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BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)
BOTANY Vol. 4
1965-1973
BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1975
DATES OF PUBLICATION OF THE PARTS
No. 1 November 1965 No: 2 May 1967 No. 3 June 1968 No. 4 May 1969 No. 5 June 1969 No. 6 August 1970 No. 7 q ‘ August I97I No. 8 . “ December 1973
ISSN 0068-2292
Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU
1*
CONTENTS ZUR NL WS
BOTANY VOLUME 4
PAGE Cuticular studies as an aid to plant taxonomy. By C. A. STACE . I The genus Elaphoglossum in the Indian Peninsula and Ceylon. By W. A. SLEDGE : i ; i ; : : es. Fungi of recent Nepal expeditions. By F. L. BALFouR-BROWNE .— 97 A synopsis of Jamaican Myrsinaceae. By W. T. STEARN : ww Lag The Jamaican species of Columnea and Alloplectus (Gesneriaceae). By W. T. STEARN : : , = §E7O New or little known Himalayan species of Swertia and Veratrilla (Gentianaceae). By H.SmitH . : : ; : eek y A survey of the tropical genera Oplonia and Psilanthele (Acantha- ceae). By W.T.STEARN . : : : ; 250 Angiosperms of the islands of the Gulf of Guinea (Fernando Po, Principe, S. Tomé, and Annobon). By A. W. EXELL . ‘ 3325
Index to Volume 4. , ; : : ‘ ; ; mee 8
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CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
BY CLIVE A. STACE
(University of Manchester) Y y }
Pp. 1-78 ; 10 Text-figures ; Plates 1-5
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) BOTANY Vol. 4 No. 1 LONDON: 1965
THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.
This paper is Vol. 4, No. 1 of the Botany series.
© Trustees of the British Museum (Natural History), 1965
TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY)
Issued November, 1965 Price One Pound Fifteen Shillings
COTIGULARSSPUDIES=ASeAN Alb TO - PLANT “TAXONOMY
By CLIVE. A. STACE
THIS review is an account of the systematic value of the epidermal characters of angiosperm leaves as they are seen on preparations of the cuticular membrane, and is based on a detailed study of about 250 species belonging to three families. The introductory sections outline the history of the taxonomic use of cuticular and epider- mal characters, the anatomical background of the features observed on cuticular membranes, the methods employed in the isolation and mounting of the latter, and a suggested routine for their formal description. The main part of the work comprises a somewhat detailed account of the cuticular characters which have been found to be useful in identification and taxonomy, and of the variability that some of these exhibit in various conditions. The precautions necessary in the use of such characters are stressed. In the final section the place of cuticular characters in modern taxonomic work is discussed.
It is hoped that these comments will help to place the systematic use of the epidermis upon a firmer and more scientific basis than has hitherto been the case.
This paper is a modified portion of a thesis submitted to the University of London for the Ph.D. degree. The work was supported by grants from the Department of Scientific and Industrial Research.
I am indebted to the Keeper of Botany, British Museum (Natural History), for providing laboratory, library and herbarium facilities for the pursuit of this work, and to Dr. A. W. Exell for his very considerable help and valuable discussion at all stages. I am also grateful to those members of the Departments of Botany and Palaeontology, British Museum (Natural History), who have helped me with several problems, and to Mr. G. Grange, Department of Botany, University of Manchester, for preparing the photomicrographs.
References (by author and date) given in the text are to the list of Special Literature at the end of the paper.
4 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
§I. PREFACE
Plant taxonomy has undergone a great many changes since it was first treated as a scientific discipline in the seventeenth century, one of the major contributing factors being the discovery of the use of new fields of investigation to supplement the old. The ultimate aims of the majority of taxonomists have naturally also changed, the original hope of cataloguing every species of plant now embracing the idea that their evolutionary and genetic interrelationships should also be expressed. I accept that taxonomists should aim at devising “natural”’ classifications which attempt to indicate the phylogenetic position of each basic unit, although it seems to me certain that this goal will be in sight only when the genetical constitution of each taxon is known and can be expressed with something approaching the precision of a chemical formula, towards which the present binomial system of nomenclature is no more than a preliminary step.
Prior to the latter part of the nineteenth century taxonomists used only morpholo- gical (i.e. macroscopic) characters in their studies, and apart from the inclusion of the broadest vegetative features they confined themselves to the reproductive organs. It was not until about 1865 that anatomical (i.e. microscopic) characters were first used, and considerably later that this became usual practice. The volumes of Engler & Prantl’s Die natiirlichen Pflanzenfamilien (1887-1915) included a relatively large amount of anatomical and vegetative morphological details, and since then taxonomic monographs have been considered incomplete without them. Modern workers are constantly employing new diagnostic characters, and work of this kind will need to continue until every structural, physiological and biochemical feature has been utilized.
Parts of the plant which have been most successfully investigated from an anatomical rather than morphological viewpoint include the wood, primary vascular system, laticifer systems, crystals, epidermis (including stomata and trichomes), floral vascular supply, sclerenchyma, pollen-grains and chromosomes. A further group of characters, such as colour, chromosomal behaviour during meiosis, antigen- antibody reactions and chromatographic analyses of various chemical constituents, in fact represent plant structure at a third (i.e. sub-microscopic) level : that of the molecule. Only a synthesis of all this and other evidence will provide anything approaching a complete picture of each species, and thus an ideal classification. Even today there is a great reluctance by many taxonomists to make real use of any but macroscopic characters, and there are a large number who believe that vegetative features are to be used only as subsidiaries to evidence from floral studies.
Of all the types of somatic anatomy studied it seems clear that the less superficial layers of the plant have provided the best evidence for taxonomists, the wood, where present, perhaps being the most useful of all. The leaf epidermis, or its cuticle, has received relatively little attention from neobotanical taxonomists and most of the studies undertaken have been concerned with somewhat specialized groups of plants, e.g. Gramineae. Palaeobotanists, however, have used epidermal characters in classification since the beginning of this century, and today their use is
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 5
practised as a routine procedure, although here the groups of plants concerned have usually been non-angiosperms. In recent years much more attention has been paid by both neobotanists and palaeobotanists to their use in angiosperms, but most of the studies are far from adequate and the subject is still in its infancy.
Except in fresh material of a relatively small proportion of species the leaf epidermis of modern angiosperms is extremely difficult or impossible to remove in toto. In fossil species, moreover, it is frequently not or only fragmentarily preserved, and most of the early work on fossil leaf forms was carried out solely with the aid of leaf impressions. The discovery that the cuticle, which covers the whole of the leaf epidermis, bears an imprint of the epidermal features was therefore of great importance, especially as it often remains on fossil leaves in a well-preserved state long after all cellular layers have disintegrated. Moreover, except in plants with a very thin cuticular covering, the cuticle can be removed from the fossil or modern leaf (whether fresh or as a herbarium specimen) in a continuous sheet, to which end several methods have been employed. Whilst some epidermal features are lost in cuticular preparations the cuticle does provide some extra characters, and the study of the cuticle rather than the epidermis places the work on a consistent and precisely definable basis which is comparable with palaeobotanical investigations. The present survey is concerned with the cuticle itself, although epidermal studies are of course highly relevant.
The level of knowledge now reached with regard to the systematic anatomy of the epidermis or cuticle is quite impressive in some groups, such as Gramineae or gymnosperms. In the dicotyledons and other monocotyledons, however, little extensive work has been attempted. Most investigations have been aimed at separating small groups of species or genera with little or no attempt at studying the degree of variation found within the taxa which are being separated. Much of the literature in the dicotyledons consists of short sections subsidiary to much wider studies of a more general nature. Work on fossil or recent plants by palaeo- botanists has been solely for the purpose of identifying fossils by means of a compari- son with living species rather than to investigate the intrinsic features of either group, and scarcely any mention is found of the possibilities of the presence of environmental or other variation. Many of the better surveys, even excluding the gymnosperms or Gramineae, appear to have been undertaken on plants with leathery, somewhat xeromorphic leaves (e.g. Lauraceae, Magnoliales). Such leaves have a thick cuticle which shows the epidermal cell pattern clearly, and for some reason these xeromorphs usually have a less variable pattern than mesomorphic species. Studies of the latter types have mostly been of a very limited scope, and in many cases a quite inadequate amount of material has been used as representative of each species.
Thus there is a considerable gap in the literature with regard to both the variation of cuticular patterns within a species and the types of characters and their degree of taxonomic usefulness which are to be found in ‘“‘ normal’ mesomorphic di- cotyledons. The series of investigations upon which this article is based was aimed
6 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
directly at helping to fill this gap. The survey was intended to be intensive rather than extensive, for, whilst this narrows the range of experimental material, it was just this close study of a limited group of plants which was needed. Extensive investigations have been undertaken in the past by a few workers, mainly palaeo- botanists, but they have invariably been too superficial for general use. The families of plants chosen for this programme, Combretaceae, Rhizophoraceae and Avicenniaceae, are almost wholly tropical. They contain about 550, 120 and 11 species in twenty, sixteen and one genera respectively, and although all are woody in habit the species show a great variety of growth forms, including trees, shrubs and lianes of all degrees of specialization occurring in all types of tropical habitat. All three families, of which the first two are regarded as very closely related but the third as very remotely so, contain mangroves. It is suggested that the results gained from the intensive study of these families have enabled a good number of general conclusions to be drawn which are applicable to all or most groups of dicotyledons.
A soundly based theory of cuticular patterns can be put to a good number of uses besides the obvious applications in identification, taxonomic research and phylogenetic investigations. These include peat stratigraphy, pharmacognostical analyses and animal foodstuff research. Only after thorough examination of all aspects of epidermal and cuticular variation can systematic cuticular evidence be legitimately used in these fields, and it is hoped that the following sections will go some way towards realizing that aim.
§2. HISTORICAL INTRODUCTION
A brief historical survey of the study of epidermal and cuticular anatomy throws much light on the situation at the present day, and is a considerable help in under- standing the areas in which work is now most needed.
The earliest reference to fossil cuticles is apparently the note by Brodie (1842) “When the sandstone is freshly broken the epidermis of the fossil frequently peels off ...’’, but this observation was not followed up. Three years later Goeppert & Berendt (1845) figured cuticular fragments of one species of conifer and two species of angiosperm which they found preserved in amber. These figures appear in a large volume primarily concerned, amongst plants, with twigs and leaf impressions, and this is the case with the later cuticular diagrams of Schleiden (1846), Unger (1853) and Wessel & Weber (1856). In all four cases the figures are crude and highly diagrammatic, and except where the material has been available for subse- quent study the true identity of the plants is obscure. For example the leaves described by Unger as a Potamogeton have been found to belong to a member of the Lovanthaceae, whilst those figured by Schleiden are not identifiable at all (Edwards, 1935). Wessel & Weber described and figured a number of dicotyledons which they referred, in most instances, to living species, whilst Goeppert & Berendt more advisedly used generic names, such as Alnites, which were not applicable to living plants.
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY if
The first really important work on the systematic treatment of cuticles was that of Bornemann (1856), describing fossil cuticles of cycads from the neighbourhood of Thiiringen. He was the first person to realize the exact nature of the cuticles which he studied, and went to great pains to explain the relation between them and the leaves which bore them. He wrote: “ The outline of the cell walls of the epidermis is almost always shown on the (cuticular) membrane by a network of dark brown lines, which apparently represent the cell walls, but which are much thinner than these would have been. These brown lines are to be regarded as parts of a homogeneous cuticle .. .”’ (transl.).
From then onwards a large number of works, from short notes to large volumes, began to appear on the subject of fossil cuticles. Even so, systematic studies on the surface features of angiosperms generally made use of the whole epidermis, and such investigations in the latter half of the nineteenth century were very numerous. One of the earliest of these was the investigation by Prillieux (1856) into the types of peltate trichomes found in various members of the Oleaceae, and he was actually able to separate all 23 species examined on the structure of these trichomes alone.
The first taxonomist to make constant use of anatomical characters for his diagnoses was apparently Bureau (1864) in his revision of the Bignoniaceae, although these characters naturally formed but a very minor part of the descriptions. Soon after this anatomical characters began to appear more widely in taxonomic treat- ments, reaching a climax in Engler & Prantl’s Die natiirlichen Pflanzenfamilien (1887-1915). Conversely anatomists were becoming more aware of the systematic value of many of their studies, and a number of extremely useful surveys appeared at that time, several of which are unknown to many present-day workers. Weiss (1865) produced a monumental account of the sizes, numbers and distribution of stomata in a large number of dicotyledons ; Bokorny (1882) surveyed the distri- bution and occurrence of pellucid spots in dicotyledon leaves, classifying them according to their causes ; Bachmann (1886) gave a beautifully illustrated account of all the types of peltate hairs known, in systematic arrangement ; Grob (1896) provided a very useful early account of grass-leaf epidermides ; and von Minden (1899) thoroughly covered the different types of water-secreting organs (e.g. water stomata, hydathodes) found in dicotyledons. It was perhaps this productive period which led Fritsch (1903) to summarize the most important systematic anatomical characters to which taxonomists might pay more attention, and in many instances his comments still apply today. Solereder’s encyclopaedic volumes on dicotyledon systematic anatomy, first published in 1898-99, represent the epitome of anatomical works at that time, and together with a supplement were translated into English by Boodle & Fritsch (1908). They remain a standard reference today, one of their most valuable aspects being the detailed survey of the principal families in which a wide range of anatomical characters are to be found.
Although works on fossil cuticles from Bornemann’s (1856) time onwards were equally numerous very few of great value were produced before the present century,
8 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
and scarcely any of these were concerned with angiosperms. Schenk (1869-71) included some drawings of cuticles in his studies of Wealden plants, but of course none of these was an angiosperm. Zeiller (1882) wrote what must be regarded as the first review on cuticular studies, and his article brings together much useful information with many supplementary original facts on cuticles of fossil ferns and conifers. Perhaps the most important of the early cuticular studies, however, were those of Nathorst (1907-12). His Paldobotanische Mitteilungen represents the first work which reported a long series of cuticular studies on a large number of species, and should be considered as the consolidation of this subject as a truly scientific discipline.
One of the most important features of Nathorst’s work is the improvement in various techniques, and these were quickly assimilated by palaeobotanists elsewhere in Europe and in America. This resulted in a considerable number of studies entirely devoted to cuticles, rather than studies using cuticular characters as sup- plementary evidence as had previously been usual. Thompson (1912), for instance, studying Cretaceous conifers, re-examined the genus Frenelopsis and concluded that Zeiller (1882) was wrong in believing that its stomata had the unusual number of four guard-cells : in fact these four cells are the subsidiary cells, the guard-cells being quite normal.
The first noteworthy cuticular investigations by British workers were those of Thomas & Bancroft (1913) on cycads. They compared cuticles of fossil and recent species with an emphasis on stomatal characters, and thus uncovered a considerable amount of detail regarding the relationships of the fossil types. Some of the first good photomicrographs of cuticles were included in this publication, and others were produced by Wills (1914) who reported on fossils from British coal-measures. Thomas (1930) later produced an article supplementing the knowledge of Mesozoic cycadean fronds set out in the earlier work. To exemplify the spread of cuticular studies from continental Europe Holden’s (1915) paper may be cited, being one of the first works on Asian cuticles. From her study of Indian fossil conifers she concluded that epidermal leaf characters may often be of great value in specific determination, but that they seem of little use as phylogenetic evidence.
By far the most important cuticular investigations are those of Florin, who for the past forty years has produced a large number of excellent accounts of recent and fossil gymnosperm cuticles, together with first-class photographs and drawings. One of his earliest important papers (Florin, 1920) was concerned with a comparison of fossil conifer cuticles with those of modern conifers, and most of his subsequent work has been developed along these lines. Florin’s most important work on modern conifers (1931) includes descriptions of the cuticles of almost every known species, with particular reference to stomata, and a section is also included on the stoma types of the major gymnospermous fossil and recent groups for comparative purposes. Later (1938-45) he produced an even larger work on Upper Carboniferous and Lower Permian conifers, including full details of their cuticles, but this work also embraces the reproductive structures and other vegetative organs than the
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 9
leaves. Perhaps his most important work on non-coniferous cuticles was the account of the stomata of Mesozoic cycads (Bornemann’s subject in 1856) and Bennettitales, attempting to shed some light on the relationships of these groups (Florin, 1933). More recently Florin has devoted most of his time to studying conifer cone-structures, and it is fortunate that he continued to investigate conifer relationships rather than turn his attention to angiosperm cuticles as Edwards (1935) once suggested.
Bandulska (1923) investigated the cuticular anatomy of dicotyledon and conifer leaves found in Eocene deposits at Bournemouth, and was able to place one of the latter into an already known fossil species. The dicotyledons, which were un- identified at the time, were placed into a new fossil form-genus, Dicotylophyllum. In her following four papers Bandulska described a series of cuticles of modern dicotyledons and in several genera she placed species which she had previously described under Dicotylophyllum. These genera were Nothofagus and Fagus in the Fagaceae (1924) ; Amba, Lindera, Litsea and Neolitsea in the Lauraceae (1926) ; Cinnamomum also in the Lauraceae (1928) ; and Tristania and Rhodomyrtus in the Myrtaceae (1931). This work was the first important investigation of modern dicotyledon cuticles to be attempted, and it clearly showed that with a great deal of patience conclusive and positive results can be obtained from this approach.
Odell (1932) published a highly controversial paper which not only condemned the identification of fossil leaves by their gross form (shape and venation), but also severely attacked the use of cuticular characters for this purpose. She apparently originally attempted to identify the older Eocene fossils of the Bournemouth deposits, where Bandulska had first investigated the younger Eocene leaves, but concluded that the variation shown in all characters which were normally used to this end made it an impossible task. The bulk of her paper consists of a series of conclusions drawn from the study of 170 species of modern angiosperms, but un- fortunately she does not state the extent of her sample of each species. Her method of argument is to take each character used in cuticular differentiation separately, discuss the evidence for the fact that this character is known to vary under environ- mental or other conditions, or that completely unrelated species are identical (or, conversely, closely related species quite different) in that character, and conclude that it is taxonomically worthless. One could, of course, equally well do this with any other character that has ever been used taxonomically, but because (for example) Ranunculus ficaria L. possesses about seven to twelve petals one cannot conclude that petal number is of no diagnostic importance, either in this or in any other group.
It is perhaps worth mentioning two of the twelve or so characters which Odell considered, as more or less random examples. With regard to the presence and types of subsidiary cells around the stomata she states that the three main types recognized by Solereder are not found especially in any taxonomic groups, but are distributed variously in different genera and families. Moreover in some species only the stomata on the stems are associated with subsidiary cells. Thus “ the presence or absence of subsidiary cells bordering the stoma can be of little systematic value’’. In the case of non-glandular hairs Odell mentions that the arrangements
ste) CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
and numbers per unit area on a leaf vary tremendously, and are affected by environ- mental factors such as sunlight, humidity, wind and altitude. Their abundance also varies on different parts of the same plant or even of the same leaf. Types of hair often differ within a family, and there is often more than one type even on one leaf, so that the types of hair are not connected with taxonomic categories. She therefore decided that ‘‘ clothing hairs cannot be used in the identification of the vegetative parts of Angiosperms ’’, and that their “ structure has been found of no diagnostic value’”’. Her final conclusions were that “‘ any feature of the epidermis of the vegetative parts of living Angiosperms is unsatisfactory for diagnostic work ’’, and that “‘ the modern method of naming fossil Angiosperms from a combination of the form, venation, and epidermal structure of their vegetative organs is quite inadequate for specific or even generic diagnosis’.
In almost all instances Odell’s statement of fact must be regarded as correct, but it is in some of her extraordinary conclusions that she is obviously in error. If she was attempting to show that it is in general impossible to place a vegetative portion of a plant into its correct family or order on a single epidermal character (or even on a combination of such characters) then one would readily agree with her, but to assert that each of the twelve or so characters is ‘‘ of no diagnostic value ’’, or “cannot be used in determinative work”’, is obviously quite illogical. Odell appears to have overlooked two important basic points in drawing her conclusions. First of these is that the compilation of diagnoses and hence the identification of species should not be undertaken with a single character, but with a combination of as many characters as possible. Although two families, genera or species may not be separable by a single character a combination of several characters will often enable separation to be effected. Secondly a range of form is often more important taxonomically than is a fixed type. For example the possession of a variable number of petals by Ranunculus ficaria is a notable taxonomic feature of this species, not a reason to doubt the systematic value of petal number. Further criticism of Odell’s remarks is unnecessary. In general one would agree with her that cuticular characters are usually not useful at the family level or above, but it is unfortunate that she does not differentiate between those cases where the characters are useful in phylogenetic classification and those where they are useful for identification.
Belying Odell’s claim that stomatal structure ‘‘ cannot be regarded as of systematic value’ just four of several publications may be cited. Carolin (1954), working with Dianthus, found that contrary to the usual findings (see below) neither stomatal size nor frequency could be used to indicate the level of ploidy. However, within the diploid and within the hexaploid groups most investigated species could be separated by a series of size, frequency and morphological characters. Stebbins & Khush (1961) surveyed the ‘‘ stomatal complex ”’ of most families of monocotyledons. They found that characters such as the number and type of subsidiary cells gave valuable clues to the affinities of the plants, so that each family had a predominant type or arrangement. Watson (1962) found that in the Epacridaceae the stomatal distribution and orientation on the perianth and leaves, and also the broad structural types of stomata, were mostly differentiated at the generic or a higher level, and in
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY Wr
the Evicaceae (Watson, 1965) the stomata provided equally important taxonomic evidence although in several instances there was little or no correlation with the usually accepted classification.
Pharmacognosists have frequent need to identify leaves or fragments of leaves used as herbs, and, as the number of species is relatively small, epidermal features have often made identification possible. Timmerman (1927) produced an early analysis of one of the characters used in pharmacognosy, namely stomatal frequency. Using Datura she found that stomata were more frequent at the leaf apex than in the middle, whilst the ratio on the two epidermides also altered in different parts of the leaf. She therefore concluded that stomatal frequency was generally of no use for identifying leaf fragments in this genus, although she was able to separate one species from the rest of those examined by this character. In the same year, working from a different angle, Salisbury (1927) also investigated the constancy of stomatal frequency, adding environmental effects to the positional factors considered by Timmerman. Salisbury found much the same variation as Timmerman, except that the proportions on the two epidermides did not appear to alter. He found that if the proportion of stomata to epidermal cells (the stomatal index, i.e. 100S/E +S) was measured instead of the stomatal frequency the variation produced by environ- ment and position could be completely cancelled, apart from the effect of humidity. He was able, in fact, to measure the environmental humidity by means of the stomatal index. This very simple information was of extreme importance to taxonomists, and the stomatal index is now used as a routine procedure in palaeo- botany (cf. Harris, 1944). Salisbury’s use of the stomatal index had been fore- shadowed, however, by Loftfield (1921), who noticed that the ratio of stomata to epidermal cells often remained constant when the stomatal frequency varied.
A less reliable method of overcoming the variation in stomatal frequency was also used by Baranov (1924), who found that the stomatal frequency of the middle portion of leaves half-way up the stem was equal to the average frequency of all parts of all leaves. This does not take into account the fact that most fossils are only fragmentary, nor the effects of environment.
Working much later than the above, Gupta (1961) also found a considerable variation in stomatal frequency on leaves from different parts of the plant, and demonstrated a negative correlation between this and the lamina size. He thus used an Absolute Stomatal Number, the product of the stomatal frequency and the lamina size, as a constant, this being comparable to the stomatal index.
Sax & Sax (1937) and Sax (1938) reported on some very interesting and now almost classic studies on the effect of polyploidy on stomatal size and frequency. They found a positive correlation between each of these two variables and polyploidy, and could usually separate the diploids and tetraploids in any given series of plants. Modern Floras now often contain characters such as stomatal size and stomatal index in the separation of closely related species where one is a polyploid (e.g. Rorippa nasturtium-aquaticum (L.) Hayek and R. microphylla (Boenn.) Hyland.).
12 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
The great mass of anatomical work which marked the end of the last century seems to have largely ceased with the publication of Solereder’s work and supplement (1908). Rehfous (1917), however, provided a very useful survey of work on stomata. In the 1930’s systematic anatomy, together, in fact, with taxonomic studies in general, began to regain popularity, and from then on the number of systematic studies has increased quite steadily.
Meyer (1932, 1932a) investigated the Alismataceae, firstly for taxonomic and se- condly for phylogenetic reasons. Most of the genera and species Meyer examined could be separated by anatomical features, but he found little evidence suggesting close relation between the Alismataceae and Ranunculaceae ; the stomata, for instance, are quite different. This work was connected with the commencement of a Systematische Anatomie der Monokotyledonen by Solereder & Meyer (1928-33), of which only a very small proportion appeared.
Prat (1932) published an extremely important work surveying the systematic value of grass epidermides, this family without doubt being the most amenable amongst the angiosperms to epidermal classification. Several important taxonomic changes have been foreshadowed in the Gramineae by epidermal considerations, such as the exclusion of Nardus from the Hordeeae or of Eragrostis from the Festuceae, a situation only elsewhere paralleled in some gymnosperms. Many more recent works on this subject have appeared, but the exceptional nature of the Gramineae precludes the necessity of their being considered fully here. The most important work is that of Metcalfe (1960), which is a complete survey of almost all genera and of all the litera- ture. Some recent papers which show the amazing systematic value that can confidently be placed on minute features of grass epidermides are those of Tateoka et al. (1959) and Borrill (1961). In all cases, however, students of grass epidermal anatomy have used the whole epidermis in preference to the cuticle.
In the last twenty years a great many systematic anatomical studies have been undertaken, and the resulting publications provide sections on epidermal and cuticular characters varying in length with their taxonomic importance in the group concerned. Mention may be made of the work of Bailey & Nast (1944, 1948) in the Winteraceae, Illiciaceae and Schisandraceae, Heintzelman & Howard (1948) in the Icacinaceae, Morley (1953) in the Melastomataceae, Tomlinson (1959) in the Musaceae, and Paterson (1961) in the Epacridaceae. The degree of use which can be obtained from the epidermis varies considerably in these groups, as would be expected. Thus Tomlinson was able to use a number of general and special epidermal characters to great effect in delimiting at the generic level, but Bailey & Nast, who were two of the few anatomists to isolate the cuticle and study it as such, found that apart from its very distinctive appearance throughout the Winteraceae it did not furnish good taxonomic characters.
Over forty years after Solereder’s volumes (1908) Metcalfe & Chalk (1950) published along similar lines a modern survey of dicotyledon systematic anatomy. The literature survey is essentially concerned with work done since Solereder’s time, and many details of structure mentioned by the latter are omitted where no further
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 13
examinations have been undertaken. The greatest improvements and additions are to be found in the sections on secondary-xylem structure, upon which emphasis is justifiably placed. With regard to the epidermis and cuticle Solereder’s work is therefore frequently the more informative. In reviewing characters of taxonomic importance Metcalfe & Chalk obviously consider wood and associated tissues to be of the highest importance, and epidermal characters are dismissed in a few paragraphs. With regard to stomata they quite correctly state that the number per unit area is too variable to be of any great importance, but it is strange that they do not mention the usefulness of the stomatal index. More recently Metcalfe has commenced a comparative Anatomy of the Monocotyledons, of which to this date volumes have appeared on the Gramineae (Metcalfe, 1960) and the Palmae (Tomlinson, 1961). Without such an important basis as Solereder’s dicotyledon volumes the compilation of a systematic anatomy of the monocotyledons is a considerably greater task, and when completed will probably prove even more valuable.
In 1954 Metcalfe published a paper which may be compared with one by Fritsch (1903) half a century before. This set out to indicate the main lines along which systematic anatomy could be developed, and in the next year (1955) he summarized recent work on this subject in the monocotyledons, especially the grasses. More recently Metcalfe (1963) has elaborated his opinions on the desirable trends in modern comparative plant anatomy.
From about the time of Bandulska’s work (1923-31) onwards a considerable number of articles on fossil dicotyledons began to appear in the literature, in several cases obviously directly inspired by her publications. Stockmans (1932) reported on two species discovered in Belgium : a Litsea and a Dewalquea. The first genus, a member of the Lauraceae, had been discovered in England by Bandulska (1926), whilst the extinct genus Dewalquea had been previously found in Ireland by Johnson & Gilmore (1921). Discussions by Stockmans and Johnson & Gilmore, in both cases well illustrated, on the affinities of this genus produced little concrete theory, several diverse families being mentioned. Since not all of the species of Dewalquea possessed peltate scales on the leaf it was suggested that they were perhaps representatives of two different genera. Whether or not this is true it must be remembered that other genera known today (e.g. Combretum and Rhododendron) possess scaly and scale-less species.
Hofmann produced a valuable series of papers from 1926 to 1932 (cf. Hofmann, 1932) on plant remains in early Tertiary formations. In her earlier works she attempted to give names of living genera or species to the cuticles, and in 1926 she described fossil plants in ten living genera from these deposits. In 1932, however, she changed to the more advisable system of describing these old leaves in a fossil form-genus. Her choice of name for this was Folium, which is nomenclaturally superfluous owing to Bandulska’s early name Dicotylophyllum for the same purpose. Edwards (1935) justifiably comments that her descriptions of new fossil leaves are somewhat inadequate, but she gives some good photographs.
Kubart (1927) found great difficulty in distinguishing between the fossil leaves of
I4 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
Fraxinus (Oleaceae) and Umbellaria (Lauraceae) which he discovered in North America. He pointed out, however, that had the cuticles been preserved there would have been no difficulty in discrimination as the epidermal structure of the two genera is quite different. In support of this he figured a series of recent species of each genus, showing the great divergence between them. Also working on Ter- tiary plants, Straus (1930) was able to identify a number of species with modern genera, and in the cases of the two better-preserved genera (Juglans and Populus) he gave some good high-powered photographs. Straus pointed out the need for microscopic examination of the cuticle in cases where two fossil leaves appeared to be closely similar.
In recent years the great majority of works on fossil cuticles has been concerned with pre-Tertiary plants, mostly gymnosperms. Perhaps the most important publications have been those of Harris (1942—>), who has covered a wide range of topics on the Jurassic flora of Yorkshire in several scores of articles, many of these including cuticular studies which have often proved of considerable diagnostic value.
Cookson (1953) reported on a relatively recent (early Tertiary) cycad from Australia which, on the basis of its cuticular characters, could be referred to the genus Macrozamia, which still occurs in Australia today. Cuticles of this fossil and of the living M. hope: W. Hill ex F. M. Bailey are shown, and due to the great similarity between them the former is named M. hopeites Cookson.
Apart from Europe and North America some of the most important cuticular studies have emanated from India, and as examples two of the papers on the Indian Glossopteris flora by Srivastava and his colleagues may be cited. Srivastava (1956) produced a most thorough account of all the species of Glossopteris, Gangomopteris and Palaeovittaria, with very clear descriptions, drawings and photographs. All species could quite easily be separated on cuticular characters alone, a whole series of features, concerning epidermal cells, stomata and venation, being employed. Surange & Srivastava (1956) concurrently considered the generic limits in this group of leaves. They concluded that the three genera, which were delimited on frond shape, could be better divided into six genera using the cuticular characters. These six genera did not correspond in any way to the three established genera, and pending further information no new generic names were proposed.
Apparently the only general review of cuticular studies in angiosperms is that of Edwards (1935). He gives an extremely useful survey of the literature to that date, which has obviated the necessity of very detailed accounts of the earlier work in this introduction. The account treats recent and fossil monocotyledons and dicoty- ledons in four separate sections, those on monocotyledons being mostly concerned with grasses. Many of his citations are annotated with some sort of assessment on their importance or validity, but in the case of Odell’s (1932) paper he quite justifi- ably records strong protest with a series of objections, mostly in the form of counter- evidence. One would concur with almost all of his opinions on this and other papers, but as mentioned previously I prefer to attack Odell’s deductions and conclusions rather than her statement of fact. In several instances it is obvious that Edwards
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 15
held almost as extreme ideas as Odell, though in a completely opposite direction. He was clearly convinced of the tremendous and widespread usefulness of cuticular characters in the angiosperms, which is perhaps to be expected of a palaeobotanist whose cuticular studies had mostly been concerned with gymnosperms. Had he made extensive investigations in the angiosperms he would have met, in many instances, with a much wider range of variation than is found in any gymnosperm, and than he seemed to admit as existing in the angiosperms. For instance, in criticizing Odell’s statement that the leaves of Campanula latiloba A. DC. and Inula salicina var. denticulata Borbas are identical, he found that there were in fact good microscopic differences between them, and he also suggested that Odell’s ‘‘ own drawings do not suggest identity of epidermal structure’. Odell’s figures differ only in the thickness and degree of undulation of the epidermal cell walls, and had Edwards been familiar with the range of variation of the cuticles of many angio- sperms he would have realized that most species show a wider range of variation than Odell’s two drawings. Thus, although in fact the two species do apparently differ in cuticular structure (fide Edwards), Odell’s figures certainly do not suggest it. Apart from a few criticisms such as the above, Edwards’s paper is a valuable and critical review. His conclusion may be quoted: “ given careful and critical work on well-preserved material, together with a detailed comparison with a wide range of living forms, results obtained from a study of fossil angiosperm cuticles will be as valuable as those derived from any other fossil remains, and certainly far more reliable than those founded on leaf impressions alone ”’
Edwards did not mention any articles concerned solely with modern dicotyledon cuticles, and the first that the writer has traced is the account by Rao (1939) of the order Magnoliales in the sense of Hutchinson (1926), a very heterogeneous assembl- age of families about which Rao concludes: “‘ From the point of view of epidermal studies, the Magnoliales do not represent a co-sanguinary group, but indicate a convergence ’’. Rao’s paper presents a detailed description of the cuticles of over 50 species representing more than half the genera of the order, but since he admitted that in most cases only a single leaf of each species was examined the results must be treated with extreme caution. Objection must be made to some of Rao’s con- clusions, although a detailed criticism will not be attempted here. Rao implied, for instance, that the cuticular striations found in some species of Magnoliales are phylogenetically related to those of the Cycadales. However, representatives of many dicotyledon families possess cuticular striations which are identical with those of the Magnoliales. Even if cuticular striations in the two orders were phylogeneti- cally related, and this seems unlikely and in any case could never be proved, no such inference could be drawn merely from their presence in both groups. Rao strongly stressed further evidence of a connexion between the Magnoliales and gymnosperms from his stomatal studies. Using Florin’s (1931) two categories of stomatal de- velopment Rao claimed that two genera of Magnoliales, Euptelea and Cercidiphyllum, possessed haplocheile stomata, which according to Rao were otherwise virtually unknown in the dicotyledons yet were typical of all gymnosperms except the Bennettitales, Welwitschiales and Gnetales. Later work has shown many of Rao’s
16 CUTICULAR STUDIES AS AN. AID TO_PLANT TAXONOMY
conclusions regarding stomatal types to be erroneous, and a modern stomatal survey of the Magnoliales is badly needed. Jalan (1962), for example, has found that the stomata of Schisandra are haplocheilic (as, in fact, they are in a great many dicotyledons), and the same is true of Gnetum (Maheshwari & Vasil, 1961). In Magnolia and Michelia species the stomata on the leaves are syndetocheilic, but those on the flowers haplocheilic (Paliwal & Bhandari, 1962). The need for develop- mental studies rather than observations of mature structure cannot be over- emphasized, and the approach of several Indian workers in this field is encouraging (cf. Pant, 1965). Further reference to this is made in the section on stomata.
Ahmad (1964, 1964b, 1964c) has investigated the cuticular and epidermal anatomy of 17 genera of Solanaceae, and found that a number of characters concerning the stomata, trichomes, epidermal anticlinal cell walls and cuticular striations are very useful in identification. The same characters enabled 28 species of Solanum and 23 of Cestrum to be differentiated.
Sinclair & Dunn (1961), on the other hand, working in the monocotyledons, appear to have made conclusions regarding the usefulness of cuticular characters at the family and generic levels which are not justified on present information.
It is perhaps largely by chance that the Lauraceae have been studied more frequently than any other dicotyledon family with regard to their cuticular charac- ters. Studies already mentioned have been those of Bandulska (1926, 1928), Kubart (1927) and Stockmans (1932), and there have been other less important ones besides. Kostermans (1957), in his taxonomic studies of the Lauraceae, mentioned the useful- ness of cuticular characters, but pointed out the care necessary in the interpretation of the results of these investigations. He was referring mostly to Bandulska’s work. Marlier-Spirlet (1945) investigated the epidermal anatomy of Cinnamomum more thoroughly than did Bandulska, and produced many figures. He also showed vertical sections of the leaf to demonstrate the derivation of various features of the cuticular preparations which were not readily interpretable from surface views alone. It was found to be possible to separate many of the eighteen species in- vestigated, and three broad groups were recognized, containing four, three and eleven species respectively. The most useful taxonomic criteria appear to be the shape of the upper epidermal cells and the types of stomata. A continuation of this study would be of considerable interest and value. Dilcher (1963) undertook a “cuticular analysis’ of an Eocene species of Ocotea, and after a study of its variation and of the living taxa concluded that it should be considered a distinct species.
Martin (1955) used cuticular anatomy to a most novel and useful end to which there appear to be no subsequent references. Previously Parkinson & Fielding (1930 ; fide Martin, 1955) had compiled notes of some epidermal characters of the plants used as cattle food, using the solid leaf fragments. In order to study the diet that sheep choose in various pastures Martin examined the rumen contents and faeces of these animals to ascertain quantitatively the species of plants re- presented by their cuticles. In the upland pastures in which the work was pursued
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 7
the number of species was rather limited and few species were closely related ; thus Martin was able to separate all forty likely plants by cuticular anatomy and deter- mine them quantitatively in the faeces. He gives brief descriptions and photographs of twelve representative species. Of these forty species only sixteen were recognized in the stomach contents or faeces, including four grasses, two sedges, three rushes, two mosses and five dicotyledons. Except in the case of two species of Juncus each genus was represented by only a single species, although other genera were represented by two or more species among the originally investigated group of forty plants. It seems likely, however, that some or many mesomorphic herbs would never be represented due to their extremely thin cuticle, although the only known cutinase enzyme has been isolated for certain solely from Penicillium spinulosum Thom (Heinen, 1960, 1961).
This work is a further indication of the economic use to which cuticular studies may be put, and emphasizes the need to discover the basic principles underlying the possible application of this method in various directions.
§3. ANATOMICAL INTRODUCTION
Some idea of the precise relationship between the cuticle and the underlying epidermal cells is essential to a full understanding of the nature of the patterns seen on acellular cuticular preparations and their interpretation in terms of a cellular layer.
Although Bornemann (1856) obviously understood the cuticular-epidermal relationship fully the first good general anatomical accounts are those of de Bary (1871, 1884), and Héhnel (1878) appears to be the first to have described the chemistry of the cuticle. During the past thirty years our knowledge of the cuticle and of its essential constituent, cutin, has steadily grown, accelerated by two major technical advances. The first of these, polarizing microscopy, became available in the 1920’s and its use is shown in the work of several Continental writers. The second, electron microscopy, a post-war development, has been utilized in cuticular studies mainly by Roelofsen and a number of American workers.
In all vascular plants the basic chemical component of all the cell walls is cellulose, whose long-chain molecules exist as long compound microfibrils. In the inter- fibrillar spaces are a number of substances, including other polysaccharides, fatty materials, water and inorganic and organic solutes. Separating two such walls of adjacent cells is a cellulose-free layer known as the middle lamella, consisting basically of calcium and magnesium pectates. The outer face of an epidermal cell, however, is unusual in being adjacent to the environment rather than to another cell, and it is on this face that the cuticle is found. The precise structure of the latter was first ascertained by means of polarizing microscopy (e.g. Meyer, 1938 ; Roelof- sen, 1952) and was confirmed later by electron microscopy (Roelofsen, 1959).
External to the normal cellulose cell wall of the outer face of each epidermal cell is found a thin layer of pectic material which is presumably continuous with the middle lamellae of the anticlinal walls of the underlying epidermal cells (Fig. 1).
18 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
Outside this is a usually two-layered wall, which according to the nomenclature of Roelofsen (1952, 1959) may be termed the cuticular membrane. The inner portion of this, the cuticular layer, is composed essentially of a cellulosic framework between the microfibrils of which are encrusted large amounts of cutin. The outer part is a usually thinner layer, the cuticle proper, which lacks cellulose and is composed mainly of cutin, which is adcrusted on to the cuticular layer. Esau (1953) refers to the process of encrustation as cutinization, and that of adcrustation as cuticulari- zation. The above terminology is adopted hereinafter. In a number of plants, usually those with leathery leaves, the four layers exterior to each epidermal cell lumen are simplified insofar as a cellulose-free pectic layer is absent, the cutin-free innermost cellulose wall gradually merging into the outermost cellulose-free cutin layer. Rarely are all four layers as sharply defined as suggested in Fig. 1, but in general it can be said that the innermost layers lack cutin and the outermost layers lack cellulose. Chemically cutin consists of a number of highly polymerized long- chain hydroxy-fatty acids.
cuticle
cutin only
PSE TENS :
cutin + cellulose
4 path
oh ets ay diame, Pars ak ais
Pa increasing cutin
2*,* increasing cellulose
cuticular layer
pectic layer
1 cellulose ? cellulose
cellulose only only cell wall
? possible termination of middle lamella
cuticular flange \
middle lamella ee wee
|__— middle lamella
. ? possible boundary between cutin-cellulose and cutin-free layers
|
|
| a |
f middle lamella |
cell outline Ge
observed on J vas
cuticular preparations
ies
Ce ere ee outline of
subepidermal cells
ji ae, outline of
epidermal cells
ae original extent
of cell walls
extent of original cell wall
Fic. 1. The organization and interpretation of cuticular membranes and preparations : sections of epidermides with normal and abnormally extensive cutinization are shown, together with diagrams showing the significance of the patterns observed on the respective cuticular preparations.
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 19
Where the outer walls of two epidermal cells meet, the pectic layer, if present, may form a T-shaped juncture with the middle lamella of the vertical walls of the cells. In most cases, however, the cuticular layer projects down between the two cellulose cell walls for some way, forming the three-dimensional network of so-called cuticular flanges (Fig. 1). The length of the cuticular flanges varies from their virtual absence to their reaching all the way to the inner wall of the epidermis. In some leaves a greater proportion of the epidermis may be cutinized, in extreme cases the cell walls on all faces of the epidermal cells and even the outer walls of the subepidermal layer being encrusted with a certain amount of cutin. This internal cutinization is a feature of many xeromorphic plants.
Cuticular flanges are formed by the cuticular layer alone, and the cuticle remains a thin uninterrupted membrane. Indeed, the cuticular membrane as a whole is a continuous and virtually extra-cellular sheet, and should be thought of as something additional to the cell wall. It completely covers the leaf, usually including any trichomes which may be present, being interrupted only by the stomata. In a herbaceous plant it appears to surround the whole of the aerial shoot, including the apical meristems and flowers, although some workers have stated that certain trichomes lack a cuticular covering. With regard to the root system there is less certainty, opinions differing whether it is only the root tip and root hairs or nearly all the root that lacks a cuticular membrane (Lee & Priestley, 1924 ; Priestley, 1943 ; Esau, 1953). At each stoma the cuticular membrane follows the contours of the guard-cells, and often ends on their inner walls. In some cases, however, it is more extensive, covering also the adjacent mesophyll cells. Cutinized cell walls have also been found in more internal regions of the plant (cf. Arzt, 1934), and Lamarliére (1906) found a “‘ cuticule interne ”’ lining the aerenchyma of water plants.
Right from the time that the leaf is a minute primordium in the apical meristem it is covered by the rudiments of the cuticular membrane in the form of a very thin and pliable yet continuous fatty layer. These fatty substances obviously migrate through the cell wall from the cytoplasm, although there are no visible pores in the former for the passage of the cutin. On exposure to air the cutin hardens to a varnish-like cuticle by polymerization. A reasonably hardened cuticular membrane is present long before the epidermal cells have attained full size, and the problem is thus one of increasing the surface area of a relatively non-stretchable substance. Some workers suggest that the epidermal cells may grow in surface area only at their edges, so that the already hardened cuticular membrane remains undisturbed (e.g. Schieferstein & Loomis, 1959).
Not only is wax present within cell walls and cuticular membranes but it is also found in a fairly pure state on the surface of the cuticle of many plants. In recent years these deposits, which are the cause of the “‘ bloom ”’ on the surfaces of many leaves, have been studied in considerable detail, since they are important to the understanding of water repellency. Fairly comprehensive accounts are given by Mueller et al. (1954), Schieferstein & Loomis (1956, 1959), Scott et al. (1958), Juniper & Bradley (1958), Roelofsen (1959) and Juniper (1959, 1959a, 1960). It is
20 CUTICULAR, STUDIES AS AN AID TO PLANT TAXONOMY
interesting to note that the type of deposit (frequently variously shaped rods and plates) appears to be largely characteristic of each plant species. The deposits, when washed, dissolved or brushed off growing leaves, re-form quite rapidly and in exactly the same form as previously, and although their abundance varies with the environment and other factors their non-varying form is obviously in some way genetically regulated. What determines the form which the wax assumes is of course unknown, and moreover different parts of the plant (e.g. stem and leaf epidermides) may have totally different patterns. This subject is obviously of some taxonomic interest though doubtfully of any widespread practicability, and it is not considered further in this review.
In studying the cuticular membrane various methods may be employed to isolate it from the rest of the leaf. All make use of one or both of two important structural features of the epidermis which will have become apparent from the preceding paragraphs : firstly, the cuticular membrane is in most plants separated from the cellulose cell wall within by a thin layer of pectic material ; and secondly, the cuticle and the bulk of the mature cuticular layer are essentially composed of a substance (cutin) of very different chemical structure from the main constituent of the cell wall (cellulose). Isolation of the cuticular membrane may be effected with either of two types of substance : an enzymic preparation, either a pectinase which digests the pectic layers and so frees the cuticular membrane from the underlying cells and these from each other (Orgell, 1955), or a cruder hemicellulase-cellulase- pectinase mixture which additionally digests the cell walls (Skoss, 1955) ; or a macerating solution, composed usually of strong acids, which completely dissolves away all polysaccharide leaving only cutin unaffected. Very much later the acid also dissolves the cutin. The enormous resistance of cutin to the effects of age, micro-organisms and chemicals is, in fact, its greatest attribute with regard to the present study. Only one organism (Penzcillium spinulosum) is known to be able to break down cutin (Heinen, 1960, 1961). The advantage of the above enzymic method is that the cutin is always completely unchanged, and in biochemical studies of the cuticular membrane this method is the only one to be recommended. On the other hand the macerating solutions are very much quicker and simpler to use, and the speed of reaction may be adjusted more easily. The cuticular membranes are also better for microscopic examination since they are considerably cleaner.
The upper surface of such preparations is smooth, or minutely striate or papillose, whereas the lower surface is protracted into a reticulate series of flanges which represent the position of the epidermal cell walls. When viewed from above or below the cuticular membrane is much thicker at the position of these flanges, which therefore show up, especially if the cutin is stained, as a series of narrow darker lines representing the shapes of the original epidermal cells (Fig. 1). It is important to realize that these lines only show the position of the central region of each double cell wall, and that the original walls were present for a considerable thickness on either side of them. In those cases where there are no cuticular flanges the cuticular preparations will of course lack the cell outlines, but it is apparently rare in woody plants for at least the faintest and most fragmentary of outlines not to be present.
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 2I
As mentioned previously some plants have a heavily cutinized epidermis, the whole of the outer, vertical and inner walls and even the top of the vertical walls of the subepidermal layers being encrusted with cutin. In these cases the preparations have cuticular flanges which are very long and join up at their bases, and in surface view a second fainter outline, that of the subepidermal layer, may be visible below the epidermal outline. There is some evidence that the cuticular membrane is more or less completely lacking in cell outlines, or is not even able to be isolated in a continuous sheet, in many though by no means all herbaceous dicotyledons. In these cases the whole epidermis, or plastic moulds of it, must be studied.
§4. MATERIALS AND METHODS
For the purposes of obtaining cuticular preparations leaves may be used in any state (fresh, spirit-preserved or dried) and it is fortunate that herbarium specimens even over 100 years old produce at least as satisfactory preparations of the cuticular membrane as fresh leaves, exactly the same techniques being used. It is important, if studies of cuticular membranes are to be used as routine taxonomic procedure, that the methods involved in their isolation should be simple, quick and reliable.
With regard to the choice of material it is essential to ensure that a representative sample of each species is examined, and this involves a considerable amount of preliminary investigation into the variability of the taxa concerned. Thus the amount of material which can be considered truly representative will vary greatly, but in the present study I found it necessary to examine only one leaf of each specimen, except in cases of obviously different-aged leaves. The number of specimens of each species which was investigated varied according to the amount of material available, the degree of macroscopic variation apparent, and the ecological and geographical range of the species concerned. Three to five specimens were usually found to be a sufficient sample, but care was taken to select those with the widest morphological and ecological variation. In many cases, however, fewer specimens were available, while on the other hand special circumstances sometimes necessitated the examination of up to twenty.
The first task in investigating the taxonomic use of a new character is obviously to check it with the other characters already used, and for this correctly named material is absolutely essential. In the present series of investigations the families concerned are rather well covered in the literature, although the lack of knowledge concerning two genera necessitated a complete revision of all the species (Exell & Stace, 1963), and in some families this problem is so marked that it would become the limiting factor. Other genera of the three families studied still badly require revision. Whatever the state of knowledge of the classification of the taxon concerned differences of opinion may exist and occur, and a careful record of the actual specimens examined is essential. All the permanent slides prepared by me are marked with the collector, the number, year and country of collection, and the herbarium from which the material was obtained. This enables the identifica- tion to be checked whenever required.
22 CUTICULAR STUDIES AS) AN AID! TO PLANT TAXONOMY.
Many different macerating techniques have been employed to isolate the cuticular membrane. Enzymic maceration is not suitable for taxonomic work as the condi- tions (temperature, pH, concentration) have to be carefully controlled and the method is very slow, and in fact in many thick leaves a clean preparation cannot be obtained. Of the milder chemical macerating solutions Eau de Javelle (CaCl, and K,CO, solutions) is the most used (Nathorst, 1907 ; Bather, 1908 ; Bandulska, 1923), but for present purposes strong acids are preferable. Schulze’s Solution (Schulze, 1855), concentrated HNO, with a few KCIO, crystals added, is the most popular, and several improvements and variations have been suggested (e.g. Bather, 1908 ; Thomas, 1912 ; Thomas & Bancroft, 1913 ; Gothan, 1915 ; Harris, 1926). I found, however, that a slight modification of Jeffrey’s Solution (Jeffrey, 1917), equal parts of 10% chromic and bench concentrated nitric acids, gave the best preparations in the shortest time, and it has been used without exception. This mixture was variously diluted to conveniently regulate the time taken for macera- tion to about sixteen hours, i.e. overnight, and in all 250 or so species examined the method was entirely successful. Care must be taken not to allow maceration to continue for too long since over-macerated cuticular membranes are unsuitable for future manipulation and examination.
Parts of leaf macerated
Cuticular preparations obtained
arts irs
, u iad
iy eos
| ae
Te margin
midrib
u = cuticular membrane from upper epidermis
| = cuticular membrane from lower epidermis
Fic. 2. Diagram illustrating the origin of cuticular preparations.
For purposes of conformity identical regions of each leaf were macerated (Fig. 2), so that three pieces of cuticular membrane were made into separate permanent mounts.
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 23
In some cases it is necessary to examine the cuticular pattern of species not avail- able for maceration, or of species whose cuticular membrane cannot be removed as a continuous sheet. For this purpose the preparation of a plastic epidermal cell mould can prove of value, the plastic, usually cellulose acetate or nitrate, being either painted on to the leaf as a solution or pressed on as a thin sheet and then partly and temporarily redissolved (e.g. Nathorst, 1907, 1907a ; Bather, 1907 ; Thomas, 1912 ; Long & Clements, 1934 ; Sinclair & Dunn, 1961), making use of the fact that the outer surface of the cuticle usually follows the contour of the epidermal cells to some extent. The version of this method considered most suitable is that of North (1956), who used the formula 2 gms. of cellulose acetate and o-7 gms. of gentian violet in 100 mls. of acetone, the stain improving the contrast. Herbarium specimens, whether dry or boiled, rarely give such good results as fresh leaves, but they are usually quite usable. The method is useless for most thick- cuticled leaves, which do not show the lines of the epidermal cells on the outer cuticular surface, and for pubescent leaves, although in the latter case the hairs can sometimes be removed. Occasionally transparent cell moulds are found already prepared on herbarium specimens in the form of flakes of old dried herbarium gum, which may show the epidermal cell outline very clearly.
Frequently sections of leaves must be prepared in order to interpret the observed cuticular patterns. Fresh or spirit-preserved leaves are naturally easier to section, but herbarium leaves of great antiquity, after boiling in water for a while, usually provide good sections when cut by hand or with a freezing-microtome. It is often more expedient to isolate the whole epidermis rather than the cuticular membrane alone, this method being particularly applicable to Gramineae. Lactic acid is generally used to separate the epidermis from the subepidermal layers, and the epidermis may then be stained with any general botanical stain (Clarke, 1960).
The technique employed to prepare permanent mounts was again designed to meet the three requisites of speed, simplicity and reliability. The best stain was found to be Sudan IV (1% solution in 70% alcohol) which has the advantage of being progressive, so that overstaining does not occur, but the disadvantage that dehydration and thus mounting in canada-balsam or similar media is impossible since the stain is re-dissolved from stained preparations by absolute alcohol. Thus glycerin-jelly was used as the mountant, and although this is not completely per- manent most preparations are still perfect after at least thirty years and can at any time be remounted. Since the cuticular preparations are acellular sheets rather than pieces of tissue the number of stages from water to alcohol, alcohol to water, and water to glycerol can be reduced to a minimum, and each stage to a few seconds. The procedure adopted for preparing permanent mounts may be summarized : leaf fragments macerated in suitably diluted Jeffrey’s Solution in an open watch- glass ; cuticular membranes washed in water, re-macerated in undiluted solution and re-washed in water if some cellular tissue remains, and transferred to a slide in 70% alcohol ; 70% alcohol replaced by Sudan IV solution for fifteen to thirty minutes ; slide irrigated with water, with 50% glycerol, and then with 100% glycerol ; and cuticular membranes finally mounted in glycerin-jelly.
24 CUTICULAR STUDIES AS AN AID TO-PLANT TAXONOMY
Cuticular preparations can be kept indefinitely, before staining and mounting, in 70% alcohol ; placed in water they become excellent cultures of bacteria, fungi and protozoa (which feed on the adhering polysaccharides, not on the cutin) in a day or two.
§5. SURVEY OF CUTICULAR CHARACTERS
The present section is a catalogue of the main characters which are present in cuticular preparations, together with some idea of the extent of their variation from species to species. In general the ranges of variation discussed indicate those normally encountered without examining plants of an extreme facies or with a very specialized adaptation, so that no mention is made of unusual extremes of structure or of features of very restricted occurrence. Any combination and number of these characters may be of diagnostic value in a particular group of plants. The first task is thus to survey the features found on cuticular membranes, and then to investigate the degree of variation each exhibits. After analysing the causes of variation the systematic value can be ascertained. Most of the comments made have been drawn from my experience in investigations of the three families mentioned previously, and, whenever relevant, examples are usually given from these groups.
Such a general and, it is hoped, purely objective survey of cuticular characters has not been previously compiled, and thus the different terms which have been introduced into the literature are quite uncorrelated. In many cases, for instance, several descriptive terms exist for a single situation (e.g. cuticular flanges, teeth, pegs or ribs). When an acceptable and non-ambiguous descriptive term is already available this has been adopted, but in some cases terms had to be coined, or existing unsuitable ones replaced. All accepted terms are printed in italics, and it is hoped that this will encourage nomenclatural stability.
Of the characters observed during microscopic examination of cuticular prepara- tions some are quite easily observed on whole leaves with an ordinary hand-lens, or with the naked eye. In these cases microscopic examination usually gives a more quantitative or precise assessment of the character, although some points (such as the colour or “ texture’ of the indumentum, e.g. silky, bristly or velvety) may be lost with this medium. Other characters are wholly microscopic in nature, and it is convenient here to treat the two groups separately.
A. MACROSCOPIC CHARACTERS.
The four features discussed here are treated only from the microscopic viewpoint, although preliminary study of a leaf with a hand-lens or low-power binocular microscope should always be undertaken.
(1) Leaf venation.
The venation of a leaf, second to its general shape, is frequently the most obvious character, and descriptions of all new plants contain (or should contain) a note of its organization. The type of venation is often typical of broad taxonomic groups. In
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 25
general, dicotyledons have leaves (or leaflets) with a single central main vein which is a continuation of the petiole, if this is present. This is known as a midrib, costa or main vein. In some leaves two or more main veins may be present, although that situation is more characteristic of monocotyledons. Branching from the midrib are a number of lesser veins or nerves of successively smaller order. Those branching directly from the midrib may be termed Jateral veins, and they are usually very numerous. However, a limited number, from two or three to a score or more, are generally much more conspicuous than the rest and are here described as major lateral veins, as opposed to the less distinct minor lateral veins. In most descriptions “lateral veins’’ usually refers to the former only. The number of major lateral veins, their spacing, whether they arise in pairs or independently on either side of the midrib, the angle at which they arise, the curvature along their length, and their method and position of ending are all taxonomic features.
midrib
minor lateral vein
major lateral vein
was IOS KY MLM YZ < D> LMALZSY LIES IMGT A SEALY A SARA SS TPES secondary vein veinlet termination tertiary vein
venule
Fic. 3. Terminology of leaf venation.
Such a venation system is pinnate, but palmate systems also exist. In the latter case there are a number of main veins all arising from a single point and diverging outwards, and each usually has an accessory venous system resembling that of a pinnately veined leaf.
Generally, arising from the lateral veins is a series of secondary veins, from these tertiary veins, and so on. Unlike the branching of a tree, however, the smallest veins, the venules or veinlets, in most cases do not possess terminations, but usually join up to form an extensive venule reticulum. The number of orders of veins between the midrib and the venules is usually one or two only, but it may be more. The term venule is reserved for the smallest veins with no directional tendency. What-
26 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
ever the number of orders of veins, however, there is in these types of leaves usually a system of venules which emanates directly from the midrib, and almost always one from all other orders of veins. The venation system (Fig. 3) may thus be represented :
Secondary Tertiary a a veinn——————>_—-veins »veins, etc.
Venule reticulum
The above systems of venation, terminated by a venule reticulum, are termed closed, but open systems, where a reticulum is not present, are found in some dicotyledons.
The spaces in the venule reticulum are known as areolae, veinlet-islands, vein- islets or inter-reticular spaces. Their shape and size vary considerably and, although difficult to measure, are often the cause of the different appearance of various leaves. Levin (1929) found that the average “ vein islet area ’’ is of considerable taxonomic importance.
Although most, or all, venules join up to form the venule reticulum there are often a number which end blindly, projecting into areolae, these being termed veznlet terminations (venule terminations). Hall & Melville (1951) calculated the “ veinlet termination number” (number of veinlet terminations per square millimetre of leaf surface) and found it of diagnostic importance. It was stated to be uncorrelated with the number of areolae per square millimetre, and less variable over different parts of the same leaf and hence of greater systematic value. Later work by the same authors (1954) confirmed that the veinlet termination number was more constant than the number of areolae, but found that the two showed a slight positive correlation, and both a negative correlation with leaf size. Gupta (1961) improved Hall & Melville’s methods by devising the “absolute vein-islet number” and “absolute veinlet termination number ’’, the product of the respective numbers and the leaf area. This, of course, cancels the variation due to leaf size and, although more laborious to calculate, should always be used when whole leaves are available. In certain disciplines, such as pharmacognosy and palaeobotany, however, whole leaves are often not available ; in these cases the non-absolute numbers should be used with the greatest caution.
Leaf venation is treated from a different aspect below, when its position as a cuticular character will become clear.
(2) Distribution of trichomes.
The very different degrees of pubescence shown by plants have given rise to a great variety of descriptive terms (e.g. hirsute, pubescent, pilose, ciliate, puberulent, etc.) which unfortunately have no universally recognized meaning. Using cuticular
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 207
preparations the precise distribution and frequency of hairs and other trichomes can be measured, i.e. the individual trichomes can be counted and their positions noted. Taking into account the enormous variation that is encountered, only differences of a considerable degree are likely to be of taxonomic value, and the extremes of a range must be noted.
Precise data may thus sometimes replace such statements as “ almost glabrous ’ or “sparsely pubescent ’’, but in general their compilation will not be worth the labour involved. Besides variation apparently uncorrelated with other factors the degree of pubescence is dependent more than most characters upon the environ- mental conditions and the age of the leaf.
Of much more value than simple trichome frequency is the relative frequency of trichomes on different parts of the same leaf. As is well known, hairs are usually more abundant on the midrib and veins of a leaf, and these regions are usually the last to become glabrous as the leaf ages. However, this is by no means always the situation. In the Combretum psidioides Welw. aggregate, for example, there are three taxa (treated by Exell (1961) as subspecies of C. psidioides) which differ only in their distribution of indumentum on the lower leaf surface : one taxon has a densely pubescent venule reticulum and sparsely pubescent to glabrous areolae (subsp. psidioides) ; the second a glabrous venule reticulum and tomentose areolae (subsp. kwinkiti (De Wild.) Exell) ; and the third a tomentose venule reticulum and areolae (subsp. dintert (Schinz) Exell). Moreover, a further species in this aggregate, C. grandifolium F. Hoffm., differs not only in its larger leaves but also in having glabrous areolae and a sparsely pubescent to glabrous venule reticulum. This taxon, demonstrating the fourth combination, appears to be a further sub- species of C. psidioides. Dr. Exell and I have found that all four taxa belonging to this group have apparently no other constant differences, either macroscopic or microscopic, than the distribution of hairs. In many plant groups the distribution of hairs on the leaf margin is frequently also a diagnostic character.
The broad type of trichome, e.g. sessile glands, stalked glands, branched hairs, peltate hairs, long straight hairs or short curly hairs, etc., is also a character often used without resorting to the use of a microscope, although this is not to be encour- aged. Small differences, often vital, may be not apparent with a hand-lens or binocular microscope, and very distinct taxa can easily be classed together. The key to the genera of Sapotaceae by Hutchinson & Dalziel (1931), for instance, is completely misleading due to a mis-statement on the type of hair present. I have found that the hairs described as ‘‘ simple (not medifixed) ”’ (as opposed to medifixed, stalked and thus “ T-shaped ”’ hairs) are in fact medifixed but sessile. Such errors would have been avoided if a microscope had been employed. There have been many erroneous statements concerning the trichomes of the Combretaceae. A hand-lens should only be relied upon when two or more very different types of trichomes are being searched for (e.g. in Epilobium), or special distinctive trichomes are sought (e.g. in Urtica).
As previously mentioned, characters of the texture or overall appearance (includ- ing colour) of the indumentum are lost in cuticular preparations.
,
28 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
(3) Cork-warts and similar structures.
Very frequently there are to be found on the cuticular preparations of leaves structures which are usually referable to wounds of one sort or another, but which, in some cases, represent cork-warts or related structures. Cork-warts are not abundant in nature, and they are usually constant in occurrence, so they are frequently useful in identification. In the mangrove genera of the Rhizophoraceae, for instance, they are present in all species of Rhizophora but in none of the other three genera. In West Africa, moreover, they are larger in R. mangle L. than in the other two species of that genus found there, and are said to be diagnostic in the field (Keay, 1953). They appear as small convex warts of corky tissue up to about 2 mm. across.
Microscopically the position of a cork-wart on the cuticular membrane is marked by a circular or less regular hole surrounded by modified epidermal cells (Plate 1 A). The latter are usually, though not always, thicker-walled and much smaller than the normal epidermal cells, and they are arranged in strict rows radiating for some distance from the hole. It appears as if the radiating cells are meristematic, being produced torepairthehole. These areas are thus very characteristic and conspicuous.
Unfortunately three types of wounds are extremely similar to the cork-warts. Purely mechanical accidents are marked by variously shaped, but usually long and scar-like, areas, the surrounding cells being identical to those of the cork-warts. In many cases, presumably in the case of more ancient wounds, or perhaps when only part of the outer wall of the epidermis has been removed .or damaged, a very thin cuticular membrane, usually scarcely stainable, has grown over the wound, this being either amorphous or with very thin cell walls. The cells on these areas, if present, are usually normal-sized, although small cells would have been expected if they were meristematic. Insect punctures are also frequently found on leaves, these more often resembling cork-warts than accidental injuries, and probably not being distinguishable from the former. They are of the same order of size as a small cork-wart, and usually smoothly circular in outline. They were also reported by Orgell (1955), who observed an identical structure. Moreover he found that they frequently became occupied by corky or gummy plugs, showing a very close analogy to cork-warts. The third type of wound which I have observed is a fungal attack, found in the form of hemispherical shining excrescences on both leaf surfaces of one specimen of Buchenavia kleinti Exell. The hole left by these fruiting bodies after maceration is surrounded by the radiating rows of small cells typical of cork-warts (Plate I B).
Although cork is more or less resistant to the macerating solutions used in the present study, owing to its chemical similarity to cutin, it is not preserved on the cuticular preparations as the wart or plug is not connected to the cuticular membrane, but falls away during maceration. Where a wound has healed over, and occasionally a cork-wart has dropped out, however, the tissues may become covered with a thin cuticular membrane, as observed above.
Thus, although wounds have no intrinsic taxonomic value, they are important as they closely resemble cork-warts, and thus render the latter considerably less
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 29
useful in systematics. This equally applies to macroscopic and microscopic studies.
Other types of structures found on leaves, although often not macroscopic characters, are sufficiently similar to cork-warts to be mentioned here. In many cases they may be referred to excretory glands (e.g. salt glands, chalk glands, sessile hydathodes, etc.), which may be macroscopically very conspicuous. The actual gland is frequently only very thinly covered with a cuticular membrane, and the adjacent cells may be radially arranged or otherwise modified. Holes may also be found in the cuticular membrane due to the presence of modified epidermal cells (e.g. calcified or silicified cells) which have no cuticular covering, but these are usually surrounded by normal epidermal cells, as are the terminations of long idioblasts. Cells containing crystals frequently break through to the surface of the leaf, and these are often surrounded by radially arranged cells, as in wounds. In this connexion the minute pimples and translucent spots common on some leaves and often used by taxonomists may be mentioned. They are not epidermal characters : the former are due to abnormally large epidermal or subepidermal cells, usually containing crystals ; the latter to similar cells which interrupt the mesophyll. In fact in most cases the translucent spots correspond with small pimples, although either one may be present in the absence of the other, or rarely both exist together on the same leaf but are unconnected.
Wound-like structures are frequently present on cuticular preparations in such a regular manner that they are almost certainly not wounds, but no macroscopic equivalent is visible and no microscopic subepidermal structures seem to be connected with them. In Terminalia catappa L., for example, all specimens examined showed ragged holes along the leaf margin (Plate 1 c), and T. chebula Retz. possesses similar areas there and on the lower epidermis (but not the upper). In T. bursarina F. Muell. similar areas are present in the major lateral vein axils on the upper epidermis, corresponding to the presence of domatia on the lower epidermis. They are caused by the growing of the domatia (see below) right through the leaf to the upper surface.
Besides hydathodes, other water-secreting organs are the water-stomata, which are usually distinctly different from the normal stomata in some way (e.g. by their smaller or larger size). They are, of course, not macroscopic structures, although they are in some families united into groups which appear as small surface spots (Solereder, 1908). In the two Combretaceous mangrove genera, Lumnitzera and Laguncularia, large stomata which may be water-stomata are found scattered over the leaf surface. Usually they are surrounded by enormous subsidiary cells, and around the latter the epidermal cells may be radially arranged (Plate 1 D). In extreme cases the radially arranged cells form an extensive area resembling those around a wound or cork-wart, and indeed in some instances the large stoma has disappeared, leaving a hole. Solereder (1908), in fact, states that stomata may become underlaid with a development of cork which ultimately erupts to give rise to a cork-wart. Cork-warts have not been recorded from the Combretaceae, however, and any connexion between the observed structures and cork is conjectural.
30 CUTICULAR STUDIES AS AN AID:TO PLANT TAXONOMY
In addition Lwmmnitzera possesses wound-like areas along the leaf margin (cf. Terminalia catappa) which appear to be distinct from those developing around the large stomata, and which are also unconnected with the shallow marginal pits thought to be domatia.
Finally, the presence in many instances of thin-walled areas, often only a few cells in size, on the leaf epidermis may be noted. In some cases the component cells are much more or much less undulate-walled than the rest of the epidermal cells. Their origin is unknown.
”
It may thus be seen that the presence of “ spots ’’, “‘ pimples ”’ or “ warts ”’ on leaves may be due to any one of a whole range of epidermal (or subepidermal) structures, which only a microscopic examination will determine. Their study is, however, hampered by the presence of various types of wounds and punctures of external cause which may extremely closely resemble the structural features and thus limit the determinative value of the latter.
(4) Domatia.
The term domatium is used to denote a small cavity of very varying form, found on almost any vegetative part of the plant, which is inhabited, or thought to be inhabited, by some type of animal. Domatia have been almost completely neglected from a taxonomic viewpoint in the past, and so limited is our knowledge of them that they are treated rather fully here. It will be seen that on occasions they may be of great diagnostic significance.
Domatia vary from cavernous passages in rhizomes, stems or petioles to minute pits or hair-tufts on the under surfaces of leaves. The former are usually occupied by ants, and the literature concerning these so-called myrmecodomatia is fairly extensive (see Uphof, 1942). The smaller domatia, which may be up to about 4 mm. in diameter, are often said to be occupied by mites or similar arthropods, although good evidence for this is usually lacking. They are, however, called acarodomatia, a term suggested by Lundstrém (1887), and the plants on which they occur acarophytes or acarophilous plants. Elliot (1911) stated that the mites could be seen scurrying away from the appropriate regions of the leaves of many common English trees when the former were disturbed with a needle, and Mani (1964) reported that acarodomatia may be inhabited by mites such as species of Tarsonemus.
The literature relevant to acarodomatia is remarkably scanty, the only important papers being a biological study by Lundstrém (1887) and a review by Penzig & Chiabrera (1903). The latter account is excellent from many aspects, and thus any sort of systematic review is unnecessary here. The authors reported acarodomatia from about 425 species and 175 genera in 44 families, all woody dicotyledons. They are most abundant in the Rubiaceae, in which Penzig & Chiabrera recorded 54 acarophilous genera. As these authors predicted, since 1903 many additional genera have been shown to possess acarodomatia, but few are of very special note.
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 31
An exception is Dioscorea, which is the only monocotyledonous genus and also the only herbaceous one known to possess acarodomatia.
Apart from the fact that they are more or less confined to woody dicotyledons there is no connexion whatsoever between the type of leaf and the presence or type of domatia. They do perhaps tend to be rarer on leaves with inconspicuous venation, but they are not absent from these and any generalization would have a number of exceptions.
The commonest types of acarodomatia are small pockets or pits in the axils of the main veins with the midrib on the lower leaf surface. Since the whole domatium, whatever its shape, is lined with epidermis throughout, cuticular preparations provide excellent transparent models of the domatia and afford the best method of studying their anatomy. The preparations can be studied both with a high-power and a binocular microscope. The domatia are very conspicuous on the cuticular preparations since, where there is an extended piece of leaf tissue overarching the main bulk, the cuticular membrane overlaps and so is three layers in thickness.
There is in all probability no system of classification that will accommodate all types and grades of domatium. The usefulness of some sort of classification is, however, obvious, and the one which I use to cover leaf domatia has proved relatively satisfactory. Two main types of domatium are recognized (Fig. 4):
Marsupiform (pocket-shaped) domatium (domatium marsupiforme) (Plate 2 A, B).
All grades, from an extremely highly developed “ pocket ”’ to a situation which is scarcely able to be called a domatium exist. They are found in the axils of the major lateral veins with the midrib, or more rarely of secondary veins with major lateral veins. Ina leaf with very conspicuously raised veins a cuticular preparation will always show folding of the epidermis of the veins over that of the areolae, and this effect is of course doubled at a vein branch, especially the branchings from the midrib. Leaves of Buchenavia reticulata Eichl., for instance, show this to a marked degree. In some leaves this folding is only evident in the major lateral vein axils, and here a V-shaped suggestion of a domatium is to be seen, as, for example, in Terminalia mollis Laws. Penzig & Chiabrera (1903) classed this species as domatium-bearing, but the case is debatable. In the genus Tervminalia, as probably in many others, a whole series of types from that of T. mollis to the most complex pocket-shaped domatium is to be seen. This gradation can be visualized as the gradual filling-in of the notch of the V to form a V-shaped triangular pocket where all traces of the ‘‘ arms” of the V have disappeared. These latter types bear no relation at all to the conspicuousness of the veins, often being found on leaves with an almost planar venation. The basal point of the triangle points towards the vein axil, and the mouth of the domatium, forming one side of the triangle, away from it. Thus the depth of the domatial cavity, which lies between the leaf surface and the triangular area of tissue, is parallel to the leaf surface. The mouth of the pocket- shaped domatium is therefore at its broadest and most distal point.
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
=a
I, primary-axillary lebetiform. 2, primary-axillary marsupiform. 3, 4, vertical sections of 1 and 2 respectively, at angle bisecting the vein axil. 5, primary-axillary hair-tuft. 6, primary-axillary pubescent-marsupiform.
Fic. 4. Types of domatia :
7, marginal lebetiform, in section (Lumnitzera). 8, baso-laminar revolute (Strephonema).
g, baso-laminar revolute (Dioscorea). 10, twinned supernumerary lebetiform (Terminalia
catappa).
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 33
Lebetiform (bowl-shaped) domatium (domatium lebetiforme) (Plate 2 Cc, D).
As with the preceding type all grades of bowl-shaped domatia are to be found, the simplest of which might be visualized as a shallow depression in the leaf surface. Such shallow depressions are found in Lumnitzera, on the leaf margin, and have been referred to as domatia by Backer (fide van Steenis, in litt.). In most cases, however, bowl-shaped domatia are found in the axils of veins. In Terminalia plagata Merr. well-developed domatia are present in the major lateral vein axils, but in the secondary vein axils reduced forms, the shallow depressions visualized above, are to be found. A series may again be traced where the opening of the hollow becomes smaller, and/or the hollow itself becomes deeper and wider, the genus Terminalia showing all types of intermediates. As noted by Penzig & Chiabrera (1903), in some cases (e.g. J. microcarpa Decne.) the pore of the domatium may be at the level of the rest of the leaf surface, the pit being entirely internal, whereas in other cases (e.g. T. catappa) the pit is only partially sunken into the leaf tissue, the pore being at the apex of a small dome. In the best-developed bowl-shaped domatia the pore is extremely small in proportion to the large pit beneath (e.g. T. microcarpa). In the present type the depth of the domatium is thus at right-angles to the leaf surface. The pore is usually in the centre of the domatium as viewed from above, but it may be placed elsewhere. In some abnormal forms of domatium in T. catappa there are two pores. The pore may be circular, elliptic or slit-like : in the last case it is most frequently found on elongated domatia, the elongation of both domatium and pore being parallel to the lateral vein.
The above two types of domatium are nearly always distinct, and it is suggested that they arose independently, from the overarching veins and the shallow hollow respectively. Intermediates, although rare, are apparently present, but they are thought more likely to represent the points of convergence than of divergence. As shown above, increasing stages in complexity of both types can be traced which are difficult to interrelate. In apparently intermediate types (e.g. Terminalia moluccana Lam.) the bowl-shaped domatium has a large pore which is distal to the vein axil, and the cuticular membrane is often not trebled at its distal edge. The trebled membrane thus forms a crescent rather than a circle, and moreover the depth of the domatium appears to be parallel to the leaf surface, although it may not be so. This type obviously closely resembles the pocket-shaped domatium. The latter may also approach the intermediate situation by the narrowing of the opening. The domatium of Buchenavia fanshawei Exell & Maguire, which is classed as the bowl- shaped type, may possibly be a modified pocket-shaped type as all other species of Buchenavia which possess domatia have the latter type.
Two other types of domatium may be recognized : the first often very well defined ; the second of a highly dubious nature.
Revolute (rolled-margin) domatium (domatium revolutum).
In this type the leaf margin is inrolled, or folded under like the flap of an envelope. Inrolled domatia are found in such plants as Hevea brasiliensis (Kunth) Muell. Arg.
24 CUTICULAR: STUDIES AS AN AID TO PLANT TAXONOMY
(Penzig & Chiabrera, 1903) and Sivephonema spp. (De Wildeman, 1923 ; confirmed by my own examination). The inrolling is present at the base of the lamina, just above the petiole on either side. The degree of inrolling varies considerably, and the status of the domatium is thus dubious. In the flap-like domatia a specialized portion of the lamina is folded under ; if this is folded back it forms a regularly shaped projection from the leaf margin. It was found in dicotyledons (e.g. Ilex sp.) by Lundstrém (1887) and later in Dioscorea by De Wildeman (1904). The latter author described the structure in D. smilacifolia De Wild., in a new species (D. acarophyta De Wild.), and subsequently, from 1912 to 1914, in ten other new taxa. Burkill (1939) showed that these twelve represented three species only : D. smilacifolia, D. minutiflora Engler and D. pynaertit De Wild., but he discovered domatia in a fourth closely related species, D. praehensilis Benth.
Hair-tuft domatium (domatium fasciculatum).
The vein axils of the otherwise sparsely pubescent or glabrous leaves of a large number of species of trees may be densely pubescent or pilose, and these tufts of hairs have been considered to be domatia by Lundstrém (1887) and others. If mites do take refuge in them this may be accepted. Amongst British trees Tilia, Alnus and Corylus are examples. In this type there is no modification of leaf tissue other than the development of the hairs. Although many trees have leaves which are at first densely pubescent and later scarcely so, and the last areas to lose the indumentum are the veins of the lower leaf surface, the hair-tuft domatia are often well defined, even if their association is dubious. They are frequently found in connexion with the slightest development of a pocket-shaped domatium.
Lundstrém (1887) recognized more or less the same four types of domatia as I do, although a much smaller range of forms was considered. Penzig & Chiabrera (1903) closely followed Lundstrém, but described six types. The “ hollow ’’-shaped (fossette) and “‘ pocket ’’-shaped (tasche or borsette) domatia were each divided into two categories according to the presence or absence of hairs. It is not, however, advisable to distinguish between glabrous and pubescent types because the difference is apparently of no consequence and every type of intermediate is to be found. Penzig & Chiabrera appear to separate the “ hollows ” from the “ pockets ”’ in that the former are sunken into the leaf tissue with their openings on the level of the leaf surface, whilst the latter (which may be either bowl-shaped or pocket-shaped in my system) are raised above the leaf surface. Intermediates are, however, as common as either extreme, and the classification is considered inferior to that used in the present study. It so happens that Lundstrém’s scheme does not commit itself to agreement with either Penzig & Chiabrera’s system or mine, since all his ‘hollows ’’ were in fact wholly sunken into the leaf, no raised hollows (as in Terminalia catappa) being noted.
The latinized terms have been given for the convenience of concise description. The position of the domatia can also be described by the following terms : marginal, for those at the leaf margin (e.g. Lumnitzera) ; primary-axillary, for those in the
CULICULAK STUDIES-AS AN, AID TO PLANT TAXONO My, 35
major lateral vein axils (the most common type) ; secondary-axillary, for those in the secondary or lesser vein axils (e.g. Tilia spp., Terminalia oreadum Diels and T. plagata) ; baso-laminar, for those on either side of the lamina base (e.g. Quercus vobur L., Strebhonema spp. and Dioscorea spp.) ; extra-axillary, for those just below each major lateral vein axil (e.g. Rudgea lanceolata Benth.) ; and supernumerary, for those not associated with any particular part of the lamina (e.g. Terminalia catappa, where they are found scattered up the side of the midrib apart from the vein axils ; and T. grandiflora Benth. and Conocarpus spp., where they are found very sparsely scattered over the lower leaf surface). They are occasionally also found elsewhere : for example in Schinus terebinthifolia Raddi domatia revoluta are found on leafy wings to the rhachis of the pinnate leaves ; and in one or two genera they have been reported from young twigs and petioles.
In most cases the epidermis lining and around the domatia is little or not modified. In the axils of major lateral veins there is usually a patch of small straight-walled isodiametric cells, and when a primary-axillary domatium is present this area is merely enlarged, and this type of epidermis lines the domatium. In some cases, however, the cells do differ in some way from those surrounding the domatium, usually being smaller. In Terminalia catappa the adjacent cells are often arranged in radial files for a short distance. In Conocarpus erectus L. var. erectus the epidermis is absolutely identical in, near and away from the domatia, even to the extent of the possession of stomata. In C. erectus var. sericeus Forsstr. ex DC. the epidermis is very densely pubescent, but that lining the domatia is more or less glabrous, as in var. erectus.
In many cases the domatia are associated with an abnormal amount of hair development in, on or around the actual structure. In some species, e.g. Thiloa gracilis (Schott) Eichl., the hairs are of a peculiar type not found elsewhere on the leaf. Ina few species gland-like structures are to be found in the domatium. These were noted in Anacardium by Lundstrém (1887), and in the Combretaceae they have been found in Buchenavia parvifolia Ducke and Terminalia archboldiana Exell, though not very frequently. The nature and function of their secretion, if any, are unknown. It is just possible that the deeply sunken glands found on the leaves of Laguncularia are homologous with domatia, but this is not likely as they are found on both epidermides and have an extremely narrow slit-like opening at the surface.
The examples cited above clearly show that domatia may have a considerable value in taxonomy, although they have rarely been used. An extreme case is per- haps that of Buchenavia fanshawei which has well-developed domatia lebetiformia. Of the other 22 species examined in this genus 13 have domatia marsupiformia and nine lack domatia altogether. Thus much use of this fact was made in constructing a key to the genus (Exell & Stace, 1963). In the Combretaceae domatia revoluta are found in all six species of Stvephonema, which is usually treated as a separate subfamily from the other genera, all of which lack these domatia. The remaining genera belong to three groups, of which two contain eight genera each. In one of these groups (Terminalieae) four genera possess domatia (three lebetiform and three
36 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
marsupiform); whilst in the other (Combreteae) all eight genera possess marsupi- form domatia only. The third group (Laguncularieae) consists of three genera, of which one possesses peculiar marginal lebetiform domatia. Thus in the Combretaceae the occurrence of the various types of domatia to some extent agrees with the generally accepted subdivision of the family. This is not the usual situa- tion, however. Penzig & Chiabrera (1903) recorded all six of their types from the Rubiaceae, and four from several genera, and in general domatia are useful in identification at the specific level only.
There is no reason not to believe that all specimens of a domatium-bearing species do possess domatia, although these may be sparser in some cases and may be lacking altogether on some parts of the plant such as the sucker shoots. Bloembergen (fide van Steenis, 7m litt.) found that Alangium chinense (Lour.) Harms and A. kurzti Craib possess domatia in most of Asia, but not in Malaysia, but, as this situation has never been reported elsewhere, the absence of domatia in Malaysia or the identity of the species in the two areas needs careful checking. In the genus Terminalia, of which I investigated 160 species, no such geographical variation was noted. For example T. catappa, a native of Asia and Australasia, possesses very typical domatia whether in its native countries or planted in Africa or America ; and T. arjuna (Roxb.) Wight & Arn., a native of India, lacks them there or when planted elsewhere in Asia, Africa or America. <A different type of geographical variation, however, was discovered in Tervminalia. Of the 160 species studied 50 (31°%) possess domatia, of which 16 (32°) have domatia marsupiformia and 34 (689%) domatia lebetiformia. However, in Australasia 21 (62%) of the 34 examined species possess domatia, and the figures for other continents are : Malaysia 19 (58%) out of 33 ; Asia 4 (13%) out of 34 ; Africa r (3%) out of 39 ; and America 7 (23%) out of 30. Thus as one travels east or west from the islands between Australia and Malaysia the percentage of domatium-bearing species drops sharply. The proportion of the two types of domatia does not significantly differ from continent to continent.
Extremely little is known about the formation or function of acarodomatia, but despite suggestions to the contrary it is obvious that mites play no part in their formation. It is more likely that the mites inhabiting the domatia (if this is a regular occurrence) are in a purely chance association. It is possible that they may enlarge the domatia somewhat once they have taken possession of them, and this may be the cause of the deep domatia in Terminalia bursarina which sometimes reach the upper epidermis. There is no necessity, however, to consider that this is not anatural plant process. Lundstrém (1887) claimed that the domatia of Psychotria, Coprosma and Rhamnus developed in the absence of mites, but, although he was doubtless correct, his experiments have never been repeated under the exacting conditions necessary to exclude all mites. Mani (1964) also stated that the mites did not participate in domatium formation. Bailey (1924) showed that American myrmeco- domatia developed in the absence of ants. Whilst it is true that seedlings (and often suckers) lack domatia, these developing only at a later stage, seedling and sucker leaves often differ in many ways from the other leaves, and van Steenis (1953) states that in Nothofagus domatia develop by the young sapling stage.
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 37
Good evidence for the non-participation of mites in domatium formation is that domatia are definitely formed in predisposed places: e.g. at the bases of the laminae in Dioscorea, or just below each major lateral vein axil in Rudgea. If the domatia were formed by mites they would not be so distributed, and neither would they be so constant in their presence (on all individuals in all geographical areas) and structure. In the case of primary-axillary domatia these are not always found in all major lateral vein axils. They are always better-developed towards the lamina base, but in some species, e.g. Terminalia bursarina, T. grandiflora and Nothofagus menziesit Oerst., they are found only in the lowest axil on each side of the midrib, or in the latter two sometimes also in the next axil. Furthermore, only certain species in any one genus may possess domatia, e.g. only two species out of about 40 in Nothofagus (van Steenis, 1953) and four out of several hundred in Dioscorea (Burkill, 1939). The domatia are often very hairy, and since many mite-produced galls also possess this character, definitely as a direct result of the animals’ activities, it has been suggested that pubescent domatia are also formed in this way. There is no reason to believe this, however, and in most Combretaceae, at least, the pubescent domatia (often the only pubescent parts of the leaf) are relics of a wholly pubescent juvenile state, and the hairs may wear off even here at senescence.
The very young leaf develops domatia marsupiformia by an extension of the pocket tissue over the vein axil, but, if domatia lebetiformia evolved in the manner previously suggested, ontogeny does not repeat phylogeny as the pits do not appear to develop by an extension of tissue encroaching over a hollow. In the cases examined (especially Conocarpus erectus) the pit forms internally, and at first is only marked by a primary axillary pimple. Later a pore develops in the top of the pimple, and this enlarges to a varying degree. The hollow appears to develop internally. Different stages may be seen on one leaf, development being acropetal. This is an important argument against any participation by mites in domatium formation. Holtermann (1893), however, seems to suggest that the pits of Conocarpus do develop by means of an epidermal intucking, and not as I have observed.
The situation regarding domatia has been discussed at some length, although numerous other points could have been included, since little is known of them in general, and they have been largely neglected by taxonomists. With regard to their position and structure they may be very useful in identification, but their biological significance is highly obscure. They are doubtless often used as a shelter by mites and other similar minute arthropods, but in the great majority of cases no animals have been connected with them. This is in marked contrast to the case regarding myrmecodomatia. The prefix “‘ acaro-’”’ is thus best rejected and replaced by one suggestive of the organ on which the domatia are found, e.g. phyllodomatia.
B. MICROSCOPIC CHARACTERS. (1) Epidermal cells.
There are a large number of directions of variation to be found with respect to the epidermal cells, even omitting the several characters which are not apparent from a purely surface examination, but they have rarely been used by taxonomists.
38 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
In certain regions the epidermal cells are modified, notably over the veins and leaf margin and around the stomata and trichome-bases. This section deals only with the areas not so modified.
Although the shape, size and other characters of the epidermal cells exhibit a wide range of genotypic variation, which in many cases has definite taxonomic applica- tion, the cells certainly show an extreme degree of phenotypic variation which is elsewhere only paralleled by characters such as pubescence and petal colour, etc.
One of the most obvious features of the epidermal cells is their shape ; this is often quite different on the two epidermides. This fact is often of systematic importance, although Odell (1932) decided that it indicated that cell shape was “of no diagnostic value’’. The cells are usually sodzametric, i.e. more or less the same in length and width, with any scattered elongated cells randomly orientated. In some dicotyledon leaves, however, as well as in many groups of monocotyledons, the epidermal cells are predominantly elongated. This feature usually occurs in narrow leaves, the cells lying parallel to, or less often at right-angles to, the leaf axis. In cases of isodiametric-celled epidermides some species have a characteristically low (e.g. Gutera senegalensis J. F. Gmel., Plate 4 D) or high number of walls to each cell, this factor markedly altering the appearance of the epidermis.
In some species the epidermal cells are arranged in small groups separated by thicker than normal cell walls, where it appears that an originally single cell has retained its identity after many subsequent divisions. Species of Avicennia show this feature well. A related phenomenon is the anticlinal division of the epidermal cells, where cells are divided by one or more thin straight walls which cut a normal- sized cell into two or more parts. Such secondary division (i.e. occurring after normal epidermal cell division has ceased, and not being accompanied by an increase in size) is often taxonomically important. In the genus Laguncularia, for example, it is absolutely constant, and it is not well developed elsewhere in the family Combretaceae.
The term “ wall” is here used as an abbreviation for the apparent cell wall produced by the cuticular flanges, and it should always be borne in mind that the actual cell wall was present for some distance on either side of these. For this reason some workers prefer the term “ cell outline ’’.
The anticlinal walls of the epidermal cells are either straight, curved or variously undulate or sinuate (cf. Plate 3 A, B). This is undoubtedly of taxonomic use in some instances : it has been used as a major character in Lemna and is useful in many groups of Combretum. For some unknown reason straight-walled epidermal cells are commoner in xeromorphic plants than in mesomorphic ones, which typically have undulate cell walls. The relative advantages and disadvantages of undulate walls are unknown, but Wylie (1943), who attributes a conductive function to the epidermis, suggests that their advantage is to increase the area of contact between adjacent cells. If this is so it is surprising that undulate walls are not commoner in xerophytes. The type of undulation is very various ; the “ waves’’ may be
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 39
most easily measured in exactly the same terms as light waves, i.e. in the three variables frequency, amplitude and wave-length. In the present study the frequency was usually expressed in terms of wave-lengths per wall, but in some cases it seems that the number of peaks (i.e. half wave-lengths) per cell may be a less variable value. There may be considerable or very little variation in the degree of undulation on one leaf, but the undulation often markedly decreases towards the leaf margin and veins. It will be seen from Fig. 5 that the three variables mentioned above aptly express all the characters of the undulations but one, i.e. the shape of the waves. These may usually be described as U-shaped, V-shaped or Q-shaped. In some species peculiar shapes of undulation are found with ornamentations, in the form of knobs, ridges or T-shaped thickenings, on the outer side of the crest of each wave. Curved epidermal cell walls may be described as having half a wave-length.
Frequency Wave-length Amplitude
ey ae
oa. . i aye i se SS 0:5 2x y
Fic. 5. Types of cell-wall undulation.
The exact causes of epidermal cell-wall undulation are unknown, the two main _ theories involving the suggestions that it is due to the tensions set up between the mesophyll and the epidermis, and that it is caused by the method of hardening of the differentiating cuticular membrane (Watson, 1942). It has been found that whereas many leaves have the epidermal cells undulate-walled throughout
40 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
the height of the cells, some have only the outer part of the wall undulate, and thus the second hypothesis is by far the more likely. Sifton (1963) has provided further evidence supporting Watson’s theory. Amongst species examined in the present study the situation where only the outer part of the wall is undulate is very con- spicuous in Ramatuella. Watson, however, was able to produce either condition in mature ivy leaves by varying the illumination during development, so that the character is probably rarely of taxonomic use. In some species with straight walls the isolated cuticular membranes show the tips of the cuticular flanges to be irregu- larly undulate, but this is probably due to the effect of removing the epidermal cells. It is especially noticeable in Conocarpus erectus.
As mentioned above, the cell shape is often different on the two epidermides, the chief varying factor being the degree of undulation. In almost all undulate-walled leaves the amount of undulation is greater on the lower than on the upper epidermis. In many cases the former is markedly undulate-walled and the latter straight-walled. Odell’s (1932) remark that a few leaves of Acer pseudoplatanus L. may show the reverse of this situation, which the bulk of the leaves exhibit, needs to be checked. Some species, however, do possess a markedly more undulate-walled upper epidermis than lower, e.g. Terminalia microcarpa, T. plagata and others.
Epidermal cells vary tremendously in size, average widths being about 20-30 un. Even on typically large-celled epidermides, however, small cells are usually to be found scattered. Some epidermides, in fact, are characterized by their extremely variable-sized cells. That cell size might be of taxonomic value is indicated by the knowledge that polyploids usually have larger cells than diploids, although this is better shown by the stomata than other epidermal cells. Solereder (1908) mentions the fact that certain species are characterized by a particularly small- or large-celled epidermis, and this has been noted in the present study. Avicennia marina var. vumphiana (Hallier f.) Bakh., for example, has a notably smaller-celled epidermis than other varieties of this species. Lumnitzera littorea (Jack) J. O. Voigt has a smaller average cell size than has L. racemosa Willd., although some overlap occurs and this character, like all others, fails to separate the two species on surface cuticular characters in all cases. The cell size is frequently different on the two epidermides, as is cell shape. Solereder (1908) states that in general the upper epidermis is composed of larger cells than the lower, but the opposite was found in the present study. Characters such as the comparative sizes of parts of the same leaf are extremely useful as the different parts are all equally affected by external conditions, e.g. the presence of upper epidermal cells twice as large as the lower epidermal cells is a better diagnostic character than the presence of upper epidermal cells of an average length of 40 uw. The cell size usually decreases towards the leaf margin and apex.
Although the boundary between the cuticular membrane and the cellulose cell wall is usually a smooth line, interrupted only by the cuticular flanges, in some species regular cellulose pegs protrude outwards into the cuticular membrane, which becomes very thin in these regions. This may or may not be accompanied by an extension outwards of the cell lumen in the corresponding position. In either case small
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 41
elliptic or circular areas are seen on the cuticular membranes where the thin membrane is less deeply stained. Such areas are very well developed in Combretum obanense (Baker f.) Hutch. & Dalziel and certain of its relatives, but in all other species of this genus they are unknown. The thin areas are frequently present in the bulge of each undulation, and in C. obanense they are so conspicuous that the normal differentiation between cuticular flanges and non-flange areas is entirely masked, except on the veins. The epidermis in this case appears to be composed of minute cells about 5-10 pw across, with extremely thick walls. Other situations have been described where the outer epidermal cells contain large pores, the cell contents having been lost. The origin of the pores in the cuticular preparations of Cinnamomum obtained by Marlier-Spirlet (1945) is obscure.
The surface of each epidermal cell may be flat or convex, or bear a small conical process. The latter is known as a papilla, and the epidermides or cells which bear papillae are said to be papillate, but unless the papillae exceed a certain length they are not seen on surface preparations of cuticular membranes. They may, however, be viewed on folded-over pieces of membrane. All gradations from simple conical papillae to long structures better described as hairs exist, and if they are long enough to become folded over on cuticular preparations they are visible in surface views. They are often considered to be taxonomically important. The papillae are usually centrally placed on the epidermal cell, but they may be found on one corner of each cell, as in Combretum zenkeri Engler & Diels. Elsewhere in this large genus papillae have been recorded in less than half a dozen species, e.g. C. lanceolatum Pohl ex Eichl., and in all cases are of the central type.
(2) Leaf venation.
The epidermal cells above and below the veins are usually variously modified, so that leaf venation is a cuticular character. Characters of the venation which are microscopic alone, as opposed to the type of venation (which is a macroscopic character), are the extent and nature of the modification of the epidermal cells.
The midrib and the lateral, secondary, etc. veins are usually enclosed or capped by sclerenchyma, which may or may not reach either epidermis. In any case the epidermis above and below these veins is usually markedly modified, in the majority of cases the cells being elongated parallel to the veins. These elongated cells are usually rectangular, and, even if the epidermal cells of the areolae have undulate walls, straight-walled. In general the more xeromorphic a leaf the less conspicuous are the veins on the epidermis. Usually the degree of modification of the epidermis is proportional to the prominence of the veins, hence there tends to be a much greater modification on the lower than on the upper epidermis, although this is not always so. In most cases a more modified epidermis has more narrowed and elongated cells over the veins. The ends of the cells may be truncate, oblique or almost pointed and interlocking (prosenchymatous) ; and the cells arranged end to end in numerous parallel rows, or less regularly placed. In many cases, especially when the midrib is well developed on the epidermis but the cells are not very narrow, the cells are secondarily subdivided at right-angles to the midrib, and in these cases
42 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
the cells may be squarish or even broader than long. Some leaves, most frequently rather xeromorphic leaves with a prominent midrib but rather inconspicuous veins and venules, have the epidermis modified over the former into strict rows of broader- than-long cells, the primary and secondary cell divisions being indistinguishable because all the cuticular flanges are equally thick. Examples found in the Combre- taceae are Strebhonema spp., Combretum demeuset De Wild., Terminalia laxiflora Engler and many others. This effect is almost always more greatly developed on the upper epidermis, and furthermore the cells of the veins of lesser prominence show it to a progressively lesser degree.
The lesser veins and venules showing no directional tendencies have a different anatomical structure. Above and below, or below only, many of them are bounded by a dorsiventral extension, the vein-extension or vein-rib, which consists of living cells elongated parallel to the venules and abutting on to the epidermis below and often also above. Wylie (1943) considers that these, as well as the epidermis, constitute a conductive system supplementary to the venous system. When a venule is connected to the epidermis in this way the latter is modified. Thus it may be seen that, as with the major veins and the midrib, the upper epidermis is modified to an equal or lesser extent than the lower epidermis over the lesser veins and venules. The modification of the cells over the venules (Plate 5 B) is similar to but much less developed than that over the midrib, etc. The cells are generally elongated, to varying extents, although secondary division of them is rare or absent. When the epidermis is mostly composed of undulate-walled cells the venule cells are usually straight-walled, but sometimes they are slightly, or even equally, undulate-walled. In the latter case only their elongation and, on the lower epider- mis, the lack of stomata make them distinguishable.
Thus every intermediate between the very long and narrow midrib cells to the scarcely modified venule cells exists. The cells in the centre of each vein are mostly greatly modified, the edges of the veins gradually or abruptly merging into the normal epidermal cells. The most modified midribs are several hundreds of cells wide, the smallest venules only one or two. In an average mesomorphic leaf such as those of many species of Combretaceae the following degree of modification is seen : midrib composed of about 50 parallel rows of narrow elongated straight-walled cells ; major lateral veins about half as wide, the cells shorter and wider ; secondary veins about ro-15 cells wide, the cells only about twice as long as broad ; lesser veins about 5-10 cells wide ; venules forming venule reticulum about 2-3 cells wide, the cells little longer than broad, many with slightly undulate walls. Variation in one extreme is shown by strongly xeromorphic leaves, e.g. of Laguncularia, where only the midrib is modified on the epidermis, and that not conspicuously ; and in the other extreme by species with strongly reticulate leaves, e.g. Buchenavia reticulata, where the venules forming the reticulum are 5 or more cells wide, the cells being straight-walled and somewhat narrow and elongated. In all cases the lower epidermis is more greatly modified than the upper, so that the venation of the upper epidermis of a leaf would resemble that of the lower epidermis of a more xeromorphic leaf, and so on.
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 43
From the preceding paragraphs it can be seen that the three variables mentioned (i.e. the venation pattern ; the degree to which the venation system is recognizable on the epidermis ; and the type of modification of the cells) are completely indepen- dent of each other. Because of this, caution must be exercised in interpreting the extent of the venation system seen on the epidermis as that to be found in the leaf : it is frequently not so. The smallest veins represented on an epidermis, whether they are in fact the smallest venules of the leaf or of a much larger order, tend to be of the 2—3-wide slightly elongated-cell type. Thus a venule reticulum with areolae 3000u across seen on an epidermis might represent this structure in the leaf, or smaller venules may be present but not differentiated on the epidermis. According to the excellent figures of Wylie (1943), and to my own observation, the ultimate veinlet terminations rarely have vein-extensions, and are thus rarely visible on an epidermis. Apparent veinlet terminations seen on the epidermis are due to venules which are not represented in full.
Although the vein-like structures seen on the epidermis are of course only modified epidermal cells, they are for convenience referred to as midribs, veins and venules, etc. Thus the “ upper epidermal midrib ” or “ lower epidermal venules ’”’ may be referred to in descriptions. The unmodified epidermal islands within the venule reticulum may likewise be referred to as the areolae. In cases where the venules are not well represented on the epidermis, and the venation system is thus apparently open rather than closed, the unmodified epidermal cells may be referred to as the non-venous areas. In general the differentiation of the veins on the epidermis is subject to rather more phenotypic variation than is the organization of the actual veins themselves, calling for a greater measure of caution in their taxonomic use.
(3) Leaf margin.
The epidermis is continuous over the leaf margin, connecting the upper and lower epidermides, but its cells are modified at these positions, providing useful taxonomic characters. Towards the margin the cells of both epidermides are frequently smaller and thicker-walled, and if the normal cells are undulate-walled the undulations become less pronounced. Thus the marginal cells either abruptly or gradually merge into the normal epidermal cells.
The width of the margin, of course, is partly dependent upon the thickness of the leaf, and may be of diagnostic value. In most mesomorphic leaves the margin is not very wide, often not exceeding 5 or 6 cells across. The cells are in most cases rectangular, straight-walled and variously elongated (Plate 3 c). Secondary division of these cells may be apparent, giving rise to squarish or broader-than-long cells. The margin is thus frequently very similar to a venule, and may be indistinguishable but for its continuous straight appearance and the absence of other venules branching from it.
In certain cases, especially in xeromorphic leaves, other types of leaf margin are to be found. One of the most common of these is composed of small isodiametric cells, frequently with very thick walls and more or less circular lumina. Sometimes
44 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
these are arranged into parallel rows as is normally the case, but in other leaves they may be quite irregularly arranged. All intermediates are to be found, but in some cases the margin provides valuable systematic characters. The two Com- bretaceous mangrove genera, Lummnitzera and Laguncularia, for example, are closely similar in cuticular characters, but one of the three main differences lies in the leaf margin. In Laguncularia (Plate 3 D) the cells are much smaller than those elsewhere, have thick walls and circular lumina and are arranged completely at random. In Lumnitzera, however, the marginal cells are scarcely smaller than the other epidermal cells, have little-thicker walls, have angular lumina, are rectangular and slightly to conspicuously longer than broad, and are always arranged end to end in parallel rows.
T-piece at
stomatal aperture stomatal pole
walls of guard-cells
outer stomatal ledge subsidiary
radial walls of cells
tangential
Fic. 6. Terminology of stoma as seen in surface view.
(4) Stomata.
The term stoma is here taken to include the pair of guard-cells with the stomatal aperture or pore between them. In all vascular plants there are normally two guard- cells to each stoma, these arising from a single stomatal mother-cell at an early stage of leaf development, although abnormalities with one or three or more guard-cells do rarely occur (Dehnel, 1961; Ahmad, 1964a; Pant, 1965) and may even be of diagnostic value. Other cases have been described of bistratose guard-cells which have divided by a periclinal wall (Rehfous, 1914). The guard-cells are surrounded by normal or modified epidermal cells, which in the latter case are termed subsidiary cells or accessory cells. The whole system is known as the stomatal complex or stomatal apparatus. In surface view the guard-cells, in the great majority of vascular plants, and in almost all dicotyledons, are vaguely reniform. The anticlinal wall on the side towards the pore is here described as the poral wall, and that towards the rest of the epidermis the epidermal wall. If subsidiary cells are present their anticlinal walls abutting on the guard-cells may be known as radial walls, and those adjacent to the rest of the epidermis as tangential walls (Fig. 6).
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 45
In vertical transverse section the guard-cells show an extremely various construc- tion which is surprising in view of their relative constancy in surface view (Fig. 7). The guard-cells vary in three main directions: their position relative to the subsidiary or other adjacent cells ; the relative thickenings of the various walls of the guard-cells ; and the pattern of cuticular ornamentation of the guard-cells, All these are important taxonomic characters. In most mesomorphic plants the guard-cells are on a level with or very slightly sunken below or raised above the rest of the epidermis, but in some xeromorphic plants they are often sunken to a considerable degree. In many conifers, as well as in a number of angiosperms, the guard-cells are very deeply sunken and completely overarched by the subsidiary
back cavity
sub-stomatal cavity
inner stomatal ledge
stomatal flap NS ‘
inner walls of guard-cell
epidermal
poral wall \
4 1D by
subsidiary cell
outer wall of guard-cell
outer stomatal ledge front cavity
stomatal aperture Fic. 7. Terminology of stoma as seen in transverse vertical section.
cells, so that in a surface view the former are almost invisible, their position being marked by a ring of subsidiary cells (known as a poral ring) around a nearly circular pore. In other plants the subsidiary cells are also sunken. The thickenings of the guard-cell walls are very uneven in distribution, being connected with the physiology of the stomatal movements. In general, however, the poral and epidermal walls are rarely thickened, the former never, whilst the other two walls (i.e. the outer wall, adjacent to the environment, and the inner wall, adjacent to the inner regions of the leaf) are variously thickened. In some species, mostly xeromorphic, the thickenings are very considerable, and the cell lumen may be reduced to an extremely
46 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
small proportion of the cell volume. The amount of cuticular ornamentation of the guard-cells is frequently proportional to the thickening of the cell walls, and hence to the degree of xeromorphism. The cuticular membrane which covers the epidermis is continuous over the subsidiary and guard-cells, and either ends where the inner and epidermal walls meet or also covers a varying amount of the subepidermal cells which line the space which usually exists below the stoma (the sub-stomatal cavity). In most cases the cuticular membrane is very thin on the poral walls, but thick on the outer and inner walls, especially where each of these meets the poral wall. Here the cuticular membrane may be protracted into a pair of cuticular or stomatal ledges, ridges, horns, rims or hooks, which may be termed outer and inner respectively. This is found in many xeromorphic plants, but more frequently only the outer stomatal ledge is present. When the stoma is closed the poral walls of the two guard-cells meet, and above and below this point the outer and inner stomatal ledges also, if present, meet, or nearly so. Thus two extensions of the stomatal pore are delimited : a front cavity and a back cavity (Fig.7). Insome plants, e.g. Bruguiera spp. (Areschoug, 1902) and Ceriops spp. (Stace, 1963), there are actually two pairs of outer stomatal ledges subdividing the front cavity into two portions.
In cuticular preparations the stomata are usually well preserved, and they provide valuable systematic characters (cf. Plate 3 A, B). Because much stomatal variation is in the third dimension (depth), however, a number of characters are not usually observable on cuticular preparations. It is often very difficult to ascertain the exact relative positions of the guard-cells and their neighbouring cells and the precise structure beneath the outer stomatal ledge. The cuticular membrane below the latter, since it curves back over the inner stomatal wall, is conspicuous on many preparations as a pair of stomatal flaps. A very good idea of the degree of cuticulariz- ation and sunkenness of the guard-cells and subsidiary cells may often be obtained by examining cuticular preparations with the membranes folded over on themselves inside outermost (Plate 5 c). Also a surprising amount of structure can be observed once the many lines and ridges on the preparation have been interpreted by means of vertical sections. This approach was useful in the Rhizophoraceous mangroves, which have many diagnostic details of stomatal structure. The single known epidermal difference between Lumnitzera littorea and L. racemosa, the shape of the guard-cells and subsidiary cells in section, however, was not visible from surface views of either cuticular preparations or whole epidermides.
However, in surface view the most obvious cuticular characters are present on the outer walls of the guard-cells. The shape of the guard-cells as a whole is a useful criterion which is usually constant within species. The commonest shapes are circular, elliptic and oblong, but some stomata may even be broader than long, and sometimes angular. The stomatal poles where the two guard-cells meet may be obtuse, truncate, rounded or retuse, the last apparently being the case in many xeromorphic plants. The appearance of various thickenings on the outer guard-cell walls is also of importance. The outer stomatal ledge is usually visible from surface views, and its thickness and relative distance between the epidermal and poral walls
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 47
are characters of significance. When this ledge is directly over the poral wall of the guard-cells it forms a thick rim to the stomatal aperture which Bandulska (1923, etc.) called the poralrim. At the stomatal poles there may be developed a T-shaped thickening or 7-pzece, much used by Bandulska (1924) as a character in the Fagaceae. Sometimes only the upright or the cross-piece of this is present. In some groups there is a small thin area at the polar end of each guard-cell, being circular, elliptic or comma-shaped. Solereder (1908) mentioned several examples and Rao (1939) found them commonly in the Magnoliales. A few genera are known to possess peculiar lobe-like extensions of the guard-cell epidermal walls, and these were noted in Strepbhonema.
Perhaps of more significance than any of the above characters, however, are the presence and types of subsidiary cells around the stomata. In his various studies Florin (1931, 1933) distinguished two types of stomatal development in the gymno- sperms, based on whether or not the guard-cells and subsidiary cells originate from the same mother-cell (syndetocheile and haplocheile respectively), and these types can usually be seen in the mature state because the former has two adjacent sub- sidiary cells and the latter a variable greater number. Although it was frequently necessary for Florin to draw conclusions concerning ontogeny by examining the mature anatomy, since many of his plants were fossils, it is unfortunate that it was so because the two terms are now more often used for describing the mature anatomy than the developmental sequence, and it has been shown by many workers that the presence of subsidiary cells does not necessarily signify a syndetocheile ontogeny, nor does their absence always indicate a haplocheile one. Thus the terms syndeto- cheile and haplocheile are better abandoned, and the cells neighbouring a stoma (whether they be subsidiary cells, i.e. different from the other epidermal cells, or not) described as mesogenous or perigenous according to whether or not they arise from the same mother cell as the stoma. These terms were first used by Florin (1933). In addition Pant (1965) proposed the term mesoperigenous for those situations where one neighbouring cell is mesogenous and the rest perigenous. This terminology has the advantages that it is equally applicable to all groups of plants rather than primarily to the gymnosperms, and that it allows for the fact that mesogenous subsidiary cells may become indistinguishable from other epidermal cells, or that perigenous cells may become specialized as subsidiary cells. The last term has no
ontogenetic implications.
The earliest system of subsidiary cell classification was adopted by Solereder (1908) from the work of Prantl (1872) and Vesque (1889). Six ontogenetic categories were recognized, three of which were perigenous and three mesogenous. Due to convergence during maturation only four types were recognizable in the mature state, being termed Ranunculaceous, Rubiaceous, Caryophyllaceous and Cruciferous. Metcalfe & Chalk (1950) completely ignored the modes of development and renamed these four categories as follows: anomocytic, with no distinctive subsidiary cells (Plate 4 A); paracytic, with 2 subsidiary cells lying parallel to the guard-cells (Plate 4 c) ; dzacytic, with 2 subsidiary cells lying transversely to the guard-cells; and anisocytic, with 3 subsidiary cells of which one is distinctly larger or smaller than the other two.
48 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
These groups are often of considerable taxonomic significance, in many cases one type being characteristic of a whole genus or family. In the Combretaceae, for instance, Stvephonema is the only genus with paracytic stomata, and this agrees with the usual treatment of Stvepbhonema as a separate subfamily. Apart from the four types of situation mentioned above there are other arrangements of subsidiary cells for which no terms are generally available, although for descriptive purposes they are obviously desirable. The term actinocytic was proposed by Metcalfe & Chalk (1950) for cases where the subsidiary cells, usually four or more in number, are elongated radially to each stoma, and cyclocytic by Stace (1963), where a similar number of cells forms a narrow ring round each stoma. Both terms have subsequently been used by other workers, although they probably refer to mere modifications of the anomocytic type. Of the 19 genera of Combretaceae without paracytic subsidiary cells, in fact, two (Lumnitzera and Laguncularia) are cyclocytic (Plate 4 B) and the rest anomocytic. Bandulska (1931) also found cyclocytic subsidiary cells (“ girdle cells ’’) in various genera of Myrtaceae. More usually, however, differentiation by subsidiary cell types is only at the specific level, or is even here absent. In Anopyxis klaineana (Pierre) Engler (A. ealaensis (De Wild.) Sprague), for example, paracytic, anisocytic and anomocytic stomata are said to occur on the same leaf (Metcalfe & Chalk, 1950), and other members of the Rhizophoraceae also show varying arrangements. It seems probable that only one basic mode of development is involved, but that the degree of subdivision of the subsidiary cells finally differs, producing the different arrangements.
A reliable ontogenetic classification is highly desirable, and that proposed by Pant (1965) seems to be satisfactory. Ten categories are recognized, of which one is perigenous, three mesoperigenous, and six mesogenous. This nomenclature is quite separate from one based upon mature topography. The latter scheme would involve a larger number of categories (perhaps about 15), several of which could be derived from several different modes of development. Apart from the six terms mentioned previously one other has so far been named: f¢etracytic (Metcalfe, 1963), for cases where there are four subsidiary cells, two polar and two lateral, as in many monocotyledons. Terms such as paracytic, tetracytic, etc., could conveniently be prefixed by meso-, peri-, etc., once the mode of development was known. Members of the Rubiaceae are meso-paracytic (Pant, 1965), for instance, and of the Gramineae peri-paracytic (Stebbins & Jain, 1960; Stebbins & Shah, 1960; Stebbins & Khush, 1961). All investigated monocotyledons, in fact, have been found to be perigenous. It seems clear that the developmental studies at present being carried out mainly by Pant and his compatriots (e.g. Pant, 1965; Pant & Mehra, 1963; Pant & Verma, 1963) will provide valuable evidence for a classification of stomata which will prove to be of considerable taxonomic and phylogenetic value.
In some plants the subsidiary cells are furnished with distinct papillae, which may be absent elsewhere on the leaf. These may be quite long and overarch the stomatal aperture (Solereder, 1908).
Stomata vary considerably in size and this is often a character of some importance. The relation between the level of ploidy and stomatal size is well known, but stomatal
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 49
size does not always indicate the former (Carolin, 1954). In many species the stoma- tal size also varies on a single leaf, but this may not prevent it from being an important taxonomic feature. On many leaves abnormally large stomata are to be found, and these are often, if not almost always, water-stomata, which are supposed to secrete drops of liquid water. Their presence is noteworthy. Water-stomata, or water-pores if the surrounding cells are not obviously guard-cells, may also be smaller than the normal stomata, and sometimes occur in groups (Solereder, 1908).
The distribution and frequency of stomata are their most conspicuous characters, and are often of considerable systematic value. They are very often, of course, connected with the ecology of the species, and thus not likely to be of great phylo- genetic importance. In many hydrophytes, for example, stomata are either absent or infrequent and vestigial. In some terrestrial plants also some stomata may be vestigial, and Solereder (1908) describes situations where they may be plugged with a resinous mass. In plants with floating leaves they are usually on the upper epidermis only, and the same is true of other species with variously modified leaves which for some reason have a more concealed lower than upper epidermis, or even with apparently normal leaves (Solereder, 1908). In most dicotyledons, however, the stomata are more or less confined to the lower epidermis, although it appears to be usual to find a few scattered along the vicinity of the upper epidermal midrib of mesomorphic leaves. Solereder also noted this, and wisely pointed out that it could have ‘“‘ no great systematic value’. They are present there in most species of Combretaceae, and are frequently abnormally elongated. They are sometimes present upon the midrib itself, rather than beside it, and cannot here be functional. All intermediate situations from the complete absence of stomata on the upper epidermis to their presence as abundantly as on the lower epidermis are to be found, even within the Combretaceae alone. Lwmmnitzera spp., in fact, have stomata more abundant on the upper than on the lower epidermis. Species showing a slight increase from normal in the abundance on the upper epidermis have stomata scattered quite frequently alongside the midrib, more sparsely alongside the veins, and very sparsely in the rest of the areolae, and the further from the midrib they are situated the less abnormal is their anatomy. On the lower epidermis stomata are usually confined to the areolae, or non-venous areas, but sometimes they are to be found scattered on the midrib. They have seldom been recorded from the leaf margin of a dorsiventral leaf, or from the venules and veins of the lower epidermis.
Thus stomata are typically found in the areolae of the lower epidermis, and usually only very sparsely anywhere else. In leaves showing various xeromorphic characters, especially where a closed system of venules is not apparent on the epidermides, they may be otherwise distributed. In the latter case they are frequently found all over the epidermis, except for the veins and midrib, but in some species they may be present only in special areas, e.g. longitudinal grooves, or depressions. These latter situations are not found in the Combretaceae, although here there are some species with very prominent reticulations and thus sunken areolae. The position and types of sunken areas, the stomatal crypts or stomatal pits, may be very
50 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
useful in taxonomy (cf. Morley, 1953). They are quite independent of the individu- ally sunken stomata found in a number of xeromorphic leaves.
In leaves with stomata confined to areolae or to stomatal crypts the stomata are not regularly orientated, their long axes being quite randomly directed. In some leaves, however, especially narrow leaves or those with stomata present in longi- tudinal grooves, the stomata are all orientated parallel to the leaf axis. In a still smaller group of species the stomata are orientated at right-angles to the leaf midrib, e.g. Laguncularia racemosa (L.) Gaertn. f. These characters are usually of absolute constancy and of sufficient scarcity to be valuable in identification.
The frequency of stomata, either in the areolae or in the non-venous areas, is a much used character, and frequently misused. A large number of workers have used the stomatal frequency in order to distinguish between species, with varying degrees of success. Timmerman (1927), for example, found the character of little use in Datura since the frequency varied considerably on different portions of the leaf. Environmental and other factors also cause variation in frequency. Valuable taxonomic evidence, however, may often be obtained if the stomatal index (100S/ E+S), based on the proportion of stomata to epidermal cells, is used instead of the frequency per unit area (Salisbury, 1927), as has been shown by various workers in modern and fossil plant taxonomy. In Lummitzera, however, the two species only differ on stomatal frequency since the stomatal indices of the two species are identical, the stomatal frequency being greater in one due to the smaller size of all the cells.
(5) Types of trichomes and trichome-bases.
Despite some attempts by Odell (1932) to prove otherwise, trichome anatomy provides a very important group of taxonomic characters. The definition of trichomes as outgrowths of the epidermis is not absolute as there is every grade between these and the so-called emergences which differ in also involving some subepidermal layers in the outgrowth, but in the present context both types are included.
The extent to which trichomes are cuticular characters depends upon the degree to which they are cuticularized. In the case of most emergences and of trichomes with multiseriate basal parts a high proportion of the outgrowth is preserved on cuticular preparations, and the structure can be observed quite easily. Thus the whole or most of such outgrowths is cuticularized, only the more distal regions being sometimes not so covered. In the case of trichomes with pauci- or uni-seriate basal parts a similar proportion of the outgrowth may be cuticularized, but quite frequently only the very basal parts or even none at all possess a cuticular membrane. There are, of course, many exceptions to these generalizations ; for example the very long unicellular hairs of Macropteranthes leichhardtii F. Muell. are well cuticularized along their whole length. Where cuticularization is extensive cuticular preparations may provide the best means of studying the trichome anatomy.
Outgrowths with large multiseriate bases are not readily broken off at the base, but when this happens an irregular hole appears in the cuticular preparation. Trichomes with pauci- or uni-seriate bases are much more easily removed in toto
a —
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 51
from the epidermis, the appearance of the remaining trichome-base (i.e. the adjacent modified region of the epidermis, exclusive of any parts of the trichome itself) depending upon the original organization of the basal parts of the trichome, of which there are two main situations (Fig. 8). Firstly the trichome may rest upon a normal or modified epidermal cell or group of cells, or, in the case of some unicellular trichomes such as root-hairs, the trichome may in fact not be separated from the epidermal cell by a cell wall. In these types the former position of a lost trichome will be indicated on the cuticular membrane by a modified epidermal cell or cells, by a hole in an epidermal cell, or will not be visible at all. Secondly the base of the trichome may be inserted between the epidermal cells, or may replace an epidermal cell, so that a lost trichome leaves a pore in the cuticular preparation.
Vaal wot lay Van'ene ee ae Cane alone
—
Fic. 8. Four types of trichome-base as seen in vertical section.
Trichome-bases (hair-bases, gland-bases, etc.) of the above types are common features of cuticular preparations, especially as the trichomes are usually covered with a thinner cuticular membrane than the rest of the epidermis so that slight over-maceration removes the trichome completely. Their organization provides useful diagnostic characters, and it has been found necessary to create a terminology for the parts of the trichome-base (Fig. 9). The trichome-base cells may or may not surround a pore. In the former case the trichome-base cell walls may be termed poral, radial and epidermal, the first of these often being considerably thickened and then known as the poral rim.
52 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
A trichome occupying the pore of a trichome-base therefore consists of two parts : the foot, which is inserted into the pore ; and the body, which extends above the epidermis. The foot may thus be a single cell, a small group of cells, or a peg-like basal portion of a cell. The parts of the body of the trichome have been given very different names by various authors according to the trichome structure (e.g. stalk, head, branches, arms, disk, fringe, rim, etc.) and generalization is impossible and in
act undesirable.
hair-base cells
poral rim ae
base of poral rim
scarcely odi Fs pore enideemal modified foot-cell poral walls of hair-base cells radial
Fic. 9. Terminology of two main types of trichome-base.
There have been many attempts to classify trichomes (cf. Bachmann, 1886 ; Solereder, 1908 ; Foster, 1949 ; Seithe, 1960, 1962; etc.) but none has been generally successful. As mentioned previously, not only does each trichome type grade into the next but there is no real distinction between papillae, trichomes and emergences. The most usual division, between glandular and non-glandular types, suffers from the same disadvantages as other systems in that there is no sharp distinction between the groups, and in any case the division often separates obviously very closely related types. Furthermore, within each group or subgroup the direc- tions of variation are often quite different in various plant taxa, so that the only way to avoid the situation where a single trichome could belong to more than one group on different points of anatomy would be to create a system with many groups each with numerous parallel subgroups. It is thus quite clear that no useful general classification of trichomes can be constructed, but that instead it is advisable to create systems for each particular plant group investigated as it becomes necessary,
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 53
and to employ special descriptive terms in each case without attempting to correlate the terms from taxon to taxon. This has the advantage that newly discovered types can easily be fitted into the system at any time.
Good accounts of the ranges of variation and of the details of anatomy and func- tioning of trichomes are given by Solereder (1908) and Netolitzky (1932) respectively, and so no sort of review is given here. Instead an indication of the types of problems and the degree of taxonomic usefulness which may be involved in trichome studies is given by a brief statement of two major points of interest in the Combretaceae. In all twenty genera of this family peculiar unicellular hairs with distinctive basal components (known as ‘“‘ Combretaceous hairs ’’) occur, elsewhere being known only in a few genera of the Myrtaceae and of the Cistaceae. These are of great interest for they virtually delimit the Combretaceae from all other families except the other two mentioned above, and thus indicate a monophyletic origin of this family. The Myrtaceae, moreover, are generally considered to be closely related to the Combreta- ceae, and the hairs thus support this view. The presence of the hairs in the Czstaceae, however, is difficult to understand since this family has never been considered at all closely related to the Combretaceae. The identity of the Combretaceous and Cistaceous hairs thus needs to be investigated. In Strepbhonema, a Combretaceous genus usually placed in a separate subfamily from all the other genera, typical Combretaceous hairs are present in one of the species, thus confirming its inclusion in this family. In the other four species, however, they are absent and are replaced by hairs unique in the Combretaceae, so that the hairs also support the treatment of Strephonema as a separate subfamily.
The second point concerns the presence of “ scales’ (Plate 5 A), which are short- stalked trichomes with a disk-shaped multicellular head found in a wide range of plant families (Bachmann, 1886). These trichomes are often extremely distinctive, but in some families they intergrade with stalked globular glands, with stellate hairs, or with cup-shaped hairs. In the Combretaceae scales are found in all species of the small genera T/iloa, Calycopteris and Guiera, and in over half of the large genus Combretum. These four genera all belong to the same tribe (Combreteae), and the scales do seem to indicate, therefore, that this classification is well founded. In the Combretaceae the disk of the scales is always one cell thick, but varies in the degree and manner of subdivision, giving rise to a very wide range of types. In Rhododend- von, a genus of the very distant family Evicaceae, however, the main direction of variation is in the overall organization of the scale rather than in its division in one plane, producing an equally varied range of forms (Cowan, 1950 ; Seithe, 1960). Furthermore, in Rhododendron there are intermediates between all of the numerous (25 according to Cowan ; 43 to Seithe) types of trichomes, whereas in the Combreta- ceae there are three main types (Combretaceous hairs, scales and stalked glands) which are always quite distinct (Stace, 1965). Thus Cowan concluded that all the trichomes of Rhododendron are phylogenetically directly related, especially as they all appear to pass through identical early stages of ontogeny. Although this may be correct in Rhododendron I do not consider it to be the case in Combretum, where it is not possible to construct a ‘‘ phylogenetic tree ”’ of trichome types as did Cowan.
54 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
The stalked glands of the Combretaceae (Plate 5 B) are found in all the scale-less genera of Combreteae except Pteleopsis, in all the scale-less species of Combretum itself, and also in Conocarpus, a genus in another tribe. Their presence in the last genus and absence from Pteleopsis is puzzling, since the two tribes are otherwise well defined on a number of characters. The presence and types of stalked glands or scales are thus very useful taxonomic criteria in Combretum and in other genera of the Combreteae (Stace, 1961, 1963, 1965 ; Exell, 1963). In both Combretum and Rhodo- dendron the major taxonomic positions of trichome differentiation are at the sub- generic and sectional level, and where a section contains species with unexpectedly divergent trichome types there is good reason to doubt the homogeneity of that group.
Compared with trichome structure trichome size is relatively unimportant, but nevertheless at times very useful. For example the Combretaceous hairs of the Combretaceae are relatively very uniform in structure, but those of Macropteranthes leichhardtit are very distinct in their large size, and those of Guiera senegalensis extremely so in their very small size. The length of hairs generally is, of course, a frequently used taxonomic character.
The trichome-base frequently provides characters of importance, most notable of which is the shape (i.e. the degree of modification) of the trichome-base cells. In some cases they are scarcely modified from normal epidermal cells, but usually they are elongated radially to the pore, when this is present. If the normal epidermal cells are undulate-walled the radial walls of the trichome-base cells are very often straighter or straight, and frequently their epidermal walls also. Although the number of trichome-base cells varies considerably within one species, in many cases there is a typical range, of which 5-8 is the commonest, which may aid in the diagnosis of the plant. Some species, for example, have a characteristically low or high number of trichome-base cells. In some species the trichomes are so numerous that almost every epidermal cell is a trichome-base cell, and radially elongated. In Guiera (Plate 4 D) many upper epidermal cells are adjacent to two or three hair-bases, and there are no unmodified upper epidermal cells. The latter are thus typically elongated in two or three directions, and the cuticular preparations of this genus are quite distinct in the whole family. In many Combretum species (e.g. C. glutinosum Perrott. ex DC. and several related species) almost all of the lower epidermal cells are scale-base or hair-base cells.
Quite frequently, especially in xeromorphic species, the trichome-base cells are not radially elongated, but have become tangentially divided so that there are several rings of modified cells, the inner of which is composed of cells longer in the tangential than in the radial plane. The cuticular flanges of these walls are usually thick and straight. Sometimes the two different types of trichome-bases mentioned above form valuable diagnostic characters between species, although in other cases this is not so and too much emphasis should not be placed on them. In Terminalia glabrescens Mart., for example, both extremes and all degrees of intermediates may be found.
CUTICULAR: STUDIES AS AN AID TO PLANT TAXONOMY. 55
The size of the pore, where present, varies considerably, but in some cases may characterize species. In Anogeissus acuminatus (Roxb. ex DC.) Wall. ex Bedd., for example, the pores are very large (often over 50y. across) and are a quite distinctive character of this species. The thickness of the poral rim varies in two dimensions, i.e. towards the centre of the pore (Plate 5 A), and towards the centre of the leaf. In the former case it may result in a very small pore of only 1-2y. in diameter. The thick rims are frequently, though not always, accompanied by a considerable thickening of the radial walls of the trichome-base cells, so that a stellate pattern of thickening is obtained. In extreme cases, where the trichome-base cells are of the tangentially subdivided type, all the walls of the trichome-base are conspicuously thicker than the rest of the epidermal cell walls. The poral rim often becomes deeper by a development of cuticular membrane around the foot of the trichome, often up to the inner wall of the epidermis. The diameter of the pore is nearly always narrower at the base than at the apex, so that the sides and base of the pore can be seen in surface view.
Where there is more than one type of trichome on a leaf the types of trichome-bases bearing them are often different (Plate 5 D). In Combretum, for example, where hairs are always accompanied by either stalked glands or scales, the poral rims of the latter are much thicker and deeper than those of the hair-bases, and the two can be readily distinguished. Moreover the distribution of the scales is different from that of the stalked glands, the latter being commoner on the veins, the former in the areolae. Thus the abundance and distribution of each type of trichome can be ascertained even if the trichomes have been lost. In Conocarpus erectus there are two types of bases: those with radially and those with tangentially elongated cells. Stalked glands are found on all of the latter types, on intermediate types and on a few of the former types ; whilst Combretaceous hairs are confined to the former types of bases. Thus in this genus the difference is not so well marked.
(6) The cuticle.
Besides providing a means of studying epidermal anatomy in the absence of any cellular layers the cuticle often possesses characters of its own on its outer surface, and the cuticular membrane also varies in the degree of its development. All these characters are worthy of study as potential taxonomic criteria.
The most obvious feature is the degree of development of the cuticular membrane, as has been mentioned in several preceding sections, notably the cuticular orna- mentation of the guard-cells and trichome-bases. The thickness of the cuticular membrane over the normal epidermal cells is also a character of diagnostic importance. In most cases thicker membranes are characteristic of plants of drier habitats, and are thus a measure of xeromorphy, although this is not always so. In Lumnitzera racemosa and Macropteranthes kekwickii F. Muell., for instance, the wall between the lumen of the epidermal cells and the exterior is quite thick, as might be expected in these two xeromorphs. However, in the former the cuticular membrane is thick and has long stout cuticular flanges, whilst in the latter the cuticular membrane is
56 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
thin with scarcely any flanges, the non-cutinized epidermal wall accounting for the greater part of the thickness of the outer compound wall. Because of its relation to the environment, however, the degree of development of the cuticular membrane is liable to considerable phenotypic variation, and great care must be exercised in its use in taxonomy. The cuticular flanges show the same degrees of genetic and phenotypic variation and may also prove of diagnostic value in some cases, besides determining the over-all appearance of the cuticular preparation and whether or not the epidermal cell outlines are visible (Plate 5 c). In Combretum psidioides, for example, the cellular organization of the scales is always more or less completely indiscernible, whilst in all other closely related groups it is clearly visible on cuticular preparations. Again, xeromorphic species usually have well-developed flanges, but some species have thick cuticular membranes yet very short and narrow flanges.
The other character of the cuticular membrane is the type of minute ornamentation of the outer surface of the cuticle, this having been used by a number of past workers. The vast majority of plants have a smooth or minutely granulated cuticle when viewed with a light microscope, and no use can be made of this fact beyond the absence of any definite markings. The latter, as seen in many mesomorphic plants, take the form of a series of minute grooves and ridges which are commonly termed striations from their appearance in surface view. Their presence and degree of development do not always seem to be related to xeromorphy, although some species of Combretaceae have been found in which the degree: of striation is not constant. In their less-developed state the striations are usually found running parallel to the midrib and veins, and radially to the trichome-bases and stomata, and the presence of such a pattern is probably of little importance. In general the striations bear no relation to the cell outlines, each marking passing over a number of cells, but exceptions have been found where each striation is always contained within a single cell outline. When the striations are better developed they appear on the non-venous areas unconnected with the trichomes or stomata, and their orientation, arrangement and degree of development may be very various. Where they are found in areolae they are usually contained within one areola, i.e. they do not continue across the venules. Where the veins are less extensive on the epidermis the striations may be much longer. They are extremely conspicuous, for example, in Macropteranthes kekwickit and M. montana (F. Muell.) F. Muell., where they are found on all parts of the epidermis. In section the surface of the cuticle appears papillate. In extreme cases, several of which have been found in Combretum, the whole cuticle is covered with striations, those radial to the trichomes and stomata and those parallel to the veins being continuous over other areas of the epidermis (Plate 5 D). Sometimes the striations are so strong, and the cuticular flanges so weak, that the former obscure all signs of the cell outlines.
In Combretum zenkeri the long papillae have a conspicuously echinate cuticle. The series of raised cuticular anastomosing ridges reported by Solereder (1908) in several families appear to be rare.
EE
ha” = DLC
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 57
§6. DESCRIPTION OF CUTICULAR CHARACTERS
Just as the formal description of a new plant species follows a well-defined and well-known pattern, so should the characterization of a cuticular membrane, although hitherto this has not been the case. Two types of description are required for various purposes : a relatively full account of the cuticular membrane characters ; and an abbreviated account or diagnosis. A suggested form for the former, after many adjustments, is as follows :
1. Distribution of species ; habitat.
2. Morphology of the leaf : a. general morphology as seen with the naked eye. b. special morphology—venation pattern and prominence, trichome types and distribution, clear spots, pimples, etc.
3. Account of the cuticular membrane :
a. general features—thickness, ease and quality of preparation.
b. shape, arrangement and size of upper epidermal cells of : i. venation system ; ii. non-venous areas.
c. shape, arrangement and size of lower epidermal cells of : i. venation system ; il, non-venous areas.
d. shape, arrangement and size of margin cells.
e. distribution, frequency, orientation, size and structure of stomata, in- cluding the subsidiary cells.
f. distribution, frequency, size and structure of trichomes and trichome- bases.
g. presence of any other features, e.g. cork-warts, hydathodes, water-stomata, domatia.
h. presence of markings, e.g. striations, on the cuticle.
4. If necessary : interpretation of surface cuticular characters, primarily of the stomata, by means of leaf sections. A brief mention of the organization of the hypodermal and inner layers of the leaf might not be out of place as a general background to the cuticular characters.
Such a description is an obvious prerequisite for the use of cuticular characters in taxonomy and identification, and full allowance must be made for environmental and other variation. Characters which are thought to be likely to vary beyond the limits observed should be indicated.
For the purposes of a diagnosis the third section above, i.e. the description of the cuticular characters themselves, should alone be used, and this in an abbreviated though essentially similar form to that above. To this end certain abbreviations may be employed, comparable to lvs., fls. and fr. in other descriptions. The following are suggested : epid. (epiderm-is, -ides or -al); 1. and u. (lower and upper); cutic. (cuticle(s), cuticular or cuticular membrane(s), unless these need to be distinguished) ;
58 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
stom. (stom-a, -ata or -atal); ven. (venule(s)); ret. (reticulum); w.l./wall (wave- lengths per wall) ; ampl. (amplitude) ; isod. (isodiametric) ; 2ry (secondary) ; lat. (lateral(ly)) ; longit. (longitudinal(ly)) ; trans. (transverse(ly)) ; tang. (tangential (ly) ; rad. (radial(ly)).
In some cases these procedures may need to be modified, just as with normal taxonomic descriptions, but in general their use would lead to a greater uniformity and thus ease of reference and comparison. A diagnosis of a typically meso- morphic species is given here :
Diagnosis of Combretum fruticosum (Loefl.) Stuntz.
Cutic. medium thickness ; features clear. U.epid. : midrib broad and conspicuous, of many longit. rows of narrow elongated cells ; lat. veins conspicuous but narrower ; minor veins and some ven. distinguishable but ven. ret. absent or fragmentary ; cells of non-venous areas averaging c. 30 pu. across, isod., with straight walls, or less often with undulate walls with up to t4w.l./walland 5 wampl. L. epid. : midrib and lat. veins broader and more conspicuous than on u. epid. but minor veins and ven. not so much more conspicuous and ven. ret. still ill-defined ; cells of areolae averaging c. 20-25 yu. across, isod., with faint undulate walls with up to 24 w.1./wall and 8 wampl. Margin of a very few rows of slightly elongated rectangular, straight-walled cells. Stom. very frequent in non-venous areas of 1. epid., sparse alongside u. epid. midrib, randomly orientated on 1. epid., c. 22-30 X 15 wu; epid. walls very faint ; poral walls quite conspicuous ; stom. ledges absent ; poles mostly rounded, sometimes retuse or obtuse ; adjacent epid. cells 4-7, unmodified. Combretaceous hairs frequent on midribs and lat. veins of both epid., but sparse elsewhere ; hair-bases with pore c. 10-25 uw across ; poral rim some- what thickened ; hair-base cells mostly 5-9 and slightly rad. elongated, with straight, slightly thickened rad. walls ; internal compartments long and pointed. Scales bowl-shaped with raised convex cutic. membrane, frequent in 1. epid. areolae, sparse on 1. epid. venous areas and on u. epid., there frequently commoner on or near midrib ; scale-bases with pore c. 10-25 across ; poral rim considerably thickened ; scale-base cells c. 8-13, considerably rad. elongated with thin straight rad. walls ; scales c. 100-200 y across, circular in surface view, scalloped at margin, divided into c. 35-70 cells by rad. walls alone, few of the cells reaching the scale centre, all reaching the scale margin ; stalk of scale uniseriate.
