibus et calcare breviusculo prædito; petalo inferiore obtuso margini-
bus latissimis (4-45 mm. lato) patulis albidis apicem attingentibus sub-
spathulato ;T petalis interioribus sursum curvatis apice atropurpureis. Fructýs
maxim, 5*5 mm. longi, 3 mm. lati, subrotundo-obovati, carinati, cum mucrone
distincto persistente obtusiusculi et inferne angustati, in sicco apicis foveolis
latis obscuris dense tuberculato-rugosi.
OE ea eR
GENERA FUMARIA AND RUPICAPNOS. 261
B. platyptera, var. nov.
Exsice. Reverchon, Plantes d'Algérie, 1898 (Kabylie), No. 3, ut F. vagans,
Jord.!
A typo differt bracteis brevioribus pedicellos dimidios vix superantibus,
floribus paulo minoribus, et presertim fructibus multo minoribus circa
2:75 mm. longis ac latis eum mucrone brevissimo obtusissimis.
Hee species, quee Algeriam et Tunisiam habitat, per flores pallidos maximos
et petalorum exteriorum (preesertim inferioris) alas latissimas notabilis est.
A F. agrariá sepalis tenuibus anguste carinatis, corollis latioribus, fructuque
plus minusve mucronato densius rugoso differt. F. major, Bad. corolla
angustiore alis atropurpureis reflexis rosea et fructibus minoribus obtusioribus
precipue distinguitur, sed quod ad fructum pertinet, varietati platyptere
simillima est.
The diagnosis of this fumitory, the finest-flowered of all the genus, is
based on Boissier’s specimens, together with Cosson’s plant at Kew, the
E. G. Paris example in Herb. Mus. Brit., which shows extremely fine flowers
but unfortunately no fruit, and Reverchon’s plant in Mr. C. Bailey's
collection. The last-named bears fruit more like that of F. major than of
Boissier’s type, but the remarkably broad wings and the colouring of its
corolla render it impossible to mistake its closest affinity, and it is therefore
placed as a variety under F. atlantica.
3. FUMARIA OCCIDENTALIS, Pugsley in Journ. Bot. xlii. 217 (1904);
Fum. in Drit. 43 (1912).
Icon. Journ. Bot. l.c. tab. 462.
Tæsice. Dörfer, Herb. Norm. No. 4814!
This species is perhaps the F. vulgaris corubensis alba (sic) of Parkinson
(Theat. Bot. 287 (1640)), noted as growing in cornfields in Cornwall.
In addition to the characters already given whereby this plant may be
distinguished from F. agraria, Lag., it may be remarked that the colour of
its corolla is a elear rosy-white, quite without the yellow and greyish tints
which mar the brightness of the flowers of Lagasca's plant, and its upper
petal is less obtuse with the wings distinctly reflexed upwards as in F. major,
Bad.
4. FUMARIA RUPESTRIS, Boissier & Reuter, Pugillus, 4 (1852); Hamm. Mon.
40 (1857) ; Haussk. in Flora, lvi. 556 (1873).
Icon. Hamm. l.c. tab. 6 (mala).
Exsicc. Boissier & Reuter, Iter Algeriensi-hispanicum, 1849, Tlemcen, in
Hb. Kew! Warion, Pl. Atlant. Sel., 1878, No. 112, Tlemcen! Reverchon,
Pl. de l'Andalousie, 1890, No. 569, Ronda, ut F. agraria !
Boissier's var. laxa (F. arundana, Boiss. ap. Willkomm & Lange, FI.
Hisp. iii. 880 (1880)) appears to have been described from a shade-form,
B CONCOUN M
^"
262 MR. H. W. PUGSLEY: A REVISION OF THE
Specimens collected at Ronda by Mr. C. Bucknall in 1907 do not accord with
Boissier’s diagnosis and seem indistinguishable from well-grown African
examples.
Haussknecht’s forms diffusa and umbrosa (Flora, l.c.) also appear to be
shade-forms, and it may indeed be doubted whether Boissier’s original
account of the species was taken from a normal plant.
In well-grown individuals, the corolla of F. rupestris is pale pink, with an
occasional suffusion of purple in the wings of the upper petal. When shade-
grown, it becomes whitish, with the purple colour confined to the tips of the
inner petals as in F. agraria.
F. rupestris is remarkable among the Agrari for the narrowness of its
corolla and the wings of the outer petals, and on this ground alone might be
placed with almost equal propriety with the Murales near F. Bastardii, Its
fruit, however, although not very large, is of markedly Agrarian form.
F. rupestris var. maritima, Battandier in Bull. Soc. Bot. Fr. xxxii. 336
(1885), is apparently referable to F. bicolor, Sommier.
-
9. Fumaria BALLI, sp. nov. (Pl. 10.)
F. agraria subsp. tenuisecta, Ball in Journ. Bot. xi. 297 (1873) ; Spic-
Fl. Marocc. in Journ. Linn. Soc. xvi. 316 (1878).
Exsice. J. Ball, Iter Maroceanum, Ait Mesan, 1871, in Hb. Kew!
Fumaria annua, ut videtur, habitu laxiusenlo et satis robusto. Folia pallide
virentia, irregulariter. J-4-pinnatisecta, foliolis in lacinias parvas planas
angustas lineari-oblongas acutas vel mueronatas fissis priedita. tacemi
laxiusculi multifori (15-25-flori), pedunculos breves multo superantes. Bractew
lineari-lanceolate, obscure serrate, tenuiter acuminate, pedicellis fructi-
feris erecto-patentibus longiusculis apicem. versus modice incrassatis dimidio
breviores, sed infime interdum paulo longiores. Sepala minima, circa 2 mm.
longa, 1 mm. lata, oblonga, acuminata, obscure dentata, preter nervum
angustum viridiusculum rosea, in fructu juniore persistentia. Corolla
12-14 mm. longa, rosea, ut in F. rupestri angusta ; petalo superiore alis roseis
reflexis carinam. vix superantibus sed apicem attingentibus et calcar tenue
versus longe productis obtuso; petalo inferiore obtusiuseulo marginibus
latiusculis patulis apicem attingentibus ; petalis interioribus sursum curvatis
apice purpureis. Fructús modici, sine mucrone 2:5 mm. longi, 2 mm. lati,
subrotundi, obsolete carinati, obtusiwseuli et persistenter grosse mueronati,
inferne multo angustati, siccitate apicis foveolis obscuris sed latiuseulis
dense sed tenuiter tuberculato-rugosi.
Heee Fumaria distinetissima quie subsectionis Agrariarum flores fructüsque
habet, per folia decomposita eis F. Avalikii et F. micranthe similia mirabilis
est. Inter afħines F. rupestri proxima est, sed proeter foliorum differentiam
bracteis brevioribus, sepalis minimis et fructibus minus rugosis, grosse
mucronatis potius quam rostratis, facile distinguitur.
GENERA FUMARIA AND RUPICAPNOS, 263
F. Ballii in Mauritania habitat, ubi cl. J. Ball in regione inferiore
Atlantis Majoris in convalle Ait Mesan ad 1400 m. specimen unicum legit.
This remarkable and very rare Fumitory has been named after its
discoverer to avoid confusion with F. tenuisecta Syme, Eng. Bot. ed. 3, i. 113
(1863).
The accompanying plate has been drawn from the single known specimen
in Herb. Kew.
6. FUMARIA MAJOR, Badarro in Moretti, Bot. Ital. i. 10 (1826); Jordan,
Pugillus, 6 (1852) ; Haussk. in Flora, lvi. 552 (1873), ex parte ; Rouy &
Foucaud, Fl. France, i. 176, ut forma F. agrarie (1893). F. agraria B.
major, Hamm. Mon. 28 (1857).
F. major floribus dilute purpureis, Magnol. Hort. Monspel. 82 (1697). The
Sloane Herbarium contains, under Magnol's name, two specimens of this
species, H.S. 58/146 & H.S. 166/322, the former from Montpellier.
Icon. Reichb. Plante Critics, i. 1-113, No. 1222.
Tæsice. Billot, Fl. G. & G. No. 3308! Société Dauphinoise, 1880, Nos.
2354! et bis!
Fumaria annua, robusta, basi ramosa, internodis longis laxa et diffusa, raro
petiolis cirrhosis scandens. Folia irregulariter 2-3-pinnatisecta, foliolis in
lobos oblongos acutos vel mucronatos fissis preedita. Racemi plus minusve
laxi, 10-20- (raro plus 15-) flori, pedunculos srepissime breviusculos superantes,
Bractew lineari-oblongre, apice interdum serrulatze, cuspidate vel acuminate,
pedicellos (ructiferos apice inerassatos suberectos vel patenti-erectos subequantes.
Sepala eis F: officinalis similia, 3-3°5 mm. longa, 15-2 mm. lata, ovato-
lanceolata vel ovata, vix peltata, acuminata, irregulariter /nciso-dentata, praeter
nervum dorsalem rubicundum rosea, corolle tubo angustiora. Corolla magna,
12-14 mm. longa, rosea; petalo superiore obtuso, lato, dorsum compresso,
calcar deflexum versus suleato, alis intense atropurpureis reflexis brevibus
apicem attingentibus et carinam multo excedentibus preedito ; petalo inferiore
marginibus latis patulis roseis apicem attingentibus obtuso et sæpe sub-
spathulato ; petalis interioribus sursum curvatis apice atropurpureis.
Fructús magni, 2:5—3 mm. longi ae lati, subrotundo-obovati, obtusissimi vel
subtruneati, cum mucronulo interdum obsoleto carinati et paululum compressi,
in vivo inferne in stipitem obsoletum pedicelli apicem subequantem angustati,
siccitate apicis foveolis latis obscuris dense (uberculato-rugosi.
B. spectabilis, Rouy, Suites à la Fl. de France, i. 31 (1887). F. spectabilis,
Debeaux, Recherches Fl. Pyr.-Orient. i. 20 (1878) ; Rouy & Foucaud,
Fl. Fr. i. 176, ut forma F. agrarie (1893).
Ewsice. Magnier, Fl. Selecta No. 2387 !
A typo differt foliorum segmentis angustioribus lanceolatis vel lineari-
oblongis ; racemis densioribus; corollis pallidioribus minoribus (circa 12 mm.
264 ` MR. H. W. PUGSLEY: A REVISION OF THE
longis) petalo superiore angustius alato apiculato ; fructibus obsolete
mueronulatis.
Of F. spectabilis, Bischoff, Del. Sem. Hort. Bot. Heidelberg, p. 4 (1849), no
authentie specimen has been seen, but plants raised from seed received from
Jordan under this name seem identieal with the narrow-leaved Pyrenean
form described by Debeaux and other French authors.
Although Bischoff says of his F. spectabilis “ patria ignota," Jordan
(Pugillus, p. 6) states that it is indigenous in Istria and South Italy—an
assertion copied by later writers. The examples seen from these countries,
however, appear distinct from the French F. spectabilis and identical with the
allied forms inhabiting Greece and Syria. These are separated as a species
under the name of F. Gaillardotii, Boissier,
y. algerica, var. nov.
F. agraria var. algerica, Haussk. l.c. 555 (1873).
Jvsice. Bourgeau, Pl. d'Algérie, 1856, Maison Carrée, Alger, Cosson, 1854,
ut F. agraria! | Reverchon, PI. d'Algérie, Kabylie, 1896, No. 3, ut F. muralis!
[The sheet of this in Herb. Zurich is a mixture of the true plant with
F. flabellata, Gasp.] | Société Dauphinoise, 1879, No. 286 bis, ut F. agraria!
Foliorum segmenta quam typi paulo angustiora. Racemi primo densi
tandem elongati, multifori (ssepius plus 20-, nonn unquam 40-flori) Bracteze
pedicellis longiuseulis (5-6 mm. longis) breviores. Sepala 3-4 mm. longa.
Corolla speciosa, 13-15 mm. longa, petalo inferiore marginibus quam in typo
latioribus usque ad 3 mm. lato. Fructis obtusiusculi potius quam obtusissimi
mucronulati.
Tt is not easy to understand why Haussknecht placed this showy African
plant under F. agraria. Not only by its pink, dark-tipped flowers, but
through its toothed sepals and its densely tubercular-rugose fruits it is closely
related to the plant of Badarro.
OF the specific type two forms, perhaps varietally distinct, seem to be
widely distributed in the Riviera: one a dwarf, compact plant, with relatively
small flowers and fraits, the former very deeply coloured; the other a
stronger, laxer form, with larger but lighter-coloured flowers in laxer
racemes, and larger fruits.
When growing, the rich pink, almost black-tipped flowers of F. major are
very handsome.
Hybrid individuals between this species and F. capreolata, apparently
quite barren, have been collected and distributed from the South of France
under the name of F. Burnati, Verguin in Rouy, Rev. Bot. Syst. ii, 122
(1904).
The range of F. major extends from Western Liguria across the South
of France into Catalonia; the var. spectabilis grows in Narbonne and the
Eastern Pyrenees, also in South Russia (Astrakan, Becker, 1876!) ; the
yar. algerica in Algeria,
GENERA FUMARIA AND RUPICAPNOS. 265
7. Fumaria GarrrAnpoTIL, Boissier, Fl. Orient. i, 139 (1867). F. major.
auct., ex parte, non Badarro ; F. spectabilis, auct., ex parte, non Bischoff ?
F. drepanensis, Lojacono Pojero in Malpighia, fasc. ii.-iii. 110 (1906).
Ewsiec. Neugebauer, Fl. Exsicc. Austro-Hungarica No. 2898, ut F. major !
Heldreich, Herb. Fl. Hellenieze No. 68, ut F. major! Todaro, Fl. Sicula Exsice.
No. 222, ut F. major !
Fumaria annua, robustissima, internodis longis parce ramosa, diffusa vel
petiolis cirrhosis scandens. Folia irregulariter 2-3-pinnatisecta, foliolis in
lobos cuneiformes mucronatos, oblongos obtusos vel rarius lanceolatos acutos
fissis. Racemi laziuseuli, 10-16- (ssepius 10—12-) flori, pedunculos crassos
superantes. Bractew oblong, cuspidatie, preter nervum viridiusculum albidz,
pedicellos fructiferos suberectos apice valde incrassatos subuquantes. Sepala
3-4 mm. longa, circa 2 mm. lata, ovata, peltata, acuta, plus minusve inciso-
dentata, preter nervum dorsalem obscurum albida, corolla tubo æquilata.
Corolla 10-12 (rarius 13) mm. longa, pallide rosea; petalo superiore obtusiusculo,
calcare adseendente et alis atropurpureis reflexis apicem | attingentibus
carinamque paulo excedentibus preedito ; petalo inferiore marginibus latiusculis
patulis roseis apicem vix attingentibus subacuto ; petalis interioribus sursum
curvatis apice atropurpureis. Fructús magni, 2:5 mm. longi et paululum
latiores, subrotundo-quadrati vel subrotundi, obtusissimi truncati vel raro
subemarginati, obsolete mucronulati et carinato-compressi, in vivo, ut videtur,
in stipitem quam pedicelli apex multo angustiorem abrupte angustati, siccitate
apicis foveolis mediocribus obscuris dense tuberculato-rugosi.
He species F. majori, Bad. valde affinis est, sed differt habitu etiam
robustiore, pedunculis pedicellisque crassioribus, bracteis sepalisque latioribus,
corollis pallidioribus minoribus petalo superiore angustius alato calcareque
majore preeditis, et fructibus subrotundis potius quam obovatis magis
carinato-compressis foveolisque minoribus notatis.
F. agraria, Lag. preter floris majoris colorem formamque sepalis plus
minusve lanceolatis apice subintegris diu persistentibus et fructüs valde
compressi carina in rostrum emarginatum producta facile dignoscitur.
Floris colore et magnitudine, sepalis majoribus, fructibus retuse rostratis
F. occidentalis, Pugsley, etiam diversa est.
F. rupestris, Boiss. floribus angustis, sepalis longissimis et fructibus
acutiusculis minoribus cum F. Gaillardotii confundi non potest.
F. Gaillardotii in Syrià, in Egypto, in Cretà (Heldreich, Heracleum, 1846,
ut F. flabellata!), in Greecid, in Dalmatia, in Istria, in Sicilia et forsan in
Italià australi invenitur.
F. Gaillardotii, although obviously near F. major and generally united
with it by recent authors, seems to be a more Eastern plant which is specifi-
cally distinct owing to several minor but constant points of divergence, and
especially by its different sepals and corolla. In spiteof its wide distribution
it appears to show little variation, except in the breadth of the leaf-segments,
LINN. JOURN.— BOTANY, VOL. XLIV. Z
266 MR. H. W. PUGSLEY : A REVISION OF THE
and its general aspect is that of a coarse and rampant F. major with remark-
ably poor flowers. It is notable that neither Boissier nor Haláesy recognizes
the identity of the Greek with the Syrian form.
There is a specimen of this plant in Herb. Mus. Brit. “ F. agraria, Lag.—
In arvis prope Nebi-Seidone, circa Sidonem Syris, d. 13 Martii, 1860, coll.
Gaillardot.”
** Series Orientales.
Fructüs obscure carinati, omnino obtusi.
8. FUMARIA FLABELLATA, Gasparrini in Rendic. Accad. Scien. Napoli, i. 51
(1842) ; Hamm. Mon. 41 (1857) : Haussk. in Flora, lvi. 542 (1873).
F. alexandrina, Gasp. in Rendic. Accad. Scien, Napoli, i. 50 (1842), non
Ehrbg. ex Hammar, pro parte ; F. capreolata var, 8, Parlatore, Mon.
Fum. 77 (1844) ; F. Gasparrinii, Bubani, Fl. Pyr. iii. 276 (1901) ?
Icon. Hamm. l. c. tab. 5.
Exsice. Todaro, Fl. Sicula Exsiec. No. 221, Palermo! Reverchon, Pl.
d'Algérie, 1896 (Kabylie), No. 103, ut F. capreolata !
This beautiful fumitory, which bears a resemblance to F. capreolata owing
to its long-peduncled racemes of white, dark-tipped flowers borne on recurved
pedicels, is included by Haussknecht with his Capreolate. Since the corolla,
however, is distinctly Agrarian in form, it seems preferable to follow Hammar
in placing it here. As in F. capreolata, the pedicels are frequently straight
in shade-grown plants.
F. flabellata is very finely represented in the exsiecata “ Reverchon,
Pl. d'Algérie, 1896 (Kabylie), No. 103," as F. capreolata.
F. alexandrina, described by Gasparrini, seems to be a rampant form of
this species, as identified by Parlatore. Gasparrini, whose type of F. flabéllata
appears co be a dwarf form from a native habitat, did not recognize that the
two plants were conspecific.
F. ambigua, Lojacono Pojero, FL. Sicula, 62(1888), is reduced by Nicotra in
Le Fum. Ital. p. 60, to a variety 8. ambigua of this species, differing. from
the type by its rosy flowers and larger fruits, Authentic material of this
plant has not been examined,
Examples of F. flabellata have been noted from South Italy, Sicily, and
Algeria, but none from Spain. The specimens met with under this name
from Istria and Dalmatia have proved referable to F. capreolata, F. Gaillardotii
or F. judaica, but a Dalmatian example (Botteri, Cazziol !) exists in Herb,
Mus. Brit. under the name of F. speciosa.
An unnamed, fragmentary plant in Herb. Kew, collected by Miss R.A.
Bainbridge in South Morocco in 1907, resembles 7. flabellata in its large,
white, dark-tipped flowers, but its pedicels are straight and such fruits as it
shows nearly smooth. ‘This may prove to be an undescribed species.
bho
GENERA FUMARIA AND RUPICAPNOS. 67
9. Fumaria supara, Boissier, Diag. Pl. Or. ii. No. 8, p. 15 (1849) ; Haussk.
in Flora, lvi. 551 (1873). F. alewandrina, Ehrbg. ex Hamm. Mon. 43
(1857).
Icon. Hamm. l.c. tab. 5, ut F. alexandrina.
Egsice. Bornmüller, Iter Syriacum, 1897, No. 46, Jaffa! Letourneux,
Pl. Aigyptiacee, No. 231, Mariout !
B. insignis, var. nov.
Ezxsice. K. Untchj, Kaiserwald, Pola, 1904, ut £, Jlabellata! Pichler,
Dalmatia, in agris circa Spalato, 1870, ut F. agraria (partim) !
Racemis paucifloris pedunculos subæquantibus, /foribus multo majoribus
(12-13 mm. longis) petalo superiore latius alato subacuto et breviter rostellato,
fructibus subrotundis apice abrupte trurcatis obsolete foveolatis sine maculis
nigris a typo differt.
Hæc pulchra varietas habitat in Istria, prope Polam, et in Dalmatia, circa
Spalato, ubi ex Egypto aut Palestiná forsan introducta est.
F. flabellata longe distat pedunculis longioribus, pedicellis fructiferis
areuatis recurvis nec erecto-patentibus, sepalis magnis ovatis, et petali
superioris alis atropurpureis nee albidis.
In the case of F. judaica, Hammar's description, though taken from the
living plant, seems unsatisfactory respecting the calyx and the fruit. All the
specimens examined show very small, oblong-lanceolate sepals, as described
by Boissier and Haussknecht, and while the fruit varies in the development
of its keel, no mucronulus has been observed in any dried example that
has reached maturity. A feature of the fruit hitherto unnoticed is the
conspicuous tubercle at the base of the inner apical pit. Good flowers are
very frequently lacking in herbarium material of this species (especially of
the Egyptian plant), and it is probable that the Syrian and Egyptian forms
are not exactly identical, the fruiting pedicels in the latter (the form
described by Hammar) being distinctly shorter. It may also prove that
F. judaica occurs in Greece and has been confounded with the following
species.
An example in Herb. Mus. Brit. from Central Africa (Oudney, Denham &
Clapperton, 1822-4, as F. officinalis !) is seemingly referable to F. judaica.
hd ka
FUMARIA AMARYSIA, Boiss. & Heldr. in Fl. Orient. i. 138 (1867) ; Haussk.
in Flora, lvi. 550 (1873).
Musiec. Heldreich, Herb. Greece. Norm. No. 817!
Fumaria annua, parce ramosa, lawa, diffusa vel petiolis cirrhosis scandens.
Folia irregulariter SOM MOM foliolis in lobos oblongos mucronatos vel
lanceolatos acutos fissis, infimis breviter petiolatis. Racemi la. viusculi, sæpius
10-15-flori, pedunculos subæquantes. Bractew lineari-lanceolatæ, acuminate,
pedicellis mediocribus apice aliquanto incrassatis fructiferis subereetis paulo
breviores. Sepala 15-2 mm. longa, 1 mm. lata, ovato-lanceolata, acuta, vix
peltata, plus minusve dentata, præter nervum dorsalem rubicundum rosea,
Z 2
268 MR. H. W. PUGSLEY: A REVISION OF THE
corolla tubo multo angustiora. Corolla 9-10 mm. longa, pallide rosea ; petalo
superiore alis roseis reflexis carinam superantibus apicem. attingentibus et
calcar versus longe productis obtuso; petalo inferiore marginibus latis patulis
roseis apicem vix attingentibus subacuto, sæpe libero deflexo; petalis
interioribus sursum curvatis, breviter et obscure apice atropurpureis. Fructús
majusculi, 2'5 mm. longi et paululum angustiores, breviter subrotundo-obovati,
superne subtruncati et inferne angustati, arinå obscura paululum compressi,
in sicco apicis foveolis obscuris tubereulato-rugosi.
Hee species proxima est F. judaicw, cujus forsitan mera varietas sit, sed
F. judaica differt habitu robustiore, pedicellis vulgo longioribus apice magis
incrassatis, sepalis oblongo-lanceolatis acuminatis basi solum dentatis, corolla
majore albidà nec rosea, fructibus majoribus nitidis evidentius tuberculato-
rugosis.
F. macrocarpa, Parl. cujus corolla ei 7. amarysie subsimilis est fructibus
maximis globosis longe recedit.
F. Bastardii, Bor. subsectionis Muralium, pro quà cl. Boissier primo
F. amarysiam habuit, petali inferioris marginibus angustis presertim. sed
etiam bracteis brevioribus, sepalis majoribus serratis, fructibus subacutis vel
obtusis nee vero subtruncatis plane differt.
Heectenus F. amarysia in Greecia sola inventa est.
11. FUMARIA macrocarpa, Parlatore, Pl. Novee, 5 (1842), et Mon. Fum.
143 (1844) : Hamm. Mon. 45 (1857); Haussk. in Flora, lvi. 559 (1873).
F. megalocarpa, Boiss. & Sprunn, Diag. PI. Or. i. No. 1, p. 68 (1843).
Icon, Hamm. l. c. tab. 6.
Ewsice. Orphanides, Fl. Grieca. Exsicc. No. 20, Athens, ut F. megalocarpa !
Heldreich, Herb. Greece. Norm. No. 1002, ut F. megalocarpa ! Balansa, PI.
d'Orient, 1854, No. 128, Smyrna!
This species, recognizable at a glance by its very large, globular fruits, is
remarkable also for its extremely narrow sepals and the complete absence of
purple colouring from its corolla. In well-grown examples the pale, rosy
flowers attain a length of 11 mm., with the linear-lanceolate sepals nearly
3 mm. long. The distant and curiously recurved lower petal depicted in
Hammar's figure was noticed by Boissier as characteristic of this species, and
its spreading margins are more spathulate in outline than in any other
member of the Grandiflore.
F. owyloba, Boiss. Diag. Pl. Or. ii. No. 8, p. 14 (1849), appears to be a
shade-form rather than a true variety of F. macrocarpa.
Supsecrio Il. CAPREOLATAE.
A
Capreolate, Haussk. in Flora, lvi. 539 (1873); Pugsley, Fum. in Brit. 5
(1912) ; Hamm. Mon, 24 (1857), ut sectio, ex parte.
Pedunculi longi pedicellis swpissime apice mulio incrassatis przediti.
Bracteæ raro quam pedicelli fructiferi multo breviores, Sepala plerumque
magna, apicem versus integriuscula, raro quam coroll tertia pars breviora
GENERA FUMARIA AND RUPICAPNOS. 269
ejusque tubum latitudine ssepissime superantia. Corolla seepius angusta ;
petalo superiore alis apicem haud attingentibus acuto; petalo inferiore acuto
marginibus erectis angustis apicem haud attingentibus. Fructüs parvi,
mediocres vel magni, in vivo plerumque stipite carnoso distincto quam
pedicelli apex inerassatus angustiore priediti, siccitate læves usque ad rugosi.
The wings of the upper petal, as well as the tips of the inner ones, are dark
purple in F. capreolata, F. purpurea, F. macrosepala, and F. berberica. In
F. dubia and F. coccinea this dark colour is obscure or wanting, and in the
last-named the wings themselves are obsolescent.
* Series Eu-Capreolatæ.
Pedicelli fructiferi areuato-recurvi aut saltem patentes ; fructüs leves vel
rugulosi.
12. FUMARIA CAPREOLATA, Linn. Spec. Plant. 701 (1753) ; Hamm. Mon. 24
(1857); Pugsley, Fum. in Brit. 6 (1912). F. pallidiflora, Jordan in
Schultz Archives, 305 (1854). (PI. 9, fig. 2.)
In Hammar’s Monograph Ray's F. major scandens flore pallidiore (Hist.
i. 405 (1686)), cited by Linnzus, is omitted from the synonymy of this
species. The omission is not without justification, for Ray’s remarks betray
no acquaintance with 7. capreolata, and none of the exsiccata in the Sloane
Herbarium under his name belongs to this species, being all, except one
doubtful fragment (probably F. muralis subsp. Bore), rampant forms of
F. officinalis, as is also a specimen similarly named in the contemporary Herb.
Dubois at Oxford. It may thus be reasonably concluded that rampant forms
of F. officinalis were the recognized P. major scandens Jlore pallidiore of the
British botanists of that period. The Sloane Herbarium contains a good
example of F. capreolata (ILS. 58/146) under Bauhin’s name (Pinax, 143),
and there is a British specimen of var. Babingtonii in Herb. Dubois, labelled
“ F, scandens altera Czesalp.”
Icones. Savi, Mat. Med. tab. i, ic. 1; Sturm, Deutschiands Flora, i. 62. 13 :
Flora Danica, tab. 2359 (forma umbrosa); Hamm. l. c. tab. iii. (forma
sepalis insolite dentatis) ; Journ. Bot. xl. tab. 436, fig. 2, ut F. speciosa.
Exsice. Bourgeau, Fl. Pyr.-Espag. No. 391! Neugebauer, Fl. Exsicc.
Austro-Hungarica, No. 2899!
! Sepala typi vel paulo breviora, subintegra ; corolla rubescens.
B. spectosa, Hamm. Mon. Slügt. Fum. 25 (1854), et Mon. 25 (1857).
F. speciosa, Jord. in Cat. Grenoble, 15 (1849), et Linnza, xxiii. 507
(1850).
Exsice. Mabille, Herb. Corsicum, 1868, No. 337, ut F. speciosa !
Hæc varietas racemis laxiusculis, pedicellorum tenuiorum bracteis paulo
brevioribus, sepalis vulgo 4-5 mm. longis 2-3 mm. latis ovatis potiusquam
ovalibus frequentius subintegris, corollis primo albidis seepius tandem omnino
rubris petalo superiore parum angustato preeditis a typo differt.
210 MR. H. W: PUGSLEY : A REVISION OF THE
In addition to the relatively shorter and broader sepals of this variety and
its more highly coloured corolla, which often changes wholly from white
to deep crimson, the upper petal, when fresh, is distinetly less laterally
compressed than in typical F. capreolata, and the pedicels seem slenderer and
more flexuous, with somewhat. shorter bracts.
According to Hammar the racemes are denser than in the type, but the
examples examined show laxer racemes as noticed by Jordan (“racemis . . .
laxiusculis ").
In the living state this variety, whieh is found chiefly in Spain and
Southern France, is especially beautiful owing to the bright red and white
colouring of its flowers.
!! Sepala quam typi sepe longiora ; corolla albida vel roseo-tincta.
y. albiflora, Hamm. Mon. Slügt. Fum. 25 (1854), et Mon. 25 (1857).
F. capreolata B. flavescens, Nicotra, Le Fum. Ital. 44 (1897), excl.
syn. partim.
Husice. Heldreich, Herb. Greece. Norm. No. 1003, ut F. capreolata !
Many North European specimens that have been referred to this variety
are probably shade-forms only (forma parviflora, Haussk.) of the type, but a
few of them and some white-flowered examples from the Mediterranean
region appear to constitute a distinet variety.
6. Babingtonii, Pugsley, l.e. 9. F. pallidiflora, Babington in Journ. Linn.
Soe. iv. 157 (1860), non Jordan.
Icon. Journ, Bot. xl. tab. 436, fig. 1, ut F. capreolata.
e. devoniensis, Pugsley, l c. 10.
111 Sepala minora, magis dentata ; corolla ssepius rubescens.
E. condensata, Ball, Spic. Fl. Maroce. in Journ. Linn. Soc. xvi. 315 (1878).
tæsice. Hooker, Inzuda, Greater Atlas, S. Marocco, 1871, in Hb. Kew!
Racemi densi. Bracteæ pedicellis parum reeurvatis paulo breviores,
Sepala 4 mm. longa, 2°5-3 mm. lata, ovata, acuta, basin versus conspicue
ineiso-denta^a. Corolla 10-11 mm. longa, ut in var. speciosc. rubescens.
Fructis quam in typo majores, 2:5 mm. longi et plns 2 mm. lati.
This variety shows the dense racemes characteristic of the specific type
combined with relatively small and much eut sepals and large fruits similar
to those of var. Babingtonii.
n. Hochreutineri, var. nov.
Peere 2 d d hr inar Tavar 4 ros
Evsice. B. P. G. Hochreutiner, Voyage botanique en Algérie, 1901,
T y : 4 i
No. 18, Environs d'Alger, ut F. capreolata var. intermedia, Haussk. !
[ *olaiiv parr hi ee "c "
Sepala relatis e parva, 8—35 mm. longa, vix 2 mm. lata, oblongo-ovata,
breviter aeuminata, circumcirca parce serrato-dentata, albida ; corolla
10-11 mm. longa, ut in var. speciosd rubescens.
GENERA FUMARIA AND RUPICAPNOS. EL
This rare plant, remarkable for its relatively small, toothed sepals, bears a
general resemblance in other respects to var. speciosa, and in comparison may
be regarded as somewhat analogous with the var. brevisepala of F. purpurea.
It agrees in floral characters with Haussknecht's description of his var.
intermedia, but the plant in the Boissier Herbarium on which this name
was founded cannot be retained as a variety of F. capreolata, which
Dr. Hochreutiner’s plant evidently is, and a fresh name for the latter has
therefore been substituted.
F. platycalyx, Pomel, Nouv. Mat. Fl. Atlant. 239 (1874), is a plant of
which no material has been seen, but it is apparently a shade-form of
F. capreolata, as treated by Battandier & Trabut (Fl. de l'Algérie, i. 26
(1888)).
Of F. nemorosa, Lojacono Pojero in Fl. Sicula, 61 (1888), which is reduced
by Nicotra to a synonym of F. capreolata var. albiflora, several authentic
specimens have been seen which either belong to this variety or are shade-
forms of the specific type. The diagnosis of F. nemorosa, however, states
that the fruits are “ grosse rugulosis, —a feature recalling F. flabellata rather
than any form of F. capreolata—and so the name may be held doubtful.
The var. atrosanguinea, Brochon & Neyraut, adopted by Rouy & Foucaud
(Fl. Fr. i. 171), seems to be a form only of var. speciosa, Hammar, in which
the sepals as well as the corolla become red. It was originally issued as
F. speciosa, Jord., forma atrosanguinea, in the exsiceata “C. Magnier, Fl. Sel.
Exsice. No. 2388. Pyrénées-Orientales, 1890."
Tt may be observed that while this variety atrosanguinea seems almost
inseparable from var. speciosa, Hamm., the variety described by Rouy &
Foucaud under this latter name appears to be Hammar's type, which is not
otherwise accounted for by the collaborateurs.
The variety provincialis, Rouy & Foucaud (l. c.), seems from the description
to be a luxuriant state of F. capreolata rather than a true variety—a view
borne out by the example in Herb. C. Bailey of ** Magnier, Fl. Sel. Exsiec.
No. 456,” which is cited by the authors in illustration.
Another specimen in the same collection under this name, gathered
by Albert in 1893, at Sollies-Toucas, Var, whence Rouy & Foucaud record
var. provincialis, is, however, much more distinct and perhaps a good variety.
This has bracts barely exceeding half the length of the pedicels, and
relatively lax and slender racemes of large, pale flowers attaining 14 mm. in
length, with the upper petal very narrow and comparatively but very little
laterally compressed, much as in var. speciosa.
A further conspieuously fine plant allied to this form, but with large fruit,
is one from Corfu (Baenitz, Fl. Corcyrensis ; Kastrades, 1896, as
F. capreolata Y), which may likewise be varietally separable.
272 MR. H. W. PUGSLEY: À REVISION OF THE
Tt is not always easy, in the absence of adequate material and a knowledge
of the habitats, to justly estimate the value of the differences presented by
this widely distributed and variable species.
With regard to the geographical distribution shown at pp. 10 and 11 of
* Fumaria in Britain,” the correctness of the habitats, Canaries and
St. Helena, given both by Hammar and Haussknecht, may be doubted, the
plants intended being probably F. coccinea and F. muralis respectively.
13. FUMARIA DUBIA, sp. nov.
F. capreolata B. intermedia, Haussk. in Flora, lvi. 541 (1878).
Evxsice. Montagnes voisines du Fort L'Empereur, Alger, 1831, Herb.
Fauché, No. 1 (in Hb. Boissier) !
Fumaria F. capreolate habitu foliisque et pedunculis longis racemos densos
superantibus. Bractew pedicellos Jructiferos arcuato-recurvos subequantes.
Sepala circa 3*5 mm. longa et 2 mm. lata, ovata, peltata, acuminata, denticulata,
albida. Corolla 10-12 mm. longa, omnino rosea, ut videtur, aut roseo-albida ;
petalo superiore angusto acuto alis roseis carinam haud superantibus preedito ;
petalo inferiore marginibus angustissimis erectis acuto. — Fruetüs parri,
2:25 mm. longi et 2 mm. lati, subrotundi, ad medium latissimi, superne in
apicem subacutum et inferne in stipitem quam pedicelli apex angustiorem
æqualiter angustati, juniores plane apiculati, siccitate apicis foveolis parvis
distinctis rugulosi.
Descriptio secundum exemplar solum imperfectum in Herb. Boissier.
In montibus Algeriæ habitat.
Hae Fumaria que ut species nova dubitanter descripta est, per sepala
minora denticulata, per petali superioris alas roseas nec purpureas, et
præsertim per fructůs subacutos rugulosos a F. capreolatd, ut videtur, specifice
differt.
Secundum cl. Haussknecht hujus plantæ fructus rugulosus transitum ad
F. flabellatam denotat, sed, me judice, et fructus rugulosus subacutus et
corolla angusta sine apice purpureo rosea affinitatem cum F., coccined
monstrant.
The specific name intermedia having been already appropriated under the
genus Fumaria (Withering, Arr. ed. 3, iii. 620, t. 29 (1796)) to represent a
plant now referred to Corydalis bulbosa, DC., the present form has been
described as F. dubia.
14. FUMARIA PURPUREA, Pugsley in Journ. Bot. xl. 135 & 179 (1902) ;
Fum. in Brit, 12 (1912). F. Boræi, Babington in Journ. Linn. Soc. iv.
157 (1860), non Jordan.
Icon. Journ. Bot. xl. tab. 436, fig. 3.
B. brevisepala, Pugsley, Fum. in Brit. 13.
leon, Pugsley, l. e. fig. 1 (Journ. Bot. l. tab. 519).
eo
GENERA FUMARIA AND RUPICAPNOS. 273
There is a British specimen of F. purpurea in the Herbarium of Dillenius
at Oxford, and another still earlier (cirea 1700) in the Herb. Dubois. The
latter is labelled * F. major scandens floribus albis riche saturate purpureo
(sic). Omnibus partibus major est F. scandente nostrate. An F. altera
Cæsalp.? ”
** Series Macrosepalæ.
Pedicelli fructiferi suberecti ; fructûs rugosi.
15. FUMARIA MACROSEPALA, Boissier, Elenchus No. 7, 8 (1838); Voyage
Bot. Espagne, ii. 19 (1839-1845); Hamm. Mon. 36 (1857); .Willkomm
& Lange, Fl. Hisp. iii. 880 (1880).
Icones. Boissier, Voy. Bot. Espagne, tab. iv.; Hamm. l.c. tab. iii.
Exsicc. Willkomm, Iter Hispanicum, 1845, No. 863, Malaga! Lange,
Pl. Hisp. 1851-2, Jaen, in Hb. Kew! Huter, Porta & Rigo, Iter Hisp.
1879, No. 518, ut F.latisepala ! .
In this species, which is said to inhabit rocky slopes rather than cultivated
ground, the corolla resembles that of F. capreolata in its narrowly winged
upper petal and its peculiar reddish dorsal suffusion.
F. malaeitana, Haussk. & Fritze in Flora, lvi. 548 (1873), although described
as closely allied to F. gaditana, Haussk., seems rather, judging by the type
specimen in Herb. Boissier (Ad sepes prope Malacitanam, R. Fritze, Mai,
1873 !), as well as by the diagnostie characters, a form of F. macrosepala, as
suggested by Willkomm & Lange (lv. 881). Fritze's specimen is a piece of
a large, vigorous plant, gathered late and perhaps on cultivated ground, in
-which the mucronulus of the fruit is obscured.
A plant sent out by Porta & Rigo as F. macrosepala, Boiss. var. malacitana,
Haussk. & Fritze, (Iter IV Hispanicum, 1895, No. 24!) is luxuriant
F. macrosepala, with the sepals and corolla ranging to 7 mm. and 13 mm. in
length respectively.
This fine species is reported also for the mountains of Tetuan, in Morocco,
by J. Ball in Journ. Linn. Soc. xvi. 315 (1878). Dall's specimen is at Kew.
16. FUMARIA BERBERICA, sp. nov. (PI. 11, fig. 1.)
F. agraria var. atlantica, Ball, Spic. Fl. Marocc. in Journ. Linn. Soc. xvi.
316 (1878), non Cosson.
Evsice. J. Ball, Iter Maroccanum, Ait Mesan, 1871, in Hb. Kew, ut
F. agraria var. atlantica !
Fumaria habitu robusto sine dubio annua. Folia 2—3-pinnatisecta, foliolis
in lobos oblongos acutos mucronatosve eis 7. agraric similes fissis. — /'acemi
10-15-flori, pedunculos sulwquantes. Bractew magne, lineari-lanceolate,
acuminate, subfoliate, pedicellos fructiferos suberectos breviusculos apice
multo incrassatos conspicue superantes. Sepala 4°5-5°5 mm. longa, 2°5-3 mm.
lata, ovata, peltata, acuminata, circa basin marginemque inferiorem parce
oe
i
274 MR. H. W. PUGSLEY : A REVISION OF THE
serrato-dentata, preter nervum dorsalem latum viridem albida, corolla tubo
latiora. Corolla 11-12 mm. longa, voseo-albida ; petalo superiore alis
atropurpureis reflexis. carinam æquantibus apicemque fere attingentibus
subacuto; petalo inferiore acuto marginibus erectis. angustissimis ; petalis
interioribus apice atropurpureis, Fructés majuseuli, circa 2*5 mm. longi
ac lati, subrotundi, cum mucronulo persistente obtusissimi et inferne parum
angustati, paululum compressi sed distincte carinati, siccitate apicis foveolis
parvis obscuris dense sed haud grosse rugosi.
Heee rara species, ad gregem Capreolatarum plane referenda, F. macro-
sepale proxima. est, sed sepalis dentatis, corollis minus angustatis apicem
versus haud rubescentibus nec in siccitate viridescentibus, fructibus minoribus
sed distinctius rugosis differt.
Quanquam el. Ball hane. plantam ad F. agrarie varietatem retulit, id ab
illo dubitanter faetum esse manifestum est, nam * Planta haud omnino certa—
facies F: capreolate, sed characteres potius F. agraric scripsit. Ut etiam
F. macrosepalam certe cognovit ejus similitudinem cum /. berbericd non
vidisse paulo notabile est, et per hujus corollam in vivo latiorem forsitan
explicandum sit.
F. capreolata per flores angustiores, pedicellos areuato-recurvos fruetüsque
leves minores, F. agraria per sepala parva, eorollam omnino diversam
fructüsque rostratos grosse rugosos, et F. flabellata per corolle formam
pedicellosque arcuato-recurvos plane distant.
F. berberica in rupibus arenaceis Atlantis majoris Mauritanie australis, in
convalle Ait Mesan (alt. 1600 m.) habitat.
A second specimen in Herb. Kew (F. agraria, var. —, Hooker, Revaia,
6-7000 ft., S. Marocco, Greater Atlas, May, 1871!) may also belong to
this species, but it shows no fruit and is too fragmentary for certain
determination.
17. Fumarta coccinea, Lowe MSS. in Herb. Kew and Herb. Mus. Brit.,
sp.nov. (PI. 12.)
F. montana, Haussk. in Flora, lvi. 546 (1873), et auct., non Schmidt.
Tæsice. Bourgeau, Pl. Canar. No. 178, ut F. capreolata! Dela Perraudiere,
Canaries, 1855, in Hb. Boissier, ut XF. capreolata var. canariensis! Lowe,
Santa Cruz, Teneriffe, 1859, in Hb. Mus. Brit.! | Bornmüller, Pl. Exsice.
Canar. 1901, No. 2029! et No. 2028 (forma umbrosa), ut F. montana!
Sprague & Hutchinson, Canary Is., No. 1265!
Fumaria annua, internodis longis gracilis, plus minusve ramosa, petiolis
cirrhosis scandens. Folia sepe parvula, foliolis in lobos oblongos, subellipticos
vel late cuneatos, seepissime obtusos et nonnunquam mucronatos fissis, infimis
breviter petiolatis, irregulariter 2-(rarius 3-) pinnatisecta. Racemi pauciflori
(6-12-flori), pedunculis gracilibus paulo breviores. — Bractew lineari-subulatie,
acuminate, pedicellis fructiferis erecto-patentibus, rectis flexuosisve, apice
incrassatis, sepe breviusculis, paulo breviores. Sepala 3-4 mm. longa, circa
GENERA FUMARIA AND RUPICAPNOs. 2750
2 mm. lata, oblongo-ovata vel oblongo-lanceolata, peltata, acuta, parce dentata
vel etiam praecipue basin versus paululum laciniata, nervo dorsali obscuro
viridiuseulo albida vel roseo-albida, corolle tubum subzquantia, longe
persistentia. Corolla 8-10 mm. longa, rubra vel coccinea (in formis umbrosis
albida), in fructu juniore persistens ; petalo superiore alis rubris angustissimis
(carinam haud :equantibus) vel omnino obsoletis acuto angustissimo; petalo
inferiore marginibus angustis erectis apicem haud attingentibus acuto; petalis
interioribus prope rectis apice parum saturatioribus. Fructis parvi vel
mediocres, 2-2:5 mm. longi ac lati, subrotundo-quadrati, obtusissimi obtusi vel
rarius acutiusculi, inferne abrupte angustati, satis carinato-compressi, siccitate
apicis foveolis parvis plus minusve distinctis rugosi.
F. coceinea habitu gracili, floribus minoribus rubris angustissimis petalo
superiore fere exalato preeditis ab hujus seriei aliis speciebus differt.
F. montana, Schmidt in Beit. Fl. Cap. Verd. Inseln, p. 264, quacum
F. coccinew forma umbrosa a cl. Haussknecht auctoribusque recentioribus
confusa est, habitu robustiore, foliolis longius petiolatis cum lobis
angustioribus, bracteis longissimis, floribus minoribus semper albidis, petali
inferioris subspathulati marginibus patentibus fructuque majore subgloboso
plane earinato distinguitur et ad subsectionem Latisepalarum referenda est.
F. coccinea habitat in insulis Canariensibus ubi in montibus sylvaticis
invenitur.
This very distinct fumitory, which has been distributed at intervals from
the Canaries under different names, can only be placed among the Capreolate,
and is remarkable for its very narrow and nearly unicolorous corollas.
In the living state, when not shade-grown, the flowers appear to be bright red
in colour, and have been referred to by collectors as crimson, * coccinea,”
and * sanguinea."
The fruits of this species show considerable variation in form, for while
those of Lowe's original specimens are quite rounded-obtuse, other herbarium
material furnishes a transition to the subacute-fruited form lately obtained
by Messrs. Sprague & Hutchinson.
Haussknecht’s identification of this plant with Schmidt’s F. montana
seems attributable to the similarity in size and colour of the flowers when
F. coccinea is shade-grown, both species lacking the dark corolla-tip that is
venerally prevalent in the genus. There is also some resemblance in the
D
sepals, but in other respects the two plants appear widely different.
Sussrcri0 II. MURALES.
Murales, Haussk. in Fiora, lvi. 513 (1873). Capreolate, Hamm. Mon. 24
(1857), ut sectio, ex parte ; Medie, Pugsley, Fum. in Brit. 15 (1912).
Pedunculi plus minusve breviusculi, eis F. bicoloris et F. sepium exceptis.
Pedicelli quam in subsectionibus prioribus sepius minus incrassati, nisi in
F. Thuretii. Bracteæ longitudine variabiles sed pedicellos fructiferos raro
276 MR. H. W. PUGSLEY: A REVISION OF THE
sequantes. Flores racemorum recentiorum quam in prioribus sæpe pauciores
minusque explicati. Sepala dentata, serrata vel integriuseula, raro corollie
tertià parte longiora vel ejus tubo plane latiora. Petala exteriora obtusa vel
acuta, inferius marginibus angustis qui apicem vix attingentes plerumque
erecti rarius patentes sunt. Fructis parvi aut modici, forme diversissime,
siccitate tuberculato-rugosi, rugulosi vel etiam læves.
In this subsection the wings of the upper petal, as well as the tip of the
inner ones, are dark purple except in F. bicolor and some forms of F. Bastardi.
The term Medic, which was applied to this difficult group in “ Fumaria in
Britain," has been abandoned in favour of Murales, Haussk. as further
investigation sufficiently shows that F. media, Loiseleur,* Notice,’ p. 101 (1810),
a plant whose identity has given rise to much discussion, represents only a
rampant form of F. officinalis, Linn. and has no connection with the species
of this subsection. — Loiseleur refers to its retuse fruits in his diagnosis and
again in his subsequent remarks, and the figure which he quotes (Vaillant,
Bot. Par. tab. x. fig. 4)—an exceptionally good drawing but without fruits—
bears a raceme of dorsally compressed flowers, with small sepals, which, if
not a form of F. officinalis, can only be referred to F. major, Bad.
De Candolle, Reichenbach, and Boreau clearly regard F. media, Lois. as
closely allied to X. oficinalis, and Parlatore (Mon. Fum. p. 59) shows at
some length that they are not separable as species, a view more recently
maintained in Rouy & Foucaud’s ‘Flore de France’ and Nicotra's * Le
Fumariacee Italiane, The confusion that has arisen through the connection
of this name with F. muralis and its allies seems largely due to Hammar’s
interpretation of F. media (Mon. Fum. p. 28), which, combined with his
association of F. Bastardii with F. muralis, forms the most serious error in
his generally excellent work, and is carefully corrected by Haussknecht.
The further extension of Loiseleur’s name by Willkomm & Lange
(Fl. Hisp. iii. p. 882) to cover the majority of the varied Spanish forms of
this group argues a treatment of this subsection quite unequal to that of the
remainder of the genus.
*'Series Sub-Agrariz.
Sepala parva ; petali inferioris margines angusti, patentes ; fructüs rugosi.
18. FUMARIA BICOLOR, Sommier ex Nicotra in Le Fum. Ital. 55 (1897) ;
Fl. Giglio, 7 (1900). F. capreolata var. —, Duthie in Journ. Bot. x.
208 (1812) ; F. rupestris var. maritima, Battandier in Bull. Soc. Bot.
Fr. xxxii. 330. (1885); F. Loiseleurit. Clavaud £. leronensis, Burnat,
Fl. Alp. Marit. 69 (1892).
Evsice. Burnat, St. Marguerite, Cannes, in Hb. Boissier & Hb. Zurich,
ut F. Loiseleurii B.leronensis! Townsend, St. Marguerite, 1872, in Hb. Kew,
ut F. maculata! Duthie, Fl. Melit. Exsiec. 1874, in Hb. Kew & Hb.
Mus. Brit., ut F. capreolata, var. — !
Ho. x4 E
GENERA FUMARIA AND RUPICAPNOS. 211
Fumaria annua, habitu gracili, diffusa vel petiolis cirrhosis scandens.
Folia glaucescentia aut viridia, foliolis in lobos lanceolatos acutos vel
oblongos mucronatos fissis irregulariter 2-3-pinnatisecta. Racemi pauciflori
(8-12-flori), pedunculis tenuibus spe incurvatis breviores. Bractew oblonge,
cuspidate, pedicellis fructiferis gracilibus flexuosis interdum suberectis interdum
patentibus triplo quadruplove breviores. Sepala minima, 2-2:25 mm. longa,
circa 1 mm. lata, ovalia, vix peltata, breviter acuminata, plus minusve dentata,
nervo dorsali rubro rosea, coroll tubo multo angustiora. Corolla 10-13 mm.
longa, fere recta, albida vel roseo-albida, tandem preter petalorum interiorum
apicem omnino rosea, ut in racemo singulo flores juniores albidi maturiores
rosei videantur; petalo superiore subacuto alis roseis reflexis apicem vix
attingentibus carinam eequantibus vel paulo excedentibus calcar breviuseulum
versus longe productis ut in F. Bastardii ; petalo inferiore marginibus angustis
patentibus roseis apicem haud attingentibus acuto; petalis interioribus apice
atropurpureis. Fructús parvi, 2-2:25 mm. longi et DL:5-1:75 mm. lati,
subrotundo-ovati, subacuti sed inferne parum angustati, obscure carinati et
paululum compressi, siccitate dense sed haud grosse rugos apice foreolis
angustis sepius nigris et nonnunquam maeulá laterali nigra notati.
Hee species, a cl. Sommier in *Florula del Giglio’ eta cl. Burnat in ‘Flore
des Alpes Maritimes’ bene descripta, in subsectione Agrariarum, ad quam per
corolla formam accedit, a cl. Nicotra locata est. In aliis characteribus, sicut
in habitu gracili et in fructüs formá, subsectionem Muralium plane refert, et,
me judice, F. Bastardii, Bor. (et preecipue varietati vaganti) proxima est.
F. bicolor racemis paucifloris, pedunculis tenuibus longioribus, bracteis
sepalisque minimis, corollá angustiore fere rectá, petali superioris calcare
breviore, petali inferioris marginibus paulo latioribus, fructibus minoribus et
densius rugosis a F. Bastardii distinguitur.
F. rupestris, Boiss. habitu robustiore, bracteis longis setaceo-acuminatis,
sepalis majoribus lanceolatis, fructibus rugosioribus multo majoribus plane
differt.
F. amarysia, Boiss. bracteis multo longioribus, petalis interioribus apice
brevius atropurpureis, fructibus multo majoribus obovatis subtruncatis longe
recedit.
F. bicolor in insulis italicis Giglio, Elbá, Montecristo, Capraià et Giannutri,
atque in insulà Sta. Margarita prope Cannes Galloprovincie dumeta nec
culta habitat; in insulà Melité etiam in loeis saxosis umbrosis atque in
scopulis maritimis Algeriz invenitur. In herbario Kewensi exemplar
imperfectum “F. Petteri Rchb. Sicilia, Parlatore, No. 137” forsitan ad
hane speciem relatum sit.
This rare fumitory, one of the few species not affecting disturbed ground,
is especially interesting to British botanists in that it was discovered in 1872
almost simultaneously at Cannes by the late Mr. F. Townsend, who pointed
218 MR. H. W. PUGSLEY : A REVISION OF THE
it out to M. Burnat, and at Malta by Mr. J. F. Duthie. The identity of
F. bicolor and F. Loiseleurii B. leronensis seems to have been hitherto over-
looked.
19. Fumaria Basrarpi, Boreau in Duchartre Rev. Bot. ii. 359, excl. var. 8
(1846-7); Fl. du Centre, ed. 2, 28 (1849), et ed. 3, 34 (1857) :
Pugsley, Fum. in Brit. 35 (1912). F. confusa, Jordan in Cat. Dijon
(1848), et Linnæa, xxiii. 469 (1850) ; Pugsley in Journ. Bot. xl. 173
(1902); F. media B. confusa, Hamm. Mon. 28 (1857), non F. media,
Loiseleur ; F. Gussonei, Boiss. ex Haussk. in Flora, lvi. 513 (excl.
F. Petteri, Rchb. in syn.), in sensu lato (1873).
leones. Hamm. l.c. tab. 3, ut F. media B. confusa; Journ. Bot. xl.
tab. 436, fig. 6, ut F. confusa.
Eesice, Billot, Fl. G. & G., No. 1206 ! et No. 3307 bis! Schultz, Herb.
Norm. No. 605, ut F. confusa !
Fumaria annua, habitu satis robusto, sepe ramosissima, suberecta vel diffusa
vel rarius petiolis cirrhosis scandens, Folia irregulariter 2-3-pinnatisecta,
viridia vel glaucescentia, foliolis in lobos oblongos acutos mucronatosve quam
F. muralis angustiores fissis praedita. Racemi laviuseuli, multiflori (vulgo
15-25-flori), pedunculos superantes. — Dractee lineari-oblong:e, cuspidate,
pedicellis fructiferis longiusculis suberectis vel erecto-patentibus duplo vel
etiam triplo breviores. Sepala circa 3 mm. longa, V9 mm. lata, ovalia, vix
peltata, acuta, plus minusve circumcirca serrata, preter nervum. dorsalem
viridiusculum rosea, corollie tubo angustiora, inferne remota, sepe in fructu
juniore persistentia. Corolla 10-11 (raro 12) mm. longa, preter. petalorum
interiorum. apicem atropurpureum rosea ; petalo superiore obtuso vel acuto,
a latere compresso (dorso satis angusto), etiam in gemmis haud spathulato,
alis roseis reflexis calcar versus longe productis siepe carinam excedentibus
apicemque attingentibus prodito; calcar sepalis longius: petalo inferiore
marginibus angustis patulis apicem vix attingentibus subaeuto. Fructis
modici, circa 25 mm. longi ae lati, rotundati, obtusiusculi vel rarius obtusis-
simi, paulo compressi sed obscure carinati, in vivo inferne in stipitem
obscurum qui pedicelli apicem parum incrassatum æquat vel excedit sæpissime
vi! angustati, siccitate apicis foveolis latis paululum obscuris satis rugosi.
! Petalum superius omnino roseum.
B. patens, comb. nov.; F. Gussonei, var. patens, Haussk. l.e. 518.
Easice. Mabille, Herb. Corsicum, 1868, No. 338, ut F. Gussonei !
Planta laxa sed robusta, foliorum lobis quam in typo paulum latioribus.
Bractew acuminata, intimæ pedicellos fructiferos breviusculos flewuosos scope
patulos sub:equantes, relique. dimidio breviores. Corolla 9-10 mm. longa.
Fructüs circa 2 mm. longi ac lati, ut in var. Gussonei subglobosi et leviter
rugosi. Aliter ut in ty po.
ie
GENERA FUMARIA AND RUPICAPNOS, 21
y. vagans, var. nov.
F. vagans, Jord. in Cat. Grenoble (1849), et Linnea, xxiii. 508 (1850);
Hamm. l.c. 46.
Icon. Hamm. l.c. tab. 6, ut F. vagans.
Bracteæ oblongze, cuspidate, apice deuticulate, pedicellis fructiferis triplo
breviores. Sepala 2-25 mm. longa, 1°5-2 mm. lata, rotundato-ovalia,
breviter acuta, nervo dorsali inerassato carinata. Petalum superius vix a
latere compressum, alis roseis patenti-reflexis carinam paulo superantibus
apicemque vix attingentibus subaeutum. Fructús circa 2:25 mm. longi, 2 mm.
lati, subrotundo-ovati, acutiusculi, inferne in stipitem obscurum quam pedicelli
apex paulo angustiorem angustati, tenuiter rugost, Aliter ut in typo.
This plant, though near the specific type, is separable by its more carinate
sepals, by the form of its corolla, which when fresh is broader and less
laterally compressed, and especially by its smaller, more acute and more
finely rugose fruits. Rouy & Foucaud’s variety occidentalis of F. vagans (FI.
France, i. p. 174) seems from the description to be a shade-form only.
!! Petalum superius apice atropurpureum.
8. Gussonei, Pugsley, Fum. in Brit. 40 (1912). F. Gussonei, Boiss.
Diag. Pl. Or. ii. No. 8, p. 13 (1849); Hamm. l.c. 34. (PI. 9, fig. 3.)
Exsice. Billot, Fl. G. & G., No. 1109, ut F. Gussonei! Fiori &c., Fl. Exsice.
Ital. No. 827, ut F. serotina var. Gussonei! — Lojacono, Pl. Ital. Select.
No. 74, ut F. Gussonei !
Varietas scepius satis nana foliis plus minusve glaucis.
longa, 1-2 mm. lata, quam typi paulo latiora, Corolla 9-10 (raro 11) mm.
longa; petalo superiore alis atropurpureis sepe obtuso; petalo inferiore
marginibus quam in typo angustioribus. Fructis satis parvi, cirea 2 mm.
longi et lati, subglobosi, rotundato-obtusi et inferne plane angustati, siccitate
quam in typo apicis foveolis obseurioribus tenwius rugos?.
Sepala 2-3 mm.
e. hibernica, Pugsley, Fum. in Brit. 41 (1912).
Icones. Eng. Bot. Suppl. 2976, ut F. confusa. Pugsley, l.c. fig. 5 (Journ.
Bot. l. tab. 519).
The E.B.S. figure was drawn from a Scilly specimen collected by Townsend.
Although not very characteristic, the acuminate purple tip of the upper petal
is sufficiently indicated.
The Spanish plant represented in the exsiccata “ Sennen, P1. d'Espagne,
No. 228,” as F. vagans, closely resembles this variety except for its smaller,
acuter fruits.
é. affinis, var. nov.
F. affinis, Hamm. Mon. Slügt. Fum. 47 (1854), et Mon. 39 (1857).
Icon. Hamm. Mon. tab. 5, ut F. affinis.
Crsiccs Lange, Almeria, 1852, in Hb. Boiss. !
Hammar's description and figure of this little-known plant recall the small-
flowered form of F. major, Bad., but Lange's specimen from Almeria in
280 MR. H. W. PUGSLEY: A REVISION OF THE
Herb. Boissier clearly belongs to F. Bastardii, as determined by Haussknecht.
In the colour of the corolla it resembles var. Gussonei, but the flowers are
larger with more broadly winged outer petals; and the fruits are relatively
large and very obtuse, approaching those of X. major in shape but showing
no mucronulus in the dry state.
The specimen of F. affinis in Herb. Kew, received from Lange, does not
agree with that in Herb. Boissier and with Hammar's diagnosis, but seems to
be a large-flowered example of typical F. Bastardii.
q. benedicta, comb. nov.
F. Gussonei var. benedicta, Nicotra in Nuov. Giorn. Bot. Ital. nov. ser. iv.
312 (1897); Le Fum. Ital. 53 (1897).
Icon. Nuov. Giorn. Bot. Ital. l.e. tab. 10.
This variety is remarkable for the black apical pits and lateral markings
of its fruit, resembling those of F. bicolor, It is also characterized by its
rampant habit, relatively broad leaf-segments, apically thickened pedicels,
and rather narrow sepals and wings to the upper petal.
Plants agreeing in all respects with Nicotra’s description and plate were
collected at Tangier by Rev. J. Roffey in the spring of 1916.
F. Jordanii, Gussone, Enum, Pl. Ins. Inar. (1854), seems from authentic
specimens to be a variation of var. Gussonet with flowers coloured as in the
specific type, and F. serotina, Guss. (L. c.) is perhaps a sha le-form of the same
plant. The former is stated to have fruits sometimes spotted at the apex.
There is Ischian material of both of these plants in Herb. Kew received from
Gussone himself.
F. recognita, Lacroix in Bull. Soc. Bot. France, vi. 551 (1859), described
as intermediate between /. Bastardii and F. Borei, is represented in
Herb. Kew by a specimen received from Lacroix in 1861 which is typical
F. Bastardii, with finely developed flowers and unusually long sepals. Other
material observed under this name shows large, dark-tipped flowers resembling
those of var. affinis, of which F. recognita is made a synonym by Rouy &
Foucaud.
F. pia, Nicotra in Att. Acc. Dafnica, Acireale, Ser. 2a, i. 5 (1905), may
also belong to this species, but its inadequate diagnosis renders its deter-
mination impossible in the absence of authentie specimens.
A form resembling the specifie type, but with remarkably small flowers
(only about 8 mm. long), was colleeted at Gibraltar by Major Wolley-Dod
in 1913.
F. Bastardii is one of the most difficult species of the genus and is unique
among the Grandijlorw owing to some of its forms having the inner petals
only tipped with purple, while in others this colour pervades also the wings
of the upper petal. In fruit, likewise, it is remarkably variable, the rugosity
being appreciably finer in the varieties patens, vagans, and Gussonei than in
GENERA FUMARIA AND RUPICAPNOS. 281
the type. On the other hand, the general habit, the oval, serrate sepals, and
the long-winged corolla are so constant and so marked in the plants brought
together by Haussknecht under this species, and it is so evident from
Herbarium material that they are largely connected by intermediates, that
they cannot with confidence be kept specifically apart.
The distribution of this chiefly Mediterranean species is given at p. 41 of
* Fumaria in Britain." Of its varieties, 8. patens was described from Corsica,
and y. vagans occurs in Southern France; var. Gussonei is common in Italy
and the neighbouring islands, and grows also in Algeria, Spain, and Greece,
as well as in Britain; var. hibernica is exclusively British ; var. affinis is
only certainly known from Spain; and var. benedicta has been seen from
Sicily, Sardinia, and Morocco.
A hybrid (apparently barren) between this species and F. muralis subsp.
Borwi has been observed in Guernsey.
** Series Eu-Murales.
Sepala magnitudinis variabilis; petali inferioris margines angustissimi,
erecti vel rarius patentes ; fructüs leves vel rugulosi.
20. Fumarta Munsyi, Boissier & Reuter, Pugillus, 5 (1852): Haussk.
in Flora, lvi. 536 (1873).
Exsiee, Reuter, Oran in suburbio La Mosquée, 1849, in Hb. Boiss.!
Boissier & Reuter, Prov. Oran, 1849, in Hb. Boiss, ut F. flabellata!
G. Munby, Oran, in Hb. Kew, ut F. capreolata !
Fumaria robusta sed lava, plus minusve elongata, interdum petiolis cirrhosis
scandens. Folia foliolis in lobos anguste oblongos acutos vel ovato-
oblongos mueronatos (forma major, Haussk.) irregulariter 2-3-pinnatisecta.
Racemi laviusculi, multi-(20-25-) flori, pedunculos breviusculos superantes.
Bracter anguste lineari-oblongse, acuminate, pedicellis fructiferis 5 mm.
longis, tenuibus, flexuosis, patenti-erectis vel patentibus tertiá parte breviores.
Sepala 2-4 mm. longa, 1°5-2°5 mm. lata, subrotundo-ovata vel ovata, peltata,
breviter acuminata, circumcirca (presertim ad basin) irregulariter dentata,
nervo dorsali viridi valde carinata, albida. Corolla speciosa, 12-13 mm.
longa (14 mm. in f. majore), pallide rosea; petalo superiore alis atropurpureis
reflexis carinam paulo excedentibus apicemque vix attingentibus obtuscusculo,
'aleare quam sepala longiore praedito ; petalo inferiore marginibus angustis
patulis pallidis apicem vix attingentibus acuto, sepe libero deflexo ; petalis
interioribus sursum curvatis apice atropurpureis, Fructús minimi, vix 2 mm.
longi ac lati, subrotundi, subacuti vel obtusi, circa medium latissimi, paululum
compressi et obscure carinati, siccitate apicis foveolis parvis paulo obscuris
sublieves.
LINN. JOURN.—BOTANY, VOL. XLIV. 2
282 MR. H. W. PUGSLEY: A REVISION OF THE
Quod ad formam pertinet, fructus F. Munbyi satis variabilis videtur, nam
quanquam in exemplaribus Boissieranis subacutus in specimine Herbarii
Kewensis rotundato-obtusus est.
Hee rara species, que propter flores magnos cum fructibus minimis
conjunetos notabilis videtur, per florum spectabilium racemos longos
F. Martinii proxima est, sed pedunculis longioribus, bracteis longioribus
angustioribusque, sepalis dentatis carinatis, petali inferioris marginibus
latioribus, et fructibus minimis plane differt.
F. muralis (sensu lato) racemis brevioribus, sepalis ecarinatis, petalo
superiore subspathalato, inferiore sine marginibus patulis, fructibus seepissime
majoribus distinguitur. Preterea, hoe species (sensu stricto) per habitum
gracilem et flores minores recedit, dum subspecies neglecta bracteas breviores,
sepala integriuscula, corollas minores, fructtisque obtusiores magis obovatos
habet, atque subspecies Boræi, ad quam F. Munbyi a cl. Hammar, descriptionem
in Boissier, Diag. Pl. Or. Ser. 11. i. p. 17 (1853), sequente, relata. est, sepalis
majoribus, petali superioris calcare breviore, fructibus multo majoribus obovatis
obtusissimis facile dignosei potest.
F. apiculata, Lge. habitu minus robusto, pedunculis brevissimis, pedicellis
suberectis, floribus minoribus acutioribus, sepalis angustioribus paucidentatis,
fructibus apieulo persistente preeditis separanda est.
F. Bastardii, Dor. per bracteas breviores, per sepala ovalia serrata, et per
fructüs majores rugosos nee sublieves a F. Munbyi longe distat.
F. Munbyi provinciam Oran in Algeria habitat.
21. Fumaria MARTINI, Clavaud in Act. Soc. Linn. Bordeaux, tom. XLII.
5° série, tom. 2, p. Ixix (1888). F. paradoxa, Pugsley, Fum. in Brit.
31 & 74 (1912).
Icon. Pugsley, l.c. fig. 4 (Journ. Bot. l. tab. 519), ut F. paradowa.
FExwsice. Bourgeau, Pl. d'Espagne, 1863, ut F. Bastardii! Magnier, FI.
Select. No. 1075!
Since the publication of this plant as a new species under the name of
F. paradoxa, it has been found that in 1888 a communication respecting
Martin's plant mentioned at page 74 of “ Fumaria in Britain” was made by
Clavaud to the Linnean Society of Bordeaux, as cited above, which defines
its salient features and its position in the genus, and may fairly be regarded
as a valid publication of the name F. Martinii. It is therefore proposed to
accept this name, reducing XF. paradowa to a synonym.
It curiously happens that Clavaud's communication was due to the plant
collected by Martin (C. Magnier, Fl. Selecta, No. 1075) being referred by
certain botanists to F. major, Badarre, just as occurred at first with the
British form (Fum. in Brit., p. 55), and it may thus be of interest to
reproduce some of Clavaud's observations.
GENERA FUMARIA AND RUPICAPNOs, 2823
* J'avais indiqué pour cette dernière plante . . . le nom de F. Martini,
en prévenant M. Magnier qu'il ne fallait y voir qu'une forme du stirpe
Capreolata voisine du F. Borei, Jord. et du F. muralis, Sond. et qu'elle
n'avait rien de commun avec le F. major, Bad. . . . M. Magnier n'en
conserva pas moins sur ses étiquettes le nom de F. major, Bad., en m'infor-
mant qu'il s’appuyait sur l'opinion de deux éminents botanistes . . . apres
+. HE m'a adressé une étiquette imprimée conforme à ma détermination
.. C Chacun peut voir et lire dans la Monographie d’Olof Hammar quels
sont les caractères distinctifs trés nets des F. capreolate et des F. agrariw
e. Je me plais à croire que les botanistes un peu au courant du genre
Fumaria n'auront pas de peine à reconnaitre que la plante de M. Martin
appartient au groupe du F. capreolata et wa rien de commun avec le F. major,
Bad... . Notre plante n'a d'analogie réelle qu'avee le F. Borwi, Jord.,
muralis, Sond. et speciosa, Jord., et j’ajoute qu’elle se distingue de ces trois
dernières formes elles-mémes par plusieurs caractères. . . . En somme, si,
comme croyaient Koch et Sonder, la terminaison ogivale du fruit a une
sérieuse importance, c'est, du F. muralis que notre plante se rapproche le
plus, malgré la dimension trés différente de ses fleurs. Si, au contraire, ce
caractère de Pachaine n'était guère ici qu'un. accident, comme il se pourrait,
notre plante se séparerait bien peu du F. Borai, Jord.”
It will thus be seen that Clavaud was the first to notice and to describe the
true affinities of this fumitory, and that the restoration of his name, F. Martinii,
is clearly due to him. In emphasizing its likeness to F. muralis and F. Borwi,
however, he scarcely allows sufficient importance to the long, lax, short-
peduncled racemes that lend it the general facies of F. major.
Rouy (Fl. France, i. 172) refers Martin's plant to F. flabellata, Gasp.—a
species lacking even the superficial resemblance that connects it with
F. major.
When growing, F. Martinii is a remarkably beautiful plant owing to its
long and graceful racemes and its brilliantly coloured flowers.
It is evidently distributed through the Iberian Peninsula, whence its range
extends through Western France to Southern Britain. Its Spanish localities
at present known are Navalmoral, Estremadura (Bourgeau, 1863, as
F. Bastardii!), Calatayud, Aragon (Vicioso, 1906, as F. affinis!), Ameyugo,
Ayelas, Pancorbo, Castile (Sennen, 1906, as F. muralis var. Paui!), and
Almeria, Andalusia (Huter, Porta & Rigo, 1879, as F. muralis!) In
Portugal it occurs at Vimioso, Pinello (Mariz, 1888, as F. agraria!), and
probably in other localities, where it has been confused with F. muralis
subsp. Borwi. The only French stations known are Romorantin, Loir-et-
Cher (Martin, 1884 !), Gatteville, Manche (Tardieu, 1880, as F. Boræi !) and
Rennes, Bretagne (Hb. Kew, as F. capreolata!). In Britain, in addition
to Penryn, Cornwall (Davey, 19041), it grows in the island of Guernsey
(Pugsley, 1914 !).
242
284 MR. H. W. PUGSLEY : A REVISION OF THE
22. Fumaria sEPIUM, Boissier, Diag. Pl. Or. Ser. rr. No. 1, p. 16 (1853) x
Hamm. Mon. 27 (1857); Haussk. in Flora, lvi. 525 (1873).
F. muralis var. platycarpa, Rouy & Foucaud, Fl. Fr. i. 173 (1895).
(PI. 9, figs. 8, 9.)
Vesice. Boiss. & Reuter, Algeciras, 1849, in Hb. Boiss.! Wolley-Dod,
Fl. Calpensis (Algeciras), Nos. 1711! et 201!
Fumaria annua, robusta, ramosa, elongata, ssepe petiolis cirrhosis longe late-
que scandens. Folia irregulariter 2-3-pinnatisecta, foliolis in lobos oblongos,
euneatos vel ovato-rotundos plus minusve obtusos et sepius longiuscule.
mucronatos fissis prædita. Racemi lawiusculi, pauciflori (0—12-flori), pedunculis
gracilibus vulgo strictis rarius paulo incurvatis breviores. Bractew anguste.
lineari-lanceolate, setaceo-acuminatie, pedicellos fructiferos 5 mm. longos,
tenuissimos, erecto-patentes, strictos vel flexuosos infimi subwquantes, ceterie-
breviores. Sepala 4—5 mm. longa, 2-2:5 mm. lata, ovata (in formis umbrosis
angustiora, lanceolata), peltata, plus minusve acuminata, basi parce dentata
aliter subintegra, preter nervum dorsalem viridem albida, corolla tubo
wquilata. Corolla magna, 12-14 mm. longa, speciosa, roseo-allida, priesertim
apicem versus late rubescens (in floribus serioribus pallida et interdum
conspicue minor); petalo superiore subacuto, vix dorsum compresso,
raleare longo deflexo alisque intense atropurpureis reflexis carinam paulo
excedentibus sed apicem vix attingentibus prædito; petalo inferiore marginibus
angustissimis erectis vel subpatentibus apicem haud attingentibus acuto, sæpe
libero deflexo ; petalis interioribus albidis vel rubescentibus apice atropurpureis.
Fructis parvi, cirea 2 mm. longi ae lati, subrotundi, obtusissimi, subcompressi,
sarinati, in vivo inferne in stipitem paulo obscurum quam pedicelli apex
paulisper angustiorem contracti, siccitate l;rves apicis foveolis parvis notati.
Deseriptio seeundum exemplaria viva ad Algeciras (in loco classico) a
Major Wolley-Dod collecta, Martio, 1913.
B. gaditana, var. nov.
F. gaditana, Haussk. l.c. 547; Willkomm & Lange, Fl. Hisp. iii. 879
(1880).
Exsice. Boissier & Reuter, Tangier, 1549, in Hb. Boiss. ! Boissier & Reuter,
Grazalema, Prov. Granada, 1849, in Hb. Boiss. ! Wolley-Dod, Fl. Calpensis,
Nos. 1629 ! 1647 ! 1701!
Racemi densiores, vulgo 10-16-flori; pedunculi pediceliique robustiores.
Bractew quam in typo breviores latioresque, lineari-oblongee, acuminate,
pedicellorum longitudinem dimidiam paulo superantes. Sepala 3-4 mm.
longa, 1:5-2 mm. lata, oblongo-ovata, acuta, saltem basin versus parce
irregulariter. dentata. Corolla typi sed tandem sepe omnino rubescens.
Fruetis quam in typo majores, 2:5 mm. longi ac lati, subrotundo-quadrati.
Hoc pulchra species, eujus primum forma umbrosa floribus depauperatis
a cl. Boissier et a cl. Hammar descripta est, nune e plantà normali depingitur.
Inter F. capreolatam et F. muralem medium fere tenet, sed ab alterà racemis
i
Leeuw ON Omm
GENERA FUMARIA AND RUPICAPNOS, 285
paueifloris, pedicellis tenuibus nec recurvatis, sepalis minoribus, sine stipite
distincto fractibus, ab alterà foliorum segmentis latioribus, pedunculis
longioribus, sepalis subintegris, corollis angustioribus albidis nee roseis plane
differt.
F. macrosepala et F. berberica subsectionis Capreolatarum pedicellis
erassioribus, bracteis sepalisque majoribus, fructibus majoribus rugosis nec
leevibus facile separand:e sunt.
Speciei typus hactenus in Algeciras Bæticæ solum inventus est, sed varietas
gaditana freti Gaditani contrarias partes in BeeticA et in Mauritania habitat.
Exemplar nune in herbario Mus. Brit. a el. R. P. Murray ad sepes prope
(intram Lusitaniæ collectum ad hane speciem, ut videtur, etiam referen-
dum est.
This fine but local fumitory was originally described by Boissier from a
depauperate form, collected at Algeciras late in the season, in which, as
sometimes happens with other species, the sepals were particularly narrow.
The flowers of this material were so poor that its identity with other
specimens obtained the same year by Boissier & Reuter at Tangier,
Grazalema, and Gibraltar does not appear to have been recognized by
Boissier or later by Haussknecht, although the Tangier example in Herb.
Boissier is labelled F. sepium in the handwriting of Reuter.
These last-named plants, as represented in Herb. Boissier, are more or less
normal, though collected late, and form the basis of Haussknecht’s
F. gaditana, which he seems to have regarded as in no way related to the
F. sepium from Algeciras. But it is clear that the original F. sepium is a
depauperate shade-form, and its specific identity with /. gaditana, first
suggested by Willkomm & Lange (/.¢.), may be confirmed by a comparison
with * Porta & Rigo, Iter IV Hisp. 1895, No. 610” from Algeciras and the
specimens collected in 1913 by Major Wolley-Dod in Boissier’s original
locality, some of whieh are shade-forms plainly showing a transition to the
state of the plant as found by Boissier. As Boissier’s name antedates
F. gaditana, it must necessarily stand for the species, and as the plant of
Algeciras, which is thus the specific type, seems to differ in minor details
from that found at Gibraltar and elsewhere, this latter form has been
distinguished as a separate variety gaditana.
23. FUMARIA MURALIS, Sonder in litt. ap. Koch, Synopsis Fl. Germ. ed. 2,
Appx. 1017 (1845); Haussk.in Flora,lvi. 523 (1873) ; Pugsley in Journ.
Bot. xl. 175 (1902), et Fum. in Brit. 16, 22 & 74, excl. F. apiculata
in syn. (1912). F. capreolata B. Burchellii, DC. Syst. ii. 183 (1821) ;
F. media y. muralis, Hamm. Mon. 29 (1857), non F. media, Lois.
Icones. Fl. Danica, tab. 2473 ; Hamm. l.c. tab. 4, ut F. media y. muralis
(fructu obtuso) ; Journ. Bot. xl. tab. 436, fig. 4.
Exsicc. Billot, Fl. G. & G. No. 2807! Mandon, Pl. Mader. No. 5!
Bourgeau, Pl. Canar. No. 1173, ut F. officinalis !
A
3
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MR, H. W. PUGSLEY : A REVISION OF THE
Fumaria annua, habitu semper gracili et sepius exiguo, interdum in arvis.
nana, suberecta, ramosior, interdum diffusa vel petiolis cirrhosis scandens.
Folia irregulariter 2-3-pinnatisecta, nonnunquam plus minusve glauca, foliolis
(sepius parvis) in lobos oblongos mucronatos vel lanceolatos acutos fissis.
Racemi laviusculi, pauciflori (ssepissime sub-12-flori), pedunculos graciles
rectos ineurvatosve fere wquantes. Bractew lineari-lanceolate, acuminate,
pedicellis fructiferis tenuibus qui plerumque recti et erecto-patentes sed varius
flexuosi et paulo reeurvati sunt tertiá parte breviores. Sepala 3—4 mm. longa,
1:5-2 mm. lata, ovata, peltata, sepissime acuminata, preesertim basin versus
et in margine inferiore plus minusve dentata, preter nervum dorsalem
viridiuseulum albida vel roseo-albida, latitudine corollæ tubum sequantia.
Corolla 9-10 mm. longa, rosea; petalo superiore dorsum compressiusculo,
haud lato, alis atropurpureis reflexis carinam superantibus apicemque fere
attingentibus apiculato et, ut in floribus junioribus saltem videtur, spathulato ;
"ileare sepala fere æquante ; petalo inferiore marginibus. erectis angustissimis
acuto vel apiculato ; petalis interioribus sursum curvatis apice atropurpureis,
Fructis parvi vel etiam minimi, circa 2 mm. longi (apiculo incluso) et
latitudine paulo minores, subrotundo-ovati vel subrotundi, usque ad maturitatem
acutiusculi vel obtusi brevissime apiculati, in stipitem prope obsoletum inferne
angustati, parum compressi et obscure carinati, in sicco apicis foveolis parvis
sepius paululum obscuris læves.
! Fructtis subglobosi sine apiculo obtusi.
B. lata, Lowe, Fl. Mader. i. 15 (1868).
Herba virescens, foliorum segmentis latiusculis, racemis 8—15-floris, sepalis
subintegris, corollis usque ad 12 mm. longis, lwte roseis apice atropurpureis,
Jructibus obtusis subglobosis obsolete rugulosis.
Lowe's types of this Madeira variety, notable for its large, bright flowers
with almost entire sepals, are in Herb. Mus. Brit.
[y. Lebelii, Rouy & Foucaud, Fl. Fr. i. 173 (1893) ; non vidi.
This plant appears to be chiefly notable for its dense racemes and broad
bracts and sepals. |
8. Lower, Pugsley, Fum. in Brit. 17 & 23 (1912).
In addition to the points of distinetion already assigned to this variety,
its sepals are commonly broader and less acuminate than in the specitic
type.
Lowe's specimens referable to this variety are in Herb, Mus. Brit.
)
1! Fructis subrotundo-ovati, acutiuseuli vel brevissime apiculati.
e. decipiens, Pugsley, Fum. in Brit. 23 (1912).
GENERA FUMARIA AND RUPICAPNOs. 287
F. Borai 8. muraliformis, Clavaud, Fl. Gironde, p. 49 and PI. iii, B 2 (1882),
appears from authentic material to be a form of F. muralis closely allied to
var. Lowei, and his B. verna and y. serotina early and late states of the same
form.
The present remarkable distribution of P. muralis is shown at pages 23
&74 of “ Fumaria in Britain.” Another link in the chain of habitats extending
from north-western Europe and the Atlantic Islands round South Africa to
the East Indies and New Zealand is supplied by a specimen in Herb, Mus.
Brit. (erroneously referred to F. parviflora, Lam.) from Utakamand, in the
Nilghiris of Southern India.
The variety decipiens occurs in Western France (Reverchon, Roche-sur-
Yon, Vendée, 1882, as F. Borwi!) as well as in Britain.
23a. Subspecies NEcLECTA, Pugsley. Fum. in Brit. 20 & 24 (1912).
Icon. Pugsley, l.e. fig. 2 (Journ. Bot. l. tab. 519).
23b. Subspecies Borat, Pugsley in Journ. Bot. xl. 132, 175 & 180 (1902) ;
Fum. in Brit. 18 & 25 (1912). F. Borei, Jordan in Cat. Grenoble
(1849), et Pugillus, 4 (1852) ; Haussk. in Flora, lvi. 520 (1873) ;
F. Bastardii B. major, Boreau in Duchartre, Rev. Bot. ii. 359
(1846-7); F. media a. typica, Hamm. Mon. 28, excl. syn. partim
(1851), non F. media, Lois. (PI. 9, fig. 10.)
Icones. Curtis, Fl. Londinensis, ii. tab. 145 (fase. vi. 47), ut F. capreolata ;
Smith, Eng. Bot. 943, ut F. capreolata ; Hamm. l.c. tab. 2, ut F. media a.
typica : Journ. Bot. xi. tab. 436, fig. 5.
Zrsice. Billot, Fl. Œ. & G. Nos. 2209 et 2209 bis, ut F. Borwi! Schultz,
Herb. Norm. No. 1007, ut F. Borat!
Fumaria ssepius robusta et satis ramosa, in arvis suberecta vel plus minusve
diffusa, in sepibus murisque elongata et seepe petiolis cirrhosis valde scandens.
Folia irregulariter 2—3-pinnatisecta, viridia (raro plus minusve glauca),
foliolis in lobes oblongos vel late euneiformes, acutos vel mucronatos fissis.
Racemi laviusculi, haud multiflori (sub-15-flori), pedunculos subequantes.
Bractew lineari-lanceolatze, acuminate, pedicellis paululum crassis rectis erecto-
patentibus (aliquanto flexuosis vel recurvatis in formis umbrosis) paulo
(interdum tertià parte) breviores. Sepala 4-5 mm. longa, 25-3 mm. lata,
ovata vel subrotundo-ovata, peltata, acuta vel breviter acuminata, sepissime
basin versus irregulariter dentata sed raro subintegra, preter nervum
viridiuseulum albida vel roseo-albida, corollae tubo latiora et inferne contigua.
Corolla 10-12 mm. longa, rosea vel raro rubra (forma rubens); petalo
superiore lato, dorsum compresso, alis .atropurpureis reflexis. carinam
superantibus apicemque sepe attingentibus acuto, apiculato vel etiam obtuso,
in gemmis ante alarum reflexionem spathulato, calcare sepalis breviore ;
petalo inferiore marginibus angustissimis erectis rarissime patulis acuto, ssepe
288 MR. H. W. PUGSLEY : A REVISION OF THE
libero deflexo ; petalis interioribus sursum curvatis apice atropurpureis.
Fructis modici, circa 2:5 mm. longi, 22:25 mm. lati. obovati vel subrotundo-
obovati, obtusissimi, paulo compressi sed obscure carinati, et in stipitem
obscurum quam pedicelli apex haud latiorem inferne angustati, siccitate apicis
foveolis parvis distinctis leres aut leviter rugulosi.
