Annals of the Missouri Botanical Garden Volume XLVII 1960 Annals of the Missouri Botanical Garden A Quarterly Journal containing Scientific Contributions from the Missouri Botanical Garden and the Henry Shaw School of Botany of Washington University in affiliation with the Missouri Garden. Information The Annals of the Missouri Botanical Garden appears four times during the calendar year: February, May, September, and November. Four Beginning with Volume 45, 1958: Subscription Price $12.00 per volume Single Numbers 3.00 each Content* of previous issues of the Annals of the Missouri Botanical Garden are listed in the Agricultural Index, published by the H. W. Wilson TABLE OF CONTENTS Calamophyton bicephalum, A New Species from the Middle Devonian of Belgium Suzanne Leclercq and Henry N. Andrews, Jr. and Phytogeny in the Orchidaceae Robert L. Dressier and Calaway H. Dodson 25-68 The Genus Schismatoglottis (section Philonotion) in America George S. Bunting 69-7 1 Taxonomica. II 73-8 o Panama. Part IV, Fascicle 2 (Chloranthaceae to Proteaceae) -Robert E. Woodson, Jr., Robert W. Schery and Collaborators 81-203 The Strengthening System in the Stem of Maize William H. Murdy 205-226 Studies Involving Sustained Treatment of Maize with Gibberellic Acid I: Further Notes on Responses of Races Norton H. Nickerson and Thomas N. Embler 227-242 Studies Involving Sustained Treatment of Maize with Gibberellic Acid II: Responses of Plants Carrying Certain Tassel-Modifying Genes Norton H. Nickerson 243-261 Flora of Panama. Part IV, Fascicle 3 (Loranthaceae to Polygonaceae) .Robert E. Woodson, Jr., Robert W. Schery and Collaborators 263-359 General Index to Volume XL VII 361-373 MISSOURI BOTANICAL GARDEN STAFF Frits W. Went Waldo G. Fechner, Oscar E. Glaessner, C. Ranlet Lincoln, Norton H. Nickerson, Carroll W. Dodge, Trifon von Schrenk, Mycologist Assistant Curator Museum of ROBERT L. DrESSLER, Economic Plants ** 1 S«tertC«*tar George B. Van Schaack, John D. Dwyer, Robert E. Woodson, Jr., BOARD OF TRUSTEES Robert Brookings Smith Vice-President Leicester B. Faust Second Vice-President Henry B. Pflager Howard F. Baer Dandzl K. Catlin Sam'l. C. Davis Dudley French Henry Hitchcock John S. Lehmann Robert W. Otto Warren McK. Shapleigh EX-OFFICIO MEMBERS George L. SeSoi Stratf. 3rd Lee Morton, Edward C. Donnelly Ethan Raymond R. Tucker A. H. Shepley, Oscar E. Glaessner, Secretary Volume XL VII Annals of the Missouri Botanical Garden FEBRUARY, 1960 Classification and Phylogeny in the Orchidaceae ..... Robert L ay H. Dodson 25-68 The Genus Schismatoglottis (section P ica ;,-:■ Miscellanea Taxonomica. II. .... 73-80 Annals of the Missouri Botanical Garden A Quarterly Journal containing Scientific Contributions from the Missouri Botanical Garden and the Henry Shaw School of Botany of Washington University in affiliation with the Missouri Botanical Garden. Garden are listed i Annals of the Missouri Botanical Garden FEBRUARY, 1960 CALAMOPHYTON BICEPHALUM, A NEW SPECIES FROM THE MIDDLE DEVONIAN OF BELGIUM SUZANNE LECLERCQ and HENRY N. ANDREWS, Jr. Introduction The fossil plant material on which the present study is based was collected by S. Leclercq in 1949 and 1952 from a Middle Devonian horizon in eastern Belgium. The Middle Devonian in Belgium is divided into the Couvinian and the overlying Givetian and our specimens come from the lower part of the latter. The fossiliferous horizon is well exposed in one of two quarries (the north one) known locally as the "Carrieres Brandt". They are situated on the upper part of the western slope of a hill lying 400 meters south of the junction of the rivers "La Gileppe" and "La Vesdre" at the small village of Goe which is in turn located about 30 kilometers east of Liege and 20 kilometers from Germany. The plant- bearing horizon is a lens-shaped deposit of numerous greenish and grey-blue shale layers 0.1 to 1.1 meters thick which are interbedded with arkose strata. It has been shown that the Middle Devonian in this region is a littoral formation deposited in shallow marine conditions (Aderca, 1932, p. 14; Liegeois, 1956, p. 98). The fossil remains are abundantly distributed through the shales but the quality of preservation varies with the different layers. In most of them large specimens are encountered while in some the material is more fragmentary. The plants are found for the most part as well preserved compressions and occasionally as petri- factions in which the plant tissues are impregnated with iron hydroxide. The "Goe flora" is a large one and includes many specimens referable to the Protoarticulatae. Some of them are large, splendid specimens representing the (1) 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN typical habit of Calamophyton; others are more fragmentary and it is prob- lematical as to whether they should be referred to this genus or to Hyenia. It is, moreover, very possible that some will prove to lie outside the limits of both genera. As is often the case in paleobotany our knowledge of these presumed early articulates has accumulated as the result of the work of several investigators and the quality of preservation and relative fragmentary state of the plants dealt with has varied greatly. Due chiefly to faulty preservation previous interpretations are not entirely correct and it seems fair to assert that, in general, the plants are not as well known as some of the published accounts would lead one to believe. The present investigation was initiated in part from the discovery of the elaborate organization of the sporangiophore of the Upper Devonian sphenopsid cone Eviostachya hoegi Stockmans which was described in detail by Leclercq in 1957. The unexpected complexity of the sporangiophore "head" of this plant and the availability of well preserved specimens of the Middle Devonian Proto- articulates suggested the desirability of reinvestigating the latter. Our study is based on several Calamophyton specimens which display the typ- ical habit of that plant and the preservation is such that we have had an oppor- tunity to contribute somewhat to a better understanding of the morphology of the leaves and spore-bearing appendages. The latter have proven to be much more complex than was supposed and we have been able to add appreciably to our knowledge of the branching pattern of the leaves. It is thus necessary to review the previous studies, at least briefly, particularly with reference to the presumed differences between Calamophyton and Hyenia. Previous Studies The genus Calamophyton was established in 1926 by Krausel and Weyland with C. primaevum as the type species. It is reconstructed as a plant with an upright main stem which branches initially in a more or less digitate fashion; the primary branches, although predominantly monopodial, may also give rise to nearly equal dichotomies (see their plate 15, fig. 2). The leaves are once or twice forked and apparently less filiform than those of Hyenia. In view of our own description that follows it is perhaps significant to emphasize this difference as noted by Krausel and Weyland (cf. their text figs. 26 of Calamophyton and 16a, 16b of Hyenia). Somewhat stouter, unforked sterile appendages were borne on the "main stem" and on the basal part of the primary branches. The fertile shoots are similar to those of Hyenia but each arm of the bifurcated sporangiophore usually bears but one sporangium (cf. their text fig. 27 of Calamophyton and text fig. 22 of Hyenia). Additional specimens were described by the same authors in their 1929 contribution. In 1932 Aderca described and figured a fine sterile specimen of Calamophyton primaevum which was obtained at the same outcrop from which the new material described herewith was obtained. This specimen will be referred to again but it may be pointed out that it was erroneously determined as Hyenia elegans (see LECLERCQ & ANDREWS CALAMOPHYTON BICEPHALUM 3 Aderca's pi. 2, fig. 4), although the mistake was corrected by Krausel (1932, p. 277). In 1940 Leclercq reported C. primaevum, as well as a new species, C. renieri, from the Middle Devonian of Belgium. She noted (see pp. 18, 33, 35) that most of the leaves in both species were more frequently divided and more filiform than those described for the above mentioned German specimens (cf. Leclercq, 1940, text fig. 9 with Krausel and Weyland's figures). The differences between the foliage of Calamophyton primaevum and Hyenia elegans as indicated by Krausel and Weyland in the text figures cited above is quite striking; those of Calamophyton show a leaf that is narrow, almost wedge-shaped with a notched tip and it is this figure that has been used in textbooks (see, for example, Hirmer, 1927, fig. 400, 401; Arnold, 1947, fig. 55; Walton, 1953, fig. 38; Smith, 1955, p. 241; Magdefrau, 1956, fig. 61). Actually, Krausel and Weyland's restoration does not bear out this exaggerated difference and in their description the leaves of Calamophyton are described as "schmallineal, ein- bis mehrfach gegabelt" (1926, p. 141). Unfortunately their specimens are no longer available for comparison but we shall present additional evidence which suggests that it is very doubtful whether the two genera can be distinguished on the basis of their leaves when only fragmentary specimens are available. The fertile appendages (sporangiophores) of the two genera were described by Krausel and Weyland as being very similar. In the case of Hyenia each branch of the sporangiophore is said to bear two or three sporangia while only one is found in the corresponding position in Calamophyton (cf. their text fig. 22 for Hyenia and text fig. 27 for Calamophyton) . In her 1940 study of certain Belgian protoarticulates Leclercq noted that in Calamophyton each branch of the sporangi- ophore usually bore two sporangia; in the Hyenia specimens that she then had available the sporangiophores were noted to have several slender, bifurcating distal appendages. This suggested a greater complexity than was reported for the German fossils and is interesting in the light of our present observations which have been made on specimens of vastly better preservation. As to the general habit of the plants in the two genera, although Krausel and Weyland's original restoration of Hyenia (1926, pi. 16) shows a plant rather similar to that of Calamophyton, it was later recognized that the leafy and fertile shoots of Hyenia were borne on a relatively massive rhizome (Krausel and Weyland, 1932, fig. 3). Studies that confirm this type of habit have been made on Hyenia specimens from Norway (Hoeg, 1945) and Belgium (Leclercq, 1940) and these authors have both supplied restorations that indicate a conspicuous rhizome with slender upright shoots which are usually unbranched. In summary it may be said, in the light of our present knowledge, that it is very doubtful whether the upright shoots of Hyenia can be differentiated from the distal parts of Calamophyton and only the general habit of the two remains as a distinguishing character that may be used with assurance. There is thus some question as to whether the two should be generically separated. It may also be noted that the anatomy of the two is not sufficiently well known to shed much light on the problem. 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN Materials on Which the Present Study Is Based; Study Technique It is evident from the above summary that much remains to be learned about these fossil plants and it is particularly necessary that we have better preserved specimens in order to eliminate the doubt that exists concerning the morphology and taxonomy of these presumed early members of the articulate group. Our investigations have not solved all the problems but we have been able to make the following contributions which add to an understanding of the morphology of Calamophyton: 1. The sterile appendages (leaves) dichotomize two to four times and it is doubtful whether they can, with certainty, be distinguished from those of Hyenia. 2. The sterile appendages are three-dimensional in their branching pattern. 3. The fertile appendages are much more complex than previous accounts 4. Both fertile and sterile appendages may be borne on the same branch; there is, however, a marked tendency for the fertile appendages to be aggregated together into an apparently specialized branch system, but not into a clearly defined "cone". The specimens that have been studied and the names assigned to them are as Calamophyton bicephalum Leclercq and Andrews, sp. nov.: Nos. 5011/609, 5012/337, 5009/588, 5007/346. In all cases the first of each dual number is the permanent one assigned to the fossil and the second is the number originally given in the field). Calamophyton primaevum Krausel and Weyland: No. 5006, previously de- scribed by Aderca, 1932, pi. 11, fig. 4. Since the technique that is employed in the investigation of these fossils, although very simple, is perhaps not widely known, a brief description seems appropriate. The plants are preserved as compressions and both fertile and sterile appendages were three dimensional in their branching patterns. Since they also possessed some rigidity a single appendage may extend through several millimeters of sediment, thus it is not possible to apply a transfer technique and remove the entire specimen. What is revealed when the rock is first split gives at best a partial picture of an appendage as we shall point out in some detail in the descrip- tions. It is thus necessary to follow, through both part and counterpart, a selected appendage in order to determine its three-dimensional form. In the case of the fertile appendages, due to their small size and complex branching pattern, it often becomes necessary to excavate the matrix in terms of fragments only a few microns in diameter. Mechanical vibratory devices are much too coarse for such a delicate degagement and it is accomplished with steel needles and a small, light weight hammer. The size of the needles and the shape of the point will depend on the nature and delicacy of the operation. Invaluable information may thus be obtained with careful, painstaking work and the only costly element involved is Gross Morphology of the Branch System Reference will be made first to figure 1 (No. 5012/337) which is the largest LECLERCQ & ANDREWS CALAMOPHYTON BICEPHALUM 5 and most complete specimen insofar as the branch system as a whole is concerned. This consists of a major axis two cm. in diameter which divides repeatedly; since it is shown natural size in the figure it does not seem necessary to cite measure- ments for the various branch orders. It should be noted that the ultimate basal part of the plant is not preserved nor has this been identified in any other speci- mens. We shall, however, as a matter of convenience in description, refer to this basal part of the specimen as the main stem, and the first division members as the primary branches. Although the initial branching of the main stem in a more or less digitate fashion seems to be highly characteristic of Calamophyton there is some variation in the organization of this cluster of primary branches. In figure 1 the main stem divides to form two unequal branches and the latter continue to dichotomize more or less equally throughout the length of the specimen. In the specimen shown at the upper left of figure 7 the main stem produces, almost simultaneously, seven primary branches of nearly equal diameter; the main stem of the specimen in the lower right of this figure starts with an equal dichotomy and the primary branches quickly divide again. In figure 14 the main stem produces three primary branches, although it might be interpreted as an initial unequal dichotomy in which the larger branch to the right dichotomizes again about two mm. above the initial division. The rather broad diagonal white band running through the upper part of the main stem represents a slight fault in the rock. It is important to note that the main stem in fig. 1, and to a lesser degree the basal portion of the primary branches, shows the transverse bands which have been cited as a characteristic feature of Calamophyton. Several of these striations are preservation artifacts although some suggest an internal structure possibly com- parable with the higher articulates; they are not evident in the upper branches of the specimen. Less clearly defined bands are present in the larger specimen (upper left) of fig. 7; we have not observed the transverse bands in the other specimens at our disposal. It also seems significant that in the basal part of this specimen (fig. 1) the striations are not correlated with any regularity in the distribution of the sterile appendages; thus if it is assumed that the transverse striations indicate nodes the leaves are scattered irregularly through the internodal region. Therefore we do not feel that there is sufficient evidence to regard this character as distinctive of Calamophyton and there is not adequate reason for comparing them precisely with the characteristic nodal structure of articulates such as the calamites and Equisetum. The Sterile Appendages (Leaves) A. On the main stem The sterile appendages or leaves on the main stem are somewhat more robust than those borne higher up on the plant. Some appear as simple unforked struc- tures, others are branched while some represent only the basal portions of branched leaves. The best preserved examples were found on specimens Nos. 5012/337 (fig. 1) and No. 5011/609 (part and counterpart, figs. 20 and 21). Some leaves of 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN No. 5012/3 37 are reproduced at a magnification of X 5 on plate 1; they represent the appendages shown in fig. 1 at points a (fig. 2), b (fig. 3), c (fig. 4) and d (fig. 5). It will be noted that the most profusely branched leaf reached four orders of divisions and was found higher up on the stem than the other examples One leaf was found on the main stem of the right hand specimen shown in figure 7 (point a) which contained a clearly defined vascular strand which could be followed almost to the tip. This leaf is shown enlarged in fig. 1 1 and a portion of the vascular strand is shown in fig. 10; the tracheids are apparently of the annular type. In specimen No. 5011/609 (fig. 20) the leaves display a tendency toward a whorled arrangement but it seems significant to note that the transverse bands, previously considered as characteristic of Calamophyton, are not evident here and, as we have noted above, the transverse bands in the stem in fig. 1 are not cor- related with any regular arrangement of the leaves. B. On the branch system above the main stem It is important to note that, prior to degagement, specimens give a misleading picture of the leaves since only about half of the appendage is exposed by the initial split of the rock specimen; this has been noted for other Devonian fossils in previous studies by Leclercq (1940, 1957). Consequently a considerable number of leaves were degaged or excavated with a small hammer and fine steel needles. Leaves were selected for detailed study that appeared to be well preserved and that were clearly identifiable on both part and counterpart. The leaf was degaged away, starting at the distal end, and followed toward the point of attachment to the stem until a branch was encountered. When this was done to both part and counterpart no question remained as to whether the entire leaf had been revealed. This resulted in the partial destruction of some leaves but the results obtained fully justified the procedure. In several instances we were fortunate in obtaining the desired information without destruction of the leaf. Figure 8 shows a leaf from specimen No. 5009/ 588 (fig. 7, b) which appeared initially with the Y-shaped terminal portion (a) attached at point b. Due to a fortunate cleavage of the rock it was possible to remove this terminal portion intact and a comparable branch c was revealed directly underneath. The chip was glued in position as shown in the photo, thus preserving the entire leaf. In summary then, we are dealing with a leaf that forked at point b to form two equal branches {a and c) and the latter in turn dichoto- mize at right angles to the first dichotomy. A restoration of the leaf is shown in text fig. 1. Several other fortunate excavations of this sort revealed the same three-dimensional branching pattern without destruction of the leaf. Several other examples, also taken from specimen No. 5009/588, will be men- tioned to record variations in the gross morphology of these leaves: LECLERCQ & ANDREWS CALAMOPHYTON BICEPHALUM 8 ANNALS OF THE MISSOURI BOTANICAL GARDEN Text figure 2 shows another leaf from the right hand specimen in figure 7 (e) ; this is similar to the preceding example but the initial dichotomy takes place closer to the proximal end and the secondary forking results in shorter branches. Figure 12 shows a leaf attached at point c in figure 7. This was revealed initially, partly in side view and partly in surface view, and no further excavation was conducted. The leaf divides first at a, and then both branches divide at right angles to the first division at b. At the upper of the b divisions one of the two secondary branches was lost but this is indicated in the restoration drawing given in text fig. 4. The upper portion of figure 13 shows another leaf from the same specimen (fig. 7, d) . In this case the entire leaf was exposed initially and it is included to illustrate an example in which four orders of branching are evident; it is shown in text fig. 3 as an aid in interpretation. This represents the maximum degree of branching that we have observed in the leaves of Calamophyton. Other leaves with four orders of branching have been carefully uncovered on the part and counterpart of specimen No. 5009/588 (figs. 9, 19). When well preserved they appear similar to the one shown in fig. 13 and text fig. 3. Another example (fig. 5) of one with four orders of branching is shown in the fertile specimen No. 5012/337 at point d. It is important to note that the leaves inserted in a similar position on a sterile branch system were small and wedge-shaped. Many other examples could be cited but they would only duplicate the results obtained from those cited above. The ones chosen reveal the typical branching pattern and the variations that are encountered with reference to the number of orders of branching. There are numerous examples of the type shown in figure 8 and occasional variants from this pattern reveal a lack of forking of the secondary branches but the examples shown in figures 9, 12, 13 and 19 with three or four orders of branching seem to be most typical. It is pertinent next to summarize the evidence that indicates the leaves are three-dimensional structures. In most cases it could be observed that the first dichotomy resulted in two branches that were directly superimposed; this is well shown in figure 8 where branch a was directly above branch c. In the second division, however, the resultant two segments lie apparently in the same bedding plane. In most cases where two or more orders of branching were present we are virtually certain that the orientation of successive branchings was at right angles, except possibly the fourth order. Additional evidence comes from examples such as the one in figure 12 (text fig. 4) in which irregular fracturing of the rock revealed a leaf initially in side and surface view. There is next the question of the cross-sectional shape of these appendages; they are shown in the restorations as having been terete and we feel that there is substantial evidence to indicate that this was the case. First, the leaves must have had considerable rigidity to have retained their three-dimensional form during fossilization. As many degagements reveal, one primary segment pushed down into the accumulating sediments so that the leaf as a whole was not flattened into one plane. It seems most unlikely that this would have been possible if the leaves had been thin, laminate structures. LECLERCQ & ANDREWS CALAMOPHYTON BICEPHALUM 9 Second, in well preserved specimens the thickness of the carbonaceous film is somewhat greater than might be expected if the leaves were laminate. Third, for whatever correlation purposes it may be worth, it may be noted that several early land plants that are known in the petrified state (Rhynia, Horne- ophyton, at least some Coenopterids) possessed a shoot system in which the ultimate branches are terete. The Fertile Appendages (Sporangiophores) The quality of preservation of the fertile appendages varies considerably and this is due in part to their relative state of maturity at the time of fossilization. Our most significant evidence has come from specimen No. 5011/609 in which the preservation is excellent and the sporangiophores retain most or all of their sporangia. In contrast to this, specimen No. 5012/337 seems to be one in which dehiscence took place some time prior to fossilization; most of the sporangia are shriveled, fragmentary or missing. Our basic description will, therefore, deal with a well preserved and apparently complete appendage from No. 5011/609. This is supplemented with a description of a second appendage of this specimen as well as several from No. 5012/337, partly to illustrate minor variations in the morphology of the fertile appendage and differences that result from the preservation of specimens at different states Specimen No. 5011/609 The fertile branch system as a whole is shown natural size in figure 21, the general morphology of the specimen being essentially the same as that found in the specimens shown in figs. 1, 7 and 14. It consists of an upright main axis of about 7 mm. in diameter which, at the top, divides into three primary branches which are of about the same size. One of the branches is dichotomized. The ultimate basal and terminal portions of the specimen are not preserved. Below the point of division the main axis bears sterile, rigid appendages. These represent for the most part only the basal portion of the whole appendage; some are more complete and display a part of the more distal, divided portion. These organs are similar to the "basal leaves" found on the main axis of specimen No. 5012/337 (fig. 1). The primary branches of the specimen are fertile throughout the portion that is preserved (fig. 29). They bear many whorls of sporangiophores only, there being no associated bracts. It has not been possible to determine the number of sporangiophores in a whorl; there are at least three and possibly as many as six. It was also difficult to decide whether, in successive whorls, the appendages were superimposed or alternate, though it is more probable that they were superimposed. It is important to note that here, as in the case of the sterile appendages, the initial exposure of the specimen gives a misleading picture of the general mor- phology of the sporangiophore. The latter is a three-dimensional structure and half or more of it may be covered by the rock matrix. In order to be sure of 10 ANNALS OF THE MISSOURI arriving at a correct understanding of the sporangiophore many of them were carefully uncovered under a binocular microscope using needles and a light weight hammer as described above. Successive stages in the degagement of a selected sporangiophore are illustrated in figures 23 to 27. Figure 22 represents the counterpart (specimen No. 5011/ 609B) of the initial stage shown in fig. 23. As an introduction to the detailed description it may assist the reader to note briefly the over-all branch pattern: the fertile appendage divides into two segments or "heads" (referred to as upper and lower), each of which bore three short side stalks and each of these in turn terminates in a pair of sporangia; the distal tip of each of the two segments is usually bifurcated but not always so. Thus the sporangiophore as a whole, composed of two similar heads, supports twelve sporangia if all develop. In figures 22 and 23 the upper segment (I) is shown from its base up to the ultimate preserved tip. This segment supports two side stalks, A and B, which are attached on either side at slightly different levels; two sporangia are borne on each stalk (fig. 23 and text fig. 6). Figure 24 reveals the third stalk, C, of the upper segment and one of its sporangia is visible; the second one is still overlain by stalk B although its distal part appears beside the two sporangia of the B stalk. Text figure 6 shows this upper segment or head with the three bifurcated stalks and their six sporangia in the position in which they were preserved. In figure 24 a portion of the lower segment or head may be seen behind the upper one. Figure 25 shows the lower segment (II) in connection with segment I as well as the three stalks A, B, and C, which are borne by it (segment II) ; this is also shown in text figure 7. Stalk A has two sporangia in connection; stalk B appears with one sporangium attached and the second lies under segment I; stalk C is just coming into view at this level. In figures 25, 26 and 27 the second sporangium of stalk B has been revealed by removing parts of segment I. Stalk C, which was recurved under segment II, has been uncovered by destroying parts of the latter; it is now clearly revealed and has one sporangium suspended in an anatropous position on each of its two divisions or pedicels. Text figure 7 shows this lower segment (II) with its three bifurcated stalks and their six sporangia. Text figures 6 and 7 when superimposed show the entangled stalks and sporan- gia of the entire sporangiophore as it was embedded in the sediment; text figure 8 is a reconstruction of the sporangiophore as we believe it appeared before fossiliza- tion and in natural position. Thus, in summary, the sporangiophore was a small, adaxially inclined appendage attached to the branch axis. At a point about one third of the way from its proximal end the sporangiophore branched into an upper and lower segment. Each segment was erect and rigid and supported three short side stalks fixed at two different levels; the first stalk was given off at a different level than the two others which appear generally to be given off simultaneously. Each of the three stalks is slightly bifurcated at its tip with a sporangium attached to each bifurcation. Usually the distal, rigid portion of each segment of the sporangiophore terminates in an irregular dichotomy. The sporangia were cylindrical sacs 2.0-2.7 mm. long and 0.5-0.7 mm. in LECLERCQ & ANDREWS CALAMOPHYTON BICEPHALUM 11 . Lower segment of same sporangiophore (II) ; see figs. MISSOURI BOTANICAL GARDEN diameter and pointed at the distal end. They were apparently strongly constructed for despite the entangled position in which they occur they are often neither broken nor opened. It is possible that the wall of the sporangium was rather thick, perhaps as a result of several layers of cells or having been fossilized when im- mature; the latter seems most likely. The mode of dehiscence has not been positively ascertained. Careful examina- tion of the sporangia on the holotype specimen (No. 5011/609) has revealed no distal pore but on a few sporangia a median longitudinal line is found on the ventral side (fig. 28) which may represent the place of dehiscence. It is evident from the numerous sporangiophores we have studied that their organization was essentially the same throughout; there are, however, some slight variations and in view of the rather surprising complexity of the fertile appendages as contrasted with previous accounts of Calamophytons it seems pertinent to sup- plement the description given above with somewhat briefer considerations of several other examples. LECLERCQ & ANDREWS CALAMOPHYTON BICEPHALUM 13 Text figure 9 is a camera lucida drawing of an appendage (No. 5011/609) prior to any treatment. The sporangiophore stalk divides into the two divisions; a considerable part of I is present but only the basal part of II. Referring to segment I, branch B is shown with its two sporangia Bl and B2 and one sporangium of the C branch is present. In order to expose segment II, which appeared to dip down into the rock, sporangium Cl was degaged away; after working through the sediment underneath sporangium Cl the II segment was revealed as shown in text fig. 10 and fig. 31. Three branches (A, B and C) were found departing from segment II. Branch C terminates in a single sporangium, Cl, but judging from its mode of attachment this is one of an original pair; branch B is terminated by one clearly defined sporangium (B2) and there appears to be another beneath it; the third branch (A) could not be followed. Text figure 11 and fig. 30 show the counterpart of the appendage; only seg- ment I is present. The sporangia Bl, B2 and Cl are readily correlated with their position as shown on the other face of the specimen (text fig. 9); there is some question as to the sporangium labeled Al and a fifth one appears to the right of sporangium B2 whose position on the segment is undetermined. Text figure 12 brings this information together as a semi-restoration. It may be noted that there were not less than eight sporangia borne by the entire appendage and the number was probably closer to ten or twelve. Specimen No. 5012/337 Some significant information was obtained from this specimen (fig. 1) con- cerning the general branching pattern of the fertile appendages but, as noted above, very few sporangia could be identified intact due to the fact that dehiscence had apparently taken place some time prior to fossilization. It should be noted that there is no counterpart of this specimen; when it was found during the quarrying operations an extended search was conducted for the other side but it was appar- ently completely shattered and lost. The three sketches shown in text figs. 13-15 show an appendage before and after degagement, and a semi-restoration respectively. The initial appearance was essentially as shown in text figure 13 aside from a slight amount of degagement that was necessary to expose branches B and C. The segment was then degaged away, starting at the distal end and extending down to the point indicated by line — o. The underlying segment (II) was then detected and when completely excavated it appeared as shown in text fig. 14. The semi-restoration of this appendage (text fig. 15) thus shows it as consist- ing of two main branches, I and II; II produces at least two secondary branches and terminates in a forked tip; I produces three secondary branches. All of the sporangia had been shed prior to fossilization. A second example (fig. 6 and text fig. 16) is included since it gives some interesting information concerning the variation in the general morphology of the fertile appendages and particularly the ultimate terminations. It is virtually certain that the upper portion was attached as indicated by the dotted line but this was not positively demonstrated. The essential features of this appendage 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN seem to be as follows: the main stalk branches into two segments which in turn divide, each giving off several side branches, and terminate in unforked distal tips. Although specimen No. 5012/3 37 is the largest one in the collections and the most completely preserved, with reference to the main branch system as a whole, none of the ultimate terminations of the branches are preserved; these probably extended out at least 10 or 15 cm. beyond what is shown in the photo (fig. 1). The ultimate appendages, both sterile and fertile, are for the most part rather poorly preserved and we interpret it as a plant, or portion of a plant, that had shed its spores and undergone partial decay prior to fossilization. Several fertile appendages are sufficiently well preserved as to offer evidence of their relationship with the new species C. bicephalum. Diagnosis Calamophyton bicephalum Leclercq and Andrews, sp. nov. Plants consist of a main stem, the basal portion of which is unknown, which divides in a digitate fashion into two to several branches; these branches in turn divide in more or less equal dichotomies. Transverse striations are occasionally present on the main stem and lower portions of the primary branches. The shoot system, above the main axis, appears to be predominantly fertile or sterile. The sterile appendages (leaves) are somewhat more robust on the main stem than above, with a single vascular bundle, and dichotomize one to four times in a three-dimensional pattern and attain a length of about one cm. Fertile appendages are aggregated to form a specialized branch but not in definite cones; a fertile appendage consisted of a basal stalk which divided into an upper and lower seg- ment; each segment bore three short side branches which terminate in a slight bifurcation with a pendulous sporangium attached to each, resulting in a total of twelve sporangia; each of the two main segments terminates in a short dichotomy. Sporangia are cylindrical, pointed at the distal end, 2.0-2.7 mm. long by 0.5—0.7 mm. in diameter, with possibly a line of dehiscence on the ventral side. Holotype: No. 5011/609. Paratypes: No. 5012/337, No. 5009/588. All pre- served in the Laboratoire de Paleontologie Vegetale, Universite de Liege. Horizon: lower Givetian, Middle Devonian. Locality: Carrieres Brandt (north quarry), Goe, Belgium. Discussion With the exception of the general organization of the branch system the new species of Calamophyton described here presents characters, particularly in the nature of the ultimate fertile and sterile appendages, that are quite new. In these features our plant differs sharply from Calamophyton as portrayed in all recent text-books. It was quite surprising to find these appendages so complex and it might be supposed that we are actually dealing with plants that should not be identified with this genus. Yet it may be noted that our specimens showed, on the initial split of the rock, rudimentary leaf -like organs (sterile appendages) which are small and once or twice divided apparently in one plane as previously described for C. primaevu being once forked and in on to the sporangiophore of C. e same holds true for the sporangiophore; for impression before degagement (figs. 22, 23) of plane. This aspect is apparently very similar (Leclercq, 1940) with the difference that the 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN small axis of the sporangiophore of C. bicephalum is prolonged as an erect append- age. Thus, in order to determine whether C. primaevum K. and W. actually had three dimensional leaves a specimen of this species (figs. 17, 18) formerly described by Aderca (1932) was selected for re-investigation. Figure 15 shows two leaves that were selected as they appeared initially; both dichotomize and of the two resultant branches the left one in both undergoes another forking and, in the case of the lower leaf the left fork is again apically notched. Beginning at their apical extremeties the two leaves were degaged away down to the point indicated by the lines o o, at which point a branch was found dipping down into the rock matrix. When completely exposed these underlying branches appeared as shown in figure 16 distal to the line o o. The latter were thus directly beneath the branches shown in figure 1 5 and in both leaves the second dichotomy was at right angles to the first; these leaves are shown restored in text figure 5. Thus, in view of the identical morphology found in C. bicephalum we consider the three dimensional branching to have generic value. The prominent characters of C. bicephalum are the three dimensional sterile and fertile appendages. The former was a fairly rigid, terete structure with somewhat varied form; wedge-shaped ones, notched at their apex, were found along the lower portions of the primary branches. Higher up the branching of the leaf became more pronounced until the "mature" stage, characterized by four orders of branching, was attained. Attention may be called to the fact that such mature leaves were found on the lower portion of the primary branches of fertile twigs. It may next be of interest to consider the possible homology of the sterile and fertile appendages. In 1936, Eames noted that "from the evidence provided by Hyenia and Calamophyton it is apparent that 'leaves' and the fertile tips in this group are homologous". Our study indicated that this is undoubtedly true if one compares a "mature" leaf with a sporangiophore. The latter divides into two main branches, referred to previously as superior and inferior segments, which may correspond to the initial dichotomy of the sterile appendage. Beyond this point the fertile appendage is more complex in that each segment produces three side branches each of which bears a pair of sporangia while the segment proper is elongate and usually terminates in a forked apex. Whether or not the sterile appendages may be termed "leaves" is perhaps a matter of opinion but there would seem to be no doubt that they functioned as such. The present study is an initial one dealing with the fossil plants from the Goe locality. The difficulty of working with Devonian plants is emphasized by this investigation but in no less degree it is demonstrated that reliable information may be obtained from such ancient land plants when they are well preserved. It seems pertinent to add a few comments concerning previous work with both Hyenia and Calamophyton since they are undoubtedly closely related. Of the several species of these two genera that have been described to date none is known The problem of making significant comparisons with previously described species is rendered difficult by the fact that during the recent war the material LECLERCQ & ANDREWS CALAMOPHYTON BICEPHALUM 17 described by Krausel and Weyland (1926, 1929, 1932) was destroyed although two fragments of the fertile branch system of C. primaevum, from Dr. Krausel's collections, are preserved in the Department of Geology of the University of Cologne. The two fragments on which C. renieri (Leclercq, 1940) is based are preserved at the University of Liege. In the light of what has now been found in C. bicephalum it may be desirable to rework, with the degaging technique, the available material of the two other species to try to determine the real value of the three specific epithets. It is, however, questionable whether the remaining speci- mens of the earlier described species are sufficiently well preserved to allow effective comparison, consequently our only course seemed to be to assign a new specific name to the Goe specimens. In the light of what is now known it seems safe to assert that the only dependable difference between Hyenia and Calamophyton lies in the general habit of the two. Even this is complicated by the fact that we do not know anything about the basal part of the shoot system of Calamophyton. Our specimens do, however, suggest a habit that may be worth a few words of speculation. It seems likely that C. bicephalum consisted of upright shoots that were predominantly fertile (fig. 21) or sterile (fig. 7, lower right). If this was the case it seems unlikely that the fertile shoots would have had sufficient photosynthetic tissue to have maintained themeselves independently. In explanation it may be supposed that both were borne either on a common erect small "trunk" that would have given the plant a shrub-like appearance, or they were borne on a common rhizome with a relationship comparable to that found in the modern Equisetum arvense. It is expected that further studies of the Goe fossils will contribute toward solving this and other problems relating to both Calamophyton and Hyenia. Summary The material on which this study is based came from the lower Givetian (Middle Devonian) of Belgium. A new species, Calamophyton bicephalum Leclercq and Andrews, is created in view of the complexity of the fertile appendages which contrast with homologous structures in the previously described species C. primaevum Krausel and Weyland and C. renieri Leclercq. The investigation has dealt chiefly in elucidating the structure of the sterile and fertile appendages. The sterile appendages (leaves) are shown to be three dimensional organs that forked once (young leaves) to four times (mature leaves) and were probably rather rigid and terete in cross section. The three dimensional structure of these appendages is considered to have generic value. The fertile appendage branched into upper and lower segments; each was erect and supported three curved side stalks attached at slightly different levels; each of these stalks terminated in a pair of pendulous sporangia. Thus the entire appendage bore twelve sporangia when all developed. The sporangia were cylindrical and pointed at the distal end, with possibly a line of dehiscence in the ventral side. 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN Acknowledgements Je desire exprimer ma gratitude au Fonds National de la Recherche Scientifique de Belgique pour les deux campagnes de fouilles qu'il a subsidises en 1949 et 1952 et au cours desquelles j'ai recolte le materiel ici etudie. Je remercie le Patrimoine de l'Universite de Liege qui m'a permis de recevoir le Professeur Henry Andrews pendant quatre mois a mon laboratoire. A Monsieur J. Damblon vont mes sinceres remerciements pour Pattention et le soin avec lesquels il a execute la reconstitution du sporangiophore reprSsente figure 8 du texte. Je suis particulierement reconnaissante a Monsieur Jose Lewalle, Assistant, et a Monsieur Maurice Discry, Preparateur-technicien, de Paide continue qu'ils ont apportee a 1'execution des nombreux documents photographiques necessites par cette etude. (Professeur S. Leclercq) This investigation was carried out during my tenure as a Fellow of the John Simon Guggenheim Memorial Foundation; thanks are also due the National Science Foundation for aid in defraying publication costs and in other ways. (HNA) Literature Cited Handbuch der Palaobantanik. Munich, pp. 1-708. . Contributions to the Devonian flora of western Norway. III. usel, K. and H. Weyland, 1926. Beitrage zur Kenntnis der Devonflora. T. . 1929. Beitrage zur Kenntnis der Devonflora. in. Senckenberg. natui . 1932. Pflanzenreste aus dem Devon. III. Uber Hyenia N 274-280. lercq, S. 1940. Contribution a l'etude de la flore du Devonien de Liegeois, R. 1956. Excursion dans le Mesodevonien rj. Soc. Geol. de Belgiq, Sci., Mem. 4, 14 (fasc. 3 956. Excursion dans le Me ' Ja Vesdre. Ann. W Tw.nl A. Vol. II. Bryophytes and Pteridophytes. idy of fossil plants. London, pp. 1-201. ; ig. 1. A large specimen shown natural size. Figs. 2-5. I :s a-d, respectively, on specimen shown in fig. 1 ; all ma ' preserved fertile appendage; see also text fig. 16 and LECLERCQ & ANDREWS— CALAMOF ANNALS OF THE MISSOURI BOTANICAL GARDEN & CPLANATION OF Plate PLATE II Calam n bicephalum Lech 7-13: specimen N :rcq and Andre Two specimens 9. Leaf showin a leaf shown ir a in fig. 7; X 3. Fi, 1; X 62. Fig. 11. ;. 12. Leaf from F 8. Leaf taken g; X 5. Fig. 1 Leaf with va: :oint c in fig. 7 LECLERCQ & ANDREWS— CALAMOPHYTON BfCEPHALUM #' f f i LECLERCQ & ANDREWS CALAMOPHYTON BICEPHALUM 21 Calamophyton bice^hahim Led men No. 5007/346. XI. en No. 5006, X5, shown as they appeared originally, fig. 15 were degaged away. These two leaves are ANNALS OF THE MISSOURI BOTANICAL GARDEN Explanation of Plate PLATE IV Calamophyton bicephalum Leclercq and Andrews Fig. 19. Leaf showing four orders of branching from specimen No. 5009/5 8! Fig. 20. Main stem of specimen No. 5011/609, XI. Fig. 21. Holotype specim 22-27 show stages in the excavation of a fertile appendage from specimen No. 501 for explanat.on see text (p. 10); all X5. Fig. 28. Portion of fertile appendag specimen No. 5011/609, X5. [ 60'); 26 27 28 LECLERCQ & ANDREWS— CALAMOPHYTON B1CEPHALUM LECLERCQ & ANDREWS— CALAVIOPHYTON BICEPHALUM LECLERCQ & ANDREWS CALAMOPHYTON BICEPHALUM 23 Explanation of Plate PLATE V Calamophyton bicephalum Leclercq and Andrews Fig. 29. Upper portion of holotype specimen No. 5011/609 (as I enlarged three times. Figs. 30, 31. Part and counterpart of a fertil specimen No. 5011/609, X5; see also text figures 9-12 and description i CLASSIFICATION AND PHYLOGENY IN THE ORCHIDACEAE ROBERT L. DRESSLER and CALAWAY H. DODSON Introduction The Orchidaceae form one of the largest families of ; one of the most fascinating by reason of their diversity and specialization in floral structure. A satisfactory classification of the orchids into tribes and subtribes is not yet available. The most commonly used system, that of Schlechter, has for some time not been in accord with the rules of nomenclature, and has many features which may be questioned on botanical grounds. The recent International Botanical Congress has clarified the rules concerning the nomenclature of categories between family and genus, and provides an occasion for a reevaluation of nomen- clature in the Orchidaceae. In reviewing the groups within the Orchidaceae we have, of necessity, made a number of observations on relationships and probable phylogeny within the family. These form the final portion of this paper. While many workers have described new genera and species of orchids, there has been very little monographic work, and we may safely say that the family has really been very little studied, considering its size and complexity. Until there has been a great deal more systematic study of the family it will not be possible to present a finished system of tribes and subtribes. Consequently, no new taxa are presented in this paper, but we have attempted only to review and evaluate pre- vious systems of classification, and to present a tentative system, with synonymy. Swartz, in 1880, first divided the orchids into those with a single fertile anther and those with two fertile anthers, thus providing the basis for the subfamilial divisions now recognized. Lindley, in his "Orchidearum sceletos" (1827), was the first to divide the family into tribes. In this work he recognized eight tribes. Later, in "The Genera and Species of Orchidaceous plants," only seven tribes were maintained, with "sections" or "divisions" recognized under some of these. Reichenbach never presented a detailed system of orchid classification, and his categories were vague and inconsistent in both rank and spelling (1852, 1884). Bentham (1881), in preparing a system for Genera Plantarum, recognized only five tribes, and delineated 27 subtribes under these. Pfitzer (1887) criticized the classification of Bentham and offered a revised system, based primarily on vegetative features. While the rank of Pfitzer's categories was not very clearly indicated, 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN one finds that the groups with names ending in "-inae" are referred to as "Tribus." Thus Pfitzer's classification recognized 32 tribes, with a number of subtribes. This same system was followed in "Die Natiirlichen Pflanzenfamilien," with only minor changes. Pfitzer's classification was the basis for Schlechter's posthumously published "System der Orchidaceen" (1926). Schlechter, however, recognized only four tribes and treated the remaining tribes and subtribes of Pfitzer all as subtribes, of which he enumerated eighty. Some of Schlechter's subtribes were characterized in earlier papers (1911, 1915) as "Gruppen," but only in the 1926 paper were they treated as subtribes. In studying Schlechter's work, one often feels that his system was published in unfinished form, and that he might have presented a much more coherent system, had he lived to complete it. Mansfeld (1937) has reviewed Schlechter's system and made some modifications, but most recent authors have followed Schlechter's original system with little change. Recently Hawkes and Heller (1959) have presented a list of subtribes and genera in which they recognize no less than 88 subtribes. Schlechter might well be characterized as a "splitter"; he followed very narrow concepts at all levels of his classification. Subsequent workers have reduced a large proportion of his genera and species to synonymy, but most have accepted his tribes and subtribes with little question. Actually, many of the subtribal bound- aries drawn by Schlechter have proven to separate closely related genera. In several cases genera assigned to different subtribes have proven interfertile. While Schlechter recognized eighty subtribes in the subfamily Orchidoideae and several others have since been proposed, we tentatively recognize only about forty in our proposed revision. We believe that future study may further reduce the number of subtribes to be recognized. While the rules of nomenclature permit an almost excessive number of categories between genus and species (subgenus, section, sub- section, series and subseries), there are relatively few categories between family and genus. Where Schlechter's subtribes seem useful, even though too closely related or too poorly defined, we have indicated them as "alliances." These are not intended to have formal nomenclatural status. To recognize them as subtribes would tend too much toward taxonomic inflation, and would tend to obscure the really close relationships which exist within the family. Some authors have cast doubt on the validity of genera which are interfertile. While we do not believe that a fertility criterion (alone) can be applied for generic status in the orchids, we do feel that interfertile genera should not be placed in separate subtribes. In every case where authentic hybrids have been reported between subtribes, however, the morphological evidence, alone, favors their union into a single group. In those cases where we feel quite sure that closely related genera were separated in Schlechter's system, we have united them into a single subtribe. In other cases, however, we have deferred judgment because of insuf- ficient familiarity with the plants involved. The main difference between Schlechter's system and our own is that he em- phasized differences, while we are seeking resemblances. His system was primarily analytical and aimed at identification (though often faulty for that purpose), while our own is synthetic, as we believe these higher categories should be. This DRESSLER & DODSON CLASSIFICATION IN ORCHIDACEAE 27 is not to belittle the key features used by Schlechter. Where valid, these are still available for keying groups within the subtribes, but relationships are, we hope, more clearly shown in our system. In many cases the key features chosen by Schlechter will separate only a portion of the genera in closely related or artificial groups. There are primarily three features of the International Code of Botanical No- menclature which affect the choice or form of names used in this paper. In the case of subfamilies, tribes or subtribes, the name of any taxon which includes the type of the next higher taxon must be based on the same stem as the name of the next higher taxon. Thus "Ophrydeae" and "Platantherinae," for example, must be replaced by Orchideae and Orchidinae, respectively. This requirement, which was only recently added to the rules (by the 1959 Congress) causes a few changes, but should cause no confusion, and in general makes the taxonomy at this level more logical. The requirement of the suffixes -oideae, -eae and -inae for subfamilies, tribes and subtribes has been in the rules for some time, but has often been ignored by those dealing with orchid nomenclature. The same is true of the principle of priority, which demands the use of names proposed by Bentham, where these differ from those used by later workers. Tribal Delineation The separation and chai is relatively clear. Each forms ; recognized. In the subfamily Orchidoideae the situation is rather different. The tribe Orchideae is distinctive and rather easily characterized, though clearly related to the Neottieae. The distinction of further tribes is much less clear. In Schlechter's system all of the genera with mealy or sectile pollen (except the Orchideae) are grouped in the tribe Polychondreae (= Neottieae), while the genera with hard, waxy pollinia are separated as the more advanced tribe Kero- sphaerae (= Epidendreae). As Mansfeld has shown, the distinction between mealy and waxy pollinia is neither practical nor natural. Some genera of the Bletiinae have, according to Mansfeld, mealy pollinia. Certainly most of the remaining genera have rather soft pollinia. Some genera of the Sobraliinae have hard pollinia and this subtribe shows close affinity to the Thuniinae and the Epi- dendrinae. Similarly, the Arethusinae have mealy pollinia, but Crybe and Jimensia (Bletilla) are closely related to Bletia. There is a complete series ranging from free pollen grains to the hard ceraceous pollinia of the Oncidiinae and Sarcanthinae. Any arbitrary degree of cohesion chosen as a dividing line would split natural genera and subtribes. Mansfeld (1937) placed the Arethusinae and Sobraliinae in the Epidendreae, and this action is supported by the morphological studies of Hirmer (1920). Such a system, though, leaves no practical way of distinguishing the Epidendreae and Neottieae and is, we believe, still unnatural. The subtribes Vanillinae, Pogoniinae and Gastrodiinae seem to show much closer relationship to the Sobraliinae and Arethusinae than to the other subtribes of the Neottieae. By placing these subtribes in the Epidendreae one achieves a system which is both 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN more natural and more practical. By this arrangement the great majority of the primitive Epidendreae (with mealy pollinia) have incumbent, operculate anthers, like those of Phajus or Cattleya (see fig. 3D). The position of the anther in the more advanced Epidendreae is extremely diverse, but these are easily distinguished by the truly hard pollinia. A few species of primitive Epidendreae, such as Triphora and some species of Epistephium and Elleanthus, have erect anthers. Thus the position of the anther is not a fool-proof key feature, but it does seem to provide a better practical separation, as well as a more natural classification. Several authors have distinguished the tribes Epidendreae and Vandeae on the basis of pollinia structure, but a clear distinction proves to be difficult. The presence or absence of a stipe is one feature which has been used for this purpose, but some genera which would unquestionably belong in the "Vandeae" have little or no stipe, while a few other genera which are not closely related have stipes or stipe-like structures (Genyorchidinae, Thecostelinae, some species of Polystachya) . Another feature which is characteristic of the "Vandeae" is the presence of super- posed pollinia; yet Coelogyne and some species of Thuniinae and Polystachya have more or less superposed pollinia, but seem otherwise not referable to the "Vandeae." There seems to be no feature or combination of features which will serve to separate the more advanced Orchidoideae into two clear-cut main groups. There is a general trend from plants with terminal inflorescence and relatively simple pollinia to those with a lateral inflorescence and highly specialized pollinia, but there is no sharp break and the relationships seem too reticulate to admit the separation of two tribes on this basis. The subtribes of the "Vandeae" (Cyrtopodiinae to On- cidiinae as listed on p. 29) seem to represent a relatively homogeneous and more specialized offshoot from some of the several evolutionary lines within the re- mainder of the Epidendreae. All of these subtribes appear to have been derived from more or less Eulophia-like ancestors. For this reason, it is sometimes con- venient to consider them as a unit, and a better understanding of the subtribes related to the Cyrtopodiinae may indicate a sharper break between the two main divisions of the Epidendreae than is now evident. While the Epidendreae form a natural and closely knit group, the Neottieae are more diverse in terms of relationship. Even with the removal of the Vanillinae, Pogoniinae and Gastrodiinae, it is not certain that the Neottieae form a really natural group. Including all of these genera in one tribe because they all possess mealy pollen is somewhat comparable to a hypothetical grouping of Vanilla, Selenipedium, and Apostasia into a single taxon because of seed characteristics. Mealy pollen, like the sclerotic seed coat and the lateral anthers of the Cypripedi- oideae, is a feature which was doubtless found in all orchids at an early stage in orchid evolution. Its occurrence in two or more otherwise dissimilar groups is scarcely strong evidence of relationship. Since the present paper is primarily a review of orchid classification, any major reorganization of the Neottieae or the Cypripedioideae must be deferred for the present. The possible groupings within these taxa are discussed in the final section on tribal phylogeny (p. 62). DRESSLER & DODSON CLASSIFICATION IN ORCHID ACEAE 2 Subfamily Cypripedioideae Tribe 5. Epidendreae (Continued) Tribe 1. Apostasieae Subtribe Coelogyninae Tribe 2. Cypripedieae Epidendrinae Subfamily Orchidoideae a . Epidendrum alliar Tribe 3. Neottieae b. Eria alliance Subtribe Limodorinae c. Polystachya allian Chloraeinae d. Glomera alliance Rhizanthellinae e. Podochilus allianc* Pterostylidinae f. Arpopbyllum allia Pleurothallidinae Neottiinae Adrorhizinae Thelasiinae Diuridinae Ridleyellinae Cryptostylidinae Liparidinae Prasophyllinae Dendrobiinae Genyorchidinae Spiranthinae Thecostelinae a. Tropidia alliance b. Goodyera alliance Cyrtopodiinae c. Spirant hes alliance Catasetinae d. Cranichis alliance Cymbidiinae Tribe 4. Orchideae Sarcanthinae Subtribe Epipogiinae Stanhopeinae Orchidinae Maxillariinae Disinae a. Zygopetalum alliai a. Disa alliance b. Lycaste alliance b. Satyriutn alliance c. Maxillaria alliance Coryciinae Pachyphyllinae Tribe 5. Epidendreae Cryptocentrinae Subtribe Vanillinae Oncidiinae Gastrodiinae a. Oncidium alliance Pogoniinae b. Ornithocephalus all Sobraliinae c. Dichaea alliance Thuniinae Arethusinae Subtribes of uncertain position Bletiinae Grobyinae Collabiinae Pachyplectrinae Proposed Phylogene List Above is given a list of the tribes and subtribes which we recognize, followed by a tentative key to tribes. We have, in nearly all cases, changed the endings of the group names to accord with the rules of nomenclature. Very few of them were published in the appropriate form, though status was clearly indicated. We have attempted to arrange the groups in a "phylogenetic" sequence; that is, we 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN have tried to place the more primitive members at the beginning of each group, and we have tried to place closely allied groups together, where possible. The number of subtribes which should be immediately adjacent to the Epidendrinae in any "natural" system, but only two can be so placed. "We have indicated groups of related subtribes by lines, but interrelationships within the Epidendreae are better shown in figure 1 (p. 51). Keys to continued reliance on unworkable keys. Since previously published keys to the Orchidaceae have proven to be inadequate in many features, we have prepared these keys de novo, as much as possible. Comparison will show Schlechter's key to be much simpler in many respects. To this we can only reply, "Yes, but his key doesn't work." There will surely be many sections in these keys where a similar complaint is justified. It is extremely difficult to write a workable key to the orchids of the world. The person working in a single continent has an easier time of it, and would do well to prepare his keys independently, rather than trying to adapt either these or Schlechter's keys. An English translation of Schlechter's key, with some revision, is given in Withner's recent book (Schweinfurth, 1959). Key to Subfamilies and Tribes style or the anther and stij ;mT connected by ttzzsrs r«rSift"2 ). Perianth essentially regula r, the lip never deeply saccate; fert ile anthers 2 or 3, spicuous. flattened' median i ). Pollinia 2 to 8, hard, waxj l^'tyle^relative'ly'thicl 1Z™^ Pollinia 2 or 4, soft, me deciduous; leaves usually herbaceous, not articulate; growth usu S^STwiS } - Sw.'sass^ ^ba'se of The" pollinia 6 ). Anther terminal and operculate (incumbent) or rarely erect, l Actantbus) ; stems without corms or other thickenings Synonymy and Discussion In the following section we give keys to subtribes and 1 K*mi or .su.ll} more or less Neottieae ist the subtribes alpha- DRESSLER & DODSON CLASSIFICATION IN ORCHIDACEAE 31 betically under the tribes, with synonymy and discussion where appropriate. We have not given full citations with the names of tribes and subtribes, but the papers containing new tribes or subtribes are indicated by an asterisk in the bibliography. We estimate that there are about 600 distinct, valid genera in the Orchidaceae. A really critical enumeration of genera is not yet possible. The lists given here are based primarily on Schlechter (1926), and doubtless contain some genera which do not merit recognition, while omitting others which should be listed. Only for the Epidendrum and Oncidium alliances can we indicate with some confidence the genera which will be maintained by critical revision; and, even here, there are genera of which we have not yet seen adequate or living material. If these alli- ances are representative of the family, the total number of valid genera may be well under 600. Subfamily CYPRIPEDIOIDEAE Lindley (Diandrae Kunth, Pleonandrae Pfitzer [1903], Apostasioideae Wettstein) Tribe 1. APOSTASIEAE R. Brown (Pulverae Blume, in part) Some authors have excluded the Apostasieae as a separate family. We, how- ever, agree with Rolfe (1909), J. J. Smith (1934), Mansfeld (1934) and Holttum (1953), that these plants cannot logically be excluded from the family without also excluding the Cypripedieae, and we feel that neither action is desirable. A classification which excludes the Apostasieae because they are inconspicuous and retains the Cypripedieae because they are showy is scarcely acceptable. It is possible, of course, that detailed study will show the Apostasieae to be basically different from other orchids in some features. As far as present knowledge goes, they are primitive orchids, and quite as closely related to some Neottieae as these are to the other orchids. As Godfery (1932) indicates, there is little evidence of close relationship between the Apostasieae and Cypripedieae, even though they show the same basic plan of flower structure. It is quite possible that the current subf amilial division is artificial. Apostasia, Neuwiedia. Tribe 2. CYPRIPEDIEAE The four genera of ladyslippers form a relatively uniform relic group. They are markedly divergent from most other orchids in that the median anther is repre- sented by a large shield-like staminode. In spite of their differences, the ladyslippers are orchids in good standing. The three abaxial stamens, resupination, reduction in seed structure, mycorrhizal relationship, and the less obvious features which pre- dispose the family to evolution as epiphytes are themes which run throughout the family. The primitive features to be found in Selenipedium are strongly reminis- cent of those found in other primitive orchids, though they do not, of course, necessarily indicate close relationship. Mansfeld (1937a) notes some resemblances between the Cypripedieae and Epipactis. Cypripedium, Paphiopedilum, Phragmipedium, Selenipedium. 