ANNALS OF THE MISSOURI BOTANICAL GARDEN VOLUME 74 1987 The Annars, published quarterly, contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden, St. Louis. Papers origi- nating outside the Garden will also be accepted. Authors should write the Editor for information concerning arrangements for publishing in the ANNALS. EDITORIAL COMMITTEE GroncE K. Rocers, Editor Missouri Botanical Garden MarsHALL R. CROSBY Missouri Botanical Garden GERRIT DAVIDSE Missouri Botanical Garden Jonn D. DWYER Missouri Botanical Garden & St. Louis University PETER GOLDBLATT Missouri Botanical Garden HENK VAN DER WERFF Missouri Botanical Garden Colophon This volume of the Annars of the Missouri Botanical Garden has been set in APS Times Roman. The text is set in 9 point type while the figure legends and literature cited sections are set in 8 point type. The volume has been printed on 70# Centura Gloss, an acid-free paper designed to have a shelf-life of over 100 years. Centura Gloss is manufactured by the Consolidated Paper Company. Photographs used in the Annars are reproduced using 300 line screen halftones. The binding used in the production of the Annars is a proprietary method known as Permanent Binding. The Annars is printed and distributed by Allen Press, Inc. of Lawrence, Kansas 66044, U.S.A. © Missouri Botanical Garden 1988 ISSN 0026-6493 Volume 74 E D 2 Ë: = e t Volume 74, Number 4 | Annals of the Spring 1987 Missouri Botanical Garden The Annals, published quarterly, contains papers, primarily in systematic botany, con- tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the Garden will also be accepted. Authors should write the Editor for information concerning arrangements for publishing in the ANNALS. Instructions to Authors are printed on the inside back cover of the last issue of each volume. Editorial Committee George K. Rogers and Nancy R. Morin Marshall R. Crosby Coeditors (this issue), Missouri Missouri Botanical Garden Botanical Garden Gerrit Davidse Janice Wilson: sy 5.3 Missouri Botanical Garden Editorial Assistant, Missouri Botanical Gardens o es | John D. Dwyer - S Missouri Botanical Garden & Saint Louis University m Peter Goldblatt —— Missouri Botanical Garden Ux Henk van der Werff _ Missouri Botanical Garden . Eleven, P.O. Box 299, St. Louis, MO 63166. Sub- SSN ery charge, $35 per volume. Four issues per vol- - Volume 74 Annals Number 1 of the NZ 1987 Missouri Botanical Garden SYSTEMATIC EMBRYOLOGY OF THE ANISOPHYLLEACEAE! HIROSHI TOBE? AND PETER H. RAVEN? ABSTRACT An embryological study of Anisophylleaceae, which comprise Anisophyllea, Combretocarpus, Poga, and Polygonanthus, and which have traditionally most often been referred as a tribe or subfamily to : PARIE t osome of bretocarpus is reported as n = 8, that of the ~~ apii genera as n = 7. Embryologically Anisophylleaceae are diversified and show differences from genus to genus, but they are clearly distinct m raceae in having their combination of con RUE baies states, including persistent nucellar tissue at least until i early stages of seed sp tng tou thin two cell-layered inner integument Poga and P In trast to Rhizophoraceae, Anisophylleaceae agree almost completely w ith Myrtales in their embryological features of the o rder. Embryological of A e Hd = `° "a = ° ° evidence iode: supports the bonu ily and, decus from other lines of evidence, suggests a Myrtale. n affinity for the family. Proposed assignments Anisophylleaceae to Rosales or to Cornales a " not sup ported. An analysis of similarities in Character states in the four genera the ra one leading to Anisophyllea a nd Combretocarpus, and the other leading to Poga ápd Politak anthus. Combretocarpus, with which Anisophyllea shares 3 few synapomorphies, is most specialized wit hema the family in having many apomorphies. In contrast, om and Polygonanthus share many ples morphies, most of which are also common to EE Anisophylleaceae, as defined here, consists of referred Polygonanthus to Euphorbiaceae (Ducke, four genera and 34 species, Anisophyllea (30 spp.), | 1932, 1933; Kuhlmann, 1940), Olacaceae (Croi- Combretocarpus (1 sp.), Poga (1 sp.), and Polygo- zat, 1939), Saxifragaceae (Baehni & Dansereau, nanthus (2 spp.) (Airy Shaw, 1973; Cronquist, 1939), or to its own family, Polygonanthaceae 1981, 1983). In contrast with the stable assign- (Croizat, 1943). Despite these, there is little doubt, ment of three other genera, various authors have on the basis of morphological and wood anatom- ! Supported by grants to the junior author from the U.S. National Science Foundation and in part by the Grant-in-Aid Scientific Research to the senior author from the Ministry of Education, Science and Culture, Kalkman, Duncan W. Thomas, and Mohd Shah Bin Mohd Noor for their collection and arrangement of the excellent material used in this study, and to Rolf Dahlgren, L. A. S. Johnson, Adrian Juncosa, and Richard Keating for their useful comments regarding this manuscript. ? Biological Laboratory, Yoshida College, Kyoto University, Kyoto 606, Japan. ? Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. ANN. Missouni Bor. GARD. 74: 1-26. 1987. TABLE |. Studied taxa and collections. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Taxa Collections Anisoph "d om JB (Jack in 1984, no voucher Singapore. Bukit Timah Nature Reserve. Sidek Bin Kiah & Tan Yam Leong s.n. Singapore. Botanic Garden, ides iei Sidek Bin Kiah s.n. in 1984, no voucher; Mohd Shah s.n. in 1984, no voucher. ass ag Maxwell Hill, Perak. B. C. Stone 15403, (KLU, MO). Brunei. A. M. Juncosa s.n. in 1981, no voucher. Anisophyllea sp. Combretocarpus rotun- Mi n Brunei Poga oleosa Pierre Cameroon. D. W. Thomas 3494, (MO). nue ay Kuching, Sarawak. P. Chai s.n. in 1981, 1983, and 1985, no voucher. . M. Juncosa s.n. in 1983, no voucher. Cameroon. Korup Natl. Park. D. W. Thomas 2273, (MO). Nigeria. Awi, Akamkpa. J. O. Ariwaodo s.n. in 1983, (FHI 99607). Polygonanthus amazoni- Brazil. Along the Rio Paca, Amazonas. J. Zarucchi 3138, 3184, (US). ical evidence, that Polygonanthus fits well in An- isophylleaceae, together with the three other gen- era that have tra iar been placed there (see Kuhlmann, 1944; Pir Rodrigues, 1971; Van Vliet, 1976). Of the ur genera of this family, Anisophyllea is relatively widely distributed in tropical Africa and Asia, also occurring in trop- ical South America; Combretocarpus is restricted to West Malaysia, Poga to tropical West Africa; and Polygonanthus to the Amazon Basin of Bra- zil (Pires & Rodrigues, 1971) The relationships of Anisophylleaceae have been controversial. A traditional view, and the one most widely accepted, is that Anisophylle- aceae have close affinities with Rhizophoraceae, and they often have been considered a tribe or subfamily within a broadly conceived Rhizopho- raceae (Bentham & Hooker, 1865; Baillon, 1877; been treated as a distinct family, Anisophylle- aceae have generally been considered closely re- Thorne (1983) placed Rhizophoraceae (com- posed of two subfamilies: Rhizophoroideae and Anisophylleoideae) in the Cornales. Recently, however, Cronquist (1981, 1983) concluded that Anisophylleaceae were not closely related to Rhizophoraceae and assigned them to izophoraceae sensu a stri Dahlgren (1983), who also denied any close re- lationships between Anisophylleaceae and Rhi- d idee pared „Anisophylleaceae in Cor- s(C its own ; see also Dahl- + t Dh; 1 1 (NÁ 4:4 or iis gren & Thorne, 1984). In the light of these diverse opinions, we have attempted to determine whether Anisophylle- aceae are actually closely related to Rhizophora- ceae sensu stricto or not, or whether they might even be grouped together as one family. If the two groups are not closely related, what are their respective affinities? Anisophylleaceae have been studied to a very limited extent, particularly regarding pes ana- tomical characteristics. Their wood anatomy, however, has been studied relatively intensive (Marco, 1935; Geh & Keng, 1974; n Vliet, 1976). Based on a comparison of the Mog anat- omy of the two groups, Van Vliet (1976) sup- ported a broad definition of Rhizophoraceae in- cluding Anisophylleaceae. In contrast, Behnke (1984), basing his conclusions on the features of their sieve-tube plastids, suggested Anisophyl- leaceae were quite distinct from Rhizophoraceae sensu stricto. He found that both Anisophyllea and COMMIELOE HS pave S-type plastids (con- J plastids (containing protein) that are character- istic of Rhizophoraceae sensu stricto. Another interesting distinction between the two groups, which has been known for some time, is that all four genera of Anisophylleaceae are aluminum accumulators; whereas the genera of Rhizopho- raceae sensu stricto are not (Chenery, 1948; Kukachka & Miller, 1980). Although embryo- logical information has been extremely useful in suggesting relationships at this level (see Tobe & 1987] Raven, 1983), almost no information is available on Anisophylleaceae. The only published data on ovule morphology (of **Anisophylleia zeylan- ica") is that of Karsten (1891) nearly 100 years ago; however, most of these observations seem to be incorrect, as we shall discuss subsequently. Vaughan (1970) described the mature seed coat structure of Poga, providing a drawing; Geh and Keng (1974) reported on the endosperm in the seeds of Anisophyllea and Combretocarpus. Ex- cept for these fragments of information, appar- ently nothing has been reported about the em- bryological features of Anisophylleaceae. In this paper, we present an overall study of the embryology of Anisophylleaceae, which is intended to provide information bearing on their relationships and systematic position. We have studied Anisophyllea and Combretocarpus in de- tail, and Poga and Polygonanthus to a lesser de- gree. Important features have been noted for all genera and are presented here. MATERIALS AND METHODS All four genera, Anisophyllea, Combretocar- pus, Poga, and Polygonanthus, were investigated in this study. The species we studied are listed in Table 1 together with their voucher infor- mation. Flower buds and fruits in all stages of development were collected and fixed in FAA (5 parts stock formalin; 5 parts glacial acetic acid; 90 parts 70% ethanol); however, female buds of Poga oleosa and fruits of Polygonanthus ama- zonicus were not available. Herbarium material of Anisophyllea and Combretocarpus was studied to supplement our observations of fruits and seeds. Preparations of microtome sections for obser- vation were made following standard paraffin techniques. After dehydration through a tertiary- butyl alcohol series, the samples were embedded in Paraplast with 56—58?C mp. Flower buds of Anisophyllea disticha and fruits of Poga oleosa were too hard to be sectioned without being soft- ened initially. Therefore, after these structures were trimmed to expose their tissues, the embed- ded samples attached to blocks were soaked in a mixture of a 10:3:90 glycerol: 10% Aerosol OT: water (Schmid & Turner, 1977) for at least several days at 20—25?C and then sectioned. Se- rial sections 6-10 um thick were stained with Heidenhain's hematoxylin, safranin, and fast- green FCF and were mounted in Entellan. Ma- ture seed coats of Anisophyllea sp. (D. W. Thom- as 3494, MO) and Poga oleosa, which were too thick and hard to be sectioned by standard par- TOBE & RAVEN —ANISOPHYLLEACEAE 3 affin techniques, were embedded in a JB-4 plastic and stained with 0.1% Toluidine Blue. In order to count the number of cells in mature pollen, we attempted to use safranin-staining of the grains (Tobe & Raven, 1984). We failed to obtain any staining ofthe pollen nuclei, however, probably because a thick exine hinders the infil- tration of dye. Consequently, we counted the number of cells in the pollen using microtome- sectioned pollen grains. The expressions we have used for the frequency of different shapes of mi- crospore tetrads follow those of Schmid (1982). OBSERVATIONS ANISOPHYLLEA R. BR. The embryological characteristics were basi- cally the same in the two species studied, one from Africa and one from Asia, The features reported in the foll to be common to both species, , unless particular vo dis are given. Anther and microspores. The anther is tetra- sporangiate. The wall prior to maturation com- prises basically five cell layers: an epidermis, an endothecium, two middle layers, and a tapetum (Fig. 1); the wall formation therefore conforms to be the Basic type (Davis, 1966: 10). The anther wall, however, often has only one middle layer, which shares a histogenetic origin with the ta- petum. The tapetum is glandular (Fig. 2). At one point in their development, the cells of the ta- petum become 2-nucleate, but subsequently the two nuclei fuse with each other. During matu- ration, the middle layer(s) degenerate and the epidermal cells are stretched tangentially while the cells of the endothecium become more or less enlarged (Fig. 2). Eventually, the endothecium develops fibrous thickenings. Although the epi- dermis persists, it is often collapsed on the en- dothecium (Fig. 3). Anther dehiscence takes place by longitudinal slits (Fig. 3). The connective tis- sue between the two microsporangia of each the- ca is completely disorganized before an anther dehisces (Fig. 3). Meiosis in a microspore mother cell is accom- panied by simultaneous cytokinesis, and the re- sultant microspore tetrads, on the basis of 50 selected tetrads (of Anisophyllea disticha), are **usually" (92%) tetrahedral, **very occasionally" (6%) decussate, and “rarely” (2%) isobilateral. The pollen grains are two-celled at the time of shedding (Fig. 4 Chromosomes. Since pollen mother cells be- tween the telophase of meiosis I and the meta- WCIC found 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 ES 1-5. Anisophyllea. — 1, 2, 4, 5. A. disticha. —3. A. sp. (D. W. Thomas 3494, MO).— 1. Transverse CN (TS) of a young anther showing the five cell-layered wall structure. Bar — 10 um.— 2. TS of an older anther with degenerating middle layers (arrow). Bar = 10 um.—3. TS of a developed anther. Its wall consists of the fibrous Mna epum and the epidermis. Bar = 50 um.—4. Two-celled mature pollen at the time of shedding. Arrows indicate nuclei of the two cells. Bar = 10 um.— 5. Chromosomes of a pollen mother cell at a stage between Victus I and metaphase II. n = 7. Bar = 10 um. ep, epidermis; et, endothecium; ml, middle layer; t, tapetum; mc, microspore mother cell. 1987] phase of meiosis II were fixed by chance and included in microtome sections, we were able to count the chromosome number of Anisophyllea for the first time: A. disticha has n = 7 (Fig. 5) Megagametophyte and nucellus. The ovule is anatropous. A single archesporial cell differ- entiates beneath the apical dermal layer of the nucellus (Fig. 6). The archesporial cell divides periclinally into two: the upper primary parietal cell and the lower sporogenous cell (Fig. 7). The primary parietal cell divides periclinally, and its derivatives further divide anticlinally and peri- clinally, forming parietal tissue with three to five layers above the embryo sac. The sporogenous cell develops into a megaspore mother cell and undergoes meiosis, giving rise to a linear tetrad of megaspores. A triad of megaspores may also be formed by suppression of the second, mitotic division on the micropylar side. In the mega- spore tetrad (or triad), the chalazal megaspore functions (Fig. 8). A functional megaspore de- velops successively into a 2- (Fig. 9), 4- (Fig. 10), and 8-nucleate embryo sac (Fig. 11). Thus the mode of the embryo sac formation is of the Po- lygonum type. The synergids are slightly hooked polar nuc which is positioned near the egg apparatus (Fig. 13). Consequently an organized mature embryo sac has only five nuclei or cells: an egg cell, two synergids, and two polar nuclei (as a single cen- tral nucleus; Figs. 12, 13). uring megasporogenesis and megagameto- genesis, apical epidermal cells of the nucellus di- vide periclinally, and their daughter cells also repeat periclinal divisions. As a result, a four to six cell-layered nucellar cap is formed above the embryo sac (Fig. 14); Karsten (1891) also illus- trated such a nucellar growth in “Anisophylleia zeylanica." The nucellar cap and the other nu- cellar tissue, both of which enclose the embryo sac, persist into younger stages of fruit devel- opment (Figs. 11, 14, 17). There is no case in which the nucellar tissue degenerates before fer- tilization so that the embryo sac directly borders on the integument. Integument. The ovule has a single integu- ment (Figs. 7, 15), although Karsten (1891) de- scribed the ovule of * "Anisophylleia zeylanica" nucellar tissue surrounding the embryo sac as the TOBE & RAVEN—ANISOPHYLLEACEAE 5 inner integument and the (only) true integument as the outer integument. A micropyle is always formed by the integu- ment, excepting one very unusual case in whic the integument did not grow beyond the nucellar apex (Fig. 16). In this respect as well, Karsten (1891) seems to have erred: he considered the persistent, lateral nucellar tissue to be the inner integument, which he concluded did not enclose the nucellar apex. In fact, Karsten concluded that a micropyle is not formed in “‘Anisophylleia zey- lanica.” Referring to Karsten’s drawing of ovules and descriptions, we also mistakenly character- ized the ovule of Anisophyllea not only as being bitegmic but also as having a nucellar beak (which actually was the well-developed persistent nu- cellar cap; see Tobe & Raven, 1983) The integument is about five to seven cells thick in Anisophyllea disticha (Fig. 15) and about four to five cells thick in A. sp. (Fig. 17). The thickness of the integument is not different from one part of ovule to another, and therefore the cross section of the ovule is nearly circular (Fig. 17). A raphe bundle ramifies oblique-laterally toward the chalazal end (Fig. 18). Therefore in cross section the ovule or fruit has four to five vascular bundles at the peripheral part of the integument of testa (Fig. 19). Throughout the development of the ovule or fruit, the integument or seed coat is thickened by secondary multiplication. However, the in- nermost cell layer never differentiates toward the so-called endothelium Fertilization, endosperm, and embryo. De- spite their multiovular condition in Aniso- phyllea (usually four and rarely three ovules per ovary), fruits were always one-seeded. Fertiliza- tion is porogamous. Endosperm formation is of the Nuclear type (Fig. 20). Because of incom- pleteness of our fruit sample, we could not con- firm whether or not wall formation takes place in free endosperm nuclei. Hand-sectioned ma- ture seeds of the two species we studied lacked endosperm (Fig. 21). Concerning the presence of endosperm in seeds, Hou (1958) described that in Anisophyllea disticha seeds consist of a solid ody, of which the main part is formed by a thick, hard albumen. Geh and Keng (1974) stated that in Anisophyllea disticha, the entire undiffer- entiated embryo is embedded in endosperm; consequently, they characterized the seed of Ani- sophylleae (Anisophyllea and Combretocarpus) as albuminous. Based on results of our obser- vations, however, it seems that what Hou thought 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FIGURES 6-14. Anisophyllea. —6, 7, 9-13. A. disticha. — 8. A. sp. (D. W. Thomas 3494, MO).—6. Longitudinal Mice (LS) of an ovular primordium with the 1-celled archesporium. Bar = 10 um.— 7. LS of a young ovule with the primary parietal cell. Bar — 10 um.— 8. LS of a young ovule with the con megaspore. Arrows the 2-nucleate embryo sac stage. Bar = 10 um.— 10. LS of an older ovule at the 4-nucleate Vigo sac stage. One of the two nuclei at the micropylar side appears in the next section. Bar — 10 um.— 11. LS of a nearly 1987] to be the thick, hard albumen was actually the embryo itself, and that the seed that Geh and Keng observed was too young to confirm the endosperm condition. Although we did not pursue the whole process of embryogenesis either, the development of cotyledons. Geh and Keng (1974), however, re- ported two protuberances on the apical part of the embryo in Anisophyllea disticha, which they interpreted as two cotyledons. We conclude that the cotyledons of Anisoph dps either develop in- completely or are essentially absent in Aniso- phyllea. No hypostase is a after fer- tilization. Mature seed and seed coat. The mature seed is narrowly cylindrical, 13.0-13.5 mm long and 3.8-4.0 mm thick in Anisophyllea disticha, whereas it is ovoid or elliptical in outline, 13.0— 13.8 mm long and 6.0—6.4 mm thick in A. sp. (Fig. 21). In the young seed, the seed coat appears to be constructed of a thick, massive tissue, with ple inner layer about 25—30 cells thick (Fig. 23). The cells of the outer epidermis are thick-walled and cuboid, whereas those of the underlying multiple inner layer are also thick- walled but extremely stretched tangentially. COMBRETOCARPUS HOOK F. Anther and microspores. The anther is tetra- ing maturation, the cells of the epidermis are somewhat enlarged and become tanniferous; the cells of the endothecium are also enlarged; the middle layers degenerate (Fig. 25). The tapetum is glandular, and its cells become 2-nucleate. The TOBE & RAVEN—ANISOPHYLLEACEAE 7 two nuclei in a tapetal cell later are fused with each other. Thus the mature anther wall is com- posed of the persistent but somewhat collapsed epidermis and the fibrous endothecium (Fig. 26). By the time of anther dehiscence, the connective tissue between two microsporangia of each teca degenerates completely. After dehiscence by lon- gitudinal slits, the anther wall is remarkably re- flexed (Fig. 2 Meiosis in ihe microspore mother cells is ac- companied by simultaneous cytokinesis. The shape of the resultant tetrads, on the basis of the examination of 50 selected tetrads, is “usually” (78%) tetrahedral, **occasionally" (1496) decus- sate, and “very occasionally" (896) isobilateral. The pollen ien are two-celled at the time of shedding (Fig. Chromosomes. “Pollen mother cells at the metaphase of meiosis I happened to be fixed and appeared in microtome sections. On the basis of those sections, we observed the chromosomes of Combretocarpus for the first time and deter- mined n = 8 (Fig. 29). Size differences seem to be present among those eight chromosomes. Megagametophyte and nucellus. The ovule is anatropous and crassinucellate. The archespo- rium is nearly always 1-celled (Fig. 30). A multi- cellular archesporium may very rarely differen- tiate—an ovule or young fruit containing twin ambs sacs was very ay observed (Fig. 3 Th lly into two: the upper primary parietal “cell and the lower sporogenous cell. The primary parietal cell may or may not divide further periclinally; if it does so, a two cell-layered parietal tissue is formed. The sporogenous cell develops into a megaspore mother cell (Fig. 31). After enlarging in volume, J ecome 2-nucleate (Fig. 32). spore is functional. Then, while the micropylar megaspore degenerates, the two nuclei in the — mature ovule at the 8-nucleate embryo sac stage. Of the eight nuclei, two polar nuclei are fused into a single central nucleus, and three antipodal cells are degenerating. Bar — 50 um.— 12, 13. Tw owing the egg apparatus and the central nucleus. Bars = 10 um mature ovule sho o serial LSs of a part of —14. Same as Figure 11, but at a lower magnification. Note that the nucellar tissue is persistent and that no cell layer of the integument shows differentiation into an endothelium. Bar = 50 sporogenous cell; fc, functional megaspore; n, nucleus of the em um. arc, SA cell; p, primary parietal cell; s, c; eg, egg cell; ega, egg apparatus; cn, central nucleus; ant, antipodal cell; sy, synergid; nuc, nucellar tissue; pig integument; cp, nucellar cap. 8 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURES 15-23. Anisophyllea. — 15, MO).— 15. Longitudinal section (LS) of a young ovule. Note that the avuke i is unite 18-20, 22. A. disticha.—16, 17, 21, 23. A. ics (D. W. Thomas 3494, Bar = 50 um.—16. LS um. —17. fae ries (TS) of a mature LS of a mature ovule tangentially cut through a raphe showing the ramification of a raphe bundle. Bar = 100 um.— 19. TS of a young seed. Note that the thick seed coat contains several vascular bundles at the peripheral part. Bar = 1 mm.— 20. LS ofa young seed containing a proembryo and free endosperm of an unusual mature ovule lacking a micropyle. Bar = 50 ovule. Bar = 50 um.— 18. 1987] TOBE & RAVEN— ANISOPH Y LLEACEAE 9 functional chalazal megaspore separate from each ther: one moves toward the micropylar end, while the other moves toward the chalazal end (Fig. 33). Each nucleus divides successively to form a 4- and an 8-nucleate sac (Fig. 34). Thus the embryo sac formation conforms to the bispor- ic Allium type. The synergids are slightly hooked, and the antipodals are ephemeral, disappearing before fertilization. Two polar nuclei do not fuse with each other until fertilization takes place; they are positioned near the egg apparatus. A mature embryo sac just before fertilization is composed of five nuclei or cells: an egg cell, two synergids, and two polar nuclei. Embryo sacs characteristically accumulate an abundance of starch grains (Fig. 35). The starch grains, which begin to accumulate from the 2-nucleate embryo sac stage, are most abundant in the 8-nucleate embryo sac stage but disappear after fertilization. During megasporogenesis and megagameto- genesis, the nucellar tissue does not show any particular differentiation and persists at least un- til the earliest fruit stages (Figs. 36, 38). Apical dermal cells of the nucellus do divide periclinally (Fig. 37), and their daughter cells also repeat peri- clinal divisions, thus forming a nucellar cap four to six cell layers thick above the embryo sac (Fig. 36) Integument. The ovule is unitegmic (Figs. 31, 36). The growing integument is about four or five cells thick (Figs. 36; see also Fig. 38). No differ- ence in thickness exists between the different parts of the ovule. Therefore, except for the raphe, the cross section of ovule is nearly circular (Fig. 38). The integument is not vascularized. Neither sec- ondary multiplication of the integument nor dif- ferentiation of the innermost cell layer into a so- called endothelium occur. e integument elongates beyond the nucellar rogamous. After fertilization, the fruit elongates remarkably (Fig. 39). Endosperm formation is of the Nuclear type (Fig. 40). In the early stages, free endosperm nuclei are located around the proembryo (Fig. 40) and at the peripheral region of the embryo sac (Fig. 43). Because of incom- pleteness of our fruit sample, we could not ob- serve to what degree an amount of endosperm increases later. The mature seeds lack endosperm (Figs. 41, 42). We did not investigate embryo- genesis in detail but can state, on the basis of our observations of a few microtome-sectioned proembryos, that it proceeds normally (Fig. 40). ithin the mature fruit of Combretocarpus, the embryo is elongate and nearly circular in cross section (Figs. ). e embryo is cotyledonous with so small cotyledons ele a long hypocotyl (Fig. 41 The hypostase is not differentiated even after fertilization. Mature seed and seed coat. The mature seed is linear, 9.5-10.4 mm long and 1.2-1.3 mm thick; it contains several vascular bundles in the raphe (Fig. 42), which are derived by ramifica- tion from a raphe bundle. These bundles are re- stricted to the raphe, never entering the integu- ment or testa. In the young seed, the seed coat is composed of a tanniferous outer epidermis and a multiple inner layer, which degenerates (Fig. 43). Even- tually, in the mature seed, the seed coat com- prises only the outer epidermis, which is formed of pigmented, cuboid cells (Fig. 44) POGA PIERRE Anther and microspores. The anther is tetra- sporangiate. The wall prior to maturation com- prises five cell layers: an epidermis, endotheci- um, two middle layers, and a tapetum (Fig. 45). Wall formation conforms to the Basic type. Dur- ing maturation, cells of the epidermis as well as of the endothecium enlarge, while the middle layers degenerate (Fig. 46). The tapetum is glan- dular, and its cells become 2-nucleate (Fig. 46). The two nuclei in a tapetal cell are not fused with each other. The mature anther wall is composed of the persistent epidermis and the endothecium. The epidermis is tanniferous, and the endothe- cium develops fibrous thickenings (Fig. 47). The anther dehisces by longitudinal slits. By the time of dehiscence, the connective tissue between two < nuclei. Bar = 20 um.— 21. Longitudinal hand-section ofa mature seed. ^ Bar 44 4 14 ar = 5 mm.— 22. LS of a young seed showing a thick seed coat. Bar = 200 um.— 23. LS of a mature seed coat that is formed by both the multiple inner layer and the conspicuous outer epidermis. Bar — 40 um. in, integument; nuc, nucellar tissue; rb, raphe bundle; b, vascular bundle; pem, proembryo; fe, free endosperm nucleus; em, mbryo ; SC, seed coat; epi, epidermis of seed coat; inl, multiple inner layer. 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 d i reflexed. Bar = 50 um.— 28. Two-celled mature pollen at pi time of shedding. Bar = 10 um.— romosomes of pollen mother cell at metaphase I. n — & Bar = 2 um. ep, epidermis; et, endothecium; ml, middle layer; t, tapetum; mc, microspore mother cell. 1987] microsporangia of each theca degenerates com- l tely. Meiosis in the microspore mother cell is ac- companied by simultaneous cytokinesis. The shape of the resultant tetrads, on the basis of the examination of 50 selected tetrads are “usually” (86%) tetrahedral, “occasionally” (12%) decus- sate, and “rarely” (2%) isobilateral. The pollen grains are 2-celled at the time of shedding (Fig. 48 Chromosomes. Using serially sectioned pol- len mother cells that were fixed at the later pro- phase of meiosis I, we observed the chromo- somes of Poga for the first time and determined the chromosome number n = 7 (Figs. 49-51). Nucellus and integuments. Although female flowers were not available, we confirmed by us- ing mature ovules and very young fruits that the nucellar tissue enclosing the embryo sac persists at least until the early stages of fruit development (Figs. 52, 54). No hypostase is differentiated even after fertilization. The ovule is bitegmic, i.e., possessing the outer and the inner integument (Fig. 52). The outer integument is originally about four or five cells thick, and the inner integument two cells thick (Fig. 53). The cells of the outer epidermis of the outer integument, which later become those of the outermost layer of the exotesta, are conspic- uously enlarged into cuboid cells. The raphe bun- dle ramifies and vascularizes the outer integu- ment. In a cross section of a young fruit, six to eight vascular bundles in addition to several raphe bundles are observed in the testa (Fig. 54). The lom is formed by both integuments (Fig. 5 l and embryo. We could not ob- serve either the mode of endosperm formation or embryogenesis. But we can say at least that the mature seed completely lacks endosperm (Figs. 55, 56) as Vaughan (1970) described, and that the embryo does not have cotyledons. Con- cerning the cotyledons, Vaughan (1970) men- tioned that they are fused. However, judging from the resemblance in exomorphology of the em- bryo with Anisophyllea, it seems that Poga also lacks cotyledons from the beginning. Mature seed and seed coat. The fruits are al- ways one-seeded. The mature seed is 20.0-22.5 mm long and 12.0-13.5 mm thick, is ovoid and slightly suppressed toward the raphe-antiraphe direction, and has a thick, dark brown seed coat (Figs. 55, 56). TOBE & RAVEN—ANISOPHYLLEACEAE 11 In the young seed, the seed coat is composed only of a thick testa and lacks a tegmen. It seems that, during the process of seed development, the inner integument or tegmen is crushed, while the outer integument or testa increases in thickness by secondary multiplication. Within the young testa, a differentiation into a multiple outer layer and a multiple inner layer can be observed (Fig. 57). The multiple outer layer is about 8 cells thick and has cells that are more or less enlarged. In contrast, the multiple inner layer is 7—10 cells thick, with the cells stretched tangentially (Fig. 57). The structure of the mature seed coat ba- sically does not differ from that of the young seed coat. In the mature seed coat, however, the walls of the constituent cells are thickened, and the multiple inner layer occupies nearly one-third of the whole thickness ofthe testa, with the multiple outer layer occupying the remaining two-thirds (Fig. 58). Because our microtome sections of the seed coat were not very good, we could not ex- amine the details of cell structure. Vaughan (1970), however, gave a drawing of the anatom- ical structure of the testa, which consists of an inner layer that is about 12 cells thick and an outer layer about six or seven cells thick. Refer- ring to Vaughan (1970), Corner (1976) described the outer epidermis of the multiple outer layer as composed of cuboid cells with slightly thick- ened, lignified walls, and the other cells of the multiple outer layer as thin-walle The mature seed coat structure of Poga, which is bitegmic, seems comparable with that of An- isophyllea, which is unitegmic. In both genera, the mature seed coat contains a similar (probably identical) multiple inner layer, which is charac- teristically composed of tangentially stretched cells with thick walls. The only evident difference between the mature seed coat structure of Poga and that of Anisophyllea lies in thickness of the outer layer, i.e., about six or seven cells thick in Poga (see Vaughan, 1970) and one cell thick (out- er epidermis only) in Anisophyllea. In other words, T seed coat of Anisophyllea, like that of oga, may also be constructed principally of the 34 or “outer Integument”), which of course is not differentiated in the single integument of Anisophyllea. Therefore the seed of unitegmic Anisophyllea and even of unitegmic Combreto- carpus (with a mature seed coat consisting only ofthe outer epidermis) may be regarded as testal, and the seed of bitegmic Poga can also be defined in this way. 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FIGURES 30-38. _Combretocarpus rotundatus. — 30. Longitudinal section (LS) of an ovule with the 1-celled archesporium. Bar = 10 um.—31. LS of a young ovule with the primary parietal cell and the megaspore mother cell. Note that the ovule has only a single integument. Bar = 10 um.— 32. LS ofa young ovule with the Beers gii dyad. Note that each megaspore has two nuclei. Bar = 10 um.—3. LS of an ovule at the 2-nucleate embryo sa stage. Bar — 10 um.— 34. LS of a nearly mature ovule at the 8- nucleate embryo sac stage. Bar = 10 uv idee 1987] POLYGONANTHUS DUCKE Anther and microspores. The anther is basi- cally tetrasporangiate. The microsporogenous tissue, however, is occasionally transversely di- vided by a septum composed of tapetal cells (Fig. 59). Although we could not determine the modes of anther wall formation, the wall prior to mat- uration comprises five cell layers: an epidermis, endothecium, two middle layers, and a tapetum. During maturation, the middle layers degener- ate, while the cells of both the epidermis and the endothecium become enlarged (Fig. 59). The ta- petum is glandular, and its cells become 2-nu- cleate before degeneration (Fig. 60). The two nu- clei in a tapetal cell do not fuse with each other. Eventually the mature anther wall is composed of a persistent epidermis, whose cells are some- what collapsed in places, and a fibrous endothe- cium (Fig. 61). The connective tissue between two microsporangia of each pius b completely before the anthers dehi pollen grains are 2-celled at the time of shedding (Fig. 62). Chromosomes. Using serially sectioned mi- crospore mother cells that were fixed at the late prophase of meiosis I, the chromosomes of Po- lygonanthus were observed for the first time. Throughout, at examination and reconfirmation in many cells, we determined the chromosome number of P. amazonicus as n = 7 (Figs. 63-65). Megagametophyte and nucellus. Although our observations are fragmentary, we were able to observe some aspects of the process of mega- sporogenesis and megagametogenesis in Polygo- nanthus. The ovule is anatropous and crassinu- cellate. At least one parietal cell is cut off above the megaspore mother cell (Figs. 66, 67). Al- though the mode of embryo sac formation was not determined, the 2- nucleate a sac of ct from ‘hit ‘of Combretocarpus rotundatus Gi de- velops a bisporic A//ium type embryo sac) (Fig. TOBE & RAVEN —ANISOPHYLLEACEAE 13 68; compare with Fig. 33). The accumulation of starch grains in the embryo sac, which is char- acteristic of Combretocarpus, does not occur in Polygonanthus amazonicus. The antipodal cells are probably ephemeral, because they are absent in the organized mature embryo sacs (Fig. 69) The nucellar tissue enclosing the embryo sac is persistent until at least the stage of fertilization (Fig. 69). Periclinal divisions occur in the apical dermal cells of the nucellus. Therefore the nu- cellar cap is probably formed by derivatives of the apical dermal cells. Integuments. The ovule is bitegmic, i.e., it has both an outer and an inner integument (Fig. 66). The outer integument is initially about five cells thick, but later becomes seven to nine or more cells thick because of secondary multipli- cation (Fig. 66). The inner integument, in con- trast, is two cells thick (Fig. 67). In the later stages, u g t creases in cn t 69). Although we could not observe any stages of the development of the seed coat, it seems very unlikely that the inner integument or tegmen contributes to its structure when mature. The raphe bundle ramifies throughout the outer integument, which is there- fore vascularized. In cross section, seven or eight bundles in siii to several raphe bundles are observed (Fig he be is ‘fered by both integuments (Fig. 69). DISCUSSION Our own results on the embryology and chro- mosome numbers of Anisophylleaceae, together with some data on ovule and seed morphology published earlier (Karsten, 1891; Vaughan, 1970; Geh & Keng, 1974), are presented in Table 2. On this basis, we summarize the embryological features of Anisophylleaceae as follows Anther tetrasporangiate, but occasionally polysporangiate because of insertion of tapetal septa (Polygonanthus); anther wall with five cell — Polarized view of the same as that shown in Figure 34, showing a conspicuous accumulation of starch grains in the embryo sac. Bar = 10 um.—36. LS ofa mature ovule with an | organized embryo sac. Note that the nucellar tissue is persistent. Bar = 50 um ofay divisions occurring in apical epidermal cells of the nucellus. Bar = = 10 umn. — —38. Transverse section ofa young seed with twin embryo sacs. Bar cell; in, integument; n, nucleus O um. arc, archesporial cell; p, primary p of the embryo sac: sy, a m pos eta ja purius a cell; nuc, nucellar cell; mc, megaspore mother tissue; pd, periclinal cell division; sc, seed coat; es, embryo ANNALS OF THE MISSOURI BOTANICAL GARDEN MEO v | [T Q) £ L hI IGURES 39—44. deer dud rotundatus. — 3 i —40. LS of a proembryo with m endosperm nuclei surrounding it. Bar — 50 um. .—41. fruit. Bar — el i wath a cotyledonous embryo. Not e that the mature seed is exalbuminous. Bar = 1 mm.— of a matur 42 “atuntami section of a mature seed with several vascular bundles at the raphe. Bar = 500 um.—43. LS of a young seed coat Bar = 10 um.—44. LS ofa mature seed coat. Bar = 10 um. pem, proembryo; fe, free endosperm nucellus; cot, cotyledon; em, embryo; sc, seed coat; rb, raphe bundle layers, its formation of the Basic type; anther epidermis persistent, consisting of more or less collapsed cells; endothecium persistent and de- veloping fibrous thickenings; middle layers ephemeral; tapetum glandular, its cells 2-nu- tually fused in Anisophyllea and Combretocar- pus, but not in Poga and Polygonanthus. 1987] TOBE & RAVEN—ANISOPHYLLEACEAE 15 FIGURES 45-51. Poga oleosa. —45. Transverse section (TS) of a young anther showing the wall structure with . TS of an older w). Bar = 10 five cell layers. Bar = ofa ther with degenerating middle layers (arro m.—47. TS of a nearly mature ther wall consists of the fibrous endoth m and the epidermis. Bar = 100 um.—48. Two-celled mature pollen at the time of sheddi ndicate nuclei of the two cells. Bar = 10 e serial sections of pollen mother cell at the late prophas win romosomes of n um.— 49-51. Thre S 7. Seven chromosomes are numbered | to 7. Bars = 10 um. ep, epidermis; et, endothecium; ml, middle layer; t, tapetum Cytokinesis in the microspore mother cells si- Gametic chromosome number n = 8 in Com- multaneous; microspore tetrads tetrahedral, de- — bretocarpus, n = 7 in Anisophyllea, Poga, and cussate, or isobilateral; pollen grains 2-celled Po/ygonanthus. when shed. Ovule anatropous and crassinucellate; arche- 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURES 52-58. Pogaoleosa.—52.1 itudinal section (LS) of ote that the ovule is a and has a persistent nucellar tissue. Bar = 100 um.— 53. Transverse section Mes + a mature ovule showing the inner integument, with two cell layers, and the eae hg oy with four cell layers. Bar = 10 um. M TS ofa = 500 um 5. Longitudinal hand-section of the mature sced. Note that the mature seed is exalbuminous. Bar = 5 mm.— — 56. Transverse hand-section of the mature seed. Bar = 5 mm.— 57. TS of a young seed coat. Note that there is no tegmen and that the testa is E into the multiple i inner layer and the multiple outer layer. Bar = 100 um.— 58. LS of a mature seed coat. Bar = m. ii, inner integument; oi, outer integument; es, embryo sac; nuc, nucellar tissue; rb, eee bundle: b, vascular bundle; sc, seed coat; em, embryo; inl, multiple i inner layer; oul, multiple outer layer 1987] TOBE & RAVEN—ANISOPHYLLEACEAE 17 Ficures 59-65. Polygonanthus amazonicus. — 59. Longitudinal section is a developed anther. Arrow indi- cates the a septum dividing a microsporangium. Bar = 50 um.—60. Transverse section of a young Tax š Bar = 10 um.— 61. TS of a mature anther at the time of dehiscence. ‘Its wall consi of the fibrous endothec and the ME Bar = 50 um.— 62. Two-celled mature pollen at the time of shedding. Arrows indicate nuai of the two cells. Bar = 10 um.—63-65. Three serial sections of pollen mother cell at E. prophase I showing chromosomes of n = 7. Seven chromosomes are numbered 1 to 7. Bars = 10 um. t, tapetum; ep, epidermis; et, endothecium. 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FiGURES 66-70. Polygonanthus amazonicus.—66. Lon gitudinal section (LS) of a young phon Note that the ovule is bitegmic. Bar = 50 um.— 67. stage. Bar — 10 um. — ol, Outer integument; p, pri imary parietal cell; mc, geie Same as that shown in Figure 66, that the inner integument is two cells thick. Bar = 20 4 9. LS of a mature ovule. Aies in: the nucellar tissue is persistent. outer integument has vascular but shown at higher magnification. Note at the 2- ien ons embryo sac Bar = 50 um.— 70. bundles. ii, inner integument, r cell; n, nucleus of the embryo sac; es, m.—68. LS of an ovule ore mother mbryo sac; nuc, nucellar tissue; rb, raphe bundle; b, vascular bundle. ari l sus ine: off a primary parietal mbryo formation of the Polygonum oe » nisoph opk or the Allium type (Combre- tocarpus), synergids slightly hooked; antipodals ephemeral; polar nuclei fused before fertilization (Anisophyllea) or not fused (Combretocarpus). Nucellar tissue not degenerating at least until the early stages of seed development; apical dermal nucellar cells dividing periclinally, forming a nu- cellar cap, chalaza without a hypostase. Ovule unitegmic (Anisophyllea and Combre- tocarpus) or bitegmic (Poga and Polygonanthus); in bitegmic ovules, the inner integument two cells thick and the outer integument thicker; outer integument vascularized due to ramification of raphe bundles, but not vascularized in Combre- 1987] tocarpus, micropyle formed by either the one uclear type; seed exalbuminous; mode of embryogenesis not determined; embryo (potentially) dicotyledonous with a long hypo- cotyl, having either small cotyledons (Combre- tocarpus) or rudimentary and/or no cotyledons (Anisophyllea and Poga). Seed coat testal (Poga) or logically testal (Anisophyllea and Combreto- carpus); mature seed coat formed by the outer epidermis alone (Combretocarpus), both the out- er epidermis and the multiple inner layer (Ani- sophyllea), or both the multiple outer layer and the multiple inner layer (Poga). RELATIONSHIPS WITH RHIZOPHORACEAE Although, as shown in Table 2, some of the embryological features of Anisophylleaceae are diverse, the family is consistent enough in most such characteristics to allow a more critical com- parison with Rhizophoraceae than has hitherto been possible. In summary, these two families share only a few embryological features. They do agree, for example, in having a crassinucellate, bitegmic ovule and the Nuclear type of endo- sperm formation; but a combination of these and other shared features is widespread among many other unrelated families of the angiosperms as well. In contrast, psi Sana ipie from Rhi- zophoraceae in som portant embryological features. First of all, in i yaya the nu- ceae the nucellar tissue is ephemeral, disappear- ing completely by the time of fertilization (see Karsten, 1891, for Rhizophora, Ceriops, Bru- guiera, and Carallia; Cook, 1907, for Rhizopho- ra; Carey, 1934, for Rhizophora; Mauritzon, 1939, for Gynotroches; Juncosa, 1984a, 1984b, for Bruguiera and Cassipourea). Therefore in Rhizophoraceae the Midi sac borders directly on the inner integum Secondly, in ee NR (Poga and Po- lygonanthus) the inner integument is character- istically two cells thick, whereas in Rhizophora- ceae it is much thicker. Indeed, an inner integument with four to eight layers has been illustrated by Karsten (1891) for Bruguiera, Ceri- ops, and Carallia, by Carey (1934) for Rhizopho- ra, and by Mauritzon (1939) for Bruguiera and TOBE & RAVEN — ANISOPHYLLEACEAE 19 Gynotroches. Juncosa (1984a, 1984b) described the inner integument of Bruguiera exaristata as initially being about 10 cells thick and that of Cassipourea elliptica as being about five to eight cells thick. In addition, a specialization of the innermost cell layer of the inner integument into an endothelium has been reported in some in- land genera of Rhizophoraceae, including Car- allia (Karsten, 1891), Gynotroches (Mauritzon, 1939), and Cassipourea (Juncosa, 1984a). An en- dothelium is never formed in Anisophylleaceae. Thirdly, the mature seed is exalbuminous in Anisophylleaceae, but albuminous in Rhizopho- raceae. The presence of abundant endosperm in mature seeds has been reported for Rhizophora (Cook, 1907; Carey, 1934; Juncosa, 1982), Ceri- ops (Carey, 1934), and Cassipourea (Juncosa, 1984a Some critical differences in embryo and seed coat morphology might also be added It distin- guishing / (see Comer, 1976). 2 evel, studies on those ll too limited to allow this. Further studies on the embryology of Rhizophoraceae, including embryo and seed coat morphology, are needed to clarify the dif- ferences between this family and Anisophylle- aceae To sum up, despite insufficient information on the embryology of Rhizophoraceae, the available data indicate that Anisophylleaceae differ sig- nificantly from them. If Anisophylleaceae were included as a tribe or subfamily, Rhizophoraceae sensu lato would be defined very broadly. With the support of exclusive occurrence of the nature of aluminum accumulation (Chenery, 1948; Ku- kachka & Miller, 1980); alternate, exstipulate leaves; three or four free styles (Geh & Keng, ai Anisophylleaceae and Rhiz closely related and warrants regarding Aniso- phylleaceae as a distinct family. SYSTEMATIC POSITION OF ANISOPHYLLEACEAE Cronquist (1981, 1983) has proposed assigning Within Rosales, (Hydrangeaceae, Columelliaceae, Grossulari- aceae, Greyiaceae, Bruniaceae, and Alseuosmi- aceae). Of these, only Grossulariaceae have been 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE 2. Embryological and chromosomal data of Anisophylleaceae. [VoL. 74 Number of integu- ments Thickness of integ- uments when bi- tegmic i.i. 2 cell-layered; 0.i. 4—5 cell-lay- ered Character Anisophyllea Combretocarpus Poga Polygonanthus Anther and microspores: Number of sporan- 4 4 4, sporangium occa- gia sionally divided Anther wall devel- Basic type Basic type Basic type opment Anther epidermis Persistent Persistent a Persistent Endothecium Fibrous Fibrous Fibro ibrous Tapetum Glandular Glandular Cur Glandular Number of tapetal 2 2 2 2 nuclei Tapetal nuclear fu- Occur Occur Not occur Not occur sion Cytokinesis in Simultaneous Simultaneous Simultaneous Simultaneous meiosis Shape of micro- Usually tetrahe- Usually tetrahe- Usually tetrahe- ? spore tetrad dral, very occa- dral, oo dral, occasional- sionally decus- ly dec ly decussate, : very aie rarely isobilater- isobilateral isobilateral a Mature pollen 2-celled lled 2-celled 2-celled Chromosomes: Base number x27 x=8 x=7 x27 Megagametophyte and nucellus: Ovule curvature Ana Anatropous Anatropous Anatropous Nature of nucellus Crassinucellate Crassinucellate Crassinucellate Crassinucellate Archesporium 1-celled arly al ? ? 1-celled, very rarely 2-celled Thickness of pari- 3-5 cell-layered 1-2 cell-layered ? ? etal tissue Shape of mega- Linear Linear ? ? spore tetrad Functional mega- Chalazal cell Chalazal cell ? ? spore Pattern of embryo Polygonum type Allium type T ? sac formation Synergids Slightly hooked Slightly hooked ? ? Antipodal cells Ephemeral Ephemeral ? Probably ephemeral r of nuclei 5 5 ? 5 or 5 cells in ma- e embryo sac Accumulation of Not occur Occur ? Not occur starch grains in embryo sac Nucellar tissue Persistent Persistent Persistent Persistent Nucellar cap orme Formed ? Probably formed Hypostase Not formed Not formed Not formed Not formed Integuments: l l 2 2 1.1. 2 cell-layered; o.i. about 5 cell- layered 1987] TABLE 2. Continued. TOBE & RAVEN—ANISOPHYLLEACEAE 21 Character Anisophyllea Combretocarpus Poga Polygonanthus Vasculature Present Absent Present Present Micropyle forma- By the only integu- By the only integu- By both integu- By both integu- tion ment ment ments ments Differentiation of Not occur Not occur Not occur Not occur endothelium Fertilization, endosperm, and embryo: Path of pollen tube Porogamous Porogamous ? ? Tem forma- Nuclear type Nuclear type ? ? ius in ma- Absent Absent Absent ? ture see Embryogenesis ? ? ? ? Embryo in mature e Or Cotyledonous Not cotyledonous ? seed ot cotyledonous Size of cotyledons un small (rudi- Small — ? when present mentary) Mature seed and seed coat: Shape of seed Narrow-cylindrical Linear Ovoid and slightly ? (A. disticha); suppressed to- ovoid or ellipti- ward raphe-an- cal (A. sp.) tiraphe direction Size of seed 13.0-13.6 mm 9.5-10.4 mm long 20.5-22.5 mm ii. = 3.8-4.0 and 1.2-1.3 mm long, 12.0-13.5 m. (A. diam. mm wide pus 13.0- 13.8 mm long and 6.0-6.4 mm diam (A. sp.) Tegmen — — Ephemeral ? Whole thickness of 26-31 cell-layered 1 cell-layer 17-20 cell-layered ? see C Thickness of inner 25-30 cell-layered — 7-10 cell-layered 7 layer o Thickness of outer l cell-layer 1 cell-layer About lO cell-lay- ? layer of SC ered i.i., inner integument; o.i., outer integument. relatively well studied embryologically, whereas the others have been studied little or not at all Grossularine h exalbuminous seeds (Cronquist, 1). On the other hand, so- phylleaceae resemble Grossulariaceae (princi- pally Ribes, from which most data are available) in nearly all features of anther and microspore development; in their anatropous, bitegmic, and crassinucellate ovule; Polygonum-type embryo sac; ephemeral antipodal cells; inner integument with two cell layers (see Davis, 1966; Corner, 1976; Cronquist, 1981, for data on Grossulari- aceae). However, Anisophylleaceae differ from Grossulariaceae in several embryological fea- tures. For example, the tapetal cell is basically 2-nucleate in Anisophylleaceae, but multinu- cleate in Grossulariaceae; the integument is vas- cularized in Anisophylleaceae, but not vascular- ized in Grossulariaceae; endosperm formation is of the Nuclear type in Anisophylleaceae, but of the Cellular or the Helobial type in Grossulari- aceae; the tegmen is ephemeral in Anisophylle- aceae (Poga), but persists in Grossulariaceae; the seeds are non-arillate in Anisophylleaceae, but arillate in Grossulariaceae (see 3j que 1926; Davis, 1966; Corner, 1976, for data on Gros- sulariaceae). Therefore it seems that available R In contrast, Dahlgren (1983) pum Aniso- 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN phylleaceae in the Cornales, which comprise 27 families including Hydrangeaceae (and five fam- ilies whose position is uncertain; see also Dahl- gren & Thorne, 1984). Of 27 families, nine have either not been studied embryologically, or have been studied only to a very limited degree. Of the 18 remaining families, nearly all share a uni- tegmic ovule, ephemeral nucellar tissue, endo- thelium, Cellular type of endosperm formation, and albuminous seed. The Cornales thus seem to be very well defined by a combination of those shared embryological features. Anisophylle- aceae, which lack any of those characteristic em- bryological features of the Cornales (almost cer- tainly unitegmic ovule in Anisophyllea and Combretocarpus), seem clearly distinct from the Cornalean families and do not belong in that order. We would rather suggest Myrtalean affinities for Anisophylleaceae. Embryologically, Aniso- phylleaceae agree almost completely with Myr- tales, and in fact share the eight ordinal embry- ological features (see Tobe & Raven, 1983, 1984): 1) anther tapetum glandular, 2) ovule crassinu- cellate, 3) inner integument two cells thick, 4) micropyle formed by both integuments, 5) an- tipodal cells ephemeral, 6) endosperm forma- tion— Nuclear type, 7) seed exalbuminous, and 8) mature pollen 2-celled. One might point out a fusion of tapetal nuclei (in Combretocarpus and Anisophyllea), formation of the nucellar cap, and testal seed as features distinguishing Anisophyl- leaceae from Myrtales. However, nuclear fusion in the tapetal cells is undoubtedly a secondary characteristsic that evolved in two genera of An- isophylleaceae. Indeed Poga and Polygonanthus, both of which have many primitive features, as will be discussed later, have unfused tapetal nu- clei. The nucellar cap, which is formed by deriv- atives of the apical nucellar dermal cells, is com- monly observed in Combretaceae (Myrtales; Venkateswarlu & Rao, 1972). A seed coat lacking a tegmen is frequent in Melastomataceae (Myr- tales; Corner, 1976). Anisophylleaceae may dif- fer from Myrtales in having embryos with re- duced or rudimentary cotyledons and a long hypocotyl. Such an embryo morphology seems to result in hypogeal germination, which is re- ported in at least Anisophyllea disticha (Geh & Keng, 1974). The peculiar embryo morphology and germination habit may suggest a specialized position of Anisophylleaceae. However, embryos devoid of cotyledons are recorded in many un- related families, a majority of them growing in [VoL. 74 ecologically specialized habitats (Natesh & Rau, 1984, review). Study of embryogenesis and or- ganogenesis in seeds through germination seems to be needed for the elucidation of the ecological significance of such specialized embryos in An- isophylleaceae. Except for the difference in em- bryo morphology, there seems to be essentially a perfect correspondence in embryological fea- tures between Anisophylleaceae and Myrtales. Viewing other reproductive and vegetative character states, Anisophylleaceae lack both the intraxylary phloem and the vestured pits, which are regarded as characteristic features defining the Myrtales (Van Vliet, 1976; Van Vliet & Baas, 1984; Dahlgren & Thorne, 1984). However, the occurrence of S-type sieve-element plastids in Anisophyllea and Combretocarpus, in contrast with the P-type plastids in Rhizopt agrees with Myrtales (Behnke, 1982, 1984). Tricolpor- ate pollen morphology in Anisophylleaceae (as well as in Rhizophoraceae) is of the basic type found in the Myrtales (Erdtman, 1966; see also Dahlgren & Thorne, 1984). Aluminum accu- mulation characteristic of Anisophylleaceae (un- known in Rhizophoraceae) is known to occur in Crypteroniaceae and Melastomataceae of Myr- tales (Chenery, 1948; Kukachka & Miller, 1980). Thus, considering a considerable number of coincidences (in reproductive anatomy) in con- trast with a limited number of differences (in wood anatomy), in conjunction with support by sieve-element plastid type, palynology, and alu- minum accumulation, it seems that Anisophyl- leaceae are closely related to Myrtales. Depend- ing on how we interpret the lack of intraxylary phloem and vestured pits in Anisophylleaceae, it might even be justifiable to place Anisophyl- leaceae in the Myrtales. According to Van Vliet and Baas (1984), the combined occurrence of intraxylary phloem and vestured pits is very re- stricted in the dicotyledons; in fact, except for the Myrtales, this combination is found only in part of the Gentianales, Thymelaeales, Poly- galales, and Polygonales. Outside these orders, either one of these features (but not both) spo- radically occurs in many different groups of di- cotyledons (see Van Vliet & Baas, 1985: 784, fig. 1). Only a few orders are characterized by con- sistent possession of one or both of those two wood anatomical features. Therefore it does not seem that the lack of intraxylary phloem and vestured pits in Anisophylleaceae necessarily precludes a possibility of close affinity with Myr- tales. Based on total evidence now available, we 1987] would suggest that Anisophylleaceae be placed near Myrtales. Perhaps Anisophylleaceae rep- resent one of the groups that diverged from a common ancestral stock with Myrtales and then spread widely. The position of Anisophylleaceae will be evaluated better as the Rosiflorae or the Rosales, which are considered phylogenetically basic in position with respect to Myrtales, are understood better embryologically. INTERRELATIONSHIPS AND EVOLUTION OF THE GENERA Because of many shared embryological fea- tures, as shown in Table 2, as well as of shared vegetative and some other shared reproductive features (see also Geh & Keng, 1974; Van Vliet, 1976), there is no doubt that Aniopibiled. Com- bretocarpus, Poga, and Polygonanthus are mono- phyletic. Despite the lack of data about several features in Poga and Polygonanthus, the avail- able embryological data are now enough to allow us to compare all four genera. Of these, Combretocarpus is the most distinct. It has a gametic chromosome number of n = 8, Allium type embryo sac, nonvascularized integ- ument, starch grains in the embryo sac, cotyle- donous embryo, and thin mature seed coat one cell layer thick. In contrast, Anisophyllea, Poga, and Polygonanthus have a chromosome number of x = 7, Polygonum type embryo sac (unknown in Poga and Polygonanthus), vascularized integ- ument, no starch grains in the embryo sac, non- cotyledonous embryo (unknown in Polygonan- thus), and thick mature seed coat (unknown in Polygonanthus). Combretocarpus agrees with Anisophyllea only in having fused tapetal nuclei and a unitegmic ovule. On the contrary, Aniso- phyllea differs from Poga and Polygonanthus in sharing neither bitegmic ovules nor distinct ta- petal nuclei as well as in not sharing a multiple outer layer in the mature seed coat (though un- certain in Polygonanthus). Polygonanthus differs from Poga only in its occasionally divided mi- crosporogenous tissue. Except for this, there is no essential difference between Poga and Poly- gonanthus, as far as the data available are con- cerned. In order to clarify phylogenetic interrelation- ships ofthe genera, it seems necessary to evaluate each of the characters showing differences be- ua mbryologi phylleacca following Eldredge and Cracraft (19 ° + 4 o TOBE & RAVEN—ANISOPHYLLEACEAE 23 of character state similarities (i.e., synapomor- phies or symplesiomorphies). Of the embryolog- ical features, the Polygonum type embryo sac formation (Anisophyllea) that is characteristic of a majority of the dicotyledons (Davis, 1966) is undoubtedly primitive to the Allium type man- ner (Combretocarpus), and also bitegmy (Poga and Polygonanthus) is primitive to the unitegmy (Anisophyllea and Combretocarpus, Bouman, 1984). Concerning the vasculature of the integument, there 1s no consensus regarding whether or not the vascularized integument represents an ar- chaic condition. Bouman (1984) suggested that there seems to be a general relation between the size of ovules or seeds and the degree of vascu- larization. As far as Anisophylleaceae are con- cerned, the vascularized integument or testa (Ani- sephyllet, Poga, and Polygonanthus) i is baee he ii vaseclaeieed one Fe aR E bretocarpus has multiple vascular pieds in vin raphe of the mature seed (see Fig. 42). This vas- cular condition in Combretocarpus is probably derived from the condition seen in the three oth- er genera by suppression of vascular extension into the integument, because Combretocarpus has the thin integument that eventually becomes the one cell-layered seed coat at maturity. In this connection, the thick mature seed coat or testa is probably primitive (symplesiomorphous) to the thin, one cell-layered mature seed coat. Com- pared with Poga, Anisophyllea lacks a hypo- oga; Innerlayerand the hypodermal tissue ofthe mul- tiple outer layer. Corner (1976, vol. 1: 57) has considered the limitation of a multiple mechan- ical tissue (probably like that of Poga) into one cell-layered as one of specialization trends of seed coat. Following Corner, we may be able to pos- tulate that the successive or simultaneous re- duction of the multiple inner layer and the hypo- dermal tissue of the multiple outer layer had occurred in the seed coat evolution of Aniso- phylleaceae so that only the epidermis was per- sistent, as in Combretocarpus. Although we did not observe the anatomy of the testa of Polygo- nanthus, it was confirmed that the (outer) integ- ument shows a seconda dition that is cle ombretocarpus. Therefore it seems very likely that Polygonanthus would form a mature seed with as thick a testa as in Poga. 24 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE 3. Evolutionary trend of karyological and some embryological characters in Anisophylleaceae. Plesio- Characters morphy Apomorphy 1. pei of integu- 2 l 2. Pio nuclei Not fused Fused 3. “Outer layer" of the Thick Thin, 1 cell “thick” seed coat thick (thickness) 4. Chromosome number =7 n=8 5. Pattern of embryo sac Polygonum Allium type formation type 6. “Inner layer" of the Present Absent mature seed coat 7. Accumulation of Absent Present starch grains in de- veloping embryo 8. Present Absent sac Vasculature of integu- ments The accumulation of starch grains in the de- veloping embryo sac (Combretocarpus) is known to occur in many unrelated families of dicoty- ledons (see Davis, 1966). Even within a family, however, their occurrence is in general restricted y by particular groups in many unrelated families, probably because of the necessity of dif- ferent metabolic activity during megagameto- genesis. Embryos with moderately developed cotyle- dons are almost universal among dicotyledons and therefore must be primary. On the contrary, because of their restricted occurrence (see Natesh & Ram, 1984), embryos lacking cotyledons seem to be secondary in the evolutionary trend. In this respect, it might be interpreted that an embryo with small cotyledons (Combretocarpus) seems less specialized than that which lacks cotyledons, or at the most has rudimentary ones (Aniso- phyllea and Poga). Such differences in the degree of size reduction of cotyledons, however, may be share a conspicuous hypocotyl, a truly significant and unusual feature It may be difficult to determine whether the chromosome base number of Anisophylleaceae is n = 7 or 8. Noticeably n = 8 occurs only in [Vor. 74 Combretocarpus, a genus that is furnished with many advanced and fewer primitive character states as discussed above. In contrast, n — 7 is shared by Anisophyllea, Poga, and Polygonan- thus, all of which — particularly the latter two— retain a combination of primitive character states. Thus it seems likely that n = 7 is the base number of the family, and n = 8 the derived. The fusion of tapetal nuclei (Anisophyllea and t O — (Poga and Polygonanthus), as discussed arlier. These nuclei also remain distinct in most Made, (Tobe & Raven, 1983). The results of our evaluation of these character states are summarized in Table 3. On this basis, we constructed a cladogram illustrating the evo- lutionary interrelationships of the genera (Fig. 71). The cladogram indicates that the proto-Ani- sophylleaceae, a hypothetical ancestor of the family, had nearly all of the embryological fea- tures that are presently retained by Poga and tually uncertain in Poga an tegmy, vascularized integument, thick seed coat consisting ofa multiple inner layer and a multiple outer layer, no starch grain accumulation during megagametogenesis, and non-fused tapetal nu- clei. An tral luti y li toh diverged into two main branches: one leading to Combretocarpus and Anisophyllea, and the other leading to Poga and Polygonanthus. In the for- mer branch, the ovule became unitegmic; tapetal nuclei fusion has been generalized, and the thick- ness of the multiple outer layer was reduced into one cell layer (i.e., the outer epidermis); all three characters are synapomorphies common to Combretocarpus and Anisophyllea. This branch further diverged into two branchlets: one leading to Combretocarpus, and the other leading to Ani- sophyllea. In the branchlet leading to Combre- tocarpus, chromosome base number changed to n = 8; the Allium type embryo sac and unitegmy were derived; complete reduction of seed coat tissue except for the outer epidermis (i.e., of both a multiple inner layer and the hypodermal tissue of the original thick seed coat) and reduction of mentary vasculature occurred nearly si- e. B ge B O ri gam respect to embryolog ing change has pique in the other branchlet 1987] e ° SY S o n ` C „O "2 S) d o [— 5 [ . 1- pe 4 e 3 =C 2 F 1 FiGuRE 71. A cladogram illustrating postulated evolutionary interrelationships of the genera of Aniso- phylleaceae. Corresponding characters numbered 1 to 8 and their evolutionary states are shown in Table 3. Shaded rectangles indicate apomorphies. *Character state uncertain in Polygonanthus; "Character state un- certain in Poga. leading to Anisophyllea, or in the other main evolutionary line leading to Poga and Polygo- nanthus. Consequently, Combretocarpus differs sub- stantially from the three other genera and is ap- parently the most specialized member of the family. In contrast, most plesiomorphies are re- ferred to Poga and Polygonanthus, which both retain a combination of primitive embryological character states common to the ancestor of Ani- sophylleaceae. Anisophyllea stands in a more or less intermediate position between Combreto- carpus, on the one hand, and the group com- prising Poga and Polygonanthus, on the other. TOBE & RAVEN—ANISOPHYLLEACEAE 25 Indeed Anisophyllea shares most of its archaic features with Poga and Polygonanthus but shares its apomorphies, including unitegmy and tapetal nuclear fusion, with Combretocarpus. Phylogenetic interpretations of the infrafamil- ial relationships as discussed above are primar- ily based on embryological character state eval- uation and chromosome number. The cladogram shown in Figure 71 is to be regarded as limited in this respect and provisional. The evolutionary trend in certain characters (such as chromosome number and seed coat structure) might be the opposite of what we have proposed, and certain characters (such as tapetal nuclear condition) might be of much less fundamental significance than others. Earlier studies on Anisophylleaceae he whether or not Polygonanthus was closely related to the other three genera (for instance, Pires & Rodrigues, 1971) or whether or not Anisophyl- leaceae (*Anisophylleae") should be excluded from Rhizophoraceae (for instance, Van Vliet, 1976). We now regard both of these questions as definitively solved and hope that our phyloge- netic diagram will stimulate further research on the family from various other points of view and will thus be improved as a result of these inves- tigations. LITERATURE CITED AIRY SHAW, H. K. 1973. A Dictionary of the Flow- ering Plants and Ferns. 8th edition. Cambridge Univ. Press, 7 mbridge BAEHNI, C. & P. DANSEREAU. 1939. Polygonanthus, genre de Tire. ce Bull. Soc. Bot. France 86: 183-186. BAILLON, H. E. 1877. Histoire des Plantes. 6: 292- 294. BEHNKE, H.-D. 1982. Sieve-element plastids of Cy- rillaceae, Erythroxylaceae and Rhizophoraceae: tids. Plant Dub Evol. 141: 31—39 ' 1985]. Ultrastructure of sieve-element plastids of Myrtales and allied groups. Ann. Mis ouri 824-831. Bot. Gard. 71: BENTHAM, G. & J. D. HOOKER. 1865. Genera Plan- tarum 1(2): 677-683. he ovule. Pp. 123-157 in B. M. ology of Angiosperms. Further investigations on the em- bryology of viviparous seeds. Proc. Linn. Soc. New 1948. Aluminum i in the plant world. L Kew Bull. 1948: 173-183. 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN Cook, M. T. 1907. The embryology of dr s mangle. xc Torrey Bot. Club 34: 271-277 -— E. J. 1940. Wayside Trees of Malaya, me 1. yes Print., Singa n "1976. The Seeds of Dicotyledons, Volumes 1 & 2. Cambridge Univ. Press, Cambridge CRoizaT, L. M. C. 1939. Polygonantheae (Olaca- ceae). Bull. Soc. Bot. France 86: 5-7. 43. Polygonanthaceae. Cact. Succ. J. (Los 64. . 1981. An Integrated System of Clas- sification of Flowering Plants. Columbia Univ. Press, New York. 983. Some jc era in the dicotyle- Nord. J. Bot. 3: 75-83. DAHLGREN, R. 1983. Tim aspects of angiosperm evolution and macrosystematics. Nord. J. Bot. 3: 119-149. —— & R. F. THORNE. 1984. [1985]. The order Myrtales: circumscription, variation, and relation- ships. Ann. Missouri Bot. Gard. 71: 633—699. Davis, G. L. 1966. Systematic Embryology of the Angiosperms. John Wiley & Sons, New York. Ducks, A. 1932. Polygonanthus. Notizbl. Bot. Gart. Berlin-Dahlem 11: 345-346. 933. Polygonanthus. Arch. Jard. Bot. Rio de Janeiro 6: 62—63. ELDREDGE, N. & J. CRACRAFT. 1980. Phylogenetic Patterns and the Evolutionary Process. Columbia Univ. Press, New York. ERDTMAN, G. 1966. Pollen Morphology and Plant Taxonomy. Angiosperms. Hafner Publishing Co., New York. GEH, S. Y. RH. KENG. 1974. Morphological studies on some inland Rhizophoraceae. Gard. Bull. Straits Settlem. 28: 183-220. Hou, D. 1958. Rhizophoraceae. /n Van Steenis (ed- itor), Flora Malesiana 5: 429-493. JUNCOSA, A. M 82. Developmental morphology of the embryo and seedling in Rhizophora mangle L. (Rhizophoraceae). Amer. J. Bot. 69: 1599-1611. 984a. Embryogenesis and seedling devel- opment in Cassipourea elliptica (Sw.) Poir. (Rhi- zophoraceae). Amer. J. Bot. 71: Embryogenesis and developmental morphology of the seedling in Bruguiera exaris- tata rin Hou (Rhizophoraceae). Amer. J. Bot. 71: 91. ud G. 1891. Über die Mangrove-Vegetation im malaysische Archipel. Eine morphologische- biologische Studie. Biblioth. Bot. (Stuttgart) 22: 11-18, 31-41. Au^ tud J. C. 1940. Polygonanthus punctulatus. Reunion Sul-Amer. Bot. 944. Nota prévia. Rodriguésia 8: 65. [Vor. 74 SUEACHEN B. F. & R. B. MILLER. 1980. A chemical E Int. Assoc. Wood Anatomist Bull n.S., 109. Manco, H. F. 1935. Systematic anatomy of the woods of the Rhizophoraceae. Trop. Woods 44: 1-20. MAURITZON, J. 39. Contributions to the embryol- ogy ofth d d Myrtales. Acta Univ Lund. 35: 1-121. E H. 1964. Rhizophoraceae. Pp. 357-359 . Melchior (editor), A. Engler's Syllabus der Plarzenfamilien. II. Gebrüder Borntraeger, Ber- Ste S. & M. A. RAU. 1984. The embryo. P snc in B. M. Johri (editor), p of A sperms. Springer-Verlag, Ber Nice F. 6. Anatomie der jo M Samen. Gebrüder Brontraeger, Berlin. Pires, J. H. & W. A. RODRIGUES. 19 Notas sóbres os géneros Polygonanthus e Anisophyllea. Acta Amazonica 1(2): 7-15. 1922. The Flora of the Malay Pen- insula, Volume 1. L. Leeve & Co., Londo SCHIMPER, A. F. W. 1893. Rhizophoraceae. In Engler & K. Prantl (editors), Die Natürlichen deer E Wilhelm Engelmann, Leipzig 3(7): 4 SCHMID, 1 nn Descriptors used to indicate abun- dance and frequency in ecology and systematics. Taxon 31: 89-94. . TURNER. 1977. Contrad 70, an ef- fective softener of herbarium material for anatom- ical study. Taxon 26: 551-552. TAKHTAJAN, A. L. 1980. Outline of the classification of flowering plants ee Bot. Rev. (Lancaster) 46: 225-3 THORNE, R. F. 1983. Proposed new realignments in 117. n embryological analysis of the Myrtales: its definition and char- acteristics. Ann. Missouri Bot. Gard. 70: 71-94. & 1 The number of cells in the pollen of Melastomataceae (Myrtales). Bot. Mag. Tokyo 97: 131-136. VAN VLIET, G. J.C. M. 1976. Wood anatomy of the Rhizophoraceae. Leiden Bot. Ser. 3: 20-75. AAS. 1984.[1985]. Wood anatomy and classification of the Myrtales. Ann. Missouri Bot. Gard. 71: 783-800 70. The Structure and Utilization all Ltd., London. en ological studies i in some ` Combretaceae. Bot. Not 125: 161-179. TRAPLINERS IN THE TREES: HUMMINGBIRD POLLINATION OF ERYTHRINA SECT. ERYTHRINA (LEGUMINOSAE: PAPILIONOIDEAE)! DaviD A. NEILL? ABSTRACT Erythrina sect. Erythrina Mdh re) specie tributed principally in Mesoam nators— were observed at 17 ciun P 13 spe pollinators were all "high- reward traplining" ada behavior, including in arduus short- € d es t defense by a hummingbird of a ui tree is piri eui The traplining hummii n flo ral visit tors—includin s of hummingbird- viae i -= and shrubs, dis- g nectar - sins southern Mexi xico an pos Rica e high-reward traplining systems involving hermit hummingbirds and understory plants such as Heliconia (Musa- ae). Erythrina L. (Leguminosae: Papilionoideae) comprises about 112 species distributed throughout the tropical regions of the world and extending into warm temperate areas in both the northern and southern hemispheres (Krukoff & Barneby, 1974; Neill, in press). Most species are trees or shrubs, but about 10 species occurring in climates with pronounced dry and/or cool sea- sons are perennial herbs with large, woody root- stocks. Erythrina species occur in a very wide variety of habitats, from lowland tropical rain forests to very arid subtropical deserts to high- land coniferous forests above 2,500 m in ele- vation. Erythrina species have red or orange flowers and copious nectar and are adapted to pollina- tion by nectarivorous birds. Two distinct syn- dromes of ornithophily are evident. All 42 of the World species and 15 of the 70 New World ine birds cannot hover efficiently or for any length of time, and the inflorescences of passerine-pol- linated Erythrina are oriented in such a way that the birds can perch while feeding on nectar from the flowers. The corolla standard is usually broad and the flowers are open, with exposed repro- PER S parts. Pollen is deposited on the feeding bird’s breast (Cruden & Toledo, 1977). In con- trast, 55 of the New World species of Erythrina (nearly half the genus) are pollinated by hum- mingbirds (Trochilidae), which occur only in the “pseudotube,” concealing the wing and keel pet- als as well as the reproductive parts. The flower resembles the tubular corollas of many gamopet- alous hummingbird-pollinated plants, but in Er- ythrina the pseudotube is not sealed on the ven- tral side where the margins of the corolla standard meet. The inflorescence axis of the humming- ' Discussions with Peter Feinsinger, Peter Raven, and Héctor Hernandez were helpful in the planning and execution of this research and in the interpretation of the results. Frederick Neill’s Tae and companionship all during the fieldwork in Mexico was invaluable. Francisco Guerrero and Rom Velásquez, biologists at El Sumidero National Park, helped with the fieldwork there. Alina Chacón, Mary Merello, Gloria Hoch, and Donna Krausz assisted with the preparation of the manuscript and figu and Missouri Botanical Garden. The research was funded by grants from the National Science Foundation (DEB 81-20386), the Jesse Noyes Smith Foundation and the Organization for Tropical Studies for fieldwork in Costa Rica, and Elizabeth Neill ? Missouri Botanical Garden, St. Louis, Missouri 63166. ANN. Missouni Bor. GARD. 74: 27-41. 1987. 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN bird-pollinated species is erect and the flowers are oriented outward, providing no perch for the hovering hummingbirds. The American | are included in six different sections of subg. Er- ythrina; these I believe to have been derived from passerine-pollinated groups by convergent evo- lution in several independent lineages. The larg- est by far of the hummingbird-pollinated groups is sect. Erythrina (36 species) with its center of diversity in northern Central ; outh- ern Mexico (Neill, in press). Most species a sect. Erythrina are canopy or subcanopy trees, ranging in height from 5 m in semiarid scrub to 25 m or more in lowland rain forest or cloud forest. A few species inhabit the understory and light gaps of wet forests. Many have restricted ranges and are edaphic specialists; an example is E. tux- tlana, which grows only on limestone outcrops in lowland wet forests in southeastern Mexico. A number of field studies of pollination in Er- ythrina have been conducted in recent years, in- cluding observations of passerine- pira as well as ł 1974; Toledo & ] Hernández, 1979; ED à Toledo, 1979, 1982; Cruden & Toledo, 1977; Feinsinger et al., 1979; Morton, 1979; Steiner, 1979; Guillarmod et al., 1979). An extensive sur- vey of hummingbird pollination of Erythrina tree pecies, however, was lacking prior to the re- search reported here. 5 Ree | 11; 4 2 9 HUMMINGBIRD FORAGING BEHAVIOR: A REVIEW Hummingbird species differ in size, bill mor- phology, and foraging behavior as do the floral morphology and flowering patterns of the plant species they visit (Feinsinger & Colwell, 1978). The two principal behavioral types are territo- rialists, which feed at large patches of flowers, defen ing the flowers against usurpers; and trap- relatively small, short- billed birds, and “hig reward" trapliners, birds with relatively i. bodies, high energetic requirements, and long or curved bills. High-reward trapliners are the most "specialized" of hummingbirds in the sense that they have the highest fidelity to particular plant species. These are the only birds capable of pol- [VoL. 74 linating plants with iene pula flowers, and they ceto plant species with low population densities. The most well-studied of the high-reward trapliners are the hermit hummingbirds (Phaethorninae), which forage principally in lowland and mid-elevation wet forest understories on widely spaced, nectar- rich herbs and shrubs. Stiles (1978, 1981) noted that very few hum- mingbird-pollinated plants are canopy trees; in contrast, many ornithophilous plants of the Old World, pollinated by passerine birds, are trees. Stiles reasoned that this difference is related to the social systems of the two bird groups. Pas- lly forage in large flocks, whereas hummingbirds are virtually always sol- itary. A large concentration of flowers on a large tree would be parceled up into feeding territories pollination. A large flock of pass contrast, could quickly exhaust the resources of even a large tree, and the flock would be com- pelled to move on to the next tree, thus effecting cross-pollination. Erythrina is an exception to the general paucity of hummingbird-pollinated canopy or subcan- opy trees. It should be instructive to determine by observations in the field whether Stiles’s pre- diction holds true for hummingbird-pollinated Erythrina. If hummingbirds parcel the crown of a tree into several feeding territories, they may reduce intertree pollen flow; alternatively, hum- mingbird-pollinated Erythrina trees may possess adaptations that reduce territorial behavior and promote intertree movement of the humming- bird pollen vectors. In the present study of hummingbird polli- nation in natural populations of trees of sect. Erythrina, several questions were addressed: Do the hummingbirds that pollinate Erythrina be- have as territorialists or as trapliners? How spe- cialized are Erythrina pollinators? Another goal was to assess the potential for pollen transfer between different species of Erythrina. Experi- mental hybridization studies (Neill, in press) in- dicate that species of sect. Erythrina are highly interfertile, but unless the hummingbird pollen vectors carry pollen from one species to another, hybridization will not take place under natural conditions. This assessment required informa- tion about the flight and foraging patterns of the birds, and whether the Erythrina species shared the same pollinators or had different, host-spe- cific pollinators. 