Fig. 2: Short-cells and hairs (a: nodular-cells; b: dumbbell-shaped cells; c: cross-cells; d: micro-hairs; e: prickle-hairs)
By URSULA SCHOLZ
Citation: Scholz, U. 1981. Monographie der gattung Oplismenus (Gramineae). Phanerogamarum monographiae Tomus XIII. J. Cramer, Vaduz, Germany. 217 pp. With 46 figures and 2 tables. Englush translation by Anthony McIntyre, Spencer Atkins, and Felix Tweraser.
Published by A.R Gantner Kommandit community, FL-9490 VADUZ; © 1981 A.R. Gantner Verlag K.G., FL-9490 Vaduz;
Printed in Germany by Strauss & Cramer GmbH, 6945 Hirschberg 2
ISBN 3-7682-1292-0
Table of contents
12.1 Description of genus
12.2 Key to the species
12.3 Conspectus
(O. aemulus, O. affinis; O. baronii, O. burmanii; O. compositus, O. flavicomus, O. gracillimus,O. humbertianus, O. hirtellus) [Order alphabetized here; not in presentation].
13. Taxonomic structure and geography of the genus – changes and discussion
13.1 General and introductory views
13.2 Over view of distributional areas
13.3 The species of Oplismenus sect. Scabriseta
13.4 The species of Oplismenus sect. Oplismenus
13.4.1 Oplismenus hirtellus
13.4.2 Oplismenus compositus
13.4.3 Oplismenus aemulus
13.5 Comparisons of the taxa in tropical and extra tropical regions
13.6 The disjunct distribution of Oplismenus hirtellus subsp. undulatifolius
14. Doubtful taxa
15. Excluded taxa
16. Acknowledgments
17. Summary
Comment from Barkworth
18. Literature cited
In the secondary literature, Oplismenus is frequently ignored. Estimates of the the number of species fluctuate between 10 and 40. Earlier, morphologic-systematic research of the genus was usually geographically restricted; they focused on specific areas of circulation. (E.g.. Hitchcock & Chase: America; Honda: Japan). Mez began a monographic study of the genus in the 1920s. However this study was never finished. It remained unpublished and is found as an unfinished manuscript in the Botanical Museum of Berlin-Dahlem.
The apparent lack of agreement among modern floristic-works, the strong polymorphism of the species and, last but not least, the disjunct subtropical region of some taxa prompted me to undertake a monographic study of the genus.
Early in my work, a study of Oplismenus by Davey & Clayton (1977) was published based on discriminant analyses. From the results it can be seen that a systematic organization of the taxa is very difficult and that, because of this, it is necessary to attempt a fundamental study of the plants from all geographical regions. Moreover, the varying circumscriptions and organization of the genus by earlier authors necessitated a thorough study of the synonyms.
In the relevant floristic works, six different species are repeatedly mentioned: Oplismenus undulatifolius for Europe and Asia, O. compositus for Asia, O. burmanii for Asia and Africa, O. hirtellus for Africa and America, O. rariflorus and O. setarius for America. The present work should analyze whether different species are being studied or if infra-specific taxa should be recognized. The most important characteristics of the wide-ranging taxa just mentioned will be studied and critically considered in relation to their taxonomic placement, evolutionary history, and geographical origin.
The present monographic study duplicates the above mentioned monograph by Mez in some points. Thisis especially true of the status changes that were recommended by Mez. A new organization of the genus Oplismenus will be presented, including a discussion of questions that remain concerning the taxonomy that could not be answere in this study.Within the scope of this work, approximately five thousand specimen documents were evaluated, from the following herbaria: A, B, Bern, Bish, BM, BR, C, CGE, E, F, FI, FR, G, GOET, HBG, K, L, LD, LE, M, MO, NY, OXF, P PNH, PR, PRC, TI, US, W, Z, ZT (Abbreviations, see Index herbariorum).
Special attention was paid to making certain that the the type specimens of all the names included in Oplismenus were located and studied. In total, 61 type specimens were discovered, such that most names could be clearly placed. Where type samples were not available for different reasons, special note was taken. In this case the placement relies either on the original description or on the statement of authors who have seen the type. This approach did not always permit a decision to be made; names that could not be placeed were placed under taxa dubia.
As far as the scale of specimen materials was concerned they were clearly different down to the individual taxa, despite the total number being so large. For the widely distributed species of Oplismenus compositus, O. burmannii and O. hirtellus, hundreds of specimens were available, while for the more narrowly distributed taxa there was only a minimal number of specimens.
The study was largely confined to the study of morphological attributes. Complete cytological or physiological analyses were not within its scope. Therefore, these kinds of studies from other authors could only be referenced. The measurements of individual organs were statistically evaluated. In most cases either mean values or primary value ranges could be defined; these included approx. 70% of all values for the character. Extreme values are generally found within parenthesis. The descriptions of the taxa are based almost entirely on my own observations. They include, however, some information from additional studies for the same taxa published by authors listed in the special section of this work. In addition, common names, economical impact, ecology, sociology and illustrative evidence were taken from the referenced sources.
The distribution of the taxa was verified in all important floristic works. The attached maps show the distribution based on specimens, As the majority of the herbarium specimens originate from the 19th century and the beginning of the 20th, it is not clear whether populations still exist at the locations shown. In literature, hints are occasionally found that allow a confining of the area.
The author had the opportunity to go on research excursions to Togo and to Switzerland in order to observe some taxa in their natural location, so that some observations concerning the life forms and the ecology as well as sociological statements could be used in this work.
The generic name Oplismenus was described by Palisot de Beauvois, Flore d’ Oware et de Benin 2: 14, pl. 68 (1810) with the nomenclatoral type of the genus, Oplismenus africanus. Most authors give 1807 as the publication date for the Flore d’ Oware et de Benin, as this date appears on the title page. From Stafleu (1967) it can be seen that the Flore d’ Oware et de Benin appeared in individual printings from 1804 until 1818 and that the section that applies to Oplismenus first appeared in September 1810.
Robert Brown, Prod. Fl. Nov. Holl: 194 (1810) described the genus under the name Orthopogon. According to Stafleu (1967), this work appeared in April 1810, therefore the name Orthopogon has priority over Oplismenus. Consequently, the need arose to either start using the seldom used name Orthopogon as a correction or else to conserve the generally well known name Oplismenus. Smith & Kerguelen (1976) suggested, on the grounds that the name Oplismenus is used by almost all authors, that the genus is widely spread in all tropical and subtropical areas and that all the species described by Orthopogon are also published under Oplismenus, that this name be conserved. In regard to the conservation of the genus name Oplismenus The Committee for Spermatophyta voted 12:0 for conservation (Taxon 27: 287; 1978). Hence Oplismenus is the correct name, by conservation.
Arduino (1764) described the taxon that is became known as Oplismenus undulatifolius, under the name “Paniculum undulatifolium”. The genus name Paniculum would have had priority over Oplismenus and Orthopogon, if it were valid. It can, however, be shown that this name is an an error, because the illustrations and type specimens that belong to it are correctly labeled as Panicum undulatifolium. Arduino also gives no indication that the described species can be separated from the genus Panicum.
Poiret (1816) inadvertently spelled Oplismenus as “Ophismenus”. Schlechtendal (1861-62) and Hasskarl, according to Chase (1911) nameed the genus “Hoplismenus”, derived from the Greek root of the word όπλίσμενος: the armed, so named due to the glume with awns. As this case revolves around an orthographic variation, the name Hoplismenus is taken to be a homonym of Oplismenus.
Steudel (1854) listed the generic name Hekaterosachne with one species H. elatior. The description itself does not indicate that it is Oplismenus. The type could not be studied, but various authors (Cheeseman 1906; Dalla Torre & Harms 1900) equate the genus with Oplismenus and the New Zealand species H. elatior with Oplismenus undulatifolius. According to my research O. undulatifolius is not found in the Australian region, so it must refer to O. compositus, O. hirtellus subsp. imbecillis or O. aemulus . In light of these facts the classification remains doubtful.
Rumpf (1750) coined the pre-Linnean name Hippogrostis amboina. This genus name does not, despite common belief, correspond to Oplismenus. Rumpf, Herb. Amb. VI: 14 (1750) described two species, namely H. major and H. minor. Both species differ only in size and are combined as Hippogrostis amboinica. Hippogrostis amboinica is depicted in table V, f.2 and corresponds neither in picture nor in description to Oplismenus. Merrill (1917) referred to the depicted plant as Ischaemum timorensum Kunth. In table V, f. 3 of Rumpf (1750), a species of Oplismenus-species is depicted; it corresponds to O. burmannii, so far as one can determine from a rather roughillustration of the habit. The accompanying text leads to the conclusion that a species is being discussed, which Rumpf labels as “eius denotat speciem”. Merrill certifies it as Oplismenus compositus, which according to his own data, is not included in Rumpf’s herbarium. Oplismenus burmannii, however, is present as evidence,but it doesn’t correspond to Merrill’s above mentioned depiction. The name Hippogrostis can however, under no circumstances be applied to the plant displayed in f.3. Nevertheless O. Kuntze (1891) needlessly picked up this name and changed the combinations of multiple species from Oplismenus to Hippagrostis (Hippogrostis).
The genus Oplismenus was not always unanimously defined. Palisot de Beauvois (1810) identified seven species in addition to the type species O. africanus. Persoon (1805) had previously put these together as two species groups: “Spic. composita, spicul. compressis secundis” . Valid combination changes of these species that Persoon quotes from Panicum, are first found in Palisot de Beauvois, Essai d’une nouvelle Agrostographie: 53 (1812). They are as follows: O. bromoides, O. burmannii, O. compositus, O. elatior, O. helvolus and O. hirtellus. In addition, the species O. foliaceus and O. undulatiflolius were named. The designation of “foliaceus” is clearly due to a typographical error in which O. loliaceus is named a synonym, as only Panicum loliaceum Lam. appears in the index of the Agrostrograph.
The case of O. undulatiflolius is more complicated. In text S. 54 O. undulatifolius – like the other speciesl – is listed as nomen nudum. In the Index s. 168 “Panicum undulatifolium And (Ard.)” is listed as a synonym for O. burmannii, and “Panicum undulatifolium ? L.” is listed as a synonym of O. undulatifolius. As no Panicum undulatifolium exists this combination is invalid (Niles & Chase 1925; Becherer 1929).
The type species of the genus O. africanus was described, illustrated, and nameed as a separate species next to Panicum hirtellum L. and Panicum loliaceum Lam. by Palisot de Beauvois. No voucher specimen is cited. Two specimens can, however, be studied, as they were well known to Palisot de Beauvois and have comments on them: “types de la Flora d’ Oware et de benin” (G) and “dedit Palisot de Beauvois” (LE). Both plants are similar in their habit (very delicate), they are however, relatively strongly differentiated in their inflorescence characteristics. The specimen 1 from Geneva corresponds to the depiction in the Flore d’oware et de Benin and should therefore be considered the lectotype. The specimen 2 from Leningrad is intermediate between O. hirtellus subsp. fasciculatus and subsp. setarius.
Like Palisot de Beauvois, R. Brown also tightly circumscribed his genus Orthopogon (Greek origin: όρθός straight, πώγων beard) and only compiled species under it that have awns in the outer three glumes and whose spikelets are pressed together from the sides. He lists Orthopogon compositus (= Panicum compositum L.) and three further species that he described, Orthopogon aemulus , Orthopogon flaccidus and Orthopogon imbecillis.
The genus Oplismenus is described with similar circumscriptions by Roemer & Schultes (1817), Raddi (1823), Nees von Esenbeck (1829 and 1841) , Bentham & Hooker (1883), Domin (1915), Hitchcock (1913 on following pages), Koidzumi (1925) and Honda (1924 and 1930). In contrast to this circumscription, which we consider to be Oplismenus s. str., are the interpretations of Kunth in Humbolt, Bonpland & Kunth (1816) and Kunth (1833), Desvaux (1831 and E. Fournier (1816) who sxpanded the genus to include the genus now known as Echinochloa as a section of Oplismenus. Even so Sprengel (1825) also recognized Orthopogon. Later however, like the earlier Poiret (1816) and after him Steudel (1854), Sprengel reduced Orthopogon to a section of Panicum, while Trinius, in earlier works (1820) accepted Orthopogon s. str. Mez (1917 and 1921) accepted Oplismenus (s. str.), but added to it some species that belong in different genera.
Schechtendal (1961-62) divided Oplismenus into two sections, base on characterisics of the awns. Species Oplismenus sect. Orthopogon (= sect. Oplismenus) have strong, red-gold, smooth awns, whereas members of Oplismenus sect. Scabrista have delicate, whitish, scabrous awns. This division appears sensible as the make-up of the awns is an important criterion, both in physiological as well as in dispersal.
Davey & Clayton (1977), in their study of some of the species of the genus, adopted new interpretations. They analyzed the species O. compositus, O. hirtellus, O. undulatifolius O. aemulus , O. imbecillis, O. rariflorus and O. setarius according using discriminate analysis (Cooley & Lohnes 1971). They compared all the species and attempted to separate them. They concluded that some species are easy to separate when one considers them within individual geographical regions, such as America, Africa, Asia, and Australia. When comparisons included specimens from multiple regions, however, they found some species to be non-separable species, O. hirtellus, O. compositus and O. undulatifolius. They concluded that there were no distinct boundaries between these three species.
The historical consideration of the genus proves clearly how difficult the separation of the species from one another is. This was made especially evident through the very objective research methods of Davey & Clayton. From that one may doubtless conclude that through a classical systematic approach no fully satisfying results can be expected.
Because t he systematic placement of Oplismenus was not one of the objectives of this study, only the basic approach of Pilger (1940), Hsu (1965) and Butzin (1969, 1970 a and b) are discussed. They placed the genus within the subfamily of Panicoideae, based on different criteria.
Oplismenus belongs to the subfamily of Panicoideae that in general, for example according to Pilger (1940), is characterized by havomg spikelets with two florets, one being reduced or staminate and the other bisexual. [It is the bisexual floret that is meant by the phrase "fertile floret"). Individual or paired spikelets are similar, either without awns or with a few simple awns. The lemma and palea of the fertile floret are commonly indurate. Handle-shaped silica cells and two-celled micro-hairs with smaller end-cells are often present.
Oplismenus is unusual among the predominantly awned tribe Panicoideae, but its inclusion in the subfamily is not disputed. Pilger placed Oplismenus in a group of awned genera that included Chaetium Nees, Stereochlaena Hackel, Achritchaete Pilger, Oryzidium C. E. Hubbard & Schweickerdt, Poecilostachys Hackel and Oplismenopsis (L.) Parodi.
Butzin (1969; 1970 a, b) attempted to clarify the classification of the Paniceae for which he gave the following diagnostic features: Awned glumes, leaves with evident transverse venation, placement of the hila, spikelet orientation, spikelet compression, and structure of the inflorescence. Through the preferential evaluation of these characteristic complexities, Oplismenus close to Acroceras Stapf, Streptostachys Desv., Poecilostachys Hackel and Chloachne Stapf. Clearly Oplismenus is different because of the relatively smooth but sticky awns that are are present in many species. As the species of the genus Oplismenus sect. Scabriseta have clearly dentate awns this characteristic cannot be used, by itself, for circumscribing the genus. The notched glumes that are especially noticeable in O. affinis offer an additional characteristic for circumscribing the genus because the only other awned species of Paniceae exhibiting this characteristic is Mesosetum Steud.
In studying the leaf venation of Paniceae, Butzin (1970 a) determined that there is evident transverse venation in Oplismenus. In many cases this proves true, but there are, however, many plants in which simple parallel venation exists. Consequently, this characteristic cannot be used ito distinguish Oplismenus from its relatives.
Prior to the above treatments, additional attempts had been made to subdivide Paniceae, not only according to morphological criteria, but also anatomical and cytological criteria. Hsu (1965) set up a new organization based on characteristics of the lemma epidermis of the lemma, the form of the lodicule and the structure of the base of the style. These charateristics place Oplismenus closest to Panicum L., Ichnanthus P. Beauv. and Echinochloa P. Beauv.
Oplismenus includes both perennial and annual species. As no special survival organs and no wooded parts are developed, the perennial taxa must be hemicryptophytes. In published accounts, O. flavicomus, O. gracillimus, O. compositus, O. hirtellus and O. undulatifolius are usually mentioned as perennial taxa. The annuals appear to be O. affinis and O. burmannii. I was able to check this for O. burmannii in Togo, where a specific site was researched during multiple seasons. It is difficult to differentiate morphologically between the annual and perennial species. In some cases stronger growth hints at a multi-year life-type.
In observing the morphology of the genus, it is necessary to determine which criteria are vital to the definition of the species and other taxa and which are irrelevant. It becomes evident that what is an important distinguishing for some groups of two or more taxa may be uniimportant for other groups.
The hairiness of the vegetative area cannot be used as a definitive characteristic to differentiate species or subspecies, as examples of varying hairiness appear in all taxa. The furrows, the culms, the upper and lower sides of leaves, the sheath (the edge as well as the flat part), and the ligule can all have hair. The length of the single-celled, undivided hairs (so-called macrohairs) varies from 0.5 to 5 mm. Shorter hairs appear mainly on the culm, on the tip of the ligule, and mixed with longer hairs on the leaf blade. They stand straight out from their base and have a wide base that gradually turns into a thin hair point. Longer hairs are found on the edge of the sheath, sometimes on the flat portion of the sheath, and on the leaf blade.
The density of the hair also varies greatly. Especially the edge of the sheath and occasionally the leaf blade can have dense hair, so dense that they appear velutinous. These hairs either stand up or are bent over. The cylindrical microhairs also need to be mentioned. They are present in almost all taxa on the lower and upper sides of the leaves. Noticeably dense hair on multiple parts of the plant is found in O. hirtellus and O. compositus. The leaf sheaths are, mostly, hairier on the nodes and the points than on other areas. A constant, dense hairiness of the flat portion of the sheath and also the inflorescence rachilla appears only in O. hirtellus subsp. undulatifolius, enabling this characteristic in combination with others to be used in distinguishing the taxon. The length of the hair growth of the leaf-ligule is variable within most taxa, only O. hirtellus subsp. capensis and some plants of O. hirtellus subsp. acuminatus and subsp. psilostachys have ligules that are glabrous on the surface.
The length of the leaves varies more or less greatly among all the taxa (between 2 and 16 cm); only O. hirtellus subsp. microphyllus is characterized by consistently short leaves.
The leaf shape varies greatly as well. Only O. hirtellus subsp. imbecillis is noticeably different in having small lanceolate leaves.
The leaf epidermis exhibits basically all the characteristics that are typical for Panicoideae, a group of predominantly tropical grasses. The most important taxa of the genus were studied, to determine whether the their characteristics could be used for a systematic organization and whether the tropical or extra-tropical morphological groups exhibit differences in the epidermis. Gross (1896) studied the epidermis of Oplismenus undulatifolius. Grob's system is the basis for the following treatment. All the parts of the epidermis, the upper parenchyma as well as the ribs, were studied on both the upper and lower side of the second leaf below the inflorescence. The leaf pieces were 0.5 X 0.5 cm; they were boiled and coated with nail polish. After lengthy drying the polish was carefully pulled off with tweezers and studied in a drop of water under the microscope. An exact imprint of all parts of the epidermis was there, only the longer hairs were broken, being apparent only as diffuse hair base. This method is a simple form of Sampson's (1961) procedure. On a total of 50 samples the shape and length of the stomates, the long-cells the short-cells, the other forms which includes silica cells, the microhairs and the prockles were studied. The occurrence of long- and short-cells as well as microhairs was considered separately for the parenchyma and vascular tissue.
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| Fig. 1 Overview of epidermis | Fig. 2: Short-cells and hairs (a: nodular-cells; b: dumbbell-shaped cells; c: cross-cells; d: micro-hairs; e: prickle-hairs) |
Many different shapes of silicacells were found, as well as, for tropical grasses, typical cylindrical, two-celled micro-hairs that lay on the epidermis with the base cell orthogonally broken off, were found (Fig. 2).
The epidermis can be divided into two areas. (Fig. 1): 1) The costal region over the vascular bundle; and 2) inter-costal regions over parenchyma tissue. The fundamental cells of the costal regions are elongate, small and strongly rippled and are present in all taxa. No other cells were found on either surface of O. hirtellus subsp. tschushimensis, or on the lower surface of O. hirtellus subsp. undulatifolius. In all other taxa, the row of fundamental-cells is interrupted by silicified short-cells. These short-cells can be dumbbell-shaped or nodular and may occur singly or in groups in an undisturbed order in one or two rows on the costa-field. Grob alludes especially to the rows of only silica-short-cells. Dumbbell-shaped cells can alternate irregularly with nodular cells. The nodular cells are symmetrically or irregularly spread. The length of the silica-short-cells ranges from 17.4 to 35.4 μm.
Occasionally cross-shaped silica cells and/or small pickle hairs are seen in the costal region. The prickle hairs are 29 to 95.7 μm long, thick walled and single-celled; however their form does not correspond to Gross (1896) hairs depicted in Abb. X, 52. A clear prickle point is seldom recognizable; mostly the hairs on the point are bluntly rounded off.
The area between two costa strips is referred to as the inter-costal region. Its fundamental cells are rectagular,, more or less cylindrical cells, whose walls are flat-edged or have a weak to moderate ripple. In extreme cases the cells have a length of 46.4 to 113.1 μm. In addition to the fundamental cells, there are short cells that are almost square shaped. Grob defines these as “short” long-cells as they neither anatomically nor chemically differ from remaining long-cells.
Stomates are located in a row along the edge of the inter-costal region, between the fundamental cells. They are found in varying numbers, commonly on the lower surface, less commonly on the upper surface. The stomates are usually oval but occasionally round; sometimes cylindrical-small shapes appear. They are (20.3)29-52.2 μm long and 14.3 to 31.9 μm wide. Grob states that one of the adjoining cells is commonly silicified. This could not be confirmed by the above study.
Like the apertures true short cells are found along the edge of the intercostal region. They have silicified walls and are of characteristic form. Occasionally silica-cells are also present elsewhere in the total intercostal region, where they are loosely spread among the long-cells. They are thinly spaced, mostly cross-shaped and have a diameter of 11.8 to 29 μm. On the lower surface of O. hirtellus subsp. tsushimensis, subsp. undulatifolius and O. affinis var. Iand on the upper surface of O. burmannii var. burmannii no silica cells are found. Prickle hairs appear sometimes in the intercostal region of O. hirtellus subsp. capensis (both surfacese), subsp. setarius (lower surface) and O. affinis var. humboldtianus. More commonly cylindrical, delicately walled angular-hairs can be observed, whose end cells in many cases can be lost in older leaves. Gross (1896) statement that angular hairs in the genus Oplismenus appear predominantly on the lower surface of the leaf, was not confirmed. They appeared equally often on the two surfaces.
When the epidermal structure of individual taxa is compared, no taxon characteristics appear. As shown in table 1, the distribution of each characteristic is so broad that there is overlap in all features. Therefore the characteristics of the leaf's epidermis can be given no systematic value. It also demonstrates that it is not even possible to distinguish and tropical and extra tropical forms. In O. hirtellus subsp. tsushimensis and subsp. undulatifolius, the silicate-cross-cells that are typical of tropical grasses are uncommon and slightly modified.
Voucher specimens for leaf studies.
The rachis, primary branches, secondary branches, spikelets, glumes and lemma may all be hairy. The hairs on the rachis and primary branches grow from the raised edges and can be long or short (up to 3 mm) and flexible. The spikelet's pedicel usually has a few long hairs that may extend to the middle of the spikelet. In some species (O. affinis and O. burmannii in the American region) there is an obvious augmentation of the pedicel hairs so that the thick or moderately hairy spikelets are even more densely hairy.
Furthermore the glumes and lower lemma vary individually in their hairiness. In most cases the hairs grow from the edge of the glumes with some coming from the rounded backs. Oplismenus affinis and O. burmannii var. lanatus have very strong and thick spikelet hairs. These hairs are as long as or longer than the glumes themselves. Especially noticeable is the ring shaped hair growth of the lemma of O. affinis, which is almost exclusively concentrated on the middle area and is not seen in any other species. Thick, but mostly short hair growth is sometimes found on the spikelets of O. aemulus var. aemulus and var. densiflorus, O. hirtellus subsp. imbecillis f. lanceolatus, O. hirtellus subsp. fasciculatus and O. compositus. The hairs are mostly appressed to the spikelets and are short. The second lemma and palea are always glabrous.
The absence or presence of dense inflorescence hair helps in distinguishing O. aemulus var. flaccidus from var. aemulus .
The inflorescence of the taxa is made up of multiple partial inflorescences that form a compound structure. Oplismenus hirtellus subspp. acuminatus, capensis, and psilostachys have fewer side branches than the other taxa. The secondary inflorescences in the other taxa are racemosely constructed; they are separated from one another and grow alternately out of the triangular rachis.
In general, the lateral branches are longest in the lower portion of the inflorescence and shooter to strongly reduced in the upper portion. The distances between successive lateral branches (rachis internodes) also become shorter towards the top of the inflorescences. Two exceptions to this are O. aemulus var. densiflorus and O. hirtellus subsp. tsushimensis in which there are few lateral branches and the rachis internodes are essentially the same length throughout the inflorescence, making the inflorescence appear compacted and clumped. The constancy of this feature makes it a useful characteristics for distinguising these taxa from the all the others.
Each spikelet pair includes one shortly pedicillate spikelet and one almost sessile spikelet. Schlechtendal (1861-62) found some patterns concerning the placement of the spikelet pairs on a side branch. In taxa with long side branches like O. compositus the spikelet pairs sit clearly behind one another and are usually even clearly separated from one another. Taxa with short side branches like O. hirtellus subsp. undulatifolius have the spikelet pairs in clusters. This is not always th case. For instance, O. hirtellus subsp. hirtellus, with spikelet pairs close together, and O. aemulus vars. aemulus and flaccidus, with well spaced spikelet pairs, have lateral branches that are almost the same length. The length of the lateral branches is not therefore decisive, rather the number of spikelet pairs per-side branch. Fig. 3 shows a diagram view of multiple longer and shorter side branches with the location of their spikelets. If a longer side branch has many spikelet pairs, the spikelet organization is thick with multiple rows, while a shorter side branch with fewer spikelets gives a different impression. Schlechtendal (1861-62) relates these differences to the partially or fully reduced spikelet of the pair. In reality, particularly in O. compositus, single spikelets can be reduced to a glume with a long awn or even reduced completely to awns. The organization of spikelets cannot, however, be attributed to this phenomenon alone, because when the spikelet pairs are not reduced they may be either densely or loosely packed on the lateral branches.
The partial or complete reduction of the awns on the first lemma, in addition to the organization of the spikelets, helps differentiate between O. hirtellus, O. aemulus and O. compositus.
The shortly pedicellate spikelets consist of two awned glumes, a lower floret that is sterile or staminate and usually has an awned lemma but no palea, and an upper floret that is enclosed by the upper lemma and palea.
The individual taxa exhibit great variation in spikelet length. Oplismenus gracillimus differs from all other taxa by having a spikelet length of 1.5-1.8 mm. Spikelet length, in combination with other characters, also serves to differentiate other pairs of taxa, for example O. burmannii (2)2.8-3.5 mm and O. flavicomus 3.8-4 mm; O. affinis var. affinis (3.3)3.5-4(4.5) mm and O. affinis var. humboldtianus 3-3.3 mm; O. compositus var. compositus (3.3)3.6-5.2 mm and O. compositus var. rariflorus (2.5)2.6-3.2 mm. Aside from these, some subspecies of O. hirtellus (subsp. acuminatus, subsp. capensis and subsp. psilotachys) can be separated from the subspecies with smaller spikelets using spikelet length in combination with other attributes.
Glume shapes varies within each taxon but is nearly always lanceolate. Oplismenus baronii and O. gracillimus differ from all the other taxa in having short, broad glumes. There is overlap in glume length relative to each other and the first lemma. In O. flavicomus, the second glume is clearly the longest of the three but in all other species the first lemma, or sometimes the first glume, is longest.
The number of veins in the glumes and lower lemma usually varies but the the two paleas always have two veins and the second lemma five veins. Only O. hirtellus subsp. undulatifolius is constant with respect the develops a constant number of veins (first glume 3 veins, second glume 5 veins, lower lemma 7 veins) which makes it possible, in combination with other attributes, to distinguish this taxon from all other taxa in the genus.
The differing structure of the awns is important attribute in distinguishing the two sections. Awn length works is a distinguishing characterisits only for O. aemulus var. densiflorus; for all other taxa, the range of variation is so great that this attribute is useless for taxon delimitation. The starting point of the awn, either below the the glume tip (as in O. humbertianus) or at the tip as an extension of the complete glume (in all other taxa) also separates both these taxa from all other taxa.
Observation of living plants, collected in Ticino, confirmed that the persistent, almost smooth awns secrete an adhesive substance when the fruit is ripe. This phenomenon, which is found only in Oplismenus among grasses, lead to a more detailed study of the awn anatomy and the chemical composition of the adhesive secretion (see sect. 7.3.2). The plants of O. hirtellus subsp. undulatifolius that were collected in Ticino and the awns of O. burmannii that were taken from herbarium specimens were used to study the phenomenon. The scabrous awns of O. burmannii have a smaller diameter than the smooth awns and are uniformly scleremacymetic. No clear layering was recognizable.
The awns of O. hirtellus subsp. undulatifolius were studied in early in development of the spikelet (Panel 1, Fig1) and when the fruits were mature (Panel 1, Fig. 2). The sections show a strong anatomical difference of the awn tissue. Three zones can be determined: 1. The outer epidermal layer: a layer of ordered palisade cells with thick outer walls, thin side and back walls. The cells are plasmatic. In the older cells, the plasma is concentrated in the outer section of the cell, and the core is noticeably larger than in younger awns. 2. Adjacent to the epidermis is a multi-rowed layer of thick walled, sclerenchymatized cells; on the cell walls deposits of cellulose are found. 3. The inside of the awn contains xylem cells of the vascular bundle with a large lumen. The profiles of the awns were inspected for lignin with phloroglucinol-hydrochloric acid. The xylem part turned red, all other cell walls remained uncolored.
The composition of the awns shows that the adhesive secretion is created in the epidermis layer and is secreted from there, but no pores could be seen in the cross sections. For comparison, photos of the outer surface were taken with a raster electron microscope (Panel 1, fig.3.) and (Panel 2, fig.1.-4.). It was determined that the awns upper surface was uneven, but uniform in its cell structure. The enlarged black spots in Panel 2, Fig. 4. cannot be interpreted as exit holes. Aside from the normal form of the awn in Panel 2, Fig.1. – 4. are, occasionally, awns that separated into 2 to 3 small teeth (Panel 1, Fig. 3.).
As no openings or clearly defined raised cell can be found, it must be determined that the adhesive secretion is secreted through thin spots of the outer epidermal layer.
After three months of drying, ripe awns of O. hirtellus subsp. undulatifolius were examined chemically to determine the structure of the adhesive secretion. The adhesive characteristics remained present, even after extended drying of the material. For very old samples, however, no adhesive awns could be observed. The fact that they had produced it was, however, indicated by the presence dirt stuck to the specimen. As the adhesive characteristic is at least partially present aftwer washing in water, it cannot be a glycoside or polysaccharide.
The solubility of the substance in different media was studied. It was determined that the adhesive substance did not dissolve with water. It dissolves well in diethyl ether and petroleum benzene and even better in ethanol, which indicates that it is most soluble in organic, polar dissolvers. It is, therefore, not a saccharide, polysaccharide, or mucopolysaccharide. It must be a lipophilic substance.
The possibility that the secretion was a terpenoid was examined next. For this purpose the following preliminary tests were be performed: 1. A drop of concentrated sulfate was applied to the awn. The substance on the upper surface of the awn took on a red-brown coloring. This reaction occurs among others in the presence of terpinene or steroids. Also the microtome cross section was stained with concentrated sulfate. The adhesive substance took on a red coloring but cells were not colored. This means that the secretioin is not present within the awn. To compare, sulfate was added to the DC made extract concentrate of the substance. No red coloring developed. 2. The reaction from Liebermann – Burchard with terpenoid was performed. The sample should turn olive-green for a positive reaction. A very faint tinge of green-brown developed. The reaction yielded no more definite results when repeated.
3. The reaction from Brieskorn and Briner with triterpene and sterine was performed. The sample should, for a positive result, turn red with triterpene and brown with sterine. It turned a faint shade of red-brown. This reaction was also inconclusive.
All reactions with terpinene or steroids occurred with only partially positive results. Placing this adhesive secretion in this elemental group can therefore only be done with great reservation.
Studies with help of the thin-layer-chromatography:
Thin-layer-chromatography offers the possibility of determining whether the adhesive secretion is a single substance or a combination of substances.
For this purpose the awns of the lowest glume were separated and approx. 100 every 15 sec. were swung in petroleum benzene. Thereafter the sample was compressed. The DC-study was performed with DC-silica gel plates (product: 60f – 254).