§7. VARIATION OF CUTICULAR CHARACTERS
The main disadvantage of many taxonomic characters is that they will vary independently of the genotype, and it is thus important to discover to what extent and under which conditions this variation willoccur. In all groups certain characters vary so much that they are useless in taxonomy, but since these characters are not the same ones in every plant group every character is worthy of study. With regard to cuticular characters variation is due to three major causes : the age of the leaf, or rather its degree of maturity ; the environment in which the plant is situated ; and the position of the leaf upon the plant, that is to say the internal and external environment of the leaf. In general the characters affected by these three factors are those of size, frequency and degree rather than the actual anatomy or organi- zation of the particular structures. Thus the majority of the characters previously discussed show relatively little of this type of variation. Much of the variation shown within species is of course genetically controlled ; this must be equally carefully studied, and, if possible, distinguished from the environmental and develop- mental variation.
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 59
In the case of variation due to age it is obvious that in extremely young leaves every cuticular character would be quite unlike that character on a mature leaf, and such immature leaves are usually of no taxonomic use with regard to either macro- scopic or microscopic features. The present section therefore only deals with leaves that are likely to be chosen for cuticular examination, i.e. leaves which are over half-grown.
The relevant literature shows a generalized sequence of the development of the various cellular layers of a typical simple leaf to be established (Avery, 1933 ; Esau, 1953 ; Slade, 1957 ; etc.). The leaf primordium arises as a bud-like outgrowth of the apical meristem of the stem, and develops by its own apical meristem into a finger-like projection which corresponds to the petiole and midrib. The lamina later grows out from this as a pair of lateral wings, this activity of a marginal meri- stem usually commencing before the cessation of that of the apical meristem, in most cases before the leaf is a millimetre long. When apical growth has ceased the leaf elongates by intercalary growth. The thickness of the lamina in terms of cell number is determined at a very early stage in development, when the leaf is a few millimetres long, and subsequent increase in cell number is solely by anticlinal divisions. Very soon after the lamina has begun to develop the major lateral veins appear in its central portion, branching from the midrib and keeping pace with the lamina extension by intercalary growth. The lesser veins and venules develop similarly at progressively later stages. Recent work (Pray, 1963) has shown that the veinlet terminations also develop in the same manner, and not by dissociation of parts of the venule reticulum in the expanding leaf as was formerly thought to be the case.
Before the leaf approaches its mature size cell division ceases altogether, the remaining increase in leaf size being by cell expansion alone, and, although some of the layers are distinguishable before cell division ceases, most of them acquire their characteristic form after this point. The first parts of the leaf to end cell division are the two epidermides, followed by the spongy mesophyll and lastly the palisade mesophyll. Not all regions of the leaf stop producing more cells at the same time, moreover, the leaf apex reaching maturity first and the base last, and superimposed on this sequence the leaf margin continues cell division longer than the midrib region. The tip of the leaf is thus the most mature, and the marginal parts the least. Since the epidermis is the first layer to show a cessation of cell division, cell enlargement shows its greatest development there. Stomata develop from epidermal initials soon after the number of cells in the lamina thickness has been established, and their mature shape is attained in most cases before that of the other epidermal cells. The times and rates of trichome development usually appear to run parallel to those of the stomata, and in almost all the cases investigated trichomes and stomata had reached their mature structure before the leaf was its mature size. Thus one of the latest features of the epidermis to appear is the mature epidermal cell shape and size.
Examination of data from a wide range of taxa has led to the conclusion that there are six cuticular characters which are regularly to be expected to show sufficient
60 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
environmental and developmental variation to pose serious taxonomic difficulties. These are discussed separately.
A. EPIDERMAL CELL SIZE. |
A number of workers (e.g. Yapp, 1912 ; Salisbury, 1927 ; Watson, 1942) have found that the epidermal cells are larger on leaves in more humid or more shaded situations, and Odell (1932) cites workers who also discovered a decreasing cell size with excess carbon dioxide, drier air, drier soil and greater altitude. In general these size differences are proportional to a change in leaf area. The cell size also decreases with an increased height of insertion upon the axis of many herbs (Yapp, 1g12 ; Stober, 1917 ; Odell, 1932 ; Turrell, 1942 ; Ashby & Wangermann, 1950 ; etc.). Although this might seem to reflect the more shaded and humid environment of lower leaves Ashby & Wangermann (1950) concluded that the difference in cell size was not dependent upon the climate when the leaf unfolds, even though in Ipomoea at least the later the time of seed sowing the larger were the epidermal cells, but found (1950a) that the factors affecting this character operate mainly by in- creasing the period of cell division in upper leaves. Ashby (1948) discovered that the decrease in cell size was accompanied by a decrease in leaf area and cell number, and although the latter were affected by water supply the former was not, since plants in dry and wet conditions produced the same-sized epidermal cells. This is contrary to the results of most of the above workers.
Some investigations into the relation between cell size and leaf maturity were made by me on a number of species, using North’s cellulose acetate film technique (North, 1956). In the two main species investigated, Quercus robur and Carpinus betulus L., the leaves grow by cell division and enlargement until they are about one-third their mature size, when cell division ceases and subsequent leaf enlarge- ment is entirely due to cell enlargement. In species with undulate epidermal cell walls, e.g. C. betulus, the cells appear for a time to be enlarging more rapidly than the leaves. This apparent anomaly is due to the fact that cells can increase in length and breadth without any increase in area while the wall undulations are developing (cf. Fig. ro).
Secondly, the nature of the abnormally small leaves found at the base of most sucker shoots was investigated in Quercus robur, Buchenavia capitata (Vahl) Eichl. and Platanus x hybrida Brot., cells from fully mature leaves of as wide a size range on the same sucker as possible being examined. No significant cell-size difference whatsoever was found, an increase in mature leaf size of a factor of three producing no increase at all in the cell size in all three species. Representative figures obtained from Quercus and Buchenavia are:
Quercus Buchenavia Liv‘size Av. cell lgth. Li. size Av..cell lgth. 9:0 X50 cm. 30°0 wp 3°5 X20 cm. 42°5 wb 6-0 X3°0 cm. 28-0 up 2°5 XI°3 cm. 41°5 w 4°5 X3°0 cm. 310 p I-0 X0°6 cm. 40°5
3°93 X20 Cm: 28°5 w
ee
a. =
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 61
This is in direct contrast to the results obtained by Turrell (1942) and others working with herbs showing typical heteroblastic development (i.e. varying leaf types up the stem) where the upper and lower leaves differed in a number of size characters besides the overall leaf size, and thus indicates that the small leaves at the base of suckers do not represent normal heteroblastic growth. Since different-sized leaves on a single sucker differ in cell number but not cell size the factors regulating the mature leaf size must take effect early on by determining the point at which cell division stops. The same results were obtained from abnormal suckers on coppiced trees of Quercus which possessed leaves over twice as long and twice as broad as those on normal suckers, yet which had epidermal cells of the same size.
Finally, the average epidermal cell sizes of different parts of a single leaf were measured, when in every case the cells increased in size with the distance from the leaf apex and from the leaf margin. Representative measurements for Quercus vobur are : extreme leaf apex, 29 X22 uw ; extreme lamina base, 36 x31 w ; extreme margin, half-way from base to apex, 30 X20 uw ; half-way from base to apex and from margin to midrib, 34 X27 u.
These observations merely serve to emphasize the variability of epidermal cell size not only with age and minor genetic variation but with the environment, position of the leaf on the shoot and the position of the cells in the leaf. However, there does appear to be less variation than might be expected with respect to the cells of leaves on a single shoot which attain different sizes at maturity, and this is encouraging from the angle of systematic anatomy. Should the character appear to be of taxonomic use great care must be taken to use strictly comparable mature material, and to ascertain the degree of variation to be found. Thus with herbarium material this feature should not be used unless very well marked.
An additional complication, the secondarily subdivided epidermis, is unlikely to cause any confusion as the nature of the extra walls is usually obvious (see §2 above). They presumably develop long after normal cell division has stopped, and represent a renewal of meristematic activity.
A very similar situation to the above exists regarding stomatal size, although in this case the leaf size is not wholly and directly dependent upon the character. Measurements indicate, in fact, that the stomata may become mature in size and anatomy before the epidermal cells have fully expanded, and so the use of absolutely mature leaves may not be so essential. Various workers, especially those previously mentioned, have found that stomata become longer nearer the base of a herbaceous plant, in shade, in moist air and in moist soil. These results are not always capable of repetition, however, and some workers (e.g. Odell, 1932) found no difference in stomatal size on sun and shade leaves. Thus the variation of this character in the particular group concerned should be fully ascertained before it is taxonomically employed, but the evidence does suggest that the average stomatal size is less subject to phenotypic variation than that of the epidermal cells.
62 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
B. STOMATAL FREQUENCY.
Since the number of stomata that appear on a leaf is determined at the end of the period of cell division, long before a leaf reaches its full size, the stomatal frequency expressed as the number of stomata per unit area clearly decreases as the leaf expands. Again, therefore, only mature leaves are suitable for comparative measurements of stomatal frequency. Moreover the stomatal frequency often varies considerably on different parts of the same leaf, and on different leaves of the plant. Salisbury (1927) found that the lowermost leaves of a plant, and the basal and the midrib regions of the leaf, had a lower stomatal frequency, which was reflected to an equal extent on both epidermides when the upper epidermis possessed stomata. The increased frequency of stomata on higher (smaller) leaves of herbs has also been noted by Yapp (1912), Stober (1917), Rea (1921) and Turrell (1942), but Odell (1932) and others have found that the frequency increases towards the base and midrib of each leaf.
Environmental factors also regulate the stomatal frequency, the latter being lowered by humid air, wet ground, shade from sun and wind, lower altitude, and a lesser concentration of carbon dioxide (fide Odell, 1932).
Fortunately, however, the above variation can apparently be almost entirely cancelled by recording stomatal frequency in terms of the proportion of stomata to epidermal cells :
No. of stomata No. of stomata + No. of epidermal cells
<I00 =—Stomatal Index.
Since the variation in stomatal frequency caused by the above factors is apparently due to the increase or decrease in epidermal cell (and to a lesser extent stomatal) size, but not to cell number, the stomatal index is not affected. Salisbury (1927), who was the first to use this value, found that one factor did, however, alter the stomatal index—the humidity. It is not known how widespread this phenomenon is, but it seems that the diverse variation shown by stomatal frequency can often be more or less overcome. More work is needed, however, to establish that the stomatal index is always as invariable as has been claimed : Yapp’s (1912) data, for example, would suggest that it is not. Gupta (1961) also used a value which cancelled the effect of the position and environment—the absolute stomatal number. This is obtained by calculating the product of the stomatal frequency per unit area and the leaf area, and can therefore be used as an alternative to the stomatal index. Reyenga & Karstens (1964) described an unusual situation in the sepals of Hydrangea, which are small and pink during anthesis, but which afterwards turn green and enlarge considerably. This period of growth includes not only the formation of new stomata but also the disintegration and disappearance of some of the original ones, the stomatal index rising due to the predominance of the former process. Moreover, the original stomata have actinocytic subsidiary cells with radial cuticular striations, whilst those of the later stomata are anomocytic and not striated. Furthermore, Dehnel (1960) reported in Begonia the disintegration of
CULICULAR STUDIES-AS AN AID TO PLANT TAXONOMY 63
stomata which have been wounded. Phenomena of this kind are probably exceptional, however, and need rarely enter into consideration.
Rea (1921) found that in Campanula rotundifolia L. the hydathodes showed the same positional and environmental variation as the stomata.
C. TRICHOME FREQUENCY.
The general situation regarding trichome density is similar to that of stomatal frequency in that usually the number of organs is determined when the leaf is very young and the frequency thus drops as the leaf expands. Thus a densely pubescent young leaf may develop into a sparsely pubescent mature leaf. Yapp (1912), however, found that in Filipendula ulmaria (L.) Maxim. there was an additional development of hairs after the leaf began to unfold, the original pubescence being developed in the leaf bud. As in the case of stomata the pubescence often varies from leaf to leaf, from one part of a leaf to another, and under a variety of environ- mental conditions. The variation in trichome frequency on different parts of one leaf is a well-known and important taxonomic character. Yapp (1912) and Stober (1917) both found that the pubescence on plants generally rose with increasing distance up the aerial shoot : in many plants, in fact, the radical leaves are almost glabrous and the upper cauline leaves conspicuously pubescent. The inner radical leaves are also usually more pubescent than the outer.
The same workers, in general, as those who have found a variation in other characters, have also documented the variation of trichomes in different environ- mental conditions (fide Odell, 1932). Hairs have been found to be more abundant in greater sunlight, greater wind exposure, drier air, drier soil and greater altitude. McDougall (1927), however, reported that far more hairs developed in reduced sunlight in Lactuca biennis (Moench) Fernald (L. spicata auct.). It is possible that in some of these cases the proportion of hairs to cells does not vary, but at least in many instances it does. In general there is no possibility, therefore, of the use of a trichome index comparable to the stomatal index ; trichome density is a much more variable character than stomatal frequency, and its taxonomic use is thus more restricted. Only differences of a considerable degree or differences in the relative distribution of trichomes are reliable. The one advantage of a measurement of hair density in terms of hairs per cell rather than of hairs per unit area is that the former cancels the effect of age in any one leaf, except as noted below. As in the case of stomata the number of hairs per epidermal cell may remain constant as the leaf expands, and it is unfortunate that this value does not also remain constant in different environmental conditions and positions on the plant.
A further complicating factor concerned with trichome frequency is the phenomenon of glabrescence : the gradual loss of hairs with age. As explained previously most hairs may be localized on cuticular preparations by the presence of a hair-base, the centre of this often being occupied by a pore. It was thought that the number of hair-bases per epidermal cell might remain constant throughout the life of a leaf, even if most of the hairs themselves dropped off. Unfortunately
64 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
in most cases this was found to be untrue. Observations were made chiefly on a number of species of Buchenavia, Conocarpus and Combretum ranging from extremely small young leaves, from which cuticular membranes were difficult to prepare, to senescent leaves that had begun to fall from the trees. In the species of Buchenavia examined, notably B. capitata and B. kleinii, the very young leaves are uniformly densely pubescent or sericeous, whilst the very old leaves are mostly glabrous with a few hairs still remaining on the midribs and major lateral veins. Microscopic examination of the former showed that almost every epidermal cell was adjacent to one or more hair-bases, but as the leaf expanded and the hairs began to fall off the hair-bases gradually became occluded, and in the mature state the majority were not visible at all. The more actively growing (i.e. the more immature) a leaf the more quickly occluded are unoccupied hair-bases. Occlusion is effected by the hair-base cells encroaching inwards until no pore remains. When loss of the trichomes is delayed until after leaf growth has ceased, however, the vacated hair-bases often remain distinct thereafter, as is the case with the diagnostically important scales of Combretum spp., although this is not true of the hairs of Conocarpus erectus var. sericeus.
Thus the use of trichome frequency as a taxonomic character must be preceded by an exhaustive series of observations on a wide range of material in order to discover the cuticular changes accompanying growth and senescence, as well as the degrees of phenotypic variation.
D. EPIDERMAL CELL SHAPE.
The epidermal cells gain their characteristic shape gradually from the time that cell division ceases onwards, and in general this is completed before the process of cell expansion has ended. Certain types of cell shape, however, may be finally assumed after the leaf has reached full size. The only important example of this is the undulation of the lateral epidermal cell walls. Since the undulations occur equally on either side of an imaginary mid-line, i.e. into each of the two adjacent cells, the apparent length of the cell can increase (by twice the value of half the amplitude) without any increase in volume (Fig. 10).
original length = |
{\ ra original breadth — b ] Ney LWA original area = |b
original cell wall
i final cell wall nea AY) rralioagth ns vy
final breadth = b+x final area = Ib
Fic. 10. Diagram illustrating growth of cell-wall undulations.
In most cases examined, however, the development of undulations takes place whilst the leaf is expanding, and can give rise to the apparent phenomenon that the
I i ea i OO
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 65
cells are growing faster than the leaf. The final amplitude of the undulations is usually reached as the cells (and the leaf) attain full size, although the maximum growth of the undulations ceases some time before this. In Buchenavia capitata, however, the leaves at flowering time, which are fully grown but considerably thinner and more pubescent than when mature, have conspicuously less undulate walls than those at fruiting time, and in this case at least the undulations do increase in amplitude without any increase in cell area. This is most apparent on the upper epidermis. According to Watson (1942) it is the stresses set up by the hardening cuticular membrane which cause the plastic cell walls to become undulate, and when the cells have become mature their walls are too rigid to allow further increase of undulation. It is thus clear that absolutely mature leaves must be examined in order to ascertain the degree of undulation of the cell walls.
Much variation in this undulation has been noted in various environmental conditions, strongly undulate to straight walls often being observed within a single species. Brenner (1900), amongst the earlier workers, found that considerable differences could be found between plants in “‘ normal ’’ and damp conditions : in two species of Cvassulaceae the normally straight-walled cells were converted to markedly undulate-walled cells, and in a third species the exactly opposite result was obtained. A considerable number of workers have noted that the amplitude of the undulations increases with increased shade, humidity and soil dampness (Odell, 1932). Watson (1942) found that epidermal cells of Hedera in sunny situations had 6-3 crests per cell, whilst those in shade had 8-7. Moreover the latter extended throughout the height of the cells, whereas the former were confined to their outer edge. This was explained on the basis that in the sun the cells become mature and rigid more quickly, so that the hardening cuticular membrane does not have an effect to such a great depth as in the shade.
Yapp (1912), Stober (1917) and other workers (fide Odell, 1932) have found that the undulation of the cells of many species decreases the higher the point of insertion on the stem, in some cases radical leaves having undulate-walled cells and cauline leaves straight-walled cells.
With regard to other features of cell shape Brenner (1900) claimed that Crassula portulacea Lam. has strongly papillose cells in damp conditions, but flat cells in “normal ”’ situations.
Few other characters of the epidermis show as much variation as the cell-wall undulation, and although this has been used taxonomically in a number of groups its use requires the most extreme caution.
E. LEAF VENATION.
Variation in the actual venation of leaves is apparently not very frequent, and when it does occur not drastic. The characters found to vary, if any, are exemplified by the situation found in Medicago sativa L. by Turrell (1942), where the larger leaves, which had larger epidermal cells and stomata than normal, possessed larger areolae,
66 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
i.e. the number of veins and venules was not different, but the spaces between them were larger. Levin (1929) and Hall & Melville (1951) claimed that the areola size was diagnostically important, but the latter authors (1954) and Gupta (1961) later found that it in fact varied with leaf size.
The most variable venation character with regard to the present study is the degree to which the various orders of veins are represented on the epidermis. This feature appears as the epidermal cells are enlarging, but its exact time of develop- ment probably varies from group to group. In Buchenavia capitata and B. kleinit, for instance, it develops earlier on the lower epidermis, where it is quite well developed well before the cells are full-sized, than on the upper epidermis, where it has not fully developed when the leaves are fully grown: in the latter case both the smaller venules and undulations of the cells develop as the full-sized leaf matures. In fact it is the non-appearance of undulations on the venule cells which causes them to be distinguished so easily.
No comments concerning the phenotypic variation of the epidermal veins seem to have been made in the literature, but considerable variation was in fact found in a number of species. Since the species in which this was noted were all of tropical origin the reasons could not be definitely explained, but] there is little doubt that they are partly environmental and positional in origin. The differences were concerned with the size of the smallest venules distinguishable on the epidermis, and thus with the sizes of the areolae and the degrees to which the venous system appeared to be open or closed. Usually the range of variation within a species gives extremes with different-sized areolae only, but in some cases a species shows open and closed venous systems of all degrees.
It appears that the environmental conditions producing a leaf with fewer veins discernible on the epidermis in fact act primarily by producing a thicker leaf due to the larger cells and inter-cellular spaces. This in some way seems to permit fewer of the vein extensions to extend from the venules as far as the epidermis, so that fewer venules produce epidermal cell modifications. Greater shade and humidity are the commonest conditions which cause these effects.
F. DEVELOPMENT OF CUTICULAR MEMBRANE.
The mature thickness of the cuticular membrane is not attained until a fairly late stage, when cell enlargement is completed. The membrane is, however, thick enough to be isolated 7m toto even before cell division has stopped, and the cuticular flanges are then well enough developed to show quite clearly the outlines of the cells. Nevertheless absolutely mature leaves are essential for comparative taxonomic purposes.
Furthermore environmental conditions are known to affect the cuticular membrane thickness and cuticular-flange length. Skoss (1955) found that the “ cuticle ’’ was thicker in leaves exposed to the sun, and the same was discovered by a number of other workers who also correlated a thicker cuticular membrane with a drier soil, drier air, more exposure to wind, a higher altitude and other factors (fide Odell, 1932).
t
ee
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 67
Also the cuticular membrane is usually thicker on the upper stem leaves than on the lower stem or radical leaves (Yapp, 1912) but Stober (1917) reported that in some plants it was thickest on the lower stem and upper radical leaves. This cuticular thickness is accompanied by a thickening of the outer epidermal cellulose cell wall.
Variation may also be caused by other, unknown, conditions, since Stevens (1932) noted a fluctuation in the cuticular thickness on cranberry fruits (Vaccinium macrocarpon Aiton) from year to year which was not correlated with fruit size or climate, and which was exactly repeated in 33 different varieties of the plant. In Lumnitzera racemosa, a mangrove with a normally very thick cuticular membrane, a single specimen was encountered with very abnormally thin leaves due to the sparse development of the usually abundant water-storage tissue. This specimen had a very much thinner cuticular membrane than usual, but the epidermal anatomy was otherwise quite typical, and the habitat gave no clue to the reason for the peculiarity.
Conspicuous cuticular striations are absent from most leaves, but when they are _ present they do not develop until a relatively late stage, and their usefulness is restricted to mature leaves. They are, however, of great taxonomic value in some groups, being highly characteristic of three of the four species of Macropteranthes, for example. No very young leaves of markedly striate species of the Combretaceae were available, but Martens (1934) has followed the development of the cuticular striations on certain petals. There is, however, a considerable variation in the degree of development of the cuticular striations in some species, and full account of this must be taken. In Combretum molle R. Br. ex G. Don, for example, the only known gathering from South West Africa is most distinctive in its extremely conspicuously striated cuticle, the striations being so strong as to obscure the cell outlines (Stace, 1961). In tropical Africa, however, where this species is very abundant, all degrees of striation are found from the previous situation to the presence of only a few striations placed radially to each trichome-base, a situation present in many or even most species of the genus. Since the cell outlines in the latter cases are very clear, the two extremes have an entirely different aspect. The cause of this variation is, of course, unknown, but would seem likely to be environ- mental as the climate of the South West African habitat is relatively very dry.
G. SUMMARY.
It is clear that a great deal of variation is to be encountered within one species in certain cuticular characters, although little or none may be found in others. Varia- tion may be attributed to minor genetic (genotypic) differences or to certain measurable influences, both genotypic and phenotypic, of three main types— developmental, positional and environmental. Obviously variation due to all of these factors must be fully considered before the characters are utilized taxonomically.
In many or most plants the lower leaves are in a different environment from the upper, and their structure often reflects this. In Filipendula ulmaria, for example, the lower leaves are in a much more shaded and humid situation, and the type of
68 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
variation from the upper leaves which they show is similar to that which is shown by upper leaves of other plants which are in a wholly damper environment, although this is not to say that the characteristics of the lower leaves are actually caused by their environment, as are those of upper leaves of plants in damp, shaded situations. The characters of these leaves are nevertheless to be regarded as relatively meso- morphic, and those of leaves in drier conditions as xeromorphic. The commonest mesomorphic cuticular characters are : a thinner cuticular membrane and shorter flanges ; more undulate epidermal cell walls ; fewer hairs ; less densely situated stomata ; larger epidermal cells and stomata ; and larger areolae.
A third level of differentiation of mesomorphic and xeromorphic characters, beyond various parts of one plant and plants of one taxon, is between different species of one larger taxon. In this case the characters are genetically regulated, and they differ to some extent from those mentioned above, mainly in that they are concerned with structure to a greater extent than size and frequency, but they do show a number of similarities. The xeromorphic characters as exemplified by the Combretaceae are: a thicker cuticular membrane and longer cuticular flanges ; straighter cell walls ; more hairs (in a group in which hairs are of frequent occurrence the more xeromorphic species are usually the most pubescent, but in other groups xeromorphs are frequently extremely glabrous, e.g. mangroves) ; lesser development of veins and venules on the epidermis ; presence of stomata on both leaf surfaces (in those groups in which there is a tendency for stomata to appear on the upper epidermis the character is usually most developed in the most xeromorphic species, but in other groups the stomata of xeromorphs are frequently more limited in distribution on the lower epidermis alone, e.g. they may be confined to grooves or crypts) ; more sunken guard-cells ; and, within the anomocytic stomatal group, the tendency to develop distinctive epidermal cells adjacent to the guard-cells.
It is the task of the taxonomist to distinguish between phenotypic and genotypic variation. In the former case the problem is to correlate it with specific environ- mental or other factors and to ascertain the degree of the variation, and in the latter to decide at what taxonomic levels it should be recognized, if at all.
§8. GENERAL CONCLUSIONS
It cannot be too heavily emphasized that there are no guiding principles enabling assessment of the taxonomic usefulness of a given character in a taxon in which it has not been previously utilized ; neither are there any means of predicting the characters most likely to prove of greatest diagnostic importance in an uninvestigated group. It is thus unwise and unjustifiable to draw on experience in one taxon and to apply it to another, even if the two plant groups are very closely related. In the Combretaceae, for instance, the flowers and trichomes provide the most useful systematic characters in the tribe Combreteae, the fruits being relatively uniform in structure ; whereas in the closely related Terminalieae the fruits and the epidermis are the only organs showing a large number of known taxonomic criteria. The diagnostic value of pollen-grains, chromosomes and wood anatomy in this
———————
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 69
family has as yet been little studied. Furthermore, in general, characters tend to vary and assume a greater or lesser taxonomic importance quite independently of one another, and there is no single character which universally or even usually overrides all others in importance. Even in the case of the chromosomes, for which the latter has most frequently been claimed, there are very many taxa in which characters other than the number and morphology of these organelles are of greatest significance, indicating that the genetic information in the chromosomes is no more manifested in their over-all shape than in that of the flowers, fruits, leaves or other organs.
There is, therefore, no reason to doubt the validity of epidermal features merely because in some groups they do vary greatly or do not agree with differentiation by other means. If it is possible for two taxa to differ solely on the shape of the stamens or leaves then it is possible for two others to be distinguished only on the form of their hairs or epidermal cells. It is very doubtful, however, whether a difference in a single feature of one organ (whatever that organ) will differentiate between taxa at a higher level than “‘ minor genetic variants’, and almost certainly not between “species”? as normally understood. In reaching taxonomic conclusions evidence from as many sources as possible should be incorporated into the catalogue of information which will lead to the proposed classification. No facet of the plant, morphological, physiological, biochemical or genetic, should be omitted. This becomes abundantly clear when the case of taxa whose systematic position varies with the character selectively utilized (e.g. Pteleopsis and Calycopteris in the Com- bretaceae) is considered.
Undoubtedly minor exceptions to these generalizations exist, though by no means disproving the rule. There does appear, for instance, to be some degree of cor- relation between the diagnostic value of the epidermal cells and stomata and the degree of xeromorphy of the taxon. Most groups which have been found to possess taxonomically valuable epidermal characters are xeromorphic to some extent, e.g. gymnosperms, Lauraceae, Magnoliaceae, Ericaceae, Epacridaceae, various mangroves, and others. This is perhaps an expression of the idea that plants which are more highly specialized tend to be less variable, potentially as well as actually. It seems obvious that the greater the complexity of an organ the more features there are to be used by taxonomists, and thus the greater the diagnostic value of that organ. There is, in addition, the commonly held belief that characters of apparently little or no differential survival value (e.g. pollen-grain sculpturing, delimitation of cells of certain trichomes) are those with the greatest systematic and phylogenetic significance.
Attempts have often been made to generalize on the taxonomic level at which various diagnostic characters differentiate. Fritsch (1903), for example, concluded that gland anatomy was more indicative above the generic level, and the presence or absence of papillate epidermal cells at the specific level. Edwards (1935), how- ever, pointed out that in different groups stomatal frequency can be a family, generic or specific criterion, or of no systematic importance at all. This is true of
70 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
almost every character as the many exceptions to Fritsch’s and others’ claims testify. Thus, apart from noting that in a particular family or other group certain characters are more important than others at the various taxonomic levels little or no valid generalization can be made. A good example illustrating this point is the possession of a very densely or very sparsely pubescent leaf epidermis : in Ramatuella it serves to distinguish between R. argentea Kunth and R. virens Spruce ex Eichl. ; in Hibiscus vitifolius L. between subsp. vitifolius and subsp. vulgaris
Brenan & Exell ; in Conocarpus erectus between var. sericeus and var. erectus ; and —
in many species of Epzlobium between no more than various phenotypic modifications. In these cases, therefore, the level at which the character differentiates is presumably dependent upon the other characters with which it is correlated, and with the causes of the differences. Similarly, the presence of stomata on the upper epidermis is important at the family level within the mangrove habit, but amongst the non- mangrove members of the Combretaceae it is of little significance, varying from species to species or often even infraspecifically. Stomatal size is frequently directly the result of the level of ploidy, and indicates a second reason for the fact that characters differentiate at varying taxonomic levels : namely that taxonomists have often failed to accord similar ranks to groups of taxa showing exactly the same differences. Thus Rorippa microphylla and R. nasturtium-aquaticum, Galium palustre L. subsp. palustre and subsp. tetraploideum Clapham, and Ranunculus
ficaria var. ficaria and var. bulbifer Marsden-Jones are three pairs of taxa of which —
the first is diploid and the second tetraploid, although the resultant stomatal size differences apparently show differentiation at three distinct levels, if the taxonomic ranks assigned above are accepted}.
The need to investigate exhaustively all aspects of the genotypic and phenotypic variation of a “‘new’”’ character has been previously stressed. In cases where characters have been used before this preliminary survey has been undertaken the identifications attempted have often proved faulty, and have cut across decisions based on other characters. When the potentiality of the character is understood, however, and accurate identification is possible, the taxonomy of the group may on occasions be improved in the light of the discoveries. The present cuticular study has resulted in several such proposals in the classification of the Combretaceae. It is not to be expected that the new evidence will cause a drastic re-classification to become desirable, but rather that the new character might so shift the balance of the evidence from all sources that in some places taxonomic adjustments seem advisable. Entirely new classifications based on one type of evidence are usually shorter-lived than the original.
Epidermal characters, then, are likely to provide not only a means of identification (especially of sterile material) but also as valuable taxonomic and phylogenetic clues as most other characters, and they exhibit much the same characteristics with respect to their pattern of variability. Edwards (1935) commented that the
1 Rorippa microphylla was in fact first described as a variety of R. nasturtium-aquaticum, and Ranunculus
ficavia var. bulbifer has been raised to subspecific rank as R. ficaria subsp. bulbifer (Marsden-Jones) Lawalrée.
CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY 71
cuticular differences between closely related species or infraspecific taxa are usually slight or absent. This is by no means always so. When all the differences between two closely allied taxa are analysed it is usually found that a few characters are conspicuously different, many slightly or scarcely so, and others identical. Thus, although it is most likely in such a case that the cuticular characters will also be only slightly or not different, the same is true of any other single feature. Occasion- ally, however, it is just that particular character being studied that shows the greatest difference between the two taxa. Several examples where this is true of the cuticular characters have been discovered within the Combretaceae. It seems that a knowledge of the relative usefulness and an understanding of the level of differentiation of cuticular characters in the angiosperms as a whole will only be objectively obtained when cuticular studies have been absorbed into the general practice of plant taxonomy.
I do not claim that cuticular characters are of any outstandingly fundamental or all-important significance, as did some of the early exponents of the taxonomic use of pollen-grain or chromosome features. They are, however, to be regarded as characters of undoubted importance as further pieces of the jigsaw of complete systematic evidence, and at times they are in fact of greater value in identification and taxonomy than are any other characters of which we know.
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STEENIS, C. G. G. J. VAN (1953). Results of the Archbold Expeditions. Papuan Nothofagus. Journ. Arnold Arb. 34 : 301-374.
GULICULAR SLUDIES AS-AN ALD O° PLANT, TAXONOMY 77
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78 CUTICULAR STUDIES AS AN AID TO PLANT TAXONOMY
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Bull. B.M. (N.H.) Bot. 4, PLATE 1
A, Rhizophora mucronata Lam.: lower epidermis of leaf, showing cork-wart (x 420). B, Buchenavia kleinii Exell : upper epidermis of leaf, showing wound caused by fungal attack (420). C, Terminalia catappa L.: leaf margin, showing characteristic hole (420). D, Lumnitzera racemosa Willd. : lower epidermis of leaf, showing water-stoma ( X 420).
er Fe -
Bull. B.M. (N.H.) Bot. 4, 1 PLATE 2
a”
A, Buchenavia grandis Ducke : lower epidermis of leaf, showing marsupiform domatium (x80). B, B. capitata (Vahl) Eichl. : lower epidermis of leaf, showing marsupiform domatium (x60). C, Terminalia catappa L. : lower epidermis of leaf, showing lebetiform domatium (x80). D, Conocarpus erectus var. sericeus Forsstr. ex DC. : lower epidermis of leaf, showing lebetiform domatium ( x 60).
Bull. B.M. (N.H.) Bot. 4, 1 PLATE 3
os Se. ast <A. @ ne they
an oe,
A, Rhizophora mangle L. : lower epidermis of leaf, showing strongly cuticularized stomata and straight epidermal cell walls (x 420). B, Buchenavia fanshawei Exell & Maguire : lower epidermis of leaf, showing thinly cuticularized stomata and undulate epidermal cell walls ( x 420). C, Combretum nigrescens King : leaf margin (170). D, Laguncularia racemosa (I..) Gaertn. f. : leaf margin ( x 170).
I aS eo
A, Ramatuella virens Spruce ex Eichl. : lower epidermis of leaf, showing anomocytic stomata (x420). B, Laguncularia racemosa (L.) Gaertn. f. : lower epidermis of leaf, showing cyclocytic stomata (x 420). C, Strephonema sericeum Hook. f. : lower epidermis of leaf, showing paracytic stomata ( x 420). D, Guiera senegalensis J. F. Gmel. : upper epidermis of leaf, showing numerous simple hair-bases ( x 420).
Bull. B.M. (N.H.) Bot. 4, 1
PLATE 5
A, Thiloa glaucocarpa (Mart.) Eichl. : lower epidermis of leaf, showing scale with strongly cuticularized scale-base ( x 420). B, Combretum obovatum F. Hoffm. : lower epidermis of leaf, showing stalked glands on the raised venule reticulum (170). C, Ceriops tagal (Perrott.) C. B. Robinson : lower epidermis of leaf, showing stoma and cuticular flanges in profile on cuticular membrane folded inside outermost ( x 420). D, Quisqualis hensir
(Engler & Diels) Exell : upper epidermis of leaf, showing cuticular striations and three types of trichome-base ( x 420).
9 Lage Sea ieee ae ee a ts a, oe | ,
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n
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4
W. A. SLEDGE
zs eS é 1 MAY 1967 ¢ ~ THE GENUS ELAPHOGLOSSUM IN THE Gy eS INDIAN PENINSULA AND CEYLON
BY
W. A. SLEDGE.
(University of Leeds) “VW
Pp. 79-96
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) BOTANY Vol. 4 No. 2 LONDON : 1967
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THE GENUS ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON
By W. A. SLEDGE
In a comment on the systematic list of ferns in a paper by Manton and myself on the cytology and taxonomy of the pteridophytes of Ceylon, I expressed the view (Phil. Trans. Roy. Soc., ser. B. 238 : 158 (1954)), that the names currently employed for the four species of Elaphoglossum were in need of revision. It subsequently became apparent that both the taxonomy and the nomenclature of the Indian species also required re-examination.
The names employed in Beddome’s works are mostly those originally given to African or tropical American plants with which Indian ones were at that time considered identical. The same names are now used in a more restricted sense and the discrimination of several species with more limited distributions within the areas of the aggregate species has left in doubt the correct names to be assigned to Indian and Ceylon species. So far as I am aware the only paper on Indian species of Elaphoglossum which has appeared since the time of Beddome is one by Biswas (Bull. Misc. Inf. Kew 1939 : 237-241 (1939)) in which two new species are described. Both of these supposed new species however are unnecessary renamings of species already described by Blume and Feée.
In his Ferns of Southern India, Beddome described under the name Elaphoglossum conforme a species which appears to be not uncommon in the Nilgiri Hills. Biswas, who described it as new, was correct in rejecting this identification but appears not to have questioned the identification of another species growing in the Nilgiri and adjoining hills which Beddome and later authors called E. stigmatolepis (Fée) Moore ; indeed he makes no reference anywhere in his paper to this species. Beddome’s FE. conforme was in fact based on specimens of the same species as that first described by Fée as Acrostichum stigmatolepis, and it is therefore Beddome’s E. stigmatolepis which requires a new name. In his Handbook to the Ferns of British India, Ceylon and the Malay Peninsula (1883), Beddome amplified the distribution credited to E. conforme to include the Himalayas, Ceylon and Malaya. The plant from the Himalayas is EF. marginatum, which had long been regarded as a synonym of E. conforme but which Biswas correctly reinstated as a distinct species. Neither E. stigmatolepis nor E. marginatum occur either in Ceylon or Malaya and these regions were included through errors of identification and do not cover a third species un- accounted for amongst those described by Beddome.
The Indian specimens which Beddome originally referred to E. laurifolium (changed in his Handbook to E. latifolium) are identical with Blume’s Javan Acrostichum angulatum, which name he cited as a synonym of EF. laurifolium in his Ferns of Southern India. This is the other species which Biswas needlessly described as a new species. E. angulatum is an easily recognized species on account of its long- creeping rhizome and very distinctive rhizome scales, yet most Ceylon gatherings of
BOT. 4, 2. 6§
82 ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON
this species are identified in herbaria not as EF. laurifolium but as E. conforme. The true Acrostichum laurifolium of Thouars is confined to Tristan da Cunha and Gough Island, and another species from Ceylon and southern India with short- creeping rhizome, closely placed fronds and different rhizome scales, which is usually labelled E. laurifolium in herbaria, is E. commutatum, a species which was founded by Mettenius partly on a Ceylon gathering made by Thwaites but which is not illustrated in Beddome’s works.
As regards the ferns that Beddome called E. sguamosum, this name was currently used to cover superficially similar plants ranging from tropical America eastwards to southern India and Ceylon. African, Madagascan and Mascarene representatives of the aggregate species have since been split off from Jamaican plants, whence the type was described. The Nilgiri and Ceylon plants which were united under the same name by Beddome are certainly distinct from one another and as neither can be matched by African or Mascarene gatherings, I have described these as two new species, using names already given to herbarium specimens at Kew by Krajina but never published by him.
Of the names used by Beddome, only E. spatulatum stands unchanged and even this very distinct species has recently been incorrectly excluded as a Ceylon species by Madame Tardieu-Blot (Not. Syst. 15 : 432 (1959)), and the equally distinct Mada- gascan EF. schizolepis has erroneously been substituted for it.
The confusion which has surrounded the application of so many of these names is discussed in more detail later in this paper under the individual species ; but it may be useful to list here the illustrations and names given in Beddome’s Ferns of Southern India together with their correct identifications :
F.S.I. t. 196 as E. viscosum = FE. stelugerum (Wall. ex Baker)
Alston & Bonner F.S.I. t. 197 as E. squamosum = E. nilgiricum Krajina ex Sledge F.S.I. t. 198 as E. conforme = E. stigmatolepis (Fée) T. Moore F.S.I. t. 199 as E. stigmatolepis = E. beddomei Sledge F.S.I. t. 200 as E. laurifoium = E. angulatum (Blume) T. Moore F.S.I. t. 213 as E. spathulatum = E. spatulatum (Bory) T. Moore
[ E. marginatum (Wall. ex Fée)
T. Moore
No illustration in F.S.I. 4 E. commutatum (Mett.) Alderw.
van Rosenb. | E. ceylanicum Krajina ex Sledge
At the time of writing, a proposal (Anderson in Regn. Veg. 40 : 18 (1965)) to con- serve the generic name Elaphoglossum Schott ex J. Smith (Hooker’s Journ. Bot. 4: 148 (Aug. 1841)) against Aconiopteris C. Presl (Tent. Pterid. : 236 (1836)) is under consideration. It has been approved by the Special Committee for Pteridophyta (Taxon 15 : 333 (1966)) but awaits ratification.
My grateful thanks are tendered to the Directors and Curators of the herbaria at the British Museum (Natural History) (BM), the Botany School, University of Cambridge (CGE), the Royal Botanic Garden, Edinburgh (E), the Royal Botanic
ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON 83
Gardens, Kew (K) and the Division of Systematic Botany, Peradeniya, Ceylon (PDA) for access to, or for the loan of, specimens in their charge.
KEY TO THE SPECIES
Fronds coriaceous or at least stiff in texture, margins with a cartilaginous border, surfaces glabrous or with minute scales. Rhizome long-creeping, fronds spaced. Rhizome scales papery, broad, pale brown; lamina 2-5-4:5 cm. broad 1. E. angulatum Rhizome scales not papery, dark brown or black ; lamina 1-5—2-5 cm. broad 2. E. stigmatolepis Rhizome short-creeping, fronds crowded. Rhizome scales entire, linear (5-10 times as long as broad), stipes much shorter than lamina ‘ . 3.£E. beddomer Rhizome scales more or less fimbriate, narrow (4-5 times as long as broad), stipes up to half as long as lamina. Rhizome scales dark brown, 5 mm. long, acuminate 4. E. marginatum Rhizome scales brown, 10 mm. long, attenuate with crisped tips 5. E. commutatum Fronds thickly herbaceous or soft in texture, margins without a cartilaginous border, surfaces conspicuously scaly or hairy. Scales of stipe and lamina stellate with long rays : ; 6. E. stelligerum Scales not stellate. Scales ovate with long marginal teeth. Fronds densely paleate, the lower surface wholly obscured by imbricating scales ; scales of stipe broad, mostly concolorous 7. E. nilgiricum Fronds paleate but neither surface obscured by scales; scales of stipe narrow, mostly with their borders and teeth dark-coloured 8. E. ceylanicum
Scales filiform, hair-like : ; g. E. spatulatum
1. Elaphoglossum angulatum (Blume) T. Moore, Index Fil. : 5 (1857).
Acrostichum angulatum Blume, Enum. Pl. Jav. : 101 (1828) ; Fl. Jav., Fil. : 25, t. © (1829).— Fée, Mém. Fam. Foug. 2: 32 (1845).—Kunze in Linnaea 24: 248 (1851).—Christ in Neue Denkschr. Schweiz. Ges. Naturw. 36 (1, 2) : 46, 49 (1899).
Olfersia angulata (Blume) C. Presl, Tent. Pterid.: 234 (1836).
Acrostichum conforme sensu Hook., Sp. Fil. 5 : 198 (1864) pro parte ; non Swartz.—Hook. & Baker, Synops. Fil. : 401 (1868) pro parte.
Acrostichum marginatum sensu Thw., Enum. Pl. Zeyl.: 380 (1864) ; non Wall. ex Fée.
Elaphoglossum laurifolium sensu Bedd., Ferns S. Ind. : 67, t. 200 (1864) ; non T. Moore.
Elaphoglossum latifolium sensu Bedd., Handb. Ferns Brit. Ind. : 416 (1883) ; non J. Smith.
Elaphoglossum ogatai C. Chr., in Dansk. Bot. Ark. 9 (3) : 67 (1937).
Elaphoglossum krajinae Biswas in Bull. Misc. Inf. Kew 1939 : 240, t. 1 fig. 2 (1939).
Rhizome long-creeping with stipes I-3 cm. apart, clothed with pale, rufous-brown, papery, thin-walled, broadly ovate, acute scales, 5 X 2 mm., with subentire margins. Fronds stalked 15-35 (45) cm., stipe of sterile fronds up to 15 cm. or more, often as
84 ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON
long as the lamina but sometimes only half or less their length, scaly, the scales like those of the rhizome but with irregularly fringed margins ; lamina elliptic, 2-5-5 cm. broad, apex acute, base narrowed and shortly decurrent on the stipe, margins with a cartilaginous, revolute border, lower surface with scattered, minute, brown, more or less stellately-laciniated scales, midrib with a few larger irregularly laciniate scales ; texture coriaceous. Fertile fronds on stalks about as long as or longer than the sterile ones, 2-3 cm. broad. Spores 27-30 x 18-21.
SoutH Inp1A: Anamallays, 1,800 m., on trees, “ F.S.Ind. f. 200 Elaphoglossum laurifolium”’ Beddome (BM). Kudiabad, Sispara Rd., 2,400 m., Nov. 1883, Gamble 13473 (K). Bear Shola, Kodaikanal, Pulney Hills, rg May 1898, Bourne 4972 (K). Kodaikanal Waterfall, Pulney Hills, 30 May 1898, Bourne 4973 (K).
CEYLON: Thwaites C.P. 1311 (BM; CGE; K; PDA). Gardner 1165 (BM; CGE, Nuwara Eliya, on trees, 1,500-2,100 m., Sept. 1844; E; K). Gardner 1310 (E; K). Nuwara Eliya, Thomson (K). Same locality, Freeman 369A, 370B, 371C (BM). Same locality, Day in Herb. Henderson (E.). Same locality, creeping over tree trunks, 9 May 1906, Matthew (K). Pedrotalagala, 2,100 m., on mossy trees, 26 Dec. 1950, Holttum (SING). Adams Peak, on trees, 2,100 m., 14 Feb. 1908, Matthew (K). Same locality, 1,950 m., 14 Dec. 1950, Sledge 615 (BM). Namunukula, Uva Province, 1,950 m., 24 Feb. 1954, Sledge 1208 (BM). 1855, R. W. Rawson 1039 (BM). Mrs. Carr ex Herb. T. Moore (K). Wallin Herb. Henderson and in Herb. Neill Fraser (E).
South India (Nilgiri, Pulney and Anamallay Hills), Ceylon, Sumatra, Java, Borneo, New Guinea, Philippine Islands, Tonkin, Formosa.
The long-creeping rhizome and broad, membranous, pale brown or rufous-brown, concolorous scales readily distinguish EF. angulatum from all other Asiatic species. It differs also from all other Indian or Ceylonese species in having the veins united by a connecting strand at their extremities—a character of the Aconiopteris (E. nervosum (Bory) Christ) group. This structural feature is however invisible in living and herbarium specimens and can only be seen after treatment with a clearing agent. Its spores are smaller than those of the other marginate-leaved species occuring in India and Ceylon. E. stigmatolepis, which it most resembles in growth habit, may be distinguished by its dark rhizome scales and the narrower fronds appearing punctate beneath on account of the more peltate scales and the black points of attachment left where these have been shed. |
Ceylon gatherings of this fern are usually named FE. conforme and Indian ones either E. conforme or E. laurifolium (or E. latifolium). They certainly resemble African E. conforme but they agree far better and are indeed indentical with authentic specimens of Blume’s A. angulatum at Kew and with other gatherings from Java. Kunze (loc. cit.) long ago reached the same conclusion when he referred a Nilgiri gathering (Schmid 65) to this species. A. angulatum was later merged in A. conforme by Hooker in the Species Filicum, and subsequent authors followed Hooker in their treatment of Ceylon and Indian plants.
Beddome’s illustrations of what he names F. conforme and E. laurifolium in his Ferns of Southern India and the descriptions in his Handbook have been the greatest
ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON 85
source of confusion surrounding the taxonomy of Indian Elaphoglossums. In the former work his illustration (t. 198) and description, purporting to be A. conforme Swartz, were based on plants on a sheet in the Kew collection annotated by him “type of tab. 198 F.S.I.”’. The specimens on this sheet represent unmistakable though small and not very characteristic examples of EF. stigmatolepis (Fée) Moore. (Beddome’s illustration (t. 199) of what he calls E. stigmatolepis Fée represents another species.) The two different scales depicted represent examples taken from the rhizome and, in the case of the shorter and broader one with markedly jagged margins, from the stipe or midrib of the frond. The enlargement of a portion of the underside of the frond shows stellate scales quite different from the minute peltate scales with shortly fimbriate margins which characterize EF. stigmatolepis, and this drawing could not possibly have been made from any of the fronds on the sheet referred to. There is moreover no reference in the accompanying description or in the later description in the Handbook to the presence of stellate scales on the fronds. The earlier work similarly contains no description of the form or colour of the rhizome scales, but in the Handbook these are referred to as “ blackish ”’ in colour, a description which is applicable to the specimens on the sheet and appropriate for F. stigmatolepis (and even, though somewhat less accurately, for E. marginatum, which, from the amplified distribution assigned in the Handbook to “ conforme’’, he now included in that species) but which is completely false for E. angulatum or for the true African E. conforme.
It is clear from the evidence provided by Beddome’s own specimens at Kew and the British Museum that he was unfamiliar with FE. angulatum for, whereas E. stigmatolepis (i.e. Beddome’s E. conforme) is stated to be “‘ very common ”’ and is represented by several gatherings of his, he appears to have collected E. angulatum once only. This single gathering is in the British Museum collection, the sheet being labelled by Beddome ‘“‘ Anamallays. 6,000 ft., on trees. Elaphoglossum laurifolium (Thouars) F.S.Ind. f. 200”. The description of E. laurifolium in the Ferns of South. India is followed by the citation of ‘‘ FE. angulatum Bl. En. 2o1 [sic] : Id. FI. Jav. 25, t.6”’ in synonomy and the reference to the rhizome as ‘“‘ wide-creeping with bright chestnut or golden scales’’ and the other distinctions attributed in the Handbook to his E. conforme and E. latifolium (the name there substituted for E. laurifoliwm of the Ferns of Southern India) become explicable when it is realized that his E. conforme is based on E. stigmatolepis and his E. laurifolium (or E. latifolium) is based on E. angulatum.
Christensen states (in Gard. Bull. Str. Settl. 7 : 2gr (1934)) a propos of Blume’s figures of his A. conforme (i.e. E. commutatum) and A. angulatum, “I am by no means sure that the two ‘ species ’ illustrated by Blume are really different’. And so it might seem from the plates themselves, for whilst the plate of the latter is an admirable representation of E. angulatum, that of the former is less characteristic of E. commutatum particularly in the spacing and small size of the fronds. Blume’s drawings and descriptions of the rhizome scales of his two species however indicate such considerable differences and they agree so very closely with the scales of E. commutatum and E. angulatum that I have no hesitation in accepting the descriptions and the plates as illustrations of these two species. Fée’s plate of A. laurifoliwm does not represent the same species, It is a Mascarene plant which has since been
86 ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON
described by Mme. Tardieu-Blot (Not. Syst. 15 : 433 (1959)) as E. alstont.
The type of E. ogatai is at the British Museum. Of this Christensen (Dansk Bot. Ark. 9 (3) : 67 (1937)) says “‘ it is extremely like E. angulatum (Bl.) Moore . . . differs however in its remarkably thin texture and the distinctly but shortly decurrent lamina’’. Tagawa, who is familiar with the living plant, states (in Mem. Coll. Sci. Univ. Kyoto, ser. B, 20: 28 (1951)) that its fronds are thin-coriaceous and that no distinction can be drawn in the other respects. One of Bourne’s Indian plants (Bear Shola, Pulney Hills) also has quite thin fronds, and the more ovate rather than elliptic fronds which look distinctive in the type specimen of EF. ogatai can be matched by a specimen of Matthew’s from Adam’s Peak, Ceylon. I therefore accept Tagawa’s reduction of E. ogatat to a synonym of EF. angulatum.
Biswas’s E.. krajinae is quite clearly no more than a redescription of E. angulatum, which he evidently never took into consideration when separating Indian and Ceylon plants from African E. conforme.
2. Elaphoglossum stigmatolepis (Fée) T. Moore, Index Fil.: xvi, 15 (1857), 368 (1862).—Christ in Neue Denkschr. Schweiz. Ges. Naturw. 36 (1, 2) : 52 (1899).
Acrostichum stigmatolepis Fée, Mém. Fam. Foug. 2 : 62, t. 24 f. 2 (1845).—Hook., Sp. Fil. 5 : 216 (1864). Elaphoglossum conforme sensu Bedd., Ferns S. Ind.: 67, t. 198 (1864) ; Handb. Ferns
Brit. Ind.: 416, fig. 247 (1883) pro parte; non J. Smith. Elaphoglossum ballardianum Biswas in Bull. Misc. Inf. Kew 1939: 239, t. 1 fig. I (1939).
Rhizome long-creeping, stipes 5-10 mm. apart, clothed with ovate-lanceolate scales, 3-5 X I-2 mm., brown at the base and blackish above, with subentire or sparsely fringed margins and acute tips. Fronds stalked, 15-35 cm. long, stipes of sterile fronds variable in length, sometimes as long as the lamina but commonly much less, scaly, at least when young, with rather broad scales with jagged margins; lamina narrowly elliptic, 1-5-2:5 cm. broad, apex acuminate, base narrowed and decurrent on the stipe, margins with a narrow, revolute, cartilaginous border, midrib sparsely scaly beneath with short broad scales like those on the stipe, lower surface clothed with minute, dark, peltate scales with fimbriate margins, upper surface with scattered scales of the same form when young, becoming smooth with age; texture stiffly mem- — branaceous to subcoriaceous. Fertile fronds on stipes normally exceeding those of the sterile fronds, 1-2 cm. broad. Spores (39) 42-45 (48) x 27-30 mM.
SoutH InpiA: “Ind. Orient Neilgherries, 159’ (CGE, ?isotype). Nilgiris, 2,100 m., “ type of tab 198 F.S.I.’”’, Beddome (K). Same locality, 1860, Beddome 5 (K). Same locality, 2,100 m., Oct. 1889, Gamble 21420 (K). Same locality, 2,100 m. Nov. 1870, Henderson (E). Same locality, Wight 50 (K). Same locality, on trees and rocks, 1842, Gough 3234/58 (K). Ootacamund, 2,100 m., Oct. 1885, Gamble 16963 (BM). Same locality and date, Gamble in Herb. Blanford (E). Same locality, 2,100 m., Aug. 1885, Gamble 1666r (K). Same locality, Babcock (K). Same locality, Cockburn 85 (BM). Dodabetta, Nilgiri District, 2,550 m., June 1883, Gamble 12038 (E). Kudiabad, Sispara Road, 2,400 m., Nov. 1883, Gamble 13474 (K).
ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON 87
Moir Point, Pulney Hills, 2,340 m., stream, July 1937 and May 1938, Avoticasamy 10 (BM).
South India (Nilgiri and Pulney Hills).
Elaphoglossum stigmatolepis differs from the other marginate-leaved species in its narrower, less coriaceous fronds, which are more tapering above and studded beneath with minute disc-like scales with shortly fimbriate margins. The points of attach- ment to the lamina appear as black dots when the scales have been shed. These are referred to by Fée as “petits points qui paraissent étre de nature glanduleuse: ne serait-ce pas la base persistante des squames?”’. Young fronds show more sparsely distributed scales on the upper surface and in older fronds, where they have been shed, similar minute black scars are recognizable. Fée does not allude to these in his description but they are correctly shown in his illustration.
The rhizome scales are more pointed than those of E. angulatum, pale brown at the base but almost black, and often somewhat glossy, in the upper parts, and hence through imbrication they appear darker than the uniformly pale, rufous-brown and thinner, more papery scales of EF. angulatum. As in that species, the stipe bears broad, brown scales with jagged margins and these are continued on to the lower part of the midrib. The spores are considerably larger than in FE. angulatum, agreeing in size with those of E. marginatum, from which, and from E. commutatum, it differs in its more strongly creeping rhizome, in its frond form and texture, and in its scale characters.
Fée’s description of E. stigmatolepis was based on a gathering ‘‘ in herb. de Lessert., [sic] sub no. 159’’. The collector’s name was evidently not stated on the label but the provenance is given as “in Indiis orientalibus, Neilgherries”’. There is in the Cambridge University collection a sheet bearing a complete and well-preserved plant with three sterile and one fertile frond attached to a long-creeping rhizome 15 cm. long. No collector’s name is inscribed on the label but this also reads “‘ Ind. orient Neilgherries 159’. The specimen agrees so exactly with Fée’s description and plate that it is almost certainly an isotype. Wight 50 in herb. Kew is also an excellent match for Fée’s illustration and the Cambridge plant.
In Fée’s description the fronds are said to be “‘ membranaceis, papyraceis ’’ and he does not include this species in the coriaceous-leaved group to which A. margina- tum, A. conforme and A. laurifolium are referred. At the same time the fronds are described as narrowly revolute, and revolute edges are commonly associated with the stiff-leaved species. I have seen no living plants of E. stigmatolepis but to judge from herbarium specimens the fronds are thin but stiff and are distinctly less coriaceous than those of the other marginate-leaved species.
E. ballardianum Biswas is merely a redescription of E. stigmatolepis and inaccurate in the statement that the fronds lack a border. One of the gatherings which Biswas cites is labelled Acrostichum stigmatolepis and another has had the same name pen- cilled on the sheet, yet Biswas makes no reference to this species anywhere in his paper, though he could hardly have failed to note its description in the literature which he cites or that it was founded on plants from the identical area of his own Supposed new species.
88 ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON
3. Elaphoglossum beddomei Sledge, sp. nov.
Elaphoglossum stigmatolepis sensu Bedd., Ferns S. Ind.: 67, t. 199 (1864) ; Handb. Ferns Brit. Ind.: 418 (1883) ; non T. Moore.
Acrostichum stigmatolepis sensu Baker in Hook. & Baker, Synops. Fil., ed. 2: 521 (1874) ; non Fée.
Rhizoma repens, palezs linearibus brunneis marginibus integris 6-9 mm. longis basi Imm. latis vestitum, stipitibus crebrioribus. Frondes stipitatae, 20-35 cm. longae ; stipites frondium sterilium plerumque breves et 3-6 cm. longi, vel, si longiores, dimidia parte longitudinis laminae breviores, juventutes paleaci paleis linearibus fimbriis marginalibus paucis ; lamina coriacea, anguste elliptica, 2-3 cm. lata, in apicem acutum diminuta, bas: gradatim decrescens et in alas angustas aut cristas in stipite decurrens, marginibus angustis revolutis cartilagineis, infra paleis minutis stellatis vestita, supra in frondibus junioribus paleis similibus vestita, in vetustioribus glabrescens. Frondes fertiles eas steriles longitudine tota aequantes sed stipes quam lamina aequaliter longus ; lamina I-5—2:5 cm. lata. Sporae (36) 39-42 (45) x 24-30 [.
SouTtH Inp1A: Anamallays, 1,050 m., “ F.S.Ind. f. 199 Elaphoglossum stigmatolepis Kunze ’’, Beddome (BM, holotype). Palghat Hills, 1,500 m., Beddome (K). Nilgiri Hills, 2,100 m., Beddome (K). Same locality, Beddome 6 (K). Same locality 1,500 m., 1870, Henderson (E). Sispara Ghat, Nilgiri District, Nov. 1883, Gamble 13418 (K). South India (Nilgiri, Anamallay and Palghat Hills).
Elaphoglossum beddomei differs from E. stigmatolepis in its longer and narrower, uniformly brown rhizome scales, the margins of which are entire or. at most with an occasional marginal fimbriation. Its shortly stalked more coriaceous fronds are decurrent on the stipes, forming narrow wings or ridges often extending to the base, and the minute scales on the under surfaces of the fronds are stellately laciniated. The stipes bear linear scales markedly different from those of E. stigmatolepis, in which they are short and broad and are continued on to the midrib of the lower surface of the fronds.
Beddome’s illustration well portrays the habit of this species. In one of the three sterile fronds depicted the narrow wing extending downwards from the decurrent lamina to the base of the stipe is correctly shown. Sometimes the stipes are con- siderably longer than those illustrated, and then the decurrent wings merge into ridges on the sides of the stipes. The fringed scale represents one of those clothing the stipe, as the rhizome scales have margins which are mostly quite entire though a few fimbriations may sometimes be present.
When Beddome first used the name E. stigmatolepis for this species he added a query both after the name and following the citation of Fée’s figure. Baker’s annotations on Beddome’s specimens at Kew and his citation of Beddome’s illustra- tion show that this was the plant which he had in mind when adding Acrostichum stigmatolepis to the second edition of the Synopsis Filicum and, following Baker, the query was dropped by Beddome in his Handbook. Two distinct species however occur in the Nilgiri and adjoining hills in South India and the true E. stigmatolepis is the plant which was subsequently distinguished by Biswas as E, ballardianum, He
ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON 89
evidently accepted the present species, on the evidence of Beddome’s illustration and the named sheets at Kew, as being E. stigmatolepis, though he makes no reference to this species anywhere in his paper but compares E. ballardianum with E. conforme, to which its relationship is certainly more remote.
4. Elaphoglossum marginatum (Wall. ex Fée) T. Moore, Index Fil.: 8, 11
(1857), 361 (1862).
Acrostichum marginatum Wall., Numer. List: 2, n. 17 (1829), nom. nud.
Acrostichum marginatum Wall ex Fée, Mém. Fam. Foug. 2 : 31 (1845), excl. syn. Blume.
Acrostichum conforme sensu Hook., Sp. Fil. 5: 198 (1864) pro parte.—Hook. & Baker, Synops. Fil. : 401 (1868) pro parte.
Elaphoglossum conforme sensu Bedd., Handb. Ferns Brit. Ind. : 416 (1883) pro parte minore et excl. fig. 247; non J. Smith.—Tagawa in Mem. Coll. Sci. Univ. Kyoto, Ser. B, 20 : 29
I95I). Bain Christ in Bull. Herb. Boiss. 6 : 867 (1898) ; in Neue Denkschr. Schweiz. Ges. Naturw. 36 (1, 2) : 51 (1899).
Rhizome stout, short-creeping, stipes close, clothed with dark brown, thick-walled, narrow, lanceolate, attenuate, shortly and irregularly fringed scales, 4—5 * I-I:5 mm. Fronds stalked, 20-50 cm., stipe of sterile frond up to half as long as lamina but often much less, deciduously scaly ; lamina elliptic, 2-4 (5) cm. broad, apex acute, base gradually narrowed and decurrent on the stipe, margins with a translucent cartilaginous border, lower surface with scattered minute, more or less stellately laciniated, brown scales, upper surface glabrous; texture coriaceous. Fertile fronds on longer stalks than the sterile ones, lamina slightly narrower. Spores 42-45 X 27-30 M.
NEPAL: Sheopore, 1829, Wallich 17 (K). Wallich 774 (BM). Sheopuri Lekh, Katmandu, 1,350 m., 19 Aug. 1954, Stainton, Sykes and Williams 6917 (BM). Between Torke and Okhaldunga, 1,950 m., 2 Nov. 1954, Zimmermann 1997 (BM).
SIKKIM : Darjeeling, 2,280 m., 9 Aug. 1875, C.B. Clarke 26913 (K). Same locality, 1,800 m., Oct. 1881, Gamble 9884 (K). Kohima, 1,800 m., 21 Oct. 1885, C. B. Clarke 40971 (K). Neebay (?), 2,250 m., 16 Oct. 18—, C. B. Clarke 25337 (BM). Jeylep Rd., 1,800 m., Nov. 1880, Gamble 9938 (K). Treutler 988 (K). 2,100 m., Sept. 1882, Levinge (K). Jerdon (K).
_ Buutan: Kancham, Punakha, 1,950 m., 28 Aug. 1914, Cooper 3058 (BM). Trashiyangsi Valley, Tobrang, on rock faces, dense jungle, 29 Aug. 1934, Ludlow & Sherriff 902 (BM; E).
AssAM : Sohra River, 1,500 m., 16 Oct. 1872, C. B. Clarke 19239 (BM; K). Same locality, 1,500 m., 28 Nov. 1871, C. B. Clarke 14778 (K). Same locality, 1,350 m., 15 Oct. 1872, C. B. Clarke 18826 (K). Vale of Rocks, Sept. 1886, C. B. Clarke 45828 (K). Suruseem, Khasia, 1,200-1,800 m., 26 May 1850, Hooker & Thomson (K). Kalapanee, Khasia 1,650 m., 6 Aug. 1850 and 28 Oct. 1850, Hooker & Thomson (K). Assam, 1870-80, G. Mann (K). Ukhrul, Manipur State, on rocks in the forest, 1,800 m., 12 Aug. 1948, Kingdon Ward 17935 (BM).
Himalayas eastwards from Nepal, Sikkim, Bhutan, Khasi Hills, Manipur, Yunnan, Formosa,
go ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON
Elaphoglossum marginatum differs from the other coriaceous-leaved species in its thicker, more shortly creeping rhizome with more crowded stipes. E. angulatum is — readily distinguished by its long-creeping habit with well-spaced fronds and much broader, thinner and light brown rhizome scales. E. stigmatolepis also differs in its more rampant habit, its narrower fronds, which are more tapering above and below, and by the different form of the scales on the lower surface. Both Fée and Moore included Java in the distribution of E. marginatum through inclusion of the plants later distinguished by Mettenius as Acrostichum commutatum. From this E. mar- ginatum is best distinguished by its stouter rhizome and more clustered fronds and its darker coloured rhizome scales, which lack the long-attenuate flexuose and curled apices and long marginal fimbriations of E. commutatum.
Christ makes no mention of Acrostichum marginatum in his monograph of Elapho- glossum, but his E. fuscopunctatum described from Yunnan I consider the same as Wallich’s plant. There are specimens at Kew and the British Museum of Henry 9158, on which Christ’s species was based, and these are, to me, indistinguishable from some Himalayan gatherings of E. marginatum (e.g. Hooker & Thomson, from Khasia in Herb. Kew; Ludlow & Sherriff 902 from Bhutan in Herb. Edin.). The scales in several other collections of E. marginatum give a conspicuously fusco- punctate appearance to the undersides of the fronds. Tagawa (loc. cit.) reduced both to forms of E. conforme.
5. Elaphoglossum commutatum (Mett.) Alderw. van Rosenb., Malayan Ferns, Supple 2.427. (2917): Acrostichum conforme sensu Blume, Enum. Pl. Jav. 2 : [275] (1828) (errore “‘ aemulum”’ in p. tor); Fl. Jav., Fil.: 23, t. 5 (1829); non Swartz.—Hook., Sp. Fil. 5: 198 (1864) — pro parte.—Hook. & Baker, Synops. Fil.: 401 (1868) pro parte. Acrostichum marginatum Fée, Mém. Fam. Foug. 2 : 31 (1845) pro parte, incl. syn. Blume. Elaphoglossum laurifolium sensu T. Moore, Index Fil. : 359 (1862) pro parte, quoad specim. ex India, Ceylon, Java; non T. Moore, op cit.: xvi (1857). Acrostichum laurifolium sensu Thw., Enum. Pl. Zeyl.: 380 (1864) ; non Thou. Acrostichum commutatum Mett. apud Kuhn in Ann. Mus. Bot. Lugd.-Bat. 4: 292 (1869). | Rhizome creeping, stipes rather close, clothed with narrow, brown scales with — fimbriate margins and long-attenuate, flexuose and twisted tips, 10 X 2 mm. Fronds ~ stalked, 15-30 (50) cm., stipe of sterile fronds variable in length, up to half as long as the lamina but often much less, scaly when young with spreading, linear, fimbriate, — dark brown scales and with smaller, appressed, stellately laciniated scales, but be- — coming smooth with age; lamina elliptic, 2-5 (6) cm. broad, apex acute, base — gradually narrowed and decurrent on the stipe, margins with a translucent, cartila- — ginous, revolute border, lower surface with scattered, minute, more or less stellately — laciniated scales, upper surface with similar scales when young, becoming glabrous; texture coriaceous. Fertile fronds on stalks normally exceeding those of the sterile ones and sometimes the stipes longer than the sterile fronds, 1-5-3-0 cm. broad. Spores 33-36 X 24-27 [. SoutH Inp1A: Bolampatty Hills, 1,500 m., Beddome (K). Nilgiri Hills, 1,200 m., Nov. 1870, Henderson (E).
ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON gt
CEYLON: Thwattes C.P. 1310 (BM; CGE; E; K; PDA). Hoolankande Peak, Matale, 10 Jan. 1862, Brodie (E). Nuwara Eliya, Freeman 368 A (BM). Kabot’s Gap, Rangala, 8 Sept. 1927, Alston 2375 (PDA). Knuckles, 1881, no collector’s name (PDA). Ramboda Pass, 1,650 m., epiphytic on tree trunk by stream in jungle, 17 Dec. 1950, Sledge 646 (BM). Hoolankande, 1,350 m., 20 Jan. 1954, Sledge roro (BM). Jungle above Le Vallon Estate, 1,500 m., g Feb. 1954, Sledge 1116 (BM). Parawella Falls, Kandapola, near Nuwara Eliya, 1,425 m., 19 March 1954, Sledge 1329 (BM). Robinson 20 (K). Central Prov., Naylor Beckett (K). 1871, Randall in Herb. R. W. Rawson 3285 (BM). R. W. Rawson 1027-1802 (BM). Henderson (E). Anderson (E).
South India (Nilgiri Hills), Ceylon, Java.
Elaphoglossum commutatum is intermediate in habit between E. marginatum and E. angulatum, with less crowded stipes than the former but with shorter internodes than the latter, in which the fronds are 1-2 cm. or more apart. It differs from E. marginatum in its rhizome scales as described under that species, and in its smaller spores. From EF. angulatum, apart from its much less widely creeping rhizome, it differs in its markedly narrower, darker brown, finely acuminate and strongly fimbriate rhizome scales. The stipes bear similar linear dark-coloured scales inter- mixed with small appressed stellate scales like those of the under surface of the blades, which spread down the stipes, though in old fronds the stipes may be practi- cally naked. In E. angulatum the broad, pale brown scales which clothe the rhizome spread up the stipe and are continued on the lower part of the midrib of the lamina, becoming narrower and more laciniated upwards and gradually merging into the minute stellate scales of the under surface. Stellate scales are absent on the stipes of E. angulatum.
Mettenius based his description (loc. cit.) on plants of Blume’s from Java and on Thwaites C.P. 1310 from Ceylon. His references to Thwaites’s numbered collection and to the description and figure of Blume’s A. conforme and his own accurate description of the rhizome scales fix the identity of the species. Van Alderwereldt van Rosenburg validly transferred the specific epithet to Elaphoglossum though the plants he had in mind were doubtfully the same as those of Mettenius. The species has been treated in most works as a synonym of FE. laurifolium (Thou.) Moore, which was described from Tristan da Cunha. Christensen (Res. Norw. Sci. Exped. Tristan da Cunha, 1937-38, 6 : 19 (1940)) expressed the view that EF. laurifolium was endemic to Tristan da Cunha. Through the kindness of Mr. James Dickson I have been able to examine good specimens of E. laurifolium collected by him on Tristan da Cunha in 1962. These and other gatherings from Tristan and from Gough Island stand well apart from Ceylon and South Indian plants in their wide-creeping habit with long internodes, their dark brown-black rhizome scales, which lack the fim- briated edges and crisped apices of E. commutatum, and their larger spores (45-48 x 30-33 ju).
E. commutatum stands between E. laurifolium and E. callifolium, the latter being a larger plant with bigger rhizome scales (up to 2 cm. long), which have entire margins. Backer and Posthumus (Varenflora voor Java: 25I (1939)) maintained E, commutatum (sub. nom. E, laurifolium) and E. callifoliwm as distinct species,
92 ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON
though they express the view that the former may be no more than a form of the latter. I have no doubt as to Ceylon and South Indian plants being distinct from E. callifolium but I am much less sure if plants from Java and elsewhere in Indonesia which have been called E. laurifolium are identical with those from India and Ceylon. Specimens so named from Java, Sumatra, Borneo and Indo-China have rhizome scales the margins of which are less irregular and they lack the frizzed tips of those of E. commutatum. They approach F. callifolium in these respects though the scales are narrower and shorter.
I suspect E. permutatum Alderw. van Rosenb. (Bull. Jard. Bot. Buitenz., ser. 2, 16 : 13 (1914)) to be synonymous with E. commutatum. There is an isotype (Matthew 696 from Mt. Sago, Sumatra) at the British Museum and this has rhizome scales with fimbriate margins and crisped apices although the description states that the scales are “‘integerrimis’”’. In its other characters the cotype does not appear to me to be significantly different from E. commutatum.
6. Elaphoglossum stelligerum (Wall. ex Baker) T. Moore ex Alston & Bonner in Candollea 15 : 216 (1956). Acrostichum neritfolium Wall., Numer. List: 2, n. 16 (1829), nom. nud. Acrostichum stelligerum Wall., op. cit.: 65, n. 2167 (1830), nom. nud. Elaphoglossum stelliger'um T. Moore, Index Fil.: 15 (1857), 368 (1862), nom. nud. Acrostichum viscosum sensu Hook., Sp. Fil. 5 : 220 (1864) pro parte ; non Swartz. Elaphoglossum viscosum sensu Bedd., Ferns. S. Ind.: 67, t. 196 (1864); Handb. Ferns
Brit. Ind.: 420 (1883) ; non J. Smith.
Acrostichum stelligerum Wall. ex Baker in Hook. & Baker, Synops. Fil, ed. 2 : 521 (1874). Acrostichum yunnanense Baker in Bull. Misc. Inf. Kew 1898 : 233 (1898). Elaphoglossum yunnanense (Baker) C. Chr. in Contrib. U.S. Nat. Herb. 26: 327 (1931).
Rhizome creeping with stipes close together, clothed with linear, attenuate, dark castaneous, glossy scales, 3-5 X 0:5 mm., with occasional fimbriations in their lower parts, elsewhere with entire or obscurely dentate margins. Fronds stalked, 20-40 cm. long, stipes of sterile fronds densely scaly with a mixture of stellate scales with slender rays and lanceolate scales with long-fimbriate margins; lamina narrowly elliptic 1-2°5 cm. broad, apex acute, base gradually narrowed and decurrent on the stipe, lower surface with abundant, loose, reddish-brown, stellate scales with long rays which diverge from the leaf surface, the midrib with similar scales intermixed, especially in the lower part, with broader-centred ones with long-fimbriate margins ; upper surface stellate-pubescent when young, becoming more or less smooth with age, the scales paler than those on the lower surface ; texture thin but firm. Fertile fronds on longer stipes than the sterile ones, lamina narrower, to I cm. broad. Spores 42-45 X 30 pM.
NEPAL : 1820, Wallich 16 (BM; E; K). Wallich ex coll. Hooker f. & Thomson 96 (E). Panapa, 1,546 m., 6 Sept. 1954, Zimmermann 1036 (BM). Between Maneb- hanjyang and the Sun Kosi, 1,500 m., 5 Nov. 1954, Zimmermann 2058 B (BM).
SIKKIM: 600 m., Beddome (BM). 2,400 m., 1868, Henderson (E). 1871, Levinge (K). Darjeeling, 1871, Hope (E). Same locality, coll. Levinge, ex Herb. Hope (E). Same locality, Oct. 1873, C. B, Clarke 21486 (K). Goke, 17 July 1880, Levinge (E).
OC
ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON 93
BuutTan: Towards Kolepani, 3 Nov. 1835, Griffith (K).
AssAM: Khasia, 1,350 m., Hooker & Thomson (E ; K). Same locality, 1,200 m., 16 Sept. 1850, Hooker & Thomson 2285 (K). Same locality, 1,200 m., 16 Nov. 1871, C. B. Clarke 14678 (K). Jowye, Jainteas, 1,200 m., 19 Oct. 1867, C. B. Clarke 5938 (K). Mariao, 1,200 m., 7 Nov. 1871, C. B. Clarke 15369 (BM; E). Bogapani, Khasia, 1,050 m., 29 Oct. 1871, C. B. Clarke 16367 (K). Same locality, 1,200 m., July 1890, G. Mann (E).
SoutH InpIA: Pulney Mts., Sept. 1836, Wight 51 (K). Same locality, 1,800 m., Beddome (BM ; K). Periya Shola, Pulneys, 21 Apr. 1898, Bourne 4971 (K). Kodai- kanal, Pulney Hills, 1,960 m., May 1937, Avoticasamy 7 (BM). Nilgiri, 1860, Beddome 4 (K). Anamallays, 1,350 m., “ F.S.Ind. f. 196°’, Beddome (BM). Same locality, 1864, Henderson (E). Shevaroy Hills 1,200 m., 1898, coll. J. W. Furrell 66 ex Herb. Hope (E). Massif du Shevaroy, ro Oct. 1939, Faucheux (BM). Honey Rock stream, Yercaud, Shevaroy Hills, 1,100 m., 25 Oct. 1962, Ghatak G1g5A (BM).
Yunnan, Indo-China.
Elaphoglossum stelligerum is readily distinguished from all other Indian species by its loose, rufous-brown, stellate scales with long rays which are not appressed to the surfaces of the stipes or fronds. The linear, dark-coloured, somewhat glossy rhizome scales are also markedly different from those of other species. It is closely related to E. petiolatum (Swartz) Urban, described from Jamaica and formerly credited with a pantropic distribution.
I see no significant difference between F. stelligerum and Baker’s Acrostichum yunnanense. Baker’s description states “ad A. stigmatolepidem, Fée, magis accedit; frondibus linearibus ad marginem paleis parvis ciliatis et paleis facialibus profunde stellatim fissis differt’’; but the comparison with E. beddomei—A. stigmatolepis sensu Baker being FE. beddome: mihi—is incomprehensible since the plant Baker had before him is manifestly far closer to, and is indeed in my opinion specifically identical with, E. stelligerum. Ching evidently dismissed it as a valid species since there is a second sheet of Henry 10370, the type gathering of A. yunnanense, in the same folder at Kew which he has labelled E. petiolatum (Sw.) Urban. Christensen upheld E. yunnanense (in Contrib. U.S. Nat. Herb. 26 : 327 (1931)) but his comments on the distinction between it and EF. petiolatum are equally applicable to EF. stelligerum, which he did not distinguish. Nor do I agree with Holttum (Fl. Malaya 2: 455 (1954)) that Malayan specimens agree better with the type of E. yunnanense than with South Indian specimens. They are more robust than either Henry’s Yunnan collection or Indian gatherings.
E. stelligerum is very closely related to E. blumeanum (Fée) J. Sm. from Malaysia and E. salicifoliwm (Willd. ex Kaulf.) Alston from the Mascarene Islands (type from Réunion) and Africa, all being formerly included in E. petiolatum Swartz, which was described from Jamaica. Malaysian plants are larger than those from India and the scales on the two surfaces of the sterile fronds are dissimilar. Indian and Mascarene plants have the scales on the two surfaces alike and differences between them are so slight that they probably represent no more than geographical variants. The names are retained however pending a much-needed revision of the whole E. petiolatum complex.
94 ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON
7. Elaphoglossum nilgiricum Krajina ex Sledge, sp. nov.
Elaphoglossum squamosum sensu Bedd., Ferns S. Ind.: 67, t. 197 (1864) ; Handb. Ferns Brit. Ind.: 420, fig. 251 (1883) excl. specim. ex Ceylon; non J. Smith.
Rhizoma breviter repens, paleis anguste lanceolatis vel linearis, pectinato-dentatis, fusco-castaneis, marginibus nigris, nitidis, vestitum. Frondes caespitosae, breviter stipitatae, 5-25 (40) cm. longae ; stipites paleis ovatis vel late ovatis, pallido-ferru- ginels in margine dentibus setiformibus praelongis, densissime obtecti; lamina anguste oblonga vel longitudo-ligulata, 1-2 cm. lata, apice obtusa, in basi sensim attenuata, in marginibus et utraque superficie copiose squamosa, infra paleis brun- neis, mollibus, lanceolatis ad late ovatis, margine dentibus setiformibus fimbriata, confertis imbricatis occultata, supra paleis similibus minus confertis, pallidioribus, aliquibus paleis in ambitu plus-minusque orbiculatis sed in margine radiis tenuibus longis instructis; textura crassiter herbacea. Forma frondium fertilium ut in sterilibus.
SouTH INpIA: Sisparah Ghat, Nilgiris, 1,500 m., “type of tab 197. FSI.”, Beddome (K, holotype). Same locality, 1,800 m., Nov. 1883, Gamble 13468 (K). Same locality, Miss Cockburn 8&1 (BM). Nilgiri, 1860, Beddome 7 (K). Pykara Falls, Nilgiri: District, 1,800 m., June 1883, Gamble 12009 (K). Nediwattan, Nilgiri Hills, 1,800 m., May 1866, Gamble ex Herb. Blanford (E). Nilgiri Hills, Henderson (E).
Endemic to Nilgiri Hills.
The type of Elaphoglossum hirtum (Swartz) C. Chr. was described from Jamaica. This name, or the synonym E. squamosum (Swartz) Urban, was formerly used in a collective sense to cover plants ranging from tropical America eastwards to southern India and Ceylon, which are the only regions in Asia whence plants have been found. Several species based on collections from Africa, Madagascar, and the Mascarene Islands have since been given independent rank. As South Indian and Ceylon plants not only differ from each other but cannot be matched satisfactorily with any Jamaican, African or Mascarene gatherings which I have examined, I have adopted the unpublished names affixed to the Kew sheets by Krajina in 1937.
Specimens from the Nilgiri Hills are very uniform and they differ consistently from Ceylon plants in the scales of the stipes being markedly broader with their margins and setiform teeth rarely dark-coloured as in E. ceylanicum, save at the base of the stipe adjacent to the rhizome. They also differ in their much more densely paleate fronds, the lower surfaces of which are hidden beneath the closely imbricating pale-brown scales.
Beddome’s illustration (Ferns S. Ind.: t. 197 (1864)), is a fair representation of the species except that the stipes are unaccountably portrayed as smooth and glabrous whereas his description states that they are “ very scaly’ and both in his own and all other gatherings they are invariably densely paleate with patent scales. The isolated, long, falcate and apparently glabrous frond depicted on the left-hand side of the plate is meaningless and, not surprisingly, is omitted from the reproduction of this plate in the Handbook.
|
ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON 95
8. Elaphoglossum ceylanicum Krajina ex Sledge, sp. nov.
Rhizoma breviter repens, paleis linearibus pectinato-dentatis fusco-castaneis vel nigris vestitum. Frondes caespitosae, breviter stipitatae, 5-25 cm. longae ; stipites paleis anguste lanceolatis, attenuatis, acutis, ferrugineis, in margine dentibus setiformis praelongis, plerumque marginibus dentibusque fuscis, obtecti; lamina oblongo-elliptica, r-2 cm. lata, in apice acuta vel acuminata, in basi sensim attenuata, in marginibus et utraque superficie paleis tenuibus brunneis; supra pallidioribus vestita, sed infra, praeter in juventute, non paleis occultata.
CEYLON: Gardner 1164 (BM; CGE on trees in dark forests at Nuwara Eliya, Sept. 1844; K, holotype). Thwaites, C.P. 3292 (BM; CGE; E; PDA). Central Province, on trees and rocks, Naylor Beckett gor (K). Peacock Hill, Pussilawa, 1870, Randall in Herb. R. W. Rawson 3284 (BM). Pundaloya, 1,500 m., Freeman 372A, 374A, 375B (BM). Nuwara Eliya, Freeman 373B (BM). Walker (K). Robinson 22 (K). Rocks by stream at Lonach near Norton Bridge, Central Province, goo m., 13 Dec. 1950, Sledge 598 (BM). Ramboda, on mossy rock by stream in jungle, 1,650 m., 17 Dec. 1950, Sledge 648 (BM). Kuda Oya, Ramboda, on rocks in shade by stream through jungle, 1,725 m., 28 Dec. 1950, Sledge 769 (BM).
Endemic.
The distinctions between this species and FE. nilgiricum are referred to under the previous species. The fully developed fronds are acute at the apex and green in colour since neither surface is covered by a continuous coating of scales as is the lower surface of the frond in E. milgivicum. Young fronds however are very scaly. In
_ both species the lamina scales vary in shape, the body of the scales being mostly
broadly to narrowly ovate in outline but some are more or less orbicular. All have the margins fringed by long setiform teeth.
E. ceylanicum seems to come closest to E. deckenw var. rufidulum (Willd. ex Kuhn) Tardieu-Blot (Not. Syst. 15: 430, t. 4 f. 6-10 (1959) =F. hirtum var. rufidulum (Willd. ex Kuhn) C. Chr. in Dansk. Bot. Ark. 7 : 170 (1932)) from Madagascar and Réunion. In that species, however, the fronds are borne on long stipes and the shape of the rhizome scales and their marginal teeth are different.
Plants from Madeira and the Azores have jet black scales on the stipes and under surface of the costae, intermixed with ferrugineous ones with and without thickened and darkened margins. Such scales are not present in E. nalgiricum or E. ceylanicum. Specimens from Madeira and the Azores are closest to E. splendens (Bory) Brackenr. from Réunion and Mauritius but are better treated as a separate species.!
1 Elaphoglossum paleaceum (Hook. & Grev.) Sledge, comb. nov. Acrostichum paleaceum Hook. & Grev., Ic. Fil. 2: t. 235, Alph. Index et Syst. Index (1831) (errore “ vestitum ’’ in expl. tab.). It would seem that Hooker and Greville realized during the preparation of the Icones Filicum that Lowe’s manuscript name Acrostichum vestitum was a later homonym. Its appearance in the text
accompanying the plate, in spite of the fact that A. paleaceum is used on the plate and in the indexes, is to be regarded as an error.
96 ELAPHOGLOSSUM IN THE INDIAN PENINSULA AND CEYLON :
9. Elaphoglossum spatulatum (Bory) T. Moore, Index Fil.: 14 (1857).—_ Bedd., Ferns S. Ind.: 71, t. 209 (err. 213) (1864). ; Acrostichum spatulatum Bory, Voy. Mers d’Afriq. 1 : 363, t. 20 f. 1 (1804).—Fée, Mém. Fam.
Foug. 2: 51, t. 14 f. 3 (1845). 1 Acrostichum acutum Fée ex Kuhn in Linnaea 36: 44 (1869). Rhizome creeping with stipes rather close together, clothed at the apex with filiform, rufous scales. Sterile fronds up to 10 cm. long, stipes about equalling the lamina in length and covered with patent, reddish, setaceous scales ; lamina lanceolate to rhomboid-lanceolate, I-1-5 cm. broad, base narrowed into the stipe, apex usually acute sometimes obtuse, margins and both surfaces covered with hair-like scales ; texture rather thickly herbaceous. Fertile fronds smaller than the sterile ones and on much longer stipes, lamina broadly oval or almost orbicular, conduplicately folded when young and often splitting at the apex when mature.
CEYLON: Thwaites C.P. 989 (BM; E; K; PDA). Adam’s Peak, Moon (BM). Nuwara Eliya, Rawson W. Rawson ro4r (BM). Same locality, Freeman 376A, 377B (BM). Sita Eliya, Hakgala, March 1885 (PDA). Hawa Eliya, 25 Aug. 1926, J.M. de Silva(PDA). Kuda Oya, Ramboda Pass, 1,700 m., inshady places on rocks by stream in jungle, 28 Dec. 1950, Sledge 771 (BM). . 1,500—1,800 m. “‘ Type of tab 209 FSI”. Thwaites in Herb. Beddome (K). 1870, Thwaites in Herb. Henderson (E). On wet rocks at about 1,800 m., Hutchison (E.). Wall (E). Robinson 23 (K). Gardner (K).
Africa (N. & S. Rhodesia, Transvaal, Natal, Cape), Madagascar, Réunion.
A very distinct species, much smaller than all other Indian and Ceylon species. Closely related plants, which have often been treated as belonging to the same species, occur in tropical America and Tristan da Cunha ; but Christensen (Res. Norw. Sci. Exped. Tristan da Cunha, 1937-38, I (6) : 20 (1940)) considered the species on that island, FE. obtusatum (Carm.) C. Chr., to be “ certainly quite different from E. spathula- tum””’.
Madame Tardieu-Blot (Not. Syst. 15: 432 (1959); in Humbert, Fl. Madag. Polypod. 2 : 49 (1960)) has erroneously transferred the Ceylon records of E. spatulatum to E. schizolepis (Baker) Christ. The type of the latter—from Madagascar—is a Kew and is quite unlike any Ceylon plant.
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FUNGI OF RECENT NEPAL EXPEDITIONS
- 3 JUL1968 a»
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BY
FRANCES L. BALFOUR-BROWNE,
Ph. 97-141; 4 Text-figures
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) BOTANY Vol. 4 No. 3
LONDON: 1968
THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become vyeady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.
In 1965 a@ separate supplementary series of longer papers was instituted, numbered serially for each Depariment.
This paper 1s Vol. 4, No. 3 of the Botany serves.
World List abbreviation: Bull. Br. Mus. nat. Hist. .(Bot.).
© Trustees of the British Museum (Natural History) 1968
TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY)
Tssued 24 June, 1968 Price Nineteen Shillings
FUNGI OF RECENT NEPAL EXPEDITIONS By FRANCES L. BALFOUR-BROWNE
Tue fungi recorded below were collected during four general botanical expeditions made in the Nepal highlands by (x) J. D. A. Stainton, W. R. Sykes and L. H. J. Williams in 1954, (2) J. D. A. Stainton in 1956, (3) A. H. Norkett in 1961-62 and (4) J. D. A. Stainton in 1962. Expeditions (x) and (4) were in Central Nepal while the other two were mainly in Eastern Nepal.
The collections were made at altitudes between 300 m. and 5,000 m. At the highest levels were Conifer and Rhododendron forests with Evergreen Oaks coming in from below, and the lower levels were characterized by rice terraces, bamboos, ferns and Castanopsis forest. Sheals & Inglis (1965) give details of the local geog- raphy and the nature of the terrain encountered in Expedition (3).
In this account references have been restricted to the authorities of the names used and to the more significant or well-known synonyms. For additional references to most of the species the revised (1960) edition of Butler & Bisby’s indispensable Fungi of India by Vasudeva should be consulted.
In some groups the delimitation of genera is very controversial and consequently the classification and the names employed are very much a personal matter of opinion with very little agreement amongst workers. This is particularly so for the Polyporaceae. During the last twenty years new systems of classification for this group have been published by Pilat (1936-42, Europe), Cunningham (1947, 1947-50, 1965, New Zealand), Corner (1932, 1953, general), Overholts (1953, America), Imazeki (1943, Japan), Donk (1933, 1960, general), Singer (1962, general), Bondart- zev (1953, Russia), Nobles (1958, Canada), Pinto-Lopez (1952, Portugal), Kotlaba & Pouzar (1957, Europe), Teixeira (1962, Brazil). These systems have been based on a variety of criteria: general anatomical and morphological structure, hymenial structure, hyphal structure, biosystematics, hyphal thickening and the presence or absence of clamps, and physiological characters. Significant as are these characters, nevertheless owing to the difference in emphasis placed upon them by different authors, considerable difficulty arises in attempting to derive a stable or consistent nomenclature. Here, therefore, the well-known, old or mainly Friesian subdivisions have been used for the Polyporaceae and the less conservative names included in the synonymy.
Recently suggested, but still tentative, relationships of certain agarics (so hitherto regarded) with polypores rather than with other agarics, or vice versa, are also not taken up here. The second edition of R. Singer’s The Agaricales in Modern Taxo- nomy (1962) should be referred to for new ideas on the classification of this group.
As regards the Clavariaceae, these have been named by Dr. E. J. H. Corner of Cambridge, and each identification for which he is responsible is indicated by his initials in parentheses.
In mountainous countries such as Nepal with an annual rainfall in some areas of 500 cm., the difficulty is to get the plants dry. The tendency is therefore to press them too enthusiastically, with the consequence that some of the agarics and
BOT. 4, 3 7
100 FUNGI OF RECENT NEPAL EXPEDITIONS
Clavariae were tissue-paper thin and their hyphal structure indiscernible. To off-set this it would have been valuable to have had some of the soft and fleshy fungi preserved in fluid, and full notes as to colour, texture and shape when fresh are always much desired. For the rest, the material was in good condition and together represents the largest collection so far from this difficult and until recently almost inaccessible region: 160 species excluding some immature and over-ripe specimens which so far have resisted identification. Previous records consist of about two dozen species collected by Sir Joseph D. Hooker over a century ago and described by Berkeley (1850, 1851, 1852, 1854) and about 70 species collected by Polunin, Sykes and Williams in 1952 and reported in a previous number of this journal (Balfour-Browne, 1955).
All the specimens cited in this paper are in the herbarium of the British Museum (Natural History).
PHYCOMYCETES ALBUGINACEAE
ALBUGO BLITI (Biv.) Kuntze, Revis. Gen. Pl. 2 : 658 (1891).—Best. Biga in Sydowia 9 : 347 (1955). Uredo bliti Biv., Stirp. Rar. Sicil. 3 : 11 (1815). ° Cystopus bliti (Biv.) De Bary in Ann. Sci. Nat., Sér. 4, Bot. 20 : 131, tab. 13 figs. 13-15 | 1863). | i ee Schwein. in Trans. Amer. Phil. Soc., New Ser., 4 : 292 (1832). Cystopus amaranthi (Schwein.) Berk. in Grevillea 3 : 58 (1874). NeEpaAL: Bhurungdi Khola, 1,600 m., on Amaranthus lividus L., 20th May, 1954, Stainton, Sykes & Williams 5342. | Distribution: Worldwide. For a recent review of the genus Albugo consult Bestagno Biba (tom. cit.: 339-58). —
PERONOSPORACEAE SCLEROSPORA GRAMINICOLA (Sacc.) Schroet. in Cohn, Krypt.-Fl. Schles. 3 (1) : 236 (1886).
Protomyces graminicola Sacc. in Nuov. Giorn. Bot. Ital. 8 : 172 (1876).
NEPAL: Bongakhani, 2,130 m., on grass, 22 Aug. 1954, Stainton, Sykes & Williams
3943. Distribution: America, Europe, Africa, India, Australia.
PYTHIACEAE PH ¥YTOPHTHORA INFESTANS (Mont.) De Bary in Journ. Roy. Agric. Soc., Ser. 2, 12 : 24) (1876). Botrytis infestans Mont. in L’Institut, Sect. 1, 13 : 313 (1845).
NEPAL: Murigurja Gad, 2,500 m., on potato, 27 July 1954, Stainton, Sykes & Williams 3654. Distribution: Worldwide.
FUNGI OF RECENT NEPAL EXPEDITIONS 101
ASCOMYCETES MORCHELLACEAE
MORCHELLA ELATA Fries, Syst. Mycol. 2 : 8 (1822).
NEPAL: Siklis, north of Pokhara, 3,000 m., on rotten tree trunk, 21 Apr. 1954, Stainton, Sykes & Williams 4956. Distribution: America, Europe, India, China, Japan, Australia.
HELVELLACEAE HELVELLA CRISPA Fries, Syst. Mycol. 2 : 14 (1822). NEPAL: Lete, 2,600 m., beneath conifers, 27 Aug. 1954, Stainton, Sykes & Williams
7501. Distribution: Worldwide; previously recorded from Nepal in 1955.
HUMARIACEAE ALEURIA AURANTIA (Fries) Fuckel in Jahrb. Nass. Ver. Naturk. 23-24 : 325 (1870).
Peziza aurantia Fries, Syst. Mycol. 2 : 49 (1822).
NEPAL: Ghar Khola, 2,130 m., 14 June 1954, Stainton, Sykes & Williams 5759. Distribution: Worldwide.
GEOGLOSSACEAE
GEOGLOSSUM AFFINE (Durand) Sacc. & Trav. in Sacc., Syll. Fung. 19 : 756 (1910).— Maas Geest. in Persoonia 4 : 23 (1965).
Gloeoglossum affine Durand in Ann. Mycol 6 : 420 (1908).
NEPAL: Annapurna Himal, Seti Khola, 3,830 m., on shady banks, 28 July 1954, Stainton, Sykes & Williams 6540. South of Gurjakhani, 4,000 m., among dwarf Rhododendron, 17 Aug. 1954, Stainton, Sykes & Williams 3879.
Distribution: United States, Himalayas.
These specimens were examined and identified by Dr. Maas Geesteranus.
MITRULA ROSEA Lloyd, Mycol. Not. 61 : 885, t. 129 fig. 1529 (1919).
NEPAL: Gosainkund, Malemchi, 2,800 m., on bare earth, 30 May 1962, Stainton 3789.
Distribution: India.
Pale rose coloured ascophores about I cm. tall with smooth or contorted caps.
SCLEROTINIACEAE RUTSTROEMIA sp. cf. FIRMA (Fries) Karst. in Bidr. Kann. Finl. Natur & Folk 19 : 108 (1871).
Peziza firma Fries, Syst. Mycol. 2 : 117 (1822). Ciboria firma (Fries) Fuckel, Symb. Mycol. : 312 (1869). Phialea firma (Fries) Gill., Champ. Fr., Discom, : rot, t. 74 fig. 2 (1883).
102 FUNGI OF RECENT NEPAL EXPEDITIONS
NEPAL: Dhankuta Province, near Mahe, 1,300 m., on dead twig, 20 Sept. 1961, Norkett 5175 B.
Distribution (of R. firma): Worldwide.
The fungus agrees well with this species except that its spores are somewhat short, 9-II xX 4-5 yw. The material, however, is barely ripe.
HY ALOSCYPHACEAE Perrotia malemchiensis Balfour-Browne, sp. nov. «(Fig. I.)
Apothecia sparsa, superficialia, sessilia, carnosa, uda pallide brunnea, sicca cinna- barina, extus villosa, 500-1,000 yw diam.; setae cylindraceae ad basim + cohaerentes apice liberae et acutae, septatae, minute granulosae, pallide ochraceae; asci clavati, recti vel curvuli, octospori, 80-110 < Q-Io “4; sporae clavatae vel fusoideae, hyalinae, 5-7 septatae, 20-30 X 4-5 “; paraphyses filiformes, septatae, hyalinae.
Ad ligna et cortices arborum frondosarum.
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Fic. 1. Perrotia malemchiensis Balfour-Browne. A, asci containing ascospores; B, paraphyses; C, ascospores; D, external hair; E, apothecium. Stainton 3768.
FUNGI OF RECENT NEPAL EXPEDITIONS 103
NEPAL: Gosainkund, Malemchi, 2,400 on a dead tree, 29 May 1962, Stainton, 3768 (holotype).
This differs from related species not only in colour and measurements but in spore septation: P. fusca Miill. & Dennis, I-septate; P. lutea (Phill.) Dennis, up to 31- septate; P. himalayensis Mill. & Dennis, 3-septate.
DERMATEACEAE Mollisia dhankutae Balfour-Browne, sp. nov. (Fig. 2.)
Apothecia superficialia, sessilia, usque ad 1-5 mm. diam., disco sordide flavido- albo; excipulo fusco-brunneo, pseudoparenchymatico; asci cylindraceo-clavati, octospori, poro jodo tincto, 80-90 x g-I0 #; ascosporae biseriatae, elongato-
1Op
Fic. 2. Mollisia dhankutae Balfour-Browne. A, asci and paraphyses; B, ascospores; C, apothecia. Norkett 7751 A.
104 FUNGI OF RECENT NEPAL EXPEDITIONS
fusoideae, uniseptatae, hyalinae, 16-20 X 3-5-4 “; paraphyses filiformes, septatae, 2 #4, supra usque 3°5 /U.
In ramis siccis Bambusae.
NEPAL: Dhankuta Province, Taplejung district, Sanghu, 1,750 m., on dead bambco stump, 19 Nov. 1961, Norkett 7751 A (holotype). Sanghu, 2,060 m., 9 Nov. 1961, Norkett 7349.
Although very close to Mollisia caesia var. andina Dennis (Kew Bull. 14 : 441 (1960)) from Venezuela, it differs in the absence of a whitish margin to the receptacle, in the slightly larger and more acutely pointed spores and in the different host. It differs also from Cenangella bambusicola Rick (Broteria 5 : 37 (1906)), which was described from living bamboo in South America, in that the paraphyses do not form an epithecium and the apothecia are not at first white, but externally very dark brown with an off-white disc.
OSTROPACEAE
VIBRISSEA TRUNCORUM Fries, Syst. Mycol. 2 : 31 (1822).
NEPAL: Rambrong, Lamjung Himal, 4,500 m., on dead roots of Rhododendron, 10 July 1954, Stainton, Sykes & Williams 6269..
Distribution: America, Europe. Apparently not previously recorded from Nepal or neighbouring countries, possibly because it is easily overlooked, being small and generally found on roots, frequently submerged in water, or on debris.
Apothecia scattered or in groups, with orange discs, 3-5 mm. diam., seated on pale stalks, blackish below, 15 xX 2-3 mm. Asci 250-300 x 6-8 mw; ascospores acicular, hyaline, multiseptate, 180-200 X 1°5 w. Paraphyses branched, filiform with spherical heads.
DIATRYPACEAE
DIATRYPE CHLOROSARCA Berk. & Broome in Journ. Linn. Soc. Lond., Bot. 14 : 123 (1873). NEPAL: Tumlingtar, Arun ravine, on dead twigs, 23 Dec. 1961, Norkett go6o. Distribution: Ceylon and India. The Nepal collection agrees with the type material described from Ceylon: spores 7-9 X 2-2°5 ww, hyaline to light brown.
HY POCREACEAE
cf. BALANSIA ANDROPOGONIS Syd. apud H. & P. Syd. & Butl. in Ann. Mycol. 9g :
395 (1911).—Patel, Gokh. & Kulk. in Indian Phytopath. 4 : 65 (1951).
NEPAL: Mayangdi Khola, 1,000 m., on inflorescence of Chrysopogon aciculatus, 4 Sept. 1954, Stainton, Sykes & Williams 4137.
Distribution (of B. andropogonis): India, Philippines.
The inflorescence of the host is so deformed and shrouded by the fungus as to give the appearance of quite a different genus of grass. Only the Ephelis stage of the fungus is represented: conidia acicular, 20-24 X I°5 y.
FUNGI OF RECENT NEPAL EXPEDITIONS 105
EPICHLOE CINEREA Berk. & Broome in Journ. Linn. Soc. Lond., Bot. 14 : 111
(1873).
NEPAL: Taplejung district, Dhankuta Province, Sanghu, 2,000 m., on an indeter- minable grass haulm, 17 Oct. 1961, Norkett 5696 C.
Distribution: Previously recorded from Ceylon. Sydow and Butler’s record from Mysore (Ann. Mycol. 9 : 394 (I911)) appears to be a different species, as Petch (Ann. Roy. Bot. Gdns. Peradeniya 7 : 88 (1920)) has noted.
PHYLLACHORACEAE CATACAUMA REPENS (Corda) Theiss. & Syd. in Ann. Mycol. 13 : 383 (1915). Sphaeria repens Corda, Icones Fung. 4 : 42, tab. 9 fig. 123 (1840). Phyllachova repens (Corda) Sacc., Syll. Fung. 2 : 597 (1883). NEPAL: Phewa Tal, 800 m., on fallen leaves of Ficus religiosa, 6 May 1954, Stainton, Sykes & Williams 5254. Distribution: Cuba, Natal, India. Several species of Catacauma have been described from Ficus spp. but this Nepal collection is quite typical of C. repens.
XYLARIACEAE HyPoxYLON MULTIFORME (Fries) Fries, Summa Veg. Scand. : 384 (1849). Sphaeria multiformis Fries, Syst. Mycol. 2 : 334 (1823).
NEPAL: Taplejung, Mewa Khola, 2,750 m., on moss-covered branch (? birch), 22 Jan. 1962, Norkett 9300. Ganesh Himal, Ankhu Khola, 2,130 m., on rotting log, 12 May 1962, Stainton 3694.
Distribution: Widespread in Northern hemisphere; previously recorded from Nepal.
HyYPOXYLON TRUNCATUM (Schwein.) J. H. Mill. in Trans. Brit. Mycol. Soc. 17 : 130
(1932); Monogr. World Sp. Hypoxylon : 95 (1961).
Sphaeria truncata Schwein. in Schr. Naturf. Ges. Leipz. 1 : 44 (1822).—Fries, Syst. Mycol. 2 : 442 (1823).
NEPAL: Bakhri Kharka, north of Pokhara, 2,000 m., 25 Apr. 1954, Stainton, Sykes & Williams 5065.
Distribution: Tropical and semitropical; America, Africa, China, Japan.
Miller (loc. cit., 1961) should be consulted for an account of this species and its considerable synonymy.
XYLOSPHAERA HYPOXYLON subsp. ADSCENDENS (Fries) Dennis in Bull. Jard. Bot. Bruxelles 31 : 124 (1961).
Sphaeria adscendens Fries in Linnaea 5 : 537 (1830).
Xylosphaera adscendens (Fries) Dennis in Kew Bull. 13 : 102 (1958).
Xylaria hypoxylon f. tropica H. & P. Syd. & Butl. in Ann. Mycol. 9 : 418 (1911).
Xylaria hypoxylon var. tropica (H. & P. Syd. & Butl.) Balf.-Browne in Bull. Brit. Mus. (Nat. Hist.), Bot. 1 : 216 (1955).
BOT. 4, 3. 7§
106 FUNGI OF RECENT NEPAL EXPEDITIONS
NEPAL: South of Gurjakhani, 3,500 m., 18 Aug. 1954, Stainton, Sykes & Williams 3897.
Distribution: South America, West Indies, Africa, India, Nepal, Indonesia.
This is the fungus previously recorded from Nepal as Xylaria hypoxylon var. tropica (Balfour-Browne, loc. cit.). Dennis (loc. cit., 1961) considers that it should be treated as a separate subspecies of Xylosphaera hypoxylon Dumort. Dennis had not seen the type specimen of Xylaria hypoxylon forma tropica and left open the question of whether that name was a synonym of Xylosphaera hypoxylon subsp. adscendens. I too have not seen the type, but I follow Dennis in considering that this is a separate subspecies.
XYLOSPHAERA MELLISSII (Berk.) Dennis in Kew Bull. 13 : 104 (1958); in Revista Biol. r : 186 (1958). Hypoxylon mellissit Berk. in Melliss, St. Helena : 379 (1875).
Xylaria mellissit (Berk.) Cooke in Grevillea 11 : 85 (1883). Xylaria arbuscula Sacc. in Michelia 1 : 249 (1878).—J. H. Mill. in Bothalia 4 : 265 (1942).
NEPAL: Dhankuta Province, Chainpur district, Tumlingtar, Sabhaya Khola, 600 m., ‘‘ growing out of a niche in rock ’’, 8 Dec. 1961, Norkett 8471 A.
Distribution: Generally common in the tropics and subtropics, but not previously reported from the Himalayan region.
The Nepal collection consists of branched stromata with long slender stalks and short cylindrical heads; ascospores are 14-16 X 4°5 MU.
XYLOSPHAERA POLYMORPHA (St. Amans) Dumort., Comment. Bot. : 92 (1822).— Dennis in Bull. Jard. Bot. Bruxelles 31 : 140 (1961).
Sphaeria polymorpha St. Amans, Fl. Agenaise : 520 (1821). Xylaria polymorpha (St. Amans) Grev., Fl. Edinensis : 355 (1824).
NEPAL: Arun Valley, Sabhaya Khola, 2,300 m., on tree trunk in forest, 7 Sept. 1956, Stainton 1607.
Distribution: Worldwide.
For a recent account and complete synonymy see Dennis (loc. cit.).
XYLOSPHAERA TELFAIRII (Berk.) Dennis in Kew Bull. 13 : 106 (1958); in Bull. Jard. Bot. Bruxelles 31 : 119 (1961).
Sphaeria telfairiit Berk. in Ann. Nat. Hist. 3 : 397 (1839). Xylaria telfaivit (Berk.) Fries in Nov. Acta Reg. Soc. Sci. Upsal., Ser. 3, 1 : 127 (1851).
NEPAL: Arun Valley, Kasuwa Khola, 2,800 m., on tree trunk in forest, 1m Sept. 1956, Stainton 1619. Arun Valley, Hatiar, 2,600 m., 21 Aug. 1956, Stainton 1408.
Distribution: Africa, Ceylon, India, Indonesia, Australia.
For a recent account of the species, Dennis (loc. cit.) should be consulted.
FUNGI OF RECENT NEPAL EXPEDITIONS 107 UsTULINA DEUSTA (Fries) Petrak in Ann. Mycol. 19 : 279 (1921).
Sphaeria deusta Fries, Syst. Mycol. 2 : 345 (1823). Ustulina vulgaris Tulasne frat., Sel. Fung. Carp. 2 : 23, tab. 3 figs. 1-6, (1863), nom. superfl.
NEPAL: Bakhri Kharka, north of Pokhara, 2,000 m., 25 April 1954, Stainton, Sykes & Williams 5056. Distribution: Worldwide.
CORONOPHORACEAE
CORONOPHORA EPISTROMA Syd. apud. Syd., Mitter & Tand. in Ann Mycol. 35 : 231
(1937).
NEPAL: Sanghu, gulley below camp, 2,000 m., on dead twig, 21 Dec. 1961, Norkett 6390.
Distribution: Originally recorded from Allahabad, and not reported since.
This species was originally described as parasitic in the stromata of Haplosporella phyllanthina and again in the present gathering it is growing within the remains of a fungal pycnidium, but the identity of the latter could not be determined, nor that of the host twigs.
PLEOSPORACEAE
FENESTELLA FENESTRATA (Berk. & Broome) Schroet. in Cohn, Krypt.-Fl. Schles. 3(2) : 435 (1897). (Fig. 3).
Valsa fenestvata Berk. & Broome in Ann. Mag. Nat. Hist., Ser. 3, 3 : 366, t. 10 fig. 14 (1859). Fenestella princeps Tulasne frat., Sel. Fung. Carp. 2 : 207 (1863), nom. superfl.
NEPAL: Dhankuta district, Chitre, on dead twig, 2,000 m., 20 Sept. 1961, Norkett 5159 A.
Distribution: N. America, Europe.
The fungus occurs on an unidentified dicotyledonous twig. The pseudothecia ‘are superficial, + stipitate, arising in small groups on an inconspicuous stroma. Asci cylindrical, 250 x 20 w approx. Ascospores 30-50 X 12-14 m“, broadly fusi- form, dark brown, the end cells being + hyaline; there are 3 main and several lesser transverse septa and several longitudinal septa.
Petrak (Sydowia 8 : 165 (1954)) describes Cucurbitaria pakistanica from Choa -Saidan Shah, on Acacia modesta. This resembles the Nepal fungus in general structure but its spores are considerably smaller, the average size being 13-22 x 8-10 py.
VENTURIACEAE REHMIODOTHIS OSBECKIAE (Berk. & Broome) Theiss. & Syd. in Ann. Mycol. 12 : 192 (1914).
Dothidea osbeckiae Berk. & Broome in Journ. Linn. Soc. Lond., Bot. 14 : 134 (1873).
108 FUNGI OF RECENT NEPAL EXPEDITIONS
Fic. 3. Fenestella fenestrata (Berk. & Broome) Schroet. A, asci containing ascospores; B, ascospore; C, stroma with pseudothecia. Norkett 5159 A.
NEPAL: Dhankuta Province, Taplejung district, Sanghu, 2,000 m., on Osbeckia sp., 4 Oct. 1961, Norkett 5696 D. Distribution: Previously recorded only from Ceylon.
POLYSTOMELLACEAE SCHNEEPIA sp. cf. DISCOIDEA (Racib.) Racib. ex Theiss. & Syd. in Ann. Mycol. 13 : 203 (I9I5). Parmularia discoidea Racib., Parasit. Algen & Pilze Java’s 2 : 21 (1900). NepAL: Dhankuta Province, Taplejung district, Sanghu, 1,600 m., on fern, Davallia sp., 7 Jan. 1962, Norkett 8552.
Distribution (of S. discoidea): Previously recorded on Polypodium longissimum from Java.
‘
FUNGI OF RECENT NEPAL EXPEDITIONS 109
The Nepal material is not quite ripe, but appears to represent the above species. Its rounded black stromata occur on the underside of the fronds and are somewhat raised in the centre. The asci and the I-septate brownish spores, which are not fully mature, measure approximately 35 x 8 w and Io X 4 ym respectively.
Several black stromatic Ascomycetes with 1-septate spores have been described on ferns. Careful comparison of type and authentic material is needed and would probably reveal that a number of names are synonyms.
BASIDIOMYCETES USTILAGINACEAE CINTRACTIA CARICIS (Pers.) Magnus in Verh. Bot. Ver. Prov. Brandenb. 37 : 79 (1806). Uredo caricis Pers., Syn. Meth. Fung. : 225 (1801). NEPAL: Lamjung Himal, 4,500 m., on inflorescence of Cyperaceae, 14 July 1954, Stainton, Sykes & Williams 6350. East of Chalike Pahar, 4,500 m., 4 Aug. 1954,
Stainton, Sykes & Williams 3743. Distribution: Worldwide.
FARYSIA OLIVACEA (DC.) H. & P. Syd. in Ann. Mycol. 17 : 41 (1919). Uredo olivacea DC., Fl. Frang. 6 : 78 (1815).
NEPAL: Bhurungdi Khola, 1,600 m., smut balls in inflorescence of Carex cruciata Wahl., 20 May 1954, Stainton, Sykes & Williams 5336. Near Lumsum, 2,300 m., 24 Oct. 1954, Stainton, Sykes & Williams 9127.
Distribution: Worldwide.
SPHACELOTHECA HYDROPIPERIS (Schumach.) de Bary, Vergl. Morph. Biol. Pilze : 187 (1884).
Uredo hydropiperis Schumach., Enum. PI. Saell. 2 : 234 (1803).
NEPAL: Southwest of Gurjakhani, 3,300 m., in flower of Polygonum campanulatum, 20 Oct. 1954, Stainton, Sykes & Williams 9104. Dhankuta Province, Taplejung district, Sanghu, 2,000 m., 25 Nov. 1961, Norkett 7532 A.
Distribution: Worldwide on Polygonum sp. Previously recorded from Nepal in
1955.
SPHACELOTHECA REILIANA (Kiihn) Clint. in Journ. Mycol. 8 : 141 (1902). Ustilago veiliana Kiihn in Rabenh., Fung. Eur. : No. 1998 (1875).
NEPAL: Murigurja Gad, 2,500 m., on Zea mays, 27 July 1954, Stainton, Sykes & Williams 3653. Distribution: Recorded in most countries where maize is grown,
110 FUNGI OF RECENT NEPAL EXPEDITIONS USTILAGO BISTORTARUM (DC.) Korn. in Hedwigia 16 : 38 (1877). Uredo bistortarum DC., Fl. Frang. 6 : 76 (1815).
NEPAL: East of Chalike Pahar, 4,830 m., on leaves of Polygonum affine, 22 Sept. 1954, Stainton, Sykes & Williams 4541.
Distribution: Worldwide.
Forming in this material elongated pustules over the under surface of the leaf lamina, and not occurring on the margins. The spores are pale purple, angular to globose, 10-16 w, minutely verrucose.
This species, common in most parts of the world, appears not to have been re- corded previously from India or neighbouring countries.
Ustilago horde: (Pers.) Lagerh. in Mitt. Badischen Bot. Ver. [2](59) : 70 (1889), non Ustilago hordei Bref. (1888).
Uvedo segetum var. hordei Pers., Syn. Meth. Fung. : 224 (1801).
NEPAL: Gurjakahni, 2,800 m., on barley inflorescence, 1 June 1954, Stainton, Sykes & Williams 2941. Tegar, north of Mustang, 6 Aug. 1954, Stainton, Sykes & Williams 2252.
Distribution: Worldwide.
The name used for this species is illegitimate, being a later homonym of Ustzlago hordet Bref. (in Nachr. Klub Landwirthe Berl. 221 : 1593 (28 June 1888)), which was based on the loose smut of barley, as a name for which it antedated U. nuda (Jensen) Kellerm. & Swingle (see below). Before the correct names of these two species can be established, it is necessary to fix the application of the name Uredo segetum Pers. (Syn. Meth. Fung. : 224 (180r)), and also to investigate the status and application of the names Uredo carbo-tritict and Uredo carbo-hordei quoted by Kellerman & Swingle (in Annu. Rep. Exp. Stat. Kansas State Agric. Coll. 2 : 262, 269, 278 (r890)) from a work by Philippar not available to me. I therefore adopt for the present the names in current use.
USTILAGO MORINAE Padw. & Azmat. Khan in Mycol. Pap., Imp. Mycol. Inst. Io : 1
(1944).
NEPAL: Above Seng Khola, 4,500 m., in flowers of Morina, 25 June 1954, Stainton, Sykes & Williams 3254.
Distribution: Recorded from Kashmir in 1944 on Morina longifolia.
Ustilago nuda (Jens.) Kellerm. & Swingle in Annu. Rep. Exp. Stat. Kansas State Agric. Coll. 2 : 277 (1890).
Ustilago segetum var. nuda Jens. in Journ. Roy. Agric. Soc., Ser. 2, 24 : 406 (1888). Ustilago nuda Rostr. in Tidsskr. Landgkon 8 : 745 (1889), nom. nud.
NEPAL: Chimgoan, north of Tukucha, 3,000 m., on barley in field, 3 June 1954, Stainton, Sykes & Williams 904. Gurjakhani, 2,830 m., on wheat, I June 1954,
FUNGI OF RECENT NEPAL EXPEDITIONS I1l
Stainton, Sykes & Williams 2940. Tamur valley, Mewa Khola, on wheat, 17 May 1956, Stainton 343.
Distribution: Worldwide.
For the status of this name, see under Ustilago horde: above.
USTILAGO PIPERI Clint. in Proc. Boston Soc. Nat. Hist. 31 : 382 (1904).—G. W. Fisch., Man. N. Amer. Smut Fungi : 291, fig. 123 D (1953).
NEPAL: Near Seng Khola, 4,600 m., on Polygonum rumicifolium, 9 Aug. 1954, Stainton, Sykes & Williams 3809.
Distribution: United States of America.
The Nepal collection agrees well with this species; it forms large pustules on the underside of the leaves; the spores are pinkish purple, spherical or subspherical, 7-9 w diam., marked with striae. This species differs from U. bistortarum in the distinctly smaller spores.
TILLETIACEAE MELANOTAENIUM SELAGINELLAE Henn. & Nyman in Warb., Monsunia I : 2 (1900).
NEPAL: Dhankuta Province, Taplejung district, Sanghu, on Selaginella sp., 27 Oct. 1961, Norkett 6551 A.
Distribution: Previously recorded from Tjibodas in Java.
In the present collection the smut occurs mostly on the basal portion of the leaves, but occasionally it covers the whole leaf surface. Spores dark brown, globose, coarsely verrucose, 15-18 «~ diameter.
MELAMPSORACEAE
COLEOSPORIUM BARCLAYENSE Bagchee in Ind. Forest Rec., New Ser., Bot. 4 : 53
(1950).
NepaL: Annapurna Himal, Mardi Khola, 4,160 m., on Senecio alatus Wall. ex DC., 20 Sept. 1954, Stainton, Sykes & Williams 8528.
Distribution: India.
The teleutospore stage is represented in this collection. Morphologically, the Coleosporium species on Senecio are difficult to separate, but, on the basis of inocula- tion experiments carried out by Bagchee (loc. cit.) on rusts on different species of Senecio, that on S. alatus would appear to be his C. barclayense.
COLEOSPORIUM CAMPANULAE (Pers.) Kickx, Fl. Crypt. Flandres 2 : 54 (1867).— Gaiim. in Beitr. Krypt.-Fl. Schweiz 12 : 113, fig. g9—100 (1959).
Uredo campanulae Pers., Syn. Meth. Fung. : 217 (1801).
NEPAL: Near Jagat, 2,800 m., on Campanula sp., 5 July 1954, Stainton, Sykes & Williams 3376. Above Lumsum, 3,700 m., on Campanula seedlings, 23 Oct. 1954, Stainton, Sykes & Williams 9122. Rambrong, Lamjung Himal, 3,000 m., on Lobelia seguinit var. doniana (Skottsb.) Wimm., 27 Oct. 1954, Stainton, Sykes & Williams
112 FUNGI OF RECENT NEPAL EXPEDITIONS
8306. Annapurna Himal, Mardi Khola, 2,600 m., on Lobelia sp., 20 Sept. 1954, Stainton, Sykes & Williams 8531. Baglung, 1000 m., on Wahlenbergia gracilis DC., 20 Apr. 1954, Stainton, Sykes & Williams 548.
Distribution: Widely represented in the Northern hemisphere and previously recorded from Nepal.
The uredospores occurred on stems as well on both leaf surfaces. A greater proportion of the spores on Wahlenbergia were oblong and elongated compared with those on Campanula. Probably two biological forms are represented. Gaiimann (loc. cit.) should be consulted for an account of at least six such forms which have been isolated from different genera and species of Campanulaceae.
COLEOSPORIUM INULAE Rabenh. in Bot. Zeit. 9 : 455 (1851).—P. & H. Syd., Monogr. Ured. 3 : 609 (1915).—Vasud. in Butl. & Bisby, Fungi of India, Rev. Ed. : 85 (1960).
Uredo inulae Kunze in Klotsch, Herb. Viv. Mycol., No. 589 (1844), nom. nud.
NEPAL: Pasgam, 1,500 m., on Inula cappa DC., 25 June 1954, Stainton, Sykes & Williams 5938.
Distribution: Europe, North Africa, Asia Minor, India.
The uredo stage is represented and this chiefly on the upper surface of the leaves, with very few pustules on the lower.
COLEOSPORIUM PEDICULARIDIS Tai in Farlowia 3 : 100 (1947).
NEPAL: Taglung, south of Tukucha, Kali Gandaki, 4,000 m., on leaves of Pedicu- laris sp., 22 Sept. 1954, Stainton, Sykes & Williams 7987. Panchasi, 2,300 m., on Pedicularis sp., 15 Oct. 1954, Stainton, Sykes & Williams 8944.
Distribution: China.
There is little doubt that the Nepal fungus represents this species. The uredosori are chiefly on the under surface of the leaves, light yellow and soon pulverulent. The uredospores are coarsely and densely echinulate, 18-30 x 12-20 mw, oblong to irregularly globose.
COLEOSPORIUM PLECTRANTHI Barcl. in Journ. Asiatic Soc. Bengal 59 (2): 89 (1890). NEPAL: Sattewati, 2,000 m., on Plectranthus sp., 12 Oct. 1954, Stainton, Sykes & Williams 8950. Distribution: India, Japan.
COLEOSPORIUM SENECIONIS (Pers.) Fries, Summa Veg. Scand. : 512 (1849). Uredo farinosa var. senecionis Pers., Syn. Meth. Fung. : 218 (1801).
NEPAL: Above Lumsum, 3,160 m., on Senecio graciliflorus DC., 10 Sept. 1954, Stainton, Sykes & Williams 4327. Dhankuta Province, Milke Danda, 3,160 m., on Senecio graciliflorus DC., 16 Nov. 1961, Norkett 7121 A.
Distribution: Worldwide.
Uredospore stage. See note under Coleosporium barclayense.
“are
Ai nc om
FUNGI OF RECENT NEPAL EXPEDITIONS 113
HYALOPSORA POLYPODII (Pers.) Magn. in Ber. Deutsch. Bot. Ges. 19 : 582 (1902). Uredo linearis var. polypodii Pers., Syn. Meth. Fung. : 217 (1801).
NepaL: Annapurna Himal, Seti Khola, 4,160 m., on Polypodium malacodon, 3 Aug. 1954, Stainton, Sykes & Williams 6603. Distribution: North America, Europe, India, Japan.
MELAMPSORA sp. cf. HIRCULI Lindr. in Acta. Soc. Fauna & Flora Fenn. 22 (3) : 19
(1902).
NepaL: Annapurna Himal, 4,150 m., on Saxifraga moorcroftiana Wall., 2 Aug. 1954, Stainton, Sykes & Williams 6587.
Distribution (of M. hirculi): Europe (Finland, Russia, Switzerland).
Although this species so far has apparently only been recorded from Europe, the Nepal collection agrees very closely with the original description of M. hirculi, but the fungus occurs not only on the under surfaces, but more frequently on the upper leaf surfaces, which become considerably discoloured and blotchy. Uredospores are globose, ovate or ellipsoid, minutely verrucose, 16-25 x 14-18 mw; paraphyses are abundant and capitate or clavate, 35-60 < 10-20 with a thick wall. The teleuto- spores are brown, oblong, 35-55 X 8-12 yu.
MILESINA sp. cf. EXIGUA Faull in Journ. Arnold Arb. 12 : 218 (1931).
NepaL: Dhankuta Province, Taplejung district, 1,950 m., on Dziplazium sp., g Jan. 1962, Norkett 8642 A.
Distribution (of M. exigua): Poland, Japan and the Siberian coast.
Uredosori only are represented; they occur in brown discoloured areas on both sides of the fronds. Many of the spores are somewhat irregularly polygonal in shape. They measure 28 x 15 mu (average) and are quite smooth.
PUCCINIACEAE
FROMMEA DUCHESNEAE (Arth.) Arth. in Bull. Torrey Bot. Club 44 : 504 (1917); in N. Amer. Fl. 7 : 731 (1925).
Kuhneola duchesneae Arth. in N. Amer. FI. 7 : 185 (1912).
Frommea obtusa [var.] duchesneae (Arth.) Arth., Man. Rusts U.S. & Canada : 93 (1934).— Gaum. in Beitr. Krypt.-Fl. Schweiz 12 : 1177 (1959).
Frommea obtusa-duchesneae Vienn.-Bourg. in Rev. Path. Vég. Entom. Agric. Fr. 33 : 38 (1954), nom. superfi.
NEPAL: Ghar Khola, 1,600 m., on Duchesnea indica, 14 June 1954, Stainton, Sykes & Williams 5767.
Distribution: Nepal, and North America and France, where the host is naturalized.
It seems that Arthur in 1934 (loc. cit.) had second thoughts about treating the rust on Duchesnea indica as a full species and, describing it as a “ less robust form ”’ compared with F. obtusa, regarded it merely as a variety of the latter species.
In the Nepal collection the uredo-sori are abundant, the uredospores measure
BOT. 4, 3. 788
114 FUNGI OF RECENT NEPAL EXPEDITIONS
9-14 x 18-20 yw, distinctly small for F. obtusa (which occurs on Potentilla sp.) but in agreement with the spores described on Duchesnea from America by Arthur and from France by Viennot-Bourgin. No other spore form was present on the Nepal plants. In the absence of firm evidence to support synonymy with F. obtusa the name F.. duchesneae is retained here. This is the first record of the rust in what is considered to be the host plant’s centre of origin.
GYMNOSPORANGIUM PADMARENSE Balf.-Browne in Bull. Brit. Mus. (Nat. Hist.) 1 :
205, fig. 2 (1955).
NEPAL: Near Gurjakhani, 3,000 m., on Juniperus wallichiana, 3 June 1954, Stainton, Sykes & Williams 2969.
Distribution: Nepal.
The material under consideration is old, the sori are broken up and the teleuto- spores have lost their pedicels.
GYMNOSPORANGIUM CUNNINGHAMIANUM Barcl. in Sci. Mem. Med. Off. Army Ind. 5 : 78, t. 1-3 (1890). NEPAL: Village south of Chakure Lekh, 6 Apr. 1952, Polunin, Sykes & Williams 1870. Distribution: India, Nepal.
PHRAGMIDIUM INCOMPLETUM Barcl. in Journ. Asiatic Soc. Bengal 59 (2) : 83 (1890).
NEPAL: Siklis, north of Pokhara, 3,000 m., on Rubus sp., 21 Apr. 1954, Stainton, Sykes & Williams 4950.
Distribution: India.
The uredospore stage only is present, and it agrees well with the original account, except that it is chiefly epiphyllous. Uredospores have a thick, 3-4 mw, epispore, which is warted. There are no paraphyses.
PHRAGMIDIUM sp. cf. NEPALENSE Barcl. in Journ. Asiatic Soc. Bengal 60 (2) : 220
(x891).
NEPAL: Bhuji Khola, 2,800 m., on Potentilla nepalensis, 16 Oct. 1954, Stainton, Sykes & Williams 9058.
Distribution (of P. nepalense) : India.
Uredosori only are represented. In the original description it was not stated whether the epispore is smooth or echinulate. Padwick & Azmatullah Khan (Mycol. Papers, Imp. Mycol. Inst. 10 : 4 (1944)) record this species and describe the uredospores as finely echinulate. This agrees with those on the present collection, and the measurements are similar.
PUCCINIA CARICIS var. HIMALAYENSIS (Barcl.) Padw. & Azmat. Khan in Mycol. Papers, Imp. Mycol. Inst. 10 : 9 (1944).
Aecidium urticae var. himalayense Barcl. in Sci. Mem. Med. Off. Army Ind. 2 : 29, t. 4, 5 figs. 8-17 (1887); in Journ. Asiatic Soc. Bengal 56 (2) : 368 (1887).
FUNGI OF RECENT NEPAL EXPEDITIONS 115
NEPAL: Jagat, 2,600 m., on Urtica sp., 5 July 1954, Stainton, Sykes & Williams 3364.
Distribution: India, Himalayas.
Cf. Padwick & Azmatullah Khan (loc. cit.) for an account of the somewhat con- fused synonymy.
PUCCINIA FAGOPYRI Barcl. in Journ. of Bot. 28 : 261 (1890). NEPAL: Dhankuta Province, Taplejung district, Sanghu, 2,000 m., on Fagopyrum,
ir Nov. 1961, Norkett 7173 A. Distribution: India.
PUCCINIA GENTIANAE (Strauss) Link in L., Sp. Pl., Ed. 4, 6 (2) : 73 (1825). Uredo gentianae Strauss in Ann. Wetter. Ges. 2 : 102, t. 11 fig. 33 (1811).
NEPAL: South of Gurjakhani, 4,000 m., on Gentiana sp., 8 June 1954, Stainton, Sykes & Williams 3071.
Distribution: Widely distributed in the northern Hemisphere.
Only the aecidial stage is represented in the Nepal material.
PUCCINIA GLUMARUM (J. K. Schmidt) Erikss. & Henn. in Medd. K. Landtbr.-Akad. Exp. 38 : 141 (1896).
Uredo glumarum J. K. Schmidt in Allgem. Okonom.-Tech. Flora 1 : 27 (1827).
NEPAL: Gurjakhani, 2,800 m., on Triticum vulgare, 1 June 1954, Stainton, Sykes & Williams 2939. Distribution: Worldwide.
PUCCINIA GRAMINIS Pers., Syn. Meth. Fung. : 228 (1801).
NEPAL: Ghasa, Kali Gandaki Valley, 2,500 m., on Berberis sp., 31 May 1954, Stainton, Sykes & Williams 5499; same locality and host, 13 June 1954, Stainton, Sykes & Williams 5750.
Distribution: Worldwide.
Some of the pustules were a bright pink and considerably swollen.
PUCCINIA LANTANAE Farl. in Proc. Amer. Acad. Arts & Sci. 18 : 83 (1883).—G. Laund. in Mycol. Papers, Commonw. Mycol. Inst. 89 : 43 (1963).
NEPAL: Sanghu, Dhankuta Province, 2,000 m., on Justicia diffusa, 15 Oct. 1961, Norkett 6124.
Distribution: North and South America, India, Indonesia, Philippines, China, Japan.
PUCCINIA LEUCOPHAEA H. & P. Syd. & Butl. in Ann. Mycol to : 258 (1912).
NeEpac: Ghar Khola, 2,000 m., on ha tae coccinea, 14 June 1954, Stainton, Sykes & Williams 5758.
116 FUNGI OF RECENT NEPAL EXPEDITIONS
Distribution: India.
Aecidial stage only and chiefly epiphyllous; only a few isolated sori on the lower surface and on the petioles. In the original account the aecia were said to be hypophyllous.
PUCCINIA POLYGONI-AVICULARIAE Pers., Syn. Meth. Fung. : 227 (1801).
Puccinia polygont Alb. & Schwein., Consp. Fung. : 132 (1805), nom. superfl—Géaum. in Beitr. Krypt.-Fl. Schweiz 12 : 775 (1959).
NepaL: Dhankuta Province, Taplejung district, Sanghu, 2,000 m., on Polygonum nepalense, 25 Nov. 1961, Norkett 7530 A.
Distribution: Worldwide.
The teleutospore stage is represented. This species and P. polygoni-amphibu are united by some mycologists but treated as separate by others. The two species are maintained here not only on biological grounds but also on account of definite small morphological distinctions, which apply not only to European plants, as generally stated, but also to American specimens as exemplified in the British Museum herbarium. In P. polygomi-ampibu, usually occurring on Polygonum amphibium, P. lapathifolium and related species, the teleutosori remain for long covered by the epidermis and form small pimply pustules, and the spores are © frequently somewhat bent and easily lose their pedicels. On the other hand, in P. — bolygoni-aviculariae, usually on Polygonum dumetorum and P. convolvulus, the , pustules rapidly burst through the epidermis of the host, leaving smooth black cushions of straight, stalked teleutospores.
In the present instance the host, Polygonum nepalense, resembles in general — appearance and texture Polygonum convolvulus, and the teleutospores and sori of the fungus agree exactly with those described for P. polygont-aviculariae.
PUCCINIA PULVERULENTA Grev., Fl. Edin. : 432 (1824).—Gaiim. in Beitr. Krypt.-F1. Schweiz I2 : 929 (1959). NEPAL: Near Dogadi Khola, 4,300 m., on Epilobium sp., 23 June 1954, Stainton,
Sykes & Williams 3226. Distribution: Worldwide.
PUCCINIA USTALIS Berk. in Hook., Journ. Bot. 6 : 207 (1854). Puccinia songarica Jacz. in Hedwigia 39 : (130), fig. 1 (1900). NEPAL: Rambrong, Lamjung Himal, 4,000 m., on Ranunculus sp., 29 June 1954,
Stainton, Sykes & Williams 6016. Distribution: India, Turkestan, Mongolia.
WOT atta ttm Leaman matte
RAVENELIA EMBLICAE Syd. apud H. & P. Syd. & Butl. in Ann. Mycol. 4 : 438 (1906). —P. & H. Syd., Monogr. Ured. 3 : 293 (1914). NEPAL: Chainpur path, Tumlingtar, 800 m., on Phyllanthus emblica, 21 Dec. 1961, — | Norkett 8100.
FUNGI OF RECENT NEPAL EXPEDITIONS 117
Distribution: Previously recorded from India and Burma.
Ravenelia phyllantht Mundk. & Thirum. (Mycol. Papers, Imp. Mycol. Inst. 16 : 24, fig. 18 (1946)) seems to be synonymous. This was described on Phyllanthus polyphyllus from Mysore.
UREDINALES—Form Genera
AECIDIUM CRINI Kalchbr. in Grevillea 11 : 26 (1882).—Mundk. & Thirum. in Mycol. Papers, Imp. Mycol. Inst. 16 : 16 (1946).
Aecidium amaryllidis H. & P. Syd. & Butl. in Ann. Mycol. 10 : 274 (1912).
NEPAL: Dana, Kali Gandaki Valley, 1,600 m., on Crinum amoenum Roxb. ex Ker-Gawl., 13 June 1954, Stainton, Sykes & Williams 5738. Distribution: India, South Africa.
AECIDIUM INFREQUENS Barcl. in Journ. Asiatic Soc. Bengal, 59 (2) : 105 (1890).
NEPAL: Nr. Dogadi Khola, 4,300 m., on Geranium collinum Steph. ex Willd., on open slopes, 23 June 1954, Stainton, Sykes & Williams 3227.
Distribution: India, Nepal, Japan.
The aecidia cover considerable areas of the lower side of the leaves, showing as light brown patches on the upper surface. Each aecidium is 250-300 w diam. The size, ornamentation, colouring of the aecidia and aecidiospores correspond exactly with those originally described for A. infrequens by Barclay on a Geranium sp. (? nepalensis) from Simla. The present fungus agrees also with the details given for A. sanguinolentum on other Geranium spp. in Finland, Russia and America by Lindroth (Bot. Notiser 1900 : 241), and he suggested this might be a synonym of Barclay’s rust.