1987] MATERIALS AND METHODS FLORAL BIOLOGY AND BREEDING SYSTEMS Observations on phenology and other details of Erythrina floral biology, as well as experi- mental studies of genetic self-incompatibility, were made on wild populations in Mesoamerica and on cultivated trees in Hawaii. The experi- mental methods for the breeding system studies are described by Neill (in press). OBSERVATIONS OF FLORAL VISITORS Floral visitors to 17 populations of 13 species of hummingbird-pollinated Erythrina trees were observed in Mesoamerica. These included four wet season-flowering species in Costa Rica, July— September 1981, and nine dry season-flowering species in southern Mexico, January—April 1983. All but one of the species is in sect. Erythrina; the exception is E. gibbosa from Costa Rica, in the monotypic sect. Gibbosae. Flowering and pollination patterns in this species are very sim- ilar to those in sect. Erythrina. For the 13 species a total of 195 person-hours of observation was conducted. For a population the number of observation hours ranged from 2.5 to 45.5. Most of the observations were made between dawn and 12 P.M., because floral visitor activity is usually greater in the early morning and drops substantially by noon. Some obser- vations were conducted in the late afternoon Saen avian floral after the midday lul At the beginning y each observation day I counted the number of open flowers on each tree. For each bird visit to a tree crown, I recorded the time, duration of visit, number of flowers probed, and direction of arrival and departure. I judged qualitatively the frequency of the visi- tor's contact with the reproductive parts of the flower, as a measure of potential pollination ef- ficacy. Many of the non-pollinating passerine bird visitors actually destroyed or removed a consid- erable number of flowers daily, reducing the number available to subsequent visitors to the tree. This activity was recorded and entered into the daily flower censuses. NECTAR PRODUCTION I sampled daily nectar production in four species in sect. Erythrina: E. globocalyx and E. NEILL— ERYTHRINA POLLINATION 29 costaricensis inhabiting wet forests in Costa Rica, and E. chiapasana and E. goldmanii in dry for- ests of Chiapas, Mexico: Because the inflores- cences in the tree crowns were difficult to reach from the ground, repeated sampling of nectar secretion of individual flowers over the course of a day was not possible; only a day’s total pro- duction was sampled. In the late afternoon, flowers due to open the following day were bagged with mosquito net- ting. These *'first-day" flowers were removed at about 4 P.M. the following day, and nectar vol- ume from each flower was measured by repeated probing with a 10- or 25-ml calibrated micro- pipette. Ns production for *second-day" flowers of . goldmanii and E. chiapasana was estimated in the following manner: inflorescences were bagged with mosquito netting in the late after- noon as for first-day flowers, and nectar was al- lowed to accumulate in the isolated flowers for two days. Total nectar accumulation was mea- sured at 4 P.M. on the second day. The mean first-day accumulation of nectar subtracted from the mean 2-day accumulation yields the esti- mated second-day nectar production. (This method assumes that nectar removal does not influence secretion.) All bagged flowers fell off by the morning of the third day. Sugar concentration expressed as percent su- crose equivalence (Bolten et al., 1980) was mea- sured for each flower or for the pooled nectar of several flowers using an American Optical model 10431 temperature-compensated hand refrac- tometer. With the figures for nectar volume and sugar content, the mean caloric value of the nec- tar per flower was calculated for each population sampled. RESULTS FLORAL BIOLOGY AND BREEDING SYSTEMS In most passerine-pollinated species of Ery- thrina the stamens and stigma are well separated at anthesis and the flowers are homogamous, i.e., pollen is released from the anthers and the stigma close proximity, tightly enclosed in the floral pseudotube, and the flowers are protandrous. On the first day of anthesis the staminal fila- 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 1. Observations of avian floral visitors to Erythrina tree species. Nectar Thieves & Robbers Humming- Species Locality' Trees? Hours? Pollinators birds Passerines 1. E. ameri- Mexico: Oaxaca 4 9 Heliomaster — Icterus galbula cana Coatlán constantii 2. E. berte- Mexico: Chiapas 5 3.5 Anthraco- Anthraco- Icterus macu- roana x alenque thorax pre- thorax pre- lialatus E. folkersii tii vostii Pheucticus lu- dovicianus 3. E. bere- Mexico: Veracruz 2 2 Eugenes ful. | — — nices Tequila gens 4. E. chiapa- Mexico: Chiapas 8 15.5 Heliomaster — Icterus wagleri sana El Sumidero constantii 5. E. chiapa- | Mexico: Chiapas 6 18 Eugenes ful- Hylocharis Diglossa barit- sana Teopisca gens leucotis ula Icterus galbula 6. E. coch- Costa Rica: Heredia 6 20 Heliomaster Chalybura — leata La Selva longirostris urochrysia 7. E. costari- Costa Rica: Puntarenas 1 5 Heliomaster Phaeochroa — censis San Vito de Java longirostris cuvierii 8. E. gibbosa Costa Rica: Alajuela 2 11 Phaethornis Lampornis — Monteverde guy hemileucus 9. E. globo- Costa Rica: San Jose 5 8.5 Eugenes ful- — — calyx Las Nubes gens Campylop- terus hemi- leucurus 10. E. gold- Mexico: Chiapas 12 45.5 Heliomaster — Icterus gularis manii El Sumidero constantii I. pectoralis I. wagleri 11. E. folkersii Mexico: Veracruz 2 3.5 Phaethornis — — Los Tuxtlas supercilio- Sus Campylop- terus hemi- C. curvipennis 12. E. folkersii Mexico: Chiapas 2 45 Phaethornis Amazilia tza- Icterus galbula Palenque supercilio- I. prostheme- SUS las 13. E. lanata Mexico: Oaxaca 2 4.5 Heliomaster — — erto Escondido constantii 14. E. lanata Mexico: Jalisco 2 7.0 | Heliomaster — Cassiculus Chamela constantii melanicte- rus 15. E. pudica Mexico: Chiapas 8 16.0 Heliomaster — Icterus gularis El Aguacero constantii 16. E. tux- Mexico: Chiapas 3 15.0 Heliomaster Amazilia Coereba flave- tlana Malpaso constantii cyanoceph- Anthraco- ala Cyanerpes lu- thorax pre- A. tzacatl cidus vostii Icterus gradu- acauda 1987] NEILL—ERYTHRINA POLLINATION 31 TABLE 1. Continued. Nectar Thieves & Robbers Humming- Species Locality! Trees Hours? Pollinators irds Passerines Eugenes ful- Pheucticus lu- gens dovicianus 17. E. tux- Mexico: Veracruz l 5.5 | Campylop- Eupherusa — tlana Uxpanapa terus curvi- imi pennis l asa we locality and voucher data for each observation are listed in Appendix. 2 Number of indi rved. vidual trees obse 2 Paeon Tua of observation ments are fully grown and pollen is released from the stamens situated near the apex of the corolla within the pseudotube. At this stage the style is shorter than the stamens and the stigma is not receptive. The style and ovary continue to elon- gate during the night after the first day of flow- ering. By the second day, when most ofthe pollen has been removed by floral visitors, the stigma, now receptive and with a sticky exudate on its surface, is held a few mm beyond the anthers, just inside the mouth of the pseudotube at the apex of the corolla. Each flower, then, is func- tionally male on the first day and functionally female on the second. Unpollinated flowers usu- ally abort after the second day but sometimes remain for a third or fourth day before aborting; the stigma appears to remain receptive during this time. The inflorescence of sect. Erythrina is a pseu- doraceme with fascicles of three flowers each ar- cence, and usually one or two fascicles of flowers open each day. An individual inflores- cence, with 30-50 fascicles, blooms for two or three weeks. An inflorescence in “full bloom" is composed of three to nine functionally female flowers at the bottom (in one to three fascicles), three to six functionally male flowers in the fas- cicles just above the females, and above them floral buds at progressively younger develop- mental stages. Inflorescence development in an individual Erythrina tree is staggered, so a tree often blooms for two to three months or more. Blooming among trees in a population is also staggered, so a pop- ulation is often in bloom for four to five months annually or even longer. Some species remain in bloom for a shorter period, one to two months within a population. Detailed, multi-year phe- nological data for any particular site, however, is not available. Most species of sect. Erythrina flower during the dry season, from January to May in Me- soamerica, and are leafless when in flower. Leaves flush, in general, after flowering has ceased and while fruits are developing, prior to or just after the onset of the rainy season in May or June. Some species flower during the rainy season, June to October. These species also usually shed all their leaves before flowering and flush a new set of leaves as flowering ceases. This behavior is unusual: few other tree taxa in the Neotropics, especially in very wet forests, shed leaves during the rainier portion of ai year and retain them during the drier portio The data presently E nsi: suggest that all Erythrina species are genetically self-compatible (Neill, in press). The fitness of progeny resulting from self-pollination is significantly lower than that of progeny resulting from outcrossing; out- crossing appears to be predominant in the breed- ing systems of Erythrina. Some seed set from geitonogamous pollinations and even from oc- casional autogamy, which probably occurs in natural populations. FLORAL VISITORS The avian visitors to the flowers of most ob- served Erythrina por ions include d h mingbird and passerine “‘illegi on, gi m linators (Table 1). Pollination records for each Erythrina species are summarized in Table 2, which encompasses prior reports on humming- bird-pollinated tree species: Erythrina lanceo- 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 TABLE 2. Species of hummingbirds observed as legitimate pollinators on 15 species of Erythrina. Abbre- viations in parentheses refer to the countries in which species were observed (M = Mexico, C = Costa Rica, and T = Trinidad). Hubiniasbirds Erythrina Species _ " G S 6 š € 2 “< _ = < 6 3 4 = 3 RE š s S ç B š Bo Q S a uw S S S 8 š gs 8 3 S BB E Š 8 S & ÈA &8 S S S E 8 § Š S S š 8 3S E $S B E S E i $ š 3 & & S S $ s Š l š & g 3S BES Š uj nj bJ bj ud us yy ye ye ye S SQ OR og Heliomaster constantii + + + + + + H. longirostri + + + + Eugenes fulgens + + + + + pylopte hemileucurus + + + C. curvipennis + + Anthracothorax prevostii + + + Phaethornis guy + + Ph. superciliosus + Glaucis hirsuta + lata in Costa Rica (P. Feinsinger, pers. comm.) and E. pallida in Trinidad and Tobago (Fein- singer et al., 1979) For this discussion I have adopted a modified version of Inouye’s (1980) terminology for floral larceny: “nectar robbers” make a hole or oth- erwise damage floral tissue to gain access to the nectar, while cludes pollination. The distinction between nec- tar thieves and robbers is important because rob- bers may damage the ovary or stylar tissue and often destroy or remove the entire flower, and thus may have a much greater effect in reducing the reproductive potential of the plant than do nectar thieves. Billl bt 4 : :1 - from Ridgway, 1911) of the hummingbird species observed as pollinators and illegitimate visitors to Erythrina trees are compared in Figure 1. Legitimate pol- linators all have bills longer than 28 mm, where- as nectar thieves and robbers, with one excep- tion, have bills shorter than 22 mm POLLINATORS Nine species of hummingbird were observed as legitimate pollinators of the 15 species of Er- ythrina trees. All ofthe hummingbird pollinators may be characterized as long-billed, high-reward trapliners, although some behaved as territori- alists or even nectar thieves on certain occasions, as detailed below. I observed two species of hermit humming- birds (Phaethorninae) pollinating Erythrina: Phaethornis guy and P. superciliosus. A third hermit species, G/aucis hirsuta, was reported as a pollinator of E. pallida in Trinidad by Fein- singer et al. (1979). The hermits visited princi- pally the smaller understory species of Erythrina but rarely were in the forest canopy. Their for- aging behavior at understory Erythrina is similar to that documented for understory herbs such as Heliconia (Stiles, 1975) or shrubs such as Aphe- landra (McDade, 1984). The remaining six Erythrina pollinators are non-hermits (Trochilinae); all are long-billed, high-reward trapliners which forage like hermits, but usually in the forest canopy or open areas rather than in the understory. Heliomaster con- stantii is the principal or sole known pollinator of at least six Erythrina species in the dry forests on both the Pacific and Caribbean slopes of Me- NEILL— ERYTHRINA POLLINATION ERYTHRINA FLORAL VISITORS: HUMMINGBIRDS s NS NN. \\ \ \ x 30_N NN NY w ail S A N AAAA s2 \ \ \ \ \ YX all AN - \\\\\ \\V all NN N N N N \ N NN N N N \ y>FFFOOQOA Nectar Thieves & Robbers FIGURE 1. Bill lengths of Bop éd. visitors to Erythrina trees, including pollinators and legitimate visitors. Ph gu = Phaethornis guy; Ph s Heliomaster longirostris; Gl hi = = Phaethornis superciliosus, He co = Glaucis hirsuta; Ca he = Campylopterus hemileucurus; Eu fu = Heliomaster constantii; He lo E genes fulgens: Ca cu = grey pif curvipennis, An pr = Anthrocothorax prevostii, Ph cu = Phaeochroa cuvieri Amazilia tzacatl, Am ba = Heliothryx baro soamerica, and from E. pallida in Trinidad (Feinsinger et al., 1979). Eugenes fulgens is found mostly in the highlands of Mesoamerica above 1,500 m. It is the principal pollinator of at least four highland species of Erythrina, and I also observed it below 1,000 m pollinating E. tux- tlana in southern Mexico. Campylopterus hemi- leucurus 1s a bird of wet forests from near sea level to 1,800 m and has been observed polli- nating three Erythrina species in such habitats. Campylopterus curvipennis was observed as a pollinator of two Erythrina species in low- to mid-elevation wet forests of southern Mexico. Anthracothorax prevostii was the least consis- tent Erythrina pollinator. This bird of low- to = Amazilia cyanocephala; Hy le = Hylocharis leucotis; Eu ex = [eredi eximia; He mid-elevation humid fi as a legitimate pollinator and contacted the re- productive parts of Erythrina like the other long- billed trapliners. More frequently, however, An- thracothorax was a nectar thief, as described below. The species of hummingbirds that pollinate sect. Erythrina all behave in a similar manner when feeding at the flowers. The hummingbird first approaches the inflorescence and hovers to align its bill precisely with the axis of the first flower it is to visit (Fig. 2). It inserts its bill at the apex of the flower; at full penetration the reproductive parts of the flower always contact the bird’s throat, upper chest, or the base of the 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURES 2-5. liomaster constantii a self to feed at flowers of e lanata; dry forest near the Pacific coast, Pochutla, Oaxaca, Mexico. Bill i pine-oak forest, Teopisca. Chia upper throat. — 4-5. inserted at apex of corolla. — 3. pas, Mexico. At full insertion of the bill, pollen is deposited at base of bill or on Hylocharis wasps a short-billed nectar thief, approaching and fora Hummingbird pollinators and nectar thieves of Erythrina sect. Erythrina.—2. Pollinator He- Eugenes fulgens pollinating E. chiapasana; highland ging at flowers of E chiapasana; same 2 sss as in Figure 3. The bill is inserted into the mouth of the calyx or the slit of the corolla **pseudotube" to obtain nectar; reproductive parts of the flower are not contacted. bill (Fig. 3). The bird remains hovering at this position for up to five seconds, then withdraws and moves to another flower in the inflorescence or to another inflorescence. The pollination records summarized in Table 2 indicate that there is no species-specific, one- to-one relationship of Erythrina species and pecies. The ecological and geo- birds have been observed at several species of Erythrina at different localities in the birds’ range, and more than one pollinator has been recorded for many of the Erythrina species. The bird species are quite similar to one another behav- iorally and morphologically, and the tree species are also quite similar to one another in terms of floral morphology, flowering behavior, and nec- tar rewards (the last is discussed below). Ery- thrina sect. Erythrina and related groups are ev- idently adapted to pollination by the high-reward trapliner guild of hummingbirds as a whole. This, however, is a small subset of all hummingbirds. 1987] In Mesoamerica, there are very few additional hummingbird species with the appropriate mor- phology and behavior for Erythrina tree polli- nation, other than the nine species listed in Table 2. (Doryfera ludoviciae in the highlands of Costa Rica and Panama may be the only high-reward trapliner in the region not reported as an Ery- thrina pollinator [P. Feinsinger, pers. comm.].) the local community level, often only a single high-reward traplining hummingbird is present so that many populations of Erythrina are pollinated by a single bird species. In the lowland dry forests of Pacific Mesoamerica, He- liomaster constantii is the only appropriate hum- mingbird present, so it is undoubtedly the sole pollinator of the Erythrina tree species restricted to Pacific dry forests. The hummingbirds are not strict specialists on Erythrina. Well-studied species such as Eugenes fulgens have been reported visiting a number of other plant species. The two Heliomaster species may be more specialized as Erythrina foragers than are the other hummingbird genera. During the course of his extensive community-level studies of hummingbirds, P. Feinsinger (pers. comm.) observed Heliomaster longirostris on Trinidad to feed only at Erythrina pallida and at the apocynaceous vine Mandevilla hirsuta; while at Monteverde, Costa Rica, Heliomaster constantii visited exclusively different species of the same two plant genera. NECTAR THIEVES The nectar thieves of Erythrina sect. Erythrina chrysia (Tab le 1). The short bills of these birds preclude them from reaching the floral nectar by inserting their bills at the apex of the corolla. Nectar thieves take advantage ofthe incomplete- ly sealed tube of the Erythrina corolla. They ap- proach the flower from below (Fig. 4) and, often with some struggle and manipulation of their bills and bodies, slip their bills into the proximal end of the ventral slit of the pseudotube formed by the corolla standard, or into the mouth of the calyx (Fig. 5). They are thus able to gain access damaging tivity is concentrated at the base of the corolla, and thieves were never observed to contact the NEILL— ERYTHRINA POLLINATION 35 reproductive parts of the flowers at the apex of the corolla. Unlike the pollinators, thieves fre- quently grasp the flowers with their feet when feeding and clamber over the inflorescence to reach adjacent flowers. They may damage the surface of the corolla somewhat, but they do not appear to damage the ovary itself. Small nectar thieves may lack the power to pierce the thick Erythrina perianth as do the larger-bodied nectar robbers. Most of the hummingbird nectar thieves are small species with bills under 21 mm long and bodies weighing less than 6 g. An exception is Anthracothorax prevostii, whose bill length of 28 mm is within the low end of the range of the legitimate pollinators and whose body size of 10 g is equivalent to that of the pollinators. A. pre- vostii was occasionally seen visiting Erythrina flowers in the manner of a true pollinator, but more often it behaved as a nectar thief in a man- ner similar to the smaller opportunistic birds. NECTAR ROBBERS: HUMMINGBIRDS Nectar-robbing hummingbirds have shorter bills than the pollinators but are larger in body size and more powerful than the nectar thieves. The robbers pierce the calyx or base ofthe corolla with their needle-like bills to gain access to Er- ythrina nectar. I observed the large (12 g) hummingbird Phaeochroa cuvierii repeatedly robbing flowers of a roadside Erythrina costaricensis near San Vito de Java in southern Costa Rica. Hovering beneath or beside the inflorescence, the bird placed the tip of its bill against the fleshy calyx and with three or four sharp thrusts punctured through the calyx to plunder the nectar. Usually the robber hovered while piercing the flower, but sometimes it perched on the inflorescence. Flow- ers strewn on the ground below the tree had up to six puncture holes through the underside of the calyx, indicating that Phaeochroa returned to a single flower several times to drain it of nectar. Most pierced flowers were soon aborted, and some had signs of damage to the ovary caused by the robber's bill. On two successive mornings, a Phaeochroa repeatedly robbed an E. costari- censis tree that was also visited at intervals by a legitimate pollinator, Heliomaster longirostris. When on occasion the two birds arrived to fee at the tree simultaneously, a territorial fight en- ued. In the same region of southern Costa Rica o 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN where I made the foregoing observations of E. costaricensis, Skutch (1971) reported that Heli- omaster longirostris pollinated the flowers of Er- ythrina berteroana while Phaeochroa cuvierii and another short-billed hummer, Heliothryx bar- american rainy season, August-November, while E. berteroana flowers during the dry season, late December to March. The similarity of visitation patterns reported for these two species with dif- ferent flowering phenologies in the same region (they are not strictly sympatric) suggests that to- gether they support the same pollinators and il- legitimate visitors in succession for a consider- able portion of the year NECTAR ROBBERS: PASSERINE BIRDS AND PARROTS All of the | non. -hummingbird visitors to d thrina sect. E erally FROM De reproductive potential of the flowers they visit. These robbers include passer- ine birds in the families Icteridae, Coerebidae, and Fringillidae, and the non-passerine parrot family Psittacidae. Icterids and coerebids are le- gitimate pollinators of some Erythrina species in the Neotropics, but on sect. Erythrina they are strictly parasitic. I observed seven species of orioles (Icterus spp.) robbing eight species of Erythrina in Mexico and the icterid e eines Neige. one Erythrina speci rioles were the most fre- quently ED p all robbers a Expl the most complex behavior to obtain the nectar. Typically, an oriole would pluck a flower with its bill, then hold it against a branch with one foot and jab its bill into the mouth of the calyx to reach the nectar. After plundering the flower, the oriole would drop it and pluck another. The sumed daily by the orioles. Sometimes an oriole would impale a flower on a thorn of an Erythrina tree branch to hold it in place while the oriole imbibed the nectar; the impaled flowers were left hanging on the branch Oriole species vary in their degree of special- ization as nectar feeders (Stiles, 1981). Some ev- idently obtain a high proportion of their caloric requirements from floral nectar, at least during certain seasons of the year. Orioles are important [Vor. 74 legitimate pollinators of some Erythrina species such as the widespread E. fusca (sect. Duchas- saingia) (Morton, 1979) and the central Mexican endemic E. oliviae (sect. Olivianae) (Toledo & Hernandez, 1979). These passerine-pollinated Erythrina species are presumed to represent an ancestral condition with respect to the hum- mingbird-pollinated groups including sect. Ery- thrina. Orioles and other icterids are known to behave as nectar robbers, in a similar manner to their behavior at Erythrina, at other plant species in the Neotropics, including banana (Musa par- adisiaca), which was introduced from the Old World tropics (Skutch, 1954). In the case of Er- ythrina, the evolutionary relationship of the ori- oles’ nectar-robbing behavior of the humming- bird-pollinated species to their legitimate Senay of the putatively is pean pollin n. Did nm switch to Sud c bbs after having evolved nectar-feeding behavior as legitimate pollinators, or was the order reversed? everal species of the honeycreeper family (Coerebidae) are nectar robbers of Erythrina flowers. I observed the flower-piercer Diglossa baritula robbing E. chiapasana in the highlands of southern Mexico by holding the corolla with its specialized hooked upper mandible and pierc- ing it with its lower mandible to extract the nec- tar. Hernández and Toledo (1979) observed sim- ilar behavior by the same bird species at Erythrina leptorhiza, an herbaceous species of highland central Mexico. Two coerebid bird species, the shining hon- eycreeper Cyanerpes lucidus and the bananaquit Coereba flaveola, were nectar robbers of Ery- thrina tuxtlana. These birds sometimes pierced calyces in the manner of Diglossa, and at other times slipped their bills into the calyx mouth without puncturing it, in the manner ofthe short- billed hummingbird nectar thieves. Like the ori- oles, these two honeycreepers that behave as rob- bers of hummingbird-pollinated Erythrina species are also important legitimate pollinators of passerine-pollinated Erythrina including E. poeppigiana in Trinidad (Feinsinger et al., 1979) and E. megistophylla in Ecuador (Steiner, 1979). migrant rose-breasted grosbeak Pheucti- cus ludovicianus (Fringillidae) nectar-robbed a living fencepost row of hybrid Erythrina berte- roana x E. folkersii and natural populations of E. folkersii and E. tuxtlana, all on the Atlantic slope of Chiapas, Mexico. Unlike the other pas- L 1987] NEILL—ERYTHRINA POLLINATION 37 TABLE 3. Daily nectar production in flowers of Erythrina sect. Erythrina. X Sucrose Species Nectar Volume Equivalence X Calories (Locality) X ul + s.d. Wt/Vol per Flower N A. First-day flowers E. costaricensis (San Vito de Java) 36.4 + 14.7 29% 43.8 10 E. globocalyx (Las Nubes) 31.4 + 19.5 22.8% 29.3 10 E. chiapasana (El Sumidero) 29.6 + 12.0 27.3% + 3.9% 33.3 12 E. goldmanii (El Sumidero) 31.9 + 14.6 28.9% + 2.2% 38.2 10 B. Two-day accumulation of nectar E. chiapasana (El Sumidero) 49.8 + 26.1 27.3 56.0 6 Estimated caloric production of econd- flower (B-A): 22.7 E. goldmanii (El Sumidero) 52.3 + 31.1 68.1 9 "wn m caloric production of second-day flower (B-A): 29.9 serine robbers, grosbeaks actually consumed flo- varied relatively little within or among popula- ral tissue as well as nectar. Usually they plucked tions (2396-2996). The calculated mean caloric the flower and either crushed the calyx with their value of the nectar per flower ranged from 29 to bills and dropped the flower or consumed the 43 cal among the sampled populations, with an entire flower. At times grosbeaks merely bit off overall mean of 36 cal. the end of the corolla (and pistil), leaving the Nectar continued to accumulate on the second flower attached with the calyx and corolla stump. day of flowering in the bagged flowers of E. chia- On several occasions I observed the short-billed pasana and E. goldmanii. The estimated caloric hummingbird Amazilia tzacatl follow a foraging production of the second-day flowers (overall grosbeak and insert its bill into the decapitated mean = 27 cal) was somewhat less than in the Erythrina corolla tube to extract the remaining first-day flowers, but this may have been due to nectar. inhibition of production by the accumulation of On a number of occasions I observed parrots large nectar volumes in the protected flowers, in (Psittacidae) consume immature seeds of Ery- the absence of removal by nectarivores. thrina trees, but never the flowers. Skutch (1971), The results of the nectar sampling from the however, reported the orange-chinned parakeet different species were similar enough to allow a Brotogeris jugularis to be an important nectar rough estimate of the daily caloric production robber of Erythrina berter insouthern Costa per flower for hummingbird-pollinated sect. Er- Rica. The parakeets plucked the flowers with their — ytArina in general. For purposes of the discussion bill or feet, bit through the calyx to extract the below, an average caloric production of 35 cal nectar, and dropped the flowers without consum- per flower per day is assumed as an approxi- ing floral tissue. mation. This is somewhat less than the values reported by Stiles (1975) for hermit-pollinated NECTAR PRODUCTION AND CALORIC VALUE Heliconia species (48—141 cal) but is an order of magnitude or greater than the production typical of plants pollinated by short-billed generalist hummingbirds (Feinsinger, 1978). Daily nectar production per flower in the sam- pled populations of Erythrina chiapasana, E. costaricensis, E. globocalyx and E. goldmanii is shown in Table 3. Within populations, the vari- ance in nectar production per flower was high (for example, the range in E. globocalyx was 10— Macmillen and Carpenter (1977) derived a 67 ul). The mean nectar volume for each of the regression equation for the 24-hour energy costs four populations (30-36 ul) was quite similar, of nectar-feeding birds, based on empirical data however. The sugar concentration of the nectar on basal metabolic rates and energetic costs of HUMMINGBIRD ENERGETICS AND NECTAR REWARDS 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN flight for hummingbirds, Hawaiian honeycreep- ers, and African sunbirds. Using this equation, the daily energetic cost for a Heliomaster weigh- ing 8.0 g is calculated to be 10.9 kcal. This is the equivalent of 311 Erythrina flowers at 35 cal/ flower. (Hummingbirds do gain some nourish- ment by consuming arthropods, so the actual daily nectar consumption of an Erythrina pol- linator may be somewhat less.) Erythrina species in sect. Erythrina — even large canopy trees at the peak of their blooming pe- riod — do not generally produce as many as 300 flowers per day. Therefore an individual is prob- ably not “worth” defending as a feeding territory by a hummingbird; several trees must be visited daily to satisfy the bird's energetic requirements. Intertree movement of the foraging birds, and consequent pollen flow between trees, is evi- dently promoted by the limited number of flow- ers produced on an individual tree. In contrast, me extended blooming period of Erythrina and £41 of the nectar resource promotes the high fidelity of visitation exhibited by the traplining pollinators. This syndrome is exem- plified by my data on flowering behavior and pollination observations of populations of sev- eral different species of Erythrina, discussed be- w. Erythrina cochleata. Erythrina cochleata is a 25-m tall canopy tree at La Selva Biological Sta- tion, a tropical wet forest site in Costa Rica. I attempted to locate every reproductively mature individual of this species in an area of about 2 km? at La Selva and found a total of 10 trees. This species is confined to the alluvial terraces of the rivers and major streams, so on a large scale the trees were clumped, but no individual was less than 50 m from its nearest conspecific neighbor. Erythrina cochleata flowers during the wet season, and the population was in flower continuously at least from May through Septem- ber 1981 I observed flower production and floral visi- tors for six days at four different individual trees. The trees averaged 112.5 (range 84-181) open flowers per day. The pattern of floral visitation each day was very consistent. The only pollinator and regular visitor was Heliomaster longirostris. Each morning between 7:00 A.M. and 8:30 A.M. a solitary Heliomaster would arrive, visit as few as four to as many as 45 Erythrina flowers, and depart. These visits were repeated at sporadic intervals during the morning. (Birds were not [Vor. 74 tagged, so subsequent visits may have been made by different individual birds.) On only two occasions was more than one hummingbird seen at a time in an Erythrina cochleata crown, and both times the interloper (once another Heliomaster, once a nectar-thiev- ing Chalybura urochrysia) was chased away by the Heliomaster. Heliomaster longirostris is considered an **un- common" bird at La Selva (Slud, 1960), yet I saw this species every time I looked for it at flowering Erythrina trees. Evidently the small, scattered population of Erythrina cochleata sup- ports the nutritional requirements of, and re- ceives consistent pollination service from, a small population of Heliomaster for a period of several months each year. What the Heliomaster hum- mingbirds do when the trees cease flowering is own. They may migrate to populations of other Erythrina species on the Atlantic slope of Costa Rica, such as E. steyermarkii, that flower during the dry season, or they may forage at other canopy flowers such as Mandevilla spp. vines (besides Erythrina, there are no other hum- mingbird-pollinated canopy-level trees in the re- ion). Erythrina goldmanii. The habitat and pop- ulation structure of Erythrina goldmanii at Can- on del Sumidero National Park in Chiapas, Mex- ico, where I observed this species, are very different from those of E. cochleata at La Selva, but the pollination systems of the two species are quite similar. Erythrina goldmanii is a dry forest species and is rather scrubby, rarely attaining a height of over 6 m. At El Sumidero, on a slope above the semi- arid basin ofthe Rio Grijalva, E. goldmanii forms dense populations of small trees in disturbed sec- ondary forest. I made observations in a 2 ha plot containing 54 plants. Most trees had only one or two inflorescences in bloom, with less than 10 flowers per tree; the largest tree had 36 flowers. In all there were 475 flowers in the 2 ha plot at peak flowering. Heliomaster constantii was the only pollinator and the only hummingbird visitor seen at Ery- thrina goldmanii in over 45 hours of observa- tion. At least three Heliomaster individuals were beyond the boundaries of the plot. Their fidelity to Erythrina was very high: only twice did I see 1987] a Heliomaster visit any other plant, and then only for single floral probes Orioles were Beguen visitors to Erythrina goldmanii, and they destroyed an estimated 21% of the flowers daily in the manner described pre- viously. There may have been competition for nectar resources between the orioles and the liomasters, but I never observed any aggressive interactions between orioles and hummingbirds. pecies of Erythrina sect. Erythrina are usually allopatric, being separated by elevation and hab- itat; but at El Sumidero two species come into contact. Erythrina goldmanii inhabits the dry lower slopes at about 800 m, and E. chiapasana occurs in the moister forest on top of the plateau at 1,100 m. Heliomaster constantii visited Ery- thrina chiapasana just as it did E. goldmanii less in the intermediate zone, on the upper slopes of the El Sumidero escarpment (Neill, in press). The birds evidently do not discriminate among Erythrina species when the species occur together. Heliomaster hummingbirds are cer- tainly the pollen vectors implicated in interspe- cific gene flow between Erythrina species at El Sumidero. Erythrina tuxtlana. One final observation indicates that traplining hummingbirds will sometimes behave as facultative territorialists if they are given the opportunity. I observed floral visitors to a 20 m tall Erythrina tuxtlana in mid- elevation wet forest near Malpaso, Chiapas. The tree had a broad-spreading crown with 1,400 open flowers. According to the estimates of nectar pro- duction in other species, this should have been ofthe nee into bading tert ories and maintained thet hen not feeding, each bird generally perched within its territory, and many aggressive interactions en- sued when one bird crossed into another’s ter- ritory. This was the only instance of consistent within-tree territoriality I observed in any Ery- thrina population. CONCLUSIONS: Is IT COEVOLUTION? The flowers of Erythrina sect. Erythrina pro- vide a rich nectar resource that is fed upon by many species of birds besides the legitimate pol- NEILL—ERYTHRINA POLLINATION 39 padi The pollinators, however, constitute a mall guild of “high-reward traplining" hum- ied about eight species in Mesoamerica. hese are mostly non-hermits of the subfamily Trochilinae. Predominant among these is the ge- nus Heliomaster. The pollinators are highly faithful visitors to Erythrina, which provides them with a consistent nectar resource for long periods. The limited caloric value of nectar pro- duced per tree per day usually precludes the maintenance of permanent feeding territories at a single tree by the hummingbird visitors. The consequent nomadic or **traplining" behavior of the hummingbirds and the dispersal patterns of the pollen they transport among the scattered individual E rythrina trees may be a critical factor summary, is a canopy-level analogue of the high- reward traplining pollination systems of Heli- conia and similar understory plants. In this sense the pollination system of sect. Erythrina, togeth- er with the other hummingbird-pollinated sec- tions of Erythrina trees (sects. Stenotropis, Pseu- do-edules, Gibbosae, and Corallodendra; cf. Neill, in press) may be unique. Hummingbird-polli- nated canopy and subcanopy trees are in them- selves uncommon (Stiles, 1978), and I know of no other genus of canopy trees besides Erythrina that is adapted to pollination by the traplining guild of hummingbirds. To what extent have species of sect. Erythrina and their hummingbird pollinators coevolved? To what extent is this a specialized mutualism? Feinsinger (1983) indicated that a highly Occurrence; although te may oer ona particular plant ost hermit species forage on a number of different plants. Similarly, most hermit-pollinated plants are ser- viced by several species of hermits, although shorter-billed birds are excluded as pollen vec- tors. If one's definition of coevolution requires a high degree of one-to-one species specificity in such mutualistic interactions, then hermits and hermit-pollinated plants cannot be considered very *'tightly coevolved." Feinsinger (1983) con- pem Em *most hermits, hermit- like hum- 4 1 4 CJ GNE EL on LLUD coevolution between two divae - proud of species." 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN Species of sect. Erythrina vary in the degree of specificity of their association with the hum- mingbird pollinators. Erythrina species of the dry forests of the Pacific slope are pollinated ex- clusively by Heliomaster constantii, so the plant’s fitness is directly dependent on the behavior and morphology of a single bird species. The oppor- tunity for the plant to evolve adaptations to spe- cific traits of the bird is clear. Heliomaster con- stantii, however, feeds upon and pollinates a number of Eryt t the bird’s geographic range, andi it also feeds ı upon and pol- linates at least one other plant genus (Mandevilla; Feinsinger, pers. comm.). Although there is un- oubtedly a temporary sort of exclusivity in the Erythrina-Heliomaster association in certain ecological communities at certain seasons of the year, the association cannot really be considered an obligate mutualism The plant-pollinator association is less specific for Erythrina species of highland and wet-forest communities, where several species of traplining hummingbirds often visit and pollinate an in- dividual tree on a single day. Hummingbirds such as Eugenes and Campylopterus visit a rather wide variety of plants besides Erythrina. Even in these cases, however, such plant-pollinator associa- tions involving high-reward traplining hum- mingbirds are much more exclusive than those involving short-billed generalist hummingbirds and een E plants. g the species of sect. Erythrina, there is little difitreptianon in floral morphology, flow- patterns, the pollinators—the pollination system of all species is quite similar. Rather, species are dif- ferentiated by their restriction to distinct climatic edaphic conditions. This large group of species, as a whole, has evolved a particular set of adaptations to the guild of high-reward trap- lining hummingbirds. LITERATURE CITED BOLTEN, A. B., P. FEINSINGER, I. BAKER & H. G. B 1980. On the calculation of sugar ease in flower nectar. Oecologia 41: 301-304. CRUDEN, W. R. & V. M. TOLE 1977. Oriole pol- lination of Erythrina breviflora bari evidence for a polytypic v Pla Syst. Evol. 126: 393-4 03. FEINSINGER, P. 1978. Ecological interactions between plants and birds in a successional tropical com- munity. Ecol. Monogr. 48: 269-287. —. 1983. Coevolution and pollination. Pp. 282- [Vor. 74 310 in D. Futuyma and M. Slatkin (editors), Co- evolution. Sinauer Assoc., Sunderland, Massachu- setts. & R. K. CoLweELL. 1978. Community orga- nization among pongo a S edi birds. Amer. Zool. Ls -795. . B. LiN ian A. SWARM & J. A. W 1979. Visio e the ecilination biology of res Erythrina species on Trinidad 2 Tobago. Ann. Missouri Bot. Gard. 66: 451-47 GUILLARMOD, A. J., R. A. JUBB & C. 1 SKEAD. 1979. Field studies m six southern African species of Erythrina. Fo Missouri Bot. Gard. 66: 521-527. HERNÁNDEZ, H. 1982. Female sterility in Ery- thrina M. Allertonia 3(1): 72-76. . TOLEDO. 1979. The role of nectar robbers and pollinators in the reproduction of Er- ythrina leptorhiza. Ann. Missouri Bot. Gard. 66: 512-520. ———— & ————. 1982. Floral biology of Erythrina Ma d p the evolution of pollination sys- n American species of the genus. Allertonia —84. (1: Inouye, D. W. 1980. The vr iid of floral lar- ceny. E 61: 1251-1253. KRUKOFF, B. A. & R. C. Sr 1974. A con- spectus a the genus Erythrina. Lloydia 37: 332- LINHART, Y. B. 1973. Ecological and behavioral de- terminants of pollen dispersal in hummingbird- pollinated Heliconia. Amer. Naturalist 107: 511- McD DADE, L. A. 1984. Systematics and reproductive MORTON, E. S. 1979. Effective pollination of Ery- coev ived behavioral manipulation? Ann. Mis- ouri Bot. Gard. 66: 482-489. Mina. D. A. In press. j studies on species relationships in Erythrina (Leguminosae: Papilio- noideae). Monogr. Syst. Bot. Missouri Bot. Gard. 1911. The birds of North and Middle I part . 1954. Life Histories of Central Amer- ican Birds. Cooper Ornithological Club 1971. A Naturalist in Costa Rica. Univ. of Florida Press, Gainesville. SLup, P. 1960. The birds of Finca “La Selva," Costa Rica: a tropical wet forest locality. Bull. Am. Mus Nat. Hist. 121: 49-148. STEINER, K. E. 1979. Passerine pollination of Ery- thrina megistophylla Diels (Fabaceae). Ann. Mis- souri Bot. Gard. 66: 490—502. . 1975. Ecology, or phenology a i e Costa Rica y 5 9 Ecological pet volutionaty implica- tions of bird pollination. Amer. Zool. 18: 603- 615. 1987] 1981. one aspects of bird-flower co- evolution, with particular reference to Central ouri Bot. Gard. 68: 323-351. 974. Observations on the relation- p S led 487. ERNÁNDEZ. 19 gud iie oliviae: a new case n oriole pollination i r Mexico. An Missouri Bot. Gard. 66: 503-51 APPENDIX Locality and Voucher Data for Erythrina Populations Used in Observations of Floral Visitors (Numbers correspond to those listed in Table 1 1. E. americana Miller. Mexico: Oaxaca, 5 km E of San Pablo Coatlan. 16?12'N; 96?47'W, Elev. 1,450 m. Disturbed gallery = with Taxodium, and cultivated fields. Tree to along intermittent stream. 9-10 Feb. 1983. Neill 5421. 5424. 2. E. berteroana Urban x E. folkersii Krukoff & Moldenke. Mexico: ERAS 3 km S of Palenque. Elev. 100 m. 17?28'N; 92?00'W. Fencepost bordering field. Tree to 8 m. 18 March 1983. = ill 5533. . E. berenices Krukoff & Barneby. Mexico: Vera- cruz, Tequila. 19245'N; 97°03’ W. Elev. 1,650 m. Coffee plantation; Premontane Wet Forest. Tree to 12 m. Jan. — 9 Neill 5381. 4. chiapasana Krukoff. erg: Chiapas, El Su al National Park, Km 14-16. Elev. 1,100 m. 16?47'N; 93°06'W. Disturbed L. Dry Forest, transition to moist mixed Quercus forest. Tree 15 m Mar ch 1983. Neill 5455, 5458, 5465. Krukoff. Mexico: Chiapas, 13 km 6. E. cochleata “naran Costa Rica: ‘Heredia, La Selva Biological Station. Elev. 84°00'W. Tropical Wet Forest. Tree 25 m. 15-18 Aug., 18-21 Sept. 1981. Neill 5015, 5101. . E. costaricensis Micheli. Costa Rica: Puntarenas, San Vito de Java, Las Cruces Botanical Garden. Elev. NEILL— ERYTHRINA POLLINATION 41 1,200 m. 8*45'N; 82*55'W. Premontane Rain Forest, roadside. Tree 6 m. 11-12 Sept. 1981. Neill 5099. 8. E. gibbosa Cufodontis. Costa Rica: Alajuela, up- per Penas Blancas Valley, below Monteverde Reserve. Elev. 1,400 m. 10?20'N; 84?45'W. Premontane Rain Forest; edge of pasture. Tree to 4 m. 4—6 Sept. 1981. Ne x 5057. . E. globocalyx Porsch & Cufodontis. Costa Rica: "s Jose, Las Nubes. Elev. 1,700 m. 9°53'N; Z vials Fencepost row, border of pasture. Tree to 8 m; sporadic wo p 14 Aug., 25 Sept. 1981. Neill 5035. 5142. E. goldmanii Standley. Mexico: Chiapas, El Sumidero National Park, Km 7. Elev. 900 m. gni N; 93°06'W. Tropical Dry Forest; secondary, disturbed scrub. Tree 8 m. 25 Feb.-1 March 1983. Neill, MAT E folkersii Krukoff & Moldenke. Mexico: Ve- ux ur tree to 8 m. 28 Jan 12. E. folkersii koff pas, Palenque Archaeological Site. 17?29'N; 92?01'W. Tropical Wet Forest; forest edge. Tree 5 m. 19 March 1983. Neill clk 13. E. lana ae Mexico: Oaxaca, 37 km W of Puerto Escon dido. ev. m. 15?90'N; 97?20'W. Tropical Dry sid. scrub. Tree 6 m. 13 Feb. 1983. "t 430. a Rose. Mexico: Jalisco, Chamela Bio- local Station. Elev. 250 m. — N; 105?03'W. Tree 13 Jan. 1983. Neill 532 E. pudica Krukoff & M Mexico: Chiapas, Rio de la Venta, Cascada El Aguacero. Elev. 750 m. 16°46'N; 93°33'W. Tropical Dry Forest, scrub. Tree 6 m. 27, 31 March 1983. Neill 5512. 16. E. tuxtlana Krukoff & Barneby. Mexico: Chia- pas, 25 km N of Ocozocuautla. Elev. 700 m. 16?48'N; 93?25'W. Premontane Wet Forest; karst limestone. Tree to 20 m. 28 March, 9 April 1983. Neill 5486, 5621. Barneby. Mexico: Ve- ra 90 m. 17?11'N; 94?39'W. Tropical Wet Forest; kars limestone. Tree 15 m. 16— 17 April 1983. Neill 564 A COMPARISON OF THE DIVERSITY, DENSITY, AND FORAGING BEHAVIOR OF BEES AND WASPS ON AUSTRALIAN ACACIA! PETER BERNHARDT? ABSTRACT Twenty-seven bee taxa and 24 wasp taxa were collected on the open inflorescences and/or extra- floral nectaries of eight Ac acia species in Victoria, taxonomic diversity, bees outnumbered wasp foragers by 88% of Australia. Despite this superficial similarity in the combined catch of winged d, with the short-tongued Halicti- dae and Colletidae comprising the largest unit of native Apoidea o lossum (Halictidae) and Leioproctus (Colletidae) comprised i Th the genera Lasioglos. o short-ton foraging female bees o % of the combined catch of the tw Acacia species. No wasps, however, were collected on var. retinodes o f A. retinodes e families e number of bee n the Acacia species studied. Pollen taxa collected on er through late autumn. Polylectic foraging bee o % of the on extra-floral nectar before foraging on nectarless inflorescences. The density and taxonomic rediere d of wasps remained highest on the Acacia species that offered the greatest volume of sucrose- “rich, extra floral nectar (i.e., A terminalis). B Australia than are wasps. The direct influence of wasps on polyad oan appears to be nominal except in those Acacia species bearing functional extra-floral nectaries Winged Hymenoptera (bees and wasps) have been observed to forage frequently on inflores- cences of Australian Acacia. In contrast to the Psyllidae and some Coleoptera, bees and non- parasitic wasps are not destructive to the small flowers that comprise an Acacia head or spike (Bernhardt, 1982). Instead evidence suggests that Hymenoptera are often pollinators of some Aca- cia species in southeastern Australia (Bernhardt, 1982; Bernhardt et al., 1984; Knox et al., 1985). The flowers of all Australian Acacia examined thus far are nectarless (Bernhardt, 1982; Bern- hardt et al., 1984; Bernhardt & Walker, 1984, 1985; J. Kenrick & G. Beresford, pers. comm.). ar When the anthers Pen is eight polyads pres- ent in each anther are extruded, or partially ex- truded, from their respective sacs (Kenrick & Knox, 1979; Knox & Kenrick, 1982). Conse- quently, bees, wasps, and certain flies easily col- lect polyads from the synchronously opening flo- rets in an inflorescence. Female bees are known to collect Acacia poly- ads to feed to their larvae. Foraging bees remove polyads from the anthers via thoracic vibration of whole inflorescences (Buchmann, 1983) or by scraping anthers directly with their forelegs (Vo- gel, 1978), or both (Bernhardt & Walker, 1984, 1985). Acacia species are usually self-incompat- ible. Seed set tends to occur only when pollina- tors move between genotypes belonging to the same species (Knox & Kenrick, 1982; Bernhardt et al., 1984; Kenrick et al., 1984b). Capture records of insects foraging on Acacia in southeastern Australia and analyses of their pollen loads suggest that Hymenoptera are often more important as polyad vectors than are either ! Research was conducted at the Plant Cell Biology Research Centre of the School of Botany, University of Melbourne under the supervision of R. B. Knox. unding was provided by the Australian Research Grants Scheme and the Australian Department of Education (CPPER). I thank A. Heisler and the rangers of the National Parks Service, Victoria (Brisbane Ranges, Cape Schanck) for their cooperation. This study would not have been possible without the timely D of J. Kenrick, G. Beresford, R. Ma m of Victoria identified Hymenoptera a Walker of the National Mus rginson, P. O'Neal, and T. Hough. J. nd sent pedet wasps to other Knox and D. M. Australian authorities. I am nu grateful for the continuing interest and mPp port R. B. Calder. C. D. Michener provided a most valuable critique of the o riginal man 2 Department of Biology, Saint Louis University, 3507 Laclede, St. Louis, Missouri 63103. ANN. MIssouRI Bor. GARD. 74: 42-50. 1987. 1987] beetle or fly taxa. Calliphorid and syrphid flies transport polyads from Acacia inflorescence to inflorescence without damage, but they may oc- cur on Acacia inflorescences at lower density and diversity than Hymenoptera (Bernhardt et al., 1984; Knox et al., 1985). Excluding the rare documentation of pollina- tion by birds or marsupials (see review by Turner, 1982; Knox et al., 1985) the flowering behavior, floral presentation, and polyad presentation of most Acacia species would be expected to favor a system of generalist entomophily (Bernhardt, 1982; Bernhardt et al., 1984). That is, all insects that forage for polyads have immediate access to the inflorescences of Acacia and could effect deposition of polyads on respective stigmas. Therefore the purpose of this study was to de- termine which groups within the Hymenoptera were major vectors of Acacia polyads with suf- ficient fidelity to regularly effect seed set. To ac- complish this end the density and taxonomic di- versity of polyad foragers were compared to their respective activities on Acacia inflorescences. MATERIALS AND METHODS Acacia species and study sites. Eight Acacia were selected to determine interspecific and in- traspecific foraging preferences of Hymenoptera. The species of Acacia may be found in flower throughout the year (Kenrick et al. 1984a, 1984b; Bernhardt, 1982) with the majority flowering from August through October (Costermans, 1983). Therefore the eight selected species rep- resented the 12-month flowering season of the genus but emphasized the period of intensively overlapping floral phenology from the last month of winter (August) until the second month of spring (October). The periods of fieldwork, study sites, and habitats of each Acacia species are list- ed below. Descriptions of floristic alliances fol- low Specht and colleagues (1979). 1) Acacia longifolia Willd. 31/vi11/84—28/1x/84. Langwarren Reserve: Tall shrubland with dis- rupted epacrid heath (Bernhardt, 1986). 2) A. mearnsii De. Wild. 8/xi/84—2 1/xi/84. Cor- anderrk Reserve: Moist sclerophyll wood- land/shrubland (see Bernhardt & Calder, 1981). 3) A. mitchelii Benth. 16/vii/82-21/1/83. Bris- bane Ranges National Park: Dry sclerophyll woodland/shrubland (see Bernhardt & Walk- er, 1984). 4) A. myrtifolia Willd. 19/viii/82-7/x/82. Bris- BERNHARDT — AUSTRALIAN ACACIA 43 bane Ranges National Park: Dry sclerophyll woodland/shrubland (see Bernhardt & Walk- er, 1984). 5) A. paradoxa DC. (syn. A. armata R. Br.). 12/ 1x/84-31/x/84. Brisbane Ranges National Park: Dry sclerophyll woodland/shrubland (see Bernhardt and Walker, 1984). 6) A. pycnantha Benth. 11/viii/82-16/ix/82. Brisbane Ranges National Park: Dry sclero- phyll woodland/shrubland (see Bernhardt & Walker, 1984). 7) A. retinodes var. retinodes Schdl. 16/i/82—17/ 1/82. Grampians National Park: Montane and valley dry sclerophyll forest/shrubland with adjacent epacrid heaths (see Bernhardt & Walker, 1985) 8) A. retinodes var. uncifolia J. Black. 15/xi/81— 5/11/82. Cape Schanck National Park: Coast- al calcareous dune flora consisting of tall shrubland and invasive, naturalized shrubs and herbs (see Bernhardt et al., 1984). 9) A. terminalis Macbr. 18/iii/83-28/iv/83. Er- ica-Moe (south Gippsland) and Boolah Boo- lah State Forest. Moist sclerophyll woodland/ forest with a rain forest element (see Knox et al., 1985). Analysis of Hymenoptera. The foraging be- havior of Hymenoptera was observed and re- corded over the respective periods of fieldwork every day or every other day. Insects were col- lected selectively from 8 A.M. until 2 P.M. as for- aging behavior becomes negligible by mid after- noon. Insects were collected only if they were ob- served foraging on the open inflorescences of Acacia and/or taking nectar from extra-floral nectaries on the leaves or phyllodes of A. /on- gifolia, A. myrtifolia, A. pycnantha (Bernhardt & Walker, 1984), and A. terminalis (Knox et al., 1985). Foraging is defined here as the active re- moval of polyads from anthers or the probing of flowers and extra-floral nectaries with mouth- parts (Bernhardt et al., 1984). Terminology for bouts of foraging by bee taxa follows Michener (1979). Insects were killed communally in jars con- species carried by insects, each insect was placed on a clean glass slide and “bathed” in a couple of drops of 100% ethanol. When ethanol had evaporated, the white residue remaining on the 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 1. Bee taxa collected on the inflorescences of Acacia. Acacia Species on Which Bee Taxon Was Captured* No. Bee Taxon LO ME MI MY PA PY RR RU TE Bees Anthophoridae: Exoneura (Brevineura) spp. 0 4 0 0 0 0 0 0 0 4 Exoneura (Exoneura) spp. 0 18 0 0 0 0 2 0 11 31 35 Apidae: Apis mellifera” A l l 5 4 A A A A Colletidae: Amphylae 0 0 0 0 0 0 0 0 1 l Calli om 0 0 0 0 0 0 0 0 4 4 Callomelitta spp. 0 0 l 2 0 0 l 0 l 5 Euhesma spp 0 0 0 0 0 3 8 0 0 11 E a al spp. l 0 0 0 2 0 0 0 0 3 Hylaeus sp. 0 0 0 0 0 0 l 0 0 l Leionrocs metallescens 0 0 0 0 0 0 0 14 0 14 . plumo 0 0 0 0 0 0 0 l 0 l ene s (E uryglossidia) spp. 0 0 0 0 42 0 2 0 0 44 Leioproctus (Leioproctus) spp. 3 0 2 12 7 23 19 0 2 79 Trichocolletes sp. 0 0 0 0 l 0 0 0 0 | 1 164 Halictidae: Homalictus brisbanensis 0 2 0 0 0 0 0 10 0 12 H. demissus 0 0 0 0 0 0 l 0 0 l H. dixon 0 0 0 0 0 0 0 0 5 5 H. holochorus l 0 0 0 0 0 0 0 0 l H. megastigmus 0 0 2 0 7 0 l 0 0 9 H. oxoniellus 0 0 0 0 0 0 0 4 0 4 H. punctatus 0 0 0 l 0 0 1 0 0 2 Lasioglossum (Australictus) spp. 0 12 0 0 0 0 0 0 0 12 (Austrevylaeus) spp 0 0 0 0 0 0 0 0 l l La. (Chilalictus) spp. 0 9 12 5 2 2 6 l 4 41 La. (Parasphecodes) spp. 2 l 9 9 22 6 16 71 22 158 Nomia spp. 0 5 5 0 0 0 0 0 0 _10 256 Megachilidae: Megachile sp. 0 0 0 0 0 0 0 2 0 2 2 Grand Total 7 51 31 29 83 34 58 103 51 457 a LO = A. -e ME = A. mearnsii, MI = A. mitchelii, MY = A. myrtifolia, PA = A. paradoxa; PY = A. pycnantha: RR = etin mie var. retinodes; RU - A. ret. var. uncifolia; TE — A. terminalis. — Abundant oe more than five bees caught but far more than that number were identified on sight on this Acacia species). slide was mounted in one or two drops of Cal- dividual unless 25 polyads could be counted in berla's fluid (Ogden et al., 1974). Identification that single sample. Taxa producing monad pol- of pollen was made under light microscopy. len were not counted as present in a sample un- However, since insects were killed in the same less 25 individual grains were identified on a jar, contamination of pollen species was possible. single slide (Bernhardt et al., 1984). Insects Therefore an Acacia species or species of Epa- washed of pollen were air dried, numbered, place cridaceae was not counted as present on an in- in individual glassine bags, and identified. 1987] BERNHARDT — AUSTRALIAN ACACIA 45 TABLE 2. Characteristics of the secretion of extra-floral nectar in four Acacia species.* Characteristics of the Nectary or Nectar LO MY PY TE Nectary secretes throughout the flowering seasons of + + + Nectary secretes > 1 ml every morning? = = = + Nectar hexose ric ? = + — Nectar sucrose rich or sucrose dominant ? + = + ? Derive > Observations and collections of nectar made bet from Bernhardt (1982), Bernhardt and Walker AR and Knox et al. (1985). tween 7-9 A LO = A. longifolia, MY = A. myrtifolia; PY = A. pycnantha; TE = A. terminalis. RESULTS BEES Bees foraged for polyads on each of the eight Acacia species (Table 1). A total of 457 bees were caught on the eight Acacia species. The greatest numbers of bees collected belonged to the short- tongued family of Halictidae. Of all five families the Halictidae offered the greatest number of gen- era collected. Lasioglossum (subgenus Parasphe- codes; Halictidae) species were collected in great- er numbers on Acacia compared to all other locally abundant on half of the Acacia species studied and could be identified effortlessly on sight (Table 1). This bee was the dominant for- ager on A. /ongifolia. Five bee families were identified on eight Aca- cia species. However, seven out of the 27 bee taxa represented single captures. The long- tongued families Apidae and Megachilidae were represented by only one taxon each. The third long-tongued family, Anthophoridae, was rep- resented by two subgenera of Exoneura (Table 1). All remaining bee taxa belonged to the short- tongued families Halictidae and Colletidae. Eighty-four percent of the Colletidae collected belonged to the genus Leioproctus s.l. Eighty- three percent of the Halictidae belonged to the genus Lasioglossum s.l. (Table 1). Lasioglossum (subgenus Parasphecodes) species and A. mellif- era were the only bee taxa collected on each of the eight Acacia species (Table 1). Leioproctus (subgenus Leioproctus) Lud were caught on seven out of eight Acacia speci Representatives from the Ë wiis Antho- phoridae, Apidae, ae i and Halictidae were recorded as foraging on the extra-floral nectaries of A. longifolia, A. myrtifolia, A. pycnantha, and A. terminalis (Tables 1, 2). There was no evi- dence that Acacia inflorescences growing on shoots bearing extra-floral nectar received great- er numbers of bees (or mare bee taxa) compared to those A (Table 1). There was no obvious correlation between the number of bee taxa collected on an Acacia species and the habitat in which the species was studied (Table 3). At the intraspecific level, though, the dry sclerophyll woodlands and heaths of the Grampians National Park offered a wider fauna for A. retinodes var. retinodes than did the coastal dunes of Cape Schanck (Table 3). The number of bee taxa below the family level collected on four Acacia species sympatric through the Brisbane Ranges National Park varied from five to eight. No more than four families of bees were collected on any Acacia species (Tables 1, 3). Variation in the number of bee taxa on an Acacia species did correlate positively with the flowering seasons of the plants and increased from mid winter (July) until the end of autumn (May; see Table 4). The number of bee taxa collected terminalis (Table 4). The greatest number of bee taxa was collected on the summer-flowering A. retinodes var. uncifolia (Table 4). The majority of bee taxa foraging on the eight Acacia species visited a wide range of plants for pollen and/or nectar (Tables 4, 5). Technically these bees must be classified as polylectic for- agers (Armstrong, 1979; Michener, 1979). From mid winter until late spring, and then from au- tumn to early winter, 21-55% of the bees cap- tured on any Acacia species carried the polyads of that species mixed with the pollen of one or more sympatric species. Furthermore 63-91% of the bees captured on Acacia from late spring to late summer carried the pollen of one or more sympatric plants mixed with Acacia polyads (Ta- ble 4). Bees collected on A. pycnantha showed 46 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE 3. Variation in the number of bee taxa col- lected at different sites. No. Bee Taxa Acacia (No. Site? Taxon Families) Brisbane Ranges MI 7 (3) MY 6 (3) PA 8 (3) PY 5 (3) Cape Schanck RU 8 (4) Coranderrk Res ME 8 (3) Erica-Moe; Bub on TE 10 (4) Grampians RR 12 (4) Langwarren Reserve LO 5 (3) ? See study sites for description of habita A. ae. PA = t. A, retinodes var. ` retinodes. LO = A. longifolia. the lowest level of polylectic foragers, whereas bees collected on A. mearnsii showed the highest (Table 4 Sixty-three percent of the bees bearing loads of Acacia polyads that were mixed with the pol- len of other plants carried monads of one or more genera of Myrtaceae (Table 5). Ninety-two per- cent of the bees carrying mixed pollen loads car- ried the pollen of plants that bear floral nectaries Table 5) WASPS Wasps were collected on all Acacia species ex- cluding A. retinodes var. retinodes. Only 58 wasps were collected on the Acacia species representing seven wasp families (Table 6). Sixty-four percent of the wasps were divided equally between the Sphecidae and Tiphiidae. The greatest numbers belonged to Cerceris s.l., but these insects were collected only on A. paradoxa and A. terminalis Table 6) A total of 24 wasp taxa below the family level were collected on Acacia. Eleven of these rep- resented single captures. No wasp taxon was re- corded on more than two Acacia species. Acacia species offering extra-floral nectar appeared to attract a minimum of twice as many wasp taxa as those lacking extra-floral nectar (Tables 2, 6). Wasps observed and collected on A. /ongifolia, A. myrtifolia, and A. terminalis consistently flew to the extra-floral nectary before attempting to forage on the nectarless florets. [Vor. 74 TABLE 4. Flowering patterns of Acacia species as correlated with the taxonomic diversity and foraging behavior of the bees. No. Bee No. Bees axa on Bearing* Flowering Period Acacia Acacia and Acacia Species sp. Polyads Ratio’? Mid Winter-Early Spring A. longifolia 22 (9) 0.40 A. pycnantha 41 (9) 0.21 Mid Winter-Mid Spring A. myrtifolia 29 (16) 0.55 Early Spring-Late Spring A. paradoxa 96 (50) 0.52 Late Spring A. mearnsii 8 59 (45) 0.91 Early Summer-Mid Summer A. mitchelii 7 32 (25) 0.78 Early Summer-Late Summer A. retinodes var. retinodes 12 67 (56) 0.8 A. retinodes var. uncifolia 8 121 (77) 0.63 Autumn-Early Winter A. terminalis 10 50 (16) 0.32 a The first number in the column refers to the total number of bees caught on the particular Acacia sp. that carried polyads of Acacia on their bodies. The second number, in parentheses, refers to the total number of +L 4 a Te | $ 1 4 1 L 11 y EMT NP genera, ° The ratio = number of bees that carried Acacia use plus pollen of other genera divided by total mber of bees that carried Acacia polyads. Acacia terminalis offered more extra-floral nectar per gland on a daily basis than did the three other nectariferous species (Table 2). Aca- cia terminalis recelved 33—66% more wasp taxa than were captured on the other three nectarifer- ous species. Wasps foraged selectively on Acacia species am dunchona ind Pora Hectanes. These te id spring and from autumn to early mns (Tables 4, 6). Consequently the density and diversity of wasps on Acacia was heaviest during the coldest seasons. Seventeen wasp taxa carried the polyads of at least one of the Acacia species on which they were caught: Antamenes sp., Anthobosca sp., 47 1987] BERNHARDT—AUSTRALIAN ACACIA TABLE 5. Comparative frequencies of pollen from other sr phyte famili identified on bees carrying Acacia polyads. No. of Bees Bearing Acacia Polyads Mixed wh of Other Families Captured on each Acacia Spermatophyte Family? LO ME MI MY PA PY RR RU TE Total Compositae + — 0 4 0 0 3 I 6 26 0 40 Dilleniaceae — 0 0 3 0 4 l 0 0 0 8 = semaine + 0 0 0 l 3 l 0 0 3 8 umin Tanaman PNA l 4 0 0 23 0 0 0 0 28 Liliaceae s.l. + 0 6 0 0 2 0 0 0 0 8 Myrtaceae + 2 47 19 2 26 0 71 43 15 225 inaceae — 9 0 0 0 1 l 0 0 0 11 Pittosporaceae + 0 0 0 0 0 0 2 0 0 2 Proteaceae + 0 0 0 4 0 3 0 0 0 7 Rhamnaceae + 0 0 0 12 2 4 0 0 0 18 * + = bees bearing pollen of genera with nectariferous flowers; — = bees bearing poll fg tł tarl flowers. LO = A. longifolia, ME = A. mearnsii, MI = A. mitchelii MY = A. myrtifolia, PA = A. paradoxa; PY = A. pycnantha; RR = A. retinodes var. retinodes, RU = A. ret. var. uncifolia, TE = A. terminalis. Cerceris antipodes, Cerceris sp., Labium sp., Lis- sopimpla excelsa, Lissopimpla i Lophocheilus s sp., Phyma re tiles pygi- . Sp. near nonilicornis, pd us Sp., Pseudozethus sp., Rhagigaster PASEAR A Sphex sp., Tachynomia moerens, and the unidentified species of Tiphiidae. Mixed loads of Acacia poly- ads plus the pollen of sympatric plants were con- fined to those angiosperm families with nectar- iferous flowers: Compositae, Epacridaceae, Myrtaceae, Rhamnaceae, and Solanaceae. DISCUSSION Wasps versus bees as Acacia pollinators. The asps, however, comprise only 11% of the total population of foraging Hymenoptera. This supports previous reports that bees forage for Acacia polyads in far greater numbers than do wasps (Bernhardt, 1982; Bernhardt et al., 1984; Bernhardt & Walker, 1984, 1985; Knox et al., 1985). Female bees appear to forage primarily for polyads and secondarily for extra-floral nec- tar (Bernhardt & Walker, 1984; Knox et al., 1985). The results presented in this paper indicate that wasps tend to forage preferentially for extra-flo- ral nectar. Large quantities of extra-floral nectar fail to assure polyad transport by wasp foragers (Knox et al., 1985). Almost equal numbers of bees and wasps foraged for nectar and polyads on valley populations of A. terminalis, but bees consis- tently carried polyads of A. terminalis more often than did wasps (Knox et al., 1985). The low den- sity of wasp populations on flowering Acacia, combined with the foraging preferences of these insects, suggests that wasps are of secondary im- portance to seed set, at best. Under certain cir- cumstances, pollen-eating flies are probably su- perior polyad vectors compared to wasps (Bernhardt et al., 1984) This does not mean to suggest that wasps are always nominal pollinators ofthe Australian flora. Unfortunately few studies have been done to as- sess the selective pressure of wasp foraging on other angiosperms (Armstrong, 1979). Waspsare important pollinators of nectariferous and nec- tarless (pseudocopulatory) Orchidaceae in Aus- tralia (Beardsell & Bernhardt, 1982). Wasps may also pollinate the nectariferous Epacridaceae (Bernhardt, pers. obs.; Knox et al., 1985). Of course, wasps are important pollinators of Pro- sopis (Simpson et al., 1977), but these mimosoid shrubs bear floral nectaries. Bee diversity on Acacia species. Although all representatives from all five families of Apoidea found in Australia (Armstrong, 1979) were iden- tified on Acacia in this study, some families were more common than others. This is partially ex- plained by biogeography. The Apidae are poorly represented in Australia (Michener, 1979), and the native genus Trigona is uncommon south of the Tropic of Capricorn (K. Walker, pers. comm.). 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 TABLE 6. Wasp taxa collected on the inflorescences of Acacia species. Acacia spp. on Which Wasp Was Captured No. Wasp Taxon LO ME MI MY PA PY RR RU TE Wasps Braconidae: Apanteles sp. l 0 0 0 0 0 0 0 0 1 l Eumenidae: Antamenes sp. 0 3 0 0 0 0 0 0 0 3 Pseudozethus sp. 0 0 0 0 0 0 0 0 5 5 Unidentified sp. 0 0 l 0 0 0 0 0 1 9 Ichneumonidae: Biconus sp. 0 0 0 1 0 0 0 0 0 1 Labium spp l 0 0 0 0 0 0 0 3 4 Lissopimpla excelsa 0 0 0 1 0 0 0 0 0 l Lissopimpla sp 0 0 0 0 0 0 0 0 2 2 8 Pergidae: Neourys sp. 0 0 0 0 0 l 0 0 9 l l Pompilidae: Chirodamus sp. 0 0 0 0 0 0 0 0 1 l Pompilius sp. 0 0 0 0 0 0 0 0 1 1 2 Sphecidae: Cerceris antipodes 0 0 0 0 0 0 0 0 7 Ji Cerceris spp. 0 0 0 0 1 0 0 0 6 7 n sp. 1 1 0 0 0 0 0 0 0 2 Rhopalum 0 0 0 0 0 0 0 0 l l Sphex sp. 0 0 0 l 0 0 0 0 0 mt 18 Tiphiidae: Anthobosca sp. 0 0 0 0 0 0 0 3 5 E anilitatus 0 0 0 l 0 0 0 0 2 Phymatothynnus pygidalis 0 0 0 0 1 0 0 9 l Phymatothynnus sp. nov. n. nonilicornis* 0 0 0 0 0 3 0 0 0 3 Phymatothynnus 0 0 0 1 0 0 0 0 0 l hagigaster comparatus? 0 0 0 0 0 2 0 0 0 2 Tachynomia moerens* 0 0 0 0 0 2 0 0 0 2 Unidentified spp. 2 0 0 0 1 0 0 0 0 3 19 Total No. Nerd Acacia specie 5 4 1 5 2 10 0 3 28 58 Total No. a Taxa/ Acacia species 4 2 l 5 2 6 0 l 9 a Refers to wasp taxa caught en asa = A. longifolia, ME = A. mea : MI = A. mitchelii; MY = A. myrtifolia, PA = A. paradoxa; PY = A. pycnantha; RR = A. retinodes var. wna ri RU = A. ret. var. uncifolia; TE = A. terminalis. 1987] Trigona may be important in the pollination of paleotropical Acacia through northeastern Aus- tralia (Bernhardt, 1982; Armstrong, 1979). Biogeography alone, however, cannot explain the comparative dearth of anthophorid and megachilid bees on Acacia species in southeast- em Australia, While it is true that neither family ity of Colletidae in Australia (Michener, 1979), there are far more megachilid and anthophorid taxa in Victoria than were sampled in this study (Michener, sidered families of long-tongued bees. Although they may be common throughout the habitats described in this paper, evidence suggests that they forage preferentially on flowers offering pol- len and nectar (Armstrong, 1979). In particular, Megachilidae often are associated with zygo- morphic flowers, particularly papilionoid le- gumes (Michener, 1965; Armstrong, 1979). The only anthophorid bees relatively common on Acacia were Exoneura. These bees were largely confined to the only two arborescent Acacia species studied bearing bipinnately compound leaves: A. mearnsii and A. terminalis. This is probably coincidental, but it would be worth- while exploring “presumed” foraging preferences that Exoneura may express towards sections of Acacia s.l. Climatic conditions may also influence the di- versity of bee taxa on Acacia with overlapping distributions. During the spectacular drought of 1982 and 1983 no specimens of Leioproctus (subgen. Euryglossidia) were collected on Acacia in the Brisbane Ranges (Bernhardt & Walker, 1984). The Brisbane Ranges had a cool, wet spring in 1984, and in that year Leioproctus (subgen. Euryglossidia) far outnumbered previously abundant Leioproctus (subgen. Leioproctus). Considering the broad polylecticism of the bee taxa on these eight Acacia species, one cannot refer to any bee taxon as an “acacia bee" as some entomologists speak of **clover bees,” “orchid bees,” and “squash and gourd bees” (Michener, 1974, 1979). The very absence of floral and ex- tra-floral nectar in so many Australian Acacia species forces bees to forage on other sources. Oligolecty continues to decline as bees exploit co-blooming sources of nectar and additional pollen in competition with Acacia. Lasioglossum (Halictidae) and Leioproctus (Colletidae) remain the predominant polyad for- agers of many Acacia species (Bernhardt, 1982; Bernhardt et al., 1984; Bernhardt & Walker, 1984, = oO BERNHARDT—AUSTRALIAN ACACIA 49 1985; Knox et al., 1985). Lasioglossum species may be more important pollinators than Leio- proctus species. Lasioglossum were caught on all Acacia species studied and were collected in greater numbers than were Leioproctus. Parsphecodes and Chilalictus were the most commonly collected subgenera on six out of the eight Acacia species studied. Parasphecodes bees outnumbered Chilalictus bees. These allied sub- genera may exert preferential foraging patterns when nectarless angiosperms have overlapping flowering periods. From later winter through late spring Acacia species and some Hibbertia species (Dilleniaceae) have H unes flowering pe- riods. Lagio, en. Chilalictus species forage iq e on yrs (Bernhardt, 1984, 1986), but Parasphecodes forage in greater ni Des on Acacia (Bernhardt & Walker, 1985). Polylecticisms and the role of polylectic bees in Acacia cross-pollination. The Halictidae and Colletidae are both families of short-tongued bees that forage for nectar on flowers with shallow widely distributed family of nectariferous angio- sperms in Australia (Costermans, 1983). Most shrubby Myrtaceae flower through the warm spring-summer months when the greatest num- ber of bees (both short- and long-tongued) are active. Out of the four most important nectarif- erous families that were patronized most fre- quently by bees in this study, three have flowers with short-tubular or shallow-bowl perianths (Faegri & van der Pijl, 1979): Myrtaceae, Com- positae, and Rhamnaceae. The keeled “flag” flowers of the papilionoid legumes are also vis- ited by short-tongued bees probably because the weight of comparatively large-bodied Leioproc- tus and Lasioglossum triggers access to the con- cealed nectaries. Bees that pollinate nectarless flowers tend to belong to polylectic genera: Apis (Michener, 1974), Bombus (Buchmann, 1983), Centris (Frankie et al., 1983), Lasioglossum (Bernhardt, 1984; Bernhardt & Burns-Balogh, 1986). It is not unusual for an individual bee that belongs to a taxon in which foragers are not fed by siblings to visit one to four nectarless but pollen-rich species and six to eight nectariferous species dur- ing flowering seasons (Macior, 1968; Bernhardt & Montalvo, 1979; Bernhardt, 1984; Bernhardt & Burns-Balogh, 1986). Acacia species in Australia are, in general, mass- flowering plants that are also highly self-incom- 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN patible. Seed set will not occur unless bees move spontaneously from one shrub to another. The absence of floral nectar forces polyad foragers to break their bouts on Acacia species with trips to nectariferous flowers growing on nonphyloge- netically related plants (Bernhardt & Walker, 1984, 1985). When the bees are replete with chemical energy (i.e., nectar) they may return to Acacia, but in the mosaic distribution of scle- rophyll shrubs it will probably be to a shrub dif- ferent from the one abandoned for nectar. LITERATURE CITED ARMSTRONG, J. A. 1979. Biotic pollination mecha- nisms in the Australian flora—a review. New Zea- . BERNHARDT. 1982. Pollination biology of Australian terrestrial orchids. de l 183 ¿n E. G. Williams, R. B. Knox, J. H . Gilbert & P. Bernhardt (editors), Pollination 82. Plant Cell Biology Research Centre, Univ. of Mel- rne, Australia. Maid P. 1982. Insect pollination of Australian a. Pp. 85-97 in E. G. Williams, E. Knox, J. ilber & P. Bernhardt (editors), Pollination '82. Plant Cell rid Research Centre, Univ. of n Austra 4. The s sete biology of Hibbertia sri oS PI. Syst. Evol. 147: 267-277. Bee-pollination in Hibbertia fascicu- lata Dilniaeae. Pl. nr Evol. 152: 231-241. 1986. Floral mimesis » Thelma Mene: Br. Pl. Syst. Evol. 151: 187- D. M. CALDER. 1981. The floral ecology of pasau populations of Amyema pendulum and yema T (lanta 2). Bull. Torr. Club 108: 213-230. & . MONTALVO. 1979. The pollination ecolo ogy of Echeandia macrocarpa (Liliaceae). Brittonia 31: 64-71. K. WALKER. 1984. Bee foraging on three patric species of pueda Acacia. Int. J. Entomol. 26: 322-330. 85. Insect foraging on Acacia ee var. . retinodes. Int. J. Entomol. 27: 97- l . KENRICK & R. B. KNox. 1984. Pollination biology ay is breeding system of Acacia reti- es (Leguminoseae: Mimosoideae). Ann. Mis- souri Bot. Gard. 71: 17-29. BUCHMANN, S. L. 1983. Buzz pollination in angio- rms. Pp. 73-114 in C. E. Jones & R. J. Little (editors), Handbook of Experimental Pollination Biology. Van Nostrand wee ias Inc., New York. COSTERMANS, L. tive Trees and Shrubs of sm Australia. "Rigby Publishers, Austra- maa N DER Pur. 1979. Principles of Pollination Y Ecology, 3rd edition. Pergamon, Ox- ford/New Y [VoL. 74 FRANKIE, G. W., W. A. HABER, P. A. OPLER & K. S. Bawa. 1983. you and organization of Little (editors), Handbook of Experimental Pol- lination Biology. Van Nostrand Rheinhold Inc., New York. Kenrick, J. & R. B. KNox. 1979. Pollen develop- ment and cytochemistry in some Australian species of rer ore J. Bot. 27: 412-427. L & R. B. Knox. 1984a. Self-incom- ud in Acacia. —a pre- or post zygotic mech- anism? Pp. 146-153 in E. G. Williams & R. B. Knox (editors), Pollination '84. Plant Cell Biology Research Centre, School of Botany, Univ. of Mel- bourne, Australi ERN ali HARDT, ` R. MARGINSON, G. BERESFOR & R. B. KNOX. : in E. Pollination '84. Plant Cell Biology Research Cent Wigs of Botany, Univ. of Melbourne, Austra lia. Knox, R. B. & J. KENRICK. 1982. Pol dup —_— in Suus to the breeding system of 4 411—418 in D. Mulcahy & E. Ottaviano editor Pollen Biology. oleis Holland Press, Amsterdam ,P HARDT, R. MARGINSON, BERESFORD, I. BAKER & H. G. BAKER. 1985. Ex- tra- in nectaries as sr ashi Sd bird polli- ation t. 72: 1185- 196. MacioR, L. W. 1968. Bombus (Hymenoptera, Api- dae) queen foraging in relation to vernal pollina- tion in Wisconsin. Ecology 49: 20-25 MICHENER, C. D 65. A classification of the bees of the Australian and South see regions. Bull. Amer. Mus. Natur. Hist. 130: 62. . 1974. The Social ake of the Bees. Belk- nap Press of Harvard Univ., Cambridge, Massa- chusetts. 1979. Seer dd of the bees. Ann. Mis- souri Bot. Gard. 16: 277-347. pling Airborne Pollen. “Hafner, New York. Simpson, B. B., J. L. NEFF & A. R. MOLDENKE. 1977. Pp. 84-107 in B. B. Simpson (editor), wa atl Its Biology in Two Desert Scrub Ecosyste Dowden, Hutchinson & Ross, Inc., rai ei Pennsylvania SPECHT, R. A., E. M. RAE & V. H. BOUGHTON. 1974. Conservation of major plant communities in Aus- tralia and Papua New Guinea. Austral. J. Bot. Suppl. Ser. 7: 1-667. TuRNER, V. 1982. Non-flying mammal pollination: an opportunity in appen Pp. 110-121 in E. G. Williams, R. B. Knox, J. H. Gilbert & P. Bernhardt (editors), Pollination '82. Plant Cell Biology Re- search Centre, oe Botany, Univ. of Mel- bourne, Austra VoGEL, S. 1978. c T shifts from reward to deception in pollen flowers. Pp. 89-96 in A. J. Richards (editor), x Pollination of Flowers pied Linn. Soc. Symp. No. 6, Academic Press, FLOWER LONGEVITY AND PROTANDRY IN TWO SPECIES OF GENTIANA (GENTIANACEAE)! C. J. WEBB AND JAN LITTLETON2 ABSTRACT Both Gentiana saxosa and G. serotina are protandrous. When flowers open, pollen is presented extrorsely around the closed stigma for one to six days. As the stigma opens, t e toward the corolla lobes. The length of the female phase, and therefore reproductive flower life, is ae hakai by pollination, although in both species the corolla may remain fresh for longer than one month. Fresh ale- i i flowers pollinated on their tenth day of stigma presentation seme no om ipii they appeared fresh. Senescence of unpollinated flowers differed between species: in G. saxosa the flowers remained open and gradually deteriorated, but in G. serotina the flowers asus d diei before full senescence. Pollination-induced flower senescence has been demonstrated for a number of other angiosperms, and the usual reactions to pollination are corolla abscission, color change, or wilting. In Gentiana, the closed corolla enfolds the large superior ovary and may serve to protect it from predators as well as prevent further pollinator visits. Pollination-i maintenance costs by ensuring that the flower functions no longer than neces sary. One correlate of this =o in hermaphroditic flowers is protandry, which ensures pollen dispatch before flower clos Floral senescence may be either time-depen- dent (endogenous) or exogenous (usually polli- nation-induced). However, there have been few detailed investigations of the factors that deter- mine floral senescence, and hence, floral longev- ity (Primack, 1985). Most such studies have con- ceptrated on corolla tor changes or other t and follow p signal senescence (e.g., Arditti et al., 1973; yee ditti, 1976; Gottsberger, 1971; Gori, 1983; Strauss & Arditti, 1984; Casper & La Pine, 1984; Halevy, 1984), and less attention has been afforded struc- tural changes such as wilting, flower closure, and corolla abscission (Mayak & Halevy, 1980). Most of this research is concerned with the proximate determinants of floral longevity rather than the evolution of particular responses (but see Stead & Moore, 1979; Gori, 1983; Casper & La Pine, 1984; Devlin & Stephenson, 1984). The paucity system and ultimately for the plant’s overall re- productive strategy. For instance, in hermaph- rodite flowers pollination-induced flower senes- cence will limit the duration of pollen and stigma ! We thank I. C. Brown and garden staff at Lincoln, presentation and so may influence or be influ- enced by the extent and nature of dichogamy (Lloyd & Webb, 1986). New Zealand species of Gentiana (in the southern group of Philipson, 1972) are protan- drous (Thomson, 1881; Simpson & Webb, 1980; Webb, 1984a). Their large, relatively simple owers make them particularly suitable subjects for experimental studies of flower function. This paper describes the response of flowers of two species to pollination and reports the results of experiments to determine the functional dura- tion of male and female phases in terms of pollen presentation and seed production. MATERIALS AND METHODS The two species of Gentiana selected were those that grew best in cultivation. Gentiana saxosa Forster f. grows naturally in coastal sites of south- ern South Island and Stewart Island, New Zea- land; plants were collected from Curio Bay, Southland. enti serotina Cockayne occurs in grassland in h Island; plants were collected from Lake Lyndon, Canterbury. The plants were grown in clay pots in an insect- proof cage in greenhouses at Lincoln, Canter- Canterbury, for maintaining the plants in cultivation, J. Miles for spen with pye and P. Brooke for pp aaa 2 = > We are grateful to L. F. Edgar, Daph- Lively, nd D. oyd for co mments on a draft e manu 2 Botany n Department an E and Industrial Meu y see Bag, Christchurch. New Zealand. ANN. Missouni Bor. GARD. 74: 51-57. 1987. 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE 1. Results of controlled pollinations to de- termine self-compatibility in Gentiana saxosa and G. serotina. % Seeds Pro- u ap- duced ber of sules per ow- ro ap- Treatment ers duced sule G. saxosa Cross- 10 10 94.2 pollinated Self- 10 10 94.2 pollinated Unpollinated 10 0 — G. serotina Cross- 5 5 57.9 pollinated Self- 5 5 67.6 pollinated Unpollinated 5 0 — bury. Field observations were made at Lake Lyn- n. To test for self-incompatibility and autogamy 10 flowers in G. saxosa and five flowers in G. serotina were assigned to each of three treat- ments: self- and cross-pollination by hand, and unpollinated (Table 1). Flowers were tagged in- dividually and observed daily. When capsules matured, they were harvested and good seeds and aborted seeds or undeveloped ovules were counted. The duration of male and female phases was determined for individual tagged flowers. Pollen and stigma presentation were observed daily un- til petals withered and turned brown. Of 61 flow- ers of G. saxosa observed daily, five were cross- pollinated on each of the first to tenth days of their female phase, and 11 left unpollinated. Of 34 flowers of G. serotina, eight were pollinated on each of the first, fifth, and eighth days of their female phase, five on the tenth day, and five flowers were left unpollinated. For G. serotina, three flowers from each of the first, fifth, and eighth day pollinations were collected two days after pollination, the stigma dissected out, fixed, stained with aniline blue, and examined under fluorescent microscopy to determine the extent of pollen germination and pollen tube growth. For the remaining flowers, the responsiveness of each to pollination was quantified as the number of days until (a) the flower closed completely, (b) closed to its greatest extent (for flowers that closed [Vor. 74 incompletely), or (c) the number of days until the corolla withered and turned brown (for flow- ers that never closed). When capsules were ma- ture they were harvested and seed production scored. At Lake Lyndon, 10 fresh, unbagged, female- phase flowers of G. serotina were cross-pollinat- ed by hand, and ten were left unpollinated. These flowers were examined the following day. RESULTS Flower form, phenology, and pollina- tion. Both Gentiana saxosa and G. serotina are perennial herbs with a central leafy rosette and stout taproot. The flowers are on annual, leafy, lateral flowering branches that bear from one to eight flowers in G. serotina and one to 30 or more in G. saxosa. The corolla is deeply 5-lobed, white with translucent stripes on the lobes and upper tube (Fig. 1), and greenish-yellow toward the base of the tube. The five stamens are attached to the corolla tube; the anthers are purple and the pollen is cream to brownish-yellow. Nectar is produced between the bases of the filaments near the base of the tube. The central ovary contains an av- erage of 29 ovules in G. saxosa and 47 in G. serotina (Webb, 1984a). The flowers lack a well- defined style; the stigma is 2-lobed, distinctly papillate, and dry. In the field, G. saxosa blooms in summer and autumn (January to May), and G. serotina from late summer to autumn (February to May). Un- der greenl diti G hed peak flowering in December and G. serotina in March. The flowers are sweetly scented and visited on fine days by a range of insects. At Lake Lyndon, G. serotina was most frequently visited by syr- phid flies and solitary bees (Lasioglossum sor- didum). During anthesis, flowers opened com- pletely only on fine days, and almost completely closed at night. On wet or cold, dull days few new flowers opened, and in those that had opened previously the corolla lobes did not spread com- etel = Both G. saxosa and G. serotina are self-com- patible (Table 1); however, flowers that were not pollinated and from which insects had been ex- cluded failed to produce any seed, so biotic pol- lination is necessary for seed set. Protandry. The flowers of both species are distinctly protandrous. In neither was there any obvious synchrony of male and female phases among flowers within a plant or even a flowering branch, so that within large plants pollen and 1987] WEBB & LITTLETON—GENTIANA 53 FiGures 1-8.