1. Attempt:
Starting point (1.5 cm): 10, 20, 40 substance pad. Isolating distance: 10 cm. Eluent: a: petroleum benzene 80%, diethyl ether 20%. b: petroleum benzene 80 %, diethyl ether 15%, glacial acetic acid 5%. Spraying medium: sulfuric acid-methanol mixture; carbonization at 130° C.
Analysis: For three starting points of the eluent a three spots with the following Rf-values can be differentiated: Rf 1: 6.75 Rf 2: 8.6 Rf 3: 9.75
For eluent b five spots can be defined: Rf 1: 0.5 Rf 2: 3.5 Rf 3: 3.75 Rf 4: 9.5 Rf 5: 9.8.
This study shows, with certainty, that the adhesive liquid of the awns is a heterogeneous substance with at least five components.
2. Attempt:
Starting point (1.5 cm): The thickened dried substance is re-dissolved with a small amount of petroleum benzene and applied in concentration. Isolating distance: 10 cm. Eluent: c: petroleum benzene 80%, diethyl ether 15%, ethanol abs. 5%. d: petroleum benzene 80 %, diethyl ether 10%, ethanol abs. 5%, glacial acetic acid 5%. Spraying medium: sulfuric acid-methanol mixture; carbonization by 130° C.
Analysis: In this attempt eluents with a higher polarity were used. The separation of the substance is hereby unclear. For Eluent c four spots with the following Rf-values can be determined: Rf 1: 3.6 Rf 2: 4.15 Rf 3: 9.45 Rf 4: 9.65.
For the attempt with eluent d, five definable spots with similar Rf-values can be defined: Rf 1: 4.75 Rf 2: 5 Rf 3: 5.6 Rf 4: 9 Rf 5: 9.65.
3. Attempt:
This attempt is similar to attempt 2, however, only eluent d was used. Spraying medium: anisaldehyde 1 part, methanol 17 parts, sulfuric acid 2 parts. After spraying the plate was dried at 100° C and analyzed under UV-light.
Analysis: Five possibly even seven spots could be located. Both lower spots were colored red (Rf 1: 0.3 Rf 2: 0.55). The upper spots were Blue-gray. – In the primary florescence; spots 3, 4 and 5 appear with a red tail. They were blotted out at 254 nm UV, which indicates the presence of phenolical groups. In the secondary florescence; spots 3, 4 and 5 appear gray-brown with a yellow-brown tail.
From all of these attempts it can be determined that the adhesive secretion is a heterogeneous lipphilic substance. Some results suggest that it is a terpenoid, others results cast doubt on this conclusion, so the question of what the substance is must remain open.
In this context the work of Ohmoto (1967) can be cited. He researched triterpene in Gramineae. His research, unlike this analysis, included all the herbal plants. From Oplismenus he isolated cylindrin, isoarborinol, friedelin, campesterol, sitosterol and stigmasterol.
The florets of the spikelets are made of two oblong or wide-obovate frail lodicules that reach a length of 0.4 mm (Fig. 4a). Three stamens are present; they have an oblong-straight shape and are yellow. The ovary is oblong and carries on its tip a divided long style, which phases out into two incidental, purple-red, feathery branched stubs.
The caryopsis is oblong oval and approx. 2 to 2.5 mm long. The hilum is narrow and approimately. 1/3 the length of the caryopsis (Fig. 4 b).
The stamens of Oplismenus are only pushed half way out of the spikelets when ripe and they open longitudinally. The pollen grains are yellow, mostly polygonal due to meiosis disruptions and are flattened on the sides, seldom round. Their cross-sectional dimension varies between 26.1 and 43.5 μm. Different sized grains were found, even within a single anther. It is probable that such pollen sterile; that can, however, be determined with fertility tests from the Maneval-Fuchsin-solution. The above mentioned details make it conceivable that reproduction is often apomictic. The exceptions were confirmed by Chiguryaeva (1976), who describes the correlation between apomixis and polymorphism, as well as between apomixis and polyploidy. Both phenomena are present in Oplismenus.
Scnarf (1931) and Shadowski (1926) performed embryological studies on O. undulatifolius. These showed that the erect embryo sac next to the egg apparatus has, remarkably, two pole cores and an antipodal apparatus of 10 to 18 cells, which form for pollination and separates from the rest of the embryo sac. Many of these cells have two nuclei, their plasma is heavily vacuolized. After pollination the entire antipodal system dies.
8.1 Photoperiodism
A study by Mathon (1970) showed that O. hirtellus has an ambiphotoperiodic reaction of O. hirtellus. The, until now, seldom studied phenomenon of ambiphotoperiodisim states that some plants develop especially well under extreme photoperiodical conditions, namely under short days- (10 hrs.), or under continuous light (24 hrs.), which is the case for O. hirtellus. In contrast under intermediate conditions (16 hrs. of light) inflorescence development was heavily delayed.
Kirchner et al. (1908-12) describe O. undulatifolius as a typical shade plant that sets its leaves perpendicular to the rays of light.
Hofstra (1972) and Smith & Brown (1973) determined that Oplismenus has C-3 type of photosynthesis, and determined a clear dependence on the ecology of the plants. The C-3 type, which manifests itself anatomically through the lack of chloroplasts in the vascular bundle-sheath of the grass blades, appears in the subfamily of Panicoideae only in shade plants and appears to indicate the intermediate position of these plants between the purely tropical taxa, that belong to the C-4 type, and the taxa of other subfamilies that grow predominantly in temperate climates. This is especially notable considering the tropical-sub-tropical-Mediterranean expansion of the genus Oplismenus.
The species of the genus Oplismenus are almost exclusively forest grasses that are found either in dense or moderate shade. Open grassland or places with lots of sun are rarely populated. In contrast forests along rivers or streams and swampy shaded areas are favored. Only O. burmannii is occasionally found in locations with more sun.
Many taxa are commonly encountered as weeds in shady coffee or coco plantations. Their effects as a weed must, however, be relatively minimal. Only Ponert (1977) mentions that the weed O. burmannii, which was introduced in Caucasus, was carefully exterminated to prevent its expansion.
The complete spikelet with both glumes with awns, the sterile lemma with short awns and the fertile lemma and palea that go hard when the fruit is ripe, serves as a diaspore. The awns of the spikelet play a major role in the epizoochoric dispersal of the fruit.
The awns of the species from Oplismenus sect. Scabriseta function through mechanical anchoring of the dentate awn, a mechanism that is seen often in the Poaceae. The awns of the specie of Oplismenus sect. Oplismenus function through a principle of adhesive shoots. The adhesive effects were studied on masculine hairs, on skin and on sheep's wool and were present in all three cases.
9. Chromosome count and Hybrids
For the widely distributed taxa, chromosome counts were found from different authors in the sixties and seventies. The base number is considered to be x=9. For the Asiatic forms of O. burmannii Sharma & Hevi (1959) reported 2n=18, for O. compositus Mehre & Chaudhary (1974) obtained n=9. Diploid plants appear, however, to be in the minority; all the other published numbers point clearly to olyploidy.
For O. burmannii, tetraploid, hexaploid and octoploid plants are known (2n=36, Mehra & Sharma 1973; 2n=54, Koshla & Mehra 1973; 2n=27, Kammacher et al. 1973). This was true of both Asiatic and African material.
Kammacher et al. (1973) attempted to show that octoploid plants of for O. hirtellus developed more robustly than hexaploid plants. They studied multiple plants from the Ivory Coast and determined that they have three giant-forms with 2n=72 and one dwarf-form with 2n=54. I examined their voucher specimens (Adjanohoun 907, 942, 943, 946). Based on my examination, all four plants belong to O. hirtellus subsp. hirtellus, i. e. that they have long side shoots and one of the spikelet organizations typical of that subspecies. This leaves open the fundamental question as to whether O. hirtellus subsp. hirtellus is just one of the polyploid forms of the frail O. hirtellus subsp. fasciculatus. As no live plants were available for cytological research, the size of the pollen grains of the robust plants and the dwarf form studied Kammachers et al. and some additional especially frail specimens of O. hirtellus subsp. fasciculatus were studied. The size of the pollen grain varied between 31.9 and 34.8 μm for all the plants studied. IN other words, there was not strong correlation between pollen size and the robustness or otherwise of the plants.
Notable connections for O. compositus are pointed out by Mehra & Sharma (1977). The authors indicate a hybrid population O. compositus x O. burmannii with a chromosome count of 2n=54. In my opinion the number 54 alone is not proof of successful hybridization, as it is also mentioned in many studies from other authors. A study of the morphological characteristics given by Mehra & Sharma revealed, however, that the relevant plant was clearly intermediate. The arrangement of the awns played a major role here and such plants with intermediary awns are seen in the hybrid species O. xthwaitesii. The explanation suggested by Mehra & Sharma that hybrids are among these was also confirmed through further cytological studies. The pollen was up to 70% sterile, the organization of chromosomes was not homologous. Missing chromosomes were definable in the daughter generation.
In conclusion it can be said that Oplismenus is mostly polyploid. The polyploids do not, however, appear to have larger pollen grains or a more robust habit than diploid plants.
In central Europe taxa Oplismenus are occasionally kept as house plants, as they appear very decorative due to the sometimes heavily rippled leaves. Horticultural forms with green-white long-striped leaves were cultivated. In van Houtte, Flores des Serres 17:5 (1867-68) an example of this cultivated form is depicted.
Most of the cultivated material available to the author belongs taxonomically to Oplismenus sect. Oplismenus, specifically to the O. hirtellus and O. aemulus . A more specific placement is impossible due to the possible reduction of the form by cultivating for specific attributes.
Occasionally also plants of Oplismenus sect. Scabriseta are cultivated. A form of O. burmannii with striped leaves originated in Kew and was placed in literature under the name O. burmannii var. albidum N.E.Br.
Schlechtendal (1851) described O. gracilis as species of unknown origin found in gardens. He placed it in Oplismenus sect. Scabriseta and distinguished and said that it differed from all other taxa in having a single vein in the lower glume and 3 veins in the lower lemma. As the type was not available, the position of the species could not be determined. It is included under Taxa dubia.
11. Studies of Attribute Correlations
In this portion of the study, I examined the correlation between various pairs of attributes. The goal was to eventually use this method to create more criteria for taxon definitions. Through graphical representations it will be shown how large the region of variation is for individual attributes and the extent to which such relations are useful.
11.1 Leaf Length / Spikelet Length
In the Figs. 5- 8, leaf length is show in relation to spikelet length for three taxa. Figure 5 diagram represents O. hirtellus subsp. fasciculatus, subsp. hirtellus and O. compositus var. compositus, which are taxa with short, medium length and long side shoots. The distribution of values is large for all three taxa; a correlation of the two attributes is clearly definable for O. hirtellus subsp. hirtellus. The crosses mark the approximate center of the statistical distribution.
Fig. 6 represents the taxa O. hirtellus subsp. setarius, O. compositus var. compositus and var. rariflorus. Attribute correlations are not available. However, this combination allows a more or less clean separation of O. compositus var. compositus and O. hirtellus subsp. setarius and of O. compositus var. compositus and var. rariflorus.
Fig. 7 represents three taxa with short side shoots. While O. hirtellus subsp. setarius and subsp. fasciculatus are relatively well separated, O. hirtellus subsp. undulatifolius cannot be separated from either of the other taxa through this attribute correlation.
The taxa in the Fig. 8 diagram have an even greater amount of overlap; even their statistical centers are close.
From this set of studies it is clear that the chosen attributes are, in general, not able to separate the taxa by themselves. Only in combination with other attributes does their value in this regard begin to increase.
11.2 Length of the inflorescence branches/ Spikelet Length
In a second attempt the attributes “length of the inflorescence branches” and “length of the spikelets” were studied in relation to each another. Naturally only the taxa that have well developed side shoots are considered in this section. In the Fig. 9 diagram O. hirtellus subsp. fasciculatus, subsp. hirtellus and O. compositus var. compositus were compared, while the Fig. 10 diagram compared O. aemulus , O. compositus var. compositus and var. rariflorus. There is no significant correlation between the two attributes. Nevertheless, the graphical depiction makes it clear that the individual attributes (especially the length of the side shoots) has a broader distribution than other attributes.
Through combinations of attributes all taxa can be more or less well separated, which boils down to the key determining factors.
In connection with the leaf anatomy a further attempt is made in the following section to relate attributes and discover correlations. Even when such microscopic attributes do not qualify for use in a key, they offer the possibility of clarifying the related structures.
11.3.Length of Stomates/ Leaf Length
Leaf length alone cannot distinguish the taxa. There are some taxa, however, that can be grouped together based on their similar leaf lengths. The mean value of the stomate length should be related to this. Taxa with short leaves (2 to 6 cm long) are represented in Fig. 11 group 1. The mean length of the stomates is 38.3 to 41.5 μm. Group 2 covers taxa with medium length leaves (4 – 11 cm) and medium stomates of 38.9 to 44.9 μm. The values are almost completely overlapping. Oplismenus hirtellus subsp. fasciculatus has the highest value (44.9 μm) and tends already toward the third group, in which the taxa with large leaves (6 to 14 cm) are combined. Here the mean values for the stomates are 43.5 to 46.4 μm. With that can be pointed out a clear enlargement of the aperture to taxa with longer leaves.
Oplismenus P. Beauv., FL. D'Oware et de Benin 2: 14 (1810, t.p. 1807), nom. cons.; Orthopogon, R. Br., Fl. Nov. Holl. 194 (1810); Hippagrostis Rumpf, Herb. Amboin. 6: 14, t.5 (1750) ex. O. Kuntze, Rev. Gen. 777 (1891); Paniculum Ard., Anim. Bot. Sp. Alt. 14 (1764), err. Orthogr. “Panicum”
Plants perennial or annual, procumbent and thick carpet forming grasses, inflorescences borne above the leavesfloret stems. Roots fine, few and branching. Culm 15-105 cm tall, round in cross-section, lightly grooved, glabrous or hairy; basal branching intravaginal; rooting at the nodes.
Leaf sheaths open, with overlapping edges, grooved, mostly hairy on edges and occasionally over the body; ligules truncate, strongly ciliate (0.2)0.4-1.5(2.5) mm long; blades ovate to lanceolate, acuminate, commonly more or less strongly rippled, base asymmetric; edges rough; lower leaves usually smaller than those above.
Inflorescence racemose or twice-racemose; branches triangular in cross-section, usually terminating in a single spikelet, hairy or glabrous, bearing alternating spikelet pairs in clumps or on 0.5 -10 cm long secondary branches. Secondary branches asymmetrically triangular, with alternating spikelet pairs on two surfaces, terminating in a single spikelet. Spikelet pairs either sparse, densely packed, or clumped, one spikelet subsessile, the other shortly pedicellate.
Spikelets 1.8-5.5 mm long, lower floret reduced or staminate, upper floret bisexual; glumes lanceolate, dentate, sometimes rounded; glumes lanceolate, dentate, the outer veins coalescing with the inner veins distally, then all the longitudinal veins coalesing with the well developed midvein, cross venation occasionaly evident, weak; lower glume 0.8-5 mm long, with 3-7 veins, veins coalescing distally, awned, awn 1.5-17 mm; upper glume 1-4.5 mm long, with 3-9 vein, veins coalescing distally; lower lemma 1.5-5.3 mm long, sometimes dentate, with 7-11 veins, sometimes awned, awn to 4.3 mm long; lower palea absent or weakly developed, truncate, and 2-veined. Lemma and palea of upper floret glossy and glabrous, enclosing the flower, hard when ripe.
Lodicules 2,, about 0.4 – 0.6 mm long, frail, elongated or broadly ovate and wide in the upper part, sometimes clearly bilabiate.stamens 3; ovaries elongated-ovular with two terminal styles and purple-red, feathery divided style branches.
Caryopsis glossy, tightly encircled by the upper lemma and palea; hilum elongated, about1/3 as long as the caryopsis.
Type species: Oplismenus africanus P. Beauv.
Species 9; 4 endemic in Madagascar, 1 new world, 1 confined to Oceania and Australia, 3 distributed in all tropical and partially subtropical areas of the world.
1a |
Awns frail, whitish, scabrous |
|
|---|---|---|
| 1b | Awns hardy, red-gold, bare, adhesive when ripe |
|
| 2a | Awns originate under the top of the glumes, glumes acuminate, or dentate. |
O. humbertianus |
| 2b | Awns terminal, glumes duel toothed or entire |
|
| 3a | Glumes notched on the tip, awns originating from the sinus; spikelets densely hairy |
O. affinis |
| 3b | Glumes entire on tip, awn terminal; spikelet thickly or fairly heavy hairy |
|
| 4a | Upper glume clearly longer than the lower glume and lower lemma |
O. flavicomus |
| 4b | Lower lemma longer than the glumes |
|
| 5a | Glumes oblong-lanceolate, clearly longer than wide | O. burmanii |
| 5b | Glumes oblong-round, almost as wide as long |
|
| 6a | Lowest side shoot up to 1.5 cm long; spikelets 2.7 – 3 mm long |
O. baronii |
| 6b | Lowest side shoot 5 – 6 cm long; spikelets 1.5 – 1.8 mm long |
O. gracillimus |
| 7a | Spikelet pairs not overlapping, sometimes distant; lower lemma usually unawned; secondary inflorescence branches (1.5) 3 – 10 cm long |
O. compositus |
| 7b | Spikelet pairs, clumped lengthwise or organized behind one another in multiple rows; secondary inflorescence branches absent or up to 3 cm long |
8 |
| 8a | Lower lemma shortly awned; spikelet pairs clumped or den on lateral branches; spikelets 2.2 – 5.6 mm long, if shorter than 3 mm, the spikelets always clumped |
O. hirtellus |
| 8b | Lower lemma unawned, sometimes shortly pointed; spikelet pairs thick or clearly imbricate but not distant; ; spikelets 2.4 – 3.2 mm long |
O. aemulus |
Oplismenus P. Beauv.
sect. Scabriseta Schlecht., Linnaea 31: 301 (1861-62).
O. affinis Schult.
--O. affinis Schult. var. affinis
--O. affinis var. humboldtianus U. Scholz
O. baronii Camus
O. burmannii (Retz.) P. Beauv.
--O. burmannii (Retz.) P. Beauv. var. burmannii
--O. burmannii var. lanatus (Buse) Backer
--O. burmannii var. multisetus (Hochst. ex. A. Rich.) U. Scholz
O. flavicomus Mez
O. gracillimus Mez
O. humbertianus Camus
sect. Oplismenus
O. aemulus
--O. aemulus (R. Br. ) Roem. & Schult. var. aemulus
--O. aemulus var. flaccidus (R. Br. ) Domin
--O. aemulus var. densiflorus U. Scholz
O. compositus (L..) P. Beauv.
--O. compositus (L..) P. Beauv. var. compositus
--O. compositus var. rariflorus (Presl) U. Scholz
--O. compositus var. sylvaticus (Lam.) U. Scholz
O. hirtellus (L.) P. Beauv.
--O. hirtellus (L.) P. Beauv. subsp. hirtellus
--O. hirtellus subsp. acuminatus (Nees) U. Scholz
--O. hirtellus (L.) P. Beauv. subsp. capensis (Hochst.) Mez ex U. Scholz
--O. hirtellus subsp. fasciculatus U. Scholz
--O. hirtellus subsp. imbecillis (R. Br.) U. Scholz
----O. hirtellus f. imbecillis
----O. hirtellus f. lanceolatus U. Scholz
--O. hirtellus subsp. japonicus (Steud.) U. Scholz
--O. hirtellus subsp. microphyllus (Honda) U. Scholz
--O. hirtellus subsp. psilostachys (Honda) U. Scholz
--O. hirtellus subsp. setarius (Lam.) Mez ex Ekman
--O. hirtellus subsp. tsushimensis (Honda) U. Scholz
--O. hirtellus subsp. undulatifolius (Ard.) U. Scholz
Oplismenus affinis Schult., Mant. II: 273 (1824). Panicum schultesii Steud., Nomencl. 2.ed. II: 263 (1841). - Type: "In insula Sancta Martha", Bertero 2579, M, Holotypus.
Panicum francoi Steud., Syn. Pl. Glum. I: 44 (1854). – Oplismenus mollissimus Hochst. Ex Steud., Syn Pl. Glum. I: 44 (1854), pro syn. – Type: “Oaxaca” Franco s.n., (W?); n.v.
Key to the varieties:
| 1a | Spikelets usually 3.5 -4 mm long, very hairy; lower lemma with a dense hairy collar in the middle; glumes with a rounded tip | var. affinis |
| 1b | Spikelets 3 – 3.3 mm long, glumes densely hairy; lower lemma usually only slightly hairy; glume tip usually acuminate | var. humboldtianus |
Oplismenus cristatus K.B. Presl, Rel. Haenk. I: 323 (1830); Lamson-Sribner, F., Studies on American grasses: 7 (1897). – Panicum cristatum (K. B. Presl) Steud., Syn. Pl. Glum: I: 46 (1854). Oplismenus burmannii (Retz.) P. Beauv. F. cristata (K. B. Presl) Hier. Ex Peter, Feddes Repert. Spec. Nov. Regni Veg. Beih. 40, 1 A: 222 (1938) – type: Mexico: Haenke s. n. (PR); Location “Luzon” indicated by an additional label.
Distribution of O. affinis |
 |
Annuals, frail or hardy grass; culm 15 to 50 cm high, bare or haired in the grooves. Leaf blades ovate-oblong to oblong-lanceolate, mostly densely airy, especially on the base, usually long rippled on the edge, (0.6) to 0.9 to 1.2 to (1.6) cm wide, (2) to 3 to 5 to (7) cm long. Sheath thin or thick long hairs around the edge, likewise haired or bare on the flat. Ligule has a hairy collar on the end, 0.6 to 1.1 mm long. Inflorescences racemose placed together, compact; rachis 3 – 7 cm long; side shoots upright, leaning over or distant. Rachis as well as side shoots thick and long hair growth; the hairs are 2 to 3 mm long. The length of the side shoots decrease from the bottom up, the lowest is 0.5 to 1.5 cm long and carries unilaterally 10 to 20 spikelets, in thick succession. Separation of side shoots is very short, inflorescences almost connected. Spikelets paired, mostly surrounded by a thick collar of long hair around the base. Spikelets oblong-lanceolate, (3.3) to 3.5 to 4 to (4.5) mm long, thick long hair, flattened or almost round on the back. Lower glume lanceolate, (2) to 2.2 to 3.3 to (3.6) mm long, (0.7) to 1 to (1.2) mm wide, very thick, long hairs on the edges and on the flats, (3) to 5 veins, deeply notched on the tip, dentate rounded; with awn. Upper glume lanceolate, 2.6 to 3.4 to (3.8) mm long, (0.8) to 0.9 to 1.2 to (1.4) mm wide, very thick and long hairs on the edges and on the flats, (5) to 7 veins, deeply notched on the tip, dentate rounded; with awn. Lower lemma oblong tapered distally , (3.1)3.3-3.8 to (3.9) mm long, (1.6)1.9-2 (2.3) mm wide, has a thick hairy collar in the middle region, 9 veins, little or no notch on the tip. Lower palea absent. Caryopsis pale or brownish, approx. 2 mm long. Awns frail, whitish and dentate; awn of Gluma I (7) to 8 to 15 to (17) mm long, of Gluma II (2.4) to 3.8 to 7 to (10) mm long and of Lemma I (0.4) to 0.9 to 1.6 to (2.4) mm long (Fig. 12).
Distribution: Tropical Regions of the New World.
Ecology: In shady woods.
Economic Impact: Weed in coffee plantations.
Anthesis: September to December.
Common names: Lacate (Mexico); Gram. De conejo (Central America).
Selected voucher specimens: Brazil: Conias and Maranho prov. (Martius s. n.; M); prov. Pianhy (Martius s. n.; M). - Columbia: Santa Marta (G). - Costa Rica: Altos "de Santa Barbara (H. Pittier 1078; M) Bomca (Tonduz 4466; M) Prov. Guanacaste (Jimenez 2727; NY); Nicoya (Tonduz 13758; G). – Dominican Republic: Manie1 de Ocoa (Türckheim 3610; PR). - Ecuador: west of Duran (Schimpff 1068; G); Guayaqui1 (Jameson 365; LE). – El Salvador: Dep. Sonsonate, Armenia (Standley 23486; NY). - Guatemala: Alta Vera Paz (Türckheim 473; G); Chi1ion near Majatenango (Bernoulli 468; G); Dep. Santa Rosa (Heyde & Lux 6276; G); Osuna (ZT); Tiquisati (N. Johansson s. n.; G). - Honduras: Dep. Morazan (J. G. Hawkes 1994; G). - Mexico: Chiapas, Mapastepec (Tateoka 1005; US); Guatu1co (Liebmann 375; US); State of Guerrero (Y. Mexia 8718; B); Chilpancingo (Conert s. n.; FR); Between Huixtla and Tapachula (W. Boege 1079; FR); La Correa (Lang1asse 440; G) Maria Madre (Ferris 5674; US); Sierra de Tonalo (Purpus 7412; US); Sonora (Scott Gentry 4820; US); Yucatan, Calotmul (G. F. Gaumer 2429; G); Izama1 (G. F. Gaumer 1038; G); San Anselmo (C. F. Millspaugh 2428; G). - Nicaragua: Grenadu (P. Levy 24; G) Managua (Zelaya 38; NY). - U.S.A.: Lower California [If this is a translation of Baja California, it is in Mexico], Miraflores (Brandegee 75; NY). - Venezuela: Caracas (Funck 418; G); Tovar (A. Fendler 1705; G).
var. humboldtianus U. Scholz, nom. nov.
Op1ismenus humboldtianus Nees, Agrost. Bras.: 264 (1829), nom. illeg. - Op1ismenus humboldtianus Nees var. genuinus E. Fourn., Mex. PI. 2: 37 (1886), nom. illeg. - Type: Brazil, Bahia: Nees s. n., M, Lectotypus.
Op1ismenus affinis K. B. Presl, Rel. Haenk. I: 323 (1830), nom. il1eg. - Oplismenus pres1ii Kunth, Enum. Pl. I: 141 (1833). - Type: Panama, Ganomenzes: Haenke s. n., PR, Holotypus.
Hippagrostis burmannii O. Kuntze, Rev. Gen.: 777 (1891), non Panicum burmannii Retz. - Supporting Source: Kuntze 2126; NY.
Orthopogon burmannii Spreng., Syst. I: 306 (1824, t.p .. 1825). non Panicum burmannii Retz. - Panicum 1appaceum Wi1ld. ex Spreng., Syst. I: 306 (1824, t.p. 1825), pro syn.
Frail plants, 18 to 25 cm high; Spikelets 3 to 3.3 mm long; Lower glume 2-2.4 mm long, 0.7 to 1 mm wide, 5 to 7 veins, deeply notched on the tip, pointed dentate; Gluma II 2.1 to 2.6 mm long, 1 to 1.2 mm wide, deeply notched on the tip, pointed or blunt dentate; Lemma I 2.9 to 3.2 mm long. 1.7 to 1.9 mm wide, Slightly notched on the tip. Glumae thick and long hair, Lemma I fairly heavy to little hair.
Distribution: Tropical regions of the New World.
Ecology: In Rainforests; in Deciduous Forests (Mexico).
Anthesis: September to January.
Selected Supporting Specimens: British Honduras: All Pines (W. A. Schipp 783; G). - Costarica: (Tonduz 1481; LE). – Dominican Republic: (Bertero s. n.; M). - Ecuador: Guayaquil (Spruce 6326; OXF). – El Salvador:: Dep. Santa Ana (Weberling 2681; M). - Guatemala: Dep. Jutipa (W. C. Shannon 3672; G). - Haiti: Vallee des Trois-Rivieres (E. L. Ekman 5103; G). - Mexico: Atoyac (Kerber 166; LE); Veracruz (J. A. Purpus 2893; M); Morelos, near Yautepec (C. G. Pringle 11330; B). - Panama: Culebra, Canal Zone (A. S. Hitchcock 421; G); Chiman (Lewis et al. 3254; NY); Chagres, Isthmus of Panama (A. Fendler 273; OXF); prov. Panama (Killip 4191; NY). - Venezuela: Prov. de Carabobo (J. Linden 1.559; G); Maracay (C. Vogl 121; M).
Oplismenus baronii Camus, Bull. Soc. Bot. France 75: 911 (1928). – Type: Madagascar: Baron 909, P, Holotypus.
Frail plants, 20 to 25 cm high; Culm bare or slightly hairy; Leaves oblong-lanceolate, 0.2 to 0.3 cm wide, 1.5 to 2.5 cm long, bare or slightly hairy; Sheath overall bare or slightly hairy around the edges. Ligule with a more or less heavily haired collar on the end, 0.4 to 0.7 mm long: inflorescence 3.5 to 4 cm long, with multiple side shoots. The lowest side shoot is 1.5 cm long and has 10 to 15 spikelets thickly behind one another. The rachis and the side shoots are slightly hairy. The distance between the lowest two side shoots is 0.5 cm. Spikelets in thick succession. Rachis as well as side shoots have little hair. The distance between the bottom two side shoots is 0.5 cm. Spikelets arranged in pairs, ovate-lanceolate, 2.7 to 3 mm long, little hair growth. Gluma I oblong-rounded, 1.3 to 1.6 mm long, 1.2 to 1.3 mm wide, 3 veins, with awn. Gluma II oblong-rounded, 1.8 to 2.2 mm long, 1.3 to 1.5 mm wide, 3 veins, with awn. Lemma I oblong-rounded, 2.4 to 3 mm long, 2.1 to 2.2 mm wide. Palea I narrow, 2 veins, frail. Awns frail, whitish and dentate; Awn of Gluma I 5 to 7 mm long, of Gluma II 0.8 to 1.5 mm long. The Lemma I is only pointed.
Distribution: Madagascar.
Supporting Source: Madagascar (Baron 909; P).
Oplismenus burmannii (Retz.) P. Beauv., Ess. Agrost.: 54 (1812); Bor, N.L. in Rechinger, K. H., Fl. Iranica: 484 (1970): Bosser, J., Gram. Pat. et Cult. Madag.: 353; Cooke, T., Fl. Pres. Bombay 2: 443 (1958); Gamble, J. S., Fl. Pres. Madras 3: 1232 (1967); Haines, H. H., Bot. of Bihar and Orissa 3: 1045 (1961); Hara, H., Fl. Eastern Himalay: 369 (1966); Komarov, V. L., Fl. U.S.S.R. 2: 34 (1934); Koorders, S. H., Exkursionsflora von Java 1: 137 (1911); Mitra, N. J., Flow. Pl. of Eastern Ind. 1: 197 (1958); Nakai, T., Fl. Koreana 2: 349 (1911); Prain, D., Bengal Plants 2: 883 (1963); Ridley, H. N., Fl. Malay Penins. 5: 221 (1925); Robyns, W., Fl. Agrost. Congo Belge & Ruanda-Urundi: 150 (1934); Roxburg., W., Fl. Ind. 1: 298 (1832); Stapf, 0., in Prain, D., Fl. Trop. Afr. 9: 636 (1934); Stewart, R. R., Fl. West Pakistan: 120 (1972); Tzvelev, N. N., Poaceae U.S.S.R.: 655 (1976).
| 1a | Spikelets glabrous; lowest inflorescence branches 2 cm long | var. multisetus |
| 1b | Spikelets hairy; lowest inflorescence branch usually only 1.5 cm long | 2 |
| 2a | Spikelets fairly densely hairy, hair short. | var.burmanii |
| 2b | Spikelets silvery, densely hairy with long hair. | var. lanatus |
Distribution: Tropical to subtropical regions of the Old and New World (Fig. 15).
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Panicum burmannii Retz., Obs. III: 10/1783). – Orthopogon burmannii (Retz.) R. Br., Prod. Fl. Nov. Holl.: 194 (1810). - Type: India, Madras: Konig s. n., LD, Holotypus; C. Isotypus.
Panicum bromoides Lam., Tab. Encyel. I: 170 (1791). - Oplismenus bromoides (Lam.) P. Beauv., Ess. Agrost.: 54 (1812). - Type: "Ile de France", Commerson s. n., C, HOlotypus; FI, Isotypus.
Hybrids: Oplismenus thwaitesii Hook., f. in Trim., Fl. Ceylon 5: 169 (1900). = O. burmannii x O. compositus.
Icones: Bosser, J., Graminees des paturages et des cultures a Madagascar: 353, f. 133 (1969). - Burmann, N. L., Fl. Ind.: 24, t. 13, f. 1 (1768) ("Panicum hirtellum"). - Koechlin, J., Fl. Gabon 5: 59 (1963). - Reichenbach, L., Ic Fl. Germ. I: t. 28, f. 1410 (1834). - Rose Innes, R., Manual of Ghana Grasses: 192, f. 62 (1977).