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN Subfamily ORCHIDOIDEAE (Monandrae Kunth) Tribe 3. NEOTTIEAE Lindley (Granulosae Blume [in part], Pulverae Blume [ir part], Goodyereae King & Pantling, Listereae King & Pantling, Polychondrea< Schltr., Epipactieae Hatch [illigitimate, because nomenclaturally superfluou when published] ) Key to Subtribes saprophytes in Neottia) (north temperate) and Aphyllorchis) ; lip usually divi ■ es often basal; lip no hinged, with a ret oitzr;: 11 U VtODORINAE Lip nge, actively motile (scnue) v ^l ut ^x ely motile (Sout V.::: plants with Cu Lisdcla roots and mn usually without distinct wings or audicle-like stipe a, habit van i' ' ..P RA SO 'HYLIIVXF ,., ;, 8 len by the enfolding lip base; sepals or petals relative Chloraeinae Pfitzer (Caladeniinae Pfitzer, Thelymitrinae Pfitzer, Acianthin; Schltr., Corysanthinae Schltr., Megastylidinae Schltr., Corybasinae Mansf. [nc validly published, unless later (1954) reference to Corysanthinae Schltr. 1 taken to validate its publication as a new name. Corysanthinae, however, a valid name, though based on a synonym] ) This is the group which includes most of the bizarre Australian genera. Tl striking modifications have led to the naming of several subtribes, but these do n( seem tenable, at least as previously delimited. Thelymitra is distinctive because < its nearly regular perianth, which is frequently blue, but it is closely allied to tl DRESSLER & DODSON CLASSIFICATION IN ORCHIDACEAE 33 other genera through Calocbilus, Adenochilus and Glossodia. Some species of Thely- mitra are unusual for this group in the attachment of the rostellum or viscidium to the apex of the pollinia, but this is not consistent within the genus. Some species of Acianthus are very unusual in the form of the column and the position of the anther (operculate), but A. reniformis is more representative of the subtribe in these features; Mansf eld's action in grouping Acianthus with Caladenia, thus seems correct. Corybas is closely related to Caladenia. The three American genera Chloraea, Bipinnula and Asarca are somewhat distinctive in habit, but the other American genus, Codonorchis, is closely related to both Chloraea and Caladenia. Most of the Australian genera show a characteristic pitted, conic anther. Acianthus, Adenochilus, Asarca, Bipinnula, Burnettia, Caladenia, Calocbilus, Chilo- glottis, Chloraea, Codonorchis, Corybas, Epiblema, Eriochilus, Glossodia, Leptoceras, Lyperantbus, Megastylis, Rimacola, Thelymitra, Townsonia. Cryptostylidinae Schltr. Cryptostylis is a distinctive genus, though without striking key features. Some species mimic insects and are involved in the strange relationship of pseudo- copulation. This subtribe, the Diuridinae and the Prasophyllinae seem to form a distinct group with column structure similar to that of the Spiranthinae. The degree of relationship to the Spiranthinae is uncertain. Coilochilus, Cryptostylis. Diuridinae Bentham This group is of special morphological interest because of the large staminodia and because the filament and style are scarcely united into a column. Some Spiran- thinae show similar columnar structure, though without the staminodia. Diuris, Orthoceras. Limodorinae Bentham (Cephalantherinae Pfitzer, Epipactiinae Godfery) The Limodorinae include genera which are quite primitive in some respects, and, being largely European, they have been studied much more than other primitive orchids. Aphyllorchis, Cephalanthera, Epipactis, Limodorum. Neottonae (Listerinae Schltr.) These genera are distinctive in the sensitive rostellum, which forcibly extrudes a viscid droplet when touched. The anther is either erect or somewhat incumbent on the subequal rostellum. These genera show some resemblances to the Limo- dorinae, and, like that group, have relatively large chromosomes (Duncan, 1959). Listera, Neottia. Prasophyllinae Schltr. Mansfeld placed these genera with Thelymitra, but they are distinctive in aspect, and unusual in the possession of stipes. These are generally described as 34 ANNALS OF THE MISSOURI caudicles, but are not derived from the pollinia (see Vermeulen, 1959). This group seems more nearly allied to the Diuridinae and Cryptostylidinae. Corunastylis, Goadbyella, Microtis, Prasophyllum. Pterostylidinae Pfitzer (Drakaeinae Schltr.) These Australian genera are remarkable for their sensitive, motile lip, but very closely allied to the Chloraeinae, and especially to Chiloglottis. Caleana, Drakaea, Pterostylis, Spiculaea. Rhizanthellinae Rogers Rogers placed this group near the Gastrodiinae, but the form of the lip, column and anther indicate that they are allied to Caladenia. Cryptanthemis, Rhizanthella. Spiranthinae Bentham (Corymbidinae Bentham [based on Corymbis, an ortho- graphic variant of Corymborchis], Cranichidinae Pfitzer, Physurinae Pfitzer [based on Pbysurus L. C. Rich., nom. nud.], Tropidiinae Pfitzer, Maniellinae Schltr.) We feel confident that the Spiranthinae and Cranichidinae should be merged. These groups have much the same floral structure, and we see little justification for separating them, especially if some American authors are correct in reducing Schlechter's Spiranthinae nearly to a single genus. The genera included in the Goody era alliance are somewhat distinct in habit (rooting at the nodes, rather than roots fascicled) , but agree well in floral features. Here, too, there seem to be too many genera. If future study should indicate the advisability of segregating the group as a separate subtribe, the name might be based on the familiar genus Goodyera. The Tropidia alliance seems to have the strongest claim to subtribal distinction, but no differences in floral structure have been demonstrated. In habit, these genera resemble Palmorchis, of the Sobraliinae, and Apostasia. The earliest subtribal name, Corymbidinae, should be changed in form, if it is to be used. Key Tropidia alliance not strongly plicate 2 .. ..SMra a. Goodyera alliance: Anoectoc ;~topus, Cystorchis, Dicero- stylis, Dossinia, Erythrodes (Physurus), Eucosia, t: tiylh, Goodyera, Gymnochilus, Haemaria, Herpysma, Hetaeria, Hylophila, Kublhasseltia, Lepidogyne, Macodes, Moerenboutia, My rmecbis, Odontochilus, Orchipedum, Papuaea, Platylepis, Tubilabium, Vrydagzynea, Zeuxine. b. Cranichis alliance: Altensteinia, Basken ill clla, Pontbeiva, Porpbyrostacbys, Prescottia, Pseudocent centrum, Stenoptera, Wull- schlaegelia. DRESSLER & DODSON CLASSIFICATION IN ORCHIDACEAE 35 c. Spiranthes alliance: Centrogenium, Eury styles, Lankesterella, Manniella, Pelexia, Sarcoglottis, Sauroglossum, Spiranthes. d. Tropidia alliance: Corymborchis, Tropidia. Tribe 4. ORCHIDEAE (Granulosae Blume. [in part], Ophrydeae Lindley, Epipogieae Parlatore) Here, except for the inclusion of the Epipogiinae in this tribe, we have fol- lowed essentially the classification of Bentham, which seems to be the best. There are still several problems in the morphological interpretation of flower structure in the Orchideae. Until this is better understood, it is difficult to consider the evolution or relationships of the group. As Godfery (1933) and Swamy (1949) have indicated, the Orchideae are clearly more highly specialized than the majority of Neottieae, and should not precede them in a phylogenetic scheme. Key to Subtribes Leafless 525 r'v i - erect or incun , narrowl y attached to the thetrumn" cept for Sihvrck ■«); the anthe r erect or rechnate, 2£* ~ZLi::tL: :;;. d :: ;:!::: ;jatt&z whh he the m base ; P per ] imes spurred (Old Wt divided L £533 Coryciinae Bentham (Disperidinae Schltr.) Ceratandra, Corycium, Disperis, Pterygodium. Disinae Bentham (Satyriinae Pfitzer) is a single subtribe. The position of the elongate (Old World)' Satynum alliance a. Disa alliance: Brownleea, Disa, Schizodium. b. Satyrium allianci \ilvorchis} Epipogiinae Schltr. Recent authors have placed these genera in the Neottieae, but the persistent anther and the sectile pollinia with basal caudicles indicate a much closer affinity with the Orchideae. Godfery (1933) followed Parlatore (1858) in treating 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN Epipogium as the type of a separate tribe. In Epipogium aphyllum the anther is incumbent and the pollinia are parallel with the caudicles, which attach to a viscidium near the apices of the pollinia. It is not clear that the incumbent anther indicates a relationship with the primitive Epidendreae; the peculiar orientation of the caudicles suggests that it is a derived condition in this species. Rohrbach (1866) gives detailed illustrations of E. aphyllum. The anther is erect in the autogamous E. roseum (as it is in the young bud of E. aphyllum) and the caudicles are apparently functionless. The detailed structure of Stereosandra is not well known, but the anther is erect and the pollinia bear caudicles. These genera probably are not very closely related to the Gastrodiinae, though, being sapro- phytes, they are superficially similar. Epipogium, Stereosandra. Orchidinae (Angiadeniae Parlatore, Habenariinae Bentham, Serapiadinae Ben- tham, Ophrydinae Bentham & Hooker, Gymnadeniinae Pfitzer, Androcory- dinae Schltr., Huttonaeinae Schltr., Platantherinae Schltr.) The union of the Platantherinae and Habenariinae of Schlechter is certainly appropriate, when there is yet disagreement as to whether or not Habenaria and Vlatanthera are distinct genera. This group is seriously in need of monographic attention. The extreme "splitting" of European workers combined with the (equally unrealistic) extreme "lumping" of recent American workers has led to Aceratorchis, Aceras, Acrostylia, Amitostigma, Anacamptis, Androcorys, Arnottia, Bartholina, Benthamia, Bicornella, Bonatea, Brachycorythis, Centrostigma, Chamaeorchis, Coeloglossum, Cynorchis, Dactylorchis, Deroemera, Diphylax, Diplacorcbis, Diplomeris, Dithrix, Galea; i icnia, Gymnadenia, Habenaria. \\ vmimnm, Himanto glossum, Holothrix, Huttonaea, Leucorchis, Loroglossum, Neobolusia, Neotinea, Neotti- anthe, Nigritella, Ophrys, Orchis, P, v, Platanthera, Platycoryne, Ro .us, Sch-wartzkopffia, Serapias, Traunsteinera, Tylostigma. Tribe 5. EPIDENDREAE Lindley (Cereaceae Blume, Arethuseae Lindley, Gastro- dieae Lindley, Malaxideae Lindley, Vandeae Lindley, Kerosphaerae Schltr., all but 4 of Pfitzer's 31 tribes [1887], Sturmieae Pfitzer [1897]). inflorescence axillary s LndXlI/L a!lv ,r„ id; plants autophytes < =tSM ,S ' " W ' Vi ' 0; flow, r v;r ODI ^ F nd grass-like (No rth Am. n,, and >ase, deciduous (Amen ica and Africa).... rticulated, pa 6 DRESSLER & DODSON CLASSIFICATION IN ORCHID ACEAE 37 t7 [ pLrotTJlr l hl "' m a kw ^ " ° ;•/,.,/■;, 'rum, Den drobium, Coelo g yne Inflorescence lateral, usually basal Leaves plicate 9 mre^hooTs^fALr 1 ' K U,,KSS lll " >,S *'"'' '' tCn,U " rh tllC im '" ,UU ' """ The pollinia naked, without caudicles (seealsc , PsenJern ,, Dcndrob iinae) (widespread). The pollinia bearing caudicles ....... ....... ,1 Pollinia 8 12 Not with the above combina, The ovary articulated to the ANNALS OF THE MISSOURI BOTANICAL GARDEN ulbs usually of a single internode (rarely absent in the Zygopetatum alliance). ). Pollinia 4 or (usually) 8, clavate or laterally flattened, withe ut stipe and "^^^ Poll.ma usually with a stipe 2 7). Pollinia 2 or 4; plants usually with corms, sometimes with elongate pseudobulbs and basal inflorescence; lip often hinged or spurred (widespread) Cyrtopodiina Pollinia pseudob utb; P lip tS of!en h s^T bTtd^ h^^Tl^ ually lateral on the red; anther usually 5). Lipimn obile, often with antenna-like processes; flowers usually racemose (America). solitary lly hinged at base and mobile, without antenna-like pr cesse= flowers often 2-8 (usually 4). clavate or laterally flattened, often without caudicle or Pollinia 2, or 4 and superposed, usually with stipe and viscidii 3 3). Pollinia naked, without caudicles, usually without a viscidium never with stipe or Pollinia not naked, usually with distinct viscidium and stipe or caudicles. 3 2 or 4; inflorescence various (widespread, predominant! Pollinia 8; inflorescence paniculate (New Guinea) RlDLEYELLINA Tot "slighTor Tbslt^nT tr^ti e 'Z^^ freqUer Column base of column more or less united into a tubular strv Lip free from column, not forming a tubuhr structure (Old Wor ld) Genyorchidina ). Pollinia 4, the caudicles inconspicuous; roots extremely thick (Ceylon and South Pollinia 8, with a long caudicle; roots not especially thick (A ; a) Thelasiina D). Pseudob Cymbidiina 6) ' withT . Clinand differ^ted'^rom^thf st^e C °| Um e n ' c imm ° bile; visdla\, \' cogardneria, Otostylis, Paradisianthus, Pescatoria, Promenaea, Stenia, Wareella, Zygopetalum. Oncidiinae Bentham (Notyliinae Bentham, Adinae Pfitzer, Aspasiinae Pfitzer, Ionopsidinae Pfitzer, Odontoglossinae Pfitzer, Trichopiliinae Pfitzer, Brachti- inae Schltr., Campanemiinae Schltr., Cochliodinae Schltr., Comparettiinae Schltr., Dichaeinae Schltr., Lockhartiinae Schltr., Macradeniinae [Mansf .] Schltr., Ornithocephalinae Schltr., Papperitziinae Schltr., Pterostemminae Schltr., Saun- dersiinae Schltr., Telipogoninae Schltr., Trichocentrinae Schltr.) It is in this group that we have committed the most wholesale reduction of subtribes. These changes, however, are required not only by the rapidly accumu- lating evidence of intergeneric fertility (see Moir, 1959), but by the patterns of morphological variation as well. Some of the "genera" placed in separate sub- tribes by Schlechter can not be distinguished by any feature known to us. ANNALS OF THE MISSOURI BOTANICAL GARDEN a. DlCHAEA This genus is distinctive in habit cephalus alliance. The pilose colum: found in Telipogon and allied genera. b. Oncidium alliance iification, that there would ances, those with spurs and those without. The presence or absence of a spur, however, is not consistent within the genera, and these groups are nearly all interfertile. We have successfully crossed Comparettia with Oncidium, Trichocentrum, Trichopilia and Ionopsis. Rod- riguezia, similarly, is interfertile with a wide range of genera. Trichopilia is super- ficially distinct, in that the base of the lip enfolds the column, but this same feature is found in some species of Miltonia. Notylia is unusual in the dorsal position of the anther, but this, too, is inconsistent. Lockhartia is distinctive in habit, but has no other consistent feature to distinguish it from the other genera of this group. Pterostemma is poorly known, but its monopodial habit can not be taken to exclude it from this subtribe. Ada, Amparoa, Aspasia, Brachtia, Brassia, Capanemia, Caucaea, Chaenanthe, Coch- lioda, Cohniella, Comparettia, Diadenium, Erycina, Gomesa, Hybochilus, Ionopsis, Leochilus, Lockhartia, Macradenia, Mesospinidium, Miltonia, Neodryas, Neokoehleria, Notylia, Odontoglossum, Oncidium, Papperitzia, Petalocentrum, Polyotidium, Plectrophora, Pterostemma, Quekettia, Rodriguezia, Rodrigueziopsis, Roezliella, Rusbyella, Sanderella, Saundersia, Scelochilus, Sigmatostalix, Solenidium, Systeloglossum, Sutrina, Theodorea, Trichocentrum, Trichopilia, Trizeuxis, Warmingia. c. Ornithocephalus alliance These genera are distinguished from the Oncidium alliance only by the number of pollinia, but they do form a somewhat distinctive group. There is no way, however, to separate the Telipogoninae and the Ornithocephalinae. Many of the Telipogoninae are distinctive in that they apparently mimic insects (as in many species of Ophrys), and in the pilose column and hooked viscidium, but these latter features are matched by Cordanthera and some species of Dipteranthus. Centroglossa, Chytroglossa, Cordanthera, Cryptarrhena, Dipteranthus, Dipterostele, Ornithocephalus, Phymatidium, Platyrhiza, •eros,Zygostates. Pachyphyllinae Pfitzer Though distinguished by the differently shaped column and consistently mono- podial habit, these genera seem closely allied to the Maxillariinae. Pachyphyllum DRESSLER & DODSON CLASSIFICATION IN ORCHIDACEAE 47 and Chrysocycnis, especially, appear to be closely allied, and resemble each other in habit, inflorescence and 3 -winged ovary. Centropetalum, Pachyphyllum. Pleurothallidinae Bentham glossum, Restrepia, Scaphosepalum, Stelis, Yolanda. Pogoniinae Pfitzer (Nerviliinae Schltr.) We follow Mansfeld in separating the Vanillinae and the Pogoniinae, and in- cluding Nervilia in the Pogoniinae. The flower structure of Nervilia is quite comparable to that of the other Pogoniinae, and most species were first described under Pogonia. The habit is somewhat distinctive. These genera are similar in floral structure to both the Vanillinae and the Sobraliinae. The pollen is quite mealy, but they lack the distinctive seed structure of the Vanillinae. Cleistes, Isotria, Lecat:- - \cnilhi, Pogonia, Pogoniopsis, Psilo- chilus, Triphora. RlDLEYELLINAE Schltr. The habit of this genus is suggestive of Bulbophyllum, and the poll to be naked, though the floral structure is otherwise r Thelasiinae. Ridleyella. Sarcanthinae Bentham (Aeridinae Pfitzer) This is the great group of primarily Old World monopodial orchids. In floral specialization and complexity they parallel the American Oncidiinae and are not easily "keyed" from the monopodial Oncidiinae, though there is probably no close relationship. Note that this group must be known as the Vandinae if the tribe Vandeae is maintained. Hawkes and Heller (1959) list the "subtribe Campylo- centrinae." This name is listed by Hoehne in Flora Brasilica (12 1 :23,39), but we cannot find that it has ever been formally proposed, nor can we find any reason that it should be. Abdominea, Acampe, Adenoncos, Aerangis, Aerantbes, Aerides, Ambrella, Ancistror- rhyncbus, Angraecopsis, Angraecum, Ankylocheilos, Arachnis, Armodorum, Ascocenlrum, Ascochilopsis, Ascoglossum, Barombia, Bathiea, Beclardia, Bogoria, Bolusiella, Bonniera, Crossangis, Cryptopus, Cyrtorchis, Dendrophylax, Diaphanantbe, Dmklageella, Dipfo- centrum, Diploprora, Dryadorchis, Eggelingia, I <\ Eurychone, Finetia, FUxgen , Holcoglossum, Hymenorchis, Jumellea, Lemurella, ,, Microsaccus, Microtatorcbis, Mystacidium, Nephrangis, Neobathiea, Oconia, Oeoniella, Omoea, Ornitho- chilus. PelatdK , I'hormangis, Phragmorchh, Plectrelminthes, Podangis, Polyrbiza, Pomatocalpa, Porphyrodesme, Rangaeris, Renantbera, 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN Kenantherella, Rbipidoglossum, Khynchostylis, Robiquetia, Saccolabiopsis, Saccolabium, Sun ant!;:.-, S us, Schoenorchis, Solenangis, Spbyrarhynchus, Stauro- chilus, Stauropsis, Taeniophyllum, T* mum, Triceratorhynchus, Trichoglottis, Tridactyle, Uncifera, Vanda, Vandopsis, Ypsilopus. Sobraliinae Schltr. (Palmorchidinae Schweinf. & Correll [not validly published]) These genera were included in the Neottieae by Schlechter, but the pollinia are more or less waxy in many species, and the flower structure agrees well with that of the Epidendrinae and Thuniinae. Mansfeld actually assigns the Thuniinae to this subtribe, though we believe this action to be questionable, at least without more detailed knowledge of the Thuniinae. Schweinfurth has evidently abandoned the use of the subtribe Palmorchidinae, which was proposed without description. In no case did Schweinfurth and Correll assign Corymborchis to the Palmor- chidinae, an action which would scarcely have been defensible (see, however, Hawkes & Heller, 1959). Dicer at ostele, Elleantbus, Palmorchis, Sertifera, Sobralia, Xer orchis. Stanhopeinae Bentham (Gongorinae Schltr.) This group is not clearly distinguished from the Maxillariinae {Ly caste alli- ance), though the more bizarre genera {Stanbopea, Coryantbes, etc.) are easily distinguished. More study of this group is needed. Acineta, Chaubardia}, Cirrbaea, Coeliopsis, Coryantbes, Endresiella, Gongora, Gorgo- glossum, Houlu ; ..i, Lueddemannia, Lycomormium, Paphinia, Peristeria, Polycycnis, Scblimia, Sievekingia, Stanbopea, Trevoria. Thecostelinae Schltr. The flower structure of Thecostele is quite bizarre, but adequate knowledge of all species may indicate that it should be included in the Genyorchidinae. Thelasiinae Schltr. The Thelasiinae are small Asiatic plants which are distinctive in habit and inflorescence. The pollinia have been described as having a stipe, but this appears to be an unusually long caudicle (Mansfeld, 1937b). Chitonanthera, Octarrhena, Oxyanthera, Phreatia, Rbyncbopbreatia, Tbelasis. Thuniinae Schltr. (Claderiinae Mansf. [not validly published] ) Tall Asiatic plants with slender stems and rather showy flowers, in habit these plants (especially Arundina) closely resemble the Epidendrinae. In the structure of the column, however, they more closely resemble Coelogyne and Pbajus. Brom- headia was placed in the Polystachyinae by Schlechter, but is surely out of place there (see Ridley, 1891). It resembles Dilochia in habit and appears to resemble Claderia in flower structure. Unfortunately, we have seen too little of all these genera. The group is unusually diverse in the structure of the pollinia, and it is possible that further study will show it to be unnatural. The subtribe Claderiinae was proposed without description. Arundina, Bromheadia}, Claderia, Dilochia, Thunia. DRESSLER & DODSON PHYLOGENY IN ORCHIDACEAE 49 Vanillinae Bentham These genera are, in several features, among the most primitive in the sub- family. They are included in the Pogoniinae by Schlechter, but the habit and distinctive seed structure would seem to justify their separation, as indicated by Mansfeld. They also show some affinity to the Sobraliinae, but there is little evidence of close alliance with the Neottieae. Duckeella, Epistephium, Eriaxis, Galeola, Vanilla. SUBTRIBES OF UNCERTAIN AFFINITY Grobyinae Schltr. This genus has been placed near the Cymbidiinae in most systems, but this seems to be questionable. We have not seen adequate material, but a comparison with the Maxillariinae may be in order. Grobya. Pachyplectrinae Schltr. This genus is known only from New Caledonia. It apparently belongs to the Neottieae, but Schlechter's description is not adequate for more exact placement. It may be allied to the Diuridinae and Cryptostylidinae. Pachyplectron. Phylogeny There are two extremes in the variation patterns shown by living plant groups, depending apparently on rates of evolution and amount of extinction. At one end of the spectrum stand such families as the Magnoliaceae, Annonaceae and Nym- phaeaceae. These families have the appearance of being "old" groups, in which evolution is proceeding at a leisurely pace and extinction has greatly affected the pattern of variation. Such groups as the cacti and the Euphorbiaceae-Euphorbieae represent the other extreme. They show little evidence of great age, show signs of relatively rapid diversification in geologically recent time, and give much less evidence of extinction. Our classification of genera and higher groups is in large measure based on extinction, and so we have very different problems with these two types. The delimitation of genera and tribes within the Magnoliaceae or the Nymphaeaceae is not difficult. We are often at a loss, though, to understand their relationships or phylogeny. As should now be clear, the orchids are near the other end of our spectrum in these features. Genera are often difficult to define, and higher categories within the family seem even worse. These hazy boundaries between tribes and subtribes, however, may give clues to the patterns of phylogeny within the family. We do not mean to imply that living groups can often be derived from other living groups, but one can find excellent evolutionary series for nearly every morpho- logical feature within the orchids. In recent years there has been a healthy skepticism concerning phylogenetic schemes (see especially Sporne, 1959). Even 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN when one can find a good morphological series in some feature, it is often difficult to decide in which direction or directions evolution has occurred. This problem is not so serious in a highly derived group such as the Orchidaceae. In comparing Cephalanthera and Oncidium there can be little doubt as to which is derived and which is primitive. In nearly every feature in which these two differ, it is Cephalanthera which is the ordinary monocot, easily comparable with other mono- cot families, 9 carcely be understood without comparing it with the less specialized members of the family. One would scarcely expect to derive whorls of separate stamens and pistils from the column of the advanced Epiden- dreae, but the evolution of the column from primitively free parts is easily under- stood and partially documented among the living orchids. As an actively evolving group in which the patterns of evolution are unusually clear, the orchids are especially appropriate for evolutionary study and may throw a good deal of light on parallelism, polyphylesis and other problems which plague the biologist dealing with apparently more ancient groups. When one studies the relationships of the subtribes, a pattern takes shape, and one may form a rather clear idea of the over- all evolutionary patterns for the family. The pattern of relationships for the Epidendreae (fig. 1) scarcely provides a classic dendrogram, but there are clear indications of primitive groups and some indications of the patterns of evolution which may have occurred. These may best be considered by discussing evolution in particular features of the plant. As Holttum (1955) has shown, the predominant growth form in a wide range of monocots is the sympodium, and we may reasonably consider this to be the primitive condition in the orchids. The majority of primitive orchids have a rather ordinary monocot habit, as for example Cephalanthera, with a short rhizome and erect, non-thickened annual stems with scattered, spiral leaves and a terminal inflorescence. The origin of the majority of other sympodial growth forms by phyletic shortening and/or thickening of the stems is readily envisioned. Of special interest is the monopodial habit, in which the stem has unlimited apical growth and roots are not restricted to the basal portion. In some systems this has been considered to be the distinguishing mark of the Sarcanthinae. We find, however, that the monopodial habit occurs in the Vanillinae, Cymbidiinae, Maxil- lariinae, Pachyphyllinae, Cryptocentrinae, Thelasiinae and Oncidiinae, and possibly in other groups as well. In some of these groups we have a good graded series from sympodial plants to related monopodial types. Not only does the monopodial habit appear to have evolved independently in many groups, but its evolution seems to have followed somewhat different patterns in different cases. In the Vanillinae and the Sarcanthinae it has apparently been the simple retention of apical growth in the members of a sympodium (with lateral inflorescence). In some of the Maxillariinae and allied groups it appears to be the suppression of pseudobulbs on a leafy rhizome. In other cases, such as Maxillaria valenzuelana, and some Oncidiinae {Oncidium pusillum complex, Rodriguezia spp.), the monopodial habit :i LLARI INAE DRESSLER & DODSON PHYLOGENY IN ORCHIDACEAE 51 THECOSTELINAE \ THELASI INAE •E.TCN.DINU J plEUR0THALL|D|NA[ DHMoainuE ! / mmnmn shmmim <«!°"»" ' EpTdEMDR IIJir^-__ ~J \ ' PACHTPHYLLIN4E LIPMIDIUE . inJV. \ „--' \ J,, ; „*" ""^BLETIINAE CYRTOPODI INAE SOBRALMNAE / N. / \ ** ARETHUSINAE ^COLLABI INAE\ seems to have evolved by the retention of a permanent juvenile form. In hybrid swarms involving Rodriguezia refracta (sympodial) and a related monopodial species one may find a complete series ranging from those plants which are mono- podial only in the seedling stage to those which are monopodial for the life of the plant. Kranzlin (1923) attempted to retain the monopodial habit as the distinguishing feature of the Sarcanthinae by classing Dichaea, Lockbartia, Pterostemma and the Pachyphyllinae as the heterogeneous "Pseudomonopodiales," pointing out that Lockbartia and some species of Dichaea are not strictly monopodial, in that the stems are erect, and of more or less limited growth, with roots and branches arising only at the base. While this is true, a very strict definition of the monopodial habit would also disqualify some Sarcanthinae (ex. Mystacidium distichum), and still leaves some undoubted monopodia in Om tnd species of Dichaea. Though of considerable morphological and evolutionary interest, the monopodial habit has limited value as a taxonomic criterion. Saprophytic orchids are found in all three tribes of the Orchidoideae, and in at least twelve different subtribes. The majority of orchids pass through a sapro- phytic seedling stage, which may last for months, especially in terrestrial species. Thus, the evolution of a completely saprophytic life cycle in many different groups of orchids is not surprising. The wholly saprophytic orchids pose special taxonomic problems. The adaptations for saprophytism drastically change the vegetative features of the plant, thus obscuring some of the characteristics normally used in determining relationships. It may be that even the reproductive features are affected by these adaptations. In Corallorhiza, for example, we find the polliniar 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN apparatus to be simpler than in the related Oreorchis and Tipularia. In the absence of these closely allied genera, its relationship to the Cyrtopodiinae would be much less clear. The saprophytes are difficult to cultivate and poorly represented by herbarium specimens, which further complicates their study. Autogamy is fre- quent, and an autogamous saprophyte is nearly the ultimate in taxonomic difficulty. A special key to the saprophytic genera of orchids is needed. PSEUDOBULBS AND CORMS A great many orchids, and especially the epiphytic groups, show variously thickened stems or "pseudobulbs." While these structures are quite diverse in form, they fall into a limited number of morphological types and seem to show some evolutionary trends. One of these seeming trends is from pseudobulbs (or corms) of several or many inter nodes to pseudobulbs of a single internode (as in Bulbophyllum, Maxillariinae and Oncidiinae). Reference to the chart of relation- ships (fig. 1) suggests that there may be two basic patterns for the origin of pseudobulbs. In the majority of the Bletiinae, Cyrtopodiinae and related groups (the right side of fig. 1), pseudobulbs seem to have been derived phyletically from more or less corm-like structures, as in Bletia and Phajus. These thickened stem- bases may be found in either terrestrial or epiphytic groups, while the pseudobulbs of a single internode are restricted to primarily epiphytic groups. The other main pattern for the derivation of pseudobulbs appears to be shown by the Epidendrinae and some related groups (left side of fig. 1). In these groups corm-like structures are not found, and the pseudobulbs appear to have evolved by a phyletic thickening of the entire aerial shoot (as in Dendrobium, Barkeria, etc.). The more derived members of these groups may also possess pseudobulbs of a single internode, which are morphologically indistinguishable from those of the Oncidiinae or Maxillariinae (ex. Bulbophyllum, some species of Dendrobium). The evolutionary trends in leaf type seem rather clear, and have already been outlined by Rolfe (1909-1912). The primitive type of orchid leaf is probably non-articulate, wide (more or less elliptic) and plicate, of convolute vernation. The trends toward an articulate leaf which is narrow (more or less ligulate) and conduplicate, have probably occurred independently in several phyletic lines, and appear to be strongly correlated with the epiphytic habit. It is interesting that the monopodial orchids all have strictly conduplicate leaves, with the partial exception of Vanilla, which has convolute vernation, but the fleshy leaves of Vanilla are by no means plicate, and the mature leaf usually appears conduplicate. This points out the imperfect correlation between conduplicate leaves and duplicate vernation. The correlation is generally good, however. We have used plicate and conduplicate in the present paper, as terms descriptive of the mature leaf and therefore more readily determined. There are, of course, transitional stages, such as the many- veined but conduplicate leaves of most Cymbidiinae, which appear superficially It is probable that the primitive orchid flower was a relatively unspecialized monocot flower with equal and similar perianth parts, similar to that now found in the Apostasieae. Such a hypothetical primitive orchid flower is sketched in fig. 2. The only unusual feature of the simpler orchid flowers is a certain degree of zygomorphy in the androecium. The three anthers which are found in the orchids are all on the abaxial side of the flower and are members of two different whorls. The adaxial stamens were evidently lost at an early stage in orchid evolu- tion. In some groups of ancestral orchids, as in the living Cephalanthera, the viscid matter of the relatively unspecialized stigma may have served to glue the pollen masses of the median anther to the pollinating agent on its withdrawal from the flower. Such a relationship was surely the basis for the evolution of those tribes in which only the median anther is functional. A tremendous diversity exists in the form of the column and anther among living orchids. So great is this diversity in form and position that it is very difficult to use a precise termi- nology for the anther and its parts. The column itself has clear dorsal, ventral and lateral aspects (though morphologically confused by resupination, in which the adaxial side of the flower becomes ventral). 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN R D ^ # c F Fig Diaqr minat c longitudin 1 section of orchid co Pollen is •..eWvilid/p. • i. 'The • . to the base o :," : ^ Sp!r.i , M stipules fully amplex occasionally in Brosimum). scarcely longer than broad, h. Unarmed trees, pistillate heads :ads more or less conduplicate-flabellate, c at the base and perigynous; leaves =omimngl«^wh (155) ANNALS OF THE MISSOURI BOTANICAL GARDEN slightly fleshy drupe The genus Ogcodeia occurs in Costa Rica and may yet be found in Panama. It is related to Perebea and Helicostylis, but is distinguished by the burr-like pistillate inflorescences and fruits; the heads are involucrate as in Perebea but the flowers epigynous and reduced in number as in Helicostylis. The East Indian and Micronesian genus Artocarpus is represented in cultivation by the breadfruit (A. communis Forst.) and the jackfruit (A. heterophyllus Lam.). Both are large and handsome shade trees and are monoecious, the staminate flowers in stout cylindrical spikes and the pistillate in great spherical or cylindrical heads somewhat reminiscent of the northern Osage orange [Madura pomifera (Raf.) Schneid.] on a larger scale. The jackfruit has undivided leaves; breadfruit leaves are leathery, deep glossy green and incised like those of some monstrous oak. Breadfruit, which was the principal cargo of the ill-fated H.M.S. "Bounty" in 1789, still is an important article of food for many people in Panama, both for the starchy pulp of the fruit and for the large nut-like seeds of some varieties (breadnuts) ; by the Spanish-speaking people it is known as drbol de pan and fruta de pan. Jackfruits are edible but of inferior quality. 1. CHLOROPHORA Gaud. Chlorophora Gaud. Bot. Voy. Freycinet 509. 1830. Dioecious trees, the branches occasionally with stout axillary spines. Leaves alternate, distichous; stipules lateral and about half-amplexicaul. Inflorescences axillary, the staminate spicate, the pistillate capitate; flowers associated with spatulate glandular-carinate bracts. Staminate flowers with 4 essentially free tepals; stamens 4, the filaments rather elongate, involute before anthesis, the anthers 2 -celled, extrorse. Pistillate flowers with 4 basally coherent tepals, hypogynous, the pistil with a simple filiform lateral stigma. Fruit a loosely coherent slightly fleshy syncarp. Two species — the following and Ch. excelsa (Welw.) Benth. & Hook. f. of western tropical Africa. (156) flora of Panama (Moraceae) 117 OF THE MISSOURI BOTANICAL GARDEN 1. Chlorophora tinctoria (L.) Gaud. Bot. Voy. Freycinet 508. 18 30. Morus tinctoria L. Sp. PI. 986. 1753. Morus xanthoxylon L. Syst. Nat. ed. 10. 2:1267. 1759. Broussonetia tinctoria (L.) HBK. Nov. Gen. & Sp. 2:32. 1817. Broussonetia plumerii Spreng. Syst. Veg. 3:901. 1826. Morus tataiba Veil. Fl. Flum. 10: t. 21. 1827. Madura tinctoria (L.) D. Don, ex Steud. Nomencl. ed. 2. 2:87. 1841. Madura plumiera D. Don, ex Steud. loc. cit. 1841. Morus plumiera (D. Don) Burm. ex Steud. loc. cit. 1841. Broussonetia xanthoxylon (L.) Mart, in Flora 24: 2 Beibl. 10. 1841. Broussonetia / it. 1841. Madura sempervirens Ten. Cat. Hort. Bot. Nap. 87. 1845. Madura xanthoxylon (L.) Endl. Gen. Pi. Suppl. 4 2 :34. 1847. Madura chlorocarpa Liebm. in Danske Vidensk. Selsk. Skr. 5 ser. 2:314. 1851. Madura Polyneura Miq. in Mart. Fl. Bras. 4 1 :154. 18 53. Madura affinis Miq. loc. cit. 155. 18 53. Madura subintegerrima Miq. loc. cit. 157. 1853. Madura velutina Bl. Mus. Bot. Lugd.-Bat. 2:82. 1856. Trees, reputedly to 30 m. in height. Leafy twigs rather slender, somewhat flexuose, glabrous, developing a yellowish gray bark. Leaves alternate, distichous, petiolate, the blade quite variable, broadly oval to ovate, obovate or oblong, the tip usually subcaudate-acuminate, the base usually strongly inequilateral and more or less cordate, closely serrate to entire, 3-12 cm. long, 2-6 cm. broad, firmly membranaceous, glabrous to minutely puberulent above and below, the petiole 5- 10 mm. long, slender; stipules lateral, leaving a scar about half-surrounding the stem or less, narrowly lanceolate, 2-10 mm. long. Staminate spikes slender, 3-10 cm. long, bearing innumerable densely congested flowers interspersed with minute puberulent spatulate glandular-carinate bracteoles about 6 mm. long: tepals 4, broadly oval, minutely puberulent, about 3 mm. long; stamens 4, the filaments about 5 mm. long, strongly involute before anthesis, the anthers broadly oval, about 1.5 mm. long; pistillode about 1 mm. long. Pistillate heads globose, about 1 cm. in diameter, very shortly pedunculate, the flowers very densely congested and interspersed with spatulate glandular-carinate bracteoles about 6 mm. long: tepals 4, narrowly spatulate, about 6 mm. long, minutely puberulent; ovary superior, obovoid, about 4 mm. long, the lateral filiform densely papillate stigma about 10 mm. long. Fruit a very slightly fleshy, globose aggregate to about 2 cm. in diameter. Southern Mexico to Argentina; Antilles, at low elevations. Mora or morillo and macano; fustic. The wood is extremely durable and resistant to insects and decay according to Allen (Allen, P. H. The Rainforests of Golfo Dulce. Gaines- ville, Fla. 1956). canal zone: between France Field and Catival, Standley 30301; Balboa Heights, Killip 3000; Darien Sta., Standley 31600. chiriqui: Puerto Armuelles, Woodson & Schery 809, Stern & Chambers 136; Progreso, Cooper & Slater 237. darien: Garachine, Pittier 5605. Panama: Las Sabanas, Zetek 3684; Panama, Standley 26833; Rio Tapia, Standley 2820O; Rio Tecumen, Standley 2938 1; Chepo, Pittier 4710. flora of Panama (Moraceae) 119 2. TROPHIS P. Br. Trophis P. Br. Hist. Jam. 357. 1756; L. Syst. Nat. ed. 10. 1289. 1759, nom. Bucepbalon L. Sp. PI. 1190. 1753, nom. rejic. Skutchia Pax & Hoffm. ex Morton, in Journ. Wash. Acad. Sci. 27:306. 1937. Dioecious trees and shrubs, the branches unarmed. Leaves alternate, distichous; stipules lateral. Both staminate and pistillate inflorescences secund racemes and spikes, the small congested or relatively distant flowers interspersed with small, shortly stipitate, peltate bracts. Staminate flowers: tepals 4, united or essentially free to the base, with an obvious pistillode; stamens 4, the filaments much longer than the tepals and strongly inflexed before anthesis, later sharply reflexed, the anthers broadly oval, introrse. Pistillate flowers epigynous; tepals 4, minute; style central and deeply 2-lobed. Fruit a small fleshy 1 -seeded drupe. About 6 species ranging from southern Mexico to the Amazon basin, and in the Antilles. Trophis chorizantha Standi., with less congested staminate spikes and racemose pistillate inflorescences is to be expected in western Panama. 1. Trophis racemosa (L.) Urb. Symb. Ant. 4:195. 1905. Bucepbalon racemosum L. Sp. PI. 1190. 1753. Trophis americana L. Syst. Nat. ed. 10. 1289. 1759. Trophis americana /3 ramon Bur. et y meridionalis Bur. in DC. Prodr. 17:253. 1873. Sahagunia urophylla Donn. Sm. in Bot Gaz. 40:11. 1905. Trees to about 15 m. in height. Leafy twigs rather slender, somewhat flexuose, glabrous, developing a yellowish gray bark. Leaves alternate, distichous, petiolate, the blade broadly oval to elliptic-oblong, occasionally irregularly pandurate, the tip subcaudate-acuminate, the base essentially equilateral and rounded to broadly obtuse, entire or somewhat serrate-undulate toward the tip, 6-20 cm. long, 2-9 cm. broad, firmly membranaceous, glabrous, the petiole 5-7 mm. long; stipules lateral, narrowly lanceolate, about 5 mm. long. Staminate spikes slender, 3-5 cm. long, bearing innumerable densely congested flowers interspersed with minute shortly stipitate peltate bracts: tepals 4, somewhat united at the base, broadly oval, about 2 mm. long, minutely papillate; stamens 4, the filaments about 3 mm. long, the anthers about 0.5 mm. long. Pistillate spikes rather short and strongly secund, 1.0-1.5 cm. long, densely and minutely ferruginous-puberulent, bearing rather few and distant sessile truncate conic flowers about 2 mm. long: perianth lobes minutely trigonal; stigma lobes about 1.5 mm. long. Fruits globose or ovoid, Southern Mexico to the Amazon basin. Ojoche macho, lechosa, ramon, gallote, morillo; breadnut. bocas del toro: Changuinola Valley, Cooper & Slater, 4, 4", Seibert 1 58 1, 1583; Almirante, Cooper 349; Old Bank Island, Von Wedel 2075-, Water Valley, Von Wedel ■ r 5pO; Rio Cricamola between Finca St. Louis and Konkintoe, Woodson, Allen d Seibert 1^24. canal zone: Barro Colorado Island, Standley 41080, 41082; Culebra, P 3627; between France Field and Catival, Standley 30296, 30235; Gamboa, Pittier 6636, 6637, 6652; Rio Pedro Miguel, near East Paraiso, Standley 29063; Gatun, Standley 27287; Fort Sherman, Standley 30970. chiriqui: San Bartolome, Woodson & Schery 866. (159) 120 ANNALS OF THE MISSOURI BOTANICAL GARDEN flora of Panama (Moraceae) 121 3. SOROCEA A. St.-Hil. By WILLIAM C. BURGER Sorocea A. St.-Hil. in Mem. Mus. Paris 7:473. 1821. Pseudosorocea Baill. in Adansonia 11:296. 1875. Balanostreblus Kurz, emend. Hutchinson, in Kew Bull. 1918:152. 1918. Dioecious trees, the branches unarmed. Leaves alternate, distichous; stipules lateral. Both staminate and pistillate inflorescences racemes and spikes; the small, congested or relatively distant flowers interspersed with small, shortly stipitate, peltate bracts. Staminate flowers: tepals 4, united or essentially free to the base, imbricate in bud, without an obvious pistillode; stamens 4, the filaments about as long as the tepals and scarcely inflexed before anthesis, the anthers broadly oval, extrorse. Pistillate flowers hypogynous to perigynous: tepals 4, completely united or minutely 4-lobed; ovary superior to inferior, the style central and deeply bifid. Fruit a small 1 -seeded drupe. About a dozen species ranging from Costa Rica to Argentina, a. Leaves glabrous beneath, narr ..bovate, rarely 1. Sorocea affinis Hemsl. Biol. Centr.-Amer. Bot. 3:150. 1883. Trees to about 15 m. Leafy twigs rather slender, slightly flexuose, glabrous, developing a yellowish gray bark. Leaves alternate, distichous, petiolate, the blades narrowly elliptic to elliptic -obovate, abruptly subcaudate-acuminate, the base equilateral and acutely cuneate, more or less serrate-undulate toward the tip, 6-20 cm. long, 2—8 cm. broad, firmly membranaceous, glabrous, the petiole about 2-10 mm. long; stipules narrowly lanceolate, about 5 mm. long. Staminate racemes 2-8 cm. long, bearing numerous rather closely spaced flowers interspersed with minute peltate bracts, minutely puberulent-papillate: tepals 4, somewhat united at the base, broadly oval, about 2 mm. long. Pistillate raceme rather short, 1-3 cm. long, elongating in fruit, minutely puberulent-papillate, bearing 8 or more shortly pedicellate flowers about 2 mm. long, the perianth thickened above the middle of the flower. Drupes subglobose, about 8 mm. broad, green with orange tip or bright red when fully mature. Presently known only from lowland forests of Panama but probably extending south into Colombia. bocas del toro: vicinity of Chiriqui Lagoon, Von Wedel 1103, 1390; Water Valley, Von Wedel 598A, 961, 1715; Almirante, Cooper 554. canal zone: Quebrada Melgada, Steyermark 1 7494; Gatun, Johnston 1582, 1693; Gamboa, Standley 28 41 2; Las Cruces, Seibert 576; Barro Colorado Island, Aviles 26, 52, 84, Seibert 566, Bailey 2? Bailey 538, Starry 314, Woodworth & Vestal 323, Bangham 558, 479> Salvoza 966, Standley 4<>797> :. chiriqui: El Boquete, Pittier 3045. cocle: El Valle, Seibert 464. colon: Fato (Nombre de Dios), Pittier 3836; Paler, 4123. darien: Ensenada Guayabo, Stern & Chambers 180. Panama: Arraijan, Wood- son, Allen 8 Seibert 1383; Taboga Island, Hayes 658. (161) ANNALS OF THE MISSOURI Sorocea pubivena Hemsl. Biol. Centr.- Clarisia mollis S Trees to about 20 m. Leafy twigs rather thick (2-4 mm.), short-pubescent becoming glabrescent. Leaves alternate, distichous, petiolate, the blade broadly elliptic to elliptic-oblong, caudate-acuminate, the base equilateral and obtusely cuneate, entire to serrate-undulate, 10-30 cm. long, 5-10 cm. broad, firmly mem- branaceous, shortly pubescent beneath, the petiole about 8-20 mm. long; stipules narrowly acute, 6-8 mm. long. Staminate spikes 5-12 cm. long bearing numerous sessile flowers interspersed with minute peltate bracts; tepals 4, somewhat united at the base, about 2 mm. long, filaments broad and slightly connate at their base. Pistillate racemes short, elongating in fruit to 14 cm. bearing numerous flowers about 2 mm. long, the perianth thickened above, adnate to the ovary for about half its length. Drupes subglobose to ovoid, about 10 mm. broad. Costa Rica to Panama. bocas del toro: vicinity of Chiriqui Lagoon, Von Wedel logo (Type of Clarisia II.); Almirante, Cooper & Slater 28. chiriqui': Progreso, Cooper & Slater 174; vicinity of Gualaca, Allen 5050. flora of Panama (Moraceae) 4. CLARISIA R. & P. Clarisia R. & P. Prodr. Fl. Peruv. & Chil. 128. 1794. Sahagunia Liebm. in Kon. Danske Vidensk. Selsk. 2:316. 1851. Soaresia Fr. Allem. in Arch. Palestr. Scientif. Rio Jan. 1:142. 1858. Acanthinophyllum Fr. Allem. in Rev. Brazil. 1:368. 1858. Dioecious trees, the branches unarmed. Leaves alternate, distichous; stipules lat- eral. Both staminate and pistillate in- florescences secund racemes and spikes, or the pistillate subcapitate or in some species reduced to paired or solitary axillary flowers, bearing small, shortly stipitate, peltate bracts as in Trophis and Sorocea. Staminate flowers with a single stamen and an indefinite vestigial peri- anth, the anther broadly oval. Pistillate flowers epigynous, with 4 minute superior perianth lobes and an inconspicuous but persistent basal involucre of several pel- tate bracteoles, the style central and deep- ly 2-lobed. Fruit a small 1 -seeded drupe. About 8 species ranging from south- ern Mexico to Brazil and Peru according to Lanjouw (in Rec. Trav. Bot. Neerl. 33:254. 1936.). 1. Clarisia panamensis Woodson, spec. nov. Arbores usque ca. 10 m. altitudine attigentes ramulis sat validis glabris maturi- tate cortice tenue brunneo-griseo tectis. Folia alternata disticha lamina late obovato-elliptica apice obtuse subcaudato-acuminata basi obtusa 10-15 cm. longa 5-7 cm. lata membranacea glabra margine integra venis utroque latere ca. 10-12 petiolo valido ca. 1 cm. longo; stipulis lateralibus anguste lanceolatis ca. 1 cm. longis. Inflorescentiae femineae racemosae usque ca. 6-florae rhachide in fructu ca. 2 cm. longo glabro floribus ignotis. Inflorescentiae ac flores masculae ignotae. Fructus brevissime pedicellati late obovoideo-ellipsoidei ante maturitate usque 2.5 cm. longi 1 cm. crassi apice styli lobis persistentibus linearibus recurvatis ornati basi bracteis involucratis peltatis pluribus minutissime puberulis cincti. region north of El Valle de Anton, alt. 1000 m., September 27, 1946, P. H. I relationship of C colombiana (Rusby) Lanj. s known only from a single staminate specimen ANNALS OF THE MISSOURI BOTANICAL GARDEN (H. H. Smith 424) which has smaller, narrowly obovate, strikingly caudate leaves. The fruits of C. panamensis differ both in shape and in size from those of any other Clarisia presently known. Also to be expected in western Panama is C. mexicana (Liebm.) Lanj., which is known from the Golfo Dulce region of Costa Rica. It may be distinguished readily from C. panamensis by its rather narrowly oblong leaves and small globose 5. BROSIMUM Sw. Brosimum Sw. Prodr. Veg. Ind. Occ. 12. 1788, nom. conserv. Alicastrum P. Br. Hist. Jam. 372. 1756, nom. rejic. Piratinera Aubl. Hist. PI. Guinn. Fr. 2:888. 1775, nom. rejic. Galactodendrum Humb. Relat. Hist. 2:108. 1819. ore, in Trans. Linn. Soc. 2 ser. 4:473. 1895. perhaps occasionally dioecious trees. Leaves alternate, dis- chous; stipules lateral to fully amplexicaul. Inflorescences globular to turbinate, flora of Panama (Moraceae) 125 typically monoecious with 1-2 central pistillate flowers surrounded by numerous minute staminate commingled with more or less conspicuous peltate bracts, but infrequently dioecious. Pistillate flowers: epigynous, the perianth vestigial, the stigma deeply 2-lobed and apparently proterogynous. Staminate flowers with a definite 4-lobed perianth or the perianth vestigial or lacking; stamens 4, 2 or 1, the anthers oblongoid and basifixed to circular and peltate, dehiscing longitudinally to circumscissilly. Fruit a false drupe. Probably about 24 species ranging from southern Mexico to Argentina, and in the Antilles. A genus most interesting in its floral modifications and very badly in need of revision, as are most Moraceae. The first three species of the enumeration which follows would be referable to Brosimopsis if a narrow generic concept were followed consistently in this treatment. I have indicated in a previous paragraph that I am not in sympathy with such a view and follow it in other instances merely in deference to previously established custom. In this case, however, custom is not greatly involved. Amongst the Panamanian genera of Moraceae alone no less than four parallel reductions of the male flowers to a vestigial perianth and a single stamen may be observed: Trophis — » (Sorocea) -» Clarisia; Brosimopsis -» Brosimum; Perebea -» Castilla; Olmedia — » Pseudolmedia. The eight Panamanian species also show a most interesting transition in the structure of the anther from a rather commonplace basifixed type with longitudinal dehiscence to a most unusual peltate body with perfectly symmetrical circumscissile dehiscence, aptly described by Aublet (loc. cit. sub Piratinera) as "en forme de champignons". The series is illustrated in the accompanying figures. In the following text two new Costa Rican species are described, since they have been collected only a few miles from the Panamanian border and probably will yet be discovered in our flora. a. Flowers not wholly concealed before anthesis by the weakly expanded, Leaves relati vely large, 2- U , 3 -times as long as broad, gh .brous; ',r,e, the .ccd vely "small, 3- "softly cellous beneath; the seed about 1 .i -' .. , ■ .:...., • ■ stem; leaves rat: rongly unequal 1 roncealed before vers wholly c anthesis by the stron ;ly t r ■ anded'. th a vesthia! or obsolete perian -nplex. ' fully amplexicat »; heads globose. ANNALS OF THE MISSOURI BOTANICAL GARDEN lies 1.5-3.0 cm. long; flowering peduncles stout; anthers uely inserted on the filament, clearly longitudinal in dehis- . Leaves broadly ovate to ob] 1. Brosimum ojoche Woodson, spec. nov. Arbores magnae usque 35 m. altae ut dicitur, ramulis sat tenuibus glabris. Foliorum lamina oblonga vel obovato-oblonga apice obtuse acuminata basi late obtusa ca. 9-18 cm. longa 4-7 cm. lata firme membranacea glabra, petiolo 5-15 mm. longo; stipulis lateralibus ovato-lanceolatis 5-7 mm. longis cicatrice ramulum ca. dimidia cingenti. Inflorescentiae capituli usque 5—7 mm. pedunculati globosi ca. 1 cm. diametro metientes nostri prosus masculi an semper dioeci; florum masculorum tepalis 4 subliberis aequalibus ca. 1.5 mm. longis, staminibus 4 valde exsertis antheris oblongoideis apice setoso-appendiculatis basifixis longitudinaliter dehiscentibus ca. 1 mm. longis. Flores feminei ignoti. Fructus globosi ultra 2.5 flora of Panama (Moraceae) 127 Costa Rica: cartago: forets de Turrialba, alt. 570 m., Tonduz 8338 (US); paturages de Turrialba, alt. 650 m M Nunez 16353 (US); puntarenas: Esquinas forest, region between Rio Esquinas and Palmar Sur de Osa, alt. 75 m., Jan. 30, 1951, P. II. Allot 5809 (US, holotype) ; Hacienda Liscano, vallee du Bani, alt. m., Pittkr 12082 (US). A northern extension of B. ojoche may be represented by five specimens from British Honduras which are discussed briefly under B. terrabanum. This species and two which follow, although at present known only from Costa Rica, have been collected so close to the Panamanian border that it seems quite likely that sooner or later they will be found in Panama. The Allen specimen was identified previously as B. terrabanum Pittier, while the Nunez and Brenes specimens were determined as B. costaricanum Liebm. by Pittier, who himself included his own collection within Helicostylis montana Pittier which, as it happens, is synonymous with true Brosimum costaricanum. It is curious that Pittier, although particularly interested in Brosimum, twice described species of that genus as species of Helicostylis, a genus which has yet to make its appearance in Central America as far as I am aware. Brosimum ojoche, the specific epithet of which is taken from the vernacular name noted upon three of the cited sheets, is easily distinguished from B. costa- ricanum by the characters of the key. It appears more than somewhat questionable to me that the specimens from Cartago and those from Puntarenas could reasonably represent a single species but the foliage of the four sheets is strikingly similar. The Allen specimen is in flower while the three others are in fruit (in a rather pitiful state of disintegration). 2. Brosimum costaricanum Liebm. in K. Dansk. Vidensk. Selsk. Skr. ser. 5. 2:334. 1851. Helicostylis montana Pittier, in Contrib. U. S. Nat. Herb. 20:96. 1918. Brosimum sapiifolium Standi. & L. Wms. in Ceiba 3 :40. 1952. Trees up to 30 m. tall, the younger twigs relatively slender, softly white- puberulent to glabrate. Leaves relatively small, rather narrowly obovate-oblong, the tip rather abruptly subcaudate-acuminate, the base obtuse or rounded, 8-18 cm. long, 2-4 cm. broad, membranaceous, softly white- tomentellous or puberulent on the veins beneath; petioles about 5 mm. long. Inflorescences possibly dioecious, globose, the staminate with peduncles about 1 cm. long, very slender, about 1 cm. in diameter. Staminate flowers: tepals 5, nearly free and essentially equal, about 2 mm. long; stamens 3-4, widely exserted, the anthers basifixed, oblongoid, about 1 mm. long. Pistillate flowers presently unknown. Nearly mature fruits globose, about 1.5 cm. in diameter, with persistent peltate bracts, the seed about 1 cm. long. Costa Rica and Panama, in lowland forests. chiriqui: Paso de Quebrada Gato, between Hato Jobo and San Felix, Pittier 5426; Progreso, Cooper & Slater 188. Brosimum ramonense Standi, of eastern Costa Rica, which may be expected in western Panama, differs from B. costaricanum chiefly in the scant and appressed ferruginous pubescence of the leaves. (167) ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. Brosimum allenii Woodson, spec. nov. Arbores magnae ut dicitur usque ca. 40 m. altitudine attingentes vulgo palo de vaca incolarum appelatur quod latice potabili, ramulis crassiusculis internodiis congestis glabris. Folia late ovato-elliptica abrupte brevissime acuminata basi rotunda 9-20 cm. longa 5-12 cm. lata coriacea omnino glabra, petiolis crassis ca. 1.5 cm. longis; stipulis amplexicaulibus ovatis longe acuminatis ca. 1.5 cm. longis cicatrice ramulum transverse cingenti. Inflorescentiae globosi ca. 1.5 cm. diam. metientes nostri monoici flore unico feminei cum masculis floribus multis, pedun- culo crasso perbrevi ca. 8 mm. longo; riorum masculorum tepalis 5 inaequalibus ca. 1 mm. longis, staminibus 2 valde exsertis antheris oblongoideis basifixis longitudinaliter dehiscentibus ca. 1 mm. longis. Fructus non visi. Costa Rica: puntarenas: Esquinas forest, region between Rio Esquinas and Palmar Sur de Osa, alt. 75 m., Jan. 30, 1951, P. H. Allen 5813 (US, holotype). Mr. Allen reports that the latex of this species, which he associated with B. utile, the famous "cow tree" first discovered by von Humboldt, is "of good flavor, rather like condensed milk, good in coffee." I believe that B. allenii is sufficiently distinguishable from true B. utile through the characters of the preceding key. (168) FLORA OF PANAMA (Moraceae) 4. Brosimum guianense (Aubl.) Huber, in Bull. Mus. Goeldi 5:337. 1909. Piratinera guianense Aubl. Hist. Pi. Guian. Fr. 2:888; 4: t. 340. 1775. Piratinera panamr \>. U. S. Nat. Herb. 20:100. pi. 7. 1917. Brosimum panamense (Pittier) Standi. & Steyerm. in Field Mus. Publ. Bot. 23:40. 1944. Trees to about 25 m. tall, essentially glabrous throughout. Leaves oblong- obovate, abruptly and obtusely acuminate to subcuspidate, the base broadly obtuse to rounded, entire, 5-12 cm. long, 2.5-4.0 cm. broad, firmly membranaceous, the petiole about 5 mm. long; stipules lateral, about 5 mm. long. Heads broadly obconic to turbinate, usually somewhat lobed or convolute, about 7-10 mm. broad, the slender peduncle about 1.0-1.5 cm. long. Staminate flowers with a low vestigial perianth and 1 stamen with a rather thick filament about 2 mm. long and an obliquely inserted broadly oval anther up to 1 mm. long, associated with suborbicular shortly-stipitate peltate bracts. Pistillate flowers 1-3. Mature fruit British Honduras to Venezuela. I forests. ANNALS OF THE BOTANICAL GARDEN san blas: Puerto Obaldia, Pittier 4336. A photograph of Aublet's collection at the British Museum shows that Pittier was correct in associating P. panamensis with P. guianensis; they probably are conspecific. Pittier 4336 is represented in the U. S. National Herbarium by three sheets, each with several inflorescences. I have been unable to detect the plural pistillate flowers which Pittier considered so important in maintaining Piratinera distinct from Brosimum, either in the remnants of the inflorescences which Pittier himself supposedly dissected or in an additional one which I dissected myself. One head, however, does indeed appear to be pistillate (without external evidence of staminate flowers), but I have not had the temerity to dissect it. Here, as in other species of Brosimum in Panama, the heads would appear frequently to be unisexual. As Standley expresses it pointedly in the cited references, "In typical Brosimum there is only one pistillate flower, in Piratinera two; but with ordinary mens it is difficult to find even one pistillate flower, to say nothing vering two, and the difference is at least not a practical one." 5. Brosimu i Contrib. U. S. Nat. Herb. 20:102. 1918. Trees 20-25 m. in height, with young twigs rather stout and with rather close internodes, minutely and indefinitely puberulent when young, eventually develop- ing a rather thick periderm. Leaves rather broadly oblong, abruptly and acutely acuminate, rounded at the base, 10-25 cm. long, 3-10 cm. broad, subcoriaceous, sparsely puberulent to glabrate, the petioles stout, 5-15 mm. long; stipules amplex- icaul, lanceolate, long-acuminate, 2-3 cm. long, pilosulose, leaving oblique scars nearly encircling the stem. Inflorescences globose, monoecious, about 1 cm. in diameter, the peduncle of about equal length, rather stout, staminate flowers: without an obvious perianth and with a single stamen with a broadly oval anther (170) FLORA OF obliquely affixed to the filament, obviously longitudinal in dehisce] depressed-globose, 2.0-2.5 cm. in diameter. Costa Rica to Colombia. san blas: hills of Sperdi, near Puerto Obaldia, Pit tier 4345, 4418. This is von Humboldt's "cow tree" (palo de vaca) with potable lai Helicostylis latifolia latifolium Standi, bernadetteae Woodson, nom. Contrib. U. S. Nat i m. tall, the branches grayish brown, inconspicuously puberu- lent to glabrate. Leaves elliptic-obovate, broadly acuminate-subcuspidate to obtuse or rounded at the tip, broadly cuneate to rounded at the base, 6-12 cm. long, 3-6 cm. broad, with about 12-18 pairs of rather crowded veins, subcoriaceous, glabrous, the petioles about 4-7 mm. long; stipules almost fully amplexicaul, about 5-7 mm. long. Inflorescences subglobose, ours staminate, about 5 mm. in diameter at anthesis, the peduncle about 4-5 mm. long. Staminate flowers without an obvious perianth; stamen 1, the anther excentrically peltate, about 1 mm. in diameter, dehiscing by 2 basal valves. Pistillate flowers and fruit unknown. Panama, in lowland forests. ANNALS OF THE MISSOURI BOTANICAL GARDEN CANAL ZONE: B arro Colorado Island , Standby 41156, Kenoye Pittier 6584. panaj ia: Alhajuela, Chag res Valley, Pittier 3488. This species so closely resembles B. alicastrum that th nificant difference n the structure on the anthers had escape the slight but very sig- it was called to my attention by our artist for the flora of Panama, Mrs. Bernadette Velick. Nevertheless, the difference also apparently was noticed by Pittier and its significance over-emphasized as the synonymy testifies. Mrs. Velick was unable to detect the perianth segments described by Pittier, nor could I. After an examination of the type specimen of B. latifolium Standi. (Eggers I572I, F) , which bears a single immature pistillate head, I have concluded that it should be referred to B. alicastrum Sw. 7. Brosimum alicastrum Sw. Prodr. Veg. Ind. Occ. 12. 1788. Brosimum latifolium Standi, in Trop. Woods 42:26. 