! Flores fere typi ; fruetüs subquadrati, rugulosi.
B. ambigua, Pugsley in Journ. Bot. xl. 178 & 180 (1902,; Fum. in
Brit. 26 (1912).
Varietas typi habitu sed foliorum lobis angustioribus lanceolatis, sepalis
angustioribus acuminatis, corollis paulo minoribus. — Fructs fere 2:5 mm.
longi, latitudine paulisper minores, subquadrati et inferne in stipitem pedicelli
apicem saltem æquantem viv angustati, quam in F. Borvi aliis formis magis
compressi, in sicco apicis foveolis obscuris leviter rugulosi.
`
y. hispanica, var. nov.
vesicc. Porta & Rigo, Iter II Hisp. 1890, No. 105, Almeria, Barranco
del Caballar, ut F. media, var. muralis !
Varietas bracteis lanceolatis acuminatis pedicellos patulos patenti-recurvosve
æquantibus vel infimis paulo superantibus; fructibus 2°25 mm. longis, 2 mm.
latis, subrotundo-quadratis, obtusissimis, inferne paululum angustatis, plane
rugulosis. Aliter ut in typo.
This plant is remarkable among the forms of the subspecies Borei for its
large, broad bracts and distinctly rugulose fruits.
!! Flores minores ; fructûs subrotundo-obovati, fere læves ;
plantæ graciliores,
6. gracilis, Pugsley, Fum. in Brit. 19 & 26 (1912).
e. britannica, Pugsley, Fum. in Brit. 19 & 27 (1912).
Zesice. Marshall, No. 2915, ut F. Boræi var. serotina !
€. longibracteata, Pugsley, Fum. in Brit. 19 & 27 (1912), ut sub-
varietas,
Hybrids between this subspecies and. A. officinalis, quite barren and inter-
mediate in eharaeters, have been observed in Guernsey and in the south of
England.
The subspecies Borei is a rare plant except in the British Islands and the
north and west of France, but there are specimens from Hesse, Germany,
and from Sweden in Herb. C. Bailey, and one from Norway in Herb. Kew.
Coutinho records it for Portugal, and it occurs in "pain, where it is reported
by Haussknecht to grow commonly about Gibraltar and elsewhere in
Southern Andalusia. As Major Wolley-Dod, however, recently failed to
find it in that district, and met with F. sepium in abundance, it is probable
that these two plants have been confused.
GENERA FUMARIA AND KUPICAPNOS. 289
The records for Morocco and Algeria, though supported by the authority
both of Hammar and Haussknecht, may also be doubted, the plants observed
in these two countries being probably F. sepium and F. Munbyi respectively.
Hammar's records for the Canaries and South Africa doubtless refer to true
F. muralis, to which likewise (perhaps var. læta) Haussknecht’s Madeira
locality in all probability belongs. The recognition of the subspecies Bort
and of F. muralis as Italian plants by Nicotra (Le Fumar. Ital.) seems due
to confusion with forms of F. Bastardii.
x FUMARIA PAINTERI, Pugsley, Fum. in Brit. 29 (1912).
Icon. Pugsley l.c. fig. 3 (Journ. Bot. l. tab. 519).
Although of presumably hybrid origin (F. muralis subsp. Borei x
F. officinalis ?) this plant appears to merit recognition as a more or less
established form, differing from other known /wmaria-hybrids in its general
development of perfect fruits.
24. lF'UMARIA APICULATA, Lange, Ind. Sem. Haun. 23, et Ann. Sci. Nat.
Ser. 1v. ii. 371 (1854) ; Hamm. Mon. Sliigt. Fum. 36 (1854), et Mon.
31 (1857). F. Reuteri, Boiss. Diag. Pl. Or. ii. No. 8, p. 13 (1849), et
Pugillus,4 (1852),ex parte; Haussk. in Flora, lvi. 538 (1873) : F. inedia
6. apiculata, Willkomm & Lange, Fl. Hisp. iii. 882 (1880), non
F. media, Lois.
Icon. Hamm. l.c. tab. 4.
Eesicc. Lange, Escorial, 1851, in Hb. Boiss.! Reuter, Castella prope
Miraflores ad radices Sierree de Guadarrama, in Hb. Boiss., ut F. Reuteri!
Durieu, Pl. Sel. Hispano-Lusit. Sec. 1, Asturicæ, 1835, No. 413, Cangas de
Tineo, ut F. capreolata! Lange, Cordoba ad sepes, 1852, in Hb. Kew,
ut F. Borat!
Hoe species hispanica, quae a cl. Hammar eum icone bene descripta est,
quanqaam cum F. Reuteri confusa, sine dubio F. muralem (sensu stricto)
maxime refert. A F. murali foliorum lobis angustioribus, racemis bre-
vissime peduneulatis, pedicellis fere erectis nec erecto-patentibus, sepalis
angustioribus ovato-lanceolatis paucidentatis, petalo superiore angustato
acuto, fructibus longioribus cum apiculo minimo persistente obtusiusculis
separari potest.
F. Reuteri, planta Granatensis quie ad F. 7 huretii, Boiss. a cl. Haussknecht
translata est, foliorum laciniis etiam angustioribus, pedicellis patentibus,
sepalis late oblongis, corollis minoribus (raro 10 mm. longis), fructibus
minimis subacutis stipitatis plane differt.
The closely cohering petals mentioned in Hammar’s diagnosis are
characteristic of weak or shade-grown plants in this as in other species.
In good flowers the lower petal of F. apiculata is deflexed and free, as is well
shown in the Durieu example in Herb. Kew.
BENED nets ari UT TT T bd TEF vU.
290 MR. H. W. PUGSLEY : A REVISION OF THE
This fumitory has been the subject of considerable eonfusion. It was first
included in 1849 in the original acconnt of F. Reuteri, a species founded by
Boissier on two Spanish plants, one from Castile and the other from the
Sierra Nevada in Andalusia. Five years later a second species, also
founded on a Castilian plant, was established by Lange as F. apiculata.
In Hammar’s Monograph these two species are retained, the description of
F. Reuteri being taken from Boissier’s original diagnosis and a specimen,
probably Andalusian, sent by him to Fries, while F. apiculata is described
and figured from living, cultivated plants presumably grown from seed
obtained from Lange. Haussknecht, who examined the original material of
these plants in Herb. Boissier, detected that the two Spanish examples cited
for F. Reuteri were essentially different, and retained Boissier’s name for the-
Castilian plant, referring the Andalusian one to the Greek F. Thuretii,
which it closely resembles. At the same time he made F. apiculata, Lge.,
of which Herb. Boissier also contains an original specimen, a synonym of
F. muralis, Sond. This arrangement is reflected in Willkomm & Lange’s
‘Flora Hispanica’ (except that F. apiculata there becomes F. media 8.
apiculata), but the authors do not appear familiar with the Castilian form
placed under F. Reuteri or with Boissier’s Andalusian plant.
It is evident from the types in Herb. Boissier that Haussknecht correctly
separated the two plants on which F. Reuteri was founded by Boissier.
But the original specimens also show with nearly equal certainty that the
Castilian F. Reuteri is conspecifie with F. apiculata, Lge., described from the
same district, and th t these two plants, while allied to F. muralis, are
distinet from it and from all other described forms. They agree well with
Hammar’s diagnosis of F. apiculata, except that the fruiting pedicels are
markedly suberect, as noted by Lange, instead of * erecto-patentes,” and the
fruit perhaps less oval than ovate.
The nomenclature of these plants is involved, for of the two forms
originally included by Boissier under 7. Reuteri, that inhabiting Andalusia
is apparently described under this name by Hammar, while Haussknecht
restricts the name to the Castilian form. It is clear that this latter plant
was the first to be segregated (as F. apiculata, Lange), and hence it seems
preferable to apply Boissier’s name to the Andalusian form and to maintain
F. apiculata, Lange, for that discovered in Castile. Moreover, although
both plants were originally cited for /. Reuteri by Boissier, it is probable
from the terms of his diagnosis that it was actually taken from the Andalusian
one which he himself collected, rather than from the Castilian plant gathered
by Heuter which he united with it.
25. Fumaria Perrert, Reichenbach, Icones Fl. Germ. iii. 1 (1838) ; Parlatore,
Mon. Fum. 85 (1844), ex parte ; Visiani, Fl. Dalmatica, iii. 98 (1852),
et Suppl. i. 118 (1872) : Hamm. Mon. 32 (1857), ex parte : Schlosser
& Vukot. Fl. Croatica, 205 (1869).
GENERA FUMARIA AND RUPICAPNOS. 291
Icon. Reichb. l.c. tab. 2, fig. 4453 /.
Kwsice. Pichler, Mte. Marian bei Spalato, 1885, in Hb. Zurich and Hb.
C. Bailey! Huter, Fl. Exsiec. Austro-Hungarica, No. 2900, cult., orig. Clissa
in insula Lesiná !
Fumaria annua, gracilescens, multo ramosa, diffusa, vix scandens. Folia
irregulariter. 2-3-pinnatisecta, foliolis flabellatis longiuscule petiolatis in
lobos sæpissime oblongos mucronatos fissis. Racemi laviusen!i, haud multifori
(vulgo sub-12-, raro ad 18-flori), brevissime pedunculati aut subsessiles.
Bractew lineari-lanceolatz, acuminate, albidee, pedicellos fructiferos apice in-
crassatos patulos vel erecto-patentes (rarius paulo recu rvos) subeguantes. Sepala
35-45 mm. longa, 2-275 mm. lata, ovalia vel subrhombea, conspicue peltata,
subacuta, preesertim medium versus irregulariter uni- vel paueidentata, rarius
integriuseula, nervo dorsali viridi spe lato albida, corollæ tubo paululum
latiora. Corolla 9-10°5 mm. longa, rosea; petalo superiore angustato subacuto
alis atropurpureis reflexis apicem raro attingentibus carinamque viridem
abrupte terminatam vix sequantibus et calcare longo adscendente preedito ;
petalo inferiore marginibus angustissimis erectis subacuto ; petalis interioribus
fere rectis apice atropurpureis. ucts mediocres, 2°5-2°75 mm. longi et
2 mm. lati, ovati vel fere turbinei, plus minusve apice attenuati, acutiusculi,
prope basin latissimi et in stipitem obscurum quam pedicelli apex angustiorem
abrupte angustati, carinati et paulo compressi, siccitate apicis foveolis
angustis satis distinctis rugulosi.
Hiec species racemis fere subsessilibus, sepalis subrhombeis, corollà anguste
alata, fructibus ovatis fere turbineis plus minusve attenuatis aeutiusculis inter
affines notabilis est.
Apud el. Boissier in * Flora Orientalis, i. 137 (1867), specimina F. Petteri
dalmatica a cl. Petter accepta F. Gussonei, Boiss. affines vel forme sunt,
sed hee planta, que vero Fumarie Bastardii, Dor. solum varietas est, racemis
plane peduneulatis, sepalis serratis multo minoribus, petalo superiore latius
alato, petali inferioris marginibus patulis, fructibus subrotundis obtusis
longe distat.
A F. murali, ond., quee in Koch, Synopsis Fl. Germanicie, ed. 2, p. 435,
ad hane speciem relata est, F. Petteri racemis subsessilibus, sepalis subrhombeis
paucidentatis, petalo superiore angustius alato, fructibus ovatis attenuatis
plane separatur.
Racemi laxi floribus magnis multiflori et fructûs qui quidem acutiusculi
sed subrotundi nec ovati sunt etiam Fumariam Martinii, Clav. sejungunt.
F. Munbyi, Boiss. racemis multifloris longius peduneulatis, sepalis magis
dentatis, corollis majoribus cum inferioris petali marginibus patulis, fructibus
subrotundis duplo minoribus facile distinguitur.
F. apiculata, Lge. pedicellis suberectis, sepalis ovato-lanceolatis, petalo
superiore acutiore sed alis latioribus preedito, fructibus apiculatis levibus
cum hae specie confundi non potest.
*
292 MR. H. W. PUGSLEY : A REVISION OF THE
In exemplaribus Fl. Exsice. Austro-Hungarica, No. 29001, flores fruetüsque,
ut in fumariis cultis expeetandum est, minores sunt. quam in speciminibus
agrestibus quae el. Th. Pichler in vineis ad montem Marian prope Spalato
collegit.
F. Petteri montem Marian prope Spalato, insula Lesinæ Clissam et forsan
locos alios in Dalmatia habitat.
The application of the name +. Petteri, Rchb. to this apparently rare
Dalmatian plant has been decided on only after careful consideration.
Although it appears from Hammar’s note (Mon. p. 33) that Petter sent out
various fumitories under this name, it seems fairly certain that the Dalmatian
plant on which Reichenbach founded his species was that cited in Visiani's
Fl. Dalmatica, viz. :—* F. media, Petter, Bot. Wegw. No. 421, non Lois.”
(1832), which was collected by Petter at Spalato. Hammar regards this
plant as conspecific with /. Thuretii, Boiss., but Boissier (l.e. supra) entirely
dissents from this view, regarding F. Petteri, Rchb. as allied rather with
F. Gussonei. Haussknecht reduces F. Petteri, Rchb. to a synonym of
F. Gussonei, and Haláesy (Fl. Greee. i. 48 (1901)) uses the name in preference
to F, Gussonei for Boissier’s plant. Koch again likened F. Petteri, Rchb. to
F. muralis, which he first referred to it (/. c, supra).
An explanation of these differing views is afforded by the later Spalato
specimens collected by T. Pichler and referred to F. Petteri, Rehb., which
agree with Reichenbach’s account, taken in conjunction with the diagnoses in
the Floras of Dalmatia and Croatia, and are no doubt identical with Petter’s
original plant described hy Reichenbach. It may be seen from these
specimens that while the species produces flowers resembling those of
F. Thuretii and similarly borne in subsessile racemes, thus accounting For
Hammar’s union of the two forms, its habit and foliage are very different and
rather recall F. muralis and F. Gussonei, as was noticed by Koch and
Boissier respectively. ‘The ovate, attenuate form of the fruit in F. Petteri,
moreover, is distinctly different from that obtaining in either of these allied
plants, and hence there seems sufficient reason for maintaining it as a separate
‘species.
Hammar's figure of F. Petteri (Mon., tab. iv.) seems compounded of this
and the following species, the foliage and corolla recalling F. Petter’, and
the pedicel, sepal and fruit /. T huretii.
*** Series Sub-Latisepale.
Sepala majuscula ; petali inferioris margines angustissimi, erecti aut rarius
subpatentes obscure apice dilatati; frucetüs minimi, plus minusve rugulosi :
foliorum laeiniz relative angustze ; flores minores in racemis subsessilibus.
26. Fumaria THureri, Boissier, Diag. Pl. Or. Ser. ut. No. 1, p. 15 (1853) ;
Fl. Orient. i. 137 (1867) : Haussk. in Flora, lvi. 494 (1873), excl. loc.
GENERA FUMARIA AND RUPICAPNOS. 293.
hisp. ; Haláesy, Fl. Gree. i. 46 (1901). F. Petteri, Hamm. Mon. 32.
(1857), ex parte, non Reicbb.
Evsiec. Heldreich, Herb. Greece. Norm. No. 1005 !
Fumaria annua, habitu satis robusto, basi precipue ramosa, nunc suberecta
et ssepe nana nunc in formis umbrosis diffusa subscandens. Folia irregulariter
2-3-pinnatisecta, glaucescentia, foliolis (sepius parvis) in lobos oblongos
mucronatos vel lineari-oblongos acutos fissis. Racemi sepius. multi- (15-35-),
flori, sub anthesin densi, tandem elongati, brevissime pedunculati vel subsessiles..
Bractew sublanceolatze, acuminate, pedicellos fructiferos breviusculos apice-
incrassatos, ut in F. capreolatd arcuato-recurvos (in formis umbrosis vulgo-
rectos divaricatos), subequantes. Sepala 2°5-3 mm. longa, 1:5-2 mm. lata,
ovata vel rarius subrhombea, peltata, acuta, parce irregulariter repando-
dentata, preter nervum dorsalem viridiusculum albida vel subrosea, coroll»
tubum subequantia. Corolla circa 9 mm. longa, intense rosea, ei F. Petteri
subsimilis ; petalo superiore angustato subacuto alis atropurpureis reflexis apicem
raro attingentibus carinamque viridem subtruncatam vix zequantibus et
calcare adscendente przedito ; petalo inferiore marginibus angustissimis erectis
acuto ; petalis interioribus apice atropurpureis. /ructis minimi, circa 2 mm.
longi ac 1:51:75 mm. lati, ovati vel subrotundo-ovati, acutiusculi et inferne
in stipitem quam pedicelli apex angustiorem contracti, paulum compressi et
obscure earinati, siccitate apicis foveolis minutis distinctis plus minusve
rugulosi.
! Corolla quam in typo major ; foliorum laciniæ angustiores.
B. Heldreichii, Boiss. Fl. Orient. i. 137 (1867) ; Haláesy, l.c. 46 (1901).
F. Heldreichii, Boiss. Diag. Pl. Or. Ser. u. No. 1, p. 16 (1853) ;
Hamm. l. e. 33 (1857).
Exsice. Haussknecht, Iter Graecum, 1885, Athenæ, in valle Cephissi, in
Hb. Mus. Brit., ut F. Heldreichii !
Varietas habitu subscandente et foliis pallide virentibus in lacinias lineari-
oblongas fissis. Dractere lineari-lanceolate, pedunculos fructiferos recurvos
sequantes. Corolla plus 9 mm. longa, pallide rosea. Fructús minimi, vix
2 mm. longi et paululum angustiores, subrotundi et breviter acutiusculi,
levissime rugulosi. Aliter ut in typo.
The exsiceata of this variety, “ Heldreich, Herb. Græc. Norm. No. 9038,”
as represented in Herb. Mus. Brit., is a lax and vigorous plant with suberect
fruiting pedicels and fruits less globose than ovoid. The station whence it
was obtained, Mt. Pentelicos, produces a number of forms of this variable
species.
y. deflexa, var. nov.
F. deflexa, Heuffel in Flora, xxxvi. 619 (1853).
Kxsice. Heldr. Herb. Greece. Norm. No. 1005/, ut F. Thuretii £. fleribundat
294 MR. H. W. PUGSLEY : A REVISION OF THE
Planta typo robustior, internodis longis sepe scandens. Folia glauca,
laciniis — lineari-oblongis planis. Racemi — sub-30-flori, floribus | cernuis
laviusculi ; bracteæ lineari-lanceolate pedicellos fructiferos arcuato-recurvos
plus minusve incrassatos zequantes vel etiam superantes. Corolla 9-10 mm.
longa, rosea, quam in typo major. Fructis typi, nisi apicis foveolis paululum
majoribus.
The fruits of this form are described by Heuffel-as * subrotundis obtusis,"
but those observed in recent specimens from the Banatus (Degen, prope
Orsovam, 19041), as well as in Heldreich’s Greek exsiecata, are of the
subrotund-ovate, subacute form seen in the specitie type.
! 1 Corolla quam in typo haud major ; pedicelli fructiferi raro recurvi.
ò. pikermiana, var. nov.
F. pikermiana, Boiss. Diag. Pl. Or. Ser. 11. No. 6, p. 9 (1859) ; Fl. Orient.
i. 137 (1867) ; Haussk. l. e. 493 (1873) : Haláesy, Le. 48 (1901).
.— leon. Stefani, Major & Barbey, Samos, tab. 5 (1891), ut F. pikermiana.
Hesiec, Guieciardi, No. 3189, Pikermi Attiem, in Hb. Boiss, ! Heldreieh,
No. 771, Chelidoni Attics, in Hb. Boiss., ut F. bracteata! Pichler, PI.
Gree. Exsicce., Euboa, 1876, in Hb. Mus. Brit., ut F. Thuretii var. glauca!
Haussk. Iter Græcum, Laurion, 1885, in Hb. Mus. Brit., ut F. pikermiana !
Caulis multo ramosa; foliorum laciniæœ lineari-oblongw sæpius obtuse
mucronatæ. Racemi lawiuseuli bracteis sublanceolatis pedicellos incrassatos
patulos vulgo superantibus. Flores intense rosei, eis typi similes sed minores ;
corolla 7-8°5 mm. longa. Fructis minimi, vix 2 mm. longi, ovati, acutiusculi.
Aliter ut in typo.
The material examined of this variety consists of branches of large plants
gathered late and showing poor flowers. Although described as a distinct
species by Boissier, and maintained as such both by Haussknecht and
Halácsy, it is difficult to see how it essentially differs from F. Mhuretii, of
which it possesses the characteristic narrowly-winged coroila without (so far
as can be seen) any spathulate dilation of the lower petal. The length of its
bracts are sometimes equalled in other forms of this species, and it may be
doubted whether its small flowers and spreading pedicels are not merely the
result of a depauperate condition such as sometimes produces similar features
in F. capreolata. The figure in ‘Samos’ also portrays the general features of
F. Thuretii, excepting its recurved pedicels ; and the absence of wings to the
upper petal of the enlarged flower in this plate seems due to an artist’s error,
the wings being sufficiently indicated in the flowers of the racemes. In the
absence of good material and in deference to the views of botanists acquainted
with the living plant, F. pikermiana is retained as a variety.
The Zurich collection contains a number of varying depauperate or shade-
forms of F. Thuretii, mostly much branched, and showing ample foliage and
bo
95
GENERA FUMARIA AND RUPICAPNOS.
relatively few-flowered racemes with small flowers and spreading fruiting
pedicels. Some of these very closely approach this variety pikermiana.
e. thasia, var. nov. í
HKesice. Sintenis & Bornmüller, Iter Turcicum, 1891, No. 447 (Limenas,
Insula Thasos), ut F. Thuretii! Dornmüller, Iter Anatolicum tertium, 1899,
No. 4046 (Mudania, Bithynia), ut F. anatolica !
Planta gracilis, scandens, racemis sub-20-floris, pedicellis fructiferis
gracilibus, sepius irregulariter sub-patentibus nee recurvis, bracted lanceolata
acuminata plane longioribus ; typi sepalis et corolla, nisi petali superioris alis
atropurpureis apicem attingentibus carinamque superantibus ; fructibus minimis,
vix 2 mm. longis et 1:5 mm. latis, subrotundo-ovatis, subacutis, in sicco
minute rugulosis.
Hee species variabilis ad Fumariam Petteri accedit, sed foliorum laciniis
minoribus angustioribusque, pedicellis saepissime arcuato-recurvis, floribus
minoribus, sepalis magis ovatis, fructibus minimis nec inferne abrupte angus-
tatis neque apice attenuatis specifice differre videtur.
F. Thuretii in Griecià, ubi copiose crescere videtur, in Cypro, et in insulis
Principum prope Byzantium invenitur ; varietates /Zeldreichii et pikermiana
in Greeid ; var. defleva in Greecid, in. Banatu, et forsan ia Hercegovina et
in Montenegro; var. thasia in insula olim turcicà Thasos, et in Bithynia.
This species, both as F. Thuretii and F. pikermiana, is placed by Haussknecht
in the subsection Latisepalw of the Parviflora, although included by Hammar
among the Capreolate. Owing to its relatively fine leaf-cutting and large
sepals it certainly approaches the Latisepala, but the examination of good
and abundant material shows its floral features to be essentially those of
the section Grandiflora.
27. Fumarta Revver, Doissier, Diag. Pl. Or. ii. No. 8, p. 13 (1849), et
Pugillus, 4 (1852), ex parte; Hamm. Mon. 35 (1857), excl. loc. not.
Castell, pro parte. F. Thuretii, Boiss. (ex parte) ap. Haussk. in
Flora, lvi. 494 (1873) ; F. parviflora B. segetalis, Hamm. l. e. 17, et
Willkomm & Lange, Fl. Hisp. iii. 885 (1880), pro parte; F. segetalis,
Joutinho, Fl. Portugal, 246 (1913).
Jesice. Boissier, Sierra Nevada prope San Geronimo, in Hb. Boiss. ! Ball,
Iter Hisp. 35, ex regione subalpinà montium S. Nevada, in jugis Cerro
Tesoro et La Cartajuela, 5—6000 ft., in Hb. Kew, ut F. bætica ! Reverchon,
Pl.d'Andalousie, 1889, Sierra de Ronda, ut F. media var. apiculata! Reverchon,
Pl. d'Espagne, 1891, No. 572, Sierra de Segorbe, Valence, ut F. Bastardii!
Fumaria annua, gracilescens, multo ramosa, suberecta vel diffusa, haud
scandens. Folia irregulariter 3-pinnatisecta, foliolis in lacinias parvas confertas
oblongas vel lineari-oblongas acutas vel mucronatas interdum subcanaliculatas
fissis. Racemi lawiusculi, 10-18-flori, tandem elongati, brevissime pedunculati
vel subsessiles. Bractew lineari-oblongze, cuspidatze, viridescentes, pedicellis
296 MR. H. W. PUGSLEY : A REVISION OF THE
fructiferis apice parum incrassatis flexuosis irregulariter patentibus nec
recurvis paulo breviores. Sepala 275-3 mm, longa, 15-2 mm. lata, late oblonga,
peltata, breviter acuta vel obtusa mucronata, basi sepius. subtruncata,
integriuscula vel parce irregulariter denticulata, preter nervum dorsalem
viridiusculum albida, corolle tubo zequilata. Corolla 9-10 mm. longa, pallide
rosea; petalo superiore subaeuto, calcare magno longicollo deflexo sepala
superante et alis atropurpureis reflexis apicem attingentibus carinamque
paululum superantibus prodito ; petalo inferiore marginibus angustissimis
suberectis nonnunquam ut in sectione. Parviflorarum apice paululum dilatatis
subacuto ; petalis interioribus fere rectis apice atropurpurels. Fructis
minimi, angusti, circa 2 mm. longi, 1°5 mm. lati, ovato-elliptici, juxta medium
latissimi, in apicem acutiuseulum vel fere acutum et in stipitem angustissimum
subeequaliter angustati, paulo compressi e£ plane carinati, siccitate apicis
foveolis minutis distinctis leviter. rugulosi. :
Hæc rara planta, ab auctoribus multo eonfusa et a el. Haussknecht ad
F. Thuretii relata, ei sine dubio valde affinis est sed habitu minus robusto,
foliorum laciniis confertis minoribus, racemis cum pedicellis patentibus haud
multifloris, sepalis latioribus integriuseulis, corollis pallidioribus petalo
superiore latius alato caleareque longiore preedito, fructibus semper angustis
stipitatis satis differt.
F. Reuteri in regione subalpinà montium in Sierra de Ronda, in Sierra
Nevada, et in Sierra Segorbe Hispanic australis, atque in Sierré de Serpa
Lusitanis habitat, ubi indigena certe videtur.
The confusion that has oceurred between F. Reuteri and F. apiculata, Lge.,
giving rise to a complicated synonymy, has been dealt with under the latter
species.
A specimen collected at Granada by Lange was associated with 7. parviflora
by Hammar as a variety segetalis (Mon. 1. e.), although at first referred to
F. Reuteri. Lange himself also named this plant F. Reuteri (in Kjoeb.
Vidensk. Meddel. 65 (1865)), but afterwards in the * Flora Hispanica? he
followed Hammar, remarking, however, “forsan specifice distinguenda."
Hammars diagnosis of his var. segetalisis tco briet for certain determination,
but Lange’s clearly points to a plant specifically distinct from F. parviflora ;
and on its discovery and recognition in Portugal it was raised to specific
nk as F. segetalis in Coutinho’s recent * Flora de Portugal? Professor
Coutinho has been kind enough to send an authentic Portuguese specimen
which confirms its identity with the original Andalusian 7. Reuteri of
Boissier, and F. parviflora B. segetalis, Hamm. and F. segetalis, Coutinho,
must accordingly be added to the synonymy of that species, with the
modification ** pro parte? in the case of the former, Hammar having also
deseribed the plant in question under the valid name F. Reuteri.
F. Reuteri approximates, both in flowers and in foliage, to the species of the
section Parviflora, and more than any other member of this subsection shows
GENERA FUMARIA AND RUPICAPNOS. 297
a slight apieal dilation of the lower petal. It is retained among the
Grandiflore, however, as it is obviously much more closely allied to
the preceding species F. Thuretii than to any member of the small-flowered
group.
Srctto IT. PARVIFLORA.
Parviflore, Pugsley, Fum. in Brit. 45 (1912). — Officinales, Hamm. Mon.
9 (1857) ; Angustisectee, Haussk. in Flora, lvi. 404 (1873).
Foliorum laciniz plane aut canaliculate, lanceolatze vel anguste oblongee
usque ad lineares vel setaceæ (F. montand, Schmidt, except), nonnunquam
mucronate. Flores rarissime quam in F. oficinali, L. majores, plerumque
minores, nec plus 9 mm. longi ; petalum superius apice gibbum format cujus
margines (alee) vulgo sursum reflexi sed plus minusve erecto-patentes, rarius
patuli sunt ; petalum inferius apice gibbum monstrans qui margines patentes
apieem versus plane dilatatos habet plus minusve spathulatum est. Petala
interiora ssepius parum curvata, quam in seetione I relative latiora.
SussECTIO IV. LATISEPALZE.
Latisepale, Haussk. in Flora, lvi. 493 (1873) ; Pugsley, Fum. in Brit. 54
(1912). Oficinales, Hamm. Mon. 9 (1857), ut sectio, ex parte.
Pedunculi breves (F. montand excepià) pedicellis plerumque apice valde
incrassatis preediti. Bracteæ, nisi in F. rostellatá, pedicellos fructiferos vulgo
superantes. Sepala magna, ssepe parum dentata, longitudine corolle tertiam
partem saltem æquantia, ejusque tubo latiora. Petalum inferius quam in sub-
sectionibus sequentibus obscurius spathulatum. Fructis minimi usque ad
majusculi, plus minusve subrotundi, siccitate rugosi, rugulosi vel subleves.
In this subseetion the wings of the upper petal are dark purple like
the tip of the inner ones in F. Araliki, F. micrantha, F. bracteosa, and
F. rostellata, but sometimes obscyrely so in F. micrantha. In F. montana
the dark colouring appears to be absent not only from the wings of the upper
petal but from the tip of the inner ones.
28. FUMARIA MONTANA, J. A. Schmidt, Beitr. Flora Cap Verd. Inseln,
963 (1852). (Pl. 11, fio. 29
Eesice. Schmidt—In rupestribus montium Ins, S. Antonii, März 1851, in
Hb. Vindob. !
Fumaria annua, ut videtur, habitu modice robusto, satis ramosa, diffusa,
petiolis cirrhosis scandens. Folia foliolis in lobos oblongos vel cuneatos,
mucronatos, planos, obtusos vel rarius emarginatos fissis, infimis longiuscule
petiolatis, irregulariter 2—3-pinnatiseeta. Racemi pauciffori (10-12-flori),
lawiusculi, pedunculos rectos crassiusculos subequantes. Bractew lineari-
lanceolate, acuminate, longissimse, pedicellis fructiferis apice incrassatis
LINN. JOURN.—BOTANY, VOL. XLIV. 25
298 MR. H. W. PUGSLEY : A REVISION OF THE
flexuosis et sepius ut in F. capreolatd arcuato-recurvis semper longiores.
Sepala 3-3°5 mm. longa, 1°5-2 mm. lata, lanceolato-ovata, acuminata, peltata,
grosse serrato-dentata, preter nervum viridiuseulum | albida, corolla tubo
paululum latiora. Corolla 6-7 mm. longa, roseo-albida; petalo superiore
subacuto alis albidis patenti-reflexis apicem vix attingentibus et carinam
viridem haud sequantibus preedito ; petalo inferiore marginibus patentibus
albidis subspathulato ; petalis interioribus, ut videtur, omnino albidis.
Fructûs majusculi, 2:5 mm. longi et 2°75 mm. lati, subglobosi, cirea medium
latissimi, in apicem rotundato-obtusum et in stipitem obscurum equaliter
angustati, paululum compressi sed insigniter carinati, in sicco dense sed
tenuiter rugosi, nitidi, et apicis foveolis parvis angustis distinetis notati.
The diagnosis of this very rare fumitory, known only from the Cape Verde
Island of S. Antonio and unique in its combination of small flowers and
broad leaf-segments, is taken from Schmidt’s specimen at Vienna, which has
been lent for examination through the kindness of Dr. Zahlbruckner. This
specimen now shows very few developed flowers, but such as remain are
quite small, with the subspathulate lower petal elearly visible. The essential
smallness of the flower is emphasized in Schmidt's very fair description,
although, influenced probably by the broad leaf-segments, he contrasts it with
F. agraria rather than any member of the Parviflore. Without more ample
material it is perhaps impossible to determine with certainty the plant’s exact
affinities, but as not only the floral characters, both of sepals and corolla, but
also the form of its fruit recalls the Latisepale, it seems preferable to place it
in this group, in spite of its foliage, rather than to regard it as an Agrarian
species with a degenerate corolla. In the dried specimen the peculiar
unevenness of the leaf-margins alluded to by Schmidt does not now seem
traceable.
F. montana is said to inhabit the rocks of the higher mountains of
S. Antonio, probably at about 5000 ft. alt. (the highest point in the island is
7400 ft.), and even at this height, in the latitude of Cape Verde, the humid
atmosphere may aecount in some measure for the development of its foliage.
The occurrence of an endemic species of the Parviffore on the west side of
Africa is somewhat unexpected, but it may be noted that another species of
this subsection, F. bracteosa, Pomel, is chiefly North African.
The relationship of F. montana with the endemic species of the Canaries
that has been referred to it, is dealt with under F. coccinea.
29. FUMARIA KRALIKII, Jordan in Cat. Dijon, 19 (1848), et Linnwa, xxiii.
471 (1850) ; Hamm. Mon. 23 (1857). F. anatolica, Boiss. in Pinard,
Pl. de Carie Exsice. (1842), nomen solum, et Diag. Pl. Or. ii. No. 8,
p. 14 (1849) ; Haussk. in Flora, lvi. 505 (1873).
Icon. Hamm. l. c. tab. ii.
Exsice. Sintenis & Bornmiiller, Iter Turcicum, 1891, No. 97, Dedeagatsch,
ut F. anatolica ! Schultz, Herb. Norm. Nov. Ser. Cent. 20, No. 1912, Nisch,
GENERA FUMARIA AND RUPICAPNOS. 299
ut F. anatolica ! Heldreich, Pl. Exsicc. Grecis, No. 3479, ut F. anatolica
(var. minor ?)! Sintenis, Iter Thessalonicum, 1896, No. 433, ut F. anatolica !
This very pretty fumitory has the aspect, when dried, of a miniature
F. purpurea owing to its recurved, purplish flowers with large, whitish sepals.
It also recalls F. Thuretii in the abruptly terminated keels of the outer petals
and the slight development of the wings of the upper one, but it may be
readily distinguished by the subspathulate lower petal of its much smaller
flowers, which do not exceed 7 mm. in length, as well as by its much more
obtuse fruits.
F. Kralikii is well described by Jordan (loc. cit.), who is largely followed
by Hammar, but its bracts are oblong or linear-oblong rather than oblong-
obovate, the wings of the upper petal are narrow and do not cover the keel,
and the very small, subglobose fruits (L:5-1:75 mm. long and broad), which
show distinct if very small apical pits, are less “leves” than “subleves vel
leviter rugulosi In some forms the racemes are at first very short and
dense, though afterwards elongating.
In addition to the habitats cited by Haussknecht, this species grows in
Bulgaria (Herb. Mus. Brit.) and in Armenia (Herb. Kew), and would thus
seem to oceur throughout the Balkan Peninsula and Asiatie Turkey, with the
exception of Mesopotamia and Arabia. It is also represented in Herb. Mus.
Brit. from the Crimea (Herb. Pallas, ut F. officinalis, partim), the label
noting that it grows copiously on rocks.
30. FUMARIA MICRANTHA, Lagasca, Elench. Hort. Matrit. et Gen. Spec. PI.
Nov. 21 (1816); Hamm. Mon. 21 (1857); Pugsley, Fum. in Brit.
54 (1912). F: densiflora, DC. Cat. Hort. Monsp. 113 (1813), et
Prodr. Syst. Nat. i. 130 (1824), ex parte; Haussk. in Flora, lvi. 507
(1813) ; F. calycina, Bab. in Trans. Bot. Soc. Edinb. i. 34 (1844) ;
F. mucronulata, Schur, Enum, Pl. Transsilv. 38 (1866) ?
F. tenuifolia, Gerard, Herb. 928 (1597), described from plants found in
cornfields between Charlton and Greenwich. In Johnson’s ‘Gerard’ the
description was changed to fit Platycapnos spicatus, Bernh., with which it
was generally confused by succeeding authors although apparently distin-
guished by Morison (Hist. ii. p. 262 (1680)) as F. vulgaris minor tenuifolia.
Icones. Eng. Bot. Suppl. 2876 ; Hooker, Icones, iv. tab. 363; Hamm. l. e.
‘tab. ii.
B. dubia, var. nov. F. micrantha forma dubia, Pugsley, l.c. 57
& 58 (1912); F. micrantha y. Parlatoriana, Rouy & Foucaud, FI.
Fr. i. 179 (1893) ? non Boissier.
Icon. Fl. Danica, tab. 2472, ut F. micrantha.
Exsice. Ridley & Fawcett, Wareham, in Hb. Mus. Brit.! Billot, Fl. G. & G.
No. 709, ut F. densiflora !
2B2
300 MR. H. W. PUGSLEY : A REVISION OF THE
In addition to the distinguishing features pointed out in “ Fumaria in
Britain," this variety is usually somewhat less robust and more freely
branched than the specifie type. Rouy & Foucaud's description of var.
Parlatoriana seems to refer to this plant rather than that so named by
Boissier.
[y. littoralis, Rouy & Foucaud, Fl. Fr. i. 179 (1893); n. v. F. littoralis,
Du Mortier, Bull. Soc. Roy. Bot. Belg. vii. 359 (1868). This plant is allied
to the preceding variety, being much branched and having similar relatively
narrow sepals; but it appears to be separable owing to its broader leaf-
segments and short racemes of pale flowers. |
It is noteworthy that prior to the appearance of Grenier & Godron’s ‘ Flore
de France’ in 1847, F. densiflora DC. was not associated by botanists with
this species. This may be seen not only from De Candolle’s works, but in
Sibthorp & Smith’s ‘Flora Greea,’ vii. p. 63, in Moris's ‘Flora Sardoa,’ i. p. 90,
and in Arnott’s paper on the genus in Rep. Bot. Soc. Edinburgh, p. 104
(1840).
F. micrantha is shown in “ Fumaria in Britain”
to be eminently a
Mediterranean species. The varieties dubia and littoralis are at present
known only from Western Europe.
31. FUMARIA BRACTEOSA, Pomel, Nouv. Mat. Fl. Atlant. 239 (1874).
F. densiflora B. bracteosa, Batt. et Trabut, Fl. de l'Algérie, i. 29 (1888) ;
F. Parlatoriana, Kralik, Pl. ZEgypt. Exs. (nomen); F. micrantha
B. Parlatoriana, Boiss. Fl. Orient. i. 137 (1867) ; F. densiflora
B. Parlatoriana, Haussk. in Flora, lvi. 510 (1873).
Exsice. Letourneux, Pl. Aigypt. No. 229a, Mariout, 1879, ut F. densiflora
(partim) ! Bornmüller, Iter Syriacum, 1897, No. 43, Jaffa, ut F. Parlatoriana!
Sintenis & Rigo, Iter Cyprium, 1880, No. 175, ut F. parviflora !
Fumaria annua, caulibus crassis elongatis robusta, parce ramosa, diffusa,
suberecta, vix scandens. Folia glauca foliolis in lacinias angustissimas
lineari-setaceas canaliculatas acutas fissis 2—4-pinnatisecta. Racemi floriferi
densissimi, fructiferi laxiores, multiflori (20—30-flori), pedunculos crassos
brevissimos multo superantes. — Bractee magne, lineari-oblonge, acuminate,
viridescentes, pedicellis fructiferis erecto-patentibus brevissimis (cirea 2 mm.
longis) apicem versus conspicue incrassatis plane (siepe. subduplo) longiores.
Sepala l:5-2 mm. longa, *75-1:25 mm. lata, rotundato-ovata vel ovato-
lanceolata, peltata, acuminata, denticulata, nervo dorsali roseo albida. Corolla
minima, 3:5—4:5 mm. longa, pallide rosea; petalo superiore. subacuto alis
atropurpureis carinam conspicuam viridem haud sequantibus apicemque vix
attingentibus atque calcare brevissimo fere gibbiformi ; petalo inferiore
marginibus angustis roseis subspathulato ; petalis interioribus apice atropur-
pureis. Fructûs modici, circa 2 mm. longi et lati, subrotundi, obtusissimi vel
subtruncati, parum. compressi sed conspicue carinati, siccitate apicis foveolis
obseuris plane rugosi.
ae TT SL
GENERA FUMARIA AND RUPICAPNOS. 301
Hee Fumaria que F. micranthe faciem eximie exhibet et pro varietate
ejus speciei a pluribus auctoribus habetur, per flores generis minimos breviter
calearatos notabilis est, et hoc charactere cum foliorum laciniis setaceis,
pedicellis brevissimis, fructibusque conspicue carinatis specifice differre
videtur.
F. bracteosa Palestinam (Jaffa (Bornmüller) ! et prope Beersheba
(Hb. Mus. Brit.)!), Mesopotamiam (Schlüfli, prope Bagdad, in Hb. Kew !),
insulam Cyprum, Egyptum (prope Alexandriam!) et Algeriam (Faure, prope
Oran !) habitat.
This, the smallest flowered of all fumitories, is shown as a distinct species
rather than a variety of F. micrantha, chiefly on account of its corolla,
whieh is not only remarkably small but of characteristic form with a
much diminished spur. This feature is quite constant in the different
herbarium material examined, some of which shows good and abundant
flowers, and there seems no sufficient reason for regarding the plant as a
depauperate form or condition of F. micrantha.
32. FUMARIA ROSTELLATA, Knaf in Flora, xxix. 290 (1846) ; Hamm. Mon.
20 (1857) ; Haussk. in Flora, lvi. 510 (1873). F. transsilvanica, Schur
(=F. macrosepala, Schur, non Boiss.), Enum. Pl. Transsilv. 38
(1866), ap. Haussk. l. c.
keon. Hamm. l. c. tab. ii.
Ewsicc. Celakovsky, &c. Fl. Exsice. Austro-Hungarica, No. 2902 !
An authentic example of F. prehensilis, Kit. in Hb. Mus. Brit. is identical
with this species, but according to Parlatore specimens of F. micrantha were
sent out by Kitaibel under this name, and the posthumous diagnosis of
F. prehensilis in Linnza, xxxii. p. 493 (1863), shows evident confusion
between F. rostellata and F. officinalis. It thus seems clear that Hammar
and Haussknecht were wise in passing over Kitaibel's name (Ind. Pl. Hort.
Pesth, p. 10 (1812)), though anterior to both F. micrantha and F. rostellata.
Another name that appears to have been generally overlooked is F. Sturmii,
Opiz in Naturalientausch, x. p. 267 (1825). This is referred to F. micrantha in
the ‘Index Kewensis,’ but both the diagnosis and the habitats cited recall
F. vostellata, and if this identification can be confirmed from authentic
exsiecata, the name F. Sturmii must be used in preference to F. rostellata.
Knaf's description of this species is accurate and fairly complete, and is
largely followed by Hammar. The fruiting pedicels are slenderer and the
bracts smaller than in the other plants of this subsection, and the pedicels are
by no means always short as stated in Hammar's diagnosis. In some agrestal
forms with highly coloured flowers in very floriferous racemes the wings of
the upper petal are much developed and extend almost to the apex, which is
then no longer rostellate though still acute. Such plants are possibly
varietally distinct.
302 MR. H. W. PUGSLEY : A REVISION OF THE
There is also some degree of variation in the fruit, which is sometimes
plainly longer than broad and occasionally very shortly acute. Its surface,
when dry, is usually finely rugulose.
The range of F. rostellata, a plant of Central Europe, extends as far north
as Lithuania (Grodno, in Hb. Kew !), and in addition to the habitats cited by
Haussknecht, it occurs in Serbia (Hb. Kew) and in Bulgaria (Hb. Mus. Brit.).
Sunskcrio V. OFFICINALES.
Officinales, Haussk. in Flora, lvi. 404 (1873) ; Pugsley, Fum. in Brit. 45
(1912) : Hamm. Mon. 9 (1857), ut sectio, ex parte.
Pedunculi breves vel breviusculi, nisi in /. Boissieri, pedicellis ssepius
apice parum incrassatis preediti, Bractew pedicellis fructiferis breviores
(F. mierostachys exceptà). Sepala haud magna, plus minusve dentata, raro
corollie tertià parte longiora, ejusque tubo (nisi in F. microstachys) angustiora.
Fruetüs modici aut maximi, conspicue lati, truncati vel retusi, siccitate
rugosi rarius rugulosi.
In F. officinalis and F. microstachys the wings of the upper petal are normally
dark purple like the tip of the inner ones, but this colouring is obscure in
F. cilicica and F. Boissieri.
33. FUMARIA OFFICINALIS, Linn. Spec. Plant. 700 (1753); Hamm. Mon. 9
(1857) ; Haussk. in Flora, lvi. 404 (1873); Pugsley, Fum. in Brit. 45
(1912). F. vulgaris, Bubani, Fl. Pyr. iii. 278 (1901).
F. purpurea et F. flore albo, Gerard, Herb. 927 (1597); F. vulgaris,
Parkinson, Theatr. Bot. 287 (1640).
Icones. Woodville, Med. Bot. ii. tab. 88 (floribus pallidis ad var. elegantem
accedens) ; Smith, Eng. Bot. 589 ; Curtis, Fl. Lond. i. tab. 147 (fasc. ii. 52)
(forma agrestis, Haussk.) ; Wagner, Pharm.-Med.-Bot. tab. 59 ; Fl. Danica,
tab. 940 (f. floribunda, Pet. ex Haussk.); Hayne, Gewiichse, v. tab. 4;
Sturm, Deutschl. Fl. i. 62, tab. 14, ut F. officinalis var. major ; Svensk Bot.
i. tab. 42 : Reichb. Icones Fl. Germ. iii. tab. 3, fig. 4454, cum var, scandente ;
Hamm. l.c. tab. i; Clavaud, Fl. Gironde, pl. 4, fig. 1.
As pointed out in the general remarks on the subsection Murales, F. media,
Loiseleur, ‘Notice’ p. 101 (1810), is closely related to this species. Loiseleur
separates it from X. officinalis by its more rampant habit, ample foliage, and
larger, paler flowers ; and it seems best regarded as a large-flowered state of
the plant figured by Reichenbach, without diagnosis, as F. officinalis var.
scandens, which was reduced to a forma scandens in ** Fumaria in Britain,”
p. 51. It is perhaps also the a. vulgaris of Koch, Syn. Fl. Germ. ed. 2, App.
p. 1017 (1845), and of Hammar (Mon. p. 10), but it is not clear whether
these authors intend to distinguish under this name a variety separable from
the specific type.
GENERA FUMARIA AND RUPICAPNOS. 303
! Racemi quam in typo breviores, relative pauciflori ; sepala minora.
B. minor, Koch in Sturm Deutschl. Fl. i. 62, No. 14 (1833) ; Syn. Fl.
Germ. ed. 2, App. 1018 (1845); Hamm. l.c. 10 (1857); Haussk.
l.c. 419 (1873) ; Pugsley,/. c. 52 (1912).
Icon, Sturm, Deutschl. Fl. i. 62, tab. 14.
Hwsicc. Orphanides, Fl. Greca, No. 519, ut F. officinalis var. laxiflora !
Glauca, tenella, diffusa et sspe ramosissima. Racemi lawiusculi, 10—20-
Hori. Kepala parva, circa 2 mm. longa; corolla quam typi paulo minor
pallidiorque. Fructtis ut in typo latiores quam longi, 2 mm. Jongi et 2:5 mm.
lati, retusi.
The comparatively short and few-flowered racemes and the small sepals
characteristic of this variety were first accurately diagnosed by Haussknecht,
but the former feature and the relatively small, pale flowers seem sufticiently
shown in Koch’s original figure.
In Herb. Mus. Brit. there are Crimean specimens from Herb. Pallas
seemingly referable to this variety labelled * In rupibus Taurie ubique."
It may be inferred from this that the plant grows in natural stations in that
country, and this appears to be confirmed in M. Bieberstein’s * Fl. Taurico-
Caucasica,’ ii. 147 (1808), where it is observed, under F. officinalis, * Habitat
in ruderatis et arvis, varietas minor in saxosis."
It is probable that F. Meyeri, Steven, Enum. Pl. Phaner. in Taurià sponte
crescentium in Bull. Soc. Bot. Mose. xxix. i. 288 (1856), is identical with
this variety.
y. Wirtgenii, Haussk. l.c. 411 & 420 (1873) ; Pugsley, l.c. 52 (1912).
F. Wirtgenit, Koch, Syn. Fl. Germ. ed. 2, App. 1018 (1845).
Icon, Pugsley, l.c. fig. 6 (Journ. Bot. l. tab. 519).
Exsice. Dörter, Herb. Norm. No. 4601, ut F. Wirtgenii !
Habitu foliisque variabilis, sed ssepe typo gracilior foliorum laciniis paulo
angustioribus. acemi laæiusculi, vulgo 10—20-/fori ; pedicelli quam in typo
breviores et crassiores, rarius plus minusve elongati. Sepala parva, circa
2 mm. longa et 1 mm. lata, ssepius albida et vix acuminata. Corolla quam
typi paulo minor pallidiorque, petalorum ezteriorum alis apice valde dilatatis
ut petala superius emarginatum et inferius truncato-spathulatum fiant.
Fructüs 2—2:5 mm. longi et viw latiores, rotundato-truncati sed haud retusi,
seepissime apiculo parvo persistente przediti, circa medium latissimi et inferne
in stipitem quam in typo minus obscurum angustati, siccitate apicis foveolis
obscuris plane rugosi.
The exsiecata, Billot No. 1603 (F. Wirtgenit, Koch), as represented in
Herb. Boissier, clearly belongs to this variety and is not identical with the
example similarly labelled in Herb. Mus. Brit. which is referred to var.
elegans in “ Fumaria in Britain."
204 MR. H. W. PUGSLEY : A REVISÍON OF THE
!! Racemi longi, multiflori ; foliorum lacini quam in typo angustiores.
ò. densiflora, Parlatore, Mon. Fum. 53 (1844); Haussk. l. e. 421
(1813). F. densiflora, DC. Cat. Hort. Monsp. 113 (1813), et Prodr.
Syst. Nat. i. 130 (1824), ex parte ; F. officinalis var. floribunda, Koch,
Syn. Fl. Germ.ed.2, App. 1018(1845), ex parte; Hamm. l. c. 10 (1857);
F. oficinalis var. pycnantha, Loret & Barr. Fl. Montp. ed. 1, 32
(1876) ; F. parviflora, Sibth. & Smith, Fl. Græc. Prodr. ii. 50 (1813),
et Fl. Græc. vii. 63 (1830) ? non Lamarck.
Icon. Ñibth. & Smith, Fl. Greee. vii. tab. 668, ut F. parviflora.
Evsiec. Bourgeau, Pyr.-Espag. No. 394, ut F. officinalis ! Welwitsch, Fl.
Lusit. Nos. 136 & 764, ut F. officinalis var. parviflora !
The F. parviflora of Sibthorp & Smith was referred by Hammar to
F. micrantha, to which the description, at least as to its subglobose fruits,
may be held to apply. The figure, however, has evidently been drawn from
an example of this variety of F. officinalis, which it very well portrays, except
for the obscurely shown fruits. Both the corolla and the sepals are widely
different from those of any form of F. micrantha.
It is also probable, though not equally certain, that this variety is the
F. officinalis var. densiflora of Moris's * Flora Sardoa,’ i. p. 90 (1837).
e. elegans, Pugsley, l. c. 52 (1912). F. media, Reichb. Icon. Fl.
Germ. iii. 1 (1838), nee Lois. nec aliorum.
leon. Reichb. l. c. tab. 2, fig. 4453, ut F. media.
Exsice. Billot; Fl. G. & G. No. 214, ut F. officinalis! Fiori & Béguinot,
FI. Exsiec. Ital. ii. No. 1050, ut F. officinalis !
This plant is well marked by its ample glaucous foliage with narrowly
lanceolate segments, by its long racemes of usually pale flowers, and by its
small fruits borne on long and slender pedicels subtended by relatively short
bracts. These features do not agree with those of Loiseleur’s F. media, but
are emphasized by Reichenbach, either in his plate or his diagnosis, where
the plant is referred to as a “species constantissima.”
Continental examples occasionally show finer flowers than those from
which the description in “ Fumaria in Britain ” was taken, and present some-
what the appearance of a rampant state of var. densiflora.
Hammar cites Reichenbach’s figure of F. media for his var. scandens, but
his description depicts a different plant with broad leaf-segments and large
fruits, whose affinity may be held uncertain.
The variety albiflora, Parlatore, Mon. p. 53, with which the author identifies
F. Gasparini, Bab. in Trans. Bot. Soc. Edin. i. 36 (1844), is referred by
Hammar to var. minor. No material of this species has been seen, however,
with the white flowers by which var. albiflora is distinguished, and it seems
probable that this character was taken from a shade-form only.
Of the var. banatia, Haussk. l.e. 421 (1873), no authentic material has
been examined, and its identity with Reichenbach’s var. scandens seems to
require further investigation.
GENERA FUMARIA AND RUPICAPNOS. 305
The varying states of this fumitory, as it is seen in cornfields in spring,
in gardens in summer, and when shade-grown, are carefully distinguished by
Haussknecht under the three form-names, floribunda, Peterman, agrestis, and
umbrosa ; and the parallel variations of the variety Würtgeni? are similarly
dealt with, as also those of the species F. Schleicheri, Soy-Will., F. Vaillantü,
Lois., and F. parviflora, Lam. These variations, which Haussknecht remarked
in the species familiar to him, probably occur generally throughout the genus
where the climatic conditions are favourable for the continued propagation
and growth of the plants throughout the summer.
The distribution of F. officinalis, as given by Hammar, is amended by
Haussknecht and in * Fumaria in Britain.” It is pre-eminently the common
European Fumitory, but both in Asia and North Africa it is scarcer and less
widely spread than F. parviflora.
Haussknecht further enumerates a number of widely scattered localities
for the varieties minor, Wirtgenii, and densiflora ; and it may be added that
var. elegans occurs not only in Britain, France, and Germany, but also in
Italy and Spain.
A hybrid of the variety Wirtgenii with F. Vaillantii var. Chavinit has
been described from the Var in Southern France as x F. Albertii in Rouy &
Foucaud's Fl. de France, i. p. 178 (1893) ; and in the same year another
description of F. oficinalis x Vaillantii (F. abortiva) from Thessaly was pub-
lished by Haussknecht in Mittheil Thür. Bot. Ver. N. Folge, iii. & iv.
p. 102 (Symbolæ ad floram græcam). There is an authentic example of this
latter plant in Herb. Kew. F. officinalis x parviflora and F. officinalis x
micrantha are veferred to at page 50 of * Fumaria in Britain.”
34. FUMARIA CrLICICA, Haussk. in Flora, lvi. 423 (1873). F. officinalis y.
tropidocarpa, Buser in Fl. Orient. Suppl. 26 (1888).
Exsice. Kotschy, Iter Cilicico-Kurdieum, 1859, No. 42, ut F. officinalis
var. floribunda! Bourgeau, No. 32, Trebizond, 1862, in Hb. Boissier, ut
F. officinalis! Balansa, Vallée de Djimil, Lazistan, 1866, in Hb. Boissier, ut
F. officinalis! Zorab, No. 417, Erzerum, ut /. officinalis! Darbes, Syria
Libanotica, 1881, in Hb. Zürich, ut F. oficinalis! Pl. Auraniticee, Postian
ap. Coll. Syriens. Prot., El Kafr, 1886, ut F. officinalis var. tropidocarpa !
Sintenis, Iter Orient., 1894, No. 5495, Armenia Turcica !
In addition to authentic material of Kotschy’s plant on which this species
was founded, the herbaria at Kew and the British Museum contain excellent
Armenian examples collected by Sintenis, which are labelled as referred by
Haussknecht to F. cilicica.
In these examples the characters of this remarkably floriferous and
somewhat showy fumitory are well seen, and they generally confirm the
accuracy of Hausskneckt’s unusually full diagnosis.
W c iius
306 . MR. H. W. PUGSLEY : A REVISION OF THE
After an examination of the exsiccata cited it may be remarked that,
compared with 7. officinalis, F. cilicica seems chiefly notable for its very long
and dense racemes, bearing from 30 up to 80 flowers and much exceeding the
short and thick peduncles. Its bracts, also, are more subulate and more
finely pointed, while the sepals are relatively small and narrow and, at least
in some cases, acuminate rather than mucronate. The corolla is rather larger
than in F. officinalis (8-9 mm. long), with broad, short, obscurely purple
wings to the upper petal much exceeding the keel and extending to its apex;
the spur ascending, longer and more curved than in F. officinalis ; and the
lower petal less spathulate, and subacute with spreading margins scarcely
reaching the apex. The apical pits of the fruit are roundish and shallow,
and the persistent apiculus, while apparently invariably present, is sometimes
very short, giving the fruit a strong resemblance to that of F. officinalis.
On this ground Buser seems to have reduced this plant to a variety of that
species.
It will be seen from the exsiccata cited that F. cilicica has a wide distribu-
tion in Asiatic Turkey.
35. Fumaria Boissier, Haussk. in Flora, lvi. 424 (1873); Buser in Fl.
Orient. Suppl. 27 (1888).
Eusice, Haussk. Terek Mesopotamiæ borealis, 1867, in Hb. Boissier !
This species, described by Haussknecht from plants which he himself
collected in Northern Mesopotamia, seems from the exsiccata in Herb.
Boissier—the only material examined—to be separable with difficulty from
F. cilicica.
In the Boissier specimen, which shows good foliage and flowers though only
immature fruit, the leaves, racemes, pedicels, bracts, and sepals appear to be
essentially those of F. cilicica; and although the outer petals are more
broadly winged and the apiculus of the young fruit is longer than in the
Kotschy gathering which forms the type of F. cilicica, yet it is clear from
the more recent material referred to this species that these features are liable
to an appreciable range of variation.
Although Buser retains F. Boissieri as a species while he reduces A. cilicica
to a variety of F. officinalis, it is apparent that he recognizes the very close
affinity of the three plants.
36. FUMARIA microstacuys, Kralik ex Haussk. in Flora, lvi. 552 (1873).
F. judaica B? microstachys, Buser in Fl. Orient. Suppl. 29 (1888).
Eesice. Kralik, Alexandrie, Egypt, 1847, in Hb. Boissier! et Hb. Kew!
Gaillardot, Pl. d'Egypte, No. 437, Alexandrie, 1870, ut F. Vaillant ?!
Schweinfurth, Basse-Egypte, No. 267, Alexandria, 1890, ut F. Judaica!
Fumaria, ut videtur, annua, satis robusta, diffusa, tandem ramosissima.
Folia plus minusve glauca, foliolis in lacinias lineari-oblongas, planas, acutas vel
obtusas raro mucronatas fissis 3-4-pinnatisecta. Racemi ia«iusculi, pauciffori
GENERA FUMARÍA AND RUPICAPNOS., 307
(6-12-flori), primarii pedunculos subequantes, seriores pedunculos brevissimos
multo superantes. Bractew subulato-lanceolate, breviter acuminate, pedicellos
fructiferos erecto-patentes brevissimos (2 mm. longos) apice valde incrassatos
sube quantes vel superantes. Sepala circa 15-2 mm. longa, *15 mm. lata,
sepius ovata, acuminata, dentata, albida. Corolla 5-7 mm. longa, pallide rosea ;
petalorum exteriorum carinis abrupte terminatis, alis atropurpureis mar-
ginibusque roseis in exemplaribus visis haud bene explicatis; petalis
interioribus apice atropurpureis. —ructüs mawimi, 3 mm. longi et paululum
latiores, brevissime obcordati, retusi, supra medium latissimi et inferne in
stipitem obscurum multo angustati, parum compressi sed conspicue carinati,
in sicco apicis foveolis latis obscuris rugos.
Hee Fumaria distinetissima propter fruetüs maximos cum F. judaieá vulgo
confusa est. In exemplaribus visis ores haud satis explicantur, sed corollas
formam sectionis Parviflorarum exhibere manifestum est.
F. judaica foliorum laciniis majoribus latioribus, pedunculis pedicellisque
longioribus, sepalis angustioribus, corollis albidis multo majoribus eum petalo
inferiore haud spathulato, fructibus paulo minoribus, obscurius carinatis,
magis tuberculato-rugosis longe distat.
F. microstachys in /Egypto solo (prope Alexandriam) hactenus inventa est.
This remarkable fumitory, of whose position in the genus Haussknecht
expressed some doubt owing to lack of adequate material, and of whieh
good flowering specimens are still wanting in British herbaria, is unique
among the Purviflome on account of its very large fruits. In habit it
evidently approaches F. officinalis and F. Vaillantii, which in some degree
it probably resembles also in flower, but its fruits recall F. macrocarpa and
F. judaica of ihe Grandiflore, and it thus seems a somewhat anomalous
intermediate between these widely different species. As a member of the
Parviflore it seems best placed in the subsection Officinales on account of
the moderate size of its sepals and the broad form of its fruits.
Sussgcrio VI. MICROSEPALZE.
Microsepale, Pugsley, Fum. in Brit. 59 (1912). Officinales, Hamm.
Mon. 9 (1857), ut sectio, ex parte; Parviffore, Haussk. in Flora, lvi. 441(1873).
Pedunculi plus minusve breves, rarius obsolescentes (F. australi excepta),
pedicellis incrassatis tenuibusve præditi. Braetez variabiles. Sepala minima
(nisi in F. Janke et F. australi) vel etiam absentia, sepissime insigniter
dentata, vulgo corolle quartá parte breviora ejusque tubo sepissime
multo angustiora. Fructüs parvi vel modici, rarissime conspicue lati, siccitate
rugosi vel rugulosi.
The blackish-red wings of the upper petal so generally characteristic of the
genus are well marked in the deep rose corollas of F. Schleicheri and F. Janke,
and in the lighter flowers of F. microcarpa. They become less marked in
308 MR. H. W. PUGSLEY : A REVISION OF THE
F. abyssinica, and are generally obscure in F. indica and F. Schrammii,
while in F. Vaillantii the dark colouring is fairly distinct in some forms
and quite absent in others.
In F. parviflora the normally white corolla usually shows a small external
bloteh only of purple on the wings of the upper petal, recalling the coloration
of F. occidentalis; and in F. asepala the purple tinting, if present at all, is
confined to the tips of the inner petals.
* Series Ambigua.
Flores rosei, 5-7 mm. longi; sepala 1:5-2 mm. longa; petali superioris
ale sursum reflexe vel obsolescentes deflexæ.
37. Fumaria AbYssINICA, Hamm. Mon. 19 (1857); Haussk. in Flora, lvi.
492 (1873).
Jeon. Hamm. l.c. tab. vi.
Egsice. Schimper, Iter Abyss., sec. secunda, 1842, No. 1347, Demerki, ut
F. officinalis! Schimper, Abyss. 1863, No. 1429 (shade-form) ! Ankober,
1841, in Hb. Kew! Fl. Colon. Eritrza, 1902, No. 207, in Hb. Mus. Brit. !
Schweinfurth, 1889, No. 1675, El Ejan am Schibam (2700 m.), Arabia Felix,
in Hb. Kew !
There is good material in Herb. Mus. Brit. of. the plant on which this
species was founded. As pointed out by Haussknecht (/.c.), the fruit is
scarcely obtuse, as stated in Hammar’s diagnosis, but rather mucronulate
when young, and at maturity subacute and very shortly apiculate, much as
in some forms of F. parviflora, though the keel is less marked than in that
species. In the Eritrzan exsiccata a somewhat more rounded-obtuse, though
still apiculate form prevails. The fruit is 2-2:25 mm. long and about
equally broad, with very shallow apical pits.
It may further be remarked that the leaf-segments in F. abyssinica are
more generally linear-oblong than oblong-lanceolate ; and the racemes, which
are usually sub-12-flowered in Schimper's examples but show as many as 20
flowers in the Eritrean exsiccata, are much longer in fruit than the very
short peduncles, except occasionally when shade-grown. The bracts seem
to be relatively broad—linear-oblong rather than linear—and fully as long
as the short (2 mm. long), suberect pedicels. The ovate, acuminate sepals
are about 1°5 mm. long and 1 mm. broad ; the purple wings of the obtuse
upper petal are reflexed upwards as in X. officinalis or in F. Sehleicheri, but
are normally rather narrow and scarcely cover the keel ; and the lower petal
appears less distinctly spathulate than in most of the other species of the
subsection.
In the Arabian specimen at Kew the wings of the outer petals are but
little developed, and the obscurely apiculate fruit is more rounded-obtuse
than in the Abyssinian type.
GENERA FUMARIA AND RUPICAPNOS. 309
The range of F. abyssinica is now known to extend from Abyssinia to the
Italian coast colony of Eritrea and across the Red Sea to Arabia Felix.
There is also an immature specimen from Uganda in Herb. Mus. Brit.
(Bagshawe, Ruchiga, No. 396 !) which seems referable here. It appears to
occur only at a considerable altitude. F. abyssinica is further recorded on
Haussknecht’s authority for Gabes in Murbeck's ‘Contributions Fl. du Nord-
Ouest de Afrique,’ p. 5 (1897).
The material collected in the Kilimandjaro district of East Africa as
F. abyssinica is referred to a distinct species F. australis.
38. FUMARIA AUSTRALIS, sp. nov. (Pl. 13.)
Exsice. Volkens, Fl. des Kilimandscharo, 1893, Nos. 953 & 1333, Marangu
(2800 m.) et Fuss des Kifinnka (2700 m.), ut F. abyssinica! Whyte,
Nandi District, Tropical E. Africa (2000 m.), 1898, in Hb. Kew!
Fumaria verisimiliter annua, habitu satis robusto, internodis longis
elongata, petiolis cirrhosis scandens. Folia glauca, foliolis in lacinias planas
lineari-lanceolatas vel lanceolatas, acutas, longe mucronatas vel aristatas
fissis 2-3-pinnatisecta, Racemi 12-20-flori, floriferi conspicue densi, etiam
fructiferi pedunculis mediocribus paulo breviores, rarius eos subsequantes.
Bractee magne et conspicue late, oblongs, cuspidate, sparsim serrulatw,
pedicellis fructiferis brevissimis (circa 1:5 mm. longis), crassis, erecto-patentibus
subduplo longiores. Sepala circa 2 mm. longa, 1 mm. lata, oblongo-ovata,
acuta, irregulariter dentata, rosea. Corolla 5—7 mm. longa, rosea ; petalo supe-
riore alis roseis angustissimis fere obsoletis sepissime deflexis apicem haud
attingentibus acuto, calcare brevi parum curvato ; petalo inferiore marginibus
patentibus apicem vix attingentibus subacuto subspathulato ; petalis interiori-
bus apice atropurpureis. Fructûs modici, circa 2:25 mm. longi ac lati,
subrotundi, superne cum apiculo brevissimo persistente obtusiusculi, inferne
in stipitem latum parum angustati, paululum compressi sed plane carinati, in
sicco apicis foveolis obscurissimis dense rugosi.
Hee Fumaria, que ad F. abyssinicam relata est, habitu laxiore scandente,
foliorum laciniis fere aristatis nonnunquam latioribus, racemis densis longius
pedunculatis, bracteis majoribus, et preesertim corollis acutis petalique
superioris alis angustissimis deflexis nec sursum reflexis satis differt.
Ab hujus subsectionis speciebus aliis per corolle formam subexalatam
F. australis :equaliter distat. Petali superioris ale fere obsolete eis
F. coccinec similes sed specierum duarum characteres reliqui omnino diversi
sunt.
F. australis in regionibus Kilimandjaro et Nandi Africe orientalis crescit,
The exsiceata collected by Volkens, though elongate, show highly coloured
and what seem to be normally developed flowers, and appear to be the usual
form of a southern endemic species.
310 MR. H. W. PUGSLEY : A REVISION OF THE
39. FUMARIA JANK E, Haussk. in Flora, lvi. 491 (1873).
Exsice. Janka, Szekelyhid com. Bihar Hungaris, 1861, in Hb. Boiss. !
The only material examined of this rare Hungarian plant is the specimen
from Janka in Boissier’s Herbarium. This shows poor flowers, but has
the appearance of a distinct species agreeing with Haussknecht’s diagnosis
except in the length of the pedicels, which seem distinctly shorter than those
of F. Schleicheri.
F, Janke is evidently somewhat intermediate between F. rostellata and
F. Schleicheri, and is retained in this subsection in accordance with the views
of Haussknecht.
** Series Eu-Microsepala.
Flores rosei vel albidi, raro plus 6 mm. longi; sepala minuta (5-15 mm.
longa) vel absentia; petali superioris ale sursum reflex: erecto-patentes
vel patule.
40. FUMARIA ScCHLEICHERI, Soyer- Willemet, Observ. Pl. France, 17 (1828) ;
Haussk. in Flora, lvi. 411 &485 (1873). F. Vaillantii, subsp. Schleicheri,
Rouy & Foucaud, Fl. Fr. i. 181 (1893) ; F. Laggeri, Jordan, Pugillus,
7 (1852) ; F. tenuiflora, Fries Herb. ex Hamm. Mon. 13 (forma
umbrosa), et F. Vaillantii B. Laggeri, Hamm. Mon. 15 (1857):
F. acrocarpa, Peterman (nomen ?).
Icon. Hamm. l.c. tab. i, ut F. Wirtgenii (sic).
Exsice. Schultz, Herb. Norm. No. 2109! Borbas, Fl. Exsice. Austro-
Hungarica, No. 2903 IT!
Fumaria annua, gracilescens, satis ramosa, suberecta vel diffusa, rarius
petiolis cirrhosis scandens. Folia plus minusve glauca, foliolis in lacinias
planas lineari-oblongas vel lineari-lanceolatas, acutas rarius mueronatas fissis
2-(rarius 3-) pinnatisecta. Racemi floriferi densiusculi, swpissime 12—20-,
rarius plus 20—flori, fructiferi laxiores, primarii pedunculos graciles subaquantes
seriores plane superantes. Bractew lanceolato-subulaize, acuminate, pedicellis
fructiferis suberectis gracilibus (4 mm. longis) apice ipso incrassatis fere triplo
breviores. Sepala circa 1 mm. longa,5— 15 mm. lata, ovata, acuminata, irregula-
riter inciso-dentata,rosea,corollz tubo plane angustiora. Corolla5—6mm. longa,
saturate rosea (in sicco sub-violacea) ; petalum superius obtusum alis atropur-
pureis sursum reflexis carinam viridem excedentibus apicemque attingentibus
et egleare longo curvato preditum ; petalum inferius marginibus patulis roseis
apicem vix attingentibus obtusiusculum spathulatum; petala interiora apice
atropurpurea fere recta. Fructûs modici, circa 2 mm. longi ac lati, subrolundi,
superne cum apiculo brevi persistente rotundato-obtusi (rarissime sine apiculo),
inferne parum angustati, paulo compressi et plane carinati, siccitate apicis
foveolis parvis paulo obscuris rugosi.
B. supina, var. nov.
F. supina, Janka in Termesz Füzetek, i. 30 (1877).
Egsicc. Janka, Noszoly et Feketelak, Transsilvaniz centralis, 1876, in
GENERA FUMARIA AND RUPICAPNOS. 311
Hb. Kew! Szépligeti, Fl. Hungarica, Budapest, Schwabenberg, 1888, in
Hb. Mus. Brit., ut F. Schleicheri !
Typo simillima, sed petalo superiore alis latioribus carinam conspicue
superantibus obtusissimo, petalo inferiore marginibus latioribus apice obovato,
fructibus paululum minoribus vix 2 mm. longis ac latis leviter rugosis.
F. Schleicheri foliorum laciniis latioribus, racemis longius pedunculatis
pedicellis longioribus gracilibus breviter bracteatis, floribus saturate roseis,
sepalis majoribus, petali superioris haud emarginati alis atropurpureis reflexis
nee patulis a F. Vaillantii et a F. Schrammii differt.
F. officinalis habitu robustiore, bracteis longioribus, floribus sepalisque
majoribus, fructibus latioribus retusis truncatisve facile distinguitur.
This pretty species, which is normally a very distinct plant, characterized
especially by its deeply coloured flowers and its apiculate fruits borne on long,
slender, and shortly bracteated pedicels, has been the subject of much
confusion since it was first distinguished by Soyer- Willemet. Hammar, who
overlooked Soyer-Willemet/s description, seems to have been but slightly
acquainted with it; and he failed to recognize the identity of Peterman’s
shade-grown F. aerocarpa from Leipzig, which he described as F. tenuiflora,
Fries, with the Swiss plants usually distributed by Lagger as F. Laggert,
Jordan, and reduced in the Monograph to a variety of F. Vaillantii.
It was not till Haussknecht dealt with these plants that Soyer-Willemet’s
name was re-established and the essential features of F. ScAleicheri satisfac-
torily diagnosed in a tabular form in which they are contrasted in detail
with F. officinalis, F. Wirtgenii, and F. Vaillantii—the species with which
F. Sehleicheri had been previously hopelessly confused. This elucidation is
a careful and accurate piece of work, clearly showing Haussknecht’s intimate
knowledge of these plants, but his identification of F. Laggeri, Jord. with
F. Chavinii, Reuter, rather than with F. Schleicheri, cannot be accepted, as
explained at page 68 of “ Fumaria in Britain." Haussknecht’s determination
was perhaps influenced by the fact that in Herb. Boissier Lagger's specimen
of F. Laggeri is F. Chavinii.
Even in quite recent years other species have occasionally been mistaken
for F. Schleicheri, as in the exsiecata Fl. Austro-Hungarica, No. 2903 I,
where F. Schrammii has been sent out under this name.
According to Haussknecht, authentic material of F. carinata, Schur,
Enum. Pl. Transsilv. p. 38 (1866), the identity of which is indeterminable
from the author’s description, belongs to this species, but it is referred to
F. Vaillantii in Simonkai’s Transylvanian Flora.
The distribution of F. Schleicheri is shown by Haussknecht to extend from
Montpellier and Nice, and from the mountains of Savoy across Central
Europe and Russia to the Caucasus and the Altai District of Central Asia.
It is also recorded for the Spanish province of Valencia (cf. Willkomm,
312 MR. H. W. PUGSLEY : A REVISION OF THE
Fl. Hisp. Suppl. 311 (1893) ). It is one of the hardiest species of the genus,
growing at an altitude of nearly 6000 feet in the Pennine Alps; and there
are specimens in Herb. Kew from Siberia Altaica (Ledebour & Ludwig, as
F. Vaillantüi !).
Janka's F. supina has been reduced to a variety of this species as the
authentic specimen from Janka at Kew appears to show all the essential
features of F. Schleichert and scarcely to differ except in its broadly winged
corolla. It is reduced to a synonym of F. Schleichert by Simonkai.
F. Schleicheri x F. Vaillantii var. Chavinit.
Exsicc. Brunies, Zernez, Ofenpass Gruppe, Graubünden, Switzerland,
1903, in Hb. Zurich, ut F. Vaillantii var. Laggeri !
Planta robusta, elongata ; folia laciniis paululum latioribus quam in
F. Vaillantii var. Chavinii preedita. Racemi 20-30-flori, longi, laxiusculi,
peduneulos plane superantes. Bracteæ lineari-oblonge, acuminate, pedi-
cellorum gracilium suberectorum dimidium paulo excedentes. Sepala 1 mm.
longa, ovata, longe acuminata ; corolla cirea 6 mm. longa, læte rosea, petali
superioris alis purpureis et ut in F. Schleicheri sursum reflexis. Ovaria
apiculata ; fruetüs omnino abortivi.
Exemplaria Fumariw Schleicheri et F. Vaillantii var. Chavinii in hae
regione collecta in herbario Zurich sunt.
41. FUMARIA microcarpa, Boissier ex Haussk. in Flora, lvi. 490 (1873).
F. Schleicheri B. microcarpa, Buser in Fl. Orient. Suppl. 27 (1888).
Exsice. Huet du Pavillon, Erzerum, 1853, in Hb. Boiss., ut F. parviflora
var. fl. roseis! Maunsell, Van, Asiatic Turkey, 1899-1900, in Hb. Mus. Brit.!
Becker, Pl. Astrachanice, Sarepta, 1879, in Hb. Mus. Brit., ut F. Vaillantii!
This plant, diagnosed as a species with some doubt by Haussknecht from
Huet du Pavillon's specimen in Herb. Boissier, is clearly very closely allied
to F. Schleicher’, as Haussknecht remarks. The resemblance is seen in the
foliage, in the dark-tipped flowers with the wings of the upper petal reflexed
upwards, and in the sub-rotund, apiculate fruits. But F. microcarpa differs
in its dwarfer and more erect habit, its much shorter pedicels with relatively
long braets, its smaller and less deeply coloured flowers with more curved
inner petals, and its smaller fruits (about 1:75 mm. long and broad).
In some degree F. microcarpa may be regarded as intermediate between
F. Schleicheri and F. Schrammii, the latter of which it resembles in its
small, apiculate fruit. Its corolla, however, quite laeks the marked dorsal
compression, with spreading wings, which characterizes the flowers of
F. Schrammii and F. Vaillantüi.
'The more recent material collected by Maunsell and Becker, and now at the
British Museum, appears to be conspecific with the type of F. microcarpa
in Herb. Boissier, although these later plants show shorter peduncles and
GENERA FUMARIA AND RUPICAPNOS, 313
the Sarepta examples somewhat longer pedicels and larger corollas (up to
5°5 mm. long).
It may thus be concluded apparently that F. microcarpa is a local species,
more or less generally distributed from Armenia northwards to the basin of
the lower Volga.
The exsiecata labelled “F. microcarpa, Boiss., Huet du Pavillon, Erzerum,
1853,” in Herb. Mus. Brit. (a fragment) and in Herb. Kew are not identical
with the type in Herb. Boissier and are referable to F. Schrammii and
F. Vaillanti respectively.
Other Asiatic specimens annotated as similarly named by Haussknecht
(Bornmiiller, Pl. Anatol. Orientalis, 1890, No. 1877, Amasia!) appear to
belong to F. Vaillantii var. conferta, or (Dong uM Pl. Lydi: et Cariæ, No.
9015, Smyrna !) to F. Schrammii.
42. FUMARIA INDICA, sp. nov.
F. Vaillantit var. indica, Haussk. in Flora, lvi. 443 (1873); F. parviflora,
Wight & Arnott, Prod. Fl. Penins. Ind.-Orient. i. 18 (1834),non Lamk. ;
F. parviflora var. Vaillantii, Hooker fil. & Thomson, Fl. Indica, i. 258
(1855) ; F. parviflora subsp. Vaillantii, Hooker fil. Fl. British India, i.
128 (1872).
Icon. Wight, Illustr. Ind. Bot. i. tab. 11, ut F. parviflora.
Ezxsicc. Hohenacker, Pl. Ind. Or. No. 1488, ut F. parviflora! Hooker,
No. 206, Behar, ut F. Vaillantii (f. umbrosa)! Clarke, Nos. 26864
& 26903, Pubna & Beanleah, Bengal, ut F. parviflora ! Watt, Fl. N.W.
Himalayas, No. 852, Pangi, ut F. parviflora ! Watt, Fl. Gangetic Plain,
No. 2156, Bankipore, ut F. parviflora !
Fumaria annua, sepius caulibus crassis habitu robusto vel elongato, suberecta
vel diffusa, verisimiliter haud scandens. Folia vulgo ampla, fwniculoidea,
plus minusve glauca, foliolis in lacinias longas lineares vel lineari-oblongas
(raro paulo latiores in formis umbrosis) confertas planas acutas fissis 2—4-
pinnatisecta. Racemi 15-25-flori, floriferi densiusculi, fructiferi elongati, laxi,
pedunculos crassos seepissime subdi superantes, rarius subsessiles. Bractew
lanceolato-subulat:e, tenuiter acuminate, pedicellos (vulgo 2-2:5 mm. longos,
rarius ad 4:5 mm. elongatos) erecto-patentes apice multo incrassatos subaquantes
vel inferiores nonnunquam longiores. Sepala circa 1:5 mm. longa, *50—1 mm.
lata, lanceolata vel ovato-lanceolata, acuminata, plus minusve inciso-dentata,
rosea, in fructu juniore szepe persistentia. Corolla 5-6 mm. longa, rosea ; petalo
superiore haud lato alis roseis (rarius purpurascentibus) plus minusve sursum
reflexis carinam superantibus apicemque attingentibus obtuso, et calcare longo
curvato non adscendente przdito ; petalo inferiore marginibus patulis
apicemque attingentibus obtuso spathulato, ssepe libero deflexo; petalis inte-
rioribus apice atropurpureis sursum curvatis. Fructús mediocres, 22:25 mm.
longi, 2:25-2:5 mm. lati, subrotundo-quadrati, subtruncati et CEP obscure
retusi (in statu juniore cum apiculo brevi qui rarissime persistens est),
LINN. JOURN.— BOTANY, VOL. XLIV. 20
314 MR. H. W. PUGSLEY : A REVISION OF THE
inferne abrupte angustati, paululum compressi sed valde (priesertim apicem
versus) carinati, in sicco apicis foveolis obscuris rugosi.
Forma montana.
Exsicc, Wallich, No. 14368, Kamaon! Collett, No. 457, Simla, ut
F. parviflora! Aitchison, No. 216, Afghanistan !
Planta humilior, compacta; foliorum minorum laciniis brevioribus
erassioribus subeanalieulatis ; bracteis pedicellos breviores sepius multo
superantibus.
Fumaria indica per folia vulgo magis decomposita laciniis longioribus
predita, per racemos longiores prope constanter pedunculatos, per sepala
plerumque majora, per corollam angustiorem plane curvatam, minus dorso
compressam, petali superioris alis sursum reflexis potius quam patentibus, et
per fruetüs subtruncatos valde catinatos sepissime majores a F. Vaillantii,
a F. Schrammii, et a F. parviflord differt.
Preterea F. Vaillantii habitu graciliore, foliorum laciniis latioribus, petalo
superiore emarginato eum calcare adscendente distinguitur ; F. Schrammii
habitu gracili, racemis paucifloris, floribus petalo superiore emarginato
pallidis, fructibus rugulosis apiculatis insuper separanda est ; et F. parviflora
foliorum laciniis vulgo angustioribus canaliculatis, floribus albidis vel roseo-
tinctis nee vero roseis, petali superioris alis sepius externe purpureo-maculatis
caleareque semper adscendente etiam notata est.
F. Sehleicheri et F. microcarpa habitu graciliore, foliorum laciniis multo
latioribus relative sparsis, floribus constanter apice omnino atropurpureis,
fructibus minoribus rotundatis semper apiculatis longe distant.
F. indica in Indià extratropicd in planitie, in montibus subtropicis et in
peninsul: montibus Nilghiri occurrit, atque in cultis planta communis est ;
etiam in Seindià, in Afghanistan, in Kashmir, et sub montibus Himalaya in
Kumaon, in Napaliá et in Bhotan invenitur.
Ultra regiones Indis hee species in Beloochistan (Duthie, No. 8568,
Quetta, in Hb. Mus. Brit., ut 7. parviflora (abnormalis) !) et in Persia
(Kotschy, No. 133, Abu Schier, in Hb. Mus. Brit.! Polak, Iter Persicum,
1882, Kaswin, in Hb. Kew, ut F. Vaillantii! Kotschy, No. 45, prope
Teheran, 1843, in Hb. Kew solo, ut /. parviflora!) crescit; etiam ad
septentrionem in Turkestan (Regel, Iter Turkestanicum, Baldschuan, 1883,
in Hb. Mus. Brit., ut F. Vaillantii !), in Songariâ (Krassnow, FI. Iliensis,
1886, ut F. Vaillant, partim !), in Orenburg (Haussk.) et in Mongoliá
(Meyer, Pl. Mongoliz, No. 720, Chugutcehak !).
The distinetive appearance of this plant, which is now well represented at
Kew from numerous Índian localities, was remarked so long ago as 1834 by
Wight & Arnott (l.c. i. p. 18), who refer it to F. parviflora with the remark :
* In the Indian plant the flowers are rose-coloured and approach closely to
F. Vaillantii.^ Hammar places it under F. Vaillantii on account of its flat
leaf-segments and obtuse fruits, but notes that its leaf-cutting is much finer
GENERA FUMARIA AND RUPICAPNOS. 315
than in the type; and the same peculiar features are recognized by
Haussknecht in his diagnosis of F. Vaillantii var. indica. Like his
predecessor, however, Haussknecht does not seem to have remarked the
characteristic corolla or to have closely examined the fruits.
It is evident from their labels that some more recent Indian collectors have
hesitated what name to apply to this plant, and Aitchison, who found it a
common weed on the Afghan frontier, refers it to F. officinalis, seemingly
distinguishing it from F. Vaillantii, which he collected as a rare species in
the Kurrum Valley.
F. indica is remarkable for its great altitudinal range, extending from the
lowland Ganges basin to at least 9000 ft. in the N.W. Himalayas. As might
be expected, examples obtained in the warm regions differ considerably from
montane specimens, The form of the plains is usually a large, robust plant,
with long leaf-segments, giving it a fennel-like aspect, and frequently
elongate pedicels. The hill-plant is naturally dwarfer, and shows relatively
small leaves, with thicker and somewhat channelled segments. It also has
commonly very short and thick pedicels, much exceeded by the bracts, rather
finer flowers, and probably relatively larger sepals. As these differences are
such as would be expected to arise solely from the varied environment, the
mountain plant has been distinguished merely as a form.