— 1-7. Protandry and reaction to pollination in flowers of Gentiana serotina (bars = 0.5 cm 1. First open, male phase.—2. Fourth corolla. — after pollination of fresh Pani phase flower.— 6. eighth day of female phase.— (bar = 0.5 cm) stigmas were presented simultaneously in differ- ent flowers and geitenogamy could occur. When a flower first opens, the anthers have already dehisced to present pollen ext ly near the center of the flower and the stigmatic lobes are tightly closed (Fig. 1). Later in the male phase, the corolla lobes open further, the stigma begins to open, and the stamens move outwards toward the corolla (Fig. 2). In G. saxosa, the stigma may open and stamens recurve late on the first day of anthesis, or as late as the fifth day. The mean duration of the male phase is 1.41 days (N = 57). In G. serotina, the stigma opens and stamens recurve between the third and seventh days (mean =4.73 days, N = 30). In G. saxosa, an- thers have usually moved halfway toward the corolla when the stigma opens and have reached the corolla lobes in up to three days after that. In G. serotina, the anthers reach the petals and usually wither one or two days after the stigma opens (Fig. 3), and in some cases they eventually protrude between the corolla lobes. Although pollen is no longer presented in a central position by the time the stigma opens, there may be some overlap between pollen and stigma presentation within the flower, especially in G. saxosa, for which the male phase is shorter. However, under 4. Flower closing, one day after arcade of fresh female-phase flower. — Flower incompletely closed, five days after pollination on 7. Unpollinated flower, 18 days.—8. Unpollinated flower of G. saxosa, 30 days day, stigma opening.—3. Seventh day, 1 igma open, anthers near 5. Flower closed, two days field conditions, little pollen usually remains by the time the stigma opens. When the stigma opens the two lobes occupy the position in which pollen was presented in the newly opened flower (Figs. 1, 3). The duration of the female phase is dependent on pollination. Reaction of the flower to pollination. Both species showed similar reactions to pollination. When fresh, first-day, female-phase flowers were pollinated, they partially closed that night and failed to reopen the following day (Fig. 4). Two days following pollination the corolla lobes were imbricate to form a neat, tight structure similar to, but slightly larger than a late bud (Fig. 5). The petals slowly turned brown and withered as the capsule reached maturity after about four weeks. When the flower was pollinated on the second to fifth days of the female phase, the corolla re- acted as quickly and folded as neatly as it did following first-day pollination. After the fifth day, u „ei close or closing incomipietely third of the flowers pollinated after the fifth day of their female phase never closed completely. There was some variation among flowers of the 2 4 6 8 10 DAY OF FEMALE PHASE FIGURE 10. Average percentage of ovules produc- 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 17 100., Z 5 ; Oso | | C5 O Z p =) c z C60 . T / O = / a = 10 4 40 OQ na ul S TE n WY 20 | D o o IN S O te ae 0 T T T T *—. 1 O = (O > < e o NEVER R POLLINATED DAY OF FEMALE PHASE ` E9. Days taken for flowers of Gentiana sax- osa (solid dots) and G. serotina (open squares) to close or wither NE pollination on different days of emale phas same age, but the trend to decreasing reactivity with increasing age was clear in both species (Fig. ). In the field, flowers of G. serotina reacted sim- ilarly to those in the greenhouse. Ten fresh fe- male-phase flowers were all closed on the day following hand pollinations, whereas eight of 10 flowers not pollinated by hand had reopened. Under fluorescent ai ee all three stig- mas from flowers of G. serotina that had been pollinated on the first day of the female phase had numerous germinating pollen grains and many well-formed pollen tubes penetrating the stigma. Of the three stigmas pollinated on their fifth day, two showed good pollen germination and the third had only a few germinating grains. Two of the three stigmas that had not been pol- linated until their eighth day had no pollen ger- minating, and the third had good germination. The proportion of ovules that developed into good seeds was clearly affected by the age of the production for flowers pollinated later than the fourth or fifth days of the female phase (Fig. 10). In G. saxosa, flowers pollinated on the first to fourth days of the female phase consistently ma- ing seed for flowers of Gentiana saxosa (solid dots) and : í Y fall : 11: 4i am ALS ferent days of female phase (mean + standard error for G. saxosa, mean only for G. serotina). tured a high proportion of their ovules. Those pollinated on the fifth to seventh days were more variable in seed production, and seed production was related to reactivity of individual flowers. For example, a flower pollinated on the fifth day failed to close and produced no seed, whereas a flower pollinated on the seventh day closed al- most completely and had 25% seed production. Few flowers pollinated on the eighth day of the female phase, and none pollinated on the ninth or tenth days peoduced soea, The panem of seed production for fl in G. serotina was similar (Fig. 10), but the small- er sample size does not allow a detailed analysis. Thus, although the corolla and stigma may appear fresh for much longer, the functional fe- male phase in both species is only four to eight days. Adding together male and female phases, one gets a reproductively functional flower life of five to 12 days for G. saxosa, and six to 14 days for G. serotina. The reaction of flowers that were never pol- linated differed between species. In G. serotina, the corolla lobes continued to diverge as the flow- er aged, but eventually began to close (Fig. 7), finally reaching a position similar to that of a late-pollinated flower (Fig. 6) before turning recurved position and in all but one of 11 flowers showed no sign of returning to a closed position (Fig. 8) and turned brown after an average of 17 days. In both species some flowers had white, 1987] turgid corollas for longer than a month, although the stigmas were dull and blackened by this time and the stamens had altogether withered. DISCUSSION Floral longevity of individual flowers may be defined in terms of reproductive function as the period over which a flower is able to receive and/ or dispatch pollen. However, outside this period, flowers may perform functions not directly re- lated to their own reproductive success, as in the case of supplemental pollinator attraction pro- vided by late buds and older flowers. Thus, a more practical definition of floral longevity in terms of reproductive success of whole plants may be the period for which a flower is able to attract pollinators. For many plants, particularly tropical species, floral longevity appears to be a predetermined, endogenous characteristic with many such species having 1-day flowers (Dob- kin, 1984; Primack, 1985). In some such species, non-induced floral changes, which indicate that the flower is no longer reproductively functional, may occur before full senescence (indicated by abscission or wilting of petals). In others, flowers may be reproductively functional until full se- nescence (Gori, 1983). The second major pattern is provided by those species in which flower life may be curtailed by pollination (Kerner, 1902; Arditti, 1976; Stead & Moore, 1979; Devlin & Stephenson, 1984). It is important to note that for species in which floral changes are pollina- tion-induced, the potential flower life is still en- dogenously determined and the pollination-in- duced changes occur within this outer limit. Gori (1983) gnized and ized able data on five basic types of floral change that follow pollination or indicate the end of repro- ductive function before full senescence: color change, termination of odor and/or nectar pro- duction, change in flower orientation, collapse of flower parts, and corolla abscission. To this may be added flower closure as described here for Gentiana, and reported by Kerner (1902) for Mammillaria glochidiata. In fact, flower closure is likely to be the response in many species in which petals close regularly at night or in dull weather. Although floral changes may prevent or deter pollinator visits, the reproductive, func- tional end of flower life occurs when no viable pollen is available for dispatch and the stigma ceases to be receptive. In Gentiana serotina and G. saxosa, the end of reproductive flower life is indicated either by flower closure or the loss of WEBB & LITTLETON — GENTIANA 55 responsiveness of the flower to pollination. Once this occurs, pollen does not germinate on the stigma, and no seeds are produced. The begin- ning of flower life is marked by the corolla lobes opening to expose the dehisced anthers; all pollen is normally lost before the beginning of the fe- male phase, so pollen availability does not de- termine the end of flower life, although it may affect flower longevity if the male phase is pro- longed by a paucity of pollinator visits as in Lo- belia cardinalis (Devlin & Stephenson, 1985). When full senescence ofa flower finally occurs, floral changes as listed above may occur. In ad- dition, the whole flower may be aborted, even if pollination has occurred, especially in species that use flower abortion as a means of maternal reg- ulation (Lloyd, 1980; Bawa & Webb, 1984). Within a species, it is possible that the floral changes that occur following pollination may dif- fer from those of flowers that are never polli- nated. In both Gentiana saxosa and G. serotina, fresh female-phase flowers closed in response to pollination; in G. saxosa, but not G. serotina, the reaction of never-pollinated flowers differed in that flower closure did not occur and petals with- ered in a recurved position. However, the end of functional pollen and stigma presentation in never-pollinated flowers occurred well before the morphological change of flower closure in G. se- rotina and corolla wilting in G. saxosa. The per- sistence of these flowers well beyond their re- productively functional life may add to the overall floral display of the plant. Pollination-induced changes in flowers have been interpreted as signals that direct pollinators to unvisited flowers (Allen, 1898; Kerner, 1902; Arditti, 1976; Stead & Moore, 1979; Casper & La Pine, 1984) or help to conceal the developing seeds within pollinated flowers from predators (Allen, 1898), or simply minimize costs of flower maintenance by retaining the perianth no longer than is necessary (Kerner, 1902). Gori (1983) considered three aspects of the first alternative: avoidance of pollinator interference within pol- linated flowers, increasing the pollinator's for- aging efficiency and so increasing the residence time on the plant, and increasing ppilinatiog: ef- ficiency by restricting p productively fonction) flowers. In ‘the buo species of Gentiana studied here, corolla closure clearly signals that the flower is unavailable for visits; in fact it precludes visits, and the corolla also tightly enfolds the developing ovary until it is almost mature, making it less accessible to 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN predators. Field experiments, as described by Gori (1983) for Lupinus, would be necessary to determine the precise function of pollination-in- duced flower closure in Gentiana. In many in- sect-pollinated species, the end of flower life is signalled by corolla abscission and the devel- oping ovary is either inferior or enclosed within infolded bracts or calyx lobes as in Malvaviscus arboreus (Webb, 1984b). Corolla closure, rather than abscission, may be necessary to protect the ovary in Gentiana because the large superior ovary extends well beyond the small calyx lobes. In terms of natural selection, flower life for a particular species is likely to be a trade-off be- tween the cost of maintaining the flower in a receptive state and the probability that it has been pollinated. Thus, as originally suggested by Kerner (1902), flower life is likely to be longer for species that normally experience unpredict- able weather conditions allowing fewer suitable times for pollinator visits, for species with few, large flowers per plant, and for species that are obligately outcrossing. These suggestions are supported by data on flower life that show that many tropical plants, particularly those flowering in the lowlands d the dry season, have 1-day , 1984; Primack, 1985). Under contrast, plants of higher altitudes and of tem- perate areas have long-lived flowers, as might be expected when many days are unsuitable for flower visits. That many large orchid flowers with complex outcrossing mechanisms are long-lived is to be expected because they may have low rates of flower visitation. There are three strategies that might to some extent ameliorate the difficulty of unpredictable pollination. First, if flower life is pollination-de- endent, as in the two species of Gentiana de- ied here, then flower maintenance costs are minimized. There is, however, probably a cost in for pol- lination-induced floral changes (Gori, 1983). Second, under conditions unfavorable to polli- nators, autogamy may occur (Kerner, 1902; Fae- gri & van der Pijl, 1979); the evolution of autog- amy often involves loss of dichogamy or herkogamy, and also a reduction of flower life (Morin, 1983). In some species selfing may be delayed until an opportunity for outcrossing has been provided or may even be a direct response to unfavorable weather conditions. In Gentiana lineata, the flowers open fully in warm sunny weather suitable for insect pollination, but on [Vor. 74 wet or dull days the flowers open only partly so that the anthers dehisce directly onto the stigma (1984). T during rather unfavorable to pollination, or flowers may open only on suitable days with the floral parts protected during inclement weather (Kerner, 1902). When floral changes are pollination-induced, flowers are likely to be non-dichogamous or pro- tandrous because HOWEN: closure, are c sion, and other t rtail the time over which pollen can be pene (Lloyd & Webb, 1986). The pollination-induced flower closure described here for Gentiana can be effective only if flowers are protandrous; pro- togyny would preclude pollen presentation, and adichogamy would severely limit the male phase. Flower life may appear to be the simple result of proximate causes—especially of weather con- ditions and pollinator availability. However, the average flower life for a population, the particular response of flowers to pollination, and the cues used to determine the time of senescence, must all be the result of natural selection. Yet, the selection of these factors has seldom received the attention of reproductive biologists in spite of the important part they play in determining the success of plants as pollen or seed parents. LITERATURE CITED ALLEN, G. 1898. Flashlights on Nature. Grant Rich- ards, London. ARDITTI, J. 1976. Post-pollination phenomena in or- chid flowers. The Orchid Review 84: 261- OGAN & A. V. CHADWICK. 1973. Post- pollination phenomena in orchid flowers. IV. Ef- fects of ethylene. Amer. J. Bot. 60: 883-888. Bawa, K. S. & C. J. WEBB. 1984. Flower, fruit and seed abortion in tropical forest trees: implications for the evolution of paternal and maternal repro- ductive patterns. Amer. J. Bot. 71: 736-751. Casper, B. B. & T. R. LA Pine. 1984. Changes in corolla color and other floral characteristics in Cryptantha humilis (Boraginaceae): cues to dis courage pollinators? Evolution 38: 128-141. DevLin, B. & A. G. STEPHENSON. 1984. Factors that influence the duration of the staminate and pis- tillate phases of Lobelia cardinalis flowers. Bot. Gaz. 145: 323-328. Sex differential floral lon- gevity, nectar secretion, and pollinator foraging in a protandrous species 72: 303-310. D 1984 i d resident nectarivores FAEGRI, K. & L. VAN DER PUL. . Oecologia 64: 245-254. 1979. The Principles 1987] of Pollination Ecology, 3rd edition. Pergamon Press, Oxford. GonRi, D. F. 1983. Post-pollination phenomena and tive floral changes. In . Jones & R. J. Little (editors), Handbook of Experimental Pol- lination Biology. Van Nostrand Reinhold, New GOTTSBERGER, G. 1971. Colour change of petals in ow arboreus flowers. Acta Bot. Neerl. 20: —388. unre A. H., C. S. WHITEHEAD & A. M. KOFRANEK. 1984. Does pollination induce abscission of Cy- clamen flowers promoting ethylene produc- tion? Plant Physiol. 75: 1090-1093. KERNER VON MARILAUN, A. 1902. The Natural His- tory of Plants, odes F. W. Oliver (translator). Blackie & Son, Lon LrLovp, D. G. 1980. Sexual strategies in plants. I. An hypothesis of serial adjustment of maternal in- vestment during one reproductive session. New Phytol. 86: 69-79. . J. WEBB. 1986. Avoidance of interfer- ence between the presentation of pollen and stig- mas in angiosperms. I. Dichogamy. New Zeal. J. 2 1980. Flower senes- V. Thimann (editor), Senescence in Plants. CRC Aes Boca Raton, Florida. Morin, N. 1983. Ap of Githopsis (Campan- ulaceae). Syst. Bot. 8: 436-468. PHILIPSON, W. R. 1972. The generic status of the WEBB & LITTLETON—GENTIANA 57 southern hemisphere gentians. Adv. Pl. Morph. 1972: 417-422. PRIMACK, R. B. 1985. Longevity of individual flow- SCHOEN, D. The selection of cleistogamy and heteromorphic diaspores. Biol. J. Linn. . 23: 303-322. SIMPSON, M. J C. J. WEBB. 1980. Germination in udies on flower longevity in Digitalis. Pollination induced corolla abscission in Digitalis flowers. Planta 146: 409— 414. STRAUSS, M. S. & J. ARDITTI. 1984. Postpollination phenomena in orchid flowers. XII. Effects of pol- lination, emasculation, and auxin treatment on flowers of Cattleya porcia *Cannizaro' and the ros- tellum of Phalaenopsis. Bot. Gaz. 145: 43-49. THoMsoN, G. M. 1881. On the fertilization, etc., of New Zealand flowering plants. Trans. Proc. Ne Zeal. Inst. 13: 241-288. 984a. Constraints on the evolution of tematics. Academic Press Canada, Don Mills, On- tario. 1984b. Hummingbird pollination of Mal- vaviscus arboreus in Costa Rica. New Zeal. J. Bot. 22: 575—581 NOTES ON THE BREEDING SYSTEMS OF SACOILA LANCEOLATA (AUBLET) GARAY (ORCHIDACEAE)! PAUL M. CATLING? ABSTRACT To document breeding systems in the Mensa neotropical terrestrial orchid, Sacoila paula Nati pollination and seed development were studied in the field and in cultivated plants. In the s ern Florida study area, plants of var. /anceolata were not pollinated ; and hummingbird pollinators were apparently absent. When plants from ea were abundant in southern Ontario, huoyalagbini- ñollinstiop was observed on numerous occasions and the incidence of pollination was approximately 90%. Pollination experiments demonstrated a reliance on pollen vectors in plants of var. lanceolata from central Guyana, where a variety of hum- mingbird pollinators are available, but the plants of the same variety from southern Florida were found to be agamospermic. Examination of serial sections of ovaries in successive developmental o in portions of the tropical range. Plants of S. ' lanceolata var. paludi cola from southern Florida were found to self-pollinate. I ponnani indepe in southern Florida populations of S. lanceolata var. lanceolata and v dia and the apparent absence of races totally nul upon pollen vectors are aoc With pollinator-paucity Sacoila lanceolata (Aublet) Garay var. lanceo- lata is a reddish- to orange-yellow- or occasion- ally pale greenish-flowered, terrestrial orchid with a broad neotropical distribution extending from northern Florida and northern Mexico to north- ern Uruguay (Luer, 1972). be urpis: var. paludicola Luer is restricted to extreme southern Florida [the Siar eines Strand in Collier Co. (Luer, 1971a), Kendall Hammock in Dade Co. (pers. obs.), and from one location in the Everglades (McCartney, 1985)] and the Ca- ribbean region (based on examination of speci- mens at AM varieties are representative of the bird pollina- tion syndrome (Pijl & Dodson, 1966). Casual observations of seed development in a few cultivated plants that were maintained in the absence of any potential pollen vectors suggested that both var. /anceolata and var. paludicola may hk ll; + £ 4 4 +1 3l southern Florida. Since the breeding systems of tropical and subtropical terrestrial orchids are relatively poorly known, a study was undertaken to document the breeding systems of S. /anceo- lata. METHODS FIELD STUDIES OF POLLINATION IN SOUTHERN FLORIDA In order to gather information on natural pol- lination, populations of S. /anceolata var. lan- ceolata were observed for an overall total of 20 hours in southern Florida during mid-May 1978 and 1984. The presence of potential pollinators, including hummingbirds and large bees partic- ularly, was noted. Large bees are the regular pol- linators of some related species (Pijl & Dodson, 1966; Catling, 1983) Since pollination of orchid flowers involves removal of the pollinarium from the anther, it is possible to obtain an approximation of the incidence of pollination by recording pollinaria ! Living plants of Sacoila lanceolata var. paludicola ki provided by C. A. Luer, H. Brown, and G. Matous. C. A. Luer and V. R and N. H. Wil ? Agriculture Canada Brownell assisted with field work (University of nd C. A. Luer (Missouri Botanical Garde Useful criticisms were provided by S. C. H. Barrett n), N. R. Morin (Missouri Botanical Garden), k. liams (Univ. of Florida). W. Wojtas kindly anda with the wir pes wor Research Institute, Centr: rm, Ottawa, Ontario, Canada KIA 0C6. ANN. MISSOURI Bor. GARD. 74: 58-68. 1987. 1987] FIGURE 1. x. B, d pen Een ers, Immokalee, Collier Co., Florida, CATLING—BREEDING SYSTEMS OF SACOILA 59 1cm pou . S. lanceolata var. lanceolata.—A. Frontal view of flowers, ps oisi Co., —B. y la spike of cultivated plant ey central Guyana, South Am —D. ca, 13 May.—C. S. z var. e ned spike, Side of flow vated plant from Fahkahatchee Strand, Collier e. Florida, 15 Febru removal. This is more easily recorded than the presence of pollen tubes in the deteriorating col- umn of faded flowers. It is, however, only an approximation since pollinaria removal is oc- casionally (although apparently rarely) prevent- ed by malformation or drying out of the viscid- ium or by the difficulty of attaching to an existing excess load of pollinaria. The incidence of pol- lination in southern Florida, as determined by the absence of the pollinaria in faded flowers, 60 ANNALS OF THE MISSOURI BOTANICAL GARDEN was determined by examining 500 flowers from a total of 100 inflorescences representing eight locations. STUDIES OF POLLINATION OF TRANSPLANTS Plants from the Sarasota and Immokalee areas of southern Florida were observed for approxi- mately 33 hours after transplanting into a rural setting in southeastern Ontario, Canada, in late May 1984. The presence of potential pollinators was again recorded. The approximate incidence of pollination was determined by pollinaria re- moval (see above) in five groups of five experi- mental plants each, including a total of 429 flow- POLLINATION EXPERIMENTS Pollination experiments were performed on plants maintained in a glass house in Ottawa in April and May 1984 from which potential pol- linators (insects, hummingbirds) were excluded. Voucher specimens are contained in the Agri- culture Canada herbarium of the D. Research Institute in Ottawa, Ontario (DAO The experiments were performed on ra groups of plants: 1) 60 plants of S. /anceolata var. lan- ceolata, four from each of 15 localities in south- ern Florida distributed in a broad band, ca. 30 km wide, from Sarasota south to Immokalee; 2) three plants of S. /anceolata var. lanceolata from near Mahdia, central Guyana, South America; and 3) six plants of S. /anceolata var. paludicola including three from the Fahkahatchee Strand, Collier Co. and three from Kendall Hammock, ade Co., both in southern Florida Starting from the base of a spike, each se- pollinated, the fourth pollinated with pollen from a different flower on the same plant (geitonoga- mous pollination), and the fifth was pollinated with pollen from a flower on a different plant (cross-pollination). With the exception of the plants from Guyana, cross-pollination was car- ried out using plants from different localities. Each ovary was appropriately tagged to indicate the treatment of the corresponding flower: The Hull ber as the percentage of embryo sacs containing em- bryos. The latter value was estimated by emp- well [Vor. 74 tying the capsule and determining the presence or absence of embryos in the first 200 embryo sacs observed. This was followed by a scan of a few thousand to make certain that the value ob- tained was representative. If the value was con- sidered not representative, two more samples of 200 were scored and the average percentage was then recorded. ANATOMICAL STUDY Each of the three groups of plants used in the pollination experiments was studied anatomi- cally to dete ermine whether or not peeudogamy tre equiring was operating and to determine the method of agamospermy in plants producing seed without any pollination. Serial sections of ovaries were examined in successive developmental stages. Ovaries fixed and preserved in formalin-acetic acid-alcohol (FAA) were dehydrated, embedded in paraffin, sectioned, mounted, and stained us- ing a safranin-hematoxylin combination (Lillie, 1969; Jensen, 1958). Mature seeds of experi- mental plants were stained for examination using the differential stain technique described by Ow- czarzak (1952). The percentage of polyembryo- nic seed was determined in the same way as the percentage of embryo sacs containing embryos (see above). (asexual seed STUDY OF DRIED MATERIAL An attempt was made to determine the dis- tribution of breeding systems through the ex- amination of herbarium material from AMES, MICH, SEL, FTG, and DAO (acronyms from Holmgren et al., 1981). This is made possible, to a degree, by association of various morpho- logical features with a specific breeding system. Plants were considered agamospermic if they possessed any polyembryonic seed. Relatively small flower size was considered potentially in- dicative of autogamy (see Ornduff, 1969). Lack of expansion of some ovaries in fruiting material was considered suggestive of obligately cross- pollinated races. RESULTS FIELD STUDIES OF POLLINATION IN SOUTHERN FLORIDA In southern Florida, insects (including larger bees) were present but were not observed visiting the flowers of S. lanceolata var. lanceolata. No 1987] CATLING — BREEDING SYSTEMS OF SACOILA 61 E2. Ruby-throated yanmar (Archilochus colubris) pollinating Sacoila lanceolata var. lanceolata ida, 25 May FIG fede: som ota Co., Flori . Female probin ower directly through the floral tube. Female withdrawing with a pale yellowish Sdn attached to the tip of the bill. — C. Male probing a dower from the right side below the lip.—D. Male probing flower directly. hummingbirds were observed. There was no evi- dence of pollination, in the form of pollinarium removal, in any of the eight southern Florida locations. STUDIES OF POLLINATION OF TRANSPLANTS In southeastern Ontario, large insects includ- ing large bees were common, but they totally ignored the flowers. Ruby-throated Humming- birds (Archilochus colubris) were also common. pollinators. Females were observed visiting in- florescences on 25 separate occasions and males on 46. A visit usually involved probing three to five flowers on an inflorescence and visiting one to three inflorescences in a group of five. Visits were 15 minutes to two hours apart and were most regular in the morning between 9 A.M. and 11:30 A.M. Visits by one or more females (i.e., female visits) involved a direct probing of the flowers (Figs. 2A, 3A), the pollinia becoming at- tached, by the elongate cushion-type viscidium that sheaths the rostellum (Fig. 3C, D, E, and Greenwood, 1982), to the distal portion of the bill (Figs. 2B, 3B). Male visits involved probing from the side and below the lip approximately 50% of the time (Fig. 2C, D). Probably as a con- sequence of this “robbing,” male visits resulted 5 experimental plants, which were observed being pollinated by the hummingbirds, the incidence of pollination was 90% (1.e., 38 of 429 flowers). 62 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 oe | 10 mm | FIGURE 3. a flower of Sacoila (alent see var. lanceolata. — A. B tw Column viewed from below viewed from Co., Florida POLLINATION EXPERIMENTS Pollination was clearly necessary for seed de- in the absence of pollination. Pollination at any time is followed by fading in two or three days. In S. /anceolata var. lanceolata from southern Florida, seed developed regardless of pollinar- ium removal (Table 1), and these plants were clearly autonomously agamospermic. The flow- ers lasted 5-15 days in the absence of pollination, but pollination resulted in fading within two days. Seed development occurred in undisturbed flowers of S. /anceolata var. paludicola, but not when the pollinarium was removed from the flower at an early stage (Table 1), indicating it | the di (darkenedi = C D E iagrams of a pM eager (Archilochus colubris) ia aen pollinia from ird with bill inserted in flower, prior pward movement —B. Bird with pollinia attached near tip | of bill. — afte below showing viscidium (darkened) and pollinia. All diagrams based on material from Sarasota autogamy (self-pollination) or pseudogamy (pol- lination-induced agamospermy) in this variety. Self-pollination occurred after the flowers had en open for one or two days. The flowers are short-lived, lasting for approximately five days. The mechanism of self-pollination is a simple lengths of the pollinia and the basal part of the column (Fig. 4A). ANATOMICAL STUDY The Guyana plants of var. lanceolata and plants of var. paludicola from southern Florida dem- onstrated a normal sequence of pollination, pol- 1987] CATLING — BREEDING SYSTEMS OF SACOILA 63 TABLE 1. Pollination experiments with Sacoila lanceolata. No./96 No./% No. of No. of Ovaries Ovaries % Sacs with Treatment Plants Ovaries Expanded* with Seed* Embryos* S. lanceolata var. lanceolata—Guyana Undisturbed 3 6 0/0 0/0 0 Pollen removed 3 6 0/0 0/0 0 | botes 3 6 6/100 4/66 50-100 Geitonogamous-pollination 3 6 6/100 4/66 60-100 Batic 3 6 6/100 4/66 60-100 S. lanceolata var. lanceolata—South Florida Undisturbed 60 200 200/100 200/100 90-100 Pollen removed 60 180 180/100 180/100 97-100 Self-pollination 60 180 180/100 180/100 96-100 Geitonogamous-pollination 60 180 180/100 180/100 97-100 Cross-pollination 60 180 180/100 180/100 97-100 S. lanceolata var. paludicola—South Florida Undisturbed 6 25 25/10 25/10 95-100 ollen removed 6 12 0/0 0/0 0 Self-pollination 6 12 12/100 12/100 95-100 Geitonogamous- eee 6 12 12/100 12/100 95-100 Cross-pollinatio 6 12 12/100 12/100 95-100 * As a percentage of the total ovaries tested. len tube growth, fertilization, and embryo de- velopment (see Catling, 1982). In plants of var. lanceolata from Florida, the gametophyte de- generated and one or more cells of the inner in- tegument proliferated to become embryos. The proliferation of these cells was indicated by the more deeply staining protoplasts, relatively large nuclei, and well-developed nucleoli (Fig. 6). Pro- liferation had initiated in the ovaries of flowers open for five days and may have initiated earlier but was not evident in mature ovaries with buds on the point of opening. Using ovaries of in- creasing age within an inflorescence, it was pos- sible to trace the darkly staining and proliferating micropylar integument to embryos in mature seeds. The seed resulting from adventitious em- bryony (Fig. 7C) differs from that resulting from fertilization (Fig. 7A, B) in having a proportion of the seeds polyembryonic (range 28—95%, mean 71.3%, standard deviation 17, based on 15 flow- ers representing 15 different southern Florida populations). ive capsules from flowers cross-pollinated on (range 56-94%, mean 73.5%, standard deviation 17.9; range 30-84%, mean 57.2%, standard de- viation 23.5, respectively). STUDY OF DRIED MATERIAL Many more collections and some supporting fieldwork would be necessary to allow a reliable assessment of the distribution of different breed- ing systems throughout the range of S. lanceo- lata. However, an appraisal of the extent to which the breeding systems of the southern Florida que p ible with the ma- terial aiid: Agamospermy, as evidenced by polyembryon- ic seed, was found in all fruiting plants from Florida and in some plants from Guatemala, Be- lize (British Honduras), and Bolivia. In some cases this was associated with non-opening (i.e cleistapogamous, see Sheviak, 1982) flowers (for example, district of Peten, Guatemala, 12 June 1933, C. L. Lundell 3896, MICH). The occur- rence of agamospermy within the main tropical range is not clearly associated with relatively high altitude, a situation that involves some of the same effects as high latitude, and the sample is not large enough or sufficiently well documented to reliably reflect a trend. Relatively small flowers (lateral sepals 14—17 mm long), suggestive of self-pollination, were characteristic of plants from a number of Carib- bean islands (for example, Bahamas, St. Vincent, based on specimens at AMES and FTG). Limited ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Ry L 0.5mm | | 0.5mm | A. Column of Sacoila lanceolata var. lanceolata viewed diy below (left) and diagrams of cross- rida. — B. sections (right) based on material from Sarasota Co., Flo n of Sacoila t var. paludicola viewed from below and diagram of cross-section (right) of flowers esse sa from Collier Co., Florida. C,-C,, cross-sections illustrated on the right-hand side of the figure: P, pollinarium; PG, pollen ebrius S, stigmatic surface; V, viscidium. fruit set, suggestive of obligate cross-pollination, was restricted to Costa Rica and northern South America. DISCUSSION amospermy is uncommon in the orchid y reported only in and wari, 1952; Ca ding: 1982), Prasophyllum (Bates, 1984a), Microtus (Bates, 1984b), Paracaleana (Jones, 1977), Dactylorhiza (Dressler, 1981), Zygopetalum (Dressler, 1981), and Pterygodium (Schelpe, 1970). Autogamy is much more prev- alent than agamospermy in vascular plants gen- erally and in the orchid family, where it has been reported in over 60 genera (Catling, unpubl. data). In S. lanceolata var. lanceolata, it is not known whether agamospermy is facultative. Although there is no evidence of seed resulting from cross- fertilization in terms ofa lower incidence of poly- 1987] e 0.5 mm CATLING — BREEDING SYSTEMS OF SACOILA 65 J » ^ e Cross-section of column of S. /anceolata var. paludicola, from Collier Co., Florida. This section IGU ] represents the position C,—C, in Figure 4B. AC, anther cap; P, pollinarium; PG, pollen germinating; S, stigmatic surface. embryony, this may have been the result of cross- pollinations using genetically identical individ- uals with some degree of self-incompatibility. Al- though the crossings involved plants from dif- ferent localities in southern Florida, it is conceivable that the different populations had the same agamospermic origin and are geneti- cally identical. Pollination by hummingbirds in S. lanceolata is to be expected on the basis of various floral features such as the tubular, horizontally orient- ed, reddish flowers without internal marking and lacking odor (Austin, 1975; Pijl & Dodson, 1966), but no previous observations of hummingbird pollination have been reported. The only feature of characteristically bird-pollinated orchid flow- ers that is lacking is a dark pollinarium (Dressler, 1971), that of S. /anceolata being pale yellow. Although the observation of hummingbird-pol- lination reported here involved transplanted and cultivated plants, there seems to be no reason to doubt that these observations are indicative of the pollination mechanism in various other sit- uations where one or more species of humming- birds are available. The suggestion of a lack of natural pollinators in southern Florida pineland is of interest insofar as it may provide an explanation for the polli- nator-independent breeding systems. Although self. nollination and iated with pollinator-paucity and colonization of new territory (for example, Allard, 1965; Jain, 1976; Lloyd, 1978) the actual absence of pollinators has not often been well established (for example, Kevan, 1972); nor has “newness” been quanti- fied. In the southern Florida pinelands in May, Fr- ythrina herbacea and Ipomea microdactyla are among the few characteristic hummingbird blos- soms available, and these species were not ob- served near colonies of var. /anceolata. Only one species of hummingbird (A. colubris) is present, 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURE 6. A, B. Cross-sections of ovules of S. lanceolata var. lanceolata from Sarasota Co., Florida.—C. Longitudinal section. g — degenerate gametophyte, ii — proliferating inner integument, oi — outer integument. Material from Sarasota Co., Florida. and by April and May it has already migrated localized (Sprunt, 1954), and there is very little evidence of breeding in southern Florida (Rob- ertson, 1974). Thus there is relatively little to attract hummingbirds to the vicinity of many SS. lanceolata colonies in southern Florida during the May blooming period, and there is only one potential hummingbird pollinator, which is ap- parently not abundant. The var. paludicola also exists in the absence of a diverse hummingbird- pollinated flora in southern Florida, although a few bromeliads exhibiting the syndrome do flow- er simultaneously in the hammocks. The situa- tion over much of the range of var. /anceolata is quite different since many different humming- bird species are available and there is a diversity and continuity of hummingbird-pollinated blos- soms on a year-round basis (Grant & Grant, 1968). Populations of five other tropical orchids (£ p- that promotes self-pollination. This is in contrast to parts of (or all of) the warmer continental neo- tropics where these same species have a column structure adapted to cross-pollination, or where this latter column structure is at least predomi- nant (Luer, 1971b, 1972; Pijl & Dodson, 1966; pers. obs.). Pollinators are not clearly lacking in the case of these species. It is possible that con- tinual near extinctions due to frost (Singleton, 1936; Stowers & Le Vasseur, 1983), hurricanes (Craighead & Gilbert, 1962; Alexander, 1967) and sea-level fluctuations (Fairbridge, 1974; 1987] CATLING — BREEDING SYSTEMS OF SACOILA 67 FIGURE 7. C. Seed of S. lanceolata var. lanceolata from Sarasota Co., Flo Long, 1974) resulted in a strong selection for colonizing ability, regardless of pollinator-avail- nator-paucity is considered to provide an ade- quate explanation for their pollinator-indepen- dent breeding systems in southern Florida. LITERATURE CITED ALEXANDER, T. R. 1967. Effect of hurricane Betsy on the southeastern Everglades. Florida Acad. Sci Quart. J. 30: 10- ALLARD, R. W. 1965. Genetic systems associated with OHA species. Pp. 49- 76 in H. G. Baker & G. C. Stebbins A, B. Seed of Sacoila lanceolata var. paludicola. — A. Collier Co., Florida. — B. Dade Co., Florida. — rida. (editors), The Genetics of Colonizing Species. Ac- ademic Press, New York. AUSTIN, D. F. 1975. Bird flowers in the eastern United States. Florida Scientist 38: 1-12 BATES, R. 1984a. Apomixy in four south Australian species of Prasophyllum R.Br. J. Native Orchid Soc. South Austr. 8(4): 38-39 1984b. The genus Microtus R.Br. (Orchida- ceae): a taxonomic revision with notes on biology. J. Adelaide Bot. Gard. 7(1): 45-89 CATLING, P. M. 1982. Breeding systems of north- eastern North American Spiranthes (Orchida- ceae). ae J. Bot. 60(1 2): ador 3039. . Pollination of northeastern North Ame erican uem e (Orchi pres Canad. J. Bot. 3 V.C. GILBERT. 1962. The effects orida. DRESSLER, R. L. TL Dark pollinia j in humming- 68 ANNALS OF THE MISSOURI BOTANICAL GARDEN bird-pollinated orchids, or do hummingbirds suf- fer from strabismus? Amer. Naturalist 105: 80- 83 ———, The Orchids, Natural History and Classification. Harvard Univ. Press, Cambridge, Massachusetts. FAIRBRIDGE, R. W. 1974. The holocene sea-level rec- ord in south Florida. Pp. 223-232 in P. J. Gleason (editor), Environments of South Florida: Present and Past. Miami Geological Society Mem GRANT, K. A. & V. GRANT. 1968. Hummingbirds and Their Flowers. Columbia Univ. Press, New GREENW he = 982. Viscidium types in the ani ou (Mex.) 8(2): 283-310. HOLMGREN, P UKEN & E. K. SCHOFIELD. 1981. Index Herbariorum, ~ 2 The Herbaria of the World. Dr. W. Junk, B JAIN, S. K. 1976. Theevolution "e NER plants. Annual Rev. Ecol. JENSEN, W. A. Bo tanical Mini NM Prin- ciples and Practice. W. H. Freeman and Co., San Francisco. 1977. Miscellaneous notes on Australian to synonymy. The Orchadian 5: 126-1 128 KEvAN, P. G. 1972. Insect eta of high arctic flowers. J. Ecol. ~~ 83 LirLig, R. D. H. J. Fai S Biological Stains. The Williams and Wilkins Co., Baltimore Lrovp, D. G. 78. Demographic factors and self- fertilization in plants. Pp. 67-88 in O. T. Solbrig (editor), Demography and the Dynamics of Plant Populations. Blackwell, Oxford Lona, R. W Ç southern Florida. Pp. 28—36 in P. J. Gleason (ed- itor), Environments of South Florida: Present and Past. Miami Geological Society Mem. 2 Origin of the vascular flora of [VoL. 74 d C. A. 1971a. A variety of Spiranthes lanceolata n Florida. Florida Orchidist 14: 19. 1971b. Abnormal development of the ther—a report of two cases. Florida Orchidist 14: 26-29. 72. The Native Orchids of Florida. New York Botanical Garden, Bronx, New York MAHESHWARI, . Polyembryony in angio- sperms. Palaeo- botanist (Lucknow) 1: 319-329. McCartney, C. 1985. Orchids of Florida—the or- chids of Everglades National Park— 1. Amer. Or- chid Soc. Bull. 54(3): 265-2 ORNDuFF, R. 1969. Reproductive biology in relation to ipei Taxon 18: 121-133. A. OWCZARZAK, 952. Arapid method for mounting rige BUDE p Technol. 27: 249-251. Pu, L. C. 1966. Orchid Flowers: Their Pollination and Evolution. Univ. of Miami Press, Coral Gables, Florida. ROBERTSON, R. 1974. The southern Florida avifauna. Pp. 414-4 52 in P. J. Gleason (editor), Environments of South dine: Present and Past. Miami Geological Soc SCHELPE, E. A. 1970. Seif pollination, aie ain d African Orchid Society Journal, March: 9- D SHEVIAK, C. J. 1982. Biosystematic study of the Spi- ranthes cernua complex. New York State Museum (Albany) Bull. 448, 73 pp. SINGLETON, G. 1936. The freeze of 1934. Florida Acad. Sc. Proc. 1: 23- SPRUNT, A., JR. 1954. Florida Bird Life. Coward- McCann Inc. and National Audubon Society, New Yor STOWERS, D M., JR. & M. LEVASSEUR. 1983. The Florida freeze of 13 January 1981: an impact study of west-central Florida. Florida Scientist 46: 72- 83 FLOWER AND FRUIT BIOLOGY IN SOUTHERN SPANISH MEDITERRANEAN SHRUBLANDS! JAVIER HERRERA? ABSTRACT Flower and fruit biology was studied in a coastal, southern Spanish serok community composed of 30 plant taxa. Data on breeding systems; rewards offered to vectors; flower, fruit, a xa in the community have insect- pollinated hermaphroditic rphology. Dioecious species are relatively well represented fruiting intensities are reported. Most ta flowers that are largely unspecialized in mo nd seed sizes; and (27% of the total), as are vertebrate-dispersed species (43%). Bagging experiments demonstrated that apap were required for maximum fruit production, but the existence of ee rent systems was not t indicia was investigated, it was found that sprouting taxa ted. When the relationship between fruiting intensity and the words perform vegetative had, on average, lower fruit production than those that were unable to sprout. Low fruit production is Polen a relation to reproductive allocation trade-offs. Mediterranean-type vegetation has been the subject of research for investigators who wish to h 1 } in OP(YOTra he ically distant areas with similar environmental factors (Specht, 1969; Mooney & Dunn, 1970; Cody & Mooney, 1978). Also, a great emphasis has been put upon plant development and the adaptive features of plants in this hia seasonal climate oe & Parsons, 1973; oney et al., 1974; Kummerov, 1983). Cares lit- tle is kapun. however, about other characteris- ticsin the biology of mediterranean species, such as their reproductive biology. The paucity of in- formation is particularly noticeable with respect to plants living in the Mediterranean region itself (but see C. M. Herrera, 1981, 1984; Jordano, 1982, 1984, for plant-frugivorous bird relation- ships, and J. Herrera, 1985, for nectar secretion patterns in scrub). Some information is available for mediterranean areas in America, Australia, and South Africa (for example, Moldenke, 1975; Specht et al., 1981; Kruger, 1981). But in spite of this, our present knowledge of the reproduc- tive biology of mediterranean shrublands is low compared to our knowledge of tropical (for ex- ample, Frankie et al., 1974, 1983; Heithaus, 1974; Bawa, 1979; Bawa & Opler, 1975; Opler et al., 1980) and temperate plant communities (for ex- ample, Mosquin, 1971; Kevan, 1972; Pojar, 1974; Reader, 1977; Primack, 1983). This paper presents part of a study designed to investigate the reproductive biology of a southern Spanish sclerophyllous scrub commu- nity. Flower and fruit features in a number of taxa are used to elucidate reproductive patterns. The relationship between sprouting behavior (the production of new stems from established Kd zomes, lignotubers, or burls; James, 1984) an pollination—reproduction variables is also inves- tigated. Pollination relationships at the com- munity level will be dealt with elsewhere (J. Her- rera, in prep.). STUDY AREA AND METHODS This study was conducted in the Donana Bi- ological Reserve (Donana National Park, Spain). The reserve is located on the Atlantic coast of southwestern Spain, in an area with a mediter- ranean-type climate where the vegetation is com- posed mainly of mediterranean sclerophyllous shrublands with some planted pine woods. An- nual precipitation averages 537 mm. Mean an- nual temperature is 16°C, January being the cool- est month (9.8°C) and July the hottest (24.6°C). Temperatures rarely descend below zero, and summer drought covers five months on average (May through September). The soil is sandy, and maximum elevation above sea level is 3 Basically the vegetation encompasses two types t of the underground water table is relatively near 'This study received financial support from the Spanish Comision Asesora de Investigacion Cientifica y Técnica (CAICYT), through a grant (82/264) to Salvador Talavera (Departamento de Botanica, Facultad de Biologia, Universidad de iii i ae Dr. P. E. Gibbs for correcting the English, and two reviewers for helpful comments on the ma ? Departamento de Bout jm de Biologia, 41080 Sevilla, Spain. ANN. Missouni Bor. GARD. 74: 69—78. 1987. 70 ANNALS OF THE MISSOURI BOTANICAL GARDEN soil surface, the scrub becomes dominated by hygrophytic species (heath). Otherwise, a highly xerophytic scrub vegetation is found (see Allier et al., 1974; Rivas-Martinez et al., 1980). The most representative species, both in hygrophytic and xerophytic scrub, were selected for study giv- ing a total of 30 taxa. From December 1982 through March 1984 the study plot was visited weekly, and data on flowering phenology, pol- lination relationships, and flower-fruit biology of the species were gathered. Here I report only on aspects of flower and fruit biology. Individual plants or branches were marked be- fore flowering, and their area estimated from maximum and minimum diameters. Flowers were counted every week during the time of study, nd the maximum numbers registered for the plants of a species were averaged to give an es- timate of flower production per unit area. Fresh samples of flowers and fruits of all taxa were taken to determine the most outstanding external features, such as dimensions, color, and sex. Ten to 20 flowers of each species were mea- sured (length, maximum and minimum diame- ters), and 20 to 500 complete flowers without their pedicels were air dried and weighed. The maximum dimension of a flower was found to be directly correlated with its dry weight (r, = 0.806, N = 30, P < 0.001). Therefore dry weight has been used subsequently as an estimate of flower size, since it is less dependent on the ar- termined from external features and visitor cen- suses (J. Herrera, in prep.). Flower forms were referred to those of Faegri and van der Pijl (1979). Pollen production and the number of ovules in the ovary of each flower were determined for 10 to 15 flowers per plant species. Ovules were counted under a dissecting microscope. Pollen production was assessed in the same flowers by macerating one or two anthers in a known vol- ume of detergent-safranine solution and count- ing the number of grains in 10 replicates of 5 ul. Pollen-ovule ratios of hermaphroditic species were then compared with those given by Cruden (1977). To investigate nectar secretion, flowers were observed in the field and in the laboratory under a dissecting microscope. In doubtful cases, the arrival of insect visitors to flowers was precluded by bagging branches with white nylon mesh; flowers were examined after 24 hours and, when possible, the accumulated nectar was quantified. [VoL. 74 The volume per flower was measured by the length of the column, in mm, inside 5 ul micro- pipettes. Concentration, on a weight : weight ba- sis, was measured in two ATAGO refractometers (models N1 and N2) compensated to ambient temperature. The milligrams of nectar sugar plus diluted solids (Inouye et al., 1980) were com- puted by the product of volume x concentration (Bolten et al., 1979) Percent fruit production in open pollinated flowers of the species was estimated from flower and ripe fruit counts on marked branches. In the same individual plants that were used to estimate percent fruiting, some branches were also bagged with white nylon mesh (or with glassine paper bags in anemophilous taxa) to determine fruiting intensities when pollinators were excluded. Bag- ging experiments were not carried out with dioe- cious taxa. Samples of fruit were collected and oven dried to a constant weight, and the number of seeds, along with the weights ofthe whole fruit, all seeds, and that of an individual seed were averaged and recorded. Notes on fruit and flower predation were also taken and, when possible, the agents responsible were identified. The ability to perform vegetative regeneration after complete destruction of aboveground bio- mass (sprouting) was assessed for every species. PAS on ou of these Species w were tns estudy area on plants damaged by fire, M or human disturbance during the years 1982 through 1985 RESULTS FLOWER BIOLOGY The names of the species studied, together with their most distinctive floral features, are sum- marized in Table 1. Our set of mediterranean plants consists of 30 taxa in 17 families and 25 genera. Most families contribute one or two species, with only Cistaceae, Leguminosae, La- biatae, and Ericaceae being relatively well rep- resented (5, 4, 3, and 3 species, respectively). Pollination by insect vectors is dominant in the community, where entomophilous species account for more than 80% of the total (Table 1). Only five species s rely on wind for pollen dis- dioecious species are relatively well represented (27%). Thymus tomentosus is the only species with a different sexual condition (gynodioecy). 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JO 1431M AIP ‘MA ‘WO 1940] ‘JA ‘Pow uoneuripod ‘Wd :urojsÁs 3urpoo1q *Sg) sds qnuos 0ç Jo saInquNe [eJOLJ “| 114V L 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN | Oo S. 2^ a e m. ° š, > = se. "Ka 2 c 2r ° e, > 4 A S1 á u. 7 0 FLOWER DRY wees wm El. Relationship een the weight of an individual flower and the mean number of flowers per square meter in peak blo ien plants. Triangles — sprouting taxa; circles = nonsprouting taxa. Flower sizes (as dry weight of the complete flow- er) in the sample range between 2.2 mg for the female flowers of Pistacia lentiscus and 65.5 mg for Cistus salvifolius. More than 5096 of the taxa have flowers of 5 mg or less in dry weight. Floral dry weight is not independent of the breeding system in this set of species. If taxa are segregated in two categories, less or more than 5 mg (a value that is near the median in the frequency distri- bution), dioecious taxa predominate in the lower class (8 out of 9), whereas hermaphroditic taxa are evenly distributed among the two classes (10 and 11, respectively). The differences are statis- tically significant (G = 5.04, df = 1, P < 0.025). The predominant flower morphology is of the dishbowl type (Faegri & van der Pijl, 1979), which is present in the Cistaceae and also in all the entomophilous dioecious taxa. Species in the La- biatae have short, tubular corollas; while the le- gumes present typical flag blossoms. The only long-tubed flowers are those of Lonicera pericly- menum. Generally speaking, flowers are small and/or have corollas that impose little or no re- striction on floral visitors (except for L. pericly- menum and the legumes). All species in the Le- guminosae and three in the Cistaceae have yellow flowers, which is the most common color (nine of total); whereas six species have white corollas. Many of these yellow- or white-flowered taxa offer pollen as the main reward to pollinators (Table 1). There are five species with pink flow- ers, which provide predominantly nectar as the reward. [VoL. 74 Among entomophilous taxa, pollen and nectar are offered by approximately the same number of species (12 and 13, respectively). However, since wind-pollinated taxa are also occasionally used as a pollen source by insects (pers. obs.), this food material is more readily available in our community than is nectar, at least in terms of plant taxa. Furthermore, we have succeeded in quantifying the secretion in only six out of 13 species in which nectar is the main reward. These species were Daphne gnidium (0.17 mg sugar/ flower per 24 hr.), Erica ciliaris (0.08 mg), La- vandula stoechas (0.15 mg), Lonicera pericly- menum (2.00 mg), Rosmarinus officinalis (0.20 mg), and Rubus ulmifolius (1.20 mg). In the re- maining seven species the yield of nectar sugar per flower per 24 hr. was too scarce to be quan- tified with our method (presumably less than 0.08 mg). Maximum concentration values ranged be- tween 9.5% (Erica ciliaris) and 60% (Rubus ul- mifoliusy, volumes per flower on a daily basis ranged between 0.9 ul (Lavandula stoechas, Daphne gnidium) and 10 ul (Lonicera pericly- enum). Pollen yield of the individual flowers is given in Table 1. The lowest is that of Armeria velutina (10? grains) and the highest that of Myrtus com- munis (599 x 10?), which has insect-pollinated, nectarless flowers of the brush type common in the Myrtaceae. Since pollen production is not independent of flower size, a more correct esti- mate of staminate effort on a per flower basis would be the number of pollen grains produced per milligram of flower dry weight (Relative Sta- minate Effort, RSE). Defined in this way, RSE is minimum for entomophilous taxa such as Ar- meria velutina (500) and vain periclymenum (524). The former has dimorphic pollen grains and stigmas, and the latter a the most spe- cialized (long-tubed, highly nectariferous) flowers in the sample. Maximum values of RSE are found in anemophilous species such as Corema album (2.3 x 10°) and Erica scoparia (120 x 10). Mean RSEi y higher for than for entomophilous ones (U — 121, N = 5, 25, P < 0.001). Values of pollen-ovule ratios for hermaphro- ditic species are given in Table 1. In most taxa this ratio is greater than 2,000 and thus referable to the allogamous class of Cruden (1977). Only three species have pollen-ovule ratios lower than ,000. Mean number of flowers for plants in peak bloom are also shown in Table 1. A highly sig- nificant relationship exists between flower num- 1987] HERRERA—FLOWER & FRUIT BIOLOGY TABLE 2. Fruit and seed characteristics for 29 scrub species. Numbers in parentheses are sample sizes. DE. Mean Dry Weight (mg) Pre- Species Type! of Seeds Whole Fruit All Seeds Individual Seed dation Armeria velutina D l 3 (50) 1 l (50) E Asparagus aphyllus F 1.2 46 (100) 23 19 (122) * Calluna vulgaris D 3.6 (100) 0.2 0.1 (325) = Chamaerops humilis F l 1,364 (18) 781 781 (18) + Cistus libanotis D 22.3 59 (20) 23 1 (445) + Cistus salvifolius D 51 137 (12) 53 1 (612) + F 3 57 (30) 35 12 (90) = Cytisus grandiflorus D 6.9 214 (20) 39 6 (137) = Daphne gnidium F l 18 (75) 8 8 (75) = Erica ciliaris D 17.9 5 (80) 0.2 0.01 (1,150) + Erica scopari D 6.5 1 (60) 0.1 0.02 (1,000) + Halimium commutatum D 2.6 33 (20) 14 (52) + Halimium halimifolium D 25.7 59 (20) 15 0.6 (514) = Helianthemum croceum D 10.3 34 (15) 15 1 (155) + Helichrysum picardii D l 0.6 (750) 0.6 0.6 (750) on Lavandula stoechas D 2.3 4 (50) l 0.6 (114) + Lonicera periclymenum F 2.3 52 (30) 15 7 (30) = Myrtus communis F 5.7 108 (75) 56 10 (430) = Osyris alba F l 157 (30) 90 90 (30) + Osyris quadripart F l 131 (75) 73 73 (75) - Phillyrea angestfola F l 35 (100) 12 12 (100) — Pistacia lent F l 22 (80) 13 13 (80) = Rhamnus oem F 1.6 63 (30) 26 16 (30) = Rosmarinus officinalis D 2.9 4 (20) 2 0.6 (59) + Rubus ulmifolius F 40.4 213 (30) 87 2 (30) Smilax asper. F 1.4 78 (100) 51 36 (140) — 2. i genistoides D 2.5 59 (15) 13 5 (38) + Ulex m D 2.1 22 (20) 7 3 (35) + Ulex xn an D 2 47 (10) 12 6 (20) + ' D, dry fruits; F, fleshy, vertebrate-ingested fruits. ber and flower size (dry weight); this relationship can be easily appreciated in Figure | (r= —0.826, = 27, P < 0.001). T of flowers by insect predators, mostly noctuid larvae, coleopteran larvae, and oth unidentified insects, was observed in only four species (Table 1). O + FRUIT BIOLOGY Table 2 summarizes various characteristics of the fruits in our set of mediterranean plant species. Fruit production in Thymus tomentosus was so sparse that we were unable to gather a reasonable sample of fruits and seeds; this species has there- fore been excluded from the analyses. According to the way in which their seeds are dispersed our taxa are easily divided in two groups: those whose seeds are dispersed by vertebrates (13 taxa) and those whose seeds are dispersed by the wind or in a largely passive way (16 taxa). The former have fleshy fruits, while the latter have dry fruits (mostly capsules, legumes, or achenes). Seed col- lecting by ants has been observed in a few taxa, but data are not conclusive enough to recognize a third (ant- -dispersed) class. Most vertebrate- number of seeds per fruit ranges between l and 51, and the fruit size (fruit dry weight) between 0.6 mg (Helichrysum picardii) and 1,364 mg (Chamaerops humilis). The lightest seeds are those of Erica ciliaris (2 x 10? mg) and the heaviest those of Chamaerops humilis (781 mg). Predation of fruits, mainly by Curculionidae, lar- vae of Tortricidae, Noctuidae, and parasitic hy- menopterans, is far more common than flower predation (55% and 13% of species pectively). Damage by predators was observed more fre- quently on dry than on fleshy fruits (G — 4.02, df = 1, P < 0.05) 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE 3. Results of independence tests performed between the type of fruit (dry or fleshy) and other vari- ables of flowers and fruits. Class intervals considered were weight of fruit, less than 20 mg, 20-40 mg, 40- 60 mg, or more than 60 mg; weight of the paneer seed, less than 10 mg, or more than 10 mg; num seeds per fruit, less than 2, 2-4, or more than 4. Independence of G df P i (hermaphrodit- oec 11.72 1 «0.001 Weight ier 804 3 «0.05 Weight of cu seed 23.32 | «0.001 Seeds per fruit 13.88 2 <0.001 G, value of the statistic in the G test of independence; df, degrees of freedom. Results of tests for the independence of fruit type and a few variables of flowers and fruits are given in Table 3. There is a significant tendency for species with hermaphroditic flowers to pro- duce dry fruits, and for species with unisexual flowers to produce fleshy fruits. The indepen- dence hypothesis of fruit type and breeding sys- tem (Bawa, 1980) is thus rejected (Table 3). The only gynodioecious taxon in the sample has been included (conservatively) in the dioecious class for this analysis. The independence hypothesis for the type of fruit and the variables weight of fruit, weight of an individual seed, and seeds per fruit are all rejected. In fact, fleshy fruits tend to be, on average, heavier than dry ones (U = 185.5, N = 16, 13, P < 0.05) and to have fewer and heavier seeds (U = 203, N = 16, 13, P < 0.001). Note that we are reporting dry weights, so that differences are not due to the high water content of fleshy, vertebrate-ingested fruits. Fruiting intensities for 28 of the studied species can be seen in Table 4. Data are not available for two dioecious species, Corema album and Rhamnus lycioides. Fruit production after exclu- sion of pollinators is also reported for hermaph- roditic taxa. Percent of fruit production in open pollinated flowers ranged between 2% for Thy- mus tomentosus and Daphne gnidium, and 9296 for Halimium commutatum. Bagging of flowers in most cases had a clear effect of decreasing fruit production—to levels as low as 0-196 in 15 species. There was just one taxon that was un- affected by exclusion of pollinators (Daphne gni- dium, 296 fruit production under both treat- ments). Some fruit was produced (10—3196) in bagged flowers of a few taxa (Calluna vulgaris, [Vor. 74 Erica scoparia, Myrtus communis, and Rosma- rinus officinalis), but even in such cases polli- nating vectors were needed for fruiting to arrive at its maximum. REPRODUCTIVE TRADE-OFFS Figure 2 shows the relationship between per- cent fruiting and the quotient of the fruit dry weight to the flower dry weight (fruit : flower size ratio hereafter), which indicates about how high the increase in size is from the first reproductive unit to the second. This variable will be em- ployed below to investigate some patterns of re- productive allocation in our set of species. A highly significant negative relationship exists be- tween percent fruit production and fruit : flower size ratio (r = —0.6780, N = 26, P « 0.001, log transformed data). Thus species in which there is a great increase in dry weight during the tran- sition from flower to fruit are those with the low- est percent fruiting, whereas species in which this process involves little gain in weight have rela- tively greater fruiting success. Species known to perform vegetative regen- eration (sprouting) are PEPE, from non- sprouting ones in Table 4, and their respective distribution in the relationship defined by per- cent fruit production and fruit: flower size ratio is Shown in Figure 2. Sprouting taxa tend to have low values of fruiting and high values of fruit: flower size ratio. Mean percent fruit production is not, however, significantly different for sprout ing and nonsprouting taxa (U = 109, = 14, 12, P > 0.2); differences in mean dry weight increase from flower to fruit are weakly signifi- cant (U = 117.5, P > 0.05). Species in the Eri- caceae lower the coherence of the sprouting group, since they are well-known sprouters but have relatively low fruit: flower size ratios. If the Er- icaceae are removed, the sprouting group is en- tirely composed of taxa with few-seeded, verte- brate-ingested fruits; with the exclusion of ericaceous taxa, the nonsprouting group overlaps entirely with the dry fruit group, and the sprout- ing group does likewise with the fleshy fruit group. Differences in mean percent fruit production and fruit: flower size ratio are now significant (U = 101.5, P < 0.05; U= 116.5, P < 0.002, N = 12, 11, respectively). Figure 1 shows an inverse relationship be- tween the number of flowers at peak blooming and UNE dry WORK Both sprouting and non- enly distributed along this contia so dat i mean number of flowers at 1987] HERRERA— FLOWER & FRUIT BIOLOGY 75 TABLI Fruit production in open pollinated and bagged flowers of the studied species. Numbers in paren- theses indicate the number of flowers, and N the number of plants. The sample of the only gynodioecious species (Thymus tomentosus) includes three hermaphrodites and four female plants. Species known to be capable of sprouting are marked with an asterisk. Fruit Production (%) Species Open Pollinated Bagged N Armeria velutina 59 (478) 3 (476) 5 Asparagus aphyllus* 24 (804) — 6 Calluna vulgaris* 88 (100) 24 (50) 5 Chamaerops humilis* 4 (4,560) — 7 Cistus libanotis 55 (5,822) 0.2 (538) 4 Cistus salvifolius 50 (3,014) 0 (92) 10 Cytisus grandiflorus 12 (150) 0 (134) 4 Daphne gnidium* 2 (23,749) 2 (2,660) 10 Erica ciliaris* 45 (147) 1 (105) 10 Erica scoparia* 89 (85) 31 (65) 5 Halimium commutatum 92 (100) 0 (100) 5 Halimium halimifolium 41 (70) 1 (1,259) 5 Helianthemum croceum — 0 (171) 3 Helichrysum picardii 68 (1,308) 1 (1,443) 5 Lavandula stoechas 69 (200) 0.4 (1,040) 10 Lonicera periclymenum* 8 (1,537) 4 (211) 4 Myrtus communi. 68 (857) 23 (78) 4 Osyris alba 7 (1,385) — 8 Osyris quadripartita* 5 (2,400) — 5 Phillyrea angustifolia* 14 (1,229) 0 (735) 5 Pistacia lentiscu 16 (1,368) — 5 Rosmarinus Pi daa 31 (256) 10 (797) 5 Rubus le 78 (931) 0 (158) 10 Smilax a 9 (4,053) — 5 ADU genistoides 40 (115) 0.2 (651) 4 Ti us tomentosus 2 (210) 0 (210) 7 Ulex minor 16 (238) 0 (221) 10 Ulex parviflorus 5 (310) 0 (471) 5 peak blooming is not significantly different be- tween them (U — 101, P 0.1); neither is mean flower dry weight significantly different (U = 113.5, P > 0.1, N = 14, 13). Both sprouting and nonsprouting taxa may be many- and small- or few- and big-flowered. DISCUSSION The 30 mediterranean plant species studied represent a relatively small sample ofthe regional flora (more than 2,000 taxa for southern Spain, of which nearly 300 are woody). Furthermore, shrublands have many different and diverse species compositions in southern Spain, depend- ing on elevation, rainfall, edaphic factors, etc., so that the results reported here should be ex- tended only with caution to other scrub com- munities in the region. Nevertheless many ofthe studied taxa are widespread, and the community they form is undoubtedly a clear example of coastal scrub on sandy soils, which is character- istic of other areas in southern Spain. In the studied community a sizeable hetero- geneity in reproductive traits was likely to occur, since 30 plant species were distributed among 17 families. However, certain groups (virtually Cis- did others. Hence a phylogenetic component in the reproductive patterns recognized should not be ruled out, in addition to an ecological com- ponent and to the fact that the plants form a steady, long-lasting community achieving repro- duction year after year. Several aspects in the reproductive patterns are not restricted to our particular community. The relative dominance of taxa in which nectar yield is low or even zero, 76 ANNALS OF THE MISSOURI BOTANICAL GARDEN 100r aa °. Ë ^ s ° ° a = ° t SOF p’ — ° . 3 e u T ô ° A A Eom, ur A, 2 0 1 FRUIT WEIGHT: FLOWER WEIGHT(log) URE 2. Relationship between fruit: flower dry species; black triangles = species in the family Erica- e. for example, is general in the region (J. Herrera, 1985). The consequences of this factor upon the i gena relationships will be dealt with else- ere (J. Herrera, in prep.). Dispersal by inan- imate or vertebrate agents creates a dichotomy that is also found in other mediterranean-type communities (Bullock, 1978, chaparral; C. M. Herrera, 1984b; Jordano, 1984, southern Spain). nities (Reader, 1977; Ruiz-Zapata & Arroy 1978; Bawa, 1979; Bawa & Beach, 1983) un nators are needed for maximum fruit produc- tion. Further investigations will determine the extent to which incompatibility systems occur in shrublands of southern Spain. Sprouting is also a common regenerative strat- egy of woody plants in many and diverse eco- systems (James, 1984). It is particularly impor- tant in regions of a mediterranean-type climate that experience summer droughts and frequent fires. This aspect has received much attention in of a continuum) have been recognized: the “seeder” strategy consists of ob- ligate seed reproduction and incapacity to form new stem sprouts after destruction of aerial parts; the *sprouters" can produce stem or root sprouts that enable repeated shoot production despite frequent damage (Malanson & Westman, 1985; see James, 1984, for an extensive review). It ap- pears that both strategies also are found among sclerophyllous species of the Mediterranean Ba- [Vor. 74 sin. In our community most plants fall clearly into one category or the other, although species in the Ericaceae could be easily included in both: they produce plenty of seeds and are also able to sprout, which was reported some time ago for Calluna vulgaris (Kayll & Gimingham, 1965). It has been hypothesized that pollination-re- production variables may be associated with each of the sprouting or nonsprouting strategies. In seeders, for example, the pollination system must be efficient enough to assure the release of nu- merous seeds (which could open a way to self- compatibility), produce numerous flowers highly attractive to pollinators, and offer a greater re- ward per flower than sprouters (Carpenter & Recher, 1979). In contrast, sprouting taxa are proposed to have a high rate of outcrossing, al- though often with a low production of flowers, which would be relatively low rewarding (Fulton & Carpenter, 1979; Carpenter & Recher, 1979). "ee hypotheses are supported in part by our dat ^ high rate of outcrossing has to be expected in the sprouting group of species, since many dioecious taxa are included in that group. There are no evident signs, however, of a high incidence of self-pollination in the nonsprouting group. In contrast, mean percent fruit production was sig- nificantly higher for nonsprouters than for sprouting species, which the hypothesis outlined above. Low pollination efficiency due to unisexuality can be reasonably proposed to explain low fruit production in the sprouting group, but we wish to put forward a comple- mentary view. Percent of fruit production was found to be inversely related to fruit : flower size ratio (i.e., the increase in mass needed to produce a fruit from a flower). Such a relationship sug- gests the existence of a trade-off between the en- ergy that a plant allocates to an individual fruit or seed and the number it can successfully form. Low fruit production in vertebrate-dispersed, big- seeded species may reflect low pollination suc- cess and/or the impossibility of developing every fruit with pollinated ovules, due to the relatively igh costs involved in the ripening process. It has been demonstrated, for example, that some species experience low fruit production despite good levels of pollination, which is apparently due to intrinsic regulatory mechanisms (Lloyd, 1980; Wyatt, 1981; Casper & Wiens, 1981; Cas- per, 1983; Bookman, 1983; Bawa & Webb, 1984; see Stephenson, 1981, for a review). Flower size and number showed a negative relationship too, 1987] but the predicted tendency for sprouting taxa to appear at one end of this continuum has not been detected by Wells (1969) pointed out that the capacity to sprout vegetatively from underground parts is probably an ancestral trait. James (1984), how- ever, suggested that sprouting cannot always be seen as an ancestral trait in woody dicotyledons. In the present study the capacity to sprout is associated with other traits, such as the produc- tion of fleshy, vertebrate-ingested fruits, heavy seeds, and low fruit production, along with a relatively high incidence of dioecy (species in the genera Asparagus, Chamaerops, Lonicera, Osy- ris, Pistacia, Rhamnus and Smilax, for exam- ple). This group of taxa has subtropical affinities: sclerophyllous species in these genera existed well before the Pleistocene and the evolution of true mediterranean climatic conditions (Raven, 1973; Axelrod, 1975; Pons, 1981). In contrast, species in the genera Armeria, Cistus, Cytisus, Hali- mium, Lavandula, Rosmarinus, or Thymus are nonsprouters; and the high number of taxa en- demic to the Mediterranean Basin makes clear that species in these genera originated much more recently (Quezel, 1978, 1981; Pons, 1981). The capacity to sprout is thus lacking in the typically mediterranean taxa but is jiu in the more mediterranean ones. This supports s (1969) iat pene se is an ancestral trait, a it may indicate that only those "tertiary flora" subtropical sclerophyllous taxa with a capacity to sprout were able to survive the shift to seasonal dryness associated with the mediterranean climate. LITERATURE CITED ALLIER, C. F., F. GONZALEZ-BERNALDEZ & L. RA- MIREZ-DíAz. 1974. Reserva Biológica de Doñ- ana. Ecological Map. Estación Biológica de Don- 1975. Evolution and biogeography ° Madrean- Tethyan sclerophyll vegetation. re Mi issouri Bot. Gard. 284-334. Bawa, K. S. 979. 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JAMES, S. . Lignotubers and burls—their struc- ture, function and ecological significance in med- iterranean ecosystems. Bot. Rev. 50: 225-266. JORDANO, P. 1982. Migrant birds are the main seed 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN Oikos 38: 183-193. 1984. Relaciones entre plantas y aves frugi- voras en el matorral mediterráneo del área de Doñana. Ph. D. Thesis. University of Sevilla, Se- H. GIMINGHAM. 1965. = regeneration of Calluna vulgaris after fire. J. Ecol : 729-734. KEELEY, 77. Seed production, seed popula- tions i in soil and seedling production after fire for ing chaparral shrubs. Ecology 58: 820-829. 1972. Insect yaa of high Arctic . 60: 831 Rd owth and flowering ! he r Veios and Related Shrublands. ER mster OM ipee iid i ` 1983. Comparative phenology of mediterranean-type plant communities. Pp. 300- 317 in F. J. Kruger, D. T. Mitchell & J. U. M. Jarvis (editors), Mediterranean-type Ecosystems. Springer-Verlag, Berlin LLovp, D. G. 1980. Sexual strategies in plants. 1. 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New Phytol. 88: 375-385. REPRODUCTIVE SYSTEMS AND CHROMOSOME RACES OF OXALIS PES-CAPRAE L. AND THEIR BEARING ON THE GENESIS OF A NOXIOUS WEED! ROBERT ORNDUFF? ABSTRACT In its native South Africa, Oxalis pes-caprae is represented by diploid and tetraploid races; the short-styled, d So uth ual proportions in natural populations. The diploid and tetraploid races of the species have a well- developed incompatibility system associated with their floral trimorphism. Outside South Africa, the is there evi idence of any aggressive superiority of the short- styled morph i in the South Africa. W h Afri asa eed dn the diploids and tetraploids within South Africa, is unknow Oxalis pes-caprae L. is a troublesome and widespread agricultural and garden weed, par- ticularly in areas of the world with a mediter- ranean climate such as central Chile, the Medi- terranean basin, parts of Australia, and California (Salter, 1944; Young, 1958; Munz, 1959; Mi- chael, 1964). The species is native to southern Africa, where it is variable (Salter, 1944), and where it is distributed from Namibia (South West Africa) southward to the Cape region and around the Indian Ocean coast at least as far north as the Knysna area, sometimes ranging well inland. There it occurs as a “well-behaved” native of relatively undisturbed sites as well as a weed, and it is particularly common in vineyards and along roadsides. In southern Africa, the species is tristylous (Fig. 1), but throughout most of its exotic range it is represented by a short-styled form, which is pentaploid (25 — 35), and which reproduces asexually via bulbils. In parts of its exotic range, tetraploid and presumably sexual populations also occur, though less commonly than the sterile pentaploid. In view of the importance of this species as a weed, certain features of the reproductive biology es rica, but apparent less successful and chromosome cytology of plants originating in South Africa were studied in the hopes of elu- cidating the events that led to the origin of the aggressive weedy races from the native races. MATERIALS AND METHODS Bulbs collected from natural populations of Oxalis pes-caprae in the Cape Province region in South Africa in 1970 and 1971 were later grown at Berkeley for chromosome studies. One col- lection was provided by Sherwin Carlquist. Chromosome numbers were determined by ex- amining microsporogenesis in flower buds of these cultivated plants preserved in 3: 1 ethanol: acetic acid and stained in acetocarmine. Bulbs collected by Peter Goldblatt from two localities in 1984 provided plants used in an artificial crossing program to determine the presence and and a vineyard at Rustenberg, near Stellenbosch, Cape Province. The two localities are ca. 50 km from each other. Crosses were made in the spring of 1985 and 1986 by transferring pollen from ! Supported in part by grants from the National eir Foundation, the Council for Scientific and Industrial ke Research of South Africa, and the Committee on R Carlquist, T. I. Chuang, Rivka cea = Peter Goldblatt n, and Patricia Watters for assisting this a in various 0. Kim E. Steiner, David E. Sym rch, University = lifornia, Berkeley. I thank Sherwin , R. Harry Koga, Peter W. Mic = se Donald Pfister, ? Department of Botany, unity of California, Berkeley, California 9472 ANN. MIssouRI Bor. GARD. 74: 79-84. 1987. 80 ANNALS OF THE MISSOURI BOTANICAL GARDEN aie 5 mm Mid Long FIGURE 1. ignation of each floral form the text are given to the iw of the figure. dehisced anthers to stigmas under insect-free conditions. Seed set was determined by collect- ing nearly mature capsules and counting the seeds released by them in seed packets. Pollen size for the three morphs was determined by mounting i grains in aniline blue-lactophenol and mea- ring them with an ocular micrometer; pollen stalna bility was determined by counting the number of stained grains in a sample of 100 mounted in this medium RESULTS Chromosome numbers. Seven ofthe nine na- tive populations examined were tetraploid (n = 14); two were diploid (n = 7; Table 1). Previously published reports, summarized in Table 2, in- dicate only tetraploidy and pentaploidy for the species. Weediness. The two diploid populations and four of the tetraploid populations were growing under disturbed conditions such as roadsides, vineyards, or grainfields and were considered weedy; the others occurred in undisturbed con- ed (field data MIURI Udta ditions and are lacking for one tetraploid Bauen Table 1). Morph ratios. Morph ratios have been re- ported earlier (Ornduff, 1974). In two popula- [VoL. 74 long -level mid-level l £ short-level Short Illustrations d gynoecia and androecia of the three floral forms of Oxalis pes-caprae. The des- s given below each illustration. The stigma and anther level designations used in tions, morph ratios of Longs : Mids: Shorts were 1:1: 1, both populations were considered, with some doubt, to be non-weedy. In four popula- tions, morph ratios deviated from equality. In one of these, Longs were deficient; in another, Mids; in third, Shorts; and in the fourth, Longs and Mids were greatly outnumbered by Shorts. Three of these four populations were character- ized as weedy (two for which chromosome counts are available were tetraploid). Pollen size and stainability. Pollen size of diploids and tetraploids is trimorphic (Table 1). Pollen from the long-level anthers is largest, that from mid-level anthers is smaller, and that from short-level anthers is smallest. Pollen from an- thers at equivalent levels in different morphs was generally of di same size duin a population, but there were i pol- len size from equivalent anther sets. There was no correlation between differences in pollen size and in chromosome number of the diploids and tetraploids. Pollen from presumed pentaploids collected in California was extremely variable in size, even from an individual anther. Pollen stainability of diploids and tetraploids was vari- able, but was mostly over 6096. One Short tet- d (7038, Table 1) had pollen with 1296 and 25% stainability from its two sets of anthers. One lege Ave., Berkeley 1987] ORNDUFF—ONXALIS PES-CAPRAE TABLE l. Chromosome numbers, pollen size (um), and pollen stainability in Oxalis pes-caprae. Origin of Pollen ollection (authors Chrome: (size; standard deviation; stainability in percent) or collector's field some Floral Long-level Mid-level Short-level umb Number Morph thers Anthers Anthers Cape Province, South Africa: 7035: Worcester, river- n= 14 Long 41.7; 1.83 (—) 35.8; 2.03 (—) bank, non-weedy Mid 45.7; 2.65 (—) 33.0; 2.05 (—) 7038: Klapmuts, road- n = 14 Short 54.1: 3.94 (12%) 41.2; 3.10 (25%) side, weedy 7041: Stellenbosch, — Short 49.5; 3.20 (66%) 41.8; 1.83 (69%) vineyard, weedy 7096: Mamre Road Sta- n=7 Mid 46.4; 1.95 (99%) 33.7; 1.91 (98%) tion, roadside, weedy 7248: Gouda/Hermon, n] Mid 42.7; 3.94 (—) 31.1; 2.20 (—) roadside, weedy 7292: KL on yman — Long 41.8; 1.83 (8990) 42.7; 4.84 (88%) ry, field, non- Mid 50.1; 2.38 (86%) 38.9; 1.83 (7890) eed Short — 50.4; 4.15 (93%) 41.5; 2.93 (98%) 7301: — n" n = 14 Long 39.0; 1.83 (78%) 30.3; 3.92 (79%) baan, field, non- Mid 46.2; 1.68 (83%) 31.2; 1.54 (81%) weedy Short 48.3; 5.73 (81%) 34.4; 2.03 (77%) 8053: Calvinia, grain n=14 Long 42.6; 2.42 (65%) 36.2; 2.52 (60%) field, weedy id 52.2; 2.67 (64%) 35.0; 1.81 (40%) Short — 47.2: 2.75 (74%) 38.0; 2.55 (73%) 8055: Clanwilliam, — Long 38.2; 2.49 (75%) 32.6; 2.07 (68%) veld, non-weedy Mid 45.1; 2.89 (94%) 32.6; 2.30 (86%) Short 48.3; 2.87 (88%) 38.8; 2.83 (82%) Goldblatt s.n.: Rustenberg, n = 14 Long (90%) —-;— (84%) near Stellenbosch, Mid —;— (6696) —;— (62%) abandoned farmland Short — — (70%) —;— (8196) Goldblatt s.n.: Noordhoek, n = 14 Long —;— (93%) —;— (88%) Cape Peninsula Mid —;— (81%) —;— (79%) Short —;— (88%) -— (7896) Carlquist s.n.: South- n= 14 Long 40.2; 2.46 (80%) 32.0; 1.93 (46%) western Cape Prov- Mid 50.8; 4.84 (92%) 34.5; 2.01 (96%) ince Short 49.7; 4.37 (67%) 41.8; 3.71 (63%) California, U.S.A.: Ornduff s.n.: Univ. — Short —;— (62%) —;— (87%) California campus, — Short —;— (58%) —;— (51%) Berkeley R. Dulberger s.n.: as — Mid =; — (49%) —;— (5896) above R. Dulberger s.n.: Col- — Mid —;— (32%) —;— (29%) set ofanthers ofa few other collections (e.g., 8053 Mid, Carlquist s.n. Long) had pollen with low stainability. The two collections used in the crossing program had pollen stainabilities ex- ceeding 62%. Presumed Short pentaploids col- lected in California had pollen stainabilities ranging from 29% to 87%. However, these pollen grains were variable in size, and the total number per anther was reduced compared with pollen production of diploids and tetraploids as esti- mated from the density of the pollen grains on the prepared slides. 