Polunin, Sykes & Williams 4765, recorded in 1955 in this Journal (Balfour-Browne, 1955) as A. infrequens is microscopically similar to the fungus now reported, but differs in that the aecidia are grouped in orbicular patches with a small bare spot in the centre of each patch, i.e., as described by Lindroth for the less heavily in- fected specimens of his A. sanguinolentum. The host of this earlier collection, which has now been identified also as G. collinum, is more elegant and slender. Possibly the heavy rust infection is responsible for the coarser growth of the host specimen of Stainton, Sykes & Williams 3227.
A. sanguinolentum, in consequence of inoculation experiments from Geranium spp. of European origin (Lindroth, loc. cit.), has been described as a stage in life history of Puccinia polygoni-amphibii Pers. However, until experiments are made using specimens of the Nepal fungus, no certain conclusions can be drawn as to its relation- ships or alternative host plants.
In the meantime A. infrequens Barcl. is the name preferred on grounds of distri- bution and would be also on grounds of priority should this species prove to be identical with A. sanguinolentum,
118 FUNGI OF RECENT NEPAL EXPEDITIONS
AECIDIUM SCUTELLARIAE Syd. apud H. & P. Syd. & Butl. in Ann. Mycol. 5 : 504
(1907).
NepaL: Ghar Khola, 1,800 m., on Scutellaria scandens D. Don., 14 June 1954, Stainton, Sykes & Williams 5763.
Distribution: Himalayas.
A similar fungus, Aecidium scutellariae-indicae Dietel, has been described from Japan on Scutellaria indica var. japonica. This may be identical with the above — species (P. & H. Syd., Monogr. Ured. 4 : 115 (1923)). |
| |
PERIDERMIUM ORIENTALE Cooke in Ind. Forester 3 : 91 (1877) “ orientalis ”’ Aecidium complanatum Barcl. in Journ. Asiatic Soc. Bengal 59 (2) : 101 (1890).
NEPAL: Dhaibungkot, 1,600 m., on dead pine needles, 31 May 1949, Polunin o4t. Near Beni, 1,300 m., on needles of Pinus longifolia, 23 May 1954, Stainton, Sykes & Williams 2794.
Distribution: India, Nepal, Bhutan.
UREDO HYPERICI-MYSORENSIS Petch in Ann. R. Bot. Gdns. Peradeniya 6 : 213 (1917). NEPAL: Sanghu, Dhankuta Province, 1,400 m., on Hypericum sp., 12 Nov. 1961, Norkett 7088. Distribution: Ceylon, India.
AURICULARIACEAE
AURICULARIA DELICATA (Fries) Henn. apud Bresad., Henn. & Magn. in Engl., Bot. Jahrb. 17 : 492 (1893). Laschia delicata Fries in Linnaea 5 : 533 (1830). Laschia tremellosa Fries, Summa Veg. Scand. : 325 (1849). NEPAL: Ranipauwa, north of Beni, Kali Gandaki, 1,000 m., 3 Sept. 1954, Stainton, Sykes & Williams 7629. Distribution: mostly tropical; America, Africa, India, Australia, Pacific.
AURICULARIA MESENTERICA Pers., Mycol. Eur. 1 : 97 (1822).
NEPAL: Arun Valley, Num, 1,500 m., on tree trunk, 30 Aug. 1956, Stainton 1459.
Distribution: America, Europe, India, Indonesia, Australia.
The fructifications are broadly attached, many more or less disciform, and they therefore superficially resemble A. peltata Lloyd. However the hairs are much ~ longer, up to 500 yp.
f
»
AURICULARIA POLYTRICHA (Mont.) Sacc. in Atti R. Ist Veneto, Ser. 6, 3 : 722 (1885).
Exidia polytricha Mont. in Bélang., Voy. aux Indes-Or. 2 : 154 (1834). Hirneola polytricha (Mont.) Fries in K. Vet.-Akad. Handl. 1848 (1) : 146 (1849).
NEPAL: Tamrang Khola, 2,300 m., on branch of tree, 21 Nov. 1961, Norkett 7889. — Distribution: Worldwide,
FUNGI OF RECENT NEPAL EXPEDITIONS 119
TREMELLACEAE
GUEPINIA HELVELLOIDES (Fries) Fries, Elenchus Fung. 2 : 31 (1828).
Tremella helvelloides Fries, Syst. Mycol. 2 : 211 (1822).
Phlogiotis helvelloides (Fries) Martin in Amer. Journ. Bot. 23 : 628 (1936). Tvemella rufa Pers., Mycol Eur. 1 (1) : 103 (1822).
Gyrocephalus rufus (Pers.) Bref. in Unters. Gesammtgeb. Mykol. 7 : 131 (1888).
NEPAL: Taglung, Kali Gandaki, 3,500 m., 22 Sept. 1954, Stainton, Sykes & Williams
7990. Distribution: North America, Europe, China, India.
TREMELLA MESENTERICA Fries, Syst. Mycol. 2 : 214 (1822).
NEPAL: Dhankuta Province, near Mahe, 1,300 m., on dead tree, 20 Sept. 1961, Norkett 5175 D. Distribution: Worldwide.
EXOBASIDIACEAE
EXOBASIDIUM sp.
NEPAL: Lete, Kali Gandaki Valley, 3,800 m., on Rhododendron campanulatum, 4 June 1954, Stainton, Sykes & Williams 5607. Above Sauwala Khola, 3,800 m., on Rhododendron lepidotum, 15 Sept. 1954, Stainton, Sykes & Williams 4430. Near Lumsum, 2,300 m., on Rhododendron seedlings, 24 Oct. 1954, Stainton, Sykes & Williams 9132.
Specific identification could not be made as the collections in all cases were very over-ripe. Several species of Exobasidium have been described on Rhododendron. References to the literature on Exobasidium can be found in Sundstrém (Phytopath. Zeitschr. 40 : 213-17 (1960)) and in McNabb (Trans. R. Soc. N.Z., Bot. 1 : 267
(1962)). AGARICACEAE ARMILLARIA MELLEA (Fries) Kummer, Fir. Pilzk. : 134 (1871).
Agaricus melleus Fries, Syst. Mycol 1 : 30 (1821).
NeEpAL: Arun Valley, Kasuwa Khola, on tree trunk in forest, 11 Sept. 1956, Stainton 1618. Distribution: Worldwide.
CLITOCYBE TABESCENS (Fries) Bresad., Fung. Trident. 2 : 84, t. 197 (1900). Agaricus tabescens Fries, Hymenomyc. Eur., Ed. 2 : 111 (1874).
NEPAL: Lete, Kali Gandaki Valley, 2,600 m., in leaf mould at base of Pinus chylla, 3 June 1954, Stainton, Sykes & Williams 5551. Distribution: Worldwide.
120 FUNGI OF RECENT NEPAL EXPEDITIONS
CoPRINUS COMATUS (Fries) Gray, Nat. Arrang. Brit. Pl. 1 : 633 (1821). Agaricus comatus Fries, Syst. Mycol. 1 : 307 (1821).
NEPAL: Chimgaon (north of Tukucha), Kali Gandaki, 4,500 m., 17 July 1954, Stainton, Sykes & Williams 1846. Distribution: Worldwide.
COPRINUS DISSEMINATUS (Fries) Gray, Nat. Arrang. Brit. Pl. 1 : 634 (1821).
Agaricus disseminatus Fries, Syst. Mycol. 1 : 305 (1821). Psathyrella disseminata (Fries) Quél. in Mém. Soc. Emul. Montbéliard, Ser. 2, 5 : 153 (1872) (reimpr. quam Champ. Jura Vosg. : 123 (1872)). NEPAL: Arun Valley, Hatiar, 2,300 m., on fallen tree in forest, 20 Aug. 1956, Stainton 1394. Distribution: Worldwide.
CREPIDOTUS MOLLIS (Fries) Staude in Festg. Mitgl. XIX Versamml. deutsch. Land- und Forstwirthe Coburg : 71 (1857) (reimpr. quam Schwaémme Mitteldeutschl. :
71 (1858)). Agaricus mollis Fries, Syst. Mycol. 1 : 274 (1821). NEPAL: Arun Valley, Kasuwa Khola, 2,800 m., on tree trunk in forest, 11 Sept. 1956, Stainton 1617. Distribution: America, Europe, China, Japan, Australia. Very badly crushed in pressing but the layers of parallel and gelatinous hyphae were readily observed; spores 9 X 5 “, smooth.
GOMPHUS FLOCCcosuS (Schwein.) Sing. in Lloydia 8 : r40 (1945). Cantharellus floccosus Schwein. in Trans. Amer. Phil. Soc., New Ser. 43 253. (2832).
NEPAL: Above Sauwala Khola, 3,300 m., on earth bank in Quercus forest, 13 Sept. 1954, Stainton, Sykes & Williams 4375. Distribution: Recorded from North America, China, Japan, as well as from Nepal
(1955).
LACCARIA LACCATA (Fries) Cooke in Grevillea 12 : 70 (1884). Agaricus laccatus Fries, Syst. Mycol. 1 : 106 (1821).
NepaL: Arun Valley, Barun Khola, 4,000 m., in short grass, pinkish brown, 15 Sept. 1956, Stainton 1662. Distribution: Worldwide.
LACTARIUS PUBESCENS (Krombh.) Fries, Epicrisis Syst. Mycol. : 335 (1838).
Agaricus pubescens Krombh., Naturg. Abbild. & Beschreib. Essb., Schadl. & Verdacht.
Schwamme 2 : 24, t. 13 figs. 1-14 (1832).
FUNGI OF RECENT NEPAL EXPEDITIONS 121
Nepav: Arun Valley, Barun Khola, 4,000 m., in short grass, 15 Sept. 1956, Stainton 1660.
Distribution: Apparently worldwide, but it is uncertain how many records under the name of the coarser L. torminosus (Fries) Gray, from which many mycologists have not separated this species, refer to it. Very few species of Lactarius have as yet been recorded from India or any of the neighbouring countries.
LEPIOTA ERMINEA (Fries) Gill., Hyménomycétes : 59 (1874).
Agaricus ermineus Fries, Syst. Mycol. 1 : 22 (1821).
NEPAL: Mathand, near Pokhara, 1,120 m., on shady bank, ‘“‘ white except top of cap which is brown ’’, 22 June 1954, Stainton, Sykes & Williams 5852. Distribution: Europe, India, Australia.
MARASMIUS CRINIS-EQUI F. von Muell. ex Kalchbr. in Grevillea 8 : 153 (1880).
Marasmius equicrinis F. von Muell. ex Berk. in Journ. Linn. Soc. Lond., Bot. 18 : 383 (1881), nom. superfi. NEPAL: Murigurja Gad, 2,500 m., on dead vegetation near ravine track, 27 July 1954, Stainton, Sykes & Williams 3647. Distribution: America, India, Ceylon, Philippines, Australia.
Panus polychrous (Lév.) Singer ex Balfour-Browne, comb. nov.
Lentinus polychrous Lév. in Ann. Sci. Nat., Ser. 3, Bot. 2 : 175 (1844). Lentinus vellereus Berk. & Curt. in Journ. Linn. Soc. Lond., Bot. 10 : 301 (1868). Lentinus kurzianus Currey in Trans. Linn. Soc. Lond., Ser. 2, Bot. I : 120, t. 20 fig. I1 (1876). Panus polychrous Sing., Agaricales in Modern Taxonomy, Ed. 2 : 172 (1962), nom. invalid. NEPAL: Tumlingtar, Sabhaya Khola, 600 m., on dead tree, 20 Dec. 1961, Norkett S108. Distribution: Cuba, India, Nepal, Ceylon, Philippines, Australia. A few additional synonyms are given by Singer (loc. cit.).
PANUS TIGRINUS (Fries) Sing. in Lilloa 22 : 275 (1951). Agaricus tigrinus Fries, Syst. Mycol. 1 : 176 (1821). Lentinus tigrinus (Fries) Fries, Epicrisis Syst. Mycol. : 389 (1838).
NEPAL: Midam Khola, Chisankhu, 650 m., on dead tree trunk, 4 May 1954, Stainton, Sykes & Williams 5214. Distribution: Worldwide.
PHOLIOTA SQUARROSA (Fries) Kummer, Fiihr. Pilzk. : 84 (1871). Agaricus squarrosus Fries, Syst. Mycol. 1 : 243 (1821).
NEPAL: Chimgaon (north of Tukucha), Kali Gandaki, 3,500 m., in forest, at base of conifer, 14 Sept. 1954, Stainton, Sykes & Williams 7830. Distribution: North America, Europe, Japan.
122 FUNGI OF RECENT NEPAL EXPEDITIONS
SCHIZOPHYLLUM COMMUNE Fries, Syst. Mycol 1 : 330 (1821).
NEPAL: Ranipauwa (north of Beni), Kali Gandaki, 1,000 m., on tree, 12 Sept. 1954, Stainton, Sykes & Williams 7818. Tumlingtar, Sabhaya River, Chainpur district, 600 m., 12 Dec. 1961, Norkett 8480. Hinwan Khola, Chainpur, 660 m., on dead stick, 21 Dec. 1961, Norkett 9024.
Distribution: Worldwide.
HYDNACEAE HERICIUM ERINACEUS (Fries) Pers., Mycol. Eur. 2 : 153 (1825). Hydnum erinaceus Fries, Syst. Mycol. 1 : 407 (1821).
NEPAL: Above Sauwala Khola, 3,300 m., on Quercus in thick forest, 15 Sept. 1954, Stainton, Sykes & Williams 4415. Chimgaon, Kali Gandaki, 3,300 m., 14 Sept. 1954, Stainton, Sykes & Williams 782r.
Distribution: America, Europe, India, Japan, China.
HYDNELLUM ZONATUM forma VESPERTILIO (Berk.) Coker & Beers, Stipitate Hydnums of Eastern U.S. : 80 (1951).
Hydum vespertilio Berk. in Hook., Journ. Bot. 6 : 167 (1854).
NEPAL: Taglung, Kali Gandaki, 3,500 m., 22 Sept. 1954, Stainton, Sykes & Williams
799I. Distribution: America, Europe, India.
POLYPORACEAE
AMAURODERMA RUGOSUM (Blume & Nees) Torrend in Broteria, Ser. Bot. 18 : 127 (1920). . Polyporus rugosus Blume & Nees in Nov. Act. Phys.-Med. Acad. Caes. Leop.-Car. 13 : 21, t, 7 (1826): NEPAL: Dhankuta Province, below Sanghu, on roots of bamboo, 1,800 m., 27
Feb. 1962, Norkett 10233. Distribution: Mostly tropical; Madagascar, India, Ceylon, Java, Philippines.
GANODERMA APPLANATUM (Pers.) Patouill. in Bull. Soc. Mycol Fr. 5 : 67 (1889).— Humphr. & Lewis in Philipp. Journ. Sci. 45 : 514 (1931).
Polyporus fomentarius var. applanatus Pers., Mycol. Eur. 2 : 80 (1825). Polyporus applanatus (Pers.) Wallr., Fl. Crypt. Germ. 2 : 591 (1833).
NEPAL: Bakhri Kharka, north of Pokhara, 1,800 m., on rotten tree trunk, 24
April 1954, Stainton, Sykes & Williams 5060. Taglung, Kali Gandaki, 3,300 m.,
19 Oct. 1954, Stainton, Sykes & Williams 8198. Dharan Bazar, Terai forest, south ~
of Gopa Gurkha Camp, 250 m., on old tree, 27 Feb. 1962, Norkett 10234. Distribution: Worldwide.
FUNGI OF RECENT NEPAL EXPEDITIONS 123 GANODERMA LUCIDUM (Fries) Karst. in Rev. Mycol. 3 (9) : 17 (1881). Polyporus lucidus Fries, Syst. Mycol. 1 : 353 (1821).
NEpAL: Bakhri Kharka, north of Pokhara, 2,000 m., on rotten tree trunk, 25 April 1954, Stainton, Sykes & Williams 5075. Distribution: Worldwide
FoMEs PECTINATUS (Klotzsch) Gill., Hyménomycetes : 686 (1874). Polyporus pectinatus Klotzsch in Linnaea 8 : 485 (1833).
NEPAL: Between Bakhri Kharka and Rambrong, 2,300 m., on rotten tree trunk, 26 Apr. 1954, Stainton, Sykes & Williams 5081. Distribution: America, Europe, India, Australia, Philippines.
FOMES MARGINATUS (Fries) Gill., Hyménomycétes : 683 (1874). Polyporus marginatus Fries, Syst. Mycol. 1 : 372 (1821).
NEPAL: Taglung, Kali Gandaki, 3,300 m., on forest tree, 11 July 1954, Stainton, Sykes & Williams 1751. Also at 3,500 m., 22 Sept. 1954, Stainton, Sykes & Williams
7992. Distribution: America, Europe, India, Nepal, China, Japan.
POLYPORUS ARCULARIUS Fries, Syst. Mycol. I : 342 (1821).
Polyporellus arcularius (Fries) Pilat in Kav. & Pilat, Atlas Champ. 3 : 75, t. 30-31, fig. 18 (1936).
var. ARCULARIUS.
NEPAL: Midam Khola, Chisankhu, 660 m., 4 May 1954, Stainton, Sykes & Williams 5210. Kabre, Kali Gandaki, 2,000 m., 13 June 1954, Stainton, Sykes & Williams 5742. Dhankuta Province, Chainpur district, Tumlingtar, 600 m., on dead trunk, 13 Dec. 1961, Norkett 8109 B; 8815 A, and on 14 Dec. 1961, Norkett 8107 B.
Distribution: Worldwide; previously recorded from Nepal.
var. STRIGOSUS Bourd. & Galz., Hymenomye. Fr. : 532 (1928).
NEPAL: Chipli, North of Pokhara, 2,600 m., on rotten tree trunk, 18 Apr. 1954, Stainton, Sykes & Williams 4882.
Distribution: Worldwide.
Distinguished by its marginal hairs.
POLYPORUS PARGAMENUS Fries, Epicrisis Syst. Mycol. : 480 (1838).—Overh., Polyp. U.S., Alaska & Canada : 336 (1953).
NEPAL: Sanghu, Milke Danda Forest, 2,900 m., on old dead tree, 16 Nov. 1961, Norkett 7129.
124 FUNGI OF RECENT NEPAL EXPEDITIONS
Distribution: Widespread in temperate and tropical regions.
The material is in good condition but not sporing. There is much confusion over the use of this name, P. biformis Klotsch and P. cervinus Fries. It is hoped to make a more critical study of the problem shortly. Meanwhile Overholt’s interpretation of P. pargamenus is adopted.
POLYPORUS CORRUGATUS Pers. apud Gaud. in Freyc., Voy. aut. Monde Uranie & Physicienne, Bot. : 172 (1826). Earliella corrugata (Pers.) Murrill in Bull. Torrey Bot. Cl. 34 : 468 (1907).
Polystictus persoonii Cooke in Grevillea 14 : 85 (1886). Daedalea sanguinea Klotsch in Linnaea 8 : 481 (1833).
NEPAL: Chainpur district, Tumlingtar, Dhankuta Province, 600 m., on dead tree, 13 Dec. 1961, Norkett 8109 E.
Distribution: West Indies, India, Nepal, East Indies and throughout most of the tropics. AE
POLYPORUS PICIPES Fries, Epicrisis Syst. Mycol. : 440 (1838).—Overh., Polyp. U.S., Alaska & Canada : 262 (1953).
Polyporellus picipes (Fries) Karst. in Bidr. Kann. Finl. Natur. & Folk 37 : 31 (1882).— Pilat in Kav. & Pilat, Atlas Champ. 3 : 99, t. 44 fig. 1-3, t. 46 fig. b, fig. 24, p. 105 fig. B (1937).
NEPAL: Ghar Khola, 3,100 m., on dead trunk, 15 June 1954, Stainton, Sykes & Willams 5769. Arun Valley, Kasuwa Khola, 2,800 m., 11 Sept. 1956, Stainton 1624.
Distribution: Worldwide.
POLYPORUS SQUAMOSUS Fries, Syst. Mycol. I : 343 (1821).
NEPAL: Rambrong ridge, north of Pokhara, 3,300 m., on rotten tree trunk, 27 Apr. 1954, Stainton, Sykes & Williams 5103. Distribution: Worldwide.
POLYPORUS SULPHUREUS Fries, Syst. Mycol. 1 : 357 (1821).
Grifola sulphurea (Fries) Pilat in Beih. Bot. Centralbl. 52 (B) : 39 (1934). Laetiporus sulphureus (Fries) Bondartz. & Sing. in Ann. Mycol. 39 : 51 (1941).
NEPAL: Chimgaon, Kali Gandaki, 3,500 m., 14 Sept. 1954, Stainton, Sykes & Williams 7828. Arun Valley, Kasuwa Khola, 2,800 m., on tree in forest, 11 Sept. 1956, Stainton r62r.
Distribution: Worldwide; previously recorded from Nepal.
Ce ee ee “4 ay
POLYPORUS ZONALIS Berk. in Ann. & Mag. Nat. Hist. 10 : 375, t. ro fig. 5 (1843). 7
NEPAL: Karelung, Madi Khola, 660 m., on rotten branch, 23 June 1954, Stainton, Sykes & Williams 5911.
FUNGI OF RECENT NEPAL EXPEDITIONS 125
Distribution: tropical and semi-tropical; Central and South America, Cuba, India, Indonesia, China, Australia. Resembles the type but is a little thicker. Spores globose and no cystidia.
POLYSTICTUS AFFINIS (Blume & Nees) Fries in Nov. Act. Soc. Sci. Upsal., ser. 3, I: 75 (1851).
Polyporus affinis Blume & Nees in Nov. Act. Phys.-Med. Acad. Caes. Leop.-Car. 13 : 18, t. 4 (1826). Microporus affinis (Nees) Kuntze, Revis. Gen. Pl. 3 (2) : 495 (1898).
NepaL: Arun Valley, Sashaya Khola, 660 m., on tree trunk in forest, 4 Sept. 1956, Stainton 1578. Dharan Bazar, Terai Forest, south of Gopa Gurkha Camp, 250 m., 27 Feb. 1962, Norkett 10224.
Distribution: widespread in tropical and subtropical regions; previously recorded from Nepal.
POLYSTICTUS CINNAMOMEUS (Gray) Sacc., Syll. Fung. 6 : 210 (1888).
Strilia cinnamomea Gray, Nat. Arrang. Brit. Pl. 1 : 645 (1821).
Polyporus cinnamomeus (Gray) Fries, Epicrisis Syst. Mycol. : 468 (1838).—Overh., Polyp. U.S., Alaska & Canada : 386 (1953).
Coltricia cinnamomea (Gray) Murrill in Bull. Torrey Bot. Cl. 31: 343 (1904).
NEPAL: Taglung, Kali Gandaki, 3,500 m., in wood, 22 Sept. 1954, Stainton, Sykes & Williams 7991.
Distribution: Worldwide; previously recorded from Nepal.
Very close to P. perennis but distinguished by its more uniform and silkier cap.
POLYSTICTUS HIRSUTUS (Fries) Fries in Nov. Act. Soc. Sci. Upsal., ser. 3, I : 86 (1851).
Polyporus hirsutus Fries, Syst. Mycol. 1 : 367 (1821). Coriolus hirsutus (Fries) Quél., Enchir. Fung. : 175 (1886).—Bourd. & Galz., Hyménomyc. Fr. : 561 (1928).
NeEpac: Dhankuta Province, Milke Danda Forest, 260 m., 29 Nov. 1961, Norkett 8307 A. Distribution: Worldwide.
POLYSTICTUS PERULA (Fries) Fries in Nov. Act. Soc. Sci. Upsal., ser. 3, I : 73 (1851).
Polyporus perula Fries, Syst. Mycol. 1 : 349 (1821).
Polyporus xanthopus Fries, Syst. Mycol. 1 : 350 (1821).
Polystictus xanthopus (Fries) Fries in Nov. Act. Soc. Sci. Upsal., ser. 3, 1 : 74 (1851). Microporus perula (Fries) Hariot in Bull. Soc. Mycol. Fr. 7 : 206 (1891).
NEPAL: Rupakot Tal, 800 m., on rotten branch, 5 May 1954, Stainton, Sykes & Williams 5233. Arun Valley, Hinwan Khola, 800 m., on rotten log, 4 Sept. 1956,
126 FUNGI OF RECENT NEPAL EXPEDITIONS
Stainton 1533. Sanghu, 3,000 m., on tree stump, 2 Oct. 1961, Norkett 5540. Chain- pur district, Tumlingtar, 600 m., 13 Dec. 1961, Norkett 8109 C.
Distribution: Widespread in tropical and sub-tropical areas; previously recorded from Nepal.
POLYSTICTUS SANGUINEUS (Fries) Fries in Nov. Act. Soc. Sci. Upsal., ser. 3, I : 75
(1851).
Polyporus sanguineus Fries, Syst. Mycol. 1 : 371 (1821). Pycnoporus sanguineus (Fries) Murrill in Bull. Torrey Bot. Cl. 31 : 421 (1904).
NEPAL: Midam Khola, Chisankhu, 660 m., 4 May 1954, Stainton, Sykes & Williams 5218. Midam Khola, Karelung, 600 m., 23 June 1954, Stainton, Sykes & Williams 5910. Kusma, 660 m., 2 Nov. 1954, Stainton, Sykes & Williams 9270. Sanghu, 3,000 m., 6 Oct. 1961, Norkett 5709.
Distribution: Mostly tropical and sub- tropical.
The Nepal material is thin and smooth and conforms with P. sanguineus. P. cinnabarinus Fries, at one time considered to be a synonym, has been shown to be distinct on the basis of cultural interfertility tests (McKay in Mycologia, 51 : 465-
73 (1959)).
POLYSTICTUS SUBAFFINIS Lloyd, Mycol. Not. 40 : 550, fig. 755 (1916).
NEPAL: Surauti Khola, 660 m., on dead bamboo, 12 Aug. 1954, Stainton, Sykes & Williams 6869.
Distribution: Japan, Java, Madagascar.
The present collection appears to agree completely with Lloyd’s species as he figured it from Umemura’s Japanese specimen, but it is doubtful whether this species is distinct from Polystictus affinis (Fries) Fries.
POLYSTICTUS TABACINUS (Mont.) Sacc., Syll. Fung 6 : 280 (1888). Polyporus tabacinus Mont. in Ann. Sci. Nat., Ser. 2, Bot. 3 : 349 (1835).
NEPAL: Siklis, north of Pokhara, 2,100 m., on rotten tree trunk, 22 Apr. 1954, Stainton, Sykes & Williams 4974. Gurjagaon, 3,000 m., on dead tree, 25 Sept. 1961, Norkett 5481. Milke Danda Forest, 3,000 m., on old dead tree, 16 Nov. 1961, Norkett 7127.
Distribution: South America, Africa, India, East Indies, China, Australasia.
g-I0 pores per mm., setae subulate, dark brown. This species differs from
P. iodinus in having smaller pores.
POLYSTICTUS TEPHROLEUCUS (Berk.) Sacc., Syll. Fung. 6 : 275 (1888). Trametes tephroleuca Berk. in Hook., Journ. Bot. 6 : 165 (1854). Coriolus tephroleucus (Berk.) Bondartz., Trutov. Ghrib. Evr. Chasti $.S.S.R. & Kavk. :
492, fig. 126 (1953). Coriolus favoliporus Pilat in Bull. Soc. Mycol. Fr. 52 : 313, t. 3 figs. 3-4 (1937). Trametes favolipora (Pilat) Pilat in nam & Pilat, Atlas Champ. 3 : 267, t. 182, fig. 105
(1939).
al Tie,
FUNGI OF RECENT NEPAL EXPEDITIONS 127
NEPAL: Lulo Khola, 16 Sept. 1952, Polunin, Sykes & Williams 3448. Near Lumsum, 2,300 m., on dead tree stump, 24 Oct. 1954, Stainton, Sykes & Williams 9137.
Distribution: Asia: Kazakstan, India and East Nepal.
Fine specimens but the pores in Stainton, Sykes & Wulliams 9137 are mostly discoloured owing to a mycelial growth over the hymenium.
POLYSTICTUS VERSATILIS (Berk.) Fries in Nov. Act. Soc. Sci. Upsal., Ser. 3, 1 : 92 (1851).
Trametes versatilis Berk. in Lond. Journ. Bot. 1 : 150 (1842).
Polyporus versatilis (Berk.) Romell in Bih. K. Svensk. Vet. -Akad. Handl. 26, (3, 16) : 35 (1901).—Lloyd, Mycol. Not. 50: 703, figs. 1049-50 (1917).—Overh., Polyp. U.S., Alaska & Canada: 325 (1953).
NEPAL: Ghar Khola, 2,600 m., 3 May 1954, Stainton, Sykes & Williams 5444. Maikot, 2,600 m., on stump, 4 July 1954, Stainton, Sykes & Williams 3363. Tum- lingtar, Chainpur, 600 m., on old tree in ravine, 9 Dec. 1961, Norkett 8693. Dharan Bazar, Terai forest, south of Gopa Gurkha Camp, 230 m., 27 Feb. 1962, Norkett 10232.
Distribution: America, Madagascar, India, Malaya, Indonesia, Japan, China.
POLYSTICTUS VERSICOLOR (Fries) Fries in Nov. Act. Soc. Sci. Upsal., Ser. 3, 1 : 86 (1851).
Polyporus versicolor Fries, Syst. Mycol. 1 : 368 (1821). Coriolus versicolor (Fries) Quél., Enchir. Fung. : 175 (1886).
NEPAL: Ranipauwa, Kali Gandaki, 1,000 m., 3 Sept. 1954, Stainton, Sykes & Williams 7637. Lete, Kali Gandaki, 2,800 m., 17 Sept. 1954, Stainton, Sykes © Williams 7893. Bakhri Kharka, Pokhara, 2,000 m., 25 Apr. 1954, Stainton, Sykes & Williams 5073. Arun Valley, Hatiar, 2,600 m., 21 Aug. 1956, Stainton r4z2. Taplejung, above Sanghu, Milke Danda Forest, 3,000 m., 16 Nov. 1961, Norkett 97124 A. Mewa Khola, 2,700 m., 23 Jan. 1962, Norkett 9345 and 9345 A.
Distribution: Worldwide; previously reported from Nepal.
DAEDALEA UNICOLOR Fries, Syst. Mycol. 1 : 336 (1821).—Overh., Polyp. U.S., Alaska & Canada : 125 (1953).
Coriolus unicolor (Fries) Patouill., Ess. Tax. Fam. & Genr. Hyménomye. : 94 (1900).
NEPAL: Sanghu, 3,000 m., 1 Nov. 1961, Norkett 6728. Distribution: America, Europe, North Africa, China, Australia.
TRAMETES CERVINA (Schwein.) Bresad. in Ann. Mycol. 1 : 81 (1903).
Boletus cervinus Schwein. in Schrift. Naturf. Ges. Leipz. 1 : 96 (1822). Coriolus cervinus (Schwein.) Bondartz., Trutov. Ghrib. Evr. Chasti S.S.S.R. & Kavk. :
493, fig. 127 (1953). Polyporus biformis sensu Berk. in Ann. & Mag. Nat. Hist. 3 : 392 (1839), non Klotzsch.
128 FUNGI OF RECENT NEPAL EXPEDITIONS
NEPAL: Ulleri, north of Kusma, Kali Gandaki, 2,600 m., on tree in wood, 1 Nov. 1954, Stainton, Sykes & Williams 8275.
Distribution: Europe, Russia, India, Ceylon, China, Australia.
For nomenclature of this fungus see note under Polyporus pargamenus (p. 123).
TRAMETES GIBBOSA (Fries) Fries, Epicrisis Syst. Mycol. : 492 (1838).
Daedalea gibbosa Fries, Syst. Mycol. 1 : 338 (1821). Pseudotrametes gibbosa (Fries) Bondartz. & Sing. in Ann. Mycol. 39 : 60 (1941).
NEPAL: Siklis, north of Pokhara, 21 Apr. 1954, Stainton, Sykes & Williams 4957. Distribution: Europe, Africa, India, China. .
LENZITES BETULINA (Fries) Fries, Epicrisis Syst. Mycol. : 405 (1838).
Daedalea betulina Fries, Syst. Mycol. 1 : 333 (1821). Trametes betulina (Fries) Pilat in Kav. & Pilat, Atlas Champ. Eur. 3 : 327, t. 220, fig. 142
(1940).
NeEpAL: Arun Valley, Hatiar, 2,600 m., on tree trunk, 21 Aug. 1956, Stainton r4r2. Siklis, north of Pokhara, 2,500 m., on rotten tree trunk, 19 Apr. 1954, Stainton, Sykes & Williams 4926.
Distribution: Worldwide.
LENZITES PALISOTII (Fries) Fries, Epicrisis Syst. Mycol. : 404 (1838).
Daedalea palisotii Fries, Syst. Mycol. 1 : 335 (1821) “‘ Palisoti’’. Daedalea applanata Klotzsch in Linnaea 8 : 481 (1833).
Lenzites vepanda Fries, Epicrisis Syst. Mycol. : 404 (1838).
Lenzites applanata (Klotzsch) Fries, Epicrisis Syst. Mycol. : 404 (1838).
NeEpaAL: Arun Valley, Sabhaya Khola, 800 m., on tree trunk, 3 Sept. 1956, Stainton 1576. Dhankuta Province, Chainpur district, Tumlingtar, 600 m., on dead branch, 13 Dec. 1961, Norkett 8109 A.
Distribution: Widespread, especially in the southern hemisphere.
LENZITES SUBFERRUGINEA Berk. in Hook., Journ. Bot. 6 : 134 (1854). .
Gloeophyllum subferrugineum (Berk.) Bondartz., Trutov. Ghrib. Evr. Chasti $.S.S.R. & Kavk. : 50 (1953).
NEPAL: Lete, Kali Gandaki Valley, 2,600 m., on rotten tree trunk, 12 June 1954, . Stainton, Sykes & Williams 5730; same locality, 8 July 1954, Stainton, Sykes & Williams 1639. Taglung, Kali Gandaki, 27 Aug. 1954, Stainton, Sykes & Williams — 7494.
Distribution: India, Nepal, Philippines, Japan. |
No spores were found in any of the gatherings. Stainton, Sykes & Williams 5730 has a grey cap, 1639 has grey cap with wide brown margin, and in 7494 the cap is” entirely brown.
, nd
FUNGI OF RECENT NEPAL EXPEDITIONS 129
THELEPHORACEAE CORTICIUM CAERULEUM (Pers.) Fries, Epicrisis Syst. Mycol. : 562 (1838).
Thelephova caerulea Pers., Mycol. Eur. 1: 147 (1822).—Fries, Elench. Fung. 1 : 202 (1828).
NEPAL: Ganesh Himal, Ankhu Khola, 2,800 m., on bark in broad-leaved forest, 17 May 1962, Stainton 3731. Distribution: America, Europe, India, Australia, Japan.
HYMENOCHAETE MOUGEOTII (Fries) Cooke in Grevillea 8 : 147 (1880). Thelephora mougeotii Fries, Elench. Fung. 1 : 188 (1828).
NepaL: Annapurna Himal, on branches of Rhododendron campanulatum, 30 Aug. 1954, Stainton, Sykes & Williams 6641. Rambrong, Lamjung Himal, on branch of Betula utilis, 7 July 1954, Stainton, Sykes & Williams 6202. Mewa Khola, 1,300 m., on dead wood, 1 Feb. 1962, Norkett 9182. Ganesh Himal, Ankhu Khola, 2,800 m., on bark in broad-leaved forest, 17 May 1962, Stainton 3732. Ganesh Himal, Mailung Khola, 4,000 m., 20 May 1962, Stainton 3744.
Distribution: Europe, India, Nepal, Australia, New Zealand, China.
HYMENOCHAETE RHEICOLOR (Mont.) Lév. in Ann. Sci. Nat., Ser. 3, Bot. 5 : 151 (1846).
Stereum rheicolor Mont. in Ann. Sci. Nat., Ser. 2, Bot. 18 : 23 (1842).
Stereum tenuissimum Berk. in Hook., Lond. Journ. Bot. 6 : 510 (1847).
Hymenochaete sallei Berk. & Curt. in Journ. Linn. Soc. Lond., Bot. 10 : 333 (1868).
Hymenochaete tenuissima Berk. apud Berk. & Curt. in Journ. Linn. Soc. Lond., Bot. Io : 333 (1868) nom. nud.
Hymenochaete tenuissima (Berk.) Berk. & Broome in Journ. Linn. Soc. Lond., Bot. 14 :
67 (1873). Stereum elegantissimum Spegazz. in An. Soc. Cient. Argent. 17 : 78 (1884).
NEPAL: Dhankuta Province, Taplejung district, Sanghu, 2,000 m., 17 Oct. 1961, Norkett 5696 B; same locality, 15 Nov. 1961, Norkett 7112 C. Distribution: North and South America, Africa, West Indies, India.
HYMENOCHAETE RUBIGINOSA (Fries) Lév. in Ann. Sci. Nat., Ser. 3, Bot. 5 : 151 (1846).
Thelephora rubiginosa Fries, Syst. Mycol. 1 : 436 (1821).
NEPAL: Chainpur district, Tumlingtar, 600 m., on dead tree, 13 Dec. 1961, Norkett 8109 D; same locality, 16 Dec. 1961, Norkett 8567. Bistribution: Worldwide.
130 FUNGI OF RECENT NEPAL EXPEDITIONS HyYMENOCHAETE TABACINA (Fries) Lév. in Ann. Sci. Nat., Ser. 3, Bot. 5 : 152 (1846).
Thelephora tabacina Fries, Syst. Mycol. 1 : 437 (1821). Stereum tabacinum (Fries) Fries, Epicrisis Syst. Mycol. : 550 (1838).
NEPAL: Annapurna Himal, on branches of Rhododendron campanulatum, 3 Aug. 1954, Stainton, Sykes & Williams 6642. Near Dogadi Khola, 4,300 m., on dead shrubs, 11 Aug. 1954, Stainton, Sykes & Williams 38178.
Distribution: Worldwide.
LOPHARIA CRASSA (Lév.) Boidin in Bull. Soc. Mycol. Fr. 74 : 479 (1958). |
Thelephora crassa Lév. in Ann. Sci. Nat., Ser. 3, Bot. 2 : 209 (1844).
Stereum umbrinum Berk. & Curt. apud Berk. in Grevillea 1 : 164 (1873). Hymenochaete vinosa Cooke in Grevillea 8 : 149 (1880). | Laxitextum crassum (Lév.) Lentz in U.S. Dept. Agric., Agric. Monogr. 24 : 20 (1955). |
NEPAL: Dhankuta Province, Dhankuta, near Mahe, 1,300 m., 20 Sept. 1961, Norkett 5157 C. Sombu, 1,600 m., 23 Sept. 1961, Norkett 5289. Dhankuta Pro- vince, Taplejung district, Sanghu, 3,000 m., 3 Oct. 1961, Norkett 5639 A; same locality, 2,000 m., 15 Nov. 1961, Norkett 7112 C.
Distribution: America, Europe, Africa, India, Australia, New Zealand.
STEREUM OSTREA (Fries) Fries, Epicrisis Syst. Mycol. : 547 (1838).
Thelephora fasciata Schwein. in Schrift. Naturf. Ges. Leipz. 1 : 106 (1822). Thelephora ostrea Fries, Elench. Fung. I : 175 (1828).
Thelephora versicolor var. fasciata (Schwein.) Fries, loc. cit.
Stereum fasciatum (Schwein.) Fries, Epicrisis Syst. Mycol. : 546 (1838).
NEPAL: Arun Valley, Khandbari, 2,300 m., on rotting log, 31 Aug. 1956, Stainton TAG2.
Distribution: widespread; previously recorded from the Himalayas.
Since Fries did not treat Thelephora fasciata Schwein. as a separate species in his Elenchus, which is part of the starting-point for the Fungi caeteri, the epithet of his IT’. ostrea must be adopted when the two names are regarded as synonyms.
STEREUM HIRSUTUM (Fries) Gray, Nat. Arrang. Brit. Pl. 1 : 653 (1821).
Thelephora hirsuta Fries, Syst. Mycol. 1 : 439 (1821).
NEPAL: Chipli, north of Pokhara, 3,600 m., 18 Apr. 1954, Stainton, Sykes & | Williams 4885. Arun Valley, Hatiar, 2,600 m., on rotting log, 2r Aug. 1956 3 Stainton r4ro. Arun Valley, Sibrung, 27 Aug. 1956, Stainton 1447. Kasuwa Khola, © 3,300 m., 12 Sept. 1956, Stainton 1637. Milke Danda Forest, 3,000 m., 16 Nov. — 1961, Norkett 7125. Sanghu, 2,300 m., 23 Nov. es Norkett 8196.
Distribution: Worldwide.
FUNGI OF RECENT NEPAL EXPEDITIONS 131
STEREUM ROSEO-CARNEUM (Schwein.) Fries, Acta Soc. Sci. Upsala, Ser. 3, I : 112 (1851).
Thelephora roseo-carnea Schwein. in Schrift. Naturf. Ges. Leipz. 1 : 107 (1822). Laxitextum roseo-carneum (Schwein.) Lentz in U.S. Dept. Agric., Agric. Monogr. 24 : 22
(1955).
NepaAL: Dhankuta Province, Taplejung district, Sanghu 2,060 m., 15 Nov. 1961, Norkett 7112 B.
Distribution: North and South America, Japan, China.
The fructifications are pinkish buff, resupinate on twigs; the paraphyses have branching tips; spores 8 X 4:5 y.
STEREUM SANGUINOLENTUM (Fries) Fries, Epicrisis Syst. Mycol. : 549 (1838). Thelephora sanguinolenta Fries, Syst. Mycol. 1 : 440 (1821).
NEPAL: Gurjakhani, 3,160 m. on small branches, 30 July 1954, Stainton, Sykes & Williams 3678. Distribution: North America, Europe, South Africa, Australia, New Zealand.
STEREUM SUBPILEATUM Berk. & Curt. apud Berk. in Hook., Journ. Bot. 1 : 238 (1849).—Lentz in U.S. Dept. Agric., Agric. Monogr. 24 : 36 (1955).
Stereum insigne Bresad. in Nuovo Giorn. Bot. Ital. 23 : 158 (1891). Xylobolus subpileatus (Berk. & Curt.) Boidin in Rev. de Mycol. 23 : 336 (1958).—Lentz in Sydowia 14 : 118 (1960).
NEPAL: Bakhri Kharka, north of Pokhara, on rotten tree trunk, 24 Apr. 1954, Stainton, Sykes & Williams 5059. Milke Danda Forest, Dhankuta Province, 2,800 m., 2 Nov. 1961, Norkett 6806 A; same area, 3,000 m., 16 Nov. 1961, Norkett 7124.
Distribution: America, Europe, India, Indonesia, China, Japan.
Lentz and Boidin should be consulted for modern interpretations of this aggregate species.
STEREUM sp. cf. suULCATUM Burt apud Peck in New York State Mus. Annu. Rep.
54, 1, App. I : 154 (1901).
NEPAL: South of Gurjakhani, 3,600 m., on tree in Abies forest, 16 Aug. 1954, Stainton, Sykes & Williams 3868.
Large robust sporophores superficially like those of a large S. princeps (Jungh.) Lév. but having a tomentose sulcate pileus, coarse, more or less parallel skeletal hyphae intermingled with generative hyphae, and no acanthophyses as such, but merely some slightly granular cystidial hyphae. The spores, irregularly globose, smooth or very faintly punctate and amyloid, 4-5-6 ~ diameter, resemble those of S. sulcatum Burt and those of S. taxodii Lentz & McKay (Mycologia 52 : 262 (1960)), two species recently transferred by H. L. Gross to his genus Echinodontium (Myco- path. & Mycol. Appl. 24 : 8, 11 (1964)). The*Nepal fungus however differs in the
132 FUNGI OF RECENT NEPAL EXPEDITIONS
absence of large encrusted cystidia and in the possession of large flabelliform reflexed pilei. In view of the large and conspicuous fructifications it would seem unlikely that this fungus has not been recorded previously. I therefore defer describing it as new to science.
THELEPHORA CARYOPHYLLAEA Fries, Syst. Mycol. 1 : 430 (1821).
Phylacteria caryophyllea Patouill., Hym. Eur. : 154 (1887), nom. nud. Phylacteria caryophyllea (Fries) Patouill. ex Bourd. & Maire in Bull. Soc. Mycol. Fr. 36:
76 (1920).
NEPAL: Taplejung district, above Sanghu, 2,000 m., amongst moss on earth, 12 Oct. 1961, Norkett 5927 A. Distribution: Worldwide.
VARARIA RHODOSPORA (Wakef.) G. H. Cunn. in Proc. Linn. Soc. N.S.W. 77 : 291 (1953); in N.Z. Dept. Sci. Industr. Res. Bull. 145 : 100 (1963).
Stereum duriusculum sensu Bresad. in Ann. Mycol. 6 : 43 (1908), non Berk. & Broome.
A sterostromella rhodospora Wakef. in Kew Bull. 1915 : 372 (1915).—Banergee in Journ. Ind. Bot. Soc. 14 : 45 (1935).
A sterostromella dura Bourd. & Galz. apud Bourd. & Maire in Bull. Soc. Mycol. Fr. 36:
74 (1920). Dichostereum durum (Bourd. & Galz.) Pilat in Ann. Mycol. 24 : 223 (1926).
NEPAL: Sanghu, 820 m., 9 Nov. 1961, Norkett 7324.
Distribution: America, Europe, Africa, India, Japan, Australia, New Zealand.
The species was recorded from India by Banergee. The Nepal collection con- sists of tough, resupinate thalli covering earth beneath tree roots, about 2-3 mm. thick, mid-brown, of stratose context. Only a few basidia were observed embedded in the dichophysoid paraphyses; spores globose, straw-coloured, echinulate, 5-6 yu diameter; context hyphae brown, dendroidly and dichotomously branched, most markedly and densely in the hymenial layer, where they form the brown dendro- physes and dichophyses; intermingled with them are finer, readily stained hyphae. Rogers & Jackson (Farlowia I : 309 (1943)), treat Dichostereum durum as a synonym of Vararia pallescens (Schwein.) Rog. & Jacks. Type material of Thelephora pal- — lescens Schwein. in the B.M. Herbarium differs in several particulars, notably its — finer context, and would appear to be not merely a different growth-form, as these ~ authors suggested, but a distinct species.
CLAVARIACEAE
CLAVULINA MUSSOORIENSIS Corner, Thind & Dev in Trans. Brit. Mycol. Soc. 41 : 204 t. 8 fig. 3, text-fig. r (1958). NEPAL: Near Gurjakhani, 2,800 m., among grass on open slope, 28 July 1954, Stainton, Sykes & Williams 3670.
Distribution: India. (E.J.H.C.)
ee
FUNGI OF RECENT NEPAL EXPEDITIONS 133
Clavulina alta Corner, sp. nov.
Receptacula ad 11 cm. alta, alba, sicco luride flava; stipite 2-6 cm. x 3-8 mm., bene evoluto; ramulis inferioribus polychotomis v. applanato-multifidis, superior- ibus r mm. latis dichotomis v. cristatis, axillis inferioribus 3-6 mm. latis. Sporae 95-14 X 7:5-9 4, subglobosae, lacrymiformes v. pyriformes, apiculo 1 yw longo. Basidia 6:5—7°5 lata, bispora. Hymenium incrassatum; cystidiis nullis; hyphis subhymenialibus 6-17 yu latis, fere pseudoparenchymaticis. Hyphae 3-12 y latae, fibulatae, tenue tunicatae, cellulis potius brevibus.
NEPAL: Chimgaon, north of Tukucha, Kali Gandaki, 3,500 m., on ground beneath conifers, 14 Sept. 1954, Stainton, Sykes & Williams 7825. (Herb. Mus. Brit. holotype). This resembles C. rugosa (Fries) Schroet. in the large spores and wide sub- hymenial hyphae, and C. cristata var. coralloides Corner in the form of the fruit-body. I have not seen such a distinct form before, and the spores are constantly rather
narrow for their length. (yee.
Lentaria macrospora Corner, sp. nov.
Receptacula ad 10 cm. alta, gregaria v. caespitosa, carneoflavida; stipite ad 25 X 2-4 mm., axillas inferiores polychotomas versus plus minus dilatato; ramulis superioribus teretibus dichotomis strictis ascendentibus, 1-2 mm. latis. Sporae 20-30 X 3°7-5'5 #, hyalinae, cylindricae, v. subclavatae, saepe curvatae v. Sig- moideae, et allantiformes, tenue tunicatae, haud amyloideae. Basidia ad 45 x g-10°5 mw; sterigmatibus 2—4, 7-8 w longis. Hymenium incrassatum; cystidiis nullis. Hyphae 2°5-7 uw latae, fibulatae, tunicis ad 0-5 m# vix incrassatis; in mycelio 2:5—4 latae, fibulatae, monomiticae, tunicis tenacibus sed vix incrassatis, passim partibus ampulliformibus ad 15 y latis inflatae, crystallis sphaeroideis 2—7 yw latis inter hyphas numerosis.
NEPAL: Tamur Valley, Ghunsa, east of Walungchung Gola, 4,300 m., on ground under conifers, 27 July 1956, Stainton 1145. (Herb. Mus. Brit. holotype).
This resembles in shape and colour the common tropical L. surculus (Berk.) Corner but the spores are much longer, the hyphal walls are scarcely thickened, and the habitat seems to be humicolous. Many basidia had 1-3 long spiculiform sterig-
mata, but they may have been abnormal and formed after collection. (E.J.H.C.)
RamarIA aff. BOTRYTOIDES (Peck) Corner, Monogr. Clavaria, Ann. of Bot., Mem. I : 562 (1950). Clavaria botrytoides Peck in N.Y. State Mus., Mus. Bull. 94 : 49, t. 93 figs. 5-7 (1905).
NEPAL: Taglung, south of Tukucha, Kali Gandaki, 3,300 m., on ground beneath trees, Ir July 1954, Stainton, Sykes & Williams 1691. Distribution (of Ramaria botrytoides): America, southern parts of Australia and
Tasmania, Japan; when the species is interpreted in a wide sense. (Ey 2G.)
134 FUNGI OF RECENT NEPAL EXPEDITIONS
RAMARIA SUECICA (Fries) Donk in Med. Bot. Mus. Herb. Rijks Univ. Utrecht 9g : 105 (1933).—Corner, Monogr. Clavaria, Ann. of Bot., Mem. 1 : 629 (1950).
Clavaria suecica Fries, Syst. Mycol. 1 : 469 (1821). Clavariella suecica (Fries) Karst. in Bidr. Kann. Finl. Natur. & Folk 37 : 187 (1882).
NEPAL: South of Gurjakhani, 3,300 m., on damp shady forest bank, 18 Aug.
1954, Stainton, Sykes & Williams 3902. Distribution: Europe, China, Canada, Northern U.S.A. (pL)
RAMARIA FLACCIDA (Fries) Ricken, Vadem. Pilzfr.: 254 (1918). Clavaria flaccida Fries, Syst. Mycol. 1 : 471 (1821).
NEPAL: Taglung, south of Tukucha, Kali Gandaki, 3,300 m., in pine wood, 22
Sept. 1954, Stainton, Sykes & Williams 7971. Distribution: America, Europe, South Africa, Australia, China, Japan. (E.) FC)
RAMARIA OBTUSISSIMA (Peck) Corner, Monogr. Clavaria, Ann. of Bot., Mem. 1 : 609 (1950). Clavaria obtusissima Peck in N.Y. State Mus., Mus. Bull. 167 : 39 (1913).
NEPAL: Taglung, south of Tukucha, Kali Gandkai, 3,000 m., beneath conifers, 12 July 1954, Stainton, Sykes & Williams 1790. Distribution: U.S.A. and Canada.
The pink form is represented. (ESAS
RAMARIA SUBAURANTIACA Corner apud Balf.-Browne in Bull. Brit. Mus. (Nat. Hist.), Bot. I : 200 (1955). NEPAL: North of Barse, 4,000 m., on Abies stump, 14 Aug. 1954, Stainton, Sykes & Williams 3851. Distribution: Tibet. Spores 10:5-I5 xX 5-6 mw, rather coarsely subverrucose. No clamps. (E.J.C)
CYPHELLACEAE Chromocyphella bryophyticola Balfour-Browne, sp. nov.
Ee lee ng
Fungus cupulatus, cupulis sessilis, levis, griseo-albis, 0-5 mm. diam.; contextis
tenuis, mollis, 15 ~ latis ex hyphis elongatis efformatis; hymenio levo, brunneo;
basidiis cylindricis, 15-16 x 4-5 4; sporis globosis, 5~7 « diam., brunneis, punctatis. NEPAL: Sanghu, 1,800 m., on moss, Pterobryopsis, and on an intermingled liver- wort, on shady rock, 8 Nov. 1961, Norkett 7292. (Herb. Mus. Brit. holotype).
FUNGI OF RECENT NEPAL EXPEDITIONS 135
This fungus is not unlike Cyphella chromospora Patouill. (Tab. Anal. Fung. I : 19, fig. 32 (1883)), but the spores are larger. It differs from Chromocyphella burtit Bridge Cooke (Sydowia, Beiheft 4 : 137 (1961)), in its smaller basidia and its definitely punctate spores.
This appears to be the first record of a “‘ Cyphella ’”’ in the Himalayan region.
SCLERODERMATEACEAE SCLERODERMA AURANTIUM Pers., Syn. Meth. Fung. : 153 (1801).
NEPAL: Mathand, near Pokhara, 1,160 m., on shady bank, 22 June 1954, Stainton, Sykes & Williams 5851.
Distribution: Worldwide.
Spores 8—Io y, reticulated.
LYCOPERDACEAE
BovisTa sp. cf. BOVISTOIDES (Cooke & Massee) Ahmad, Gasteromycetes W. Pakistan, Publ. Dept. Bot. Univ. Panjab 11 : 16 (1952).
Mycenastrum bovistoides Cooke & Massee in Grevillea 16 : 26 (1887).
NEPAL: Above Dogadi Khola, amongst short grass on exposed slope, 21 June 1954, Stainton, Sykes G& Williams 32rt.
Distribution (of Bovista bovistoides): India.
The Nepal collection appears to be very close to this species but has very slightly _warted spores, whereas Ahmad described them as smooth. The capillitium threads agree in being chestnut brown and unpitted.
BoVISTA ECHINELLA Patouill. in Bull. Soc. Mycol. Fr. 7 : 165 (1891).
Bovistella echinella (Patouill.) Lloyd, Mycol. Notes 23 : 286, t. 89 figs. 1-2 (1906). Lycoperdon echinella (Patouill.) Ahmad in Journ. Ind. Bot. Soc. 20 : 138 (1941).
NEPAL: Chainpur district, Tumlingtar, 600 m., on earth near base of cliff of Sabhaya River, 9 Dec. 1961, Norkett 8682 A.
Distribution: North and South America, Jamaica, Europe, Pakistan.
The spores are smooth, not echinulate as described by Patouillard, but show “lines” beneath the outer membrane which at a certain focus appear like spines; the pedicels are mostly 6 yw long.
GEASTRUM FIMBRIATUM (Fries) E. Fisch. in Engl. & Prantl, Nat. Pflanzenfam., 2 Aufl., 7a : 73 (1933).
Geaster fimbriatus Fries, Syst. Mycol. 3 : 16 (1829).
NepaLt: Tukucha, Kali Gandaki, 3,600 m., 12 Oct. 1954, Stainton, Sykes & Williams 8212. Distribution: America, Europe, Africa, India, Australia.
136 FUNGI OF RECENT NEPAL EXPEDITIONS
GEASTRUM HARIOTI (Lloyd) E. Fisch. in Engl. & Prantl, Nat. Pflanzenfam., 2 Aufl., 7a : 73 (1933). Geaster hariotit Lloyd, Mycol. Not. 25 : 311, t. 99 figs. 7-9 (1907).
NEPAL: Tumlingtar, Sabhaya Khola, 600 m., on earth beneath bamboo, 20 Dec. 1961, Norkett goo2.
Distribution: South America, West Indies, Europe, Ceylon, Australia (according to Cunningham, v. below).
The eight specimens from Nepal resemble the descriptions of Geastrum hariotii very closely in being non-hygroscopic, in having a sessile endoperidium, dark, sulcate peristome and small, 3—3:5 “ diam., minutely verrucose spores. But the surface of the endoperidium is furfuraceous or granular rather than pitted as des- cribed by some authors (Cunningham, Gasteromycetes Austral. and N.Z. : 165
(1942)). LYCOPERDON PYRIFORME Pers., Syn. Meth. Fung. : 148 (1801). NepAL: Tukucha, Kali Gandaki, 3,500 m., 12 Sept. 1954, Stainton, Sykes @
Williams 7806. Distribution: Worldwide.
FUNGI IMPERFECTI SPHAEROPSIDACEAE CONIOTHYRINA AGAVES (Durieu & Mont.) Petr. & Syd. in Beih. Rep. Spec. Nov. Reon. Ver, 42° 322 (1927).
Phoma agaves Durieu & Mont. in Mont., Syll. Pl. Crypt.: 271 (1856). Coniothyrium agaves (Durieu & Mont.) Sacc., Syll. Fung. 3 : 318 (1884).
NEPAL: Sanghu, 2,000 m., on Agave, 17 Nov. 1961, Norkett 7154; Sombu, 1,600
m., on Agave, 23 Sept. 1961, Norkett 5313. Distribution: America, South Europe, Africa, India.
NECTRIOIDACEAE
ASCHERSONIA sp. cf. VIRIDANS (Berk. & Curt.) Patouill. in Bull. Soc. Mycol. Fr. FFAS (A801);
Hypocrea viridans Berk. & Curt. in Journ. Linn. Soc. Lond., Bot. 10 : 376 (1868). Aschersonia disciformis Patouill. in Bull. Soc. Mycol. Fr. 8 : 136 (1892). Aschersonia viridula Sacc. in Ann. Mycol. 11 : 547 (1913).
NEPAL: Phewa Tal, 800 m., on leaves of Castanopsis sp., 8 May 1954, Stainton, Sykes & Williams 5272 (a). .
Distribution (of A. viridans): Central America (Trinidad, Vera Cruz, Cuba, Ecua- dor), Brazil, Mexico. ;
The Nepal fungus was growing on white fly. It differs from previous accounts in the greater number of pycnidia to each stromatic cushion, which is completely dotted over with the greenish ostioles; conidia 12-16 X I-52 yp.
FUNGI OF RECENT NEPAL EXPEDITIONS 137
Another Aschersonia also was collected on Castanopsis leaves; this appears to be close to A. caespiticia Syd. (in Engl., Bot. Jahrb. 54 : 260 (1916)), but differs in the rough surface of the tubercles and its small basal cushions. Tilhar, 3 Nov. 1954, Stainton, Sykes & Williams 9251 (a).
LEPTOSTROMATACEAE
MELASMIA SALICINA Tulasne frat., Sel. Fung. Carp. 3 : 119, t. 15 figs. 15-17 (1865).
NEPAL: East of Chalike Pahar, 4,160 m., on Salix sp., 25 Sept. 1954, Stainton, Sykes & Williams 4587.
Distribution: not previously collected in this part of the world but its perfect state, Rhytisma salicinum Fries, has been recorded from the Punjab.
MELANCONIACEAE
Mastigonetron americanum (Mont.) Balfour-Browne, comb. nov.
Pestalotia americana Mont. in Gay, Hist. Chile, Bot. 7 : 481 (1850).—Guba, Monogr. Monochaetia & Pestalotia : 268 (1961).
Seiridium liquidambaris Berk. & Curt. apud Berk. in Grevillea 2 : 154 (1874).
Mastigonetron fuscum Klebahn in Myc. Centralbl. 4 : 18, fig. 37-38 (1914).
Monochaetia liquidambaris (Berk. & Curt.) Guba, loc. cit. (1961), nom. synon.
NEPAL: Taplejung district, Sanghu, Tamrang Khola, 2,000 m., on dead twigs, Ig Oct. 1961, Norkett 6319 A.
Distribution: North and South America.
The acervuli, about 0-5 mm. in diameter, are scattered over the twigs and resemble lenticels in appearance. The conidia, 20-27 x g—I0°5 m, are dark brown, unicellular, ovoid or ellipsoid, each with a hyaline apical appendage 30-40 x 2 m, and a pedicel I2 X 2 “, approximately.
The fungus agrees exactly with Klebahn’s species and apparently also with the type of P. americana (in spite of Montagne’s description of the conidia as biseptate) since Guba, loc. cit., states that ‘‘ Montagne’s drawings of the fungus and my study show dark colored 1-celled ellipsoid conidia’. The drawing mentioned is apparently unpublished.
In view of its unicellular conidia the species cannot be included in either Mono- chaetia or Pestalotia. As for Seiridium Nees & Henry, (Syst. Pilze : 18, t. 3 (1837)), arguments can be brought forward for retaining it as an earlier name for Mastigone- tron if one regards as accurate the elder Nees’s (Syst. Pilze & Schwamme : 22 (1816), t. I fig. 19 (18r7)) description and drawing of unicellular appendaged conidia. This non-septate condition of the conidia was accepted by Berkeley & Curtis when they selected this genus for the fungus on Liguidambar. Alternatively, Seiridium can be regarded as an older name for Monochaetia if Fries’s (Syst. Mycol. 3 : 473 (1832)) statement that his examination of Nees’s material showed multiseptate conidia is taken as correct. The original collection appears to be lost but other collections since then and reputed to be the same species,*i.e. S. marginatum, are invariably
138 FUNGI OF RECENT NEPAL EXPEDITIONS
described as having a brown septate conidium with a transparent apical seta. The probable explanation for this discrepancy is that Nees chanced to examine and illustrate the unripe fungus, i.e. before the conidia became septate, while they were still unicellular, spindle-shaped and contained grey granular protoplasm. Fries, on the other hand, and all subsequent workers have described the mature fungus, which represents what is now generally regarded as typical Monochaetia. Taking this latter view, or better still treating Seividium as a nomen confusum, this generic name cannot be used in place of Mastigonetron and therefore the correct name for the Nepal fungus becomes Mastigonetron americanum, as cited above.
STILBACEAE
ARTHOBOTRYUM NILGIRENSE Subram. in Proc. Ind. Acad. Sci., Sect. B, 46 : 324,
fig. I (1957).
NEPAL: Chainpur, Tumlingtar, 660 m., on old bamboos, 15 Dec. 1961, Norkett 8868.
Distribution: Previous record and original description from bamboo, Sim’s Park, Coonoor, Madras, 1956.
Podosporium himalensis Balfour-Browne, sp. nov. (Fig. 4).
Synnemata atrobrunnea numerosa et effusa ex hyphis parallelibus juxtapositis efformata. Conidia atrobrunnea, I-12 septata, obclavata, leniter curvata et irregu- | lariter disposita, 9-I5 X I6-I00 pm.
NEPAL: Lamjung Himal, 4,500 m., black woolly growth on branches of Rhododen- dron campanulatum, 14 July 1954, Siainton, Sykes & Williams 6341. (Herb. Mus. Brit., holotype). ;
The general form of the synnemata is similar to that described by Subramanian (Journ. Ind. Bot. Soc. 35 : 73 (1956)) for his Prathoda saparva; the conidia however are different and the conidiophores hardly distinct from the hyphae and not geni- — culate. The Nepal material resembles an extremely luxuriant form of Podosporium rigidum Schwein., originally described from Carolina. The hyphae are hormiscium- — like and very probably any portion breaking away can regenerate fresh growth, — independently of conidial reproduction. i
STILBUM CINNABARRINUM Mont. in Ann. Sci. Nat., Ser. 2, Bot. 8 : 360 (1837). i Stilbum lateritium Berk. in Ann. Nat. Hist. 4 : 291, pl. 8 fig. 2 (1840).
NEPAL: Tumlingtar, by shore of Sabhaya Khola, 600 m., 11 Dec. 1961, Norkett 8107 C.
Distribution: North and South America, Cuba, Dominica, Africa, Nepal, India, — Ceylon, Australia.
Pleonectria pseudotrichia (Schwein.) Wollenw. is its perfect form. The present collection occurs on unnamed bark and is very sparing.
FUNGI OF RECENT NEPAL EXPEDITIONS 139
anes Seas es; Res &e. SAN Q
Conetme. ee Cee es oS =
ia 8 Cr. ans
ci i. Oo me
seca aN Se
=
Zee, aes [ —t = a
ne
2S 0
Fic. 4. Podosporium himalensis Balfour-Browne. A, general aspect; B, conidia and conidiophorous hyphae. Stainton, Sykes & Williams 6341.
STILBUM INCONSPICUUM Currey in Trans. Linn. Soc. Lond., Ser. 2, Bot. 1 : 129 (1876).
Stilbum kurzianum Cooke in Grevillea 16 : 71 (1888).
NEPAL: Sanghu, 2,000 m., 17 Nov. 1961, Norkett 7153.
Distribution: India.
On dead twigs of Rosa sp. Synnemata 3-4 mm. tall, pale orange soon becoming cinereous; conidia rod-shaped 7-9 Xx 3 /.
140 FUNGI OF RECENT NEPAL EXPEDITIONS
TUBERCULARIACEAE EPICOCCUM ANDROPOGONIS (Rabenh.) Schol-Schwarz in Trans. Brit. Mycol. Soc. AZ PAT COME: iT ALORG): Cerebella andropogonis Rabenh. in Bot. Zeit. 9 : 669 (1851).
NEPAL: Bhadauri, near Pokhara, 2,000 m., on inflorescence of a grass, I Nov. 1954, Stainton, Sykes & Williams 8326.
Distribution: Worldwide but chiefly in the tropics and subtropics.
This genus has been revised recently by Langdon (Mycol. Commonw. Mycol. Inst. Papers, 61 : 1-18 (1955) under the name Cerebella), and by Schol-Schwarz (tom. cit. :
149-173).
EPICOCCUM PURPURASCENS Schlechtend., Fl. Berol. 2.: 136 (1824).—Link in L., Sp. Ply ede Ay Ou(Z) TOS (1825).
Epicoccum nigrum Link, loc. cit—Schol-Schwarz in Trans. Brit. Mycol. Soc. 42 : 170 (1959). ;
NEPAL: Argam, near Pokhara, 830 m., on leaves and stems of Acrocephalus indicus (Burm.) O. Kuntze, 10 Sept. 1954, Stainton, Sykes & Williams 7146. Distribution : Worldwide.
SPE CLAS toi ho RA TUR
BALFOUR-BrRowneE, F. L. (1955). Some Himalayan Fungi. Bull. Brit. Mus. (Nat. Hist.), Bot. 1 : 189-218.
BERKELEY, M. J. (1850). Decades of Fungi. Hook., Journ. Bot. 2 : 42-51, 76-88, 106-112.
(1851). Decades of Fungi. Hook., Journ. Bot. 3 : 39-49, 77-84, 167-172, 200-206.
(1852). Decades of Fungi. Hook., Journ. Bot. 4 : 97-107, 130-142.
(1854). Decades of Fungi. Hook., Journ. Bot. 6 : 129-143, 161-174, 204-212, 225-235.
BonpDartzeEV, A. S. (1953). Tvrutov’ie grib’i Europei skoi chasti S.S.S.R. i Kavkaza. [{Poly- poraceae of the European part of the U.S.S.R. and the Caucasus.| Leningrad.
But Ler, E. J. & BisBy, G. R. (1960). Fungi of India. 2nd ed., revised by R. S. VASUDEVA, New Delhi.
CorNER, E. J. H. (1932). The fruit-body of Polystictus xanthopus. Ann. of Bot. 46 : 71-111.
(1953). The construction of Polypores. I. Introduction. Phytomorphology 3 : 152-167.
CUNNINGHAM, G. H. (1947). Notes on the classification of the Polyporaceae. N.Z. Journ.
Sct. & Technol. 28, A : 238-251.
(1947-1950). New Zealand Polyporaceae. N.Z. Dept. Sci. Industry. Res. Bull. 72-83.
(1965). Polyporaceae of New Zealand. N.Z. Dept. Sci. Industr. Res. Bull. 164.
Donk, M. A. (1933). Revision der niederlandischen Homobasidiomycetae—Aphyllophora-
ceae. II. Med. Bot. Mus. Herb. Rijks Univ. Utrecht 9 : 1-278. .
(1960). The generic names proposed for Polyporaceae. Persoonia 1 : 173-302. ,
ImazEkI, R. (1943). [Genera of Polyporaceae of Nippon.} In Japanese. Bull. Tokyo Sct. Mus. 6, I-III.
KotvaBa, F. & Pouzar, Z. (1957). Pozndamky k tiidéni evropskych choroSt. [Notes on the classification of European pore Fungi.] Ceskd Mykol. 11 : 152-170.
Noses, M. (1958). Cultural characters as a guide to the taxonomy and phylogeny of the Polyporaceae, Canad. Journ, Bot. 36 : 883-926.
i
BUNGI-OF RECENT NEPAL EXPEDITIONS 141
OveRHOLTsS, L. O. (1953). The Polyporaceae of the United States, Alaska and Canada. Ann Arbor, Mich.
PivAt, A. (1936-1942). Polyporaceae. In Kavina, C. & PiLAt, A., Atlas des Champignons de l'Europe, 3. Praha.
Pinto-Lopez, J. (1952). ‘‘ Polyporaceae ’’ Contribuicgao paraa sua biotaxonomia. Mem. Soc. Broter. 8 : 5-191.
SHEALS, J. G. & INGLIs, W. G. (1965). The British Museum (Natural History) expedition to east Nepal 1961-62. Introduction and lists of localities. Bull. Brit. Mus. (Nat. Hist.), Zool. 12 : 95-114.
SINGER, R. (1962). The Agaricales in modern taxonomy. 2nd ed. Weinheim.
TexieErrA, A. R. (1962). The taxonomy of the Polyporaceae. Biol. Rev. Cambr. Phil. Soc.
37 : 51-81.
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A SYNOPSIS OF JAMAICAN MYRSINACEAE
BY WILLIAM THOMAS STEARN Karak -
Pp. 143-173; 25 Text-figures; Plates 6-13
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‘$ QD 10 JUN 1969
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) BOTANY Vol. 4 No. 4 LONDON : 1969
THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical serves.
Paris will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.
In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department.
This paper is Vol. 4, No. 4 of the Botany series.
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Issued 30 May, 1969 Price £1 4s.
A SYNOPSIS OF JAMAICAN MYRSINACEAE
By WILLIAM T. STEARN
SUMMARY
The family Myrsinaceae is represented in Jamaica by three genera, Ardisia, Wallenia and Myrsine (including Rapanea). Recognition of the species presents especial difficulty in Wallenia, which is dioecious; some taxa are known only from specimens with male flowers or female flowers or fruits, i.e. in only one of three states needed for adequate discrimination; few are known in all three; 15 species, two occurring also in Cuba, the others endemic, are listed here. Avrdisia is represented by eight species, one (A. solanacea) introduced and naturalized; Myrsine by two, which Mez placed in Rapanea. Keys, synonymy and statements of distribution are given. Ardisia brittonit and A. byrsonimae are described as new; A. vosea Urban (1908), non A. rosea King & Gamble (1905), is renamed A. urbanii.
INTRODUCTION
STUDENTS of the family Myrsinaceae tend to be censorious of the work of Carl Mez (1864-1944) on this difficult and to most botanists unattractive family, wherein, as C. L. Lundell (19662) has remarked, “‘ most generic characters are weak and those _ separating species are often weaker.’’ Nevertheless, they must needs be grateful to Mez for his comprehensive survey of its West Indian members published in Urban’s Symbolae Antillanae 2 : 389-433 (1901) and for his world monograph in Engler’s Das Pflanzenreich IV .236 (1902), in which he provided a helpful basic classification, co-ordinated the extensive literature, and enumerated and described the material then available. The subsequent great increase of specimens in herbaria and a more critical attention to matters of nomenclature and typification than prevailed then have, however, made necessary many alterations and some drastic revisions of Mez’s treatment; moreover, some characters emphasized by him in his keys have been found impracticable. E. Y. Hosaka, for example, in his account of the Hawaiian species of Myrsine sensu lato (in Occ. Pap. B.P. Bishop Mus. 16 : 25-76 (1940)) concluded that Rapanea, Suttonia and Myrsine, kept as separate genera by Mez, overlap so much in their characters that they should be combined in one genus, Myrsine, as indeed they were by Bentham & Hooker in 1873; H. C. D. de Wit (in Bull. Jard. Bot. Bruxelles 27 : 233-242 (1957)) has united Heberdenia with Ardisia. On the other hand, Lundell (in Wrightia 3 : 88 (1963)) has raised Mez’s Ardisia sect. Synardisia to generic rank as Synardisia. A few examples may suffice to indicate the nomenclatural rectifications needed. The name Ardisia coriacea Swartz (1788), based on Jamaican material, was misapplied by Mez to a species which does not occur in Jamaica, i.e. A. obovata Ham. (A. guadalupensis Griseb.). The name Ardisia sanguinolenta Blume (1825) is the correct name for A. zollingeri A.DC. (1844): A. wallichii A.DC is the correct name for A. sanguinolenta Wall. ex Mez (1902). Merrill has called attention (in Lingnan Sci. Journ. 11 : 50 (1932) and in Contrib. Arnold Arb. 8 : 131 (1934)) to Mez’s misinterpretation of A. humilis Vahl (1794), which, according to Merrill, is the same as A. hainanensis Mez (1902), while BOT, 4, 4 8
146 A SYNOPSIS OF JAMAICAN MYRSINACEAE
A. humilis sensu Mez is A. ellaptica Thunb. (1795) (syn. A. obovata Blume (Dec. 1825—March 1826), non A. obovata Ham. (Nov. 1825)). As regards the East Asiatic Myrsinaceae, a much needed nomenclatural and taxonomic overhaul has been made by E. H. Walker (in Philipp. Journ. Sci. 73 : 1-258 (1940), with supplementary notes in Journ. Arnold Arb. 23 : 344-355 (1942), in Bot. Mag. Tokyo 67 : ro5—11T1, 155-162, 203-213, 247-255 (1954) and in Quart. Journ. Taiwan Mus. 12 : 161-194 (1959)). Unfortunately, there exists no comprehensive survey of the American M-yrsinaceae later than Mez’s of 1902, although many new species have been published since by Urban, Standley and C. L. Lundell, who has also revised the genus Parathesis (Lundell, 1966d) and the Guatemalan members of the family (Lundell, 1966a). The author of a local Flora is accordingly obliged to make a study more or less independent of Mez’s work and to extend his enquiries beyond his own geographical area. The following synopsis of Jamaican Myrsinaceae is a by-product of preparing a more detailed account for the Flora of Jamaica; descriptions and particulars of localities to be given there in detail are either summarized below or omitted.
Mez in 1901 based his account of the Myrsinaceae occurring in Jamaica on 14 wild and two cultivated gatherings of Ardisia, 15 of Wallenia, 13 of Rapanea (i.e. Myrsine), i.e. 42 in all. For the Flora of Jamaica I have examined at least 120 Jamaican gatherings of Ardisia, 130 of Wallenia, and 50 of Myrsine, i.e. in all more than 300 gatherings. This increase in material has made evident the distribution of the Myrsinaceae in Jamaica, but has not greatly simplified the problems of their classifica- tion. Thus, despite much field work, the female flowers of Wallenia clusioides, W. crassifolia, W. discolor, W. erythrocarpa and W. xylosteoides and the male flowers of W. discolor, W. elliptica, W. erythrocarpa, W. punctulata and W. sylvestris have apparently missed collection; they have not been observed in the material studied, which, in addition to that of the British Museum (Natural History), London (cited below as BM), has included the specimens of the Systematisch-Geobotanisches Institut, Universitat G6ttingen (GOET), the Institute of Jamaica, Kingston, Jamaica (IJ), the Royal Botanic Gardens, Kew (K), the New York Botanical Garden (NY) and the University of the West Indies, Mona, Jamaica (UCWI) and the lecto- type of Wallenia grisebachii at the Botanische Staatssammlung, Munich. Thanks are here expressed to the Directors of these Institutions for their invaluable co-operation in lending or making available for study this extensive material. Nevertheless, the following account as regards the status and distinctive characters of some taxa is necessarily provisional.Text-figures 21-25 have been drawn by Joanna C. Webb (Mrs. D. Erasmus).
HISTORY
The first members of the Myrsinaceae to be recorded from Jamaica, indeed from all America, were Ardisia tinifolia and Wallenia laurifolia gathered by Hans Sloane in 1688 or 1689. He collected the Avdisza “on the Mountains near Mr. Elletson’s Plantation in Liguanee”’, i.e. in the parish of St. Andrew, Jamaica, and published it in his Catalogus Plantarum: 169 (1696) and Voyage to the Islands Madera etc. 2 : 98, t. 205 f. 2 (1725) as “‘ Jasminum forte, arboreum, foliis laurinis obtusis latioribus
A SYNOPSIS OF JAMAICAN MYRSINACEAE 147
atrovirentibus, flore pentapetalo racemoso purpureo reflexo’’. Concerning this, Sloane wrote: “ This Tree riseth to about thirty Foot high, having a Clay or Ash- colour’d pretty smooth Bark; its Twigs are set about with Leaves which are very smooth, of a dark green Colour, having a quarter of an Inch long Footstalks, being four Inches long and two broad in the Middle, where broadest, having an eminent middle Rib. The Flowers stand on several little Branches, being pentapetalous, purplish, bow’d back, having yellow Stamina in their Middle.”’ Olof Swartz found the same species “in aridis submontosis Jamaicae’”’ during his stay in the West Indies from 1784 to 1786, and gave it the binomial Ardisia tinifolia when founding the genus Ardisia in his Nova Genera et Species Plantarum seu Prodromus : 48 (1788).
Swartz referred five species to Ardisia: 1. A. tinifolia from Jamaica; 2. A. coriacea (now regarded as conspecific with A. tinifolia), its provenance vaguely recorded by him as “ India occidentalis ”’; 3. A. serrulata (now Parathesis serrulata (Swartz) Mez) from Hispaniola, these three being of his own collecting; 4. A. laterifolia (now Stylo- gyne laterifolia (Swartz) Mez), vaguely recorded as from “ India occidentalis ”’; 5. A. parasitica (now Grammadenia parasitica (Swartz) Griseb.) from “ Montserrat ”’ (later corrected to Dominica) ; these last two were known to him only from specimens in the herbarium of Sir Joseph Banks, now at the British Museum (Natural History), as the asterisk against the diagnosis indicates. Later authors have accepted A. tintfolia as the type-species of Ardisia. Swartz’s description of the species in his Flora Indiae Occidentalis 1 : 468 (1797) is fairly detailed and accompanied by a reference to his “Icon. fasc. 3”. Unfortunately, this third fascicle of his [cones Plantarum incognitarum (fasc. I, 1794; fasc. 2, 1800) was never published, although Swartz sent the illustrations and text to Schreber in Erlangen for publication. Schreber died in 1810, Swartz in 1818; the Palmische Verlagsbuchhandlung in Erlangen published a note in Flora (Regensburg) 3 : 144 (March 1820) offering these illustrations to whoever would undertake their publication, but no one responded. Their subsequent history is unknown until about 1890 when the Berlin botanist Ignaz Urban acquired a volume containing presumably all that had survived of Swartz’s 200 drawings of West Indian plants. They are cited in Urban’s Symbolae Antillanae (cf. r : 164 (1898)) as “‘ Sw. Ic. ined ’’, and in Fawcett & Rendle’s Flora of Jamaica (1 : xviii, etc. (I9I0)) as ‘‘ Sw. Icon. ined:”’. They could not be traced at the Botanisches Museum, Berlin-Dahlem after the 1939-45 war and hence were believed to have been destroyed by bombing in March 1943, as stated in Sitwell and others, Great Flower Books : 77 (1956). Luckily, however, Urban had sold this precious volume in 1922 (cf. Journ. Bot. (Lond.) 60 : 361 (1922)) toC. A. Lindman in Stockholm and it is now preserved in the library of the Royal Swedish Academy of Sciences; Plates 6 and 7 are reproduced here from these drawings by permission of the Royal Swedish Academy of Sciences, to which grateful acknowledgement is made. Sets of photographs of Swartz’s unpublished drawings have been deposited in the libraries of the British Museum (Natural History), London, the Institute of Jamaica, Kingston, Jamaica, and the Hunt Botanical Library, Pittsburgh, U.S.A. Swartz’s coloured drawings representing Ardisia tinifolia and A. coriacea, together with A. serrulata and A. parasitica, leave no doubt as to the application of the names. Swartz applied the name A. coriacea to a Jamaican plant (without evident veining
148 A SYNOPSIS OF JAMAICAN MYRSINACEAE
of the leaves but otherwise very similar to his A. tinifolia) which was described by Mez in rgo1 as A. harrisiana; hence, as pointed out by Urban (Symb. Antill. 3 : 330 (1902); op. cit. 8 : 519 (1921)), Mez used the name A. coriacea wrongly for a species of the Lesser Antilles, i.e. A. obovata (A. guadalupensis).
According to Sloane’s account (Voy. Nat. Hist. Jam. 1 : 234 (1707)) of the Jamaican plant now known as Wallenia laurifolia, but recorded by him as “‘ Bryonia nigra fruticosa, foliis laurinis, floribus, racemosis, speciosis’’, he ‘‘ gathered it, if I rightly remember, in St. Maries, near Cabeg¢a del oro in the North-side of this Island amongst the Woods ”’; he described the flowers simply as “ at the top of the Branch... standing in a bunch together, being many very beautiful and small’’. On this species Swartz in 1788 (Nov. Gen. & Sp. Pl. : 2, 31) founded the genus Wallenia, naming it in honour of Matthew Wallen, from whom Swartz received much hospitality while in Jamaica. Meanwhile, Jacquin had published in his Selectarum Stirpium Americanarum Historia : 17 (1763) the provisional designation Petesioides laurifolium referring to incomplete material collected by him in Santo Domingo (Haiti) between 1755 and 1759, and Petesioides was accordingly cited in the 1952, 1956, and 1961 editions of the International Code of Botanical Nomenclature, under No. 6304, as a nomen genericum rejiciendum necessitating the conservation of Swartz’s Wallenia. Linnaeus had, however, ruled in Critica botanica : 34 (1737) that “‘ nomina generica in otdes desinentia, e foro Botanico releganda sunt ”’ (generic names ending in oides are to be banished from the botanical forum). Hence, when Linnaeus himself used designations such as Baccharioides, Bannisteroides, Cornutioides, Euonymotdes, Hibiscoides, Oxycoccotdes, Senecioides etc. in his Flora Zeylanica (1747) for obscure plants and when loyal followers of Linnaean method and nomenclature such as Jacquin, Loefling and Rottboll used designations such as Celosioides, Diodiodes, Malpighioides, Sideroxyloides, Staehelinoides and Viscoides, they did not intend them to be permanent names for the genera concerned but merely as provisional or token names indicating resemblance; they intended that these -ozdes designations should later be replaced by correctly formed names. Jacquin’s description of his Petesioides indicates that he possessed only a specimen with functionally female flowers; he believed that it represented a new genus which could not be properly established until fruits became available; accordingly, in his own words, “‘ ego interea ab habitu nomen fluxum imposui’”’ (I meanwhile, from its habit of growth, have given it a perishable name). Such token designations are in a different category from the pre-Linnaean generic names ending in -o7zdes, such as Capnotdes, Ficoides, Nymphotdes, brought into post-Linnaean use as definite generic names by Miller, Hill, Adanson, Medicus and others. In a like spirit Rottb6ll in 1772 described Schoenoides and Scirpoides as new genera to which names would be given later; in 1773 he named the first Kyllinga, the second Fuirena. Swartz thus acted rightly according to the procedure of his times in replacing Petesioides by Wallenia and Jacquin evidently approved Swartz’s action, for he himself adopted the name Wallenia in his Plantarum variorum Horti Caesaret Schoenbrunnensis Descriptiones 1 : 13, t. 30 (1797). Valid publication of ‘‘ Petesiodes Jacq.’”’ as a real generic name dates from Otto Kuntze’s adoption of it in 1891 (Revis. Gen. Pl. 2 : 402).
In 1788 Swartz named, as Samara coriacea (Nov. Gen. & Sp. Pl. : 32), a third
A SYNOPSIS OF JAMAICAN MYRSINACEAE 149
Jamaican member of the family, referring it to the Linnaean genus Samara, which is now included in Embelia Burman f. This species, found “in sylvis montium Jamaicae australis ’’, is congeneric with Rapanea guianensis Aublet, as Swartz noted, but Rapanea itself is not well separated from Myrsine L.
The collections including Myrsinaceae made in Jamaica during the first half of the nineteenth century by Macfadyen, Alexander, McNab, Nathaniel Wilson, March, and Purdie and used by Grisebach in preparing his Flora of the British West Indian Islands : 392-397 (1861) provided the material for publication there of Wadllenia clustifolia, W. venosa, Ardisia xylosteoides, and A. clustoides, and the record of A. humilis (actually A. solanacea) as naturalized in Jamaica. Grisebach’s book (cf. Stearn, 1965) seems to have given the impression for many years that no further collecting need be done in the West Indies. William Harris’s collecting from 1894 onwards brought about a new era in the botanical exploration of Jamaica, revealing the existence there of a multitude of new species which had escaped the notice of earlier collectors (cf. Urban, Symb. Antill. 6 : 70-86 (1909)). In 1895 Krug & Urban, working out Harris’s collections from the Blue Mountains, Jamaica, published two further species, Myrsine acrantha and Ardisia densiflora. Mez in 1901 recorded from Jamaica four species of Ardisia, A. harrisiana being proposed as new, eight species of Wallenia, and four of Rapanea. At the same time he defined more satisfactorily the genera, by transferring to Wallenia some species which Grisebach had included in Ardisia, although he separated Rapanea from Myrsine, reducing the latter to four African and Asiatic species typified by M. africana L.
The acquisition of new Jamaican material, mostly collected by Harris and by N. L. Britton of the New York Botanical Garden, and the re-study of the older material seen by Mez led Urban between 1908 and ro15 to describe as new Ardisia dictyoneura, A. troyana, Wallenia calyptrata, W. corymbosa, W. discolor, W. elliptica, W. erythro- carpa, W. punctulata and W. sylvestris and Britton to describe Petesioides sub- verticillata (now Wallenia subverticillata). Urban’s descriptions are wonderfully accurate, clear and detailed and are accompanied by helpful critical notes in which he made the most of inadequate material, but as usual he never provided a key to aid their recognition. The construction of such a key has indeed proved very difficult, partly because some taxa are only known in one of their three possible states—thus of the 16 species of Wallenia distinguished here, only six have been collected in their staminate, pistillate and fruiting states—but also because of the variation among these plants, which makes every gathering seem a little different from other gatherings. This is discussed further below under Wallenza.
GEOGRAPHICAL DISTRIBUTION
Four, if not five, of the Jamaican members of the Myrsinaceae occur outside the island. These are Ardisia compressa, which extends from Mexico to northern South America, Myrsine coriacea, which extends from Cuba over Jamaica, Hispaniola, Puerto Rico and the Lesser Antilles, and from Mexico over Central America, to Bolivia and Argentina, Wallenia laurifolia, which occurs also in Cuba and Hispaniola, and W. subverticillata, which also occurs in Cuba. Ardisia densiflora is closely allied
150 A SYNOPSIS OF JAMAICAN MYRSINACEAE
to A. spicigera of Mexico and A. gentle: of British Honduras; indeed, according to Lundell (1966a) the three are conspecific. The ornamental Asiatic species A. solanacea (Text-fig. 8) has become naturalized in several places. All the others appear to be endemic to Jamaica, although Myrsine acrantha may probably be found on other islands; specimens of it are likely to be referred to Myrsine (Rapanea) guianensis. Members of the family may be found in Jamaica at any altitude from sea-level to the top of Blue Mountain Peak (7,400 ft., 2,250 m.).
Although it would be premature to define types of distribution within Jamaica on the basis of present material, this is adequate to indicate relative ranges. Thus Myrsine acrantha (Text-fig. 20), M. coriacea (Text-fig. 19), Arvdisia tinifolia (Text- figs. 1-4), Wallenta laurifolia (Text-fig. 17), and W. venosa (Text-fig. 14) have comparatively wide ranges extending both east and west. A few are localized in the east, i.e. Avdisia brittoni: (Text-fig. 5), Wallenia sylvestris (Text-fig. 18), and W. subverticillata (Text-fig. 9). Some others are restricted to the higher parts of the Blue Mountains, i.e. Avdisia densiflora (Text-fig. 6), Wallenia calyptrata (Text-fig. 13), W. crassifolia (Text-fig. 10), and W. fawcetti (Text-fig. 16). As far as is known, Ardisia byrsonimae (Text-fig. 5), A. urbanit (Text-fig. 5), Wallenta corymbosa (Text- fig. 11), W. elliptica (Text-fig. 9), and W. punctulata (Text-fig. 13) are restricted to the middle of the island. Avdisia dictyoneura (Text-fig. 6), Wallenia grisebachu (Text- fig. 15), and W. purdieana (Text-fig. 16) occur both in the middle and west, Ardisia compressa (Text-fig. 7), Wallenia clusioides (Text-fig. 9), and W. erythrocarpa (Text- fig. 12) only in the west. As there has obviously been no bias in making the collec- tions here analysed, the species such as Arvdisia brittoni (Text-fig. 5), A. byrsonimae (Text-fig. 5), A. urbanii (Text-fig. 5), Wallenia discolor, W. elliptica (Text-fig. 9), W. erythrocarpa (Text-fig. 12), and W. punctulata (Text-fig. 13), which are known only from single gatherings or from gatherings made at the same place, must be of very limited range.
Specimens are cited below under Jamaican parishes, proceeding usually from west to east or from north to south within the parish.
FAMILY CHARACTERS
The family Myrsinaceae as represented in the West Indies consists of small trees or shrubs, with the leaves usually alternate, rarely whorled, and without stipules. The flowers are regular, bisexual or unisexual, with parts in fours or fives, and are grouped in axillary or terminal racemes or panicles sometimes condensed into fascicles or umbels. The calyx is persistent, with segments free or basally connate, imbricate or valvate. The corolla is usually gamopetalous but in Myrsine acrantha parted to the base. The stamens are inserted on the corolla, opposite to and the same number as the lobes of the corolla, with the anthers opening introrsely by lengthwise slits or apical pores. The ovary, superior in West Indian genera, is one-chambered with a free central placenta, into which the few or many ovules are sunk, appearing as slight or distinct bulges on its surface; the style may be short or long, with the stigma undivided or lobed or fimbriate. The fruit in West Indian genera is a more or less globose drupe. The family as a whole includes about goo
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156 A SYNOPSIS OF JAMAICAN MYRSINACEAE
species and occurs in the tropics and sub-tropics of the Old and the New World. Of its 30 or so genera, nine are represented in the West Indies as a whole but only three in Jamaica. Those absent from Jamaica are Parathesis (Cuba, Hispaniola, Puerto Rico), Stylogyne (Lesser Antilles), Conomorpha (Lesser Antilles), Weigeltia (Lesser Antilles), Cybianthus (Trinidad) and Grammadenia (Puerto Rico, Lesser Antilles).
KEY TO JAMAICAN GENERA
Flowers in usually terminal (sometimes axillary and then long-pedunculate) racemes or panicles; style slender; stigma inconspicuous.
Flowers bisexual, with stamens and style both well developed in the same flower; corolla c. 6-12 mm. long; ovules numerous, scattered in several series over the placenta. ; ‘ 1. Ardisia
Flowers unisexual, dioecious, the male flowers with well developed exserted stamens, the female flowers with abortive included stamens but exserted styles; corolla 1-5-6 mm. long; ovules 3-4 in a single horizontal row on the
placenta . ; , . 2. Wallenia Flowers in umbellate clusters on very short, scaly axillary spurs; style short or none (in American species) ; stigma conspicuous : : , F 3. Myrsine
t. ARDISIA Swartz
ARDISIA Swartz, Nov. Gen. & Sp. Pl. : 3, 48 (1788); Fl. Ind. Occ. 1 : 467 (1797); nomen conservandum No. 6285. Lectotype: A. tinifolia Swartz.
KEY TO JAMAICAN SPECIES
Inflorescences all axillary; pedicels 1-4 cm. long, clustered almost umbellately at the end of the peduncle; anthers 5-6 mm. long. : . 8.A. solanacea
Inflorescences terminal (sometimes with a few axillary inflorescences below) ; pedicels mostly 1-7 mm. long, sometimes to 12 mm. long but then racemosely armangeg along the lateral peduncle; anthers 1-5-5 mm. long:
Lateral peduncles compound, i.e. bearing 3 to 4 secondary peduncles each umbellately 2-6-flowered; leaves 7-16 cm. long, the blades narrowly obovate to oblanceolate; filaments of stamens 0-5 mm. long . . 7. A. compressa
Lateral peduncles simple, i.e. directly bearing 3 to 28 pedicelled flowers but no secondary peduncles; leaves mostly less than 12 cm. long, the blades various; filaments of stamens I-5 mm. long:
Leaves closely clustered at the tips of shoots, less than 5-5 cm. long; inflorescence contracted, less than 4 cm. long and broad, the branches 3-6-flowered, the pedicels 1-4 mm. long; central Jamaica ; ; . 6. A. byrsonimae
Leaves more widely spaced, some or all more than 5-5 cm. long; inflorescence more than 4 cm. long and broad, the branches usually many-flowered, the pedicels 2-15 mm. long:
A SYNOPSIS OF JAMAICAN MYRSINACEAE 157
Inflorescence somewhat fastigiate, the densely many-flowered branches ascending at c. 40°; corolla c. 5 mm. long; anthers c. 1-5 mm. long, much shorter than the filaments; Blue Mountains : . 5.A. densiflora
Inflorescence looser, the branches spreading at c. 50-60°; corolla 7-12 mm. long; anthers 2-5-5 mm. long, equalling or longer than the filaments:
Pedicels 1-4 mm. (rarely to 5mm.) long; leaf-blades with well-marked reticulate venation : 4. A. dictyoneura Pedicels 4-15 mm. long; leaf blades not ‘markedly reticulate: Sepals 2-5-4 mm. broad: Anthers 4:5-5 mm. long, apically dehiscent; sepals almost or quite
glabrous; eastern Jamaica ‘ . 2, A. brittoni Anthers 3-3°5 mm. long, laterally dchiccent atest to the base; sepals ciliate; western Jamaica. : : : : a. At urbanii
Sepals 1-2 mm. broad: Sepals ovate or broadly ovate: Anthers 3-3-5 mm. long. ; : . I. A. tinifolia var. I Anthers 4-5 mm. long : : A. tintfolia var. 2 Sepals mostly narrowly ovate or oblong-lanceolate: Leaf-blades mostly less than 6 cm. long; sepals 2-2-5 mm. long A. tinifolia var. 3
Leaf-blades 7-12 cm. long; sepals 2-5-3 mm. long A. tintfolia var. 4
Subgenus r. ARDISIA
ARDISIA subgenus ARDISIA; subgenus typicum generis.
Ardisia Swartz, Nov. Gen. & Sp. Pl. : 3, 48 (1788), sensu stricto. Pickeringia Nutt. in Journ. Acad. Sci. Philad. 7 : 95 (1834). Ardisia sect. Euardisia Endl., Gen. Pl. : 736 (1839). Ardisia subgenus Pickeringia (Nutt.) Mez in Urban, Symb. Antill. 2 : 396 (1901); in Engler, Pflanzenr. IV. 236 : 79 (1902). Lectotype: A. tinifolia Swartz. Inflorescence always terminal; branches simply racemose. Stamens with con-
spicuous filaments. Species all American.
I. Ardisia tinifolia Swartz, Nov. Gen. & Sp. Pl. : 48 (1788); Fl. Ind. Occ. 1 : 468 (1797).—Griseb., Fl. Brit. W. Ind. Is. : 396 (r861r).—Mez in Urban, Symb. Antill. 2 : 400 (rg01); in Engler, Pflanzenr. IV. 236 : 83, fig. rr F—K (1902). (Text-fig. 23G, H; pl. 6, 7.)
Ardisia coriacea Swartz, Nov. Gen. & Sp. Pl. : 48 (1788); Fl. Ind. Occ. 1 : 470 (1797). Ardisia harrisiana Mez in Urban, tom. cit. : 401 (1901); in Engler, tom. cit. : 83 (1902). Ardisia troyana Urban, Symb. Antill. 5 : 455 (1908). “* Jasminum forte, arboreum, foliis laurinis obtusis latioribus atrovirentibus, flore pentapetalo racemoso purpureo reflexo ’’ Sloane, Cat. Pl. Jam.: 169 (1696); Voy, Jam. Nat. Hist. 2: 98, t. 205 fig. 2 (1725). The species Ardisia tinifolia as here accepted includes plants diverging in form and size of leaf, in number of flowers, shape of sepals and length of anthers. Selected specimens, e.g. Harris 8741 (type-collection of A. troyana) from near Troy, Central
158 A SYNOPSIS OF JAMAICAN MYRSINACEAE
Jamaica, with very small leaves, Proctor 16621 from Orange River Valley, St. James, with comparatively elongated sepals, and Loveless 1198 from the Long Mountain, near Kingston, with long anthers, could reasonably be regarded as exemplifying independent species and were indeed treated by the writer as such until the difficulty of finding stable combinations of characters and of allocating specimens intermediate in one character or another to these supposed taxa made it more practical to place them all in one species, within which four major populations (Text-figs. 1-4) are more or less distinguishable though not easily defined. They are simply given numbers below. Swartz’s original diagnosis of his A. tinifolia in 1788 is brief but the more detailed description in his Flora Indiae Onentalis, and his unpublished illustration (Pl. 6, see p. 147) and authentic specimens (BM) clearly establish the identity of the plant he had in mind. A representative modern gathering is Harris 12087, collected between Constant Spring and Bardowie, St. Andrew, with veining on the lower surface of the leaf plainly evident. It typifies the population (var. I below) found in the southern parishes of Jamaica, with an altitudinal range of 50 ft. (15 m.) to 3,500 ft. (1,070 m.). Here belongs Swartz’s A. coriacea with veining on the lower leaf-surface faint or not evident, likewise represented by an unpublished drawing (Pl. 7). Diverging from typical A. tinifolia principally by having somewhat longer anthers is the population (var. 2 below) characteristic of the Long Mountain near Kingston, at 500 ft. (150 m.) to 1,000 ft. (300 m.). Small short-petioled leaves evidently veined characterize plants of central Jamaica (var. 3 below) named A. troyana by Urban, which also differ from typical A. tinifolia in having shorter branches of the inflorescence. These occur at 700 ft. (210 m.) to 2,500 ft. (750 m.). Their range slightly overlaps that of a population (var. 4 below) with larger longer-petioled leaves, looser many-flowered inflorescences and slightly shorter filaments, found in the northern parishes of Jamaica from almost sea-level to 700 ft. (210 m.). They occur in the same area at Sherwood Content, Trelawny, and merit particular study there in the field.
Together the plants here placed in Ardisia tinifolia thus give the impression of a population within which morphologically and geographically differentiated taxa have been formed but have not achieved complete independence.
Var. 1.
A. tinifolia Swartz, loc. cit. (1788). A. coriacea Swartz, loc. cit. (1788). A. harrisiana Mez, loc. cit. (190T).
Leaf-blades mostly elliptic, sometimes narrowly obovate, the apex obtuse or acute, 5-12 cm. long, 1-5-7 cm. broad. Sepals imbricate, ovate or broadly ovate, rounded, c. 1:5-2:5 mm. long, 1:2-2 mm. broad. Anthers 3—3:5 mm. long.
St. ELIZABETH: Proctor 15419 (BM; IJ), Proctor 11344 (BM; IJ).
CLARENDON: Proctor 22739 (BM; IJ), Proctor 9712 (IJ), Proctor 10233 (IJ), Stearn 1044 (BM), Adams 8424 (UCWI).
St. ANN: Alexander (K).
ST. CATHERINE: Proctor 9310 (BM; IJ), Proctor 8373 (IJ), Proctor 22739 (IJ).
St. ANDREW: Britton 3461 (NY), Stearn 59 (BM), Proctor 11428 (IJ), Harris 5970
A SYNOPSIS OF JAMAICAN MYRSINACEAE 159
(BM; NY), Harris 11137 (NY; UCWI), Newill 5449 (IJ), Harris 5926 (NY), Harris 12081 (BM; NY; UCWI), Harris 9033 (BM; NY; UCWI), Powell 850 (IJ), Proctor 23694 (IJ), Harris 6600 (BM; NY; UCWI: type-collection of Ardisia harrisiana Mez), Britton 2987 (NY), Proctor 24511 (BM), Proctor 23982 (BM).
St. THomas: Adams 12579 (UCWI).
Var. 2.
Leaf-blades elliptic or broadly elliptic, a few broadly obovate, the apex obtuse or rounded, 4-9 cm. long, 2-6cm. broad. Sepals imbricate, ovate or broadly ovate, c. 2-2°5 mm. long, 1:5-2 mm. broad. Anthers 4-5 mm. long.
KINGSTON and St. ANDREW: Britton 377 (NY), Robbins 1496 (UCWI), Proctor 7386 (IJ), Webster 4986 (BM), Loveless r1g8 (UCWI), Yuncker 17315 (BM), Anderson & others 626 (UCW]I).
Var. 3. Ardisia troyana Urban, loc. cit. (1908).
Leaf-blades elliptic, the apex obtuse, c. 3-10 cm. long, 1-5-6 cm. broad (usually less than 6cm. long, 3-5cm. broad). Sepals narrowly ovate, 2-2:-5 mm. long, I-1°5 mm. broad. Anthers 3—3°5 mm. long.
TRELAWNY: Harris 8741 (BM; NY; UCWI: type-collection of A. ¢voyana Urban), Britton 605 (NY), Harris 9473 (NY; UCWI), Howard, Proctor & Stearn 14677 (BM), Howard, Proctor & Stearn 14679 (BM; IJ), Proctor r1060 (BM; IJ), Proctor 22543 (BM; IJ), Proctor 24458 (BM; IJ), Proctor 25676 (IJ).
CLARENDON: Harris 12769 (NY; UCWI), Proctor 9772 (IJ), Proctor 10233 (IJ).
MANCHESTER: Proctor 25625 (BM; IJ).
Var. 4.
Leaf-blades narrowly obovate or elliptic, the apex acute or shortly acuminate, 5-12 cm. long, 2-6 cm. broad. Sepals scarcely or not imbricate, ovate to obovate- lanceolate, obtuse, c. 1:5-2°5 mm. long, 1-1-5 mm. broad. Anthers c. 3 mm. long.
HANOVER: Proctor 11280 (BM; IJ).
WESTMORELAND: Harris 10220 (BM; K; NY; UCW]).
St. JAMES: Harris 10318 (BM; NY; UCWI), Britton & Hollick 2355 (NY), Proctor 16621 (BM; IJ), Proctor 24292 (BM; IJ).
TRELAWNY: Proctor 11062 (BM; IJ).
St. Mary: Loveless 2437 (UCWI), Proctor 7546 (IJ), Yuncker 17838 (BM).
2. Ardisia brittonii Stearn, sp. nov. (Text-fig. 21; Pl. 8a)
Arbor parva ad 4:5 mm. alta, vamulis hornotinis primo ferrugineis pulverulentisque, annotinis nigrescentibus glabrisque. Folia dissita, c. 4-8 mm. inter se distantia, magna, patentia, longipetiolata; lamina oblongo-elliptica, apice obtusa, margine integra, basi cuneata in petiolum attenuata, 8-17 cm. longa, 3-8 cm. lata, glabra, utrinque punctata, coriacea, venis numerosis utrinque prominulis marginem versus reticulate anastomosantibus; petiolus 1-2:2cm. longus. Inflorescentiae terminales multiflorae laxae, c. 15cm. longae, 10cm. latae, supra glabrae, inferne minute
BOT. 4, 4 9
160 A SYNOPSIS OF JAMAICAN MYRSINACEAE
pubescentes, ramulis patentibus ad gcm. longis racemose 8—12-floris; pedicelli inferrores ad 1-5cm. longi. Calyx c. 4-5-5 mm. longus; sepala imbricata parum inaequalia, ovata vel late oblonga, rotundata, c. 4-4-5 mm. longa, 2-5-3 mm. lata, punctata, margine glabra. Corolla rosea, c. 1-3 cm. longa; petala in 1/4 altitudinis coalita, apice obtusa, parte libera oblonga c. 4mm. lata, per anthesin revoluta, nigro-punctata et lineolata. Stamina quam petala breviora, c. 1-5 mm. supra corollae basin inserta; antherae anguste triangulares, c. 4-5-5 mm. longae, apice obtusa rimis brevibus dehiscentes, basi cordatae; filamenta 3 mm. longa. Ovarium late ovoideum; stylus c. 9 mm. longus, stigmate minuto,
Fic. 21. Dissected flower of Ardisia brittonit Stearn (Harris & Britton 10558), X 3.
St. THomaAs: Bath to Cuna Pass, Sept., 1908, Harris & Britton 10558 (NY, holotype: BM; K; UCW]).
This species of eastern Jamaica has been named in honour of Nathaniel Lord Britton (1859-1934), founder of the New York Botanical Garden and author of many botanical works on the eastern United States and the West Indies; see National Acad. Sct. U.S.A. Biogr. Mem. 19 : 147-202 (1938) for a biography by E. D. Merrill and a bibliography by J. H. Barnhart.
Ardisia brittonw differs from the widespread A. tinzfolia principally in its broader sepals and in having somewhat larger flowers and longer anthers than is usual in A. tinifolia. It agrees more closely with A. urban of central Jamaica but may be distinguished from that by its glabrous calyx and its longer anthers, which dehisce only at the apices.
3. Ardisia urbanii Stearn, nom. nov. (Text-fig. 22.) Ardisia vosea Urban, Symb. Antill. 5 : 456 (1908); non A. vosea King & Gamble (1905).
TRELAWNY: Harris 9419 (BM; NY; type-collection), Harris 9418 (BM; NY; UCWI), Britton 659 (NY).
Ardisia urbanii is known only from collections made at Troy on the southern side of the Cockpit Country by Harris & Britton in 1906 at about 1,600 ft. (530 m.). The
A SYNOPSIS OF JAMAICAN MYRSINACEAE I61
size of the calyx (with sepals 3-5 mm. long, 2-5-4 mm. broad) distinguishes it from A. tinifolia; the ciliated sepals and the anthers dehiscent to the base distinguish it from A. brittonii of eastern Jamaica. According to Harris, it forms a small tree to 25 ft. (7-5 m.) high, with rosy flowers. It is named in honour of Ignaz Urban (1848- 1936), the author of many important works on the West Indian flora; see Ber. Deutsch. Bot. Ges. 48 : (205)-(225) (1931) for a biography and bibliography by Th. Loesener.
Fic. 22. Dissected flower of Avdisia urbanii Stearn (Britton 659), X 4.
4. Ardisia dictyoneura Urban, Symb. Antill. 6 : 28 (1909).
HANOVER: Harris 10345 (BM; NY; UCWI; type-collection), Proctor 11316 (BM; IJ), Britton 2277 (NY).
WESTMORELAND: Proctor 7321 (BM; IJ).
St. JAMES: Proctor 22979 (BM; IJ).
TRELAWNY: Proctor 10610 (BM; IJ).
MANCHESTER: Proctor 11028 (BM; IJ), Proctor 11036 (BM; IJ), Proctor 18299 (IJ), Stearn 351 (BM).
St. ANN: Purdie (K), Proctor 22835 (BM; IJ), Proctor 26724 (BM), Stearn 567 (BM), Proctor 11896 (IJ), Howard & Proctor 15039 (IJ), Proctor 7494 (IJ).
St. CATHERINE: Howard & Proctor 13620 (IJ; NY).
Ardisia dictyoneura is a species of western and central Jamaica at 1,000—3,000 ft. (300-g00 m.). The type collection was made by ‘Harris in 1908 at Fray Woods in Hanover. It has often been collected in fruit, only twice in flower, and is notable for its profusely and finely reticulate-veined coriaceous mostly obovate or narrowly obovate leaves and its comparatively large fruits, 6-5-8 mm. in diameter when mature, borne on pedicels only 1-4 mm. long.
BOT. 4, 4 9§
162 A SYNOPSIS OF JAMAICAN MYRSINACEAE
5. Ardisia densiflora Krug & Urban in Notizbl. Bot. Gart. & Mus. Berlin r : 79 (x895).—Urban, Symb. Antill. 1 : 385 (1899); op. cit. 6 : 28 (r1909).—Mez in Urban, Symb. Antill. 2 : 398 (1901); in Engler, Pflanzenr. IV. 236 : 82 (1902).— C. L. Lundell in Fieldiana, Bot. 24 (8) : 141 (1966). (Text-fig. 23F.)
Ardisia spicigera Donn. Smith in Bot. Gaz. 27 : 434 (1899), fide Lundell, loc. cit. (1966). Ardisia gentlei C. L. Lundell in Field & Lab. 13 : 11 (1945), fide Lundell, loc. cit. (1966).
St. ANDREW: Harris 5227 (K; UCWI), Harris 5431 (BM; K; type collection), Harris 7657 (BM; K; NY; UCWI), Harris 6578 (BM; UCWI), Harris 10029 (BM; NY; UCWI), Harris 6077 (NY ; UCWI), D. Watt 8225 (NY; UCWI), Maxon & Killip 1343 (NY), Proctor 24511 (BM; IJ), Proctor 25610 (BM; IJ).
The discovery in 1894 of Avdisia densiflora in the Blue Mountains of Jamaica, on which it grows at about 3,000—4,000 ft. (g00—1,200 m.), was one of the early results of the renewed collecting in Jamaica at the end of the nineteenth century. It isa well-marked species with a densely many-flowered inflorescence, the branches of which are ascending, almost fastigiate, and which bear up to 25 small white flowers on pedicels r-6 mm. long. The filaments of the stamens are much longer than the longitudinally dehiscent anthers. According to Lundell, A. spicigera from Mexico and A. gentle: from British Honduras are both conspecific with A. densiflora, which thus has a fairly wide but sparse Central American distribution comparable with that of Achimenes erecta (Lam.) H. P. Fuchs, a gesneriad restricted in Jamaica to the Blue Mountains.
6. Ardisia byrsonimae Stearn, sp. nov. (Text-fig. 23A—E; Pl. 8b).
Arbor parva ad 4:5m. alta, vamulis hornotinis ferrugineis pulverulentisque, annotinis cinereis glabrisque. Folva pro genere parva, ad apices ramulorum conferta, I-5 mm. inter se distantia, ascendentia, brevipetiolata; Jamina obovata, apice rotundata interdum emarginata, margine integra, basi cuneata, c. 3-5-5 cm. longa, I*3-3'5 cm. lata, glabra, utrinque punctata, coriacea, venis vix prominulis; petzolus 2-5mm. longus. IJnflorescentiae terminales fere sessiles multiflorae densae, c. 3 cm. longae et latae, primo pulverulentae, ramulis ascendentibus ad 3 cm. longis racemose 3-6-floris; pedicelli 1-4 mm. longi. Calyx c. 3 mm. longus; sepala imbricata, ovata, rotundata, c. 2-2-5 mm. longa, 1-5-2 mm. lata, margine scariosa ciliataque. Corolla rosea, c. 7-8 mm. longa; pefala in 1/4 altitudinis coalita, apice acuta, parte libera lanceolata, 2—2:5 mm. lata, per anthesin recurvata, punctata. Stamina quam petala breviora, c. I°5-2 mm. supra corollae basin inserta; antherae anguste triangulares, c. 3-5°5 mm. longae, apice mucronatae et rimis brevibus quasi poratim dehiscentes, basi cordatae; filamenta c. 2mm. longa. Ovarium late ovoideum; stylus c. 8-9 mm. longus, stigmate minuto. Fructus transverse late ellipsoideus, c. 4-5 mm. altus, 5-6 mm. latus, glanduloso-punctatus, styli basi c. r mm. longa coronatus.
CLARENDON: Peckham Woods, 760 m., Dec. 1917, Harris 12799 (BM; NY; UCWI), same locality, Aug. 1954, Webster & Proctor 5428 (BM), same locality, Jan. 1955, Proctor 9743 (BM; IJ), same locality, Dec. 1955, Stearn g (BM, holotype; K), Proctor 11408 (IJ).
A SYNOPSIS OF JAMAICAN MYRSINACEAE 163
Fic. 23. Avrdisia byrsonimae Stearn: A, fruiting specimen (Stearn 9), x 4; B, buds (Proctor 9743), X 2; C, dissected flower (Stearn 9), xX 3; D, anther (Stearn 9), x 5; E, fruit (Stearn 9), X 4. Ardisia densiflora Krug & Urban: F, dissected flower (Harris 10029), x 4. Ardisia tinifolia Swartz; G, dissected flower (Harris 12081), x 3; H, placenta with ovules (Harris 12081), x 20. Ardisia solanacea Roxb.; I, dissected flower (Harris 11637), calyx and ovary x 1I+5, corolla and stamens x 2.
Ardisia byrsonimae is a slow-growing tree endemic to wooded limestone hill-tops in upper Clarendon, Jamaica. It differs from other Jamaican species in its compara- tively small leaves crowded at the ends of the shoots and its contracted inflorescences with very short pedicels. From A. tinifolia var. 3 (A. troyana Urban), likewise inhabiting the western interior of Jamaica, it may also be distinguished by its
164 A SYNOPSIS OF JAMAICAN MYRSINACEAE
markedly imbricate and more rounded sepals. The epithet byrsonimae was given to this species by N. L. Britton under Jcacorea but never published by him. It refers to a resemblance in habit between it and certain West Indian species of Byrsonima (Malpightaceae), notably B. lucida (Mill.) Rich., with small obovate leaves usually clustered at the ends of the shoots.
Subgenus 2. ICACOREA (Aublet) Pax
ARDISIA subgen. IcACOREA (Aublet) Pax in Engler & Prantl, Nat. Pflanzenfam. 4 (I): 94(1889).—Mez in Urban, Symb. Antill. 2 : 391(r907).
Icacorea Aublet, Hist. Pl. Guiane Franc. 2, Suppl. : 1 (1775). Ardisia sect. Icacorea (Aublet) Griseb., Fl. Brit. W. Ind. Is. : 395 (1861).
Type (by monotypy): Avdisia guianensis (Aublet) Mez (Icacorea guianensis Aublet).
Inflorescences terminal or both axillary and terminal; branches compoundly racemose, bearing several umbellate or corymbose clusters. Stamens with short filaments. Species all American.
7, Ardisia compressa Kunth, Nov. Gen. & Sp. Pl. 3 : 245 (1819).—Mez in Urban, Symb. Antill. 2 : 393 (1901); in Engler, Pflanzenr. IV. 236 : 89 (1902).—C. L. Lundell in Fieldiana, Bot. 24 (8) : 139 (1966).
Ardisia decipiens sensu Griseb., Fl. Brit. W. Ind. Is. : 395 (1861); non A. DC. Icacorea compressa (Kunth) Standley in Contrib. U.S. Nat. Herb. 23 : 1110 (1924).
HANOVER: Proctor 10013 (BM; IJ), Harris 10253 (BM; K; NY), Britton & Hollick 2129 (NY), Britton & Hollick 2164 (NY).
WESTMORELAND: Proctor & Mullings 22060 (BM; IJ).
St. JAMES: Proctor 16166 (BM; IJ).
Ardisia compressa ranges from Mexico to northern South America, including Trinidad, but is not recorded from any other West Indian island. Humboldt and Bonpland collected the type near Caripe, Venezuela. Most continental specimens have the apex of the leaf more gradually tapered than have Jamaican specimens, which approximate most to some from British Honduras. The Jamaican specimens are all from the west of the island. The British Museum (Natural History) possesses an unlocalized specimen from Jamaica collected by William Wright who was in Jamaica in 1765-76 and 1782-85, living most of the time at Hampden, in Trelawny, and Mez suggests that this was of cultivated origin, which, however, seems unlikely. It is easily distinguished from other Jamaican species by its more intricately branched inflorescence, the lateral peduncles of which bear three or four secondary peduncles each with two to six small, whitish, brown-flecked flowers; the filaments of the stamens are about 0:5 mm. long, and much shorter than the anthers.
Subgenus 3. TINUS Mez
ARDISIA subgen. Tinus Mez in Engler, Pflanzenr. IV. 236 : 124 (1902). Lectotype: Ardisia elliptica Thunb. (A. humilis sensu Mez, pro parte; non Vahl).
A SYNOPSIS OF JAMAICAN MYRSINACEAE 165
Inflorescences axillary. Stamens with very short filaments. Species Asiatic and Australian, one often naturalized elsewhere.
8. Ardisia solanacea Roxb., Pl. Coast Coromand. I : 27, t. 27 (£795).—Sims in Curt. Bot. Mag. 40 : t. 1677 (1814).—Mez in Urban, Symb. Antill. 2 : 402 (IgoT); in Engler, Pflanzenr. IV. 236 : 132 (1902).—E. H. Walker in Philipp. Journ. Sci. 73 : 58 (1940). (Text-fig. 23 I.)
Ardisia humilis sensu A.DC. in DC., Prodr. 8 : 129 (1844); non Vahl.—Wight, Ic. Pl. Ind. Br. 4 : t. 1212 (1848).—Griseb., Fl. Brit. W. Ind. Is. : 396 (1861).—C. B. Clarke in Hook.
f., Fl. Brit. Ind. 3 : 529 (1882).—De Wild., Ic. Select. Hort. Thenen. 1 : 79, t. 19 (1900). Icacovea solanacea (Roxb.) Britton, Fl. Berm. : 284 (1918).
ST. JAMES: Proctor 16461 (BM; IJ).
St. Mary: Harris 11637 (NY; UCWI), Proctor 20615 (BM; IJ), Adams 12076 (UCW]).
St. ANDREW: Grabham (NY), Adams 10094 (BM).
PoRTLAND: Powell 717 (BM; IJ; NY), Proctor 16546 (BM; IJ; NY), Proctor 19723 (BM; IJ; NY), Marble 852 (NY), Yuncker 18826 (BM), Adams 11482 (BM; UCWI).
St. THomas: Nichols 191 (NY), Powell 854 (IJ), Harris & Britton 10585 (NY; UCWI), Proctor 9217 (BM; IJ), Webster & Wilson 5218 (BM; IJ), Howard, Proctor & Stearn 14809 (BM; IJ), Proctor 16450 (BM; IJ), Yuncker 17523 (BM), Yuncker 18826 (BM; UCW]), Harris 6040 (NY), Britton 3555 (NY), Adams ro471 (UCWI).
A native of India, Malaya and China, long cultivated in tropical gardens, this has become naturalized here and there in Jamaica, usually along streams or on shaded road banks, and is sometimes called “ Blackberry” or “ Craingcraing’’. It is easily recognized by its axillary few-flowered inflorescences with comparatively large pale rose flowers clustered almost umbellately at the end of the long peduncle.
2. WALLENIA Swartz
WALLENIA Swartz, Nov. Gen. & Sp. Pl. : 2, 31 (1788); Fl. Ind. Occ. 1 : 247 (1707); nomen conservandum No. 6304.
Petesioides Jacq. ex Kuntze, Revis. Gen. Pl. 2 : 402 (1891), nom. superfl.
Type (by monotypy): W. laurifolia Swartz (Petesioides laurifolium Jacq.)
Wallenia belongs to the tribe Myrsineae characterized by having the ovules few and arranged in a single series around the placenta. Both it and Myrsine can be readily distinguished by their small flowers, with the corolla 1-5-6 mm. long, from Ardisia of the tribe Ardisieae, characterized by having numerous ovules arranged in several series or apparently scattered over the placenta. The loose many-flowered inflorescence of Wallenia readily distinguishes it from Myrsine (Rapanea) which has flowers crowded in axillary clusters. The Jamaican species all belong to the sub- genus Wallenia (subgenus Euwallenia Mez) which is confined to Jamaica, Cuba and Hispaniola and has an erect terminal paniculate inflorescence; the other subgenus, Homowallenia Mez, with pendulous axillary racemose inflorescences, occurs in Cuba, Hispaniola, Puerto Rico, and the Lesser Antilles but not in Jamaica.
166 A SYNOPSIS OF JAMAICAN MYRSINACEAE
The dioecious nature of Wallenia possibly militates against the formation of well- defined taxa within the genus; it certainly makes difficult the correlation of characters in herbarium material. The corolla of male flowers is much longer than the calyx; the anthers are borne on long exserted filaments. The corolla of female flowers is shorter or little longer than the calyx; the filaments are very short and the anthers aborted. Consequently female flowers are much less conspicuous than male flowers and are liable to be passed over by collectors as not being fully developed. Thus there is rarely available, from the same locality, male and female flowering material and mature fruiting material obviously belonging to the same species, which would provide the means of correlating other material. Instead some members of the genus are only known in one of these three possible states. The variability of the genus, presumably maintained by “ out-breeding”’, is such that two gatherings rarely match completely. Mez and Urban accordingly tended to describe each new gathering as representing a new species distinguished from its congeners by a com- bination of subtle rather than well-marked characters, the latter being few within the subgenus Wallenia (Euwallenia Mez). The lack of comparable material makes it difficult to assess the worth of such characters. In preparing the present account it has not been possible to devise a key making full use of floral characters and unfortu- nately the vegetative characters used tend either to overlap or to be difficult to express without appearing too definite and exclusive or too vague and hence useless. Thus while the key should lead to the identification of typical material of the species, it would be too optimistic to expect conclusive results with all specimens. Wallenia laurifolia occurs in Cuba, Hispaniola and Jamaica, W. subverticillata in Cuba and Jamaica; the other species accepted here are apparently endemic to Jamaica.
KEY TO JAMAICAN SPECIES
Leaves with base broadly rounded and slightly cordate, paired or in whorls of three together, fairly large (3-17 cm. broad), the petiole short (less than 6 mm. long)
or lacking:
Leaves rounded at the apex, 7-17 cm. broad: Leaves sessile. ; : : : . I. W. clusiotdes Leaves with 3-6 mm. long petiole ; . 2. W. elliptica
Leaves narrowed towards the acute or obtuse apex, 3-6 cm. broad 3. W. subverticillata Leaves with base cuneate or attenuate, rarely rounded (the petiole then I cm. or more long), usually alternate, only occasionally paired or whorled, the petiole short or long: Leaves reticulate-veined beneath, with slightly raised cross-veins between the main lateral veins: Petiole short, c. r-8 mm. long; blades under 11 cm. long: Leaves mostly broadest at the middle and gradually narrowed from there to the apex: Petiole very short, 1-4 mm. long; blade inconspicuously punctate beneath 4. W. fawcettit
A SYNOPSIS OF JAMAICAN MYRSINACEAE 167
Petiole longer (5-8 mm. or more long); blade conspicuously punctate beneath ; . 10. W. venosa Leaves mostly broadest above the middie with the apex rounded or obtuse: Leaf-blades 2—8 cm. long, less than 3:5 cm. broad; crest of Blue Mountains 5. W. crassifolia Leaf-blades larger (5-11 cm. long) : Leaves coriaceous, firm, greyish or glaucescent; female calyx c. I-4¢ mm. long . ; k : 6. W. corymbosa Leaves chartaceous, thinner: Leaves drying light green; female calyx c. 1-5 mm. long rr. W. purdieana Leaves drying brownish; female calyx 1 mm. long 7. W. xylosteoides Petiole longer (c. 8-30 mm. long); blades mostly more than 11 cm. long: Leaf-blades nearly three times as long as broad, mostly narrowly elliptic: Base of blade narrowly cuneate or attenuate . . 8. W. erythrocarpa Base of blade rounded or abruptly contracted . 12. W. grisebachu Leaf-blades about twice as long as broad, mostly elliptic or narrowly obovate: Lateral veins spreading outwards from the midrib at an angle of about 40° 9. W. punctulata Lateral veins spreading outwards from the midrib at an angle of about 50-70°: Male flower with calyx 1-5-2°5 mm. long, corolla 2-5-4 mm. long; persis- tent fruiting calyx not more than 1-5 mm. long: Leaves usually densely blackish-punctate and somewhat mottled beneath, strongly veined, the base cuneate . 0. W. venosa Leaves inconspicuously punctate and not mottled beneath, lightly veined, the base cuneate or rounded or abruptly contracted: Leaves drying light green; inflorescence glabrous, loose; male flower with corolla yellowish, not punctate; style about 0-8 mm. long 11. W. purdieana Leaves drying brownish; inflorescence minutely pubescent, compact ; male flower with corolla reddish, distinctly punctate; style
about 15mm. long. ; 12. W. grisebachu Male flower with calyx c. 3 mm. iene eoroile 5-6 mm. long; persistent fruiting calyx about 2:5 mm. long , ; 13. W. calyptrata
Leaves pinnately veined beneath, i.e. without raised cross-veins between the main lateral veins: Blades of some or all leaves abruptly contracted at base into the petiole; inflorescence usually minutely pubescent .. 12. W. grisebachit Blades of all leaves gradually narrowed into the petiole; inflorescence glabrous: Blades of leaves mostly less than 12 cm. ae 6 cm. broad; petiole not more than I cm. long i 14. W. laurifolia Blades of leaves up to 25 cm. long, 1 EL-crn, broad, mostly more than 12 cm. long, 6 cm. broad; petiole 1-2 cm. long:
168 A SYNOPSIS OF JAMAICAN MYRSINACEAE
Fruits vertically ribbed ‘ 15. W. sylvestris Fruits not ribbed ; : : : : ‘ . 16. W. discolor
1. Wallenia clusioides (Griseb.) Mez in Urban, Symb. Antill. 2 : 4rr (rgor); in Engler, Pflanzenr. IV. 236 : 245 (1902).
Ardisia clusioides Griseb., Fl. Brit. W. Ind. Is. : 396 (1861) excl. specim. Alexander,
HANOVER: Harris 10333 (BM; NY), Proctor 10028 (IJ), Proctor 26590 (IJ).
WESTMORELAND: Purdie (K, lectotype), Harris 10210 (BM; NY).
ST. JAMES: Harris 9177 (BM; NY).
ST. ELIZABETH: Harris 9761 (NY), Britton 1110 (NY), Proctor 26530 (IJ).
W. clusioides is a species of western Jamaica notable for its very large sessile and subcordate leaves 15-31 cm. long, 7-17cm. broad. It forms a small tree or tall shrub 6~20 ft. (2-6m.) high. The female flowers are unknown. The greenish- yellow male flowers have the calyx 1-5-2 mm. long, the corolla about 2-5 mm. long.
2. Wallenia elliptica Urban, Symb. Antill. 6 : 30 (1909). (PI. ga.)
Ardisia clusioides Griseb., Fl. Br. W. Ind. Is. : 396 (1861) pro parte, quoad specim. Alexander. St. ANN: Alexander 569 (K; NY; type-collection), Britton & Hollick 2776 (NY). An obscure species described by Urban on a specimen collected by Alexander at Grierfield near Moneague, St. Ann, W. elliptica is provisionally kept apart from W. clustoides on account of the shortly petioled leaves. The male flowers are unknown. The female flowers have the calyx and corolla about 1-5 mm. long.
3. Wallenia subverticillata (Britton) Ekman ex Urban, Symb. Antill. 9 : 412 (1925).—Alain in Leén & Alain, Fl. Cuba 4 : rrr (1957). (Text-fig. 24K.)
Petesioides subverticillatum Britton in Bull. Torrey Bot. Club 37 : 355 (1910).
PORTLAND: Proctor 22102 (IJ).
ST. THomas: Britton 3937 (NY), Harris & Britton 10694 (NY, holotype; UCWI), Harris & Britton ro7or (K; NY), Britton 4049 (NY), Proctor 28681 (BM).
Wallenia subverticillata, a species of mountain woods at 1,000-2,000 ft. (300— 600 m.), is distinguished by having its leaves grouped in pairs or threes; they are almost sessile and are much narrower and more attenuate towards the apex than those of W. clusioides but are nevertheless extremely variable in shape and size, e.g. 5-20 cm. long, 3-5 cm. broad on the same plant. The type was collected by Harris and Britton on a slope below Big Level, eastern St. Thomas. Rather surprisingly, specimens of what appear to be the same species have been collected in Las Villas, Cuba, by Ekman. The calyx of male flowers is 2-2-5 mm. long, of female flowers about r mm. long; the corolla of male flowers is about 2-5 mm. long, of female flowers about r mm. long, slightly longer than the calyx.
4. Wallenia fawcettii Mez in Urban, Symb. Antill. 2 : 408 (190r); in Engler, Pflanzenr. IV. 236 : 244 (1902). (PI. gb.)
A SYNOPSIS OF JAMAICAN MYRSINACEAE 169
PorRTLAND: Maxon & Killip 714 (NY).
PoRTLAND or ST. THOMAS: Harris 5422 (NY, type collection).
St. THomAs: Proctor 9684 (IJ).
From other relatively small-leaved and short-petioled species, Wallenta fawcetti may be recognized by its more attenuate leaves, which are gradually narrowed above the middle to a usually acute apex. It is known only from the crest of the Blue Mountains, at about 5,000—5,500 ft. (1,500-1,700 m.); the type was collected by Harris near Portland Gap but whether on the Portland or St. Thomas side is un- certain. The calyx of male flowers is about 1-5 mm. long, of female flowers about Imm. long; the corolla of male flowers is about 2-5 mm. long, of female flowers unknown.
5. Wallenia crassifolia Mez in Urban, Symb. Antill. 2 : 409 (1901); in Engler, Pflanzenr. IV. 236 : 244 (1902). (PI. roa.)
Wallenia venosa Griseb., Fl. Brit. W. Ind. Is. : 394 (1861) pro parte, excl. specim. Wilson.
St. ANDREW: Proctor 9890 (IJ; NY), Proctor 9524 (BM; IJ), Maxon & Killip 1026 (NY), Philipson 932 (BM), E. G. Britton 3866 (NY), Purdie (K, holotype).
St. THomas: Webster & Wilson 5445 (IJ), Proctor 9207 (IJ), Stearn ror (BM), Proctor 11458 (IJ).
Wallenia crassifolia inhabits mountain rain forest along the crest of the Blue Mountains between 4,750 and 6,500 ft. (1,400-2,000 m.). That the smallness of its elliptic or narrowly elliptic leaves is not simply a reduction occasioned by its high mountain environment is shown by the occurrence of the large-leaved and undoubtedly distinct W. calyptrata in the same habitat. The type-locality of W. crassifolia is Morces Gap, where it was first collected by William Purdie in 1843. The calyx of male flowers is about 2 mm. long, of female flowers about 1 mm. long, the corolla of male flowers about 4 mm. long, of female flowers unknown.
6. Wallenia corymbosa Urban, Symb. Antill. 5 : 457(1908). (Text-fig. 24A—J.)
TRELAWNY: Harris 8720 (BM; NY; UCWI; type-collection), Harris 9095 (NY; UCWI).
CLARENDON: Harris 10864 (NY), Harris 10996 (NY), Harris 12781 (BM; NY; UCWI), Harris 12803 (NY; UCW]).
St. ANN: Howard & Proctor 15037 (IJ), Howard & Proctor 15170 (BM; IJ), Proctor 26739 (BM).
Typical Wallenia corymbosa inhabits the central hills of Jamaica at about 2,000— 2,500 ft. (600-760 m.), probably extending well into the Cockpit Country, and is notable for its thick greyish or glaucescent narrowly obovate leaves.
The type-locality is on Crown Lands near Troy, where William Harris collected it at 2,500 ft. (760 m.) in 1904. Subsequent collections in the same area have provided an unusually adequate representation of it in male and female flower and in fruit. The flowers are described as “ pale yellow with red spots”’ or “‘ cream with red
170 A SYNOPSIS OF JAMAICAN MYRSINACEAE
stripes’’. The calyx of male flowers is 1-5-2 mm. long, of female flowers about I-4.mm. long, the corolla of male flowers 3-4 mm. long, of female flowers about 2°5mm. long. The fruits are about 3:5 mm. broad.
Fic. 24. Wallenia corymbosa Urban: A, female flowering specimen (Harris 12803), X 4; B, female flower (Harris 12803), xX 6; C, dissected calyx and gynoecium of female flower (Harris 12803), x 8; D, dissected corolla of female flower with aborted stamens (Harris 12803), X 8; E, placenta of female flower (Harris 12803), X 20; F, male flower (Harris 12819), X 8; G, dissected corolla of male flower with aborted gynoecium (Harris 12819), x 8; H, anther of male flower (Harris 12819), x 20; I, fruit (Harris 12781), X 5; J, leaf (Harris 12781), x 4. Wallenia subverticillata (Britton) Ekman: K, leaves (Harris & Britton 4049), x 4. Wallenia grisebachii Mez: L, leaf-base (Proctor 8247), X 3.
A SYNOPSIS OF JAMAICAN MYRSINACEAE LT
7. Wallenia xylosteoides (Griseb.) Mez in Urban, Symb. Antill. 2 : 409 (1901); in Engler, Pflanzenr. IV. 236 : 244 (1902). (Pl. rob.)
Ardisia xylosteoides Griseb., Fl. Br. W. Ind. Is. : 395 (1861) tantum quoad pl. Jamaic.
ST. ANN: Alexander 568 (K, lectotype).
Grisebach reported his Ardisia xylosteoides from “‘ Haiti! ; Mexico! ; New Granada! ”’ as well as Jamaica; these records cannot apply all to one species. Mez in Igor restricted the epithet xylosteoides to Jamaican material collected by Alexander on St. Ann’s Road, Moneague. The same species is represented by Jamaican Plants r118 (UCWI) and rraz (K) without locality. The calyx of female flowers is about i mm. long, the corolla scarcely longer. Male flowers and fruit are unknown. The epithet indicates a resemblance in leaf-form to Lonicera xylosteum L., the Xylosteum of pre-Linnaean authors.
8. Wallenia erythrocarpa Urban, Symb. Antill. 6 : 29 (1909). (Pl. rra.)
HANOVER: Harris 10343 (BM; NY; UCWI; type-collection).
WESTMORELAND: Harris 7080 (UCWI).
This species is still only known from material collected by Harris, the type from Fray Woods at 1,650-1,800 ft. (500-550 m.) in 1908; he gathered other material at Grandvale at 500 ft. (150 m.) in r898. It is notable for its large but relatively narrow long-petioled leaves with prominent reticulate veining, their narrowly elliptic-oblong shape and their size (11-21 cm. long, 4:5—7:5 cm. broad) distinguishing it from W. venosa. The calyx in fruit is about rmm. long. Male and female flowers are un- known.
9. Wallenia punctulata Urban, Symb. Antill. 7 : 322 (rgt2). (Pl. rrb.)
MANCHESTER: Britton 3281 (NY; type collection).
Wallenia punctulata is another little-known species, known only from the type- collection gathered by N. L. Britton in 1908 between Brown’s Town and Porus, but distinguished from other species with prominently reticulate-veined leaves by its _ more oblique lateral veins which pass outwards from the midrib at an angle of about 40°. The calyx in fruit is about 2 mm. long, the corolla of female flowers according to Urban about 1-7 mm. long.
10. Wallenia venosa Griseb., Fl. Brit. W. Ind. Is. : 394 (1861) quoad specim. Wilson.—Mez in Urban, Symb. Antill. 2 : 4r0 (Igor); in Engler, Pflanzenr. IV. 236 : 245 (1902).—Urban, Symb. Antill. 7 : 322 (1912). (Pl. 13a.)
HANOVER: Britton 2425 (NY), Britton 2292 (NY), Britton 2326 (NY). WESTMORELAND: Harris 10248 (BM; NY).
TRELAWNY: Harris 8768 (BM; NY; UCWI).
ST. ELIZABETH: Harris 9925 (BM; K; NY; UCWI).
MANCHESTER: Britton 3712 (NY).
CLARENDON: Proctor 18400 (BM; IJ), Proctor 26514 (BM; IJ), Proctor 27637 (BM;
TJ).
St. ANN: Proctor 26514 (BM).
172 A SYNOPSIS OF JAMAICAN MYRSINACEAE
ST. ANDREW: Harris 5633 (BM; NY; UCWI), Harris 6126 (BM), Harris 16032 (BM; NY; UCWI), Adams 11845 (UCWI).
St. THomas: Harris & Britton 10745 (NY), Britton 3963 (NY), Britton 3988 (NY), Britton 4145 (NY).