Annuals, mostly frail grass; culm (16) to 25 to 45 to (65) cm high, Usually hairy in multiple grooves and more so on the nodes. Leaves light green, smooth or lightly rippled, oblong - lanceolate or ovate-oblong, usually has long white hairs, 0.6 to 1.3 to (1.6) cm wide, (2) to 3 to 7 to (8) cm long. Middle vein is weak, side veins 2 to 3 on both sides; cross veins present. Sheath thin or thick long hairs around the edge, usually sparse long hairs or bare on the flat. Ligule usually has a hairy collar on the end, 0.6 to 1.2 to (1.8) mm long. Inflorescences racemose placed together; rachis (3) to 4 to 8 to (11) cm long, sometimes hairy; side shoots 3 to 10, stand upright at an angle to the rachis. The length of the side shoots decreases from the bottom up, the lowest is (0.2) to 0.5 to 1.5 to (3) cm and carries unilaterally (3) to 10 to 20 to (25) spikelets, in thick succession, they are in pairs, usually encircled at the base by a collar of long hair that almost reaches the length of the spikelets. Spikelets oblong-lanceolate, (2) to 2.8 to (3.5) mm usually fairly heavily haired, flattened or almost round on the back. Gluma I oblong-oval, (1.3) to 1.9 to (2.4) mm long, 0.7 to 0.9 to (1.2) mm wide, fairly heavy hair growth, 3 to 5 to (7) veins, with awn. Gluma II similar in shape, (1.5) to 2 to (2.3) mm long, (0.7) to 1.2 to (1.5) mm wide, fairly heavily haired, 5 to 7 veins, with awn. Lemma I oblong, (2) to 2.6 to (3.5) mm long, (1) to 1.7 to (2.4) mm wide, (5) to 7 to 9 to (11) veins, pointed or with a short awn. Palea I not present. Caryopsis oblong-lanceolate, convex on the back, almost flat on the front. Awns frail, whitish and dentate; awn of Gluma I (3) to 6 to 11 to (15) mm long, of Gluma II (0.4) to 3 to 6 to (9) mm long; Lemma I has either a short point or a short awn - 0.6 to 2 to (2.3) mm (Fig. 14).
Ecology: In shady, half shady and open forests; forests along rivers, dry forests and thorn patches, occasionally in the savannah between shaded rock outcroppings and crops; sandy soil on basalt.
Sociology: Under Palms, Acacia campylocantha, Tamarindus indica, Trichilia emetica, Eucalyptus, Ficus, Bridelia macrantha, Mango. Companion plants: Impatiens, Pilea, Ponzolchia arachoindea.
Economic Impact: Weed in coffee, cocoa, banana and kapok plantations, in peanut and corn fields. Removal resource: 0.4 % Perepod (fungicide).
Anthesis:In tropical regions: Usually August to December or March to May. – In subtropical regions: September to November.
Pests: Tilletia vittata (Berk.) Mundkur, Tilletia vittata var. burmannii Mishra; Ustillaginoidea oplismeni Ramakrishnan & Sundaram; Dub.: Ustilago oplimeni Viennot-Bourgin
Chromosomes:
2n = 18: Koshla (1972); Mitra & Datta (1967); Narayan & Muniyamma (1972); Saxena & Gupta (1970).
2n = 28: Larson, fide Kammacher et al. (1973).
2n = 36: Mehra & Sharma (1973); Pohl & Davidse (1971); Reeder (1971).
2n = 44: Mehra & Chaudhary (1974).
2n = 46: fide Kammacher et al. (1973).
2n = 54: Koshla & Mehra (1973).
2n = 72: Kammacher et al. (1973).
Remarks: Deviant forms with very small leaves (up to 1.5 cm long) are known in: India, Silhet: Dehlus s. n., CGE, Cameroon, Yaunde: Zenker & Staudt 515, Z. probably handled these plants as potential problems or as adjustments to step-plants.
Host (1805) listed a Panicum hirtellum for Europe. Host's herbarium specimens from the Hostchen Garden are, however, O burmanii. However, according to Reichenbach (1846) and Mertens & Koch (1823) they were taken from a native location (swampy spots on the seaside by Aquilegia). It appears in this case to be a one time find, perhaps a carryover from the distribution area in India. The clearly recognizable species has never been found in Europe again.
Long ago the species was once found in Caucasus. This appearance can be explained by anthropological carryover, in connection with the introduction of new subtropical grain forms (Komarov 1934). Ponert (1977) references a report from Makasvili (1936), about a 1930 population covering a 20 by 30 sq m area that was destroyed in order to prevent the expansion of this weed. This was probably a success as, apart from one other example from 1940 (Achanastilivi s. n.; K) no other plants were found in the Herbarium from Caucasus. For this reason the historical occurrence of the species in Caucasus was not included on the distribution map.
Selected Supporting Sources: Ethiopia: (Chiovenda 320; FI); 11 km SW of Bure (I. Frils et al.. 1904; K); Erytrea - Amasen, Fil-Fi1 (A. Pappi 5426; P) Gambela (0. Parker E. 454; K); 55 km W of Lekemti '(W. de Wilde 8926; BR). - Angola: Go1ungo Alto (Welwitsch 7252; 3003; BM). - Annobon_I.: (M. Rose 592; P). – Equatorial Guinea: Riomuni (J. A. Guerra 3879; FR)-. - Botswana: Kasane (A. Blair Rains 73; K). - Burundi: Bururi (J. Lewalle 4565; BR). – S. Antao: Cape Verde (R. J. Lowe s. n.; K). – Ivory Coast: Abengonzom (G. Roberty 12612; Z, 12638; G); near Abidjan (A. S. B. 13, 518; K); Singrobo (Adjanohoun 375 A; K).J:. – Fernando Po: (E. Guinea 576; K). - Gabun: Booue (J. Anton-Smith 295; K). - Gambia: W. of Jabang (A. Blair Rains 25; K), - Ghana: Anum (R. Rose Innes 31173; K); At Ho (Phillips 30552; K); Chebi (R. Rose Innes 30871; BR); Fosu (E. O. Asal 1775; K); Kumasi (BannermannBruce 4423; K). - Guinea: Badabon (R. Schnell 7552; K); Conakry (Maclaud 116; P); Dyeke (J. T. Baldwin j. 9654; K); Kouria (A. Chevalier 14924; K); Kousankoro (Adam 7078; P); Macenta (H. Jacques-Felix 1296; Pl. - Cameroon: Bankim (A. Meurillon 1069; P); Buea (F. D. Maitland 30; K); Douala and M. Bamboutos (A. Meurillon 52; P); Kribi (J. Schröder s. n.; FR) Victoria (Winkler 57; PRC). (F. D. Maitland 69; K); Yabom (R. Letouzey 7948; P); Yaounde (J. & A. Raynal 9422; P). - Kenya: (J. A. Allen 115; K); 20 m. S of Mombasa (D. Napper 1272; K); Gede. 60 m. N of Mombasa (A. Bogdan 4716; K); Kwale Distr., 18 m. S of Mombasa (R. B. Drummond & J. H. Hemsley 3998; FI); Malindi (Echlin 3; K); Marrarani. Boni (J. B. Gillespie 267; K); Nairobi (M. E. Church 80; K). - Liberia: Vonjama Distr. (J. 1. Baldwin 12030; K). – Madagascar: Central (G. W. Parker s. n.; FI); lhosy (H. Humbert 14451; G); North (M. Bernier 49; G); Tananarivo (J. M. Hildebrandt 4105; W). - Malawi: Distr. Dedza (E. A. Banda 439; BM); Chipata M. (W. C. Verboom 979; K); Citala R. (G. Jackson 488; K); Nyassa-See (Simons 1876; BM): near Zambia (B. Cormack 160: K). - Mozambique: Malema (A. R. Torre & J. Paiva 11189; BR); Manica e Sofala (Garcia 622; BM). - Namibia: Distr. Grootfontein North (H. Merxmüller & W. Giess 2025; M). - Niger: Nupe (Barter 1267; GOET). - Nigeria:· Prov. Abeokuta, Distr. Glokemeji (G. Jackson 3161168; K); Abiya (A. Blair Rains 246; K); 40 m. W. of Benin (Oshogbo 22; BM); Distr. Egba (Wit & Daramola 838; K); Gembu (Ekwuno 77232; K); lbadan (I. T. Swarbrick 2881; K); Lagos (W. McGregor 100; K); Distr. Orlu (Emwiogbon & Onyeachusim 65905; K); North, Panshanu Pass. (D. W. Lawlor & I. B. Hall 223; K). – Pemba I: Mahwikwi (J. H. Vaugham 205; BM). - Zambia: Abercorn Distr. Chilongolwelo (M. MacCallum Webster A 295; K); Kalomo (W. L. Astle 2276; K); Luangwa Valley (W. L. Astle 4636; K); Lukusuzi (J. A. Sayer 1160; K); Mapanza (E. A. Robinson 1358; M); Distr. Mporokoso, Kundabwika Falls (J. B. Phipps et. al. 3166; K); Mu-Lungushi (J. Kornas 3501; K), Mwembeshi (D. Vesey-Fitzgerald 4037; BM); Namwala (W. L. Astle 2956; K); Ngoni, Fort Jameson Distr. (W. C. Verboom 572; K). - Senegal: (Hudelst 590; OXF); Casamance (G. Roberty 6415; G), (J. G. Adam 18186; K); Bassin de la Gampie (R. P. Berhaut 2780; p); Jardin de Honu (A. Chevalier 44,097; P); Thies-Diourbel (R. P. Berhaut 4054; P); Tiazoye (G. Roberty 6287; Z). – Sierra Leone: Bumbuna (N. W. Thomas 3343; K); Freetown (D. Gledhill 503; K): Kambia (J. C. Deighton 806; K); Keneme (J. C. Deighton 417; K); Koinadugu Distr. (H. Mead 232; K); Newton (J. C. Deighton 1453; K); Njaja (Morton & Jarr 2359; K). - Socotra: Tamarid (G. Schweinfurth 422; K). S. Thome: Nova Moka (A. Moller 140; K). - Equatorial prov. (J. Wyld 267: BM). - Tanzania: Chimala Hospital (R. A. Nicholson 213: K); lringa Distr., Lukosi R. (J. Procter 3341; K); Milepa (D. P. Pielon 139; K); Shinyanga (B. D.;Burtt 2583; BR); Songea Distr., Rovuma R. (E. Milne-Redhead & P. Taylor 9561; B); Wala R. (R. Böhm 99; W);Ufipa Distr., Kisa-Rukwa (M. McCallum Webster 176; K). - Togo: Anecho (P. G. Mahoux 313; P); N of Palime. Daye Povohi (H. Scholz 126; B). – Peoples Republic of Congo: Brazzaville (Coomay s. n.; BR). - Zaire: Bambesa (P. Gerard 4.636; BR): Route de Boma (R. Devred 3351; BR); Gandayika (L. Liben 2792; BM); 50 km NE of Lubumbashi (S. Lisowski 591; K); Prov. Katanga, Kundelungu (de Witte 6015: K): Kialo (A. Thiebaut 295 b; BR); Kimpako (H. Vanderyst 5333; BR): Kisantu (R. Germain 2059; K); Leopoldville (W. Fasseaux 1071; K); Popokabaka (L. Pauwels 2279; BR); Stanley Distr., Mokoba (H. Vanderyt 3780; K); Kiunda, Route de Tumba a Thysville (P. Compere 2123: BR); Thysville (Bequaert 7733; BR). – Central African Republic: Haute-Kotto (G. le Testu 3.272; P). - Zimbabwe: Darwin Distr. (J. B. Phipps 2441; M); Gokwe Oistr. (BM); Kariba· Distr. (BM); Lomagundi Distr. (J. B. Phipps 2489; M); Marandellas (J. B. Phipps 1532: K); Mrewa (S. Mari 920; K); Mtoko (J. B. Phipps 2533: K): Salisbury (F. Eyles 3429; K); Selukwe Distr. (H. M. Biegel 2042; BM); Umtali (A. O. Crook 911; BR): Urungwe: Ruesi R. (R. Goodier 553; K); Victoria Falls (Simon 2141; K); Walobo Distr. (0. B. Willen 5816; BM).
Burma: Kyauktan (Po Khant 1346; K); Rangoon (C. E. Parkinson 15192; K); Syriam (H. S. McKee 5793; K). - Ceylon: Hambantola Distr. (F. R. Fosberg 50166: K): Paranatotopola (R. G. Cooray 691 20728; K); Vavuniya Distr. (D. Clayton 5298; K). - India: M. Abu (E. Blatter 2714: K); Ananthapura (A. Meebold 10484; . K); Assam Shillong (Thakur Rup Chand 8256; K); Bengal (Mokim 1520; G); Baidyanath (J. O. Nusker 1179; M); Bihar (E. Janaki 1453; B); Bombay (R. R. Fernandez 773; K); Hiran R. (K. T. B. Hodd 251; K); Madras, Bison Hill (5099; K) Madras; Calicut (K. Rungache s. n.; K); Dehra Dun (U. Singh 492; K); Madras, Ganjam Distr. (V. Narayanaswami 4682; K); Ghatoli (B. M. Wadhwa 7551; K); Prov. Khasia, Mairong (Schlaginweit 127; PR); Malad (G. L. Shah 801; K); Morabad (J. Thomson 8)25; K); Surguja State (N. Koelz 19034; K); W. Himalaya Saharanpur (J. F. Duthie s. n.; K); Orissa, Jojomura (H. F. Mooney 2; K); Sikkim, Selim (C. B. Clarke 36709; K); Travancore (M. Rama Rao 1807; K). - Indonesia: Kl. Sunda I., Alor (0. Jaag 1367; ZT); Celebes, Prov. Minahassa (S. H. Koorders 17625 b; K); Seran, Boano (Kornassii 1322; K); Wetar, Tarra (5. Bloembergen 3726; PNH); Bali; M. Prapat (Kostermans et al. 52; K). - Nepal: Butwal (Stainton et al. 8847; K); Godovari, S of Kathmandu (A. O. Schilling 1083; K); Okhaldhunga (A. Zimmermann 862; G); Singpur (D. Schaaf 117; K); de Thari a Ganga (A. Zimmermann 1250; G). – New Guinea: Saroa (Murhy S. n.; G). - Philippines: Prov. Rizal, Luzon (E. D. Merrill 830; G);- Lamao R., Mt. Mariveles, Luzon (R. S. Williams 135; K); Prov. Batangas, Luzon (M. O. Sulit 22474; PNH); Antique Prov., Panay (V. Aligaen 22147: PNH). - Thailand: Chiengmai (T. Sφrensen et al. 5927; K). – U.S.S.R.: Georgia, Adsharistan, Transcaucasus (Achanastilivi s. n.; K).
Brazil: Mattogrosso (Pilger 339; B). - Colombia: Dep. Cauca (A. Asplund 10525; G); El Tambo(K. v. Sneidern 165; G). - Costa Rica: Reventazon R. (G. Cufondoti 566; G). - Cuba: Papayo (E. L. Ekman 10335; G). Dominican Republic: Constanza (H. Türckheim 2883; M);·Ecuador: Ambato (E. Heinrichs 140; G); Guayaquil (E. Asplund 15988; G); Naranjapata (Schimpff 545; G); Pichincha Prov. (I. Holmgren 860). – El Salvador: Dep. Morazan (J. M. Tucker 441; G). - Guatemala: Dep. Quezaltengango (L. O. Williams 22990; G); San Jose (Brenning 445; B); Dep. San Marcos, Palo Gordo (L. O. Williams 25992; G). - Mexico: Cuernavaca (C. G. Pringle 6209; PRC); Jalisco (J. R. Reeder 2350; FR); Morelia, Rincon (Arsene 38;.G); Puebla (H. J. Conert s. n.; FR); Valee de Mexico (Bourgeau 1301; G). - Nicaragua: Dep. Matagalpa (L. O. Williams 23396; G). - Panama: Prov. Panama (H. Pittier 6822; G); Sabanas, near Chepo (A. A. Hunter & P. H. Allen 13; G). - Peru: Dep. Huanuco (E. Asplund s. n.; G).
var. lanatus (Buse) Backer, Handb. Fl. Java 2: 172 (1928).
Orthopogon burmannii (Retz.) R. BR. var. lanatus Buse in Miq., Pl. Jungh.: 370 (1854). - Type: Java, Pidjugan: Junghuhn s. n., n. v.
Panicum album Poir., Encycl. Suppl. IV: 274 (1816). - Oplismenus albus (Poir.) Roem. & Schult., Syst. Veg. II: 890 (1817). - Orthopogon albus (Poir.) Nees ex Steud., Syn. Pl. Glum. I: 44 (1854), pro syn.; Nees ex Miq., Fl. Ind. Bat III: 442 (1857, t.p. 1855). - Type: Java: Thunberg s. n., FI, Holotypus.
Oplismenus javanicus Klotzsch ex Schlecht., Linnaea 31: 309 (1861-62), nom. nud. - Type: Java, LE (dub.).
Panicum hirtellum Burm. f. non L.: Fl. Ind.: 24, t. 13 (1768).
Leaf sheath thick and long hair on the edge, on the flat dense or sparsely hairy. Rachis usually thick and long hair growth. Side shots 0.5 to 1.2 cm long, 10 to 16 spikelets in placed thick succession on the lowest. Spikelets 3 to 3.5 mm long, thick long hair; glumes entire, not notched on the tip (s. O. affinis).
Distribution: Indonesia
Selected Supporting Sources: Chad: Lac Tschad (A. Chevalier 6067; P). - Indonesia: Java: Batavia (C. Schröter s. n.; ZT), (Junghuhn s. n.; L), (v. Steenis 5298; K), (de la Savinierre 1568; G), Labillardiere (P. B. Webb s. n.; K); Semarang, Telomajo (Koorders 27777 b; K); Kl. Sunda I., Alor (0. Jaag 1017; PNH). - Japan: (9777; L). Vietnam: (M. Germain s. n.; K); Massif du Grand Eperon (Poilane 579; K); near Saigon (M. Godefroy-Lebeuf s. n.; K).
var. multisetus (Hochst. ex A. Rich.) U. Scholz, novo stat.
Panicum (Oplismenus) multisetum Hochst. ex. A. Rich., Tent. Fl. Abyss. II: 377 (1850). - Type: Ethiopia: Schimper 149, BM, Holotypus, FI, Isotypus.
Inflorescence not hairy, side shoots 0.5 to 2 cm long, carrying 5 to 15 spikelets, arranged in succession. Spikelets 2.4 mm long, never with hair.
Distribution: Ethiopia
Supporting Source: Ethiopia: Tacozethal by Djeladjeranne (Schimper 1469; FI).
Oplismenus flavicomus Mez, Noizbl. Bot. Garden, Berlin-Dahlem 7: 55 (1917). – Type: Madagascar. Forsyth-Maajor 208, B, Holotypus; G, Isotypus.
Description: Perennial grass; culm 30 to 45 cm high. Leaves lanceolate-linear, 0.6 to 0.9 cm wide, 6.5 to 10 cm long; Mid-vein clearly defined. Sheath lightly wedged open in the upper part, very long (up to 5mm) sparse hair on the edge, sparse and long on the flats, thick long hair on the upper part. Ligule Hairy collar on the end, 0.7 to 1.4 mm long. Inflorescences racemose, placed together, 10 to 11 cm long, no hair, somewhat coiled. Side shoots upright, heavily haired. The length of the side shoots decreases from the bottom up, the lowest is 1.5 to 2 cm long and carries, unilaterally approx. 20 spikelets, arranged in clumps or oblong clumps. The distance between the lowest side shoots is 3 cm. Spikelets placed together in pairs, lanceolate, 3.8 to 4 mm long, either without hair or only a little on Gluma I. Gluma I oblong, 2.6 to 2.9 mm long, 1.6 mm wide, 3 to 5 veins, with awn. Gluma II longest glume of the spikelet, oblong, 3.7 to 3.9 mm long, 1.9 mm wide, 5 veins, with awn. Lemma I oblong, 3.6 to 3.8 mm long, 1.5 to 2.3 mm wide, 7 veins, sometimes with awn. Awns frail, yellowish and dentate; awn of Gluma I 3.3 to 8 mm long, of Gluma II 1.1 to 1.3 mm long. Lemma I has no awn or has an awn with a length of 0.7 mm. |
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Distribution: Madagascar.
Supporting Source: Madagascar, Ambohimitombo forest (Forsyth-Major 208; B).
Oplismenus gracillimus Mez, Notizbl. Bot garden, Berlin-Dahlem 7: 55 (1917). – Type: Nossi-be: Boivin s. n., B. Holotypus; W, Isotypus.
| Probably perennial grass; culm 30 to 40 cm high. Leaves oblong-lanceolate, 0.4 to 0.7 cm wide, 5 to 7 cm long. Sheath not wedged open in the upper part, long thick hair only on the edge. Ligule hairy collar on the end, 1.1 to 1.5 mm long. Inflorescences racemose, placed together, 12 to 20 cm long, no hair. Side shoots upright, leaning away. The length of the side shoots decreases from the bottom up, the lowest is 5 to 6 cm long and carries 11 to 22 spikelets loosely in succession. The distance between the lowest side shoots is 2 cm. Spikelets placed together in pairs, rounded, 1.5 to 1.8 mm long, without hair. Gluma I rounded, 0.8 to 1 mm long, 0.7 mm wide, 5 to 7 veins, with awn. Gluma II 1.1 to 1.3 mm long, 0.6 to 1.1 mm wide, 9 veins, pointed. Lemma I oblong, 1.5 to 1.7 mm long, 0.6 to 1.1 mm wide, 9 veins. Awns frail, whitish and dentate; awn of Gluma I 8 to 9 mm long. Gluma II only has a point and Lemma I has no awn. |  |
Distribution: Nossi-be (an island to the northwest of Madagascar)
Supporting Source: Nossi-be: Boivin s. n.; B.6.
Oplismenus humbertianus Camus, Nat. Malagache 5: 146 (1953). – Type: Madagascar, Vallee de l'Ifasy: Humbert & Capuron 25889; P, Holotypus.
Culm bare or little hair. Leaves oblong, 0.6 to 1 cm wide, 3 to 5 cm long, heavily haired on the upper and lower sides. Sheath bare on the flats, mid-length hairs along the edge. Ligule has hairy collar, 1.1 mm long. Inflorescence 5 to 6 cm long, hangs over partially. The side shoots are 1 to 2.5 cm long, the lowest carries 10 to 15 spikelets clearly arranged in succession. The distance between side shoots is 0.2 to 0.5 cm, The Inflorescence is thereby heavily pressed and compact. Spikelets arranged in pairs, the sitting one is usually reduced to an awn; 3.7 mm long, very little hair. Gluma I linear-lanceolate, 3.6 mm long, 0.9 mm wide, 5 veins, with awn; the awn originates 0.6 mm below the tip of the glume. Gluma II lanceolate, 3.4 mm long, 1.2 mm wide, 7 veins, with awn; the awn originates 0.4 mm below the tip of the glume. Lemma I oblong-lanceolate, 3.4 mm long, 1.9 mm wide, 9 veins, with awn; the awn originates a little below the tip of the lemma. Palea I narrow, 2 veins. Awns frail, white-reddish, dentate. Awn of Gluma I 16 mm, of Gluma II 1.1 mm and of Lemma I 0.3 mm long.
Distribution: Madagascar.
Supporting Source: Madagascar: Vallee de l'Ifasy, Distr. Ambilobe (Humbert & Cauron 25889; P).
Oplismenus sect. Orthopogon (R. Br.) Schlecht., Linnaea 31: 301 (1861-62).
Oplismenus aemulus (R. Br.) Roem. & Schult. ‘O. aemulans' , Syst. Veg. II: 487 (1817).
Key to the Varieties:
| 1a | Rachis glabrous; spikelets 2.4 to 3.1 mm long and clearly successive; awn of lower glume 2.4-3.1 mm long | var. flaccidus |
| 1b | Rachis and usually the primary branches of the inflorescence densely hairy | 2 |
| 2a | Awn of lower glume 6 - 9 mm long; awn of upper glume 1-1.5 mm long; lower lemma pointed or blunt; primary inflorescence branches clearly separated from one another; spikelets clearly successive | var. aemulus |
| 2b | Awn of lower glume 1.5 - 3.5 mm long; awn of upper glume 0.2- (1.1) mm long, lower lemma unawnd; primary inflorescence branches densely packed, the inflorescence appearing clumplike; spikelets in thick succession | var. densiflorus |
Orthopogon aemulus R. Br., Prod. Fl. Nov. Holl. 194 (1810). - Panicum aemulum (R. Br.) Steud., Nomencl. 2.ed. II: 252 (1841). – Oplismenus setarius (Lam.) Roem. & Shult. var. aemulus (R. Sr.) Bailey, Quensland Fl. 6: 1838 (1902). - Oplismenus aemulus (R. Br.) Roem. & Schult. var. pilosus Domin, Biblioth. Bot. 85: 328 (1915). - Type: Australia, Keppel Bay: R. Brown 6132, BM, Holotypus.
Oplismenus aemulus (R! Br.) Roem. & Schult. var. lasiorhachis Domin, Biblioth. Bot. 85: 329 (1915). - Type: Australia, rainforests in the Tambourine Mountains: Domin s. n.; n. v. (PR?).
Oplismenus burmannii (Retz.) P. Beauv. var. intermedius Honda, Bot. Mag. Tokyo 38: 191 (1924). - Oplismenus compositus (l.) P. Beauv. var. intermedius (Honda) Ohwi, Acta Phytotax & Geobot. 2: 35 (1942). - Type: Taiwan, Kobayashi 486, TI, Lectotypus; Owatari s. n., Sasaki 19, TI, Syntypi.
Oplismenus compositus (L.) P. Beauv. f. pubescens F. Brown, Bernice P. Bishop Mus. Bull. 84: 68 (1931). - Type: Marquesas (1.), Nukuhiva, Taipi Val: F. Brown 594; BlSH, Isotypus.
Usually frail grass; leaves oblong-lanceolate, 0.4 to 1.1 cm wide, 3 to 8 cm long, More or less thickly haired. Sheath long thick hair growth on edges. Ligule has short hairy collar, 0.4 to 1.1 mm long. Inflorescences racemose, placed together, 8 cm long. Culm has long thick hair; side shoots usually have long thick hair as well; the lowest is 1.5 to 2.5 cm long and carries approx. 15 spikelets more or less in thick succession. Spikelets placed in pairs, oblong-lanceolate, long or short thick hair, 2.8 to 3.2 cm long. Awns robust, yellowish, bare; the awn of Gluma I is 6 to 9 mm long, of Gluma II 1 to 1.5 mm long. Lemma I is pointed or blunt. Distribution: Tropical East Asia and Australia.
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Supporting specimens: Australia: Brisbane (Clemens s. n.; U. S.); Moreton Bay (F. v. Mueller s. n.; W); N. Queensland (N. Michael 920; E); N. S. Wales, Byron Bay (Meebold 3380; M). - New Caledonia: Babrongho, Sentani (C. Versteegh 4742; PNH). – New Hebrides: Aneitum (Cuming 17; G). – Phillipines: Luzon, Baguio (Sulit 7602; PNH).
var. flaccidus (R. Br.) Domin, Biblioth. Bot. 85: 328 (1915).
Orthopogon flaccidus R. Br., Prod. Fl. Nov. Holl.: 194 (1810). - Oplismenus flaccidus (R. Br.) Roem. & Schult., Syst. Veg. II: 487 (1817). - Panicum flaccidum (R. Br.) Steud., Nomencl. 2. ed. II: 256 (1841). - Type: Australia, Newcastle Distr.: R. Brown, 6131, BM, Holotypus.
Usually frail grass; leaves ovate-oblong or oblong lanceolate, 0.8 to 1.1 cm wide, 5 to 7 cm long. Sheath sparse thin hair on the edge. Ligula has a hairy collar, 0.6 to 1.6 mm long. Inflorescences racemose, placed together, 7 cm long.; culm and side shoots bare. Side shoots 0.5 to 3 cm long; the lowest carries 6 to 15 spikelets more or less in thick succession. Spikelets placed in pairs, oblong lanceolate, sometimes a little hairy, (2.2) to 2.4 to 3.1. to (3.7) mm long. Awns robust, yellowish and bare; the awn of Gluma I 2.6 to 7 mm long, of Gluma II 0.2 to 1.9 mm long; Lemma I has a short point or is blunt. Distribution: Tropical East Asia and Australia. Distribution: Tropical East Asia and Australia. Ecology: In rainforests and treed savannah. Sociology: Under Casuarina, Pandanus, Melaleuca; with Ottochloa nodosa. Anthesis: April to November. Remarks: An especially small leaved and frail form is well known from Ceylon: Sittampalam s. n., HBG. |
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Selected Supporting Sources: Australia: (Sieber 73; LE); Cape York. Queensland (L. J. Brass 18214; L); Middle Percy Isl., Queensland (M. Lazarides 5645; B); Hastings R., near Richmond R. (Beckler s. n.; M) Lord Howe I. (Marder s. n.; G) Ceylon: (Walker s. n.; G). – Community-I.: (Forster s. n.; M). - Indonesia: Bali (Kosterman et al. s. n.; NY); Java; Mt. Gede (Hochreutiner 1198; G); Tongatabu (T. B. Cartwright s. n., OXF). - Malaysia. Borneo, M. Kinabalu (J. & M. S. Clemens 32613; G). - Mauritius: (Commerson 135; LD). – N. Caledonia: (Germain s. n.; W). - N. Zeeland: (Hooker 1754; G); Tutu (L). Philippines: Luzon, Mt. Mariveles (Merrill 6987; G); Lepanto; Mt. Data (M. Ramos & G. Edano s. n.; W). - Samoa: (S. J. Whitmee s. n.; W); Nu'ulua (A. Whistler 1935; B). - Taiwan: Tammi (U. Faurie 117; PNH).
var. densiflorus U. Scholz nov. var. – Type: New Guinea: W. Vink 16332, L, Holotypus.
Diagnosis: Lamina foliorum ovata – lanceolata vel lanceolata; inflorescentia racemiformis, rami coarcti. Spiculae ovatae – lanceolatae, dense contingentes; arista glumae I 1.5 – 3.5 mm longa, glumae II 0.2 – (1.1) mm longa. Lemma I muticam.
Ovate oblong, sometimes oblong leaves; side shoots arranged in thick succession, an almost clumped inflorescence; Spikelets inserted in thick succession, ovate-oblong. Awn of Gluma I 1.5 to 3.5 mm long, of Gluma II 0.2 to (1.1) mm long; Lemma I has no awn. Distribution: New Guinea, Australia. Common name: Sunsba-Magngak (New guinea); Tauri (New Guinea: Enga-language). Ecology: Weed in cultivated fields.
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Oplismenus compositus (l.) P. Beauv., Ess. Agrost.: 54 (1812); Bor, N. In Rechinger, K. H., Fl. Iranica: 484 (1970); Bosser, J., Gram. Pat. et Cult. Madagascar: 355 (1969); Cooke, T., Fl Pres. Bombay 3: 443 (1958); Diels, F. L. E., Fl. Centr. China Bot. Jahrb. Syst. 29: 223 (1901); Engler, A., Plant World of East Africa: 104 (1904); Gamble, J. S., Fl. Pres. Madras 3: 1231 (1967); Hara, H., Fl. Eastern Himalaya: 369 (1966); Haines, H. H., Bot. Bihar and Orissa 3: 1045 (1961); Koorders, S. H., Exkursionsflora von Java 1: 138 (1911); Mitra, N. J., Flow. PI. East. Ind. 1: 197 (1958); Prain, D., Bengal Plants 2: 883 (1963); Ridley, H. N., FI. Malay Penins. 5: 221 (1925); Stapf, O. in Prain, D., Fl. Trop. Afr. 9: 634 (1934); Stewart, R. R., Fl. West Pakistan 120 (1972).
| 1a | Lowest inflorescence branch 2.5 to 3 cm long; spikelets 2.4 to 4.1 mm long, arranged in thick succession; Mauritius | var. sylvaticus |
| 1b | Lowest inflorescence branch 1.5 to 10 cm long; spikelets 2.6 to 5.2 mm long, loosely arranged in succession. | |
| 2a | Plants usually frail; spikelets 2.6 to 3.2 mm long; inflorescence branches 1.5 to 5 to (7) cm long | var. rariflorus |
| 2c | Plants robust; spikelets (3.3) to 3.6 to (5.2) mm long; inflorescence branches 3 to 10 cm long | var. compositus |
Panicum compositum L. Spec. PI. 57 (1753). - Orthopogon compositus (L.) R. Br., Prod. FI. Nov. Holl.: 194 (1810). – Orthopogon remotus (L.) Trin., Fund. Agrost.: 181 (1820), nom. illeg. - Hippagrostis composita (L.) O. Kuntze, Rev. Gen.: 777 (1891). - Type: Ceylon, Hb. Hermann (BM?)
Panlcum aristatum Retz . Obs. IV: 17, (1786-87). Typus: China: Wennerberg s. n. (LD?).
Panicum composito-proximum Rottl. ex Willd., Ges. Naturf. Fr. N. Schrift. 4: 224 (1804), - Oplismenus indicus Roem. & Schult., Syst. Veg. II: 484 (1817). - Panicum certificandum Steud., Syn. Pl. Glum. I: 44 (1854). - Type: India, Bengal: Rottler 869, M, Holotypus.