43. FUMARIA VariLLANTII, Loiseleur in Desvaux, Journ. Bot. ii. 358 (1809),
et Notice, 102 (1810) ; Hamm. Mon. 14 (1857) ; Haussk. in Flora,
lvi. 411 & 441 (1873) ; Pugsley, Fum. in Brit. 66 (1912). F. Camerarii,
Bubani, Fl. Pyr. iii. 281 (1901).
Icones. Vaillant, Bot. Paris, tab. 10. fig. 6 ; Sturm, Deutschl. FI. i. 62,
tab. 15; Reichb. Icon. Fl. Germ. iii. tab. 1. fig. 4452; Hamm. l.c. tab. i
(f. sepalis latioribus) ; Clavaud, Fl. Gironde, Pl. 4. fig. 3.
Exsice. Billot, Fl. G. & G. No. 215 bis! Schultz, Herb. Norm. No. 414!
Fl. Exsiec. Carniol. No. 2871!
Hammar's diagnosis of this s; ecies is fairly satisfactory, but he does not
seem to have appreciated the sparse and long-petioled foliage characteristic
of the typical form, which was noticed by the older botanists Vaillant and
Loiseleur. It is possible that his description was partly based on examples
of the variety Chavinii, which, judging from British herbaria, appears to be
the prevalent if not the only form growing in Scandinavia.
In addition to its omission to emphasize the slender habit and bipinnatisect
leaves peculiar to typical F. Vaillantii, the Monograph is inaccurate respecting
the sepals, whieh are usually lanceolate instead of subrotund-ovate, and
frequently persistent on the young fruit rather than caducous. The flowers
of the typical form are normally of a purplish-pink colour; and the
ascending spur of the upper petal is a characteristic feature in this species, as
202
316 MR. H. W. PUGSLEY : A REVISION OF THE
in F. parviflora, which is not mentioned in the diagnoses in “ Fumaria in
Britain."
The mucronulus of the young fruit, which sometimes lasts almost to
maturity on the living plant, although invisible in the dry state, is also
inadvertently unnoticed in “ Fumaria in Britain.” Four forms, based
chiefly on the variations of the fruit, are distinguished by Clavaud
(Fl. Gironde, 51 (1882) ).
F. Vaillantii var. gracilis, Knaf in Flora, xxix. 292 (1846), is referred by
Haussknecht to a shade-form of the specific type.
B. conferta, Haussk. l. c. 445.
Exsice. Heldreich, Isbarte, 1854, ut F. Vaillant !
Herba robusta, compacta et satis ramosa ; folia laciniis obtusiusculis haud
sparsis 2-3-pinnatisecta. Racemi breves plerumque subsessiles ; pedicelli
haud graciles, brevissimi (circa 1 mm. longi). Corolla minima, 4-5 mm.
longa, pallide vosea, petalo superiore alis roseis plus minusve emarginato,
calcare ut typi curvato adscendente. Fructis minimi, vix 2 mm. longi et
latitudine paulo minores, subrotundi vel subrotundo-obovati, sine apiculo
obtusi. Aliter ut in typo.
The type of this variety is represented in Herb. Mus. Brit., and also at
Kew, where two good examples exist.
Other exsiceata, as “ Bornmüller, Pl. Anatol. Or. No. 1877," referred to
F. microcarpa, Boiss., apparently agree most nearly with this variety.
y. Chavinii, Rouy & Foucaud, Fl. Fr. i. 181 (1893) ; Pugsley, l.c. 70
(1912). F. Chavinii, Reuter, Cat. Pl. Genève, ed. 2, 10 (1861);
F. Vaillantii, Bab. in Eng. Bot. Suppl. 2877 (1844), partim (fl. roseis) ;
F. Vaillant B. Laggeri, Haussk. l. c. 442, nec Hamm. nec 7. Laggert,
Jord.
Icon. Eng. Bot. Suppl. 2877, partim (fl. roseis), ut F. Vaillantii.
Evsice. Billot, Fl. Exsice. Cont. No. 3508, ut F. Chavinii! Bourgeau, Pl.
d'Espagne, No. 2108, ut F. Vaillantii ! Schultz, Fl. G. & G. 2nd Cent. No. 6,
ut F. Vaillantüi !
Quam typus planta robustior et erectior, sepius parce ramosa et nunquam
intricato-ramosissima. Folia irregulariter 2—3-pinnatisecta, foliolis quam in
typo brevius petiolatis, laciniis lineari-oblongis linearibusve, acutis, confertis
potius quam sparsis. Racemi 10-20- (raro 25-) flori, pedunculos breves
superantes. Pedicelli quam typi plerumque longiores (3 mm. longi), flexuosi,
suberecti vel rarius erecto-patentes. Sepala quam in typo minus persistentia.
Corolla circa 6 mm. longa, dilute sed ssepius læte rosea, petali superioris valde
emarginati alis latissimis, sepe subserratis, roseis vel raro obscure purpureis.
Fructds quam typi sepe paululum majores, cirea 2°25 mm. longi et 2 mm.
lati, subrotundo-ovati, obtusi vel etiam subretusi, siccitate dense rugosi.
In this variety, which was feebly defined by Reuter but more accurately
diagnosed by Haussknecht and by Rouy & Foucaud, the flowers are usually
GENERA FUMARIA AND RUPICAPNUS. 317
light rose-coloured with very little, if any darker tinting in the wings of the
upper petal. These wings, as well as the margins of the lower petal, are
frequently more strongly developed than in the specific type. The fruit of
var. Chavinii, besides being often rather larger and more rugose than in
typical F. Vaillantii, differs further in being normally plainly longer than
broad.
There is good material of this variety at Kew from the locus classicus near
Geneva.
F. Vaillantiiy. ochroleuca, Knaf in Flora, xxix. 292 (1846), is probably, for
the most part, a pale-flowered form of this variety.
ô. venetica, var. nov.
Crsiec. Fiori, Béguinot, Pampanini, Fl. Exsice. Ital. No. 565, Venetia,
G. Rigo, ut F. Vaillantii!
Planta glauca, ut in var. Chavinii robusta, et satis ramosa. Folia ampla,
irregulariter 2-3-pinnatisecta, foliolis in lacinias confertas, lineari-lanceolatas
vellineares, acutas fissis. Racemi densiusculi, multi-(sub-20-) flori, quam in
typo paulo longius pedunculati. Bractew late, lineari-oblonge, cuspidate,
pedicellis breviusculis (circa 2 mm. longis) erecto-patentibus paulo breviores.
Sepala saltem 1 mm. longa, *50-"66 mm. lata, ovato-lanceolata, plus minusve
serrata, nervo dorsali viridiusculo rosea aut roseo-albida, caduca. Corolla ut
in var. Chavinii, sed petali superioris alis angustioribus obscure purpureis.
Fructis plane latiores quam longi (2 mm. longi et 2:25 mm. lati), subtruncati,
siccitate cum apicis foveolis latis rugosi.
Hee varietas var. Chavinii proxima est, sed pedicellis brevioribus erecto-
patentibus, bracteis sepalisque latioribus, corolla alis angustioribus obscurius
colorata, fructibus multo latioribus distinguitur.
F. officinalis var. Wirtgenii foliorum. segmentis latioribus, sepalis corollis
fructibusque majoribus conspicue differt.
F. Schleicheri pedicellis gracilioribus et multo longioribus, bracteis mini-
mis, floribus saturate roseis petali superioris alis valde reflexis, fructibus
apiculatis rugulosis plane recedit.
F. Vaillantii var. venetica in cultis prope Torri del Benaco, Rivoli et
Veronam in Venetia Italie septentrionalis habitat, ubi frequentissime a cl.
Rigo ut F. Laggeri collecta est. Hæc planta non solum ut F. Laggeri et
F. Vaillantii sed etiam sub nomine F. Schleicheri (Fiori, ete., Fl. Exsice. Ital.
No. 566!) distributa est. Exemplar hujus varietatis, ut videtur, sub nomine
F. Laggeri prope Weimar (Kastenberg) a Dr. Torge collectum in herbario
Kewensi est.
An element of doubt exists as to whether the above three varieties are best
placed as such under F. Vaillantii, and further study of the group seems
desirable, especially as the features of the floral organs can be readily
mistaken owing to their small size.
318 MR. H. W. PUGSLEY : A REVISION OF THE
The short racemes of minute flowers seen in var. conferta are not unlike
those of F. Schrammii, which is treated as a separate species, but its fruit
characters seem to bring it to F. Vaillantii, with which it also agrees in the
form of the corolla-spur.
The specific type, peculiar chiefly for the distant and sparing leaflets noted
by its earliest describers, looks widely different from well-marked examples
of var. Chavinii, not only in foliage but also in its slenderer and more
branching habit, its differently coloured corolla with apparently a more
ascending spur, and its distinctly broader fruits. But Haussknecht’s remarks
that various intermediates are to be found seems correct from the evidence
of dried material, and this renders it difficult to define the two forms as
distinct species.
The variety venetica is most closely related to var, Chavinii, which it
resembles in habit and foliage. These features, combined with its broad
fruits, somewhat recall those of F, officinalis, and it might be confused with
the small-flowered varieties of this Species on a cursory examination.
Fumaria Vaillantii is one of the most widely distributed species of the
genus, extending from Spain across Europe to the Altai Mountains of
Central Asia and to the Indian frontier. In Kurope it appears to be less
plentiful in the Mediterranean region than in the central portions of the
Continent, while in Asia it occurs from the Aigean to Beloochistan,
Afghanistan and Kashmir, as well as further north from the Caspian
eastwards to Chinese Songaria. Ledebour’s specimens at Kew from the
Altai Mountains are a mixture of this species and F. Schleicheri.
The occurrence of F. Vaillantii in North Africa and the Canaries, though
reported by both Hammar and Haussknecht, seems open to doubt. It is
indeed recorded by Ball for one Moroccan station (Journ. Linn. Soc. xvi.
p. 214 (1878)), but Battandier & Trabut (FI. de l'Algérie, p. 29) remark that
they have not seen this species from Algeria, and Munby's Algerian
specimens at Kew under this name belong to F. parviflora. More recently,
however, a Tunisian habitat has been reported by Murbeek. (Contr. FI.
Nord-Ouest. Afrique, p. 5 (1897)). The examples from the Canaries similarly
named, both in Herb. Kew and Herb. Mus. Brit., are referable to Z. parviflora.
The specific type, originally described by Loiseleur from a French plant,
seems to be chiefly found in Western Europe on the calcareous soils of
France and Britain. It also grows in the Tyrol (Hb. Kew D), in Albania
(Hb. Kew !), in Asia Minor (Hieropolis, Haussk. in Hb. Mus. Brit. !), and
in Persia (Teheran, Kotschy, in Hb. Kew!) ; while still further east a
similar form, sometimes with broader sepals, prevails, which is represented by
the exsiccata Jacquemont, No. 723, Kashmir, in Herb. Kew! and Aitchison,
No. 210, Kurrum Valley, Afghanistan, in Herb. Kew, as 7. parviflora !
The variety Chavinii, which is not confined to caleareous ground, appears
from herbarium material to be the prevalent form, though not always equally
GENERA FUMARIA AND RUPICAPNOS. 319
well marked, of Central and Northern Europe, and is the only form seen
from Scandinavia. It also grows in Spain and in France, as well as in
Britain, but no undoubted examples of it have been identified from the more
eastern parts of the specific range.
44. FUMARIA ScHRAMMII, sp. nov.
F. Vaillantü var. Schrammii, Haussk. in Flora, lvi. 444 (1873) ; Rouy &
Foucaud, Fl. Fr. i. 181 (1893) ; F. parviflora a. tenuifolia aa. Schrammii,
Ascherson, Verhandl. Bot. Ver. Prov. Brandenburg, v. 221 (1863).
asice. Bourgeau, Pl. d'Espagne, 1850, No. 538, ut F. Vaillantii!
Filarszky,Fl. Exsicc. Austro-Hungarica, No. 29031, ut P. ScAleicheri! Schneider,
Iter Baleanicum, 1907, No. 144, Bulgaria prope Varna! Bornmüller, Pl.
Lydi et Cariæ, No. 9015, Smyrna, ut F. Schleicheri var. microcarpa ! Born-
miiller, Pl. Anatol. Or. 1889, Nos. 127 & 135, et 1890, No. 1739! Kotschy,
No. 55, Teheran, 1846, ut F. Vaillantü !
Fumaria annua, sepissime gracilis, caulibus tenuibus nune elongatis
sparsis nunc ramosissimis compactis preedita, raro petiolis cirrhosis scandens.
Folia plus minusve glauca, sepius ampla, foliolis tenuibus longe petiolatis in
lacinias lineari-lanceolatas vel fere lineares (swpius satis longas) planas
acutas fissis 2—2-pinnatisecta. Racemi breves, pauci-(6-15-) flori, breviter
peduneulati, subsessiles vel etiam sessiles. Bracter | anguste, lineari-
subulate, tenuiter acuminate, pedicellis fructiferis erecto- patentibus brevissimis
(circa 2 mm, longis) sepius gracillimis subduplo breviores. Sepala *5— 15 mm.
longa, *25—4 mm. lata, sublanceolata, inciso-dentata, albida, facile caduca.
Corolla 5-55 mm. longa, pallide lilacina; petalum superius alis lilacinis vel
subpurpureis fere patentibus emarginatum, valde dorso compressum, calcare
deflexo parum curvato præditum ; petalum inferius marginibus lilacinis
patentibus subtruncatum spathulatum; petala interiora apice atropurpurea
fere recta. Fructús parvi, l:15-2 mm. longi et lati, subrotundi vel subrotundo-
ovati, cum apiculo brevissimo persistente obtusiusculi vel obtusi, inferne in
stipitem angustum obscurum contracti, paulo compressi et plane carinati, in
sicco apicis foveolis distinctis rugulosi.
B. orientalis, var. nov.
Exsicc. Brotherus, Pl. Caucasice, 1881, No. 426, Carthalinia, in Hb.
Mus. Brit., ut F. Vaillantii! Huet du Pavillon, Erzerum, 1853, in Hb.
Mus. Brit., ut F. microcarpa! Komarow, Samarkand, 1892, in Hb. Kew,
ut F. asepala! Gilgit Exped., No. 191, in Hb. Kew!
Herba robustior, sepius basi ramosissima, foliolis quam in typo sparsioribus.
Racemi plerumque subsessiles, raro pedunculati. Corolla 5-6 mm. longa,
petali superioris valde emarginati alis pallide purpureis. — Fructs 2 mm.
longi et lati, cum apiculo distincto (quam ty pi longiore) persistente obtusissimi,
inferne abrupte angustati, quam in typo minus compressi, in sicco rugosi
potius quam rugulosi.
330 MR. H. W. PUGSLEY : A REVISION OF THE
Hee species Fumariwe Vaillant? cujus formam umbrosam sæpe refert
simillima est, sed per foliorum lacinias longiores, per racemos breviores
sæpe subsessiles, per corollas lilacinas petali superioris calcare parum curvato,
per fruetüs minores persistenter apiculatos minus rugosos specifice differre
videtur. :
F. parviflora, Lamk. habitu robustiore, foliorum segmentis angustioribus
plus minusve canaliculatis, bracteis sepalisque latioribus, corollis albidis
petalo superiore haud emarginato calcare adscendente, fructibusque valde
carinatis rugosis seepius mucronulatis plane differt.
F. Sehrammii Hispaniam (Reuter, Madrid, in Hb. Kew !), Galliam
(Bentham, Avignon, in Hb. Kew! Heverchon, Briançon et Vaucluse,
in Hb. Kew! Lille, Haussk. Gironde, Rouy & Foucaud), Helvetiam
(Graubünden, in Hb. Zurich!), Germaniam (Ascherson & Benitz,
Brandenburg, in Hb. Kew! Thuringen !), Hungariam (Filarszky, No. 2903 1!),
Bulgariam (Schneider, Varna !, Macedoniam (Salonika, Haussk.), et
Tauriam (Hb. Pallas!) Europe habitat; in Asia etiam Anatoliam
(Bornmiiller, Smyrna! et Amasia!), Armeniam (Huet du Pavillon, Erzerum
—var. orientalis !), Transcaucasia (Brotherus, Carthalinia—var. orientalis !),
Persiam (Kotschy, Teheran! Bunge, Astrabad—var. orientalis !), Turkestan
(Komarow, Samarkand—cum var. orientali!), Chitral (Relief Expedn.,
No. 15875, in Hb. Mus. Brit., ut F. parviflora !), Gilgit (Expedn. No. 191,
in Hb. Kew—var. orientalis !), et Kashmir (Winterbottom, in Hb. Kew !).
A compact and densely branched Spanish plant in Herb. Zurich, labelled
without date * F, cæspitosa Loscos Exsicc. Fl. Arag. n. Castelseras. Leg.
Loscos " differs from the original plant sent out by Loscos as F. cwspitosa
and is a form of F. Schrammit var. orientalis. The specimen of F. cwspitosa
in Herb. Boissier also belongs to this form.
It is only after some hesitation that F. Sehrammii has been described as a
species distinct from F. Vaillanti, to which it is evidently very closely
alied and of which it will be seen to possess almost the same general
geographical distribution. While resembling its ally, however, it seems to
keep constantly distinct in several partieulars, and no intermediate material
has come under observation concerning the identity of which any reasonable
doubt can be entertained.
Haussknecht, in describing F. Schrammii as a variety of F. Vaillantii,
mentions it as a remarkable plant that may prove to be a separate species,
and it seems probable from the notations on Bornmiiller’s labels that at a later
date both he and Ascherson were confirmed in this view.
The variety orientalis has been placed under F. Schrammii owing to its
similarly coloured flowers and persistently apiculate fruit, but the latter
organ differs considerably in form from that of the type, and it may prove,
when better material of the variety is available, that the two plants are not
conspecific.
GENERA FUMARIA AND RUPICAPNOS. 321
45. FUMARIA ASEPALA, Boissier, Fl. Orient. i. 135 (1867); Haussk. in Flora,
lvi. 461 (1873).
Egsicc. Bornmüller, Pl. Anatol. Orient., 1893, No. 1737, Amasia, ut F.
parviflora f. erecta ! Wiedemann, Anatolia, in Hb. Kew, ut F. parviflora!
Bornmüller, Iter Persico-Turcicum, 1892-3, No. 3136, Angora, Anatolia, ut
F. parviflora ! Kotschy, No. 17, Mons Taurus, 1836, ut F. parviflora (par-
iim)! Kotschy, Iter Cilicico-Kurdicum, 1859, No. 41, in Hb. Mus. Brit., ut
F. parviflora! — Sintenis, Iter Orientale, 1888, No. 282, Mesopotamia!
Stapf, No. 2158, Schiras, 1885, in Hb. Kew!
Fumaria annua, humilis, caulibus gracilibus suberectis haud scandentibus
smpius valde ramosis predita. Folia glauca, irregulariter 2- (raro 3-)
pinnatisecta, foliolis paulo sparsis in lacinias lineari-oblongas planas acutas basi
plus minusve attenuatas fissis. Racemi floriferi brevissimi, fructiferi tandem
paulo elongati, pauciffori (6-12-flori), subsessiles aut raro brevissime pedun-
culati. Bractew lineari-oblonge, acuminate, albidz, pedicellos fructiferos
erecto-patentes brevissimos (1-2 mm. longos) haud crassos sepe superantes sed
nonnunquam breviores. Sepala absentia aut rarissime (Sintenis, Iter Or. No.
282, partim) minutissima vix videnda. Corolla 4-5 mm. longa, alba ; petalo
superiore valde dorsum compresso, alis latis albis patentibus vel etiam deflexis
ultra carin: crass viridis apicem productis conspicue emarginato, atque
calcare vix curvato preedito ; petalo inferiore lato, carinæ crasse viridis
marginibus albis patentibus apicem versus abrupte dilatatis et ultra apicem
productis spathulato-emarginato, petalum superius extra calcar simulante ;
petalorum interiorum apice (interdum obscure) purpureo ab exteriorum alis
subcelato. Fructis parvi, circa 2 mm. longi et subseque lati vel paulo angus-
tiores, subrotundi vel subrotundo-ovati, paulo compressi et plane carinati, cum
apiculo brevissimo persistente obtusi et inferne paululum angustati, siccitate
apice minute sed plane bifoveolati rugulosi.
B. compacta, Haussk. l. c. 461 (1873).
Exsicc. Aucher-Eloy, Herb. d’ Orient, No. 4051, Ispahan, ut F. Vaillantii !
Kotschy, No. 45, Teheran, 1843, in Hb. Mus. Brit. (non Hb. Kew), ut
F. parviflora! Bunge, Iter Persicum, 1859, inter Teheran et Tabriz, ut
F. parviflora! Polak, Iter Persicum, 1882, Dauletabad, in Hb. Kew!
Planta nana, caulibus curtis suberecta. Foliorum lacinim anguste
oblanceolate, obtuse vel rarius mucronate. Racemi constanter brevissimi,
vulgo 4-6-flori. Flores typi, sed petalorum interiorum apex viridescens nec
purpureus. Fructis subrotundi, nonnunquam quam in typo paulo majores.
y. aprica, var. nov.
Exsicc. Bornmüller, Iter Persico-Turcicum, 1892-3, No. 3234, Anatolia
(Siwas), ut F. parviflora f. aprica ! Kotschy, Iter Cilicico-Kurdicum, 1859,
No. 41, in Hb. Kew, ut F. parviflora!
M
bo
n2
MR, H. W. PUGSLEY : A REVISION OF THE
Foliorum lacini angustiores, lineares nec lineari-oblonge. — Bractez quam
in typo angustiores. Corolla dorso et petalorum exteriorum marginibus
angustioribus presertim apicem versus rubescens. Fructüs minimi, 1°75 mm.
longi et 1*5 mm. lati, subrotundi. Aliter ut in typo.
Hee species que Fumariv Vaillantii habitum foliaque sed F. parviflore
flores albidos habet non solum calyce abortivo sed etiam petalis duobus
exterioribus valde emarginatis altero alterum simulante notabilis est.
F. Vaillantii racemis longioribus pedunculatis, sepalis lanceolatis, corollis
majoribus roseis caleare adscendente petaloque inferiore truncato nec
emarginato, fructibus sine apiculo persistente rugosis ; F. parviflora habitu
robustiore, foliorum magis decompositorum laciniis angustioribus vulgo
canalieulatis, racemis longioribus, sepalis szepissime ovatis rarissime obsoletis,
corollis vulgo majoribus calcare adscendente petalisque exterioribus haud
emarginatis, fructibus sepissime rugosis valde carinatis insuper distinguitur.
F, Schrammii habitu laxiore. foliorum laciniis et racemis longioribus,
sepalis sublanceolatis, corollis majoribus lilacinis petalo superiore solo
emarginato satis differt.
F. indica habitu robusto, foliorum laciniis longis angustis, racemis longis
peduneulatis, floribus major ibus roseis, petalis exterioribus haud emarginatis,
fructibus truncatis rugosis facile separanda est.
This distinct though inconspieuous fumitory, remarkable for its strongly
compressed flowers with the outer petals almost equally winged, is widely
distributed in Asiatic Turkey and extends eastwards into Persia. The variety
compacta seems to occur chiefly in the latter country, but it is also recorded
by Haussknecht for two Syrian localities.
46. FUMARIA PARVIFLORA, Lamarck, Encycl. Méthod. ii. 567 (1788) ; Hamm.
Mon. 16 (1857) ; Haussk. in Flora, lvi. 456 (1873) ; Pugsley, Fum.
in Brit. 60 (1912). F. tenuifolia, Roth, m fasc. ii. 82 (1800).
Clusius’ Capnos cretica floribus candidis fused macula (Hist. Rar. Pl. lib. vi.
p. ecix (1601)) appears to be this species. It is cited by C. Bauhin (Pinax,
p. 143 (1623)) as a synonym of Fumaria minor folio oblongo capillaceo, which,
according to A. P. de Candolle (Bull. Hb. Boissier, Tom. iv. 2™ Série, No. 3,
p. 304 (1904)), is represented in Bauhin’s Ho by F. parviflora.
Icones. Sturm, Deutschl. Fl. i. 62, tab. 16; Reichb. Icon. Fl. Germ. iii.
tab. 1. fig. 4451 (f. sepalis majoribus) ; Hamm. l.c. tab. ii; Clavaud, Fl.
Gironde, Pl. 4. fig. 4
Ewsice. Heldreich, Herb. Grec. Norm. No. 1206, ut F. parviflora
f. umbrosa ! Wirtgen, Pl. Sel. Rhenan. fasc. iv. No. 159! Bourgeau, PI. Pyr.-
Espagnol. No. 389! Bourgeau, Pl. d'Espagne, 1851, No. 1014! Fiori &
Béguinot, Fl. Exsiec. Ital. No. 1051 (ad var. Symei accedens)!
GENERA FUMARIA AND RUPICAPNOS. 323
Forma Vivianii.
F. leucantha, Viviani, Fl. Corsieze Sp. Nov. Diagn. 12 (1824) ; F. parviflora
var. leucantha, Clavaud in Act. Soc. Linn. Bordeaux, xxxv. 4° sér. v. 276
(1881), et Fl. Gironde, 52 (1882), ex parte.
Eæsice. Pichler, Fl. Exsice. Austro-Hungarica, No. 87, Spalato, ut
F. 4 arviflora! Reverchon, Pl. de Sardaigne, No. 91, ut F. parviflora (Ë. fol.
laciniis latioribus) !
Fructibus.obtusis nec acuminatis nee emarginatis a typo differt.
In the ease of this species Hammar’s diagnosis is generally accurate, but
he appears to have overlooked the characteristie broad bracts, which he
describes as linear, as well as the normally obtuse but not emarginate upper
petal, with its purple bloteh on the outer or lower side of the wings.
The form regarded as the specific type, in accordance with the general
view of authors, is the more robust plant, wiih laxer and less glaucous
foliage, whiter and somewhat narrowly winged corollas, and shortly
mucronulate fruits, as described in “ Fumaria in Britain” (F. parviflora var.
leucantha, Clavand, l.c., excl. forma Vivianii). The rotundate, ascending
spur common to all the forms of the species is not mentioned, through
inadvertence, in that description. It may be added that the leaves in this
species are commonly 3-4 rather than 2—3-pinnatisect, and the corolla is not
often more than 5 mm. in length. |
Flores albi, vix roseo-tincti ; sepala seepissime albida; petali
superioris alee quam in typo angustiores.
B. Symei, Pugsley, l. c. 65 (1912).
F. Vaillantii, Bab. in Trans. Bot. Soc. Edin. i. 36 (1844) ; et Eng. Bot.
Suppl. 2877 (1844), ex parte, non Lois.
Icon. Eng. Bot. Suppl. 2877, ut F. Vaillantii (partim—f. flor. albis).
Easice. Reverchon, Pl. de Andalousie, No. 557, ut F. parviflora !
In addition to the short leaf-segments, relatively large sepals, and finally
retuse fruits characteristic of this variety, it may be distinguished by its
racemes, which, both in flower and in fruit, are normally laxer than in the
other forms of this species.
Other examples collected in Spain (Bourgeau, Pl. d'Espagne, 1850,
No. 531 1), in Morocco (Hooker, Casa Blanca, 1871, in Hb. Kew !), and in
the Canaries (Lowe, No. 21bis, Orotava, 1858, in Hb. Mus. Brit., ut F, parvi-
flora fl. albis!) are closely related to, if indeed separable from this variety.
y. latisecta, Haussk. l. c. 460 (1872).
Exsice. pud No. 410, Mt. Sinai, 1835, ut F. parviflora ! Macdonald,
Arabia Petræa, 1849, in Hb. Kew, sub F, parviflora |
Planta satis robusta, suberecta, haud scandens ; folia vix glaucescentia,
ampla, laciniis linearibus vel oblongo-linearibus, 1-2 mm. (ad 3 mm. ap.
Haussk.) latis, fere planis, obtusis obtusiusculisve prodita. /tacemi sessiles,
324 MR. H. W. PUGSLEY: A REVISION OF THE
densi (etiam fructiferi), quam folia opposita multo breviores; bracteæ
pedicellos brevissimos (circa 1 mm. longos) plane superantes. Corolla
4:5-5 mm, longa, albida ; petalo superiore anguste alato. Fructús parvi, vix
2 mm. longi ac lati, subrotundi, obtusi vel subretusi, brevissime apiculati,
siccitate subleves, Aliter ut in typo.
This variety, which is well represented at Kew, is notable for its ample
foliage with broad and nearly flat leaf-segments, for its very short and
dense racemes, and for its nearly smooth instead of rugose fruits.
Tommasini's specimen from Trieste, in Herb. Boissier, included in Hauss-
knecht’s citations for this variety, shows poor flowers and fruits, but has the
aspect of a form of F. Vaillantii, as labelled. A few other European
exsiccata, as “ Reverchon, Pl. de Sardaigne, No. 91," approach this variety
in foliage.
8. persica, var. nov.
Egxsice. Sintenis, Iter Transcaspico-Persicum, 1900-01, No. 1548, ut F.
asepala ! Bornmüller, Iter Persicum alterum, 1902, No. 6112, Persia borealis,
ut F. parviflora! Kotschy, No. 314, Schiras, 1842, ut F. parviflora! Fl.
Baluchistan, No. 3521, ut F. parviflora (partim—pl. fl. albis)! Kotschy,
No. 17, Mons. Taurus, 1836, in Hb. Kew, ut X. parviflora (partim)! Postian,
Pl. Mont. Syriz borealis, Aintab, in Hb. Mus. Brit., ut F. asepala !
Planta robusta, ramosior, haud scandens; /olia plus minusve glauca,
ampla, laciniis linearibus vel oblongo-linearibus (raro plus 1 mm. latis),
confertis, brevibus, crassiusculis, subcanaliculatis, acutis subacutisve prodita.
Racemi sepissime sessiles, breves, floriferi densi, fructiferi paulo elongati ;
bracteze pedicellos brevissimos (circa 1 mm. longos) multo superantes.
Sepala absentia aut minuta (circa *5 mm. longa), triangulari-ovata, acuminata,
albida. Corolla minima, 4-4°5 mm. longa, alba; petali superioris alis
angustis vulgo sine maculá purpurea ; petalorum interiorum apice purpureo
vel viridiusculo, Fructús 2 mm. longi et paululum latiores, subrotundo-
obovati, obtusissimi vel subtruncati, obscure brevissime apiculati, in sicco rugosi.
Hoe varietas que per foliorum amplorum lacinias breviores confertas
et per fructüs obtusissimos a typo longe distat varietati latisect« similis est,
sed foliorum laciniis brevioribus angustioribus, sepalis absentibus vel minutis,
corollà minima, fructibusque obtusissimis rugosis potius quam sublevibus
satis differre videtur.
! ! Flores tandem roseo-tincti ; sepala rosea ; petali superioris alæ
sæpius latiores.
e. glauca, Clavaud in Act. Soc. Linn. Bordeaux, xxxv. 4 sér. v. 277
(1881), et Fl. Gironde, 53 (1882); Nicotra, Le Fum. Ital. 69
(1897). F. glauca, Jordan, Pugillus, 8 (1852) ?
Ezsice, Schultz, Herb. Norm. No. 415 ter, ut F. parviflora! Bourgeau,
Env. de Toulon, No. 18, in Hb. Kew, ut F. parviflora! Kralik, Pl. Tunet,
GENERA FUMARIA AND RUPICAPNOS. 325
1854, No. 8, in Hb. Mus. Brit., ut F. parviflora! Bornmüller, Iter Persico-
Turcieum, 1892-3, No. 2030, Persia borealis, ut F. parviflora !
Planta quam typus humilior, haud scandens. Folia valde glauca, laciniis
anguste linearibus, brevibus, canaliculatis, seepe crassiusculis. Racemi floriferi
densi; flores quam in hujus speciei aliis formis magis patentes. Corolla
usque ad 5 mm. longa, sæpius lete roseo-tincta ; petali superioris ale quam in
typo latiores, presertim apicem versus dilatate ; petalo inferiore swpe sub-
truncato et libero deflexo. —ructás vix 2 mm. longi latique, subrotundi,
obtusiusculi brevissime apiculati, rarius subrotundo-ovati acutiusculi. Aliter
ut in typo. i
The corolla of this pretty variety differs somewhat in shape from that
of the other forms of this species, and at the time of flowering is generally
less erect in the raceme.
Plants occur (cf. Bourgeau, Pl. d'Espagne, 1849, No. 22, in Hb. Mus.
Brit. !) having the habit and flowers of var. glauca with the more attenuate
fruit of var. acuminata.
£. acuminata, Clavaud, ll. c. 277 et 53 (1881 et 1882) ; Pugsley, l.c.
64 (1912).
Icon. Smith, Eng. Bot. 590, ut F. parviflora.
Exsice. Schultz, Herb. Norm. Nos. 415 & 415 bis, ut F. parviflora!
Sintenis, Iter Trojanum, 1883, No. 301, Kenkoei, ut F. parviflora !
Planta ut var. glauca humilis, haud scandens. Folia intense glauca,
laciniis tenuiter linearibus vel subcapillaceis canaliculatis. Racemi floriferi
densiusculi. Corolla relative majuscula, 5—6 mm. longa, tandem roseo-tincta ;
petalorum exteriorum margines quam in typo paululum latiores. Fructis
2:25-25 mm. longi et 2 mm. lati, subrotundo-ovati, cum apiculo brevissimo
persistente acutiusculi vel acuminati,in sicco apicis foveolis fere obsoletis rugosi.
This variety is notable not only for its more acuminate fruits but for
its very fine, almost capillary leaf-segments ; and its flowers are somewhat
larger than those of any other form of the species.
q. sinaitica, Haussk. l. c. 460 (1873).
Ewsice. Boissier, Jardin du Sinai, Arabia Petræa, 1846, in Hb. Boissier !
Planta nana, vix 4 em. alta, satis ramosa. Folia intense glauca, inferiora
caules fere æquantia, laciniis obtusis brevissimis (superioribus vix 1 mm.
longis) preedita. Corolla roseo-tincta. Fructis ut in typo.
This seems to be a very rare plant, known only from Mt. Sinai.
0. indicoides, var. nov.
Exsicc. Bornmüller, Iter Persico-Turcicum, 1892-3, No. 9, Persia australis
(Farsistan), ut F. parviflora, transiens ad F. segetalem, Hamm.! Bagdad,
Samaur, (ex Herb. Schiifli) in Hb, Kew! Russell, Aleppo, in Hb. Mus.
Brit. !
326 MR. H. W, PUGSLEY : A REVISION OF THE
Planta internodis longis robusta, suberecta vel diffusa. Folia glauca,
laciniis linearibus vel oblongo-linearibus, breviusculis, acutiusculis, canalicu-
latis. Racemi etiam floriferi haud densi ; bracteæ pedicellos crassos breves
(circa 1:5 mm. longos) plane superantes. Sepala 1-15 mm. longa, subrotundo-
ovata, acuminata, dentata. Corolla vix 5 mm. longa, dilute rosea; petali
superioris alis latis omnino roseis; petalorum interiorum apice viridiusculo
potius quam purpureo. Fructis 2-2:25 mm. longi et lati, subrotundo-obovati,
obtusissimi, obscure brevissime apiculati, priesertim apicem versus conspicue
carinati, in sicco apicis foveolis obscuris rugosi.
Hee varietas per flores sine apice atropurpureo obscure roseos et per
fruetüs majores obtusissimos apicem versus conspicue carinatos a F. parvi-
fore aliis formis separanda est et transitum. ad F. indicam denotare videtur,
sed foliorum laciniæ breviores, racemi subsessiles et presertim. corolla
brevior alis patentibus caleareque rotundato adscendente prodita ejus eum
F. parviford propinquitatem certissime monstrant,
An example in Herb. Mus. Brit. (Haussknecht, Iter Orient. Schiras, ut
F. parviflora Y), with small rosy-white flowers and obscure sepals, seems inter-
mediate between this variety and 8. persica.
Hammar's variety segetalis (Mon. p. 17), distinguished by sepals one
fourth as long as the corolla and as broad as its tube, has been shown under
F. Reuteri to belong to that species and not to F. parviflora,
Haussknecht, who does not seem to have seen Lange’s specimen on which
this variety was founded, regards it as a form of true F, parviflora, and
cites for it several fresh exsiccata, of which those from Segura de la Sierra,
Murcia, and from Aintab have been examined, and are found to resemble
closely Clavaud’s variety glauca.
The original specimens of F. cvspitosa, Loscos (Loscos, Series Exsicc.
Fl. Aragon, Centuria prima, No. 2) are referred by Haussknecht in Oesterr.
Bot. Zeit. xxvii. p. 50 (1877), to F. parviflora f. erecta, Haussk. This plant
is admitted as a distinet species in Willkomm & Lange's FI. Hispaniea, iii.
p. 884, but in their Supplement (p. 311) it is reduced to a variety of F.
Vaillantii with the remark * vix varietas dicenda." The example in Herb.
C. Bailey is an early-flowering form of /. parviflora, probably identical
with the material seen by Haussknecht, but as already noticed under
F. Schrammii, the specimens in Herb. Boissier and Herb. Zürich are quite
different, so that some confusion presumably occurred among the plants sent
out under this name. The original description by Loscos and by Willkomm
& Lange point, on the whole, to a form of F, Vaillantii.
Another plant allied to F. parviflora is F. Trabuti, Battandier & Trabut,
Fl. de l'Algérie, i. 29 (1888), of which no material has been seen. It is’
described as follows :— :
Fumaria verisimiliter annua caulibus curtis robustis, Folia inferiora
GENERA FUMARIA AND RUPICAPNOS. JP
multa, conferta, angusta, oblonga, laciniis curtis lanceolatis paulo latis haud
canaliculatis predita, eis sectionis Jetroeapnos quarumdam specierum
similia ; folia superiora ut in F. parvijlord. Flores satis magni, 6 mm. longi,
albi; sepala lata. Fructis magni, obtusi, apiculati. Habitat in provinciá
Oran Algeriæ.
A further variety of F. parviflora, established as S. macrocarpa in J. Perez
Lara's Florula Gaditana, pars 5, p. 64, in Anales de Hist. Nat. Soc. Espagn.
(1898), is said to liave rosy flowers and fruits two or three times as large as
in the type.
Like F. Vaillantii, F. parviflora is one of the most widely spread members
of the genus, having a somewhat similar range extending from Spain across
Europe to Afghanistan and Beloochistan. It is clearly, however, a more
southern species and one more impatient of cold. In Europe it is commonest
in the Mediterranean region, and does not occur so far to the north and east
as F. Vaillantii, being little known in Russia except in the Crimea
(Herb. Pallas). Similarly in Asia it is absent from the distriets between the
Caspian and Mongolia, where F. Schleicheri, F. Vaillantii, and F. Sehrammii
have all been found. On the other hand, it is known from all the countries
of North Africa from Morocco to Egypt, as well as from the Canaries.
The commonest form of this polymorphie plant is no doubt the specific
type, varying considerably in the form of its fruit and connected with most
of the varieties by numerous intermediate forms. This occurs from the
Canaries, Morocco and Spain eastwards to Syria and Mesopotamia. The
variety Symei is known from Britain and Spain, and probably grows
elsewhere ; var. acuminata is mostly a plant of Western Europe, but has been
collected also in Asia Minor; var. glauca is pre-eminently a Mediterranean
form, growing on the African as well as the European side and extending
into Syria and Persia. Haussknecht’s varieties latiseeta and sinaitica are
both Arabian forms, probably collected in natural habitats, and examples
which he obtained in the desert near Bagdad seem almost identical with var.
latisecta, which Haussknecht records also for Dalmatia. The two remaining
varieties, both characterized by very obtuse fruits though dissimilar in other
respects, are the most eastern forms of the species. The variety persica
appears to bea prevalent form in Persia, growing also in Beloochistan, Syria,
and the Taurus Mts. ; var. indicoides, which is presumably a rarer form,
ranges, so far as is known, from Syria across Mesopotamia to Southern
Persia.
338 MR. H. W. PUGSLEY : A REVISION OF THE
RUPICAPNOS.
Rupicapnos, Pomel, Mat. Fl. Atlant. 16 (1860) ; Nouv. Mat. Fl. Atlant. i. 240
(1874). Fumaria sect. Petrocapnos, Cosson & Durieu in Bull. Soc. Bot.
France, ii. 305 (1855) ; Willkomm & Lange, Fl. Hisp. iii. 878 (1880) ;
Cosson, Comp. FI. Atlant. ii. 80 (1883-1887).
Plant: perennes caudice decumbente caulibusque brevissimis aut rarius annuw
namv caule suberecto brevissime ramoso. Folia in speciebus perennibus
pleraque subradicalia, in omnibus semper longe petiolata, foliolis breviter
petiolatis vel subsessilibus 1-2-pinnatisectis vel inciso-flabellatis irregulariter
pinnata (R. muricarid exceptá) ; segmenta secundaria decurrentia ; petioli
nunquam cirrhosi. Inflorescentia pedicellis fructiferis gracilibus elongatis
tandem deflexis racemoso-corymbiformis. Bracteæ pedicellis fructiferis
multoties breviores. Corolla uni- vel inæeque bicalcarata ; petalerum ex-
teriorum margines apicem versus in statu juvenili jam multo explicati ut in
genere Sarcocapnos ; petalum superius subplanum potius quam semi-cylin-
drieum, ealearatum, superne gibbum format qui apicem vix attingit et
margines sepissime patentes in limbum latum interdum dilatatos nunquam
purpureos habet sed non alas in jugum laterale productas ; petalum inferius
inferne gibbum apicem vix attingentem marginibus patentibus cireundatum
formans sepe basi ipsá breviter calearatum vel saccatum fit; petalorum
interiorum nervus medius apicem versus sepe conspicue alatus est. Floris
characteres reliqui utin genere Fumarid. Fructus monospermus, indeli.scens,
nuciformis, semper mucronatus vel rostratus sed haud apice biforeolatus
(sine rimis in mesocarpio) est ; exocarpium tuberculis rugosum ; endocarpium
omnino ad mesocarpium adhwrens ; semen apice rotundatum vel paulo depressum,
sine rugá longá infra mieropylam.
Perennial, or more rarely annual plants, always with short stems, growing
in rock-clefts. Inflorescence corymbiform with pedicels which lengthen in
fruit and become deflexed so as to carry the seeds to the cavities of the rocks
on which the planis grow. Lower petal often shortly spurred. Fruit
without apical pits.
In the perennial species of Rupicapnos the root is quickly elongated and
the leaves are at first all radical, forming a close tuft. After the first year,
the plant develops a thick, branched, decumbent rootstock, from which arise
very short, branched stems, bearing long-petioled leaves and subterminal
corymbiform racemes. The foliage is usually less decompound than in
Fumaria, and the leaf-segments are much more shortly petioled and more
decurrent, the secondary petioles never being cirrhose. The flower-buds of all
the species resemble those of the genus Sarcocapnos rather than Fumaria,
having well developed margins to the two outer petals, while the spur of the
upper one is very small. This spur, as seen in perfectly developed flowers,
varies greatly in different species and is of importance for affording specific
GENERA FUMARIA AND RUPICAPNOS, 329
characters, but in shade-grown plants of some species it does not always
attain its normal development. So far as can be judged from the limited
material examined, the corolla is more uniform in the species of this genus
than in Fumaria, and shows little tendency to cleistogamy.
CONSPECTUS OF SPECIES.
Secrion I. MURICARIA.
Suberect annual plants with cauline leaves and few-flowered racemes. Flowers small or
rather so, 4-8 mm. long, white; outer petals rarely dilated apically into a spreading, sub-
orbicular limb, little longer than the inner petals, which are tipped with dark purple. Fruit
very large, or at least of moderate size. |
* Leaves 2-trisect. Racemes subsessile.
l. R. muricaria. Flowers 7-8 mm. long ; lower petal spathulate. Fruit very large, sub-
rotund, marginally muricate, with long, large beak,
** Leaves 2-pinnatisect, Racemes peduncled.
2. R. delicatula, Leaves with multifid leaflets. Flowers 4 mm. long; outer petals
apically dilated. Fruit subglobose, muricate, apiculate.
8. R. longipes. Leaves deltoid. Flowers 6-8 mm. long; lower petal dilated into a
suborbicular limb. Fruit very large, subrhomboid, muricate, with
large, broad beak. Pedicels very long.
4. R. pretermissa. Leaves deltoid. Flowers 6-7 mm. long; lower petal spathulate.
Fruit rather large, ovate-elliptical, finely rugose, with compressed
beak.
5. R. sublevis. Leaves oval, with closer leaflets, Flowers... Fruit of moderate
size, narrowly obovate, finely rugose, with small beak.
SrcrtIon II. SARCOCAPNOIDES.
Decumbent perennial plants with quinate-pinnate, mostly subradical leaves and many-
flowered racemes. Flowers small or rather so, 4-10 mm, long; outer petals each apically
dilated into a spreading, suborbicular limb, distinctly longer and broader than the inner
petals, which are tipped with dark purple. Fruit small.
6. R. sarcocapnoides. Lobes of leaflets elliptical, Flowers 4-5 mm. long, white; lower
petal not saccate. i
7. R. Heboudiana. Lobes of leaflets cuneiform. Flowers 9-10 mm. long, rosy-white
lower petal saccate.
Secrion III. TRIPTERYX.
Decumbent perennial plants with 2-3-pinnatisect, mostly subradical leaves and many-
flowered racemes. Flowers small or rather so, 4-10 mm. long, whitish and more or less
tinted with yellow; outer petals more or less apically dilated into spreading, suborbicular
limbs, little longer than the inner petals, which are conspicuously winged and at most
obscurely tipped with dark purple. Fruit small to rather large.
* Flowers 4-5 mm. long; lower petal not saccate.
! Lower petal gradually dilated apically.
8. R. numidica. Leaves subdeltoid with remote leaflets, Spur of upper petal very
small, straight. Fruit oblong-obovate.
LINN. JOURN.—BOTANY, VOL, XLIV. 2D
330 MR. H. W. PUGSLEY: A REVISION OF THE
9, R. Cossonit. Leaves oval with closer leaflets. Spur of upper petal rather large,
curved. Fruit subrotund-obovate.
!! Lower petal more abruptly dilated apically ; spur of upper petal moderate.
10. R. tenuifolia. Leaves oblong, with finely cut leaflets. Wings of inner petals not
exceeding outer petals. Fruit subrotund.
11. Reaput-platalee. Leaves subdeltoid, with remote, cuneiform segments, Wings of inner
petals much exceeding outer petals. Fruit elliptical.
** Flowers 6-10 mm. long; lower petal subsaccate.
12. R. erosa. Leaves oblong; leaflets with broad segments. Spur of upper petal
rather long. Fruit subrotund, sometimes rather large.
Secrion IV. CALLIANTHOS.
Decumbent perennial plants, usually larger than those of the other sections, with
9-3-pinnatisect, mostly subradical leaves and many-flowered racemes. Flowers large,
12-16:5 mm, long, white or pale purple; outer petals never apically dilated into a spreading,
suborbicular limb, little exceeding the inner petals, which are tipped with dark purple;
lower petal saccate. Fruit very large to moderate in size.
Subsection AFRICAN E.
Lobes of leaf-segments relatively narrow. Flowers white.
13. R. africana. Leaves subdeltoid, with narrowly oblong lobes. Sepals ovate-
lanceolate. Spur of upper petal very long. Fruit of moderate
size, obovate-elliptie, shortly mucronate.
14. R. decipiens. Leaves deltoid, with subelliptical lobes. Sepals oval. Spur rather
long. Fruit very large, obovate, shortly mucronate.
15. R. platycentra. Leaves deeply divided, with small, obovate lobes, Sepals orbicular.
Spur rather long. Fruit . . . oval-elliptic.
16. R. cerefolia. Leaves more decompound, with small, narrow lobes. Sepals ovate-
lanceolate. Spurlong. Fruit of moderate size, elliptic, acuminate.
17. R. speciosa. Leaves with broadly oblong lobes. Sepals orbicular. Spur short.
Fruit . . . lanceolate-obovate, long-mucronate.
Subsection POMELIANZE.
Lobes of leaf-segments broad. Flowers pale purple.
18. R. Pomeliana. Leaf-segments subimbricate, with rounded lobes. Sepals orbicular.
Spur very short. Fruit rather large, subrotund-obovate.
19. R. oranensis. Leaf-segments with broad lobes but not imbricate. Sepals small,
M ovate. Spurlong. Fruit large, oval-obovate.
90. R. ochracea. Leafsegments not imbricate, with small, obovate lobes. Sepals
small, lanceolate. Corolla smaller, with long spur. Fruit narrow,
oblong.
Secrio I. MURICARIA.
Plante annus, suberectæ. Racemi pauciflori. Flores parvuli, rarius
parvi, 4-8 mm. longi, albi ; petalis exterioribus quam interiora ssepius fere
recta apice plane alata atropurpurea vix longioribus, marginibus eorum
apice subpatentibus nonnunquam in limbos suborbiculares dilatatis; stylus
Mri dii id wn ED ae eee 7 IS T
GENERA FUMARIA AND RUPICAPNOS, 291
malleiformis lobis divaricatis (an semper ?). Fructüs maximi ad modici,
sepius valde compressi, pericarpio quam in sectionibus sequentibus
verisimiliter tenuiore preediti.
RUPICAPNOS MURICARIA, Pomel, Nouv. Mat. Fl. Atlant. i. 245
Fumaria muricaria, Batt. & Trabut, Fl. d'Alger. i. 26
(PI. 14, fig. 1.)
Hasive, Chevallier, Pl. Saharze Alger. 1902, No. 397, Ghardaia, ut Fumaria
longipes !
Rupicapnos, ut videtur, annua, radice longa et caule suberecto (4-8 em.
alto) brevissime ramoso preedita. Cotyledones in statu foliaceo maxime,
circa 4 em. longæ, lineari-oblanceolate, tenuiter acuminatie, diu persistentes.
Folia caulina haud crassa, omnia longissime petiolata, eum petiolo 5-7°5 em.
longa, deltoidea, foliolis subtrifidis longiuscule petiolatis et segmentis
secundariis in lacinias parvas oblongas vel lanceolatas longe mucronatas vel
fere aristatas irregulariter multifidis trisecta. Racemi corymbiformes, pauci-
(2-5-) flori, brevissime pedunculati vel subsessiles, foliis plus duplo breviores.
Bractew 1-2 mm. longæ, subulate, tenuiter acuminatze ; pedicelli Jructiferi
filiformes, apice vix inerassati, flexuosi, ad 15 mm. longi. Sepala parva,
circa 1 mm. longa, ‘5—6 mm. lata, ovata vel lanceolata, vix peltata, acuta,
parce dentata vel subintegra, albida. Corolla 7-8 mm. longa, preeter carinas
virides alba vel leviter roseo-tineta ; petalo superiore apice marginibus
latiusculis subpatentibus anguste oblongo, truncato obtusissimo vel etiam retuso,
caleare gracili longo deflexo, [nectario flexuoso longissimo ap. Pomel];
petalo inferiore apice marginibus latiusculis patentibus spathulato, obtusissimo,
basi verisimiliter obscure saccato; petalis interioribus paulo sursum curvatis
apice plane alatis atropurpureis. Stylus malleiformis. Fructis maximi, sine
rostro 2°75-3 mm. longi et 3-3°25 mm. lati, subrotundi sed latiores quam
longi, cum rostro magno 1:5-2 mm. longo quadrangulato-subulato obtusissimi
et inferne subtruncati, valde compressi et acute carinati, pericarpio satis
tenui priediti, in sicco (sine apicis foveolis) carinam versus subregulariter
grosse et acute tuberculato-rugosi vel muricati, infra rostrum et ad fructüs
medium vix tuberculati.
Hee Rupicapnos distinetissima per folia trisecta, per racemos corymbi-
formes subsessiles paucifloros et prsesertim per fructüs maximos grosse
rostratos carinam versus muricatos ab generis omnibus aliis speciebus facile
dignoscitur.
R. muricaria Metlili (ap. Pomel), Ghardaiam et forsan alios locos in
regione Sahare Algeriensis habitat, ubi in fissuris rupium crescit.
This species is unique in the genus on account of its trisect leaves, its
subsessile, 2-5-flowered corymbs, and its very large, long-beaked fruits,
muricate only towards their margins. Chevallier’s exsiccata, which form the
basis of the foregoing description and from which the accompanying plate is
2D2
332 MR. H. W. PUGSLEY : A REVISION OF THE
drawn, were presumably referred to Fumaria longipes, Cosson, owing to that
plant being originally described as an annual with somewhat similar flowers.
Their agreement with the account of X. muricaria furnished by Pomel is
unmistakeable.
[2. RuricAPNos DELICATULA, Pomel, Nouv. Mat. Fl. Atlant. i. 246 (1874) ;
n. v. Fumaria delicatula, Batt. & Trabut, Fl. d'Alger. i. 26 (1888).
Rupicapnos annua, Folia foliolis multijidis in lobos ssepius parvos obovatos
vel oblongos, obtusos vel subacutos, divaricatos fissis verisimiliter 2-pinna-
tisecta. Racemi corymbiformes, breves sed floribus pluribus quam in
R. muricarid priediti, Bractew lanceolate, acuminate ; pedicelli. capillares,
apice inerassati, Sepala lanceolata, denticulata. Corolla minima, 4 mm.
longa, albida ; petalis exterioribus apice. dilatatis, calcare superioris brevissimo
recto paululum attenuato; nectario brevissimo : petalis intericribus, ut
videtur, apice purpureis. Fructis subglobosi, leviter apiculati, parum com-
pressi, siccitate acute tuberculato-rugosi vel muricati.
R. delicatula rupes calciferas ad Ksar-el- Maiam Algerie australis habitat.
This plant appears to differ considerably from the other known annual
species, and some of its features recall R. tenuifolia. Judging from Pomel's
knowledge of these plants, however, it is unlikely that he would have
confused it with a perennial species, although his diagnosis makes no mention
of its different habit; and the species has therefore been provisionally
adopted. |
3. RUPICAPNOS LONGIPES, Pomel, Mat. Fl. Atlant. 16 (1860) ; Nouv. Mat.
Fl. Atlant. i. 245 (1874). Fumaria longipes, Coss. & Dur. in
Bull. Soc. Bot. Fr. ii. 305 (1855; ; Batt. & Trabut, Fl. d'Alger. i. 26
(1888) ; F. numidica var. longipes, Coss. Comp. Fl. Atlant, ii. 82
(1883-1887). (PI. 15, fig. 1.)
Easicc, Balansa, Pl. d'Algérie, 1853, Mnechounés, near Biskra, in Hb.
Mus. Paris (partim), ut F. longipes !
Rupicapnos, ut videtur, annua, radice longá et caule suberecto (5-10 em.
alto) brevissime ramoso prodita. Folia caulina, haud crassa, omnia
longe petiolata, eum petiolo 5-12 cm. longa, deltoidea, foliolis infimis
breviter petiolatis, reliquis subsessilibus, et segmentis secundariis in laci-
nias oblongas lanceolatasve aeutas irregulariter fissis 2-pinnatisecta subquinata,
vel primaria simpliciora pinnata. acemi corymbiformes, pauci-(3—8-)
flori, cum pedunculo (ad 35 mm. longo) foliis multo breviores. Bractew
1-2 mm. longs, ovales, acute, subintegre ; pedicelli fructiferi fere fili-
formes, apice incrassati, flexuosi, longissimi (usque ad 60 mm. longi).
Sepala cirea. 2 mm. longa, 1 mm. lata, ovalia, peltata, acuta, subintegra
vel leviter crenato-dentata, nervo dorsali viridiusculo alba. Corolla
GENERA FUMARIA AND RUPICAPNOS. SoD
6-8 mm. longa, alba ; petalo superiore apice marginibus latis subpatentibus
ultra carine apicem productis obovato-oblongo, truncato vel retuso,
carinà crass, viridi vel purpurascente, calcare longiusculo | deflexo ;
petalo inferiore (quam superius latiore) apice marginibus latis patentibus in
limbum suborbicularem obtusissimum vel retusum abrupte dilatatis et carina
viridi predito, basi subsaccato ; petalis interioribus fere rectis apice plane
alatis atropurpureis. — Fructüs maximi, sine rostro 3:5 mm, longi, 3 mm. lati,
subrhomboidel, cum rostro magno (1 mm. longo), latissimo, striato, quadrangu-
lato, compresso subacuti et inferne attenuati, conspicue compressi et acute
carinati, pericarpio satis tenui præditi, siccitate preter rostrum. omnino
grosse et acute tuberculato-rugosi vel muricati.
R. longipes in fissuris umbrosis rupium ad Mnechounés, prope Biskram, ad
Elkantaram (Cosson) et in Montium Aurasiorum regione montaná inferiore
prope Beni Souik (Cosson) in provincià Constantinà Algeriw invenitur.
This species was originally described by Cosson & Durieu (l. c.) as a
Fumaria, and was founded on exsiecata collected by Balansa near Biskra and
by Gallerand near Beni Souik, both in the southern part of the province of
Constantine. The annual nature of the plant and its floral characters were
defined in the original description, but no mention was made of its peculiar
fruit ; and later, in the Compendium, it was reduced by Cosson to a variety
of his perennial F. numidica. Pomel admitted it as a species, although the
actual plant was unknown to him.
The sheet of Balansa’s Biskra plant named Fumaria longipes in Herb.
Mus. Paris contains four specimens, two of which agree with Cosson’s
diagnosis, so far as it goes, and possess the remarkable fruit described above.
From these the foregoing deseription and the accompanying figure have
been taken. The other two specimens, which are not alike, possess some-
what similar foliage but are clearly shown by their widely different fruits to
be specifically distinct. These are separated under the names of X. preter-
missa and R. sublevis.
4. RUPICAPNOS PRETERMISSA, sp. nov. (PI. 14, fig. 2.
Zæsice. Balansa, Pl. d'Alger. 1853, Mnechounés, near Biskra, in Hb.
Mus. Paris (partim), ut Fumaria longipes !
Rupicapnos annua, Aabitu foliisque R. longipedis sed, ut videtur, planta
minor. Racemi corymbiformes, pauct-(3-5-) fori, cum pedunculo (sub-20 mm.
longo) foliis plus duplo breviores. Bractew 1-1:5 mm. longe, oblonge,
apice truncate sepius tridentate ; pedicelli fructiferi filiformes apice paululum
incrassati, flexuosi, usque ad 30 mm. longi. Sepala cirea 1 mm. longa, 15 mm.
lata, suborbicularia, peltata, subacuta, irregulariter crenato-dentata, alba.
Corolla 6-7 mm. longa, alba ; petalo superiore apice marginibus latinsculis
subpatentibus oblongo, truncato emarginatove, cariná crass’ purpurascente,
calcare gracili longiusculo deflexo; petalo inferiore apice marginibus latiusculis
B
224 MR. H. W. PUGSLEY : A REVISION OF THE
patentibus spathulato, obtusissimo vel retuso, carina. viridi priedito, basi
verisimiliter subsaccato ; petalis interioribus fere rectis apice modice alatis
atropurpureis. Fructis majusculi, cum rostro 3 mm. longi, 2-2:25 mm. lati,
ovuto-elliptici, superne in rostrum latiusculum compressum angustati subaeuti,
inferne abrupte angustati (haud attenuati), valde compressi et acute carinati,
siccitate omnino tuberculato-rugulosi.
Hiec species X. lonyipedi plane affinis est, sed per pedicellos fructiferos
subduplo breviores eum bracteis truncatis, per sepala minora magis dentata,
per corollas angustiores et per fructüs subduplo minores, leviores, inferne
haud attenuatos satis differt.
R. prvtermissa in fissuris umbrosis rupium ad Mnechounés prope Biskram,
in provincia Constantinà Algerie, ubi cum Æ. longipede crescit, a cl. Balansa
inventa est.
This rare and obseure plant has been described and figured from the
single specimen included in Balansa’s sheet of Fumaria longipes in Herb.
Mus. Paris, to which reference has already been made. Its points of
distinction from R. longipes were apparently overlooked both by Balansa and
by Cosson, owing probably to the similarity of its foliage and the incon-
spicuous nature of its few-flowered clusters of small, pale flowers. It
appears to possess, however, a number of minor differences in addition to
the widely divergent fruit.
9. livricAPNOS SUBLEVIS, sp. nov. (Pl. 15, fig. 2.
Hesicc. Balansa, Pl. d'Alger. 1852, Mnechounés, near Biskra, in Hb. Mus.
Paris (partim), ut Fumaria longipes !
Rupicapnos verisimiliter annua, caule suberecto (circa 4 em. alto)
brevissime ramoso priedita. Folia pleraque caulina, haud crassa, longe
petiolata, cum petiolo subalato 4-6 em. longa, ovalia vel subdeltoidea, foliolis
infimis subpetiolatis reliquis sessilibus omnibus paulo confertis, et segmentis
secundariis in lacinias lanceolatas «cutus irregulariter fissis 2-pinnatisecta
subquinata, Racemi corymbiformes, pauci-(4—8-) fori, cum pedunculo (ad
15 mm. longo) foliis subduplo breviores. Bractew 15-2 mm. longi, oblonga,
acutz, interdum apice sparsim dentate ; pedicelli fructiferi filiformes apice
inerassati, flexuosi, usque ad 40 mm. longi. Flores haud visi. — Fructás
modici, cum rostro 2 mm. longi, 1175-2 mun. lati, anguste obovati, superne in
rostrum parvulum breviter angustati, subacutz, inferne multo attenuati, satis
compressi et plane carinati, siccitate ¿infra rostrum omnino leviter tuberculato-
rugulosi.
Hee species, cujus flores non visi sunt, habitu ad X. longipedem accedit
sed folia ut in X. Cossonit fissa et fructûs rugulosos modicos nec maximos
monstrat, K. pretermissa pedicellis brevioribus, bracteis truncatis,
fructibus magis compressis ovato-ellipticis potius quam obovatis etiam
differt.
GENERA FUMARIA AND RUPICAPNOS. 335
R. sublevis in fissuris rupium ad Mnechounés prope Biskram in provincia
Constantináà Algerie, ubi cum œX. longipede et R. pretermissd crescit,
invenitur.
The diagnosis of this rare plant is taken from the solitary example on
Balansa’s sheet of Fumaria longipes in Herb. Mus. Paris, which unfortunately
now shows no flowers. Except for the slightly winged petioles, its foliage
bears a resemblance to that of R. Cossonit rather than R. longipes, and its
narrow, finely rugose fruit is very distinct.
SgcTI0 II. SARCOCAPNOIDES.
Plante perennes, decumbentes. Racemi multitlori. Flores parvi vel
parvuli, 4-10 mm. longi, albi vel roseo-albidi; petalis exterioribus quam
interiora fere recta apice modice alata atropurpurea conspieue longioribus et
latioribus, in limbos patentes suborbiculares apice dilatatis ut in genere
Sarcocapnos ; stylus fere malleiformis lobis curvatis subpatentibus. Fructtis
parvi.
RUPICAPNOS SARCOCAPNOIDES, Pomel, Mat. Fl. Atlant. 16 (1860) ; Nouv.
Mat. Fl. Atlant. i. 245 (1874). Fumaria sarcocapnoides, Coss. & Dur.
in Bull. Soc. Bot. Fr. ii. 306 (1855) ; F. numidica subvar. sarco-
capnotdes, Coss. Comp. Fl. Atlant. ii. 82 (1883-1887). (PI. 16.)
Zesice. Balansa, Pl. d'Alger. 1853, No. 995, in Hb. Mus. Paris (partim),
ut Fumaria corymbosa !
Rupicapnos perennis sed probabiliter a primo anno florens, caudice brevi
crassiusculo ramoso decumbente et caulibus brevissimis preedita, Folia
pleraque subradiealia,. plus minusve glauca, haud crassa, 6-10 cm, longa
(petiolo incluso), longe petiolata, oLlongo-deltoidea, foliolis breviter petiolatis
trifido-flabellatis et segmentis secundariis in lobos oblongo-ellipticos subacutos
irregulariter fissis quinato-pinnata. Racemi corymbiformes, circa 20-jlori,
supe fureati, cum pedunculo gracili foliis subduplo breviores. Bractew circ
15 mm. longs, late ovales, cuspidate, apice dentate ; ad tg fructiferi
capillares apice paululum incrassati flexuosi, infimi usque ad 25 mm. longi.
Sepala circa 1 mm. longa, 15 mm. lata, late Bin vel suborbicularia, peltata,
acuta, dentata, alba. Corolla minima, 4-5 mm. longa, carinis crassis viridibus
alba ; petalis exterioribus quam interiora multo (saltem 1 mm.) longioribus et
latioribus, superiore marginibus latissimis patentibus ultra carin: apicem
productis apice in limbum magnum suborbicularem vel seepius obcordatum
dilatato, calcare brevi (1 mm. longo) obovato recto ; inferiore, ut superius,
apice in limbum patentem suborbicularem dilatato, basi haud saccato ; petalis
interioribus vectis apice plane dilatatis modice alatis atropurpureis. Stylus
fere malleiformis lobis curvatis subpatentibus. Fructús parvi, sine mucrone
circa 2 mm. longi et 1:5 mm. lati, oblongo-obovali, cum mucrone brevi obtusi
336 MR. H. W. PUGSLEY : A REVISION OF THE
et inferne multo angustati, satis compressi et plane carinati, siccitate omnino
dense tuberculato-rugulosi.
Hee species distinetissima per petala exteriora interioribus longiuscule
longiora et latiora affinitatem conspicuam eum genere Sarcocapnos monstrat
sed fructum monospermum tuberculatum generis Rupicapnos habet.
K. sarcocapnoides regionem montanam Algerie australis in provincia
Constantinà (Djebel Tougour prope Batnam) habitat, ubi in fissuris rupium
crescit.
This plant was originally described by Cosson in 1855 from a specimen
observed among the exsiecata sent out by Balansa under the number 995
and otherwise referred to Fumaria numidica, Cosson noting that the plant
constitutes a trie passage between the genera Fumaria and Sarcocapnos
owing to the form of its corolla. Later, however, in Illustr. Fl. Atlant. i. 15
(1882), Cosson remarks that F. sarcocapnoides appears to be a shade-form
only of F. numidica, adding that shade-forms of Sarcocapnos similarly
deviate from the type; and in the Compendium it becomes a sub-variety of
.F. numidica. It is not easy to understand how shade-conditions could
transform the corolla-characters of F. numidica into those of F, sarcocapnoides,
developing the margins of the outer petals while rendering depauperate the
wings of the inner ones, and as moreover it is not only in their flowers that
the two plants differ, the earlier view of Cosson is here followed.
R. sarcocapnoides seems to bea very rare plant, and it is not known to have
been collected except by Balansa, from whose specimen at Paris the above
diagnosis is taken and the accompanying figure drawn. It was unknown
to Pomel, who adopted the species from Obaban:
[7. Rurrcarxos Repovpiana, Pomel, Nouv. Mat. Fl. Atlant. fasc. ii. 379
(1876) ; n. v. Fumaria Reboudiana, Batt. & Trabut, Fl. d'Alger. i. 26
(1888).
Rupicapnos perennis, fragilis, X. sarcocapnoidi affinis. Folia viridia,
foliolis in segmenta cuneiformia vel oblonga, obtusa fissis quinato-pinnata.
Bractew oblongw ; pedicelli apice incrassati. Sepala obovata vel suborbicularia,
valde dentata. Corolla 9-10 mm. longa, roseo-albida ; petalis exterioribus
marginibus latis patentibus ¿n limbos suborbiculares nne dilatatis, superioris
calcare obovato longiusculo, inferioris basi valde saccatá ; petalis interioribus
apice atropurpureis haud conspicue. alatis. — Fructás parvi, obovati, breviter
mucronati, in sicco tuberculato-rugosi.
R. Reboudiana rupes ad Bou-Taleb (Reboud) Algeriz australis habitat.
This plant appears distinct from Pomel’s brief description, and is retained
as a species although no material of it has been seen. It is evidently, as
stated by Pomel, a rélalively large-flowered ally of R. sarcocapnoides. |
GENERA FUMARIA AND RUPICAPNOS. 337
Srorio III. TRIPTERYX.
Plante perennes decumbentes. Racemi multiflori. Flores parvi vel
parvuli, 4-10 mm. longi, albidi, plus minusve luteo-tincti; petalis exterioribus
quam interiora fere recta apice ut interdum exteriora superant conspicue
alata, haud atropurpurea (nisi interne vel brevissime ad apicem ipsum). vix
longioribus, marginibus eorum apice patentibus sepe in limbos suborbieulares
dilatatis; stylus fureeeformis lobis adscendentibus. Fructüs parvi ad
majusculi.
8. RurrcAPNOs NuMIDICA, Pomel, Mat. Fl. Atlant. 16 (1860), ex parte ;
Nouv. Mat. Fl. Atlant. i. 243 (18:4). Fumaria numidica, Coss. &
Dur. in Bull. Soc. Bot. Fr. ii. 306 (1855), ex parte; Coss.
Illustr. Fl. Atlant. i. 15 (1882), ex parte; Comp. Fl. Atlant. ii. 81
(1883-1887), ex parte ; Batt. & Trabut, Fl. d'Alger. i. 25 (1888), ex
parte. (Pl. 9, fig. 6.)
Exsicc. Cosson, Constantine, 1853, in Hb. Mus. Paris, ut F. numidica!
Choulette, Fragm. Fl. Alger. Constantine, 1855,in Hb. Kew, ut F. numidica !
Soc. Dauph. No. 285, Constantine, 1574, Dr. Reboud !
Rupicapnos perennis sed interdum a primo anno florens, caudice brevi
crassiusculo ramoso decumbente tandem vestigiis petiolorum emarcidorum
donato et caulibus brevissimis preedita. Folia pleraque subradicalia, crassius-
cula, glaucescentia, 6-15 (raro ad 25) em. longa (petiolo incluso), longe
petiolata, subdeltoidea, foliolis 2-3-paribus haud confertis breviter petiolatis
et segmentis secundariis in lacinias satis remotas oblongas vel cuneatas
obtusas ssepe leviter mucronatas irregulariter fissis 3-3-pinnatisecta. Racemi
corymbiformes, densi, multi-(20-30-) fori, breviusculi, cum pedunculo (ad
4 em. longo) foliis saltem subduplo breviores. Bractew 1-15 mm. longe,
lanceolate, acuminatie, obscure denticulate ; pedicelli fructiferi fere capillares
apice incrassati flexuosi infimi usque ad 25 mm. longi. Sepala circa 15 mm.
longa, *75 mm. lata, oblongo-lanceolata, peltata, acuta, basin versus parce
dentata, albida. Corolla minima, circa 4 mm. (4-5 mm. ap. Pomel) longa,
carinis viridibus vel lutescentibus albida ; petalo superiore apice marginibus
latiusculis subpatentibus oblongo vel anguste obovato, obtusissimo vel truncato,
calcare minimo brevissimo (vix 1 mm. longo) recto ; petalo inferiore apice
marginibus latis patentibus sensim dilatato cuneato-obovato obtusissimo, basi
haud saccato; petalis interioribus fere rectis apice ald lata lutescente
rotundatá inter petalorum exteriorum margines quos paululum superat
subpatulá preeditis obscure atropurpureis. Fructûs modici, sine mucrone
2-75 mm. longi, 2 mm. lati, oblongo-obovati (obovati ap. Pomel), eum mucrone
brevi obtusissimi et inferne attenuati, satis compressi et plane carinati,
siccitate omnino dense tuberculato-rugosi.
R. numidica que per flores minimos apice dilatatos brevissime calearatos
338 MR. H. W. PUGSLEY : A REVISION OF THE
facile distinguenda est, in Algeria rupes caleiferas prope urbem Constantinam
et forsan alios locos habitat.
Pomel is here followed in restricting the Fumaria numidica of Cosson,
which includes several distinct plants collected in different Algerian stations,
to the form growing in the neighbourhood of Constantine, the first locality
cited by Cosson in his original diagnosis of F. numidica.
There is a flowerless specimen of this plant in Herb. Kew, collected by
Bové at Constantine in September, 1859, in which the leaves are more than
25 em. in length.
9. RUPICAPNOS COSSONIL, sp. nov.
Fumaria numidica, Coss. & Dur. in Bull. Soe. Bot. Fr. ii. 306 (1855),
ex parte; Coss. Illustr. Fl. Atlant. i. 15 (1882), ex parte; Comp.
Fl. Atlant. ii. 81 (1883-1887), ex parte ; Batt. & Trabut, Fl. d'Alger.
i. 25 (1888), ex parte.
Icon Cosson, Illustr. Fl. Atlant. i. tab. 9, figs. 1-11, ut F. numidica.
Ewsice. Balansa, Pl. d'Algor. 1853, No. 995, in Hb. Mus. Paris (partim),
Hb. Kew, Hb. Mus. Brit. et Hb. Manchester, ut F. corymbosa !
Rupicapnos perennis, eaudice brevi crasso ramoso decumbente tandem
vestigiis petiolorum emarcidorum donato atque caulibus brevissimis vel fere
nullis pr:edita. Volia pleraque radicalia, crassa, plus minusve glauca, cum
petiolo 5-10 em. longa, longe petiolata, oblonga vel ovalia, foliolis (2-3-
paribus) satis confertis, breviter petiolatis vel supremis subsessilibus, et
segmentis secundariis in lacinias subdivaricatas oblongas obtusas obscure
mucronatas vel acutinsculas irregulariter fissis 2-3-pinnatisecta. Racemi
subeorymbiformes, meu/ti-(15-30-) fori, pedunculis paulo breviores, cum
pedunculo (ad Sem. longo) folia sulerquantes vel in formis umbrosis breviores.
Bracter 1-175 mm. longi, lanceolata, acute vel acuminatie, parce serrulata;
pedicelli fructiferi graciles (Fere filiformes), anguste clavati, apice incrassati,
flexuosi, infimi usque ad 25 mm. longi. Sepala 1-15 mm. longa, ‘5-1 nun.
lata, ovalia vel ovata, peltata, subacuta, plus minusve crenato-dentata, nervo
dorsali viridiusculo albida. Corolla parva, 4-5 mm. longa, haud gracilis,
preter carinas virescentes albida; petalo superiore apice marginibus latius-
culis patentibus anguste obovato. obtusissimo vel etiam retuso, calcare satis
longo adscendente paululum curvato ; petalo inferiore lato marginibus latis
patentibus apice sensim dilatato cuneuto-obovato obtusissimo (apice quam
petalum superius paulo latiore), bast Laud saccato ; petalis interioribus fere
rectis apice vix coalitis alâ laté lutescente proesertim ad apicem dilatatà inter
petalorum exteriorum margines quos paulo superat subpatulà præditis
verisimiliter interne et apice ipso brevissime purpureo-tinctis. Stylus
furezeformis. — F'ructs modici, sine mucrone 2-2:5 mm. longi, 1175-2 mm.
lati, sulbrotundo-obovati. vel obovati, cum mucrone distincto obtus?, plane com-
pressi et valde carinati, siccitate omnino dense tuberculato-rugosi submuricati.
viis. M 1 "^ Ge o. ae s GIU METUT ETE TL TENETE EAE TT | Sos
GENERA FUMARIA AND RUPICAPNOS. 339
Heee species Æ. numidien affinis est sed foliolis laciniisque minus remotis,
racemis relative longioribus, pedicellis crassioribus, corollis majoribus longius
calearatis, fructibus cum mucrone longiore latioribus differt.
R. Cossonii fissuras rupium ad Djebel-Tougour, prope Datnam, in provincia
Constantiná Algerie habitat.
Cosson’s original Fumaria numidica is partly founded on this plant and
the plate in his * IHlustrationes Flore Atlantice ^ is evidently drawn from it.
As Pomel, however, has adopted this specific epithet for the form found at
Constantine, and Dalansa's plant from Batna appears to be an essentially
distinct species with several points of difference, a new name becomes
necessary, and it therefore seems fit to commemorate Cosson, who was the
first to distinguish and describe the small-flowered species of the genus.
10. RUPICAPNOS TENUIFOLIA, Pomel, Nouv. Mat. Fl. Atlant. 1. 244 (1874).
Fumaria tenuifolia, Batt. & Trabut, Fl. d'Alger. i. 26 (1888);
F. numidica, Coss. Comp. Fl. Atlant. ii, 81 (1883-1887), ex parte.
Egsice. Dourgeau, PI. d'Alger. 1856, No. 242, ut F. numidica !
Rupicapnos perennis caudiee brevi crassiusculo ramoso decumbente et
caulibus brevissimis ut in Æ. Cossonit. Folia pleraque radicalia, vix crassius-
cula, plus minusve glauca, cum petiolo longo gracilescente 5-20 cm. longa,
anguste oblonga, foliolis sepius 3-paribus breviter petiolatis et segmentis
secundariis (interdum subflabellatis) in lacinias confertas lineari-oblongas,
vel oblongas in foliis primariis, acutas vel obtusiusculas (parvas obovatas
obtusas ap. Pomel) irregulariter fissis. 2-3-pinnatisecta. Racemi corymbi-
formes, densiuseuli, nulti-(20—40-) fori (pauciflori ap. Pomel), cum pedunculo
gracili (ad 5 em. longo) joliis subduplo breviores nisi in formis nanis.
Bractew 1-15 mm. longi, oblonge, acuminate ; pedicelli fructiferi capillares
apice inerassati flexuosi infimi ad 20 mm. longi. Sepala circa 1:5 mm. longa,
*5— 15 nun. lata, lanceolata (ovata ap. Pomel), vix peltata, acuta, parce
denticulata, albida. Corola minima, 4-4°5 mm. longa, prster carinas
lutescentes vel virescentes albida; petalo superiore apice marginibus latis
patentibus in limbum suborbicularem obtusissimum dilatato, calcare breviusculo
satis gracili fere recto sed apice dilatato, [nectario paulo arcuato calcar
medium attingente ap. Pomel] ; petalo inferiore apice marginibus latis in
limbum suborbicularem obtusissimum vel truncatum abrupte dilatato, basi haud
saccato ; petalis interioribus fere rectis apice alâ latá lutescente rotundatà
patulà petalorum exteriorum margines subæquante præditis haud atro-
purpureis. Fructûâs parvi, sine mucrone 1:75-2 mm. longi, 1:5-L:75 mm.
lati, subrotundi vel subrotundo-obovati (obovati ap. Pomel), cum mucrone
gracili obtusissimi, paululum compressi et obscure carinati, siccitate omnino
dense tuberculato-rugosi.
Hee species duabus priecedentibus (X. numidicw et R. Cossonii) atlinis est,
sed per folia graciliora magis dissecta, per petala exteriora in limbos
340 MR. H. W. PUGSLEY : A REVISION OF THE
suborbiculares dilatata ct per fruetüs minores gracile mucronatos separanda
est.
R. tenuifolia regionem montanam Algeriw australis in provincia Oranensi
ad Itimam (Pomel), prope Stitten et forsan ad alios locos habitat.
The diagnosis of R. tenuifolia has been taken from Bourgeau’s exsiccata
at Kew and in Herb. Manchester, which, while differing in some particulars
from Pomel’s original description, as indicated, seem to be essentially
conspecific although perhaps varietally separable. The relationship of the
two forms is indeterminable in the absence of authentic material of the
Itima plant seen by Pomel, which appears to differ chiefly by its broader
leaf-segments, fewer-flowered racemes, broader sepals, and less rotund
fruits.
11. RuricAPNOs CAPUT-PLATALEE, Pomel, Nouv. Mat. Fl. Atlant. i. 244
(1874). Fumaria caput-plataler, Batt. & Trabut, Fl. d’Alger. i. 2:
(1888). (PI. 9, fig. 7.)
Eusice. Murbeck, Iter Alger.-Tunetanum, 1896, Dyr-el-Kef, Tunetia media,
in Hb. Mus. Brit., ut F, numidica var. longipes !
Rupieapnos perennis, glauca, caudice crassiusculo et caulibus abbreviatis
ut in Æ. numidied. Folia pleraque subradicalia, crassa, eum petiolo
longissimo 10-15 em. longa, subdeltoidea, foliolis flabellatis infimis petiolatis
reliquis ssepius fere subsessilibus in segmenta remota cuneiformia vel obovata
obtusissima rarius obscure mucronata fissis irregulariter. 2-pinnatisecta sub-
quinata. Racemi primo corymbiformes cito elongati, densi, conspicue multi-
(30—50-) fori, cum pedunculo ad 8 em. longo folia paulo superantes vel
parum breviores. Bractew 1-1-5 mm. longi, lineari-lanceolate, acuminate,
subintegro ; pedicelli fructiferi Jiliformes apice inerassati longissimi (infimi
usque ad 50 mn. longi). Sepala circa 2 mm. longa, 1-1:5 mm. lata, ovata,
peltata, acuminata, integriuscula vel basi parce dentata, albida. Corolla
parva, 4-5 mm. longa, preter carinas virescentes albida ; petalo superiore
apice marginibus latis patentibus late obovato vel suborbiculari truncato vel
obtusissimo, calcare breviusculo fere recto, [nectario brevissimo calcar vix
ineunte ap. Pomel] ; petalo inferiore apice marginibus latissimis patentibus
in limbum suborbicularem truncatum dilatato, basi haud saccato ; petalis
interioribus fere rectis apice alâ latissimá lutescente rotundatà in spatulee
forma inter petalorum exteriorum margines quos longe superat patente
preeditis interne obscure atropurpureis. Fructis mediocres, sine mucrone
circa 2°25 mm. longi et 2 mm. lati, fere. elliptici, eum mucrone obtusiusculi,
satis compressi sed parum carinati, siccitate omnino dense tuberculato-
rugosi.
R. caput-platalew valde R, numidiee affinis, sed per alarum latitudinem in
petalis interioribus que petalorum exteriorum margines longe superant
mirabilis est. Foliorum segmentis cuneiformibus obtusioribus, racemis
GENERA FUMARIA AND RUPICAPNOS. 341
conspicue multifloris eum pedicellis fructiferis ad 50 mm. longis et fructibus
ellipticis ab hujus sectionis speciebus aliis insuper differt,
Hæc species ad El Ghicha in regione montaná Djebel Amour Algerim
australis (Pomel) primum inventa est atque prope Dyr-el-Kef Tunetim
medize et forsan in aliis locis crescit.
This remarkable plant has been described from Murbeck’s exsiccata in
Herb. Mus. Brit., collated with the diagnosis of Pomel, with which it
substantially agrees. The fanciful name “caput-platalew” or “ spoonbill’s
head” is perhaps not inapt owing to the great apical dilation of the inner
petals.
Murbeck’s reference of his exsiccata to Fumaria numidica var. longipes,
Cosson, is probably due to the great length of the fruiting pedicels, which
characterizes this plant in common with X. longipes. In every other respect,
however, R. caput-platalee is widely different from the annual species
distinguished by Cosson.
12. RUPICAPNOS EROSA, Pomel, Nouv. Mat. Fl. Atlant. i. 243 (1874).
Fumaria erosa, Batt. & Trabut, Fl. d'Alger. i. 26 (1888) ; F. numidica,
Coss. & Dur. in Bull. Soc. Bot. Fr. ii. 306 (1855), ex parte ; Coss.
Illustr. Fl. Atlant. i. 15 (1882), ex parte; Comp. Fl. Atlant. ii. 81
(1883-1887), ex parte.
Exsice. Kralik, Pl. Alger. 1858, No 3, ut F. numidica ! Paris, Iter Bor.-
Afric. 1886, No. 10, ut F. numidica! Chevallier, Pl. Saharz Alger. 1897,
No. 147, ut F. numidica !
Rupicapnos perennis R. numidice habitu sed caulibus nonnunquam
erassioribus. Folia pleraque subradicalia, crassiuseula, glaucescentia, cum
petiolo longo 3-15 em. longa, oblonga, foliolis (vulgo 3-paribus) sepius
brevissime petiolatis et segmentis secundariis in /ohos obovatos vel rotundatos
obtusissimos (vix mucronatos) divarieatos sape incisos aut rarius oblongos
subacutos fissis (pinnulis subflabellatis et earum lobis seepe alternantibus)
irregulariter 2-3-pinnatisecta. Racemi corymbiformes, 10—20-flori, cum pedun-
culo 2-5 em. longo foliis multo breviores vel in formis apricis nanis folia sub-
wquantes, Bractew 1-15 mm. longe, lineari-lanceolat« , acuminatee ; pedicelli
fructiferi filiformes apice abrupte inerassati flexuosi infimi usque ad 25 mm.
longi. Sepala 1:5-2:5 mm. longa, 1-1:5 mm. lata, ovata vel oblonga, peltata,
acuminata, plus minusve dentata, albida. Corolla 6-10 mm. longa, quam in
hujus sectionis aliis speciebus relative angustior, preter carinas crassas
apicem haud attingentes lutescentes vel pallide virescentes alba ; petalo
superiore apice marginibus latiusculis patentibus oblongo-obcordato retuso,
caleare longiusculo deflexo ad basin angustato ; petalo inferiore marginibus
latis patentibus in limbum suborbicularem retusum abrupte dilatato, basi
obscure subsaccato ; petalis interioribus fere rectis apice ald latá lutescente
presertim ad apicem dilatatà angulatà subpatula petalorum | exteriorum
342 MR. H. W, PUGSLEY : A REVISION OF THE
margines subequante præditis verisimiliter interne nec externe purpureo-
tinctis. — Fructás parvuli, sine mucrone 2-2:25 mm. longi et circa 2 mm. lati,
subrotundo-obovati vel subrotundi, cum mucrone breviusculo quandrangulato
obtusissimi et inferne in stipitem obscurum angustati, paulo compressi et
plane carinati, siccitate omnino dense tubereulato-rugosi.
B. major, var. nov.
tesice. Clary, no. 33, Ghada de Ghernenta, 1888, in Hb. Mus. Paris, ut
Fumaria numidica (Rupicapnos erosa, Pomel) !
Planta multo major habitu laxiore. Folia haud crassa, viridia, cum petiolo
longo gracili 20-30 em. longa, subquinata. Racemi densi, multi-(15-30-)
Hori, cum pedunculo (ad 8 em. longo) folia subduplo breviores, Bractew
minima, circa 1 mm. long, ovato-lanceolatw, acuminate, obscure denticulatz ;
pedicelli fructiferi capillares, apice incrassati, infimi ad 40 mm. longi. Sepala
parva, 1-1°5 mm. longa, *75-1 mm. lata, acuta. Corolla 8-10 mm, longa.