The lectotype of this widespread species is Wilson 388 (GOET, K), collected in 1858 without specified locality, which was designated by Mez in 1go1. Wallenia venosa has prominently reticulate-veined leaves with black dots beneath sometimes sO numerous as to give it a somewhat mottled appearance. The calyx of male flowers is 1-5-2 mm. long, of female flowers 0-8 mm. long, the corolla of male flowers about 3mm. long, of female flowers about Imm. long. The Alexander specimen (GOET) from the Blue Mountains cited by Grisebach under W. venosa belongs to W. crasstfolia.
11. Wallenia purdieana Mez in Urban, Symb. Antill. 2 : 408 (rg0r); in Engler, Phlanzenr. 1V. 236: 243-(r902)... (Pl. r2a:)
HANOVER: Harris 10305 (BM; UCWI), Proctor r13or (IJ).
ST. JAMES: Proctor 22992 (BM; IJ).
TRELAWNY: Harris 9096 (UCWI), Britton 510 (NY), Proctor 15663 (BM; IJ), Howard, Proctor & Stearn 14663 (BM; IJ).
MANCHESTER: Purdie (K, type), Proctor 16118 (BM; IJ), Howard, Proctor & Wagenknecht 20520 (NY), Harris & Britton 10601 (BM; NY; UCWI), Robertson 5416 (UCWI).
Wallenia purdieana is a species of western and central Jamaica between 1,000 ft. (300 m.) and 3,000 ft. (g00m.). The type was collected by William Purdie in Manchester, but without specified locality. The leaves on drying retain their green colour unlike those of other species which become brownish or grey. The calyx of male flowers is about 2 mm. long, of female flowers about 1-5 mm. long, the corolla of male flowers about 3 mm. long, of female flowers about 1-5-2 mm. long.
12. Wallenia grisebachii Mez in Urban, Symb. Antill. 2 : 411 (1901) quoad specim. Wullschlégel 1338; in Engler, Pflanzenr. IV. 236 : 245 (1902).—Urban, Symb. Antil. 6°31 (1900); (Text-ig. 24 Pl 225.)
Wallenia laurifolia sensu Griseb., Fl. Brit. W. Ind. Is. : 394 (1861) pro parte; non Swartz.
ST. JAMES: Proctor 22154 (BM; IJ).
St. ELIZABETH: Howard & Proctor 13745 (IJ; NY), Howard & Proctor 13755 (IJ).
TRELAWNY: Perkins 1385 (K).
MANCHESTER: Wullschlagel 1338 (Munich, lectotype), S. Brown 300 (NY), Robertson 5470 (UCWI), Adams ror16 (UCWI1), Adams 11766 (UCW]).
CLARENDON: Proctor 8247 (IJ).
St. ANN: Britton & Hollick 2776 (NY; UCWI).
The species W. grisebachit as accepted here consists of plants with fairly large coriaceous leaves having the rounded base contracted abruptly into the petiole and becoming brownish on drying. Mez separated W. grisebachit from among material included by Grisebach in W. laurifolia. Urban further divided this, restricting the name W., grisebachw to the taxon represented by Wullschldgel 1338 collected at
A SYNOPSIS OF JAMAICAN MYRSINACEAE 173
Fairfield, Springfield, in Manchester parish, between 1847 and 1849. The inflores- cence is compact; the flowers are rusty brown. The calyx of male flowers is 2 mm. long, of female flowers 1-5-2 mm. long, the corolla of male flowers about 4 mm. long, of female flowers about 1-5 mm. long.
13. Wallenia calyptrata Urban, Symb. Antill. 5 : 458 (1908); in Fedde, Repert. Sp. Nov. 13 : 469 (1915).
St. ANDREW: Shreve s.n. (NY), J. A. Harris & Lawrence 15490 (NY), Adams 7440 (UCW]).
PORTLAND: Proctor 26717 (BM; IJ).
St. Tomas: Proctor 9638 (IJ), Proctor 9683 (IJ; NY), Stearn 1ro4 (BM), Proctor 11457 (IJ).
Although the type-gathering of W. calyptrata made by Rehder in 1903 at Morces Gap has not been seen, the agreement of Urban’s description with the other material collected along the crest of the Blue Mountains leaves no doubt as to its identity. Its range here seems to be between 5,000 ft. (1,500 m.) and 7,000 ft. (2,100 m.) in mountain mist-forest. It has prominently reticulate-veined leaves 8-14 cm. long, 3°5-6 cm. broad, and larger male flowers than other species. The calyx of male flowers is 3-3°5 mm. long, of female flowers 2-5 mm. long, the corolla of male flowers 5-6 mm. long, of female flowers 3 mm. long.
14. Wallenia laurifolia Swartz, Nov. Gen. & Sp. Pl. : 31 (1788); Fl. Ind. Occ. 1: 248 (1797).—Mez in Urban, Symb. Antill.2:407 (1901); in Engler, Pflanzenr. IV. 236 : 243 (1902).—Urban, Symb. Antill. 8 : 521 (1921r).—Alain in Leon & Alain, Fl. Cuba 4 : rro, fig. 42 (1957).
Petesioides laurifolium Jacq., Select. Stirp. Amer. Hist. : 17 (1763), nom. invalid. Wallenia clusiifolia Griseb., Fl. Brit. W. Ind. Is. : 394 (1861).
“ Bryonia nigra fruticosa, foliis laurinis, floribus racemosis speciosis ’’ Sloane, Cat. Pl. Jam. : 106 (1696); Voy. Jam. Nat. Hist. 1 : 234, t. 145 f. 2 (1707).
HANOVER: Proctor 10035 (BM; IJ).
MANCHESTER: S. Brown 138 (NY), Proctor 24175 (BM; IJ), Crawford 775 (NY), Farr & Sanderson 22334 (IJ), Harris & Britton 10613 (NY; UCWI), Howard & Proctor 14964 (IJ), Howard, Proctor & Wagenknecht 20520 (BM; IJ), Harris 12862 (BM; NY), Howard, Proctor & Stearn 14708 (BM; IJ).
CLARENDON: Harris 11188 (NY; UCWI).
ST. ANN: Stearn 603 (BM), Proctor 11907 (IJ), Proctor 8632 (IJ), Proctor 16142 (IJ).
ST. CATHERINE: Britton 2616 (NY).
St. Mary: Sloane, Herb. IV. 494 (BM), Proctor 7554 (NY; IJ).
ST. ANDREW: Harris 6880 (BM; NY; UCWI). — :
PORTLAND: Stearn 538 (BM), Proctor rr84r (IJ), Wight 195 (NY), Maxon & Killip 830 (NY), Harris & Britton 10767 (K; NY), Proctor 11347 (BM; IJ), Proctor 11352 (IJ), Proctor 16259 (IJ), Howard, Proctor & Stearn 14758 (BM; IJ), Stearn 226 (BM; UCWI), Britton 4r2r (NY).
ST. THomas: Britton 4109 (NY), Powell 1647 (BM; IJ).
174 A SYNOPSIS OF JAMAICAN MYRSINACEAE
This widespread species, first described from Hispaniola and also known from Cuba, is mainly distinguished by its relatively inconspicuous veining with the areas between the main lateral veins flat, not reticulate with prominent veinlets. The material from the upper part of the John Crow Mountains exemplified by Proctor 9822 gathered at 1,500—2,500 ft. (450-750 m.), with very small leaves (to 6 cm. long) and reduced inflorescence, differs much in appearance from that of the coast, exemplified by Stearn 226, but these differences are assumed to result from the diversity of habitat, as small leaves approaching these are to be found on some specimens from elsewhere referred to this species, e.g. Howard, Proctor & Stearn 14708 from Manchester, Proctor 8632 from St. Ann, and Proctor 10035 from Hanover, and inter- mediates occur moreover on the John Crow Mountains. The calyx of male flowers is I°5-2°5 mm. long, of female flowers about 1-5 mm. long, the corolla of male flowers 2°5-3 mm. long, of female flowers about 1-5 mm. long.
15. Wallenia sylvestris Urban in Fedde, Repert. Sp. Nov. 13 : 468 (1915). (Pl-13b:)
PORTLAND: Proctor 10477 (IJ; NY), Proctor 16258 (IJ), Proctor 5266 (BM; IJ), Proctor 5244 (IJ).
PORTLAND or St. THomas: Harris & Britton 10720 (NY, type-collection), Harris & Britton ro690 (K; NY; UCWI), Harris & Britton 10776 (NY; UCWI).
This species is apparently confined to mountain woods on the John Crow Mountains at 1,500-2,500 ft. (450-750 m.). The type-locality is the southern end of the John Crow Mountains near the boundary of the parishes of Portland and St. Thomas. It is notable for its large long-petioled leaves with elliptic or narrowly elliptic (rarely obovate or narrowly obovate) blades up to 22 cm. long, 9 cm. broad. The calyx and corolla of female flowers are about 2mm. long. The male flowers are unknown. The subglobose fruits are slightly punctate and vertically ribbed.
16. Wallenia discolor Urban, Symb. Antill. 6 : 29 (1909).
St. ANN: Alexander (fide Urban).
Described by Urban from fruiting material collected by Alexander, probably in St. Ann but without recorded locality, this appears to be distinguished from W. laurifolia by its larger leaves (18-25 cm. long), shorter pedicels (0-5-3 mm. long) and coarsely punctate but not ribbed fruits. Male and female flowers are unknown.
3. MYRSINE L.
MyrsINE L., Sp. Pl. 1: 196(1753), Gen. Pl. ed. 5: 90(1754). Type (by monotypy): M. africana L.
The genus Myrsine as accepted here includes the groups kept apart by Mez as Myrsine, Rapanea and Suttonza.
KEY TO JAMAICAN SPECIES
Young shoots and petioles minutely pubescent; leaf-blades mostly narrowly elliptic and mostly less than 2:5 cm. broad; corolla lobes joined for the lower 0-5 mm. ormore . : : , 5 ; ‘ , ; . 1. M. coriacea
A SYNOPSIS OF JAMAICAN MYRSINACEAE 175
Young shoots and petioles glabrous; leaf-blades mostly obovate or narrowly obovate and mostly more than 2:5 cm. broad; corolla lobes free to the base 2. M. acrantha
1. Myrsine coriacea (Swartz) R. Br. ex Roem. & Schult. in L., Syst. Veg., ed. nov. 4 : 511 (1819).—A. DC. in DC., Prodr. 8 : 99 (1844). (Text-fig. 25E—H.)
‘Samara coriacea Swartz, Nov. Gen. & Sp. Pl. : 32 (1788); Fl. Ind. Occ. 1 : 261 (1797).
Caballeria ferruginea Ruiz & Pavon, Syst. Veg. Fl. Peruv. & Chil. 1 : 280 (1798).
Myrsine ferruginea (Ruiz & Pavon) Spreng. in L., Syst. Veg., ed. 16, 1 : 664 (1825).
Myrsine laeta sensu Griseb., Fl. Brit. W. Ind. Is. : 392 (1861); non A.DC.
Rapanea coriacea (Swartz) Mez in Urban, Symb. Antill. 2 : 428 (1901); in Engler, Pflanzenr. LV..236. + 380° (1902);
Rapanea ferruginea (Ruiz & Pavon) Mez in Urban, Symb. Antill. 2 : 429 (1901); in Engler, Pflanzenr. IV. 236 : 381 (1902).—Urban, Symb. Antill. 8 : 522 (1921).—Britton & Wilson in Sci. Surv. Puerto Rico 6 : 61 (1925).—Alain in Leén & Alain, Fl. Cuba 4 : 112, fig. 43 (1957).—J. F. Macbr. in Publ. Field Mus. Nat. Hist., Bot. 13 (5) : 168 (1959).—Little & Wadsworth, Common Trees of Puerto Rico : 432, fig. 203 (1964).
ST. JAMES: Norman 50 (BM).
TRELAWNY: Webster, Proctor & Powell 5355 (BM; IJ), Britton 486 (NY), Adams 12413 (UCWI).
MANCHESTER: Proctor 21920 (BM; IJ), Britton 3216 (NY), Adams 8466 (UCWI), Proctor 23118 (BM; IJ).
CLARENDON: Proctor 9723 (IJ), Proctor 8427 (IJ), Proctor 15905 (BM; IJ; UCWI).
St. ANN: Proctor & Howard 14992 (IJ), Howard, Proctor & Stearn 14607 (BM; IJ), Proctor 7471 (BM; IJ), Skelding 3016 (UCWI), Adams 12698 (UCWI).
ST. CATHERINE: Howard & Proctor 13599 (IJ).
ST. ANDREW: Adams 12675 (UCWI), Britton 147 (NY).
PoRTLAND: Philipson 933 (BM), Proctor rorro (IJ; NY), Howard, Proctor & Stearn 14767 (BM; IJ).
St. THomas: Webster & Wilson 5454 (IJ), Stearn 111 (BM), Proctor 11463 (IJ), Maxon 9442 (US), Proctor 1108 (IJ), Stearn 505 (BM).
A widely ranging species, recorded in the West Indies from Cuba, Hispaniola, Jamaica, Puerto Rico and the Lesser Antilles and on the American mainland from Mexico to Peru, Chile, Bolivia and Argentina, this is commonly known as Rapanea ferruginea, a name based on Peruvian material; Swartz’s Samara coriacea, based on Jamaican material being, in my opinion, conspecific, his epithet is adopted here. It is sometimes called “‘ colic wood’ in Jamaica, according to Proctor.
2. Myrsine acrantha Krug & Urban in Notizbl. K. Bot. Gart. Mus. Berlin 1 : 79 (1895); in Urban, Symb. Antill. 1 : 380 (1899). (Text-fig. 25A—D.)
Myrsine coriacea sensu Griseb., Fl. Brit. W. Ind. Is. : 392 (1861); non (Swartz) R. Br.
Rapanea guianensis Mez in Urban, Symb. Antill. 2 : 431 (1901); in Engler, Pflanzenr. IV. 236 : 392 (1902) pro parte, quoad pl. Jamaic.; non Aubl.
Rapanea acrantha (Krug & Urban) Mez in Urban, Symb. Antill. 2 : 433 (1901); in Engler, Pflanzenr. IV. 236 : 376 (1902).
ICH
176 A SYNOPSIS OF JAMAICAN MYRSINACEAE | /
Fic. 25. Myrsine acrvantha Krug & Urban: A, fruiting specimen (Robertson 5372), x 4; B, flower (Harris 8295), X 10; C, dissected flower with aborted gynoecium (Harris 8295), x 10; D, fruit (Harris 7061), x 5. Myrsine coriacea Swartz: E, leaf (Stearn 111), X $; F, leaf (Stearn 505), x 4; G, dissected corolla with stamens (Stearn 111), X 10; H, dissected calyx with gynoecium (Stearn 111), X Io. 5
WESTMORELAND: Proctor & Mullings 22061 (BM; IJ). St. ELizABETH: Britton 1320 (NY), Proctor 7754 (BM; IJ), Howard & Proctor 13682 (IJ). ; MANCHESTER: Howard, Proctor & Wagenknecht 20521 (BM; NY; UCW]I), Robertson 5372 (BM; K; UCWI), Harris 8295 (BM; NY; UCWI), Proctor 21908 (BM; IJ),” Proctor 11619 (BM; IJ), S. Brown 159 (NY), Adams rorr4 (UCWI), Proctor 23130
(IJ).
ahd i eke ea
A SYNOPSIS OF JAMAICAN MYRSINACEAE 177
ST. ANDREW: Harris 6072 (NY), Harris 5398 (K; NY; UCWI; type-collection of Myrsine acrantha), Harris 5398 bis (BM; UCWI), A. von der Porten 2035 (IJ), Harris .5528 (NY; UCWI), Loveless 1502 (IJ; UCWI).
PORTLAND: Proctor 19737 (BM; NY; UCWI).
St. THomas: Proctor 9580 (BM; IJ).
Myrsine acrantha has hitherto been regarded as a rare and obscure species, known only from the type and topotype collections gathered by Harris in 1894 and 1895 at Strawberry Hill Cottage high in the Blue Mountains of St. Andrew. Mez distin- guished it by its free petals from the two other Jamaican species accepted by him under the names Rapanea ferruginea and R. guianensis. Thus according to his own generic key in the Pflanzenreich IV. 236: 15 (1902) it should be referred to the otherwise Pacific Ocean group Suttonia! In fact all the Jamaican specimens hitherto referred to Myrsine (or Rapanea) guianensis agree with M. acrantha in having obovate or narrowly obovate leaves, flowers borne on very short bract-covered lateral shoots, and free petals. Myrsine guianensis (Aubl.) Kuntze does not occur in Jamaica; it was described by Aublet in 1775 (Hist. Pl. Guiane Frang. 1 : 121; 3 : t. 46) as Rapanea guianensis from French Guiana and is accepted by Mez as ranging from Florida over the West Indies to the Guianas and over Mexico and Central America to Bolivia. Typical M. guianensis as represented by specimens from French Guiana (Aublet! Sagot 913! Martin!) is notable for its large obovate leaves up to 10 cm. long, 7 cm. broad, with a glossy almost varnished upper surface and distinctly punctate lower surface; plants of similar character occur in Bolivia, Colombia, British Guiana, and
| Surinam. Very similar to this but distinguishable by the leaves being marked with short lines beneath is M. trinitatis A.DC. occurring in Trinidad, Tobago, Grenada, Martinique and Puerto Rico. Plants from Hispaniola, Cuba, Jamaica, the Bahamas and Florida stand apart from these in their smaller leaves, rarely more than 8 cm. long, 2-5 cm. broad, less glossy above, punctate but not lined beneath. Whether more than one taxon is represented in this mostly sterile material is uncertain. Some specimens from Cuba, e.g. Ekman 15603 (Oriente province), and from Haiti, e.g. E. C. Leonard 8456 (Departement du Nord) in their leaves and short spurs unfortunately lacking flowers, closely resemble the Jamaican M. acrantha. In the Bahamas and Florida the plants of this alliance as represented by A. Curtiss 44 from New Providence, Bahamas, and R. A. Howard 8047 and Tracy 7454 from Florida have their five corolla-lobes fused at the base for about 0-4-0-5 mm., as figured by
| Small (Man. Southeast Fl. : 1029 (1933)). They represent a species distinct from
| M. acrantha for which the correct name appears to be Myrsine punctata.*
* Myrsine punctata (Lam.) Stearn, comb. nov. Sideroxylum punctatum Lam., Tabl. Encyc. & Méth., Bot. 2: 42 (1794) [from Carolina, Michaux]. Bumelia punctata (Lam.) Roem. & Schult. in L., Syst. Veg., ed. nov. 4 : 498 (1819). Myrsine floridana A.DC. in Trans. Linn. Soc. London 17 : 107 (1834); in DC., Prodr. 8 : 98 (1844) {from Florida]. , Rapanea guyanensis sensu Small, Man. Southeast FI. : 1029, fig. (1933); non Aublet.
SPECIAL LITERATURE
GrisepacH, A. H.R. 1859-64. Flora of the British West Indian Islands. London. Hosaka, E. Y. 1940. A revision of the Hawaiian Species of Myrsine (Suttonia, Rapanea), Myrsinaceae. Occ. Pap. Bernice P. Bishop Mus., Honolulu. 16 : 25-79.
178 A SYNOPSIS OF JAMAICAN MYRSINACEAE
LUNDELL, C. L. 1966a. Flora of Guatemala: Myrsinaceae. Fieldiana, Bot. 24 (8) : 135-200. |
1966b. The genus Parathesis of the Myrsinaceae. Contrib. Texas Res. Found. Bot. Studies, —
5. Renner, Texas. |
Mez,C. 1901. Myrsinaceae. Urban, Symb. Antill. 2 : 389-433.
1902. Myrsinaceae. Engler, Pflanzenreich IV. 236 (Myrsinaceae).
STEARN, W. T. 1965. Grisebach’s Flora of the British West Indian Islands: a Soar cioa and bibliographical introduction. Journ. Arnold Arb. 46 : 243-285.
Swartz, O. 1797. Flora Indiae occidentalis 1 : 247-249, 468-469.
Urpan, I. 1898. Bibliographia Indiae occidentalis botanica. Urban, Symb. Antill. I : 3-195.
1909. Incrementa siphonogamarum florae jamaicensis. Urban, Symb. Antill. 6 : 70-131,
Wa ker, E. H. 1940. A revision of Eastern Asiatic Myrsinaceae. Philipp. Journ. Sct.
73 : 1-258 1954. Concerning the Myrsinaceae (‘‘ Ardisiaceae’’) of Japan. Bot. Mag. Tokyo 67: IO5-III, 155-162, 203-213, 247-255.
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4
‘THE JAMAICAN SPECIES OF COLUMNEA AND
ALLOPLECTUS (GESNERIACEAE)
BY
WILLIAM THOMAS STEARN
British Museum (Natural History) Ai |
Pp. 179-236; 29 Text-figures, Plates 14-21
BULLETIN: OF
THE BRITISH MUSEUM (NATURAL HISTORY) BOTANY. Vol. 4 No. 5
LONDON : 1969
THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), imstituted in 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.
In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department.
This paper is Vol. 4, No. 5 of the Botany series.
World List abbreviation Bull. Br. Mus. nat. Hist. (Bot.).
© Trustees of the British Museum (Natural History) 1969
TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 10 June, 1969 Price {1 8s.
THE JAMAICAN SPECIES OF COLUMNEA AND ALLOPLECTUS (GESNERIACEAE)
By WILLIAM T. STEARN With Notes on CYTOLOGY and CHEMOTAXONOMY By BRIAN D. MORLEY
SUMMARY
Eleven species of Columnea L., together with hybrids between two species (C. rutilans and C. urbanii), and two species of Alloplectus Mart. occur in Jamaica. All are endemic and have distinctive ranges within the island. Coluwmnea proctorii and C. urbanii, here described as new species, have previously been misidentified.
The present paper deals with their history and morphological characteristics, the typification of the names Columnea, Achimenes and Alloplectus, the definition of the genera Columnea and Alloplectus, and the effect of pollination by humming-birds on their evolution; it also provides notes on cytology and chemotaxonomy as well as a key, descriptions and enumerations of speci- mens. Diagnostic features of pubescence and calyx are illustrated. A survey by methods of numerical taxonomy (taxonometrics) using both a wide range of characters and a reduced range gave a classification of the species in general agreement with one reached by traditional methods.
INTRODUCTION
THE following paper on the Jamaican species of Columnea and Alloplectus is the out- come of the preparation of the text on the Gesneriaceae for Fawcett and Rendle’s Flora of Jamaica, volume 6. A study of all the available herbarium material before my visit to Jamaica in 1955-56 (cf. Stearn, 1959) indicated the need for further collecting and study of the species, despite the excellence of Morton’s careful and critical revisions of 1944, and I gathered specimens of them whenever I came across them. Thanks to the co-operation of Mr. George R. Proctor of the Institute of Jamaica I was able to see most of them in their type localities or nearby. Subsequent collecting by Proctor has further increased our knowledge of their distribution. In 1965 Dr. Brian Morley visited Jamaica to study the cytology of the genus Columnea, using wild material. His field work revealed the existence of hybridization between C. rutilans and C. urbanii and he was able to introduce most of the species into cultivation at the Royal Botanic Gardens, Kew. Study of the abundant material now available made evident the need also to consider the definition of the genera Columnea and Alloplectus on a wider basis. It provided data which appeared suitable for testing taxonometric analysis as a means of classifying the species. These matters are dealt with below.
THE TYPE OF COLUMNEA
The genus Colwmnea L. (sensu stricto) comprises more than 100 species, all native to tropical America. It attains its greatest morphological diversity and breaks up BOT. 4, 5. II
182 THE JAMAICANSPECIESOFCOLUMNEA ANDALLOPLECTUS
into its greatest number of species in northern South America and Central America: 17 species, for example, are recorded from Costa Rica alone. In the West Indies as a whole it is represented by some 16 species, one of which, C. scandens L., is the type-species by monotypy. Charles Plumier collected this between 1689 and 1697 “in locis insulae martinicanae ’’, i.e. in Martinique, Lesser Antilles, and founded a new genus upon it dedicated to the Italian botanist Fabius Columna (i.e. Fabio Colonna, 1567-1050). He published a brief description, together with an engraving showing floral details, in his Nova Plantarum Americanarum Genera : 28, t. 33 (1703) as follows:
“ Columnea est plantae genus flore monopetalo, personato, cujus labium superius non nihil fornicatum & excavatum, inferius vero tripartitum. Ex calyce autem surgit pistillum, posticae floris parti, ad instar clavi infixum, quod deinde abit in fructum globosum, mollem, seminibusque plenum exiguis, & oblongis.
Columneae species sunt.
Columnea scandens, phaeniceo flore, fructu albo, Columnea scandens, flore lutescente, fructu albo.”’
Plumier had, however, drawn up a much more detailed description and had made a coloured drawing depicting the flowers as red, the original of which is at the Muséum National d’Histoire Naturelle, Paris, but of which there are copies in the University Library, Groningen, the Herbarium Library of the Royal Botanic Gardens, Kew (Plum. MSS. 2 : 117), and the Department of Botany, British Museum (Natural History), London (Plum. MSS. 2 : 104). That now at Groningen forms part of the set of copies of Plumier’s drawings originally made for Boerhaave, which passed after his death to Johannes Burman, Linnaeus’s friend, at Amsterdam. In 1756 Burman published an engraving made from it, excellently portraying the habit of the whole plant, in Plantarum Americanarum Fasciculus quartus: t. 89, Lamarck’s account in the Encyclopédie méthodique, Botanique 2 : 66 (1786), which is fuller than any previously published, was based on Plumier’s manuscript material in Paris. Plumier’s plant with yellowish flowers is assumed to be a rare colour form.
Linnaeus himself possessed no specimens of Colwmnea and he based his description of the genus in his Genera Plantarum: 373 no. 921 (1737), 5th ed.: 286 no. 710 (1754) and likewise his protologue of C. scandens in the Species Plantarum 2 : 638 (1753) primarily on Plumier’s published text and figures, although it is evident that between the preparation of the first edition (1737) and the second edition (1742) of the Genera Plantarum he saw herbarium material of a Colwmnea which enabled him to amend — his 1737 statement “‘ STAM: Filamenta duo (pingit, an quatuor) ’’, based on Plumier’s © figure which portrays only two stamens, to ““ STAM. Filamenta quatuor, quorum duo— longiora, sub labio superiore recondita. Amntherae simplices”’ in 1742. He may well have seen material during his visit to Parisin 1738. In any event he stated “ Habitat in Gallia aequinoctiali”’, meaning the French possessions in tropical America. The traditional use of the name Columnea scandens L. for the red-flowered species occurring in Martinique is thus well-founded. This ranges in the Lesser Antilles from Montserrat, by way of Martinique, St. Vincent and Grenada, to Trinidad and |
Tobago, but does not occur in the Greater Antilles. :
:
; _ :
4
THE JAMAICAN SPECIES OFCOLUMNEAANDALLOPLECTUS 183
SLOANE’S JAMAICAN MATERIAL
The first record of any species now included in the genus Columnea was made not
_ by Plumier but by Hans Sloane and resulted from his stay in Jamaica from December
,
1687 to March 1689. He found “in montosis Insulae Jamaicae sylvis’’ a species
_ which he listed as ‘‘ Rapunculus fruticosus foliis oblongis integris villosis ex adverso
sitis flore purpureo villoso”’ in his Catalogus Plantarum quae in Insula Jamaica
_ sponte proveniunt : 58 (1696). This entry would be completely cryptic but for the
description and the rather crude engraving in his later Voyage to the Islands Madera...
_ and Jamaica i : 157, t. 100 f. r (1707) and the specimen in Herb. Sloane 3 : fol. 21
at the British Museum (Natural History) on which these were based. Sloane’s artist Everard Kickius unfortunately failed to distinguish the calyx from the corolla,
_ both being very hairy, but examination of Sloane’s specimen shows his plant to be
Columnea fawcettit (Urban) Morton, which was not distinguished as a species until 1944. Sloane probably collected it on Mount Diablo.
THE TYPE OF ACHIMENES
Patrick Browne, failing to connect Linnaeus’s genus Colummnea with any Jamaican plant, established a new genus Achimenes in his Civil and Natural History of Jamaica : 270, t. 30 (1756). He included two species. The first of these, his ‘“‘ Achimenes major, herbacea, subhirsuta, oblique assurgens’”’ to which he referred Sloane’s “ Rapunculus fruticosus . . .” is undoubtedly a Columnea, probably C. fawcettit, since he mentions it as “‘ most commonly met with in the woods of New Liguanea and St. Ann’s”’ and describes the “‘ divisions of the cup’, i.e. calyx-segments, as “ pinnated at the sides, somewhat like those of the garden rose’’. Browne’s generic description was based on this species, as is evident, for example, from his account of
_ the irregular corolla: “limbus erectus, in quatuor lacinias inaequales, inaequaliter
sectus; Jacinta superior recta, latiuscula, profunde crenata, sive bifida; laterales
oblongae & a superiori oblique decendentes; infima anguste patula, ad medietatem - floris incisa, longissima.”’ Hence Achimenes P. Browne (1756) as to type is congeneric
s i)
x
”?
with Columnea L. Browne’s second species ‘‘ Achimenes minor, erecta, simplex .. . is the plant commonly known as Achimenes coccinea (Scop.) Pers. or A. pudchella (L’Hérit.) Hitchc., but whose correct name is A. erecta (Lam.) H. P. Fuchs. Browne regarded it as agreeing ‘“‘in the most essential parts fof the flower] with the fore-
_ going ’’, although it has an almost regular corolla. When Persoon dealt with these _ two Brownean species in his Synopsis Plantarum 2 : 164 (1806) he referred Browne’s first species to Columnea, as Burman had already done in 1756 and Swartz in 1788, _ but he retained the name Achimenes for the second species, at the same time providing _ a new generic description referring to this alone: “ Cal. superus 5-phyllus. Cor. infundibuliformis; limbus planus, 5-lobus, subaequalis. Rudimentum filamenti 5-ti. _Antherae connexae. Caps. bilocularis.’”’ Under the name Achimenes Persoon in fact established a new genus distinct from Achimenes P. Browne (sensu stricto) and the name Achimenes has been consistently used for it. This now includes about 25
species, of which 13 are in cultivation, together with many cultivars, some of hybrid
184 THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS
origin. Being a later homonym of Achimenes P. Browne, it needs conservation; the type-species of Achimenes Pers. is A. coccinea (Scop.) Pers., i.e. A. erecta (Lam.) HPs Buchs:
THE TYPE OF ALLOPLECTUS
The genus Alloplectus was founded by Martius, Nov. Gen. Sp. Pl. Brasil. 3 : 53 (1829), with the following differential character: “Calyx liber, coloratus, penta- phyllus, foliolis imbricatis, plus minus connatis, duobus interioribus. Corolla infera, tubulosa vel claviformis, rectiuscula, limbo brevi quinquelobo aut quinquedentato. Stamina quatuor didynama, cum quinti postici rudimento minimo, e basi tubi. Annulus hypogynus in glandulam posticam tumens. Capsula baccans, ovata, coriacea, unilocularis, bivalvis, seminibus numerosis oblongis.”” He included two Brazilian species, both described and illustrated in detail, A. sparsiflorus Mart. and A. circinatus Mart.
In the list of conserved generic names in the International Code of Botanical Nomenclature the name Alloplectus Mart. (1829) is conserved under No. 7860 against Crantzia Scop. (1777) and Vireya Raf. (1814) and its lectotype given as A. sparsiflorus Mart. (= A. Murtellus (Schott) Preston), although A. circinatus would have been a better choice since Martius saw no mature fruits of A. sparsiflorus but had both flowering and fruiting material of A. civcinatus and described the fruits and seeds of Alloplectus from this species. His two Brazilian species are, however, closely allied and differ in several characters from many species now referred to the genus Allop- lectus. If, however, a broad concept of the limits of Alloplectus is accepted, as in the present paper, Dalbergaria Tussac (Fl. Antilles 1 : 141 (1811-13) must be added to its synonomy as a nomen rejiciendum, since its type, D. phaenicea Tussac, loc. cit. (1811-13) is the same Hispaniola species (Alloplectus sanguineus (Pers.) DC., Prodr. 7 : 546 (1839))* as Rafinesque’s Vireya sangutinolenta, i.e. the type of Vireya Raf. already listed as a nomen rejiciendum. The short-lived Palermo Sfecchio delle Scienze o Giornale enciclopedico di Sicilia,t wherein (I : 194 (June, 1814)) Rafinesque published Viveya, is an extremely scarce periodical and his account is accordingly reprinted here:
“VIREYA. Cal. 5partitus, lac. equal. laciniatis, Corolla peripetala tubulosa 5dentata, basi gibbosa; Stam.4. didynamica inclus. Ov. liberum oblong. basi monadeno, stigma 2lamellat. Capsula 1.locul. 4valve polysperma, semina recept. — 4. longitudinal. affixa. Planta, fol. oppos. flor. axillaris. — Oss. Mi. fu comunicato — questo nuovo genere del Sign. Turpin col. barbaro nome di Caonabo, al quale ho sostituito il nome dell’ egregrio moderno naturalista e filosofo Virey. Si appartiene — alla mia seconda Classe Mesogynia, 5. Ordine Epidia, Famiglia Didynamia: ha delle affinita colli generi Browallia, e Lindernia.
Vireya sanguinolenta. Caule erecto simplex, foliis oppositis, alternis multo major, oblongis integris acutis sub-villosis subtus sanguinolentis, florib. axillarib. solitaris sessilis. — Oss. Nasce nell’Isola di S. Domingo offerisce le singolari particolarita di
* Martius, Nov. Gen. & Sp. 3 : 57 (1829) suggested that Besleria sanguinea Pers. belonged to Alloplectus but he did not make the combination A. sanguineus attributed to him by Jackson (Index Kew. 1 : 84
(1893)), Urban (Symb. Antill. 2 : 358 (1901)), and Morton (Contr. U.S. Nat. Herb. 29 : 4 (1944)). + The British Museum (Natural History) now possesses a photostat copy.
THE JAMAICAN SPECIESOFCOLUMNEAANDALLOPLECTUS 185
avere alternativamente le foglie dieci volte pit: grande delle opposte, e coperte al di sotto di una larga macchia sanguinosa: ha le corolle d’un giallo livid6.” Vireya Raf. is thus a synonym of Dalbergaria.
RECORDS OF COLUMNEA AND ALLOPLECTUS IN JAMAICA
The first post-Linnaean author to deal with the Jamaican species of Columnea was Olof Swartz, who travelled extensively in Jamaica between 1784 and 1786 and succeeded in finding an extraordinary number of rare plants. His journeys took him into the parishes of Trelawny, Hanover, St. Elizabeth, St. Andrew, St. Catherine and St. Thomas. His collections included three new species of Columnea, i.e. C. hirsuta, C. hispida and C. rutilans, of which he published diagnoses in his Nova Genera et Species Plantarum seu Prodromus : 94 (1788) and detailed descriptions in his Flora Indiae Occidentalis 2 : 1080-1086 (1800). Under his C. hirsuta he cited as synonyms Sloane’s “‘ Rapunculus fruticosus . . .”” and Browne’s ‘‘ Achimenes major herbacea .. .”, but from the significant note in his description, “‘ Hirsuties pulchre articulata,” and his specimen in the British Museum (Natural History) it is clear that he based his C. hirsuta on material of the species growing high in the Blue Mountains which has conspicuous multicellular hairs on the leaves. To this species the name C. hirsuta has always been attached. Swartz himself ascended Blue Mountain Peak in 1785 and, while a guest of Matthew Wallen at Coldspring, St. Andrew, in the Blue Mountains, he had other opportunities of collecting the species. In the west of Jamaica he collected C. rutilans, conspicuous for its leaves red beneath. Swartz’s third species, C. hispida, was long enigmatic. He himself collected it only in fruit “in montibus altis Jamaicae occidentalis, ad rupes umbrosas’’. Its flowers in fact remained unknown until February 1957; the flowering material collected by Wullschlagel in Manchester which Grisebach in 1862 (Fl. Brit. W. Ind. Is. : 465) referred to C. hispida belongs to another species, C. urbanit. Swartz’s own descrip- tion and his specimens at the British Museum (Natural History) and the Rijksmuseet, Stockholm, show C. hispida to be a very distinct species and justify Morton’s com- ment in 1944: “‘ From description I am not able to identify this with any of the species known from Jamaica. It is perhaps a valid species, not collected again.”’ Swartz, as was unfortunately his custom, gave no precise locality. In 1784 he travelled across western Jamaica from Montego Bay on the north coast to Savanna- la-Mar on the south coast and it would seem probable that he then came across this very rare and local species, for White Rock Hill, St. James, where Proctor and I had the good fortune to rediscover it in March 1956, lies near a route between these towns.
Grisebach’s account of Coluwmnea in his Flora of the British West Indian Islands : 464-465 (1862) added a further species, C. argentea Griseb., notable for its sericeous leaves and other parts, which the missionary Heinrich Rudolf Wullschlagel (1805-64) of the Moravian Brothers had collected in 1849 at Nazareth, Manchester. Evidently unaware of Grisebach’s publication, Hanstein described it anew in 1865 as C. wullschlaegeliana. This again is a very local species. In 1956 I was overjoyed to find it growing among rocks near the Moravian Chapel at Nazareth, which must have been the very place where Wullschlagel gathered it. As noted above, Grisebach included under C. hispida Swartz a species here named C, urbanit.
186 THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS
Grisebach placed three Jamaican species in the genus Pterygoloma, which was founded by Hanstein in 1854 (Linnaea 26 : 211) on a species described and illustrated in Bot. Mag. 72 : t. 4250 (1846) as Alloplectus repens Hook. and there stated to have been collected by William Purdie “‘ in the ascent of the Sierra Nivada, Santa Martha ”’, Colombia. The first of these, which Grisebach erroneously identified with Ptery- goloma repens (Hook.) Hanstein and enumerated as such, is a dwarf creeping species, with flowers not so markedly zygomorphic as in typical species of Columnea; it had been collected in Jamaica both by Wullschlagel and Purdie. In 1gor Urban distin- guished it as Columnea jamaicensis. Grisebach’s two other species, P. pubescens from western Jamaica and P. cristatum from central Jamaica, based on collections by Wilson, Purdie, Alexander and Wullschliagel, agreed with the first in their low creeping habit but differed in having almost cylindrical or clavate corollas with very short almost equal rounded lobes. They are easily distinguished from one another by the calyx-segments, entire in P. pubescens, pinnatifid with spreading lobes in P. cristatum. Fawcett listed all three under Alloplectus (see p. 234) in 1893. The association of the three accords with the results of a taxonometric analysis of the group (see p. 213).
Nothing more was published on the Jamaican species of Columnea until 1901, when Urban included them in his ‘‘ Enumeratio Gesneriacearum ” (Symb. Antill. 2 : 344-388 (1901)). He revised nomenclature, listed synonyms and specimens, and amended previous descriptions in the careful and scholarly manner which makes his contributions to West Indian botany so valuable as quarries of information, but as usual he failed to provide a key to the plants concerned, an omission which detracts so much from the practical utility of his work. He recorded five species as Jamaican, expressing his opinion about their distinctness by the comment ‘“‘ Fortasse omnes Columneae species in Jamaica obviae formis intermediis inter sese conjunctae sunt, C. jamaicensi excepta’’. There are indeed some specimens intermediate between recognized species, but these would appear to result from hybridization. All the species accepted by Urban have definite characters associated with distinctive
geographical ranges which justify their separation. C. hirsuta he regarded as a very —
variable species, within which, despite the limited material then available, he distinguished seven taxa. On the basis of the much more abundant and adequate material now at hand, it seems more reasonable to refer Urban’s varieties and forms of C. hirsuta to four species as follows:
C. hirsuta Swartz C. hirsuta Swartz C. hirsuta Swartz
var. genuina (p. 361) var. pallescens (p. 362) var. concolor (p. 362) var. subintegra (p. 362) forma wullschlaegelit (p. 362) = C. urbanii Stearn
I
forma hansenii (p. 363) = ? C. harnsu (Urban) Britton ex Morton
forma harrisit (p. 363) = C. harristi (Urban) Britton ex Morton
var. fawcettii (p. 363) = C. fawcettit (Urban) Morton
———— ee
THE JAMAICANSPECIESOFCOLUMNEA ANDALLOPLECTUS 187
In 1909 Urban added another species, Columnea brevipila, collected by Harris on Bluefields Mountains, Westmoreland.
Morton in 1944 (a, b) revised the West Indian species of Columnea and Alloplectus. With much more material for study than Urban had, most of it collected by Harris and by Britton, he described seven Jamaican species of Colummea in detail, raising two of Urban’s varieties of C. hirsuta to specific rank as C. harrisia and C. fawcetti and adding a new species, C. subcordata, from Windsor, Trelawny. Unlike Urban, he provided a key to all the West Indian species of both Columnea and Alloplectus. Examination of living material during my visit to Jamaica in 1956 (cf. Stearn, 1959) confirmed the taxonomic soundness of Morton’s work, already evident from a prior survey of herbarium material.
DEFINITION OF COLUMNEA AND ALLOPLECTUS
The difficulties of defining the American genera of Gesneriaceae, among them Columnea and Alloplectus, are reflected in the different views of authors on their generic limits. As stated by Leeuwenberg (1958 : 293), “‘ there are several cases of close inter-relationship of species usually placed in separate genera, rendering the boundaries between those genera vague or even untenable. Judging from these cases alone one would be compelled to unite large groups of genera or even complete tribes under one genus; this is particularly true for the Columneinae. The typical species of the genera are however so different that such a procedure would be quite undesirable. But part of those generic limits are based rather on convenient charac- ters than on conspicuous discontinuities of features. A consequence of this state of affairs is the presence of what may be called ‘ borderline species ’, i.e. species that have some characters of one and some of an other genus.”’ Such a species in Jamaica is Columnea jamaicensis, referred by Grisebach to Pterygoloma, by Fawcett to Alloplectus, by Urban to Columnea. The two Brazilian species (see above) on which Martius founded the genus Allopflectus in his Nova Genera et Species Plantarum Brasiliensium 3 : 53, t. 223 (1829) are markedly different from Columnea in their narrowly tubular rather ventricose corolla with the limb almost regular and very shortly lobed, i.e. shallowly 5-toothed or 5-lobed, the lobes almost equal, in their free anthers and their rather fleshy 2-valved capsule, but these distinctions have not remained clear-cut as more and more species of the group have been discovered. A few examples will serve to indicate the lack of correlated differences. Thus a species such as Columnea aureonitens Hook. (Bot. Mag. 73 : t. 4294 (1847)) has a narrowly tubular corolla with very short almost equal lobes, but four of the anthers cohere as in typical zygomorphic species of Columnea, while C. kalbreyert Hook. f. (Bot. Mag. 108 : t. 6633 (1882)) has a zygomorphic corolla but the four anthers are completely free from one another. C. guianensis Morton, illustrated by Leeuwenberg (1958 : 387, fig. 16), agrees with many typical species of Colwmnea in having the leaves of a pair markedly unequal in size, but its corolla is narrowly cylindrical with a small almost regular limb of five short erect lobes, the two dorsal ones however connate, and the anthers are free from one another as in a typical Alloplectus. The same is true of C. sanguinea (Pers.) Hanstein (Dalbergaria phaenicea Tussac) with
188 THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS
markedly unequal leaves but a regular cylindrical corolla and anthers cohering only in pairs. Alloplectus savannarum Morton (illustrated by Leeuwenberg, 1958 : 365, fig. 9) has unequal leaves and cohering anthers but a cylindrical corolla and a two- valved capsule as in typical Alloplectus. A possible distinction based on the form of the corolla tube, i.e. gradually widened and not contracted at the mouth in Columnea, ventricose with a contracted mouth in Alloplectus, is obscured by such species as Columnea filipes Oliver (Hook., Ic. Pl. 25 : t. 2428 (1896)) with ventricose tube but connate anthers. Characters of the fruit, an almost globose berry in typical Columnea, a capsule separating into two fleshy or thick valves in typical Alloplectus, might provide means of separation, but A. domingensis with a regular corolla has a berry-like fruit like a typical Columnea and the fruits of many species placed currently in Columnea or Alloplectus are unknown; the correlation of carpological, habit and floral characters in the group as a whole is accordingly doubtful. Attempts to separate the two genera by referring those with a coloured calyx and free anthers to Alloplectus, the type of which, A. /uirtellus, has a dark red calyx, and those with a green calyx and connate anthers to Columnea are frustrated by such a species as C. scheideana Schlecht. (illustrated in Bot. Mag. 70 : t. 4045 (1843)) with a large purplish-red calyx as in Alloplectus but a markedly zygomorphic 2-lipped corolla and connate anthers typical of Columnea. Anisophylly is likewise unavailing, since the type-species of both Columnea and Alloplectus have the two leaves of a pair equal.
In short there is no apparent discontinuity between Columnea and Alloplectus. O. Kuntze in 1891 (Rev. Gen. Pl. 2 : 470) united the two, together with Hypocyrta and Nematanthus, into one genus, Columnea (sensu lato), which would now include about 220 species, defined as follows: ‘‘ Ovarium omnino superum. Disci glandula postica magna ceterae 0 vel parvae. Stamina 4. Antherarum loculi paralleli. Filamenta basi dilatata unilateraliter connata cum corolla breviter connata. Calyx 5-partitus vel alte 5-fidus. Corolla apice variabilis exappendiculata. Fructus subbaccatus vel demum bivalvis.’’ This course certainly has simplicity to recommend it as regards nomenclature but obscures the special character of Coluwmnea (sensu stricto) as a highly developed group. The prevailing gamopetalous type of corolla found in Gesneriaceae and allied families consists of a fairly long tube with a spreading limb which is shorter than the tube and slightly or distinctly two-lipped; the two upper lobes form an upper lip while the lower lobe and the two lateral lobes together form a lower lip which serves as a landing platform for insects seeking nectar at the — base of the tube. The stamens are usually included within the corolla-tube and self- pollination is often possible; it occurs, for example, in Alloplectus species under — cultivation. The fruit is usually a capsule. AJdloplectus (sensu stricto) represents — this general unspecialized type. The corolla of Columnea (sensu stricto) represents a reversal of the trend towards development of an expanded lower lip able to support the weight of a bee. In Colwmnea the upper side of the corolla has become prolonged forward, the two uppermost lobes being fused and enlarged into a hood sheltering the exserted anthers and stigma; the two lateral lobes are reduced and they spread either sideways or upwards; the lower lip is also reduced and turned down or back. By these features, together with the horizontal poise of the flowers and their usually conspicuous red or yellow colouring, the flower of Columnea (sensu stricto) has
THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS 189
become perfectly adapted to pollination by nectar-seeking humming-birds on the wing (Text-fig. 5). The reduced lower lip and lateral lobes offer no obstacle to the bird as it flies under the lengthened upper lip and thrusts its beak and tongue down to the abundant nectar at the base of the tube, while its wings beat unimpeded on both sides of the lip. The anthers and stigma, being directly under the upper lip, are so placed that they touch the top of the head of the hovering bird. The coherence of the anthers ensures that they deposit the maximum of pollen in one area on the bird’s head where the stigma of another flower will first come into contact with it. The corolla as a whole is firm enough in texture to sustain vibration by the wing-beats of a humming-bird (mostly 20-25 beats per second, but ranging from 8 to 80 according to the size of the bird; cf. Greenewalt, 1960). Such a flower well exemplifies “ the syndrome of ornithophily ”’ as characterized by Faegri & van der Pijl (1966 : 109). This has been achieved by the successful association of several divergences from the almost regular flower typical of Alloplectus, which divergences have evidently also occurred separately and independently, as the anomalous intermediate species mentioned above indicate.
The proposal is accordingly made that Colwmnea should be defined more narrowly than is customary, by including within it only species of this ornithophilous habit, characterized by having a distinctly bilabiate red or yellow corolla, with the four upper (posterior) lobes fused and extended into a galea, the lower (anterior) lobe narrow and bent downwards, and having exserted stamens with the four anthers initially connate. Those species not satisfying this definition should be transferred to Alloplectus* or possibly other genera. Thus Alloplectus might likewise be more narrowly defined and restricted to species with a regular corolla and a capsular fruit akin to the type-species. To accommodate the species not fitting into this and Columnea (sensu stricto), Dalbergaria and possibly also Pterygoloma might then be restored.
A situation of this kind, in which a well-marked specialized offshoot, such as Columnea (sensu stricto), is still connected to a generalized stock, such as Alloplectus
* Acceptance of this viewpoint makes it necessary to transfer to Alloplectus the following West Indian, Guianaian and Venezuelan species allocated to Columnea by Morton and by Leeuwenberg: Alloplectus affinis (Morton) Stearn, comb. nov. Columnea affinis Morton in Fieldiana 28 : 529 (1953).
Alloplectus aureonitens (Hook.) Stearn, comb. nov. Columnea aureo-nitens Hook. in Bot. Mag. 73 : t. 4294 (1847).—Leeuwenberg in Act. Bot. Neerl. 7 : 383 (1958).
Alloplectus calotrichus (Donn. Smith) Stearn, comb. nov. Columnea calotricha Donn. Smith in Bot. Gaz. 40 : 9 (1905).—Leeuwenberg, tom. cit. : 385 (1958).
Alloplectus cubensis (Urban) Stearn, comb. nov
Columnea sanguinea var. cubensis Urban, Symb. Antill. 2 : 359 (1901). Columnea cubensis (Urban) Britton in Torreya 5 : 215 (1905).—Morton in Contr. U.S. Nat. Herb. 29 : 5 (1944).—Alain in Leon & Alain, Fl. Cuba 4 : 472 (1957).
Alloplectus guianensis (Morton) Stearn, comb. nov.
Columnea guianensis Morton in Bull. Torrey Bot. Club 75 : 564 (1948).—Leeuwenberg, tom. cit. : 386 (1958).
190 THE JAMAICANSPECIES OF COLUMNEA AND ALLOPLECTUS
(sensu lato), by a few species showing intermediate stages of development, does not permit of a solution both logical and convenient and hence proves disconcerting to a tidy-minded taxonomist intent on emphasis of discontinuity, but it is all the more interesting and worthy of study as indicating possible lines of evolution within the group. The most practical procedure would seem to be to recognize as genera both the large heterogeneous groups manifesting a wide range of characters occurring in many different combinations without consistent correlations and also at the same time the allied apparently monophyletic groups separated from them by possessing consistent correlations of characters whereby these form biologically important or conveniently distinguishable entities, the few intermediate species being referred to the more heterogeneous groups. The case of Alloplectus and Columnea parallels in some ways that of Lilimm and Nomocharis (cf. Sealy, 1950) where a solution according with this viewpoint has proved satisfactory.
The Jamaican species, apart from Columnea jamaicensis, fit readily into Columnea with a strongly bilabiate corolla and Alloplectus with a regular cylindrical corolla. C. jamaicensis is like A. pubescens and A. grisebachianus in its low creeping habit and has the bilabiate character of the corolla less marked than in typical Columnea, but the tube expands gradually, the upper lip consists of two fused lobes with two spreading lateral lobes, and the four anthers are connate. These associated floral characters justify the allocation of C. jamaicensis, and hence of Pterygoloma, to Columnea, as defined here, even though on the sum of its other characters, as is shown by taxonometric analysis, it has more in common with the Jamaican species of Alloplectus.
CHARACTERS DISTINGUISHING THE SPECIES
The Jamaican species of Columnea and Alloplectus are soft-wooded plants of —
creeping or sprawling habit, rather sparingly branched, with stems which may even reach a length of several metres in some robust specimens, the leaves then being clustered near the growing tip and the branchlets marked with the conspicuous scars of the fallen leaves. Columnea jamaicensis (Pl. 14), Alloplectus pubescens and A. grisebachianus have slender creeping branchlets only 1-3 mm. thick, whereas the others have much stouter branchlets about 5-10 mm. thick, initially erect but drooping downwards with age when the plants grow as epiphytes on trees. The leaves are paired, those of a pair being almost equal in size and shape, as in C. urbana (Pl. 16), to markedly unequal, as in C. proctorii (Pl. 17) and C. rutilans (Pl. 21), the smaller leaf of the pair having a shorter petiole as well as a shorter blade and often falling earlier than the larger leaf. This anisophylly, although used by Leeuwenberg
(1958 : 295) in his study of Guiana Gesneriaceae to separate Alloplectus sect. Allo-
plectus with the leaves of a pair almost equal from Coluwmnea sect. Collandra with leaves of a pair markedly unequal, varies in the Jamaican species, being apparently constant in some, such as C. hispida, C. subcordata and C. harrisii, but variable in
others such as C. fawcettit and C. hirsuta, which may have the leaves of a pair almost —
equal or markedly unequal. The difference between the large and small leaves is particularly evident in C, rutilans, C. proctorti and C, subcordata,
OO —
a
.. > ¢
THE JAMAICANSPECIESOFCOLUMNEA ANDALLOPLECTUS 191
The proportions as well as the actual dimension of the leaf-blade provide characters of some diagnostic value, as Morton demonstrated by measuring 50 representative - leaves of each species when he had enough material available. Thus in Columnea jamaicensis, C. hirsuta and C. subcordata, the blades average about two times as long as broad, in C. fawcettii, C. brevipila and C. harrisit, between 2-4 and 2-7 times as long as broad, and in C. rutilans and C. argentea, about 3-4 to 3-7 times as long as broad. The leaf-base is remarkably asymmetric in C. harris and C. hispida C. rutilans stands apart from the others in having the underside of the leaf either completely red or else conspicuously red-veined, and this red colouring is evident in some of the hybrids between C. rutilans and C. urbanit.
Diversity of hair-covering provides features of diagnostic value, notably the hair- covering on the upper surface of the leaf-blade, where four main types of hairs may be distinguished (Text-fig. I, p. 192):
1. Hairs erect, 8-g-celled, red, to 3-5 mm. long, e.g. Columnea Iispida (Text-fig. 1A), in which also occur minute erect colourless, 3-celled hairs.
2. Hairs erect, 5—10-celled, colourless, 0-8—2:5 mm. long, e.g. C. hirsuta (Text-fig. IC), C. proctorit, C. subcordata, C. urbanii (Text-fig. 1B); in C. hirsuta these are associated with minute almost appressed 2-celled hairs.
3. Hairs bent forward and somewhat or almost appressed, 4-6-celled, colourless, scarcely 1 mm. long, e.g. C. argentea (Text-fig. 1F), C. harrisw (Text-fig. rE) and C. rutilans (Text-fig. 1D), in all of which they are associated with minute almost appressed 2-celled hairs.
4. Hairs bent forward, almost appressed, 2-celled, colourless, 0-2-0-4 mm. long, exclusively present in C. brevipila (Text-fig. 1D) and C. fawcettw (Text-fig. 1H), associated with longer hairs in C. argentea (Text-fig. 1F), C. hirsuta (Text-fig. IC), C. harristi (Text-fig. 1E), and C. rutilans (Text-fig. 1G).
Suppression of the long hairs in C. hirsuta would produce leaves like those of C. fawcetti.
The flowers are axillary, with one to five in an axil, on pedicels 1-3 cm. long, varying from species to species in the type of hair-covering.
The most useful characters for specific distinction are provided by the calyx (Text- fig. 2). The segments may increase a little in size after flowering as the fruit matures but retain their shape, being, for example, about 12 mm. long in flower but 16 mm. long in fruit in Columnea jamaicensis, about 2:5 cm. in flower but 3-5 cm. in fruit in C. rutilans. They are mostly between 2 and 3cm. long. C. subcordata (Text-fig. 2G) stands apart from the other species, in which the five segments are free almost to the base, by having four segments united for about the lower third of their length and the fifth almost free.
The segments are linear and entire in C. hispida (Text-fig. 2A) and lanceolate or narrowly lanceolate and entire or scarcely toothed in C. argentea (Text-fig. 2C), C. brevipila, C. harrisii, C. proctorii (Text-fig. 2B), C. urbanit (Text-fig. 2H) and A. pubescens (Text-fig. 29G). In C. hirsuta and C. fawceettii (Text-fig. 2D; Text-fig. 28B) the segments bear one to three remote but usually well-marked teeth on each margin. In C. rutilans (Text-fig. 2F) and A. grisebachianus (Text-fig. 29C) they are
192 THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS
laciniate in the lower part, with 3-5 prominent teeth on each margin. These differences in shape, together with differences in hair-covering, enable most species to be recognized by the calyx alone. The hairs of the calyx are red in C. hispida, C. fawcettii, C. hirsuta, C. hispida and C. rutilans, colourless in the others.
The corolla as mentioned above (p. 186) is small, narrowly tubular and regular in Alloplectus grisebachianus (Text-fig. 29A, B) and A. pubescens and slightly 2-lipped
A 'B ‘D
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WW
urbanii
WK ee KK é
: argentea
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Fic. 1. Hairs on upper leaf-surface of: A, Columnea hispida Swartz; B, C. urbanii Stearn; C, C. hirsuta Swartz; D, C. brevipila Urban; E, C. harrisit (Urban) Morton; F, C. argentea Griseb.; G, C. rutilans Swartz; H, C. fawcettii (Urban) Morton. (All drawn from living plants cultivated at Kew.)
———e—e—e—eEeEeeEeeeeeeeeeeee ee ee
a
THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS 193
in Columnea jamaicensis (Text-fig. 28H), but larger and strongly 2-lipped in the other species. It is completely yellow in C. argentea (Text-fig. 5), C. brevipila, C. hispida C. subcordata and C. urbanii (Text-fig. 28G) but longitudinally striped with red in the others, including many of the hybrids between C. rutilans and C. urbanit.
The filaments of the stamens are glabrous in Columnea argentea and C. brevipila but pubescent in most other species.
Fic. 2. Calyx of: A, Columnea hispida Swartz (Stearn 449); B, C. proctorii Stearn, with apex of pedicel (Stearn 451); C. argentea (Stearn 359); D, C. fawcettit (Stearn 157); E, C. jamaicensis Urban, a (Wullschldgel 1290), b (Stearn 928); F, C. rutilans Swartz (Stearn 453); G, C. subcordata Morton (Stearn 472); H, C. urbanii Stearn (Stearn 361).
1904 THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS
Certain species can thus be easily recognized by distinctive and unusual characters. For example, Columnea jamaicensis has small leaves about 3-5 cm. long, a short calyx and a small corolla not strongly bilabiate. C. vutilans stands apart from other species in having its leaves red below and its calyx-segments partly laciniate. C. subcordata has a bilabiate calyx with four segments fused for their lower third. C. fawcett with toothed calyx-segments and C. brevipila with entire calyx-segments differ from other species in having only very short appressed 2-celled hairs on the upper surface of the leaf. C. argentea has relatively narrow leaves silvery-grey with long somewhat appressed sericeous hairs. C. hispida is notable for the long erect reddish hairs covering the upper side of the leaf as well as the calyx and for the relatively small corolla.
CyTOLOGY By Brian D. MORLEY
The chromosome numbers of species of Columnea have been reported by Rogers (1954), Eberle (1956), Fussell (1958), Lee (1962a, b, 1964, 1966, 1967), Morton (1963), Lee & Grear (1963) and Morley (1967). Investigation of the chromosome numbers of ten Jamaican species and four natural hybrids selected from a hybrid swarm between C. rutilans and C. urbani confirms the reports for C. brevipila, C. fawcetti and C. hispida by Lee (1964, 1966) and shows that all the Jamaican species havea diploid number of 2n = 18 and that their basic number is the same as that of other Columnea species, x = 9. The plants studied were as follows:
Plant Species Locality reference no. 2n C. hirsuta Hardwar Gap, St. Andrew—Portland. Morley 33 18 C. fawceettit Mount Diablo, St. Ann. Morley 2 18 (coll. E. Lodge) C. rutilans Ramgoat Cave, Trelawny. Morley 3 18 Morley 35 18 C. rutilans Near Catadupa, St. James. Morley 20 18 C. argentea Shooters Hill, Manchester. Morley 36 18 C. urbana Top Hill, Manchester. Morley 14 18 Morley 442 18 C. urban Near Newport, Manchester. Morley S.6 18 @ C. urbanu Rose Hill Village, Manchester. Morley S.7 18 C. proctoria Near Troy, Trelawny. Morley S.17 18 (coll. C. D. Adams) C. proctori1? Ramgoat Cave, Trelawny. Morley 5 18 g C. hispida Near Catadupa, St. James. Morley 40 18 © C. harris Seven Rivers, St. James. Morley 23 18 C. harrisw Near Leamington, Westmoreland. Morley S.11 18 © C. brevipila Cho Cho Gulley, Westmoreland. Morley 24 18 | C. jamaicensis Near Leamington, Westmoreland. Morley 27 18
(coll. C. D. Adams)
THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS 195
C. jamaicensis Near Leamington, Westmoreland. Morley S.10 18
C. rutilans x Top Hill, Manchester. Morley 16, 44,45 18 urbanu hybrid 4
C. rutilans xX Top Hill, Manchester. Morley 47 18 urbanu hybrid 6
C. rutilans x Top Hill, Manchester. Morley 15, 46 18 urbanu hybrid 8
C. rutilans x Top Hill, Manchester. Morley 48 18 urban hybrid 11
A. grisebachianus Mount Diablo, St. Ann. Morley s.n. 18
(coll. E. Lodge)
Although the chromosomes are small, they display sufficient morphological diversity to allow recognition of the following three karyotype series. A metacentric position is defined as an approximate position of 50% along the length of a chromo- some and submetacentric as 60 %-—70 % along a chromosome.
Series r. Karyotypes with large chromosomes, i.e. longer and thicker, where the smallest chromosome is always longer than broad. Two pairs of chromo- somes with submetacentric centromere position, seven pairs of chromo- somes with metacentric centromere position; see Text-fig. 3.
Species: C. hirsuta, C. fawcettii, C. urbani: (from an area north of Mandeville).
Series 2. Karyotypes with large chromosomes, i.e. longer and thicker, where the smallest chromosome is always longer than broad. Three pairs of chromosomes with submetacentric centromere position, six pairs of chromosomes with metacentric centromere position; see Text-fig. 3. Species: C. urbanit (from an area south of Mandeville), C. brevipila, C.
rutilans, C. jamatcensis.
Series 3. Karyotypes with small chromosomes, i.e. shorter and thinner, where the smallest chromosome is always as long as broad. Three pairs of chromo- somes with submetacentric centromere position, six pairs of chromosomes with metacentric centromere position; see Text-fig. 4.
Species: C. argentea, C. hispida, C. harrisit, C. proctorit.
Series 1 karyotypes characterize the species Columnea hirsuta and C. fawcettit, which are morphologically and biochemically distinct from other Jamaican species and are only found in eastern Jamaica. Most C. urbanit karyotypes belong to Series 2, with the exception of one plant which had a Series 1 karyotype.
Although Columnea jamaicensis is included in Series 2, it is unlike any other Jamaican species, having a different phenotype and a karyotype with two pairs of satellites. The metacentric satellite pair of C. urbanii (66.1460) requires further analysis.
BOT. 4, 5 12
196 THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS
Karyotypes of Jamaican species of Columnea. x2500
A 66-991 hirsuta. PERS RSRRSERVFRRASS ot 7g) 66-1627 fawcettii. A aoe tsuge Z HAR FRRR ORAS 66:1042 urbanii. ee oo te Ra 0 a woes Whe SIRS SBesss 66-1460 urbanii. a ‘ VE Peas SERS SR Seuzes oe ) 66:960 brevipila. m HO ya Va Mm Si Obes sa SS SS as iaee ie) 661084 rutilans.
es F Aa RA AS CHER SAMS eee 66-1468 jamaicensis.
esas SSRs ses eee
~
Fic. 3. Karyotypes of some Jamaican species of Columnea.
The hybrids between Columnea rutilans and C. urbanii have karyotypes inter- mediate in morphological details between those of the parents; see Text-fig. 4.
The Jamaican species with smaller and more asymmetric karyotypes, i.e. Columnea argentea, C. hispida and C. harrisii in Series 3, have the more limited distributions in Jamaica.
While, on evidence so far available, no changes in chromosome number have accompanied the evolution of Columnea, changes in chromosome structure have occurred, as is evident from gross morphological studies of Jamaican and Central American Columnea karyotypes.
THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS
66-1119
Karyotypes continued.
argentea.
S2usas SESS ee 3u cata
66-1180
hispida.
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66-1465
66-1152 Ssansa
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CHEMOTAXONOMY
By Brian D. MORLEY
197
te Salaas
SGIUGAH
Karyotypes of some Jamaican species and hybrids of Columnea.
Cultivation of the Jamaican species of Columnea at Kew enabled Dr. J. B. Harborne to carry out on fresh leaves three chromatographic tests for flavones, orobanchin, and cinnamics, and also to test for saponin.
The substances detected
198 THE JAMAICAN SPECIES OF COLUMNEA ANDALLOPLECTUS in these tests were characterized as follows:
I. Flavone a: Rf 0-35 in butanol-acetic acid-water, Rf 0-05 in 5 % acetic acid; colours in ultra-violet light +- NH: dark brown-bright yellow; possibly luteolin 7-glucoside.
Flavone b: Rf 0-13 in 5% acetic acid; colours as flavone a.
Flavonec: Rf 0-23 in 5% acetic acid; colours as flavone a.
Flavone d: Rf 0-25 in butanol-acetic acid-water, Rf 0-05 in 5 % acetic acid; colours as flavone a, but more intense.
II. Ovobanchin: A complex ester of caffeic acid (see Harborne, 19662). III. Czinnamics,
Blue Spot: Rf 0-69 in butanol-acetic acid-water; colours in ultra-violet
light: colourless—blue; possibly a p-coumaric acid ester.
Cinnamics, Turquoise Spot: Rf 0-24 in butanol-acetic acid-water. IV. Saponin: Detected by the presence of frothing when an aqueous leaf
extract was shaken up.
These tests yielded the following results:
I II III IV Flavones Orobanchin Cinnamics Saponin fe =e A f; ts an Plant reference Blue Turquoise Species number Se {Dieta t a spot spot C. hirsuta Morley 33 + + —- — os —- — -- C. fawcettit Morley 34 + + —- — ao —_ —_ — C. proctorit Morley S.17 - —- —- — + as -- + C. urbanit Morley 12 - - - = + + 4- + C. harrisit Morley 6 + + — + + + + — C. argentea Morley 36 - — tr — a + _ C. hispida Morley 40 — tr tr — + + _ —_ C. rutilans Morley 3 —- tr —- = a — _ _ C. brevipila Morley 24 tr -—- —- — ++ _ _ _ C. jamaicensis Morley 27 + -—- — — “ - Where + = present; — = absent; tr = trace.
The leaf-flavone attributes of Columnea hirsuta and C. fawcettii are similar and distinct from those of the other species; they support their taxonomic separation from other Jamaican species on the basis of calyx characters. Certain cytological features of the two species also distinguish them from the others in Jamaica. C. urbanit and C. proctorit have similar leaf-flavone attributes, and this parallels their morphological similarities. The leaf-flavone attributes of C. urbanii and C. brevipila are nearly similar, which adds some evidence for their general affinity to that provided by morphological and cytological data. The remaining species show a reticulate pattern of flavone attributes. C. harrisii was found to have a flavone attribute not observed in leaves of the other Jamaican species tested.
THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS 199
The tests for cinnamics and saponin give results in accordance with those for flavones in the case of the species pairs Columnea Mirsuta and C. fawcettw, and C. urbanit and C. proctorit, the first pair being negative and the second positive. The resemblance in flavone attributes between C. urbani and C. brevipila is not paralleled in the case of cinnamics and saponin. C. rutilans was found to have columnin in the leaves, a 3-desoxyanthocyanin, as reported by Harborne (19660), which is responsible for the red pigmentation of the abaxial leaf surface in this species.
Columnea argentea and C. harrisit have common saponin and cinnamics attributes paralleling the densely hairy leaves found only in these species; thus the sensitivity of the biochemical tests is shown by their discrimination of particular species within a group of closely related endemic species. As noted above, they provide further evidence of affinity between C. hirsuta and C. fawcetti1, between C. urban and C. proctorii, and to a lesser extent between C. brevipila and C. urbani1, and between C. argentea and C. harnisit.
Investigation of the cinnamics and saponin attributes of material from a hybrid swarm at Top Hill, Manchester, gave the following results. The hybrids are arranged in a series suggested by their degree of morphological agreement relative to their parents; the biochemical attributes broadly parallel this arrangement.
Cinnamics Plant r A ~ reference Blue Turquoise Taxon number spot spot Saponin C. rutilans Morley 19 — — — Hybrid 11 Morley 48 + — — Hybrid to =6©Morley 46 — _ alr Hybrid 6 Morley 47 + _ _ Hybrid 4 Morley 8 a — + Hybrid 1 Morley 43 + = + C. urbani = Morley 31 + + +
The leaf-flavone attributes of Columnea rutilans and C. urbanii are too similar to warrant their examination in the hybrids, but the saponin and cinnamics attributes of the hybrids were found to differ. The turquoise spot attribute in C. urbani1 was not found in any of the hybrids and it may be genetically recessive. It is possible that backcrossing has occurred in the hybrid swarm and this may have complicated a simple pattern of gradation of the parental saponin and cinnamics attributes as found in hybrids 10 and 11. With a sufficiently comprehensive set of tests it might be possible to trace a backcrossing system in the hybrid swarm, particularly if synthetic backcrosses were made and used as marker plants.
Harborne (19662) reported that biochemical tests distinguished between the sub- families Gesnerioideae and Cyrtandroideae. The present study has shown that biochemical tests are also sensitive at specific and intraspecific levels; leaf flavones provide attributes for the identification of Jamaican species of Colummnea, and leaf- saponin and cinnamic attributes identify different hybrid phenotypes in a hybrid swarm,
200 THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS
Harborne (19660) has suggested that there may be correlation between the posses- sion of 3-desoxyanthocynanins and ornithophilous pollination in the Gesnerioideae, as brightly coloured flowers are attractive to birds and plants synthesizing those pigments in larger quantities would have a selective advantage. |
ECOLOGY AND DISTRIBUTION
All the Jamaican species of Columnea are endemic. Although they do not vie in~ brilliancy of colour with such conspicuous Central American species as C. crassifolia, — C. gloriosa, C. hirta, C. microphylla and C. oerstediana (cf. Moore, 1957; Raymond, 1965), they are nevertheless attractive and interesting plants. They often grow as epiphytes high on trees and are then usually associated with bromeliads, notably clumps of Hohenbergia, thus easily passing unnoticed, but they also grow on rocks or stone walls when the rainfall is high enough to permit the luxuriant growth of mosses and the deposit of humus. Evidently they need good drainage and a moist or intermittently moist atmosphere. |
The large zygomorphic yellow or red flowers have protruding stamens and style, and secrete nectar at the base of the corolla-tube and are visited by humming-birds. Three species of humming-bird (Trochilideae) inhabit Jamaica (cf. Bond, 1947), namely the Jamaican Mango Humming-bird (Anthracothorax mango) with a beak about 2-5 cm. long, the Streamertail (Tvochilus polytmus) with a beak about 2 cm. long and the Vervain Humming-bird (Mellisuga minima) with a beak about 9 mm. long. Of these, the Streamertail has been observed visiting flowers of Columnea (cf. Morley, 1966). The corolla-tube in most Jamaican species of Columnea averages about 2 cm. long; indeed the general size of the corolla appears to be correlated with the length of the beak and head of the Streamertail (Text-fig. 5), which has thus probably been a major factor in the evolution of the Jamaican species, apart from C. hispida and C. jamaicensis. If the Streamertail became extinct, most of the Jamaican species of Columnea would probably also become extinct.
Humming-birds are strongly territorial in their habits. Thus a single bird feeds within a limited area and drives other humming-birds from it, thereby restricting pollination of ornithophilous flowers to plants within this feeding area (cf. Pitelka,
Fic. 5. Flower of Columnea argentea and head of Streamertail Humming-bird (Trochilus polytmus).
THE JAMAICANSPECIESOFCOLUMNEA ANDALLOPLECTUS 201
1942). Grant & Grant (1966 : 55), for example, record humming-birds each indi- vidually guarding its own patch of Zauschneria latifolia in flower and chasing other humming-birds from its preserve. These territorial habits of humming-birds presumably favour the inbreeding of plant colonies, which may be very important in the earlier stages of species development. As, however, unlike bees, humming-birds have no flower constancy but dart indiscriminately from one nectar-yielding species to another, they are likely to cross-pollinate quickly any two species flowering at the same time in a given feeding area. The numerous seeds contained in the white or pink berries of Columnea are presumably distributed from tree to tree by other birds; the association of Columnea with bromeliads may possibly result from birds drinking water and collecting insects at the bromeliad pitchers. Ants, however, find the seeds attractive, according to Morley, as they do those of other Gesneriaceae, and thus could aid dissemination over the same tree or rocky slope.
Although the Jamaican species have much the same ecological preferences, they differ markedly in range (Text-figs. 6-19) and exhibit the general tendency of the family Gesneriaceae towards much local endemism. Longitude 77° 20’ approxi- mately halves the island of Jamaica. West of it occur Columnea rutilans, C. harrisit, C. jamaicensis, C. urbanii (with a slight extension of range east of it), C. hispida, C. proctorii, C. brevipila, C. subcordata and C. argentea, to the east of it C. fawcettiw (with a slight extension of range west of it), C. hirsuta and Alloplectus pubescens, as also Achimenes erecta (Text-fig. 20), the early history of which is entangled with that of Columnea. Alloplectus grisebachianus has a central position. Hybridization is easily effected in cultivation but rarely occurs in the wild because, although the ranges of some species overlap, usually only one grows in a particular locality. Dr. Brian Morley collected at Top Hill, Manchester, near Walderston (Text-fig. 16), in a man-disturbed habitat, where both C. rutilans (Text-fig. 17) and C. urbanit (Text-fig. 9) occur, a series of intermediate plants evidently resulting from their hybridization. There may also be some ecological separation. Thus C. hirsuta (Text-fig. 14) grows mostly between g00 m. (3 ,000 ft.) and 1,500 m. (5,000 ft.), although descending in Portland and St. Thomas to 600 m. (2,000 ft.). C. fawcetti (Text-fig. 15), which is closely allied and has been long confused with it, grows mostly between 400 m. (1,400 ft.) and goo m. (3,000 ft.) in the same general area as C. hirsuta but extends far beyond the range of this. Some species are clearly very restricted in range, e.g. C. argentea (Text-fig. 12) and C. hispida (Text-fig. 7), though the reasons are not evident.
TAXONOMETRIC SURVEY
Before the computer-aided techniques of numerical taxonomy (taxonometrics) can be used with confidence to classify taxa belonging to difficult or obscure groups, by assessing the relative over-all similarity of these taxa and then arranging them hierarchically in accordance with their degree of similarity and divergence, it is necessary to test the validity and sensitivity of such techniques by applying them to better understood groups already satisfactorily classified by traditional methods and represented by adequate material. A computer-aided technique which for such a group produces a classification essentially similar to or as reasonable as a well
202 THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS
CoLuMNEA
7 so" 78 jo” 77° jo ]
JAMAICA JAMAICA
$——_—______
= | |
JAMAICA
ae" 78° jo” 77° ” je” 78° jo" 37°. 3
JAMAICA
JAMAICA
ah. e 78° se 772 ae T
JAMAICA | JAMAICA
Se eS
2a,
12 argentea 13 brevipila
Fics. 6-13. Distribution of Columnea in Jamaica: Fig. 6, C. jamaicensis ; Fig. 7, C. hispida; Fig. 8, C. proctorit; Fig. 9, C. urbanii; Fig. 10, C. subcordata; Fig. 11, C. harvrisii; Fig. 12, C. argentea; Fig. 13, C. brevipila,
THE JAMAICAN SPECIESOFCOLUMNEAANDALLOPLECTUS 203
CoLuMNEA eo 78° 30° 77° eo so 78° jo 730 jo" JAMAICA JAMAICA -- - - ---——_}+—- —|s"
i y oe ae jo" 78° 3 7 3 15 fawcettii Je. 78° 50" : (ihe je je 13° 10" ize jo” JAMAICA JAMAICA 30 +——__-__] 46] a -—- -—_ —4 —_—_3e “| — has 4 30° 7 yo" 77° 30° jo" 78 jo 77 3 16 hybrid (rutilans x urbanii) 17 rutilans ALLOPLECTUS so" 78° so 77° 30° so 78° io" iE jo"
j | | JAMAICA JAMAICA jo} —-—-— -—___._f—___7 == 4 Meee jy Jo a — {39 I le wince I
jo 78 30" 77 3 40 78 30" 77 3 18 grisebachianus 19 pubescens ACHIMENES Jo 78° je" 77° 30° | | JAMAICA 2 ——————— er i er Wee ie Seat 3°
20 erecta Fics. 14-20. Distribution of Columnea, Alloplectus and Achimenes in Jamaica: Fig. 14, Columnea hirsuta; Fig. 15, C. fawcettii; Fig. 16, C. rutilans x urbanii; Fig. 17, C. rutilans; Fig. 18, Alloplectus grisebachianus; Fig. 19, A. pubescens; Fig. 20, Achimenes erecta.
204 THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS
grounded existing one obviously commends itself for use in other investigations. Conversely a technique which makes nonsense when dealing with the known is unlikely to do better with the unknown. Application of taxonometric methods to different groups has given results of varied acceptability. Thus a comparison by Kendrick and Weresub (1966) of members of orders within the Basidiomycetes, using 67 haphazardly assembled equally weighted characters, resulted in a haphazard classification, seemingly of no value for any purpose. Watson, Williams & Lance (1967), using 38 characters, produced a classification of the genera of Ericales which bears only a limited resemblance to the one generally accepted; they regard, however, the new scheme as more convincing, “ since it provides a convenient summary of the information put into it and makes sense of observations other than those on which
| .
it was based, while the traditional scheme does neither.’’ Sheal’s taxonometric — survey of mesostigmatid mites of the Hypoaspis-Androlaeleps complex, using 62 — characters, gave results in good agreement with a recent revision by traditional
methods (Sheals, 1965; cf. also British Museum (Natural History), 1966). Likewise
an arrangement of the genera of megascolecoid earthworms (Oligochaeta Megas- —
colecidae) by computer technique, using 43 characters, gave results coinciding largely with one of the previously proposed classifications (Sims, 1966).
A major difficulty in numerical taxonomy is finding enough attributes for computa- tion; obviously the smaller the number, the greater the risk of results distorted by abnormalities. Between 40 and 100 characters have been postulated as necessary to give results statistically valid (Sokal & Sneath, 1963), but in practice it is often difficult or impossible to provide so many characters; the diversity within small groups does not stretch that far. Hence it is desirable to ascertain empirically whether a much smaller number of characters will yield results which, even if not so detailed and meaningful as those derived from a large number of characters, can
nevertheless be helpful by grouping together organisms with over-all resemblance —
and separating those which differ. It is also desirable to ascertain the extent to which a reduced set of selected characters and an equally reduced set of characters picked at random agree in their results with each other and with a maximum set.
The information available about the Jamaican species of Columnea and Alloplectus seemed to be broadly enough based and adequate enough in quantity to provide such comparisons and to test the ability of a computer-aided taxonometric procedure to classify these species. By including cytological, biochemical and anatomical data along with gross morphological data it was possible to tabulate 51 characters varying from species to species, i.e. constant within a given species but not for the whole group. This full set of characters was the basis of one computation; 27 of them had been used in a key and thus formed a reduced set weighted in favour of readily observed macroscopic characters, on which indeed most angiosperm taxonomy is necessarily based. These were used for a second computation. For comparison,
27 characters were taken at random, i.e. by assigning number to each of the full 51
characters and picking blindfold 27 cards out of 51 bearing these numbers; 13 were found to coincide with those of the other set. These were used for a third computa- tion. Such characters for coding purposes may be divided into three groups (cf. Watson, Williams & Lance, 1967): qualitative (or discrete) two-state (or dichotomous),
THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS 205
in which a character exists in one or other of two states; qualitative multi-state, in which a character exists in one of three or more states; quantitative (or continuous) comprising characters within which several measurable or assessable though less clear-cut and sometimes overlapping states may be distinguished. For some characters their allocation to one or other of these groups is anpteny Those used in this survey were as follows:
TWO-STATE
1.* Stem thickness: less than 5 mm.; more than 5 mm.
2.* Leaves: isophyllous; anisophyllous.
3.* Leaf base: symmetric; asymmetric.
4.* Leaf underside: green; red.
5.* Hairs of leaf: colourless; red.
6. 2~-3-celled hairs on leaf upperside: absent; present.
7. Other hairs on leaf upperside: 4—7-celled; 8—10-celled.
8.* 2-3-celled hairs on leaf underside: absent; present.
9. Other hairs on leaf underside: 4-7 celled; 8—10-celled. 10. Pedicels: always 1; 1-several.
11. Hairs of pedicels: colourless; red.
12.* Calyx-segments: all free; 4 fused, 1 free.
13.* Calyx: green; wholly or partly red.
14.* Hairs of calyx: long; short.
15. Long hairs of calyx: erect; otherwise.
16. Short hairs of calyx: few; many.
17.* Hairs of calyx: colourless; red.
18.* Corolla: completely yellow; yellow and red.
19.* Corolla length: 3 cm. or less; 4 cm. or more.
20.* Corolla-tube length: 1 cm.; I-4—2:5 cm.
21. External hairs of corolla: 2-6- celled; 6—12-celled. 22. External hairs of corolla: colourless; red.
23. External hairs of corolla: appressed; spreading. 24. External hairs of corolla: few; many.
25. Internal hairs of corolla: 1-3-celled; 3—6-celled. 26.* Filaments of stamens: glabrous; hairy.
27. Non-glandular hairs on style: absent; present.
28. Submetacentric chromosome pairs: 2; 3.
29. Saponin: absent; present.
30.* Corolla: regular; zygomorphic.
31. Anthers: almost square; oblong.
MULTI-STATE
32. Flavone: none; A; B; C.
206 THE JAMAICAN SPECIESOFCOLUMNEAANDALLOPLECTUS
QUANTITATIVE
33.* Leaf-blade length|breadth: 1-5-2; 2:4—2°7; 3°4-4.
34.* Leaf-blade length: under 5:5 cm.; 5*5-I12 cm.; more than 12 cm.
35. Length of long petiole: almost nil; to 1 cm.; to I-5 cm.; to 3-5 cm.
36.* Vein pairs of leaf: 3; 4-5; 5-7; 7-8.
37.* Hair density on leaf upperside per sq. cm.: 0-25; 26-100; 101-400; 401-800; 801-1,600.
38.* Hairs on leaf upperside: erect; strongly curved; appressed.
39. Hair density on leaf underside per sq. cm.: 0-25; 26-100; 101-400; 401-800; 801—1,600.
40.* Hairs on leaf underside: erect; strongly curved; appressed.
41. Palisade layer: in upper 4; at middle; in lower 3.
42. Cell width in palisade layer: narrow; medium; broad.
43. Pedicel length: 2-10 mm.; to 25 mm.; to 30 mm.
44.* Hairs on pedicel: appressed; ascending; erect.
45.* Length of calyx-segments: I-I-4 cm.}; I-5-I-Q cm.; 2—2°4 cm.
46.* Calyx-segments: entire or almost so; toothed; laciniate.
47.* Length of upper lip of corolla: nil; 2-2-4 cm.; 2°5-3 cm.; 3:I-3:4 cm.
48. Length of mid lobe of corolla: nil; 3-5 mm.; 10-15 mm.; 16-18 mm.
49. Length of lower lip of corolla: nil; 1 cm. or less; I-5 cm. or more.
50.* Ovary: glabrous; hairy above; completely hairy.
51. Cinnamic: absent; scanty; abundant.
These characters were ascertained from living and herbarium material and coded numerically, e.g. hairs colourless = 0, hairs red =1, hairs appressed = 0, hairs strongly curved = 1, hairs erect = 2, then tabulated as a species/data matrix, all being included in the full matrix, but only the 27 marked * in the reduced matrix of selected characters (reduced key matrix). For many quantitative characters, an average from a number of specimens was taken and the coding based on this. Computation of the above data on an Orion electronic computer at the Rothamsted Experimental Station, Harpenden, Herts., England, by G. Ross, using Gower & Ross’s program CLASP, gave the “ median sort ’’ results expressed as dendrograms (phenograms) in Text-figs. 21-23.
Since certain species are obviously closely linked in their morphological characters, e.g. (1) Columnea hirsuta and C. fawcettii, (2) C. subcordata and C. proctorit, (3) C. argentea, C. brevipila and C. harrisit, and others are relatively isolated, e.g. C. hispida, C. jamaicensis, A. grisebachianus, it was postulated before computation that, unless the resulting dendrograms linked and separated such species accordingly, they would have little or no relevance to the actual situation within the genus. In fact all three dendrograms make these associations and distinctions. The dendro- gram (Text-fig. 21) from the full matrix (51 characters) achieves, however, the most satisfactory differentiation, then comes that from the reduced random matrix (Text-fig. 22), both being superior to that from the reduced key matrix of 27 charac- ters (Text-fig. 23). Thus, whereas the dendrogram from the reduced key matrix
Ee
THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS 207
(Text-fig. 23) simply places together at a similarity coefficient (phenon level) of 80% C. fawcettii, C. harris, C. hirsuta, C. proctoria and C. subcordata without further distinction and separates from them at the 75% level C. argentea and C. brevipila, the full one and the reduced random one put these species in three groups, the full one (Text-fig. 21) at the 75 % level, the random (Text-fig. 22) at the 70 % level, i.e. (1) C. hirsuta and C. fawcetti together, (2) C. subcordata and C. proctori with C. urbanit, (3) C. argentea and C. brevipila with C. harrisi1; the dendrogram from the full matrix goes further by associating C. hirsuta, C. fawcettu and C. rutilans. In
65 70 75 80 85 90 95
i l l ambiguus
pubescens
grisebachianus
hispida
argentea
brevipila
harrisii
fawcettii
hirsuta
hybrid 6
rutilans
hybrid 10
hybrid 4
urbanii
proctorii
subcordata
jamaicensis
Fic. 21. Dendrogram showing the percentage of similarity of 17 species and hybrids of Columnea and Alloplectus computed from a matrix of 51 characters (cf. Fig. 24).
208 THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS
these instances, a taxonometric method has thus created from an assemblage of numerous unweighted characters a classification basically the same as one made by traditional methods. All dendrograms emphasize the distinctness of C. jamatcensis from other Jamaican species of Columnea by separating it at the 65% (full and reduced random matrices) and 70% (reduced key matrix) levels; no dendrogram, however, indicates any resemblance of this to the Alloplectus species A. ambiguus, A. grisebachianus and A. pubescens. All associate the hybrids with one or other of their parents. The superiority of the dendrogram based on 27 random characters (Text-fig. 22) over that based on 27 selected characters (Text-fig. 23) would appear to result from the first including biochemical, anatomical and cytological characters as opposed to the purely morphological characters of the second. The difference between them is in line with the general view of taxonomists that certain characters
65 70 75 80 85 90 95 100
| | | | | ] | ]
ambiguus
pubescens
grisebachianus
harrisii
argentea
brevipila
fawcettii
hirsuta
hybrid 6
hybrid 10
rutilans
hybrid. 4
urbanii
pee een mee Oe ee Oe L ns stibcordata
proctorii
jamaicensis
hispida
Fic. 22. Dendrogram showing the percentage of similarity of 17 species and hybrids of Columnea and Alloplectus computed from a matrix of 27 characters taken at random (cf.
Fig. 25).
THE JAMAICANSPECIESOFCOLUMNEA ANDALLOPLECTUS 209
reflect more strongly than others the genetic composition of taxa but it also indicates that such characters are not all of them consciously used by taxonomists.
A dendrogram provides a convenient expression of relationship by degree of similarity, since this similarity presumably arises from and expresses a similarity of genes, but it suffers from the disadvantages inevitable in a linear sequence. A more revealing view of relationships may be obtained by producing from the similarity matrix, by means of Gower’s method of Principal Co-ordinates Analysis (Gower,
65 70 75 80 85 90 70 | | ] | ] | |
ambiguus
pubescens
grisebachianus
argen tea
brevipila
hybrid 4
urbanii
fawcettii
harrisii
hirsuta
proctorii
subcordata
hybrid 6
hybrid 10
rutilans
jamaicensis
hispida
Fic. 23. Dendrogram showing the percentage of similarity of 17 species and hybrids of Columnea and Alloplectus computed from a matrix of 27 characters used in a key for their recognition (cf. Fig. 26).
210 THE JAMAICANSPECIESOFCOLUMNEA ANDALLOPLECTUS 6 | 4 od COoLUMNEA > 3 rutilans T e hybrid 6 hybrid 10 hirsuta ® 2 ——_ b . . fawcettii revipila a harrisii oe = @ —§subcordata | ° T e ® argentea hybrid 4 . hispida . ~4 3 “2, -| 4‘ 4 | i 2 3 L 7 T T J — T 7 e 7 urbanii -| -2 jamaicensis -3 T e e 4 ambiguus -4 _ e e . ubescens grisebachianus 5 P ALLOPLECTUS Fic. 24. Vector diagram showing relative similarity of 17 species and hybrids of Columnea
and Alloplectus computed from a matrix of 51 characters (cf. Fig. 21); the vertical co-ordinates are the vectors corresponding to the first latent roots, the horizontal co- ordinates those of the second latent roots, of the transformed matrix.
THE JAMAICANSPECIESOFCOLUMNEA ANDALLOPLECTUS
rutilans
hybrid 6
211
hirsuta Cotumnra 3 4 hybrid 10 ®@ e e fawcettii 2 «< brevipila e proctorii | 4 : hispida @ hybrid 4 e -4 -3 =? ai | 2 3 4 = 4 ¢ ss + my $ Sa + + harrisii @ ®@ subcordata e argentea @ | a urbanii x <2 j jamaicensis @ . grisebachianus e -3 4 4 = e Ms ambiguus pubescens 8 “5 ALLOPLECTUS 4a Fic. 25. Vector diagram showing relative similarity of 17 species and hybrids of Columnea
and Alloplectus computed from a matrix of 27 characters taken at random (cf. Fig. 22); co-ordinates as in Fig. 24.
BOT. 4, 5.
13
212 THE JAMAICAN SPECIES OF COLUMNEA ANDALLOPLECTUS
1966), a scatter diagram with the scaled first and second vectors of the transformed matrix as co-ordinate axes. In this vector diagram the distances between the units are proportional to the square root of their dissimilarity. Those resulting from the full and reduced matrices (Text-figs. 24-26) differ very much less than might be expected; indeed the major differences between the three are in the relation of the units to the axes and not to one another, as becomes evident by rotating the diagram of the reduced key matrix (Text-fig. 26) through about 45 degrees to the left and by
COLUMNEA
@ rutilans
4° hybrid 10
hirsuta 3 fawcettii e 3 e
hybrid 6 @
Z e.
brevipila
harrisii @@ @ hybrid 4 Pierrent e ® subcordata argentea
urbanii
jamaicensis e
a ambiguus
® grisebachianus
ALLOPLECTUS pubescens @
Fic. 26. Vector diagram showing relative similarity of 17 species and hybrids of Columnea and Alloplectus computed from a matrix of 27 characters used in a key for their recognition (cf. Fig. 23); the right to left co-ordinates are derived from the first latent roots, the left
to right co-ordinates from the second latent roots (cf. Fig. 24).
THE JAMAICANSPECIESOFCOLUMNEAANDALLOPLECTUS 213
transposing from left to right and vice versa the units of the diagram of the reduced random matrix (Text-fig. 25). They all convey much the same general information. In all three diagrams, Columnea lirsuta and C. fawcetti are brought closer to C. yutilans than is apparent in the dendrogram from the reduced matrices; C. rutilans and C. urbanw are well isolated and their hybrids spaced between them. All diagrams place fairly close together C. proctor, C. brevipila, C. subcordata, C. argentea and hybrid 4, an assemblage which should, however, on morphological grounds be separated into three groups. The configuration of species becomes here more meaningful if the third latent vector is brought into consideration by being treated as a third dimension. Thus, in the data derived from the full matrix, the third latent vector of — 0-25 for C. proctori brings this close to C. subcordata with a vector of — 0-24, while C. brevipila, with a vector of 0-25, and C. argentea, with a vector of 0-23, stand apart from them both and from hybrid 4, with a vector of 0-11; consideration of the third latent vector thus separates them into three groups agreeing with their general morphological characters.
The units on the vector diagrams resolve into a general pattern which appears significant. The typical species of Columnea with large zygomorphic flowers stretch together across the upper part of the diagram of the full matrix (Text-fig. 24) in a sequence passing from C. urbanii with green isophyllous leaves and entire calyx- segments to C. rutilans with markedly anisophyllous leaves red below and laciniate calyx-segments, and they undoubtedly form a natural group, the evolution of which has probably been partly controlled by the Jamaican endemic Streamertail Humming- _ bird (Text-fig. 5). From this group C. hispida stands somewhat apart, and in view of itsshort and more dingy corolla it probably has a different pollinating agent. Far beloy: them and well separated from each other in every diagram are the species of Allopiectus, i.e. A. ambiguus, A. grisebachianus and A. pubescens, probably self- pollinated, with C. jamaicensis brought much closer to them than to the typical species of Columnea, a placing which agrees well with its general morphology but is not evident from the dendrograms.
The vector diagrams give indeed the general impression that, whereas the typical Columnea species of Jamaica can well be regarded as derivatives of a single stock, C. jamaicensis represents an independent and florally less developed offshoot of Alloplectus.
A dendrogram simply indicates degree of similarity without necessarily placing the units of a particular group into a meaningful sequence within it, whereas a scatter diagram places them in two-dimensional relation to one another and hence can be used as a better basis for arranging the units in a sequence indicative of resemblance and possibly of genetic affinity. The dendrogram derived from the full matrix arranged in this way (Text-fig. 27) by reference to the corresponding scatter diagram (Text-fig. 24) gives an arrangement which coincides almost exactly with the taxo- nomic arrangement previously made by traditional methods.
This survey thus confirms the view that taxonometric methods when applied to an adequate body of data can produce a grouping of species certainly as satisfactory as one obtained by traditional methods and possibly more suggestive of their course of development; moreover such groupings can result from the computation of much
214 THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS
less than the postulated necessary 40 characters, particularly when the characters used are diverse enough to epitomise the whole organization of the species.
SYSTEMATIC ACCOUNT MATERIAL STUDIED
This account has been based on the collection in the Department of Botany, British Museum (Natural History), London (BM) together with material lent by the
65 70 75 80 85 90
95
] | | | |
rutilans hybrid 6 hybrid 10 hirsuta fawcettii brevipila argentea harrisii proctorii subcordata hybrid 4 urbanii hispida ambiguus grisebachianus pubescens
jamaicensis
Fic. 27. Dendrogram showing phenetic affinity of 17 species and hybrids of Columnea and Alloplectus computed from a matrix of 51 characters and arranged in a sequence based on a vector diagram from the first and second latent roots (cf. Figs. 21, 24).
THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS 215
Systematisch-Geobotanisches Institut, Universitat, Gottingen (GOET), the Science Museum, Institute of Jamaica, Kingston, Jamaica (IJ), the Royal Botanic Gardens, Kew (K), the Botanische Staatssammlung, Munich (M), the New York Botanical Garden (NY), the Department of Botany, University of the West Indies, Mona, Jamaica (UCWI) and the United States National Herbarium, Smithsonian Institu- tion, Washington, D.C. (US).
Specimens are listed by the parishes of Jamaica, beginning with Hanover and Westmoreland in the west and ending with Portland and St. Thomas in the east (cf. Stearn, 1959 : 134, fig. 1a).
As an indication of how our knowledge of the Jamaican species of Columnea and Alloplectus has grown, it is of interest to note that Grisebach (in 1862) distinguished seven species on 15 gatherings, Urban (in Ig0I) seven species on 20 gatherings, Morton (in 1944) ten species on 54 gatherings; the present account distinguishing 13 species has been based on 190 gatherings. Our understanding of the group in Jamaica would appear more likely now to benefit from field observations on pollina- tion and dissemination than from further collecting.
ACKNOWLEDGEMENTS
Grateful thanks are here expressed to those in charge of the above herbaria for the opportunity to study their material. I am indebted especially to Mr. George R. _ Proctor for his help to me while collecting Columneas and other plants in Jamaica. Dr. Brian D. Morley has generously made available the results of his extensive cytological and other research and contributed the sections on cytology and biochemistry above. He and Miss Barbara Heywood also assisted me in ascertaining and tabulating the characters used in the taxonometric analysis above, which is the result of computation by Mr. Gavin J. S. Ross at the Department of Statistics, Rothamsted Experimental Station; it has benefited much from the advice of Dr. J. G Sheals. Mr. D. Erasmus drew the text-figures. The co-operation of them all is much appreciated.
KEY TO JAMAICAN SPECIES OF COLUMNEA AND ALLOPLECTUS
Corolla tubular or narrowly funnel-shaped, the limb not markedly 2-lipped, the lobes of the limb only 2-8 mm. long; leaf-bearing branches slender (1-3 mm. diam.) ; all leaf-blades small (mostly under 5 cm. long, 3 cm. broad):
Calyx-segments laciniate with four or five prominent teeth either side; central Jamaica ; 2. Alloplectus grisebachianus Calyx-segments entire or Gath: one or fine cmtuve teeth either side: Leaves and calyx densely hirsute; corolla-lobes almost uniform, about 1-2 mm. long; eastern Jamaica . , ; . 1. A. pubescens Leaves and calyx apparently glabrous but with ciliate margins and short inconspicuous hairs on surface; corolla-lobes forming two lips, the lower lip about 6 mm. long; western Jamaica : . I, Columnea jamaicensts
216 THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS
Corolla markedly 2-lipped, the lobes of the limb rcm. or more long; leaf-bearing branches stout (5-10 mm. diam.) ; leaf-blade of at least one leaf of a pair larger (longer than 5 cm. or broader than 3 cm.):
Leaf-blades partly or completely red beneath or coloured red along the veins: Calyx-segments deeply toothed along the margin, the lowermost teeth to 5 mm. long; larger leaf-blades three to four times as long as broad 11. C. rutilans Calyx-segments entire or shortly toothed, the teeth to 2 mm. long; larger leaf- blades about two times as long as broad . C. rutilans * urbani
Leaf-blades green or silvery beneath, not red-veined or red: Hairs on upper side of leaf minute, 0-2—-0-3 mm. long, 2-celled, inconspicuous: Calyx-segments entire, green, inconspicuously hairy with short rather appressed hairs:
Pedicels with minute appressed hairs 0-4 mm. or less long; Westmoreland, St. Elizabeth ; ; . 8.C. brevipila
Pedicels with longer somewhat spreading hairs to 1 mm. long; Manchester
C. rutilans x urbani Calyx- segments with one to three linear teeth to 2 mm. long on each margin, red in upper part, densely hirsute ; ; - 9. C. fawcettu Hairs on upper side of leaf 1 mm. or more long, 3-8- celled, conspicuous: Calyx sericeous with dense appressed silvery hairs:
Leaf-blades densely sericeous beneath, three to five times as long as broad, with four or five primary veins either side, almost symmetric at base; corolla yellow . 6. C. argentea
Leaf-blades with short curved hairs not completely covering the surface beneath, about two to three times as long as broad, with six to eight primary veins either side, markedly asymmetric at base; corolla red and yellow : . 6.C. harnsi
Calyx not sericeous but hispid or 1 pilose with long, spreading, often glandular, hairs:
Calyx-segments with one to three linear teeth on each margin and with attentuate red tips; eastern Jamaica : : : g. C. hirsuta
Calyx-segments entire or only minutely toothed, green, the tips not attenuate; central and western Jamaica:
Leaves and calyx with conspicuous red hairs; corolla about 2-5 cm. long 2. C. hispida
Leaves and calyx with colourless (not red) hairs; corolla 4—5 cm. long: Four calyx-segments joined at base for 5 to 8 mm., the fifth (posterior) one free from them; leaf-blades sometimes subcordate at base, usually asymmetric . . 5. C. subcordata All calyx-segments free almost to the base; leaf-blades rounded to
cuneate at base:
Leaves of a pair markedly unequal in size, the largest asymmetric at base and usually 8-14 cm. long, hirsute below with abundant long spreading hairs; corolla striped red and yellow throughout
4. C. proctorit
THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS 217
Leaves of a pair almost equal in size, the largest symmetric at base and at most 8cm. long, usually much less, strigillose below with very short appressed hairs; corolla yellow or with red markings at base only:
Corolla yellow; style yellow. : ; . 3. C. urban
Corolla yellow with red stripes on base, or, if entirely yellow, style
red : : : : . C. rutilans x urbani COLUMNEA L.
CoLuMNEA L., Sp. Pl. 2 : 638 (1753); Gen. Pl. ed. 5 : 286 (1754).
Shrubs. Leaves opposite, those of a pair equal to markedly unequal, usually shortly petiolate. Flowers solitary or fasciculate in leaf-axils, with or without bracts. Calyx 5-partite. Corolla inferior, erect in the calyx, gibbous but not spurred at base, the tube gradually expanded upwards, long, the limb 2-lipped, the upper lip 3-lobed with the middle lobe formed from two united lobes into a notched galea or hood and with two lateral lobes spreading from it, the lower lip entire and recurving or reflexed. Stamens 4, didynamous, the filaments united in the lower part, free and filamentous above, the anthers of all four stamens cohering at anthesis, their loculi parallel, longitudinally dehiscent; fifth stamen represented by a staminode. Disc gland large and dorsal, sometimes accompanied by two to four smaller glands. Ovary superior; stigma usually 2-lobed. Fruit a fleshy berry; seeds minute, longi- tudinally or obliquely striate.
Type-species by monotypy: C. scandens L.
The above description does not cover the anomalous non- Jamaican species with
regular corolla mentioned above (p. 187).
I. Columnea jamaicensis Urban, Symb. Antill. 2 : 359 (z901).—Morton in Contr. U.S. Nat. Herb. 29 : 3 (1944).—Stearn in Proc. Linn. Soc. Lond. 170 : 141 (1959). (Pl. 14; Text-figs. 2E, 6, 28H.)
Pterygoloma repens sensu Griseb., Fl. Brit. W. Ind. Is. : 464 (1862) pro parte, quoad pl. Jamaic.; non Hanstein.
Shrub of dwarf creeping habit, the stems grey, slender, c. 2 mm. thick, pilose with reddish hairs when young, later glabrous. Leaves paired, those of a pair unequal, sparsely setose on both surfaces, dark green above, paler green below, the larger leaf of the pair long-stalked, the blade narrowly ovate, asymmetric, with the apex shortly acuminate or acute, the margin almost entire or remotely serrulate, the base cuneate, 2°5-5°5 cm. long, I-23 cm. broad, the smaller leaf soon falling, short- or long-stalked, the blade 0-5-3 cm. long, 0:5-1-5 cm. broad; petiole of larger leaves 5-11 mm. long, of smaller leaves 1-3 mm. long. Flowers solitary in leaf-axils. Pedicel 1-5-3 cm. long, strigillose with very short appressed hairs. Calyx divided to the base, reddish or green, sparsely and shortly pilose and ciliate outside; segments narrowly ovate or lanceolate, acute, 8-5—12 mm. long in flower, to 16 mm. long in fruit, entire or with one or two minute teeth on each margin. Corolla tubular, gradually widened, 2-5-
218 THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS
3 cm. long, crimson or orange yellow streaked with red bands, the tube much longer than the calyx, the upper lip about 0-5-1 cm. long, 3-lobed, the lower lip 5-8 mm. long, narrowly oblong. Stamens slightly exserted; filaments glabrous; anthers about 1mm. long. Ovary glabrous except for a few sparse hairs near the apex; style glandular-pilose. Berry white, c. 5 mm. across.
WESTMORELAND: Springfield, 1849, Wullschlégel 1290 (M; GOET, type of C.
yamaicensis). Bluefields Mountain, four miles from the sea, 600 m., Purdie (K)
—
Sa SWANS;
S S
: = SSS
3
Fic. 28. Columnea fawcettii (Urban) Morton: A, flowering shoot; B, calyx-segment; C, stamens; D, anthers; E, anther showing insertion of filament; F, gynoecium, with Columnea urbanii Stearn; G, flower (Stearn 361, spirit material).
Columnea jamaicensis Urban: H, flower (Stearn 928, spirit material).
disc-gland (Stearn 255).
THEJAMAICANSPECIESOFCOLUMNEA ANDALLOPLECTUS 219
Three miles west of New Works, Bluefields Hill, 680 m., July 1954, Farr 5643 (IJ). Two and a half miles northeast of Bluefields P.O., 700m., July 1954, Webster & Wilson 1954 (AA). Kentucky Hill, Bluefields Mountain, 600-760 m., March 1908, Harris 10200 (BM; NY). Woodstock, near Newmarket, 420 m., Sept. 1907, Britton 1571 (NY; US), Harris 9836 (NY). Same locality, Oct. 1960, Proctor 21518 (BM). Near Woodstock, 360 m., Dec. 1965, Adams 12794 (BM). One mile west-north- west of Hopewell, 530 m., Nov. 1955, Proctor 11232 (BM; IJ).
TRELAWNY: Cockpit Country, at Black Banana Hole, five miles north of Accom- pong, 330 m., April, 1956, Stearn 928 (BM). Red Mud Hole, six miles north-north- west of Accompong, 330 m., April 1956, Stearn 964 (BM).
Urban based his C. jamaicensis on specimens collected by Wullschlagel at “ Springfield’, presumably Springfield in Westmoreland parish, although there are at least 20 other places in Jamaica named Springfield. Grisebach had earlier (Fl. Brit. West. Ind. Is. : 464 (1892)) listed this as Pterygoloma repens (Hook.) Hanstein, which is figured in Botanical Magazine 72 : t. 4250 (1846) as Alloplectus repens from Colombia. Among Jamaican species of Coluwmnea it is notable for its very slender creeping stems, small leaves and almost tubular narrow corolla with a short only slightly bilabiate limb. The type-specimen (Wullschlégel 1290, GOET) has red narrowly ovate sepals about 12 mm. long, 4-6 mm. broad at anthesis, and similar specimens have been collected in Westmoreland. Specimens collected by me in the Cockpit Country diverge in having green lanceolate sepals 8-5—10 mm. long, 3-5-4 mm. broad at anthesis. It grows on damp walls, moss-covered rocks and trees, in western Jamaica (Text-fig. 6). :
Hooker’s Alloplectus repens, i.e. C. repens (Hook.) Hanstein (in Linnaea 34 : 395 (1865)) was based on a cultivated plant said to have been introduced by Purdie from Colombia. It comes so close to C. jamaicensis, however, as to arouse suspicion that Hooker may have had an unusual variant of C. jamaicensis mislabelled as to origin. If no such plant as Hooker’s occurs in Colombia, then the correct name for the Jamaican species will be C. repens. Some of Purdie’s Colombian specimens have been attributed to Jamaica by Grisebach; hence the possibility of some of his Jamaican material being attributed to Colombia.
2. Columnea hispida Swartz, Nov. Gen. & Sp. Pl. : 94 (1788); Fl. Ind. Occ. 2: 1085 (1800).—Griseb., Fl. Brit. W. Ind. Is. : 465 (1862) excl. descript.—Urban, Symb. Antill. 2 : 364 (1g0r).—Stearn in Proc. Linn. Soc. Lond. 170 : 141 (1959). (Pl. 15; Text-figs. x A, 2 A, 7.)
Shrub, the stems brownish grey, stout, hispid with stiff spreading hairs to 3 mm. long. Leaves paired, those of a pair markedly unequal, dark green above, paler green below, hirsute on the upper surface and at the margin with long erect reddish hairs to 4mm. long, hirsute on the lower surface along the veins with similar long hairs and hispid on the surface itself with short colourless hairs to 0-5 mm. long, the larger leaf of the pair relatively long-stalked, asymmetric, the blade more or less elliptic or obovate, with the apex obtuse, the margin remotely and shallowly serrate or almost entire, the base asymmetric, its lower side broadly cuneate, its other side narrowly
220 THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS
cuneate and arising from the midrib about 5-10 mm. higher, 6-15 cm. long, 4—7 cm. broad, with four to six curved veins impressed each side of the midrib above, the smaller leaf 2-6 cm. long, 1-5—3-5 cm. broad; petiole of larger leaf 1-2 cm. long, of smaller leaf 5-10 mm. long. Flowers up to three in a leaf-axil. Pedicels c. 5 mm. long, densely hispid. Calyx divided to the base, very pale green but hispid outside with long reddish hairs; segments very narrowly oblong, c. 15 mm. long in flower to 2cm. long in fruit, 3 mm. broad, entire. Corolla strongly 2-lipped, dull golden yellow, c. 2-5 cm. long, densely hispid on the back with hairs to 3 mm. long, the tube a little shorter than the calyx, c. r cm. long, the upper lip c. 1-5 cm. long and trilobed with a notched middle lobe and two rounded lateral lobes c. 4 mm. long, the lower lip c. cm. long, 4 mm. broad, recurved. Stamens exserted; filaments minutely and sparsely hairy; anthers c. 2mm. long. Ovary covered with long ascending bristles; style glandular-pubescent.
ST. JAMES: White Rock Hill, one mile south of Sweetwater, 600 m., March 1956, Stearn 449 (BM). Same locality, Feb. 1957, Proctor 16158 (BM; IJ). Half mile south of Point P.O.,500—550 m., 1961, Proctor 22577 (IJ). Near Catadupa (fide Morley).
UNLOCALIZED: Swartz (BM, type-collection).
This species was described by Swartz from fruiting specimens collected by himself “in montibus altis Jamaicae occidentalis, ad rupes umbrosas ”’ and it remained thus vaguely localized and not again collected, despite the wide-ranging activities of Purdie, Harris and Britton in Jamaica, until found by Proctor and myself in 1956, growing on shaded limestone cliffs in the west of the island (Text-fig. 7), as indicated by Swartz. The flowers were unknown until 1957. It has since been cultivated at Kew and Montreal botanic gardens. From other Jamaican species it is easily distinguished by the long reddish hairs on the upper surface of the leaves and by the small flowers.
3. Columnea urbanii Stearn, sp. nov. (Pl. 16; Text-figs. 1B, 2H, 9, 28G.)
Columnea hispida sensu Griseb., Fl. Brit. W. Ind. Is. : 465 (1862) pro parte maj.; non Swartz.
Columnea hirsuta sensu Hanstein in Linnaea 34 : 413 (1865) pro parte quoad Wullschlagel 959; non Swartz.
Columnea hirsuta var. subintegra {. wullschlaegelit Urban, Symb. Antill. 2 : 363 (1901).
Suffrutex, caulibus ascendentibus 4-6 mm. crassis, primo pilis patentibus ad 2 mm. longis hirsutis, demum griseis glabratis, internodiis 0-5-4 cm. longis, cicatricibus prominentibus. Folia ad apicem ramulorum conferta, per paria aequalia, subsessilia, plerumque late elliptica sed interdum rotunda vel late obovata, apice obtusa vel rotundata, margine subintegra, basi late cuneata raro asymmetrica, 3-8 cm. longa, 2°5-5'5 cm, lata, in vivo utrinque pallide viridia, nervis lateralibus utroque latere costae 4-5 arcuatis, supra pilis erectis incoloribus plerumque 5-cellularibus c. 2 mm. longis hirsuta, subter pilis plerumque multo brevioribus appressis c. 0-5 mm. longis strigulosa; petiolus c. 1-4 mm. longus. Flores axillares 1-2. Pedicelli c. 8-15 mm. longi, hirsuti. Calyx viridis, extus hirsutus; segmenta libera subaequalia lanceolata acuta, anthesi 10-15 mm. longa, post anthesin ad 25 mm. longa, 3-5-5 mm. lata, integra. Corolla valde bilabiata, c. 5 cm. longa, flava, dorso pilis patentibus ad
THE JAMAICAN SPECIESOFCOLUMNEAANDALLOPLECTUS 221
2:5 mm. longis hirsuta; tubus basi postice saccatus gradatim ampliatus c. 22 mm. longus calyce fere duplo longior; labium superum c. 18-25 mm. longum, trilobatum, lobo medio emarginato c. 10mm. lato, lobis lateralibus anguste triangularibus patentibus; labium inferum c. 18 mm. longum, 3 mm. latum, recurvatum. Stamina exserta, filamentis glabris, antheris oblongis c. 1-5-2 mm. longis. Ovariwm apice hirsutum; stylus pubescens.
Shrub, the stems ascending, grey, stout, 4-6 mm. thick, hirsute, with spreading hairs to 2mm. long when young, later glabrous. Leaves paired, those of a pair almost equal, short-stalked or almost sessile, the blade mostly broadly elliptic but varying from rotund to broadly obovate, with the apex obtuse or rounded, the margin almost entire, the base broadly cuneate, sometimes asymmetric, 4-12 cm. long, 2:5-5°5 cm. broad, light green on both surfaces, with 4-5 curved veins each side of the midrib impressed deeply above, hirsute above with erect mostly 5-celled hairs c. 2mm. long, strigose below with numerous short hairs c. 0:5 mm. long, together with some longer hairs; petiole 1-4 mm. long. Flowers I or 2 in a leaf-axil. Pedicels c. 8-15 mm. long, hirsute. Calyx divided to the base, green, densely hirsute outside, less so inside; segments lanceolate, acute, 10-15 mm. long in flower, 3-5 mm. broad, to 25 mm. long in fruit, entire. Corolla strongly 2-lipped, pale yellow, c. 5 cm. long, hirsute on the back with long spreading hairs to 2-5 mm. long, the tube nearly twice as long as the calyx, c. 22 mm. long, the upper lip c. 18-25 mm. long, trilobed, with a notched middle lobe c. 10mm. broad and two spreading narrowly triangular acute lateral lobes, the lower lip c. 18 mm. long, 3 mm. broad,
recurved. Stamens exserted; filaments glabrous; anthers c. 1-5-2 mm. long. Ovary
hirsute at the apex; style pubescent.
MANCHESTER: Top Hill, 820 m., March 1966, Morley 325 (BM). Fairfield, 1840, Wullschldgel 959 (GOET; M; type-collection of C. hirsuta var. subintegra f. wull- schlaegelit). Along road between Fairfield and Huntley, 800-900 m., Jan. 1957, Proctor 16109 (IJ). Huntley, 900 m., 25 Feb. 1956, Stearn 361 (BM, holotype), Proctor 11621 (IJ). Near Christiana, May 1901, Fawcett 8145 (UCWI). Upper
slopes and summit of Heron’s Hill (Shooter’s Hill, Martin’s Hill), goo—940 m.,
March 1958, Proctor 17489 (BM; IJ). Banana Ground, 900 m., Dec. 1961, Adams tor27 (UCWI). Oatlands, north-west of Rose Hill, 830m., Jan. 1961, Proctor 21915 (BM).
St. ANN: Mason River district, three to four miles northwest of Kellits, 760 m., March 1957, Proctor 16201 (BM; IJ).
CLARENDON: Peckham Woods, 760 m., March 1954, Proctor 8422 (IJ). Knox College, Spaldings, 850 m., March 1952, Proctor 6345 (IJ).
This species was first collected by the Moravian missionary Wullschlagel in 1849 and identified by Grisebach with the very different C. hispida. Urban recognized
its distinctness but, having only Wullschlagel’s limited material at hand, described it
briefly as a form of C. hirsuta, which is confined to the Blue Mountains of Jamaica. In a living state the leaves are a bright green with the veins deeply impressed above, and the flowers are pale yellow. It grows among both rocks and trees, and sometimes on stone walls, in central Jamaica (Text-fig. 9). Fairfield, where Wullschlagel collected his specimens, and Huntley, where I collected mine, are about five miles
222 THE JAMAICAN SPECIES OFCOLUMNEA AND ALLOPLECTUS
apart and near the western boundary of Manchester. It is named in honour of Ignaz Urban (1848-1931), the most scholarly, industrious and scientifically prolific of the many botanists who have studied West Indian plants; a short survey of his career by R. A. Howard will be found in E. Carroll and S. Sutton, Cumulative Index to the nine Volumes of the Symbolae Antillanae : 1-5 (1965). Among hybrids between this species and C. rutilans are some plants clearly intermediate, others approximating more closely to one or other parent species, probably as a result of back-crossing. Those nearest to C. urbanii in leaf-size and pubescence may nevertheless usually be distinguished from it by having the base of the leaf-blade more attenuate and the corolla partly red, although sometimes they agree with it in almost all characters but have only minute bristles on the upper surface of the leaf.
4. Columnea proctorii Stearn, sp. nov. (Pl. 17, 18; Text-figs. 2B, 8.)
Suffrutex, caulibus ascendentibus c. 5 mm. crassis, pilis patentibus ad 3 mm. longis hispidis, internodiis longis, cicatricibus prominentibus. Folia ad apicem 5-14cm. longum ramulorum disposita, per paria valde inaequalia, folio majore breviter petiolato asymmetrico, lamina ovata ad obovata, apice acuta vel breviter acuminata, margine subintegra, basi uno latere rotundato, altero cuneato, 5—I0 cm. longa, 2:5-6-5 cm. lata, pallide viridia, nervis lateralibus utroque latere costae 4-6 arcuatis supra profunde impressis, utrinque pilis erectis incoloribus plerumque 5-cellularibus 1-5—-2:5 mm. longis hirsuta, folio minore fere sessili orbiculato I-3°5 cm. longo latoque, similiter vestito; petiolus folii majoris 2-8mm. longus. Flores axillares 1-3. Pedicelli 12-20 mm. longi, dense hirsuti. Calyx viridis, extus pilis longis glandulosis hirsutus; segmenta libera subaequalia anguste lanceolata acuta, anthesi c. 2 cm. longa, post anthesin c. 3 cm. longa, 2-3 mm. lata, integra. Corolla valde bilabiata, c. 4-5 cm. longa, flava rubrovittata, dorso pilis patentibus glandulosis ad 2mm. longis dense hirsuta; tubus basi postice saccatus gradatim ampliatus c. 18-25 mm. longus calycem aequans; labium superum c. 18 mm. longum, trilo- batum, lobo medio emarginato c. 7 mm. lato, lobis lateralibus anguste triangularibus vel fere lanceolatis obtusis patentibus; labium inferum c. 12 mm. longum, 4mm. latum, recurvatum. Stamina exserta, filamentis glabris, antheris oblongis 2 mm. longis. Ovarium basi glabrum, supra medium pubescens; stylus glanduloso- pubescens.
Shrub, the stems grey, stout, c. 5 mm. thick, hispid with spreading hairs to 3 mm. long. Leaves occupying the terminal 5-14 cm. of the shoots, paired, those of a pair markedly unequal, light green, hirsute on both surfaces with erect colourless hairs I‘5-2'5 mm. long, the larger leaf of the pair short-stalked, asymmetric, the blade more or less ovate to obovate, with the apex acute or shortly acuminate, the margin almost entire, the base asymmetric, its lowest side rounded, the other cuneate and arising from the midrib about 5 mm. higher, 5—10 cm. long, 2:5-6:5 cm. broad, with four to six curved veins each side of the midrib impressed deeply above, the smaller leaf almost sessile and orbicular, 1-3-5 cm. long and broad; petiole of larger leaf 2-8 mm.long. Flowers 1-3 in a leaf-axil. Pedicels 12-20 mm. long, hirsute. Calyx divided to the base, light green, hirsute with long glandular hairs, the segments
!
THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS 223
narrowly lanceolate, acute, c. 2cm. long, 2-3 mm. broad in flower to 3 cm. long in fruit, entire. Corolla strongly 2-lipped, pale yellow with longitudinal red stripes, c. 4-5 cm. long, densely hirsute on the back with long spreading glandular hairs to 2mm. long, the tube almost as long as the calyx, c. 18-25 mm. long, the upper lip c. 18 mm. long, trilobed, with a notched middle lobe about 7 mm. broad and two spreading narrowly triangular to almost lanceolate obtuse lateral lobes, the lower lip c. 12mm. long, 4mm. broad, recurved. Stamens exserted; filaments glabrous; anthers 2mm. long. Ovary glabrous in the lower part, pubescent in the upper part style glandular-pubescent.
HANOVER: Ramble, March 1905, Fawcett 8916 (UCWI).
St. JAMES: Chatsworth district one and a half miles north-north-west of Maroon Town, 480 m., March 1956, Stearn 451 (BM, holotype; K), Proctor 117753 (IJ). White Rock Hill, one mile south of Sweet Water, 600-670 m., Feb. 1957, Proctor 16160 (IJ). Same locality, Dec. 1962, Proctor 23009 (BM; IJ).
ST. ELIZABETH: Maggotty Falls, by the Black River, 120 m., May 1956, Stearn 1039 (BM), Proctor 15382 (IJ).
TRELAWNY: Cockpit Country at Black Banana Hole, about five miles north of Accompong, 330 m., April 1956, Stearn 938 (BM). Cockpit Country, near Red Mud Hole, about six miles north-north-west of Accompong, 330 m., April 1956, Stearn 957 (BM; K). Cockpit Country, at Ramgoat Cave, 460 m., Jan. 1956, Howard, Proctor & Stearn 14645 (BM; IJ). Miss Laura’s Hill, Wilson Valley district, one mile north of Warsop, 600-670 m., Jan. 1964, Proctor 24452 (BM; IJ). Island View Hill,
Wilson Valley district, one and a half miles north of Warsop, 600-670 m., March
1960, Proctor 20756 (BM; IJ). Five miles northwest of Troy, 530 m., Sept. 1963, Adams 12420 (BM).
MANCHESTER: Half a mile northwest of Christiana, 900 m., Oct. 1958, Proctor 18288 (BM; IJ). Same locality, Feb. 1960, Proctor 20595 (BM; IJ). One mile southeast of Pike, g00 m., Oct. 1958, Proctor 18308 (BM; IJ).
This species is widespread in the Cockpit Country and adjacent areas of western
_ Jamaica (Text-fig. 8) and is apparently more northern in distribution than C. urbanit,
from which it differs in having the leaves of a pair markedly unequal in size and hirsute below, and the corolla usually striped longitudinally with red; the five free calyx-segments distinguish C. proctorii from C. subcordata. I have named it in honour of George Richardson Proctor, author of A Preliminary Checklist of Jamaican Pteridophyta (1953), Additions to the Flora of Jamaica (1967) and other publications on Jamaican plants, whose collecting activities, extended since 1949 over almost the whole of Jamaica, have so notably increased knowledge of the distribution of species within the island and in whose company in 1956 I had the privilege of collecting the type material of this species.
5. Columnea subcordata Morton in Contr. U.S. Nat. Herb. 29 : 6 (1944).— Stearn in Proc. Linn. Soc. Lond. 170 : 141 (1959). (Pl. 19; Text-figs. 2G, Io.)
Shrub, the stems grey or brownish, stout, c. 5 mm. thick, hispid when young with long mostly spreading brownish multicellular hairs c. 2-3 mm. long. Leaves paired, those of a pair unequal, all short-stalked, green and hirsute with sparse long whitish
224 THE JAMAICANSPECIESOFCOLUMNEA ANDALLOPLECTUS
multicellular hairs 0-8—-1-8 m ~. long on both surfaces, the blade of the larger leaf of the pair more or less oblong-elliptic, with the apex abruptly and shortly acuminate, the margin entire and long-ciliate, the base subcordate and slightly oblique, 8-14 cm. long, 4:5-6:5 cm. broad, the blade of the smaller leaf (which usually falls quickly) broadly ovate or broadly obovate, 2:5-3:5 cm. long, 2 cm. broad; petiole 3-10 mm. long. Flowers 1-4 in a leaf-axil. Pedicels 1-5-2 cm. long, hirsute. Calyx 2-lipped, the four anterior segments joined from the base for 5 mm. (in flower) to 8 mm. but the posterior segment free to the base, light green, hispid with long whitish hairs on both surfaces, 1-5-2:2 cm. long, the four free segments narrowly oblong-lanceolate 2°5-4°5 mm. broad, acute, the posterior segment linear-tapering, all entire. Corolla strongly 2-lipped, light yellow, glandular-pilose, 4-4-8 cm. long, the tube mostly covered by the calyx, the upper lip c. 2:5 cm. long with a deeply notched middle lobe and two spreading oblong lateral lobes, the lower lip c. 1-5 cm. long, narrowly oblong, reflexed. Stamens exserted; filaments pubescent; anthers 2mm. long. Ovary sericeous; style glandular-pilose.
TRELAWNY: Mount Ridgway Road, Windsor, roo—150 m., April 1931, G. S. Miller 1556 (US, type). Windsor House estate, 120 m., March 1956, Stearn 472 (BM; K).
Columnea subcordata stands apart from all other Jamaican species of Columnea in the fusion of the four anterior calyx-segments into one for the lower third of their length, with the posterior one remaining completely free. It probably arose as a mutation from C. proctoriz. At the type-locality, Proctor and I found it growing high on trees, the sparingly branched stems hanging to a length of six feet. The leaves are dark green above, the hairs colourless, the corolla light yellow (Chinese yellow). It is evidently confined to a small area of western Jamaica (Text-fig. 10).
6. Columnea harrisii (Urban) Britton ex Morton in Contr. U.S. Nat. Herb. 29 : Lo (1944).—Stearn in Proc. Linn. Soc. Lond. 170 : 141 (1959). (Text-fig. 1E, II.)
Columnea hirsuta var. subintegra f. harrisii Urban, Symb. Antill. 2 : 363 (1901).
Shrub, the stems grey, fairly slender, pilose with whitish hairs. Leaves paired, those of a pair unequal or almost equal, green and sericeous with numerous slightly arched more or less appressed multicellular hairs on both surfaces, the larger leaf of an unequal pair asymmetric, the blade more or less narrowly elliptic-oblong, with the apex shortly acuminate, the margin remotely and shallowly toothed, the base rounded on the outer side, acute on the inner side, 7-5-13 cm. long, 3-5 cm. broad, the smaller leaf broadly or narrowly ovate or almost orbicular, 1-5—4:5 cm. long, I:0-2:3 cm. broad; petiole of larger leaves 3-8 mm. long, of smaller leaves I-3 mm. long. Flowers 1 to 3 ina leaf axil. Pedicels 5-25 mm. long, hirsute. Calyx divided to the base, green, densely sericeous on both surfaces; segments lanceolate to narrowly lanceolate, 1-8-2-5 cm. long, c. 4 mm. broad, entire or with one or two minute teeth on each margin. Corolla strongly 2-lipped, yellow striped with red, 3-8-5 cm. long, sericeous outside with long whitish hairs, the tube almost covered by the calyx, the upper lip c. 2-3 cm. long with a notched middle lobe and two spreading narrowly- oblong lateral lobes, the lower lip c. 12-15 mm. long, narrowly oblong, recurved. Stamens exserted; filaments glabrous; anthers c. r-8mm. long. Ovary sericeous; style glandular-pilose.
THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS 225
HANOVER: Belvedere, 160m., Jan. 1899, Harris 7522 (BM, isotype). Same locality, March 1903, Fawcett 8480 (BM; NY). Hills near Kempshot, March 1908, Britton 2438 (NY; US). Shettlewood, 350-450 m., Feb. 1961, Proctor & Mullings 22016 (BM; IJ).
WESTMORELAND: Newmarket, Sept. 1907, Britton 1576 (NY). Woodstock, 400 m., Aug. 1907, Harris (UCWI). Same locality, 360-420 m., Oct. 1960, Proctor 21519 (BM; IJ).
St. JAMES: One mile southwest of Montpelier, 110 m., Feb. 1961, Proctor & Mullings 22010 (BM; IJ).
Columnea harrisi1, although placed by Urban under C. /irsuta, is abundantly distinct in its sericeous pubescence, entire calyx-segments and glabrous stamens, its distribution, and its ecology, as pointed out by Morton. It is a lowland species confined to the west of the island (Text-fig. 11), where it is epiphytic on large trees and often associated with bromeliads. Its name commemorates William Harris (1860-1920), for many years Superintendent of Public Gardens in Jamaica, whose extensive collections from 1890 onwards led to the discovery in Jamaica of approaching 400 species above those recorded for the island in Grisebach’s Flora
(1859-64).
7. Columnea argentea Griseb., Fl. Brit. W. Ind. Is. : 465 (1862).—Urban, Symb. Antill. 2 : 361 (1g01).—Morton in Contr. U.S. Nat. Herb. 29 : 9 (1944).— Stearn in Proc. Linn. Soc. Lond. 170 : 141 (1959). (Pl. 20; Text-figs. 1F, 2C,
Bika.)
C. wullschlaegeliana Hanstein in Linnaea 34 : 414 (1865).
Shrub, the stems stout, c. 5 mm. thick, silky at the apex with white hairs. Leaves paired, those of a pair almost equal, the blade very narrowly elliptic, narrowly elliptic or oblanceolate, with the apex shortly acuminate, the margin remotely and shallowly toothed, almost entire, the base rounded or cuneate, 7-5—17 cm. long, 25-4 cm. broad, grey-sericeous on both surfaces; petiole 4-10 mm. long. Flowers I to 4 ina leaf axil. Pedicels 5-25 mm. long, sericeous. Calyx divided to the base, densely sericeous on both surfaces; segments equal, oblong-linear, c. 14-18 mm. in flower to 20-24 mm. long in fruit, 2-4 mm. broad, entire. Corolla strongly 2-lipped, pale yellow, 4-5 cm. long, densely white-sericeous outside, the tube slightly longer than the calyx, c. 2-0-2-5 cm. long, the upper lip c. 2-0-2-5 cm long, with a notched middle lobe and two narrowly triangular lateral lobes, the lower lip 15-18 mm. long, oblong-linear, reflexed. Stamens exserted; filaments glabrous; anthers 1-5 mm, long. Ovary sericeous; style glandular-pilose. Berry ellipsoid, pink, c. 11 mm. long, 8 mm. broad.
MANCHESTER: Nazareth, 1849, Wullschldgel 959 (GOET, holotype of C. argentea; M, holotype of C. wullschlaegeliana). Nazareth, among rocks behind the Moravian Church, one mile south of Maidstone P.O., 880 m., Feb. 1956, Stearn 359 (BM; K:; topotype of C. argentea and C. wullschlaegeliana), Proctor 11615 (IJ). Alumina Jamaica property near Kirkvine Works, Shooters Hill, 380-460 m., Jan. 1958, Howard & Proctor 14944 (BM; IJ). Bath district, along Somerset Road southeast of
226 THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS
Johns Hall, 700 m., Jan. 1961, Proctor 21883 (BM; IJ). New Green, Sept. 1908, Britton 3755 (NY). Clones, Ballynure to Malton, 700 m., April 1962, Adams 11018 (BM).
Columnea argentea is an attractive and easily recognized species, notable for the silvery and silky pubescence which clothes the relatively long and narrow leaves, the calyx and the outside of the pale clear yellow corolla. At the type-locality I found it growing among rocks. Howard & Proctor record it as epiphytic on clumps of Hohenbergia in trees. All collections have come from within ten miles of one another in central Manchester (Text-fig. 12).
8. Columnea brevipila Urban, Symb. Antill. 6 : 41 (1909).—Morton in Contr. U.S. Nat. Herb. 29 : 9 (1944).—Stearn in Proc. Linn. Soc. Lond. 170 : 141 (1959). (Text-fig. 1D, 13.)
Shrub, the stems grey, stout, 5-8 mm. thick, strigillose when young with minute white appressed hairs and with lines of more villous hairs between the leaf-insertions, later glabrous. Leaves paired, those of a pair almost equal, shortly stalked, the blade narrowly oblong-elliptic or narrowly elliptic, with the apex shortly acuminate, the margin remotely and obscurely toothed, the base cuneate, 5-11 cm. long, 2—5 cm. broad, green and strigillose on both surfaces with minute whitish appressed hairs c. 0-2 mm. long; petiole 7-11 mm. long. Flowers I to 5 in a leaf-axil. Pedicels 1-2 cm. long, strigillose. Calyx divided to the base, green, minutely strigillose on both surfaces; segments narrowly lanceolate, acute, 2-3 cm. long, 3-5 mm. broad, entire. Corolla strongly 2-lipped, completely yellow, 4-5-5 cm. long, sericeous out- side with long whitish hairs, the tube mostly covered by the calyx, the upper lip c. 2-2-5 cm. long, with a strongly notched middle lobe and two spreading and ascending oblong lateral lobes, the lower lip to 1-5 cm. long, narrowly oblong, reflexed. Stamens exserted; filaments glabrous; anthers 2mm. long. Ovary densely hirsute; style glandular-pilose.
WESTMORELAND: Kentucky Hill, Bluefields Mountains, 600-760 m., March 1908, Harris torg9 (NY; UCWI; US; type-collection). Hills west of Bognie, Bluefields Mountains, 700-760 m., Feb. 1961, Proctor & Mullings 22043 (BM). Teague Gully, 300-400 m., Feb. 1961, Proctor & Mullings 22062 (BM). Cairn Curran district, two and a half miles west-north-west of Hopewell, 530 m., March 1956, Stearn 433 (BM; K), Proctor 11719 (IJ).
St. ELIZABETH: One mile north of Redgate, 150 m., March 1956, Stearn 438 (BM).
Columnea brevipila agrees with C. fawcettit in having the upper surface of the leaf beset with minute appressed hairs but differs completely in its entire sepals. It is confined to southwestern Jamaica (Text-fig. 13). Some hybrids between C. rutilans and C. urbanii come close to it in general character but differ in having their pedicels and sepals clothed with spreading hairs.
9. Columnea hirsuta Swartz, Nov. Gen. & Sp. Pl. : 94 (1788); Fl. Ind. Occ. 2: 1080 (1800) excl. syn.—Hook. in Curtis, Bot. Mag. 58 : t. 3081 (1831).—Griseb., Fl. Brit. W. Ind. Is. : 465 (1862) excl. parte-—Hanstein in Linnaea 34 : 413
THE JAMAICAN SPECIESOFCOLUMNEAANDALLOPLECTUS 227
(x865) excl. parte.—Urban, Symb. Antill. 2 : 361 (1901) excl. var. subintegra & var. fawcettit.—Morton in Contr. U.S. Nat. Herb. 29: 7 (1944).—Stearn in Proc. Linn. Soc. Lond. 170 : 141 (1959). (Text-fig. 1C, 14.)
Eusynetra bicolor Raf., Fl. Tellur. 2 : 57 (1837), nom. superfl.
Columnea hirsuta var. genuina Urban, tom. cit. : 361 (1901).
Columnea hirsuta var. pallescens Urban, tom. cit. : 362 (1901). Columnea hirsuta var. concolor Urban, tom. cit. : 362 (1901).
Shrub, the stems brownish grey, moderately slender, densely hirsute when young with long multicellular hairs. Leaves paired, those of a pair unequal, green and hirsute on both surfaces with long multicellular flaccid hairs together with short 2-3-celled hairs, the blade of the larger leaf oblong-elliptic to narrowly obovate, with the apex acute or shortly acuminate, the margin shallowly serrate, the base rounded or cuneate, 4:5-12 cm. long, 2:5-5:5 cm. broad, the blade of the smaller leaf elliptic or broadly elliptic, 2-6 cm. long, 1-8—3-8 cm. broad; petiole c. 15 mm. long. Flowers I-3 in a leaf-axil. Pedicels 1-2 cm. long, hirsute. Calyx divided to the base, red or green, densely hirsute on both surfaces with long multicellular red or colourless hairs; segments linear, slightly uneven, c. 2 cm. long in flower to 4 cm. long in fruit, 2-4 cm. broad, remotely toothed with one to three teeth on each margin. Corolla strongly 2-lipped, red with broad yellow longitudinal stripes, c. 4-5-5 cm. long, the tube mostly covered by the calyx, c. 2cm. long, the upper lip c. 2-5 cm. long, with a deeply notched middle lobe and two spreading narrowly triangular lateral lobes, the lower lip c. 2:2cm. long, oblong-linear, reflexed. Stamens exserted; filaments sparsely pubescent; anthers 1-8-2-5mm. long. Ovary sericeous; style densely glandular- pilose. Berry white.