82 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 2. Published chromosome counts of Oxalis pes-caprae. Chromosome Number Locality of Population Examined Reference 2n = 28 Cape Town, South Africa Marks, 1956 India (garden plants) Mathew, 1958 Madeira Borgen, 1974 South Australia Oram in Symon, 1961 Western Australia Michael, 1964 2n = 34 India (Punjab) Bir and Sidhu, 1978, 1979, 1980 Sidhu, 1979 2n = ca. 35 Italy Vignoli, 1935, 1937 2n = 35 Unknown South Australia Western Australia Cape Town, South Africa Franklin in Michael, 1964 Franklin in Michael, 1964 Compatibility relationships. Table 3 presents ent plants from Rustenberg and Noordhoek. Le- gitimate pollinations are those between anthers and stigmas at equivalent levels; all other pol- linations are termed illegitimate (Darwin, 1877). Only 4% of the illegitimately pollinated flowers produced capsules compared with 87% of the legitimately pollinated flowers. The average number of seeds per FEE eine jus legitimate crosses was 15.9; fi t it was 0.2. Self-pollinations dud 0-0.2 ied per pollination. Intermorph, illegitimate polli- nations produced 0—0.5 seeds per pollination. In- termorph, legitimate pollinations produced 13.0- 20.9 seeds per pollination. The two populations used in the crossing program produced similar results for all classes of crosses. Seed production of Shorts following legitimate pollinations was slightly greater than that ofthe other two morphs. oth populations are tetraploid (Table 1). DISCUSSION In its native range in South Africa, Oxalis pes- caprae is represented by indigenous weedy and non-weedy races with a conventional tristylous floral morphology, including pollen-size tri- morphism, and a fully pia trimorphic in- compatibility system. viduals are strongly self-incompatible and oaa cross-polli- nations produce little or no seed. Legitimate pol- linations produce considerable quantities of seed. The pentaploid form has been reported from that country as well (Michael, 1964), but it is unclear how common this race is there. In South Africa, tetraploids are apparently more common than diploids, but weedy populations of both are known. Unequal morph ratios may be more common in weedy South African races than in non-weedy ones and are likely a conse- quence of vegetative propagation that is en- hanced by physical disturbances of the habitat during agricultural or road-building activities. Morph ratios were found in which Longs, Mids, and Shorts, respectively, were deficient in num- bers. Each unequal morph ratio differed from the others, with no morph(s) predominating overall. Thus there is no basis from observation of native races that accounts for the fact that the aggressive morph outside South Africa is short-styled. Over most of its exotic range of distribution, Oxalis pes-caprae appears to be represented by a fairly sterile, pentaploid short-styled morph. As early as 1887, Hildebrand noted the preva- lence of this short-styled form and its lack of seed set. Henslow (1891, 1910) also noted these fea- tures and described in some detail the means of vegetative reproduction later amplified by Galil (1968). Where introduced, the species is distrib- uted by human agents and, in places, by other animals such as the mole-rat (Galil, 1967) or by birds (Young, 1958). Despite the high level of pollen sterility of the short-styled pentaploid, it apparently reproduces occasionally by seed. This may result from self- or geitonogamous pollinations in populations where Shorts alone are represented. Illegitimate pollinations of sexual Shorts carried out in the present study produced small amounts of seed, which offers support for this suggestion. Vignoli (1937), in an embryological study of the species, noted rare sexual reproduction in the pentaploid but concluded also that apogamy may rarely oc- cur in this race. Another line of evidence for 1987] TABLE 3. ORNDUFF=— OXALIS PES-CAPRAE 83 Results of legitimate and illegitimate pollinations in two populations of Oxalis pes-caprae. The first figures summarize results for the Rustenberg population; the second figures summarize results for the Noordhoek populatio Number of Average Style Length (9 parent) x Number of Flowers Number nther Level/Style Length Flowers Producing umber o of See rent)! Pollinated Capsules Seeds Obtained Pollination Self-pollinations (all illegitimate): L x m/L selfed 18; 55 0; 6 0; 34 0; 0.6 L x s/L selfed 20; 12 0; 0 0; 0 0; 0 M x I/M selfed 38; 33 0; 1 0; 5 0; 0.2 M x s/M selfed 23; 35 0; 0 0; 0 0; 0 S x VS selfed 18; 20 1; 0 1; 0 0.1; 0 S x m/S selfed 15; 12 0; 0 0; 0 0; 0 Intermorph, illegitimate pollinations: L x s/M 26; 20 0; 0 0; 0 0; 0 L x m/S 23; 13 1; 2 10; 7 0.4; 0.5 M x s/L 25; 29 0; 2 0; 8 0; 0.3 M x I/S 15; 26 0; 1 0; 1 0; 0 S x l/M 15; 21 0; 3 0; 9 0; 0.4 S x m/L 25; 30 1;4 17; 6 0.7; 0.2 Intermorph, legitimate pollinations: x l/M 18; 48 12; 41 279; 738 15.5; 15.4 L x I/S 27; 34 23; 30 430; 583 15.9; 17.2 M x m/L 33; 37 29; 38 413; 533 12.5; 14.4 M x m/S 54; 42 46; 34 797; 546 14.8; 13.0 S x s/L 13; 22 11; 20 222; 460 17.1; 20.9 S x s/M 11; 29 11; 25 255; 588 23.2; 20.1 Summary of all legitimate, illegitimate pollinations: Legitimate 156; 212 132; 188 2,396; 3,448 15.4; 16.3 Illegitimate 261; 306 3; 19 28; 70 0.1; 0.2 tation is as follows: “L x m/L selfed” means a long- styled flower (L) was self-pollinated with its own s from the mid-level (m) set of anthers. ‘ means a long-styled flower was pollinated with pollen from the short-level stamens of a mid-styled flower. Figure l illustrates the three flower types occasional sexual reproduction of this form comes rwi e southern Italy by Vignoli (1935) and in central California by Dulberger (pers. comm.). It also has been suggested that in Western Australia, where the pentaploid sometimes exists in mixed populations with tetraploids, hybridization be- tween the two may occur (Michael, 1964). The variability of the species there would suggest fre- 1961) and and Madeira (Mathew, 1958; Borgen, 1974), but it is unknown whether these are composed of more than one morph. Although the occurrence of tetraploids and pentaploids in Australia as a consequence of in- dependent introductions is documented (Mi- chael, 1964), it is possible to explain the origin of tetraploids within pentaploid populations by another mechanism. The pentaploids proque some viable pollen (Table i Vignoli, & Sidhu, 1980; Michael, 1964). The illustrations and discussion of microsporogenesis in penta- ploids by Vignoli (1935, 1937) indicate that pol- len grains with n = 14 can be produced by such plants. If megasporogenesis also results in eggs with 71 — 14, fertilization of these eggs by diploid sperm would result in a tetraploid zygote. The place and mode of origin of the weedy 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN pentaploid race are uncertain. Lower (1963) sug- gested that it may have originated outside South Africa. A 5x chromosome count reported by Mi- chael (1964) indicates that this race is present in South Africa, but it is apparently not common (although this remains to be documented). The origin of the weedy pentaploid race can only ten- tatively be reconstructed, but I believe that it likely occurred in South Africa. The simplest ex- planation for the origin of pentaploidy is that it resulted from the union of an unreduced gamete of a tetraploid plant with a haploid gamete from a diploid plant. Since diploids are unknown out- side South Africa, this event must be assumed to have occurred in South Africa. Although dip- loids and tetraploids are not known to be sym- patric in South Africa, they have been collected very near each other, so sympatry may occur. Weediness clearly preceded the occurrence of pentaploidy since sexual diploids and tetraploids are frequently aggressively weedy: they com- monly occur in cultivated fields and along road- sides. Pentaploidy itself has not conferred weedi- ness on the species. Likewise, prodigious means of vegetative reproduction occur in diploids and tetraploids with a fully developed sexual appa- ratus, so that the almost exclusively vegetative propagation characteristic of the pentaploid is a condition that likewise preceded the origin of pentaploidy. Present evidence, although scanty, suggests that the largely sterile pentaploid race is less success- ful as a weed in South Africa than are the sexual diploids and tetraploids. Outside South Africa, however, the pentaploid seems to prevail, pos- sibly as a consequence of its greater competitive success Over sexual races under exotic conditions (as, for example, seems to be the case i in Austra- lia) or of introduction that led to this race being more abundant than its apparent diploid and tetra- ploid precursors. There is a possibility that there are competitive differences among the chromo- somal races of Oxalis pes-caprae, or among its morphs (as suggested ae io ns Tue re- 11 suggests the possibility That: in these, the sexual apparatus may be impaired and that asexual mechanisms are more important in their repro- ductive mode. Both suggestions merit study. Nevertheless, the sequence of events leading to the origin of pentaploidy and the routes of hu- man-aided migration and introduction of this species to other continents will probably never be fully reconstructed. [VoL. 74 LITERATURE CITED Bir, S. S. & M. SipHu. 1978. In IOPB chromosome number reports LX. Taxon 27: 223-231. 1979. Cytological observations in weed flora of orchards of Patiala TA Punjab. Recent Res. Pl. Sci. (New gs li) 7 1-271. jon aul studies on weed flora k cultivable lands of Patiala district, Punjab. J. Palynology 16: 85- BORGEN, L. romosome nu mbers of Maca- ronesian flowering plants II. Norwegian J. Bot. 21: 195-210. DARWIN, C. The Different Forms of Flowers on Plants of the Same Species. John Murray, Lon- on. Gaui, J. 1967. On the dispersal of the bulbs of Oxalis cernua Thunb. by rare (Spalax eh- bii Nehring). J. Ecol. 55: 787-7 968. Miren dispersal in Oxalis cernua. m J. Bot. 55: 68-7 HrENsLOW, G. 1891. On the northern distribution of Oxalis cernua Thunb. Proc. Linnean Soc. Lond. 1890-1891: 31-36. 1910. Remarkable instances of plant disper- 2-361. eh oy. Hort. Soc Mes BRAN Experimente über die ge- aie ake Fortpflanzungsweise der Oxalisar- ten. Bot. Zeit —6; 33-40. in H. F. rt on Oxalis pes-caprae L. 963. n South Africa. Unpublished document supplied by Waite Institute, Glen Osmond, South Austra- lia. Marks, G. E. 1956. Chromosome numbers in the genus Oxalis. New Phytol. 55: 120-129. MATHEW, P. M. 1958. Cytology of Oxalidaceae. Cy- tologia 23: 200- MICHAEL, P. W. 196 e identity and origin of ifornia Press, Berkeley and Los Angeles. ORNDUFF, R. 1974. Heterostyly in South African plants: a conspectus. J. South African Bot. 40: 169- 187. ios J. R. 1973. Soursob (Oxalis pes-caprae L.) n We stern Australia: its life history, distribution, Distributional and Cyt tological Flora of Cultivable Fields of . E. The species of Oxalis established n South Australia. Trans. Royal Soc. S. Aus -71. TN L. 1935. Ricerche preliminari di citologia sull'Oxalis cernua Thunb. Nuovo Giorn. Bot. Ital. 42: 668-669. — P. 1937. Fenomeni reproduttivi di Oxalis cer- nua Thunb. Lavori Inst. Bot. Palermo 8: 5-30. Y AMASHITA, K. 1935. Zytologische Studien an Oxalis I. Jap. J. Genet. 11: 36. Young, D. P. 1958. Oxalis in the British Isles. Wat- sonia 4: 51—69. FLORA OF THE VENEZUELAN GUAYANA — II JULIAN A. STEYERMARK! ABSTRACT A total of 18 species, four subspecies, and one variety are newly described from the Venezuelan a & = x = = = 3 Q = < a ~ o e = = t Q Fu , Kp, w^ Dp lss] Q = = & > Q = = S m = a8 [^] e 3 > r is (Ochn Y UCVIlySidUCav) phologically closely d but —— ape yos subspecies isolated on different sandstone descri table mountains. A key to the subspecies provided, as well as a ie to the BE acted species s of Bonnet BROMELIACEAE Brocchinia oliva-estevae Steyerm. & Lyman B. Smith, sp. nov. TYPE: VENEZUELA. Bolivar: summit of Auyan-tepui, extreme north end above Angel Falls, Dec. 1984, Francisco Oliva Esteva s.n. (holotype, VEN). Figure 1. lanta parva caulescens florifera 3.7 dm alta, caule s, nervis paullo obscuris; scapo folioso, scapi bracteis lanceolato-ligu- latuis 6-8 cm longis, 1.4—1.7 cm latis; VENE ag erecta laxe bipinnatim Mr 2.1 dm longa tenui, arsim brunneo-furfuracea xem. axibus c pedicellis 1— 2: pro ngis minute furfuraceis; petalis albidis lanceo- latis pesce al Med res tuse € acutis haud Baguicutatis 3 m longis, longis 0.5 mm in vario inferiore aban oeda 3 mm longo; ovulis aes e ia appendicibus cau- datis Caulescent, small, herbaceous plant, flowering 3.7 dm tall; stem erect, 13 cm tall. Leaves as- cending, pale dull green both sides with about 10 parallel, slightly darker longitudinal lines be- coming bronzy where entering sheath, submem- branous, flexible, concave above, convex below, ibed Sauvagesia marahuacensis is tia. not contracted at base, the larger ones ligulate- lanceolate, acute, mucronate, 10 cm long, 1.7- 1.8 cm wide, lower leaves ovate, acuminate, 1.5— 2.8 cm long, 1-1.4 cm wide, the upper ones lon- ger than the internodes, minutely pale lepidote. Scape foliose, scape bracts lanceolate-ligulate, 6— 8 cm long, 1.4-1.7 cm wide. Inflorescence erect, laxly bipinnately paniculate, 2.1 dm long; rachis slender, covered with a scattered brown furfura- ceous indument, with 5 short, racemose, lateral axes 3.5-6 cm long, each axis 15-17-flowered, the uppermost part ofthe inflorescence elongated to 10 cm; primary bracts subtending the lower three axes ascending, lanceolate, acute, 3 cm long, 0.8 cm wide, entire; floral bracts oe ah acute to acuminate, 3—3.5 mm long, 0.7 wide at the base. Flowers shortly pedicellate, E l- 2 mm long, Medie Eco = pale een, lanceolate, acute, 3—4. ong, | mm wide, dorsally die -DTE in lower half. nguiculate, Anthers suborbicular-oblong, 0.5 mm long, ba- sally bilobed; filaments white, 1.2 mm long. vary inferior, pale green, subclavate-cylindric, 3 mm long, 1 mm wide at summit, 0.7 mm wide at base, pale brown furfuraceous; ovules caudate- appendaged at both ends. This species of Brocchinia is characterized by its bipinnate inflorescence with lepidote, simple, ' Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, ANN. MissouRi Bor. GARD. 74: 85-116. 1987. FIGURE |. ANNALS OF THE MISSOURI BOTANICAL GARDEN Brocchinia oliva-estevae.— A. Habit.—B. Flower with bract. [VoL. 74 1987] racemose axes, primary bracts extending from !⁄4 to 4 the length of the rachis, shortly pedicellate flowers, non-unguiculate petals, and narrow sub- membranous, slightly nerved, short leaves only 10 cm long by 1.8 cm wide, which are not con- tracted toward the base. It is most closely related to B. cowanii L. B. Smith of Cerro Moriche, Terr. Fed. Amazonas, Venezuela, in having a bipin- nate inflorescence with non-unguiculate petals, but differs from that taxon in the scape bracts shorter than the internodes and in the shorter sepals and petals. It is a pleasure to name this interesting species for Mr. Francisco Oliva Esteva, Venezuelan landscape architect, an avid student of Brome- liaceae, and author of several books on orna- mental plants of Venezuela. PIPERACEAE Peperomia marahuacensis Steyerm., sp. nov. TYPE: VENEZUELA. Territorio Federal Ama- 65°23'W, 2,520-2,620 m, 26-27 Feb. 1985, Julian A. Steyermark & Bruce Holst 130742-A (holotype, MO; isotype, VEN). Figure 2C Herba effusa; caulibus elongatis 8 dm longis, 2.5-3 scentes ramorum spicuis nervis duobus lateralibus inconspicuis, supra nervis principalibus pilosulis, demum glabris, subtus nervis sparsim pilosulis pilis patentibus 0.2-0.5 mm longis ceterum glabris vel glabratis, marginibus parte suprema '4-!6 ciliolatis ceterum glabris; petiolis 5-20 mm longis, laminis 2—3-plo brevioribus pilosis. Sprawling herbaceous plant with stems elon- gated to 8 dm, 2-3 mm diam., sparsely puber- ulous except at the densely pubescent younger tips. Leaves alternate, orbicular- to triangular- ovate, acuminate at apex, truncate or broadly rounded at base, larger blades 3-4.5 cm them conspicuous and 2 lateat nerves Tant, Lal brouson theu the main nerves, de surface glabrous or da brate, but the nerves sparsely pilosulous with spreading hairs 0.2-0.5 mm long; leaf margins ciliolate in the uppermost '4-!5, elsewhere gla- brous; petioles 5-20 mm long, '4—'2 length of leaf blade, pilose. STEYERMARK-— VENEZUELAN GUAYANA 87 From the related P. foveolata Steyerm. of Ce- cent along the nerves of the upper and lower surfaces, and in the shorter, pubescent petioles, which are proportionately shorter in relation to the length of the leaf blade. Compared with P. peltoidea Kunth, it differs in the non-peltate, non- ace, but mainly glabrous below on the leaf egi itself, with leaf margins ciliolate only in the uppermost !4—!5 , and in the petioles short- er than the leaf blades. Peperomia gentryi Steyerm., sp. nov. TYPE: VENEZUELA. Territorio Federal Amazonas: Cerro de la Neblina, Camp V, valley north base of Pico pus pee 0?49'N, 66°0’W, 1,250 m, 21-24 M 1984, Ronald Liesner & Brian opie: 16901 (holotype, MO; iso- type, VEN). Herba repens; foliis alternis late ovatis vel subrhom- infimis 5-7-plinerviis praeter margines superiores cili- atos glabris; spicis geminis 2-3.5 cm longis; drupis el- lipsoideo-ovoideis rostratis. Repent herb with elongated glabrous stems 2 mm diam.; internodes 4—9 cm long. Leaves al- ternate, broadly ovate to subrhombic-ovate, acute to subacute at apex, truncate to broadly obtuse at base, 5-7-plinerved, the innermost lateral nerves forking off the midrib within the lower- most 5-11 mm, 4.5-9 x 3-5.5 cm, moderately pilose-ciliate on the upper margins; petiole 3.5— 7 cm long. Inflorescence with paired spikes; pri- mary peduncle 2.8-5 cm long, minutely puber- ulent; secondary peduncle subtending the spike .5-2 cm long, sparsely puberulent 2 or glabrous; spikes 2-3.5 cm long, 1.5-2 m iam. Fruit basally attached, p spin ovoid, rostrate, the body 0.5 x 0.3 mm, the beak 0.2-0.3 mm long; stigma anterior near base of beak. — aratype. Same locality as type, 12 Apr. 1984, Gen- try & Stein 46542 (MO, VEN). Figure 2A, B This species is related to both P. distachya and P. schwackei. From P. distachya (L.) A. Dietr. this species differs in the 5—7-plinerved broadly ovate to subrhombic-ovate leaves, which are acute to subacute instead of acuminate at the apex and truncate to broadly obtuse at the base. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 13mm i > R £ E A ant i | "d 5 w S E ; X š z NÁ ` € | k | \= N / < =—— A S | M FIGURE 2. Peperomia. A, B. P. gentryi. — A. Habit. — B. Beaked fruit. —C. P. marahuacensis. — Habit STEYERMARK 1987] — VENEZUELAN GUAYANA 89 From P. schwackei C. DC. of southern Brazil, it differs in the 5-7-plinerved, acute to subacute leaves and longer secondary peduncle, which supports the spike. Piper gentryi Steyerm., sp. nov. TYPE: VENEZUELA. erritorio Federal Amazonas: Cerro Ne- blina, trail due south from Base Camp, stunted swampy forest, alt. 140 m, 23 Apr. 1984, Alwyn Gentry & Bruce Stein 46887 (holotype, MO). Frutex 1.5-metralis, internodiis 2-3.5 mm diam. gla- bris; foliis grosse rugosis lanceolato-ellipticis vel ob- longo-ellipticis apice acutis vel acuminatis basi subae- quilateraliter acutis vel subacutis 1 7-22 cm longis, 5.5— 7.5 cm latis supra glabris subtus ad nervos hirsutulis ceterum glabris, nervis lateralibus utroque latere 11- 15 totis pinnatim venosis; petiolis 3 12 mm longis breviter hirsutulis; spicis 2 cm longis, 6-7 mm latis; pedunculis 3 mm longis glabris; E floriferis fim- briatis. Shrub 1.5 m tall, internodes 2-3.5 mm diam., glabrous. Leaves lance-elliptic or oblong-elliptic, acute to acuminate at apex, subequally acute or subacute at base, 17-22 cm long, 5.5-7.5 cm wide, broadest at the middle, pinnately nerved throughout to apex, glabrous above, punctate, glabrous below on leaf surface but hirsutulous with short, spreading hairs on the midrib and lateral nerves, grossly rugose both sides, venation sulcate above, elevated below, the areoles formed by the tertiary veins averaging 5-7 mm diam .; lateral nerves 11—15 each side, spreading at an angle of 15—20°, sulcate above, subelevated be- low, anastomosing at 5-10 mm from the margin; petioles 5-12 mm long, shortly hirsutulous, vagi- nate to the base of the blade. Spikes erect-as- cending, 2 cm long, 6-7 mm wide; peduncle 3 above, depressed at summit, 2 mm diam., gla- species is related to P. holtii Trel. & daa from it differs in the glabrous stems and nae more numerous lateral nerves, and floral bracts conspicuously fimbriate. V OCHYSIACEAE A REEVALUATION OF THE GENUS EUPHRONIA (VOCHYSIACEAE) In his publication on a revision and taxonomic position of the genus Euphronia, Lleras (1976) showed that the generic name Lightia must be replaced by the earlier Euphronia. Furthermore, he transferred the genus from the Trigoniaceae, where it had been placed previously, to the chysiaceae, the family he considered as showing the closest affinity. Although two species had previously been rec- ognized under Lightia in Warming's treatment of the genus (1875), Lleras considered these taxa as constituting one variable taxon, which he des- ignated E. hirtelloides Martius ex Martius & Zucc. His decision in considering this a polymorphic species was based on what he judged as extreme variability in leaf morphology and lack of geo- graphical or other *'consistency." While preparing a taxonomic treatment for the flora of the Venezuelan Guayana, I have con- cluded that three distinct taxa are involved wor- thy of specific recognition. Consistent differences have been found in characters such as type of inflorescence, length of pedicels, apex of calyx lobes, and quality of pubescence present on the exterior of the calyx lobes and pedicels. More- over, the three taxa demonstrate distinct geo- graphical areas to which they are limited. KEY TO THE SPECIES OF EUPHRONIA la. Calyx lobes caudate-acuminate, the exterior portion pes by a dense, uniform, ap- ed, lanuginose tomentum; inflorescence x, covered with an appressed lanuginose tomentum .......... ee Ib. Calyx lobes pee 1 pe tuse, or subro date at apex, the rior portion with he se, ascending to spreading hairs throughout or at least on midrib; flowers 3—23, racemose, the pe e arising y Ped an oard rachis; pedicels 1.5-5 mm long, shorter than the ca- lyx, rr loosely ascending to spreading pu- bescence 2a. FI am rimus paie ml the. epre branch, mainly (7-)15-23-flowered, 4- mi — with shortly appressed tomen- tu E. hirtelloides š Florife erous portion of rachis si shortly N [^n (-4.8) cm long, mainly ovate to ovate-elliptic, gen- FiGURE3. Euphronia.—A. E. hirtelloides. — A. Habit of edu branch. — AA. Calyx. — B. E. guianensis. — B. Habit of flowering branch. — BB. Calyx.— C. E. acuminatissima. —C. Habit of Busen: branch. — CC. Calyx 1987] < ° 8 ~~ STEYERMARK— VENEZUELAN GUAYANA zuphronia guianensis o suphronia hirtelloides A suphronia acuminatissima W l n? AV nz FIGURE 4. Map showing distribution of Euphronia. KC | es? erally rounded to obtuse at apex, 1.75- 2 times longer than broad; inner calyx white borders; exterior of calyx lobes with dense, loosely ascending to spreading hairs over the entire surface .... E. guianensis Euphronia acuminatissima Steyerm., sp. nov. TYPE: VENEZUELA. Territorio Federal Ama- zonas: Rio Orinoco, Sabana Cumare on right bank of Caño Cumare, Río Atabapo, 20 km g.l pie 43762 (holotype, MO; NY). Figures 3C, CC, 4 Frutex vel arbuscula 0.2—5-metralis; foliis oblongo- ellipticis vel lanceolato-ellipticis apice rotundatis vel obtusis Ap cuspidatis basi acutis vel subacutis 2- 4.5 cm longis, 1-2 cm latis; inflorescentia subfascicu- lata plerumque (2-)3-6-flora pedunculata, pedunculo mm longo; pedicellis 7- 8r mm longis, indumento ena lanceolatis apice caudato- acuminati S | 7-9 mm longis, indumento lanuginoso pilis adpressis munitis. Subshrub to small tree 0.2-5 m tall; stems gray-white lanuginose, becoming glabrescent or glabrous on older or lower portions. Leaves co- riaceous, dark green above, white below, oblong- elliptic to lance-elliptic, rounded to obtuse at the abruptly cuspidate apex, acute to subacute at base, 2-4.5 cm long, 1—2 cm wide, 2.6-3 times longer than broad, gray-floccose above on young leaves, becoming glabrous on older leaves, white-pan- nose below; petioles 2-6 mm long, gray-white lanulose. Inflorescence terminal or axillary, sub- fasciculate, sessile or shortly pedunculate, (2-) 3—6-flowered, the flowers crowded at or near the summit of the abbreviated rachis; peduncle 0.5— 1 cm long, together with the rachis gray-white lanuginose. Bracts subtending pedicels subulate, caudate-acuminate, 3.5-4 mm long, 0.4-1 mm wide, lanuginose, more persistent than in other species. Pedicels 7-8 mm long, nearly equaling the calyx, densely gray-white lanuginose with ap- pressed, more or less uniform indument. Calyx 10-12 mm long, densely gray-white lanuginose without, the hairs E appressed; n cam- panulate-turbinate, 2.5—4 mm long, 3-5 mm wide at summit; lobes ' rowls lanceolate, maine to a long caudate acuminate apex, 7-9 mm long, 92 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1.5-2.5 mm wide, gray-white lanuginose with- out, minutely sericeous within. Petals lavender or lilac, spatulate, rounded at 2 d (15-2)18-23 mm long, (4-)6—9 wide near summit, 1.5 mm wide at or aa, middle, loose- ly pilose within along middle, glabrous else- where, sericeous dis bearing shorter hairs upper port glabrous elsewhere. Anthers oblong, a chavs bilobed basally, 2.5 mm long; filaments 9-11 mm long, the sterile one retrorsely pilose, the others glabrous. Style 10 mm long, antrorsely pilosulous. Fruit cylindric, 13-14 mm long, gray-white lanuginose. Distribution. Mainly on white sand savannas of the Territorio Federal Amazonas, Venezuela, at altitudes of 100-150 meters. Paratypes. VENEZUELA. TERRITORIO FED AMAZONAS: Rio Guainia, Sabana El Venado, left bank l m abo NY); same locality, Maguire & Wurdack ches (NY): base of Cerro Yapacana, 125-130 m, Maguire, Cowan & Wurdack 30479-A, 30509 (NY); Bajo Rio Cha alrededores de Canaripó, a unos 20 km al E de la con- fluencia con el Río Orinoco, 4?03'N, 66?49'W, 98 m, Huber 1070, 1877 (NY, VEN); Rio Ventuari, frente al caserío de Canaripó, 4?09'N, 66°50’W, 100 m, Huber 2441 (NY, VEN); Río Orinoco, poco más río abajo de Santa Barbara, 4°02'N, 67°15'W, 100 m, Huber 2471 (NY, VEN); E del Cafio Perro de Agua, a unos 30 km al SE de la confluencia Orinoco-Ventuari, 3?47'N, 67*00'W, 100 m, Huber & Tillett 2824 (NY, VEN); 10 km al S del Río Autana, 15 km al SW del Cerro ie 4*44'N, 67?33'W, 100 m, Huber 4063 (NY, VEN); Ca San Miguel between Limoncito and Caño Ikebeme (about 70 km from river mouth), 100-140 m, Wurdack & Adderley 43249 (NY); middle Caño Yagua, NE and base of Cerro Cucurito, 3°36'N, 66°34'W, Huber & Tillett 2925 (VEN); 20 km NW of Yavita, headwaters of Caño Pimichín, 3?1'N, 67°33’W, 120 m, Huber & Medina 5947 (VEN); 4 km west of Serranía He Cuao, 4°59" N, 67°32'W, Huber & Tillett 5293 (VEN); Rio 2021 (MO, TFA V). BOLIVAR: T a 133282 This species differs markedly from both E. hir- telloides and E. guianensisin the longer, caudate- acuminate calyx lobes and fewer-flowered, subfasciculate inflorescence with longer pedicels. The pubescence, moreover, is quite different from either of the other species, with both pedicels and the exterior of the calyx lobes covered by a short, appressed, finely lanuginose indument, the other two taxa having a pubescence of longer, loose, spreading to ascending hairs on the ped- [Vor. 74 icels and on a part or whole of the outer surface of the calyx lobes. Euphronia guianensis (R. H. Schomb.) H. Hal- ier, Meded. Herb. Leid. 35: 13. 1918, in obs.: Lightia guianensis. Lightia guianensis Schomb. in Linnaea 20: 757. 1847; Warm in Martius’s Flora Brasiliensis 13(2): 122. 1875. Figures 3B, BB, 4 Shrub or tree (0.2—)2—10 meters tall, branches gray-lanuginose, becoming glabrous below or in age. Leaves coriaceous, gray to silvery white be- low, shortly petiolate to subsessile, mainly ovate to subovate, generally rounded or obtuse at a minutely mucronate apex, less frequently sub- acute, chiefly rounded to obtuse at base, (0.6-) 1.5-3 (4.8) cm long, (0.5—)1—2(-2.5) cm wide (rarely sterile shoots of juvenile branches elliptic- oblong, 7 x 3-3.5 cm), revolute, densely white tomentose below with elevated tomentose mid- rib, young leaves gray-white floccose above, old- er ones mainly glabrous above with narrowly sulcate midrib, sometimes white-tomentose along upper midrib; petioles 1-4 mm long, densely gray tomentose. Inflorescence terminal or axillary, mainly simple and short racemose or occasion- ally the axis with a branch at base, 2-6 cm long including the peduncle, the flowering portion 1- 4 cm long, 1-2 cm wide, 3-8-flowered; peduncle 0.6—2.5(-4) cm long, together with the rachis densely tomentose. Bracts subulate, 4-5 mm long, tomentose, caducous. Pedicels 1-5 mm long, shorter than the calyx, densely tomentose with subspreading to loosely ascending hairs. Calyx 6—9 cm long, tube shallowly campanulate, 1.5— 3 mm long, 3-4 mm wide at summit, densely gray tomentose with short spreading-ascending hairs; calyx lobes slightly unequal, broadly lan- ceolate to oblong-lanceolate, acute, the outer lobes dull gray nearly throughout with narrow paler margins, the inner lobes with conspicuous broad white margins and a narrow gray-green median zone, the longer inner lobes 6.5-7 mm long, 3 mm wide, the shorter outer lobes 4.5-5.5 mm long, 2.5-3 mm wide. Petals bluish, purplish, or rose (varying according to different collectors), spatulate, rounded at apex, unguiculate, 10-15 mm long, m wide, shortly and inconspic- uously appressed-pubescent without to glabres- cent, conspicuously long pilose within except near apex. Anthers narrowly oblong, 2-2.5 mm long; filaments 10-11 mm long, the sterile one re- trorsely pilose, the others glabrous. Style 8-10 1987] mm long, antrorsely pilose. Fruit oblong-cylin- dric, 9-20 mm long, gray short-tomentose. me. “‘Curataquilla’’; *'sacarai- y savannas and open sandstone exposures of the eastern portion of Bolivar in the Venezuelan Guayana from the re- gion of the Gran Sabana at Cerro Roraima west- ward to the summit of Cerro Guaiquinima, and in the adjacent Pakaraima Mountains of Guy- ana, at altitudes of 300—-1,200 m, ascending to 1,600 m on the slopes of Ptari-tepui. [Cardona 2726 from the summit of Auyan-tepui at an al- titude of 2,500 meters was cited as Euphronia hirtelloides by Lleras (1976). It is actually Myrtus alternifolia Gleason, as cited in Flora of Auyan- tepui (Steyermark, 1967).] Specimens examined. GUYANA. Mt. Ayanganna, Pakaraima Mountains, between Chinowieng and Chi- Chi Landing, 1,000 m, Maguire, Bagshaw & C. K Maguire 40647 (NY); Kamarang Station, Pakaraima Mountains, 500 m, Maguire & Fanshawe 32614 (NY). VENEZUELA. BOLIVAR: Gran Sabana, El Dorado-Sta. Elena road, 2.5 km before turnoff at San Rafael, 1,030 m, Luteyn, Lebrón-Luteyn & Steyermark 6294 (MO, NY); km 146 along El Dorado-Sta. Elena road, 1,280 m, Luteyn, Lebrón-Luteyn & Steyermark 6291 (MO, NY); between Santa Teresita de Kavanayen and base of Ptari-tepui, 1,220 m, Steyermark 60307 (F, NY); between Ptari-tepui and Sororopán-tepui, 1,615 m, Steyermark. 60274 (F, MO); Cerro Manacauaray, head- 1,100 m, Cardona 2609 (NY); alrededores de Sta. Elena de Uairén, Lasser 1273 (NY, VEN); Gran Sabana, 27 km N of Kama-Merü, carretera El Dorado-Sta. Elena road, 5?30'N, 61?20'W, 1,300 m, Holst, Steyermark & Manara 2222 (MO); 5 km east of Kavanayen, 1,200 m Maguire 33717 (NY); Uarupata, Maguire 33283 NY); Kamarang head, Gran Sabana, 800—950 m, Ma- guire 33293 (NY); región de los rios Icabaru, Hacha y qe sin nombre, 45 NY ue re del E an 3°41" g"W. "Sieve E. Dunsterville 113455, 113119-A (NY : : 175 south of El Dorado, 1,200 m, Steyermark 111 296 (NY, VEN); Uriman, 300 m, Steyermark 75330 (F, MO, NY); Río Caroní below Urimán, 393 m, Stey- ermark & Wurdack 3 (F, MO, NY); región of Canaima, 6°15'N, 62°47'W, 200—500 m, Agostini 258 (NY, VEN); Hacha ponas Canaima, Prance 16550 (NY, US): be- tween Luepa and Kavanayén, 1,317-1,375 m, Badillo & Forsa 6255 (MY); km 135-1 37 south of El Do- rado, Badillo & Holmquist 6196 (MY); 148 km south of El Dorado, 1,350-1,400 m, Steyermark & Dunster- ville 104162 (MY, VEN); Cerro Akurimá, Sta. Elena, STEYERMARK —VENEZUELAN GUAYANA 93 Tamayo 2699 (US); between Parupa and Kavanayén, eek 792 (VEN); 7.5 km NE of Santa Elena, 4?40'N, 61?4'W, 880 m, Steyermark & Liesner 127592 (MO, UB This species is characteristic ofthe eastern por- tion of the Venezuelan Guayana in the state of Bolívar, where it is a common ied on rocky open sandstone outcrops and savanna Warming (1875) gave only a brief jm of Lightia guianensis, mentioning a few salient characters such as “foliis minoribus, ovatis v. obovatis, basi rotundato-cuneatis; racemis bre- vivissimis 4-6-floris" in differentiating this species onia hirtel- e glabris" has been found to be incorrect. Careful examination of specimens from eastern Vene- zuela shows the filaments of the fifth sterile sta- men to be retrorsely pilose and the four fertile stamens as glabrous, characteristic of the genus. Although united by Lleras with E. hirtel/loides (loc. cit.), E. guianensis is amply distinct not only in its shorter, fewer-flowered racemes and small- er, differently shaped leaves, as mentioned by Warming, but also in the hairs of the outer part of the calyx lobes and pedicels uniformly loosely ascending to spreading. Moreover, dried speci- mens manifest broad white marginal zones on the inner calyx lobes with only a relatively nar- rower gray-green central midrib. This latter char- acter is in contrast to the more uniformly gray- green calyx lobes of E. hirtelloides with only the central median portion with a loose, dense, as- cending sericeous pubescence. e specimens of Prance 16550 and Agostini 258 from the Canaima region need some com- ment. The material of Prance is sterile with the leaves larger than usual, and that of Agostini has inflorescences (up to 7 cm long) and peduncles (3.5—4 cm long) longer than usual, thus attaining dimensions similar to those of E. hirtelloides. However, the short flowering portion (2-3 cm) is like that of E. guianensis. Euphronia hirtelloides Martius, Nov. Gen. et Sp. : 121,4, 73. 1825. Lightia licanioides Spruce Warm. in Mart rasiliensis Lightia licanioides. Figures 3A, AA, Shrub or tree mainly 2-10 m tall, branches gray lanuginose, becoming glabrous below. Leaves coriaceous, gray-green above, white be- 94 ANNALS OF THE MISSOURI BOTANICAL GARDEN low, shortly petiolate, lance- to oblong-elliptic or ovate, shortly acute at apex, acute to obtuse at base, 4.5-7.5 cm long, 1.5—4 cm wide, densely and closely white tomentose below with elon- gated midrib, g i tomentose in the sulcate groove; petioles 5-7 mm long. Inflorescence terminal or axillary, simply racemose, or with 1 or 2 lateral axes branching from the base, 5-11.5 cm long including the pe- duncle, flowering portion 4-10 cm long, 2-2.5 cm wide, (7—)15—23-flowered; peduncle 1-4 cm long, together with the rachis gray-lanulose. Bracts lanceolate, acute, 3.5—4 mm long, 0.8-1 mm wide, sericeous without, caducous. Pedicels 1.5-5 mm long, mainly shorter than the calyx, with dense, loosely ascending to spreading hairs. Calyx 6-9 mm long, tube shallowly campanulate, 2 mm long, 2-4 mm wide at summit, gray-tomentel- lose, lobes unequal, inner ones oblanceolate-ob- ovate, shortly acute, broader than the outer and broadened above the middle, outer ones broadly iid more narrowed above the middle, 2.5-)5-8 mm long, 2-3 mm broad, sericeous both sides, more abundantly long-sericeous with longer, looser hairs on outer midrib, rather uni- formly gray-sericeous elsewhere, externally with shorter appressed hairs. Petals rose, purplish, or lavender, the limb spotted with violet (fide Clark and Maquirino), spatulate, rounded at base, un- guiculate, short-sericeous or sometimes glabres- cent, long pilose within at base and median por- V11IIVillllwo YY hite ile one retrorsely pilose, the others glabrous. Style 11-12 mm long, antrorsely pilose. Fruit cylin- dric, 15-23 mm long, 5-6 mm wide, subobtusely trigonous, gray lanate. Distribution. Amazonian Brazil in Estado Amazonas and Territorio do Roraima, Ama- zonian Colombia in Vaupés and Caquetá, and Territorio Federal Amazonas of southern Ven- ezuela, at altitudes of 100—150 meters. Specimens examined. | VENEZUELA. TERRITORIO FE- DERAL AMAZONAS: ad flumina Casiquiari, Vasiva et Pa- cimoni, 25-26 Feb. 1854, Spruce 3413-x (type of Ligh- tia licanioides; isotype, NY; photo ; Carlos de Rio Negro, 1°56’N, 67°03'W, 119 m, Clark & koa Oe 7338, 7363 (NY); NW base of rro Yapacana, epee a 34525 (MO, NY), 34546 (NY); Sa avann rro Yapacana, m p & Wurdack 305 94 EN Coro . AMAZONAS: Río Caquetá, Araracuara sa- vannas, rpe C. K. Maguire & Permie 44153 ; Vaupés: lower Río Parana-pichuna, at Mitú, x 9 [Vor. 74 RAZIL. AMAZONAS: Rio Curicuriary, affluent Rio Negro, Ducke 337 (GH, MO, NY, US); same locality, Ducke 159-A, 23869 (NY, US); Manaus-Caracarai Road (BR-174), km 115, campina adjacent to Igarapé Lajes, Zarucchi, Almeida & rae eee 2544 (NY); Rio Negro, Preto, Fróes 22753 (M i iuxi 200 km above mouth, 2 of Rio Negro, Prance et al. 15502 (NY, US); Rio Negro to 608 (NY). Territorio Federal Rora nco, Sáo José de 51 (MO, NY, US); Caracarahy, Rio Branco, Ducke ] 407 (GH, NY, US); Rio Negro, Rio Tea 40 km above mouth, village Bacuri, Kubitzki et al. 79. 240 (US). In his description of Lightia licanioides (=Eu- phronia hirtelloides), Warming (1875) incorrect- ly described all five filaments of the stamens as retrorsely pilose, and plate 22 (1875) depicts three of the stamens with retrorse pubescence. How- ever, dissection of specimens pertaining to this taxon reveals the retrorse pilosity present only on the fifth sterile filament, while the other four fertile ones are glabrous. As indicated in the pres- ent key to the species and in comments under E. guianensis, E. hirtelloides is distinct in having a more elongated, many-flowered inflorescence; larger, more acutely tipped leaves; and the pu- bescence of pedicel and outer surface of calyx lobes of a different type. It should be noted here that the calyx of Euphronia is gamophyllous [Lleras (1976) describes the calyx as consisting of five sepals], consisting of a calyx tube and five lobes, as originally defined by Martius and Zuc- carini (1826), Robert Schomburgk (1847, as Lightia), and Warming (1875, as Lightia). AQUIFOLIACEAE Ilex liesneri Steyerm., sp. nov. TYPE: VENEZUELA. Territorio Federal Amazonas: Depto. Ata- erro Marahuaca, 1-2 km N of 3?43'N, 65°31'W, 1,100 m, 8-9 Mar. 1985, Ronald Liesner 18469 (ho- lotype, MO; isotype, VE rutex 1-2 m altus glaber; foliis anguste elliptico- lanceolatis ad apicem obtuse attenuate eque angustatis 4.5-9 cm lon- =f 0.5-1.5 cm latis subtus sparsim punctulatis, ad margines crenato-serrulatis, utroque latere 5-13-cre nulato; inflorescentiis axillaribus et lateralibus solitae inflorescentiae pedicellis filiformibus sub anthesi 4— mm n longis sub fructu 4-8 mm longis; floribus 4- meris; f. calucic Zarucchi 1986 (GH). mm longis, 1.5 mm latis; floribus 9: calycis lobis 0.5 1987] STEYERMARK — VENEZUELAN GUAYANA 95 FIGURE 5. Ilex liesneri. — A. Habit of fl ill fl —B. Habit of flowering [o wasa showing terminal inflorescence. — C. Habit of deos branch. —D. Detail of portion of ins leaf "Pies showing punctation, revolute margin, and crenulations. — E. Pyrenes, lateral view, left; dorsal view, right. — Pistillate flower. — G. Staminate flower mm longis, 1.2 mm latis; petalis suborbicularibus ro- Shrub 1-2 m tall with slender branches. Leaves tundatis 1.5 mm longis, 2 mm latis; fructu subgloboso — narrowly elliptic-lanceolate, narrowed above to vel ovoideo-subgloboso 7 mm longo, 7 mm lato; pyre- nis 4-5 trigonis 5 mm longis, 3.5-5 mm latis, dorso à long obtuse apex, narrowed below to a long 3—5-costatis. acute base, 4.5-9 cm long, 0.5-1.5 cm wide, 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN sparsely punctate below, midrib narrow, sulcate above, slightly elevated below, lower surface enervate, upper surface 7-8-nerved on each side of midrib, subhorizontally spreading, anasto- mosing 1-2 mm from margin, remotely crenu- late-serrulate with 5-13 depressed crenulations on each margin. Petioles 5-15 mm long. Inflo- rescence axillary and lateral, solitary, trichoto- mously cymose, pedunculate; cymes simple or compound, few-flowered (usually 7) when simple with one flower on the central axis and three each on the two lateral axes, when compound each of the axes 2-3-flowered, the axes divaricately spreading, 1-3 mm long; pedicels 4-8 mm lon Bracts subtending pedicels spreading, 0.1 mm long. Peduncle slender, 0.8-1.5 mm long. Flow- ers 4-merous. St te flowers: calyx lobes sub- orbicular, rounded, 1.8 mm long, 1.5 mm wide; anthers suborbicular, 0.8 x 0.8 mm; filaments 0.8 mm long; pistil rudiment ovoid, 1 mm long. Pistillate flowers: calyx lobes 0.5 mm long, 1.2 mm wide; petals suborbicular, rounded, 1.5 mm long, 2 mm wide; ovary ovoid or subglobose, 1— 1.5 mm long, constricted slightly at summit into a short style 0.2-0.3 mm long; stigma prominent, ovoid-capitate. Fruit subglobose or ovoid-sub- globose, 7 x 7 mm; pyrenes 4-5, trigonous, 5 mm long, 3.5-5 mm wide, dorsally 3—5-costate. Paratypes. VENEZUELA. TERRITORIO FEDERAL AMAZONAS: Depto. Atabapo, Cerro Marahuaca, Ri Yameduaka arriba, 3°38'N, 65?28'W, 1,225 m, Liesner 17604 (MO, VEN); “Sima” Camp, wiegen pors tion along eastern branch of Cerro Negro, ahuaca, 3°43'N, 65°31'W, 1,140 m, Stevermark & Holst 130575, 130422, 130424 (MO, VEN). Figure 5 This species is distinguished by the narrowly lance-elliptic, crenulate, punctate leaves, which are long attenuate at each end, the 3—5-dorsally ribbed pyrenes, the slender, solitary peduncles from both axillary and lateral buds, and the fili- form pedicels on trichotomous cymose inflores- cences. OCHNACEAE Tyleria apiculata Sastre (Fig. 6A-G) This species was recently described by Sastre (Phytologia 59: 313-314. 1986) with only a brief description based upon a single specimen, which I collected. A later expedition in 1985 to Cerro Marahuaca by Bruce Holst, Ronald Liesner, and [VoL. 74 me resulted in the collection of more ample ma- terial. In view of these additional collections, the following more detailed description can be fur- nished. Shrub or small tree 2-3 m tall. Leaves erect, crowded, those on sterile branches clustered at the apex, scattered below the inflorescence on fertile branches, subobtuse at apex, sometimes with a minute mucro 2 mm long, the apically clustered leaves on sterile branches 7-8 cm long, 5-10 mm wide, those on fertile branches often shorter and 6-13 mm wide, gradually narrowed to the sessile base, glabrous on both surfaces, margins finely serrulate-ciliate with closely as- cending purplish hairs 0.5-0.7 mm long, lateral nerves finely parallel from base to apex, strongly ascending to apex. Stipules ovate-triangular or lanceolate, obtuse to acute, 4-10 mm long, 2 mm wide, finely parallel-veined. Inflorescence panic- ulate, terminal, many-flowered, 12-14 cm long, 5—7.5 cm wide in the basal half, 3.5- 4 cm wide in upper half; flowers pedicellate, pedicels fili- form, 10-13 mm long, articulate 1-2 mm above base, dilated below apex, spreading in fruit. Se- pals membranous, ovate or elliptic-ovate, ob- tuse, 8 mm wide, 5 mm long. Petals pink, ob- ovate, rounded at apex, narrowed to the base, 15 mm long, 11 mm wide. Anthers linear-oblong, 4 mm long, acutely apiculate, subsessile; fila- ments 0.3 mm long. Staminodes spatulate, rounded at apex, 7 mm long, 2.5 mm wide above, adnate 1 mm above the base with dimorphic lateral appendages, the longer appendage deeply cut into an elongated subulate, simple, distal seg- ment 5.5-6 mm long attached to a ligulate mul- tifimbriate portion, and an inner proximal short- er appendage 5 mm long with lateral fimbrillate segments. Pistil 9 mm long; style subulate, 5 mm ong; ovary narrowly conic, 4 mm long, | mm wide. Capsule oblong-conic, 10-11 mm long. — Specimens examined. VENEZUELA. TERRITORIO FE- L AMAZONAS: Depto. Atabapo, Cerro Marahuaca, below Salto Los Monos on tributary of headwaters of Rio Iguapo, 3°35'N, 65?23'W, 1,500-1,600 m, 11 Mar. 1985, Liesner 18511 (MO, VEN); Rio Yameduaka ar- riba, 3°38’N, 65?28'W, 1,225 m, 17-18 Feb. 1985, Liesner 17624 (MO, VEN); “Sima Camp,” southcen- tral portion of forested slopes along eastern branch of Caño Negro, 3?43'N, 65?31'W, 21-22, 24 Feb. 1985, Steyermark & Holst 130565 (MO, VEN); below Salto Los Monos on tributary of headwaters of Río Iguapo, 3°35'N, 65?23'W, 1,500 m, 13-14 Oct. 1983, Steyer- mark 129649 (P, holotype; MO, VEN, isotypes). Sastre notes that the species is well marked by its 1987] STEYERMARK— VENEZUELAN GUAYANA 97 FiGure 6. Tyleria sewers — A. Habit of flowering branch.—B. Sterile leafy branch showing terminal clusters of leaves. — C. Detail of leaf s lower surface. — D. Anther. — E. Anther and staminode, ventral view.— F. Staminode, dorsal view. nih Cap apiculate anthers and contrasts it with T. spectabilis species in the piis pedicels in terminal panier Mag. & Wurd. and T. floribunda Gl. because of its broader and lon sho shorter hon It also resembles T. linearis Gl. of ad- er petals, and d larger fistibriate lateral appendages of is jacent Cerro Duida in leaf shape but differs from that — staminodes 98 ANNALS OF THE MISSOURI BOTANICAL GARDEN Sauvagesia marahuacensis Steyerm., sp. nov. TYPE: VENEZUELA. Territorio Federal Ama- zonas: Depto. Atabapo, Cerro Marahuaca, Rio Yameduaka arriba, 3?38'N, 65?28'W, 1,225 m, Ronald Liesner 17677-A (holo- type, MO; isotype, VEN). Figure 7. Suffrutex 0.5-metralis, caulibus virgatis; foliis sub- sessilibus anguste linearibus vel subulatis apice cus- nervis elevatis striatis; floribus ad apices ramulorum pedicellis 3-3.5 mm oe sepalis lanceolatis acutis 5- 5. 5 mm m longis, = a m latis; saree era 7-7.5 mm longis, supra mediu 5 mm latis; uri 5 coronae squamulis X HABI corona l-seriata basi cum filamentis in columnam brevem 0304) mm latam coalita, squamulis quinque 1.2-1.5 mm longis, laminis subulato-spathulatis apice rotundatis m 7-0. 9 mm longis i in stipitem 0. 5m m lon- gum 2.7 mm ORR 0.2-0.25 mm latis. Virgate subshrub 0.5 m tall. Stems dichoto- mously branched or 3—6-verticillate, densely fo- liose toward the apex, leafless for most of the length below, 2-3 mm diam. Stipules densely crowded, overlapping, ge toe ei lanceolate, 2.5-3 mm long, 1 mm wide, dorsally carinate, conspicuously pectinate from base to apex with 15-18 subulate appendages on each side, each one tipped by a filiform, white cilium 1 mm long, lower appendages often deciduous. Leaves sub- 15 times longer than broad, cuspidate at apex, inconspicuously narrowed at base, margins revo- lute, minutely glandular serrulate, each margin with 10-11 minute, appressed-ascending teeth 0.1 mm long, upper surface marked with small transverse depressions. Flowers numerous near the apex, conspicuously pedicellate; pedicels 3— 3.5 mm long. Sepals lanceolate, acute, 5-5.5 mm long, 1.7 mm wide. Petals white, obovate, round- ed above, narrowed to a subcuneiform base, 7- 7.5 mm long, 4-5 mm wide. Stamens 5; anthers linear, 2.1-2.7 mm long, 0.2-0.25 mm wide; fil- aments 0.6 mm long. Staminodes 5, subulate- spatulate, rounded at apex, 1.2-1. m long, arrowed basally, the claw 0.5—0.7 mm long, 0.2 mm wide, the laminar portion 0.7—0.9 mm long, the summit of the laminar portion attaining one- third to one-quarter length of anther. Ovary ovoid, 1 mm long; style subulate, 2.9-3 mm long. [Vor. 74 This species differs from S. guianensis (Eichl.) Sastre and varieties in the narrower linear-subu- late leaves 10-15 times longer than broad with depressed areas on the upper surface and in the long-pedicellate, more numerous flowers con- spicuous at the ends of the leafy stems as con- trasted with the solitary, sessile or barely pedi- cellate flowers largely hidden amongst the leaves of S. guianensis. SAUVAGESIA GUIANENSIS AND VARIATIONS A study of Sauvagesia guianensis (Eichl.) Sastre (1970) reveals much variation not only in details of leaf morphology but in those of staminodial form as well. This taxon (sensu lato) is distrib- uted in the region of the Guayana Shield, with its greatest concentration in the Venezuelan uayana, but with outliers in adjacent Guyana and Colombia (Sastre, 1970). It was originally described by Eichler as Leit- gebia guianensis (op. cit.), based upon a collec- tion by Richard Schomburgk from the savannas of Guyana. Gleason (1931) referred collections of Tate from the summit of Cerro Duida to this taxon. In 1946 Lasser described a Steyermark collection, also from the summit of Duida, as Leitgebia gleasoniana. At this time Lasser (1946) associate helps collection from Cerro Pa- raque (Sipapo) as conspecific with the Steyer- mark type. Sastre (1970) eventually transferred Leitgebia guianensis to Sauvagesia and referred later col- lections originating from various parts of the Venezuelan Guayana as one taxon. The vari- ability of this taxon becomes obvious upon more detailed study. It is noted that specimens origi- nating from p Gran Sabana of eastern Vene- zuela and adjacent Guyana near the type locality have leaves ain fewer times longer than broad, relatively shorter and with shorter-point- ed apices than those from the summits of various table mountains westward in Venezuela. More- over, the staminodes of specimens from the east- ern sector of the range have shorter laminar por- tions. Additionally, a surprising degree of variation is shown between plants collected from various sandstone table mountains. For exam- ple, plants from the summit of Cerro Guaiqui- nima in Bolivar have leaves with only 2-4 glands on sach margin that appear in the upper three- other parts of the range ‘manifest 6-14 glands on each mar- gin, which are distributed from one-fourth to 1987] STEYERMARK— VENEZUELAN GUAYANA 2.6mm Le = 11mm > NM j N V Y KU SE > A = Z z Z AU MU nd [LL — Du o dA E = 16mm 2.9mm 13mm 2.7mm 11.5mm 9.5mm 2.7mm 7.5mm 5.5mm FIGURE7. A-F. Sauvagesia marahuacensis. — A. Habit. — B. Leaf, lower side. — C. Leaf, upper side. — D. Detail of upper apical portion of lower side of leaf. —E. Petal.—F. Po rion of stamens and staminodes attached to membran eek baal Sauv ] j sp. ensis:—A!'. Leaf, lower side. we Apical portion of rtion of ig pee showing anther fer staminode. — B! t. . gui aeni subsp. gleasoniana:—B. p f, lower side. — B?. Leaf, upper side. — B?. Apical conta of leaf. — ^. Mar- i C'-C*. S sipapoensis: — C'. Leaf, lower side. — C". Marginal leaf glands. — C^. Apic droecium showing anther and staminode.— D'-D*. S. guianensis subsp. guaiquinimensis: —D'. Lea per , Up side. — D?. Leaf, lower side. — D?. Apical portion of leaf. — D+. Marginal leaf glands. — D5. Portion of androecium showing anther and staminode. — D*. Petal. 100 halfway up the leaf margin to the apex. Further- more, petals of the Cerro Guaiquinima plants have acute to subacute apices, whereas those on specimens elsewhere are rounded. Moreover, in contrast to the usually encountered elongated, linear-ligulate, narrowly spatulate, or narrowly elliptic laminar portion of the staminode, plants from the summit of Cerro Sipapo (Paraque) have developed a suborbicular type about as broad as ng. Results obtained from a study of the available material in NY and VEN herbaria indicate that Sauvagesia guianensis has undergone differen- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 tiation in various portions of its geographical range resulting in populations showing a diver- gence in various characters. This geographical isolation on several of the sandstone table moun- tains is recognized in the present study as cor- related with taxonomic characters sufficiently distinct as to be considered of subspecific sig- nificance. The following key encompasses the principal differences by which these variations may be rec- ognized within S. guianensis as well as distin- guishing them from the related S. marahuacen- Sis. KEY TO SAUVAGESIA GUIANENSIS AND SUBSPECIES, AND RELATED SAUVAGESIA MARAHUACENSIS la. Flowers on pedicels 3—3.5 mm long, numerous and conspicuous near apices of the leafy stems; leaves linear-subulate, 0.8-1 mm wide, 10-15 times longer than broad; upper leaf surface with v l depressed areas S. marahuacensis essile = wers subs, or on pedicels to 2 mm long pedicellate, few and mainly concealed amidst the leaves; leaves linear-oblanceolate, 1-3 mm wide, 3-10 times longer than broad; upper leaf surface ere A depressed areas s 3-3.5 m m long, 1-1.5 mm wide; Colombia ...... 2b. Saino des 1-1.9 mm long, 0.2-0.5 mm wide; Venezuela 2 Gu E oe subsp. araracuarensis 3 3a. = liar glands 2-4 on en margin, the lowest starting at 4 ne E below the apex; petals acute to acute at a ianensis subsp. guaiquinimensis w e ands 6-14 on e margin, the lowest starting from He the distance below the apex; minar p of staminode suborbicular, about as broad as long, 0.5 x 0.4-0.5 mm; upper aieh of leaf blade with a conspicuous puncticulate appearance c. ianensis subsp. sipapoensis 4b. Laminar portion of staminode elongated, longer ie broad, linear-ligulate, elliptic-spat- 0.2-0.3 m ulate, or narrowly Dm 0.8-1.9 x m; upper surface of leaf blade lacking a puncticulate appear: 5a. Foliar — 6-1 “Ë on amam ina ma oblanceolate or narrowly oblanceolate, 8- 11 mm ong, 1.5-3 mm wide, 3—4 to bluntly pointed; ane ee pr stamino 8 mm a ong mes longer than broad; leaf apex shortly acute de 0. Fi is subsp. guianensis io o mm long, 1-2 mm wi S. gu . Foliar glands 11-14 on Mrd margin; leaves linear or Uns ane n 10-18(-22 , 7-10 times To than broad; leaf apex prolonged, acu- minate; laminar portion a staminode 1.2-1 m m . S. guianensis subsp. gleasoniana la. Sauvagesia guianensis (Eichl.) Sastre subsp. guianensis. Sauvagesia guianensis (Eichl.) Sastre, Caldasia 10: 570. 1970. Leitgebia guianensis Eichl. in Martius's Flora Brasi- liensis 13(1): 413. pl. 83, fig. 2. 1871. TYPE: Guyana. Rich. Schomburgk 1553. Figure A'—A* Leaves oblanceolate, acute to bluntly pointed at apex, 6-11 mm long, 1.5-3 mm wide, 3.5-4 times longer than broad, foliar glands (6—)7-11 spicuous among the leaves; subsessile to 2 mm long pedicellate. Petals obovate, rounded at apex, 6.8-7.5 mm long, 5 mm wide. Anthers 2.5-3 mm long. Staminode lamina narrowly to broadly el- liptic-spatulate, rounded to subacute, 0.8-0.9 mm long, 0.3 mm wide, the narrower lower portion about equaling the lamina in length. Distribution. Guyana and adjacent south- eastern Venezuela, Bolívar. Additional E gee) examined. VENEZUELA. BOLÍVAR: km 1 fEl Dorado, ade of Rio Sacaica, 1,200 m, Soda Md VEN); 52 km N o ma-meru, carre Do ts Sta. Elena, 5?40'N, 61?25'W, 1,300 m, P9» Steyermark & Manara 2201 (MO, VEN). 1987] lb. Sauvagesia guianensis subsp. gleasoniana asser) Steyerm., comb. nov. Leitgebia gleasoniana Lasser, Bol. Acad. Ci. Venez. 9: 246. 1946. TYPE: VENEZUELA. b ades Federal Amazonas: Cerro Dui Savanna Hills, Aug. 1944, Cane 2 (holotype, VEN; isotype, F). Figure 7B'!—B>. Leaves linear or linear-lanceolate, aristate long- pointed at apex, 10—18(—22) mm long, 1-2 mm wide, 7-10 times longer than wide; foliar glands 11-14 each margin, ascending outwardly from margin, elongated, 0.3-0.5 mm long; upper sur- face usually marked with iUe transverse stria- tions. Flowers solitary, few en and incon- spicuous among the leaves, iban to 1-2 mm pedicellate. Petals obovate, rounded at apex, 8 mm long, 4.5 mm wide. Anthers 2.3-2.9 mm long. Staminode 1.2-1.9 mm long, the laminar portion linear-ligulate or narrowly spatulate, ob- tuse or rounded, 0.3-1.2 mm long, 0.2 mm wide, much longer than wide, the lower stipitate por- tion 0.5-1.3 mm long. Distribution. Summits of Cerro Duida and uachamacari, Territorio Federal Amazonas, Venezuela. Additional specimens examined. | VENEZUELA. TERRITORIO FEDERAL AMAZONAS: Cerro Duida, 3?40'N, 65°45'W, 1,500 m, Steyermark, Brewer- Carias & Lies- ner 124569 (MO, VEN); plateau of Huachamacari, 3?50'N, 65?43'W, 1,720 m, 1 Mar. 1985, Liesner 18057 (MO, VEN); Savanna Hills, Cerro Duida, Steyermark 58248 (F, VEN). lc. Sauvagesia guianensis subsp. sipapoensis Steyerm., subsp. nov. TYPE: VENEZUELA. TERRITORIO FEDERAL AMAZONAS: Serrania Sipapo, cumbre, sección speed 5?N, Vli 1,500 m, Ste ark, Bre Carias & Liesner ee. eae Mk iniit VEN). Figure 7C!—C4. A subsp. guianense squamulorum laminis suborbic- ularibus apice late rotundatis 0.5 mm longis, 0.4— 0.5 mm latis, foliis supra conspicue cellulosis re- cedit Leaves linear-lanceolate, ending in a long cus- pidate apex, 10-13 mm long, 2-2.5 mm wide, 5-6 times longer than broad; foliar glands short, 7-10 each margin, incurved-appressed, the low- est ones beginning * the distance below the apex; upper surface with a conspicuous cellular ap- pearance. Flowers solitary, few, subsessile, in- conspicuous among the leaves. Petals obovate, rounded at apex, 5.5 mm long, 4 mm wide. An- STEYERMARK — VENEZUELAN GUAYANA 101 thers 2-2.5 mm long. Staminode with the lam- inar portion suborbicular, broadly rounded, 0.3— 0.5 mm long, 0.4—0.5 mm wide, stipitate lower portion 0.7-0.8 mm long. Distribution. Summit of Cerro Sipapo, Ter- ritorio Federal Amazonas, Venezuela Id. Sauvagesia guianensis subsp. guaiquinimen- sis Steyerm v E quinima, cumbre, sector occidental, cerca del borde cubierto con bosque bordeando una sabana, 5?45'N, 63?43'W, 1,540 m, 27 May 1978, Steyermark, Berry, G. C. K. & E. Dunsterville 117498 (holotype, MO; iso- type, VEN). A subsp. guianense atque ceteris petalis apice acutis vel subacutis e latere 2- 4 secus longitudinem s superiorem 3⁄4 Lx recedit. Leaves linear-oblanceolate, acute at apex, 6— 9.5 mm long, 1.5-2 mm wide, 4—6 times longer than wide; foliar glands 2-4 each margin, in- serted along the upper 34 of the margin, ap- pressed, short; upper surface with few or no transverse lines but with a cellular appearance. Flowers solitary, few, hidden and inconspicuous among the leaves, subsessile. Petals obovate, acute at apex, 5.5 mm long, 2 mm wide above middle. Anthers 2.7 mm long. Staminode 1.6 mm long, the laminar portion ligulate-oblong or ligulate-obovate, rounded or subobtuse at apex, 1.1 mm long, 0.3-0.4 mm wide, narrowed to a stipitate portion 0.5 mm long. Distribution. Known only from the summit of Cerro Guaiquinima, Bolivar, Venezuela. Paratypes. VRNDACHEA, Cerro Tag e cumbre, sector suroeste , $38 N, 63?45'W, 1,650 m, Steyer- mark, Berry, G. C. K. & E. ud 117434 (MO, VEN); Cerro ar er cumbre, sector suroeste- central, 5?45'N, 63?35'W, 950 m, Steyermark, Berry, G. C. K. & E. Dunsterville 117474 (MO, VEN). Figure D'-D*. le. Sauvagesia g b (Sastre) Steyerm., stat. nov. o mt gui- anensis var. araracuarensis Sastre, Bull. Mus. Nat. Hist. Paris 35: 1978. TYPE: COLOMBIA. Com. Amazonas: Rio Caqueta, Araracuara, camino a La Sabana, segunda sabana de Tibeyes, 500—600 m, 5 Jan. 1977, Sastre & Reichel 5139 (holotype, P; iso- types, COL, G). 102 Staminodes larger than in the other subspecies, 3-3.5 mm long, 1-1.5 mm wide. Distribution. Known only from savannas over sandstone soils at the type locality. THEACEAE (BONNETIACEAE) Since the publication of “Flora of the Vene- zuelan Guayana — I” (Steyermark, 1984), the fol- lowing additional new taxa are described in Bon- netia. iin bolivarensis Steyerm., sp. nov. TYPE: EZUELA. Bolivar: Ptari-tepui, cumbre, sai N, 61?47'W, 2,400 m, 19 Nov. 1984, Otto Huber 9818 (holotype, VEN; isotype, MO) Frutex l- metralis; foliis dense tonan Ep oblongo obtu: e ) 2.5-5 x 0.8-1.7 cm; sepalis 11- 12.5 x pecs mm; petalis 21-22 x 14-16 mm; stylis tribus 3-3.5 mm lotes fere usque ad basem divisis. Leaves crowded at summit of branches, ob- long-lanceolate, subsessile, narrowed to an ob- tusely acute apex, obtuse at base, (2-)2.5-5 x 0.8-1.7 cm, faintly impressed-nerved on both sides or the lateral nerves not evident, midrib flower oblong-lanceolate, acute to obtuse, 11-12 x 4.5 mm, dorsally carinate, setulose marginally with dark setae 1 mm long; sepals lance-oblong, subacute, 11-12.5 x 4-5 mm, obtusely dorsally keeled basally and apically; petals white, sub- cuneately obovate, subtruncate apically with un- equally rounded sides, narrowed to the base, 21- 22 mm long, 14-16 mm wide at summit, 4 mm wide at base. Stamens numerous, multiseriate; filaments 5 mm or less long; anthers 0.5-0.8 x 0.6 mm; pistil 9 mm long; S 3, 3-3.5 mm long, divided about 73 way dow The larger flowers and larger, prt leaves differentiate this taxon from B. chimantensis Steyerm., B. tepuiensis Kobuski & Steyerm., and B. toronoensis Steyerm. In its deeply 3-parted style it differs from B. tepuiensis. Bonnetia guaiquinimae Steyerm., sp. nov. TYPE: m, 26 May 1978, Julian A. Steyerm Berry & G. C. K. & E. sche 117421 (holotype, VEN; isotype, MO). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 12: =k ` la: hl non lan rutex 1.5-metralis; f: ceolatis Ma oblanceolatis apice acutis basi subobtusis vel obtus 6 x 1.3-1.6 cm subtus enerv petalis 20 longis; stylo subulato apice leviter 3-lobato Leaves coriaceous, entire, oblong-lanceolate to oblanceolate, acute at apex, gradually narrowed to a subacute or subobtuse base, 5-6.5 x 1.3- 1.6 cm, enervate below, the midrib subimpressed below, lateral nerves elevated above; petiole 1— 2 mm long. Sepals coriaceous, suborbicular-ob- ovate, rounded at a shortly cuspidate apex, 10— 12 mm long, 6-8 mm wide above the middle, 3-4 mm wide at base; petals white, obovate, nar- rowed to a subunguiculate base, 20 mm long, 15-20 mm wide at the summit, 2-3 mm wide at base; filaments distinct, 3.5-7 mm long. Anthers 0.7 mm; pistil 9 mm long; style merely 3-lobed at apex. This taxon is characterized by the shallowly 3-lobed style, entire, oblong-lanceolate, acute leaves, which are enervate beneath. distinguished from B. chimantensis Steyerm. by the larger petals, shallowly 3-lobed style and larg- er leaves enervate below. From B. toronoensis Steyerm. it differs in the larger sepals and petals and larger, entire leaves enervate below, while from B. tepuiensis and subsp. minor Steyerm. it is separated by the larger, minutely mucronate sepals, longer filaments, and entire leaves. — Bonnetia cane up Sp. nov. TYPE: VENEZUELA. var: Ptari-tepui, cumbre, 5°45'N, 61°45’ w, 2,360-2,420 m, 23 Feb. 1978, Steyermark, Carrefio, McDiarmid & Brewer-Carias 115645 (holotype, VEN; iso- type, MO). Frutex 2.5 m; foliis sessilibus seer apice acutis majoribus 3.5-4.5 x 1-1.5 cm minute obscureque ser- rulatis; floribus solitariis breviter eR pedun- culis teretibus vel subteretibus 4—6 mm longis; sepalis lanceolatis vel suboblanceolatis acutis 12413 x -5 mm; petalis luteis obovatis apice rotundatis 16-18 x 12 mm; antheris 0.8 x 0.7 mm; stylis tribus 6 mm longis fere usque ad basem divisis. Leaves coriaceous, lanceolate, acute at apex, slightly narrowed to the base, the larger ones 3.5- 4.5 x 1-1.5 cm, obsoletely pinnately nerved, midrib slightly elevated below, microscopically and obscurely serrulate. Flowers solitary, short- pedunculate; peduncle terete or subterete, 4—6 mm long; bracts immediately subtending flower narrowly oblanceolate, acute, 11-12 x 3-4 mm; 1987] sepals subcoriaceous, lanceolate or suboblanceo- late, acute, 12-13 mm, 10-striate, mi- nutely ciliolate, obtusely carinate; petals yellow, obovate, rounded above, 16-18 mm long (pre- anthesis), 12 mm wide near apex, 2 mm wide at base. Filaments 1.5-3.5 mm long (pre-anthesis); anthers 0.8 x 0.7 mm; pistil 10 mm long; 3 styles 6 mm long, free nearly to the base. From the yellow-flowered B. wurdackii Ma- guire, this species differs in the larger, lanceolate, and acute leaves, longer sepals, larger petals (even in bud), longer filaments, and longer style branches. The leaves, furthermore, do not man- ifest the pale punctate stomata that are clearly shown in B. wurdackii. From B. tristyla Gleason it is easily distinguished by the short pedicels, smaller floral parts and smaller leaves; while from B. huberiana Steyerm. it is well separated by the larger, lanceolate leaves, shorter pedicels, and larger floral parts. Bonnetia tristyla Gleason subsp. nervosa Stey- erm., subsp. nov. TYPE: VENEZUELA. Terri- torio Federal Amazonas: Cerro Yapacana, below summit, 3?45'N, 66?45'W, 825 m, 5 May 1970, Julian A. Steyermark & George Bunting 103153 (holotype, VEN). Frutex vel arbor 1.5—4-metralis; foliis supra costa media necnon nervis lateralibus nonnullis conspicue manifesteque elevatis, nervis lateralibus supra inae- qualiter prominentibus; petalis 25-38 x 15-22 mm. Leaves suboblong, oblong-oblanceolate, or ob- long-obovate, obtuse to rounded at apex, nar- Petiole absent or 1-2 mm long. Peduncle 3-4.3 (—6.5) cm long. Sepals 12-20 mm long, the outer 12-15 mm long, the inner 15-20 mm long. Petals 25-38 x 15-22 mm. Mee TERRITORIO FEDERAL o Yapacana, summit, 1,000-1,200 m, 5 May 1970, Steyermark £ Bunting 103103 (VEN); Cerro ieee 2 Jan. | , Maguire, Cowan & Hat dack 30632 (N B. Cerro Yapacana, 1,200 Maguire, Cowan & Wurdack 30665 (NY. VEN); Gene Paratypes. AMAZONAS: Cerr STEYERMARK— VENEZUELAN GUAYANA 103 bie 30 km al SSW de Ocamo, 2°31'N, 65?23'W, 440-600 m, 1-2 Mar. 1984, Steyermark, Berry & De- lascio 130405 (MO, VEN); Cerro de la Neblina, Caño Grande SSW of Cumbre Camp, 1,050-1,100 m, 25 Dec. 1957, Maguire, Wurdack & C. K. Maguire 42498 (NY, VEN); Cerro de la Neblina, summit, Canon Grande slopes E of Cumbre Camp, 1,200-1,300 Maguire, Wurdack & C. K. Maguire 42235, 42179 (NY, VEN); Cerro Avispa, Río Siapa, summit, od N, 65°51'W, 1,510 m, G. C. K. & E. Dunsterville s.n. (VEN); same locality and data, Cardona 3098 (VEN); Serrania de Vinilla, 20 km SW of Ocamo, 2°20'N, 65°22' 6 m, Huber 6168 (VEN); E side of Río Siapa, 65°41'W, 600 m, Huber 6006 (VEN). ue This taxon differs from typical B. tristyla Glea- son in having smaller petals, upper surface of the leaf blade with the midrib prominently elevated, and unequally prominent pairs of lateral nerves, with 6—8 of the pairs prominently elevated al- ternating with finer, lightly impressed interme- diate ones. In B. tristyla subsp. tristyla the large petals 35-42 x 27-30 mm contrast with the smaller ones of subsp. nervosa. Moreover, in subsp. tristyla the upper midrib is shallowly de- pressed, not elevated, and all the lateral nerves of the upper surface of the leaf are equally in- conspicuous and lightly impressed. The leaf blades in typical B. tristyla tend to be larger (5-) 8-15 x 2.5-5.0 cm as contrasted with 4-8 x 1.5-3.5 cm in subsp. nervosa. The base of the leaf blade in B. tristyla subsp. tristyla is cuneately rowed, whereas that of subsp. nervosa is usu- ally slightly obtusely curved or rounded above the junction with the petiole. The geographical ranges of the two subspecies are distinct. Typical B. tristyla, described origi- nally from Cerro Duida, occupies the northeast- ern sector of the range, from Cerro Duida an Marahuaca east to the Meseta de Jaua and Sari- sarinama in Bolivar. Bonnetia tristyla subsp. ner- vosa, in contrast, is found in the more western sector, ranging from Cerro Yapacana south to Cerro Vinilla, Avispa, Aracamuni, and Neblina. The only other species of Bonnetia having large yellow petals, B. steyermarkii Kobuski, is easily distinguished from B. tristyla by the larger acute sepals subtended by 4—6 large sepaloid bracts and the broader, acutely angled ancipital peduncle. The following key incorporates the newly de- scribed taxa of Bonnetia with those previously described by the author in 1984. KEY TO THE SPECIES AND SUBSPECIES OF BONNETIA la. e phan eral nerves of upper leaf surface elevated, impressed on lower surface; leaf blades mainly 2-4 ns 2.5 cm B.t tepuiensis subsp. tepuiensis 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 2b. Lateral nerves of rpg leaf surface faintly impressed, mostly not evident on lower surface; | .8-1.4 cm ades 1-2 x eaf “epp subsp. minor lb. Style divided into 3 anas nearly or all the way to the base, or shallowly 3- lobed at the a a. Style shallowly 3-lobed B. pip 3b. Style divided into 3 branches, parted halfway or more to the base. Petals white or pin 5a. Petals 21-22 mm n long; mg subacute, 1 1-12.5 mm long; leaves obtusely acute at apex, the larger 4-5 x 1.2-1.7 cm ... B. bolivarensis cA i-a subacute, 1.2-3.1 x 0.5-1.3 cm . Petals 9-12 mm long; ra obtuse or rounded at apex, 8-9 mm long; Tee) acute to 6a. mihi in a terminal rosette; upper leaf surface enervate or nerves faint; petals B. chimantensis m broad 6b. pes imbricately extending on the branch below its tip; upper leaf surface im- pressed-nerved; petals 9-10 mm broad 4b. Petals yellow. toronoensis 7a. Agro elongate, 3-9 cm long, often surpassing the leaves, ebracteat 8 ds elev 8b. Petals iiir 2.5-3.8 x 7-3 cm; lateral nerves of upper leaf surface ails impressed but ted; upper midrib shallowly depressed B. tri a subsp. tristyla ll sty 1.5-2.2 cm; lateral nerves of upper leaf eiie unequally impressed, 6-8 pairs iba elevated; upper midrib — B. tristyla ges nervosa N c er. . Peduncle lacking or at most 1.6 cm long, usually hidden among the leaves or mu 9a. socii linear-oblanceolate, 2.5-7 mm wide; petals 8 mm long; peduncle 8-16 m Sea 9b. Leaves broadly ME Vasin, or oblong-oblanceolate, 8-15 mm wide venas n vegetative shoots som wider); peduncles 3-6 mm long. 102. Leaves broadly ons ioe acute, only slightly narrowed at the base, len nearly the same widt or most of ngth, 3.5-4.5 x 1-1.5 cm; leaves finely teed. -nerved mnn not pale punctate beneath, stomata not manifest; sepals 12-13 mm B. ptariensis o = manifest; sepals 9-10 mm Al LISSOCARPACEAE Lissocarpa "m Steyerm., sp. nov. TYPE: NEZUELA. Territorio Federal Amazonas: Depto. MD Cerro Marahuaca, riverine forest upstream from “Sima” Camp, along branch of Cano Negro, southcentral portion, 3?43'N, 65?31'W, 28 Feb.-1 Mar. 1985, 1,140 m, Steyermark & Holst 130880 (ho- lotype, MO; isotype, VEN). r 10-metralis glabra; foliis subcoriaceis oblon- Sc cm latis, nervis lateralibus numerosis cena 13 mm renee floribus solitariis. bibracteatis supra ax- illari pedicellis 3-3.5 mm velit bins ovatis apice rotundatis 2.5 mm longis, 2-2. zm m latis; corollis cylindricis peru maturo) m longis, basi 1.5 mm latis, medio m latis, lobis. 4 ligulato-oblongis rotundatis 4 x 2 mm, coronae segmentis lanceolatis acutis 1.5-2 mm longis; calyce ig og 6 mm longo, calyce 5 mm Feud l. = m lato, lobis 4 suborbicularibus rotun 2 es is, 2 mm latis, minute glandulari-ciliolatis; ruc anguste elliptico-oblongo in prominentiam obtu ong ; Leaves ias or obovate, obtuse, rounded, or subacute at apex, con- o the m broadest above the middle, 1 1-3.5 x 0.4-1 (- e beneath, pale punctate beneath, the stomata B. wurdackii panua T abrupte angustato basi rotundato 3-3.3 m longo, 1.2-1.7 cm lato, 2.2-2.5 plo longiore quam ue Tree 10 m tall, glabrous throughout. Leaves subcoriaceous, dark green above, paler below, oblong-elliptic to elliptic-ovate, obtusely acu- minate at apex with acumen 0.8-1.2 cm long, obtuse to subacute at base, 7.5—13 cm long, (2-) 3-5.5 cm wide, glabrous both sides; midrib el- evated above, impressed or less elevated below; lateral nerves numerous, faint, about equally prominulous on both sides as the loosely tertiary reticulate venation; petiole 5-13 mm long. Flow- ers solitary, superaxillary or axillary on new or old branches, pedicellate, bibracteate at base; pedicels 3-3.5 mm long; bracts ovate, rounded at apex, 2.5 mm long, 2-2.2 mm wide. Corolla S constricted basal part 2 mm long; 4 corolla lobes ligulate-oblong, rounded, 4 mm long, 2 mm wide; 8 coronal divisions lanceolate, acute, 1.5-2 mm long. Calyx and hypanthium 6 mm long; calyx STEYERMARK 1987] —VENEZUELAN GUAYANA 105 5 mm long, 1.5-2 mm wide, the cylindric tube 4 mm long, 1.5 mm wide; 4 lobes suborbicular, rounded at apex, 2 mm long, 2 mm wide, mi- E glandular-ciliolate on margins. Style 1.5 m long, upwardly thickened to a subtruncate- L. stigma. Fruit yellow-green, narrowly el- liptic-oblong, abruptly narrowed to a bluntly tri- angular protuberance at apex, rounded at base, 3-3.3 cm long, 1.2-1.7 cm wide, the subtending bracts more or less persistent. This species differs from L. benthamii in the differently shaped fruit which is 2.2-2.5 times longer than broad and narrowly elliptic-oblong, and in the solitary flower instead of the few- to several-flowered, subracemose inflorescence. Additionally, its occurrence in the montane for- est on the slopes of Marahuaca at an elevation of 1,140 m is in contrast to the lower altitudes where L. benthamii has been collecte The differences suggested by Gleason (1926) to differentiate L. guianensis Gl. from L. ben- thamii cannot be maintained with respect to the more prominent upper midrib and conspicu- ously reticulate veinlets on both surfaces, sup- posedly characteristic of L. guianensis. Exami- nation of material collected by Liesner from San Carlos de Rio Negro, Venezuela, type locality for L. benthamii, indicates variation in these char- acters, some specimens showing prominently raised upper midribs but with only obscure ter- tiary venation on the ape tolar Space (Liesner 8692; Liesner & Clark 9 1117) have more jeticulain veinlets on the lower surface. Similarly, specimens de- termined by White as L. guianensis (Maguire 34618, 34907) show only obscure venation on the lower surface. The leaves of the Marahuaca specimens are prominently reticulate-veined on both surfaces with the upper midrib manifestly elevated. The leaves are relatively small in size as use with either L. guianensis or L. bentham Although L. benthamii and L. guianensis can- not be well separated on vegetative characters, the larger flowers of L. guianensis may be the best difference in distinguishing the two taxa. Pending further collections of flowering material, the two taxa may at present be considered as separate species. S RUBIACEAE Chomelia stergiosii Steyerm., sp. nov. TYPE: VENEZUELA. Bolívar: Anacoco, Río Cuyuní, entre puesto de la Guardia Nacional Acara- bisi and Anacoco, 2 Aug. 1981, Basil Ster- gios & Gerardo Aymard 2804 (holotype, VEN; isotype, PORT). Arbor 8-metralis, ramulis glabris; foliis elliptico- ovatis vel lanceolato- ellipticis apice e» tuse acutis basi bri e pedunculo 2.5 cm longo glabro; floribus sessilibus; ca- tulis vel reflexis oblongo- spathulatis apice rotundatis 3-5 mm longis (1—)1.7-1.8 mm latis, intus dimidio inferiore sparsim pilosulis; hypanthio extus sparsim a corolla hypocrateriformi, tubo 20 mm longo, m lato ubique glabro, lobis lineari-ligulatis 4 mm loe 1.2-1.5 mm latis. Tree 8 m tall, branches slender, glabrous, spines axillary, 18-19 mm long. Leaves membranous, elliptic-ovate to lance-elliptic, obtusely acute at apex, acute to subobtuse at base, glabrous above, glabrous below except barbellate in the leaf axils and sometimes sparsely pilosulous along some of the lateral nerves, 4.5-7 cm long, 2-4 cm wide, with minute dark dots moderately scattered be- neath; lateral nerves 4—5 each side, slender, slightly sulcate above, faintly impressed below ascending, faintly anastomosing near margin; petiole 4-5 mm long, ciliate on upper margins, elsewhere glabrous, canaliculate above. Stipules triangular-ovate, acute, appressed-pubescent without, ciliolate at apex, 2-3 mm long. Inflo- rescence terminal, long-pedunculate, congested- cymose with 3-6 flowers, ebracteate; peduncle 2.5 cm long, filiform, glabrous; flowers sessile, the central one solitary and sessile, the others on short lateral axes 0.5 mm long. Calyx lobes 4, unequal, the two larger ones oblong-spatulate, rounded at apex, narrowed in the basal third, 3— 5 mm long, (1-)1.7-1.8 mm wide in upper part, | mm wide in lower third, spreading to reflexed, sparsely minutely pilosulous within in lower half, glabrous without, two smaller lobes ligulate, ob- tuse, 2 mm long, 0.8 mm wide, spreading to reflexed; hypanthium clavate-turbinate, 2 mm ong, | mm wide above middle, sparsely pilo- sulous with loosely spreading hairs. Corolla sal- verform, tube 20 mm long, | mm wide just below summit, 0.7 mm wide at base, glabrous within and without, lobes linear-ligulate, rounded at apex, 4 mm long, 1.2-1. wi within, strigillose without mainly above m Anthers slightly exserted, linear, 2 mm long, 2 mm wide, glabrous. Style 13.5 mm long, gla- brous. 106 This species is sympatric with C. delascoi Stey- erm. but is distinguished from that taxon in the much longer corolla, foliaceous, manifestly un- equal, and spreading calyx lobes, which are sparsely pilosulous on the interior surface of the larger pair, and larger leaves. It differs from C. polyantha Blake in the glabrous exterior of the corolla tube, the longer corollas, and the longer, spreading calyx lo I take pleasure in naming this species for Dr. Basil Stergios, director of the herbarium of PORT, who has. activated a well- organized. collecting leagues, has established a in the Venezuelan Llanos of Edo. Portuguesa. Coccocypselum croatii Steyerm., sp. nov. TYPE: VENEZUELA. Bolivar: vicinity of Icabaru, 4°19'N, 61?44"W, 600 m, 25 Jul. 1982, Thomas B. Croat 54112 (holotype, MO). , Herba radicans, caulibus repentibus 2 mm | diam. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 out. Fruit blue, 4 mm long, 7 mm wide, glabrous. This species is related to C. condalia R. & P. of Peru, from which it differs in the larger ovate leaves, shorter petioles, more elongate setose cil- la on the leaf margins, and more numerous, ar- cuately curved lateral nerves. Pagameopsis Steyerm. Recent collections of Pagameopsis have ne- cessitated a reevaluation of the specific and sub- specific elements within the genus. Pagameopsis maguirei Steyerm. subsp. pusillus Steyerm., subsp. nov. TYPE: VENEZUELA. Territorio Federal Amazonas: Depto. Rio Negro, Cerro de la Neblina, Camp III, NW Plateau, 13.5 km ENE of Base Camp, 0°54'N, 66°4'W, 1,750-1,850 m, 16-18 Feb. 1984, Ronald Liesner 16013 (holotype, MO; iso- type, VEN). Figure 8. Suffrutex 0. 1-0. 2- metralis; foliis ad apices! ramorum bris, subulatis elongatis 8-10 mm longis glabris, foliis petiolatis, petiolis 3-4 mm longis, marginibus superi- oribus setoso-ciliatis ceterum glabris; laminis obtusis vel rotundatis 4—4.5 cm longis, 1.5-3 cm latis supra costam mediam pilis setosis munitis atque marginibus dense adpresso-setoso-ciliatis, ciliis 0.5-0.7 mm longis ceterum m glabris, nervis lateralibus Wtfoque latere 7-9 arcuato nase ie longipedunculatis ara floris, pedunculo 2. in 3. cm pos E anthesim 1-1.5 mm longo; calycis lobis lineari-lanceo- latis acutis 2.5-3 mm longis, 0.7—0.9 mm latis omnino glabris; = (immatura) coerulea 4 mm longa, 7 m lata glab Creeping herbaceous plant with rooting gla- brous stems 2 mm diam.; stipular sheath gla- brous except densely ciliate on the summit, pro- longed into a glabrous, elongated, subulate appendage 8-10 mm long. Leaves petiolate; pet- ioles 3-4 mm long, upper margins setose-ciliate, otherwise glabrous; leaf blades ovate, shortly acute at apex, obtuse or rounded at base, 4—4.5 cm long, 1.5-3 cm wide, upper midrib setose- ciliate, margins densely idi s setose-ciliate, the incurved hairs 0.5-0.7 m glabrous; lateral nerves 7-9 each ascending, elevated below. Inflorescence long pe- dunculate, 5-7-flowered; peduncle 2.5-3 cm. Flowers subsessile: aa thium and calyx a brous; hypanthium in post-anthesis 1-1.5 m long; calyx lobes linear-lanceolate, acute, 2.5.3 mm long, 0.7—0.9 mm wide, glabrous through- elliptici vel lan- ceolatis apice obtusis vel subobtusis ‘die Ibis l- 2.5 cm longis, 4-7 mm latis, supra dense adpresso- hirsutulis subtus marginibus dense hirsutulis, nervis is d tis |. m hemisphericis 7-30 mm lo dunculatis; pedunculo erecto 2: 5-4. 5 cm longo la is A mm crasso, ces 4-5 ramorum bre vium inflorescentiae, quoque glomerulo 34. floribus; genes 4—5-meris 4 mm lon- gis, tubo lobisque extus praeter basin glabram dense strigosis, lobis 4—5 Melee uus subobtusis 2 mm longis intus dense lanulosis; bg den lineari- lanceolatis acutis extus den trigoso- “hirsutis intus glabris in sinubus una sra ie mun Dwarf ligneous plant 0.1—0.2 m tall. Leaves densely crowded in an apical rosette, narrowly lance-elliptic or lanceolate, narrowed to an ob- tuse or subobtuse apex, narrowed to a subsessile base, 1-2.5 cm long, 4-7 mm wide, densely ap- pressed-hirsutulous above, densely hirsutulous below and on margins; midrib elevated below, not evident above; lateral nerves not manifest; petiole 2-3 mm long, scarcely distinguishable from the leaf base. Stipular sheaths closely 4—5 short axes, the lowest axes 8-15 mm long, 107 STEYERMARK— VENEZUELAN GUAYANA 1987] EY 3 te a tte wok Pees Se ae Mee ig tror mE Y Ll ris, =,” ` SZ, x? > Á X > “ x, 17 I» es yea ore ee tes! FIGURE 8. Pagameopsis maguirei subsp. pusillus. — A. Habit.—B. Flower, with subtending bracts.—C. Co- rolla, interior view.—D. Calyx and hypanthium, with pistil detached from calyx, interior view. 108 densely hirsute; rachis with 1-3 pairs of hori- zontally spreading bracts 3 mm long, 1 mm wide, hirsutulous without, glabrous, the inflorescence bibracteate at base with the bracts foliose, linear to oblong-lanceolate, subobtuse, 8-15 mm long, 2-3 mm wide, densely pubescent above, sparsely or moderately so below. Peduncle 2.5-4.5 cm long, 1-1.5 mm thick. Flowers in dense nearly sessile glomerules of 3-4 at the ends of short axes; 2 bracts subtending the base of the calyx navic- ular, oblong-lanceolate, 1.5-1.8 mm long, 0.8-1 mm wide, glabrous and glandular within. Calyx lobes slightly unequal, , linear-lanceolate, acute, 1.5-2 mm long, 0.5-0.6 mm wide, dense strigose-hirsute without and on margins, gla- brous within with 1 gland between the sinuses of the lobes. Corolla 4 mm long, densely strigose without except where glabrous at base; tube 2 mm long; lobes 4-5, ligulate-lanceolate, subob- tuse at apex, 2 mm long, densely lanulose within. This taxon differs from the other known taxa of Pagameopsis in the very small, narrow, dense- ly hirsute leaves and dwarf habit. It is closely related to P. maguirei Steyerm. subsp. neblinen- sis in having apical leaf clusters, but differs in the smaller alone m with 3-5 axes, and in the dense pubescen abrous im of subsp. pusillus has been found on nearby Cerro Avispa of Venezuela. Pagameopsis maguirei subsp. pusillus var. gla- brus Steyerm., var. nov. A subsp. pusillus var. pusillus foliis glabris recedit. TYPE: VENEZUELA. Territorio Federal Amazonas: summit of Cerro Avispa, Río Siapa, 1?30'N, 65°51'W, 1,510 m, 5 Dec. 1972, Cardona 3096 (holotype, NY; isotype, VEN) Another variation of Pagameopsis maguirei from adjacent northern Brazil is here described. KEY TO THE SPECIES, SUBSPECIES, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Pagameopsis maguirei subsp. neblinensis Stey- erm. var. pirapucuensis Steyerm., var. nov. TYPE: BRAZIL. Serra Pirapucü, 1,300 m, 26 Jan. 1966, Nilo T. Silva & Umbelino Brazao 60880 (holotype, NY). Frutex; foliis caulinis dispersis haud omnino api- calibus, longitudinem caulis 2.5-4 cm oc occupantibus, oblanceolatis apice subacutis 8-8. 