1935. Trees to about 30 m. tall, the young branches slender, grayish brown, glabrous or essentially so. Leaves oval to oblong-elliptic, obtuse to shortly subcuspidate- acuminate at the tip, broadly obtuse to rounded at the base, 5-15 cm. long, 2-6 cm. broad, with about 12-18 pairs of slender veins, firmly membranaceous, glabrous, the petioles about 5-7 mm. long; stipules almost fully amplexicaul, about 5 mm. long. Inflorescences subglobose, monoecious or dioecious, 3-6 mm. in diameter at anthesis, the peduncle slender, about as long as the heads or somewhat shorter, flowers with a very indefinite vestigial perianth; stamen 1, the antl Southern Mexico to Ecuador; Cuba and Jamaica, at low elevations. CHiRiQuf : Progreso, Cooper & Slater 2jo, 263. Brosimum bernadetteae, B. alicastrum and B. terrabanum appear to be well founded upon the basis of flower and inflorescence structure, but the leaves are not particularly distinctive. At the present time it is difficult to define their natural ranges not only because of the rather few fertile specimens available in herbaria but also because of the large number of sterile specimens labeled either as B. alicastrum or B. terrabanum which add considerable confusion and would much better have been left upon the trees. Brosimum 'itinera terral basinym, no Pittier, in Contrib. U. S. Nat. Herb. Lundell, in Carnegie Inst. Wash. Publ. 471 Trees up to about 25 m. tall, the young branches moderately stout, reddish brown, indefinitely pilosulose to glabrate. Leaves broadly ovate to oblong-ovate, narrowly subcaudate-acuminate at the tip, broadly obtuse to rounded at the base, 10-25 cm. long, 3-9 cm. broad, with 18-22 pairs of prominent veins, subcoriaceous, glabrous, the petioles quite stout, about 1 cm. long; stipules almost fully amplex- icaul, about 1 cm. long, minutely pilosulose. Flowering heads apparently usually dioecious, subglobose, 4-6 mm. in diameter at anthesis, the peduncles very slender, two or three times longer than the heads. Staminate flowers with an indefinite vestigial perianth; stamen 1, the anther circular, centrally peltate, about 1 mm. in diameter, dehiscing circumscissilly. Fruit unknown. (173) 134 ANNALS OF THE MISSOURI BOTANICAL GARDEN Costa Rica and Panama (doubtfully in Guatemala and El Salvador), in lowland bocas del toro: Farm 8, region of Almirante, Cooper 441. Lundell misapplied B. terrabanum to include two specimens from British Hon- duras, Gentle 17 37 and Schipp 522, which, together with three others of the same collectors deposited in the herbarium of the Missouri Botanical Garden (Schipp J 360; Gentle 3307, 3440) appear to me probably to represent a northern extension of B. ojoche. 6. BATOCARPUS Karst. Batocarpus Karst. Fl. Colomb. 2:67, 1863, emend. Woodson, non sensu Fosberg in Proc. Biol. Soc. Wash. 55:101. 1942. Dioecious trees. Leaves alternate, distichous; stipules lateral. Staminate inflo- rescences in discoid involucrate heads, the bracts and bracteoles basifixed, not peltate. Pistillate inflorescences in globose heads without involucral bracts nor obvious bracteoles. Staminate flowers with a deeply 4-lobed perianth and 4 stamens with broadly oval longitudinally dehiscent anthers apically attached to the fila- ments. Pistillate flowers with a fleshy tubular accrescent perianth, the ovary superior and with a central deeply 2-lobed papillate stigma. Fruit a globose rather fleshy syncarp. Two well-authenticated species, the following and B. costaricensis Standi, and L. Wms., which may be expected in western Panama. 1. Batocarpus orinocensis Karst. Fl. Colomb. 2:67. pi. 134. 1863. Trees to about 10 m. in height, the young twigs slender, minutely puberulent when very immature, becoming glabra te. Leaves broadly elliptic to oval-obovate, rather abruptly subcaudate-acuminate, broadly obtuse to rounded at the base, 8-20 cm. long, 4-7 cm. broad, entire, membranaceous, glabrous, the petiole slender, about 1 cm. long; stipules about 5 mm. long, narrowly lanceolate. Staminate heads discoid, about 1 cm. in diameter, prominently involucrate with several broadly ovate minutely tomentellous bracts about 2 mm. long, the peduncle about 8 mm. long. Pistillate heads globose, about 2 cm. in diameter, the peduncle about 1 cm. long, without a definite involucre. Panama to Amazonian Colombia and Peru. bocas del toro: Buena Vista Camp on Chiriqui Trail at 12 50 ft., Cooper 601. The staminate heads of this specimen (US) indeed superficially resemble those of Brosimum costaricanum Liebm. as originally identified, but dissection discloses the absence of the peltate bracteoles essential to Brosimum. The shape and size of the leaves and the character of the stipules agree startlingly with the excellent plate of a pistillate specimen of Batocarpus orinocensis provided by Karsten, and there appears to me no doubt concerning the identity of the two. flora of Panama (Moraceae) 135 136 ANNALS OF THE MISSOURI BOTANICAL GARDEN If this be true, our specimen thus provides the first information concerning the staminate structures of the genus, known previously only from Karsten's plate and the citation of an additional pistillate specimen from Peru by Fosberg (loc cit. 1942), since the second species, Batocarpus costaricensis Standi. & L. Vms., also is known only from pistillate specimens. This view is in conflict with that of Fosberg that Batocarpus and Anonocarpus Ducke, a monotypic genus, are congeneric. Anonocarpus amazonicus Ducke (in Archiv. Jard. Bot. Rio Jan. 3:39. 1922) fortunately was described with full details of both staminate and pistillate inflorescences, the former being elongate spikes of naked 1 -staminate flowers. Since the pistillate inflorescences of Bato- carpus and Anonocarpus have much in common, as noted by Ducke, it is easy to sympathize with Fosberg's impulse to merge the two. This now becomes less plausible since inflorescence structure is such an inflexible criterion of genera in Moraceae. Batocarpus costaricensis, presently known from western Puntarenas Province, Costa Rica, and to be expected in western Panama, is well distinguished from B. orinocensis by the prominently repand-serrate leaves and nearly sessile pistillate 7. PEREBEA Aubl. Perebea Aubl. Hist. PI. Guian. Fr. 2:952; 4: t. 361. 1775. Mikania Neck. Elem. Bot. 2:217. 1790, non Willd. Dioecious trees. Leaves alternate, distichous, entire to rather obscurely undulate-serrate toward the tip; stipules fully amplexicaul. Inflorescences axillary, solitary or clustered, involucrate, discoid. Staminate more or less united; stamens 4, nearly included, the anth flowers numerous and all fertile; tepals 4, free or somewhat united, accrescent and somewhat pulpy in fruit but essentially free and united only at the base; ovary superior to subinferior, the style central, the 2 stigma lobes short and broad to narrow and filiform. Fruit a more or less fleshy, weakly united syncarp. Perhaps 4-6 species from Costa Rica to Brazil and Bolivia. Of the related South American genera may be mentioned Helicostylis, which differs from Perebea in the exinvolucrate inflorescences, the pistillate being most peculiar in having only one or very few of the epigynous central flowers fertile and the peripheral flowers reduced to a sort of enveloping involucre composed of the accrescent fleshy sterile perianths, a. Leaves predominantly cordate or subcordate, the stipules 2-4 cm. long; flora of Panama (Moraceae) 137 (177) 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. Perebea guianensis Aubl. Hist. PL Guian. Fr. 2:952; 4: /. 361. 1775. Perebea castilloides Pittier, in Contrib. U. S. Nat. Herb. 13:43 8. *. 80-81. 1912. Perebea pseudopeltata Mildbr. in Notizbl. 10:184. 1927. Perebea tessmannii Mildbr. loc. cit. 185. 1927. Perebea laevigata Standi in Trop. Woods 16:36. 1928, nom. nud. in syn. Perebea glabrata Standi, in Field Mus. Publ. Bot. 4:201. 1929. Perebea acantbogyne Ducke, loc. cit. 11:579. 1932. Small trees infrequently to 20 m. tall, with rather stout fiexuose ferruginous- hirsutulous branches. Leaves broadly oblong-elliptic grading toward obovate or ovate, the apex abruptly and narrowly subcaudate-acuminate, the base predom- inantly more or less deeply cordate, infrequently broadly obtuse or rounded in young leaves, entire to indefinitely undulate-serrate toward the tip, 1 5-40 cm. long, 7-15 cm. broad, firmly membranaceous, more or less ferruginous-hirtellous, par- ticularly beneath, the petiole rather stout, 5-10 mm. long; stipules narrowly ovate-lanceolate, long-acuminate, 2-4 cm. long, densely sericeous. Staminate heads about 1.0-1.5 cm. in diameter, the peduncle about 1 cm. long. Pistillate heads superficially similar to the staminate with about 25-50 flowers. Fruiting heads oblate-ovoid, about 5-6 cm. broad, the achenes about 1 cm. long. Panama to Brazil and Bolivia, in lowland forests. Reported as cerillo by Standley, ule and cuacho by Proctor Cooper. bocas del toro: Cricamola, Cooper 523; Punta Pefia, near Chiriqui Grande, Pittier s. n. canal zone: west of Pina Base Camp, Limon Bay, Johnston 1607; Frijoles, Standley 2 7453- san blas: Perme, Cooper 634. The association of these plants with Aublet's name is made with some misgiv- ings. Perebea guianensis was excellently and unmistakably illustrated by Aublet, but it is doubtful whether the holotype exists at Paris, since Trecul apparently was not able to find it and my friend Dr. F. G. Meyer did not find an isotype at the British Museum. Dr. Meyer did find a staminate specimen, apparently of this species collected by Martin in 1804, a duplicate of which is now available not only at the British Museum (and presumably Paris?) but at the Missouri Botanical Garden. This sheet corresponds generally to those cited from Panama on the one hand and on the other to several collections from South America which, because they are now distributed under other names, I shall cite both in support of my interpretation and for the use of others: Brazil: Krukoff 4827, 7041, 7209, 8076, 8350, Ducke 1221 (24-5-1940) Peru: Klug 2719, K. C Lorenz 3210—2W deposited in the herbarium of the Missouri Botanical Garden. For positive identi- fication fruiting specimens from Panama are necessary, and I must confess that some of the Panamanian sheets are noticeably more pubescent than these from Brazil and Peru. 2. Perebea xanthochym a Karst. Fl. Colomb. 2:23. /. 112. 1862. Perebea integrifolia Karst. loc. cit. 1862. Castilloa markhamiana J. Collins, Rept. Caoutch. 12. /. 3. 1872. Perebea markhamiana (J. Collins) Benth. ex Hook, f . in Trans. Linn. Soc. 2 ser. 2:211. 1886. Perebea hispidula Standi, in Ann. Missouri Bot. Gard. 29:350. 1942. (178) FLORA OF PANAMA (Mo Small trees to about 10 m. tall, with rather slender and scarcely flexuose hispidulous to glabrate branches. Leaves oblong-elliptic, subcaudate, acuminate to abruptly subcuspidate at the tip, the base broadly obtuse to rounded, usually more or less conspicuously undulate-serrate toward the tip, infrequently subentire, 8-28 cm. long, 3-9 cm. broad, rather inconspicuously appressed-puberulent beneath to nearly glabrous, the petiole about 5 mm. long; stipules lanceolate, 5-10 mm. long. Staminate heads about 5 mm. in diameter, sessile or subsessile. Pistillate heads usually with about 5-10 flowers about 5 mm. in diameter, the peduncle rather slender, 3-5 mm. long, fruiting heads subsessile, about 1 cm. in diameter, the achenes to about 1 cm. long. Costa Rica to Colombia, in lowland forests. Island, Von Wedel 1035. These sheets agree very well with Karsten's plate which illustrates both P. xanthochyma and P. integri folia (the chief differentium of which would appear to be sex!), and with a fragment of Karsten's isotype of the former (F). 8. CASTILLA Cerv. Castilla Cerv. in Gac. Lit. Mexico, Suppl. 7. 1794. Monoecious or dioecious trees (cf. O. F. Cook, in Science, ser. 2. 18:437. 1903; H. Pittier, in Contrib. U. S. Nat. Herb. 13:253. 1910), the branches unarmed. Leaves alternate, distichous, closely and minutely ciliate-denticulate to essentially entire; stipules fully amplexicaul. Inflorescences axillary, solitary or clustered, involucrate, the staminate discoid and conduplicate-flabellate, the pistil- late discoid or subglobose. Staminate flowers essentially unorganized and naked, with stamens bearing oval anthers apically affixed to the filament. Pistillate flowers more or less coherent, with a fleshy accrescent conic- tubular perianth and a subinferior ovary, the stigma 2-lobed almost to the base of the central style. Fruit a more or less fleshy syncarp. Probably no more than 4-6 species extending from southern Mexico to Bolivia. Pittier (loc. cit. 1910) provides a very difficult key to 10 species which I have found to be quite impractical when applied to herbarium specimens; I suspect that it would also present problems in the field. From the evidence available to me, I can visualize at most two scarcely distinguishable species in Panama. The genus has been of sporadic interest in the past as a possible source of commercial pistillate flowers half < reddish pulp in fruit.... Leaves obtuse to acuu pistillate flowers cohere 140 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. Castiixa elastica Cerv. in Gac. Lit. Mexico, Suppl. 7. 1794. Castilla panamensis Cook, in Science, n. ser. 18:438. 1903. Trees 5-10 m. tall, the young branches densely hirsute with spreading (less frequently appressed) golden hairs, eventually glabrate. Leaves oblong-obovate, more or less cordate and usually not strongly inequilateral at the base, subcuspidate- acuminate at the tip, 20-30 cm. long, 10-14 cm. broad, membranaceous, minutely and closely ciliate-denticulate, both surfaces golden spreading-hirsute but par- ticularly beneath, the petioles about 1 cm. long; stipules 3-6 cm. long. Inflores- cences in clusters of 2-4 in catenate series of the upper leaf-axils, occasionally solitary. Staminate heads conduplicate-reniform, about 1.5 cm. long and 2 cm. broad, the peduncle about 1 cm. long. Fruiting heads thickly discoid, 4-5 cm. in diameter, about 1.5 cm. thick, sessile or subsessile, the component flowers half or more coherent, developing an orange or reddish pulp at maturity. Southern Mexico to Colombia and perhaps southward; in moist forests at low elevations. Ule-ule, bule, caucho, mastate bianco. bocas del toro: Laguna de Chiriqui, Hart 140; Chiriquicito, Seibert 155I; Punta Rovalo, Seibert 1566; Almn- nth Dale Farm, Seibert 1586, 1587, 1588. canal zone: old site of Gorgona, Maxon 6783; Las Cascadas Plantation, Standley 25697, 29497, Cook # Martin 46; Ancon Hospital grounds, Maxon 6769, Gaillard s. n. cocle: El Valle de Anton, Allen 3625. darien: along Sambu River, Pittier 5526; Boca de Pau- arando, Pittier 5714. Panama: Rio Tapia, Standley 28048, 26186, 26177; Juan Dias region, Maxon & Harvey 6748; Arraijan, Maxon tf Cook 7032, 7033; Rio Tecumen, Standley 29374; Tumba Muerto Road, near Panama, Standley 20818. This species has been of more than passing i nearly two centuries, but has never becom 2. Castilla tunu Hemsl. in Hook. f. Icon. PL ser. 4. 7: pi 2651. 1901. Castilla fallax Cook, in Science n. ser. 18:438. 1903. Trees 5-20 m. tall, the young branches densely hirsute with appressed rather dingy-brown hairs, eventually glabrate. Leaves broadly elliptic to oblong-obovate, at the base, 15-40 cm. long, 7-15 cm. broad, minutely and closely ciliate- denticulate to subentire, membranaceous, appressed-hirsute above and below to glabrate, the petiole about 1 cm. long; stipules 4-5 cm. long. Inflorescences paired or solitary in the upper leaf-axils. Staminate heads conduplicate-reniform, about 2 cm. long and 2.5 cm. broad, sessile or subsessile. Fruiting heads oblate-globose, 2-3 cm. in diameter, quite sessile, the component flowers coherent only toward the base, greenish and nearly dry at maturity. British Honduras to Panama, in moist forests at low elevations. ina and Yaviza, indefinite, Cooper 284. This species, according to Allen, "is reported as NOT being used for rubber" in Darien, which Pittier confirms from his experience in Costa Rica. According flora of Panama (Moraceae) 142 ANNALS OF THE MISSOURI to Pittier, the species also is known to occur in Chiriqui, in the vicinity of 1 In spite of the faults which both Pittier and Cook have found with Her description and plate, there appears to me little doubt that C. fallax is a super 9. POULSENIA Eggers Poulsenia Eggers, in Bot. Centralbl. 73:65 [err. 49]. pi J. 1898. Inophloem Pittier, in Journ. Wash. Acad Sci. 6:113. 1916. Monoecious trees, the young branches usually more or less aculeate, rarely un- armed. Leaves alternate, distichous; stipules fully amplexicaul. Both staminate and pistillate inflorescences of globose or subglobose heads. Staminate flowers many: perianth of 4 scarcely united tepals; stamens 4, with broadly oval, basifixed anthers. Pistillate flowers few (3-9): perianths tubular-conic, shortly 4-dentate, strongly coherent in the head; ovary superior, the stigma deeply 2-lobed, widely exserted. Fruiting head a fleshy syncarp. One species ranging from British Honduras to Bolivia. 1. Poulsenia armata (Miq.) Standi, in Trop. Woods 33:4. 1933. Olmedia (?) armata Miq. in Seem. Bot. Voy. Herald 196. 1854. Poulsenia aculeata Eggers, in Bot. Centralbl. 73:66 [err. 50]. 1898. Inophloem armata (Miq.) Pittier, in Journ. Wash. Acad. Sci. 6:113. 1916. Coussapoa rekoi Standi, in Contrib. U. S. Nat. Herb. 20:211. 1919. flora of Panama (Moraceae) 143 Trees to 25 m. tall, all young parts scattering!/ hispid and aculeate to glabrous and smooth. Leaves broadly and rather obliquely oval or elliptic, the tip obtuse or rounded to shortly subcuspidate-acuminate, the base broadly obtuse, 8-40 cm. long, 4-20 cm. broad, entire, subcoriaceous, the petiole 1-3 cm. long; stipules 2-3 cm. long. Staminate heads globose, without an obvious involucre, many-flowered, about 1 cm. in diameter. Pistillate heads subglobose or ovoid, without an obvious involucre, 3-9-flowered, about 1.5 cm. in diameter. Mature fruiting heads bocas del toro: Changuinola Valley, Cooper & Slater 88; Almirante, Cooper 455. canal zone: Alhajuela, Pit tier 3731 ; Gatiin Valley, Pit tier s. n.; Barro Colorado Island, Shattuck 1 104, Carpenter 75. colon: Dos Bocas, Fato Valley, Pit tier 4202. darien: Pittier reports that the bark is very thick and fibrous and is soaked and pounded into cloth (mastate) by the Darien Indians, a use previously reported by Seemann. 10. OLMEDIA R. &P. Olmedia R. & P. Fl. Peruv. & Chil. Prodr. 129. t. 28. 1794. ? Maquira Aubl. Hist. Pi. Guian. Fr. Suppl. 2:36; 4: t. 389. 1775. Dioecious trees. Leaves alternate, distichous; stipules wholly amplexicaul. Staminate inflorescences involucrate discoid heads. Staminate flowers definitely organized; perianth campanulate, broadly 4-dentate; stamens 4, the anthers basi- fixed, oblongoid. Pistillate inflorescences with a solitary prominently bracteate- involucrate flower, the ovary superior, the stigma with 2 widely exserted filiform lobes. Fruit a small slightly fleshy false drupe. One species in Central America; the South American species are impossible to ascertain at present because of our faulty knowledge. 1. Olmedia aspera R. & P. Fl. Peruv. & Chil. Prodr. 129. t. 28. 179 4. Rather small trees or shrubs to about 6 m. tall, the young twigs rather slender, not conspicuously flexuose, densely to sparsely hispidulous, eventually glabrate. Leaves obovate- or oblanceolate-elliptic, rarely irregularly lyrate, narrowly subcaudate-acuminate at the tip, acutely or obtusely cuneate at the base, frequently more or less falcate, 7-20 cm. long, 2-9 cm. broad, usually more or less con- spicuously and irregularly serrate-undulate toward the tip, firmly membranaceous, scabrous or hispidulous to glabrous, the petioles about 5-7 mm. long. Staminate inflorescences solitary or in clusters of 2-4, about 1 cm. in diameter, the peduncles about 5 mm. long. Pistillate inflorescences solitary or in small axillary clusters, the peduncles about 5-7 mm. long, the involucral bracts ovate, 2-4 mm. long, persistent. bocas del toro: Chiriqui Lagoon, Von Wedel 1043, 1094; Water Valley, Von Wedel 896, 941, 1582, 1- jo. can ' 6747 > Johnston 1584, Standby 27209; Quebrada Salamanca, Steyermark Sf Allen 17142; Rio Pina-Rio Media divide, Johnston J 703l Quebrada La Palma, Dodge (3 Allen I?3 6 5; Mindi Hills, Johnston 1733; Darien Station, Standley 31597; Cerro Gordo, near Culebra, Standley 26015; Gorgona, Maxon (18}) ANNALS OF THE MISSOURI BOTANICAL GARDEN 47 '44; Barro Colorado Island, Standley 31450, Woodworth & Vestal 372, Abbe 46, 666, Salvoza 930, 939, Bangham 521, 530. chiriqui: San Felix, Pittier 529I; Paso Quebrada Gato, Pittier 5425; Hato del Jobo, Pittier v/-H. coci ii: El Valle de Anton, Allen 1736. darien: Pinogana, Pittier 6578. Panama: Rio Tecumen, Standley 29454; Chepo, Pittier 4707. This is the only Panamanian species of Moraceae where I have been able to find anything comparable to the familiar leaf polymophism of the northern mulberries. 11. PSEUDOLMEDIA Tree. Pseudolmedia Tree, in Ann. Sci. Nat. 3 ser. 8:129. t. 5. 1847. Olmediopsis Karst. Fl. Colomb. 2:17. t. 109. 1862. Dioecious trees. Leaves alternate, distichous; stipules fully amplexicaul. Staminate inflorescences in discoid involucrate heads of numerous extremely re- duced flowers represented by a solitary stamen with an oblongoid basifixed anther (1*4) flora of Panama (Moraceae) 145 and an extremely vestigial perianth, intermingled with more prominent spatulate bracteoles. Pistillate flowers solitary with an involucre of several closely imbricated bracts, perigynous or epigynous; perianth tubular with 4 minute dentate lobes; stigma narrowly 2-lobed, widely exserted. Fruit a false drupe subtended by the persistent involucre. 4-5 species of the Antilles, Central America and northern and western South PSEUDOLMEDIA S (Sw.) Griseb. Fl. Brit. W. Ind. 152. 1859. Brosimum spurium Sw. Prodr. 12. 1788. Pseudolmedia havanensh Tree. In Ann. Sci. Nat. 3 ser. 8:130. 1847. Brosimum caloxylon Standi, in Trop. Woods 17:11. 1929. Trees 8-20 m. tall, the young branches relatively slender, glabroi ovate- to oblong-elliptic, rather abruptly and bluntly subcaudate-ac subcuspidate, the base obtuse to rounded, firmly membranaceous 1 entire, 9-15 cm. long, 3-5 cm. broad, glabrous or essentially so, the petiole 3-5 mm. long; stipules vary narrowly lanceolate, long-acuminate, 1-2 cm. long. Staminate heads usually paired, sessile or subsessile, about 8-10 mm. in diameter, the bracts broadly obtuse, minutely puberulent. Pistillate heads sessile, the flowers ovoid, about 2 mm. long, softly puberulent, the stigma lobes about equally long. Drupes broadly ovoid, fleshy, about 1 cm. long and broad. British Honduras to northern South America; Antilles. Reported as cuqua by Cox, bloodwood and cacique by Proctor Cooper. £ocas del toro: Buena Vista Camp on Chiriqui Trail, Cooper 6o? '; "Bocas del Toro Region", Cox s. «.; Cricamola Valley, Cooper 535. All three collections are quite sterile, hence their assignment to this species is not infallible. It is true that these leafy sheets could be matched fairly well by some South American species of Brosimum or even Ficus. But this guess appears 146 ANNALS OF THE MISSOURI 12. FICUS [Tourn.] L. By GORDON P. DEWOLF, Jr. Ficus [Tourn.] L. Gen. Plant, ed. 5. 482. 1754. OZ««i<« Raf. Sylva Tell. 58. 183 8. Urostigma Gasp. Nov. Gen. Fie. 7. 1844. Pharmocosycea Miq. in Hook. Lond. Journ. Bot. 7:64. 1848. (Synonyms not referring to American species not cited.) A genus (at least so far as the American species are concerned) of soft-wooded, generally smooth-barked trees and shrubs with milky or opalescent latex. Many species begin life as epiphytes, or epiliths, which may eventually, through the coalescence of their roots, completely encircle the trunk of, and strangle, their host. Leaves entire and spiral in native species — rarely opposite and sometimes toothed or lobed in the Old World. Stipules long or short, enfolding the buds, generally quickly deciduous but rarely persistent, leaving a scar surrounding the stem. Flowers unisexual, borne over the inner surface of a hollow, globose, more or less fleshy structure (the receptacle or "fig") the apical pore (ostiole or orifice) of which is closed by a series of interlocking bracts. The female flowers are of two kinds: functional females, generally sessile and maturing into viable achenes, and sterile females (gall flowers), generally stalked and functioning as incubators for the larvae of a wasp which pollinates the fertile flowers. Male, female and gall flowers are completely intermixed in American species. The figs are generally solitary or paired, borne among the leaves but sometimes on specialized short shoots behind the leaves in American species. In Old "World species they may be borne similarly or in leafless racemes or panicles on the trunk and larger branches. There are about 750 species in the tropics of both hemispheres. In the American tropics the genus is represented by two subgenera (of three) and about 70 species. In addition to the native species of Panama, numerous exotics are cultivated, particularly F. elastica, F. nitida and the banyan, F. religiosa, and the root- climbing ivy-like F. pumila. These usually can be distinguished by their persistent aerial and prop roots which are infrequent in the native species. The common edible fig with deeply lobed leaves, F. carica, is seldom encountered in Subgenus urostigma (Gasp.) Miq. Most species of fig are disseminated by birds which eat the more or less fleshy receptacles and wipe their beaks clean of the sticky pulp upon the branch of a convenient tree. The small seeds embedded in the pulp deposited in this way germinate epiphytically, the seedlings grasping the supporting tree with their tentacles of aerial roots which in time reach the soil. Meanwhile the woody body of the fig develops rapidly and the supporting tree eventually is destroyed by the weight of its triumphant epiphyte. For this reason the name matabalo (tree-killer) often is anplied to the strangler-figs of Central America. The appropriate Spanish words hipo for the fleshy receptacles and hiynero for the tree are current in Panama. flora of panama (Moraceae) 147 Subgenus pharmacosycea Miq. The Pharmacosyceas seldom, if ever, begin life as epiphytes. They are large, free-growing trees of the forest and of second growth. The subgenus was originally erected for a few American species but it is now known that about 46 Australasian species belong here as well. In any consideration of a taxonomic treatment of tropical American plants it is important to realize that botanically scarcely one quarter of tropical America has been adequately explored. Literally thousands of square miles of rainforest and campo have either never been investigated by botanists or have not been so for fifty to one hundred years. Even so, our herbaria are full of alleged species whose only claims to distinction, when they are examined critically, are the political boundaries separating them. With the exception of the West Indies, Mexico, parts of Central America and some areas in northern South America, what knowledge we have of the lowland evergreen forests seems to be restricted to the vegetation of the riverbanks. In addition, few, even of modern collectors, have bothered to make field notes — or, if they have made them, have not bothered to transcribe them onto labels to accompany their specimens. The net result is that, although there is a good deal of (literally scrappy) material in our herbaria, we can get only the most vague idea either of distribution or of variation of the species involved, and we know practically nothing of the ecology of the plants. As a final example and indict- ment, Ficus nymphaeaefolia is apparently a fairly common species from Panama to the State of Amapa in Brazil at the mouth of the Amazon. Even as tropical trees go, it must be striking in appearance. Yet the only description of this species as a living plant that I have been able to find is in Humboldt's "Personal Narrative ..." of his travels in the American tropics in 1799-1804. The floral characters of American figs, unlike those of many Asian and African species are disconcertingly uniform. The precise form and structure of the fig itself is, however, particularly significant. In addition, microscopic vegetative characters have considerable taxonomic significance. These characters were first investigated by Otto Renner in 1907. More recently they have been exploited and practical techniques for their elucidation devised by Professor E. J. H. Corner, to whom I am greatly indebted for knowledge of them. Over the past three years I have been able to see nearly all of the types of American species of Ficus preserved in European herbaria. More than a year ago, I was able to present to the Board of Research Studies of the University of Cambridge a taxonomic and nomenclatural revision of the species of Ficus, proposed on the basis of allegedly American material from the time of the publication of the Species Plantarum in 1753 to the first proper taxonomic revision of the genus by Miquel in 1847-48. This involved a taxonomic study of all the species known from Brazil, northeastern South America and the West Indies, as well as a cursory survey of the species of the Andes and of Central America and Mexico. Over and above the courtesies extended to me by the directors and staffs of all I have visited or corresponded with (which will be acknowledged 148 ANNALS OF THE MISSOURI BOTANICAL GARDEN explicitly in a forthcoming paper) , I am particularly indebted to the directors of the New York Botanical Garden and the United States National Museum, who allowed me to borrow certain specimens in their care to use as a basis for study of the taxonomy of the species. I am also in particular debt of gratitude to the American Academy of Arts and Sciences who generously made a grant to allow me Some months ago, Dr. R. E. Woodson, Jr., hearing of my interest in the genus, sent me for comment a manuscript treatment which he had prepared for the flora of Panama. It is a pleasure to record that, with one or two minor excep- respondence, Dr. Woodson asked me to prepare a fresh treatment of Ficus for the Flora. The following descriptions and synonomy are the result. I have seen little Panamanian material for any species. The descriptions are drawn from the material which I have seen, modified freely from Dr. Woodson's manuscript. The specimens cited are quoted directly from that manuscript. The nomenclature and synonymy are based (with one or two exceptions) on my own studies. what beaked below 1 gs borne in pairs in the le , : Y^u-i ar glandular frequently pre subgenus urostigma) Figs cylindrical, to 20 mm. long, with dense ling yellow- :..:■■ d. Stipules glabrous; figs dd. Stipules appressed-pube e. Leaves smooth above : n the\ eins beneath. ee. Leaves scabrid above. ,s 8-30 mm. in : 1 ,-' . luncle. FLORA OF PANAMA (Moraceae) . Figs 15- JO mm. in diameter. ' a shelf or socket on which figs are borne, basal bracts fused to the base of fig for about 5 mm 18. F. tuerckheimii "' fi^ToA^m^'iniKJmeMr.!. .' !lJ. F. trigonata 1. Ficus maxima P. Mill. Gard. Diet. ed. 8 (Ficus no. 6). 1768. Ficus citrifolia hort. ex Lamck. Encycl. Meth. (Bot.) 2:494. 1786, quoad syn. excl. typ (in herb. Lamarck) . Ficus laurifolia hort. ex Lamck. loc. dt. 2:495. 1786, quoad syn. excl. typus (spec. herb. Desfontaines) . Ficus virens Dryand. in Ait. Hort. Kew. ed. 1. 3:451. 1789, quoad syn. excl. typus. Ficus anthelminthica Rich, ex DC. Ess. Pi. Med. 267. 1804 (fide herb. Richard), noi nud. Ficus radula H. & B. ex Willd. loc. cit.' 1 144. 1 806. Pharmacosycea grandaeva Mart, ex Miq. in Hook. Lond. Journ. Bot. 7:70. 1848. Pharmacosycea guyanensis Miq. loc. cit. 67. 1848. Pharmacosycea glaucescens Liebm. in Kong. Danske Vidensk. Selsk. Skr. ser. 5. 2:332. 18! Pharmacosycea hernandezii Liebm. loc. cit. 1851. Pharmacosycea rigida Miq. in Seem. Vot. Voy. Herald. 195. 1854, nee Desf. nee Jack. Pharmacosycea mexicana Miq. in Versl. en Med. Kon. Akad. Wetensch. 13:415. 1862. Pharmacosycea pseudoradula Miq. loc. cit. 414. 1862. Ficus suffocans Griseb. Fl. Brit. W. Ind. 150. 1864. Ficus glaucescens (Liebm.) Miq. Ann. Mus. Bot. Lugd.-Bat. 3:300. 1867. (189) 150 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ficus hernandezii (Liebm.) Miq. loc. cit. 1867. Ficus parkeri Miq. loc. cit. 1867. Ficus me xic ana 299. 1867. Ficus pseudoradula (Miq.) Miq. loc. cit. 1867. Ficus coybana Miq. loc. cit. 300. 1867. Ficus guadalajarana S. Wats, in Proc. Amer. Acad. 26:151. 1891. Ficus finlayana Warb. in Urb. Symbol. Antill. 3 :487. 1903 (pro parte) . Ficus picardae Warb. loc. cit. 484. 1903. Ficus rubricosta Warb. loc. cit. 486. 1903. Ctbrida Warb. loc. cit. 485. 1903. Ficus ulei Warb. ex Ule, in Engl. Bot. Jahrb. 40:141. 1907, nom. nud. Ficus plumieri Urb. in Fedde, Repert. 15:158. 1918. Ficus bopiana Rusby, in Mem. N. Y. Bot. Gard. 7:23 0. 1927. Ficus ulei Rossb. in Fedde, Repert. 42:60. 1937. Ficus vicencionis Dugand, in Caldasia 2:385-386. 1944. Small to large trees 5-30 m. tall. Twigs 3-5 mm. in diameter, glabrous or pubescent, with a thin, scurfy, yellowish brown periderm. Stipules to 25 mm. long, narrowly deltoid, glabrous, puberulent, or pubescent at the base. Lamina 2.5-12 cm. wide X 6-24 cm. long, elliptic, broadly elliptic, lanceolate, oblanceo- late, or obovate, glabrous above, glabrous or puberulent beneath; apex blunt, acute, departing from the midrib at an angle from 0°-30°; basal veins departing from the midrib at an angle of 40°- 60°; intercostals slightly raised. Petiole 5-40 mm. long, %-% the length of the lamina, the epidermis generally scurfy. Figs 10-25 mm. in diameter, globose, or sub-globose, glabrous or puberulent, sometimes with a stalk above the basal bracts 2-7 mm. long, borne among the leaves; color green or yellow, sometimes mottled darker; peduncle 2-25 mm. long, glabrous or pubescent; basal bracts 1-2 mm. long, deltoid, glabrous or pubescent; orifice 1-2 mm. in diameter, flat, or the bracts slightly outflexed. Southern Mexico to and through the Amazon basin, lowlands to about 1000 ft. , bocas del toro: Changuinola Valley, Cooper & Slater 92, Dunlap 408. canal zone: south of Ft. Sherman, Johnston l6j6; Balboa, Standley 25611, 27006; Corozal, Standley 26829; Ancon, Pittier 2728; Culebra, Pittier 2211. cocle: Penonome, Williams 251. Panama: San Jose Island, Johnston 449, 533, 1338, 1383; Taboga Island, Standley 27976; Chepo, Pittier 4757; Juan Dias, Standley 30481. This seems to be a relatively common species in lowland forest and in lowland hill forest from Cuba and southern Mexico to, and through, the Amazon basin. In the low mountain rain forest in Jamaica this species occasionally overtops the forest canopy (Apsey, 1953), and it forms an element of the flora which springs up on waste land left after removal of bauxite in western Jamaica (Howard and Proctor, 1957). Despite the multitude of names which have been proposed for various examples of this species, it does not seem to be particularly variable. There is, of course, considerable variation in leaf size (which is reflected, at least to some extent, in lateral vein number) between young and old plants and shoots, and, of course, leaves from vigorous shoots seem to be generally more strongly acuminate than leaves from mature twigs. The oldest legitimate name for this species as here defined is Ficus maxima Mill. (190) flora of Panama (Moraceae) 151 There is no specimen from Miller's herbarium in the British Museum (NH) , so the name must be typified on Ficus indica maxima, folio oblongo ... of Sloane's "Catalogus ..." 189 (1696) and "Voyage ..." 2:140, t. 223 (1725). There is no specimen in the Sloane collection at the B.M. (NH) and the illustration is certainly not diagnostic, but the description is good and sufficiently specific, so that I have no hesitation in designating it the nomenclatural type of Miller's name and following Fawcett and Rendle (Fl. Jam. 3 1 :48-49. 1914) in referring it to the plant which for a century has been known as Ficus suffocans Griseb. Ficus protensa (Griseb.) Hemsl. was included in the synonymy of Ficus glaucescens by Standley, but I do not know on what grounds. No type was cited in the original description and I have no record of having seen a specimen so labeled. The description is certainly not diagnostic of this taxon. I think that it is unwise to doubt Grisebach's assignment of his plant to the subgenus urostigma without being able to demonstrate his error from the original material. I have not seen authentic material of Ficus guadalajarana, S. Wats. It is included here on the authority of Dr. Woodson. I have seen the types of all the 2. Ficus Tonduzii Standi, in Contrib. U. S. Nat. Herb. 20:8. 1917. Ficus macrocyce Pittier, sensu Dugand. Large buttressed trees to about 15 m. tall. Twigs 4-7 mm. in diameter, glabrous, the epidermis frequently exfoliating, with a thick greyish brown peri- derm. Stipules 25-60 mm. long, narrowly deltoid, glabrous. Lamina 6-15 cm. wide X 12-30 cm. long, broadly ovate, elliptic or obovate; apex rounded acute, acute or very shortly and bluntly acuminate; base rounded or rounded cuneate; lateral veins sometimes very broad and conspicuous, 9-1 5 pairs, departing from the midrib at an angle of 10° or less; intercostals conspicuous, but not much raised; basal veins departing from the midrib at an angle of 15°- 30°, continuous with the very conspicuous submarginal vein. Petiole 20-80 mm. long, 4-5 mm. thick, V2-V& the length of the lamina. Figs 20-35 mm. in diameter, globose, glabrous or minutely scabrid, borne among the leaves; color not noted; peduncle obsolete to 10 mm. long, sometimes with a stalk above the bracts to 5 mm. long; basal bracts 2-3 mm. long, very broadly deltoid; ostiole about 2 mm. in diameter, plane with the surface of the receptacle, or produced in a beak 5-10 mm. high. Costa Rica to northern Colombia, near sea level to about 640 m. canal zone: Barro Colorado Island, Wetmore Gf Abbe 303, Carpenter 5, Shattuck 199; Obispo, Standley 31706. chiriqui: Hiqueron, Cooper & Slater 249. 3. Ficus macbridei Standi, in Publ. Field Mus. Nat. Hist. (Bot.) 13:305. 1937. Ficus torresiana Standi, loc. cit. 18:3 07. 1937. Trees to 15 m. tall. Twigs 7-9 mm. in diameter, glabrous or very shortly brown-pubescent, the epidermis sometimes exfoliating. Stipules 50-75 mm. long, narrowly deltoid, glabrous or minutely pubescent. Lamina 12-28 cm. broad X 26-42 cm. long, broadly ovate; apex acute or slightly acuminate; base rounded or (191) 152 ANNALS OF THE MISSOURI BOTANICAL GARDEN emarginate; lateral veins 12-16 pairs departing from the midrib at an angle of 10°-30°; basal veins 1-3 pairs, departing from the midrib at a similar angle or at an angle from 30°- 50°; intercostals slightly prominent. Petiole 20-85 mm. long, glabrous or pubescent, %— % the length of the lamina. Figs 15-25 mm. in diameter, obovoid, glabrous or pubescent, borne among leaves ( ? ) ; color not noted, but in dried material with yellowish macula tions; peduncle obsolete; basal bracts 2- 3 lobed, about 2 mm. long; orifice about 2 mm. in diameter, slightly crateriform. This species, so far as I know, has not yet been found in Panama. It is found in Costa Rica and Peru, however, and should be looked for in areas between. 4. Ficus insipida Willd. Sp. PL ed. 4. 4:1 143. 1806. Ficus glabrata HBK. Nov. Gen. & Spec. 2:47. 1818. Ficus adhatodaefolia Schott, ex Spreng. Syst. Veg. ed. 16. 4 (App.) :409. 1827. in us anihclminthica Mart, in Spix et Mart. Reise in Brasilien 3:1158. 1831. Pharmacosycea vermifuga M ^0. 1848. Pharmacosycea angustifolia Liebm. in Kong. Danske Vidensk. Selsk. Skr. ser. 5.2:333. 1851. View I amifuga (Miq.) Miq. Ann. Mus. Bot. Lugd.-Bat. 3:300. 1867. Ficus segoviae Miq. loc. cit. 1867. :dnia S. Wat. in Proc. Am. Acad. 26:151. 1891. Ficus krugiana Warb. in Urb. Symbol. Antill. 3 :487. 1903. Ficus crassiuscula Warb. ex Standi, in Contrib. U. S. Nat. Herb. 20:12. 1917. Ficus werckleana Rossb. in Fedde, Repert. 42:60. 1937. Ficus boyacensis Dugand, in Caldasia 1 4 :31. 1942. Ficus crassa Kl. & Karst. ex Dugand, in Caldasia, l 4 :35-36. 1942. Large or small buttressed trees, 8-40 m. tall. Twigs 2-6 mm. in diameter, generally glabrous, with a thin reddish or greyish brown periderm. Stipules 30- 125 mm. long, narrowly deltoid, generally glabrous. Lamina 2-11 cm. wide X 5-25 cm. long, lanceolate to broadly elliptic, essentially glabrous, generally glossy above, sometimes slightly inequilateral; apex blunt or acute, to acuminate; base cuneate, round cuneate, rounded or emarginate; lateral veins 10-25 [-30] pairs, departing from the midrib at an angle of 40° or less; basal veins departing from the midrib at an angle from 30°-60°; intercostals slightly prominent. Petiole 10-65 mm. long, %-% the length of the lamina, the epidermis not exfoliating, though frequently wrinkled. Figs 1 5-3 mm. in diameter, globose with or with- out a stalk above the basal bracts, 1-6 mm. long, glabrous or pubescent, borne among the leaves; color green or yellowish-green; peduncle 3-22 mm. long, thin or stout; basal bracts three, 1-3 (-5) mm. long, deltoid or semicircular; orifice flat, or somewhat crateriform or mammillate, 2-4 mm. in diameter, 1-2 mm. high. Southern Mexico to Southern Brazil, in lowland forests. bocas del toro: Almirante, Cooper 444. canal zone: upper Chilibre River, Seibert 1 512; Ancon, Piper 6006; Chagres River, Muenscher 12284, 12291% Ft. Sherman, Johnston 1510; Barro Colorado Island, Bailey & Bailey 479, Carpenter 3, Standley 4093 1 ' Wetmore & Abbe 126. chiriqui: Rio Dupi, Pittier 2539. colon: Rio Fato, Pittier 3880. darien: Boca de Cupe, Williams 679; Yaviza, Allen 4589. This is, undoubtedly, the most widely distributed of the Pharmacosyceas, though with the exception of one doubtful record it does not occur in the West Indies. It ranges, on the continent, from northwestern Mexico to Paraguay. In Mexico FLORA OF PANAMA (Mo 153 and Central America it seems to be a component of the arid or sub-arid forests — but, since Standley notes that figs are usually left standing when forests are cleared and since much of the accessible forest of Central America and Mexico is not virgin, it seems unwise to try to guess as to the natural habitat. Standley records, of the populations which he called "mexicana," that they were stranglers. This is, presumably, a very unusual state of affairs for any plant of this subgenus. Both Dugand (Caldasia 3 12 : 133-148. 1944) and Little (in sched.) report the use of the latex of this species as a vermifuge in Colombia. The south Brasilian vernacular name figueira purgante and Miquel's epithet vermifnga for material from this same area, suggests that its medicinal usage is widespread. In addition, Archer (in sched.) records that Para the latex is sometimes used in the coagulation of the latex of Hevea. 5. Ficus popenoei Standi, in Field Mus. Publ. Bot. 4:301. 1929. Fiats tolimenm Standi, he. cit. 17:177. 1937. Trees to 25 m. tall. Twigs 4-6 mm. in diameter, fulvous spreading pubescent, eventually glabrate and developing a moderately thick greyish brown periderm. Stipules about 10 mm. long, densely spreading fulvous pubescent. Lamina 5-15 cm. long X 4-10 cm. wide, broadly oval to obovate-oblong, pubescent below and scabrid above; lateral veins 5-9 (-10) pairs, departing from the midrib at an I angle of 20°- 40°; basal veins departing j at a similar angle; intercostals somewhat prominent. Petiole 5-15 mm. long, pubescent, Vq-Yiq the length of the lamina. Figs 14-22 mm. long X 9-16 borne among the leaves; color green, brown tinted (yellow-brown pubescent in dried material) ; peduncle 3-5 mm. long, pubescent; basal bracts broadly deltoid 2-5 : ng, pubesc about 2 slightly umbonate. British Honduras to central Colombia, in lowland forests. 15169. PA] Island, Johnston 483, Dr. Johnston reports that the habit of this species is quite variable: some may be stranglers, while others may germinate in the soil; a few develop a banyan-like stilted habit, but this is exceptional. This seems to be a relatively common plant in Central America which has been identified, in the past, as Ficus velutina. There may be some vegetative similarity, but the cylindrical, densely yellow-brown pubescent figs are quite distinctive. (193) ANNALS OF THE MISSOURI BOTANICAL uiniaefolia A. Richard. Fl. Cub. Fanerog. 3:221. t. 72. 1850. Urostigma eugeniaefolium Liebm. in K. Danske Vid. Sclsk. Skrivt. 5 5 er. 2:329. 1851. Urostigma oerstedianum Miq. in Seem. Bot. Voy. Herald 196. t. 36. 1854. Urostigma liebm an v. una m Miq. U. cir. 195. 1854. Urostigma chiriquianum Miq. in Versl. en Med. Kon. Akad. 13:412. 1862. mims (Miq.) Miq. in Ann. Mus. Bot. Lugd.-Bat. 3:298. 1867. ) Miq. loc cit. 1867. Ficus oerstedianu:. .299. 1867. Ficus eugeniaefolia (Liebm.) Hemsl. Biol. Centr.-Amer. Bot. 3:144. 1883. ifl Urb. Symbol. An till. 3:464. 1903. Ficus omphalophora Warb. loc. cit. 466. 1903. Ficus wilsoni Warb. loc. cit. 467. 1903. Shrubs or small to medium sized trees to 30 m. tall. Twigs 2-3 mm. in diameter, glabrous, with a rather close brownish grey periderm. Stipules 4-15 mm. long, glabrous, ciliolate or glaucous. Lamina 1-5 cm. wide X 1-7-1 1 cm. long, elliptic or obovate; apex rounded, acute, or short, abruptly acuminate; base cuneate or narrowly rounded, sometimes somewhat emarginate; lateral veins 6-14 pairs, departing from the midrib at an angle from 20°- 30°; basal veins departing from the midrib at a similar angle; intercostals not, or only somewhat, prominent. Petiole 4-10 mm. long, %-% the length of the lamina. Figs 3-7 mm. in diameter, globose, glabrous, borne among the leaves; color reddish; peduncle 1-5 mm. long, glabrous or minutely puberulent; basal bracts two, 1-2 mm. long, ovate, glabrous or minutely puberulent; orifice 1-2 mm. in diameter, somewhat mammillate. Guatemala to Colombia, at intermediate or higher elevations; and in the Bahama Islands and the Greater Antilles. Mat a palo. canal zone: Barro Colorado Island, Wilson 122, Carpenter 64, Woodworth # Vestal 658, Bangham jog. chiriqui: Gualaca, Allen 5026; Boquete, Davidson 61 2; Caldera, \l\en 2228. darien: Cana, 26243. veraguas: Rio Canazas, Allen I 5 8. This is an exceedingly variable taxon, the West Indian plants called Ficus jacquiniaefolia and F. sintenisii being, superficially, very different in appearance. There is, however, no important character by which they can be circumscribed and it seems wise, therefore, to include them here. Ficus perforata L. is based on /. 132, f. 2. of Burmann's edition of Plumier's plates. Neither Warburg nor Urban were sure of the identity of the plant, but I believe that Warburg was correct in suggesting that it is the plant which he called F. omphalophora, the common shrubby fig of the Lesser Antilles. F. wilsoni of Jamaica is indistinguishable, save only that the stomates are somewhat larger. 7. Ficus hartwegii (Miq.) Miq. Ann. Mus. Bot. Lugd.-Bat. 3:299. 1867. Urostigma hartwegii Miq. in Hook. Lond. Journ. Bot. 6:545. 1847. Ficus colubrinae Standi, in Contrib. U. S. Nat. Herb. 20:16. 1917. Trees, 10-35 m. tall. Twigs 3-4 mm. in diameter, appressed pubescent to flora of Panama (Moraceae) 155 glabra te, soon developing a rather thin yellowish brown periderm. Stipules to 20 mm. long, deltoid, appressed pubescent. Lamina 2.5-8 cm. wide X 3.5-13.5 cm. long, broadly elliptic or obovate, appressed pubescent on the veins; apex short abrupt acuminate; base rounded, emarginate or cuneate; lateral veins 4-6 (-8) pairs, departing from the midrib at an angle from 20°-30° (-40°); basal veins departing from the midrib at an angle from 30°-40° (-60°); intercostals only slightly prominent. Petiole 6-30 mm. long, ( %-) %-ty the length of the lamina, appressed pubescent. Figs 6-7 mm. in diameter, globose, glabrous, borne among the leaves; color pale red; peduncle obsolete; basal bracts two, to 2 mm. long, semi- circular, pubescent; ostiole to 2 mm. in diameter, plane in a very slight rim of receptacular tissue. Guatemala and British Honduras to Colombia, in lowland forests and thickets. bocas del toro: Old Bank Island, Von Wedel 1999; Cricamola Valley, Cooper 519; Almirante, Cooper 404. canal zone: Fort Sherman, Johnston 1626, 1748, Standley 3H22; Fort Randolph, Standley 28676; France Field, Standley 30339; Barro Colorado Island, Shattuck 239, Carpenter 20, Woodworth 8 Vestal 57 1, Kenoyer 319. cocle: EI Valle de Anton, Allen 2000, 2478, 2889. The basal veins appear to be three but are accompanied on either side by a According to Dugand (Caldasia 2 9 : 274-275. 1943) this is an extremely common fig in the central Andes of Colombia between 500 and 1600 m. altitude. 8. Ficus dendrocida HBK. Nov. Gen. & Spec. PI. 2:46. 1817. Ficus elliptic* HBK. loc. cit. 1817. 1 ^ - :' M 1 > -"! Ficus deniroctonia Spreng. Syst. Veg. ed. 16. 3:780. 1826. Large spreading trees, to 30 m. tall, frequently with many air roots. Twigs 2-3 mm. in diameter, pubescent, developing a rather thick dark brown periderm. Stipules 5—8 mm. long, narrowly deltoid, appressed pubescent. Lamina 2.5-5 cm. wide X 5-8.5 cm. long, elliptic or slightly obovate, scabrid above, pubescent beneath; apex acute; base rounded; lateral veins 5-8 (-12) pairs, departing from the midrib at an angle from 30°-40°; basal veins departing from the midrib at an angle 30°- 50°; intercostals somewhat raised. Petiole 6-15 mm. long, %r%o the length of the lamina. Figs 5-6 mm. in diameter, globose, pubescent or glabrescent, borne among the leaves; color not noted; peduncle obsolete; basal bracts 1-2 mm. long, connate, semicircular, pubescent; ostiole 1-2 mm. in diameter, flat. Panama and Colombia, perhaps to northern Brazil. Panama: margins of tidal swamp, Pacora, Allen 3446. This is a species which seems to be almost completely confined to the drainage of the Rio Magdalena in Colombia, below 700 m. altitude. What little we know of its variation is due to the efforts of Dr. A. Dugand (Caldasia 2 8 : 169-171. 1947) , who notes that it is extremely variable in size, shape, texture and pubescence of (195) 156 ANNALS OF THE MISSOURI BOTANICAL GARDEN 9. Ficus pertusa L. f. Suppl. Plant. 442. 1781. Ficus padifolia HBK. Nov. Gen. & Sp. 2:47. 1817. Ficus complicata HBK. loc. cit. 48. 1817. Ficus ciliolosa Link, Enum. Pi. Berol. 2:450. 1822. folia Link, loc. cit. 1822. foUa Link, loc. cit. 1822. ■ >!ia Schott, ex Spreng. Syst. Veg. ed. 16. 4 (App.) :409. 1827. udia Hook, et Arn. Bot. Beech. Voy. 310. 1841. Ficus arpazusa Cai s. 15. 1842. Ficus cerasifolia Kunth & Bouche, Ind. Sem. Hort. Berol. 16. 1846. Ficus planicostata Kunth & Bouche, loc. cit. 1846. Ficus periplocaefolia Kunth & Bouche, loc. cit. 1846. Ficus consanguinea Kunth & Bouche, loc. cit. 1846. Urostigma erythrostictum Miquel, in Hooker, Lond. Journ. Bot. 6:540. 1847. Urostigma geminum Ruiz ex Miquel, in Hooker, Lond. Journ. Bot. 6:547. 1847. Urostigma scheideanum Miquel, in Hooker, Lond. Journ Bot. 6:539. 1847. Pharmacosycea peruviana Miquel in Hooker, Lond. Journ. Bot. 7:72. 1848. Urostigma baccatum Liebm. in Kong. Danske Vidensk. Selsk. Skr. 5 ser. 2:327. 1851. Urostigma sapidum Liebm. in Kong. Danske Vidensk. Selsk. Skr. 5 ser. 9:327. 1851. Urostigma turbinatum Liebm. in Kong. Danske Vidensk. Selsk. Skr. 5 ser. 9:327. 18 51. Urostigma sulcipes Miq. in Vers. Med. Akad. Amsterdam, 13:413. 1862. Ficus ochroleuca Griseb. Fl. Brit. W. Ind. Isl. 151. 1864. Ficus faydeni Miq. Ann. Mus. Bot. Lugd.-Bat. 3:219. 1867. Ficus baccata (Liebm.) Miq. loc. cit. 3:299. 1867. Ficus erythrosticta (Miq.) Miq. loc. cit. 3:298. 1867. Ficus liebmanniana Miq. loc. cit. 1867. Ficus gemina (Ruiz, ex Miq.) Miq. loc. cit. 1867. Ficus sapida (Liebm.) Miq. loc. cit. 1867. Ficus sulcipes (Miq.) Miq. loc. cit. 1867. it. 1867. Ficus fasciculata S. Wats, in Proc. Amer. Acad. 24:78. 1889, non King (1888). Ficus sonorae S. Wats. loc. cit. 1889. Ficus immersa Warb. in Bull. Soc. Bot. Fr. 59 (Mem. 3) :641. 1912. (1913), nom. nud. Ficus palmicida Pittier, in Bol. Soc. Venez. Cienc. Nat. 4 30 :69. 1937. Ficus peruviana (Miq.) Rossb. in Fedde, Repert. 43:61. 1937. Ficus kanukuensis Standi, in Lloydia. 2:174. 1939. Ficus tarapotina Warburg, mss. Large spreading trees to 30 m. or more tall. Twigs 1-2 mm. in diameter, glabrous, or pubescent, developing a yellowish grey periderm. Stipules 5—7 mm. long, narrowly deltoid, glabrous or minutely puberulent. Lamina 1-5 cm. wide X 2.5-12 cm. long, elliptic or lanceolate; the acumen 5-20 mm. long; base c seldom rounded and slightly emarginate; lateral veins (4-) 6-12, departing from the midrib at an angle from 20°- 60°; intercostals somewhat raised. Petiole 5—30 mm. long, %-% (- 1 /i 2 ) as long as the lamina. Figs 5-18 mm. in diameter, globose, glabrous, borne among the leaves; color pink or yellow or green with brown flecks; peduncle 2-5 mm. long, glabrous; basal bracts 1-4 mm. long, semi- circular, glabrous; ostiole 1-2 mm. in diameter, crateriform, the bracts sunken into the receptacle or surrounded by a raised ring of receptacular tissue. Southern Mexico and Jamaica to Paraguay. In lowland to highland forests and savannas. Mata palo t higuito. (196) flora of Panama (Moraceae) This, and Ficus citrifolia are undoubtedly the most widespread of the American species of the genus. This species, which is immediately recognized by its peculiar crateriform ostiole and narrowly lanceolate or elliptic leaves, differs from F. citrifolia in being absent from all of the West Indies except Jamaica, and being very common in Mexico and Central America (The Jamaican population is obviously related to the Central American!). There is a very definite distinction between populations on the basis of fruit size, but there is no sharp discontinuity the populations on the Caribbean coast of South America from Venezuela east to the Amazon, and in the Amazon basin, are relatively small fruited, while the Jamaican, Mexican, Central American, and southern and central Brazilian popu- lations are relatively large fruited. With a few exceptions, in the island of Jamaica, the Brazilian population has larger figs than the Central American. This seems to me a peculiar, and probably quite significant pattern of distribution, but of what, or how, I cannot say. It goes without saying that an intensive study of this species in the field should be most rewarding. The synonymy of this species is extensive but relatively simple. The name Ficus pertusa is based on a specimen from Surinam which represents the small fruited populations of that area. F. subtriplinervia, F. erythrosticta, F. gemina, F. tarapotina and F. kanukuensis all refer to this same general population. F. padifolia is the earliest name for the relatively large fruited populations in Colombia, Central America, Mexico and Jamaica. F. complicata, F. lancifolia, F. ochroleuca, F. faydeni, F. baccata, F. sulcipes, F. sapida and F. turbinata all refer to the same general population. Ficus arpazusa is the earliest name for the plants of south Brazilian and eastern Peruvian regions, which Warburg called F. immersa. 10. Ficus trachelosyce Dugand, in Caldasia 4:69. fig. 14. 1942. Strangler trees to about 25 m. tall, the young branches slender, glabrous, devel- oping a thin yellowish periderm. Leaves oblong-elliptic, narrowly subcaudate- acuminate, broadly obtuse at the base, 6- 12 cm. long, 2.5-5.0 cm. broad, rather closely pinnate-veined, submembrana- ceous, glabrous, the petiole 1.5-2.0 cm. long, slender; stipules narrowly lanceolate, subcoriaceous, about 1.5 cm. long. Re- ceptacles paired at the nodes, subglobose, about 15 mm. broad, glabrous, the osteole THE MISSOURI BOTANICAL GARDEN collar about 5 mm. long; involucral long; peduncle rather slender, about rid Colombia, probably also in southwestern Costa Rica, in lowland chiriqui: Puerto Armuelles, Allen 6295. The above description was prepared by Dr. Woodson. I strongly suspect that this is simply a form of Ficus pertusa. 11. Ficus bullenei I. M. Johnston, in Sargentia 8:113. 1946. Trees to 21 m. tall; twigs 3-4 mm. in diameter, frequently conspicuously angled, densely spreading reddish-brown pubescent, eventually developing a rather thick grey periderm; stipules 10-20 mm. long, deltoid, densely appressed, reddish- brown pubescent; lamina 45-110 mm. wide X 100-200 mm. long, oblong-obovate to broadly obovate or oval, densely spreading pubescent, especially below, becom- ing glabra te above; apex rounded, acute, or minutely acuminate; base rounded, rounded cuneate, or cuneate, sometimes emarginate; lateral veins 4-8 pairs, departing from the midrib at an angle of 20°- 40°; basal veins departing from the midrib at a similar angle or from 40°- 60°; intercostals not prominent; petiole 10-30 mm. long, %-%n tne length of the lamina. Figs about 15 mm. in diameter, globose, minutely and densely spreading pubescent, borne among the leaves; color not noted; peduncle 1-4 mm. long; basal bracts two, 4-5 mm. long, ovate-deltoid; orifice about 2 mm. in diameter, surrounded by a raised collar of receptacular tissue, 2-3 mm. high. Panama, in lowland forests and savannas. 1 1837. Panama: San Jose Island, Johnston 23 1, 389, 2. Ficus citrifolia P. Mill. Gard. Diet. ed. 8. Ficus no. 10. 1768. Ficus pedunculata Dryand. in Ait. Hort. Kew. ed. 1. 3:450. 1789. Ficus populifolia Desf. Tabl. ed. 1. 239. 1804, nom. mid. (fide herb. ! Ficus populnea Willd. Sp. PL ed. 4. 4:1141. 1806. Ficus laevigata Vahl, Enum. 2:183. 1806. Ficus leiitiginosaVM, loc. cit. 1806. Ficus gigantea HBK. Nov. Gen. & Spec. 2:48. 1817. rt. Berol. 2:448. 1822. Fktu exinOa Schott, ex Spreng. Syst. Veg. ed. 16. 4 (App.) :410. 182: Ficus pyri folia Desf. Cat. Hort. Par. ed. 3. 413. 1829. Ficus catesbaei Steud. Nomencl. ed. 2. 1:636. 1840. Ficus botryapioides Kunth & Bouche, Ind. Sem. Hort. Berol. 15. 1846 Ficus syringae folia Kunth & Bouche, loc. cit. 1846. Urostigma angtt . Lond. Journ. Bot. 6:539. 1 8< Urostigma amazonicum Miq. loc. cit. 541. 1847 Ficus brevifolia Nutt. Sylva, ed. 1. 2:3. 1854. Ficus surhtamemh Miq. 1 Ficus angustifolia (Miq.) Ficus amazonica (Miq.) Miq. loc. cit. 1867. Ficus thomaea Miq. loc. cit. 299. 1867. Ficus populoides Warb. in Urb. Symbol. Anti flora of Panama (Moraceae) ] Ficus brittonii Boldingh, Fl. Curasao 20. 1914. Ficus hemsleyana Standi, in Contrib. U. S. Nat. Herb. 20:29. 1917 (non King, 1887). Ficus guaranitica Chodat, in Bull. Soc. Bot. Geneve. 2 ser. 11:254. 1919 (1920). Ficus turbinata Pitt, in Bol. Soc. Venez. Cienc Nat. 4 30 :61. 1937 (non Willd. 1806). l;na Dugand, in Caldasia l i :66. 1942. Ficus standleyana Dugand, loc. cit. 2:441. 1944. Ficus antimanensis Pitt. Mss. Ficus dugandii auct. Trees to 16 m. tall, or in exposed places, a shrub 0.3-2 m. tall. Twigs : mm. in diameter, glabrous, with a thin yellowish periderm. Stipules 5-30 n long, narrowly deltoid, glabrous, sometimes somewhat cm. wide X 2.5-20 cm. long, lanceolate, ovate, elliptic ovate, oblong, or obovate; apex acute to acuminate; base rounded, rounded-cuneate cuneate, or truncate, frequently emarginate or subcordate; lateral veins 4-16 pairs, departing from the midrib at an angle from 10°-40°; basal veins, departing from the midrib at a similar angle, or to 60°; intercostals not, or scarcely, prominent. Petiole 7-70 mm. long, slender, l/ 2 -l/ 7 (-%) the length of the lamina. Figs 6-12 [-15] mm. in diameter, globose, glabrous, borne among the leaves; color reddish or yellowish when ripe; peduncle 2-18 mm. long, glabrous or puberulent; basal bracts two, 2-3 mm. long, broadly deltoid to semicircular, with a hyaline margin, glabrous or puberulent; orifice 2-3 mm. in diameter, flat or very slightly raised. Florida to Paraguay, in lowland forests. canal zone: Chagres, Fettdler 286; Limon Bay, Johnston 1533, 1548; Culebra, Pit tier 2318; Balboa, Standley 25473, 25580; Barro Colorado Island, Standley 41129. Panama: San Jose Island, Johnston 109, 494, 680, 752, 1063, 1077, nog, 1168, 1177, 1381, J 3^7, I3$9> 1390; Taboga Island, Standley 27887. F. hemsleyana Standi. Standley published this name as a substitute for the epithet verrucosa of Liebmann. Standley apparently did not see the type, for the material that he has cited which I have seen (Tonduz 11576; Fendler 286) repre- sents a form of F. citrifolia. Oersted's plant is a very different thing, perhaps allied to F. petiolaris or F. obtusifolia. Ficus citrifolia, as I understand it, is a species which ranges from Florida to Paraguay. Though there seems to be little direct evidence, I presume that this is frequently a species of disturbed habitats. Specimens from relatively high altitudes (600-1700 m.) in the Greater Antilles and Venezuela and Colombia (F. populoides, F. turbinata, F. subandina) may represent a separate taxon, though present infor- mation suggests that they are only ecological forms. There is not enough informa- tion at present to evaluate their status. The leaf shape is generally ovate, with the base rounded and/or more or less emarginate. The leaf bases may vary from round to emarginate on the same twig. Along the northern coast of South America and the eastern coast of Central America, from the Guianas to Honduras, the leaf form seems to be predominantly oblong (F. angusti folia, F. surinamensis, F. hemsleyana, F. standleyana) . Forms occur, however, which are transitional to the normal leaf form of the species. 160 ANNALS OF THE MISSOURI BOTANICAL GARDEN The type of Ficus citrifolia is in the Banksian Herbarium at the British Museum (NH). It consists of a twig with two leaves and a peduncle with the remnants of a fig. There is no doubt that it represents this common lowland fig of the West Indies. The Catesby plant to which Miller makes reference is not a particu- larly good representation, but N. L. Britton refers to it here. 13. Ficus trigonata L. Pi. Surinam. 17. 1775. Ficus crassinervia Desf. ex Willd. Sp. Pi. ed. 4. 4:1138. 1806. Ficus berteroi Warb. in Urb. Symbol. Antill. 3:468. 1903. Ficus eggersii Warb. loc. cit. 469. 1903. ■■';.' Warb. loc. cit. 1903. Ficus hartii Warb. loc. cit. 458. 1903. Ficus coombsii Warb. loc. cit. 456. 1903. Ficus mitrophora Warb. loc. cit. 457. 1903. Ficus yucatanensis Standi, in Cont. U. S. Nat. Herb. 20:33. 1917. Ficus pittieri Standi, loc. cit. 31. 1917, pro parte, quoad paratypus. Ficus campbellii I. M. Johnston, in Sargentia 8:114. 1949. v I. M. Johnston, loc. cit. 117. 1949. Ficus fawcettii Britton, Mss. Large, spreading trees, to 15-25 m. tall. Twigs 2-5 mm. in diameter, glabrous or pubescent, developing a greyish-yellow periderm; stipules 15-27 mm. long, deltoid, glabrous or pubescent. Lamina 3-12 cm. wide X 4.5-25 cm. long, elliptic, or generally somewhat obovate; apex sometimes acute or even slightly acuminate, but generally rounded; base cuneate, rounded, or emarginate, sometimes slightly inequilateral; lateral veins (5-) 6-11 pairs, departing from the midrib at an angle from 20°-40°; basal veins 1-2, departing from the midrib at an angle from 30°- 50°; intercostals slightly prominent. Petiole 7-50 mm. long, %-% the length of the lamina. Figs 10-14 (-30) mm. in diameter, globose, glabrous or pubescent, borne among the leaves; color green when immature, red when mature; peduncle obsolete, or 2-10 mm. long, glabrous or puberulent; basal bracts two, 2-5 mm. long, deltoid or semicircular, sometimes variously split, variously glabrous, ciliolate, or puberulent; orifice 1-3 mm. in diameter, plane or slightly umbonate. Greater Antilles and the Carribbean coast of Central America from Yucatan to Colombia. canal zone: Gamboa, Pittier 2602; Chagres, Fendler 285; south of Fort Sherman, Johnston 1640. cocle: El Valle de Anton, Allen 198 1; Penonome, Williams 404. Panama: San Jose Island, Johnston 3, 49$, 695, 721, 730, 701, o?o, tot 2. TF2T, Ti2^. 1 127, 1353, 1384; Erlanson 106, 206; Miller 1910. This is apparently one of the common figs of the Greater Antilles. While there is considerable vegetative variation within the species, there seems to be no discernable geographical restriction of the variation and, consequently, there seems to be little justification in recognizing a distinct species on each island. Besides this, it seems to be fairly common in Central America. I have not seen all of the material cited by Standley under the name Ficus velutina, but the Panama collection (Pittier 335 g) is surely this species. flora of Panama (Moraceae) 161 Ficus paraensis (Miq.) Miq. in Ann. Mus. Bot. Lugd.-Bat. 3:298. 1867. Ficus panamensis Standi, in Contrib. U. S. Nat. Herb. 20:15. 1917. / >bora Benoist, in Arch. Bot. (Caen) Bull. 3:171. 1929 (1931). Ficus haughtii Standi, in Publ. Bot. Field Mus. Nat. Hist. 17:170. 1937. Ficus uberrima Standi, loc. cit. 177. 1937. Ficus putumayonis Dugand, in Caldasia 1 4 :62. 1942. Ficus arukensis Standi, in Bull. Torr. Bot. Club 75:295. 1948. Ficus manabiensis Standi. Mss. Shrubs, or small to large trees to 10 m. tall. Twigs 2-7 mm. in diameter, glabrous or minutely puberulent, developing a greyish-brown, striate, periderm. Stipules to 35 mm. long, narrowly deltoid, glabrous or minutely puberulent. Lamina 2.5-9 cm. wide X 6-27 cm. long, oblong, oblong-elliptic, oblong- oblanceolate or oblanceolate, glabrous; apex acuminate, the acumen commonly 5- cuneate; lateral veins 9-18 pairs, departing from the midrib at an angle from 10°- 30 ; basal veins, at least the strongest, departing from the midrib at an angle from 40°- 60°; intercostals scarcely, or somewhat, prominent. Petiole 5-35 (-80) mm. long, i/ 3 -% (-1/^) the length of the lamina. Figs 10-18 mm. in diameter, globose to sub-pyriform, glabrous or minutely puberulent, borne among the leaves; color greenish or yellowish, sometimes with red streaks and sometimes with crimson basal bracts; peduncle obsolete to 3 mm. long; basal bracts 2-5 mm. long, ovate, connate, puberulent; ostiole 2-4 mm. in diameter, umbonate, to 3 mm. high. Southern Mexico to Peru and northern Brazil, in lowland forests. canal zone: Barro Colorado Island, Bailey I colon: Rio Fato, Pittier 389 3, 3908. panam/ 1 1 34, 1174 I3&5> I409- The ripe receptacles appear to be olive green with purplish stripes and promi- nent purplish osteoles. Johnston (in Sargentia 8:118-119, 1949, sub F. pana- mensis), notes that although a strangler, F. paraensis does not appear to kill its As here understood, Ficus paraensis is a common and widespread species in the lowland rainforest, ranging from southern Mexico (fide Standley) along the Caribbean coast to Panama, thence along the western coast of South America at least as far as Ecuador, and along the eastern coast of South America to the mouth of the Amazon, and westward from this to the foot of the Andes. There are a good many collections, particularly from Central America and the western part of the Amazon basin, with rather thick (5-7 mm. in diameter) twigs. In these same areas, and on the western coast of South America, forms with strongly emarginate (or cordate) leaf bases are also common. I suspect that these collec- tions represent juvenile plants or shoots. Throughout the area of distribution, mention is made that ants build nests on the stems and branches of plants of (201) ANNALS OF THE MISSOURI BOTANICAL 15. Ficus i P. Mill. Gard. Diet. ed. 8. Ficus i Ficus numphaeifolia L. Mant. Plant. Alt. 305. Ficus nymphoides Thunb. Ficus Genus Dissert, Ficus ierensis Britton, in Bull. Torr. Bot. Club Ficus anguina R. Benoist, in Bull. Mus. Hist. > Ficus duquei Dugand, in Caldasia 1 4 :42. 