Fructûs majusculi, sine mucrone circa 2*5 mm. longi et 2:75 mm, lati,
subrotundi, siccitate grosse et dense tuberculato-rugosi. Aliter ut in typo.
Hee planta ab hujus sectionis aliis speciebus per foliorum 2-3-pinnati-
sectorum lobos latos divaricatos, per corollas angustiores pallidiores multo
majores caleare longiore petaloque inferiore subsaccato præditas, et per
fruetüs subrotundos breviter mucronatos distinguenda est.
KR. erosa regionem montanam Djebel Amour Algeriw australis prope
El Beida (ap. Pomel) et ad Laghouat habitat; varietas major hactenus ad
Ghada de Ghernenta solum inventa est.
The oldest specimens of this plant, colleeted at Laghouat by Kralik, and
also others in Herb. Kew received from Reboud, are late gathered and
more or less exhausted individuals bearing small flowers, but are none the
less clearly conspecific with the much finer examples obtained more recently
in the same district by Paris and Chevallier. It is noteworthy that the
foliage of the weaker plants shows relatively narrow and acute segments,
The variety major is a much larger plant, with finer flowers, almost
recalling the section Callianthos. It appears, however, to possess all the
essential features of the Laghouat form, except that its bracts and sepals
are smaller and broader,
Srerro IV. CALLIANTHOS.
Plante perennes decumbentes eis sectionum aliarum vulgo majores,
Racemi multiflori. Flores magni, 12-16:5 mm. longi, albidi vel pallide
purpurei; petalis exterioribus quam interiora apice sursum curvata modice
alata atropurpurea vix longioribus, marginibus eorum apice deflexis patentibus
vel sursum recurvatis in limbos suborbiculares nunquam dilatatis : stylus
malleiformis lobis divaricatis, Fructûs modici ad maximi,
ESL dE s 2a T Ec
GENERA FUMARIA AND RUPICAPNOS. 343
SunsEcrIo AFRICANA.
Foliorum lobi relative angusti. Flores albidi.
13. RtPICAPNOS AFRICANA, sp. nov. (non R. africana, Pomel).
Fumaria africana, Lamarck, Encycl. ii. 569 (1788) ; Coss. & Dur. in
Bull. Soc. Bot. Fr. ii. 305 (1855), ex parte ; Coss. Comp. FI. Atlant. ii,
80 (1883-1887), ex parte; F. corymbosa, Desfontaines in Act. Soc.
Hist. Nat. Paris, i. 26 (1792), et Fl. Atlant. ii. 124 (1800), non
F. corymbosa, Hammar ; Rupicapnos graciliflora, Pomel, Nouv. Mat.
Fl. Atlant. i. 241 (1874) : Fumaria graciliflora, Batt. & Trabut, Fl.
d'Alger. i. 25 (1888).
Icon. Desfontaines, l.c. i. tab. 6, ut F. corymbosa.
Egxsice. Bourgeau, Pl. d'Alger. 1856, No. 181, Nedroma, in Hb, Mus.
Paris (fructibus plerisque exclusis) et in herb. aliis (partim), ut F. africana !
Rupicapnos perennis, caudice erassiuseulo ramoso decumbente siepe paulo
elongato tandem vestigiis petiolorum emarcidorum donato caulibusque semper
brevibus. Folia pleraque subradiealia, plus minusve glaucescentia, 10-25
em. longa (petiolo gracilescente incluso), longe petiolata, oblongo-deltoidea,
foliolis ambitu fere ovatis vulgo breviter petiolatis et segmentis secundariis
subflabellatis in lacinias oblongas vel lineari-oblongas acutas rarius obtusas
mucronatas irregulariter fissis 2-pinnatisecta subquinata, Racemi corymbi-
formes, 15-20-/lori, cum pedunculo (ad 6 em. longo) foliis subduplo breviores.
Bractew 1°5-2°5 mm. longze, lanceolate, tenuiter acuminate (denticulate ap.
Pomel) ; pedicelli fructiferi filiformes apice incrassati, infimi ad 50 mm. longi.
Sepala circa 8 mm. longa, 9 mm. lata, ovalia vel ovato-lanceolata, peltata,
acuminata, leviter dentata vel ad basin dentibus angustissimis sublaciniata
(subintegra ap. Pomel), preeter nervum dorsalem latum viridiusculum albida.
Corolla maxima, 14-16 mm. longa, gracilis, carinis viridibus albida ; petalo
superiore apice marginibus parum dilatatis subpatentibus vel deflexis non-
nunquam leviter purpureo-tinetis anguste oblongo obtusissimo, calcare longo
(circa 5 mm.) deflexo basi angustato apice rotundato, [nectario longissimo
haud ineurvato ap. Pomel] ; petalo inferiore marginibus patentibus apice vix
dilatatis lineari-oblongo, obtusiusculo, ad basin ipsam inconspicue (valde ap.
Pomel) gibboso-saceato ; petalis interioribus apice sursum curvatis modice
alatis atropurpureis. Stylus malleiformis. — /ructis modici, sine mucrone
circa 2:5 mm. longi, 2 mm. lati, obovato-elliptici (elongato-elliptiei ap. Pomel),
cum muerone brevissimo obtusissimi vel obtusi, inferne in stipitem angustum
obseurum breviter angustati, paulo compressi et acute carinati, siccitate
omnino tenuiter tuberculato-rugosi.
Hee species prope Mascaram (Desfontaines) et in rupibus dolomiticis
ad Nador prope Tiaret (Pomel) in parte orientali atque prope Tlemcen
(Desfontaines) et ad Nedromam in parte occidentali provinciæ Oranensis
344 MR. H. W. PUGSLEY: A REVISION OF THE
in Algeria crescit; etiam in locis aliis hujus provincie sine dubio videnda
est.
This plant was discovered by Desfontaines prior to 1788 near Tlemcen and
Mascara, and specimens of it were brought to France for cultivation. Its
earliest description appears in Lamarck's Encyclopedia under the name of
Fumaria africana, and four years later it was re-described by its discoverer
as F. corymbosa, Both of the descriptions are in some detail and in. virtual
agreement, and with these and Desfontaines’ very fair figure, the diagnostic
characters of the species can be determined with some accuracy. The plant
is evidently one with rather finely cut leaf-segments, ovate sepals, slender
whitish corolla with a long spur, and slightly pointed fruits. It is not known
whether any original specimen of Lamarck’s or Desfontaines’ is now in
existence.
In 1855 Cosson, in establishing the section Petrocapnos of Fumaria, again
briefly describes this species, but extends its definition to include the purple-
flowered plant occurring about Oran; and Hammar, in his Monograph of
Fumaria, bases his description on the Oran form.
In Pomel’s revision of his genus Rupicapnos (Nouv. Mat. Fl. Atlant. i.
p. 240) six species of this group are established, and the name R. africana,
with which Fumaria africana Lamarck is identified, is restricted to a
purple-flowered form, characterized by very broad leaf-segments, large,
orbicular sepals, and a very short spur to the upper petal. This plant is
said to grow at Garrouban, Tlemcen, Gorges de la Tafna, Nedroma, and
Oran, Tlemcen being one of Desfontaines’ stations. Only one undoubted
example of Pomel’s plant, however, obtained in the first-named of these
localities, has come under observation, the exsiccata examined from Tlemeen,
Nedroma, and Oran being elearly different.
It is not easy to see what led Pomel to apply the epithet africana to this
apparently uncommon purple-flowered form rather than to Desfontaines
white-flowered plant, but it is likely that when Pomel’s account of these
plants was written, he had not been able to consult the works of Lamarck
and Desfontaines, a reference to which would have shown the distinctness of
the original X. africana from the form to which he restricted this specific name,
As Article 48 of the current Rules of Nomenclature requires that original
specific epithets must be retained (or restored) for the species to which they
were first given, it becomes necessary reluctantly to differ from Pomel
and to regard his Rupicapnos africana as a stillborn name, transferring
it to the plant originally described by Lamarck. This appears to be
conspecific with Pomel's X. graciliflora.
The above diagnosis of this species is based on Bourgeau’s sheet of
F. africana from Nedroma (No. 181) in Herb. Mus. Paris (excluding the
detached fruits, which are a mixture), together with the small examples
on the sheets of the same set in Herb, Manchester, Herb. Kew, and Herb.
GENERA FUMARIA AND RUPICAPNOS 345
Mus. Brit. These have been collated with the original accounts of Fumaria
africana, F. corymbosa, and Rupicapnos graciliflora by Lamarck, Desfontaines,
and Pomel respectively.
The admixture of this species and of the purple-flowered Oran form on
Bourgeau’s sheets of F. africana may be taken as evidence that these two
plants grow in company at Nedroma, as has already been seen sometimes to
happen in the case of some of the small-flowered species.
14. RUPICAPNOS DECIPIENS, sp. nov.
Fumaria corymbosa, Boissier, Voy. Bot. Esp. ii. 19 (1839-1845), non
Desfontaines ; F. africana, Willkomm & Lange, Fl. Hisp. iii, 878
(1880), non Lamarck; Coss. Comp. Fl. Atlant. ii, 80 (1883-1887), ex
parte.
Egsice. Huter, Porta & Rigo, Iter Hisp. 1879, No. 516, ut F. africana !
Reverchon, Pl. d'Andalousie, 1850, Grazalema, ut F. africana ! Porta & Rigo,
Iter IV Hisp. 1895, No. 23, ut F. africana !
Rupicapnos perennis, glauca, radice longissimo, caudice crassiusculo
ramoso decumbente et caulibus semper brevibus ut in R. africand. Folia
pleraque subradicalia, glaucescentia, plus minusve carnosa, 10-25 em. longa
(petiolo incluso), longissime petiolata, deltoidea, foliolis subdeltoideis (duobus
infimis multo majoribus petiolatis reliquis seepissime subsessilibus) et seg-
mentis secundariis subflabellatis in lobos oblongos ellipticos vel euneatos acutos
vel ohtusos vix mucronatos irregulariter fissis 2-pinnatisecta subquinata.,
Racemi corymbiformes, 70—23-/lori, cum pedunculo (ad 5 em. longo ssepius
multo breviore) foliis subduplo breviores. Bractew 2-3 mm. longs, albido-
virescentes, oblong, acute vel acuminate, interdum sparsim serrulatze ;
pedicelli fructiferi graciles, apice abrupte incrassati, flexuosi, longissimi
(infimi ad 65 mm. longi). Sepala 2°5-4 mm. longa, 2-3 mm. lata, late ovata
rarius suborbicularia, peltata vel cordata, acuta, basin versus leviter crenato-
dentata, nervo dorsali lato viridiusculo albida. Corolla maxima, 12-16 mm.
longa, carinis viridiusculis albida ; petalo superiore marginibus apice plane
dilatatis subpatentibus sæpe pallide purpurascentibus fere oblongo vel oblongo-
obovato, obtusissimo vel etiam retuso, cariná apicem versus attenuata, calcare
modico (cirea 4 mm. longo) vix eurvato apice rotundato; petalo inferiore
marginibus patentibus apice valde dilatatis spathulato obtusissimo, ad basin
ipsam smpius gibboso-saccato ; petalis interioribus sursum curvatis apice
modice alatis atropurpureis. — Fructüs maximi, sine mucrone 3-825 mm.
longi, cirea 3 mm. lati, subrotundo-obovati vel subrotundi, cum mucrone brevi
quadrangulato obtusi vel obtusissimi et inferne parum angustati, paulo com-
pressi et valde carinati, siccitate omnino dense et elevato tuberculato-rugosi.
B. mauritanica, var. nov.
Exsice. J. Ball, Tetuan, 1851, in Hb. Kew, ut Fumaria corymbosa!
Hooker, Tetuan, 1871, in Hb. Kew, ut F. africana !
LINN. JOURN.— BOTANY, VOL. XLIV. 2€
346 MR. H. W, PUGSLEY : A REVISION OF THE
"Margines petalorum exteriorum. quam in typo paulo latiores. Fructis
sine mucrone brevissimo 3—3:25 mm. longi, 2-2°5 mm. lati, oblongo-obovati, eis
iypi multo angustiores. Aliter ut typus.
Hee rara planta certe R. africane valde affinis est, sed foliis brevioribus
deltoideis lobis latioribus preeditis, bracteis sepalisque latioribus, corolla
minus gracili, petalorum exteriorum marginibus apice latioribus, calcare
breviore, fructibus majoribus valdius tuberculatis separanda est atque
specifice differre videtur.
R. decipiens in fissuris rupium, ad 400 m. adscendens, regionis Sierra de
Alora et Pizarra in provineià. Malacitand atque prope Grazalemam in pro-
vinciâ Gaditanà Hispaniw invenitur; varietas mauritanica prope Tetuan in
Mauritaniá et forsan ad Aloram cum typo crescit.
The specimens of this plant collected originally by Boissier at Alhaurin
(= Alora ?), now in the herbaria at Kew and the British Museum, are shade-
forms with relatively broad leaf-segments and poorly developed flowers.
Their fruits are found—rather unexpectedly—to differ from those of the
exsiccata collected at Alora by Huter, Porta & Rigo (No. 516), and rather to
resemble those of the Tetuan form.
[15. RUPICAPNOS PLATYCENTRA, Pomel, Nouv. Mat. Fl. Atlant. i. 242
(1874) ; n. v. Fumaria platycentra, Batt. & Trabut, Fl. d" Alger. i. 25
(1888).
Foliorum segmenta in lobos parvos obovatos vel lanceolatos profunde fissa.
Bractew oblonge, denticulate ; pedicelli apice inerassati. Sepala orbicularia,
valde dentata, corollae tubum :equantia. Corolla 14 mm. longa, albida; petali
superioris margines apice paulo dilatati, calcar circa 4 mm. longum vix
eurvatum apice abrupte rotundatum, nectarium flexuosum, ineurvatum ;
petala interiora. apice purpurea. — Fructás cum mucrone. quadrangulato
ovali-elliptici, siccitate tuberculato-rugost.
Heee species rupes ad Kef Iroad prope Toucriam in Algeria habitat.
lhis plant is maintained as a separate species in deference to the views of
Pomel, although no material of it has been available for examination. It
appears to be closely allied to R. decipiens, differing chiefly in its more finely
divided foliage and its strongly toothed, orbicular sepals. ]
16. RUPICAPNOS CEREFOLIA, Pomel, Nouv. Mat. Fl. Atlant. i. 242 (1874).
Fumaria cerefolia, Batt. & Trabut, Fl. d'Alger. 1.25 (1888) ; F. africana,
Coss. Comp. Fl. Atlant. ii. 80 (1883-1887), ex parte, non Lamarck.
Zesiee. Cosson, Djebel Zaccar, prés Milianah, 1854, in Hb. Kew, ut
Fumaria corymbosa! Joad, Milianah, 1873, in Hb. Kew, ut F. africana !
Cosson, Milianah, 1875, in Hb. Kew, ut F. africana! Soe. Dauphinoise,
No. 672, Milianah, ut F. africana!
Rupicapnos perennis, plus minusve glauca, caudice crassiusculo caulibusque
AMENS * Ebo oxc. 200102 M
GENERA FUMARIA AND RUPICAPNOS. 347
brevibus ut in X. africand. Folia pleraque subradicalia, glaucescentia,
10-20 em. longa (petiolo gracili incluso), longe petiolata, oblonga, foliolis
3-paribus subdeltoideis brevissime petiolatis et segmentis secundariis in
lacinias satis remotas, parvas, lineari-oblongas, acutas mueronatasve profunde
incisis. 2-3-pinnatisecta. Racemi racemoso-corymbiformes, 20—25 flori, cum
pedunculo (ad 7 em. longo) foliis sepissime paulo rarius subduplo breviores.
Bractew 15-4 mm. longe, oblonga (lineares ap. Pomel), acuminatze ; pedi-
celli fructiferi filiformes apice vix incrassati longissimi (infimi ad 60 mm.
longi). Sepala 3 mm. longa, l:5-1: 75 mm. lata, ovata vel lanceolato-ovata,
peltata, acuta vel acuminata, plus minusve irregulariter dentata vel basi
laciniata, preter nervum dorsalem angustum viridiusculum albida. Corolla
maxima, sepe gracilis, 14-16:5 mm. longa (12 mm. ap. Pomel), carinis
gracilibus viridiusculis albida vel roseo-albida : petalo superiore marginibus
apice paulo dilatatis sursum recurvatis vel patentibus anguste oblongo obtusis-
simo, calcare longo (circa 5 mm.) deflexo ad basin angusto sed apicem versus
rotundato ; petalo inferiore marginibus latis patentibus apice satis dilatatis
subspathulato obtuso, nonnunquam ad basin ipsam obscure saccato ; petalis
interioribus sursum curvatis apice modice alatis subtruncatis breviter sed læte
atropurpureis. LE ructiis modici, sine mucrone circa 2:5 mm. longi et 2 mm.
lati, obovato-elliptici, cum mucrone longo acuminati, valde compressi et plane
carinati, siccitate omnino grosse tuberculato-rugosi.
Hæc pulchra et distinctissima species per foliorum magis decompositorum
lacinias parvas angustas, per corollas maximas roseo-albidas spice breviter
sed conspicue atropurpureo-maculatas et per fructüs modicos subellipticos
acuminatos inter hujus sectionis alias species notabilis et facile distinguenda
est.
R. cerefolia prope Milianah atque Mazis (ap. Pomel) in provincia Algeriensi
Algeric invenitur.
The above description, which has been drawn up from the exsiecata cited,
agrees with Pomel’s diagnosis except that in the material examined the
bracts and sepals seem rather broader, and the corolla is larger, the flowers
being especially fine in Soc. Dauph. No. 672, and the specimen at Kew
collected by Joad. Pomel’s diagnosis seems to have been taken from poorly
grown plants, and is corrected by Battandier & Trabut (7. c.), who observe that
the flowers of F. cerefolia are very large, not less than 15 mm. in length.
The flower and dissections of Fumaria africana in Cosson's Illustr. FI.
Atlant. i. tab. 9, figs. 12-20, appear to have been drawn from a plant of
this species.
17. RuPiCcAPNOs speciosa, Pomel, Nouv. Mat. Fl. Atlant. i. 241 (1874).
Fumaria speciosa, Batt. & Trabut, Fl. d Alger. i. 25 (1888) ; F. africana,
Coss. Comp. Fl. Atlant. ii. 80 (1883-1887), ex parte, non Lamarck.
Crsicc. Warion, Saida, 1872, in Hb. Kew (sine fructibus), ut F. africana!
Te - i, ER
348 MR. H. W. PUGSLEY : A REVISION OF THE
Rupieapnos perennis habitu R. africane. Folia fere ut in R. african
(maxima ap. Batt. & Trab.) sed oblonga, segmentis secundariis subflabellatis
in lacinias latiuseule oblongas, subacutas vel mucronatas fissis 2-pinnatisecta.
Racemi F. africane. Bractew 2-4 mm. longe, lanceolate, acuminate ;
pedicelli fructiferi filiformes apice incrassati. Sepala cirea 3 mm. longa ac
lata, fere orbicularia (corollae tubum subeequantia), peltata, acuta vel breviter
acuminata, valde dentata, albida. Corolla magna, 14-15 mm. longa, carinis
viridibus albida ; petalo superiore marginibus apice vix dilatatis dilute pur-
purascentibus oblongo obtuso, calcare brevi (circa 3 mm. longo) apice rotundato,
[nectario lineari incurvato ap. Pomel] ; petalo inferiore marginibus patentibus
apice paulo dilatatis l/neari-oblongo obtuso, basi gibboso-saccato ; petalis interi-
oribus apice violaceo-atropurpureis. — Fructis verisimiliter modici, eum mucrone
longo lanceolato-obovat?, siccitate omnino valde tuberculato-rugosi.
Hee species habitu foliisque X. africanam refert, sed per foliorum seg-
menta latiora, per sepala orbieularia nee ovalia, per corollam haud gracilem
calcare brevi preeditam satis differt. 2. decipiens foliis deltoideis, sepalis
ovalibus, corolla longius calearata, fructibus multo majoribus facile separanda
est, atque KX. cerefolia per folia magis decomposita laciniis parvis angustis
prodita, per sepala angusta, per corollam longe calearatam apice breviter
purpureo-maeulatam et per fructüs subellipticos acuminatos omnino diversa
est.
R. speciosa rupes ad fluminis Mine cataractam prope Mascaram (ap.
Pomel), ad Saidam et forsan ad locos alios in provincia Oranensi Algerie
habitat.
No authentie or satisfactory material of this plant has been seen, but
Warion's Saida example at Kew, although in a fragmentary condition,
appears clearly referable to it and has been cited accordingly. The specific
diagnosis given above is based on the work of Pomel, collated with this
exsiccata. Another example at Kew, one of two labelled * F.corgymbosa—
Munby, Tlemcen,” may also belong to this species, which it closely resembles
in foliage, while its fruits, which are not large, agree in form with Pomel’s
description. Unfortunately this specimen now shows no trace of flowers.
Sussectio POMELIANZE.
Foliorum lobi lati. Flores pallide purpurei.
18. RurrcAPNos PoMELIANA, sp. nov. (PI. 9, fig. 4.)
R. africana, Pomel, Nouv. Mat. Fl. Atlant. i. 240 (1874), nomen
abortivum ; Fumaria africana, Coss. Comp. Fl. Atlant. ii. 80 (1883—
1887), ex parte; Batt. & Trabut, Fl. d'Alger. i. 25 (1888) ; non
F. africana, Lamarck.
Exsice. Bourgeau, Pl. d'Alger. 1856, Garrouban, in Hb. Mus. Paris (pro
maj. parte), ut Fumaria corymbosa !
GENERA FUMARIA AND RUPICAPNOS. 349
Rupicapnos perennis, glauca, caudice crassiusculo, ramoso, brevi vel paulo
elongato, decumbente, tandem vestigiis petiolorum emarcidorum donato,
caulibusque semper brevibus. Folia pleraque subradicalia, glauca, carnosa,
10-20 em. longa (petiolo incluso), longe petiolata, oblonga, foliolis late
obovatis breviter petiolatis et segmentis secundariis subimbricatis raro
alternantibus in lobos rotundatos vel obovatos obtusos vix mucronulatos sæpe
incisos irregulariter fissis. 2-pinnatisecta subquinata, Racemi corymbiformes,
multi-(circa 20-) flori, cum pedunculo (ad 6 em. longo) foliis subduplo
breviores. Bractew 2-275 mm. longe, late ovales, cuspidatz, apicem versus
dentate ; pedicelli fructiferi graciles (fere filiformes), apice incrassati,
flexuosi, infimi ad 25 mm. longi. Sepala 3-35 mm. longa, fere 3 mm. lata,
suborbicularia (coroll tubo latiora), peltata vel cordata, breviter acuminata,
presertim basin versus inciso-dentata, prseter nervum dorsalem latum rubi-
cundum purpurascentia. Corolla magna, 12-14 mm. (15 mm. ap. Pomel)
longa, plus minusve gracilis, pallide purpurea ; petalo superiore marginibus
apice parum dilatatis subpatentibus vel deflexis (nonnunquam paulo sursum
recurvatis) anguste oblongo obtuso, cariná viridiusculà petali apicem vix
attingente, calcare brevissimo (2-3 mm. longo) rotundato curvato; petalo
inferiore marginibus patentibus apicem versus vix dilatatis lineari-oblongo nec
subspathulato, obtusiusculo, ad basin ipsam gibboso-saccato ; petalis interioribus
sursum curvatis apice modice alatis atropurpureis. Fructús majusculi, sine
mucrone 2:5-2:75 mm. longi ac lati, verisimiliter subrotundo-obovati (fere
ovales ap. Pomel), cum mucrone brevi obtusissimi et inferne angustati, satis
compressi et valde carinati, siccitate omnino grosse et elevato tuberculato-
rugosi.
Hæc planta per foliorum lobos latissimos rotundatos, per bracteas latis-
simas atque per flores purpureos brevissime calcaratos sepalis suborbicularibus
preeditos ab hujus gregis speciebus aliis specifice differre videtur.
R. Pomeliana rupes ad Garrouban et forsan ad locos alios in provincia
Oranensi Algerie habitat.
This plant is identified by Pomel with the original Fumaria africana,
Lamk. It has been shown, however, that the descriptions of Lamarck and
Desfontaines can be definitely determined to refer to another species, albeit
the name of F. africana was subsequently extended by Cosson to cover in an
aggregate sense practically all the forms of the present section. As the
specific epithet “ africana? must be retained for the form originally so
described, and the plant so named by Pomel is distinet and has not been
otherwise distinguished, a fresh epithet becomes necessary for the latter. It
is therefore proposed to commemorate Pomel in its new name.
It is noteworthy that while Pomel's diagnosis of this plant agrees well
with the Garrouban exsiccata in Herb. Mus. Paris, the examples seen from
other localities which he cites (Tlemcen, Nedroma, and Oran) show different
characters and seem to belong to other species. Of these, the Oran plant
350 MR. H. W. PUGSLEY : A REVISION OF THE
which is less sparingly represented in herbaria than the other forms of the
genus and agrecs with none of Pomel’s diagnoses, is accordingly separated
as a new species. It is of course possible that the forms from these localities
that came under the notice of Pomel were identical with the Garrouban
example, for it is almost certain from the mixtures observed in exsiccata
that different species of this genus grow in company. But if so, it is curious
that Pomel, who apparently knew Oran well, should not have met with the
plant that has been most generally obtained there by other collectors.
19. RUPICAPNOS ORANENSIS, sp. nov. (Pl. 9, fig. 5.)
Fumaria africana, Coss. & Dur. in Bull. Soc. Bot. Fr. ii. 305 (1855), ex
parte; Coss. Comp. Fl. Atlant. ii. 80 (1883-1887), ex parte, non
Lamarck; F. corymbosa, Hammar, Mon. Fumar. 42, excl. syn. ct loc.
Alhaurin (1857), non F. corymbosa, Desfontaines.
Icon. Hammar, l. e. tab. v, ut F. corymbosa (sed petalo inferiore haud
saccato).
Ewsice. Balansa, Pl. d'Alger. 1852, No. 337, ut F. corymbosa! | Munby,
Pl. Alger. 1850, No. 38, ut F. corymbosa! Bourgeau, Pl. d'Alger. 1856,
No. 181, Nedroma (partim, sed non in Hb. Mus. Paris), ut /. africana !
Faure, Pl. d'Alger. 1904, Oran, ut F. africana!
Rupicapnos perennis sed a primo anno jam florens, glauca, radice elongata,
caudice crassiusculo ramoso decumbente brevi vel paulo elongato, caulibus
semper brevibus nonnunquam fere nullis. Folia pleraque subradicalia,
glauca vel glaucescentia, plus minusve carnosa, 10-30 em. longa (petiolo
incluso), longe petiolata, oblonga, foliolis fere ovatis infimis breviter petiolatis
reliquis subsessilibus et segmentis seeundariis subflabellatis sepius alternan-
tibus in lobos obovatos vel oblongos obtusos mucronatos vel subacutos sæpe
incisos irregulariter fissis 2-pinnatisecta. Racemi corymbilormes, multi-
(10—30-, ssepius plus 20-) fori, cum pedunculo (ad 5 em. longo) foliis seepissime
plus duplo breviores. — Dractee 15-3 mm. longæ, oblonga, acute rarius
acuminate, interdum sparsim serrulatæ, verisimiliter purpurascentes ; pedi-
celli fructiferi graciles (Fere filiformes), apice incrassati, flexuosi, infimi usque
ad 40 mm. longi. Sepala minima, 2-2:5 mm. longa, l:5—1*15 mm. lata, ovata,
peltata vel cordata, acuta, basin versus irregulariter crenato-dentata, preter
nervum dorsalem latum rubicundum purpurascentia, facile caduca. Corolla
maxima, 12-16 mun. longa, ut videtur carinis viridibus pallide purpurea ;
petalo superiore marginibus apicem versus paulo dilatatis sepius sursum
recurvatis purpurascentibus oblongo obtusissimo, calcare longiusculo (4-5 mm.
longo) curvato apice rotundato (in formis umbrosis breviore) ; petalo inferiore
marginibus latiusculis patentibus apicem versus sensim paulo dilatatis lineari-
oblongo vix subspathulato obtusiusculo, ad basin ipsam sepissime plane gzbboso-
succato; petalis interioribus sursum curvatis apice modice alatis atropurpureis.
GENERA FUMARIA AND RUPICAPNOS. 351
Stylus malleiformis. Fructis magni, sine mucrone 3-3:25 mm. longi, 2'5-
2:15 mm. lati, ovali-obovati, eum mucrone conspicuo obtusissimi et inferne multo
angustati (supra medium latissimi), plane compressi et valde carinati, siccitate
omnino grosse tuberculato-rugosi.
Hee species foliorum lobos latos et flores pallide purpureos R. Pomeliane
habet sed braeteis sepalisque multo angustioribus, petali superioris ealcare
subduplo longiore et fructibus plane longioribus satis differre videtur.
R. oranensis vicinitatem oppidi Oran, Nedromam, Tlemcen ut videtur, et
probabiliter alios locos in parte septentrionali provincia Oranensis Algerire
habitat.
This plant, which is better represented in the herbaria consulted than any
other member of the genus, is rather inexplicably unnoticed by Pomel, unless
it is intended to he included with the previous species X. Pomeliana, as may
perhaps be inferred from the localities cited. It is not easy to adopt this
view, however, seeing how clearly Pomel’s description accords with the
features of the Garrouban plant in Herb. Mus. Paris and how widely it
differs in severai particulars from œR. oranensis. On the other hand, as
already remarked under R. Pomeliana, it seems impossible that Pomel was
unacquainted with this plant in the neighbourhood of Oran.
In shaded situations the foliage of X. oranensis becomes lax with relatively
thin leaflets, somewhat acutely and by comparison narrowly lobed, while its
flowers tend to be reduced in size, with no upward recurving of the margins
of the upper petal, and a shorter, straighter spur. These features may be
seen in some of the specimens sent out by Balansa, by Bourgeau, and by
Faure. Munby's plants, on the other hand, are mostly very dwarf examples,
probably collected in dry and exposed rock-fissures, and their clusters of
well-developed flowers almost equal the much reduced leaves, of which the
segments are thick in texture and very small.
Judging from exsiecata, the flowers of R. oranensis appear to be of a paler
purple colour than those of Æ. Pomeliana, and they are less concolorous, the
margins of the upper petal being sometimes distinctly flushed with a deeper
tint, recalling the coloration of the flowers of Fumaria agraria.
There is a specimen of this plant at Kew from Munby’s herbarium, associ-
ated with another which may be X. speciosa and labelled: Tlemcen,” but it
appears uncertain from the arrangement of other examples on the sheet
whether it really came from this habitat or whether a transposition of
specimens has not at some time taken place.
[20. RUPICAPNOS OCHRACEA, Pomel, Nouv. Mat. Fl. Atlant. i. 242 (1874) ;
n. v. Fumaria ochracea, Batt. & Trabut, FI. d'Alger. i. 25 (1888) ;
F. africana, Coss. Comp. Fl. Atlant. ii. 80 (1883-1887), ex parte,
non Lamarck.
Foliorum segmenta in lobos parvos obovatos vel oblongos spathulatos fissa.
:
:
y
P
-—— Dey ETE
352 MR. H. W. PUGSLEY: A REVISION OF THE
Bractew lanceolata, integre ; pedicelli graciles apice breviter incrassati.
Sepala minima, lanceolata, valde denticulata, quam corolle tubus multo
angustiora, Corolla circa 12 mm. longa, gracilis, pallide ochraceo-purpurea ;
petali superioris margines apice parum dilatati ; calcar longiusculum (circa
4 mm. longum), apice rotundatum ; petala interiora apice purpurea. Fructis
angusti, ewm mucrone gracili fere oblongi, inferne in stipitem distinctum
angustati, siccitate rugosi. .
R. ochracea rupes calciferas ad Goudjilam prope Tiaret in parte orientali
provinciæ Oranensis Algeriæ habitat.
This plant, of which no material has been seen, seems essentially distinct
from the other species of this section owing to its peculiarly coloured and
much smaller flowers, and its narrow fruits. It may possibly be regarded as
intermediate in characters between X. oranensis and KR. erosa. |
EXPLANATION OF THE PLATES.
PLATE 9.
Fig. 1. Flower of Fumaria agraria, Lag., with transverse section near the apex.
Fig. 2. Flower of F. capreolata, Linn., do. do.
Fig. 3. Flower of F. Gussone?, Boiss., do. do.
Fig. 4. Flower of Rupicapnos Pomeliana, sp. nov, — do. do.
Fig. 5. Flower of R. oranensis, sp. nov., do. do.
Fig. 6. Flower of R. numidica, Pomel, do. do.
Fig. 7. Flower of 2. caput-platalee, Pomel, do. do.
Fig. 8. A fully developed flower of Fumaria sepium, Boiss.; drawn from a specimen col-
lected at the locus classicus.
Fig. 9. A depauperate flower of the same.
Fig. 10. A completely cleistogamic flower of F. muralis subsp. orci; from a cultivated
specimen.
Detached flowers all enlarged x 2:5. A=keel; B=wing or margin.
PLATER 10.
Fumaria Ball, sp. nov., natural size, with detached flower and dried fruit ( x 2:5)
drawn from the unique specimen at Kew.
E
Prat 11,
Fig. l. Fumaria berberica, sp. nov., natural size, with detached flower and dried fruit
(x 25); drawn from a specimen at Kew.
Fig. 2. F. montana, J. A. Schmidt, natural size, with detached flower and dried fruit
(x 2:5); drawn from Schmidt’s original specimen at Vienna,
PLATE 12.
Fumaria coccinea, sp. nov., natural size, with detached flower and dried fruit
(x25); drawn from a specimen collected by Messrs, Sprague and Hutchinson
(no, 1265),
PUGSLEY. JOURN. LINN. Soc., Bor. VoL. XLIV. PL. 9.
H. W. P. del. Grout, se.
1-3& 8.10. FLOWERS OF FUMARIA. 4-T, FLOWERS OF RUPICAPNOS.
PvGsLkr. JOURN. LINN. SOC.» Bor. VoL. XLIV. PL. 10
H. W. P. del. Grout, ec.
FUMARIA BALLII, sp. nov.
PUGSLEY. JOURN. LINN. S0C: Bor. VOL. XLIV. PL. 11.
H. W. P. del. Grout, sc,
1, FUMARIA BERBERICA, sp. nov. 2, F. MONTANA, J. A. Schmidt.
H. W. P. del.
JOURN. LINN. SOC., Bor. VOL. XLIV. PL. 12.
FUMARIA COCCINEA, sp. nov
PUGSLEY JOURN. LINN. SOc., Bor. VOL. XLIV. PL. 13.
Jr. W. P. del. Grout, sc
FUMARIA AUSTRALIS, sp. nov.
PUGSLEY. JOURN. LINN. SOC., Bor. VoL. XLIV., PL. 14.
H. W. P, del. Grout, sc.
1, RUPICAPNOS MURICARIA, Pomel. 2. R. PRAZTERMISSA, sp. nov
PUGSLEY. JOURN., LINN. Soc., Bor. VOL. XLIV. PL. 15.
H. W. P. de). Grout, sc.
1. RUPICAPNOS LONGIPES, Pomel. 2, R. SUBLZEVIS, sp. nov.
PUGSLEY. JOURN. LINN. Soc., Bor. VoL. XLIV. PL. 16.
H, W. P. del.
RUPICAPNOS SARCOCAPNOIDES, Pomel.
GENERA FUMARIA AND RUPICAPNOS. 3853
PLATE 13.
Fumaria australis, sp. nov., natural size, with detached flower and dried fruit
(X 25); drawn from a specimen at Kew (Volkens, no. 953).
PLATE 14.
Fig. 1. Rupicapnos muricaria, Pomel, natural size, with detached flowers and dried fruit
(x 25); drawn from a specimen in Herb. Mus, Manchester (Chevallier, no. 397).
Fig. 2. R. pretermissa, sp. nov., natural size, with detached flowers and dried fruit (x 2°5) ;
drawn from a specimen in Herb. Mus. Paris.
Prater 15.
Fig. 1. Rupicapnos longipes, Pomel, natural size, with detached flowers and dried fruit
(x 2:5); drawn from a type-specimen in Herb. Mus, Paris.
Fig. 2. R. sublevis, sp. nov., leaf and fruit (x 2:5); drawn from a specimen in Herb. Mus.
Paris. :
PLATE 16.
Rupicapnos sarcocapnoides, Pomel, natural size, with detached flowers and dried
fruit (x 2:5); drawn from a type-specimen in Herb, Mus. Paris.
LINN. JOURN —BOTANY, VOL. XLIV. ` 2F i
MR. W. H. PUGSLEY : A REVISION OF THE
INDEX
SPECIES AND VARIETIES RECOGNIZED AS VALID IN THE
FOREGOING REVISION ON PAGES 259-327, 331-352.
A star * denotes a name here published for the first time.
Fumaria, Linn., 250-327.
abyssinica, Hamm., 308.
agraria, Lag., 259.
— var. chilensis, Parl., 259.
— — mauritanica, Haussk., 259.
amarysia, Boiss. & Heldr., 267.
apiculata, Lange, 289.
asepala, Boiss., 821.
— var. aprica, Pugs.*, 321.
— — compacta, Haussk., 321.
atlantica, Coss. & Dur., 260.
— var. platyptera, Pugs.*, 261.
australis, Pugs.*, 309.
Ballii, Pugs.*, 262.
Bastardii, Bor., 278.
— var. affinis, Pugs.*, 279.
— — benedicta, Pugs.*, 280.
— — Gussonei, Pugs., 279.
— — hibernica, Pugs., 279.
— — patens, Pugs.*, 278,
— — vagans, Pugs.*, 279.
berberica, Pugs.*, 273.
bicolor, Somm., 276.
Boissieri, Haussk., 306.
bracteosa, Pomel, 300.
capreolata, Linn., 269.
— var. albiflora, Hamm., 270.
— — Babingtonii, Pugs., 270.
— — condensata, Ball, 270.
— — devoniensis, Pugs., 970.
— — Hochreutineri, Pugs.*, 270.
— — speciosa, Hamm., 269.
cilicica, ZZawssk., 305.
coccinea, Lowe *, 274.
dubia, Pugs.*, 972.
flabellata, G'asp., 266.
Gaillardotii, Boiss., 265,
indica, Pugs.*, 313.
Fumaria
. indica f. montana, Pugs.*, 314.
Janke, Haussk., 510.
judaica, Boiss., 267.
— var. insignis, Puys.*, 267.
Kralikii, Jord., 298.
macrocarpa, Parl., 268.
macrosepala, Boiss., 273.
major, Bad., 263.
— var. algerica, Pugs.*, 264.
— — spectabilis, Rouy, 263.
Martinii, Clav., 982.
micrantha, Lag., 299.
— var. dubia, Pugs.*, 299.
— — littoralis, Rouy § Fouc., 300.
microcarpa, Boiss., 312.
microstachys, Kralik, 306.
montana, J. A. Schmidt, 297.
Munbyi, Boiss. & Reut., 281.
muralis, Sond., 285.
— subsp. Boræi, Pugs., 287.
— — — var. ambigua, Pugs., 288.
— — — — britannica, Pugs., 288.
— — — — gracilis, Pugs., 288.
— — — — hispanica, Pugs.*, 288.
— — — — longibracteata, Pugs., 288,
— — neglecta, Pugs., 287.
— var. decipiens, Pugs., 286.
— — læta, Lowe, 286.
— — Lebelii, Rouy § Fouc., 986.
— — Lowei, Pugs.. 286.
occidentalis, Pugs., 261.
officinalis, Linn., 302.
— var. densiflora, Parl., 304.
— — elegans, Pugs., 304.
— -— minor, Koch, 303.
— — Wirtgenii, Zaussh., 303.
Painteri x, Pugs., 989.
GENERA FUMARIA AND RUPICAPNOS. 255
Fumaria
parviflora, Zam., 322.
— f. Vivianii, Pugs.*, 323.
— var. acuminata, Clav., 325.
— — glauca, Clav., 524.
— — indicoides, Pugs.*, 325.
+ — latisecta, Haussk., 523.
— — persica, Pugs.*, 324.
— — sinaitica, Haussk., 325.
— — Symei. Pugs., 323.
Petteri, Reichd., 290.
purpurea, Pugs., 272.
— var. brevisepala, Puys., 272.
Reuteri, Boiss., 205.
rostellata, Knaf, 301.
rupestris, Boiss. § Reut., 261.
Schleicheri, Soy.- Will., 310.
— var. supina, Pugs.*. 310.
Sehrammii, Pugs.*, 319.
— var, orientalis, Pugs.*, 319.
sepium, Zoiss., 284.
— var. gaditana, Pugs.*, 284.
Thuretii, Boriss., 292.
— var. deflexa, Pugs.*, 203.
— — Heldreichii, Boriss., 295.
— — pikermiana, Pugs.*, 294.
— — thasia, Pugs.*, 295.
Vaillantii, Zois., 315.
Fumaria
316.
— — conferta, Haussk., 316.
— — venetica, Pugs.*, 317.
Rupicapnos, Pomel, 328-352.
africana, Pugs.*, 943.
caput-plataleee, Pomel, 340.
cerefolia, Pomel, 346.
Cossonii, Pugs.*, 338.
decipiens, Pugs.*, 345.
— var. mauritanica, Pugs.*, 345.
delicatula, Pomel, 332.
erosa, Pomel, 941.
— var. major, Pugs.*, 342.
longipes, Pomel, 332.
muricaria, Pomei, 331.
numidica, Pomel, 337.
ochracea, Pomel, 351.
oranensis, Pugs.*, 350.
platycentra, Pomel, 946.
Pomeliana, Pugs.*, 548.
prietermissa, Pugs.*, 333.
Reboudiana, Pomel, 336.
sarcocapnoides, Pomel, 335.
speciosa, Pomel, 547.
sublievis, Pugs.*, 334.
tenuifolia, Pomel, 339.
Vaillantii, var. Chavinii, Rouy § Fouc.,
ii ER mi a
Rea ee mee
aes
PETITES RR ee ee ees
aE re ees
MR. C, E, SALMON ON A HYBRID STACHYS, BDF
A Hybrid Stachys. By C. E. Sarmon, F.L.S.
(With one Text-figure.)
[Read 17th January, 1918.]
For some years I have grown in my garden Stachys germanica raised from
Oxfordshire seed kindly given me by Mr. S. H. Bickham (see Watson Bot.
Ex. Club Rep. 1912-13, p. 405), and close by several flourishing clumps of
Stachys alpina which came to me from Mr. J. W. White's garden at Clifton,
and whose origin was the only British station in West Gloucestershire
(Journ. Bot. 1897, p. 380 ; 1898, p. 209).
Early last summer I noticed, in close proximity to the above, a
partieularly vigorous plant sending up several stems and bearing root-leaves
that in colour and hair-clothing could be neither those of germanica nor
alpina, and later, when the plant was in full bloom, which occurred just
after the period when ©. alpina was at its best and before that of
S. germanica, it became obvious that I had an interesting hybrid between
these two species.
The colouring of the plant, as a whole, was almost intermediate between
the silvery-grey tone of germanica (caused by the abundance of its white
silky tomentum) and the comparatively green tint of alpina.
On its strong stout stems, between three and four feet high, overtopping
both its parents (as they grow in my garden), an abundance of bright flowers
were borne, larger than those of either alpina or germanica and of a different
colour. Those of alpina are of a dull purple, inclining to brown, and
germanica bas corollas of a decidedly pleasing pink tint ; the flowers of the
hybrid were clear bright purple, slightly paler near the throat.
The next step was to ascertain what was known and had been written
about this hybrid, and I found it had been described as long ago as 1789
when William Aiton (then of the Royal Gardens, Kew) gave it specific rank
as Stachys intermedia, with the following diagnosis :—* S. verticillis multi-
floris, calycibus subpungentibus, foliis oblongis subcordatis crenatis, caule
sublanato. Oblong-ieav’d Stachys. Nat. of Carolina. Introd. about 1762.
Fl. June & July” *.
This deseription is very good as far as it goes, but there must be some
error as regards the country of origin ; neither S. germanica nor S. alpina
(and consequently the hybrid) are known to occur in America. An
authentic specimen, probably sent by Aiton himself, may be seen in the
British Museum Herbarium labelled: “ S. intermedia. Hort. Kew. Julio
1764," and this matches my examples extremely well.
: * Hort. Kewensis, ii. 301.
LINN, JOURN,—BOTANY, VOL. XLIV.
bo
Q
358 MR. C. E. SALMON ON
Since the above was written, Mr. Britten kindly examined this specimen
and reported that after the words “S. intermedia” on the sheet, some
writing I could not decipher was certainly * MSS.", and that this refers to
the Solander MSS. in which the description is found and upon which
* Hortus Kewensis," ed. 1, was based. Mr. Britten added this interesting
note :—* The name on the sheet and the description in the MSS. (vol. xiii.
p. 67) are entirely in Solander’s hand, so that if you like, you can cite the
plant as of [Solander in] Aiton, Hort. Kew. ii. 301.
“ Solander, in his MSS., added, later than the description, * 1764 in Carolina.
Catesby. Consequently I hunted up Catesby’s plants and in Herb. Sloane
212, f. 29, I found his specimen, also named by Solander .... Whatever
this may prove to be, the Kew Gardens plant is the type of S. intermedia as
the reference to Catesby was added later.”
I have looked at this American plant and believe that it is a broad-leaved
form of Stachys palustris, a frequent species in the Southern States, whereas
S. germanica and alpina are unknown there.
I presume that the plant that occurred in my garden used to be in cultiva-
tion at Kew, as it is included in the Kew Handlist of Herbaceous Plants,
ed. 1, p. 445, 1895 (as S. alpina, L., var. intermedia), but it has apparently
died out and is not mentioned in the 1902 (2nd) edition. I cannot find the
plant included in any of our nurserymen's catalogues that I have come
across, nor have I seen it in any gardens with which I am acquainted.
Nothing is known of it at the Cambridge Botanie Garden.
On the Continent, however, where the two parents are more widely and
abundantly distributed under natural conditions, the hybrid is apparently
of quite frequent occurrence in gardens. M. Rouy states that it is often
cultivated in gardens (Fl. Fr. xi. p. 308, 1909), and M. Correvon, the well-
known hortieulturist of Chéne-Bourg, Geneva, writes :—* They all cross
very often even in nature. We have in Switzerland alpino-lanata and
alpino-germanica."
Briquet has an interesting note, under S. alpina in his * Labices des Alpes
Marit.” par. 2. 240, 1893, where he mentions that it crosses particularly
readily with S. germanica, giving rise to a series of intermediate plants,
usually sterile, of which many are cultivated in gardens on account of their
beautiful spikes of flowers and velvety foliage.
He mentions the various names that have been given to these hybrids, and
remarks that although no specimens have yet been found in his distriet in
a wild state, it is quite likely to occur, as the parents are both found.
He explains Bentham’s remark in DC. Prodr. xii. 465 : “J see no definite
limit between S. germanica, S. lanata, and . . . . S. alpina," owing to the
fact that that author recognized no hybrids in the genus Stachys.
Rouy (I. c.) has attempted to segregate the hybrid accordingly as it varies
towards one parent or the other, using the name /ntermedius, Ait., for the
aggregate plant, with synonyms S. sibirica, Link, and S. rosea, Hohen,
=
pa.
À HYBRID STACHYs. 359
The form which approaches the germanica parent he called “ a. digeneus
nob.", founded upon the S. digenea, Legué (in Bull. Soc. Bot. Fr. xl. p. 213,
1893), and to this he assigned the synonyms S. germanica var. intermedia,
Boiss., and S. curviflora, Tausch.
The form which has more affinity with S. alpina, Rouy called “8. para-
doaus nob.", which he considered synonymous with S. alpina var. intermedia,
Benth., and, doubtfully, S. urticifolia, Tausch.
If we follow Rouy, my plant must be placed under his a. digeneus, as
it possesses a preponderance of the characters of germanica and agrees
satisfactorily with M. Legué's original description (l. c.).
As regards the synonyms mentioned by the authors quoted above, the
following notes were made when endeavouring to work out my specimens at
the British Museum.
S. sibirica, Link.—This is described in Enum. Hort. Bot. Berol. pars 2,
p. 109 (1822), and from the description it is possible it may refer to the
hybrid under discussion, but if the figure in Sweet, Brit. Fl. Garden, i. t. 100
(1825) correctly portrays Link's plant (and it is so labelled), then that is a
form with remote whorls of flowers even at the summit of the stem, and the
clothing of the plant less silky in nature than in my specimens.
Reichenbach’s Ic. Crit. iv. t. 328 also figures the S. sibirica of Link, but
that again differs from my plant in many respects, which tallies far better
with t. 327 in the same work, labeled S. intermedia, Ait.
S. rosea, Hohen.— Boissier (Fl. Orientalis, iv. 720, 1879) gives this as one
of the synonyms of his S. germanica var. intermedia, and indicates that he
has seen a type-specimen. It must not be confused with the S. rosea of the
latter author (op. cit. 725). Hohenacker's description (Enum. Talysch, 300,
1838) is meagre in the extreme, practically a “nomen nudum," and I have
not been able to see any specimens.
S. alpina, L., var. intermedia, Bentham in DC. Prodr. xii. p. 465 (1848).
This has a short description—*' major, foliis rugosioribus subtus interdum
sublanatis," and a large number of synonyms. We cannot, I think,
conclude as Rouy has done, that because the * variety " is placed under
alpina it indicates a form of the hybrid that approaches more to that parent.
Indeed, Bentham adds—** An ad S. germanicam melius referenda ??
Of the synonyms he gives, two strike me as being remarkable—
“ S. cretica, Linn. spec. p. 812, fide deseriptionis. S. orientalis, Linn. spec.
p. 813, fide exempl. Tournef. in h. Banks.”
As regards the former, Index Kewensis (ii. 972, 1895) indicates that
S. cretica, L., Sp. pl. 581 (the Bentham synonym refers to ed. 2 of Sp. pl.)
equals * germanica, alpina”: the Linnean description, however, will not do
at all for the hybrid under discussion, but seems to correspond with the fine
figure in Sibthorp, Fl. Grzeca, vi. t. 558 (1827) labelled S. cretica.
360 MR. 0. E, SALMON ON
Concerning S. orientalis, here again Ind. Kew. (op. cit. 973) refers the
Linnean plant to “alpina” ; neither the description nor the plate in Sibthorp
(op. eit. t. 560) of S. orientalis bears this out. Moreover, the Tournefortian
example in Herb. Mus. Brit., quoted by Bentham, is clearly not of close
relationship with germanica, alpina, or intermedia.
S. germanica, L., var. intermedia, Boiss.—I think it is clear from the
description given (loc. cit.) that Boissier would include under his name that
form of the hybrid which shows more obviously the germanica parent.
It might be possible to arrange the rather puzzling synonymy of the
hybrid and the varying forms as follows :—
The * aggregate " hybrid :
S. sibirica, Link (1822).
S. alpina, L., var. intermedia, Benth. (1848). :
S. alpina x germanica, Briq. (1893).
* Regregate " A h
(nearer germanica). J S. intermedia, Sol. in Ait. (1789) *,
S. curviflora, Tausch (1831) (fide Rouy).
S. rosea, Hohen. (1838). non Boiss.
S. germanica, L., var. intermedia, Boiss. (1879).
S. digenea, Legué (1893).
S. intermedius, Ait., var. digeneus, Rouy (1909).
* Begregate ” B \
(nearer alpina). S. paradoxa, Rouy (1893).
? S. urticifolia, Tausch (1831).
S. intermedius, Ait., var. paradoxus, Rouy (1909).
It may perhaps be of interest to record in tabular form the characteristics
of my hybrid as eompared with the corresponding features of S. germanica
and S. alpina, observed from growing and dried material :—
S. germanica. 5S. digenea. S. alpina.
Plant c. 75 cm. in height or Plant c. 107 em. in height or Plant c. 60 em. in height or
more; stem covered with more; stem provided with more; stem hairy with +
an abundance of dense copious white silky + — spreading hairs, the shorter
white silky + spreading spreading pubescence mixed ones (particularly near sum-
pubescence. No glandular with shorter glandular mit of stem) glandular.
hairs. Whole plant not hairs. Whole plant scented Whole plant with strong
strongly scented and silvery- as in alpina and grey-green S. sylvatica-like scent and
grey in appearance, in appearance. green in appearance.
we
* It is not the S. intermedia of Tenore (Fl. Nap. i. p. xxxiv, 1811-15), which is
S. heraclea, as Tenore himself afterwards admitted,
:
S. germanica.
Leaves (basal) ovate-oblong,
acute, provided with dense
white silky non-glandular
pubescence (and thus grey-
ish in appearance) more
abundant on lower surface ;
petioles similarly clothed.
Upper leaves and bracts
ovute- or linear-lanceolate,
decidedly more densely
silky than basal.
All leaves thick in tex-
ture.
Flowers in whorls forming a
dense terminal spike with
a few separated whorls
below.
Calyx 10-11
obeonic, teeth
acuminate, usually rather
shorter in proportion to
calyx than those of alpina,
Whole calyx (and teeth)
clothed with long white
ascending silky hairs, with
sessile glands but no glan-
dular hairs.
min. long,
triangular-
Corolla pale purple or rose-
pink; upper lip arched,
c. $ as long as calyx, with
long white silky hairs pro-
jecting considerably beyond
tip; lower lip slightly
longer than upper, + entire
or crenulate.
Hairs in corolla - tube
about halfway down.
Stiyma white.
Ripe seeds smooth, almost
black (blackish brown),
c. 2 mm. long.
A HYBRID STACHYS.
S. digenea.
Leaves (basal) ovate-oblong,
+ obtuse, green and hairy
above, usually grey-green
below with hairs and +
silky pubescence and also
sessile glands; petioles
more silkily hairy than in
alpina,
bracts than in
alpina and broader than in
Upper leaves and
narrower
germanica, decidedly more
silkily hairy than basal.
Leaves slightly thick in
texture.
Flowers in whor!s laxer than
those of germanica and
closer near top of spike than
those of alpina.
Calyx 10-12 mm. long, ob-
conie, teeth triangular-
acuminate, as in germanica.
Whole calyx (and teeth)
clothed with long white
ascending silky hairs, not
quite so dense as in ger-
manica, with sessile glands ;
short glandular hairs pre-
sent on calyx-teeth, rarely
on tube.
Corolla bright purple, paler
near throat, larger than in
germanica or alpina ; upper
lip arched, c. $ as long as
calyx, with long white
hairs (not quite so dense
as in germanica) projecting
considerably beyond tip;
lower lip considerably
longer than upper, + entire
or crenulate, broader than
in germanica or alpina.
Hairs in corolla- tube
about ł way down.
Stigma almost white, faintly
tinged with purple.
Ripe rather more
coarsely rugulose than in
alpina, brown, c. 23 mm.
long.
Only one seed usually
fertile in each calyx.
seeds
LINN. JOURN.—BOTANY, VOL. XLIV.
361
S. alpina.
Leaves (basal) ovate, obtuse,
green, paler on underside,
hairy above, rather less so
beneath where there are
sessile glands; petioles
hairy. Upper leaves and
bracts ovate-acute, only
slightly more hairy than
basal.
Leaves thin in texture.
Fiowers in whorls almost all
separated, with a few aggre-
gated ones at summit.
Calyx 10-11 mm. long, cam-
panulate, teeth broadly tri-
angular (more abruptly
pointed than in germanica)
acuminate or mucronate,
Whole calyx (and teeth)
hairy with ascending-
patent hairs, mixed with
shorter glandular ones ;
sessile glands also present.
Corolla dull brownish-purple ;
upper lip flat, c. } as long
as calyx, with dense hairs,
shorter than those of ger-
manica, projecting beyond
tip slower lip considerably
longer than upper, + emar-
ginate.
Hairs in corolla-tube near
the base.
Stigma purplish.
Ripe seeds usually minutely
and irregularly rugulose,
brown, c. 2? mm. long.
2n
302 MR. €. E. SALMON ON A HYBRID STACHYS.
If one sums up the various differences enumerated above, it will be seen
that by far the larger proportion of the characters of the hybrid in my garden
are those of germanica rather than alpina, the last-named showing n" in
(1) the strong peculiar scent of the leaves, (2) the presence of glandular
hairs, and (3) the eharacteristie seed.
Fic. 1
a. Stachys germanica, flower aud seed.
b. S. digenea, ditto.
c. S. alpina, ditto.
All enlarged.
It is, I think, rather a remote hope that the hybrid described may be
found in a wild state in these islands, for the only station for Stachys alpina,
which is in West Gloucestershire, lies some 30 miles away, in a west-south-
westerly direction, from the nearest known locality for 5. germanica n
Oxfordshire.
I do not know, however, the limits as regards the carriage of pollen by
insects and the distances proved to have been covered.
I wish. to thank Mr. Britten, Mr. Skan, and Mr. J. W. White for help
in the preparatiow of this note, and to say that I shall be happy to supply
seeds of the hybrid as far as the stock will allow.
ON SOME CRITICAL SPECIES OF ECHIUM. 363
A Revision of some Critical Species of Echium, as exemplified in the
Linnean and other Herbaria ; with a Description of Echium judwum,
a new Species from Palestine. By C. C. Lacarra, F.L.S.
[Read 6th June, 1918.;
INTRODUCTORY.
THE five papers collected under the above title are offered as a quarry, from
which any future monographer of the genus Echiwm may dig material.
They are :—
1. Five Critical Species of Echium: E. judeum, mihi, E. australe,
Lam., Æ. Coineyanum, mihi, E. pyenanthum, Pomel, f. salman-
ticum, Lag., pp» 366-379.
2. The Genus Echium in the Herbaria of Tournefort, Jussieu, and
Lamarck, pp. 379—384.
3. The Echia of Sibthorp's Herbarium, pp. 384-392.
4. The Linnean Species of Echium, pp. 392-427.
5. The Echia of Miller's * Gardener’s Dictionary,’ pp. 427—438.
The European Echia have been very badly treated by Linnæus. Out of
the nine species mentioned by him or existing in his herbarium, one,
rubrum, Jacq., though represented by no less than three specimens, seems
entirely to have escaped his notice, the older synonym for it being twice
quoted, but each time under a different species. Two others, ereticum and
riolaceum, are such hopeless entanglements of contradictions that neither
name can be used and both must be abandoned. Lusitanicum, though
recognisable with certainty from the synonyms and the specimen, is Sail
BEPARI There only remain vulgare, italicum, pyrenaicum, and planta-
gineum, of which the descriptions are quite clear, though with regard to the
last three tedious discussions are necessary to clear away the cobwebs spun
by later botanists.
Subsequent authors never studied this genus seriously till de Coincy nd
itin hand. His valuable papers, some of ise posthumous, are scattered in
Morot’s * Journal de Botanique, in Bull. Herb. Boiss., in Act. Congr. Internat.
1900, and elsewhere. His death was a grave loss, but unfortunately he
never consulted the Linnean specimens, or those of other old herbaria in
England, nor does he quote a sufficient number of well-known exsiccata
to bile his views quite clear in all instances. I have been obliged to differ
LINN. JOURN,—BOTANY, VOL. XLIV., 2
364 MR. C. C. LACAITA : A REVISION OF
with him altogether in the eases of Æ. australe and E. pyrenaicum, and to
a lesser degree in regard to Æ. pustulatum. He seems to have devoted most
attention to the species of Algeria, but his chief discovery was the difference
in the arrangement of the scales in the corolla-tube, from which the veins
take their origin. In accordance with this he divides the genus into the
Sections Gamolepis and Eleutherolepis (* Sectionnement du genre Echium”
in Act. Congr. Internat. Bot. 1900, pp. 346-350), though admitting that one
or two species, particularly Æ. maritimum, Willd. (E. confusum, Coincy),
are ambiguous and variable in respect of that character.
The structure of the inflorescence in the genus Echium was studied by
Kaufmann (see his paper * Ueber die Entwickelung der Cyma scorpioidea
bei den Borragineen” in Bot. Zeit. xxvii. p. 886, 1869). What the older
authors called “spikes” in this genus are really what the Germans term
“ Wickeln.” The Latin cincinnus is employed to express the same idea, but
I do not find that any satisfactory English equivalent has come into use.
“Curl,” which is the literal translation of cincinnus does not exactly corre-
spond to Wickel. I would suggest “furl,” which is all the more suitable
because, not being a word in common use, it does not suggest any false
connotation *, At any rate, the verb “unfurl” exactly expresses what
happens to the cincinni as they develop from bud to fruit. The word
'eyme," used by de Coiney and other moderns, is not suitable, for,
although the structure is one of the eymose arrangements, the word itself,
as applied to the visible result of the structure, conveys to the mind a
picture very different from that presented by the inflorescence of any
Echium.
De Coiney, in the same paper on *Sectionnement,? and in his note on
E. simplex in Bull. Hb. Boiss. 2, iii. p. 276 (1903), has given an admirable
account of the arrangement of the flowers in the cincinnus, explaining how
the difference in the several lacini of each ealyx depends on the exact
position of the flower relatively to the rachis, on which their disposition is
distichous and subunilateral.
Most European species of Echium have been described as biennials,
though Coutinho, Fl. Port. (1913), more judiciously marks them © or g
with the exception of lusitanicum (Broteri), polycaulon (salmanticum), and
rosulatum, which are certainly perennial, and parvifforum, which is always
annual. But arenarium, australe, davum, grandiflorum, italicum, planta-
* I must enter a protest against the growing pedantic misuse of the term “ connote,”
especially by writers on scientific subjects, in the sense of plain English “mean” and
occasionally even in the sense of * denote." The name Echium denotes all the plants of the
genus, but connotes the generic characters. The use of the word as an English logical
term was introduced by James Mill, and is explained in Mill's ‘Logic, i. 9, 856. — Connotare
was used by the schoolmen in the 14th Century, but not in the precise sense of the Mills,
See, farther, in Murray's Eng. Dict. sub voc, “ connotative” and “ connote,”
SOME CRITICAL SPECIES OF ECHIUM. 365
gineum, pomponium, pustulatum, pyrenaicum, rubrum, tuberculatum, and
vulgare seem to behave as annuals or biennials, according to circumstances.
In December, 1917, I was able to observe carefully large numbers of
italicum, pyrenaicum, plantagineum, and pustulatum around Rome and near
Taranto. In every case I found that the plant which had flowered in the
summer was a dead skeleton with a rotten root. There were no living plants
of any of those species, except baby seedlings, which would undoubtedly
flower and perish in their turn within 12 months. But in cultivation, and
in climates with a less prolonged summer drought, the evidence seems to
point to a biennial habit, with two possible flowering seasons. /chium
angustifolium, Mill., on the other hand, appears to have taken the opposite
course, for whilst Miller grew it as an annual at Chelsea, we are assured by
Haláesy that in Greece it (Z. elegans) is perennial.
The presence of hairs on the filaments, an excellent distinctive character
for one or two species, has been made too much of by de Coincy for certain
others, in which the hairs are very obscure and very few, and not constantly
present. On the other hand, he has perhaps underrated differences in the
nueules, whieh serve to separate true vulgare from its southern substitutes,
The size of the corolla can only be used with reservation as a specific
character, for in most, if not all, the species, sexual variation affects the size,
sometimes to an extreme degree, as in so-called Æ. Wierzbickii and in the
pistillate form of Æ. setosum, Vahl. Colour is more reliable, but dangerous
to depend on in dried specimens, especially if these have been exposed to the
action of any acid. And it is to be borne in mind that the blue kinds often
throw white or pink individuals.
Echium rubrum stands by itself on account of its undivided style with
a capitate stigma. It is strange that this character should not have been
noticed by Jacquin himself or by De Candolle in the * Prodromus. It would
be an exaggeration to create a new monotypic genus on this ground, but it
necessitates some modification of the received generic definitions of chium.
De Candolle, Prodr. x. p. 16, says: “ stigma bipartitum, stylo nempe apice
bilobo, lobis filiformibus " ; the accurate Bentham, in Gen. PI. ii. p. 863,
“stylus filiformis, apice breviter 2-fidus, stigmatibus parvis." —Gürke in
Engler’s * Pflanzenfamilien,’ iv. 3, p. 128, merely translates Bentham's Latin
into German, without addition or subtraetion of a syllable.
In the course of my researches I have seen all the specimens of the genus
in the following herbaria, besides my own :—
]. Kew.
2. British Museum.
3. Sibthorp’s, Sherard’s, and Dillenius’s plants at Oxford.
4. The Bailey herbarium now in the Museum of Manchester Uni-
versity.
212
366 MR. C. C. LACAITA : A REVISION OF
5. Herb. Mus. Paris, including the collections of Tournefort, Lamarck,
Jussieu, and de Coiney. I had not time to go through the speci-
mens of herb, Cosson, now at the Museum, which are indispensable
for Algerian species.
. Herb. Bonaparte which includes herb. Rouy.
. Erbario Centrale Italiano at Florence, and Desfontaines's specimens
in herb. Webb.
. The herbarium of the University of Rome.
. Tenore's and Gussone’s herbaria at Naples.
-1 7;
ec ooo
I have been sent a few specimens through the kindness of M. Sudre of
Toulouse, Prof. Coutinho of Lisbon, and Prof. Henriques of Coimbra. To
these gentlemen I must express my gratitude, and above all to M. Henri
Lecomte and his assistants at the Paris Museum, and to Prince Roland
Bonaparte, who so generously allows access to his magnificent collections.
My thanks are also due to many in Italy and at home, particularly to
Dr. Daydon Jackson, whom I have troubled with endless small inquiries,
and to Mr. Wilmott at the British Museum for the assistance of his younger
eyes and for counsel, perhaps not always followed, as to technicalities of
nomenclature.
L
FIVE CRITICAL SPECIES OF ECHIUM.
ECHIUM JUDÆUM, sp. nov.
Among the specimens of Fehium from Palestine in the herbaria of the
British Museum and Kew are several labelled plantagineum (or violaceum in
the same sense), although differing altogether from that species in their
indumentum, which is not homogeneous but dimorphous, in the shape of the
bracts, and in the corollas which are more obliquely cut, but less wide at
the throat, and covered with soft pubescence instead of being glabrous with
only some long hairs on the veins and ciliæ on the margin. In fact, the
nearest species is not E. plantagineum but E. grandiflorum, Desf., to which
E. judeum bears a considerable resemblance, but the filaments are glabrous,
at any rate, as seen without the microscope, whereas in grandiflorum they
are constantly furnished with scattered transparent hairs, longer than the
breadth of the filament itself.
E. judeum, mihi; Sect. Fleutherolepis, Coincy.
Radix fusiformis, ille Æ. vulgaris similis. Caules plerumque simplices,
erecti, c. 35 em. alti, indumento dimorpho, se. pubescentia creberrima,
brevi, grisea, et pilis longioribus, sparsis, albis, substrigosis. Folia
radicalia (in exempl. unico ex Hierosolyma presentia) petiolata, ovalia,
medio nee basi latiora, 14 x 41 em. ; penninervia, nervis utrinque 6-8
]
?
SOME CRITICAL SPECIES OF ECHIUM. 367
ante marginem evanidis, non anastomosantibus ; caulina inferiora petio-
lata, anguste elliptico-oblonga ; 5-10 em. longa, 1-2 cm. lata ; suprema
sessilia, sepe exacte lanceolata. Foliorum indumentum caulino simile,
sed pilis rarioribus tuberculo mediocri insidentibus. Tubercula magis
quam in 7. grandifforo conspicua, illis E. pustulati minora. Inflores-
centia e cincinnis axillaribus simplicibus quaquaversis constans, virgineis
valde scorpioideis, in fructu elongatis erecto-patulis. Flores in cin-
cinnis 20—40, sessiles, distichi sed unilaterales, bracteati sed infimo ssepe
ebracteato (folio caulino supposito bractere officio functo), initio conferti,
serius inter se remotiusculi. Bractez lanceolate ; nonnullæ basi parum
latiores at in acumen products. Calyx sericeo-pilosus, pilis albis, usque
ad 2-3 mm. longis, pro genere mollibus, haud aut vix strigosis ; lacinie
lineares, usque ad 12 mm. long:e, sed tubo corollino multo breviores,
post anthesin non ampliate nec (ut in Æ. amano) longe concrete, tubo
tantum in fructu incrassato. Corolla ore obliquo, ille Æ. grandiflori
haud absimilis, in sicco 3-4 em. longa; pubescentia minuta, albo-grisea,
et secus nervos pilis nonnullis longioribus vestita, margine non ciliata.
(‘olor in sicco violaceo-purpureus vel fasciis roseis et pallide ceeruleis
alternantibus. Stamina inclusa, vel duo longiora ultra labium inferius
brevissime exserta, filamentis glaberrimis violaceis, antheris ovoideis,
impuberibus amæœne ceruleis, serius lutescentibus. Stylus hirsutius-
culus, violaceus, apice ad cirea 2 mm. (multo minus profunde quam in
E. amano) bifidus, initio vix aut parum exsertus, postea elongatus.
Nucule videntur (perfectas non vidi) parve, 3 mm. longze, apiculate,
latere interiore acute carinate, tuberculato-rugosw, pallide. Herba
exsiccatione virido-grisea, nec, ut in Æ. grandifforo, tabacina.
Differt ab Æ. grandifloro, Desf.:—(1) Foliorum tuberculis majoribus ;
(2) Bracteis lanceolatis nec lato basi ovatis aut subhastatis; (3) Pilis
cincinnorum multo copiosioribus, albis, longioribus, mollioribus, vix strigosis ;
(4) Calyeis laciniis longioribus; (5) Filamentis glaberrimis ; (6) Herba
exsiccatione non tabacina.
Ab E. plantagineo, L. (etiam Palestine incola) longius distat indumento
dimorpho, foliis caulinis bracteisque basi non semiamplexicaulibus, corollis
magis obliquis sed fauce minus dilatato, passim pubescentibus, non vero
ciliatis, staminibus brevioribus.
Habitat in Palestina. Exemplaria vidi sequentia in herbariis Kewensi
et Musei Britannici.
1. Jerusalem, Haram Court ; leg. M. J. Fox, ii. 1867 (nom. Æ. violaceo).
2. Kallirhoe ; leg. W. A. Hayne, iii. 1872 (nom. Æ. violaceo). 3. Magdala;
leg. B. T. Lowne, 1863-1864 (nom. E. violaceo). 4. Jericho in desertis :
Bornmüller, It. Syriacum, 1897, no. 1136 (nom. Æ. plantagineo). 5. Jericho
in stony places ; Plant. Palæst. Maris Mortu : leg. Meyers & Dinsmore
(nom. E. plantagineo). 6. Beirut; Pl. Syriw littoralis ex herb. Postian,
no, 234, v. 1875 (nom. Æ. plantagineo).
368 MR. C. C. LACAITÀ : A REVISION of
ECHIUM AUSTRALE.
What is Echium australe, Lam. Tll. i. p. 412 (1791) ? De Coiney, in
Morot's Journ. Bot. xiv. p. 326 (1900), identifies it with a plant from
southern Spain, which I shall call Echium Coincyanum, distributed in
Bourgeau’s exsiccata for 1849, no. 334, and for 1852, no. 1625. In a foot-
note he remarks, “ Cette espèce a été confondue tantôt avec I E. ereticum, L.,
tantôt avec le grandifforum, tantôt avec l'angustifolium, Lam. ou Salzm.,
tantôt avec le pustulatum laxum, Herb. Boiss.” In spite of my great respect
for de Coiney's knowledge of the genus Eehium, of which he has been the
only serious student hitherto, I cannot doubt that in this case he has himself
only increased the confusion, for Lamarck’s species is quite unlike the
Bourgeau specimens, being, in fact, a garden form identical with the specimen
of ereticum in the Linnean herbarium *, and undoubtedly derived from some
variety of X. grandiflorum, Desf., probably from that which is found on the
borders of France and Spain.
Consequently, the names Æ. grandiflorum, Desf. (1798), and E. macran-
thum, Roem. & Sch. (1809) +, must be replaced by that of Lamarck, which,
moreover, cannot give way to J£ creticum, L., for, although the Linnean
* See my paper on the Echia of hb. Linn., p. 396.
T The name of grandiflorum for Desfontaines's species was altered to macranthum by
Roemer and Schultes, Syst. iv. p. 20 (1819), “ob Æ. grandiflorum. antiquius Andr. et Vent.”
Rouy, Fl. Fr. x. p. 309, adopts this alteration, also quoting the still earlier Æ. grandiflorum,
Salish, But Æ. grandiflorum, Salisb. Prodr. p. 115 (1796), was only proposed by him as an
equivalent for X. orientale, L, It was therefore still-born and must be ignored. Æ. grandi-
lorum, Andr. Bot. Rep. tab. xx. dated May Ist, 1798, is identical with £. formosum, Pers.
Syn. i. p. 163 (1805), now referred to a different genus as Lobostemon formosus, Lehm., and
E. grandiflorum, Vent. Malmaison, p. 97 (1803), is identical with the grandiflorum of
Andrews,
The preference of Andrews's name is based on the assumption that it was published earlier
than the first volume of Desfontaines's * Flora Atlantica.’ But of this I have not been able
to find any evidence. Many copies of the ‘Flora Atlantica’ bear the date “anno VIII"
on the titlepage of both volumes, which has led to that work being referred to the year
1800. These titlepages are false. Earlier copies—for instance, that of the Linnean Society—
bear the date * anno sexto reipubliez gallicæ” in both volumes, which is untrue for vol. ii.
Evidence of the true dates is to be found in Cat. Bibl. Banks, vol. v.—itself published in
1800, where at p. 71 the first vol. is said to be of “an VI” and at p. 214 the second is
referred to “an VIL" Moreover, on p. 458 of the Linnean Society's copy of vol. ii. there
is a note in Sir J. E. Smith's hand, “ Sept. 25, 1799,” proving that that volume cannot have
come out later than 1799. Now “an VI” ran from Sept. 1797 to Sept. 1798, so that
E. grandiflorum in vol, i. p. 166, may well have appeared before May Ist, 1708. Neverthe-
less, it is strange that Desfontaines himself, in Hist. Arbr. i. p. 177 (1809), should have used
the name grandiflorum in Andrews's and Ventenat's sense without any allusion to his own
grandiflorum. At any rate, the priority of Andrews's name is so doubtful that, on the
strictest interpretation of the rules, it does not justify the abandonment of so well-known
and suitable a title as grandiflorum, Desf., for the Algerian Echium, especially as its rival is
à Lobostemon.
SOME CRITICAL SPECIES OF ECHIUM. 369
specimen is identical with Lamarck’s type, E. ereticum, Sp. Pl., is an intricate
and inextricable mixture of the species represented in the herbarium with
the totally different Æ. angustifolium, Mill. (non Lam.), = Æ. elegans, Lehm.,
and is therefore a nomen confusum that cannot be employed for any
species *, So much the better, for creticum would be an objectionable title
for australe, which is not found in Crete or in Syria, whence Linnæus says
that he had his seeds of ereticum.
Probably de Coiney was led astray by the occasional occurrence of
E. australe with smaller corollas than those of the typical Æ. grandiflurum
of Algeria ; indeed, neither Lamarck’s nor Linnzeus’s specimen has the very
long corollas of that form, for which, if a distinctive name is demanded,
I think it should be Æ. australe, Lam., forma macranthum (Roem. & Sch.),
Coutinho, or, perhaps better, forma grandiflorum (Desf.). But the corollas
are similarly cut in both ; the difference in size is not specific or varietal,
nor associated with different geographical areas. It is parallel to the
variation in corolla-size, often, perhaps always, sexual, that is so well known
in Z. vulgare, I. plantagineum, and E. maritimum, and is most remarkably
displayed in the Egyptian Æ. setosum, Vahl +. But not all cultivated
individuals of australe have the reduced corollas. Sweet, Brit. Fl. Gard.
t. 101 (April 1, 1825), describes and figures Æ. australe raised from seed
received from Germany under that name, showing very long corollas.
E. australe differs from Æ. Coincyanum in the much broader and differently
shaped leaves, ovate, or oval narrowing at the base, not oblong; in their
indumentum, which, though varying much in individuals, has not the soft
pubescence, “duvet très fin” of de Coincy, to form a carpet beneath the
tubercle-borne sete; and in the tobacco-like colour of the dried plant,
whereas Coincyanum keeps greyish. The corollas are differently cut and of
a different colour, the rich red tinge of australe being traceable even in old
exsiecata. The filaments are conspicuously and constantly hairy; not
pubescent, but furnished with long, slender, flexuose, scattered hairs, easily
seen even in sicco. Sweet, loc. cit., mentions “very hairy filaments.” The
hairs on those of Coincyanum are extremely scarce, not present in every
flower, perhaps not even in the majority, and very difficult to observe in
herbarium specimens. The fruiting calyces are much wider apart in
australe, and the nucules remarkably large, as in Algerian grandiflorum,
twice the size of those of Coinceyanum. It is unfortunately impossible to
distinguish how much of de Coiney's elaborate description of australe is
drawn from Bourgeau’s specimens, and how much from Lamarck’s or others
of true australe.
Lamarck gives very short diagnoses of his species of Kehium; of his
no. 1860, he only says: “ Æ. australe foliis caulinis ovatis, utrinque attenu-
* See my Echia of hb. Linn., infra. p. 396.
T See de Coincy in Morot, Journ. Bot. xv. p. 322 (1901).
310 MR. C. C. LACAITA : A REVISION OF
atis, staminibus corollam wquantibus. Ex Eur. austr.? Æ. lusitanicum?”
He is obviously describing a garden plant, whose native home he does not
know. Seanty though the diagnosis may be, the description of the leaves is
sufficient to exclude with certainty Æ. Coincyanum, as represented by the
Bourgeau exsiecata, which have oblong stem-leaves, wider or narrower, but
not ovate nor attenuate at both ends. This character, in fact, excludes all
European and Mediterranean Kehia, except E. grandiflorum and LE. rosulatum,
Lange.
Let us follow the advice of Rouy, Fl. Fr. x. p. 309, note, who, though he
adopts de Coincy’s wrong identification of australe with Bourgeau’s speci-
mens, nevertheless recommends us to interpret Lamarck’s scanty diagnosis
by means of Poiret's good description in Dict. Encyc. viii. p. 672 (1808),
which is so precisely applicable to Lamarck’s type of australe * and to the
creticum of hb. Linn. that it is worth transcribing :— Il y a de si grands
rapports entre cette espèce et l'/eAum grandiflorum, qu'on serait tenté
d'attribuer leurs différences aux localités ou à ia culture. Celle-ci a assez
généralement ses fleurs plus petites et ses feuilles plus rudes, Ses tiges sont
droites, hautes d'un à deux pieds, rameuses, parsemées de poils rudes,
allongés, cendrés ; garnies de feuilles ovales, un peu oblongues, rudes, médio-
crement hérissées de tubercules et de poils roides, épars ; à peine veinces,
entières, ciliées à leurs bords, rétrécies presque en pétiole à leur base, un peu
aiguës ou obtuses à leur sommet; les inférieures beaucoup plus amples :
les supérieures plus étroites et plus courtes ; celles des derniers rameaux et
les florales sessiles, élargies à leur base, presqu'acuminces.
* Les fleurs sont disposées en épis allongés, feuillés, très hérissés; le calice
divisé en cinq découpures droites aigués ; la corolle d'un grandeur médiocre,
un peu pileuse, une fois plus longue que le calice, de couleur violette ou d'un
rouge pourpre; le tube aussi long que le calice ; le limbe tronqué oblique-
ment, à cinq lobes inégaux; les ¢tamines aussi longs que la corolle,
légèrement pileuses, ainsi que le style : le stigmate bifide.
“Cette plante croit dans les contrées méridionales de l'Europe. On la
cultive au Jardin des Plantes de Paris. (V. v.)
In addition to Bourgeau's examples, de Coincy also quotes for his compound
australe the specimen so named in Lamarck’s herbarium and another in that
of Jussieu t. He consequently enlarges his description to include these as
well as Bourgeau’s, altering the character of the stem-leaves to “ ovales ou
oblongues ” so as to cover both species. But he was aware that Lamarck’s
and Jussieu's specimens were not identical with Bourgeau’s as appears from
the following observation : * L’australe existe dans l'herbier du créateur de
* See my note on Lamarck’s herbarium, specimen no. 15, infra, p. 384.
T Both of these I have examined and found identical with each other. One of those in
bb. Jussieu, no. 6619, is labelled as coming from the Jardin des Plantes, “ex hort. reg.
Paris "
SOME CRITICAL SPECIES OF ECHIUM. 311
l'espèce, mais, comme il arrivait souvent aux anciens botanistes, il l'a établie
sur un exemplaire qui porte les signes évidents de culture. Il s'ensuit que
certains caractères ont pu étre altérés, c'est ce qui est arrivé pour les feuilles
qui sont ovales dans les échantillons des herbiers Lamarck et Jussieu et
oblongues dans ceux récoltés par Bourgeau." Just so; but the remark
about the leaves is going too far ; the species which has been modified by
cultivation is the essentially similar “ grandiflorum,” not the essentially
different Coineyanum.
Then de Coiney goes on: “son inflorescence lâchement paniculée lui
assigne une place à part et ne convient à aucune des espèces qui ont été
confondues avec elle." This is strikingly true of the Lamarck and Linnean
specimens, and also of the more luxuriant individuals of Algerian grandi-
florum, but does not hold good for Bourgeau’s plants, in which the
inflorescence shows no marked difference from that of the forms called
pustulatum by writers on the flora of Spain.
This should be enough to clear the Bourgeau specimens out of the way,
and to exclude the possibility of Æ. Coincyanum being the progenitor of
Lamarck’s australe. It remains to consider from what form of grandiflorum
that garden plant ean have arisen. E. grandiflorum occurs in five geo-
graphical areas: 1. Algeria and the adjacent regions; 2. Sardinia aud
Corsica ; 3. The French department of the Var; 4. North-Eastern Spain
and the adjacent district of France; 5. Portugal. Areas 4 and 5 are the most
likely to have supplied the seed, and I should have said preferably 4, where
individuals almost identical with the cultivated plant occur, but that Lamarck
himself has hinted at Portugal. Area 3 is most improbable, because the
variety that predominates there is very unlike Lamarck's.
1. The Algerian plant is very abundant and practically only differs from
Lamarck’s type in its somewhat longer corollas. Desfontaines says : “ affine
I. australi Lam. differt foliis levibus aut tuberculis vix conspicuis conspersis,
corolla duplo triplove majore." The relative size of the corolla is here much
exaggerated. Many, though not all, Algerian specimens show leaves with
enough tubercles to be undistinguishable from those of Lamarck. As far as
habit is concerned, that garden form looks like an extreme case of the
more diffuse and broad-leaved Algerian specimens from rich situations.
M. Battandier, the father of Algerian botanists, writes “les feuilles varient
assez de dimensions suivant que le sol est plus ou moins fertile." There are
plenty of Algerian specimens in the herbaria. I will only quote Reverchon,
Pl. Alg. 1896, no. 68, from Bougie, and Faure’s exsicc. from Oran à Santa
Cruz, as luxuriant examples, the latter with smaller flowers than usual.
The middle stem-leaves of these are trom 2 to 3 cm. broad.
2. I have not seen enough material from these islands to speak of the form
that occurs there. Moris indicates creticum, in the sense of australe, from
Sardinia, Fl. Sard. iii. p. 128, tab. xevii., and his specimens bear out the
TE NOTARIIS
Bez MR. C. C. LACAITA : A REVISION OF
determination. In Corsica the species seems to be exceedingly rare. I have
only seen one specimen, in hb. Mus. Paris ex hb. Grenier labelled
* FE. ereticum, sommet calcaire plus haut du Pic de Pigno sur Nouzo.”
Viviani, Fl. Cors. Diagn. p. 3 (1824), quotes macranthum = grandiflorum
from Bonifacio.
3. The specimens from the district between Hiéres and Nice belong to a
quite special narrow-leaved variety, which I think merits a name and might
be called var. provinciale. The stem is erect and the inflorescence narrow.
The root-leaves measure 6-10 em. by 1-2 cm.; those of the stem are only
about 5 mm. broad. Their indumentum is softer, and their appearance when
dried paler and more buff-coloured than in the other forms. The corollas
are of full Algerian size. Of this variety I have seen many examples from
the Esterel range, from Agay, Fréjus, Antibes, Roquebrune, Bormes, Hiéres,
and from the summit of the Maures range *à la Sauvette.” I do not think
that Lamarck’s or Linnzeus's plant can have arisen from a form so different
in leaf-shape, habit, and general facies.
4. The plants of. the Spanish borderland are mostly rather diffuse, with
broad leaves, rather rougher than those of the Algerian plant, and with
corollas of very variable size, some as long as in Algeria, others at least one-
third shorter. These are probably the origin of the cultivated plant. But
the shorter corolla, as said before, does not indicate a specific difference.
Some of the following exsiecata are labelled australe, some creticum, some
grandiflorum :—
(a) from France.
1. Soc. Dauph. no. 3822 from Banyuls-sur-Mer (in hb. Lacaita).
2. Banyuls ; ex. hb. Loret, hb. Timbal-lagrave, and hb. Gautier.
3. Cerbére à la Tour du Midi (in hb. Bonaparte).
4. Vallée de Consolation près Collioure (hb. Mus. Paris).
(b) from Spain.
1. Cadaques (Catalonia); Sennen Pl. D'Esp. no. 828 as *ereticum
forma grandiflorum."
2. Rosas near Gerona (in hb. Bonaparte).
3. Figueras (lib. Bonaparte).
4. Cambrils (Tarragona) (hb. Bonaparte).
5. Benicarló (Castellon de la Plana) (hb. Bonaparte).
6. Port Bou (hb. Bonaparte).
9. I regret that I have not been able to see any specimens from Portugal,
but Coutinho has dealt with the species so well in his Fl. Port. p. 510 (1913),
that we may safely accept his view, from which it is clear that Lamarck’s
plant might have come from Portugal. Coutinho distinguishes two varieties
of australe: (a) genuinum, with branching habit and corollas of 13-20 cm.,
SOME CRITICAL SPECIES OF ECHIUM. 313
which would correspond to the garden form under discussion, and (b)
macanthum with simple or nearly simple stems and corollas of 20-30 mm.