5 cm longis, 1.3-1.5 cm vel saltem 1 parte costa media inferiore costae mediae pilosis, subtus c -10mm tis, moderatim sir pud strigosis, pee dense hirsutis. Shrub; leaves arranged along the uppermost length of the stem, not wholly apical, oblanceo- late, subacute at apex, 8-8.5 cm long, 1.3-1.5 cm wide, appressed-pubescent above or at least pi- lose on the lower part of the midrib, abundantly substrigose or pilose below, otherwise sparsely puberulous; petioles abundantly strigose or sub- strigose throughout. Stipular sheaths elongated, 7-10 mm long, wide, moderately to densely strigose, the summit densely hirsute. This taxon differs from the other varieties of subsp. neblinensis in the more densely strigose stipular sheaths, which are more densely hirsute at the summit. From var. neblinensis it differs in having the leaves dispersed along the stem length or 4 cm, and in the pilose petioles and lower surface. From subsp. neblinensis var. angustifoli- us it differs in the leaves pubescent below and above at least along the midrib or surface. From subsp. maguirei var. maguirei it may be differ- entiated by the pubescent upper midrib and low- er leaf surface. In order to accommodate the above newly de- scribed taxa, the following revised key to Pa- gameopsis is offered. AND VARIETIES OF PAGAMEOPSIS la. Verte: aiia spaced along the upper part of n for 2.5-11 cm; stipular sheath elongated, usually longer than broad or as DR as broad, POD 2a. aa ‘of leaf margins EE. haire di 4: very e 0.5—1 mm long; inflorescence with ner 3-5 densely flowered eee on i2 pairs of lateral l. axes, the whole inflorescence 1.5-4 cm ong, 2-4 cm wide; leaves mainly 175-3!4 times longer P. ee N og . Cilia of leaf m long; inflorescence branche the numerous margins papi 1 or deniers to ascending, less than 0.5 m glom mall rules borne on 7 or more branched axes, whole inflorescence 5— TE. cm long, 3-6 cm wide; pen mainly (2%-)3—6 times longer than broad 3a. Lower d surface glabrous Or mainly so, sometimes pie upper leaf surface glabrous 4 a. Ud5 »511U1 ti lobe nd mm 1 long, i 1. 1.25 mm wide . lobe S GCNSC Milivilvuss NS P. naue subsp. maguirei var. maguirei 4b. Petiolar base Meade above, gl to sparsely pubescent below; interior of calyx lobes 1987] STEYERMARK — VENEZUELAN GUAYANA 109 with short strigose pubescence, the exterior with pies hirtellous pubescence; corolla 2.2-3 mm long, 0.7-1 mm wide lo 3b. Lower leaf surface pubescent; upper | magui rei su € neblinensis var. Mise E labrescent, the midrib pubesc maguirei ake sp. neblinensis var. pirapucuensis lb. Leaves apically ee = the be AR 0.5- d 5 cm length of stem; stipular sheath c tracted, m than lon s broad as long, 1-6 mm lon 5a. Leaves 2-10 cm ae 0. 7-1. 8 cm wide; bracts subtending base of inflorescence ve "d mm long, 4T mm wide; inflorescence branching into numerous sap glomerules borne in 7 or more branched axes, the whole inflorescence 5-11 cm long, 3 m wide; shru cA Ss long, 2-3(-5) mm wide; inflorescence with o pairs of lateral axes, the whole inflorescence 0. T d cm er . Leaves 1-2.5(-3) cm long, 0.4—0.7 cm wide; Pra ts subtendin g base y 3-5 yaspa i glomerules borne on 2 long, 2-3 cm wide; d ucl ligneous plant 6a. Upper and lower leaf surfaces densely hirsute „u P. maguirei subsp. pusillus var. pusillus 6b. Upper and lower leaf surfaces glabrous P. maguirei subsp. pusillus var. glabrus Psychotria guanchezii Steyerm., sp. nov. TYPE: Guanchez 3644 (holotype, VEN; TFAV). Figure 9. isotype, Suffrutex 1.5-metralis, ramis glabris; stipulae vagina uobus 16-20 utroque latere divuehis pate ibus; inflores- centia terminali pedunculata, pedunculo gracili erecto 3 cm longo, 1 mm lato minute sparsimque puberu- puberul entibus; bracteis sub fasciculis florium ligulatis vel lanceolatis 2.5-3 mm longis utrinque minute den- seque puberulentibus; calyce hypanthioque 1.5-2 mm opan; calycis lobis deltoideis obtusis 0. 3 mm longis, m latis; i 5—6 mm lon- a extus praeter basim glabram ipsa dense puberulenti pilis patentibus 0.1-0.2 mm longis munita "i dimi- a parte superiore puberula ceterum glabr: Slender subshrub 1.5 meters tall, branches gla- brous. Stipular sheath 2—2.5 mm long, 1.8-2 mm wide, glabrous, terminating on each side in 2 subulate minutely ciliolate teeth 3 mm long. Leaf base, glabrous both sides, 4—9.5 cm long, 0.8-2 ide, midrib elevated ni impressed bove; lateral nerves 16—20 side, divari- cately spreading at an angle of 5— ioe faintly anas- tomosing at 1-2 mm from margin; petiole 3—9 mm long, glabrous. Inflorescence small, cy- mosely umbellate, terminal, 1.2-1.8 cm long, 1.3- wide, on erect, slender peduncle 4—4.3 cm long, 1 mm wide, minutely and sparsely pu- berulent; axes of inflorescence 4, three lateral ones spreading, 4-6 mm long, 0.5-1 mm wide, one central axis erect and longer, 7-8 mm long, 1 mm wide, all moderately to abundantly puber- ulent with spreading unequal puberulence, the longest hairs 0.1 mm long; 3 lateral axes 5—6- flowered, 4th axis 7-8-flowered. Bracts subtend- ing base of inflorescence divaricately spreading, subulate, 2.5-2.7 mm long, 0.3 mm wide, mi- nutely and moderately puberulent; bracts sub- tending flower clusters ligulate to lanceolate, acute to obtuse, 2.5-3 mm long, 0.7 mm wide, mi- nutely densely puberulent on both sides. Calyx and hypanthium 1.5-2 mm long; hypanthium shallowly campanulate, l x 1 mm; ca m bt wide. Corolla Infundibuliform, 5—6 mm long; aie 2.2-3.5 mm ng, 0.8 mm wide at base, 1.3 mm wide at ori- fice, dency puberulent, except at very base, with short spreading hairs 0.1—0.2 mm long, pubes- cent in upper half within from staminal insertion to orifice, elsewhere within glabrous; lobes 4, lan- ceolate, subacute, 1.7-2.5 mm long, 1.3 mm wide, densely puberulous without. Anthers linear, 1.1— 1.2 mm long, included in uppermost part of co- rolla tube; filaments 0.8 mm long, inserted half- way up corolla Tube Style 4.2 mm long, aah w ü d, dilated, exserted, 0.8 m This species is related to P. capitata ^"; & P. but is reduced in the size of all parts. Moreover, the densely puberulous calyx and minute, puber- ulous corollas, small inflorescence with slender peduncle, and narrow leaves are noteworthy dis- tinguishing characters of the taxon. From P. pi- resii Steyerm. of Amapá, Brazil, it is differen- d =] 110 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 4mm yi! À Ade dad j key "tul ,^ i UE L. fed C FiGURE9. Psychotria guanchezii. — A. Habit. — B. Corolla. — C. Calyx and hypanthium, with position of disk indicated. — D. Corolla, pre-anthesis. 1987] tiated by the smaller, 5-merous corollas, shorter stipular teeth, and smaller inflorescence wit fewer axes. Psychotria ronaldii Steyerm., sp. nov. TYPE: VENEZUELA. Territorio Federal Amazonas: Depto. Atabapo, Cerro Huachamacari, for- ested slope, 3?39'N, 65?42'W, 600-750 m, 4 Mar. 1985, Ronald Liesner 18214 (holo- type, MO; isotype, VEN). Figure 10. Frutex 2-metralis, caulibus glabris; stipularum va- lateralibus 4—6 patent briato- PRA pep pe edun culo erecto 1.8—2.5 l- longo, mm crasso glabro; floribus sessilibus cient s" videtur n ullis; h mm longo glabro, calyce breviter 5- lobato, dentibus late deltoideis vel fere truncatis; corolla subinfundibulifor- .5 mm longa extus glabra, intus infra insertionem “Yuhu ways pilosulo ceterum glabro, lobis 5. Shrub 2 m tall; stems glabrous. Stipular sheaths broadly semicircular, rounded or slightly acute at apex, buff-pubescent within and on the apical margins, 2.5 mm high, 5 mm wide. Leaves broadly elliptic-obovate, caudate-acuminate at apex with acumen 2 cm long, acutely narrowed to a decurrent base, 22-30 cm long, 8-10.5 cm wide, glabrous both sides; lateral nerves 14-17 each side, widely spreading at an angle of 15° 20°, elevated below, tertiary venation beneath finely grossly reticulate. Petioles 1—1.5 cm long, glabrous. Inflorescence terminal, paniculate, subhemispheric, slightly broader than long, 2 cm long, 2.5-3 cm wide at base; lateral axes divar- icately spreading, the lower ones 7-8 mm long, the upper 2-4 mm long; base of axes with fim- briate-pubescent scars; axes terminating in 7—10- flowered clusters. Peduncle erect, 1.8—2.5 cm long, 1-1.5 mm thick, glabrous. Flowers in groups of 7-10. Calyx and hypanthium 2.1 mm long, gla- brous; hypanthium | mm long, | mm wide; calyx | mm long, 1.2 mm wide, the border shallowly toothed; calyx teeth broadly deltoid to nearly truncate. Corolla cream-colored, subinfundibu- liform, 4.5 mm long, 2 mm wide above, | mm wide at base, glabrous without, glabrous within except for a pilosulous zone at the antheriferous STEYERMARK— VENEZUELAN GUAYANA 111 level; lobes 5, a lanceolate, subacute, 1.5 mm long, | mm w The immediate P > 80% (3) 0.01854 90% > P > 80% (4) 0.0096 98% > P > 95% Level of significance is set at 0.05. The null hypoth- esis is proved in each tively. Figures 8 and 9 show Anaphase II (AII) diagonal bridge and Telophase II (TII) excluded chromosomes, which also occur at a total fre- quency of 6.36% while normal AII movement occurs at a frequency of 93.64%. dere anal- yses of these data (Table 2) show that pollen viability percentage and the frequencies of AI and AII normal movements and the formation of 10 chromosome bodies per PMC during the first prophase to metaphase (93.56%, 93.64%, 93.51%, and 93.39%, respectively) are not sig- nificantly different; that the frequency of for- mation of nine bodies per PMC is not signifi- cantly different from the frequency of occurrence of laggards and excluded chromosomes at TI; and that the total frequencies of formation of nine and 11 bodies, AI bridge formation and TI chromosome exclusion, AII diagonal bridge for- mation and TII chromosome exclusion, and per- centage pollen abortion are all not significantly different. In the formation of nine bodies, the hexavalent second largest pair (cf. chromosome bodies in Figs. 2-4). When this association occurs, the first 1987] and second large pairs become about equal in length at MI (see Fig. 4; see also mitotic chro- mosome lengths of the two large and eleventh ofits own (in 11 separate bivalents), or associates with any homologues other than the second larg- est, meiotic behavior becomes erratic and meiot- ic products are inviable. DISCUSSION Some representatives of Urginea in other areas have been shown to have B chromosomes (De Wet, 1957; Jones & Smith, 1967), but accessory chromosomes have not been reported in any of the west tropical African materials. Where they have been reported, these accessory chromo- somes are known to be different from the auto- somal members of their complement. They are heterochromatic, their numbers vary within populations, their transmission at mitosis is not regular, and at meiosis there has not been any mechanism of transmission comparable to that shown in U. altissima, where association of the extra pair with a specific pair of autosomes is the mechanism for successful transmission of the ex- tra chromosomes into the spores and thereby into the next generation of the plant. Hence the extra pair of chromosomes in U. altissima can- not be seen as essentially inert as in the cases of the B chromosomes reported in other species. The formation of a hexavalent involving the extra pair and two other pairs may indicate some genetic affinities between the three pairs involved (Battaglia, 1964), or show the probable origin of the extra pair (Wedberg et al., 1968). However, the formation of an association between the extra , the second large pair, with a much higher NN than in the hexavalent formation, and leading to successful meiotic behavior, may identify the true origin of the extra pair. This extra pair cannot be regarded as accessory because it is indistinguishable from the other pairs in mitotic behavior and structure (cf. chromosomes in Clarkia unguiculata, and C. williamsonii in Mooring, 1960, and Wedberg et al., 1968, respectively). The extra pair behaves normally at mitosis as in the large B chromo- somes of Brachychome lineariloba (Carter & Smith-White, 1972), but its successful transmis- sion from one generation of the plant to another depends upon a meiotic mechanism of its as- sociation with a specific pair of autosomes in the OYEWOLE— URGINEA ALTISSIMA 129 complement. Hence, it differs from the B chro- mosomes of B. lineariloba. The origin of ie extra pair is undoubtedly betrayed in its association with the second large pair — only such association leads to viable spore formation. The mode of origin of the extra pair is probably by fragmentation at a heterochro- matic region along the second (short) arm of the parent (second large) autosomal pair. This is why the point of breakage on the otherwise acentric fragment is able to exercise secondarily a spindle- fiber organizing function at mitosis for effective polar movement—and, hence, normal autoso- mal behavior— but which is incidentally too weak to effectively organize the movement of the frag- ments at meiosis whenever they form a bivalent of their own. They then remain excluded on the equatorial plane after polar movement of the other chromosomes. Again, in the formation of 11 normal biva- lents, incomplete pairing or total asynapsis in the extra pair, which would lead to early repulsion ofits members prior to anaphase movement, was not observed at all. That is, the extra pair shows synapsis (and probably chiasma formation). This means that (1) the extra pair contains essential 1 ing, (2) the extra pair came from homologous segments of a homologous pair of parent chro- mosomes, and (3) the extra pair carries genic matter essential to the survival and success of the plant. This is why its transmission is effected by a genetic mechanism. The failure of this mechanism ultimately leads to the formation of inviable meiotic products. The behavior of this extra pair during cell di- vision is of interest. The incidence of fragments within chromosome complements has been re- ported, both in nature and in experiments, by tution, or rejoin with broken ends of other chro- mosomes (Lea, 1946; Hair, 1952; Giles, 1954). If restitution is to occur, it occurs immediately after the breakage and is dependent on oxidative enzyme metabolism (Wolff & Luippold, 1955). The extra pair of chromosomes (fragments) in U. altissima has healed broken ends in somatic cells and hence behaves as a normal autosomal pair during mitosis. During meiosis, however, the ends either remain healed and the pair forms a bivalent of its own, which becomes excluded 130 during polar movement, or the broken ends be- come reactivated and thereby the pair associates with other chromosome pairs. Such a behavior has not been reported earlier. Rees (1958), how- ever, working on Scilla, reported the behavior of chromosome fragments in pollen mother cells. In Scilla, the fragments become attached either to their parent autosomes or to any other chro- mosomes, depending on the relative distance of the fragments to one or the other. The attach- 1 ment may | yn- chronous attachment must take place within a sticky matrix. As no sticky matrix was found in U. altissima, there was no evidence to suggest that the association of the extra pair with the second large autosomal pair was anything but synchronous. Hence the situation in U. altissima differs from that of Scilla in that it shows clear evidence of a genetically controlled mechanism of association, thereby stabilizing the resultant genetic system to ensure a high percentage fer- tility of sexual reproduction. This is in keeping with the suggestions of Blackwood (1956) and Rutishauser (1956). It will be interesting to find out what initiates the mechanism of healing and reactivation of the broken ends of the fragments and the second large autosomal pair during mitosis and meiosis. It is likely that the stage of development at which the dissociation occurs is the interphase between the end of spore formation and the beginning of spore development, while the reactivation of the broken ends would take place at the onset of meiosis in t ual reproduction are = generally controlled hor- monally, these events are likely to be part of the effects of the hormonal control, which in itself is genetic. This allows repetition of this event whenever necessary. In this case, the extra pair of chromosomes should be regarded as fragments that are geni- cally essential to the survival and success of the genetic system of the plant species. By this frag- mentation, an otherwise somatic s of2n — 20 has become 2n = 22. This — 22 should, however, be seen as 2n = 20 + 2 ff. he evolution of this new number may be con- nected with the morphological differentiation and differential ecological preference that have led to ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 the recognition of U. a/tissima sensu stricto from its relatives U. gigantea and U. viridula Baker in the U. altissima complex dcus 19752). LITERATURE CITED BATTAGLIA, E. 19 Cytogenetics of 6-chromo- somes. Cytologia 17: 245-299, BLACKWOOD, E. 1 The inheritance of 8-chro- CARTER, C. R Brachychome lineariloba. 3. Accessory chromo- omes. Chromosoma (Berl.) 39: 361-379 Du C. D. & L. F. LA Cour. 1942. The Handling of Chromosomes. George Allen & Un- win, London. & A. P. WYLIE. 1955. Chromosome Atlas of Flowering Plants. George Allen & Unwin, Lon- don DE WET, J. M.J. 1957. Chromosome numbers in the Scilleae. Cytologia 22: 145-159. GILes, N. H. 54. Radiation- oua chromosome aberrations in Tradescantia. 713-762 in A. Hollaender (editor), cee Biology, Volume 1. McGraw Hill, New York Hair, J. B. 1952. The origin of new chromosomes in Agropyron. Heredity 6, Vn een on Chromosome Breakage]: 215-2 a 1967. i i TUE of the Liliaceae: . The karyotypes of twenty-five tropical mid ‘Kew Bull. 21: 31-38. LEA, D. E. . Actions of Radiations on Living Cells, 2nd edition. Cambridge Univ. Press, Cam- bridge ge. MARENAH, L. J. & J. H. W. HOLDEN. 1967. Karyotype studies in Avena: I. The karyotype of Avena i cultivar Condor. Chromosoma 22: 456-464 1960. Cytogenetic studies of Clarkia unguiculata: II. merary chromosomes. OYEWOLE, S. O. 1975a. Taxonomic treatment of the Urginea altissima (L.) Baker oo in West Af- rica. Bol. Soc. Brot. 49: 163-172. 1975b. Cytotaxonomic studies in the genus Urginea Stein in West Africa: k Karyotype anal- ysis in U. altissima Baker, U. gigantea ire Oyewole m U. viridula Biker (emend. ). Bo Brot. 49: 2 Rees, H. 1958. pes rential behavior of chromosomes a9 taneous chromosome breakage in Trillium : 367 Translocation heterozygotes a nd supern ary ch osom populations of Clarkia wil- pment Evolution 22: 93-107. Worrr, S. & H POLD. 1955. Metabolism and AE A E Science 122: 231-232. CYTOTAXONOMIC STUDIES IN THE GENUS URGINEA STEIN IN WEST AFRICA. III. THE CASE OF URGINEA INDICA (ROXB.) KUNTH IN NIGERIA! S. O. OYEWOLE? ABSTRACT Urginea indica (Roxb.) Kunth is a morphologically variable species of wide distribution in the Old World. In West Tropical Africa, it has The large form been considered to consist of a large form and a dwarf rm was investigated morphologically in as field and in form. the laboratory, as well as karyo- ue in the field. E differ in both vegetative wed differences in ecological an orp ere fo preferences. Biometrical analysis of vegetative and fora peer showed that the four variants relim hat they are similar but not identical in karyomorphology. Art inary karyotype analysis showed Sek crosses between them failed to suc viable seeds. It is concluded that U. indica is an incipient polyspecies or species complex. Urginea indica (Roxb.) Kunth is one of the four species of Urginea recognized in the latest revision of Liliaceae in Flora of West Tropical Africa (Hepper, 1968). It is distinguished from the other species by its globose capsule. It is wide- spread in the Old World tropics, inhabiting sa- vanna vegetation of tropical and subtropical areas of Africa, southern India and further east (Thi- selton-Dyer, 1898). In Nigeria, it is widespread in the central segment of the country, occurring between latitudes 7°N and 10°N. It occurs in open, heavy soil with a top layer of humus or in clay soil of seasonally flooded plains. It occurs in different ecological niches in var- (1968) observed this morphological variation and contended that there are at least two different forms of this variable species: a large form and a dwarf form. These two are easily distinguish- able in the field. The large form has light green leaves whose undersurface, immediately out of the bulb, is pinkish; the reproductive shoot is also pinkish, 45-150 cm tall, with not less than 15 flowers in the lax, racemose inflorescence. The tepal is also pinkish with a greenish keel. Leaves and flowers are never borne together. The dwarf form, on the other hand, has dark green leaves; the reproductive shoot is green, generally less than 40 cm tall, with flowers usually ranging be- tween one and 12; the tepals are yellowish green with a green keel. Leaves and flowers are never borne together. Field studies of this taxon during vegetative growth and flowering revealed that Hepper’s contention was a rather conservative estimate. The present paper therefore aims at establishing the taxonomic status of U. indica through morphological studies, starting with the large form MATERIALS AND METHODS Natural populations of the large form were studied morphologically during several field trips. Representative samples were collected and brought into cultivation in nurseries first at the University of Ibadan (southwestern Nigeria), then at Ahmadu Bello University, Zaria (northcentral Nigeria) and later at the University of Ilorin (westcentral Nigeria). Each asec site was visited at least twice—duri the vegetative growth period (May to Mic and during the flowering and fruiting period ewe to March). Altogether about 200 bulbs were brought into cultivation. Four morphological varias were identified from differen collection. Titled A-D, they were grown on ad- jacent nursery beds. No intermediate forms were encountered, even where the variants grew to- gether or in contiguous sites. Records of mor- phological features were kept and carefully fol- lowed in order to identify any environmentally induced features. These collections have been in cultivation since 1972/73. Each bulb is normally not more than 3—4 inches ! The financial support from the "pn edad à iS Ilorin Senate Research Grant No. 8.184.22, especially for the ed. final phase of this work, is gratefully acknow ? Department of Biological Sciences, oe of Ilorin, Ilorin, Nigeria. ANN. Missounmi Bor. GARD. 74: 131-136. 1987. 132 deep in the soil. Soil samples were usually taken along with each bulb, and both texture and com- position of the soil were determined later in the laboratory. The type of vegetation in which the population was found was also note M hological cl teristi divided into qualitative and quantitative features. The qual- itative features are bulb shape, shoot color, in- florescence color, leaf form, perianth color, ovary shape, ovary color, filament, style, and anther color. These were assessed visually. The quan- titative features are shoot and inflorescence height, number of flowers in the inflorescence, pedicel length, tepal length and width, length of ovary, style and stigma, anther, filament, and leaf, and leaf width. The leaf length-to-width ra- tio, called leaf ratio (l/w), was calculated for each meters or with millimeter graph paper. The shoot was measured from its base at bulb surface to its hile th ight was taken from the base of the first flower to the tip of the shoot. Both measurements were taken after the last (youngest and apical) flower had either opened or withered to ensure that the reproductive shoot had stopped elongating. Floral parts from newly opened flowers were dissected out for measure- ments. T 43 g£ + 1 CIV HIANY in order to investigate leaf cuticular surface pat- tern, leaf margin, number of veins, structure of veins, stomatal structure, and the general pattern of tissue distribution in the lea Chromosome number and karyomorphology were studied using root tip squashes. Root tips were harvested at about 8 A.M., pretreated for one hour in saturated aqueous solution of p-dich- lorobenzene, fixed in fresh 1:3 acetic alcohol (glacial acetic acid and absolute ethanol) and stored for at least 30 minutes at about — 4°C before hydrolysis. Hydrolyzed root tips were squashed in 2% acetic orcein. RESULTS The major collection areas are indicated in Figure 1. Except for Groups B and D, which were found growing together in a wide expanse of land, different groups were found in different popu- lations and in different ecological niches. All the populations encountered were found growin either in large numbers or as a few individuals dispersed in open savanna, with very light grass and forb cover and a few short, scattered trees ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 and shrubs. The soil was heavy clay with or with- d/or pebbles. The top layer was dark humus. All the populations were found in flood plains with little organic topsoil during the rains. In the dry season, the soil was hot, dry, hard and cakey. Figure 2 illustrates the vegetative morphology of the four groups, while Table 1 contains a sum- mary of all the morphological features investi- ated. Leaves are produced from the onset of the rains, and the plants remain vegetative for most of the rainy season. The leaves dry up towards the end of the rains. The early annual savanna fire of October to November burns the dry leaves, and reproductive shoots may be produced any time from two weeks after the burning. The ma- ture flower opens into a bell shape; the pedicel curves back to carry the flower face downwards at anthesis and until after pollination, after which the pedicel straightens up. The flower withers if not fertilized. When fertilization occurs, how- ever, the young fruit enlarges while the fading tepals close over it and shrivel into a little cap on top of the fruit by the time of maturation. Groups B and D are most similar, being dis- tinguished by shoot and inflorescence height, leaf length and form, and number of flowers per in- florescence only. D is distinguished from C by the lengths of tepal, style and stigma, filament and anther, filament color, bulb shape, shoot and inflorescence color, and ovary shape and color. B differs from A in shoot height and color, in- florescence height, lengths of pedicel, style and stigma, filament, anther, ovary shape, and fila- ment color. A and C are distinguished by shoot height, inflorescence height and color, number of flowers, lengths of pedicel, tepal, anther and leaf, leaf width, and ovary shape. A and D differ in shoot and inflorescence height, number of flowers, lengths of tepal, style and stigma, fila- ment and leaf, filament color, leaf form and width, ovary shape, and bulb shape. Figure 3 illustrates variations in leaf surface patterns, leaf margi out st tinctly different from the others in each of the features exhibited The leaf surface pattern is similar in all, show- ing minute crenation, which is most noticeable in variant D but less so in variants C, A, and B, respectively (see Fig. 3: A,, B,, C,, and D,). A and D have similar epidermal cell size, type, and arrangement as well as palisade cell size, form, 1987] OYEWOLE-— URGINEA INDICA 133 A p EET L L L RENT za " 4 7 8 9 10 t! 12 14. *13 139 M12 «Kano c E UNAUA . 1 » Zaria I! " ` p cya X DQuChi grs Kadtna `. "m T au i p. d OS a (FR. J a “Ge = ? š: ot 7t "6 6 a m ( «^ | 3 4 5 é 7 8 9 (0 m 12 13 i4 L L 2 2 A. A L 2 [B], 3C] & A.D. O denotes areas of major sampling of 1[A], routes during trips. FIGURE 1. and arrangement. They differ in leaf margin, being short and acute in A, the adaxial and abaxial epidermal layers being separated to the margin by the palisade, which aborts on a marginal epi- dermal cell, while leaf margin in D is projected with both epidermal layers coming together out- side the palisade and terminating with a marginal epidermal cell. They also differ in the distribu- tion of phloem tissue in the vein, being bicol- lateral in A but only collateral in D. In B and C, the epidermal cells are short and isodiametrical, the palisade cells are also short and less tightly Map of Nigeria showing areas of major sampling of the large form of U. indica. arranged, and the veins have large xylem vessels with conspicuous bicollateral phloem tissue. The leaf margin shows progressive elongation, being long and more acute in B than in A. In B, both epidermal layers do not close completely beyond the palisade before terminating in a marginal epi- dermal cell; in C, leaf margin is projected with rounded tip as in D and the epidermal layers barely close up beyond the palisade before ter- minating in a single marginal cell. Finally, the arrangement of the large metaxylem vessels is similar in A, B and D but differs in C. Undehisced anther color Pin Dirty white in Dirty white in Dirty cream 134 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 TABLE 1. Data on morphological features. Taxa Characters A B C D Bulb shape Small, edium, Medium, ovoid Large, spherical spherical spherical Shoot color reen Pink Light green Pink Inflorescence color Pink Pink Yellowish cream Pink Leaf form Short, Short, re- Long, coiled Long, straight coiled flexed Perianth color Pinkish Pink with Pinkish brown Pink with yellowish brown yellowish with green keel keel ith green keel green keel Ovary shape Pyramidal Globose Pyramidal Globose Ovary color reen Green Green Light green Filament color Pink Yellowish Pinkish Yellowish Style color ink i i Pink Creamy white Shoot height (cm) 45-60; 52.9 70-90; 76.9 100-140; 123.8 100-140; 119.9 Inflorescence height (cm) 15-20; 17.6 24—30; 27.0 50-60; 54.7 50-70; 59.5 Number of flowers 15-20; 17 15-20; 17 20-30; 24 0-30; 25 Pedicel length (mm) 22-23; 22.6 35-40; 37.1 30—40; 33.9 *(30-) 50-70; 60.3 Tepal length (mm) 12-14; 12.8 12-13; 12.6 15-16; 15.5 1-13; 12.1 Tepal width (mm) 44.5; 4.2 4-6; 5.2 3.5—4.5; 4.0 4.5; 4.5 Ovary length (mm) 4.5—5.5; 4.9 5-6; 5.3 4.5-5.0; 4.9 5-6; 5.5 Style + stigma (m 6.5-7.5; 7.0 5-6; 5.3 *(6.5—)7.0; 7.0 5-6; 5.5 Filament length (mm) 8.5-9.5; 9.0 5; 5.0 *(8.5—)9.0; 9.0 5-6; 5.6 Anther length (mm) 2; 2.0 2.5-3.5; 3.0 2.5: 2.5 3; 3. Leaf length (cm) 20-25; 22.7 25-35; 31.2 50-80; 67.3 50-70; 61.1 Leaf width (cm) 0.8-1.3; 1.0 0.8-1.3; 1.1 1.4-2.6; 2.1 1.0-2.4; 1.6 Leaf index (l/w) 17-31; 23.6 24-43; 29.8 22-52; 32.6 26-56; 39.5 * [ndicates infrequent deviating measurements. All four groups have a somatic chromosome number of 2n — 20. They have similar but not identical karyomorphology. An analysis of the karyotypes is the subject matter of a separate report (Oyewole, 1986). Artificial crossing be- tween the four groups failed to produce any hy- brid fruits. DISCUSSION Urginea indica has been described as a vari- able species (Hepper, 1968; Morton, 1961). Mor- phological variation was maintained even under uniform cultivation, suggesting that the variation d i with ecological preference rather than being ubiquitous in all populations, as would be ex- pected if the variations were due to polygenic effects (Dobzhansky, 1951; Huxley, 1942; Math- er, 1943). In U. indica, two distinct forms, B and D, were found together in the same population area without intermediates, indicating that the differences between them are not environmen- tally induced, while each group has a distinct karyotype (Oyewole, 1986). Hence variation in this case is not just a case of polymorphism. The correlation between the external morpho- logical variations and the leaf epidermal and me- sophyll features strongly supports the idea that this taxon is not just a single species. These an- atomical features are genetically controlled and, under the same environmental conditions, still maintain their differences. The importance of such anatomical features in species delimitation has been amply emphasized by Carlquist (1959) and Metcalfe (1963) and exhaustively demon- strated in many other works (for example, Prat, 1932; Church, 1949; Serensen, 1953; Borrill, 1959, 1961; Oyewole, 1971; Adeyemi, 1981). OYEWOLE— URGINEA INDICA FIGU ay e Vegetative morphology of the four groups (A-D) of the large form of U. indica. Horizontal ^s ar represents 4 m.—3. Leaf surface patterns: 1 — Leaf surface, Koi quede cell structure and the palisade layer; leaf margin; and 3— Leaf vein structure. Diagonal bar represents 25 um 136 Speciation, in the words of Dobzhansky (1951), is “that stage of evolutionary process at which the once actually or potentially interbreeding ar- ray of forms becomes segregated in two or more separate arrays which are physiologically inca- pable of interbreeding.” Recent views on specia- tion emphasize the relationship of the organism and the environment as the controlling factor (Hutchinson, 1959; Lewis, 1969). Thus adaptive radiations often occur when a species enters an unoccupied habitat with diverse open niches or when a population acquires a new complex of adaptive characters that enables it to exploit available environment more efficiently (Steb- bins, S as recorded for the Axonopus com- es Gledhill, 1966). Hence it is clear that U. indica, in which there are four morpho- logically distinct, genetically isolated forms even within the so-called large form, is not simply one phenotypically plastic genotype. It is significant that these forms exist side-by-side in nature or at least within the same geographical location and climatic condition while maintaining their identity both reproductively and morphologi- cally. Obviously their karyotypes resemble one another. However, they are biological entities. It is untenable to regard the hitherto U. indica as a single species (Lewis, 1969); rather it must be recognized as a species complex. The evolution- ary history of U. indica may possibly be similar to that of A/buca nigritana and the U. altissima complex in the same family (Gledhill & Oye- wole, 1972; Oyewole, 1975, respectively). LITERATURE CITED ADEYEMI, F. A. 1981. Biosystematic Studies of Some Nigerian Taxa of Anthericum Lin. and Chloro- phytum Ker-Gawl (Liliaceae). P D. Thesis. Uni- fais of Ibadan, RILL, M. 1959. A biosystematic study of some Glyceria species in Britain. I: Taxonomy. Wat- sonia 3: 291-298. 961. Epidermal characteristics in the dip- loid subspecies of D. glomerata. J. Linn. Soc. Bot. 8 56: 453-458. CARLQUIST, S. 1959. Anatomy and systematic posi- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 s of Centaurodendron and Yunquea (Compos- tae). Britonnia 10: 78-93. CHURCH, G. L. A D. study of Gly- ceria and Puccinellia. Amer. J. Bot. 36: 155-165. DoBzHANSKY, T. 1951. Genetics and the Origin of Species, 3rd edition. Columbia Univ. Press, New Yor GLEDHILL, D. 6. Cytotaxonomic revision of the A W) Beauv. complex. Bol. WOLE. "1972. The taxonomy of Albuca in West Africa. Bol. Soc. Brot. 46: 149- 170. HrPPER, F. N. 1968. Notes on West African mono- cotyledons. Kew Bull. 21: 493-498. HUTCHINSON, G. E. 1959. Homage to Santa Rosalia or why are there so many kinds of animals? Amer. Naturalist 93: 145-159. HuxLEY, J. 1942. Evolution: The Modern Synthesis. Harper and Brothers, p York. Lewis, H. 1969. Speciatio y and evolution. Taxon 18: 21-25. MATHER, K . 1943. Polygenic inheritance and natural selection. Biol. Rev. 18: 32-64. METCALFE, C. R. Comparative anatomy as a modern botanical discipline. Advances Bot. Res. 8. OYEWOLE, S. O. I Biosystematic Studies in the Genus A/buca L. with Particular Reference to Those Species Occurring in Nigeria. Ph.D. Thesis. Uni- un of Ibadan, Nigeria. axonomic treatment of the Urginea altissima (L. : Baker complex in Nigeria. Bol. Soc. rot. 49: -17 poe studies of the genus Africa. IV. Population dif- ype indica (Roxb.) Kunth. Ann. Missouri Bot. Garden 7-14 PRAT, H. 1932. piderme des Graminees: étude anatomique et EE P Ann. Sci. Nat. Bot. 14: 117-324. e T. 1953. revision of the Greenland es of Puccinellia. Parl. Medd. Gronl. 136(3): 46. Nn G. L., JR. 1971. Processes of Organic Evo- lution. Prentice Hall, Englewood Cliffs, New Jer- sey. THISELTON-Dy ER, W. T. (editor). 1898. Pp. 424—426 in Flora of Mrs Africa, Volume 8. L. Reeve & Co., London CYTOTAXONOMIC STUDIES IN THE GENUS URGINEA STEIN IN WEST AFRICA. IV. POPULATION DIFFERENTIATION AND KARYOTYPE VARIATION IN URGINEA INDICA (ROXB.) KUNTH! S. O. OYEWOLE? ABSTRACT Qualitative and quantitative studies of eS - over 250 individual plants of the variable Sedes Urginea indica (Roxb.) Kunth were carried o he + The plants were sampled from 23 collection ot tips were used for mitotic preparations. Ten otypes were ` recognized, four of which represented the first phenotype while the remaining Six ggre essive in its exploitation of various the demands of each ecological niche and a device to isolate the individual gene pools. Morphological variability is an undisputed at- tribute of a species composed of sexually repro- ducing individuals in a large panmictic popula- tion. However, the assumption that continuous populations were spatially fluid, panmictic, and genetically homogeneous has been assailed by the results of extensive works on both plants and animals (Epling & Dobzhansky, 1942; Selander et al., 1969; Bradshaw, 1972; Jones, 1973; Schaal, 1975). This assumption has now been largely replaced by - Anita that, particularly i in plants, many exten l semi- d ne This, according to Linhart et al. (1981), may be due to the effect of diver- sifying selection in heterogeneous environments and/or highly restricted gene flow as a result of spatial isolation. There is abundant evidence in support of each of these two phenomena in the process of speciation. Evidence of restricted gene flow has led to the assumption that, within con- tinuous populations, there exist small clusters of genetically related individuals (Bradshaw, 1972; Levin & Kerster, 1974). In spite of this, the ex- istence of distinct correlated discontinuities in the phenotypic characteristics of a continuous the existence of distinct segmentation in the ge- netic structure of the population (cf. Oyewole, 1971). Such segmentation can be maintained only by a number of factors, chief among which is an intrinsic isolation mechanism. The present work analyzes the results of stud- ies of the genetic structure of morphological vari- ants of the variable species or species complex, Urginea indica (Roxb.) Kunth. The morpholog- ical differentiation among the large form is the subject of the third in a series of studies of the genus (Oyewole, 1986). MATERIALS AND METHODS Populations were sampled in the wild, and plants were cultivated in experimental sites (Oyewole, 1986). Collection sites are illustrated in Figure 1. The distribution of the species, be- tween latitudes 7?N and 10°N, spans the decid- uous woodland and savanna of the Southern Guinea Savanna vegetation zone. Over 250 plants were collected from 23 sites in 17 sampling areas. Plants from distinct populations were grown to- gether under the same experimental conditions; seven distinct morphological groups were rec- ognized. Four of the forms (A, B, C and D) rep- resent the large form, while E, F and G belong to the dwarf form. Root tips were harvested between 8 and 9 A.M., pretreated for one hour in sat. aq. 1,4-dichlo- robenzene, fixed, and treated for mitotic squash preparations following the popio eth- ods (Darlington & LaCour, 1969). Chromosome counts were taken at random Malas every prep- aration. Chromosome measurements were taken from not less than 100 cells, at full mitotic meta- ! I gratefully acknowledge the technical assistance I received from the Department of Botany, University oo ss des Kenya, especially the patience with which Scolastica W. Karari, departmental secretary, typed th manu 2 esa m of Biological Sciences, University of Ilorin, Nigeria. ANN. MISSOURI Bor. GARD. 74: 137-143. 1987. 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 L ah. p a L. L L i À — ra 4 7 8 E) IO ft 12 (4. [13 151 4 -12 [| 4 EU (o - [0 « de be @ © 9 aOa tO 10; © N 1 s * 63 g : 1 7 A FG (^ ^i, e A2, AA), A AL, ° e í Zs t8, 4G XD, TE, PF oG) s. 3 4 Š G 7 8 9 (o II (2 13 (4. FicunE |. Map of Nigeria showing areas of major sampling sites. phase, in each morphological group. Homolo- kenir gous chromosomes were easily identified in each set of measurements from comparative chro- mosome lengths and the relative lengths of chro- mosome arms. Measurements were recorded in order of magnitude for the haploid set. Data were pooled for each group, from which average chro- mosome length, the relative chromosome-arm length, and the position of the centromere were determined. ower buds of appropriate age were collected between 7 and 10 A.M., immediately incised and fixed, and the anthers squashed and stained. As many inflorescences as were available in each group were sampled, and meiotic stages from pachynema to telophase II were examined in not less than 100 pollen mother cells (PMCs) in those groups that flowered (not all the groups have flowered in cultivation). Figure 2 contains somatic metaphase comple- ments of the various groups. All the groups have 2n = 20. Karyotype data is summarized in Table 1. Each morphological group is represented by a different karyotype. The total length of chro- matin material, at metaphase, of each karyotype differs from the others. The karyomorphology is similar, although intrinsic differences abound (Fig. 3). Chromosome classification, using the chromosome index (ratio of long arm to short arm), is according to Levan et al. (1964). The chromosomes in each complement were classi- fied as long (6.0 um and above), medium (4.0- 5.9 um) and short (below 4.0 um). Details of the meiotic study will be presented in a subsequent part of this series. Group A. This group is represented by four OYEWOLE— URGINEA INDICA KARYOTYPE VARIATION 139 1987] "uerpouiqns = uis 'uerpoul = W EUW qns = 1S FEUI = 1 989 089 €L ct ç CHL TES tet 9ct c8 ç L6'S 9U1IOSOUI -oJqo/u12uo[ unguroiuo ULIN 8c LEI L6'S€I cç CvI 8t 80I pr srl 9€'901 9Lv8 9c' 68 9€ 9TT t€ 6II 1uouro[d -WI09/Y13UZ] urjguroJuo [£10], 1s 1s 1s 1s 1 1s 1s ws 1s 1s uongo] 219UIOU2/) t6€ 69 Lov 09 SL8 os OV 9c I6 Sv 9n[eA 4 PIP Ort Scv ce 88v 0't $c $c Le t L9'€ q13uo| ungtuoiuO OI 1s 1s 1s 1s 1 1s 1s UIS 1s 1s UOT]?20[ 2J9UIOJ]U2/) 6Ç ç 9'9 os Ov CL PI's os $t 9¢°9 OL on[eA 4 cry Sev St SLE EIS $t 0't SLT 89't Ot q13uo| ungtuo:uO 6 1s 1s 1 1s 1s 1 1s 1s 1s 1s Uone] 219Uu101]U2/) £9'e os GE 88v Os PLL os ct vr os 3n[eA 4 IL SIs ETS c6t STS L9'€ 0t Oe 90°F o't q13uo[ unetuoiuO 8 1s 1s 1s 1s 1s 1s ul 1s 1s 1s uone] 23J9UIOJI]U2.) 9t'6 £'9 09 88v 67'S [A Ol 9't IC € OL 9n[gA 4 96v gÇ ç çC ç COE ES 88't 0t ' 0t ley 0t q13uo| urjguro1g;) L 1s 1s 1s 1 1 1S 1s 1s 1 1 UOT]E2O[ 919u101U9O 6S6 0'9 09 L0°8 88L O'S 079 079 S78 0'8 onyea 4 v's tps Secs LIv c6 `ç Sv $t $t £9 v SP yisus] ur]guro1q;) 9 1s 1s 1s 1s 1s 1 1s 1s 1s 1 UOT]E2O| 219UIOJ]U2;) vs $9 $9 os 98'$ €8 OL 09 9'9 06 9n[gA 4 69'6 $9'6 £9°¢ St 09 SLY OV S'E SLY os q13uo| urjguro1q;) ç 1 1 1s 1 1 1s 1s 1s 1s 1s UOT]E2O[ 3139UIOJ])U2/) PLL LOL 09 06 PIL L9'9 9'ç ces Sv 079 onyea 4 c8'9 8L OL 0's £r CLS Ot SLY ç ç ç ç q13uo| ur]guro1q;) v 1 1 1 1 1 1 1 1 1s 1 UOT]£29[ Ə19u10nUu2O 601 $8L £€ IT 9L 9978 CL 076 $€L8 OL L9'6 NLA 4 I8'8 £9'8 $c 6 OTL 9$'6 £19 os 88v SL 0'8 yisua] urguo1g;) € 1 1 1 1 1 1 1 1 1 1 uongoo[ 919u1011u9O 9'cI 8'8 $'OI CL SL Ol € Ol CLI L9'8 88'8 076 9n[eA 4 £€ IT St'6 SII $8 cL Il $8 88'9 STL 88'6 (OKORI q13uo| uyewoIyo [4 1 1 1 1 1 1 1s 1s 1 1 UOT]£20| Ə919u101lU9O sc SOTI oll L9'6 Sc TI $8 $9 $9 906 001 onyeAa 4 £€'cI SO ZI Sel c8'6 Vel $6 SL SL 9€'01 OTT yisus] urietuoiuO I 5 i d a Ə d "V "V ‘Vv 'V son3o| -OWO0H '(voipui »2u184/]) erep sdAjOAIeyY jo Areurumg "| a18v 1 Ar > P Ae : MI : * tt *) Y B ~ ` ` N, a < ` i y mo "m a. x 3 . N ^ À — © * - M " v^ ` ux F N 5 P y , ` i e 3 — ee: 3 š i a NC t B b " Q O 7 etaphase co ple - O pe 3 ^. d O O ° 2 d are d O g one C š and A, O g = = g able pe A eprese : O Ü JUO of 2 400 epa ° 1987] in one sampling site about 80 km to the east of A E A, karyotype consists of chromosomes whose average lengths vary between 3.67 um and 11.0 um, with an average total chromatin length of 119.34 um per complement. The complement consists of three long, six medium, and one short pairs of chromosomes, all with terminal and sub- terminal centromeres. Endomitosis frequently Occurs in the root cells. Meiosis is normal, and 10 bivalents are regularly formed. Even though each complement of A,, A, and A, could be resolved into pairs of similar chro- mosomes, members of such pairs are by no means identical. None of the individuals of these three karyotypes has flowered since they were brought into cultivation; hence these hypothetical pair- ings could not be verified in actual meiotic pair- ing. A, consists of chromosomes whose average lengths range from 3.31 um to 10.56 um, with an average chromatin length of 116.36 um per complement. The complement consists of six long, 10 medium, and four short chromosomes, all with subterminal-terminal centromeres. The longest two of the chromosomes have a centric region as wide as the short chromosome arm length. A, chromosomes range in average length from 2.5 um to 7.5 um, with an average chromatin length of 85.26 um per somatic complement. There are five long chromosomes in the com- plement, four of which resolve into two pairs while the fifth is associable with a shorter chro- mosome. This long chromosome has a conspic- uous secondary constriction (arrowed in Fig. 2, A4). The whole complement consists of five long, four medium, and 11 short chromosomes, all of which have their centromeres in the subtermin- al-terminal region, except the smallest two pairs which have submedian centromeres. A, chromosomes vary in average length from 2.5 um to 7.5 um. They have an average chro- matin length of 84.76 um per somatic comple- ment. The complement consists of four long, six medium, and 10 short chromosomes, all with subterminal-terminal centromeres except a short pair with median centromere. Group B. This is represented by only one karyotype. Chromosomes vary in average length from 3.0 um to 9.5 um, with an average chro- matin length of 106.36 um per somatic comple- ment. The complement consists of three long, three medium, and four short pairs. All the chro- OYEWOLE— URGINEA INDICA KARYOTYPE VARIATION Ailit anto Ay È Jinna A2 GUA by than sees uns A3 EITAYTTITTTTIN A4 [ Diu Hillia B Whun C inus Mini D T Ilii lilii ua E MMM F HTTP G URE 3. Idiograms ofthe various karyotypes rep- Eu by the groups. Horizontal bar represents 30 mosomes have subterminal-terminal centro- meres. The longest pair has a wide centric region. Meiosis is normal except for the infrequent early separation of members of a small pair Group C. Chromosome length varies from an average of 4.88 um to 13.1 um, with an av- erage chromatin length of 148.44 um per somatic complement. There are five long and five me- dium pairs of chromosomes in the complement. All the chromosomes have subterminal-termi- nal centromeres. The longest pair has a centric region as wide as its short arm. Multivalents are frequently formed during meiosis. Group D. Populations of this group are sym- patric with those of Group B. The karyotype con- sists of chromosomes with subterminal-terminal centromeres. The average chromosome length varies from 3.5 um to 9.82 um, with an average chromatin length of 108.48 um per somatic com- plement. There are three long, three medium, and four short pairs of chromosomes. The two 142 longest pairs have a wide centric region. Meiosis is regular. roup E. The karyotype of this group con- sists of chromosomes with subterminal-terminal centromeres. Chromosomes vary in length from 4.25 um to 13.5 um and have an average chro- matin length of 142.52 um per somatic comple- ment. There are four long and six medium pairs of chromosomes. The first and third long pairs have a wide centric region. Meiosis is regular. Group F. The karyotype consists of chro- mosomes that vary in length from 4.1 um to 12.05 um, with an average chromatin length of 135.97 um per somatic complement. The com- plement is made up of four long and six medium pairs, all with subterminal-terminal centro- meres. The fifth pair has a secondary constriction on the long arm. Meiosis is regular. Group G. The karyotype consists of chro- mosomes whose average lengths vary between 4.14 um and 12.33 um, with an average chro- matin length of 137.28 um per somatic comple- ment. The complement consists of four long and six medium pairs, all with subterminal-terminal centromeres. Meiosis is regular. DISCUSSION The similarity in the morphology ofthe karyo- types is obvious. The differences in the karyo- morphology of the different populations seem minute, but they are basic and do underlie the differences in the external morphology of each retained. It is, however, evident that changes have occurred (or are occurring) in this taxon that ma be correlated with the morphological differentia- tion of the populations. The recognition of dif- ferent karyotypes that correspond to different ; the presence of more than one karyotype in an otherwise mor- phologically uniform unit raises interesting ques- tions about evolutionary phenomena. This mor- phologically uniform unit, Group A, is interesting. $ ete see Kobe Ku fate, panied by external morphological differentia- tion, but it is partly correlated by habitat pref- the A,, A, and A, complements clearly plements represent natural hybrid swarms that ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 introgressively identify with A,, which is likely to be one of the putative parents. The different karyotypes are correlated with differences in ex- ternal morphology and ecological preferences. Those forms that inhabit different ecological niches may have differentiated in response to differences in ecological demands, while those which inhabit same or similar ecologic niches must be fundamentally different genetically in order to retain their individual identities mor- phologically. In both cases, karyotype differen- tiation seems to have resulted in reproductive isolation by which the different forms are main- tained in nature he differences in the amount of chromatin material may have resulted from (or led to) karyotype differentiation. Difference in the chro- matin material is correlated with both morpho- logical differentiation and ecological preferences (note A, and A,, B and D are in the same niches; A, and A,, and E, F, and G are in similar niches; and C and E are in different niches). The morphological variability of this species has long been recognized, but no prior attempts have been made to distinguish the different forms beyond the arbitrary categorization of “large” and "dwarf." This is probably due to the fact that no field collection ever contains both floral and vegetative features together, as well as to the wide east-west distribution of the species. The aggressive exploitation of different niches by dif- ferent biotypes has resulted in the differentiation of the morphological forms within the species' broad areas of distribution. The isolation of spe- cific biotypes, forming small clusters of individ- uals, in such ecological niches probably led to the accumulation of favored genes and/or mod- est chromosomal changes and eventually to the specific karyotypes that are associated with spe- cific morphological forms as well as with specific ecologic niches (Wright, 1940; Bush et al., 1977; Bengtsson, 1980). There is no doubt that mor- phological differentiation in these populations is more obvious than differentiation in chromo- some morphology, suggesting that chromosome repatterning may have been mild and might have involved only small segments in gene/gene block rearrangement. Hence this species seems to com- prise a stable polymorphism in which the differ- ent forms have attained reproductive isolation and genetic stability, and hence each form has retained its morphological identity. This case is therefore different from that of Agrostis tenuis 1987] (Bradshaw, 1959) or Elymus rechingeri (Heneen & Runemark, 1962) but seems to be similar to that of the diploid neospecies of Clarkia (Lewis, 1973) Flower formation and fruit development are a common feature with most of the morphological forms, especially in nature. Preliminary studies of meiotic behavior have shown regularity of pol- len formation in most of them under cultivation. However, this apparent sexual reproduction is coupled with vigorous vegetative propagation by axillary bulb formation in varying degrees in all forms. It is therefore necessary to have a closer look at the reproductive biology of the entire species in order to ascertain the extent of actual sexual reproduction and the mechanism of pol- lination in each form. Hybridization experi- ments between the different forms, which will hopefully shed light on their genetic divergence, are in progress. Thus far, preliminary results of such experiments show successful n cross- ing between only two forms, A, an LITERATURE CITED BENGTSSON, B. O. 1980. Rates of karyotype evolution in placental saci Hereditas 92: 37-47. 1959. Population differentiation cal differ- HAW, A. in Agrostis pns Sibth. I. Morphologi entiation. New Phytol. 58: 208-227. 1972. Some evolutionary consequences of being a f Evol. Biol. 5: 25-47. BusH, G. L., S. M. Case, A. C. WILSON & J. L. PATTON. 1977. Rapid speciation and chromosomal evo- lution in mammals. Proc. Nat. Acad. Sci. 74: 3942- BRADS OYEWOLE— URGINEA INDICA KARYOTYPE VARIATION 143 ar e C. D. & L. F. LACougR. 1969. The Han- ing of Chromosomes, 6th edition. George Allen Lon .& T. DOBZHANSKY. 1942. Genetics of nat- ural populations. VI. Microgeographical races in Lynanthus parryae. Genetics 27: 317-331. HENEEN, W. K. & H. RUNEMARK. 1962. Chromo- some polymorphism and morphological diversity n Elymus pie fa u Sedis 48: 545-564. dones J. S. Ecological genetics and natural selection in molluscs. Science 182: 546-551. LEVAN, A., K. FREDGAR & A. A. SANDBERG. 1964. Nomenclature for centromeric position on chro- mosomes. Hereditas 52: 201-220. Levin, D. A. AND H. W. KERSTER. 1974. Gene flow in seed plants. Evol. Biol. 7: 179-220. Lewis, H. 1973. The origin of diploid neospecies in Clarkia. Amer. Naturalist 107: 161— i LINHART, Y. B., J. B. MITTEN, K. B. STURGEON & M. L. Davis. 1981. Genetic ara in space and time " a population of ponderosa pine. Heredity 46: 407—426. OYEWOLE, E O. Biosystematic Studies in the Genus A/buca L. with Particular Reference to Those Species Occurring in Ix . Ph.D. Thesis. Uni- versity of Ibadan, Nig 86. Cytotaxonomie studies in the genus Ur, rein ea Stein in West Africa. III. The case of Ur- on pe Ra fi Kunth i in n Nigeria. Ann. Mis- -136. ran mi E e p s structure and local dif- ferentiation Ç: Liatris cylindracea. Amer. Natu- ralist 109: 511-528. SELANDER, R. K : Y. YANG & W. G. Hunt. 1969. olymorphism in esterases and hemoglobins in wild populations of the house mouse. Studies in Genetics 5: 271—328. WRIGHT, S. 1940. Breeding structure of populations in relation to speciation. Amer. Naturalist 74: 232- 248. NEW TAXA OF OENOTHERA L. SECT. OENOTHERA (ONAGRACEAE)! WERNER DIETRICH? AND WARREN L. WAGNER? ABSTRACT everal new ionis are described in advance of a complete monograph of Oenothera sect. Oenothera imannia. u the plains to eastern Colorado and Tex unz to O. laciniata subsp. p outcrossing species, O. are differentiated from the oo ae fees O. pubescens, for a total of three species assigned to subsect. Nutantigi Subsection Raimannia is n divided ge two series: series Candela, established for the species with densely flowered spikes bearing two ers opening per day, inflorescences without ore flow lateral branches, acute to rounded petals, and straight floral tubes (O. rhombipetala, O. heterophylla, O. clelandii, O. bifrons, and O. curtissii); and to ema rgin exicana, O. laciniata described for relictual populations i in southeastern series inflorescences usually interrupted by lateral branches, usually only o nate petals, and floral tubes curved upward , O. drummondii, and O. humifusa). The new iiec Oenothera flare is aimannia is restricted to pla D with loose ne flower ope ng pe prior to anthesis (O. grandis, O. fa lfurr. n bivalents o me differentiated from its presumed S O. grandis and pem the morphologically similar and dein atric O. laciniata. The ew combinat mondii su thalassaphila is made for the populations in Baja California oie set as O. G ee Over 20 years of cultivation of nearly 150 strains for experimental hybridizations, cytolog- ical examination, study of breeding systems, and extensive study of herbarium materials, has led to revision of the species referred to subg. Rai- mannia (Rose ex Britton & A. Brown) Munz by Munz s 1965). Extensive analysis of cross- as resulted in realignment ofa group RS 75 species into sect. Oenothera, which in- cludes the species referred by Munz to his subg. Oenothera and subg. Raimannia (Stubbe & Ra- ven, 1979). In turn, this large group of basically intercrossable species was further subdivided into ve smaller crossing groups that form highly fer- tile hybrids and usually have compatible plas- tids. The first of these groups comprises the South American species, which Dietrich (1977) placed into subsect. Munzia W. Dietrich (45 species). The second, subsect. Oenothera (13 spp.), was outlined by Raven et al. (1979). The third and p is subsect. Emersonia n & W. L. Wagner (four spp.), revised in 1985 by Dietrich et al. The fourth and fifth groups, subsect. Raimannia and the new subsect. Nutantigemma described in this paper, will be the subject of an upcoming detailed revision. Prior to publication of the revision, the present paper makes the new subsection and species available for regional floras and concur- rent work on Oenothera, including DNA restric- tion mapping and studies of flavonoids, pollen, and seed anatomy. SUBSECTION NUTANTIGEMMA Populations occurring in montane sites from the western United States south to South Amer- ica, referred by Munz (1935, 1965) to a variety laciniata. This widespread entity is a permanent translocation heterozygote treated by us as O. pubescens Willd. ex Spreng. Two new Mexican species, closely related to Oenothera pubescens, one from the cape region of Baja California and one in the southern Sierra Madre Occidental, were detected during the study of herbarium specimens and cultivation of nu- merous strains in the experimental gardens (Stubbe & Raven, 1979). They are described here as O. breedlovei (bivalent-forming) and O. tam- ! This work has been supported by a series of grants from the National Science Foundation to Peter H. Raven. ? Botanisches Institut der apis META Universitatsstr. 1, D-4000 Düsseldorf 1, West Germany. ? Bernice P. Bishop Museum, P.O. B ANN. MissouRi Bor. GARD. 74: 144-150. 1987. 19000-A, Honolulu, Hawaii 96817, U.S.A. 1987] rae (presumably bivalent-forming). Oenothera tamrae is known only from the type collection and has not been grown in the experimental gar- den. All three species are placed into the new subsect. Nutantigemma. Oenothera L. sect. Oenothera subsect. Nutanti- gemma W. Dietrich & W. L. Wagner, sub- sect. nov. TYPE: O. pubescens Willd. ex Spreng. Oenothera sect. ribs ia sensu Ser. ex DC., Prodr. 4: 46. 1828, pro part a co sensu Wooton & pese Contr. U.S. Natl. erb. 16: 150. 1913, M br sensu Rose, 2 ‘Natl . Herb. 8: 330. 1905, pro parte; sensu T Sprague $ Riley, Bull. Misc. Inform. 200. 1921, a d fa mania an u (Rose ex Britton & A. wn) Munz, Amer. J. Bot. 22: 645. 1935, pro r. nios sect. Anogra sensu Tidestrom, Fl. Ariz. & x. 273. 1941, pro part Oenothera Sue Raimannia sect. Raimannia sensu ose ex Britton & A. Brown) Munz, N. Amer I. 5: 105. 1965, pro Oenothera sect. Oenothera subsect. Raimannia sensu (Rose ex Britton & A. Brown) W. Dietrich, Ann. Missouri Bot. Gard. 64: 612. 1977 [1978], pro parte. Her haps annuae vel probaliter biennes, erectae, ro- sulatae, simplices vel caules principales ramosi ramis We areas rosula adscendentibus. Folia rosulae anguste eb'anceolata, profunde partita vel n integra, acuta; el elliptica, a nguste oblanceolata vel oblanceolata vel lanceolata vel an- acut folia strigillosa vel villosa, raro glanduloso- pubescen- mplices vel in tiaca post anthesin. Styli 276.5 cm longi, stigmate sub th : Py e 1 + 1 th M4 ed x. — Capsulae cylindricae, 1.8-4.6 cm longae. Semina am- bito elliptica vel rotundata, brunnea vel atrobrunnea, saepe atromaculata, 0. .9— : x mm longa, 0. 6- 1 mm cras- sa. Planta | el struc- turaliter UE LIEN heterozygotica complexa. Numerus gameticus chromosomaticus, n = 7 The principal reason for according these three species (O. pubescens-group of Stubbe & Raven, 1979) subsectional status is the sterility of hy- DIETRICH & WAGNER—NEW TAXA OF OENOTHERA 145 brids in crosses between subsections Raimannia and Nutantigemma. They range from western Texas west to Arizona and southeastern Cali- fornia, south to Mexico and Guatemala, and one species extends to the Andes of Colombia, Ec- uador, and Peru south to the province of Junin. The nodding flower buds on the species of sub- sect. Nutantigemma clearly differentiate them from those of subsect. Munzia and subsect. Rai- mannia. This feature is shared with the white- flowered sect. Kleinia Munz, sect. Anogra (Spach) Endl., sect. Ravenia W. L. Wagner, occasionally with Oenothera caespitosa Nutt. subsp. nava- Jjoensis W. L. Wagner, Stockhouse & Klein [sect. Pachylophus (Spach) Endl.], and with the yellow- flowered sect. Eremia W. L. Wagner. In Oeno- thera fruticosa L. [sect. Kneiffia (Spach) Endl.], which has yellow flowers, the shoot apices, but not the flower buds, are bent downward; a similar condition occurs in O. speciosa Nutt. [sect. Xy- lopleurum (Spach) Endl.]. Nodding buds appear to represent, at least for the most part, plesio- morphy rather than convergence. Another important argument for considering this group a subsection is that these three species are distributed completely allopatrically from subsect. Raimannia. The species of series Nut- antigemma grow exclusively in montane habi- tats, from 3,900 m down to approximately 1,500 m, while the species of subsect. Raimannia grow only at lower elevations. Oenothera breedlovei is self-compatible and bivalent-forming; O. pubes- cens is a permanent translocation heterozygote forming a ring of 14 chromosomes in meiosis I and with 40-70% pollen fertility; O. tamrae is probably an outcrossing bivalent-former, based on 90-1009 fertile pollen in the type collection. KEY TO THE SPECIES OF SUBSECT. NUTANTIGEMMA la. Stigma elevated above the anthers at anthe- sis; pollen ca. 90-100% fertile; Laguna Mts., ide California, Mexico, and Nayarit, Mex- = Lower leaves deeply parted almost to the midrib; mature buds up to 5 mm in di- ameter: Laguna Mts., Baja California, Mexico O. breedlovei 2b. Lower leaves not parted to the midrib; mature buds rad mm in diameter; Na- yarit, Mexic O. tamrae 1b. Stigma surroun hace I" the EE see ca. 40-70% fertile; Arizona, Texas, co (ex- E s California), [i ti Colombe o Per O. pubescens 146 Oenothera breedlovei W. Dietrich & W. L. Wag- ner, sp. nov. TYPE: Mexico. Baja California: glong Interior val- jer: [La Laguna], south of Pico La Aguja, Sierra La Laguna, 6,300-6,700 ft., 22 Oct. 1977, D. E. Breedlove 43362 (holotype, MO- 2695034). Herbae annuae vel biennes, erectae, rosulatae, 2-3 dm altae. Folia profunde partita vel remote vadoseque dentata. Gemmae vetustiores nutantes. Tubi florales 2.4—4 cm longi. Sepala 1.2-2.2 cm longa, saepe rubro- egli villosa et glanduloso-pubescentia; apices se- paloru mm longi. Petala 1.6-3.5 cm longa, flava. Cou: 1.8-4.6 cm longae, E s Samina 1.2-1.6 mm longa, 0.6—0.7 mm crassa, saepe Mm Numerus gameticus chromosomaticus, n = 7; planta chromosomatice structuraliter homozygotica, npa compatibilis. Erect annual or probably biennial herbs, form- ing rosettes; stems 2-3(-5) dm long, simple or with a branched main stem and arcuating lateral branches arising from the rosette, usually flushed with red, densely strigillose, sometimes also scat- tered villous. Rosette leaves narrowly oblanceo- late, 4-12 cm long, 0.5-2 cm wide, pinnately o remotely and shallowly dentate, the apex acute, gradually narrowed to the petiole; cauline leaves very narrowly elliptic to lanceo- late, 2-5 cm long, 0.5-2 cm wide, usually pin- nately parted, the apex acute, the base narrowly cuneate, short-petiolate to sessile; bracts lanceo- late to narrowly ovate, 1.5-3 cm long, 0.5-1.5 cm wide, deeply parted to remotely shallowly dentate, the apex acute, the base broadly cuneate to narrowly cuneate; leaves and bracts densely strigillose and sparsely villous. Inflorescence usu- ally with lateral branches, the young buds erect, the older ones nodding, becoming erect just be- fore opening. Flowers usually | per spike opening near sunset each day. Floral tube flushed with red, 2.4—4 cm long, ca. 1 mm diam., sparsely to densely strigillose, villous, and glandular puber- ulent. Mature buds oblong, 3-5 mm diam. at the base. Sepals yellowish, often flushed with red, also red maculate and striped at the margins, scattered to densely villous and scattered to densely glandular puberulent, 1.2-2.2 cm long, the free tips 0.5-1 mm long, erect and appressed in bud, strigillose to villous. Petals yellow, broad- ly obovate to very broadly obovate, retuse, 1.6— 3.5 cm long, 1.6-3.7 cm wide. Filaments 1.1-2 cm long; anthers 4-12 mm long; pollen ca. 90- 100% fertile. Ovary 1-2.5 cm long, ca. 1.5 mm diam., densely strigillose; style 4—6.5 cm long, the visible part 1.7-2.5 cm long; stigma elevated ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 above the anthers at anthesis, the lobes 3-6 mm long. Capsule cylindrical, 1.8—4.6 cm long, 3-3.5 mm diam., densely strigillose. Seeds ellipsoid to broadly ellipsoid, brown to dark brown, often with darker flecks, 1.2-1.6 mm long, 0.6-0.7 mm diam., the surface pitted. — FD but modally outcrossing. Gametic chromosome number, n = 7 (7,, at meiotic metaphase I). This new species, known only from Laguna Mts., southern Baja California, Mexico, is named in honor of Dennis E. Breedlove (California Academy of Sciences), who has added greatly to our knowledge of Mexico through his extensive and excellent collections and to whom we are indebted for collecting material of this and many other Mexican Oenothera species for cultivation at the Botanical Institute of the University of Diisseldorf and at the Missouri Botanical Gar- Oenothera tamrae W. Dietrich & W. L. Wagner, sp. nov. TYPE: Mexico. Nayarit: Sierra Madre, near Santa Teresa, territory of Tepic, 8 Aug. 1897, J. N. Rose 2133 (holotype, US-301038; isotypes, NY, UC). Herbae annuae vel biennes, erectae, rosulatae, 2-4 dm altae. Folia partita seu remote obtuseque dentata vel quasi integra. Gemmae vetustiores nutantes. Tubi florales 3.5-4.2 cm longi, villosi et glandoloso-pubes- centes. Sepala 1.8-2.5 cm longa, rubro complana et rubro-fasciata ad margines; apices sepalorum 0. longi. Petala 2-3.5 cm longa, flava. Capsulae 4—4.5 cm longae, strigillosae et villosae. Semina 1-1.1 mm longa, 7 mm crassa Erect annual or biennial herbs, probably form- ing rosettes; stems 2-4 dm long, simple or with obliquely ascending lateral branches arising from the rosette, densely strigillose and sparsely to densely villous. Cauline leaves narrowly elliptic or narrowly lanceolate to lanceolate, 4-8 cm long, 0.8-1.8 cm wide, pinnately parted or remotely and bluntly dentate to subentire, the apex acute, the base narrowly rowly lanceolate to lanceolate, 0.7-1.5 cm wide, remotely and bluntly dentate, the apex acute, the base narrowly cuneate, sessile; leaves and bracts strigillose. Inflorescence simple or with lateral branches, nodding. Usually 1 flow- er per spike opening probably near sunset each day. Floral tube flushed with red, 3.5-4.2 cm long, 1.5-2 mm diam., sparsely villous and sparsely glandular puberulent. Mature buds cy- lindrical to narrowly ovoid, 5-7 mm diam. at the base, nodding before anthesis. Sepals yellow- T > v ° 1987] ish, often flushed with red and striped red at the margins, the pubescence as on floral tube, 1.8- 2.5 cm long, the sepal tips ca. 0.5 mm long, erect in bud, strigillose. Petals yellow, very broadly obovate, retuse, 2-3.5 cm long, 3-4 cm wide. Filaments 1.5-1.7 cm long; anthers 6-9 mm long; pollen ca. 90-10096 fertile. Ovary 1.8-2.6 cm ong, ca. 2 mm diam., densely strigillose and densely villous, the apex also glandular puber- ulent. Style 5.3-6.4 cm long, the visible part 1.8— 2.2 cm long; stigma elevated above the anthers at anthesis, the lobes 5-8 mm long. Capsule cy- lindrical, 4—4.5 cm long, 3-4 mm diam., the pu- bescence as on ovary but less dense. Seeds broad- ly ellipsoid, brown with dark red flecks, 1-1.1 mm long, ca. 0.7 mm diam., pitted. Chromo- some number unknown. This rare new species, known only from the type locality, near Santa Teresa in the Sierra Madra, Nayarit, Mexico, is named in honor of Tamra Engelhorn Raven, botanist and wife of Peter H. Raven. The description is based entirely on the type collection made by J. N. Rose i 1897. SUBSECTION RAIMANNIA The yellow-flowered species assigned by Munz (1935, 1965) to his subg. Raimannia from the central and eastern United States are now con- sidered to comprise subsect. Raimannia. Stubbe and Raven (1979), when considering the realign- ment of sect. Oenothera, included O. grandis (Britton) Smyth, O. laciniata Hill, O. drummon- dii Hook., O. humifusa Nutt., O. heterophylla Spach, and O. rhombipetala Nutt. ex Torrey & A. Gray in subsect. Raimannia. Work toward the overall revision of the subsection has led to the recognition of two series within it. Oenothera sect. Oenothera subsect. Raimannia ose ex Britton & A. Brown) W. Dietrich series Candela W. Dietrich & W. L. Wagner, series nov. TYPE: O. rhombipetala Nutt. ex Torrey & A. Gray. d sect. III uha. Nouv. Ann. Mus. Hist. Nat. 347. 1835, pro p Oenothea 8E ecco ias sensu S. d Proc. Amer. . Arts 8: 574. 1873, pro PT Ren sensu r Bull. Teer Bot. ‘Club 23: 172, 1896, pro part Raimannia sensu Rose, Contr. U.S. Natl. Herb. 8: 330. 1905, pro parte. Oenothera subg. Raimannia sensu Munz, Amer. J. Bot. DIETRICH & WAGNER—NEW TAXA OF OENOTHERA 147 22: 645. 1935, pro parte; sensu iig & Riley, Bull. Misc. Inform. 200. 1921, prop Oenothera subg. Raimannia sect. RA sensu Munz, N. Amer. FI. II. 5: 105. 1965, pro parte. erbae annuae vel biennes, probaliter breviter pe- ae apu e rosula Kaa e ua, ramis later iem g ae elliptica vel late elliptica, 1.1-1.9 mm longa, 0 mm crassa, plerumque atromaculata. Series Candela comprises a clearly defined group of five closely related species that occur in sandy soil in open places such as fields, prairies, roadsides, and open woods, from southern South Dakota, Minnesota, and Michigan south to Tex- as and southeastern New Mexico and east to Louisiana, Alabama, northern Florida, and outhern Georgia. They all have the presumably derived characters of relatively densely flowered spikes on which two or more flowers open every evening, unlike the species of series Raimannia, which nearly always produce only one flower per branch each day, a plesiomorphic characteristic. The spikes of series Candela never have lateral shoots, as 1s often the case in series Raimannia. The fully grown buds of series Candela are nar- rowly oblong, whereas those of series Raimannia are lanceoloid in the large-flowered structural homozygote species and oblong to ellipsoid in the small-flowered heterozygote species. In series C t. also clearly differentiate series Candela from se- ries Raimannia; those of series Candela are acute to rounded at the apex, in contrast with those of series Raimannia which are truncate to emar- ginate. Further, the capsules of series Candela are relatively short and thicker toward the base, while those of series Raimannia are on average longer and nearly cylindrical. All species of series Candela appear to be biennials. In contrast to this specialized habit, the species of series Rai- mannia have evolved an annual habit except O. drummondii and O. humifusa, which inhabit sand dunes and have retained the generalized peren- nial habit. Three of the species, Oenothera heterophylla, O. bifrons D. Don, and O. f- incompatible bivalent-formers, and O. W. Dietrich, Raven & W. L. Wagner and O. 148 curtissii (Rose) Small are complex structural het- erozygote species, presumably derived from an ancestor similar to O. rhombipetala. Oenothera sect. Oenothera subsect. Raimannia (Rose ex Britton & A. Brown) W. Dietrich series Raimannia (Rose ex Britton & A. Brown) W. Dietrich & W. L. Wagner, comb. & stat. nov. Based on Raimannia Rose, Contr. U.S. Natl. Herb. 8: 330. 1905, ex Britton & A. Brown, Ill. Fl. N. U.S., 2nd edition, 2: 596. 1913. Oenothera subg. Rai- mannia (Rose ex Britton & A. Brown) Munz, Amer. J. Bot. 22: 645. 1935. Oenothera subg. Raimannia sect. Raimannia (Rose ex Brit- ton & A. Brown) Munz, N. Amer. FI. II. 5: 105. 1965. Oenothera sect. Oenothera sub- Gard. 64: 612. 1977 [1978]. TYPE: O. lacin- iata Hill. Onagra sensu Moench, Methodus 1: 675. 1974, pro Oenothera sect. Onagra sensu Ser. ex DC., Prodr. 4: ; , pro parte. Oenothera sect. pig cba sensu Fischer & Meyer, In- dex Sem. rt. Petrop. 2: 44. 1835, pro parte. are bons I Spach, Hist. Nat. Vég. 4: 353. 1835, dan BE. ae III & IV sensu Spach, Nouv. Ann. Mus. Hist. Nat. 4: 347. 1835, pro parte. Oenothera § P E sensu S. Watson, Proc. Amer. . Arts 8: 574. 1873, pro parte. Per en sensu Raimann, Nat. Pflanzenfam. III. 7: 14. 1893, pro parte; sensu Small, Bull. Torrey Bot. Club 23: 172. 1896, pro Oenothera sect. Oenothera sensu Tidestrom, Fl. Ariz. & Mex. 272. 1941, pro parte. Erect to procumbent, annual or perennial herbs yo ceolate to oblanceolate, narrowly elliptic, elliptic, narrowly obovate or narrowly oblong, parted to subentire, the lobes + dentate, the apex acute, the base narrowly cuneate to acute to almost ses- sile; all leaves strigillose or strigillose and villous, sometimes also glandular puberulent. Inflores- cence lax, often with lateral branches, usually only 1 flower per spike opening near sunset each day. Floral tube 1.5-5 cm long, yellowish, often ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 flushed with red, rarely also red-flecked, strigil- lose or glandular puberulent, or villous and glan- dular puberulent, usually curved upward. Sepals greenish to yellowish, often flushed with red or red- -stripe att e margins d-flecked, the pubescence usually as on floral tube, the sepal tips 0.3-5 mm long, erect and appressed or spreading in bud, sometimes separated. Petals very broadly obovate, truncate to emarginate at apex, 0.5-4.5 cm long, 0.5-5.5 cm wide, yellow, sometimes pale yellow, fading red to orange after anthesis. Style 2-7.5 cm long; stigma elevated above the anthers at anthesis or surrounded by the anthers and pollen shed directly onto it. Cap- sule cylindrical, 2-5.5 cm long, 2-5 mm diam. Seeds ellipsoid to subglobose, brown, sometimes with darker flecks (O. drummondii, O. humi- fusa), 0.8-2 mm long, 0.3-0.9 mm diam. Self- incompatible (1 sp.) or self-compatible and mod- ally outcrossing (1 sp.) to modally autogamous (2 spp.), or permanent structural heterozygotes (2 spp.). Gametic chromosome number, n = 7 (7,,, ©14 or intermediate chromosome configu- rations at meiotic metaphase I Series Raimannia of subsect. Raimannia is comprised of six species occurring in open, sandy, and disturbed places, sometimes on dunes, from North Dakota south to Texas and east to the Atlantic Coast, in Mexico along the Gulf Coast, and disjunct in southern Baja California. The species exhibit considerable variation; among them, however, only Oenothera drummondii can be subdivided into two geographically separated subspecies. Typical of the species of series Rai- mannia are loose inflorescences, which often have lateral branches, and upward-curving flower buds. Comparisons with series Candela were made in the discussion of that series. The distribution of series Raimannia is essen- tially the same as that of series Candela but ex- tends farther east, to the Atlantic Coast, and ex- tends south to the state of Campeche, Mexico, along the coast of the Gulf of Mexico. Oenothera drummondii Hook. subsp. thalassaphila (Bran- degee), comb. nov. is disjunct, occurring along the Pacific Eid at the southern tip of Baja Cal- ifornia, Mex x. puits (Britton) Smyth, O. edie riae (described as new below), Spach, and O. drummondii form bivalents whereas O. laciniata Hill and O. humifusa Nutt. are permanent structural heterozygotes. In this section only O. grandis is self-incompatible; all 1987] other species, both Divalent- formers and com- plex structural het are self-compatible and largely autogamous. Oenothera falfurriae W. Dietrich & W. L. Wag- ner, sp. nov. TYPE: Grown from seeds and cultivated in the Botanical Garden of Düs- seldorf, Germany, 2 July 1981, cult. no. 81- 115 from seeds collected in U.S.A. Texas: Brooks Co., 13.3 mi. S of junction of High- ways 281 and 285 in Falfurrias, 10 May 1978, K. Allred & R. Shaw 2021 (holotype, MO- 3332203; isotypes, DUSS, M, MO) Herbae annuae, erectae vel parum decum- bentes, rosulae foliis paucis, 1-4 dm altae. Folia ita .5—4 cm longi, villosi et glanduloso-pubescentes. Sepala 1—2.2 cm longa, viridi-flava, immaculata vel rubro- maculata; apices sepalorum minuti, 0.5-2 mm longi. Petala 1.3-2.5 cm longa, flava vel pallide flava. Capsulae 2-4.5 cm longae, 2-2.5 mm cras- sae, strigillosae, villosae et glanduloso-pubes- entes. Semina 0.8-1.4 mm longa, 0.3-0. crassa. Numerus amas chromosomaticus. n = 7: planta chromosomatice structuraliter ho- mozygotica, autocompatibilis. Erect to decumbent annual herbs, forming a rosette with only a few leaves; stems 1-4 dm long, usually simple, densely to sparsely strigil- lose, villous and sometimes also glandular pu- berulent. Rosette leaves oblanceolate, 5-12 cm bracts elliptic, narrowly ovate to lanceolate, 2— 4.5 cm long, 0.5-2.5 cm wide, dentate or sub- entire to pinnately lobed, narrowed to the base, subsessile; all leaves densely to sparsely villous and glandular puberulent, especially on the mid- rib of the lower surface and along the margin, usually also sparsely to moderately strigillose. Inflorescence lax, simple or with lateral branches, usually only 1 flower per spike opening near sun- set each day, erect at anthesis. Floral tube 2.5- 4 cm long, densely to sparsely villous and glan- dular puberulent. Mature buds lanceoloid to nar- rowly ovoid or oblong-ovoid, 0.4—0.6 cm diam. DIETRICH & WAGNER — NEW TAXA OF OENOTHERA 149 at the base. Sepals green to greenish yellow, sometimes with red spots, the pubescence as on the floral tube, 1—2.2 cm long, the sepal tips 0.5- 2 mm long, erect in bud, strigillose and villous. Petals yellow, broadly obovate, 1.3-2.5 cm long, 1.4-2.7 cm wide, the apex truncate to slightly retuse. Filaments 10-17 mm long; anthers 4—5 mm long; pollen ca. nen fertile. Ovary 1- 1.7 cm long, ca. 1.5 mm diam., densely villous, d E and glandular Sera aim style 3.5-5 m long, the visible part 1.2-3 cm long; stigma deis elevated above the anthers at anthesis, the lobes 3-7 mm long. Capsule cylindrical, 2- 4.5 cm long, 2-2.5 mm diam., the surface pitted. Seeds brown, ellipsqid, 0. an l. 4 mm long, 0.3- 0.6 mm diam. Self ible, modally autog- amous. Gametic chromosome mimber: n=7(7y at meiotic metaphase I). Oenothera falfurriae, named after Falfurrias, Brooks County, Texas, where the type was col- lected, is endemic to open sandy sites in south- eastern Texas. Its range is nearly the same as those of O. bifrons and O. mexicana. When Die- trich first detected the species, the specimens were treated as hybrids between O. grandis and O. laciniata since they were somewhat intermediate between these species. Seed samples collected by K. Allred and R. Shaw made it possible to cul- tivate this species at the Botanical Institute in Dusseldorf, and it soon became obvious that the plants were by no means hybrids, but instead represented an undescribed bivalent-forming species. All plants examined formed 7,, in mei- OSIS, Bs no individuals grown from seed resem- bled either O. laciniata or O. grandis. The in- i ca collection numbers of Allred and Shaw represent population samples of several plants from which seeds were taken and sowed sepa- rately: 2016, 2020, and 2021 contained O. fal- furriae and O. laciniata; 2018 contained O. fal- furriae and O. mexicana. Seeds taken from plants of O. falfurriae produced only O. falfurriae and those of O. laciniata produced only O. laciniata. Oenothera falfurriae differs from O. grandis in its self-compatibility and smaller petals, which are intermediate in size between those of O. /a- ciniata and those of O. grandis. Stigmas in the closed mature buds are only slightly raised above the anthers, suggesting that self-pollination is common in O. falfurriae. Also, the shape of the buds is more or less oblong, in contrast with the lanceoloid buds of O. grandis, and the sepals in O. falfurriae are very delicate and pressed to- 150 gether in bud, whereas in O. grandis they are often spreading, longer, and thicker. tan 3 fal is narrowly presumably relictual. It appears to maintain itself distinct from the other species of series Rai- mannia with which it grows sympatrically — O. grandis, O. laciniata, and O. mexicana — by pos- sessing a unique plastome. Artificial crosses made by Dr. Behn at the Botanical Institute in Düs- seldorf showed that crosses between O. drum- mondii or O. humifusa and O. falfurriae as the staminate parent produced pale seedlings that failed to grow beyond the cotyledon stage. Sim- ilarly, the seeds of crosses between O. grandis and O. falfurriae did not germinate at all (Behn, pers. comm.). Also, since crosses between O. drummondii, O. humifusa, or O. grandis and O. laciniata produce completely green and viable offspring, we can assume that similar crossing barriers exist between O. falfurriae and O. laci- niata, based on the pattern of such relationship in Oenothera sect. Oenothera generally. Oenothera drummondii Hook. subsp. thalassa- phila irap be Dietrich & W. L. Wag- ner, comb. Based on Oenothera tha- lassaphila dde Univ. Calif. Publ. Bot. 10:185. 1922. Oenothera drummondii Hook. var. thalassaphila (Brandegee) Munz, Amer. J. Bot. 22: 651. 1935. rvPE: Mexico. Baja California Sur: San José del Cabo, 12 Mar. 1892, T. S. Brandegee 216 (lectotype, UC- 107674; see Munz, Amer. J. Bot. 22: 651. 