Oplismenus decompositus Nees in Endl., Prod. Fl. Norf.: 19 (1833). - Panicum peninsularum Steud., Syn. Pl. Glum. I: 44 (1854). – Panicum compositum Rottl. ex Steud., Syn. Pl. Glum. I: 44 (1854), pro syn. - Type: India: Wight prop. 62, NY, Holotypus.
Panicum bidentulum Steud., Syn. PI. Glum. I: 45 (1854). - Orthopogon junghuhnii Nees ex Steud., Syn. Pl. Glum. I: 45 (1854), pro syn.; Nees ex Miq., Fl. Ind. Bat. III: 444 (1857, t. p. 1855). - Panicum bidentatum Steud., Syn. Pl. Glum. I: 45 (1854), err. pro P. bidentulum Steud., - Oplismenus junghuhnii (Nees ex Miq.) Boerlage, Ann. Jard. Bot. Buitenzorg 8: 63 (1890). -Type: Rebo Running: Nees S. n., B, Lectotypus.
Orthopogon compositus (l.) R. Br. var. glabrescens Buse in Miq., Pl. Jungh.: 370 (1854). - Type: Java, Merapi: Junghuhn s. n.; L, Holotypus:
Panicum gonyrrhizum Steud., Syn. Pl. Glum. I: 44 (1854). Orthopogon gonyrrhizus (Steud,) Miq., FI. Ind. Bat. III: 443(1857, t. p. 1855). - Type: Java: Goering 199, P, Holotypus.
Panicum longeracemosum Steud., Syn. Pl. Glum. I: 45 (1854). - Orthopogon longeracemosum (Steud.) Miq., Fl. Ind. Bat. III: 443 (1857, t. p. 1855). - Type: Java: Zollinger 915, P, Lectotypus; 719 (P?). Syntypus, n. v.
Oplismenus compositus (L.) P. Beauv. var. lasiorhachis Hack., Bur. Gov. Lab. 35: 81 (1905). - Type: Philippines, Separation Point, Paraguai: Merrill 826 (PNH, destr.?), n. v.
Oplismenus formosanus Honda, Feddes Repert. Spec. Nov. Regni Veg. 20: 361 (1924). - Type: Taiwan: Hayata s. n., TI, Holotypus.
Oplismenus owatarii Honda, Feddes Repert. Spec. Nov. Regni Veg. 20: 361 (1924). - Oplismenus compositus (L.) p, Beauv. var. owatarii (Honda) Ohwi, Acta Phytotax. Geobot. 11: 35 (1942). - Type: Taiwan, Buizan: Matuda-Bizi 316, TI, Lectotypus; Owatari s. n., (TI?), Syntypus, n. v.
Oplismenus patens Honda, Feddes Repert. Spec. Nov. Regni Veg. 20: 360 (1924). - Oplismenus compositus (L.) P. Beauv. var. patens (Honda) Ohwi, Fl. Jap.: 149 (1953). - Type: Japan, Oshima: Uchiyama s. n., TI, Lectotypus; Kuroiwa s. n., (TI?), Syntypus. n. v.
Oplismenus compositus (L.) P. Beauv. f. glabratus F. Brown, Bernice P. Bishop Mus. Bull. 84: 68 (1931). - Type: Marquesas, Nukuhiva: F. Brown 736, BISH, Lectotypus; 927 (BISH?), Syntypus, n. v.
Oplismenus undulatifolius (Ard.) Roem. & Schult. var. elongatus Honda ex Nakai, Koryo Shikerin No Ippan: 80 (1932). - Type Nakai 13237, TI, Isotypus.
Oplismenus coreanus Nakai, Bull. Sei. Nat. Mus. 31: 140 (1952), p. p., nom. illeg. ,- Type: n. v. (TI?)
Oplismenus burmannii auct. non Retz.: Mez in Perkins, Frag. Fl. Phil.: 144 (1904): Thwaites, Enum. Pl. Zeylan.: 358 (1854), Supporting Source: Thwaites s. n., W.
Hybrids:
S. u. Oplismenus burmannii (Retz.) P. Beauv.
Ic.: Bosser, J., Gram. Pat. et Cult. Madagascar: 355 f. 133 (1969). – Fröman, B. & Persson,S., An ill. Guide to the Grasses of Ethiopia: 359 L 219 (1974). - Hsu, K. - S., Fl. Taiwan 5: 564 f. 1433 (1978). - Pilger, R., Panicoideae in A. Engler, Die Nat. Pflanzenfamilien 14 e: 48 f. 25 (1940).
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Perennial, usually hardy plants; culm 30 to 106 cm high. leaves oblong-lanceolate or – less often – ovate-oblong, hairy or bare , (0.4) to 0.8 to 1.6 to (2.2) cm wide, (3) to 4 to 9 to (16) cm long. Sheath long or short haired along the edge, seldom hair on the flat, loose long or short. Ligula hairy collar on the end, (0.3) to 1.3 to (2) mm long. Inflorescences racemose, placed together, the rachis is flat to triangular, between 6 and 32 cm long, almost always bare. Side shoots 6 to 10, sanding out at an angle. The length of the side shoots decrease from bottom to top, the lowest is 3 to 10 cm long. The side shoots are far apart, usually with at least the distance of their own length. The lowest shoot carries 8 to 20 to (40) spikelets unilaterally, arranged loosely to very loosely in succession (separation of up to 1 cm between spikelet pairs). Spikelets placed in pairs, dark green, sometimes purple overrun, lanceolate, (3.3) to 3.6 to (5.2) mm long, bare or hairy. Gluma I oblong-oval, (1.8) to 2.6 to (3.7) mm long, (0.7) to 1.2 to (1.6) mm wide, (3) to 5 veins, with awn. Gluma II oblong-oval, (1.8) to 2.5 to (3.8) mm long, (1) to 1.5 to (1.9) mm wide, (5) to 7 veins, with awn or occasionally with a short point. Lemma I oblong, (2.7) to 3.3 to (5) mm long, (1.5) to 2.1 to (2.9) mm wide, with a short point (0.2 – 0.4 mm) or blunt. Palea I thin, translucent, 2 veins, sometimes missing. Caryopsis Oblong, convex on the back, flattened on the front. Awns robust, yellow-reddish, bare, becomes adhesive when fruit is ripe; awn of Gluma I 3 to 15 mm long, of Gluma II 0.7 to 5 mm long.
Distribution: Tropical and subtropical regions of the Old and New World.
Common name: Taru (Pogatumo-language, Sumo); Tepe (Orne-language, Mafoka); Sihuhu Mairoro (Orokaiva-language) – New Guinea.
Ecology:In humid Rainforests: along rivers in cloud forest and secondary forests; in bamboo thickets and in reed vegetation; on granite.
Sociology: Under Newtonia, Artocarpus, Northea, Eugina uniflora, Morinda citriflolia, Leucaena; Quercus (Mexico). Associates plants: Philippines: Panicum philipes, Echinochloa colona, Carex, Sida, Hyptis.
Economic impact: Weed in coffee plantations.
Pests: Claviceps viridis Padwick & Azmatullati; Helminthosporium oplismeni Sawada & Katsuki; Phakopsora oplismeni Cummius; Phyllachora oplismeni-compositi Sawada; Phyllachora oplismeni Syd. var. major Batista; Physopella oplismeni B. U. Patil & Thirumalachar;.
Anthesis: January to December; especially march to May and October to December.
Chromosomes:
n = 27: Koshla (1972).
2n = 54: Koshla & Mehra (1973); Mehra & Sharma (1973).
n = 36: Gupta (1971).
2n = 72: Avdulov fide Bor (1960); Gould & Soderstrom (1974); Hsu (1967); Narayan & Muniyanna (1972).
Remarks: More hairy forms are appearing: 1. Culm, leaves, flats of the sheath have thick long hair: Mozambique, Manica e Sofala: Chase 7436; BM; Tansania, Mahenge: Schlieben 1997; HBG; Samoa, Upolu: Whistler 1615; B. - 2. Leaves, body of the sheath has thick long hair: Java, Semarang: Koorders 27780 b; B; Malaysia, Sabah: Nooteboom 1193; B; Thailand, Fang: Sφrensen et al. 1519; C; Payab: Sφrensen et al. 3045; B; Japan, Kyushu: Togasi 1475; B. - 3. Spikelets partially, rachis and side shoots have thick, long hair: Australia: Müller s. n.; G; India: Griffith s. n.; OXF; Bacungan, Edano 222; PNH.
Another form that has deviated from O. compositus var. compositus is the species described by Honda as Oplismenus patens. It is distinguished by especially wide leaves (2 to 3 cm): Ceylon: Koenig s. n.; M; Faurie 6407; G; Japan, Tokyo: Tagshi 7276; L; Java: Nagler 15; B; Vietnam, Tonkin: Balansa 1605; G.
Selected Voucher Specimens: Ethiopia: Eritrea-Amassen , Fil-Fil (A. Pappi 5426; FI); Ghinda (A. Pappi 4526; FI); Jimma (Massa 280; FI). - Kenya: Kisumu, Nairobi (R. A. Dümmer 1640; BR); Kwale Distr. Longomwagandi (F. Magogoa & P. Glover 940; BR). - Madagascar: Tamatare (M. Goudot s. n.; G). - Malawi: Chipata M. (W. C. Verboom 971; 8M); Kyimbila (A. Stolz 1239; Z);Dedza Distr., Nchisi For. (I. D. Chapman 1311; 8M); Kota-Kota Distr,. (J. Brass 17044; BR). - Mozambique: Sofala Prov., Chimoio (N. C. Chase 6874; BR); Between Nicuadala und Narral (A. R. Torre 4427; BM); Serra da Morrumbala, Massingire (A.R. Torre 5-231; BM). – Pemba I: Mkumbua (J. H. Vaughan 211; BM). - Sambia: Dlstr . Mporokoso, Kundabwika Falls (J. B. Phipps & D. Versey-Fitzgerald 3159; BM). - S.Africa: Prov. Cape, Distr. Port St. Johns (M. Y. Wells 3392; BR); Mariepskop (H. van der Schijff 4325 A;W). - Tansania: Usambara, Gonja (C. Holst 4241; PR); Usambara; Lutini (C. Holst 3279; P); Mahenge Distr., Sali (H.J. Schlieben 1997; M); Massagati. Ruhudje (H. J. Schlieben 1179; HBG); East-Usambara, Amant (A. Peter 20189; W); Zansibar, Muyuni (J. H. Vaughan 2208; BR). - Zimbabwe: Chihi Distr., Madzivire Dep. (R. _B. Drummond 7895; BM); Gokwe Distr., Lasame R. (M. Y. Bingham 640; BM); Nyumquarara Valley (H. B. Gilliland 1549; BM); Selukwe (H. Wild 4290; BR); M. Sindinda Distr. (H. Wild 1901; FI); Umtali, Mennine R. (N. C. Chase 4485; BM). - Australia: Queensland, Cape York Penins~ (J. Brass 19318; G); Queensland, Rockingham Bay (F. Müller s. n.; G). - Burma: Myitkyina (R. O. Belcher 840; G); Kyajkiyu, Ramree I.. (G. C. Wallace 9022; B); Fort Stedman (A. Huk s. n.; M). - Ceylon: Kandy, Roseneath (F. Ballard 1007; B); (Thwaites 913; FR). - China: Fukien Prov. Hinghwa (lin Pi 6424; W); Hainan, Tao (C. Wang 35294; G); Kiangsi; Lungnan Distr. (S. K. Lau 4449; G); Prov., Kwangtung, Tao poo (S. S. Sin et al. 113; B); Prov. Kwangtung, Wan Shan Toi (C. O. Levine 10172; PNH); Yunnan, Ma Shan, Mekong Yangtze Div. (G. Forrest 132.00; E). – Fidii I: Ovalan (A. V. Hugel 99; CGE). - India: Assam, Tirap R. Valley (R. O. Belcher 22 C 903; G); Calcutta (P. Drain S. n., G); Cote Coromandel (M. Belanger 149; G); Darjeeling (C. B. Clarke 24832; G); East Himalaya (Griffith s. n.; OXF); Himalaya (Hooker s. n.; OXF); N. W. Himalaya, Mussoorie (R. R. Stewart 11421; G); Malangor (Hohenacker 368; G); Saharanpur, Rajpur (J. J. Duthie 7720; G); Sikkim (I. D. Hooker s. n.; M); Shembaganur (Foreau 1778; M). - Indonesia: Borneo, Bangarmassing (J. Motley 151; CGE); Borneo, Banguey I. (P.' Castro & F. Melegrito 1486; G); Borneo, Mt. Kinabalu (Clemens 51051; G); Borneo, Penibukan (Clemens 51567; HBG); Borneo, Sarawak (A. Brooke 8148; G); Borneo, Serawai (Winkler 100; HBG); Borneo, Tenompok (Clemens 30289; HBG); Celebes, Provo Minahassa (S. H. Koorders s. n., L); Java, Manau, near Buteng (Kostermans & Wirawan 496; G); Salak, near Bogor (Kurz 1837; M); Java, ldjengebirge (C. Schröter 7; ZT); Java, Merapie (Junghuhn s. n.; L); Java Pengalengan (Hochreutiner 1329; G); Java Gede Tji-Beureum (Hochreutiner 15; G); Java Viah Tanabong (E. de la Savinierre 1637; G.l; Sumatra, Buidjei, Deli (L. Martin S. n.; M); Sumatra, Laut Kawar (Frey-Wyssling 8053; ZT); Sumatra, Asahan (A. Krukoff 4056; J); Sumatra, Tiga Dolok (Surbeck s. n.; ZT); Kl. Sunda I., Alors (0. Jaag 327; ZT); Lombok (J. Elbert 1893; PNH); Timor (0. Jaag 53; ZT). - Japan: Kyushu, Ambo (M. Togasi 1475; G); Ryukyu, Taketomi (F. R. Fosberg 37617; L). -Malaysia: Langkawi, Sungai Raya (E. 1. H. Corner S. n.; PNH). - Mollukken: Amboine (Webbs. n.; G). - Nepal: away from Azidunga toward Belzat (A. Zimmermann 2117; G). – N Calendonia: (G. Bonati 391; HBG); Yahoue; (1. Franc 2143; G). – New-Guinea: Kiwori (Murphy s. n.; G); Waikaiuna (L. J. Brass 25394; PNH ); Sepik Distr., Mori (P. J. Darbyshire & R. D. Hoogland 8085; B). - Pakistan: Saidpur (R. R. Stewart 23667; W). - Philippines: Luzon: Ilocos Norte Prov. M. Quebrada (G. E. Edana 17856; PNH); Provo Rizal, Antipolo (E. D. Merrill 200; PRC); Provo Sorsogon. Irosin (E. D. Merrill 15347; B); Mindanao: Agusan Prov., Butan (D. R. Mendoza 42021; PNH); Cotobato prov. (H. Kerr 35211; PNH); Davao Prov. M. McKinley (G. E. Edana 1026; PNH); Mindoro: Bongabon (E. Maliwanag 171; PNII); Mt. Yagaw (M. D. Sulit & H. C. Conklin 16896; PNH); Guimaras: (A. Usteri s. n.; ZT); Leyte: Takuranga R. (B. Santos 39931; PNH); Palawan: Tarateon R. (G. E. Edano 14160; PNH); Puerto Princesa (G. E. Edano 222; PNH). - Panay: Ilvico Prov. (A. T. Taleon 33866; PNH). - Polillo: (R. B. Fox 9117; PNH). Samar: M. Capotoan (G. E. Edano 15576; PNH); Mt. Purog (G. E. Edano 15455; PNH). - Samoa: Upolu (Reinecke 13; L); Lanuto See (Hochreutiner s. n.; G). – Sevchellen I: (C. Jeffrey & A. Maulinie 790; P). - Thailand: Doi Sutap (Hosseus 305; M); Soi Dao, Chantaburi (N. L. Bor 329; C); Kanchanaburi (C. F. V. Bensehom 3506; L). - Vietnam: Vallee de Lankok, Tonkin (B. Balansa 1605; G). - Ecuador: Guayas (A. S. Hitchcock 20429; US); Guayaquil (E. Asplund 15925; G). - Hawaii: Halawa Valley (5. Ishikawa 134; L). - Mexico: La carrea (Langlasse 444; US); Jalisco (R. Me Vaugh 14190; G); Morelia (Arsene 39; G); Sierra Cordiellera (H. Galeotti 5847; G). - Panama: Canal Zone, Fort San Lorenzo (E. L. Tyson et al. 3704; MO); Prov. Chiriqui (G. Davidse 10, 154; MO). - Venezuela: Prov. Caracas (Funck & Schlim 161; G).
var. rariflorus (K. B. Presl) U. Scholz. nov. stat.
Oplismenus rariflorus K. B. Presl, Rei. Haenk: I: 320 (1830); Hitchcock, A. S., Contr. U.S. Natl. Herb. 22,3: 131, f. 24 (1920), North Amer. Flora 17,4: 309 (1931); - Panicum parciflorum Steud., Syn. Pl. Glum. I: 45 (1854). – Qplismenus hirtiflorus K. B. Presl in Kew Index (err. pro O. rariflorus K. B. Presl. - Type: "Acapulco", Haenke S. n. (PR?), n. v.; PR: Labeled as Type: "in Bengalia circa Calcuttam", Helfer "annls 1838-40".
Oplismenus latifolius Haenke ex Steud., Nomencl. 2. ed. II: 220 (1841). - Type: Peru: Haenke s. n.; n. v.; fide Hitchcock (1920).
Oplismenus liebmannii E. Fourn., Mex. Pl. 2: 38 (1886). - Type: Mexico: "in campis inter la
Galera et Pochutta", Liebmann 372, C, Lectotypus; Mexico, Zacuapan: Liebmann 373, C, Syntypus; Liebmann 374, Syntypus.
Oplismenus thiebautii E. Fourn., Mex. Pl. 2: 39 (1886). - Type: Mexico, Acapulco: Thiebaut 1074, P, Holotypus.
Oplismenus polliniifolius Honda, Feddes Repert. Spec. Nov. Regni Veg. 20: 362 (1924). - Type: Taiwan, Hokuto; Nakahara s. n., TI, Lectotypus; Miyake 24, TI, Syntypus, n. v.Ic.: Hitchcock, A. S., Contr. U.S. Natl. Herb. 22,3: 132, f. 24 (1920). - Matuda, E., Las Gramin. del Estado de Mexico: 54 (1958), ("Oplismenus hirtellus"). - Swallen, J. R., Fl. Guatemala 2: 229, f. 72. Fieldiana: Bot. 24. 2 (1955).
Usually frail grass: culm 30 to 50 cm high, bare or haired. Leaves ovate-oblong, bare or with little hair, (0.5) to 0.8 to 1.4 to (2.6) cm wide, (3) to 4 to 8 to (13) cm long. Sheath short, thick or sparse hair along the edge, usually bare on the flats. Ligula usually has a hairy collar, (0.2) to 0.4 to 1.3 to (2.2) mm long. Inflorescence placed together, racemose; rachis 5 to 15 cm long, usually bare; side shoots up at an angle and out from the main axis. The length of the side shoots decreases from bottom to top, the lowest is 1.5 to 5 to (7) cm long and carries 8 to 20 spikelets arranged in loose succession. Spikelets placed in pairs, heavily haired at the base. Spikelets oblong- lanceolate, (2.5) to 2.6 to 3.2 mm long, usually without hair. Gluma I oblong, (1.8) to 1.9 to 3.2 mm long, (0.9) to 1.1 to 1.2 to (1.4) mm wide, 3 to 5 veins, with awn. Gluma II oblong, 2 to 2.4 to (2.9) mm long, (1.2) to 1.3 to 1.6 to (1.9) mm wide, 5 to 7 veins, with awn. Lemma I oblong, (2.4) to 2.7 to 3.2 mm long, (1.5) to 1.9 to 2.1 mm wide, 7 to (9) veins. Palea I is not present. Caryopsis 2.5 mm long. Awns hardy, yellowish and bare; awn of Gluma I (2.7) to 4.2 to 8 to (13) mm long, of Gluma II 0.2 to 1.2 to (1.7) mm long. Lemma I has a short awn (0.4 to 0.6 mm long) or is blunt. Distribution: Tropical regions of the New and Old World. |
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Ecology: In moist, shady areas - coppices and along rivers or streams.
Sociology:Under Pterocarpus (Sunda I.), Pinus, Quercus (Mexico). With Tristania sp. and Imperata cylindrica.
Notes: Here also there are especially hairy forms: 1. Inflorescence and inflorescence branches are thick and long-haired: Ceylon: Reynard s. n.; B; India: Waitz s. n.; L; Maligodam: Rebu s. n.; G; Malaysia: Kampong: Sinclair 39114; E; Philippines, Luzon: Merrill 738; B; Samoa, Upolu: Whistler 1909; B. – 2. Spikelets are thick and long-haired, inflorescence are short-haired: Philippines, Luzon: Ramos 200/00; K. – 3. Spikelets, inflorescence branches, blades, and sheaths are thick-haired: Australia, Queensland, Middle Percy I.: Lazarides 5617.
Selected Voucher Specimens: Columbia: Dep. del Valle, Alto del Dinde (Cuatrecasas 22940; US); Highland of Popayan (Lehmann 454; L). – Ecuador: Prov. Chimborazo, Huigra (Camp 3176; US); Prov. Guayas, Guayaquil (E. Asplund 16646; G). - Mexico: Jalisco, Mazamitla (US); Jalisco, Zapotlan (Hitchcock 7237; US); Michoacan, Cerro de Carboneras (King & Soderstrom 4837; US); Michoacan, Morelia (Arsene 3115; US); ,Oaxaca, Sierra de San Felipe (C. G. Pringle 4944; M); Rincon del Carmen, Temascaltepec (Hinton 1952; W); Veracruz, Banderilla (Soderstrom 471; US). - Peru: Dep. Piura, Huancabamba (Ferreyra 10876; US). - Socotra I.: Wadi Dilab (G. Schweinfurth 587; P). - Australia: Queensland, Cairns (Helms 1158; US); Queensland, Port Mackay (A. Dietrich 92; B). - India: Bengal, near Calcutta (W. Helfer s. n.; LD). -Indonesia: Kl. Sunda I., Wetar, Klisana (S. Bloemberger 3834; PNH).
var. sylvaticus (Lam.) U. Scholz nov. stat.
Panicum sylvaticum Lam. Encycl. IV: 743 (1798). – Oplismenus sylvaticus (Lam.) Roem. & Schult., Syst. Veg. II: 481 (1817). – Orthopogon sylvaticus (Lam.) Miq., Fl. Ind. Bat. III: 443 (1857, t. p. 1855). – Type: “Ile de France”, Commerson s. n., L. Holotype.
Andropogon undatus Jacq., Coll. III: 237 (1791, t. p. 1789). – Polinia undata (Jacq.) Spreng., Plant. min. cogn. Pugillus II: 12 (1815). – Oplismenus jacquini Kunth, Rev. Gram. I: 44 (1829), nom. Illeg. – Panicum undatum (Jacq.) Steud., Nomencl. 2. ed. II: 264 (1841). – Type: Mauritius, n. v.
Orthopogon pratensis Spreng., Syst. Veg. I: 306 (1824, t. p. 1825). – Oplismenus pratensis (Spreng.) Schult., Mant. II 597 (1824). – Panicum pratense (Spreng.) Steud., Nomencl. 2. ed. II: 261 (1841). – Type: Mauritius: St. Vincent 43, B-WILLD, Lectotype.
Mostly robust grass; Culm 25 to 50 cm high. Blades elongated egg-shaped to oblong lanceolate, 0.5 to 1.2 cm wide, 4 to 8 cm long. Sheath sometimes short-haired on the edges, otherwise bare. Ligule have a hairy collar, 0.7 to 1.1 mm long. Inflorescence compact, racemose; rachis approx. 10cm long, bare. Inflorescence branches angle out, but remain straight.The lengths of the inflorescence branches get smaller higher up. The bottommost is 2.5 to 3 cm long and has 10 to 20 thick consecutive spikelets. Spikelets in pairs, 2.4 to 4.1 mm long. Awns robust, redish, hairless. Lemma I have no awn or are only short and pointed.
Distribution: Mauritius
Selected Voucher Specimens: Mauritius: (E. B. Blackburn s. n.; CGE; Bojer s. n.; M; sieber 13; M; Sieber 205; G).
Oplismenus hirtellus (L.) P. Beauv., Ess. Agrost.: 54 (1812); Adams, C. D., Flow. Plants of Jamaica: 185 (1972); Angely, J., Fl. Anal. e Fintogam. do est. de S. Paulo 6: 1191 (1970); Clayton, W. D. in Hepper, F. N. (ed.), Flora of West Tropical Africa ed. 2, 3, 2: 437 (1972); Hitchcock, A. S., Contr. U. S. Natl. Hebr. 22, 3: 129 (1920); Robyns, W., Fl. Agrost. Congo Belge & Ruanda-Urundi 2: 147, p.p. (1934); Schnell, R., Ic. Plant. Afr. 2: 47 (1953); Stapf, O. in Prain, D., Fl. Trop. Afr. 9: 631 (1934).
| 1a | Inflorescence branches missing or very short (0.3 cm); Spikelets in pairs on the culm or clustered on the inflorescence branches | |
| 1b | Inflorescence branches 0.5 to 3 cm long; Spikelets in pairs, clustered, consecutive thick multi-rowed, or in consecutive pairs | |
| 2a | Inflorescence branches 0.5 to 0.7 cm long; Spikelets in a clustered arragement, 3 to 3.4 to (4.5) mm. | subsp. fasciculatus |
| 2b | Inflorescence branches 1 to 3cm long; Spikelets arranged thickly consecutive, 3.3 to 3.6 to (5) mm long | subsp. hirtellus |
| 3a | Inflorescence branches absent | |
| 3b | Inflorescence branches very short; Spikelets in a clustered arrangement | |
| 4a | Plants delicate; blades up to 2 cm long, lanceolate; Spikelets 2.7 to 3.1 mm long | subsp. microphyllus |
| 4b | Plants delicate to robust; blades 3 to 7 cm long; Spikelets 3.7 to 5.6 mm long; ligules for the most part hairless | |
| 5a | Plants delicate; spikelets 4 mm long, hairy; blades 3 to 4 cm long; East Asia | subsp. psilostachys |
| 5b | Plants delicate to robust; spikelets 3.7 to 5.6 mm long; blades 3 to 7 cm long | |
| 6a | Plants robust; Spikelets in the upper part of the inflorescence in pairs, on the lower part up to seven, sparsely haired or hairless | subsp. acuminatus |
| 6b | Plants delicate to robust; Spikelets always in pairs, always hairless; Ligule glabrous | subsp. capensis |
| 7a | Blades lanceolate; spikelets 2 to 5 on the lowest inflorescence branch (?secondary) | |
| 7b | Blades elongated ovals to oblong lanceolate; spikelets on the lowest inflorescence branch (?secondary), in clustered arrangement | |
| 8a | Blades 2 to 5 cm long; spikelets sparsely hairy | subsp. imbecilis f. imbecilis |
| 8b | Blades 4 to 9 cm long; spikelets shortly but densely hairy | subsp. imbecilis f. lanceolatus |
| 9a | Sheaths and culm long, densely hairy; lower glumes 3-nerved, upper glumes 5-nerved, lower lemmas 7-nerved; spikelets 3 to 7 on the lowest inflorescence branch (?secondary); blades elongate-ovate | subsp. undulatifolius |
| 9b | Sheath and culm usually glabrous, rarely hairy; lower glume (3)- to 5-nerved, upper glume (5)- to 7-nerved, lower lemma 9- to 11-nerved; 4 to 25 spikelets on the lowest inflorescence branch (?secondary) | |
| 10a | Spikelets (2.2) to 2.7 to (3.3) mm long, blades oblong; plants delicate | subsp. setarius |
| 10b | Spikelets (2.8) to 3.2 to 3.7 to (4.1) mm long, blades for the most part elongate-ovate | |
| 11a | Spacing between the inflorescence branches 0.2 cm, seldom more, spacing does not increase on lower parts; inflorescence compact. | subsp. tsushimensis |
| 11b | Spacing of the lowest inflorescence branches 1 to 3 cm, going upward spacing decreases; inflorescence loose | subsp. japonicus |
Oplismenus hirtellus subsp. hirtellus
Panicum hirtellum L., Syst. 10. ed.: 870 (1759). – Orthopogon hirtellus (L.) R. Br., Prod. Fl. Nov. Holl.: 194 (1810). – Typus LINN, fr. microfiche.
Panicum loliaceum Lam., Tabl. encycl. meth. I: 170 (1791). – Oplismenus foliaceus (Lam.) P. Beauv., Ess. Agrost.: 54 (1812), sphalm. Oplismenus loliaceus (Lam.) P. Beauv., Ess. Agrost.: 168, 170 (1812); Ettinghausen, E., Beitrag zur Kenntnis der Nervatur der Gramineae: 413 (1866). – Orthopogon loliaceus (Lam.) Spreng., Syst. Veg. I: 306 (1824, t. p. 1825). – Panicum foliaceum (Lam.) Steud., Nomencl. 2. ed. II: 256 (1841), pro syn. Hippagrostis loliacea (Lam.) O. Kuntze, Rev. Gen.: 777 (1891). Oplismenus compositus (l.) P. Beauv. var. loliaceus (Lam.), Hack. in Stuckert, Anales Mus. Nac. Hist. Nat. Buenos Aires, ser. 3,6: 438 (1906). - Oplismenus hirtellus (L.) P. Beauv. subsp. loliaceus Mez ex Peter, Feddes Repert. Spec. Nov. Regni Veg., Beih. 40,1 A: 219 (1938). - Oplismenus hirtellus (l.) P. Beauv. f. foliaceus Stehle, Carribean Forest., Suppl. 6: 283 (1945), cit. err. (v. Chase & Niles (1962) - "not verified"). - Type: Philippines: Commerson s. n.; n. v. (P?).
Panicum velutinum G. F. N. Meyer, Prim. Fl. Essequ. 51 (1818). - Orthopogon velutinus (G. F. N. Meyer) Spreng., Syst. Veg. I: 306 (1824, t. p. 1825). - Opl ismenus velutinus (G. F. N. Meyer) Schult., Mant . II: 271 (1824). - Type: Essequibo (Suriname: "Sophienburg": G. F. N. Meyer (LE?), n. v.
Panicum oahuaense Steud., Nomencl. 2. ed. II: 260 (1841), nom. nud. - Oplismenus oahuaensis Nees & Meyen ex Steud., Nomencl. 2. ed. II: 220 (1841), pro syn. - Type: Sandwich I.: Meyen s. n., BR, Lectotype.
Oplismenus chondrosioides E. Fourn., Mex. Pl. 2: 39 (1886). - Type: Mexico: Liebmann 367, C, Lectotype; Bourgeau 1668, Schaffner 281 b, n. v., Syntypes.
Oplismenus compositus Bauer, Ill. Fl. Nov. Holl: t. 135 (1813), non Panicum compositum L. - Beleg: Norfolk I.: Bauer s. n., W. Milium undulatifolium Moench, Meth. Pl. 202 (1974), non Panicum undulatifolium Ard. – Beleg: cult. Hort. Marburg, n.v.
Ic.: Fröman, B. & Persson, S., An ill. Guide to the Grasses of Ethiopia: 360, f. 220 (1974). - Hitchcock, A. S., Contr. U. S. Natl. Herb. 22,3: 130 f. 23 (1920).
Perennials, mainly robust plants; Culm 28 - 70(95) cm high. Blades oblong lanceolate but sometimes elongated ovals, (0.5) 0.9 - 1.8 (2.5) cm wide, (2)5 - 12(17) cm long. The midrib is prominently over the base, there are 3 - 5 side nerves on each side, and 5 - 7 intermediate nerves. Sheath is around the edges thick, long and loose, or short haired, in the middle sometimes long and loose, but usually glabrous. Ligules usually have a hairy collar, (0.4)1 - 2(2.5) mm long. Inflorescence compact, racemose, (5) 7 - 18(20) cm long, seldom very hairy, usually short. Inflorescence branches 3 - 10, stand straight up or deviate to the side a small amount. The lengths of the inflorescence branches get smaller, the higher up one goes. The lowest is (0.5)1 - 3 (3.5) cm long and has secundly (8)15 - 20 to (30) spikelets that are thickly arranged in a consecutive fashion. The spacing between the two bottom most inflorescence branches is (1)2 - 4 to (5). Spikelets are in pairs, oblong lanceolate, (2.6)3.6(5) mm long, loosely haired or hairless. Lower glume oblong, (1.6)2.9(5) mm long, (0.9)1.3(1.5) mm wide, (3) to 5 to 7-nerved, with awn. Upper glume oblong, (1.5)2.5(3.7) mm long, (1.1)1.6(2.1) mm wide, (5)7(9)-nerved, with awn. Lemma I oblong, (2.5)3.3(4.3) mm long, (1.5)2.1(2.8) mm wide, (7)9(11)-nerved, with awn. Palea I slender, 2-nerved, or missing. Caryopsis oblong lanceolate, 2 to 2.5 mm long. Awns robust, redish-yellow and hairless. When the fruit become the ripe, the awns become adhesive or sticky. The lengths of the awns are as follows: (7.8) mm long, and Lemma I (0.2) 0.4 - 1.3 (4.3) mm long. |
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Distibution: Tropical regions of the Old and New World.