This may correspond either to the Algerian grandifforum or to var. provinciale,
but such closer determination is immaterial as far as the plant of Lamarck is
concerned.
Among the old specimens in the British Museum there are several that
agree with Lamarek's type *. on which Solander has altered his original
determination of eretieum to australe, perhaps regarding the name creticum
as belonging to the very different Z. creticum angustifolium rubrum C. B. P.—
E. elegans, There are also two specially interesting examples of /. australe
Lam. in hb. Gay at Kew, both from the herbarium of L. C. Richard, On
both Gay has noted “ proeul. dubio cultum " and added on one label, “ Je
soupçonne que cette plante est provenue des graines de PE. grandiflorum,
Desf.," and on the other “ J'ai vu cette plante dans l'herbier du Muséum,
dans ceux de Desfontaines, de Jussieu, de Delessert, etc. ‘Tous les
échantillons qui s'y trouvent ont été cultivés. Celui de l'herb. du Muséum
a été pris au Jardin des Plantes le 3 Juillet, 1813. Tous sont remarquables
par leurs feuilles larges, amincies à la base, et par leurs braetées trés
allongées, les inférieures larges et Foliacées." — Gay's herbarium also contains
the two specimens of. Bourgeau's quoted by de Coincy. The determination
on their labels, “ E. angustifolium ? Lam. Salzm.," is due to Gay himself, who
was a very close observer, and proves conclusively that he knew they were
not F. australe, Lam., as grown in the Jardin des Plantes and as represented
in herb. Juss.. Gay could not have seen Lamarck’s type, for in his time
Lamarck’s herbarium was in Germany.
It will be seen from what has been said that I do not accept the arrange-
ment in Rouy, Fl. Fr. x. pp. 309, 310, where australe and macranthum are
treated as distinct species on the ground of different size of the corollas and
of the nueules, without any reference to different leaf-characters. But Rouy
has obviously copied his measurements from those of de Coincy, quoting
Bourgeau’s specimens together with Soc. Dauph. 3822 for australe. The
nucules described for that species as “ trés-petites, 2-23 mm.” obviously
belong to Coincyanum and not to australe, Lam. Lamarck’s type-specimen
of australe has no ripe nucules ; see p. 384 belowf.
* See a list of these in my Echia of hb. Linn., infra, pp. 401, 402.
+ Since the above was written I have grown E. grandiflorum at Selham in Sussex from
seed sent by M. Battandier of Algiers. Two sowings were made. The earlier flowered in
July and resembled the Algerian plant in habit and size of corolla. But the later-sown
plants, which did not begin to flower till September and continued till the hard frosts, were
leafy and diffuse, with smaller corollas, exactly resembling the creticum of Herb. Linn. and
confirming the opinion expressed in the text as to the specific identity of australe, Lam.,
grandiflorum, Desf., and creticum, Herb. Linn.
314 MR. C. C. LACAITA: A REVISION OF
Ecnuiuw CorNCYANUM, nom. nov.
Echium Coincyanum, mihi, — E. australe, Coiney, Rev. Esp. Crit. Echium in
Morot’s Journ. Bot. xiv. p. 326 (1900), quoad exempla Bourgeauana, Rouy,
Fl. Fr. x. p. 309 (1901), non Lam. Ill. i. p. 412 (1791), nee Poir. Dict.
Encycl. viii. p. 672 (1808) =F. ereticum, Nym. Consp. Fl. Eur. p. 515 (1881),
et. Willk. et Lange, Prodr. Fl. Hispan. ii. p. 487 (1870), pro maxima parte,
non L.— E. angustifolium, Salzm. (et aliorum) in schedis, non Lam.
Tn my note on X. australe, Y have pointed out that de Coiney confused two
different species under that name, having united to the true australe, Lam.,
a plant best known from certain exsiccata of. Bourgeau, to distinguish which
I propose the name Coneyanum. It would have been prsterable to call this
species after M. Bourgeau but for the existence of Æ. Bourgeanum, an
arborescent species iom Teneriffe,
Echium Coineyanum is based on Bourgeau's no. 334 of the year 1849 from
the Tajo de Ronda in Andalusia, no. 1625 of 1852 from the Sierra San Felipe
de Jativa, and I may add, though it is not mentioned by de Coincy, his
identical no. 989 of 1850 from the Sierra de Segura. The Kew herbarium
contains two examples of eaeh of these numbers in excellent condition.
No. 334 was determined by J. Gay as “ E. angustifolium? Lam. Salzm.”
meaning that though uncertain whether it really is Lamarck’s angustifolium
(which it is not) he considered it the same plant that Salzmann had collected
“ad vias circa Malagam abunde ” and labelled * Æ. angustifolium, Lam. DC.”
This determination of Gay’s proves that he knew the plant not to be
E. australe, Lam. of herb. Juss. ete., and as grown in the Jardin des
Plantes. The similar naming of nos. 989 and 1625 is due to Cosson.
I have examined these species with some care. The stem-leaves are not
oval but oblong ; the corollas are large, 20-25 mm., mostly blue (in sicco),
not dull reddish as in Æ. australe, and less obliquely eut. The presence of
hairs on the filaments of E. Coineyanum is not quite certain and should be
carefully observed on the living plant. De Coincy says “au moins un des
trois filets postérieurs poilus, ordinairement tous les trois,” and in a footnote
“ Pai trouvé des exceptions.” It is impossible to say whether these
observations of his were made on specimens of Coincyanum or on true
australe, but I think more probably on Coineyanum. In Bourgeau’s
specimens I could find none on the first examination, when two corollas
were boiled for dissection, although I was assisted by the defter fingers and
younger eyes of Mr. Hutchinson of the Kew herbarium. Subsequently in a
corolla opened without boiling I found a very few sparse hairs on two
filaments. My apparently contradictory results seem to agree with
de Coiney’s. The filaments of australe, on the contrary, are always
remarkably hairy.
SOME CRITICAL SPECIES OF ECHIUM. ato
Besides the three Bourgeau numbers, which must be regarded as the types,
I consider the following specimens to be referable to Æ. Coincyanum * :—
1. Salzmann in hb. Gay ; ad vias circa Malagam ; as F. angustifolium,
Lam.
. Willkomm anno 1845, no. 902, from Malaga as pustulatum.
3. Porta et Rigo, it. ii. hisp., no. 151, from Almeria as ereticum.
Wolley-Dod anno 1912, nos. 412, 473, and 610, from Gibraltar at
the Mediterranean steps as ereticum.
5. Roffey, March 1, 1916 (in hb. Mus. Brit.), from Tajo de Ronda.
bho
Apr
I have met with no specimen of Coineyanum in the old herbaria earlier
than that of Salzmann.
ECHIUM rycsANTHUM — E. ANGUSTIFOLIUM, Lam. (1791), non
Mill. (1768), nec Thunb. (1811).
Echium angustifolium, Mill. Gard. Dict. (1768), and E. angustifolium, Lam.
Ill. i. p. 412 (1791), were wrongly synonymised by Poiret in Diet. Encycl.
viii. p. 671, since when Miller's species has been entirely ignored or quoted
without any discussion of its identity.
It is undoubtedly the plant known to the old botanists as Æ. ereticum
angustifolium rubrum, C. B. P., and, as I hope I have established in my notes
on Miller's Echia, is the oldest name for E. elegans, Lehm., plentiful in
Greece and the Levant. But Æ. angustifolium, Lam., is a totally different
species from Spain, closely allied to Æ. humile, Desf., and figured in Barr. ic.
1011 as Lycopsis angustifolia minor hispanica. After Lamarck’s time it
became a source of perplexity and was confused by Salzmann, J. Gay, and
Cosson with /, Coincyanum, mihi. Eventually Rouy rediscovered the species
in 1879 at Jativa and at Hellin, the latter station being close to Tobarra,
whence Lamarck had the type-specimen still to be seen in his herbarium.
Rouy describes it fully in his Exe. Bot. ii. p. 16 (or in Bull. Soc. Bot. Fr.
xxix. p. 123). It has been discussed at length by de Coiney in Morot's
Journ. Bot. xiv. p. 106 and xvi. p. 215.
* A good many other Spanish specimens of * pustulatum " and *ereticum" probably
belong to Coincyanum. Collectors meeting with plants that look like possible pustulatum
in Spain should make quite sure in vivo whether the filaments are perfectly glabrous, as in
true pustulatum, Sibth. Porta & Rigo, It. iii. no. 142, seems to be a mixture, being an
instance of the bad practice of those collectors in distributing specimens from more than one
locality under the same number. This no, 142 is labelled “ supra Calpe,” which is on the
coast between Valencia and Alicante, and also * Sierra de la Fuensanta," which is inland
near Murcia.
On a specimen in lib. Cosson from S.W. Morocco leg. Mardochée, 1875, Rouy has pencilled
“australe.” It is certainly not australe Lam., but I think it is Coineyanum, which very
probably extends to Morocco. I had not time to search for it in Cosson's magnificent
herbarium of North African plants now at the Paris Museum.
216 MR. €. C. LACAITA : A REVISION OF
Lamarck’s species is very near Æ. humile, Desf. (1791), which is not
known from Spain, although Porta and Rigo distributed angustifolium, Lam.,
under that name. De Coincy separates these two specifically in Journ. Bot.
xiv., but in his Enum. Echium Fl. Atl. in vol. xvi. Æ. humile and some other
North African chia are subordinated as varieties to Z. angustifolium, Lam.
Unquestionably the name angustifolium, Lam., must give way to angusti-
folium, Mill., and Lamarek's species receive a new name, Now de Coincy
identifies with angustifolium a certain E. pyenanthum, Pomel, which is the
name substituted by that author in Nouv. Mat. Fl. Alp. fase. 2, p. 276
(1876) * for his earlier X. densiflorum, described fully in Nouv. Mat. p. 92
(1874), but abandoned as a name owing to the existence of E. densiflorum,
DC. Cat. Monsp. p. 108 (1813), a shrubby species introduced from “Teneriffe
or Madeira," though apparently no longer found there.
E. pycnanthum, Pomel, is therefore the lawful name by which the angusti-
folium of Lamarck should be cailed, or, if reduced to a variety, Æ. humile,
Desf., var. pycnanthum (Pomel), nov. comb. (= E. angustifolium, Lam.. var.
pyenanthum, Coincy ).
EcHICM SALMANTICUM.
The identity of this species can only be established by cireumstantial
evidence, as no authentic specimens are known and all that Lagasea says
of his species in Nov. Gen. et Spec. p. 10 (1816) is *foliis radicalibus
lanceolatis ; staminibus longissimis glabris, corollæ fauce subpervia.
E. Lusitanicum, Linn. ? Hab, cirea Salmanticam. Augusto floret.”
The purport of this note is to make known J. Gay’s description ew vivo of
the plant cultivated in his day as Mehium salmanticum, and to support the
position taken up by Coutinho in Bol. Soc. Brot. xxi. p. 115 (1905), where
E. salmanticum of Lagasca is identified with Æ. polycaulon, Boiss., Diagn. xi.
p. 92 (1849), rather than with Æ. lusitanicum, L.= E. italicum, Brot.=
FE. Broteri, Samp. Boissier’s species was described from a plant in herb.
Pavon from the valley of Plasencia in Estremadura, and carefully
distinguished by him from Æ. lusitanicum, L., as represented in Jussieu’s
herbarium. Bat practically this plant is only known from specimens
collected by Bourgeau (no. 2467 of the year 1863) on the banks of the river
Plasencia. These were distributed at first as Æ. vulgare, but the label was
afterwards corrected by Reuter to Æ. polycaulon.
The arguments in favour of the identification of E. salmanticum with
* In the separate copies p. 296, but p. 40 where included in Bull. Soc. Sci. Alg. for
1876. There is no heading * ZZ. pycnanthum,” but a single line introducing the name forms
the last of the paragraph describing E. onosmoides. Battandier had referred E. pyenanthum
to E. sericeum, Vahl, a very different species, in Batt. et Trab., FI. Alg. p. 609 (1888), but
in Fl. Synopt. Alg. et Tun. p. 235 (1904), and in the suppl. to Fl. Alg. p. 67 (1910),
de Coincy’s opinion is accepted and pyenanthum is quoted as synonymous with angustifolium,
Lam.
SOME CRITICAL SPECIES OF ECHIUM. aui
E. polycaulon are: (1) The relative proximity of Salamanca to Plasencia,
whereas lusitanicum has only been found hitherto in northern Portugal and in
Galieia in three localities in the frontier province of Orense (see Merino,
Fl. de Galicia, ii. p. 155, 1906) ; (2) That a plant identical with Bourgeau's
specimens of polycaulon from Plasencia existed under thename of salmanticum
in several botanical gardens long before Boissier described his species, and in
fact within 10 to 20 years of Lagasca’s publication, thus raising a strong
presumption that the seeds had come from Lagasca himself.
Of these early garden specimens of. salinantieum identical with polycaulon
the following have come to my knowledge: (1) Herb. Boiss. ex Hort.
Genev., leg. Duby anno 1822; teste Buser ex Coutinho, 1. c. ; (2) Leipzig,
anno 1835 ; teste Braun in litt, ined. ad J. Gay; (3) Hort. Neap. Novembri
1836 ; misit Tenore in Herb. Gay ; (4) Hort. Carlsruh., unde semina misit
Braun ad J. Gay anno 1839; (5) Hort. Paris. in Herb. Gay, anno 1843 ;
(6) H. R. Neap. in herb. Ten.
The difference between Æ. salmanticum = polycaulon and E. lusitanicum, L.
= Broteri, Samp., was recognised by J, Gay in his MS. notes in Herb. Kew
before the publication of polycaulon. In addition to the specimen of
lusitanicum received from A. de Jussieu, to which I refer in my notes on
E. lusitanicum of Herb. Linn., Gay’s herbarium contains two good examples
of a plant grown in the Jardin des Plantes in 1843 from seed sent from
Carlsruhe by A. Braun on Feb. 1, 1839, as Æ. salmanticum, Lag. Both are
identical with Bourgeau’s no. 2467 referred to above. Now lusitanicum and
polycaulon resemble each other in their remarkable method of growth ;
Coutinho, Fl. Port. ii. p. 499, says they are perennial plants producing
lateral stems below the rosette of basal leaves *. This had been described
for lusitanicum by Brotero, Fl. Lus. i. p. 290, as follows : * Caules quinque
ad duodecim ex eadem radice, sub rosula foliorum radicalium, primum
obliqui, dein ineurvati, erectiusculi ; folia radicalia in orbem prostrata,
pedalia et longiora, ad medium biuncialia aut latiora, lanceolata." Gay
observed the same character in the specimens he saw alive in the Jardin
des Plantes, but not in the scraps of lusitanicum from the herbaria of
Tournefort and Jussieu. He consequently thought that his cultivated plant
must be identical with Brotero’s italieum — lusitanicum, L., and suggested
for it, but fortunately did not publish, the name Æ. Broterianum.
On Feb. 1, 1839, Braun writes from Carlsruhe to Gay: ‘ Von Echium
salmanticum schicke ich Samen ; gehen sie Ihnen auf, so bekommen Sie die
merkwürdigste Species aus diesem Genus, die von allen andern günzlich
abweicht durch ihren Wuchs. Als Sie meine “chia im Jahre 35 durchsahen,
schrieben Sie an ein schlechtes Exemplar aus dem Leipziger Garten: * Species
* Another perennial species from Spain and Portugal, Æ. rosulatum, Lange, has the same
habit,
378 MR. C. C. LACAITA : A REVISION OF
mihi obscura, Æ. lusitanico, L., similis sed corollis dimidio majoribus violaceis-
que non carneis certe diversa. Forte Æ. plantagineum parviflorum." This
suggestion is of course impossible, and the rest of Braun's letter is devoted
to distinguishing his salmanticum from plantagineum, with a rough sketch to
show the dissimilar method of growth. The seeds germinated and on 25th
June, 1843, Gay gathered in the Jardin de l'école de médecine de Paris one
of the two specimens now in his herbarium, which at first he took to be
& FE. lusitanicum, L., et Lehm. E. italicum, Brot.,” annotating it“ E. caulibus
ex una radice pluribus, rosule centrali subjectis." But in a further note of
July 2nd he modified the synonymy as follows: “Echium Broterianum, Gay
=F. italicum, Brot, sed Echium amplissimo folio lusitanicum, "Tourn. =
E. lusitanicum, Poir., est planta longe alia. Longe alia videtur Æ. caule
simplici foliis caulinis lanceolatis, ete., Royen, E. lusitanicum, Linn. et Lehm.,
cujus laciniæ calycinæ ex Lehmanno lanceolate sunt et acuminatæ, non vero
(que Broterianæ) obtusiusculæ oblongæ vel ellipticæ.” He seems to have
thought there were three species where only two exist and erred in referring
Brotero's italicum to the plant from the Carlsruhe seeds instead of to the
other kind, Æ. lusitanicum.
On the same day, July 2nd, he gathered the other specimen now in his
herbarium from the same seeds, and described it ex vivo as follows (it is the
only deseription of this species from the living plant) :—
* Echium salmanticum, Lag. En pleine terre, ou il forme une énorme
touffe de 2-3 pieds de haut. Ab ÆEchio lusitanico herb. Jussi:ani diversis-
simum. (1) Inflorescentia quasi racemosa non paniculata (spicis partialibus
scilicet brevissimis non elongatis); (2) calyce quam corolla plus dimidio
breviore, non corollam subsequante, phyllis ellipticis vel oblongis, non
lanceolatis aeutatis, margine et ad carinam ciliatis, ceeterum glabris,
non sparsim et longius setosis; (8) corolla ampliore multo imprimis latius
aperta (limbo campanulato non cylindraceo-conico). Pili eaulini uniformes,
longi, patentissimi, firmuli, hasi non bulbosi neque pube ulla breviore inter-
mixta. Corolla violaceo-ceerulea, adhue clausa vinoso-rosea, 34 lin. longa,
ore maxime hiante 3 lin. lato, lobis limbi duobus superioribus paulo longiori-
bus. Filamenta longe exserta, corollam plus duplo longa (sic). Caules ex
una radice plurimi, spurii (revera rami radicales), rosulee centrali (caule
haud evoluto) subjecti!” Then on a much later ticket he added
* FE, polycaulon, Boiss. ? "
The above account should be compared with Coutinho’s description of
polycaulon (salmantieum) in Bol. Soc. Brot. xxi. p. 111, and will be found
to agree in essentials, There is a photograph of a herbarium specimen of
E. polycaulon in Rouy’s Ilustr. Pl. Eur. Rar. tab. xli. It is strange that no
one as yet should have taken the trouble to rediscover /. salmanticum in such
an accessible locality as the neighbourhood of Salamanca. Both this plant
and Æ. lusitanicum seem to have been lost from the gardens where they once
were grown,
SOME CRITICAL SPECIES OF ECHIUM. 379
There is an Æ. vulgare, var. salmanticum, Coiney, in Morot’s Journ. Bot, xiv.
p. 304 (1900), which must not be confused with Æ. salmanticum, Lag. The
specimen in de Coincy’s herbarium, gathered by him at Salamanea on
May 28th, 1892, on which his var. salmanticum is based, is very poor, hardly
in flower, and I should have thought altogether too scanty to justify a new
name. I rather doubt its being a form of vulgare.
TE
THE GENUS ECHIUM IN THE HERBARIA OF TOURNEFORT,
JUSSIEU, AND LAMARCK.
These three famous herbaria, now in the Muséum d'Histoire Naturelle at
Paris, are so important for the genus Echium that the following list of the
specimens they contain may be useful. The first two have been examined
and annotated by that minutely observant botanist, J. Gay. The third was
not accessible to him, owing to its strange adventures. J, B. Monnet,
chevalier de la Marck, ended his life in straightened circumstances in 1829,
A few years before (1824) his herbarium had been bought, at the suggestion
of Alexander von Humboldt, by J. A. C. Roeper, who took it with him to
Bale, and thence to Rostock, where he filled the chair of botany. From
Roeper it was bought by the Grand Duke of Mecklenburg for the University
of Rostock, and there remained till 1887, when it was offered for sale to pro-
vide funds of which the University was in need, and was purchased, with
the help of the French government, for the Museum of Paris * -—
I. Hers. TOURNEFORT.
no. 584 Sine nomine aut loco ; leaves only ; undeterminable.
» 985 “ Echium majus et asperius flore albo," labelled by Gay in 1834
“ E. pyrenaicum Desf.," which it is not. The specimen consists
of the upper part only of two stems, and is a wide-spreading form
of E. italicum, L., which the French botanists often confuse with
E. pyrenaicum.
586 “ Echium majus et asperius flore dilute purpureo Bot. Monsp.,”
labelled by Gay “ E. pyrenaicum Desf.," which in this case is
correct. It consists of two branches broken off a stem.
987 no label of origin, but labelled by Gay * Verisimiliter Ech. am-
plissimo folio lusitanicum (specimen minus evolutum)= Echium
lusitanicum Linn. Poir. etc." This is exactly the lusitanicum of
herb. Linn. =Æ. Broteri, Sampaio.
* This information is collected from Bureau, “Sur l'entrée de l'herbier de Lamarck au
Muséum d'Histoire naturelle," in Comptes Rendus, Ac. Sci. Par. 104, p. 187 (1887), and
Bonnet, * L'herbier de Lamarck” in Morot's Journ. Bot. vi. p. 129 (1902).
LINN, JOURN.—BOTANY, VOL. XLIV, 2K
MR. C. C. LACAITA : A REVISION Or
* Ehium vulgare paniculatum crispum,” not labelled by Gay. A
lusus or deformity of W. vulgare. The plant is in bud only.
“ Echium ereticum angustifolium rubrum O. D.," labelled by Gay
« FE. ereticum a Poir. Dict.” This is precisely Æ. angustifolium,
Mill.== E. elegans, Lehm.
“ Echium sylvestre hirsutum, maculatum C. Bauh.
FE. rubrum, Jacq.
“ Echium maritimum. Insularum. Stechadum, flore maximo ceruleo,”
labelled by Gay **Je erois que cette forme rentre dans le grandi-
lorum. Dest., qui selon moi, constitue une espèce bien distinete
du plantagineum?” He is right; it is the narrow-leaved form of
grandiflorum that is found here and there in Dept. Var.
* Echii genus in maritimis insult. Porquerolles,” root-leaves only :
undeterminable.
no name but “Fouqué,” labelled by Gay “Echium orientale folio
oblongo molli et cinericeo, Tourn. Cor. 6? E. pustulatum 'Ten.* in
herb. mus. Par. non Sibth.” Then Chaubard notes ** /ehiwm
vulgare pilis adpressis Viv. Fl. Agen. Il n'est pas d'orient, mais
de la France méridionale." The specimen is one of the forms that
replace typical vulgare in southern France, sometimes determined
by French botanists as pustulatum, sometimes as «tuberculatum,
though not the tuberculatum, Hoffmg. et Link, nor the true
pustulatum of Sibthorp. See my notes on Sibthorp’s herbarium.
“ Echium Cupani,” labelled by Gay “Echium calycinum | Viv."
which it is.
no name, but labelled by Gay “ Echium vulgare L." to which |
”
is unmistakably
assent.
no name, but labelled by Gay “ Echium plantagineum (hortense fol.
floralibus solito majoribus, flore paulo minore).” I assent.
no name ; no label by Gay; a branched plant in fruiting state,
apparently of the group that represent vulgare in the south, but
I eannot venture on a close determination.
no name and no label by Gay ; it is ZZ. flavum, Desf.
no name ; an unknown hand has suggested in pencil Æ. arenarium,
Guss., which is right.
“ Echium zEgyptie, flore magno suave rubente," labelled by Gay
“ Echium Rauwolfii Delil.," which is right.
Tournefort’s herbarium contains no specimen of his Æ. orientale,
verbasci folio, flore maximo campanulato.
SOME CRITICAL SPECIES OF ECH!UM. 381
II. Hers. JussrEv.
no. 6601 Echium unnamed “ex hort. Bagatelle 1788.” Only the upper half
I
6603
6604
6605
6606
6607
6611
6612
6613
6614
6615
6616
6617
6618
6619
of a plant which is obviously Æ. lusitanicum herb. Linn.— Brotert,
Samp.
contains three sheets with no original name, all labelled by Gay
“ pyrenaicum,” but only two belong to that species, the third
being italicum, L.
labelled by Gay “Æ. italicum L., Willd., Lehm et aliorum nonnull.,”
is either true italicum, L., or glomeratum, Poir. For lack of time
I could not examine carefuliy.
two sheets :
(1) “E. amplissimo folio lusitanicum. Ech. lusitanicum Jolio
cubitali T. hb. Isnardi.”
(2) “ E. amplissimo folio lusitanicum” ex herb. D. Charles.”
Both specimems are undoubtedly the lusitanicum of herb. Linn.
“no. 48 Leh. undulatum Pourr. Gallice, envoyé par M. Pourret
1802," labelled by Gay “ forte E. lusitanicum L.,” which it is,
being identieal with no. 6605.
two pieces :
` (1) * E. rubrum” is rightly corrected by Gay to E. vulgare.
(2) “ E. ereticum angustifolium rubrum ex hb. D. Charles? is
vulgare corollis voseis.
“Echium Insularum Stechadum” is rightly labelled by Gay
“E. plantagineum,” for it is not the plant so-called in
Tournefort’s herbarium no. 591.
two sheets, ex herb. Isnard and ex herb. Charles, both named
“ Echium creticum latifolium rubrum," but both are E. planta-
gineum and have been so labelled by Gay.
“ Echium orientale” is plantagineum as labelled by Gay. This
must be the specimen alluded to by Poiret, Dict. Encye. viii.
p. 673, where he says that he has seen Tournefort’s type of
E. orientale in hb. Jussieu.
“ Echium ex hisp." do. do.
* Echium ex hort. R. Paris ” do. do.
sine nomine, do. do.
* Echium bonariense ” do. do.
sine nomine, a scrap do. do.
two sheets :
(1) “E. australe Lam. ex hort. reg. Paris, Desf." marked
“bene” by Gay.
(2) sine nomine aut loco, Both are precisely the creticum of
herb. Linn.
2: K 2
382 MR. C. C. LACAITA : A REVISION OF
no. 6620 sine nomine “ex h. r. Paris," labelled by Gay “E. grandiflorum
Desf.?" This also is identical with Z. ereticum of hb. Linn.
4 6621 “ FE. foliis angustis et villosis T. R. H. 136 ex hb. Isnardi," no note
by Gay. There are four sheets of a much branched plant. This
synonym is referred by Jussieu in Barr. p. 16 to Anchusa angustis
villosis fol. hisp. Barr. Ie. 577, which is Echium hispanicum Asso,
Mant. Stirp. Arag. p. 162 (1781). As I had not seen Pau's
specimens of Asso’s Echium at the time T was in Paris, I could
not recognise it in these sheets, and dare not express an opinion
from memory.
, 6622 two sheets :
ED A hispanicum verrucosum annuum anguiiifoliuh ex hb. P.
et hb. Charles” is identical with no. 6621.
(2) * E. rosmarinifolium (? Roris-marini folio) T. EH. 120 6x
hb. Charles," may also be hispanicum, Asso, or the Spanish plant
commonly called pustulatum. I cannot say, for the reason
mentioned above. Both (1) and (2) are labelled by Gay
* E. tuberculatum Hfg. & Lk. ? " which they are not.
„ 6630 six sheets :
(1) “E. ereticum angustifolium rubrum ex hb. Isnardi, ^
labelled by Gay “ E. creticum a Poir. ex hb. Desf.,” is identical
with hb. Tournefort no. 589. It is Æ. angustifolium, Mill.
— E. elegans, Lehm. I have examined the specimens in
Desfontaines's herbarium at Florence referred to by Gay.
There are three, on which Gay has noted “Ex hisce tribus
speciminibus suum Æ. creticum elaboravit Poiret in Dict. viii.
p. 670, quorum duo Syriaca, meum ined. Æ. Tournefortii, tertium
JEgyptiaeum E. prostratum. Delil. Æg. sistit." Nos. 1 and 2
came from Labillarditre, who collected in Syria, but they are
sine loco. They are E. angustifolium, Mill.— E. elegans, Lehm.
No. 3, sent from Egypt by Delile himself, is typical Æ. sericeum,
Vahl, which is Delile's prostratum.
(2) sine loco ; my MS. abbreviated note is illegible.
(3) “misit D. Thunberg e Tripoli" is apparently also
angustifolium.
(4) sine loco, labelled by Gay “Je ne puis distinguer cette
plante du E. plantayineum.” This is certainly not angustifolium,
but is either plantagineum or maritimum. I did not examine it
minutely.
(5) and (6) are both angustifolium= elegans.
I cannot now explain the lacune in the numbers between 6607 and 6611
and between 6622 and 6630, I may have accidentally missed some sheets,
_
.
S
p
14.
hg aes M COMPONE T, TUN
SOME CRITICAL SPECIES OF ECHIUM. 383
III. Hers. Lamarck.
* Echium vulgare” sine loco. It is.
* Echium rubro flore montis aurei," presumably Mont D'Or. It is vulgare
fl. roseo. ^
* Echium plantagineum Vahl” is E. plantagineum, L., a specimen with
fine large radical leaves.
* Echium elongatum Lam. an Echium majus et asperius fl. albo? Ses
fleurs sont d'un blanc un.peu incarnat., les corolles sont courtes" An
unknown later hand has added * Echium glomeratum Poir.,” which it
is not, being precisely Æ. italicum, L. (as already noted by de Coiney)
=altissimum, Jacq.
* Echium fruticosum L.” is Lobostemon fruticosus, Buek.
. “ Echium argenteum Berg. 40 et Lam. Ill., Pluk. Tab. 341. f. 8” is
Lobostemon argenteus, Buek, the plant figured by Jacquin Hort.
Scheenb. i. p. 34, as E. fruticosum, but not the plant named
E. argenteum in the Linnean herbarium (see my paper on the
Echia of Linnæus, infra, p. 396).
* Echium capitatum” is Lobostemon capitatus, Buek.
“ Echium capitatum var. 8? is Lobostemon spherocephalus, Buek.
* Echium spicatum” is Lobostemon spicatus, Buek.
. “ Echium strictum Lam. Ill.” is E. strictum, L. fil.
. “ Echium falcatum Lam. Ill." with a label in later writing ** Lophost.
glaber Buek," which apart from the mis-spelling, is probably right.
2. * Echium candicans L. suppl.” is that species.
3. “ Echium italicum . . . . (illegible) Pernegul. Echium asperrimum Lam.
Ill." is precisely E. pyramidale, Lap., with whose type-specimens I
have compared it, and therefore is Æ. pyrenaicum, Desf.
* Echium angustifolium Lam. Ill. E. hispanicum Tobarra = Barr, Ic.
1011." This is the type of Lamarck’s species, which Poiret wrongly
identified with Æ. angustifolium, Mill. Salzmann, J. Gay, and Cosson
misinterpreted this name, applying it to a very different species,
E. Coineyanum, mihi. But Rouy re-discovered the true plant in
1879 not far from Tobarra, and later collectors have distributed it
as E. humile, Desf. But de Coincy, who discusses it fully in Morot’s
Journ. Bot. xiv. p. 106, and xvi. p. 215, points out that it is not true
humile. The name angustifolium suits admirably, but is unavailable
owing to the priority of Miller's angustifolium. It must therefore be
called E. pyenanthum, Pomel, Nouv. Mat. Fl. Atl. fasc. 2, p. 40 (but
p. 296 in some copies, where the paging is consecutive with the
author's earlier publication), where the name is substituted for
E. densiflorum, Pomel, owing to the existence of the earlier Æ. densi-
florum, DC. Barr. Ie. 1011, Lycopsis angustifolia minor hispanica,
284 MR. C. €. LACAYITA : A REVISION -OF
certainly represents the plant of Lamarck, who, however, was wrong
in quoting for it ££. hispanicum, for which see note on no. 6621 of
Herb. Jussieu,
15. ** Echium australe Lam. illustr.,? with a later label in an unknown hand
“ Echium lusitanicum ?,” a ridiculous suggestion that may be dis-
missed. This, again, is an important type. It is absolutely identical
with the specimen of E. creticum in herb. Linn. No doubt a garden
plant, originally derived from E. grandiflorum, Dest., but not from
the narrow-leaved, upright form of that species found in Dept. Var.
The flowers are badly dried and look smaller than they really are.
There are no ripe nueules, so that de Coincy’s description of them in
Morot, Journ. Bot. xiv. p. 327, where he mixes up Lamarck’s australe
with Æ. Coincyanum, mihi, cannot have been derived from Lamarck’s
type.
16. * Echium creticum Lam. ill," labelled by a later hand * Hehium cre-
ticum?” Three examples: one “ex D. Sonnerat," the others sine
loco. "They are all Æ. plantagineum, L., to which Lamarck’s creticum
has long ago been referred.
17. * Petit rameau détaché d'une sommité fleurie d'un. Echium frutiqueux
des Canaries," labelled by a later hand ** chium aculeatum Poiret,”
which it may well be.
ILI.
THE ZCHIA OF SIBTHORPS HERBARIUM.
The specimens in Sibthorp's herbarium, now at Oxford, were originally
tied up in bundles corresponding to Sibthorp’s different journeys and the
districts from which they came. But the individual specimens are without
any indication of locality or determination by Sibthorp himself, the only
writing on the sheets being in Smith's hand, except in the caseof plants from
the island of Zante, which were not collected by Sibthorp, but purchased
from a druggist of the island, who has written on some of them the local
Greek name.
Smith tells us, in Rees’s Cyclopedia sub voce Sibthorp, that the plan of the
* Prodromus was drawn out by Dr. Sibthorp, but nothing of the ‘Flora Græca’
except the figures was prepared, nor any botanical descriptions. ‘* The final
determination of the species," he says, ** the distinction of such as were new,
and all eritieal remarks have fallen to the lot of the editor." He has not
always been very happy in these determinations and remarks. In the genus
Echium alone he has fallen into two grave errors. But we must remember
that he had not the copious material collected by later travellers, which
makes it so easy for us to criticise his work.
SOME CRITICAL SPECIES OF ECHIUM. 385
Smith has not attempted to determine all the specimens. Only seven
species of Echium are enumerated in Fl. Gr. Prodr. i. pp. 124-126 (1806),
and of the fourteen examples of that genus in the herbarium only seven have
been named by Smith. Nine of the fourteen bear a printed label with
“J, Sibthorp, M.D.,” indicating that they were actually collected by him.
Three others are marked, apparently in a clerk’s hand, * Herb. Sibthorp.
South of Europe. Qy.,” and the remaining two are from the Zante collec-
tion mentioned above. From a copy of the * Prodromus" in the possession of
the Linnean Society, it appears that at one time authentic Sibthorpian speci-
mens of his Æ. hispidum and E. creticum, presented by Dr. Daubeny, existed
in the Society’s general South European herbarium. Unfortunately, these
were sold by auction on November 10th, 1863, being comprised in lot 82
with Welwitsch’s Portuguese collection and Dr. Prior’s east Mediterranean
plants. The lot only fetched 34 shillings, but the purchaser’s name is
not known. The loss is peculiarly unlucky, as those two species are the
very ones about which Smith went wrong. He appears to have taken
the names of the three new species of the ‘ Flora Graeca’ (X. pustulatum,
E. hispidum, and E. diffusum) from some list of Bauer's figures which he
found among Sibthorp’s papers, but which does not seem to have been pre-
served. Sundry rough lists exist among Sibthorp’s MSS., but they contain
none of the above names. A list “intended to form the outline of the * Flora
Greeea’” only includes three Lehia, viz., no. 164, Æ. italicum, no. 165,
E. vulgare, and no. 166, Æ. creticum, without localities; and a list of
Thracian plants mentions “ K. violaceum in campis Thraciv” and ** E. cre-
ticum in campis circa Byzantium.” There is nothing in the lists to connect
these names with any particular specimens.
It seems that Bauer did not draw—or, at any rate, did not complete—the
figures of the * Flora Groca from live plants. His originals exist at Oxford.
They have been most faithfully reproduced by Sowerby in the published
plates. There are also many sheets of Bauer’s pencil-sketches of the dif-
ferent parts of the organs of the plants, perhaps done in the field, but at any
rate from freshly-gathered specimens. Portions of several species belonging
to different genera are found on the same sheet. These sketches are marked
all over with numbers indicating the precise tints to be afterwards applied
in the coloured pictures. The tints must have been very numerous, as the
numbers run to upwards of 200.
I have to thank Mr. Druce for much of the above information and for the
facilities: afforded for the examination of the specimens enumerated and
discussed below.
(1.) ECHIUM VULGARE, L., labelled ** Herb. Sibth. South of Europe. Qy.”
and not named by Smith. It is a large specimen with the upper half of the
stem cut off, but I have little doubt that it is Æ. vulgare, which is quoted in
386 ' MR. C. C. LACAITA : A REVISION OF
Prodr. i. p. 125 for Byzantium and Laconia. The latter station seems to me
very doubtful for typical vulgare, whieh is not found on the shores of the
Mediterranean.
(1I.) Ecnruu rusrULATUM, labelled by Smith “ Heh, pustulatum Sibth. list
of figures.” This is certainly the plant represented in Fl. Gr. tab. 180,
although in a more advanced state than the figure. In Prodr., l c., and FI.
Gr. ii. p. 68, Sibthorp is said to have found this species “in Sicilia tantum.
Nevertheless, it agrees perfectly with the example collected by him in the
kingdom of Naples, now in Herb. Banks, but wrongly referred by Smith to
E. hispidum, whose expression “in agro Neapolitano” must, I think, be
understood to mean “in the kingdom of Naples," not “in the vicinity of
Naples,” where that precise form is not found, though it occurs in southern
Calabria as well as in Sicily. Among some rough notes of Smith's at Oxford
I have come across “ Mehium hispidum Sibth. from Naples; stem very bristly ;
flowers smallish, blue," which is sufficient to exclude the real hispidum, It is
possible that Sibthorp himself may at some time have confused pustulatum
with the very different Greek plant figured as hispidum. Modern specimens,
which entirely agree with both the above Sibthorp specimens, are :—
1. Todaro, Fl. Sic. no. 931, from Messina, a locality also quoted by
Gussone, Fl. Sic. Syn. i. p. 232, for E. pustulatum.
2. Thomas in Hb. Gay at Kew, from Calabria, anno 1816.
3. Arcangeli in Hb. Kew, from Cape Spartivento in Calabria.
4. Tenore in Hb. Kew, from southern Italy, without precise locality.
I am unable to see any real distinetion between the above and two well-
known exsiecata from Nicolosi on the slopes of Etna, viz. :—
5. Strobl, Fl. Ztn., 24. vi. 1872.
6. Lojacono, Pl. It. Select. no. 72.
Nevertheless, these were referred to by their collectors to a form that
Gussone, l.c., and Lojacono, Fl. Sie. iv. 2, p. 76, supposed to be distin-
guishable from pustulatum, and wrongly referred to Æ. tuberculatum, Hoffmg.
et Link, quoting Nicolosi as a locality. "True Æ. tuberculatum, Hotfmp. et
Link, as interpreted by de Coincy in Morot, Journ. Bot. xiv. p. 303 (1900),
and by Coutinho, Fl. de Portugal, p. 500 (1913), and represented by the
following specimens, does not exist in any part of Italy * :—
* Alph. de Candolle remarks in Prodr. x. p. 19, under Æ. tuberculatum, “omnia ex verbis
cl. auct. Nemo specimina authentica vidit et omnes de hac specie disposuerunt. Pater
diversas plantas ex Italia, Gallia et Hispania sub hoc nomine in herb. consociavit, sed
communio dissocianda.” This judicious observation has been overlooked by many French
and Italian authors, e. g., by Loret, Glanes d'un Botaniste, in Bull. Soc. Bot. Fr. vi. p. 406
(1859), who, though he properly declines to identify the so-called pustulatum of southern
France with Sibthorp's pustulatum, unfortunately refers it to tuberculatum, Hoffmg. & Link.
SOME CRITICAL SPECIES OF ECHIUM. 387
7. Schultz, Herb. Norm. no. 1429; S. José prope Conimbricam,
Maio 1882, leg. Moller (in Herb. Bailey at Manchester).
8. Fl. Lus. Exs. Hort. Bot. Conimbr. no. 110; Coimbra, Quinta das
Maias, Aprili 1886, leg. Moller, but named * pustulatum " (Herb.
Mus. Brit.).
9. J. Daveau, Herb. Lusit. 1878; Penna de Pau, environs de
Lisbonne (Herb. Kew).
10. Burchell, Cat. in Lusit. lect. no. 565, as E. vulgare (Herb. Kew).
The common Echium that replaces Æ. vulgare in Istria and the greater
part of continental Italy is by no means identical with Sibthorp’s type
of pustulatum, and is quite different from the Portuguese tuberculatum,
though usually known by one or other of these names. It seems inter-
mediate between vulgare and pustulatum, and may well be known as
E. vulgare var. grandiflorum, Bert., under which name it has been admirably
described by Bertoloni, Fl. It. ii. pp. 348, 350. It is the pustulatum of
Koch and of many Italian authors, but the tuberculatum of Gussone’s
Neapolitan herbarium. J. Gay noticed the difference between true pustu-
latum of Sibthorp and this form, which he proposed to call E. Tenoreanum.
His MS. notes attached to the Calabrian specimen of pustulatum in his
herbarium (no. 2 above) are interesting. “Æ. pustulatum, Fl. Gr. Prodr.
planta sicula. Fl. G. t. 180 optima... . Sibthorpii icon. in Fl. Graec. meum
calabrieum specimen optime refert; rami floriferi longiuseuli et omnes
partes hispidissime ; sed tota planta nimium viridis, quod forte uni pictori
tribuendum. Unum quod Parisis vidi sieulum E. pustulati specimen,
in hb. Fontanesii exstat, in monte Ætna a D. Schouw lectum. Hnic in
sesquipedalem longitudinem porrecto et simplicissimo spicule sunt 11,
brevissime vix unciales, extrorsum arcuate, unde habitu certe differt.
Hoc vero non nisi ab state juniore pendere videtur. Convenit vero cum
His treatment of the subject loses much of its value owing to his conception of pustulatum
being based on a Bourgeau specimen from southern Spain, which is not Sibthorp’s species,
and has been referred by de Coincy to E. pycnanthum (angustifolium, Lam., non Mill.),
but his remarks on the French plant are interesting: “ Quoique PE. pustulatum de Toulouse
et de la Flore de France ait un facies un peu différent de celui de l'E. vulgare ordinaire, et
qui suffit le plus souvent pour empécher de le confondre avec lui, on óprouve néanmoius,
lorsqu'on étudie chaque organe, une sorte d'impossibilité d'y reconnaitre des caractères
stables et vraiment spécifiques. Celui notamment qui est relatif à la forme de la panicule
est tellement variable, et par suite si peu distinetif, qu'il me parait sans valeur." He does
not seem to have been aware that the nucules of these southern plants are not those
of typical vulgare. Rouy, on the other hand, Fl. Fr. x. p. 317, being well aware that the
French plant is not the true Portuguese tuberculatum, refers the former back to pustulatum
treated as a subspecies of vulgare: but that he had no clear idea of pustulatum appears from
his quoting both Bourgeau, no. 1314, and Todaro, no. 931, as representing it, though these
two exsiccata belong to different species.
388 MR. C. ©. LACAITA: A REVISION OF
méo. Echium pustulatum Ten.* in herb. Mus. Paris (in arvis siccis circa
Neapolim a Tenorio lectum) alia planta est, /ccAio vulgari longe atftinior,
foliorum pube eum hoe vulgari conveniens, diversa satis in bracteis, foliorum
margine et eosta dorsali rigidioribus lucidis, strigulis brevissimis rigidis
adpressis bractearum dorsum occupantibus, etc."
This intermediate var. grandiflorum is represented by the following,
among many other, examples :— Ò
11. Portici al Granatello (near Naples), from Tenore as Æ. vulgare in
Hb. Kew.
12. Ibidem, from Gussone as Æ. tuberculatum in Hb. Kew.
13. M. Gargano, from Tenore as E. vulgare in Hb. Kew.
14. “ In collibus aridis totius Dalmatie," Unio It. 1829 as X. pustu-
latum in Hb. Kew.
15. Veglia (Dalmatia), leg. Bauer as Æ. pustulatum in Hb. Kew.
16. Zara (Dalmatia), leg. A. Braun as £. pustulatum in Hb. Kew.
The last three obviously represent Koch's conception of E. pustulatum.
Like typical vulgare this form occasionally occurs with pink corollas, when
it has been mistaken by Italian botanists—e. y., Bertoloni, Fl. It. ii. p. 347—
for Æ. angustifolium, Mill. (hispidum Sibth.), which is not Italian t.
Although I have looked through—somewhat hurriedly—all the French
specimens of Echium in the Paris Museum and those of M. Rouyt, I have
found none among them that are undoubtedly identical with Sibthorp’s
pustulatum. Most so-named Spanish examples are referable either to
E. Coincyanum, mihi — E. creticum, Willk. et auctt. hisp., non Linn., or to
E. hispanicum, Asso§ = Anchusa angustis villosis foliis hispanica, Barr. Ic. 577,
but I cannot speak positively as to others.
The systematic value of Æ. pustulatum as represented by Sibthorp’s type,
and its relation to the other forms referred to, remains doubtful. What
is clear is that the vulgare of northern and central Europe is replaced in the
Mediterranean, not by one equivalent form, but by sundry * little species,”
differing from each other as much as from typical vulgare, and that these
have hitherto been very superficially studied. Herbarium specimens are
* Tenore usually sent out this form from the vicinity of Naples as vulgare, sometimes as
tuberculatum, occasionally, as in the specimen here referred to, as pustulatum. True
pustulatum he sometimes sent out under that name but sometimes as tuberculatum, so that
his specimens, for which a precise locality is rarely indicated, are valueless for nomenclature,
I have seen Schouw's Etna plant in hb. Desfontaines, now at Florence; it is like no. 5
above, and undistinguishable from Sibthorp's specimen.
T See my observations on Æ. Sibthorpii in Nuov. Giorn. Bot. It. xxv. p. 136 (1918).
} Rouy’s fine herbarium is now the property of Prince Roland Bonaparte, who most
generously welcomes those who wish to study his magnificent collections.
$ Mant. Stirp. Indig. Arag. p. 162 (1781). A copy of this rare work is in the library
at Kew. Specimens distributed by Pau are in the Bailey herbarium at the Manchester
Museum.
a RS ii... bau
SOME ORITICAL SPECIES OF ECHIUM. 389
usually very unsatisfactory in this genus. They should be collected with
the complete radical leaves before flowering, then in flower, and again with
fully developed fruiting spikes. Both true pustulatum and vulgare var.
grandiflorum differ from our English vulgare in a more branching habit, which
however is not developed in weaklings ; in corollas normally larger, of a
different blue, which shows purple streaks or a slight purple tinge ; in
stamens not so far exserted * ; and particularly in the nutlets, which, instead
of being merely rugose with inconspicuous, if any, tubercles, are remarkably
papilloso-tuberculate. This last character has been rather cavalierly treated
by de Coincy, op. cit. p. 303, but if it can be established as constant it would
be decisive in favour of a specific separation from vulgare of the forms in
which it occurs. But it requires further observation on a sufficient number
of fresh specimens in different regions.
There is one other point to notice. De Coincy, at pp. 303 and 323,
distinguishes vulgare and pustulatum, treated as a variety of vulgare, as
having glabrous filaments, whilst in tuberculatum, Hoffmg. et Link, * at least
one of the three posterior filaments is hairy, and usually all three." Though
hairy filaments are very important in some other species, notably in grandi-
florum, Dest., they afford so feeble a character in tuberculatum (which is quite
distinguishable on other grounds) that it has been deliberately ignored by
Prof. Coutinho, The hairs are very few and very weak, and extremely difficult
to see in any herbarium specimen—indeed, I very much doubt the constancy
of their presence. Yet they are not always entirely absent, as they are in
all forms of vulgare and in all specimens of pustulatum that I have examined,
I have refrained from dissecting the few remaining corollas of Sibthorp’s
type, but as far as can be seen they are glabrous, and so they are in Todaro’s
identical plant.’ Nevertheless, in the detail of tab. 180 at least two of the
tilaments are shown as hairy, and I have verified that they were so drawn by
Bauer in his original sketch. The kind of hairs figured, sparse and slender,
suggests that Bauer must have really seen them. They are not the kind of
pubescence that an artist would be likely to introduce apart from actual
observation, I am unable to explain this matter further.
(IIL) ECHIUM ANGUSTIFOLIUM, Mill., labelled by Smith * Echium hispidum
list of figs. Naples. Sibth. in H. Banks." This is precisely E. hispidum
of Fl. Gr. tab. 181. The synonymy is Æ. angustifolium, Mill. Gard. Dict.
(1768), non Lam. — E. hispidum, Prodr. Fl. Gr. p. 125 (1806) = Æ. elegans,
Lehm. Asperif. p. 459 (1818) — Æ. Sibthorpii, R. et S. Syst. iv. p. 26 (1819)
= E. sericeum var. hispidum, Boiss. Or. iv. p. 207 (1879).
* The phrases “included” and “exserted” have been used ambiguously of the stamens
of the irregular corollas of this genus. By “included,” I understand shorter than the lower
lip of the corolla; “exserted” should, I think, mean exceeding the upper lip. Stamens
that exceed the lower, but fall short of the upper, lip may fairly be said to equal the corolla
and be called sub-exserted.
390 MR. C, €, LACAITA : A REVISION OF
Lehmann created the name elegans and Roem. & Sch. that of Sibthorpii
(before he had seen Lehmann’s publication of the preceding year) for Sibthorp’s
species, owing to the existence of Thunberg’s hispidum of 1794 (Prodr. Pl.
Cap. p. 33), the much earlier hispidum, Burm. fil. Fl. Cap. Prodr. p. 5
(1768), being reduced at the same time by Lehmann to a synonym of
E. capitatum, L. Both these are Cape Lobostemons, but in any case the
priority belongs to Miller's name, the identity of which with Sibthorp’s
E. hispidum I have maintained in my notes on the Echia of Miller's
* Gardener's Dictionary.’
This species, which is the E. creticum II. of. Clusius and the E. ercticum
angustifolium rubrum of C. Bauhin and of Tournefort, is plentiful in Crete
and in many parts of Greece. By Linnmus it has been mixed up with a
totally different species under the confused name of E. ereticum. Smith
was misled into supposing that Sibthorp had obtained the plant “in agro
Neapolitano” by his false determination of the specimen in Herb. Banks,
now at the British Museum, which, as stated above, is not hispidum but
pustulatum. It is possible that Sibthorp himself at some period may have
confused this South Italian pustulatum with the Greek species figured as
E. hispidum. A similar confusion had been made by Miller (see my
note on his E. angustifolium).
(IV.) ECHIUM PARVIFLORUM, Moench, labelled by Smith * Echium ereticum.?
(V.) ECHIUM PARVIFLORUM, Moench, labelled by Smith in ink * Echium
ereticum” and in pencil * Sibth. at vix H [erb].L [inn]. perhaps from Naples."
Both IV. and V. are parviflorum=calycinum, Viv., no. V. being the usual
seaside plant, and no. IV. the stronger form that develops in richer soil — var.
erectum, DC, Smith's pencil note shows that the error of taking this plant
for E. creticum, L., was not due to him but to Sibthorp. I cannot account
for the false colour attributed to the corollas in Prodr. p. 126, where Smith
calls them “rubro-violacei,” and so they are represented in tab. 183,
whereas in most cases they are pale blue. The false identification in
De Candolle's * Prodromus; x. p- 22, of creticum, Fl. Gr., with creticum, L.,
is explained by the observation in a footnote “ plantam Linnæi et ic. Fl. Gr.
non vidit pater nec ego."
(VL) ECHIUM PLANTAGINEUM, L., labelled by Smith “ Echium plantagineum
H. L.? It is quite typical; the corollas have dried partly blue, partly
purple.
(VIL) ECHIUM PLANTAGINEUM, L., from Zante, not named by Smith, but
“no. 45 BovóóyAeecov." In this the corollas have dried blue.
(VIII.) ECHIUM PLANTAGINEUM, L., also from Zante, not named by Smith,
but labelled * 234 Botydwooor.” In this case the corollas have dried pale
SOME CRITICAL SPECIES OF ECHIUM. 391
pinkish purple as shown in Fl. Gr. tab. 179. E. plantagineum frequently
tends to this colour in the Ionian islands. In the Atlantic islands, io judge
by the numerous examples in Herb, Kew, it seems usually to be a rich
purplish blue.
(IX.) ECHIUM PLANTAGINEUM, L. Three small pieces on one sheet, on
which is written in a clerk’s hand * Herb. Sibthorp. South of Europe Qy?”
These have not been named by Smith.
(X.) EcHIUM ITALICUM, L., labelled by Smith * E. italicum H. L? A
small piece in fruit, very hirsute.
(XL) ECHIUM ITALICUM, L., not named by Smith. On the sheet is written
“ Herb. Sibthorp. South of Europe Qy?” Both X. and XI. are true
E. italicum, not E. pyrenaicum, which does not seem to grow in Greece.
(XIL) ECHIUM ARENARIUM, Guss., not named by Smith. It shows the
typical small dark blue corollas with style as well as stamens included.
This species is plentifully represented in modern herbaria from Crete and
from Attica. Halácsy, Consp. Fl. Gr. ii. p. 340, wrongly identifies
E. diffusum, Sibth. et Sm., with E. arenarium.
(XIII.) ECHIUM DIFFUSUM, Sibth. et Sm., labelled by Smith * Echium
diffusum Fl. Gr. tab. 182.” The specimen is in fruit.
(XIV.) ECHIUM DIFFUSUM, Sibth. et Sm., not named by Smith, but with
the printed label “ J. Sibthorp M.D.,” showing it to have been collected by
Sibthorp. It is identical with no. XIII. These two specimens differ
toto cælo from E. arenarium in their reddish corollas with conspicuously
exserted style and stamens as long as the upper lobes. The fruiting calyces
also differ from those of arenarium. It is true that in Fl. Gr. ii. p. 69,
E. diffusum is described “staminibus corolla brevioribus? and tab. 182 shows
them about as long as the corolla. But in these specimens they are longer
than they are drawn by Bauer and certainly not shorter than the corolla.
The colouring of that figure and the words in the text, “flores punicei,
precedenti (sc. Æ. hispido) similes," exclude Æ. arenarium. In fact,
E. diffusum is precisely Æ. sericeum, Haláesy, loc. cit. p. 339, non Vahl,
quoted by that author for the plant from the sandy shore at Canea, more
correctly referred by DC., Prodr. x. p. 23, to E. diffusum. Identical
specimens from the isle of Naxos may be seen in Herb. Gay at Kew, ticketed
“ E. diffusum Smith. In Naxo insula ann. 1829 ; Despréaux sine nomine
in herb. Deless. — Frustula haecce speciminis multicaulis dedit Guillemin.
Schedule Despréauxiane nomen Æ. arenarium Guss. propria manu addidit
Gussone, sed falso, ui mihi videtur. Species valde affinis E, setoso, Vahlii,
392 MR. C. C. LACAITA : A REVISION OF
an satis diversa? — Diversissima a setoso Delile seu arenario Guss.,” and
again in Heldreich’s exsiecata from that island under the name of
Hi. arenarium B. Sieberi. This var. Sieberi was so named by De Candolle,
Prodr. x. p. 21, and is based on an Egyptian specimen collected at the
Pyramids by Sieber, which I have not seen. The Egyptian exsiccata
most like Sibthorp’s plant and those from Naxos are a prostrate form of
KE... sericeum, Vahl, from Ramleh, with subexsert stamens, which differ from
Sibthorp’s difusum in possessing the adpressed pubescence of typical
sericeum. J
Boissier, Fl. Or. iv. p. 207, treats E. diffusum as a variety of sericeum,
quoting it from Cos and Cyprus as well as from Crete and Naxos. This is
right from his point of view, as he also treats Æ. angustifolium, Mill.,
as var. elegans of F. sericeum. But if angustifolium and sericeum are kept up
as separate species and d//usum submerged, it is a problem to which of the
former diffusum should be referred as a prostrate variety. The chief
difference between those two species is in the indumentum—adpressed and
rather silky in sericeum, but patent and hispid in angustifolium. That of
difjusum seems intermediate, but to my eyes nearer to that of angustifolium,
where it would also be preferable to place diffusum on geographical
grounds. It seems best to retain diffusum as a species until much more
copious material is available.
The synonymy therefore is Æ. diffusum, Sibth. et Sm., Prodr. i. p. 125
= E. sericeum, Halácsy non Vahl =Æ. sericeum, Vahl, var. diffusum, Boiss.
= E. arenarium var. Sieberi, Heldr., exsice. non DC.
IV.
THE LINNEAN SPECIES OF ECHIUM.
The chia of Linnzeus and his son comprise six South African species now
referred to the genus Lobostemon; one from Asia Minor transferred by
Boissier to his monotypic genus AMegacaryon; three shrubby species of
Echium from the Atlantic islands, and nine European herbaceous species.
It is only with the last that I propose to deal fully, as the Atlantic species
and the Megacaryon offer no difficulty, and I am not sufficiently acquainted
with the South African flora to say much about the Lobostemons, though
two of these, nos. 1 and 6, raise problems of interest.
Not all these nineteen species are represented in the Herbarium, which,
however, contains three not mentioned in the Linnean writings, viz.:
Lobostemon montanus, Buek, chium rubrum, Jacq.. and a small Echium from
Spain which I cannot determine with any confidenee.
I shall first enumerate the species, then the specimens, then offer some
general remarks, and finally discuss the European species. As the sheets in
the herbarium are not numbered, they are referred to by letters of the
TENERE S
SOME CRITICAL SPECIES OF ECHIUM. 393
alphabet, as the use of numbers might lead to confusion should the sheets
themselves be numbered at some future time :—
(L) LonosrEMON"*.
l. Echium argenteum, L. Mant., p. 202, non Lobostemon argenteus,
Buek in Linnæa, x. p. 133. This is specimen O in the herbarium,
labelled Æ. argenteum. It has been examined by Mr. N. E. Brown,
who confirms that it is not the L. argenteus of Buek, but a
species not otherwise known, which he has described and named
Lobostemon magnisepalum in a paper read before the Linnean
Society on 6 February, 1919, which will appear later in this
Journal. The herbarium contains no specimen corresponding
to ££. argenteus, Buek.
2. Echium capitatum, L. Mant., p. 42— Lobostemon capitatus, Buek,
l.c. p. 143. Specimen Q.
3. Echium fruticosum, L. Sp. Pl. (1753) p. 139 = Lobostemon
fruticosus, Buek, l. c. p. 134. Specimens RID,
. Echium levigatum, L. Sp. Pl. (1762) p. 199=Lobostemon
levigatus, Buek, l. c. p. 139. Specimens U, V.
Echium spicatus, L. fil., Suppl. p. 132— Lobostemon spicatus, Buek,
l.c. p. 145. Specimen W.
6. Echium, unnamed = Lobostemon montanus, Buek, l.c. p. 152, on
the authority of Mr. N. E. Brown. Specimen P.
nS
Or
(I1.) MEGACARYON.
The herbarium contains no specimen of Echium orientale, Sp. Pl. p. 139,
and it is practically certain that Linnæus never saw the plant in any form.
Its identity depends entirely on the synonym quoted from Tournefort,
« Echium orientale, verbasci folio, flore maximo campanulato,” and that
author's account of it in his * Voyage au Levant,’ ii. p. 248 (1717), or iii.
p. 83, in the English translation of 1741. It is first mentioned by Linnzeus
in Hort. Cliff. p. 43, “ Echium caule ramoso, foliis caulinis ovatis, floribus
solitarüs ex alis. Tourn. Cor. 6; itin. 3, p. 94. Crescit in Oriente." The
diagnosis in Sp. Pl. is a mere transcript of this with the word “lateralibus ”?
substituted for “ex alis." Tournefort found his plant at Grezi on the road
from Trebizond toward Armenia. His description and the locality make it
certain that it is the species for which Boissier created his monotypic genus
* I have not included in the Lobostemon list an Echium glabrum, Linn., Epist. ed. van
Hall, p. 27 (1880), where that name is given in a letter to J. Burman, of Oct. 4th, 1758,
for a plant received from him. The diagnosis runs, “Æ. glabrum, caule levi, foliis lanceo-
latis nudis, margine carina apiceque scabris,” with synonyms cf Pluk., Old., and Herm. On
the strength of these synonyms, Richter, Cod. Linn. p. 156, identifies this plant with
E. levigatum, published two years after this letter was written. E. glabrum, Vahl, and
E. glabrum, Thunb., are of course much earlier than this Æ. glabrum, Linn., of 1830,
394 MR. C. C. LACAITA : A REVISION OF
Megacaryon in Pl. Or. Nov. Decas i. p. 7 (Feb. 1875), based on Bourgeau's
specimen from Macka. Boissier does not there allude to Linnsus or
Tournefort, but in Fl. Or. iv., published later in the same year, he altered
the name from Megacaryon armenum to M. orientale, quoting | chium
orientale, L., and Tournefort’s synonym.
The synonymy therefore is Megacaryon orientale, Boiss. Or. iv. p. 204
(1875) — M. armenum, Boiss. Pl. Or. Nov. Dec. i. p. 7 (1875, Feb.) = Onosma
megalospermum, Boiss. in Bourg. exsicc. = Echium orientale, L. Sp. Pl. p. 140
(1753) =, grandiflorum, Salisb. Prodr. p. 115 (1796), non Desf.
Boissier says “ex unica corolla exsiecata in specimine fructifero fausto
caso superstite florem descripsi," but since then fine specimens in flower
and fruit have been collected by Sintenis, It. Orient. 1892, nos. 4159 and
4859, from two places in Paphlagonia. These may be seen in Herb. Kew
and in Mus. Brit. Poiret, Diet. Encyc. viii. p. 673, says that he has seen
Tournefort’s type in Herb. Juss. He no doubt was alluding to specimen
no. 6613, which, although labelled orientale, is nothing but X. plantagineum.
There is no example of Tournefort’s species in his own herbarium. That
Linneus never saw a specimen is certain, not only from his giving no
diagnosis of his own and merely quoting Tournefort, but also from an MS.
note in his interleaved copy of Sp. Fl. ed. 2, where he quotes for Z. orientale
Tournefort’s figure, and also Trew, Pl. Rar. tab. 1, published in 1768.
Trew's figure, which seems to represent Æ. grandiflorum, Desf., is so utterly
unlike Megacaryon orientale, that no one who had ever seen an example of
that striking plant could have connected this figure with it.
(IIL) Ecutum. (A) Atlantic shrubby species.
l. Echium candicans, L. fil. Suppl. p. 131. Specimen A, which is
actually the type referred to by Linn. fil.
2. Echium giganteum, L. fil. ibidem. No specimen.
3. Echium strictum, L. fil. ibidem. No specimen.
(IV.) Ecntum. (B) European herbaceous species.
1. Echium creticum, L. Sp. Pl. (1753) p. 139. Specimen N.
2. Echium italicum, L. ibidem. Specimen H.
3. Echium lusitanicum, L. ibidem. Specimen M.
4. Echium plantagineum, L. Mant. p. 202. Specimen C.
5. Echium pyrenaicum, Desf. (L. Mant. p. 334). Specimen I.
6. Echium rubrum, Jacq. Not mentioned anywhere by Linnzeus, but
represented in the herbarium by the three unnamed specimens
J, K, L.
T. Echium violaceum, L. Mant. p.42. Specimen F.
8. Kehium vulgare, L. Sp. Pl. (1753) p. 139. Specimen B,
9. Echium —— ? Specimen G.
Teo UIROS ES a
SOME CRITICAL SPECIES OF ECHIUM. 395
The Linnean herbarium contains two covers marked Ecnmruw. In these
there are 23 sheets, of which only thirteen bear specific names in the hand-
writing of Linneus; two more are in that of hisson ; but eight are unnamed.
In two instances there are two sheets pinned together, although bearing quite
different plants. As there is no evidence that they were so pinned by
Linnsus himself, this raises no presumption that he supposed the pinned
specimens to belong to the same species.
Of the 23 specimens one is a minute example of Lithospermum apulum ;
nine are Lobostemons from South Africa; one is Echium candicans from
Madeira, and the remaining twelve are European species.
For the sake of clearness, all writing on the sheets is here quoted in italics
between inverted commas, and followed by the letters L., L. fil., or J. E. S. ;
the last standing for Sir J. E. Smith, all whose notes are written in pencil.
List OF SPECIMENS.
A. Inscript. “ candicans. Madera Mason,” L. fil., and a tergo “ Echium
plantagineum thyrsiflorum in rupis” (sic) “ altioribus, Mason ex
litteris? L. fil. A fine specimen.
B. Inscript. “ 3. vulgare,” L. Itis sine loco. ,
C. Inscript. “ plantaginifolium H. U.," L., indicating that it is a cultivated
specimen from Hort. Ups.
D. Inscript. * A. 39,” L., and “ Was pinned to vulgare. E. orientale
H. B.?” J. E. S. Sine loco, but probably received from Alstroemer.
E. Inscript. *.L. 152," L. Sine nomine aut loco, but the inscription
indicates a plant received from Loefling, and therefore from Spain or
Portugal. Below this sheet is pinned sheet G.
F. Inscript. * E. violaceum H. U.," L. Cultivated specimen from Hort.
Ups.
G. Inscript. * 452a," L., and a tergo “ Echium montanum parvum, flore
magno. n montibus Espartal. Lo...” L. The last three letters
of the collector’s name are undecipherable, but he was probably
Loefling. This sheet is pinned below sheet E.
H. Inscript. “ 2 italicum,” L. Sine loco. Below this sheet is pinned
sheet J.
I. Inscript. “ pyrenaicum H. U.," L., et a tergo “flores purpurei," L.
A cultivated specimen from Hort. Ups.
J. Inscript. “ e" L., signifying that it came from Gerber, who collected
in South Russia. This sheet is pinned below sheet H.
K. No inscription.
L. No inscription.
M. Inscript. “ Echium lusitanicum folio amplissimo Tourn." in the hand-
writing of Jan Burman, then “ lusitanicum” in that of Linn. fil.,
followed by “?” J. E. S.
LINN. JOURN.—BOTANY, VOL, XLIV, 2I
396 MR. C. C. LACAITA: A REVISION OF
N. Inscript. * ereticum 4,” L. and a tergo “ Echium creticum,” L.
O. Inscript. “ Echium argenteum 145,” L. The number is doubtless that
of the specimen in the list of those sent by Tulbagh to U psala about the
year 1767, which was published as a Supplement to the * Proceedings
of the Linnean Society for 1917-1918. In that list No. 145 is said
to come from the Swartberg, but is there referred by Linnzeus to
Zehiwum fruticosum.
P. Inscript. “ Sp. 86,” L., indicating that it came from Sparrman, who
collected in South Africa.
Q. Inscript. “ capitatum,” L.
R. Inscript. “ Echium fruticosum,” L.
S. Inscript. “ fruticosum 98,” L., which again suggests Sparrman.
T. Inscript. “ fruticosum,” L.
U. Inscript. “ levigatum,” L., and “H. D. et Willdenow,” J. E. S.
meaning Herb. Banks. —
V. Inscript. “levigatum,” L.
W. Inscript. ** Echium spicatum,” I. fil.
Z. No inscription by Linnæus, but * Lithosp.apulum?” J. B.S. Smith's
determination seems correct.
Hpi
w.
(I.) ECHIUM CRETICUM of Linnæus is an inextricable compound of two
distinct species :—
1. E. creticum angustifolium rubrum, C. B. P. = E. angustifolium, Mill.*
(1768), non Lam. =Æ. hispidum, Sibth. & Sm. (1806) =Æ. elegans, Lehm.
(1818) = Æ. Sibthorpii, Roem. et Sch. (1819).
2. E. creticum latifolium rubrum, C. B. P. — E. creticum, herb. Linn.
=F, australe, Lam., non Coincy, a garden form certainly not derived from
any native of Crete or the eastern Mediterranean, and. not specifically dis-
tinguishable from Æ. grandiflorum, Desf.
The herbarium specimen is identical with that of X. australe in hb.
Lamarck, which I have examined. I have discussed Lamarck’s species in a
preceding paper, which should be read with the present observations. It
only differs from E. grandiflorum, Desf., in having corollas about one-third
smaller. Its identity with the species of Desfontaines has already been
claimed by Moris, who visited the Linnean herbarium and remarks in Fl.
Sard. iii. p. 128, * Herbar! specimen e planta luxuriante.” De Coiney
declined to grapple with the problem of the identity of E. eretieum, L. In
May 1918, I received seeds of qrandifforum from Prof. Battandier of
Algiers, which by the end of July had produced plants in my garden at
* The justification of the revival of Miller's name, which has been overlooked. will be
found in my paper on the eAia of Miller's * Gardener's Dictionary,’ infra, p. 497.
SOME CRITICAL SPECIES OF ECHIUM. 397
Selham, Sussex, identical with the Linnean specimen. The later sowing
shows diffuse lower branches lying on the ground, which explain the * eaule
procumbente" of the misquotation (see below) in Sp. Pl. p. 139. The
corollas vary in the same plant from 28 to 35 mm., but hardly attain the
largest Algerian size.
The herbarium specimen of Æ. ereticum consists of two pieces on the same
sheet, possibly from the same plant. The larger is the upper part, about
35 em. long, of a plant in flower with a branched stem. The six branches
vary from 11 to 15 em. in length. They are therefore approximately equal,
giving an appearance of a diffuse habit. In this piece the lower calyces are
as much as 2 em. apart and barely 1 cm. long. They are exceeded by the
bracts, the lower of which are sub-foliaceous, but the middle and upper
exactly match those similarly placed in Æ. grandiflorum, Desf., being broadly
ovate at the base and acuminate, with the two sides conspicuously unequal.
The smaller piece is a single branch in fruit, 26 cm. long, but not yet fully
extended as the uppermost corollas are still in flower. This has fruiting
ealyces only 1 em. apart, but 1-14 em. long with unequal laciniæ.
The corollas are very oblique, shaped as in Æ. grandiflorum, about 2:5 em.
long in sicco, and less than 1 em. wide at the throat, softly pubescent outside
without long hairs on the veins ; the lower lobes shortly and closely ciliate,
but the upper without cili. The stamens are shorter than the upper lobes
and are best described as subexsert; they are very hairy. The colour of the
corola is now a dark dingy purple (quite unlike that of the specimen of
violaceum, which has dried blue), and has obviously been red in the living
plant. The leaves vary from ovate to oval, being much broader (8-10 em.
by 3-4 em.) than in any of the specimens of grandiflorum from south-eastern
Franee, but only a little broader than in some of those from the Spauish
border and from Algeria. The lateral veins are conspicuous, as shown in
Desfontaines's figure of grandiflorum. Their indumentum is dimorphous,
with few long bristles arising from inconspicuous tubercles, It is, in fact,
that of grandiflorum, but scantier, which I attribute to the effect of cultiva-
tion. As in that species the dried plant is the colour of tobacco.
To sum up: this plant appears to be a garden form of grandiflorum with a
more widely branching habit and wider leaves than normal. Though
identical in shape, pubescence, and eolour, the corollas are not so large as in
fine specimens of grandiflorum. Desfontaines’s figure (Fl. Atl. tab. 46) shows
a less widely branching plant with narrower steni-leaves than the Linnean
specimen of ereticum, which is well represented by the figure in Sweet’s Brit.
Flower Garden, tab. 101 (April 1st, 1825), under the name of E. australe.
This was a garden plant grown from seeds received from Germany under
that name. Sweet’s description is well worth consulting.
Specimens corresponding with that of Linnzeus are not uncommon in the
old herbaria, They all are undoubtedly taken from garden plants.
212
398 MR. C. C. LACAITA : A REVISION OF
The question now arises whether the name Æ. ereticum, L., should be used
either for Æ. australe, Lam. = E. grandiflorum, Desf., or for E. angustifolium,
Mill. =F. elegans, Lehm., or whether it should not rather be entirely dis-
carded as a nomen confusum. The following considerations will, I think,
establish that it must be disearded :—
It will be best to begin with the history of the name eretieum, which first
appears in Clusius, Hist. ii. p. 164 (1601). He describes two species as
E. creticum I. and E. creticum II., both from garden plants. His descrip-
tions are not in themselves such as to allow of certainty, but it is pretty
clear that No. I. is the ereticum of herb. Linn, and No. II. is Æ. angustifolium,
Mill. He observes * bina porro alia Æchii genera, Cretico semine, quod
mittebam, nata, creverunt Joanni ab Hogelande, anno MDXCIII....
accipiebam autem semina, ex quibus hæ plante nate, non Kehit appellatione
sed Anchuse, cujus semen e Creta allatum esset." The use of the mood in
* esset ” indicates that though the seeds were supposed to have come from
Crete, this was uncertain. This little turn of the Latin did not escape
Linnzeus, who says in Hort. Cliff. of No. II. “crescit forte in Creta." The
Cretan origin may be accepted for No. II. but not for No. I., as no Echium
resembling the Linnean specimen or those of the other old herbaria has ever
been found in the island, Æ. ereticum, Fl. Gr. being notoriously K. parvi-
Illorum, Moench.
The next mention of the name is by C. Bauhin, * Phytopinax,’ p. 489 (1596),
where, distinguishing Æ. rubrum ereticum from L. vulgare, he says “ minus
est, sed ramosius, paucioribus floribus ; eleganter rubent; tota planta
Lyeopsin alteram anglieam Lobel icon. p. 579 plurimum refert." The
allusion to Lobel's figure, which represents Æ. italicum or E. pyrenaicum
(see my notes on Miller's Echia), shows that Bauhin meant Clusius No. II.
= E. angustifolium, Mill. Afterwards, in the * Pinax^ of 1623, p. 254, he
enumerated without diagnosis two species, Æ. creticum latifolium rubrum and
LE. creticum angustifolium rubrum. These are the names which constantly
recur in later authors and obviously correspond to Clusius's creticum I. and
ereticum II.
It is uncertain, and indeed immaterial, to which of these two should be
referred the Echium | Candice flore pulchre rubente of J. Bauhin, Hist. iii.
p. 589 (1651), distinguished from Æ. vulgare as * magis ramosa, flores forma
similes, nempe hiantes, sed colore diverso pulchro, scilicet rubente." Though
I think it more probably corresponds to ereticum I.
Linneeus first speaks of Æ. ereticum, in Hort. Cliff. p. 43 (1737), as follows:
“ Echium caule simplici, foliis caulinis linearibus, floribus spicatis ex alis.
E. ereticum angustifolium rubrum Bauh. Pin. 2545 Boerh. lugdb. i. p. 194.
7. creticum 2 Clus. Hist. 2, p. 145. Crescit forte in Creta, unde Clusius
semina habuit," The Hort. Cliff. plant is therefore identical with Æ. angusti-
k
i
:
]
SOME CRITICAL SPECIES OF ECHIUM. 399
folium, Mill., to the exclusion of E. creticum latifolium, which explains why
the specimen of the latter in Herb. Hort. Cliff. bears no name.
We next find in Hort. Ups. p. 35 (1748) “ Echium calycibus fructiferis
distantibus [1]; E. caule simplici f. caulinis linearibus etc. Hort. Cliff. 43 [2];
E. ereticum angustifolium et latifolium rubrum Bauh, Pin. 254 [3]: Æ. rubro
flore Raj. hist. 499 [4]: Hab. in Syria unde semina habui [5]; Differt (sc.
ab. KE. italico, specie priecedenti) caule magis ramoso, magis folioso, foliis
magis ovatis, corollis maximis rubris [6].” So here confusion begins. To
analyse the six statements I have numbered : [1] is his new diagnosis which
he afterwards repeats in Sp. Pl. It is very characteristic of the herbarium
specimen, of some examples of X. plantagineum, and also of E. grandiflorum,
but is inapplicable to K. angustifolium. [2] Can only apply to E. angusti-
folium. [3] Mixes up Bauhin’s two very distinct species. [4] Is quoted in
error, as this synonym belongs to /. rubrum, Jacq. See my notes on that
species. [5] Seeds of Æ. angustifolium may well have come from Syria,
where that kind is plentiful, as well as in Greece and Crete, but neither
angustifolium nor any other Syrian plant could have given rise to the
herbarium specimen *. [6] The whole of this observation only agrees with
the herbarium specimen, admitting of Clusius No. I. The **corollis maximis
rubris" is very important. It is consistent with only two species, viz.
E. amenum, Fisch. & Mey., and £. australe, Lam., var. macranthum, Coutinho
= E. grandiflorum, Desf. But E. amenum is out of the question, so that
grandiflorum is the only alternative. Both * maximis" and “rubris” are
fatal to Æ. eretieum, Fl. Gr., which notoriously is Æ. parviflorum, Moench,
and “ maximis " excludes Æ. angustifolium.
We now come to Z4 creticum, Sp. Pl. p. 139, where there is no fresh
diagnosis but only the quotation of three synonyms, that from Hort. Ups. f,
that from Hort. Cliff., and the Æ. ereticum latifolium et angustifolium, C. B. P.
Also * Habitat in Creta? and the obs. ** stamina non longiora labio breviore
corolla.” The note about the stamens agrees fairly well with the herbarium
and with Æ. grandiflorum, but excludes Æ. angustifolium.
Some further light (?) is thrown by two incidental remarks in Mant. ii.
p. 202, where Linnzeus says of Æ. plantagineum “ corolle violaceze, non
cæruleæ Æ. vulgaris nec breviores Æ. cretici? and “bracteæ longitudine
* [t is possible that Linneus may have received the seeds of his herbarium specimen
from his frequent correspondent Gouan, who might easily have obtained E. australe from
Southern France and forwarded it under the name of creticum without mentioning its origin.
In the British Museum there is a specimen like that of Linnæus from Hort. Gouan, unnamed
by him, but labelled by Solander first ereticum and subsequently lusitanicum ! !
f Linneus bere strangely misquotes his own Hort. Ups. by altering fructiferis to fructes-
centibus and adding the words caule procumbente, which are not found in Hort. Ups.
400 MR. C. C. LACAITA : A REVISION OF
calycis non subulatze Æ. vulgaris nec calyce longiores E. eretici”? The remark
about the bracts agrees with the herbarium and excludes Æ. angustifolium.
The difficulty is the phrase * nec breviores Æ. eretici”? There must be some
slip of the pen or of the printer. The most naiural interpretation of the
words is “not shortish as those of Æ. creticum”*. But that would be in
direct conflict with fact and with the * corollis maximis ” of Hort. Ups. We
should have to suppose that his thoughts had shifted to X. creticum angusti-
folium, for its corollas are smaller than those of 77. vulgare and much smaller
than in Æ. plantagineum. But this supposition does not seem compatible
with the immediately following remark about the bracts. The only alterna-
tive is to suppose either that cretici is a misprint for cretico, or that the word
ilis has dropped out after breviores, and to interpret “not shorter than
E. creticum” or “ not shorter than those of X. ereticum." This would make
good sense and fit the facts, but it would put a great strain on the words as
they stand.
Lastly, in Mant. p. 334 (1771), Linnæus added “ E. eretieum ; corolla
saturate rubra, tubo calycibus breviore. Filamenta apice parum pilosa.’ The
hairy filaments exelude angustifolium in which they are glabrous, but are
most characteristic of australe and of its large-flowered form. The hairs can
be seen in the herbarium specimen, of which also the corollas look as if they
had been “saturate rubr," judging by their present dull dark reddish
appearance. The phrase “tubo calycibus breviore," however, presents a
slight difficulty, as it does not seem compatible with “ corollis maximis," Tt
may be meant only to indicate that the calyx-segments are remarkably long,
as indeed they are in grandiflorum and in the herbarium specimen f.
Can we be surprised that, after Linneeus had so hopelessly mixed up two
species which bis predecessors had kept distinct, his followers should have
found themselves perplexed? It would be sheer waste of paper to go
through all the ways in which the name creticum has been used subsequently ;
de Coincy calls it “Vinévitable creticum que tous les anciens botanistes ne
manquaient pas d'appliquer à tous les Echium qu'ils ne savaient pas nommer."
It will be enough to say that the creticum of Poiret i, of Lehmann, and of
* Is it possible that these words may have led Sibthorp, and consequently Smith, to apply
the name ereticum to E. parviflorum, Moench ?
T The way in which authors speak of the corolla tube being longer or shorter than that
of the calyx in this genus is very vague, as they do not define the point at which they con-
sider the tube of the corolla to end and the limb to begin. As a rule, there is no marked
separation.
1 Poiret, Diet. Encyc. viii. p. 671, makes two varieties of Æ. creticum, referable respec-
tively to E. angustifolium, Mill, and to E. sericeum, Vahl, wrongly quoting (together with
sundry other misapplied synonyms) Æ. ereticum latifolium rubrum, Tourn., for the first, and
E. creticum angustifolium rubrum, Tourn., for the second, with the observation, * La
première recueillie en Syrie par M. de Labillardiére, est bien celle de Tournefort, comparée
avec son herbier, et la seconde celle de Forskal. V. s. in herb. Desfont." "This statement is
criticised as follows by Gay in a note in his herbarium: “ Poiret a décrit son creficum sur
deux échantillons conservés dans l'herbier Desfontaines et rapportós de Syrie par Labillar-
SOME CRITICAL SPECIES OF ECHIUM. 401
de Candolle’s ‘Prodromus?’ are inextricable muddles of plants and of
synonyms.
The conclusion then is :—
(1) The Linnean specimen and other similar garden forms should be
called E. australe, Lam., not E. ereticum.
(2) E. ereticum, L., is a confusion of two totally distinct species, and a
confusion so intricate that the name cannot be used for either of
them, or indeed for any species, but must be rejected as a nomen
confusum.
The following old specimens, all undoubtedly garden plants, are the same
as that of the Linnean Herbarium. All except the last two are at the
British Museum :—
1. From Hort. Cliff., unnamed.
2. Herb. Miller, labelled in Miller's hand “ Æ. calycibus fructescentibus
ete. H. U. 35 ; E. ereticum latifolium rubrum” and by Solander
“ E. creticum.” This is the smaller piece on a sheet, the larger
piece on which is Æ. plantagineum, although the names seem
intended to apply to both pieces.
. Chelsea garden, no. 1367 of 1749, labelled ** Æ. ereticum angustifolium
rubrum C. B. P4" “ angustifolium” being obviously a slip of the
pen for * latifolium.”
4. Herb. Leche ; two pieces, one labelled originally “ lusitanicum ? but
subsequently * eretieum L. H. S." (Linuzi Hortus Siccus), the other
labelled * ereticum Sp. Pl. 139.”
Hort. Gouan, unnamed by him but labelled by Solander first “ ereti-
cum ” and subsequently * lusitanicum.”
Herb. Pallas, labelled originally “ violaceum” and then by Solander
LI
os
or
for)
also “violaceum”? There are two pieces on the sheet, one being
creticum, Herb. Linn. but the other plantagineum.
7 and 8. Herb. Pallas, both labelled originally ‘ creticum,”
then by
Solander first “ ereticum ” but subsequently “australe.”
9. Herb. Pallas, labelled originally “ E. creticum hortense,” then by
Solander first “ ereticum ? and subsequently “australe.” There are
two pieces on the sheet, one being ereticum, Herb. Linn., but the
other is Æ. amaenum, Fisch. & Mey.
diére. Il y rapporte l'E. cret. latif. rubr. de Tournefort, et en lisant sa descr. on croirait
qu'il a vu cette plante dans l'herbier de Tournefort. I] est pourtant certain qu'elle n'y est
point. A sa place on trouve IE. cret. angustif. rubr. Tourn. que Poiret rapporte mal à sa
var. B. et qui est identique avec les échantillons de Labillardiére." Gay is right, but in
herb. Desf., besides the Labillardiére specimens from Syria, which are just E. angustifolium,
Mill., there is one of Delile's from Egypt, which is typical Æ. sericeum, Vahl (see my notes
on herb. Jussieu, no. 6630). Now typical sericeum has even narrower leaves than angusti-
folium, which is the probable explanation of how Poiret came to misepply the two Bauhin
phrases adopted by ‘Tournefort.
ERNST.
402 MR. C. C. LACAITÀ : A REVISION OF
10. Herb. Smith * ex Herb. Davall. 1802.”
11. In the Fielding Herbarium at Oxford, an old example sine loco marked
“ ereticum.”
These specimens reveal how vague were Solander’s notions of his great
master’s species of Echium. There are sundry other old specimens at the
British Museum marked * ereticum ? which are neither creticum of the
herbarium nor angustifolium, Mill. For instance :—
a. Herb. Sloane, vol. 198, p. 50, from Petiver’s * Plant:e Rayan’ marked
** E. eret. latif. rubrum” is E. plantagineum.
b. Herb. Sloane, vol. 306, p. 154, from Herb. Uvedale, similarly labelled,
is Æ. lusitanicum (E. Broteri).
c. Hort. Cliff. labelled ** Æ. creticum latifolium rubrum” and * creticum ”
is E. plantagineum.
d. Hort. Cliff., labelled “ Æ. creticum angustifolium rubrum” (though
identical with the preceding specimen) is also plantagineum.
(II.) ECHIUM ITALICUM and ECHIUM PYRENAICUM are best dealt with
together, though I consider them to be quite distinct species, italicum being
identical with Æ. altissimum, Jacq., but E. pyrenaicum with £. pyramidale,
Lap.
The diagnosis of italicum, without the name, first appears in Hort. Ups.
p. 85 (1748) as X. corollis viz calycem eacedentibus, margine villosis, with three
synonyms: (1) Æ. majus et asperius flore albo, Bauh. pin. 254. Raj. hist. 498 ;
(2) Eehii. altera species, Dod. pempt. 631: and (3) Lycopsis Bauh. pin. 255.
There follows the obs. ** Differt ab chio vulgari, Fl. Suec. 158, foliis palli-
dioribus, angustioribus, levioribus ; racemis minoribus, corollis cinereis
s. ceruleo-exalbidis vix calyce longioribus," also “ Habitat in Anglia, Italia.