1935). The separation of Oenothera drummondii subsp. thalassaphila, which is restricted to dunes of coastal southern Baja California del Sur, Mex- ico, from subsp. drummondii depends on a com- bination of characters since no single morpho- logical feature separates them clearly. Oenothera drummondii subsp. thalassaphila always grows ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 for several years, as is demonstrated by the con- sistent presence of nonflowering shoots and large taproots on the older plants. By contrast, O. drummondii subsp. drummondii is basically an annual, seldom overwintering for a second sea- son; it usually has only a few nonflowering shoots or none, and the development of its taproot is considerably weaker than in subsp. thalassaphi- la. In general, the habit of subsp. drummondii is more upright than that of subsp. thallassaphila, which has prostrate to ascending stems. In ad- dition, the calyx of subsp. thalassaphila often has red spots and lacks glandular hairs, whereas the calyx of subsp. drummondii only rarely has red- dish spots and is often glandular puberulent. The sizes of the capsules and seeds are also modally distinct: in subsp. thalassaphila the capsules are 2-4 cm long and 2.5-5 mm in diameter, and the seeds are 1.5-2 mm long and 0.7-0.9 mm in diameter; in subsp. drummondii the capsules are 2.5-5.5 cm long and 2-3 mm in diameter, and the seeds are 1.1-1.7 mm long and 0.5-0.8 mm in diameter. LITERATURE CITED a W. 7 [1978]. The South American ecies of Oe es sect. Oenothera (Raimannia, | fein. Onagraceae). Ann. Missouri Bot. Gard. 64: 425-626. RAVEN & W. L. WAGNER. 1985. Re- vision of Oenothera sect. Oenothera subsect. uci eur SEP nied Syst. Bot. 10: 29-48. Muwz, P. A. . 1979 [1980]. An outline of the systematics of Oenothera sub- sect. Euoenothera (Onagraceae). Syst. Bot. 4: 242- 252 STUBBE, W. & P. H. Raven. 1979. A genetic contri- bution to the taxonomy of Oenothera sect. Oe- nothera (including subsections Euoenothera, Emersonia, Raimannia and Munzia). Pl. Syst. Evol. 133: 39-59 A NEW COMBINATION AND NEW SUBSPECIES IN OENOTHERA ELATA KUNTH (ONAGRACEAE)! WERNER DIETRICH? AND WARREN L. WAGNER? ABSTRACT The new combination of Oenothera elata Kunth subsp. hookeri (Torrey & A. Gray) W. Dietrich & ri County, ed fro razos Texas. It appears to be a rare relictual entity most closely related to O. ati subsp. monte ys which occurs disjunctly some 680 km to the west of subsp. texen These names are made avallable in anticipa- tion of their use in regional floras and concurrent studies of flavonoids, cytology, and pollen mor- phology in advance of a detailed revision of Oe- nothera subsect. Oenothera (Dietrich & Wagner, in prep.). Detailed presentation of data and dis- cussions will be given in the revision. Munz (1949, 1965) divided the large-flowered, bivalent-forming, outcrossing populations of sect. Oenothera subsect. Oenothera from the western United States south to Panama into two species, O. hookeri Torrey & A. Gray and O. elata Kunth. During the past ten years we have reevaluated the variation pattern of these plants by a detailed study of a large number of herbarium specimens, fieldwork, and study of plants cultivated in the experimental gardens at Diisseldorf. These stud- ies have shown that the two entities are extremel act diameter, and modally in several other features. It was therefore suggested by Raven et al. (1979), in an outline of the systematics of Oenothera sect. Oenothera subsect. Oenothera (formerly Eu- oenothera), that they should be considered con- specific Oenothera elata can be subdivided into four subspecies: subsp. e/ata with a scattered distri- bution from Guanajuato, Mexico, to Costa Rica and Panama in Central America; subsp. hirsu- tissima (A. Gray ex S. Watson) W. Dietrich, which Occurs in the western United States from Wash- ner, 1983); subsp. texensis subsp. nov., known only from one collection in Brazos County, Tex- as; and subsp. hookeri (Torrey & A. Gray) comb. et stat. nov. occurring in moist coastal and slight- ly inland sandy and bluff sites in California from Marin County south to San Diego County. The four subspecies of Oenothera elata can be distinguished with the following key. KEY TO THE SUBSPECIES OF OENOTHERA ELATA la. Stem, leaves, and ovary (capsule) exclusively ) with red; ps s tips of the capsule distinct ee 2b. Stem usually green; the = e of th capsule indistinct z Ma ture buds (excl. oo ae nar- rowly lanceolate in outline 5-5 cm "Pa ve tips 2-3 mm long; petals m long; capsule 5-6.5 c ur bcn undulate; leaves mem- branous; plant in d asa to . texensis - ceolate in outline, a cm more than 10 dm tall ............. bsp. elata Ib. Stem and ovary (capsule) predominantly with erect pubescence (short and long villous), stem without glandular hairs . . subsp. Airsutissima 4b. Stem in the region of the inflorescence with glandular hairs 5a. Sepals green or - flushed with red, without or with indistinct pustulate hairs, sparsely to scattered villous; ' This work has been supported by a series of grants from the National Science Foundation to Peter H. Raven. ae d Institut der Universitat Düsseldorf, Universitatsstr. 1, D-4000 Düsseldorf 1, West Germany. ernice P. Bishop Museum, P.O. Box 19000-A, Honolulu, Hawaii 96817, U.S.A. ANN. Missouni Bor. GARD. 74: 151-152. 1987. 152 plant in cultivation id than dm p. hirsutissima ; Sepals always flushed us red, w ith wn c vation not more than 8 dm tall ..... subsp. hookeri The plants described here as O. elata subsp. texensis are known only from a single herbarium specimen and from material cultivated at Düs- seldorf. In 1981 Dietrich collected a seed of an unusual Oenothera from a TRT herbarium spec- imen. From the large flowers it appeared to rep- resent O. grandiflora L'Hér., but when plants were grown in Düsseldorf they clearly repre- sented O. elata. Further, based on its strigillose pubescence it appeared to be subsp. Airsutissima; however, other characters clearly separated it from both strigillose pubescent subspecies of O. elata, subspp. elata and hirsutissima. The stems of the Brazos County plants are always green, the capsules are 5—6.5 cm long, the buds are narrowly lanceolate, and the leaves are membranous. These plants grow tall in cultivation, to 2 m or more. Oenothera elata Kunth subsp. texensis W. Die- trich & W. L. Wagner, subsp. nov. TYPE: Grown from seeds taken from herbarium specimen TRT-205991 and cultivated at the Botanical Garden of the University of Düs- seldorf, 12 Sept. 1984, cult. no. Stubbe 84- 204; original source, U.S.A. Texas: Brazos Co.,ca. 17 km NW of Navasota River bridge on Hwy. 6 in vicinity of Peach Creek cutoff, 25 Oct. 1978, P. M. Catling & K. L. Intosh (holotype, MO-3332204; isotypes, DUSS, M) Herbae biennes, erectae, in culturam usque ad 20 si. Folia undulata m flava, 4.5-5.5 cm longa. Stylus longus, stigmate sub anthesi supra unde edipi) Capsulae 5-6.5 cm lon- gae, strigillosae. Numerus gameticus chromesomati- cus, n = 7; 1 uis ires zem homozygotica (7 bivalenta in metaphasium primum meiosis), autocom- patibilis ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Munz (1949, 1965) recognized two subspecies of his Oenothera hookeri from moist coastal or slightly inland sites in California: subsp. hookeri and subsp. montereyensis. He referred plants with a bushy habit, blunt buds, free sepal tips 1-2.5 mm long, and sepals usually 2-2.5 cm long to subsp. montereyensis, whereas plants with a less bu hed habit, attenuate buds, free sepal tips S referred to subsp. hookeri. shown that these characters represent intra- and inter-populational variation and, further, ap- pear to vary independently. The bushier habit of some coastal plants appears to be an adaptation to wind and salt spray. All of these populations are here treated as members of one coastal sub- species of Oenothera elata distinguished from the other three subspecies of O. elata primarily y its densely glandular puberulent and long- villous buds. Oenothera elata Kunth subsp. pp dol & A. Gray) W. Dietrich & W. Wagner, comb. et stat. nov. Oenothera hookeri Torrey & A. Gray, Fl. N. Am 49 0. TvPE: California [without stir sa n cality] ouis s.n. (holotype, Oenothera hookeri Torr. & A. Gra subsp. monterey- ensis Munz, Aliso 2: 14. 1949. TYPE: United States. California: Monterey Co., 0.2 mi. S of mouth of Alder Creek, 6 Nov. 1934, C. B. Wolf6223 (RSA- 12778, holotype (not seen); isotypes, GH, NY, OM). LITERATURE CITED Munz, P. A. 1949. The Oenothera hookeri group. Aliso 2: 1-47. ————. 1965. ney Fl. N. Amer. II. 5: 1-278 RAVEN, P. H., W. D dole ipiis Tees An outline of ib atics of Oenothera sub- sect. peepee ae Syst. Bot. 4: 242- WAGNER, W.L. New species and combinations in the genus Oenothera heey ia Ann. Mis- souri Bot. Gard. 70: —196. A REVISION OF MEZILAURUS (LAURACEAE)! HENK VAN DER WERFF? ABSTRACT The neotropical genus Mezilaurus (Lauraceae), which consists of 18 species and is best represented in the drainage area of the A from Licaria and reinstated on its caatingae van der mazon, is revised. Clinostemon, a genus of two species recently separated own, is included here in Mezilaurus. Eight sp Werff, M. duckei van der Werff, M. glaucophylla van der Werff, M. micrantha van ecies, Mezilaurus der Werff, M. opaca Kubitzki & van der Werff, M. due ped van 5 e M. pyriflora van der Werff, and M. quadrilocellata van der Werff are described as new. One me, M. thoroflora van der Werff, and a new combination, M. mahuba Tree van ton Werff a are pere a Mezilaurus comprises 18 species, occurring from Costa Rica to southern Brazil. The majority of the species are found in the drainage area of the Amazon River and adjacent Guayana. The type species, M. navalium, is restricted. to the to insufficient material) are shrubs from the cer- rado vegetation. Fourteen species are Amazo- nian and have been reported from a variety of habitats. Ten Amazonian species, ranging from shrubs to tall trees, occur in terra firme vegeta- tion, frequently in xeromorphic forests on white sand. One species, M. mahuba, is restricted to flooded forest. Three species are known to occur in secondary vegetation (M. thoroflora, M. syn- andra and M. lindaviana), although it is not clear whether these are typical of secondary vegetation or were left standing when the primary forest was vesting. Lauraceae, widely used for timber, are therefore frequently found as isolated trees in pastures or similarly disturbed habitats. No hab- itat information is available for the Colombian species, which is only known from Chigorodó in northern Antioquia. The Costa Rican species oc- curs in wet lowland forest near the Pacific Coast. Most species of Mezilaurus are collected in- frequently, and I have seen more than ten col- lect ctions for only two species (M. itauba and M. many Mezilaurus species are ave doubt explains the paucity of collections. The genus is greatly undercollected and much more material is needed for a better taxonomic un- derstanding. The main use of Mezilaurus is for timber. The species are locally well known and their hard wood is much used for boat building and con- struction. Mez (1889) mentioned that the fruits of M. itauba are edible. On the label of Fróes 12152 (Mezilaurus pyriflora) it is stated that the wood causes injuries to the skin, presumably a kind of dermatitis. The present revision was undertaken because it became clear that the genus C/inostemon should be merged with Mezilaurus. This led to a closer look at recent collections, during which several undescribed species were found. I now recognize 18 species in Mezilaurus, which more than dou- bles the number of species recognized by Kos- termans (1938 MATERIALS This study is based both on older collections, already cited by Kostermans (1938), and recent collections personally selected during visits to major American and European herbaria or re- ceived on loan with other unidentified Laura- ceae. In my experience the genus was unrecog- nized in many herbaria. Unfortunately, I have not yet had the opportunity to visit the leading Brazilian herbaria. I fully expect that such visits will yield additional taxa, not only from the Am- in this group of small-flowered Lauraceae. ! I thank the curators of AAU, BM, BR, C, F, G, HUH, K, L, NY, U, and US for the loan of specimens. Dr. A. Gentry suggested improvements in the text, and Dr. J. Dwyer corrected the Latin descriptions. John Myers made the illustrations. ? Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. ANN. MissouRi Bor. GARD. 74: 153-182. 1987. TAXONOMIC HISTORY The first species of Mezilaurus was published by Allemáo (1848) as Silvia navalium. Meissner (1864) recognized that Silvia Allemào was a later homonym of Silvia Vellozo and proposed the new name Silvaea. Unfortunately, Silvaea Meissner is a later homonym of Si/vaea Phillipi. O. Kuntze (1891) proposed another new name, Mezia, to replace Silvia Allemào, but the name Mezia O. Kuntze was predated by Mezia Schwacke, a genus of Malpighiaceae. Finally, Taubert (1892) proposed the name Mezilaurus, which Mez (1892) accepted. He included seven species in the genus. Pax (1897), overlooking the publication of Mezilaurus, proposed Neosilvia as new name for Silvia Allemao. Neosilvia is therefore a superfluous name. The name Mezi- laurus was not universally accepted at first. Even Mez (1904, 1920, 1924) used the name Silvia again. Ducke (1930, 1935) published several new species under the generic name Silvia. Koster- mans's revision (1938) definitively established the use of Mezilaurus Taubert During the nineteenth century, several species now placed in Mezilaurus were described in oth- er genera, mostly in genera now included in Li- caria. Meissner (1864) described three species in Acrodiclidium and one in Oreodaphne (a syn- onym of Ocotea). Bentham in Hooker (1878) transferred two of these to Misanteca (a synonym of Licaria) and Bentham and Hooker (1880) placed the other two (including the type species of Mezilaurus) in Endiandra, an Old World ge- nus. Mez (1889) recognized six species in Me- zilaurus (as Silvia), the type species, the four species described by Meissner (1864), and a sixth species now included in Licaria. Kostermans (1938) accepted eight species of Mezilaurus, four of Mez's species and four described since 1889 by Mez and Ducke. Allen (1964) described two new species of Mezilaurus from Venezuela. Kuhlmann and Sampaio (1928) published the monotypic genus Clinostemon based on Acro- diclidium mahuba Samp. Kostermans (1938) did not accept this genus and placed it in Licaria. Later Allen (1948) described a new species closely related to Licaria mahuba and noted the resem- blance to Mezilaurus. Recent investigations (Ku- bitzki et al., 1979) have shown that these two species do not belong in Licaria and are more closely related to or congeneric with Mezilaurus. In this paper both are included in Mezilaurus. The eight species recognized in Kostermans's ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 monograph are all maintained in this publica- tion. The increase in the number of Mezilaurus species is partly due to the inclusion of Clino- stemon in Mezilaurus and partly due to recent collections that represent undescribed species. GENERIC RELATIONSHIPS One of the main taxonomic difficulties in Lau- raceae is that many of the genera are poorly de- fined. This is reflected in the various infrafamilial classification schemes and the frequency with which species are transferred between various genera. During the last 30 years generic relation- ships within the Lauraceae have been discussed in three papers (Kostermans, 1957; Hutchinson, 1964; and Richter, 1981) Kostermans (1957) attached much importance to the fruit (with or without cupule) and less importance to the number of anther cells; he placed Mezilaurus in the subtribe Beilschmie- diinae of the tribe Perseeae, close to Endiandra, and noted, as did Bentham and Hooker (1880), a similarity to Endiandra. He also stated that Endiandra and Mezilaurus differ in anther shape and the positions of their leaves. Additional dif- ferences are the very fine reticulation of the leaves and the large, spreading tepals in many species of Endiandra (comparable to tepals in Nectan- dra), which are very unlike the small, erect, scale- like tepals in Mezilaurus. These differences great- ly outweigh the similarities between Endiandra and Mezilaurus (number of fertile stamens, number of anther cells, fruit more or less without cupule) and Endiandra is probably not a close relative of Mezilaurus Hutchinson (1964) considered the number of anther cells more important than the develop- ment of the cupule. He placed Mezilaurus next to Misanteca, but the distribution data given un- der Misanteca strongly suggest that Hutchinson, as did Bentham and Hooker (1880), included in his Misanteca two Brazilian species here includ- ed in Mezilaurus. As can be seen from the taxonomic history of Mezilaurus, its species frequently have been placed in Licaria (or its synonyms Acrodiclidium and Misanteca). These two genera have in com- mon that they are the only neotropical Lauraceae with three fertile, two-celled stamens. The gen- eral flower shape of species belonging to these two genera can be quite similar and a generic separation based solely on flowers can be ve difficult. Fortunately, other characters readily 1987] identify the genera: Mezilaurus has the leaves always clustered at the tips of the branches, Li- caria never; in Mezilaurus the cupule consists of a small, platelike disk, in Licaria it grows into a rather large, double-rimmed cup, and the inflo- rescence of Mezilaurus is a double raceme (Figs. 3, 8C), an inflorescence type never found in Li- caria. Kubitzki et al. (1979) partly enumerated these differences and discussed the placement of Licaria mahuba (Samp.) Kostermans and L. ma- guireana Allen. They concluded that these species did not belong in Licaria and resurrected the generic name inostemon to accommodate them. Clinostemon was considered a close rela- tive of Mezilaurus, the only difference being the presence of staminodia in Clinostemon and their absence in Mezilaurus. The two Clinostemon species also share edi obovate leaves with an abruptly rounded bas The taxonomic DR of absence/pres- ence of staminodia in defining genera of Laura- ceae is open to discussion. In some genera stam- inodia are consistently present (Persea, Phoebe); in others they may be present or absent (Aiouea, Aniba, Licaria, Ocotea). This suggests that a ge- neric separation based only on presence/absence of staminodia is weak, especially because the staminodia are small, ca. 0.5 mm, and not easy to find. The discovery oftwo undescribed species in Colombia and Costa Rica with the leaf shape and size of a Mezilaurus, but with staminodia like Clinostemon, is another reason to place Cli- nostemon in synonymy under Mezilaurus. Richter (1981) amply discussed the wood and bark anatomy of the Lauraceae. He found that within the Lauraceae, Mezilaurus occupied an isolated position and was easily recognized both on wood and bark characters. He also found that the wood (bark was not available) of the two Clinostemon species was either undistinguisha- iz ae erah earlier in iie ed et ES (1979). The great similarity in wood ana plus the iso- lated position of Mezilaurus/ Posi aodio in the Lauraceae is the second reason to merge these two genera. Richter (1981) also found that Li- caria and Endiandra are, as far as wood anatomy is concerned, not closely related to Mezilaurus. The genus most closely related to Mezilaurus by wood characters is Anaueria, a monotypic genus incompletely known from a few collections in Brazil and Peru. Kostermans (1952) and Hutch- WERFF—REVISION OF MEZILAURUS 155 inson (1964) both placed Anaueria in Beilsch- miedia; Kostermans mentioned, without giving details, that he did so as a result of studying additional herbarium material from Rio de Ja- neiro. Neither Anaueria nor Beilschmiedia is likely to be confused with Mezilaurus, since they have flowers with six or nine fertile stamens and never have clustered leaves. Two species described in this paper merit ad- ditional comments. Mezilaurus ines ie and M. glaucophylla have an unusual di tion, being only known from northern eade and Costa Rica. All other Mezilaurus species oc- cur in the Amazonian forests or other parts of Brazil. Secondly, M. quadrilocellata and M. glaucophylla have staminodia (as do the species formerly placed in Clinostemon) and the leaf shape of Mezilaurus species (quite unlike the species formerly included in Clinostemon). Thus, they link Mezilaurus with Clinostemon. More- over, they are the only species in the genus with four anther cells on each stamen. Given the im- portance frequently attached to the number of anther cells and the number of fertile stamens, the presence of three four-celled anthers (not found in any other New World Lauraceae) could be sufficient for the recognition of a new genus. However several other genera include species with two-celled and four-celled anthers, and because other characters (leaves clustered at branch tip, shape of the inflorescence) point toward Mezi- laurus, I include these species in Mezilaurus. As a result of the transfer of Clinostemon and the inclusion of Mezilaurus quadrilocellata and M. glaucophylla, my concept of Mezilaurus is wider than has been used by previous authors. I regard as diagnostic characters the leaf position (clustered at the tips of branches), the small, platelike cupule (but fruits from most species are not yet known), the type of inflorescence (a dou- ble raceme), and the presence of three fertile sta- mens. Richter (1981) discussed diagnostic wood s bark characters. n, Mezilaurus shows in floral char- acters a strong resemblance to Licaria. However, differ in characters of leaf position, inflorescence type, and number of fertile anthers. A close re- lationship between Mezilaurus and Endiandra is very unlikely. Currently available information indicates that Mezilaurus, including Clinoste- 156 mon, is endemic to the Neotropics, and that it occupies an isolated position in the family MORPHOLOGY AND TAXONOMIC CHARACTERS Mezilaurus species range from small trees or shrubs (the cerrado species) to tall forest trees much valued for their timber. The twigs are gen- erally thick, show conspicuous leaf scars, and are The leaves in all species are pin- nately veined. The lateral veins frequently arch upward and become connected with the more distal lateral vein. The texture of the leaves is variable; most species have chartaceous leaves, but a few have coriaceous leaves in which sec- ondary and tertiary venation is poorly visible. Conspicuous gland dots in the leaves occur rare- ly. The leaves generally turn dark upon drying. Characteristic for the genus is the fact that the leaves are always clustered at the tips of the branches. Young shoots grow initially rapidly without beanie leaves; after this elongating period, leaves are formed at the tip of the young branch. Such branches may have several clusters of leaves, representing different growing seasons. I will call this growth pattern long shoot-short shoot. Under unfavorable conditions (several Mezilaurus species are reported from white sand forests or caatinga forests) the difference between the long shoots and short shoots becomes less pronounced and the growth pattern may seem a succession of short shoots, with only one cluster of leaves at the tip of the branches. However, I think the difference between long shoot-short shoot or short shoot growth pattern is quanti- tative, not qualitative. Species with clustered leaves of the long shoot- short shoot pattern occur regularly, but not fre- quently, in several other neotropical genera of Lauraceae (Aniba, Endlicheria, Nectandra, Oco- tea, Phoebe, and Pleurothyrium). However, only in Mezilaurus is this clustered leaf pattern char- acterisitic or dominant. The non-Mezilaurus species with clustered leaves are rarely confused with Mezilaurus; even in vegetative state they are readily separated by conspicuous pubescence or leaf color differences. Only one species, Ocotea rubra, very closely resembles Mezi/aurus in ster- ile state. However, its flowers with nine four- celled stamens and fruit with a large cupule make identification easy early all species of Mezilaurus have elliptic to obovate leaves. The base of the leaves, how- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 ever, offers some useful characters. Four species, ezilaurus mahuba, M. pyriflora, M. thoroflora, and M. duckei, have large leaves (to 60 cm long) which become gradually narrowed toward the base; at the base the leaves are abruptly nar- m: becoming rounded or even cordate. Three pecies, M. subcordata, M. quadrilocellata, and M glaucophylla, have an obtuse or rounded leaf base with a distinct petiole, 2-6 cm long. In these three species the leaf shape is slightly obovate or ovate. In all other species (with the exception of M. crassiramea), the leaf base is gradually atten- uate or decurrent on the petiole. Most of these species have a petiole. Three species, however, M. caatingae, M. crassiramea and M. decurrens, have sessile leaves or nearly so. In some collec- tions of M. crassiramea the leaf base is gradually narrowed, in others it is rather abruptly nar- rowed. In this species the petioles, if present, are less than 1 cm long Pubescence. There is not much variation in pubescence within Mezilaurus. Two species, M. crassiramea and M. lindaviana, have erect pu- bescence on the lower surface of the leaves; this can be quite sparse in M. /indaviana, however. The other species have varying amounts of ap- pressed pubescence on leaves, stems, terminal buds, and inflorescences. These varying amounts of pubescence have little diagnostic value. Inflorescence. The inflorescence of Mezilau- rus consists of acompound raceme (dibothryum, see Weberling, 1981, 1985). This inflorescence type is present without modifications in M. cras- siramea, M. lindaviana, M. mahuba, M. pyri- flora, M. thoroflora, and M. duckei. Short tertiary axes are sometimes present in M. glaucophylla and M. quadrilocellata. In the other species the inflorescences are smaller and the flowers are not evenly distributed along the lateral branchlets of clustered flowers are a derive worth noting that the species with a well-devel- oped dibothryum have larger (often much larger) inflorescences and have usually smaller flowers than the species with clustered flowers. A di- bothryum is a rare inflorescence type among oth- er neotropical Lauraceae (if it ever occurs outside of Mezilaurus). The inflorescence type of one Mezilaurus species, M. decurrens, is not known due to the fragmentary nature of the single available spec- imen. 1987] Flowers. In the following discussion I con- sider only mature flowers. Because it is difficult to tell whether a flower is mature or not in Me- zilaurus, I define a mature flower as one with opened anther cells. In buds or young flowers diagnostic characters are often difficult to see. The flowers of Mezilaurus are unusually dif- ficult to dissect. In addition to their small size, the floral tube contains much mucilage, which has usually hardened during drying. Softening the flowers requires boiling for at least one hour. After softening, the mucilage becomes viscid and sticky and the dissected floral parts frequently stick tightly to the floral tube. Therefore, I have rarely relied on characters of the floral parts to separate taxa, especially because other characters are available. Dimensions given for flowers and their parts should be accepted with some reservation. In order to dissect the flowers, it is necessary to boil them. During the boiling the flowers swell; the degree of size increase depends on the duration of boiling. I have recorded flowers that measured 3 mm dry as swelling to 1.7 mm after boiling. Similar increases in size were found for the sta- mens as well. Only rarely have I used flower sizes in the key and, in these cases, flower sizes are taken from dry flowers. i i are acu small and equal. The exceptions are M. cophylla and M. pedem in S the outer three tepals are smaller than the inner three. With the exception of three species (Mezilau- rus caatingae, M. palcazuensis, and M. mahuba), all species have clearly pedicellate flowers. Stam- inodia (which are not easy to find due to their small size) are present in six species (M. duckei, M. glaucophylla, M. mahuba, M. pyriflora, M. quadrilocellata, and M. thoroflora), and only M. mahuba has glands at the base of the fertile sta- mens. The most interesting variation of floral struc- tures 1s found in the shapes and positions of an- thers. In most neotropical Lauraceae the anthers have the shape of erect stalked plates with the anther cells on the inner or outer surface (for instance, in Persea, Phoebe, Nectandra, and Oco- tea). In several of the genera with two-celled an- thers the plate shape of the anthers is less pro- nounced or lost and the anther cells are situated at or near the tip of the anthers (Aniba, Licaria). In several species of Licaria the stamens are shaped like columns with the anther cells situ- ated near the apex and here the difference be- WERFF—REVISION OF MEZILAURUS 157 tween filament and anther has disappeared. In a few species of Mezilaurus (M. duckei, M. glau cophylla, M. pyriflora, and M. edv san en a similar arrangement is found. Here the stamens remain included in the erect tepals and the anther cells are situated at the tips of the stamens. In M. glaucophylla and M. quadrilocellata, the only species with four anther cells on each stamen, the tips of the stamens are flattened and form a small platform on which the anther cells are sit- uated. Thus, by looking from the outside in the flowers, one sees the 12 anther cells as small pores. In these four species the anther cells are situated iim and also open apically; that is, the flaps open upwar Mezilaurus ot is the only species with the anther cells situated introrse or introrse-lat- eral; in all remaining species the anther cells are extrorse, situated on the outer face of the stamen. Here the anther cells are more or less exserted, tend to be relatively large, and, most interesting, the stamens develop a dorsal ridge on which the anther cells are situated. The ridge is usually about as long as the anther cells and appears as an outward-facing hump near the tip of the stamen. The anther cells open towards the crest of the ridge, where the flaps are situated back-to-back. Such anther cells have been described as opening “laterally” in the literature, but in order to avoid confusion with laterally situated anther cells (which occur, for instance, in Pleurothyrium), I will call this type of opening ‘“‘back-to-back.” n decurrens the dorsal ridges are not strongly dexeloned and the anther cells are hard- ly exserted. In the remaining species the anther cells are clearly exserted. e most extreme development is found in M. ahuba, where the dorsal ridges with the anther fro greatly exserted anther cells does not correspond with variation in other characters. In fact, the extremes (M. pyriflora with immersed anther cells, M. mahuba with greatly exserted anther cells) are very similar in other characters such as leaf shape, leaf size, and type of inflorescence. In a few species (M. crassiramea, M. linda- viana, M. palcazuensis, M. sprucei, M. subcor- data, and M. synandra) the filaments of the sta- mens are fused in a ring or a short tube. This is a useful character, but because it is hard to rec- ognize, I have not used it in the key. It is most easily seen on young fruits. When the filaments are free, they are visible at the base of the fruit; 158 ANNALS OF THE MISSOURI BOTANICAL GARDEN when the filaments are united, they are visible as a small cap on top of the fruit. REPRODUCTIVE BIOLOGY In a study of the reproductive biology of some neotropical Lauraceae, Kurz (1983) and Kubitz- ki and Kurz (1984) reported that a Clinostemon species here described as Mezilaurus duckei showed synchronized dichogamy with a pro- nounced protogyny. In this system, two flowering morphs are found. In the A morph, flowers open in the morning and expose the receptive stigmas. During this phase, no pollen is released. Around midday, the stigma wilts and is no longer receptive. The male phase, when the anthers shed the pollen, takes place in the afternoon. An individual flower thus lasts only one day. In the B morph, flowers open in the afternoon, when the stigmas are receptive. Pollen is released during the morning of the fol- lowing day. During the fieldwork, Kurz was able to observe only two flowering Mezilaurus trees and these turned out to be both A morphs. However, he studied A and B morphs of other Lauraceae species and found that for fertilization, cross- pollination between A and B morphs was nec- essary. Selfing of A or B morphs did not result in seed set. It is likely that these findings apply to Mezilaurus as well. The Mezilaurus flowers observed by Kurz were visited by four species of small (2-3 mm) Trigona bees (Meliponinae). Mezilaurus flowers are not [Vor. 74 known to produce nectar and it is likely that pollen is the only reward for their pollinators. TAXONOMIC TREATMENT Mezilaurus Taubert, Bot. Centralbl. 50: 21. 1892. TYPE: M. navalium (Allemao) Taubert. Silvia Allemào, Dissertatio, Rio de Janeiro. 1848, non Conc. Silvaea Meissner, DC. Prodr. 15: 84. wa Natürlichen Pflanzenfamilien, Nachtrag zu Teil II-IV. 1897, nom. super Clinostemon Kuhlm . & Samp., B s. Nac. Rio de Janeiro 4(2): 57. 1928. TYPE: ys ep a w (Samp.) uhlm. & Samp. Shrubs to tall trees, mostly South American, but with one species in Costa Rica. Leaves al- ternate, usually congested at the apex of the twigs, entire. Petioles often swollen at base. Inflores- cences axillary, sometimes seemingly terminal, few- to many-flowered, forming a compound ra- ceme (dibothryum); flowers clustered at the tips of the inflorescence branchlets in several species. Bracts and bracteoles deciduous. Tepals 6, equal or nearly so, small, scalelike, usually erect. Fertile stamens 3, representing the third whorl, 2-celled (in two species 4-celled). Staminodia present or absent. Staminal glands present in one species. Filaments free or connate; anther cells usually extrorse and exserted, situated on a dorsal ridge. Ovary ellipsoid to ovoid, included in the flower tube. Fruit ellipsoid, situated on a small, plate- like cupule. KEY TO MEZILAURUS la. Stamens 4-celled; N. Colombia and Costa Rica 2 ter tiary FF quadrilocellata N. Colombia ....... 2b. Inflorescence gray ron idi glabrous or with few basal hairs; tertiary ventu on upper leaf surface raised; Cost M. P = Stamens 2-celled; S. America 'E " the Andes 3a Leaves d narrowed toward base, usually abruptly rounded there, generally large, ex- nes 25c +} 4 uetlal Amazon 1 To1 3a E POLI J from fl M. mahuba 5 4b. esum pedicellate; anthers not hooklike exserted; not occurring in flooded forest Sa. Leaves elliptic or slightly obovate, rounded at tip; anthers not exserted at anthesis M. duc s Sb. Leaves strongly obovate, acute or acuminate at tip; anthers exserted or not ........ 6a. Ant hers exserted at anthesis; pedicels 4-8 mm long Anthers included at anthesis; pedicels 1.5-2 mm lon 3b. Leaves decurrent or obtuse at base, generally ue rarely exceeding 20 cm Leaves M. oda SM M. pyriflora 7 7a pubescent below; twigs visibly pubescent a owers pubescent; shrub or id tree in cerrado vegetation U U M. crassiramea Flo abrous; tree in rain fore M. lindaviana 7b. Leaves glabrous below; twigs not visibly pubesce 9 f tip acute (in M. micrantha apex is iid but present) 10 9b. Leaf tip rounded 12 1987] 10a. Flowers sessile; Per WERFF—REVISION OF MEZILAURUS 159 10b. Flowers pedicellate; "Brazil or Peru . lla. Leaves coriaceous; pedi 11b. Leaves — wa: pedicels (5)10-15 n long abrous 12a. Flow M. palcazuensis 11 M. micrantha M. sprucei 13 cels ca. 2 mm Flowers sessile; caatinga forests along Rio Negro ... j caatingae 13b. Flowers en rain forests of S. Brazi 12b. Flowers pubesce ll u. ge 14a. Leaf base yee or rounded; anther cells (lateral) uen N 14b. Leafb 11 š *4 th + subcordata 1 a. 16b. Leav l wn o l M S Mezilaurus caatingae van der Werff, sp. nov. TYPE: Brazil. Amazonas: Rio Negro, Sao Fe- lipe, caatinga on sandy soil, tree, 15 m, 27 Sept. 1952, Fróes 28761 (holotype, MO). Figures 1, A & B; 2 Arbor, 15 m. Ramuli teretes, glabri. Folia coriacea, $ ne ‘ad apices. ramulorum, glabra, margine invo- 0-15 x Poe super laevia nitidaque, subtus opaca costa elevata nervis lateralibus et venatione immersa. Petioli . 1 cm lon axillaris pubes scens (basis inflorescentia tantum adest). Flores labri vel basi leviter minute adpresse pubescentes, ronis, sessiles, conferti ad apices ramulorum inflo- rescentiae sie m longi. Tepala 6, aequalia, erecta, pa 0 m). Stamina 3, per anthesim exserta, | , extrorsis lateraliter des ntibu us. Filamenta libra, pubescentia, antheris ae- e re sm Tubus per glaber. Ovarium gla- brum, globosum, 0.5 m ylus 0.6 mm longus, per anthesim a Fructus Med Tree, 15 m. Twigs terete, glabrous. Leaves clustered at the tips of biais glabrous, cori- aceous, obovate, 10-15 x 4—5 cm, the tip round- ed, the base gradually debi to the short pet- iole, the margin inrolled, the upper surface smooth, slightly lustrous, the lower surface with a raised midvein; lateral veins (8-12) and reti- culation + immersed; petioles ca. 1 cm, the lam- ina decurrent as two narrow ridges. Inflorescence axillary, minutely appressed pubescent, panicu- lately branched, broken on the specimen seen. Flowers glabrous or with some minute pubes- cence at the base, sessile, clustered at the tips of the inflorescence branchlets, 2.2 mm long, ob- 15a. Tepals erect; : flower tube not ‘constricted at apex flowers cup-shaped .. . Flowers 1.5 x 1.5 mm; upper leaf surface PE M. ALULUISUC Leaves sessile or nearly so; anthers scarcely exsert- m M. decurrens 1 cm long; anthers M. itauba . Tepals incurved; Ere tube constricted at apex; flowers depressed globos 17a. Flowers 2 x iege m; upper leaf surface shiny and with raised reticulation ynandra the reticulation not raise opaca conic. Tepals 6, very small, 0.2 mm long, equal, erect. Stamens 3, exserted at anthesis, | mm long, the anthers 2-celled, extrorse, opening back-to- back; filaments free, as wide as anthers, pubes- cent. Floral tube glabrous. Ovary globose, gla- brous, 0.5 mm. Style slender, 0.6 mm long, exserted at anthesis. Fruit unknown. In leaf shape Mezilaurus caatingae resembles M. itauba and M. decurrens. The sessile flowers separate it immediately from these species, how- ever. Vegetatively, the recurved leaf margin is diagnostic. This character also occurs in M. mi- crantha, but this species has a bluntly acute leaf apex, never rounded as in M. caatingae. Mezilaurus crassiramea (Meissner) Taubert ex Mez, Arbeiten Kónigl. Bot. Gart. Breslau 1: 112. 1892. Oreodaphne crassiramea Meis- sner, DC. Prodr. 15(1): 117. 1864. Silvia crassiramea (Meissner) Mez, Jahrb. Kónigl. Bot. Gart. Berlin 5: 106. 1889. Mezia cras- siramea (Meissner) Kuntze, Revis. G 2: 574 TYPE: Brazil. Goias: Sects d’Ourada, Pohl 1463 (W, K, G-DC, U—this the only specimen seen). Figures 1, C & D; 2. Small trees, to 6 m tall. Twigs terete, thick, the older ones with a thick and conspicuous cork layer, the young tips with a dense, light brown tomentum. Leaves clustered at the tips of the twigs, almost sessile (petioles to 3 mm long), ovate or slightly obovate, the tip rounded, the base 160 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Figure 1. A-B. Mezilaurus Ap in — A. Leaf. — B. Flower. C-D. M. crassiramea. —C. Leaf. — D. Flower. E-F. M. decurrens. — E. Leaf. — F. Flower. G-H. M. itauba. — G. Leaf. — H. Flow rounded or gradually narrowed and abruptly rounded, with margins frequently recurved, 10 x 5 cm, the upper surface pubescent, but becoming glabrous with age, the lower surface rather dense- ly pubescent with pale brown hairs; lateral veins 10-15 pairs, leaving the midvein under almost 90*: veins and the final reticulation immersed on upper surface, raised on lower surface. Inflores- 1987] WERFF—REVISION OF MEZILAURUS O 100 200 300 400 500 600 miles 80 70 60 50 40 FIGURE 2. Distribution of Mezilaurus caatingae (), M. crassiramea (8), M. decurrens (A), and M. duckei cences in the axils of small bracts, seemingly ter- minal, 7-12 cm long, about as long as leaves, pedicellate, puberulous or with very short pu- bescence, the fl ged spicatel lateral branchlets; branchlets and flowers sub- tended by puberulous, ovate bracts, these ca. 0.1 mm long, the flowers brown puberulous, ca. 2 mm long; pedicel ca. 2 mm. Tepals 6, equal, ca. 0.8 mm long, erect, ovate. Stamens 3, exserted; anthers 2-celled, the cells small, on dorsal ridge of anthers, opening back-to-back; filaments fused into a tube. Ovary ovoid, ca. 1 mm, the style along the gradually narrowed, ca. 1 mm, exserted beyond anthers. Staminal glands and staminodia lacking. Young fruit globose, 6 mm diam., seated on swollen pedicel, the tepals persisting. Vernacular name. Cumbuquinha (fide Rat- ter). Additional specimens examined. BRAZIL. GOIÁS: Serra Dourada, Anderson 10003 (F, NY, MO, US). MATO GROSSO: 7 km SW of Xavantina, Ratter et al. 805 (MO); ca. 270 km N of Xavantina, Ratter 1293 (MO). 162 Mezilaurus crassiramea is a well-defined species known from a few collections in cerrado vegetation. Diagnostic characters are the thick corky twigs, the pubescent leaves, and puberu- lous flowers. It can, as many cerrado species, withstand fire; the Ratter collections come from trees with charred or fire-blackened trunks. Mezilaurus decurrens (Ducke) Kosterm., Med- ed. Bot. Mus. Herb. Rijks Univ. Utrecht 25: 40. 1936. Silvia decurrens Ducke, Trop. i 35 Yale No. 20999 (lectotype, RB, not seen; isolectotype F, fragm. U). Figures 1, E & F; 2 Large tree. Twigs minutely puberulous toward apex, the terminal bud with yellowish, appressed pubescence. Leaves clustered at the tips of branches, glabrous on both surfaces, elliptic or narrowly elliptic, 15-25 x 5-7.5 cm, the tip rounded, the base gradually decurrent onto the petiole, this 1-2 cm long; laminae coriaceous, opaque, the reticulation not raised, rather lax. Lateral veins not strongly developed, 7-12 pairs, immersed above, slightly raised below. Midrib thick, dark, raised above, more conspicuously so on lower surface. Inflorescences axillary, subter- minal, pyramidal, 10-18 cm long (fide Koster- mans), appressed pilose. Flowers subglobose or obconical, ca. 2-2.5 mm long, 2 mm wide, ap- pressed pubescent. Tepals 6, equal, erect, trian- gular, wider than long. Fertile stamens 3, 2-celled, 1-1.5 mm long, slightly exserted, pubescent; fil- aments connate, wider than anthers; anther cells extrorse, opening back-to-back. Ovary subglo- bose, densely pubescent (except base), ca. | mm long, the style slender and briefly exserted. Fruit unknown. Mezilaurus decurrens is rare and known to me with certainty only from the type collection. At first glance it appears quite similar to M. itauba, but differs in several subtle characters. The leaves of Mezilaurus decurrens do not have the raised reticulation and minute gland dots of M. itauba, and their leaf bases taper more gradually into the tiga Better differences are found in the flow- ers: in M. decurrens the anthers are scarcely ed and the filaments are connate, whereas in M. itauba the anthers are greatly exserted and the filaments free. A few collections that I place in Mezilaurus ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 itauba are close to M. decurrens in leaf outline (Ducke 681, Foldats 3613), but I attach more diagnostic value to the raised reticulation and the greatly exserted anthers Mezilaurus duckei van der Werff, sp. nov. TYPE: Brazil. Amazonas: Reserva Florestal Ducke, Aleusio 98 (holotype, US). Figures 2, 3. Arbor, 20 m alta. Ramuli crassi, cicatribus conspi- a : uper obscura, subtus pude Peti li crassi, ca. e pyram persistentibus et puberulis. Flon ores spicatim secus ra- mulos inflorescentiae dispositi, frequenter reflexi, tur- binati, ca. ] mm vel pubescentia laxa ad basim. Tepal — ca. 0.3 mm longa. Stamina fertilia 3, ca. m longa, filamentis pubescentibus, liberis, latioribus eae antheris glabris 2-locellatisque. Staminodia 6, parva, ca. 0.2 mm longa. Ovarium conicum, glabrum ca. 0.4 mm longum, stylo ca. 0.7 mm longo. Fructus ignoti. Tree, 20 m. Twigs thick, 1 cm diam. 5 cm below the tip, with conspicuous leaf scars, the tip very finely brown tomentellous. Leaves clus- tered at the tips of branches, slightly hien 25-30 x 8-10 cm, the tip rounded, gradua narrowed toward the base, the base abruptly nar- rowed; upper surface glabrous, opaque, venation immersed, the tertiary venation scarcely visible; lower surface minutely puberulous when young, secondary and tertiary venation raised, the mid- rib strongly raised and thick; petioles 0.5 cm thick, cences axillary, ca. branched, the branchlets 3-4 cm long, the u ones slightly shorter than the lower ones, mi- nutely brown puberulous. Bracts and bractlets pilose, persisting at anthesis, the bracts 1.5 mm long, bractlets ca. 0.7 mm long. Flowers pedi- cellate, pedicels ca. 2 mm long, glabrous. Flowers glabrous, cup-shaped, | mm long, tepals 6, small, 0.3 mm long, erect. Fertile stamens 3, ca. 0.5 mm long, the filaments free, pubescent, wider than the glabrous, 2-celled anthers. Staminodia , ca. 0.2 mm long. Ovary conical, glabrous, ca. 0.4 mm long, the style ca. 0.7 mm long. Fruit unknown. 1987] WERFF—REVISION OF MEZILAURUS FIGURE 3. Mezilaurus duckei. — A. Habit. — B. Flower. 163 164 Common name. Itauba abacate. Paratype. BRAZIL. AMAZONAS: Reserva Florestal Ducke, tree nr. 116, Rodrigues 8203 (NY). This species is named after Adolpho Ducke, an outstanding botanist and collector who made numerous excellent collections of Lauraceae in Amazonian Brazil. It is fitting that this new species is only known from the forest reserve dedicated to Ducke Menzilaurus glaucophylla van der Werff, sp. nov. TYPE: Costa Rica. Prov. San José: Zapatón de Puriscal, tree, 9 m, Zamora & Poveda 1014 (holotype, MO; isotypes, F, CR). Fig- ure 4 r, 20 m. Ramuli teretes, cicatribus insignibus viter €—— vel triangulares, cinereo-pubescentes, . Folia ad apices ramulorum conferta, obovata, 2. subtus glauca, apicibus basibusque illares, pyramida atae, cinereo-strigosae. Pedicelli ad 1.5 mm longi, glabri vel basim aliquot pilis. Flores glabri, il ees in sicco ca. 1 mm longi et lati. Tepala 6, per anthesin erecta vel paullo i incurvata; inaequalia, 3 exteriora interioribus breviora, late deltoidea. Sta- mina 3, 4-locellata, filamentis latitudine antheris ae- quantibus, ca. 1 mm longa, ca. 0.7 mm lata, tepalis exterioribus opposita. Staminodia 3, ca. 0.5 mm longa, li ind a tepalis interioribus opposita. Tu- bus flor vadosus, intus glaber. Ovarium glabrum, silipscideum. sensim in stylo attenuatum, ovarium stylusque ca. | mm longus. Fructus ignotus. Tree, to 20 m tall. Twigs terete, with conspic- uous leaf scars, minutely pubescent, the terminal bud white appressed pubescent; bark on older twigs soft and flaking. Leaves clustered at the tips of the branches, alternate, obovate, charta- ceous, glaucous and laxly and minutely puber- ulous below, the tip rounded or acute, the base obtuse or acute, 15-25 x 8-12 cm, the midrib and lateral veins immersed, but the tertiary ve- nation slightly raised above, glabrous except for the puberu midrib; midrib and lateral vein rescences axillary, pyramidate, to 15 cm long, gray strigose. Pedicels ca. 1.5 mm long, glabrous ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 or with few gray, appressed hairs at the base, subtended by 2 small, deltoid, strigose bracts, ca. 0.2 mm long. Flowers glabrous, more or less cup- shaped, ca. | mm long and wide when dry. Tepals 6, at anthesis more or less erect, the outer three smaller than the inner three, broadly deltoid. Sta- mens 3, all 4-celled, the filaments as wide as the anthers; glabrous or with a few hairs at the base, ca. 1 mm long, 0.7 mm wide, situated opposite the outer tepals; tips of the stamens curved in- ward; anther cells positioned on the upper part. Inner tepals pushed apart at anthesis and expos- ing anther cells in their sinuses. Staminodia 3, alternating with stamens, lanceolate, strigose, ca. 0.5 mm long. Ovary glabrous, ellipsoid, gradu- ally narrowed into style, ovary and style ca. 1 mm long. Floral tube shallow, glabrous inside. Fruit unknown. types. CosrA RICA. PUNTARENAS: Osa Penin- E in E t W. of Rincón, Hammel et al. 15214 (MO; duplicates to be distributed). There is no doubt that Mezilaurus glauco- phylla and M. quadrilocellata are closely related. They differ from the other Mezilaurus species in the following characters: distribution (the only species north of the Andes); presence of four- celled anthers; leaves more or less glaucous be- low; unequal tepals; and the inflorescence not strictly a dibothryum, but sometimes with short tertiary axes. These two species could be regard- ed as forming a new genus, based on their four- celled stamens and unequal tepals, but it should be mentioned that several other Mezilaurus species have small tepals that one cannot very well judge to be equal or not. If additional dif- ferences separating M. glaucophylla and M. quadrilocellata from the other Mezilaurus species are found (in cupule shape, for instance), it might be better to treat them as a separate genus, but for the time being, I prefer to include them in Mezilaur In addition to the THeteneps mentioned in Table 1, the few a that Mezilaurus elaucophylla has larger, "innt leaves and larger inflorescences than M. quad- rilocellata. suggest Mezilaurus itauba (Meissner) Taubert ex Mez, Arbeiten Kónigl. Bot. Gart. Breslau 1: 112. 1892. Acrodiclidium itauba Meissner, DC. Prodr. 15(1): 86. 1864. Endiandra itauba (Meissner) Benth. & Hook., Gen. Pl. 3: 154. 1880. Silvia itauba (Meissner) Pax, Natür- lichen Pflanzenfamilien 3(2) 123. 1889. 1987] WERFF—REVISION OF MEZILAURUS 165 FicunE 4. Mezilaurus glaucophylla. — A. Habit. — B. Flower and bud seen from above. Mezia itauba (Meissner) Kuntze, Rev. Gen. Acrodiclidium itauba Meissner var. amarella Meissner, à DC. Prodr. 15(1): 86. 1864. Type: Brazil. Para: Š " : A. PE: Pl. 2: 574. 1891. TYPE: Brazil. Para: Santa- Santarem, Spruce 646 Qectotype not chosen, Dij; rem, Spruce 643 (lectotype, K, fide Koster- i mans, BM, C, fragm. F). Figures 1, G& H; 5. Oreodaphne hookeriana Meissner, DC. Prodr. 15(1): 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 1. Comparison of Mezilaurus glaucophylla with M. quadrilocellata. M. glaucophylla M. quadrilocellata Inflorescence gray strigose rufous tomentellous Pedicels few basal hairs, otherwise glabrous rufous tomentellous Tertiary venation on upper leaf surface Terminal bud aise densely gray pubescent immerse brown tomentellous 131. 1864. Type: Brazil. Para: Santarem, Spruce 669 (not seen. ezilaurus anacardioides deri Taubert ex Mez, Arb. Bot. Garten Breslau 1: qas Acrodi- clidium anacardioides Meissner, m r. 15(1): a kyi Misanteca anacardioides eee . & Hook., Gen. Pl. 3(2): 155. 1880. Silvia d (Meissner) Mez, Jahrb. Bot. Gart. Berlin 5: 108. 1889. Mezia anacardioides (Meis- sner) Kuntze, Revis. Gen. Pl. 2: 574. 1891. TYPE: Venezuela. Amazonas: San Carlos de Río Negro, Spruce 2961 (lectotype not chosen, BM, G). Silvia polyantha Mez, Bull. Herb. Boiss. 2 sér. V: 233. . TYPE: Brazil. Amazonas: Moura, Rio Ne- gro, Ule 6055 (holotype, B, not seen, isotype, G). Silvia rondonii Mez et Hoehne, Bot. Archiv VI: 230. 4. TYPE: Brazil. Mato Grosso: near Tres Bu- ritys, Kuhlmann 1976 (K, not seen). Large trees to 35 m tall, rarely shrubs. Twigs terete, glabrous or nearly so, the terminal bud appressed pubescent, the bark rather thin and aking. Leaves clustered at the tips of branches, firmly chartaceous or coriaceous, glabrous at ma- turity, elliptic or slightly obovate, ca. 15 x 5 cm, the base gradually narrowed into petioles, the tip rounded, the reticulation on both surfaces slight- ly raised, the midrib and lateral veins (7-12 pairs) immersed above, raised on lower surface, the lateral veins arching upward and fading near the margin. Lower leaf surface densely and minutely gland dotted; petioles glabrous, with swollen bas- es, 1-2.5 cm long. Inflorescences axillary, sub- terminal, narrowly pyramidate, 5-10 cm long, laxly appressed pubescent. Flowers subumbel- lately arranged at tips of lateral branchlets, laxly and minutely appressed pubescent. Pedicels mi- nutely appressed pubescent, 2-4 mm long. Bracts and bractlets deciduous. Flowers hemispherical, 1.5-2 mm long, the 6 equal tepals erect (rarely spreading), wider than long, the anthers exserted. Fertile stamens 3, 2-celled, ovate-elliptical, ca. the cells large, opening back-to-back. Ovary el- lipsoid, pubescent, the style exserted. Flower tube pubescent. Staminal glands and staminodia lack- ing. Fruit an ellipsoid berry, ca. 2 x 1 cm, sub- tended by a small, platelike cupule. Selected additional specimens examined. SURINAM. Boschreservaat, sectio O, tree No. 760, Boschwezen 3088 (NY). BRAziL. PARÁ: Rio Tapajos, Villa Braga, Ducke RB 17537 (G, US). PARÁ: Serra dos Carajas, M. G. Silva 2909 (MO); Rio Jari, Monte Dourado, N. T. Silva 1041 (NY). PERU. MADRE DE DIOS: Tahuamanü, Diaz 17 53-96 (MO). Borivia: San Francisco, 50 km from Pto. Rica Pando, E. Menesk 626 (MO) Mezilaurus itauba is the most frequently col- lected and widest ranging species of the genus. In addition to the countries listed, it has been reported from French Guiana, based on a Mé- linon collection I have not seen. Although there s some variation in degree of pubescence and leaf shape throughout its range, M. itauba is ad- equately characterized by free staminal fila- ments, pubescent flowers, and glabrous leaves rounded at the tip and gradually narrowed ba- sally. The anthers, with large cells, are also long exserted for their size. The numerous gland dots on the lower leaf surface are best seen on dena young leaves; on mature, more coriaceous ve they are often scarcely visible. Alencar 55 (MO) is included in M. itauba with hesitation; it differs somewhat in leaf shape, but in the absence of floral differences I regard it as M. itauba. However, when more collections are sci it might turn out to be a new species. ood of Mezilaurus itauba is hard and much pie for construction. Mez (1889) reported that the berries are edi Mezilaurus lindaviana Schwacke et Mez, Arb. B arten Breslau 1: 112. 1892. TYPE: Bra- zil. Amazonas: Rio Branco, Schwacke 7080 (lectotype, chosen by Kostermans, B, not seen). Figures 6, A & B; 7 Misanteca duckei Samp., Commissao Linhas Telegr. Estrat. Matto Grosso Amazonas, Publ. 56 (An- f 1917. Silvia duckei M wurdackiana C. K. Allen, Mem. New York rden 10: 56. 1963. TYPE: Venezuela. Bo- a Hato La Vergaretia, Wurdack & Guppy 91 (holotype, NY, isotype, US). 1987] WERFF—REVISION OF MEZILAURUS 167 O 200 400 600 800 1000km L ee eee —— === O 100 200 300 400 500 600 miles 30 70 60 FIGURE 5. Distribution of Mezilaurus itauba. Tree, to 25 m. Twigs thick, round, witha rather thick corky layer, the tips with brown velutinous pubescence. Leaves alternate, clustered at the tips of the branches, chartaceous or coriaceous, ob- ovate, 8-17 x 4-8 cm, the tip rounded or very shortly acute, the base cuneate or abruptly rounded; young leaves hirsute, becoming gla- brous above (except on midrib and primary veins) at maturity; venation impressed above, the mid- rib, secondary and tertiary venation raised be- low; secondary veins about 10 pairs; petioles short, thick, brown-tomentose, 5-8 mm long. In- 50 40 florescences axillary, mostly subterminal on branches, pyramidal, tomentellous, about as long as the leaves; branchlets patent, 1-3 cm long, the flowers spicately arranged and often somewhat recurved. Flowers white or yellow-green, fra- grant, glabrous, 1.2 mm long. Tepals 6, equal, triangular, ca. 0.2 mm long, 0.4 mm wide. Fertile stamens 3, 1 mm long; filaments (ca. 0.5 mm connate, narrower than anthers, pubescent; an- thers shortly exserted, 2-celled, 0.5 mm long, the cells situated on a dorsal ridge, opening back-to- ack. Ovary ellipsoid, 0.8 mm long, with slender 168 FIGURE 6. F. M. micrantha. —E. Leaf.— 1 E i +h +h bre | mens t basal glands. Staminodia lacking. Fruit ellipsoid, 2 x 1.2 cm, subtended by a small platelike cupule (ca. 3 mm diam.). QC] "TT | Pe LPT ANA: Kanuku Mountains, For. Dept. Brit. Vies 5804 (NY), same inpet Smith 3208 (F. MO, US). BRAZIL. NAS: Rio ids nco, Boa Vista, Ducke 1336 (F, NY, US) A ing of Macapa, Fróes & Black 27453 (NY). es Pinsha de Santarem, Fróes 30976 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 3cm A-B. “yayawa ee —A. Leaf.—B. Flower. C-D. M. mahuba. —C. Leaf. — D. Flower. E- (NY, US); Rio Jari, Monte Dourado, E. Oliveira 4749 (NY), same location, N. T. Silva 996 (NY, US). Mezilaurus lindaviana is somewhat variable in leaf shape but is clearly characterized by the combination of pubescent leaves and glabrous flowers. Mezilaurus crassiramea, a similar species with pubescent leaves, has pubescent flowers and is known only as a shrub or a small tree in cerrado vegetation. 1987] WERFF—REVISION OF MEZILAURUS 169 200 400 600 800 1000km eS es eee eee eee | O 100 200 300 400 500 600 miles 80 70 40 FIGURE 7. Distribution of Mezilaurus lindaviana (6), M. mahuba (A), and M. micrantha (gp. Silva 2403 (NY, 2 sheets, MO) is included in M. lindaviana as an aberrant collection; it differs from other collections of that species in having sparse appressed pubescence. Other characters (leaf shape, size, flowers) point to M. /indaviana and I feel that a single collection with unusual pubescence need not be given taxonomic status. Kostermans (1938) cited as type of M. linda- viana Schwacke 7080 — Glaziou 19798 and gave Serra d'Antonio Pereira in Minas Gerais as the type locality. Schwacke and Mez (Mez, 1892) cited only Schwacke 7080 as type collection and gave as type locality “in campis ad Rio Branco." It is likely that Glaziou distributed duplicates of the Schwacke collection under his own name with incorrect locality data, as he did with other col- lections (Wurdack, 1970). Therefore, I ignore the reference of M. lindaviana as occurring in Minas Gerais, as cited by Glaziou (1905-1913) and Kostermans (1938). Mezilaurus mahuba (Samp.) van der Werff, comb. nov. Basionym: Acrodiclidium mahuba Samp., Commissao Linhas Telegr. Estrat. 170 Matto Grosso Amazonas, Publ. 56 (Annexo 5, Bot. Parte X): 14. 1917. Clinostemon ma- huba (Samp.) Kuhlm. & Samp., Bol. Mus. Nac. Rio de Jan. 4(2): 57. 1928. Licaria ma- T (Samp.) Lundell, Wrightia 4: TYPE: Brazil. Para: Gurupa, Varzea do Rio Amazonas, Ducke MG 16538 = RB 17582 (isotype, U). Figures 6, C & D; 7. Large trees. Twigs thick, glabrescent, with dense brown tomentum when young. Leaves large, 20- 40 x 12-15 cm, obovate, clustered at the tips of branches, glabrous above with the exception of the puberulous midrib, softly pubescent below, the apex rounded, the base cuneate; venation immersed on upper surface; midrib, lateral veins (15-23 pairs) and tertiary venation raised below; petioles thick, 5 mm diam., 2-3 cm long, densely and minutely tomentose. Inflorescences subter- minal, densely puberulous, large (20-35 cm long), minute, incurved. Fertile stamens 3, all 2-celled; filaments densely strigose, with 2 glands attached a little above the base; anthers glabrous, strongly curved outside the flower tube. Staminodes 9, ca. 0.5 mm long, lanceolate, strigose. Ovary gla- brous, globose, ca. 0.5 mm long, the style ca. 1 mm long, curved at the tip. Fruit ellipsoid, 3.5 cm long, seated on a small, glabrous disk. . — examined. BRAZIL. AMAPA: o Juruxi-Mazagao, B. V. Rabelo 2715 (MO). PARA: is Ducke RB 1 7583 (U, US), Ducke 1234 (NY, MO, US); Trapiche Hypolito, Krukoff 5870 (BR, NY, MO); Ilha de Pará, Mori et al. 16510 (MO), Belem, Murça Pires 1488 (NY). AL 7 L 2 4° of its peculiar anthers. When naq it is espi similar to M. thoroflora. The latter, however, has shorter petioles and a fine and closely appressed pubescence on young twigs and leaves, whereas in M. mahuba the petioles are longer and the pubescence consists of spreading hairs. Mezilau- rus mahuba is a species known only from sea- sonally inundated forest in the states of Pará and Amapa pa. The few available collections of Mezilaurus mahuba clearly show two phases in the floral development, probably corresponding with the male and female phase as described by Kubitzki and Kurz (1984) for several other species of Lau- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 raceae. The three sheets of Krukoff 5870 all have flowers with the anthers recurved and tightly pressed against the flower, almost hiding the an- ther cells (female phase), while the three sheets of Ducke 1234 have only flowers with the anthers spreading and free of the flower, fully exposing the anther cells (male phase). The inflorescences of these specimens are large, and it is interesting that the flowers on an inflorescence all appeared to be in the same phase of development, quite unlike what I have seen in species of Ocotea and Nectandra with large inflorescences. Mezilaurus micrantha van der Werff, sp. nov. TYPE: Brazil. Amazonas: Manaos, Reserva Florestal Ducke, Rodrigues & Coelho 7555 (holotype, NY). Figures 6, E & F; 7. Arbor, 20 m alta. Ramuli teretes, glabri vel prope mm atra. Venat , leviter e d esposa triangularia, er 0.6 mm longa, per anthe liberis, den ibus. Antherae 2- locellatae, lo- cellis extrorsis, tae ventrali staminis omnino pubes- centi. Ovarium glabrum, sensim in stylo attenuatum, ovarium stylusque | mm longus. Fructus ignoti. Tree, 20 m tall. Twigs terete, glabrous or, near the apex, with some appressed pubescence, the terminal bud sericeous; the bark rather thick. Leaves clustered at the tips of the branches, co- riaceous, drying blackish, glabrous at maturity, ut when young with some appressed pubes- cence, elliptic, ca. 10-15 x 3.5-5 cm (exclusive of petiole), the base gradually narrowed atten- uately into the petiole, the tip blunt but not rounded, the margins revolute; veins and retic- ulation not or scarcely raised on upper surface; midvein and main lateral veins (5-8 pairs) raised on lower surface, but reticulation not obvious. Petioles to 2 cm long, glabrous at maturity. In- florescences axillary, subterminal, to 5 cm long, pyramidate, with some appressed pubescence. Flowers arranged subumbellately at the ends of the lateral branches, appressed pubescent. Ped- 1987] icels minutely appressed pubescent, ca. 2 mm long at anthesis. Bracts and bractlets deciduous. equal, triangular, erect. Stamens 3, 0.6 mm long, exserted 0.3 mm at anthesis, the filaments free, densely put ; anthers 2-celled, glabrous, the cells extrorse, opening back-to-back; ventral side of the anther entirely pubescent. Floral tube pu- bescent. St staminodia lacking Ovary glabrous, gradually narrowed into style, the ovary and style ca. 1 mm long, the style exserted at anthesis. Fruit unknown. Paratypes. marge AMAZONAS: Manaos, Reserva Floresta Ducke, W. Rodrigues 3190 (NY); AM-1, Km , W. Rodrigues 7066 (NY). Mezilaurus micrantha is rather similar to M. itauba; it differs in having smaller flowers, blunt but not rounded leaf tips, lack of gland dots on the leaves, slightly revolute leaf margins, and nearly black dried leaves. These characters are not strong individually, but taken together they allow identification of flowering as well as sterile collections. The flowers of this species are among the smallest I have seen in the genus, hence its specific epithet. Mezilaurus navalium (Allemáo) Taubert ex Mez, Arbeiten Kónigl. Bot. Gart. Breslau 1: 112. 1892. Silvia navalium Allemáo, Dissertatio, Rio de Janeiro. 1848. Silvaea navalium (Al- lemao) Meissner, DC. Prodr. 15(1): 84. 1864. Endiandra navalium (Allemao) Benth. & Hook., Gen. Pl. 3: 154. 1880. Mezia nava- lium (Allemáo) Kuntze, Revis. Gen. Pl. 2: 574. 1891. TYPE: Brazil, Rio de Janeiro, A/- lemáo, s.n. (holotype, R, not seen). Figures B, 9 Tall trees, to 25 m. Branches terete, glabrous, the tips with appressed, short hairs, the terminal bud densely gray strigose. Leaves clustered at branch tips, narrowly elliptic, 10-12 x 3-3.5 cm, somewhat coriaceous, glabrous on both surfaces or with few appressed hairs along the midrib, the tip rounded, the base sharply acute, lateral veins not strongly developed, 10-15 pairs, the upper surface dull, smooth, lower surface with slightly elevated reticulation; petioles ca. 1 cm long. In- florescences axillary, glabrous or with few scat- tered hairs, 3-5 cm long, the flowers clustered at the ends of the lateral branches. Flowers gla- brous, ca. 2 mm long. Pedicels 2-3 mm long, glabrous. Braots deciduous. Tepals 6, equal, erect, WERFF—REVISION OF MEZILAURUS 171 scalelike, ca. 0.3 mm long. Stamens 3, ca. 1.5 mm long; filaments free, strigose; anthers exsert- ed, glabrous, the 2 large cells positioned on a dorsal ridge, opening back-to-back, slightly di- vergent, exposing the exserted stigma. Ovary el- lipsoid, glabrous, ca. 2 mm long, including the exserted stigma. Staminal glands and staminodia lacking. Immature fruits subtended by the small tepals, occasionally the stamens visible at the base of the young fruit. Additional specimens examined. BRAZIL. RIO DE JANEIRO: Theresopolis, Glaziou 11470 (C, G, U), Gla- ziou 11473 (C); Petropolis, Glaziou 12124 (C, G), Gla- ziou 12125 (C, G, MO, U, US). Mezilaurus navalium is the only Mezilaurus species known from the Atlantic rain forests in southern Brazil. The wood is hard and much used for naval construction. Diagnostic charac- ters, in addition to its distribution, are the gla- brous flowers and leaves with rounded or blunt apices. Mezilaurus opaca Kubitzki & van der Werff, sp. nov. TYPE: Peru. Depto. Loreto: Prov. Re- quena, Distr. Jenaro Herrera, trocha al Rio Yaveri, cerca al Arboretum de Jenaro Her- rera, 15 m tree in low forest, flowers greenish yellow, 20 Aug. 1976, Revilla 1226 (holo- type, MO; isotype, HBG). Figures 8, C & D; 12 0 m. Ramuli glabri, sed apicibus foliiferis e tertia immersa; aequalia, parva, ca. 0. x mm, minute va 2-loculatis. Ovarium glabrum, ovoideum, sensim in stylo attenuatum. Fructus ignoti. Tree, 15 m. Twigs glabrous, rather thick (5-7 mm diam. immediately below the leaves), with conspicuous leaf scars, the bark gray. Tips o branches with dense, brown, sericeous pubes- cence. Leaves alternate, clustered at tips of branches, ca. 20 x 9 cm, young ones with some appressed pubescence, glabrous at maturity, chartaceous, elliptic, the tip rounded or bluntly 172 Ficure 8. A-B. Mezilaurus navalium. — A. Leaf. — B. M. palcazuensis. — E. Leaf. — F. Flower. acute, the base gradually narrowed onto the pet- iole, green, opaque above, the midvein elevated, the secondary veins slightly elevated, the tertiary venation more or less immersed, not easily vis- ible; lower surface also opaque but with the ve- nation more elevated; petioles 3—4 cm long, with narrow wings of the decurrent laminae. Inflores- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Flower. C-D. M. opaca. —C. Habit. — D. Flower. E-F. cences compound racemes in the axils of decid- uous bracts, slightly longer than the petioles, when young rather densely appressed pubescent, at an- thesis less so. Bracts and bracteoles deciduous. Flowers clustered at the tips of inflorescence branchlets, with some appressed pubescence, de- pressed globose, constricted at the apex, ca. 1.5 x 1987] WERFF—REVISION OF MEZILAURUS 173 — 200 400 600 800 1000km r—r——— sh O 100 200 300 400 500 600 miles 80 70 60 FIGURE 9. (4), and M. navalium (©) 1.5 mm (including the exserted stamens). Tepals 6, equal, pointing inwards, wider than long, ca. 0.4 mm wide, ca. 0.2 mm long, with some ap- pressed pubescence. Floral tube short, ca. 0.5 mm long, with a pubescent ring in the upper part. Stamens 3, 2-celled, ca. 1 mm long, the anthers glabrous, exserted at anthesis, the cells on a dor- sal ridge, the valves opening back-to-back; fila- into the style; style exserted beyond stamens; stigma a small plate. Fruit unknown. 50 40 Distribution of Mezilaurus palcazuensis (@), M. pyriflora (8), M. quadrilocellata (), M. sprucei O). Mezilaurus opaca is rather similar to M. syn- andra, but differs from that species by its smaller flowers and the leaf characters mentioned in the key. Béguin et al. (1985) reported M. synandra from the Jenaro Herrera Arboretum. It is pos- sible that this specimen represents M. opaca, but I have not seen it. Mezilaurus palcazuensis van der Werff, sp. nov. TYPE: Peru. Cerro de Pasco: Selva Central, Palcazu Valley, elev. 300-600 m, 7 Dec. 174 1984, Hartshorn, Quijano & Mateo 2691 (holotype, MO). Figures 8, E & F; 9. Arbor, 25 m. Ramuli teretes, glabri vel prope apicem adpresse pubescentes, cicaticibus foliarum conspicuis funds is. Gemma terminalis sericea. Folia chartacea, ad adulta glabra, juvenalia mid sse x iru vel anguste obovata, ap- ice acuta, basi at l ed cm. Costa ele- : i aud conspi- M forescentia ore niculata, 4cm lon cua. Peti oli ca. | cm F n ils sessilibus vel e sessilibus. Flores minuti aree s, obconi .l.5m n gi. Tepala tamina 3, mm longa, per anthesin exserta. Antherae 2- locellatae, gla- brae, extrorsae parvam pubescentem in superficie. ventrale. Ovarium globosum, glabrum. Fructus ignoti Tree, 25 m. Twigs terete, glabrous or with some appressed pubescence near tip, with conspicuous leaf scars. Terminal bud sericeous. Leaves char- taceous, clustered at the tips of branches, with some appressed pubescence when immature, gla- brous when mature, obovate or narrowly ob- ovate, the tip acute, the base gradually narrowed into the petiole, 10-15 x 3-4 cm; costa raised on both surfaces; secondary veins and reticula- tion immersed, not obvious; petioles ca. 1 cm long. Inflorescence axillary, 4 cm long, panicu- late, minutely appressed pubescent, the flowers clustered at the tips ofthe lateral branches, sessile or nearly so. Flowers minutely appressed pubes- cent, sessile, ca. 1.5 mm long. Tepals 6, erect, 0 ong. Fertile stamens 3, 2-celled, exsert- ed at anthesis; anthers extrorse, situated on a dorsal ridge, opening back-to-back; filaments united, glabrous. Staminal glands and stamino- dia lacking. Fruit unknown. Mezilaurus palcazuensis is only known from the holotype, consisting of a small twig with few leaves and one inflorescence. It is therefore quite likely that the description does not embrace the morphological variation of this species. Note- worthy features are the acute leaf tips and the sessile flowers, both unusual characters in the genus. The leaves are also thinner than in other species of Mezilaurus, but with only one speci- men available, it is not certain whether this is a distinguishing character. Similar sessile flowers occur also in M. caa- tingae, known from caatinga forest along the Rio Negro, which differs in having larger flowers and coriaceous, rounded leaves with inrolled mar- gins. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Mezilaurus pyriflora van der Werff, sp. nov. TYPE: Brazil. Amazonas: Sao Paulo de Olivenga, basin of creek Belem, 26 Oct.-11 Dec. 1936, Krukoff 8711 (holotype, NY; isotype, MO, GH). Figures 10, A& B; 9 5 m alta. Ramuli crassi, ad 1 cm diametro, et subtus eleva per im- 5 ° e = 2 p e. "S e lateralibus. Inflorescentia axillaris, subterminalis, an- guste pyramidalis, ramulis basalibus perlongioribus quam terminalibus, adpresse pubescens. Flores parvi mm x 0.9 mm), glabri, pyriformes. Pedicelli pu- bescentes, 1.5-2 mm longi. Stamina fertilia 3, inclusa, 0.8 mm i celli pa Ovarium depresse globosum, glabrum, 1.2 m ongum. Ped a pA Glandulae filamentorum nullae. Fructus igno Tree, to 25 m tall. Twigs thick (diam. 1 cm 4- 5 cm below tip), terete, with conspicuous leaf scars, the tip with gray, minute and appressed pubescence, becoming glabrous with age. Ter- minal bud densely gray pubescent. Leaves clus- tered at tips of branches, firmly chartaceous, ob- ovate, the basal half of the lamina widening very gradually, the apical half rather abruptly wid- ened, large (40-60 x 14-18 cm at maturity), the tips acute, gradually narrowed towards the base, but the base abruptly narrowed, rounded or al- most subcordate, mostly glabrous, with some ap- venation scarcely visible above, raised below; petioles ca. 1 cm long, 5 mm thick, densely ap- pressed pubescent, the lamina decurrent as two narrow ridges. Inflorescences axillary, subter- minal, pyramidal, the basal branchlets much longer than the terminal branches (the longest ca. 10 cm long, decreasing to ca. 1 cm), the main axis and branchlets with appressed pubescence; bracts and bracteoles pubescent. Flowers ar- ranged spicately along branchlets, glabrous, often reflexed, pear-shaped, ca. 0.9 mm long, 0.9 mm wide. Pedicels pubescent, especially near the base, 1.5-2 mm long at anthesis. Fertile stamens 3, ca. 1987] WERFF— REVISION OF MEZILAURUS 175 FicuRE 10. A-B. Mezilaurus pyriflora. — A. Leaf. — B. Flower. C-D. M. quadrilocellata. — C. Leaf. — D. Flower. E-F. M. sprucei. —E. Leaf. — F. 0.8 mm long, included; filaments free, pubescent; Paratype. BRAZIL. AMAZONAS: Fróes 12152 (NY). anther cells minute, terminal, opening towards the tip. Ovary isis E glabrous, ca. Mezilaurus pyriflora is known only from two 1.2 mm wide, ca. ong. Staminodia 6, collections from the vicinity of São Paulo de Oli- representing the outer 6 um mens, ini ca. venga. Leaf shape, the short, pubescent pedicels, m long. Staminal glands lacking. Fruit un- the included, short anthers and the pear-shaped known. flowers (with the apical valves of the anthers 176 mimicking the dried crown of a pear) separate this species from M. mahuba and M. thoroflora. The Fróes collection in NY bears the annotation “wood causes injury to the skin.” The two collections of Mezilaurus pyriflora had been annotated as Euphorbiaceae and Ochna- ceae; because Krukoff's collections were widely distributed, it is possible that duplicates of M. ified in additional herbaria Lud Gd Mezilaurus quadrilocellata van der Werff, sp. nov. TYPE: Colombia. Antioquia: Chigorodó. Tree, 20 m. Flowers white. 100-200 m, M. Garcia Barriga 17626 (holotype, GH; iso- types, AAU, US). Figures 10, C & D; 9 Arbor, 20 m. Ramuli crassi, 4-5 mm diametro, te- retes, glabri apicibus foliiferis tomentellis. Folia con- ferta ad apices ramulorum, glabra, elliptica vel leviter obovata, chartacea, basi cuneata, apice rotundata, ca. 5x 8cm, petiolis 2-4 cm longis, tomentellis, margine cartilaginea, leviter incrassata. Venatio super immersa, subtus tomentella costa manifeste elevata, nervis la- teralibus (6—8) elevatis, nervis basalibus marginem at- tingentibus. Inflorescentiae axillares, foliis breviores, anguste pyramidatae, 6—9 cm longae, tomentellae. Flo- res parvi, m longi, pedicellis 2-3 mm longis, tomentellis. Tepala 6. 3 interioria 3 exterioribus duplo o ,c nga, interiora o m nga, omni abra. Stami longa, lamentis pubescentibus, sine glandu antheris bescentia. Ovarium ellipsoideum, glabru gum, sensim in stylum attenuatum. Fructus ignoti. Tree, 20 m tall. Twigs thick, 4-5 mm diam. immediately below the leaves, terete, glabrous except for the rufous tomentellous leaf-bearing apex. Leaves clustered at tips of branches, gla- brous, elliptic or slightly obovate, ca. 15 x 8 cm, the base cuneate, the apex rounded, the margins y thickened, the venation immersed above, the midrib prominently raised and tomentellous on lower surface, the lateral veins (6—8) less prominently raised, the basal ones reaching the margin, the upper ones arcuate and not reaching the margin; tertiary venation slight- ly raised, petioles 2-4 cm long, rufous tomen- tellous. Inflorescences axillary, shorter than the leaves, narrowly pyramidate, 6—9 cm long, to- mentellous. Flowers small, ca. 1.5 mm long; ped- icels 2-3 mm long, tomentellous; flower tube and tepals glabrous outside. Tepals 6, the outer ones half as long as the inner ones, erect, with tips curved inward; outer tepals ca. 0.6 mm long, ovate, inner ones ca. 1.2 mm long, ovate, gla- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 brous. Ovary ellipsoid, gradually narrowed into the style, 1 mm long, glabrous. Fertile anthers 3, ca. 1 mm long; filaments rather densely pubes- cent; anthers 4-celled, glabrous, the tips bent in- ward, forming a flat shield exposed at anthesis; anther cells situated on this shield. Fertile an- thers alternating with 3 small (ca. 0.5 mm), slen- der, densely pubescent staminodia. Fruit un- known Mezilaurus quadrilocellata is known only from the type collection in northern Colombia, not far from the Panamanian border and the Caribbean. Further discussion is given under M. glauco- phylla. Mezilaurus sprucei (Meissner) Taubert ex Mez, Arbeiten Kónigl. Bot. Gart. Breslau 1: 112. 1892. Acrodiclidium sprucei Meissner, DC. Prodr. 15(1): 86. 1864. Silvia sprucei (Meiss- ner) Mez, Jahrb. Kónigl. Bot. Gart. Berlin 5: 119. 1889. Mezia sprucei (Meissner) Kuntze, Revis. Gen. Pl. 2: 574. 1891. TYPE: Brazil. Amazonas: San Gabriel de Cacho- eira, Rio Negro, May 1852, Spruce 2323 (lectotype, chosen by Kostermans, K; iso- types BM, C, NY, U). Figures 10, E & F; 9. Mezilaurus maguireana C. K. Allen, Mem. New York Bot. Garden 10: 58. 1963. TYPE: Venezuela. Ama- zonas: Río Guainía, Maroa, Maguire et al. 41698 (holotype, NY; isotype, GH). Small tree, to 10 m tall. Twigs terete, glabrous, raceous, glabrous, elliptic, 12 x 5 ) cm, the base acute, the apex acute or acuminate; lat- eral veins 10-15 pairs, slightly elevated or im- mersed on upper face, raised on lower surface, the reticulation slightly raised on both surfaces; petioles to 3 cm long, flat, bordered by a narrow ridge, the very base of the petiole round and branchlets, ca. 2 mm long, the pedicels to 1.5 cm long. Tepals 6, equal, erect, ca. 0.5 mm lon Stamens 3, ca. 1.2 mm long, exserted; anthers free and somewhat divergent, 2-celled, the cells large, opening back-to-back; filaments pubes- cent, connate. Ovary ellipsoid, ca. 0.8 mm long, abruptly narrowed into the slender, ca. 1 mm 1987] long, style. Flower tube pubescent within. No staminal glands or staminodia. Fruits not seen. po Bate examined. VENEZUELA: 30 km f Puert achucho, Guanchez 255 (TFAV). AMAZONAS: ee Carlos de Río Negro, Clark & Ma- guirino 7784 (MO), 8091 (MO); Cerro Neblina base camp, Gentry & Stein 46836 (MO); BRAZIL. AMAZONAS: Río Negro above Camanaus, Prance et al. 16042 (NY). PERU. LORETO: Iquitos, near Picuruyacu, Revilla 106 (G, MO); Requena, Jenaro Herrera, Vasquez & Jara- millo 984 (MO). Mezilaurus sprucei can be recognized easily by its glabrous, acute or acuminate leaves, lax in- florescences, and especially by the flowers with long pedicels. The Revilla collection from Peru has short pedicels (5 mm long) but agrees in other characters with Mezilaurus sprucei. I tentatively place Mezilaurus maguireana in synonymy under M. sprucei. The type does not agree completely with the typical M. sprucei; the inflorescences are stiffer, the flowers have pedi- cels only 5 mm long, the reticulation is less raised on the upper leaf surface, and the leaf apices are less acuminate. These differences are only of de- gree and the few collecti of M. sprucei at hand probably do not show the i range of variation in the species. In what I consider important char- acters (acute leaves, long petioles, glabrous, sub- umbellately arranged flowers, and connate fila- ments), M. maguireana agrees with M. sprucei. I found that in old flowers the anthers become divergent, although the filaments remain con- nate. ee dodge (Ducke) Kosterm., Med- i . Rijks Univ. Utrecht 25: 40. 1936. Silvia ao Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 115. 1930. TYPE: Brazil. Pará: dry upland forest of Jumanda River, Ducke RB 19974 (holotype, RB; not seen; isotype, U). Figures 11, 12. Medium-sized tree to 20 m. Twigs terete, with conspicuous leaf scars, the terminal bud densely and minutely appressed pubescent, this disap- pearing rapidly as twig matures. Leaves clustered near apices of branches, rarely few leaves per- sisting on older twigs, the leaves glabrous, cori- aceous, slightly ovate or elliptic, ca. 15 x 7 cm, ioles 3-5 cm, rarely only 1 cm long. Inflorescences axillary near tips of twigs, pyram- idal, rather laxly flowered, appressed tomentel- WERFF—REVISION OF MEZILAURUS 177 lous, 5-12 cm long; branchlets slender, distant, to 1.5 cm long, the flowers clustered near their tips. Bracts and bracteoles deciduous. Pedicels slender, tomentellous, 1-1.5 mm long. Flowers globose, puberulous, ca. 1.5 mm long. Tepals 6, equal, erect, ca. 0.6 mm long. Stamens 3; ca. 1 mm long; filaments ca. 0.7 mm, pubescent, con- nate; the 2-celled anthers exserted like small horns from the flower tube, glabrous; anther cells large, lateral-introrse, opening toward the tip. Ovary vin ca. ] mm long; style exserted, the stig- nute. Staminal glands and staminodia T Fruit (fide Kostermans, 1938) ellipsoid, .5-3 cm long, 1.5 cm diam., subtended by a small, platelike cupule (4-5 mm diam.) with sub- persistent tepals. Additional specimens examined. PERU. MADRE DE DIOS: Tambopata, Gentry et al. 46116, 45952 (MO). Mezilaurus subcordata is collected rarely and known to me only from an isotype and two recent collections in Peru. Very possibly it is not a true disjunct—it may also occur in the intervening area, since a tree with green flowers 1.5 mm long can easily be overlooked. The Peruvian collec- tions come from a tree plot in which every tree was sampled, regardless of whether it was fertile or not. The description is based on the U isotype. The Peruvian specimens differ in having glabrous flowers and slightly thinner leaves. I find these differences too weak for recognition of a new taxon, but additional collections may show the Peruvian plants to be distinct. Mezilaurus subcordata can be recognized readily by its long petioles and rounded leaf bas- es; it is also the only species in the genus with lateral-introrse anther cells. Mezilaurus synandra (Mez) Kosterm., Meded. Manáos, dry upland forest near Pensados, Ule 8835 (lectotype, B, fide Kostermans, > seen; isolectotype, L). Figures 13, A& B Tree, to 15 m tall. Twigs terete, glabrous, often with conspicuous leaf scars, the terminal buds yellowish strigose-sericeous. Leaves clustered at tips of branches, glabrous, elliptic or slightly ob- ovate, 8-20 x 4-10 cm, the base cuneate or acute, the tip rounded, lateral veins 10-15 pairs, + im- mersed above, slightly raised below, the tertiary ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FIGURE 11. venation reticulate and slightly raised on both surfaces, the midvein raised below, triangular in diameter; petioles conspicuous, 3-5 cm long, gla- brous, flat or with 2 narrow ridges on upper side. Inflorescences subterminal, much shorter than leaves, ca. 3 cm long, the flowers in small (4—5 flowered) clusters at the ends of few lateral branchlets, minutely strigose. Bracts and bract- lets early deciduous, not seen. Flowers sparsely and minutely strigose, depressed globose, as wide as long or wider than long, the flower tube nar- rowed toward the tip (ca. 2 mm wide and 1.7 mm long on unpressed flowers). Tepals 6, equal, erect, scalelike, ca. 0.4 mm long, ca. 0.8 mm wide. Stamens 3, ca. 1.9 mm long, exserted, the Mezilaurus subcordata. — A. Habit. — B. Flower. anthers glabrous, 2-celled, the cells opening lat- eral-apically or apically; filaments connate, pu- bescent. Ovary pupescent ea. ES mm dong; ae ca. 0.3 mm long. St lacking. Fruit ovoid, ca. A cm long, 1 cm wide, subtended by a small, platelike cupule. Additional specimens examined. BRAZIL. AMAZONAS: anáos, Igarape da Cachoeira, Baixa do Tarumã, Cha- gas s.n. = MG 21.108 (NY); Manáos, Parque 10 de Novembre, Coelho s.n. = INPA 3934 (NY); Manáos, Pensador, Ducke 233 (F, NY), Ducke 233, second col- pe (US); Manáos, Ducke RB 23964 (G, US); Man- s, Pensador, Ducke RB 25092 (US); Manáos, Cach- oeira, Alta do Tarumã, Rodrigues & Lima 30 Y). Reported by Béguin et al. (1985) from the Arborétum Jenaro Herrera, Loreto, Peru 1987] WERFF—REVISION OF MEZILAURUS 179 o 200 400 600 800 1000km € ere | r———mcCàs1xa” | O 100 200 300 400 500 600 miles 80 70 60 50 40 FIGURE 12. Distribution of Mezilaurus opaca (), M. subcordata (W), M. synandra (B), and M. thoroflora (e). Mezilaurus synandra is only known from dry, Mezilaurus thoroflora van der Werff, nom. nov. low forest on terra firme, near Manaus and one Basionym: Licaria maguireana Allen, Bull. collection in Peru. Several collections indicate Torrey Bot. Club 75: 315. 