St. ANDREW: Moresham River, quarter mile above Hermitage Dam, 350 m., March 1956, Stearn 412 (BM). MHardwar Gap, 1,200 m., Sept. 1952, Robbins 1511 (UCWI). Summit of Catherine’s Peak, 1,540m., Nov. 1952, Proctor 7341 (IJ). West slope of Catherine’s Peak, 1,300m., Jan. 1961, Proctor 21954 (BM). Catherine’s Peak, Jan. 1888, Eggers 3603 (US, type-collection of C. hirsuta var. pallescens). Vicinity of St. Helen’s Gap, 1,475 m., March 1920, Maxon & Killip 591 (US). Morces Gap, 1,520m., July 1903, Nichols 15 (K; US). Same locality, 1,500 m., March 1920, Maxon & Killip 1722 (US). Along trail between Cinchona and Morces Gap, 1,520 m., Feb. 1941, Burrowes 3563 (IJ). Near Cinchona, 1,600 m., June 1896, Fawcett 6366 (BM, type-collection of C. hirsuta var. concolor). Cinchona, windward slopes, Feb. 1915, Harris & Lawrence C 15158 (US).
PORTLAND: Trail from Morces Gap to Vinegar Hill, 1,175-1,500 m., March 1920, Maxon & Killip 694 (US). Near Mabess River, 780 m., Dec. 1905, Harris 9123 (NY). Lower eastern ridge of Mossman’s Peak, 1,600-1,700 m., May 1926, Maxon 9665 (US). East slope of the John Crow Mountains two miles southwest of Ecclesdown, 600 m., March 1951, Proctor 5703 (IJ). Same locality, Jan. 1956, Howard, Proctor & Stearn 14790 (BM). Upper north slope of Silver Hill Gap, along track toward Big Level, 1,100 m., Feb. 1963, Proctor 23255 (BM; IJ).
St. THomas: Near Whitfield Hall, 1,400 m., Jan. 1956, Stearn 123 (BM). Portland Gap, 1,640 m., Dec. 1954, Proctor 9615 (BM). Same locality, Jan. 1956, Stearn 94 (BM). Northwest slope of Yallahs Hill, 600 m., April 1956, Stearn 677 (BM).
BOT. 4, 5. 14
228 THE JAMAICAN SPECIES OFCOLUMNEAANDALLOPLECTUS
UNLOCALIZED: 1777, Shakespear (BM). W. Wright (BM). Swartz (BM, type- collection of C. hirsuta). 1843, Lane (K). Purdie (K). March (K).
An eastern Jamaican species characteristic of the Blue Mountains (Text-fig. 14), Columnea hirsuta is easily distinguished from its closest ally, C. fawcetti, by the long multicellular hairs on the upper leaf-surface. The leaves green beneath separate these species at a glance from C. vutilans, which has also more deeply toothed calyx- segments.
Usually C. hirsuta grows as an epiphyte, the vegetative shoots clinging like ivy to mossy tree trunks while the flowering shoots spread horizontally outwards, but it may sometimes be found growing terrestially among rocks.
10. Columnea fawcettii (Urban) Morton in Contr. U.S. Nat. Herb. 29 : 8 (1944).— Stearn in Proc. Linn. Soc. Lond. 170 : 141 (1959).—Raymond in Mém. Jard. Bot. Montréal 57 : 13, t.6 (1965). (Text-fig. rH, 2D, 15, 28.)
Columnea hirsuta sensu Griseb., Fl. Brit. W. Ind. Is. : 465 (1862) pro parte; non Swartz.
Columnea hirsuta var. fawcettii Urban, Symb. Antill. 2 : 363 (1901).
Rapunculus fruticosus foliis oblongis integris villosis etc. Sloane, Cat. Pl. Jam. : 58 (1696); Voy..Nat. Hist.. Jam. 1°3-557,-t. 100-f. © (4767):
Shrub, the stems grey, fairly slender, pilose with whitish hairs. Leaves paired, those of a pair equal to very unequal, green and shortly strigillose on both surfaces, the hairs of the upper surface 2-celled, 0-3-0-4 mm. long, the blade of the larger leaf mostly elliptic or narrowly elliptic, with the apex shortly acuminate, the margin remotely and obscurely toothed, the base asymmetric, cuneate, 6—10-5 cm. long, 2:5-4:5 cm. broad, the blade of the smaller leaf in shape like the larger, 3-4-5 cm. long; petiole of larger leaves 8-16 mm. long, of smaller 2-4mm. long. Flowers I to 3 in a leaf-axil. Pedicels 12-15 mm. long, hirsute. Calyx divided to the base, red, densely hirsute on both surfaces; segments slightly uneven, narrowly lanceolate, 2-3-2°8 cm. long, 3-4 mm. broad, remotely toothed, with two or three linear teeth to 2mm. long on each margin. Corolla strongly 2-lipped, red with yellow longi- tudinal stripes, c. 4-5-5 cm. long, sericeous with long whitish hairs, the tube covered by the calyx, the upper lip c. 2-8 cm. long, with a strongly notched middle lobe and two spreading narrowly triangular lateral lobes, the lower lip c. 2 cm. long, narrowly oblong, reflexed. Stamens exserted; filaments slightly pubescent; anthers c. I-5 mm. long. Ovary sericeous; style glandular-pilose. Berry ovoid, white.
Sr. ANN: Interior Road, near Browns Town, Feb. 1897, Fawcett 6587 (BM, type- collection). Dry Harbour Mountains, two miles west of Albion, 760 m., Jan. 1956, Stearn 157 (BM). Vicinity of Lydford Post Office, Governor’s Hill, 560 m., Dec. 1953, Howard & Proctor 13570 (IJ). Reynolds Mine Area near Lydford Post Office, 360-420 m., Jan. 1956, Howard, Provtor & Stearn 14602 (BM). Averham Park, 560 m., March 1956, Stearn 585 (BM). Soho, 430 m., May 1915, Harris 11993 (NY). St. Ann’s Road, four miles from Moneague, Dec. 1849, Alexander (K). Union Hill, near Moneague, April 1908, Britton & Hollick 2741 (NY). Grierfield, near Moneague, April 1908, Britton 2659 (NY). Lower part of road to Holly Mount, 400 m., March 1956, Stearn 576 (BM). Near Camperdown School, one and a half miles southwest of Gibralter, 420 m., March 1956, Stearn 568 (BM; K).
THE JAMAICANSPECIESOFCOLUMNEAANDALLOPLECTUS 229
St. CATHERINE: Vicinity of Holly Mount, Mount Diablo, 750m., May 1904. Maxon 2334 (US). Same locality, Feb. 1905, Harris 8896 (K; NY). Mount Diablo, Feb. 1916, Ridley (K). Upper slopes of Mount Diablo, 500-800 m., Feb. 1920, Maxon & Killip 429 (NY; US). Between Point Hill and Juan de Bolas Peak, 610-820 m., March 1958, Yuncker 18374 (BM).
St. ANDREW: Cooper’s Hill, Red Hills, 670-760 m., March 1954, Proctor 8488 (IJ; NY). Same locality, Feb. 1956, Stearn 255 (BM; K).
PORTLAND: Port Antonio-Moore Town Road, March 1924, Norman 225 (BM). Uncommon Hill, above Fruitful Vale Post Office, 670-760 m., April 1954, Proctor 8579 (IJ). Seamen’s Valley, 150-250 m., Feb. 1920, Maxon & Killip 81 (NY; US). Alligator Church district, one mile south of Seamen’s Valley, 70-150 m., March 1960, Proctor 20640 (BM). Northwest slope of Joe Hill, 300-670 m., April 1955, Proctor roro2 (IJ). Same locality, Feb. 1956, Stearn 243 (BM). Vicinity of Millbank, 200-300 m., Feb. 1920, Maxon & Killip 133 (NY; US). East slope of John Crow Mountains, one to one and a half miles southwest of Ecclesdown, 300-460 m., April 1954, Proctor 8603 (IJ). Same locality, 460-600 m., Aug. 1954, Webster 5579 (BM). Same locality, Jan. 1955, Proctor g798 (IJ). Ten miles southwest of Priestmans River, 460-760 m., Jan. 1951, Proctor 5255 (IJ).
St. THomaAs: Maccasucker Bump, 825-1,025 m., June 1926, Maxon 9523 (US). Ridge east of Cuna Cuna Gap, 750-840 m., June 1926, Maxon 9407 (US). North slope of Corn Puss Gap, 600 m., Jan. 1945, Barry 1058 (IJ). Sulphur Creek, above Fountain Inn [Bath Fountain], 150-200 m., Nov. 1957, Yuncker 17486 (BM).
UNLOCALIZED: Sloane, Herb. 3 : fol. 21 (BM). 1859-60, Metcalfe (BM). Purdie (K).
Columnea fawcettit is the most widespread Jamaican species of Columnea (Text-fig. 15) and was the first to be collected and recorded, having been gathered by Sir Hans Sloane as long ago as 1688-89 and illustrated by him, albeit crudely, in 1707. Never- theless, because it closely resembles C. hirsuta, it remained without varietal or specific designation until rg01, when Urban named it C. hirsuta var. fawcettit; not indeed until 1944 was its true status recognized, Morton then giving Urban’s variety specific rank as C. fawcettii. It differs from C. hirsuta principally in having only short appressed hairs on the upper surface of the leaves. Its name commemorates
William Fawcett (1851-1926), from 1880-86 assistant botanist at the British Museum, a period during which the botanical collections were transferred from Bloomsbury to South Kensington, then from 1887 until his retirement in 1908 Director of Botanic Gardens and Plantations in Jamaica. When in Jamaica he became especially interested in the Orchidaceae of the island and his monograph, prepared in collabora- tion with A. B. Rendle (1865-1938), was converted into the first volume (published in 1910) of the Flora of Jamaica. He worked continuously on this Flora at the British Museum (Natural History) until his death in 1926.
Ir. Columnea rutilans Swartz, Nov. Gen. & Sp. Pl. : 94 (1788); Fl. Ind. Occ. 2 : 1083 (1800).—DC., Prodr. 7 : 542 (1839).—Griseb., Fl. Brit. W. Ind. Is. : 465 (1862).—Hanstein in Linnaea 34 : 415 (1865).—Urban, Symb. Antill. 2 : 363 (rg0r).—Morton in Contr. U.S. Nat. Herb. 29 : 5 (1944).—Stearn in Proc. Linn. Soc. Lond. 170 : 141 (1959). (Pl. 21; Text-figs. 1G, 2F, 17.)
BOT. 4, 5. 14§
230 THE JAMAICAN SPECIESOFCOLUMNEA ANDALLOPLECTUS
Shrub, the stems grey, stout, when young strigose or almost tomentose with reddish hairs, later sparingly strigose. Leaves paired, those of a pair unequal, the upper surface dark green and strigillose with short multicellular hairs, the lower surface red-veined or completely red, pilose and almost sericeous, the larger leaf of the pair long-stalked, the blade lanceolate or narrowly lanceolate, with the apex shortly acuminate, the margin remotely serrulate, the base asymmetric, acute, 7-18 cm. long, 2-5-6 cm. broad, smaller leaf of the pair soon falling, short-stalked, the blade 2-4-5 cm. long, 1-5—2-5 cm. broad; petiole of larger leaves 1-2-5 cm. long, of smaller leaves 3-5 mm. long, sericeous. Flowers 1 to 4 in a leaf-axil. Pedicels 1-2 cm. long, strigillose. Calyx divided to base, pilose on both surfaces with long red hairs; segments 2—2:5 cm. long in flower to 3-5 cm. long in fruit, pinnately cleft with three acute teeth on each margin. Corolla strongly 2-lipped, red with longi- tudinal orange stripes, 4:5-5:5 cm. long, the tube covered by the calyx, the upper lip to 3 cm. long, with a strongly notched galea and two spreading narrowly triangular lateral lobes, the lower lip to 2:5.cm. long, narrowly oblong, reflexed. Stamens exserted; filaments glabrous or slightly hairy, pale yellow; anthers 2:5 mm. long. Ovary sericeous; style glandular-pilose. Berry white, globose, c. 12mm. broad, pilose.
HANOVER: Northwestern slope of Dolphin Head, May 1906, Harris 9265 (NY; US). Interior summit slopes of Dolphin Head, 460-520 m., April 1955, Proctor 10036 (BM; IJ). Same locality, March 1960, Proctor 20689 (BM; IJ).
St. JAMES: Chatsworth district, one and a half miles north-north-west of Maroon Town, 490 m., March 1956, Stearn 453 (BM).
St. ELIZABETH: Near Accompong, 400 m., April 1956, Stearn 1005 (BM). Cooks Bottom, east of Elderslie, 490-520 m., May 1960, Proctor 20864 (BM).
TRELAWNY: Near Troy, 600 m., April 1906, Harris 9370 (BM; K; NY). Same locality, 600-660 m., June 1906, Maxon 2852 (NY; US). Bartley’s Run, three miles north-north-west of Troy, 520 m., April 1956, Stearn 927 (BM). ‘Tyre district, two miles north of Troy, 530 m., March 1955, Proctor 9926 (IJ). Six miles from Troy, April 1954, Robertson 1058 (UCWI).
MANCHESTER: Feb. 1844, Purdie (K). Near Christiana, May 1901, Fawcett 8144 (UCWI). Heron’s Hill (Shooter’s Hill, Martin’s Hill), gt0o-940 m., March 1958, Proctor 17482 (BM). Top Hill, 820 m., March 1966, Morley 344 (BM).
CLARENDON: Peckham Woods, 750 m., May 1912, Harris r10g2 (NY; US). Same locality, March 1954, Proctor 8424 (IJ). Same locality, May 1955, Robertson 2034 (K). Knox College, Spaldings, 850 m., May 1949, Dignum 34 (IJ). Bull Head, March 1903, Fawcett 8465 (UCWI).
UNLOCALIZED: 1787, Swartz (BM, type-collection).
Columnea rutilans is a western Jamaican species (Text-fig. 17) easily recognized by the conspicuous red colouring on the under-side of the leaves and the calyx. The calyx-segments are, moreover, deeply and pinnately lobed and clothed with long reddish hairs. Although occasionally epiphytic, it mostly grows in pockets of humus on limestone rocks. Despite its distinctness from other species, it hybridizes with C. urbanii when, as at Top Hill, Manchester, it grows at the same place.
THE JAMAICAN SPECIES OFCOLUMNEA ANDALLOPLECTUS 231
HYBRIDS Columnea rutilans < urbanii (Text-fig. 16).
A number of hybrids between ornamental cultivated species of Columnea has been raised in gardens, among those named for horticultural purposes being C. x banksii Lynch (C. oerstediana Klotzsch ex Oerstedt x C. schiedeana Schlechtendal), C. x euphora H. E. Moore (C. gloriosa Sprague x C. lepidocaula Hanst.), C. x lemoiner Garnier (C. glabra Oersted x C. magnifica Oersted) and C. x vedrariensis Mottet (C. magnifica x C. schiedeana). Unless, however, plants of two species grow together or within the territory of one individual humming-bird, hybridization is unlikely often to take place in the wild. Jamaica provides the first recognized example of its occurrence.
In April, 1962, Dr. C. D. Adams collected at Top Hill near Walderston, Manchester (Text-fig. 16) a Columnea (Adams 11085; UCWI) which agreed with C. rutilans in having its leaves red beneath but diverged in its shorter leaves with longer hairs on the upper surface and its toothed rather than laciniate calyx-segments. He surmised that it might be a hybrid of C. rutilans with another species. He also collected between Contrivance and Top Hill a Columnea (Adams 11077; UCWI) with leaves like C. urbani but flowers longitudinally red-striped which likewise did not fit any recorded species. Earlier, in 1924, C. Norman (Norman 69; BM) collected near Mandeville, Manchester, an anomalous specimen.
Investigation of the Top Hill area by Dr. Brian D. Morley in 1966 revealed the existence here of both C. rutilans and C. urbanii and a series of intermediate plants evidently derived from their hybridization. These two species are so distinct in the shape, size, colour and pubescence of their leaves and calyx-segments, as well as the colour of the corolla, that an isolated intermediate specimen of unknown provenance could easily be taken to represent an independent species not closely connected with either. Such a specimen, for example, is Morley 334, hybrid 6, having elliptic leaf- blades 6-9 cm. long, 3-5-4:5 cm. broad, reddish below, also slightly toothed calyx- segments covered with minute bristles and long reddish multicellular hairs, and a yellow corolla with similar reddish hairs. The ranges of C. rutilans and C. urbanii overlap in Manchester and upper Clarendon and it is within these parishes that the plants combining their characters occur, sometimes where neither of the two species is now present. This may be due to the disappearance of these species but it may also result from the dispersal of seed of hybrid origin by birds and later by self- pollination and inbreeding of a colony thus established outside the original locality. Morley collected ten nothomorphs at Top Hill in which the characters of C. rutilans and C. urbanii were variously mingled.
The following gatherings also appear to show traces of hybrid origin, although much closer to C. urbanii than to C. rutilans:
MANCHESTER: Grounds of West Indian Training College near Mandeville, 760 m., March 1958, Proctor 17455 (BM; IJ). Providence, one and a half miles west-south- west of Newport, 800-820 m., Jan., 1961, Proctor 21905 (IJ). Rose Hill, 760 m., Oct. 1965, Morley 274C (BM). Moorlands estate, two miles northeast of Spur Tree,
232 THE JAMAICAN SPECIES OF COLUMNAE AND ALLOPLECTUS
760 m., Jan. 1956, Howard, Proctor & Stearn r47or (IJ). Newport, 680-760 m., Feb. 1966, Powell & Green 1663 (IJ). CLARENDON: Peckham Woods, 760 m., March 1956, Robertson 2720 (UCWI; IJ).
ALLOPLECTUS Mart. ALLOPLECTUS Mart., Nov. Gen. & Sp. Pl. Brasil. 3 : 53 (1829), nom. cons.
Shrubs or herbs. Leaves opposite, those of a pair equal to unequal, usually long- petiolate. Flowers solitary or fasciculate in leaf-axils or borne in short axillary racemes, with bracts. Calyx 5-partite. Corolla inferior, erect or oblique in the calyx, gibbous but not spurred at base, the tube cylindric or ventricose, contracted or not expanded in the throat, the limb regular or almost so with five short rounded lobes. Stamens four, didynamous, the filaments united in the lower part, free and filamentous above, the anthers free or cohering in pairs, their loculi parallel, longi- tudinally dehiscent. Disc gland large and dorsal, sometimes accompanied by smaller glands. Ovary superior; stigma usually 2-lobed. Fruit a capsule or berry; seeds minute, longitudinally or obliquely striate.
Lectotype (see p. 184): A. sparsiflorus Mart. (= A. Mirtellus (Schott) Preston).
The genus Alloplectus as defined above comprises about 80 species, native to tropical
America, including the Caribbean islands.
t. Alloplectus pubescens (Griseb.) Fawcett, Provis. List Flow. Pl. Jam. : 28 (x1893).—Urban, Symb. Antill. 2 : 358 (x902).—Morton in Contr. U.S. Nat. Herb. 29 : 15 (1944). (Text-figs. 19, 29gG—H.)
Pterygoloma pubescens Griseb., Fl. Brit. W. Ind. Is. : 464 (1862). Columnea pubescens (Griseb.) Kuntze, Rev. Gen. Pl. 2 : 472 (1892).
Shrub of dwarf creeping habit, the stems grey, slender, 2-3 mm. thick, hirsute with colourless hairs. Leaves paired, those of a pair almost equal to markedly unequal, short-stalked, green and densely hirsute on both surfaces, some of the hairs on the upper surface glandular, the larger leaf with the blade elliptic or narrowly obovate, symmetric, the apex acuminate, the margin inconspicuously denticulate, the base cuneate, 4-5-8 cm. long, 2-3 cm. broad, the smaller leaf often half as long as the larger leaf; petiole 2-10 mm. long. Flowers solitary in leaf-axils. Pedicel 7-15 mm. long, hirsute. Calyx divided to the base, green, hirsute on both surfaces with — colourless glandular hairs; segments narrowly elliptic, acuminate, c. I cm. long, entire. Corolla tubular, slightly swollen at the middle, c. 2 cm. long, reddish orange, the tube c. 4 mm. broad at the middle, strigose outside, the lobes almost equal, I-2 mm. long. Stamens included; filaments glabrous; anthers c. I-rmm. long. Ovary glabrous; style glabrous. Berry pink or purplish, c. 5 mm. broad.
PORTLAND: 610 m., Wilson (K; GOET, type). John Crow Mountains, one and a third miles south-west of Ecclesdown, 460-610 m., Aug. 1954, Webster 5569 (BM; IJ). Same locality, Dec. 1955, Proctor 11346 (IJ). Same locality, Jan. 1956, Howard, Proctor & Stearn 14792 (IJ).
THE JAMAICAN SPECIES OF COLUMNAE AND ALLOPLECTUS 233
Alloplectus pubescens is a rare and inconspicuous species confined to the rain forest of the mountains in the extreme east of Jamaica (Text-fig. 19) and differs from other West Indian species of Alloplectus in its entire sepals, which are hirsute on both surfaces. The corolla is variously described as “ pale orange’’, “‘ orange’’ and “pale salmon-pink ”’, the fruit as “ pink ”’ or “ light rose-purple ”’.
Fic. 29. Alloplectus grisebachianus (Kuntze) Urban: A, flowering shoot (Proctor 22648, 16539); B, section of flower after anthesis (Harris 555); C, calyx (Harris 555); D, anthers (Harris 555); E, anther, back view (Harris 555); F, gynoecium (Harris 555); Alloplectus pubescens (Griseb.) Fawcett: G, calyx (Wilson); H, seed (Webster 5569). Alloplectus ambiguus Urban: J, section of ovary (after H. E. Moore, 1957).
234 THE JAMAICAN SPECIES OF COLUMNAE AND ALLOPLECTUS
2. Alloplectus grisebachianus (Kuntze) Urban, Symb. Antill. 2 : 357 (1901).— Morton in Contr. U.S. Nat. Herb. 29 : 18 (1944). (Text-figs. 18, 29gA—F.)
Pterygoloma cristatum Griseb., Fl. Brit. W. Ind. Is. : 464 (1862).
Columnea grisebachiana Kuntze, Rev. Gen. Pl. 2 : 472 (1892).
Alloplectus cristatus (Griseb.) Fawcett, Provis. List Flow. Pl. Jam. : 28 (1893); non A. cristatus (L.) Mart. ex G. Don (1838).
Alloplectus grisebachianus var. ochrotrichus Urban, Symb. Antill. 5 : 497 (1908).—Morton,
loc. cit. (1944). Columnea hunnewellu L. B. Smith in Rhodora 39 : 275 (1937).
Shrub of dwarf creeping habit, branched, sometimes forming a bush with descending shoots, the stems grey, slender, 2-3 mm. thick, densely hirsute when young with reddish hairs or colourless hairs, the leaf-scars prominent. Leaves paired, those of a pair mostly unequal, short- or long-stalked, green and hirsute on both surfaces with long rather appressed hairs, the larger leaf of the pair with the blade elliptic or ovate, slightly or distinctly asymmetric, the apex shortly acuminate, the margin shallowly denticulate, the base rounded, 2:5-6cm. long, 1-5-3 cm. broad, the smaller leaf sometimes half as long as the larger leaf and falling earlier; petiole 3-10 mm. long. Flowers solitary in the leaf-axils. Pedicel 2-10 mm. long, hirsute. Calyx divided to the base, green, hirsute outside with long reddish hairs or colourless hairs, glabrous inside; segments laciniate, c. 10-12 mm. long, with two to five prominent teeth on each side. Corolla tubular, slightly swollen at the middle, c. 2cm. long, dull purplish red, the tube c. 5-6 mm. broad at the middle, hirsute outside with long spreading hairs and very minute appressed hairs, the lobes almost equal, c. 1mm. long. Stamens included; filaments glabrous; anthers c. r mm. long. Ovary glabrous; style minutely pubescent in the upper part. Berry white, c. 7mm. broad.
TRELAWNY: near Troy, 610m., May 1903, Harris 8549 (BM; US; lectotype collection of Alloplectus grisebachianus var. ochrotrichus).
MANCHESTER: Olive River, Christiana district, gro m., Oct. 1901, Harris 8413 (BM).
CLARENDON: Peckham Woods, 750 m., Dec. 1917, Harris 12766 (K; US). Same locality, Dec. 1955, Stearn ro (BM).
St. ANN: Albion, March 1850, Purdie 555 (K, lectotype of Pterygoloma cristatum). Dry Harbour Mountains, two miles west of Albion, 600 m., Jan. 1956, Stearn 158 (BM; K). Mount Diablo, Feb. 1916, Ridley (K). Same locality, Jan. 1938, Hunnewell (BM; K; US). Union Hill, near Moneague, 750 m., Britton & Hollick 2744 (NY). Hollymount, Feb. 1905, Harris 8891 (NY). Same locality, Sept. 1906, Britton 718 (NY). Along the road to Hollymount, Mount Diablo, 800 m., Aug. 1957, Proctor 16539 (BM). Ramble, Claremont, 450 m., Jan. 1898, Fawcett & Harris 7168 (BM).
UNLOCALIZED: Alexander (GOET). Wullschlégel (GOET)
The transfer by Fawcett in 1893 of Grisebach’s epithet cristatus from Plerygoloma to Alloplectus created a later homonym of Alloplectus cristatus (L.) Mart. ex G. Don, based on Besleria cristata L. (1753). In 1892, however, Kuntze united Alloplectus, Nematanthus and Hypocyrta with Columnea and as he could not use Grisebach’s
THE JAMAICAN SPECIES OF COLUMNAE AND ALLOPLECTUS 235
epithet owing to the prior claim of Linnaeus’s published under Besleria, which resulted in the name Columnea cristata (L.) Kuntze, he coined the new name Columnea grisebachiana for Pterygoloma cristatum Griseb. Thereby he provided a legitimate epithet for use under Alloplectus. For a biography and bibliography of August Heinrich Rudolf Grisebach (1814-79), the G6ttingen professor of botany whom the name commemorates, see Journ. Arnold Arb. 46 : 245-260 (1965).
Alloplectus grisebachianus is restricted to the central uplands of Jamaica (Text-fig. 18). The typical and commonest form of the species is notable for the long reddish multicellular hairs which clothe the branchlets, petioles, leaf-blades, pedicels, calyx and corolla. To this belongs the type-specimen of Columnea hunnewellit L. B. Smith collected by Hunnewell and Griscom in March 1936, on Mount Diablo. In speci- mens collected by Harris at Troy (Harris 8549) and at Olive River (Harris 8413) the hairs are colourless and Urban distinguished them as A. grisebachianus var. ochro- trichus in 1908. A gathering made by Hunnewellin January 1938 on Mount Diablo includes, however, both forms; the loss of colour in the hairs has probably taken place independently in several localities.
Alloplectus ambiguus Urban is a species not of Jamaica but of Puerto Rico; because it is adequately known from wild and cultivated material and is allied to the Jamaican species, it has been included above in the taxonometric survey (pp. 207-214) to increase the representation of Alloplectus and in Fig 29 (p. 233) to illustrate the placentation of these plants.
SPECIAL LITERATURE
Bonn, J. 1947. Field Guide to Birds of the West Indies. Ed. 2. New York.
British Museum (NaTurRAL History). 1966. Report on the British Museum (Natural History), 1963-1965. London.
Burman, J. 1756. Plantarum Americanarum Fasciculus Quartus. Amsterdam & Leiden.
EBERLE, P. 1956. Cytologische Untersuchungen an Gesneriaceen. Chromosoma 8 : 285-316.
Farcri, K. & VAN DER Pijt, L. 1966. The Principles of Pollination Ecology. Oxford.
Fawcett, W. 1893. A Provisional List of the Indigenous and Naturalised Flowering Plants of Jamaica. ‘Kingston, Jamaica.
FusseELL, C. P. 1958. Chromosome numbers in the Gesneriaceae. Baileya 6 : 117-125.
Gower, J. C. 1966. Some distance properties of latent root and vector methods used in multivariate analysis. Biometrika 53 : 325-338.
GRANT, V. & GRANT, K. A. 1966. Records of hummingbird pollination in the western American flora. Aliso 6 : 51-66.
GREENEWALT, C.H. 1960. Hummingbirds. New York.
GriseBAcH, A. H. R. 1859-64. Flora of the British West Indian Islands. London.
Harsorne, J. 1966a. Caffeic acid ester distribution in higher plants. Z. Naturforsch.
21B : 604-605.
1966b. Comparative biochemistry of the flavonoids—II. 3-desoxyanthocyanins and
their systematic distribution in ferns and gesnerads. Phytochemistry 5 : 589-600.
KeEnprick, L. B. & WeREsSvuB, L. K. 1966. Attempting neo-Adansonian computer taxonomy at the ordinal level in the Basidiomycetes. Syst. Zool. 15 : 307-329.
236 THE JAMAICAN SPECIES OF COLUMNAE AND ALLOPLECTUS
Lez, R. E. 1962a. Chromosome numbers in the Gesneriaceae. Baileya 10 : 33-45.
1962b. Achimenes coccinea, a tetraploid; and additional counts of chromosome numbers
in the Gesneriaceae. Bazleya 10 : 146.
— 1964. Additional chromosome numbers in the Gesneriaceae. Baileya 12 : 159.
— 1966. Additional chromosome numbers in the Gesneriaceae. Baileya 14 : 35-36.
— 1967. Additional chromosome numbers in the Gesneriaceae. Baileya 15 : 118.
— & Grear, J. W., Jr. 1963. Additional chromosome numbers in the Gesneriaceae. Batleya 11 : 131.
LEEUWENBERG, A. J. M. 1958. The Gesneriaceae of Guiana. Acta bot. neerl. 7 : 291-444.
Moore, H. E. 1957. African Violets, Gloxinias, and their relatives. New York.
Morey, B. D. 1966. Coluwmnea and aspects of its evolution. Sci. Notes & News, Jamaica
2: 13-15.
1967. In A. Léve, IOPB chromosome number reports. Taxon 16 : 215-222.
Morton, C. V. 1944a. The West Indian species of Columnea. Contr. U.S. natn. Herb.
29 : I-I5.
1944b. The West Indian species of Alloplectus. Contr. U.S. natn. Herb. 29 : 15-19.
1963. Another Columnea new to cultivation. Baileya 11 : 23-27.
PITELKA, F. A. 1942. ‘Territoriality and related problems of North American hummingbirds. Condor 44 : 187-204.
RayMonp, M. 1965. Species of Columnea and Trichantha in cultivation in the Montreal Botanic Garden. Mém. Jard. bot. Montréal 57.
Rocers, O. M. 1954. Chromosome counts in Gesneriaceae. Batleya 2 : 14-18.
SEALY, J. R. 1950. Nomocharis and Lilium. Kew Bull. 1950 : 273-297.
SHEALS, J. G. 1965. The application of computer techniques to Acarine taxonomy. Proc. Linn. Soc. Lond. 176 : 11-21.
Sims, R. W. 1966. The classification of the Megascolecoid earthworms: an investigation of © Oligochaete systematics by computer techniques. Proc. Linn. Soc. Lond. 177 : 125-141.
Soka, R. R. & SNEATH, P. H. A. 1963. Principles of Numerical Taxonomy. San Francisco
& London. STEARN, W. T. 1959. A botanist’s random impressions of Jamaica. Proc. Linn. Soc. Lond, 170 : 134-147.
1968. Observations on a computer-aided survey of the Jamaican species of Columnea and Alloplectus. In V. Heywood, Modern Methods in Plant Taxonomy: 219-224.
Swartz, O. 1788. Nova Genera et Species Plantarum seu Prodromus. Stockholm, Uppsala & Abo. UrBaNn, I. 1go0r. Enumeratio Gesneriacearum. Symb. Antill. 2 : 344-388.
1909. Nova genera et species IV. Symb. Antill. 6 : I-55. } Watson, L., Wittiams, W. T. & Lance, G. N. 1967. A mixed-data numerical approach to- angiosperm taxonomy: the classification of Ericales. Proc. Linn. Soc. Lond. 178 : 25-35+—
eg ee
W. T. STEARN, D.Sc., Sc.D.
Dept. of Botany
BritisH Museum (NATURAL HIsTory) CROMWELL RoAD
Lonpon, S.W.7
Bull. Br. Mus. nat. Hist. (Bot.) 4, 5 PEATE Mia
tog
BaresH Mosq b ONATIRAL taserced Tks huts date las | | Lobel |
HERBARIUM MUSE! BRITARNIC! henna
KY \ N
bd
Sf FLORA OF TAMAICA
Colom AA Ey Tt & ia Lene. St &
VOR. Roe det WY, Ste aun has ‘esnntamonto : HG
INSTITUTE OF JAMAICS FLORA OF JAMAICA Parish: WESTMORELAND
Name Columnea jamaicensis Urb. tocatity: 1 mile W.N.W. of HOPEWELL.
Altitude: 1750 ft. pate Movember 21, 1955 Habitas, Wooded limestone hillside.
Trailing epiphyte; calyx red; corolla Neles)yellow, streaked with orange.
olicetor George Ry Preetor so, 41232
jamaicensis
Columnea jamaicensis Urban (Proctor 11232; representative specimen in Herb. Brit. Mus. (Nat. Hist.), London).
| BOT. 4, 5. 15 |
PLATE 15
4 Br. Mus, nat. Hist. (Bot.) 4, 5
‘(uopuoT ‘(SIE] “3@N) ‘SN “WIG ‘q4uoZy {zz4vmg) ZUIVMG vprdsiy vauwmnjoy Jo adAyo[oOFY
‘puowAry “JA 0VOYg
‘Uspley) WUv}OY [VI}UOW! 7 Pozearz[N zjIeMg vpidsiy vauwmnjoy jo yueyd Surary
081) SEot :Z bid (8841) Ab
16 ppldsiy Vauwajey
NYWIDRdS AdAL TRIOM GS Ug “Sstwmog |,
.
y
ull. Br. Mus. nat. Hist. (Bot.) 4, 5
ts
ey 1 2 ym iNaresat History! ROTOR APE
yy 1 3i 4 8 Lbidhe hatibaiba
redeataferad as
| Revres Mute
i
HERBARIUM MUSET @RTTANNICT cata
diteite
HOLOTYPE Columnea urbanii
Stearn
FLORA OF JAMAICA
Coluwnea " &P-
panisn Manchester aur yoooft ware 2S. FL. page to Ituntiey ‘ On
leaves crrwded nth Mert wa
youks bu road at Ei ps af gh oets' | Besekt 4reen “pp sede Sedat mostly So. elites :
» week me bing ow lamer de a dect 0-5 mn lms, the
aie waved sebwoen Eke Yeing tsrelia i ~ ate qe! sts is Procter weil nf Cou. WT Sream 80. 36 {
Columnea urbanii Stearn (Stearn 361; holotype in Herb, Brit, Mus. (Nat. Hist.), London).
PEATE 16
Bull. Br. Mus. nat. Hist. (Bot.) 4, 5 PEATE 17
ea ee | JAMAICA .
HERBARIUM MUSE! BRITANNIC! Jf ‘paceman set te
HOLOTYPE Columnea proctorii
Stearn
FLORA OF JAMAICA Co { Um ne aw laws api)
| fe St. Sumas aut, 1600 FE ware 10 Mack 195% we Chatswarth district, 14 miles NN. Ww. ¥ Mareen Town
ighutts we th Loney dranche ad & Pres dom Meing ob se line pith im presses BS wales Wath Grey ce Hociee ee |
Orlin sdlew asarlate with leract obinal sh W.T Steamy and ckripes - pelle Gallows no, asi ¥ tabs
ee oe
Columnea proctorii Stearn (Stearn 451; holotype in Herb. Brit. Mus. (Nat. Hist.), London).
proctori i
Bull. Br. Mus. nat. Hist. (Bot.) 4, 5
mT Mussa NOTOGRAPH
| 17| ile ‘|
ISOTYPE Columnea proctorit
Stearn
FLORA OF JAMAICA
CoLunen
fans St. Janes ALT imcatny Ghabeworth district Raroon Tow.
ight green
polles yalisw. CO. WL oT. SEARS
HERBARIUM MUSEI BRITANNICA i ee
PLATE 18
Columnea proctorii Stearn (Stearn 451; isotype in Herb, Brit. Mus. (Nat. Hist.), London).
} PLATE ro Bull. Br. Mus. nat, Hist. (Bot.) 4, 5
4
TYPE SPECIMEN Columnea subcordata
Morton im Contr, V, §, Nat, Hevb. 29: 6 (i944)
UNITED STATES NATIONAL MUSEUM
PLANTS OF JAMAICA
7 ? x. Dut fic y aay f [te ef Vins Whidsor, Parish of Trelawny; altitude 100 to 150 meteors
Ne Ving 4 GERRIT S. MARR, Cothenter aye F
Columnea subcordata Morton (Miller 1556; holotype in U.S. Nat. Herb., Washington).
Sig
Bull. Br. Mus. nat. Hist. (Bot.) 4, 5 PLATE 26
Iennstanealaenslanaabisrshinsslopuslistsliunntauay) MBRAARIUM MUSEI BRITAWNICE :
FLORA OF JAMAICA Solames angenten @ise. (TorotyPs) Panis Manchester atr, 2900 f%. oars 25. IL, lose
Nasareth, par rooks behind the Moravian OM Gawd tk tap or B10.
Shoots crey, to 70 om. long; leares *pfendinc, silky, the longest vo 17 oa. long, 5 om. broad;
bem os say _— corelle pale yellows fruit » Pink, ll mm. Long, 8 ms. broad, cot, WF, seraan we, 20S
Columnea argentea Griseb. (Stearn 359; topotype in Herb. Brit. Mus. (Nat. Hist.), London).
Bull. Br. Mus. nat. Hist. (Bot.) 4, 5
PIT ree er ephingag } ] Perm Ng ong
I i | Biovounany atten 1 2 : ‘a
} bth tt 4d
AMAL Re yeas tye HERBARIUM mics i #
FLORA OF JAMAICA
PARTSH 9%» oe 2 sATE
OO ALITY
(0k, WF. STRgRM
Coll, orBsiw No. 1095
rutilans
Eg
PLATE 21
Columnea rutilans Swartz (Stearn 1005; representative specimen in Herb. Brit. Mus. (Nat.
Hist.), London).
OR LITTLE KNOWN HIMALAYAN SPECIES OF ce TIA AND VERATRILLA (GENTIANACEAE
“ .
H. SMITH
B LLETIN. OR a aces
USEUM (NATURAL HISTORY) — aaa | Vol. 4 No. 6
FT le Aire ees Ts
=
NEW OR LITTLE KNOWN HIMALAYAN SPECIES OF SWERTIA AND VERATRILLA
(GENTIANACEAE)
BY HARRY SMITH, Institute of Systematic Botany, University of Uppsala
Pp. 237-258 ; 7 Text-figures, Plates 22-37
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) BOTANY Vol. 4 No. 6 LONDON: 1970
THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be com- pleted within one calendar year.
In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department.
This paper is Vol. 4 No. 6 of the Botanical series.
World List abbreviation Bull. Br. Mus. nat. Hist. (Bot.).
© Trustees of the British Museum (Natural History), 1970
TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 18 August, 1970 Price me
NEW OR LITTLE KNOWN HIMALAYAN SPECIES OF SWERTIA AND VERATRILLA GENTIANACEAE
By HARRY SMITH
SUMMARY
TEN species of Swertia : S. acaulis, S. grandiflora, S. virescens, S. pseudo-hookert, S. staintonit, S. assamensis, S. crossoloma, S. sblendens, S. gracilescens and S. burmanica are described as new. The first eight of these, with S. hookert C. B. Clarke and S. candelabrum H. Smith, form a distinct group within Swertia sect. Swertia and a key is provided for this group. The new name S. franchetiana is proposed for S. stricta Franch., a later homonym of S. stricta Collett & Hemsl. Kingdonwardia codonopsidoides Marq. is shown to be a synonym of Swertia racemosa (Griseb.) C. B. Clarke and reasons are given for not maintaining Kingdonwardia as a genus distinct from Sweriia. Extensions of range are recorded for S. kiharae Kitam. and the previously described species mentioned above.
Certain errors in Franchet’s original account of Veratrilla are pointed out, Swertia mekongensis Balf. f. & Forr. is treated as a synonym of Veratrilla batlloni Franch. and Swertia burkiliana W. W. Smith is transferred to Veratrilla. Extensions of range of both species are recorded. |
INTRODUCTION
Until this century the world’s herbaria contained few specimens of plants from the Himalayan region. In the last few decades, however, a very considerable amount of material has been collected in Tibet, Assam, Bhutan, Sikkim, and, more recently, Nepal, much of this in the course of expeditions under the auspices of the British Museum (Natural History). This paper deals with those specimens of the genera Swertia and Veratrilla that have been collected in those countries and that either represent new species or provide new information about species previously described. Except for a few specimens from the Royal Botanic Garden, Edinburgh, indicated by “‘ (E)’’, all those cited are in the herbarium of the British Museum (Natural History) ; there are duplicates of the great majority in Uppsala.
A*
240 HIMALAYAN SWERTIA AND VERATRILLA
SWERTIA L. SWERTIA L., Sp. Pl. 1 : 226 (1753) ; Gen. Pl., ed. 5 : 107 (1754).
Sectio SWERTIA
Swertia subgen. Euswertia C. B. Clarke in Hook. f., Fl. Brit. Ind. 4 : 127 (1883), nom. invalid. Swertia sect. Euswertia Gilg in Engl. & Prantl, Naturl. Pflanzenfam. 4 (2) : 88 (1895), mom. invalid.
SWERTIA KIHARAE Kitam. in Acta Phytotax. & Geobot. 16 : 133 (1956). (Plate 22 ; Text-fig. I.) CHITRAL : Gangallat Gol, S.W. of Chitral, 2,700 m., amongst shrubs at edge of
stream, petals and filaments white, anthers black, 18 June 1958, Stainton 2708. Golan Gol, 3,000 m., edge of stream, white, 13 July 1958, Bowes-Lyon 63.
b
a c Fic. 1. Swertia kihavae Kitam. : a, corolla lobe with nectary and stamen ; b, calyx ; c, ripening capsule (Stainton 2708). All x5. These are interesting finds of a species originally described from Afghanistan. No species of the Swertia aucheri Boiss. group has been recorded until now from the Indian sub-continent. |
THE SWERTIA HOOKERI GROUP OF SPECIES
The botanical explorations in the remote parts of the Himalayas during this — century have shown that Swertia hookeri C. B. Clarke, originally described from a single collection from Sikkim, is not an isolated species but is instead a member of a complex group consisting of not less than ten distinct species, all obviously of close — affinity. Unfortunately the material in herbaria is not rich ; collectors do not relish | putting such robust and unwieldy plants in the press.
HIMALAYAN SWERTIA AND VERATRILLA 241
This group is well differentiated from other members of the section Swertia by the following characters : 4-merous fairly large to very large flowers, single nectary, robust growth, thick hollow stem, which can be 0-8 m. high or more (the abbreviated S. acaulis H. Smith excepted), and seeds globose and unwinged or much flattened and winged.
A distinctive character of unusual nature divides this group into two parts, each with five species. In the first part the margin of the leaves and sepals is glabrous, though a cellular papillosity is visible at a high magnification (text-fig. 2, b). In the other the margin is cut into hair-like, chlorophyllaceous lacinules only 0-3-0°5 mm. long (text-fig. 4, b, c). No specimen has been found that deviates in the slightest way from one or other of these two patterns. The species development seems to have followed parallel lines in these two groups. In both we have species with naked or with fimbriate nectaries, and species with cymose or with pyramidal inflorescences. Whether there is a similar parallelism in types of seed cannot yet be determined ; there are four species of which there is not available ripe seed nor ovules so far advanced that the shape of the seed can be safely presumed. At present we know that globose wingless seeds are present in both groups but flattened winged seeds are so far known only in the first.
The following is a key to this Group of species:
Margin of leaves and sepals entire, with obsolete cellular papillae (text-fig. 2, b):
Nectary naked: Inflorescence cymose, many-flowered ; seed flattened : Plant 20-80 cm. tall ; inflorescence of distant axillary cymes, rarely
moderately branched ; seed discoid, circularly winged . S. hookert Plant subacaulous, stem at most 2 cm. tall ; seed oval, the obtuse ends unequally and irregularly winged ‘ : : : S. acaulis
Inflorescence pyramidal ; seed globose, not winged :
Large robust plant 60 cm. or more tall ; petals reddish, 3 cm. long by 1-6
cm. broad ; inside base of sepals with a ring of black hairs S. grandiflora Plant 20-30 cm. tall ; petals greenish, 1-2-2-0 cm. long by o- 7-1-2 cm. broad ; inside base of sepals glabrous . : : . SS. virescens Nectary densely fimbriate ; plant 30-40 cm. tall ; inflorescence pyramidal ; flowers yellowish-greenish, with glabrous and scarcely winged pedicels; seeds globose, not winged ’ ; . S. pseudo-hookert Margin of leaves and sepals minutely and somewhat irregularly lacinulate, lacinules
chlorophyllaceous, hair-like, 0-3-0-5 mm. long (text-fig. 4, b, c) : Nectary naked, its margin smooth :
Stem deeply 5-grooved ; inflorescence pyramidal, copiously branching from base and nodes ; wings of stem and pedicels ciliate ; petals 2 cm. long by I-7 cm. broad ; ; seed globose, not winged . : . S. staintoni
Stem terete or nearly So ; inflorescence of axillary cymes not or moderately branched ; wings of stem and pedicels glabrous ; petals 2-5 cm. long by 2 cm. broad ; (seed not seen) : . S. assamensis
Nectary toothed or fimbriate at the margin :
242 HIMALAYAN SWERTIA AND VERATRILLA
Margin of nectary toothed with pointed projections not more than 0-5 mm. long, less than 1/3rd of the diameter of the nectary ; flowers few on robust, curvingly ascendent branches ; , S. candelabrum
Margin of nectary fimbriate with filamentous projections 1-2 mm. long, 4 or more of the diameter of the nectary ; inflorescence pyramidal :
Connective of anthers prolonged, often claw-like ; flowers pale creamy green ; petals I-g cm. long by 1-0 cm. broad ; wings of pedicels ciliate
S. crossoloma
Connective of anthers not prolonged ; flowers brick red ; petals 2:5 cm. long by 2:0 cm. broad ; wings of pedicels glabrous except at point of junction with the decurrent sepals. : . S. splendens
SWERTIA HOOKER! C. B. Clarke in Hook. f., Fl. Brit. Ind. 4 : 127 (1883). (Plate 23 ; ©
Text-fig. 2 a—d.)
T1BET : Kongbo, Doshong La, 4,000 m., on grassy slope at base of precipice, herb to I m., calyx green, corolla greenish yellow with faint browny purple veins, filaments and ovary purple-black, anthers dull yellow, 15 July 1938, Ludlow, Sherriff and Taylor 5256. Same locality, amongst shrubbery, calyx green, corolla yellowish green, anthers and ovary dark green, 16 Aug. 1947, Ludlow, Sherriff and Elliot 14376.
NEPAL : Barun Khola, N. of Num, Arun Valley, 3,800 m., on open slopes, corolla — greenish brown, filaments white, 12 June 1956, Stainton 641. S. of Topke Gola, Arun-Tamur watershed, 4,300 m., on open slopes, corolla, filaments and anthers bronze-coloured, 9 July 1956, Stainton 903.
SIKKIM : Changu, 3,700 m., in turfy level space on hill, red flowers, spikes 60-90 cm., 8 Sept. 1913, R. E. Cooper 832. North Chakung Chu, 4,000 m., on peat hillside, I m. tall, red flowers, many whorls, 20 Sept. 1913, R. E. Cooper 918.
BHUTAN : Ju La Mangde Chu, 4,300 m., in open alpine zone, calyx green, corolla brownish red, 19 July 1949, Ludlow, Sherriff & Hicks 16892. Parshong, 4,000—3,800 m., tall, red, 27 July 1914, R. E. Cooper 1993.
Clarke’s original description, although based on only two gatherings, is very accurate and complete and there can be no doubt about the identity of his plant.
Swertia acaulis H. Smith, sp. nov. (Plate 24 ; Text-fig. 2e.) Swertia acaulis H. Smith ex Nilsson in Grana Palyn. 7 : 111 (1967), nom. nud,
Omnino glabra, radix validus verticalis, caulis valde abbreviatus 1-2 cm. altus, flores numerosos, cymosos, subaequaliter 3-4 cm. longe pedicellatos edens. Habitu S. multicaulis D. Don' subsimilis sed distat : floribus multo majoribus albis (nec caeruleis), nectario perfecte nudo (nec fimbriato), seminibus complanatis, ambitu ovalibus, in apicibus inaequaliter alatis (nec subglobosis exalatis).
Folia rosularia desunt, caulina aggregata, numerosa, lanceolata, acuta, trinervia,
1 C. B. Clarke in Hook. f., Fl. Brit. Ind. 4 : 129 (1883) says that S. multicaulis is 5-merous, whereas it is 4-merous, as correctly stated in the original diagnosis by D. Don, Prodr. Fl. Nepal : 128 (1825).
HIMALAYAN SWERTIA AND VERATRILLA 243
5-7 cm. longa, ad 0-9 cm. lata, lamina in petiolum subaequilongum sensim angustata. Flores 4-meri ; calyx fere ad basin fissus, lobi ovati, acuti, 11-16 mm. longi et 5—7 mm. lati ; corollae tubus perbrevis, lobi obovati, rotundato-truncati, apice minute lacerulati, 18 mm. longi et 11 mm. lati, nectario nudo nigro ; stamina tubo adnata, filamentis liberis 11 mm. et thecis 2 mm. longis, connectivo antherarum fere 0-5 mm. longe aristulatim protracto ; ovarium anguste ovatum in stigmas sessiles angustatum ; semina complanata, ambitu ovalia, I-0x 0-6 mm. magna, in apicibus valde inaequa- liter alata.
{
d e
Fic. 2. Swertia hookeri C. B. Clarke : a, corolla lobe with nectary and stamen ; b, margin of leaf, with chlorophyllaceous area dotted ; c, middle part of rosular petiole (Ludlow, Sherriff & Elliot 14376) ; d, seed from above and in cross section (holotype) Swertia acaulis H. Smith : e, seed from above (Lall Dhowj 0291). b X50,e XI0,a,c,d X5.
NEPAL : Foot of Rolwaling, 3,700-5,500 m., yellow, 1930, Lall Dhowj 0291 (holo- type). Dudkund, 5,500 m., yellow, 24 Aug. 1932, K. N. Sharma E504. Wabak Khola, east of Num, Arun Valley, 4,600 m., on open slopes, petals white, filaments and anthers blue, 12 Aug. 1956, Stainton 1334.
244 HIMALAYAN SWERTIA AND VERATRILLA
Swertia grandiflora H. Smith, sp. nov. (Plate 25.)
Planta 60 cm. vel ultra alta S. candelabro H. Smith subsimilis, sed distat : margine foliorum et sepalorum integro (nec lacinulato), floribus majoribus 3 cm. longis, calycis lobis late ovalibus obtusis (nec cordato-triangularibus, acutis).
Caulis robustus ad 1-6 cm. diametiens, ut rami et pedicelli modeste quadrialatus, glaber. Inflorescentia pyramidalis, ramis robustis, paucifloris e nodis 2 arcuatim adscendentibus. Folia margine integra, rosularia ad 18 cm. longa et 6 cm. lata, 7-13 nervia, lamina elliptica apice acuta in petiolum latum brevem vel longiorem
angustata, caulina perpauca ad 9:5 cm. longa et 6 cm. lata, sessilia, nervisc.g Flores —
patuli ad 5:5 cm. diametientes, rubroviride striati ; calycis tubus perbrevis basi interno breviter nigro-barbatus, lobi inaequales, duo majores ad 1-9 cm. longi et 15 cm. lati, ovales, obtusi vel subobtusi, glabri ; corollae tubus brevis, lobi oblongi, apice rotundati, ad 3 cm. longi, superiori parte 1:5-1-7 cm. lati, nectario nudo sub- quadrangulari margine paullo elevato cincto ; stamina tubo adnata, filamentis liberis crassis 10 mm. longis, antheris caeruleis 3 mm. longis ; ovarium conicum stigmatibus parvis sessilibus coronatum ; capsula matura (e Ludlow & Sherriff 1040) 2 cm. longa, valvis apice recurvantibus, pedicellis ad 4-10 cm. prolongatis ; semina ovali-globosa, exalata, 1-7-2‘:0—1-4-1°6 mm. magna, testa levi.
BuHUuTAN : Shingbe, Me La, 4,000 m., on open hillside, very common, flowers striped dull red and green, 14 June 1949, Ludlow, Sherriff & Hicks 20732 (holotype). MeLa 4,300 m., open grassy hillside above tree line, 6 Oct. 1934, Ludlow & Sherriff 1040.
Swertia virescens H. Smith, sp. nov. (Plate 26 ; Text-fig. 3c, d.) Swertia virescens H. Smith ex Nilsson in Grana Palyn. 7 : 112 (1967), nom. nud.
Species 15-20 cm. alta, S. hookeri C. B. Clarke affinis sed distat : planta multo minore, inflorescentia pyramidale, floribus vix 1-5 cm. longis, foliis omnibus obtusis (nec subacutis) caulinis sessilibus (nec longe petiolatis).
Caulis glaber, quadrialatus, superiori dimidia parte 3-nodosus, e nodo infimo rami 4, pluriflori, suberecti, florendi tempore 4-6 cm. longi editi, in nodo supremo flores numerosi cymosi, omnes graciliter pedicellati, pedicellis glabris modeste quadrialatis. Folia rosularia numerosa, lamina ovali, obtusa, 7-nervia, margine integra, c. 5 cm. longa et 2°5 cm. lata, in petiolum ad 1-5 cm. longum et 1 cm. latum, margine densius ciliato, attenuata, folia caulina ovato-ovalia sessilia. Floves virescentes, brunneo- striati, nutantes ; calycis tubus perbrevis, lobi ovati, acuti vel subobtusi, ad 10 mm. longi et 5—7 mm. lati, margine integri ; corollae tubus vix 2 mm. longus, lobi cuneato- obovati, apice rotundati et minutissime erosulati, ad 12 mm. longi et 5~7 mm. lati, nectario nudo subrotundato ; stamina tubo 1-5 mm. alte adnata, filamentis liberis 5 mm. longis, antheris 2:3 x 1-7 mm. magnis ; ovarium anguste ovatum, stigmatibus subsessilibus orbicularibus ; capsula, semina non visa, ovulis semievolutis, ut videtur, subglobosis exalatis.
BuuTAN : Tsampa (Tolegang), Pangotang, 4,400 m. on open hillside, locally common, corolla green striated red brown, 3 July 1949, Ludlow, Sherriff & Hicks 19323 (holotype).
HIMALAYAN SWERTIA AND VERATRILLA 245
The following two collections are closely allied to Swertia virescens : Tibet, Chick- char, Tsari, 4,600 m., on open grassy hillside above rhododendrons, pale cream green, filaments dull green, anthers dark brown, stigma pale green, 13 June 1936, Ludlow & Sherriff 2149 ; Bhutan, Tsampa, Pangotang, 4,300 m., in open wet stony place,
d
Fic. 3. Swertia pseudo-hookeri H. Smith : a, corolla lobe with nectary and stamen ; b, seed - (Ludlow, Sherriff & Hicks 19414). Swertia virescens H. Smith : c, corolla lobe with nec- tary and stamen ; d, middle part of rosular petiole (Ludlow, Sherriff & Hicks, 19323). All x 5.
corolla white with striations inside, 16 June 1949, Ludlow, Sherriff & Hicks 19168. These are both low-growing but very robust plants that are just beginning to flower ; their flowers are bigger than those of Swertia virescens and differ in colour, and the
246 HIMALAYAN SWERTIA AND VERATRILLA
striation of the petals is hardly noticeable. They probably represent an undescribed species which cannot be published as new on the unsatisfactory material yet available.
Swertia pseudo-hookeri H. Smith, sp. nov. (Plate 27 ; Text-fig. 3a—b.)
Swertia pseudo-hookert H. Smith ex Nilsson, Grana Palyn. 7 : 111 (1967), nom. nud.
= “pp @ eaten
Planta 30-40 cm. alta, caulis teres, inflorescentia laxa e nodis 3-4 anguste pyra-
midalis. Habitu S. vivescentt H. Smith subsimilis sed nectario longe nigro-fimbriato (nec nudo) inter alia distincta.
Folia rosularia ad 12 cm. longa, lamina elliptica subacuta c. 5 cm. longa et 1-5 cm. lata, margine integra, in petiolum latum margine glabrum laminam longiorem vel breviorem sensim attenuata, folia caulina sessilia, sursum decrescentia, in nodis paria. Flores albo-virides marginem versus purpureo-striati, 1-5 cm. longe pedicellati,
pedicellis glabris obsolete alatis ; calycis tubus brevissimus, lobi subaequimagni ovati —
obtusi, c. 10 mm. longi et 5~7 mm. lati, margine integri ; corollae tubus perbrevis, lobi obcuneati, apice truncati et parum erosulati, 1-5 cm. longi et 0-9 cm. lati, nectario ovali c. 2 mm. longe nigro-fimbriato, basi non clauso ; stamina tubo adnata, filamentis liberis 8 mm. longis et thecis 3:0x 1-6 mm. magnis ; ovarium ovatum,
stigmatibus parvis subsessilibus ; capsula (e Ludlow, Sherriff & Hicks 19801) angusta, —
I°5 cm. longa, carpellis apice excurvantibus, seminibus ovato-globosis exalatis c. I'4xX 1-2 mm. magnis.
BuuTAan : Tsampa, Marlung, 4,400 m., on open grassy hillside, corolla pale green, finely striated purple, 12 July 1949, Ludlow, Sherriff & Hicks 19414 (holotype). Upper Mangde Chu, Saga La, 4,700 m., 14 July 1949, ravines in alpine zone, herb 60 cm. or more in height, calyx pale green, corolla greenish white finely striated purple at edges of segments, filaments purplish, anthers black, ovary and style greenish blue, 14 July 1949, Ludlow, Sherriff & Hicks 16817. Mangde Chu, Nam- dating, 4,300 m., on steep open hillside, flowers over, 3 Oct. 1949, Ludlow, Sherriff & Hicks rg80r. Pumthang, Tibdeh La, 4,000 m., on peaty slopes, showy pyramid of white and pale blue flowers, 1 Sept. 1915, R. E. Cooper 4769.
Swertia staintonii H. Smith, sp. nov. (Plate 28 ; Text-fig. 4.)
Swertia staintonii H. Smith ex Nilsson in Grana Palyn. 7 : 112 (1967), nom. nud.
Planta c. 30 cm. alta, inflorescentia pyramidalis e basi et nodis 3-4 ramosa, ramis ,
et ramulis suberectis. Species S. crossolomae H. Smith subsimilis sed distat : caule, praesertim inferiori parte, alte 5-sulcato (nec subtereti), floribus majoribus, nectario nudo (nec fimbriato), connectivo antherarum subplano (nec protracto, saepe unci- nato).
Caulis, rami et pedicelli ciliato-alati, alis caulinis in sulcis nec in angulis locatis. Folia rosularia numerosa, subacuta, 6-10 cm. longa et 1-2 cm. lata, lamina in margine dense lacinulata, lacinulis robuste piliformibus 0-3-0-4 mm. longis, in petiolum subaequilongum sensim angustata, petiolo in margine laxe ciliato ; folia caulina infimi nodi 5-verticillata, subacuta, lanceolata, c. 7x 2 cm. magna, breviter petiolata. Flores rosei (e collectore) 1-2 cm. longe pedicellati ; calycis tubus
HIMALAYAN SWERTIA AND VERATRILLA 247
perbrevis, lobi ovato-triangulares, acuti, dense lacinulati, 1-5 cm. longi et I cm. lati, marginibus basalibus in alas pedicelli decurrentibus ; corollae tubus vix 2 mm. longus, lobi obcuneati margine fere integri, ad 2 cm. longi et 1-7 cm. lati, nectario nudo basi rotundato apice subacuto ; stamina tubo 1-5 mm. alte adnata, filamentis liberis r cm. longis, antheris 3x2 mm. magnis minutissime apiculatis ; ovarium anguste ovatum, stigmatibus orbicularibus subsessilibus ; semina (ex Cooper 1994 et 2990) subglobosa, exalata, I-5—1I-9 x I-2-I-4 mm. magna.
SUMO IY Reon Vw Wherry
ee sia Wy
Fic. 4. Swertia staintonit H. Smith : a, corolla lobe with nectary and stamen ; b, margin of leaf ; c, the same, with chlorophyllaceous area dotted (Stainton 755) ; d, cross section of lower part of stem (Cooper 1994). a,b X5,c X50,d x2}.
NeEpaL : Arun Valley, Chhoyang Khola, west of Num, 3,800 m., on open slopes, corolla pink, filaments black, anthers white, 23 June 1956, Stainton 755 (holotype).
BuuTAN : Parshong Timpu, 4,000 m., on peaty gravel, common in this one place,
flowers white, 27 July 1914, R. E. Cooper 1994(E). Same loc., in alpine turf, pyramid of white flowers, 25 Sept. 1914, R. E. Cooper 2990(E).
I have no doubt that Cooper’s two gatherings belong to the same species as Stainton’s. All three have exactly the same unusual morphological organization.
248 HIMALAYAN SWERTIA AND VERATRILLA
The difference in colour is of little account. The Cooper specimens are both in a far advanced fruiting condition ; the flowers are shrivelled, the leaves withered or gone, the capsules mostly empty, but the important characteristics are still there. The peculiar arrangement with the deeply groved stem and the 5-verticillate lower cauline leaves is exactly the same as in the Stainton plant. So are also the shape and margin of the leaves and sepals and the size and form of the petals and nectaries.
Cooper’s notes on the labels “‘ pyramid of white flowers ’’ and “‘ common in this one place’ show that he was well acquainted with the plant. It is certainly to be supposed that he also collected flowering specimens, but no such specimen ever reached Edinburgh. Possibly some packages of his got lost on their way during the troubled year of 1915.
Swertia assamensis H. Smith, sp. nov. (Plate 29.)
Planta robusta c. 30 cm. alta, habitu S. hookert C. B. Clarke subsimilis, sed inter alia distat : floribus majoribus, marginibus foliorum sepalorumque dense lacinulatis (nec integris).
Caulis glaber modeste alatus. Folia omnia et sepala lacinulata, lacinulis pili- formibus vix 0-5 mm. longis ; folia rosularia ad 9 cm. longa, lamina elliptica sub- obtusa 2-2°5 cm. lata, in petiolum subaequilongum sensim attenuata, petiolo margine sparse ciliata. Flores in cymis 3 distantibus, raro breviter ramosis, dispositi, subpatuli, magni, c. 1 cm. longe pedicellati, pedicellis quadrialatis glabris ; calycis lobi duo majores triangulari-ovati ad 13 mm. longi et basi 12 mm. lati, duo minores angustiores et breviores ; petala late obcuneata ad 2-5 cm. longa et apicem versus 2 cm. lata, margine minutissime lacerulata, nectario singulo, nudo, margine basali membranaceo modice elevato ; stamina tubo brevissimo adnata, filamentis liberis 10 mm. longis, thecis robustis 4 mm. longis ; stigmata subsessilia ; capsula, semina non visa.
AssAM : Poshing La, 4,000 m., on cliff ledges and turf slopes, “ (= 13797)”, 22 July 1938, Kingdon-Ward r4o008 (holotype). Bhutan frontier, Orka La, 3,400- 4,000 m., abundant in alpine pastures where cattle feed, comes up in colonies where scrub is cleared, height 25-35 cm., flowers yellow, streaked red, 7 June 1938, Kingdon-Ward 13797.
SWERTIA CANDELABRUM H. Smith in Notes Roy. Bot. Gard. Edinb., 26 : 255, pl. 34, fig. 8 d—-f (1965).
A specimen which may belong to this species is : Bhutan, Sakden, Nyuksang La, 3,700 m., open hillside among rhododendron scrub, pale greenish yellow streaked pale reddish purple, 6 July 1934, Ludlow & Sherriff 601. It differs from the type of S. candelabrum, which comes from central Nepal, in the colour of its flowers, whitish blue in the type, and in the membraneous margin of the nectary, which is less lacini- ate. In other respects it is very similar ; its water-swelled ovules are ellipsoidal and not winged.
> EE
HIMALAYAN SWERTIA AND VERATRILLA 249
Swertia crossoloma H. Smith, sp. nov. (Plate 30 ; Text-fig. 5.)
Planta florendi tempore 22 cm. alta, S. staintonii H. Smith subsimilis sed distat : caule subtereti (nec alte sulcato), nectario fimbriato (nec nudo), connectivo anther- arum protracto, saepe uncinatim curvato (nec subplano), floribus minoribus.
Cc
Fic. 5. Swertia crossoloma H. Smith : a, corolla lobe with nectary and stamen ; b, outline of calyx lobe ; c, margin of leaf (Ludlow & Sherriff 421). All x5.
Caulis subteres, basin versus 4-alatus, alis ciliatis ; inflorescentia pyramidalis e basi et nodis 3 ramosa, ramis, ramulis et pedicellis ciliato-alatis. Folia rosularia ad II cm. longa, lamina late lanceolata, acuta, ad 2 cm. lata, in petiolum subaequilongum sensim attenuata, margine crebre lacinulata, lacinulis piliformibus 0-3—-0-4 mm. longis. Flores pallide albo-virescentes (e collectore) ; calycis lobi subaequales, c. 7 mm. longi et 3-5 mm. lati, ovato-triangulares, subobtusi vel acutiusculi, margine irregulariter lacinulati, basin versus etiam ciliati, marginibus basalibus in alas pedicelli decurrenti- bus ; corollae lobi subobovales ad 1-9 cm. longi et 1-0 cm. lati, superiori parte minute, deorsum grossius lacerulati, nectario orbiculari circum longe fimbriato ; stamina tubo adnata, filamentis liberis 10 mm. longis, thecis c. 3 x 2 mm. magnis, connectivo antherarum protracto saepe uncinatim curvato. Capsula, semina non visa.
250 HIMALAYAN SWERTIA AND VERATRILLA
BuutANn : Me La, 4,300 m., open grassy hillside, pale creamy green, striped blue at base, 6 Aug. 1933, Ludlow & Sherriff 421 (holotype).
Swertia splendens H. Smith, sp. nov. (Plate 31; Text-fig. 6.)
Planta plus quam 40 cm. alta (parte basali deficiente), ex affinitate S. crossolomae H. Smith, a qua distat : planta multo majore, inflorescentia laxiore, floribus lateritiis, 2°5 cm. longis (nec virescentibus, 1-7 cm. longis), corollae lobis late obovatis sub- truncatis (nec ovalibus), connectivo antherarum abbreviato (nec uncinatim pro- tracto), fimbriis nectarii inflexis 1 mm. longis (nec erectis 2 mm. longis).
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Fic. 6. Swertia splendens H. Smith : a, outline of calyx lobe ; b, base of corolla lobe with nectary ; c, anther ; d, ovary ; e, margin of leaf (Ludlow & Sherriff 654). All X5.
Caulis (pars florifera solum adest) robustus, obsolete quadri-alatus, alis sparsissime sed robuste ciliatis ; nodi3 inflorescentiae pyramidalis distantes, rami infimi arcuatim adscendentes 2—5-flori, in nodo supremo flores plures (ad 10) cymosi. Folia rosularia (duo adsunt) spathulata, 7-nervia, lamina obovata, subacuta, 8 et 12 cm. longa, ¢. 5°5 cm. lata, margine crebre lacinulata, lacinulis piliformibus c. 0-3 mm. longis, in petiolum latum, subaequilongum attenuata, petiolo in margine sparse, basi intus
HIMALAYAN SWERTIA AND VERATRILLA 251
densius ciliato. Flores ampli lateritii 1-3-5 cm. longe pedicellati, pedicellis quadri- alatis glabris ; calycis tubus perbrevis, lobi triangulari-cordati, majores ad 1-6 cm. longi, 1-0 cm. lati, margine minute lacinulati basin versus etiam ciliati ; corollae tubus brevis, lobi late obovati, 2-3 cm. longi et 2:0 cm. lati, margine apicaliter minutissime lacerulati, nectario orbiculari fimbriis inflexis 1 mm. longis cincto ; stamina tubo 2 mm. alte adnata, filamentis liberis 11 mm. longis, thecis 3-5 mm. longis, apicibus thecarum connectivo abbreviato liberis ; ovarium ovatum, stylo cum stigmatibus parvis 2 mm. longo ; capsula, semina non visa, ovula, ut videtur, subglobosa exalata.
T1BET : Milakatong La, near Tawang, 4,000 m., open stony hillside, brick red, 14 July 1934, Ludlow & Sherriff 654 (holotype).
sectio OPHELIA
Swertia sect. Ophelia (D. Don ex G. Don) Gilg in Engler & Prantl, Naturl. Planzenfam. 4 (2) : 88 (1895).
Ophelia D. Don ex G. Don, Gen. Syst. 4 : 178 (1837).
Swertia series Ophelia (D. Don ex G. Don) Benth. & Hook., Gen. Pl. 2 : 817 (1876).
Swertia subgen. Ophelia (D. Don ex G. Don) C. B. Clarke in Hook. f., Fl. Brit. Ind. 4 : 121 (1883).
Swertia franchetiana H. Smith, nom. nov. (Plate 32.)
Swertia stricta Franchet in Bull. Soc. Bot. France 46 : 322 (1900) ; non S. stricta Collett & Hemsl. in Journ. Linn. Soc. Lond., Bot. 28 : 91 (1890).
TrBET : Hills north of Lhasa, 3,800 m., on open sandy banks, flowers pale blue violet striated darker, 31 Aug. 1943, Ludlow & Sherriff 9907. Lhakang, 3,700 m., open hillside, very pale blue, veins mauve, 2 Sept. 1933, Ludlow & Sherriff 518. Kongbo, Lhatra Chu (Pangkar), 4,100 m., on dry ground, calyx green, corolla bluish white, filaments blue, anthers very dark brown, style green, stigma very dark brown, 7 Sept. 1947, Ludlow, Sherriff & Elliot 15694. Kongbo, Temo La, 3,200-3,400 m.,
| On grassy banks in forest, calyx green, petals pale watery blue with dark blue veins, | two lateral lines of purple hairs at base, stamens pale blue, anthers purple, ovary | light green at base, darker above, 7 Sept. 1938, Ludlow, Sherriff & Taylor 6992. | Kongbo, Tsangpo Valley, Pe, 2g00 m., in grass by streams, calyx green, corolla slaty
blue veins darker, 23 Aug. 1947, Ludlow, Sherriff & Elliot 14436.
Franchet described this species from western China ; it has not previously been recorded outside China proper.
Swertia gracilescens H. Smith, sp. nov. (Plate 33.)
Planta hapaxantha, S. paniculatae Wall. affinis, sed gracilior, flores minores, emaculati, eleganter dispersi (nec in cymas modice contracti), folia angustiora, rarissime ad 3 mm. lata.
252 HIMALAYAN SWERTIA AND VERATRILLA
Caulis erectus ad 35 cm. altus, basin versus ad 2 mm. crassus, in lineis decurrentibus — brevissime ciliolatus vel glabrescens, e nodis 10-13 pyramidatim ramosus, ramis — numerosis, gracilibus, inferioribus maximis ad 10 cm. longis et 0:5 mm. crassis, suberectis vel subpatentibus, sursum gradatim deminuta. Folia caulina lanceolato- linearia, uninervia, media maxima ad 2:5 cm. longa, circ. 2 mm. lata, marginibus glabra. Flores albi, emaculati, 5-6 mm. longi, 5-meri, uninectariati, ¢racillime 4-6 mm. longe pedicellati ; calycis tubus perbrevis, lobi attenuato-lineares, 4-5-5 mm. longi, basi 0-5-1 mm. lati, emucronati ; corollae tubus minus quam I mm. longus, lobi lanceolato-ovati, acuti, ad 5:5 mm. longi et 4 mm. lati, nectario glabro, orbicu- lato, margine superiori elevato ; stamina tubo 0-7 mm. alte adnata, filamentis liberis non dilatatis 3:5 mm. longis, thecis 0-8 x 0-3 mm. magnis ; ovarium anguste ovoideum in stylum 1-5 mm. longum contractum, stigmatibus vix 0-5 mm. longis. Capsula, semina non visa.
Fic. 7. Swertia burmanica H. Smith : corolla lobe with nectary and stamen. xX 5.
NEPAL : Bhuji Khola, 2,100 m., open slope by track, flowers white, 15 Oct. 1954, Stainton, Sykes & Williams 9049 (holotype). Near Gurjakhani, 2,600 m., exposed grass slope, flowers white, 17 Sept. 1954, Stainton, Sykes & Williams 4470. Kali Gandaki Valley, Taglung, south of Tukucha, 3,700 m., on grass slopes, petals and filaments white, anthers black, 22 Sept. 1954, Stainton, Sykes & Williams 7970.
;
HIMALAYAN SWERTIA AND VERATRILLA 253
Swertia burmanica H. Smith, sp. nov. (Plate 34 ; Text-fig. 7.) Swertia burmanica H. Smith ex Nilsson in Grana Palyn. 7 : 110 (1967), nom. nud.
Planta hapaxantha Swertia racemosae (Griseb.) C. B. Clarke affinis et habitu similis, tamen distat : foliis subcoriaceis, lineari-lanceolatis, media parte (nec basin versus) latissimis, ad 2:5 cm. longis et 0-5—0-7 cm. latis, pagina inferiori conspicue flavescente ; floribus amplioribus ad 1-5 cm. longis, calyce turbinato (nec campanulato), tubo calycis breviore, I-5-2-5 mm. longo (nec 2-5 mm.) ; corolla fere ad basin filamentorum partita, lobis late ovalibus, acutis, minute mucronulatis, margine obsolete micro- ciliatis, ad 13 mm. longis et 10 mm. latis, tubo 5-plo longioribus (nec sublongioribus), indusio nectarli robusto, squamiforme, margine libero partim incurvante breviter laciniato, laciniis ecoloratis (nec tenero, magis diviso, laciniis vulgo nigrescentibus) ; filamentis deorsum valde incrassatis, basi approximatis sed non connatis (nec basi fere 1 mm. longe tubiforme connatis).
Burma : Mt. Victoria Ridge, 2,600 m., a gregarious weed growing all along the ridge and on turf slopes in thousands, flowers 5-petalled, purple, 26 Oct. 1956, Kingdon-Ward 22747 (holotype).
On the type sheet there are 3 individuals present. Two of them are more or less densely ciliate on the wings of the stem, on the surface and base of the under-side of the leaves, and on the calyx. The third specimen is nearly glabrous except on the
wings of the stem. The degree of hairiness seems to be of small taxonomic import-
ance, as is also the case in S. racemosa (below).
SWERTIA RACEMOSA (Griseb.) C. B. Clarke in Hook. f., Fl. Brit. Ind. 4 : 124 (1883).
Swertia racemosa Wall., Numer. List : 154, n. 4377 (1831), nom. nud.
Ophelia racemosa Griseb., Gen. & Sp. Gentian. : 319 (1838).
Kingdon-Wardia codonopsidoides Marquand in Journ. Linn. Soc. Lond., Bot. 48 : 207 (1929) ; in Hook. Icon. Pl. 32 : t. 3163 (1932).
T1BET : Kongbo, Pa La (Tra La), 3,700 m., in colonies in damp shady situations, flowers pale dull purple, 26 Sept. 1924, Kingdon-Ward 6205 (isotype of Kingdon-Wardia codonopsidoides). Kongbo, Bo La, 4,300 m., under bushes in light forest, calyx light green, corolla purplish white, filaments pale blue, anthers black, 9 Sept. 1947, Ludlow, Sherriff & Elliot 15705. Kongbo, Nambu La, 4,400 m., on grassy banks in juniper forest, corolla pinkish mauve, 31 Aug. 1938, Ludlow, Sherriff & Taylor 6951. Tsari valley, 4,000-4,300 m., on alpine meadow slopes, 11 Sept. 1935, King- don-Ward 12324. Tsari, Podzo Sumdo, 3,500 m., by the side of a stream in dense forest, corolla palest mauve, 5 Sept. 1936, Ludlow & Sherriff 2544.
NEPAL : Deoli Patan, S.E. of Jumla, 3,800 m., growing on open slopes, corolla mauve, stamens blue, 5 Sept. 1952, Polunin, Sykes & Williams 3189. Rohagaon, Suli Gad, 3,400 m., open slopes, corolla mauve, nectaries at base of lobes green with fringe of dark blue hairs, filaments blue at apex, anthers brown, style blue, 14 Sept. 1952, Polunin, Sykes & Williams 3386. Near Seng Khola, 4,100 m., grass gullies, flowers pale mauve, 3 Oct. 1954, Stainton, Sykes & Williams 4688. Near Phagune
254 HIMALAYAN SWERTIA AND VERATRILLA
Dhuri, 3,500 m., sheltered gully, flowers lilac, 12 Oct. 1954, Stainton, Sykes & Williams 4799. Above Dogadi Khola, 4,600 m., exposed earth pockets near ridge, flowers mauve, calyx and pedicels dark purple, 12 Aug. 1954, Stainton, Sykes & Williams 3834. Same loc., 4,000 m., among dwarf Rhododendron, flowers light purple, 1 Oct. 1954, Stainton, Sykes & Williams 4661. Dhorpatan, 2,900 m., among shrubs on bank, flowers mauve, 16 Oct. 1954, Stainton, Sykes & Williams 9056. Above Sauwala Khola, 3,400 m., grass slopes on ridge, flowers mauve, 13 Sept. 1954, Stainton, Sykes & Williams 4381. Same loc., 4,000 m., among short grass on ridge top, abundant, flowers mauve, 13 Sept. 1954, Stainton, Sykes & Williams 4389. N.W. of Gurjakhani, 3,500 m., grassy slope, flowers white, 20 Sept. 1954, Stainton, Sykes & Williams 4502. Same loc., 3,700 m., near grazing encampment, abundant, flowers dark mauve, 20 Sept. 1954, Stainton, Sykes & Williams 4512. Kali Gandaki Valley, Samargaon, north of Tukucha, 4,900 m., on steep grass slopes, calyx mauvish green and hairy, corolla, filaments and anthers pale mauve, 16 Aug. 1954, Stainton, Sykes & Williams 7249. Kali Gandaki Valley, Sangdah, north of Tukucha, 4,100 m., open grass slopes, calyx green, corolla, filaments and anthers whitish blue, 18 Aug. 1954, Stainton, Sykes & Williams 7309. Kali Gandaki Valley, Tukucha, 3,700 m., open grass slopes, calyx greenish mauve, corolla and filaments pale mauve, lower leaves reddish green, 9 Sept. 1954, Stainton, Sykes & Williams 7735. Same loc., 3,700 m., open grass slopes, calyx greenish white, corolla and filaments white, anthers black, 9 Sept. 1954, Stainton, Sykes & Williams 7737. Same loc., 3,100 m., open grass slopes, corolla white, 15 Oct. 1954, Stainton, Sykes & Williams 8157. Kali Gandaki Valley, Taglung, south of Tukucha, 3,700 m., open grass slopes, petals and filaments white, anthers black, 20 Sept. 1954, Stainton, Sykes & Williams 7945. Sabze Khola, 4,000 m., among low bushes in moist ground and by stream side, flowers pale lilac or mauve, leaves and stem stained red, 6 Sept. 1950, Lowndes Lr4g1. Khangsar, 4,300—4,600 m., open dryish hillside, in great quantity, flowers pale lilac or mauve, leaves and stem stained red, 9 Sept. 1950, Lowndes L1502. Bimtakothi, 4,000 m., damp hillsides, flowers whitish, 12 Sept. 1950, Lowndes L1539. Baraha Pokhri ridge, 4,000 m., open grassy hillside, flowers greyish white, 19 Sept. 1950, Lowndes L1526. Mardi Khola, Annapurna Himal, 3,500 m., among shrubs, corolla pale blue, 17 Sept. 1954, Stainton, Sykes & Williams 8465. Rambrong, Lamjung Himal, 3,700 m., on open hillside, corolla pale blue, 18 Sept. 1954, Stainton, Sykes © Williams 8631. Kyangjin Ghyang, 4,000 m., grassy ground, pale purple, O. Polunin Tg09. ‘‘Gossain Thar’ [? = Gosainkund], Wallich 4377 (isotype of Swertia racemosa). Gosainkund, 4,900 m., 28 Oct. 1935, F. M. Bailey’s collectors 60. E. of Num, Maghang Khola, Arun valley, 3,500 m., on open slopes, corolla mauve, 15 Sept. 1956, Stainton 1737. Sabel, 4,000 m., 18 Sept. 1937, K. N. Sharma 63/94. Chocho, 3,500 m., 27 Sept. 1937, K. N. Sharma 56/94. Gupchet, 3,700 m., 13 Sept. 1935, F. M. Bailey’s collectors s.n. Chilung Pati, 4,100 m., 25 Oct. 1935, F. M. Bailey's collectors 49. Rolwaling valley, 3,800 m., 28 Sept. 1960, Swan R go.
SIKKIM : Changu, 3,700 m., on loamy and sandy banks, blue, 25 cm. tall, 27 Sept. 1913, Cooper 954. BuutAN : Lingshi, 4,100 m., shrubbery, flowers dull purple, 24 Sept. 1967, Bowes
EL as
HIMALAYAN SWERTIA AND VERATRILLA 255
Lyon 5135. Thimpu, 3,100 m., on dry loam, blue to purple flowers, 25 cm. tall, slender, 5 Nov. 1914, R. E. Cooper 3549. Kopub, 3,400 m., in mould and leaves, etc., at edge of forest, flowers pale blue and white, 24 Sept. 1914, R. E. Cooper 2r14r. Lhabja, 3,800 m., on moist soil under forest, flowers blue, 30 cm. tall, 22 Sept. 1914, R. E. Cooper 2187. Waitang, Tsampa, 4,000 m., in dense vegetation of steep hillside, flowers pale pink with a tinge of blue, up to 40 cm. tall, 19 Sept. 1949, Ludlow, Sherriff & Hicks 19753. Tolegang, Tsampa, 4,300 m., among dwarf rhododendron on the open hillside, corolla pale slaty blue, 10 Sept. 1949, Ludlow, Sherriff & Hicks 19730.
Assam : Near Orka La, 3,700—4,000 m., alpine turf slopes along the ridge, flowers purple, 26 Sept. 1938, Kingdon-Ward 14292.
The genus Kingdon-Wardia is said by its author to be intermediate in position between Gentiana and Swertia, differing from the former by the presence of a fimbriate nectary at the base of each corolla lobe and from the latter by “ the conspicuous calyx tube surmounted by triangular lobes and the well-marked corolla tube.”” The distinction from Gentiana certainly holds true, but the separation from Swertia is not convincing. Marquand was obviously not acquainted with Swertia racemosa, a species little known in 1928 and represented in the Kew herbarium at that date by only the Wallich type specimen and by the specimen from Tibet to which Burkill
had given the manuscript name Swertia codonopsidoides.
The only statement in the diagnosis of Kingdon-Wardia that does not fit Swertia vacemosa is : “ corolla circaad}lobata’’. This statement is, however, not correct ; a more exact one would be : corolla ad 4 vel paullo ultra lobata. The figure subse- quently published by Marquand in Hooker’s Icones shows the lobes and the tube to be of equal length, but the previous incorrect statement is repeated in the text. In the isotypes that I have seen my own measurements show that the lobes are in fact constantly a trifle longer than the tube, exactly as in Swertia racemosa, with which species Kingdon-Wardia codonopsidotdes is undoubtedly conspecific.
Swertia racemosa has a slightly isolated position in the genus, differing from other species of Swertia sect. Ophelia by the less divided corolla and by the well developed calyx tube, but it is not divergent in any other respect. Its affinity with such species as S. ciliata (G. Don) Burtt and S. hispidicalyx Burk. is indicated by the shape of the nectary, stamens and style. It is even closer to S. burmanica, which is very similar in appearance. The calyx and deeply cleft corolla of that species conform to the normal pattern found in S. sect. Ophelia and it thus links S. racemosa with the other members of the section.
Plentiful material of Swertia racemosa has been collected in recent years, especially by the British Museum expeditions to the Himalayas. In that herbarium there are 32 gatherings from Nepal, 1 from Sikkim, 6 from Bhutan, 5 from S.E. Tibet and 1 from Assam. This ample material shows that S. racemosa has an unusually wide range of variability. The corolla lobes vary from distinctly to only slightly longer than the tube ; the calyx varies from 5 to 2 mm. in length, the calyx lobes are usually much longer than the tube and distinctly unequal, but they may also be shorter than the tube and of subequal size, and the angles between the lobes vary from broadly
256 HIMALAYAN SWERTIA AND VERATRILLA
rounded to narrowly acute ; the stigma is sometimes subsessile, but commonly the style is more or less prolonged and may reach a length of 5 mm. ; the petals average in length 11 mm. (5-14) ; the calyx, the margins and upper surface of the leaves vary from densely or sparingly long- or short-ciliate to perfectly glabrous. All these variations seem to be independent of each other, and the extremes are connected by intermediate stages. To name such indistinct forms would be meaningless. It may nevertheless be mentioned that really long-styled specimens are not found outside Nepal, but this may be due to the fact that the plant has been collected over 30 times there and only 13 times elsewhere.
VERATRILLA (Baillon) Franchet
VERATRILLA (Baillon) Franchet in Bull. Soc. Bot. France 46 : 310 (1900). Swertia sect. Veratrilla Baillon in Bull. Mens. Soc. Linn. Paris 1 : 730 (1888).
At a session of the Linnean Society of Paris on 6 April 1888, Baillon demonstrated a gentianaceous plant collected by Delavay in Yunnan. He gave a description, pointing out that it had certain characters in common with Je@eschkea, Obolaria, Bartonia and Gentiana, but concluded “‘. . . Swertia proprement dits, et c’est a ce genre que le Veratrilla, si singulier qu’il puisse paraitre, devra se rapporter comme section.’”’ This must be regarded as valid publication of the name Sweriza sect. Veratrilla. The sectional epithet was chosen on account of the resemblance of this plant to Veratrum L. Baillon proposed no specific name for this plant.
In the following year, Baillon (Hist. Pl. 10 : 142 (1889)) expressed uncertainty about the taxonomic status of the Delavay plant, saying in a footnote to Swertia “aut generis sectio, aut generis proprium erit Verairilla H. Bn... ., cujus flores diceci, seepe 4-meri. . . Species asiaticae multae novae hanc plantam cum Frasera et Swertiis legitimis nihilhominus connectunt.”
Franchet, who had at an earlier date, probably in 1885, given a manuscript name under Swertia to the Delavay specimen in the Paris herbarium, obviously became convinced that it represented a separate genus and published it in rgoo at that rank and, ignoring his earlier manuscript epithet, published the name Veratrilla baillonu for the species. I quote his diagnosis in extenso ; there are two statements in it that should be altered. It seems as if Franchet had by that time lost interest in the plant ; the description is not of the same high class as one is used to from him :
“ Veratrilla, an generis sectio an genus proprium Baillon, Hist. des pl. X, 142 et Bull. Soc. Linn. de Paris, 729.
“Flores diceci, sepius 4-meri ; corolla tubuloso-rotata, staminibus inferne cum tubo connatis juxta sinum tantum liberis, brevibus ; stylus brevis (vix I mm.) stigmate bilobo, lobis ovatis patentibus ; capsula ovata, breviter attenuata ; semina late ovata, levia.—Planta glabra, elata veratriformis ; folia inferiora longe petiolata, omnia opposita, anceolata [sic], inferiora vaginata ; inflorescentia e racemis densis pyramidam longam, angustam efformantibus ; corolla luteoviridis, 6 mm. longa et lata.
“V. Bailloni sp. nov.
HIMALAYAN SWERTIA AND VERATRILLA 257
“‘ Hab.—La Chine occidentale, Yunnan, dans les broussailles, au-dessus du col de Yen-tze-hai, alt. 3500 métres (Delavay) ; prairies humides de Tsang-chan, prés de laréte (Delavay n. 147).
‘“‘ Genre remarquable, a cause de sa dioicité ; les étamines sont insérées comme celle des J@schkea.”’
Baillon said in his original account “‘ ses fleurs sont ordinairement tétraméres, plus rarement pentaméres, principalement dans le pied femelle’ and in the Histoire des Plantes ‘‘ flores. . . saepe 4-meri”’, and Franchet said “ flores. . . saepius 4-meri ”’, but I have examined 16 different collections and have never found a single flower that was not tetramerous. Kingdon-Ward, however, recorded pentamerous flowers in a field note (see below).
Franchet’s description of the seeds reads ‘‘semina late ovata, levia’’. A more accurate one would be : semina complanata, ambitu subovalia, circum anguste alata c. 3X1‘5-2 mm. magna, ala 0-3-0:5 mm. lata.
The great difference in appearance between the male and female plants must also be pointed out. The female plant has the long narrow pyramid of Franchet’s description ; each bract supports one or two short upright branches bearing fewer than ten pedicelled flowers. The inflorescence of the male plant is robust and spreading, several hundreds of small flowers without pedicels being massed together on the stout and more or less patent branches. This difference is well demonstrated by the specimens of this species from China that are illustrated as plates 35-36. Those from Tibet or Assam are not so satisfactory for photography.
VERATRILLA BAILLONII Franchet in Bull. Soc. Bot. France 46 : 311 (1900) (‘‘ bailloni”’). (Plates 35, 36.)
Swertia mekongensis Balf. f. & Forr. in Notes R. Bot. Gard. Edinb. 4 : 80, t. 19 (1907).
S.E. TiBet : Mt. Kenichunpo, Salween and Irrawaddy divide, 4,600 m., limestone crags, alpine region, plant fleshy, flowers yellow, May—July 1932, Rock 21938.
Assam : Orka La, Bhutan frontier, 4,000 m., pastures, flowers 4-merous, sometimes 5-merous, Kingdon-Ward 13725.
Veratrilla batllonii had not previously been recorded outside China proper, the identity of Swertia mekongensis with it not having been recognized. It is now recorded for S.E. Tibet and Assam.
Veratrilla burkilliana (W. W. Smith) H. Smith comb. nov. (Plate 37.)
Swertia burkilliana W. W. Smith in Journ. Asiat. Soc. Bengal, new ser. 7 : 78, t. 1, fig. i-iii (1911). Veratrilla burkilliana H. Smith apud Nilsson in Grana Palyn. 7 : 113 (1967), nom. nud. S.E. Tiset: Tsari, Tomtsang, 4,300 m., on open grassy hillside, corolla dull purple margined with green and slightly streaked with greenish purple, anthers blue | purple, 23 June 1936, Ludlow & Sherriff 2204. Same loc., 4,300 m., on open grassy hillside, flower over, 13 Sept. 1936, Ludlow & Sherriff 2578. Chubumbu La, Langong,
B
258 HIMALAYAN SWERTIA AND VERATRILLA
4,000 m., common in open swampy meadows, corolla pale green, tinged dull slate blue at base, 2 June 1938, Ludlow, Sherriff & Taylor 3943.
SIKKIM : Changu, 3,700 m., in turf on lake side, green flowers, black marks, blue stamens, I July 1913, R. E. Cooper g2. Same loc., 3,700 m., on lake border, greenish flowers with spots, 20 Aug. 1913, R. E. Cooper 565.
BHUTAN : Pung La, 3,700 m., open marshy places near top of pass, flowers blue marked with green, 9 July 1949, Ludlow, Sherriff & Hicks 20903.
This second species of Veratrilla agrees exactly in all generic characteristics with V. baillonii, but it is quite distinct specifically. The plant is much smaller and less robust, only 15-30 cm. tall as against 30-50 cm. The cauline leaves are shorter and relatively broader. The seeds are slightly smaller, nearly circular in outline and with a slightly broader wing. The difference in appearance between male and female plants is strongly marked in this species also. The excellent figure accompanying the original description of the species represents the female plant. In the male the inflorescence is spreading, up to 4 cm. wide by 6 cm. long.
Dr, H. SmitH
INSTITUTE OF SYSTEMATIC BOTANY OF THE UNIVERSITY OF UPPSALA BOTANICAL GARDEN AND BOTANICAL MUSEUM
UPPSALA I
SWEDEN
Bull. Br. Mus. nat. Hist. (Bot.) 4, 6 PLATE 22
-
i $ z 3 = é FLORA OF “HiT EAL Loe. Alt. Date Gm bacieay bei Qeoek aie 1 {kage taw yw . Sus d Corraay tinted. 3. af ides rt ehlea. iki, ore : pial " planale Adod " iL. Swertia kiharae Kitamura me ve Coll. J. D. A. STAINTON No.2708
Swertia kihavae Kitam. (Stainton 2708 in Herb. Brit. Mus.)
Bull, Br. Mus. nat. Hist. (Bot.) 4, 6
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PLATE 23
FLORA OF i A oN 1437 j . atc. f ’ Lat i ix. ’ Vis ws ar 7 2 ae at) 1° one
Lhe OE Remo ee =a
Swertia hookeri C. B. Clarke (Ludlow,
Swertia hooker! C.B, Clarke
Sherriff & Elliot 14376 in Herb. Brit. Mus.)
Bull. Br. Mus. nat. Hist. (Bot.) 4, 6 PLATE 24
ly
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Bull. By. Mus. nat. Hist. (Bot.) 4, 6
1 gl
71s Muswum (Naroral History) : Oger ~ | Laduuladula daca
HOLOTY PE Swertia grandiflora H. Smith
weet
Swertia grandiflora H. Smith (Ludlow,
PLATE
a
FLORA OF [3
Coll. &. Lyupiow, G. Sueraire & J. H, Hice
Sherriff & Hicks 20732 ; holotype in Herb. Brit. Mus.)
2
5
Bull. Br. Mus. nat. Hist. (Bot.) 4, 6
|! i | T 4| vy TT ih 2 1 be Miter)
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HOLOTY PE Swertia virescens H. Smith
Swertia virescens H. Smith (Ludlow, :
PEATE
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Determ H. SMITH, Upsate 1960.
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Sherriff & Hicks 19323 ;
holotype in Herb. Brit. Mus.)
20
Bull. Br. Mus. nat. Hist. (Bot.) 4, © PEATE
FLORA OF BHUTHR Ne 19414
Ai 14 Soe: Date
A 7 Ag boot 1 4 es oe oe) A Doe ay ty. Manat WS ? Pa Call Lug wv SiHeReire & J. Wicks _ Typus HOLOTY PE
Swertia pseudo-hookeri H. Smith
ig SR At BES Bd Ea i
fers Museum (Narueal Histomy «
Protograrti | {| | ‘ | | | 3
Swertia pseudo-hookeri H. Smith (Ludlow, Sherriff & Hicks 19414 ; holotype in Herb. Brit. Mus.)
aah
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Bull. Br. Mus. nat. Hist. (Bot.) 4, 6
rT 7 | i 2 fir fs Museum (Natvea Heerosy ‘ ft
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HOLOTY PE Swertia staintonii H. Smith
Swertia staintonit H. Smith (Stainton
FLORA OF EASTERN NEPAL
Alt Date
Coll. J. D. A. STAINTON No. 7
See
FOI
holotype in Herb. Brit. Mus.)
PLATE
28
Bull. Br. Mus. nat. Hist. (Bot.) 4, 6 PLATE 29
HOLOTY PE a it is H. Smit
eh fac al aie We , é env Msn Narn. Miron P nies
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Swertia assamensis H. Smith (Kingdon-Ward 14008 ; holotype in Herb. Brit. Mus. The printed number on the ticket has been deleted in blue ink and 14008 substituted, but this has not reproduced on the photograph.)
Bull. Br. Mus. nat. Hist. (Bot.) 4, 6
Pyar
i
| i 2 atisn Museum (Narorat History PHOTOGRAPH i 1 2 | | tun wuluislis
HOLOTY PE Swertia crossoloma H. Smith
Swertia crossoloma H. Smith (Ludlow & Sherriff 421 ; holotype
1 Gal os a Bel Dome Yo
in Herb. Brit. Mus.)
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Bull. Br. Mus. nat. Hist. (Bot.) 4, 6 Piers
og Ty abs 12 t ‘s
1 2 (Barris Museum (NaTurAt History) PHOTOGRAPH
oe Sa ae ae i
Q HOLOTY PE
Swertia splendens H. Smith
F. LUDLOW—G. SHERRIFt
CAT ONG L » Locality MAW AT ENE LA No §54
New “Tawarc 0 + Alt. ece 82 Te Date is ies Description
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Swertia splendens H. Smith (Ludlow & Sherriff 654 ; holotype in Herb. Brit. Mus.)
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Bull. Br. Mus. nat. Hist. (Bot.) 4, 6
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Description
Very parks bmw, Vewin wane.
A cinclilh carve aL A . Swertia franchetiana H. Smith ) & "Satna pina ome, ene ont mnt ee and nal eal saat et telat gee
Swertia franchetiana H. Smith (Ludlow & Sherriff 518 in Herb. Brit. Mus.)
32
Bull. Br. Mus. nat. Hist. (Bot.) 4, 6 Pie Ag E33
1.4414.)
Detorm. H, SMITH, Dpsate 950
WX Mandin Softee 1 Ubiona
A HOLOTYPE -_ > —_— Swertia gracilescens i A. Smith ees Bi
: FLORA OF No FO.G. } hana Khelo, alt. ], O06 1G Pate 15} 10) 4454. Lat, : Long. r) cc“ ' , } . yaw she it ba, Cech, ' nag
4 q ' Theuats hey H ' ' '
Swertia gracilescens H. Smith (Stainton, Sykes & Williams 9049 ; holotype in Herb. Brit. Mus.)
Bull. Br. Mus. nat. Hist. (Bot.) 4, 6 PLATE 34
HOLOTY PE Swertia burmanica H. Smith
22747
Swertia burmanica H. Smith (Kingdon-Ward 22747 ; holotype in Herb. Brit. Mus.)
Bull. Br. Mus. nat. Hist. (Bot.) 4, 6
STITTITITTTT rrr yg } 1 j sa Museum (NATURAL Hirsroey): PHOTOGRAPH
i | | J |
Veratrilla baillonit Franchet
,
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irk OG AG
ORY F
Veratrilla baillonit Franchet, female plant (Yw 735r4 in Herb. Brit. Mus.)
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| | | |
dw) BA
JERA OPLONIA AND
)
(ACANTHACEAE
NTH
W. T. STEARN
OF
ye
ATURAL HISTORY Vol. 4 No.
N
A SURVEY OF THE TROPICAL GENERA PLONIA AND PSILANTHELE (ACANTHACEAE)
BY WILLIAM THOMAS STEARN
——— xref
\
Pp. 259-323 ; 18 Text-figures ; Plates 38-47
BULLETIN OF __ THE BRITISH MUSEUM (NATURAL HISTORY) BOTANY Vol. 4 No. 7 , LONDON : 1971
THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), «stituted in 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.
In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department.
This paper is Vol. 4, No.7 of the Botanical serves.
World List abbreviation Bull. Br. Mus. nat. Hist. (Bot.).
© Trustees of the British Museum (Natural History), 1971
TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 10 August 1971
Price £2.60 |
ee
A SURVEY OF THE TROPICAL GENERA OPLONIA AND PSILANTHELE (ACANTHACEAE)
By WILLIAM T. STEARN
SUMMARY
On grounds of priority the generic name Oplonia Raf. (1838) replaces Anthacanthus Nees _ (1847) ; both have the same type, Justicia spinosa Jacq. The genus Psilanthele Lindau (1897) is restricted to its type-species, P. eggersit Lindau of Ecuador ; the Peruvian and Jamaican species put in Psilanthele by Lindau are transferred to Oplonia, the supposed distinction between the ‘“‘ Rahmenpollen ”’ of the one and “‘ Spangenpollen ”’ of the other being illusory. Forsythi- _opsis Baker (1883), hitherto considered endemic to Madagascar, is here united with Oplonia, as they agree in habit, floral structure, capsule, and pollen. The genus Oplonia thus extended consists of one species in Peru, eight in the West Indies, none in Central America, and five in Madagascar, which, like Cuba and Jamaica, is a major centre of endemism. Recognition of taxa within the group is very difficult owing to extensive but seemingly uncorrelated variation in habit of growth, pubescence of shoots, and size and shape of leaves. The spines provide useful diagnostic characters. Floral dimorphism, i.e. the occurrence of long-stamened short-styled flowers and of short-stamened long-styled flowers on different individuals of the same taxon in the “same district, is demonstrated, possibly for the first time in the Acanthaceae ; otherwise the _ genus manifests very little floral diversity. Most species grow in dry open habitats as part of _ scrub forest. Their evolution would appear to be from a mesophytic spineless South American _ ancestor, resembling the small-flowered species of Odontonema, by progressive reduction in size of leaf, by contraction of the inflorescence from a narrow panicle or raceme into a few-flowered _ axillary fascicle or a solitary axillary flower, and by conversion of lateral branches into spines, by which changes they became xerophytic. The most primitive species on this assumption are f O. grandiflora of Peru and O. jamaicensis of Jamaica. Parallels in distribution with Neriacanthus, é Heppiella, Huerta and Purdiaea indicate ancient floristic links between the Greater Antilles and _horth-western South America independent of Central America and the Lesser Antilles and _ Suggest that a major part of the Greater Antillean flora may be descended from plants of a land- _™ass now sunk which occupied the position of the present Caribbean Sea. The presence of congeneric species in tropical America and Madagascar but not in continental _ Africa, as exemplified by Oplonia, Oliganthes, and Ravenala, is attributed to continental drift and subsequent extinction on the African mainland, as there exist genera with ranges inter- ‘Mediate between the extreme discontinuity of these and the continuous range of, e.g., Sabicea. _ New nomenclatural combinations are: Oplonia acicularis (Justicia acicularis Swartz), O. _armaia (J. armata Swartz), O. grandiflora (Psilanthele grandiflora Lindau), O. linifolia (Forsythi- Opsis linifolia Benoist), O. microphylla (J. microphylla Lam.), O. minor (F. minor Benoist), 0. minor var. meridionalis (F. vincoides var. meridionalis Benoist), O. minor var. vestita (F. vin- Coides var. vestita Benoist), O. nannophylla (Anthacanthus nannophyllus Urban), O. purpurascens (4. purpurascens Griseb.), O. vincoides (J. vincoides Lam.). O. tetrasticha var. polyece from Oriente Province, Cuba, and O. acuminata and O. puberula, both from Madagascar, are described as thew ; Psilanthele minor Lindau is renamed O. armata var. pallidior. Descriptions, synonymy and distributional data are provided. As a guide to the capacity of computer-aided taxonometric (numerical taxonomic) methods to facilitate the sorting of relatively intractable specimens into groups definable and acceptable as taxonomic units, two of these methods (using programmes CLASP and Taximetric Similarity
262 A SURVEY OF THE TROPICAL GENERA
Graph-clustering) were applied to Oplonia specimens providing a matrix of 24 unweighted characters. The results, expressed as stages of clustering, as a dendrogram and as a minimum scanning tree converted into a pictorialised scatter diagram, demonstrated their value as aids to understanding assemblages wherein most distinctions lie in a diversity of minute characters neither simultaneously observable nor easily correlated. The stage-by-stage clustering, with progressive reduction of the degree of similarity necessary for admission of specimens to clusters, portrayed graphically by the Taximetric Similarity Graph-clustering programme, commended itself as a time-saving substitute for part of the process of searching for correlations.
HISTORICAL INTRODUCTION
THE genus Oflonia is a small group of woody much-branched usually spiny small- flowered tropical plants belonging to the family Acanthaceae subfamily Ruelloideae tribe Odontonemateae. The flowers are solitary in the leaf-axils or borne in com- paratively few-flowered axillary fascicles with minute bracts, this much-reduced inflorescence distinguishing Oplonia from Odontonema Nees (Thyrsacanthus Nees), which has terminal racemes or panicles. The calyx, as in Odontonema, is cleft into five narrow segments. The corolla (Text-fig. 1, B) is zygomorphic, usually lilac or red,
Fic. 1. Oplonia armata var. pallidioy Stearn: A, Habit, x 2/3; B, corolla, x 1; C, flower of long-styled form cut lengthwise, x 2 ; D, pollen grain, x 300 ; E, capsule, X I (Harris 10257).
but sometimes white, with a slender tube and a two-lipped limb, the upper lip two- | lobed, the lower lip three-lobed. There are two stamens, which may be exserted or included, and two staminodes. One species, Oplonia grandiflora, grows in northern
OPLONIA AND PSILANTHELE (ACANTHACEAE) 263
Peru ; the others are West Indian or Madagascan. Most of these can be distin- euished from other West Indian members of the Acanthaceae by their opposite axillary spines (Text-fig. 2), which are modified shoots, not modified leaves as are the paired spines of Barvleria and Barleriola. They supply the major taxonomic features of the species and vary from the forbidding rigid four-ranked stiletto-like thorns up to 3 cm. long of O. tetrasticha to the short prickles, sometimes under 4 mm. long, of O. nannophylla. The generic names Opflomia and Anthacanthus and the epithets acicu- laris, armata, polyece and spinosa all refer to this armature.
Although pre-Linnaean collectors gathered a few specimens of Oplonia microphylla in Antigua and Jamaica, the first species to be scientifically recorded was O. spinosa collected by Nicolaus Joseph von Jacquin near Port-au-Prince, Haiti, in 1757-58 and published by him in 1760 as Justicia spinosa (Pl. 43 a). On this species Rafinesque founded the genus O¢lonia in 1838, his characteristically contracted diagnosis being as follows :
“987. OPLONIA R. (weapons) diff. Justicia, cor. hypocraterif, tubo tereto, limbus 5 lobo, bilabiato, lab. sup. bilobo plano. Type Ol. spinosa R. Justicia do. L. auct. Prickly shrub, and probably all the prickly American sp. J. micro- phylla, armata, acicularis &c. ”’
Probably Rafinesque never saw any of the plants concerned, which are all West Indian and were then rare in herbaria. However he never hesitated about founding new genera on the illustrations and descriptions of more cautious authors* and he could have drawn more than enough information for his diagnosis from Vahl’s Enumeratio Plantarum 1 : 167-169 (1804), where J. spinosa, J. microphylla, J. armata and J. acicularis are described. The epithets spinosa, armata and acicularis pre- sumably suggested the generic name Oplonza, derived, like the name Oflopanax (Torr. & Gray) Miq., from the Greek ézAov, “implement of war, weapon, armour ”’ and thus cognate with the word “ hoplite’”’ ; Hoplonia would indeed have been a better rendering. Rafinesque’s Flora Telluriana (Philadelphia, 1837-38) is a very rare work, of which apparently only two copies were available in Europe until the publication in 1946 of a photographic facsimile. Certainly C. G. Nees von Esenbeck knew nothing of it when in 1847 he separated these same prickly species from Justicia and Evanthemum and named the genus Anthacanthus, from av@os “ flower’’ and axavOos, “spine”. Nees distinguished within Anthacanthus a section Armati consisting of the spine-bearing West Indian species (A. spinosus, A. armatus, A. acicularis, A. microphyllus, A. lycioides, A. cuneatus, A. emarginatus and a “‘ species dubia ’’, A. sprengelii, actually a member of the Rubiaceae) and a section Inermes consisting of unarmed species (A. vincoides from Madagascar, A. stnuatus and A. vepandus from the New Hebrides) which could well have been left in Evanthemum
*In his Flora Telluriana 4 : 64, n.986 (1838), Rafinesque with customary recklessness also published
a genus Crateola to include two little-known species, ‘‘ Types the J. vincoides, J. parviflora Ortega non
Retz.’’, of which he had read the brief descriptions in Vahl, Enum. Plant. 1 : 166-167 (1804), and charac-
terized it as ‘“‘diff. Justica, cor. hypocraterif. tubo brevis basi inflato, limbo plano 5 lobo vel 5 partito
equalis”’. As Vahl described the corolla of Justicia vincoides Lam. as “ infundibiliformis ’ ‘(its tube is
in fact narrowly cylindric and not swollen at base) but stated that J. parviflora Ortega had ‘‘corollae
tubus basi ampliatus ’’, I accordingly designate J. parviflora as the lectotype of Crateola Raf. Nees» placed this obscure species in the genus Dipteracanthus Nees (1832), although Hemsley suggested that
it might belong to Hemigraphis Nees (1847). The name Crateola thus typified cannot displace either
Oplonia or Forsythiopsis ; should it compete with Hemigraphis, the latter name should be conserved.
264 A SURVEY OF THE TROPICAL GENERA
(see p. 320) as accepted then.* In 1920 Britton and Millspaugh designated Antha- canthus spinosus (Justicia spinosa Jacq.) as the lectotype of Anthacanthus. Since, however, the genus Anthacanthus as originally circumscribed by Nees included the designated type of Oplonia Raf., the name Anthacanthus was technically a super- fluous name when published and its type has always been Justicia spinosa Jacq. (see Int. Code Bot. Nomencl. Art 7, Note 4.)
Subsequent botanists dealing with the family Acanthaceae have maintained the genus and, with the exception of Otto Kuntze, have adopted the name Anthacanthus for it. In 1839 Endlicher divided the genus Evanthemum into three groups of un- specified rank, namely Euranthemum of the East Indies, Planetanthemum of Africa and Australia and Hesperanthemum of the West Indies. He did not list their species but his group Hesperanthemum, comprising ‘‘ frutices antillani spinosi’’ with ‘‘ pedun- culi axillares, subuniflori’’, obviously refers to the group typified by Evanthemum spinosum (Jacq.) R.Br. ex Roem. & Schult., ie. Justicia spinosa Jacq. Kuntze raised this group to generic rank in 1891 as Hesperanthemum, thereby creating a superfluous new name. At the same time, obviously knowing nothing about the plants themselves covered by this nomenclatural exercise, Kuntze described as Jasminum coeruleum the only one he himself had collected on St. Thomas in the West Indies. As pointed out by Green (1969), there are no Jasminum species native to the New World, J. fluminense Vell. being a naturalized African species, and Kuntze’s Jasminum coeruleum is here regarded as conspecific with Oflonia spinosa.
Although the genus Oflonza is easily recognised and defined, the species are far otherwise. Gustav Lindau in 1895 described it as having “‘ 6 (oder mehr) schwer zu trennende, westindische Arten’’. The difficulty of separating the nine species named by earlier authors led him in 1g00 to reduce them to three distinguished on characters so unreliable that he might just as well have been content with one. Lindau regarded A. armatus as a species endemic to Jamaica, which it is, but he treated A. acicularis, in fact also a Jamaican endemic, as “ eine sehr variabele Species” extending from the Bahamas over Cuba, Jamaica, Hispaniola, St. Thomas and Antigua to Guadeloupe and Marie Galante. To A. spinosus he attributed a similar range, from the Bahamas over the Greater and Lesser Antilles to Guadeloupe and Martinique. Specimens with puberulous flowers he referred to the one, those with glabrous flowers to the other, a procedure of no value whatever.
Meanwhile Lindau described in 1897 a new genus Psilanthele, based on a new species , P. eggersit, collected by Baron Henrik Eggers (Eggers 15129) in the province Manabi of Ecuador, which has very small flowers, the corolla being about 4 mm. long, in 10-20-flowered raceme-like panicles terminating axillary shoots (Pl. 47). The generic name, unexplained by Lindau, is evidently from yrds, “‘ smooth, bare”, av@7An, “ blossom, panicle’’. Since he considered its pollen to be of the “ Rah- menpollen ”’ type typical of the Graptophylleae, whereas Anthacanthus and Odonto-
* As accepted by Nees, the genus Evanthemum L., of which the type species is E. capense L., included not only species of Evanthemum sensu stricto (Daedalacanthus T. Anderson) but also of Pseuderanthemum Radlkofer (‘‘ Evanthemum ”’ sensu T. Andersoni, non L.), as explained by Stapf in Bot. Mag. 135 : t.8239 (1909) and Ruspolia Lindau (cf. Milne-Redhead in Kew Bull. 1936 : 255-274 (1936) ). Oplonia, Psilanthele and Odontonema are related to Pseudervanthemum and not to Evanthemum, of which one species, E. pulchellum Andrews (E. nervosum (Vahl) R. Br. ex Roem. & Schult., Daedalacanthus nervosus (Vahl) T. Anderson), is naturalized in the West Indies.
OPLONIA AND PSILANTHELE (ACANTHACEAE) 265
nema were characterized by “‘ Spangenpollen ’’, Lindau did not associate it with the latter but with Carlowrightia, Anisacanthus and Harpochilus.
In 1904 Lindau added to his genus Psilanthele a second species, P. grandiflora (Pl. 42) collected by Ernst Ule (Ule 6493) in north-eastern Peru, which differed from the type-species in having larger flowers, the corolla being about I cm. long, in con- densed axillary fascicles. He noted that this had two minute staminodes and “ pollinis granula typica, fere globosa, c. 38 uw diam.’’ In the following account, Psilanthele is restricted to its type species and P. grandiflora is transferred to Oflonia.
The extensive collections made by William Harris in the interior of Jamaica brought to Lindau’s attention a species agreeing in its contracted inflorescence with the Peruvian P. grandiflora but having larger flowers, the corolla being nearly 2 cm. long, although with “ pollinis granula iis Graptophyllearum conformia, sub- globosa, 46-50 yw diam.”’ Lindau named this P. jamaicensis in 1908. That year Harris collected material of an allied plant on the coast of Jamaica which Lindau described in 1912 as P. minor (Text-fig. 1). He then pointed out that Psilanthele resembled Anthacanthus so closely in habit and flower that only the constant lack of spines in the former and a difference in pollen separated them : “‘ Psilanthele zeigt stets deutlichen Rahmenpollen, Anthacanthus nur Spangenpollen’’. A Jamaican specimen without spines (Harris 10224) and identical with P. minor but possessing “Spangenpollen”’ he accordingly referred to Anthacanthus. Lindau took this as evidence of the over-riding importance of the character of the pollen grain, as opposed to general resemblance, in the classification of the Acanthaceae. With equal cogency he could have taken it to exemplify the danger of over-emphasis on a slight difference in the marking of the pollen grain when countered by resemblances in everything else.
Increase in the number of specimens available for study has increased, not lessened, the difficulty of defining taxa within this group. When travelling and collecting in Jamaica in 1955-56 (cf. Stearn, 1959) I collected specimens of these plants in many parts of the island. A low-growing white-flowered species occurring in the parishes of St. Thomas and Portland, subsequently identified as Swartz’s Justicia acicularis (now Oflonia acicularis), was easily recognizable in the field as distinct from the other members of the genus ; a taller-growing species, now referred to O. microphylla, with
very small leaves and pale lilac flowers found on coastal limestone near Kingston
has also a distinctive appearance. The large leaves and numerous flowers of O. jamaicensis set this apart from the others, as regards which there appeared to be no evident pattern of variation with convenient discontinuities. Instead the speci- mens collected merely indicated the existence of numerous local populations, probably themselves more or less homogeneous and with their own assemblage of minute characters but not well differentiated from other local populations. Preparation of an account of the Acanthaceae for the continuation of Fawcett and Rendle’s Flora of Jamaica made it necessary to classify this bewildering and exasperating material. Questions of nomenclature then arose, which involved taxonomic decisions ; whether, for example, early names based on material from other West Indian islands could be applied to Jamaican taxa. In dealing with these it became painfully clear that a revision of the group over its whole range from the Bahamas to the Grenadines and
266 A SURVEY OF THE TROPICAL GENERA
into Peru could not be avoided. The present paper is the outcome of this survey and is based on about 250 gatherings. In 1900 Lindau based his account on some 60 ~ gatherings ; Nees von Esenbeck in 1847 had about 12 gatherings. The close resem- blance of Jamaican species placed in Oflonia and Madagascan species placed in Forsythiopsis was quite unexpected and did not become evident until long after the completion of the account of the American group.
TAXIONOMIC CHARACTER
‘
Lindau (1900) wrote of his ‘‘ Anthacanthus acicularis”’ that it was “‘ eine sehr variabele Species, sowohl was Behaarung wie auch Blattgrésse betrifft. Dadurch, dass die Dornen bald lang bald kurz, bald gerade abstehend oder im Winkelabstehend, bald etwas gebogen, bald sehr diinn, bald holzig dick sind, wechselt der Habitus ungemein.’’ Within this he included the plants distinguished by other authors as Anthacanthus microphyllus, A. cuneatus, A. lycioides, A. purpurascens and A. tetra- stichus, although not, as it happens, any specimens of the true A. acicularis! He thus dismissed as having no taxonomic value their diversity in form and length of spines. Of his A. sPinosus he wrote similarly ; ‘“‘ Ebenso wie vorige Art sehr variabel im Habitus. Die Veranderlichkeit erstreckt sich auf die Stacheln und die Blatter ”’. He was left with “ flores + puberuli”’ as the distinguishing feature of A. acicularis and “‘ flores semper glaberrimi”’ of A. sbinosus. The characters mentioned above cover the major variation within the genus. They may be summarized as follows :
Habit of Growth. Most members of the genus are shrubs up to 2 metres high. The extremes are represented by Oplonia nannophylla (Pl. 45) of Cuba, a shrublet only 4-8 cm. high with internodes 1-3 mm. long, and O. armata var. pallidior of Jamaica, which may grow into a small tree. Possibly each species when living has a distinctive habit. As noted above, I found O. acicularis easily recognizable in Jamaica by its low, much- branched growth as well as by its white flowers.
Pubescence of Branches and Spines.
With age the shoots of Oflonia lose their original epidermis, this being replaced by a thin bark, and become glabrous. As in Thymus (Labiatae) and Epilobium (Ona- graceae), the young four-sided stems may be holotrichous, i.e. with hairs on all sides, or goniotrichous, i.e. with the two opposite sides immediately below the leaves glabrous and the two opposite intervening sides hairy, their position thus alternating from node tonode. The hairs themselves may be relatively long (up to 0-5 mm. long) or much shorter (down to o-I mm. long), dense or sparse, horizontally spreading or slightly or distinctly upcurved.
Variation in the pubescence of shoots is well illustrated by a series of specimens belonging to a small-leaved taxon (here included in Oplonia microphylla) which grows on the dry limestone over a limited area of south-eastern Jamaica near Kingston (Text-fig.2J-M). The type-specimen of Anthacanthus cuneatus Nees (Macfadyen s.n. in Herb. Kew) exemplifies one extreme : white spreading hairs up to 0-5 mm. long cover densely all four sides of the shoots, while the spines are densely pubescent with
OPLONIA AND PSILANTHELE (ACANTHACEAE) 267
Fic. 2. Spines and leaves of Oplonia : A,B,C, O. spinosa (Jacq.) Raf. (A, Buch 1851, Haiti ; B, Curtiss 133, New Providence ; C, Marie-Victorin et al. 21625, Cuba) ; D,E, O. tetrasticha (Griseb.) Stearn var. tetvasticha (D, Proctor 16430, Cuba ; E, isotype, Wright 3067, Cuba) ; FG, O. tetrasticha var. polyece Stearn (F, Clemente 5014, Cuba ; G, holotype, Clemente 5657, Cuba) ; H,I, O. acicularis (Swartz) Stearn (H, authentic specimen, Swartz, Jamaica; I, Stearn 498, Jamaica) ; J,K,L,M, O. microphylla (Lam.) Stearn (J, isotype, Richard, St. Croix ; K, Howard rog16, Grenadines ; L, Avaque-Molina & Barkley 22] 448, Jamaica ; M, holotype of Anthacanthus cuneatus, Macfadyen, Jamaica) ; N, O. nannophylla (Urban) Stearn (holotype, Ekman 16532, Cuba) ; O, O. armata var. pallidioy Stearn (Stearn 863, Jamaica) ; P,Q, O. purpurascens (Griseb.) Stearn (isotypes, Wright 3066, Cuba) ; all x 14. Cystoliths are present on all leaves but are indicated only in figs. H and I.
268 A SURVEY OF THE TROPICAL GENERA
shorter hairs. Similar abundant pubescence of branches and spines occurs in Araque-Molina & Barkley 22]448 and Proctor rorg5, both gatherings from near localities in Kingston and St. Thomas ; Proctor 9971 and Asprey 2059 from St. Catherine and Proctor 18268 from St. Thomas have the shoots likewise densely hairy but the spines glabrous ; Harris 9606, Proctor 11436, Stearn 66, and Stearn 83o, all from the Long Mountain, Kingston and St. Andrew, have two opposite sides densely hairy but the other two sides almost glabrous and the spines glabrous. In Harris 8609 from St. Andrew the hairs are much more abundant on two opposite sides than on the intervening sides but the hairs are only 0-I to0-2 mm. long. Thus three types of hair covering, which in some genera, e.g. Tiymus, would be taken as evidence of specific distinctness, here occur on plants of similar habit growing under similar conditions in the same region.
Spines.
The spines of Oflonia (Text-fig. 2 ; Pl. 45) are reduced shoots ; hence they arise singly and opposite one another in the axils of leaves, each node having but one pair. The apparent exception, Grisebach’s Anthacanthus bispinosus with two pairs of spines at a node, does not belong to Oplonia, its spines being modified leaves, and proves to be Barleriola solanifolia (L.) Oerstedt, specimens of which may be found in herbaria determined as Anthacanthus spinosus (e.g. J. H. Jack 5596) and A. tetra- stichus (e.g. A. Gonzales 672). Spines appear to be constantly absent from the mesophytic O. jamaicensis and O. grandiflora. Among the specimens referred to O. armata var. pallidior some have no spines, other have well developed spines ; indeed the same bush may bear spiny and spineless shoots, as on Stearn 863 from St. Elizabeth. Spines are usually well developed in the xerophytic species. They then supply some conspicuous diagnostic characters. Thus spines spreading horizontally but with a slight downward curve or curved so strongly as to be hooked or deflexed characterize O. spinosa as accepted here. Such spines are found on specimens collected from the Bahamas to the Virgin Islands but not on any Jamaican specimens. Very long, straight, usually stout and rigid spines, varying between I cm. and 3:5 cm. in length, distinguish O. tetrasticha of Cuba ; spines more than 1-4 cm. long are un- known outside Cuba. Spines very slender for their length, i.e. 8 mm. to 14 mm. long but about 0-3 mm. thick, help to give O. acicularis of Jamaica its distinctive habit. In O. microphylla they are shorter and more rigid. The development of pairs of short subsidiary spines on the ordinary spines is characteristic of O. tetrasticha var. polyece. The Madagascan species have no spines.
Leaves. The largest leaves of the genus are found in O. grandiflora (Pl. 42) and O. jamaicensts where they vary on the same plant from 2 cm. to 10 cm. in length and to 3 cm. or moreinwidth. At the other extreme are O. nannophylla, O.linifolia and O. microphylla with leaves only 2-10 mm. long. On many specimens the leaves seem fairly constant in size and shape but on others there may be remarkable diversity. Thus a gathering made on St. John, Virgin Islands (Robertson 26), here referred to O. spinosa, has the leaves almost circular or very broadly ovate, the length and breadth being almost the same with the greatest breadth at or below the middle. On
OPLONIA AND PSILANTHELE (ACANTHACEAE) 269
a gathering by A. C. Smith (A. C. Smith 10562) from Virgin Gorda, Virgin Islands, also referred to O. spinosa, some leaves are likewise broadly ovate but others are elliptic or narrowly obovate, i.e. proportionally narrower with the greatest breadth at or above the middle. Such specimens connect with the narrowly elliptic and oblanceolate leaves of O. spinosa on specimens collected in Haiti and the Bahamas. The apex of the leaf is acute to acuminate in O. acuminata and O. grandiflora. The Madagascan O. linifolia has linear leaves. The others have the apex itself rounded and usually slightly or distinctly indented, sometimes with a distinct mucro.
The veining (Pl. 43b) is evident in some specimens, faint or invisible in others, and seemed at first to offer characters of possible taxonomic value. Using the clearing technique described by Arnott (1959), preparations were made of leaves from a diversity of plants, these inserted in a microfiche viewer, and their veining then drawn on a vastly enlarged scale. It proved to be brachydromous in all, i.e. with about 6 to 11 lateral veins on each side of the midrib running slightly obliquely almost to the margin, then bending abruptly forward and joining the bend of the lateral vein immediately above, thus making a closed series of loops more or less parallel to the margin. Variation in the number of veins and their angle of diverg- ence from the midrib was slight.
Number of Flowers. The inflorescence of Oplonia occasionally develops into a short lateral raceme but is usually reduced to a sessile axillary fascicle, which may bear up to 15 flowers in the large-leaved O. jamaicensis but is commonly only one- or two-flowered in species with leaves less than 2:5 cm. long.
Pedicels. The pedicels are mostly much longer than the calyx and up to x cm. long. In O. purpurascens, however, the flowers are almost sessile with pedicels 1-1-5 mm. long and thus shorter than the 3-4 mm.-long calyx. Specimens of O. microphylla some- times have pedicels as short, but longer pedicels are usually to be found on the same plant. They are usually glabrous, but occasional individuals have hairs on the pedicels.
Calyx. The calyx is parted almost to the base into 5 narrow segments which are linear in O. grandiflora but very narrowly triangular (subulate) in the others. The calyx is usually about $5 mm. long. Its length does not seem correlated with other char- acters. In specimens referred to O. armata var. pallidior it varies from 2 mm. (Howard, Proctor & Stearn 14740, Norman 100) to 5 mm. (Stearn 863).
Corolla
The corolla (Text-fig. 1, 3) is surprisingly uniform in size and form and consists of a slender tube slightly expanded above the insertion of the stamens on its upper part and a spreading two-lipped limb with two ascending more or less fused upper lobes and three descending distinct lower lobes. Tube and limb are almost equal in length. In 0. grandiflora, so inappropriately named, the tube is only 4-4-5 mm. long, in O. acicularis 4-6-5 mm., in the others 6-10 mm. long. Although Lindau clutched at the
270 A SURVEY OF THE TROPICAL GENERA | ; presence or absence of hairs on the corolla in despair of finding any other character — for specific distinction, this seems of trivial value. The hairs are very small and — are present or absent without correlation with any other characters.
The red colour of the corolla of O. armata var. armata readily distinguishes it in a living state from all other members of the genus. In these the corolla is mostly a bluish lilac or bright purple having at the base of the lower lip a pale area marked with dots or streaks of purple, which presumably serve as nectar guides. In O. acicularts the corolla is normally white.
Length of Stamens and Style In some flowers of Oflonia the stamens project conspicuously from the tube of the corolla, in others they scarcely reach beyond its mouth (Text-fig. 3). This diversity
Fic. 3. Floral dimorphism in Oplonia armata (Swartz) Stearn ; flower cut lengthwise : A,B, O. armata var. pallidior Stearn (A, Adams 11147, Round Hill, Clarendon, Jamaica ; B, Adams 11143, same locality) ; C,D, O. armata var. armata (C, Adams 11062, between Aenon Town and McKay, Clarendon, Jamaica ; D, Adams rro60, same locality). Flower, Xo 2 MEAVES SGT:
seemed at first to offer welcome specific characters but undoubtedly expresses floral dimorphism, merely separating individuals within the same population, such as occurs in many gamopetalous families, notably the Rubiaceae (cf. Burck, 1883-4 ; Bahadur, 1963), although not previously recorded for the family Acanthaceae. Thus
OPLONIA AND PSILANTHELE (ACANTHACEAE) 271
in the specimens of south-eastern Jamaica mentioned above as referred to O. micro- phylla and agreeing closely in habit, some have short stamens and long styles, others long stamens and shorter styles. For example, Proctor 18268 from St. Thomas, Harris 8609 from Kingston and St. Andrew, Asfrey 2059 and Proctor 9971 from St. Catherine have the filaments about 1-2-1-5 mm. long, the style 7-9 mm. long. Araque-Molina & Barkley 22]448 from St. Thomas, Proctor 11436, Stearn 66 and Stearn 830 from Kingston and St. Andrew have the filaments about 5-5-5 mm. long, the style 4-5-5 mm. long. Both types occur on the Long Mountain near Kingston. Among specimens of O. armata var. pallidior, Stearn 726 has the filaments 1-5 mm. long, the style 10 mm. long, but Proctor 16655 has the filaments 5:5 mm. long, the style 6 mm. long. Likewise, of specimens of O. armata var. pallidior collected by Dr. C. D. Adams on Round Hill, Clarendon on 13 May 1962, Adams 11147 has long- exserted stamens and a short style, Adams 11143 short stamens and a long-exserted style (Text-fig. 3, A,B). Similar dimorphism is found in O. armata var. armata collected by Adams between Aenon Town and McKay, Clarendon, Adams 11062 (with light crimson corolla) having long-exserted stamens and a short style, Adams rro60 (with rich-violet corolla), short stamens and a long-exserted style (Text-fig. mC, D).
The same floral dimorphism is evident in Cuban material of O. spinosa all from the Moa region of Oriente province. Thus Victorin, Clemente & Alain 21621 and 21625 have the filaments about 2 mm. long, the style 10 mm. long, but in Clemente 3665 and 3646 the filaments are about 4:5 mm. long, the style about 5-6-5 mm. long. Such floral dimorphism probably occurs throughout the genus ; several specimens agreeing in other respects and from the same district or, better, the same locality are needed to demonstrate this. Its development is of great evolutionary interest. Its presence in Oplonia raises the question whether it may not have been overlooked elsewhere in the family and have thus led to unjustified separation of long-stamened and short- stamened forms as different species, although, as Darwin (1877 : 136)* pointed out, in the Rubiaceae ‘‘ this character, which is not even of specific value in the hetero- styled species, is often of generic value in other members of the family ’’. It also occurs in Madagascan specimens formerly placed in the genus Forsythiopsis. Thus in a gathering of O. minor (Humbert 2506) one specimen has the filaments about 2-5 mm. long and the style 7 mm. long, another has the filaments about 5 mm. long and the style 5 mm. long. A specimen of O. vincoides collected near Fort Dauphin by Cloisel (Cloisel 209) has the filaments about 3 mm. long, the style about 10 mm. long, while another, also from Fort Dauphin, collected by Scott Elliott (Scott Elliott 2586) has the filaments about 6 mm. long, the style 6-8 mm. Thus heterostylous floral dimorphism probably occurs throughout the genus.
Pollen
Lindau attributed ‘‘Spangenpollen” to Oflonia and ‘‘ Rahmenpollen”’ to Psilanthele, using this as a generic distinction. Actually the difference between “ Rahmenpollen’”’ and ‘‘ Spangenpollen”’ is by no means clear-cut. The pollen grains of typical species of Oplonia such as O. microphylla (Text-fig. 4) and O. armata
* The “‘ Faramea”’ described by Darwin has been discussed by H. G. Baker, ‘‘ Pollen dimorphism in the Rubiaceae ’’, Evolution 10 : 23-31 (1956), 340 (1956) and proves to be Rudgea jasminoides Muell.-Arg.
272 A SURVEY OF THE TROPICAL GENERA
var. pallidior (Pl. 38) are spheroid or prolate, 30-50 » m. in diameter, with three germ pores each in a longitudinal furrow (mesocolpium) reaching almost to the pole, the grain thus being primarily three-furrowed (tricolpate), but with a shorter subsidiary longitudinal furrow (pseudocolpus) on each side of the main furrow ; all nine furrows are separate and do not join. They are “ Spangenpollen’’, as figured by Lindau (1893 : 38, fig. I L, t.2 fig. 62 ; 1900 : 173, fig. 12, 13), since the subsidiary furrows remain distinct and do not continue towards the poles and there fuse. In ‘‘ Rah- menpollen ”’, as figured by Lindau (1893 : fig. 1 O, t.1 fig. 49-51 ; 1900 : 173, fig. Io, Ir) and attributed by him to Psilanthele, the two adjacent subsidiary furrows meet near the poles, thus joining into three pairs. If these furrows converge but do not quite touch, then presumably the grain belongs to the “‘ Spangenpollen’”’ type (cf. Pl. 38, 40) ; if they join, then it belongs to the “‘Rahmenpollen”’ type (cf. Pl. 39).
Fic. 4. Pollen grains (‘‘ Spangenpollen ’’) of Oplonia microphylla (Lam.) Stearn (Stearn 66 ; drawing based on Stereoscan photograph), x 700.
This distinction is not always easy to observe even on grains stained and magnified x 1500, yet it is the only one, following Lindau, by which the Jamaican species referred to Psilanthele can be distinguished generically from Anthacanthus, i.e. Oplomia. Actually there is no difference in pollen grains between Oflonia and Psilanthele. Photographs made with the scanning electron microscope (Cambridge Instrument Co. Stereoscan), at magnifications between x 500 and x 5,700, of pollen grains of Oplonia microphylla (Stearn 66), O. armata var. armata (Proctor 21407), O. armata var. pallidior (Stearn 726), O. acicularis (Britton 2894), O. jamaicensis (Harris 8968), O. grandiflora (Ule 6493), the Madagascan O. vincoides (Baron 1937, McWhirter & Capuron 184) and Psilanthele eggersti (Eggers 15129) show them to be essentially uniform in form, although varying in size, with their subsidiary furrows sometimes joining but usually not quite reaching each other. The type species of Psilanthele (P. eggersit of Ecuador) differs from Lindau’s Peruvian and Jamaican species not in pollen grains but in its loose inflorescence and very small flowers with the two loculi (thecae) of the anther separated a little, not absolutely parallel and contiguous as in Oplonia. Hence the genus Psilanthele is here restricted to P. eggersit and the other species are transferred to Oplonia.
a; 2
OPLONIA AND PSILANTHELE (ACANTHACEAE) 273
Variation in size of pollen within a species can be illustrated from Jamaican material of O. microphylla, measurements taken by Dr. S. Sengupta from Io grains each of 8 specimens being as follows : _ 1. Proctor 18268 : diameter range 33-3 um—36-0 ym ; spheroid. 2. Proctor 24186 : diameter range 34:2 wm-—36-0 wm ; spheroid. 3. Stearn 66: diameter range 31:5 wm-—32:4 wm ; prolate, with length range 43°2 pm—50°4 pm. 4. Webster, Ellis & Miller 8364 : diameter range 28-8 wm—29°7 wm ; prolate with length range 45-0 um—49°5 wm. 5. Harris 9606 : diameter range 28-8 wm—29-7 wm ; prolate, with length range 40°5 pm—43°2 pm. 6. Araque-Molina & Barkley 22]448 : diameter range 32:4 um—36-0 um ; spheroid. 7. Stearn 830 : diameter range 35:2 um-—37°8 wm ; prolate-spheroid, with length range 38-7 um—40°5 yum. 8. Proctor 9971 : diameter range 27:0 ym—28-8 ym ; prolate-spheroid, with length - range 28-8 wm-—32-4 wm.
TAXONOMIC PROCEDURE
As noted above, the plants of this group show diversity in habit, in length, form and direction of spines, hairiness of branches, form and size of leaves, number of flowers at an axil, length of pedicel, length of calyx, colour and pubescence of corolla, and length of filaments and style, without there being much readily evident corre- lation between the characters of one organ and those of another. The difficulty of perceiving associations is increased by the smallness of the organs themselves, which require examination one by one with a lens ; each character thus presents itself to the mind as a separate entity not linked with others. Lindau’s treatment of 1900 brought together under one species a multitude of very unlike specimens while at the same time referring almost identical specimens to different species. Dissatisfied by this and by the evident confusion, I decided to ignore for the time being all literature about the group and all determinations on the herbarium sheets, and instead to begin anew by treating each specimen as if it were a new and independent taxon, tabulating its characters and then trying to ascertain how these were correlated with others. This proved a laborious and time-consuming procedure, despite the independent use of three complementary techniques, i.e. punched cards, metroglyphs and correlation coefficients. Firstly, characters of potential utility were recorded on marginally punched cards. These made it possible to check the distribution of characters and to test likelycombinations, usually with negative results ! Secondly,a chart was made by expressing a few salient characters of each specimen (e.g. the size and shape of the leaves, the type of pubescence and the colour of the corolla) as pictorialized scatter diagrams or metroglyphs (cf. Text-fig. 7 B), following Edgar Anderson (1952 ; 1957 ; see also Hatheway, 1962.) Thirdly, with the help of Dr. P. H. A. Sneath, a range of 33 character states on 68 specimens were coded for computer-aided single-linkage cluster analysis ; Sneath then calculated their correlation coefficients (cf. Sneath, 1962) and produced a dendrogram indicating overall similarity in the characters taken. Those available in Oplonia fell far below the postulated minimum of forty characters
274 A SURVEY OF THE TROPICAL GENERA
and some unexpected associations resulted. Metroglyphs seemed to provide the best method of grasping the essentials of this complex taxonomic situation, punched cards a means of quickly retrieving the information, correlation coefficients a means of grouping objectively the specimens or taxa. In fact none of these methods gave a satisfying arrangement by itself. They resulted, however, in bringing together piles of specimens having many characters in common, which it had been difficult or impossible to associate by simple observation, since there were too many inconspicuous variables concerning too many specimens for them all to be kept in mind at the same time. It then became possible to take more or less homogeneous clusters of specimens and associate other specimens with them, later uniting these groups with other groups as the discontinuities between the groups were obliterated or bridged. Be- fore this process merged the twenty or so provisional West Indian taxa into one, the dominance of an easily observable feature in a particular group and its absence from others, e.g. very small leaves, hooked or decurved spines, very long spines, numerous flowers in a fascicle, or red corolla, became evident. Most of the material could then be placed into groups characterized by these features ; ambiguous specimens were referred to them on a basis of general resemblance to already placed specimens from the same island or district. The result has led me to accept nine generally recogniz- able but not always clearly distinguishable American groups treated below as species.