Common Names: Luhunda (Bahundi language, Zaire); Matunda-Muindu, Muinza (Nord du Mayombe, Zaire); Tudama-dama (Region de Kisantu, Zaire); Lakasi (South Zaire). Tamakoesji (Kar., South America); Herbe à barbes (Martinique).
Ecology: In shady forests and in forests along rivers or streams.
Sociology: Under Aleurites, Tectona, Cola nitida, Albizzaiz; in the Raphia palm grove
Economic importance: Weed in forest plantations
Chromosomes:
2n = 54 : Kammacher et al. (1973); Tateoka (1965).
2n = 43, 57, 60 (63) : fide Kammacher et al. (1973).
2n = 72: Kammacher et al. (1973); Reeder (1967); Tateoka (1965).
2n = 90: Pohl & Davidse (1971).
Notes: There are multiple densely hairy haired forms: 1. Culm, blades, sheaths, and infloresence shafts thick and long-haired: British Honduras, El Cayo Distr.: Gentle 9036; G; Cuba, Bayata: Ekman 3310; G. – 2. Infloresence shafts thick and long-haired: Peru, Pampayacu: Macbride 5026; G; Tonga I., Kao: Yuncker 15,876; G.
Selected voucher specimens: Argentina: Tucuman (Roth s. n.; NY); Dep. Trafi (M. Sillo 3356; G). - Bolivia: North Yungas (O. Buchtien 778; G); Lake Rogagua (White 1203; NY). - Brazil: Baixo Amazonas (Goeldi 104; NY); Braco Joaquim, Mata (Reitz & Klein 3.163; M); 5 km above Labrea (G. T. Prance s. n.; M); Obidos, Prov. Parà (R. Spruce s. n.; OXF); Serra do Caparao (Chase 814; NY); Torres (B. Rambo 45997; B). – British Honduras: Toledo Distr., Cero (P. H. Gentle 6967; G); El Dorado Road (P. H. Gentle 6983; G); Manga Camp (P. H. Gentle 6519; G). - Chile: (Poeppig 2007; G). - Colombia: Santa Marta (H. Smith 2168; G); Dep. Santander, Tona (Killip & Smith 19471; NY). - Cuba: East (C. Wright 751; G), (Poeppig s. n. W). - Dominican Republic: (M. Poiteau s. n.; G); Prov. La Vega (H. C. Allard 14203; B). - b: Guayaquil (Hartweg 705; G); Prov. Pichincha, W from Quito (E. Asplund s. n.; G); Quito (W. Jameson 166; G). - Guadeloupe: (Krauss s. n.; G). – French Guiana: (Leprieur s. n.; G). – Hawaii I.: Oahu, Sacred Falls (O. Degener s. n.; B); Hawai (W. Hillebrand s. n.; B); Maui (G. Spence 138; L). - Martinique: (Sieber 263; PR). Mexico: Cuernavaca (C. G. Pringle 6203; G); State of Morelos (A. S. Hitchcock 423; G); Veracruz (L. Williams 8609; PR). - Nicaragua: Dep. Matagalpa (L.O. Williams et al. 24936; G). - Panama: R. Chagres ( P. N. Allen 4114; G); Paraiso (M. Wagner s. n.; M). - Peru: Dep. Loreto, Tarapoto (PR). - Puerto Rico: Maricao (P. Sintenis 72; G). - Paraguay: Cordillera de Altos (K. Fiebrig 366; G). - Trinidad: (Sieber s. n.; G). – Venezuela: Valle die Aragna (W); Tabay (W. Gehringer 544; G); E from Merida (B. & F. Oberwinkler 13426; M).
New Guinea: W Sepik Distr., M. Amdutakin (C. Kalkman 5308; L).
Ethiopia: Afresa (A. Vatova 693; FI); 5 km from Asbe-Tefferi (16; FI); Amasen, Faghenat (A. Pappi 5326; G); Godjam, Bahar Dar. (O. Sebald 452; B); Mega (R. Corradi 1252; FI); Valle di Mugar Bolè (H. Sonford 42; FI); 9 km N from Jimma (Mooney 6036; FI). - Burundi: (J. LewaIle 4588; G). - Ivory Coast: (G. Roberty 12610; Z); 11 km E from Akribelekrou (G. Roberty 12740; G); Bouroukrou (A. Chevalier 16972; K); Kokondekro (Adjanohoun 322 A; K); 15 km E from Béréby (Oldeman 564; BR); Thiébissou (Leeuwenberg 2091; BR); Touba (8215; K). Gabon: Sibange Farm (M. Dinklage 551; HBG). - Ghana: Kumasi (G. Roberty 12890; G). - Cameroon: (Büsgen s. n.; B); Bipinde (G. Zenker 1214; Z); Buea (F. W. H. Migeod 24; K); Dehane (M. Dinklage 409; HBG); Cameroon M. (Dunlap 66; K); between Makay and Nemeyong (R. Letouzey 11790; P); Esele (T. D. Maitland 1037; K); Mbalmayo (H. Jacques-Félix 9137; P); 40 km S from Ndikinimeki (R. Letouzey s. n.; BR); Victoria (Preuss 1305; K). Kenya: Mara Masai Res. (Bally 5374; G). - Liberia: Boporo Distr. (I. T. Baldwin 10368; K); 20 m from Kakatown (A. White s. n.; K); Nimba (J. G. Adam 20242; K); So (D. H. Linder 1130; K). Mozambique: Manica e Sofala (Garcia 830; BM). - Nigeria: Prov. Benin (J. P. M. Brenan & P. W. Richards 8471; K); Calabar R. (D. R. Rosevear 25; K); lbadan, Gambari For. Res. (C. F. A. Onochie 34949; K); Kabba Road (A. C. Parsons 26; K); Lagos (J. M. Dalziel 1324; K); Mambilla Plat. (J.D. Chapman 2846; K). – Republic of South Africa: Cap Prov. Omcoma (Drege s. n.; P); Bokfontein (A. O. D. Mogg 14271; BR); Kentani (Pegler 2035; Z); Magoebaskloof (B. de Winter 95; B); Natal, Buchanan (Munro s. n.; W); Natal, Durban (A. Rehmann 8649; W); Pondoland (Bachmann 148; Z); Shilovane (Junod 2322; Z); Wilhelmina Gebergte (H. S. Irvine & al. 55750; B). St. Thomas: S. Vicente (J. Espirito Santo 92; BR); Santa Anna (A. Moller 142; K); Boa Estrada (A. Chevalier 14339; bis). – Sierra Leone: Magboloko (R. R. Glanville 99; BM); Lolehun (Katta 27; BR).- Tanzania: Irangi (B. D. Burtt 1572; BM); Kilimanjaro (Geillnger 3897; Z); Kokola, Ngurdoto Nat. Parc (Greenway & Kanuri 11.963; BR); Usambara (C. Holst 458; B). - Togo: Bassar (de Plaen 50; BR); Klouto (de Plaen 57; BR). - Uganda: Bukiga (J. D. Snowden 1213; BM). - People's Republic of Congo: Niari, near Mouyondzi (B. DesCoings 5607; P). - Zaire: Terr. Dibaya (L. Liben 2957; BM); Parc Nat. Albert (J. Lebrun 8114; B); Kisanga (E. Detilleux 567; BR); North Kivu, Virunga Mountain Range (U. Stauffer 409; Z); Marungu (Vanden Prande 282; BR); Nioka Res. (Taton 297; BR); Parc Nat. de l'Upemba (de Witte 3556; BR); Walikale (R. Gutzwiller 2673; BR); Wamba (H. Gallens 4099; BR). – Central African Republic: Mbaiki (C. Tisserant 3459; P). - Zimbabwe: Distr. Inyanga (O. West 4842; BM).
subsp. acuminatus (Nees ex Steud.) U. Scholz nov. stat.
Panicum acuminatissimum Steud., Syn. Pl. Glum. I: 45 (1854). - Oplismenus acuminatus Nees ex Steud., Syn. Pl. Glum. I: 45 (1854), pro syn. –
Panicum balfourii Baker, Fl. Mauritius: 438 (1877), nom. superfl. - Oplismenus hirtellus (L.) P. Beauv. var. acuminatus (Nees ex Steud.) Mez ex Peter, Feddes Repert. Spec. Nov., Beih. 40,1 A: 220 (1938). - Type: India: Wight s. n., W, Holotype.
Panicum (Oplismenus) barbifultum Hochst. ex Schlecht., Linnaea 31: 307 (l861-62). - Type: India, M. Nilagiri, Hohenacker 1279; HBG, Isotype.
Perennial plant, Culm 30 to 50 cm high, hairless. Blades oblong lanceolate, (0.3) to 0.5 to 1 to (1.7) cm wide, 3 to 8 to (12) cm long. Sheath on the edges thick or loose or short and thickly haired, in the middle very rarely long and loosely haired. Ligule often with a hairy collar, 0.4 to 1.1 to (1.5) mm long. Infloresence simply racemose; Main shaft 5 to 10 to (14) cm long, bare. Inflorescence branches are not present, or very seldom under but never over 0.5 cm long. Spikelets stand in pairs, directly on the the main shaft, or up to 7 on short inflorescence branches; spikelet pairs are separated from one another. Spikelets (3.6) to 4.3 to (5.5) mm long, usually hairless. Lower glume lanceolate, (2.4) to 3.4 to (4.1) mm long, (1) to 1.5 to (1.9) mm wide, 5-nerved, with awn. Upper glume lanceolate, (2.4) to 3.2 to (4.5) mm long, (1.4) to 1.8 to (2.1) mm wide, 5- to 7-nerved, with awn. Lemma I elongated, (3.8) to 4.3 to (5.2) mm long, (1.9) to 2.1 to (2.4) mm wide. 7- to 9-nerved, with awn. Awns robust, redish and bare, becomes sticky when ripe; awn from the: Lower glume 4 to 13 mm long, Upper glume 2.6 to 3.2 mm long, and lemma I 0.4 to 1.1 to (1.9) mm long.
Distribution: India, Madagascar.
Notes: There is also a different form that has the following characteristics: blades linear; lowest inflorescence branch 2.2 cm long, with 8 spikelets that are arranged in a consecutive order: Bourbon: 150; LD.
Selected vocuher specimens: Madagascar: Ambatowang (L.); India: (Griffith s. n.; L;); (Wight 1653; CGE); Himalaya (J. J. s. n.; G); Kaity (R. F. Hohenacker 1279; HBG); M. Khasia (J. D. Hooker & J. J. s. n.; G). Nepal: (Wallich s. n.; LD).
subsp. capensis (Hochst.) Mez ex U. Scholz, nov. stat.
Oplismenus (Orthopogon) capensis Hochst., Flora, Neue Reihe 4. Jahrg. 1,7: 114 (1846). – Panicum kraussii - Steud., Syn. Pl. Glum. I: 45 (1854). - Oplismenus africanus P. Beauv. var. capensis (Hochst.) Stapf in Thiselton-Dyer, Fl. Cap. 7: 417 (1899). - Type: South Africa "ad rivulos in sylvis Knysna“, Krauss s. n., W., Lectotype, M., Isotype.
Oplismenus simplex K. Schum. ex Engler, Sitzungsber. Königl. Preuss. Akad. Wiss., Phys.-Math. Kl. 1: 48 (1894), nom. nud. - Oplismenus africanus P. Beauv. var. simplex (K. Schum. ex Engler) Stapf in Thiselton-Dyer, Fl. Cap. 7: 417 (1899). - Type: Siouth Africa: Dohne Mountain, in a wooded Kloot: Galpin 2449, (PRE?), n. v.
Oplismenus bromoides Baker, J. Linn. Soc. Bot. 21: 452 (1885), non ‘Panicum bromoides Lam.' – Oplismenus bakeri Schinz in Dur. & Schinz, Consp. Fl. Afr. V: 771 (1985). – Type: Madagascar: Baron 3213, (K?), n. v.
Oplismenus hirtellus Chippindall, Grasses and Past. of S. Afr.: 362, p.p. (1955), non Panicum hirtellum L.
Mainly a delicate plant; Culm (15)25 - 35(50) cm high. Blades oblong lanceolate - lanceolate, rarely elongated egg-shaped, (0.2)0.4 - 0.8(1.6) cm wide, (2)3 - 7(11) cm long. Sheath along the edges thick or loose and long-haired, in the middle rarely loose and long haired. Ligule with a hairy collar, (0.3)0.5 - 1(1.8) mm long. Infloresence simply racemose; rachis (2)5 - 8(13) cm long, bare. Secondary inflorescence branches not present. Spikelets arranged in pairs, directly on the rachis, very rarely up - 4 spikelets together; spikelet pairs are separated from one another, the undermost 1 - 3(4) cm. Spikelets (3.7)4.5(5.6) mm long, lanceolate, always hairless. Lower glume lanceolate, (2.2)3.3(4.2) mm long, (1.2)1.6 - 1.9 mm wide, 5-(7-) nerved, with awn. Upper glume lanceolate, (2.6)3.4(4.5) mm long, (1.3)1.9 - to (2.9) mm wide, (5-)7-(9-) nerved, with awn. Lemma I oblong, (3.3)4(5.3) mm long, (1.6)2.2(2.6) mm wide, (5-)(7-)9-(11-) nerved, with awn. Lower palea usually missing. Awn robust, redish and bare, becomes sticky when ripe; awn: from Lower glume (5)8 - 10(15) mm long, Lower glume (1)1.5 - 2.2 (5) mm long, and lower lemma 0.4 - 0.7 (2) mm long. |
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Distribution: Tropical and southern Africa.
Ecology: In shady and partly shady rainforests and gallery forests.
Ecology: Under Cupressus.
Anthesis: January to December, usually March to July.
Notes: There is a known form from Madagascar that differs somewhat: Spikelets 5.3 mm long, bare, Lower glume pointed end notched in, bottommost side shaft 1 cm long with 8 spikelets, arranged consecutively. Madagascar: Bojer s. n. – This specimen is known as “Oplismenus rigidus Boj., ad prior. Trin.” A literary description is not known of to the author.
Selected voucher specimens: Cameroon: Cameroon M. (Johnston s. n.; K). - Kenya: M. Abendare (L. Saroglia 223; FI); Kwale Distr. (R. B. Drummond & J. H. Hemsley 3924; FI). - Liberia: Nimba M. (I.G. Adam 20078; K). - Madagascar: Massif de l'Ankaratra (R. Devary et al. 4567; B); South Betsilo (J. M. Hildebrandt 4003; G); Vallée de la Manambola (H. Humbert 13211; G). - Malawi: Nyassa Hochl., Kyimila (A. Stolz 1181; B); near Lake Kaulime (P. J. Tyrer 954; BR) ; Rumpi Distr., Mwenembe (Chapman 109; BM). - Nigeria: Obuden Plat. (Tuley; K). - Zambia: Nyika Plat. (F . White 2715; BR). – S. Tomé: Vanhulst (A. W. Exell 235; BM). - Republic of S. Africa: (Burchell 3159; P); Div. Albany, Grahamstown (F. A. Rogers 27566; Z); Boschberg (MacOwan 1508; Z); Cape of Good Hope, near George (R . Schlechter 2222; FR); Natal, Drakensberg Range (H. Humbert 14886; P); Dlinza For., Eshowe (J. H. Ross 1998; M); Kranskop (R. G. Strey 4813; M); Qudeni For. (Fischer & Schweickerd 116; BR); Swaziland, Pigg's Peak (R. H. Compton 27826; M); Transvaal, Houtbosh (A. Rehmann 5736; .Z); Transvaal, Shilovane (A. Junod 1227; G); Westfalia Estate (J. J. Bos 1139; M); Zoutpansberga. (W. Rauh 3074; M.). - Tanzania: Iringa Distr. Dabage (R. Polhill & S. Paulo 1455; B); Mufindi (S. A. Renvoize & R. A. Abdallah 1929; BR); Bez. Morogoro, Uluguru-Geb. (H. J. Schlieben 3622; BR); Njombe Distr., Mapala (E. Milne-Redhead & P. Taylor 10794; B). - Uganda: West Prov., Bwamba (R. Ross 1097; BM). - Zaire: Elisabethville (H. Humbert. & P. Quarré 16840; P); Parc. Nat. Albert (J. Lebrun 8620; Fl). - Zimbabwe: M'Besa (H. B. Gilliland 1731; 8M); Distr. Melsetter (S. K. Simon & J. F. Ngoni 1340; BM); Vumba M. (R. B. Drummond 5078; BM); Distr. Wedza (H. Wild 6359; BM).
subsp. fasciculatus U. Scholz nov. subsp. – Type: Zaire, Ituri, Semliki à l'Est de Beni: H. Humbert 8993, B, Holotype.
Oplismenus africanus P. Beauv., Fl. d'Oware et de Benin 2: 14 (1810, t. p. 1807). – Panicum africanum Poir., Encycl. Suppl. IV: 275 (1816). - Orthopogon africanus Sweet, Hort. Brit. ed.1: 448 (1827). - Type: Oware et Benin: P. Beauv. s. n., G, lectotype.
Oplismenus brasiliensis Raddi, Agrost. Bras.: 40 (1823). - Panicum raddianum Steud., Syn. Pl. Glum. I: 45 (1854) - Type: Brazil, CorCovado: Raddi s. n., Fl, Holotype.
Oplismenus depauperatus E. Fourn., Mex. Pl. 2: 38 (1886). - Type: Mexico: F. MüIler 2019, NY, Lectotype, Schaffner 207, Galeotti 5847, Syntypi (BR?), n. v.
Oplismenus aristulatus Burcham, Contr. U. S. Hatl. Herb. 30: 419 (1948). -Type: New Britain: Burcham 138, US, Holotype.
Panicum hirtellum Lam. non L., Tabl. encycl. méth. I: 170 (1791). - Oplismenus hirtellus Roem. & Schult., non L. Syst. Veg. II: 481 (1817); Chippendall, L. K. A., Grasses and Pastures of S. Africa: 362, p.p. (1955); Robyns, W., Fl. Agrost. du Congo Belge et du Ruanda-Urundi 2: 147 (1934).
Hippagrostis hirtella O. Kuntze non L., Rev. Gen.: 777 (1891).
Ic.: Jacqes-Félix, L., Les Gram. d'Afrique trop. 1: 244, f. 172 (1962). - Koechlin, J., Flore du Gabon 5: 59 (1963). - Robyns, W. F. Agrost. du'Congo Belge et du Ruanda-Urundi 2: 149, f. 33 (1934).
Diagnosis
Lamina foliorum lanceolata vel ovata-l anceolata, 3 - 10 cm longa; inflorescentia 6 - 12 cm longa, racemi erecti, 3 - 8, prope apicem inflorescentiae reducti, basim 0,5 - (1,5) cm longi; 7 - 15 spiculae racemis basalibus in fasciculatis aggregatae.
Perennial, usually robust grass; Culm (20) to 30 to 50 to (100) cm high, hairy or bare. Blades usually oblong lanceolate, sometimes elongated oval, occasionally loosely short-haired, (0.4) to 0.5 to 1.5 to (1.9) cm wide, (2) to 3 to 10 to (12) cm long, midrib more noticeable, side nerves 2 to 4 on both sides, often with cross nerves. Sheath around the edges loose or thick haired, in the middle seldom loosely long haired. Ligule usually with hairy collar, (0.3) to 0.5 to 1.8 mm long. Infloresence grouped together, racemose, rachis (4) to 6 to 12 to (15) cm long, very seldom loose haired. Inflorescence branches 3 to 8, standing straight up, but distanced from one another; the length of the inflorescence branches gets shorter, the higher up one goes, the bottomost is usually no longer than 0.5 (seldom up to 1.5) cm and has (5) to 7 to 15 to (25) spikelets, clustered in a spherical manner to an oblong clustered arrangement. The spacing of the two bottommost inflorescence branches is 1 to 3 to (4) cm. Spikelets are arranged in pairs, however, sometimes indistinct because they are arranged in such a thickly clustered fashion. Spikelets oblong lanceolate, (3) to 3.4 to (4.5) mm long, usually the 3 bottommost glumes are haired. Lower glume oblong, (1.3) to 2.5 to (4.5) mm long, (0.7) to 1.3 to (1.8) mm wide, (3-) to 5- to (7-) nerved, with awn. Upper glume oblong (1.6) to 2.6 to (3.5) mm long, (0.8) to 1.5 to (2.1) mm wide, (5-) to 7-nerved, with awn. Lemma I elongated oval, (2.4) to 2.9 to (4.5) mm long, (1.5) to 2 to (2.6) mm wide, (5-) to (7-) to 9-nerved, with awn. Palea I 2-nerved, occasionally not present. Caryopsis elongated, plano-convex. Awn robust, redish gold, bare, becomes sticky when ripe; awn from the: Lower glume (4) to 5 to 10 to (12) mm long, Upper glume (1) to 1.5 to 2.6 to (4.8) mm long, and lemma 0.4 to 1.1 to (1.9) mm long. |
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Distribution: Tropical regions in the Old and New World.
Ecology: In shady rainforests, gallery forests, bog forests, secondary forests; in lighted places; in dry forests (Bolivia), Campos (Brazil). On gneiss, quartzite, and sand.
Sociology: Under Agramomum, Tectona, Parkia, Albizzia gummifera, Lantana, Tecla nobilis, Phoenix reclinata; with Conopharyngia holstii and Sapium sp.
Economic impact: Weed in coffee and cocoa bean farms.
Pests: Cladochytrium replicatrum J. S. Karling; Tilletia vittata (Berk.) Mundkur
Anthesis: January to December; usually March to May and October to December.
Notes: There are also some known densely hairy forms as following. 1. Culm thick and long haired, blades thick and short haired, and sheaths loose-long haired: Burundi, Bururi: Lewalle 3501, G; Cameroon: Maitland 83, K; Zaire, Leopoldville: Bequaert7384; BR. – 2. Sheaths and infloresence shafts thick and long haired: Bolivia, Tipuani: O. Buchtien 5331; G. – 3. Blades thick and long haired: Cameroon, Buea: Maitland 853; K ; Dohm 4147; K; Ecuador: Heinrichs 734; M. – 4. Culm, sheath, and infloresence shafts haired, inflorescence branches not present or up to 0.2 cm long, number of spikelets 2 to 6: Zaire: Sita 639; P. The form mentioned in 4. and also some plant specimens found in Togo are a in between form of the the european O. hirtellus subsp. undulatifolius.
Selected voucher specimens: Argentina: Colonia Benites, Chaco (N. Rojas Acosta 17677; G); A. G. Schulz 7361; G}; Dep. El Dorado, San Pedro (A. Schinini & A. Fernandez 5924; ZT); Corrientes, Dep. General Plaz (T. S. Ibarrola 3735; B); Dep. Mburucuyá, Prov. Corrlentes (T. M. Petersen 2677; G); Guayculec, Zormosa (Jörgensen 3098; B). - Bolivia: La Paz valley (C. Troll 1586; M); Mapiri Reg. near Saramipiuni (O. Buchtien 1; M); Prov. Sara, Dep. Santa Cruz (J; Steinbach 5681; G). - Brazil: Rio das Cortas, Prov. Bahia (Martin 19; M); Santa Catarina (Reitz &, Klein 3106; G); Igarapé, Parimé (P. v. Luetzelburg 21495; M); Minas Gereas Serra do Angico (Y. Mexia 5642; G); Corcovada, Staat Rio (P. v. Luetzelburg 6043; M). – British Guiana: Basin of Rupununi R. (A. C. Smith 2368; G); Georgetown (A. S. Hitchcock 16597; G); El Cayo Distr. (P. H. Gentle 8647; G); Toledo Distr., San Antonio (P. H. Gentle 7551; G); Stann Creek Distr. (P. H. Gentle 8159; G): - Cubà: Vento, Prov. Havana (A. H. Curtiss 593;93; M); Prov. Pimar del Rio (E. L. Ekman 12829; G); Prov. Santa Clara, Colonia Limones Prov. (C. G. Pringle 76; M). - Dominican Republic: Provo. Barahona, Lutto bei Rincôn (M. Fuertes 1282; M). - Ecuador: Guayaquil (Jameson 896; G); Prov. Los Rios, R. Plata (E. Asplund 5316; G). - Grenada: Georges (W. E. Broadway s. n.; PR). - Guatemala: Dep. Peten, Tikal (E. Contreras 418; G). - Haiti: Massif de la Hotte (E. L. Ekman 1149; G). – Hawaii I.: Oahu, Nuuanu Park (A. S. Hitchcock 14057; G); Oahu, M. Tantalus (L. D. Whitney 3897; M). Honduras: Dep. Santa Barbara (C. Thieme 5581; M). Jamaica: St. Andrew (C. D. Adams 8335; FR). - Martinique: (Sieber s. n.; B). - Mexico: Fortin (E. Kerber 316; FR); R. Piedros (Hiaram S. n.; B); Tlapacoyan (C. Troll 199; M); Corodba, State Vera Cruz (A. S. Hitchcock 422; G). - Nicaragua: Dep. Matagalpa (L. O. Williams et al. 23893; G). - Panama: Gatun Lake, Canal Zone (A. S. Hitchcock s. n.; G). - Paraguay: Sierra de Amambay (E. Hassler 11992 b; G). - Peru: La Divisora, Huánco (E. Asplund 12591; B); R. Domingo (J. F. Macbride 4251; G). – Sandwich I.:(M. Gaudichaud s. n.; L). – Surinam: Augustus Falls (B. Maguire 24765; G). - Trinidad: (Z. H. Hart 3304 b; G); Port of Spain (A. S. Hitchcock 601; G). – U. S. A.: Louisiana (Engelmann s. n.; HBG).- Venezuela: Cerro Santa Ana (A. Steyermark & A. Braun 94617; B); Chivacóa (Vareschi & Pannier 2570; M). New Guinea: Bernhard Camp, Idenburg River (L. J. Brass 13719; A); Matafuna Bay (P. F. Stevens 50143; A); Monts Point, Willaumez Penn. (J. R. Croft et al. 41383; A); Morobe Distr., Finschhafen (J. & M. S. Clemens 484; BR).
Ethiopia: Eritrea, Rediti (Baldrati s. n.; FI); Kaffa Prov. Jimma (W. Burger 3634; FI); Mega (R. Corradi 1293; FI); Wolieso, Stoa Provo (H. F. Mooney 7611; Fl). - Angola: (J. Gossweiler 5737; BM); (Nolde 671; BM); Distr. Huilla (Welwitsch 2696; BM); R. Cuango (A. W. Exell & E. A. Mendonca 313; BR); Lunda (Gossweiler 13984; B); Ngongola (Wellen 509; BR). - Annobon: Pico del Centro (F. Melville 163; BM). - Burundi: Nyamagana (J. Lewalle 3422; BR); Bubanza (Reekmans 647; BR); Bururi (M. Reekmans 5069; B); Kanyinya (J. Lewalle 5553; G). – Cape Verde I.: Sao Antao (A. Chevalier 45434; P) Cote d'Ivoire: Adiopodoumé (W. de Wilde 1114; K); 5 km N from Béréby (R. Oldeman 504; K); Kokondekro (352 A; K); Bassin du Sassandra et Bassin du Haut Cavally (R. Port~res s. n.; BR); Singrobo (Adjanohoun 317 A; K); Touba (8214; K). – Fernando Po: Moka (A. S. Boughey 56; K); (Vogel 140; K). - Gabon: (M. Thollon 654; P); Eleki (N. Hallé & G. Cours. 6097; P); M. de . Cristal (N. Hallé & J. F. Villiers 4905; P); 10 km SW from Hdjole (N. Hallé 1825; P); Koltang, 30 km E from Libreville (M. G. Gilles 66; p); Mayombe (Le Testus. n.; P); Mouila - Ndèndé (Waltier 6; P). - Ghana: Ankagul (J. B. Hall 3064; P); Ashanti (F. R. Irvine 5080; K); Wacri Tafo (H. E. Box 23; B). - Guinea: (J. G. Adam s. n; P); Vallée du Baffing (H. Pobéguin 1737; P); Kindia (H. Jacques-Félix 2103; P); M. Nimba (R. Schnell 3717; P). - Cameroon: Batanga (M. Dinkl age s. n.; HBG); Eseka (P. Bambs 1317; BR); Douala (A. Meurillon 616; P); Johann-Albrechtshöhe (Staudt 450; G); Nanga (J. Koechlin 7117; P); Sadolkoulay (J. & A. Raynal 12235; B); 10 km S from Ngaoundéré (F. J. Breteler 590; G); 24 km ESE from Nyabesan (J. & A. Raynal 10227; P); Yaoundé Station (Zenker & Staudt 483; K). - Kenya: M. Aberdare (T. Fries 621; BR); M. Kenya (F. White 1355; BM); Nyanza Prov., Kericho Distr. (R. A. Maas Gesteranus 5798; B). - Liberia: West Prov., Boporo Distr. (J. T. Baldwin 12031; K); R. Coss (M. Dinklage 2313; B); Ganta (W. J. Harley 1706; K); Gbarnga (Daniel & Prior 458; P); Nimba (J. G. Adam 20386; K); Sanokwele Distr. (W. J. Harley 1054; K); Vonjama (J. T. Billdwin 9867; K). - Madagascar: N from Maroambihy (H. Humbert 23.368; G). - Malawi: Distr. Rumpi, Nyika Platt (Simon et al. s. n.; BM); Distr. Port Herald (Chapman 2094; BM). - Mocambique: Garuzo (H. B. Gilliland 1807; BM). - Nigeria: Jos Plat. (F. N. Hepper 1071; K); Lagos (O. Olufsen 818; K); Mambilla (M. Oche 75; K); Olokemeji For. Res. (A. Jones et al. 4935; Il); Sapoba (Keay & Onochie 21618; K); Usonigbe (Keay & Onochie 21597; K). - Principe : Terreiro Velho (A. W. Exell 503; BR). - Zambia: 12 km W from Kawabwa, Ntimbacushi Falls (E. A. Robinson 2350; M); Distr. Solwezi, Lunga R. (I. B. Edwards 545; 8M); Victoria Falls (B. K. Simon & R. Hill 2140; P); Banks of Zambesi (F. A. Rogers 207; 13M). – S. Tomé: Lago Amelia (Rose 221; Pl. - Senegal: Stat. Lauma (R. P. Berhaut 2779; 8R); Sikasso (M. Vuillet 506;P). – Sierra Leone: (T. S. Gardner 54; BM); Lester Peak (G. F. Scott Elliott 3854; BM); Njala (F. C. Deighton 1496; BM). –Republic of South Africa: (J. Z. Drege 108; P); Natal, Clavimont (G. Medley Wood 6044; Z); Natal, Durban Prov. (J. Bolus 14648; B); Omcomas (Drege 433 b; Pl. - Tanzania: Korogwe Distr. (H. G. Faulkner 965; B); Mahenge (H. J. Schlieben 1996; G); Morogoro, UI uguru M: (H. J. Schlieben 3797; Z); Usambara (Peter 10027; W). - Tenerife: (Ventenat s. n.; G). - Togo: Near Gapé (F. J. Breteler 7297; B); Sokodé (F. Schroder 159; K); Anié River by Soutouboua (H. Scholz 102; B); NW Palimé, Klouto (H. Scholz 131; B). - Uganda: (Dümmer 311; Z); Ankole (J. D. Snowden 1420; BM); Distr. Bugichu, Bulecheke (T. D. Maitland 1211; B); Masaka (J. D. Snowden s. n.; BM); Distr. Mengo (H. C. Hawkins 428; BM); Sonso R. (W. J. Eggeling 2287; BR). - People's Republic of Congo: Divenie (J. Koechlin 2204; p); 20 km from Sibiti in the direction of Komono (de Nere 1991; P); Brazzaville (A. Chevalier 27362; P). - Zaire: Bambili (Seret 248; BR); Reserve de Djugu (Smeyers 108; BR); Eala (Staner 1434; G); 8 km SE from d'Elisabethville (A. Schmitz 492; BR); Parc. Nat. Garamba (H. de Saeger 1441; BR); between Iwama and Isandja (C. Everard 2847; BR); Kalehe (Pierlot 2702; BR); 11 km de Lumumbashi (J. J. Symoens 12853; M); Kisantu (Vanderyst 299; B); Idjwi I. in the Lac Kivu (H. Humbert 8360; BR); Lukula (R. Devred 1793; BR); Sao Maluku (H. Breyne 2182; BR); Mwene Distr., Prov. Kasai (Dandoy 74; BR); Parc Nat. Albert (J. Lebrun 9862; BR); Yambata (H. Montchal 103; BR). - Central African Republic: (H. Breyne 1082; BR); Bambari (C. Tisserant 429; BM); Bonar (J. C. Bille 2201; P); Dauzat (J. C. Bille 1951; P); Terr. Mahagi (R. Christiaensen 1129; BR); Nola, Mambére R. (G. J. H. Amshoff 7055; BR); R. Kombala, Ouaka (P. P. Tisserant 2317; P). - Zimbabawe: Chirinda For. (A. A. Obermeyer 2169; BM); 40 km S from Marandellas (B. K. Simon et al. 1825; BM); Distr. Mazoe (M. Bingham 1424; BM).
subsp. imbecillis (R. Br.) U. Scholz nov. stat.