1948. Misanteca that it occurs in secondary vegetation. When maguireana (Allen) Lundell, Wrightia 4: 100. flowering, M. synandra can be recognized easily 1969. Clinostemon maguireanum (Allen) by its short inflorescences and broad, depressed- Kurz, J. Arnold Arbor. 60: 520. 1979. TvPE: globose flowers, which otherwise occur only in Guyana: Mazaruni Station, Forestry Dept. M. opaca. Fruiting or sterile collections are rath- Brit. Guiana 2956 (F220) (holotype, NY; er similar to M. itauba, which has, however, a isotype, K). Figures 13, C & D; 12. larger inflorescence, generally smaller leaves, and the upper leaf surface dull, not shiny, with less Tree, ca. 25 m tall. Twigs thick, terete, the tip prominently raised reticulation. densely and minutely appressed pubescent, be- 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 3cm FIGURE 13. A— B. Mezilaurus synandra. — A. Leaf. — B. Flower. C-D. M. thoroflora. — C. Leaf. —D. Flower. coming glabrous with age. Terminal bud yellow- ish pubescent. Leaves clustered at the tips of branches, coriaceous, obovate, gradually nar- rowed toward base, the base abruptly rounded, the apex acute or shortly acuminate, 30-60 x 12-18 cm, pinnately veined, 15-25 pairs of lat- eral veins, these immersed above, but elevated below. Midrib thick, ca. 7 mm wide, raised on 1987] both surfaces. Tertiary venation not visible above, but raised below. Lamina glabrous above, except along midrib, sparsely appressed pubescent be- low, especially near the base and along midrib. Inflorescences in axils of aborted leaves, subter- minal, appressed gray-pubescent when young, with few hairs at anthesis, 30-50 cm long, lateral branchlets 2-3 cm long over the entire length of cylindrical inflorescence, these with scattered hairs, bracts and bractlets, strigose, persisting at anthesis. Flowers clustered toward the tips of Üranchlets, glabrous, ellipsoid, ca. 1.5 mm long, wide, the anthers ca. 0.5 mm exserted, pedicels 4-8 mm long. Tepals 6, equal, erect, scalelike. Fertile stamens 3, ca. 1 mm long, the filaments free, pubescent, ca. 0.5 mm long, the 2-celled anthers glabrous, exserted, the large cells bescent, the outer six slightly sagittate, the inner three lanceolate. Infructescence ca. 60 cm long, the fruit ellipsoid, ca. 1.7 x 1 cm, subtended by a thin platelike cupule, ca. 0.6 cm diam. Paratype. Guyana, Mazaruni Station, Forest Dept. Brit. Guiana 2704 (K). Mezilaurus thoroflora is known only from a few collections in Guyana. Diagnostic for this species are the long-pedicelled flowers, the large exserted anthers, and the uniform, short, lateral branchlets of the inflorescence. Vegetatively, M. roflora The illustration in Kubitzki et al. (1 979) of M. thoroflora (as Clinostemon maguireanum) is not representative of material I have seen. The two collections from the Forest Department of Brit- ish Guiana do not have such pronounced cordate leaf bases and do not have the lower branchlets of the inflorescences much longer than the upper ones, as shown in Kubitzki et al. (1979). Kostermans (1938) cited two collections (Monteiro Costa 323, Kaufmann 605, both in F) under M. lindaviana. In my opinion these spec- imens belong to M. thoroflora or are very close to it. They have considerably smaller leaves than the type of M. thoroflora, but leaf shape is quite similar. Monteiro Costa 323, a flowering speci- men, shows exserted anthers and staminodia, and this clearly indicates M. thoroflora. Pedicels are shorter and leaves smaller than in M. thoroflora, WERFF—REVISION OF MEZILAURUS 181 but it is likely that the few available collections do not show the full range of vegetative variation. Because the combination Mezilaurus magui- reana already exists, it was necessary to create a new epithet for Licaria maguireana. The epithet thoroflora is derived from the Crest "thoros," semen, and “‘flos,”’ flower, in reference to rembia between the small, long-pedi- celled flowers and spermatozoa. IMPERFECTLY KNOWN SPECIES Mezilaurus sp. A. A collection made by J. da Silva Costa (RB 1807 96) in the State Mato Grosso, Brazil, prob- ably represents an undescribed species. The lo- cality data suggest it was collected in cerrado vegetation as a 9 m tree. The most distinguishing characters are found in the leaves, which are densely gland-dotted on the upper surface, char- taceous, and have a few appressed hairs on the lower surface. The young flower buds are gla- brous. This is clearly not the other cerrado species, M. crassiramea, which has pubescent leaves, and the gland-dotted upper leaf surface has not been found in other Mezilaurus species. I prefer to wait with a formal description until flowering material is available. LITERATURE CITED ALLEMAO, F. 1848. Dissertatio, Rio de Janeiro. ALLEN, C. K. 1948. Lauraceae. In B. Maguire et al. Plant explorations in Guiana in 1944, chiefly to the Tafelberg and the Kaieteur plateau — III. Bull. 7 9 a e. In B. Maguire et al. The of the Guayana Highland, V. Mem. New uid) 44-1 . D. R. SPICHIGER & J. MiBGEe. 1985. Las 3: 146-164. London. Ducke, A. 1930. Plantes nouvelles ou peu connues de la Région Amazonienne IV. Arch. Jard. Bot. Rio de Janeiro 5: 101-188. 35. Notes on the itaüba trees: the Ama- zonian species of the genus Silvia Allem. Trop. Woods 42: 18-21. 1905-1913. Plantae Brasiliae centralis . The Genera of Flowering Plants. Clarendon Press, Oxfor KOosTERMANS, A. J. G. H. 1938. LO TR of the Lau- raceae III. Recueil Trav. Bot. Néerl. 35: 56-129. 182 ————. 1952. A historical survey of the Lauraceae III. ; Sci. Res. (Jakarta) 1: 141-159. Lauraceae. Reinwardtia 4: 192-256. a K. & H. Kurz Synchronized di- chogamy and dioecy in neotropical Lauraceae. Pl. Syst. Evol. 147: 253-266. . RICHTER. 1979. Reinstate- ment of Clinostemon (Lauraceae). J. Arnold Ar- bor. 60: 515-522. KUHLMANN, J. G. & A. DE SAMPAIO. 1928. Clino- stemon, Kuhlm. et A. Samp., n. gen. de Lauraceas da Amazonia. Bol. Mus. Nac. Rio de Janeiro 4: KUNTZE, O. 1891. Revisio Generum Plantarum II. Zi Kurz, H. 1983. 5 ue Gat- tungen neotropischer Laur und Revision der Gattung Licaria patei, "Thesis, Laives of Hambur; MEISSNER, C. F. 1864. Lauraceae. /n A. de Candolle, Prodromus Systematis Naturalis Regni Vegeta- bilis 15: 1-260. MEZ, C. 1889. Lauraceae Americanae. Jahrb. Kónigl. Bot. Gart. Berlin 5: 1-556. March ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 . 1892. Spicilegium Laureanum. Arbeiten Kónigl. Bot. Gart. Breslau 1: 71-166. . 1904. Additamenta monographica 1904. Bull. Herb. Boissier. sér 2. 5: 233-244. 9 Additamenta monographica 1919. Re- pert. Spec. Nov. Regni Veg. 16: 305-309. 1924. Additamenta monographica 1924. Bot. Arch. 6: 230-231. Pax, F. 1897. Die Natürlichen Pflanzenfamilien. Nachtrag und Register zu Teil II-IV. Anatomie des sekundáren Xy- lems und der Rinde der Lauraceae. Sonderbande des Naturw. Ver. Hamburg, 5, Verlag Paul Parey, Hamburg. TAUBERT, P. 1892. [Review of: Revisio Generum Plantarum by O. Kuntze.] Bot. Centralbl. 50: 17- 24. WEBERLING, F. 1981. Morphologie der Blüten und der EC sies Verlag Eugen Ulmer, Stuttgart. . 1985. r In foreszenzmorphologie der Lau- as rroneous X dahi in Glaziou collections of Vid Bean iniu] Taxon 19: 911- 913. Volume 73, No. 4, pp. 653-831 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN, was published on 30 1987. |. Portraits of Botanists —— The Missouri Botanical Guia has begun issuing a series of patos“ “Portraits Salas are reproduced from the Garden’s Archives, and brief information a CONTENTS Systematic Embryology of the Anisophylleaceae Hiroshi Tobe & Peter H. Raven _... 1 nosae: Papilionoideae) David A. Neill Australian Acacia Peter Bernhardt ... Flower Longevity and Protandry i in Two cae of Gentiana (Gentianaceae) C. J. Webb & Jan Littleton .. Notes on the Breeding XAR of Sacoila senweatia (Aublet) Garay (Orchidaceae) Paul M. Catling .... Flower and Fruit Sui in DES Spe Belarus Sinbad: - Javier Herrera w... U. T... U... Reproductive Systems and SOME SEN of Oxalis pes-caprae L. G Their en on the Genesis of a Noxious Weed Robert Ornduff Trapliners in the Trees: Hummingbird Pollination of Erythrina Sect. Erythrina (Legumi- £ A Comparison of the Diversity, Density, and F. oraging Behavior of Bees and Wasps on ` iy 51 j € 79. 85 pel : Flora of the Venezuelan Guayana—ll Julian A. Steyermark SE A New Spee of f Jatropha ere) from Nicaragua Grady gx. ges B. Faden & D. R. Hunt — »psis with XI eer ' (Commelinaceae) $ Webster i d 17 š 2 5e Chromosome Numbers B Madagascar Plants — Elisabeth R abakonandrianina & Gerald s p Gr. 1: ; Cytotaxonomic. Studies i in the Cems: Urginea Stein i in West Africa. IL Karyotype Evolution in Urginea altisima (L. ) Baker FO Oyewole ' Cytotaxonomic Studies i in the Genus Gris Stein in West Africa. HL The Casa of Urginea di | indica. (Roxb.) Kunth i in Nigeria. SN Io Oyewole ; Cytotaxonomic Studies i in the Genus Urginea Stein in West (Doe IV. P. entiation -and EAS Virin + in S an > uite une K no MA 126. To: opulation Differ- n. š Š 2 seene. Taia Warren Volume 74 Number 2 Volume 74, Number 2 Summer 1987 Annals of the Missouri Botanical Garden The Annals, published quarterly, contains papers, primarily in systematic botany, con- tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the Garden will also be accepted. Authors should write the Editor for information concerning arrangements for publishing in the ANNALS. Instructions to Authors are printed on the inside back cover of the last issue of each volume. Editorial Committee George K. Rogers Editor, Missouri Botanical Garden Janice Wilson Editorial Assistant, Missouri Botanical Garden Marshall R. Crosby Missouri Botanical Garden Gerrit Davidse Missouri Botanical Garden John D. Dwyer Missouri Botanical Garden & . Saint Louis University -Peter Goldblatt Missouri Botanical dnt. | Henk van der Werf Missouri panal Garden | For pI Saut contact Eleven, P.O. Box 299, St. Louis, MO 63166. Sub- ` scription price is $75 per volume U.S., $80 Canada and Mexico, $90 all other countries. Airmail deliv- ery one $35 | per volume. » Four i issues tpe vol- m | | The ANNALS OF THE MissoURI BOTANICAL GN x 7 (ISSN 0026-6493) is published quarterly by the . Missouri Botanical Garden, 2345 Tower Grove Av i enue, St. Louis, MO 63110. Second class postage i alioa Louis, Modum EE Er Box 299, s. Lois MO 63166 Volume 74 Number 2 1987 Annals of the Missouri Botanical Garden WZ VASCULAR EPIPHYTISM: TAXONOMIC PARTICIPATION AND ADAPTIVE DIVERSITY! DaviD H. BENZING? ABSTRACT Vascular epiphytes carci few qualities beyond occurrence in tree crowns that identify them as a those of terrestrials native to comparably arid or infertile sites. Moisture supply, m arily because their phylogenetic origins and life styles in forest canopies feature, determines where a particular type of epiphyte will survive. Epiphytes constitute about 10% orchids for epiphytism are identified. R families in tree crowns are less clear, but some possibilities are offered. relations and, Bm mineral nutrition may explain why epiphytism has rarely, if ever, preceded branch parasiti EPIPHYTISM: DEFINITION AND BREADTH Epiphytes are plants that spend much or all of their lives attached to other plants. Qualifying forms range from microbes to angiosperms; both aquatic and terrestrial vegetation provide me- chanical support. Interaction with phorophytes can range from incidental to physiologically in- timate; primarily rain-fed “atmospheric” bro- meliads and orchids (Figs. 3, 4) are anchored by a few roots (sometimes only one), whereas con- tact is almost complete for the largely endophytic dwarf mistletoes. (The designation “epiphyte” is here reserved for free-living vascular species; it does not include hemiparasites.) This report has two purposes: to describe taxonomic participa- tion in epiphytism and to explore the extraor- dinary proliferation in arboreal habitats by some families and higher taxa. First, epiphytes are ex- amined in terms of basic characteristics and im- portant biological distinctions. Plants considered to represent a specific eco- logical category usually share key qualities that ! The first four papers of this issue were presented as part of the 32nd Annual perd Symposium, The Biology of Epiphytes, held at the Missouri Botanical Garden 18-19 October 1985, an nd s 11392, Gerrit Davidse, Principal Investigator grant BSR 831 —— in part by nto the Missouri Botanical Garden. ? Oberlin College, Oberlin, Ohio 44074, U.S.A. ANN. Missouni Bor. GARD. 74: 183-204. 1987. ounda 184 set them apart from other vegetation. Occurrence on the same general type of substratum and, more importantly, utilization of comparable resource bases by similar mechanisms require consider- able, often conspicuous, convergence. All botan- ical carnivores possess traps designed to attract and process fauna for food; all lianas feature slender habit and novel vascular anatomy; all vernal ephemerals from temperate deciduous forests generate simplified shoots bearing helio- philic, short-lived foliage. In contrast, anchorage in tree crowns— sometimes even inability to sur- vive terrestrially—exhibited by so many of the approximately 25,000 epiphytic species appears to have little unifying basis. No growth form, seed type, identity of pollen vector, water/carbon balance regimen, source of nutrient ions, nor re- source procurement mode is shared by all me- chanically-dependent species. Moreover, char- acteristics of epiphytic and soil-based flora overlap broadly, as do important aspects of their habitats. Growing conditions for epiphytes, as indicated by rooting media, microclimate, and stability and dispersion of substrata, are diverse and similar to those for much terrestrial vegetation. Like soils, which range from equable to dry and infertile, canopies of everwet to microphyllous to cactus forests 1 Dose me bi severe stress on resident flora. Problem d dispersal in tree crowns are probably similar to those experienced by cliff dwellers and plants native to other pre- cipitous sites. A naked bark surface must com- plicate water and mineral balance just as do thin soils and rock faces. At the other extreme, how- ever, the everwet forest's rotting trunks or branches (Fig. 2), debris-filled knotholes, and ant nests (Fig. 7) probably offer to certain humi- philous epiphytes resources equal to those avail- able at humid, fertile, terrestrial sites. In effect, heterogeneity and similarity to other habitats complicate definition of the epiphytic biotope and thwart attempts to understand why tree crowns offer the only recourse to such a variety of vascular plants. From another vantage point, canopy-dependent species seem to be no more versatile than soil-rooted plants; most will sur- vive only under narrowly proscribed circum- stances (e.g., rigid confinement to twigs rather than larger axes, humus as opposed to “uncon- ditioned”’ bark, dark instead of better-illuminat- ed sites) Many taxonomies have been formulated to classify mechanical dependence. Criteria most ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 often used are extent of arboreal presence (fa- cultative vs. obligate forms), nature of depen- dency on supporting vegetation (mistletoes vs. epiphytes), exposure requirement (heliophiles vs. sciophytes), habit (e.g., tank forms, myrmeco- phytes), and type of substratum (e.g., humi- phobes, ant-nest forms). A further criterion rec- ognizes two types in a special group that normally taps soil part of the time—the hemiepiphytes. Primary hemiepiphytes, a group which includes stranglers, begin life on bark and later produce soil roots; secondary hemiepiphytes (Fig. 11) germinate on the ground and then become can- opy-dependent as older roots and basal stems of vining shoots decay. Features more directly re- flecting peculiarities of vegetative function as- sociated with epiphytic life (e.g., presence of ab- sorptive trichomes, carbon fixation pathway) have not generally been employed to distinguish epiphytic vegetation despite their utility for ex- plaining type of microsite and how existence in that kind of space is sustained. ne criterion above all others may prove meaningful to those concerned with the me- chanics of epiphytism: temporal access to mois- ture. While adequate mineral nutrition and ex- posure are no less critical than are suitable water relations to long-term survival of an epiphyte, avoiding drought injury is the more immediate challenge. Constant adjustment to rapidly chang- ing moisture supply must be accomplished via appropriate stomatal and photosynthetic re- sponses. There are just two types in this new taxonomy: continuously-supplied (CS) and pulse- supplied (PS) forms. Moisture supplies are steady for CS species tapping deep, absorbent media (Figs. 2, 5) or catchments created by the plant itself (Figs. 8, 9). The most self-contained mem- bers of the canopy-dependent flora (PS epi- phytes) draw water from relatively nonabsorbent substrata or other sources subject to quick drying (Figs. 3, 4). Epiphyte life history has been influenced by many selective forces, including substratum dis- tribution and stability. Patchiness in the epi- phytic biotope ranges from gross patterns due to hyperdispersion of tropical-forest trees to finer- grained discontinuity in the scattered arrays of suitable bark within individual hosting crowns. Mortality continues to be high during and fol- lowing seedling establishment. Disturbance is le- thal when supporting bark fragments exfoliate, inhabited twigs fall, infested trees collapse, an (less common but broader in extent) natural dis- 1987] BENZING— VASCULAR EPIPHYTISM 185 ed on. ing in a rotting bra zomes pearing traps of sik ales i ucl a tank brom ges NE id ot as, Venezuela.— artially dissected shoot of the bromeliad Brocchinia tatei on Cerro Neblina, Venezuela, mic.— 2. Young specimen of ing electron micrograph of the bi of Campylocentrum pachyrrhizum and two small bromeliads growing on a small branch in south Florida. asters ravage whole communities. Effective patch life (patch duration relative to the interval need- ed for a resident plant to reproduce) must be especially short for a multitude ofepiphytes whose fecundity is limited by drought and poor nutrient sources. It is this group that possesses an unusual combination of stress and r-selected characters (Benzing, 1978). In contrast, CS species, with their relatively regular resource supplies, should exhibit less emphasis on shortening the life cycle and channeling scarce commodities into repro- ductive rather than vegetative tissue. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 1987] ADAPTIVE VARIETY IN CANOPY DEPENDENTS Some adjustments to conditions in tree crowns are predictable and obvious— viz. capacities for aerial dispersal and holdfast — but others are more cryptic and varied. Depending on taxon and hab- itat, epiphytes exhibit various mechanisms for carbon fixation and procurement of moisture and nutrients. Essential elements are drawn from re- markably diverse sources, including several not ormally available to plants. A survey in some Indian forests (Sengupta et al., 1981) revealed greater nitrogenase activity in the phyllosphere of epiphytes than of terrestrials, including host- ing trees. Impoundments built of roots or shoots provide access to nutrients via intercepted can- opy fluids and litter (Figs. 8, 9) and the necessary detritivores and saprophytes these catchments attract (Benzing, 1986a). Carnivory is rare in the epiphyte synusia (Fig. 1), while trophic myr- mecophily (Fig. 12) is common and may, in fact, be entirely restricted to canopy habitats (Thomp- son, 1981; Givnish et al., 1984). Where substrata are deeply penetrable and moist, absorptive or- special permeabilities to match the character of the media (organic and acidic) they so often ex- oit. Water/carbon balance is also assured by con- siderable mechanistic variety. At one extreme, some canopy- -dependent plesdanhyes: alterne croclimate dictates: Quite the opposite, im- poundments provide continuous drought relief for hundreds of species of bromeliads (Fig. 8), fewer orchids, some other ts (Fig. 9), and ferns. Turnover of the entire leaf surface cued by impending seasonal drought is exemplified by members of several orchid genera (Fig. 2). Cras- sulacean acid metabolism (CAM) may be better represented in the forest canopy than in any other habitat type; stable carbon and hydrogen isotope data as well as more direct measurements of fix- ation pathway indicate that every known vari- ation on this mechanism exists in epiphyte flora. Occasional subjects—the shootless orchids, for BENZING— VASCULAR EPIPHYTISM 187 instance—exhibit novel machinery for gas ex- ange (Cockburn et al., 1985). Unusual osmotic qualities and related stomatal sensitivities ap- pear to rank among the most unique of the func- tional peculiarities of epiphytism (Benzing, 1986a). Survival is often predicated on extensive storage capacity, economical water use, and abil- ity to rebound rapidly from drought-imposed stress. Water balance is aided by special absorp- tive tissues that prolong contact with transitory fluids via mini-impoundment (e.g., the velamen of orchid roots; Fig. 10; Benzing & Pridgeon, 1983). Like other stress-tolerant plants, PS epi- phytes grow slowly, a characteristic which mod- erates resource requirements but heightens vul- nerability to habitat patchiness, disturbance, and other phenomena that oblige heightened fecun- ty. Habits of epiphytes are often specialized in ways other than those associated with impound- ment and myrmecophily. Abbreviation and merger of functions is common in the especially xerophytic forms, particularly the PS epiphytes; the most spectacular reductions occur in Bro- meliaceae and Orchidaceae (Benzing & Ott, and paralleled by complementary shoot m fication. Just about every intermediate condition between profuse and very sparse rooting exists throughout the complex of more than 500 species. Corresponding shoot changes center on a pro- gressively diminished impoundment capacity and an indumentum elaborated to the point where, among atmospheric forms, absorbent trichomes densely cover all foliar and most stem surfaces (Fig. 3). At this stage of specialization, no other organs are needed for uptake function. Orchi- daceae (subtribe Sarcanthinae) exhibit a com- parable progression except that here a telescoped, leafless shoot now supports production of green roots and the periodic axillary scape (Fig. The loss of axial differentiation seen exclu- sively in PS epiphytes may not be coincidental. Species that grow attached to impenetrable me- dia or hang free in the atmosphere occupy hab- itats as uniform as those colonized by prevas- — FIGURES 5-8. Neblina, Venezuela.— 6. the base of a tree in undisturbed lowland rain forest in Amazonas, Vene uela. in ant-nest carton in lowland Venezuelan rain forest. orida —5. A mixed colony of angiospermous epiphytes rooted in deep dense growth of bryophytes supporting ipd tolerant hymenophyllaceous ferns at 7. Seed humus on a palm on Cerro lings of epiphytes rooted —8. Tillandsia ey M a tank bromeliad, in south 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 , ` - _ CORTEX 1987] cular plants or modern aquatics. In such an environment, PS epiphytes need not maintain the division of labor between shoot and root that would be required for life on soil and, in fact, may sacrifice fitness by retaining a body plan inherited from terrestrial ancestors. Material and of energy econor vegetative Exo in organ system while other parts are lost may promote fitness where stress and high mortality place a premium on fecundity. Complex functional tradeoffs obliged by extreme a reduction in Bromeliaceae and J 1 (Benzing & ps 1981; Benzing et al., 1983 Somewhat less spectacular abbreviations leading to single-leaf ramets parallel increased exposure and stress tolerance in additional, predominantly epiphytic, orchid lines (e.g., Dendrobium, Epi- dendrum ). Although extensive studies have been made of orchid pollination, seed germination, and seed- ling nutrition, and some work has been done on mistletoes (e.g., Bernhardt, 1983), bromeliads (Smith & Downs, 1974), and a few other epi- phytes, life history characteristics of canopy-de- pendent vegetation as a whole remain obscure. What data are available indicate that nonuni- formity is the rule once again. An exception is the usual pattern of extended iteroparity; only a few Tillandsia and Vriesea species flower just once. Pollinators include all the common vectors except wind. Because air turbulence is adequate to disperse most epiphyte seeds despite their greater mass compared with pollen, absence of wind pollination must relate to other factors, most likely to the expense of required reproductive material or hyperdispersion of populations. Syn- dromes featuring specialized animals or unusual foraging activities are more common here than in other tropical synusiae. Breeding systems vary from those obliging strict outcrossing via traplin- ers to autogamy. Some minor patterns may be emerging, however. Orchids seem to be predom- inantly allogamous, while many ant-nest inhab- itants and Marcgraviaceae are selfers (Gentry & Dodson, 1987). The widely-held view that, on average, epiphyte populations are more frag- BENZING — VASCULAR EPIPHYTISM 189 mented than those of terrestrials and hence sub- ject to special genetic structuring and propensity for cladogenesis needs to be verified. If they are not, neither a uniform nor unique reproductive pattern should be expected. Seed is dispersed by various agencies. Among families, the vectors are most often animals (via fleshy fruits), but when species are counted, g wind-borne species (more than 80% of the total), seeds bearing elab- orate appendages are uncommon. Attempts to relate seed form and mass to mobility in aerial habitats can be complicated because disparate agents sometimes provide service for the same species. Dischidia seeds bear a plumose coma but also an oily eliasome to attract ants (short-range vectors; Docters van Leeuwen, 1954). Nest-in- habiting Codonanthe has a fleshy berry contain- ing myrmecochorous seeds. Other subtleties of epiphytic reproduction are just as easily over- looked — one flat side on otherwise fusiform Hyd- nophytum seeds reduces vulnerability to dis- lodgement by stemflow (pers. obs.). Seed size differential between terrestrial and epiphytic rel- atives is not consistent. Madison (1977) reported more massive seeds in soil-based compared with canopy-based Araceae and Cactaceae, but epi- phytic gesneriads definitely do not produce light- er seeds than do numerous terrestrial relatives (H. Wiehler, pers. comm.). Surprisingly, micro- sperms of terrestrial orchids are more buoyant than those of test epiphytes (Stoutamire, 1974). Rockwood's survey (1985) of 365 species in eight families from diverse communities in Costa Rica, Panama, and Peru yielded additional, unexpect- ed results. Average seed mass of 59 epiphytes (orchids excluded) was lighter than that of tree seeds but heavier than that of terrestrial herbs and shrubs. Table 1 tabulates angiosperm families con- taining more than 50 epiphytic species and lists characteristics that contribute to their success in tree crowns. Figure 13 summarizes adaptive di- versity within free-living, canopy-dependent, vascular flora relative to local moisture supply. Species with character states assigned to the far — FIGURES 9-12. section through the root of E pidendrum ] rise illustrating the major tissues, x 250.— —9. A “trash basket" Anthurium in lowland rain forest in Amazonas, Venezuela.— 10. Cro 11. Hemiepiphytic aroid in rain forest at Río Palenque, Ecuador. within the swollen hypocotyl. o [Vor. 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN 190 'S[ejop 10} 1X21 29g “SILIIQEYOIONW ut Ajpiqe rea? IMISIOW o1 Surp1oooe sayAydida 1e[noseA JO sajeis IILILYI Jo IUNIO `E] TANDIJ ` - =o Senma Wvo 31v9I1180 WY2-£t£2 , 9NI19A9-NWVO pe m — =e = - - S31AHdOO3WHSAW OIMdOHN1 i — I—MÀ — a — ç SINYdn250 N3QHVD 1S3N-1NV ——— r—— — — — — — - ———— — —— —— —— — o — — a a -- NOILONf3 3^11YV1393A Q3NISWOO' a — — — — — — — - ( 1853803 ANG) ( 183804 1YNOSV3S ) ( 1834804 NIYH/Qn012 ) 1N3L11lWH31NI A 13Q0IM Alddns G3IIddNS AISNONNILNOD SIdHVDONON 9Iddv2A10d SNOIYVA T1VWINV ONIM SHYOLDFA 3SHu3AIOQ AHQAHOTINIOd H9IH 1VILN310d 31N10S Ssnonai»2aa Wvo £2 329N3H343Hd WNINOWWY VZIHHHODAW H1MOHO MOIS Q3Sv8-NOIL1VAIIdID3Ud QaSv8-SnwWnH 31949 3417 *W31SAS 9NIQO33u8 :lySu3dSiIq a¥Odsvia :NOI1VNIT11Od NOI19nqOtudaqd :39NV 1V8 NOSYVI/HY3LUM *NOILIH ARN NOILONNA 3^AI1V1393A NOILONGIY 3AI1V1393A AHdHOWOUu3X 1N3WONñ0OdWI 1vWwHuadid3 31AHdid3iW3H AHVWIHd Snoaova8uaH AQOOM [LIGVH :3unix31 SHNLONYLS 3AI1V1393A 1987] right of the grid (indicated by lines) are usually PS types. Those on the extreme left experience continuous supply punctuated only by the un- common, normally brief, drought. Epiphytes bearing character states assigned predominantly to the left or right of Figure 13 can occur together. Pulse-supplied and CS forms inhabit humid sites, although only PS epiphytes occupy tree crowns in drier locations. Twig and free-hanging “mist” types in rain forests, for instance, receive mois- ture and nutrients exclusively via periodic flow, so their supply is intermittent. A nearby tank or humus epiphyte experiences considerably less discontinuity between rain events. Species situ- ated on highly exposed sites in humid forests or on most anchorages in arid woodlands exhibit slow growth, long-lived photosynthetic organs, and foliar or root surfaces modified to prolong contact with passing fluids (i.e., thick velamina or absorbing trichomes). Vegetative bodies are often much abbreviated, another means of im- proving economy. Woodiness seems to be sus- tainable only where moisture is abundant. Features associated with regular resource sup- ply are diverse. Hemiepiphytes are continuously supplied, at least while rooted in soil; habits may be woody or herbaceous. Nutrient i drawn from various types of humus by roots and ab- sorptive leaves of various CS forms. Carnivory is possible for the tank former. Ant-nest gardens are scarcer in the drier forests, but trophic myr- mecophily is widely available. Too little is known about mycorrhizae and nitrogen nutrition in the tosynthetic pathway, first supplemented with CAM-cycling and then ever greater reliance on nocturnal fixation, seems to play a diminishing role from left to right. Precise ordering of CAM variations in Figure 13 and the proposed signif- icance of CAM-cycling as a mechanism that *poises" the plant for drought (Ting, 1985) should be considered provisional at this time. Poikilo- hydry is feasible only on relatively humid sites for reasons described below. Deciduousness (e.g., Catasetum: Fig. 2) occupies an intermediate po- sition, occurring where drought is predictable (seasonal) and not too severe. High solute po- tential has been documented among CS and PS epiphytes (Harris, 1918; Spanner, 1939; Sinclair, 1983) although its effect on stomatal sensitivity has not been examined extensively. Vegetative features seem to have no bearing on type of pol- BENZING— VASCULAR EPIPHYTISM 191 linator used. Dispersal syndromes correlate only weakly with habitat humidity or resource pro- curement mechanisms. Anemochory is found everywhere, but a pulse-supplied nature in- creases the probability that an angiosperm will be wind-dispersed. Propagules that pass safely through a bird's gut are generally larger (more expensive) than the smaller anemochores, a re- lationship that may help explain why wind-dis- persed Tillandsia and Orchidaceae dominate the most stressful epiphytic habitats. Without small seeds, low productivity would probably result in low generative capacity for the PS epiphyte. The few monocarpic bromeliads are relatively well- provisioned impounders rather than stress-tol- erant atmospherics; the latter’s resource pro- curement capability is probably inadequate to support semelparity (Benzing & Davidson, 1979). In summary, epiphytes engage in a broadly similar rather than a narrowly proscribed way of life. Many other ecological groups are better de- fined because members subjected to less varied environmental constraints conform to more co- herent adaptive syndromes. Inconsistency in the epiphyte synusia is a function of habitat breadth (numerous possibilities for adaptive specializa- tion)and th ftree crown occupants. More than one daina com- bination can be associated with specific types of rooting media and climates, and no single key feature or suite of features is evident. Vegetative henomena that seem to be proportionately bet- ter represented i in canopy- than in soil-dependent patchiness and disturbance. Modes of pollen and seed conveyance are also various among epi- phytic plants, with few, if any, characters unique to these taxa. Indeed, most features of canopy- dependent plants occur among terrestrial flora as well, although not necessarily in the same com- binations. SYSTEMATIC OCCURRENCE AND THE MULTIPLE ORIGINS OF EPIPHYTISM Approximately 10% ofall vascular plant species are at least occasionally epiphytic, but distribu- tion among higher taxa is uneven (Gentry & Dodson, 1987). Broad involvement is particu- [VoL. 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN 192 so1Áudido Aqqniys *po1oo1-snuinu peru so1&udooounur orqdoi g soa1Audido Áqqnuus *pa1oo1-snuiny peau SIoptOA? 1ugnoJp *sajÁudido uopue3-1sou *sa1AudooouniAui orgdou L SIopioA? 1u8noip *so1Audooounuir orqdou] ‘soyAydida po1001-snui -ny pe13u33 pue uopie3-1s2N so1Audido uopue3-1s2N so1Aud -IdƏ 12xseq-use1 ‘SWOJ WY) *8uunpuo-1gdnojp ‘SWOJ uonoərmsəi: :sa1Audooounikui orgdoj L s1o[Sue]s K[[e£uorse2oo *uisnÁud -Iidoruiou Areurud 'ÁpooA, soyAydido po1001-snuiny [e19u28 *so)Audidoruiou Arepuosas so1Aydids po -1001-snuinu u33jo “SUIUTA so1Audidaruiau Ápoo M (SULIOJ WY) 8uunpuo-1]8nojp) so1&udido Sq ‘saiAydida po1oo1-snumnu peru (sououdsoun?) so1&ud -Ido Sd 'so1Áudido JUL L sə1Kudidə pəo1ooi-snuinu [&19u28 pue 193seq-usen *sa1Áudidorurou Kiepuooog saidydido on&udoros *po1oou-snuinqu jeru Áuqdaouirodox ‘WYO ssousnonpioop ‘sue 10J SIoquieqo poouq :juourpunoduuroJoe JA ssou -snonpioəp :S]U? 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Arboreal monocots far outnumber dicots, even exceeding Filiopsida (44 vs. 2796) for proportional occur- rence in forest canopies. Of the two large lycopod genera, Lycopodium is about one-third epiphytic (143 spp.) while Se/aginella —a considerably larger, heterosporous group— contains only five such species (Beitel, 1979). A majority of Psi- lotophyta regularly inhabit tree crowns. Gym- nosperms, on the other hand, are largely confined to soil, in part no doubt because of their heavy seeds and costly wind pollination. Orchidaceae have been more successful than any other lineage in canopy habitats. Two out of three epiphyte species are orchids; about 7096 of the family is canopy-dependent. Two other large monocot groupswithar ra- ceae, especially Anthurium, Tiladandron. nd Rhaphidophora, and Bromeliaceae, more than half of which anchor on bark. Canopy-dwelling dicots are disproportionally represented by Cac- taceae, Ericaceae, Gesneriaceae, Melastomata- rsatility within participating clades is not consistent. Compo- nent taxa may be diverse or uniform in the mech- anisms that foster habitation in tree crowns (Ta- ble 1). In all, about 80 vascular families contain at least one canopy-dependent member, but there are conspicuous, occasionally puzzling, omis- sions. Some very large, ecologically diverse groups with numerous tropical members are little rep- resented in, or even absent from, canopy floras. Leguminosae may be constrained by fruit and seed types that offer little potential for aerial dis- persal. Poaceae (and all Cyperales) may be al- most entirely terrestrial owing to consistent wind pollination and the graminoid habit, which does not lend itself to evergreen drought endurance. Plausible reasons for the paucity of epiphytes this family. These CAERpHODP e n the more advanced taxa. The same is true of ferns, but again, there are exceptions. Two members of the primitive order Ophioglossales root in hu- mus impounded by persistent palm leaf bases ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 and Platycerium fronds. If Psilotum and Tme- sipteris of the bigeneric Psilotophyta are indeed sole remnants of the Silurian/Devonian psilo- phyte complex, epiphytism is possible for the most primitive vascular relics. Epiphytism has probably evolved indepen- dently from terrestrial stock in every participat- ing family of seed plants and most families of ferns. It has arisen more than once in most of those spermatophyte taxa containing canopy-de- pendent species on different continents (Gent & Dodson, 1987), indicating lI Spit for arboreal life is rather fundamental in s major clades. Most diverse of ic Wi bros types, phylogenetically speaking, are the humi- philes whose drought liability obliges access to moist organic debris such as that covering older bark exposures in everwet forests. Virtually every family containing epiphytes includes at least one such species; most contain no other type. Where more exacting conditions (e.g., exposed bark and twig surfaces) prevail, few families are repre- sented, although these are sometimes speciose (Benzing, 1978). Neotropical ant nests provide specialized substrata for a small myrmecocho- rous flora which often offers extrafloral nectar. Perhaps additional traits as yet unrecognized that might affect capacity to root in cartons are mostly limited to Araceae, Bromeliaceae (specifically subfamily Bromelioideae), Cactaceae, Gesneri- aceae, Marcgraviaceae, Orchidaceae, and Piper- aceae. Some Asclepiadaceae, Melastomataceae, and Rubiaceae are part of similar but less well- defined symbioses in Australasia (Janzen, 1974). Those trophic myrmecophytes that exhibit un- equivocal modifications for ant occupancy all be- long to Asclepiadaceae, Bromeliaceae, Melas- tomataceae, Orchidaceae, Polypodiaceae, and Rubiaceae (Huxley, 1980). Rubiaceae seem to be most specialized for the relationship (Fig. 12). Stranglers—about 300 in all—come from just eleven families of dicots, most notably Moraceae (Ficus). Secondary hemiepiphytes have vining habits, and indeed the majority belong to groups with a scandent tendency (e.g., Araceae, Cyclan- thaceae, Marcgraviaceae). Bromeliaceae largely account for the tank formers (Fig. 8). Trash-bas- ket habits (Fig. 9) are present in a modest number of families; the numerous PS epiphytes are al- most exclusive to Bromeliaceae (mostly Til- landsia) and Dreidaceue (many subtribes). Both parallelis much sada in the rise of epiphytism. Par- o, 1987] allelism is illustrated by CAM, which features a y mechanism (incorporation of CO, via 8-car- boxylation) fundamental to regulation of ionic and osmotic balance through metabolism of ma- late and perhaps other organic ions (via a bio- chemical pH-stat). Heightened enzyme activity, appropriate malic acid mobility and storage ca- acity, phase-shifted stomatal behavior, and oth- er associated characters enabling high water use efficiency have emerged in close to 30 families and more than once among some of those con- taining epiphytes— Bromeliaceae and Orchida- ceae, for instance. Inherently slow relative growth rates, great longevity of whole plants and their individual parts, along with underlying oligotro- phic physiology in canopy- and soil-based vege- tation alike, probably also reflect common po- tential realized under similar selective impetus. Somewhat more unique features based less on homology than on novel design opportunity are, among others, devices to effect impoundment and/or accommodate ant occupancy. On bal- ance, gresource use appear to be based on relatively few. widely available, tures effecting resource procureme mo numerous and varied origins, s. likely based on convergence. PREDISPOSITION AND ADAPTIVE CANALIZATION Compared with vertebrates, vascular plants hibit varied habits, tolerances, and life histories. Quite commonly, natives of disparate habitats even belong to the same genus. Difference in evo- lutionary pattern partly reflects the higher plant's greater physiological plasticity aided by contin- uous turnover of both vegetative and reproduc- tive organs. Selection may occur among ra- owe even tissues within individual rgans— during the life of a single clone (Walbot & Cullis, 1985). Evolutionary tempo is rapid enough to effect speciation in as little as 104—105 years—or perhaps a few decades! —in Orchida- ceae (Gentry & Dodson, 1987). Reputedly, only a small number of genes need to be involved to alter structure, function, and ecological tolerance (Gottlieb, 1984; but see Coyne & Lande, 1985). BENZING— VASCULAR EPIPHYTISM 195 Some angiosperm lineages are quite conserva- tive, however, as if for them, too, entry into cer- tain adaptive zones restricts access to others. Cases of evolutionary canalization abound in Magnoliophyta. Somatic conservatism is fre- quent within genera and an invariant adaptive mode can typify entire, albeit small, specialized families (e.g., Sarraceniaceae, Lemnaceae). More impressive evidence for constraints on direction but not speciation in plant evolution is provided by greater redundancy in other families. Exam- ples include halophytism in Chenopodiaceae, ruderalism in Brassicaceae, drought-enduring xerophytism in Cactaceae and, of course, epi- phytism in Orchidaceae. No single life style char- acterizes the entire clade in any of these exam- ples, but particular themes turn up too often to deny underlying family-wide dispositions. Suc- cess in a particular environment is predicated on a novel set of features that imparts ability to counter major external constraints. Once the es- sential components were perfected and integrat- ed, whether by serendipity or some optimizing mechanism, each ancestral stock became better equipped to enter one, and less so another, adap- tive zone. Since accessible space was widely available in each case, considerable cladogenesis ollowed, sometimes through agencies unrelated to vegetative performance, hence native substra- tum. Orchidaceae achieved unequalled specia- tion in humid, neotropical forest canopies be- cause of behavior peculiar to male euglossine bees. Adaptive syndromes are comprised of struc- tural, functional, and phenological components which can be identified through consistent oc- currence in participating species. Ruderals suc- tats. Components of this well- ma re ah ii are: | ical and root tissue; vigorous S nud iran d small, long-lived, often light-sensitive, propa- gules; and self-compatibility. Here, such traits as low shoot/root ratios, woodiness, and extensive succulence are never found because they are in- consistent with ephemeral life histories. Al- though the habits of vascular halophytes range from large trees (mangroves) to succulent terres- trial and submerged herbs, certain physiological phenomena are common to all because potentia for desiccation is always one of the predominant 196 problems in saline media. Osmotic compensa- tion and compartmentation of specific solutes provide the only means for maintaining water relations in hyperionic habitats. In every case, potentially toxic ions, taken in from the envi- ronment to lower tissue water potential, must be sequestered in vacuoles at concentrations above those encountered in glycophytes, and they must be balanced across the tonoplast by osmotica of plant origin. Salt balance can be fine-tuned fur- ther by taxon-specific features. Chenopod halo- hytism is aided in salty surroundings by mul- ticelled vesicular hairs which excrete excess Na* an ^ succulence provides for ion dilution. The presence of organic solutes, including pro- line, in other salt-tolerant taxa instead of the be- taines found in Chenopodiaceae reveals that ments is not complete. Likewise, the abundant malic acid synthesized by all CAM plants at night for subsequent daytime genera- tion of an internal CO, supply is usually decar- boxylated through only one of the three existing reactions: mediation by NAD- or NADP-depen- dent malic enzyme or PEP carboxykinase. On balance, floristic variety generally reflects the equability and heterogeneity of a particular adaptive zone, all other variables being equal. Fertile, moist soils and abundant light have . wide convergence—ruderals are a good ample. Weeds in crop fields belong to many nde reflecting easy access to a collection of required character states. Tropical forest cano- pies have been similarly colonized but only to a certain degree. Whereas many lineages have evolved traits that allow growth on the organic arboreal “soils” of everwet forests, few have in- vaded more demanding zones, as demonstrated by narrower taxonomic participation in PS epi- phytism (Benzing, 1978). Inherited —in effect, phyletic — constraints de- termined which ancestors of modern families could generate epiphytic derivatives. Inherent barriers are complex and little understood, but their existence in plants is becoming recognizable (e.g; Kochmer & Handel, 1986; Hodgson & MacKey, 1986). Potential to express key com- ponents of an adaptive syndrome was not suffi- cient to assure its establishment. Universal oc- currence of CAM-like function, for instance, did not guarantee adoption of CAM by all xero- phytes nor by all aquatic macrophytes in the soft- water lakes where nocturnal fixation is favored ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 by limited carbon supply (Keeley, 1981; Rich- ardson et al., 1984). Moreover, each character of a syndrome must be free of pleiotropic and link- age relationships with maladaptive or internall incompatible traits. Biomechanical compatibil- ity is essential; for instance, plants with vigorous cambia may rarely employ CAM, probably be- cause costly woodiness is not sustainable without greater capacity for carbon gain. ittle is known at this point about inertia in the genetic, epigenetic, and biomechanical inter- play among phenomena responsible for salt bal- ance in halophytes, resource uptake and pro- cessing in short-lived eutrophs, or water economy in CAM plants. But occurrence among related genomes does demonstrate that some characters affecting vegetative performance and thereby suitability in particular habitats — e.g., sites of ni- trate reduction (shoot as opposed to root), ability to adjust osmotically under various degrees of stress, and presence of particular ion pumps and More narrowly restricted throughout Magnoliophyta, often to a single family or order, are features such as capacity to form a particular type of mycorrhiza, nodulate with Rhizobium, synthesize cardiac glycosides or betaines rather than anthocyanin pigments, and utilize sorbitol rather than sucrose for phloem transport. Access to vital characters during plant evolu- tion has been affected by related impacts on pho- ton, water, and nitrogen use efficiency (Raven, 1985a). Constraints on evolutionary opportunity have been strongest where drought, shade, or infertile substrata accentuate premiums on water, energy, and nutrient economy respectively. Type of nitrogen source, for instance, imposes different demands, depending on where (in which organ) processing takes place, how much water and en- ergy is needed per unit of product, and the en- vironmental context (is light or moisture scarce or abundant?). Calculation of comparative costs creted by NO, assimilators. Although the am- monium-to-protein pathway is least expensive in terms of energy consumption, overriding fac- tors may still dictate another choice even where NH, is in greatest environmental supply. Soil is the usual sink for H: generated by NH,* use. 1987] Indeed, owing to the immobility of protons in phloem, terrestrial plants process most of their acquired NH, in roots, a potential limitation with special relevance for epiphytism. If this is the universal rule, then what compensation, if any, accompanied root system reduction in ad- vanced Bromeliaceae? Evidence indicates that NH, is the predominant form of nitrogen in at least some tropical-forest canopies (e.g., Curtis, 1946). Conceivably, the absence of similar mor- phological diminution in nonbromeliad lineages is in part related to their less flexible nitrogen metabolism, although the role bromeliad foliar trichomes have played in obviating absorption by roots cannot be ignored in such comparisons. Perhaps slow-growing plants like the atmospher- ic bromeliads metabolize N at such low rates that complications are avoided. Either the internal biochemical pH-stat is adequate for disposal or excess protons are dumped from trichomes when shoots are wette Similarly, land plants have evolved several mechanisms to effect osmotic balance. Here, cost escalates with deployment of inorganic (e.g., Na*, Cl), then organic nonnitrogenous, and finally nitrogenous solutes, especially on infertile me- dia. Again, nitrogen is an important currency but perhaps no less so than the energy and water spent for its acquisition and processing. In a somewhat different vein, fungal biomass has to be supported by the mycorrhizal plant; but re- turns in phosphorus, and sometimes water and other nutritive ions, justify the investment in all but the most fertile habitats. Comprehensive cost accounting, through knowledge of functional in- compatibilities and the hereditary and epigenetic phenomena controlling access to key traits, is necessary to interpret patterns of radiation. It is at these levels of plant performance that many of the tradeoffs, economies, and accommoda- tions mo the evolutionarily-stable strat- egy oc HISTORICAL BASIS FOR CANOPY DEPENDENCE It is currently impossible to explain fully why one particular lineage developed canopy depen- ence while another did not. But partial answers are available in some cases; several of the more notable ones are discussed below. Four questions provide focus: Why are proportionally more ferns than seed plants epiphytic? Why do so many BENZING— VASCULAR EPIPHYTISM 197 monocots, particularly orchids, dwell in cano- pies? Why have several families of dicots with bvious advantage by basic habit or water balance profile hun a ie so widely there? Fi- nally, why is branch parasitism relatively un- common? Ferns. Homosporous pteridophytes are suc- cessful in tree crowns, where they usually occupy lower strata, because of small diaspores. A sec- ond, less obvious, factor is a capacity to tolerate substantial drying and deep shade. Poikilohydry is pronounced in exceptional taxa (Polypodium polypodioides), but many other filicaleans exhibit desiccation tolerance superior to that possessed by most seed plants. A fern's pattern of drought resistance, unlike that of most CAM plants, is particularly compatible with occurrence deep in the forest. For instance, ultrathin fronds of Hy- menophyllaceae (Fig. 6) probably photosaturate at very low fractions of full insolation and they can survive considerable desiccation. Greater ex- posure might be tolerable, and upper as well as lower strata heavily colonized, but for the trade- off associated with poikilohydry. Resurrection is adequate for countering the occasional brief drought every umid forest ascent now and then, but frequent d another matter. A regulated water ebum e on thick (ex- pensive, opaque) epidermal barriers and greater diffusive control is critical on markedly arid sites because photosynthesis is more likely than is res- piration to be curtailed by severe water deficits. d moisture supply be too intermittent and = i O 9 ! Raven (1985b) cited rates of physiological pro- cesses, including photosynthesis and transpira- tion at the low end of the ranges reported for tracheophytes, as a reason why ferns are so well equipped to inhabit shady, drought-prone loca- tions. Nevertheless, a modest invasion of drier lo- cations has been possible for higher ferns. One enabling mechanism here is drought avoidance via seasonally deciduous foliage (e.g., PAlebo- ium aureum). Occurrence in some stressful Australasian sites is possible for evergreen Pyr- Drymoglossum piloselloides, Pyrrosia longifolia, and P. angustata (Wong & Hew, 1976; Sinclair, 1983; Hew, 1984) along with drying character- 198 istics more reminiscent of a resurrection plant than a typical desiccation-resistant xerophyte. A thorough examination of ferns with regard to microclimates, substrata, and water and carbon relations in both gametophyte and sporophyte stages will be necessary to place discussion of the evolution of pteridophytic epiphytism on a firm- er foundation. Liliopsida as a whole. Orchids account in large measure for the immense numbers of epiphytic species, but monocots would be disproportion- ately common in tree crown habitats even with- out them. Bromeliaceae and Araceae rank sec- ond and third. The ratio of monocot to dicot species in tree crowns is 5: 1, but itis 1 : 4 overall. Although Araceae, Bromeliaceae, and Orchida- ceae are the most successful families in forest canopy habitats, there is no common adaptive theme. Two photosynthetic pathways in many variations, tank and trash-basket impound- ments, myrmecophytism, foliar trichomes, vel- phytic monocots. A peculiar body plan, shared to some extent with the higher ferns but less so with dicots, may have offered special class-wide opportunity. Monocots in particular, but dicots as weil, are often sectoralized in the sense that individual shoots operate as collections of relatively inde- pendent, serially aligned, physiological units (IPUs; sensu Watson & Casper, 1984). Tracer studies have shown that meristems may receive fixed carbon mainly from nearby leaves, some- times only those attached at the subtending nodes. Longitudinal segmentation seems to be more characteristic of dicots, and may also help ex- plain why several different habits and associated ecological tendencies have been emphasized in one or the other ofthe two classes. Partitionment into vertical compartments is evidenced by movement of labeled photosynthate among leaves and associated buds along a single or- thostichy. Xylem supply is similarly restricted. Secondary thickening may eventually obliterate conductive barriers to lateral flux imposed by the dicot primary body, peice where eu- ure as previously noted, r set of limitations, especially in arid habitats. Monocots with their more reticulate '**atactostele," best known today ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 in palms (a group portraying the **Raphis Prin- ciple"; Tomlinson ) appear to possess unique capacity for functional integration on one hand and, on the other, habits that permit ex- tensive vegetative renewal in mimimal space. Perhaps especially fortuitous is the unusual abil- ity to coordinate remote sources and sinks, and a related capacity to localize effects of damage and maintain vascular supply to organs that a more rigid system might be forced to abandon or underutilize. This kind of flexibility was il- lustrated by itl mature ramets, returned to sink status by shading or defoliation (e.g., Callaghan, 1984; Welker et al. icd. remained alive rather than self-pruned, as commonly occurs in forest trees. Historically, peers of this sort may have fostered ex- traordinary architectural plasticity n special implications for novel evolution. Recall that sev- eral large epiphytic monocot lineages have undergone major vegetative reorganization un- equaled in dicot counterparts. The exceptionally well-developed horizontal segmentation just described was undoubtedly important in the evolution of stress-tolerant epi- phytic monocots, perhaps ferns as well. The ad- vantages of a rhizomatous sympodial habit and potential physiological autonomy of the single phyton (a morphological unit composed of a leaf and associated adventitious root(s), bud(s), and subtending stem segment) were most accentuat- ed during orchid phylogeny. Sequential produc- tion of reduced shoots (in effect, shoots com- posed largely of single expanded phytons in extreme cases) is a recurrent theme in this family. Adult shoots of Dendrobium ultissimum consist of nothing more than strings of stubby, closely placed, leafless pseudobulbs. The broader suit- ability of architecture based on serial renewal via determinate shoots generated from closely-placed meristems is illustrated by occurrence beyond the monocots. Somewhat less condensed ver- sions of the same general arrangement exist mong mechanically-dependen , some Gesneriaceae) and ‘Lycopodium. Differentiation of roots into feeder and holdfast types (Fig. 9), a useful division of labor for the vine or epiphyte, seems to be more common in monocots than elsewhere. onorchid monocots. Bromeliaceae, with far fewer species and almost exclusively neotropical distribution, nevertheless rival Orchidaceae for variety of epiphytic life styles. Tank habits have dao5oo5vo vv hen eo 1987] evolved independently in two subfamilies, and in all three if Brocchinia is correctly assigned to Pitcairnioideae (Benzing et al., 1985). A rosulate shoot was required for each transference of ab- sorptive role from root to shoot. Ancestry was apparently mesic in both Tillandsioideae and nk shoots are associated with ating soil substitutes in leaf bases, are funda- b CAM plants. Specialization for PS iphytism, in effect for greater stress tolerance, Y: proceeded farthest in Tillandsioideae by way of the derived atmospheric forms (Benzing et al., 1985). Here, absorbing trichomes (Fig. 3) are per- fected to the highest degree whilst the vegetative apparatus is reduced to simplest form. Epiphytic Bromeliaceae, more than most, were clearly adapted to endure rigorous conditions by the presence of a suitable epidermal appendage and habit in precursors. Here, epiphytism is based on a modified shoot (Fig. 8) with the foliar tri- chome as its keystone feature (Table 1). Absorp- tive function might be possible in glabrous leaf bases when long-term contact with moist, nutri- tive tank fluids is maintained, but the rapid up- take required to sustain a rootless, nonimpound- ing PS bromeliad would be impossible without an extraordinary foliar indumentum. Myrme- cophytism, and perhaps a single case of carnivory (Catopsis berteroniana; Givnish et al., 1984), are also associated with specialized trichomes and inflated leaf bases (Benzing, 1970). Hypotheses concerning how the bromeliad foliar epidermis may have acquired its current function and im- portance are described elsewhere (Pittendrigh, 1948: Benzing et al., 1985). (Briefly: contrary to Pittendrigh’s proposition that absorptive func- tion would only emerge under drought selection, Benzing et al. posited a mesic, infertile, ancestral habitat where the foliar epidermis evolved pri- marily to promote acquisition of nutrient ions from impounded humus or perhaps animal prey.) Bromeliad seeds are disseminated by birds (Bro- melioideae) or wind (Pitcairnioideae and Til- landsioideae). Pollination syndromes are diverse and apparently not associated with either tank or atmospheric habit. Aroid, by comparison with bromelioid or or- chidoid, epiphytism is neither as advanced nor as versatile. There are no reports of CAM here, and overlapping foliage that might mitigate drought lacks the water-tight quality possessed BENZING— VASCULAR EPIPHYTISM 199 by inflated bromeliad leaf bases. Trash-basket catchments (Fig. 9) sometimes trap falling litter but little moisture. Pots fail to produce velam ving the most drought; tolerant Orchidaceae, nor is there any indication that these organs can match foliage in photosyn- thetic vigor. Seasonally deciduous leaves on green or tuberous stems occur in Philodendron and Re- musatia respectively, but these are minor themes with few participating species. Arboreal exis- tence in Araceae is based predominantly on two mechanisms, both humus-based: impoundment, seen in short-stemmed Anthurium and some Philodendron (Fig. 9); and secondary hemiepi- phytism (Fig. 11), a more widespread phenom- enon most often M. xin in Philodendron. nd t d vining hab- its incorporating progressive dieoff of proximal stem regions, appear to be the central vegetative features responsible for aroid expansion in can- opy habitats (Table 1). Both sympodial (e.g., Philodendron) and monopodial (Pothos) habits are found in the hemiepiphytes. Ant nests are utilized by some Anthurium and veo ei Water and nutritional relations exhibit no ob- vious unusual modHE onis for arboreal life but neither have b losely. Baccate fruits are an integral part of the syndrome, but they occur throughout the family without habitat re- striction. Pollinators range from beetles to eu- glossines. Specialized pollen vectors may have encouraged enlargement of Anthurium and pos- sibly other genera. Cyclanthaceae, the only other nonorchid monocot family with a sizable epi- phyte contingent (about 6696 of Asplundia), mostly penetrate the forest canopy as rooted climbers and secondary hemiepiphytes. True epiphytism occurs in Sphaeradenia and Stele- stylis (G. Wilder, pers. comm.). Stems and inter- nodes are shorter than those of related hemiepi- phytes. Orchid monocots. Orchidaceae owe their nu- merical superiority among epiphytes to an ex- ceptionally propitious set of vegetative and re- productive features and to extraordinary cladogenesis promoted by specialized pollina- tion syndromes (Benzing & Atwood, 1984). Veg- etative mechanisms vary tremendously accord- ing to the taxon's native substratum and ecoclimatic conditions. But there are several im- portant attributes common to all canopy-depen- dent family members that, in some form, pre- disposed early stock for arboreal life. For example, 200 epiphytic Orchidaceae possess specialized roots varying in photosynthetic performance and water relations, depending on structure and metabo- lism; uptake is enhanced in all cases by a non- living velamen which imbibes precipitation and contained solutes for subsequent sorption through transfer cells in an underlying exodermis (Benz- ing et al., 1983; Benzing & Pridgeon, 1983; Fig. 10). This same mantle effectively retards desic- cation injury from both short-term and extreme drought (Benzing, 1986a). Velamina simply em- bolize air in order to break the hydraulic contin- uum that, if left intact, would allow matric forces in adjacent drying substrata to dehydrate living epiphyte tissue. A green cortex supplements shoot photosynthesis and is the major site of carbon gain for the plant in exceptional cases (Fig. 4). Hyperovulate gynoecia and aggregated pollen characterize most of the family. Microspermy— up to millions of tiny, lightly provisioned seeds per capsule—requires fungal intervention for germination. High-fidelity, long range, but often inefficient pollinators promote ethological iso- lation leading to plant speciation (Benzing & At- wood, 1984). Pre-epiphytic orchid stock probably possessed structurally modified, locally suberized epider- mis/hypodermis layers as do most extant terres- trial family members and some other monocots (e.g., Zea, Allium, Amaryllis). Similar root spe- ae is sl eee m dicots. Microspermy and associ were prob- ably also CREE ina terrestrial context, as sug- gested by the current habitats in which all other such Hetesoiznphic: plant hoops occur (e-g.. Production of numerous tiny diaspores, subsis- tence on transitory resource supplies, and main- tenance of high water and nutrient use efficien- cies set the stage for migration to forest canopies, including many uninhabitable by less stress-tol- erant epiphytes. Evolution of pheromonelike fragrances and specialized floral morphology tightened relationships with specific hymenop- terans and dipterans, and assured extensive pro- liferation of several clades that happened to be canopy- -dependent. Large es ba related ort-range vectors with r no known propensity for exclusive foraging suggest that substratum-specific factors have also been important. As in a number of other diverse communities of sessile organisms, coexistence of densely packed epiphyte popula- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 tions may be favored on substrata subject to in- termediate disturbance (Connell, 1978; Benzing, 1981, 1986b). Radiation in other speciose, can- opy-dependent taxa visited by opportunistic or sedentary pollinators (e.g., Anthurium, Peper- omia) may be due to similar phenomena. Dicots. Magnoliopsida are, on the whole, poorly disposed to epiphytism — just 2% of species ry is often predicated on a single theme and no other family incorporates the diverse resource hibited by epiphytic Bromeliaceae and Orchi- daceae. Except in Marcgraviaceae, dicot terres- trials always outnumber confamilial epiphytes. In tree crowns, Peperomia ranks first in size among successful dicot genera and even families, a statistic fostered by pantropical distribution, the presence of CAM variations (Sipes & Ting, 1985), and high-volume production of small ad- hesive fruits. Habits range from shrubby to mi- nute and creeping. Moraceae also owe much of their major epiphytic presence to a single genus (Ficus) with a similar broad range; here the stran- gler habit provides the vegetative basis for suc- cess. Rampant speciation within a relatively nar- row adaptive profile has again been encouraged by circumtropic range and host-specific polli- nators—in this case, the fig wasps. According to Ramirez (1977), the moraceous strangling habit evolved *'as a response to lack of light at the forest level." Necessary attributes for success in- cluded presence of viscid hyaline coats on seeds that would germinate only on moist humus, long aerial roots, water-use-efficient seedlings, and dispersal by winged vertebrates. Marcgraviaceae and Clusiaceae are additional single-strategy families, emphasizing secondary and primary hemiepiphytism respectively. Stranglers also oc- cur in Schefflera (Araliaceae), Posoqueria (Ru- biaceae), Metrosideros (M eae) and else- where, but they are atypical among confamilial canopy-based taxa Most epiphytic Asclepiadaceae belong to closely related, succulent, vining Dischidia and oya. Flasklike leaves of D. rafflesiana and sev- eral other species enclose nests; ants provide dis- persal service for many more. Forms with less specialized foliage regularly root in or grow against ant debris, providing clues as to how ant-fed rel- atives evolved. Dome-shaped leaves of D. col- lyris grow tightly pressed against bark, providing 1987] shelter for Iridomyrmex colonies (Huxley, 1980). Photosynthesis involves CAM and/or CAM-cy- cling (Kluge & Ting, 1978). Cactaceae became equipped for the canopy-dependent synusia through drought selection in terrestrial habitats. Fleshy, small-seeded fruits and climbing habit would eventually favor life in canopies. Move- ment into tree crowns appears to have involved some reversals. Originally aphyllous, stems of the most advanced epiphytic forms (which hap- pen to be natives of humid forests—e.g., Zygo- cactus, Ripsalis) have lost their armature and become much flattened, or narrowed if still terete (e.g., Hatiora), presumably to improve perfor- n shade. Family-wide CAM is probably mands requires uptake by long-lived roots from more or less continuous supplies in tree fissures or soil (the secondary hemiepiphytes). Despite the extreme drought tolerance of many terrestrial relatives (e.g; Maxillaria, Ferrocactus), epi- phytic Cactaceae seemingly never colonize the most demanding bark and twig exposures. Less obvious is the basis for high epiphyte suc- cess in Ericaceae, Gesneriaceae, Melastomata- ceae, and nonmyrmecophytic Rubiaceae. Most canopy-dependent members in all four families grow exclusively on humus mats in humid for- ests. Woody habits and sclerophyllous foliage, sometimes complemented by storage tubers, characterize Ericaceae and Melastomataceae. About one-half of the rubiaceous epiphytes (Hydnophytum, Myrmecodia) supplement min- eral nutrition and store moisture via ant-inhab- ited, swollen hypocotyls (Fig. 12). Basically herbaceous Gesneriaceae feature broader growth- form variety, and several genera contain CAM plants. Like some Peperomia, these gesneriads exhibit trilayered mesophyll that may signal un- usual photosynthesis. Substrata are more diverse in this family, ranging from ant-carton to less specialized humus. Codonanthe and related gen- era, along with some hemiepiphytic cacti, are probably the best drought-insulated of the dicot epiphytes. Baccate fruits provide seed mobility in most cases, although Rhododendron, a few gesneriad genera, and large proportions of Me- lastomataceae and Rubiaceae ripen wind-borne seeds. Ant-associated species are myrmecocho- rous. Representation in canopy habitats varies among these families, ranging from 4% to 35% BENZING— VASCULAR EPIPHYTISM 201 of all genera in Rubiaceae and Ericaceae respec- tively (Madison, 1977). The ericad statistic 1s all the more ing the family’s size ane numeraus temperate taxa. The other three and neal ly OI exclusively moist- tropical, hence have had greater access to arbo- real habitats. Breadth and depth of specialization for canopy dependence is further indicated by comparing the total of exclusively epiphytic gen- era containing two or more species with the num- ber which include soil-based species as well. Ges- neriaceae are most canopy-adapted by this measure with 13 genera meeting each criterion, while Ericaceae is least so with only four terres- trial-free genera out of 22 containing epiphytes; the largest of the four contains only eight species. Vaccinium is especially noteworthy for its wide range throughout Old and New World boreal to Melastomataceae and Rubiaceae are 8:12 and The influence of geography in determining which families would contribute the most species to the forest canopy flora (Gentry & Dodson, 1987) is apparent in Magnoliopsida. Epiphytic Gesneriaceae are centered in the neotropics where tribe Episcieae and particularly genera like Dry- monia, Columnea, Dalbergaria, and Trichantha have radiated extensively in tree crowns. Spe- ciation here, as in epiphytic Liliopsida (Araceae, Bromeliaceae, Orchidaceae), appears to involve specialized American pollinators (humming- birds and euglossines). Paleotropical counter- parts are far fewer but still number more than 100 species. Canopy-dependent ericads are also disproportionally neotropical and ornithophi- lous. All but a small fraction of the peperomias are American, although little is known about the group’s reproductive biology. Because 73% of Costa Rica’s large fern flora roots in tree crowns (Wagner & Gomez, 1983), it appears that neo- tropical epiphytism is not all pollinator-related. Wide availability of moist montane forest in the northern Andes and Central America, which is reputedly amenable to fine niche partitionment (Gentry & Dodson, 1987), has also been impor- tant for expansion of American forms. Melasto- mataceae and Rubiaceae number among the few heavily epiphytic p families that failed to generate more n Old World epiphytes. Scrutiny of le aasan systems and pertinent vegetative characters might help ex- 202 plain why mechanically-dependent New World portions of these two families have failed to undergo as much cladogenesis as have many co- occurring epiphytic pteridophytic and angio- spermous groups. Chemical peculiarities of substrata or unusual mycorrhizae may be responsible for uneven epi- phyte development among higher monocot and dicot taxa. Ericaceae, Melastomataceae, and Or- chidaceae exhibit family-wide affinity for acidic, often humid and organic, infertile soils. Little is probably often scarce. Substrata in everwet forest where most epiphytes live tend to be sodden, at least moderately acid, and certainly organic. Use of NH,* rather than oxidized nitrogen by plants native to such substrata may have been a pre- disposing character for epiphytism. Ericads are notably deficient in nitrogen reductase, a sign of long dependence on reduced nitrogen. Ericaceae were perhaps especially well positioned for can- opy invasion through formation of mycorrhizae of a type seen in some extant terrestrials that mobilize nitrogen and phosphorus from sterile, organic soil (Stribley & Read, 1975; St. John et al., 1985). Terrestrial Orchidaceae are also strongly mycorrhizal, but the advantages, if any, that fungi provide canopy-dependent adults re- main little studied and controversial (Hadley & Williamson, 1972; Sanford, 1974; Benzing & Friedman, 1981). Broad surveys of epiphyte roots and nitrogen-cycle microbes in canopy substrata could prove rewarding. A general work-up of tree- crown media as nutrient sources is much needed. Mistletoes. Restricted taxonomic participa- tion in branch parasitism is made all the more intriguing by the widespread abundance of mis- tletoes, some of which range farther north than any vascular epiphyte. Virtually the entire com- plement—about 1,300 species— belongs to San- talales. Why many thousands of plants from so many other higher taxa root nowhere but on bark yet never invade host vasculature is puzzling. Occurrence in Santalales of terrestrials with haustoria, including primitive Loranthaceae, but no epiphytes suggests that branch parasitism arose from root parasitism. Perhaps more direct aerial transitions were precluded by biomechanical constraints. Mistletoes transpire profusely even during drought in order to acquire sufficient ni- trogen from host xylem (Ehleringer et al., 1985). This is the antithesis of an epiphyte’s usual con- servative water use pattern. Failure by so many ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 epiphytic lineages to achieve parasite status sim- ply because invasive organs are difficult to ac- quire seems unlikely in light of the diverse soil- based species that tap any nearby roots. More plausible is the theory that continuous function during a historical crossover between the hig solute potential/low maximum turgor/sensitive stomata pattern of the epiphyte and the three to five times more concentrated osmotica (Harris & Lawrence, 1916; Harris, 1918) and correlated foliar conductance patterns of the parasite would require buffering, which is less available in can- opy than in terrestrial habitats. Most of the root function in the conventional manner, drawing upon soil moisture and thus reducing the liability to water loss of transitional forms. SUMMARY Epiphytes do not constitute a narrowly defined group of plants on either taxonomic or functional grounds, nor is their habitat usually unique in its important physical characteristics. Growing conditions in both humid and arid forest cano- pies overlap with those at ground level, as do adaptive mechanisms in canopy- and soil-de- pendent flora. Substrata exhibiting the most unique and powerful constraints on plant success occur in the driest exposures. Circumvention of dependence on soil by epiphytes is varied. Modes of moisture and nutrient acquisition are the most unusual, especially among PS forms. Ways in which resources are conserved and deployed are often variations on, and perhaps sometimes identical to, patterns present in terrestrial rela- tives. No single key feature—like CAM, im- poundment habit, or animal dispersal—nor combination of key characters underlies epiphyt- ism. Solutions are many, a condition favoring broad taxonomic composition of mechanically- dependent synusiae. In contrast, failure of a large tropical group to evolve epiphytic members may relate to the presence of a few well-entrenched, maladaptive features. Evolutionary inertia prob- ably also reflects little-understood genetic con- straints and functional incompatibilities that block emergence of enabling syndromes even though some of their mechanistic components are already well established. Cladogenesis and adaptive radiation for life on varied substrata have been more closely allied in some epiphytic taxa than in others. Most fam- ilies, even some containing hundreds of epi- 1987] phytic species, have entered forest canopies through a single vegetative theme. Propitious re- lationships with pollinators, intermediate dis- turbance, and narrowly circumscribed substra- Orchidaceae are exceptional for their diverse vegetative mechanisms. These two families do well under equable conditions, but they are es- pecially dominant in stressful environments be- cause they can maintain enough generative pow- er to compensate for mortality imposed by habitat patchiness and disturbance. LITERATURE CITED BEITEL, J. M. 1979. 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Crassulacean acid metabolism and crassulacean acid metabolism modifications in Peperomia camptotricha. Plant 77: 59-63. Downs. 1974. iiri queen Pt. Un ntersuchungen uber den Warme und Wasserhaushalt von Myrmecodia und Hya- n tum. Jb. Wiss. ia 88: 243-283. STOUTAMIRE, W. P. Terrestrial orchid seed- ds ngs. /n C. L. MEE (editor), The Orchids, Sci- ntific Studies. Wiley- "Md cred New York. SEE D. P. & D. J. READ. 1975. Some nutritional aspects of the biology xe ericaceous mycorrhizas. Pp. 195-207 in F. E. Sanders, B. Mosse & P. B. Tinker (editors), Endomycorrhizas. Academic Press, New York THOMPSON, J. N. 1981. Reversed animal-plant in- teractions: Hei evolution of insectivorous and ant- fe ed plants. Biol. J. Linn. Soc. 16: 147-155. Tina, I. P. Us pon qiu metabolism. Ann. Rev. Plant. Physiol. 36: 595-622. TOMLINSON, P. B. 19 Development of the stem . Dickison (editors), Contempo Problems in Plant Anatomy. Academic Press, London. WAGNER, W. H. & L. D. 1983. Pteridophytes (Helechos, Ferns). Pp. 311-318 in D. H. Janzen (editor), Costa Rican "Natural uon Univ. Chi- cago Press, Chicago. WaLBOT, V. & C. A. CuLLIS. 1985. Rapid genomic change in higher cane Ann. Rev. Plant Physiol. 96. GÓMEZ. WATSON, M. A. & B. B. CASPER. 1984. Morphoge- in dud Ann. Rev. id Sst 15: 233-258. WELKER, J. M., E. J. , D. D. Bri esi D. GOESCHL. 1985. . ort among veg- etative tillers within two bunchgrasses: ie with carbon-11 ME Pie ae 67: 209-212. WoNG, S. C. & C. S. HEw. 6. Diffusive resistance, titratable acidity, and eo fixation in two tropical epiphytic ferns. Amer. Fern J. 66: 121-124. DIVERSITY AND BIOGEOGRAPHY OF NEOTROPICAL VASCULAR EPIPHYTES! ALWYN H. GENTRY AND C. H. DopsoN? In his classic work Schimper (1888), empha- sizing the taxonomic diversity of epiphytes, list- ed 33 families and 232 genera of epiphytes. Until very recently, subsequent authors have generally accepted Schimper’s figures (Richards, 1957; Jo- hansson, 1974). However, epiphytism (here used in a broad sense to include hemiepiphytes, see Kress, 1986) was recently reported to exist in 65 different vascular plant families (56 families ex- luding ferns), 38 of these with epiphytes in the Neotropics (Madison, 1977). Our own data (Ap- pendix 1) and additional records compiled by Kress (1986) now record 83 vascular plant fam- ilies with epiphytic species.* At least 876 genera include at least one epiphytic species and there are perhaps 29,000 epiphytic species, ca. 10% of all vascular plants (Table 1). Thus at first sight epiphytism seems a very widespread and suc- cessful life-style, which very many unrelated taxa ave evolved. However, a closer examination suggests that even though there are both many species and higher taxa of epiphytes, few of the higher taxa account for most of the species. Burger (1985), for example, emphasized that relatively few lin- eages have been able to enter the epiphytic niche, presumably because of the complex suite of ad- aptations needed. Thus even though it is true that the evolution of an epiphytic habit has been a relatively common feature of vascular plant evolution, it is equally true that very few of the taxa that have evolved an epiphytic habit have radiated successfully to produce other epiphytic species (Table 2). In most of the epiphyte-con- taining families, epiphytism is a rather insignif- icant anomaly. Indeed, eliminating a mere 85 such “oddball” species from the roster of the world’s epiphytes removes 31 families from the epiphytic ranks. Only 32 seed plant families have as many as five or more epiphytic species, 26 of these with epiphytes in the Neotropics. It is on the 42 families (Table 3) that contain = in the Neotropics that this paper will Even though this analysis of epiphyte posset and distribution is largely focused on the Neo- tropics, a few comparisons with the Paleotropics are instructive. There are actually slightly more families with epiphytes in the Paleotropics (43) than in the Neotropics (42), with all of the pa- leotropical epiphytic families having epiphytic dipped in Australasia but only 15 in Af- and Madagascar. If only the 32 seed plant families with five or more epiphytic species are considered, there are also roughly equal repre- sentations of epiphyte-containing families in the Neotropics (26) and Australasia (25) but only about half as many in Africa (14). At the species level the story is very different. There are many more epiphytes in the Neotrop- ics, at least half again as many as in Australasia and six times as many as in Africa. Althoug similar numbers of genera and families evolved epiphytism in the different regions, subsequent speciation as epiphytes was dramatically greater in the Neotropics. A major objective of this pa- per, then, will be to try to explain why there is so much epiphyte diversity in the Neotropics. EPIPHYTE FAMILIES One approach to an overview of neotropical epiphyte diversity is a taxonomic one. Table 3 summarizes the neotropical epiphytic seed plant flora by family. Of the 42 families represented by at least one habitually epiphytic species in the Neotropics, the Orchidaceae are by far the most important with ten times as many epiphytic neo- ! We thank the National Science Foundation (INT- ps 6840; BSR-834- ei for — of our Ecuadorian fieldwork. Drs. H. Bedell, T. Croat, B. Hammel, J. Lut binson provided unpublished data on disteibuHon ‘of epiphytism in bru r, L. B. Smith, M. Dillon, and taxonomic specialties. We n, S. Renner, P. Tay H. R especially thank B. Hammel for making available oe data on the habit composition of the La Selva ; B. Burger r, T. Croat, B. Hammel, M. Madis cata Botanical Garden, P.O. Box 299, St. L n, S. er and E. Zardini for much technical ipe in its prepar ouis, Missou ri 63166, and E. Zardini for reviewing the manuscript; U.S.A. 3 Th set used in this paper was s derived a Su that of Kress (1986) and is mostly based on Madison’ s figures as modified in Appendix ANN. Missouri Bot. GARD. 74: 205-233. 1987. 