1942 Trees 7-35 m. tall, glabrous. Twi, minutely puberulent. Stipules to 40 m or minutely puberulent, rwigs 4-13 mm. in diameter, glabrous or l mm. long, lanceolate or deltoid, glabrous Lamina 6.5-20 cm. wide X 9-30 cm. long, nate; base cordate; lateral veins (4-) 5-11 pairs, departing from the midrib at an angle from 10°-40°; basal veins departing from the midrib at an angle of 20° or less; inter- costals not prominent. Petiole 30-200 mm. long, V 2 - X A the length of the lamina. Figs 15—25 mm. in diameter, globose, minutely puberulent or glaucous, borne among the leaves,; color greenish, with purple spots; peduncle obsolete to 5 mm. long; basal bracts 4-19 mm. long, deltoid or ovate, ostiole 2—3 mm. in diameter, probably plane with the surface, but in dried material fre- quently raised, or surrounded by a more or less distinct ring of receptacular tissue. Costa Rica to State of Amapa, Brazil, in lowland forests. Bailey 3/3; Madden Reservoir, Muenscher 1 2299. This seems to be a fairly common species on the lower slopes (below 1000 m.) of the mountains behind the coast, and in the lowland forests, from Panama to the mouth of the Amazon. Humboldt found it in the coastal mountains of Venezuela and recorded it as having many buttress-like aerial roots. The species seems to be quite uniform throughout its range, though there are a few minor variations. The few specimens I have seen from Panama have the apex of the leaf very shortly terial and specimens from Trinidad have und in material from other areas, but the Some of the Colombi larger basal bracts to the fig than are foun Trinidad material differs from the Colombi crystal cells in the veins. that : (202) flora of Panama (Moraceae) 16. Ficus obtusifolia HBK. Nov. Gen. & Sp. 2:49. 1817, Urostigma involutum Liebm. in K. Dansk. Vid. Selsk. Skrivt. 5 ser 2- Urostigma bonplandianum Liebm. loc. cit. 323. 1851. . Bot. Lugd.-Bat. 3:298. 18 Irees to about 20 m. tall. Twigs 5-6 mm. in diameter, glabrous or pubescent, with thick wrinkled grey periderm. Stipules to 30 mm. long, deltoid, glabrous. Lamina 3.5-12 cm. wide X 9-25 cm. long, oblanceolate to broadly obovate, glabrous; apex rounded or bluntly acute; base cuneate; lateral veins 5-11 pairs, departing from the midrib at an angle from 20°-40°; basal veins departing from the midrib at an angle from 40°-50°; intercostals not, or scarcely, Petiole 10-30 mm. long, l/ 6 -l/ 9 the length of the lamina. Figs 15- diameter, globose, glabrous or minutely puberulent, borne among the leaves; c not noted; peduncle obsolete to 8 mm. long, puberulent; basal bracts two, mm. long, semicircular, connate, sometimes more or less split, finely pubesci orifice 3-5 mm. in diameter, flat or somewhat umbonate. Southern Mexico to northern Peru, in lowland forests. canal zone: Daytoi Standley 28729; Barro Colorado i blas: Perme, Cooper 641. This is apparently a fairly widespread species, ranging from Central Mexico to northern Guatemala, thence along the Pacific Coast through Costa Rica and Colombia to northern Peru. It ranges in altitude from near sea level to 1500— I have seen the HBK. type in Paris and the Liebmann types from Copenhagen. There can be no question but that these are conspecific. The name Ficus obtusifolia was first used by Roxburgh in the Hortus Ben- galensis in 1814. It was here a nomen nudum, not validly published, so that it does not invalidate the publication of Ficus obtusifolia, HBK. in 18 17. Roxburgh's name was not validly published until 1832. I have not seen the type of Ficus proctor-cooperi Standley. That name is included in synonymy here on the authority of Dr. Woodson. 17. Ficus costaricana (Liebm.) Miq. in Ann. Mus. Bot. Ludg.-Bat. 3:298. 1867. Urostigma costaricanum Liebm. In K. Dansk. Vid. Selsk. Skrivt. 5 ser. 2:322. 1851. Ficus kellermannii Standi, in Contrib. U. S. Nat. Herb. 20:18. 1917. Trees 12-15 m. tall. Twigs 3-5 mm. in diameter, pubescent or glabra te, developing a rather thin brownish grey periderm. Stipules 12-25 mm. long, broadly deltoid, glabrous, persistent. Lamina 3-7.5 cm. wide X 6-18 cm. long, oblanceolate or oblong-elliptic, frequently somewhat inequilateral, glabrous; apex rounded, acute, or shortly blunt acuminate; base rounded or rounded cuneate, emarginate; lateral veins 5-12 pairs, departing from the midrib at an angle from (203) ANNALS OF THE MISSOURI BOTANICAL GARDEN 20°- 30°; basal vains departing from the midrib at an angle of 50°- 60°; inter- costals prominent. Petiole 5-30 mm., %-% the length of the lamina. Figs 10- 12 mm. in diameter, depressed globose; glabrous, borne among the leaves; color reddish with darker spots; peduncle obsolete; basal bracts 5-10 mm. broadly deltoid, glabrous. Guatemala to Panama, at low or intermediate elevations. canal zone: Barro Colorado Island, Woodworth & Vestal 572, Salvoza 937, Standley 41148. chiriqui: David, Pittier 2826, 2835; Caldera, Pittier 3348. The persistent stipules and the oblate reddish receptacles with rather thin flesh are outstanding traits of this species. It also is represented by S. Hayes 838 (sine 1 Standi, in Contrib. U. S. Nat. Herb, i 13. 1917. Ficus isophlebia Standi, loc. cit. 14. 1917. Hcus jimcnesti Standi, loc. cit. 1917. Trees to 30 m. tall. Twigs 4-6 mm. in diameter with crowded internodes, glabrous, developing a rather thick reddish-brown periderm. Stipules 15-20 mm. long, narrowly deltoid, glabrous, sometimes glaucous. Lamina 3-10.5 cm. wide X 4-17 cm. long, broadly oval or ovate, glabrous; apex rounded or bluntly acute; base rounded cuneate, rounded, truncate, sometimes more or less emarginate or subcordate; lateral veins 7-10 pairs, departing from the midrib at an angle from (15°-) 25°-40°; basal veins departing from the midrib at a similar angle or up to 70°; intercostals not prominent. Petiole 20-65 mm. long, 2-3 mm. thick, %- flora of Panama (Moraceae) 165 Vs (-y 5 ) the length of the lamina. Figs 9-14 mm. in diameter, depressed globose, glabrous, borne among the leaves; color not noted; peduncle obsolete; basal bracts 10-15 mm. long, rounded deltoid, surrounding the receptacle and adherent to it for about 5 mm. British Honduras to Panama and possibly southward, in lowland forests. ?ittl°r C 282i^ T ° RO: ?Uert0 ArmuelleS ' Woodson * Schery, 812. cHiaiouf: David, Possibly more than one species is represented here, but I doubt it. The recep- tacles are so closely sessile as a rule that the growing twig beneath them develops curious persistent sockets which remain long after the fruit has fallen. 19. Ficus da vidsoniae Standi, in Field Mus. Publ. Bot. 22:15. 1940. Trees to 30 m. tall. Twigs stout, glabrous, with a thick brown periderm. Stipules about 10 mm. long, deltoid. Lamina 4-7 cm. wide X 8-15 cm. long, oblanceolate or broadly obovate, glabrous; apex rounded or obtuse; base cuneate or broadly cuneate; lateral veins 8-12, departing from the midrib at an angle of 20°- 30°; basal veins departing from the midrib at an angle of 50°- 60°; intercostals not prominent. Petiole 10-20 mm. long, i/ 8 the length of the lamina. Figs 8-10 mm. in diameter, globose, glabrous, borne among the leaves; color unknown; peduncle obsolete; basal bracts broadly rounded, deltoid, about as long as the receptacle. El Salvador to Panama, in higher-elevation forests. i Hagen 2095. cocle: La For this species I have seen only a leaf of Davidson 688. On the basis of general shape and proportions, confirmed by the structure of the lower epidermis, this seems to be fairly closely related to Ficus clusiaefolia Schott, ex Spreng. of the eastern coast of South America from Venezuela to Rio de Janiero. 13. POUROUMA Aubl. Pourouma Aubl. Hist. PI. Guian. Fr. 2:891. 1775. Puruma St.-Hil. Expos. Fam. 2:313. 1805, var. orth. Dioecious trees, frequently epiphytic and strangling when juvenile, the branches usually stout and hollow. Leaves spiral, basifixed, entire to deeply lobed; stipules fully amplexicaul and leaving a scar completely surrounding the stem. Inflores- cences repeatedly cymose, the flowers rather indefinitely glomerate at the ends of the branches. Staminate flowers with 4 nearly free tepals and 4 free stamens. Pistillate flowers with a fleshy cupular or tubular perianth, epigynous; stigma obscurely 2-lobed. Fruit a somewhat fleshy false drupe of moderate size. Perhaps 30 or more species from southern Mexico to Bolivia and Brazil. In Panama and Costa Rica the trees are known locally as guarumo, a name applied to certain other quite unrelated plants with a milky or mucilaginous sap. (205) 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN The species of Pourouma are perhaps the most ambiguous and vexatious of American Moraceae, a dubious distinction. The representation from Panama is extremely meagre and very little, if anything, is known of the variability of the species of the genus in any part of its range. The flowers apparently display little in the way of systematic variation. Consequently the species which follow are ' usually 7- to 9-lobed '...'.'. ' ' '.. ' ^ ' 4. P. oraria 1. Pourouma radula R. Ben. in Bull. Mus. Hist. Nat. Paris 28:320. 1922. Trees of moderate size, the young branches relatively slender, minutely fulvous- tomentellous to glabrate. Leaves variable, entire and ovate-elliptic to broadly 3-lobed with the median lobe divided about halfway to the petiole and scarcely constricted at the base, obtuse to broadly acute at the tip, rounded to cordate at the base, 10-25 cm. long and 6-15 cm. broad, firmly membranaceous, scabridulous above, densely and minutely fulvous- tomentellous beneath, the petiole 2-12 cm. long, rather slender, densely and minutely fulvous- tomentellous. Inflorescences about as long as the subtending petioles or somewhat shorter, densely and minutely fulvous-tomentellous. Fruits broadly ovoid, about 1.5 cm. long, minutely fulvous- tomentellous. Panama to the Guianas, in lowland forests. The leaves of this specimen coincide fairly well with those of the type specimen of P. aspera Tree, tracings of which were sent me by M. Leandri of Paris. How- ever Trecul describes the pubescence of P. aspera as consisting of "pilis albis adpressis". 2. Pourouma johnstonii Woodson, spec. nov. Arbores ut dicitur ca. 7-12 m. attingentes, ramulis crassis fistulosis dense rubiginoso-papillatis pilis hispidis interspersis. Folia latissime 5-lobata basi amplec- tante cordata supra scaberula subtus pallidiora inter venas venulasque hirtellas minute appresse arachnoideo-villosula, lobis breviter acuminatis per longetudinem ca. dimidiam basi conjunctis parte libra medii ca. 20 cm. longa 15 cm. lata, 2 exteriorum ca. 5 cm. longa et lata sinibus obtusis, petiolis ca. 20 cm. longis minute rubiginoso-papillatis pilis hispidis interspersis; stipulis anguste laceolatis ca. 8-10 cm. longis extus rubiginoso-papillatis pilis hispidis interspersis. Flores sexum (206) flora of Panama (Moraceae) 1714 (MO, HOLOTY Closely related Trees to about glabrate. Leaves at maturity deeply 3-lobed, long and broad, the lobes divided about two-t oad relocation between Gorgona and Gatun, V it tier 2286. -aria, but probably distinct as set forth in the key. . Ben. in Bull. Mus. Hist. Nat. Paris 28:320. 1922. tall, the branches stout, hollow, appressed-sericeous the (207) broadly cordate, about 10-2 lirds to the petiole, the media » above, pale and densely pa , the petiole 20-30 cm. long, c 168 ANNALS OF THE MISSOURI BOTANICAL GARDEN appressed-sericeous; stipules narrowly lanceolate, about 8-10 cm. long, densely ap- pressed-sericeous. Peduncles paired at the nodes, slightly shorter than the subtend- ing petioles, densely appressed-sericeous below, rubiginous-papillate above. Neither staminate nor pistillate flowers known from Panama. Fruits broadly compressed- ovoid, about 1.5 cm. long and 1 cm. broad, minutely puberulent. Guatemala to the Guianas, in forests of i cocle: north of El Valle de Anton, Allen 3742. darien: Cana, Williams 983. Closely related to P. palmata Poeppig & Endl., in which the leaf base is tyr lria Standi. & Cuatr. in Caldasia 7:301. 1956. Apparently trees of moderate size, the young branches stout and hollow, densely rubiginous-papillate interspersed-hispid to glabrate. Leaves at maturity deeply 7- to 9-lobed about two-thirds or more to the petiole, deeply amplectant- cordate at the base, about 25-35 cm. long and broad, the median lobe strongly contracted at the base, scaberulous above, paler and densely arachnoid-vilosulose beneath, the petiole about 15-25 cm. long, densely rubiginous-papillate sparsely interspersed- hispidulous; stipules oblong, 8-10 cm. long, densely rubiginous-pulverulent interspersed hispid. Peduncles paired at the nodes, the peduncle somewhat shorter than the subtending petiole, rubiginous-papillate sparsely interspersed-hispidulous. Staminate flowers unknown. Fruits compressed-ovoid, about 1.5 cm. long and 1 cm. broad, densely rubiginous-papillate interspersed-hispidulous. Panama and Colombia. "Panama": Hayes 348, 862. Reminiscent of P. cecropiae folia Mart, ex Miq. of the Amazon basin, but the leaf lobes are not as deeply divided nor as many as in that species. Shattuck 260 from Barro Colorado Island may represent a juvenile leaf form. 14. COUSSAPOA Aubl. Coussapoa Aubl. Hist. PL Guian. Fr. 2:955. 1775. Dioecious trees, usually epiphytic when juvenile. Leaves spiral, basifixed, entire to more or less definitely undulate or crenate; stipules fully amplexicaul, leaving a scar completely surrounding the stem. Inflorescences usually paired at the nodes, the staminate dichotomously compounded small globular heads, the pistillate superficially simple or (in C. magnifolia) obscurely compound heads. Staminate flowers with 4 (-3 ) free tepals and 2 completely fused stamens. Pistillate flowers tubular or clavate, the limb thickened and calyptriform and minutely porous much as in Cecropia, the ovary superior, with a barely exserted penicillate stigma. Fruit a scantily fleshy syncarp. About 30 species ranging from southern Mexico to Brazil and Peru. Coussapoa resembles Ficus in its frequently epiphytic germination and strangling habit, but it is less aggressive in the latter trait. The hollow branches frequently are inhabited by ants. (208) Panama (Moraceae) glabrate beneath . 3. C. brevipes 1. Coussapoa magnifolia Tree, in Ann. Sci. Nat. 3 ser. 8:98. 1847. Coussapoa nymphaeifolia Standi, in Proc. Biol. Soc. Wash. 37:50. 1924, as to specimens cited, in part, not as to type. Coussapoa cbagresiana A. D. Hawkes, in Phytologia 3:30. 1948. Trees to about 10 m. tall, the branches stout and hollow, minutely ferruginous- tomentellous to glabrate. Leaves broadly oval, broadly rounded at the tip, more or less deeply cordate at the base, 15-30 cm. long, 10-25 cm. broad, entire or indefinitely undulate, firmly membranaceous, minutely puberulent to glabrate beneath, the venation subpalmate and rather distant, with 6-8 pairs of upper secondaries, the tertiaries prominently dendroid and forming a coarse and irregular reticulum, the petioles 5-15 cm. long; stipules about 3-5 cm. long, minutely ferruginous- tomentellous. Staminate inflorescences much shorter than the sub- tending petioles, about 3- to 4-times dichotomous, the heads nearly 1 cm. broad at anthesis. Pistillate inflorescences obscurely compound, the heads 2- to 4-lobed, 4-5 cm. broad at anthesis, the peduncles simple, about 2-3 cm. long. Panama to Peru, in lowland forests. canal zone: Chagres, Hayes 354; Fort San Lorenzo, Allen 5/ 20; west of Limon Bay, Johnston 1768; Barro Colorado Island, Kenoyer 321, Woodworth & Vestal 606. Hayes 354., the type of C. cbagresiana, was cited by Standley following the description of C. nymphaeifolia, but the type of the latter from Costa Rica, Cook & Doyle 757, has foliage with more typically pinnate, crowded venation and the pistillate heads are quite sessile. The Panamanian plants seem to coincide exactly with a Pavon isotype photographed at Berlin by Dr. J. Francis Macbride. 2. Coussapoa panamensis Pittier, in Contrib. U. S. Nat. Herb. 18:226. 1917. Trees up to about 30 m. tall, the branches relatively slender, glabrous or essentially so. Leaves ovate, obtuse at the tip, obtuse or rounded, rarely truncate or obscurely cordate at the base, 10-30 cm. long, 7-15 cm. broad, subcoriaceous, cinereous beneath, the venation closely and typically pinnate with 10-20 pairs of secondaries, the tertiary venation densely parallel and forming a delicate and uni- form reticulum, the petioles 2-8 cm. long; stipules 2-6 cm. long, densely and minutely puberulent-papillate. Staminate inflorescences about equaling or much shorter than the subtending petioles, 2- to 4-times dichotomous, the heads about 5 OF THE MISSOURI BOTANICAL GARDEN . broad at anthesis. Pistillate inflorescences apparently simple, the heads about n. broad at anthesis, the peduncles 2-6 cm. long, minutely puberulent. Southern Mexico to Panama and probably extending into northern South ;rica, in lowland forests. flora of Panama (Moraceae) 171 bocas del toro: Chaguinola Valley, Dunlap 299; Rio Cricamola, Woodson, Allen & Seibert 1893, Cooper 538; Coca Cay, Von Wedel 2879; Water Valley, You We del 1733. canal zone: Fort Sherman, Standley 30924; Rio Medio, M,llvr 1718; Gatun, Hayes 414, 986, 1008; Barro Colorado Island, Standley 41170, Kenoyer 314, Woodworth i Vestal 445 ; 660. colon: Rio Fato, Pittier 3892. Panama: San Jose Island, Johnston 606, 652, 3. Coussapoa brevipes Pittier, in Contrib. U. S. Nat. Herb. 18:225. 1917. Epiphytic shrubs, the branches relatively stout, glabrate. Leaves broadly oval, obtuse to rounded at the tip and at the base, crenate, 10-17 cm. long, 6-11 cm. broad, subcoriaceous, inconspicuously arachnoid beneath to nearly glabrate, the venation rather typically and closely pinnate, with about 11 to 14 pairs of secondaries, the tertiary venation densely parallel and forming a delicate uniform reticulum, the petioles 4-6 cm. long; stipules 4-7 cm. long, minutely and densely arachnoid-puberulent. Staminate inflorescences unknown. Pistillate inf] apparently simple, subsessile, the heads about 1 cm. broad at anthesis. Panama in lowland forests. Known only from the type collection. san blas: Sperdi, near Puerto Obaldia, Pittier 4386. A puzzling species, possibly present in Colombia also. 15. CECROPIA L. 1 Loefl. Iter Hisp. 272. 1758, nom. conserv. nk and branches stout and hollow. Leaves spiral, fully amplexicaul and leaving a scar of spadicose spikes in digitate perianth tubular with a somewhat thickened porous rillate flowers: perianth tubular with thin shredding walls and a fleshy porous circumscissile operculum, the penicillate stigma barely exserted. Fruit a minute achene. Possibly in excess of 80 species of tropical America; a genus badly in need of revision as in most Moraceae, the species of which are probably not as difficult to distinguish as has been supposed. Cecropia is one of the most conspicuous and picturesque trees of moderate size in clearings and thicket areas because of its rapid growth and long-petioled digitate leaves; it is seldom if ever found in established forests. The hollow trunks and branches almost invariably are the homes of myriads of aggressive ants. It is commonly known as guarumo, with various modifications. Young seedlings of Cecropia present what may be an interesting instance of juvenile reversion. The first stem leaves of C. peltata are subtended by lateral stipules and are undivided although with more or less undulate-serrate margins; next ensue leaves with fully amplexicaul stipules and basifixed blades which are palmately divided into 3 segments as in many species of Pouroiima. Finally the excentrically peltate mature foliage is attained. (211) 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 60. Cecropia peltata FLORA OF PANAMA (Mo Cecropia bumboldtiana Kl. in Linnaea 20:530. 1847. Cecropia arachnoidea Pittier, in Contr. U. S. Nat. Herb. 18:226. 1917. Cecropia asperrima Pittier, loc. cit. 227. 1917. Trees 6-20 m. tall with stout, seldom-branching trunks, the young branches stout, hispidulous to glabrate. Mature leaves divided about midway to the center or scarcely beyond, scabridulous above, paler and densely arachnoid-villosulose to glabrate beneath, the lobes usually 9-1 1 , not contracted toward the base or scarcely so, obtuse to rounded at the tip, the petioles densely and minutely hirtellous, with a ferruginous-velutinous basal pulvinus; stipules about 6-9 cm. long, hirtellous. Staminate spadices in clusters of 12-30, 3-5 cm. long, about 4 mm. in diameter, with slender hirtellous stipes about 3-5 mm. long, the spathes 4-6 cm. long shortly before anthesis, broadly conic-oblongoid, densely white-arachnoid-villous, the common peduncle 7—10 cm. long after anthesis, slender, minutely hirtellous. Pistillate spadices in clusters of 4-6, at anthesis about 4-5 cm. long and 5 mm. in diameter, in fruit about 5-10 cm. long and 1 cm. in diameter, subsessile or with very inconspicuous stipes, the spathes as in the staminate inflorescences, the common peduncle about 7-8 cm. long at anthesis, somewhat accrescent in fruit. Southern Mexico to northern South America and in the Greater Antilles, chiefly in clearings and thickets at low elevations. canal zone: Frijoles, Piper 5837; Gamboa, Standley 28477; Matachin, Pittier 4056; Culebra, Pittier 4060; Tabernilh. Pittier - -??; Barro Colorado Island, Kenoyer 310, Seibert 56$. Panama: Sa km 154, 155, 7$, 47*, Erlmson 233, Harlow 87. 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN The leaves of this species, as in all Cecropias, vary so greatly in size and frequently so large that only a very poor impression is imparted by herbari specimens. In C. peltata mature leaves may attain a diameter of nearly one \ 2. Cecropia longipes Pittier, in Contr. U. S. Nat. Herb. 18:227. 1917. Trees to about 10 m. tall with stout seldom-branching trunks, the young branches very stout, densely gray-hispid to glabrate. Mature leaves divided about midway to the center or less, scabridulous above, paler and subarachnoid-puberulent beneath, the lobes 9-13, not contracted toward the base or scarcely so, broadly obtuse to rounded at the tip, the petioles very stout, densely gray-hirtellous, with the ferruginous velamen of the basal pulvinus more or less concealed by inter- spersed white longer hairs; stipules about 9 cm. long, ferruginous-hirtellous with interspersed longer white hairs. Staminate spadices in clusters of 50-60, 4-12 cm. long and about 2 mm. in diameter, with slender gray-hirtellous stipes about 2 cm. long, the spathes unknown, the common peduncle about 10-12 cm. long, gray- hirtellous after anthesis. Pistillate spadices in clusters of 8-12, at anthesis about 8-9 cm. long and 5 mm. in diameter, in fruit about 10-12 cm. long and 1 cm. thick, very shortly stipitate, the spathes about 5-6 cm. long, gray-hirtellous, the common peduncle 5-6 cm. long, rather irregularly but conspicuously gray- hirtellous, somewhat accrescent in fruit. Presently known only from Panama, in thickets and secondary forests at low elevations. Bailey 421. The extremely long pendulous pistillate peduncles are most impressive. The mature leaves may attain a diameter of nearly a meter. 3. Cecropia eximia Cuatr. in Rev. Acad. Colomb. Cienc. 6:287. 1945. Trees to about 10 m. tall with stout seldom-branching trunks, the young branches very stout, shortly hispidulous to glabrate. Mature leaves divided nearly to the center, smooth and when juvenile sparsely arachnoid-villous above, paler and minutely canescent beneath, the lobes about 9, conspicuously contracted toward the base, broadly acute or obtuse at the tip, the petioles very stout, densely arachnoid-villous with the basal pulvinus ferruginous-velutinous; stipules about 6 cm. long, ferruginous-velutinous with inspersed longer canescent hairs. Stami- nate inflorescences of about 10 spadices 12-15 cm. long with slender basal stipes about 1.5 cm. long, the common peduncle about 15-20 cm. long, arachnoid- villosulose to glabrate. Pistillate spadices in clusters of about 5, at anthesis 8-10 cm. long and about 8 mm. thick, with enlarged subcalyculate pruinose basal stipes about 1 cm. long, the spathes oblongoid, long-apiculate, about 10-12 cm. long and 2-3 cm. thick shortly before anthesis, gray-hirtellous, the common peduncle about 8-10 cm. long, gray-hirtellous. (214) flora of Panama (Moraceae) 175 Costa Rica to Colombia, in lowland thickets and secondary forests. bocas del toro: Almirante Region, Slater 48. canal zone: area west of Limon Bay, etc., Johnston 1758. At one time I suspected that an earlier name for this species might be C insignis Liebm., described very inadequately from San Juan de Nicaragua. In the her- barium at Copenhagen, which is the chief depository of Oersted's collections, there is no specimen labeled as C. insignis. However, 5 sheets of Oersted 5861, one of which is noted as having been collected at San Juan de Nicaragua, are referable to C. peltata L. in the broad sense employed here. The description of the staminate inflorescence is taken from a Costa Rican specimen collected in the region of Limon. The Panamanian specimens are pistillate, as in the Colombian type. Three leaf specimens from the Canal Zone and deposited in the U. S. National Herbarium, possibly represent juvenile forms of this species: Piper 5847, 584.0, and Killip 1 21 7 3. It really is something of a crime, or at least a misdemeanor, to collect leaves of Cecropia unaccompanied by 4. Cecropia maxonii Pittier, in Contrib. U. S. Nat. Herb. Trees 10-12 m. tall t very stout, minutely cin< center, scabridulous and irregularly pellucid-maculate above, paler and inconspicu- ously hirtellous beneath, the lobes 9-11, conspicuously contracted toward the base, broadly obtuse to rounded at the tip, the petioles rather stout, densely and minutely puberulent, with the basal pulvinus densely ferruginous-velutinous. Staminate spadices in clusters of 12-15, up to about 10 cm. long and 4 mm. thick, with slender minutely puberulent-papillate stipes about 5-10 mm. long, the spathes oblongoid, shortly apiculate, about 12-14 cm. long shortly before anthesis, minutely subarachnoid-puberulent, the common peduncle about 5-7 cm. long shortly before anthesis, minutely pilosulose. Pistillate spadices about 1 cm. long in mature fruit (our specimen fragmentary) , the common peduncle about 6-7 cm. long. Known only from the type locality, in highland secondary forest. chiriqui: Boquete, Maxon 5132, Davidson 862. The maculation of the leaves is most peculiar and possibly is pathological, but I have seen nothing comparable to it in other specimens of Cecropia. It may be similar to the cystolith maculations of Urticaceae. 5. Cecropia obtusifolia Bertol. Fl. Guat. 39. 1840. Cecropia schiedeana Kl. in Linnaea 20:53 1. 1847. Cecropia bicolor Kl. loc. cit. 1847. Cecropia digitata Ten. ex Miq. in Mart. Fl. Bras. 4 1 :149. 18 53. % Schott, ex Miq. loc. cit. 148. 1853. 'ogeleri Burret, in Notizbl. 9:49. 1924. nexicana var. macrostachya Donn. Sm. i (215) ANNALS OF THE MISSOURI BOTANICAL GARDEN Trees to about 10 m. tall or more, with stout seldom- branching trunks, the young branches stout, ferruginous-hirtellous to glabrate. Mature leaves divided nearly to the center, scabridulous above, paler and minutely cinereous beneath, the lobes usually 9-13, conspicuously contracted toward the base, rounded to shortly acuminate at the tip, the petioles rather slender, minutely ferruginous- hirtellous, with the basal pulvinus densely ferruginous-velutinous. Staminate spadices usually in clusters of 3, 12-16 cm. long and about 4 mm. thick, subsessile or with slender puberulent stipes to about 5 mm. long, the spathes 12-20 cm. long shortly before anthesis, narrowly oblongoid-fusiform and frequently somewhat falcate, rather sparsely arachnoid- villosulose, the common peduncle about 5-10 cm. long, relatively slender. Pistillate spadices in clusters of 3-4, 25-30 cm. long and about 5 mm. thick at anthesis, somewhat accrescent in fruit, sessile or with a very indefinite stipe, the spathes about 16-20 cm. long shortly before anthesis, flora of Panama (Moraceae) 177 narrowly oblongoid-fusiform and frequently somewhat falcate, sparsely arachnoid- villosulose to glabrate, the common peduncle 8-14 cm. long, nearly glabrous. Southern Mexico to northern South America, in lowland clearings, thickets and secondary forests. bocas del toro: Water Valley, Von Wedel 771, 155Q. canal zone: Las Cruces Trail, Hunter & Allen 462; Gatun, Standley 27304; Masambi, Pittier 2673; Darien Station, Macbride 2697; Summit, Harvey 5282; Moj M /qc ; Taber- Uer 3826; Barro Colorado Island, Wetmore & Abbe 128, 204, Wood-worth d Vestal 644, Kenoyer 309. darien: Pinogana, Allen 43 II. veraguas: Santa Fe to Rio Santa Maria, Allen 4422. 16. DORSTENIA L. Dorstenia L. Spec. PI. 121. 1753. Kosaria Forsk. Fl. Egypt.-Arab. 164. 1775. Sychinium Desv. in Mem. Soc. Linn. Paris 4:216. 1826. Succulent or subsucculent herbs. Leaves stipules lateral. Flowers common discoid or radiate head, the obsolete perianths 1 with the receptacle. Stamina te flowers with 1-3 minute stamens, the filaments inflexed before anthesis. Pistillate flowers with the ovary included and the excentric stigma shortly 2-lobed. Receptacles accrescent and fleshy in fruit, at length extruding the minute achenes. Perhaps 75 or more very diverse species of the tropics of both hemispheres. In Panama only one species is presently known. Dorstenia contrajerva L. Sp. Pi. 121. 1753. Dorstenia houstoni L. loc. cit. ed. 2. 176. 1762. Dorstenia contrajerva var. houstoni (L.) E. Bur. in DC. Prodr. 17:259. 1873. Dorstenia contrajerva ssp. tenuiloba S. F. Blake, in Contr. U. S. Nat. Herb. 24:2. 1922. Dorstenia contrajerva var. tenuiloba (S. F. Blake) Standi. & Steyerm. in Field Mus. Bot. Succulent or subsucculent acaulescent or subacaulescent herbs. Leaves densely crowded, long-petioled, of relatively small or moderate size, extremely variable in dimensions and outline, commonly deeply pinnatifid, basifixed, usually scabridulous or inconspicuously puberulent. Receptacles on rather long slender peduncles, centrally peltate, variously radiate to nearly quadrangular, accrescent in fruit to as much as 5 cm. in diameter. Southern Mexico to northern South America and in the Antilles, in moist thickets and forests from sea level to about 2,000 m. Widely known as contra- yerba in Central America, which is a reflection of its use as a febrifuge. bocas del toro: Water Valley, Von Wedel 1530. canal zone: Las Cruces, Seibert 574; Ancon, Pittier 2748. cocie: Ccrro V.ille Chiquitn, Seibert 502. darien: Cana, Williams 938. Panama: Isla Taboga, Woodson, Allen & Seibert I JOS; Juan Diaz, Maxon & Harvey 6716. Far more frequent than the rather few citations imply. (217) 178 ANNALS OF THE MISSOURI BOTANICAL GARDEN flora of Panama (Urticaceae) URTICACEAE By ELLSWORTH P. KILLIP Monoecious or dioecious herbs, shrubs, or trees, sometimes armed with stinging hairs and spines, usually bearing cystoliths; leaves simple, alternate or opposite (those of adjacent nodes or of a pair often unequal and dissimilar), entire, toothed, or rarely lobed, stipulate; flowers unisexual, small, greenish or greenish white, in unisexual or androgynous clusters, the perianth 2-5 -lobed or parted or sometimes wanting; stamens as many as the perianth lobes and opposite them; ovary superior, 1 -celled; style undivided, the stigma penicillate-capitate or filiform; fruit an achene, sometimes enclosed in the persistent perianth. About 40 genera of the tropics and temperate zones of both hemispheres, including the stinging nettles, picas and ortigas. Here also may be included the Asian hemp and hashish plant, Cannabis sativa L., known as marijuana in Latin America, an erect herb with watery sap and digitately 5- to 9-divided leaves and inconspicuous dioecious flowers. Despite some attempts of the authorities, mari- juana is grown more or less openly and used as a constituent of narcotic cigarettes. Pilea Lindl. Coll. Bot. pi. 4. 1821, nom. conserv. Adicea Raf. Anal. Nat. 179. 1815, nom. rejic. often succulent, without stinging hairs; stipules intra-axillary, deciduous sistent; leaves opposite, usually petiolate, toothed or entire, those of a pai or markedly unequal, similar in shape or very dissimilar; flowers in unise androgynous clusters, these solitary or forming cymes or panicles; st: flowers 4 (rarely 2 or 3) -parted, the pistillate normally 3 -parted, the segment usually larger than the lateral ones; stigma sessile, penicillate; compressed, orbicular or ovate. 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN d. Leaves all distin ee. Smaller leaves oblong; stem drying dark; plant usually pubes- ' dedduous. eaVeS ee. Larger leaves ovate or elliptic, 4-7 cm. wide, their petiol s ff. Stem glabrous; leaves rhombic-ovate, generally not more than 2 cm. long, flat 11. P. auric benes minute, scarcely more than 0.5 mm. long, only slightly (220) flora of Panama (Urticaceae) 181 1. Pilea microphylla (L.) Liebm. in Dansk. Vid. Selsk. Skrift. V. 2:296. 1851. Parietaria microphylla L. Syst. ed. 10. 1308. 1759. Pilea muscosa Lindl. Coll. Bot. pi. 4. 1821. Glabrous, succulent, much-branched herb, up to 30 cm. high, variable in size and habit; leaves usually crowded throughout the length of the branches, entire, succulent, those of a pair unequal, the larger prevailingly obovate, up to 1 cm. long, petiolate, obtuse or subacute, tapering at base, the smaller orbicular or obovate-orbicular, rarely more than 3 mm. long, sessile or short-petioled; cystoliths linear, transverse across the leaf-blade; plants monoecious, rarely dioecious, the flower clusters very small, sessile or nearly so; achenes ovate, about 0.3 mm. long. Throughout the American tropics, up to 2500 m. altitude, occurring in a variety of habitats, as in crevices of stone walls or pavements, on mossy roots or rocks, or on the forest floor; often cultivated as a border plant or in pots. bocas del toro: Changuinola Valley, Dunlap 358, 387; Chiriqui Lagoon, von Wedel 24.87. canal zone: Balboa, Standley 28554, 29384 (cultivated; distributed as P. I ; Ancdn, Piper 6017; Fort Sherman, Standley 3094$; ^"o Colorado Island, Killip 40003; Shattuck 196 (cited as P. serpyllacea) ; Chagres, Fendler 259. colon: Porto Bello, Pittier 2447. Panama: near Panama, Standley 26888, 29807, Heriberto 67; between Las Sabanas and Matias Hernandez, Standley 31834; Chepo, Pittier 4460. A related species, P. herniarioides (Sw.) Lindl., may occur in Panama. It is distinguished by rhombic-orbicular leaves, the upper two or three pairs rosulate at the tip of the branches, the flower-clusters sessile within these rosettes. Pilea serpyllacea (H.B.K.) Liebm., also a related species, has been reported from Panama, but the records were apparently based on robust P. microphylla. 2. Pilea parietaria (L.) Blume, Mus. Bot. Lugd. 2:48. 1856. Pilea ciliaris Wedd. in Ann. Sci. Nat. III. 18:209. 1852. Plant herbaceous or suffrutescent; leaves of a pair (especially the petioles) somewhat unequal, the blades thin, elliptic or ovate-lanceolate, 2-8 cm. long, 1-3 cm. wide, rarely longer, acuminate at apex, rounded or subacute, sometimes sub- auriculate at base, 3 -nerved, entire, usually cilia te, glabrous or sparingly strigillose with pellucid hairs, usually punctate beneath, the cystoliths linear or fusiform, usually inconspicuous; plants monoecious, the panicles 1 or 2 to an axil, shorter than the leaves, pedunculate, androgynous, or the upper almost wholly staminate and the lower pistillate; achenes minute, barely 0.5 mm. long. Guatemala to Panama, on shaded banks and in moist forests, at altitudes between 1000 and 2000 m.; also in the West Indes. chiriqui: Quebrada Velo, alt. 1800 m., Woodson 6 Schery 242; Chiquero, Boquete District, Davidson 549. 3. Pilea imparifolia Wedd. in Ann. Sci. Nat. III. Bot. 18:212. 1852. Decumbent or subrepent herb with numerous erect branches, essentially gla- 82 ANNALS OF THE MISSOURI BOTANICAL GARDEN brous throughout; leaves of a node unequal and dissimilar, the larger rhombic-ovate to elliptic-oblong or obovate, 1-6 cm. long, 0.8-2 cm. wide, narrowed at apex, acute or attenuate at base, sessile or short-petioled, crenate- serrate above middle, the smaller leaves obovate-orbicular or orbicular-reniform, 0.6-1.5 cm. long, strongly asymmetrical, subentire; cystoliths linear on upper leaf surface, unequally puncti- form beneath; plants dioecious, the cymes sessile or subsessile, few-flowered; achenes about 1 mm. long. Panama to northern Peru, eastward to the Guianas and Amazonian Brazil; usually in dense forests. cocle: trail to Las Minas, north of El Valle de Anton, alt. 1000 m., climbing on trees and boulders, Allen 2470; El Valle de Anton, 1000 m., on tree trunk, Alston 8791. This species is common in the forests of the Choco, Colombia, where it shows much variation in shape and size of the leaves. Doubtless it occurs in Darien. 4. Pilea cornmanae Killip, in Jour. Washington Acad. Sci. 15:292. 1925. Plant erect, up to 60 cm. high, glabrous throughout except the leaves, the stem as well as the foliage densely covered with elevated linear cystoliths; stipules orbicular or ovate, persistent; leaves coarsely serrate, dark green above, paler beneath, thin, sometimes sparingly strigillose above with hyaline hairs, the leaves of a node unequal and somewhat dissimilar, the larger ovate-lanceolate, 2-7 cm. long, 1.5-3 cm. wide, acuminate at apex, acute and often oblique at base, the petioles up to 4 cm. long, the smaller leaves broadly ovate or suborbicular, 1.5-2 cm. long, 1-1.5 cm. wide, acute; plants monoecious or dioecious, the flower-clusters unisexual, pedunculate; stamina te heads globose, about 1 cm. wide, the segments long-caudate; pistillate flowers loosely clustered in subglobose heads; achenes broadly ovate or suborbicular, about 1 mm. long. In wet forests of western Panama and San Jose Province, Costa Rica, between 1300 and 2000 m. alt. chiriqui: Holcomb's Trail, along Rio Caldera, Killip & Cornman (Killip 3543, type) ; Cerro Horqueta, v on Hagen 2? von Hagen 20I0. 5 . Pilea trian aeana Wedd. in DC. Prodr. 1 6 1 : 1 1 1 . 1869. Stem repent below, at length erect, up 1 puberulent or strigillose above; leaves of £ upper very unequal, oblong or oblong-lanc< at base, petiolate, crenate-serrate, glabrous, or sometimes pilose on the nerves beneath, the larger leaves 7-15 cm. long, 2-6 cm. wide, the smaller ones 1.5-4 cm. long, 0.5-1.3 cm. wide; plants dioecious; staminate flowers in dense globose heads which are solitary or form a branched cyme, the tips of the segments about 1.5 mm. long; pistillate flowers in lax, pedunculate panicles, the achenes 1-1.5 mm. long. Panama and adjacent parts of Colombia, up to 1000 m. alt. 1. alt., Allen 239$; flora of Panama (Urticaceae) 183 6. Pile a chiriquina Killip, in Jour. Washington Acad. Sci. 15:291. 1925. Fleshy herb, up to 1 m. high, slightly suffrutescent toward base, glabrous throughout; leaves dark green above, paler, sparingly punctate beneath, crenate- serrate, triplinerved, those of a pair unequal and dissimilar, the larger oblanceolate, 3-9 cm. long, 1-2 cm. wide, caudate-acuminate, subauricular at base, suboblique, the petioles up to 5 mm. long, the smaller leaves ovate or ovate-lanceolate, 0.7-3 cm. long, 0.5-1.5 cm. wide, acute, obliquely subcordate, sessile or subsessile; cystoliths minute, linear; plants dioecious, the flowers in compact peduncled cymes, the staminate perianth segments with filiform tips, the pistillate cymes smaller, the achenes broadly ovate, about 1 mm. long. Panama; in wet highland forests, between 1400 and 2100 m. alt. bocas del toro: Robalo Trail, northern slopes of Cerro Horqueta, Allen 4014, 4915 (only $ collection known), chiriqui: upper Rio Caldera, Killip 3546 (type); Bajo Chorro, Woodson & Schery 668; slopes of Cerro Horqueta, Allen 4825. 7. Pilea donnell-smithiana Killip, in Jour. Washington Acad. Sci. 15:292. Erect herb, about 1 m. high, glabrous throughout; leaves ovate or elliptic, crenate-serrate nearly to base, rounded or subauriculate and often oblique at base, those of a node very unequal, the larger 10-20 cm. long, 4-7 cm. wide, caudate- acuminate, the petioles up to 2.5 cm. long, the smaller leaves up to 3 cm. long and 1.5 cm. wide, their petioles up to 5 mm. long; plants dioecious, the staminate flowers in small clusters in a small, few-branched panicle, the pistillate in few- flowered, short-peduncled, subglobose heads; achenes about 2 mm. long. Costa Rica and adjacent parts of Panama; in wet forests at 1300 to 1700 m. alt. A related species, P. costaricensis Donn. Sm., may occur in western Panama, material from Bajo Chorro, Chiriqui (Davidson 56 and 717), with very young inflorescence, perhaps being referable to it. 8. Pilea ptericlada Donn. Sm. in Bot. Gaz. 31:121. 1901. Usually a stout herb, somewhat suffrutescent below, erect, up to 30 cm. high, glabrous throughout or fuscous-pubescent on the petioles and leaf -nerves; stipules orbicular-ovate, persistent; leaves of a node similar and subequal, oblong-elliptic or somewhat oblanceolate, 5-20 cm. long, 3-7 cm. wide, acuminate, cuneate-attenuate at base and sometimes long-decurrent on the petiole, coarsely crenate-serrate above middle, triplinerved, the petioles up to 1 cm. long; cystoliths mostly punctiform; plants dioecious, the inflorescence cymose-paniculate, borne in the upper axils, shorter than the leaves; staminate flowers long-pediceled, whitish, the lobes short- mucronate; pistillate flowers sessile, the achenes ovate, 1.5-2 mm. long, suboblique. Costa Rica and Panama. 184 ANNALS OF THE MISSOURI BOTANICAL GARDEN bocas del toro: Fish Creek, vicinity of Chiriqui Lagoon, von Wedel 2203, 2237, 2318. darien: Cana, alt. 750 m., R. S. Williams 8lO. canal zone: between Peluca Hydrographic Station and Quebrada Peluca, along Rio Boqueron, Steyermark d Allen 17252 (plant smaller in every way). 9. Pilea gracilipes Killip, in Jour. Washington Acad. Sci. 15:294. 1925. Plants glabrous throughout; stem repent at base, the branches erect, up to 40 cm. high; leaves of a node similar in shape and subequal (or the smaller about three-quarters as long as the larger), lanceolate or ovate-lanceolate, 1-8 cm. long, up to 3 cm. wide, acuminate at apex, acute or somewhat rounded at base, crenate- serrate, the cystoliths linear, faint; plants monoecious or dioecious, the staminate and pistillate inflorescences often borne in the same axil, their peduncles very slender; staminate flowers in globose heads, 5-7 mm. wide, the pistillate in loose glomerules forming an interrupted spike, or racemose-paniculate; achenes lance- ovate, 1-1.5 mm. long. Guatemala to Panama; in wet mountain-forests, 1500 to 4000 m. alt. chiriqui: Cerro Horqueta, 1700-2100 m., Pittier 3230, 5426 (type); Volcan de Chiriqui, Woodson, Allen d Seibert 894, Woodson d Schery 449, Davidson 1 027; Monte Lirw, Seibert 293; Bajo Chorro, Woodson d Schery 659; Quebrada Velo, Woodson d Schery 243, 269; between Casita Alta and Cerro Copete, Woodson d Schery 347. 10. Pilea rugosissima Killip, in Proc. Biol. Soc. Washington 52:28. 1939. Stem 30-100 cm. high, densely appressed-strigose; stipules oblong, persistent; leaves ovate or ovate-lanceolate, 1.5-6 cm. long, 1.5-3 cm. wide, acuminate at apex, rounded or cordulate at base, sharply serrate, strongly rugose, appressed- strigose between the veins of the upper surface, appressed-setose on the nerves and globose, peduncled head, the perianth about 5 mm. long, including prominent Panama. chiriqui: Bajo Chorro, Boquete District, about 1800 m. alt., Davidson 335 (type). 11. Pilea auriculata Liebm. in Dansk. Vid. Selsk. Skrift. V. 2:299. 1851. Stem repent at base, with many slender, erect or decumbent branches; stipules ovate, persistent; leaves rotund-rhombic, 8-20 (rarely to 50) mm. long and wide, cuneate at base, coarsely crenate, strigillose with a few hyaline hairs above, the leaves of a node similar and subequal but one usually with a shorter petiole; plants usually dioecious, the flowers in small clusters up to 1 cm. wide, their peduncles generally exceeding the adjacent petioles; staminate flowers nearly 4 mm. long, including appendages about 1.2 mm. long; pistillate flowers with 1 of the segments larger than the others, auriculate; achenes broadly ovate, nearly 2 mm. long. Southern Mexico, Guatemala, Costa Rica (where common), and western Pan- ama; wet forests from 1500 to 3000 m. alt. 1 934> Woodson d Schery 346, of Panama (Urticaceae) 185 (Sw.) Wedd. in Ann. Sci. Nat. III. 18:255. 1852. isk. Vet. Akad. 8 :63, pi. I, f. 2. 1787. Stem repent or trailing, rooting at most of the nodes, villosulous; stipules per- sistent; leaves almost orbicular, 5-12 mm. wide, crenate, usually strigillose with stiff, hyaline hairs on both surfaces, the cystoliths inconspicuous; plants dioecious or occasionally monoecious, the staminate flowers usually clustered in the uppermost axils, the pistillate in dense, short-peduncled cymes; achenes barely 0.4 mm. long. West Indies to Venezuela and Amazonian Peru; sometimes cultivated. den, Standley 28555, 41IQI. i Denkschr. Akad. Wiss. Math. Naturw. (Wien) Slender, erect annual, 10-30 cm. high, the stem simple, glabrous; stipules very small, soon deciduous; leaves rhombic-elliptic or ovate, 1-4 cm. long, 0.8-2.5 cm. wide (extremes up to 6 X 4 cm.), acute at apex, cuneate at base, serrate except toward the base, thin-membranous, glabrous, or sparsely hyaline-strigose above; plants monoecious, the inflorescences androgynous, cymose-paniculate, shorter than the petioles, borne at nearly every node, the staminate flowers relatively few; achenes suborbicular, not more than 0.5 mm. long. 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN Widely distributed in the American tropics, from Mexico and the Lesser Antilles to Chile, Argentina, and Brazil, up to 1500 m. alt. bocas del toro: Changuinola Valley, in garden, Dunlap 399. canal zone: Las Cascadas Plantation, near Summit, Standley 25764. This species has been confused with P. pumila Gray, of the United States; the plant from Santiago de Veraguas, cited as P. pumila by Seeman (Bot. Voy. Herald 194), is undoubtedly P. hyalina. 14. Pilea pallida Killip, in Jour. Washington Acad. Sci. 15:295. 1925. Plant erect, 30 cm. or more high, essentially glabrous throughout; leaves elliptic-lanceolate, 8-20 cm. long, 3-7 cm. wide, caudate-acuminate, acute at base, sometimes suboblique, serrate, densely covered with fusiform and punctiform cystoliths, slightly silvery-lustrous and punctate beneath, the petioles up to 7 cm. long; plants dioecious; stamina te flowers densely congested in globose, sessile clusters, the perianth 1.5-2 mm. long including tips about half as long; pistillate flowers in much-branched cymes, shorter than the petioles, the achenes about 0.6 mm. long, black, minutely papillose. Panama and Costa Rica; in forests, 1000-1400 m. alt. bocas del toro: Sibubi Falls, Sixaola Valley, Rowlee & Rowlee 376 (type), cocle: El Valle de Anton, 1000 m. alt., Alston 8797. 15. Pilea involucrata (Sims) Urban, Symb. Antill. 1:298. 1899. Urtica involucrata Sims, Bot. Mag. 51: pi. 248 1. 1824. Pilea chrysosplenioides Wedd- in Ann. Sci. Nat. III. 18:231. 1852. Pilea pubescent var. involucrata Wedd. in DC. Prodr. 16 1 :153. 1869. Stem repent at base, the branches ascending, up to 30 cm. high, appressed- hirsute above; leaves massed at end of branches, obovate or rotund-ovate, up to 5 cm. long and 3.5 cm. wide, rounded at apex, rounded or subauriculate at base, short-petioled, finely crenate, ciliate, strigillose with appressed hyaline hairs (rarely glabrate) above, sparingly to densely pubescent beneath, especially on the nerves, the cystoliths on the upper surface linear or fusiform and confined to the margin; plants monoecious or dioecious, the cymes generally unisexual, sessile or subsessile, shorter than the leaves; achenes minute, less than 0.5 mm. long. Panama and the West Indies to Colombia and Venezuela, usually below 1000 m. alt.; cultivated in the United States. canal zone: Empire, Crawford 574- Salamanca Hydrographic Station, Rio Pequeni, Woodson, Allen & Seibert 1600. 16. Pilea acuminata Liebm. in Dansk. Vid. Selsk. Skrift. V. 2:302. 1851. Stem subrepent at base, the branches erect, 20-40 cm. high, glabrous, or sparingly pubescent in upper part; leaves ovate-lanceolate, 4-10 cm. long, 2-3 cm. wide (extremes to 15 X 6 cm.), acuminate, cordate or somewhat narrowed at base, coarsely serrate, sometimes bullate, usually strigillose with a few hyaline hairs above, hirsutulose on the nerves beneath; plants usually dioecious, the cymes (226) flora of Panama (Urticaceae) 187 diffusely branched, with peduncles up to 6 cm. long; staminate flowers with filiform appendages; achenes minute, barely 0.5 mm. long. Mexico to Colombia; in wet shady places, between 600 and 1000 m. alt. cocle: El Valle de Anton, Allen 1737, Alston 8792. 17. Pilea pubescens Liebm. in Dansk. Vid. Selsk. Skrift. V. 2:302. 1851. Stem repent, at length erect and usually with several erect or ascending branches, brownish-pubescent; leaves usually massed at the end of the stem or branches, wanting or much reduced below, broadly ovate, up to 7 cm. long and 6 cm. wide, petiolate, usually coarsely crenate-serrate, often ciliate, sparingly strigillose with hyaline hairs or rarely glabrous above, hirsutulous beneath on the nerves and veins, the cystoliths linear and fusiform above, obscure beneath; plants monoecious or occasionally dioecious, the flowers in dense globose clusters forming a panicle up to 7 cm. long, slender-peduncled, the panicles usually androgynous Pilea pubescens evidently has a wide distribution in the lowland tropics although doubtless much material has been wrongly referred to it. Direct comparison has been made between the Panama specimens here cited and the type, collected in southeastern Brazil by Lund and generously lent the author by the Botaniske Museum, Copenhagen, and there is perfect agreement. ocle: El Valle de Anton, 2. BOEHMERIA Jacq. Boehmeria Jacq. Enum. Pi. Carib. 9. 1760; Stirp. Amer. 246, pi. 157. 1763. Trees, shrubs, or perennial herbs, unarmed and without stinging hairs; leaves opposite or alternate, toothed, 3 -nerved, those of the adjacent nodes sometimes unequal and dissimilar; plants monoecious or dioecious, the flowers in globose, usually unisexual clusters in the axils of the leaves or forming a spike; staminate flowers 3 or 4 (rarely 5) -parted; pistillate flowers tubular, contracted at the throat, 2-4-toothed or entire, the stigma filiform, the achene enclosed in the persistent, 5 enlarged perianth. 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. Boehmeria cylindrica (L.) Sw. Prodr. 34. 1788. Urtica cylindrica L. Sp. PL 984. 1753. Erect herb up to 1.5 m. high, the stem simple or branched; leaves opposite on the stem, alternate on the branches, narrowly lanceolate, ovate-lanceolate, or or subtruncate at base, coarsely dentate or serrate-dentate, glabrous, or puberulent beneath on the nerves; flower-clusters sessile in an erect or ascending spike which often is leafy toward the tip; plants monoecious; staminate perianth 4-lobed; pistillate perianth rotund-obovate, 1.5-2 mm. long and broad, 4-toothed, com- pressed, nearly sessile. Eastern Canada; eastern and southern United States; Mexico to Panama; West Indies; southeastern Brazil. 2. Boehmeria aspera Wedd. in Arch. Mus. Paris 9:349. p. II, f. 24-28. 1856. Shrub or suffrutescent herb, 1-3 m. high; leaves alternate, those of the adjacent nodes very unequal and somewhat dissimilar, crenate-serrate, coriaceous, strongly bullate, smooth or hispid above, densely cano-hirsute beneath, the larger lanceolate, 4-15 cm. long, 1-4 cm. wide, long-acuminate, the smaller leaves ovate, 1-5 cm. long, 1-2.5 cm. wide, acute; plants monoecious, the flower- clusters predominantly pistillate; staminate perianth 4-lobed; pistillate perianth about 1 mm. long, sericeo- pubescent at throat, bidentate. Costa Rica to Peru. chiriqui: vicinity of Bajo Mona and Quebrada Chiquero, alt. 1500 m., Woodson & Scbery 560. 3. Boehmeria ulmifolia Wedd. in Ann. Sci. Nat. IV. 1:202. 1854. Boehmeria jallax var. ulmifolia Wedd. in DC. Prodr. 16 1 :198. 1869. Shrub or small tree up to 6 m. high, with slender, more or less pubescent branches; leaves serrate, thin, appressed-hispidulous above, pilosulous or puberulent beneath, those of adjacent nodes strongly dimorphic and unequal; larger leaves elliptic-ovate, 5-15 cm. long, 1.5-5 cm. wide, cuspidate-acuminate, acute or rounded at base, somewhat oblique, with slender petioles 5-20 mm. long, the smaller leaves (often soon deciduous) orbicular or subreniform, 5-10 mm. long and broad, usually emarginate at base, sessile; plants monoecious or dioecious, the flower-clusters axillary, generally unisexual; staminate perianth 4-lobed; pistillate perianth about 2 mm. long, stipitate, very slightly compressed, bidentate. Southern Mexico to Panama, from about 1000 to 2000 m. alt., ascending higher in western Panama. bocas del toro: Robalo Trail, northern slopes of Cerro Horqueta, alt. 1800-2100 m., Allen 4gi3 (tentatively referred here), chiriqui: upper Chiriqui Viejo va'i of Monte Lirio, alt. 1300-1900 m., Seibert 183; Potrero Muleto to summit, Volcan de Chiriqui, alt. 3500-4000 m., Woodson & Schery 465. flora of Panama (Urttcaceae) 189 4. Boehmeria cuspidata Wedd. in Arch. Mus. Paris 9:345. 1856. Boehmeria ramiftora var. cuspidata Wedd. in DC. Prodr. 16 X :\97. 1869. Slender shrub or suffrutescent herb, up to 3 m. high, the stem appressed- pubescent; leaves of adjacent nodes similar but unequal, cuspidate-acuminate, coarsely serrate, entire toward the cuneate or slightly rounded, suboblique base, appressed-hispidulous above, sparingly pilosulous beneath, the larger leaves 6-15 cm. long, 2-5 cm. wide (smaller on the branches), the petioles slender, up to 5 cm. long, the smaller leaves 3-5 cm. long, 1.5-2.5 cm. wide, the petioles up to 1.5 cm. long; plant monoecious (or dioecious?), the flower- clusters axillary; staminate perianth 3-lobed; pistillate perianth elliptic, about 1.5 mm. long, narrowly margined, bidentate. Southern Mexico to Panama. Panama: hills above Campana, alt. 600-800 m., Allen 1310. 5. Boehmeria pavonii Wedd. in Ann. Sci. Nat. IV. 1:202. 1854. Shrub or small tree, 2-6 m. high, the branches finely appressed-pubescent; . 64. Boehmeria ulmifolia 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN leaves alternate (those of the adjacent nodes similar or nearly so though quite unequal) , finely serrulate or sometimes subentire, subcoriaceous, finely appressed- hispidulous above, usually appressed sericeo-pubescent on the nerves and veins beneath and puberulent in the areoles, the larger leaves oblong-lanceolate or elliptic- 4-16 cm. long, 1.5-4.5 cm. wide, attenuate-acuminate, the petioles i. long, the smaller leaves similar though sometimes ovate, subsessile, and cute; plants dioecious, rarely monoecious, the clusters generally unisexual; e perianth 4-lobed; pistillate perianth attenuate at base, narrowly margined. emala and Panama; Peru and Bolivia. QUI: EI Boquete, alt. Davidson 830. 3. FLEURYA Gaud. . Voy. Uran. Bot. 497. 1826. lally with stinging hairs; leaves alternate, petiolal Annual herbs, clusters in large panicles; staminate perianth 4-5 -parted, with 4-5 stamens- pistillate perianth with 4 imbricated segments, the stigma papillose, at length hooked, the achenes oblique, compressed. flora of Panama (Urticaceae) 1. Fleurya aestuans (L.) Gaud. Voy. Uran. Bot. 497. 1826. Urtica aestuans L. Sp. PI. ed. 2. 1397. 1762. Plant generally terrestrial, erect, simple or few-branched, up to 2 m. high, stinging hairs usually present, the stem sometimes glandular; leaves broadly o rarely ovate-lanceolate, up to 18 cm. long and 12 cm. wide, acu( usually rounded at base, coarsely and sharply dentate, thin-membranous, the petioles up to 12 cm. long; panicles often much longer than the petioles; achenes 1-1.5 mm. long. Widely distributed in the tropics at low elevations; often occurring in waste bocas del toro: vicinity of Chiriqui Lagoon, von Wedel 1269, 2848. canal zone: San Pablo, Pittier 4.063; Ancon, Pittier 2 75 l; Balboa, Standley 25286, 25601, 2698 I; Gamboa, Standley 28305. chiriqui': Puerto Armuelles, Woodson & Schery 849. darien: Boca de Cupe, Allen 873. Panama: Bella Vista, Kill Corozal road, Standley 26/65; Tumba Muerto road, Standley 29707; Las Sabanas, Paul 32; Chepo, Pittier 4767. veraguas: Isla de Coiba, Mendez 128. 4. URERA Gaud. Urera Gaud. Voy. Uran. Bot. 496. 1826. Trees or shrubs, rarely scandent, usually with stinging hairs or stout prickles; stipules free, or more or less connate; leaves alternate, petiolate, 3- or 5-nerved at base, penninerved above, the cystoliths punctiform, linear, or wanting; plants usually dioecious, the flowers in axillary, dichotomous or irregularly branched cymes or panicles; stamina te flowers with a 4-5 -parted perianth and 4-5 stamens and a rudimentary ovary; pistillate flowers with 4 equal or unequal segments, the stigma penicillate-capitate, persistent; achenes straight or oblique, at least partially sur- rounded by the fleshy, enlarged perianth. aa. Leaves n"" Tobed, 'denTce, cremate, 'or "ubentTr^ b. Achenes more • ne or sinuate- 1. Urera laciniata (Goudot) Wedd. in Ann. Sci. Nat. III. 18:203. 1852. Urera girardiniodes Seem. Bot. Voy. Herald 194. 1854. ' Low tree, shrub, or erect herb, 1-4 m. high, the branches densely covered with stout bristles or spines; leaves 15-35 cm. long and wide, deeply incised-lobed (lobes acuminate, entire or few-toothed), spiny on the nerves beneath, membranous, glabrescent; inflorescence paniculate, up to 20 cm. long; staminate flowers in glomerules, the pistillate in glomerules or often distinct, sessile; achenes about 1.5 mm. long, suboblique. (231) 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN Costa Rica to Venezuela and Peru, at low altitudes. canal zone: Las Cruces, Seemann 872 (type of 17. girardiniodes) . 2. Urera baccifera (L.) Gaud. Voy. Uran. Bot. 497. 1826. Urtica baccifera L. Sp. PL ed. 2. 1398. 1762. Coarse, erect, subligneous herb, or a shrub or small tree, 1-6 m. high, the stem usually covered with short, stout prickles; leaves broadly ovate or round-ovate to oblong-ovate, up to 35 cm. long and 25 cm. wide, rounded or cordate at base, coarsely dentate or irregularly sinuate-dentate, usually with scattered stinging hairs above and with such hairs or prickles on the nerves beneath; plants dioecious, the flowers in much-branched cymes; fruit succulent, 3-5 mm. long, white or rose- Widely distributed in tropical America. The hairs are very painfully stinging. canal zone: Ancon, Piper 600O; Fort Sherman, Standley 31160. COLON: Lagarto, Cowell 257. darien: Pinogana, Allen 926; Marraganti, R. S. Williams 602. Panama: Taboguillo Island, Miller 2001; sabanas near Panama, Paul 361 ; Juan Diaz, Standley 30536; Rio Tecumen, Standley 26692, 26732; Rio Tapia, Standley 28244. 3. Urera caracasana (Jacq.) Griseb. Fl. Brit. W. Ind. 154. 1859. Urtica caracasana Jacq. Hort. Schoenbr. 3 :71. pi. 396. 1798. Urtica verrucosa Liebm. in Vidensk. Selsk. Skr. V. 2:295. 1851. Urera jacquini Wedd. in Ann. Sci. Nat. III. 18:200. 1852. Shrub or small tree, 2-10 m. high, with elongate branches, the young twigs, petioles, inflorescence, and leaf nerves usually armed with stinging hairs; leaves broadly ovate to ovate-lanceolate, up to 30 cm. long and 25 cm. wide, acuminate, cordate or cordulate at the base (basal lobes sometimes overlapping), crenate- dentate, scabrous, sometimes verrucose, above, more or less pubescent beneath; plants dioecious, the cymes regularly dichotomous; staminate flowers sessile in distinct, compact glomerules; pistillate flowers sessile or pedicillate in compact glomerules (rachis and segments sometimes becoming enlarged), the segments usually densely covered with white, punctiform cystoliths; achenes 0.5-1 mm. long. Widely distributed in continental tropical America from sea level to 2000 m. altitude. The species is here interpreted in a very broad sense, and the specimens cited show considerable variation in shape and texture of the leaves, presence or absence and nature of the foliar cystoliths, degree of pubescence, arrangement of the flowers, and size of the achenes. The American species of Urera are greatly bocas del toro: Changuinola Valley, Dunlap 174; Nievecita, Woodson, Allen & 'ibert 1849; Darkland, von Wedel I; Water Valley, von Wedel 686. canal zone: amei Hill, Vittier 3804. chiriqui: Puerto Armuelles, Woodson tf Schery 863; between iso Ancho and Monte Lirio, upper Rio C . Allen 1471 (variant de- ribed as Urtica :ro Ga lera Chorcha, near Gualaca, Allen 5063; Bajo horro Boquete District, Davidson 349. col6n: Rio FattS, Pittier 3899. darien: Boca i Cupe, Allen 911 R. S. Williams 716; Cana, R. S. Williams 8l 5 ; Yape, Allen 8S5; — , Allen 4592. Panama: Juan Diaz, Standley 30487; Arenoso, flora of Panama (Urticaceae) 193 4. Urera elata (Sw.) Griseb. Fl. Brit. W. Ind. 154. 1859. Urtica elata Sw. Prodr. 37. 1788. Shrub or small tree, 2-4 m. (occasionally to 6 m.) high, sometimes becoming scandent; leaves ovate-oblong to elliptic-ovate, up to 30 cm. long and 12 cm. wide, acuminate at apex, rounded at base, sometimes slightly emarginate, crenate- serrate, crenulate, or subentire, glabrescent, or occasionally hirsutulous on the principal nerves beneath; plants dioecious, the inflorescences dichotomous, much shorter than the petioles, staminate flowers sessile in glomerules, glabrous; pistillate flowers short-pediceled, usually borne singly. With much the same distribution in Panama as the preceding species; also in Jamaica and Colombia. Two forms are represented by the specimens cited below; one has relatively small leaves which bear punctiform cystoliths densely massed all over the upper surface, and agrees best with type material of U. elata in the British Museum; the other has leaves generally more than 1 5 cm. long, the cystoliths being short-linear and radiating from the center of the areoles to the veinlets. This latter variant has been confused with U. alceifolia (Poir.) Gaud., of French Guiana, which tot be separated from the commoi 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN bocas del toro: Changuinola Valley, Dunhp 241; Rio Cricamola, between Finca St. Louis and Konkintoe, alt. 10-50 m., Woodson, Allen & Seibert 1923. canal zone: Hayes 750; Barro Colorado Island, Standley 41132, Kenoyer 396; Madden Dam, Allen 2008. chiriqui': vicinity of Bajo Mona, alt. 1500-2000 m., Woodson tf Schery 593, Woodson, Allen & Seibert 1005; near Monte Lirio, R, I h riq u \ dley, G. White 63; Progreso, Cooper & Slater 176; El Boquete, alt. 1000-1300 m., Maxon 4.992, Pittier 2939. Valle de Anton, trail to Las Minas, alt. 1000 m., Allen 2461, 2/12; vicinity of Finca Tomas Arias, alt. 600 m., Allen 3617. colon: Rio Fato, alt. 10-100 4189. darien: Cana, R. S. Williams 794. 5. POUZOLZIA Gaud. Pouzolzia Gaud. Voy. Uran. Bot. 503. 1826. Leucococcus Liebm. in Dansk. Vid. Selsk. Skrift. V. 2:311. 1851. Margarocarpus Wedd. in Ann. Sci. Nat. IV. 1:203. 1854. Low shrubs, rarely woody vines; stipules free, often persistent; leaves alternate (in American species), entire or rarely toothed; petiolate, trinerved, densely covered with punctiform cystoliths above; plants usually monoecious, the flowers in small clusters, in the leaf axils or forming spikes; staminate perianth depressed- globose in bud, 3-5-parted or -lobed, the stamens 3-5 ; pistillate perianth tubular, contracted at the throat, 2-4 toothed, completely enclosing the ovary, strongly nerved, the stigma pubescent on one side, the achenes crustaceous, shiny. a. Leaves white-tomentose beneath, long-petiolate 1. p. guatemalana 1. Pouzolzia guatemalana (Blume) Wedd. in DC. Prodr. 16 1 :233. 1869. Boehmeria guatemalana Blume, Mus. Bot. Lugd. Bat. 2:206. 1856. Slender shrub; branches hirsutulous; leaves lanceolate or ovate-lanceolate, 6-15 cm. long, 2-4.5 cm. wide, caudate-acuminate, obtuse or acute at base, entire, some- times bullate, asperate above, pilose on the nerves and veins and white-tomentose elsewhere beneath; petioles 2-5 cm. long, very slender; glomerules androgynous; staminate flowers 4-merous, the pistillate 4-toothed. Guatemala to Panama, lowlands of the Atlantic side; rare. bocas del toro: beyond Rio San Juan, Changuinola Valley, Dunhp 457. 2. Pouzolzia obliqua (Poepp.) Wedd. in Arch. Mus. Paris 9:405. 1857. Margarocarpus obliquus Poepp. et Wedd. Ann. Sci. Nat. IV. 1:204. 