TAXONOMETRIC SURVEY
The most difficult task when classifying apparently rather uniform biological material, such as that of Oflonia, wherein most distinctions lie in a diversity of minute characters neither simultaneously observable nor easily correlated, is that of putting the specimens into small relatively homogeneous groups which can later be joined into larger groups as defining attributes become evident ; obviously then, the study of such material would be eased by the use of an efficient computer-aided method of assembling the specimens into successively united groups (clusters) characterized at first by high overall resemblance between the members of each group, a resemblance which would decrease progressively as more groups were united. Hence it seemed worthwhile, as a guide for future procedure in dealing with intractable groups like Oplonia, to apply such methods to an assemblage of specimens of Oplonia.
Availability of computers and exercise of mathematical ingenuity have now led to the creation of possible numerical methods bewildering in their number and claims (cf. Johnson, 1968, for comment and extensive bibliography ; also Cole, 1969). Assessment of their respective merits for a given task must depend upon practical experience. They need to be tested by application to group upon group (cf. Stearn, 1969).
The following observations relate to a method of cluster analysis based on graph theory, to which Professor D. J. Rogers (then at the New York Botanical Garden, now at the University of Colorado) drew my attention in 1964, after the present study, mostly done in 1961-62, had been virtually completed. This method has since been expounded mathematically by Estabrook (1966) and applied by Wirth, Estabrook
—
OPLONIA AND PSILANTHELE (ACANTHACEAE) 275
and Rogers (1966) to Orchidaceae subtribe Oncidiinae, by Irwin and Rogers (1967) to Cassia sect. Apoucouita and by Prance, Rogers and White (1969) to Chrysobalanaceae tribe Hirtellae. It groups objects into clusters, defined by similarity of their mem- bers, with stage-by-stage reduction in the degree of similarity necessary for objects to be placed in the same cluster. It is a process wherein hitherto unclustered objects are progressively admitted into earlier-formed clusters through linkage by resem- blance to objects already in the cluster. Thus small clusters become linked to larger clusters ; as a cluster enlarges through incorporation of other clusters, so the amount of similarity between all members of the resulting cluster lessens at each stage until ultimately the objects all come together into one large cluster. The computer print- out for the whole process records it as a series of stages of clustering. The taxo- nomist must decide which of the intermediate stages, when there exist several or many clusters of specimens, have the most relevance to his needs.
The first task, as in other taxonometric procedures, consists of listing all the available characters and their states differing from specimen to specimen ; these are then tabulated for each specimen being studied. Comparison by the computer pairs the specimens according to their mutual similarity. This stage, the first partition, divided the material into the basic classes. The principles governing procedure, as stated by Estabrook (1966), are :
“ (1) a biological classification for a collection of objects is a series of partitions for this collection, classes under later partitions consisting wholly of classes under earlier partitions. (For example, genera consist wholly of species.)
“ (2) For any given partition in this series, two similar objects should not be placed in different classes (this assumes that no knowledge of evolution is available).
“ (3) The classes of a given partition should be isolated from one another ; that is, there should be some phenotypic discontinuity between members of different classes. ”’
As the original clusters (disjoint partitions) merge into the ultimate single cluster
(conjoint partition), by coming together at given degrees of lessening similarity, the
greater the overall resemblance between specimens the earlier will be the stage at which they are linked into a common cluster ; conversely, the greater their divergence so much the later will be their linkage. Part of the task of preparing a taxonomic revision of a group has always been a search for such resemblances and divergences,
followed by hierarchic ranking of the sub-groups detected. A machine technique
quickly displaying these relations can thus significantly lessen the labour of research ; it may even indicate correlations of characters which otherwise would not have been perceived.
To test this computer-aided hierarchical clustering technique based on graph theory, the characters, 26 in all, of 35 specimens of Oflonia representing the geo- graphical and morphological extremes of the group as well as numerous intermediate states were tabulated by me and then computed (using Taximetric Similarity Graph- Clustering programme on CDC 1604 computer at New York)* by Rogers, who con- verted the computer print-out into a series of diagrams (subgraphs), each represent-
* This study was run on a Control Data 1604 under the direction of D. J. Rogers of the Taximetrics Laboratory then located at the New York Botanical Garden but now at the University of Colorado, Boulder, Colorado, U.S.A. The programmes are operative on IBM 7000 series + 360/75 and CDC 6400.
B
276
A SURVEY OF THE TROPICAL GENERA
ing one partition of 21 stages (levels) of clustering. These specimens (to which the scientific name now adopted has been added) were as follows :
0M ON AU BW DN H
HH H
12: rae 14. £5:
16; 17. 18. 19. 20. 21:
22: 23. 24. 25,
26. 27. 28. 29. 30.
31. 32% 33-
34. 35.
. Britton 2877 ; . Stearn 550 ;
. Britton 2894 ; . Proctor 8433 ; . Stearn 454 ;
. Harris 8968 ; . Proctor 16430 ; . Leon 3326 ;
. Curtiss 133 ;
. Robertson 26 ; . Stearn 726 ;
Proctor 21407 ;
Box 757 ;
Howard rogt6 ;
Araque-Molina & Barkley 22] 448 ;
Proctor 9971 ;
Stearn 830 ;
Norman Ioo ;
Stearn 863 ;
Ekman 15042 ;
Marie-Victorin, Clemente & Alain 21625 ;
Leonard & Leonard 13674 ;
Valeur 724 ;
Ekman 16534 ;
Clemente 5657 ;
Buch 1851 ; Proctor 10236 ; Schomburgk 107 ; Stearn 486 ; Wright 3067 ;
Clemente 5014 ; Stearn 455 ; Proctor 8830 ;
Stearn 498 ; Ule 6493 ;
Jamaica ; Jamaica ; Jamaica ; Jamaica ; Jamaica ; Jamaica ; Cuba ; Cuba ; Bahamas ; Virgin Islands ; Jamaica ; Jamaica ; Antigua ;
Petit St. Vincent ;
Jamaica ; Jamaica ; Jamaica ; Jamaica ; Jamaica ; Cuba ;
Cuba ; Hispaniola ; Hispaniola ; Cuba ; Cuba ;
Hispaniola ; Jamaica ; Hispaniola ; Jamaica ; Cuba ;
Cuba ; Jamaica ; Turks & Caicos
Islands ; Jamaica ; Peru ;
Ove OS 7 O00 OO OS eS
O. acicularis
O. acicularis
O. acicularis
O. jamaicensis
O. jamaicensis
O. jamaicensts (isotype)
. tetrasticha var. tetrasticha . tetrasticha var. tetrasticha . spinosa
. spinosa
. armata var. pallidior
. armaia var. armata
. microphylla
. microphylla
. microphylla
. microphylla
. microphylla
. armata var. pallidior
. armata var. pallidior
. tetrasticha var. tetrasticha
. spinosa
. spinosa
. microphylla
. nannophylla (type)
O. tetrasticha var. polyece (type)
O. spinosa
O. armata var. armata
O. spinosa
O. armata var. armata
O. tetrasticha var. tetrasticha (isotype)
O. tetrasticha var. polyece
O. armata var. armata
O. spinosa O. acicularis O. grandiflora (isotype)
Study of Rogers’s series of diagrams (subgraphs) in association with the specimens themselves revealed particularly the utility of the middle stages (levels) of clustering
OPLONIA AND PSILANTHELE (ACANTHACEAE) 277
as an aid to perceiving and distinguishing the main components of the group. Text-fig. 5, for example, represents the clustering reached at stages 6 and 12. The first partition (stage 1) made according to a very high degree of resemblance brought together the almost identical Jamaican specimens (Nos. 15, 16, 17) now referred to O. microphylla. By stage 6 Jamaican specimens (Nos. I1, 12 and 18, 19) now referred to O. armata had been placed in two clusters, Lesser Antillean specimens (Nos. 13, 14) referred to O. microphylla in another, Cuban specimens (Nos. 7, 8) referred to O.
STAGE 6 STAGE 12 ler S iv 7 {zo {ei} 28) S
eel
z s[4}-—{s]3 3 [s}—_{4} +]
Fic. 5. Stages 6 and 12 of 1-22 sequence in cluster analysis of 35 specimens of Oplonia (listed by numbers on p. 276) by graph-theory method as computed by D. J. Rogers ; A, specimens classified as O. armata ; J, as O. jamaicensis ; M as O. microphylla ; S as O. spinosa ; T as O. tetrasticha.
tetrasticha and Jamaican specimens (Nos. 4, 5) referred to O. jamaicensis in yet others. By stage 12 the grouping and linking have gone far enough to make a significant arrangement of most of the specimens. Thus Nos. 13, 14, 15, 16, 17, and 23 when put side by side agree in having very small leaves and short ascending spines and constitute a taxon for which the earliest epithet is microphylla ; Nos. 7, 8, 20, 25 and
278 A SURVEY OF THE TROPICAL GENERA
30, all from Cuba, with their long straight horizontal spines form another group, for which the epithet is tetrasticha ; Nos. 9, 21, 22, 26, and 28 with their mostly curved — horizontal or deflexed spines likewise form a group, to be called spinosa ; Nos. 4, 5_ and 6, all from Jamaica, with large leaves and many flowers represent the easily distinguishable jamaicensis. Among the specimens unclustered at this stage, No. 24 (nannophylla), Nos. I, 2, 3 and 34 (acicularis) and 35 (grandiflora) strike the eye as” well distinct from the others, while Nos. 32 (armata), 10 (spinosa) and 33 (armata) show resemblance to some specimens already clustered. The last specimen to enter
was No. 35 (grandiflora), from Peru, which accords with its morphological and_ geographical distinctness. The logical coherence of this grouping made me regret
that the technique used had not been available much earlier ; it would have provided
a means of grouping fairly quickly the specimens into the taxa ultimately adopted without so much time-consuming search for correlations. The computer run dealing with this material took 3 minutes 47 seconds, an infinitesimal amount of time com-
pared with that of ascertaining and tabulating data and that of expressing and con-
sidering the results. The method commends itself for its graphic portrayal of classi-
fication as a stage-by-stage process and corresponds closely enough in its working to
the mental acts of a working taxonomist to be able to serve as a time-saving substi-
tute for some of them. Thus in dealing with a large confused group it is convenient
to pick out various types and use these as focal points to which specimens can be referred by resemblance, thereby creating assemblages from which concepts of
taxa can take shape and definitions be formulated. This computer method creates
such assemblages in the same typological manner, presents them to the eye pictorially,
and thus prepares the way for defining their circumscription and ascertaining the
characters by which they can be distinguished.
The reasonableness of the classifications of genera (cf. Sims, 1966) and species (cf. Wilkinson, 1967 ; Stearn, 1969) produced by computation using the computer programme CLASP indicated the desirability of testing it as a means of classifying individual specimens, i.e. of arranging them into groups to be treated as species or subspecies, by taking each specimen as a separate operational taxonomic unit (OTU) for association with other specimens on a basis of overall shared characters, as had been done using the different technique outlined above. Accordingly essentially the same characters (24) of the same specimens (35, plus one of O. purpurascens from Cuba earlier inadvertently omitted) were coded and tabulated for computation as follows :
Two-sTATE (BINARY)
tT. a= tall ; low.
2. Internodes : 5 mm. long or less ; more than 5 mm. long. 3. Hairs of stem: present ; absent.
4. Hairs of stem: short ; long.
5. Hairs of stem: on 2 sides ; on 4 sides.
6. Spines : present ; absent.
7. Shines : short ; long.
8.
Spines : fine ; stout.
OPLONIA AND PSILANTHELE (ACANTHACEAE) 279
Qg. Spines : simple ; branched. 10. Leaf-base : cuneate ; rounded. 11. Leaf-apex : emarginate ; not emarginate. 12. Leaf-apex : mucronate ; not mucronate. 13. Flowers : I or 2 ; more than 2. 14. Peditcel : shorter than calyx ; longer than calyx. 15. Pedicel : pubescent ; glabrous. 16. Sepals : subulate ; linear. 17. Corolla : pubescent outside ; glabrous. 18. Corolla-tube: to 4:5 mm. ; more than 4:5 mm.
MULTI-STATE (QUALITATIVE)
19. Spines : deflexed or curved ; horizontal ; ascending. 20. Leaf: broadest below middle ; at middle ; above middle. 21. Corolla: white ; blue ; red.
QUANTITATIVE
22. Leaf length,
maximum ;_ less than I2 mm. ; 12-25 mm. ; more than 25 mm. 23. Leaf breadth,
maximum: less than 6 mm. ; 6-18 mm. ; more than 18 mm. 24. Sepallength: 2-2:5 mm. ; 2-5-4 mm. ; more than 4 mm.
A few specimens, being incomplete, did not provide all the information desired and these deficiencies, it was understood, had later to be kept in mind since they might affect not only the subsequent placing of such specimens but also the linkage of others ; thus Nos. 20 and 24 lacked flowers.
Computation of this data was made by Mr. G. J. S. Ross at the Rothamsted Experimental Station, Harpenden, Herts. using an Orion electronic computer. The resulting classification, as expressed in a dendrogram (cf. Text-fig. 6) and a minimum spanning tree (cf. Text-fig. 7), agreed in general with that obtained by other means and undoubtedly would have been very helpful if initially available. The anomalous placing, however, of some specimens led to re-examination of their coding. This showed that a few attributes had been originally defined somewhat ambiguously or too arbitrarily ; thus some spines intermediate in length between the undoubtedly long spines of O. tetrasticha and the undoubtedly short spines of O. microphylla had been classified as ‘‘long’’ for some specimens but “ short ’”’ for others, thereby causing inconsistency and so making non-valid or exaggerated distinctions ; more- Over a few attributes, it must be confessed, had been erroneously coded. Since variability within a population may not be evident from the relatively few specimens available in herbaria for some groups, characters were taken, from the individual specimens, which a priori might have been of taxonomic value but which are now known to vary from individual to individual within a population, e.g. the presence of hairs on two or four sides of the stem, on the pedicels and on the outside of the
280 A SURVEY OF THE TROPICAL GENERA
corolla. In asmall sample, using few characters, these may have a distorting effect and, when the aim is to define populations, would seem best ignored. In the dendro- gram a specimen (No. 6) of the very distinct O. jamaicensis was not associated as closely with Nos. 4 and 5 as it should have been, simply because it had pubescent instead of glabrous shoots and flowers. The anomalous placing of specimens caused by the unsatisfactory coding of a few characters indicates the closeness of affinity
70 80 90 |
35 grandiflora a 6 jamaicensis r 5 jamaicensis 4 jamaicensis 32 armata 23 microphylla 11 armata 27 armata 10 spinosa tetrasticha 25 tetrasticha = 8 tetrasticha — 22 spinosa
16 microphylla | 15 microphylla 17 microphylla 14 microphylla 13 microphylla 33 spinosa 9 spinosa — 28 spinosa 26 spinosa 21 spinosa
20 tetrasticha?
36 purpurascens —— 24 nannophylla a 19 armata 18 armata 29 armata 12 armata aot 30 tetrasticha
7 tetrasticha 34 acicularis acicularis 4 _r— 2 acicularis
t— 1 acicularis
Fic. 6. Dendrogram showing overall phenetic similarity of 36 specimens of Oplonia (listed by numbers on p. 276) as computed from a matrix of 24 characters by G. J. S. Ross.
OPLONIA AND PSILANTHELE (ACANTHACEAE) 281
S 10 A 27 31 T 24 36 20 28 35 T © ° © . 8 N P T? s iS Ks + 30 M 17 wa M - i = 13 14 Pd J J mu 9ls « at
Fic. 7. (A). Minimum spanning tree uniting 36 specimens of Oplonia (listed by numbers on p. 276), the lengths of segments indicating phenetic distances between specimens as computed from a matrix of 24 characters by G. J. S. Ross ; A, specimens classified as O. armata ; Acas O. acicularis ; G. as O. grandiflora ; J as O. jamaicensis ; M as O. micro- phylla ; N as O. nannophylla ; Pas O. purpurascens ;S as O. spinosa ; T as O. tetrasticha.
at 29 \ | / < 1 / Wg =aV; --- Vz --- Ad _
wv A a
(B). Part of above minimum spanning tree converted into pictorialized scatter dia- gram ; A leaves broadest below middle, 0 at middle, vy above middle ; ; upward, spines ascending, | downward, spines downwards curved,—spines horizontal ; @ flowers white.
282 A SURVEY OF THE TROPICAL GENERA
within the group. It also serves as a warning of the narrow margin of error in treat- _ ing closely knit groups and the need for care in defining attributes so that essentially — similar ones are similarly coded ; inconsistencies in these as well as slight errors of — observation or recording may have their effect disproportionately magnified through — lack of compensating other characters within a low range of characters. Taking, however, the dendrogram (Fig. 6) as a whole, it emphasizes the distinctness of the — Peruvian O. grandiflora (No. 35) from the Caribbean group and it brings together enough specimens of O. acicularis (Nos. I, 2, 3, 34), of O. armata (Nos. 12, 18, 19, 29), of O. jamaicensis (Nos. 4, 5, 6), of O. microphylla (Nos. 14, 15, 16, 17), of O. spinosa (Nos. 9, 21, 26, 28, 33), and of O. tetrasticha (Nos. 8, 25, 31, and 7, 30) to form a basis for classification, though it is not acceptable as a whole without re-allocation of some specimens.
A minimum spanning tree (cf. Gower & Ross, 1969 ; Ross in Cole, 1969) constructed from the same data provides alternative pictorial expression of phenetic linkage and distance (Text-fig. 7) when, as here, principal co-ordinate analysis does not help very much because the first five latent roots are nearly equal and a plain vector diagram (Text-fig. 8) consequently has no clearly informative pattern and gives a confused impression. It should be noted that, in a minimum spanning tree, the linkages be- tween units and the distances separating them along the branches are significant, not the direction and relative spacing of branches which are simply a matter of conven- ience in drawing the diagram. Thus, in Text-fig. 7A, Nos. 1, 2, 3 and 34 are closely and directly linked but their proximity to Nos. 36 and 22 is purely coincidental and has no significance, because these are only distantly and indirectly linked, i.e. through No. 12. Here O. grandiflora (No. 35) is again given an isolated position. O. jamaicensis (Nos. 4, 5 and 6) links on to O. spinosa (Nos. 22, 26, 21, 33, 9) and so to O. armata (Nos. 19, 12, 29, 27, 18, II, 32) which has a central pivotal place in the genus and from which some others such as O. acicularis (Nos. I, 2, 3, 34), O. tetrasticha- (Nos. 7, 30, 25, 31, 8) and O. microphylla (Nos. 17, 13, 14, 16, 15) as well as O. nanno- phylla (No. 24) and O. purpurascens (No. 30) could be derived. Anomalously placed specimens are No. 20 (incomplete, now referred to O. tetrasticha with doubt), No. 28 (now referred to O. sPinosa), No. 10 (now referred to O. spinosa) and No. 23 (now referred to O. microphylla). .
A minimum spanning tree or principal co-ordinate analysis graph can be converted into a pictorialized scatter diagram more helpful for correlations of characters simply by adding to each position representing a specimen certain symbols indicating character states likely to have taxonomic importance, e.g. a triangle for leaves broadest below the middle, an inverted triangle for those broadest above the middle, a red mark for red flowers, a blue one for blue, lilac or purple flowers, a dot for white ones, small symbols for small leaves, larger for large leaves, etc. (cf. Text-fig. 7B).
To utilize the results of these computer-produced diagrams in practical taxonomy the simplest procedure is to lay out the specimens on a large table in a sequence or disposition corresponding to the diagrams and then by scanning and study to find the major characters diagnostic of the groups and the discontinuities between them. Marked discontinuities are almost nonexistent in Oplonia but the minimum spanning tree (Text-fig. 7A) can then be seen to have eight major elements. Excluding No. 35
OPLONIA AND PSILANTHELE (ACANTHACEAE) 283
(O. grandifiora) and Nos. 5, 4, 6 (O. jamatcensis) on account of their very large leaves and No. 24 (O. nannophylla) for its condensed growth, the most evident common denominator for the specimens assembled in each other group is the character of the spine, even though in coding this accounted for only 4 of the attributes. Using spine characters for diagnosis and now considering also the provenance of the specimens,
T+3 ©30 036 024 016 4 Saale 29 = 014 12 oi re e ; e @ o7 tee, 20 13 03 034 32 Ce Oo er ee ey 7 el LJ aa | Je 330 «=T el] e?] + e022 ©26 027 35¢e 6e 5e + @4 1-6
Fic. 8 Vector diagram showing lack of easily discernible patterns of correlated variation in 36 specimens of Oplonia (listed by numbers on p. 276) as computed from a matrix of 24 characters by G. J. S. Ross ; the vertical co-ordinates are the vectors corresponding to the first latent roots, the horizontal co-ordinates those of the second latent roots, of the trans- formed matrix.
No. 28 from Hispaniola with downward-curved spines and No. 10 from the Virgin Islands with some spines likewise downward-curved fit into the spinosa group and No. 23 from Hispaniola with fine ascending spines fits into the microphylla group. The same result would have been attained had spine characters received special weighting when coded.
284 A SURVEY OF THE TROPICAL GENERA
Long experience has convinced taxonomists that characters are of unequal value ~ in classification ; to produce a reasonable systematic arrangement by taxonometric — methods they could well be unequally weighted, but unfortunately such weighting ~
cannot usually be made before decision as to the nature of the group. Hence, since the main purpose of these methods is to aid decision-making, a priori weighting or advance decision would reduce their effectiveness ; characters need to be weighted a posteriori according to their distribution in the groups formed on the basis of overall similarity. Apparently the best course would be to accept as a basis the main groupings produced from an unweighted coding, then to investigate the content of these groups and modify them, if need be, according to the concepts of unequal taxonomic value of characters which emerge from this study.
GEOGRAPHICAL DISTRIBUTION AND ECOLOGY
The situation thus demonstrated in Oflonia parallels in some respects that de- scribed for the perennial species of Ziziphora (Labiatae) by I. C. Hedge (1961). The general habitat of these plants of the Near and Middle East is arid steppe and dry alpine slopes ; that of Oplonia is generally dry rocky slopes. In Ziziphora the plants are small-leaved and have small zygomorphic flowers showing sexual states which had not previously been recognized as such, again much as in Oplonia. The occur- rence within a single Ziziphora population of forms having very diverse hair-covering, together with scarcely correlated differences in habit of growth, proportion of leaf, length of calyx and shape of calyx-teeth, led Hedge to the conclusion that, despite some forty specific names applied to these plants, they form one actively evolving genetic unit in which certain phenotypic trends are apparent but are not yet marked by discontinuities justifying their specific or infra-specific designation. He found it impossible to construct a satisfactory key to the variants of Ziziphora but accepted a number of reference points within the multiplicity of trends in variation. This likewise is largely true of Oplonia. Many combinations or intergradations of char- acter are possible but some are represented by few or no individuals. Zzziphora inhabits a continuous landmass extending from Bulgaria over Turkey, the Caucasus and northern Persia to the Western Himalaya and the Altai mountains. The main territory of Oflonza is the Antillean chain of islands, most of which have long been separated from one another and others, e.g. the Lesser Antilles, have always been independent (cf. Alston, 1952 ; Schuchert, 1935). Thus conditions here favour differentiation. Cuban species, such as Oplonia nannophylla and O. tetrasticha, and Jamaican species, such as O. acicularis, O. armata and O. jamaicensis, individually possess morphological features unknown in the other islands. There are, however, forms extending from the Bahamas to the Virgin Islands and from Cuba to the Grenadines which do not make a homogeneous series or even a cline and do not break up into consistently distinct populations, but which include here and there forms so different that considered apart from the whole assemblage they would seem to merit specific rank and have indeed sometimes been givenit. Ina genus of closely allied plants, independent homologous and parallel variation is to be expected ; thus similar-looking plants of different immediate phylogeny may occur in areas remote from one another. The classification adopted here is, it is hoped, less arbitrary than
OPLONIA AND PSILANTHELE (ACANTHACEAE) 285
Lindau’s but is likely to be arbitrary nevertheless in that, by attempting to assemble specimens linked by overall resemblance into groups which can be recognized by a -few fairly well-marked and constant features and be named according to the nomen- clatural types coming within their circumscription, no account is or can be taken of parallel or convergent evolution ; moreover not all specimens manifest clearly the chosen diagnostic features. With these reservations the genus, as stated above, has been divided into 14 species, which although not always easy to distinguish seem to have geographical as well as morphological justification.
The genus Oflonza as defined and classified here consists of one species (O. grandt- flora) in northern Peru, three species (O. acicularis, O. armata and O. jamaicensis) confined to Jamaica, three species (O. nannophylla, O. purpurascens and O. tetrasticha) confined to Cuba, two species of much wider Antillean range, i.e. O. spinosa, which extends from the Bahamas over Cuba, Hispaniola and Puerto Rico (but not Jamaica) to the Virgin Islands, and O. microphylla, which extends from eastern Jamaica to the Grenadines in the Lesser Antilles, and, by the inclusion of Forsythiopsis, also five species from Madagascar. They usually inhabit the drier regions of these islands, their normal habitats being on limestone cliffs or rocky slopes thinly covered with scrub forest and thus well-drained, open and sunny. The xerophytic features characteristic of most species, i.e. the marked development of spines and reduction of leaf size and inflorescence, accord with their ecology but are uncommon within the family Acanthaceae as a whole. It is essentially these features of reduction which separate them from the small-flowered species of Odontonema, such as O. nitidum, which are of mesophytic habit, with relatively large leaves, numerous flowers in racemes or narrow panicles and no spines. Psilanthele eggersii of Ecuador approaches closest to Odontonema. Within Oplonia itself, O. grandiflora of Peru, O. jamatcensis of Jamaica and O. acuminata of Madagascar approximate most to the presumed archetype of the group in having a mesophytic habit, relatively large leaves, no spines and several or many flowers in axillary fascicles which are evidently condensed racemes.
The occurrence of species showing relatively primitive features, as Pszlanthele eggersit and Oplonia grandiflora, within the region of northwestern South America (northern Peru, Ecuador and Colombia ; Text-fig. 9), which has been an area of major development for the Acanthaceae—thus 39 native genera of Acanthaceae, with _ 365 species, are recorded from Colombia by Leonard (1951-58)—points to this as being the area where Oflonia had its origin, even though it has reached its greatest morphological diversity in Cuba and Jamaica. It misses the Central American main- land entirely. In this it is far from unique phytogeographically. As Seifriz (1943) mentions, the plant life of Cuba shows great similarity to that of South America, less to Central America and little to North America, and the same is probably true of the other Greater Antillean islands, Jamaica, Hispaniola and Puerto Rico. There are certainly floristic links between South America and the Greater Antilles which are unlikely to have been made by way of a former land connection between Honduras and the Greater Antilles (regarding which cf. Schuchert, 1935 ; Asprey & Robbins, 1953). Thus the genus Neriacanthus (Acanthaceae) consists of N. grandiflorus Leonard in Colombia, N. lehmannianus (Lindau) Lindau in Colombia and Peru, and N. purdieanus Benth. in Jamaica. Heppiella (Gesneriaceae) has about ten species in
286 A SURVEY OF THE TROPICAL GENERA
tropical South America, none in Central America, H. corymbosa (Swartz) Urban in south-eastern Jamaica and H. cubensis Morton in Cuba (Oriente province). Huertea (Staphyleaceae) has one species, H. glandulosa Ruiz & Pavon, in Peru, none in Central America, and one, H. cubensis Urban, in Cuba and Hispaniola. Of the genus Purdiaea (Cyrillaceae), according to J. L. Thomas (1960, 1961), there is one species, P. nutans Planchon, in Peru, Colombia and Venezuela and another one, P. belizensis (Smith & Standl.) J. L. Thomas, in British Honduras, whereas Cuba possesses ten species, of which one, P. cubensis (A. Rich.) Urban, grows in the Isle of Pines and Pinar del Rio province and of which the other nine are endemic to Oriente province.
= wag
Fic. 9. Distribution of Oplonia and Psilanthele in tropical America ; 1, West Indian species of Oplonia ; 2, Oplonia grandiflora in Peru ; 3, Psilanthele eggersit in Ecuador. (Goode Base Map, University of Chicago).
Thomas (1960) states : ‘‘ there are several trends of specialization in Purdiaea which indicate that the South American species, P. nutans, is the most primitive species in the genus’’. The Melastomataceae provide further examples of floristic linkage. Thus Clhidemia grisebachii Cogn. of Jamaica and C. cruegerana of Trinidad and Venezuela are closely related, likewise Miconia nubicola Proctor of Jamaica and certain species of the Venezuelan coastal Cordillera and the Andes with no close relatives elsewhere (J. J. Wurdack in litt.). Hemnrtetta sessilifolia Triana occurs in Jamaica, Trinidad and Venezuela, H. ramiflora (Swartz) DC. in Jamaica, Trinidad, Venezuela, the Guianas and Brazil.
Such facts taken from a diversity of families speak for a former direct way of floristic migration between north-western South America and the Greater Antilles in which Central America had little or no part. Despite the limited collecting then available, Grisebach as long ago as 1865 alluded to such a ‘‘ besondere Wanderungs-
OPLONIA AND PSILANTHELE (ACANTHACEAE) 287
linie ’’ connecting the Andes of South America with the mountains of Jamaica and Cuba. Croizat (1952 : fig. 71), arguing largely from the distribution of some Cacta- ceae, also postulates a Peru—West Indies track across Venezuela direct to the Greater Antilles.
Moreover it seems improbable that the rich and diversified endemic flora of the Greater Antilles evolved on so limited an area as this flora now occupies and likewise improbable that the ancient floristic connections of the Greater Antilles and northern South America were made by long-distance leaping dispersal across the great expanse of the Caribbean Sea, although floating seeds carried by sea currents must have taken a number of coastal plants from island to island and light air-borne seeds, e.g. of
var, roar wees nee,
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7 CAYMAN ISLANDS
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SARBUDA
DOMINICA sure is q ; 4 SERRANA CAYS Q marriniaue be > CARIBBEAN SEA ostucia x BARBADOS gf Ssrvince Pes ARUBA yp GRENADINES - . {CURAGAS D GRENADA = oe
NO Roque aca DLANQUIELA a BONAIRE bd 2 TOBAGO aed TORTUGA SARS RITA
Fic. 10. Distribution of Oplonia nannophylla a ; O. spinosa ®@.
members of the Apocynaceae, have been swept over the area by high winds. In general, the biological facts seem to favour the hypothesis that “‘ until the end of the Cretaceous a continental landmass occupied the position of the present-day Caribbean Sea ”’ (cf. Weyl, 1965), and this landmass (Caribia) subsided during the Cretaceous and Tertiary periods. The submergence of almost the whole area, including Jamaica and Cuba, by Oligocene times would have led to the extinction of the major part of its flora but much of the present Greater Antillean flora may nevertheless be descended from the plants of this lost territory which presumably both contributed to and received from the flora of South America and contributed to that of Central America. Such a hypothesis provides a plausible background to facts of Caribbean phyto- geography at present otherwise inexplicable, among them the distribution of Oplonza. Croizat (1952 : 368) has indeed gone so far as to describe “‘ the tale of dispersal in the Caribbeans”’ as “‘ the record how a land once filling the extant sea crumbled front by front, thereby creating huge gaps in distribution. ”
288 A SURVEY OF THE TROPICAL GENERA
Jamaica and Cuba are the major islands of development for Oplonza. show different patterns of distribution (see Text-figs. 10-14).
The species
In Jamaica, O. microphylla (Text-fig. 12) inhabits the very hot and dry honeycomb limestone along the southern coast from St. Thomas to St. Catherine, which has a
'SLA OF PINOS EAT
cs NAGUA i cf 4 fo Turks is
Aer 1SLANDS - b t | a ,O@mMINICAN
cAICoSs om SQ
o t : 300 miles P + 500 Km GRAND v Bh y= zs cveurnera pay TORTUGAS ° 0 v e ANDROS s aris. . oe 2 San SaLvacg® boas Q (WaT on 13. r > &. PLANA CAYS * “ar Pos
swan oy JR BLIC i gga" \ 40, J 3 ot er st Laren te a. eke ant preaye HISPANIOLA av te" 2. roses 9 a sce Q maariniaue 4 oO" LUCIA a \ ented o Ow CENT — ARUBA RGRENADINGS ~ BONAIRE os TOBAGO gs poses nga 2 Fic. 11. Distribution of SnenE purpurascens @ ; O. microphylla @. 30° 78 ° 30° 77 - ay
JAMAICA
Fic. 12. Distribution of Oplonia acicularis © ; O. jamaicensis @ ; O. microphylla (in
Jamaica) @.
characteristic scrub forest flora described by Asprey and Robbins (1953). O. acicularis (Text-fig. 12), with an evident preference for moister and even somewhat shaded habitats, has a remarkably disjunct distribution, one area in the east and another in
————
OPLONIA AND PSILANTHELE (ACANTHACEAE) 289
the west but not in the middle of the island. O. jamazcensis (Text-fig. 12), considered the most primitive Antillean species, grows only in the moist interior upland region. O. armata (Text-fig. 13), as accepted here, presents much diversity in habit and leaf- form with geographical differentiation mostly in colour of the corolla. Red flowers occur only on plants inhabiting the interior (var. avmata), paler bluish or lilac flowers only on those nearer or at the coast (var. pallidior).
3" 78° 30" Te 30"
JAMAICA
5 ° < 10 is 20
30"
Fic. 13. Distribution of Oplonia armata ; var. pallidior ®@ ; var. armata ©.
50 100 150 miles
| =
Fig. 14. Distribution of Oplonia tetrasticha ; var. tetrasticha @ ; var. polyece .
Cuba divides conveniently into four botanical regions (cf. Conde, 1952, also Alain, 1953, 1958) ; 1 : Occidente, comprising the provinces of Pinar del Rio, Habana and Matanzas ; 2: Las Villas, the former province of Santa Clara ; 3 : Camaguey ; 4:Oriente. O. purpurascens (Text-fig. 11) is apparently endemic to the mogotes, the precipitous limestone “‘ haystack hills’”’ (cf. Seifriz, 1943 : 394 ; Lehmann, 1954) of Pinar del Rio around Vifiales. O. nannophylla (Text-fig. 10) occurs in Pinar del Rio, Habana and Las Villas. O. tetrasticha var. tetrasticha (Text-fig. 14), with simple spines, likewise occurs in Pinar del Rio, Habana, Las Villas and Oriente, but is largely
290 A SURVEY OF THE TROPICAL GENERA
replaced in Oriente province, rightly described by Croizat as a ‘“‘ hotbed of endemism”’ by O. tetrasticha var. polyece with branched spines. Oflonia has not yet been collected in Camaguey. In Oriente occur also plants referred to O. spinosa (Text-fig. 10) (see p. 307), a species which extends over the low-lying Bahama Islands northward to Grand Bahama and New Providence and eastward to the Virgin Islands. Within Oriente province it appears to be confined to the Moa region on the north coast.
O. microphylla (Text-fig. 11), which has not colonized the Bahamas, extends the range of the genus beyond the Greater Antilles to the Grenadines in the Lesser Antilles, which are of volcanic origin, but it does not reach Barbados, Tobago or Trinidad. The absence of the genus from these three islands, which lie on the continental shelf of north-western South America, as also from Venezuela, makes it as improbable that it migrated from South America by way of the Lesser Antilles as that it did so along the isthmian route leading to the presumed Antillean-Honduran land bridge.
From the above survey it would appear that the genus Oplonia may have had its origin in South America from mesophytic shrubs resembling Psilanthele eggersii or small-flowered species of Odontonema, that it reached the Greater Antilles by way neither of Central America and the Honduran land-bridge nor of the Lesser Antilles but by a former direct long-vanished link, and that its major development has taken place in Jamaica and Cuba with little change in floral structure but with increasing vegetative adaptation to xerophytic conditions.
PHYTOGEOGRAPHICAL LINKAGE OF TROPICAL AMERICA AND MADAGASCAR
A remarkable discontinuity of range associated with little morphological diver- gence becomes apparent when the American species of Oplonia are compared with the Madagascan species currently referred to Forsythiopsis. Since their resem- blances in habit, floral structure, capsule and pollen together leave no characters available for generic separation, they are here placed in the same genus despite their geographical remoteness. Use of different generic names for the American and Madagascan plants respectively has obscured their affinity. Baker would surely never have described Forsythiopsis as an independent genus in 1883 had he not then compared Baron’s Madagascan specimens, the type material of Forsythiopsis baront Baker, with Ruellia instead of with Anthacanthus (i.e. Oplonia), to which Nees von Esenbeck had already in 1847 referred Lamarck’s conspecific Justicia vincoides, likewise from Madagascar. Moreover, the pollen of Forsythiopsis, as is evident from that of the holotype of F. baront (Baron 1937), the holotype of F. linifolia (Humbert & Swingle 5507), an authentic specimen of F. australis (Scott-Elliott 270) and other specimens (McWhirter & Capuron 184, Humbert 2606) studied with the scanning electron microscope at magnifications between x 500 and 5,400, agrees with the pollen of West Indian Oflonia specimens ; it is prolate or spheroid, 3-colporate, each colpus flanked by a pseudo-colpus extending almost to the poles, the surface pitted with minute lumina (Pl. 41); it in no way resembles the reticulate pollen of Ruellia. Similar pollen occurs in Pseuderanthemum and also the continental African genus Ruttya, within which C. B. Clarke would have incorporated Forsythiopsis, but this has larger flowers with monothecous (unilocular) anthers (whence Hoch-
OPLONIA AND PSILANTHELE (ACANTHACEAE) 291
stetter’s name Haplanthera), possibly derived from a Pseuderanthemum- or a For- sythiopsis-like stock with dithecous (bilocular) anthers.
45° 46 48 50°
re 12 S ra’ 12
14 \ 14
18 18
e . Tananarive e
20°
22
24
25°
Statute Miles O 20 40 60 80 100
50°
Fic. 15. Distribution of Oplonia in Madagascar ; Oplonia vincoides @ ; O. minor var. minor @; O. minor var. meridionalis © ; O. minor var. vestita * ; O. linifolia mw; O. acuminata + ; O. puberula a.
In Madagascar (Text-fig. 15) the species grow mostly within Humbert’s phyto- geographical domains Sud and Ouest (cf. Blanc, 1971, for maps and climatological data), arid and semi-arid regions with open xerophytic scrub and savanna vegetation;
Cc
-~ gk
292 A SURVEY OF THE TROPICAL GENERA
they thus appear to have much the same ecological requirements as most West Indian species of Oflonia. Madagascar, with some 400 species of Acanthaceae to an area of 228,000 square miles, has been a major centre of development for the family, out. rivalling even Colombia, with some 370 species to 462,000 square miles. No species congeneric with Oplonia (including Forsythiopsis) are known from continental Africa. In manifesting such discontinuity Oplonia does not stand alone. There are other links, few but definite, between the floras of tropical America and Madagascar which testify to a much wider primaeval range of certain’ groups forming part of Perrier de la Bathie’s ‘‘ elément austral”’ in the Madagascar flora, “‘ témoins résiduels d’une flore antique en voie d’extinction ” (Perrier de la Bathie, 1936 : 139-140). Thus Ravenala (Musaceae) has one species, R. madagas- carvensis Sonnerat, the well-known “ Traveller’s Tree’’ in Madagascar ; its only close ally is a species of tropical America (Brazil, Guiana), R. guianensis (L. C. Rich.) Petersen ; the latter having five stamens (instead of six) can be put in a genus by itself as Phenakospermum guianense (L. C. Rich.) Miq., but, whether the two are treated as forming one genus with two species or as two monotypic genera, their affinity and their distinctness from all other genera remain beyond dispute. Oliganthes (Compositae), with 9 species in Madagascar and 12 in tropical America but none elsewhere, provides another example of such linkage, as do Clara (Herreria Ruiz & Pavon, non Adans. ; Liliaceae) with 5 or 6 species in South America and Herreriopsis with one species in Madagascar. Gladiopappus (Compositae) of Madagascar is allied exclusively to the tropical American genera Richterago (Seris Less. non Willd.), Hyalis and Hecastocleis. These anomalous ranges only become explicable when considered in relation to those of genera or groups of closely allied genera occurring not only in tropical America and Madagascar but also in continental” Africa. For example, the genus Sabicea (including Stipularia; Rubiaceae) has about 50 species in tropical America (including Peru, Ecuador, Colombia and the West Indies), 85 in continental tropical Africa and 5 in Madagascar. A year before the publication of Alfred Wegener’s book on continental drift, and long before it had become generally known, characters common to various American and African species of Sabicea led Wernham (1914) to the opinion that “‘ the genus existed simul- taneously in Africa and America as an ancestral race (Primosabicea) with lax in- florescence, represented at the present day by e.g. S. venosa in Africa and S. paraensis in America ”’. It has maintained a continuity of range in Africa, unlike, for example, the genus Hirtella (Chrysobalanaceae) with approaching 100 species in tropical America, 2 in tropical East Africa, of which one occurs in Madagascar but none in West Africa. Mendoncia (Acanthaceae) has about go species in tropical America, 4 in tropical West Africa (but not East Africa) and 3 in Madagascar. Symphonia (Gutitferae) is remarkable in having one species (S. globulifera L.f.) common to tropical America and tropical West and Central Africa and 15 species recorded from Madagascar. An even more remarkable range has been attained and maintained by Christiana africana DC. (Tiliaceae ; cf. Kew Bull. 17 : 501 (1964)), which occurs in tropical America, tropical West, Central and East Africa and Madagascar ; the one other species of the genus is Brazilian. Genlisea (Lentibulariaceae), with I5 species, also occurs in tropical America, tropical continental Africa and Mada-
*
OPLONIA AND PSILANTHELE (ACANTHACEAE) 293
gascar. Savia (Euphorbiaceae) has two main centres of diversity, with 12 species in the West Indies and 9 in Madagascar, but their discontinuity is slightly lessened by 2 species in Brazil and one in South Africa ; one species extends into southern U.S.A. The Velloziaceae display a likewise significant pattern of distribution linking tropical America, continental Africa and Madagascar, as noted, for example, by Camp (1947). The family consists of Vellozia and Barbaceniopsis, which are exclusively tropical American, and Barbacenia (sensu lato, including Xerophyta) which has about 75 species in tropical America, about 40 species in continental Africa, 3 in Madagascar, one in southern Arabia. Several consistent features distin- guish American Barbacenia (sensu stricto) from African Xerophyta ; one African species (Vellozia elegans Oliver of Natal), in the opinion of L. B. Smith and E. Ayensu (in litt.), constitutes an intermediate monotypic genus (Zalbotia Balf. f., non S. Moore) combining several primitive features, which suggest that the family originated in Africa ; but whether the African species are included in Barbacenia or separated from it generically is irrelevant from a phytogeographical standpoint in view of the undoubted affinity of the American and African groups. A somewhat like affinity is that of Aphelandra (Acanthaceae), having about 200 species in tropical America, with Stenandriopsis having 5 species in tropical West Africa, one in East Africa and 8 in Madagascar (cf. Heine in Aubréville, Fl. Gabon 13 : 99 ; 1966). The genera Agarista (Evicaceae) and Agauria provide another example ; Agarista (cf. Stevens, 1970) has about 34 species, mostly in South America but with a few in Central America and the southern United States ; the closely allied genus Agauria, according to Sleumer (1938), consists of one polymorphic species divisible into 20 intergrading infraspecific taxa, a few occurring widely separated in West Africa, more on the East African mountains, most of them in Madagascar.
Equally significant is the existence of other groups connecting tropical America and Madagascar which have more restricted ranges evidently resulting from the
fragmentation and reduction of former more extensive ranges. Thus the genus
Erblichia (or Piriqueta sect. Erblichia ; Turneraceae) has one species in Central America, one in southern Africa, three in Madagascar. Another phytogeographic- ally relevant group is that formed of three closely allied monotypic genera of Composttae described by Lawralrée (1943), i.e. Eleutherantha in tropical America,
-Hoffmanniella in Cameroons and Congo, Exomiocarpon in Madagascar. A rather
similar linkage is to be found in the Dichapetalaceae with Stephanopodium in tropical America, Tapura in tropical America and continental Africa, Falya in Madagascar. The Canellaceae, one of the nine exclusively Amerafran families*, exemplifies the same general floristic linkage, with Canella, Cinnamodendron and Pleodendron in tropical America, Warburgia in tropical East Africa and Cinnamosma in Madagascar. The genus Carpodiptera (Tiliaceae) does not reach Madagascar but nevertheless manifests a distribution of the same type as Hirtella by having species in Central America and the West Indies, tropical East Africa, and the Comoro Islands
* Angiosperm families occurring both in tropical America and Africa but not known elsewhere are Bromeliaceae, Canellaceae, Caricaceae, Humiriaceae, Hydnovaceae, Mayacaceae, Rapataceae and Vochysi- aceae (cf. Vester, 1940 : 527-529, 567-569, Maps 238-241, 245-248). Loasaceae and Velloziaceae occur in
tropical America and Africa but extend from east Africa into southern Arabia (cf. Vester, 1940 ; maps 243,244). To these can be added Strelitziaceae if separated from Musaceae.
294 A SURVEY OF THE TROPICAL GENERA
north of Madagascar (cf. map in J. Hutchinson, Evolution and Phylogeny of Flowering Plants : 232 (1969)). .
There are also families and genera occurring both in tropical America and West Africa (cf. Hepper, 1965) which do not reach Madagascar but nevertheless emphasize the floristic affinity of the two continents. One such is Cienfuegosia (cf. Fryxell, 1969) in the Malvaceae with 16 American and 8 African species.
Taken together these diverse ranges form a series manifesting stage by stage the reduction from a wide continuous range, such as that of Sabicea (America, conti- nental Africa, Madagascar), by way of intermediate discontinuous ranges, such as those of Erblichia and Hirtella, to the extremely discontinuous ranges of Ravenala, Oliganthes and Oplonia. These floristic linkages must have had their origin in the far distant past, presumably long before the end of the Cretaceous period, when the continuity of the Amerafran landmass (cf. maps in Bullard, Everett & Smith, 1965 ; Smith & Hallam, 1970 ; Dietz & Holden, 1970), later separated into South America and Africa, still provided the opportunity for (or the narrowness of the South Atlantic Ocean made no obstacle against) the spread of extreme western- originated groups eastward across that vast landmass, the interior of which must have been very dry, however, before separation, and the spread of eastern-originated groups westward, since climatic conditions were presumably more favourable for the evolution and preservation of diversity towards the periphery than towards the centre. Ifso, the groups concerned have kept their generic characters, or at least many characters indicative of affinity, essentially unchanged for probably go million years or more ; indeed Hawkes and Smith (1965) have argued from the taxonomy and distribution of Bromus, Gossypium and Solanum that these genera existed already in the early Cretaceous, some 100 to possibly 135 million years ago.
Madagascar has certainly been isolated for a long time. If, as stated by Flories (1970), the rift which became the Mozambique Channel separating Madagascar from continental Africa had been completed by the end of the Cretaceous period, i.e. some 70 million years ago, at which time modern orders and families of angiosperms were already differentiated (cf. Muller, 1970), then the ancestors of the Madagascan plants whose nearest allies now inhabit tropical America must have already acquired the characters still common to the Madagascan and American plants or at least a close genetical similarity determining later parallel development. This conclusion, if correct, implies the origin of the Acanthaceae long before that part of the Miocene epoch, some 12-20 million years ago, to which belongs the earliest available palaeontological evi- dence of its existence, i.e. pollen of the Justicia-type named Multiareolites formosus and of the Tvichanthera-type named Multimarginites vanderhammenit (cf. Germeraad et al., 1968: 302, t.6) from Venezuela; phytogeographical evidence indicates a Cretaceous origin.
Acceptance of the view that the angiosperms originated on the Gondwanaland continent and that many genera existing today had acquired their distinctive characteristics before it broke into several continental landmasses (cf. Melville 1966 ; Schopf, 1970) harmonizes many otherwise seemingly inexplicable facts of angiosperm distribution. The drifting apart of Africa and South America, together with the associated far-reaching climatic disturbance, must have led to the extinction
OPLONIA AND PSILANTHELE (ACANTHACEAE) 295
or reduction of range of some groups while promoting the development and spread of others. Evidently those groups such as Oliganthes and Ofplonia, now with representatives in tropical America and Madagascar but not elsewhere, owe the isolation of these to continental drift which divided the original stock into American and African populations, to the spread of the African population across Africa and over to Madagascar or of a Madagascar-originated population across Africa to America, to climatic changes, which led to extinction in continental Africa, and to peripheral survival in America and Madagascar, with the larger area and more varied habitats of tropical America favouring greater specific development and range extension.
ACKNOWLEDGMENTS
The preparation of this survey of Oflonia and Psilanthele would probably have been simpler had a revision of the whole group been planned from the start instead of undertaken reluctantly as the only way of elucidating nomenclatural problems affecting Jamaican species. No attempt has been made to borrow for study all the material available from islands other than Jamaica. My thanks are due to the directors and curators of the following institutions for the privilege of examining specimens in their charge : the Arnold Arboretum of Harvard University, Cambridge, Mass. (cited as A) : the Field Museum of Natural History, Chicago (F) ; the Gray Her- barium of Harvard University (GH) ; the Science Museum, Institute of Jamaica, Kingston, Jamaica (IJ) ; the Royal Botanic Gardens, Kew (K) ; Herbario de la Salle, Habana (LS ; temporarily deposited in NY) ; the Institute Botanique, Université de Montréal, Montreal (MT) ; the New York Botanical Garden, New York (NY) ; Laboratoire de Phanérogamie, Muséum National d’Histoire Naturelle, Paris (P) ; the Botanical Department, Naturhistoriska Riksmuseum, Stockholm (S) ; Depart- ment of Botany, University of the West Indies, Mona, Jamaica (UCWI).
I am also grateful to Brother Alain (H. Liogier), formerly of the Colegio de la Salle, Habana, for details of distribution in Cuba, to the Director, Botanisches Museum, Berlin-Dahlem, for photographs and to Dr. G. Wagenitz for critical details of the types of Evanthemum emarginatum and E. lyciotdes in the Willdenow Herbarium, to ' Professor G. Erdtman, Stockholm-Solna, for preparations of pollen grains, to the late Mr. J. B. Evans, British Museum (Natural History), also for preparations of pollen _ grains and of cleared leaves, to Dr. José de Js. Jiménez, Santiago de los Caballeros, for specimens from the Dominican Republic, and to Mr. G. R. Proctor, Institute of Jamaica, for companionship and help on many a field trip in Jamaica and for much information about its floristics and geography. Dr. H. Heine and Prof. J. Leandri gave valuable help in mapping the distribution of Oflonia in Madagascar.
The taxonometric section of this paper owes its existence to the co-operation of Dr. P. H. A. Sneath, British Medical Research Council, Mr. J. G. Gower and Mr. G. J. S. Ross of the Rothamsted Experimental Station, Harpenden, Herts., England and Professor D. J. Rogers, University of Colorado, Boulder, Colorado, U.S.A. ; their computation and analysis of the data supplied and preparation of diagrams have proved stimulating and informative and this acknowledgement is only a mild expression of gratitude for their help.
The photographs of pollen grains illustrating this paper have been made with the
296 A SURVEY OF THE TROPICAL GENERA
““ Stereoscan ’’ electron microscope in the Electron Microscopy Unit, British Museum ~ (Natural History) and I thank Miss P. A. Sims and Dr. S. Sengupta for their valuable technical assistance in this and other palynological work.
Miss Beatrice M. Corfe drew Text-fig. 1, Miss Joanna C. Webb (Mrs. D. Erasmus) — Text-figs. 2 and 3, Mr. Derrick Erasmus Text-figs. 4 and 16 and Miss Victoria — Goaman Text-figs. 17 and 18.
PSILANTHELE Lindau PsILANTHELE Lindau in Bull. Herb. Boiss. 5 : 663 (1897).
Shrubs without spines. Leaves small, shortly petiolate, entire ; cystoliths present. Flowers small, pedicellate in short-pedunculate few-flowered loose raceme-like cymose inflorescences terminating short axillary branches. Bracts minute. Calyx 5-parted, c.2 mm. long, the tube very short, the segments almost
Fic. 16. Flower of Psilanthele eggersit Lindau (Eggers 15129) cut lengthwise.
equal, narrow, acute. Corolla zygomorphic, bluish, c.5 mm. long ; tube narrow, cylindric, almost straight ; limb spreading, 2-lipped, the upper lip shortly 2-lobed, the lower lip 3-parted, the segments almost equal. Stamens 2, inserted above the middle of the corolla-tube ; filaments short ; anthers with 2 spurless equal disjunct loculi ; staminodes 2, extremely minute. Pollen prolate, 3-colporate, each colpus flanked by 2 pseudocolpi, reticulate with minute lumina. Ovary with 2 ovules in each loculus ; style relatively long. Capsule clavate, gape 4-seeded. Type- species by monotypy : Psilanthele eggersti Lindau.
Only known species the following : 1. Psilanthele eggersiiLindauin Bull. Herb. Boiss. 5 : 664 (1897).—Mildbraed apud
Diels in Bibl. Bot. 116 : 148 (1937). (Text-fig. 16 ; Pls. 40a, 46). Shrub to 2 m. high ; branches slender, ascending, sparsely and minutely pubescent,
OPLONIA AND PSILANTHELE (ACANTHACEAE) 297 | red-brown (fide Diels) when young, becoming yellowish with age. Leaves short- | stalked, the blade narrowly elliptic or lanceolate, the apex acuminate, the base | attenuate, c. 1-5~5 cm. long, 5-15 mm. broad, glabrous and with numerous cystoliths | above, glabrous below, thin, the veining evident. Flowers in 16~-20-flowered inflorescences I-2 cm. long ; pedicels 2-3 cm. long. Calyx c.2 mm. long, the seg- ments very narrowly triangular. Corolla blue (fide Lindau) or white with red streaks on lower lip (fide Diels), minutely pubescent, c.5 mm. long, the tube 2-5 mm. long, the segments 2:5 mm. long. Stamens almost included, the filaments 0-7 mm. long, the anthers 1 mm. long.
Ecuapor : Manali: probe Hacienda El Recreo, 16 July 1898, Eggers 15129 (F ; K ; isotypes). Chimborazo : im Gebiet des R. Chanchan bei Naranjapata, 530m., Sept. 1933, Diels r1g2 (fide Mildbraed, loc. cit).
The above account is based on Lindau’s original description and isotypes in the Kew and Chicago herbaria. As the type was collected by Baron Henrik Franz Alexander Eggers (1844-1903) in western Ecuador near the coast at El Recreo (0° 27’S) south of Cape Pasado, and Mildbraed records it from central Ecuador at Naranjapata (2° 17'S), it would seem to be a species widely distributed in Ecuador but rarely collected, the flowers being inconspicuous and its habit of growth in no way remarkable. Eggers was a Danish professional soldier, who served in Mexico and the Danish West Indian islands (St. Croix, 1869-72, 1873-74, St. Thomas, 1872-73, 1874-1885) and who managed while so engaged to collect botanical speci- mens on many West Indian islands. In 1891-99 he travelled and collected in South America (Venezuela, Ecuador). An account of his career will be found in Urban, Symb. Antill. 3 : 40-43 (1902).
A photograph of the holotype (formerly in Berlin-Dahlem) is available in Field Museum of Natural History, Chicago, photograph 8737.
OPLONIA Raf. OpLonié Raf., Fl. Tellur. 4 : 64 (1838).
Evanthemum c. Hesperanthemum Endl., Gen. Pl. : 706 (1839).
Anthacanthus Nees in DC., Prodr. 11 : 460 (1847) nom. superfi., excl. § 2.—Griseb., Fl. Brit. W. Ind. Is. : 457 (1862).—Benth. in Benth. & Hook., Gen. Pl. 2 : 1097 (1876).—Lindau in Engler & Prantl, Nat. Pflanzenfam. iv. 3b : 335 (1895) ; in Urban, Symb. Antill. 2 : 214 (1900).—Britton & Millsp., Bahama FI. : 402 (1920).—Britton & Wils. in Sci. Surv. Porto Rico & Virgin Is. 6 : 212 (1925).—Alain in Leén & Alain, Fl. Cuba 4 : 498
(1957).
Forsythiopsis Baker in Journ. Linn. Soc. Lond., Bot. 20 : 218 (1883).—Benoist in Humbert, Fl. Madag. 182 : 106 (1967).
Hesperanthemum (Endl.) Kuntze, Revis. Gen. Pl. 2 : 490 (1891), nom. superfi.
Shrubs or small trees, mostly with opposite axillary spines, sometimes spineless. Leaves usually small, shortly petiolate or almost sessile, entire, often fascicled when two or more pairs arise on the same extremely short axillary branch ; cystoliths present, often abundant. Flowers small, pedicellate, solitary and axillary, or in sessile or short-pedunculate axillary fascicles. Bracts minute. Calyx 5-parted, 3-6 mm. long, the tube very short, the segments almost equal, narrow, acute.
298 A SURVEY OF THE TROPICAL GENERA
Corolla zygomorphic, white, bluish, purple or red, 8-18 mm. long ; tube narrow, ~ cylindric, almost straight, slightly expanded above ; limb spreading, 2-lipped, the upper lip 2-lobed, the lower lip 3-parted, the segments almost equal ; aestivation imbricate. Stamens 2, inserted above the middle of the corolla-tube, included or exserted ; filaments slender, short or long ; anthers with 2 parallel spurless equal — loculi ; staminodes 2, minute. Pollen spheroid or prolate, 3-colporate, each colpus flanked by 2 pseudocolpi, pitted with minute lumina. Ovary with 2 ovules in each loculus ; style long, the stigma slightly thickened and almost entire. Capsule clavate, stipitate, 2—4-seeded. Seeds somewhat flattened, almost circular in outline, 2-3 mm. broad, borne on acute retinacula.
Type-species by original designation : Oplonia spinosa (Jacq.) Raf. based on Justicia spinosa Jacq.
Species variable and difficult to define, fourteen accepted here, one from Peru, eight West Indian, ranging from the Bahamas and Cuba to the Grenadines in the Lesser Antilles, and five Madagascan.
As Brizicky (1969) has pointed out, it is impossible to decide the rank of Endlicher’s subdivisions of genera.
KEY TO SPECIES OF OPLONIA
Spines up to 3 cm. long and mostly more than 1-5 cm. long, spreading horizontally or
slightly ascending, straight, rigid ; Cuba : ; : 6. O. tetrasticha Spines all unbranched : : . 6a. var. tetrasticha Spines mostly bearing 2-4 subsidiary ne : ; ; 6b. var. polyece
Spines absent, or, if present, mostly less than 1-2 cm. long, or, if longer, deflexed or curved or very slender : Spines present, some or all slightly or distinctly curved, spreading almost horizont- ally or recurving ; Bahamas to Virgin Islands : 4. O. spinosa Spines absent or, if present, straight and spreading horizontally or ascending : Shrub of very dwarf habit, 4-8 cm. high, with internodes 1-3 mm. long; Cuba. 9. O. nannophylla Shrub taller and looser in habit, the internodes more than I cm. long : Pedicels shorter than the calyx: Spines present ; western Cuba . ; : 8. O. purpurascens Spines absent ; Madagascar. : . : : 13. O. minor Pedicels some or all longer than the ee Corolla white ; stamens always included, the filaments c.o-8 mm. long, shorter than the anthers ; low spreading shrub under 60 cm. high, with leaves mostly broadest below the middle ; Jamaica 7. O. acicularts Corolla red, purple, pale violet or blue, very rarely white ; stamens exserted or included, the filaments 1-5~7 mm. long, longer than the anthers ; shrubs mostly 1-2 m. or even small trees to 6 m. high, with leaves variable but often broadest at or above the middle: Leaves all very small (not more than 12 mm. long, 6 mm. broad): Leaves linear; Madagascar . ; : . . 14.0. linifolia
OPLONIA AND PSILANTHELE (ACANTHACEAE) 299
Leaves elliptic to narrowly obovate: Spines present ; Jamaica to the Grenadines . 5. 0. microphylla Spines absent ; Madagascar 12. O. minor
Leaves larger, some or all more than 12 mm. long: or, if less, then broadest
at or below the middle:
Apex of leaf acuminate or acute ; calyx segments about as long as
the 4:5-5-mm. long corolla tube:
Flowers 7-10 at each axil ; leaf blade narrowly cuneate at base ;
Peru ; : oy ee oe grandiflora Flowers 1-3 at each arate mleag blade roduded at base ; Madagascar . : 10. O. acuminata
Apex of leaf usually obtuse o or rounded: pirels: acute ; calyx segments shorter than the 5—10-mm. long corolla tube: Corolla segments longer than tube ; style hairy ; Madagascar: Mature leaves mostly 3-9 cm. long: Calyx segments narrowly triangular, sparsely hairy. 11. O. vincoides Calyx segments narrowly oblong, densely hairy. 12. O. puberula
Mature leaves less than 3 cm. long . ’ : 13. O. minor Corolla segments slightly shorter than tube ; style glabrous; Jamaica:
Leaves mostly 3-10 cm. long, 1-8—4:5 cm. broad ; flowers in each fascicle numerous Ses ; interior of Jamaica, mostly above 500m. . ; ; 2. O. jamaicensis
Leaves smaller, mostly 1-2-5 cm. iene. 0-4-2 cm. broad, if longer then broadest above the middle and flowers solitary ;
flowers solitary or few (2-4) in each fascicle. 3. O. armata Corolla red ; leaves often broadest above the middle; plants of the interior . : : . . 3a. var. armata
Corolla pale purple, lilac or almost white: leaves mostly broadest at the middle ; plants occurring below 500 m., usually coastal 3b. var. pallidior
. _—
I. Oplonia grandiflora (Lindau) Stearn, comb. nov. (Pls. 40b, 42). Psilanthele grandiflora Lindau in Bull. Herb. Boiss., Sér. 2, 4 : 401 (1904).
Shrub 1-4 m. high ; branches stout, greyish, at first sparsely pilose with long hairs, later glabrous ; spines absent. Leaves short-stalked, the blade very variable in size in the same fascicle, mostly lanceolate, sometimes narrowly elliptic, the apex acute, the base narrowly cuneate, c.2—10 cm. long, 0-5-3 cm. broad, glabrous except for sparse more or less appressed hairs on the veins below and along the margin, the veining evident with 6-8 lateral veins each side of the midrib ; petiole 5-10 mm. long. Flowers 7-10 in sessile axillary fascicles ; pedicels 1-2 cm. long, glabrous. Calyx 45-6 mm. long, the segments linear, acute. Corolla white with violet markings (fide
300 A SURVEY OF THE TROPICAL GENERA
————
E. Ule), glabrous, the tube 4-4-5 mm. long, the upper lip slightly 2-lobed, c. 5 mm. —
long, the lower lip 3-parted with obtuse segments to 6:5 mm. long, 4 mm. broad. Stamens included, the filaments c. I mm. long, the anthers scarcely 1 mm. long. Endemic to Peru.
Peru : San Martin : prope Fuan-Guerra ad Tarapoto, “‘ Strauch 1-4 m., Bl. weiss —
mit violetter Zeichnung ’’, Oct. 1902, E. Ule 6493 (K, isotype).
O. grandiflora is a geographically isolated species differing from other members of the genus in its acute leaves, its linear more or less parallel-sided calyx-segments, which equal the corolla-tube in length, and its smaller corollas, but it approaches O. jamaicensis in its comparatively large and broad leaves and many-flowered fascicles. There seems no adequate basis for the generic separation of this Peruvian species from the Jamaican species which Lindau associated with it under the name Psilanthele, even though this raises the problem of the presence of ancient South American elements in the Jamaican flora (see p. 285).
When transferred to the genus Oflonia, the epithet grandiflora is misleading as the species has smaller flowers than any other. Lindau, however, contrasted it not with members of Oplonia (Anthacanthus) but with Psilanthele eggersit, which has even smaller flowers.
A photograph of the holotype (formerly in Berlin-Dahlem) is available in Field Museum of Natural History, Chicago, photograph 8738.
2. Oplonia jamaicensis (Lindau) Stearn, comb. nov. (PI. 39). Psilanthele jamaicensis Lindau in Urban, Symb. Antill. 5 : 501 (1908).
Sparse straggling shrub to 4 m. high ; branches rather stout, light brown and minutely pubescent with very short upcurved hairs when young, later greyish and glabrous, somewhat broadened and flattened below the nodes ; spines usually absent. Leaves short-stalked, the blade narrowly obovate, oblanceolate or elliptic, the apex rounded or obtuse and often emarginate, sometimes mucronate, the base cuneate or attenuate, 3-10 cm. long, 1-8-4:5 cm. broad, glabrous above, sometimes sparsely
pubescent along the nerves below, the veining usually prominent below ; petiole
c.5 mm. long. Flowers 4-15 in sessile axillary fascicles ; pedicels 5-12 mm. long, minutely pubescent or glabrous. Calyx c.5 mm. long, the segments very narrowly triangular or subulate. Corolla purplish or lavender blue with purple blotches at the base of the lobes, pubescent, the tube c.10 mm. long, the segments 7-9 mm. long. Stamens almost included or long-exserted, the filaments c.2 mm. or 6 mm. long, the anthers 1-5 mm. long. Capsule c.1-5-2 cm. long.
Endemic to inland woods of Jamaica at about 350-800 m.
JAMAICA : St. James : White Rock Hill, 1 mile S. of Sweetwater, 640 m., 10 Mar. 1956, Stearn 454 (BM ; S), Proctor 11745 (IJ). Trelawny : Tyre, near Troy, 600 m., 24 Apr. 1906, Harris 9362 (BM ; K ; NY) ; 3 Apr. 1917, Perkins 1323 (K). 2 miles N. of Troy, 530 m., 14 Mar. 1955, Proctor 9949 (IJ) ; 4 mile N. of Troy, 480 m., 14 Mar. 1955, Proctor 9924 (IJ). Manchester : Vicinity of Mandeville, Feb. 1910, S. Brown 264 (NY). Clarendon : Peckham Woods, 760 m., May 1912, Harris rro60 (NY) ;
|
OPLONIA AND PSILANTHELE (ACANTHACEAE) 301
To Mar. 1954, Proctor 8433 (BM ; IJ) ; May 1955, Robertson 1992 (UCWI). Aenon Town to McKay, 600 m., Apr. 1962, Adams 11034 (UCWI). St. Ann : Mason River District, 3-4 miles NW. of Kellits P.O., 2 Mar. 1957, Proctor 16216 (IJ). Along road to Hollymount, Mount Diablo, 820 m., 6 Aug. 1957, Proctor 16523 (IJ) ; 680 m., Mar. 1962, Adams 10830 (UCWI). St. Catherine : Hollymount, Mount Diablo, 760 m., 11 Aug. 1908, Harris 8968 (NY, lectotype ; BM ; F) ; 15 Feb. 1905, Harris 8888 (BM; NY).
This species was first collected by Swartz but was confused, it would seem, with O. armata. It stands apart from other Jamaican species in its large leaves and numerous flowers. The corolla is described as “lavender blue’”’ by Perkins, as “ purple ”’ or “ violet ” or “‘ pale lavender ”’ by Proctor, as “ purple ’’ by Robertson.
3. Oplonia armata (Swartz) Stearn, comb, nov.
Justicia armata Swartz, Nov. Gen. & Sp. Pl. : 13 (1788) ; Fl. Ind. Occ. 1 : 28 (1797).—Vahl, Enum. Pl. 1 : 168 (1804).
Evanthemum armatum (Swartz) R. Br. ex Roem. & Schult. in L., Syst. Veg., ed. nov. I : 176 (1817).
Anthacanthus armatus (Swartz) Nees in DC., Prodr. 11 : 460 (1847).—Griseb., Fl. Brit. W. Ind. Is. : 457 (1862).—Lindau in Urban, Symb. Antill. 2 : 217 (1900).
Hesperanthemum armatum (Swartz) Kuntze, Revis. Gen. Pl. 2 : 490 (1891), excl. syn. Neesii.
Sparse straggling or erect shrub or small tree (in var. pallidior) ; branches slender, arching or spreading, pubescent when young on two opposite sides with short or very short ascending or upcurved hairs, the other two opposite sides almost or quite glabrous ; sfines often absent but when present slender, rigid, 5-10 mm. (rarely to 20 mm.) long, spreading at an angle of c.75°, glabrous or minutely pubescent. Leaves short-stalked, very variable in size and shape, the blade ovate or narrowly ovate, elliptic or narrowly obovate or oblanceolate, the apex obtuse or rounded, usually emarginate, the base cuneate or attenuate, 0-5-4 cm. long, 0-3-2 cm. broad, glabrous above, sometimes sparsely pubescent along the nerves below, the veining below scarcely evident to prominent ; petiole 2-5 mm. long. Flowers solitary or 2-4 in sessile axillary fascicles ; pedicels 5-15 mm. long, glabrous or minutely pubescent. Calyx 4-6 mm. long, the segments very narrowly triangular or subulate. Corolla deep pink, rose or red-purple (in var. armata), lavender-blue, pale purple or almost white (in var. pallidior), pubescent or glabrous, the tube 8-10 mm. long, the segments 4-8mm. long. Stamens almost included or long-exserted, the filaments c.1-5-2 mm. or 5—7 mm. long, the anthers c.1-1'-5 mm. long. Capsule c. 1-5-2 cm. long.
Endemic to Jamaica, var. pallidior at sea-level to 400 m., var. armata at 150-850 m.
3a. Oplonia armata var. armata (Text-fig. 3 C, D ; Pl. 43b).
Shrub 1-5-3 m. high. Leaf-blades narrowly ovate, narrowly elliptic or narrowly obovate, the base cuneate or attenuate. Flowers mostly solitary, sometimes 2—4 in a fascicle. Corolla red, variously described as deep rose-pink, red-purple, cerise and crimson.
Forma 1. Leaf-blades mostly narrowly ovate, I-2°5 cm. long, 0-5—1°5 cm. broad. Corolla deep pink or bright cerise.
302 A SURVEY OF THE TROPICAL GENERA
JAmAIcA : St. James : White Rock Hill, 1 mile S. of Sweetwater, 640 m., 10 Mar, 1956, Stearn 455 (BM). St. Elizabeth : Balaclava, 150 m., 25 Mar. 1917, Perkins 1413 (GH ;K). Trelawny : Boothe district, 3 miles N. of Troy, 490 m., 14 Mar. 1955, Proctor 9957 (IJ). Troy, 13-18 Sept. 1906, Britton 589 (F ; NY). Manchester : Halifax, 26 July 1952, West & Arnold 896 (BM). Clarendon : Peckham Woodland, 760-850 m., 2 Mar. 1910, Harris 10877 (GH ; K ; NY ; UCWI ; US).
Forma 2. Leaf-blades varying from narrowly ovate to narrowly obovate, mostly more or less elliptic, 0-7—2-5 cm. long, 0-4-1 cm. broad. Corolla deep rose or red- purple (‘‘ deep lilac ’’).
JAMAICA : Trelawny : End of Crown Lands Road coming from Peckham, 300-600 m., Mar. 1954, Robertson 2079 (UCWI). St. Elizabeth : Ipswich, 150-210 m., 7 Mar. 1917, Harris 12508 (BM ; GH ; K ; NY ; UCWI; US). Maggotty, 5 July 1928, Orcutt 2123 (BM). St. Ann: Moneague Hills, 1850, Alexander (K). Somerton district, 500 m., 28 Mar. 1956, Stearn 608 (BM) ; Proctor rrg2r (IJ). Manchester : Mandeville, Oct. 1886, Hart 1545 (K). Clarendon : Aenon Town to McKay, 700 m., Apr. 1962, Adams 11060 (UCWI), Adams 11062 (UCWI) ; Feb. 1961, Adams goo2 (UCWI). Peckham Woods, 300-600 m., Mar. 1956, Robertson 2485 (UCWI).
Forma 3. Leaf-blades mostly narrowly obovate, I-2:5 cm. long, 0-4-1 cm. broad. Corolla rose-pink, red-purple, bright rose or crimson.
JAMAICA : Trelawny : Island View Hill, Wilson Valley district, 14 miles N. of Warsop, 600-670 m., 13 Oct. 1960, Proctor 21407 (BM ; IJ) ; 30 Apr. 1964, Proctor 24826 (BM ; IJ). Ramgoat Cave District, 450 m., 4 July 1955, Howard & Proctor 14406 (BM ; IJ ; LS) ; 19 Jan. 1956, Howard, Proctor & Stearn 14674 (A ; BM ; IJ). Norwood to Bailey, 770 m., Apr. 1963, Adams 12474 (UCWI), 12479 (UCWI). Man- chester : Glasgow near Troy, 400 m., 18 Sept. 1906, Harris 9478 (BM ; NY ; UCWI). Troy to Oxford, Sept. 1906, Britton 678 (NY). Clarendon : Peckham Woods, 760 m., 26 May 1955, Proctor 10236 (BM ; IJ) ; May 1955, Robertson 1985 (UCWI). Sz. Thomas : Between Whitehall and Big Hill, 300-900 m., 12 Feb. 1953, Proctor 7668 (IJ ; LS; NY). Gorge of Plantain Garden River, 330 m., 15 Mar. 1956, Stearn 486 (A ; BM ; UCWI).
3b. Oplonia armata var. pallidior Stearn, stat. & nom. nov. (Text-figs. 1, 2 O, 2A, bs Ple3s)., Anthacanthus acicularis sensu Lindau in Urban, Symb. Antill. 2 : 214 (1900) pro parte
min., tantum quoad specim. Harris 6324 ; non (Swartz) Nees. Psilanthele minor Lindau in Urban, Symb. Antill. 7 : 383 (1912).
Shrub or small tree to 6 m. high. Leaf-blades mostly ovate or elliptic, sometimes broadly elliptic or narrowly obovate, the base rounded to attenuate, 0-5-4 cm. long, 0-3-2 cm. broad, the veining faint or not evident beneath. Corolla pale purple or lilac or almost white with purplish markings.
JAMAICA : Hanover : Green Island, Fish River, in coastal thicket, 15 Mar. 1908, Harris 10257 (BM ;F ; K ; NY ; US ; isotypes of Psilanthele minor). Upper S. slope —
OPLONIA AND PSILANTHELE (ACANTHACEAE) 303
of Dolphin Head, 370-490 m., 11 Apr. 1955, Proctor roor8 (BM ; GH). Coastal thicket, Pedro Point, Mar. 1908, Britton 2184 (NY). Wooded limestone sea-cliffs 4 mile NW. of mouth of the Great River, 3-15 m., 15 Sept. 1957, Proctor 16655 (BM ; IJ). Lucea to Montego Bay, Mar. 1908, Britton 2195 (NY). Westmoreland : Sea- coast, Negril, 10 Mar. 1908, Harris 10224 (NY ; UCWI ; US) ; Britton & Hollick 2073 (NY). Little Bay, 15 m., Dec. 1961, Adams ro169 (UCWI). St. James : Glen Devon Road, Montego Bay, 1924, Norman 100 (BM). Hills overlooking Ironshore estate, 60-240 m., 16 Feb. 1963, Proctor 23236 (IJ). St. Elizabeth : Coast about 2 miles E. of White House, 22 Feb. 1958, Yuncker 18012 (BM ; F ;S). Vicinity of Merriman’s Point, in dry coastal woodland, 9 m., 10 May 1956, Stearn 863 (A ; BM ; UCWI) ; Proctor 15337 (IJ). Near Fort Charles, 8 Jan. 1959, Robertson 5390 (UCWI). Be- tween Southfield and Pedro Cross, 360 m., g May 1956, Stearn 843 (BM). Lover’s Leap, Santa Cruz Mountains, Sept. 1907, Britton 1151 (IJ). Yardley Chase, lime- stone cliff, 490 m., 4 Sept. 1907, Harris 9675 (NY ; UCWI; US). 1 mile NW. of Alligator Pond, 14 Sept. 1954, Howard & Proctor 13807 (IJ). 0-6 mile W. of Lititz, 240 m., 21 Jan. 1956, Howard, Proctor & Stearn 14740 (A ; BM ; IJ) ; Sept. 1907, Harris 9675 (NY ; US). Kaiser mine area S. of Gutters, 13 Sept. 1954, Howard & Proctor 13760 (IJ ; NY). Manchester : Alligator Pond, 1 May 1896, Harris 6324 (NY ; UCWI). Near Great Bay, 3 miles SE. of Alligator Pond, 15 m., 3 Apr. 1956, Stearn 641 (BM). 1 mile ESE. of Gods Well, on N. side of Round Hill, go m., 3 Apr. 1956, Proctor 11942 (BM ; IJ). St. Ann: Dry coastal thorn forest near the sea, Reynolds bucket line, 20-21 Dec. 1953, Howard & Proctor 13577 (IJ). Fort Point, Dry Harbour (Discovery Bay), 3 m., 12 Apr. 1956, Stearn 726 (BM). 1 mile W. of Discovery Bay, 30 m., Apr. 1960, Adams 6855 (UCWI). Clarendon : E. end of Port- land Ridge, 15 m., 18 Mar. 1943, Lewis rog (IJ). Round Hill near Milk Pen, 45 m., May 1962, Adams 11143 (UCWI), rrr47 (UCWI). Milk River, 3-6 m., 5 June 1965, Proctor 26894 (BM).
O. armata as accepted here includes plants of such diversity in habit, size and shape of leaf, and colour of corolla that Lindau referred them to three species and two genera. The numerous specimens collected in Jamaica since Lindau’s publications have increased rather than diminished the difficulty of classifying them. Swartz’s original diagnosis of his Justicia armata reads : “‘ I. fruticosa, aculeata, foliis oblongis emarginatis coriaceis nitidis. Jamaica’’. The description in his Flora Indiae Occidentalis attributes to it ‘‘ Caulis fruticosus, 3-4 pedalis . . . Folia breviter petio- lata cuneato-obovata, emarginata, nervosa... Aculei gemini... spinaeformes... Flores axillares, solitarii, pedunculati, coccinei ”’ and gives its habitat as ‘‘ in rupibus montium Jamaicae septentrionalis’’. Red-flowered plants of this character occur in the interior of Jamaica, e.g. at White Rock Hill, which may possibly be the type- locality, as it stands by a route between Montego Bay and Savanna La Mar ; Swartz travelled across the island from the one to the other during his stay in Jamaica (cf. Stearn, 1965). The number of flowers may vary up to four in a fascicle. Leaves of such red-flowered plants vary in size from 0-2 cm. to 2:5 cm. long, and display aston- ishing diversity in shape : three main types, described above as forms 1-3, may be distinguished under var. armata. Except in colour of corolla, it does not seem possible to distinguish from these a very variable assemblage of plants, here put
304 A SURVEY OF THE TROPICAL GENERA
under var. pallidior, occurring mostly at or near the coast and having pale purple, — lavender-blue or almost white flowers ; one of these, in a spineless state, was described — by Lindau in 1912 as Psilanthele minor, the specific epithet of which is certainly in- appropriate for a variant of O. armata forming “a small tree up to 20 feet high in— coastal thickets ’’, as noted by Harris for the type-collection (Harris 10257) ; Harris” recorded it as having “‘ flowers pale blue’”’. This, however, is also evidently allied closely to plants inhabiting the interior of Jamaica and possessing larger leaves and | more numerous flowers (4-1 5 in a fascicle) described by Lindau in 1908 as P. jamai- censis. The latter occurs in at least one locality, White Rock, with O. armata var. armata {. 1 ; according to the limited material from this place, they here keep distinct, the difference in flower colour probably serving to prevent cross-pollination.
Swartz’s description of his Justicia armata as having solitary red flowers excludes O. jamaicensis. A specimen presented by him to Sir Joseph Banks as his J. armata unfortunately lacks corollas. It has glabrous shoots, spines about 5 mm. long, solitary flowers with pedicels to 1 cm. long and calyx about 5 mm. long, but oblan- ceolate or narrowly obovate leaves, varying from 2-5 cm. long, I cm. broad to 5-5 cm. long, 2cm. broad. No specimen exactly corresponding to this has been seen among the numerous collections now available from Jamaica. In its solitary flowers it agrees with O. armata as accepted here on the basis of Swartz’s description ; in its large leaves it comes closer, however, to O. jamaicensis. It might possibly represent a hybrid between them. Its existence suggests that O. jamaicensis could be united with O. armata, but in view of the distinctness of all the other specimens they are here kept apart. In Swartz’s herbarium at the Riksherbarium, Stockholm, there are four pieces labelled “‘ armata’’, evidently part of the same gathering, one piece with a spine to I cm. long, another with four pedicels arising from the same axil. This agrees with O. jamaicensis. The descriptions in Swartz’s Flora Indiae Occidentalis for the most part are evidently based on notes made directly from living plants in Jamaica and, as far as they go, are quite accurate. The explanation of the discrep- ancy between Swartz’s description and his now available material may be that he described his armata from one living plant and kept dried specimens from another.
4. Oplonia spinosa (Jacq.) Raf., Fl. Tellur. 4 : 65 (1838). (Text-figs. 2 A, B, C ; Pl. 43a).
Justicia spinosa Jacq., Enum. Syst. Pl. Carib. : 11 (1760) ; Select. Stirp. Amer. : 2, t.2 fig. I (1763) ; Select. Stirp. Amer. Ic. Pict. : 7, t.2 (1780).—L., Sp. Pl., ed. 2, 2 : 1663 (1763) excl. pl. Milleri—Swartz, Obs. Bot. : 12 (1791).—Willd. in L., Sp. Pl., ed. 4, 1 : 100 (1797). —Vahl, Enum. Pl. 1 : 167 (1804).
Evanthemum spinosum (Jacq.) R.Br. ex Roem. & Schult. in L., Syst. Veg., ed. nov. 1 : 176 (1817).
Evranthemum emarginatum Link in Jahrb. Gewachsk. 1 (3) : 47 (1820).—Schult., Mant. I : 154 (1822).
Anthacanthus eee (Jacq.) Nees in DC., Prodr. 11 : 460 (1847).—Lindau in Urban, Symb. Antill. 2 : 216 (1900) excl. parte.—Britton & Millsp., Bahama FI. : 403 (1920).— Urban, Symb. Antill. 8 : 655 (1921).—Britton & Wils. in Sci. Surv. Porto Rico & Virgin Is. 6 : 212 (1925).—Alain in Leén & Alain, Fl. Cuba 4 : 500 (1957).—P. S. Green in Kew Bull. 23 : 275 (1969).
OPLONIA AND PSILANTHELE (ACANTHACEAE) 305
Anthacanthus spinosus var. horridus Nees, loc. cit. (1847). Anthacanthus emarginatus (Link) Nees, tom. cit. : 461 (1847). Jasminum coeruleum Kuntze, Revis. Gen. Pl. 2 : 410 (1891). Hesperanthemum spinosum (Jacq.) Kuntze, tom. cit. : 490 (1891).
Shrub of variable habit, usually erect, sometimes dwarf but up to 3 m. (9 ft.) high : branches ascending, slender, glabrous on 2 opposite sides and pubescent on the 2 intervening sides with short spreading hairs or densely and minutely pubescent all round, sometimes with the hairs on two sides slightly longer ; spines curved or directed downwards slightly or strongly, sometimes almost straight and horizontal, 4-12 mm. long, at base c.0-5-0-7 mm. thick, glabrous or minutely pubescent. Leaves short-stalked, the blade very variable in shape and size, mostly narrowly elliptic or narrowly obovate, sometimes broadly ovate or elliptic, the apex obtuse or rounded and almost truncate, entire or emarginate, the base cuneate, c.8-20 mm. long (including the 1-1-5 mm. long petiole), 3-9 mm. broad, glabrous, the veining below faint or evident. Flowers solitary or 2-6 in sessile axillary fascicles ; pedicels 3-9 mm. long. Calyx 3-5 mm. long, the segments narrowly triangular or subulate. Corolla bluish purple or lavender, glabrous or pubescent, the tube c. 6-g mm. long, the segments 6-7 mm. long. Stamens included or exserted, the filaments 1-5-2 mm. or 6 mm. long ; anthers c.1-8 mm. long. Style gQ-10 mm. or 6 mm. long. Capsule c.I:2 cm. long.
Bahamas, from Grand Bahama, New Providence and Andros, over eastern Cuba, Hispaniola and Puerto Rico, to the Virgin Islands, mostly on coastal limestone, but ascending to goo m.
Bauamas : Grand Bahama : Barnett’s Point, 5-13 Feb. 1905, Britton & Millsbaugh 271r (F). New Providence : 13 Mar. 1888, Eggers 4487 (BM: K). Near Nassau, 24 Mar. 1903, Curtiss 133 (BM ; F ; GH ; K) ; 8 Mar. 1905, Wight 174 (GH ; K). Point Montague, 27 Aug. 1904, Britton & Brace 311 (F). Rifle Range, 3 Feb. 1905, E. G. Britton 3279 (F). Andros : Mangrove Cay, Aug.—Sept. 1906, Brace 4926 (F) ; 16-19 Jan. 1910, Small & Carter 8471 (F ; K). Fresh Creek, 10 June 1890, Northrop & Northrop 396 (F ; GH) ; 12 Sept. 1937, Matthews 131 (K). Nicols Town, 8 Apr. 1890, Northrop & Northrop 396A (K). Eleuthera : fide Britton & Millspaugh, loc. cit. (1920). San Salvador (Watling Island) : Graham Harbour to Cockburn Town, 15 Mar. 1907, Britton & Millspaugh 6192 (F). Cockburn Town and vicinity, 12-13 Mar. 1907, Britton & Millspaugh 6099 (F). Rum Cay : Sclater’s Hill, 4 Dec. 1905, Brace 3949 (F). Port Nelson, 23 Mar. 1966, Gillis 6298 (A). Exuma: Cay N. of Wide Opening, 18 Feb. 1905, Britton & Millspaugh 2783 (F). Crooked Island : Landrail Point, Jan. 1906, Brace 4560 (F). Long Cay : Dec. 1905, Brace 4108 (F.). Acklins Island : Spring Point, Dec. 1905—Jan. 1906, Brace 4281 (F.) Caicos Islands : Bellemont, North Caicos, 2 Mar. 1911, Millspaugh & Millspaugh 9188 (F). Jackson- ville, East Caicos, 26 Feb. 1911, Millspaugh & Millspaugh 9083 (F). Vicinity of Cockburn Harbour, South Caicos, 17 June 1954, Proctor 8830 (IJ). Great Inagua Island : Between Matthew Town and Lighthouse, 14 Oct. 1904, Nash & Taylor 1053 (F ; K). Salt Pond Hill, 12 Oct. 1904, Nash & Taylor 993 (F ; K).
CuBA : Oriente : Mayari, Charrascos cerca de la crestas del Cristal, 2~7 Apr. 1956, Alain, Acuna & Lopez Figueiras 5701 (LS), 5703 (LS), 5708 (LS), 58r0 (LS). Cerro
306 A SURVEY OF THE TROPICAL GENERA
de Miraflores, Cananova, July 1942, Léon & Mentero 21152 (LS). Moa, lieux rocheux, 16-23 Apr. 1943, Marie-Victorin, Clemente & Alain 21485 (LS ; MT), 21778 (LS). Moa, Chemins de Cayo Chiquito, 18 May 1944, Clemente 3646 (LS ; MT). Moa, bois en arriére de la Mine Franklin, 20 May 1944, Clemente 3665 (LS ; LT). Moa, Playa la Vaca, Apr. 1943, Marie-Victorin, Clemente & Alain 21503 (LS). Rio Yagrumaje, 16-23 Apr. 1943, Marte-Victorin, Clemente & Alain 21621 (LS), 21625 (MT). Moa, chemin de la mine Delta, a l’est de la vallée du Rio Cayoguan, Marie- Victorin & Clemente 21778 (LS ; MT). Moa, Orillas del Rio Cayoguan, 13 July 1949, Clemente & Alain 880 (LS). Moa, curso del Rio Cayoguan, 13 July 1949, Alain & Clemente 856 (LS). Sierra de Nipe, 24 July 1914, Ekman 2179 (S). Sierra de Nipe, headwaters of Rio Piloto, 750 m., 15 May 1915, Ekman 5701 (S) ; 375 m., 3 June 1924, Ekman 9170 (S). Headwaters of Brazo Dolores, 925 m., 15 Oct. 1914, Ekman 3117 (S). Charrasco, Pefia Prieta, Magdalena, 30 July 1953, Alain 3491 (LS).
Haiti : Without precise locality, Poiteaw (BM ; P) ; Schomburgk 107 (BM). Département du Nord Ouest : Vicinity of Jean Rabel, 1-13 Mar. 1929, Leonard & Leonard 13674 (A ; GH). Vicinity of Port 4 ’Ecu, Mar. 1929, Leonard & Leonard 13876 (K). Département de l Artibonite : Plaisance, Gonaives, 400 m., May 1900, Buch 375 (IJ). Département del Ouest : Duchené, Port au Prince, 800 m., Mar. 1920, Buch 1851 (IJ). Massif de la Selle, Port au Prince, 300 m., 21 Apr. 1926, Ekman 5923 (K). Massif de la Hotte, gr. Morne Rochelois, Miragoane, 25 Mar. 1927, Ekman 7896 (S).
DOMINICAN REPUBLIC : Without precise locality, 1852, Schomburgk (BM). Puerto Playa, sea-level, 1 May 1959, Jimenez 3931 (BM). Opposite San Gabriel, S. side of Samana Bay, sea-level, 10 April 1921, Abbott 1293 (GH). Barahona, Enriquillo (Petit Trou), sea-level to 100 m., 15 Feb. 1922, Abbott 1756 (GH).
PuERTO Rico : Aquadilla: Cape Borinquen, 2 Mar. 1929, Britton & Britton 9283 (F ; GH). Quebradillas, 2 Mar. 1914, Britton, Britton & Cowell 1958 (F). Bayamon: Hills near Bayamon, 20 m., 23 Feb. 1900, Heller 4688 (A; F; GH). Humacao : Punto Cabalero, 29 Sept. 1886, Sintenis 5202 (BM; GH). Arecibo : Prope Utuado ad Los Angelos, 17 Jan. 1887, Sintenis 59173 (K). Vieques Island : Cayo Puerto Real, 7, Feb. 1914, Shafer 2753 (GH).
St. THomaAs : Without precise locality, Herb. Willdenow 276 (B, type of Eranthe- mum emarginatum ; BM, photo) ; Feb. 1874, Kuntze (K, isotype of Jasminum coeruleum) ; Feb. 1887, Eggers 630 (A ; P). Cowells Hill, Jan. 1881, Eggers 203 (K).
ST. JOHN : Without precise locality, 1 March 1957, Robertson 26 (A).
VIRGIN GorDA : North Sound, 29 Oct. 1918, Fishlock 6 (A ; GH). Eastern slope of Virgin Peak, 150-420 m., 15 Apr. 1956, A. C. Smith 10562 (A ; K ; IJ).
St. Crorx : Without precise locality, L. C. Richard (P) ; May 1896, Ricksecker 335 (F ; GH). E. slope of Mount Eagle, 230 m., 21 Dec. 1959, Proctor 20448 (A ; BM ; IJ). Near Kingshill, 20 Jan. 1961, Howard 15658 (A).
Jacquin’s epithet spinosa is the earliest one given to a member of the group ; his original diagnosis under Justicia spinosa being but three words long, ‘‘ Justicia monanthera, spinosa’’, this epithet has excusably been applied to a diversity of
OPLONIA AND PSILANTHELE (ACANTHACEAE) 307
plants. The term “ monanthera”’ as used here means that the two loculi (thecae) of the anther are at the same level and opposite one another, thus obviously being part of one anther, in contrast to the term “ dianthera ”’ referring to stamens in which the two loculi are at different heights, the filament thus appearing to possess two anthers. Jacquin’s diagnosis served to distinguish his species from all the species until then referred to Justicia ; it fits, however, almost all the members of the genus Oplonia and its application would be obscure indeed but for the detailed description and the illustration published three years later in his Selectarum Stirpium American- arum Historia. This establishes the name as based on a shrub up to 1:5 m. high with more or less elliptic or narrowly ovate leaves about 1-5 cm. long and cuneate at base, with spreading spines shorter than the leaves, one to four flowers at an axil, a short calyx only half the length of the corolla-tube, and a purple corolla, its habitat being
| “in fruticetis & arbustis maritimis circa Portum Principis in Domingo’’. There | can be no doubt that a plant collected at Duchené, Port au Prince, Haiti, by W. Buch
(Buch 1851 in IJ) represents Jacquin’s Justicia spinosa. While some spines on this
| specimen spread horizontally, most of them curve markedly downwards and they
vary from 8 to15 mm.inlength. The leaves range from I to 2 cm. in length, 2:5 to 7mm. in width. There does not seem any justification for separating specifically from this gathering other Haitian specimens with downwards curved spines, e.g. Buch 375 from Gonaives, which has, however, small ovate leaves rounded at base, and Leonard & Leonard 13674 from Jean Rabel, which has lanceolate, narrowly elliptic and narrowly oblanceolate leaves. Buch’s Duchené material links up, however, with comparatively large-leaved specimens from the Moa district of Cuba (e.g. Marie- Victorin, Clemente & Alain 21778) on which the leaves vary from elliptic to ovate and narrowly obovate and from 7 mm. to 35 mm. long. The range of variation within a a series of specimens collected in the Moa district does not make it feasible to separate specifically the plant from St. Thomas on which the names Evanthemum emarginatum Link and Jasminum coeruleum Kuntze are based. Leonard and Leonard’s small- leaved material from Jean Rabel in turn links up with the small-leaved plants characteristic of the Bahamas. There is indeed surprisingly little difference between some specimens from New Providence, Bahamas (e.g. Curtiss 133) and some from St. John, Virgin Islands (e.g. Robertson 26).
5. Oplonia microphylla (Lam.) Stearn, comb. nov. (Text-figs. 2 J—M, 4; Pl. 44).
Justicia microphylla Lam. in Tab. Encycl. Méth., Bot. 1 : 37 (1791).—Vahl, Eclog. Amer. 2:1 (1798) ; Enum. Pl. r : 168 (1804).
Evanthemum microphyllum (Lam.) R.Br. ex Roem. & Schult. in L., Syst. Veg., ed. nov. 1: 176 (1817).
Evanthemum lycioides Link in Jahrb. Gewachsk. 1 (3) : 47 (1820).
Anthacanthus microphyllus (Lam.) Nees in DC., Prodr. 11 : 461 (1847).—Griseb., Fl. Brit. W. Ind. Is. : 457 (1862).—Duss in Ann. Inst. Colon. Marseille 3 : 427 (1897).
Anthacanthus lycioides (Link) Nees, loc. cit. (1847).
Anthacanthus cuneatus Nees, loc. cit. (1847).
Hesperanthemum microphyllum (Lam.) Kuntze, Revis. Gen. Pl. 2 : 490 (1891).
Anthacanthus acicularis sensu Lindau in Urban, Symb. Antill. 2 : 214 (1900) pro parte ; non (Swartz) Nees.
308 A SURVEY OF THE TROPICAL GENERA
Erect much-branched spiny shrub to 1-5 m. high ; branches spreading almost horizontally or ascending, slender, brownish or grey when young, glabrous on 2 opposite sides and pubescent on the 2 alternating sides, with short or moderately long spreading or ascending whitish hairs 0-I-0-3 mm. long, or pubescent all round, the internodes 3-15 mm. long ; sfines ascending or almost horizontally spreading, ~ straight 3-12 mm. long, glabrous or pubescent. Leaves short-stalked, the blade elliptic or narrowly elliptic or narrowly ovate to oblanceolate or narrowly obovate, the apex rounded or almost truncate, slightly emarginate and mucronate, the base cuneate, c.3-12 mm. long (including the 0-5-1 mm. long petiole), 1-5-6 mm. broad, — glabrous above, glabrous or occasionally pubescent below along the midrib, the veining faint not evident. Flowers solitary or paired ; pedicels 2-7 mm. long, glabrous. Calyx 2-3-5 mm. long, the segments narrowly lanceolate. Corolla pale purple, lilac or almost white with purple dots on the lower lip, minutely pubescent, the tube c.6~7 mm. long, the segments 5-6 mm. long. Stamens almost included or long-exserted, the filaments c.I:5 mm. or 4-5 mm. long, the anthers 1-5 mm. long. Capsule c.I-2 cm. long ; seeds c.2 mm. long. }
South-eastern Jamaica, Hispaniola and Puerto Rico, Lesser Antilles south to the Grenadines, at sea-level to about 600 m., mostly under 200 m. ;
Jamaica : Without precise locality, 1730, Houstoun (BM) ; W. Wright (BM) ; Macfadyen (K, type of Anthacanthus cuneatus). St. Catherine : Port Henderson Hills, Nov. 1954, Asprey 2059 (UCWI). Lazaretto, above Port Henderson, 30-75 m., 27 Mar. 1955, Proctor 997t (BM ; IJ). Kingston & St. Andrew : E. of Militia Rifle Range, 45 m., 8 Jan. 1904, Harris 8609 (NY). Long Mountain, S. side, 90 m., 3 July 1907, Harris 9606 (BM ; K ; NY ; UCWI) ; 90-220 m., 19 June 1959, Webster, Ellis & Miller 8364 (BM). W. slope of Long Mountain, along road to Wareika, 75-370 m., 7 May 1954, Farr 5133 (IJ) ; 140 m., 3 Jan. 1956, Stearn 66 (A ; BM; UCWI) ; Proctor 11436 (IJ) ; 150 m., 7 May 1956, Stearn 830 (A ; BM). Gorge of the Cane River, 75-120 m., 22 May 1955, Proctor 10195 (IJ) ; 29 Oct. 1957, Yuncker 17215 (F). St. Thomas : Four Mile Wood, between Bull Bay and Grant’s Pen, 150 m., 28 Nov. 1958, Proctor 18268 (A ; BM ; IJ). Grant’s Pen, 6 m., Dec. 1959, Adams 5665 (UCWI). Yallahs, 30 m., 16 June 1952, Avaque-Molina & Barkley 22] 448 (BM ; IJ). Vicinity of Gypsum mines, 1 mile N. of Eleven Mile, 150 m., 8 Nov. 1963, Proctor 24186 (BM).
Harri : Département de lV Artibonite : André, Gonaives, 200 m., Apr. 1899, Buch 144 (IJ). Département de l’Ouest : Plaine Cul-de-Sac, near Gauthier, 24 Feb. 1925, Ekman 3321 (GH; F;K;S). Département du Sud : Prope Port a Piment, Aug, 1917, Ekman 698 (A;S). Inter Cahauane et Aux Anglais, July 1917, Ekman 378 (S).
DoMINICAN REPUBLIC : Without precise locality, 1852, Schomburgk 37 (BM). Barahona : Barahona, Las Salinas, 40 m., May 1911, Fuertes 1105 (A ;P). Santiago? distr. of San José de las Matas, 600 m., 12 Oct. 1931, Valeur 724 (A ; BM ; F ; K ; 5S). |
Pico Diego de Ocampo, 1220 m., 3 Mar. 1957, Jimenez 3455 (BM).
PuERTO Rico : Ponce : Ponce to Playa Las Cucharas, 13 Mar. 1913, Britton, Britton & Marble 1948 (A ; F). Ponce to Penuelas, 22 Mar. 1906, Britton & Cowell 1271 (F). Culébra : Mar. 1906, Britton & Wheeler 23 (F). Vieques : Jan. 1876, Eggers (GH).
} |
OPLONIA AND PSILANTHELE (ACANTHACEAE) 309
St. Croix : Without precise locality, L. C. Richard (P, isotype of Justicia micro- phyla). TorTOLA : Coastal hill, 1918, Fishlock 105 (K).
ST. BARTHELEMY : Morne de Grand Fond, 250 m., 11 Oct. 1951, Le Gallo 311 (A ; LS; NY). Grand Platon, 150-200 m., 30 Oct. 1952, Le Gallo 1057 (LS ; NY) ; 12 Nov. 1953, Le Gallo 2180 (A).
ANTIGUA : Without precise locality, Wullschlagel 452 (GOET) ; Nicholson (K). Santa Maria Hill, 25 Apr. 1937, Box 757 (A ; BM). Little Cove near Half Moon Bay, July 1960, D. R. Harris 34 (BM).
} GUADELOUPE : La Désirade : Duchatssang 461 (GOET) ; 10-150 m., Nov. 1935, Stehlé 424 (IJ) ; 100 m., Stehlé & Stehlé 7ogr (A) ; 8 June 1960, Proctor 21212 (A).
GRENADINES : Petit St. Vincent : East end, Mar.-Apr. 1950, Howard 10916 (A ; BM). Little Martinique : Mar. 1950, Howard 10933 (A ; GH).
O. microphylla was published by Lamarck in 1791 under the name Justicia micro- phylla with an extremely brief diagnosis and a vague indication of its provenance as follows : ‘‘ J. fruticosa, foliis obovatis coriaceis, spinis axillaribus brevissimis, | pedunculis unifloris lateralibus. Ex insulis Caribaeis. Richard. an justicia armata Swartz.”’ The type-specimen in Lamarck’s herbarium, now at the Muséum National d'Histoire Naturelle, Paris, gives no more precise information about its place of collection but fortunately it matches so exactly a localized specimen, under another hame, in L. C. Richard’s own herbarium, likewise at Paris, that they presumably formed part of the same gathering made “‘ in fruticetis sepibus Stae Crucis ”’, i.e. on the island of St. Croix, Virgin Islands, which Richard visited between February 1786 and November 1787 (cf. Urban, Symb. Antill. 3 : 111 (1902) ). Richard’s material has light brown branches which are almost glabrous on two opposite sides but pubescent on the other two sides with moderately long whitish hairs to 0-3 mm. long ; the short stout slightly ascending spines are up to 3 mm. long ; its numerous, narrowly obovate leaves are 4-10 mm. long, 3-6 mm. broad, and sometimes have a few hairs along the midrib below. The small leaves and open habit, with branches spreading laterally from the main stem at an angle of about 60°-80°, give it a distinctive appearance which it shares with other specimens from the Lesser Antilles ; these diverge from it principally in having somewhat longer (5-12 mm. long) spines. On the dry coastal limestone of south-eastern Jamaica grow similar small-leaved plants with, however, more erect branches which ascend from the main stem at an angle of about 50° or less. There is much variation in degree of hairiness within this Jamaican population. Anthacanthus cuneatus was founded on a gathering by James Macfadyen with markedly hairy shoots, spines and leaves ; among modern Jamaican collections this is represented by Avaque-Molina & Barkley 22]448 from Yallahs, St. Thomas. It does not seem possible, however, to make any satisfactory distinction between this and other less hairy Jamaican specimens of the same area : distinctions between specimens from Jamaica and those from Hispaniola and Puerto Rico are likewise not clear-cut. Hence the epithet microphylla is here used to cover them all.
310 A SURVEY OF THE TROPICAL GENERA
6. Oplonia tetrasticha (C. Wright ex Griseb.) Stearn, comb. nov.
Anthacanthus tetrastichus C. Wright ex Griseb., Cat. Pl. Cub. : 198 (1866).—Alain in Leén & Alain, Fl. Cuba 4 : 500, fig. 219 (1957).
Hesperanthemum tetrastichum (C. Wright ex Griseb.) Kuntze, Revis. Gen. Pl. 2 : 490 (1891).
Anthacanthus acicularis sensu Lindau in Urban, Symb. Antill. 2 : 214 (1900) pro parte min. ; non (Swartz) Nees.
Much branched spiny shrub, up to about 2 m. high ; branches stout, yellowish when
young and pubescent all round or only on two opposite sides with short spreading © hairs, the internodes 5-15 mm. long ; spines horizontally spreading or very slightly —
ascending straight, rigid, 1-3-5 cm. long, to 1 mm. thick, simple (var. tetrasticha) or branched (var. polyece), glabrous or minutely pubescent. Leaves short-stalked, the
blade narrowly obovate to narrowly elliptic, the apex rounded, slightly mucronate, — the base cuneate, 6-10 mm. long (including the 1-3 mm. long petiole), 2-3 mm. broad ~ much shorter than the spines, glabrous, the veining not evident. Flowers solitary — in the axils ; pedicels 4-6 mm. long, glabrous. Calyx c.3-5 mm. long, the segments ~
subulate. Corolla lilac or “‘bright violet with white purple-dotted patch in throat ”’ (fide Proctor), glabrous or pubescent, the tube 4:5-10 mm. long, the segments c.3-5 mm. long. Stamens included or exserted, the filaments c.1 or 4 mm. long, the anthers c.1-5 mm. long.
Endemic to Cuba.
6a. Oplonia tetrasticha var. tetrasticha (Text-figs. 2 D, E).
Spines all simple, not branched.
CuBA : Without precise locality : 1860-64, C. Wright 3067 (BM ; K ;S, isotypes). Pinar del Rio : Cuabales al Este de la Loma de Cajalbana, La Palma, 10 June 1950,
Alain & Acuna 1395 (LS) ; 3 Apr. 1954, Alain 3909 (LS). Norte del Pan de Guajaibén, —
5 Apr. 1952, Alain 2423 (LS). Habana : Guanabacoa, Lomas de las Jatas, 3 June 1923, Ekman 16530 (S). Cojimar, 24 May 1914, Ekman rior (S). Columbia,
Habana, 22 June 1912, Ledn 3326 (LS; P). Valle del Almendares, Dec. 1908, Leon
465 (LS). Loma S. dela Jata, Guanabacoa, 4 Jan. 1912, Ledn 2865 (LS). Loma de Machado, Jibacoa, 2 Jan. 1929, Leén & Roig 13756 (LS). Matanzas : Camarioca, 1849, Rugel 249 (K). Cerca de la Boca de Canasi, 17 Oct. 1928, Leén 13696 (LS). Las Villas (Santa Clara) : Sabana de Motembo, 2 Jan. 1919, Leén & Fortem 8579 (LS). Loma de Belén, cerca de Santa Clara, 28 June 1931, Ledn 14947 (LS). Cienfuegos Bay, dry east-facing slopes of Loma San Juan, 11 July 1947, Wood & Atchison 7416 (IJ). Gavilan, Cienfuegos, 13 Mar. 1928, Jack 5815 (A; LS). Alto de Farallén, Ensenada de Guajimico, S. de Cienfuegos, 21 Mar. 1953, Alain, Clemente & Wal- singham 2839 (LS). Guajimico Bay, on coast S.E. of Cienfuegos, 3-15 m., 23 Mar. 1957, Proctor 16430 (IJ). Oriente : El Cobre,6Oct. 1916, Ekman 7832 (S). Santiago, in limestone hills W. of Playa Siboney, 8 Nov. 1917, Ekman 8723 (S). Santiago, pr. Aquadores, 1922, Ekman 15631 (S). Sierra de Nipe, between Rio Bayate and Rio Piedra, 8 Sept. 1922, Ekman 15042 (BM ;S).
OPLONIA AND PSILANTHELE (ACANTHACEAE) 311
6b. Oplonia tetrasticha var. polyece Stearn, var. nov. (Text-figs, 2 F, G.)
Spinae primariae rectae lignosae I-3:5 cm. longae plerumque ramosae spinas secundarias plerumque 2 interdum 4 geminatas I-12 mm. longas vel tantum gemmas minutas geminatas ferentes.
Primary spines straight, I-3-5 cm. long, usually branched and bearing 2-4 paired spines I-12 mm. long or in their place minute paired buds.
CuBA : Oriente : Sierra Maestra, on the edge of Arroyo Corojo, a tributary of Yara, near Nagua, c.150 m., 5 Aug. 1922, Ekman 14735 (BM). Cuabales de serpentina, Holguin, May 1939, Leén & Carabia 18981 (LS). Pinal Mayari, Sierra de Nipe, 7 Apr. 1941, Leén, Marie-Victorin, Clemente & Alain 19920 (LS). Renté, Bahia de San- tiago, I May 1943, Clemente 2680 (LS). Club de Punta Gorda, Bahia de Santiago de Cuba, 1 Mar. 1953, Lopez Figuetras gtr (LS). Manigua costera de Santiago de Cuba, Mar. 1954, Lopez Figuetras 1131 (LS). Antigua linea de Firmeza, Santiago de Cuba, May 1946, Clemente 5014 (LS). Manigua de Ciudamar, June 1946, Clemente 5039 (BM ; LS). Camino viejo del Morro, Oct. 1947, Clemente 5657 (BM, holotypus ; LS) ; July 1943, Clemente 2906 (LS). Manigua de Aguadores, Santiago de Cuba, May 1946, Clemente 5021 (LS). Punta de la Gaviota junto a la Playa de Siboney, Santiago de Cuba, 24 Feb. 1952, Lopez Figueras 359 (LS). Farallon de la Perla, 14 Feb. rg11, _ Shafer 8772 (GH ; K). La Perla, prope villam Monte Verde, 1859, C. Wright 369 (GH ;K). Las Ninfas, Guantanamo, Feb. 1919, Hioram 2354 (LS). Macambo, Via Azul, 28 Dec. 1954, Alain & Lopez Figueiras 4210 (LS). Charrascos de Jauco, July 1924, Leén 1191 (LS). Region de Baracoa, entre Jauco y Cajobabo, July—Aug. 1924, Leén 11987 (LS), 12079 (LS), 12031 (LS).
The long and rigid sharply pointed spines projecting from the branches in four ranks (whence the epithet ¢etrasticha) have earned this species the Cuban vernacular name “‘ no-me-toques ’’. They are longer than in any other species of Oplonia. The type collected by Charles Wright somewhere in western Cuba has simple stiletto-like spines, as have other specimens collected outside of the Oriente province. Within that province, along the south coast, there is a marked tendency for these major spines to bear a pair or two pairs of subsidiary spines which are usually short but may be up to 12 mm. long. Numerous specimens with such branched spines have been collected around Santiago de Cuba. Many of them have the shoots hairy on two opposite sides only, the two sides alternating with these being glabrous, but some (e.g. Lopez Figueivas 359 from Siboney, near Santiago de Cuba) are hairy all round. The leaves of main shoots vary from 5 to 12 mm. in length ; those on the spines are much smaller. The plant would appear to initiate a lateral branch, then halt its development after producing a pair or less often two pairs of short spines, then stop its development entirely by making it terminate as a sharp spine. The few corollas examined of this variant have been glabrous, whereas the few corollas examined of typical O. tetrasticha have been minutely pubescent. A specimen of the Baracoa region, Oriente, collected by Leén (Leén 12031), appears at first to have long simple spines as in typical O. tetrasticha, but these bear nevertheless a pair of minute buds corresponding to the secondary spines of other specimens collected in this area by Leén. The Oriente population with branched spines is accordingly here distin-
312 A SURVEY OF THE TROPICAL GENERA
guished as var. polyécé, the epithet from zodv- (‘many ”’) and -nKn (‘‘ pointed ”’) being formed by analogy with edjKn ‘(‘‘ well-pointed’’) and Aerrnxn (“ fine-pointed”’). Usually the spines are more than 1-5 cm. long. A specimen from Habana province, ; collected by Ekman (Ekman rror) at Cojmar, which it does not seem possible to refer to any species but O. tetrasticha, nevertheless has straight slightly ascending spines : with the smallest down to 3 mm., the longest to 2 cm. long. Another Ekman | k ;
gathering (Ekman 15042) from Oriente province has more slender spines than is typical and approaches O. microphylla.
7. Oplonia acicularis (Swartz) Stearn, comb. nov. (Text-fig. 2 H, I).
Justicia acicularis Swartz, Nov. Gen. & Sp. Pl. : 13 (1788) ; Fl. Ind. Occ. 1 : 29 (1797).— ~ Vahl, Enum. Pl. 1 : 169 (1804). { Evanthemum aciculare (Swartz) R. Br. ex Roem. & Schult. in L., Syst. Veg., ed. nov. I : 177 ©
(1817). Anthacanthus acicularis (Swartz) Nees in DC., Prodr. 11 : 460 (1847) excl. parte. Anthacanthus jamaicensis Griseb., Fl. Brit. W. Ind. Is. : 457 (1862), nom. superfl. Hesperanthemum aciculare (Swartz) Kuntze, Revis. Gen. Pl. 2 : 490 (1891). ; Anthacanthus spinosus sensu Lindau in Urban, Symb. Antill. 2 : 216 (1900) pro parte min. ; non (Jacq.) Nees.
Low spreading much branched spiny shrub, mostly under 30 cm. high ; branches slender, brownish when young and pubescent with minute upward-pointing hairs on 2 or all sides ; spines very slender, 8-14 mm. long, c. 0-3 mm. thick. Leaves short- stalked, the blade ovate to narrowly lanceolate, the apex obtuse, mucronate, the base cuneate or rounded, 7—25 mm. long (including the c.1 mm. long petiole), 4-10 mm. broad, glabrous, the veining not evident. Flowers solitary in the axils ; pedicels c.3-4 mm. long, minutely pubescent. Calyx 2-2-4 mm. long, the segments very narrowly lanceolate or subulate, ciliate. Corolla white, glabrous, the tube cylindrical c.4-6:5 mm. long, the limb spreading with segments oblong, c.4-6 mm. long, 2:5- 3 mm. broad, obtuse. Stamens included, inserted on the upper third of the corolla- tube, the filaments c.o-8 mm. long, the anthers I-¢ mm. long. Capsule c.I-5 cm. long ; seeds c.2-5 mm. long, rugose.
Endemic to Jamaica.
Jamalca : Without precise locality : 1862, March (K) ; Swartz (S, type collection) ;
Wilson 453 (GOET ; K). Westmoreland : Between Grange Hill and Glasgow, Mar.— Apr. 1908, E. G. Britton 2894 (NY). Hills near Bulstrode, 12 Mar. 1908, E. G. Britton 2877 (K ; NY). Portland : Gorge of the Swift River at Eden, $ mile N. of Paradise, 30 m., I9 Mar. 1956, Stearn 550 (BM), Proctor 11866 (IJ). Navy Island, June 1897, Fredholm 3073 (NY ; US). Friendship Valley district near Ecclesdown, 330-360 m., 23 May 1967, Proctor 27920 (IJ). Priestman’s River, Nov. 1886, Hart 1545 (K) ; 3 m., 13 Aug. 1957, Proctor 16574 (IJ) ; 18 Mar. 1958, Yuncker r8g2zr (IJ). Law- rence Bottom, 370 m., Mar. 1961, Adams 9258 (UCWI). St. Thomas : Big Level district, S.E. end of John Crow Mountains, 450-600 m., 16 Mar. 1956, Stearn 498 (BM), Proctor 1818 (IJ).
Although O. acicularis has been recorded (as Anthacanthus acicularis) from many West Indian islands, through confusion with other species, it is, in fact, a Jamaican —
OPLONIA AND PSILANTHELE (ACANTHACEAE) 313
endemic characterized by low growth, slender needle-like spines, mostly ovate leaves and small white flowers which make it easily distinguishable in a living state from other Jamaican species. It often grows in somewhat shaded places. Swartz’s original diagnosis of his Justicia acicularis reads : “I. fruticosa, diffusa, spinosa, spinis setaceis, floribus pedunculatis axillaribus solitariis. Jamaica’”’. Later he supplemented this in his Flora Indiae Occidentalis with a detailed description, giving particulars such as “ Fruticulus pedalis . . . Folia petiolata, opposita ovata minora, apice acumine brevissimo . . . Spinae oppositae, setaceae . . . Flores . . . parvi, albidi’”’ etc., which together apply only to the Jamaican species described above. It has a remarkably disjunct distribution within the island, most specimens having been collected on the extreme east and a few in the west but none in the middle.
8. Oplonia purpurascens (Griseb.) Stearn, comb. nov. (Text-fig. 2 P, Q.)
Anthacanthus purpurascens Griseb., Cat. Pl. Cub. : 197 (1866).—Alain in Leén & Alain, FI. Cuba 4 : 500 (1957).
Hesperanthemum purpurascens (Griseb.) Kuntze, Revis. Gen. Pl. 2 : 490 (1891).
Anthacanthus acicularis sensu Lindau in Urban, Symb. Antill. 2 : 214 (1900) pro parte min. ; non (Swartz) Nees.
Shrub ; branches ascending, slender, purplish, glabrous on 2 opposite sides, minutely pubescent on the 2 other sides when young ; spines ascending at an angle of c.48-58°, slender, straight, 4-8 mm. long, at base 0-5 mm. thick. Leaves short- stalked, the blade obovate or oblanceolate, the apex obtuse, entire or slightly emarginate, the base narrowly cuneate, 6-15 mm. long, 1-5-6 mm. broad, glabrous, the veining scarcely evident ; petiole 1-3 mm. long. Flowers solitary or paired in the axils ; pedicels c.I-1-5mm.long. Calyx c.3-4 mm. long, the segments very narrowly lanceolate or subulate. Corolla “‘ pale purple’”’, minutely pubescent, the tube c.8—9 mm. long, the segments c.6mm.long. Stamens included.
Endemic to Cuba.
CuBA : Without precise locality, 1860-64, C. Wright 3066 (BM ; K ; S, isotypes). Pinar del Rio : Sierra de Vifiales, 8 June 1923, Ekman 16582 (BM). Mogote de la _ Bandera, Vifiales, g Mar. 1924, Ekman 18658 (LS). Ensenada de los Muertos, _ Mogote de la Bandera, Vifiales, 29 Mar. 1953, Alain 2860 (LS). Ensenada de la Bandera, Mogote de la Bandera, Vifiales, Mar. 1953, Alain 2892 (LS).
O. purpurascens is a species characteristic of the mogotes (“‘ haystack hills” or “ Kegel-karst ’’) of north-western Cuba. The very short pedicels distinguish it from allied species. Possibly here belongs a specimen collected in the district of Cienfuegos, Santa Clara by R. Combs (Combs 567 ; K).
9. Oplonia nannophylla (Urban) Stearn, comb. nov. (Text-fig. 2 N, Pl. 45). Anthacanthus nannophyllus Urban in Fedde, Repert. Sp. Nov. 20 : 350 (1924).
Low spreading shrublet branched from the base, 4-8 cm. high ; branches spreading or arching, slender, minutely pubescent when young, the internodes 1-3 mm. long ; spines almost horizontally spreading or ascending, straight or very slightly curved,
314 A SURVEY OF THE TROPICAL GENERA
slender, 2:5-I1I mm. long, at base to 0-5 mm. thick. Leaves short-stalked, almost — sessile, much shorter than the spines, the blade obovate or narrowly elliptic, the — apex obtuse, the base broadly cuneate, 2-5 mm. long, I-3 mm. broad, glabrous, the cystoliths numerous, the veining not evident, rigid ; petiole c.o-5 mm. long. Flowers unknown.
Endemic to Cuba.
CuBA : Pinar del Rio : Cuabales, Oeste de la Loma de Cajalbana, La Palma, 3 Apr. © 1954, Alain 3907 (LS) ; Feb. 1953, Alain 2757 (LS). Habana : prope Guanacaboa in Lomas de la Yatas, 3 June 1923, Ekman 16534 (S, type). Las Villas (Santa Clara) : Manigua, Sabana de Motembe, 28 Aug. 1922, Léon 11385 (GH ; LS).
O. nannophylia is an inconspicuous plant of extremely reduced habit and probably slow growth, with minute leaves and short internodes, which has not yet been collected in flower. It is apparently restricted to north Cuba between 80° and 84°W.
— t t 42 B C
A . 4
Fic. 17. Leaves of Oplonia from Madagasgar. A, B, O. vincoides ; C, O. acuminata ; D. O. minor var. meridionalis.
10. Oplonia acuminata Stearn, sp. nov. (Text-fig. 17 C)
Frutex 1-3 m. altus, inermis ; yamuli hornotini in sicco atri glabri [vel pubescentes], annotini graciles pallidi, internodiis plerumque elongatis ad 9 cm. longis. Folia manifeste petiolata ; lamina ovata, apice acuminata, basi rotundata in petiolum ad cm. longum abrupte angustata, c. 2-5 cm. longa, acumine 5—9 mm. longo inclusa, 1-4 cm. lata, glabra [vel supra sparse subter magis praecipue secus venas puberula], cystolithis tantum sub lente visibilibus sed numerosis, papyracea, venis arcuatis conspicuis. Flores axillares 1-3 ; pedicelli 7-9 mm. longi, glabri. Calyx c.5 mm. longus, sparse puberulus [vel glaber], segmentis subulatis. Corolla alba, bilabiata ; tubus c.4:5 mm. longus, sparse puberulus ; labium superum ascendens c. 8 mm.
OPLONIA AND PSILANTHELE (ACANTHACEAE) 315
longum, segmentis duobus supra medium coalitis quaamobrem bilobatum lobis liberis -¢.3'5 mm. longis 2 mm. latis ; labium inferum descendens trifida, segmentis anguste ovatis acutis c. 8-g mm. longis, 4 mm. latis ; segmenta extra intraque praecipue apicem versus puberula. Stamina in fauce affixa, exserta ; filamenta laevia, in typo 5°5 mm. longa ; antherae purpureae, c.2 mm. longae ; staminodia minuta. Ovarium glabrum ; stylus puberulus, c.12 mm. longus. [Capsula c.1-+7—2+5 cm. longa].
Shrub 1-3 m. high ; branches slender, glabrous or pubescent when young ; spines absent. Leaves short-stalked, the blade ovate, the apex acuminate, the base rounded and abruptly narrowed into the petiole, 2-5 cm. long (including the 5-9 mm. long elongated tip), I-4 cm. broad, almost glabrous or else pubescent particularly below along the veins ; petiole 5-10 mm. long. Flowers 1-3 in the axils ; pedicels 7-9 mm. long, glabrous. Calyx c.5 mm. long, sparsely puberulous or glabrous, the segments subulate. Corolla white, puberulous, the tube c.4-5 mm. long, the segments 8-9 mm. long, 4 mm. broad, the upper two united to above the middle. Stamens ex- serted, the filaments (on the type) c.5-5 mm. long, smooth, the anthers c.2 mm. long. Style puberulous. Capsule 1-7—2:5 cm. long.
Endemic to western Madagascar.
MADAGASCAR : Environs de Majunga ; arbuste 1-3 m., fleur blanche a 2 lévres, la supérieure entiére, l’inférieure trilobée ; Nov. 1914, Perrier de la Bathie 12908 (P, holotype). Vallée du Fiherenana, 4 15-25 kil. en amont de Tuléar ; arbuste 2 m., corolle 4 lévre inférieure blanche, lévre supérieure blanc-verdatre, anthéres pour- pres ; vers 200 m., Mar. 1934, Humbert 14357 (K).
This is separated from O. vincoides by its somewhat broader, ovate, distinctly acuminate leaves which are also thinner in texture. The two upper segments of the corolla are united for more than half their length to form an apically bilobed upper lip. The only two known localities are some 500 km. apart. Details added from Humbert’s specimen are cited in the Latin description in square brackets e.g. [vel pubescentes].
11. Oplonia vincoides (Lam.) Stearn, comb. nov. (Pl. 41, 46 ; Text-fig. 17A, B).
Justicia vincoides Lam. in Encycl. Méth. Bot. 1 : 628 (1785).—Vahl, Enum. Pl. 1 : 167 (1804). Evanthemum vincoides (Lam.) R. Br. ex Roem. & Schult. in L., Syst. Veg., ed. nov. 1 : 176
(1817). Anthacanthus vincoides (Lam.) Nees in DC., Prodr. 11 : 461 (1847). Forsythiopsis baronit Baker in Journ. Linn. Soc. Lond., Bot. 20 :219, t.27 (1883) ‘‘ baroni’’. Forsythiopsis australis Scott Elliot in Journ. Linn. Soc. Lond., Bot. 29 : 37 (1891). Forsythiopsis vincoides (Lam.) Benoist, Cat. Pl. Madag., Acanth.: 15 (1939).—Benoist in Humbert, Fl. Madag. 182 : 106, fig. 18 no. 10-13 (1967), excl. var. minor.
Much-branched shrub 1-3 m. high ; branches slender, pilose when young ; spines absent. Leaves short-stalked, the blade narrowly elliptic to narrowly obovate, the apex usually obtuse, often emarginate, the base cuneate or attenuate, 2-9 cm. long, 0-6-3 cm. broad, conspicuously veined, puberulous when young, later almost glabrous ; petiole 2-6 mm. long. Flowers solitary or 2-3 in axillary fascicles ;
316 A SURVEY OF THE TROPICAL GENERA
pedicels 1-3 cm.long. Calyx 4-5-6 mm. long, the segments very narrowly triangular, sparsely puberulous. Corolla white, blue or violet, glabrous or pubescent, the tube c.5-8 mm. long, the segments c.8-12 mm. long, 3:5—5 mm. broad, the upper two free to the lower 2-5 mm. Stamens almost included or long-exserted, the filaments c.I-5-3 mm. or 6-8 mm. long, papillose, the anthers 1-5-2 mm. long. Style puberulous. Capsule c.2-2°5 cm. long.
Endemic to Madagascar.
MADAGASCAR : Without locality, but apparently from Fort Dauphin, Commerson (P, type collection of Justicia vincoides). Fort Dauphin, Scott Elliot 2586 (BM ; K, lectotype of Forsythiopsis australis), 2701 (K) ; Clotsel 209 (BM). 10 km. W. of Fort Dauphin, 0-12 m., 2 Sept. 1968, McWhirter & Capuron 184 (K). Central Mada- gascar [Imerina], Baron 1937 (K, type of F. baronit). Ankorafantsika, Ambalo- Boeni, 10 Oct. 1950, Ramamoufisoa 2058 (P).
Lamarck described his Justicia vincotdes from herbarium material collected in Madagascar by Philibert Commerson, who spent the last three months of 1770 botanizing around the French settlement, Fort Dauphin, in the extreme southeast of the island. Here, therefore, must be its type-locality. No specimen now exists in Lamarck’s private herbarium and possibly he never possessed one but based his description of J. vincoides on the Commerson specimen still existing in the her- barium of the Museum National d’Histoire Naturelle, Paris (Pl. 46). In 1847 Nees referred it to the genus Anthacanthus after studying another Commerson specimen in Vahl’s herbarium at Copenhagen. The epithet vinco1des was suggested by a general resemblance in leaf to a periwinkle, probably Vinca minor L. The leaves of the Commerson specimen in Paris are elliptic, narrowly elliptic or narrowly obovate, cuneate at base, obtuse at the apex, with a minute apiculus, 2-4 cm. long, 7-15 mm. broad, prominently veined and glabrous. The flowers are clustered in short-peduncled axillary cymes. The corolla is pubescent inside and out, the tube about 5 mm. long, the segments about 8 mm. long, 4 mm. broad ; the two segments of the upper lip are fused for the lower 2-5 cm. The filaments of the stamens are about 1.5 mm. long. Likewise short stamens occur on a specimen collected near Fort Dauphin by Cloisel (Cloisel 209). It is almost identical with the specimens collected near Fort Dauphin by Scott Elliot on which in 1891 he based his Forsythiop- sis australis ; these, however, have filaments about 5 mm. long. This Madagascan species thus manifests the same floral dimorphism, some individuals having long stamens and short styles, others short stamens and long styles, as the West Indian species of Oplonia.
In 1883 Baker founded the genus Forsythiopsis with one species, F. baronit, on specimens collected by the English missionary Richard Baron in central Madagascar, the precise locality in the Imerina region unfortunately not recorded. Whereas in the Fort Dauphin material of O. vincoides the leaves are evidently mature, on Baron’s specimen the leaves are only partly developed, the longest only 2:5 cm. long, I-2 cm. broad, which may be associated with a local difference of climate. The corolla is pubescent, with the tube about 7 mm. long, the segments about 10-11 mm. long, 4°5 mm. broad, the two segments of the upper lip being united for about the lower
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OPLONIA AND PSILANTHELE (ACANTHACEAE) 317
2mm. The filamentsofthestamens are about 2:5 mm.long. A specimen with similar immature leaves was collected by Ramamoufisoa. Benoist in 1939 equated it with Lamarck’s Justicia vincoides and Scott Elliot’s Forsythiopsis australis. Baker compared his Forsythiopsis with Ruellia and Benoist in 1967 similarly placed it in the ‘‘ Tribu des Ruelliées ” next to Strobilanthus. On account of its 3-colporate usually prolate pollen, with a pseudocolpus (colpoid streak) each side of the colpus, and other characters, including imbricate aestivation, it belongs, however, to the subfamily Ruelloideae tribe Ruellieae subtribe Odontoneminae as defined by Breme- kamp (1965), not to his subtribe Rwuellinae or subtribe Strobilanthidinae, and it cannot be separated from Oflonia. The unlikelihood of Madagascan material having any affinity with American and the consequent neglect of comparison together account for the acceptance of Forsythiopsis as an endemic Madagascan genus.
Although the application of the epithet vincotdes to the taxon of the Fort Dauphin area with narrowly elliptic leaves up to 9 cm. long, 3 cm. broad, is thus certain, the circumscription of the species presents much difficulty. Thus Benoist in 1930 described a Forsythiopsis minor with obovate or narrowly obovate leaves only 2-7 mm. long, I-4 mm. broad, which in 1967 he treated as F.. vincoides var. minor. In1949 he described under F. vincoides two varieties, one with glabrous leaves, his var. meri- dionalis, the other with densely hairy leaves, his var. vestita, both with obtuse leaves not more than 3 cm. long. He also included in F. vincoides material with ovate distinctly acuminate leaves. His 1967 concept of F. vincoides thus embraced several distinguishable taxa, which seem to represent at least three species.
12. Oplonia puberula Stearn, sp. nov. (Text-fig. 18).
Frutex (liana caule crasso, fide Humbert et Capuron), inermis ; vamuli pallidi, hornotini puberuli, annotini decorticantes glabri, internodis ad 5 cm. longis. Folia breviter petiolata ; Jamina anguste obovata, apice rotundata, obtusa vel acuta, basi cuneata, immatura c.2-4:5 cm. longa, I-2 cm. lata, supra sparse puberula, subter praecipue secus venas multo magis puberula, venis arcuatis conspicuis utroque latere costae 5-7 ; petiolus ad 6 mm. longus, dense puberulus. Flores axil- lares plerumque 4 ; pedicelli 5-10 mm. longi, dense puberuli. Calyx 2-5-3 mm. longus, omnino dense puberulus, segmentis anguste oblongis obtusis. Corolla violacea (fide Humbert et Capuron), bilabiata ; tubus c.7-8 mm. longus, puberulus ; labium superum ascendens c. 12 mm. longum, segmentis duobus fere ad medium coalitis quamobrem bilobatum lobis liberis c. 7 mm. longis, 3 mm. latis ; labium inferum descendens trifida, segmentis ellipticis obtusis c. 12 mm. longis, 5-5 mm. latis ; segmenta extra intraque puberula. Stamina in fauce affixa, exserta ; fila- menta valde papillosa, in typo 4 mm. longa ; antherae olivaceae (fide Humbert et Capuron), c.1-5 mm. longae ; staminodia minuta. Ovarium glabrum ; stylus dense puberulus, c.12 mm. longus.
Shrub (a thick-stemmed liane according to Humbert & Capuron) ; branches puberu- lous when young ; spines absent. Leaves short-stalked, the blade narrowly obovate, the apex rounded, obtuse or acute, the base cuneate, about 2-4-5 cm. long, I-2 cm.
318 A SURVEY OF THE TROPICAL GENERA
broad in an immature state, above sparsely puberulous, below much more thickly — puberulous, particularly along the veins ; petiole to 6 mm. long. Flowers about 4 in — the axils ; pedicels 5-10 mm. long, densely puberulous. Calyx 2-5-3 mm. long, densely puberulous all over, the segments narrowly oblong. Corolla violet, puberu- — lous, the tube 7-8 mm. long, the segments c.12 mm. long, 3-5-5 mm. broad, the upper ~ two united almost to the middle. Stamens exserted, the filaments (on the type) — c.4 mm. long, the anthers c.1-5 mm. long. Style densely puberulous. Endemic to northern Madagascar.
Fic. 18. Oplonia puberula Stearn. A, leaf; B, flower in front view: C, corolla in longitudinal section; D, calyx; E, gynoecium.
MapacGascar : Vallée inférieure de l’Androranga, affluent de la Bemarivo (nord- est) aux environs d’Antongondriha, Mont Anjenabe, forét ombrophile sur gneiss ; grande liane a tige épaisse, écorce subereuse, corolle violette, anthéres olivatres, style blanc ; 750 m., 3-7 Nov. 1950, Humbert & Capuron 24103 (P, holotype ; BM).
This taxon comes very close to O. vincoides, from which it may be distinguished
aE Ay
OPLONIA AND PSILANTHELE (ACANTHACEAE) 319
by its narrowly oblong calyx-segments completely and densely covered outside with very short erect hairs ; those of O. vincoides are narrowly triangular with much sparser hairs not completely covering their surface. It is everywhere hairier than O. vin- coides and has a slightly shorter calyx and slightly larger corolla. There would also appear to be a difference in habit if Humbert and Capuron’s description of it as a large liane is correct and applies to other specimens. On these grounds it is here given separate specific rank. The type-gathering evidently represents a long-styled state ; the style itself is more densely hairy and slightly longer (i.e. c.12 mm. long as against 8-9 mm.) than in long-styled O. vincoides.
13. Oplonia minor (Benoist) Stearn, comb. nov.
Forsythiopsis minor Benoist in Bull. Soc. Bot. France 76 : 1033 (1930).
Forsythiopsis vincoides var. meridionalis Benoist in Mém. Soc. Bot. France 1949 : 94 (1949). —Benoist in Humbert, Fl. Madag. 182 : 108 (1967).
Forsythiopsis vincoides var. vestita Benoist, loc. cit. (1949).—Benoist, loc. cit. (1967).
Forsythiopsis vincoides var. minor (Benoist) Benoist in Humbert, Fl. Madag. 182 : 109 (1967).
Shrub ; branches slender, pubescent when young ; spines absent. Leaves short- stalked, the blade obovate or narrowly obovate, the apex obtuse, the base cuneate, c.2-30 mm. long, I-Io mm. broad, puberulous when young, later glabrous or puberu- lous ; petiole c.1 mm. long. Flowers solitary or paired ; pedicels 7-9 mm. long. Calyx 2-4 mm. long, the segments subulate. Corolla pale violet or lilac, pubescent, the tube c.5—6 mm. long, the segments c.8—10 mm. long, 3~4 mm. broad, the upper two united at base for the lower third of their length. Stamens almost included or long-exserted, the filaments c.2:5 mm. or 5 mm. long, papillose, the anthers c.1-5 mm. long. Style puberulous or almost glabrous. Capsule c.1-5 cm. long.
Var. minor
F. minor Benoist, loc. cit. (1930).
Leaves 2-9 mm. long, I-4 mm. broad, puberulous when young, later glabrous.
_ Calyx 2-3 mm. long, glabrous.
MADAGASCAR : Fort Dauphin, Behara, 10 July 1926, Decary 4273 (BM ; K). Environs de Tuléar, 10-200 m., Sept. 1924, Humbert & Perrier de la Bathie 2506 (K). Many other localities are cited by Benoist (1967).
Var. meridionalis (Benoist) Stearn, comb. nov. (Text-fig. 17D).
F. vincoides var. meridonalis Benoist, loc. cit. (1949).
Leaves 12-25 mm. long, 4-10 mm. broad, becoming glabrous. Calyx c.3 mm. long, almost glabrous.
MADAGASCAR : Fort Dauphin, c. 1890, Cloisel 9g (BM). Amborombe, 7 Aug. 1931, Decary 9118 (BM; K). Massif de l’Andohahelo, au-dessus de Pisopiso
(Beake), 1000-1100 m., Jan. 1934, Humbert 13676 (K, isotype of F. vincoides var. meridionalis).
320 A SURVEY OF THE TROPICAL GENERA
Var. vestita (Benoist) Stearn, comb. nov.
F.. vincoides var. vestita Benoist, loc. cit. (1949).
Leaves 8-30 mm. long, 3-15 mm. broad, puberulous on both surfaces, densely —
so along the veins below. Calyx 2-5-6 mm., puberulous.