Orthopogon imbecillis R. Br., Prod. Fl. Nov. Hall.: 194 (1810). - Oplismenus imbecillis (R. Br.) Roem. & Schult., Syst. Veg. II: 487 (1817). - Panicum imbecille (R. Br.) Trin., Sp. Gram. Ic. II: t. 191 (1828). – Oplismenus compositus (L.) P. Beauv. var. imbecillis (R. Br.) F. M. Bailey, Queensland Grasses: 19 (1888). - Oplismenus setarius (Lam.) Roem. & Schult. var. imbecillis (R. Br.) Benth. ex Hack. in Warb., Bot. Jb. Syst. 13: 259 (1891). – Oplismenus undulatifolius (Ard.) Roem. & Schult. var. imbecillis (R. Br.) Hack., Govt. Lab. Publ. 25: 82 (1905); Hayata, B., Fl. Mont. Form.: 235 (1908); Hsu, K.-S., Fl. Taiwan 5: 566 (1978); Merrill, E. D., Enum Phil. Flow. Pl. 1: 72 (1925). - Type: Australia: Brown 6133, K, Holotype, n. v.; LE, E, Isotype.
Oplismenus undulatifolius (Ard.) Roem. & Schult. var. lanceolatus Domin, Biblioth. Bot. 85: 329 (1915). - Type: Australia; N. O. Queensland: Domin s. n. (PR?), n. v.
Oplismenus undulatifolius (Ard.) Roem. & Schult. var. mollis Domin, Biblioth. Bot. 85: 329 (1915). - Type: Australia: South Queensland: Tambourine Mountains: Domin s. n.; N. S. Wales: Tweed River: Guilfoyle s. n.; Syntypi (PR?), n. v.
Distribution of O. hirtellus subsp. imbecillis f. imbecillis. |
O. hirtellus subsp. imbecillis (R. Brown 6133, LE; Australia; Type). |
Delicate grass: Culm approx. 25 cm high. Blades lanceolate, (0.3) to 0.4 to 0.6 cm wide, 2 to 5 cm long, sparsely haired or hairless. Sheath around the edges short or long haired, bare in the middle. Ligule frayed or short haired in the upper edge, 0.3 to 0.8 mm long. Infloresence uncomplex, racemose, approx. 5 cm long. Rachis bare, inflorescence branches seldom present, the undermost being up to 0.3 cm long. Spikelets in clusters, (2) to 5 to (10) grouped together, oblong, the Lower glume and Upper glume sometimes moderately thickly haired, (2.7) to 3.1 to (3.6) mm long. Lower glume (1.5) to 2.1 to 2.7 mm long, (0.7) to 1 to (1.4) mm wide, 3- to 5-nerved, with awn. Upper glume (1.9) to 2.3 to (2.9) mm long, (1.1) to 1.4 to (1.8) mm wide, 5- to 7-nerved, with awn. Lemma I (2.4) to 2.9 to (3.4) mm long, (1.3) to 1.9 to (2.2) mm wide, (7-) to 9-nerved, with awn. Awn redish, robust, bare; awn of the: Lower glume (5) to 7 to (10) mm long, Upper glume (1.7) to 2.3 to (3.3) mm long, and lemma I (0.4) to 0.7 to(1.5) mm long. Caryopsis elongated oval.
Distribution: Tropical East Asia (especially in the Philippines)
Common Names: Kinswawa (Wahgi language, East Asia).
Selected voucher specimens: Bermuda: Hamilton I (Taylor 49-1162; G). Fiji: Ngau (Smith 7925; L); Ovalan, Lovoni Valley (A. C. Smith 7318; L). - Indonesia: Celèbes, Prov. Minahassa (S. H. Koorder 19790; L); Sunda I., Lombok (J. Elbert 1248; L); Palau, M. Luisualumonogui (T. Hosakawa 6877; l); W: Sumbawa, M. Batulanteh (Kostermans 18307; L); Molukken, Ternate (D. R. Pleyte 19; L); Timor, M. Perdido (v. Steenis 18273; L); Borneo, M. Kinabalu (Clemens 40726; G). - Malaysia: Sabah, Borneo (Noteboom 1415; L). - New Guinea: Morobe Distr. (Clemens 1202; L); Papua, Rambuso (l. J: Brass 2803; L); Subdistr. Popondetta (A. N. Millar 1906; l). - New Zealand: Kermandec I. (W. R. Sykes 10/K; l): - Norfolk Island.: M. Pitt (R. D. Hoogland 6592; L). - Philippines: Bohol (M. Ramos 4297; W); Leyte: M. Jamagdan (G. E. Edana 12050; PNH); Luzon: Mariveles, Prov. Bataan (E. D. Merrill 739; W); Distr. Lepanto, M. Data (E. D. Merrill 4511; W); Rizal Prov., M. Tokduanbanoy (Ramos & Edano 48617; B); Prov. Sorsogon, Irosin (E. D. Merrill 17105; W); Mindanao: Davao (G. E. Edana 11052; PNH); Lake Lanao (Clemens 639; W); Zamboanga del Norte (C. & Ch. Frake 35977; PNH); Tawitawi: Sulu Prov. (Ramos & Edana 44120; W). – Samoa: Falealupo (A. Whistler 105; L): - Society Islands.: (J. W. Moore 62; L); Taiti (Kastolsky s. n.; LE).
f. lanceolatus U. Scholz nov. f.
-Type: Australia: “between Wolumba and Merimbula, S. of Bega”, R. Pullen 4039, L, Holotype.
Oplismenus imbecillis (R. Br.) Roem. & Schult. var. morrisonenis Honda, Bot. Mag. Tokyo 38: 190 (1924). – Type: Formosa: Kawakami & Mori 1845; TI, Holotype.
Diagnosis: Lamina foliorum lanceolata (3) - 4 - 7 - (9) cm longa; spiculae saepe breviter et dense pilosae.
Delicate or robust grass; Blades lanceolate, (3) to 4 to 7 to (9) cm long; Spikelets usually short, thick haired. Distribution: Tropical East Asia (Indonesia – Austrailia). Selected voucher specimens: Australia: (Sieber s. n.; L); Key I. (H. Jensen 112; C); Cape York Penins. (L. J. Brass 19156; G); Queensland, Endeavor R. (v. Mueller s. n.; M); North Kennedy (B. Hyland 04151; L); Mossman Distr., Rumula (Mukee 9091; L). – Fiji I.: Ovalan, Lovoni (A. S. Smith 7318; W). – Maluku: Morotai (H. J. Lam. 3614; L). |
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subsp. japonicus (Steud.) U. Scholz, nov. stat.
Panicum japonicum Steud., Flora 24: 18 (1846). Oplismenus japonicus (Steud.) Honda, Bot. Mag. Tokyo 38: 189 (1924; (Beleg: Japan, Okinawa: Fashiro 214, TI). – Oplismenus undulatifolius (Ard.) Roem. & Schult. var. japonicus (Steud.) Koidzumi, Bot. Mag. Tokyo 39: 302 (1925); Hsu, K.-S., Fl. Taiwan 5: 566 (1978). – Type: Japan: Göring s. n. (P?), n. v.
Oplismenus coreanus Nakai, Bull. Sci. Nat. Mus. 31: 140 (1952), p.p., nom. illeg.
Panicum burmannii Franchet, A. & Savatier, L., Enum. Pl. Jap. II: 160 (1879), non Retz.
Culm: (10) to 35 to (60) cm high, usually hairy in some grooves. Blades egg-shaped oblong, seldom oblong or oblong-lanceolate, (0.3) to 0.9 to 1.5 to (2.5) cm wide, (2) to 4 to 8 to (12) cm long. Sheath thick or loosely long haired around the edges; usually bare in the middle, sometimes loosely or thickly haired. Ligule with or without hairy collar, 0.4 to 1.2 mm long. Infloresence uncomplex to grouped, racemose; main shaft (4) to 6 to 11 cm long, usually bare, occasionally hairy; inflorescence branches not present or very short. Spacing between the bottommost inflorescence branches 1 to 3 cm. Spikelets arranged in pairs, located directly on the main shaft or in clusters of 3 to 10 on the short inflorescence branches (up to 10 cm); laceolate, (2.8) to 3.6 to (4.1) m long, fairly thickly haired. Lower glume oblong, (1.8) to 2.5 to (3) mm long, (0.8) to 1.1 to (1.5) mm wide, 3- to 5-nerved, with awn. Upper glume oblong, (2) to 2.5 to (2.9) mm long, (1.1) to 1.4 to (2) mm wide, 5- to 7-nerved, with awn. Lemma I oblong, (2.9) to 3.3 to (3.8) mm long, (1.5) to 2 to (2.3) mm wide, 7- to 9-nerved, with awn. Pelea I usually not present. Awn robust, redish, bare, becomes sticky when ripe; awn of the: Lower glume (5) to 7 to 13 to (15) mm long, upper glume 1.8 to 5 to (6) mm, and lower lemma 0.3 to 1.7 mm long. Distribution: Japan, China, Korea. Common Names: Chizenai-Sara (rippled bamboo), Chidjimi-zasa (Japan). Ecology: In damp, shady locations; bank of rivers or streams. Anthesis: February to November, mainly September to October. Chromosomes: 2n = 54: Tateoka (1967). Selected voucher specimens: China: Shanhai (Faber s. n.; W); Kiangsi, Lu Shan (H. C. Cheo 289; E); Lokchong (Y. Tsiang 1244; E); Paita (Licent 9649; W); Prov. Shansi, Yao-shan (E. Licent 12786; 101); Shantung Prov., Lung Tung (C. Y. Chiao; E); Yunnan (R. P. Maure 7534; W); Mekong Salween Divide (G. Forrest 14633; W); Yün-shan, Wukang (Handel-Mazzetti 12480; E). - Japan: Annori (U. Faurie 6975;W); Kirifuri, Nikko (J. Matsumura 349; TI); Nikko (C. Schröter s. n.; ZT); Kyoto (I. Ohwi s. n.; ZT); Kyushu (A. Takenouchie s. n.; B); Miyanoskata (J. Bisset 2613; E); Honshu, Prov. Sagami (Yo Momiyama 146; TI); Hakone (J. Matsumura 21; 101); Sapporo (U. Faurie 1260; W); Sendai, Honshu (W); Shizuoka (J. Sugimoto s. n.; B); Tokyo (H. Honda s. n.; TI); Tsoyasaki (A. Takenouchi s. n.; B); Yatsuo, Honshu (So Kirino 381; M); Yokohama (Maximowicz s. n.; W). - Korea: Cheju Do (Taquet 9049; W); Fusan (U. Faurie 815; W). |
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subsp. microphyllus (Honda) U. Scholz. nov. stat.
Oplismenus microphyllus Honda ex Ui, Kishu Shokubutsu Shi: 278 (1929), nom. nud., Honda, J. Fac. Sci. Univ. Tok., Sect. 3, Bot. 3: 274 (1930), descr. – Oplismenus undulatifolius (Ard.) Roem. & Schult. var. microphyllus (Honda) Ohwi, Bot. Mag. Tokyo 55: 546 (1941); Hsu, K.-S., Fl. Taiwan 5: 566 (1978). - Type: Japan, Honshu, I. Itsukushima: Ogawa 6, TI, Lectotype; Japan, Honshu, in monte Kiyozumiyama: Nakai s. n., TI, Japan, Prov. Kii Nachi: Ui 14, TI,
Syntypes: Oplismenus minor Merrill, Phillip. Gov. Lab. Publ. 17: 9 (1904). - Type: Philippinen, Luzon, Mt. Mariveles: Merrill 3203, W, Holotype.
Delicate, small grass; Blades egg-shaped oblong to oblong, 0.3 to 0.9 cm wide, 1.5 to 2 to (4) cm long, bare or covered with a few long, white hairs. Sheath loosely long haired around the edges, in the middle usually bare, sometimes loosely long haired. Ligule usually without a hairy collar or covered with a few short hairs, 0.4 to 0.7 mm long. Infloresence simply racemose, 4 to 5 cm long. Inflorescence branches not present. Spikelets solitary or aranged in pairs directly on the main shaft, separated from one another; lanceolate, 2.7 to 3.1 mm long, very sparsely haired. Lower glume oblong, 2.2 mm long, 0.8 to 1.2 mm wide, 3-nerved, with awn. Upper glume oblong, 2.2 mm long, 1.2 to 1.3 mm wide, 5-nerved, with awn. Lower lemmaoblong, 2.6 to 2.9 mm long, 1.5 to 1.9 mm wide, 7-nerved, with awn. Pelea I not present. Awn robust, redish, bare, becomes sticky when ripe; awn of the: Lower glume 6 to 8 mm long, Upper glume 1.6 to 3 mm, and lower lemma 0.7 to 1 mm long.
Distribution: East Asia.
Voucher specimens: Indonesia: Borneo, M. Kinabalu (J. & M. S. Clemens 40175; HBG). – Japan: Honshu: Itsukushima I. (Ogawa 6; TI); Kiyozumiyama (Nakai s. n.; TI); Prov. Kii Nachi (Ui 14; TI). – Philippines: Luzon: Mt. Mariveles (Merill 3203; W).
subsp. psilostachys (Honda) U. Scholz, nov. stat.
Oplismenus psilostachys Honda, Bot. Mag. Tok. 41: 337 (1927). – Type: Formosa, Buizan-Zyagan: Matuda-Bizi 287; TI, Holotype.
Delicate grass; Blades lanceolate, 0.3 cm wide, 3 to 4 cm long. Sheath long and thickly haired around the edges. Ligule without hairy collar, 0.4 mm long. Infloresence uncomplex, racemose, 7 cm long, bare, inflorescence branches not present; Spikelets postioned singularly or in pairs in the rachis, lanceolate, 4mm long. Lower glume elongated, 2.8 mm long, 1 mm wide, 3-nerved, with awn. Upper glume elongated, 3.1 mm long, 1.5 mm wide, 5-nerved, with awn. Lower lemma elongated, 3.9 mm long, 1.9 mm wide, 7-nerved, with awn. Pelea I not present. Awn robust, redish, bare; awn of the: Lower glume 7 mm, Upper glume 1.6 mm, and lower lemma 0.7 mm long.
Distribution: Taiwan.
Voucher specimens: Taiwan: Buizan – Zyagan (Matuda 287; TI).
subsp. setarius (Lam.) Mez ex Ekman, Ark. Bot. 13: 33 (1913). –
Panicum setarium Lam., Tabl. Encycl. meth. I: 170 (1791). – Oplismenus setarius (Lam.) Roem. & Schult., Syst. Veg. II: 481 (1817); Adams, C. D., Flow. Pl. Jamaica: 185 (1972); Hitchcock, A. S., Contr. U. S. Natl. Herb. 22,3: 126 (1920), North Amer. Fl. 17,4: 308 (1931); Long, R. W. & Lahela, O., Fl. Trop. Florida: 164 (1971); Small, J. K., Man. S. E. Flora: 82 (1933). - Orthopogon setarius (Lam.) Spreng., Syst. Veg. I: 306 (1824, t p. 1825). - Oplismenus compositus (L.) P. Beauv. var. setarius (Lam.) Bailey, Queensland Grasses: 19 (1888). - Hippagrostis setaria (Lam.) O. Kuntze, Rev. Gen.: 77 (1891). - Type: Richard s. n., P-LAM, Holotype.
Orthopogon hirtellus Nutt., Gen. Am. Pl. I: 55 (1818), non 'Panicum hirtellum L.' – Orthopogon parvifolius Nutt., Gen. Am. Pl. 55, in errata (1818). - Oplismenus parvifolius (Nutt.) Kunth, Rev. Gram. I: 45 (1829). Panicum nuttallianum Steud., Nomencl. 2. ed. II: 260 (1841). - Type: Florida (PH?), n.v.
Panicum hirtellum Muhlenb.," Descr. ubi.: 100 (1817), non L.
Setaria hirtella Schult., Mant. II: 276 (1824), non 'Panicum hirtellum L.'.
Ic.: Burkart, A., FI. ilustr. de entre Rios (Arg.) 2: 335 (1969). - Cabrera, A. L., Fl. Prov. Buenos Aires 2: 517, f. 135 (1970). - Fournet, J., Fl. ili. Phanerogam. Guadeloupe: 145 (1978). - Hitchcock, A. S., Contr. U. S. Natl. Herb. 22: 128 f. 22 (1920). - Radford, A. E. & Ahles, H. E., Bell, C. R., Man. Vase. Fl. Carolinas: 131 (1968). - Swallen, J. R., Fl. Guatemala 2: 229, f. 72 (1955). - Wiggins, I. L. & Porter, D. M., FI. Galapagos: 861 (1971).
Usually a delicate plant; Culm 15 to 40 cm high. Blades elongated, seldom elongated-lanceolate, (0.3) to 0.5 to 1 to (1.2) cm wide, 2 to 6 to (8) cm long, sparsely haired or bare. Sheath usually only around the edges thick, long, or short-haired, seldom in the middle loosely long-haired. Ligule with hairy collar, (0.4) to 0.6 to 1.1 to (1.5) mm long. Infloresence usually hairless, (3) to 4 to 8 to (13) cm long, with 3 to 5 to (8) very short (up to 0.3 cm) inflorescence branches. Spikelets grouped together in pairs, clustered in groups of 4 to 10 to (15). The spacing between the two bottommost inflorescence branches is 1 to 2 to (3) cm. Spikelets lanceolate, (2.2) to 2.7 to to (3.3) mm long, moderately thickly haired. Lower glume elongated, (1.4) to 1.9 to (2.2) mm long, (0.7) to 1 to (1.3) mm wide, (3)- to 5- to (7)- nerved, with awn. Upper glume elongated, (1.5) to 2 to (2.4) mm long, (0.9) to 1.2 to (1.6) mm wide, 5- to 7- nerved, with awn. Lower lemmaelongated, (2.1) to 2.6 to (3) mm long, (1.1) to 1.8 to (2.2) mm wide, 7- to 9- nerved, with awn. Pelea I is existent or not present. Awn robust, redish, bare; awn of the: Lower glume (2.7) to 4 to 9 to (10) mm, Upper glume 1 to 2 to (2.6) mm, and lower lemma (0.2)0.4 - 0.7(1.5) mm long. Caryopsis: elongated-oval, approx. 2.5 mm long. |
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Distribution: Subtropical and tropical regions of the New World.
Vernacular names: Running mountain grass, wavy-leaf basket grass (USA); Z'erbe à barbes (Martinique).
Ecology: In moist, shady forests, in coppices along inland lakes, in marshy areas and along rivers and streams.
Sociology: Often present with Erythrodes querceticola (Orchidaceae).
Economic significance: Weed in coffee bean farms.
Anthesis: March to December; usually September to November.
Chromosomes: n = 36: Reeder (1968).
Notes: In west Africa there are few specimens that are most likely coincidental mutations of O. hirtellus subsp. fasciculatus that appear similar to subsp. setarius. These are most likely mutations since it is not reasonable to assume that it is a former floristic relationship.
Selected voucher specimens: Argentina: Prov. Corrientes, Dep. Mburucuyá (T. Myndel Petersen 2678; G); Dep. Mercedes, Colonia Pellegrini (C. Quarin & A. Schinlni 1070; ZT); Isla Martin Garcia (L. R. Parodi 4653; L); San Ignacio (K. Duiroga 47; G). – Barbados I.: Welshman's Hall Gulley (Carter s. n.; CGE). - Brazil: (Sellow s. n.; G); Curitiba, Pinheirinho (G. Hatschbach 8917; B); Morro da Caixa (P. R. Reitz 4.569; L); Obidos, Prov. Para (R. Spruce 50; M); Paraopeba (E. P. Herniger 32501; M); South (Camargo 63691; B). - Cuba: Campo Florido (Leon 4135; NY); Sierra de Nipe, ,R. Piloto (E. L. Ekman 2091; G); Trinidad M. (A. Gonzales 161; FR); Sierra de Vigne, R. Jimbambay (E. L. Ekman 9871; G). – Dominican Republic: (Bertero s. n.; M); Macoris (Taylor 235; NY); Prov. Monte Cristy (E. J. Valeur 482; G); Prov. San Juan (Howard 8741; B). – French Guiana: Cayenne (W). - Guadeloupe: (Bertero s.n.; G), - Guatemala: La Vega (Heyde & Lux 6275; C). - Haiti: Anse Galette (Leonard 3038; NY). - Honduras: Dep. La Paz (Molina 24090; NY). - Jamaica: Hordware (Harris 11843; NY); Kingston {A. S. Hitchcock 600; G); Ewarton at Linstead (A. S. Hitchcock 9415; L); Portland(C. D. Adams 11.537; FR). - Mexico: Valle de Orizaba (Schaffner 163; W); near Mirador, Prov. Veracruz (Satorius s. n.; FR). - Paraguay: Between R. Apa und R. Aquidaban (K. Fiebrig 4231; L); Cordillera de Altos (E. Hassler 3934 a; G); E from Caaguazu (B. Balansa 159 a; L); Dep. Paraguari (Sparre & Vervoost 751; FR); Pilcomayo (E. Hassler 74; G); Yaguason (B. Balansa 2733; L). – Puerto Rico: (Poiteau s. n .; G); Cayey (M. Kuhn 2225; G); Jabucoa (P. Sintenis 4953; PR); Mayagües (P. Sintenis 72 b; M); Ponce (A. A. Heller 6303; L). - Uruguay: Montevideo (J. Arechavaleta 65; ZT); Dep. Treinta y Tres (Herter 1090; M). – U.S.A.: Arkansas: Hulton (B. F. Bush 982; CGE); Florida: Chipola R., Jackson County (Biltmore 2885 a; PR): Jacksonville (A. H. Curtiss 5301; W); St. Johns R. (A. H. Curtiss 3595; FR); Jupiter (A. H. Curtiss s. n.; G); Tallahassee Leon County (V. Nash 2524; G); Georgia: Savannah City Hall, Chatham County (J. Swanberg s. n.; G); Louisiana: (C. W. Short s. n.; PR); New Orleans (C. Baehin 246: G); Missouri: St. Louis (Fista s. n.; L); Ocean Spring (S. M. Tracy 4533; G); Pearl R. County (F. H. Sargent 8310; G); S. Carolina: Cahoun County (Y. F. Logue & J. Bozeman 2133; B); Texas: (F. Lindheimer 210; OXF); Anderson County (I. Palmer 14376; B); Rabb Palm Grove (R. Runyon 3399; M); Rusk County (Vinzent 54; HBG).- West Indies: St. Thomas: Signal Hill (Eggers 174; FR).
subsp. tsushimensis (Honda) U. Scholz, nov. stat.
Oplismenus tsushimensis Honda, Feddes Repert. Spec. Nov. Regni Veg. 20: 360 (1924). – Type: Japan, Kyushu, Prov. Tsushima: Hirata 39, TI, Holotype.
Sometimes a robust plant; Culm hairy, occasionally thick and long, 20 to 40 cm long. Blades elongated-eggshape, or more seldom, elogated-lanceolate, (0.5) to 0.8 to 1.5 cm wide, (3) to 4 to 6 to (10) cm long. Sheath loosely or thick and long-haired around the edges and in the middle, rarely hairless in the middle. Ligule with or without a hairy collar, 0.8 to 1.5 mm long. Infloresence racemose, rachis usually thick, long-haired, 4 to 14 cm long, with short inflorescence branches (up to 0.7 cm long). Spikelets grouped together in pairs, clustered in groups of 4 to 10 on the inflorescence branches. The spacing between the inflorescence branches remains the same the whole height of the plant, usually around 0.2 cm long. Spikelets lanceolate, (3) to 3.2 to (3.7) mm long, always hairy. Lower glume elongated, (1.9) to 2 to (2.3) mm long, (0.9) to 1 to (1.2) mm wide, 3- to 5-nerved, with awn. Upper glume elongated, (1.9) to 2 to (2.6) mm long, (1.2) to 1.4 to (1.6) mm wide, 5- to 7-nerved, with awn. Lower lemmaelongated, (2.7) to 2.9 to (3.4) mm long, (1.6) to 1.9 to (2.2) mm wide, 7- to 9-nerved, with awn. Pelea I usually not present. Awn robust, redish, bare, becomes sticky when ripe; awn of the: Lower glume 6 to 7 to (15) mm long, Upper glume (2) to 2.3 to 2.9 to (4,6) mm long, and lower lemma (0.2) to 0.4 to 1.1 mm long. Distribution: East Asia. Anthesis: September to November. Selected voucher specimens: China: Kwangsi (Tsang 22884; W). - Japan: (Tanaka 16; W); Sendai (Jisiba s. n.; W); Hondo, Ueno Park, Tokyo (J. Ohwi 164; ZT); Tokyo (Matsumura 223; W); Prov. Musashi (Makino 8263; BR).
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subsp. undulatifolius (Ard.) U. Scholz, nov. stat.
Panicum undulatifolium Ard., Animad. Sp. alt.: 14 (1764); Ascherson, P. & Graebner, P., Syn. Mitteleur. Fl. 2: 74 (1898); Ettinghausen, E., Beitrag zur Kenntnis der Nervatur der Graminae: 413 (1866). – Oplismenus undulatifolius (Ard.) Roem. & Schult., Syst. Veg. II: 482 (1817); Bor, N. L. in Rechinger K. H., Fl. Iranica: 485 (1970); Conert, H. J., Gramineae. In Conert, H. J., Hamann, U., Schultze-Motel, W., Wagenitz, G. (ed.), Hegi, G., Ill. Fl. von Mitteleuropa 1 (3): 69 (1980); Hsu, K. S., F1. Taiwan 5: 566 (1978); Komarov, V. L., Fl. U.S.S.R. 2: 34 (1934); Lorenzoni, G. G. F1. Veg. Friuli: 113 (1967); Mayer, E., Blütenpfl. im slowenischen Gebiet: 378 (1952); Oberhofer, E., Der insubrische Vegetationskomplex: 141 (1964); Tzvelev, N. N., Poaceae U.S.S.R.: 655 (1976). - Orthopogon undulatifolius (Ard.) Spreng., Syst. Veg. I: 306 (1824, t. p. 1825). - Orthopogon undulatus Link, Hort. Berol. 1: 202 (1827), nom. superfl. - Type: Arduino s. n., M, Holotype; C, Isotype.
Oplismenus undulatifolius P. Beauv., Ess. Agrost.: 54 (1812), nom amb.
Orthopogon bolosii Vayreda, Cavanillesia 4: 61 (1931), nom. nud. - Type: Spain, Olot: Bolós 5997, W, Holotype.
Oplismenus coreanus Nakai, Bull. Sci. Nat. Mus. 31: 140 (1952), p.p., nom. illeg.
Panicum burmannii auct. non Retz.: Balbis, Misc: Bot. I 8: 80 (1804). - Beleg: Italy, Piemont: Balbis s. n ., C. - Marschall Bieberstein, Fl. Taur. I: 51 (1808).
Panicum hirtellum auct. non L.: All., Fl. Pedemont. 11: 240 (1785); Scop., Del. Florae Faunae insubr. III: 72 (1788), Wulfen ex Jacq., Coll. I: 263 (1787, t. p. 1786). Beleg: Wulfen s. n., W, M.
Ic.: Komarov, V. L., Fl. U.S.S.R. 2: pl. 2, f 10 a-c (1934). - Kiem, J., Ber. Bayer. Bot. Ges. 49: 7 (1978). - Reichenbach, L., Ic. Fl. Germ. I: tb 28, f. 1209 (1834).
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Perennial, sometimes delicate plant; Culm (17) to 20 to 37 to (50) cm high. Blades elongated-eggshaped, seldom elongated lanceolate, short-haired, (0.6) to 0.8 to 1.5 to (2.5) cm wide, (2) to 3 to 9 to (12) cm long, very wavy, midrib very noticeable above the base, becoming finer towards the end, side nerves 3 to 4, intermediate nerves 7. Sheath almost always thickly long-haired in the middle and around the edges. Ligule with or without hairy collar, 0.5 to 1 cm long. Infloresence racemose, almost always thickly long-haired, (3) to 4 to 8 to (18) cm long, with very short (0.5 cm long) or missing inflorescence branches. Spikelets positioned alone or in pairs, arranged in clusters 2 to 7 to (10). Spacing between the two bottommost inflorescence branches (0.5) to 1 to 3 to (6) cm. Spikelets lanceolate, (2.6) to 3.3 to (3.8) mm long, somewhat hairy. Lower glume elongated, (1.5) to 2.1 to (2.7) mm long, (0.7) to 1 to (1.3) mm wide, 3-nerved, with awn. Upper glume elongated, (1.5) to 2.2 to (2.7) mm long, (1) to 1.4 to (1.7) mm wide, 5-nerved, with awn. Lower lemmaelongated, (2.2) to 3 to (3.6) mm long, (1.5) to 1.9 to (2.2) mm wide, 7- to (9)- nerved, with awn. Palea I almost always not present. Awn robust, redish-yellow, hairless, becomes sticky when ripe; awn of the: Lower glume (4) to 7 to 11 to (13) mm, Upper glume (2) to 2.4 to 3.8 to (5) mm, and lower lemma(0.2) to 0.4 to 0.8 to (1.3) mm long. Caryopsis: elongated oval.
Distribution: Southern Switzerland, northern Italy, U.D.S.S.R., Iran, Turkey, India, China, Japan. – Temperate-subtropical zone. [Discovered in U.S.A. in 1996].
Vernacular names Welligblättriger Geradbart (wavy-leaved straight barb), welligblättrige Grannenhirse (wavy-leaved millet awn).
Ecology: In shady chestnut and moist alden forests; between cobblestones (seldom); 0 to 1000 m.
Sociology: Companion plants: Brachypodium sylvaticum, Vinca minor, Salvia glutinosa, Hedera helix, Asplenium sp., Adianthum nigrum, Corylus avellana, Viburnum opulus, Acer campestre, Ulmus campestris, Ostrya carpinifolia, Pinus sylvestris, Picea excelsa.
Anthesis: East Asia: February to November, usually July; Europe: July to November, usually September.
Chromosomes: (Data may refer to specimens of O. hirtellus subsp. japonicus)
2n = 54: Gould & Soderstrom (1974); Koshla & Mehra (1973); Tateoka (1965), (1967).
n = 27: Koshla (1972).
2n = 90: Davidse & Pohl (1974).
Selected voucher specimens: China: Hunan Prov., Ma-Ling-Tung (C. S. Fan &Y. Y. Li 464; G) Kweichow, Kwangsi (Y. Tsiang 648 b; W). - Iran: Asterbad, Bender Ges. (P. Sintenis 1492; G); Bandar-e-Pahlavi (J. & A. Bornmüller 8335; B); North (Butse s. n.; W). - Italy: South Tyrol: Bozen, Kaiserau (Hausmann s. n.; B); (C. v. Hepperger s. n.; W); (Lauter 100; FR); Rodlerau (Hausmann s. n.; B); Valsugana (Faichini s. n.; ZT); Niederlanaer Au by Meran (J. Vetter s. n.; W); Vallée d'Aosta (E. Thomas s. n.; W); Vallesia (Hagenbach s. n.; M); Aquilegia (Putterlick s. n.; W); Cadro (Siegfried s. n.; Z); near Chiavenna, Prov. Sondrio (SDt. Lager s. n.; G); Gardasee (Pittoni s. n.; W); Karfreit (Correns s. n.; M); Gargiano (P. Bigo 770; B); Grantola (Franzoni s. n.; W); Luganersee, St. Margherita (S. Vautier s. n.; G).; Prov. de Novare (L. Terretaz s. n.; G); Lago d'Orta, Piemonte (A. F. Negri s. n.; ZT); Pallanza (A. W. Peipers s. n.; FR); Pavia (G. Sieber-Gyti s. n.; ZT); M. Bracco, Valle du Po (B); Staragora by Görz (I. Glowacki s. n.; B); Stresa, Lago Maggiore (C. Bicknell; B); Montello (P. A. Sanadorz 45; IT); Prov. Torino, Ivrea (Vaccari 1402; Z); area around Turin (Reichenbach s. n.; W); Friaul, Udine (Tommasini s. n.; W); Veneta (R. Pampanini s. n.; G); Veneta, St. Maria di Teletto (Pampanini s. n.; Z). – Japan: Kiusiu (G); Musashi, Tokyo (J. Palacky s. n.; PR); Tokyo (K. Onuma s. n.; ZT); Umgaisha ob. Nikko (C. Schröter s. n.; ZT). - Yugoslavia: Dalmatlen (FR). Korea: Cheju Do (Taquet s. n.; G). - Nepal: Mugu Karnali Valley (O. Polunin et al. 5240; E). - Switzerland: Jardin de Genève (G); Agnuzzo, Luganersee (A. Becherer s. n.; G); Arcegno (P. Aellen s. n.; B); Bellinzona (Mittelholzer s. n.; ZT); Bignasco (G. Kohler s. n.; G); Brusino Arsizio (E. Baumann s. n.; Z); Riva, San Vittale (G. Kummer s. n.; ZT); Cassarate Canyon, Sonvico (W. Koch 54/80; IT); Gordola (Müller-Argau s. n.; ZT); Gudo (D. Rapin s. n.; CGE); Between Locarno and Gordola (E. Fisek s. n.; ZT); Locarno, Madonna del Sasso (St. Arnold 53; Z); Lugano (L. Mari s. n.; Z); Betwwen Lugano and Agno (Cornaz s. n.; Z); Val Maggio (Muret s. n.; Z); Malcantone (E. Streuli 129; ZT); Moesa R. (J. L. Terretaz s. n.; G); Arasio by Montagnola(E. Berger 136; ZT); Musano (Z); Ponte Brolla (M. Rikli s. n.; ZT); Ponte Tresa (F. v. Tavel s. n.; ZT); Sementina (L. Favrat s. n.; ZT); Graubünden: Calania, Misox (G. Walser s. n.; Z); Valle Erono (H. Seitter Sargans s. n.; ZT); Gano (G. Walser s. n.; ZT); San Vittore (H. Seitter Sargans s. n.; ZT); Sorte (H. Kunz s. n.; ZT). - Turkey: Amanus M., Kizil-Dagh, Col de Selderin (M. P. Delbès 39; G); Behizè Lazistan (B. Balansa 1547; G); Giresun (P. Zurse 4133; G); Trabzon (Handel-Mazzetti 215; W). – U.D.S.S.R.: Alazan (Reichenbach s. n.; W); Kaukasus, Gagra (M. Rikli s. n.; W); Jurjewo (G. Woronow 1146; W); Prov. Baku, Distr. Lenkoran (N. Pastuchov 404 b; HBG).