1854. (Boehmeria obltqua Poepp., nom. nud. as synonym) . Shrub or small tree, 1-5 m. high, densely hirsute nearly throughout, the branches sometimes scandent; leaves oblong or oblong-lanceolate, 2-15 cm. long, 1-5 cm. wide, entire, attenuate-acuminate at apex, oblique and usually emarginate at base, scabrous above, the petioles up to 1 cm. long; flower clusters androgynous or unisexual; staminate flowers 4-merous; achenes light brown or white, the styles flora of Panama (Urticaceae) bocas del toro: vicinity of Chiriqui Lagoon, von Wedel 1 434, 2597; Valley, Dunlap tfo. canal zone: Barro Colorado Island, Kctuner 327, A, //<•< -'.//'; Balboa, Standley 23410, 25587, 29265; Mamei Hill, Pittier 3792; Rio Pedro Miguel, Standley 29976; Las Cascadas Plantation, Standley 25783; Cerro Gordo, near Culebra, Standley 25972; Gamboa, Pittier 6651, Standley 28306; Darien Station, Standley 31606; Las Cruces Trail, Standley 29070; Cocoli, Riley 128. Leucococcus occ Shrub or sr (Liebm.) Wedd. in Arch. Mus. Paris 9:410. 1857. ill tree, up to 5 m. high, the branches densely hirsutulous to nearly elliptic-lanceolate to broadly lanceolate-ovate, 7-15 cm. long, 3-9 linate at apex, cuneate or rounded at base, entire, appressed- with stiff scattered hairs, more or less pilosulous beneath, especially md at the margin; petioles slender, 2-8 cm. long; glomerules androgynous, Honduras thickets at loi arely unisexual, :o Colombia and Venezuela; elevations. :a de Cupe, 40 m. alt., Allen 86, -ed, the in Puerto Rico. In forests and ■anama: Taboga Island, Pittier 3530, 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN 6. PHENAX Wedd. Phenax Wedd. in Ann. Sci. Nat. IV. 1:191. 1854. Unarmed shrubs or suffrutescent herbs; leaves alternate, petiolate, toothed, rarely entire, the cystoliths mainly punctiform; plants monoecious or dioecious, the flowers in dense, sessile, axillary clusters; staminate flowers usually 4-lobed; pistillate flowers without a perianth, the small achene sessile and subtended by a brown, scarious bractlet, the stigma elongate-filiform, persistent. 1. Phenax angustifolius (H.B.K.) Wedd. in Ann. Sci. Nat. IV. 1:193. 1854. Boehmeria angustifolia H.B.K. Nov. Gen. & Sp. 2:34. 1817. Shrub or suffrutescent herb, up to 3 m. high; leaves narrowly lanceolate, 5-15 base, thin membranous, glabrous above, very sparingly strigillose on the nerves and veins beneath, the petioles very slender; plants dioecious; achenes minute, about 0.3 mm. long, the styles 4-5 mm. long. Costa Rica to Peru and Bolivia, usually at elevations of less than 800 meters. bocas del toro: Changuinola Valley, Dunlap 37 1, cocle: vicinity of El Valle, 100-800 m. alt., Allen 764. 2. Phenax mexicanus Wedd. in Ann. Sci. Nat. IV. 1:193. 1854. or subcoriaceous, the petioles up to 5 cm. long, usually much shorter; plants monoecious; achenes about 0.8 mm. long. Southern Mexico to western Panama, usually at elevations of more than 800 chiriqui: El Boquete, Maxon 4995, Pittier 2881, Davidson 452; Finca Lerida to Pefia Blanca, Woodson 8 Schery 312; valley of the upper Rio Chiriqui Viejo, P. White 340. A related species, P. hirtus (Sw.) Wedd., a common plant of the tropics, apparently has not yet been collected in Panama. 3. Phenax rugosus (Poir.) Wedd. in DC. Prodr. 16 x :235 38 . 1869. Procris rugosa Poir. in Lam. Encycl. 5:628. 1804. Boehmeria rugosa Pers. Synops. 2:556. 1807. Shrub, 1-3.5 m. high, the stem stout, densely hirsute; leaves ovate, 3-12 cm. long, 1-7 cm. wide, acute or acuminate at apex, rounded or rarely cordulate at base, crenate-serrate, rugose, usually villous-tomentose beneath; plants monoecious, the glomerules very dense; achenes about 0.5 mm. long. (236) flora of Panama (Urticaceae) Southern Mexico tc > Venezuela and Bolivia, usually occurring above 1000 chiriqui: El Boquet central valley of Rio Chii e, Pittier 2Q02, Davidson 544; vicinity of "New Switzerlan iqui Viejo, Allen 1370. 7. MYRIOCARPA Benth. Myriocarpa Benth. Bot. Voy. Sulph. 168. 1844. Unarmed trees or shrubs, without stinging hairs, leaves alternate, large, toothed or rarely subentire, 3 -nerved, petiolate, bearing conspicuous cystoliths; plants dioecious, rarely monoecious; staminate flowers in small, contiguous glomerules, forming slender, dichotomous spikes, the perianth 4-parted; pistillate flowers in slender, elongate, dichotomous, sub-unilateral spikes, rarely in panicles, without a perianth but subtended by 2 or 4 bractlets, the achenes stipitate or sessile. (237) 198 ANNALS OF THE MISSOURI Killip, in Proc. Biol. Soc. Washington 1. Myriocarpa 40:29. 1927. Myriocarpa longipes var. yzabalensis Donn. Sm. in Bot. Gaz. 16:13. 1891. Shrub or tree, 2-10 m. high; leaves elliptic-ovate or broadly ovate, up to 35 cm. long and 18 cm. wide, acuminate at apex, rounded or slightly narrowed at base, denticulate, bearing numerous conspicuous linear cystoliths which radiate from the center of the areoles, glabrous above, glabrous, or pubescent on the nerves beneath, the petioles up to 35 cm. long; staminate spikes up to 10 cm. long; pistillate spikes very slender, pendent, up to 60 cm. long, the bractlets 2, soon deciduous, the achenes scabrid with sparse short hairs, eciliate, at length black and sometimes lustrous. Guatemala to Panama, generally at elevations of less than 1000 meters. bocas del toro: Changuinola Valley, Dunlap 30O; Chiriqui Lagoon, von Wedel 14.II, 1445, 1505, 1692, 1699, 1886; Columbus Island, Skutch 5; Almirante, Cooper 414. canal zone: Chagres, Fendler 280; Barro Colorado Island, Shattuck 242, 494, Aviles 27, Standley 31451, 40922; Frijoles, Killip 3379, Standi 27194; Gamboa, Standley 28400; Obispo, Standley 31687; Fort Sherman, Standley 30936; Santa Rita Trail, Cowell II 3; Rio Indio, Dodge & Allen 1 7289; Quebrada Salamanca, Dodge, Steyermark & Allen 1 6997. chiriqui: El Boquete, Maxon 4946, Pittier 2878, 5157- cocle: El Valle, Allen 4218. colon: Porto Bello, Pittier 2487. Panama: Rio Tapia, Standley 26213, 28277; Rio Tecumen, Standley 29368. flora of Panama (Proteaceae) Trees, shrubs or rarely herbaceous, rarely dioecious. Leaves alternate, rarely opposite or whorled, simple or compound, sometimes heteromorphic, pinnately veined; estipulate. Inflorescence a head, spike or raceme, axillary or terminal, bracteate, the bracts persistent or caducous, subtending 1 to several flowers. Flowers bisexual or unisexual by abortion, monochlamydeous, cyclic, hypogynous, actinomorphic or zygomorphic. Tepals free or variously connate, petalaceous, tetramerous, valvate in bud. Stamens 4, antipetalous, the anthers dehiscing longi- tudinally, the filaments variously adnate to the perianth. Pistil 1 , often surrounded basally by glands or a disc, often stipitate, the ovary superior, 1-carpellate, 1-loculate, the ovules 1-many, with 2 integuments, the placentation parietal or the ovules pendulous, the style 1, slender, the stigma 1, lateral or terminal. Fruit a follicle, achene, samara or drupe; seeds lacking endosperm, sometimes winged. A family primarily restricted to the dry regions of the Southern Hemisphere, consisting of about 55 j and South African. Eight gei iera are represented in the New World of which two are indigenous to Panama. A number of species are c ultivated in numerous parts of Central America for their extremely showy infloresi :ences. At present I have no evidence of cultivated species in Panama. The following treatment i; > based on a revision of the American Proteaceae by Dr. H. Sleumer (in Bot. Jahrb. 76:139-211. 1954). a. Leaves alternate, simple or pinnat ely compound (heteromorphic), entire or dentate; inflorescences racem glands; fruit a follicle, obliquely ■ 1. ROUPALA Aubl. Roupala Aubl. Pi. Guyan. Franc. 1:83,/. 32. 1775. Leinkeria Scop. Intr. 345, n. 1607. 1777. Khopala Schreb. Gen. PI. 1:62, n. 144. 1789. RoM* J. F. Gmel. Syst. 2:225. 1791. Kupala Vahl, Symb. 3:20. 1794. Shrubs or trees. Leaves alternate, heteromorphic, simple or pinnately com- pound, dentate to entire, petiolate. Inflorescence spikes or racemes, axillary or terminal. Flowers bisexual, actinomorphic. Tepals free. Stamens inserted at about the middle of the perianth, the anthers oblong-linear, subsessile to sessile, connective sometimes broadly but briefly produced. Disc of 4 hypogynous glands. Ovary with 2 pendulous ovules, the stigma terminal. Fruit a follicle, obliquely 2-valved, with 2 winged seeds. An American genus containing about 50 species distributed from the Isthmus of Tehuantepec southwards to Peru, Bolivia and Brazil (Rio Grande do Sul) . One species is known from Panama. (239) 200 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. Roupala Montana Aubl. PL Guyan. Franc. 1:83. 1775. Roupala Pyrifolia Salisb. & Knight, in Knight Prot. 102. 1809. Rhopala media R. Br. in Trans. Linn. Soc. 10:191. 1810. Rhopala dentata R. Br. loc. cit. 192. 1810. Rhopala complicata Kunth, in HBK. Nov. Gen. & Sp. 2:153, /. ng. 1817. Rhopala ovalis Pohl, PL Bras- Ic. 1:107, t. 86. 1827. Rhopala macropoda Klotzsch & Karst. in Linnaea 20:473. 1847. Rhopala gardneri Meissn. in Mart. Fl. Bras. 5 1 :83, t. 31, f. I. 1855. Rhopala martii Meissn. loc. cit. 87, /. ?2 , /. 3. 1855. Rhopala martiiwxT. loc. cit. 1855. Rhopala frondosa Rich, ex Meissn. in DC. Prod. 14:427. 1856, in syn. Rhopala tomentosa a integrifolia Meissn. loc. cit. 428. 1856. Rhopala bohnctu:*; a Meissn. loc. cit. 430. 1856. Rhopala yrr#j . foe tit. 43 5. 1856, in synon. Roupala borealh Hemsl. Biol. Centr. Am. Bot. 3:78, /. 76. 1882. Roupala darienensis Pittier, in Contr. U. S. Nat. Herb. 18:228. 1917. Roupala pana,,: 2:9. 1917. Roupala discolor Rusby, Descr. N. Sp. Am. PL 12. 1920. Roupala disshntlh Pit.,.:. B. 1 w Wn./ Cuk Nat. 5:303. 1939. Roupala repanda Lundell, in Am. Midi. Nat. 29:472. 1943. Roupala montana var. dentata (R. Br.) Sleumer, in Engl. Bot. Jahrb. 76:173. 1954. Trees to 20 m. tall, the young stems terete, ferrugineous-strigillose and glabres- cent. Leaves heteromorphic; juvenile leaves pinnately compound, generally larger than the adult forms, the number of leaflets extremely variable even on a single branchlet, margin generally coarsely serrate or sometimes undulate, the veins con- spicuous; adult leaves ovate, 5-12 cm. long, 2-9 cm. broad, acute or acuminate at the apex, cuneate to more or less obtuse at the base and often decurrent upon the petiole, subcoriaceous or coriaceous, ferrugineous-strigillose and glabrescent above and below, the costa plane to immersed above, emersed below, the primary lateral veins numerous, conspicuous, the margin entire, undulate to variously dentate or serrate, often repand; petiole 1-6 cm. long. Inflorescence racemose, terminal or more often axillary, multi-flowered, the rhachis 6-15 (-18) cm. long, tomentose, the flowers borne singly or in pairs. Tepals linear-oblanceolate, 7.0-8.5 mm. long, about 1 mm. broad, widely reflexed, strigillose without, glabrous within; stamens inserted about at the middle of the perianth, the anthers linear-oblong, 2-3 mm. long, 0.5 mm. broad, the filaments about 0.5 mm. long, glabrous; hypogynous glands about 0.5 mm. tall, carnose, glabrous; ovary about 1 mm. long, 1 mm. in diameter, densely strigose, the style about 3 mm. long, glabrous, the stigma narrowly clavate; pedicel to 3 mm. long, strigillose. Fruit a flattened follicle, often minutely spurred at the base, 2.5-4.0 cm. long, about 1.5 cm. broad, glabrous, the winged seeds oval, 1.5 cm. long, 0.8 cm. broad. Distribution of the species essentially that of the genus. Flowering in January and February in Panama. between Corozal and Ancon, Pittier 263O; Ancon Hill, Standley 26370, precise locr 1: ~ »— - ete and C '580. coci Allen 2514; between Las ] (240) 9yet 1068, 1070. chiriqui: Sabana de la Tortuga, * Caldera, Pittier 3341 ; Boquete, Davidson 1061, Pit, 152; El Valle flora of Panama (Proteaceae) 201 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN darien: Cana, Williams 739; Rio Pirre, Pittier 6975. Panama: San Jose Island, Perlas archipelago, Gulf of Panama, Johnston 331, 348, 1285, 1292, 1293, 1368; near Matias Hernandez, Standley 28949; Taboga Island, Standley 2800I; Tumba Muerto Road, near Panama, Standley 29739; Rio Las Lajas, Allen 1600; Punta Paitilla, Standley 26242. Sleumer recognized three varieties of R. montana in his recent revision. One variety, var. tomentosa, is not treated herein, nor are its synonyms as it is restricted to Brazil. The two varieties which concern us, var. montana and var. dentata, are both represented in Panama. The character which serves to distinguish these varieties is "Folia subintegra vel undulata" for var. montana and "Folia bene obtuse subcrenato-serrata" for var. dentata. Young shoots and suckers have pinnately compound leaves, the number of leaflets is quite variable although the older gener- ally have more than the younger, all leaflets have a serrate margin. This pinnately compound leaf is a typical juvenile form. The mature form is a simple leaf with serrate to entire margin. Although flowering is usually evidence of a mature shoot it is not necessarily evidence that the subtending leaves have attained the mature form. A few specimens show the transition from simple leaves with serrate margin to subentire or entire margin while others show that the larger stems have leaves with undulate or entire margins and the young stems have leaves which are quite serrate. It is my belief that the simple leaf with serrate margin is transitional between the compound leaf and the simple leaf with entire margin. Finally, in many collections some of the leaves are entire while others are variously serrate- needless to say this leads to the condition in which many collections and some specimens are classified in two separate varieties. Accordingly, I have not recognized var. dentata of Sleumer. 2. PANOPSIS Salisb. Panopsis Salisb. in Knight, Prot. 104. 1809. Andriapetalum Pohl, PL Bras. Ic. 1:113. 1827. And ripe talum Schott ex Endl. Gen. PI. 342. 1836. Trees. Leaves alternate, opposite or verticillate, entire, simple. Inflorescences racemes, terminal or axillary, bracts small or 0. Flowers bisexual, actinomorphic. Tepals free. Stamens inserted below the middle of the perianth, the anthers oblong, connective apiculate, the filaments obvious. Disc cupuliform, 4-lobed. Ovary subsessile, with 2 pendulous ovules, the stigma terminal. Fruit indehiscent with a thick woody pericarp and 1 exalate seed. An American genus distributed from Costa Rica southwards to Peru, Bolivia and Brazil. It contains about 11 species of which 1 is known from Panama. Andnpetalum suaveolens Kl. & Karsten, ex Klotzsch, in Linnaea 20:472. 1847. Panopsis costancensis Standi, in Journ. Wash. Acad. Sc. 17:164. 1927. Panopsis mucronata Cuatr. in Lloydia 13:202, /. I. 1950. Trees to 12 m. tall, the young branches terete, ferrugineous-strigillose glabrescent, conspicuously lenticellate. Leaves elliptic, oblong-elliptic or oblar (242) flora of Panama (Proteaceae) 203 olate, 4-17 cm. long, 1.5-5.0 cm. broad, obtu at the apex, cuneate to obtuse at the base, coriaceous, ferrugineous-strigillose and glabrescent above and below, the costa plane above, emersed below, the primary- lateral veins numerous, extremely reticulate, conspicuously emersed; petiole 2-8 mm. long. Inflorescence racemose, terminal or axillary, multi- flowered, the rhachis 6-15 cm. long, ferrugineous-strigillose, the flowers borne singly or to 4 per cluster. Tepals linear-oblanceolate, 7-8 mm. long, 0.5-0.75 mm. broad, revolutely reflexed, strigillose without, glabrous within; stamens inserted at the base of the perianth, the anthers oblong, 1 mm. long, 0.5 mm. broad, the filaments liguliform, about 6.5 mm. long, 0.5 mm. broad, glabrous; pistil long- conical, the ovary to 2 mm. long, 0.5-0.75 mm. in diameter, densely hirsute, the style 4-5 mm. long, glabrous, the stigma narrowly clavate; pedicel 4-8 mm. long, ferrugineous- strigillose. Fruit ellipsoid to ovoid, 3.5-6.0 cm. long, 2-4 cm. in diameter. Distributed in Costa Rica, Panama, Venezuela, Colombia and Ecuador. In Panama flowering in May between 1000 and 2000 meters. According to G. White the wood ranks about 5th in Panama as a building material and is very hard and heavy. Known in Panama as aguacatia. chiriqui': Rio Chiriqui Viejo, G. White 112; trail from Paso Anch upper valley of Rio Chiriqui Viejo, Allen 1483; Boquete, Davidson 630, Pittier 2954. ■ 5'.??. MISSOURI BOTANICAL GARDEN Volume XL VII Annals of the Missouri Botanical Garden Annals Missouri Botanical Garden Annals of the Missouri Botanical Garden SEPTEMBER, 1960 THE STRENGTHENING SYSTEM IN THE STEM OF MAIZE* WILLIAM H. MURDY Introduction The species Tea mays is characterized today by an almost overwhelming range of variability; a consequence attributed in part to its unique history in association with ethnic groups of man in the New World, and in part, to former introgression with the genus Tripsacum and the present ability to exchange germ plasm with Euchlaena (teosinte; a genus of putative hybrid origin between Zea and Tripsacum). The concept of race has been advanced as a step toward a natural classification of this variability. Anderson and Cutler (1942), largely responsible for the evolution of this concept, defined a race loosely as a "population of maize plants differing from other groups by a great number of genes and sharing enough char- acters in common to be easily recognized as a distinct group". For the past several years, the Rockefeller Foundation in collaboration with Latin American countries has been performing the arduous task of classifying indigenous races of maize in Central and South America. The character of scle- roticness, as expressed by features such as stalk stiffness, the coarseness of leaves and cob induration, is one of many polygenic characters by which races of maize have been found to differ. The magnitude of tissue induration is frequently accepted as evidence of teosinte or Tripsacum contamination, since highly sclerotic plants often exhibit other plant characters regarded as "tripsacoid" in nature (Wellhausen, 1952). Moreover, segregates from maize- teosinte crosses often 206 ANNALS OF THE MISSOURI BOTANICAL GARDEN possess an inordinate degree of tissue induration (Mangelsdorf and Cameron, 1942). Galinat et al. (1956) state that "extensive experiments with maize-teosinte hybrids have demonstrated that genes responsible for lignification occur on many, if not all, of the chromosomes of teosinte". The character of sclerification, when properly evaluated, should add materially to our knowledge concerning the origin of modern maize and its past and present relationship with Euchlaena and Tripsacum. The present study was undertaken in order to shed light on the following basic questions: (1) What stem tissues become sclerified to make up the strengthening system and how do these tissues differ in amount and arrangement from the base of a stem to its apex? (2) What is the range of variability within and between races of maize in respect to the strength- ening system? (3) What relationship exists between the strengthening system of maize and that of Euchlaena and Tripsacum} One of the foremost requisites of good breeding stock is an ability to stand up well and contribute a fair share of stalk-stiffness in crosses. Several anatomical studies, based on the lower internodes of inbred lines, have established a correlation between lodging resistance and the nature of sclerotic tissue (Magee, 1948; Hunter and Dalbey, 1937). The results of these investigations are valuable to the corn breeder, but have only an indirect bearing on the nature of sclerotic, strengthening systems in vigorous open-pollinated races. Materials and Methods Six maize races were selected from the "Standard Exotics" described by Ander- son and Brown (1951), which represent the extremes of variation in Zea mays growable in the corn belt. Many plants of each race were grown under essentially uniform field conditions at the Pioneer Hi-Bred Corn Company, Des Moines, Iowa. Following is a brief description of each race. Gourdseed (Brown and Anderson, 1948)— An old southern dent variety typical of the kind of maize commonly grown in southeastern United States in colonial times. This particular stock was collected by Dr. W. L. Brown in 1946 at Grape- vine, Texas. Plants are generally tall, having many internodes, few tillers, and greatly condensed ears and tassels. Chromosome knob numbers medium high (5-7) ; plants mature slowly (approximately 94 days from planting to silk in Iowa). Northern flint (Brown and Anderson, 1947)— Representative of the maize once grown almost exclusively in the northern and eastern sectors of the United States until superseded by early corn-belt varieties. Plants vary somewhat through- out their range, but share many features in common including low chromosome knob numbers (0-2), eight-rowed ears, few, but long internodes (those above the ear not decreasing in length as in the case of Gourdseed) , several tillers shorter in height than the main stem, and a rapid growth rate (from planting to silk in Iowa takes approximately 66 days) . Argentine pop (Anderson and Brown, 1951) — A small-eared, prolific pop corn race of Paraguay and Argentina. Unlike the other five races, Argentine pop has not likely been contaminated by either North American Tripsacum or teosinte. Furthermore, its ear is closest to the concept of prehistoric South American maize MURDY STRENGTHENING SYSTEM IN STEM OF MAIZE 207 based on archeological evidence. It grows slowly in Iowa and at maturity consists of a main shoot with many short internodes relatively equal in length, plus a few tillers shorter than the main culm. Planting to silk requires approximately 105 Chapalote (Wellhausen, 1952)— A pop corn race grown in the coastal lowlands of northwestern Mexico. Plants are "tripsacoid" in appearance in the possession of several long, ear-bearing tillers approaching in height the main shoot, long inter- nodes, and long narrow leaves. Wellhausen classified this race as a primitive, indigenous pop corn with little evidence of teosinte contamination. Chromosome knob number is variable; the average is approximately six. About 90 days are required from planting to silk in Iowa. Papago (Anderson and Cutler, 1942) — Seed of this race was obtained from the Papago Indians in the desert south of Tucson, Arizona. It is typical of the maize grown by desert-dwelling Indians and similar to archeological maize grown by the prehistoric Basketmakers — the first agriculturalists of this region to leave a record (Anderson and Blanchard, 1942). Its habit resembles that of Chapalote; long ear-bearing tillers, long internodes, and long narrow leaves. Chromosome knob number is medium high and growth rate quite rapid (approximately 77 days from planting to silk in Iowa). Zapalote chico (Wellhausen, 1952) — An extremely sclerotic race grown in the coastal lowlands of southern Mexico. Plants are coarse and indurate, lack tillers, and have many short internodes above the ear. Chromosome knob number is high (12-16) and Wellhausen considers this race to be highly contaminated with teo- sinte germ plasm. It matures rapidly in its native habitat and is comparatively independent of day length when grown in Iowa. Planting to silk requires approx- imately 81 days. Six plants of Florida teosinte (Euchlaena mexicana), from a commercial source, were used in this study; two were grown in Iowa and four in Florida. Plants from two separate clones of Tripsacum dactyloides were sampled; one growing in Iowa and the other located in St. Louis, Missouri. A single plant of Tripsacum lanceo- latum collected in Guatemala by Dr. E. Anderson was also included in this study. Just after anthesis, 15 plants of each maize race were selected in an attempt to adequately sample the range of morphological variability within races. Plants were collected when completely mature and subsequently subjected to morphological and anatomical analysis. In making anatomical sections of desiccated stems, internodal discs about 2 cm. long were boiled in water, with aerosol added, for approximately 4-5 hours; placed in 50 percent alcohol overnight; sectioned with a sharp razor; stained with safranin and fast green; and permanently mounted in piccolyte, a synthetic mounting medium. Maturation of a Maize Stem ; of alternating segments, or :r is made up of an internode, with a leaf at its upper end and a bud at its base. Final maturation of a phytomer i the leaf blade, passes down the leaf sheath and finally down the internode 208 ANNALS OF THE MISSOURI BOTANICAL GARDEN below. Successive phytomers, however, mature acropetally up to, but not includ- ing, the tassel peduncle, which together with its tassel does not await the ascending wave of maturation, but matures relatively early at a time when the internodes immediately below are still immature. Normally there is no leaf associated with the tassel peduncle, thus, internodal tissue maturation begins in the upper portion of this internode. By measuring internode lengths of genotypically uniform hybrid plants at various stages of maturity, one is able to obtain a picture of the onto- genetic pattern of stem development (fig. 1). Note the plant examined ten weeks after planting; it is still relatively immature, nevertheless, the leafless peduncle is already 1 1 cm. long, almost half its assumed maximum length, while the two inter- nodes immediately below, associated with leaves, are merely 2 cm. in length, only one-ninth their assumed maximum length at maturity. A similar pattern of maturation was reported by Prat (1935) for stems of Secale cereale. He noted that the terminal internodes bearing the inflorescence elongated more rapidly than the subterminal internode. By subjecting these two internodes to mechanical stress, he found the uppermost to be more resistant. He concluded that rapid maturation accompanied rapid elongation and that both processes occurred faster in the terminal internode than in the one below. It is likely that the maturation of a maize stem conforms to a pattern wide- spread in the Graminae. Knowledge of this pattern is essential to an understanding of maize stem anatomy, for the rate at which an internode grows and matures has a direct bearing on its internal structural features. Strengthening Tissues in a Maize Stem In herbaceous monocots generally, mechanical stem strength is provided by thick-walled sclerotic tissue. The process of cell sclerification entails the deposition of a thick, cellulosic secondary wall and subsequent impregnation of primary and secondary walls with lignin. Tissues that contribute most to the strength of a maize stem can be classified in three categories: (1) sclerenchyma associated with vascular btmdles; (2) hypodermal sclerenchyma; (3) sclerified parenchyma. Vascular bundle sclerenchyma. A maize stem in cross-sectional view (plate VI, D) shows a scattered arrangement of vascular bundles surrounded by paren- chymatous ground tissue. Those bundles near the periphery of the stem are crowded, small and generally provided with heavy sclerotic sheaths, whereas the more central ones are widely-spaced, larger in size, and lack massive sheaths. The presence of two distinct vascular systems in maize has previously been reported. Both Esau (1943) and Cutler and Cutler (1948) reported two systems in the stem. The latter authors referred to the outer system as the "peripheral" and the inner as the "central". Two quite dissimilar systems have also been noted in the ear (Lenz, 1948; Laubengayer, 1949; Reeves, 1950) and tassel (Kumazawa, 1940). MURDY STRENGTHENING SYSTEM IN STEM OF MAIZE 209 The leaves of maize are vascularized by large, early-maturing bundles and smaller late-forming bundles which alternate with the larger ones. At a node, the large leaf bundles pass some distance into the stem at a right angle before assuming a vertical or oblique course down through the internode below. Ontogenetically, the large bundles originate at the node and simultaneously differentiate in two directions; up into the leaf and down through the internode before either of these organs have completed their elongation. Only when acropetal differentiation of these large bundles reaches the tip of the leaf blade, do the small alternating 210 ANNALS OF THE MISSOURI BOTANICAL GARDEN bundles begin to differentiate basipetally in the distal part of the leaf blade, develop- ing at a time when most elongation of leaf and stem has been completed (Sharman, 1942). At a node these small bundles pass into a ring of vascular tissue, formed by the lateral anastomosis of peripheral strands from the internode above, and their sheaths become merged with those of the ring bundles. Just below the node the peripheral ring of vascular tissue breaks up into distinct bundles, which pass down the periphery of the internode below. Hence, the majority of peripheral stem bundles, which play a leading role in support of the stem, are prolongations of late- maturing leaf bundles. Some of the smallest bundles found in the peripheral region of the stem have an even later origin in the intercalary meristem region at the base of a leaf sheath. Esau (1943) illustrated the origin of such bundles within the massive sheaths of large, early-maturing bundles. All vascular bundles of appreciable size are associated with a sheath of pro- tective fiber cells, heaviest at the protophloem and protoxylem poles (plate VI, B) . Esau (1943) described the ontogeny of the vascular bundle in maize. She pointed out that the fibers of the sheath originate mainly from the procambium which also gives rise to the vascular tissue itself; in addition, tissue adjacent to a bundle may contribute cells to the sheath in its somewhat advanced stage of development. The amount of such accretion varies with the size of the bundle and its location in the plant. The vascular bundles with associated sheath sclerenchyma comprise the "back- bone" of the skeletal structure supporting the stem of a maize plant. In the form of strengthening rods passing discretely through parenchymatous ground tissue the bundles contribute little to the support of the plant. It is only when they are arranged in various structural patterns that their mechanical function is most effective. The peripheral bundles with prominent sheaths, may, by the scarifica- tion of inter-bundle parenchyma, form a continuous supporting cylinder, at least in the lower internodes (plate VI, A). Similar peripheral bundles in upper inter- nodes are frequently welded to the epidermis by a bridge of sclerotic tissue derived in part from the bundle sheath and partly from the hypodermis (plate VII; Hypodermis. Immediately beneath the epidermis of a maize stem is a zone of thick-walled, elongate fiber cells (plate VI, A). This tissue, commonly known as a hypodermis, is lacking or poorly developed in lateral and foliar organs. Fre- quently the hypodermis is very thick in its radial dimension and makes a substantial contribution to the strength of certain parts of the stem. Ontogenetically, the subepidermal meristematic cylinder which gives rise to the hypodermis probably originates from the "flank meristem" of Ledin (1954). Popham (1951) refers to this histogen as a peripheral meristem forming a cylinder between the protoderm, which gives rise to the epidermis, and central meristem, which develops into pith and procambial strands. In ontogeny, parenchyma cells underlying the epidermis undergo longitudinal divisions and as the internode elongates, these initials do not undergo transverse division, but grow rapidly in the MURDY STRENGTHENING SYSTEM IN STEM OF MAIZE 211 axial direction resulting in the development of long narrow fibers (Hayward, 1938). Hypodermal fibers isolated from plants of the race Papago average 1.2 mm. in length and are 100X longer than broad. During growth, the tips of the fibers push past one another resulting at maturity in a very strong non-storied peripheral cylinder. Accentuated development of hypodermal sclerenchyma is found just above the point where a leaf or lateral branch departs from the main stem. This fact may have a significant bearing on the problem of intense sclerification of the cob rachis where the internode above each lateral branch (spikelet pair) is greatly abbreviated. Sclerified parenchyma. Parenchymatous tissue in the outer region of a stem fre- quently becomes thick-walled and lignified to an extent that differs greatly among internodes of a single plant. When well-developed, the sclerotic parenchyma takes the form of a thick, subepidermal, sclerotic cylinder pervaded by peripheral vascular bundles (plate VII; A, B). Changes in the Strengthening System of an Individual Stem Structural changes in anatomy from one internode to another have not been described for maize, but are known in other grasses. Prat (1935) found internodes to vary in their ability to withstand mechanical pressures. The lower inter- nodes were strongest in this respect with resistance diminishing acropetally. Athanassoff (1928) described the anatomy of successive internodes of a wheat culm. He found that internodes differed in the relative amount and arrangement of sclerenchyma, parenchyma, and vascular tissue. He also noted that internodes differed in respect to their time of maturation. Anatomical sections of six internodes from the stem of a single plant are pic- tured in plate VII. The principal structural differences among internodes include the following: variation in size, shape, and sheath development of peripheral vascular bundles; differences in the relationship of these outer bundles with sub- epidermal sclerenchyma; and, changes in the extent of hypodermis and sclerified parenchyma development. Peripheral vascular bundles. The peripheral bundles are very small in the lower- most internode (plate VII, A), but increase in size acropetally. In the internodes below the ear (plate VII; A, B, C) peripheral bundle sheaths, though massive, show very little centrifugal development, and between sclerenchyma at the proto- phloem end of the bundles and sclerenchyma of the hypodermis is a zone of thin- or thick-walled parenchyma. Peripheral bundles in the internodes above the ear (plate VII; D, E, F) are large in size and rhomboid in shape. These bundles have a much greater development of sheath sclerenchyma at the proto- phloem pole which, being confluent with the hypodermis, results in the formation of sclerotic "bridges" connecting the peripheral bundles to the epidermis. The small, peripheral bundles in the lower internodes (plate VII; A, B) evidently matured after these internodes had ceased to elongate. They have little protophloem and protoxylem and are relatively unstretched, as evidenced by the 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN lack of a protoxylem lacuna (Esau, 1943). Their complete dissociation from the hypodermis and sub-epidermal parenchyma infers that their development in the internode occurs after these outer stem tissues have matured. In the internode just below the lowest ear (plate VII, C) peripheral bundles are somewhat larger than in lower internodes. In addition, they possess protoxylem and the middle bundle is somewhat stretched. Therefore, up to this point, there is an acropetal narrowing of the time lag between the maturation of the outer internodal tissues and the peripheral bundles. From the next internode on into the tassel the anatomy of the stem undergoes a striking change (plate VII, D). In these upper internodes, peripheral bundles differentiate prior to the completion of internode elongation and at a time when the outer tissues of the internode are still immature and capable of further division and elongation. As a result, these bundles are large, with well developed protoxylem and protophloem, and a large protoxylem lacuna. Moreover, bundle sheaths are well developed at the protophloem pole and confluent with hypodermal sclerenchyma. Except for size, the peripheral bundles in the peduncle and tassel (plate VII; E, F) are comparable to those of the inter- nodes immediately below and are similarly bridged to the epidermis. The structural variation of vascular bundles in the corn stem described above is attributable to differences in the relative period of development and maturation of outer stem tissues and the vascular bundles descending through them from leaves above. Starting with the lowermost internodes where the peripheral bundles develop some time after the completion of internode elongation, there is a gradual change in this relationship whereby the development and maturation of the two tissues is progressively more closely approximated. At a point where the lowermost ear is borne, the development and maturation of the two tissues is coincident. Sharman (1943) has pointed out that the time taken for the internode to complete its development increases with the maturity of the plant. He noted that "in the upper internodes there may be a lapse of four or more plastochrones between the full expansion of the lamina and internode." In other words the development and maturation of leaves with their included vascular strands, occurs at a fairly uniform rate acropetally, whereas the maturation of tissue in successively higher internodes progressively slows down. An internode pattern of this plant (fig. 4, lower right) shows that the lower internodes (aside from the first two above the ground), which on the basis of anatomy were found to mature rapidly, are elongate, whereas the slower-maturing, upper internodes have relatively short lengths. Consequently, the changes in the length of internodes coincides with the changes in structural anatomy and both reflect the particular growth behavior characteristic of this plant. Hypodermis. The greatest development of hypodermal sclerenchyma occurs in the middle internodes of the plant (plate VII; C, D), diminishing in extent below (plate VII; A, B) and above, where it is found only opposite bundles in peduncle and tassel internodes (plate VII; E, F). The lower internodes and tassel mature more rapidly than internodes between and those just beneath the peduncle are the slowest to mature. It is precisely in these short, upper internodes, having the , that the hypodermis has its greatest development. MURDY STRENGTHENING SYSTEM IN STEM OF MAIZE 213 Sclerified parenchyma. In the upper internodes intervening parenchyma between peripheral bundles is firm, but relatively thin-walled and unlignified. Intense sclerification of ground parenchyma occurs in internodes quite low on the plant (plate VII; A, B). The lower internodes are the earliest to originate and have a longer period of time to deposit cell wall material. In addition, there is a great basipetal flow of sugar through these lower internodes especially in the early stages of the plant's growth. The upper internodes have a longer period to differentiate fiber cells, but a shorter period for these cells to deposit cell wall material prior to final desiccation. Furthermore, after fertilization of the ear a great amount of sugar is channeled into the developing kernels causing a drain on the availability of sugars required for the synthesis of cellulose in late-developing internodes. Racial Variation in the Strengthening System of Maize Hypodermis. In all races examined (except Argentine pop) hypodermal tissue develops to its greatest extent in the upper stem internodes between ear and tassel. It is suggested that prolonged activity of the peripheral meristem in these upper, slower-maturing internodes results in the differentiation of a great number of hypodermal fiber cells prior to tissue maturation. Races differ, however, in the extent of hypodermal tissue development, which may in part result from a dis- in growth behavior. For example, plants of the race Gourdseed consis- tently have a thicker hypodermis (greater radial number of fiber cells) than do plants of the race Northern flint (plate VIII; B, D) . As a race, however, Gourdseed is characterized by having a slow rate of growth and a series of short slow-maturing internodes above the ear shoot — the region of greatest hypodermal development in all races. Northern flint plants, on the other hand, have a rapid growth rate and the upper internodes are long and relatively quick to mature. Therefore, when comparing the potential ability of these two races to produce hypodermal scleren- chyma, their respective growth patterns must not be overlooked. Although in most races the upper internodes invariably mature more slowly than middle and lower ones, the race Argentine pop is characterized by a very uniform rate of growth resulting in all internodes being short and relatively slow to mature. In plants of this race middle internodes of the stem show the greatest extent of hypodermal development (plate IX, B) . Growth rate does not wholly account for the hypodermal differences observed among races. Papago, the race with the thickest hypodermis (plate VIII, A) , is one of the most rapid to mature. In this race, however, the unusually heavy hypo- dermis is associated with deep-lying peripheral vascular bundles and, both features may have definite selective value for such a semi-xerophytic race in diminishing the extent of water loss from stem tissues. The amount of hypodermal tissue development in plants of a particular race is fairly uniform. Where intra-racial differences do occur, they are accompanied by changes in the overall scleroticness of the plant. When the width of the hypo- 214 ANNALS OF THE MISSOURI BOTANICAL GARDEN dermal zone is measured in the internode of its greatest development, races of maize analyzed line up from greatest to least in the following manner: Papago (plate VIII, A) ; Gourdseed (plate VIII, B) ; Zapalote chico (plate VII, D) ; Chapa- lote (plate VIII, C) ; Argentine pop (plate IX, B) ; Northern flint (plate VIII, D). Hypodermal tissue is not a prominent feature in the stems of teosinte plants examined (plate VIII, E). Tripsacum likewise lacks an appreciable development of continuous hypodermal tissue. The strengthening system of Tripsacum, however, is characterized by separate, peripheral strands of procambial tissue which at maturity form sclerotic connections between peripheral vascular bundles and epidermis. Thin-walled chlorenchymatous tissue lies between adjacent girders. $ $ MURDY STRENGTHENING SYSTEM IN STEM OF MAIZE 215 Vascular bundles. The vascular bundles found in any cross section of a corn stem vary in size, shape, and also in their amount of associated sheath sclerenchyma (plate VI, D). Just inside the outer ring of heavily sclerotic bundles is a zone of strands, intermediate in character between the small, often reduced peripheral strands and the large, greatly stretched central ones. This ring of bundles is most easily discerned in the upper internodes of a corn stem and within a particular internodal cross section bundles are remarkably uniform in all their features. Determinations of bundle size were carried out on five such bundles (one appears on plate VI, B) from the fourth internode below the peduncle in each plant studied. Bundle size refers only to the dimensions of the vascular tissue within the bundle sheath. Vascular bundle size varies somewhat among plants of a single race, but to a greater extent from plants of one race to those of another. Nevertheless, in both instances there exists a strong correlation between the diameter of a particular internode and the size of its contained bundles. The relationship of bundle size to stem diameter for five races of maize plus teosinte and two species of Tripsacum is diagrammed in fig. 2. Each symbol represents the average bundle dimensions of a race with the relative amount of sheath sclerenchyma in black. The races Papago (10), Northern flint (8), Zapalote chico (9), and most Gourdseed plants (6) show a heavy development of sheath sclerenchyma in proportion to bundle size, especially when compared with Chapalote (7), Argentine pop (5), and teosinte (4). Relative to bundle size, sheath sclerenchyma is weakly developed in teosinte, greatly developed in Tripsacum dactyloides (2), and massive in Tripsacum lanceolatum (3). Peripheral stem bundles become progressively smaller and more reduced in successively lower internodes. This change is accompanied by an increase in their associated sheath sclerenchyma. Moreover, the walls of the fiber cells constituting the bundle sheath become thicker in successively lower internodes. All plants of the race Zapalote chico conform to the structural pattern shown by the plant pictured in plate VII in that most of the peripheral bundles of upper internodes are connected to the epidermis by a sclerenchymatous bridge. A similar pattern, however, is not found in all races of maize. For bridging to occur in any plant internode it is necessary that the peripheral bundles lie close to the epidermis and that sufficient bundle sheath sclerenchyma develops at the phloem Papago has deep-lying peripheral bundles and though heavily sclerotic, none are bridged to the epidermis in any stem internode. Plants of the race Chapalote vary in their incidence of bridging, but the variation is correlative with peripheral bundle distance from the epidermis. On the other hand, proximity of periph- eral bundles to the epidermis is not solely responsible for the incidence of bridging, either in a single stem or in a race. For example, peripheral bundles in all internodes of a teosinte stem lie close to the epidermis, but bridging only occurs in the peduncle and upper one or two internodes (plate X, A). In respect to races, a 216 ANNALS OF THE MISSOURI peduncles of Zapalote chico plants studied have over fifty per cent of their periph- eral bundles bridged to the epidermis, even when these bundles lie 150 microns from the epidermis (a distance greater than that of unbridged Chapalote plants). On the other hand, the peripheral bundles in the stems of Argentine pop lie relatively close to the epidermis, but are unbridged. Considering only the phenomenon of bridging in maize, teosinte and Tripsacum, the following classification can be made. 1. No bridging in any internode of the plant. This condition is found in Argentine pop (plate IX; A, B) and in the North American pop corn race, Ladyfinger. 2. Bridging in the tassel, tassel peduncle and 2-3 internodes below. North- ern flint, Zapalote chico (plate VII; A-F) , Gourdseed (plate IX; C, D) , Chapalote, and teosinte (plate X; A, B.) 3. Bridging in all plant internodes. Tripsacum dactyloides (plate X; C, D) and Tripsacum lanceolatum. The lack of bridging in Argentine pop may be due to the distance of peripheral strands from the epidermis, paucity of sheath sclerenchyma, or both. It is possible, however, that it is in part a result of a particular pattern of growth, basically different from both Tripsacum and the races of maize which exhibit bridging. Argentine pop plants have a very slow growth rate and at maturity each plant consists of a great number of short internodes differing little in length (fig. 4). The longest internode occurs in the lower one-third of the plant. Beyond this point successive internodes gradually diminish in length up to the tassel peduncle, which itself is seldom longer than the internode next below and barely, if at all, exserted beyond the uppermost leaf sheath. Peripheral bundles are uniformly small and unstretched throughout the length of the plant. This infers that in each internode the peripheral bundles mature after the completion of tissue elongation. Hence, in all internodes of the stem, internodal tissue and peripheral bundles mature at a rather uniform rate, the former keeping ahead of the latter. The presence of bridging in upper plant internodes but not in those below has been described for Zapalote chico (plate VII) and attributed to an extreme retardation in the rate of maturation of internodes above the ear shoots. A similar change occurs in the slow-maturing race Gourdseed, which like Zapalote chico, is characterized by a series of very short internodes between the ear and tassel. Northern flint and teosinte grow rapidly and the upper internodes slow down only slightly in their rate of growth and maturation (fig. 4). As in Gourdseed and Zapalote chico, the peduncle and upper one or two internodes exhibit bridging. Such bridging is likely the result of a speeded-up maturation of peripheral bundles coupled with a slight decrease in stem tissue maturation, for the upper 2-3 leaves are always very much shorter than those below. Hence, the peripheral bundles have a much shorter distance to differentiate before reaching the stem. In the case of Zapalote chico, the change in peripheral bundle configuration is abrupt; in Northern flint and teosinte it is gradual. For example, in one plant of teosinte, MURDY STRENGTHENING SYSTEM IN STEM OF MAIZE 217 : bridged to the epidermis, In the genus Tripsacum, the formation of girder sclerenchyma is an outstand- ing feature of its strengthening system, occurring throughout the length of the plant (plate X; C, D). It is also a common structural arrangement in grasses generally. Hayward (1938) stated that stems so constructed have a "continuous zone of hypodermis variable in thickness within the epidermis and enclosing longi- tudinal bands of chlorenchyma". More precisely, the outer portion of such a stem consists of a parenchymatous cortex, regularly interrupted by girders of sclerotic tissue, which link peripheral bundles with the epidermis (plate X, C). Only in lower internodes do peripheral bundles become connected laterally by the scarifica- tion of parenchyma lying between them (plate X, D). In these lower internodes the longitudinal bands of parenchyma appear to be "imbedded" in a continuous mass of hypodermal sclerenchyma, but actually, the isolation is circumscribed laterally by girder sclerenchyma and within by sclerified parenchyma. Percival (1921) stated that "a portion of the outer face of the chlorenchyma bands is always in immediate contact with the epidermis". This is true for the upper internodes of Tripsacum but in lower internodes the epidermis is underlain by a single layer of large hypodermal cells. -^o^ %^f ^ir> 15^> UPPER LOWER ARG.POP ZAP.CHICO TEOSINTE TRIPSACUM 218 ANNALS OF THE MISSOURI BOTANICAL GARDEN The peripheral bundles in Tripsacum lie close to the epidermis and their matura- tion in the stem either coincides with that of outer stem tissue or the procambium itself extends to the epidermis. Tripsacum has a rapid growth rate and at maturity consists of a relatively small number of extremely long narrow internodes cul- minating in an even longer peduncle (fig. 4). Maturation processes in this genus must be investigated before an attempt is made to compare its pattern of growth In summary, the strengthening system found in the lower internodes of most maize races studied parallels the condition found in an entire Argentine pop plant. The upper internodes, however, have a structure resembling that of Tripsacum. Figure 3 diagrammatically represents the basic differences in bridging described in this section. Sclerified parenchyma. In all plants examined the lower the internode the wider the zone of sclerified parenchyma. In lower internodes sclerified parenchyma forms a radially thick, sclerotic cylinder traversed by vascular bundles (plate VII, A). Higher up (approximately 8 internodes below the tassel peduncle) it is found chiefly between peripheral vascular bundles (plate VI, B; plate IX, B, D; plate X, B, D). In the fourth internode below the tassel peduncle interfascicular scleren- chyma is found only in Papago (plate VIII, A) and in the tassel peduncle sclerified parenchyma is never found (plate IX, A, C; plate X, A, C). A high degree of intra-racial variation occurs in regard to the extent of parenchyma that becomes sclerified, however, races which consistently were weak in the development of sclerified parenchyma include Argentine pop (plate IX, B), Chapalote, and Northern flint, whereas it was always prominent in Gourdseed (plate IX, D), teosinte (plate X, B), Papago, and Zapalote chico. Its greatest development was found in Tripsacum (plate X, D). The Strengthening System of the Lateral Ear Shoot The lateral ear-bearing shoot, or shank, is anatomically similar to the main stem except for its lack of a well-developed hypodermis and the presence of numer- ous small, peripheral bundles, which originate in the excessively wide leaf sheaths or husks. The very sclerotic race Zapalote chico does have a hypodermis in the shank and at maturity internodes have a smooth surface. In other races, there is little or no hypodermis present and with desiccation the collapse of parenchymatous tissue beneath the epidermis results in the formation of externally visible grooves and ridges. In addition to sclerotic tissue, shank strength depends on such factors as length of shank, length of internodes, diameter of internodes and weight of the ear. There is a great difference in the anatomy of internodes of a single shank depending on their length. Consequently, it is difficult to compare the short shank internodes of Gourdseed, for example, with the very long shank internodes of Northern flint. Furthermore, the amount of sclerenchyma in a shank bearing a fertilized ear is much greater than in one which bears an unfertilized ear. SYSTEM IN STEM OF MAIZE 219 Vascular bundles in the shank are generally provided with a heavy sclerotic sheath, which frequently prevents crushing of vascular tissue when arching or bending occurs. The overall strength of the shank, however, depends on the strength of tissue between the bundles or the extent of ground tissue sclerification. The races Zapalote chico and Papago, very sclerotic in respect to their stems, have a great amount of sclerified parenchyma in their shank internodes. In plants of these two races, shanks varied in length from 7-28 cm., but all were erect at maturity. Plants of the race Gourdseed characteristically have short shanks (6- 16 cm.) and their internodes have a fair amount of sclerified parenchyma. Never- theless, all shanks of this race were bent or broken due in large measure to the combination of a very heavy ear and narrow lower internodes. Both Northern flint and Chapalote have relatively long shanks (11-32 cm.). 220 ANNALS OF THE MISSOURI BOTANICAL GARDEN At maturity all shanks of Chapalote plants were erect, whereas shanks of Northern flint plants were either bent or broken. The long shanks of Northern flint have few internodes and grow rapidly, while on the contrary, those of Chapalote have many internodes and a slow rate of growth. Anatomically, shank internodes of Chapalote have a wide zone of sclerified parenchyma, but there is practically none of this tissue present in the internodes of Northern flint. Since the stems of both races are quite similar in respect to sclerenchyma, the greater shank strength in Chapalote is probably due to the much longer period cells are able to deposit wall material before final maturation and desiccation. Argentine pop has small ears borne on relatively short shanks which have many internodes, grow slowly and have fairly strong interbundle connections. Conclusion The tissue which strengthens and supports a maize stem can be subdivided into three categories: long fibers of the hypodermis; short fibers of peripheral bundle sheaths; and thick- walled, lignified parenchyma. The amount and disposition of these three tissues differs in a single stem and the manner of change from the base of a plant to its apex is dependent on the plants growth behavior. As a result, the relative importance of each tissue in contributing strength changes in different parts The greatest sclerification of ground parenchyma takes place in the lower internodes — the earliest on the plant to mature. This tissue diminishes acropetally and occurs only between peripheral bundles in upper middle internodes and not at all in the tassel, peduncle and the two to three internodes below. Middle and upper internodes are strengthened principally by hypodermal sclerenchyma, which develops its greatest radial thickness in the slow-maturing internodes between ear and tassel peduncle. The importance of peripheral bundle sclerenchyma increases acropetally. In middle internodes these bundles are interconnected laterally by sclerified parenchyma and form a continuous sclerotic cylinder within the hypo- dermis. Higher up, the lateral connections are weaker but sheath sclerenchyma developing at the phloem end of a bundle may connect with the hypodermis or epidermis forming a subepidermal girder. The basic growth pattern of all maize races is similar in that elongation and maturation of leaves and internodes occurs acropetally with the former generally preceding the latter. The rate of leaf maturation is fairly even, however, each successively higher internode elongates and matures at a slower rate than the one next below so that an upper internode may still be elongating while its associated leaf is already fully mature. Differences in the time of maturation of stem tissue and vascular bundles entering the stem from leaves leads to changes in the struc- tural anatomy of the stem from base to apex. Just as races differ in their pattern of growth, the manner and time of structural changes in the stem differ from plants of one race to those of another. MURDY STRENGTHENING SYSTEM IN STEM OF MAIZE 221 A race such as Argentine pop with a uniform rate of growth shows only slight changes in its structural anatomy from one internode to the next, whereas Zapalote chico with a rapid initial growth rate, which is abruptly slowed down above the position of an ear shoot, has a corresponding sudden change in structural anatomy. Northern flint with a rather even but rapid growth rate undergoes changes similar to those in Zapalote chico but much more gradually. In the upper internodes of such a plant the length of leaves decreases along with the time of stem maturation. The maize races studied were classified for overall scleroticness on the basis of their hypodermis, peripheral bundle sheaths and sclerified parenchyma and the result was in agreement with field observations on standability. All plants used in the present study were harvested after final maturation and desiccation, at which time it was noted if stems were standing, bent over or broken. Results of this observation are presented below. No. of plar Gourdseed Argentine pop 3 7 5 20% weak The introduction of teosinte germ plasm into maize reportedly leads to an increased induration of all plant parts (Mangelsdorf and Cameron 1942; Galinat et al. 1956) . Teosinte plants used in the present study were not especially indurate; in fact, they were less sclerotic than most of the maize races examined. Two plants do not represent teosinte as a whole; nevertheless, they do show that the range of variation within the genus Euchlaena includes quite unsclerotic plants. Both species of Tripsacum were very sclerotic, but T. lanceolatum much more so than T. dactyloides. The arrangement of sclerotic tissue in Tripsacum is similar internodes it is distinctly different from both Euchlaena and maize. Literature Cited nd H. C. Cutler. 1942. Races of Zea Mays: I. Their recognition and classification. 222 ANNALS OF THE MISSOURI BOTANICAL GARDEN Esau, K. 194J. Ontogeny of the vascular bundle in Zea Mays. Hilgardia 15:327-68. Evans, M. W., and F. O. Grover. 1940. Developmental morphology of the growing archaeological maize. Bot. Mus. Leaflets, Harvard Univ. 17:101-123. Hunter, J. W., and N. E. Dalbey. 1937. A histological study of stalk breaking in m Laubengayer, R. A. ale inflorescence of Zea Mays. i of Zea Mays in relation to i Jour. Heredity 2 cival, J. 1921. The Wheat Plant. E. P. Dutton Company, ] Explanation of Plate PLATE VI Structural features of a maize stem. A, Transection of the peripheral region of an internode showing hypodermis (HD), peripheral vascular bundles (PB), and thick-walled parenchyma between the latter (X100). B, Transection of a vascular bundle (X107) (., \n atypical vascular bundle lacking well-developed metaxylem vessels. Bundles of this kind were found in the tassel peduncles of all Zapalote plants (X107). D, Transection ol a maize stem taken from a middle internode which shows the arrangement of vasculai B =, ~V////. F E ■■ -STRENGTHENING SYSTEM IN STEM OF MAIZE MURDY STRENGTHENING Explanation PLATE i ANNALS OF THE MISSOURI BOTANICAL GARDEN PLATE VIII i internode below the t /t i MURDY STRENGTHENING SYSTEM IN STEM OF MAIZE 225 THE MISSOURI BOTANICAL GARDEN Teosinte (A, B) and Tri |§|g| ■ J D ° STUDIES INVOLVING SUSTAINED TREATMENT OF MAIZE WITH GIBBERELLIC ACID I: FURTHER NOTES ON RESPONSES OF RACES* NORTON H. NICKERSON and THOMAS N. EMBLER cultivar. Norton H. Nickerson, Missouri Botanical Garden, 2315 Tower Grove . 