MADAGASCAR : Basse valléee du Fiherenana, 50-200 m., Nov. 1933, Humbert 11535
(K, isotype of F. vincoides var. vestita). Forét de Besomaty entre le Fiherenana et VIsahaina (Mangoky), 750-800 m., Oct. 1933, Humbert 11272 (P). La Table Tuléar, Dequaire 329 (P). Au delta de la Linta, au nord d’Itampolo, 1-10 m., Aug. 1928, Humbert & Swingle 5382 (P).
The plants assembled here under the name O. minor are primarily distinguished from O. vincotdes by their much smaller leaves and somewhat smaller flowers,
Since they vary between themselves and merit study with more extensive material, —
it has seemed best to indicate variation as above, following Benoist, with some hesita- tion and doubt as to the validity of these taxa.
14 Oplonia linifolia (Benoist) Stearn, comb. nov.
Forsythiopsis linifolia Benoist in Notulae Syst. Paris 12 : 11 (1945).—Benoist in Humbert, Fl. Madag. 182 : 109, fig. 18 no. 14 (1967).
Shrub ; branches slender, glabrous ; spines absent. Leaves sessile, linear, the apex acute, the base cuneate, 5-10 mm. long, scarcely I mm. broad, glabrous. Flowers solitary ; pedicels 3-6 mm. long, glabrous. Calyx 2 mm. long, glabrous, the segments narrowly triangular. Corolla lilac, pubescent, the tube c.5 mm. long, the segments c.6 mm. long. Stamens slightly exserted ; filaments c.1 mm. long ; anthers c.1 mm. long. Style puberulous. Capsule c.1-5 cm. long.
MApDAGASCAR : Bassin de la Linta, plateau calcaire a l’est du delta, 200-250 m., August 1928, Humbert & Swingle 5507 (P, holotype).
This minute and rare species, only known from one locality in southern Madagascar, stands apart from all others by its very narrow linear leaves.
EXCLUDED SPECIES
Anthacanthus bispinosus Griseb. in Mem. Amer. Acad. Arts & Sci., New Ser. 8 : 525 (1862) from Cuba (Oriente) = Barleriola solanifolia (L.) Oerst. ex Lindau in Engler, Bot. Jahrb. 18 : 63 (1893), based on Barleria sclanifolia L., Sp. Pl. 2 : 636 (1753) ; cf. Lindau in Urban, Symb. Antill. 2 : 204 (1900). Oecersted described the genus Barleriola in Vidensk. Medd. Naturh. For. Kjébenh. 1854 : 136 (1855) but did not publish the combination Barleriola solanifolia often cited from there.
Anthacanthus repandus (Forst. f.) Nees in DC., Prodr. 11 : 462 (1847), based on Justicia repanda Forst. f., Florul. Ins. Austr. Prodr. : 3 (1786), from Tanna, New Hebrides = Evanthemum repandum (Forst. f.) R.Br. ex Roem. & Shult. in L., Syst. Veg., ed. nov. I : 175 (1817) ; cf. Seem., Fl. Vit. : 185 (1866) = Pseuderanthe- mum repandum (Forst. f.) Guillaumin in Ann. Mus. Colon. Marseille, ser. 6, 5-6 : 49
(1948).
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OPLONIA AND PSILANTHELE (ACANTHACEAE) 321
Anthacanthus stnuatus Nees in DC., Prodr. 11 : 462 (1847), based on Justicia sinuata Vahl, Symb. Bot. 2 : 11 (1791), nom. superfl. = Evanthemum longifolium (Forst. f.) Seem., Fl. Vit. : 185 (1866), based on Justicia longifolia Forst. f., Florul. Ins. Austr. Prodr. : 4 (1786), from Tanna, New Hebrides. = Pseuderanthemum longifolium (Forst. f.) Guillaumin in Bull. Mus. Hist. Nat. Paris, Ser. 2, 14 : 456 (1943).
Anthacanthus sprengelit Nees in DC., Prodr. 11 : 461 (1847), nom. superfl. based on Evanthemum acanthodes Spreng. in L., Syst. Veg., ed. 16, 1: 88 (1824), from Hispaniola = Scolosanthus acanthodes (Spreng.) Urban, Symb. Antill. 1 : 481 (1900). This belongs to Rubiaceae.
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——— Za
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WittraM T. STEARN, D.Sc., Sc.D. Department of Botany
British Museum (NaturAL History) CROMWELL Roap
Lonpon, S.W.7
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(b)
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2850. 3000.
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7.
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BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY
BOTANY | VOLUME 4
Cuticular studies as an aid to taxonomy, By C. A. STACE. 1965. _ :
The genus El phoglossum in the Indian Peninsula and Ceylon. By W.A. SLE AGE, 1967.
Fungi of recent Nepal expeditions, By F. L. Batrour-BROWNE, 1968.
A synopsis of Jamaican Myrsinaceae. By W. T; STEARN, 1969.
The Jamaican species of Coluwmnea and Alplectus (omer) By W. T. STEARN. a ,
fia aed: By H. SMITH. 1970.
A survey of the tropical genera Oplonia and Poilanthele (Acanthaceae). By W. T. STEARN. nyt ase
KW. EXELL
_ BULLETIN OF.
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ay | — Vol. 4 No. 8 : 1973,
jae ae eae ~
le >. C, tog aieass 6
_ ANGIOSPERMS OF THE ISLANDS OF THE“ GULF OF GUINEA (FERNANDO PO, PRINCIPE, S. TOME, AND ANNOBON)
BY
ARTHUR WALLIS EXELL
————
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BISGIOSPERMS OF THE ISLANDS OF THE GULF OF GUINEA (FERNANDO PO, PRINCIPE, S. TOME, AND ANNOBON)
By ARTHUR W. EXELL
| THE time has come when it seems useful to give an up-to-date list of the flowering plants of the Guinea Islands. There have been many supplements etc. since the | publication in 1944 of the Catalogue of the Vascular Plants of S. Tomé (with Principe and Annobon) (1),1 many changes recorded in the second edition of the Flora of West Tropical Africa (10, 12, 17, 18, 21), various publications by Portuguese authors (8, 9, 13), by Emilio Guinea (2, 3, 4, 5, 6, 7) and Antonio Escarré (19), as well as changes proposed by specialists in certain families or genera. Furthermore, there has been considerable additional collecting.
Fernando Po has now been added because the only way to get a clear picture of the flora of that island is to extricate the information from F.W.T.A. ed. 2 (10, 12, 17, 18, 21) and, as will be seen from the present work, this is at times incomplete.
FLORA OF FERNANDO PO
A clear distinction must be made between Fernando Po, only 32 km from the mainland, and the other three islands. The latter (Principe, S. Tomé and Anno- bon) can scarcely be classified as ‘oceanic islands’ and as I know of no appropriate term in use I shall call them ‘maritime islands’ and Fernando Po a ‘littoral island’. Of the three maritime islands only Annobon, the smallest and the furthest from the mainland, has some of the characteristics of an oceanic island, especially in its high proportion of Pteridophyta and of widespread species of Angiosperms.
The flora of Fernando Po is unlikely to have been much affected by the narrow strip of water separating it from the continent. Fairly convincing proof of this will begivenlater. I have noticed, however, in Africa that even a short stretch of water
May prove a barrier to many Leguminosae (except for littoral species). I attribute this to the comparatively heavy seeds unsuitable for wind distribution and not, as far as I know, much distributed by birds. On the three maritime islands, although the list of Leguminosae may seem extensive, if all the introduced species, the lit- toral species carried by ocean currents and the doubtfully indigenous cosmopolitan, pan-tropical or American weeds are excluded, the family is poorly represented com- pared with the adjacent mainland. Although Fernando Po has more Leguminosae, its flora still seems impoverished with respect to this family.
The Compositae, excellently distributed by wind, are quite well represented in Fernando Po but poorly so on the maritime islands when only species with some claim to be indigenous are considered. This apparent discrepancy is almost cer- tainly due to the presence of montane or submontane grassland (Moka, etc.) in Fernando Po and its almost complete absence in the maritime islands.
? For references, see Bibliography, pp. 333-336.
328 ANGIOSPERMS OF THE ISLANDS
Rubiaceae, Euphorbiaceae and Orchidaceae are well represented both in Fer- — nando Po and in the maritime islands.
Endemism in Fernando Po, as befits a littoral island, is much lower than in the maritime islands. For the relevant figures and the relationship between the islands see the Statistical Summary below.
As usual in floristic comparisons one can seldom isolate the statistical behaviour due to separate factors but in comparing these islands at least differences due to — climate are probably minimal, though Annobon is rather drier than the other three.
INTRODUCED PLANTS
All introduced plants of which I have seen records or specimens have been in- cluded. Species are marked with an asterisk in the Systematic List whenever there | is a high probability that they are not indigenous. There are of course very many — other species (mainly weeds) where the evidence is not conclusive. One reason | for including the introduced species in the list is to facilitate the naming of future — collections as a glance at the list may save a lot of trouble.
The introduced species fall into the following main categories :
(1) The successful introductions grown as crops for export, such as cocoa and coffee.
(2) Unsuccessful introductions for the same purpose. In the early days of | Portuguese colonization there are records of many attempted introductions (wheat, ; rye, etc.) at a period when there was no experience of tropical agriculture to act as } a guide. Many such species probably no longer exist on the islands.
(3) Species introduced as shade plants for cocoa or as ground cover.
(4) Species grown as local food plants Cae breadfruit, pineapples, etc.).
(5) Ornamental plants. |
(6) Accidentally introduced tropical heen weeds (usually clearly not in- _ digenous). |
(7) A few European or cosmopolitan weeds (such as Plantago major) very improb- ably indigenous.
There is of course another large category of weeds and widespread species which are probably introduced but could be indigenous.
It will be noticed that there are relatively far more introduced species for S. Tomé (and to a lesser extent for Principe) than for the other two islands. The reasons for this are historical. Both S. Tomé and Principe have been highly cultivated for about four centuries. By 1550 two-thirds of S. Tomé were said to be under cul- tivation and these islands were probably the first places where tropical agriculture — for export was carried on. Fernando Po was much less exploited and I do not know of any serious export from Annobon. Nor has this small island many introduced ornamental plants.
STATISTICAL SUMMARY
I have been interested for over forty years in the problem of comparing floras of similar or different sizes inhabiting areas of similar or different sizes in order to. obtain Quotients of Affinity and Quotients of Similarity (see especially Exell and
OF THE GULF OF GUINEA 329
Wild in Kirkia 2 : 108-130 (1961)). In the work cited we discussed in some detail
the various quotients which have been employed at different times and the justifi-
cation for the use of the simple formula referred to as QA (EW) which is the ex- pression of the number of species common to two floras as a percentage of the smaller one. This is used in the present work and is here simply called OA (Quotient of Affinity). After further thoughts about this formula (since 1961) I have now
| the impression that it tends to become less meaningful when a very small flora is
compared with a very large one. In the ultimate case, when the flora of an island or region is compared with the flora of the whole of the rest of the world we get a OA which is exactly the inverse of the proportion of endemism in the small flora (i.e. with no endemics the QA becomes 100). Thus when there is a very great disparity of size between the floras under consideration the relative abundance or absence of endemism begins to loom very large in the QA figures and to obscure other factors.
Where the OA employed is at its most useful and most accurate is in comparing the flora of a given region with those of neighbouring regions not differing greatly from it or from each other in size. I have borne this in mind in the statistical comparison of the floras of the four islands with the mainland flora and confined this comparison to the Southern Nigeria and West Cameroon regions, and a com- bination of the two (which already begins to get too large). To extend the com- parison to the whole tropical West African flora would be to approach the situation I have indicated above where the figures would do little beyond reflecting the differ- ent degrees of endemism on the different islands and be merely an immensely laborious way of arriving at such a result.
The Quotients of Affinity of the islands with one another and with the mainland regions are set out in Table I. It should be stated that the figures given in this
TABLE I[
Quotients of Affinity of the flora of the islands of the Gulf of Guinea and the adjacent mainland
FP P ST A SN wc SN+ WC FP. — 590 47°8 56°25 68:5 715 89:0 P 59°0 aa 67°9 44°7 73°2 60-2 78-9 ST 47°8 67°9 = 715 64°4 52°6 719 A 56°25 44°7 715 — 721 55°8 779
FP, Fernando Po, 1105 species; P. Principe, 314 species; ST, S. Tomé, 601 species; A, Annobon, 208 Species ; SN, Southern Nigeria, ‘S. Nig.’ of the Flora of West Tropical Africa; WC, West Cameroon, Brit. Cam.’ ‘[Br.] Cam.’ and ‘W. Cam.’ of the various volumes of the Flora of West Tropical Africa.
table are not absolutely correct because of slight adjustments made to the sys- tematic list right up to the time of going to press. The mathematical discrepancy is, however, negligible, especially in view of the considerable and unavoidable subjective factor involved in choosing which taxa to exclude (such as possibly introduced or too doubtfully identified species).
If the QA figures between the various islands are compared with similar statistics given in the past (see especially Exell and Wild, Joc. cit. Table I, column QA (EW))
330 ANGIOSPERMS OF THE ISLANDS
it will be seen that relationships between Principe, S. Tomé and Annobon have —
not altered appreciably although the total number of species recorded for each
island has increased. The figures for Fernando Po have, however, changed sig-
nificantly. This is mainly due to a complete re-appraisal of the Fernando Po
flora in the second edition of the Flora of West Tropical Africa. This has resulted
in a great increase in the QA’s between Fernando Po and the three maritime islan
correlated with a marked decrease in the percentage of species considered endemic
to Fernando Po (from 12 per cent in 1944 to 3-6 per cent now) ; this is due partly
to taxonomic changes (reduction of endemics to synonymy), partly to the dis-
covery on the mainland of species formerly thought to be endemic and partly to an increase of non-endemic species recorded. The same process has occurred to a- lesser extent in the other three islands (fairly significantly in Annobon) as Table Ite |
shows. 4 |
TaBLeE II gq
Percentage of species considered as endemic in the flora of the islands of the Gulf of Guinea
1944 1971 Fernando Po 12:0 3°6 (3°8) Principe 27 9°9 (13°7) S. Tomé 19°4 15°4 (17°8) Annobon 14°8 9-9 (1)
The figures in brackets in the last column are those obtained if species occurring on more than one island but not elsewhere are reckoned as endemics.
The decrease in Annobon is due almost entirely to the fact that the recorded non- endemic flora has nearly doubled since 1944.
Relatively few new endemic species have been described to counterbalance the tendencies mentioned above. There is little hope, however, that the present figures represent anything like finality. There are many more discoveries still to make.
In the past there have not been sufficiently accurate data available to work out the QA’s between the various islands and selected regions of the mainland. This has now been largely remedied with regard to Southern Nigeria and West Cameroon by the publication of the second edition of the Flora of West Tropical Africa.
It is perhaps worth mentioning that the statistics were first worked out sepa-
rately for the Dicotyledons and Monocotyledons but no significant difference was _
found. The Monocotyledons do not behave statistically as a separate unit. The Orchidaceae, for example, with many endemics, behave much as the Melastomata- ceae, Rubiaceae and Euphorbiaceae while the Gramineae and Cyperaceae behave very like the Leguminosae and the Compositae.
It is now easiest to consider each of the four islands separately.
Fernando Po
The QA of Fernando Po with the adjacent coastal region as a whole (SN + WC of Table I) is 89. This is very high indeed. For comparison, the highest QA
OF THE GULF OF GUINEA 331
obtained in a statistical analysis of a sample of the flora of the region covered by Flora Zambesiaca was 87 between the Eastern Division of Rhodesia and the southern part of Mozambique, two adjacent and floristically closely related regions on the mainland of Africa. The only QA higher than 89 that I have yet recorded is 92-4 between the floras of Guernsey and Alderney (Channei Islands). This OA of 89 between Fernando Po and SN + WC gives statistical proof of what has long been recognized, that Fernando Po does not have an insular flora but is floristically part of the mainland.
The OA 71-5 of Fernando Po with West Cameroon is slightly higher than the OA 68-5 between the island and Southern Nigeria, indicating a fairly equal deriv- ation of the flora of Fernando Po from West Cameroon and Southern Nigeria, but, if the species contributing to these two QAs are studied in detail, the impression is gained that the slight difference in favour of West Cameroon should really be greater than it is for it is certainly based on more significant distribution patterns, Among the species common to Fernando Po and Southern Nigeria there are far more widespread species and weeds, and many of these probably occur also in West Cameroon, which appears to be somewhat under-collected.
Principe
The QA of this island’s flora with that of the adjacent coastal region as a whole (SN + WC) is 78-9, showing a high affinity between the two, but clearly indicating that the flora of Principe is more insular than that of Fernando Po. The QA with Southern Nigeria is 73:2 and that with West Cameroon 60-2, very considerably different from the figures for Fernando Po, for they show that Southern Nigeria is a more important source than West Cameroon for the flora of Principe. The OA of 59-0 with Fernando Po and 67-9 with S. Tomé are both fairly high, as com- pared for example with a QA of 47-8 between Fernando Po and S. Tomé, and seem to indicate some derivation of the flora of Principe from the two larger islands
between which it lies.
S. Tomé
The QA between this island and the adjacent coastal region as a whole (SN + WC) is 71-9, appreciably lower than the islands nearer the mainlaind and probably fairly typical of an insular flora with considerable endemism. The QA 64:4 with Southern Nigeria is considerably higher than the QA 52:6 with West Cameroon, indicating that Southern Nigeria is the more important as a source of the flora of S. Tomé.
Annobon
This, although further from the mainland than S. Tomé, has a higher QA, 77:9, - with the adjacent coastal region as a whole (SN + WC). Here we begin to get, I believe, a partial ‘oceanic island’ effect, a somewhat impoverished flora with a higher proportion of widespread species and rather fewer endemics. The affinity with Southern Nigeria (QA 72-1) is much higher than that with West Cameroon
(QA 55°8).
332 ANGIOSPERMS OF THE ISLANDS
A special feature of the Annobon figures is the high QA, 71-5, with S. Tomé, and this appears to be a real affinity, indicating that the small Annobon flora has been partly derived from the much larger and comparatively near-by island of S. Tomé,
The QA of 56:25 between Annobon and Fernando Po seems abnormally high for that between the two most widely separated islands, especially when it is compared with OA 47-8 between S. Tomé and Fernando Po. Detailed study indicates that is
partly an ‘oceanic island’ effect, the flora of Fernando Po being more than five
times the size of that of Annobon, and partly the result of a few anomalous dis- tributions, which may not all be genuine.
General
There has been no change in the general hypothesis that in the main each island received its flora independently from the adjacent regions of the mainland, each receiving a considerably different selection so that the QAs with the mainland are usually higher than those between one island and another. Each island mostly produced its own endemics. There are relatively few cases of species endemic to more than one island and some of these may not survive future investigation and further collecting (i.e. they may yet be found on the mainland). A few, especially those common to S. Tomé and Annobon, are probably genuine.
I hope that it will be apparent that the comparisons given above, and in fact most of the discussion, would be almost impossible without using the QA concept. Whether the OA used here is the best one available is not so certain and any sug- gested improvement will always merit examination.
SYSTEMATIC LIST
This list is mainly a compilation of published records of taxa from the islands but I have been fortunate in getting assistance, for which I am most grateful, from specialists in a number of families and from other colleagues. Acknowledgements are made after the bibliography. The list is very condensed in form but it is possible from the bibliographical references and some short notes appended under the species to obtain fairly full nomenclatural data and records of specimens.
An asterisk * indicates an introduced species. It is often impossible to be certain whether a taxon such as a widespread weed is indigenous or not. It is fairly certain that all the species marked * are introduced but it is by no means certain that all the species not so marked are truly indigenous.
The bibliographical references are given by number in the central column. To avoid unnecessary complication, where infraspecific categories are now recognized, the older bibliographical references have been placed where they almost certainly belong. This usually consists of putting the reference under the typical subspecies or variety when only the specific name was originally given in the work cited. The same has been done in a few cases where it seems quite clear to which infraspecific taxon (other than the typical one) the work cited must refer even when this was not mentioned or had not been established at the time.
tt ee ee
OF THE GULF OF GUINEA 333
The letters FP, P, ST, A in the right-hand column indicate Fernando Po, Prin- cipe, S. Tomé and Annobon respectively and the letter E in the right-hand margin indicates an endemic taxon.
In the case of Fernando Po (FP) I have indicated all records additional to or omitted from the second edition of the Flora of West Tropical Africa (except in the case of introductions). These fall into very different categories. Some are evident accidental omissions (when, for example, the type is from Fernando Po), some are new and probably correct records, others are very doubtful indeed (especially those to which I have given question marks).
ANNOTATED BIBLIOGRAPHY OF WORKS PUBLISHED FROM 1944 ONWARDS
References to works prior to 1944 will mainly be found in 1, 10, II, 12, 17, 18, 21.
Different numbers are given to the separate volumes (or parts) of the second edition of the Flora of West Tropical Africa in order to maintain chronological sequence and to facilitate reference.
I. A. W. Exell. Catalogue of the Vascular Plants of S. Tomé (with Principe and Annobon). 1944. References to works prior to 1944, species references and fairly full synonymies.
/ Contributors (Angiosperms) other than the author were: Leguminosae (partly
by E. G. Baker and D. Hillcoat) ; Rubiaceae (by G. Taylor) ; Apocynaceae (by W. R. Philipson) ; Asclepiadaceae (partly by C. Norman); Loganiaceae (by W. R. Philipson) ; Acanthaceae (partly by E. Milne-Redhead) ; Labiatae (by G. Taylor) ; Euphorbiaceae (partly by J. Hutchinson) ; Ulmaceae (by G. Taylor) ; Moraceae (by G. Taylor) ; Urticaceae (by G. Taylor) ; Cannabaceae (by G. Taylor) ; Mono- cotyledones (by J. E. Dandy, with V. S. Summerhayes assisting with Orchidaceae).
2. E. Guinea Lopez. Ensayo Geobotdnico de la Guinea Continental Espanola. 1946. The relevant part is the ‘Catalogo Floristico’, pp. 218-368, in which Fernando Po records are given and some references made to the other islands.
3. E. Guinea (Lopez). Agrostigrafia fernandina e hispano-guineanae. An. Agr. Territ. Espan. del Golfo de Guinea 1947, publ. 13: 95-164. 1948.
4. E. Guinea (Lopez). Juncaceae, Cyperaceaeque Fernandopoinae. An. Jard. Bot. Madrid 9 : 335-360. 1950.
5. E. Guinea (Lopez). Melastomataceae Fernandopoanae. Bol. Real Soc. Esp. Hist. Nat. 49: 175-187. 1951.
6. E. Guinea (Lopez). Asteraceae (Compositae) Fernandopoanae. An. Inst. Bot. Cavan. I : 301-340. I9g5I.
7. E. Guinea Lopez. Esquema de la Vegetacién de Fernando Péo. Confer- encia Internacional de Africanistas Occidentales (1951). Circular Paper.
IQ5I.
334 ANGIOSPERMS OF THE ISLANDS
This includes some lists of plants collected by Guinea in Fernando Po. The work is repeated in Spanish (41-45), French (52-56), English (62-66) and Portu- guese (73-77). The Araliaceae (Polyscias fulva, Schefflera mannii, S. ledermannit) are omitted from the English version.
8. L.G.Sobrinho. Vascular Plants from S. Tomé. Port. Act. Biol. (B), 3: 392- ]
40I. 1953.
The date on the title-page of the separate is 1952 but it was apparently published ' in 1953 (see p. 392). This work records plants collected in Principe and S. Tomé —
by d’Oliveira and Noronha in 1951.
;
g. L.G.Sobrinho. Vascular Plants from Ano Bom and Principe Islands. Port,
Act. Biol. (B) 4: 177-190. 1953.
This records plants (now at LISU) collected by F. Newton in Annobon in 1893 and a few specimens collected by de Souza Jr in Principe in 1880. The existence of these specimens was unknown to me at the time of the publication of (1). There are many new records from Annobon.
10. J. Hutchinson and J. M. Dalziel. Flora of West Tropical Africa I (1), ed. 2, |
revised by R. W. J. Keay. 1954. Authors of various families other than Keay are indicated in the work. I am much indebted to this excellent second edition and the Fernando Po records have been largely extracted from this and the succeeding volumes.
11. A.W. Exell. Supplement to the Catalogue of the Vascular Plants of S. Tomé (with Principe and Annobon). 1956.
This brought (1) up to date by including the records in (8) and (9) and a consider- able collection made in S. Tomé by Espirito Santo. Acknowledgement to col- laborators is made in the introduction. This opportunity is taken to correct two formulae on p. 4 in the discussion on statistics. These should read: (1) the increase in number of species by adding area B to area A = T — a = a log, [(A + B)/A] and (2) similarly the increase of species by adding area A to area B = T—0= a log, [(A + B)/B].
12% VAS, (10).. VOh.. Tt (2)e0 TO50:
13. H. Lains e Silva. Sado Tomé e Principe e a Cultura do Café. 1958.
This important work has much information about the vegetation of the two islands, the ecology and the introduced species. It should be noted that the ‘ Nomes latinos das especies vegetais’ on p. 405 does not refer only to species occurring in the islands but includes a few species from other parts of the world mentioned in the text. There area number of newrecords. I have not myself seen the specimens of these.
14. A. W. Exell and A. Rozeira. Aditamento a Flora das Ilhas de S. Tomé e do Principe. 6 Conferencia Internacional dos Africanistas Ocidentais (Ilha de S.
Tomé, 1956) 3: 77-92. 1958.
: 4
ae ;
OF THE GULF OF GUINEA 335
Spermatophyta by A. W. Exell. This mainly consists of new records from the collections of A. Rozeira in Principe and S. Tomé. I was not shown any proofs of this publication and regret a number of typographical errors for which I was not responsible.
15. A.W. Exell. Additions to the Flora of S. Tomé and Principe. Bull. I.F.A.N. 21, sér. A, 2: 439-476. 1959. Mainly an account of new records from collections by Th. Monod made in Principe and S. Tomé in 1956.
16. A. W. Exell. Angiosperms of the Cambridge Annobon Island Expedition. Bull. Br. Mus. nat. Hist. (Bot.), 3 : 93-118 (1963).
This listed new and interesting plants collected by F. A. Melville and T. C. Wrigley in Annobon in 1959. Certain families not by the author were contributed as follows: Loganiaceae (by J. Lewis) ; Solanaceae (by W. T. Stearn) ; Piperaceae (by W. T. Stearn) ; Loranthaceae (by W. T. Stearn) ; Orchidaceae (by V. S. Summerhayes) ; Gramineae (by W. D. Clayton).
17. As (10) but revised by F. N. Hepper. Vol. 2. 1963. 18. As (17). Vol. 3 (x). 10968.
19. A. Escarré. Aportaciones al Conocimento de la Flora de Fernando Poo. I— Araliaceae, Umbelliferae. 1968. II—Piperaceae, Urticaceae. 1969.
20. W.W. Sanford. The Orchid Flora of Equatorial Guinea in relation to that of West Africa. Mutt. Bot. StSamml. Miinch. 10 : 287-298. 1971.
wr. As (18). Vol. 3 (2). 1972.
22. J. H. P. de Barros Ferreira. Contribuicéo para o Estudo das Malvaceas de S. Tomé e Principe. Garcia de Orta, 11 : 27-55. 1963.
23. J. H. P. de Barros Ferreira. Contribuigéo para o Estudo das Begoniaceas de S. Tomé e Principe. Garcia de Orta, 13 : 525-543. 1965.
24. J.H.P.de Barros Ferreira. Contribuicéo para o Estudo das Melastomataceae de S. Tomé e Principe. Garcia de Orta, 16: 69-86. 10968. Includes Tristemma thomensis Barros Ferreira published by the author on p. 66 of the same volume.
25. J. do Espirito Santo. Nomes crioulos e vernaculos de algumas plantas de S. Tomé e Principe. Bolm cult. Guiné port. 24: 193-211. 1969.
In spite of the title the plants recorded are all from S. Tomé. The author states in a footnote to the first page of (27) that the ‘prospeccdo botanica’ of 1968 did not get to Principe. Numerous spelling mistakes, etc. are mainly corrected in the Corrigenda to (27).
26. J. do Espirito Santo. Algumas Plantas medicinais e venenosas de S. Tomé e Principe. Bolm cult. Guiné port. 24: 917-940. 1969. Notes to (25) apply equally to this.
336 ANGIOSPERMS OF THE ISLANDS
27. J. do Espirito Santo. Aditamento a Flora de S. Tomé. Bolm cult. Guiné
port. 25: 489-5II. 1970.
New records are mainly introduced species but there are a few interesting in- digenous ones. Erythroxylum emarginatum, Ilex mitis and Alangium chinense add new families to those recorded from S. Tomé. The specimens are at LISJC and most of them also at LISC. This work corrects many mistakes in (25) and (26) but — adds a few more most of which are fairly obvious.
ACKNOWLEDGEMENTS
S. Balle: Loranthaceae. J. Léonard : Euphorbiaceae.
J. P. M. Brenan: various. Maria Candida Liberato : Leguminosae. J. F. M. Cannon: Araliaceae, Umbelliferae. J. B. Marshall: Compositae.
W. D. Clayton: Gramineae. E. Milne-Redhead : various.
J. E. Dandy: various. D. Napper: Acanthaceae.
A. Escarré: Piperaceae, Urticaceae and R. M. Polhill : Leguminosae.
various. J. Raynal: Cyperaceae.
A. & R. Fernandes: Barringtoniaceae, Mela- stomataceae, Onagraceae.
F. N. Hepper: various.
C. Jeffrey : Compositae, Cucurbitaceae.
E. Launert: various.
N. K. B. Robson: Hypericaceae.
R. Ross: for much help throughout. W. W. Sanford : Orchidaceae.
P. Taylor: Amaranthaceae.
NEW NAMES PUBLISHED IN THIS WORK
Pauridiantha camposu (G. Tayl.) Exell, comb. nov. (Rubiaceae). Psychotria hierniana Exell, nom. nov. (Rubiaceae).
DICOTYLEDONES
I. ANNONACEAE
ANNONA ARENARIA Thonn. * ANNONA CHERIMOLA L.
*ANNONA GLABRA L. * ANNONA MURICATA L. * ANNONA RETICULATA L. * ANNONA SQUAMOSA L. ARTABOTRYS MACROPHYLLUS Hook. f. ARTABOTRYS Sp. FP: Miuldbraed 6412 *ASIMINA TRILOBA (L.) Dunal *CANANGA ODORATA (Lam.) Hook. f. & Thomson CLEISTOPHOLIS PATENS (Benth.) Engl. & Diels FP: Bokoko, Burchardt. ENANTIA CHLORANTHA Oliver MONODORA BREVIPES Benth. MONODORA MYRISTICA (Gaertn.) Dunal Not recorded from FP in tro. MOoNODORA TENUIFOLIA Benth. MONODORA ZENKERI Engl. Not in Io.
Not recorded from FP in ro.
I Sper
25 SL
Ero Si
1, 2.16, 25:26. -Fe PP Sik
1, 16,.25 PSEA
I, 25 ST A.
2, 10 FP E Be
I ee
2, 11;-25 Bey Sr
2 FP
Io FP
T.2,t0 EPP
1, 2,7, 25, 26 EP PSE
2. 10 FP
2 FP
OF THE GULF OF GUINEA
OXYMITRA GRACILIPES Benth. OXYMITRA GRACILIs (Hook. f.) Sprague & Hutch. Unona millenii Engl. & Diels Not recorded from FP in Io. OXYMITRA HIRSUTA (Benth.) Sprague & Hutch. POLYALTHIA OLIVERI Engl. Popowla sp. FP: Bokoko, Mildbraed 6869 UvaRIA sp. UVARIODENDRON CONNIVENS (Benth.) R. E. Fries Uvaria connivens Benth. UVARIODENDRON FuSCUM (Benth.) R. E. Fries Uvaria fusca Benth. UVARIODENDRON LETESTUI (Pellegrin) R. E. Fries Uvaria mannii Hutch. & Dalz. XYLOPIA AETHIOPICA (Dunal) A. Rich. XYLOPIA AFRICANA (Benth.) Oliver FP: Moka, Mildbraed 7025. Not recorded from FP in 10. XYLOPIA QUINTASII Engl. & Diels X. striata Engl. Not recorded from FP in to.
2. MONIMIACEAE
GLOSSOCALYX LONGICUSPIS Benth. XYMALOS MONOSPORA (Harv.) Baillon ex Warb.
3. LAURACEAE
*CINNAMOMUM BURMANNI (C. & T. Nees) Blume *CINNAMOMUM CAMPHORA (L.) T. Nees & Eberm. *CINNAMOMUM ZEYLANICUM Garc. ex Blume *PERSEA AMERICANA Miller
P. gratissima Gaertn. f.
4. HERNANDIACEAE HERNANDIA BENINENSIS Welw. ex Henriq.
5. MYRISTICACEAE
COELOCARYON PREuSSII Warb. FP: Bokoko, Mildbraed 6856. Not recorded from FP in Io. *MyYRISTICA FRAGRANS Houtt. PYCNANTHUS ANGOLENSIS (Welw.) Warb. P. kombo (Baillon) Warb. Not recorded from FP in tro. PYCNANTHUS MICROCEPHALUS (Benth.) Warb. STAUDTIA KAMERUNENSIS Warb. FP: Bokoko, Mildbraed 6883, on which the record in 2 is probably based. Not recorded from FP in ro. STAUDTIA PTEROCARPA (Warb.) Warb. STAUDTIA STIPITATA Warb. S. gabonensis Warb. Not recorded from FP in 1o.
Io
es oN
2 25 O ie o2 LO
4 XN al °
2, 10 ae)
I Te25
E213 E133 10, 25 2
15:2,-7,,10
¥,.16;-1Y
2, 10
» 25
FP LD ng Pr ST FP ST ba FP FP PsP FPP St FP Sst FP FP ST ST lige & FP PSTA FP ST FP SL PEO Sf BE EP. ST FP
SE Me
338 ANGIOSPERMS OF THE ISLANDS
6. RANUNCULACEAE
CLEMATIS HIRSUTA Guillemin & Perrottet 8, 12,15 CLEMATIS SIMENSIS Fresen. Io
C. altissima Hutch. 2 RANUNCULUS EXTENSUS (Hook. f.) Schube ex Engl. 2, 10
‘Rubus extensus Schube ex Engl.’, Index Kewensis, Suppl. 1: 370 (1906), cited from FP but entirely mythical; Ranunculus extensus was undoubtedly
intended. THALICTRUM RHYNCHOCARPUM Dillon & Rich. 10 T. mannii Hutch. 25:9 7. NYMPHAEACEAE NYMPHAEA LOTUS L. 10 N. lotus var. sinoeensis Stapf 2 8. MENISPERMACEAE CHASMANTHERA DEPENDENS Hochst | CISSAMPELOS OWARIENSIS Beauv. ex DC. ite) DIOSCOREOPHYLLUM VOLKENSII var. FERNANDENSE (Hutch. & Dalziel) Troupin 12 D. fernandense Hutch. & Dalziel 2 D. tenerum var. fernandense (Hutch. & Dalziel)
Troupin 10 JATEORHIZA MACRANTHA (Hook. f.) Exell & Mendonga 2, 10 PENIANTHUS LONGIFOLIUS Miers 2, 10 STEPHANIA ABYSSINICA (Dillon & Rich.) Walp. var.
ABYSSINICA 2, 10 STEPHANIA CYANANTHA Welw. ex Hiern 12 STEPHANIA DINKLAGEI (Engl.) Diels 15 STEPHANIA LAETIFICATA (Miers) Oliver 2
Not recorded from FP in 10. The holotype, Mann 236, is from FP.
SyYRRHEONEMA FASCICULATUM Miers 2°10 TRICLISIA CORIACEA Oliver 2-10 TRICLISIA MACROPHYLLA Oliver 24TO
Recorded as endemic to FP in 10, but the distribution has since been found to extend to Sierra Leone and Cameroons Mt.
Q. ARISTOLOCHIACEAE
ARISTOLOCHIA LITTORALIS Parodi A. elegans Mast. II PARARISTOLOCHIA GOLDIEANA (Hook. f.) Hutch. & Dalziel 2, 10
I0. PIPERACEAE
PEPEROMIA FERNANDOPOIANA C.DC. 2, 7; 10,15, 19 PEPEROMIA HYGROPHILA Engl. 1, 10, 30
P. annobonensis Mildbr. 1; 2 PEPEROMIA KAMERUNANA C.DC. Io, 19 ‘Peperomia lucida’ 7
Probably P. pellucida was intended.
OF THE GULF OF GUINEA
PEPEROMIA MANNII Hook. f. ex C.DC. PEPEROMIA MOLLERI C.DC. PEPEROMIA PELLUCIDA (L.) Kunth
PEPEROMIA REFLEXA (L.f.) A. Dietr. PEPEROMIA THOMEANA C.DC. PEPEROMIA VACCINIFOLIA C.DC. PEPEROMIA sp.
PIPER CAPENSE L.f.
PIPER GUINEENSE Schumacher
P. clusti C.DC. PIPER UMBELLATUM L.
P. subpeltatum C.DC.
II. PAPAVERACEAE *ARGEMONE MEXICANA L.
I2. FUMARIACEAE *FUMARIA MURALIS Sonder ex Koch
I3. CAPPARACEAE
BUCHHOLZIA CORIACEA Engl.
B. macrophylla sensu Espirito Santo CAPPARIS TOMENTOSA Lam. CLEOME RUTIDOSPERMA DC.
C. ciliata Schumacher
‘GYNANDROPSIS GYNANDRA (L.) Briq. Pedicellaria pentaphylla Schrank Not recorded from FP in to. RitcuHIEA ERECTA Hook. f. RITCHIEA POLYPETALA Hook. f. R. fragrans sensu Mildbr. ? R. cf. glossopetala Gilg FP: Bokoko, Burchardt. R. obanensis Hutch. & Dalziel
I4. CRUCIFERAE
*BRASSICA JUNCEA (L.) Czerniak *BRASSICA OLERACEA L. *CAPSELLA BURSA-PASTORIS (L.) Medik. CaRDAMINE AFRICANA L. C. africana var. pubescens Hook. f. CARDAMINE HIRSUTA L. *CoRONOPUS DIDYMUs (L.) Smith *DIPLOTAXIS TENUISILIQUA Delile *LOBULARIA MARITIMA (L.) Desvaux *RAPHANUS SATIVUS L. Rorippa HUMIFUSA (Guillemin & Perrottet) Hiern Nasturtium humifusum Guillemin & Perrottet Not recorded from FP in ro.
2,.10;.19
L,akO. 10
tier Pale faa fs ee @ 19, 25, 26
I, 15
I
Io, 19
I
i 2 .Or LT 216
To 27 elOn LO: 25
2
[107 .5,, 1G, 25, 26
2
I, 25; 26
27
25
I
16
T0219; 10,1 te 25
I
2, 10
135 2
I
Ey 10; 54 2
2.7 T6 1 Oe
I
FP | au oe aaa top ede ate ST ST. FP A PPAR St BRP ST FPPSTA ST oT Si ST FPP SLA 1s aa & 1 2 FP og 8 FP Sr. 1 oo Sole ST ST ST of ST FP
339
340 ANGIOSPERMS OF THE ISLANDS
RoriIpPa INDICA (L.) Hiern Nasturtium sinapis (Burm. f.) O. E. Schulz Rorippa sinapis (Burm. f.) Ohwi & Hara
*RORIPPA NASTURTIUM-AQUATICUM (L.) Hayek Nasturtium officinale R.Br.
*SINAPIS ALBA L.
I5. VIOLACEAE
HyYBANTHUS ENNEASPERMUS (L.) F. v. Muell. RINOREA ANGUSTIFOLIA (Thouars) Baillon R. molleri Brandt R. ardisiiflora sensu Guinea See Tennant in Kew Bull. 16: 412 (1962). RINOREA CHEVALIER! Exell RINOREA DENTATA (Beauv.) Kuntze R. bipindensis Engl. RINOREA INSULARIS Engl. RINOREA LONGISEPALA Engl. Not recorded from FP in Io. RINOREA THOMENSIS Exell VIOLA ABYSSINICA Steudel ex Oliver V. emirnensis (Engl.) Guinea
16. POLYGALACEAE
CARPOLOBIA GLABRESCENS Hutch. & Dalziel
I7. CRASSULACEAE
*BRYOPHYLLUM PINNATUM (Lam.) Oken B. germinans sensu Guinea CRASSULA ALSINOIDES (Hook. f.) Engl. Not recorded from FP in 10. The type is from FP. KALANCHOE CRENATA (Andr.) Haw.
KALANCHOE CRENATA Var. VEREA (Jacq.) Cufod. in Bull.
Jard. Bot. Brux. 27 : 714 (1957). ST: Menyharth 24 (WU).
KALANCHOE LACINIATA (L.) DC. FP: Moka, Exell 796 (BM.). in Io.
18. HYDRANGEACEAE
*HYDRANGEA Sp.
19. PODOSTEMACEAE
TRISTICHA TRIFARIA (Bory) Sprengel T. hypnoides (St.-Hil.) Sprengel
20. CARYOPHYLLACEAE
CERASTIUM INDICUM Wight & Arn. DRYMARIA CORDATA (L.) Willd.
SAGINA ABYSSINICA Hochst. ex A. Rich. STELLARIA MANNII Hook. f. *STELLARIA MEDIA (L.) Vill.
FP : Guinea 2057.
Not recorded from FP
16
Eire
15
15, 25, 26 La
2
2, 10
2, 10
E> 21,258,286
2,7
110, tts 10027
Io
1, 2,0; 10, Ff,
13, 25 2, 10
¥,°%6,37;. 43
I
ST FP ST FP FP ST A ST FP P FP ST FP FP y : FPPSTA & FP ; I sTA @ ST FP . | ST? FP FP FP PST ‘ FP FP ST FP ST
—
OF THE GULF OF GUINEA
21. AIZOACEAE SESUVIUM PORTULACASTRUM (L.) L.
22. PORTULACACEAE
PORTULACA OLERACEA L.
PORTULACA QUADRIFIDA L. *TALINUM TRIANGULARE (Jacq.) Willd.
23. POLYGONACEAE
*ANTIGONON LEPTOPUS Hook. & Arn.
A. covdatum Martens & Galeotti POLYGONUM NEPALENSE Meissner
P. alatum (Buch.-Ham. ex D. Don) Sprengel POLYGONUM NYIKENSE Baker POLYGONUM SALICIFOLIUM Brouss. ex Willd. PoLYGONUM SENEGALENSE Meissner RUMEX ABYSSINICUS Jacq.
Not recorded from FP in to.
24. PHYTOLACCACEAE
PHYTOLACCA DODECANDRA var. APICULATA Baker & Wright
25. CHENOPODIACEAE
CHENOPODIUM ALBUM L. *CHENOPODIUM AMBROSIOIDES L.
26. AMARANTHACEAE
ACHYRANTHES ASPERA L. Not recorded from FP in fo. ACHYRANTHES BIDENTATA Blume AERVA LANATA (L.) Juss. ex J. A. Schultes FP: Guinea 1123. Not recorded from FP in to. ALTERNANTHERA MARITIMA (Mart.) St.-Hil. Telanthera maritima (Mart.) Moq. FP: Guinea 875. Not recorded from FP in Io. _ALTERNANTHERA PUNGENS Kunth A. repens (L.) Link, non A. repens J. F. Gmelin ALTERNANTHERA SESSILIS (L.) R. Br. ex DC., Cat. Hort. Monspel.: 77 (1813). AMARANTHUS GRAECIZANS L. A. blitum sensu auctt. The record in 13 is probably A. graecizans. AMARANTHUS HYBRIDUS subsp. INCURVATUS (Timeroy ex Gren. & Godron) Brenan A. paniculatus L. Not in ro. AMARANTHUS SPINOSUS L. AMARANTHUS VIRIDIs L. Not recorded from FP in to. *CELOSIA ARGENTEA L. Probably introduced.
2
By 2) 10; 23:55; 16, 25
13 L572, 0.0L, 10.825
t;. 41,13
I
Lan
dg 2.35 255.20
Lal -
Zz, TOe25
2,10 2 (‘Aurea’)
16
I, 14
t; 11,%3 1,-0:..10, 11.16
13
2
2,10} 13;,10 Tene. LO
1,2
ST FP SF oT FP. -St EE SSE EF FP A ST EP: Sr ST ST I ogee FP STA PP FP FP PSEA esr FPPSTA ST FP FPPSTA EP SLA Pes cot
341
342
CELOSIA BONNIVAIRII Schinz CELOSIA LAXA Schumacher CELOSIA LEPTOSTACHYA Benth. CELOSIA TRIGYNA L. CYATHULA ACHYRANTHOIDES (Kunth) Mogq. CYATHULA CYLINDRICA var. MANNII (Baker) Suesseng. C. mannii Baker CYATHULA FERNANDOPOENSIS Suesseng. & Friedrich CYATHULA PEDICELLATA C. B. Clarke CYATHULA PROSTRATA (L.) Blume DEERINGIA AMARANTHOIDES (Lam.) Merr. *GOMPHRENA GLOBOSA L. *TRESINE HERBSTII Hook. NOTHOSAERVA BRACHIATA (L.) Wight FP: Escarré 361, 373, 375. Not recorded from FP in 10. PHILOXERUS VERMICULARIS (L.) Beauv. FP: Escarré 360. Not recorded from FP in Io. PUPALIA LAPPACEA (L.) Juss. Not recorded from FP in Io. SERICOSTACHYS SCANDENS Gilg & Lopr.
27. BASELLACEAE
*BASELLA ALBA L. Probably introduced.
28. GERANIACEAE
GERANIUM ARABICUM Forssk. G. simense Hochst. ex A. Rich. See Laundon in Bol. Soc. Brot., sér. 2, 35 : 59 (1961).
29. OXALIDACEAE OXALIS CORNICULATA L,
*OXALIS CORYMBOSA DC.
30. AVERRHOACEAE * AVERRHOA CARAMBOLA L,
31. BALSAMINACEAE
*IMPATIENS BALSAMINA L. IMPATIENS BUCCINALIS Hook. f, IMPATIENS FILICORNU Hook. f. (Omitted from index to 10; text p. 162.)
IMPATIENS HIANS Hook. f. IMPATIENS KAMERUNENSIS Warb. IMPATIENS MACROPTERA Hook. f. IMPATIENS MANNII Hook. f.
I. affinis Warb. IMPATIENS MANTEROANA Exell IMPATIENS NIAMNIAMENSIS Gilg
I. dichroa Hook. f. ex Gilg IMPATIENS SAKERANA Hook. f. IMPATIENS TAMSIANA Exell IMPATIENS THOMENSIS Exell
ANGIOSPERMS OF THE ISLANDS
Io
2, 10
beer Aor Le pony bs Oa At I-10
2:10
Io
2
Io
Io
Te LOs 13. 16
I, 19 Ts 25/0; 0325
Io
2,4; LO
Lal
2 TOn3U 12: 13 WEY, 3; 25
Lal
2; 135, 25
13 : Oe 2, 10
FP FP os at fo FP A PP EP FP FP FPP ST A tod BE oS) ST FP EP. St FP Sia boy STA FP FPP STS PST FP ‘SF st st EP. FP FP FP FP P FP FP P SE
‘en . yp oft
ico co am
eo
*7AWSONIA INERMIS L.
*PUNICA GRANATUM L.
LupwIGIA ERECTA (L.) Hara Jussiaea erecta L.
OF THE GULF OF GUINEA
32. LYTHRACEAE
33. PUNICACEAE
34. ONAGRACEAE
LUDWIGIA HYSSOPIFOLIA (G. Don) Exell
Jussiaea linifolia Vahl
LUDWIGIA LEPTOCARPA (Nutt.) Hara
Jussiaea leptocarpa Nutt. LUDWIGIA OCTOVALVIS (Jacq.) Raven subsp. OCTOVALVIS Jussiaea suffruticosa L.
Ludwigia pubescens (L.) Hara FP: San Carlos, Mildbraed 678r.
Neither Ludwigia nor Jussiaea is recorded from FP
in 10.
35. HALORAGACEAE
*LAUREMBERGIA Cf. TETRANDRA (Schott) Kanitz
36. THYMELAEACEAE
DICRANOLEPIS DISTICHA Planchon
D. mannii Baillon
DICRANOLEPIS GRANDIFLORA Engl.
DICRANOLEPIS STENURA Gilg ex Engl. Not in 10, but probably = D. vestita.
DICRANOLEPIS THOMENSIS Engl. & Gilg
DICRANOLEPIS VESTITA Engl. PEDDIEA FISCHERI Engl. FP: Melville & Wrigley 632 (K.).
FP in fo.
PEDDIEA PARVIFLORA Hook. f. PEDDIEA THOMENSIS Exell
BoERHAVIA COCCINEA Miller B. diffusa sensu Exell pro parte et sensu Lains &
Silva.
BoERHAVIA COCCINEA var. viscosa (Lag. & Rodriguez)
Moscoso
37. NYCTAGINACEAE
B. viscosa Lag. & Rodriguez
Not recorded from FP in tro. BokERHAVIA DIFFUSA L.
*BoOERHAVIA PANICULATA Rich. *BOUGAINVILLEA GLABRA Choisy *BOUGAINVILLEA SPECTABILIS Willd.
*MIRABILIS JALAPA L.
*GREVILLEA ROBUSTA Cunn.
38. PROTEACEAE
Not recorded from
II
Io
2, 10
2, 10
16
L320
2
1-16,-25,-20 I
2
Ey: 2, 13 Lc? 20
13
P FP STA ST Pol Yb BES St FP FP BP FP ST FP FP FP SE PST A FP PSi 7 ST FP EPP st FP STA ST
343
i
344 ANGIOSPERMS OF THE ISLANDS
39. DILLENIACEAE
*DILLENIA INDICA L. £27 ST TETRACERA ALNIFOLIA Willd. I; 2,7, 10, 26 EPP A TETRACERA ERIANTHA (Oliver) Hutch. 2, 10 FP
40. PITTOSPORACEAE PITTOSPORUM MANNII Hook. f. subsp. MANNII 2, 10 PF,
41. BIXACEAE
-
*BIXA ORELLANA L. HO) 40, 42,350 Pee oe
25, 26 42. FLACOURTIACEAE (incl. SAMYDACEAE) CALONCOBA GLAUCA (Beauv.) Gilg 10 FP *CALONCOBA WELWITSCHII (Oliver) Gilg 2 FP CAMPTOSTYLUS MANNII (Oliver) Gilg - 10 FP Caloncoba mannii (Oliver) Gilg 2 CASEARIA BARTERI Mast. I, II, 25 Sit CASEARIA MANNII Mast. I P Ey FLACOURTIA FLAVESCENS Willd. II ST ? HOMALIUM AFRICANUM (Hook. f.) Benth. 2 FP ? Record probably based on a misidentification of H. letestui. HOMALIUM HENRIQUESII Gilg ex Engl. II Si E H. africanum sensu Exell I ; HOMALIUM LETESTUI Pellegrin 10 FP H. dolichophyllum Gilg ex Engl. yes | HOMALIUM SARCOPETALUM Pierre 10 FP ra LINDACKERIA DENTATA (Oliver) Gilg FP 7 FP: Bokoko, Burchardt. Notrecorded from FP in ro. i ONcoBA SPINosA Forssk. I, 2, 25, 26 FP ST f Not recorded from FP in to. OPHIOBOTRYS ZENKERI Gilg it, 25, 27 ST
43. PASSIFLORACEAE
ADENIA CISSAMPELOIDES (Planchon ex Benth.) Harms 2, 10, 26 FPP? SPz ADENIA CYNANCHIFOLIA (Benth.) Harms 2, 10 FP Ee it ADENIA LOBATA (Jacq.) Engl. 112540; 12 FPPSTA — BARTERIA FISTULOSA Mast. 2, 10 FP Fz BARTERIA NIGRITANA Hook. f. 10 FP Fz *PASSIFLORA EDULIS Sims Teel Ee25 SE . *PASSIFLORA FOETIDA L. 4,8, 10; 25,26 FP PSS *PASSIFLORA QUADRANGULARIS L. I P Se
44. CUCURBITACEAE
General reference: C. Jeffrey, ‘Key to the Cucurbitaceae of West Tropical Africa, with a guide to the localities of rare and little-known species’ in Journ. W. Afr. Sci. Assn. 9 : 79-97 (1965). STA
CAYAPONIA AFRICANA (Hook. f.) Exell t,.9; 20, IF
CITRULLUS LANATUS (Thunb.) Matsum. & Nakai EPPS z C. vulgaris Schrader 2 Colocynthis citrullus (L.) Kuntze I |
Not recorded from FP in Io.
OF THE GULF OF GUINEA
CoccINIA BARTERI (Hook. f.) Keay Physedva barteri (Hook. f.) Cogn. CUCUMEROPSIS EDULIS (Hook. f.) Cogn. Not recorded from FP in Io. CucUMIS METULIFER E. Meyer ex Naudin *CUCUMIS SATIVUS L. *CUCURBITA MAXIMA Duchesne DIPLOCYCLOsS PALMATUS (L.) C. Jeffrey Bryonopsis laciniosa Naudin pro parte LAGENARIA BREVIFLORA (Benth.) Roberty Adenopus breviflorus Benth. Not recorded from FP in 1o. LAGENARIA SICERARIA (Molina) Standley L. vulgaris Ser. *LUFFA ACUTANGULA Roxb. LUFFA AEGYPTIACA Miller L. cylindrica M. Roemer MomorDICaA CABRAE (Cogn.) C. Jeffrey Dimorphochlamys mannii Hook. f. MomorviIca CHARANTIA L.
MoMoRDICA CHARANTIA Var. ABBREVIATA Ser.
MomorpIca CISSOIDES Planchon ex Benth. MomorpDIca FOETIDA Schumacher
M. mannii Hook. f.
MoMmorDICA MULTIFLORA Hook. f. OREOSYCE AFRICANA Hook. f. PEPONIUM VOGELI!I (Hook. f.) Engl.
P. bracteatum (Cogn.) Cogn. RAPHIDIOCYSTIS MANNII Hook. f. RUTHALICIA LONGIPES (Hook. f.) C. Jeffrey
Physedra longipes Hook. f.
*SECHIUM EDULE Swartz TELFAIRIA OCCIDENTALIS Hook. f.
ZEHNERIA CAPILLACEA (Schumacher) C. Jeffrey
Melothria capillacea (Schumacher) Cogn. ZEHNERIA GILLETII (De Wild.) C. Jeffrey
Melothria capiliacea sensu Exell ZEHNERIA MINUTIFLORA (Cogn.) C. Jeffrey
FP: Wrigley 516 (K). Not recorded from FP in to.
ZEHNERIA SCABRA (L. f.) Sonder Melothria minutiflova sensu Exell M. fernandensis Hutch. & Dalziel
45. BEGONIACEAE
BEGONIA AMPLA Hook. f.
BEGONIA ANNOBONENSIS A.DC. BEGONIA BACCATA Hook. f. B. crateris Exell
I agree that B. crateris is no more than a form of B.
baccata. BEGONIA EMINII Warb. BEGONIA FUSIALATA Warb.
16
I, 10 16 T2025,
16, 25 2, 13
2
1 LOO 2
2, 10
13.2,°0 10,13; 25, 26
5 a Ras ©
ste)
10
.
~
Ne NNHNYH NNN on o°o
| °
2, 10
133595025; 25
EU 4.) 20) 2Y, 23
P5T,. 25
t. 13,25
10 23
FP FP ST? ST EP EP. S65 BPP ST A EP, SIA FP FP °StA FP FP PST A BP PST. A FP FP PP FP FP St | od EP oS ae FP FP ST FP PE SL EP PSIA PST A ST FP
345
E
346 ANGIOSPERMS OF THE ISLANDS
BEGONIA FURFURACEA Hook. f. BEGONIA GILGI Engl. BEGONIA GLADIIFOLIA Engl. BEGONIA LORANTHOIDES Hook. f. B. henriquestt C.DC. BEGONIA MACAMBRARENSIS Exell BEGONIA MANNII Hook. B. epiphytica Hook. f. B. excelsa Hook. f. BEGONIA MOLLERI (C.DC.) Warb. BEGONIA OXYLOBA Welw. ex Hook. f. BEGONIA POCULIFERA Hook. f. BEGONIA PRISMATOCARPA Hook. BEGONIA SESSILIFOLIA Hook. f. BEGONIA THOMEANA C.DC. BEGONIA sp. I BEGONIA sp. 2 BEGONIA sp. 3 BEGONIA Sp. 4 *BEGONIA sp. BEGONIA sp. cf. MACROPODA Gilg
46. CARICACEAE *CARICA PAPAYA L.
47. CACTACEAE
*OPUNTIA DILLENII (Ker-Gawl.) Haw. O. horrida Salm-Dyck
RHIPSALIS BACCIFERA (J. S. Miller) Stearn R. cassutha Gaertn.
48. OCHNACEAE
OcHNA MEMBRANACEA Oliver O. quintasii (Tieghem) Exell O. cf. gilgiana Engl. OURATEA AFFINIS (Hook. f.) Engl. ? OURATEA BRACHYBOTRYS Gilg FP: Bokoko, Mildbraed 6866. A doubtful record. Not in Io. OURATEA BRUNNEOPURPUREA Gilg Not in fo. OURATEA CALOPHYLLOIDES Hutch. & Dalziel OURATEA ELONGATA (Oliver) Engl. OuRATEA FLAVA (Schumacher) Hutch. & Dalziel ex Stapf O. reticulata sensu Guinea OURATEA MANNII (Oliver) Engl. OURATEA MICRANTHA (Hook. f.) Hutch. & Dalziel OURATEA MOLLERI (Tieghem) Exell Campylospermum vogelit var. molleri (Tieghem) Farron in Bull. Jard. Bot. Brux. 35 : 404 (1965). OURATEA NUTANS (Hiern) Exell OURATEA QUINTASII (Tieghem) Exell Rhabdophylilum arnoldianum var. quintasti (Tieghem) Farron, tom. cit. : 391 (1965).
®
Io Io Io
23
.
-
~
, 13, 23 Io
~
13, 23 10 Io Io 10
~ ~ ~
-
HH HHH DNDN NANNNARAADND SND
al ial N w
’
4 °
I, 16, 25
FP FP FP PST ST FP sT EP FP FP PP ST ¥ ST ST SF ST FP PSTA be FPPSTA STA FP FP FP P Si FP FP FP FP FP FP PST FP sT
iii
OF THE GULF OF GUINEA
49. MYRTACEAE
*EUCALYPTUS ROBUSTUS Smith *EUCALYPTUS Sp. *EUGENIA BRASILIENSIS Lam. *EUGENIA DOMBEYI (Sprengel) Skeels EUGENIA ELLIOTII Engl. & Brehmer EUGENIA FERNANDOPOANA Engl. & Brehmer EUGENIA OBANENSIS Baker f. *EUGENIA UNIFLORA L. *MyYRCIARIA CAULIFLORA O. Berg *PsIDIUM GUAJAVA L. *SYZYGIUM AROMATICUM (L.) Merr. & Perry SYZYGIUM GUINEENSE (Willd.) DC. SyYZYGIUM GUINEENSE var. LITTORALE Keay *SyzYGIUM JAMBOS (L.) Alston
Eugenia jambos L. *SYZYGIUM MALACCENSE (L.) Merr. & Perry
Eugenia malaccensis L. SYZYGIUM STAUDTII (Engl.) Mildbr.
ST: Monod 11929.
50. BARRINGTONIACEAE
NAPOLEONAEA MANNII Miers N. cuspidata sensu Guinea
51. MELASTOMATACEAE AMPHIBLEMMA CYMOSUM var. BOLIVARI Brenan & Guinea,
nom. provis.
A. Fernandes (in litt.) suggests that this may be A.
mildbraedii. AMPHIBLEMMA MILDBRAEDII Gilg ex Engl. CALVOA CONFERTIFOLIA Exell CALVOA CRASSINODA Hook. f. & Triana CALVOA GRANDIFOLIA Cogn. CALVOA HENRIQUESII Cogn. CaLVOA HIRSUTA Hook. f. CALVOA INTEGRIFOLIA Cogn. CALVOA MOLLERI Gilg CALVoA sinuaTa Hook. f. ex Triana CALVoA aff. TROCHAINII Jacques-Félix C. ignatii-bolivari Guinea, nom. provis. C. polychoeta Guinea, nom. invalid.
Referred to in the legend of the plate on p. 179 but not in the text, and presumably the same as C.
ignatii-bolivari. CALVOA UROPETALA Mildbr.
'DICELLANDRA BARTERI Hook. f.
DICELLANDRA SCANDENS Gilg ex Engl. DINOPHORA SPENNEROIDES Benth. DissoTis BARTERI Hook. f. ex Triana DissoTIs DECUMBENS (Beauv.) Triana DissoTiIs ROTUNDIFOLIA (Smith) Triana
2 13
13
27
II
2, 10
2°10
1,.2,'25
13
E522; 13,:25,120 27
2th 325 20, 27 Io
1,2
2 Io
10
10 24
13, 24
8, 13, 24
TY, 13; 16, 24 5, 10
24
II, 15, 24
24
ie)
~ ~ . ~ - - ~ . ~
.
aon He HN A AR Re Ae WD
ae at LO.
T5655 Jed; 24
347 Pr. ST ST ST ST FP E PP PEP St ST PPP SUA oh ER cob FP FPP STA EP EP 1 ST BE E FP EP. ST E oF E ST E nh Y E FP ST E ST E P E FP A. FP ie FP P E FP EP POE
348 ANGIOSPERMS OF THE ISLANDS
GUYONIA CILIATA Hook. f.
Afzeliella bolivari Brenan & Guinea, nom. provis.
Afzeliella sp. aff. ciliata MEDINILLA MANNII Hook. f. MEMECYLON FERNANDIANUM Gilg ex Engl. MEMECYLON MEMBRANIFOLIUM Hook. f. MEMECYLON MEMECYLOIDES (Benth.) Exell M. vogelit Naudin MEMECYLON aff. STRYCHNOIDES Baker MEMECYLON TESSMANNII Gilg ex Engl. Not in ro. MyYRIANTHEMUM MIRABILE Gilg TRISTEMMA CORONATUM Benth. TRISTEMMA HIRTUM Beauv.
TRISTEMMA INCOMPLETUM R. Br.
TRISTEMMA LITTORALE Benth. TRISTEMMA MILDBRAEDII Gilg ex Engl. TRISTEMMA OREOTHAMNUS Mildbr. TRISTEMMA RADICANS Gilg ex Engl. TRISTEMMA THOMENSIS Barros Ferreira
52. COMBRETACEAE
COMBRETUM BRACTEATUM (M. A. Lawson) Engl. & Diels COMBRETUM CONFERTUM (Benth.) M. A. Lawson COMBRETUM CUSPIDATUM Planchon ex Benth.
COMBRETUM INSULARE Engl. & Diels COMBRETUM PANICULATUM Vent.
? COMBRETUM PLATYPTERUM (Welw.) Hutch. & Dalziel
A doubtful record. COMBRETUM RACEMOSUM Beauv. CONOCARPUS ERECTUS L. Not recorded from FP in ro. LAGUNCULARIA RACEMOSA (L.) Gaertn. f. Not recorded from FP in to. *QUISQUALIS INDICA L. *TERMINALIA CATAPPA L.
53. RHIZOPHORACEAE
ANISOPHYLLEA CABOLE Henriq.
ANISOPHYLLEA SETOSA Mildbr., nom. nud. FP: Bokoko, Mildbraed 6836.
ANISOPHYLLEA Sp.
CASSIPOUREA ANNOBONENSIS Mildbr.
CASSIPOUREA GLOMERATA Alston
CASSIPOUREA GUMMIFLUA var. MANNII (Hook. f. ex
Oliver) J. Lewis C. mannii Hook. f. ex Oliver RHIZOPHORA HARRISONII Leechm. R. racemosa sensu Exell RHIZOPHORA MANGLE L.
This may be R. harrisonii. The genus is not recorded
from FP in Io.
, 10 , 10
5 5 2, 10 2 2 1, 10, 24, 25
Nn eH
2, 10
24
£355,100 83; 14, 15, 24
525.8, 10,:14, 13, 14, £5, 24
1; 2,-5.7; 10; 24
2,10, 24
152.55,010
2,10
24
2 2 2, 10 I I
‘0 21, 23,55 I i
tas] eH
FE i E EEE SE
FP PST
FPP FP $f
A FP
bd
ST
ale
FP Sf FP
FP ST | FPPSTA
PST
FP?
OF THE GULF OF GUINEA 349
54. HYPERICACEAE
HARUNGANA MADAGASCARIENSIS Lam. ex Poiret 10,40, 11,13, EP Sr 25 H. paniculata Pers. 2 HYPERICUM PEPLIDIFOLIUM A. Rich. Io FP HYPERICUM REVOLUTUM Vahl FP H. lanceolatum Lam. 2; 7 10 H. sp. 10 55. GUTTIFERAE | ALLANBLACKIA MONTICOLA Mildbr. ex Engl. ee Ce) FP GARCINIA KOLA Heckel 2 FP | Not recorded from FP in tro. | *GARCINIA MANGOSTANA L. 13 Sr | GARCINIA MANNII Oliver 2 FP Not recorded from FP in ro. GARCINIA POLYANTHA Oliver E22,°10 PPP? GARCINIA sp. FP FP: Bokoko, Burchardt. MAMMEA AFRICANA Sabine E, 10,15, 27 PES Pentadesma butyraceum sensu Exell et al. HAG Gf Baws ic omy 4 Mammea klaineana Pierre 2 ? PENTADESMA BUTYRACEUM Sabine |e fe See ‘Antidesma butyraceum’, p. 352. _ SYMPHONIA GLOBULIFERA L. f. 10;-26,-27 FP SF S. gabonensis (Vesque) Pierre I
FP: Bokoko, Mildbraed 6886, a8 5. Not recorded from FP in ro.
56. SCYTOPETALACEAE
RHAPTOPETALUM CORIACEUM Oliver PT FP SCYTOPETALUM KAMERUNIANUM Engl. 25,27 ST SCYTOPETALUM sp. II ST
This may be S. kamerunianum.
57. TILIACEAE
*BERRIA CORDIFOLIA (Willd.) Burret EE B. amomunilla Roxb. CORCHORUS AESTUANS L. I STA CORCHORUS CAPSULARIS L. 2 FP Not in 12. ~CorcHorus oLitorivs L. a PE ok: Not recorded from FP in 12. Duposcra MACROCARPA Bocquet 2 EP Not recorded from FP in 12. GLYPHAEA BREVIS (Sprengel) Monachino 12 Nd GREWIA CARPINIFOLIA Juss. #513 ST GREWIA FLORIBUNDA Mast. I ST GREWIA MALACOCARPA Mast. 12 ST GREWIA OLIGONEURA Sprague 2,42 Er: TRIUMFETTA CORDIFOLIA A. Rich. 7 dan i 4 FP TRIUMFETTA ERIOPHLEBIA Hook. f. 2,12 FP TRIUMFETTA RHOMBOIDEA Jacq. 12 PR SrA
Not recorded from FP in 12.
350 ANGIOSPERMS OF THE ISLANDS
58. STERCULIACEAE
CoLa ACUMINATA (Beauv.) Schott & Endl. I, ¢, 32, 23,455 FR PSE C. pseudoacuminata Engl. 2 COLA DIGITATA Mast. I PST COLA FICIFOLIA Mast. 202 FP CoLa NITIDA (Vent.) Schott & Endl. 12 FP C. acuminata var. latifolia K. Schum. 2 C. vera K. Schum. 2 CoLaA OBTUSA Engl. & Krause 2 FP Not in 12. ? COLA SPHAEROCARPA A. Chev. I ST? Said to be endemic but a very doubtful species. COLA VERTICILLATA (Thonn.) Stapf ex A. Chev. 2 FP Not recorded from FP in 12. LEPTONYCHIA ADOLFI-FRIEDERICI Engl. & Krause 2, 52 FP LEPTONYCHIA BATANGENSIS (C. H. Wright) Burret 12 FP LEPTONYCHIA DENSIVENIA Engl. & Krause 12 re L. urophylla sensu Guinea 2 LEPTONYCHIA FERNANDOPOANA Engl. & Krause ex Mildbr., nom. nud. 12 FP FP: Bokoko, Mildbraed 6823, 6862. LEPTONYCHIA LANCEOLATA Mast. 2, 12 FP LEPTONYCHIA PALLIDA K. Schum. 12 FP MELOCHIA CORCHORIFOLIA L. I st *PTERYGOTA ALATA (Roxb.) R.Br. 2 FP ? STERCULIA BEQUAERTII De Wild. 25 SI *STERCULIA FOETIDA L. 2 FP STERCULIA OBLONGA Mast. 2,12 FP
If the genus Evibroma Pierre be maintained, the name for this is E. oblonga (Mast.) Bodard
STERCULIA TRAGACANTHA Lindl. 1:2 FP PST Not recorded from FP in 12. *THEOBROMA CACAO L,. {, 2.12.13; 25... FP Psi WALTHERIA INDICA L. Be FP: Wrigley & Melville 704 (K). Not recorded from EP PAmur2:
59. BOMBACACEAE
ADANSONIA DIGITATA L. I, 9, 25, 26 FP PSTA A. digitata var. congolensis A. Chev. Tks Not recorded from FP in 12. BoOMBAX BUONOPOZENSE Beauv. 2 FP Not recorded from FP in 12. *BOMBAX KIMUENZAE De Wild. & Dur. 2 Ba CEIBA PENTANDRA (L.) Gaertn. LS, ka: SS FP PSTA
Not recorded from FP in 12.
60. MALVACEAE
*A BELMOSCHUS ESCULENTUS (L.) Moench 22 PSF Hibiscus esculentus L. ‘24
ABELMOSCHUS MOSCHATUS Medik. 2, 22 FP Sk Hibiscus abelmoschus L. ¥
ABUTILON GRANDIFLORUM G. Don I, 22, 25, 26 ST
OF THE GULF OF GUINEA
* A BUTILON GRANDIFOLIUM (Willd.) Sweet A. mollissimum (Cav.) Sweet
See R. Fernandes in Anu. Soc. Brot. 25 : 29 (1959).
*A BUTILON VENOSUM Lemaire *GOSSYPIUM BARBADENSE L. *var. BRASILIENSE (Macfad.) J. B. Hutch. G. brasiliense Macfad. G. avboreum sensu Guinea *GOSSYPIUM HIRSUTUM L. *HIBISCUS ACETOSELLA Welw. ex Hiern HIBISCUS DONIANUS D. Dietr. *HIBISCUS MUTABILIS L. HIBISCUS PHYSALOIDES Guillemin & Perrottet *HIBISCUS ROSA-SINENSIS L, *HIBISCUS SABDARIFFA L,. *HIBISCUS SCHIZOPETALUS (Mast.) Hook. f. FP: Guinea 97. HIBISCUS SURATTENSIS L. Not recorded from FP in 12. *HIBISCUS SYRIACUS L. HIBISCUS TILIACEUS L. *MALVASTRUM COROMANDELIANUM (L.) Garcke PAVONIA URENS Cav. var. URENS
PAVONIA URENS var. GLABRESCENS (Ulbr.) Brenan
P. schimperana sensu Guinea Spa acuta Burm. f.
SIDA ACUTA var. CARPINIFOLIA K. Schum. SIDA ALBA L.,
FP: Moka, Tessmann 2817. Not recorded from FP
in 12. SIDA CORDIFOLIA L.
Not recorded from FP in 12. SIDA RHOMBIFOLIA L. SIDA RIGIDA (D. Don) D. Dietr. SIDA STIPULATA Cav. SIDA URENS L. SIDA VERONICIFOLIA Lam. URENA LOBATA L.
WISSADULA ROSTRATA (Schumacher) Hook. f.
W. amplissima var. rostrata (Schumacher) R. E.
Fries W. heterosperma Hochst. ex Mast.
61. MALPIGHIACEAE
ACRIDOCARPUS LONGIFOLIUsS (G. Don) Hook. f. FLABELLARIA PANICULATA Cav. HETEROPTERIS LEONA (Cav.) Exell
62. HUMIRIACEAE
SACOGLOTTIS GABONENSIS (Baillon) Urban
I, 22
I, 22 i Ow22
I2
Z
2.
Ts .22
PL, 15;.22,-25 Te22
£522
TON Pr 322 122
2. ray 22
I, 2, 22
-
1 Oe ir Oils be 4
1s, 22
rz
52
2
TiO 5Ore lal 2) Pt 22,025
2
T 22
E230; TT 39
1, 125 13,20; 22
I, 22
12
I, 22
1, 22
1, Oo; 4b 12413: 22, 25
15,22, 25
t, 6,48; 12
ST
Si: PST-A FP
SE Pst i PST SrA PST BP St 1 op Se
EP ESE
BY
PPa so. Si
FP
FP
PR PST A
FP
FP A
BE SST A
FPPSTIA
FP
Pe stA
BPP STA
PEE SE FP Pst
FP
351
352 ANGIOSPERMS OF THE ISLANDS
63. ERYTHROXYLACEAE *ERYTHROXYLUM Coca Lam. I ERYTHROXYLUM EMARGINATUM Thonn. 9.21; 27 64. LINACEAE (HUGONIACEAE)
HUGONIA PLATYSEPALA Welw. ex Oliver Ti2 HUGONIA SPICATA Oliver var. SPICATA 252 The type variety is endemic.
65. EUPHORBIACEAE
ACALYPHA ANNOBONAE Pax & Hoffm. Taos kO ACALYPHA CILIATA Forssk. I ACALYPHA MANNIANA Miill. Arg. 12 ACALYPHA ORNATA Hochst. ex A. Rich. 2 Not recorded from FP in 12.
ACALYPHA RACEMOSA Wall. ex Baillon 12
A. paniculata Miq. 2 ALCHORNEA CORDIFOLIA (Schumacher) Miill. Arg. D251, 25.26 ALCHORNEA FLORIBUNDA Mill. Arg. 2,42 ALCHORNEA HIRTELLA Benth. I ALCHORNEA LAXIFLORA (Benth.) Pax & Hoffm.
Lepidoturus laxiflorus Benth. I *ALEURITES CORDATA (Thunb.) R. Br. ex Steudel 13 ANTHOSTEMA AUBRYANUM Baillon Teel tok?
‘Antidesma butyraceum’
Possibly Pentadesma butyraceum was intended. If so,
it is a new record of that species from FP. ANTIDESMA LACINIATUM Mill. Arg. var. LACINIATUM 2a lke ANTIDESMA MEMBRANACEUM Miill. Arg. I ANTIDESMA OBLONGA (Hutch.) Keay
FP: Wrigley & Melville 633 (K). Not recorded from
sae as *BREYNIA NIVOSA (W. G. Smith) Small 29 Phyllanthus nivosus W. G. Smith 13 BRIDELIA STENOCARPA Miill. Arg. 71%, 25 B. micrantha sensu Exell 1323.9 Not recorded from FP in 12. CAPERONIA LATIFOLIA Pax 1,22: 12 CAVACOA QUINTASII (Pax & Hoffm.) J. Léonard 15 Grossera quintastt Pax & Hoffm. I CLEISTANTHUS LIBERICUS N. E. Brown 1, 16,. 27 C. polystachyus sensu Hutch. 41, 32,25, 27 *CODIAEUM VARIEGATUM (L.) Blume I, 16 *CROTON DRACONOPSIS Miill. Arg. I CROTON LOBATUS L. 12 CROTON STELLULIFER Hutch. I CROTONOGYNE MANNIANA Miill. Arg. subsp. MANNIANA Par Pee DISCOCLAOXYLON HEXANDRUM (Miill. Arg.) Pax & Hoffm. 2, 16 Claoxylon hexandrum Mill. Arg. 12 DISCOCLAOXYLON OCCIDENTALE (Miill. Arg.) Pax & Hoffm. 1/2084, 20 DISCOCLAOXYLON PEDICELLARE (Miill. Arg.) Pax & Hoffm. 2, 16 Claoxylon pedicellave Mill. Arg. 12
ST STA cP F FP A ST re FP FP BE Pst FP P ST ST | ab FP FP STA FP ST FPPSTA ST STA ST STA ST FP e FP 1 od ST FP
Eg EE me me =
OF THE GULF OF GUINEA
DISCOCLAOXYLON PUBESCENS (Pax & Hoffm.) Exell D. occidentale var. pubescens Pax & Hoftm. For a general discussion of Discoclaoxylon see 16. DISCOGLYPREMNA CALONEURA (Pax) Prain DRYPETES FERNANDOPOANA Brenan D. glomerata (Mill. Arg.) Hutch. DRYPETES GLABRA (Pax) Hutch. DRYPETES HENRIQUESII (Pax) Hutch. DRYPETES LACINIATA (Pax) Hutch. DRYPETES OCCIDENTALIS (Miill. Arg.) Hutch. DRYPETES PRINCIPUM (Miill. Arg.) Hutch. ELAEOPHORBIA DRUPIFERA (Thonn.) Stapf ERYTHROCOCCA ANOMALA (Juss. ex Poiret) Prain ERYTHROCOCCA COLUMNARIS (Miill. Arg.) Prain ERYTHROCOCCA MANNII (Hook. f.) Prain Athroandra mannii (Hook. f.) Pax & Hoffm. ERYTHROCOCCA MOLLERI (Pax) Prain ERYTHROCOCCA PALLIDIFOLIA (Pax & Hoffm.) Keay Athroandra pallidifolia Pax & Hoffm. EUPHORBIA CERVICORNU Baillon EvUPHORBIA GLAUCOPHYLLA Poiret Not recorded from FP in 12. EvUPHORBIA HIRTA L.
| EvuprHorBIA HYSSOPIFOLIA L.
E. hypericifolia sensu Exell *EUPHORBIA PEPLUS L. EuPHORBIA PROSTRATA Aiton
*EUPHORBIA PULCHERRIMA Willd. ex Klotzsch EUPHORBIA SCHIMPERANA G. H. A. Scheele E. ampla Hook. f.
FP: Moka, Evell845(BM). Notrecorded from FP in 12.
*EUPHORBIA SERPENS Kunth
EvuPHORBIA THYMIFOLIA L.
*EUPHORBIA TIRUCALLI L. E. rhipsaloides Welw. ?
EXCOECARIA GUINEENSIS (Benth.) Miill. Arg. Sapium guineense (Benth.) Kuntze
See J. Léonard in Bull. Soc. Bot. Brux. 29 : 138 (1959).
GROSSERA ELONGATA Hutch.
GROSSERA sp.
*HEVEA BRASILIENSIS (Kunth) Miill. Arg.
*HEVEA GUIANENSIS Aubl.
*JATROPHA CURCAS L.
*JATROPHA MULTIFIDA L.
KLAINEANTHUS GABONIAE Pierre ex Prain
MACARANGA MONANDRA Miill. Arg.
MACARANGA OCCIDENTALIS (Miill. Arg.) Miill. Arg.
MACARANGA SPINOSA Miill. Arg.
MAESOBOTRYA DUSENII (Pax) Hutch.
MAESOBOTRYA GLABRATA (Hutch.) Exell Thecacoris glabrata Hutch. Maesobotrya ‘sp. nov.’
16 I
1, 2; 32,32; 25
¥,.0,.11, 32,25, 26
1 tee)
I
I
T7911, k25 13; 25, 26
16
TO. 12:52
To 10, LOi25;-20 Tr 12
II, 25
212
2052
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15
I
I
EP
ST FP
ba cel sate haw §
baatcy &
SE FRPP St A
A PP
353
H Be
hit wt
354 ANGIOSPERMS OF THE ISLANDS
MAESOBOTRYA Sp.
Thecacoris stenopetala sensu Exell pro parte MALLOTUS SUBULATUS Miill. Arg. *MANIHOT ESCULENTA Crantz *MANIHOT GLAzIovil Miill. Arg. MANNIOPHYTON FULVUM Miill. Arg.
M. africanum Mill. Arg. MAREYA MICRANTHA (Benth.) Miill. Arg. subsp.
MICRANTHA
M. spicata Baillon MARGARITARIA DISCOIDEA (Baillon) Webster
Phyllanthus discoideus (Baillon) Mill. Arg. MICRODESMIS PUBERULA Hook. f. ex Planchon NEOBOUTONIA MANNII Benth. PHYLLANTHUS AMARUS Schumacher PHYLLANTHUS CAPILLARIS Schumacher PHYLLANTHUS MUELLERANUS (Kuntze) Exell PHYLLANTHUS NIGERICUS Brenan PHYLLANTHUS ODONTADENIUS Miill. Arg.
P. niruri sensu Exell PHYLLANTHUS PENTANDRUS Schumacher PHYLLANTHUS PHYSOCARPUS Mill. Arg. PHYLLANTHUS RETICULATUS Poiret PHYLLANTHUS TESSMANNII Hutch. PROTOMEGABARIA MACROPHYLLA Hutch.
PSEUDAGROSTISTACHYS AFRICANA (Mill. Arg.) Pax &
Hoffm. subsp. AFRICANA PYCNOCOMA MACROPHYLLA Benth. var. MACROPHYLLA
RICINODENDRON HEUDELOTII subsp. AFRICANUM (Mill.
Arg.) J. Léonard R. africanum Mill. Arg.
See J. Léonard in Bull. Soc. Bot. Brux. 31 : 398 (1961).
*RICINUS COMMUNIS L. SAPIUM ELLIPTICUM (Hochst.) Pax SPONDIANTHUS PREUSSII Engl. var. PREUSSII
TETRACARPIDIUM CONOPHORUM (Miill. Arg.) Hutch. &
Dalziel
TETRORCHIDIUM DIDYMOSTEMON (Baillon) Pax & Hoffm.
THECACORIS ANNOBONAE Pax & Hoffm. THECACORIS MANNIANA (Miill. Arg. )Miill. Arg. THECACORIS MEMBRANACEA Pax THECACORIS STENOPETALA (Miill. Arg.) Miill. Arg. TRAGIA BENTHAMII Baker TRAGIA TENUIFOLIA Benth. *TRAGIA VOLUBILIS L. UAPACA GUINEENSIS Mill. Arg.
Uapaca sp. UAPACA STAUDTII Pax
66. ROSACEAE
ALCHEMILLA CRYPTANTHA Steudel ex A. Rich. A. tenuicaulis Hook. f.
*CYDONIA OBLONGA Miller Pyrus cydonia L.
14, 15 I 2,52 re I
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FP STF
ST
OF THE GULF OF GUINEA
*ERIOBOTRYA JAPONICA (Thunb.) Lindl.
*MALUS PUMILA Miller A doubtful record.
*PRUNUS CERASUS L. A doubtful record.
*PRUNUS DOMESTICA L.
*PRUNUS DULCIS (Miller) D. A. Webb
P. amygdalus Batsch
*PRUNUS PERSICA (L.) Batsch
PYGEUM AFRICANUM Hook. f. If transferred to Prunus this becomes P. africana (Hook. f.) Kalkman
*ROSA CHINENSIS Jacq.
*ROSA LAEVIGATA Michaux
*ROSA MULTIFLORA Thunb.
*RUBUS DISCOLOR Weihe & Nees
“Rubus extensus Schube ex Engl.’ Index Kew., Suppl. 1: 370 (1906), cited as ‘in Abh. Preus. Akad. Wiss. 1891 (1892) 212.—Ins. Fernando Po’. A manifest error for Ranunculus extensus Schube ex Engl., tom. cit. : 218.
RUBUS PINNATUS var. AFROTROPICUS (Engl.) Gustafsson
Rubus afrotropicus (Engl.) Guinea *RUBUS ROSIFOLIUS Smith RUBUs sp.
67. CHRYSOBALANACEAE
ACIOA MANNII (Oliver) Engl. CHRYSOBALANUS ELLIPTICUS Solander ex Sabine ? CHRYSOBALANUS ORBICULARIS Schumacher Probably C. ellipticus. PARINARI EXCELSA Sabine A doubtful record. PARINARI MACROPHYLLA Sabine A doubtful record. PARINARI MOBOLA Oliver A doubtful record.
68. DICHAPETALACEAE
DICHAPETALUM AFFINE Planchon ex Benth.
_ DICHAPETALUM BOCAGEANUM (Henriq.) Engl. DICHAPETALUM FLORIBUNDUM (Planchon) Engl. DICHAPETALUM KAMERUNENSE Engl.
FP: San Carlos, Mildbraed 6793. Not in 12. DICHAPETALUM OBLONGUM (Hook. f.) Engl. DIcHAPETALUM SUBCORDATUM (Hook. f.) Engl. DICHAPETALUM TOMENTOSUM Engl.
D. acutifolium Engl.
FP: Bokoko, Burchardt. TAPURA AFRICANA Oliver
Not recorded from FP in 12.
69. LEGUMINOSAE ABRUS PRECATORIUS L. *ACACIA FARNESIANA (L.) Willd. ACACIA KAMERUNENSIS Gandoger A. pennata sensu auctt., pro parte
D355 I
1; :9;:33242 he 228
15
I, 25
355 ST ST? ST? ST? ST ST FP ST ST ST ST FP FP STA FP P ST ST FP FP FP ? ST? ST? ST? FP E Shee FP FP FP FP E FP FP FP STA FP ST ST
356 ANGIOSPERMS OF THE ISLANDS
ACACIA PENTAGONA (Schumacher) Hook. f. A. pennata sensu auctt., pro parte * A DENANTHERA PAVONINA L. ADENOCARPUS MANNII (Hook. f.) Hook. f. AESCHYNOMENE INDICA L. AIRYANTHA SCHWEINFURTHII subsp. CONFUSA (Hutch. & Dalziel) Brummitt Baphia confusa Hutch. & Dalziel Baphiastrum confusum (Hutch. & Dalziel) Pellegrin Baphia spathacea Hook. f. See Brummitt in Kew. Bull. 22 : 381 (1968). * ALBIZIA FALCATA (L.) Backer *ALBIZIA LEBBECK (L.) Benth. * ALBIZIA MOLUCCANA Miq. *ALBIZIA PROCERA (Roxb.) Benth. ALBIzIA ZYGIA (DC.) J. F. Macbride A. brownei (Walp.) Oliver Not recorded from FP in 12. ALYSICARPUS VAGINALIS (L.) DC. ANTHONOTHA MACROPHYLLA Beauv. Macrolobium palisotii Benth. * ARACHIS HYPOGAEA L. BAIKIAEA INSIGNIS Benth. subsp. INSIGNIS BAPHIA CAPPARIDIFOLIA Subsp. POLYGALACEA (Hook. f.) Brummitt B. polygalacea (Hook. f.) Baker BAPHIA NITIDA Lodd. ST: Espirito Santo 4422. *BAUHINIA PURPUREA L. *BAUHINIA TENUIFLORA Watt ex C. B. Clarke *BAUHINIA TOMENTOSA L, BERLINIA BRACTEOSA Benth. CAESALPINIA BONDUC (L.) Roxb. C. crista sensu Guinea *CAESALPINIA PULCHERRIMA (L.) Swartz *CAJANUS CAJAN (L.) Millsp. C. indicus Sprengel *CALOPOGONIUM MUCUNOIDES Desvaux CANAVALIA ENSIFORMIS (L.) DC. Not recorded from FP in 12. CANAVALIA MARITIMA (Aubl.) Urban C. rosea (Swartz) DC. C. obtusifolia DC. pro parte Not recorded from FP in 12. *CASSIA ALATA L. CASSIA KIRKII Oliver *CASSIA LAEVIGATA Willd. CASSIA MANNII Oliver CASSIA MIMOSOIDES L. FP: Moka, Tessmann 2868. Not recorded from FP in 12. CASSIA OBTUSIFOLIA L. C. tora sensu auctt. See Brenan in Kew Bull. 13 : 248 (1958).
tt,.13 2, 11,.%2 2 II
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P FP ST FP PSF FP of FP ST FP ST FP STA FP EP SLA FP FP BY St ST FP ST FP FP STA FP PSEA FPPSTA sT FP ST FP STA EP SE FP FP STA PSI PP St FP STA
OF THE GULF OF GUINEA
*CASSIA OCCIDENTALIS L.
CASSIA PODOCARPA Guillemin & Perrottet *CASSIA SIAMEA Lam. CASSIA SIEBERANA DC. *CASSIA SOPHERA L. *CASSIA SPECTABILIS DC. *CENTROSEMA PLUMIERI (Turpin ex Pers.) Benth. *CENTROSEMA PUBESCENS Benth. *CERCIS SILIQUASTRUM L, *CLITOREA TERNATEA L., COLOPHOSPERMUM MOPANE (Benth.) Kirk ex J. Léonard Copatfera mopane Benth. Helder Lains e Silva (13) records Copaifera mopane (‘pau ferro’) as one of the important timber trees of S. Tomé, but no one else to my knowledge has ever recorded it from the island and I have seen no specimens. CROTALARIA DONIANA Baker ST: Espirito Santo 66. CROTALARIA GOREENSIS Guillemin & Perrottet *CROTALARIA LANCEOLATA E. Meyer Not recorded from FP in 12. If correctly named, not indigenous. CROTALARIA MUCRONATA Desvaux CROTALARIA OCHROLEUCA G. Don CROTALARIA RETUSA L. P: Rozeiva 452. Not recorded from FP in 12. *CROTALARIA ZANZIBARICA Benth. C. thomensis Baker f. CRUDIA KLAINEI Pierre ex De Wild. C. senegalensis sensu Guinea CYNOMETRA MANNII Oliver
DALBERGIA ECASTAPHYLLUM (L.) Taub. *DALBERGIA MONETARIA L, DANIELLIA OBLONGA Oliver
D. thurifera sensu Exell DANIELLIA OGEA (Harms) Rolfe ex Holland *DELONIX REGIA (Bojer ex Hook.) Raf.
Poinciana regia Bojer ex Hook. *DESMANTHUS VIRGATUS (L.) Willd. DESMODIUM ADSCENDENS (Swartz) DC. var. ADSCENDENS DESMODIUM ADSCENDENS var. ROBUSTUM Schubert DESMODIUM CANvUM (J. F. Gmelin) Schinz & Thell.
D. incanum DC.
DESMODIUM GANGETICUM (L.) DC. P: Gossweiler s.n., Rose 487, Rozeiva 4220.
DESMODIUM PROCUMBENS (Miller) A. S. Hitchc.
D. ospriostreblum Steudel ex Chiov. DESMODIUM RAMOSISSIMUM G. Don
D. mauritianum sensu Exell DESMODIUM REPANDUM (Vahl) DC.
D. scalpe DC.
3
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PEP ST A. EP. of ST ST ? EPP Sr ST ST ST ST 1 Ap cote b a Bi ST BE FP | eh ST PEP or ST iil ad ST FPP SIA FP EY pout aS Se EP Se Fr St FE St. A Pr PEP Sin 4 A A PEP St A BP OSE
357
358 ANGIOSPERMS OF THE ISLANDS
DESMODIUM TORTUOSUM (Swartz) DC. ST: Espirito Santo 4203, Rozeiva 5072. DESMODIUM TRIFLORUM (L.) DC. ? D. trifolium sensu Guinea DESMODIUM VELUTINUM (Willd.) DC. D. lasiocarpum (Beauv.) DC. D. latifolium DC. Not recorded from FP in 12. DIALIUM GUINEENSE Willd. Not recorded from FP in 12. DIOCLEA REFLEXA Hook. f. DISTEMONANTHUS BENTHAMIANUS Baillon Not recorded from FP in 12. DREPANOCARPUS LUNATUS (L. f.) G. F. W. Meyer ST: Rozeiva 4403. ENTADA AFRICANA Guillemin & Perrottet Not recorded from FP in 12. ENTADA GIGAS (L.) Fawcett & Rendle E. scandens sensu Guinea ENTADA MANNII (Oliver) Tisserant Piptadenia mannii Oliver
ERIOSEMA GLOMERATUM (Guillemin & Perrottet) Hook. f. ? ERIOSEMA PARVIFLORUM E. Meyer subsp. PARVIFLORUM
A doubtful record. ? ERYTHRINA CAFFRA Thunb. * ERYTHRINA DROOGMANSIANA De Wild. & Dur. Probably introduced. ERYTHRINA GILLETII De Wild. Not in 12. *ERYTHRINA INDICA Lam. *ERYTHRINA POEPPIGIANA (Walp.) O. F. Cook ? ERYTHRINA SENEGALENSIS DC. This may be E. vogelit. *ERYTHRINA UMBROSA Kunth *ERYTHRINA VELUTINA Willd. ERYTHRINA VOGELII Hook. f. ERYTHROPHLOEUM GUINEENSE G. Don Not recorded from FP in 12. GLYCINE WIGHTII (Wight & Arn.) Verdc. G. javanica sensu Exell GRIFFONIA PHYSOCARPA Baillon *HAEMATOXYLUM CAMPECHIANUM L. INDIGOFERA ASTRAGALINA DC. ST: Espirito Santo 4214. INDIGOFERA COLUTEA (Burm. f.) Merr. var. COLUTEA ST: Espirito Santo 4228. INDIGOFERA HIRSUTA L. INDIGOFERA SPICATA Forssk. I. hendecaphylia Jacq. INDIGOFERA SUFFRUTICOSA Miller INDIGOFERA TINCTORIA L. INDIGOFERA TRITA var. SUBULATA (Vahl ex Poiret) Ali ST: Rozeiva 4416. *INGA EDULIS Mart.
13
2
BE ~ I, 9, 13
2
t Apaly fgen OF EA tera
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13, 14, 15 15 Z
I $e
1,9
2Y, 25 ¥,.0; 24,25
15, 25, 27
ST
EP ?. PST
FPPSTA
FP PST
FPP St
PST
FP
STA
PSTA
ee
OF THE GULF OF GUINEA
*INGA sp. LABLAB PURPUREUS (L.) Sweet Dolichos lablab L.
Not recorded from FP in 12.
LEPTODERRIS BRACHYPTERA (Benth.) Dunn
L. africana (Baker) Harms *LEUCAENA LEUCOCEPHALA (Lam.) De Wit L. glauca sensu Benth.
LEUCOMPHALUS CAPPARIDEUS Benth. ex Planchon LONCHOCARPUS CYANESCENS (Schumacher) Benth. LONCHOCARPUS GRIFFONIANUS (Baillon) Dunn
? Millettia thonningii sensu Guinea ST: Espirito Santo 4553. LONCHOCARPUS SERICEUS (Poiret) Kunth ex DC. MILLETTIA BARTERI (Benth.) Dunn MILLETTIA MACROPHYLLA Benth. MILLETTIA MANNII Baker Not recorded from FP in 12. MILLETTIA MILDBRAEDII Harms
FP: San Carlos, Mildbraed 6975. Not in 12. MILLETTIA SANAGANA Harms MILLETTIA THONNINGII (Schumacher) Baker *MIMOSA POLYDACTYLA Humb. & Bonpl. ex Willd. *MIMOSA PUDICA L.
M. pigra sensu Espirito Santo This may be M. polydactyla. MUCUNA FLAGELLIPES T. Vogel ex Hook. f. & Benth. MUCUNA PRURIENS (L.) DC. M. utilis sensu Guinea Not recorded from FP in 12. MUCUNA SLOANEI Fawcett & Rendle M. urens sensu Exell ORMOCARPUM SENNOIDES subsp. HISPIDUM Brenan & Léonard O. cochinchinense (Lour.) Merr. See Brenan & Léonard in Bull. Jard. Bot. Brux. 24 : 104 (1954)- _ ORMOCARPUM VERRUCOSUM Beauv. *ORMOSIA DASYCARPA Jackson OSTRYOCARPUS RIPARIUS Hook. f. PARKIA CLAPPERTONIA Keay ? P. africana sensu Guinea Not recorded from FP in 12. PARKIA OLIVERI J. F. Macbride *PELTOPHORUM PTEROCARPUM (DC.) Backer ex K. Heyne P. ferrugineum (Decaisne) Benth. PENTACLETHRA MACROPHYLLA Benth.
(Willd.)
PHASEOLUS ADENANTHUS G. F. W. Meyer PHASEOLUS LUNATUS L. *PHASEOLUS VULGARIS L. PHYSOSTIGMA VENENOSUM Balf. ? PILIOSTIGMA RETICULATUM (DC.) Hochst. Bauhinia reticulata DC. Not recorded from FP in 12.
13
oat
Tt. 2, 12, 13 2. -¥2
2,12
2, 9512
2
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25
ST FP FP PPP SL FP FP FP ST PPPS ST FP FP FP FP | ol cas ib dred & FP 50 BP 1 Sy Se i PP Sr A ST PPP Sr ST FP iS STA FP FP PSEA ST dB saat Ree ST FP EP? SP ¢
359
E?
360 ANGIOSPERMS OF THE ISLANDS
PIPTADENIASTRUM AFRICANUM (Hook. f.) Brenan
Piptadenia africana Hook. f.
FP: Bokoko, Burchardt. Notrecorded from FP in 12.
PSOPHOCARPUS SCANDENS (Endl.) Verdc. *PTEROCARPUS INDICus Willd. ST: Espirito Santo 4388. PTEROCARPUS MILDBRAEDII Harms *PUERARIA JAVANICA Benth. RHYNCHOSIA DENSIFLORA (Roth) DC. R. debilis G. Don RHYNCHOSIA MANNII Baker RHYNCHOSIA MINIMA (L.) DC. R. memnonia sensu Exell RHYNCHOSIA PREUSSII (Harms) Taub. ex Harms R. cyanosperma Benth. ex Baker pro parte RHYNCHOSIA PYCNOSTACHYA (DC.) Meikle SAMANEA SAMAN (Jacq.) Merr. Pithecellobium samana (Jacq.) Benth. SCORODOPHLOEUS ZENKERI Harms Not in 12. SESBANIA SERICEA (Willd.) Link S. pubescens DC. SOPHORA TOMENTOSA L. S. occidentalis L. *TAMARINDUS INDICA L. *TEPHROSIA CANDIDA (Roxb.) DC. TEPHROSIA NOCTIFLORA Bojer ex Baker ST.: Esptrito Santo 4324, Rozeiva 775, 1231. TEPHROSIA PLATYCARPA Guillemin & Perrottet T. flexuosa G. Don
TEPHROSIA PURPUREA subsp. LEPTOSTACHYA var.
PUBESCENS Baker TEPHROSIA UNIFLORA Pers.
ST: Espirito Santo 4636. TEPHROSIA VOGELII Hook. f.
TERAMNUS LABIALIS (L. f.) Sprengel TETRAPLEURA TETRAPTERA (Schumacher) Taub.
TRIFOLIUM RUEPPELLIANUM var. PREUSSII (Taub. ex
Baker f.) J. B. Gillett T. preussii Taub. ex Baker f. TRIFOLIUM SIMENSE Fresen. T. subrotundum Steudel ex Hochst. TRIFOLIUM USAMBARENSE Taub. T. mannii Guinea, nom. nud. URaRIA PICTA (Jacq.) DC. *VICIA FABA L. VIGNA ALBA (G. Don) Planchon ex Baker f.
VIGNA GRACILIS (Guillemin & Perrottet) Hook. f.
V. occidentalis Baker f. VIGNA MULTIFLORA Hook. f. VIGNA OBLONGA Benth.
V. marina sensu Exell
i, 6, 11, 42
12
122) 0k 25 20
Ti.2s
Rey eb ot pero teed,
20
TO;-51 405 1,093,213
FP ST ST FP ST st FP ST A FP FP FP ST FP STA ST FP PST A ST Sr ST ST ST FPP ST PST A ST se PE: FP STA ST st Pr FP FPS Sk
OF THE GULF OF GUINEA 361
VIGNA RACEMOSA (G. Don) Hutch. & Dalziel TO, 121, 13 Best:
P: Rozeiva 3932, 3991. VIGNA UNGUICULATA (L.) Walp. 16 EP. A
V. sinensis Endl. ex Hassk. FP: Mann 415. Not recorded from FP in 12. See Baker in Oliver, Fl. Trop. Afr. 2: 205 (1871).
VIGNA VEXILLATA (L.) A. Rich. 12 FP V. thonningii Hook. f. 2 ZENKERELLA CITRINA Taub. 12 FP Cynometra leptoclada Harms 2 ZENKERELLA PAUCIFLORA Harms FP Cynometra pauciflova (Harms) Harms 2 Not in 12.
70. BUXACEAE *BuXUS SEMPERVIRENS L. I ST?
71. MYRICACEAE MyRICA ARBOREA Hutch. 12 1g e
72. ULMACEAE
CELTIS GOMPHOPHYLLA Baker Si
C. duvrandii Engl. Fale. Es, 25; 27
CELTIS INTEGRIFOLIA Lam. 13 ST?
CELTIS MILDBRAEDII Engl. ¥2,25,27 ST C. soyauxit sensu G. Taylor E13
CELTIS PRANTLII Priemer ex Engl. EO. 41, 27 PSA C. insularis Rendle 2
This is considered by Letouzey (Fl. Cameroun 8 : 26 (1968)) to be C. philippensis Blanco.
CELTIS WRIGHTII Planchon FP C. brownii Rendle 12 C. prantlii sensu Guinea C. rendleana G. Taylor Oe
See Polhill in Kew Bull. 19: 141 (1964). Letouzey (Joc. cit.) considers that this also is C. philippensis
Blanco. CHAETACME ARISTATA E. Meyer ex Planchon T-12 A TREMA ORIENTALE (L.) Blume FP PSTA T. guineense (Schumacher) Ficalho se ike Fa aah 73. MORACEAE ANTIARIS AFRICANA Engl. 2 we Not recorded from FP in 12. ANTIARIS sp. FP FP: Bokoko, Mildbraed 6880. *ARTOCARPUS ALTILIS (S. Parkinson) Fosberg EPPS A A. communis J. R. & G. Forster a3 510;.25 A. incisus (Thunb.) L. f. 2 *ARTOCARPUS HETEROPHYLLUS Lam. 16, 25 FPPSI A A. integer sensu G. Taylor £313
A. integrifolius sensu Guinea 2
362 ANGIOSPERMS OF THE ISLANDS
BosQuEIA ANGOLENSIS Ficalho *CASTILLOA ELASTICA Cerv.
*CECROPIA PELTATA L.
CHLOROPHORA EXCELSA (Welw.) Benth.
DORSTENIA BARTERI Bur. DORSTENIA ELLIPTICA Bur. DORSTENIA PROREPENS Engl. Ficus ANNOBONENSIS Mildbr. & Hutch. Not in 12. Recorded from FP in 1, fide G. Taylor. FICUS CAMPTONEURA Mildbr. FICUS CAMPTONEUROIDES Hutch. Ficus CAPENSIS Thunb. *FICUS CARICA L. Ficus CHLAMYDOCARPA Mildbr. & Burret F. clavencensis Mildbr. & Hutch. *FICUS ELASTICA Roxb. FicuS ERIOBOTRYOIDES Kunth & Bouché _ var. ERIOBOTRYOIDES Ficus EXASPERATA Vahl FICUS FERNANDESIANA Hutch. FICUS GLOBICARPA Warb. ex Guinea, nom. nud. I cannot trace that this name, listed by Guinea in 2, has been validly published. FICUS KAMERUNENSIS Warb. ex Mildbr. & Burret FICUS MACROSPERMA Warb. ex Mildbr. & Burret FP: Melville & Wrigley 622 (K). Not recorded from EP in 72. FICUS MALLOTOCARPA Warb. Ficus mMucuso Welw. ex Ficalho FICUS NATALENSIS Hochst. FICUS OTTONIIFOLIA (Miq.) Miq. Ficus OvVATA Vahl FIcuSs PRATICOLA Mildbr. & Hutch. Ficus PREUSSII Warb. Not recorded from FP in 12. *FICUS PUMILA L. ‘Ficus scabra (var. Jacq. nec Wild.)’ I do not know what this is. FICUS SIDIFOLIA Welw. ex Hiern FICUS SPIROCAULIS Mildbr., nom. nud. ? F. urceolaris sensu Guinea FP: Musola, Mildbraed 7oro. FIcUS THONNINGII Blume F.. microcarpa Vahl FICUS VOGELIANA (Miq.) Miq. FIcus VoGELi (Miq.) Miq. F. vogelit var. pubicarpa Mildbr. & Burret FICUS WILDEMANIANA Wartb. Not in 12. MESOGYNE HENRIQUESII Engl. MUSANGA CECROPIOIDES R.Br.
M. smithit R.Br.
I, I, cel I,
12 I, II 2
sae ae es © yy 1 25
2
I2 it
» 25
2. %, 2%, ¥2,
13, 25 12 12
22/5 Dos. 25
12, 25
43,43
A PST ST FP PST FP FP FP FPPSTA sl FP ST ST 1 od a Na ST FP EP. Sf ST E PP st FP FE FE. Sa: FP FP FP FP FP ST FP STA FP E? FE FP FP Sf FP ST E FP ST
OF THE GULF OF GUINEA
MyYRIANTHUS PREUSSII Engl.
TRECULIA AFRICANA Decaisne var. AFRICANA
Not recorded from FP in 12.
74. URTICACEAE
*BOEHMERIA NIVEA (L.) Gaudich. BoEHMERIA PLATYPHYLLA D. Don DROGUETIA INERS (Forssk.) Schweinf. Not recorded from FP in 12. ELATOSTEMA MANNII Wedd. ELATOSTEMA MONTICOLA Hook. f. ELATOSTEMA PAIVAEANUM Wedd. ELATOSTEMA THOMENSE Henriq. Perhaps a form of E. paivaeanum. ELATOSTEMA WELWITSCHII Engl.
E. henriquesii Engl. LAPORTEA AESTUANS (L.) Chew Fleurya aestuans (L.) Gaudich. ex Miq.
LAPORTEA ALATIPES Hook. f. Not recorded from FP in 12. LAPORTEA OVALIFOLIA (Schumacher) Chew Fleurya ovalifolia (Schumacher) Dandy _ FF. podocarpa Wedd. LECANTHUS PEDUNCULARIS (Royle) Wedd. PARIETARIA DEBILIS Forster f. P. laxiflora Engl. PILEA MANNIANA Wedd. PILEA MICROPHYLLA (L.) Liebm. PILEA RIVULARIS Wedd. P. ceratomera Wedd. PILEA SUBLUCENS Wedd. PILEA TETRAPHYLLA (Steudel) Blume PovUzoLZIA GUINEENSIS Benth.
POUZOLZIA PARASITICA (Forssk.) Schweinf. Not recorded from FP in 12. PROCRIS CRENATA C. B. Robinson P. laevigata sensu Guinea URERA BATESII Rendle URERA CAMEROONENSIS Wedd.
URERA MANNII (Wedd.) Benth. & Hook. ex Rendle
URERA REPENS (Wedd.) Rendle URERA THONNERI De Wild. ex T. Dur. FP: Escarvé 451. Not in 12.
75. CANNABACEAE
*CANNABIS SATIVA L.
76. AQUIFOLIACEAE
ILEx mitis (L.) Radlk.
iz pO Oe i ig 7 A 25
to: 13 1,2. 12, 19 19
2, 12, 19 £2.10 2, 12:39 1; 33
1 ee Paes Pike a
13, 19 2
Ve 2ehlh2 3s 19, 26 19
12, 19
2
2, 12, 19
19
2, 12
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2, 12, 19
19
Taaei2.13
2, 12, 19
12, 19
19. 2E 12, 13. 19
sk 2, LO M2 ako a O LO EN20 12-10
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19
£2,297
FP FPP St PP. St FP. “Sk FP FP FP FP
ST ERP “oF PEP St A FP 1 Oe ee EP FP
ST FP Peso. FP PE BPE sol FP EPPS A PP. EP. SEA PP PST A FP FP
ST FP SF
363
364 ANGIOSPERMS OF THE ISLANDS
77. CELASTRACEAE (incl. HIPPOCRATEACEAE)
CAMPYLOSTEMON ANGOLENSE Welw. ex Oliver I2 FP HIPPOCRATEA INDICA Willd. I2 FP H. loesenevana Hutch. & Moss 2 HIPPOCRATEA VELUTINA Afzel. jist Sr. MAYTENUS ANNOBONENSIS (Loes. & Mildbr.) Exell 16 A E Gymnosporia annobonensis Loes. & Mildbr. 2 MAYTENUS BUCHANANII (Loes.) R. Wilczek PP FP: Wrigley & Melville 673 (K.) Not in 12. MAYTENUS MONODII Exell 15 i & E SALACIA cf. CUSPIDICOMA Loes. FP FP: Bokoko, Burchardt. SALACIA DEBILIS (G. Don) Walp. 2,2 FP SALACIA MANNII Oliver 212 EP SALACIA PYRIFORMIS (Sabine) Steudel 5 Geen ty SL
78. ICACINACEAE
DESMOSTACHYS TENUIFOLIUS Oliver 2, 12 FP
LASIANTHERA AFRICANA Beauv. 2a, Be
LAVIGERIA MACROCARPA (Oliver) Pierre 12 FY Icacina macrocarpa Oliver 2
NEOSTACHYANTHUS ZENKERI (Engl.) Exell & Mendonga 12 BE
RHAPHIOSTYLIS FERRUGINEA Engl. Pe
FP : Bokoko, Burchardt. Not recorded from FP in 12.
79. OLACACEAE
HEISTERIA PARVIFOLIA Smith : hey Re Bee STA
OLAX GAMBECOLA Baillon TT STA O. viridis Oliver 1,9
STROMBOSIA GRANDIFOLIA Hook. f. ex Benth. 2, 12 FP
STROMBOSIA SCHEFFLERI Engl. 12 FP
STROMBOSIA sp. I A
XIMENIA AMERICANA L., ¥;-25; 26 A
80. OPILIACEAE
UROBOTRYA MINUTIFLORA Stapf FP Opilia minutiflora (Stapf) Engl. 2 Not recorded from FP in 12.
81. LORANTHACEAE
General reference: S. Balle, ‘Les Loranthacées d’Afrique portugaise. I. Guinée et San Tomé’ in Bolm. Soc. Broter., sér. 2, 38 : 9-80, t. 1-16 (1964). ? GLOBIMETULA BRAUNII (Engl.) Tieghem FP? Loranthus zenkeri Engl. 2 Not recorded from FP in 12. Balle considers it unlikely that the species occurs in FP.
HELIXANTHERA MANNII (Oliver) Danser var. MANNII Waa a Ne sT Loranthus mannii Oliver I Loranthus ternatus Tieghem I Helixanthera mannii var. ternata (Tieghem) Balle II
PHRAGMANTHERA CAPITATA (Sprengel) Balle os Loranthus capitatus (Sprengel) Engl. 2 Loranthus incanus Schumacher 2
Phragmanthevra incana (Schumacher) Balle 12
OF THE GULF OF GUINEA 365
PHRAGMANTHERA NITIDULA (Sprague) Balle FP Loranthus nitidulus Sprague 2 Phragmanthera polyscripta Balle pro parte 12 TAPINANTHUS BANGWENSIS (Engl. & Krause) Danser a2 ie VISCUM ENGLERI Tieghem 16 A VISCUM TRIFLORUM DC. Si V. nervosum var. angustifolium Sprague I
82. SANTALACEAE THESIUM TENUISSIMUM Hook. f. 2, 12 FP
83. RHAMNACEAE
GOUANIA LONGIPETALA Hemsley 12 tN LASIODISCUS MANNII Hook. f. TA Pp LASIODISCUS MILDBRAEDII Engl. 1,12 st LASIODISCUS ROZEIRAE Exell 14 ST E MAESOPSIS EMINII Engl. 2. 32,29 BP. St VENTILAGO DIFFUSA (G. Don) Exell ae 8 ae be ST ZIZIPHUS ABYSSINICA Hochst. ex A. Rich. 25 S *ZIZIPHUS MAURITIANA Lam. 13 Sr
84. VITACEAE
AMPELOCISSUS CAVICAULIS (Baker) Planchon 2 RE Not recorded from FP in 12. AMPELOCISSUS MACROCIRRHA Gilg & Brandt 2.12 FP Stated in 12 to be doubtfully distinct from A. cavicaulis. CISSUS ADENOPODA Sprague FP FP: Melville & Wrigley 455b (K). Not recorded from FP in 12. CISSUS ARALIOIDES (Welw. ex Baker) Planchon I PYSE CIssuS ARGUTA Hook. f. 2 PP?
Recorded from FP in Fl. W. Trop. Afr., ed. 1, 1: 475 (1928) butnotin1z2. The FP plant may be C. producta.
CISSUS BARBEYANA De Wild. & Dur. 5; 42 PSf. CISSUS BARTERI (Baker) Planchon yy FP CIssUS CURVIPODA (Baker) Planchon i SE E CIssuUS DEBILIS (Baker) Planchon ae J Sa a ge Sub CISSUS DIFFUSIFLORA (Baker) Planchon 2; 12 FP CISSUS GLAUCOPHYLLA Hook. f. 2 te PP CIssUS GRACILIS Guillemin & Perrottet 22 FP CISSUS LAGENIFLORA Gilg & Brandt 12 FP CiIssUS MANNII (Baker) Planchon 12 FP CIssUS OLIVERI (Engl.) Gilg I st Cissus aff. PETIOLATA Hook. f. I Sa Cissus propuctTa Afzel. T 225 12 PP Si Cissus VOGELII Hook. f. 2, 12 FP *VITIS VINIFERA L, Ets ait &
85. LEEACEAE
LEEA GUINEENSIS G. Don 5 OA) a 73 BP aor LEEA TINcTORIA Lindl. ex Baker TIGA 25 pM E
366 ANGIOSPERMS OF THE ISLANDS
86. RUTACEAE
AFRAEGLE MILDBRAEDII Engl.
*CITRUS AURANTIFOLIA (Christm.) Swingle
*CITRUS AURANTIUM L.
*CITRUS GRANDIS (L.) Osb.
*CITRUS LIMON (L.) Burm. f. C. limonum Risso
*CITRUS NOBILIS Lour.
CLAUSENA ANISATA (Willd.) Hook. f. ex Benth.
FAGARA ANNOBONENSIS Mildbr.
FAGARA MACROPHYLLA Engl.
Not recorded from FP in 12.
FAGARA RUBESCENS (Planchon ex Hook. f.) Engl. F. melanacantha (Planchon ex Oliver) Engl. F. welwitschii Engl.
FAGARA THOMENSIS Engl.
GLYCOSMIS ? AFRICANA Hook. f.
*RUTA CHALEPENSIS L.
TECLEA VERDOORNIANA Exell & Mendonca T. grandifolia sensu Guinea
87. SIMAROUBACEAE
BRUCEA GUINEENSIS G. Don
B. paniculata sensu Mildbr. in Notizbl. Bot. Gart. (1934) is almost certainly B.
Berl. 12: 185 guineensis. Not recorded from FP in 12. HANNOA KLAINEANA Pierre & Engl. FP: Bokoko, Burchardt. *QUASSIA AMARA L,
88. IRVINGIACEAE
DESBORDESIA GLAUCESCENS (Engl.) Tieghem Not recorded from FP in 12.
IRVINGIA GABONENSIS (Aubry-Lecomte ex O’Rorke)
Baillon
89. BURSERACEAE
CANARIUM SCHWEINFURTHII Engl. Not recorded from FP in 12. CANARIUM VELUTINUM Gilg Not in 12. DACRYODES EDULIS (G. Don) H. J. Lam Pachylobus edulis G. Don P. edulis var. mubafo Engl. Not recorded from FP in 12. SANTIRIA TRIMERA (Oliv.) Aubrév. Santiriopsis trimera (Oliv.) Guillaumin ex Engl.
90. MELIACEAE
CARAPA PROCERA DC. *CEDRELA ODORATA L. C. toona sensu Exell ENTANDROPHRAGMA ANGOLENSE (Welw.) C.DC.
Not recorded from FP in 12.
13
12°25, 47 TOURS 2
25) 20527
FP E 7 FP STA FP ST FP ST FP A E FP PST ST ST E ST ST? FP FP FP ST FP P FP FP FP PST | PST | EP St ST FP
OF THE GULF OF GUINEA
GUAREA GLOMERULATA Harms 2, 12 HECKELDORA STAUDTII (Harms) Staner Guarea staudtit Harms 2 Not recorded from FP in 12. *MELIA AZEDARACH L., 2, 25 *TOONA CILIATA M. Roem. 25, 27 Cedrela toona Roxb. ex Willd. I TRICHILIA GRANDIFOLIA Oliver I TRICHILIA MACROPHYLLA Oliver ex Guinea, nom. nud., non T. macrophylla Benth. 2
Apparently an erroneous citation. TRICHILIA MONADELPHA (Thonn.) J. De Wilde
T. heudelotii Planchon ex Oliver 2o12 TRICHILIA RUBESCENS Oliver 212 TURRAEA GLOMERULIFLORA Harms E2) 10 TURRAEA VOGELII Hook. f. ex Benth. Te2eO. Lior)
13 QI. SAPINDACEAE ALLOPHYLUS AFRICANUS Beauv. TEI; 13, 255 26: FP: Bokoko, Mildbraed 6792; Musola, Muildbraed 7018. Not recorded from FP in 12. ALLOPHYLUS GRANDIFOLIUS (Baker) Radlk. Exit ALLOPHYLUS HIRTELLUS (Hook. f.) Radlk. 2°52 ALLOPHYLUS OREODRYADUM Gilg ex Mildbr., nom. nud. 2a?
FP: Mildbraed 6406. BLIGHIA SAPIDA C. Koenig I CARDIOSPERMUM CORINDUM L. I CARDIOSPERMUM HALICACABUM L. Tae a2 CHYTRANTHUS MANNII Hook. f. TLL al 35125 DEINBOLLIA INSIGNIS Hook. f. 252 ERIOCOELUM KERSTINGII Gilg ex Engl. 2
Not recorded from FP in 12. ERIOCOELUM MACROCARPUM Gilg ex Engl.
FP: Bokoko, Burchardt. Notrecorded from FP in 12.
ERIOCOELUM Sp. 52 LACCODISCUS FERRUGINEUS (Baker) Radlk. 2.52 LYCHNODISCUS RETICULATUS Radlk. 2AT 2 PAULLINIA PINNATA L, 1324012 *SAPINDUS SAPONARIA L. I
92. MELIANTHACEAE BERSAMA ABYSSINICA subsp. PAULLINIOIDES (Planchon)
Verdc. 12 B. paullinioides (Planchon) Baker 2 93. ANACARDIACEAE
*ANACARDIUM OCCIDENTALE L. 1,2; 25 ANTROCARYON KLAINEANUM Pierre 2,12 LANNEA WELWITSCHII (Hiern) Engl. I *MANGIFERA INDICA L. £2,260 PSEUDOSPONDIAS MICROCARPA (A. Rich.) Engl. Ey 2, iy ke, 23)
25
FP FP BECP Ss © ST ST BES iS FP A J I er Sdn & FP-<Sf PSE 1 oe FP PST. A PE* Sr Lede & EP FP FP FP FP FP EPPST A FP PPPSrA FP A Peers. 1 Sop nated RP.
367
368 ANGIOSPERMS OF THE ISLANDS
SORINDEIA GRANDIFOLIA Engl.
? SORINDEIA JUGLANDIFOLIA Planchon ; Record probably based on a misidentification of S. grandifolia.
*SPONDIAS CYTHEREA Sonnerat
S. dulcis Forster f.
*SPONDIAS MOMBIN L,
TRICHOSCYPHA BIJUGA Engl.
94. CONNARACEAE
AGELAEA ANNOBONENSIS Schellenb. AGELAEA COCCINEA Exell AGELAEA OBLIQUA var. CORDATA (Schellenb.) Exell AGELAEA OVALIS Schellenb. AGELAEA PHAEOCARPA Exell AGELAEA PILOSA Schellenb. FP: Melville & Wrigley 638 (K). from FP in 12. AGELAEA PRINCIPENSIS Exell AGELAEA PSEUDOBLIQUA Schellenb. AGELAEA RETICULATA Exell ByRSOCARPUS DINKLAGEI (Gilg) Schellenb. ex Hutch. & Dalziel CNESTIS FERRUGINEA DC. FP: Veitch (fide Schellenb. in Engl., Pflanzenr. IV, 127 : 33 (1938)). Not recorded from FP in 12. ? CNESTIS URENS Gilg Not in 12. CONNARUS AFRICANUS Lam. CONNARUS GRIFFONIANUS Baillon PAXIA ZENKERI Schellenb. Not in 12 and not recorded from FP by Schellenberg.
Not recorded
95. ALANGIACEAE
ALANGIUM CHINENSE (Lour.) Harms A. begoniifolium (Roxb.) Baillon
96. ARALIACEAE
*HEDERA HELIX L. POLYSCIAS FULVA (Hiern) Harms *POLYSCIAS GUILFOYLEI var. LACINIATA L. H. Bailey POLYSCIAS QUINTASII Exell SCHEFFLERA BARTERI (Seem.) Harms var. BARTERI S. henriquesiana Harms ex Henriq. Tennant (in Kew Bull. 15 : 332 (1961)) considers that this is a synonym of S. barteri var. barteri. Not recorded from FP in 12. SCHEFFLERA HIERNIANA Harms S. ledermannii Harms Tennant (loc. cit.) refers this with doubt to S. barteri var. bartevi; he does not mention S. hierniana, to which it is referred in 12. SCHEFFLERA MANNII (Hook. f.) Harms
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OF THE GULF OF GUINEA 369
97. UMBELLIFERAE
APIUM LEPTOPHYLLUM (Pers.) F. v. Muell. ex Benth. I Pr CAUCALIS MELANANTHA (Hochst.) Benth. & Hook. ex Hiern 2,9; 12,19 BE C. gracilis sensu Guinea 2 CENTELLA ASIATICA (L.) Urban ; Ty Et F220 425, HE ESE 26 CRYPTOTAENIA AFRICANA (Hook. f.) Drude 12, 19 | ee *ERYNGIUM FOETIDUM L. LE NOI. Solon (aS 1g oda CARS A Us 235. 20,-25 *FOENICULUM VULGARE Miller 2 ee HYDROCOTYLE BONARIENSIS Lam. 5 dome eae HPEPsoe HYDROCOTYLE MANNII Hook. f. 13225223419 BPP -Sil HYDROCOTYLE SIBTHORPIOIDES Lam. T2210 KP. H. monticola Hook. f. 2 *PETROSELINUM CRISPUM (Miller) A. W. Hill FP SI P. sativum Hoffm. 2 P. hortense auctt. 13 PEUCEDANUM WINKLERI H. Wolff 12, 19 FP PIMPINELLA OREOPHILA Hook. f. 2. phe, 19 10) 4 P. petitianum sensu Guinea 2 SANICULA ELATA Buch.-Ham. ex D. Don T2TO FP S. europaea var. elata (Buch.-Ham. ex D. Don) H. Wolff 2 S. europaea sensu Guinea vf
98. THEACEAE
BALTHASARIA MANNII (Oliver) Verdc. ST E Adinandra mannii Oliver Sinals Melchiova mannii (Oliver) Kobuski 15 *CAMELLIA JAPONICA L, I ot *CAMELLIA SINENSIS (L.) Kuntze ST Thea sinensis L. I
99. ERICACEAE
AGAURIA SALICIFOLIA (Commerson) Hook. f. ex Oliver aN as Me FP BLAERIA MANNII (Engl.) Engl. 7 Dees ges PP B. dicranotricha Mildbr. PHILIPPIA MANNII (Hook. f.) Alm & Fries As Pes 2 PP PHILIPPIA THOMENSIS Henriq. 5 ie Xe Se 4 Sr E *RHODODENDRON MUCRONATUM (Blume) G. Don I ST
100. EBENACEAE ? Diospyros ELLIoTII (Hiern) F. White FP Maba mannii Hiern 2 Not recorded from FP in 17. According to F. White almost certainly D. piscatoria.
DIosPyROs FERREA (Willd.) Bakh. st Maba buxifolia (Rottb.) Pers., non Diospyros buxifolia (Blume) Hiern I
DIOSPYROS MELOCARPA F. White 17 FP
370 ANGIOSPERMS OF THE ISLANDS
DIOSPYROS PISCATORIA Giirke Maba bipindensis Girke
FP: Bokoko, Mildbraed 6950. Notrecorded from FP in
17.
IOI. SAPOTACEAE
CHRYSOPHYLLUM ALBIDUM G. Don
Gambeya albida (G. Don) Aubrév. & Pellegrin CHRYSOPHYLLUM CALOPHYLLUM Exell CHRYSOPHYLLUM DELEVOYI De Wild.
C. africanum sensu Baker in Oliver, Fl. Trop. Afr.
3 : 500 (1877) et auctt. (? C. africanum A.DC.).
It is impossible to decide to what species the name Chrysophyllum africanum A.DC. applies; see J. H.
Hemsley in Kew Bull. 20 : 461-465 (1966). CHRYSOPHYLLUM HENRIQUESII Engl. CHRYSOPHYLLUM PRUNIFOLIUM Baker CHRYSOPHYLLUM Sp.
MANILKARA MULTINERVIS (Baker) Dubard
Mimusops cf. LACERA Baker
OMPHALOCARPUM ELATUM Miers
VINCENTELLA DENSIFLORA (Baker) Pierre
VINCENTELLA REVOLUTA (Baker) Pierre Bakerisideroxylon revolutum (Baker) Engl.
102. MYRSINACEAE
AFRARDISIA CYMOSA Mez AFRARDISIA STAUDTII (Gilg) Mez
FP: Musola, Mildbraed 7003 (as A. haemanthus (Gilg)
Mez). The genus is not recorded from FP in 17. MAESA BORJAEANA Henriq. M. kamerunensis Mez M. lanceolata sensu Exell MAESA LANCEOLATA Forssk. RAPANEA NEUROPHYLLA (Gilg) Mez RAPANEA THOMENSIS Exell
103. LOGANIACEAE
ANTHOCLEISTA MICROPHYLLA Wernham A. macrocalyx Philipson A. micrantha Gilg & Mildbr. ex Hutch. & Dalziel A. stenantha Philipson ANTHOCLEISTA SCANDENS Hook. f. A. exelliana Monod A. sp. nov. NvuxXIA CONGESTA R.Br. ex Fresen. var. CONGESTA Lachnopylis annobonensis Mildbr. Nuxia annobonensis Mildbr., nom. nud. NvuxXIA CONGESTA var. THOMENSIS (Philipson) J. Lewis Lachnopylis thomensis Philipson STRYCHNOS ACULEATA Solereder STRYCHNOS ISABELINA Gilg FP: Mildbraed 6353. Notin 17.
1135/27 25
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OF THE GULF OF GUINEA 371
STRYCHNOS QUADRANGULARIS Mildbr., nom. nud., non S.
quadrangularis A. W. Hill FP: Bokoko, Mildbraed 6817. Not in 17.
104. OLEACEAE
JASMINUM FLUMINENSE Vell. J. mauritianum Bojer ex DC.
JASMINUM THOMENSE Exell
*OLEA EUROPAEA L,
OLEA HOCHSTETTERI Baker
OLEA WELWITscHII (Knobl.) Gilg & Schellenb. Steganthus welwitschii (Knobl.) Knobl.
I05. APOCYANACEAE
ALAFIA MULTIFLORA (Stapf) Stapf Holalafia multiflora Stapf
*ALLAMANDA CATHARTICA L,
BAISSEA MULTIFLORA A.DC.,
Not recorded from FP in 17.
*CATHARANTHUS ROSEUS (L.) G. Don Lochnera rosea (L.) Reichenb. f.
*ERVATAMIA DIVARICATA (L.) Burkill E. coronaria (Jacq.) Stapf
FUNTUMIA AFRICANA (Benth.) Stapf Kickxia africana Benth.
FUNTUMIA ELASTICA (Preuss) Stapf Kickxia elastica Preuss
Not recorded from FP in 17. *LANDOLPHIA DAWEI Stapf *NERIUM OLEANDER L.
*PLUMERIA ALBA L,
*PLUMERIA RUBRA L.
RAUVOLFIA DICHOTOMA K. Schum. RAUVOLFIA MACROPHYLLA Stapf RAvVOLFIA vomiToria Afzel.
Not recorded from FP in 17. STROPHANTHUS BULLENIANUS Mast. STROPHANTHUS PREUSSII Engl. & Pax TABERNAEMONTANA BRACHYANTHA Stapf
Conopharyngia brachyantha (Stapf) Stapf TABERNAEMONTANA EGLANDULOSA Stapf Gabunia eglandulosa (Stapf) Stapf TABERNAEMONTANA PACHYSIPHON Stapf Conopharyngia pachysiphon (Stapf) Stapf TABERNAEMONTANA STENOSIPHON Stapf Conopharyngia stenosiphon (Stapf) Stapf TABERNAEMONTANA Sp. aff. STENOSIPHON Conopharyngia sp. nov. ? “TABERNAEMONTANA Sp. ? Conopharyngia sp. *THEVETIA NERIIFOLIA Juss. VOACANGA AFRICANA Stapf VOACANGA LEMOsI!I Philipson
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372 ANGIOSPERMS OF THE ISLANDS
106. ASCLEPIADACEAE
* ASCLEPIAS CURASSAVICA L.
‘Asclepias sericea’ I cannot trace this name. A. serica Raf., a synonym of A. syriaca L., may be intended.
*CALOTROPIS PROCERA (Aiton) Aiton f.
CYNANCHUM ADALINAE (K. Schum.) K. Schum. subsp. ADALINAE
GONGRONEMA LATIFOLIUM Benth.
Marsdenia latifolia (Benth.) K. Schum.
MARSDENIA EXELLII C. Norman
ONCOSTEMMA CUSPIDATUM K. Schum. FP: Bokoko, Mildbraed 6948. Not recorded from FP in 17.
RHYNCHOSTIGMA RACEMOSUM Benth.
TYLOPHORA OBLONGA N. E. Brown Not recorded from FP in 17. The type (Mann 277) is from FP.
TYLOPHORA OCULATA N. E. Brown Not recorded from FP in 17.
TYLOPHORA SYLVATICA Decaisne
I07. RUBIACEAE
AIDIA GENIPIFLORA (DC.) Dandy Randia genipifiora DC. AIDIA PALLENS (Hiern) G. Taylor AIDIA QUINTASII (K. Schum.) G. Taylor AIDIA WATTII G. Taylor ANTHOSPERMUM ASPERULOIDES Hook. f. ARGOSTEMMA AFRICANUM K. Schum. FP : Mildbraed 6308. Not recorded from FP in 17. AULACOCALYX LE-TESTUI (Pellegrin) E. Petit Dorothea le-testui Pellegrin BELONOPHORA COFFEOIDES Hook. f. BERTIERA ANNOBONENSIS G. Taylor BERTIERA LAXA Benth. var. LAXA BERTIERA PEDICELLATA (Hiern) Wernham BERTIERA RACEMOSA (G. Don) K. Schum. var. RACEMOSA B. montana Hiern BERTIERA RETROFRACTA K. Schum. B. obversa K. Krause BoRRERIA OCYMOIDES (Burm. f.) DC. BORRERIA PRINCEAE var. PUBESCENS Hepper BoRRERIA SCABRA (Schumach.) K. Schum. B. monticola Mildbr. BoRRERIA VERTICILLATA (L.) G. F. W. Meyer CANTHIUM DUNLAPII Hutch. & Dalziel CANTHIUM HENRIQUESIANUM (K. Schum.) G. Taylor CANTHIUM SETOSUM Hiern CANTHIUM SUBCORDATUM DC, C. glabriflorum Hiern CANTHIUM sp. aff. ANOMOCARPUM DC.
ae ey 16;.17
HoH NO
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NNN HAN HHH | “J
1,3, 22-135; 14 17
1,27
17
Peer is. 26 I
FP STA he FP FP ST st EP. St EP. FP FP FP FP ST sT ST FP FP ST ST A FP ST PEP St FP FPPSTA FP Ly PST FP ST FP sT P
Hi hi
i
OF THE GULF OF GUINEA 373
CANTHIUM sp. aff. GLABRIFLORUM Hiern 13 SL CANTHIUM sp. I P CANTHIUM sp. 13 ST CEPHAELIS MANNII (Hook. f.) Hiern 17 FP Uragoga mannii (Hook. f.) Hutch. & Dalziel 2 CEPHAELIS PEDUNCULARIS Salisb. F..13 Pot x Uragoga membranifolia Mildbr., nom. nud. 2 CEPHAELIS PEDUNCULARIS var. HYPSOPHILA (Schum. & Krause) Hepper er) FP CEPHAELIS SPATHACEA Hiern bs Ps Sih CHASSALIA CRISTATA (Hiern) Bremek. 17 FP CHASSALIA DONIANA (Benth.) G. Taylor 1p 21 ST CHASSALIA HIERNII (Kuntze) G. Taylor var. HIERNII rae Ned CHASSALIA HIERNII var. GLANDULOSA G. Taylor I P CHASSALIA KOLLY (Schumach.) Hepper 17 FP Psychotria benthamiana Hiern 2
CHASSALIA SUBHERBACEA (Hiern) Hepper 17 FP Psychotria subherbacea Hiern 2
*CINCHONA
Numerous cultivated species are grown in ST; see 1.
*COFFEA ARABICA L, ce Fas tp Gre Ge Vins Ge nme aM Sc eat i 25 *COFFEA LIBERICA Bull ex Hiern cg Pee ie Oe Corey 1 een GW aged adie 6 Re COFFEA MACROCHLAMYS K. Schum. 27. EP *COFFEA STENOPHYLLA G. Don Toy bd tS 25 Si
Many other introduced species and varieties of Coffea are listed in 2 and 13.
CORYNANTHE Sp. ? FP FP: Bokoko, Burchardt. CRATERISPERMUM MONTANUM Hiern I, 13 Fpl A OE CREMASPORA THOMSONII Hiern FP FP: Burchardt. Not recorded from FP in 17. CUVIERA SUBULIFLORA Benth. 15m pd FP DICTYANDRA ARBORESCENS Welw. ex Hook. f. 747 FP DIODIA SCANDENS Swartz 17 FP DIODIA SERRULATA (Beauv.) G. Taylor I Pst EcPOMA CAULIFLORUM (Hiern) N. Hallé in Adansonia 3: 173 (1963). ST E Sabicea cauliflova Hiern Da ST: Rose 208, fide N. Hallé (additional record). ECPOMA GIGANTISTIPULUM (K. Schum.) N. Hallé, loc. cit. 17 EE:P Sabicea gigantistipula K. Schum. 17 P: Rose 419, fide N. Hallé. GALIUM SIMENSE Fresen. a7 i GALIUM THUNBERGIANUM Ecklon & Zeyher ne FP G. biafrae Hiern 2 GARDENIA NIGRIFICANS Wernham 2 FP Not in 17. GEOPHILA AFZELII Hiern 17 P G. cordiformis A. Chev. ex Hutch. & Dalziel I Carinta cordiformis (A. Chev. ex Hutch. & Dalziel) G. Taylor II, 14 GEOPHILA NEURODICTYON (K. Schum.) Hepper 16 - Psychotria neurodictyon K. Schum. i; 2
4
374 ANGIOSPERMS OF THE ISLANDS
G£OPHILA OBVALLATA (Schumacher) Didrichsen Carinta obvallata (Schumacher) G. Taylor GEOPHILA REPENS (L.) I. M. Johnston G. uniflora Hiern Carinta uniflora (Hiern) G. Taylor HEINSIA CRINITA (Afz.) G. Taylor var. CRINITA H. pulchella K. Schum. ? HEINSIA EPIDENDRON Mildbr.
Not in 17. Probably an error for Hymenodictyon
epidendron Mildbr. (= H. biafranum Hiern). HYMENODICTYON BIAFRANUM Hiern IxXORA BREVIFLORA Hiern *ITXORA COCCINEA L. IxoRA FOLIOSA Hiern
FP: Moka, Tessmann 2896. Not recorded from FP
iT: IxXORA GUINEENSIS Benth. I. riparia Hiern IxORA NEMATOPODA K. Schum / I. vosea K. Schum. IxORA THOMEANA (K. Schum.) G. Taylor LASIANTHUS AFRICANUS Hiern LASIANTHUS aff. MANNII Wernham MASSULARIA ACUMINATA (G. Don) Bullock ex Hoyle Randia acuminata (G. Don) Benth. MITRAGYNA CILIATA Aubrév. & Pellegrin MITRIOSTIGMA BARTERI Hook. f. ex Hiern MoRINDA LONGIFLORA G. Don MoRINDA LUCIDA Benth. MORINDA MORINDOIDES (Bak.) Milne-Redh. M. confusa Hutch. MUSSAENDA ARCUATA Lam. ex Poiret MUSSAENDA ELEGANS Schumacher Not recorded from FP in 17. MUSSAENDA ERYTHROPHYLLA Schumacher Not recorded from FP in 17. MUSSAENDA ISERTIANA DC. MUSSAENDA LANCIFOLIA K. Krause MUSSAENDA POLITA Hiern
FP: Bokoko, Mildbraed 6814. Not recorded from
EP sn 59: MuSSAENDA TENUIFLORA Benth. var. TENUIFLORA
MUSSAENDA TENUIFLORA Var. PRINCIPENSIS G. Taylor
MUSSAENDA TENUIFLORA var. THOMENSIS G. Taylor MUSSAENDA aff. AFZELII G. Don NAUCLEA LATIFOLIA Smith Sarcocephalus esculentus Afzel. ex Sabine Not recorded from FP in 17. OLDENLANDIA CORYMBOSA L. OLDENLANDIA HERBACEA (L.) Roxb. OLDENLANDIA LANCIFOLIA (Schumacher) DC. OTOMERIA CAMERONICA (Bremek.) Hepper O. guineensis sensu G. Taylor Tapinopentas latifolia Verdc.