The genus Oplismenus was divided into two sections, according to the position of the awns, as Schlechtendahl (1861-62) had suggested, in Oplismenus sect. Oplismenus and in sect. Scabriseta Schlecht.
Oplismenus or Orthopogon has 160 valid species names and another 56 that belong in this group, but are currently under different generic names. Of this number, 63 are homotypical enough to be automatically added in. Another 74 are ruled out of the genus as nomina excludenda, leaving a remaining 79 species names that can in most cases be assigned through investigation of their types.
In most operating conservatories only a few species are represented or recognized. It is always good to remember that we are discussing an especially polymorphic taxon whose boundaries and definition are merely a matter of man's invention. Ascherson & Graebner (1898) write this in regards to O. undulatifolius: “Most likely only a subspecies or strain of Panicum compositum L., which is widespread in the tropics.” Likewise, Hitchcock (1909) writes the following about O. hirtellus (syn. O. setarius (Lam.) P. Beauv.) only to mention two examples: "It is quite possible that the specimen here included may be referred to distinct species. The type of P. setarium Lam. at Paris resembles Wrights 1543. The blades are short and the spikelets globose and few flowered. Wright 751 and Curtiss 268 and 593 have larger and longer blades and spikes, but some of the other specimens are intermediate." – The large similarities between O. compositus on one side and O. undulatifolius and O. hirtellus on the other are especially large. The taxa do, however, differentiate themselves through the positioning of their inflorescence and the frequent absence of the awn on the lower lemmawith O. compositus. Beyond that the arrangement of the spikelets must be taken into account. With O. hirtellus they are more or less heavily clustered, while with O. compositus they are positioned roughly in rows (s. a. Kap. 7.2.). Stapf (1934) notices a relationship in O. hirtellus between sturdy growth and spikelet size, and also between the length of the awns and the length and number of inflorescence branches on the inflorescence. He also draws upon the species Lamark refers to as Panicum loliaceum attributed its spectrum to Herbarbogen f. robustus. Such correlations are not always a given (i.e. in South America).
The polymorphism of the species most likely misled many authors to come up with new species. – After a critical review of an extensive gathering of all of the last three century's described or named taxa that can be included in the genus Oplismenus the author has come to the conclusion that there are 27 taxa that can be systematically and taxonomically noticed as being different and various classes that they can be ordered into also. The relatedness between these is represented in Fig. 42 and Fig. 43.
Fig. 42. Sect. Scabriseta |
Fig. 43. Sect. Oplismenus |
If one were to make a taxonomic classification for the individual continents, the individual genera for America, Africa, and Europe are relatively easy to keep separate. However, when the worldwide areal of the genus is contemplated the individual taxa in most cases are not so easily separated. Davey & Clayton (1977) come to the same conclusion in their mathematical analysis of the genus. It therefore appears necessary that the most described species be repositioned in the taxonomical classification of species. This process will be described more fully later in the text. The geographical differentiation is mostly carried out through the formation of subspecies. In East Asia, however, a different set of criterion must be used to fully establish the taxonomical classification. Here it is most likely a successive species definition similar to the one described by Grant (1976), in which the individual classifications remained the same. These connect links or intermediaries must lie between the taxonomical elements, seeing that the traits used to differentiate the taxa become partly broken up or replaced with a somewhat modified combination. Grant opposes taxonomical species that are either a biologically founded or an evolutionary species, as claimed by Simpson (1961). In order to be able to establish biological species or types in the genus Oplismenus cross trials had to be performed on living plant material; this is, however, outside the scope of this work and will be left to later studies. As previously mentioned from Grant, the term “biological species” only allows reference to two-parented organisms, while single-parent organisms are better referred to as “evolutionary species”. The definition of an evolutionary species states that a part of the population reproduces separate from the whole and fills its own ecological niche, or rather that it has its “own evolutionary functions”.
Because it is most likely an apomixisical reproduction method that is present in the genus Oplismenus, one could also speak of evolutionary species or types, as long as no intermediary types show a commingling of the population. Even then, however, caution is necessary. For example, if one considers the distribution of O. undulatifolius in the European region it is possible to assume that no comingling with other populations is possible, seeing that the nearest related source is found in the Caucasus, where similar conditions as in the European region exist. In contrast, in the East Asian and Indian regions a comingling with other species or types is possible. It follows, therefore, that the European taxon is only constant in its morphological characteristics because of its isolated distribution and also, therefore, exhibits no evolutionary structure of its own.
Table. Geographic distribution of the taxa
When the special diversity of the genus Oplismenus with reference to the geographical diversity is investigated, one comes to the conclusion that no one continent comes to the forefront as especially preferential.
The genus is distributed in all tropical and many subtropical areas and in the Mediterranean region. Its areal is described by Holub & Jisarék (1968) as austrotropical-pen-meridional.
In Europe, O. hirtellus is represented only by O. hirtellus subsp. undulatifolius.
In the Asian-Australian region one finds four species: O. aemulus , O. burmanii, O. compositus, and O. hirtellus, along with 14 subspecies or varieties (Tab. 2).
In Africa (including Madagascar) seven species are present: O. baronii, O. burmanii, O. compositus, O. flavicomus, O. gracillimus, O. hirtellus, and O. I, along with seven subspecies or varieties (Tab. 2).
In America, four species are present: O. affinis, O. burmanii, O. compositus, and O. hirtellus, along with eight subspecies or varieties
| With respect to diversity, the Asian-Australian region is particularly rich, having 14 taxa. Whether this is the center of evolution for the genus or if the multiplicity of species and varieties is something secondary caused by the varied climate zones that border each other so closely, is indeterminable. Of the 27 taxa that are recognized here, 9 have a wide distribution and 8 a mid-size distribution (usually a mixed areal). The taxa shown in the map on the right have a small, often only dot-sized, distribution. | Taxa with narrow distributions |
Four of the six species of Oplismenus sect. Scabriseta are endemics from Madagascar that are noticeably characterized through clearly defined features, whose development is traceable to their isolated island home.
Another member of Oplismenus sect. Scabriseta is O. affinis, restricted to America, has an inflorescence that is thick and long haired on all parts and closely-spaced inflorescence branches; the two outermost glumes and often also the lowest lemma have either pointed or blunt tips. This is a feature that is not present in any other taxon with such frequency. The species is separated into two varieties. They are present in the same areas but differ in their spikelet length and shape of the glume teeth. There are, however, a whole array of crossovers and exceptions so that a higher ranking taxonomic ranking is not needed. An intermediate form between O. affinis and O. burmanii that is often attributed to O. affinis is found in Haiti. It is a sparsely hairy plant that features distinct notched glumes (Haiti: M. la Cidre: Leonard 7542, MO).
The most widespread species of sect.Scabriseta is O. burmanii. It is easy to identify because of its distinct, generally consistent characteristics that make it easy to identify. Two taxa recognized by the author through their varied hairiness are O. burmanii var. lanatus and var. multisetus. O. burmanii var. lanatus is only present in Java; its spikelets are long, thick, and silver-haired, which is very similar to the hairing of O. affinis. The glumes are, however, not like the aforementioned species', being notched on the end. Nevertheless the variety appears to be a link between the two species.
The connection from the generally clear defined Oplismenus sect. Scabriseta to sect. Oplismenus generates a form that is described by Hooker f. in Trimen (1900) as the species O. thwaitesii. Bor (1960) names the species but has the view that it most likely is only a variety of O. burmanii. Because of the study of the types the following characteristics were noticed: Blades 3 cm long, inflorescence branches 1.5 cm long, spikelets 2.6 mm long, arranged in rows of 6; Lower glume has an awn, Upper glume is pointed, and lower lemmahas no awn. The awn is greenish, robust, and teethed. It is especially easy to notice using the awn arrangement that we are dealing with a hybrid between O. burmanii and O. compositus. The occurrence of hybrids or a tendency towards hybridization between these two species is shown also through the following sources: Java, Batavia: Schiffner 1520, L; Buitenzorg: Winterbotton s. n.; K; West Java: Preyer s. n.; B. In these sources there are 2.5 to 3 cm long inflorescence branches, spikelets arranged in rows, sparsely toothed awns, and toothless lemma. Another source (India: Heifer 558) has delicate and whitish, but toothless awns.
Oplismenus sect. Oplismenus, according to my taxonomic treatment contains 3 species, namely O. hirtellus, O. compositus, and O. aemulus . All three species include multiple variants that other authors have separated into different species. A grouping into subspecies is unsustainable seeing that numerous crossover and intermediae forms exist, making all species classificationa synthetic or unnatural or, in other words, not very meaningful. Because the species classification process in the genus Oplismenus, that is Oplismenus sect. Oplismenus, is not finished but in progress, the grouping represented here seems partly artificial because it too has imtermediates between the species recognized. Nevertheless, the taxonomy presented here is still meaningful because it makes it possible to place most plants without difficulty and it reflects geographic differences.
The most important species in Oplismenus sect. Oplismenus is O. hirtellus. The name, however, is used by different authors for different groups of plants. At one extreme, it includes only plants with short inflorescence branches, specimens with long branches being referred to O. loliaceus (Lam.) P. Beauv.; at the other extreme, plants with both long and short secondary inflorescence branches are included. On the American continents the taxon with large inflorescence branches is known as O. hirtellus, and the taxon with short inflorescence branches as O. setarius. Actually there are three different forms.
One must be able to decide which of the species names is nomenclaturally correct. The type material from Panicum hirtellum L. was investigated from a microfiche in the Linné herbarium. Using this source one finds a version with distinct long inflorescence branches. We can then conclude that Panicum loliaceum Lam. is only a synonym for Panicum hirtellum L. If we were to take the form with short secondary inflorescence branches that comes mainly from Africa and separate it taxonomically from the form with long secondary inflorescence branches, name changes would be needed. The long-branched version is distinctly set apart by the name O. hirtellus (L.); I have given the short branched plants a new name: O. hirtellus subsp. fasciculatus. This taxon is basal in O. hirtellus, as many other taxa branch off of it, including the above mentioned O. loliaceus, which is from now on will be referred to as O. hirtellus subsp. hirtellus. Also branching off of O. hirtellus is the taxon O. hirtellus subsp. setarius, which is present in Africa in small quantities but has its primary distribution on the American landmass.
The type of O. hirtellus subsp. fasciculatus is distinguished through its short inflorescence branches (the lowest being 0.5 - 1.5 cm long) and clustered spikelets. On the lowest inflorescence branch there are (7) to 15 spikelets. The inflorescence branch and the sheath surface are usually glabrous. The outer glumes are 5- to 7-nerved; the bottom lemma is 9-nerved. O. hirtellus subsp. fasciculatus comes predominantly from and is very common in tropical Africa. On the other hand, in South America the predominant form is O. hirtellus subsp. hirtellus, which is distinct with its longer inflorescence branches and different spikelet arrangement. In Asia O. hirtellus subsp. fasciculatus is seldom found with such characteristics as mentioned above. One can trace multiple development pathways of this subspecies, namely in the direction of: 1) O. hirtellus subsp. japonicus, 2) O. hirtellus subsp. capensis, 3) O. hirtellus subsp. setarius, and 4) O. hirtellus subsp. hirtellus.
Let's consider next the evolutionary path to O. hirtellus subsp. japonicus. This subspecies is found in the tropical regions of East Asia and is set apart by its short or nonexistent inflorescence branches, a diminished number of spikelets (2 to 6), and often its diminished number of nerves (3/5/7) in the area belonging to the glumes. In some rare cases one of these characteristics can be found in O. hirtellus subsp. fasciculatus on the African continent – there is, therefore, no possibilty of clearly distinguishing O. hirtellus subsp. japonicus from all other taxa. Developmentally it is most likely a crossover form.
As shown in Fig. 21 [needs to be verified in the original; it is the distribution of O. aemulus var. densiflorus] there are multiple forms that come from the developmental link between O. hirtellus subsp. and O. hirtellus subsp. japonicus, the most important being the newly recognized O. hirtellus subsp. undulatifolius. This taxon, whose main distribution is in north Italy and the subtropical area the Black and Caspian seas, exhibits in these geographical areas a strong, consistent pattern of characteristics. The inflorescence shaft, sheath surface, and also the sheath edges are always densely long-hairy. The inflorescence branches are very short or nonexistent. The spikelets are clustered in groups of 2 to 5, and an individual spikelet measures (2.6) to 3.3 to (3.8) mm long; the outer glumes are always 3- to 5-nerved and the bottom lemma is 7-nerved.
In the examination of this plant's range, O. undulatifolius has always been overlooked; the typical features of the European material are disappearing and in their place distinct characteristics of O. hirtellus subsp. japonicus have emerged. According my investigation O. undulatifolius is found in the East Asian region with the typical characteristics only when totally isolated, usually, however, one or more of the characteristics are mutated and the plants show a tendency to O. hirtellus subsp. japonicus. Examples of this can be found in the following sources: India, Suala: Duthie 10136, W (Nervature of the glumes: gl I: 5, g II: 7, l I: 9; bottommost side shaft 1 cm long); Iran, Andavar: Wendelbo & Assadi 14566; E (Plant robust, spikelets 3.7 mm long inflorescence branches 0.7 cm long, main shaft and sheath surface hairless).
Several African specimens that resemble O. hirtellus subsp. undulatifolius in many respect descend most likely from a chance mutation, and are, therefore, converging forms seeing that such a mixture of an African and European population appears to me to be highly unlikely. On specimen, Italy: Lange s. n., features a 2.5 cm inflorescence branch while all other characteristics and all other inflorescences clearly resemble O. hirtellus subsp. undulatifolius. It follows, therefore, that also the characteristic of inflorescence branch length varies and is not a good taxonomic character.
A discrepancy regarding nomenclature of this taxon is referred to by Smith (1979), who did not consider the combination based on Panicum undulatifolium Ard. with Oplismenus or Orthopogon as valid, because in the work of Ardino the binary nomenclature was not consistently used. Wilmott (1935) mentions that the binary nomenclature was used only unclearly by Ardino in the first edition of the work. In the second edition from 1764, where Panicum undulatifolium is described, it is consistently used.
From O. hirtellus subsp. japonicus another taxon is derived, this one found only in East Asia: O. hirtellus subsp. tsushimensis. It is found only in Japan and is distinguished from O. hirtellus subp. japonicus through the strong approximation of its inflorescence branches and spikelet clustering. Thereby the inflorescence functions almost like a homogeneous spike or compact raceme. Crossover forms toward O. hirtellus subsp. japonicus are also encountered. On the inflorescence brancht, an otherwise O. hirtellus subsp. tsushimensis conforming plant, the spacing between the two bottommost spikelet clusters measures 1cm (Japan, Nikko: Tateoka s. n.; E).
A third complex that deviates from O. hirtellus subsp. japonicus contains three new taxa first recognized by the author: O. hirtellus subp. imbecillis, O. hirtellus subp. microphyllus, and O. hirtellus subp. psilostachys. These taxonomic classifications were necessary since the taxa posess different morphological characteristics from the above mentioned subspecies, but are nonetheless related to them through crossover forms. In the case of O. hirtellus subp. imbecillis there is a geographic distinction with a center on the Philippines; determining such are area is not possible with the other two taxa because they are only known from the type specimens.
O. hirtellus subsp. psilostachys resemble O. hirtellus subp. imbecillis in habit but differs in having 4 mm long spikelets. This is another example that large spikelets are not always necessarily coupled together with robust growth.
O. hirtellus subsp. imbecillis is distinguished by its especially delicate habit, small lanceolate blades, and fewer inflorescences. However, some specimens of O. hirtellus subp. imbecillis, mainly from Australia, are somewhat more robust and have longer, similarly strong lanceolate blades and mainly shortly thick-haired spikelets.
O. hirtellus subsp. microphyllus, a rare taxon, is distinguished by its very small blades (none being longer than 2 cm) and spikelets that are arranged on the inflorescence branches either alone or in pairs. It is possible that O. hirtellus subp. microphyllus is either only a distressed form of O. hirtellus subp. imbecillis or it is simply a very young taxon. Between these two closely related subspecies intermediate forms are found, which possess one or the other distinguishing features, but as a whole do not allow for a distinct special or sub-special separation: (Malaysia, Sarawak: Burtt 8339, E; Philippines, Luzon: Ramos & Edano 45004, G; Japan, Sagami: Mizushima 10107, C).
We will now trace the second developmental line of O. hirtellus subsp. capensis. This taxon, which must have been formed as a result of geographical differences, is found mainly in the subtropical areas of South Africa. Outside of that it, can also be found in Madagascar, tropical central Africa, and sporadically in West Africa, where it mixes with O. hirtellus subsp. fasciculatus. O. hirtellus subsp. capensis is characterized by its always hairless ligules, nonexistent inflorescence branches, and hairless, 3.7 to 5.6 mm long spikelets that are arranged either alone or in pairs on the main shaft. If these features were consistent in overlapping areas, then it would be recognizable at the species level. This is, however, not the case and that is why it appears most likely to be a sub-species with geographical differences. Intermediary forms are also frequently found; for example, the following sources: South Africa: Luxton 414, Mo; Natal: Harrison 428, M; Cap. Prov.: Drege s. n., G; Duiwelskloof: Bos 1139, B; Mozambique, Beira: Torre 4018, BM; Zaire, Lac Kivu: Humbert 7425, P; Gabon, Franceville: Descoings 6584, P; u. a. m.
The intermediate forms are usually recognizable by their numerous spikelets, singular, longer inflorescence branches, or thicker haired ligules.
This developmental line is traceable from Madagascar by way of Mauritius to India, where a mutated form O. hirtellus subsp. capensis is found. This form is known as O. hirtellus subsp. acuminatus. Common also to this subspecies are large and only rarely hairy spikelets and greatly diminished inflorescence branches. In contrast to O. hirtellus subsp. capensis the number of spikelets in the bottom part of the inflorescence is usually 5 to 7 per cluster. The geographic relationship, involving South Africa and India is noteworthy, as it is supported by Wegner's continental drift theory.
We will now discuss the developmental branch of O. hirtellus subsp. hirtellus. This taxon is distinguished by its usually strong growth, longer inflorescence branches, and thick, multi-rowed, consecutively arranged spikelets. In Africa as well as in America there are intermediary forms in the direction of O. hirtellus subsp. fasciculatus: Bolivia, Santa Cruz: Steinbach 7010, G; Martinique, M. Parnasse: Hahn 147, G. Sometimes there are also intermediary forms found between O. hirtellus subsp. hirtellus and subsp. capensis: Zimbabwe, Umtali: Crook 880, M; Madagascar: Vignier and Humbert 1530, B: inflorescence branches 1 cm long, spikelets hairless and arranged in rows of six, ligules hairless.
Another taxon, this one widespread in America, O. hirtellus subsp. setarius should be viewed as a fourth developmental line. These are delicate plants with very short inflorescence branches and clustered spikelets that are mostly approx. 2.7 mm long, bunched together in groups of 4 to 10 to (15). The American version is partly identified as being O. undulatifolius. O. hirtellus subsp. setarius differentiates itself from this taxon by having no hair on the inflorescence shaft and sheath surface, the venation of its glumes (Gl I: 5, Gl II: 7, and L I: 9), and larger spikelet clusters. However a specimen was found in the Dominican Republic that distinctly resembles O. hirtellus subsp. undulatifolius: Gl I 3- to 5-nerved, inflorescence shaft thickly haired: Ritter s. n., W. It appears to be a mutation in the direction of O. hirtellus subsp. undulatifolius. One could, however, also assume that such taxa can develop in multiple places. The presence of O. hirtellus subsp. setarius in West Africa with unique feature similarities with the American plant also supports this view: Benin: Cotonou: Froment 1052; BR; Cameroon, Dehane: Dinklage 377, HBG. – With the close relations and the few criteria we, therefore, come to the conclusion that a polyphyletic developmental path of the species or subspecies is not to be ruled out.
We turn now to the second most widely distributed species, O. compositus. This species distinguishes itself from O. hirtellus by its longer inflorescence branches (up to 10 cm) and the absence of an awn on the lower lemma. There are three distinct varieties: O. compositus var. compositus, var. rariflorus, and var. sylvaticus. O. compositus was mentioned up to this point for only its primarily Asian distribution. It could, however, be possible that it is also present in America. This is also how O. rariflorus was described. However, as previously mentioned, it is also distributed throughout Asia. Both taxa distinguish themselves through the length of their spikelets and their differing habits.
The third taxon we need to discuss is O. compositus var. sylvaticus. It is found only in Mauritius and sets itself apart with shorter inflorescence branches (the lowest is 2.5 to 3 cm long) and more compactly positioned spikelet pairs than are found in the other two varieties. This taxon also resembles O. hirtellus subsp. hirtellus, but differs mainly by not having an awn on the lower lemma. In spite of this, this taxon appears to be a connecting link between the two species, justifying in this case a special separation. In all the remaining areas of distribution the species are easy to identify, making it advisable to keep this classification, in the which I concur with the opinion of Davey and Clayton (1977).
The third species in this section, O. aemulus, is very similar to O. compositus and also appears to be a connecting link to O. hirtellus. It is, however, very different and needs to remain in a separate taxon from O. compositus var. sylvaticus. Because of its large variability, it must be divided in three varieties, namely O. aemulus var. aemulus, var. flaccidus, and var. densiflorus. Accordingly, essentially the same treatment as Domin (1915). He used the varietal epithet “pilosus” for O. aemulus, which is distinct from the form referred to by the author as O. aemulus var. densiflorus, which is only found in New Guinea. I have not been able to trace the epithet “pilosus” used by Domin.
O. aemulus var. aemulus is distinguished by its distinct inflorescence branches (0.5 to 3.5 cm long), upon which 2.8 to 3.2 mm large spikelets are arranged consecutively, either in pairs or frequently alone. The lower lemma usually has no awn and the whole inflorescence is densely hairy. In contrast, O. aemulus var. flaccidus has glabrous inflorescence rachises and inflorescence branches, as well as usually only sparsely hairy spikelets. The species O. aemulus and O. flaccidus are mentioned in many studies as synonyms for O. imbecillis. However, they can be distinguished by the length of their inflorescence branches and the presence of awns. Sometimes, though, intermediary forms are found. Their growth habit is delicate, their blades are small and lanceolate and their lower lemmas are unawned: New Guinea, Papua: Carr 11848, L; Australia, N. S. Wales: Pullen 4039, G.
The third variety of O. aemulus is O. aemulus var. densiflorus from New Guinea. In this taxon the inflorescence branches of the inflorescence are very close together and long hairy like the rachis. The awns on the three lower bracts are, however, the most distinct of all the taxa. The awn of the lower glumes is 1.5 to 3.5 mm long and the awn of the upper glume is 0.2 to (1.1) mm long; the lower lemma has no awn. There is a unique population found in Africa that is very similar to O. aemulus var. densiflorus. The inflorescence branches are also close together, the spikelets are small (2.6 mm long), and have rather short awns: Africa: Richard s. n.; Zaire: Hens 157.
13.5. A comparison of the taxa in tropical and extra-tropical areals
We will now investigate whether the taxa of the genus in the tropical regions are morphologically different from those of the extra-tropical regions. The boundaries of the tropical areas are shown on the right, as defined by Küchler (1961) based on climate and vegetation maps. When one compares the boundaries depicted with specific Oplismenus taxa, then it becomes apparent that the taxa shown in the table below as “Tropical Taxa” have a purely tropical distribution. The taxa with a purely subtropical distribution include O. hirtellus subp. japonicus, subsp. tsushimenis, and subsp. undulatifolius. In contrast, the taxa in the second part of the table below are either primarily tropic or primarily subtropical but, in each case, are also found in the other region. The variability of the taxa is quite large in the tropical as well as the subtropical regions for the four characteristics shown. A tendency for the subtropical taxa to have short or nonexistent secondary inflorescence branches is evident in the table below, but the tendency is also present throughout the tropical taxa (O. hirtellus subsp. microphyllus, subsp. imbecillis). Thus we cannot say that in the subtropical regions the taxa are mainly delicate, small bladed, and small awned - meaning that these taxa are no more than “troublesome” plants that grow in the tropics under better ecological and climatic conditions, as Schlectendahl (1861-62) suggested based on cultivation studies. |
Tropical region according to Küchler (1961).
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Characteristics of tropical and subtropical taxa
All the taxa in Oplismenus, other than O. hirtellus subsp. undulatifolius, all have a more or less continuous distribution range. The boundaries of both tropical and subtropical areals are fluid and indicate a slow migration to larger regions. In contrast, the disjunct distribution of O. hirtellus subsp. undulatifolius is worthy of not of note. This taxon is found more or less only in subtropical regions of Eurasia. Its focus of distribution in Europe is in the southern parts of Switzerland and northern Italy where it has been found in the valleys of the Maritime Alps (Aosta), in Ticino and Graubünden, in South Tyrol near Bozen and Meran, around Lake Garda, in the area around the Po River, and in the East Italian area near Friaul. Literature records report locations on Istria (Schlosser & Farcas 1869). It appears that Degen (1936) investigated the original discovery locations and discovered that nowadays the taxon is not present at any of the given locations. The sources from the South Tyrol area are from the beginning of the century and it was no longer clear, whether the subspecies was extinct there or not. Kiem (1974) found evidence that spoke towards the species continued existence. He found that the most of original areas of growth had suffered some sort of city building, but he could report (1978) to have found an area of recent growth near Burgstall.
The southern European map depicting the taxon's distribution was authored using data from another map made by Pampini (1903) and written entries from Becherer (1966). Outside of this area, some specimens of O. hirtellus subsp. undulatifolius were found by Vayreda (1931) in northern Spain, Olot, Prov. Gerona and Becherer (1966) in the Swiss canton Geneva, Chancy, northern part of the Bouchet forest. Both specimens are most likely not from natural distribution. Becherer (1966) reports that in 1963 and 1964 multiple smaller growth colonies were found in the areas mentioned above, and that these plants could not naturally be found there, seeing that these areas are frequented by botanists from Geneva and they would have long ago discovered these colonies.
In Quatre Flores de France P. Fourier (1961) mentions the presence of the taxon on the Italian Riviera. It has not, however, been found in France.
O. hirtellus subsp. undulatifolius has another relatively well defined areal in the Caucasus Mountains with distributions in Turkey (near Rize and the Amanus mountains) and in northern Iran as well (Fig. 41). The distribution data for this area was filled in by a map authored by Grossheim (1939). Outside of these areas specimens of this subspecies have only been found in India, China, and Japan. These finds, however, do not make up a separate areal, but are rather isolated cases. The eastern open-end of this areal overlaps with the areal of O. hirtellus subsp. japonicus.
The following remarks are aimed towards explaining the disjunct distribution of O. hirtellus subsp. undulatifolius. The European distribution is neither tropical nor subtropical, so there must be some reason why a single taxon of a genus that normally grows only in tropical and subtropical regions is found here. Accordingly, two possible hypotheses can be conceived: 1) O. hirtellus subsp. undulatifolius is a Tertiary Relict, which are sometimes present in the Alps in other relationship circles. 2) The taxon migrated or was brought here after the Ice Age and established itself in places with a suitable climate.
Braun-Blanquet (1923) gives some criteria for determining whether a species is a tertiary relict. The species are systematically isolated, therefore there are not derivatives of a related species. Tertiary Relicts are very ecologically adapted but are rather one-sided. They have lost the abilities to broaden their distribution and diversity, leading to a dividing up of the distribution. Oftentimes these conditions lead to a disjuncted distribution. Wettstein (1896) brings to our attention that Tertiary plants are very fragmentarily distributed and usually consist of only one species. In some areas, seeing that there was no Ice Age since this initial Tertiary adaptation, an unbroken developmental chain has been possible (i.e., in the regions of East Asia), and many different taxa have come into existence. Engler (1879) and Gams (1936) now see a relationship between the European Tertiary Relict and the Colchian regions (Caucasus), Himalaya, and Japan. Their connecting links, however, were broken by the Ice Age so that nowadays the Tertiary plant is found only in these same areas where the species originally sought refuge from the worsening climate. Wettstein (1896) viewed certain specimens that grow in the Alps along side other species as evidence of this Tertiary adaptation. The plants related closest to them are found in tropical areas.
Before we evaluate O. hirtellus subsp. undulatifolius according to these criteria, a general description and investigation of the Insubric climate of its European distribution will be given. This area contains both Mediterranean and oceanic zones. The amount of precipitation this area receives is considerably higher than most parts of Europe and especially more than the Central Mediterranean region. Hofer (1967) reports an average precipitation amount of 1300 to 2000 mm per year. Braun-Blanquet (1961) reports 950 to 1100 mm precipitation per year for the morainal region around Ivrea and 2000 mm for Ticino (in extreme cases, even up to 2500 mm). Brockman-Jerosch (1913) reports 1700 mm precipitation per year for Locarno and Pitschmann & Reisigl (1965) report 1441 mm for the Lake Como and 1500 to 2000 mm for Lake Lugano. According to their data the area around Lake Garda also receives high amounts of precipitation. Also the average temperature for July in Lugano is reported as being 21.5 °C, and the yearly average as 11.4 °C. As is apparent, in this area there are mild winters coupled with high amounts of precipitation. There is also a low level of both cloudiness and fog. Hofer (1967) reports that Locarno has the highest actual and relative amount of sunshine for all of Switzerland. He states that good climate in southern Switzerland is due to the shielding provided by the Alps from cold air masses from the east and north and the inability of these same cold air masses to form there due to the relief-dependent air circulation.
One could also assume that O. hirtellus subsp. undulatifolius grows in places in Europe that have a nearly subtropical mini-climate. If this were the case, a distribution according to purely ecological criteria would also seem logical. In order to discover the possible beginning point of the Tertiary Relict, we will compare the taxon with a map authored by Szafer (1975) that details ecological tertiary areas of refuge. Upon viewing this map it is clear that the only area of refuge on the European continent is on the Balkan Peninsula. Other areas, these being outside of Europe, include the Caucasus, between the Black and Caspian Seas, in Central Asia, and in East Asia. Additionally, Wettstein (1896) lists other refuge areas in northern Spain and in the lowlands around the Alps.
O. hirtellus subsp. undulatifolius meets some of the requirements out lined by Braun-Blanquet (1923) for a plant to be termed a Tertiary Relict: No related taxa are found on the European continent and its characteristics vary little, which points toward an older range of plants. Also, the previously mentioned ecological specialization and adaptation are present, since larger populations of the subspecies are found only in damp, wet forested areas. The amount of precipitation must be very high. – The ability to enlarge its distribution, however, has not completely disappeared; Becherer (1966) gives report of the discovery of new specimens in a forest near Geneva which were observed over several years and showed an increase in population. Also in South Tyrol have other such new discoveries been made. How intact the northeastern Spanish areal is today is determinable. Far and wide, though, the distribution and the taxon itself are largely not expanding and it is therefore a possible candidate for recognition as a Tertiary Relict.