10, Missouri; Thomas N. Embler, Dennis-Yarmouth Regional High School, < Introduction In previously reported work (Nickerson, 1959), one hybrid, tw representatives of two exotic kinds of maize were subjected third day with four different concentrations of GA. This paper presents results from identical treatment of four other kinds of maize which differ in their behavioral responses. Materials and Methods Four kinds of maize 1 were employed in this study; they were representatives of three clearly separable races (Anderson and Cutler, 1942). A Northern Flint type (Brown and Anderson, 1947), New York Flint, was chosen to compare its behavior to that of Parker's Flint, a representative employed in the previous study (loc. cit.). Classifiable as "Southern Dent corns" (Brown and Anderson, 1948), Gourdseed (see their Plate 20) and Cherokee Dent (probably a member of their Mexican June complex; see their Plate 22) were chosen both to give a comparison between two forms whose ears were different but whose plant types were not, and to complete the study of Corn-belt ancestral types (Anderson and Brown, 1952a, 1952b) from combinations of which presumably such inbreds as L317 and CC5 investigated earlier (loc. cit.) were derived. Two ears of Cherokee Dent, one with 12 rows and the other with 14 rows, were obtained from Henry Busby, who in 1959 was about 90 years old. He farms the upper end of an isolated valley near Pollard, Arkansas. His father was a full- blooded Cherokee Indian (or Cherocow, according to his nephew, from whom these details came) who came from North Carolina by way of Tennessee; this strain of maize was brought to Arkansas with him and has been grown, but probably not in complete isolation, by his family since pre-Civil War times. It is the only corn seed: Dr. William H. Murdv, Department of 1 Gourdseed: Mr. Derrel Busby. Route 1. Pollai Dr. E. G. Anderson, California Institute of Tech 228 ANNALS OF THE MISSOURI BOTANICAL GARDEN which the family considers fit for human consumption; this fact may have helped keep it relatively uncontaminated. The family refers to this maize as "90-day corn", and often uses it for two crops each year in Arkansas. In New England, at 41° 42' North Latitude (the equivalent of central Iowa), this maize reached anthesis in 74 days during the 1959 growing season. Fully ripened ears were harvested 35 days later, but it is not known how many days preceding actual har- vest date they may have reached ripeness. Its growing season in a cooler more northerly latitude thus approximates 100 days. A distinctive South American type, Argentine Pop, was also included. This pop corn was collected among others by Cutler in 1941 (personal communication) . It has figured prominently in recent genetic work involving the reconstruction of an ancestral maize form (Mangelsdorf, 1958 a, b). According to Alava (1952), Nickerson (1953) and Anderson and Brown (1953), the original source of the strain employed in this study and used for genetical and observational work both at Johnston, Iowa, and Arcadia, California, was seed collected by Mr. Lorenzo Parodi. Plants of both Argentine Pop and Gourdseed have been studied anatom- ically by Murdy (1960). In the present study, each type of maize was planted in 15 -meter rows; plants were spaced about 25 cm. in the row, and rows were 1 Yz meters apart. In view of results obtained from randomized rows and blocks of maize in a previous study (Nickerson, 1959) there was no attempt to randomize treatments applied within each row. The total number of plants in each row was divided into five groups with as nearly equal numbers as possible, and all plants in a particular group received the same treatment. The five treatments were the same as those used before and are as follows: 1 — distilled water (controls) 4 — distilled water with 125 ppm GA Every third day for the duration of treatments one ml. of the appropriate solution containing either distilled water or water plus the above-listed concentrations of GA 2 was applied from a pipette into the apical leaf cavity of each plant. Solutions were freshly made each time, or stored no longer than three days in darkness at 19° C. Insects (corn borer and corn earworm) were controlled by hand removal and by a weekly dusting of 5% DDT ("Neudust") beginning when evidence of infestation was first noted (July 10) and continuing until elongation ceased and pollination had occurred. Fertilizer (Agrico 5-10-10) was applied at the rate of 700 lbs./acre three and six weeks after germination. Planting date was June 12, 1959; treatments began July 6 and continued until tassel emergence (Table 1). ^cultural Research Division, Eli NICKERSON & EMBLER GffiBERELLIC ACID TREATMENT I 229 Initial response in all forms was rapid. Two days after the been applied, plants in all treated groups showed increases in height; these ranged up to twice the heights of control plants. During the next two weeks, controls underwent little or no internode elongation; analysis of mature treated plants (Fig. 1) shows that early internode elongation was greatly enhanced by GA, provided that the internode tissue had not reached its final stage of differentiation by the time of treatment. An attempt was made to record the sixth node on each plant by counting all leaves from the coleoptile up (but not the coleoptile itself) when the plants were young. This fifth leaf was known from previous observa- tions to be present on mature plants. It was marked by clipping a metal staple into the leaf. Small arrows on the internode diagrams (Fig. 1) show these sixth nodes as they were determined by fifth-leaf attachment when the plants were mature. As a precaution against wind damage, many of the rows were staked and plants tied to lines beginning during the fifth and sixth week and continuing throughout the growing season. New York Flint. (Tables 1, 2, 6; Fig. 1). Higher treatments of 125 and 625 ppm tended to increase lengths of internodes below the ear most drastically; even though upper internode lengths were less than those of the lower internodes in these plants, they were still equal to or greater than upper internode lengths in control plants. Just as noted in the previous study on the closely related Parker's Flint, the 125 ppm treatment proved most detrimental, and there was a recovery toward normal by plants treated with the higher 625 ppm doses. Ear production on plants treated with 125 ppm doses was much lower than with all other treatments. Ears produced by plants treated with 625 ppm doses were much shorter and smaller than those produced by controls, but kernels were set. Tillering was inhibited entirely by treatments of 25, 125 and 625 ppm. The average number of tassel branches was markedly reduced from 15.8 on controls to 6.2 on plants treated with 125 ppm doses. Plants receiving 625 ppm doses produced nearly twice this minimum number, but their average of 1 1 branches was still only y 3 the number produced on control plants. Tassels were exserted within 2-3 days in all groups, but anthesis was first noted in plants treated with 25 ppm doses. Leaves averaged 7 cm. at their widest point on controls. On plants receiving 5 ppm doses, leaves averaged 5 cm. in width and were generally longer by 15 cm. than the same leaves on controls compared just before tassel emergence. Plants treated with 25 ppm doses had at this time leaves as wide as controls but again longer by 15 cm. In plants receiving 125 ppm treatments culm diameter was only half that of control plants; leaves were no longer than control plants, and they averaged only 3 cm. in width at their widest points. The 625 ppm treatments produced plants with quite thin culms (5-10 mm.), long internodes, and compared with controls, much narrower and shorter, occasionally rolled leaves. These rolled leaves were noted previously on plants which were eventually killed by an overdose of GA (Inbreds L317 and CC5; Nickerson, 1959) ; their rolling may be described 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN as epinastic with relation to the midrib rather than with relation to the culm, in that the upper epidermis seems to grow more rapidly than the lower to the extent that a tubular structure is formed. A picture of this bizarre feature is shown (for Argentine Pop) in Plate XI. Tassels of these plants all produced female caryopses, but in only two of the four saved for study were these functional. These female spikelets were produced along the basal portions of branches and central spikes. Brace roots were not formed as copiously on these plants as on Parker's Flint, but primordia did appear as high as 30 cm. above the ground. Their total growth averaged 2-4 cm. Tassel height was significantly increased in plants receiving 5, 25 and 125 ppm doses; it was decreased in plants receiving 625 ppm doses. Gourdseed. (Tables 1, 3, 6; Fig. 1). All plants treated with GA showed initial responses in height. Plants treated with 5 ppm doses were definitely more vigorous than controls. This vigor was first manifested in increased height; plants 6 weeks old averaged 45-60 cm. compared with an average height of 30-36 cm. for controls. This effect persisted through to maturity. Leaf number (13-14) at 6 weeks was the same for controls, 5 ppm and 25 ppm-treated plants, but with 5 ppm treatments, leaf blades were 10-15 cm. longer than the 75-90 cm. average lengths on controls; leaf blade width at widest point averaged 7-9 cm. compared to the 6-8 cm. average of control plants. The average number of ears and of primary tassel branches showed increases with both 5 ppm and 25 ppm doses com- pared with controls. Although plants treated with 2 5 ppm doses reached greater average heights than did those treated with 5 ppm doses, they were not as vigorous as controls. Culms were slightly smaller, and leaves all averaged 6 cm. in width. On plants treated with 125 ppm doses, ear number was not reduced compared to controls, height was not increased over treatments using 25 ppm and primary tassel branch number was reduced only slightly compared to controls. Total tassel height was also about equal to that of controls. Leaves were equal to controls in length, but averaged only 3-5 cm. in width and were easily broken. Plants treated with 625 ppm doses showed a marked height increase compared to controls; this increase was attributable entirely to greater elongation of lower internodes. Culms however were thinner; leaves averaged only 2-3 cm. wide and were easily broken. Ear number was reduced, tassel height was reduced approx- imately 14% and the area over which tassel branches arose was greater by 40% when compared to controls. At 1 1 weeks of age, tassels had fully emerged in all other plants, but not in this group. When they emerged, 3 out of the 5 tassels collected for study had some well-developed caryopses in basal areas of branches and central spikes. In these areas, sessile spikelets were always female and pedicellate spikelets were always male. Of the five tassels, one with caryopses and one with no caryopses were male sterile. Pollen shedding, therefore, was apparently not dependent upon the presence or absence of caryopses in the tassel. Brace root formation in control and 5 ppm-treated plants was confined to the " rely; in plants treated with 25 ppm doses, they arose on i number of nodes formed brace roots in plants NICKERSON & EMBLER GIBBERELLIC ACID TREATMENT I 231 V GO 1 fi CD 1 CD 2 A? GO 2 AP 2 w v^-Us/ CD 3 /""V \l v >v^> /vJ _^""" "N GO 5 ^W NYF, New York Flic Number 1 stands for rtical axis is internode leng row depicts node of attachrr t; GO, Gourdseed; CD, Ch distilled water treatments ent'of fiftn'S above C coleo rokee Dent; AP, Argentine its. Horizontal axis Circle denotes tassel, >op.' 232 ANNALS OF THE MISSOURI BOTANICAL GARDEN treated with 125 ppm, but the extreme lengths of the internodes in this group meant that some of the brace roots were initiated more than 40 cm. above ground. These roots seldom grew more than 3 or 4 cm., and many were only 1 cm. in length. In plants treated with 625 ppm doses, there were consistently five nodes visible on which brace roots had initiated. Root length was also seldom more than 3-4 cm., and for the majority was 1 cm. They were, however, produced as high as 80 cm. above ground. Cherokee Dent. (Tables 1, 4, 6; Fig. 1). This maize resembled Gourdseed rather closely in responses and appearances of treated and untreated plants. Greatest responses occurred in younger plants; as plant size increased, recovery toward appearance of control plants was noted. Greatest height increase was with doses of 125 ppm; relatively little effect was noted on most measurements except when doses of GA applied contained 625 ppm. At that level, number of ears, number of primary tassel branches, length of central spike and peduncle length showed decreases over the same measurements in controls; only the area over which tassel branches arise showed an increase over controls. Tassels of these plants, however, did not develop any female spikelets. Bending over occurred when the plants were 6—9 weeks of age in the 125 ppm- and 625 ppm- treated groups. The phenomenon occurred in rapidly elongating internodes 1 meter above ground. Such bent-over stalks could not be straightened without breakage. One control plant had the only two tillers observed in this population. Argentine Pop. (Tables 1,5,6; Fig. 1 ; Plate XI) . This race was highly sensitive to all doses of GA. In plants 6 weeks of age, those receiving 5 ppm doses averaged 20-30 cm. in height compared with 15-20 cm. for controls. They had 10-11 leaves visible compared to 8-9 leaves on controls. Tiller number of plants treated with 5 ppm doses was also reduced; at 6 weeks, controls had 2-3 tillers per plant and those treated with 5 ppm doses had only one tiller per plant. This difference was less at maturity, however. Among plants treated with 25 ppm, tiller number and ear number were both reduced below the corresponding values for control plants. A few silks were produced in basal areas of tassel branches and central spikes. Also, leaf width was reduced by half from averages of 5-7 cm. in controls to 2.5-3.5 cm. At 6 weeks, plants treated with 25 ppm doses were 30-37 cm- tall. Plants receiving 125 ppm doses of GA were nearly all eliminated by the treatment; some plants, however, persisted longer than others. At 6 weeks of age, 10 out of 22 plants had been killed; at 11 weeks, only 5 were surviving, and two of these had been mutilated by wind. This mutilation was likely attributable to the extremely brittle nature of these plants. Under doses of 625 ppm, plants showed a strong initial response, but at 6 weeks of age only 8 out of 22 were alive, and at 11 weeks, only one plant was alive. This latter plant was not more than 5 cm. above ground, and its stem was highly contorted (Plate XI). Both doses of 125 and 625 ppm produced plants with thin leaves. In one 125 ppm-treated plant, they were in addition so inrolled that their tips never became free; a "ladder" effect was thus obtained (Plate XI). This plant produced a tassel, never exserted, which was female below and male above; it also lacked tassel branches (Plate XI). NICKERSON & EMBLER GIBBERELLIC ACID TREATMENT I 233 Discussion and Conclusions This study, along with the previous one (Nickerson, 1959), reinforces the point that the effects of GA on maize are marked, but dependent upon both the race studied and the concentration of GA employed. Internode diagrams (Ander- son and Schregardus, 1944) for the four maizes studied were constructed as before, being in each case in Fig. 1 that of an actual plant which best represented its group of five plants studied. These diagrams were constructed from the tassel down, hence the internodes were drawn in the inverse order of their appearance, and tassels are at the same relative position on each graph. New York Flint and Cherokee Dent show the same pattern of internode elonga- tion above the points of ear attachment. Gourdseed and Cherokee Dent, however, show a much greater similarity with regard to internodal development below the point of ear attachment. Argentine Pop, because of the large number of ears formed, is not comparable to any of the other races. Its growth pattern is here the same as that reported for it by Murdy (1960) under growing conditions in Iowa: the first three internodes are increasingly longer, a plateau is then reached followed by a gradual decline in successive lengths until the longer peduncle is developed. Vertical comparison of these graphs within each column shows what concentra- tions of GA affected which internodes within each race of maize: horizontal comparisons show how the various races responded to the same concentrations of GA. In New York Flint, 5 ppm and 25 ppm treatments produced growth patterns generally alike, affecting most of the internodes above the point of ear insertion. Arrows indicate the point of attachment of the 5 th leaf at the top of the fifth internode. The coleoptile is not here counted as the first leaf. Extra internodes were consistently formed below points of ear attachment on plants treated with 125 and 625 ppm GA. An extra internode appears in the diagram of the 5 ppm- treated plant above the point of ear attachment. This observation was true for 4 of the 5 plants measured. Possibly low concentrations of GA applied early enough delay formation of inflorescence primordia for a time. This suggestion is in line with findings of Bradley and Crane (1960) on Prunus sp. cultivars, where GA prevented initiation of floral primordia but did not interfere with production of bud-scales. A comparison between diagrams of New York Flint and Parker's Flint of the previous study (loc. cit.) shows a general similarity of curvature, including the anomalous detrimental behavior under treatments of 125 ppm. The fact that New York Flint plants treated with 5 and 25 ppm doses exhibit less departure from the control growth pattern than do plants of Parker's Flint treated with these same doses may be interpreted as signifying that New York Flint has in its genetic makeup some other maize germ plasm and is not therefore as "pure" a representa- tive of Northern Flint as is Parker's Flint. Gourdseed shows remarkably little internodal pattern change under doses of 5, 234 ANNALS OF THE MISSOURI BOTANICAL GARDEN 25 and 125 ppm compared with controls; under treatments of 625 ppm, its curve is quite similar to that for 125 ppm-treated New York Flint. These results may likewise be interpreted as indicating that Gourdseed is in itself not a "pure" repre- sentative of its race, Southern Dent; its hybrid ancestry, referred to by Brown and Anderson (1948), has apparently buffered its protoplasmic reactions to GA. This same situation may be seen in the previous experiments (loc. cit.) with regard to Spancross, a known hybrid. In fact, in its general slight effect on 5, 25 and 125 ppm-treated plants, and its stimulation of early internodes followed by declines in length except for the peduncle, GA creates much the same effect on both Gourdseed and Spancross. The general beneficial effect of low doses of GA on Gourdseed have been alluded to above; it is possible that GA may find commercial importance in boosting yields on maize strains which are strongly of Southern Dent ancestry. Gourdseed also has much in common with the internode pattern of Zapalote Chico, a race also previously studied (loc. cit.). The two maizes respond nearly alike to treatments with 5 ppm and 25 ppm doses; with the higher concentration of 125 ppm, Zapalote Chico shows more sensitivity to GA than does Gourdseed. When the GA concentration was highest (625 ppm) their developmental patterns were again strongly similar. Cherokee Dent, based on its lower row number and less conspicuous denting, is also a representative with mixed genetic background. This contention is borne out by relatively slight differences in internode patterns of controls, 5, 25 and 125 ppm treatments. There are also strong resemblances between its responses and those of both Gourdseed (see above) and Zapalote Chico of the previous study (loc. cit.). Brown and Anderson (1948) mentioned that the Southern Dents were a variable lot and that on both cytological and morphological evidence they showed a strong affinity to the dent corns of central Mexico. This contention was accepted fully by Wellhausen et al. (1952). In their general internode patterns of control plants and in their similar responses to various doses of GA, these resemblances are further confirmed. Argentine Pop, as it is grown in the United States, is known to be a highly inbred race. Two sorts, based on plant height and ear size, were segregated out of the original collections independently in studies conducted at Johnston, Iowa, (Edgar Anderson, personal communication) and at Arcadia, California, (E. G. collected and observed Argentine Pop in its native range in South America, regards it as being essentially inbred there also. He also reported the taller, fewer-eared segregants as not being "uncommon". This race may therefore be considered to be a rather pure one. The present study, however, used material which for several generations has always produced small ears in large numbers. Ears of this distinctive race have been figured by Weatherwax (1954, fig. 59) and Mangelsdorf (1958 a, b; fig. 6). In its strong sensitivity to GA, Argentine Pop resembles responses of the inbreds NICKERSON & EMBLER GIBBERELLIC ACID TREATMENT I 235 CC5 and L317 studied previously (loc. cit.). Higher doses are fatal; in Fig. 1, the only survivors of the 125 ppm and 625 ppm groups are graphed simply to show the extent to which deleterious reactions may go and still allow the plant to remain alive. These two plants were collected and later photographed (plate XI). The well-documented effects of GA on reduction of tiller number and ear number were also graphically illustrated (table 5). Tassels of 25 ppm-treated plants developed a few silks and female spikelets; the one non-exserted tassel of the 125 ppm-treated survivor showed a completely female basal portion and a male but sterile tip portion (Plate XI). Its general resemblance to a basal immature inflorescence is remarkable. It may therefore be concluded that GA affects maize in a fashion depending directly upon the degree of inbreeding or homozygosity of the strain involved. Since there are now many forms of maize, and since they exhibit a vast range of human control or lack of it over their past pollinations, it seems reasonable to conclude that responses of no one form of maize are "standard". It is precisely because of such racial or varietal differences that generalized conclusions, especially where a variable cultivated plant is used as experimental material, must be based upon wide studies. As an example one might compare the explanation offered for the predominant way in which GA acts in plant elongation by Feucht and Watson (1958) with that of Greulach and Haesloop (1958). The former workers, using the cultivar Blue Lake of Phaseolus vulgaris, concluded that cell elongation was primarily responsible for GA-induced increase in internode length. The latter workers, using the cultivar Black Valentine of Phaseolus vulgaris, concluded that only cell division was responsible for GA-induced increase in internode length. Paleg and Aspinall (1958) reported differences in response to GA under the same environmental conditions in two varieties of barley. Kline (1959) , working with celery plants, showed that age of plant was also important in eliciting a particular response. He suggested it may be more important even than dosage of GA applied. Other examples might also be cited; these instances referred to above, however, reinforce the argument for caution in extrapolative interpreta- The rather marked increase in length of peduncles in most treated groups (Fig. 1; Table 6) as well as a general increase in height of tassels (Table 6) are likely attributable directly to the influence of GA on these parts when the plant is quite young. Murdy (1960) has pointed out, as has Kiesselbach (1949), that the below are still immature. The thin culms and general lack of vigor in maize plants treated with high doses of GA may be a direct result of higher-than-normal respiratory rates. Using wheat seedlings, a 20% rise in respiration under treatment with GA has been reported by Lustinec and Krekule (1959). Coulombe and Paquin (1959), working with tomato, reported rapid increase in respiration, photosynthesis and transpiration with a peak 5-6 hours after treatment, followed by a rapid decline, but higher respiration and photosynthesis rates than controls during limits of their experiment. Ormrod and Williams (1960) found that plants of Trifolium hirtum 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN showed a striking increase in acid soluble phosphorus and decrease in inorganic phosphorus as quickly as one minute after treatment with GA. Using bean plants, Linck and Sudia (1960) showed that GA-treated plants absorbed more P 32 than did controls. There thus may be a correlation between GA treatment, respiratory rate and phosphorus metabolism. The phenomenon of poor leaf separation, noted occasionally in all forms given higher treatments with GA in both this study and the previous one and illustrated by Plate XI, resembles illustrated symptoms of calcium deficiency (Hambidge, 1941). Went (1957) similarly noted this same "harp-like" structure on corn plants grown under continuous light. The brittleness of maize plants treated with higher concentrations of GA may be because of mineral imbalance within the plant. In carnations, Laurie and Kiplinger (1948) reported that overdoses of potassium will cause brittleness and snapping off at nodes. Whether the GA-induced aberrations in mineral content of maize plants also causes a potassium imbalance resulting in brittle stems and leaves is not known. It may only be concluded that overdoses of GA apparently upset mineral balances in plants. Literature Cited Alava, Reino O. 1952. Spikelet variation in Zea Mays L. Ann. Mo. Bot. Garden 39:65-96. Anderson, Edgar and W. L. Brown. 1952a. Origin of Corn Belt Maize and its genetic significance. Heterosis. Edited by John W. Gowen. Iowa State College Press, pp. 124-148. md reproduction of corn. Bull. 161. Univ. of Nebraska College of Agriculture. Lincoln, Nebraska. 96 pp. Kline, John W. 1959. Growth responses of gibberellin-treated celery plants. Bot. Gaz. 120:122- Laurie, Alex and D. C. Kiplinger. 1948. Commercial flower forcing. The Blakiston Company, Philadelphia. 550 pp. tinec, J. and J. Krekule. 1959. , P. C. NICKERSON & EMBLER GIBBERELLIC ACID TREATMENT 1 237 Paleg, L. and D. Aspinall. 1958. Inhibition of the devel Weatherwax, Paul. 1954. Indian Corn in Old Am Wellhausen, E. J., L. M. Roberts and E. Hernandez X. TABLE 1 ■ AMOUNT OF GA RECEIVED BY EACH PLAN : No. of o A r n of GA per treatme \l$ New York 8 -rowed Flint 15 75 375 1875 9175 Sourdseed 18 o 9D 450 2250 1250 Cherokee Dent 15 o 75 375 1S75 9375 Argentine Pop 19 95 475 2375 1875 TABLE 2 — NEW YORK FLINT Treatment in ppm of GA 5 25 125 625 Height (nearest cm.) 126 164 17 233 204 ■z Number of Nodes 10 12 11 11.25 10.75 \ r Number of Tillers 2.6 2.6 c i Number of Ears 2.0 2.0 2.2 0.25 0.75 Number of Primary Tassel Branches 15.8 j .6 6.2 „ Percent of Tassels Wholly Male Sterile 1 100 100 Percent of Tassels with some Functional Pistillate Florets Q 25 /5 Original Size of Population 17 17 17 18 IS Number Surviving 16 16 17 13 11 Number of wind-damaged plants 238 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE 3- GOURDSEED Treatment in ppm of GA 5 25 125 625 Height (nearest cm.) 211 220 243 243 262 -5 Number of Nodes 15.2 15.4 16.8 15.6 15 6 .S a Number of Tillers o o M Number of Ears 1.4 2.2 1.6 1.4 8 Number of Primary Tassel Branches 12.4 14.6 13.0 11.8 11 8 Percent of Tassels Wholly Percent of Tassels with some Functional Pistillate Original Size of Population 25 21 23 23 24 Number Surviving 25 20 23 24 n am ng survivors o -CHEROKEE DENT Height (nearest cm.) 218 233 255 \ Number of Nodes 1 5 8 16 16.4 Number of Tillers 0.2 s! Number of Ears 22 22 IS 20 06 Number of Primary Tassel Branches 21 8 25 6 23 18 8 16 r Percent of Tassels Wholly Male Sterile 30% Percent of Tassels with some Functional Pistillate 100% NICKERSON & EMBLER GIBBERELLIC ACID TREATMENT I 239 Treatment in pp m of GA 5 25 125 625 Height (nearest cm.) ,01 87 (3 plants 46 of Plants M Number of Nodes 14.2 13.4 15.2 13.0 Insuff. No. of Plants Number of Tillers 4.2 3.0 1.2 Insuff. No. Number of Ears 7.6 7.6 5.8 Insuff . No. *1 Number of Primary Tassel Branches 22.4 21.6 20.8 Notns sels exserted i Percent of Tassels Wholly Male Sterile Q No tassels exserted Percent of Tassels with some Functional Pistillate No tassels exserted Original Size of Population in Each Treatment Group 23 22 22 22 22 Number Surviving 23 17 14 5 1 Number of wind-damaged plants among survivors 3 2 Number of plants dying from 5 8 17 21 240 ANNALS OF THE MISSOURI BOTANICAL GARDEN f MEASUREMENTS Treatment of GA in ppm. „ 5 25 123 025 New York Flint 24.0 31.8 27.5(4)* 31.0(2) 10.0(4) J y § Gourdseed 24.0 23.6 24.8 21.2 18.0 Cherokee Deo, 2..o(3 23.0 28.2(4) 24.0(4) IMO) Argentine Pop .7.. IM14 19.4 " ~ 4i New York Flint n.4 10.2 15.2 14.8(4) 13.2(4) Gourdseed .0.4 12.4 12.8 .4.0 „., Cherokee Dent 15.6 17.4 16.8 16.2 18.3(3) ,.. 8.0 7.8 " - h i New York Flint 17.2 24.2 28.2 23.8(4) 21.8(4) Gourdseed 19.0 1M ,7.4 18.6 11.8 Cherokee Dent 25.. 21.0 24.0 22.4 14.0(3) Argentine Pop ,.0 2.8 2.8 " - 1! New York Flint ,5.4 42.0 42.7 43.8(4, 23.2 Gourdseed 34.4 36.0 37.0 35.2 ,0.4 m Cherokee Dent 42.2 40.4 43.0 40.2 37.9(3) h 26.0 27.6 27.2 " - ANNALS OF THE MISSOURI BOTANICAL GARDEN Argentine Pop. Plant on right treated with 125 ppm GA each three days for entire growing season. Note ladder-like effect of leaves joined at tips. Bulge at top containing small ear photographed at extreme left; top photo is dry ear, while lower photo is same ear gently boiled in water and partly dissected. Note male spikelets at top, paired female spikelets (2 silks at base of lower left-hand photograph indicate one pair) and lack of tassel branches. Well-developed cupules are also found on ked C indicates one in oblique view. Plant in center is contorted culm and leaves plant in group receiving 625 ppm GA. Brace roots were thin, brittle and poorly developed on both plants. Black and white checked ruler divisions are 1 cm. each; fine J I v: ■ i , . li.s.i .,i ■: I. STUDIES INVOLVING SUSTAINED TREATMENT OF MAIZE WITH GIBBERELLIC ACID II: RESPONSES OF PLANTS CARRYING CERTAIN TASSEL-MODIFYING GENES* Introduction Phinney (1956) showed that dwarf -1, a recessive maize mutant, responded to Gibberellic Acid (hereinafter called GA) in such a way as to become phenotyp- ically normal with a total amount of 60 micrograms of GA applied regularly during the growing season. Recently (Nickerson, 1960a) it has been shown that the dominant maize genes Corn-grass (Cg) and Teopod (Tp) likewise respond to sustained GA treatments in a like manner, becoming essentially normal in pheno- type. It has also been shown previously (Nickerson, 1959; Nickerson and Embler, 1960) that normal maize plants respond to sustained treatment with GA in several characteristic ways. Inasmuch as some of the growth manifestations noted under such GA treatment resembled some known tassel mutants, it was decided to grow obtainable seed of a series of tassel mutants previously studied (Nickerson and Dale, 1955) and subject them to various strengths of GA treatment throughout their growing seasons. The maize mutants 1 Tassel-seed 2 (t$ 2 ), Tassel-seed 5 (Ts 5 ), Tassel-seed 6 (Ts G ), Vestigial glume (V#), Tunicate (T«), Ramosa-1 (ra x ) and Male-sterile 1 (ms u later shown to be identical to tassel-seed 8 of Nickerson & Dale, 1955) were grown, treated and studied. Materials and Methods Planting distances and cultural methods employed are the same as those set forth in an earlier account (Nickerson and Embler, 1960). There were generally 2 or 3 lots of seed available for each of the above-listed mutants. Each of the stands of plants resulting from these seed lots was divided into four groups with as nearly equal numbers as possible; all plants in a particular group received the Concentrations of GA chosen were in the range shown by a previous study (Nickerson, 1959) to have a detectable but not drastic effect on field-grown maize plants. The four treatments employed were as follows: 1 — distilled water (controls) 2 — distilled water with 50 ppm GA * This work was supported by a grant from the National Science Foundation. 1 Thanks are hereby gratefully extended to Dr. Earl B. Patterson, Department of Botany, Uni- • aiding seed from Maize Genetics Cooperation sources for 244 ANNALS OF THE MISSOURI BOTANICAL 3 — distilled water with 100 ppm GA 4 — distilled water with 150 ppm GA Each third day for the duration of treatments one ml. of the appropriate solution containing either distilled water or water plus the above-listed amounts of GA 2 was applied from a pipette into the apical cavity of each plant. Solutions were freshly made each time, or stored no longer than three days in darkness at 19° C. Planting dates, beginning treatment dates, ending treatment dates, total numbers of plants grown and total amounts of GA applied to each mutant are shown in Table I. Tassel-seed 2 (ts 2 ). Seed stocks employed had no normal 6ibs. Because of the extreme general uniformity of plants wilhin each treatment group, measurements were made only on four single treated plants exhibiting the mutant form (Table II). Central spike length was generally decreased with increasing dosage, as were the number of primary tassel branches, the area over which they originated, and the number of ears produced. Peduncle length was about equally decreased by all three dosage levels of GA. Internode number was hardly influenced at all. Plant height showed increase with GA treatment; each group of treated plants in the field was highly uniform in total plant height, but internode diagrams (Fig. 1) show that early-formed internodes were increasingly lengthened by increasing doses of GA, while later-formed ones became much shorter than those of the distilled-water-treated mutant plant. Specimen tassels show a rapid decrease in overall size, but proportions of peduncle length, branch number, spikelet number and caryopsis development remain relatively constant (Plate XII). Tassel-seed 5 (Ts 5 ). Plants were either normal (+/+) or heterozygous (Ts 5 / + ) in genetic constitution. Figures listed under "M" on Table II are averages for 4 mutant (Ts 5 /-f-) plants; figures listed under "C" are measurements for the single control (+/-}-) plant in that group. Length of central spike was depressed solely by doses of 100 ppm GA in mutant-carrying plants; only slight changes were manifested in normal sibs. Tassel branch numbers fluctuated with no definite pattern in treated mutant-carrying plants; a consistent reduction was noted by all strengths of GA employed on normal sibs. The length of stem over which tassel branches developed was relatively consistent for both mutant- carrying and normal sibs regardless of GA concentration employed. Peduncle length of normal sibs was relatively uniform, but 50 ppm doses of GA tended to shorten this length somewhat in mutant-carrying plants, while 150 ppm doses tended to lengthen peduncles by about the same amount. Internode number increases consistently in mutant-carrying plants with increasing dosage; in normal sibs the number is uniformly increased by all GA levels employed. Ear number is increased in both mutant- carrying plants and normal sibs by doses of 50 ppm GA; further increase in GA dosage level does not decrease ear number in mutant-carrying plants, but drastically does so in normal sibs. Plant heights of mutant-carrying plants, nearly equal to those of normal sibs t n ThC ^ A emp,0yed w " kindl y wppKed by Dr. Curt Leben, Agricultural Research Division, Eli NICKERSON GIBBERELLIC ACID TREATMENT II 245 AT* v^X, Figs. 1-3. Internode dia under control (H 2 0) doses, are not as sensitive as those of normal sibs until the GA dosage reaches 150 ppm. Analysis of internode diagrams show that peduncle lengths in normal sibs are as long or longer than the longest lower internodes. The consistent dip below the peduncle and above the ear is present in all cases. "With increase in dosage, early-formed internodes become longer. Mutant-carrying plants 246 ANNALS OF THE MISSOURI BOTANICAL GARDEN show a different form from normals under control (H 2 0) treatments; with increasing doses of GA, it is the internodes above the ear and below the peduncle which become the longest. Plate XIII, No. 1 shows both control plants (left of ruler) and plants treated with 150 ppm GA doses (right of ruler). Normal sibs are above and mutant- carrying sibs are below. Increased peduncle lengths may be seen on both treated plants. Reduction in tassel branch number in normal sibs is prominent; branches on treated plants are also more loosely organized. More female florets were developed on GA-treated mutant-carrying sibs than on untreated controls, but GA did not stimulate formation of caryopses in normal sibs. Tassel-seed 6 (Ts 6 ). Plants were either normal (+/+) or heterozygous (Ts 6 / -)-) in genetic constitution. As with the foregoing, "M" in Table II refers to averages of measurements made on 4 mutant individuals while "C" refers to values for a single normal plant under the same treatment condition. Central spike length was not greatly decreased by any GA treatments in either mutant or normal forms. Tassel branches, however, were consistently increased in the mutant with increasing GA concentration and at the same time decreased in normal plants. Branching area was increased in mutants with increased GA doses, but did not vary greatly in normal plants. Peduncle lengths were greatly modified by lower doses of GA, but the high level (150 ppm) approached the average of control mutants. In normals, the 100 ppm dosage produced the most noticeable depression in length, a fact also consistent with the behavior of normal sibs of Ts 5 . Ear number was in general reduced by treatment with GA, but only the highest dose (150 ppm) affected the normal sib in this way. More internodes on mutant plants seem to have elongated with GA than without it, but no consistent trend could be noticed in normal sibs. Plant heights of mutants were increased with increasing GA dosage; doses of 150 ppm increased total height by 50%. By contrast, normal sibs with the same dosage increased only by 11%. Internode diagrams (Fig. 1) show that with distilled water, lower internodes are about the same in length and general gradual increase in both mutant-carrying and normal sibs; it is the internodes above the ear which account for their differences in height. 50 ppm treatments seem to stimulate all internodes about equally, except for a reduction in peduncle length of the normal sib. 100 ppm of GA has a greater stimulation of lower internodes but apparently tends to shorten all upper internodes. 150 ppm doses on both mutant-carrying and normal sibs show a marked effect on the early internodes with a peak length reached below the ear; each internode is then shorter up until the peduncle, which although long, does not exceed the longer internode. A com- parison between these last curves shows a considerable amount of similarity. Plate XIII, No. 2 shows tassels from mutant-carrying and normal sibs. Those on the left of the ruler received treatment with distilled water; those on the right with 150 ppm GA. Peduncle growth in mutant-carrying plants is notable, reduc- NICKERSON — 2 25 190 216 1.2 M = r^/ra, 11 21 64 IS 13 23 22 155 170 2.7 C - +/«! 50 9 2S 63 10 12 23 28 196 222 1.7 (single plant 100 15 3 39 14 10 24 11 213 256 1.7 mlntsT" .50 11 2' 25 " 20 25 » 236 229 11.5 1.2 (av. of 4) C = +/m Sl (single plant 15 22 10 14 9 15 19 166 202 12.5 1.7 50 21 21 8 16 10 14 21 181 200 14.5 1.7 100 13 2 11 14 8 12 15 18 199 205 ments)"" 150 17 16 8 7 8 9 13 17 199 219 1.7 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN Explanation of Plate PLATE XII Bot. Gard., Vol. 47, 1960 NICKERSON-GIBERELLIC ACID TREATMENT 1 XICKF.RSON— GIBERELLIC ACID TREATMENT II NICKERSON GIBBERELLIC ACID TREATMENT II iS els of Ts 5 /+ (lowe, (2) maize plants. T OF THE MISSOURI BOTANICAL GARDEN PLATE ] Tassels of Vg/+ (lower) and +/+ lants. Those left of ruler were treated w 50 ppm GA. ,-^r^ «, i • "" r \ ■"#* NICKERSON— GIBERELLIC ACID TREATMENT I sTICKERSON GIBBERELLIC ACID TREATMENT II 261 PLATE XV Volume XL VII Annals of the Missouri Botanical Garden so ■■■.;«■:■■• .-..:<■ --o Annals of the FLORA OF PANAMA ROBERT E. WOODSON, Jr. AND ROBERT W. SCHERY AND l PART IV Fascicle 3 LORANTHACEAE (C. T. Rizzini) OPILIACEAE (L. I. Nevling, Jr.) OLACACEAE (L. I. Nevling, Jr.) BALANOPHORACEAE (L. I. Nevling, Jr.) ARISTOLOCHIACEAE (H. W. Pfeiffer) POLYGONACEAE (J. A. Duke) Annals OF THE Missouri Botanical Garden FLORA OF PANAMA Part IV. Fascicle 3* LORANTHACEAE Vines, undershrubs, shrubs or rarely trees, always parasitic on roots or stems; the vines, less commonly the shrubs, bearing adventitious roots united with their hosts. Leaves usually opposite, sometimes alternate, entire, seldom reduced to scales, more or less coriaceous and inconspicuously nerved, the nervation either palmate or pinnate; without stipules. Flowers very large to minute, frequently showy because of the bright colors, androgynous or unisexual, spicate, clustered, strobilate or arranged in ternations, these in racemes, spikes, corymbs, umbels and so on, rarely solitary, subtended by bractlets either free or coalescent, or bractless; perianth simple or double; caly cuius present or absent; tepals (perianth segments) 3-6 or none; stamens as many as the tepals, reduced to staminodia in the pistillate flowers, the filaments joined to the tepals; pollen grains variable, globose or tri- angular, the exine either smooth or granulose (in some small genera it is reticulate or aculeate) ; ovary formed by the receptacle or blossom axis (thus the above mentioned calyculus is the edge of the receptacle), 1- (rarely several) -celled, sterile and poorly developed in the staminate flowers; stigma capitate, wanting in the male flowers which, however, have a slightly reduced style. Fruit (indeed a pseudofruit since it is developed from the receptacle), so far as American species are concerned, a berry (a drupe in certain Old World forms), frequently showing a milky layer containing rubber and persistent perianth parts; seed solitary, naked, the endosperm obviously developed and fleshy, rarely lacking; embryo small, with 2 rather large cotyledons. The plants of this family are either completely (when they are leafless and without chlorophyll) or partly (when they bear normal foliage leaves) parasitic on other woody plants. The roots applied to the host become haustoria and through these part or all the food supply is secured from the latter. The few erect, terrestrial, tree-like forms live upon roots and their haustoria are underground. The family comprises about 38 genera and 1,400 species, mainly tropical and subtropical but occasionally found in temperate regions. It is very homogeneous, 264 ANNALS OF THE MISSOURI BOTANICAL GARDEN yet the morphological types are well marked so that the New World forms are readily distinguishable from the Old World ones. The Central American species are more closely related to the South American than to the North American or West Indian ones. The following artificial key to genera serves only for the Panamanian entities so far known, since it has been prepared with the purpose of making the generic 1. GAIADENDRON G. Don Gaiadendron G. Don, Gen. Hist. 3:431. 1843; Eichler, in Mart. Fl. Bras. 5, pt. 2: 47. 1868 (as a subgenus of Phrygilanthm) . Tall shrubs or small trees living upon earth, but parasitizing roots of other plants. Leaves opposite, more or less coriaceous, almost always minutely, but evidently, punctate beneath. Flowers conspicuous, colored, 4-8-merous, perfect, in ternations, these in racemes or corymbs; bractlets conspicuously developed, free, persistent; pedicels often unequal. Pollen grains 3 -angled, the exine granulose. Ovary said to be 2-6-celled, but this scarcely convincing. Fruit baccate, the seeds albuminous. A chiefly Andean genus of about 6 species. The following is the only known from Central America, where it is endemic. A careful re-examination of the ovary, living or fixed, will probably show that the genus does not essentially differ from Phrygilanthus and that it must be retained, as by Eichler, under the last mentioned flora of Panama (Loranthaceae) 265 1. Gaiadendron poasense Donn. Sm. in Bot. Gaz. 56:61. 1913. Parasitic shrubs or short trees reported to reach a height o£ 3-6 m., growing on the ground, with erect, terete branches. Leaf blades slightly ovate or oblong, acutish at the base, attenuate to the blunt apex, minutely punctate beneath, fleshy- coriaceous, often drying revolute at the margins, nerveless, to 8 cm. long and 4.5 cm. wide; petioles 5-10 mm. long. Racemes axillary, solitary, 7-10 cm. long. Bractlets ovate, persistent, foliaceous, 5-12 mm. long. Flowers in ternations, the ANNALS OF THE MISSOURI BOTANICAL GARDEN lort-pedicelled, androgynous, about 15 mm. long; tepals 6—7 in number, in length. Ternations pedicellate, the pedicels about 4—6 mm. long. i Rica, Panama. cinity of Cerro Punta, 2. ANTIDAPHNE Poepp. et Endl. Antidaphne Poepp. et Endl. Nova Gen. ac Spec. 2:70. 1838. Stachyphyllum Van Tieghem, in Bull. Soc. Bot. France 42:565. 1895. Medium-sized, erect shrubs, without adventitious roots in the shoots. Leaves alternate, thickly coriaceous. Flowers minute, spicate, dioecious; spikes short (up to 10 mm. in length), sessile in the leaf axils, more or less strobiliform; bractlets scale-like, broadly imbricate, either caducous or persistent in anthesis. Staminate flowers reduced to 3-6 stamens inserted at the base of the bractlets on a small disk, the filaments elongated, unequal; anthers erect, 2-4-celled, longitudinally dehiscent; pollen grains globose, with 3 pori, the exine conspicuously aculeate. Pistillate flowers sessile, in groups of 2-5 at the bractlet axils, bearing a perigonium; tepals well or poorly developed, adnate to the ovary; style none, but the stigma capitate. Fruit a round berry, the pericarp fleshy and possessing an evident milky layer. A distinctive genus amongst South American Loranthaceae, recognized at first glance by the minute, bracteate, compact and dioecious spikes bearing naked male flowers; moreover, another striking feature lies in the aculeate (echinate) round pollen grains, since no other genus in the family shows such a character. Most peculiar too is the geographical distribution of the 4 known species. A. viscoidea Poepp. et Endl., the commonest of all, is Andean; A. fendleri (V. Tiegh.) Engler occurs in Venezuela not rarely; A. amazonensis Rizz. and A. para- ensis Rizz., both more recently detected and described, are from the Amazonian forest. Thus the 4 species behave as being vicarious, coming down from the high Andean mountains to the low Amazonian basin, the habitats never intergrading with one another. Keeping in mind the aforesaid statement, it is somewhat surprising to find A. viscoidea in Central America (Costa Rica, Guatemala and Panama) but— it must be stressed— at high altitudes (from 1,400 m. upwards). 1. Antidaphne viscoidea Poepp. et. Endl. Nova Gen. ac Spec. 2:70. 1838. A densely branched shrub, glabrous, the branches generally to 50 cm. long, terete or nearly so. Leaves almost sessile, or the petioles very short and thick, obovate to suborbicular, cuneate at the base, the nerves very prominent and openly reticulate, coriaceous, 3-7 cm. long, 2-3 cm. wide. Flowers cream- colored, the Perianth segments slightly dentate. The bractlets fall when the flowers open. Fruit oval. (248) flora of Panama (Loranthaceae) 267 This species differs from the others mainly in having shorter spikes covered ith bractlets which are deciduous by the time of anthesis. The whole genus is jrked out in Rodriguesia 18-19:221-225, pi. 26-28. 1956. Steyermark mentions a collection (Fieldiana, Bot., 24. pt. 4:63. 1946) in 3. PSITTACANTHUS Mart. Always stout and erect shrubs, lacking aerial 1 coriaceous. Flowers large, showy, brilliantly colored, hexarr ternations or binations; these in racemes, corymbs or cup which subtends the flowers. Tepals either free c at the base. Stamens unequal, alternately shorter and longer, the filaments slender; pollen grains 3 -angled, flattened, the exine granulose. Stigma capitate. Fruit baccate, large for the group, the seeds without endosperm but bearing big, and often more than 2, cotyledons. About 80 species are known, especially in tropical America, 5 of them reported from Panama. The genus is easy of recognition because of the cups subtending the flowers, which are the largest among Loranthaceae, at least the New World ones. Another species that may well be expected is P. calyculatus (DC.) Don, from Mexico and found in Costa Rica, which is closely allied to P. chrismarii Urban but distinguishable by the longer, acute and not oblique leaves. 1. Psittacanthus scheryi Woods, in Ann. Missouri Bot. Gard. 28:426. 1941. A very stout shrub up to 1.8 m. tall. Branches terete, glabrous, very thick. Leaves widely ovate or suborbicular, the rounded base scarcely contracted to the petiole, fleshy-coriaceous, the nerves poorly developed, 9-11 cm. long, 6-9.5 cm. broad; petioles about 8 mm. long. Flowers orange below, deep yellow toward the tips, 2.5-3.2 cm. long; the 2-flowered peduncles fasciculately disposed, 3-6 mm. long; pedicels about 5 mm. long; calyculus with entire margin, 2.5 mm. deep; cup about as long as the calyculus; perianth segments slightly enlarged at the base, the filaments attached above the middle. Endemic. chiriqui: vicinity of Bajo Mona and Quebrada Chiquero, Woodson & Schery j8l (type). (249) ANNALS OF THE MISSOURI BOTANICAL GARDEN A slender and handsome shrub, drying olivaceous. Branches terete, glabrous. Leaves oblong or elliptic, attenuate both to the blunt apex and acute base, in youth more or less spatulate, firmly coriaceous, nerveless or nearly so, 4-7 cm. long, 1.5- 2.5 cm. wide; petioles 2-5 mm. long. Flowers geminate 2-6 in each axil, corymbosely disposed, scarlet, 2-3 cm. long; common peduncle about 8 mm. long, the pedicels twice as short; caly cuius and cup with entire margins, 2 mm. long; perianth segments lightly dilated at the base, the filaments medially inserted. Fruit green, ellipsoid, 6 to 12 mm. long. Endemic. canal zone: road to San Carlos, Harvey 5173. cocle: vicinity of El Valle de Anton, Allen igyg (type); in the same place, Allen 3702 and 2233; marshes along R. Anton, Hunter & Allen 371. 3. Psittacanthus ALLENH Woods. & Schery, in Ann. Missouri Bot. Gard. 27:309. 1940. In habit and color very similar to P. lateriflora Woods. & Schery, but the flowers are usually longer and broader and the tepals bear well developed ligules. Branches terete, toward the tips somewhat compressed. Leaves elliptic-obovate, rounded at the apex, cuneate to the base, coriaceous, almost nerveless, 4-7 cm. long, 2-3 cm. broad; petioles 2-3 mm. long. Flowers 2-6 corymbosely arranged, orange, 3-3.5 cm. long; common peduncle about 5-8 mm. long; pedicels 5 mm. long; caly cuius and cup with entire margins, 1-2 mm. long; perianth segments or tepals dilated at the base, showing conspicuous ligules; filaments about medially attached. Fruit Endemic. cocle: vicinity of El Valle de Anton, Allen 1223 (type) ; Penonome, Williams 526. P. lateriflorus and P. allenii are easily confused, and difficult to separate, at first glance. The ligule, which the latter bears, is to be seen as a basal outgrowth of the perianth segments; one must seek for it at the inner portion of the tepal base, which is dilated about 2 mm. upwards. Moreover, the leaves of P. allenii show a well-marked tendency to be broader toward the apex. Davidson's bad specimen no. 568 (Boquete, Chiriqui) differs from P. allenii in no wise but the leaf shape, which is lanceolate. 4. Psittacanthus chrismarh Urban, in Engl. Bot. Jahrb. 24:331. 1897. Psittacanthus americanus (L.) Mart, in Flora 13:108. 1830, as to the Mexican and Central Plants erect, the branches quadrate and glabrous. Leaves subsessile (petioles 0-3 mm. long), obliquely lance-elliptic or oblong, rarely ovate, narrowed both to the blunt apex and acute base, thick and hard, faintly 3-6-nerved, 7-12 cm. long, 2-4 cm. wide. Inflorescences densely flowered, both axillary and terminally (250) FLORA OF PANAMA (Loranthaceae) 269 racemose or paniculate, 8-15 cm. long; common peduncle to 2.5 cm. long; pedicels about 1.5 cm. long; cups 1-2 mm. deep. Buds conspicuously clavate-apiculate, to 4 cm. long. Flowers red, 3-5 cm. long, in ternations; calyculus denticulate at the margin, 3-4 mm. long; perianth segments enlarged at the base, the filaments about medially adnate. chiriqui: El Boquete, Maxon 538. cocle: vicinity of Las Uvas, Allen 2796; Agua- tl r 4836. herrera: Pese, Allen 806. Panama: vicinity of San Carlos, Allen, 1 130, Harvey 5172. P. chrhmarii Urban has always been confused with P. americanus (L.) Mart., from the West Indies; nevertheless, as Urban pointed out, it is easily distinguished by the obviously 4-angled branches. (251) 270 ANNALS OF THE MISSOURI BOTANICAL GARDEN 5. Psittacanthus schiedeanus (Cham. & Schl.) Bl. in R. & S. Syst. 7, pt. 2: 1730. 1830. Loranthus schiedeanus Cham. & Schl. in Linnaea 5:172. 1830. Stout plants with thick, 4-angled, compressed branches, the branchlets often wing-margined. Leaves falcately ovate, rarely oblong, acute, rounded at the base, coriaceous, the nerves slightly impressed, to 13 cm. long, about 3-5 cm. wide; petioles 1-1.5 cm. long. Inflorescences corymbose, terminal, very densely flowered, up to 20 cm. long; common peduncle to 4 cm. long; pedicels about 2 cm. long; bracts well developed, 5-12 mm. long; cups mostly 2 mm. deep. Buds slightly clavate, briefly apiculate. Flowers yellowish-orange, 5-9 cm. long, in ternations; calyculus irregularly denticulate, 5 mm. long; perianth segments not enlarged at the base, the filaments medially attached. Mexico and Central America. chiriqui: Finca Lerida to Peiia Blanca, Woodson & Schery 308; R. Chiriqui Viejo, G. White 76 and P. White 331, 42; Casita Alta, Volcan de Chiriqui, Woodson, Allen 2? Seibert 915; vicinity of Bajo Mona and Quebrada Chiquero, Wood* 1 -j $cbery $96; trail from Cerro Punta to headwaters of Rio Caldera, Allen 1444. 4. PHRYGILANTHUS Eichl. Phrygilanthus Eichl. in Mart. Fl. Bras. 5, pt. 2:45. 1868. Shrubs or small trees living both on earth, as root parasites, and upon other woody plants. Leaves generally leathery, opposite, rarely reduced to scales. Flowers typically hermaphrodite, hexamerous, seldom 4-8-merous, showy, either solitary or arranged in ternations, these in racemes, corymbs or umbels; bracts and bractlets moderately developed, both persistent and caducous according to the species group. Stamens unequal. Pollen grains 3-lobate, the exine granulose. Ovary surrounded by a well-developed, fleshy disk. Fruit a juicy berry, the seeds The genus embraces about 30 species thriving primarily in South American highlands. Only a few have been gathered in Mexico, Central America and Australia. As said before, this genus is poorly distinct from Gaiadendron (see both figures) and they possibly will later be joined into one. 1. Phrygilanthus panamensis Rizz., n. sp. Secunda stirps ad Sect. Martiella (v. Tiegh.) Engl, attribuenda. A prima species usque adhuc cognita, Ph. pdmeri (Wats.) Engl, in Mexico vigente, primo vultu floribus racemosis atque foliis caudatis discernitur, neglecta florum magnitudine. Frutex parasiticus ramis elongatis teretibusque, ad nodos modice incrassatis, internodiis 2-4 cm. longis. Folia ovato-caudata, basi subrotundata, e medio apicem versus gradatim attenuata, extremo apice valde angustata, coriacea, nervis supra tantum subtiliter impressis, 5-9 cm. longa, 3-5 cm. lata; petiolis 1-2 cm. longis. Racemi vulgo 2 ad singulas axillas, longitudine ad 4 cm. usque, laxi, rachidi (252) flora of Panama (Loranthaceae) quadrangula; floribus oppositis, neque in ternationibus aggregatis et androgynis. Pedicelli circa 1-2 mm. longi, teretes. Bractea trangulari-acuta, 2 mm. longa; bracteolae duae pro flore, liberae a bractea et hac duplo minores, conformes. Ala- bastra cylindrica, apice optime apiculata, siccitate nigrescentia. Calyculus ad marginem membranaceum 0.5 mm. altum tantum reductus. Perigonium hex- amerum, ad summum 1 cm. longum, caeterum ut in aliis generis speciebus. Fructus latent. chiriqui: Bajo Chorro, Boquete, alt. 1800 m., M. E. Davidson 431 (US, holotype; F, isotype). On vegetative characters alone this remarkable plant resembles Struthanthus marginatus, with which it has been confused; however, it suffices to observe the pointed buds as well as the solitary flowers at each pedicel of Ph. panamensis to make ready distinction. The only other species of the Section Martiella is Ph. pdmeri (Wats.) Engl., whose leaves rounded at the apex together with its red flowers, to 4 cm. long- not to mention the inflorescence (see Latin diagnosis)— place it far from the above. 272 ANNALS OF THE MISSOURI BOTANICAL GARDEN 5. STRUTHANTHUS Mart. Struthanthus Mart, in Flora 13:102. 1830; Rizzini, in Rev. Brasil. Biol. 10 (4) : 393. 1950. Shrubs, either erect and rootless or scandent and supplied with adventitious roots, rarely vines. Leaves usually coriaceous. Flowers small, pedicelled or sessile, 5-6-merous, in ternations; these in racemes, panicles, corymbs or spikes, some- times umbellate; bractlets coalescent or free, persistent or deciduous. Stamens unequal, the filaments slender. Pollen grains of two types: fertile 3-lobate, the exine granulose; sterile globose, the exine smooth. Pistillate flowers with stami- nodia; stamina te ones with reduced style, this lacking stigma. Buds of the staminate flowers clavate. Stigma capitate. Fruit a berry, the seeds with endosperm. The * Only a few t S. Syst. 7, pt. bus polystachyus R. & P. Fl. ] An erect shrub lacking adventitious roots, as much as 2 m. high (usually shorter). Branches terete, the branchlets rather compressed. Leaves broadly ovate or almost oblong, acuminate, rounded at the base, veined on both sides, more or less coriaceous, cartilaginous-margined, to 15 cm. long, 7 cm. broad; petioles 1-2 cm. long. Ternations completely sessile. Spikes solitary or 2-3 together, 3-5 cm. long. Inside the foliar axils there are many imbricate scales (prophylls) enclosing the base of the spikes. Bractlets soon caducous and so not detectable. Flowers yellow, 3-4 mm. long. Fruit oval, blue, 5-6 mm. thick; fruiting spikes rather longer and thicker than the flowering ones, to 7 cm. long. South and Central America (however, not yet found in Brazil). An easily recognizable species. bocas del toro: von Wedel 480; Rio Cricamola, Woodson, Allen tf Seibert 1911. cocxe: north of El Valle de Anton, Allen 3721; mountains beyond La Pintada, Hunter d Allen 6 2 4 . Alnnrante, Cooper 20 5 ; Bismarck, Williams 5 6l. colon: along Rio Fato, flora of panama (Loranthaceae) 273 2. Struthanthus marginatus (Desr.) Bl. in R. & S. Syst. 7, pt. 2:1731. 1830. Lorcmthus marginatus Desr. in Lam. Encycl. 3 :596. 1789. A scandent shrub provided with aerial roots. Branches somewhat compressed and lenticellate-dotted. Leaves narrowly ovate or lance-ovate, conspicuously acuminate, rounded-obtuse at the base, firmly leathery, finely nervose on both surfaces, cartilaginous-margined, 7-12 cm. long, 3-5 cm. wide; petioles 5-10 mm. long. Ternations pedicelled. Racemes 1-3 in each axil, 3-8 cm. long. Bractlets persistent, connate. Flowers yellowish green, 3—4 mm. long. Fruit yellow or orange, 8-10 mm. long. Widespread in Brazil. Costa Rica, Panama. bocas del toro: vicinity of Chiriqui Lagoon, von Wedel 1 28 J. chiriquI: Callejon Seco, Volcan de Chiriqui, Woodson & Schery 486 and 504; vicinity of Casita Alta, Volcan de Chiriqui, Woodson, Allen & Seibert 808 and 932; Rio Chiriqui Viejo valley, Cerro Punta, G. White 34; Finca Lerida to Peiia Blanca, Woodson & Schery 292; Boquete, David- son 859; Bajo Chorro, Boquete, Davidson 392, 908. 3. Struthanthus orbicularis (H. B.K.) Bl. in R. & S. Syst. 7, pt. 2:1731. Loranthus orbicularis H.B.K. Nov. Gen. & Sp. 3:434. 1820. Vines bearing adventitious roots, with long and flexuous branches, the branch- lets twining, quadrate; the young leaves seem to serve as prehensile organs (they show petioles fully developed while the blades remain much smaller than the normal ones). Leaves broadly obovate to rounded, sometimes perfectly orbicular, coriaceous, rarely veiny, to 5 cm. long, 4 cm. wide; petioles 1-1.5 cm. long. Bractlets free, soon caducous. Ternations sessile. Flowers pedicelled, yellow or greenish yellow, 5-7 mm. long. Spikes solitary, rather stout (especially when fruiting), the rachis sharply angled, 7-15 cm. long. Fruit elliptic, purple to red- brown, about 1 cm. long. Peru, Ecuador, Colombia. Rare in Brazil. Central America, where it is more widely dispersed. bocas del toro: Rio Cricamola, Woodson, Allen & Seibert 1908; Chiriqui Lagoon, von Wedel 2663, 1065. canal zone: White T2I; vicinity of Ancon, Piper 6019; Balboa, Standley 29327; Bellavista, Macbride 2754. COCLE: vicinity of El Valle, Allen 1803; Seibert 417; vicinity of Sta. Clara beach, Woodson, Allen & Seibert 1700. Panama: Woodson, Alio, f5 Seibert r<;il; Las Sabanas, Standley 25936; San Jose Island, Johnston tr 1938. PROviNci unknown: Quebrada de Oro, Wheeler ef Zetek 12-III-1923. 4. Struthanthus rotundatus Rizz. in Rev. Brasil. Biol. 10 (4): 401. 1950. Resembling S. orbicularis in habit. Branches terete, the branchlets com- pressed. Leaves perfectly elliptic or nearly rounded, slightly attenuate at the base, almost nerveless, 4-6 cm. long, 3-4.5 cm. wide; petioles 5-15 mm. long. Ternations pedicelled. Bract- lets connate, persistent. Racemes solitary, 5-10 cm. long, the rachis compressed. Flowers deep yellow, about 7 mm. long. Fruit unknown. Brazil, Panama. 274 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 75. Struthanthus rotundai flora of Panama (Loranthaceae) In general appearance, especially by its leaves, this species is very similar to S. polyrhizus Mart, too, but differs as follows: the former has racemes as long as the leaves or longer; the latter shows corymbs half as short as the leaves. So it may be understood why they have sometimes been confused. The type (see figure in Rodriguesia, 18-19, tab. 8, 1956) conserved at the British Museum leaves no doubt as to the identity of Brazilian and Panamanian material. 6. PHTHIRUSA Mart. Phthirusa Mart, in Flora 13:110. 1830; Rizzini, in Dusenia 3 (6) : 451. 1952. Erect or scandent shrubs, the latter developing aerial roots. Leaves mostly leathery. Flowers often very tiny, tetra-hexamerous, in ternations; these in racemes, spikes, panicles or rarely subsolitary; bractlets connate. Stamens unequal, the filaments of the longer scalloped at both sides by pressure of the anthers of the shorter ones. Pollen grains of one type; triangular in shape, the exine finely granulose, showing three pores. Pistillate flowers with staminodia; staminate ones with vestigial style, this lacking stigma; androgynous flowers sometimes occur. Buds of the male flowers terete. Fruit baccate as in Strutbanthus. A genus of about 5 5 species, widely distributed throughout South America and the West Indies. It differs from the preceding by the combination of scalloped filaments, pollen grains of one type and terete male buds; thus, despite the opinion of Baehni and Macbride (Candollea 7:287-290. 1936-1938), it seems best to maintain it as a genus ( see also Rodriguesia 18-19:87-234. 1956). The following are the only Central American species. 1. Phthirusa adunca (Meyer) Maguire, in Bull. Torrey Bot. Club 75:301. 1948. Loranthus aduncus Meyer, Prim. PL Esseq. 149. 1818. Loranthus paniculate H.R.K. Nov. Gen. & Sp. 3:422. 1820. Loranthus co,' J At. 1820. Loranthus magi Linnaea 3 :219. 1828. Loranthus theobromae Willd. ex R. & S. Syst. 7:132. 1829. Strutbanthus aduncus (Meyer) G. Don, Gen. Syst. 3:414. 1834. Phthirusa theobromae (Willd. ex R. & S.) Eichl. in Mart. Fl. Bras. 5, part 2:56. 1868. Phthirusa paniculata (H.B.K.) Macbr. in Field Mus. Publ. Bot. 11:17. 1931. Scandent shrubs bearing adventitious roots. Branches terete, the branchlets more or less compressed. Leaves ovate or oblong, rounded at the base, gradually distinctly nervose, leathery, 4-7 cm. long, 2.5-4.5 cm. broad; petioles (257) 276 ANNALS OF THE MISSOURI 1-2 cm. long. Flowers pale yellow, 4-6 cm. long, paniculate, pedicelled (pedicels 1-3 mm. long, rarely none) ; filaments provided with many glandules at the angles. Fruit oblong, pedicellate, orange, to 1 cm. long. Brazil, where it is widely distributed, Peru, Venezuela (also very common there), Guianas, Paraguay, Panama. South American specimens are not different from Central American ones even in the least detail. bocas del toro: vicinity of Chiriqui Lagoon, von Wedel 2414. canal zone: vicin- ity of Miraflores Lake, G. White 158; Bellavista, Salvoza 1005. chiriqui: vicinity of Callejon Seco, Volcan de Chiriqui, Woodson & Schery 500. veraguas: vicinity of Santiago, Allen 1012. Ph. theobromae and Ph. magdalenae are not at all clearly distinct; as a matter of fact, examining the extremely rich South American material, numerous transi- tions are seen to occur from one to another. The other names above quoted are merely synonyms dealing with the priority question and deserve no more attention. 2. Phthirusa pittieri Krause, in Fedde, Repert. Sp. Nov. 15 (1-3) : 441. 1917. Known only from original description which is, however, very complete. Branches terete. Leaves narrowly ovate or ovate-oblong, long-acuminate, rounded- obtuse at the base, 12-18 cm. long, 5-9 cm. broad. Ternations in axillary spikes, these numerous in each axil. Flowers 5 mm. long. colon: in the margins of Rio Fato, Pittier 391 1. 3. Phthirusa pyrifolia (H. B. K.) Eichl. in Mart. Fl. Bras. 5, pt. 2:36. 1868. Loranthus Pyrifolius H.B.K. Nov. Gen. & Sp. 3 :441. 1820. An erect shrub without aerial roots. Branches greatly flattened, the tips reddish-scurfy. Leaves ovate or elliptic, nearly rounded at the base, obtuse toward the apex, now and then mucronate or even caudate, distinctly veiny on both sides, to 10 cm. long, 5 cm. wide; petioles about 1 cm. long. Ternations perfectly sessile. Flowers very minute, 1-2 mm. long. Spikes axillary, solitary or 2 in each axil, unbranched, reddish-furfuraceous, 5-10 cm. long. Fruit oblong, 4-6 mm. long. Panamanian specimens available agree exactly with Brazilian ones. Venezuela, Guianas, Colombia, Costa Rica, West Indies, etc. canal zone: near Vigia and San Juan on Rio Pequeni, Dodge, Steyermark & Allen 1 6571; around Las Cruces, Pittier 2623; Sutton Hayes W22; without locality, Woodworth & Vestal 493; Gatiin Lake, Bangham 441; Changuinola Valley, Dunlap 32 J. 7. ORYCTANTHUS (Griseb.) Eichl. Oryctanthus Eichl. in Mart. Fl. Bras. 5, pt. 2:87. 1868. Loranthus sect. Oryctanthus Griseb. Fl. Brit. West Ind. Isl. 311. 1860. Erect shrubs, almost always with aerial roots starting from the base of the stem. Leaves opposite, coriaceous. Flowers minute, perfect, 6-merous, more or less immersed in the rachis of the spikes; bractlets poorly devel Pollen grains strongly reticulate. Stigma capitate. Fruit ba< About 1 8 species, uniformly distributed throughout the American tropics. The nus is noteworthy for its double-perianthed flowers sunk in the axis of the 1. Oryctanthus cordifolius (Presl) Urban, in Engl. Bot. Jahrb. 24:348. 1897. Viscum cordifolium Presl, Epim. Bot. 253. 1849. A stout shrub with terete branches, the branchlets compressed, scarcely rusty- furfuraceous, becoming glabrate. Leaf blades sessile, broadly ovate, cordate at the rounded base, attenuate to the blunt apex, firmly coriaceous, nerves distinct but faint, about 7-12 cm. long, 4-8 cm. wide. Spikes axillary, solitary, to 8 cm. long, the peduncle 5-1 5 mm. long. Bractlets with free margins. Flowers or buds (259) ANNALS OF THE MISSOURI 2 mm. long, inserted obliquely on the glabn Central America. . toro: Isla Colon . rSjo. si 9 6. von Wedel 5/ 2; vicinity of Nievecita, Woodson, Quebrada Tranquila, Dodge gf Allen 17332. ia: near Matias Hernandez, Standley 28923. 2. Oryctanthus occidentalis (L.) Eichl. in Mart. Fl. Bras. 5, pt. 2:89. 1868. Loranthus occidentalis L. Syst. ed. 10. 988. 1774. Erect shrubs. Branches, terete, the branchlets slightly compressed, petioles (to 5 mm., long) and peduncles (5-10 mm. long) rusty-furfuraceous, later sub- glabrate. Leaf blades very broadly ovate, almost orbicular at the base, scarcely constricted to the apex, this obtuse, thick and hard, nervation distinct or veinless, 3-10 cm. long, 2-8 cm. wide. Spikes solitary, to 3 cm. long; rachis glabrous. Bractlets with distinct margins. Flowers or buds becoming brown, 1-1.5 mm. long, perpendicular to the rachis. Fruit ovate, 4-5 mm. long, the apex not exceeded by the margin of the persistent calyculus. West Indies, Costa Rica, Panama. Fig. 77. Oryctanthus flora of Panama (Loranthaceae) 279 canal zone: Juan Mina, Piper 5703; around Las Cruces, Pit tier 2622; Las Cascadas Plantation, Standley 25679; Santa Rita Trail, Cowell 74; without locality, Wood-worth & Vestal 492; Chagres, Fendler 135-, Barro Colorado Island, Shattuck 700. 3. Oryctanthus spicatus (Jacq.) Eichl. in Mart. Fl. Bras. 5, pt. 2:89. 1868. Loranthus spicatus Jacq. Enum. PL Carib. 18. 