Hospitatur in frigidariis, biennis.” The diagnosis and obs. therefore tell us
that (a) the corollas are small, but exaggerate their smallness, (5) that they
are villous on the margin, (c) that they are pale, grey or bluish white.
(a) and (b) admit of any of the forms that have claimed to be italicum of
Linnzeus, but (c) definitely excludes pyrenaicum, in which the corollas are
pink, with the beautifully coloured stamens so carefully described in the
Mantissa and by Lapeyrouse. And it is to be remembered that in both cases
Linnsus was acquainted with the living plant, so that we are not thrown
back on the synonyms to so great an extent as in some cases.
The habitat Italia would cover both altissimum and pyrenaicum, though the
latter only occurs there within a limited area. Anglia is impossible for
either*. To the statement biennis I shall refer hereafter.
To come to the synonyms : (1) C. Bauhin's name is undoubtedly the well-
* In quoting Anglia, Linnæus only followed a mistake of the old English botanists, which
I have traced and fully discussed in my paper on the Echia of Miller's ‘Gardener's
Dictionary.’
SOME CRITICAL SPECIES OF ECHIUM. 403
known whitish-flowered plant figured by J. Bauhin, Hist. Pl. iii. p. 586
(1651), that agrees with Linn:eus's * corollis cinereis ete.” ; and of which
an extra luxuriant individual is figured by Jacquin as altissimum. What
Ray says of this synonym is all important, for he had himself observed the
plant “in agro Salernitano et Romano, inque Etruria et Gallia Narbonensi."
The first three regions are precisely those districts of Italy where italicum =
altissimum is extremely plentiful, but where pyrenaicum does not exist.
The former is most abundant in the Roman Campagna. The traveller can
easily recognise it from the window of his railway-carriage all the way from
Bologna to far beyond Naples. If he travels on to Taranto it will accompany
him from Salerno across the watershed of Potenza and far down the valley
of the Basiento. But he will never see it in the plains of Apulia, where it is
replaced by pyrenaicum, Ray's next species, called by him “ Echium majus et
asperius flore dilute purpureo Botan. Monsp.," both italicum and pyrenaicum
being found, the latter more commonly, in the Montpellier district.
(2) Dodoens's account of his Jehii altera species * is far from clear ; but
as his figure is a mere reproductien of that of Lobel's Lycopsis altera anglica,
which first appeared in Stirp. Hist. p. 312 (1576) and is repeated in his Ic.
Stirp. p. 579, it really coincides with the next synonym (c), and is referable
to E. pyrenaicum.
(3) The Lycopsis of C. Bauhin is, as I have explained fully in my paper
on the Echia of Miller's ‘Gardener’s Dictionary, a muddle of X. plantagineum
with Æ. pyrenaicum as represented by the above-mentioned figure of Lobel.
Linnæus was obviously unaware that plantagineum was mixed up in this syno-
nym when he subsequently quoted Lycopsis for his italicum B= pyrenaicum.
Echium italicum as a name starts in Sp. Pl. p. 139 (1753), with the
diagnosis of Hort. Ups. repeated and only one synonym, Æ. majus et asperius,
Cam. epit. 728, which of course is the same as the identical Bauhin phrase,
but a 8 makes its appearance, without any diagnosis or comment, to receive
the Lycopsis synonym. Then in the second edition, p. 200, a new diagnosis is
substituted, Æ. caule erecto piloso, spicis hirsutis, corollis subequalibus, stamini-
bus longissimis, the synonyms being his own diagnosis of Hort. Ups. and those
of C. Bauhin and Camerarius, with Lycopsis again referred to var. 8, of
which now we obtain some account, to which I will return when speaking of
E. pyrenaicum. Linnæus’s later works give us no further information about
his typical italicum, though they bring fresh matter for var. 8 pyrenaicum t.
Unquestionably the plant that, according to the usual and legitimate
practice, must bear the Linnean specific name is the first form described,
especially seeing that a different name for var. 8 appears in the Mantissa and
in the herbarium for the specimen corresponding to that variety. The only
specimen in the herbarium marked italicum by Linnæus is precisely the
* P. 620 in ed. of 1583, but p. 631 in that of 1616.
t The reference to Hudson's Fl. Angl. in Sp. Pl. ed. 2, p. 1678, mentioned below, only
adds confusion, not information.
404 MR. C. C. LACAITA : A REVISION OF
Echium majus et asperius flore albo so common in Italy. Its identity with the
E. altissimum of Jacquin is proved by comparison with an authentic specimen
of the latter in the British Museum * herb. Jacq." and by numerous modern
exsiccata, of which I have selected those in my list A as being absolutely
identical with the Linnean specimen.
The Linnean specimen of italicum is an upper part only, without root or
lower leaves; the narrow inflorescence measures about 30 by 5 em. ; it is in
flower, so the cincinni are not yet fully unfurled. — Corollas and filaments now
show a dirty eream-colour. In similar specimens the fully developed radical
leaves are very long and narrow, 30 by 3 cm., even 30 by 2 em. ; the inflores-
cence in the flowering state averages 45 by 5-6 cm., even reaching 55 by
6 em., but broadening after the cincinni are fully unfurled to 45 by 8-12 em.
Jacquin’s figure, Austr. tab. 16 (in App.), shows a more than usually
luxuriant plant. In his admirable description, v. p. 35 (1778), he remarks
“corolla perpetuo alba ; filamenta alba ; antheræ cum virore flavescunt,”
which agrees with Italian examples. It may be asked how Jacquin came to
create an Æ. altissimum if it is identical with Linneeus’s italicum. The reason
is simply that from Linneus’s brief diagnosis he could not know that the
species said to live in Italy and England was his own Hungarian plant.
He makes no attempt to distinguish altissimum from italicum, which he does
not so much as name. Subsequent authors, especially the French, followed
by Kerner in Oest. Bot. Zeit. xxiii. p. 164 (1873), and in Fl. Exs. Austr.
Hung. nos. 3174 and 3175, assuming, without sufficient thought, that Jacquin’s
and Linnzus’s names must indicate different plants, had to find another to
play the part o£ italicum and unfortunately pitched upon pyrenaicum as being
the true italicum, regardless of Linn:us's own descriptions and without
dreaming of consulting nis herbarium.
The following attempt to describe is based on Italian specimens, but it
must be remembered that in the case of so large a plant, herbarium specimens
are either incomplete or are undersized individuals, which do not always
exhibit characteristic habit.
ECHIUM ITALICUM ; unicaule, sive in exempl. luxuriantibus, pluricaule *.
Caules in typo 40-100 em. alti, simplices, non nisi ramulis singulis ex
axillis preediti (in var. Biebersteinii subramosi propter ramulos inferiores
ipsos cincinnis pluribus brevissimis dissitis munitos).
* This character is claimed specially for E. albereanum, Naud. & Debeaux (=L. flavum,
Lap., non Desf.), by Roumeguére in Soc. Agr. Pyr. Or. xxii. p. 163, note, * la plante est
multitige, le type, on le sait, est constamment unitige," but the writer meant pyramidale,
Lap. not true italicum, by “le type”; Debeaux remarks, op. cit. xxiii. p. 176, “cette
espèce se distingue à premiere vue de IE. pyramidale par ses tiges élevées, nombreuses dès
la base et non unitiges.” Admirable to distinguish it from pyrenatcum — pyramidale, but
the character is common to all forms of true italicum, and -often occurs in strong plants of
E. vulgare.
SOME CRITICAL SPECIES OF ECHIUM. 405
Hamuli, infimis exceptis, simplices, usque ad 1-2 em. nudi, ibique robus-
tiores in cincinnos geminos :equales bifurcantur, debiliores cincinno
unico terminantur.
Cincinni initio brevissimi, valde scorpioidei, deflorati erecti vel erecto-
patentes usque ad 8-10 em. (15-20 em. in var. Biebersteinii) producuntur.
Inflorescentia in typo spiciformis, sub anthesi c. 45 x 5 em., post anthesin
c. 45 x 8-12 em. ; in var. Biebersteinii conica, propter axem centralem
ramos inferiores, quamvis elongatos, fere duplo excedentem. Flores,
ut in genere, distiehi, subunilaterales, confertissimi, bracteis anguste
lanceolatis calycem (laciniis inclusis} c. 6-7 mm, longum æquantibus.
Corolle parvee subregulares, c. 12 mm. longze, tubo fere recto, extus pilose,
pilis paucis longioribus strigosis intermixtis, sordide albz vel sube .
ruleo-albescentes ; filamentis concoloribus. Stamina longissime exserta
seepius corollam fere duplo superantia.
Folia radicalia angusta, longa vel longissima ; evoluta 30 x 2-3 cem.;
caulina lanceolata, superiora 4-8 em. lo:
excedentia *.
iga, cincinnos floriferos longe
Indumentum e pilis strigosis, confertissimis (precipue in var siculo),
inzequalibus sed homomorphis, in sicco lutescentibus f.
But the name italicum L., though it must exclude pyrenaicum, will cover
the breader conical form which oceurs frequently in the Balkans and in
Greece, and in Southern France seems to be more usually met with than
altissimum, from which, however, it certainly cannot be specifically separated,
having the same system of ramification with the very predominant central
axis, and the tendency, even in a greater degree, to produce subsidiary stems
from the base. It also has the long, narrow, hairy, hardly tuberculate root-
leaves, the small pale yellowish or bluish-white corollas, showing cream-
colour like the pale filaments in sicco, and the yellowish indumentum. The
outline of the inflorescence is conical rather than pyramidal, the height being
at least double that of the base; it measures about 40 em. by 20-25 cm.,
and shows a similar ratio in smaller examples. This outline is due to a larger
number of the lower axillary branches being prolonged, and themselves
bearing rather widely spaced short branchlets, each consisting of single or
twin cincinni. I am unable to discover any other difference from altissimum,
and as it does not seem really to be a geograpical race, altissimum also being
found in the Balkan region, it is almost going too far to give it a name as a
variety. However, for greater clearness I shall call it var. Biebersteinii =
* So Moench, Suppl. p. 149 (1802), says of his linearifolium = italicum, L., * caule erecto
simplici, spicis foliis brevioribus."
T E. luteum, Lap., Abr. Pyr. p. 91, is said by the author to be covered with pungent
hairs, long and. yellow, as well as the flowers; but there is not a word to suggest that the
corollas themselves are yellow, except the erroneous reference of the name to Desfontaines,
who has no E. luteum. Of course, his E. flavum was inteuded, which, however, is an
altogether different species.
406 MR. C. C. LACAITA : A REVISION OF
Echium asperrimum, MB. (non Lam.) Fl. Taur. Cauc. i. p. 135 (1808), where
tlie diagnosis, quoted from the author's Tabl. Casp. (1798), runs * E. strigo-
sum ramosum, spicis axillaribus conjugatis paniculatis.” — In vol. iii. p. 131
(1819), he adds “corolla constanter apud nos leucoph:a, extus levitur
pubescens, pilis longioribus perpaucis. Pro Æ. italico L. hoe demum habeo."
This form of italicum has unfortunately been confused by many French
botanists * with the specifically different E. pyrenaicum = E. pyramidale,
* [n judging the French specimens that I refer to one form or another of E. italicum to
belong to the same species as those from central Italy, in spite of some discrepancy in the
alleged colour of the corollas, in reducing Æ. albereanum to a mere form of E. italicum, and
in identifying the huge E. pyrenaicum of Apulia with the pyrenaicum — pyramidale of the
vicinity of Toulouse, I have had to rely entirely on herbarium specimens of the French
plants, as it has not been possible in war-time to study them in their native haunts. It is,
of course, possible that as intimate an acquaintaince with living French examples, as I have
with Italian, might modify some of my opinions.
Lapeyrouse, Abr. Pyr. pp. 89-91 (1812), mentions three species of the italicum group
for the Pyrenean region: E. italicum, E. pyramidale, and E. luteum. For italicum he quotes
the old diagnosis, mentioning that the plant grows by roadsides etc., and observes “ tiges
droites, élancées, rameaux courts égaux," which agrees well with the Linnean italicum ; but
itis difficult to understand how he can say “ fleurs distantes, secondes solitaires” of any
member of this group. — Pyramidale is spendidly described, quite in accord with Linnwus's
account of pyrenaicum. -It is “rare aux Pyrénées .... à foison le long des chemins prés
Toulouse.” Its rarity in the Pyrenees may account for its misinterpretation by some authors.
The diagnosis of luteum, being that of the totally different favum, Desf., is useless, and it is
only from the habitat and the observation that the species ean be identified, practically witli
certainty, with Æ. albereanum, Naud. & Debeaux.
Bonnet, whose views have been rather uncritically adopted by Rouy in Fl Fr. x.,
synonymises in Bull. Soc. Bot. Fr, xxv. p. 209 (1878), E. luteum as well as E. pyramidale
with E. pyrenaicum, L. Mant., quoting the exsice. Bourgenu, Fréjus, no. 285, and Billot,
no. 2395. But both these are italicum (see my list B. nos. 4 & 5), not pyrenaicum, Then
for italicum he quotes Reichb. no. 995, Jamin Pl. Alger. no. 184, and Billot 2325 bis et ter.
Now Reichb. no. 095 is true tadicum, but the other three are unquestionably pyrenaicum ! ! !
(see my lists A and C). No wonder then that Bonnet's descriptions are unintelligible and
contradictory. P
Then Timbal-Lagrave in Soc. Agr. Pyr. Or. xxiii. p. 175 (1878), whilst insisting that
pyrenaicum and italicum are different species, misinterprets both, for his italicum is really
pyrenaicum, and his pyrenaicum or pyramidale is not Lapeyrouse's plant but obviously
. albereanum, a variety of true italicum. Thus and thus only can be explained his remark
“ E. luteum n'est qu'une forme de la méme plante " (sc. pyrenaicum).
This inversion or transposition of the names prevails in Cosson's determinations, and was
subsequently adopted (blindly) by Kerner. It frequently occurs in French exsiccata.
I have examined 41 French examples of so-called italicum and pyrenaicum in herb. Mus.
Paris. Of 26 labelled acum only 11 belong to that species, the remaining 15 are
pyrenaicum. But of the 15 labelled pyrenaicum only 8 are rightly so named, the other
7 being forms of italicum. |
Such confusion has produced its natural result in the amalgamation of the two species by
Grenier & Godron, by Coste, and even by de Coincy, whose views on European species are
not so infallible as on Algerian. The amalgamation is a mere counsel of despair, due,
I think, to confining attention to differences of outline and ramification, and disregarding
other characters.
TU ae SV í r sce T DT.
SOME CRITICAL SPECIES OF ECHIUM. 407
Lap., especially by De Candolle in Prodr. x. p. 23 *, where the two are
confounded under the name of /. pyramidatum, so as to make that name,
apparently so suitable, useless to indicate the form spoken of here, not to
mention the risk of confusion between two such similar words as pyramidatum
and pyramidale.
J. Bauhin's figure of Æ. albo flore majus, already referred to, seems to
represent this variety, to which the specimens in my list B are referable.
The form of italicum called Echium luteum by Lapeyrouse, with an
inadequate diagnosis, and rediscovered in 1874 by M. Naudin in the range
of the Albères above Collioure, and by him named X. albereanum, requires
some notice, especially as it has been regarded by Rouy as closely allied to
E. pyrenaicum, with which it has no resemblance or relationship. It is
described by Debeaux and Naudin in Soc. Agr. Pyr. Or. xxiii. p. 175 (1878),
from examples cultivated by Naudin, as “ dressée-hérissée, très rameuse dans
sa moitié supérieure, de plus de 1 mètre de hauteur, et plurieaule dés la
base . . . . les leurs... . formant par leur ensemble une vaste panicule láche,
diffuse et très ramifiée.” But the specimens I have seen e loco classico at
Soréde, collected by L. Conill, of which the individuals with a broad panicle
were distributed as italicum var. pyramidale (though utterly unlike pyramidale,
Lap.), and those with a narrow inflorescence as italicum var. albereanum,
cannot be separated from Æ. italicum. Both the narrower and the broader
forms are intermediate between italicum altissimum and var. Biebersteinii as
far as ramification is concerned, nor can 1 see any other distinctive character
in sicco. There is some conflict of evidence as to the colour of the corollas,
which are stated to be bluish white or slightly pinkish by Roumeguére in Soc.
Agr. Pyr. Or. xxii. p. 163. note, but yellowish white, with an occasional
pinkish blush, but not pale blue, by Naudin, /. c.
A much more remarkable form— var. siculum, mihi = Æ. italicum, auctt.
sic.—replaces typical italicum in Sicily. Where I have collected it in the
stony fields of the province of Syracuse, it is a relatively low-growing but
broad plant, with a forest of branches, spreading horizontally, when fully
developed, in every direction ; but their arrangement is not pyramidal as
in pyrenaicum, the lower ones not markedly exceeding the middle ones. The
arrangement of the cincinni, the dull white corollas and filaments, and the
size of the flowers are those of italicum not of pyrenaicum, from which it is
also distinguished by the very yellow (in sicco) indumentum. This is extra-
ordinarily copious ; on the stem it is long, patent and rigid, and so close as
completely to conceal the surface.
Todaro’s well-known exsiccata do not show the peculiar habit, as they are
* The treatment of the genus Echium is one of the weakest parts of the * Prodromus.’
The Dorrageze were not completed by Aug. Pyr. de Candolle before his death, nor written
de novo by his son. It is to be remembered that neither father nor son saw the Linnean
specimens or even the plants of the ‘ Flora Graeca,’
408 - MR. C. C. LACAITA : A REVISION OF
only scraps of plants, gathered in a very early stage, but they have the
thick yellow coat of bristles. My list C contains the only herbarium
specimens that I have yet met with of var. siculum.
T have not seen sufficient material from Asia Minor and beyond to attempt
to determine or discuss the Oriental forms. Those that I have seen are
referable to italicum sensu lato, not to pyrenaicum.
To conclude: Mehium italicum, L., regarded as specifically distinct from
I. pyrenaicum (L.), Desf., presents at least three or four marked forms in
Europe :—
l. typicum = K. altissimum, Jacq. = K. lineavifolium, Moench, Suppl.
p. 149 (1802). See specimen list A.
2, var. luteum (Lap. pro specie) = Æ. albereanum, Deb. & Naud. See
specimen lists A and D.
3. var, Biebersteinii, mihi= E. asperrimum, M.B., non Lam.= K. pyra-
midatum, DC. pro parte. See specimen list B.
4. var. siculum, mihi — E. italicum, Guss., Lojac, et auct. sie. omn, See
specimen list C.
*
To come to Æ. pyrenaicum * ; this species first appears in Sp. Pl. ed. 1 as
var. B of italicum to correspond to the Lycopsis of C. Bauhin. Then in ed. 2,
after Lycopsis has again been quoted, there is added, * corollis minoribus,
extus presertim superne pilosis, et magis regularibus ab Æ. italico differre
videtur, nisi hiec omnia e loco." This is no help to the identification of var.
B, for in pyrenaicum the corollas have indeed more and longer hairs tban
in italicum, but they are neither smaller nor more regular. In the Appendix,
p. 1678, Linnæus refers to Hudson's Fl. Angl, which had appeared in the
* I fear that it must be called pyrenaicum, Desf. Fl. Atl. i. p. 164 (1797-1798), as the
rules will hardly allow us to say pyrenaicum, L. Mant. (1771), in spite of the occurrence of
the name in the ‘ Mantissa ’ and the use of it as specific by Linnæus in his unpublished MS.
and herbarium, and of its quotation by Willdenow, Sp. Pl. i. p. 786 (1708). These dates
are correct, notwithstanding those that appear on the titlepage of Willdenow’s first
volume, and in some copies of the ‘Flora Antlantica.’ For evidence of the true date of
Fl. Atl. i., see the note to E. australe in my paper on “ Five Critical Species of Echium,”
supra, p. 368.
Unfortunately, a date so late as 1797-1798 admits of the doubt whether this species ought
not rather to be called Æ. asperrimum, Lam. Ill. i. p. 412 (1791). I think we shall be
justified in rejecting Lamarck's name as ambiguous, though its claims are arguable.
Poiret, Diet. Encyc. viii. p. 668 (1808), who should have known what Lamarck meant, uses
the name for the compound of italicum and pyrenaicum, quoting as synonymous italicum, L.,
altissimum, Jacq., and pyrenaicum, L. Mant. Rouy also, Fl. Fr. x. p. 904, seems to consider
it ambiguous, for he does not adopt it, but quotes it pro parte for both italicum and
pyrenaicum. In Lam. Fl. Fr. ii. p. 451 (1778), we find italicum and pyrenaicum both under
E. italicum ; the former as italicum, L.— E. majus et asperius fl. albo, Toum., and the latter
as var. G— E. majus et asperius fl. dilute purpureo, Tourn. But in Ill. Z c, Lamarck only
says of asperrimum “ E, caule ramoso, pilosissimo; corollis calyce longioribus; staminibus
exsertis. Ex Europa australi. Æ. italicum, L.?" The synonym italicum is queried, and
there is no allusion to his own previous use of that name. The diagnosis would cover
various other species besides italicum and pyrenaicum. So far, therefore, asperrimunm seems
e. SOME CRITICAL SPECIES OF ECHIUM, 409
same year (1762), but the reference throws no light on either italicum or
pyrenaicum, for Hudson had altogether misinterpreted italicum and its var.
B *. neither of which grow in England or Jersey. But in Linnwus's own
interleaved copy of the 2nd ed. of Sp. Pl., now in the possession of the Linnean
Society, he has written opposite Æ. italicum the following note : “ pyrenaicum
Ech. majus et asperius flore dilute purpureo T. 135. Totum valde hispidum
ut fere urens; corolla calyce latior (sie ; obviously a slip of the pen), subin-
carnata, Filamenta corolla longiora saturate rubra. Anth. cæruleæ. Sna.
levia, apice subtrigona, quasi dente intermedio s. interiore productiori.
Lycopsis monspeliaca flore dilute purpureo Moris. blæs. Magn. monsp."
This is obviously a description from life of the plant grown in Hort. Ups., of
which a specimen labelled “ pyrenaicum " is in the herbarium. The under-
lining of the Bauh. Pin. 255 in this copy indicates that at the time he had a
specimen in the herbarium. There cannot be a shadow of doubt that the
description refers precisely to the plant called pyrenaicum by Desfontaines also
and pyramidale by Lapeyrouse, Abr. Pyr. p. 90 (1813). I have examined
Tournefort's own specimen of /. majus et asperius fl. dilute purpureo (no. 586
in his herb.) and found it identical with that of Linnzeus’s pyrenaicum and
with authentic Lapeyrouse specimens of pyramidale at Kew and Mus. Brit.
It is to be noted that in this MS. entry, Linnzeus uses the name pyrenaicum
as that of a species, and not as var. 8 of italicum. He does the same on
the sheet in the herbarium. — It is impossible to guess why in the published
description in the * Mantissa’ of 1771, p. 334, he should have headed it 8 and
not with the name pyrenaicum, though the language there used seems to
indicate that he still regarded it as specifically different from italicum.
The * Mantissa ^ varies very slightly from that MS. entry. After quoting as
synonyms for var. B those of Tournefort and Morison there follow the words
“ Echium pyrenaicum vulgo," and then * Simillimum Z. italico, hispidum et
fere urens, stimulis albis. Cor. calyce non latior, subincarnata (nec alba),
infundibuliformis, fere regularis, calyce duplo longior, extus pubescens palli-
diorque. Filam. duplo longiora, saturate rubra. Anth. cæruleæ. Sem.
lx via, apice trigona, quasi denticulo intermedio s. interiore productiore."
A most admirable description, agreeing perfectly with that given by
to be a nomen dubium to be rejected. On the other hand, Lamarck's immediately following
species, E. elongatum (which puzzled Poiret), is defined * E. caule erecto piloso, longissime
spicato; corollis vix calyeem superantibus, staminibus exsertis. Fl. albi, parvi, sessiles."
If this diagnosis be compared with that of asperrimwm it will be noticed that it contains
three characters by which italicum (altissimum) differs from pyrenaicum, viz.: the erect stem
and very long spike, the smaller corollas, and their white colour. Now Lamarck’s
herbarium contains two specimens bearing on the question. One, labelled “ E. elongatum,
Lam., an E. majus asperius fl. albo?” is E. italicum. The other, labelled * E. italicum.
E. asperrimum, Lam. IL," is pyrenaicum. Considering the way in which Poiret and Rouy
have treated asperrimum, I hardly think these labels are cogent enough to force us to
abandon the appropriate, admirably described, old name of E. pyrenaicum.
* See my Zchia of Miller's * Gardener's Dictionary,’ infra, p. 423.
410 MR. C. C. LACAITA : A REVISION OF .
Lapeyrouse of his pyramidale. The herbarium specimen, taken from a plant
grown in Hort. Ups., consists of one leaf only and two flowering branches
without the main stem, but is quite unmistakable. A list D of characteristic
specimens of Æ. pyrenaicum will be found below. The following description
is founded on French exsiecata from Toulouse, but the Apulian plant seems
to be larger, with more stiffly horizontal branches; this, however, may be due
to the impossibility of exhibiting full-sized French plants in herbarium sheets.
What I must emphasise is that in Italy pyrenaicum and italicum are totally
different, and inhabit separate areas, as was long ago pointed out by Tenore
and Gussone (cf. Ten. Fl. Nap. v. p. 330, and Guss. et Ten, in Att. Ac. Nap.
v. p. 431, 1843).
It is notorious that pyrenaicum alone is found in Algeria ; it predominates
in Spain, where italicum apparently only just crosses the border into Catalonia.
In fact, it is in southern France alone that the two species are often found in
proximity, and it is precisely there that puzzling forms occasionally occur—
whether as hybrids I cannot say.
The Balearic Islands furnish a form with extra hard bristles, but not
otherwise distinguishable. This is Æ. italicum, var. balearicum, Porta & Rigo
in N. Giorn. Bot. It. xix. p. 318 (1887), but it must be referred as a variety
to pyrenaicum and not to italicum.
ECHIUM PYRENIACUM ; (planta Tolosana) semper unieaule, caulibus
secundariis nullis, aut rarissime prresentibus,
Caules e. 50-60 cm. alti., ramosi, ramis inferioribus patentibus, (a) in
planta luxuriante usque ad 45 cm. longis, ramulos floriferos 15-20
gerentibus; ipsis ramulis aut simplicibus fere a basi floriferis, aut apice
in eineinnos geminos fissis; (b) in planta debiliore non ultra 30 em.
longis, basi usque ad 10 em. foliiferis non floriferis, superne cincinnos
abbreviatos simplices, raro gominos, gerentibus ; ramis vero superioribus,
simplicibus, ad cincinnum unicum elongatum reductis.
Cineinni virginei parum scorpioidei (multo minus quam in Æ. italico), in
ramis inferioribus laterales pauciflori, quasi ad florum glomerulos post
anthesin parum elongatos reducti ; in ramis vero superioribus simplicibus
10-20-flori, ideoque post anthesin usque ad 10-15 em. elongati, suberecti.
Infldrescentia pyramidalis, latitudine in plantis evolutis altitudinem :equante
vel etiam superante ; ramis aut horizontalibus, aut (quia inter plantas
circumstantes constrietis 7) adscendentibus, ita ut plures altitudinem
axis centralis attingant *.
* [ suppose this is the character meant by the phrase “rameaux atteignant presque tous
la méme hauteur, ce qui donne à la plante l'aspecte conique,” which Bonnet uses of his
italicum, by which, as explained in a previous note, he intends pyrenaicum. But the same
words are used by Rouy, who does not seem to have noticed that Bonnet transposes the
two names, of his own italicum which is not pyrenaicum. I am afraid this takes away
nearly all the value of Rouy’s account of these species. And, indeed, the phrase indicates
an arrangement of branches which would not produce a cone, but an inverted cone,
SOME CRITICAL SPECIES OF ECHIUM. 411
Flores in cincinnis sieut in congeneribus dispositi, in ramis inferioribus
magis, in superioribus minus conferti ; bracteis lanceolatis, basi latio-
ribus quam in Æ. italico, calycem (laciniis inclusis) usque ad 10 mm.
longum parum superantibus.
Corolla parvae, tubo leviter geniculato (‘tube coudé,” Lap.), minus quam
in X. italico regulares, et paululum quam in eo longiores (15-18 mm.)
extus pilosissiinæ, pilis albis strigosis longioribus quam in Æ. italico,
carne *, filamentis pulcherrime rubris (etiam in sieco fusco-rubris),
antheris eseruleis. Stamina exserta, sed multo minus quam in Æ. italico.
Folia radicalia angustissime elliptica potius quam lanceolata (20 x 4 cm.,
15 x 3°5 cm., 10 x 2 em.), ideoque forma ab illis Æ. italici nonnihil diversa.
Indumentum asperrimum, tactu urens, minus quam in Æ. étalico confertum,
e pilis albis non lutescentibus (ideoque tota herba etiam in sicco grisea),
valde insequalibus, nonnullis apice stellatis.
As exsiccata only consisting of branches torn off, or, if entire, of
exceptionally small inviduals, do not give an adequate idea of the normal
habit of this very large species, I add a note made on the spot from four
Apulian specimens gathered in late autumn, in seed, by the roadside near
Taranto, where the species is very abundant. The two larger are one metre
high and 70 em. broad; the two smaller measure 65 by 55 cm. The larger
plants have from 20 to 30 branches, spreading in every direction, erecto-
patent in the larger, horizontal in the smaller specimens, the lowest branches
being as much as 50cm. long. Each of the longer branches bears 10 to 20
scattered branchlets, pointing in every direction. Of these branchlets the
lower are again branched into 5 to 10 subunilateral cincinni, with their
distichous flowers on the upper surface, but the upper branchlets consist of
single cincinni. These cincinni are nearly all simple, only occasionally
bifid. Even the most developed only bear about 10 flowers, forming at first
tight glomerules, but lengthening later to about 10 em. The fruiting
calyces overlap each other, each covering about half the calyx next above.
Bracts and calyx-segments are lanceolate and very hispid. The fruiting
calyx is not enlarged nor concrete at the base. The plant is everywhere
hispid with white, spreading, stinging sete that give the whole a pale grey
appearance in autumn.
The root-leaves of the young plants first appear in November. Their
surface is dark green and almost hairless in the intervals between the
incipient tubercles, which show as lighter green pustules bearing hardly
any hairs. When radical leaves are present in herbarium specimens they
are obviously broader and shorter than in italicum, more tuberculate and less
* I am not sure that albinos of pyrenaicum may not occur occasionally, just as they do in
E. vulgare and E. plantagineum. This, or hybridity, may account for the very few
examples I have seen—all from France, where both species are found—that are not at a
glance referable to italicum or to pyrenaicum.
LINN. JOURN,—BOTANY, VOL. XLIV, 9M
Ev
412
MR. C. C. LACAITA : A REVISION OF
hairy. But the characters, by which incomplete specimens are most easily
recog
nised in herbaria, are the grey not yellow indumentum, the shorter
hairier corollas with less remarkably protuding stamens, and the dark
filaments. The difference in shape of the corolla-tube is not easy to see
n SIC
20,
List A.
Ecuivm rraticum, L.2 M. altissimum, Jacq.
1. ex Herb. Jacq. in Mus. Brit. probably cult. (as altissimum).
2. Fl. Austr. Hung., no. 3714, Promontor, Hungary (as altissimum, but
identical with no. 12 below).
3. Richter, Fl. Hung., Ofen ; also in Baenitz as from Buda-Pest
(as italicum).
4. Schultz, Herb. Norm. 2868, Wolfsthal prope Budam (as altissimum).
5. Heuffel in Herb. Gay, from the Banat (as italicum).
6. Wierzbicki, from Oravieza in the Banat (as italicum).
7. Sagorski, Fl. Herceg., form Mostar (as altissimum).
8. Sintenis, 1873, no. 302, Babadagh in the Dobrudja (as italicum).
9. Pichler, 6, vi. 1885, from Spalato (as italicum).
10, Huter, 24, v. 1867, from Ragusa (as italicum).
10 bis. Reichenbach, Fl. Germ. exsiec. no. 995, from Fiume (as italicum).
11. J. S. Mill, Rimini on the road to Ravenna (as altissimum = italicum
Lehm., non Linn.).
12. Fl. Austr. Hung. no, 3715, from Verona (as italicum, but identical with
no. 2 above).
13. Fl. Ital. Exs. no. 940, from Florence (as italicum).
14. Webb in hb. Rouy, Roma extra portam Salaram (as italicum).
15. Arcangeli in hb. Florent., from Stilo in Calabria (as italicum).
16. Reverchon, Pl. Sard. 1882, no. 245, from Tempio in Sardinia (as
italicum).
T Pl. Cors. 1878, no. 49, from Bastelica in Corsica (as
italicum).
18. Kralik, Pl. Cors. no. 699, from Zicavo in Corsica (as italicum; this
no. is quoted in Rouy, Il. Fr.).
. ex hb. Grenier in Mus. Paris, from Hyères (as italicum).
20. Thuret, 1868, no. 61, from Antibes (as /talieum).
21. Loret, 1, vi. 1851, from Le Lue (Var) (as italicum).
Naudin in Mus. Par., from Collioure with the obs, * fleurs trés petites
blane-bleuátres, trés différent de PÆ. italicum qui abonde à
Collioure,” is typical italicum showing that his conception of
italicum was not that of Linnæus. This plant became var,
albereanum, Naud,
SOME CRITICAL SPECIES OF ECHIUM. 413
23. Herb. Conill. in hb. Bailey, from Sorède in Pyr. Or. (as italieum var.
albereanum ).
24. Sennen, Pl. Esp. no. 572, from Cabanas, in Catalonia (as E. pomponium,
rar. Paut, Sen. ; it is not a form of pomponium).
List B.
Ecnivm ITALICUM, L., var. BIEBERSTEINII, mihi.
l. Unio itin., in fossis c. urbem Perpignan (as luteum, Lap.).
2. ex hh. Conill in hb. Bailey, Soréde Pyr. Or. (as italicum var.
pyramidale).
3. Soc. Cènomane, 1901, no. 554, from Sorède, identical with no. 2 (as
italicum var. pyramidale, corrected by Sudre in his herb. to luteum).
4. Dourgeau, Env. de Fréjus, no. 285, from the Esterel (as pyrenaicum,
det. Coss. This is quoted for pyrenaicum by Rouy in Fl. Fr. x.
p. 305, but is not that species).
5. Billot, no. 2325, from the Monte Pisano in Tuscany (as italicum, but
quoted by Rouy for pyrenaicum, perhaps from a confusion with
Billot, 2325 bis and ter). ;
6. Frivaldsky in hb. Hooker, from Rumelia (as asperrimum, M. Bieb.).
if in Mus. Brit., from Carlova (as asperrimum, M. Bieb).
8. Stribrny, vi. 1894, from Sadovo in Bulgaria (as italicum).
9. Adamovie, Pl. Balean. vi. no. 190, from Voden in Macedonia (as
italicum).
10. Tardent in hb. Gay, from Bessarabia (as asperrimum, M. Bieb.).
ll. Richter in hb. Kew, from Odessa (as asperrimum, M. Bieb.).
12. Count Puschkin, 1783-1803 in hb. Kew, from Caucasia (as asperrimum,
MB.).
13. Aucher Eloy, hb. d'Orient no. 2397, from Greece (as pyrenaicum).
14. Zuccarini in hb. Kew, from Nauplia in Greece (as 7. pyrenaicum, L.
= E. asperrimum, M. Bieb.).
15. Sieber in hb. Kew, from Canea in Crete (as italicum).
16. Reverchon, 1886, sine numero, from Canea (as italicum).
List C.
ECHIUM ITALICUM, L., var. SICULUM, mihi.
1. Todaro, Fl. Sic. no. 1227, from Sclafani (as italicum).
2. hb. Lacaita, from Ragusa prov. Siracusa (as italicum var. siculum).
2. hb. Florent. contains similar Sicilian specimens from Palermo, Castel-
buono and Girgenti, but I have not examined them earefully.
B xcd uL ds
414
MR. C, C. LACAITA : A REVISION OF
List D.
ECHIUM pyrenatcum, L.
ex hb. Vaillant in Mus. Paris (as Æ. majus et asperius fl. dilute purpureo).
ex hb. Pourret in Mus. Paris from Narbonne (as italicum).
Lapeyrouse in hb. Gay, sine loco (as pyramidale). Other authentic
specimens from Lapeyrouse are at Mus. Brit. and in hb. Smith.
Bentham in hb. Kew, from Villefrance près Toulouse (as pyramidale).
from Restinclières (as pyrenaicum).
. hb. Loret in Mus. Par., from Toulouse (as pyrenaicum).
from Viviers (as italicum).
Billot, no. 2325 bis, from Carcassonne (as italicum, L., non Lehm.).
ter, from Fouras in Charente Inférieure (as the last).
Magnier, Fl. Select. 1250 bis, falaises de Bidart, Basses Pyr. (as
italicum).
Reverchon & Derbez, Pl. Fr. no. 249, from Bollène (as pyrenaicum).
J. S. Mill in hb. Kew, from La Dochelle (as pyramidatum).
Bourgeau, Pl. Hisp. 1852, sine numero, from Fuente de la. Higuera,
Spain (as italicum det. Cosson).
. Gandoger, Fl. Hisp. Exs. no. 258, Renteria in Guipuzcoa (as
pyrenaicum).
. ex hb. Fernandez in hb. Rouy, Guadalajara, S ain.
Y; Jara,
hb. Rouy, from Sierra de Maimon above Velez Rubio (as italicum
altered to pyramidale).
Bourgeau, Esp. 1850, from Fuensanta by Murcia (as italicum).
_ Fl. Ital. Exs. no. 2129, from Spinazzola, Prov. Bari (as italicum var,
pyramidatum).
. Martelli in hb. Florent., from Manfredonia.
Groves in hb. Florent., from Otranto.
from Gallipoli.
Rabenhorst in hb. Kew, from Gargano (as Æ. gaditanum, Boiss.,
which is ridiculous).
Reverchon, Pl. Alger. 1897, no. 243, [rom M. Babors (as italicum var.
pyramidale).
All Algerian specimens called italicum are pyrenaicum.
E. PYRENAICUM var. BALEARICUM, mihi.
24.
Porta & Rigo, 21. viii. 1885, from Majorca (as italicum B balearicum).
(III.) ECHIUM LUSITANICUM, Sp. Pl. p. 140 (1753). — Of this species
Linnæus says little and adds no more in his later works. It is defined
corollis stamine longioribus, and two synonyms are quoted, one from Royen,
the other from Tournefort, but the latter with a note of interrogation. The
habitat “in Europa australi ” is too indefinite to give any clue ; the definition
|
3
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f
:
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f
SOME CRITICAL SPECIES OF ECHIUM. 415
is not really applicable to any European species except Æ. parviflorum,
Moench, E. arenarium, Guss., and the small-flowered form of Æ. vulgare
known as Æ. Wierzbickii. These are all out of the question and have never
been suggested as the species intended by Linnæus; we are therefore
dependent on the synonyms, and on the specimen that exists in the herbarium,
to ascertain what Linnæus meant.
Both synonyms and specimen belong to the very remarkable perennial
Echium from Portugal and north-western Spain that has been named by
Sampaio * K. Broteri, because it is the Echium italicum of Brotero, non Linn.,
=F. italicum var. lusitanicum, Hotfmg. et Link. I have flowered at Selham
in Sussex in 1918 several plants of this chium from seed sent by Prof.
Coutinho of Lisbon. In good soil from ten to twenty ascending flowering
stems issue from beneath the great central rosette of leaves, which remains
flat on the ground. The inflorescence of each of these lateral stems, when
normally developed is not unlike that of E. vulgare, but if the growth of their
axis has been ehecked from any cause the upper cincinni when fully unfurled
in fruit appear almost corymbose, as is the case in the Linnean specimen.
Brotero's description of his Æ. italicum in Fl. Lus. i. p. 290 (1804), is very
full. Link in Hoffmg. & Link, Fl. Port. i. p. 185 (1809), calls it * accurata."
Other characterisations may be read in Coutinho, as Boraginaceas de
Portugal in Bol. Soc. Broter. xxi. p. 113 (1905), in the same author's Fl.
Port. p. 499 (1913), and in Merino, Fl. de Galicia, ii. p. 155 (1906).
Coutinho gives characters to distinguish the speeies from the nearly allied
E. salmanticum, Lag.= E. polycaulon, Boiss. Brotero says: * Caules quinque
ad duodecim ex eadem radice, sub rosula foliorum radicalim, primum obliqui,
dein ineurvati, erectiusculi, bipedales et ultra. . .. folia radicalia in orbem
prostrata, pedalia et longiora, ad medium biuncialia aut latiora, lanceolata ....
spice ad quinquaginta et ultra, spiraliter reeurvie, fere ex imo caule usque
ad ejus apicem." The habit of Æ. salmanticum is thus described by J. Gay
in a MS. note on plants grown in the Jardin des Plantes : * caulibus ex una
radice pluribus, rosule centrali subjectis . . . . Il forme une énorme touffe de
2-3 pieds haut.” This character in Æ. salmanticum led Gay to suppose that
it was the plant intended by Brotero, and in his MS. notes he proposed the
name /. Broterianum for it, a misnomer fortunately not published. Gay also
knew the true Æ. Broteri, which he regarded as Æ. lusitanicum, and indicated
the distinctions. The corolle are quite different in the two species.
The habitat known to Brotero was “in sabulosis ad Villa Franca, ad Mun-
dam prope Conimbricam (Coimbra) et alibi in Beira.” Hoffmg. & Link say
* contrées élevées du Portugal, entre Caldas de Gerez et Portela do Homem
en abondance. De méme dans la Serra d'Estrella, d’ou la rivière du Mondego
la transporte dans les sites inférieurs prés de Coimbra, ete." It was collected
* [n herb. Acad, Polyt. Brot. et in Lusitano, 12 Jan. 1900, ex Coutinho in Bol. Soc. Brot.
xxi, p. 118 (1904).
416 MR. C. C. LACAITA: A REVISION OF
in Galieia by Pourret in 1802 and has since been found there by Merino in
three places in the frontier province of Orense.
The five modern specimens from Portugal I have been able to inspect
are :—
1. Herb. Hort. Bot. Conimbr. (in Herb. Kew), no. 470. Coimbra,
1878. A very imperfect example.
2. Herb. Murray (in Mus. Brit.), Caldas de Gerez, leg. R. P. Murray,
16. vi. 1887.
. Herb. Murray (in Mus. Brit.), Gerez, leg. R. P. Murray, vi. 1888.
4. A specimen kindly sent me by Prof. Coutinho, labelled : ** Echium
Broteri, Samp. In Transtagana montana, circa Marvao prope
Beiram Majo, 1913.” This is accompanied by some separate
fruiting spikes.
5. A fruiting specimen given me by Prof. Henriquez of Coimbra.
TI
All the examples in flower have remarkably protruding stamens. It must
therefore be admitted at once that the character “ corollis stamine longioribus "
is a grave ditliculty, and it will be returned to below.
To come to the synonyms :—-
A. What is X. amplissimo folio Lusitanicum, Tourn.? There is no speci-
men so labelled in Tournefort’s herbarium, but no. 587 without any original
label is undoubtedly the plant to which he gave that name, and was ticketed
by J. Gay in 1834 “ Verosimiliter X. ampliss. folio lusit. ; specimen minus
evolutum = Ech. lusitanicum Linn. Poir. ete.” I have examined this specimen;
it is precisely Æ. Broteri. Then in Herb. Mus. Paris there is a cover con-
taining two pieces, one in flower, the other in fruit, ex herb. Vaillant,
labelled by him ** E. ampliss. fol. eine and subsequently by Desfontaines
* Ech. lusitanicum Linn. ex synon.” Both pieces are Æ. Broter’. Herb.
Jussieu contains three specimens that are W. Broteri; two of them under
no. 6605, of which one from herb, Isnard is labelled ** Æ. ampliss. folio lusit.;
E. lusitanicum folio cubitali T. ; the other from herb. Charles “ Æ. ampliss.
folio lusit.” ; the third under no. 6606 is labelled “ no. 48 Ech. undulatum
Pourr. Gallice ” (sc. Galicia) “ envoyé par M. Pourret 1802.”
Herb. Gay at Kew contains a similar specimen given him by A. de Jussieu.
As Gay's note confirms the above determinations of the specimens at Paris it
is here transcribed :—
s Echium lusitanicum L. (ex. herb. Juss.) = 4. amplissimo folio Lusitanicum
Tourn. Inst. Herb. Isn.! Specimen falso nomine inscriptum in herb.
Juss. verisimiliter cultum. Adr. de Juss, dedit Xbr. 1834. £. amplis-
simo folio Lusitanicum Tourn. etiam in herb. Vaill. ... Z. undulatum
Pourr. in herb. Juss. (anno 1802 e Gallecia hispanica a Pourretio
missum) hue spectare videtur, specimen vero valde imperfectum. non
nisi inferiorem caulis partem sistit. Deest in herb. Linn. [quoting
SOME CRITICAL SPECIES OF ECHIUM. 417
Lehmann]. Exstat in herb. Tournef. sine nomine et loco. Racemi in
specimine Tournefort minus quam in meo evoluti, dimidio breviores."
Then on another slip “ Xbre. 1834. Fragment d'un échantillon conservé
dans l'herbier de Tournefort * sans nom et sans localité, que je suppose
appartenir à l X. lusitanicum de Linné. Cette méme plante se trouve
dans Vherb. de Vaillant ! sous le nom de Leh. amplissimo folio Lusit.
Tourn. Inst. = /7. tingitanum altissim. flore variegato I 0x: 3.440; no; 5.
La plante de ces deux herbiers a les épis de moitié plus courts qu'il ne
sont dans Pherbier de M. de Jussieu, mais cela tient probablement à
Page des échantillons, moins avancés, moins développés dans l'herbier de
Tournefort et de Vaillant.”
It is thus quite certain that Tournefort’s synonym refers to /. Broteri,
Samp. There are several old specimens agreeing with Æ. Broteri at the
British Museum, which bear out this conclusion and moreover indicate that
the plant was in cultivation in the 18th century. I wiil quote :—
(1) Herb. Sloane, vol. 166, p. 109, among Petiver's Hort. Sice. Hispanic etc., comm.
Salvador and labelled “ E. amplissimo folio Lusitanicum Tourn.” Though
Salvador lived at Barcelona, it is clear from other Portuguese species sent by
him to Petiver that he collected in Portugal.
(2) Herb. Sloane, vol. 198, p. 48, among Petivers Pl. Rayan, also labelled
E. amplissimo, etc.
(3) Hort. Cliff. labelled by the first hand Echium sylvestre hirsutum maculatum,
C. B. P., which words have been cancelled (by the same hand ?) and replaced
by creticum angustifolium rubrum. The unknown second hand added creticum.
This specimen is an instance of the utter untrustworthiness of the names
written in Herb. Hort. Cliff.
(4) Herb. Gronovii, labelled E. folio amplissimo etc., Town., then E. corollis stamine
longioribus, but the last phrase has been cancelled and replaced by Æ. caule
erecto piloso etc., the Linnean diagnosis cf E. italicum. This alteration points to
an observer who had noticed the agreement of the specimen with Æ. italicum,
Brot., but overlooked the fact that Brotero's is not Linn:us's italicum.
(5) Hort. Chelsea. A fine specimen labelled by Solander first ? italicum and then
Echium lusitanicum, L. M.
B. Royen’s synonym, * Æ. caule simplici, foliis caulinis lanceolatis. sericeis,
floribus spicatis lateralibus, Lugdb. p. 407," is uselessly indefinite in itself,
but Lehmann, Asperif. p. 452, under X. lusitanicum (which to him meant
E. Broteri) observes, “ asservantur in herbario Vahliano exemplaria ab ipso
Royeni Museo a Brugmanno data.” It is therefore clear that this synonym
also refers to Æ. Broteri.
The Linnean specimen bears no writing in the hand of Linn:eus himself,
but an old label in that of Jan Burman saying * chium lusitanicum folio
amplissimo Tourn." Now Burman’s entire herbarium was brought by his
* Unfortunately this fragment is missing at Kew, perhaps lost when the plants were
mounted,
418 MR. C. €. LACAITA : A REVISION OF
son to Linneus in 1760. Therefore Linnzeus cannot have seen this specimen
when he wrote the diagnosis of lusitanicum for the Sp. Pl. of 1753, but that
he saw it afterwards is proved by the name lusitanicum being underscored in
his copy of the twelfth edition of the ‘Systema Nature’ of 1767*. The
name lusitanicum now on the sheet was written by Linn. fil., who ought to
have known what his father meant, notwithstanding that Sir J. E. Smith has
pencilled a note of interrogation after the name. The specimen is unmis-
takably /. Broteri. It consists of two leaves measuring respectively 45 x 8
and 30x 14 cm., covered with rather sparse soft hairs, which on the upper
surface only are tubercle-borne. There are no stem-leayes. ‘here is also
the upper part of a fruiting stem, forming a panicle 15 x 14 em., composed
of six cincinni, erecto-patent and very much elongated in the fruiting stage,
the uppermost springing close together so as to give to the whole a false
appearance of being corymbose. Bach cincinnus is 10-15 em. long, with
30-40 or more small, almost sessile flowers. The bracts are lanceolate, equal
to or shorter than the calyces, of which the teeth are lanceolate, acute, 5-7 mm.
long. The whole inflorescence is rough with white setaceous hairs on white
tubercles. Very few corollas remain, but they have evidently been small,
subregular, and only slightly dilated at the throat. They are too much
withered for the character and length of the stamens to be defined. Owing
to the advanced stage in which it has been gathered the elongated, erecto-
patent cincinni look very unlike those of examples in an early flowering
stage. But this change of superficial appearance occurs in most species.
Are we then to decide that Æ. Broteri is the species indicated by Linnzeus
under the name lusitanicum, which therefore must be used for it? Surely
we must, seeing that this is the plant referred to in the two synonyms and
represented by the fine specimen in the herbarium, notwithstanding :—
(a) the name on the specimen not being in Linnzeus’s own hand,
(^) the phrase “ corollis stamine longioribus,"
(c) the baseless opinion of Brotero that by lusitanicum Linnzus meant
a form of vulgare,
(d) the opinion of Link, based on a mistake, that the lusitanicum of
Linnzus was £F. plantagineum,
(e) the erroneous statement of Link and Lehmann, uncritically accepted
by later authors, that Linnzeus’s herbarium contains no specimen of
E. lusitanicum.
The objections shall be taken in order :—
(a) That Linnæus did not himself write the name lusitanicum on the sheet
with the specimen may have been due to his old doubt as to the identity of
Royen’s and Tournefort’s synonyms, indicated by the query appended to the
* See Dr, Daydon Jackson’s ‘Index to the Linnean Herbarium,’ p, 8, as to Linnæus’s use
of underscoring.
eU ET
SOME CRITICAL SPECIES OF ECHIUM. 419
latter in the ‘Species Plantarum.’ I think we may be fairly sure that when
he wrote that work he had not seen any specimens that he knew to be
Tournefort's Kehium amplissimo folio lusitanicum, for the example in Hort.
Cliff. was unrecognised by him. Why he did not further deal with the
species after he had Burman's specimen it is useless to speculate. But in
all the circumstances we certainly cannot take the absence of his autograph as
any indication that he thought the specimen was not his lusitanicum.
(b) This phrase is the only serious objection to the identification of
Ai, lusitanicum with K. Broteri, the stamens of the latter being exserted,
especially the two lowest, as noticed by Brotero. But in the specimen the
very few remaining corollas are in such a state that the incompatibility of
the phrase with the plant might be overlooked. Yet we cannot rely on such
an explanation, which would throw no light on the reason for the original use
of the phrase in Sp. Pl. We shall, I think, be justified in assuming either
that longioribus was a slip of the pen or of an amanuensis for brevioribus, or
that it was a mistake due to some misunderstanding that cannot now be
traced. “Quandoque bonus dormitat Homerus."
(c) Brotero says under X. vulgare, * Echium lusitanicum L. quod passim
occurrit, est varietas staminibus corolla vix brevioribus." How could Brotero
know this with nothing but Linnzeus's brief phrase to go upon? He could
not be referring to the Wierzhickii form of vulgare, quia passim non occurrit,
least of all in Portugal. Indeed, the vulgare of Brotero is supposed by all
later authors to be tuberculatum, Hoffmg. et Link.
(d) and (e) Almost simultaneously Link took the lusitanicum of Linnwus
to mean Æ. plantagineum. His suggestion and that of Brotero are equally
untenable and mutually destructive. In 1804, the very year of publication
of the Fl. Lus., Link sent to Smith two little flowering scraps, each about
24 inches long, of X. plantagineum with this ticket :—
ES
| : i
2, Echium lusitanicum ? Ad plantagineum quoque bene convenit. F...* in Lusit.
| i
These scraps may be seen on the sheet of X. plantagineum in Herb. Smith,
who has only written against them “ Portugal, Prof. Link 1804.”
No doubt these fragments are what induced Smith to insert a query after
Linn. fil.’s determination of specimen M as Æ. lusitanicum. We can only
guess that some letter of his acknowledging them must have been misunder-
stood as meaning that in Herb. Linn. there is no specimen of Æ. lusitanicum.
Indeed, Link, in Fl. Port. ii. p. 186, went.so far as to identify not only
E. lusitanicum, Mill. (which is identical with the Linnean lusitanicum), but
even Tournefort's X. amplissimo folio Lusitanieum with £F. plantagineum,
which is manifestly absurd. He then-goes on “ Æ. lusitanicum in herbario
* I cannot read the two letters after F. ; possibly meaning “Frequens.”
120 MR. C. C. LACAITA : A REVISION OF
Linneano non exstat. — eium lusitanicum Milleri et synonymon Tournefortii
hujus loci sunt fide herbariorum." (Exactly contrary to the fact.) “ Expur-
gatur itaque Æ. lusitanicum ex Systemate. Nostram plantam esse X. planta-
gineum Linn. litteris confermavit cl. Smith." Yes; Smith's letter may
indeed have confirmed that Link’s specimen was Æ. plantagineum, but that
has no bearing on the identity of X. lusitanicum, L.!! Tt is also possible
that some specimen of Boerhaave's may have led Link or Smith to fancy that
E. lusitanicum was identical with plantagineum. Among Boerhaave’s plants
in Herb. Sloane, vol. 321, p. 63, there is a piece of plantagineum labelled
“ Anchusa lusitanica non descripta.” — Link's erroneous statement * ££, lusi-
tanieum in herbario Linneano non exstat" has been handed down to later
authors and never corrected until now. Even Lehmann says * Æ. lusitanicum
deest in herbario Linneano, quod Smith in litteris," and the error has been
accepted by de Coincy in a MS. note to the specimen of E. polycaulon in his
herbarium, and by Coutinho in Borag. Port. l. c. (suprà, p. 415).
If the arguments I have adduced are not considered sufficiently convincing
to justify the use of the name lusitanicum, L., in the sense of E. Broteri, Samp.,
the only alternative will be to make use of Sampaio's appellation and. to
reject altogether the Linnean name as a nomen confusum, for there is no
other meaning that can be assigned to it.
(IV.) ECHIUM PLANTAGINEUM is well represented by specimens C, D, E,
which are quite typical. The mark * A 39 " on specimen D indicates that it
came from one of Linnzus’s correspondents whose name began with A,
perhaps Alströmer, who collected in lands where Æ. plantagineum is plentiful.
Smith's query is accounted for by a specimen from Hort. Paris in Herb.
Banks, now at Mus. Brit., which though certainly plantagineum was wrongly
labelled orientale by Solander. Moris, who inspected the Linnean herbarium,
says of this specimen in Fl. Sard. iii. p. 128 * FE. orientale H. B. sie in Linn. !
herbario, non a Linno, aliena manu inscriptum, pertinet ad Æ. plantagineum.”
Specimen E was collected by Lófling, as indicated by the mark “ L. 152,"
and therefore came from Spain or Portugal.
This is the only Linnean species besides E. vulgare that cannot give
rise to any doubts. The description of the bracts and of the radical leaves
and the reference to Barrelier, ic. 1026, are sufficient to establish its identity.
Among other distinctive characters it possesses two that are particularly
useful, because easily verified in incomplete herbarium specimens and not
found combined in any other species. (1) The indumentum of the leaves is
not dimorphous or composed, as in most kinds, of a carpet of short softish
hairs above which longer scattered bristles project, but of bristly hairs alone.
more or less tuberculate at the base according to the age and habitat of the
plant. (2) The corolla is of a peculiar thin texture, not externally pubescent
or velvety all over, but glabrous, with hairs on the nerves only and more or
SOME CRITICAL SPECIES OF ECHIUM. 421
less ciliate on the margin. The colour in life is a varying compound of blue
and light purple, which generally dries blue but sometimes shows the purple
tint. The corollas of Æ. maritimum, Willd.= E. confusum, Coiney, which by
some authors has been carelessly thought to be a variety of plantagineum, are
smaller, pubescent all over externally, and Prussian or indigo blue in dried
specimens *,
The failure of pre-Linnean botanists to recognise Æ. plantagineum and its
subsequent confusion with Æ. violaceum have caused much trouble. The
rather wide intervals between the fruiting calyces have encouraged confusion
with Æ. eretieum, and the purplish corollas have led to the misappropriation of
the name violaceum to large flowered forms of plantagineum. Examples from
the Atlantic islands often have exceptionally large flowers—e. g., Bourgeau,
Pl. Canarienses anno 1845 no. 235, from Teneriffe, with corollas fully 3 em.
long and very wide-throated. Similar plants in Herb. Kew are all labelled
violaceum by C. B. Clarke; these should perhaps be recognised as a variety,
but not under that name.
E. grandiflorum, Dest., is very distinct in respect of every organ. Its
reduction to a variety of plantagineum, accepted even by so good a judge as
Ball (Journ. Linn. Soc. xvi. p. 575), is a reductio ad absurdum. A beautiful
species from Palestine, X. judwæum mihi, closely allied to grandiflorum, is
wrongly determined as plantagineum (or violaceum in the same sense) in all
the exsiccata I have seen: for instance, from the Haram Court at Jerusalem,
leg. M. J. Fox, Feb. 1867; from Jericho in Bornmüller's Tt. Syr. 1897,
no. 1136, and from the same place in Meyers and Dinsmore, Pl. Palæst.
Maris Mortui. True plantagineum is also found in Palestine: e. g. PI.
Jordanice ex Herb. Postian. no. 526 from Sarada and no. 527 from Wady el
Kefar ; also from Antioch, leg. Loftus no. 71.
E. plantagineum is the species which has spread most widely in regions
remote from its natural home. It is represented from the Cape of Good
Hope, from Australia, where it is plentiful in grass lands round Adelaide,
from the Eastern U.S.A., from Southern Brazil (leg. Chamisso), from
Montevideo, and from Buenos Aires. Poiret gave the name of Æ. bonariense
to the Buenos Aires plant,though it is just typical plantagineum.
But /. plantagineum is by no means uniform. Its complete specific isola-
tion should encourage a study of its forms. The radical leaves do not always
resemble those of Plantago major. Jacquin, Hort. Vind. t. 45 (1770), figures
a narrow-leaved form as * Æ. plantaginei Linn. an varietas ?" which is quoted
by Murray in Syst. Veg. ed. xiv. p. 189 as * varietas foliis radicalibus lanceolatis
longe petiolatis Jacq. t. 45." M. Battandier tells me that in Algeria
plantagineum hybridises with grandiflorum. This is a matter for further
* The annulus of the tube is quite different ; plantagineum belongs to De Coincy’s sect.
Lleuthervlepis, maritimum oscillates between Eleutherolepis and Gamolepis.
422 MR. C. C. LACAITA : A REVISION OF
investigation. So far I have not seen specimens that support this statement,
but it can only be definitely confirmed in the field.
to
In the old herbaria I have found sundry specimens undoubtedly referable
plantagineum, but bearing strange determinations, the species having
remained unrecognised until Linneus’s * Mantissa? was published :—
(1) In Herb. Morison (at Oxford) labelled “ E. eretieum latifolium rubrum
C. B. P. E. ereticum Clus.”
(2) Herb. Sloane, vol. 168, p. 12, among Banister’s collection : “4. ereti-
cum P. Alpini."
(3) Herb. Sloane, vol. 321, p. 63, among Boerhaave's specimens are two
pieces of plantagineum beside each other, labelled respectively
* Lycopsis C. B. 255. Altera anglica Lob. 684. — Echii altera species
Dod. 904 " and * Anchusa Lusitanica non descripta ab expertissimo
Dom. C. .. . (illegible) apud. quem floruit anno 1686 in Viridario
Lusitanico."
(4) Hort. Cliff. labelled by the first hand Æ. folio asperrimo et verrucoso
and by the second hand Æ. vulgare.
(5) Hort. Cliff. labelled by the first hand Æ. creticum latifolium rubrum
and by the second /. eretieum. j
(6) Hort. Cliff. labelled by the first hand 77. eretieum angustifolium rubrum
(although identical with no. 5) and by the second. eretieum.
(7) Herb. Pallas, labelled X. violaceum.
(8) Herb. Helvet. in Hb. Dick, labelled by Solander X. violaceum, L.
(9) Herb. Miller (probably), labelled in Mi.ler's hand * Æ. ereticum caly-
cibus fructescentibus remotis H. U. 85; 7. creticum latifolium rubrum
C, B. P." and by Solander Æ. ereticum. (The smaller piece on this
sheet, marked H. L. S. = Hortus Linn:ei siccus, is not plantagineum
but really Æ. ereticum, herb. Linn.)
(10) Herb. Heg. Paris, 1827, originally labelled J£. maritimum ins.
Stoechadum jl. Maximo Tourn. Inst. 136 (which is represented in
Hb. Tournef. No. 591 by a specimen of Æ. grandiflorum, Dest.),
but by Miller Æ. caule erecto hispido, foliis lin.-lane. hispido-pilosis,
semiampleaicaulibus, floribus spicatis terminalibus, and by Solander
E. orientale. This is the specimen which explains Smith's query
on the Linnean specimen D. Of course, it has nothing to do with
E. orientale, L.
Al these specimens except the first are in the British Museum.
osi. s
SOME CRITICAL SPECIES OF ECHIUM. 423
(V.) ECHIUM RUBRUM.
The three specimens J, K, L are in good preservation and quite typical of
Jaequin's species. The mark on J indicates a plant sent by Gerber, who
collected in South Russia in the area of Æ. rubrum. The unusual style, not
deeply cloven into two filiform branches as in all other European species, but
ending in a elubbed stigma, is conspicuous. The corollas still preserve the
dark red colour, as of dry blood, that is peculiar to Æ. rubrum. The species
was well known to Clusius, who has a good figure of it as /Zchium rubro flore
in Rar. Stirp. Pann. p. 681 (1583) and in Hist. ii. p. 164 (1590), with a fair
description, assigning as habitat the neighbourhood of Sopronium (Oeden-
burg) in Hungary, which is also Jacquin’s locus classicus. Unfortunately,
Clusius's name is quoted by Linnzus under Æ. violaceum, but Echium rubro
flore, Ray Hist. p. 499, identical with Clusius's plant, is given as a synonym
of E. ereticum. To the confusion of mind underlying this double false
reference may be attributed the absence of any determination by Linnæus of
his three specimens.
(VI.) ECHIUM VIOLACEUM.
Echium violaceum, L. Mant. p. 42, has most frequently been misinterpreted,
especially by British botanists, as a large-flowered form of Æ. plantagineum.
Individuals of the latter species are often determined as violaceum ; many
such with large corollas, from the Atlantic islands, having been so labelled
by C. B. Clarke in Herb. Kew. On the other hand, Grenier, in Fl. Fr. ii.
p. 524, argued that Linnzeus meant Æ. rubrum, Jacq., but the diagnosis and
the specimen in the herbarium prove both suppositions to be wrong. Moris's
statement in Fl. Sard. iii. p. 129 that Æ. pustulatum, Sibth. & Sm., was meant,
“ E, violaceum H. U. sic in Linnsei ! herbario, ramus absque foliis radicalibus,
Echium pustulatum nostram maxime refert, Ab Æ. plantagineo certe
alienum,” is nearer the mark, but I shall give reasons for thinking that the
specimen is not true /7. pustulatum, Sibth. & Sm.
Are we then to substitute /. violaceum, L., as the earlier name, for that of
whatever species this specimen may really belong to? Certainly not ;
because even if the identity of the specimen were indisputable, the name
violaceum is a confusion of the plant represented thereby with Æ. rubrum, Jacq.,
and ought therefore to be rejected as a nomen confusum in conformity with
Art. 51, clause 4, of the international rules. To do otherwise would, in view
of the continual and varying misuse of the name, only perpetuate error. It
is therefore best to discard the name once and for all.
The confusion by Linnæus of Æ. rubrum with his herbarium plant from
Hort. Ups. will become obvious if we analyse what he says in the * Mantissa.’
He begins (1) with a diagnosis: * Echium corollis stamina :equantibus ;
424 MR. C. C. LACAITA : A REVISION OF
hirsutum maculatum, Bauh. Pin. 254, and (3) E. rubro flore, Clus. Hist. ii.
p.164. Then comes (4) * Habitat in Austria." And there follows an Obs.
with five remarks: (5) "simillimum Æ. vulgari sed corolla violacea " ;
(6) “ caulis magis diffusus " ; (7) “stamina corolla non longiora, purpurea,
tamen stylo albo piloso”; (8) “corolle tubus longitudine fere calycis " ;
(9) “facies distincta.” And (10) a statement that the plant described was
grown in Hort. Ups. Of the above elements 2, 3, and 4 refer to X, rubrum
and to X. rubrum only; the remaining seven apply to the herbarium
specimen.
No. 2 (the first synonym), wrongly ascribed by Linnæus to C. Bauhin's
‘ Pinax, p. 254 (1623), is not found there, but in his * Phytopinax,' p. 490
(1596), as “ Echium sylvestre secundum hirsutum maculatum = E. rubro flore
Clus. Pann. p. 682.” It is therefore identical with the second synonym,
which is found in Clusius, Rar. Stirp. Pann. p. 681 (1583), and in his Hist,
Pl. ii. p. 164 (1590), with an excellent figure that can represent nothing but
E, rubrum, Jacq., and a fairly good account of the plant, stated to come from
Sopronium (Oedenburg) in Hungary. The “habitat in Austria” therefore
agrees with Æ. rubrum, but not with the specimen, which does not resemble
any species found in Austria or Hungary *.
Nevertheless, it is impossible to identify the violaceum of Linnæus with
I’. rubrum, because: (1) the herbarium specimen is not rubrum; (2) the
herbarium contains three excellent specimens of Æ. rubrum, though unnamed
and unrecognised by Linnzeus (sce note 7 on specimens J, K, L); (3) E. rubro
Jlore, Ray Hist. i. p. 499, which is identical with Æ. rubro flore, Clus., is not
quoted here but under Æ. eretieum, showing that Linnæus had no clear or
consistent idea of Æ. rubro flore; (4) the diagnosis and the characters
indicated in the Obs, are incompatible with Æ. rubrum. The specimen is not
rubrum, because it has a bifid style and is unlike that species in every
character. The identification with Æ. plantagineum or a large-flowered
variety of that species is obviously incompatible with the synonyms and
habitat assigned ; it must have been based by those who adopted it on the
descriptive elements, which at first sight might be thought applicable. But
Linneeus tells us that he was describing the Hort. Ups. plant represented in
the herbarium by our specimen, which is not plantagineum on account of the
very different indumentum, and the pubescent corollas that also differ in
texture and shape. The specimen is not the typical Æ. pustulatum of
tubo calycibus breviore." Then follow two synonyms: (2) K. sylvestre
* With regard to the violaceum of the German authors, it is to be noted that, as alread y
remarked by Grenier, “T'E, violaceum des auteurs allemands pas plus que leur plantagineum
ne croissent en Autriche, mais seulement en Istrie et en Dalmatie,” which in the time of
Linneeus did not form part of the Austrian territories, but were still subject-lands of
Venice,
SOME CRITICAL SPECIES OF ECHIUM, 435
Sibthorp * from Sieily because the indumentum is different, being even more
tubercular, but less hispid ; this difference, however, may be due to cultiva-
tion; the calyces and bracts also look somewhat different.
Unfortunately the specimen in question only shows the upper part of
a plant with a panicle about 30 em. long by 10 em. broad, the branches in the
lower axils, whose flowers are still in bud, measuring 5-6 em. The arrange-
ment of the flowers almost recalls that of X. rosulatum, Lange, the cincinni
being few-flowered and scattered and very obscurely scorpoid or hardly
furled. The indumentum of the stem is dimorphous, but that of the leaves
is peculiar and similar to cultivated examples of rosulatum. There is prac-
tically none of the soft pubescence which is present in so many species ; all
the hairs are bristles, larger or smaller, and the whole leaf very scabrous, not
hispid, with regularly disposed sete as in plantagineum, but with numerous
minute tubercles scattered over both surfaces, some quite bald, some bearing
inconspicuous bristly hairs. At intervals among these are larger tubercles
occasionally crowned by stronger thick bristles, but in most cases these have
been worn off or never developed, so that most of the pearly pustules, which
are very conspicuous, look naked. The uppermost stem-leaves are lanceolate
and pass insensibly into the lower bracts (which alone are visible) of the
same shape. These are exceptionally long and foliaceous, equalling or
exceeding the lower flowers. The calyx-segments are very long and
acuminate, the corollas about 2 em. long; rather less oblique than is usual
in Æ. pustulatum and not so wide at the throat as in plantagineum, nor have
they the peculiar thin texture of that species. They are pubescent all over,
not glabrous with hairs along the nerves and ciliate as in plantagineum.
They have dried blue with a paler pinkish throat. The style is hairy in its
whole length and bifid to a depth of not more than 2mm. The stamens are
sub-exsert, 7. e., rather longer than the lower but shorter than the upper lobes
of the corolla. Without dissection, which is not permissible, one cannot be
sure whether the filaments are quite glabrous. I suspect that this garden
plant originated from one of the Kehia of the Spanish peninsula, but it is
difficult to say from which.
In the old herbaria at the British Museum I have found sundry specimens
more or less like that of Linnzus; all I think garden forms, in which
cultivation has reduced the hairy coat, but not the pustules, so that the leaves
sometimes look as though they were studded with pearls. Some of these
show a peculiar lax inflorescence and diffuse habit, which may point to
I. rosulatum, Lange, as their origin. Nor can we be sure that some may
not be natural garden hybrids. None have the true radical tuft of leaves
preserved :—
* See my note on the Echia of Sibthorp's herbarium, supra, p. 386.
426 MR. C. C. LACAITA : A REVISION OF
-Ó
Herb. Sloane, vol. 306, p. 156, among Uvedale’s Plante Rayan,
as “E. hispanicum verrucosum angustifolium, Dni. Vaillant; an
FE. foliis angustis villosis Tourn." I have not detected hairs on the
filaments of this specimen.
Herb. Sherard (at Oxford); from Dr. Dillenius as “ Echium
perlatum. Vaill."
Hort. Cliff., labelled by the first hand “ E. foliis angustis et villosis
Tourn.,” but by the second “fruticosum,” which is then erased and
altered to “argenteum,” both the latter names belonging to South
nz
A^
African species now referred to Lobostemon.
Herb. Gronov., labelled “ Fehium foliis angustis et villosis Tourn. 136.
Anchusa angustis villosis foliis, Boce. Mus. 2. 84, t. 78; E. perlatum
a transcript
rs
angustifolium Hispanicum quorundam." This label is
of Boerhaave, Ind. alt. Pl. p. 194 (1720).
Herb. Miller, labelled by him * Æ. caule ramoso aspero, foliis calloso-
verrucosis, staminibus corolla longioribus ; Echium ereticum C. B. T
then by Solander first “orientale” and afterwards “ angustifolium
Mill. Dict. no. 6."
Although the labels of the above specimens may point to Æ. angustifolium,
Lam., they do not belong to that remarkable species, which has nothing to
do with angustifolium, Mill. They are good instances of the wild shots of the
or
old botanists at the names of their garden plants.
6. Hort. Chelsea, no. 2067 of 1763, labelled * Æ. corollis stamine
longioribus L. Sp. Pl.” (which is the diagnosis of Æ. lusitanicum)
and “ Æ. amplissimo folio lusitanicum Tourn.”
7. Hort. Chelsea, labelled by Solander ** Æ. ereticum," on the same sheet
as a piece of plantagineum, labelled by him * orentale."
8. Herb. Smith, labelled “ Æ. genevense Fairb.; FE. creticum? Own
garden, Marlboro’ St. 1788.
Tournefort's herbarium unfortunately does not contain a specimen of his
E. foliis angustis et villosis, nor have I been abie to determine precisely the
plants bearing that title in herb. Jussieu, no. 6621. No. 6622 ** E. Roris
marini folio Tourn. 136” is very like some of the old Mus. Brit. examples.
But the difficulty of an exact determination of the violaceum of herb. Linn.
is immaterial, for, whatever it be, it conflicts so absolutely with the diagnosis
and synonyms that the name must certainly be abandoned.
(VII.) ECHIUM VULGARE, 5p. Pl. p. 139 (1753), had already been men-
tioned in Hort. Cliff. p. 43 (1737), as X. caule simplici, foliis caulinis lanceo-
latis, floribus spicatis ex alis," and again in Fl. Suec. p. 56 (1745), as found
in Sweden, with the words of the diagnosis very slightly altered. In Hort,
SOME CRITICAL SPECIES OF ECHIUM. 427
Cliff. there is the observation, * varietates hujus fere tot quot individua,"
and the desperate suggestion that even the two following species, which are
ereticum, rubrum, angustifolium and orientale, may not be specifically different.
The specimen is the upper part of a plant, with the inflorescence 24 em. long
by only 5 em. broad, composed of over twenty simple, very scorpioid, axillary
cincinni, none of which in flower exceed 25 mm. in length. The stamens all
protrude beyond the longest part of the corolla, and have glabrous filaments.
No habitat is marked, but the specimen is exactly the W. vulgare common in
England.
To include in Æ. vulgare, L., the rather numerous forms which in southern
Europe replace this plant of Scandinavia, England, and Central Europe, and
differ from each other as much as from typical vulgare, is surely putting a
great strain on the Linnean name. (See my paper on the Lehia of Sibthorp’s
herbarium as to the relation of some of these forms to true vulgare.)
(VIII) SPECIMEN G.
This is a young specimen of one of the Spanish forms of the vulgare-
pustulatum-gaditanum group, but it is too scanty to determine more closely.
It is a very small erect plant, not over 15°5 em. high, with twofold
indumentum and smallish blue pubescent corollas, about 2 cm. long, with
style and the two longest stamens only protruding, equalling the upper
corolla-lobes.
The mark, “ 152 «" seeing that this sheet is pinned to one marked
* L. 152," indicates almost certainly that this specimen also was collected by
Lófling. I have not been able to trace the phrase, * Æ. montanum parvum
flore magno," in the old books, nor to ascertain what locality is meant by
“in mont. Espartal," although I find Cape Spartel on the African coast
spelt “ Espartel" in an old Spanish gazetteer. There is no evidence that
Loefling ever visited the African side of the Strait of Gibraltar, but if this
specimen really came from that region other possibilities as to its identity
are opened up.
V.
THE ECHIA OF MILLENS ‘GARDENER’S DICTIONARY,’
Of the seven species of chium described in Miller's eighth edition of 1768
all but the seventh give rise to troublesome questions. This innocent no. 7,
Echium fruticosum, L., is a South African Lobostemon, L. fruticosus, Buek,
of which no more need be said here. To take the other six in order : the
first two are named by Miller Echium anglicum and Echium vulgare. The
first, Æ. anglicum, of which he says * staminibus corollam æquantibus ” is
doubtless Æ. vulgare, L., var. parviflorum, St. Am., Fl. Agen. p. 82 (1821),
the small-flowered pistillate form usually known as Æ. Wierzbickii, Haberl.,
LINN. JOURN,—BOTANY, VOL, XLIV, 2y
428 MR. C. C. LACAITA : A REVISION OF
Cat. Hort. Bot. Pest.* (1827), ex Houy, Fl. Fr. x. p. 306, and in Reichb.
Fl. Exc. p. 336 (1830). It has no claim to rank as a species, or even
as a variety, being merely a sexual modification of E. vulgare occasionally
found mixed with the type.T St. Amans states that both forms of corolla
had beeu found on the same plant by the banks of the Garonne by
M. Chaubard. The varying behaviour of seedlings from the pistillate form
is discussed by Rouy in a footnote. Type-specimens of Ly Wierzbicki,
collected by Dr. Wierzbicki in the Banat, may be seen at Kew and at the
British Museum. They are very exactly figured in Reichb, Fl. Germ.
iab. 1298. fig. 3. Others with less extremely small corollas are at Kew
from St. Amans's locus classicus on the Garonne, from Aurillac in Savoy,
from Tal y Clawdd, near Ruabon, N. Wales, and elsewhere.
But Miller appears to have accidentally transposed the names anglicum
and vulgare. He was, as usual, describing under no. 1 a garden plant— one
not known to him as a British native, for he says “it grows naturally in
Xermany and Austria, from whence I received the seeds.” The only
specimen in his herbarium marked “ angliewm” is precisely the common
E. vulgare, L., with protruding stamens, which therefore represents his no. 2,
not this no. 1. Moreover, the name anglicum is derived from Lobel’s Lycopsis
anglica, which Miller quotes as a synonym of his no. 2, whereas he cites
E. vulgare, C. B. P., for no. 1, though it is quoted by Linnæus for 7. vulgare.
Hudson, Fl. Angl. pp. 69, 70 (1762), had already used the names anglicum
and vulgare, but in an opposite sense from Miller, for his vulgare is defined
“caule simplici erecto, foliis caulinis lanceolatis hispidis, floribus spicatis
lateralibus, staminibus eorollam æquantibus,” which is Miller's diagnosis of
his anglicum, whilst Hudson uses the name anglicum for the common British
plant, and defines it (quoting Lycopsis anglica Lobelit) as “caule simplici
erecto, foliis lanceolato-linearibus hispidis, floribus spicatis lateralibus,
staminibus corolla longioribus," this being the phrase which Miller applies
to his Æ. vulgare. Hudson professes to be quoting both these phrases from
Mill. Dict. Hort. edit. 6, which is not in accordance with fact, for Miller first
used them in his seventh edition of 1759. They do not oceur in the sixth
edition of 1752, where Lehium vulgare alone is named, with no allusion to the
length of the stamens. In ed. 2, p. 83 (1778), Hudson reduces Æ. anglicum
to /. vulgare var. anglicum.
Miller's second species, Æ. vulgare, is precisely JE. vulgare, L., whose
diagnostic phrase, which originally appeared in Hort, Cliff. p. 45 (1737), he
eopies with the addition of the words * staminibus corolla longioribus."
These words and his statement that “this second sort grows naturally upon
* T have not been able to see a copy of this work, and Rouy does not mention the page.
T For E, Wierzbickii, see Coincy in Bull, Herb. Boiss, ser. 2, i. pp. 789-792,
ENTE
SOME CRITICAL SPECIES OF ECHIUM. 429
chalky lands in most parts of England” establish the identity. We therefore
get the following synonymy :—
(1) Echium anglicum, Mill. E. vulgare, Huds. ed. 12 4. vulgare, L.,
var. parviflorum, St. Am.* = E. Wierzbickii, Haberl., and
(2) Echium vulgare, Mill. E. vulgare, L.-— E. anglicum, Huds. ed. 1
=H. vulgare var. anglicum, Huds. ed. 2. Bauhin’s synonym
should, of course, belong to Miller's Æ. vulgare, not to his
7. anglicum, where for some obscure reason he has wrongly
placed it.
Here the matter might end were it not for the unfortunate introduction,
as a synonym, both by Miller and by Hudson, of Lobel's Lycopsis anglica or
Lycopsis altera anglica. This is no doubt what led to Linnzus assigning
Anglia as a habitat of Echium italicum, to which species he very rightly, in
Sp. PL, transferred Lobel's name as var. 8. To trace the origin of the
error as to Lobel’s plant, and the confusion to which it has given rise,
involves a long digression.
Lobel, in his * Stirpium Adversaria nova,’ p. 249 (1570), describes a plant
under the name of Lycopsis, vel Lycapsis degener Anchusa, Æginetæ. Mathioli
Cynoglossum. He tells us in very awkward Latin that it was first shown to
him by Assatius, not far from the shore at Frontignan, and that he after-
wards raised it from seed at London in England. He also states that it has
no real similarity to Anchusa, except in the colour of the root, the hispid stem
and leaves, and the tangled hairiness of the flowers ; and that it has the
flowers of /Zehium fimbriatum f, of a dilute purple colour, with slender stamens.
Now the only Echium growing in the Montpellier neighbourhood to which
this account could apply is Æ. pyrenaicum. Lobel’s plant is figured in his
Stirp. Hist. p. 312 (1576), as “ Lycopsis altera anglica Advers. p. 249
perperam Cynoglossum Matthioli” alongside of ** Kehium sive Buglossum
silvestre.” The figure in question certainly represents Echium pyrenaicum.
The corollas are clearly intended for those of either Æ. italicum or £F. pyre-
naicum, but the broad pyramidal branching is more like pyrenaicum. The
corollas especially, as well as the ramification, forbid the identification with
E. plantagineum, which was made by ©. Bauhin and by Ray, and for other
reasons might be tempting.
The same pair of figures recur in the * Kruydtboeck,’ p. 684 (1581), and in
Ie. Stirp. p. 579, in both editions of 1581 and 1591, and are again found
in Dodoens, Stirp. Hist. p. 620 (1583), under the names of /ehiwm and
chit altera species, and in Gerard, Hist. Pl. p. 802 (1633), as chium vulgare
and Lycopsis anglica. Thus Echii altera species, Dod., is identical with
* Wrongly attributed to Schur, Enum. Pl. 'Trans. p. 470, which is much later (1866).
T I have not ascertained what plant he alludes to as E. fimbriatum.
430 MR. 6. C. LACAITA: A REVISION oF
Lycopsis altera anglica, Lob., which, be it noted, does not mean “another
English Lycopsis,” but “another Lycopsis, the English one.” An unfor-
tunate expression, for that word * Anglica," which only meant that Lobel
had grown the plant in England, misled the later authors [e. g., Parkinson,
Theatr. p. 518 (1640) ], who wrongly assumed that Lycopsis anglica must be
a native English plant. If they had read a few lines farther they would
have found Lobel saying : ** Hæc ab ea quie hic in Anglia via quæ Bathonia
et Bristolia Londinum ducit per messes læta et multa hilari videtur
aspectu, floribus tantum differt; quos non secus foliorum et alarum inter-
stitia, sed in summis caulibus saturatiore purpureo, Anchuse majoris. Ut
prior (se. Anchusa lignosior angustifolia) subrubra linet et nonnihil rubelli
affricat digitis," meaning that his Lycopsis altera. Anglica is not the common
English Vipers Bugloss, which he had admired in the fields near the
Dath Road. i
Caspar Bauhin, in his *Pinax ' (1623), enumerates on p. 254 eleven plants
under the name of /eAiwin (not all belonging to the modern genus), including
E. vulgare as no. 2. Then on p. 255 he has two species of Lycopsis, the
second of which, Lycopsis «qgyptiaea, does not concern us, but to the first,
which is “ Lycopsis " simply, he appends a long list of synonyms, and among
them are Lehii altera species, Dod., Lycopsis vel Lycapsis degener Anchusa
«Eginete, Ad., and Lycopsis altera Anglica, Lob. It is therefore certain that
C. Bauhin considered Lobel’s plant identical with his own ** Lycopsis.” But
his remark at the end of the list of synonyms lets the cat out of the bag, for
it is only intelligible of /. plantagineum and cannot apply to K. pyrenaicum.
It is “ Aliquando caule caret; Monspeliaca flore est dilute purpureo; anglica
saturatioris est purpura." Now E. plantagineum often occurs in a dwarf
form, and the colour of the corollas varies just as stated by C. B., sometimes
a pale reddish purple predominating, particularly in the Ionian Islands
(cf. Fl. Gr. tab. 179), sometimes a full purplish blue which dries blue. We
must therefore bear in mind that Lycopsis anglica, Lob., is E. pyrenaicum,
but Lycopsis or Lycopsis anglica, C. B. P., is chium plantagineum *.
J. Bauhin, Hist. Pl. iii. p. 586 (1561), absurdly says of Lobel’s two
figures, ** ambe nostrum Fchium vulgare referunt,” which they do not. Bis
own two figures are different. They represent /. vulgare and his £. albo
Jlore majus, which is Æ. italicum.
Merrett, Pinax Rer. Nat. Brit. p. 34 (1666), following Parkinson, claims
Lycopsis anglica (of Gerard and therefore of Lobel) as a British plant. He
speaks of Æ. vulgare, Ger., “in muris et locis siccis," and of “ Echium
alterum sive Lycopsis anglica as growing “cum priore," so he regarded this
asa companion form of Jehiwm. Richardson (see below) understood him to
mean var. parviflorum.
is An examination of Bauhin's herbarium at Bale would be desirable to confirm the
above.
SOME CRITICAL SPECIES OF ECHIUM. 431
Ray, Syn. Meth. Stirp. Brit. p. 75 (1690), only quotes Æ. vulgare for
England, “nimis etiam frequens," but in the addenda at p. 238 there occurs
among the plants observed in Jersey by Mr. Sherard, * Lycopsis C. B.
Echii altera. species Dod. p. 680, cujus ic. hane nostram bene repreesentat.
In the sandy grounds near St. Hilary plentifully." This means that Ray
understood Bauhin’s Lycopsis to be the species subsequently known as
Li. plantagineum, for the Jersey plant is well known to be that kind. It is
still abundant at St. Helier’s and elsewhere in the island, see Lester-Garland,
Fl. Jersey, p. 126 (1903). Specimens from Sherard’s locus classicus can be
seen in Herb. Mus. Brit. and are exactly X. plantagineum, which is figured
in E. B. tab. 2798 from the Jersey plant under the name of Æ. violaceum.