In Gam's map (Gams 1936) the zones that remained free of ice are pictured on the northern and southern boundaries of the Alps. The glaciers that entirely covered both the Ticino and South Tyrol valleys are easily recognized. It follows then that the two modern areas which are the main places of distribution for O. hirtellus subsp. undulatifolius were, according to Gams, entirely covered with glacial ice. The glacation of the area around Lake Garda is depicted by Pitschmann & Reisigl (1959). Here, also, there remain only a few, small square kilometer sized areas free of the glacial ice, but Pitschmann (1965) declares the areas around Bresciano and Bergamo in the Alps as having permanently remained free of ice. According to these findings one must either wholly reject or at least strongly doubt the hypothesis that O. hirtellus subsp. undulatifolius is a Tertiary Relict. If we use the definition given by Szafer, the taxon is not actually a Tertiary Relict, since the ice free areas do not coincide with the modern distribution. There is also just as little evidence to suggest that the population migrated out of its original area of distribution into areas of refuge from the worsening climate, and then afterwards returned to its modern area of distribution. In this case there would surely be leftover populations located in the Tertiary refuge areas (as suggested by Szafer (1975)), i.e. on the Balkan Peninsula; which is not the case.It is more likely then that the present-day distribution traces back to a post Ice Age migration of the taxon, which came out of East Asia, passed through the ecologically favorable areas of the Caucasus, and found root in northern Italy. It should, however, be made clear that the question cannot be totally settled due to a few of the above mentioned facts (isolated locations in the ecological system, disjunt areal, and ecological specialization), which all speak in favor of the taxon being termed a Tertiary Relict. Seeing that there have been no direct studies on samples of pollen finds conducted, the above mentioned ideas connected to dealing with the investigation of this question must suffice.
Hekaterosaehne elatior Steud., Syn. Pl. Glum. I: 118 (1854). - Type: New Zealand; n. v. Notes: According to the original description, this is most likely not an Oplismenus species.
Oplismenus compositus (l.) P. Beauv. var. brachyphyllus Trin., Ic. Spec. Gram.: Tb. 190 b (1828). – Type: No entry.. Notes: According to the depiction by Trinius, it is most likely a shortly branched Form of O. compositus var. rariflorus.
Oplismenus gracilis Schlecht., Linnaea 24: 649 (1851). - Panicum ischnocaulon Steud., Syn. Pl. Glum. I: 45 (1854). - Type: cult. "ab hortulano Halensi cl. Wolfhagen accepimus". Notes: Possibly a species of Oplismenus sect. Scabriseta. It distinguishes itself form all other species with its 1-nerved glumes. The awn of the Lower glume should only be 3.5 mm long, and the awn on the Upper glume, in contrast, 9 mm long. Eventually there will be a switch between the glumes.
Oplismenus setarius (Lam.) Roem. & Schult. f. sterilis F. Brown, Bernice P. Bishop Mus. Bull. 84: 68 (1931). – Type: Marquesas, Hivaoa, Feani: F. Brown 1034; BISH, Holotype, n. v. - Marquesas, Hivaoa, Atuona: F.. Brown 841, BISH, Paratype. - Marquesas, Nukuhiva, Taipi Vai: F. Brown 593, BISH, Paratype. Notes: Because these are sterile plants, it is impossible to distinctly classify them. F. Brown 841 probably does not belong in the genus Oplismenus due to it differing ligule. F. Brown 593 possibly belongs to O. hirtellus f. lanceolatus due to its vegetal characteristics and areas of distribution.
Orthopogon cubensis Spreng., Syst. Veg. I: 307 (1824, t. p. 1825). - Echinochloa cubensis (Spreng.) Schult., Mant. II: 596 (1824). - Oplismenus cubensis (Spreng.) Kunth, Rev. Gram. I: 45 (1829). - Panicum cubense (Spreng.) Steud., Nomencl. Bot. 2. ed. II: 255 (1841). - Type: Cuba n. v. Notes: = O. hirtellus (l.) P. Beauv., fide Hitchcock & Chase (1951), without Typical investigation.
Orthopogon trichoides Link, Hort. Berol. I: 203 (1827). - Type: cult. "habitat Indis orientalis?"; n. v. Notes: Most likely a culture variant of O. hirtellus subsp. undulatifolius.
Panicum brevisetum (Ness ex Steud.) Steud., Syn. Pl. Glum. I: 46 (1854). - Oplismenus brevisetus Nees ex Steud., Syn. PI. Glum. I: 46 (1854), pro syn. - Type: Guarea: Cuming s. n., in Herb. Lindley (CGE?). Notes: According to the original description, this is possibly O. affinis.
Panicum elatius L. f., Suppl.: 107 (1781). – Oplismenus elatior (L. f.) P. Beauv., Ess. Agrost.: 54 (1812). - Type: Malabaria; n. v. Notes: According to the original description, this is ver similar to O. compositus
Panicum incanum Schum. & Thonn., Beskr. Guin. Pl.: 60 (1827). - Type: Guinea: Thonning; n. v., C. Notes: It could be a studied sterile source specimen from the Thonning Herbarium, C, whose classification was not possible.
Sorghum saccharatum Host, Gram. Austr. IV: 48 (1809). Notes: = O. undulatifolius, fide Kew Index.
Oplismenus abortivus auct. ex Roem. & Schult., Syst. Veg. II: 808 (1817). - Vetiveria ziganoides (L.) Nash; fide Chase & Niles (1962).
Oplismenus abortivus (R. Br.) Desv., Opusc.: 82 (1931). = Vetiveria sp.; fide Stapf (1934).
Oplismenus angustifolius E. Fourn., Mex. Pl. 2: 40 (1886). = Echinochloa sp.
Oplismenus anomalus Mez ex Peter, Feddes Repert. Spec. Nov. Regni Veg ., Beih. 40,1 A: 200 (1938). - Type: "D. O. Afrika, Konde: Rungew", Stolz 1272, n. v. (B, destr.?) = Chloachne oplimenoides (Hack.) Stapf ex Robyns; fide Clayton (1972).
Oplismenus benthamii (Steud.) Coredem., Fl. Ile Réunion: 118 (1895). = Alloteropis paniculata (Benth.) Stapf; fide Clayton (1972), als "Panicum benthamii Steud.".
Oplismenus colonus (L.) H.B.K., Nov. Gen. et Spec.: 89 (1816). = Echinochloa colona (l.) Link; fide Gould et al. (1972).
Oplismenus crus-corvi (L.) Dumort., Obs. Belg.: 138, 151 (1823). = Echinochloa sp.; fide Chase & Niles (1962).
Oplismenus crus-galli (L.) Dumort., Obs. Belg.: 138 (1823). = Echinochloa crus-galli (L.) P. Beauv.; fide Gould et al. (1972).
Oplismenus crus-pavonis H.B.K., Nov. Gen. et Spec. I: 88 (1816). - Type: "crescit in aricis calidissimis Provinciae Cumanensis prope Bordones - Venezuela." = Echinochloa crus-pavonis (H.B.K.) Schult., fide Gould et a1. (1972).
Oplismenus cuspidatus Kunth, Rev. Gram. I: 45 (1829). = Echinochloa sp.; fide Chase & Niles (1962).
Oplismenus daltonii (Parl. ex Webb.) Schmidt, Beitr. Fl. Cap. Ver. Ins.: 136 (1852) = Echinochloa colona (L.) Link; fide Chase & Niles (1962) .
Oplismenus discolor (Trin.) Kunth, Rev. Gram. I: 45 (1829). = Panicum sp.; fide Chase & Niles (1962).
Oplismenus dubius Kunth, Rev. Gram. I: 44 (1829). = Echinochloa crus-galli (1.) P. Beauv.
Oplismenus echinatus Kunth, Rev. Gram. I: 45 (1829). = Echinochloa sp.
Oplismenus erianthos (Poir.) Kunth, Rev. Gram. I: 45 (1829). = Anthaenantia villosa (Michx.) P. Beauv.; fide Hitchcock & Chase (1917).
Oplismenus fasciculatus (Lam.) Roem. & Schult., Syst. Veg. II: 487 (1817). = Coridochloa cimcina (L.) Nees; fide Chase & Niles (1962).
Oplismenus festucaceus Mez, Notizbl. Bot. Gart. Berlin-Dahlem 7: 54 (1917). - Type: "Madagascar, in silva Amboh mitombo", Forsyth-Major 209; B. = Poecilostachys baronis Stapf.
Oplismenus festucoides (Nees) Kunth, Enum. Pl. I: 146 (1830). = Chaetium festucoides Nees; fide Chase & Niles (1962).
Oplismenus forsteri Kunth, Rev. Gram. I: 45 (1829). = Echinochloa sp.
Oplismenus frumentaceus Kunth, Rev. Gram. I: 45 (1829). = Echinochloa crus-galli (L.) P. Beauv. var. frumentacea (Roxb.) W. F. Wight; fide Gould et al. (1972).
Oplismenus helvolus (L.) P. Beauv., Ess. Agrost.: 54 (1812). = Chaetochloa sp.; fide Niles & Chase (1925).
Oplismenus hildebrandtii Mez, Feddes Repert. Spec. Nov. Regni Veg. 17: 83 (1921). - Type: Madagascar: Hildebrandt 3759, B. = Poecilostachys hildebrandtii Hack.
Oplismenus hirsutus (Roxb.) Koen. ex Schult., Mant. II: 271 (1824). = Panicum i Poir.; fide Hooker (1897) .
Oplismenus hirtus Heyne ex Roem. & Schult., Syst. Veg. II: 484 (1817). - Type: "In India orlentalis", B. Heyne s. n. = Echinochloa hirta Schult.
Oplismenus hispidulus (Retz.) Kunth, Rev. Gram. I: 44 (1829). - Type: Ind. or.: Koenig (BM?). = Echinochloa hispidula (Retz.) Keng; fide Chase & Niles (1962).
Oplismenus hispidus (Muhl.) Wood., Class-Book, ed. 2: 604 (1847). = Echinochloa walteri (Pursh) Heller; fide Gould et al. (1972).
Oplismenus holciformis H.B.K., Nov. Gen. et Spec. I: 88 (1816). - Type: "crescit in humidis montanis prope Cinapecuaro", Mexico: Humboldt & Bonpland 4362 (P?). = Echinochloa holciformis (H.B.K.) Chase; fide Chase & Niles (1962).
Oplismenus hookeri Parl., Ottova Riun. Sc. Ital.: 586 (1846). = Echinochloa sp.
Oplismenus intermedius (Vahl ex Hornem.) Kunth, Rev. Gram. I: 45 (1829). = Echinochloa sp.
Oplismenus jamaicensis Kunth, Enum. Pl. I: 147 (1833). = Echinochloa crus-pavonis (H.B.K.) Schult.; fide Gould et al. (1972).
Oplismenus javanicus Roem. & Schult., Syst. Veg. II: 891 (1817). = Panicum javanicum Poir.
Oplismenus lanceolatus (Retz.) Kunth, Rev. Gram. I: 45 (1829). - Type: "Ind. or." Koenig (BM?). = Echinochloa lanceolata (Retz.) P. Beauv.
Oplismenus lappagoides Speg., Anal. Mus. Nac. Buenos Aires 3,2: 7 (1903). = Pseudechinolaena polystachya (H.B.K.) Stapf; fide Cabrera (1970).
Oplismenus limosus K. B. Presl, Rel. Haenk. I: 321 (1830). - Typus: "In insula Luzon", Haenke s. n., PR. = Echinochloa crus-galli (L.) P. Beauv.
Oplismenus longisetus (Torr.) Kunth, Rev. Gram. I: 45 (1829). = Echinochloa walteri (Pursh) Heller; fide Gould et al. (1972).
Oplismenus magellanicus Roem. & Schult., Syst. Veg. II: 485 (1817). = Centosteca latifolia (Osb.) Trin.; fide, Manod de Froideville (1971).
Oplismenus margaritaceus (Link) Kunth, Rev. Gram. I: 44 (1829). = Echinochloa sp.; fide Chase & Niles (1962).
Oplismenus minarum Nees, Agrost. Bras.: 268 (1829). - Type: "Brasilia, Paranàn, Villa Ricca, prov. Rio Negro"; M.= Ichnanthus minarum (Nees) Doell.
Oplismenus muricatus (Michx.) Kunth, Rev. Gram. I: 44 (1829). = Echinochloa muricata (P. Beauv.) Fern.; fide Gould et al. (1972).
Oplismenus muticus R. A. Philippi, Anal. Univ. Chil. 93: 714 (1896). - Type:, "In praedia Mansel (prov. O'Higgins) orn. Nathanis. Miers Cox legi". = Echinochloa colona (L.) Link; fide Cabrera (1970).
Oplismenus nossibensis Mez, Notizbl. Bot. Gart. Berlin-Dahlem 7: 53 (1917). - Type: Madagascar, Nossibé: Hildebrandt 3354, B. = Cyphochlaena madagascariensis Hack.
Oplismenus oplismenoides Speg., Anal. Mus. Buenos Aires 9: 7 (1903). = Pseudechinolaena polystachya (H.B.K.) Stapf; fide Cabrera (1970).
Oplismenus penicillatus (Nees) Kunth, Rev. Gram. I: 45 (1829).= Panicum penicillatum Nees; fide Chase & Niles (1962).
Oplismenus pictus Kunth, Enum Pl. I: 144 (1830). = Echinochloa crus-galli (L.) P. Beauv.
Oplismenus polystachyus H.B.K., Nov. Gen. et Spec. I: 88 (1816). = Echinochloa polystachya (H.B.K.) Rojas; fide Chase & Niles (1962).
Oplismenus prostratus Edgew., J. Linn. Soc. Bot. 6: 195 (1862). = Panicum setigerum Retz.; fide Hooker (1897) .
Oplismenus pseudocolonus (Roth) Kunth, Rev. Gram. I: 44 (1829). = Echinochloa colona (L.) Link; fide Kew Index.
Oplismenus pseudoundulatlfolius (Roem. & Schult.) Kunth, Rev. Gram. I: 44 (1829). = Panicum pseudoundulatiofolium Roem. & Schult.; fide Kew Index.
Oplismenus repens K. B. Presl, Rel. Haenk: I: 321 (1830). - Type: Mexico: Haenke s. n. PR. = Echinochloa colona (L.) Link.
Oplismenus sabulicolus (Nees) Kunth, Enum Pl. I: 145 (1833). - Type: "In arenosis Parae. Sieber". = Echinochloa crus-pavonis (H.B.K.) Schult.; fide Gould et al. (1972).
Oplismenus scaber (Lam.) Kunth, Rev. Gram. I: 44 (1829). = Echinochloa crus-galli (L.) Link; fide Hooker (1897) .
Oplismenus secundus K. B. Presl, Rel. Haenk. I: 322 (1830). - Type: "In Peruviae montanis huanoccensibus", Haenke, PR, n. v. = Ichnanthus minarum (Nees) Doell.; fide Chase & Niles (1962).
Oplismenus semialatus Desv., Opusc.: 81 (1831). = Axonopus semialatus Hook. f.; fide Kew Index.
Oplismenus spectabilis (Nees) Kunth, Enum Pl. I: 145 (1833) = Echinochloa spectabilis (Nees) Link.
Oplismenus stagninus (Retz.) Kunth, Rev. Gram. I: 44 (1829). = Echinochloa stagnina (Retz.) P. Beauv.; fide Chase & Niles (1952).
Oplismenus strictus Schult., Mant. II: 272 (1824). = Arundinella Wallichii Nees ex Steud.
Oplismenus tenuis K. B. Presl, Rel. Haenk. I: 319 (1830). – Type: Mexico, Haenke, PR. = Ichnanthus tenuis (K. B. Presl) Hitchcock; fide Chase & Niles (1962).
Oplismenus tomentosus Schult., Mant. II: 272 (1824). Setaria verticillata·P. Beauv., fide Kew Index.
Oplismenus volkensii (Pilger) Mez ex Peter, Feddes Repert. Spec. Nov. Regni Veg., Beih. 40,1 A: 222 (1938). - Type: East Afrika, Kilimanjaro: Volkens 1278, B. = Achritochaete volkensii Pilger.
Oplismenus walteri (Muhl.) Kunth, Rev. Gram. I: 45 (1829). = Panicum hemitomon Schult.; fide Chase & Niles (1962).
Oplismenus zelayensis H.B.K., Nov. Gen. et Spec. I: 89 (1816). = Echinochloa crus-galli (L.) P. Beauv. var. zelayensis (H.B.K.) Hitchcock; fide Abrams (1953).
Orthopogon abortivus (R. Br.) Spreng., Syst. Veg. I: 306 (1824 t. p. 1825). = Chamaeraphis spinescens Poir.; fide Chase & Niles (1962).
Orthopogon agrostoides Trev. ex Steud., Nomencl. 2. ed. II: 234, pro syn. = Arundinella brasiliensis Raddi; fide Kew Index.
Orthopogon crus-galli (L.) Spreng., Syst. Veg. I: 307 (1824 t. p. 1825). = Echinochloa crus-galli (L.) P. Beauv.; fide Gould et al. (1972).
Orthopogon dichotomus Llanos, Fragm.: 38 (1851) = Echinochloa colona (L.) Link; fide Chase & Niles (1962).
Orthopogon echinatus Spreng., Syst. Veg. I: 307 (1824 t. p. 1825). = Echinochloa sp.
Orthopogon hirsutus Spreng. ex Steud., Nomencl. 2. ed. II: 234 (1841), pro syn. = Echinochloa spectabilis (Nees) Link; fide Chase & Niles (1962).
Orthopogon hispidus (Muhl.) Spreng., Syst. Veg. I: 307 (1824 t. p. 1825). = Echinochloa walteri (Pursh) Heller; fide Chase & Niles (1962).
Orthopogon halciformis (H.B.K.) Spreng., Syst. Veg. I: 307 (1824 t. p. 1825). = Echinochloa holciformis (H.B.K.) Chase; fide Chase & Niles (1962).
Orthopogon retzii Spreng. Syst. Veg. I: 307 (1824 t. p. 1825). = Echinochloa hispidula (Retz.) Keng; fide Chase & Niles (1962).
Orthopogon sqarrosus Spreng., Syst. Veg; I: 307 (1824 t. p. 1825). = Chamaeraphis spinescens Poir., fide Chase & Niles (1962).
Orthopogon stagninus (Retz.) Spreng., Syst. Veg. I: 307 (1824 t. p. 1825). = Echinochloa stagnina (Reti.) P. Beauv.; fide Chase & Niles (1962).
Orthopogon subverticillatus Llanos, Fragm.: 38 (1831). Echinochloa sp.; fide Chase & Niles (1962).
Here I would like to like to thank Prof. Dr. H. Scholz for his multi-faceted help with both the preparation of the manuscript for publishing and the improvement of my of scientific work methods. A special thank-you goes out to Ms. Dieckmann, a scientific illustrator with the Botanical Museum of Berlin-Dahlem, for the production of the habit and detail illustrations, an especial part of this work. I would also like to thank Dr. B. Schick for the guidance and support with the chemical analysis of the awns. My earnest thanks goes out to all directors of the above mentioned herbariums for their loaning out of needed plant specimens.
In this monograph, the genus Oplismenus was discussed morphologically, anatomically, systematically, and phytogeographically using historical sources and my own studies. .
The genus was newly divided into two sections with 9 species and 18 infraspecific taxa.
Numerous changes of status are presented 3 additional taxa described: O. hirtellus subsp. fasciculatus, O. hirtellus subsp. imbecillis f. lanceolatus, and O. aemulus ver. densiflorus.
Epidermal investigation and chemical analysis of the sticky secretion of the awns (Oplismenus sect. Oplismenus) completed the morphological-systematic conclusions. These secretions could possibly belong to the family of Triterpenes.
The disjunction distribution of O. hirtellus subsp. undulatifolius is discussed; the possbility that its European distribution is the result of its being Tertiary Relict are discussed, but no conclusion reached.
From M. E. Barkworth: Literature cited. This portion was made a separate file. The index follows but I have not attempted to link the names to the places where they are mentioned; it would take too long. A search through the file will reveal the pages where the names are mentioned. Alternatively, I recommend downloading the pdf file of the original translation. This file has been edited from that translation. Of course, anyone questioning either this file or the translation should work from the original German - and German-speaking botanist. The translators that helped create this version of Scholz' monograph were language students with no background in botany. Moreover, I asked for a rather fast translation so there was no time to discuss the result. It has helped me understand Dr. Scholz's conclusions, which is why I commissioned it; I hope that it helps others. Please let me know of errors. Mary Barkworth, August 14, 2009
Andropogon undatus Jacq.
Echinochloa cubensis Schult.
Echinochloa hirtella Schult.
Hekaterosachne elatior Steud.
Hippagrostis burmannii O. Kuntze
Hippagrostis composita O. Kuntze
Hippagrostis hirtella O. Kuntze
Hippagrostis loliacea O. Kuntze
Hippagrostis setaria O. Kuntze
Hippagrostis undulatifolia O. Kuntze
Milium undulatifolium Moench.
Oplismenus abortivus auct. ex. Roem. & Schult.
Oplismenus abortivus Desv.
Oplismenus acuminatus Nees ex Steud.
Oplismenus aemulus Roem. & Schult. ‘aemulans'
Oplismenus aemulus var. aemulus
Oplismenus aemulus var. densiflorus U. Scholz
Oplismenus aemulus var. flaccidus Domin
Oplismenus aemulus var. lasiorhachis Domin
Oplismenus aemulus var. pilosus Domin
Oplismenus affinis K. B. Presl
Oplismenus affinis Schult.
Oplismenus affinis var. affinis
Oplismenus affinis var. humboldtianus U. Scholz
Oplismenus africanus P. Beauv.
Oplismenus africanus var. capensis Stapf
Oplismenus africanus var. simplex Stapf
Oplismenus albus Roem. & Schult.
Oplismenus angustifolius E. Fourn.
Oplismenus anomalus Mez
Oplismenus aristulatus Burcham
Oplismenus bakeri Schinz
Oplismenus baronii Camus
Oplismenus benthamii Cordem.
Oplismenus brasiliensis Raddi
Oplismenus brevisetus Nees ex Steud.
Oplismenus bromoides Baker
Oplismenus bromoides P. Beauv.
Oplismenus burmannii Mez
Oplismenus burmannii P. Beauv.
Oplismenus burmannii var. burmannii
Oplismenus burmannii var. intermedius Honda
Oplismenus burmannii var. lanatus Baker
Oplismenus burmannii var. multisetus U. Scholz
Oplismenus burmannii f. cristatus Hier. in Peter
Oplismenus burmannii Thwaites
Oplismenus capensis Hochst.
Oplismenus chondrosioides E. Fourn.
Oplismenus colonus H. B. K.
Oplismenus compositus Bauer
Oplismenus compositus P. Beauv.
Oplismenus compositus var. brachyphyllus Trin.
Oplismenus compositus var. compositus
Oplismenus compositus var. imbecillis Bailey
Oplismenus compositus var. intermedius Ohwi
Oplismenus compositus var. lasiorhachis Hack.
Oplismenus compositus var. loliaceus Hack.
Oplismenus compositus var. owatarii Ohwii
Oplismenus compositus var. patens Ohwii
Oplismenus compositus var. rariflorus U. Scho1z
Oplismenus compositus var. setarius Bailey
Oplismenus compositus var. sylvaticus U. Scholz
Oplismenus compositus f. glabratus F. Brown
Oplismenus compositus f. pubescens F. Brown
Oplismenus coreanus Nakai
Oplismenus cristatus K. B. Presl
Oplismenus crus-corvi Dumort.
Oplismenus crus-galli Dumort.
Oplismenus crus-pavonis H. B. K.
Oplismenus cubensis Kunth
Oplismenus cuspidatus Kunth
Oplismenus daltonii Schmidt
Oplismenus decompositus Nees
Oplismenus depauperatus E. Fourn.
Oplismenus discolor Kunth
Oplismenus dubius Kunth
Oplismenus echinatus Kunth
Oplismenus elatior P. Beauv.
Oplismenus erianthos Kunth
Oplismenus fasciculatus Roem. & Schult.
Oplismenus festucaceus Mez
Oplismenus festucoides Kunth
Oplismenus flaccidus Roem. & Schult.
Oplismenus flavicomus Mez
Oplismenus foliaceus P. Beauv.
Oplismenus formosanus Honda
Oplismenus forsteri Kunth
Oplismenus frumentaceus Kunth
Oplismenus gracilis Schlecht.
Oplismenus gracillimus Mez
Oplismenus helvolus P. Beauv.
Oplismenus hildebrandtii Mez
Oplismenus hirsutus Keon. ex Schult.
Oplismenus hirtellus P. Beauv.
Oplismenus hirtellus subsp. acuminatus U. Scholz
Oplismenus hirtellus subsp. capensis Mez ex U. Scholz
Oplismenus hirtellus subsp. fasciculatus U. Scholz
Oplismenus hirtellus subsp. hirtellus
Oplismenus hirtellus subsp. imbecillis U. Scholz
Oplismenus hirtellus subp. japonicus U. Scholz
Oplismenus hirtellus subsp. loliaceus Mez
Oplismenus hirtellus subsp. microphyllus U. Scholz
Oplismenus hirtellus subsp. psilostachys U. Scholz
Oplismenus hirtellus subsp. setarius Mez ex Ekman
Oplismenus hirtellus subsp. tsushimensis U. Scholz
Oplismenus hirtellus subsp. undulatifolius U. Scholz
Oplismenus hirtellus var. acuminatus Mez
Oplismenus hirtellus var. acuminatus f. foliaceus Stehle
Oplismenus hirtellus var. acuminatus f. imbecilis
Oplismenus hirtellus var. acuminatus f. lanceolatus U. Scholz
Oplismenus hirtellus Roem. & Schult.
Oplismenus hirtiflorus K. B. Presl
Oplismenus hirtus Heyne ex Roem. & Schult.
Oplismenus hispidulus Kunth
Oplismenus hispidus Wood.
Oplismenus holciformis H. B. K.
Oplismenus hookeri Parl.
Oplismenus humbertianus Camus
Oplismenus humboldtianus Nees
Oplismenus humboldtianus var. genuinus E. Fourn.
Oplismenus humboldtianus var. muticus E. Fourn.
Oplismenus humboldtianus var. nudicaulis Vasey
Oplismenus humboldtianus imbecilis Roem. & Schult.
Oplismenus humboldtianus var. morrisonensis Honda
Oplismenus indicus Roem. & Schult.
Oplismenus intermedius Kunth
Oplismenus jacquinii Kunth
Oplismenus jamaicensis Kunth
Oplismenus japonicus Honda
Oplismenus javanicus Klotzsch ex Schlecht.
Oplismenus javanicus Roem. & Schult.
Oplismenus junghuhnii Boerlage
Oplismenus lanceolatus Kunth
Oplismenus lappagoides Speg.
Oplismenus latifolius Haenke ex Steud.
Oplismenus liebmannii E. Fourn.
Oplismenus limosus K. B. Presl
Oplismenus loliaceus P. Beauv.
Oplismenus longisetus Kunth
Oplismenus magellanicus Roem. & Schult.
Oplismenus margaritaceus Kunth
Oplismenus microphyllus Honda
Oplismenus minarum Nees
Oplismenus minor Merrill
Oplismenus mollissimus Hochst. ex Steud.
Oplismenus multisetus Hochst. ex A. Rich.
Oplismenus muricatus Kunth
Oplismenus muticus Philippi
Oplismenus nossibensis Mez
Oplismenus oahuaensis Nees & Mez ex Steud.
Oplismenus oplismenoides Speg.
Oplismenus owatarii Honda
Oplismenus parvifolius Kunth
Oplismenus patens Honda
Oplismenus penicillatus Kunth
Oplismenus pictus Kunth
Oplismenus polliniifolius Honda
Oplismenus polystachyus H. B. K.
Oplismenus pratensis Schult.
Oplismenus preslii Kunth
Oplismenus prostratus Edgew.
Oplismenus pseudocolonus Kunth
Oplismenus pseudoundulatifolius Kunth
Oplismenus psilostachys Honda
Oplismenus rariflorus K. B. Presl
Oplismenus repens K. B. Presl
Oplismenus sabulicolus Kunth
Oplismenus scaber Kunth
Oplismenus secundus K. B. Presl
Oplismenus semialatus Desv.
Oplismenus setarius Roem. & Schult.
Oplismenus var. aemulus Bailey
Oplismenus setarius var. imbecillis Benth.
Oplismenus setarius var. imbecillis f. sterilis F. Brown
Oplismenus simplex K. Schum. ex Engler
Oplismenus spectabilis Kunth
Oplismenus stagninus Kunth
Oplismenus strictus Schult.
Oplismenus sylvaticus Roem. & Schult.
Oplismenus tenuis K. B. Presl
Oplismenus thiebautii E. Fourn.
Oplismenus thwaitesii Hook. f.
Oplismenus tomentosus Schult.
Oplismenus tsushimensis Honda
Oplismenus undulatifolius P. Beauv.
Oplismenus undulatifolius Roem. & Schult.
Oplismenus undulatifolius var. elongatus Honda ex Nakai
Oplismenus undulatifolius var. imbecillis Hack
Oplismenus undulatifolius var. japonicus Koidzumi
Oplismenus undulatifolius var. lanceolatus Domin
Oplismenus undulatifolius var. microphyllus Ohwi
Oplismenus undulatifolius var. mollis Domin
Oplismenus velutinus Schult.
Oplismenus volkensii Mez
Oplismenus walteri Kunth
Oplismenus zelayensis H. B. K.
Orthopogon
Orthopogon abortivus Spreng.
Orthopogon aemulus R. Br.
Orthopogon africanus Sweet
Orthopogon agrostoides Trev. ex Steud.
Orthopogon albus Nees ex Steud.
Orthopogon bolosii Vayreda
Orthopogon bromoides Loud.
Orthopogon burmannii R. Br.
Orthopogon burmannii var. glabrescens Buse
Orthopogon burmannii var. lanatus Buse
Orthopogon compositus R. Br.
Orthopogon compositus var. glabrescens Buse
Orthopogon crus-galli Spreng.
Orthopogon cubensis Spreng.
Orthopogon dichotomus Llanos
Orthopogon echinatus Spreng.
Orthopogon flaccidus R. Br.
Orthopogon gonyrrhizus Miq.
Orthopogon hirsutus Spreng. ex Steud.
Orthopogon hirtellus R. Br.
Orthopogon hirtellus Nutt.
Orthopogon hispidus Spreng.
Orthopogon holciformis Spreng.
Orthopogon imbecillis R. Br.
Orthopogon junghuhnii Nees ex Steud.
Orthopogon loliaceus Spreng.
Orthopogon longeracemosus Miq.
Orthopogon parvifolius Nutt.
Orthopogon pratensis Spreng.
Orthopogon remotus Trin.
Orthopogon retzii Spreng.
Orthopogon setarius Spreng.
Orthopogon sqarrosus Spreng.
Orthopogon stagninus Spreng.
Orthopogon subverticillatus Llanos
Orthopogon sylvaticus Miq.
Orthopogon trichoides Link
Orthopogon undulatifolius Spreng.
Orthopogon undulatus Link
Orthopogon velutinus Spreng.
Panicum
Panicum acuminatissimum Steud.
Panicum aemulum Steud.
Panicum africanum Poir.
Panicum album Poir.
Panicum aristatum Retz.
Panicum balfourii Baker
Panicum barbifultum Hochst. ex Schlecht.
Panicum bidentatum Steud.
Panicum bidentulum Steud.
Panicum brevisetum Steud.
Panicum bromoides Lam.
Panicum burmannii Balb.
Panicum burmannii Marschall Bieb.
Panicum burmannii Retz.
Panicum certificandum Steud.
Panicum composito-proximum Rottl. ex Willd.
Panicum compositum L.
Panicum compositum Rottl. ex Steud.
Panicum cristatum Steud.
Panicum cubense Steud.
Panicum elatius L. f.
Panicum flaccidum Steud.
Panicum foliaceum Steud.
Panicum francoi Steud.
Panicum gonyrrhizum Steud.
Panicum hirtellum All.
Panicum hirtellum Burm. f.
Panicum hirtellurm Host
Panicum hirtellum Lam.
Panicum hirtellum L.
Panicum hirtellum Muhl.
Panicum hirtellum Scop.
Panicum hirtellum Wulfen ex Jacq.
Panicum imbecille Trin.
Panicum incanum Schum. & Thonn.
Panicum ischnocauloh Steud.
Panicum japonicum Steud.
Panicum kraussii Steud.
Panicum lappaceum Willd. ex Spreng.
Panicum loliaceum Lam.
Panicum longeracemosum Steud.
Panicum multisetum Hochst. ex A. Rich.
Panicum nuttallianum Steud.
Panicum oahuaense Steud.
Panicum parciflorum Steud.
Panicum peninsularum Steud.
Panicum pratense Steud.
Panicum raddianum Steud.
Panicum sanctae-marthae Steud.
Panicum schultesii Steud.
Panicum setarium Lam.
Panicum sylvaticum Lam.
Panicum undatum Steud.
Panicum undulatifolium Ard.
Panicum velutinump. F. N. Meyer
Pollinia undata Spreng.
Setaria hirtella Schult.
Sorghum saccharatum Host