1760. An erect shrub 50-70 cm. tall. Branches terete, the slightly compressed tips, with petioles (1-5 mm. long) and rachis, reddish-scurfy but becoming more or less glabrous. Leaf blades ovate, sometimes orbicular-ovate, rounded at the base, moderately attenuate to the blunt apex, thickly leathery, lightly nerved (rarely nerveless), to 9 cm. long, 5 cm. broad. Spikes solitary, to 2 cm. long, stouter than in the preceding; peduncles to 5 mm. long. Bractlets not distinct from the rachis. Flowers or buds 1 mm. long, oblique to the rachis. Fruit ovate-oblong, about 5 mm. long, the apex evidently exceeded by the margin of the persistent calyculus. "West Indies, South and Central America. canal zone: near Vigia and San Juan on Rio Pequeni, Dodge, Steyermark & Allen 16577; Ancon Hill, vicinity of Balboa, Seibert 380, 41O; Barro Colorado Island, Shattuck 590 and Wilson 151; Gatun Lake, Wetmore & Abbe 148; Woodworth & Vestal 494. CHiRiQuf: Gualaca, Allen 5025. cocle: Penonome, Williams 195. Panama: Tumba Muerto Road, Standley 29831 ; near Punta Paitilla, Standley 26249. The morphological facts by which O. occidentalis differs from O. spicatus are minute and tedious ones. Notwithstanding, and worth noting, additional characters may be found in the spikes: in the first species these are slender, blackish and supported by a peduncle longer than 5 mm.; in the second, the spikes are stouter, reddish-brown and short-peduncled or even sessile. 8. DENDROPHTHORA Eichl. Dendrophthora Eichl. in Mart. Fl. Bras. 5, pt. 2:102. 1868; Urban, in Engl. Bot. Jahrb. 24:370. 1897. Erect shrubs, more or less yellow, lacking adventitious roots. Leaves opposite, small or wanting. Cataphylls poorly developed. Flowers trimerous, very minute, dioecious, deeply immersed in depressions in the rachis of the spikes. Anther cells confluent, dehiscing transversely; pollen grains globose, the exine smooth. Pistillate flowers without staminodia. Staminate flowers showing no style. Fruit a berry, the seeds with endosperm. A genus of about 32 species, thriving better in high-altitudinal places, chiefly in South America and the West Indies. It is very similar in appearance to Phora- dendron, from which may be distinguished as indicated in the above key, except that its species are less robust with leaves small or reduced to scales. OF THE MISSOURI BOTANICAL GARDEN in, in Ber. Deutsch. Bot. Gesell. An erect shrub. Branches terete, the branchlets somewhat compressed, striate- rugose. Leaves obovate-spatulate, long-attenuate to the base, the apex rounded and emarginate, veinless, moderately coriaceous, 2-3 cm. long, 1-1.5 cm. broad; petioles about 5 mm. long. Spikes solitary, axillary, 1-2-jointed, 5-15 mm. long, the peduncles 0-3 mm. long. Flowers 6-seriate, 18-40 in each joint. Fruit white, globose. Costa Rica, Panama. CHiRiQuf: Bajo Chorro, Boquete, David- son 369; upper belt of Chiriqui Volcan, 2. Dendrophthora biserrula Eichl. in Mart. Fl. Bras. 5, pt. 2:104. 1868. An upright, densely branched under- shrub attaining 25 cm. in height, the branches terete. Leafless. Scales about 1 mm. long. Spikes very numerous, axil- lary, solitary, 1 -articulate, to 2.5 cm. long with 4-24 flowers, these 2-seriate; the peduncles 5-8 mm. long; the rachis quadrate. Fruit globose, white. Central America. Volcan de Chiriqui, Boquete, Pittier 3073. This species is remarkable by support- ing usually a vigorous growth of fila- mentous as well as foliaceous lichens, which give it a peculiar appearance. 9. PHORADENDRON Nutt. Phoradendron Nutt. in Journ. Acad. Philadelphia ser. 2. 1:185. 1847; Trelease, The Genus Phoradendron (Urbana, Ills.: The University of Illinois, 1916). Erect and rootless shrubs. Leaves opposite, rarely reduced to scales. Flowers as in Dendrophthora, except that they are arranged in definite rows on the spike flora of Panama (Loranthaceae) 28 joints (2 or 3 series over each side of the joints). Anthers 2-celled, dehiscing b longitudinal slits; pollen grains globose, the exine smooth. Fruit as in th preceding genus. The New World mistletoe includes about 300 species. In the Old World i is replaced by Viscum. a. Cataphylls present on all internodes of the branches. (ranches 4-angled or flattened. Spikes most Shoots prevailingly flattened or 2-edged. SfK ::-:t di :\t: d marginate. joints 4- Leaves obov oblong-lanceolate or th upright tcpals. Joints nts Leaves lance-oblong or oblong. r-.: :w.'- clo>cl> connive nt. Phoradendron pergranu Trcl. Ann. Missouri Bot. Gard. 27:. A stout shrub cymosely branching at every node; shoots terete, much dilated and somewhat compressed below the nodes. Cataphylls a single basal pair on all joints, the margin almost fimbriate. Leaves falcately oblong, attenuate to both ends, rounded at the apex, acute at the base, nerveless, coriaceous, drying somewhat (263) 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN crisped marginally, 5-9 cm. long, 1.5-3 cm. wide; petioles scarcely 2-4 mm. long. Spikes solitary, mostly 4-articulate, about 3 cm. long; peduncle 3-5 mm. long; joints slender, 5-8 mm. long, 4-8 -flowered in 4 series. Fruit rounded, varnished, the tepals erect, about 3 mm. in diameter. Endemic. cocle: vicinity of El Valle, Allen 777. 2. Phoradendron robaloense Woods., n. sp. Nomen tantum in sched. Inter affines (praesertim P. henslovii Robs.) haec species distinctissima et statim discernenda vaginis cataphyllaribus in tubum amplum 3-10 mm. longum connatis, foliis spicisque neglectis. Fruticulus dioicus circa 2.5 m. altus, ramis dichotomis teretibus; internodiis 4-8 cm. longis. Vagina cataphyllaris 3-10 mm. longa, tubulosa, in dentes 2 latos rotundatos divisa. Folia obliqua, ovato-lanceolata, basi vel obtusa vel cuneata, apicem versus longe attenuata, imo apice ex indole cuta, crasse coriacea, siccitate marginibus undulata, obsolete palmatinervia aut avenia, 8-12 cm. longa, 2^.5 cm. lata; petiolis crassis, circa 1 cm. longis. Spicae 2-3 -nae in axillis, graciles, 3- 4-articulatae, 3-5 cm. longae; pedunculis 8 mm. longis; articulis plus minusve 1 cm. longis, 4-seriatim 6-14-floris; floribus femineis. Baccae elongatae, perigonio clauso coronatae, 5-6 mm. longae. CHiRiQUi: Bajo Mona, Robalo Trail, alt. 1500-2100 m. s. m., Paul H. Allen 4838 (MO, holotype). Ph. robaloense Woods, ex Rizz. is a very striking species and is easily distin- guished from its allies (Aequatoriales Cymosae) by the tubular cataphylls, the few-flowered spike joints and the elongated fruits. 3. Phoradendron supra venulosum Trel. Genus Phoradendron. 154. 1916. Branches inconspicuously angled, compressed and enlarged below the nodes; internodes rather long (4-7 cm.). Cataphylls a single pair at each joint, 1-2 cm. above the base, parted, 2 mm. high. Leaves oblong, moderately falcate, very acuminately pointed, cuneate to the base, moderately coriaceous, obscurely 5- nerved, to 13 cm. long, 6 cm. broad; petioles about 1 cm. long. Spikes solitary, 3-5 -articulate, 4-7 cm. long; peduncle 5-7 mm. long, enclosed within a large, funnel-shaped tube formed by the basal scales; joints rather stout, to 2 cm. long, 30-50-flowered, the flowers in 6 series. Fruit apparently elongated, with connivent Previously known from Guatemala, Nicaragua and Costa Rica. darien: Crist, Cana-Cuasi Trail, Terry 1571. This specimen, insufficient for a complete study, differs from Ph. supra- venulosum, as described by Trelease, in some small details and approaches Ph. membranaceum in certain others. Future studies may perhaps determine segrega- tion as a proper species. flora of Panama (Loranthaceaej 283 MISSOURI BOTANICAL GARDEN Trel. in Ann. Missouri Bot. Gard. 27:308. 1940. Branches slender, terete, regularly forked. Cataphylls a basal pair followed by 2 or 3 others, bifid at all internodes. Leaves ovate-oblong, scarcely cuneate to the rounded base, more or less abruptly acuminate, moderately leathery, obscurely 3- 5 -nerved, thickly cartilagineous-margined, 5-7 cm. long, 2-3 cm. wide; petioles nearly lacking. Spikes solitary, 5-6-articulate, 3-5 cm. long; peduncle 3-5 mm. long; joints lightly clavate, 5-8 mm. long, with about 10 flowers in 4 ranks. Fruits subglobose, 4 mm. long, the tepals inflexed. Endemic. cocle : between Las Margaritas and El Valle, Woodson, Allen 8 Seibert 1 302 (type) . 5. Phoradendron piperoides (H. B. K.) Nutt. in Journ. Acad. Nat. Sc. Phila- delphia, ser. 2, 1:185. 1847. Viscum latifolium Sw. Fl. Ind. Occ. 1:268. 1797, non Lam. Piperoides H.B.K. Nov. Gen. & Sp. 3 :443. 1820. Phoradendron latifolium (Sw.) Griseb., Fl. Brit. West Ind. 1*1. 314. 1860. A very variable shrub, abundantly branched. Shoots slender, slightly com- pressed above, soon becoming terete. Cataphylls a single pair on every joint, 2-5 pairs on the lowest one, pointed. Leaves, at least most of them, obliquely broadly ovate, angustate both toward the base and apex, acuminate, coriaceous, marginally crisped and brown to black in drying, nerveless or obscurely pinnately veined beneath, 5-12 cm. long, 2.5-4.5 cm. broad; petioles about 5 mm. long. Spikes clustered, slender, mostly 6-jointed, 4 cm. long; peduncle nearly suppressed; joints terete, 5 mm. long, 6-20-flowered, the flowers 4-ranked. Fruits subglobose, yellow, 4 mm. in diameter, with connivent tepals. Widely distributed through the New World. bocas del toro: vicinity of Nievecita, Woodson, Allen d Seibert 1843; Rio Crica- mola, between Finca St. Louis and Konkintoe, Woodson, Allen 8 Seibert IQOO, 1904; Almirante, Cooper 93, 193. canal zone: Chagres, Fendler 1 36; Barro Colorado Island, Shattuck 1 1 38; Fuertes House, Woodworth & Vestal 663; Mindi, Sutton Hayes 616. chiriqui: El Boquete, Pittier 334$; around Caldera, Pittier 3357; Bajo Mona, Davidson 522. cocle: above Penonome, Williams 258. darien: Chepigana, Cana-Cuasi Trail, Davidson & Terry 1572. Panama: Sabanas Road, Gillespie P-40. 6. Phoradendron crispum Trel. Genus Phoradendron 77. 1916. A moderately branched undershrub 30-60 cm. tall, the branches terete. Cata- phylls a single basal pair, short. Leaves obovate to round-obovate, a little attenuate to the base, firmly membranaceous, veinless or very obscurely 3 -nerved, drying crisped, 3.5-5 cm. long, 1.5-3 cm. wide; petioles 4-6 mm. long. Spikes solitary, with 2-3 short articles, 5-15 mm. long; peduncle almost suppressed; joints 10- 20 -flowered in 4 series. Fruits white, translucent. Panama and Costa Rica. CHiRiQui: Boquete, Volcan de Chiriqui, Davidson 992. (266) flora of panama (Lorantbaceae) 285 7. Phoradendron allenii Trel. in Ann. Missouri Bot. Gard. 27:307. 1940. Very similar to Ph. crispum Trel., but the flowers are constantly more numer- ous and 6-ranked. Branches terete. Cataphylls only a basal pair on the lowermost internode of the shoots. Leaves obovate, to 6 cm. long, 2.5 cm. broad, finely 3 -nerved; petioles about 5 mm. long. Spikes solitary, stout, 2-3 -articulate, to 2 cm. long; peduncle about 4 mm. long; joints 5-8 mm. long, 20-40-flowered irregularly arranged in 6 rows. Fruits globose, 4-5 mm. in diameter, the tepals more or less erect. Endemic. 8. Phoradendron seibertii (Trel.) Rizz., nov. stat. Phoradendron corynarthron Eichl. var. seibertii Trel. in Ann. Missouri Bot. Gard. 24:187. 1937. Branches 2 -edged, the branchlets more or less 4-angled. Cataphylls a single basal pair, tubular, the margin parted. Leaves narrowly oblong, some of them almost spatulate, rounded at the apex, attenuate to the base, nerveless, coriaceous, 4-7 cm. long, 1-1.5 cm. wide; petioles to 4 mm. long. Spikes solitary, slender, 4-6-articulate, to 2 cm. long; peduncle about 2 mm. long; joints short, terete, 10-1 8 -flowered, the flowers in 4 rows. Endemic. cocle : El Valle de Anton, Seibert 411 (type) . Ph. corynarthron Eichl. always shows clavate spike joints with 4 to 8 flowers, leaves, thus being clearly different from Ph. 9. Phoradendron corynarthron Eichl. in Mart. Fl. Bras. 5, pt. 2:115. 1868. Phoradendron davidsoniae Standi., Field Mus. Publ. Bot. 22:17. 1940. A handsome yellow (drying yellowish-brown) undershrub with flattened branches, the branchlets somewhat enlarged below the nodes. Cataphylls usually two pairs, one about 0.5-1 cm. above the node and the other 1.5-2 cm. above the first, pointed. Leaves lanceolate to narrowly oblong, rarely falcate, attenuate to both ends, subacute at the apex, cuneate to the base, moderately coriaceous, more or less revolute along the margins in drying, nearly nerveless to finely 3 -nerved, 4-7 cm. long, 0.8-1.5 cm. wide; petioles about 5 mm. long. Spikes solitary, typically 3-articulate, 1-1.5 cm. long (becoming 2-3.5 cm. when fruiting); peduncle 1-5 mm. long; joints clavately 4-8-flowered in 4 series, 2-10 mm. long. Fruit round, white, 3 mm. in diameter, the tepals closely connivent. Central America, widely dispersed in Panama. CHiRiQui: vicinity of Finca Lerida, Woodson 2? Schery 220; Bajo Mona, mouth of Quebrada Chiquero along Rio Caldera, Woodson 6 Schery 294; Loma Sardina, Allen 4769; vicinity of Casita Alta, Volcan de Chiriqui, Woodson, Allen (3 Seibert 929; Boquete, Davidson 740 (type of Phoradendron davidsoniae Standi.) and 923. cocle: Aguadulce, in savannas, Pittier 4955. (267) THE MISSOURI BOTANICAL GARDEN not be confused with Ph. rigidum Urban, from Vene- spikes bearing 2-3 joints, which are 10-flowered, and Trel. in Ann. Missouri Bot. Gard. 27:307. 1940. A dioecious, very bushy shrub up to 4.5 m. tall. Branches rather stout, 2 -edged, the branchlets highly compressed. Cataphylls a single pair, strictly basal, large, up to 3 mm. long. Leaves obliquely lanceolate, very acute, cuneately long- attenuate to the base, thus almost sessile or subpetioled for about 1 cm., finely 3-5 -nerved, somewhat coriaceous, to 15 cm. long, 1.5 cm. broad. Spikes both solitary and clustered, 3-4-articulate, turbinate, the flowers 6-ranked; pistillate ones much stouter, to 4 cm. long, the joints 1 cm. long, 16-24-flowered; staminate spikes slender, to 6 cm. long, the joints 6-12 mm. long, 3 0-70 -flowered; peduncle in any case 2-3 mm. long. Fruits globose, the tepals closely inflexed. Endemic. chiriqui: vicinity of Finca Lerida, Allen 4752; vicinity of New Switzerland, Allen I 399 (type) ; Cerro Vaca, Pittier 5376. Ph. novae-helvetiae, a pretty species, is very similar to Ph. tonduzii Trel, re- ported to occur in Panama (see below) ; it may be distinguished by the 6-seriate flowers and 30-70-flowered male joints (instead of 4-ranked as Trelease says). 11. Phoradendron tonduzii Trel. Genus Phoradendron 67. 1916. Authoritatively (see Standley's Flora of Costa Rica, p. 406) but not unques- tionably Panamanian; however, it may be expected. In habit of foliage and inflorescence it is closely related to Ph. novae-helvetiae Trel., above described. They are readily distinguishable by the flower disposition and by the number of male flowers. 12. Phoradendron gracilispicum Trel. Genus Phoradendron 130. 1916. Trelease mentions a collection from Chiriqui. The following description is taken from the original {ibidem) : Branches ancipital and rather persistently 2 -keeled, the internodes 4-6 cm. long. Cataphylls only basal or followed by a second pair some 2 cm. higher. Leaves lanceolate or oblong-acute, the base cuneate, devoid of nerves, 10-16 cm. long, 2.5-4 cm. broad; petioles about 1 cm. long. Spikes often clustered, markedly slender, very long (to 7 cm.), reddish, showing about 10 thin joints some 12- flowered in 4 (6) series; peduncle 2 mm. long, often followed by 1 or 2 partly or wholly sterile joints, the scales ciliolate. Young fruit elongated, the tepals Viscum undulatum Pohl, in DC. Prodr. 4:282. 183 (268) flora of Panama (Loranthaceae) 287 (269) 288 ANNALS OF THE MISSOURI BOTANICAL GARDEN Yellow, with a brownish tint, moderately tall shrub hanging from trees. Branches ancipital and 2-keeled, especially toward the tips. Cataphylls a nearly basal pair followed by 2-3 additional ones, rather pointed. Leaves lanceolate, not rarely somewhat ovate, acuminate, under lens rugose, the nerves wanting, the margins callose- thickened and conspicuously undulate, 7-12 cm. long, 2-4 cm. wide; petioles 3-8 mm. in length, canaliculate above. Spikes many times clustered, 2-5 cm. long, with 5-9 moderate joints; peduncle none or up to 8 mm. long; joints mostly 10-flowered in 2 ranks, 2-5 mm. long. Fruits round-ovoid, smooth, 3 mm. in diameter, the tepals slightly parted. Another interesting finding of a plant, previously known only from Brazil, in Panama. The excellent specimen, mentioned below, minutely examined and compared with several Brazilian ones, agrees even in the smallest details as, for instance, in the callose-margined and finely verrucose leaves. Brazil, Panama. chiriqui: pastures around El Boquete, Pittier 2932. Ph. gracilis picum Trel. is to be distinguished from the above by it longer, slender and many-jointed spikes. 14. Phoradendron cooperi Trel. Genus Phoradendron 67. 1916. Branches rather stout, 2-sided or ancipital. Cataphylls a single, strictly basal pair, pointed. Leaves falcately lanceolate to oblong-lanceolate, long-attenuate to the somewhat obtuse apex, thin, finely 5-nerved, cuneately subpetioled for 1-1.5 cm., to 20 cm. long, 2.5 cm. broad. Spikes solitary, 3-4-articulate, about 3-4 cm. long; peduncle nearly supressed; joints 16-2 5 -flowered in 4 ranks, 8-10 mm. long. Fruits round, 3 mm. in diameter, the tepals closely meeting. Central America. chiriqui: Boquete, Davidson 580. 15. Phoradendron mucronatum (DC.) Kr. et Urb. in Engl. Bot. Jahrb. 24: Phoradendron minor (Eichl.) Trel., The Genus Phoradendron 117. 1916. Branches somewhat nodose, the branchlets sharply 4-angled. Cataphylls a single basal pair, 2 mm. long. Leaves obovate, often emarginate, cuneate to the base, coriaceous, distinctly 3-5-nerved, to 3.5 cm. long, 2 cm. broad; petioles about 2-4 mm. long. Spikes solitary, subsessile, 3-4-jointed, 5 mm. long, when fruiting to 1.5 cm. long; joints 4-6-flowered in 4 series. Fruits ovoid, tuberculate, 3-4 mm. long, the tepals nearly erect. South and Central America, West Indies. (270) FLORA OF PANAMA (Loranthaceae) 289 zed As shown by Urban, Ph. emarginatum (Brazil) only differs from Ph. mucro- natum (West Indies) by the larger leaves and longer spikes. The var. minor of the former species, with smaller leaves and shorter spikes, is intermediate between the two. Actually, there are no clear-cut differences. Southern Brazilian specimens are con- stantly stouter than Central American and West Indian ones; but the northeast- ern Brazilian material is smaller than the former and approaches the latter. In short: Ph. mucronatum is a South and Central American and West Indian rather variable entity, well ch; and easily recognizable by the obovate emarginate leaves, 4-angled branches, 4- 6-flowered spike joints and verrucos< fruit — as it is evident from the examina Fig. 81. Phoradendron mucronatum 16. Phoradendron trinervium Griseb. Fl. Brit. West Ind. Isl. 314. 1860. Phoradendron rubrum var. brevispica Eichl. and var. latifolia Eichl. in Mart. Fl. Bras. 5, pt. 2:121. 1868. Branches toward the tips rhombically quadrangulate, the branchlets somewhat compressed and swollen below the nodes. Cataphylls a single basal pair, short. Leaves narrowly obovate or elliptical-obovate, angustate to the base, very obtuse, almost nerveless despite the specific name, coriaceous, 2-5 cm. long, 1-1.5 cm. wide; petioles 2-4 mm. long. Spikes solitary or clustered, 3-4-articulate, up to 3 cm. long; peduncle 1-2 mm. long; joints slender, 10-1 8 -flowered in 4 series, 4-7 mm. long. Fruit yellow, ovate, 3-5 mm. long, the tepals erect, parted. West Indies, Panama. bocas del toro: vicinity of Chiriqui Lagoon, von Wedel 2763. 17. Phoradendron venezuelense Trel. Genus Philodendron 111. 1916. Phoradendron herrerense Trel. in Ann. Missouri Bot. Gard. 27:307. 1940. Phoradendron sonanum Trel. loc. cit. 308. 1940. A rather variable, androgynous shrub. Branches 4-lined, the branchlets quad- rangulate, slightly compressed and somewhat enlarged below the nodes. Cataphylls a single basal pair, parted. Leaves lance-oblong, more or less falcate, obtuse or nuate to the base, leathery, indistinctly (271) ANNALS OF THE MISSOURI BOTANICAL GARDEN 3-5 -nerved or nearly nerveless, cartilaginous-margined, 3-6 cm. long, rarely up to 8 cm., 7-15 mm. broad; petioles 2-5 mm. long. Spikes mostly clustered, slender, often 3 -articulate (seldom to 5 -jointed), about 1.5 cm. long, but attaining to 5 cm. when fully developed; peduncle 3-10 mm. long; joints 10- flowers 4-seriate, to 1 cm. long. Fruit globose, yellow, 3 mm. Widely distributed both in Venezuela and Panama. Also in Colombia and the West Indies, where it is rare. bocas del toro: Water Valley, von Wedel 936; Chiriqui Lagoon, von Wedel I57 8 > 2518, 2599, 2648. canal zone: Anc Balboa, Seibert 408; around Gamboa, Pittier 2604; Barro Colorado Island, near Barbour Point, Wilson 146. herrera: Pese, Allen 798 (type of Ph. herrerense Trel.). Panama: Sabanas near Chepo, Hunter 2? Allen 7 8; on Sabanas, road to Chepo, Hv ».; Rio Tecumen, Standley 26552; along Corozal Road, Standley 26869; Llanos de Panama Viejo, Heriberto 288. veraguas: hills of Sona, Allen 1033 (type of Ph. sonanum Trel.). province unknown: Sutton Hayes 829. Ph. herrerense Trel. only differs from Ph. venezuelense in the more regularly branched stem; Ph. sonanum Trel. is distinguished from the latter by some longer, 3-5- jointed spikes; in both cases the differences are too poorly developed to be (272) FLORA OF PANAMA (OpUtOC OPILIACEAE 1. AGONANDRA Miers Dioecious trees, the bark generally corky. Leaves alternate, simple, thin, usually small, ovate to elliptic, petiolate. Inflorescences axillary, racemose, bracte- ate. Flowers small, green, pedicellate. Calyx small, usually crateriform, (5- or) 4-lobed or undulate. Petals 4 (or 5 ) , free, valvate, especially caducous in pistillate flowers. Stamens 4 (or 5), antipetalous, the filaments filiform, the anthers ovoid; absent in pistillate flowers. Disc annular or cupuliform, irregularly lobed, more or less connate at the base, generally erect. Pistil 1, superior, the ovary 1-loculate, the style generally obsolete, the stigma 4-lobed. Fruit a berry. In the past pistillate flowers of this genus have been characterized by the absence of petals. Examination of the single collection from Panama clearly shows the presence of petals and also reveals their caducous nature. Approximately a dozen species distributed from Mexico to Argentina. Known from a single species in Panama. 292 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. Agonandra brasiliensis Benth. & Hook. f. Gen. 1:349. 1862. Trees to 8 m. tall, the bark rough, brown, the branchlets slender, terete, gla- brous, olive-green. Leaves ovate, 2.5-5.5 cm. long, 2-3 cm. broad, abruptly and briefly acuminate at the apex, obtuse and atenuate at the base, glabrous, the costa plane to immersed above, emersed below, the primary lateral veins prominulous; petiole 3-9 mm. long. Staminate inflorescence 30- to 75-flowered, farinose, the rhachis 3-7 cm. long. Staminate flowers (immature) : pedicel about 0.5 mm. long; calyx crateriform, about 0.5 mm. long, farinose, undulate; petals 5, ovate, about 1.5 mm. long, 0.75 mm. broad, farinose without, glabrous within; stamens 5, the filaments 1-2 mm. long, glabrous, the anthers about 0.5 mm. long and broad; disc 5-lobed, the lobes very nearly free, about 0.5 mm. long, fleshy, truncate, obscurely and irregularly lobed at the apex, glabrous; pistillode conical, about 0.75 mm. long, glabrous. Pistillate inflorescences 10- to 15-flowered, farinose, the rhachis 0.5-4.5 cm. long. Pistillate flowers: pedicel 0.5-0.75 mm. long; calyx crateriform, about 0.25 mm. long, farinose without, the margin undulate; petals 4, ovate-trigonal, about 1 mm. long, 0.5 mm. broad at the base, farinose without, indefinitely papillate to glabrous within; disc annular, about 0.25 mm. tall, irregularly lobed to the middle, glabrous; ovary doleiform, 0.5-0.75 mm. long, glabrous, the style obsolete, the stigma 4-lobed. Berry ovoid, 12-15 mm. long, 10- 12 mm. in diameter, glabrous; pedicel 5-8 mm. long. Central and South America. The single collection examined was flowering and fruiting in February-March and was collected at an altitude of 50-1000 ft. cocle: Penonome and vicinity, R. S. Williams 368. flora of Panama (Olacaceae) OLACACEAE By LORIN I. NEVLING, Jr. ;s, shrubs or perhaps vines, autotrophic or reportedly imes opposite, simple, generally entire, pin- nately veined, petiolate; estipulate. Inflorescences axillary, panicles, racemes, cymes or umbels, often fasciculate. Flowers dichlamydeous, bisexual or rarely polygamodioecious, sometimes polymorphic, actinomorphic, generally small and inconspicuous. Calyx small, gamosepalous, crateriform to campanulate, usually with a 3- to 6-toothed margin, free or basally adnate to the ovary or to the disc, sometimes accrescent. Petals 3-6, usually the same number as calyx lobes, free or connate, valvate or imbricate in bud. Disc sometimes present, simple or cupuli- form. Stamens generally the same number as, or twice the number of, petals, the anthers dehiscing longitudinally or by valves. Pistil 1, the ovary superior or rarely half -inferior, sessile, generally 2- to 5-loculate, the ovules the same number as locules, pendulous, integuments 0, 1 or 2, the style simple, sometimes hetero- morphic, terminal, elongate to obsolete, the stigma often 3 -parted. Fruit mostly a drupe, sometimes accompanied by a colorful accrescent calyx. A pantropic family of about 25 genera and 300 species. Four genera are known from Panama. A fifth genus, Chaunochiton, is known from Costa Rica, Guiana and northern Brazil. Whether this disjunction is real or is an artifact of collection is not known at this time. a. Calyx crateriform or cupuliform, free from the ovary, with 3 to 6 lobes or teeth; ovary superior; ovules with 1 or 2 integuments; fruit a drupe; plants root parasites (Ximenia) or autotrophes. 1. MINQUARTIA Aubl. Minquartia Aubl. Hist. PI. Guyan. Franc. Suppl. 4. t. 370. 1775. Secretania Mull.-Arg. in DC. Prod. 15 2 :227. 1866. Minguartia Miers, in Journ. Linn. Soc. 17:33 8. 1879. Unarmed autotrophic trees, the young branches generally densely rusty- tomentulose, the hairs branched, the older stems often perforated. Leaves alternate, (275) 294 ANNALS OF THE MISSOURI BOTANICAL GARDEN elliptic, acuminate at the apex, glabrous above, glabrescent below, with resin canals and laticiferous ducts. Inflorescences spicate, bracteate. Flowers bisexual, hypogy- nous, small. Calyx cupuliform to crateriform, small, (6-) 5 -toothed, persistent but not accrescent. Petals 5 (or 6) , connate into a campanulate corolla tube, puberulent to villous within, the lobes valvate in bud. Stamens 10 (or 12) , in two whorls, the anthers dehiscing longitudinally, the filaments filiform, adnate to the corolla tube, antipetalous generally longer. Ovary 5-merous but often reduced to 3 or 4 locules by abortion, the stigma 3- to 5-lobed. Fruit a small ovoid or Three or four species distributed in Panama, Ecuador, Guiana and Brazil. A single species is known from Panama. 1. Minquartia guianensis Aubl. Hist. PL Guyan. Franc. Suppl. 4. t. 370. 1775. Secretania loranthacea Mull.-Arg. in DC. Prod. 15 2 :227. 1866. Trees to 30 m. tall, the young branches angular, glabrous. Leaves elliptic, 8-16 cm. long, 3.0-7.5 cm. broad, abruptly and briefly acuminate at the apex, rounded or obtuse at the base, chartaceous, glabrous, the costa emersed above and below, the primary lateral nerves numerous, conspicuous; petiole 5-23 mm. long, broadly canaliculate. Inflorescence spicate, rusty-tomentulose, the rhachis elongat- ing to 5 cm. long, the flowers sessile, subtended by small ovate bracts. Calyx cupuliform, about 1 mm. long, 1.5-2.0 mm. in diameter, 5-toothed, rusty- tomentulose without; petals 5, connate into a campanulate corolla tube, the tube 1.0-1.5 mm. long, tomentulose without, black-punctate within, the lobes 1.0-1.5 mm. long, 1 mm. broad, villous within; stamens 10, inserted just below the orifice, the alternipetalous inserted slightly lower than the antipetalous, the anthers broader than long, minute, the filaments about 0.5 mm. long, glabrous; ovary globose, about 1.5 mm. in diameter, densely rusty-tomentulose, the style short. Immature drupe ellipsoid, about 7 mm. long, 4 mm. in diameter. The Panamanian specimens are sterile, for all practical purposes, and the in- complete floral description is based upon fragmentary collections from South Known as manwood and nispero negro in Panama. The wood is heavy and extremely durable. bocas del toro: region of Almirante, Cooper 497. canal zone: Gamboa, Standley 28380. CHmiQuf: Progreso, Cooper 2? Slater 312. The cited specimens are, with the exception of some very immature inflores- cences, glabrous throughout. In this respect they are unlike their South American counterparts, which are usually rusty-tomentulose throughout. 2. HEISTERIA Jacq. Heisteria Jacq. Enum. Pi. Carib. 4. 1760. Rhaptostylum Humb. & Bonpl. Fl. Aequin. 2:139, /. 125. 1809. Hesiodia Veil. Fl. Flum. 4. /. 1 40. 1825, (as Hesioda). Acrolobus Klotzsch, in Verhandl. Akad. Wiss. Berlin 236, t. ?. 1856. FLORA OF Panama (Olacaceae) 295 Raptostylus Post & O. Kuntze, Lex. Gen. Phane Pbanerocalyx Spencer Le Moore, in Journ. Bot. Unarmed autotrophic shrubs or trees. Leaves alternate, oval, elliptic or oblanceolate, glabrous, with laticiferous ducts. Inflorescences in compact axillary fascicles, multi-flowered although only 1 or 2 per axil develop into mature fruit. Flowers bisexual, hypogynous, small. Calyx small, generally crateriform, free, 5-6 toothed or lobed, accrescent, becoming coriaceous. Petals 5-6, somewhat connate at the very base, glabrous or puberulent to villous within, valvate in bud. Stamens 10 or 12 in two whorls (in Panamanian species) or 5-6 in one whorl, the anthers globose, dehiscing longitudinally, the filament filiform or liguliform, antipetalous often somewhat shorter than the antisepalous. Ovary superior, conical, 3-merous to the middle, the ovules 3, pendulous, with 2 integuments, the style short to obsolete, the stigma 3 -lobed. Fruiting calyx more or less reflexed, often brilliantly colored; drupe globose, oblate spheroid or ellipsoid; embryo embedded in the top of the endosperm, small. About 65 species, West Africa, Central and South America. Five species are known from Panama. The genus is generally divided into two sections on the basis of the number of stamens. The five Panamanian species have either 10 or 12 stamens and are therefore placed in section Euheisteria Engl, (in Engl. & Prantl Aufl. 2. Nachtr. 19. 1900). Heisteria is marked by a paucity of flowering specimens which is only partially overcome by a particularly abundant supply of fruiting specimens. Unfortunately, the fruiting structures are rather variable and thus specific delimitation is some- times obscure. 296 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. Heisteria fatoensis Standi, in Field Mus. Nat. Hist. 8:137. 1930. Shrubs or trees to 3.5 m. tall, the young branches generally angular, i strikingly flexuose, glabrous, brownish-red. Leaves ovate, 5-1 3 cm. long, 3-6 cm. broad, abruptly and briefly acuminate at the apex, rotund at the base, chartaceous, the costa plane above, emersed below, the primary lateral veins 5-6 pairs; petiole 7-12 mm. long, slightly canaliculate. Calyx crateriform, 5 -toothed, glabrous, the teeth 0.5 mm. long and broad; petals 5, deltoid, 1.75-2.0 mm. long, about 1 mm. broad at the base, glabrous without, puberulent within; stamens 10, free, the anthers to 0.25 mm. long and broad, the filaments liguliform, 1.0-1.5 mm. long, about 0.25 mm. broad, puberulent; pistil conical, 0.75-1.0 mm. long, about 0.75 mm. in diameter, glabrous; pedicel 4.5—6.0 mm. long. Fruiting calyx much shorter than the drupe, obscurely lobed, strongly reflexed, about 1 cm. in diameter; drupe oblate spheroid, 7-9 mm. long, 9-11 mm. in diameter, cherry to yellowish- red; pedicel 13-20 mm. long. Known only from Panama. Flowering and fruiting from July to October. bocas del toro: Water Valley, Von Wedel 1401. colon: Loma de Gloria, near Fato (Nombre de Dios) , Pittier 4244. Pittier reports the type collection as being "a woody vine". He may have been mistaken as vines are little known in this genus. In addition, the specimens do not have a vine-like aspect. 2. Heisteria concinna Standi, in Field Mus. Nat. Hist. Bot. 8:137. 1930. Trees to 9 m. tall, the young branches terete, glabrous, olive-green to brownish- red. Leaves ovate, 6-1 3 cm. long, 3-6 cm. broad, abruptly and briefly acuminate at the apex, acute to obtuse at the base, coriaceous, the costa plane to immersed above, emersed below, the primary lateral veins generally prominulous; petiole 7-18 mm. long, canaliculate. Calyx crateriform, 5 -toothed to the middle, about 1 mm. long, glabrous, the teeth 0.75 mm. long, 0.5 mm. broad; petals 5, elliptic, 2.0-2.5 mm. long, 1.25 mm. broad, glabrous without, the lower half puberulent to villous within; stamens 10, free, the anthers 0.25 mm. long and broad, the filaments filiform, about 1.5 mm. long, puberulent; pistil Iageniform, the ovary oblate spheroid, 0.5 mm. long, 0.75 mm. in diameter, the style about 0.75 mm. long; pedicel about 5 mm. long. Fruiting calyx shorter than the drupe, 1.5-2.5 cm. in diameter, reflexed, distinctly 5 -lobed, red; drupe broadly ellipsoid, 10-15 mm. long, 8-10 mm. in diameter; pedicel 8-10 mm. long. Costa Rica and Panama. Flowering and fruiting from December to February at altitudes to 120 meters. Heisteria costaricensis Donn. Sm. in Bot. Gaz. 19:254. 1894. Shrubs to 3 m. tall, the young branches angular, glabrous, olive-green. Leaves ear-lanceolate, 16-25 cm. long, 1.5-5.0 cm. broad, gradually tapering to the :x, acute to obtuse at the base, chartaceous, the costa immersed above, emersed (278) flora of Panama (Olacaceae) 297 below, the primary lateral veins numerous; petiole 3-13 mm. long, scarcely canaliculate. Calyx crateriform, deeply 5 -toothed, glabrous, the teeth about 0.5 mm. long, 0.5 mm. broad; petals 5, deltoid, about 2 mm. long, 1 mm. broad, glabrous; stamens 10, free, the anthers to 0.5 mm. long and broad, the filaments liguliform, about 1 mm. long, 0.25 mm. broad, glabrous; pistil conical, about 1 mm. long, 1 mm. in diameter; pedicel about 1.5 mm. long. Fruiting calyx longer than the drupe, 1.5-2.5 cm. in diameter, shallowly 5-lobed, somewhat enclosing the drupe or reflexed, red; drupe ovoid, 6-8 mm. long, about 7 mm. in diameter, vertically ribbed, blue; pedicel 6-9 mm. long. Costa Rica and Panama. In Panama fruiting from October to February. canal zone: Quebrada Lopez, Allen 2128; along Caiio Quebrado, Pittier 6825; Barro Colorado Island, Hunnewell 16422, Standley 40898, 41168. 4. Heisteria macrophyixa Oerst. in Vid. Medd. Kjoebn. 1856:40. 1857. Heisteria latifolia Standi, in Journ. Wash. Acad. Sci. 17:8. 1927. Shrubs to 2 m. tall, the young stems terete to somewhat angular, glabrous, olive-green. Leaves elliptic or broadly elliptic to oblanceolate, (5-) 10-37 cm. long, 3-8 cm. broad, acuminate at the apex, acumen to 8 cm. long, cuneate to obtuse at the base, chartaceous to subcoriaceous, the costa emersed above and below, the primary lateral veins numerous, arcuate-ascending; petiole 6-30 mm. long, canaliculate. Calyx crateriform, 5- or 6-toothed, glabrous; petals (4 or) 5, deltoid, about 1.75-2.0 mm. long, 1-2 mm. broad at the base, glabrous; stamens 10, the anthers 0.5 mm. long and broad, the filaments liguliform, 0.75-1.5 mm. long, 0.5 mm. broad, glabrous; pistil conical, 1 mm. long and broad, glabrous; pedicel 1.25-2.0 mm. long. Fruiting calyx as long as or longer than the drupe, 2-3 cm. in diameter, inconspicuously and shallowly 5-lobed, more or less reflexed, red; drupe ovoid or ellipsoid, 8-12 mm. long, 6-9 mm. in diameter, vertically ribbed, black; pedicel 8-12 mm. long. A common forest species of Central America. Known in Panama as ajicillo and fruiting from April to November. bocas del toro: vicinity of Chiriqui Lagoon, Von Wedel 1082, 12/8; Fish Creek Mts., Von Wedel 2269; Water Valley, Von Wedel 846, 913, 1538, 1 720. canal zone: between France Field and Catival, Standley 30366; Barro Colorado Island, Standley 40877. colon: along Rio Culebra, above Santa Isabel, Pittier 4156. Heisteria latifolia was proposed by Standley as a new species primarily on the basis of the very broad leaves with conspicuously elongate petioles. A continuous series between the leaves of H. macrophylla and H. latifolia can be demonstrated in Panamanian material and therefore it does not seem reasonable to maintain H. latifolia as a distinct species. The flowers appear to be rather variable, even in number of parts. A flower from Von Wedel 1538 (MO) was found to have 6 calyx lobes, 4 petals and 10 Standley reported this species as attaining 10 meters in height (Field Mus. Bot. Ser. 18 1 " 2 :409. 1937). OF THE MISSOURI BOTANICAL GARDEN . Wash. ad. Sci. 17:8. 1927. Shrubs or trees to 20 m. tall, the young branches somewhat angular, glabrous, olive-green. Leaves ovate to elliptic or rarely oblanceolate or obovate, 5-15 cm. long, 3-8 cm. broad, acute to abruptly and briefly acuminate at the apex, acute at the base, chartaceous, the costa emersed above and below, the primary lateral veins numerous; petiole 5—15 mm. long, deeply canaliculate. Calyx crateriform, 5-lobed to one-third from the base, glabrous; petals 5, deltoid, 2.25-2.75 mm. long, 1.25- 1.5 mm. broad, glabrous; stamens 10, free, the anthers about 0.25 mm. long and broad, the filaments liguliform, about 1.5 mm. long, 0.5 mm. broad, glabrous or indefinitely papillate; pistil conical, 1 mm. tall, 1.25 mm. in diameter; pedicel 3.5—4.0 mm. long. Fruiting calyx about as long as the drupe, 1.0-2.0 (-2.5) cm. in diameter, shallowly 5-lobed or undulate, generally somewhat reflexed, red; drupe ellipsoid or obovoid, 5-10 (-12) mm. long, 5-7 mm. in diameter; pedicel 10-23 mm. long. Fruiting from January to September to 2000 meters. Known as naranjillo Colorado in Panama. vs del toro: Changuinola Valley, Dunlap 444, 567, Cooper (3 Slat* nte, Cooper 569; Old Bank Island, Von Wedcl 212]; h\: ' trt 1945. canal zone: along the Trinidad River, Piti rest of Limon Bay, Gatun Locks and Gatun Lake), Johi Colon, Woodson, Allen 006; Tortuguilla cove 1523; Barro Colorado (280) flora of Panama (Olacaceae) 299 Island, Bailey d Bailey 8l, Bangham 534, Kenoyer 341, Salvoza 923, 958, Wetmore 8 Abbe 20, 99, Wilson 153, Woodworth 8 Vestal 436. chiriqui: vicinity of Puerto Armuelles, Woodson 2f Scbery 856, Stern 2? Chambers 126; trail from Paso Ancho to Monte Lirio, upper valley of Rio Chiriqui Viejo, Allen 1589; Progreso, Cooper & Slater 166. cocle: El Valle de Anton, along Rio Indio trail, Hunter & Allen 328. darien: near the mouth of Rio Yape, Allen 339. Panama: Cerro Campana, trail from Campana to Chica, Allen 2657. san blas: high hills back of Puerto Obaldia, Pittier 4289. The type specimen, Pittier 4006 (US) , is an unusual specimen for two reasons: the petals and stamens which are ordinarily caducous have persisted into young fruit formation; secondly, Standley reported 6 petals and 12 stamens for it. Examination of the immature fruit of this specimen revealed only 5 petals and 10 stamens although there appeared to be some peculiarities in the androecial whorl, namely, basal cohesion between some of the filaments. Such floral variability upon a single specimen is entirely possible and has been demonstrated in at least one other species from Panama, H. macrophylla Oerst. 3. XIMENIA L. Ximenia L. Sp. PL 1193. 1753. Heymassoli Aubl. Hist. Pi. Guyan. Franc. 1:324, /. 125. 1775. Rottboelia Scop. Introd. 233, n. 1060. 1777. Pimecaria Raf. Alsog. Am. 64. 183 8. Armed root parasitic shrubs or trees, the spines axillary, the young branches glabrous. Leaves alternate, ovate or elliptic, mucronate or acuminate at the apex, obtuse to cuneate at the base. Inflorescence umbellate, simple or compound, sometimes fasciculate, bracteate. Flowers bisexual, hypogynous, medium-sized. Calyx small, 3- to 5 -toothed, not accrescent. Petals 4 or 5, free, thickly covered with red-brown barbed hairs within, valvate in bud. Stamens 8-10, in two whorls, the anthers oblong or linear, dehiscing longitudinally, the filaments filiform. Ovary long-conical, 4-loculate above the middle, the ovules pendulous, with 1 integument, the style filiform, about as long as the ovary, the stigma minutely capitate. Drupe ovoid to ellipsoid, from yellow to purple; seed with a small embryo at the apex of the fleshy endosperm. Approximately 15 species, circumtropical; Florida, Central and South America, Antilles, Africa, East Indies, Asia, and Australia. Only one species known from 1. Ximenia Americana L. Sp. Pi. 1193. 1753. Ximenia multiflora Jacq. Enum. Pi. Carib. 19. 1762. Ximenia inermis L. Sp. Pi. ed. 2. 497. 1762. Ximenia aculeata Crantz, Inst. 2:381. 1766. Heymassoli spinosa Aubl. PI. Guayan. Fran?. 1:324, t. Ximenia elliptica Forst. f. Prod. 27. 1786. 300 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ximenia loranthifolia Span, in Linnaea 15:177. 1841. Ximenia arborescens Tussac, ex Walp. Rep. 1:377. 1842. Ximenia laurina Delile, in Ann. Sci. Nat. ser. 2. 20:89. 1843. Ximenia fluminensis M. Roem. Syn. Hesper. 22. 1846. Ximenia oblonga Lam. ex Hemsl. in Biol. Centr. Am. Bot. 1:185. 1879. apex, obtuse to cuneate at the base, subcoriaceous, glabrous or bullate, the costa immersed above, emersed below, the primary lateral veins 3-6 pairs, most conspicu- ous towards the base of the lamina; petiole 5-7 mm. long, canaliculate. Inflores- cence umbellate, simple or compound, often fasciculate, multi-flowered, the bracts small, ciliate, caducous. Calyx crateriform, to 0.75 mm. long, about 1.5 mm. in diameter, 3- or 4-toothed, ciliate; petals 4, liguliform, 7-11 mm. long, 1.5-2.0 mm. broad, greatly reflexed, coriaceous, glabrous without, densely covered with red-brown barbed hairs within, white and often purple tipped; stamens 8, sub- exserted, the anthers linear, 3.0-4.5 mm. long, 0.5-0.75 mm. broad, the filaments 3.0-4.5 mm. long, glabrous; ovary long-conical, 3-4 mm. long, 1.0—1.5 mm. in diameter, glabrous, the style 3.0-4.5 mm. long; pedicel 5-6 mm. long. Drupe ellipsoid, 2-3 cm. long, 1.5-2.5 cm. in diameter, pale yellow. A circumtropical species. flora of Panama (Olacaceae) 301 bocas del toro: vicinity of Chiriqui Lagoon, Von Wedel 1402; Water Valley, Von Wedel 1620. canal zone: vicinity of Fort Sherman, Standley 31145; Victoria Fill, Allen 1750. chiriqui: Isla Parida, Pit tier 28 1 9. darien: Patirio, on cliffs along beach, Pit tier $70S\ l icinity of La Palma, Pittier 3498. Panama: San Jose Island, Erlanson 1 49, John- ston 719, 774\ Bella Vista, Killip 39943; Nuevo San Francisco, Standley 30780. province unknown: Bahia Honda, Elmore H39; La Venta, Hunnewell 16417. Schoepfia Schreb. Geo . SCHOEPFIA Schn 1789. vt. Natur. Selsk. Kjoeb. 2 Codonium Rohr, ex Vahl, in Skrivt. Haenkea Ruiz & Pavon, Fl. Per. 3:8, /. 231. 1802. Diplocalyx A. Rich, in Sagra, Hist. Cuba 11:81. 1850. Ribeirea F. Allemao, Trab. Comm. Sc. Expl. Bot. Rio de Janeiro 29, 3 8. 1864. Schoepfiopsis Miers, in Journ. Linn. Soc. 17:75. 1878. Unarmed glabrous shrubs or trees, reportedly root parasites. Leaves alternate, entire, coriaceous. Inflorescences few-flowered racemes generally fasciculate in the leaf axils, with or without basal perular bracts and the flowers subtended by a cupule composed of 2 bracteoles and 1 bract or subtended by a single bract. 302 ANNALS OF THE MISSOURI BOTANICAL GARDEN Flowers bisexual, small. Calyx extremely small, cyathiform or crateriform, margin subentire. Petals 3-6, united into a tubular-campanulate corolla, often pubescent within, the lobes valvate, reflexed, white, yellow or red. Stamens 3-6, in one whorl, as many as the petals, antipetalous, adnate to the corolla, the anthers oval, dehiscing longitudinally, sessile or short-filamented. Ovary half-immersed, the ovules pen- dulous, lacking integument, the style thin, the stigma 2- or 3-lobed. Spurious fruit with the drupe enclosed in the adnate, accrescent calyx, eccentrically annulate at the apex; seed with the small embryo at the tip of the endosperm. About 3 8 species of the Old and New World. Only one species is definitely known from Panama. A second species, S. vacciniiflora, has been reported by Standley (in Field Mus. Nat. Hist. Bot. 18 2 :409. 1937 and in Fieldiana Bot. 24 4 :90. 1946) in Panama. The S. vacciniiflora name is in itself a confusing matter. Schoepfia vacciniiflora Planch, ex Hemsl. (Diag. PL Mex. 5. 1878) is reported from Guatemala whereas S. vacciniiflora Planch, ex Planch. & Triana (Ann. Sci. Nat. ser. 5. 15:382. 1872) is reported from Colombia. Whether or not these epithets are based upon a common species is difficult to determine. . F. Gmel. Syst. Veg. 2:376. 1791. Schoepfia americana Willd. Sp. PI. 1:996. 1798. Schoepfia arborescens Roem. & Schult. Syst. 5:160. 1819. Shrubs or trees to 8 m. tall, the young branches slender, angular, olive- green to white. Leaves ovate to lanceolate, 4-8 cm. long, 1.5-3.5 cm. broad, acute- acuminate to acuminate at the apex, cuneate-attenuate or obtuse at the base, the costa emersed above and below, the primary lateral veins 5-6 pairs, rather con- spicuous; petiole to 5 mm. long. Inflorescences 1- to few-flowered, fasciculate, the peduncle to 5 mm. long, the flowers subtended by a cupule composed of 2 bracteoles and 1 bract, the cupule 1-3 mm. in diameter, 3 -toothed, ciliate. Calyx cyathiform, about 1 mm. long, subentire, glabrous; corolla tube 2.0-2.5 mm. long, 1.5-2.5 mm. broad at the orifice, glabrous, red, the lobes 4 or 5, deltoid, 1.0-1.5 mm. long and broad, glabrous without and minutely puberulent within except for a cluster of villous hairs at the point of staminal insertion; stamens 4 or 5, inserted at the orifice of the corolla tube, the anthers 0.5-0.75 mm. long and broad, exserted, subsessile; ovary half -inferior, globose, densely papillate, the style 0.5 mm. long, the stigma bifid, included. Spurious fruit ellipsoid, 10-13 mm. long, 7-8 mm. in diameter, glabrous. Florida, Central America, Antilles and South America. flora of Panama (Balanophoraceae) BALANOPHORACEAE Fleshy root parasites, rhizomatous, lacking chlorophyll, the rhizome tuberous, digitiform, cylindrical or filiform, sometimes branched, naked or sometimes with scale-leaves, yellow to dark red in color. Inflorescence monoecious or dioecious, highly modified, of endogenous origin, above ground at anthesis, more or less erect, sessile or pedunculate (in our species), the peduncle terete, with or without an encircling, often basal sheath, scaly or naked above the sheath, the head with or without protective peltate scales, sometimes with paraphysoid trichomes or reduced pistillate flowers surrounding or subtending the flowers. Flowers unisexual, mono- chlamydeous, small. Perianth absent or more often present, of 2-4 (-8) tepals, free or connate into a tube of varying size and shape, in pistillate flowers sometimes reduced to an epigynous collar or absent. Stamens generally as many as perianth segments and opposite them, the filaments free or connate into a staminal column of varying length, sometimes very short, the anthers free or connate into a syandrium, introrse or extrorse; absent in pistillate flowers. Pistil composed of 1, 2 (or 3 ) carpels, the ovules usually 1 per carpel, lacking integument, often fused to the wall of the ovary, the styles 1 or 2, terminal, the stigma usually simple, rarely sessile; in stamina te flowers absent or reduced to a pistillode. Fruit nut-like or drupaceous, the endocarp usually hard; seed sometimes fused to the wall of the fruit, the embryo apical in the fleshy oil-rich endosperm, with or without suspensor. Approximately 18 genera, pantropic but primarily distributed in the Southern Hemisphere. Three genera are represented in Panama. 304 ANNALS OF THE MISSOURI BOTANICAL GARDEN Helosis Rich, in J Caldasia Mutis, ex Caldas, in Seeman. Nuev. Gran. 2:26. 1810, nom. rejic. Latraeophila Leandr. Sacram. ex St. Hil. in Ann. Sci. Nat. ser. 2. 7:32. 1837. Lathraeophila Hook. f. in Trans. Linn. Soc. 22:5 5. 1856. Rhizome filiform to digitiform, often intertwined, rich in starch, yellowish to brownish. Inflorescence monoecious, claviform, the peduncle thick, generally encircled at the base or at the middle with a glabrous, toothed to lobed, oblique sheath (rarely absent) , the head globose, ovoid or ellipsoid, when immature covered with 6-sided, peltate, umbonate scales beneath which occurs a dense mat of para- physoid trichomes and young developing flowers, the scales spirally arranged, deciduous, about the axis of each scale are oriented two whorls containing many pistillate flowers surrounded by a single whorl of 6 flowers which are situated at the angles of the covering scale and are either rudimentary or functionally stami- nate. Perianth in the staminate flowers composed of 3 (or 6) tepals connate in the lower half into a tube, the lobes becoming widely reflexed; in the pistillate flowers reduced to a low bilabiate epigynous collar or, according to some authors, absent. Stamens as many as the perianth lobes and opposite them, exserted, the filaments adnate to the perianth tube below, free or connate into a staminal column, generally free above, the anthers united into a synandrium, each with two posterior and a single anterior pollen sac, introrse; absent in the pistillate flowers. Pistil 2-carpellate, the ovary compressed, the ovules 2, fused to the ovary wall, generally only one maturing, the styles 2, filiform, spreading, deciduous, the stigmas globose or capitate; pistillode sometimes present in the staminate flowers, often very reduced. Fruit nut-like, compressed, with crusty epicarp and hard endocarp. Howard (in Rhodora 61:79-81. 1959) reports that in alcohol-preserved speci- mens of Helosis from the West Indies the filaments are free throughout their length whereas in herbarium specimens the filaments appear to be tightly adherent. In contrast, in herbarium specimens from Central America the filaments appear to be adnate at least at the base; this same basal adnation also is found in alcohol- preserved specimens. Three species from the Antilles and tropical America. A single species from Panama. 1. Helosis mexicana Liebm. in Forh. Vidensk. Skand. Nat. 4:181. 1844. Helosis aquatica Mutis, ex Hook f. in Trans. Linn. Soc. 22:59. 1856. Caldasia mexicana (Liebm.) O. Kuntze, Rev. Gen. 2:590. 1891. Rhizome and inflorescence more or less ferrugineous on drying. Inflorescence with the peduncle 8-16 cm. long, with or without an encircling sheath, the head ovoid or more often ellipsoid, 3-5 cm. long, 1.5-3.0 cm. in diameter, the scales about 3 mm. in diameter, stalk about 2 mm. long, the paraphysoid trichomes clavi- form, 1.0-2.5 mm. long. Staminate flowers: perianth tube tubular, 2-3 mm. long, glabrous, the lobes 3, ovate, 1-3 mm. long, 0.75-1.0 mm. broad; filaments connate, free at the apex at maturity, 0.25-0.75 mm. long, the synandrium 0.5-1.0 mm. (286) flora of Panama (Balanopboraceae) long, 0.5-0.75 mm. in diameter. Pistillate flowers based on Mexican specimens: perianth obscurely bilabiate and becoming fimbriate, minute; ovary tubular to somewhat ellipsoid, 1.0-1.25 mm. long, glabrous, the styles 1.25-1.75 mm. long, deciduous, the stigmas minutely capitate. Fruit 1.25-2.25 mm. long, about 0.5 2. CORYNAEA Hook. f. , Seibert 469. Corynaea Hook. f. in Trans. Linn. Soc. 22:3 1, /. 13, 14. 1856. Itoasia O. Kuntze, Rev. Gen. 2:590. 1891. Rhizomes nodular or rarely elongate, often lobed, lacking scales, sometimes with root-like processes. Inflorescence mostly monoecious and proterogynous or rarely dioecious, claviform, the peduncle encircled at the base with a glabrous annular to cupuliform sheath which is sometimes very obscure, naked above the sheath, the head globose or ellipsoid, when immature covered with 6-sided, peltate, umbonate scales underneath which occurs a dense mat of paraphysoid trichomes and young developing flowers, the scales deciduous prior to anthesis. Perianth of the staminate flowers tubular, conical or subcampanulate, completely connate, crenate; in the pistillate flowers reduced to a low epigynous collar or, according to some authors, absent. Stamens 3, exserted, the filaments completely fused into a (287) 306 ANNALS OF THE MISSOURI BOTANICAL GARDEN staminal column free from the perianth tube, the anthers united into a synan- drium, each with two large lateral pollen sacs, introrse; absent in the pistillate flowers. Pistil 2-carpellate, the ovary compressed, the ovules 2, fused to the ovary wall, generally only one maturing, the styles 2, filiform, spreading, deciduous, the stigmas globose or capitate; pistillode present in the staminate flowers. Fruit nut- like, compressed, with crusty epicarp and hard endocarp. A poorly known genus consisting of perhaps 4 species of Central and South America. Only 1 species is represented in Panama. 1. Corynaea crassa Hook. f. in Trans. Linn. Soc. 22:31, /. JJ, 14. 1856. Itoasia crassa (Hook, f.) O. Kuntze, Rev. Gen. 2:590. 1891. Rhizome an irregularly multi-lobed mass, becoming ferrugineous upon drying. Inflorescence pink, the peduncle with an irregularly short-lobed, basal sheath, 1.5-2.5 cm. long, vertically ribbed, glabrous, the head globose and becoming fusiform, 2—4 cm. long, 1.5-2.0 cm. in diameter, the scales 3-4 mm. in diameter, stalk 2-3 mm. long, the paraphysoid trichomes claviform, 1.5-2.0 mm. long. Staminate flowers not seen. Pistillate flowers: perianth a low epigynous collar or absent (?); ovary obovoid, 1 mm. long, 0.5-0.75 mm. broad, the styles 1.0-1.5 mm. long, the stigmas minutely capitate; pedicel to 0.5 mm. long. Fruit obovoid, about 1.5 mm. long, 1 mm. broad. flora of Panama (Balanophoraceae) 307 Collected flowering in July between 3 500 and 4000 meters. According to Woodson and Schery these plants are parasitic on bamboo roots. chiriqui: Potrero Muleto to summit, Volcan de Chiriqui, Woodson & Schery 470. 3. LANGSDORFFIA Mart. Langsdorffia Mart, in Eschweg. Journ. Bras. 2:178. 1818. Langsdorfia C. A. Agardh, Aphor. 203. 1825. Senftenbergia Klotzsch & Karst. ex Klotzsch, in Linnaea 20:460. 1847, nom. nud. in synon. Rhizome lobed, creeping or upright, often dichotomously branched, tomentose and sometimes glabrescent. Inflorescence dioecious or rarely monoecious, capitate to clavate, the peduncle terete, thick, enclosed at the base by a short, lobed or toothed sheath, above the sheath vested with many imbricate scales, the scales in the basal portion ovate to oval-lanceolate, gradually becoming progressively lanceolate towards the flower-head, the head globose, ovoid or ellipsoid, staminate generally larger than the pistillate, yellow, yellowish-red or red, the staminate flowers sub- tended by reduced pistillate flowers, the pistillate flowers agglutinate into a dense mat, ebracteate. Perianth in the staminate flowers composed of 3 or rarely 2 concave tepals, valvate; in the pistillate flowers reduced to an epigynous collar, obscurely 2- or 4-lobed. Stamens as many as the tepals and opposite them, exserted, the filaments short, connate into a staminal column, the anthers connate below and free above, each with 4 pollen sacs at the base and 2 at the apex, extrorse; absent in the pistillate flowers. Pistil 1, the ovules 1, fused to the ovary wall, the style 1, erect, filiform, deciduous, the stigma minutely papillate; pistillode absent in the staminate flowers. Fruit drupaceous, with fleshy epicarp and hard endocarp. Apparently only a single species from Mexico to southern Brazil. 1. Langsdorffia hypogaea Mart, in Eschweg. Journ. Bras. 2:178, t. 5. 1818. Langsdorffia janeirensis Rich, in Mem. Mus. Par. 8:412, t. 19. 1822. Langsdorffia moritzuma Klotzsch & Karst. in Linnaea 20:461. 1847. Thonningia janeirensis (Rich.) Liebm. in Forh. Skand. Naturf. Christ. 180. 1847. Thonningia mexicana Liebm. loc. cit. 1 847. Langsdorffia rubiginosa Wedd. in Ann. Sci. Nat. ser. 3. 14:187. 1850. Senftenbergia moritziana (Klotsch & Karst.) Eichl. in Mart. Fl. Bras. 4 2 :10. 1869, nom. Rhizome digitiform, canescent-tomentose. Staminate inflorescence clavate, the peduncle 2-5 cm. long, scaly, surrounded at the base by a lobed sheath, the lobes deltoid, 0.5-1.0 cm. long and broad, canescent-tomentose, the scales stiff, ciliate, enclosing the head when immature, lower scales ovate, about 1 cm. long, 5-7 mm. broad, upper scales lanceolate, to 3.5 cm. long, 1 cm. broad, the head ellipsoid, 5-7 cm. long, about 3 cm. in diameter. Staminate flowers based on Brazilian specimens: pedicel 6-10 mm. long; tepals 3 (or 2), free, ovoid, about 3 mm. long, 1.5-2.0 mm. broad, becoming widely reflexed, glabrous; stamens 3, the filaments about 0.5 mm. long, completely connate, the anthers connate below, free above. Pistillate inflorescence generally capitate to clavate, the peduncle and vesture as in the staminate inflorescence, the head subglobose, 2-4 cm. long, 4.5 cm. in diameter. Pistillate flowers: free but tightly agglutinated with one another above due to a (289) ANNALS OF THE BOTANICAL GARDEN nto a curious tubular or obovoid diameter, surrounding the style and h it, the perianth lobes 2, short; ovary ellipsoid, about diameter, the style 1, 0.5-1.5 mm. long, slender, spirally tely capitate. Fruit ellipsoid, about 1 mm. long, 0.5 mm. twisted, the stigma The only specimen examined was collected flowering and fruiting altitude of 2000-2500 meters. chiriqui: forested ridges south of Finca Lerida, Allen 4.773. flora of Panama (Aristolochiaceae) 309 ARISTOLOCHIACEAE By HOWARD VM. PFEIFFER Scandent lianas, herbs, or rarely trees, the wood often having broad inter- fascicular rays. Leaves alternate, spiral, estipulate, simple, palmately veined, usually entire. Flowers mostly axillary, perfect, actinomorphic or zygomorphic, hypogynous to epigynous, in racemes, cymes, solitary or in clusters and cauli- florous, rarely terminal, large and showy or small, often with fetid or resinous odor. Calyx gamosepalous, often petaloid and elaborately contorted and lobed. Corolla of 3 reniform petals in Saruma, rudimentary in some species of Asarum, absent in all other members of the family. Stamens 6 to 12 or occasionally many, in one or two whorls, free, connate or adnate to the short, fleshy, united styles. Ovary apocarpous to syncarpous, 6- to 4-carpellate and -loculate, with marginal to axile placentation; ovules many to few, anatropous. Pollen monosulcate to nonaper- turate, spherical. Fruit follicular or capsular with septicidal, rarely septifragal or irregular dehiscence, often dehiscing acropetally; seeds numerous to few, with a small basal embryo in abundant endosperm. The Aristolochiaceae, or Birthworts, are distributed throughout the tropic and temperate regions of the world. The genera are poorly defined and are in need of competent revision. The largest genus, Aristolochia, is the sole representative of the family in Panama. It is a large genus with about 300 species in tropic and temperate areas throughout the world. Eleven species are reported from Panama, but further collections will probably double this number. The leaves of Aristolochia are remarkably variable. For example, on a single shoot about 2 dm. long of Aristolochia panamensis leaves may be seen which are narrowly linear-lanceolate, broadly ovate, and broadly obtriangulate with a deeply emarginate apex. As a consequence, leaf characteristics have been used as infre- quently as possible in the keys. An exception to this general rule may be noted in that group which bears pseudostipules. The Aristolochiaceae, it will be recalled, are estipulate; however, in these species, an axillary bud produces a single, small leaf which is amplexicaul and nearly sessile. It frequently is ruffled, not having the shape of the ordinary leaves on the shoot, and looks strikingly like a pair of connate, auriculate stipules. The description of a flower as complex as that of Aristolochia is always a problem. The species differences of greatest importance lie in the elaborate inflations and expansions of the calyx tube. In an attempt to standardize the descriptions, therefore, it has seemed convenient to subdivide the calyx tube into several morphological portions and to treat each of these separately. It should be emphasized here that not necessarily all of these subdivisions are present in any An explanation of these arbitrary designations follows. As the calyx arises from the apex of the ovary, it expands into a large, inflated, usually gibbous portion called the utricle. The utricle narrows into the cylindrical portion known as the tube. This expands and usually imperceptibly merges with the expanded limb. In (291) 310 ANNALS OF THE MISSOURI BOTANICAL GARDEN the larger flowers the tube has a peculiar venation at its distal end which sub- divides the tube into two portions, conveniently described as the tube proper, and the throat. The throat is usually funnelform and as a result tends to lend greater width and expansion to the limb. One constantly finds reference in the literature to the fetid odor of Aristo- lochia flowers. While many of them are truly fetid and evil smelling, many are odorless and a few have been described as having either a sweet or a resinous odor. The flowers ostensibly are pollinated by various species of Diptera. In herbarium specimens, dissection of the utricle often reveals insects which have burrowed or forced their way between the lobes of the styles. The flowers are equipped with rigid, inward-pointing hairs throughout the length of the tube. The Diptera, doubtless attracted by the color and odor of the blooms, are trapped within the utricle since they are able to pass only inward over the hairs. The flowers are protogynous. With pollination, the anthers dehisce and the rigid hairs in the tube wilt, allowing the flies to escape and carry pollen to another bloom. Several species of Aristolochia furnish the serpentaria of medicine which is used as a tonic and as a febrifuge. Many of the plants are utilized as a native remedy for snakebite and bear the common name Snakeroot. The remarkable shapes of the calyx tubes lead to various names such as Dutch- man's Pipe, Pelican Flower and Goose Flower. The name Aristolochia, which is derived from the Greek aristos, best, and lochia, delivery, refers to its supposed medicinal properties in connection with the alleviation of the pains of childbirth; the curved flower, with summit and base together suggesting, by the doctrine of signatures, the human fetus in the womb. Standley, writing about the aristolochias of El Salvador, reports that the roots of guaco or A. anguicida are used by the natives to wash clothes, to scour out dirt, and also as a remedy for stomach ache. He further states that infusions of A. arborescens are used as a remedy for venereal diseases in the female and for dysentery in children. 1. ARISTOLOCHIA L. Aristolochia L. Sp. PI. ed. 1. 960. 1753. hotrema Raf. in Amer. Monthly Mag. & Crit. Rev. 4:195. 1819. Hocquartia Dum. Comm. Bot. 30. 1822. Dasyphonion Raf. First Cat. Bot. Gard. Transylv. Univ. 13. 1824. Cardiolochia Raf. ex Rchb. Consp. 85. 1828. Einomeia Raf. Medic. Fl. 1:62. 1828. Endodeca Raf. loc. cit. 62. 1828, sphalm. Pistolocbia Raf. loc. cit. 62. 1828. Siphisia Raf. loc cit. 62. 1828. Isipbia Raf. Medic. Fl. 2 : 2 3 2. 1830. Dictyanthes Raf. in Nipbus Raf. loc. cit Stpbidia Raf. loc. cit. 247. 1832. Ambuya Raf. Fl. Tellur. 4:98. 1836. Diglosselis Raf. loc. cit. 98. 1836. Endotbeca Raf. loc. cit. 98. 1 836, corr. Endodeca (1828). flora of Panama (Aristolochiaceae) Hexaplectris Raf. loc. cit. 97. 1836. Plagistra Raf. loc. cit. 98. 1836. Psophiza Raf. loc. cit. 99. 1836. Guaco Liebm. in Forhandl. Skandin. Naturf. 1844:203. 1847. Howardia Klotzsch, in Monatsb. Acad. Berl. 1859:607. 18 59. Lianas, rarely upright perennial herbs, shrubs or trees. Leaves alternate, petiolate, estipulate but an axillary bud often producing a clasping, broadly reni- form leaf (pseudostipule) ; blade entire or 3 to 7-lobed, often cordate, palmately or pinnately veined, variable. Flowers axillary, solitary, perfect, epigynous, zygo- morphic. Calyx gamosepalous, variously inflated, thence more or less contracted, expanding into a 1- to 3-lobed limb. Corolla absent. Stamens typically 6, the anthers sessile and adnate to the style, 2 -celled, dehiscing longitudinally. Ovary inferior, 6-loculate; placentation axile, the ovules numerous, anatropous. Styles 6, marginally connate, fleshy, with coroniform stigmatic lobes, or essentially capitate. Fruit a capsule, septicidally or septifragally 6-valved, often dehiscing acropetally. Seeds numerous, horizontally compressed in vertical rows, the embryo rudimentary in abundant endosperm. In the treatment of Aristolochia presented here, it has seemed the wisest choice to include, in addition to the species collected in Panama, accounts of those reason- ably certain to be found there. Accordingly, these species will be found incor- porated into the key to species and described with the presently known groups in Perhaps a word of caution might be of some help when attempting to key flowering material in this genus. Many of the collections of the large-flowered species are collections of buds, and dimensions derived from them must be regarded as approximate. For example, the flowers of Aristolochia grandiflora become fourteen feet long but few collections reflect this great dimension. The buds un- doubtedly develop for weeks before anthesis and few collectors are willing to return to the same location repeatedly for the sake of a single bloom; hence the measured length of the calyx tube as collected may be considerably less than the mature size. d. Leaves (other i ANNALS OF THE MISSOURI BOTANICAL GARDEN slightly narrowing iry immediately subtended by a folia 'Es-.