But the remark about Dodoens's figure is strange from Hay, who had travelled
where Æ. plantagineum is plentiful. Apparently he was not so well
acquainted with Æ. pyrenaicum, though he had seen Æ. italicum in several
parts of Italy. In the second edition of 1696, p. 119, he speaks of
Lycopsis, C. B., and Lycopsis anglica, Lob., as of two different species, the
former (with /ehii altera species again wrongly brought in) growing in the
isle of Jersey and therefore plantagineum ; but of the latter he says * Lobel
mentions another sort which he calls Lycopsis anglica, to be found plentifully
among the corn on the way between Bristol and London, which no man since
him hath been able to discover ; so that 1 conclude what he observed there
was nothing but the common chium." This conclusion is right, but Ray
misrepresents Lobel, perhaps from memory, as saying just the contrary of
what he really had said.
Then in ed. 3, p. 227 (1724), three species are definitely set up, intended
for X. vulgare, E. plantagineum (from Jersey only), and Æ. vulgare var.
parviflorum, and referred to respectively as follows : 1. X. vulgare, J. B., iii.
586, C. B. P., 254. 2. Lycopsis, C. B. P., repeating all that had been said
in ed. 2 as to this plant occurring in Jersey. 3. Echium alterum sive
Lycopsis altera, Merrett. “ We have two to be met with in the North as
well as in the South; the alterum differs from the vulgare, J. B., chiefly in
the smallness of the flower and being thicker set in the spike. “Tis
probable this may be Lycopsis anglica Lobelii." This observation is due to
Dr. Richardson, many of whose notes were embodied by Dillenius in this
3rd edition.
Meanwhile, in 1699, Morison, Hist. Ox. iii. p. 441, had described Echium
plantagineum as his no. 6 under a new name as follows: ** //chium ramosius
annuum flore suave rubente, Nobis. Lycopsis C. B. P. Echii altera species
Dod. Hee fortasse est eadem planta, quæ a Lobelio in * Adversariis, p. 249,
memoratur; et e seminibus in Insula Cæsarea vulgo Jersie a cl. Botanico
D. D. Sherard collecta, nobis orta est." He therefore did not distinguish
the Lycopsis of Lobel from that of Bauhin.
We may now turn to the old herbaria. That of Sherard, which by the
432 MR. €. C. LACAITÀ : A REVISION OF
kindness of Mr. Druce I have inspected, contains no plant from Jersey ; his
only example of /. plantagineum was sent by Trionfetti and is unnamed.
That of Dillenius, besides two pieces labelled vulyare, has one labelled
* Lycopsis C. B. Pin. In agris Blackheath.” This, though thought by
Sir J. E. Smith to be italicum is nothing but vulgare, as Mr. Druce has
already stated in ‘The Herbaria of Dillenius p. 75. But in Morison's
herbarium there is a specimen of his Sechium ramosum annuum flore suave
rubente, which should be the plant raised from seeds collected by Sherard in
Jersey, and indeed is referred to plantagineum by Vines and Druce in their
account of the Morisonian Herbarium. Unfortunately the specimen on
closer examination turns out not to be plantagineum, as is proved by the
corollas being pubescent all over and by the indumentum. But the next
specimen, labelled originally * chium ereticum latifolium rubrum C. B. P.
and accepted by Vines and Druce as Æ. eretieum, L., is precisely /. planta-
gineum. It looks as if there had been some interchange of the specimens or
their labels, and the second specimen was really the one raised from the
Jersey seeds,
The volumes of the Sloane herbarium at the British Museum are instrue-
tive. In vol. 121, containing the Plante Britannice of the Rev. Adam
Buddle (1660-1715), at p. 6, there is a scrap of /. plantagineum, labelled
“ FEehii altera species, Dod. ; Lycopsis C. D.," and then, alongside of each
other, two very similar spikes of Æ. vulgare : one the ordinary form, labelled
“ Echium flore magno. This I take to be the Lycopsis anglica, Lob. Ger. ete.,
though "tis very common”; the other var. parviflorum labelled “ Echium
Jlore parvo. This I take to be the /Zehium vulgare CO. B. ete., though very
searce.” Thus we have Ray’s three kinds on the sheet, and we also
have the first suggestion of the identification, afterwards adopted by Miller,
of ©. Bauhin's vulgare with the small-tlowered form.
In vol. 151, p. 177, among Petiver's plants, are four specimens of which
two are Æ. vulgare; the third marked “Echium anglicum flore minore, stylo
tantum evserto” is var, parviflorum, and the fourth, although labelled
* E. ramosius flore suaveolente H. Ox.," is merely the white-flowered form of
vulgare. Then in vol. 321, p. 63, among Boerhaave's specimens, is a piece
of E. plantagineum labelled “ Lycopsis C. B.; Altera anglica, Lob. ; Lehii
altera species. Dod.”
To sum up the pre-Linnean position :
1. Lycopsis altera. anglica Lobel= Kehium pyrenaicum grown by him in
London. :
3. Echii altera. species, Dod., is identical with Lobels Lycopsis altera
anglica.
3. Lycopsis, C. B. P., is Echium plantagineum, of which he knew the
pink-purple and the purple-blue variations.
SOME CRITICAL SPECIES OF ECHIUM. 433
. Parkinson in 1640 and Merrett in 1666 first apply Lobel’s name,
probably known to them through Gerard, to an English form of
He
vulgare.
This idea is taken up by Richardson in Ray, who assigns Lobel's
name to the pistillate form of vulgare from northern England,
forgetting that the figure shows protruded stamens and a totally
different habit, and refers Lycopsis, C. B. P., to the Jersey plant,
i.e. to E. plantagineum.
. Morison refers Lycopsis, C. B. P., to the Jersey plant and suggests
that Lobel's may be the same.
7. Boerhaave refers Lobel's, C. Bauhin's, and Dodoens's names all to
E. plantagineum.
Or
for)
So far, the identity of Lobel’s plant with some form of X. italicum does
not seem to have been suspected. But Linneeus, Sp. Pl. p. 139 (1753),
quotes Lycopsis, Bauh. Pin. as var. 8 of ŒE. italicum, showing that he
recognised the true significance of Lobel’s figure, but misunderstood
Bauhin's Lyeopsis. In ed. 2, p. 200 (1762), he adds some characters to
distinguish var. 8 from typical italiewn, which do not alter the position.
Only, as the English authors had quoted both Bauhin's and Lobel's names
for British plants, he not unnaturally fell into the trap and supposing
italicum in some form to be British, quotes for it * Anglia.” In the same
year, 1762, Hudson reproduces the theory that Lobel's name is one of the
English forms of vulgare, for which he now must coin a binomial name, and
therefore creates /7. anglicum, though he oddly applies that name to the
common form of FK. vulgare and the name vulgare to the rare pistillate form,
just as Buddle had done before him. He also identified Lycopsis, C. B. P.,
with the Jersey plant, but finding it already quoted by Linneus for
FE. italicum, thinks it necessary to call it italicum, using the Linnean
diagnosis of this, which is inapplicable to the Jersey plantagineum.
Linnzeus saw Hudson's work in time to insert a reference in the Appendix
of Sp. Pl. ed. 2, p. 1678, where he treats Hudson's Æ. anglicum as synony-
mous with his own Æ. italicum, and Hudson's Æ. italicum as equivalent
to his Æ. italicum B, which afterwards became Æ. pyrenaicum in the
*Mantissa. These disastrous identifications are based entirely, as so often
in Linnzeus’s work, on the old synonyms quoted by Hudson. Thus he, Miller,
and Hudson between them introduced a fresh error, that Æ. italicum in some
form isa native both of England and of Jersey. This will be dealt with
under Miller's third species.
Miller's third species is ECHIUM ITALICUM corollis viz calycem e«cedentibus,
margine villosis. Hort. Upsal. 35; Mehium majus et asperius flore albo,
C. B. P., 255, This grows naturally in the south of France, in Italy, and in
434 MR. C. C. LACAITA: A REVISION OF
the Isle of Jersey. It has never been doubted that this is Æ. italieum, L.,
Sp. Pl. p. 139 (1753), exclusa var. 8. The erroneous account of the length
of the corollas is copied from Linnzeus, who repeated it from his Hort. Ups,
in Sp. Pl. ed. 1, but not in ed. 2, where the first really intelligible account of
E. italicum occurs,
There is only one sheet of Æ. italicum in Herb. Miller. It agrees perfectly
with the specimen in the Linnean herbarium, which is precisely Æ. altissimum,
Jacq. There is no writing on the sheet in Miller's hand, the name italieum
being written in pencil by Solander. There is no other Chelsea Garden
specimen of italicum ; one bearing no. 2716 and the words * E, italicum
1776,” with a penstroke through them, is 7. plantagineum. It cannot have
been labelled italicum by Miller, for he died in 1771.
But Jersey must be exeluded from the habitat. [ts supposed presence
there is due partly to reliance on the unfortunate * Lycopsis" muddle. The
false argument was this: Lycopsis has been found plentifully in Jersey by
Sherard; Linnwus says Lycopsis is italicum and Hudson also calls the Jersey
plant italicum ; therefore italicum grows in Jersey. But the error has been
kept alive owing to confusion between the white-flowered form of vulgare
and italicum. Although vulgare is not quoted for Jersey by Lester-Garland,
he says it is frequent in Herm and common in Alderney, and it looks as if
the white form had been found in Jersey in the past, though it cannot be
claimed with certainty. In Herb. Sloane, vol. 151, p. 177, there is an
example of white vulgare among Petiver's plants bearing his label : “Hehium
ramosus flore. suaveolente, H. Ox. ; altera species, Dod. ; Lycopsis, C. B.,
Anglica, Ger. ; altera Anglica, Lob. Wall Bugloss in the Isle of Jersey."
The plant is certainly vulgare fl. albo, but it is possible that the last words of
the label may not mean that this individual came from the island.
Kchium italicum is professedly figured in E. B. tab. 2081 as “a native
of Jersey, for spocimens of which from Chelsea Garden we are obliged to
Mr. Fairbairn." Probably the seeds of the Chelsea Garden plant had not
really come from Jersey. Sowerby's original drawing at the British
Museum only says * Chelsea Garden," and does not mention Jersey. The
diawing seems to be a compromise between Æ. italicum and white-flowered
I. vulgare, of which an explanation is suggested by a sheet in Sowerby’s
herbarium at Mus. Brit, where there are two specimens, both referred
to E. B. tab. 2081, but while that on the left is true italicum, the one on
the right is white vulgare. Between the two is a piece, in bud only, from
J. Dickson’s * Hortus Siecus Britannicus; 1793-1799, marked * Echium
italicum, Jersey." As far as can be judged in so early a stage, this really
is X. italicum, but did it come from Jersey? or is it a garden plant from
seeds wrongly supposed to come from that island? If really from Jersey,
E. italicum must have existed there at that time as an escape and since
died out,
T ini ee, ee re T
SOME CRITICAL SPECIES OF ECHIUM. 435
White-flowered Æ. vulgare occurs now and then in England, but
Mr. Druce informs me that he knows no case of Æ. italicum being found
except as a casual at Ware in Herts, where foreign poultry food is brought,
and at Cothill in Berks. He has kindly allowed me to examine his specimens
from both localities. Neither is true italicum. They are a very lax form
of E. pyrenaicum, identical with Lapeyrouse’s authentic specimens of his
E. pyramidale.
Miller's fourth species is “ ECHIUM LUSITANICUM corollis stamine longioribus,
L. Sp. 200 ; Æ. amplissimo folio, lusitanicum Tourn. Instag
“It grows naturally in Portugal and Spain ; the lower leaves often are
more than a foot long and two inches broad in the middle, gradually
lessening to both ends, these are covered with soft hairs. The stalks grow
two feet high ; the flowers are in short spikes coming from the side of the
stalks; the petals of these are longer than the stamina.”
This admirable description can apply to nothing but Echium Broteri,
Sampaio ex Coutinho, as Boragineas de Portugal in Dol. Soc. Brot. xxi.
p. 111 (1905), and in Cout. FI. Ports ps 499— E lusitanicum, L. (saltem
herb.) 2 Æ. italicum, Brot. Fl. Lus. i. p. 290, and Hfg. et Lk. Fl. Port. i.
p. 185, non L. The account of the leaves is quite conclusive, They are
unlike those of any other species.
Miller cannot possibly have meant Æ. plantagineum, to which for some
inscrutable reason his Æ. lusitanicum is referred, along with Æ. amplissimo
folio lusitanicum, Tourn., by Link on p. 186. De Candolle, Prodr. x.
pp. 20 & 22, made the confusion worse by accepting this false identifica-
tion for Miller's E. lusitanicum, but not for Tournefort’s synonym nor for
E. lusitanicum, L.
For an explanation of the often-repeated false statement that the Linnean
herbarium contains no example of Æ. lusitanicum, and for a justification of
the identification of E. lusitanicum, L., with Æ. Broteri, Samp., I must refer
to my notes on the Echia of Linneus. The only g ground for doubting the
identity of. Miller's, or indeed of Linneeus’s, lusitanicum with E. Broteri is
the account which Linnzus and Miller both give of the relative length of
stamens and corolla, which certainly is not true of Æ. Broteri, but it would
also be untrue of any other Portuguese species to which Miller’s name could
be supposed to apply.
It looks as if Miller had simply copied Linnzeus’s statement without
thought. I can only suppose that Linnzeus himself made a mistake. There
is an analogous misdescription of the length of the corollas of Æ. italicum in
Sp. Pl. ed. 1, where these are quite wrongly said to be * vix calycem
B ues
There exists no specimen of Æ. lusitanicum marked “ Herb. Miller," but
there is a fine one from Chelsea Garden marked, though in pencil, by
LINN. JOURN.— BOTANY, VOL. XLIV. 20
Eom
5
i
t36 MR. C. C. LACAITA : A REVISION OF
Solander “italicum” and “ Echium lusitanicum L. M." This is precisely
E. Broteri, of which there are two other old specimens in Herb. Sloane,
both from Petiver’s collections. These are mentioned in detail in my notes
on the Hehia of Linnæus.
Miller's fifth Echium is * Eoutum enETICUM calycibus fructescentibus dis-
tantibus, caule procumbente, Linn. Hort. Upsal. 35; Echium Creticum
latifolium rubrum C. B. P. 254. The fifth sort grows naturally in Crete ;
this hath trailing hairy stalks, which grow about a foot long, and put out
several side branches, garnished with hairy spear-shaped leaves about
three inches long, and three-quarters of an inch broad, sitting close to
the stalks. The flowers come out on slender spikes upon long foot-stalks
which come from the wings of the leaves; they are large, of a reddish-
purple colour, which turns to a fine blue when they are dried; these stand
at a distance from each other on the spike. It is an annual plant which
flowers in July and decays in autumn.”
Echium ereticum, L., is a mixture of two very different species, which
Clusius had distinguished, one being W. ereticum angustifolium rubrum,
C. B. P., which is Miller's sixth sort; the other Æ. creticum latifolium
rubrum, C. D. P., which is represented by the specimen in Herb. Linn.
This is a cultivated plant, which cannot have been derived from any
Cretan kind, but is obviously a garden modification of X. grandiflorum, Dest.
(See my notes on the ehia of Linnzeus.)
According to the synonymy, Miller's Æ. ereticum should be the X. ereticum
of Herb. Linn. His statement that it grows naturally in Crete is a mere
parrot echo of what he had read. Clusius himself was uncertain about
the supposed Cretan origin. But, although the synonyms point to
E. creticum, Linn. herb., I fear that Miller mixed up with it Æ. plantagineum,
which is plentiful, though misnamed, in the old herbaria and not otherwise
referred to in his dictionary. Indeed, his English description is more
than suspiciously like Æ. plantagineum. The spear-shaped leaves he
mentions would exclude that species if he were speaking of radical leaves,
but he means the stem-leaves. The purplish flowers of plantagineum
often—not always—turn a “fine blue” when dried ; those of /. ereticum,
Linn. Herb., do not.
The only Cretan kind, other than plantagineum, that could be thought
of for a moment is Æ. parviflorum, Moench, especially in its luxuriant form,
var. erectum, DO. This, indeed, is the Æ. eretieum of the * Flora Greeea,’ but
not of Linnzus. It is, however, absolutely excluded by what Miller says of
the flowers and by the reference to the synonyms.
The specimens in the old herbaria, of which a fuller account will be found
in my notes on the Hehia of Linnæus, confirm the suspicion of a muddle.
There are none at the British Museum marked * Herb, Miller," One from
SOME CRITICAL SPECIES OF ECHIUM. 437
Chelsea Garden in 1749, no. 1367, labelled “ E. ereticum angustifolium rubrum,
C. B. P," where “ angustifolium” obviously is a slip of the pen for latifolium,
is the Æ. creticum of Herb. Linn., but another, labelled by Solander
* FE. ereticum," is a variety of E. pustulatum, S. & S., or one of the allied
Mediterranean forms. There is also a sheet with no indication of where
its contents grew, labelled in Miller's handwriting * Æ. calycibus. fruetes-
centibus ete., H. U. 35: E. ereticum latifolium rubrum" and by Solander
^ E. ereticum.” Unfortunately, it contains two pieces belonging to different
species, the smaller being really Æ. ereticum, Herb. Linn., but the larger
F. plantagineum.
Therefore I think we must consider Æ. creticum, Mill, to be a nomen
confusum. This is immaterial, as it does not affect the interpretation of
E. eveticum, L.
Miller’s sixth sort is ECHIUM ANGUSTIFOLIUM caule ramoso, aspero, foliis
calloso-verrucosis, staminibus corolla longioribus. E. creticum angustifolium
rubrum, C. B. P., 254. “This sixth sort hath branching stalks which grow
a foot and a half long, declining toward the ground ; they are covered with
stinging hairs; the leaves are four inches long and not more than half
an inch broad ; they are pretty much warted and are hairy. The flowers
grow in loose spikes from the side of the stalks, and also at the end of the
branches; they are of a reddish-purple colour, but not so large as those of.
the former sort, and the stamina of these are longer than the petal. This
is also an annual plant which grows naturally in Crete.”
From the excellent description and from the synonym this is certainly
Echium hispidum, Sibth. et Sm., Fl. Gr. Prodr. i. p. 125 (1806), Fl. Gr.
tab. 181 =F. elegans, Lehm. Asperif. p. 459 (1818) =Æ. Sibthorpii, Roem. et
Sch. Syst. iv. p. 26 (1819). Lehmann's and Roemer’s names were only created
to replace that of Sibthorp, owing to the existence of an earlier homonym,
FE. hispidum, Thunb., a Cape plant, now referred to the genus Lobostemon.
We must boldly adopt Miller's name of angustifolium, if the rule of priority
is to be adhered to. To such a course three objections might be raised. Firstly,
that it would cause two changes in accepted nomenclature, for, as E. angusti-
folium, Lam., would have to give way to Æ. angustifolium, Mill., the name of
that species too would have to be changed. Secondly, Miller calls his plant
annual, whereas elegans is stated by Haláesy in Consp. Fl. Gr. ii. p. 338, to
be perennial. Such a mistake in this genus is very easy to make, unless the
writer is well acquainted with the plant in its native haunts. Moreover,
many Mediterranean coast perennials will not survive the damp of English
autumns, even if they resist the frost of winter, and have to be treated as
annuals in English gardens, where alone Miller knew his plants. Ithink
we may fairly brush aside this objection.
Thirdly, the presence of an inconvenient specimen in Herb. Miller with
202
438 ON SOME CRITICAL SPECIES OF ECHIUM.
a ticket in his own hand: * Æ. caule ramoso aspero, foliis calloso-verrucosis,
staminibus corollis longioribus," which is the diagnosis of angustifolium and
“ E, erelicum angustifolium rubrum C. B. P." to which Solander has added
in pencil “Æ, orientale” and “ angustifolium Mill. Dict. no. 6.” Unfortu-
nately, the plant is not ereticum angustifolium rubrum and does not agree
with Miller's description. It is referable to Æ. pustulatum, which was often
confused in early times with Æ. hispidum, e. g., by Smith himself. (See my
note on the chia of Herb. Sibthorp, p. 386.) We must not be governed by the
specimen and ignore the excellent description, especially when the specimen
is a garden plant. It is common to find specimens in the old herbaria
which do not agree with the species described. The case is different from
that of X. maritimum, Willd., which is represented in that author's herbarium
by a single piece of X. plantagineum, for Willdenow has told us that he never
saw but one example of his maritimum. See de Coincy in Morot’s Journ.
Bot. xiv. p. 162, who consequently alters the name maritimum to confusum,
a proceeding in which Rouy has refused to follow him. I may quote the
apposite remarks of Bonnet in his account of Lamarck’s herbarium, Journ.
Bot. xvi. p. 137 :—** On connait les incohérences de l'herbier de Linné et
les confusions de Willdenow .... un échantillon d'herbier ne doit servir à
infirmer une description publiée que s'il n'existe aucun doute sur son authen-
ticité ; la description originale, méme incomplète ou défectueuse reste entière,
tandis que l'échantillon d'herbier est trop souvent soumis à des causes qui en
altérent l'intégrité et l'authenticité." Now, though we know that Miller
wrote ihat ticket, we do not know who at some later time gummed it to the
sheet where we now find it.
On the whole, I think we should not be deterred by these three obstacles
from reviving the neglected name, which seems to have been noticed by no
one but Poiret in Dict. Encl. viii. p. 671, where he quotes it as var. b of
E. creticum, L., and equivalent to /. ereticum angustifolium rubrum, Tourn.,
which is known to be Æ. hispidum. Poiret is, of course, wrong in assigning
as synonyms of angustifolium, Mill., Lamarek’s angustifolium and Darrelier,
ic. 1011, which represents Lamarck’s plant.
Mh. H. B. GUPPY ON PLANT-DISTRIBUTION. 439
Plant-Distribution from the Standpoint of an Idealist.
By H. B. Guepy, M.B: F.L.S.
[Read 7th February, 1918. ]
THE most interesting and at the same time the most mysterious features of
plant-distribution centre around the rise of the great families. These ancient
plant-groups belong so much to an era of other things, other ways, and other
conditions, that the employment of such terms as * genera” and “ species "
in connection with their origin seems to be almost meaningless. All the
influences that we see normally at work around us now could only relate to
the differentiation of the family-types into genera and species ; and any
theory that on such grounds endeavours to apply the present to the past in
deciphering their origin would apparently be attempting an impossible task.
One ventures to think that only the hypothesis that finds its guide to the
past in the abnormalities of the present could be of service here. This would
seem to place the pre-differentiation era, the age that witnessed the rise of
the great families, outside the field of the Natural Selection theory, and in
default of its aid to cause us to look to the Mutation hypothesis for guidance.
Yet, although Darwin came to reject the “sport,” the original scope of his
theory was large enough to admit it; and it may be,as the writer holds,
that the antithesis between the two theories is more apparent than real. Yet
the Mutation hypothesis was conceived in the spirit of Darwinism, was
framed on Darwinian lines, and was formulated in Darwinian language ;
` and it is not easy to understand how the two theories were allowed to
acquire the appearance of being mutually exclusive. There is room for both
within the boundaries of the theory of Natural Selection as Darwin first
conceived it; and there is work for both schools in its extension, its improve-
ment, and its emendation. Whilst the Mutationist would find a fruitful
feld for his labours in the era of the rise of the great family-types, the
Darwinian evolutionist would be occupied with their subsequent differentia-
tion into tribes, genera, and species.
The rise of the great families and the lesson of the Composite and the
Gentians.—Two papers of great importance from this standpoint to the
student of plant-distribution were published in the * Journal of the Linnean
Society’ in 1873 and 1888, the first by Bentham on the Composite, the
second by Huxley on the Gentians. They are important because, in dealing
with the beginnings of the distribution of these two families, they ask the
same questions and raise the same issues; and it is needless to add that,
although in one case the methods employed were those of a great botanist
and in the other those of a great zoologist, they bear in each case the
impress of a master hand. Those interested in the subject will remember
440 MR. H. B. GUPPY : PLANT-DISTRIBUTION
that two years before the publication of his paper Huxley had outlined his
views in two letters to Hooker, letters which are given in the 2nd edition
of his * Life and Letters,’ by his son (ii. 465-6, 1903).
Itis proposed to commenee this paper with a comparison of the views
held by these investiga'ors respecting the early history of the distribution of
the two families concerned. Both are agreed as to the original wide distri-
bution of the primitive forms over the world, and both credit them with
ignoring the great physical features of the globe, as at present displayed.
Broad oceans, great deserts, and lofty mountain-ranges are all out of the
reckoning in the respective discussions relating to the spread of these two
families over the earth. Bentham holds to a very wide dispersion of the
original stock of the Composite: over the world when the physical configura-
tion was very different from what it is in our day. Huxley speaks of a
* primitively continuous area of distribution," and leaves the matter there.
With reference to the state of differentiation of the original stock when it
conquered the earth, Bentham holds that the Old and the New Worlds
possessed the family at the earliest recognizable stage. Huxley hypothecated
a widely spread primitive type that subsequently differentiated over the
globe. Both, in imputing a high antiquity to the respective stocks, know-
ingly disregarded the lack of geological evidence, the one considering that
the Composite dated back to an early geological period, the other holding
that even the more specialized and consequently the more recent of the
Gentians might have lived in the Cretaceous epoch. But Huxley went
even beyond this when he assumed that the * Ur-Gentian" might be
carried back “almost as much farther as probabilities permit us to carry
the existence of flowering plants."
For neither Bentham nor Huxley were the main features of the distri-
bution of these two families concerned with means of dispersal. Huxley
makes but little appeal to them and Bentham diseredits their efficacy.
Bentham begins with a family already universally distributed, although he
implies an original centre of dispersion. Huxley would have nothing to do
with any such centres, and his plain words on this subject ought never to
be forgotten by the student of distribution. All such notions were excluded
for him in a type that followed the principle of the simpler and older the
type the greater its range.—* The facts of distribution of the Gentianem
are (he writes) not to be accounted for by migration from any centre of
diffusion to which locality can be assigned in the present condition of the
world." The problem was for him essentially a matter of the local
modification of plants at different points of a *primitively continuous”
area. Both Bentham and Huxley are at one in their conclusion that the
main features of the distribution of these two families were determined
in ages geologically remote ; and neither’s view of the early stages in the
history of their distribution leaves any room for an appeal to centres of
"T d CAN TONER S V RIO HT
FROM THE STANDPOINT OF AN IDEALIST. 441
dispersion. If behind the facts of distribution lies the cardinal principle
that the farther we trace a type back the more generalized are its characters
and the wider its range, the question of its original home is obviously not
raised.
What, we may ask, was the bearing of these views on general taxonomic
principles? Bentham was dealing with a world-ranging family holding
about a tenth of the total known number of Angiosperms and displaying
relatively few of the difficulties presented by small families with restricted
distribution. He followed orthodox lines and the results were not dis-
quieting to the systematist, although, if he had pushed his conclusions home,
a clash with prevailing practice would have happened. With Huxley it
was very different, He was concerned with a much smaller family, one
less widely distributed and displaying a preference for mountainous regions.
To it he applied the same method of postulating a wide-ranging but little
differentiated primitive type, and in so doing he raised many of the difficulties
presented by the smaller families. But his method, as he perceived, con-
siderably upset the accepted grouping of the Gentians, and in his enthusiasm
in its advocacy he contended that “a revision of Taxonomy and Distribution
from the point of view of the Evolution doctrine will hardly fail to revolu-
tionise both.” But the curious point is that as far as the early stages in the
history of the Gentians are concerned the method advocated by Huxley was
not Darwinism, as then conceived. It was pure Differentiation. With
Bentham also, although clinging to the idea of a centre of dispersion, or of
a home, for the Composite, he began with a world-ranging but slightly
differentiated primitive type, and like Huxley with the Gentians he allowed
it to work out its own lines of differentiation in the various regions of the
globe. This is Evolution on a plane; and the implication is that since the
rise of the great families in the Mesozoic ages little else has been effected.
The Differentiation hypothesis —Although this hypothesis has rarely been
formulated, there are various ways of stating it. It appears in a variety of
guises in many a monograph of the families. The writer’s mode of pre-
senting it is to associate it with another theory relating to the differentiation
of conditions, the modification of form being regarded as the response to the
progressive differentiation of conditions. But it would be possible to deal
independently with the différentiation through the ages of the family into
tribes, of the tribe into genera, and of the genus into species. Yet the two
are commonly implied, and it is hard to dissociate the idea of differentiation
of type from that of diversification of condition. There may, however, be
different ways of stating the relation. The following is the writer’s method
of doing so.
He has come to close quarters with the central problem in successive
stages. In the first stage the world, as far as plants are concerned, was
mainly a differentiating world in which wide-ranging generalized types had
442: MR. H. b. GUPPY : PLANT-DISTRIBUTION
differentiated in response to the progressive diversification of conditions
originally uniform, a world in which the family differentiated into tribes,
the tribes into genera, and the genera into species. But on perceiving that
such a theory could only explain distribution where a continuous land-mass,
not affected by unstable climatic conditions, was concerned, he came to
recognize that the operations of the differentiating agencies had been largely
controlled and directed by the divergence of the two great land-masses from
the north, a control in its turn influenced by the secular fluctuations of
climate and by the barriers that lay across the lines of migration. It may
be stated that the progressive differentiation of floras in response to diversi-
fication of conditions has long been recognized. The era of world-wide
floras, as Mr. C. Reid well puts it (‘Encyclopedia Britannica, edit. 10,
vol. xxxi. p. 432), began to pass away after the Cretaceous age, and from that
period onward plants have responded more and more to the differentiation
of conditions and have arranged themselves more and more according to
geographical boundaries and climatic zones. The general trend of events in
later ages is sufficiently indicated by the frequent application to these early
types by various writers of such epithets as “comprehensive,” * generalized,”
* mixed," * synthetic," ete.
The views long advocated by Thiselton-Dyer brought about the first
modification in the original interpretation of the theory, and the writer came
in this way to lay stress on the point that the mingling of the floras of the
eastern and western worlds might be regarded as the result of the successive
migrations to and from the north polar region under the stress of climatic
changes. The third stage was reached when he realized as a result of the
statistical treatment of the subject, which is dealt with in later pages, that
there was much in the distribution of the larger groups which the mixing of
the eastern and western floras in the north polar regions would not explain.
Though true of the smaller groups, as with species, the principle that the
community between the Old and the New World is an affair of the north
did not materialise with respect to families. On the contrary, it appeared,
with regard to families of the first rank and the groups behind them, that
the main features of. distribution would have been much the same as they
now are if the land of the globe had been gathered into a single mass.
Thus the author came to distinguish between the larger and smaller groups
in the response made to the great bi-cleavage of the land-surface of the
globe, and to restrict the influences of the existing relations of land and sea
to the smaller groups, as in the case of genera and species. This led him to
perceive that if the differentiation hypothesis was valid the families and the
larger groups behind them had. not only ignored the bi-cleavage of the land-
mass of the globe, but had been developed under conditions very different
from those in which their genera and species had been produced.
The independent behaviour of the great families with respect to existing
Tult ENE ENSA ae aoe eT E S
FROM THE STANDPOINT OF AN IDEALIST. 443
geographical conditions opened to him a prospect of removing a serious
difficulty that might have been fatal to the general theory ; and that was
the difficulty of conceiving the early stages in the differentiation of a type
in response to the first stages of the diversification of uniform conditions.
Since characters become more constant and adaptivity to present conditions
becomes less marked as one proceeds up the scale from the species to the
family, it was obvious that if the hypothesis was to stand a different order of
things had to be postulated for the development of the larger groups, an
order of things in which instability of characters was associated with
uniformity of conditions. The responses of our great family types to the
changes in environment are negligible. Yet the distant age of the Creta-
ceous that witnessed the deployment of the Angiosperms, much as we know
them now, must have been preceded by an era of great instability of floral
characters—characters on which the taxonomist has based his families and
his groups of families, and characters that have been more or less fixed
during the ages that have since elapsed.
Obviously one was here face to face with a different order of things, but
some time elapsed before further progress could be made in the elaboration of
the theory. Having abandoned the position that uniformity of conditions and
immutability of type went together, a position that was the logical sequence
of the differentiation hypothesis as at first conceived, one had either to adopt
the opposite view or throw over the theory. It was nota dilemma peculiar
to the differentiation theory, since the mutationist and the Darwinian
evolutionist experience a similar difficulty when they deal with the genesis
of the larger groups, the development of the family requiring the instability
of characters which are mainly constant under present conditions. The way
out of the difficulty was suggested on reading the account by Dr. Willis
of the extreme uniformity of conditions in which the Podostemacee and
Tristichacez live in mountain-torrents and rushing streams around the
tropics, a description of a state of things approaching the primeval state
as far as uniformity is concerned. He describes the great morphological
changes of the floral and vegetative organs under such conditions, modifica-
tions characterized as without any adaptive significance and as the result of
free mutation in every direction (Proc. Roy. Soc. vol. Ixxxvii. pp. 546, 548 ;
1914). He speaks of “the most astonishing variety of morphological
structure" under conditions of life “absolutely uniform " (bid. p. 533)
On the results of the investigation of these families for many years he builds
a powerful argument for the Mutation theory, and one can scarcely doubt
that in time he will adopt a standpoint not essentially different from that of
the Differentiation hypothesis. But what one is concerned with here is the
association of extreme uniformity of conditions with extreme instability of
type. It isa picture of the abnormal side of plant-life.
In explanation of the remarkable mutations of the floral organs displayed
a a WR SE
s
k
441 MR. H. b. GUPPY : PLANT-DISTRIBUTION
by the Podostemacez, Dr. Willis connects the dorsiventrality of these
organs with that of the vegetative organs, thus introdueing a factor appa-
rently subversive of all taxonomic principles. There is in his warning of
the insecurity of the taxonomist’s position an echo of Huxley's defiant note
respeeting the revolutionizing effect of the evolutionary doctrine on the
principles of taxonomy and distribution. Dr. Willis wields the Podostemacerm,
as Huxley wielded the Gentians, in his attack on prevailing principles. In
the ease of Huxley it was concerned with the differentiation of a wide-
ranging primitive family type, and one ean scarcely doubt that he struck a
true note in his declaration. But it applied only to the second era, the era
of differentiation, the age of influences still in operation, the age of normality,
if we may so term it ; and it ought to havea profound effect on the methods
of the taxonomist and on the prineiples of distribution. In the case of
Dr. Willis it is concerned with the abnormal side of plant-life and does not
really affect the validity of prevailing taxonomic principles. The difference
is very significant, since, in view of the position taken at the commencement
of this paper, we can in our day only look in that direction for a clue to the
influence at work during the first era in the history of the Angiosperms,
the age that witnessed the rise of the great families, the age of abnormalities,
as it may be called. It is in this first era that the Mutation theory will find
its appropriate field of investigation, and it is here that the principles
disclosed by Dr. Willis in his prolonged investigations on the Podostemaceze
wil apply.
Before proceeding. to deal in the two following paragraphs with my
interpretation of the lessons to be learned from the behaviour of the Podo-
stemacee and with their application to the first era, I may say that
Dr. Willis left room for an interpretation of the same kind, but was
prevented, as he tells me, from entering a domain of pure surmise.
Postulating for terrestrial] plants an era when uniformity in environment
was the rule—an era, one might imagine, of. great atmospheric humidity,
when persistent cloud-coverings blanketed the globe and when the same
equable temperature everywhere prevailed,—the writer pictured a plant-
organism under such conditions as behaving very much like a ship in a
calm, drifting in a morphological sense in all directions and displaying
unchecked and irresponsive variations of the floral organs of a kind very
disquieting to the taxonomist and all non-adaptive in their nature. He
came to see that such modifications would become more and more fixed as
the differentiation of conditions proceeded, the degree of mutability varying
inversely with the diversification of environment.
Stated in the language of the mutationist, this would imply that the
mutations of the floral organs of our own day represent all that remains of
the capacity for great morphological changes in the early ages of the history
of the Angiosperms. A mutation as at present recognized is non-adaptive.
FROM THE STANDPOINT OF AN IDEALIST. 445
Yet it may be adaptive in another sense—namely, in its response to the
remnant of the conditions of an age long passed away. One might regard
it as the last kick of the organism in response to what remains of the
primeval uniformity of conditions, its last effort to break through the ever-
contracting ring of the differentiating agencies. The farther we go back
the greater is the capacity for mutations and the greater will be the
mutations ; and it is argued that they ought to be more frequent and more
extensive in plant-groups of large than of small range. A large mutation
would usually be impossible nowadays except under conditions approaching
those of the early ages in their uniformity. One would look for some
approach to those times in the dense forests of tropical lowlands and in the
forests of the cloud-belt or rain-zone on tropical mountains.
The ascription of periodicity to mutability by De Vries is well known,
and appeals to this principle in elucidation of the rapid rise of the Angio-
sperms in the Cretaceous period have been made ; but the question, why
basic characters so mutable then are stable now, always remains. The
position is well put by Harshberger in his great work, *The Phytogeo-
graphic Survey of North America, p. 173, 1911. “If this periodicity of
mutation (so he writes) is recognized as an evolutionary principle, we have
a reasonable explanation for the sudden appearance of so many new forms
during the Cretaceous period, for during this stage of the development of
the vegetable kingdom, through causes yet unknown, the progenitors of the
existing phanerogams were in a high state of mutability."
A few remarks may here be made on the relation of the views of distri-
bution here advocated to the Age and Area theory of Dr. Willis. If one
for the moment ignores his adoption of the prevailing practice of building
up a family from the species, there is but little that is essentially inconsistent
with the differentiation hypothesis. Had he formally associated with his
Age and Area principle the twin principle of Rank and Range, and. all that
it implies, he could not have avoided coming into line in this matter.
Since he extends his views to the larger groups, his Age and Area theory
is of. general application, and his conception of the distribution of families,
apart from his standpoint of their genesis, might very well have been
acquired in a line of argument favouring the views supported in this paper.
His discussions of the Dilleniacez, Menispermaceze, and Podostemaces are
cases in point. (See Ann. Roy. Bot. Gard. Peradeniya, 1902, 1907;
Phil. Trans. Roy. Soc. Lond. 1915 ; Proc. Roy. Soc. 1914; Ann. Bot. 1915,
ete.) Thus he connects the origin of these three pantropical families with
the most primitive and most widely distributed genera— genera that almost
possess the range of the families. This is differentiation pure and simple.
Then again, though he departs from the principle of differentiation when
he endeavours to find the original centre of dispersion or home of the
Dilleniacexe, he comes very near it in the case of the Menispermacez in his
446 MR. H. B. GUPPY : PLANT-DISTRIBUTION
inference that the primitive genera * must have commenced when there was
still a possibility of reaching both the great land masses" (Phil. Trans.
ibid. p. 338). This recalls Bentham’s opinion respecting the Composite,
that the two hemispheres, the east and the west, must have possessed the
family at its earliest recognizable stage.
So, again, when he observes in connection with the Podostemace and
Tristichaces that “ the only widespread genera are the non-specialised ones,
whilst the more specialised the genus, on the whole, the less is its area of
distribution " (Proe. Roy. Soc. 1914, p. 545), and when he remarks that
“the larger the group and the fewer the characters on which it is based, the
greater the likelihood of its being polyphyletic "—that is to say, of arising
independently in localities remote from each other (Ann. Roy. Bot. Gard.
Perad. p. 447, 1902),—he is expressing the differentiation standpoint. But
_ the writer cannot conceive how the principle, often implied in the foregoing
remarks, that the simpler the form the wider its range, could apply to groups
that have been built up, as Dr. Willis infers, by the species taking the
generic step, the genus the tribal step, and the tribe the family step (idid.).
But, apart from this, there is a great deal that links together the two
views of distribution; yet one may add that whatever view we take of
distribution, whether that of the Darwinian evolutionist, or that of the
Mutationist, or that of the advocate of pure differentiation, we all get into
the same dilemma when we handle the larger groups. If we require for
their development the mutability or instability of the characters on which
the taxonomist bases his larger groups, characters that in our own age are
relatively immutable, we cannot look to existing prevailing conditions for
guidance in the matter. However, Dr. Willis in his account of the
astounding modifications experienced by the Podostemacew, under conditions
described by him as “unique” among plants, offers, as the writer has
already explained, a way out of the difficulty.
The statistical treatment of Distribution.—-lf the Differentiation hypothesis
is valid, we may now ask what we should be justified in expecting from a
statistical treatment of the main features of plant-distribution. If we listen
to the story of the early stages in the distribution of the Composite and
Gentianaces, as interpreted by Bentham and Huxley, we should expect that
the larger plant-groups would to a great extent ignore the cleavage of the
land of the globe into two large masses diverging from the north, and that
the response made to the existing arrangement of land and sea would
increase as we go down the differentiating scale, being least for the family
and greatest for the species. On the other hand, we should expect a
marked response of the larger plant-groups to the climatic differentiation of
the latitudinal zones.
If in our investigations with the family as our starting-point we disclose a
method and a system that could not be brought about by a procedure so
haphazard as that involved in commencing the genetie sequence with the
FROM THE STANDPOINT OF AN IDEALIST. 447
species and ending with the family, then we shall make an important step
towards the proper appreciation of the main problems concerned in dis-
tribution. That two such opposite methods should seem possible, as
beginning in one case with the species and the other with the family,
and that we should be indifferent to the result, whether it be chaos or
order, are indications of failure to appreciate what really matters in plant-
distribution.
It is by no means urgent to go back to the beginning of things, to account
for the origin of families or even of species, to seek for centres of dispersion
or the homes of plant-groups, to upset the world’s geography, or to account
for progressive evolution. What is urgent is to be able to state the main
problem, and that cannot be done without some agreement about essentials.
One cannot help thinking that if a hundred students of distribution were
asked to do this, they would view the subject from such a variety of
standpoints that the task of finding a common basis of agreement would
be exceedingly difficult. The claim of the family for priority of treatment,
which is supported in this paper, is founded on what Bentham terms in the
case of the Composite the permanence of its characters. The very per-
sistence of the family lies in the fact that these characters make little or no
response to the extreme variations of existing conditions ; and it is with the
object of emphasizing its suitability for serving as a common ground of
agreement that these pages have been written.
The response of the families to the bi-cleavage of the land-mass of the
globe.—1lt is a remarkable fact that whilst the families of the Angiosperms
. respond in a marked degree to the differentiation of the climatic zones, they
largely ignore the cleavage of the land into two great masses diverging from
the north. Of the 272 families recognized in Engler's system 192, or
70:5 per cent., occur in both the eastern and western hemispheres (Tables I.,
IL, IIL). It would almost appear, as far as their occurrence in both
hemispheres is concerned, that the general distribution of the families over
the globe would not have been very different if all the land had been one
continuous little-divided mass.
It may, however, be objected that this community of families between the
Old and the New World may be mainly restricted to those most at home in
the colder latitudes of the north where the American and Eurasian land-
masses approach each other. If this were so, there would be but little force
in the above contention that the family largely ignores the bi-cleavage of the
land. Buta glance at the columns of Table IIT. will convince one that this
behaviour of the family is just as characteristic of the warm equatorial
regions as it is of the colder northern regions. There are 120 families
restricted to tropical and sub-tropical latitudes. Of these 73, or 61 per cent.,
are found in both the eastern and western worlds. There are 52 families
that are only at home in extra-tropical regions. After removing those
confined to the southern hemisphere, there remain 36, of which 23, or 64 per
CR Pop sd eee, ae
448 MR. H. B. GUPPY : PLANT-DISTRIBUTION
cent., occur in both the Old and New Worlds, We obtain similar indica-
tions by also introducing the element of those families that are mainly,
though not exclusively, either tropical or temperate. Thus, by extending
the method employed in Tables IV. and V., we arrive at the conclusion that
whilst 69 per cent. of the families that are mainly or exclusively tropical
(158 in all) occur in both the Old and New Worlds, the proportion for
families exclusively or mainly extra-tropical in the northern hemisphere
(62 in all) is 77 per cent. (consult note at end of the paper). Under the
circumstances the difference is small, and there is little to support the
objeetion that the families common to thé east and the west gather in high
northern latitudes. But it would have been enough to point out that there
is little room for such an objection in view of the fact that the proportion of
tropical families that are common to the eastern and western worlds (69 per
cent.) is very close to the proportion obtained for the families of the Angio-
sperms in the mass (70 per cent.).
The question whether the connection by families between the Old and the
New World is chiefly a problem of the cold regions of the north, where
the great Ameriean and Eurasian land-masses converge, is sufficiently
answered by the behaviour of the seven terrestrial sub-families of the
Aracem. All of them oceur in both the eastern and the western worlds, yet
four of them are exclusively tropical, two are distributed in both the tropical
and the temperate zones, and only one (Calloidez) is restricted to cold
northern latitudes. This last seems to be the only one of the seven sub-
families that holds species common to the east and the west.
We have now raised a very interesting point. Although the families
common to the two worlds do not gather in the north, the species behave in
a very different fashion. It is there that the species common to the east
and the west mostly congregate. Thus Harshberger states that of the 364
species of phanerogamic plants found in arctic western America, 320, or
about 87 per cent., occur in temperate and arctic Asia; while of the 379
species in arctic east America, 239, or 63 per cent., are also found in the
arctic regions of Europe (‘Phytogeographic Survey of North America,’
pp. 311, 312; 1911). The community of species rapidly diminishes as we
leave the north behind until we reach the tropies, where with the exception
of a few littoral, aquatie, and marsh plants, it disappears altogether, or is
only represented by a few plants, some of them not free from suspicion as
regards man's agency.
When we have two complemental families like the Myrsinaceæ and
the Primulacesw, the first tropieal and the second temperate, it is the
temperate family that alone displays a community of species between
the two hemispheres.
The manner in which the proportion of species held in common dwindles
as we go south is well illustrated by Carer, In the author's recent book on
FROM THE STANDPOINT OF AN IDEALIST. 449
the West Indies he has shown that the proportion of species which North
America holds in common with Eurasia is 93 per cent. in the arctic regions,
40 per cent. in the sub-arctie regions, 24 per cent. in temperate latitudes,
and 11 per cent. in the southern portion of the continent. If we take the
total Carex floras of the eastern and western worlds in the northern hemi-
sphere, we find that 80 per cent. of the species held in common are arctic,
29 per cent. sub-arctic, and 11 per cent. temperate. There are about
150 species common to North America and Eurasia, and of these two-thirds
are arctic and sub-arctic species.
I am not able here to deal fully with the distribution of genera from this
standpoint, but it cannot be doubted that the behaviour of genera common to
both the eastern and western worlds will be intermediate between that of
the species and families similarly distributed. This is established below in
an analysis I have made of the list of the chief genera of the Angiosperms
(about 3150) that is given by Dr. Willis in his ‘ Flowering Plants and
Ferns,’ 1908.
It has already been shown that whilst 69 per cent. of the families that
are exclusively or mainly tropical (the subtropical regions being here
included) oceur in both the Old and the New Worlds, the proportion for
families that are exclusively or mainly extratropical in the northern hemi-
sphere is 77 per cent. On the other hand, with the genera that are mainly
or exclusively tropical the proportion found in both worlds is only 23 per
cent. (408 out of 1781), whilst with those mainly or exclusively restricted
to regions beyond the tropics in the northern hemisphere the proportion
is 42 per cent. (437 out of 1045). The genera, therefore, are in their
behaviour intermediate between the families and the species. Whilst with
the species nearly all (80-90 per cent.) of those common to the eastern and
western worlds gather in the high latitudes of the north, with the families
there is but a small tendency in this direction, and reasons have been before
given for the belief that this tendency is even smaller than is above indicated.
With the genera the proportions common to both worlds would be, as before
. noted, 23 per cent. for the tropics and 42 per cent. for the cooler latitudes of
the north, and my figures suggest that north of the warm temperate region
it would be at least 50 per cent.
Rightly interpreted, there should be a great significance in the principle
that the tendency to congregate in the north on the part of plants repre-
sented in the eastern and western hemispheres is greatest and well marked
with the species and least or non-existent with the family. The connections
in the north belong to the later stages of the differentiating process, whilst
the disconnections of the south date back to remote antiquity. One would
have imagined that during the long ages that have passed the ocean-parted
eastern and western worlds would have possessed scarcely a family in
common, except in the north. Yet, as already shown, 69 or 70 per cent,
EE A oe y 5 QM R ip ee ee
TENERAS TC eae OC |. VES CHI RS io iio uies gir,
450 MR. H. B. GUPPY : PLANT-DISTRIBUTION
/
of the exclusively and mainly tropical families are common property of the
Old and the New Worlds. All the influences that have been in operation
in a differentiating world during an incalculable period of time have in a
general sense not materially defaced the primitive family type, and the
wonder is not that the differentiating agencies have done so much but that
in this respect they have effected so little.
Yet, as we have seen, about 30 per cent. of the families do respond to
the bi-cleavage of the land represented in the American and Eurasian
hemispheres. (This applies, of course, only to the families in the mass, the
proportion, as shown in a note to Table III., being much smaller, if we
regard only the principal families.) But the differentiation, or the break-up,
of the original family-type has proceeded far more rapidly in the New than
in the Old World. In its development of new families the western hemi-
sphere displays for its size nearly twice the capacity that is exhibited by
the eastern hemisphere. Of the eighty residual families that are restricted
either to one hemisphere or to the other (see Tables I., II.), one would
have expeeted the Old World to possess by far the greater number, since
the land-areas of the two hemispheres stand to each other in proportions
exceeding two to one (O.W. 35; N.W. 15; based on data given in
Whitaker's Almanack, 1917, p. 101, the polar regions being excluded,
Australia being included in the Old World). But, to one's surprise, the
difference in the number of families peculiar to each is relatively small,
45 being appropriated by the eastern and 35 by the western hemisphere.
There will subsequently be occasion to mention this point again in associa-
tion with another remarkable contrast presented by the New World with
regard to the Old World.
[One may take this opportunity to observe that the excess in families in
proportion to its area held exclusively by the New World is apparently not
exhibited to the same degree by the genera. Of the 3150 genera of the
Angiosperms named by Willis in his * Flowering Plants and Ferns’ (1908),
the Old World appropriates 47 per cent. and the New World 25 per cent.,
about 28 per cent. being held in common. Of 529 genera belonging to
42 families dealt with in the ‘Pflanzenreich’ series, 51 per cent. are peculiar
to the Old World, 32 per cent. to the New World, and 17 per cent. are
common to both. The difference between the two worlds with regard to
their peculiar families and genera may be thus expressed. With the
families the difference would be as 9 (O.W.) to 7 (N.W), but with the
genera as 9 (O.W.) to 5 (N.W.).]
We have already observed that as many as 70 per cent. of the families do
not respond to the great bi-cleavage of the land-surface of the globe,
distributed as they are in both the eastern and western hemispheres. The
response becomes greater and greater as we proceed down the seale from
the family to the species, Thus the proportions common to the two
FROM THE STANDPOINT OF AN IDEALIST. 451
hemispheres are about 56 per cent. for the tribes, less than 20 per cent.
for the genera, and about 1 per cent. for the species.: As indicated in the
table of results (Table T.), this subject has only been sampled for the tribes,
genera, and species; but reference should there be made to the accompanying
explanatory remarks. Yet the consistency in the results leaves no doubt
that the general behaviour of the tribe, the genus, and the species is correctly
illustrated in the above percentages. Here we perceive that the connection
between the Old and the New Worlds is greatest with the family, less with
the tribe, smaller still with the genus, and least with the species. Such a
result is in perfect accord with what we should expect from the successive
differentiations of a world-ranging family-type into tribe, genus, and species,
the range contracting as one goes down the scale. The effect of the opposite
method of regarding the species as diverging into the genus, the genus into
the tribe, and the tribe into the family would be chaos.
But although 70 per cent. of the families occur in both worlds they
represent in very different degrees the community in families between the
east and the west. For instance, the original distribution of the generalized
family-type in both worlds would be best exemplified now by a family of
which all the tribes belong to both hemispheres. At the other extreme the
connection implied by the community in families between the Old and
the New World would be near its breaking-point in a family where no
tribes were the common property of both hemispheres, and where most of
the tribes were gathered together in one of them. The possibility thus
presents itself of constructing a scale representing the various stages of
detachment from a both-world distribution. Taking an imaginary family
holding thirteen tribes the writer has framed such a scale, the first grade
claiming the families where all, or nearly all, the tribes are common to both
worlds, the complete detachment being illustrated in the eighth or last
grade, where all the tribes are restricted to one and the same hemisphere.
Such a grading of families would raise many difficulties, some of which
ought not to prove insurmountable for the differentiation hypothesis. It is
not possible, however, to do much more than draw attention to this method
here. It will be sufficient to mention that the Composite, the Aracez, and
the Betulaces representing, respectively, the cosmopolitan, the tropical,
and the temperate families, would find their place in the first grade. But
the anomalies of this sort that are displayed in framing such a scale are in
themselves instructive; and it is to be doubted whether a much more
effective plan could be devised to illustrate the unequal value of families
and to emphasize the necessity of grouping all families under a few great
alliances. For instance, the Scitaminez are represented in this scale by
four families, the Marantacew, the Zingiberacese, the Musacew, and the
Cannaces, which are scattered up and down its grades. From the stand-
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452 MR. H. B. GUPPY : PLANT-DISTRIBUTION
point of the differentiation theory its place would obviously be in the first
grade, together with the Composite and the Araceæ.
Having shown that the response to the cleavage of the land into two main
masses, diverging from the north polar regions, inereases with the Angio-
sperms as we go down the differentiating scale from the family to the
species, being small with the family and very pronounced with the species,
we will for a moment direct our attention to the behaviour of the great
groups, or the cohorts, that lie immediately behind the families. As might
have been expected, the response js even less than with the families. With
the families about 70 per cent. ignore the cleavage. With the cohorts, on
the other hand, 91 per cent., or 41 out of 45, are represented in both the
eastern and western worlds (Table I.). It is noteworthy that the four
cohorts that are exceptions to the rule hold in each case only a single family,
the Cyelanthacez, the Leitneriacez, the Casuarinacem, and the Balano-
psidaceze, all of them tropical, the first two belonging to the New World and
the last twoto the Old World. They are all small anomalous families which
have puzzled the systematist in his endeavours to place them. Together
they represent the flotsam and jetsam of an ever-differentiating plant-world.
The response of the families to the differentiation of the climatic zones.—
Although the families of the Angiosperms make a relatively small response
to the bi-cleavage of the land-surface of the globe, their behaviour under
the stress of climatic differentiation has been very different. From the
tables (IV. and V.) it will be seen that nearly 60 per cent. of them are
exclusively or mainly tropical, about 30 per cent. exclusively or mainly
temperate, and about 10 per cent. fairly divided between the tropical and
the temperate zones, all the regions outside the sub-tropies being regarded as
temperate. Since the differentiation theory postulates an early age in the
history of the Angiosperms when primitive generalized types ranged
the globe and uniform climatic conditions prevailed, the later ages being
occupied with the differentiation of types in response to the diversification
of climate, it follows that the results for families above given represent a
particular stage in the detachment or individualization of temperate floras.
When we look to the future and ask ourselves what will be the ultimate
result of this gradual detachment of the temperate from the tropical floras,
we shall be obliged to confess that there is little more to expect now. We
might have looked far ahead to an age when the tropical and temperate
floras would be sharply differentiated, an age when the world would be held
by complemental families representing the independent expression of tropical
and temperate conditions on the same type. But the influence of climatic
differentiation is largely played out. Nature in the development of new
forms seems to have mainly exhausted her efforts during the Upper
Cretaceous period. That which has happened since has been principally
the effect of the differentiation of ancient types in response to the progressive
FROM THE STANDPOINT OF AN IDEALIST. 453
diversification of conditions, and all we have to look for is the assertion of
man’s predominance in his replacement of natural floras by his cultivated
plants and his weeds. The time for revolutions in the plant-world was
spent geological ages ago ; and Nature in her present operations can offer
us but little aid in unravelling the revolutionary past.
The subject of the detachment of the temperate from the tropical floras
during the ages through which the differentiating influences have reigned
supreme, brings up the question of the complemental families, those which,
although now ranked as distinct families in the tropical and temperate
zones, are so closely linked together that they may be regarded as the result
of the differentiation of the same world-ranging family-type. Thus we may
view the Primulaces in the temperate regions and the Myrsinaces in the
tropics as complemental to each other and as representing the first step
in the differentiation of a common parent type. In the same way and
with the same implications we may link the Umbelliferae (temperate) with
the Araliacee (tropical), and the Chenopodiaces (temperate) with the
Amarantacez (tropical).
But the effect of the secular differentiation of climatic conditions and
of the individualization of the temperate zones has not always been the
development of a world-ranging type into separate tropical and temperate
families. Some, like the Composite, have, as far as the retention of family
characters is concerned, defied the differentiating agencies. In yielding to
the exigencies of the differentiation of climates, though still holding the
world, they have retained in this case the essential characters of the family,
on the absolute permanence of which, in the case of the Composite, Bentham
lays stress. Others like the Scitaminez are still confined to their original
home in the equatorial regions of the globe, having failed to adapt themselves
to the newly differentiated temperate zones. ‘They have given rise to
separate sub-families, often ranked as families, in the different warm regions
of the globe, but in no sense as the result of the secular diversification of
climate.
We have remarked that the work of the differentiation of floras is largely
spent, as far as climatic influences are concerned. Yet, great as this work
has been, we are, as the writer thinks, not justified in regarding it from any
other standpoint than that of adaptivity. It is not the work that was
carried on in those remote Mesozoic ages when the larger plant-groups, now
represented by the alliances of families and by the cosmopolitan and pan-
tropical families, were first developed. The charaeters that distinguished
them then distinguish them now, and as far as their essential characters are
concerned they have made but little response to the great climatic differen-
tiation of the ages. The rise of the Xerophytes presents one of the most
important and far-reaching results in the story of distribution and differen-
tiation, Yet they are of the later and not of the earlier age; and it is
2P 2
454 MR. H. B. GUPPY : PLANT-DISTRIBUTION
questionable whether any truly natural family comparable with the great
families has been developed through the changes inducing the xerophytic
organization. They might disguise them, as in the case of the Cactacere
and of some of the Euphorbiacem ; but the essential floral characters were
produced in pre-xerophytic times. A family in its truest sense is born and
not made.
It may here be added that the process of detachment of temperate families
from the tropies has not been uniform in the two hemispheres, the east and
the west, the tendency to the differentiation or detachment of temperate
families in the Old World being far more marked than in the ease of the
New World. Thus, with the exclusively Old World families 44 per cent.,
or 20 out of 45, are restricted to the tropies, using that term as including
the sub-tropies ; whilst with the exclusively New World families the pro-
portion is as high as 77 per cent. or 27 out of 35 (Table IIT.). Thus it
also appears that the tropics of the American continents actually possess a
greater number of peculiar families than those of the Old World. This
feature of American plant-distribution is to be associated with another
feature, already alluded to and illustrated in the same iable, the New World
in its entirety owning nearly as many peculiar or endemic families as the
Old World, 35 against 45, though barely half its size. In the New World,
therefore, there has been not merely a greater development of families, but
a greater segregation of such families within the tropics. In other words,
although the process of detachment of temperate families from the tropics
has been far less effective in the New World than in the Old World, the
differentiation of new families in the American tropics has been far greater.
This is one of the lessons supplied by the western hemisphere when treated
statistically from the standpoint of the differentiation hypothesis. | Reference
has before been made to the contrast in behaviour of the genera in one of
these respects. |
Before quitting the subject of the influence of the differentiation of the
climatic zones on the development of the families, a few remarks may be
devoted to the numerical distribution of the families of the Angiosperms
in the north and south hemispheres, which is illustrated in Tables VIII.,
IX. X. The matter cannot be discussed at any length here ; but it may be
observed that the numerical apportionment is much as though the land-areas
in the two hemispheres were approximately the same in extent. Yet within
the limits of vegetation there must be at least 24 times as much land in the
northern as there is in the southern hemisphere. The differentiation of
families has thus been far more active in the south than in the north, a
result that might be attributed to the much greater isolation of the southern
land-masses.
The larger plant-groups behind the fumilies.—W e have already dealt with
the cohorts in connection with the families, and it has been shown that as
T CONOCE NM
FROM THE STANDPOINT OF AN IDEALIST. 455
far as the principle of differentiation is concerned they behave conformably
with their position in the differentiating scale just above the families.
Before one proceeds to deal statistically with the large groups of the Angio-
sperms that lie behind the cohorts, a word may be said regarding the needs
of the differentiation hypothesis in this respect. Between the 45 cohorts
and the two classes of the Monocotyledons and Dicotyledons, there is a con-
siderable break or gap in the differentiating scale, which cannot be very
satisfactorily bridged over by dividing up the Dicotyledons in a few groups
as is done below. The advocate of the differentiation theory, and the writer
ventures to think that his need is also that of the systematist, requires an
intermediate group holding about a score or two dozen great alliances, each
of them cosmopolitan or pantropical, and all families of restricted range will
have to be reduced to terms of an alliance. Here geographical considera-
tions would be foremost, and the complemental families as above described
would take a prominent part in building up an alliance which would either
possess the tropics or hold the world, the pantropical and cosmopolitan
alliances being regarded as of equal value, notwithstanding the failure of the
first-named to respond to the differentiation of the temperate zone. The
object would be to restore the original world-ranging generalized types ;
and if the number of such great alliances should exceed the limits above
named, they would serve to displace the cohorts which could then be very
well dispensed with. One would commend the use of familiar designations
for all the alliances, such as Compositee, Rubiacez, Ericales, Leguminose,
Seitaminez, Aroideze, Palme, ete. But in restoring the original type in the
shape of an alliance we should give it the name of the tropical parent form
where a cosmopolitan alliance is concerned. Any names like Geraniales
and Primulales that would seem to credit the alliance with an origin in the
temperate zones should give place to the name of the older tropical forms.
The Monocotyledons and Dicotyledons.—N otwithstanding the great disparity
inthe number of families they hold (Monocotyledons 43, Dicotyledons 229),
some interesting comparisons can be made. In the first place, it wiil be
worth while ascertaining how they stand to each other with reference to the
proportion of families occurring in both hemispheres, the east and the west.
They differ but little in this respect, since the proportion is 76'8 per cent.
for the Monocotyledons and 69°4 per cent. for the Dicotyledons (Table II.).
This approximation exists in spite of the circumstance that 30 per cent. of
the Monocotyledonous families are aquatic or sub-aquatic (Table V.). Of
the 272 families of the Angiosperms 26 are aquatic or sub-aquatic, and
of these half (13) are Monocotyledons. It would therefore appear that as
regards the occurrence of a family in both the Old and the New Worlds
it behaves the same, whether monocotyledonous or dicotyledonous, or
whether aquatic or terrestrial in habit. This independence of the proportion
of aquatic plants is also displayed, as will subsequently be shown, by the
456 MR. H. B. GUPPY : PLANT-DISTRIBUTION
subdivisions of the Dicotyledons with respect to this feature in distribution.
Sympetale and Monocotyledons are distributed in the same proportions in
both worlds, as respects their families, although the first holds only 2 per
cent. of aquatic families and the second as much as 30 per cent, Yet it is
true that the large proportion of aquatic families exercises an influence in
determining the distribution of Monocotyledons ; but, as is established below,
that influence is mainly concerned in curtailing their latitudinal extension
and affects but slightly, as already shown, their response to the bi-cleavage
of the land, as indicated by the proportion of families existing in both
hemispheres.
Although the Monocotyledons and the Dicotyledons are at one in the
similar responses of their families to the great cleavage of the land, they
differ much in the responses made by their families to the differentiation of
the latitudinal climatic zones. Whilst, as shown in Tables IV. and V., both
classes hold about the same proportion of exclusively or mainly tropical
families, 98 and 59 per cent. respectively, yet in the case of the Dicotyledons
nearly all of the residue are either mainly or exclusively temperate, while
with the Monocotyledons two-thirds of the remainder are fairly well shared
between the tropical and temperate zones. Itis thus evident that as regards
the differentiation or separation of temperate floras from the original tropical
floras, the Dicotyledons are in a much more advanced stage than the Mono-
cotyledons. The equal sharing between the tropical and temperate zones of
a family originally tropical represents the first stage in the detachment of a
temperate family. The appropriation of a family by the temperate zones
represents the last stage in the detachment of a family from its original
abode in the tropics. This last stage has been attained by 21 per cent. of
the families of the Dicotyledons and by only 7 per cent. of those of the
Monocotyledons (Tables IV., V.), the last-named having lagged behind the
Dicotyledons to a marked degree as regards the differentiation or detachment
of temperate floras. The contrast may be stated in another way. Thus,
whilst 34 per cent. of the families of the Dicotyledons are either temperate
or mainly temperate, the proportion for the Monocotyledons is only 14 per
cent. This may be due to the greater prevalence of aquatic families among
the Monocotyledons. Here the proportion is as much as 30 per cent., that
for the Dicotyledons being under 6 per cent. (Table V.). The explanation
would be that aquatic conditions present a much smaller contrast between
the temperate and tropical zones than is offered by those of land plants.
The conclusions to be drawn from the behaviour of the great. plant-groups of
the Angiosperms.—Although the present arrangement of the main land-
masses and of the oceans is largely ignored by the great plant-groups, the
response becomes more and more evident as we go down the differentiating
scale. It goes without saying that in whatever way we split up the Dicoty-
ledons, whether in two or three or four groups, all the primary groups of
TUA MEN o: HE
FROM THE STANDPOINT OF AN IDEALIST. 451
the Angiosperms, commencing with the Monocotyledons and ending with
the Sympetale, take no heed of the present distribution of land and water.
But there is a slight response for the cohorts, 9 per cent. of them being
restricted to either the New or the Old World. Of the families 30 per
cent., of the tribes about 44 per cent., of the genera at least 80 per cent.,
and of the species about 99 per cent. respond to the cleavage of the land
into two main masses diverging from the north polar regions (Table I.).
This contrast in the behaviour of the larger and lesser plant-groups implies
a very great contrast in geographical and climatic conditions. There would
seem to have been a pre-differentiation era that corresponded with geo-
graphical and climatic conditions very different from those that now prevail.
At that time generalized types ranged the globe and the conditions were
far more uniform than at present. It was an age, we imagine, when floral
mutations were relatively unchecked. After that era the age of differen-
tiating conditions began, the effect of the progressive differentiation of
conditions being to restriet more and more the play of mutation in the case
of the floral organs, so that in our age the capacity is rarely exercised. In
the pre-differentiation era the generalized type had the whole earth for its
range and uniformity of conditions for its “ mise-en-scène,” a setting that
was destroyed when climate began to individualize. During such an era
reigned other things, other ways, and other conditions.
If we were to draw a line dividing this distant era from the succeeding
ages of differentiation, we should draw it just below the great family groups
as illustrated by the Compositze and the Araceze ; and if we were to contrast
the geographical conditions, we should point to the fact that whilst the
family and the groups behind it or above it mainly or entirely ignore
_the existing arrangement of land and sea, the genus and the species are in a
sense the offspring of it. Distribution in the distant past was chiefly a story
of generalized family types. In the later ages it has been principally a
story of the genus and the species and of adaptive response to the progressive
differentiation of conditions. The failure of the larger plant-groups to
respond to the great bi-cleavage of the land-mass and their subsequent
ready response to climatic differentiation mark out the two great eras—the
pre-differentiation age and the age of differentiation that followed.
What is earliest in distribution belongs to the family and the large groups
behind it. What is recent belongs to the genus and the species. To employ
the terms “ genus” and “species” when speaking of an age different in
almost every respect from the present one is to muddy the waters, or,
rather, to confuse the issues. Such a habit assumes that the present is like
the past, that we can picture what has been from what is. Yet to think
only in terms of genera and species is to ignore the better half of the story
of the development of the plant-world. The age that witnessed the rise of
458 MR. H. B. GUPPY : PLANT-DISTRIBUTION
the great families and the age that witnessed their subsequent differentiation
are things apart, and cannot be dealt with by the same method.
If one were asked how such a view of distribution could be reconciled with
that of the animal world, one might reply that since plants and animals
have been developed on quite different plans, the plant requires an applica-
tion of the Darwinian theory of evolution, in which this distinction is
recognized. In the one case development has centred around provisions
for securing the continuance of the like. In the other it has been concen-
trated on the production of a higher order of beings culminating in Man.
The first secured, the second became possible.
Note on the sub-divisions of the Dicotyledons.
In order to make a further statistical comparison of the Dicotyledons
with the Monocotyledons, the number of families in the first being
according to Engler's system more than five times those of the second,
the Archichlamydez have been broken up into four groups, making with
the Sympetale five groups for the class. For this purpose the system of
Engler was preferred, since a scale of development is implied in the
arrangement adopted ; whilst with that of Bentham and Hooker the placing
of more than a fifth of the families there recognized in the Incomplete
makes a linear classification impracticable. Yet it was the Incomplete that
led the writer to pay attention to this matter, with the result that the group
viewed from this standpoint appears to be very far from an anomalous
group. On the contrary, when treated statistically it proves to be the most
typical, as far as percentages are concerned, of all the groups of the
Dicotyledons.
Assuming that the series—Monocotyledons, Archichlamydes, Sympetale
—represents a scale of plant-development and that the same is indicated in
the arrangement of the groups of families of the Archichlamyde:e, the writer
broke up the last-named into four groups and obtained the following
succession :—
Monocotyledons, holding 43 families.
( Group A. Cohorts 1-14 with 37 families.
Ec p TOR. 2 4d a
Aychichlamy deze J Sid s dba os
Dicotyledons E. D sc X42 8
Sympetale, holding 51 families.
Group A includes 25 of the 37 families in Monochlamyde: or Incompletze
of Bentham and Hooker. Group B comprises the cohorts Ranales,
Rheeadales, Sarraceniales, and Rosales; Group C, the cohorts Geraniales,
Sapindales, and Rhamnales ; and Group D, the cohorts Malvales, Parietales,
Opuntiales, Myrtiflorze, and Umbelliflore.
Together here we have six groups which we will term the Primaries, and
He
FROM THE STANDPOINT OF AN IDEALIST. 45
in order to give point to their statistical treatment we will assume the truth
of the implication of Engler’s system that they represent a genetic sequence
commencing with the Monocotyledons and terminating with the Sympetale.
It would have been possible to discuss this matter at considerable length;
but as the treatment is purely tentative a few general remarks will be here
sufficient, and the columns of the tables (II., IV.-VIII.) will be allowed
largely to tell their own story.
It will be at once noticed that whilst the six primary groups of the
Angiosperms behave with comparative uniformity in matters concerned
with the bi-eleavage of the land-mass, as reflected in their distribution
in the Old and New Worlds, they often present marked contrast in their
responses to the differentiation of the climatic zones. Thus, to take their
behaviour in the first case, the proportions of families occurring in both the
east and west hemispheres vary only between 62 and 77 per cent. (Table II.) ;
and if we limit the comparison to the families of world-wide distribution,
termed cosmopolitan in the table, the percentages range only between
21 and 33 (Table VII.). We find a like agreement in the proportion
of families confined either to the Old or to the New World. Thus, the
percentage of families restricted to the eastern hemisphere varies only
between 12 and 24 and of those peculiar to the western world only between
ll and 14 (Table IL). This conformity is remarkable when we reflect that
in small groups of this kind we can only appreciate general approximations
or marked deviations. The similarity in behaviour on the part of the six
primary groups respecting the distribution of their families in the eastern
and western hemispheres is quite independent of the proportion of aquatic
and sub-aquatic families in each group, which is as high as 30 per cent. for
the Monocotyledons and as low as 2 per cent. for the Sympetalze (Table V.).
Yet this similarity in behaviour disappears when we regard the response
made by these six primary groups to the differentiation of the climatic zones.
As indicated in Table V., they display great variety in the appropriation of
their families by the tropical and temperate zones. Thus Groups C and D
of the Archichlamydez are the most tropical; and Group B and the
Sympetale are the least tropical; whilst the Monocotyledons and Group A
stand between. On the other hand, the Monocotyledons are by far the least
temperate of all the groups, which is to be associated with the fact that
a much larger proportion of the families are in a transition state—that is, are
equally divided between the zones—than is the case with the other groups.
Then again Group A is in these respects the most average of the six
primary groups, approaching nearest in its behaviour to that of the
Dicotyledons in the mass and nearer still to that of the Angiosperms. It
comes closest to the Incomplete of Bentham and Hooker, a result to be
expected since the two equal-sized groups hold about two-thirds of their
families in common ; whilst the group of the Incompletz in its response to
460 MR. H. B. GUPPY : PLANT-DISTRIBUTION
the differentiation of the climatic zone is the most typical of all, making a
near approach to the Dicotyledons in their entirety and the nearest of all to
the Angiosperms in the mass (Table V.).
Comparison may now be made between the Monocotyledons and the
Sympetale as representing the extremes of the series constituted by the six
primary groups. It has been shown in an earlier page that with regard to
the detachment or differentiation of temperate floras from the original
tropical flora the Dicotyledons are far in advance of the Monocotyledons.
But, as indicated in the columns of Table V., the five groups of the
Dicotyledons exhibit considerable divergencies in their behaviour in this
respect ; and it is not possible to construct a series with the Monocotyledons
and the Sympetale at the extremes, Yet in view of the position of these
two groups at the extremes of the series accepted by some systematists,
a brief comparison may be profitable. The escape of the Sympetalie from
the tropies, as contrasted with the lagging behind of the Monocotyledons, is
illustrated in different fashions in the columns of Tables IV., V., and VII.
Nearly all the tropical families of the group last named, that is 23 out of 25,
are exclusively tropical. With the Sympetale only 14 out of the 25
tropical families are exclusively tropical. Then, again, whilst 12 out of the
43 families of the Monocotyledons are fairly well shared between the tropical
and temperate zones, with the Sympetale the number of families shared is
4out of 51, or only about 8 per cent. The bulk of the non-tropical families of
the Monocotyledons are in fact in the transition state. With the Sympetalie
the non-tropical families have in most cases reached a further stage and
are more or less completely detached from the tropics. Whilst with the
Monocotyledons the families exclusively tropical amount to 53 per cent.
und those exclusively or mainly temperate to only 14 per cent., with the
Sympetale only about 27 per cent. of the families are confined to tropical
regions and as many as 43 or 44 per cent. are exclusively or mainly
temperate. A possible explanation of the contrast in behaviour between
these two groups is supplied in the great predominance of families of
aquatic habit in the Monocotyledons (see Table V.); but this is a matter
discussed in the following note.
Note on the influence of aquatic and sub-aquatie families on the distribution
of the Angiosperms.
The apportionment of the families of aquatic habit between the larger
groups of the Angiosperms and their distribution over the eastern and
western hemispheres are illustrated in Tables V. and VI. Of the 26 aquatic
families recognized by the writer, 13, or half, are appropriated by the
Monocotyledons, 12 belong to the Archichlamydew, and only 1 to the
Sympetala, namely the Lentibulariaceæ. It has been shown that it makes
FROM THE STANDPOINT OF AN IDEALIST. 461
but little difference in the distribution of the larger family-groups in the
eastern and western hemispheres whether they hold many or few aquatic
families. Thus the Monocotyledons holding 30 per cent. of these families
and the Sympetale holding only 2 per cent. are distributed in the same
proportion over the Old and New Worlds, in each case about 77 per cent. of
their families being common to the two hemispheres (Table II.). The
aquatic families are therefore quite neutral in their influence on the general
response of the Angiosperms to the great bi-cleavage of the land-mass of
the globe.
On the other hand, they sometimes seem to have a marked influence on
the response made by families to the differentiation of the climatie zones.
Thus, as shown in the previous note, the Monocotyledons lag behind the .
Sympetale to a marked degree in the detachment of temperate families from
the tropics ; and the implication is that since the former hold as many as
30 per cent. of aquatic families, and the latter as few as 2 per cent., the
influence of the aquatic habit in checking the process of differentiation
is displayed in the diminution of the climatic contrast between the
tropical and temperate zones. Yet, although this may sufficiently
explain the lagging of the Monocotyledons in the tropics as compared
with the Sympetale, it will not explain why amongst the primary groups
of the Archichlamydez those that are most tropical, like © and D, hold
the smallest number of aquatic families (Table V.).
Note on the relative proportions of * both-world " families in the tropics and in
the extra-tropical regions of the northern hemisphere.
The statement on page 447 that 61 percent. of tropical families and 64 per
cent. of northern extra-tropical families occur in both the Old and the
New Worlds is based on data given in Table III. Here we are concerned
with families purely tropical and purely northern extra-tropical. But for
the statement that follows it the data are only partially supplied in the
Tables, as in IV., V., etc. It is there asserted that 69 per cent. of
exclusively or mainly tropical families and 77 per cent. of exclusively or
mainly extra-tropical families in the northern hemisphere are found in both
the Old and the New Worlds. But to obtain these results i& was necessary
to eliminate the southern extra-tropical elements ; and to avoid the necessity
of giving another complicated set of tables I have here given the data on
which this assertion is based. They are as follows :—
The total of 153 exclusively or mainly tropical families is made up of 120
exclusively tropieal and of 53 tropical and north temperate families but
mainly tropical. (In these connections it should be sta'ed that the tropies
include the subtropics and the north temperate all the northern extra-tropical
regions.) Of the exclusively tropical 73 and of the mainly tropical all oceur
462 MR. H, B. GUPPY : PLANT-DISTRIBUTION
in both worlds. This gives a total of 106 both-world families out of a total of
153 mainly or exclusively tropical families, or 69 per cent.
The total of 62 families exclusively or mainly extra-tropical in the
northern hemisphere is made up of 36 exclusively north temperate and
26 north temperate and tropical but mainly north temperate. Of the
exclusively north temperate 23 and of the mainly north temperate families
25 exist in both the Old and the New Worlds. This gives a total of
48 both-world families out of a total of 62 northern extra-tropical families,
or 77 per cent.
It is worth noting the manner in which, according to these results, the
both-world families tend to congregate in the intermediate region between
the tropies and the north temperate zone—that is, among the tropical and
north temperate families that are mainly tropical and among the north
temperate and tropical families that are mainly north temperate. Con-
siderable importance attaches itself to the interpretation of this tendency.
TaBLE I.
(See Explanatory remarks.)
A.—The distribution of the Angiosperms between the Old and New World.
Primaries. Cohorts. Families. Tribes. Genera. Species.
No.| °/, |No.! h [No] % |No.! % | No. | % | No. | %, |
|— DB ESS RSR S R " | PEOS |
|! Old World 2, 45, 45| 165 | 39! 94:5 | 570 | 46| 9889| 52
New World....| .. | .. | 2 45.) 35| 190 | 31| 19:5 | 519 | 41| 8874. 47
Both Worlds .. .. 100 41 9L0 192 705 80 560 168 13 197 1
6 100 45 100 272 100 159 100 1259 100 18060 100.
B.—Composite excluded.
Tribes. Genera, Species.
| N o " A No s. Né 0
| NERE OE mU
Old World .......... 38 26 970 | 51 5031 52:6
New World ........ 3 21 Noi op g9 4411 46:0
Both Worlds ........ 77 53 B0 be T 134 l4
146 100 539 | 100 9576 100 —
FROM THE STANDPOINT OF AN IDEALIST. 463
C.— Composite alone.
Tribes. Genera, Species.
Noc d Nor ey, No. Hs
Bloadn E E 1 309 49 4858 518
New Wod e ; 843 47 4463 47:5
Bock Wor ....,... 12 78 1] 63 (Uf
13 | 730 | 100 | 9884 | 100
D.— Genera.
No. ur.
Old World... 95. 1472 46:7
Now World: 3 | 800 254
Both Worlds; 1,1 | 878 91:9 |
3150 100
Explanatory remarks.—With reference to sub-table A it may be remarked
that the materials are complete for the larger groups, the primaries, the cohorts,
and the families. For the tribes the matter has been sampled. In this
respect the data for 23 families and sub-families were tabulated in a paper by
the writer published in the ‘Transactions’ of the Victoria Institute for 1907.
Those for 19 others have since been added, making a total of 42; and on these
the results for the tribes, genera, and species are based. The volumes of
the * Pflanzenreich’ series have here been used, except with the Com-
posite, where Bentham has been followed. In the early paper only the
tribes designated as such were utilised. In this sub-table all intermediate
groupings between the genera and the family are employed under this head
where it is practicable.
In sub-table B all the materials in sub-table A are used with the
exception of those for the Composite, which are treated separately in
sub-table C.
In sub-table D are given the results of an analysis made by the writer of
the list of chief genera (about 3150 for the Angiosperms) that is given by
Dr. Willis in his ‘Flowering Plants and Ferns’ for 1908. The increase in
the percentage of genera common to the eastern and western hemispheres
is due to the fact that the list is only concerned with “chief” genera;
and it is obvious that the number of genera with limited distribution
that are here excluded is very large, and would have gone far to pull
down the percentage of both-world genera. Probably in a complete list
of genera the percentage found in both worlds would be nearer that for
the Composite, where the proportion is 11 per cent.; and there is there-
fore much to support the estimate adopted in this paper of “less than
20 per cent."
NE STREET
MT AR
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PLANT-DISTRIBUTION
MR. H. B. GUPPY
46
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FROM THE STANDPOINT OF AN IDEALIST. 465
TABLE III.
The Families of the Angiosperms distributed according to hemispheres
(east and west) and to latitudinal zones.
(For the sake of comparison the results for the Cohorts are given
in the last column.) i
|
No. | Percentage. | Cohorts.
| US Ed 2d : | |
| A | | L8
E "TIENE eror | 70 957 | 96— 578°, |
Ds TrOpION 5 D NOM DEI eere 73 26:8 | ll= 245
| n "y m | Tropics and North Temperate .. 15 bb d. I= 29:2
| East e We t ] Tropics and South Temperate .. 8 3:0 | |
| He RA ter 9^ || North and South Temperate ....| 14 DIET 1g a LE]
prm | North Temperaten oi: nn aiee 9 $3, |
Kl South Temperate 2. 50 3 | 11 | ^ Tee 09
| |
| 192 70:5 Alz.91295 |
| |
| (| General |
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| Old World J Tropies and North Temperate on
or East || Tropics and South Temperate ..| 5 18
| Hemisphere. || North and South Temperate ....| 2 0-7
P Dp
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| U South Temperatat t scans. 10 | B
| 45 | 165 |
| | |
| | |
| (| General
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New World | | Tropics and North Temperate ..| 1 0:4
or West — 4| Tropics and South Temperate ..| 1 0s |
Hemisphere. | | North and South Temperate .... | n
| North ‘Temperate! 7. . acie s aee 11
| U South Temperate: >... janes 3 | Th |
| |
|
| |s5 80 | |
| | 272 | 10 | 45-10 |
|
Norr.—The Tropics include the Subtropies, and the Temperate zones include all extra-
tropical regions. It is important to remember that the families are here treated in the
mass, the small with the large. Practically all the great families would be represented
in both the east and west hemispheres. The author has made a list of the principal
families, those holding the largest number of genera and species. They number
about 90, and at least 95 per cent. of them are found in both hemispheres. The
distribution of families in the north and south hemispheres is also dealt with in
different fashions in Tables VIIT., IX., X,
466 MR. H. B. GUPPY : PLANT-DISTRIBUTION
|
|
j
|
EC Equally
PUN ME Temperate | |
Exclusively _ between and tropical Exclusiv ely| $
tropical. E: tropical and but mainly | temperate | To tal,
tropical. y temperate | temperate. | |
zones, |
LORI GS dettes » " [e]
A} 17 4 4 4 8 | 37
Archi- B! 15 5 | 4 ioe | “10 41
chlamydeæ. ) C | 18 | 11 l I 11 42
D| 33 EST 3 5 | 10 58
pre — — — ——
Total ....| 83 (46: 6'/) 97 (152) 12 (67°) 17(96',) | 39 (9199) | 178
—— i 3 SEN |
| Sympetale .| 14 (27°5°/ ) 11 (21° 6*1). 4(T8, |12 (28°5°/,) | 10 (19°6°/,)| 51
| Dieotyledons ..| 97 SN 38 aee, 16(70,) | 29 (12 704) 49 (21:4*/,) | 229
| | !
Poe see | mm SIUS MEM
Monocotyledons .| 93 (53°5°/,) | 2 (4°6°/ 12 | 12 (27: 9l) | 3 (r0) | | 9(r0 | 48
| Angiosperms ../190 (441°/,) | 40 (14 zn 28 (10:3*/) E al: 879 | 52 (19°1°/,) | 972
B. Cohorts.
: | |
| Angiosperms 10 (22-2) Io] 12 (26:7°/,) | 17 (37°8°/,)| 4(89/) | 2(4-49),) | 45
Tabi LV,
Distribution of the Families and Cohorts of the Angiosperms according to
climatic or latitudinal zones.
(This Table is supplementary to Table V.)
A. Families.
|
|
|
Explanation.—Engler’s system is followed. The temperate include all extra- -tropical
regions, and the subtropics are comprised in the tropics. Cohorts 1-14 are placed
Table V. should be
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468 MR. H. B. GUPPY: PLANT-DISTRIBUTION
Explanation of Table V.
The system of Engler is adopted, the Incompletze of Bentham and Hooker
being added for the sake of comparison. The Archichlamydem have been
divided by the writer into four groups :—A.:- cohorts 1-14; B.= cohorts
15-18; C.— cohorts 19-21; D.=cohorts 22-26. The subtropics are included
in the tropies and the extra-tropical zones are classed as temperate,
The general results are given in the first series and the details in the second
series. Thus it is there shown that in the first series the tropical families
include those that occur also in temperate regions, but are mainly tropical,
and the same with the temperate families, which include those that are
mainly as well as those that are exclusively temperate. The third series
deals with the aquatic and subaquatic families, concerning which other
particulars are given in Table VI.
In the first series the absolute numbers are enclosed in parentheses. In
the second series they are omitted, but they are given in Table IV.
By following Engler's system in the cases of the Cytinacez, the Cupu-
liferze, and the Piperacez of the Incomplete the number of families has been
inereased from 36 to 39 for that group.
Taste VI.
Distribution of Families of aquatic and subaquatic habit in the Old and
New Worlds.