17 17
2
17 2.37 At fy
Te g2500 75 25,020
17 2 7 a Ry
| oy atl
PST
fP.?P
Fr
STA
FP PST A A
FP
FP P
oles
i
OF THE GULF OF GUINEA
OXYANTHUS FORMOSUS Hook. f. ex Planchon O. breviflorus Benth. OXYANTHUS GRACILIS Hiern OXYANTHUS SPECcIOSUS DC. OXYANTHUS sp. A PARAPENTAS SETIGERA (Hiern) Verdc. Vivecta setigera Hiern Pauridiantha camposii (G. Taylor) Exell, comb. nov. Urophyllum camposii G. Taylor in Exell, Cat. Vasc. Pl. S. Tomé : 202 (1944). Holotype : Campos 71 (COI). PAURIDIANTHA CANTHIIFLORA Hook. f. Urophyllum canthiiflorum (Hook. f.) Hutch. & Dalziel PAURIDIANTHA FLORIBUNDA (Schum. & Krause) Bremek. Urophyllum floribundum Schum. & Krause U. annobonense Mildbr., nom. nud. PAURIDIANTHA HIRTELLA (Benth.) Bremek. FP: Bokoko, Mildbraed 6938. Not recorded from PP 17, PAURIDIANTHA INSULARIS (Hiern) Bremek. Urophyllum insulare Hiern PAURIDIANTHA PAUCINERVIS (Hiern) Bremek. Urophyllum paucinerve Hiern PAURIDIANTHA RUBENS (Benth.) Bremek. Urophyllum rubens Benth. PAVETTA BIDENTATA Hiern PAVETTA BRACHYCALYX Hiern FP: Bokoko, Mildbraed 6812. Not recorded from FP in 17. PAVETTA CORYMBOSA (DC.) F. N. Williams P. nitida (Schumach.) Hutch. & Dalziel Recorded from FP in Fl. W. Trop. Afr., ed. 1, but not in 17. PAVETTA GLAUCESCENS Hiern PAVETTA HISPIDA Hiern PAVETTA HOOKERANA Hiern P. exellii Bremek. PAVETTA MONTICOLA Hiern P. dermatophylia Mildbr., nom. nud. PAVETTA NEUROCARPA Benth. PAVETTA ORESITROPHA Bremek. PAVETTA OWARIENSIS Beauv. PAVETTA RIGIDA Hiern PENTAS SCHIMPERANA Subsp. OCCIDENTALIS (Hook. f.) Verdc. P. occidentalis (Hook. f.) Benth. & Hook. ex Hiern PENTODON PENTANDRUS (Schumacher) Vatke Oldenlandia macrophylla DC. POLYCORYNE FERNANDENSIS (Hiern) Keay Gardenia fernandensis Hiern POUCHETIA CONFERTIFLORA Mildbr. POUCHETIA PARVIFLORA Benth.
27
2
257
1,0, 17511 45:37 17
17
2
379 FP
FP STA
FP
PSL A
FP
STA E
PP .S7
st
FP
|e Sol SB
376 ANGIOSPERMS OF THE ISLANDS
PREUSSIODORA SULPHUREA (K. Schum.) Keay Randia exserta K. Schum. ? PSILANTHUS BICARINATUS Mildbr., nom. nud. Perhaps an error for Psychotria bicarinata Mildbr. (= Psychotria recurva Hiern). ? PSILANTHUS EPIPHYTICUM Mildbr., nom. nud. Probably an error for Psychotria epiphytica Mildbr. PSILANTHUS MANNII Hook. f. Heinsia jasminiflova sensu A. Chev. PSYCHOTRIA BIFARIA Hiern PsSYCHOTRIA BRACHYANTHA Hiern PsyCHOTRIA CALVA Hiern PSYCHOTRIA CORNUTA Hiern PSYCHOTRIA CRASSICALYX K. Krause PSYCHOTRIA EPIPHYTICA Mildbr., nom. mnud., non P. epiphytica K. Krause nec P. epiphytica Elmer FP: Mildbraed 6433, 7189. PsSYCHOTRIA ERYTHROPUS K. Schum. FP: Mildbraed 6327. Not recorded from FP in 17. PsSYCHOTRIA EUCHLORA K. Schum. PsyCHOTRIA FERNANDOPOENSIS E. Petit ? Grumilea sp. aff. sbhaerocarpa Hiern Psychotria sphaerocarpa (Hiern) Hutch. & Dalziel, non P. sphaerocarpa Wall. PsyYCHOTRIA GUERKEANA K. Schum. PSYCHOTRIA HENRIQUESIANA K. Schum. Psychotria hierniana Exell, nom. nov. Grumilea macrocarpa Hiern in Oliver, Fl. Trop. Afr. 3: 217 (1877), non Psychotria macrocarpa Hook. f. (1880). Holotype : Mann (K), from S. Tomé ? PSYCHOTRIA INSIDENS Hiern E. Petit considers this to be a new genus of Rubiaceae. The specimen of this species recorded in 17 as Mann 3or has been renumbered Mann 3ora, as there is also a specimen of P. pteropetala numbered Mann 310. PsyCHOTRIA LATISTIPULA Benth. PsyCHOTRIA LEPTOPHYLLA Hiern P. setacea Hiern PsyYCHOTRIA LUCENS Hiern PsyYCHOTRIA MALCHAIREI De Wild. PsyYCHOTRIA MOLLERI K. Schum. PsSyYCHOTRIA MOLLIPES K. Krause FP: above Basilé. Mildbraed 6360. PsYCHOTRIA NUBICOLA G. Taylor PSYCHOTRIA PRINCIPENSIS G. Taylor PsyCHOTRIA PTEROPETALA K. Schum. PSYCHOTRIA RECURVA Hiern Psilanthus carinatus Mildbr., nom. nud. PsSYCHOTRIA SCIADEPHORA Hiern FP: Bokoko, Mildbraed 6853. Not recorded from EP anor, PsYCHOTRIA SUBOBLIQUA Hiern PsyCHOTRIA THOMENSIS G. Taylor
Not in 17.
17
tA Fas © Pee x, Fae ve | I
FP
FP
ST
ST
ST
FP
hi &
hh
i
OF THE GULF OF GUINEA
PsSYCHOTRIA VENOSA (Hiern) E. Petit Grumilea venosa Hiern PSYCHOTRIA sp. FP: Musola, Mildbraed 7042, as Grumilea sp. ROTHMANNIA LONGIFLORA Salisb. Randia maculata DC. ROTHMANNIA MACROCARPA (Hiern) Keay Randia myrmecophylla De Wild. Not in 17. ROTHMANNIA OCTOMERA (Hook.) Fagerl. Randia octomera (Hook.) Hook. f. ROTHMANNIA URCELLIFORMIS (Schweinf. ex Hiern) Bullock ex Robyns R. arcuata Bremek. ? ROTHMANNIA WHITFIELDII (Lindl.) Dandy FP: Bokoko, Mildbraed 6947, identification doubtful. Not recorded from FP in 17. RUTIDEA DECORTICATA Hiern FP: Bokoko, Mildbraed 6822; Musola, Mildbraed 7012. Not recorded from FP in 17, but perhaps these specimens are R. syringoides. RUTIDEA SMITHII Hiern The Annobon plant may be R. smithti var. welwitschii. RUTIDEA SMITHII var. WELWITSCHII Scott Elliot RUTIDEA SYRINGOIDES (Webb) Bremek. RYTIGYNIA CONCOLOR (Hiern) Robyns SABICEA AFRICANA (Beauv.) Hepper Stipularia africana Beauv. Not recorded from FP in 17. SABICEA CALYCINA Benth. SABICEA CAPITELLATA Benth. SABICEA EXELLII G, Taylor SABICEA HIERNIANA Wernham FP: Mildbraed 6288. Not in 17. SABICEA INGRATA K. Schum. SABICEA INSULARIS (Wernham) G. Taylor S. annobonensis Mildbr., nom. nud. SABICEA SCHAEFERI Wernham Also in Cameroon fide N. Hallé. SABICEA URBANIANA Wernham SABICEA URCEOLATA Hepper SABICEA aff. CAULIFLORA Hiern Presumably a species of Ecpboma. SACOSPERMA PANICULATUM (Benth.) G. Taylor Oldenlandia peltospermum Hiern SCHUMANNIOPHYTON MAGNIFICUM (K. Schum.) Harms FP: Bokoko, Mildbraed 6350, 6936. Not recorded from FP in 17. TARENNA BIPINDENSIS (K. Schum.) Bremek. Ixora bipindensis (K. Schum.) K. Schum. ex Hutch. & Dalziel TARENNA CONFERTA (Benth.) Hiern TARENNA GRANDIFLORA (Benth.) Hiern TARENNA NITIDULOIDES G, Taylor
17 P2p4s 25
17 2
T5 II
BP. oot.
FP
Ee
FP
FP
SL.
KP?
SY A/
378 ANGIOSPERMS OF THE ISLANDS
TARENNA Sp. I TARENNA Sp. 2 TARENNA SP. 3 TRICALYSIA BIAFRANA Hiern FP: Bokoko, Mildbraed 6962. Not recorded from EP in) 37. TRICALYSIA OLIGONEURA K. Schum. FP: Bokoko, Mildbraed 6909; a doubtful record. Not recorded from FP in 17. TRICALYSIA PALLENS Hiern UNCARIA AFRICANA G. Don var. AFRICANA ? UROGOGA NuTANS K. Krause Presumably either Cephaelis mannii or C. peduncularis. VIRECTARIA MAJOR (K. Schum.) Verdc. var. MAJOR VIRECTARIA PROCUMBENS (Smith) Bremek.
108. CAPRIFOLIACEAE
*SAMBUCUS CANADENSIS L. FP: Santa Isabel, Mildbraed 6483.
I09Q. COMPOSITAE
* A CANTHOSPERMUM HISPIDUM DC. ADENOSTEMMA MAURITIANUM DC. ADENOSTEMMA PERROTTETII DC.
AGERATUM CONYZOIDES L.
AMBROSIA MARITIMA L. ASPILIA RUDIS Oliver & Hiern subsp. RUDIS BIDENS PILOSA L.
BLuMEA AuRITA (L. f.) DC. Not recorded from FP in 17. *CHRYSANTHEMUM INDICUM L. *CICHORIUM INTYBUS L. CONYZA BONARIENSIS (L.) Cronq. Erigeron sumatrensis Retz. CONYZA CLARENCEANA (Hook. f.) Oliver & Hiern CONYZA FLORIBUNDA Kunth Erigeron floribundus (Kunth) Schultz Bip. CONYZA PERSICIFOLIA (Benth.) Oliver & Hiern Eschenbachia persicifolia (Benth.) Exell CONYZA STEUDELII Schultz Bip. ex A. Rich. FP: Escarvré k-57. Not recorded from FP in 17. CoNYZA THEODOR!I R. E. Fries FP: Escarré e-2. Not recorded from FP in 17. CRASSOCEPHALUM BIAFRAE (Oliver & Hiern) S. Moore Senecio biafrae Oliver & Hiern CRASSOCEPHALUM BOUGHEYANUM C. D. Adams CRASSOCEPHALUM CREPIDIOIDES (Benth.) S. Moore Gynura crepidioides Benth. CRASSOCEPHALUM GRACILE (Hook. f.) Milne-Redh. ex Guinea
I I I
I, 25
2,17
1,-O7 1.135, 10, 27,25
¥5.2,40):9;. 01,3035 £7. 25.20
I
I
Ls ON0;'0, Tig 33: 17, 25, 26
2
z 3, 6, 17
6,517 2.16, 27 1, 8, 345 13,25
OF 17
I
17
OTE OTS, 67, I
6; 17
Y ST ST FP FP? FP A FP FP FP FP Por FP FP STA FPP Si A: PSt Fr FPPSTA FP ST sT Pst FP TP BRS Se FP FP EP SE FP FP FP
OF THE GULF OF GUINEA
CRASSOCEPHALUM MANNII (Hook. f.) Milne-Redh. Senecio mannii Hook. f. CRASSOCEPHALUM MONTUOSUM (S. Moore) Milne-Redh. CRASSOCEPHALUM PICRIDIFOLIUM (DC.) S. Moore CRASSOCEPHALUM RUBENS (Juss. ex Jacq.) S. Moore FP: Moka, Tessmann 2872. Not recorded from FP in 17. CRASSOCEPHALUM VITELLINUM (Benth.) S. Moore Gynura vitellina Benth. DICHROCEPHALA CHRYSANTHEMIFOLIA (Blume) DC. DICHROCEPHALA INTEGRIFOLIA (L. f.) Kuntze D. bicolor (Roth) Schlechtendal ECcLIPTA PROSTRATA (L.) L. ELEPHANTOPUS MOLLIS Kunth
*ELEUTHERANTHERA RUDERALIS (Swartz) Schultz Bip. EMILIA COCCINEA (Sims) G. Don
E. flammea Cass.
E. sagittata DC. EMILIA PRAETERMISSA Milne-Redh.
FP: fide Escarré. Not recorded from FP in 17. EMILIA SONCHIFOLIA (L.) DC. ENYDRA FLUCTUANS Lour.
*EPALTES BRASILIENSIS DC. ETHULIA CONYZOIDES L.f.
FP: Melville & Wrigley 627 (K).
EP 17.
*GALINSOGA CILIATA (Raf.) Blake
FP: fide Escarré.
*GALINSOGA PARVIFLORA Cav. GNAPHALIUM LUTEO-ALBUM L. HELICHRYSUM CyMosvuM (L.) Less.
H. fruticosum Vatke HELICHRYSUM FOETIDUM (L.) Moench HELICHRYSUM GLOBOSUM Schultz Bip. ex A. Rich. HELICHRYSUM MANNII Hook. f.
HELICHRYSUM ODORATISSIMUM (L.) Less. Achyrocline hochstettert Schultz Bip. LAcTUCA CAPENSIS Thunb. LACTUCA GLANDULIFERA Hook. f. var. GLANDULIFERA LAGGERA ALATA var. MONTANA C. D. Adams MELANTHERA SCANDENS (Schumacher) Roberty M. brownei (DC.) Schultz Bip. MICROGLOSSA DENSIFLORA Hook. f. MICROGLOSSA PYRIFOLIA (Lam.) Kuntze MIKANIA CORDATA (Burm. f.) B. L. Robinson var.
CORDATA
M. scandens sensu auctt.
MIKANIA sp.
M. scandens sensu Exell MIKANIOPSIS PANICULATA Milne-Redh.
FP: Melville & Wrigley 625 (K).
from FP in 17.
SENECIO CLARENCEANUS Hook. f
Not recorded from
Not recorded
OFL7
6,517
2
20, is)
16,17
1, TE; 53
I
1 Opel Lk sick 7: 25
TO, 17
17
13
Pe
Lise; Orc AG ED 2 gs yf
I
Lt7
I 1;°6, 53; 14, 17,
Opel 7 t,.15,.27
Lt -16 757 T2103. 7, II I 12,555; 17
7ry OPEN te
FP Feo Sh FP HP FP FP HP -ST PST A FPP SL FP A 15s Saas FP BPO Si PP PST Pe St BP ST Ber St FP re on eg Pr FP FP FP PP FP FP 1B sail Saco EPP A STA EP. “of FP
379
380 ANGIOSPERMS OF THE ISLANDS
SONCHUS OLERACEUS L. SONCHUS SCHWEINFURTHII Oliver & Hiern
FP: fide Escarré. Not recorded from FP in 17.
SPILANTHES FILICAULIS (Schumacher) C. D. Adams S. acmella sensu Exell S. caulirhiza sensu Exell
STRUCHIUM SPARGANOPHORA (L.) Kuntze
Sparganophorus vaillantiit Crantz SYNEDRELLA NODIFLORA (L.) Gaertn.
*TAGETES ERECTA L. TRIPLOTAXIS STELLULIFERA (Benth.) Hutch.
VERNONIA AMYGDALINA Delile
? V. senegalensis sensu Guinea VERNONIA BIAFRAE Oliver & Hiern VERNONIA BLUMEOIDES Hook. f.
Not recorded from FP in 17. VERNONIA CONFERTA Benth. VERNONIA INSIGNIS (Hook. f.) Oliver & Hiern
FP: Escarré e-29. Not recorded from FP in 17.
VERNONIA MOKAENSIS Mildbr. & Mattf. VERNONIA MYRIANTHA Hook. f. *ZINNIA PERUVIANA (L.) L.
110. GENTIANACEAE CANSCORA DECUSSATA (Roxb.) J. A. & J. H. Schultes
EXACUM QUINQUENERVIUM Griseb.
NEUROTHECA LOESELIOIDES (Spruce ex Progel) Baillon
SEBAEA BRACHYPHYLLA Griseb. S. multinodis N. E. Brown SWERTIA ABYSSINICA Hochst. S. clavenceana Hook. f. ‘Swertia camerunensis’ I cannot trace this name.
III. PRIMULACEAE
ARDISIANDRA SIBTHORPIOIDES Hook. f.
II2. PLUMBAGINACEAE
*PLUMBAGO AURICULATA Lam. P. capensis Thunb. PLUMBAGO ZEYLANICA L.
II3. PLANTAGINACEAE
*PLANTAGO MAJOR L. PLANTAGO PALMATA Hook. f.
II4. CAMPANULACEAE
CEPHALOSTIGMA PERROTTETII A.DC. WAHLENBERGIA ARGUTA Hook. f. WAHLENBERGIA MANNII Vatke
5,0, 25; 17
FP
17
I, 6
It
T, 05-9311, 73; 17, 25, 26
2,7
1, .6;, £1; 1:35.10, 17, 25
6
T2340). 01213.
Pies Pee SF
FP Sf
PEE SL
PPE St
FP. SrA
FP
FPP St
FP STA
FP FP
EP) cor
> >
FP
| Ota mie T
scat
a eee
4 4 4 { t
OF THE GULF OF GUINEA
II5. LOBELIACEAE
DIELSANTHA GALEOPSOIDES (Engl. & Diels) E. Wimmer 17 LOBELIA ACUTIDENS Hook. f. 2, 17 LOBELIA BARNSII Exell 1,13; 15 LOBELIA COLUMNARIS Hook. f. Da Pa LOBELIA KAMERUNENSIS Engl. ex Hutch. & Dalziel G LOBELIA MOLLERI Henriq. r,.17 MOoNOPSIS STELLARIOIDES var. SCHIMPERANA (Urban) E.
Wimmer 17
I16. GOODENIACEAE SCAEVOLA PLUMIERI (L.) Vahl I
II7. BORAGINACEAE
CORDIA AURANTIACA Baker Zh 27 *CORDIA GERASCANTHUS L. 13;.27 CoRDIA PLATYTHYRSA Baker 27 C. odorata Giirke Cordia sp. 25 FP: Bokoko, Mildbraed 6939. Not recorded from FP in 17.
CoRDIA SENEGALENSIS Juss. FP: San Carlos, Mildbraed 6932. Not recorded from
PP.in 17, CYNOGLOSSUM LANCEOLATUM Forssk. 2 Not recorded from FP in 17. EHRETIA SCROBICULATA Hiern I EHRETIA ZENKERI Giirke ex Baker & Wright 14527 E. scrobiculata sensu Espirito Santo 25 Treated in 17 as a variety of E. cymosa Thonn. HELIOTROPIUM INDICUM L. E, IF, 53,25 FP: fide Escarré. Not recorded from FP in 17. MYOSOTIS ABYSSINICA Boiss. & Reuter Pa 118. SOLANACEAE *BROWALLIA AMERICANA L. II B. demissa L. 8 *BRUNFELSIA UNIFLORA (Pohl) D. Don Franciscea uniflora Pohl 13 CAPSICUM ANNUUM L, E,:2,10, 25 Not recorded from FP in 17. CAPSICUM FRUTESCENS L. 1 Gee ies Paw Not recorded from FP in 17. *CESTRUM LAEVIGATUM var. PUBERULUM Sendtner d, At, 15, 25 Now quite spontaneous in ST. *CYPHOMANDRA BETACEA (Cav.) Sendtner I *DATURA CANDIDA (Pers.) Saff. II, 16
FP: Moka, Exell 777. Commonasa bush in the grass-
land and as a small tree in the woodland. Used by the
Bubis to make fences for their villages. *DATURA METEL L, 1; 2;.25;.26 *DATURA STRAMONIUM L, 2 DISCOPODIUM PENNINERVIUM Hochst. 237
FP FP
ST FP FP FP. Sf FP
Sr FP
ST FP. SE FP FP
P
no FPP ST FP
ST
ST FP SPA FPP SE PE? SE
ol PP SIA FP’ St |e 8 FP
i
382 ANGIOSPERMS OF THE ISLANDS
*LYCOPERSICUM ESCULENTUM Var. CERASIFORME (Dunal) Alef. L. cerasiforme L. Solanum lycopersicum L. *NICANDRA PHYSALOIDES (L.) Gaertn. Physalis peruviana sensu Guinea *NICOTIANA TABACUM L, *PHYSALIS ANGULATA L. PHYSALIS MICRANTHA Link SOLANUM ACULEATISSIMUM Jacq. SOLANUM DASYPHYLLUM Schumacher S. duplosinuatum Klotzsch Not recorded from FP in 17. SOLANUM INDICUM L. SOLANUM INDICUM var. GRANDEMUNITUM Bitter SOLANUM INDICUM var. MODICEARMATUM Bitter SOLANUM MACROCARPON L. Not recorded from FP in 17. *SOLANUM MELONGENA L, SOLANUM NIGRUM L. sens. lat. S. nodiflorum Jacq. The specimens from our area are probably all referable to S. americanum Mill. SOLANUM TERMINALE Subsp. INCONSTANS (C. H. Wright) Heine S. inconstans C. H. Wright SOLANUM TERMINALE Forssk. subsp. TERMINALE S. bifurcatum Hochst. ex A. Rich. SOLANUM TERMINALE subsp. WELwitscHir (C. H. Wright) Heine S. welwitschii var. strictum C. H. Wright SOLANUM TORVUM Swartz *SOLANUM TUBEROSUM L.
II9Q. CONVOLVULACEAE
*IPOMOEA ALBA L., Calonyction aculeatum (L.) H. D. House C. bona-nox (L.) Bojer Not recorded from FP in 17. IPOMOEA BATATAS (L.) Lam. Not recorded from FP in 17. IPOMOEA CAIRICA (L.) Sweet IPOMOEA ERIOCARPA R. Br. I. hispida (Vahl) Roemer & Schultes, non Zuccagni Not recorded from FP in 17. *IPOMOEA HEDERIFOLIA L. Quamoclit coccinea sensu Exell IPOMOEA INVOLUCRATA Beauv. IPOMOEA MAURITIANA Jacq. I. digitata sensu Exell ? Convolvulus paniculatus sensu Guinea IPOMOEA NIL (L.) Roth I. hederacea sensu Baker & Rendle FP: Buchholz. Not recorded from FP in 17.
I,2
Ty 2). 0) O;- 11, 17
16
FP PST #
FE PST
aA | Gerd oe FPP ST A
FP Sf FPPST A
EP
ST
FP
Lo
ST
FPPSTA
OS eee. el Sma
te
OF THE GULF OF GUINEA 383
IPOMOEA PES-CAPRAE Subsp. BRASILIENSIS (L.) Van 1. 2.-9,.0, 11, 43, EEE SLA
Ooststr. 17 *IPOMOEA PURPUREA Roth a PP IPOMOEA QUAMOCLIT L. 2 17 to ad Sc Quamoclit vulgaris Choisy ae IPOMOEA RUBENS Choisy | oH SF das 9 I. lilacina Blume, non Schrank 2 I. riparia G. Don I Not recorded from FP in 17, but the FP record is doubtful. *ITPOMOEA SETIFERA Poiret I ies IPOMOEA STOLONIFERA (Cyr.) J. F. Gmel. ey) FP Sf Not recorded from FP in 17. *IPOMOEA TILIACEA (Willd.) Choisy t37 P *IPOMOEA TUBEROSA L. 2 EP MERREMIA AEGYPTIA (L.) Urban I sr *MERREMIA DISSECTA (Jacq.) Hallier f. 2,47 ded §3 MERREMIA PTERYGOCAULOS (Choisy) Hallier f. 2 BP. FP: Buchholz. Not recorded from FP in 17. MERREMIA TRIDENTATA subsp. ANGUSTIFOLIA (Jacq.) Van Ooststr. A Si M. angustifolia (Jacq.) Hallier f. I MERREMIA UMBELLATA (L.) Hallier f. subsp. UMBELLATA Lee ie Op a 4, BPP 120. SCROPHULARIACEAE ALECTRA SESSILIFLORA var. MONTICOLA (Engl.) Melch. 16,17 PEP A A. communis Hemsley I CELSIA DENSIFOLIA Hook. 17 FP ‘Celsia densiflora Hook. f.’ 2 *LENDNERIA VERTICILLATA (Mill.) Britton FP Stemodia verticillata (Mill.) Boldingh iby LIMOSELLA AUSTRALIS R.Br. FP L. subulata Ives 17; L. tenuifolia Nutt. 2 LINDERNIA DIFFUSA (L.) Wettst. var. DIFFUSA i G6037 FPP ST A LINDERNIA DIFFUSA var. PEDUNCULATA (Benth.) Skan it, 16797 SEA L. stictantha (Hiern) Skan LINDERNIA VOGELII Skan 17 BP *RUSSELIA EQUISETIFORMIS Schlectendal & Cham. 71,7%3 St R. juncea Zucc. 8 SCOPARIA DULCIS L. I, 25 PSA. SIBTHORPIA EUROPAEA L. 17 FP S. australis Hutch. 2 THUNBERGIANTHUS QUINTASII Engl. I ST TORENIA DINKLAGEI Engl. 17 FP T. parviflora sensu Guinea 2 TORENIA THOUARSII (Cham. & Schlecht.) Kuntze I |e VERONICA MANNII Hook. f. Bp 27, FP I2I, LENTIBULARIACEAE UTRICULARIA MANNII Oliver X45 °45; 29 FPP SL U. bryophila Ridley pS «| UTRICULARIA STRIATULA Smith tip 6 ieee fy 4 FPP
384 ANGIOSPERMS OF THE ISLANDS
I22. GESNERIACEAE
ACANTHONEMA STRIGOSUM Hook. f. DIDYMOCARPUS KAMERUNENSIS Engl. EPITHEMA TENUE C. B. Clarke STREPTOCARPUS ELONGATUS Engl. FP: fide Escarré. Not recorded from FP in 17.
STREPTOCARPUS INSULARIS Hutch. & Dalziel
S. denticulatus Engl., non Turrill STREPTOCARPUS NOBILIS C. B. Clarke STREPTOCARPUS THOMENSIS Exell
I23. BIGNONIACEAE
* JACARANDA MIMOSIFOLIA D. Don KIGELIA AFRICANA (Lam.) Benth. K. impressa Sprague K. pinnata (Jacq.) DC. MARKHAMIA LUTEA (Benth.) K. Schum. NEWBOULDIA LAEVIS (Beauv.) Seem. ex Bur.
*PARMENTIERA ALATA (Kunth) Miers Crescentia alata Kunth
SPATHODEA CAMPANULATA Beauv.
*SPATHODEA NILOTICA Seem.
124. PEDALIACEAE SESAMUM RADIATUM Schumacher
125. ACANTHACEAE
ACANTHOPALE DECEMPEDALIS C. B. Clarke ACANTHOPALE sp.
FP: Boughey 136. ACANTHUS MONTANUS (Nees) T. Anders.
ADHATODA MACULATA C. B. Clarke ADHATODA ORBICULARIS (Lindau) C. B. Clarke ADHATODA ROBUSTA C. B. Clarke ADHATODA TRISTIS Nees AFROFITTONIA SILVESTRIS Lindau ASYSTASIA GANGETICA (L.) T. Anders. ASYSTASIA MACROPHYLLA (T. Anders.) Lindau ASYSTASIA VOGELIANA Benth. BRACHYSTEPHANUS LONGIFLORUS Lindau BRACHYSTEPHANUS OCCIDENTALIS Lindau BRILLANTAISIA LAMIUM (Nees) Benth. Not recorded from FP in 17. The record from FP in 2 may be based on Fl. West Trop. Afr., ed. 1, where the distribution is given as ‘Biafran Is.’ BRILLANTAISIA MADAGASCARIENSIS T. Anders. ex Lindau BRILLANTAISIA NITENS Lindau Not recorded from FP in 17. BRILLANTAISIA Cf. OWARIENSIS Beauv. FP: Escarré j-6.
27
17
z
Pa
ae |
| Oe Fey a ES Pe Br fs 25
2 Oe i re 403 T5
5 a
y ae
ber cuts ie a 17
2, 27
Beas ie
ras Sy
ae Be
17
a 2, Pa fe
Ps te
2,17
2) 17
I
rhe Ag Be a
17
FP FP Fev Ase FP FP
ST oT
st FP
FP FP ST PP
EP. St a
EP.
st PRE PST
FP FP
FP
OF THE GULF OF GUINEA
BRILLANTAISIA PATULA T. Anders. BRILLANTAISIA VOGELIANA (Nees) Benth. CROSSANDRELLA DUSENII (Lindau) S. Moore Not recorded from FP in 17. DICLIPTERA LAXATA C., B. Clarke DICLIPTERA VERTICILLATA (Forssk.) Christensen D. umbellata (Vahl) Juss. DISCHISTOCALYX THUNBERGIIFLORUS (T. Anders.) Benth. ELYTRARIA MARGINATA Vahl E. acaulis sensu Exell ? FILETIA AFRICANA Lindau FP: Musola, Mildbraed 7oor. Not recorded from FP in 17. This may be Asystasia decipiens Heine, but that species is also not recorded from FP. *GRAPTOPHYLLUM PICTUM (L.) Nees ex Griff. HETERADELPHIA PAULOWILHELMIA Lindau HYPOESTES ARISTATA (Vahl) Solander ex Roemer & Schultes H. insularis T. Anders. HYPOESTES TRIFLORA (Forssk.) Roemer & Schultes ‘trifolia’ in 2 HYPOESTES sp. ISOGLOSSA GLANDULIFERA Lindau IsoGLossa sp. B JUSTICIA EXTENSA T. Anders. Duvernoia extensa (T. Anders.) Lindau Not recorded from FP in 17. This may be J. laxa. JUSTICIA INSULARIS T. Anders. Justicia Laxa T. Anders. JUSTICIA MANNII T. Anders. The type is from FP but is apparently untraceable. Anderson himself named two sheets of Mann 1435 (K) as this but, according to E. Milne-Redhead (in litt.), they are definitely not the species described but are Adhadota orbicularis. JUSTICIA PREUSsSII (Lindau) C. B. Clarke FP: Musola, Mildbraed 7047. Not recorded from FP in 17. JUSTICIA TENELLA (Nees) T. Anders. JUSTICIA THOMEENSIS Lindau MENDONCIA GILGIANA (Lindau) Benoist FP: Musola, Mildbraed 6990; between Musola and Concepcion, Wrigley & Melville 549 (K). Not recorded from FP in 17. MIMULOPsSIS SOLMsII Schweinf. M. violacea Lindau NELSONIA CANESCENS (Lam.) Sprengel OREACANTHUS MANNII Benth. PHAULOPSIS FALCISEPALA C. B. Clarke PHAULOPSIS MICRANTHA (Benth.) C. B. Clarke PSEUDERANTHEMUM LUDOVICIANUM (Biittner) Lindau Evanthemum ludovicianum Biittner PSEUDERANTHEMUM TUNICATUM (Afzel.) Milne-Redh. P. nigritianum (T. Anders.) Radlk.
I
oR Fane bay Fas
2
17
16
I, 15
wae hg
LE 7 225,20 i. 2
7,
17
2.
1,17
an Sy f
17
I, 16, 25 17
2
17
2
385
st FP. Sr cP
FP PST A
FP FE St
SD id
BPP St ST E
ST
Ee
FPP ST ST E re
FP
HYP FP FP oF A FP
FP
386 ANGIOSPERMS OF THE ISLANDS
RHINACANTHUS VIRENS (Nees) Milne-Redh. var. VIRENS R. dewevrei De Wild. & Dur. This is recorded from ST in 17 but I have no evidence of its occurrence in that island. *RUELLIA GRAECIZANS Backer RUELLIA PRIMULOIDES (T. Anders. ex Benth.) Heine Endosiphon primuloides T. Anders. ex Benth. RUNGIA PAXIANA (Lindau) C. B. Clarke SCHAUERIA POPULIFOLIA C. B. Clarke SCLEROCHITON PREuSSII (Lindau) C. B. Clarke Not recorded from FP in 17. STENANDRIOPSIS GUINEENSIS (Nees) Benoist Crossandva guineensis Nees STENANDRIOPSIS THOMENSIS (Milne-Redh.) Heine Crossandra thomensis Milne-Redh. *THUNBERGIA ALATA Bojer ex Sims THUNBERGIA VOGELIANA Benth. WHITFIELDIA ELONGATA (Beauv.) De Wild. & Dur. W. longifolia T. Anders. (incl. var. perglabra (C. B. Clarke) Hutch. & Dalziel)
126. VERBENACEAE
CLERODENDRUM BIPINDENSE Giirke CLERODENDRUM CAPITATUM (Willd.) Schumacher FP: Musola, Mildbraed 7043. Not recorded from EP iwcr7: CLERODENDRUM DUSENII Giirke C. thomasit Moldenke Premna macrosiphon Baker Not recorded from FP in 17. CLERODENDRUM GLOBULIFLORUM B. Thomas CLERODENDRUM GRANDIFOLIUM Giirke FP: Mildbraed 6345. Not in 17. *CLERODENDRUM JAPONICUM (Thunb.) Sweet C. fragrans var. pleniflorum Schauer CLERODENDRUM MELANOCRATER Giirke CLERODENDRUM SILVANUM Henriq. C. buchholzit Giirke *CLERODENDRUM SPECIOSISSIMUM Van Geert ex Morren C. fallax Lindl. Not listed in 2 but see photograph following p. 352. CLERODENDRUM SPLENDENS G. Don Not recorded from FP in 17. CLERODENDRUM THOMSONIAE Ball. f. C. scandens sensu Exell CLERODENDRUM UMBELLATUM Poiret C. scandens Beauv. CLERODENDRUM VOLUBILE Beauv. *DURANTA REPENS L. *LANTANA CAMARA L. PREMNA ANGOLENSIS Giirke PREMNA MILNEI Baker STACHYTARPHETA INDICA (L.) Vahl
t1,.27 I
Ee Sr t ST FP Pe FP FP FP ST ST FP FP FP yP EP SP FP FP sT FP FP STA FP SST. FP sT FP. Fe FP ST ST ST FP FP
c #<
OF THE GULF OF GUINEA
VITEX LEHMBACHII Giirke FP: Bokoko, Burchardt. Not recorded from FP in
17.
VITEX MILNEI Pieper 17
V. cienkowskii sensu Guinea 2
127. AVICENNIACEAE
AVICENNIA GERMINANS (L.) L. 14, 15, 16
A. africana Beauv. 1, 13; 27
A. nitida Jacq. 7 iy
128. LABIATAE
ACHYROSPERMUM OBLONGIFOLIUM Baker £327
CoLEus aff. PREUssII Giirke FP: Moka, Mildbraed 7103. Coleus preussii is not mentioned in 17 although the syntypes are from Buea. It is presumably a species of Plectranthus.
*‘ Coleus tuberosus’ 2 Perhaps C. parviflorus Benth. ELSHOLTZIA sp. I HOMALOCHEILOS RAMOSISSIMUS (Hook. f.) J. K. Morton 1 Plectranthus vamosissimus Hook. f. 2 HOSLUNDIA OPPOSITA Vahl Oni ket7 H. verticillata Vahl 2 HyYPTIS LANCEOLATA Poiret 17 LEONOTIS NEPETIFOLIA (L.) Aiton f. bE EN Hh OW Be as A 25, 26 Not recorded from FP in 17. LEUCAS DEFLEXA Hook. f. 17 LEUCAS MARTINICENSIS (Jacq.) Aiton f. 17 *MENTHA ROTUNDIFOLIA (L.) Huds. 13 *OCIMUM BASILICUM L. Ee 250bS Los 27; 25 OcIMUM CANUM Sims 2557 *OCIMUM GRATISSIMUM L, 17 O. viride Willd. i-¢. 31,19, 25, 26 PLATOSTOMA AFRICANUM Beauv. b ae ee PLECTRANTHUS ASSURGENS (Baker) J. K. Morton 7 PLECTRANTHUS DECURRENS (Giirke) J. K. Morton E77. Coleus decurrens Giirke 2 PLECTRANTHUS ESCULENTUS N. E. Brown Coleus dazo A. Chev. 2 Not recorded from FP in 17. PLECTRANTHUS GLANDULOSUS Hook. f. 207.
PLECTRANTHUS INSIGNIS Hook. f. FP: Melville & Wrigley 589 (K); Mildbraed 7143. Not recorded from FP in 17.
PLECTRANTHUS PUNCTATUS L’Hérit. subsp. PUNCTATUS 17 PYCNOSTACHYS MEYERI Giirke 17 P. volkensit Giirke 2 *SALVIA COCCINEA Buc’hoz ex Etlinger I SATUREJA PSEUDOSIMENSIS Brenan 17
Calamintha simensis sensu Guinea 2
FP SrA
PP St
EP A FP FP P-ST.A
EP FP
ST FP STA
SH FPP ST A
|i ere dB
ST
387
388 ANGIOSPERMS OF THE ISLANDS
SOLENOSTEMON MANNII (Hook. f.) Baker 17 SOLENOSTEMON MONOSTACHYUS (Beauv.) Briq. subsp. MONOSTACHYUS 1; 43, 13, 37; 25 SOLENOSTEMON REPENS (Giirke) J. K. Morton 17 STACHYS ACULEOLATA Hook. f. 5 aes G2 *STACHYS ARVENSIS (L.) L. II MONOCOTYLEDONES
129. HYDROCHARITACEAE OTTELIA ULVIFOLIA (Planchon) Walp. 18
I30. POTAMOGETONACEAE
POTAMOGETON SCHEWEINFURTHII A. Benn. 18
I3I. COMMELINACEAE
ANEILEMA BENINIENSE (Beauv.) Kunth 12516 ANEILEMA DISPERSUM Brenan 18 ANEILEMA UMBROSUM (Vahl) Kunth subsp. UMBROSUM 2, 18 A. nigritianum (C. B. Clarke) Hutch. 2 BUFORRESTIA MANNII C. B. Clarke 25,38 ‘Commelina africana C. B. Clarke’ 2
Possibly C. africana L. var. mannii (C. B. Clarke) Brenan is intended. This is not recorded from FP
in 18. COMMELINA BENGHALENSIS L. 2, 16 Not recorded from FP in 18.
COMMELINA CAMEROONENSIS J. K. Morton 18 COMMELINA CAPITATA Benth. 2,18 COMMELINA CONGESTA C. B. Clarke 18
C. condensata C. B. Clarke I. 225 COMMELINA DIFFUSA Burm. f. subsp. DIFFUSA ¥,8, 14,48
C. nudiflora L. pro parte 2, 25 COMMELINA DIFFUSA subsp. MONTANA J. K. Morton 18 COMMELINA ERECTA L. subsp. ERECTA 18
C. vogelit C. B. Clarke 2 CYANOTIS BARBATA D. Don 18
C. mannii C. B. Clarke 2 PALISOTA BARTERI Hook. 2,18 ? PALISOTA BICOLOR Mast. 2, 18 PALISOTA BRACTEOSA C. B. Clarke T2555 PALISOTA HIRSUTA (Thunb.) K. Schum. 18
P. thyrsiflora Benth. 2 PALISOTA MANNII C. B. Clarke 2,18 PALISOTA PEDICELLATA K. Schum. x3 PALISOTA PREUSSIANA K. Schum. ex C. B. Clarke 18 POLLIA CONDENSATA C. B. Clarke I, 2, 8, 18 POLLIA MANNII C. B. Clarke Be) to POLYSPATHA PANICULATA Benth. 2,9, 16
P. glaucescens (C. B. Clarke) Hutch. 2 *RHOEO SPATHACEA (Swartz) Stearn
R. discolor (L’Herit.) Hance ae
STANFIELDIELLA BRACHYCARPA var. HIRSUTA (Brenan) Brenan 18
FP FPPSTA FP
FP ST
FP
EP?
FP A FP
FP
FPO: Sr FP PST A
FP FP
er FP
FP ST
a ee ee es
OF THE GULF OF GUINEA STANFIELDIELLA IMPERFORATA (C, B. Clarke) Brenan 16 1d ae A Saree Buforrestia imperforata C. B. Clarke 1, 2.43
Brenan (in Kew Bull. 14: 285 (1960)) states that one of the syntypes is from FP, but he considers that it may be S. brachycarpa var. hirsuta (see above). *ZEBRINA PENDULA Schnizl. x, 33 Psi:
I32. FLAGELLARIACEAE
FLAGELLARIA GUINEENSIS Schumacher # FP Not recorded from FP in 18.
133. ERIOCAULACEAE
ERIOCAULON ZAMBESIENSE Ruhl. 18 EP, MESANTHEMUM RADICANS (Benth.) F. K6érnicke 2, 18 FP
134. BROMELIACEAE
*ANANAS Comosus (L.) Merr. I BP PSEA A. sativus Lindl.
135. MUSACEAE
*MUSA CHINENSIS Sweet 2 FP *MUSA PARADISIACA L, tp-2p 3, 25 PSP Sil TA: *MUSA PARADISIACA Var. SAPIENTUM (L.) Kuntze 2 FP ST
M. sapientum L. ‘3
M. paradisiaca subsp. sapientum (L.) K. Schum. 25 *MUSA PARADISIACA var. VITTATA (W. Ackerm. ex
Rodigas) K. Schum. I SE
*MUSA ROSACEA Jacq. I ST *MUSA TEXTILIS Née 2 FP *MUSA VENTRICOSA Welw. 13 Si *RAVENALA MADAGASCARIENSIS Sonnerat 2 iP *STRELITZIA REGINAE Banks 2 Pr
136. ZINGIBERACEAE
AFRAMOMUM CHLAMYDANTHUM Loes. & Mildbr. 18 FP AFRAMOMUM DANIELLII (Hook. f.) K. Schum. P02. 5.9, 05 /1sh hee on 18, 25, 26 AFRAMOMUM KENIENSE R. E. Fries FP FP: Melville & Wrigley 487 (K.) Not in 18. AFRAMOMUM LIMBATUM (Oliver & Hanb.) K. Schum. 2938 FP AFRAMOMUM MELEGUETA (Roscoe) K. Schum. T2.18 FP PST AFRAMOMUM PILOSuM (Oliver & Hanb.) K. Schum. 2, 18 BPE AFRAMOMUM SCEPTRUM (Oliver & Hanb.) K. Schum. FP FP: Melville & Wrigley 555 (K). Not recorded from FP in 18. Costus AFER Ker-Gawl. U2, 53390, 26. EEO St 26 CosTUS ENGLERANUS K. Schum. 18 RP COSTUS GIGANTEUS Welw. ex Ridley Ej,015.29,-25 PST A CosTUS LUCANUSIANUS J. Braun & K. Schum. 2, 18 FP CosTUS SPECTABILIS (Fenzl) K. Schum. I ST
*CURCUMA DOMESTICA Valeton I *ELETTARIA REPENS (Sonnerat) Baillon I SAF
5
389
390 ANGIOSPERMS OF THE ISLANDS
*HEDYCHIUM CORONARIUM Koenig RENEALMIA AFRICANA Benth. & Hook. RENEALMIA CINCINNATA (K. Schum.) T. Dur. RENEALMIA GRANDIFLORA Baker
Alpinia africana Ridley RENEALMIA MACROCOLEA K. Schum.
Probably the same as R. africana.
RENEALMIA MANNII Hook. f. *ZERUMBET SPECIOSUM Wendl.
Alpinia speciosa (Wendl.) K. Schum. *ZINGIBER OFFICINALE Roscoe *ZINGIBER ZERUMET (L.) Smith
137. CANNACEAE
*CANNA INDICA L. This should perhaps be treated as a distinct African species, C. bidentata Bertol.
138. MARANTACEAE
HALOPEGIA AZUREA (K. Schum.) K. Schum. Not recorded from FP in 18.
‘Hybophrynium mannianum’
I cannot trace this name. Perhaps it is an error for H. braunianum (see Trachyphrynium below). *MARANTA ARUNDINACEA L. MARANTOCHLOA FILIPES (Benth.) Hutch. Clinogyne filipes (Benth.) K. Schum. ? MARANTOCHLOA FLEXUOSA (Benth.) Hutch. Clinogyne flexuosa (Benth.) K. Schum. This species was recorded from FP in Fl. W. Trop. Afr., ed. 1, but in 18 it was divided between M. leucantha (K. Schum.) Milne-Redh. and M. purpurea (Ridley) Milne-Redh. and neither of these species was recorded from FP.
MARANTOCHLOA LEUCANTHA (K. Schum.) Milne-Redh. FP: Melville & Wrigley 554 (K). Not recorded from FP imr18.
MARANTOCHLOA MANNII (Benth.) Milne-Redh.
Phrynium mannii (Benth.) K. Schum.
MARANTOCHLOA RAMOSISSIMA (Benth.) Hutch.
Clinogyne ramosissima (Benth.) K. Schum. SARCOPHRYNIUM BRACHYSTACHYS (Benth.) K. Schum. Phrynium brachystachys (Benth.) Kérnicke Not recorded from FP in 18. THAUMATOCOCCUS DANIELLII (Benn.) Benth. Not recorded from FP in 18. TRACHYPHRYNIUM BRAUNIANUM (K. Schum.) Baker
139. LILIACEAE
* ASPARAGUS OFFICINALIS L. CHLOROPHYTUM SPARSIFLORUM Baker GLORIOSA SIMPLEX L.
Not recorded from FP in 18.
Eos lt 2,18
re Ge ie
£;.2,.°929; 26
18
P?STA
EP
FP
PST A
FP
Prey
EPP St
aM el Le ORR IES
ee
OF THE GULF OF GUINEA
GLORIOSA SUPERBA L. Not recorded from FP in 18. WURMBEA TENUIS (Hook. f.) Baker
140. SMILACACEAE SMILAX KRAUSSIANA Meissner
I4I. ARACEAE
AMORPHOPHALLUS MANNII N. E. Brown ANCHOMANES DIFFORMIS var. PALLIDUS (Hook.) Hepper A. petiolatus (Hook.) Hutch. The plant recorded as A. difformis (Blume) Engl. in 2 is likely to be this variety. ANUBIAS BARTERI Schott ANUBIAS HETEROPHYLLA Engl. Not in 18. *CALADIUM BICOLOR (Aiton) Vent. CERCESTIS STIGMATICUS N. E. Brown C. dinklagei Engl. pro parte *COLOCASIA ESCULENTA (L.) Schott CULCASIA ANGOLENSIS Welw. ex Schott CULCASIA LANCIFOLIA N. E. Brown C. insulana N. E. Brown CULCASIA PARVIFLORA N. E. Brown CULCASIA SCANDENS Beauv. CYRTOSPERMA SENEGALENSE (Schott) Engl. NEPHTHYTIS CONSTRICTA N. E. Brown NEPHTHYTIS POISSONII (Engl.) N. E. Brown PISTIA STRATIOTES L. Not recorded from FP in 18. RHAPHIDOPHORA AFRICANA N. E. Brown A frorhaphidophora africana (N. E. Brown) Engl. RHEKTOPHYLLUM MIRABILE N. E. Brown SAUROMATUM VENOSvUM (Aiton) Kunth Jaimenostea fernandopoana Guinea & Moreno STYLOCHITON ZENKERI Engl. *? XANTHOSMA SAGITTIFOLIUM (L.) Schott This may be X. mafaffa Schott. * XANTHOSMA VIOLACEUM Schott
142. LEMNACEAE LEMNA PERPUSILLA Torrey
143. AMARYLLIDACEAE
*ALLIUM CEPA L.
*ALLIUM SATIVUM L.
*CRINUM ASIATICUM L.
CRINUM jaGus (Thompson) Dandy
C. giganteum Andr.
Not recorded from FP in 18, presumably in error, as it is recorded in Fl. W. Trop. Afr., ed. 1, under C. giganteum, on the basis of Exell 768, the sheet of which in BM bears a ticket reading ‘Seen for Revised Edition of F.W.T.A.’
FP
FP
KP
FE
FP
nay &
re
391
ts tl
392 ANGIOSPERMS OF THE ISLANDS
CRINUM NATANS Baker
HAEMANTHUS CINNABARINUS Decaisne HAEMANTHUS MULTIFLORUS Martyn *HIPPEASTRUM PUNICEUM (Lam.) Urban HYMENOCALLIS LITTORALIS (Jacq.) Salisb.
144. IRIDACEAE
ACIDANTHERA DIVINA Vaupel *NEOMARICA CAERULEA (Lodd.) Sprague Marica sabinii Lindl.
145. DIOSCOREACEAE
*DIOSCOREA ALATA L. DIOSCOREA BULBIFERA L.
Not recorded from FP in 18. *DIOSCOREA CAYENENSIS Lam. DIOSCOREA DUMETORUM (Kunth) Pax
Not recorded from FP in 18. DIOSCOREA ELEPHANTIPES (L’Hérit.) Engl. D. elephantopus Sprengel DIOSCOREA MINUTIFLORA Engl. The record from FP in 2 is probably based on a mis- identification of D. smilactfolia. DIOSCOREA PREUSSII Pax Not recorded from FP in 18. DIOSCOREA SANSIBARENSIS Pax D. macroura Harms DIOSCOREA SMILACIFOLIA De Wild.
146. AGAVACEAE
* AGAVE SISALANA Perrine *CORDYLINE FRUTICOSA (L.) A. Chev. DRACAENA ADAMII Hepper DRACAENA ARBOREA (Willd.) Link DRACAENA BICOLOR Hook. DRACAENA FRAGRANS (L.) Ker-Gawl. FP: Moka Lake, Exell 809 (BM). Not recorded from FP in 18. DRACAENA LAXISSIMA Engl. DRACAENA MONOSTACHYA Baker DRACAENA PHRYNIOIDES Hook. ? DRACAENA USAMBARENSIS Engl. Not in 18. An unlikely record unless introduced. DRACAENA VIRIDIFLORA Engl. & Krause Not recorded from FP in 18. *FURCRAEA FOETIDA (L.) Haw. SANSEVIERIA GUINEENSIS (L.) Willd. Not in 18. Record in 2 perhaps based on a mis- identified specimen of S. tvifasciata Prain. SANSEVIERIA LONGIFLORA var. FERNANDOPOENSIS N. E. Brown SANSEVIERIA METALLICA Géréme & Labroy
2:58 213 18
I
16
Ey 335. 03,.20) 25
12,13
1, 79;,10)- 23,50 1,2 2,40.0,, 13; 16;
25, 26 2 1 Per pe | 2
Bit; 13,25 25, 26 18
I I 2, 18 2
2, 18 16
FP
FP
ST
I al
OF THE GULF OF GUINEA
147. PALMAE
ANCISTROPHYLLUM OPACUM (Mann & Wendl.) Drude 2, 18 ANCISTROPHYLLUM SECUNDIFLORUM (Beauv.) Wendl. A. acutiflorum Becc. 2
Not recorded from FP in 18.
*ARECA CATECHU L. 2 BORASSUS AETHIOPUM Mart. 1, 133.25 *BUTIA EREOSPATHA (Mart.) Becc. I *CARYOTA URENS L. I *COCOS NUCIFERA L,. aks ELAEIS GUINEENSIS Jacq. be 25291325 ONCOCALAMUS MANNII (Wendl.) Wendl. 2
Not recorded from FP in 18. *OREODOXA REGIA Kunth 2 *PHOENIX DACTYLIFERA L. 2013 Popococcus BARTERI Mann & Wendl. 2
Not recorded from FP in 18. RAPHIA HOOKERI Mann & Wendl. 18 RAPHIA VINIFERA Beauv. 218 SCLEROSPERMA MANNII Wendl. 2
Not recorded from FP in 18.
148. PANDANACEAE
PANDANUS CANDELABRUM Beauv. I
Not recorded from FP in 18, although the type of P.
barteranus Rendle, cited as a synonym, comes from FP. PANDANUS THOMENSIS Henriq. ., 43,725 PANDANUS SP. nov. ?- 15
149. TACCACEAE TACCA LEONTOPETALOIDES (L.) Kuntze I 150. ORCHIDACEAE
AERANGIS FLEXUOSA (Ridley) Schlechter :
A. elegans (Rolfe) Dandy I AERANGIS GRAVENREUTHII (Kranzlin) Schlechter 18, 20 AERANGIS MEGAPHYLLA Summerh, I AERANGIS PHALAENOPSIS Schlechter
Record probably based on a misidentified specimen of A. megaphylla. There is no evidence of more than one species of Aevangis on Annobon. ANCISTROCHILUS THOMSONIANUS (Reichenb. f.) Rolfe 20 Not recorded from FP in 18.
ANCISTRORHYNCHUS CAPITATUS (Lindl.) Summerh. 18, 20 ANCISTRORHYNCHUS METTENIAE (Kranzlin) Summerh. i eae Cephalangraecum metteniae (Kranzlin) Schlechter I ANCISTRORHYNCHUS SERRATUS Summerh. 18, 20
ANGRAECOPSIS DOLABRIFORMIS (Rolfe) Schlechter ee oe ANGRAECOPSIS ISCHNOPUS (Schlechter) Schlechter 20
Not recorded from FP in 18. ANGRAECOPSIS TRIDENS (Lindl.) Schlechter 2, 18, 20
Angraecum tridens Lindl. 2 ANGRAECUM AFFINE Schlechter 20
Not in 18.
FP FP EP. ay ST ST FPPST FP PST. FP FP FR SE FP EP FP FP EPP ST Pp. ST ST FP PE FP ST BP. ST EP FP FP
? ? A A
393
394 ANGIOSPERMS OF THE ISLANDS
ANGRAECUM APOROIDES Summerh. A. distichum var. grandifolium (De Wild.) Summerh. ANGRAECUM ASTROARCHE Ridley ANGRAECUM BIRRIMENSE Rolfe Not recorded from FP in 18. ANGRAECUM CHEVALIERI Summerh. ANGRAECUM DISTICHUM Lindl. ANGRAECUM DORATOPHYLLUM Summerh. ANGRAECUM INFUNDIBULARE Lindl. ANGRAECUM PUNGENS Schlechter ANGRAECUM SUBULATUM Lindl. ANSELLIA AFRICANA Lindl. BOLUSIELLA TALBOTII (Rendle) Summerh. BOLUSIELLA sp. FP: Sanford 5919, 5920. BRACHYCORYTHIS BASIFOLIATA Summerh. BULBOPHYLLUM BEQUAERTII De Wild. BULBOPHYLLUM BUFO (Lindl.) Reichenb. f. BULBOPHYLLUM CALAMARIUM Lindl. Not recorded from FP in 18. BULBOPHYLLUM CALYPTRATUM Krdanzlin BULBOPHYLLUM COCHLEATUM Lindl. BULBOPHYLLUM COCOINUM Bateman ex Lindl. BULBOPHYLLUM COMATUM Lindl. BULBOPHYLLUM DISTANS Lindl. BULBOPHYLLUM FALCATUM (Lindl.) Reichenb. f. BULBOPHYLLUM FERNANDOPOANUM Schlechter, nom. nud. FP: Bokoko, Mildbraed 6892. Not in 18. BULBOPHYLLUM FLAVIDUM Lindl. BULBOPHYLLUM FUERSTENBERGIANUM (De Wild.) De Wild. BULBOPHYLLUM FUSCOIDES J. B. Petersen BULBOPHYLLUM GRAVIDUM Lindl. BULBOPHYLLUM INSULANUM Schlechter, nom. nud. FP : Bokoko, Mildbraed 6935. Not in 18. BULBOPHYLLUM INTERTEXTUM Lindl. BULBOPHYLLUM MANNII Hook. f. BULBOPHYLLUM MAXIMUM (Lindl.) Reichenb. f. BULBOPHYLLUM MEDIOCRE Summerh. B. recurvum sensu Dandy BULBOPHYLLUM MELANORRHACHIS (Reichenb. f.) Reich- enb. f. BULBOPHYLLUM OREONASTES Reichenb. f. BULBOPHYLLUM OXYPTERUM (Lindl.) Reichenb. f. BULBOPHYLLUM QUINTASII Rolfe BULBOPHYLLUM RECURVUM Lindl. BULBOPHYLLUM RESUPINATUM Lindl. BULBOPHYLLUM SCHULTZEANUM Schlechter, nom. nud. FP: Bokoko, Mildbraed 6951. Not in 18. BULBOPHYLLUM TENUICAULE Lindl. BULBOPHYLLUM THOMENSE Summerh. BULBOPHYLLUM VELUTINUM (Lindl.) Reichenb. f. BULBOPHYLLUM WINKLERI Schlechter Not recorded from FP in 18.
18, 20 15
I
20
18, 20 Te2zatt,. 19 I
I, 18
18, 20 213,20 2.°18;,20 TON T5520 20
15
2 LES LS 520 T,<11 1820 12, 18, 20 18, 20
1°35; 18,20 2220
1, 11, 18:20
18, 20 18, 20
2, 18, 20 2:20
18, 20 18, 20 I
16, 18, 20 18, 20 18, 20
18, 20
2, 20 T,.2;,18, 20 I
14, 15, 18 20
FP
ST
ST ST ST
eg ;
— TT ee eer.
OF THE GULF OF GUINEA
CALANTHE CORYMBOSA Lindl.
CALYPTROCHILUM EMARGINATUM (Swartz) Schlechter
CEPHALANGRAECUM ACRODONTUM Schlechter ex Mildbr.,
nom. nud. Ancistrorhynchus acrodontum Sanford, nom. nud. FP: Mildbraed 7149. Not in 18. CHAMAEANGIS THOMENSIS (Rolfe) Schlechter CHAMAEANGIS VAGANS (Lindl.) Schlechter CHAMAEANGIS VESICATA (Lindl.) Schlechter CHAMAEANGIS sp. FP: Sanford 4291. CHEIROSTYLIS LEPIDA (Reichenb. f.) Rolfe CORYMBORKIS CORYMBOSA Thouars C. welwitschii (Reichenb. f.) Kuntze CYNORKIS ANACAMPTOIDES Kranzlin
CYNORKIS CALCARATA (Reichenb. f.) Schlechter ‘Cyno-
sorchis’. FP: Moka, Tessmann 2891. Not in 18. CYNORKIS DEBILIS (Hook. f.) Summerh. CYNORKIS GABONENSIS Summerh. CYRTORCHIS ACUMINATA (Rolfe) Schlechter CYRTORCHIS HENRIQUESIANA (Ridley) Schlechter CYRTORCHIS MONTEIROAE (Reichenb. f.) Schlechter CyYRTORCHIS RINGENS (Reichenb. f.) Summerh. C. bistorta (Rolfe) Schlechter Not recorded from FP in 18. DIAPHANANTHE ACUTA (Ridley) Schlechter DIAPHANANTHE BIDENS (Swartz) Schlechter DIAPHANANTHE BREVIFOLIA (Summerh.) Summerh. Rhipidoglossum brevifolium Summerh. DIAPHANANTHE CURVATA (Rolfe) Summerh. DIAPHANANTHE KAMERUNENSIS (Schlechter) Schlechter Not recorded from FP in 18. DIAPHANANTHE PAPAGAYI (Reichenb. f.) Schlechter DIAPHANANTHE PELLUCIDA (Lindl.) Schlechter DIAPHANANTHE QUINTASII (Rolfe) Schlechter DIAPHANANTHE RUTILA (Reichenb. f.) Summerh. Rhipidoglossum rutilum (Reichenb. f.) Schlechter DIAPHANANTHE SUBCLAVATA (Rolfe) Schlechter FP: San Carlos, Mildbraed 6787. Not in 18. DISPERIS MILDBRAEDII Schlechter ex Summerh. DISPERIS REICHENBACHIANA Welw. ex Reichenb. f. DISPERIS THOMENSIS Summerh. EPIPOGIUM ROSEUM (D. Don) Lindl. EULOPHIA BOULIAWONGO (Reichenb. f.) Raynal E. oedoplectron Summerh. Lissochilus giganteus Welw.
See Raynal in Rev. Soc. Sav. Haute Normandie, 39 : 47
(1966). EULOPHIA CRISTATA (Swartz) Steudel Lissochilus purpuratus Lindl.
Not recorded from FP in 18 or 20, therefore doubtful.
EULOPHIA HORSFALLII (Bateman) Summerh.
2. tty LS)
20 18, 20
20
I I 18) 520
20 Th FS. .20 16, 18, 20 ee?
18, 20
20
Teal helo: 20 15
I
I
18
15, 18, 20
fs, Dif L5; 20, 20
Test, ero,20 18
Too Tid
1B Xo)
2, 18, 20
| oes wh
f Tt 1S, 20 2.16; 15, 20 20
18
2
2,9, 18,20
EP Sts FP FP ST Ls FP 1d 5 Os ace, 1D Same 3 © FP FP 13 see eB i ie A P te EP ST ST Br st P FP i P. A PR Se Sr PP Si BE ST BP St FP A EY EP FP
395
oes
bd tl
396 ANGIOSPERMS OF THE ISLANDS
EULOPHIA MILNEI Reichenb. f. EULOPHIDIUM LATIFOLIUM (Rolfe) Summerh. Eulophia latifolia Rolfe EULOPHIDIUM MACULATUM (Lindl.) Pfitzer E. ledienii (Stein ex N. E. Brown) De Wild. EULOPHIDIUM SAUNDERSIANUM (Reichenb. f.) Summerh. EURYCHONE ROTHSCHILDIANA (O’Brien) Schlechter GENYORCHIS MICROPETALA (Lindl.) Schlechter Polystachya micropetala (Lindl.) Rolfe GENYORCHIS PLATYBULBON Schlechter FP: Gomez Moreno. Not recorded from FP in 18 or 20. GENYORCHIS PUMILA (Swartz) Schlechter GRAPHORKIS LURIDA (Swartz) Kuntze HABENARIA ATTENUATA Hook. f. HABENARIA BARRINA Ridley HABENARIA BRACTEOSA Hochst. ex A. Rich. HABENARIA BUETTNERANA Kranzlin HABENARIA GABONENSIS Reichenb. f. HABENARIA MANNII Hook. HABENARIA MICROCERAS Hook. f. HABENARIA STENOCHILA Lindl. HABENARIA THOMANA Reichenb. f. HETAERIA HETEROSEPALA (Reichenb. f.) Summerh. HETAERIA MANNII (Reichenb. f.) Dur. & Schinz FP: Mildbraed 6400. Not recorded from FP in 18. HETAERIA STAMMLERI (Schlechter) Summerh. LIPARIS DEISTELII Schlechter LIPARIS GOODYEROIDES Schlechter Malaxis sp. LIPARIS GRACILENTA Dandy LIPARIS NERVOSA (Thunb.) Lindl. L. guineensis Lindl. LIPARIS TRIDENS Kranzlin LIPARIS WELWITSCHII Reichenb. f. LipaRIS sp. FP: Sanford 4084. MALAXIS MACLAUDII (Finet) Summerh. MALAXIS PROREPENS (Kranzlin) Summerh. Not recorded from FP in 18. MALAXIS WEBERBAUERANA (Kranzlin) Summerh. MANNIELLA GUSTAVII Reichenb. f. MICROCOELIA sp. FP: Sanford 4167. NERVILIA ADOLPHII Schlechter Not recorded from FP in 18. NERVILIA RENIFORMIS Schlechter Not recorded from FP in 18. NERVILIA UMBROSA (Reichenb. f.) Schlechter OBERONIA DISTICHA (Lam.) Schlechter ORESTIAS STELIDOSTACHYA (Reichenb. f.) Summerh. PLATYLEPIS GLANDULOSA (Lindl.) Reichenb. f.
PODANGIS sp.
18, 20 15, 18 1,51
18
yes OE
18, 20 18, 20 2, 18, 20 2
2
15, 18, 20 II, 16, 20 2, 18, 20
I 10) 18,20 18, 20
14
I; 11, 18,20 2, 18, 20
18, 20
Tutt, 18
i, 43
SRW RAS
20
I, 18,:20 18, 20
15
I
I
20
bey ere Ge Gs. 18, 20
16
20
18, 20 20
18, 20 T2013 71... 20 20
20 20
Po Sk, 2
I
ee ee Gs
1; 2,41, 76; 18. 20
I
FP ST Por FP FP FP FP ye PP SE FP PP 391. FP ST PP rr FP FP P ST st FP EEE FP r i E PPP St A FP A FP FP FP FP FP Sf FP FP FP P St ST PST E FPPSTA ST
OF THE GULF OF GUINEA
POLYSTACHYA AFFINIS Lindl. POLYSTACHYA ALBESCENS Ridley subsp. ALBESCENS
P. epidendroides Schlechter ex Mildbr., nom. nud.
P. imbricata Rolfe POLYSTACHYA ALPINA Lindl. POLYSTACHYA BICALCARATA Kranzlin POLYSTACHYA BIFIDA Lindl.
POLYSTACHYA CALLUNIFLORA Kranzlin Not recorded from FP in 18. POLYSTACHYA CALOGLOSSA Reichenb. f. POLYSTACHYA CALYPTRATA Kranzlin P. elegans pro parte in 18, quoad Sanford 4or3. Not in 18. POLYSTACHYA CAMARIDIOIDES Summerh. Not recorded from FP in 18. POLYSTACHYA CULTRIFORMIS (Thouars) Spreng. POLYSTACHYA DISTICHA Rolfe POLYSTACHYA ELEGANS Reichenb. f. Excl. Sanford 4073 in 18. POLYSTACHYA EXPANSA Ridley POLYSTACHYA FRACTIFLEXA Summerh. Not recorded from FP in 18. POLYSTACHYA FUSIFORMIS (Thouars) Lindl. P. composita Kranzlin POLYSTACHYA LAXIFLORA Lindl. POLYSTACHYA NYANZENSIS Rendle Not in 18. POLYSTACHYA ODORATA Lindl. var. ODORATA POLYSTACHYA OLIGANTHA Schlechter, nom. nud. FP: Mildbraed 7131. Not in 18. POLYSTACHYA PARVIFLORA Summerh. POLYSTACHYA POLYCHAETE Krdanzlin POLYSTACHYA RHODOPTERA Reichenb. f. Not recorded from FP in 18. POLYSTACHYA RIDLEYI Rolfe POLYSTACHYA SETIFERA Lindl. POLYSTACHYA SUPERPOSITA Reichenb. f. POLYSTACHYA TESSELLATA Lindl. POLYSTACHYA THOMENSIS Summerh. POLYSTACHYA cf. MUKANDAENSIS De Wild. POLYSTACHYA Sp. POLYSTACHYA sp. FP: Sanford 5922. RANGAERIS BRACHYCERAS (Summerh.) Summerh. RANGAERIS aff. TRILOBATA Summerh. SARCORHYNCHUS POLYANTHUS (Kranzlin) Schlechter SATYRIUM sp. FP: Tessmann 2890. SOLENANGIS SCANDENS (Schlechter) Schlechter STOLZIA ELAIDUM (Lindl.) Summerh. Bulbophyllum elaidum Lindl.
TRIDACTYLE ANTHOMANIACA (Reichenb. f.) Summerh.
Not recorded from FP in 18. 6
15
1.24, 15. 15,20
2
t,. 13,33
213620
18, 20
132,18, 15, 55, 20
20
2; 18,20 20
18, 20
2
2, 15;:20 20
210, 20 2-420
I 2, 18, 20 Ethie, 26
ilo LO
I
18, 20
Ti.2;,/ 11,18, 20 15
15
I
20
Vea TY Bed ites I
18, 20 20
15, 18 18
I
20
A FPP’Si on FP PP. 1 Se fb Ee EP EY EP FP ST FP ST FP EP PP FP FP FP sr FP EPP Sk STA P FP FP STA ST ST ST FP ST oo iy BP FP ST i 4 EP.
SY
i
398 ANGIOSPERMS OF THE ISLANDS
TRIDACTYLE TRIDACTYLITES (Rolfe) Schlechter
TRIDACTYLE aff. BICAUDATA (Lindl.) Schlechter FP: Sanford 5998.
TRIDACTYLE aff. FUSIFERA Mansf.
TRIDACTYLE Sp.
TRIDACTYLE Sp. FP: Sanford 4162.
* VANILLA CRENULATA Rolfe
VANILLA GRANDIFOLIA Lindl.
*VANILLA PLANIFOLIA Andr. —
ZEUXINE ELONGATA Rolfe The record of this from FP in 20 is apparently an error. It is not recorded from FP in 18.
151. JUNCACEAE LuUZULA CAMPESTRIS var. MANNII Buchenau
I52. CYPERACEAE
BULBOSTYLIS DENSA (Wall.) Hand.-Mazz. var. DENSA B. capillaris sensu Guinea BULBOSTYLIS ERRATICA (Hook. f.) C. B. Clarke CAREX ECHINOCHLOE Kunze C. chlorosaccus C. B. Clarke CAREX LEPTOCLADUS C. B. Clarke CAREX MANNII E. A. Bruce CYPERUS ARTICULATUS L. CYPERUS ATROVIRIDIS C. B. Clarke CYPERUS BARONII C. B. Clarke C. mannii C. B. Clarke CYPERUS COMPRESSUS L. Not recorded from FP in 21. CYPERUS CUSPIDATUS Kunth A: Melville 148 (P). CYPERUS DIFFORMIS L. CYPERUS DIFFUSUS subsp. BUCHHOLZII (Boeck.) Kiiken- thal CYPERUS DISTANS L. f. CYPERUS ESCULENTUS L. CYPERUS EXALTATUS Retz. CYPERUS HASPAN L. *CYPERUS LUZULAE (L.) Retz. ST: Perseveranga, Rose 535 (P). species probably introduced. CYPERUS RENSCHII Boeck. Not recorded from FP in 21. CYPERUS ROTUNDUS L. CYPERUS SPHACELATUS Rottb. CYPERUS SYLVICOLA Ridley ELEOCHARIS GENICULATA (L.) Roemer & Schultes E. caribaea (Rottb.) Blake ST: Perseveranga, Rose 539 (P). ELEOCHARIS MUTATA (L.) Roemer & Schultes FP: Melville 713 (K). Not recorded from FP in 21.
An American
£15, 33; 26 20
15 I 20
I, 18; 25 I, tl, 25 End. 8, 25 I, 18, 20
124583; 21
no Bh HN ee dma SSS Nn oN |
. 8 H Ww
He NHN HSH NHN RD nN i
t. 2 Oo
t.-2,°0- 11, 53; 2%
I I
14
4
aS
ee Fae Ha a Bae
I 21
4
FP..°SE
FP
FP FP
ST FP EP’? FP FP ST
FP
ST
FPP FP PST A PSE st ST ST
FP PST FP PST A
st FP Sf
FP
ree
tt eee
OF THE GULF OF GUINEA
FIMBRISTYLIS CYMOSA R.Br. F. obtusifolia Kunth FIMBRISTYLIS DICHOTOMA (L.) Vahl var. DICHOTOMA FIMBRISTYLIS FERRUGINEA (L.) Vahl Not recorded from FP in 21. FIMBRISTYLIS OVATA (Burm. f.) Kern Abildgaardia monostachya (L.) Vahl FUIRENA UMBELLATA Rottb. HyYPOLYTRUM GRANDE (Uitt.) Koyama Principina grandis Uitt. HyPOLYTRUM HETEROMORPHUM Nelmes H. africanum sensu Dandy Not recorded from FP in 21. HyPOLYTRUM PURPURASCENS Cherm. H. heterophylium sensu Guinea KYLLINGA APPENDICULATA K. Schum. ? EK. cylindrica sensu Guinea KYLLINGA ELATIOR Kunth KYLLINGA ERECTA Schumacher Not recorded from FP in 21. KYLLINGA NEMORALIS (J. R. & G. Forster) Dandy Cyperus kyllingia Endl. KYLLINGA ODORATA Vahl KYLLINGA PERUVIANA Lam. Cyperus peruvianus (Lam.) F. N. Williams Not recorded from FP in 21. KYLLINGA PUMILA Michaux Cyperus tenuifolius Dandy pro parte, quoad specim. KYLLINGA ROBUSTA Boeck. MAPANIA AMPLIVAGINATA K. Schum. M. oblonga C. B. Clarke Not recorded from FP in 21. MAPANIA FERRUGINEA Ridley M. aschersoniana sensu Dandy M. subcomposita C. B. Clarke P: Monod 12126. MaPANIA MACRANTHA (Boeck.) H. Pfeiffer M. deisteliti K. Schum. Not recorded from FP in 21. MARISCUS ALTERNIFOLIUS Vahl Cyperus sublimis (C. B. Clarke) Dandy M. umbellatus sensu Guinea MAaRISCUS DUBIUS (Rottb.) Hutch. Cyperus dubius (Rottb.) C. E. Fischer MARISCUS FLABELLIFORMIS Kunth var. FLABELLIFORMIS Cyperus tenuis Swartz MARISCUS LIGULARIS (L.) Urban Cyperus ligularis L. M. rufus Kunth MARISCUS LONGIBRACTEATUS Cherm. ST: Boa Entrada, Chevalier 13706 (P). MaRISCUS LURIDUs C. B. Clarke
FP: Moka, Tessmann 67. Not recorded from FP in
21.
13 ce a Bal Jo Gate 3 |
I, 4
2.4, 43,15 Tee?
21 1;-9) 12, 43 4
1,0, 3% 21 rots 2, 21
I
4 21
FPPSTA Hie aco
ST A
PSE:
FP-P ST
EP eS
EF.
BP
risk
FP
FPP SIA
bay a
PEP St
PPP Slax
PP: Sr
FP
399
400 ANGIOSPERMS OF THE ISLANDS
MARISCUS TOMAIOPHYLLUS (K. Schum.) C. B. Clarke PYCREUS ELEGANTULUS (Steudel) C. B. Clarke P. cimicinus sensu Guinea PYCREUS POLYSTACHYOS Rottb. Cyperus polystachyos (Rottb.) Beauv. Not recorded from FP in 21. PYCREUS SMITHIANUS (Ridley) C. B. Clarke REMIREA MARITIMA Aubl. Cyperus pedunculatus (R.Br.) Kern Not recorded from FP in 2r. RHYNCHOSPORA CORYMBOSA (L.) Britton SCIRPUS BRACHYCERAS Hochst. S. inclinatus (Delile) Aschers. & Graebner may prove to be an earlier name for this species. SCLERIA BOIVINII Steudel S. barteri Boeck. SCLERIA LAGOENSIS Boeck. S. canaliculato-triquetra Boeck. SCLERIA NAUMANNIANA Boeck. TORULINIUM ODORATUM (L.) Hooper Cyperus odoratus L. Not recorded from FP in 21.
153. GRAMINEAE
ACROCERAS ZIZANIOIDES (Kunth) Dandy AGROSTIS MANNII (Hook. f.) Stapf ? AGROSTIS TROPICA Beauv. ANDROPOGON AMETHYSTINUS Steudel A. abyssinicus sensu Guinea ANDROPOGON GAYANUS var. SQUAMULATUS (Hochst.) Stapf Not recorded from FP in 2t1. ANDROPOGON MACROPHYLLUS Stapf ANDROPOGON MANNII Hook. f. ANTHEPHORA CRISTATA (Doell) Hackel ex De Wild. & Dur. ARISTIDA ADSCENSIONIS L. *AXONOPUS COMPRESSUS (Swartz) Beauv. AXONOPUS FLEXUuOSUS (Peter) C. E. Hubbard ex Troupin *BAMBUSA VULGARIS Schrader ex Wendl. BECKEROPSIS LAXIOR W. D. Clayton B. nubica sensu Dandy Dandy followed Stapf & Hubbard (in Prain, Fl. Trop. Afr. 9 : 952 (1934)), who included in B. nubica (Hochst.) Fig. & De Not. the West African plants later segregated by Clayton as B. laxior. BRACHYPODIUM FLEXUM Nees BROMUS LEPTOCLADOS Nees B. scabridus Hook. f. Not recorded from FP in 21. CENTOTHECA LAPPACEA (L.) Desvaux
CHLOACHNE OPLISMENOIDES (Hackel) Stapf ex Robyns CHLORIS PILOSA Schumacher CHLORIS PYCNOTHRIX Trin.
2, 21 21 aoa
bey es Peay fm Se Pe
259;2r Pers 21
3
3
2
2; 321 S411, 16 QO, 11
8, II
14, 21
E38 5.13,0L0 TO;2T Eers
ee Pe A aS © Pe
21 2I
i, 2, 13, 15;:22
ie, 25
FPP ST
PPP &
FP FP
FP PSI
FP
FP PST PST
a te
° ain te SiG EN aE
OF THE GULF OF GUINEA
*COIX LACRYMA-JOBI L. CTENIUM NEWTONII Hackel *CYMBOPOGON CITRATUS (DC.) Stapf CYNODON DACTYLON (L.) Pers. Not recorded from FP in 21. CyRTOCOCCUM CHAETOPHORON Dandy C. setigerum Stapf Not recorded from FP in 21. DACTYLOCTENIUM AEGYPTIUM (L.) Beauv. DESCHAMPSIA CESPITOSA (L.) Beauv. DIGITARIA HORIZONTALIS Willd. D. velutina sensu Guinea DIGITARIA LONGIFLORA (Retz.) Pers. ECHINOCHLOA COLONUM (L.) Link ELEUSINE INDICA (L.) Gaertn.
(Roemer & Schultes)
E. indica forma unispicata Sobrinho ERAGROSTIS DOMINGENSIS (Pers.) Steudel E. linearis (Schumach.) Benth. ERAGROSTIS MOKENSIS Pilger ERAGROSTIS SUPERBA Peyr. ERAGROSTIS TENELLA (L.) Beauv. ex Roemer & Schultes Not recorded from FP in 21. ERIOCHLOA NUBICA (Steudel) Hackel & Stapf ex Thell. Not recorded from FP in 21. FESTUCA ABYSSINICA Hochst. ex A. Rich. F. schimperana A. Rich. FESTUCA CAMUSIANA subsp. CHODATIANA St.-Yves F. simensis sensu Guinea FESTUCA MEKISTE W. D. Clayton F. gigantea sensu Guinea HELICTOTRICHON MANNII (Pilger) C. E. Hubbard
HETEROPOGON CONTORTUS (L.) Beauv. ex Roemer & Schultes
*HORDEUM VULGARE L. HyYPARRHENIA DIPLANDRA (Hackel) Stapf HYPARRHENIA RUFA (Nees) Stapf var. RUFA HyYPARRHENIA RUFA var. MAJOR (Rendle) Stapf ISACHNE BUETTNERI Hackel ISACHNE MAURITIANA Kunth I. aethiopica Stapf & Hubbard LEERSIA HEXANDRA Swartz LEPTASPIS COCHLEATA Thwaites L. conchifera Hackel MEGASTACHYA MUCRONATA (Poiret) Beauv. Centotheca mucronata (Poiret) Kuntze C. urekana Guinea MELINIS MINUTIFLORA Beauv. Not recorded from FP in 21. OLYRA LATIFOLIA L, OPLISMENUS BURMANNI (Retz.) Beauv.
OPLISMENUS HIRTELLUS (L.) Beauv.
1,3, 10 2%
i 23 54l O E203, 45
7
14
7a TE 3
io.0). 01, 25
2, 3
14
13, 14
I, 3; 7; Il, 13, PS u2T
8
21
1, 14
7 ae ae |
14
3, 13, 15
TO, 17
21 2, 3
21
2,3
21
2,3
Sea 5 2%
I
13
I
tO. 3
I
FR ae ig ae See 21
T1313
4, 21
Lt; 3.97). 21
2
21
3 3 E539: 73-23
bee Pay Pas Fae S ame 2 §
E20, 28,44; 21
ne ER. Mi ee ee Oe 35.10, 25
FP. SPA FP PEP SEA EP. SE FP ST EP FP PST A ST ST EPP SrA ST FP ST BPP St FP STA FP EP FP FP PSTA ot ST STA ST EPP OL PE St FP BPE or FP BP: SEA FPP ST EP STA FPP STA
401
402 ANGIOSPERMS OF THE ISLANDS
*ORYZA SATIVA L. *OXYTENANTHERA ABYSSINICA (A. Rich.) Munro PANICUM ACROTRICHUM Hook. f.
P. acrotrichum var. pubescens Guinea PANICUM BREVIFOLIUM L,
PANICUM HOCHSTETTERI Steudel PANICUM MAXIMUM Jacq. Not recorded from FP in 21. PANICUM MONTICOLA Hook. f. P. macrophyllum Guinea ex W. D. Clayton PASPALUM CONJUGATUM Bergius
PASPALUM ORBICULARE Forster f.
P. commersonii Lam.
P. scrobiculatum var. commersonii (Lam.) Stapf PASPALUM PANICULATUM L. PASPALUM VAGINATUM Swartz
PENNISETUM HORDEOIDES (Lam.) Steudel PENNISETUM MONOSTIGMA Pilger PENNISETUM POLYSTACHION (L.) Schultes PENNISETUM PURPUREUM Schumacher ? PHRAGMITES KARKA (Retz.) Trin. ex Steudel P. vulgaris sensu Guinea ? Not recorded from FP in 21.
australis (Cav.) Trin. ex Steudel. Poa ANNUA L. POA LEPTOCLADA Hochst. ex A. Rich. PSEUDECHINOLAENA POLYSTACHYA (Kunth) Stapf PUELIA CILIATA Franchet
P. acuminata Pilger
Not recorded from FP in 21. RHYNCHELYTRUM REPENS (Willd.) C. E. Hubbard RHYNCHELYTRUM REYNAUDIOIDES C. E. Hubbard ROTTBOELLIA EXALTATA (L.) L. f. *SACCHARUM OFFICINARUM L. SACCIOLEPIS AFRICANA C. E. Hubbard & Snowden SCHIZACHYRIUM BREVIFOLIUM (Swartz) Nees ex Biise *SECALE CEREALE L. SETARIA BARBATA (Lam.) Kunth
SETARIA CAUDULA Stapf SETARIA CHEVALIERI Stapf
SETARIA LONGISETA Beauv.
SETARIA MEGAPHYLLA (Steud.) Dur. & Schinz SETARIA MILDBRAEDII Mez ex C. E. Hubbard SETARIA Sp. aff. MILDBRAEDII Mez ex C. E. Hubbard SORGHUM ARUNDINACEUM (Desvaux) Stapf
*SORGHUM DRUMMONDII (Steudel) Nees ex Millsp. &
Chase
The plant recorded by Guinea is much more likely to be this species than P.
27353
3
Pee
3
see Pan Pay Fe: Pa SP 16, 21
5,-2.3, 813; at
1:20, 86234
21
7
12.4; 9) 8, 33, IO; 21
3 WO elE ils a2 2, 3, 9, II, 14,
2523-25 2ea ak Etsy 1 4). 10 1 tape FG At eZ
I I, 2
to 4S, 21 bier. oe | I
¥2. 3,5, 0,25, 21
2T
£26, kk, £4; 25
1:0, 435-53
i2. 4, 6; $35 25
I
14
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SPOROBOLUS MOLLERI Hackel SPOROBOLUS PYRAMIDALIS Beauv. Not recorded from FP in 21. SPOROBOLUS ROBUSTUS Kunth Not recorded from FP in 2r. SPOROBOLUS TENUISSIMUS (Schrank) Kuntze S. minutiflorus (Trin.) Link SPOROBOLUS VIRGINICUS (L.) Kunth Not recorded from FP in 21. STENOTAPHRUM SECUNDATUM (Walt.) Kuntze Not recorded from FP in 21. STREBLOCHAETE LONGIARISTATA (A. Rich.) Pilger STREPTOGYNA CRINITA Beauv. S. gerontogaea Hook. f. *TRITICUM AESTIVUM L.
T. vulgare Vill. *ZEA MAYS L.
NOTE
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Whilst this work was in the press the following was received: M. C. Liberato. Flora de S. Tomé e Principé Papilionaceae. 1972. Additional records contained in it, kindly communicated by its author, are
included in this paper.
INDEX TO THE FAMILIES AND GENERA IN THE SYSTEMATIC LIST
Synonyms are printed in italics.
In the text the genera and species are arranged alphabetic-
ally in each family. When a species is not to be found under the genus in which it was originally recorded it will usually have been transferred to another genus and will probably be found under
one of the references to the original genus as a synonym.
Abelmoschus, 350 Abilgaardia, 399 Abutilon, 350, 351 Abrus, 355
Acacia, 355, 356 Acalypha, 353 ACANTHACEAE, 384 Acanthonema, 384 Acanthopale, 384 Acanthospermum, 378 Acanthus, 384 Achyranthes, 341 Achyrocline, 379 Achyrospermum, 387 Acidanthera, 392 Acioa, 355 Acridocarpus, 351 Acroceras, 400
Adansonia, 350 Adenanthera, 356 Adenia, 344 Adenocarpus, 356 Adenopus, 345 Adenostemma, 378 Adhatoda, 384, 385 Adinandra, 369 Aerangis, 393 Aeschynomene, 356 Aerva, 341 Afraegle, 366 Aframomum, 389 Afrardisia, 370 Afrofittonia, 384 Afrorhaphidophora, 391 Afzeliella, 348 Agauria, 369
AGAVACEAE, 392 Agave, 392 Agelaea, 368 Ageratum, 378 Agrostis, 400 Aidia, 372 Airyantha, 356 AIZOACEAE, 341 Alafia, 371 ALANGIACEAE, 368 Alangium, 368 Albizia, 356 Alchemilla, 354 Alchornea, 352 Alectra, 383 Aleurites, 352 Allamanda, 371 Allanblackia, 349
404
Allium, 391 Allophyllus, 367 Alpinia, 390 Alternanthera, 341 Alysicarpus, 356 AMARANTHACEAE, 341 Amaranthus, 341 AMARYLLIDACEAE, 391 Ambrosia, 378 Amorphophallus, 391 Ampelocissus, 365 Amphiblemma, 347 ANACARDIACEAE, 367 Anacardium, 367 Ananas, 389 Anchomanes, 391 Ancistrochilus, 393 Ancistrophyllum, 393 Ancistrorhynchus, 393 Ancistrorhynchus, 395 Andropogon, 400 Aneilema, 388 Angraecopsis, 393 Angraecum, 393, 394 Angraecum, 393 Anisophyllea, 348 Annona, 336 ANNONACEAE, 336 Ansellia, 394 Anthephora, 400 Anthocleista, 370 Anthonotha, 356 Anthospermum, 372 Anthostema, 352 Antiaris, 361 Antidesma, 349, 352 Antigonon, 341 Antrocaryon, 367 Anubias, 391
Apium, 369 APOCYANACEAE, 371 AQUIFOLIACEAE, 363 ARACEAE, 391 Arachis, 356 ARALIACEAE, 368 Ardisiandra, 380
Areca, 393
Argemone, 339 Argostemma, 372 Aristida, 400 Aristolochia, 338 ARISTOLOCHIACEAE, 338 Artabotrys, 336 Artocarpus, 361
INDEX
ASCLEPIADACEAE, 372 Asclepias, 372
Asimina, 336
Asparagus, 390
Aspilia, 378
Asystasia, 384, 385 Athroandra, 353 Aulacocalyx, 372 ‘Aurea’, 341
Averrhoa, 342 AVERRHOACEAE, 342 Avicennia, 387 AVICENNIACEAE, 387 Axonopus, 400
Baikiaea, 356
Baissea, 371 Bakerisideroxylon, 370 BALSAMINACEAE, 342 Balthasaria, 369 Bambusa, 400
Baphia, 356
Baphia, 356 Baphiastrum, 356 BARRINGTONIACEAE, 347 Barteria, 344
Basella, 342 BASELLACEAE, 342 Bauhinia, 356 Bauhinia, 359 Beckeropsis, 400 Begonia, 345, 346 BEGONIACEAE, 345 Belonophora, 372 Berlinia, 356
Berria, 349
Bersama, 367
Bertiera, 372
Bidens, 378 BIGNONIACEAE, 384 Bixa, 344 BIXACEAE, 344 Blaeria, 369
Blighia, 367
Blumea, 378 Boehmeria, 363 Boerhavia, 343 Bolusiella, 394 BOMBACACEAE, 350 Bombax, 350 BORAGINACEAE, 381 Borassus, 393
Borreria, 372 Bosqueia, 362
Bougainvillea, 343 Brachycorythis, 394 Brachypodium, 400 Brachystephanus, 384 Brassica, 339 Breynia, 352 Bridelia, 352 Brillantaisia, 384, 385 BROMELIACEAE, 389 Bromus, 400 Browallia, 381 Brucea, 366 Brunfelsia, 381 Bryonopsis, 345 Bryophyllum, 340 Buchholzia, 339 Buforrestia, 388 Buforrestia, 389 Bulbophyllum, 394 Bulbophyllum, 397 Bulbostylis, 398 BURSERACEAE, 366 Butia, 393 BUXACEAE, 361 Buxus, 361 Byrsocarpus, 368
CACTACEAE, 346 Caesalpinia, 356 Cajanus, 356 Caladium, 391 Calamintha, 387 Calanthe, 395 Caloncoba, 344 Caloncoba, 344 Calonyction, 382 Calopogonium, 356 Calotropis, 372 Calvoa, 347 Calpyptrochilum, 395 Camellia, 369 CAMPANULACEAE, 380 Camptostylus, 344 Campylospermum, 346 Campylostemon, 364 Cananga, 336 Canarium, 366 Canavalia, 356 Canna, 390 CANNABACEAE, 363 Cannabis, 363 CANNACEAE, 390 Canscora, 380 Canthium, 372, 373
Caperonia, 352 CAPPARACEAE, 339 Capparis, 339 CAPRIFOLIACEAE, 378 Capsella, 339 Capsicum, 381 Carapa, 366 Cardamine, 339 Cardiospermum, 367 Carex, 398
Carica, 346 CARICACEAE, 346 Carinta, 373, 374 Carpolobia, 340 CARYOPHYLLACEAE, 340 Caryota, 393 Casearia, 344
Cassia, 356, 357 Cassipourea, 348 Castilloa, 362 Catharanthus, 371 Caucalis, 369 Cavacoa, 352 Cayaponia, 344 Cecropia, 362
Cedrela, 366
Cedrela, 367
Ceiba, 350 CELASTRACEAE, 364 Celosia, 341, 342 Celsia, 383
Celtis, 361
Centella, 369 Centotheca, 400 Centotheca, 401 Centrosema, 357 Cephaelis, 373, 378 Cephalangraecum, 395 Cephalangraecum, 393 Cephalostigma, 380 Cerastium, 340 Cercestis, 391
Cercis, 357
Cestrum, 381 Chaetacme, 361 Chamaeangis, 395 Chasmanthera, 338 Chassalia, 373 Cheirostylis, 395 CHENOPODIACEAE, 341 Chenopodium, 341 Chloachne, 400 Chloris, 400 Chlorophora, 362
INDEX
Chlorophytum, 390 Chrysanthemum, 378 CHRYSOBALANACEAE,
355 Chrysobalanus, 355 Chrysophyllum, 370 Chytranthus, 367 Cichorium, 378 Cinchona, 373 Cinnamomum, 337 Cissampelos, 338 Cissus, 365 Citrullus, 344 Citrus, 366 Claoxylon, 352 Clausena, 366 Cleistanthus, 352 Cleistopholis, 336 Clematis, 338 Cleome, 339 Clerodendrum, 386 Clinogyne, 390 Clitorea, 357 Cnestis, 368 Coccinia, 345 Cocos, 393 Codiaeum, 352 Coelocaryon, 337 Coffea, 373 Coix, 401 Cola, 350 Coleus, 387 Coleus, 387 Colocasia, 391 Colocynthis, 344 Colophospermum, 357 COMBRETACEAE, 348 Combretum, 348 Commelina, 388 COMMELINACEAE, 388 COMPOSITAE, 378 CONNARACEAE, 368 Connarus, 368 Conocarpus, 348 Conopharyngia, 371 CONVOLVULACEAE, 382 Convolvulus, 382 Conyza, 378 Copatfera, 357 Corchorus, 349 Cordia, 381 Cordyline, 392 Coronopus, 339 Corymborkis, 395
405
Corynanthe, 373 Costus, 389 Crassocephalum, 378, 379 Crassula, 340 CRASSULACEAE, 340 Craterispermum, 373 Cremaspora, 373 Crescentia, 384 Crinum, 391, 392 Crossandra, 386 Crossandrella, 385 Crotalaria, 357 Croton, 352 Crotonogyne, 352 CRUCIFERAE, 339 Crudia, 357 Cryptotaenia, 369 Ctenium, 401 Cucumeropsis, 345 Cucumis, 345 Cucurbita, 345 CUCURBITACEAE, 344 Culcasia, 391 Curcuma, 389 Cuviera, 373 Cyanotis, 388 Cyathula, 342 Cydonia, 354 Cymbopogon, 401 Cynanchum, 372 Cynodon, 401 Cynoglossum, 381 Cynometra, 357 Cynometra, 361 Cynorkis, 395 “Cynosorchis’, 395 CYPERACEAE, 398 Cyperus, 398 Cyperus, 399, 400 Cyphomandra, 381 Cyrtococcum, 401 Cyrtorchis, 395 Cyrtosperma, 391
Dacryodes, 366 Dactyloctenium, 401 Dalbergia, 357 Daniellia, 357 Datura, 381 Deeringia, 342 Deinbollia, 367 Delonix, 357 Desbordesia, 366 Deschampsia, 401
406
Desmanthus, 357 Desmodium, 357, 358 Desmostachys, 364 Dialium, 358 Diaphananthe, 395 Dicellandra, 347 DICHAPETALACEAE, 355 Dichapetalum, 355 Dichrocephala, 379 Dicliptera, 385 Dicranolepis, 343 Dictyandra, 373 Didymocarpus, 384 Dielsantha, 381 Digitaria, 401
Dillenia, 344 DILLENIACEAE, 344 Dimorphochlamys, 345 Dinophora, 347 Dioclea, 358
Diodia, 373
Dioscorea, 392 DIOSCOREACEAE, 392 Dioscoreophyllum, 338 Diospyros, 369, 370 Diplocyclos, 345 Diplotaxis, 339 Dischistocalyx, 385 Discoclaoxylon, 352, 353 Discoglypremna, 353 Discopodium, 381 Disperis, 395
Dissotis, 347 Distemonanthus, 358 Dolichos, 359 Dorothea, 372 Dorstenia, 362 Dracaena, 392 Drepanocarpus, 358 Droguetia, 363 Drymaria, 340 Drypetes, 353 Duboscia, 349 Duranta, 386 Duvernoia, 385
EBENACEAE, 369 Echinochloa, 401 Eclipta, 379 Ecpoma, 373, 377 Ehretia, 381 Elaeis, 393 Elaeophorbia, 353 Elatostema, 363
INDEX
Eleocharis, 398 Elephantopus, 379 Elettaria, 389 Eleusine, 401 Eleutheranthera, 379 Elsholtzia, 387 Elytraria, 385 Emilia, 379
Enantia, 336 Endosiphon, 386 Entada, 358 Entandrophragma, 366 Enydra, 379 Epaltes, 379 Epipogium, 395 Epithema, 384 Eragrostis, 401 Evanthemum, 385 Eriboma, 350 ERICACEAE, 369 Erigeron, 378 Eriobotrya, 355 ERIOCAULACEAE, 389 Eriocaulon, 389 Eriochloa, 401 Eriocoelum, 367 Eriosema, 358 Ervatamia, 371 Eryngium, 369 Erythrina, 358 Erythrococca, 353 Erythrophloeum, 358
ERYTHROXYLACEAE, 352
Erythroxylum, 352 Eschenbachia, 378 Ethulia, 379 Eucalyptus, 347 Eugenia, 347 Eugenia, 347 Eulophia, 395, 396 Eulophia, 396 Eulophidium, 396 Euphorbia, 353 EUPHORBIACEAE, 352 Eurychone, 396 Exacum, 380 Excoecaria, 353
Fagara, 366 Festuca, 401 Ficus, 362 Filetia, 385 Fimbristylis, 399 Flabellaria, 351
Flacourtia, 344 FLACOURTIACEAE, 344 Flagellaria, 389 FLAGELLARIACEAE, 389 Fleurya, 363
Foeniculum, 369 Franciscea, 381
Fuirena, 399
Fumaria, 339 FUMARIACEAE, 339 Funtumia, 371
Furcraea, 392
Gabunia, 371 Galinsoga, 379 Galium, 373 Gambeya, 370 Garcinia, 349 Gardenia, 373 Gardenia, 375 GENTIANACEAE, 380 Genyorchis, 396 Geophila, 373, 374 GERANIACEAE, 342 Geranium, 342 GESNERIACEAE, 384 Globimetula, 364 Gloriosa, 390, 391 Glossocalyx, 337 Glycine, 358 Glycosmis, 366 Glyphaea, 349 Gnaphalium, 379 Gomphrema, 342 Gongronema, 372 GOODENIACEAE, 381 Gossypium, 351 Gouania, 365 GRAMINEAE, 400 Graphorkis, 396 Graptophyllum, 385 Grevillea, 343 Grewia, 349 Griffonia, 358 Grossera, 353 Grosseva, 352 Grumilea, 376, 377 Guarea, 367
Guarea, 367 GUTTIFERAE, 349 Guyonia, 348 Gymnosporia, 364 Gynandropsis, 339 Gynura, 378, 379
INDEX 407
Habenaria, 396 Icacina, 364 LEGUMINOSAE, 355
Haemanthus, 392 ICACINACEAE, 364 Lemna, 391 Haematoxylum, 358 Tlex, 363 LEMNACEAE, 391 Halopegia, 390 Impatiens, 342 Lendneria, 383 HALORAGACEAE, 343 Indigofera, 358 LENTIBULARIACEAE, 383 Hannoa, 366 Inga, 358, 359 Leonotis, 387 Harungana, 349 Ipomoea, 382, 383 Lepidoturus, 352 Heckeldora, 367 Iresine, 342 Leptaspis, 401 Hedera, 368 IRIDACEAE, 392 Leptoderris, 359 Hedychium, 390 Irvingia, 366 Leptonychia, 350 Heinsia, 374 IRVINGIACEAE, 366 Leucaena, 359 Heinsia, 376 Isachne, 401 Leucas, 387 Heisteria, 364 Isoglossa, 385 Leucomphalos, 359 Helichrysum, 379 Ixora, 374 LILIACEAE, 390 Helictotrichon, 401 Ixora, 377 Limosella, 383 Heliotropium, 381 LINACEAE, 352
f Helixanthera, 364 Jacaranda, 384 Lindackeria, 344
i Hernandia, 337 Jaimenostea, 391 Lindernia, 383
1 HERNANDIACEAE, 337 Jasminum, 371 Liparis, 396
} Hetaeria, 396 ; Jateorhiza, 338 Lissochilus, 395
y" Heteradelphia, 385 Jatropha, 353 Lobelia, 381 Heteropogon, 401 JUNCACEAE, 398 LOBELIACEAE, 381 Heteropteris, 351 Jussiaea, 343 Lobularia, 339 Hevea, 353 Justicia, 385 Lochnera, 371 Hibiscus, 351 LOGANIACEAE, 370 Hibiscus, 350 Kalanchoe, 340 Lonchocarpus, 359 Hippeastrum, 392 | Kickxia, 371 LORANTHACEAE, 364 Hippocratea, 364 Kigelia, 384 Loranthus, 364, 365 HIPPOCRATEACEAE, Klaineanthus, 353 Ludwigia, 343
364 Kyllinga, 399 Luffa, 345
Holalafia, 371 Luzula, 398 Homalium, 344 LABIATAE, 387 Lychnodiscus, 367 Homalocheilos, 387 Lablab, 359 Lycopersicum, 382
; Hordeum, 401 Laccodiscus, 367 LYTHRACEAE, 343
: Hoslundia, 387 Lachnopylis, 370
: Hugonia, 352 Lactuca, 379 Maba, 369, 370 HUGONIACEAE, 352 Lagenaria, 345 Macaranga, 353 HUMIRIACEAE, 351 Laggera, 379 Macrolobium, 356
i Hybanthus, 340 Laguncularia, 348 Maesa, 370 Hybophrynium, 390 Landolphia, 371 Maesobotrya, 353, 354
; Hybophrynium, 390 Lannea, 367 Maesopsis, 365
Hydrangea, 340 Lantana, 386 Malaxis, 396 HYDRANGEACEAE, 340 Laportea, 363 Malaxis, 396 HYDROCHARITACEAE, Lasianthera, 364 Mallotus, 354
388 Lasianthus, 374 MALPIGHIACEAE, 351
Hydrocotyle, 369 Lasiodiscus, 365 Malus, 355 Hymenocallis, 392 LAURACEAE, 337 MALVACEAE, 350 Hymenodictyon, 374 Laurembergia, 343 Malvastrum, 351 Hyparrhenia, 401 Lavigeria, 364 Mammea, 349 HYPERICACEAE, 349 Lawsonia, 343 Mangifera, 367 Hypericum, 349 Lecanthus, 363 Manihot, 354 Hypoestes, 385 Leea, 365 Manilkara, 370 Hypolytrum, 399 LEEACEAE, 365 Manniella, 396
Hyptis, 387 Leersia, 401 Manniophyton, 354
408
Mapania 399 Maranta, 390 MARANTACEAE, 390 Marantochloa, 390 Mareya, 354 Margaritaria, 354 Marica, 392 Mariscus, 399, 400 Markhamia, 384 Marsdenia, 372 Marsdenia, 372 Massularia, 374 Maytenus, 364 Medinilla, 348 Megastachya, 401 Melanthera, 379 MELASTOMATACEAE, 347 Melchiora, 369 Melia, 367 MELIACEAE, 366 MELIANTHACEAE, 367 Melinis, 401 Melochia, 350 Melothria, 345 Memecylon, 348 Mendoncia, 385 MENISPERMACEAE, 338 Mentha, 387 Merremia, 383 Mesanthemum, 389 Mesogyne, 362 Microcoelia, 396 Microdesmis, 354 Microglossa, 379 Mikania, 379 Mikaniopsis, 379 Millettia, 359 Millettia, 359 Mimosa, 359 Mimulopsis, 385 Mimusops, 370 Mirabilis, 343 Mitragyna, 374 Mitriostigma, 374 Momordica, 345 MONIMIACEAE, 337 Monodora, 336 Monopsis, 381 MORACEAE, 361 Morinda, 374 Mucuna, 359 Musa, 389 MUSACEAE, 389 Musanga, 362
INDEX
Mussaenda, 374 Myosotis, 381 Myrciaria, 347 Myrianthemum, 348 Myrianthus, 363 Myrica, 361 MYRICACEAE, 361 Myristica, 337 MYRISTICACEAE, 337 MYRSINACEAE, 370 MYRTACEAE, 347
Napoleonaea, 347 Nasturtium, 339, 340 Nauclea, 374
Nelsonia, 385 Neoboutonia, 354 Neomarica, 392 Neostachyanthus, 364 Nephthytis, 391
Nerium, 371
Nervilia, 396 Neurotheca, 380 Newbouldia, 384 Nicandra, 382 Nicotiana, 382 Nothosaerva, 342
Nuxia, 370 NYCTAGINACEAE, 343 Nymphaea, 338 NYMPHAEACEAE, 338
Oberonia, 396 Ochna, 346 OCHNACEAE, 346 Ocimum, 387 OLACACEAE, 364 Olax, 364 Oldenlandia, 374 Oldenlandia, 375, 377 Olea, 371 OLEACEAE, 371 Olyra, 401 Omphalocarpum, 370 ONAGRACEAE, 343 Oncoba, 344 Oncocalamus, 393 Oncostemma, 372 Ophiobotrys, 344 Opilia, 364 OPILIACEAE, 364 Oplismenus, 401 Opuntia, 346 ORCHIDACEAE, 393
Oreacanthus, 385 Oreodoxa, 393 Oreosyce, 345 Orestias, 396 Ormocarpum, 359 Ormosia, 359 Oryza, 402 Ostryocarpus, 359 Otomeria, 374 Ottelia, 388 Ouratea, 346 OXALIDACEAE, 342 Oxalis, 342 Oxyanthus, 375 Oxymitra, 337 Oxytenanthera, 402
Pachylobus, 366 Palisota, 388 PALMAE, 393 PANDANACEAE, 393 Pandanus, 393 Panicum, 402 PAPAVERACEAE, 339 Parapentas, 375 Pararistolochia, 338 Parietaria, 363 Parinari, 355
Parkia, 359 Parmentiera, 384 Paspalum, 402 Passiflora, 344 PASSIFLORACEAE, 344 Paullinia, 367 Pauridiantha, 375 Pavetta, 375 Pavonia, 351
Paxia, 368 PEDALIACEAE, 384 Peddiea, 343 Pedicellaria, 339 Peltophorum, 359 Penianthus, 338 Pennisetum, 402 Pentaclethra, 359 Pentadesma, 349, 352 Pentadesma, 349 Pentas, 375 Pentodon, 375 Peperomia, 338, 339 Peponium, 345 Persea, 337 Petroselinum, 369 Peucedanum, 369
Phaseolus, 359 Phaulopsis, 385 Philippia, 369 Philoxerus, 342
Phoenix, 393 Phragmanthera, 364, 365 Phragmites, 402 Phrynium, 390 Phyllanthus, 354 Phyllanthus, 352 Physalis, 382
Physalis, 382
Physedra, 345 Physostigma, 359 Phytolacca, 341 PHYTOLACCACEAE, 341 Pilea, 363
Piliostigma, 359 Pimpinella, 369
Piper, 339 PIPERACEAE, 338 Piptadenia, 358, 360 Piptadeniastrum, 360 Pistia, 391 Pithecellobium, 360 PITTOSPORACEAE, 344 Pittosporum, 344 PLANTAGINACEAE, 380 Plantago, 380 Platostoma, 387 Platylepis, 396 Plectranthus, 387 Plectranthus, 387 PLUMBAGINACEAE, 380 Plumbago, 380
Plumeria, 371
Poa, 402
Podangis, 396 Podococcus, 393 PODOSTEMACEAE, 340 Poinciana, 357
Pollia, 388
Polyalthia, 337 Polycoryne, 375 POLYGALACEAE, 340 POLYGONACEAE, 341 Polygonum, 341 Polyscias, 368 Polyspatha, 388 Polystachya, 397 Polystachya, 396 Popowia, 337
Portulaca, 341. PORTULACACEAE, 341
INDEX
Potamogeton, 388 POTAMOGETONACEAE, 388 Pouchetia, 375 Pouzolzia, 363 Premna, 386 Premna, 386 Preussiodora, 376 PRIMULACEAE, 380 Principina, 399 Procris, 363 PROTEACEAE, 343 Protomegabaria, 354 Prunus, 355 Prunus, 355 Pseudagrostistachys, 354 Pseudechinolaena, 402 Pseuderanthemum, 385 Pseudospondias, 367 Psidium, 347 Psilanthus, 376 Psilanthus, 376 Psophocarpus, 360 Psychotria, 376, 377 Psychotria, 373 Pterocarpus, 360 Pterygota, 350 Puelia, 402 Pueraria, 360 Punica, 343 PUNICACEAE, 343 Pupalia, 342 Pycnanthus, 337 Pycnocoma, 354 Pycnostachys, 387 Pycreus, 400 Pygeum, 355 Pyrus, 354
Quamoclit, 382, 383 Quassia, 366 Quisqualis, 348
Randia, 372, 374, 376, 377 Rangaeris, 397 RANUNCULACEAE, 338 Ranunculus, 338, 355 Rapanea, 370
Raphanus, 339
Raphia, 393 Raphidiocystis, 345 Rauvolfia, 371
Ravenala, 389
Remirea, 400
409
Renealmia, 390 Rhabdophylium, 346 RHAMNACEAE, 365 Rhaphidophora, 391 Rhaphiostylis, 364 Rhaptopetalum, 349 Rhektophyllum, 391 Rhinacanthus, 386 Rhipidoglossum, 395 Rhipsalis, 346 Rhizophora, 348 RHIZOPHORACEAE, 348 Rhododendron, 369 Rhoeo, 388 Rhynchelytrum, 402 Rhynchosia, 360 Rhyncospora, 400 Rhyncostigma, 372 Ricinodendron, 354 Ricinus, 354 Rinorea, 340 Ritchiea, 339 Rorippa, 339, 340 Rosa, 355 ROSACEAE, 354 Rothmannia, 377 Rottboellia, 402 RUBIACEAE, 372 Rubus, 355
Rubus, 338, 355 Ruellia, 386 Rumex, 341 Rungia, 386 Russelia, 383
Ruta, 366 RUTACEAE, 366 Ruthalicia, 345 Rutidea, 377
Rytigynia, 377
Sabicea, 377 Sabicea, 373 Saccharum, 402 Sacciolepis, 402 Sacoglottis, 351 Sacosperma, 377 Sagina, 340 Salacia, 364 Salvia, 387 Samanea, 360 Sambucus, 378 SAMYDACEAE, 344 Sanicula, 369 Sansevieria, 392
410
SANTALACEAE, 365 Santiria, 366 Santiriopsis, 366 SAPINDACEAE, 367 Sapindus, 367 Sapium, 354 Sapium, 353 SAPOTACEAE, 370 Sarcocephalus, 374 Sarcophrynium, 390 Sarcorhynchus, 397 Satureja, 387 Satyrium, 397 Sauromatum, 391 Scaevola, 381 Schaueria, 386 Schefflera, 368 Schizachyrium, 402 Schumanniophyton, 377 Scirpus, 400 Scleria, 400 Sclerochiton, 386 Sclerosperma, 393 Scoparia, 383 Scorodophloeus, 360 SCROPHULARIACEAE, 383 SCYTOPETALACEAE, 349 Scytopetalum, 349 Sebaea, 380 Secale, 402 Sechium, 345 Senecio, 379 Senecio, 378, 379 Sericostachys, 342 Sesamum, 384 Sesbania, 360 Sesuvium, 341 Setaria, 402 Sibthorpia, 383 Sida, 351 SIMAROUBACEAE, 366 Sinapis, 340 SMILACACEAE, 391 Smilax, 391 SOLANACEAE, 381 Solanum, 382 Solanum, 382 Solenangis, 397 Solenostemon, 388 Sonchus, 380 Sophora, 360 Sorghum, 402 Sorindeia, 368
INDEX
Sparganophorus, 380 Spathodea, 384 Spilanthes, 380 Spondianthus, 354 Spondias, 368 Sporobolus, 403 Stachys, 388 Stachytarpheta, 386 Stanfieldiella, 388, 389 Staudtia, 337 Steganthus, 371 Stellaria, 340 Stemodia, 383 Stenandriopsis, 386 Stenotaphrum, 403 Stephania, 338 Sterculia, 350 STERCULIACEAE, 350 Stipularia, 377 Stolzia, 397 Streblochaete, 403 Strelitzia, 389 Streptocarpus, 384 Streptogyna, 403 Strombosia, 364 Strophanthus, 371 Struchium, 380 Strychnos, 370, 371 Stylochiton, 391 Swertia, 380 Symphonia, 349 Synedrella, 380 Syrrheonema, 338
Syzygium, 347
Tabernaemontana, 371 Tacca, 393 TACCACEAE, 393 Tagetes, 380 Talinum, 341 Tamarindus, 360 Tapinanthus, 365 Tapinopentas, 374 Tapura, 355 Tarenna, 377, 378 Teclea, 366 Telanthera, 341 Telfairia, 345 Tephrosia, 360 Teramnus, 360 Terminalia, 348 Tetracarpidium, 354 Tetracera, 344
Tetrapleura, 360 Tetrorchidium, 354 Thalictrum, 338 Thaumatococcus, 390 Thea, 369 THEACEAE, 369 Thecacoris, 354 Thecacoris, 353, 354 Theobroma, 350 Thesium, 365 Thevetia, 371 Thunbergia, 386 Thunbergianthus, 383 THYMELEACEAE, 343 TILIACEAE, 349 Toona, 367
Torenia, 383 Torulinium, 400 Trachyphrynium, 390 Tragia, 354
Treculia, 363
Trema, 361 Tricalysia, 378 Trichilia, 367 Trichoscypha, 368 Triclisia, 338 Tridactyle, 397, 398 Trifolium, 360 Triplotaxis, 380 Tristemma, 348 Tristicha, 340 Triticum, 403 Triumfetta, 349 Turraea, 367 Tylophora, 372
Uapaca, 354 ULMACEAE, 361 UMBELLIFERAE, 369 Uncaria, 378
Unona, 337
Uragoga, 378
Uragoga, 373
Uraria, 360
Urena, 351
Urera, 363
Urobotrya, 364 Urophyllum, 375 URTICACEAE, 363 Utricularia, 383
Uvaria, 337 | Uvaria, 337 | Uvariodendron, 337 |
E. 8
Vanilla, 398 Ventilago, 365 VERBENACEAE, 386 Vernonia, 380 Veronica, 383 Vicia, 360
Vigna, 360, 361 Vincentella, 370 Viola, 340 VIOLACEAE, 340 Virecta, 375 Virectaria, 378 Viscum, 365
ARTHUR W. EXELL CHURCH GATES BLOCKLEY
MorRETON-IN-MarsH, GLOs.
INDEX
VITACEAE, 365 Vitex, 387
Vitis, 365 Voacanga, 371
Wahlenbergia, 380 Waltheria, 350 Whitfieldia, 386 Wissadula, 351 Wurmbea, 391
Xanthosma, 391 Ximenia, 364
Xylopia, 337 Xymalos, 337
Zea, 403
Zebrina, 389
Zehneria, 345 Zenkerella, 361 Zerumbet, 390
Zeuxine, 398
Zingiber, 390 ZINGIBERACEAE, 389 Zinnia, 380
Ziziphus, 365
411
INDEX TO VOLUME 4
The page numbers of the principal references and the new taxonomic names are printed in bold-face type, synonyms in ttalics. Names of intraspecific taxa are indexed only when they are new or are
synonyms.
Acanthaceae, A survey of the tropical genera,
Oplonia & Psilanthele : : 261-323 Achimenes . ‘ ‘ , 28 1tS3 Acrostichum ear : i : é 96
angulatum . ; : : ; Se 8S commutatum A : 3 . 3 go conforme . : 2 : . 83,89, 90 laurifolium : ; ; : : go marginatum s ; ; . 83, 89,90 neritfolium ‘ é ‘ : Sie OZ spatulatum : : : : 2) 290 stelligerum x : : ; : 92 stigmatolepis : . ; 3 86, 88 viscosum . . : ; : : 92 yunnanense ; : : : 2 92 Aecidium amaryllidis . ; : chee Lil] complanatum ; : : hes, crini. ; : : : ; = eh ly/ infrequens . : : : : San ly) scutellariae : : ' pena ke urticae var. himalayense : é ets Agaricaceae . : 3 ‘ 3 -. IO Agaricus comatus . 2 : : 1 a L20 disseminatus : : : : EL2O eymineus . : ; ; : Lin laccatus . : ; : : GITZO melleus : : : ; : 2 TIO mollis 3 : ‘ 2 : ~ 520 pubescens . : : : : 120 squarrosus . ; ; : : LT tabescens . ; : P 5 qe op Ate) tigvinus . : : : 2 PGBS: Albuginaceae : : : : oe LOO Albugo bliti . : : : : . 100 Aleuria aurantia . : F : . 101 Alloplectus . : : oe) 1845, 232 Alloplectus affinis : : : . 189 aureonitens : : ; . 189 calotrichus : ; : : Ae Cy) cristatus . ‘ ‘ _ ‘ aeeey cubensis . : : : ‘ . 189 grisebachianus . : : 215, 234 gvisebachianus var. pchrotrichus : Fr O51" guianensis : z ‘ ‘ . 189 pubescens . F ; F . 215,232 Amauroderma rugosum . F , . 122 Anthacanthus : , : : ~ 297 Anthacanthus acicularis . 302, 307, 310, 312, 313 arvmatus. : ‘ : ‘i . #305
bispinosus . ; : : ‘ « 320
cuneatus emarginatus jamaicensis lycioides microphyllus nannophyllus purpurascens repandus sinuatus spinosus
var. horridus sprengelit tetrvastichus vincoides
Ardisia
sect. Icacorea
Ardisia
subgen. Ardisia . subgen. Icacorea subgen. Tinus
Ardisia brittonii
byrsonimae clusioides compressa . coriacea decipiens densiflora . dictyoneura gentler harrisiana . humilis vosea solanacea . spicigera tinifolia troyana urbanii xylosteoides
Armillaria mellea . Arthrobotryum nilgirense Aschersonia viridans
disciformis viridula
Ascomycetes Asterostromella dura
vhodospora
Auricularia delicata
mesenterica polytricha .
Auriculariaceae
304,
157; 156, 157;
157; 157;
157; 156, 157,
157; 157,
3°97 305 312 307 3°97 313 313 320 321 312 395 321 310 315 156 164
157 164 164 159 162 168 164 158 164 162 161 162 158 165 160 165 162 158 159 160 I7I 119 138 136 136 136 IOI 132 132 118 118 118 118
INDEX
414
Balansia andropogonis 104 Balfour-Browne, Frances L. 99-4 Basidiomycetes 109 Boletus cervinus 127 Botrytis infestans . 100 Bovista bovistoides 135 Bovistella echinella 135 Bumelia punctata 177 Cabelleria ferruginea 175 Caeoma amaranthi 100 Cantharellus floccosus 120 Catacauma repens . 105 Cerebella andropogonis 3). ylAO
Ceylon, The genus Elaphoglossum i in 1 the Indian peninsula and Ceylon 81-96 Chromocyphella pale aca . 134 Ciboria firma ‘ IOI Cintractia caricis 109 Clavaria botrytoides 133 flaccida 134 obtusissima 134 suecica 134 Clavariaceae . 132 Clavariella suecica . 134 Clavulina alta 133 mussooriensis 132 Clitocybe tabescens 119 Coleosporium barclayense 111 campanulae 111 inulae 112 pedicularidis 112 plectranthi 112 senecionis . 112 Coltricia cinnamomea 2) a25 Columnea TOL, 247, Columnea affinis F : ; Seaicte. argentea . ‘ é : é 216, 225 aureo-nitens : ; ; : 189 brevipila_. ¢ : A . 216,226 calotricha 189 cubensis ‘ é : : . 189 fawcetti. : : ; : 216, 228 grvisebachiana 234 guianensis . : , : . 189 harrisii F E ; ‘ : 216, 224 hirsuta : ; : ? , 216, 226 hirsuta : : , 3 220, 228 hirsuta var. EORETIOR 227 var. fawcetti 228 var. genuina 227 var. pallescens 227 var. subintegra _ : : 224. 220 hispida. : : : . 216,219 hispida oa 12 ZO hunnewelli . ; é 2 234 jamaicensis P ‘ : : 215,217 proctorii . : ‘ : : 216/222 pubescens . Meee se
rutilans rutilans x ar baat
sanguinea var. cubensis
subcordata urbanii wullschlaegeliana Coniothyrina agaves Coniothyrium agaves Coprinus comatus . disseminatus Coriolus cervinus favoliporus hirsutus tephroleucus unicolor versicolor Coronophoraceae
Coronophora epistroma .
Corticium caeruleum
Crepidotus mollis .
Cyphellaceae
Cystopus amaranthi bliti .
Daedalea applanata betulina ; gibbosa palisotit sanguinea . unicolor
Dermateaceae
Diatrypacae .
Diatrype chlorosarca
Dichostereum durum
Dothidea osbeckhiae
Earliella corrugata .
Elaphoglossum in the Indian peninsula and
Ceylon
Elaphoglossum angulatum
ballardianum beddomei ceylanicum commutatum conforme fuscopunctatum krajinae latifolium . laurifolium marginatum nilgiricum ogatat spatulatum squamosum stelligerum stelligerum stigmatolepis stigmatolepis
216, 229 231
. 1389 216, 223 217, 220 . 225 136
136
120
120
127
126
125
126
127
127
107
107
129
120
134
100
100
128 128 128 128 124 127 103 104 104 132 107
124
81-96 83
INDEX 415
viscosum . ; : ; ‘ : 92 yunnanense : 5 : f SPICY Epichloe cinerea. : : : - 105 Epicoccum andropogonis : 3 . 140 nigrum : : 2 : : . 140 purpurascens. : : : . 140 Eranthemum F F : i a 207; Eranthemum aciculare . : : 2 312 armatum . : ; : : a. 301 emarginatum : : , : . 304 lycioides . : 3 : ; . 307 microphyllum . é : : * 3307 spinosum . : : : : . 304 vincoides . : : : : Sees iG: EXELL, A. W. : : : » 327-411 Exidia polytricha . : ; é . «118 Exobasidiaceae : 3 : 3 2 FIO Exobasidium s : : : ~ 19 Farysia olivacea_ . : : ‘ ol male, Fenestella fenestrata : ; : . 107 princeps . 107
Fernando Po, Angiosperms of the islands of the Gulf of Guinea (Fernando Po., Principe, S. Tomé, and Annobon) 327-411
Fomes marginatus : : : AYR! pectinatus . , : : ; . 123 Forsythiopsis : ; : : 207) Forsythiopsis australis . : : eer gLs baronit ; : : : : a ss linifolia . ‘ 2 ; ; ~ 320 minor ; : . : P 7 3a0 vincoides . : ‘ : ; 3) Sh5 var. meridionalis . ; : 578379 var. minor F : : ; - 320 var. vestita . : : : 319, 320 Frommea duchesneae_ . : : - I3 obtusa-duchesneae , : é ath obtusa [var.] duchesneae : ; Ema ABLES) Fungi Imperfecti . : , 136 Fungi of recent Nepal expeditions 99-141 Ganoderma applanatum : : . 122 lucidum . : : : : eu hee Geastrum fimbriatum . : : 2: 2235 harlotii . ‘ F : ; . 136 Geaster fimbriatus . : 135 Gentianaceae, New or little known Himalayan species of Swertia and Veratrilla 239-258 Geoglossaceae ‘ : : : i LOL Geoglossum affine . ; : 1 AOL Gesneriaceae, The Jamaican species of Columnea and Alloplectus 181-236 Gloeoglossum affine : ‘ : IOI Gloeophyllum subferrugineum . : Leo Gomphus floccosus ; : ; . 120 Grifola sulphurea . : 3 : ne ia
Guepina helvelloides : : : Se bk,
Guinea, Angiosperms of the islands of the Gulf of Guinea (Fernando Po, Principe, S. Tomé,
and Annobon) 327-411 Gymnosporangium cunninghamianum . 114 padmarense : : ; : . 114 Gyrocephalus rufus ‘ ; : = L19 Helvella crispa : : é : <7 LOR Helvellaceae . : : : ‘ = LOT Hericium erinaceus F : é - 122 Hesperanthemum . ; : oo 297) Hesperanthemum aciculare : : o2 SEZ armatum . : . : 2 a EOL microphyllum . é : ; BRE: purpurascens 3 d 3 5 - 313 spinosum . : : : . . 305 tetvastichum F 310
Himalaya, New or little known Himalayan
species of Swertia and Veratrilla 239-258 Hirneola polytricha ; : : 118 Humariaceae : : ; 4, LOX Hyalopsora polypodii : : , . 113 Hyaloscyphaceae . ; é : 2 102 Hydnaceae . 22 Hydnellum zonatum forma vespertilio . 122 Hydnum evinaceus : . F weyhe2
vespertilio . : : ; - sh22 Hymenochaete mougeotii : ‘ . 129 rheicolor . ; : F : . 129 rubiginosa . : : : ‘ . 129 sallei : : ; : : L290 tabacina . : . : ; . 130 tenuissima . : ; ; ‘. e © 120 vinosa y ; : : ; ; £30 Hypocreaceae : : ; : - 104 Hypoxylon mellissit , ‘ : : £06 multiforme : ; . ; 2 105 truncatum ‘ : : . . 105 Icacorea : : : : - 164 Icacorea compressa : : : ve LO4 solanacea . . 165 India, The genus Elaphoglossum in the Indian peninsula and Ceylon : 5 81-96
Jamaica, The Jamaican species of Columnea
and Alloplectus 181-236 Jasminum coeruleum : é ; 305 Justicia acicularis . é , : Giz
armata : : . 4 : . 301 microphylla : : : : 307, spinosa. : ‘ , : . 304 vincoides . : F : ; eagi5 Kingdon-Wardia HI Ce a ; 7. 1253 Kuhneola duchesneae ; : or ELS
416
Laccaria laccata Lactarius pubescens Laetiporus sulphureus Laxitextum crassum
vosea-carneum Lentaria macrospora Lentinus kurzianus
polychrous .
tigrinus
vellereus Lenzites applanata
betulina
palisotii
vepanda
subferruginea Lepiota erminea Leptostromataceae Lopharia crassa Lycoperdaceae Lycoperdon echinella
pyriforme .
Marasmius crinis-equi equicrinis .
Mastigonetron americanum :
fuscum Melampsora hirculi Melampsoraceae Melanconiaceae Melanotaenium eelnginellae Melasmia salicina . Microporus affinis .
perula Milesina exigua Mitrula rosea Mollisia dhankutae . Monochaetia liquidambaris Morchella elata Morchellaceae Mortey, B. D. : Mycenastrum bovistoides Myrsine . Myrsine acrantha .
coriacea
coriacea
ferruginea .
floridana
laeta .
punctata .
Nectrioidaceae
INDEX
120 acuminata
120 armata
124 var. armata .
130 var. pallidor .
131 grandiflora
133 jamaicensis
121 linifolia
120 microphylla
121 minor 4
121 var. meridionalis ;
128 var. minor
128 var. vestita
128 nannophylla
128 puberula .
128 purpurascens .
121 spinosa
137 tetrasticha
130 var. tetrasticha
135 var. polyece
135 vincoides .
136 Ostropaceae .
121 Panus polychrous
121 polychrous .
137 tigrinus
137 Parmularia discoidea
113 Peridermium orientale
III Peronosporaceae
137 Petrotia malemchiensis
111 Petesioides :
137 Petesioides laurifolium
125 subverticillatum .
125 Pestalotia americana
113 Peziza aurantia
101 firma ;
103 Phialea firma
37 Pholiota squarrosa
101 Phoma agaves
IOI Phragmidium incompletum
181-236 nepalense . 135 Phycomycetes 156, 174 Phylacteria caryophyllea 3) 75 Phlogiotis helvelloides 17745175 Phyllachora vepens
175 Phyllachoraceae :
175 Phytophthora infestans .
77, Pleosporaceae
175 Pickeringia :
177 Podosporium himalensis Polyporaceae é Polyporellus arcularius
136 picipes
Nepal, Fungi of recent Nepal expeditions 99-141
Olfersia angulata Ophelia racemosa Oplonia
Oplonia acicularis
83 253 297
298, 312
Polyporus affinis applanatus arcularius .
var. arcularius
var. strigosus . biformis cinnamomeus
208,
299, 314 299, 301 299, 301 299, 302 . 299 299, 300 298, 320 299, 307 ga 319
“a
EE
INDEX 417
corrugatus 2 ‘ . 124 Ramaria botrytoides 2 : : a. 33 fomentarius var. abplanatus : th OE flaccida. : ; , ; . 134 hirsutus . : : : : eS obtusissima : ; ; ; . 134 lucidus : : : : : Tes subaurantiaca . : ; ; . 134 marginatus é : : ; woe (123 suecica 5 : : ; . 134 pargamenus : : i ; . 123 Raphanea acrantha : : é we 7S pectinatus . : ‘ ; ‘ 123 coviacea . 9 P ‘ : cape perula : i z : d E25 ferruginea . ‘ : . 3 y ITS picipes ; = ; : : . 124 guianensis . ‘ : : , B/S yugosus. : 3 : : ee guyanensis : , : : eee) sanguineus : 2 : : ee 120 Rapunculus fruticosus... Sloane. ee ZZ squamosus : ; Z : . 124 Ravenelia emblicae : c : . 116 sulphureus é é : : . 124 Rehmiodothis osbeckiae : : On tabacinus . ; , : : 2 226 Rutstroemia firma : F : . 101 versatilis . ‘ ; ‘ ; Say, versicolor . . : : ‘ 5 aid) xanthopus . : : ; ‘ | 2S S. Tomé, Angiosperms of the islands of the Gulf zonalis ‘ . : : i - 124 of Guinea (Fernando Po, Principe, S. Tomé, Polystictus affinis . a, 3 d ce 45) and Annobon) : : . 327-411 cinnamomeus. : : 3 . 125 Samara coriacea. 3 3 : 175 hirsutus.. é : ; : . 125 Schizophyllum commune : : . 122 perula . : : : : - 125 Scleroderma aurantium . ‘ : 4 1S8 persooni . : : ; : = 24: Sclerodermateaceae 2 ; ‘ hee sanguineus : ; j : . 126 Sclerospora graminicola . : ; . 100 subaffinis . . ; : : . 126 Sclerotiniaceae ; . ; : ator tabacinus . : , 2 : . 126 Schneepia discoidea : é , . 108 tephroleucus : : : ; . 126 Seiridium liqguidambaris . : : ©, 1137. versatilis . : : , ‘ . 127 Sideroxylum punctatum . ; : i E77. versicolor . : : P : a 127, SLEDGE, W. A. ; ; : : . 81-96 xanthopus . é : ’ ; 5 eiZ5 SMITH: He |. ; : 239-258 Polystomellaceae 108 Sphacelotheca hydropiperis : : = 209 Principe, Angiosperms of the islands of the Gulf reiliana. : ‘ . 109 of Guinea (Fernando Po, Principe, S. Tomé, Sphaeria adscendens : ; : los and Annobon) ; ‘ . 327-411 deusta : : : ‘ : a LO? Protomyces graminicola . F : 100 multiformis : ‘ é : +, 105 Psathyrella disseminata . : A . 120 polymorpha : : : : . 106 Pseudotrametes gibbosa . : ; =, 128 repens ; : : 5 : a dLOS Psilanthele . : ‘ ; : . 296 telfaivtt : : : . . 106 Psilanthele eggersii 3 ‘ , . 296 truncata. ; ; : ; «(105 gvandifilora : : : F = 299 Sphaeropsidaceae . : : : es 130 jamaicensis ‘ Z : : . 300 STACE, C. A. : 3 . 3-78 minor 2 F P F 302 STEARN, W. T. , : a -236 261-323 Pterygoloma cristatum . : : e834 Stereum duricusculum . : 132 pubescens . : : : : « 232 elegantissimum . : : ; oat 20 vepens A 4 Fog et) fasciatum . : : : : ‘= 130 Puccinia caricis var. - himalayensis : . 114 hirsutum . : . : ; . 130 fagopyri_. : : : : ~ 15 insigne | ‘ F 3 ; 2 13 gentianae . Pn : ; : ees ostrea : . ‘ i : . 130 glumarum . : , : : . 115 vheicolor . : ; : : o 520 graminis . F ; : ; . 115 roseo-carneum . : : ; a MASI lantanae . 2 : F : ok sanguinolentum . : : : ceo leucophaea 5 4 , , wOLiS subpileatum ; : : : Sa! BL polygont . : : : £16 sulcatum . : ‘ ; ; Res) | polygoni- -aviculariae : : ‘ . 116 tabacinum . : : 2 : TSO pulverulenta : : ‘ : +2 216 tenuissimum : : : : HE ZO songarica . : ‘ ; : Tee bes umbrinum . : A ‘ : = 130 ustalis : ‘ : ; : SELIG Stilbaceae 72. : . 4 138 Pucciniaceae ; : . : 2 ES Stilbum cinnabarrinum : E : ESS Pycnoporus sanguineus . ; : ZO inconspicuum . 5 ; : us9
Pythiaceae . : : : : : | £00 kurzianum ; , ; . so
418
lateritium . Strilia cinnamomea Swertia . : Swertia sub. gen. Euswertia Swertia sect. Euswertia sect. Ophelia sect. Swertia sect. Veratrilla Swertia acaulis acaulis assamensis burmanica burmanica . burkilliana candelabrum crossoloma franchetiana gracilescens hookeri mekongensis pseudo-hookeri pseudo-hookeri vacemosa splendens staintonii staintonii stricta virescens vivescens
Thelephora caerulea caryophyllaea crassa fasciata hirsuta mougeotit ostvea voseo-cayrnea rubiginosa . sanguinolenta tabacina , ; versicolor var fasciata .
Thelephoraceae
Tilletiaceae :
Trametes betulina . cervina favolipora . gibbosa tephroleuca versatilis
Tremellaceae
Tremella helvelloides mesenterica rufa . :
Tuberculariaceae
Uredinales . Uredo bistortarum .
241,
241,
242, DAD 241,
242, 244i.
241,
INDEX
138 125 240 240 240 251 240 256 242 242 248 253 253 257 248 249 251 251 242 257 246 246 253 250 246 246 251 244 244
129 132 130 130 130 129 130 131 129 131 130 130 129 IIr 128 127 126 128 126 127 119g T19 119 119g 140
117 110
bliti . campanulae caricis : ; favinosa var. senecionis gentianae glumarum . hydropiperis hyperici-mysorensis inulae ; ; linearis var. polypodii olivacea veiliana . segetum var. hordei Ustilaginaceae Ustilago bistortarum hordei : morinae nuda nuda piperi reiliana ; segetum var. horde var. nuda Ustulina deusta vulgaris
Valsa fenestrata
Vararia rhodospora
Venturiaceae
Veratrilla
Veratrilla baillonii burkilliana burkilliana
Vibrissea truncorum
Wallenia :
Wallenia calyptrata clusioides . clusitfolia . corymbosa crassifolia . discolor elliptica erythrocarpa fawcettii grisebachii laurifolia laurifolia punctulata purdieana . subverticillata sylvestris . venosa Venosa xylosteoides
Xylaria arbuscula . hypoxylon f. tropica
156, 167, 166,
167, 167, 168, 166, 167, 166, 167, 167,
167, 167, 166, 168, 167,
167,
100 III 109 112 115 115 109 118 112 113 109 109 110 109 110 110 110 110 I10 111 109 110 110 107 107
107 132 107 256 257 257 257 104
165 173 168 173 169 169 174 168 171 168 172 173 172 171 172 168 174 171 169 171
106 105
INDEX
var. tropica . : F , . 105 Xylosphaera adscendens . mellissii. : : LOG hypoxylon subsp. adscendens polymorpha ; : . : = 306 mellissii telfairit : . ; petOO polymorpha
Xylariaceae . : : : : 7 metOs telfairii Xylobolus subpileatus . : : a RE
419
105 105 106 106 106
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i
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fittes
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ses ao =
saletrta peated yet eaa! Set rete 3 ‘ eeet pretties + sovsiossessgerstetiees ; icicb sashes get titsrstersssie : bebebsOtegeesecte! pectesectiesstestistenstes $333 saree ts seas sreeegeretss peseqeestsseet
“s pebesee tes iio teics \ destits sevestpuesestastistrir tle easeite
+ 33 ers
bistetsse
seittestisgite ; jeiricewest res se eetisecusee feicarsretsas siesstt
Seas sesieag east é
sietarsta se asas re sreestatstasstesesqecteea gett
sesso pic stenatatae tf adtaceesststestesenettstt
b ssiteiatcrasee teats
ro icetiteeuiesesseg ta tcsstetistrsrst3 Sp hF siptisssegasieea : rst
= eee! shitas athe siete i Sagsie: = Hagasisrstistitettetceysees tesa atbeneses bytes Betsiesette Bae Sieecaraseicos iiaeeuisteeeyiotees iotinauassereisisstiresseesesysetiabe trie-goteasters tots Seectsttersgs
tiie pestyeees
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state
tripee: eee per eesti oo cose shat eet Suapeneree Strasicareres