-J: long-acuminate; flowers small, white "■ l^seV,",';'"' ^'" [ J,:"" - ,,o "-"» mi " t « i kk ' 3°™" b Zo« n many flowered raceme, in ,,', o leaves "'•^S^.T.S, r ha :^irn J 1 :ri^ n i:;t; deeply reflexed on the utricle, < HBK. Nov. Gen. 1817. Spec. Arhtolochia clausseni Duch. in Ann. Sc. Nat. ser. 4. 2:57. 1854. Aristolochia emarginata Willd. ex Duch. in DC. Prod. 15-1:466. 1864. Arhtolochia pusilla Pohl, ex Duch. loc. cit. 1864. Aristolochia tenera Pohl, ex Duch. loc. cit. 1864. Aristolochia subclausa Wats, in Proc. Amer. Acad. 20:372. 1882. Aristolochia Pyrinea Taub. in Engl. Bot. Jahrb. 21:426. 1896. Aristolochia exigua Lindm. in Bull. Herb. Boiss. ser. 2. 1:525. 1901. Herbs, 3 to 6 dm. tall, perennial, with thickened fusiform roots. Leaves alter- nate, somewhat distichous, reniform to cordate, ca. 3 cm. broad, 2 to 3 long, glabrous. Pseudostipules absent, rers small, axillary, rectilinear, the :>e and monolobate limb nearly indistinguishable as such, ca. 2 to 4 cm. long, greenish yellow. Fruit cylindric, ca. 2 cm. long, 1 cm. wide, dehiscing acropetally. Seeds numerous, horizontally compressed, 2 mm. wide, 0.2 mm. thick. Columbia and Venezuela. To be ularifolia sought in Panama. (294) FLORA OF (Aristolochiaceaej Wash. Acad. Sci. 15:5. 1925. Decumbent subshrubs, to 6 dm. tall. Leaves alternate, spiral, very variable in shape, narrowly lanceolate to broadly long, 3 fide. Pseudostit absent. Flowers cauliflorous, borne near the soil in inconspicuous short racemes, ca. 5 cm. long along the curved axis, brownish marked with purple, the utricle narrow, thence widening into a gibbous sac, the limb sharply reflexed on the utricle, the tube reduced to a constriction of the utricle. Gynostemium of 6 closely connate styles, capitate. Fruit cylindric, ca. 3 cm. long, 1 cm. wide, dehiscing api- cally. Seeds several, pyramidal, grooved, Known only from Panama. Las Coco Standby 29515, 25/40; Rio Paraiso, abcr EastVaraiso,Standley 29006 (type) ; RioPed: Miguel, near East Paraiso, Standley 20950. east of Gualaca, alt. 500 ft., Allen 5040; vicinity of San m., Pittier 5463; San Felix, eastern Chiriqui, Pittier 5/50. 3. Aristolochia trilobata L. Sp. PL ed. 1. 2:960 Aristolochia trifida Lamk. Encyc. 1:251. 1783. Aristolochia triloba Salisb. Prod. 214. 1796. Aristolochia macroura Gomez, in Mem. Ac. Lisboa 3:77. Aristolochia caracasana Spreng. Syst. 3 :753. 1826. Aristolochia appendiculata Veil. Fl. Flum. 9: t. 98. 1827. audata Booth, ex Lindl. in Bot. Reg. /. 1 453 Aristolochia tapetotricha Lem. Illustr. Hort. 3:misc. 22. ] Howardia trifida Klotzsch, in Monatsb. Acad. Berl. 18 59:i Twining, glabrous lianas. Leaves alternate, spiral, subpalmate, 3-lobate, trun- cate at the base, glabrous, ca. 6 cm. wide, 6 cm. long. Pseudostipules present. Flowers axillary, ca. 15 to 20 cm. long, red-brown with purple stripes, the utricle elliptic, ca. 5 cm. long, 3 cm. wide, curving abruptly at the apex with slight nar- rowing into the tube, ca. 5 cm. long, 2 to 3 cm. wide, ending in the bialate limb which terminates in a long filiform extension. Seeds numerous, horizontally compressed, 6 mm. diameter, 0.75 mm. thick. Throughout tropic Central and South America. 314 ANNALS OF THE MISSOURI BOTANICAL GARDEN bocas del toro: vicinity of Chiriqui Lagoon, von Wedel 1 404, 1 Chagres, Isthmus of Panama, Fendler 445, 440. 4. Aristolochia odoratksima L. Sp. Pi. ed. 2. 2:1362. 1763 Arhtolochia martiniana Standi, in Publ. Fie (296) FLORA OF PANAMA (AHstoloch Lianas, twining, scandent, glabrous. Leaves alternate, variable cordate-hastate to sub-trilobate, membranous, ca. 15 cm. long, 12 cm. wide. Pseudostipules present. Flowers axillary, the calyx borne at an abrupt angle at the apex of the ovary, dark red, mottled and striped with lighter shades of red and yellow, the utricle ovate, 2 to 3.5 cm. long, the tube arising at a right angle from the side of the utricle, 15 to 20 mm. long, the limb thick, delicate, crenate, suborbiculate, 6 to 12 cm. diameter. Fruit cylindric to subfusiform, abruptly bent in a right angle at the apex, slightly curved, 1.5 cm. diameter, 4 cm. long, dehiscing acro- petally. Seeds numerous, horizontally com- pressed, 5 mm. diameter, 0.5 mm. thick. Southern Central America, northwest- ern South America, Amazonia. >cas del toro: Changuinola Valley, Dunlap 536; vicinity lentos, Mariano Creek, von Wedel 2903; vicinity of Chiriqui, ' 204.6; vicinity of Chiriqui Lagoon, Water Valley, von We, iui Lagoon, von Wedel 1200; Water Valley, v on Wedel 791. HBK. Nov. Gen. & Spec. 2:145. 1817. Slender, scandent lianas. Leaves alternate, ovate, acute, cordate, 3 to 5 nerved, membranous, ca. 10 cm. long, 8 cm. wide, glabrous. Pseudostipules present. Flowers solitary, axillary, rectilinear, borne on young leafy shoots, 4 to 6 cm. long, yellow spotted purple, the utricle ovoid, 1 to 1.5 cm. long, narrowing obliquely to a short tube 7.5 mm. long, acuminate. Fruit cylindric, ca. 3.5 cm. long, 1 cm. wide, dehiscing acropetally. Seeds numerous, horizontally compressed, 2 mm. diameter, 0.5 mm. thick. Central America. canal zone: Camino de Corozal, Bro. Heriberto 247. cocle: between Aguadulce and the Chico R., alt. 20 m., Pittier 5008; Penonome and vicinity, alt. 50-1000 ft., Williams 204. herrera: vicinity of Chitre, alt. 20 m., Allen Iioi. panamA: vicinity of Pacora, alt. 3 5 m., Allen 2032; Argicultural Experiment Station at Matias Hernandez, Pittier 6872; along the Corozal Road, near Panama, Standley 2677O; vicinity of Juan Franco Race Track, near Panama, Standley 27796; near Punta Paitilla, Standley 26246; Taboga Island, Standley 27029, 27963; Tumba Muerto Road, near Panama, Standley 29796; sabanas, north of Panama City. Bro. Paul 587; Panama, Hayes 781, 294, 146. ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 94. Aristolochia inflata 6. Aristolochia anguicida L. Sp. PI. ed. 2. 2:1 bia loriflcn :: Lianas, with slender stems. Leaves alternate, spiral, oblong-ovate, cordate, 7 to 10 cm. broad, 10 to 15 cm. long, glabrous. Pseudostipules usually present. Flowers axillary, rectilinear, ca. 2.0 to 2.5 cm. long, mottled purple, the utricle gib- bous, 0.9 cm. long, narrowing abruptly into the tube, ca. 1.0 cm. long, which widens gradually into a funnelform, un- equally monolobate limb. Fruit cylindric, 3 cm. long, 1.5 cm. wide, dehiscing acropetally. Seeds numerous, horizontally Fig. 95. Aristolochia anguicida compressed, 0.4 cm. diameter, 1 mm. Honduras, Guatemala and south to Venezuela and Colombia. To be sought (298) FLORA OF (Aristolochiaceaej 317 318 ANNALS OF THE MISSOURI BOTANICAL GARDEN 7. Aristolochia arborescens Linn. Sp. PL 960. 1753. Aristolochia foetens Lindl. Bot. Reg. t. 1824. 1836. Howardia foetens Klotzsch, in Monatsb. Acad. Berl. 1859:610. 1859. Lianas, with twining stems. Leaves alternate, cordate, apex acute to : membranous, 5 to 20 cm. broad, 8 to 30 cm. long, glabriusculus at maturity. Pseudostipules absent. Flowers immediately subtended by an orbiculate, perfoliate bracteole, axillary, solitary but occasionally in pairs, glabrescent, yellow, blotched with purple, the utricle broadly gibbous-clavate, ca. 30 cm. long, the tube sharply reflexed on the utricle, ca. 35 cm. long, widening into a throat distinguished by peculiar annular venation and constriction, ca. 5 cm. long, the limb obliquely funnelform, terminating in a struppate extension ca. 20 cm. long; the odor putrid. Fruit ovate- cylindric, ca. 10 cm. long, 6 cm. wide, dehiscing acropetally. Seeds Mexico south to Colombia. r 4145. darien: vicinity Pinogana, 8. Aristolochia grandiflora Sw. Prod. Veg. Ind. Occ. 3:1566. 1806, non Gomez, nee Arruda, nee Vahl. Aristolochia gigas Lindl. Bot. Reg. t. 60. 1842. Aristolochia grandiflora /3 hookeri Duch. in DC. Prod. 15-1 :473. 1864. Twining lianas, older stems ridged with cork. Leaves alternate, cordate, apex acute to acuminate, membranous, 5 to 20 cm. broad, 7 to 30 cm. long, densely pilose to velutinous becoming nearly glabrous on upper surface. Pseudostipules absent. Flowers immediately subtended by an acuminate foliaceous bracteole, borne on young stems, axillary, solitary, ca. 3 m. long, the color white blotched with purple, pilose, the utricle broadly gibbous-clavate, ca. 45 cm. long, the tube sharply reflexed on the utricle, ca. 40 cm. long, the limb obliquely salverform, terminating in a filiform extension from the medial lower edge, the odor of "old tobacco". Fruit ovate- cylindric, 10 cm. long, 4 to 6 cm. wide, dehiscing acrop- etally. Seeds numerous, horizontally compressed, 1 cm. diameter, 3 mm. thick. Panama, south throughout Amazonia. darien: vicinity of Yape, alt. 30 m., Allen 647. This species rarely is collected. It appears that practically all of the large- flowered Aristolochia collections have been named grandiflora, but are properly A. arborescens. There is a relatively sharp distinction between the two species. A. grandiflora has truly a huge flower, perhaps the largest in American tropics, a total reported axial length in excess of 4.5 meters, while A. arborescens is about one- fourth as large. The flowers of the former are velutinous without and differ in color and odor from the latter. A. arborescens has a well-differentiated throat, while A. grandiflora is not markedly endowed with this feature. 9. Aristolochia costaricensis (Klotzsch) Duch. in DC. Prod. 15-1:450. 1864. Howardia costaricensis Klotzsch in Monatsb. Acad. Berl. 1859:614. 1859. :'. Publ. Bot. 6:51. 1914. Aristolochia haughtiana Hoehne, in Arquiv. Bot. Estad. S. Paulo 2:99. 1947. flora of Panama (Aristolochiaceae) 319 ANNALS OF THE MISSOURI BOTANICAL GARDEN i 15 c hispid-pilose , spiral, cor- broad, 10 to 25 cm. long, upper surface glabrescent, lower pilose. Pseudostipules absent. Flowers axillary, solitary, ca. 7 cm. long, the color brownish-purple marbled with white, densely pilose-hispid, the utricle ellipsoid, 3 cm. long, the tube arising obliquely and abruptly from the side of the utricle near the distal end, 2 cm. long, ** gradually enlarging into the obliquely linguiform lobe variously spotted and mottled and nany fleshy excrescenses or fimbriae. Fruit ovoid, 6 cm. long, 4 cm. wide, dehiscing acropetally. Seeds i mpressed, 1 cm. diameter, 2 mm. thick. Tropic Central and northern South America. -iiauKuiiiuia valley, Dunlap l8; vicinity of Chiri- — / I228 > 1267. darien: vicinity of Yape, alt. 30 m., Allen 864. Alhajuela, Chagres Valley, alt. 30-100 m., Pittier 2326; Mamoni R., above t. 23-25 m., Pittier 4732; north of Panama City, Bro. Paul 374. province *: western Panama. Stork 18. J '* 10. Aristolochia . broad, 15 to Duch. in DC. Prod. 15-1:458. 1864. Howardia veraguerisis Klotzsch, ex Duch. I spiral, ovate, apex long deeply cordate, 12 to 20 c 30 cm. long, veins and adjacent tissues white, glabrous. Pseudostipules absent. Flowers cauliflorous in short racemose clusters, small, purple, red-reticulate, the utricle subspheric, ca. 1 cm. long, narrow- ing into the 1.5 cm. long tube which unequally enlarges to form the 2 cm. long mucronate limb, which is arcuately reflexed upon the utricle. Fruit not seen. East Panama, south to Brazilian darien: El Real, Allen 2210. flora of Panama (Aristolochiaceae) 321 l Standi. Journ. Wash. Acad. Sci. 15:5-6. 1925. Slender, scandent lianas. Leaves broadly rounded-deltoid, acuminate or acute, the base very shallowly cordate to truncate, 8 to 12 cm. broad, 10 to 20 cm. long, dark green and glabrous above, beneath light green to white, subarachnoid- velutinous to thinly puberulent. Pseudostipules absent. Flowers cauliflorous in short, few-flowered racemes, maroon finely marbled with white and pink, the utricle ellipsoid, 6 cm. long, the tube short and reflexed, thence opening into the showy limb which is notched at the upper medial edge, ca. 20 cm. wide. Fruit long-cylindric, 12.5 cm. long, 3 cm. wide, dehiscing acropetally. Seeds numerous, revolute, 3 mm. diameter, 8 mm. long, 2 mm. thick. Known only from Panama. MISSOURI BOTANICAL GARDEN bocas del toro: Changuinola Valley, Dunlap 247. cai> 2250; Barro Colorado Island, Shattuck 640 ; Barro Colorado 1 Standley 40866; hills north of Frijoles, Standley 27469 (type) ; 900 m., Wetmore & Abbe I. darien: Trail between Paya an 12. Aristolochia Standi, in Contrib. Arn. Arb. 5:< Aristolochia maxima var. cordata Standi, in Field Mus. Publ. Bot. 8:136. 1930. Twining, scandent lianai 5 , sparsely hispidulous-pubesce nt. Leave! subdistichous, obl( :>ng-spatu late, apex acute to obtuse, base typically deeply cordate, 3 to 6 cm . broad, 10 to 20 cm. long, glabrescent. Pseudostipi iles absent. Flowers solitary, a ixillary, rec :tilinear or slightly arched, dark purple-1 Drown, the utricle long-ellipsoi d, ca. 5 err l. long, the tube 3 cm. long, splitting mc molaterally ) the oblong-1 ceolate limb, long. Fruit ovate-cylindric, ca. 12 cm. long, 5 cm. wide, dehiscing acropetally. Seeds numerous, horizontally compressed, 1 cm. diameter, 2 mm. thick. Central America and northern South , Bangham 455; Cerro Gordo, near Culebra, alt. ■ 2304; Barro Colorado Island, Shattuck 413 (type); Barro Colorado Island, Shattuck 1076; Barbour-Lathrop Trail, alt. 900 m., Wetmore fif Woodworth 77; Barro Colorado Island, Zetek 3618. cocle: between Las Margaritas and El Valle, Woodson et al. 1739; above Penonome, alt. 2000-3000 ft., Williams 548. Panama: San Jose Island, Erlanson 424; San Jose Island, Johnston 726, 1253, 1 316. 13. Aristolochia ] kaL. Spec. Pled. 2. 2:1361. 1763. Aristolochia geminiflora HBK. Nov. Gen. & Spec. 2:11! Aristolochia reticulata Seem. Bot. Voy. Herald, 193. II Howardia geminiflora Klotzsch, in Monatsb. Acad. Berl Howardia goii t . 1859. Howardia hoffn;. .621.1859. Howardia maxima Klotzsch, loc. cit. 615. 1859. Aristolochia biflora Willd. ex Duch. in DC. Prod. 15-1 Aristolochia mathewsii Duch. loc. cit. 497. 1864. Aristolochia maxima a maxima Duch. loc. cit. 456. I81 Aristolochia maxima geminiflora Duch. loc. cit. 457. Aristolochia maxima 7 angustifolia Duch. loc. cit. 186 Aristolochia reticulata J. F. Holton, ex Duch. loc. cit. Aristolochia sprucei Mast, in Mart. Fl. Bras. 4-2:88. 1 Aristolochia asperifolia Ule, in Verh. Bot. Ver. Prov. Bi (304) FLORA OF Panama (Polygonaceae) Flo Pseudostipules ab: stered racemes v i purple-brown, irrowing utricle ovoid, ca. 3 cm. Ion, abruptly into the recurved tv long, which gradually widens into the strongly reflexed limb, ca. 3 cm. long. Fruit ovate-cylindric, ca. 15 cm. long, 7 cm. wide, dehiscing acropetally. Seeds numerous, horizontally compressed, 1.2 Central America and tropic South America. Matias Hernandez, Standby 31839; Taboga Island, Standley 27090, 27856; Tumba Muerto Rd., near Panama, Standley 29804; sabanas, north of Panama City, Bro Paul 517; Belleviota, Macbride 2763. Aristolochia maxima POLYGONACEAE By JAMES A. DUKE Flowers perfect or unise: on short usually articulate pedicels, solitary or fasciculate within scariose ochreolae. Calyx uniseriate or biseriate, hypogynous, of 3-6 free or partially connate tepals. greenish to red or white. Stamens usually 6-9, rarely more or less, the filaments filiform or flattened, occasionally partially adnate to the calyx; anthers 2- or 4- locular, usually versatile and introrse. Ovary superior, triquetrous or lenticular, unilocular, containing a single erect orthotropous ovule, the stigmata filiform or capitate and entire or variously fringed. Achene trigonous or lenticular, usually with a crustaceous pericarp, the embryo usually excentric in a mealy endosperm. Herbs, shrubs, trees or vines with alternate usually entire simple leaves with ochreate stipules, the stems often sulcate, hollow and geniculate. Flowers in terminal racemes or panicles, often spicate, rarely cymose, or solitary or fasciculate in the axils, usually subtended by scariose ochreolae. (305) 324 ANNALS OF THE MISSOURI BOTANICAL GARDEN This predominantly north temperate family of about 30 genera and 700 species is represented in Panama by six genera. It is economically a rather unimportant family. Fagopyrum, the buckwheat, and Rheum, the rhubarb, are crops of limited importance. Antigonon spp., Polygonum spp. and Muehlenbeckia spp. are often cultivated in Central America as ornamentals. Some species of Rumex ire troublesome lawn weeds. eous or suffruticose (spp. of Polygonum § tinaria twin- F 3-6 tepals, the inner slightly larger; stamens 5-9. ; tepals, the outer reflexed or spreading, the inner usually v folded, often tuberculate; flowers perfect -5 subequal tepals, not reflexed, closely conforming to the >y, arboreal or lianoid: calyx of 5-6 tepals. the outer ircely if at all exceeded by the tepals; tepals discrete and ; tepals 5, the outer only slightly larger than the inner; perfect; tepals cordate, discrete, becoming scariose, dioecious or polygamo-dioecious (perfect in Muehlen- llary and terminal, dioecious or perfect; o their base; scramblers (in Panama), the les or pistillodes developed; tepals forming at least at their bases; trees in Panama with bilateral wood and lenticular twigs). :eded by 3 wing-like outer tepals; outer used to form a tube free of Rumex L. Sp. PI. 333. 1753. Flowers perfect or unisexual, pedicellate, few to many in verticillate ochreolate fascicles. Perianth of 6 tepals, the outer 3 usually smaller and spreading or reflexed, the inner 3 cordate, entire or lacerate, frequently with an abaxial tubercle, usually accrescent and closely investing the achene. Stamens 6, discrete, the filaments usually shorter than the anthers, the anthers mostly 2-locular and loculicidal. Ovary trigonous with 3 spreading to reflexed filiform styles capped by fimbrillate peltate stigmata. Achene triquetrous, smooth and lustrous or rough and dull, usually included. Glabrous usually perennial herbs, occasionally becoming quite tall and shrubby. Leaves alternate, entire or rarely dentate, occasionally tending to form rosettes, the ochreae hyaline and rather tardily deciduous. Of this large predominantly temperate genus, only two species, both probably naturalized from Europe, are thus far represented in the flora of Panama. Here they tend to be weeds in lawns and pastures at higher elevations. R. costaricensis (306) flora of Panama (Polygonaceae) Rechinger, a strikingly large species, is apparently endemic to the high mountau of Costa Rica. Three other species are reported in Guatemala by Standley Steyermark (in Field Mus. Bot. 24* : 13 1. 1946) . 1 mm. long; leaves 1 326 ANNALS OF THE MISSOURI 1. Rumex crispus L. Sp. PL 335. 1753. Mostly unbranched coarse glabrous perennials with yellowish taproots. Leaves glabrous, lanceolate to oblong, apically acute, basally acute to cordate, the cauline blades mostly 10-12 cm. long by 0.5-3 cm. broad; petioles 1-5 cm. long; ochreae up to 2.5 cm. long, tardily deciduous. Inflorescences of crowded many-flowered ochreolate verticils in racemes or panicles. Flowers perfect, on pedicels mostly 5-12 mm. long; outer tepals oblong, 1-2 mm. long, scarcely accrescent; inner tepals broadly ovate to cordate, accrescent becoming 3.5-5 mm. long, 2-3.5 mm. broad, with a prominent exterior tubercule developing on 1 or 3 of them; stamens 6, discrete, the filaments mostly less than 0.5 mm. long, the anthers 1-1.5 mm. long, basifixed; ovary trigonous, the 3 styles usually reflexed, with 3 peltate fimbrillate stigmata. Achenes triquetrous, brown, lustrous, included, often capped by the persistent styles and stigmata, 2.5-3 mm. long, 2-2.5 mm. broad. chiriqui: Alto Lino, wayside roadside, 4200 ft., Bro. Maurice 883. A native of Eurasia, this is a widespread weed, and common in temperate por- tions of the Americas. In Guatemala it is called lengua de vaca and lengua de caballo. The leaves are occasionally cooked as a potherb. 2. Rumex acetosella L. Sp. PL 538. 1753. Acetosella acetosella (L.) Small, Man. SE. Fl. 446. 1933. Low delicate glabrous perennials with slender creeping rootstocks. Leaves glabrous, lanceolate-hastate, apically acute to rounded, 1-3 cm. long, 3-8 mm. broad; petioles 1-5 cm. long; ochreae mostly 0.5-1 cm. long, tardily deciduous. Inflorescences of panicles of remote few-flowered verticils. Flowers dioecious, on pedicels mostly less than 3 mm. long; outer tepals oblong to narrowly ovate, about 0.5 mm. long, scarcely accrescent; inner tepals broadly ovate, about 1 mm. long, scarcely accrescent, not tuberculate; stamens 6 discrete, the filaments less than 0.3 mm. long, the anthers 0.5-1 mm. long, bilocular and loculicidal; ovary trigonous, the 3 styles spreading, with 3 peltate fimbrillate stigmata. Achene triquetrous, brown, dull and rough, often slightly exserted, about 1 mm. long. chiriqui: Cerro Punta, ca. 6025 ft., P. White 196; Valley of the upper Rio Chiriqui Viejo, P. White 60. This species, like the former, is a native of Eurasia 1 regions of the Americas. In Costa Rica it is called 1 sistent rootstocks, this is often an obnoxious weed of lawns and pastu believed to have been imported into Central America : 2. POLYGONUM L. Polygonum L. Sp. PL 359. 1753. Flowers perfect, occasionally with tendencies toward suppression with pedicels distally articulated, usually in ochreolate fascicle Peria (308) flora of Panama (Polygonaceae) 327 subequal partially connate tepals, white, green, red or pink. Stamens 5-8 (-9), the filaments discrete, often unequal in length, occasionally adnate to the tepals, the anthers small, versatile, introrse, often isodiametric and appearing peltate. Ovary lenticular or trigonous; styles 2 or 3, terminated by capitate stigmata. Achene usually included, lenticular or trigonous, often beaked, the facies ovate or orbicular. Annual or perennial herbs, twining, occasionally shrubby. Leaves alten ally sagittate, cordate or hastate, the blades mostly exceeding the petioles; ochreae usually conspicuous, often fringed with strigose cilia. Inflorescences of terminal spicate panicles, racemes or cymes, in some species the flowers solitary or fasciculate in the axils; rhachises glabrous to sericeous, occasionally glandular. This largely temperate genus contains about 150 species, about half of which occur in North America. Many of the herbaceous species are aquatic or bog plants; some are rather obnoxious weeds. Shrubby species are often cultivated as ornamentals. The sections of the genus have often been elevated to generic status, but they are here maintained at the subgeneric level. Heterostyly, tendencies toward the suppression of one sex in the flowers, environmental modifications, and hybridization have all contributed to the difficult taxonomy of the section Per- sicaria, to which belong five of the six Panama species. Fourteen species are known from Guatemala and four from Costa Rica. Dchreae with reflexed or spreading herbace d. Tepals, leaves and ochreae conspicuously dark-punc of the leaves subglabrous for the veins, glabrous 5. P. mexicanum aa. Plants twining; leaf bases cordate; inflorescences of lax axillary, fre- quently leaf-, "ted 1. Polygonum hispidum HBK. Nov. Gen. & Sp. 2:178. 1817. Robust glutinous-hispid perennials. Leaves punctate, hispid especially on the veins, entire but ciliate on the margins, apically attenuate, the bases decurrent on the petioles; blades broadly lanceolate to ovate, 10-30 cm. long, 5-9 cm. broad; ochreae 1-3.5 cm. long, hispid, the summits capped by herbao flanges. Inflorescences of terminal continuous ochreolate subspic (W) 328 ANNALS OF THE MISSOURI BOTANICAL GARDEN panicles, the peduncles strigose, the floriferous portions mostly 3-10 cm. long and 0.5-1 cm. broad, the flowers fascicled in ochreolae 2-4 mm. long. Flowers white to dark rose, the pedicels apically articulated and about equaling the ochreolae, 3-4 mm. long; tepals 5, subequal, the outer 4-5 mm. long, the inner somewhat shorter; stamens 5, discrete, affixed just above the base of the tepals; filaments 1.5-2 mm. long, the anthers about 0.5 mm. in diameter, subrotund, versatile, thus appearing peltate; ovary lenticular; styles 2, about 2 mm. long, connate for about 1 mm., the stigmata at anthesis about equaling the anthers. Achenes lenticular, orbicular, often with a concavity on one or both facies, black, lustrous, 3-4 mm. long, the persistent styles slightly over 1 mm. long. canal zone: vicinity of Gatuncillo, Piper 5624. Panama: in swamps on edge of jungle; along road between Panama and Chepo, Dodge, Hunter ,Steyermark & Allen 1667 2. This paludal or aquatic species, occurring chiefly at low elevations, is probably indigenous to South and Central America and the West Indies. 2. Polygonum punctatum Ell. Bot. S. C. & Ga. 1:445. 1817. Polygonum hydropiper Michx. Fl. Bor. Am. 1:238. 1803. not L. 1753. Polygonum hydropiperoides Pursh, Fl. Am. Sept. 270. 1814. not Michx 1803 Polygonum acre HBK. Nov. Gen. & Sp. 2:179. 1817. Polygonum maritimum Veil. Fl. Flum. 4. /. jp. 1827. Polygonum antihaemorrhoidale f. riparium Mart. Reise. 550. 1828. Polygonum antihaemorrhoidale f. aquatile Mart. Reise. 550. 1828. Polygonum antihaemorrhoidale a riparium Mart, in Linnaea 5: litt. 41. 183 0. Polygonum antihaemorrhoidale p aquatile Mart. loc. cit. 1830. Polygonum acre a aquatile Meissn. in Mart. Fl. Bras. S 1 :^. 1855. Polygonum acre p riparium Meissn. loc. cit. 18. 1855. Polygonum acre a confer tiftorum Meissn. in DC. Prodr. 14:108. 1856. Polygonum acre p leptostachyum Meissn. loc. cit. 108. 1856. Polygonum acre y brachystach ,H,>! Meissn. loc. cit. 108. 1856. Polygonum acre S majus Meissn. loc. cit. 108. 1856. Polygonum acre e riparium Meissn. loc. cit. 108. 1856. Polygonum punctatum eciliatum Small, in Bull. Torr. Club 20:214. 1893. Polygonum punctatum robu rr Club 21 -477 1894 Persicaria punctata eciliatu - -9. 1903. Persicaria punctata robustior Small, loc. cit. 379. 1903. Persicaria robustior Bickn. in Bull. Torr. Club 36:455. 1909. Persicaria punctata var. tacubayana Nieuwl. in Am. Midi. Nat. 3:131. 1913. Polygonum robustius Fernald, in Rhodora 25:147. 1921. Polygonum punctatum var. parvum Vict. & Rousseau, in Contr. Inst. Bot. Univ. Montr. 36:13. 1940. Polygonum punctatum var. aquatile (Mart.) Fassett, in Caldasia 4:221. 1946. Polygonum punctatum var. aquatile f. stipitatum F ,ssett, 1. c cit. 223. 1946. Polygonum punctatum var. typicum F., - 1 . 1949. Polygonum punctatum var. tacubayanum (N it. 374. 1949. Polygonum punctatum var. majus (Mrin 72. 1949. Polygonum punctatum var. ellipticum Fassett, loc. cit. 375. 1949. Polygonum punctatum var. con: >. 377. 1949. Polygonum punctatum var. confertiflorum f. longicollum Fassett, loc. cit. 379. 1949. Polygonum punctatum var. littorale Fassett, loc. cit. 379. 1949. Polygonum punctatum var. parviflorum Fassett, loc. cit. 381. 1949. Polygonum punctatum var. mexicanum Fassett, loc. cit. 381. 1949. Polygonum punctatum var. riparium (Meissn.) Fassett, loc. cit. 382. 1949. (310) flora of Panama (Polygonaceaej Slender glabrous annuals or perennials, sometimes creeping and forming col- ies. Leaves punctate with dark glands, subglabrous, entire but minutely ciliate the margins, apically attenuate, the bases decurrent on the petioles; blades 330 ANNALS OF THE MISSOURI BOTANICAL GARDEN lanceolate to narrowly ovate, 2-10 cm. long, 0.5-2 cm. broad; ochreae 0.5-1.5 cm. long, strigose-ciliate only apically. Inflorescences of terminal interrupted, often drooping, spicate racemes or panicles, the peduncles glabrous, the floriferous por- tions 5-15 cm. long, about 0.5 cm. broad, the flowers fascicled in ciliate ochreolae 2-3 mm. long. Flowers mostly white to greenish, the pedicels apically articulated and exceeding the ochreolae by 1-2 mm.; tepals 5, subequal, the outer 3-4 mm. long, the inner somewhat shorter, strongly punctate, ovate; stamens typically 8, discrete, affixed just above the base of the tepals; filaments 1-1.5 mm. long, the anthers less than 0.5 mm. in diameter, subrotund, versatile, thus appearing peltate; ovary trigonous; styles 3, about 1 mm. long, connate for about 0.5 mm.; stigmata capitate. Achenes triquetrous (in Panama) , with ovate facies and inconspicuous beaks, brown to black, 2.5-4 mm. long. canal zone: Gamboa, Bro. Heriberto 37; shore, end of Barro Colorado Island, Bangham 571 ; Joan Mina, Piper 5694; in water, Darien Station, Standley 31578. chiri- > of El Valle de Anton, ca. 600 m., Allen 1986; valle of the Upper Rio < hiriqui Viujo, P. White 3S; vicinity of Boquete, ca. 1300-1700 m., Woodson, Allen 6 Seibert 1165. cocle: lower portion of valley and marshes along R. Anton, El Valle de Anton, ca. 500 m., Hunter & Allen 354; Penonome and vicinity, 50-1000 ft., along stream, Williams 60; El Valle, valley floor and lower slopes along highway, Miller 1834. colon: Ahorca Lagarto to Culebra, Cowell 391. Panama: Chepo, 100 ft., Bro. Maurice 763. province unknown: western Panama, Stark 44; Sutton Hayes 968. This, the most commonly collected and perhaps most abundant species in Panama, ranges from southern Canada to Argentina. Some local Central Amer- ican names are chilillo, chilillo de perro and canilla de pava. Decoctions of the acrid leaves are reported to be used in treating dogs suffering from mange. Those interested in infraspecific delineations are referred to the monograph of Fassett (in Rhodora 6:369. 1949.), in which twelve varieties, four to be expected in Panama, are treated. Fassett cites Panama specimens of vars. aquatile and ellipticum; vars. confer tiflorum and majus, judging from their distributions, are to be expected in Panama. Closely allied P. portoricense Bert., with eciliate ochreae and ochreolae, is widespread in tropical America and might be expected in Panama. 3. Polygonum hydropiperoides Michx. Fl. Bor. 1:239. 1803. Polygonum barbatum Walt. Fl. Car. 131. 1788. not L. Polygonum mite Persoon, Syn. PI. 1:440. 1805. Polygonum virgatum Cham. & Schlect. in Linnaea 3:45. 1828. Polygonum hydropiperoides p virgatum Meissn. in Mart. Fl. Bras. 5*:17. 1855. Polygonum hydropiperoides var. strigosum Small, in Bull. Torr. Club 9:3 5 5. 1892. Polygonum hydropiperoides macouni Small, in Mem. Dept. Bot. Col. Coll. 1:81. 1895. Persicaria hydropiperoides (Michx.) Small, Fl. SE. U. S. 378. 1903. Polygonum hydropiperoides f. leucochranthum Moore, in Rhodora 16:129. 1914. Polygonum hydropiperoides var. digitatum Fernald, in Rhodora 23:260. 1922. Polygonum hydropiperoides f. strigosum Rhodora 28:26. 1926. Polygonum hydropiperoides var. macerum Stanford, loc. cit. 26. 1926. Polygonum hydropiperoides var. sanebelense Stanford, loc. cit. 27. 1926. Polygonum hydropiperoides var. bushianum Stanford, loc. cit. 27. 1926. Polygonum hydropiperoides var. asperifolium Stanford, loc. cit. 27. 1926. Slender subglabrous perennials, occasionally creeping and rooting at the lower (312) flora of Panama (Polygonaceae) 331 nodes. Leaves punctate with inconspicuous pellucid glands, usually strigillose on the veins below, entire but minutely ciliate on the margins, apically long-attenuate, the bases decurrent on the petioles; blades narrowly lanceolate, slightly falcate, 4-15 cm. long, 0.5-1.5 cm. broad; ochreae 1-3 cm. long, with apical cilia up to 8 mm. long. Inflorescences of terminal, lax, often drooping, subspicate racemes or panicles, the peduncles glabrous; floriferous portions interrupted, 5-15 cm. long, about 0.5 cm. broad, the flowers fasciculate in ciliate ochreolae 2-3 mm. long. Flowers mauve to greenish, the pedicels apically articulated and ultimately exceed- ing the ochreolae by about 1 mm.; tepals usually 5, subequal, ovate, becoming 2-3 mm. long; stamens usually 8, discrete, affixed just above the base of the tepals; filaments about 1 mm. long; anthers less than 0.5 mm. in diameter, subrotund, versatile, thus appearing peltate; ovary trigonous; styles 3, about 1 mm. long, connate for about 0.5 mm.; stigmata capitate. Achenes triquetrous (in Panama), with ovate facies and inconspicuous beaks, brown to black, 2-3 mm. long. canal zone: Gigante Bay, Barro Colorado Island, Shattuck 840; Barro Colorado Island, Kenoyer 347. Usually inhabiting meadows and stream banks, this species ranges from Canada to Central and South America, where it is supposed to merge with P. persicarioides HBK. In regions of Guatemala it is called flor de chajutal (fide Standley & Steyermark, in Field Mus. Bot. 24 4 :126. 1946.). Intergradation, south of the continental borders of the United States, prompted Stanford (in Rhodora 28:25. 1926.) to reduce P. persicarioides HBK. to varietal status under P. hydropiperoides. In the Central American material however, P. persicarioides seems more distinct from P. hydropiperoides than does P. punctatum. Recent workers have probably rightly upheld P. persicarioides as a distinct species. It occurs at moderate eleva- tions in Guatemala, Costa Rica and South America and is perhaps to be expected 4. Polygonum ACUMINATUM HBK. Nov. Gen .&Sp.2 :178. 1817 Polygonum cuspidatum Willd. in Spreng. Syst. 2:256. 1825. Polygonum erectm m Veil. Fl. Flum. 4: t. 42. 18 Polygonum setiger urn Wedd. in Ann. Sci. Nat. 3 Polygonum floribu ndum Wedd. loc. cit. 253. U Polygonum acumh tatum a bumbol DC. Prodr. U 18 56. Polygonum acumh tatum /3 glabrescem Meissn. lo Polygonum acumh ratum y subcordatun; \tcissn. Polygonum acumh Polygonum acumh latum e brachystemon Meissn. :it. 114. plhgZZZ IZZh ■ temon Meissn. 1 latum v wcdd 14. 1856 Polygonum acumh latum B capense Meissn. loc. ci k 1856. ta (HBK.) Maza, in Per. Cub. 1896. Polygonum guatemalense Gandoger, in Bull. Bot. Soc. France 66:225. 1919. Rather robust perennials, often conspicuously strigose or cinereous. Leaves punctate with inconspicuous pellucid glands, usually strigose or cinereous, strigose on the margins, apically long-attenuate, the bases abruptly decurrent on the 332 ANNALS OF THE MISSOURI BOTANICAL GARDEN petioles; blades sessile or subsessile, lanceolate, falcate, 10-30 cm. long, 2-3.5 cm. broad; ochreae 2-3 cm. long, with apical strigose cilia up to 1.5 cm. long, strigose on the veins. Inflorescences of terminal rather rigid subspicate racemes or panicles, the peduncles strigose; floriferous portions continuous, 3-13 cm. long, almost 1 cm. broad, the flowers fasciculate in ciliate imbricate ochreolae 2-3.5 mm. long. Flowers whitish, the pedicels apically articulated and ultimately exceeding the ochreolae by 1-2 mm.; tepals 4 (-5), subequal, ovate, becoming 2.5-3.5 mm. long; stamens commonly 6, discrete, affixed just above the base of the tepals; filaments to 3 mm. long, with versatile anthers about 0.5 mm. long; ovary lenticular; styles 2, 2-3 mm. long, connate for about 0.5 mm.; stigmata capitate. Achenes lenticular, with ovate facies and a clearly delineated beak about 0.5 mm. long, brown to black, lustrous, 2-2.5 mm. long, 1.5-2 mm. broad. White l 168. Inhabiting marshes and streams of low elevations, this species ranges from Mexico through Central America and South America to Argentina. In Guate- mala it is called chilillo and chilillo de clucho. The typical varieties of P. punc- tatum and P. hydropiperoides, essentially North American varieties, grade into atypical varieties in the Caribbean countries, and here tend to approach the more southern speciees, P. acuminatum and P. persicarioides. This gradation is responsible for a serial relationship between these four species. Data suggesting this seriation are included in the following chart: XZSL P. hydropiperoides var. hydropiperoides P. persicarioides P. .eumm.tum with dark "pun'cS with pellucid with pellucid "petrous 17 iri. 1840. Antigonon platypus Hook. & Arn. loc. Antigonon cordatum Mart. & Gal. in Bi Antigonon cinerascens Mart. & Gal. loc, Corculum leptopus (Hook. & Arn.) 5 U. S. D. . ufous-pubescent tuberous vir Glabrous to densely cinereous- c broadly cordate to narrowly deltoid, mostly 3-14 cm. long, 2-12 cm. broad, glabrate to closely pubescent below; mostly 1-2.5 cm. long, glabrous to pubescent, the leaf bases occasionally de< 336 ANNALS OF THE MISSOURI BOTANICAL GARDEN on the petioles forming flanges up to 5 mm. wide. Inflorescences of axillary and terminal capreolate racemes or panicles, the rhachises glabrate to densely pubescent. Flowers pedicellate, the glabrous to densely pubescent pedicels mostly articulated below the middle; tepals pink to purplish, cordate, in fruit 8—25 mm. long, 4-20 mm. broad, glabrous to closely pubescent without, rarely stipitate-glandular within; stamens 8, connate about half their length to form a filament tube, often with gonous; styles 3, arcuate; stigmata peltate. Achenes bluntly triquetrous, usually included, about 1 cm. long. canal zone: Balboa, Standby 25533; Balboa, Standley 28551 ; Chiva-chiva trail 2 mi. above Red Tank, Maxon 6 Harvey 6612; Matachin to Las C i«cadas, Cowell J57- Panama: O. U. Experiment Station, 3 mi. e. of Panama City, Maxon, Harvey & Valentine 7088. y This species is widely cultivated and escaped and determination of its natural range would be difficult. It is cultivated in Florida where it is called "confederate vine". In the West Indies it is called corallina and corallillo. In Mexico, where it is supposed to have originated, it is called coronilla and chaclomacal. Other Central American names are confite, ftor de san miguel, san diego, bellisima, eolation and confite rojo. In the Canal Zone it is called "coralvine", coralito, enredadera and cadena de amor. In Panama and the Canal Zone, A. leptopus seems to be fairly well established. A variant commonly called A. cinerascens/' which seems to differ regularly only in its dilated petioles, might also be cultivated here. Comparison of specimens with and without the dilated petioles has revealed no correlated differences. Hence it is concluded that A. platypus (A. cinerascens) is conspecific with A. leptopus. 2. Antigonon flavescens Watson, in Proc. Am. Acad. 22:446. 1887. Finely pubescent tuberous vines. Leaves narrowly cordate to deltoid, acute, e, the blades mostly about 8 cm. long by 6 cm. broad, pubescent < veins below; petioles 1-2 cm. long, sparsely pubescent, the leaf bases decurrent on the petioles forming flanges up to 4 mm. broad. Inflorescences of axillary and terminal capreolate racemes or panicles, the rhachises rufous-pubescent. Flowers pedicellate, the minutely pubescent pedicels articulated 2-3 mm. from the rhachis, 7-10 mm. long; tepals greenish white, cordate, acute, in fruit about 10 mm. long by 4 mm. broad, glabrate; stamens 8, connate slightly less than half their length, about 2 mm. long; ovary trigonous; styles 3, arcuate; stigmata peltate. Achenes bluntly triquetrous, about 11 mm. long, slightly exserted. platypus Hook. flora of Panama (Polygonaceae) 337 This lochmophilous species is known only from Mexico and Guatemala. Due to the inferior color of its tepals, it is less likely to be cultivated than other species. In characters other than color this species is very similar to some of the narrow- tepaled forms of A. leptopus. Although the flowers of a specimen labeled A. flaves- cens, Pringle 4570, collected in Jalisco, are annotated by C. H. Thompson (Oct. 10, 1898) as "all right for A. flavescens", the fruiting tepals are broadly ovate. If this annotation is correct, there is little doubt that A. flavescens is but a color form of A. leptopus. Watson (in Proc. Am. Acad. 22:446. 1887.) offered no support- ing character for color save the narrower tepals. I strongly suspect that A. flavescens is merely a color form of A. leptopus. On the contrary, Standley & Steyermark (in Field Mus. Bot. 24 4 :106. 1946.) believe it to be a good species. 3. Antigonon guatemalense Meissn. in DC. Prodr. 14:184. 1856. Polygonum grandiflorum Bertol. Fl. Guat. 412. 1840. not Willd. 1799. Antigonon grandiflorum (Bertol.) Robinson, in Proc. Am. Acad. 44:613. 1909. Antigonon macrocarpum Britton & Small, in Britton & "Wilson, in N. Y. Acad Sci. 5 2 :266. Cinereous tuberous vines. Leaves broadly cordate, abruptly acuminate, mucro- nate, the blades 3-9 cm. long, 2.5-7 cm. broad, closely cinereous-pubescent below; petioles mostly less than 1 cm. long, cinereous-pubescent, terete, the leaf bases not decurrent. Inflorescences of axillary and terminal capreolate racemes or panicles, the rhachises cinereous-pubescent. Flowers pedicellate, the pedicels medially articulated, with a change in degree of pubescence at the point of articulation, becoming 10-15 mm. long; tepals pink, at anthesis as broad as long, about 1 cm. long, closely cinereous-pubescent without, stipitate-glandular within; stamens 8, connate about one fourth their length, about 2 mm. long; ovary trigonous; styles 3, arcuate; stigmata peltate. Mature achenes not seen. This species, an inhabitant of thickets, is often cultivated. No specimens have been seen from Panama. It has been collected in Mexico, Guatemala, Salvador, Nicaragua, Costa Rica and the West Indies. Colloquial names reported to be applied in Salvador are colacion, confite and san andres. 4. MUEHLENBECKIA Meissn. Muehlenbeckia Meissn. Gen. PL 2:227. 1840. nom. conserv. Calacinum Raf. Fl. Tellur. 2:33. 1836. Karkinetron Raf. Fl. Tellur. 5:11. 1836. Sarcogonum G. Don. in Sweet, Hort. Brit. 3:577. 1839. Conobaea Bert, in Steud. Nom. Bot. 2 1 :404. 1840. Flowers polygamo-subdioecious (perfect in some spp. outside Panama), with short pedicels articulated at the base of the tepals, 1-5 (-many) in fascicles sub- tended by telescoped ochreolae. Tepals 5 (6, 4), subequal, greenish white, slightly accrescent and becoming more or less fused with the achene basally. Stamens 8 (-10), 1-3 mm. long, inserted on the base of the tepals, discrete or forming an inconspicuous annular disc, reduced or absent in pistillate flowers; anthers 4-locular, 338 ANNALS OF THE MISSOURI BOTANICAL GARDEN introrse, versatile. Ovary trigonous, reduced or absent in staminate flowers; styles 3, short, arcuate; stigmata capitate, lobate, or fimbrillate. Achenes mostly included, bluntly to acutely triquetrous, dark, lustrous, often puncticulate. Scramblers or low matted shrubs with usually glabrous, often sulcate stems. Leaves alternate, entire, apically acuminate to acute, basally hastate, cordate or acute, the blades usually exceeding the petioles; ochreae hyaline and conspicuous, deciduous. Inflorescences of axillary panicles, racemes or glomerules, occasionally This genus of some twenty species occurs chiefly in tropical American and Australian areas, often in alpine regions. In Central America are two indigens, M. tamnifolia and M. volcanica. Only M. tamnifolia has been collected in Panama. M. volcanica, a low mat-forming alpine shrub with the flowers solitary or fascicu- late in the axils of the small elliptic leaves, inhabits the high mountains of Guate- mala and western South America from Bolivia to Ecuador. M. platyclada, with the stem conspicuously flattened, is often cultivated in the Americas as a bizarre of the Solomon Islands. (HBK.) Meissn. Gen. PL 2:227. 1840. Polygonum tamnifolium HBK. Nov. Gen. & Sp. 2:180. 1817. Polygonum flexuosum Benth. Pi. Hartw. 80. 1839. Polygonum quadrangulatum Mart. & Gal. in Bull. Acad. Brux. 10 1 :353. 1843. ckie Benthamii Endl. Gen. Suppl. 4:51. 1847. Muehlenbeckia quadrangulata Endl. loc. cit. 51. 1847. Muehlenbeckia tamnifolia a Humboldtii Meissn. in DC. Prodr. 14. 149. 1856. Muehlenbeckia tamnifolia /3 quadrangulata Meissn. loc. cit. 149. 1856. Muehlenbeckia tamnifolia y Hartwegii Meissn. loc. cit. 149. 1856. Muehlenbeckia tamnifolia 5 laxiflora Meissn. loc. cit. 149. 1856. Muehlenbeckia leptobotrys Meissn. loc. cit. 149. 1856. Muehlenbeckia eron. in Bot. Jahrb. 21:307. 1895. Sarcogonum tamnifolium Rusby, in Mem. Torr. Club 6:111. 1896. Calacinum tamnifolium (HBK.) Macbr. in Field Mus. Bot. 4:116. 1927. Calacinum leptobotrys (Meissn.) Macbr. loc. cit. 116. 1927. Clambering glabrous lianas, the branches smooth and sulcate, often angulate. Leaves glabrous, ovate, apically usually abruptly acuminate, basally hastate or cordate to acute, the blades mostly 3-8 cm. long, 1.5-4 cm. broad; petioles mostly 8-20 mm. long, glabrous, terete or canaliculate. Inflorescences chiefly of axillary racemes or panicles, the rhachises glabrous, rarely with reduced leaves. Flowers short-pedicellate, the pedicels apically articulated, 0.5-3 mm. long; tepals mostly five, greenish white, ovate, apically subacute to rounded, slightly connate basally, 1.5-3 mm. long, 1-2 mm. broad, glabrous; staminate flowers with 8 (-10) mostly separate stamens about 1 mm. long attached to the lower half of the tepals, the anthers introrse and versatile, about 0.5 mm. long, the ovary a short 3 -styled rudiment or absent; pistillate flowers with trigonous superior ovaries with ovate facies; styles 3, strongly arcuate; stigmata nmbrillate-flabellate; stamens reduced to subsessile staminodia. Achenes bluntly trigonous, usually included by the tepals and hardly separable from their bases, often capped by an amorphous mass of stylar (320) flora of Panama (Polygonaceae) long, nearly or quite iriqui: R. Chiriqui Viejo, near Monte Lino, G. White OJ; Bajo Chorro, Bo< lai-hlwH 177; vicinity of Casita Aha, V..: <>a, Ml.,, j S, -ail from Paso Ancho to Monte Lirio, Allen 1507; Bajo Mono-Robalo trail, we F Cerro Horqueta, Allen 48 1 2. province unknown: Sutton Hayes 486, g8j. 1 America to Argenti 1 of the leaves was on Often forming tangles in thickets or forests, this species ranges from southern Mexico through Central America and western Sov Costa Rica it is called bejuco Colorado. A decocti sidered anti-hemorrhagic (fide HBK. Nov. Gen. & Sp. 2:180. 1817.). It is perhaps just coincidental that most specimens from Costa Rica and Panama have rounded to acute leaf bases and in other respects match the description of M. stuebelii Lind., a Colombian variant differing chiefly by its attenuate leaf bases and smaller inflorescences exceeded by the leaves. Allen 4812 with both acute and subcordate leaf bases clearly shows that the leaf base is of little taxonomic sig- nificance in M. tamnifolia. Since many specimens with acute leaf bases have inflorescences clearly exceeding the leaves, it is concluded that M. thtfbeUi is conspecific with M. tamnifolia. 340 ANNALS OF THE MISSOURI BOTANICAL 5. COCCOLOBA P. Br. ex L. By RICHARD A. HOWARD Coccoloba P. Br. ex L. Syst. Nat. ed. 10. 1007, 1367. 1759. Taxon 3:114, 156, 233. 1954, nom. conserv. Howard, Jour. Arnold Arb. 40:176-220. 1959. Guaiabara Miller, Gard. Diet. ed. 4. 2. 1754. Coccolobis P. Br. Civ. & Nat. Hist. Jam. 209. pi. 14, f. 3. 1756. Shrubs or trees, trees with scrambling branches, or lianas; branches terete or stout, commonly strongly striate, short shoots commonly developed laterally, or the terminal shoots of limited growth becoming long shoots, pith solid or the stem hollow except at the nodes; nodes commonly swollen; ocreae characteristically developed, sheathing at first, becoming split along one or two sides, the basal portion commonly coriaceous and persistent, the apical portion commonly mem- branaceous and deciduous or entirely membranaceous and deciduous, glabrous, puberulent or silky pubescent; leaves alternate, persistent or deciduous, membrana- ceous, chartaceous or coriaceous, on deciduous trees the young leaves usually turning black on drying, varying considerably in size on the same shoots, the leaves of adventitious or juvenile shoots commonly much larger and frequently of dif- ferent shape from those of normal shoots; the petiole borne at the base of, or well above the base of, the ocrea, petioles slender to stout, commonly canaliculate above, glabrous to puberulent or pubescent; leaf blades with margins flat to slightly recurved, straight or undulate, midrib commonly keeled above and often below, primary veins straight to the margin or branched near the margin becoming reticulate, or arcuate and anastomosing, or arcuate and bifurcate-anastomosing, the secondary venation parallel or reticulate, conspicuous to obscure, leaf surface glabrous, puberulent or pilose becoming glabrate, pubescence often mixed with glandular-like bodies which are either hair bases or resinous excretions from the stomata or rarely multicellular glands; inflorescence terminal or terminal on lateral shoots, paniculate, racemose or spicate, the peduncle usually short; basal ocrea present, flowers functionally unisexual, the staminate flowers borne in clusters of 2-7, the pistillate flowers usually borne singly at each nodule on the rachis, rachis swollen at each flower cluster or terete, flowering clusters distinct or confluent, subtended by a small bract and one or more membranaceous ocreolae, ocreolae usually covering the flower bud and splitting regularly or irregularly forming a membranaceous collar or sheath, or one, two or more appendages, pedicels short or well developed, equaling or exceeding the ocreolae, hypanthium well developed or slight, perianth lobes usually 5, rarely to 7 in number, imbricate in bud, often reflexed at maturity, stamens usually 8, functional stamens exserted, non-functional stamens included, filaments commonly flared at the base and more or less united, functional pistil exserted, the non-functional pistil included, ovary strongly trigonous, glabrous, functional styles 3, or if 2 or 1 the non-functional styles filamentous, stigmatic surface often expanded; hypanthium and perianth lobes expanding in fruit, or only the perianth lobes expanding to cover the achene, fruiting hypanthium when fresh commonly brightly colored and fleshy, usually (322) flora of Panama (Polygonaceae) 341 astringent to taste, occasionally strongly fibrous; the achene trigonous in outline, black, brown or tan in color, shiny or dull, the outer wall hard, the inner layer papery, seed with ruminate endosperm, the major lobes 3, the minor lobes and involutions numerous, the embryo centrally located, the cotyledons orbicular, flat, rarely folded or contorted, the radicle small and terete. Type: Coccoloba uvifera (L.) L. (Polygonum uvifera L.), probably from closely associated and not I 342 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. Coccoloba lasseri Lundell, Contr. Univ. Mich. Herb. 6:10. 1941. Tree to 12 m. tall; branchlets thick, striate, puberulent; ocreae cylindrical and flaring at the top, to 1.5 cm. long, subtruncate, membranaceous above the petiole, densely puberulent when young, the petiole inserted at or above the middle; petioles minutely puberulent, stout, striate, 3-8 mm. long; blades obovate or oblanceolate- obovate, 12 X 5, 15X8, 19 X 10.5 cm. long and broad on fertile branches, chartaceous, glabrous above, puberulent below on the midrib and primary veins, apex rounded to obtusely- acute, attenuate from above the middle to the base, the base emarginate or rounded, primary veins 8-12 on each side; adventitious shoots and their leaves unknown; inflorescences terminating short lateral branches, rarely terminal, paniculate, the panicles sessile, composed of 3 or 4 racemes, the racemes to 30 cm. long, peduncles of racemes to 1.5 cm. long, the rachis densely puberulent striate; staminate flowers borne in clusters of 3-6, pistillate flowers borne singly bracts and ocreolae minute, less than 0.5 mm. long, the ocreolae multiple, branaceous, pedicels 1-1.3 mm. long, puberulent, hypanthium about 1 mm. contracted into a short stipe, perianth lobes suborbicular, about 1 mm. in diar fertile stamens slightly exceeding the perianth lobes, functional pistil 2-3 mm. long, the styles 2 or 3, fruiting pedicels 1-1.3 mm. long, the mature fruit nearly spherical, the perianth lobes imbricate, about ^ the length of the fruit; the achene light brown, not coronate at the apex. Common name: "uvero". Collected in flower in May, June, September and November. Collected in fruit in August and December. Recorded altitude, the tidal belt to 20 m. Ebinger q8q. s.n. Panama: Bejuco, Allen 2542- Panama Nat. Hieh .. „ . , . Latter 16638; Rfo Pacora, Bartlett & Lasser 16046 (Holotvpe, MICH.) ; Balboa to Chame, Bodge, Hunter, Steyermark tf Allen 167321 Punta \&$. province unknown: Duchassamg s. n. (collected 1851). Here regarded as endemic to Panama. Coccoloba lasseri is very similar to C. tuerckheimii, differing in its smaller leaves, smaller inflorescence and nearly spherical fruits. A study of young plants, vigorous shoot growth or adventitious growth may well prove this species to be the same as C. tuerckheimii. ':213. 1904. Large tree to 15 m. tall; branches stout, striate, the internodes frequently shortened and compacted, forming areas of short shoot growth, lateral shoots, par- ticularly flowering branches, distinct as short shoots; ocreae to 3 cm. long, large and conspicuous, coriaceous, strongly striate and persistent at the base, membrana- ceous and deciduous at the apex, puberulent becoming glabra te; petioles borne 6-15 mm. above the base of the ocreae, 1.5-3.5 cm. long, usually striate and slender, puberulent when young; blades oblong-ovate, abruptly short 1 at the apex, the base acute and decurrent on the petiole, 13X7, 25 X 1 45 X 25 cm. long and wide, thinly coriaceous, glabrous above, puberulent 1 Coccoloba tuerckheimii Donnell Smith, Bot. Gaz. flora of Panama (Polygonaceae) 343 344 ANNALS OF THE MISSOURI BOTANICAL GARDEN when young, remaining so on the midrib and veins, primary veins 7-10 on each side, occasionally nearly barbate in the axils of the midrib and the veins on each side; inflorescence generally terminal on lateral short shoots, paniculate, the ocreae coriaceous, flaring, striate, 1 cm. long, the panicle almost sessile, peduncle short, 1-3 cm. long, racemes numerous, 10-19, each 10-35 cm. long, rachis and all parts puberulent, staminate flowers in clusters of 4-7, pistillate flowers borne singly at each nodule, bracts small, broadly triangular, less than 0.5 mm. long, ocreolae shorter than to scarcely exceeding the bracts, pedicels 2-3 mm. long, hypanthium conical 0.75 mm. long, perianth lobes oblong, 1-1.5 mm. long, 1 mm. wide, fertile stamens on filaments 3 mm. long, fertile pistil 3 mm. long; fruiting pedicels 2-3 mm. long, fruit oval, 11-14 mm. long, 6-9 mm. in diameter, the apex obtuse and slightly coronate, the perianth lobes surrounding the apex of the achene, the base of the fruit constricted to a stalk 1—2 mm. long; achene dark brown, obtusely trigonous in outline. Type: von Tuerckheim 8493, Cubilqiiitz, Dept. Alta Verapaz, Guatemala. Collected in flower in August. Collected in fruit in July. Guatemala, Honduras, Nicaragua, Costa Rica, Panama. alley, Island of Potrero, Dunlap 181. colon: Dos 3. Coccoloba coronata Jacquin, Enum. Pi. Carib. 19. 1760, Select, Stirp. Am. Hist. 114, /. 77. 1763; Dugand, Caldasia 4:427. 1947; Howard, Jour. Arnold Arb. 41:40, 226, 227. 1960. Coccoloba novogranatensis Lindau, Engl. Bot. Jahrb. 13:192. 1890. Coccoloba caribaea Urban, Symb. Ant. 5:377. 1907. Coccoloba waitii Johnston, Sargentia 8:122. 1949. Shrub, 3 m., or tree, 15m.; trunk commonly muscular, bark red; branchlets terete, glabrous, gray, the nodes slightly swollen; ocreae cylindrical, the upper portion membranaceous and deciduous, the lower portion coriaceous and persistent; leaves of normal shoots with petioles 8-11 mm. long, puberulent when young, with age glabrous except in the groove, attached above the base of the ocreae; blades ovate to oblong-ovate to ovate-elliptic, 6 X 4, 9 X 4, 10.5 X 5.5 cm. long and broad, thin- coriaceous, glabrous above, glabrate below except for long hairs along the midrib at the base of the blade, the apex attenuate, the base narrowly cordate, the margin entire, midrib and primary veins inconspicuous above, promi- nent below, the primary veins 5 or 6 pairs, the ultimate venation finely reticulate; leaves of adventitious shoots generally similar but occasionally obovate and nar- rowed at the base to ovate-lanceolate or lanceolate-elliptic, the blades 11 X 7 to 18 X 15 cm. long and broad, the basal lobes of leaves of adventitious shoots often overlapping; inflorescence terminal on short lateral shoots, 3-9 cm. long, the basal ocreae 5-10 mm. long, bilobed at the apex, the rachis glabrous, flowers commonly borne singly at each node, but occasionally two together in staminate plants, the bracts oblong, 2 mm. long, 0.5-0.75 mm. broad, the ocreolae 3 mm. long, bilobed at the apex, membranaceous and persistent, the pedicels 2.5-4 mm. long, the hypanthium 0.75 mm. long, the perianth lobes ovate, 1.5 mm. long and broad, the 026) flora of Panama (Polygonaceae) 345 functional stamens 2.5 mm. long, the functional pistil 2 mm. long; fruit generally spherical with coronate lobes, occasionally narrowed at the base, 1 cm. long, 5-9 mm. in diameter; the achene brown, shining. Collected in flower in May. Collected in fruit in July, September and November. St. Vincent, Grenada, Trinidad, Tobago, Margarita, Venezuela, Colombia, Peru, Guatemala, Panama. darien: Cana, Williams Q45. herrera: Pese, Allen 802. Panama: Perlas Islands, Pedro Gonzalez, , Islands, Erlanson tfj, 550, 2l6; Harlow 41 ; John- ston 21 3, 230 (GH-type of C. waitii) 957, 1306. Although most of the specimens cited have been collected in coastal areas or at low elevations, the specimens by R. S. Williams 945 were collected at 2800 ft. altitude. This collection is young material but appears to be safely referred here. There appears to be no specimen preserved that was collected by Jacquin. However, the description and illustration are sufficiently specific and can serve as the nomenclatural type. 4. Coccoloba obovata HBK. Nov. Gen. 2:176. 1817. Coccoloba Goudotiana Wedd. Ann. Sci. Nat. Ill, 13:260. 1850. Coccoloba riparia Lundell, Contr. Univ. Mich. Herb. 6:11. 1941. Shrub of 3 m. to tree 15 m. tall; branches terete, glabrous; ocreae oblique, truncate, 7—10 mm. long, glabrous, membranaceous, striate, appressed; leaves of normal shoots with petioles 12-15 mm. long, pilose or glabrous, inserted above the base of the ocreae, blades obovate to subround-obovate, rarely ovate-lanceolate to elliptic obovate, 9X6, 10X4, 16X8 to 24 X 9.5 cm. long and wide, apex obtusely acuminate, base slightly cordate, coriaceous, turning dark brown or black on drying, tomentulose on the midrib and veins, persistent, short pilose in the axils of the veins and midrib, commonly densely covered with globose or peltate excretions associated with the stomata; leaves of adventitious shoots with petioles 3.5-6 cm. long, blades broadly elliptic-ovate to elliptic-oblong, 28 X H to 37 X 26 cm. long and wide; inflorescence terminal, solitary, densely flowered, staminate to 25 cm. long, pistillate to 17 cm. long, the flowers borne singly in the pistillate and in clusters of 4 in the staminate, peduncle, rachis, bracts and ocreolae short pilose, bracts semi-orbicular, 0.75 mm. long, ocreolae bilobed 1.5 mm. long, mem- branaceous, usually in multiples, pedicels included in the ocreolae in flower, exserted and to 1.5 mm. long in fruit, hypanthium 0.75 mm. long, the lobes ovate or nearly orbicular, to 1 mm. long, functional stamens exserted, sterile stamens included, ovary triangular, styles 3; fruit globose, obtusely-trigonous, strongly striate, the base rounded, the apex obtusely coronate, 6 mm. long, 5.5 mm. in diameter; achene tan to dark brown, trigonous. Type: Colombia, prov. Tolima, Honda, Humboldt s. n. June 1805 (Herb. Willd. 7704), isotype Paris. Common name: "Papaturra blanca", "Pena blanca". Collected in flower in March, June, July, September and December. Collected in fruit in August and December. 027) OF THE MISSOURI BOTANICAL GARDEN Costa Rica, Panama, Colombia. canal zone: Barro Colorado Island, Aviles 998, Salvoza 984, Shattuck 1063, 298; F. L. Island, Bangham 593; Salamanca Hydrographic Station, Dodge, Steyerr, Allen 16983, 16983a, Woodson, Allen & Seibert 1572; northwestern part of Cana Johnston 1519; between Tumba Vieja and Salamanca, Steyermark ti Allen 16758. qui: Cerro Galera Chorcha, Gualaca, Allen j020; San Felix, Allen 3655; betwe Felix and Cerro Flor, Allen 1 91 7; Progreso, Cooper (3 Slater 270 ; San Bertolome, W & Schery 947; between Rio Chiriqui and Remedios, Woodson, Allen 2? Seibert cocle: El Valle, Allen 2229 (MICH-holotype of C. riparia) . veraguas: b Santiago and David, Stern, Chambers, Dwyer and Ebinger 999. 5. Coccoloba parimensis Bentham, Hook. Lond. Jour. Bot. 4:626. 1845. Tree to 8 m. with scrambling branches or a liana reaching 44 m.; stems slender, striate, minutely puberulent becoming glabrous, sparsely glandular; ocreae mem- branaceous, split and flaring at the apex, 2-2.5 cm. long, puberulent, usually decidous to the petiole; petioles arising from the base of the ocrea, slender, 1.5-2 cm. long on fertile shoots, puberulent; blades oblong-elliptic to elliptic-ovate, 8.5 X4.5, 13. 5X7 to 21X15 cm. long and broad, membranaceous when young becoming thin-coriaceous when mature, flat to noticeably bullate between the veins, puberulent to short pilose on the midrib and veins, puberulent to glabrate, usually sparsely glandular between the veins, apex short acuminate to obtuse, the base rounded, primary veins 6—9 on each side; leaves of sterile shoots larger, on petioles 5-6 cm. long, the blades to 27 X 16 cm. long and broad, the apex rounded and apiculate, the base truncate to cordate; inflorescence terminal on short lateral branches, usually solitary, all parts puberulent, staminate inflorescence 15 cm. long, the flowers 2-7 at each nodule, the pistillate inflorescences rarely exceeding 10 cm. in length, the flowers borne singly at each nodule, bracts minute, 0.5 mm., ovate, acute, puberulent, ocreolae large, 3-4 mm. long, membranaceous, bilobed, each lobe apiculate to acute, pedicels equaling or slightly exceeding the ocreolae, hypanthium 0.8 mm. long, narrowed to a stipe 0.5-1 mm. at the base, perianth lobes oblong-elliptic, 2 mm. long, fertile stamens longer than the lobes and exserted, fertile pistil 2 mm. long, styles 3, non-functional parts rudimentary; fruiting pedicels 3-4 mm. long, the fruit ovate, 1 cm. long, 8 mm. in diameter, the base rounded, short stalk evident, the apex obtusely acute, not coronate, the lobes imbricate and about % the length of the fruit, smooth 2nd not striate; achene chestnut brown, smooth, not trigonous. Type: Schomburgk 5. n. Rio Parime, Brazil (Kew) . Collected in flower in June. Collected in fruit in June and August. Panama, Colombia, Peru, Brazil. Altitude to 120 m. canal zone: Barro Colorado Island, Aviles 15; L. H. & E. 2. Bailey 219, 654; Bangham 447; Salvoza 904; Shattuck 1 123; Standley 41099; Wetmore & Woodworth 859. Panama: Juan Diaz region near Tapia river, Maxon 2? Harvey 6700. flora of Panama (Polygonaceae) 347 Coccoloba uvifera (L.) L. Syst. Nat. ed. 10. 1007. 1759. 1753. lidl. Nat. 8:64. 1922. Tree of strand areas, 2-15 m. tall; branches stout, papillose to pilose; ocreae rigid, coriaceous at the base, membranaceous at the apex, 3-8 mm. long, papillose to pilose; leaves of normal shoots with stout petioles, 7-10 mm. long, puberulent to pilose, the blades orbicular to reniform, 10X10, 11X1 4, 13X18, 20X 27 cm. long and broad, thick and fleshy when fresh, coriaceous when dry, glabrous and minutely punctate on both surfaces, the apex rounded, truncate or emarginate, the base rounded to broadly cordate, one lobe extending around the petiole, the primary veins 3-5 pairs usually straight, bifurcate and weakly anastomosing near the margin, commonly barbate in the axils of the basal veins, the secondary venation minutely reticulate or obscure; inflorescence stout, 15-30 cm. long, racemose, occasionally branched at the base, the rachis puberulent, staminate flowers in clusters of 1-7, the pistillate flowers solitary at each locus, the bracts ovate, 1-1.5 mm. long, 2 mm. broad, puberulent, the ocreolae membranaceous, 1 mm. long, puberulent, the flowering pedicels 1-2 mm. long, the hypanthium 2-3 mm. long, the perianth lobes 4 mm. long, 3-4 mm. wide, the fertile stamens to 4 mm. long; fruiting pedicels 3-4 mm. long, fruit obpyriform, 1.2-2 cm. long, 8-10 mm. in diameter, narrowed at the base, rounded-truncate at the apex, the perianth rose-purple when mature; achene black. Type: Linnaean Herbarium— ( terminal leaf of young plant.) Jamaica (?) Local name: "sea grape". Collected in flower in January, February and October. Collected in fruit in March. General in the West Indies, Mexico, Central and northern South America along beaches and strand locations. bocas del toro: Almirante, G. Proctor Cooper J5l>/>o,j, U.O; w 3 ,■„„■< opIuLt, 161; myrtifolia, 156; numphadfolia, 162; / r\th,ohalanm, 95; humboldtii, 101 pad i folia, \ 5< Erythrodes, 34 Esmeralda, 47 pJdumaU!, plocefnlia, 1 Eulophidiinae, 40 Eulophidium, 41 Eulophiella, 41 Eulophiinae, 40 rWlffc, 150; | Eurycentrum, 34 158; »W«/«j, Eurychone, 47 Eurystyles, 3 5 F ««<*, 159; sk Fagaceae of Panama, 95 150; ™£/r//>/i *«/<■//>«, 156; F?yanl?n /Oft, 158; * Ficus, 146; subgenus Pharmacosycea, 147, 148; subgenus Urostigma, 146, 148; ad- hat odae folia, 152; amazonica, 158; amer- icana, 154; anguina, 162; angustifolia, mat >nn, 159; arboricida,' 155; 'arbuti- folia, 156; arpazusa, 156; arukensis, 161; baccata, 156; better oi, 160; bonplandi- §llS : ana, 163; bopiana, 150; botryapioides, 158; boyacensis, 152; brevifolia, 158; brittonii, 159; bullenei, 158; campbellii, 160; catesbaei, 158; cerasifolia, 156; cestri folia, 156; chiriquiana, 154; ri/io- /om, 156; citrifolia, 158; citrifolia, 149; Calavtndmdrum Gale, 88 ricana, 163, /&£; coybana, 150; crassa, Galeandra, 41 Galeola, 49 Galeorchis, 36 Jroc/o«M, 'l55;V«W«, 159; , ricanum, 163; ei ostictum, 156; ■ ' *«, 154; hartwegii, 1 w, 163; liebman- ensis, 161; «£«*«», 1 56; schiedeanum, I56;sulcipe s, 156; *»rfo Inatum, 156 tf, 191; baccifera, 192; cara- data, 193; , 186; laciniata, 191; numt 185; parietaria, ana, 109; j/r*gitf< Triceratorhynchus, Trichoceros, 46 Trichoglottis, 48 Trichopilia, 46 Vanillinae, 49 Vargasiella, 44 Vargasiellinae, 44 Velasquezia,153;mel, Viscum cordifolium, latifolium, 284; .o.vorcA: c\j; a