Old and New Worlds, Old World. | New World. Total.
| Monocotyledons ...... 11 1 l 13
Archichlamyde A ; a 1 l
A Ba 3 3 1 7
| L p 1 | 1 2
| s Bui 2 2
| Eympeatale- at. issos: 1 |
ES j ee T. MES |
| 18 4 4 26 |
Other details are given in Table V. and the groups A, D, C, D, are
there explained,
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470 MR. H. B. GUPPY : PLANT-DISTRIBUTION
Tasty VIII.
The distribution of the Families of the Angiosperms in the North and
South Hemispheres.
(Engler's system is followed. In the subdivisions of the Archichlamydeæ,
as adopted by the writer, group A. holds cohorts 1-14 ; B. 15-18 ; C. 19-21 ;
and D. 22-26. The tropics include the subtropics, and the temperate zones
signify all extra-tropical regions.)
Archichlamydee. |
|
Minos. | i endo Eur Sym- | Dico- ert errno
cotyledons. | All4 petale. | tyledons. ^ uui
| B. | C groups. | | |
Cosmopolitan .... 14 8|11, 9.19 457 [57-45 56 10—95'7^/, |
North and South | | : | TNT | | EOS UNUS
Temperate ..| | 3 2 | BrE T. 8 | 5 | 13 | 16— 59
North Temperate..| .. | 2) 4) j 5 gj | a 30 | 90— 74
d ril Lu revpe m wp os ]i | 16— 59
Tropics ,... ese | 93 |17|15|18,88| es | 14 | 97 | 120-441
Tropics and South | | « | | | | | x:
Temperate ..| | 3 | E 8.1 | 1 5 | 6 | M og 14— 51
South Temperate . | E |.4| 81 8|.5 IE 1 1 | 16 | 16— 59
43 | 37| 41/42/68 178 51 | 229 279—100 |
TIBLE IX.
The distribution of the 272 Families of the Angiosperms in the North and
South Hemispheres.
Cosmopolitan... isses nnn 70—25°7 °/,
, North and South Temperate ......... ees 16— 5'9°/,
| Exclusively or mainly North Temperate ....| 26— 96 fo
| Exclusively or mainly Tropical ......... | 139—51:1 9/5
| Exclusively or mainly South Temperate ... | 91— 1 ^l
272 —100:0 */,
NoTE.—The Angiosperms are here treated in the mass as in the last column of Table VIII.,
but with a little different arrangement, All extra-tropical regions are classed as
temperate,
i
$
:
pe. RTE ET” SEE SG Tert eol. 0o Cee i EE eee S
FRÓM THE STANDPOINT OF AN IDEALIST. 471
TABLE X.
The distribution of the 272 Families of the Angiosperms in the North and
South Hemispheres.
(The data in Table IIT. have here been utilised. The tropics include
the sub-tropies, and all other regions are classed as temperate.)
1 ae?
|
North Temperate in varying degree ........ | 122 or 45°/,
Tropical Sip DR aid p 220 or 81 9/5
South Temperate ,, bs ROREM 116 or 45 9/,
Norr.—The total number of families represented to a greater or less extent in temperate
regions, that is in regions outside the tropies or subtropics, is 152 or 56 per cent. The
representation in the tropics is above shown to be 220 or 81 per cent.
SUMMARY.
From a consideration of the problems of plant-distribution, the writer is
led to regard the history of the Angiosperms as resolving itself into two
principal eras :—
(1) The era that witnessed the rise of the great families, a period of
relatively uniform conditions.
(2) The era that witnessed the differentiation of these family types in
response to the differentiation of the climatic and other conditions.
It is argued that conclusions drawn from the prevailing influences now in
operation could only be applied to the differentiation of the ancient family
types—that is to say, to the second era in plant-history. It is not possible,
so it is held, to apply a theory based on the present to an age of other things,
other ways, and other conditions. Only the hypothesis that finds its guide
to the past in the abnormalities of the present can be of service to us in the
interpretation of times so different.
The subject is introduced by a reference to two papers, contributed to the
*Journal of the Linnean Society, which have an important bearing on
the subject, the one by Bentham on the Composite, the other by Huxley
on the Gentians. Then follows a statement of the differentiation hypothesis
which involves the differentiation of primitive world-ranging types in response
to the progressive differentiation of their originally uniform conditions. Allu-
sion is then made to the dilemma into which all theorists fall when they come
to handle the larger groups, the very persistence of whieh in our own age
depends on the stability of their essential characters. If stable now, why so
unstable then? We are thus forced to the conclusion that in the distant era
oe eae
472 MR. H. B. GUPPY ON PLANT-DISTRIBUTION.
.
that witnessed the deployment of the Angiosperms instability prevailed.
It was an age of mutations, free and unchecked, and an age of uniformity of
conditions, the mutability decreasing and the modifications becoming more
and more fixed with progressive differentiation of conditions, an explanation
suggested by a perusal of the accounts by Dr. Willis of his prolonged
investigation on the Podostemace:e.
The distribution of families is then treated statistically ; and it is shown
that whilst they largely ignore the cleavage of the land into two great masses
diverging from the north, they respond in a marked degree to the differentia-
tion of the climatic zones. Behind their disregard for the present arrange-
ment of continents and oceans lies the story of the first era, and behind their
ready response to climatic differentiation lies the story of the second era.
In the circumstance that the response made to the bi-cleavage of the land-
mass is absent or small with the larger groups and becomes greater and
greater as we go down the differentiating scale until it attains its maximum
in the species, is recognised the contrast of conditions between the pre-
differentiation era and the era when differentiation reigned supreme. It is
held that there isa method here disclosed that could only arise by the family
differentiating into the tribes, the tribe into the genera, and the genus into
the species, since the opposite method of commencing with the species would
produce chaos.
The paper ends with the application of the statistical treatment to the
larger groups behind the families, and it is shown that whilst the Dico-
tyledons display a much greater tendency to detachment from the tropics
than the Monocotyledons, the Sympetale stand foremost in this respect
amongst all the groups of the Dicotyledons. It may be added that there
is a large amount of material in the ten tables which from considerations
of space could not be discussed. These data have therefore to tell their
own story.
GO MAS TRO Y P'E
À MALAY FORM OF CHLOROCOCCUM HUMICOLA. 473
On a Malay Form of Chlorococcum humicola (Nüg.), Rabenh. By B. Murren
Bristot, M.Sc. (Communicated by Prof. G. S. West, F.L.S.)
(Prates 17 & 18.)
[Read 21st March, 1918.]
I. HISTORICAL.
CHLOROCOCCUM HUMICOLA was first described by Nügeli in 1849, under the
name Cystococcus humicola, Nüg.*, as a spherical unicellular alga completely
saturated with chlorophyll, except for a single lateral coloufless space, and
containing a single pyrenoid. He described its multiplication as being
by non-motile gonidia set free by a splitting of the mother-cell-wall, but
did not observe any motile cells in connection with the alga. In 1868,
Rabenhorst f. identified the genus Cystococcus, Nüg., with that which Fries
had deseribed, in 1825, as Chlorococeum, in which multiplication by biciliate
zoogonidia had been observed ; and this alteration has been supported by the
great majority of later botanists. De Toni ft, however, in 1889, included
the genera Chlorococeum, Fries (1825), and Cystococcus, Nüg. (1849), in the
genus Protococcus, Ag. (1824), with which he considered them synonymous.
Wille’s recent researches§ on Prof. C. A. Agardh’s original specimens in
the Lund Botanical Museum show, however, that Protococcus viridis, Ag.,
is identical with the alga later described as Pleurococcus Nägelii, Chod., and
that De Toni’s diagnosis of Protococcus viridis, Ag., to include Chlorococcum
humicola (Niig.), Rabenh, is quite wrong, since the formation of biciliate
zoogonidia has been definitely established in this last species.
Wille shows in the same paper that the alga described and figured by
Meneghini, in 1842, under the name Chlorococcum Monas (Ag.), Menegh. ||,
cannot be the same as Agardh’s Protococcus Monas, because Agardh's
species contains no pyrenoids, whereas, both in the description and in the
figure, Meneghini lays partieular stress on the clear spot in the periphery,
by which only a pyrenoid can be intended. Wille suggests further that
Meneghini's species was probably the same as Chlorococcum humicola (Nag.),
Habenh. If this is so, and it seems quite possible, Meneghini failed to
observe the great variation in size of the vegetative cells, and only an
examination of Meneghini's original material could prove the truth of
Wille’s suggestion; but, in any case, the specific name humicola has been
* Nägeli, ‘Gattungen einzelliger Algen, Zürich, 1849, p. 85, tab. iii. E.
T Rabenhorst, Fl. Eur. Algar. iii. 1868, p. 57.
t De Toni, ‘Sylloge Algarum,’ 1889, vol. i. p. 699.
§ Wille, N., * Algologische Notizen, xxii Christiania, 1913.
|| Meneghini, J., * Monographia Nostochinearum Italicarum.” Aug. Taurin. 1842, S. 28,
tab. iii. fig. 1.
SORE SANE | CB UNESCO T Me
414 MISS B. MURIEL BRISTOL ON A
in use for so many years that it must necessarily stand in preference to
Monas.
With the exception of the observation that the fully developed cells of
Chlorococcum humicola are conocytie *, no further study of this alga has
been carried out, and our knowledge of the species is limited to Rabenhorst’s
description given in 1868. Recently, however, an opportunity has been
afforded for a prolonged study of this alga, and the life-history has proved
to be far more complicated than Rabenhorst’s description would lead one to
expect.
II. CULTURES.
In October, 1915, a series of cultures was set up with a view to determining
what algæ, if any, ean live in soil in a resting-state, about 60 samples of soil
being taken for experiment. In connection with this work, Professor G. 8.
West very kindly provided a specimen of soil from Kajang, near Kuala
Lumpur, Malay States, which had first been air-dried and then stored
in a closed specimen-tube for about two years. Three cultures were made
from this soil, two in 50 c.c. conical flasks provided with cotton-wool plugs,
and one in a small glass box. A sterilised mineral salt solution was placed
in the sterilised vessels and about a cubic centimetre of the soil was
introduced by means of a sterilised spathula. The cultures were placed
under a glass jar in a north window, and left to grow at the temperature
of the room.
The composition of the culture-solution was as follows, and as evaporation
took place the eultures were watered with a solution of half the strength.
Potassium dihydrogen phosphate (KRH,PO,) . . 1:0 gm.
OCI nirare (NANON. ees AS qe uL vc.
Oalerum chloride (GR). 22 lale cL. c» e DD. o
Magnesium sulphate (MgSO)) . . . . . . . 03,
Sodium chloride (NaCl) Wi eu ee ee ey ys
Ferric chloride (FeCl;) Xe phan uM. D EM
Distilled water . . . Ei. lice uo ESTAS
No growth whatever took place in the cultures until nearly eight months
after their being set up, but in about the middle of June, 1916, a thin green
scum appeared round the edges of the culture-fluid. This indicates that the
initial temperature required for the germination of the alga-spores in this
Malay soil is very much higher than that required for the germination
of algæ in this country since cultures of English soils set up towards the
end of November showed considerable growth by the end of April. The
alga grew abundantly, forming not only a green gelatinous scum on the
g
gr
* West, G. S., * Algæ, Camb. Bot. Handbooks, vol. i. 1916, p. 211.
l
d
;
1
i
MALAY FORM OF CHLOROCOCCUM HUMICOLA. 475
sides of the vessel, but also flat green strata floating about in the liquid.
Repeated examination has shown that the three cultures are unialgal
éultures of Chlorococeum humicola (Nig.), Rabenh., and an excellent
opportunity has thus presented itself of making a study, not only of the life-
history, but also of the cytology of the alga.
III. Tug VEGETATIVE CELL.
The ordinary vegetative cells of the alga are spherical or subspherical, and
may be either solitary or collected together into small globular clusters or
into flat expanded strata in which the cells are imbedded in mucus. One
very important characteristic of the alga is the enormous variation in the
size of the vegetative cells which make up a stratum. A great many appear
to be fully grown when they have reached a diameter of less than 20 p, while
others continue to grow until a diameter of as much as 80, has been
attained.
A vegetative cell of diameter about 13 shows a very characteristic
structure (Pl. 17. figs. 1-4). The wall is thimand is composed of cellulose, and
it is usually of a uniform thickness throughout, though occasionally one
or more slight thickenings may be seen (fig. 3). There is a single parietal
chloroplast, which may take the form of a deep green layer lining practically
the whole of the cell-wall with a single small colourless area on one side
(fig. 2), or it may be irregular (fig. 1), having, in some cases, almost the
appearance of being divided up into several smaller chloroplasts. A large
pyrenoid is very conspicuous in the chloroplast, and in some cells, particularly
in the larger ones, two or even as many as half a dozen may be seen, though
the latter is exceptional. Starch is present in the form of minute granules
scattered throughout the cytoplasm as well as in the starch-sheaths of the
pyrenoids. As the cells increase in size, oil is formed and stored in the cells,
so that their structure cannot be so clearly made out. In the very large
cells, the chloroplast is usually continuous over the whole surface of the
cytoplasm with only one, or perhaps two, perforations, and the pyrenoids
are less conspicuous, the whole contents of the cell being masked by the
presence of a great quantity of yellow oil and by the frequent development
of a bright red pigment dissolved in the oil. When a cell has attained its
full size, the cell-wall, which up to this time has remained quite thin, begins
to thicken and become stratified, and a number of button-like excrescences
are formed usually on the inside but occasionally on the outside of the wall.
But in no case does the wall ever become thicker than about 4, and the
number of striations is usually few. «
Stained preparations of the material were made by fixing with Bouin's
solution and staining with Heidenhain's iron-alum-hematoxylin and with
Delafield^s haematoxylin. Comparatively little information could be obtained
476 MISS B. MURIEL BRISTOI. ON A
from a study of the alga stained in bulk, hence the material was imbedded in
paraffin, and microtome sections were eut about 5 w in thickness.
Sections stained with Delafield’s hematoxylin show that the cytoplasm has
a somewhat reticulate structure, but that the reticulations have no definite
radial arrangement; and in a few cases there can be seen an indication
of a parietal chloroplast. In sections stained with Heidenhain’s iron-alum-
hematoxylin the reticulate nature of the cytoplasm is not nearly so notice-
able, and the spaces between the network are seen to be filled with a granular
substance, Pyrenoids, with either spherical or polygonal pyrenocrystals and
usually narrow starch-sheaths, are conspicuous, there being often many in
the larger cells and at least one in even the smallest cells. The adult cells
are hunet (PI. 18. figs. 18 & 19), the nuclei being situated at the angles
of the retieulum, but they are so minute that they can only be clearly
distinguished under a magnification of 1435. Then, each is seen to consist
of a single small granule of chromatin, the karyosome, surrounded by
a narrow but definite nuclear space which is bounded on the outside
by an extremely thin nuclear membrane. The whole structure is not more
than 1,4 in diameter, and even under a magnification of 2820 no further
details are apparent. In very young cells there is a single nucleus exactly
similar in size and structure to those of the mature cells, to which it evidently
gives rise by repeated division; but the structures are so minute that it has
been impossible, even in a single instance, to obtain any of the details of
division,
The division of the pyrenoids usually takes place by constriction, though
multiplication by fragmentation has been observed ; and all the available
evidence indicates that nuclear division and the division of the pyrenoids are
entirely independent of one another.
IV. MULTIPLICATION BY ZOOGONIDIA.
Multiplieation by zoogonidia has been observed in cells of all sizes from
about 20 win diameter upwards, Successive bipartition of the contents of
the cell takes place, with the formation of 8, 16, or an indefinite number
of small oval uninucleate bodies, according to the size of the cell. These
acquire cilia and swarm about within the mother-cell until an aperture
appears at some place in the mother-cell-wall, when they make their way
through the aperture aud escape into the surrounding water. The zoogonidia
Mi diinad in this way are extremely variable in size even from the same
zoogonidangium, but this may be due to their being eultural forms produced
lider ; slightly abnormal conditions. Most are about 8 p long with a breadth
of about 4°5 u, but they seem to be somewhat contractile, and many are found
which are about 13 long and not more than 25g broad. They have two
equal cilia, and are usually oval or pear-shaped, while the longer ones are
UTUOTQQNMNEN S C HERE CS S D
MALAY FORM OF CHLOROCOCCUM HUMICOLA, 477
sometimes pointed at both ends. Each has a single bell-shaped chloroplast
fitted against the posterior end of the zoogonidium, though in the case of a
few elongated zoogonidia the posterior end was found to be colourless, the
chloroplast being in the form of a parietal band round the middle of the cell.
There is a red pigment-spot near the anterior end of the cell and a single
pyrenoid in the chloroplast. In some cases, owing to cultural conditions,
the escape of the zoogonidia was observed before the division of the mother-
cell-contents was complete, and a number of extremely abnormal zoogonidia
were formed ; one of these is shown in fig. 14 4. with three pairs of cilia,
three pigment-spots, and three distinet chloroplasts.
When the zoogonidia have made their escape from the mother-cell, they
swim about for a short time and then frequently fuse in pairs, either with
one another or with zoogonidia from another mother-cell. Two zoogonidia
become entangled by their cilia and fusion takes place from the anterior end
in the usual way to form a zygote having four cilia attached at the anterior
end. One case, however, was observed in which the fusion took place in
such a way as to produce a zygote with two cilia at each end, while in
another a normal zygote was seen with a small zoogonidium partially fused
to its posterior end. As a general rule, the zoogonidia were of the same
size, but a few cases were observed in whieh fusion took place between
anisogametes.
A fusion of this kind does not seem to be essential to the life-history
of the alga, for a great many zoogonidia, after swimming about for some
time, come to rest and develop in exactly the same way as the zygotes. The
cells round off, lose their cilia, and, acquiring cellulose cell-walls, develop by
gradual stages into adult vegetative cells.
V. MULTIPLICATION BY APLANOSPORES.
In certain circumstances a vegetative cell produces non-motile gonidia
instead of zoogonidia. The nuclei of the mother-cell appear to increase
considerably in size and number, and each becomes the centre of a little
mass of cytoplasm which forms an aplanospore. A great many are produced
from a single mother-cell and they become angular by compression. In
stained sections of cells about to form aplanospores the nuclei are very
conspicuous and the nuclear space is seen to be finely granular with an
occasional larger granule of chromatin. The karyosome is almost central in
position, and may be spherical or oval, or somewhat elongated with a slight
constriction in the middle. In these cells the nuclei have been observed to
divide by constriction (fig. 21), and it may be that the irregularities in shape
of the karyosome are preliminary stages in the division of the nucleus. The
occurrence of this form of nuclear division in Chlorococeum is surprising in
view of the fact that mitotic figures have been obtained for both of the allied
418 MISS B. MURIEL BRISTOL ON A
genera, Characium* and Chlorochytrium t, but it points to the fact that either
the conditions were abnormal or else the genus Chlorococcum is somewhat
primitive.
The aplanospores thus produced remain enclosed for a considerable time
within the old mother-eell-wall, which gradually disintegrates to form a
mucilaginous stratum in which the young cells are imbedded, and the alga
enters into a palmella-like condition. In this state further division of the
young cells into two or four daughter-cells may take place by successive
bipartition, and strata of considerable extent may be formed; but no
cellulose walls appear, and the cells remain imbedded in a mass of mucilage.
A sudden change in external conditions, with its accompanying physiological
shock, induces the formation of cilia either at once or after one or two
further divisions, and the cells, becoming motile, swim away rapidly from
the mucilaginous stratum. Fusion of these zoogonidia has frequently,
though not always, been observed, and develpment takes place in exactly
the same way as in the normally produced zoogonidia.
The observation of this delayed production of ejia by the palmelloid cells
is interesting in that it indicates that the aplanospores are really reduced
zoogonilia, but that the surrounding nutrient conditions are such as to be
able to support the development of a large number of individuals in a small
spaee, and to render their wider distribution unnecessary.
The zoogonidia produced in this manner are oval and are larger than those
formed normally. They vary considerably in size, being from 5 to 9°5 w in
breadth and from 9'7 to 14:5 w in length according to the number of
preliminary ‘divisions of the cell. Each contains a single bell-shaped
chloroplast lining the posterior end of the cell, and in some cases the
chloroplast is somewhat irregular in thickness forming cushion-shaped
projections into the interior of the cell. There is a bright red pigment-spot
at the anterior end, and a single pyrenoid in the chloroplast. In certain
circumstances the pa'melloid cells gradually become spherical and develop
into vegetative cells.
It is characteristic of this genus that true vegetative division does not take
place. In some cases the contents of a cell divide into two parts, but the
two daughter-cells invariably secrete new cell-walls, and the mother-cell-wall
becomes converted into mucilage. The two danghter-cells remain loosely
attached to one another, and, as a result of further similar divisions, small
clusters of cells are formed (figs. 4-8). Multiplication of this kind, which
appears to be a modified formation of aplanospores, appears to take place
* Smith, G. M., *Zoospore Formation in Characium Steboldii, A. Dr.," Ann. Bot.
vol. xxx. no. cxix. July 1916.
t Bristol, B. M., “On the Life-history and Cytology of Chlorochytrium grande, sp. nov."
Ann. Bot, vol, xxxi. no. cxxi. Jan, 1917.
CK we Oe eS eee E TSAS CST S
T ee Sen C TD» Cu eee eS
MALAY FORM OF CHLOROCOCCUM HUMICOLA. 479
chiefly in cells from 10 to 20 w in diameter ; cells larger than this multiply
either by zoogonidia or by the formation of numerous aplanospores.
VI. Curonococcva HUMICOLA IN ẸNGLISH BOILS.
Of the samples of soil taken from fifty-one localities in this country, all
but two yielded C. humicola in greater or less quantity, showing the wideness
of its distribution and the constancy with which it ocenrs in soil. Of the
two in which it did not occur, one was taken from the Broadbalk plot at the
Rothamsted Experimental Station in the year 1846, and had been kept in a
dry state for nearly eighty years before being put into the culture. A
similar sample of soil collected from the same plot in 1856 yielded the alga
in considerable quantity, hence its absence from the cultures of the 1846 soil
indicates that the limit of resistance against desiccation and of retention of
vitality for C. humicola lies somewhere between seventy and eighty years.
In no essential features did the Chlorococewm humicola found in the Malay
soil differ from that found in the English soils. The spores germinated far
more slowly and the cells are on the whole larger than the English ones, but,
even in a single culture, the variation in size of the cells is so great that
no importance is to be attached to such distinctions. For the rest, the
cytology and the life-history are the same, the dominant stage depending
on the surrounding nutrient medium.
VII. AMENDED DIAGNOSIS.
Rabenhorst distinguished the species C. humicola by the following
diagnosis :—“ Ch.strato effuso, obseure viridi pulverulento ; cellulis globosis,
magnitudine variis, plerumque numerosis in familias consociatis, tegumento
communi hyalino achroo circumvelatis ; eytiodermate tenui, state provecta
suberasso ; cytioplasmate initio dilute vel luteolo-viridi, homogeneo, demum
saturate viridi, granuloso.
* Diam. cell ad 1/126" —0:0007".
“ Propagatio fit gonidiis cytioplasmatis divisione succedanea ex ultima
generationis serie transitoria ortis, e cytiodermatis abaviwm (intellige
tegumentum extremum) rupturis excedentibus et examinantibus."
To this must now be added the following characters :—
“ Zoogonidia aut quiescunt statimque in cellulas vegetativas angescunt
aut copulatione duorum facta zygota efformant.
* Propagatio fit quoque aplanosporis plurimis (zoogonidiis redactis) in
mueilagine ex solutione cytiodermatis cellule matricalis orta inclusis.
Fieri potest ut aplanospore divisiones plures subeant ; postea aplanosporze
aut cilia extrudunt et se velut zoogonidia normalia prebent, aut tardissime
in cellulas vegetativas augescunt.
« Contentus cellularum vegeiativarum majorum sepissime pigmento rubro
obscuratur,”
180 MISS B. MURIEL BRISTOL ON A
VIIT. SUMMARY.
The material described has been obtained from cultures of a sample of
dried soil, which was sent from the Malay States about two years before the
cultures were set up.
The vegetative cells are spherical or subspherical, solitary or collected
together into mucilaginous strata, very variable in size, being from 20-80 u
in diameter, each with a thin cellulose cell-wall and a single parietal
ehloroplast containing from one to several pyrenoids and numerous starch
granules. In adult cells a quantity of yellow oil is stored, in which a bright
red pigment is often dissolved.
The cytoplasm is reticulate. The young cells contain a single minute
nucleus and one pyrenoid, both of which multiply by repeated division so
that the adult cells are coonocytie with many pyrenoids,
Propagation takes place, by successive bipartition of the contents of the
mother-cell, into 8-16 or numerous biciliate zoogonidia which may develop
asexually or may act as facultative gametes. In both cases direct develop-
ment into vegetative cells takes place.
Aplanospore-formation may also take place, preceded by the multipli-
cation by constriction of the nuclei of the mother-cell. The aplanospores
remain imbedded in a mucous stratum, and enter into a palmelloid state
in which further bipartitions may take place. Eventually, the palmelloid
cells either acquire cilia and behave as normal zoogonidia or they develop
direetly into vegetative cells.
True vegetative division does not take place, but the cell-contents may
divide into two daughter-cells which immediately acquire new cell-
walls and are set free as vegetative cells by the dissolution of the mother-
cell-wall.
Chlorococeum. humicola, differing in no essential partieulars from that in
the Malay soil, has been found to occur almost universally in English soils.
The limit of its resistance against desiccation and of its retention of vitality
has been shown, by investigations on long-dried English soils, to lie some-
where between seventy and eighty years,
In conclusion, I wish to express my thanks to Professor G. S. West for his
valuable help throughout this work.
Dotanical Laboratory,
University of Birmingham.
MALAY FORM OF CHLOROCOCCUM HUMICOLA. 481
EXPLANATION OF TIIE PLATES.
PLATE 17,
Figs. 1-10 and 13 x 825, figs. 11, 12, and 14 x 1435.
1. Small vegetative cell with an irregular parietal chloroplast and one pyrenoid.
2, Cell with chloroplast having a single small perforation and six pyrenoids.
3. Cell with small thickenings on the cel'-wall.
4. Two daughter-cells still united by a gelatinous investment produced from the
disintegrated mother-cell-wall.
5. Small cluster of cells showing the formation of two daughter-cells within the
mother-cell-wall.
Figs. 6-9. Multiplication by zoogonidia in cells of varying sizes.
Fig.
Fig.
Fig.
Fig.
Fig.
iz]
10. Empty zoogonidangium with small internal thickenings of the cell-wall.
11. Zoogonidia of different forms. py., pyrenoid ; pig., pigment-spot.
12. a-e, Successive stages in the conjugation of isogametes; d, zygote with thin
cell-wall before the loss of cilia.
13. Young vegetative cells developing from zoogonidia or zygotes.
14. a-e, Unusual conditions in conjugation. «, Formation of zygote with two cilia at
each end; 5, fusion of anisogametes; c, fusion of zoogonidium to posterior end
of young zygote; d, abnormal zoogonidium set free before division is complete;
ch, chloroplast.
PLATE 18.
Figs, 15-24. Sections to show cytological characters, x 1435.
. 15. Small cell stained with Heidenhain’s hematoxylin showing reticulate cytoplasm,
a single nucleus (v.), and one pyrenoid (py-).
.16. Do. showing two nuclei, a pyrenoid dividing by constriction, and a parietal
.
chloroplast.
. 17. Do. showing finely reticulate cytoplasm, with two nuclei and one pyrenoid.
a and b, two drawings of the same cell at different levels.
ig. 18. Larger cell showing more numerous nuclei and pyrenoids, and indications of a
parietal chloroplast.
. 19. Do. with very numerous nuclei and a number of large pyrenoids; the cytoplasm is
more distinctly reticulate than in the smaller cells.
. 90. Pyrenoids of various forms.
.91. Cell just before the formation of aplanospores, showing enlarged nuclei with
variously shaped karyosomes, c-n., nucleus dividing by constriction.
g. 22. Section through mass of young aplanospores. m., mucous investment formed by
the disintegration of the mother-cell-wall ; n., nucleus; py., pyrenoid.
23. Sections of cells in palmelloid condition. a-c, Stained with Heidenhain’s
hemtoxylin; d, stained with Delafield’s hamatoxylin, showing the reticulation
of the cytoplasm ; m., mucilaginous investment; n., nucleus; py., pyrenoid.
. 24. Section of cells as in fig. 5, with pyrenoids showing distinct plates in the starch-
sheath, an enlarged nucleus, and indications of a parietal chloroplast.
Figs. 25-81, Multiplication by means of aplanospores. X 825.
bo
c
. Group of mother-cells filled with young aplanospores which are angular by
compression, :
i
ANS. ee O,
A MALAY FORM OF CHLOROCOCCUM HUMICOLA.
'
Fig. 26. Mass of young aplanospores rounding themselves off from one another, within the
i mucilaginous envelope formed by the disintegration of the mother-cell-wall.
E successive bipartitions of the aplanospores. ——
E Fig. 28. Development of cilia by palmelloid cells after a change of medium.
motile state.
E ch. , chloroplast.
Fig. 97. Part of a mucilaginous stratum (structureless) with groups of cells formed by
Fig. 29. a, Palmelloid cell which has increased in size without further division; 4, :
: vegetative cell produced directly from an aplanospore without an intermodisin :
` Fig. 30. Zoogonidia produced from palmelloid cells, differing i in size with the number of
preliminary divisions of the palmelloid cella. pig., pigment-spot ; py., pyrenoid ;
Fig. 31. a-c, Successive stages in the fusion of these zoogonidia ; ; d, zygote Sp into
vegetative cell,
JOURN. LINN. Soc., Bor. VOL. XLIV. PL. 17.
BRISTOL,
-
Grout, phot.
B. M. B. del.
CHLOROCOCCUM HUMICOLA.
.
BRISTOL.
B. M. B. del.
V EEE CT, T ENS
JOURN. LINN. SOC.,
3 n «te Tu
WA
V
WO mano
A
J
—
fo.
CHLORQCOCCUM HUMICOLA,
ee ee CPC AEEEN vec . E ME
Bor. VoL. XLIV. PL. 18
Grout, phot.
A EUROPEAN PETRIFACTION WITH FOLIAGE. 483
Bennerrires Scorri, sp. nov., a European Petrifaction with Foliage. By.
Marie CARMICHANL Sropss, D.Se., Ph.D., F.L.S., Fellow and Lecturer
in Paleontology, University College, London University.
«PLATES 19 & 20, and 4 text-figures.)
Slot. Tomo + [Read 2nd May, 1918.
Tue particular interest of the new species of Bennettites about to be described
is mainly three-fold :—(1) It is the smallest and youngest trunk yet known ;
(2) it is the first European specimen to include well pelted young foliage ;
(3) it is well preserved, thus elucidating some anatomical details of leaf-
structure in the genus hitherto not Bieapietely known from American
specimens of. other species. It is also suggestive of the conclusion that
Bennettites produced detachable trunks adventitiously arising as buds com-
parable with those found on the living Cycas circinalis.
General Description.
The specimen had leng lain in the British Museum (Natural History) in a
number of unconnected pieces, and had been transferred without history
from the Botanical Department in 1898 to the Geological Department, where
it now is. The specimen, cut into four separate pieces and two slides long
since, had passed through many hands and been given six different and
unassociated catalogue numbers, viz., V. 4502, V. 4767, V. 4782, V. 5445,
V. 5650, and V. 8423. Some of these had been associated by Mr. W. N.
Edwards, of the Geologieal Department, British Museum.
The parts when all ought together fitted so well that there is no doubt
of their forming a single specimen. There is no record of the horizon or the
locality of its source. :
The tiny trunk was oval in horizontal outline, and rather like a somewhat
pointed pear in vertical elevation. It much resembled outwardly, both in
size and appearance, a large Williamsonia fruit, being only about 8:5 em.
in height, and with a largest diameter of 7x5 em. “Text-fig. 1 gives in
outline the appearance of the specimen in natural size. de ago it had |
been cut across about the middle, cut (1) in the figure, and the upper piece
had then been cut once vertically to this and along the’ shorter diameter,
cut (2), and then one of these resulting portions cut again in half at right
angles to this eat and along the larger diameter, cut (3). These cuts gave,
| the large basal piece A, the half of the upper half, B, and the quarters of the
upper half C and D. From these in the early days two thick lonvitudinal
sections had been: made from the faces of cuts (2) and (3).
When I began work on the specimen Dr. Smith Woodward, F.R.S. „kindly
allowed me to have a complete series of sections from the lower part A,
LINN. JOURN.—BOTANY, VOL, XLIV. 2n
iia Ya
MEME
484 DR. MARIE STOPES ON BENNETTITES SCOTTI,
which yielded sections m to r, and also a series from the upper part B, which
yielded sections « to /, see text-fig. 1, thus giving 17 sections in consecutive
series. Pieces C and D are left uncut: in them there is little doubt that
further young leaves are preserved.
Externally, as is roughly indicated in the diagram (text-fig. 1). the speci-
men showed an irregular, sheathing bract-like covering, and had a curved-in
l'ext-fig. 1.
“SO ROO A
V. 5650
PLT S RIS
Diagram of original Bennettites Scotti. Description in the text.
and hollowed base like that of a Williamsonia fruit, though the covering
* scales" were much less definitely individualised. The external appearance,
however, was so far deceptive, and the specimen was not a fruit but a vouno
trunk with its main axis surrounded only by leaf-bases, and with pets
leaves still folded in the bud towards the apex. This baby stem was still tai
too young to bear fructifications.
In cross-section the small oval axis with its ring of vascular tissue is
apparent : arising from it are the numerous and closely packed leaf-bases in
normal spiral sequence, see Pl. 19. figs. 1 & 2. Above the level of section f.
A EUROPEAN PETRIFACTION WITH FOLIAGE. 485
text-fig. 1, the axis and leaf-base tissues are replaced by ramenta, but the
thiekly packed and definitely oriented ramenta simulate leaf-bases in a
manner already noted and figured for his specimens by Wieland (1906, p. 99,
text-fig. 52).
It is in this upper region that the young leaves lie in the present specimen,
section j being the lowest section in which any of them appear. In &, in the
leaf-base area directly below j, can be seen the corresponding rachis of each
leaf rounding off with its horse-shoe shaped strands of vascular tissue.
The young leaves in the specimen occur as separate fronds in their normal
sequence, each with the pinn: folded as in the bud, and each frond lies
above the area of the corresponding leaf-base, which had narrowed and
rounded off to form the rachis in a distance of a few millimetres.
The pinne in some sections are closely packed together, and in the most
complete leaflet there are tightly packed nine pinnæ on either side ; but
none of the fronds are entire, and some of the more broken or distorted leaves
show as many as ten to sixteen leaflets on one side. Figs. 8, 9, and 12, PI. 20,
show characteristic groups of pinnw. In one instance only are the pinne
attached to the rachis (fig. 7, Pl. 20), most of the groups of pinnz lie in place,
but somewhat distorted in the midst of ramenta.
The ramenta are noticeably of two kinds :—large stout ones which are
oriented so as to form a leaf-base shaped mass, and so to lie that with the
low power they exactly simulate leaf-bases surrounding the axis; and in
addition to these are the much smaller ramenta composed of as many, but
individually very much smaller cells, which are packed in drifts between the
others and also between the true leaf-bases.
As is usual in this family, there is no sign of a main, or indeed of any
other root at the base of the trunk : the lowest section of all (7) has, however,
a curious, nearly central ring of tissue which is discussed below (p. 486).
Detailed Description.
The MAIN axis.—The woody cylinder and the general outline of the stem
of this young and apparently uncrushed axis is distinctly oral (see figs. 1 & 2,
Pl. 19), aud though the original statement of Carruthers (1870) that the
oval shape of the axis is a generic character has often been disputed, I think
that the obvious oval of this exceedingly young specimen distinetly lends
support to Carruthers’ surmise.
Though this little axis is about one-fourth the diameter of the ordinary
Bennettites trunk, its arrangements of axis, vascular strands, leaf-bases, exits
of horse-shoe shaped leaf-traces and so on, are all essentially characteristic
of the genus.
The vascular cylinder is, however, very short and proportionately thick.
At its thickest part (sections p and o) the wood has about 40 tracheids in
2R2
480 DR. MARIE STOPES ON BENNETTITES SCOTTII,
the radius, and a zone of phloem approaching in extent that of the xylem.
The radii of vascular tissue are closely ranked, and are separated by wide
medullary-ray cells into the double or triple strands typical of old Bennettites
wood. But by seetion /, the highest in which the woody cylinder is well
marked before it dies out in the stem apex, the wood is reduced to about five
elements on each radius.
The protowylems lie in radial continuity with the secondary wood, and
there are no detached groups of small tracheids such as occur in some of the
Bennettitalean stems (cf. Cycadeoidea Wielandi, Wieland, 1906, or Colym-
betes, Stopes, 1915).
The elements of the secondary wood are squarish in outline, but vary con-
siderably in the way common in Bennettitalean wood, the average size is
about 15x 20 u to 25x20 p. The tracheids in the higher sections are
noticeably smaller than in the lower ones where the wood zone is thicker.
The tracheids seem to have only the barred thickening characteristic of the
family.
In a number of places the cambium is well preserved, as one or two thin-
walled, radially narrow elements between the xylem and phloem. This is
particularly g ood in slide n.
The pith is oval, about 8x 1-1 mm. in diameter, and is composed of a
mass of soft-walled tissue through which run numerous ** gum-canals," as is
characteristic of the family. There seem to be no isolated groups of tracheids
or transfusion tissue in any part of it.
A feature of particular interest is seen in the lowest section of the series
(r, text-fig. 1), viz. a centrally placed mass of tissue quite unusual in Bennet-
titalean piths. Roughly this mass is 4L em. in diameter, circular. and
placed in the centre of the oval of the pith. In the section above this, g. the
differentiated tissue-mass is not present, but when the section is viewed with
the naked eye some distance from a black background, a central circular
area is apparent, about *5 em. in diameter and distinctly different in colour
from the rest of the pith. Under the microscope this area is not delimited
in any way, but in the central region the cells seem larger and clear, and a
little more irregular than the rest of the pith, sufficiently different perhaps
to account for “the microscopically obvious difference in colour. A few of
these cells seem to be stone cells with thiek walls.
The circle of tissue on slide r can be seen in fig. 4, Pl. 19, where at s not
quite the whole circle is shown. — It contains four definite zones : (1) an outer
circle of very large, rather irregular cells with excessively thick walls ;
between the groups of these large stone cells, small, narrow cells lie rather
as do medullary rays in a bast zone ; there is, however. no conclusive proof
that the circle is a vascular cylinder of any sort, and this appearance is
probably fictitious. (2) Within this ring is a zone of two or three cells
thick of narrow, thick-walled, dark cells, remarkably like a cambium.
VES e SOINS IRE ea eee i E CONES T ii au
A EUROPEAN PETRIFACTION WITH FOLIAGE. 487
(3) Within this is a circle a dozen or so cells thick, much more cork-like
than like wood. (4) The central space within this consists of irregular pith-
like tissue among which groups of thick-walled cells are visible, and of
which the central portions are largely disorganised.
The nature of this curious circle of tissue is a point of interest. Tts
central position and large size make it evident that it is not a chance “ pith-
bundle ” such as oceur in some cycads. It might conceivably be the dying
out of the cylinder of a large primary root, and it is most unfortunate that
there was not material for just one section below r to make clear whether
the tissues were carried down into a definite vascular cylinder or not. The
way it dies out above makes it clear that were ita vascular cylinder it had
no connection with that of the stem. Such primary roots are, I believe, not
known in the cyeadean cohorts.
A suggestion which has more to recommend it is that this i« the upper
part of a separation layer which, at some time, had separated this small stem
from another, a parent stem, from which it may have sprouted. The erratic
sprouting of seedling-like buds and their development into separate stems or
lateral branches in the living Cycas circinalis (see Stopes, 1910) seems to
offer what may be a close comparison. Were this view accepted we must
look on the young stem of this new fossil as a sprowtling and not a seedling.
While it is only wise to reserve judgment about this, it may be mentioned
that, from my experience with living cycad tissues, I incline to favour the
latter interpretation. Also Dr. D. H. Scott, F.R.S., who very kindly
examined slide r, added an argument in favour of this view, saying in a
letter to me, “ I entirely agree with your interpretation, 7. e. that the ring of
secondary tissue round the centre is a separation layer and not vascular. I
judge not only from the look of the cells, but from the fact that the tissue
inside the ring is disorganized, just as would be the case if it were a separa-
tion-layer. I imagine the layer was dome-shaped and that this section cuts
through the sides of the dome.”
The leaf-traces come off from the vascular axis as single, solid ares, with
considerable quantities of tissue forming a fan ; this spreads out and breaks
up into a circle of bundles as each passes directly out through the cortex
into the leaf-bases in the usual way.
The cortex consists of a mass of large-celled soft tissue, in which run
numerous * gum-canals " larger in diameter than the tissue-cells amid which
they lie.
As will be apparent from figs. 1 & 2, Pl. 19. the leaf-bases, though rela-
tively small compared with the type species, B. Gibsonianus, are yet large
proportionately to the size of the stem itself. The leaf-bases measure about
1x8 em. and are thus actually very little less than those of the full-sized
trunk of B. Allehini (Stopes, 1915). They are of the typical rhomboidal
shape and spiral arrangement, The ground-tissue and vascular bundles
488 DR. MARIE STOPES ON BENNETTITES SCOTTI,
show no peculiarity, but the epidermis with its thick cuticles and attached
hairs is particularly well preserved. As the details of these structures have
not been described for the already known species they are worth a few
words in the present instance. In B. Gibsonianus the limiting layer of the
leaf-base is described as “something in the nature of cork,” and sections
of the original type specimen in the British Museum show this corky layer.
A thick cork layer has been more minutely deseribed for the leaf-bases of
C. gigantea by Seward (1897). The leaf-bases of the new species now
described are still in the young condition, covered by the original epidermis.
This has a very thick cuticle which forms a noticeable band of colour in the
sections. One corner of one of the leaf-bases (in slide m) has begun to
develop a multi-layered cork, and in other parts of the same leaf-base the
epidermis with its thick cuticle is conspicuous.
The colour of the general petrifaction of the tissues ranges from a greyish
to a vandyke brown, but the cuticle stands out from this as a clear, vivid
golden-brown colour, remarkably like the cuticle of a living Cycas. It is
not unlikely, indeed recent work in another field (see Stopes & Wheeler,
1918) causes me to think it highly probable, that here we are looking not at
the mineral replacement of the cuticle, but at the actual cuticle itself, thougn
the specimen is completely embedded in a mineral matrix as is usual in
petrifactions. The cuticle is best seen in the lower regions of the leaf-bases,
where it is less frequently broken by ramental attachments than it is in the
upper regions where ramenta are so thickly crowded that the epidermis can
searcely be distinguished.
Several structures which look like stomates are to be seen in the leaf-
bases, but these may be tangential cuts through the bases of ramental
attachments.
The cork layer extends only so far as the acute corner on one side of a leaf-
base, and though it is there about 18 cells thick, it abruptly comes to an end
and the rest of the leaf-base, including the other corner, is still covered by
the original epidermis and cuticle, with here and there a cell or two dividing.
In one place the division to form the first cells of the cork cambium is
particularly clear, and shows that the cork eambium of the leaf-base is
epidermal in origin (see text-fig. 2) and thus differs from that of € . gigantea,
which is described from much older leaf-bases as being sub-epidermal ii
origin (Seward, 1897, p. 28).
The ramenta are particularly numerous on the upper parts of the leaf-
bases, where they are so crowded and are packed so closely together that
they form *false-leaf-bases" which appear to the naked eye in sections
exactly like true leaf-bases, as they are of the size and shape and have the
same arrangement as the latter. This was illustrated by Wieland (1900,
pl. 19, text-fig. 52) for his American species. The present specimen shows
this iito beautifully, and also makes clear an interesting point, that in
A EUROPEAN PETRIFACTION WITH FOLIAGE. 489
this species the ramenta are of two kinds; the very large solid-walled ones
which, curving round each other and packed together, form the pseudo-leaf-
bases, and the interstitial ramenta, very much smaller and thinner walled,
packing the spaces between the former and also between the true leaf-bases.
The marked distinction between these can be well seen in fig. 5, PI. 19,
where b represents about half a pseudo-leaf-base area, and b' part of a second,
while ir are the interstitial ramenta. :
Epidermis and cork cambium. See p. 496.
There are occasional doublings and irregularities in the cells, but the
ramenta are generally one cell-row thick, and the small ones have approxi-
mately the same number of cells as the large ones, i. e. from about 12 to 20
or more. The number of eell-rows in the ramentum is, within reason, a
specific character, as was pointed out by Wieland (1906, p. 52), and the
present species in having a single row resembles B. Gibsonianus and Cyca-
deoide« nigra.
Altogether there are three types of ramenta in this plant, the two types
just described and in addition hair-like ones on the undersides of the leaves
themselves, which will be described below.
The rounding off of the leaf-base to form the rachis takes place rapidly ;
in section j, for instance, one of the leaf-bases is very definitely reduced in
area, and is rounding off, while in section his seen a rounded rachis with
its curved horse-shoe bundle and a few pinne attached to it (P1. 20. fig. 7).
Groups of leaflets, each belonging to one frond and still folded as in the
bud, lie in their normal sequence round the stem and are best seen in section
g for example, at a level high enough up to be past the apex of the axis-
tissues, which at this level are represented only by pseudo-leaf-base ramenta.
The /ronds on the present species are clearly not adventitious, but are its
normal leaves, whether or not the young stem itself had ultimately originated
as an adventitious bud. Each frond is evidently still very young, not yet
unfolded, the pinnules being packed in regular arrangement similar to that
described by Wieland (1906, pl. 19, and 1916, pl. 58), though on a much
smaller scale, The natural size of. Wieland’s packed group of pinnules
490 DH. MARIE STOPES ON BENNETTITES sCOTTII,
forming a single frond is an area roughly L:2x 2 cm., while that of the
present species is only *5 x ^4.
The number of leaflets in each group, /. e. in each single frond, is not
precisely determinable because the groups are not quite complete ; the most.
complete in appearance has 10 pinnules on either side, neatly packed and
overlapping, but higher up as many as 15 pinne are to be seen, and this
number is probably incomplete. As Wieland (1906) pointed out, the number
of pinne in the leaves of a given species of living eyeads varies so greatly
that exact computations of the pinnæ in the fossil forms do not yield distinctive
data, The indications are that the fronds of the present species were not
only small, but had relatively few pinnæ. It must not be forgotten, however,
that a species with many pinnæ like Cycas circinalis for example, have on
the early leaves of their sprouting stems less than a fifth of the number of
pinne present on a leaf produced in maturity.
Details of Leaflet Structure.
In a general way these fossil pinnæ are unlike those of Cycas, but like
those of the other genera of cycads and like those fossil species described by
Wieland (1899, 1906, 1916) in having a series of well marked, parallel
bundles, all through the mesophyll of the leaf, each bundle surrounded by a
well developed sheath and accompanying sclerenchyma. The highest number
of bundles observed in one pinna in the present fossil, is 23 ; the pinnæ cut
lower down having as fewas 5or 6. The general plan of the middle portions
of the leaflets can be seen in fig. 12, Pl. 20, and of the corners of the leaflets
in fig. 11.
These leaflets, like those described by Wieland, have a strong upper cuticle
and sub-epidermal tissue, and the upper differs from the lower surface of the
leaflet. Beautiful as are the American specimens, all the anatomical details
are not yet clear for the leaves of this family, and the present species throws
some light on a doubtful feature of the leaf as described by Wieland,
Details of the tissues of these leaves will consequently be given.
The upper epidermis, as can be seen in figs. 10 & 11, Pl. 20, and text-fig. 3,
is regular and well developed, with thickened outer walls and a cuticle. In
the central portions of the leaflet the epidermis and the thickened tissues
beneath it seem less developed than elsewhere, of. PI. 20: fig. 12 and text-
fig. 4 with fig. 11, Pl. 20, and text-fig. 3. So far as I can observe, neither
stomates nor hairs are present on the upper surface. This agrees with the
generalisations of Thomas and Bancroft (1913) on the group.
Beneath the epidermis, particularly developed at the corners of each
leaflet, where it is several cells thick, is a thick-walled hypoderm, see /,
text-fig. 3, and figs. 10 & 11, Pl. 20. A palisade seems to be slightly,
but not very noticeably differentiated from the more irregular mesophyll,
|
——————————Mm wo
UA! DES, Vine r X CN A y- TARRE E o AL i "TOR en ae ee
A EUROPEAN PETRIFACTION WITH FOLIAGE. 491
I found no evidence of any transfusion tissue between the bundles or in the
mesophyll.
The vascular bundles are surrounded by a definite sheath (5.) of two or three
layers of thickened cells. This sheath is more noticeably thickened on the
under side in the bundles near the edges of the leaflets. The more median
bundles, and those in the middle regions of the pinnules have more delicate
sheaths. A support of sclerenchyma two or three cells thick definitely
connects the sheath with the upper hypoderm. As the sheath-walls were
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Details of leaf anatomy. See p. 496.
considerably thickened presumably they were pitted, but I have not been
able to determine any definite transfusion-like differentiation.
The bundle is entirely centripetal, and appears to consist of two or three
larger vessels and a group of protoxylems : though no bundle is perfectly
preserved, several show the small wood group lying in place directly inside
the sheath so that the quantity of wood was evidently small, and soft tissue,
presumably cambium and phloem, occupies most of the rather large bundle
space, though the greater part of this soft tissue has disintegrated. -
The lower epidermis (L. e.) lies but one or two cells distantifrom the lower side
492 DR. MARIE STOPES ON BENNETTITES SCOTTII,
of the bundle-sheath. It is a definite layer, clearly to be seen, though less
thickened than the upper epidermis, and broken by gaps of what may be
stomata as well as being very often obscured by the thick tuft of hairs
attached to it. These hairs invest the lower side of the leaflets in such a
closely packed mass that they form what at first sight appears to be a
definite tissue (see z, text-fig. 3, and figs. 9 & 11, Pl. 20). The hair-like
nature of the mass and some of their attachments to the lower epidermis can
be seen in text-fig. 2, and fig. 11, Pl. 20. Here and there the basal attach-
ments to the epidermis cells can be seen, and some hairs are cut obliquely,
but most run in a direction parallel to the leaf, so that in the transverse
section of the leaf they are also cut at right angles to their length and
present themselves as more or less circular cells, The area between one
Text-fig. 4.
4. e.
Detailed anatomy of central portion of leaf. See p. 496.
leaflet and its neighbours is packed with these cells, which consequently
together form much the appearance of a true tissue. This false-tissue of
hairs is often about equal in area and shape to the leaflet to which it belongs,
but in the middle region of the frond where the pinnules are more closely
packed (PI. 20. fig. 12, and text-fig. 4) it is eliminated, or reduced to a very
narrow zone.
Cells which seem to me to be a corresponding mass of hair-cells seem to
be interpreted by Wieland (1906, p. 83) as a “lower hypoderm,” a " heavy
sclerenehyma region oceupying all the space below the bundles," and thai
which he takes for a wavy transfusion tissue of a single row of cells seems
to me to be really the lower epidermis. All the layer below this, as
shown in the lower part of the upper half of his text-fig. p. 83, is really not
sclerenchyma within the leaf, but is massed hairs outside it. The thickened,
|
A EUROPEAN PETRIFACTION WITH FOLIAGE. 492
sclerenchyma-like appearance of the hairs which apparently assisted in mis-
leading Wieland as to their nature in his specimens, is, of course, very
noticeable, and does give the mass the look of a selerenchymatous tissue in
places : however, the thickening of the ramental walls seems a well-marked
feature of this plant, as a glance at the photograph fig. 5, Pl. 19, will
demonstrate.
Wieland’s figure (his p. 83), with which comparison should be made, is
said to be (p. 92) “extensively retouched” by himself, and the group of
selerenehyma-like cells he has drawn in what he considers the lower scleren-
chyma region, were actually, I think, hairs probably similar to those of the
present species.
A specific name must be given to this baby Bennettites, though it is not by
any means impossible that some old trunk already known may be the same
species. As the details of the foliage are not known for any other European
form, they are really the vital points in the diagnosis. Needless to say it is
very difficult to draw up a diagnosis from a single specimen, but it is a task
often laid upon a paleontologist who deals in rarities.
I name the species in honour of Dr. Scott, F.R.S., to whom all palieonto-
logists are indebted.
BENNETTITES SCOTTII, sp. nov.
The whole specimen is 8°5 em. in height, but the stem itself is only 3'5 em.
The greatest diameter is about 7x5 em. including leaf-bases. The leaf-
bases are rhomboidal, about 1x'8 em. Axis distinctly oval. Woody
cylinder oval, at its thickest part, with a pith about 1°8 x 1:1 cm., there are
40 tracheids on the radius of secondary wood. Tracheids up to 25 x 30 p.
Encireling the upper part, the leaf-bases round off to form rachises bearing
fronds still folded in the bud. Above the apex of the stem the leaf-bases
are replaced by pseudo-leaf-bases of large-celled, thick-walled ramenta.
Leaflets up to 15 and perhaps more on either side of each frond. Single
leaflets with upper and lower surfaces well differentiated ; cuticle and
thickened hypoderm on upper side; thinner epidermis with a thick weft of
attached hairs on the lower side of many pinnee. Vascular bundles parallel,
close together, from 5 to 23 in each pinna. Middle of leaflet more delicate
and less hairy than the rest ; the lower corners having several layers of
thickened hypoderm. Vascular bundles collateral with well marked bundle-
sheaths, and supporting strands of sclerenchyma. Xylem apparently only
centripetal. No transfusion tissue recognised, [The specimen too young to
bear fructifications. |
494 DR. MARIE STOPES ON BENNETTITES SCOTTII,
7
Summa ny.
A specimen eut into pieces and given a variety of numbers in the British
Museum (Natural History) when pieced together and cut into serial sections
proved to be a very small Bennettites showing well preserved structure.
Though externally the specimen was much the shape and size of a William-
sonia “ fruit,” it is a young stem with foliage still folded in the bud.
Though the stem was independent, there is evidence suggestive of the idea
that this baby had originated as a “sproutling” and not a seedling. In
connection with this a curious central cirele of tissue in the pith of the
lowest section seems best interpreted as a separation layer.
The vascular axis is very short and thick for its size, and is oval, though
apparently unerushed.
The leaves, with the pinnze folded in place, are fairly well petrified, and
are the first specimens of a European Bennettites preserved with internal
petrifaction of its attached leaves. The leaf-anatomy shows a well marked
differentiation between upper and lower leaf-surface, a series of parallel
vascular bundles with centripetal xylem, with specialised sheaths and supports
of upper and lower sclerenchyma.
A noticeable feature of many of the leaflets is the great mass of hairs,
with thick walls, forming a weft of tissue-like substance, in places as thick
as the leaf itself. It is now suggested that the area described by Wieland
as lower sclerenchyma in his American foliage was probably formed by à
corresponding hair- weft, and that his single row of * transfusion cells” was
actually the lower epidermis.
There are three types of ramenta in this new species: the thick-walled,
large-celled, pseudo-leaf-base forming ramenta ; the small-celled, interstitial
ramenta ; and the hairs, composed of chains of single cells, on the lower
leaf-surface.
The leaf-bases are still covered by the original epidermis with its thick
cuticle, save where here and there a few cells are giving rise to an epidermal
cork eambium.
The plant appears to be far too young to bear fruetifieations. The speci-
men is much the smallest and youngest member of the group of Bennettitales
so far known.
It is named B. Scotti in honour of Dr. D. H. Scott, F.R.S.
My thanks are due to Dr. Smith Woodward, F.R.S., Keeper of the
Geological Department, British Museum (Nat. Hist.) for his kindness in
acceding to my request to have sections cut of the specimen; and to
Dr. D. H. Scott, F.R.S., and Prof. Oliver, F.R.S., for the benefit of dis-
cussing the specimen. Also to the Government Grant Committee of the
Royal Society for defraying the cost of necessary photographs,
A EUROPEAN PETRIFACTION WITH FOLIAGE. 495
Authorities quoted in the Teat.
CARRUTHERS, W.—1870. * On Fossil Cycadean Stems from the Secondary
Rocks of Britain." Trans. Linn. Soc. vol. xxvi. pp. 675-708,
pls. 54-63.
Scorr, D. H.—1909. Studies in Fossil Botany. Vol. ii.: Spermophyta.
Ed. 2, pp. xiii, 355-676, text-figs. London.
SEWARD, A. C.—1897. “On Cycadeoidea gigantea, a new Cycadean Stem
from the Purbeck Beds of Portland." Quart. Journ. Geol. Soc.
vol. liii. pp. 22-36, pls. 1-5, text-figs. 1-3.
Sroves, M. ©.—1910. * Adventitious Budding and Branching in Cycas.”
New Phytologist, vol. ix. pp. 235-241, text-figs. 8-14.
—— 1915. Catalogue of the Mesozoie Plants in the British Museum
(Natural History): The Cretaceous Flora, part 2, pp. xxxvi, 360,
pls. 1-32, 112 text-figs. London. i
—— 1917. “ Roots in Bennettites.” Ann. Bot. vol. xxxi. pp. 257-259,
pl. 14.
Srores, M. C., and R. V. WHEELER.—1918. * The Constitution of Coal.”
Tuomas, H. H., and N. Bancrorr.—1913. “On the Cuticles of some recent
and fossil Cycadean Fronds.” Trans. Linn. Soc. Lond. ser. 2, Bot.,
vol. viii. part 5, pp. 155-204, pls. 17-20, 32 text-figs.
WiEkLAND, G. H.—1899. “A Study of some American Fossil Cycads.—
Part II. The Leaf Structure of Cycadeotdea.” Amer. Journ. Sci.
ser. 4, vol. vii. pp. 305-308, pl. 7. New Haven.
—— 1906. “American Fossil Cycads.” Publ. Carnegie Inst. no. 34
pp. 296, pls. 50. Washington.
—— 1916. “American Fossil Cycads, vol. ii.: Taxonomy." Publ. Car-
negie Inst. no. 34, ii. pp. vii, 277, pls. 58. Washington.
1
Explanation of the Abbreviations used in the Texl-figures.
In text-figure | letters a to r refer only to series of slides.
In text-figures 2 to 4:—
b. Baud below vascular bundle. — pv. Protoxylem.
c. Cuticle of epidermis. we. X me, 1), Upper epidermis of leaf
d. Dividing cells of cork cambium. and adjacent leaf,
e. Epidermal cells. MERdlo sheath:
g. Ground-tissue. a. Xylem.
h. Hypodermal cells. z. Zone of hairs attached to lower
Le. Lower epidermis. “surface of leaf; *
ph. Phloem.
196
Fig. 1.
Fig. 2.
Fig. 3.
A EUROPEAN PETRIFACTION WITH FOLIAGE.
EXPLANATION OF THE PLATES.
(All photographs entirely untouched.]
PLATE 19.
Photograph of transverse section, natural size. Out from level m (see text-fig. 1),
showing the small axis of the trunk surrounded hy leaf-bases.
Photograph of transverse section about once-and-a-half times natural size, Cut
from level p (see text-fig. 1), showing clearly the oval pith surrounded by the
oval vascular cylinder of the axis, broken by the leaf-traces. Attached and
associated leaf-bases surround the cortex,
Photograph of the old longitudinal section of the upper part of the axis and leaf-
bases, natural size.
Vig. 4. Photograph of part of the pith from the lowest section of the axis, level r, text-
Fig. 5
Fi. 6
Ina 7.
Fig. 8
Fig. 9
Fig. 10
Fig. 11
Fig. 12
fig. 1, showing the normal tissues of the pith and also the greater part of the
circle of unusual tissue, supposed * separation layer" s. The central cells of this
circle of tissue are much decomposed.
. Photograph of about half of the mass of large thick-walled ramenta forming a
pseudo-leaf-base area, b, and part of a second such, b', these are separated bv
drifts of small-celled ramenta, zr.
- Small part of a leaf-base much enlarged, showing epidermis with thick cuticle,
here appearing as a dark line c, broken by the insertion of a ramentum. Compare
with text-fig. 2.
PLATE 20,
Photograph of rachis, showing its horse-shoe shaped vascular bundle, On the
right a few pinnie are attached to the rachis.
. Photograph of a group of pinne surrounded by ramenta.
. Photograph of part of a similar group of pinne further enlarged. Between each
leaflet (2) a zone about the thickness of each leaflet can be seen (z). This should
be compared with fig. 11 and text-fig. 3.
. Photograph of corner of one of the pinne, showing the heavily thickened hypoderm
at the corner, the bundle-sheaths, etc.
. Photograph of the corners of two pinnz (J) showing clearly the tissue-like weft of
hairs (2) growing from the lower surface of each leaflet and packing spaces
between them about the size of the leaflets themselves, Compare this with
text-fig. 3 and ulso with fig. 9, .
. Photograph of part of a closely folded leaf, taken through the middle parts of the
pinnæ where they have less thickened tissues, and where the pinne lie closely
packed without the wefts of hairs between each. Note how near together the
vascular bundles lie. Note also the contrast between the character of the leaflets
here and in fig. 11. Compare this with text-fig. 4.
(For photographs 1, 2, & 3, Plate 19, I am indebted to the skill of Mr. I’. Pittock
of University College: the others I took myself. ]
EP & MQ.S.phot.
BENNETTITES
SCOTTII, sp.nov.
Hath coll.
TIU
JounN.LiNuNw.Soc. Bor. Vor.XLIV, Pr. 20.
Huth coll.
BENNETTITES @COTTII, sp.nov.
Tee
RELATION BETWEEN THE ALGA AND FUNGUS OF A LICHEN.
The Relation between the Alga and Fungus of a Lichen, By ROBERT
Pautsoy, F.L.S., and Somervitte Hasrines, M.S., F.R.C.5.
(PLATES 21 & 22.)
{ Read 6th June, 1918.]
Recenr writers on the subject of the relation that exists between the alga
and fungus of a lichen have supported the view that hyphe, functioning as
haustoria, penetrate the living algal cells, and gradually absorb their
contents until only empty colourless cell-walls, or walls enclosing a network
of hyphal threads, remain. Upon this conclusion is based, to a great extent,
the theory of parasitism on the part of the fungus, the algal cell being
regarded as a helot that gains little, or nothing, by reason of the close
association of the two organisms.
The literature of the subject shows a complete lack of agreement as to
what takes place after the algal cell has been penetrated, or as to whether
the cell is pierced at all in the living state. Hedlund (6) 1892 was of
opinion that the protoplasm retreated, forming a concavity before the
invading hyphal thread. Schneider (8) 1897 says, “ The haustorium, after
entering the cell, develops a much branched network which encloses but
does not penetrate the cell-plasm.” Peirce (7) 1899, affirms “ That certainly
the haustoria do penetrate the protoplasm of the gonidial cells of Ramalina
reticulata.” Elenkin (4) 1905, when developing his theory of endosapro-
phytism, concluded that ** The parasitic action of the fungus haustoria on the
algal cells has been more or less proved in comparatively few cases” ; while
Danilov (3). a translation of whose paper appears in the current June
number of the ‘Journal of Botany,’ holding views similar to those of Peirce,
claims to have shown, by drawings and photo-micrographs, that branches of
excessively fine hyphal filaments, possibly without cell-wall, or with a wall
so attenuated that it cannot be demonstrated, pierce the cell-plasm in every
direction. He gives no definite information as to the frequency of pene-
tration by hyphe in the ease of the material he used, but leaves one to
infer that the condition is to be regarded as quite normal, and therefore
frequent, for he bases his theory of the relation between the two organisms
upon his statement respecting the hyphæ entering algal cells. On the other
hand, Elfving (6) 1913 says, respecting Hvernia furfuracea, * The formation
of haustoria upon the hyphae which grow into the lumen of gonidial cells, as
represented in popular textbooks and described for other species of lichens,
was extraordinarily rare; in my material only on a single occasion have I
seen such a haustorium ; while the extremely fine haustoria described by
Danilov, as specially occurring in the present species, I have never seen.”
"The present investigation was undertaken for the purpose of obtaining
#
498 MESSRS. R. PAULSON AND S. HASTINGS ON THE
more definite information as to the frequency of penetration, if any, in some
of our common lichens. For the purpose we have used as material
Ramalina calicaris, R. farinacea, Usnea florida, Platysma glaucum, Evernia
prunastri, Parmelia sawatilis, P. sulcata, P. caperata, P. acetabulum, P. fuli-
ginosa, P. physodes, Xanthoria parietina, Physcia pulverulenta, Cladonia
pyvidata, C. coccifera, C. digitata, C. macilenta and C. sylvatica. Our speci-
mens were collected mostly in the Home Counties, either during the months
of February and March or through the autumn months of September and
October. Lichens are probably at their most active period of growth, in the
south-eastern part of this country, during the early months of the year, for
it has been found by us, that the process of multiplication of the algal cells
is exceedingly active during that season of the year; so much is this the
case, that, in growing portions of the thallus the number of gonidia under-
going change is so great that full-sized cells, in a vegetative state, are
comparatively few (PI. 21. phot. 1).
Sections eut from fresh material were examined in water io which
» per cent. of pure glycerine had been added, together with a small quantity
of 75 per cent. alcohol. Other material was fixed either with absolute
alcohol or in a chromic acid solution, prepared according to Schaffner’s
formula, viz., chromic acid 3 gr., glacial acetic acid 3 c.c., water 100 c.c.
This latter gave excellent results with little or no shrinkage. Small portions
were left in the solution for 20 to 24 hours and were then thoroughly
washed in distilled water for at least 12 hours. Fixed material was
embedded in paraffin and cut with a microtome in ribbons from 2 to Tu in
thickness.
Methyl green was found to be a most useful stain for fresh material as
the fungus becomes blue and stands out in contrast with the alga which
remains green. For fixed material we selected Donney's (1) staining
medium, Haidenhain's iron alum-heematoxylin and eyanin and erythrocin, as
differential stains.
Bonney’s triple stain was tried as an experiment, for it was previously
known to us as a stain for animal tissues only. Our object was to find
a reliable reagent that would clearly differentiate the alga and fungus, and
also show distinctly the various parts of the algal cell and the hyphal thread.
With this medium the algal cells become orange-coloured, the central so-called
pyrenoid (nucleus ?) reddish violet, and a small lateral body, surrounded bya
light area, is stained a dark purple. The hyphal protoplasm also becomes
purple. Many of the younger gonidial cells are not stained in the same
manner as the mature cells; the latter, as we have already stated, become
orange-coloured, while the former stain dark purple. We suggest that the
difference in colour is due to the fact that the large cells are cut through
when sections are made, while many of the smallest cells remain entire, and
the colour which comes out so readily from cut cells, when washing the
RELATION BETWEEN THE ALGA AND FUNGUS OF A LICHEN. 499
stained sections with orange acetone, is not so easily withdrawn from those
cells that are entire. This difference in coloration is also noticeable when
using an aqueous solution of methyl green on fresh material for, within
a few seconds, the daughter cells become bright blue, as do the hyphe, while
the older cells remain green, unless kept in the stain for several days. In
this case there is no cutting of cells, they are simply teased out of the
hyphal network ; the older cell-wall resists the entrance of the stain much
more than does the thin newly-formed wall of the young cells.
In introducing his new triple stain, 1908 (1), Bonney says : ‘ The process
is as Follows :—
Solutions.
ie Se Ge ee EE ‘25 gramme.
Pyronin Ey cee ie sn 1:0 a
Distilled water... «Sates. =. agen: gene Ce.
V.B.—Make the solution by heat, and filter. Label * Methyl-Violet-
Pyronin Solution.’
“2. To 100 c.c. of Acetone slowly add, drop by drop, from a drop bottle,
a 2 per cent. aqueous solution of Orange G. (made by heat and
filtered).
“ Whilst adding the solution of the stain, keep stirring and rubbing the
mixture with a glass rod. When the fluid has attained a pale yellow colour,
a faint cloudiness appears. Further addition of the stain increases this
until a floceulent precipitate is formed. Continue the addition of the
Orange G. drop by drop, and this precipitate presently redissolves. Imme-
diately this has taken place, filter into a bottle and label * Orange Acetone.’
Method.
“1, Fix material in Acetic Alcohol (Glacial Acetic Acid F.P. 14: 77-15? C.
1 part, Absolute Alcohol 2 parts). Alcohol, or Sublimate Solution,
Chromates and Formalin render the method useless unless subsequent
to staining the sections are treated by oxidising agents.
Stain for 2 minutes in the Methyl-Violet-Pyronin solution.
Wash rapidly in water and wipe all the slide dry except the section.
Flood the slide with Orange-Acetone. A colour-cloud comes out.
Pour off and flood again—when no more colour comes out.
5. Wash rapidly in pure Acetone, and
_ 6. Transfer to Xylol and mount.”
The whole process should not take more than five minutes, and probably
bo
ee
=
less.
` In using this triple stain we found the best results were obiained by a
slightly slower process than that suggested for animal tissues, thus the
LINN. JOURN.—BOTANY, VOL. XLIV. 28
ome
rice” |r
500 MESSRS. R. PAULSON AND S. HASTINGS ON THÉ
section remained in the Methyl-Violet-Pyronin for 5 minutes, and the
whole process took about 8 minutes.
Haidenhain's iron-alum-hematoxylin is a most useful medium especially
for bringing out the structure of the chloroplast. The so-called pyrenoid
and the small lateral body already mentioned are remarkably distinct after
the application of this stain.
Erythrocin and eyanin, which have been recommended for sections of
lichens, differentiate the two components, but the reaction is not quite reliable.
With the best results from this combination the gonidia become blue,
the so-called pyrenoid dark blue, while the hyphæ take a pinkish-red
coloration.
Gonidia were examined, in the first case, in sections cut immediately
after gathering the specimens. They were either in situ, that is, in the
normal position among the hyphal threads, or were slightly separated from
the section while it was temporarily mounted in glycerine and alcohol.
Under these conditions and after frequent washings of the entire specimen
in distilled water, a small number of stray alge were sometimes still present,
but there was no difficulty in deciding which was the true lichen gonidium,
In the species of lichens enumerated above, the shape and size of the
gonidium, and the shape and structure of the chloroplast and so-called
pyrenoid are remarkably uniform. When saying this we do not forget
that there may be considerable physiological differences in the algal cells as
stated by Chodat (2) as the results of his culture experiments. Our investi-
gations respecting the relationship between the alga and the fungus have,
however, been strictly anatomical.
Fully developed gonidia of the lichens in question have a diameter ranging
from 10 to 20 p with an average approximating to 12 p. They are spherical
except when subject to pressure, which is especially present at the time when
large numbers of new cells are being rapidly added to the gonidial layer
(Pl. 21. Phot. 1). The outer surface of the cell-wall is perfectly smooth,
The chloroplast is large and differs somewhat during the development of the
cell. In the daughter cell it is generally smooth in outline, but, as the cell
enlarges, the chloroplast becomes very minutely verrucose. It is seldom
sufficiently irregular to warrant the term stellate unless this term is
employed simply to distinguish such a form from the cup or bell-shaped
chloroplast which is commonly found in members of the Chlorophyce:e.
Such a well marked. stellate form as that figured by Chodat in the * Mono-
graphie d'Algues en Culture Pure,’ under Cystococcus Cladonio, Chod.,
have not seen in the gonidia of Cladonia pyaidata when forming
of the lichen. :
It is quite clear that the microscopic characters of the chloroplast have not
been stated with any great degree of definiteness by the older writers, and
for this reason it has been not without difficulty that a decision could be
we
a component
RELATION BETWEEN THE ALGA AND FUNGUS OF A LICHEN. 501
arrived at as to the exact shape of the chloroplast of the alga frequently
associated with many foliose and fruticose lichens, and referred to by them
as Cystococcus humicola, Nüg.
Elfving (5) in describing Jrernia furfuracea, Ach., says, “the gonidium
contains a hollow spherical chloroplast cut into on one side," but the illus-
trations in his paper “ Untersuchungen über die Flechtengonidien," 1913,
do not make this ** cut into one side " at all conspicuous.
There is no vegetative division of the gonidia that we have examined, but
increase in their number results from the production of daughter gonidia
(reduced zoogonidia) formed by the division of the protoplast into 4, 8, 16, or
32 masses. Frequent numbers are 8 and 16. Division seldom stops at 4
and rarely exceeds 16 (Pl. 21. Phot. 1, d.y.). It has not been possible, so
far, to follow the division of an individual protoplast, but in sections of
material where the formation of daughter gonidia was actively taking place at
the time of fixing, various stages, from the commencement of changes in the
mother cell to the formation of the daughter gonidia, are represented. The
newly-formed cells rapidly become spherical, secrete a cell-wall, develop a
conspicuous so-called pyrenoid, and exactly resemble in miniature a mother
cell. The wall of the mother cell is absorbed, for we can trace no rupture
of the cell-wall to allow of the escape of daughter gonidia. The empty
gonidial cells, which always occur in varying quantities in a lichen thallus,
cannot be regarded as those from which daughter gonidia have escaped, for
these latter remain in the original groups after the mother cell-wall has
entirely disappeared, and it is the growth of hyphe between and around the
newly-formed gonidia that forces them apart or binds them together into
irregular masses. The empty and partially empty cells are the result of the
disorganization of the protoplast, but such change is not by any means due
only to the effect produced by the penetrating hyphal filaments. The whole
of the contents of a gonidium may be absorbed, normally, without any
apparent penetration of the cell-wall. Under the conditions in which
gonidia exist death frequently occurs, it would appear, owing to the great
number of gonidia produced at the time of sporulation, and to the con-
sequent erowding together and lack of air. Daughter cells are often
abortive before the mother cell-wall has disappeared.
We have not found a nekral layer such as Hlenkin (4) figures in
sections of Acarospora glaucocarpa and Lecidea atrobrunea, but we have
seen a relatively small percentage of dead cells after using Chlor-Zine-
Iodine, which makes them stand out very distinctly from the living gonidia
and hyphz. Danilov states that the substance in partially empty cells is
fungoid, that is, it represents the fine hyphal filaments that penetrated the
living protoplast and absorbed its substance. It is true that the sub-
stance within a partially empty cell sometimes stains blue immediately on
the application of an aqueous solution of methyl green, as does the proto-
502 MESSRS. R, PAULSON AND S. HASTINGS ON THE
plasm of the hyphal thread, but this is owing to a difference in permeability
of the dead and living cell-walls, for the contents of a living cell are stained
similarly if left in the stain for a few days
À large central so-called pyrenoid with a diameter of about one-third that
of the cell, viz., 3 to 5, is almost invariably present (Pl. 21. Phot. 1, ».).
It is there, in the newly-formed gonidia, at a very early stage, when the
diameter of the gonidium does not exceed 2? u. It is readily seen in. gonidia
from fresh material, and stains very quiekly in that which has been fixed, so
much so, that it becomes a prominent feature in all stained preparations
except in cells about to sporulate, and in the separated masses of the
original protoplast immediately after division. We regard it as the nucleus
of the cell.
With Bonney’s or Haidenhain's stain the so-called pyrenoid (PI. 21.
Phot. 1, n.) shows a definite structure, not merely on the circumference,
where. it might be interpreted as being due to amyloid substance, but a
structure that runs throughout the whole body and is seen in all sections in
whatever direction the cut may be made. It certainly has not a distinct
erystalline form such as that which is often figured.
In some sections there are cells which we have named twin gonidia
(Pl. 21. Phot. 2). It appears that two of the eight or more daughter cells,
while still within the mother cell-wall, instead of secreting each its own cell-
wall become surrounded by a common boundary. It does not appear probable,
after the close examination of many preparations, that division of the proto-
plast into two portions only takes place. These twin gonidia are mostly
equal or nearly equal in size, but occasionally one is normal while the other
is crushed against the common cell-wall during the carly stages of develop-
ment and so remains quite small.
We regard the formation of daughter cells as a perfectly natural process
taking place abundantly at a definite period of the year. Danilov suggests
that division of the protoplast takes place after the gonidium has been pene-
trated, and that the change is probably an effort to ensure that some of the
protoplast of the invaded cell should escape contamination by the fungus
plasm. The great activity in the formation of spores at a certain datids
season of dis; year leads us to the conclusion that the formation of spores
corresponds to the similar process that takes place when isolated gonidia are
cultivated upon agar-glucose and are not subjected to any excitement from
fungoid hyphe.
The diameter of the hyphal threads of lichens varies with the species,
much more than does that of the diameter of the gonidia. Taking the
hypha of the medulla, that part where the hypha is the least modified in the
lichen thallus, we find that the diameter varies from 3 to 4:5. It may be
2p in one species, 45 in another, while the mature gonidia have the same
average size in both. The symbiont fungus differs, while the alga appears
RELATION BETWEEN THE ALGA AND FUNGUS OF A LICHEN. 503
to remain constant in a large number of lichens. On the upper surface, the
hyphæ anastomose and become so closely welded together that the separate
threads cannot be recognised, and a false tissue results (Pl. 21. Phot. 3).
The hyphz of the gonidial layer have thin walls, they branch frequently,
and are shortly jointed. These short branches, in contact with algal cells,
swell out considerably into pyriform ends, and present a large surface of
contact with the cells that they surround (Pl. 21. Phot. 3). Sometimes a
gonidium is attached to the end of one of the short hyphal branches and
looks as if it had been cut off from that branch. The reagent Cl.Zn.I.
differentiates the cell-wall of the spherical cell from that of the branch.
The hyphe of the medulla form a loose felt with abundant air spaces
(Pl. 21. Phot. 1).
We have noted two forms only of contaet between hypha and gonidium :
(1) when the gonidium is surrounded with loosely-applied hyphæ, “ the
simple contact " of Schneider, (2) the intimate contact where, by a modifi-
cation in shape, the hypha presents a large amount of surface to the gonidium
to which it attaches itself. Such branches are known as extra-cellular
haustoria. We have not found a ease that could be regarded as definite
hyphal penetration of a living cell by intra-cellular haustoria. From this it
follows that the occurrence of fine gelatinous filaments, protruding from a
penetrating hypha and forming a network surrounding and even penetrating
the chloroplast, have not been seen, consequently we are unable to accept
any theory based solely upon their existence. Cases sometimes occur which
resemble a network of threads around the chloroplast. The appearance is
due occasionally to plasmolysis, and can be produced by placing sections of
fresh material in pure glycerine. The beautiful reticulation of the cyto-
plasm (PI. 21. Phot. 2} also produces the appearance of a network of threads
under certain conditions of the microscopical illumination of the object.
It has already been mentioned that the mature chloroplast is a spherical
body covered with minute verrucose protuberances. A minute object with
such a structure is likely to give rise to the formation of a defective image if
the illumination from the substage condenser of the microscope is not perfectly
centred, and when there is a misprision of focus. A defective image may
also be produced by using a smaller cone of light than that required by the
numerical aperture of the microscope objective in use. The defective image
in each case gives more or less the appearance of threads passing over the
chloroplast.
The condition of the protoplast of the gonidium, just previous to the
formation of spores, presents features that are difficult to explain, but no
penetrating hyphz have been found in such cells.
With the microscope alone it is extremely difficult to decide even the
family of the Chlorophyces to which the green spherical gonidium, common
to a large number of lichens belongs, and as we have not attempted to carry
:
504 MESSRS. R. PAULSON AND 8, HASTINGS ON THE
out the culture method, we present the result of observations based alone
upon the microscopic image.
West (9) says: * A few of the Chlorophyceœ have become constituents of
the thalli of many lichens .....— It is mostly one or two of the unicellular
and colonial members of the Protococeacer whieh are thus found.” In the
same work he states “ Chlorococeum, Fries, 1825 [= Cystococeus, Nüg.,
1849],” so that the name so long associated with the gonidia of lichens is
now Chlorococcum humicola.
As a result of the present investigation, we have arrived at the conclusion
that the gonidium of the lichens mentioned in this paper is most probably
a species of Chlorella, and that the daughter gonidia are reduced zoogonidia.
West (9) suggests that this is not unlikely the explanation of the similar
form of reproduction in the case of free Chlorella. We do not attempt to
name the species that forms the algal symbiont, but it is perhaps worth
noting that the mature gonidium of the lichens already referred to has a
greater diameter than that of a free cell of Chlorella vulgaris, Beijer., and
that during the formation of daughter gonidia, the number of such produced
within the mother cell of the lichen gonidium is larger than the number
formed in the free cell.
There is reason for concluding that Protococeus viridis, Ag.. forms the
gonidium of very few of the common British fruticose and foliose lichens,
for we have not met with any vegetative cell-division whatever among the
gonidia of the material that we used.
While engaged on that part of our work that required magnification
of from 800 to 1000 diameters, we have used a Zeiss 2 mm. achromatic oil-
immersion objective with an achromatic oil-condenser.
The photographs were voluntarily prepared for us by Mr. J. H. Pledge,
F.R.M.S., who had at his disposal, by permission of the management, the
whole of the apparatus of the physical laboratory of the Kodak Company’s
factory at Harrow. He employed a Leitz 2mm. apochromatie oil-immersion
objective, a Beek oil-immersion condenser, Watson * Holos” oculars, x5,
x 10, and sometimes a Zeiss x2 projection eyepiece. Colour screens were
used to accentuate or suppress certain details of the preparation. We gladly
avail ourselves of this opportunity for expressing our indebtedness to him
for the skilful care exercised in the production of the photo-mierographs.
We take this opportunity of thanking Professor G. S. West, who most
generously gave us advice and kindly examined some of the preparations
from whieh the photo-mierographs had been taken.
Summary.
A summary of the investigation may be made by reference to Cladonia
digitata, var. denticulata Ach. var. monstrosa Nyl., the lichen which has been
used as material for many of our preparations,
RELATION BETWEEN THE ALGA AND FUNGUS OF A LICHEN. 505
1. The gonidium is spherical except when subject to pressure from other
gonidia. The diameter of fully developed cells ranges from 8 to 15 y.
2. The chloroplast, in the mature gonidium, has a slightly uneven surface.
3. After fixing and staining, minute reticulation of the cytoplasm is
evident.
4. The so-called pyrenoid (nucleus) is large and central and exhibits a
distinct structure throughout the substance. Its diameter is roughly one-
third that of the chloroplast.
5. A small lateral body surrounded by a light area stains darker than the
so-called pyrenoid ; it is very conspicuous in many of the preparations ; it
probably represents the centrosome.
6. Twin gonidia frequently occur.
7. There is no vegetative cell division of the gonidium.
8. The increase in the number of gonidia results from the formation of
daughter gonidia, autospores (reduced zoogonidia).
9. We have seen no penetration of gonidia by 'hyphæ.
List of Papers and Publications quoted.
1. Bonney, Vioror.—“ A New and Rapid Triple Stain.” Archives Middlesex
Hospital, vol. xiii. 1908.
2. Chopat, R.—Matériaux pour la Flore Cryptogamique Suisse, vol. iv.
tase. 2. 1913:
3. Danitov, A. N.— Ueber das gegenseitige Verhältnis zwischen den
Gonidien und dem Pilzkomponenten in der Flechtensymbiose." Bull.
Jard. Imp. Bot. St. Pétersbourg, tom. x. livr. 2. 1910.
4 ELENKIN, A. A.—‘ Neue Beobachtungen über die Erscheinungen des
Endosaphrophytismus bei heteromeren Flechten.” Bull. Jard. Imp.
Bot. St. Pétersbourg, iv. pp. 25-38. 1904.
5. Errvixca, F.—' Untersuchungen über die Flechtengonidien." Acta Soc.
Sci. Fenn. xliv. no. 2. 1913.
. HzpLoxp, I. I.—' Kritische Bemerkungen über einige Arten der Flechten-
gattungen Lecanora (Ach.), Lecidea (Ach.), und Micarea (Fr.).” Bihang
till K. Svenska Vet.-Akad. Handlingar, Band xviii. Afd. 3, no. 3. 1892.
. Peirce, G. L.—* The Nature of the Association of Alga and Fungus in
Lichens.” Proc. Calif. Acad. Sci. Bot. vol. i. 1899.
. SCHNEIDER, ALBERT.—À Text-book of General Lichenology. Bingham-
ton, N.Y. Willard N. Clute & Company. 1897.
9. West, G. S.—Cambridge Botanical Handbooks. Algæ, vol. i. 1910.
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500 RELATION BETWEEN THE ALGA AND FUNGUS OF A LICHEN.
EXPLANATION OF THE PLATES.
PLATE 2l.
Phot. 1. Cladonia digitata, Transverse section of a squamule of the thallus showing gonidia
in various stages of development.
n. Nucleus, surrounded by the chloroplast. On the circumference of the latter is a
darkly stained body, probably the centrosome. ‘The cell-wall is indicated by
a faint line against which are the cut ends of the hyphe.
dg. Group of daughter gonidia. x 1000.
Phot. 2. Section as in the above. Twin gonidia with common cell-wall. Note the retieula-
tions of the cytoplasm. The cell-wall is indicated as in Phot. 1. x 2500.
Phot. 3. Cladonia pyaidata. Transverse section of a basal squamule showing part of the
gonidial layer with pyriform ends of the hyph:e closely attached to gonidia;
they are the extra-cellular haustoria. x 1000.
PLATE 22.
Phot. Cladonia digitata. Transverse section of a squamule of the thallus showing a
159; portion of the gonidial layer next the medulla. The photographs exhibit four
5&4, planes of the same section of different depths. On the left is a mature gonidium
in a plexus of hyphæ, on the right a group of daughter gonidia being surrounded
by hyphæ. x 1000.
Paulson & Hastings Journ Linw Soc Bor VoL XLV. PL ai
J.H. Pledge Phot 3
l &2 CLADONIA DIGITATA 3 CLADONIA PYXIDATA
Paulson & Hastings Journ. Linn. Soc.Bor.VoL XLIV. Pr 22
e =
J.H.Pledge Phot
CLADONIA. DIGITATA
EXTERNAL MORPHOLOGY OF THE STEMS OF CALAMITES. 507
On the External Morphology of the Stems of Calamites, with a Revision of
the British Species of Calamophloios and Dictyocalamites of Upper
Carboniferous Age. By E. A. NEWELL AnnER, M.A., Sc.D., F.L.S.,
and F. W. LAwrrELD, D.A.*
(PLATES 23-25.)
[Read 7th November, 1918.]
CONTENTS.
Page
TV Tntroduction