206 BLE |. Taxonomic distribution of vascular epi- s (modified from Madison, 1977 and Kress, 1986). Fami- Gen- lies era with with Species i- f Epi- phytes phytes phytes Pteridophytes 13 92 2,593 Gymnosperms 2 2 4 Monocots Without orchids 80 2,657 Orchids (fide Madi- son) 500 20,000 Orchids (compiled ressler, 1981) 460 15,000 Orchids (fide Kress, 6 440 13,951 Total (fide Kress, 1986) 17 520 16,608 Total (our estimate) 17 540 22,657' Dicots 51 262 4,251 Total 83 896 29,505 ! Includes Madison’s orchid figure (see text). tropical species as runners-up Araceae and Bro- meliaceae. All three of these most speciose neo- tropical epiphyte families are monocots. One other monocot family, Cyclanthaceae, also has a significant number of epiphytes. Commelina- ceae, Rapateaceae, and Philesiaceae, although with few species, have an exclusively epiphytic genus (respectively, Cochliostemon, Epidryos, Luzuriaga, and (in our experience) Philesia). As summarized by Madison (1977), the rest of the epiphytic monocot flora of the Neotropics con- sists of single species of Burmannia and Yucca and a few Central American species of Smila- cina. Since orchids are so overwhelmingly the most diverse group of epiphytes (about 7096 of their species are epiphytic), estimates of orchid diver- sity are critical to an evaluation of epiphyte di- versity. Unfortunately, orchids are amazingly poorly known taxonomically (compare the ca. 12 orchid taxonomists with the ca. 200 system- atists specializing i in the similar-sized Compos- ler, 1981) or 17,000 (Airy Shaw, 1973) to 30,000 (Madison, 1977) or 35,000 (“some authors" fide Dressler, 1981). In the best available review Dressler (1981) counted almost 20,000 species ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 and suggested 20,000-25,000 as the best esti- mate of orchid species numbers. Since 7096 of the total number of orchid species should ap- proximate the number of species of epiphytic orchid, there should be between ca. 12,000 (from Airy Shaw's estimate) and ca. 20,000 (from Mad- ison's estimate). Unfortunately, the 8,000 species "slop" between these two estimates is as great as the total number of epiphytes in all other families combined! Madison (1977) used the higher figure for his calculations, whereas Dressler (1981) gen- erally opted for more conservative estimates of orchid species numbers. We have mostly taken the higher values since we know of many cases where Dressler's figures for species numbers are significant underestimates but none where he has overestimated. For example, Dressler suggested that there are 830 species in subtribe Epiden- drineae, whereas in Ecuador alone there are 500 species in the single genus Epidendrum and we think 1,200 species would be a better subtribal estimate. Moreover, new orchid species are being dis- covered at an astonishing rate, especially in the northern Andean region, again suggesting that Dressler's estimates of species numbers will have to be adjusted dramatically upward. For exam- ple, about 2,315 orchid species are now known from Ecuador, ca. 700 of these described only in the last 15 years. Nevertheless, more than 1,500 additional Ecuadorian orchid morphospecies" ave not been identified with any Even if the 300 unaccounted for names should all prove applicable to the unidentified speci- mens at hand, it is inevitable that most of the unidentified taxa will prove undescribed and the list of orchids for Ecuador alone will increase to well over 3,000 species. The neotropical epiphytic dicot flora is more diverse in families but much less diverse in species than the monocots. Twenty-nine dicot families have at least one habitually epiphytic species in the Neotropics. The largest of these are Pipera- ceae (ca. 500 spp.), Gesneriaceae (483 spp.), Me- lastomataceae (227 spp.), Ericaceae (ca. 300 spp.), Cactaceae (133 spp.), Guttiferae (ca. 90 spp.), and Marcgraviaceae (87 spp.). In addition there are perhaps 110 neotropical species of Moraceae stranglers in the genera Ficus and Coussapoa. The only other dicot families with more than 20 epiphytic species in the Neotropics are Aralia- ceae, Bignoniaceae, Compositae, Rubiaceae, and Solanaceae (Table In addition, Bombacaceae, though with few 1987] GENTRY & DODSON-— NEOTROPICAL VASCULAR EPIPHYTES 207 TABLE 2. Largest epiphyte families (in part from Madison, 1977). No. Genera wit No. Epiphytic Percent Family Epiphytes Species Total No. Species Epiphytes Orchidaceae 460 20,000 30,000 67 (—13,951') (—19,128') (73) Bromeliaceae! 26 1,144 ,500 46 raceae 15 1,100? 2,5004? 42 cia 53 1,023 1,100 93 Piperaceae 2 710 3,000 24 Melastomataceae 33 ca. 647? 4,770? 14 Gesneriaceae 28 598* 3,000* 20 Moraceae (incl. stranglers) 3 521 1,400 37 ricace 28 4785 4,000 23 l TN 2 400 600 67 Aspleniaceae! 1 400 675 59 Dryopteridaceae! 10 292 1,920 15 Rubiaceae 21 217 6,000 4 Lycopodiaceae! l 200 400 50 Davalliaceae! 8 139 150 10 Asclepiadaceae 6 135 2,000 T Cactaceae 25 133 2,000 [Í Cyclanthaceae 7 125° 205 61 Vittaria 9 112 11 100 Guttiferae 6 92 1,000 9 Marcgraviaceae 7 897 117 76 Araliaceae 5 73 700 10 * Wiehler, 198 5 Luteyn, pers. comm. * Hammel, pers. comm. ? Bedell, pers. comm. species, has an epiphytic genus (Spirotheca). The other 16 epiphytic neotropical dicot families are represented by only occasional epiphytic species of predominantly terrestrial genera. In addition to these angiosperm families, there are two gymnosperm families with single epi- phytic neotropical species in generally terrestrial genera and at least 838 epiphytic fern species belonging to 32 different genera, a common epi- phytic Psilotum, and some epiphytic species of Lycopodium. REPRODUCTIVE BIOLOGY A number of salient characteristics that may be critical to success as epiphytes are shared by many different neotropical epiphytic taxa. Mad- ison (1977) nicely summarized many of the fea- tures of epiphyte reproductive biology and this discussion is largely based on his. From the view- point of dispersal biology, there are three main types of epiphyte propagule. The great majority of epiphyte genera and species have tiny dust- like wind-dispersed sporochores, often with highly sculptured epidermis, to aid in air flota- tion. Such seeds, representing an extreme in r-se- lection and a high risk gamble on chance es- tablishment, are found in the two most successful epiphyte groups, orchids and ferns, as well as in such taxa as Begonia (although at least one Af- perhap capsular melastomes, although seeds of capsular melastomes may be | mm long and are not strict- ly comparable (Renner, pers. comm.). In closed- canopy tropical forests such seeds are virtually unique to epiphytes. While tiny sporochore seeds are found in some tropical weedy herbs, they are unknown among tropical forest lianas (except mostly hemiepiphytic Adelobotrys) and trees (ex- cluding tree ferns), although the pterochore seeds of genera like Chimarrhis may not be any larger. 208 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 3. Epiphytic neotropical seed plant families (in part from Madison, 1977). Number of Neotropical Epiphytic Neotropical Genera Family Species with Epiphytes Distribution of Epiphytes Gnetaceae l Gnetum also epiphytic in Malaysia Zamiaceae l mia Costa iie pM only in Neotro Agavaceae l Yucca Mexico; ann in Neotropics Araceae 1,034 Anthurium, Caladiopsis, Mon- also epiphytic in Africa and stera, Rhodospatha, Philo- Asia dendron, Stenospermation, yngonium Bromeliaceae 1,144 (fide 18 genera entirely or predomi- only in Neotropics Burmanniaceae Commelinaceae Cyclanthaceae Dioscoreaceae Liliaceae Orchidaceae Philesiaceae Rapateaceae Apocynaceae Alzateaceae Araliaceae Asclepiadaceae Begoniaceae Bignoniaceae Bombacaceae Burseraceae Cactaceae Campanulaceae Compositae Crassulaceae Ericaceae Gentianaceae Gesneriaceae Kress, 1986) 125 4 11,000 (fide Madison) 3 nantly epiphytic; 5 others with some epiphytes Burmannia Cochliostemon, Campelia Asplundia, Dicranopygium, Evodianthus, Ludovia Sp P Stelestylis, cocarpus Smilacina ca. 80 genera entirely or pre- dominantly epiphytic Luzuriaga, Philesia Epidryos, Stegolepis Mandevilla Alzatea Schefflera, Oreopanax Cynanchum Begonia Schlegelia, Gibsoniothamnus Spirotheca Bursera 25 genera entirely or predomi- nantly epiphytic Burmeistera es r Neomirandea, Pseudogynox- ys, Sen necio (Pentacalia), Sin- clairia, Tuberostylis Echeverria 18 genera entirely or predomi- nantly epiphytic; several with epiphytic species oyria 12 genera entirely or predomi- nantly epiphytic; 4 other also epiphytic in New Gui uinea only epiphytic in Neotropics only in Neotropics Ecuador also epiphytic in — Pacific, and Madagasca also epiphytic in Africa me Australasia also epiphytic in New Zea- land only in oo Costa R Costa Rica-Colombia also i in Africa and Austr mostly snobi in Malaysia also epiphytic in Africa and sia only in Neotropics only in Neotropics; mostly Andean Costa Rica Rhipsalis also epiphytic in Africa and Ceylon neotropical; mostly Andean n New Zealand and Mad- agascar also few epiphytic in Hima- layas and Madagascar also epiphytic in Australasia South America also epiphytic in Africa and Australasia GENTRY & DODSON —NEOTROPICAL VASCULAR EPIPHYTES 209 1987] TABLE 3. Continued. Number Neotropical Epiphytic Neotropical Genera Family Species with Epiphytes Distribution of Epiphytes genera with some epiphytic species Griseliniaceae 3 Griselinia Chile and Brazil; also epi- phytic in New Zealand Guttiferae ca. 90 Clusia, Clusiella, Havetiopsis, only epiphytic in Neotropics edem matopus, Quapoya, enggeria Lentibulariaceae 12 Utricularia also 2 epiphytic in Africa and Australasia Marcgraviaceae 89 all 7 genera entirely or pre- only in Neotropics dominantly Pone or miepiphyt Melastomataceae 227 7 genera sores or largely epi- also epiphytic in Africa and hytic; 5 others with some Asia epiphytic cles Moraceae 111 Coussapoa, Ficus subg. Uro- Ficus stranglers also in Afri- stigma, and 1 Pourouma ca and Australasi Myrsinaceae ca. 12 Cybianthus, Grammadenia, also epiphytic in Africa and Myrsine (Rapanea ia Onagraceae 3 Fuchsia only epiphytic in Neotropics Piperaceae ca. 500 Peperomia, Piper also epiphytic in Africa and Asia Rubiaceae ca. 57 Balmea, Coprosma, Cosmi- also epiphytic in Australasia buena, Hillia, Malanea, Ma- nettia, Psychotria, fm Eon nium, Schrader. apotaceae 1 melia Costa Rica Saxifragaceae ca. 3 PM Phyllonoma only in Neotropics olanaceae ca. 30 Juanulloa, Lycianthes, Markea also epiphytic in Malaysia (+ segregates), Solanum (Solanum, Lycianthes) Urticaceae ca. 15 Pilea also epiphytic in Indo-Ma- laysia The second most prevalent dispersal mode among epiphytes is via birds. Most bird-dis- persed epiphytes have indehiscent berry fruits but a few, including Drymonia and Clusia, have dehiscent capsules with arillate seeds. In either case the seeds tend to be smaller and more nu- merous than in related nonepiphytic taxa (Mad- ison, 1977). In some families there is a marked change in pigs mode accompanying the shift to epiphytis Bignoniaceae all epiphytic species Su two probably ogi dispersed excep- tions) are bird-disperse only one nonepi- phytic species (Synapsis Mentum: is (see Gentry, 1983). In Melastomataceae 8596 of the epiphytic species have berry ird as compared with 60% of the nonepiphytic species (Renner, 1986). The third major diaspore dispersal syndrome in epiphytes is wind-dispersal via winged or plumed seeds (pterochory and pogonochory, re- spectively). Interestingly, plumed seeds as com- with winged seeds greatly predominate among epiphytes, whereas the opposite holds true for trees and lianas, at least in mature forest species. Some of the important epiphyte taxa with pogonochore diaspores are Bromeliaceae subfamily Tillandsioideae, Asclepiadaceae, Ges- neriaceae, and Rubiaceae. In Rubiaceae some epiphytic genera have small winged seeds while others have true pogonochores; the difference be- tween these dispersal modes tends to break down in such groups, with some species having such narrow reduced n: that these effectively ap- proximate large hai Finally there are a m epiphyte taxa with such 210 miscellaneous dispersal syndromes as bat-dis- ersal (some strangler figs), exozoochory via sticky diaspores (some Peperomia), and the not readily classifiable “sloppy corn-on-the-cob" ingestion of some cyclanth fruits by Callicebus and other primates (Terborgh, pers. comm.). In general, epiphyte seeds are smaller and more numerous than those of nonepiphytic relatives. For example, Renner (1986 and pers. comm.) noted that in Melastomataceae mostly epiphytic Blakea and Topobea have ca. 1,000 seeds per fruit compared with a few deren seeds per fruit in typical nonepiphytic genera such as Miconia and Clidemia. Madison (1977) estimated that seeds of epiphytic Anthurium are typically ca. 2 mm long as compared with 4-8 mm long in ter- restrial Anthurium species. There are also excep- tions to this pattern. For example, Rockwood (1985) pointed out that in Gesneriaceae epi- phytic species actually have significantly larger seeds than do shrubs and herbs. According to Rockwood's (1985) analysis, epiphyte seed size tends to be bimodal; those groups with dust- seeds or other wind-dispersed seeds have the smallest seeds of any habit type while taxa not dispersed by wind have seeds averaging larger than those of herbs, vines, and shrubs, similar to lianas, and ga smaller than trees. Neverthe- less, since the great iid of epiphytes are wind- dipened. pon iphyte habit class as a whole is generally characterized by the smallest seeds of any habit class. A dispersal strategy em- phasizing many small seeds and chance estab- lishment is typical of the r-selection syndrome often found in weedy species. Epiphytes would seem to be most unusual in being r-selected com- ponents of mature forest ecosystems. POLLINATION Madison (1977) emphasized animal-pollina- tion as a characteristic trait shared by all angio- sperm epiphytes. While true, this is hardly re- markable in a tropical context since, with virtually no exceptions (Myriocarpon (pers. obs.), Trophis (Bawa et al., 1985), and just possibly Sorocea and a few Chamaedorea species (fide Bawa et al, 1985), all lowland tropical forest angio- sperms are animal-pollinated. Nevertheless, epi- phytes as a whole surely have a more pronounced trend toward highly specific and specialized pol- lination systems than do nonepiphytes, if for no other reason than that so many epiphytes are orchids. In addition to the well known orchid ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 pollination specializations (e.g., Dodson, 1967; van der Pijl & Dodson, 1966; Williams, this ymposium), many aroids have similar Euglos- sine-attracting scent systems. Five of the largest neotropical orchid genera have specific bee pol- linators attracted by specific scents and Anthur- ium is the largest nonorchid genus in the Neotropics. While Pleurothallis, the largest neotropical epiphyte genus, does not participate in the presumably speciation-promoting Euglos- sine-pollination syndrome, it is pollinated by the large and diverse fly genus Bradesia (CD, pers. obs.) and similar coevolutionary patterns may be involved; among other small-flowered and in- conspicuous but highly diverse orchid genera, Stelis and Lepanthes are probably pollinated by Drosophila and similar flies, and Telipogon is pollinated by pseudocopulation with tachinid flies, another very large and diversified insect taxon an Unique to the Neotropics, hummingbird-pol- lination is also much more prevalent among epi- phytes (and terrestrial herbs) than in trees or free- climbing lianas. Epiphytic taxa among which hummingbird- pollination is prevalent include Bromeliaceae, Gesneriaceae (espe- um aviaceae (Norantea, sensu lato), Rubiaceae (Ravnia, Manettia), and Cactaceae (e.g., Schlumbergera). The correlation between hummingbird-pollination and epiphy- tism is well shown by Bignoniaceae. Of the two epiphytic genera of Bignoniaceae, one (Gibson- iothamnus) is entirely hummingbird-pollinated and the other (Schlegelia) also has several hum- mingbird-pollinated species; hummingbird-pol- lination is rare elsewhere in the family. Other specialized pollination systems shown by epi- l, pe rat-pollination in B/akea chlorantha (Lumer: 1980). Perhaps more striking than the diversity of highly specialized pollination systems among neotropical epiphytes is their lack of the small, inconspicuous, generalist-pollinated flowers that characterize the great majority of trees in the wet forests where epiphytes are prevalent. The only epiphytic taxa characterized by such flowers are Araliaceae, Moraceae, Piperaceae, Myrsinaceae, and Urticac epiphytic m cialist pollinators (Stiles, 1981), are taken as an 1987] example, this pattern may be clearly seen at the community level: in lowland tropical forests hummingbird-pollination is almost exclusively confined to herbs and epiphytes. Another relevant aspect of epiphyte pollina- tion biology is that such phenomena as self-com- patibility and autogamy are apparently much more prevalent than typical in tropical lowland taxa. For example seven of seven species of B/a- kea, Topobaea, and Adelobotrys tested at Mon- LI Costa Rica were self-compatible (Lu- mer, 1980; Renner, 1986) vs. 34 of 43 tested oa melastome species in the Manaus area (Renner, 1984). In Marcgraviaceae this is carried to an extreme with all species tested being au- togamously pollinated in bud despite the elab- orate floral adaptations (Bedell, pers. comm.). In summary, epiphyte reproductive biology appears to be a unique mix of r-selection and specialization. Unlike other components of ma- ture forest communities, epiphytes share many reputedly r-selected traits with weedy herbs, es- pecially in their dispersal ecology. Yet at the same time most epiphytes have highly specialized pol- lination systems, strong niche specificity, and many other traits more characteristic of k-se- lected mature forest species. DISTRIBUTIONAL PATTERNS To this point, we may conclude that, although taxa have been more than marginally successful at speciation and adaptive radiation as epi- phytes. However, the few taxa that have suc- cessfully radiated as epiphytes have done so very prolifically. Even though remarkably similar numbers of plant families have achieved epi- phytism in the Neotropics and Paleotropics, the process of epiphyte speciation would seem to have been much accelerated in the former, to judge from the very many more neotropical species of epiphytes. w uld now like to examine some trends In epiphyte distribution that may help to under- stand not only neotropical epiphyte biogeogra- phy but also some of the continental differences in epiphyte occurrence. MOISTURE One of the most striking distributional pat- terns shown by epiphytes is a tremendous de- crease in both numbers of species and individ- GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES 211 uals in drier habitats. Although this pattern seems obvious, it is by no means well-documented. In- deed Walter (1985: 57) claimed that epiphytes, contrary to such other habit groups as lianas, are found in dry as well as wet tropical forests. At the other extreme, Schimper (1903) suggested that in areas with marked dry seasons epiphytes are either completely wanting or rare and that presence of epiphytes outside the rain forest is always a sign that the dry season is not long or is accompanied by copious dew. Data for 1,000 m? samples of western Ecuadorian dry forest (Capeira, 804 mm per year) and moist forest (Jauneche, 1,855 mm) quantify the extent of this difference (Gentry & Dodson, 1987; Table 4, Fig. 1). If our results are indicative, most plants in a wet forest are epiphytes. At Rio Palenque such a sample included 4,517 epiphytic plants rep- resenting 63% of all individuals sampled. At Capeira a mere ten epiphytic plants were includ- ed in a similar sample of dry forest, representing 0.2% of the sampled individuals. The moist for- est Jauneche site was intermediate with 116 epi- phytes constituting 4% of the individual plants sampled. The difference in epiphyte density be- tween wet and dry forest is almost 500 fold. Moreover the decrease of epiphyte density in dry forest contrasts greatly with the situation for other habit groups. The number of herbs more than doubles from our moist and wet forest sam- ples to our dry one. Contrary to Walter’s asser- tion (1985: 57), lianas double from wet to dry forest; they are much commoner yet in our moist forest sample, the latter presumably atypical since Jauneche happens to be the most liana-rich site in the Neotropics (of 45 similar samples). Shrub density also increases somewhat from wet to dry forest; unlike lianas, shrubs are only about half as abundant in the intermediate moist forest as in dry and wet forest. In contrast, the number of individual trees = 10 cm DBH, and thus the ap- parent density of the forest, changed little (52, 64, and 69 trees 210 cm DBH in wet, moist, and dry forest, respectively). Epiphytes are also important contributors to the species richness of neotropical wet forests. Indeed there are 35 epiphyte species in our 0.1 ha. wet forest sample, accounting for over a third of the sampled species. This compares with only 13 epiphytes (8% of the species) in the compa- rable moist forest sample and three (2% of the species) in the dry forest one (Fig. 2) The importance of the epiphytic contribution to species diversity is equally apparent when en- 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 4. Number of species and individuals of different habit types in 1,000 m? samples of three forests in western Ecuador. Río Palenque is wet forest, Jauneche is moist forest, Capeira is dry forest (from Gentry & Dodson, 1987). Río Palenque Jauneche Capeira No. No. No. No. No. No. Habit Group Spp. Ind. % Spp. % Ind. % Sp. 96 Ind. 9 Herbs (incl. palmettos) 50 14 1,220 17 18 11 944 34 50 29 2,854 53 Shrubs 39 11 531 7 16 10 279 10 13 8 742 14 Epiphytes (incl. hemi- epiphytes) 127 35 4,517 63 13 8 116 4 3 2 10 52 Climbers (incl. lianas; excl. hemiepiphytes) 36 10 117 2 58 34 484 17 58 34 895 16 Lianas 22.5 cm (excl. herbaceous + hemi- epiphyt.) 12 3 28 43 25 124 4 19 11 58 l Total tree spp. (incl. Ju- ven.) 114 3l 653' 9 64 38 960 34 48 28 927 17 Trees <2.5 cm DBH (= saplings + seedlings) 87 24 559! 8 48 28 672 24 38 22 750 14 Trees <10 cm, >2.5 cm 86 24 217 3 35 21 245 9 32 18 108 2 Trees 210 cm DBH 32 9 52 1 30 18 64 2 29 17 69 l Total herbs (epiphytes + herbs—woody cpi- 162 5,525 31 1,060 53 2,864 Shrub layer (shrubs + saplings) 126 1,090 65 948 51 1,492 Underlayers (< ca. 3 m) (herbs + shrubs + saplings) 176 2,310 83 1,892 101 4,346 Total 365 7,210! 169 2,783 173 5,428 ! Excluding dense patch of 123 Quararibea asterolepis seedlings. tire florulas are compared (Table 5, Fig. 3). AI- most one fourth of all the Río Palenque plant species are epiphytes (Dodson & Gentry, 1978). Similarly, in another lowland wet forest at La sea Costa Rica, dh of the species are epi- phytes (Hammel, pers. com ven in moist ed sites like Barro Cian ‘land: Panama (Croat, 1978) and Jauneche, Ecuador (Dodson et al., 1985), epiphytes constitute 12-1696 of the total flora. Only in dry forests are epiphytes rel- atively insignificant, accounting for 2-4% of the species of Capeira (Dodson & Gentry, 1987) and Santa Rosa National Park, Costa Rica (Janzen & Liesner, 1980). We conclude, contrary to Walter (1985), that epiphytes d tically in drier areas than does any other habit group, but contrary to Schimper’s (1903: 198) emphasis, a few vascular epiphytes are characteristically present in even the driest neotropical forests (e.g., Capeira with 804 mm of annual precipitation). Familial makeup of the epiphytic flora also changes with precipitation (Table 6). Many more families have epiphytic representatives in wet forests than in drier ones (Fig. 4), and many epi- phytic taxa are confined entirely to wetter forests. In the Neotropics the same families tend to be represented by epiphytes under similar climatic conditions. In the driest forests, the only epi- phytes are orchids and bromeliads, perhaps the two most specialized epiphytic families. Ferns, peperomias, and Cactaceae join orchids and bro- meliads in slightly moister conditions. The next epiphytic families to appear with increasing hu- midity are aroids, Moraceae (stranglers), and Gesneriaceae, joining representatives of all the dry forest families, each of whose number of epi- phytic species is maintained or increased. The two local florula sites with over 2,500 mm of precipitation, Barro Colorado Island (Croat, 1978) and Rio Palenque (Dodson & 1978), me remarkably similar epiphytic floras. he same seven epiphytic families are most species rich at both sites, in roughly the same 1987] GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES 213 100 9 80 4 EEB epiphytes 4 WET L* trees 63 4 DRY L herbs 60 1 53 * shrubs ° 2) 504 o? climbers = ° ç ° EJ = > Es] MOIST £ 40 1 34 34 4 4 e e ra 16 pr 20 4 17 4 € > 4 4 Pa < 1 » 4 4 9 4 |", P. e Pa P. Capeira Jauneche Río Palenque FIGURE 1. Percent of individual plants belonging to different habit groups in 1,000 m? samples of three western Ecuadorian forests. 50 : ENEB epiphytes DRY MOIST WET » Bo trees 4 40 - n r3 35 CJ herbs 4 4 3 io * shrubs a » ° M Ta & 31 Ó . o 29 ° G ~ ° š climbers ° 304 28 o? « < ° a o » P ° < < [7] e o » » » o 4 25 4 = < ° > > 2 > o 4 4 4 o » o P”) > o 4 [e] 4 4 ° P ° P. 4 96 > $9 » 204 » o < ° < < ° » o » » ° < ° < 4 ° » o 2 » ° < o 4 ° » ° > > o 4 ° < 4 ° P ° P P o 4 [e] 4 4 ° > ° > > ° < ° pi 10 4 Ç o? e ao < 4 o » ° "i » o 8 4 o 4 8 ° > ° Pa > o 4 o 4 ° > o 2 o * 1°, »* Ta » 2 1" * bar >. 96 > i * lo < ° 4 Capeira Jauneche Río Palenque FIGURE 2. Percent of species belonging to different habit groups in 1,000 m? samples of three western Ecuadorian forests. 214 MOIST saprophytes shrubs parasites & treelets & of species herbs, % epiphytes climbers trees SANTA ROSA CAPEIRA JAUNECHE B.C.. PALENQUE SELVA FiGURE 3. Percent of species in local florulas be- Peper to different habit groups. order, just as they are in Hammel’s unpublished La Selva, Costa Rica, species list. Remarkably, there are 81 epiphytic species of orchids at Río: Palenque and 82 at Barro Colorado Island (BCI), 35 epiphytic aroids at Rio Palenque vs. 30 at BCI, 18 species of epiphytic Bromeliaceae, and 14 species of epiphytic or strangler Moraceae at both sites. Only epiphytic ferns and fern allies are noticeably better represented at BCI than at Rio Palenque and only epiphytic Gesneriaceae and Cy the revers sites have three epiphytic cacti and one epiphytic melastome plus one or two epiphytic Araliaceae, Solanaceae, and begonias. Only Rubiaceae and Saxifragaceae have epiphytic members at BCI ut not Rio Palenque; only Urticaceae, Pole- moniaceae, Bignoniaceae, and Ericaceae have TABLE 5. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 epiphytic representatives in the Rio Palenque flora but not the Barro Colorado one. Again, the same epiphyte families predomi- nate at other wet sites in western Ecuador (Table 7). The exact same seven most species rich fam- ilies at BCI, La Selva, and Rio Palenque are the most species rich at Centinela, again in roughly the same order. Even at Tenafuerste (alt. 1,000 m) the five families richest in epiphytes are ex- actly the same ones that are most species rich at the other Ecuadorian wet forest sites. At the extreme wet end of the precipitation gradient there are other, as yet unquantified, changes in the epiphytic flora. In the wetter part of Chocó (precipitation > 8,000 mm) berry- fruited epiphytes like Melastomataceae, Ara- ceae, Marcgraviaceae, and Ericaceae, along with arillate-seeded Guttiferae, appear to predomi- nate to a much greater degree than at Rio Pa- lenque, while ferns (Sota, 1972) and dust-seeded orchids are more t poony fepresenieo. Possibly to establish themselves in the face of such su- aasan wa rainfa here m a aodceable habit change in the pou. d of the wettest sites. Most of the predominant Chocó epiphytic families are ac- tually hemiepiphytic, many of them woody as well. In the same sites with abundant hemiepi- phytes, free-climbing lianas become noticeably less prevalent. While the average of climbers > 2.5 cm in diameter for 0.1 ha. samples at two pluvial forest sites in the Colombian Chocó was exactly the same (68) as that for a series of 20 similar samples from neotropical lowland moist and wet forests, half the sampled pluvial forest climbers were hemiepiphytic vs. an average of Habit compositions of complete local florulas. Capeira, Ecuador and Santa Rosa National Park, Costa Rica, are dry forest; Jauneche, Ecuador and Barro Colorado Island, Panama, are moist forest; Río Palenque, Ecuador, is wet forest. anta Barro Capeira Rosa Jauneche Colorado Palenque Habit Category No. 96 No. 96 No. % No. % No. % Trees => 10 cm DBH 69 15 141 21 112 18 290 22 165 16 Small trees + large shrubs 28 6 64 10 60 10 151 11 99 9 Herbs 4 subshrubs 242 52 317 48 224 37 389 30 376 36 Epiphytes (including Pis dE 9 2 24 4 72 12 216 16 238 23 Parasites and saprophyt 4 l 6 l 4 l 12 l 6 l Lianas 46 10 52 8 81 13 149 11 87 8 Small vines 66 14 63 9 55 9 109 8 84 8 Total species 464 667 608 1,316 1,055 1987] GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES 215 TABLE 6. Familial composition of epiphyte floras in lowland forests with different precipitations. Santa La Selva, arro Jauneche, Rosa Costa pios bonia Ecuador* Costa Capeira, Makokou, Rica! or? 1,855 Rica? cuador* Gabon’ Family 4,000 mm 2,980 mm Py 750 m mm mm 1,550 mm 804 mm 1,755 mm Orchidaceae 109 81 82 33 8 5 21 76 35 30 10 — — 10 Ferns and allies 59 28 43 5 7 — 26 Piperaceae 12 19 10 4 l l — Bromeliaceae 29 18 18 6 3 2 eae 13 14 14 9 2 — 7+ Gesneriac 16 12 4 2 — Cyclanthaceae ll 8 1 — — — Marcgraviaceae 8 5 2 — — — — Guttiferae 11 4 2 — — — — Cactaceae 6 3 3 3 3 l l ace 2 3 — — — — — Araliaceae 2 2 l — — — l Bignoniacea 3 2 — — — — — Melastomataceae 2 l l — — — — Polemoniaceae — l — — — — — Solanacea 1 l 2 — — — — Urticaceae l l — — — — — Begoniaceae 2 l l — — — — laceae 4 — 1 — — — — Saxifragaceae — — l — — — — mmelinaceae l — — — — — Total epiphytes 368 238 216 72 24 9 66+ Percent of flora 24 23 16 12 4 2 6+ No. families with epiphytes 20 18 17 8 6 4 6 Ee Hammel, pers. comm. pos 4 Gentry, 1978. 978. ds 7 Hladik & ain in prep.; Florence & Hladik, 1980 and cited references. 2.71 l limbers sampled at the moist dub wet sits (Gentry, 1986). in a sense, hemi- epiphytic climbers seem somehow to replace free- climbing lianas in the wettest lowland forests and also in middle elevation cloud forests. ALTITUDINAL The epiphytic flora also changes in both di- versity and composition on an altitudinal gra- dient. The general tendency is for epiphytes to be better represented in intermediate elevation cloud forests. In the Andes the peak in epiphyte diversity appears to be between 1,000 m and 2,000 m, but it lies somewhat lower in Costa Rica and Panama. Few data are available, but a comparison of incomplete data sets for several Ecuadorian sites (Table 7) documents this trend for the lower part of the gradient. The sites for which relevant data are available are Centinela (600 m) and Tenafuerste (1,000 m), both on the western slopes of the Central Ecuadorian Andes, and Mera (1,000 m) on the eastern slope. Of the well-documented sites, Centinela has the most species of epiphytes, 337, or 3596 of the flora. This compares with 238 epiphyte species ac- counting for 2396 of the flora at nearby Rí lenque (alt. 200 m). The data for the two 1 000 m sites are less complete, with many species re- maining to be discovered. Tenafuerste, with much less cloud forest effect than Centinela or Mera, has the poorest epiphyte representation, only 31% of the flora. The extreme is pluvial Mera where few collections have been made as yet. Three NUMBER OF EPIPHYTE FAMILIES AND SPECIES VS. PRECIPITATION ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 N " bs o > o a o a à 400; 120 o - £ = @ number of families E a * number of species š u a P o a o o = 300 115 E = g 3 o ] 5 D [ " 5 n £ E ra 5 E : ° š 2oo 8 = fo 5 7 ° c z is] 2 ' S ` . - n m ° ° ° [6] — à 2 a 100} o e 45 , E: m E 1000 2000 3000 4000 Annual precipitation (mm) FIGURE 4. Increase in numbers of epiphyte species and families in local florulas as a function of precipitation. hundred twenty-two orchid species are already known and we expect that there may be as many as 200 additional epiphytes as well. In contrast, Peru’s Huascaran National Park, 3,500-5,000 m in altitude, has only seven epi- phytic species, constituting a mere 196 of the park's flora (Smith, pers. comm.). The differences between Mera and Tenafuerste point out that altitudinal and moisture effects on epiphytes are complexly interrelated. Similarly Gilmartin (1973) showed that the 17 species of Bromeliaceae that occur on both sides of the Ec- uadorian Andes occur at lower altitudes on the moister eastern slopes. Density of epiphytes, although also greatest at intermediate altitudes, does not closely parallel diversity; although unquantified, we feel that epi- phyte density in the Andes tends to be greatest around 2,000-2,500 m, again occurring at some- what lower elevations in Costa Rica and Pana- ma. Due to the high densities, epiphytes are often most conspicuous at these relatively high alti- tudes, even though relatively few species may be present. In middle elevation cloud forests, epi- phytes may make up as much as 30% of the foliar biomass and 4596 of the foliar mineral capital of a forest (Nadkarni, 1984). Contrary to what happens along the moisture gradient, familial composition of the epiphyte flora changes very little along an altitudinal gra- dient, or at least on that part of it for which we have data. The same families are important in roughly the same order. The seven families with the most epiphyte species at Centinela (600 m) are the same seven that have the most species at Río Palenque (200 m). The five families with the most species at Tenafuerste (1,000 m) are the same five that have the most species at Río Pa- lenque and Centinela. The most noteworthy dif- ference between these sites is the absence of Mo- raceae stranglers at Tenafuerste; Gesneriaceae are also conspicuously less diverse at the 1,000 m more epiphyte diversity at intermediate altitudes (four spp. at Centinela and two at Tenafuerste but only one at Río Palenque; see also Renner, 1986). A few families (e.g., Bignoniaceae) dis- appear from the middle elevation epiphyte flora and most other families have decreasing num- bers of epiphytes at higher elevations. Our only high altitude data set is for Huas- carán National Park, Peru (Smith, pers. comm.), where the seven epiphyte species, all restricted to the lower part of the park between 3,500 and 1987] 4,000 m, belong to four families. The four fam- ilies with epiphytes— Piperaceae, Bromeliaceae, Orchidaceae, and ferns—are all in the top five epiphytic families in the wet Ecuadorian sites. Of the usually prevalent epiphyte families, only Araceae is lacking. Perhaps more interesting, the epiphyte families at Huascarán are exactly the same ones that are represented at Santa Rosa, Costa Rica, except that Cactaceae is missing. Ap- parently at environmental extremes, either alti- tudinal or VOULU only these same fam- ilies that are otherw most successful as epiphytes are able to survive. The very interesting but controversial sugges- tion has been made that in the tropics diversity is generally greatest at middle elevations along an altitudinal gradient. This has been shown for ed Bay herps (Scott, 1976), insects (Janzen, ; Janzen et al., 1976), and suggested for T Greater equability is a likely controlling factor for this putative ‘“‘mid-elevation bulge” in species diversity. However, data for 0.1 ha. sam- ples of plants 22.5 cm DBH suggest that plant species diversity decreases more or less uniform- ly from the most diverse lowland wet forest sites to the least diverse high altitude ones (Gentry, 1982a, 1987c). If a mid-altitude bulge in plant species richness really does occur, it must be due largely to epiphytes. Unfortunately our data sets from middle and upper elevation forests are too incomplete to be definitive. Indeed one of us (CD) thinks that because of the increase in epi- phytes there are more plant species at middle elevations than in lowland tropical forest while one of us (AG) thinks that the decrease in species numbers of such other habit groups as lianas and trees with altitude outweighs the increased num- ber of epiphytes. In either case the role of epi- phytes in the plant community is presumably greatest in middle elevation forests. SOIL FERTILITY To our knowledge no attempt to relate epi- phyte diversity to soil fertility has been made previously. Indeed one might suppose that since epiphytes are intrinsically “insulated” from di- rect dependence on soil nutrients they would be relatively unaffected by changes in soil fertility. For example, Janzen (1974a) discussed the sym- L'a: 14° LC hat t » | i. A nce ants in low-diversity poor soil “kerangas” hab- itats in Borneo, with the implication that epi- phytes are unukialiy well represented in such GENTRY & DODSON — NEOTROPICAL VASCULAR EPIPHYTES 217 TABLE 7. Familial composition of epiphyte floras in wet forests at different altitudes in Ecuador Rio Pa- Centi- e lenque nela fuerste Family 200 m 600 1,000 Orchidaceae 81 133 68 Araceae 35 52 26 Ferns 28 38 28 Piperaceae 19 19 11 Bromeliaceae 18 23 18 M eae 14 10 — Ges ace 12 16 8 Cyclanthaceae 8 5 3 Marcgraviaceae 3 2 Guttiferae 4 9 3 Cactaceae 3 2 l Ericaceae 3 9 9 cto 2 4 — Bignon 2 2 — Melastomataceae 1 4 2 Polemoniaceae 1 l — Solanacea 1 2 1 Urticaceae l 1 l Acanthaceae — 1 = Rubiace — l — Total 238 337 181 Percent of flora 23 35 31 habitats. Whitmore (1984) also emphasized the frequency of epiphytes in these forests. It is increasingly well-documented that major changes in the diversity and floristic composition of other components of tropical plant commu- nities are associated with changes in soil fertility (e.g., Ashton, 1976, 1977, 1978; Huston, 1979, 1980; Gentry, 1987b; Gentry & Emmons, 1987). For example, there are generally fewer tree, liana, and terrestrial herb species in neotropical forests on poorer soils (Gentry, 1981; Gentry mons, 1987). We have few data with which to relate epiphyte community composition to soil fertility. One of us (AG) has compiled species lists for a series of sites on different substrates in the Iquitos, Peru, area which share a similar rain- fall and climatic regime. Of these, the site with the poorest soil (Mishana, on almost pure white sand) has the fewest epiphytes (31 epiphyte species plus a few “‘indets.”’ in a relatively inten- sively inventoried area vs. 38 identified and many unidentified at less intensively studied better-soil Yanamono), suggesting that epiphyte diversity varies with soil fertility as does the diversity of other habit groups. However, to date the sam- 218 pling of epiphytes at these sites has been much less intensive than at our Ecuadorian local florula sites and is probably too haphazard and incom- plete to make these data very meaningful. Data are also available for understory com- position and levels of flowering and fruiting for a broad array of neotropical (and paleotropical) sites (Gentry & Emmons, 1987). If the data for epiphytes are extracted from that data set, a very strong reduction in numbers of fertile species of understory epiphytes on poorer soils is apparent, paralleling the overall trend of decreased num- bers of flowering and fruiting understory Species loss of soil fertility: as soil fertility decreases, terrestrial herbs, epiphytes, understory shrubs, and lianas disappear from the understory in that sequence, leaving virtually only tree saplings and seedlings in the most severely stressed forests (Gentry & Emmon 7). Anecdotal ae also indicates that epi- phytes are much less diverse and abundant on poor soils. We have observed many fewer epi- phytes in poor soil parts of Central Amazonian Brazil, southern Venezuela, and elsewhere in the Guiana shield area than in parts of the Neotrop- ics with richer soils. Large-scale biogeographical analysis also indicates that epiphytic taxa are poorly represented in these areas compared with richer soil areas nearer the Andes and in Central America (Gentry, 1982b). In a somewhat differ- ent context, Janzen (1977) has suggested that the paucity of epiphytes in dipterocarp forests results from the generally nutrient-poor Southeast Asian soils. Among the evidence for this hypothesis cited by Janzen (1977) is the observation that trees cultivated in Malesia along roads, where dust stirred up by passing vehicles increases the nutrients — to oie soe eh in an LEE unusually poor-s large epiphyte loads compared with the native eee On balance it seems clear that the epiphytic plant community is very sensitive to soil fertility, with fewer epiphytes and fewer epiphytic species in forests on poorer soils. Indeed epiphyte di- versity may be even more sensitive to change in soil fertility than is tree or liana diversity, a sug- gestion that would accord with the idea (Gentry & Emmons, 1987) that plants (presumably in- cluding epiphytes despite their lack of direct con- tact with the soil) that are barely able to eke out a marginal existence should be more susceptible ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 to the effects of relatively slight decreases in en- vironmental favorability. LATITUDE It is well known that the presence of vascular epiphytes is a characteristic of tropical forests as compared with temperate ones. We know of only four vascular epiphytes that occur north of Flor- ida in the temperate United States— Tillandsia usneoides, T. recurvata (only in southernmost Arizona), Epidendrum conopseum, and Polypo- dium polypodioides. Even Tillandsia usneoides, the northernmost vascular epiphyte, does not reach the Mason-Dixon line. The only continen- tal United States epiphyte species that is not widespread in the tropics is Tillandsia simulata Sm., endemic to Central Florida but sometimes lümped with 7. bartramii Ell. (The Mexican range of Epidendrum conopseum is also somewhat lim- ited.) Even in very wet areas that would be full of epiphytes in the tropics, only lower plants have adopted the epiphytic habit. In intermediate sub- tropical areas a gradient of increasing epiphytism at lower altitudes is evident. The decrease in vas- cular epiphytes with increasing latitude can be clearly seen in Florida where oe South Florida (latitude 25°N) has 46+ (= 2.8%) epi- phytic Pup (Long & PEA 1971), Central Florida 1 (= 1.9%) (Wunderlin, 1982), and Florida E d State Park in northern Florida (30*50'N latitude) has only two (= 0.4%), Til- landsia usneoides and Polypodium polypodioides (Mitchell, 1963). Curiously, the decrease in vascular epiphytes with increasing latitude is not symmetrical on both sides of the equator. A number of epiphytic species and even a few endemic genera of epi- phytes occur in south temperate forests. Endemic temperate South American epiphytic genera in- clude the monotypic fern Synammia, the mono- typic cactus Pfeiffera, three monotypic Gesne- riaceae genera (Asteranthera, Sarmienta, and facultatively epiphytic Mitraria), and the Lili- aceae (or Philesiaceae) Luzuriaga and Philesia. In the Australasian region, New Zealand is es- pecially noteworthy for its autochthonous epi- phytes including genera like the monotypic fern Anarthropteris, the liliaceous Collospermum (also reaching Fiji and Samoa), the only epiphytic species of families like Cunoniaceae (with two different genera having epiphytic members), and genera like Microlaena (Gramineae) and Metro- sideros (Myrtaceae). There is even a largely epi- phytic south temperate family shared by New 1987] Zealand and Southern Argentina—Chile—Grise- liniaceae (sometimes included in Cornaceae). In the north the only noteworthy temperate epi- phytes are in the Himalayas where aberrant epi- phytic species of otherwise terrestrial genera like Ilex, Tripogon, Euonymous, Sedum, and Tha- lictrum occur. This latitudinal asymmetry was already noted by Schimper (1903), who pointed out that north temperate epiphytes are merely range extensions of widespread tropical species, whereas many unusual and distinctive epiphytic taxa occur in the South Temperate region. At the community level the same trend is ap- parent. For example, Parque Nacional El Rey in Argentina, at 24°45’S latitude, has a species list (L. Malmierca, pers. comm.) of well over 500 vascular plants including 47 species of epiphytes: 20 ferns, four orchids, three species of Rhipsalis and Peperomia, and no fewer than 17 bromeliads including 14 tillandsias. In contrast, Florida Cav- erns ih rig at 30?50'N latitude has only two epiphy its similar-sized flora of 485 native species (Michel 1963). Even in rather dry south temperate vegetations vascular epiphytes can be extremely prevalent, a situation apparently with- out parallel in the North Temperate region. For example, in the Valley of Lerma near Salta, Ar- gentina (1,200 m, ca. 700 mm ppt.), there are at least 14 angiosperm epiphytes including at least ten Tillandsia species in a flora of over 750 species (Novara, 1984). Farther south in the Valdivian region of Chile vascular epiphytes, mostly belonging to endemic genera, are conspicuous elements of local floras, ranging from six species (396 of the native flora) at relatively dry Parque Nacional Tolhuaca (38?15'S) (Ramírez, 1978) to 17 species (17% of the native flora) at very wet Fundo de San Martín (39°30'S) (Cárdenas, 1976; Riveros & Ramirez, 1978); even at 41°S there are 15 vascular epi- phyte species in Puyehue National Park (Munoz, 1980). New Zealand forests have even more epi- phytes than the Chilean ones; even well south of 40°S, about 30 vascular epiphytes are typically included on local species lists (Dawson, 1980). However, at comparable latitudes in North America there are no vascular epiphytes perate forests is unclear but presumably relates to the relatively mesic, more or less oceanic cli- mates that prevail in the Southern Hemisphere. There are more epiphytes at 25°S (47 epiphytes constituting ca. 8.5% of the flora of Parque El GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES 219 Rey, L. Malmierca, pers. comm.) as compared with 25°N (46 epiphytes ~~ ca. 2.8% of the South Florida flora, L & Lakela, 1971). At least in South America, fi prevalence of many species of the genus Tillandsia in southern forests (e.g., 14 at Parque El Rey), perhaps due purely to biohistorical reasons, is another important factor. A similar pattern occurs with lianas, which are better represented in New Zealand tha north temperate forests (Dawson, 1980). CONTINENTAL TRENDS There are several conspicuous differences be- tween the epiphytic floras of different continents. Obviously, predominantly extratropical conti- nents u TN yaaa p However, thin the trop- ics between the Neotropics, tropical Africa, and tropical Australasia. The African epiphytic flora has been widely noted to be very impoverished compared with the other two regions, presum- ably reflecting a loss of mesic-adapted species during the dry periods associated with the Pleis- tocene glacial advances at higher latitudes (Rich- ards, 1973; Madison, 1977). According to Mad- ison there are only ca. 2,400 epiphytic species in Africa, less than a sixth as many as in the Neo- tropics and a quarter as many as in tropical Aus- tralasia. Even though several families and genera with epiphytes in Madagascar were omitted from Madison’s (1977) epiphyte summary (see Ap- pendix), their inclusion does not appreciably in- crease the number of African epiphyte species. Curiously, the depauperate nature of the Af- rican epiphytic flora is not obvious at the com- munity level. For example, the 59 epiphyte species at Makokou, Gabon, constitute 5% of the total Makokou flora (Table 8; compiled from Hladik & Gentry, in prep.; Florence & Hladik, 1980, and included references). While 5% of a moist forest flora might seem fewer epiphytes than would be expected in the Neotropics, Ma- kokou is quite dry, with only 1,785 mm ofannual rainfall, and its 66 (6%) epiphytic species (in- cluding stranglers) are quite in line with the 7 (12%) epiphytic species at Jauneche and 24 (4%) at Santa Rosa (Table 9). Johansson (1974) stud- ied a relatively moist region in the Nimba moun- tains of northern Liberia and reported 153 vas- cular epiphyte species (excluding six filmy ferns and 23 “facultative” epiphytes) in his study area, up to 44 species in a single 750 m plot, and up to 22 species on a single tree. In even wetter areas 220 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 TABLE 8. Habit distributions of Makokou, Gabon plant species. Gymno- Habit Ferns sperms Monocots Dicots Total Epiphytes 26 0 31 2 59 Parasites + saprophytes 0 0 3 6 9 Climbers l 2 8 248 259 Trees 0 0 5 390 395 Herbs, shrubs, and treelets 42 0 125 251 418 Total species 69 2 172 897 1,140 like southwestern Cameroon epiphytes are al- most as prevalent as ls many similar areas of the Neotropics (AG, p obs.). That tropical Heide: is also floristically impoverished with respect to the Neotropics has only recently been realized (Raven, 1976; Gen- try, 1982b). Epiphytes account for much of the overall difference between the two regions with half again as many epiphytes in the Neotropics as in Australasia (15,500 vs. 10,200) according to Madison's (1977) figures. Moreover, at least in those lowland Asian forests we have visited, there also seem to be many fewer epiphytic in- dividuals than in comparable neotropical forests. Richards's (1936) remark is typical: “One of the most striking features of the Sarawak rain forest, especially when compared with that of tropical South America, is the poverty of the epiphytic vegetation both in species and individuals." Madison suggested that the fewer tropical Asian phytes might stem simply from lack of the extensive loud forest habitats of the Neotropics, a 1977) emphasized that m tropical Asian forests tend to have relatively nu- trient-poor soils, and, if so, our suggestion of a positive correlation between epiphytes and soil fertility might help explain the relatively low epi- phyte diversity and biomass in tropical Asia. Another important continental level difference in epiphytes is taxonomic. Orchids and ferns are the predominant vascular epiphytes nearly everywhere, but the other elements of the epi- phytic flora are often very different on different continents (Table 10). Johansson (1974) gener- alized that the African epiphytic flora is made up almost entirely of pteridophytes and orchids, whereas these groups are joined by bromeliads and Cactaceae as important epiphytic taxa in in Australasia and 15 with epiphytes only in the TABLE 9. Representation of different habits in local florulas. Barr Santa Jau- Colo- Río Pa- Capeira osa eche rad lenque Selva! Makokou Habit No. % No. % No. 96 No. % No. % No. % No. 9 Epiphyte (incl. stranglers) 8 2 19 3 72 12 216 16 238 23 368 25 66+ 6+ Parasites + saprophytes 4 l 6 l 4 l 12 1 6 | 8 1l 9 l Cli s 112 24 115 18 136 22 258 20 171 16 182 12 259 23 Trees >10 cm DBH 69 15 142 21 112 19 290 22 165 16 310 21 389 34 Terrestrial herbs, shrubs, tree- lets 270 S8 381 S8 280 47 540 41 475 45 622 42 418 37 Total species 463 667 604 1,316 1,055 1,490 1,140 ' Data from B. Hammel (pers. comm.). 1987] GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES TABLE 10. Taxonomic distribution of epiphytic taxa in some lowland florulas. Ferns Monocots Dicots Total No. No. No No. No. No. Site Spp. 96 Fam. Spp. % Fam Spp. 96 Spp. 96 Neotropics Capeira, Ecu — — 2 7 88 2 2 22 9 2 Santa Rosa, C.R 7 29 2 11 46 3 6 25 24 4 Jauneche, Ecu. 5 7 3 49 68 4 18 25 72 12 Barro Colorado, Pan. 43 20 4 131 61 12 42 19 216 16 Rio Palenque, Ecu. 28 12 4 142 60 14 69 29 238 23 La Selva, C.R 59 16 5 226 61 15 83 23 368 25 Africa Makokou, Gabon 26 44 2 31 53 3 9+ 3 66+ 6+ Asia Flora of Java 200 24 3 520 63 ll 109 13 829 — Neotropics (in the case of Campanulaceae there are also 11 Hawaiian species), Africa has only one family uniquely epiphytic. That family, Cos- taceae, is a dubious segregate of Zingiberaceae, the latter with epiphytes in Asia. At least nine seed plant families have epiphytic species in both the Neotropics and Australasia (but not Africa), but not a single one has epiphytes in both the Neotropics and Africa-Madagascar but not Aus- tralasia. There are 14 families with epiphytic species in all three of the world's main tropical regions. In total there are 33 (or 34 ir pepo is nlv continent as compared with 23 with pins doe on more than one continent: clearly most seed plant families have epiphytes on only a single continent. Many of the “epiphytic” families included in the above analysis are actually terrestrial families with one or two aberrant species adapted to epi- phytism. There are only 32 seed plant families with five or more epiphytic species. If the con- tinental representations of these 32 families are compared, seven (Bromeliaceae, Cyclanthaceae, Rapateaceae, Bignoniaceae, Campanulaceae, Marcgraviaceae, and Guttiferae) have epiphytes only in the Neotropics; six (Myrtaceae, Nepen- thaceae, Pittosporaceae, Loganiaceae, Balsami- naceae, and Zingiberaceae, sensu stricto) have epiphytes only in Australasia; 14 have epiphytes in all three tropical regions; and five (Asclepia- daceae, Ericaceae, Rubiaceae, Solanaceae, and Urticaceae) wi ge a only in the Neotrop- ics and Austr; Put B way, LES are 42 neotropical seed 4 AY J families it eaninhvtec plant families with epiphytes (Table 3) but epi- phytism is minimal in 19 of these (one to four epiphytic species in the Neotropics). Of the fam- ilies with at least five epiphytic species in the Neotropics, seven— Bromeliaceae, Cyclantha- ceae, Rapateaceae, Campanulaceae (also in Ha- waii), Bignoniaceae, Marcgraviaceae, Guttifer- ae—are epiphytic exclusively in the Neotropics, and another, Cactaceae, has only one epiphytic species widespread in the Paleotropics. Fourteen of the families with some neotropical epiphytic species have epiphytes in all three tropical re- gions— Araceae, bi ordi Liliaceae, Aralia- ceae, Begoniaceae, Com sitae, Crassulaceae, Gesneriaceae, Lent tomata- ceae, Moraceae, ME ne Piparace eae (plus, marginally, Cactaceae). In addition to exclusive- ly south temperate Griseliniaceae and Philesi- aceae, seven families—Gnetaceae, Burmanni- aceae, Asclepiadaceae, Ericaceae, Rubiaceae, Solanaceae, Urticaceae—have epiphytes only in the Neotropics and Australasia. The sharing of epiphytic taxa between Austral- asia and the Neotropics but not with Africa is a very different pattern from that normally found giosperms, where close floristic relation- ships between Africa and the Neotropics (reflect- ing a shared early angiosperm Gondwanan flora, Raven & Axelrod, 1974) or between Africa and tropical Asia (reflecting the relatively direct mi- gration route provided by today’s geography) are the general rule. Even within the same family paleotropical and neotropical epiphytes are often not closely relat- ed. For example, the paleotropical epiphytic 222 Gesneriaceae belong to subfamily Cyrtandroi- deae while neotropical ones are mostly in the endemic subfamily Gesnerioideae (Wiehler 1983). Madison (1977) noted that epiphytism has arisen independently in at least three differ- ent groups of aroids. Most epiphytic neotropical orchids belong to subtribes Pleurothallidinae, Maxillarinae, and Oncidinae, while most paleo- tropical ones belong to Dendrobiinae and Bul- bophyllinae. The few epiphytic Central Ameri- can species of Cynanchum are quite unrelated to the many paleotropical epiphytic species of As- clepiadaceae. Most epiphytic neotropical Erica- ceae belong to subfamily Vaccinioideae, half the paleotropical ones to Rhododendroideae. Most epiphytic neotropical Rubiaceae belong to tribe Cinchoneae (subfamily Cinchonioideae), most paleotropical ones to Psychotrieae (subfamily deae). ` Rubioideae th h a ee ven g piphytic sp in all three continental regions, these may not be closely related to each other. Altogether only 30 epiphyte-containing seed plant genera are found in more than one of the three main tropical re- gions, and only 14— Liparis, Malaxis, Vanilla, Polystachya, Bulbophyllum, Schefflera, Begonia, Rhipsalis, Vaccinium, Utricularia, Ficus, Myr- sine, Peperomia, Senecio —are pantropical. Twelve of the genera that occur as epiphytes on more than one continent are large, diverse, ter- restrial genera in which epiphytism has arisen occasionally. For example, there are at least two independent origins of epiphytism in Utricularia (P. Taylor, fide Madison, 1977). Genera like Gnetum, Myrsine, Burmannia, Schefflera, Be- gonia, Senecio, Gaultheria, Vaccinium, Piper, Psychotria, Solanum, and Pilea have indepen- dently derived and mostly quite unrelated epi- phytic species in | Old and New Worlds. There with epi- phytic: species in | Africa and Asia: predominantly epiphytic Medinilla (Melastomataceae) and the orchid genera Acampe, Oberonia, l (M visini naceae). We are oi with only Schefflera, Rhipsalis, Ficus, Peperomia, Liparis, Malaxis, Vanilla, Polystachya, aad Bulbophyllum as gen- era in which epiphytism is widely prevalent on all three continents. Indeed the only genera in which epiphytism in both the New and Old Worlds seems to represent a true synapomorphy are three south temperate genera (Luzuriaga, Griselinia, and Coprosma), Ficus, Rhipsalis, probably Peperomia, and possibly Schefflera. The ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 clearest cases of a shared epiphytic ancestor are Cactaceae where long distance dispersal of one species of Rhipsalis is responsible for the pattern seen today (Barthlott, 1983), and the peculiar case of Ficus where the large successful pantrop- ical subgenus (Urostigma) has specialized as stranglers. The five pantropical epiphytic orchid genera likely achieved their present distributions via long-distance dispersal of their dustlike seeds. Predominantly epiphytic Peperomia may have been originally epiphytic; it is probably ancient (cf. Burger, 1977) and in many aspects of its bi- ology, in addition to its pantropical distribution, it is anomalous among epiphytes. Schefflera is predominantly epiphytic in the Neotropics, predominantly terrestrial in the Pa- leotropics; according to Madison’s (1977) esti- mate only 65 species are epiphytic. Many species grow both as epiphytes and terrestrials; epiphyt- ism, though widespread, does not seem funda- mentally intrinsic to Schefflera and may have arisen independently in all three regions and probably within a given region as we erns, notorious for the ease of long-distance dispersal of their dustlike diaspores, contrast with the angiosperms in having most large epiphytic genera preponderately pantropical. There are even exclusively epiphytic pantropical fern gen- era including Pleopeltis, Platycerium, Ctenopte- ris, Xiphopteris, Psilotum, Vittaria, and (almost) Polypodium, there is not a single exclusively epi- phytic pantropical seed plant genus. Even in ferns the majority of the epiphytic genera are restricted to one geographical region. Only one epiphytic fern genus is disjunct between the Neotropics and Asia (Ophioglossum, a large genus with only two epiphytic species separately arisen on the two continents). Five epiphytic fern genera occur in both tropical Asia and Africa but not the Neo- tropics, three of them on Madagascar but not continental Africa; again this is a pattern without parallel in the seed plants. We may conclude that, except for the ferns, strangler figs, Peperomia, Rhipsalis, and a few orchid genera, the epiphytic floras of the different tropical regions are independently derived, even in most cases where the same family or genus is involved in different regions. While certain taxa are preadapted to an epi- largely on the peculiarities of that region’s evo- lutionary milieu. Epiphytism has arisen very many times in very many groups. However, cer- 1987] tain regions have given rise to many epiphytes; others have not. In this context, the partial ex- planation offered by Madison (1977) for the greater representation of epiphytes in the Neo- tropics—that historical accident in the distri- bution of families like Bromeliaceae, Cactaceae, species—seems largely irrelevant. Indeed, many families (and other supraspecific taxa) have evolved epiphytism in the Paleotropics as in the Neotropics; the difference is that in the Neo- tropics evolutionary experiments with an epi- phytic life-style have subsequently led to much more profuse speciation. Madison (1977) also thought that one element in explaining the con- tinental difference of epiphyte diversity is a pau- city of paleotropical nonorchid monocot epi- phytes. However, our analysis emphasizes that it is not the evolution of epiphytism itself in such taxa that is the critical factor, but rather that there has been little subsequent radiation. Why have epiphytic taxa of Zingiberaceae, Costaceae, Pandanaceae, or Liliaceae not evolved into pa- leotropical versions of Bromeliaceae or Cyclan- thaceae? We will try to analyze why this should be so in the next section. EvoLuTION or EPIPHYTE SPECIES DIVERSITY Why are epiphytes so much better represented in some habitats than in others within the Neo- tropics? Part of the answer to that question can be adduced from the diversity gradients dis- cussed above. From the patterns documented somewhat tenuously established trend toward greater diversity at middle elevations, these are exactly the trends shown by angiosperms in gen- eral (Gentry, 1982a, 1982b, 1987c). Ina previous paper, based on data extrapolated from a large array of published monographs of neotropical taxa, Gentry (1982b) concluded that plant fam- ilies belonging to different habit groups have fun- damentally different distributional patterns. Families composed mostly of canopy trees or lianas have their greatest diversity in Amazonia whereas families made up mostly of epiphytes, shrubs, or palmetto-type herbs are largely extra- Amazonian and are especially concentrated along the lower slopes of the northern Andes and to a GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES 223 lesser extent in southern Central America. For epiphytes this concentration of species diversity could have been predicted from the trends out- lined above. But why are epiphytes (and some other plants) so much more diverse in these re- gions? One reason that epiphytes are especially di- verse in wet aseasonal forests is that they are able to achieve a much finer niche partitioning, and thus a higher alpha diversity, there. Western Ec- uador provides a good example of how this phe- nomenon operates. In the evergreen Rio Pa- lenque wet forest nearly all of the epiphytes have a characteristic and usually very restricted hab- itat, occurring only in the understory, the middle tory, or the canopy. Altogether 41 species of vascular epiphytes at Rio Palenque are under- story specialists: 19 species of Araceae, one of Begoniaceae, one of Bignoniaceae, five of Cy- clanthaceae, eight of Piperaceae, one of Solana- ceae, and six ferns. However, there is not a single understory-specialist epiphyte species in the highly seasonal semideciduous moist forest at Jauneche, only a few tens of kilometers away. The presence of 41 species of understory spe- cialist epiphytes at Rio Palenque accounts for much of the difference between its diverse epi- phytic flora and the relatively depauperate one Nn constant environment at enque rs within-community microhabitat specialization by epiphytes. Thus classical ideas about the greater spatial heterogeneity of everwet tropical forests (e.g., Baker, 1970) are certainly applicable to epiphyte diversity patterns both within the tropics and on a latitudinal gradient. To some extent niche fine-tuning in constant environments also occurs in nonepiphytes. A good example is provided by Gasteranthus at Centinela, Ecuador (see Gentry, 1987b). Six species occur together sympatrically. All are ter- restrial herbs and five are strictly endemic. The Another species, G. crispus, grows only in sandy creek beds of the north part of the ridge and is not strictly sympatric with the other four species. Of the four strictly sympatric species, one, G. atratus, has switched from hummingbird- to bee- pollination and has yellow flowers completely distinct from the orange flowers of the other 224 species. Two of the strictly sympatric species both have large flowers and grow in the deep shade along creek beds, but G. macrocalyx blooms in the wet season and G. pubescens in the dry sea- son. The final species, G. carinatus, is morpho- logically differentiated by slightly smaller flowers and ecologically by growing only along the ridge top. In Gasteranthus, as in the Rio Palenque epi- phytic flora, very fine niche partitioning in a rel- atively constant climate seems an important key to maintenance of high species diversity. To the extent that such niche partitioning is related to equable montane cloud forest conditions, it might be expected to be relatively favored in the Neo- tropics. A second explanation for the great epiphyte diversity in the Andean area focuses on £-di- versity resulting from the greater microsite dif- ferentiation typical of mountainous regions. A test of this hypothesis might come from com- paring epiphyte diversity in areas with high and low microsite differentiation. The two “nudos” where the eastern and western Andean cordil- leras come together briefly at opposite ends of Ecuador provide such a test. The southern Nudo de Loja (or Sabanilla) marks the beginning of the Huancabamba biogeographic discontinuity (cf. Berry, 1982); the northern Nudo de Pasto, most- ly across the border in Colombia, marks the point where the three Colombian cordilleras diverge. n different orientations of adjacent slopes. Teli- pogon (Orchidaceae) provides a good example of how such microsite heterogeneity can multiply epiphyte species diversity. There are 37 Telipo- gon species in Ecuador and five more on the Colombian side of the Nudo de Pasto. There are 14 species on the Nudo de Loja and 16 species on the Nudo de Pasto, as compared with only 14 species in the entire intervening 600 km. More instructive, nine ofthe Telipogon species on Nudo de Loja and five on Nudo de Pasto are locally endemic, each found in a single valley or a single slope. In the much larger intervening area there are only nine endemic species, each restricted to a small area of a few hectares, in several cases representing unusual and restricted microhabi- tats similar to those around the Nudos. It seems clear that in 7elipogon diversity is associated with local speciation and microgeographic spe- cialization. Again, epiphytes are not the only ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 group to show such patterns, but they do seem especially prone to this kind of microgeographic speciation. Probably the association of accen- tuated microgeography with mountainous ter- rain, which is much more prevalent in the Neo- tropics, peu some of the intercontinental diversity differen third, not ucc mutually exclusive, potential explanation for high epiphyte diversity in the northern Andean and southern Central American cloud forests is the “evolutionary ex- plosion" hypothesis advanced by Gentry (1982b) in an attempt to explain why there are so man more neotropical than paleotropical plant species. A relatively small number of genera of epiphytes, understory shrubs, and tto-type h speciated profusely in the northern Andean re- gion, in each case giving rise to very many locally endemic and often rather poorly differentiated species. Gentry (1982b: 587) interpreted the high local endemism (cf. Gentry, 1987b) and apparent plastic genera as An m, Piper, aven- dishia as reflecting Hed balance" founde effect phenomena 977. Templeton, 1980) with major genetic reorganizations or ge- netic transilience (Templeton, 1980) optimized by small and localized populations and by the need for constant in a habitat par- titioned by mountains, local rainshadows, ver- tically shifting cyclically coalescing vegetational zones, and frequent landslides. If genetic founder effects associated with recolonization of the open areas resulting from landslides or unusually fre- quent tree falls in these geologically and ecolog- ically dynamic regions are major determinants of speciation events, then much speciation in Andean-centered taxa could well be essentially sympatric and largely random. This model thus differs from the **microgeographic speciation” model in that speciation could take place at a much finer “sympatric” scale, e.g., as coloniza- tion of a specific landslide, with many of the resultant species ecologically indistinguishable from each other, rather than each adapted to a mism imputed to be associated with tropical mountains it would be expected to be most ap- plicable in the Andean region, by far the most extensive mountain system in the world's trop- cs. We have indirect but highly suggestive evi- dence for speciation associated with genetic tran- m 1987] GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES Stanhopea jenishiana Reichb. f. FIGURE 5. silience in founder populations in several genera of epiphytes. Embrya rodigasiana (Cogn.) Dod- son provides a good example of a founder event. This species is distributed mostly in western Co- lombia, between 1,000 m and 1,500 m, ranging from west of Medellin to near Buenaventura. There is also a disjunct population in a small area of southern Ecuador near Pangui, at 1,500 m above Bomboiza and Gualaquiza; prior to re- cent widespread habitat destruction, the species was very common in the disjunct and geograph- ically very limited Ecuadorian part of its range, where it was surely introduced by a long-distance dispersal founder event. Masdevallia chonta- lensis is a similar example. Well known and oc- curring in a well-known habitat, it ranges from Guatemala to Panama with a small disjunct pop- ulation near Piñas in southern Ecuador. There are several similar cases of unsuccessful founder events by dust-seeded tropical epiphytes that have temporarily established disjunct populations in Florida, been duly recorded as members of the Florida flora, and then subsequently disap- peared. An example is Leochilus labiatus which grew for a while near Fackahatchee. The Stanhopea jenishiana complex, well char- acterized by a suite of distinctive morphological characters, provides an example of the next step in such a process. Stanhopea jenishiana Reichb. f. (Fig. 5) ranges from Cali to Popayan in the Cauca Valley of Colombia, with a disjunct pop- ulation near Piñas in southern Ecuador. There are two other small geographically isolated pop- ulations of this complex in western Ecuador, each of them specifically distinct but clearly derived from S. jenishiana and each likely resulting from a single long-distance dispersal event— Stanho- pea frymirei Dodson is endemic to a range of relatively moist hilltops near the coast in Manabi and Guayas provinces and S. embreei Dodson occurs only around 1,000 m in the Bolivar-Canar border area (Fig. 6). he herbaceous and shrubby taxa that show such patterns have relatively short generation times, providing conditions appropriate for rap- id evolutionary diversification. Under either the microgeographic or founder effect hypothesis, epiphytes, as the major herbaceous component of wet tropical forests, might be expected to show unusually rapid “evolutionary explosion" type speciation. Relatively specific pollination sys- tems constitute a second factor wansi bl Gentry (1982b) as potentially promoting un ally rapid speciation in **Andean-centered" ie with shifts in specific pollinators accompanie by coevolutionary fine-tuning of precise plant- pollinator systems apparently a common evo- lutionary theme. Again epiphytes, often char- acterized by high pollinator specificity, should be prime candidates for rapid speciation. The Stanhopea jenishiana complex (see above) is a good example of how founder events and N N ON ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURE 6. Two localized derivatives of Stanhopea jenishiana.—A. S. embreei Dodson.—B. S. frymirei odson shifts in pollination syndromes can combine to give rise to new species. The presumed ancestral species, S. jenishiana, has orange flowers and is pollinated by bees of the genus Euglossa while both of the locally endemic derivative species in central Ecuador have white (E. embreei) or straw- colored (E. frymirei) flowers and have switched to pollination by Eulaema bomboides. All three species have methyl cinnamate as the major scent attractant but each has a different set of scent- modifying compounds. Eulaema bomboides, a very effective pollinator that specializes on flow- ers producing methyl cinnamate, is endemic to central Ecuador. We interpret this situation as reflecting three different instances of long-dis- tance dispersal. In the southern Ecuador Pinas population, outside the range of E. bomboides, pollination by Euglossa was maintained and no vantage of Eulaema bomboides as a pollinator occurred with essentially the same selection op- erating in these two different founder popula- tions. Different evolutionary solutions reflected by the two distinct derivative species resulted. Scelochilus, another epiphytic genus of orchid, provides an even more intriguing indication of how rapid speciation might be in such taxa. Sce- lochilus is a genus of 34 species found mostly epiphytic in guava trees; sparsely represented in natural vegetations, it was apparently ideally pre- adapted for the special conditions provided by guava plantations. Fifteen species of Scelochilus occur in Ecuador. In 1957 one of us (CD) made an intensive study of populations of Scelochilus in an extensive guava grove near the edge of wet forest at 1,000 m altitude at km. 94 of the old Guayaquil-Cuenca road in Ecuador. Two un- described species of Scelochilus were present, ost diues ni but their flowers, though some- hat variable in each species, were very distinc- tive with no overlap whatsoever between the two species. Indeed the original study, intended to focus on hybrid introgression, had to be aban- doned because the two taxa proved to be so con- sistently differentiated. These species were de- scribed as SS. frymirei and S. embreei (Fig. 7). Fifteen years later, in idis a return visit was made to the ve. In the intervening years most of the itin Wasa had been cut and converted to pasture. As a result the habitat was 1987] very different, with a much dryer aspect and the remaining guava trees rather old and decrepit. Scelochilus was much rarer but ca. 50 plants were located in the remaining guavas. Incredibly, nei- ther S. embreei nor S. frymirei was present in 1982, but rather two different new species were found, later described as S. gentryi and S. ro- mansii. As in 1957, both of these species, veg- etatively indistinguishable, were clearly differ- entiated from each other by floral characters (Fig. 7). Both of the species present in 1982 are closely related to S. frymirei. We suspect that they may represent in situ speciation events, at least in the case of S. romansii (S. gentryi has also been found in several other localities in western Ecuador). If so, natural speciation in Scelochilus can occur in as little as 15 years! An obvious alternative interpretation is that specific limits in Scelochilus are too finely drawn, with S. gentryi and S. romansii representing part of the intraspecific variability within polytypic S. frymirei. However, this possibility seems ob- viated by the fact that the two co-occurring Sce- lochilus species of 1982 pass the test of sympatry as biologically differentiated populations, even though they are more similar to each other than to their putative ancestor S. frymirei. We favor the interpretation that the kind of genetic reor- ganization that reflects speciation in Scelochilus, a ma nonorchid epiphytes, is so labile that it can be effected in incredibly short times. If this inter- pretation is correct, then it is no wonder that many epiphytic taxa have undergone what ap- pears to be truly explosive speciation in the Neo- tropics. Obviously, it is also possible that S. gen- tryi and S. romansii immigrated to the guava grove in question sometime between 1957 and 1982 with S. frymirei and S. embryi coinciden- tally becoming locally extinct during the same time interval. But the mere fact that natural spe- ciation in fifteen years seems an equally plausible explanation for these observations is surely sig- nificant. Another line of reasoning also supports the idea that certain neotropical epiphytes have undergone explosive evolution. It is probably safe to assume that, as a general rule and despite many potential exceptions, genera with many species are those which have, on the average, undergone the most rapid speciation, Thus an examination of the largest epip might indicate some a Sr ponds, both geographic and nop, th Table 11 GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES 227 lists the 47 largest genera of vascular epiphytes worldwide (those with 90 or more epiphytic species) and provides the data for such an anal- ysis. Again orchids are preeminent. Half (22) of the 47 largest epiphyte genera are orchids. While orchids might appear to be exceptional in their unusual genetic plasticity and highly specific pol- lination systems, they must unavoidably be re- garded as the most successful practitioners of the art of being epiphytic. Biogeographical analysis T t number of large neotropical and See ses orchid genera is the same, there is a dramatic difference in the number of species that they con- tain. The nine neotropical genera are far larger, accounting for 5,240 species (average 582 spp. per genus) vs. 2,626 a (average 263) for the ten paleotropical on erns account for pum of the largest wa d genera. Not surprisingly, in view of their dia- spore vagility, nearly all of the large IE. fern genera are pantropical, showing little evi- dence of unusually rapid differentiation or spe- ciation in the Neotropics. Only one fern genus, Pyrrhosia, is restricted to the Paleotropics, none to the Neotropics. The remaining 15 of the largest epiphytic gen- era are split evenly between dicots and mono- cots. The two largest dicot genera, Peperomia and Ficus, are pantropical, although the former, at least, is better represented in the Neotropics; the other largest epiphytic dicot genera are two neotropical gesneriads (Drymonia and Colum- nea, sensu lato), two ericads (Rhododendron and Cavendishia), one each epiphytic in Old and New Worlds, and two melastome genera (Medinilla and Blakea), restricted respectively to the Old and New Worlds (ane extremely similar to each other as wellast Topobea). In ganar: here seems a reasonable numerical balance between Old and New World representation in the largest dicot epiphyte gen- era, except in Gesneriaceae. The situation is very different among the nonorchid monocot epi- phytes. Six of the seven largest genera— Anthur- ium, Tillandsia, Philodendron, Vriesia, Guz- mania, an mea —are exclusivel neotropical; only Rhaphidophora is paleotropi- cal. Thus the very many species that have evolved among certain monocot epiphyte genera seem a peculiarity of the Neotropics. Why have epiphytes been especially suscep- 228 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 SCELOCHILUS EMBREEI Dodson SCELOCHILUS FRYMIREI Dodson ]— SCELOCHILUS GENTRYI Dodson SCELOCHILUS ROMANSII Dodson & Garay 1987] tible to the rapid, even explosive, speciation that seems to have characterized a large element of the neotropical flora? At least in the case of or- chids, unusually specific pollination systems have clearly played a major role in making possible very active speciation. Like the orchids, the non- orchid epiphyte families that have speciated the most profusely in the northern Andes and south- ern Central America—Gesneriaceae, Bromeli- aceae, Ericaceae, and Araceae—all share, to a greater or lesser extent, relatively specialized pol- linators and specific palliation systems as com- pared with the “average” tree, shrub, or free- climbing liana (e.g., prevalent hummingbird or euglossine pollination). On the other hand the epiphytic taxa that have speciated most profusely are characterized, as a group, by more general- ized, higher risk dispersal strategies than typical optimal genetic transilience related to multiply replicated founder events in a dynamic and kal- eidoscopically changing habitat as Gentry (1982b) suggested, then epiphytes, characterized by their unique combination of r- and k-selected repro- ductive traits, might be uniquely equipped to react to this adaptive milieu. Like weeds, they have diaspores intrinsically adapted for quick colonization of newly available habitats. In epi- phytes, like weeds, the primary need for such adaptation is presumably a response to the nd unstable nature of their normal adapting epiphytes for rapid speciation in an environment in which the need to recolonize dynamically changing microhabits is unusually frequent. However, unlike weeds whose di- versification is generally constrained by overly generalized pollination syndromes (frequently even with loss of sexual recombination), the rich array of specialized pollination systems that characterizes the successful epiphytic taxa would seem to preadapt them for rapid, pollinator-re- lated evolutionary diversification. In this con- text, it is not likely to be an accident that the preeminently successful epiphytic family, Or- GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES 229 chidaceae, is precisely that family that has both the most specific pollination systems and the tiniest, most r-selected diaspores of any angio- sperm. Moreover, from the viewpoint of “‘acci- dental” speciation via founder effect phenome- na, the unusual genetic similarity among the potential-colonizer orchid propagules that arrive together at a specific site—due to orchid seeds’ unique sharing of fathers as well as mothers, thanks to their pollinia—might be expected to accentuate the potential for genetic drift in col- ations. Interestingly, the closest a o Andean cloud forest epiphytes is found in baceous and palmetto taxa that most closely ap- proximate the unique epiphyte combination of specific pollination systems and an r-selected high-risk dispersal mode. CONCLUSION Although many unrelated kinds of plants have evolved epiphytic habits, most of these represent isolated species or genera in otherwise terrestrial families. Only three nonfern families — Orchi- daceae, Cyclanthaceae, Marcgraviaceae (and possibly also Bromeliaceae) — are predominantly Orchidaceae, Bromeliaceae, Polypodiaceae, and Araceae—and 8996 in eight families. Very few families have been successful at undergoing ex- tensive radiation as epiphytes. Indeed over two- thirds of all epiphyte species belong to the single family Orchidaceae, and to a large degree un- derstanding epiphytic diversity is synonymous with understanding orchids. Although notably few families have been very in a local flora or 6396 of the individual plants in a given sample area. Epiphytes have speciated most profusely in the Neotropics, especially in northwest South Amer- 7. Postulated rapid KT in Scelochilus. Top two species bee described as new from 1 FIGURE 7. tibns! ed guava grove at two species, both ne Plantarum “Tropic carum. of Guayaquil-Cuenca road. In 19 and both vend d to S. frymirei. yr reer ie dh Orchids of Ecuador, Icones ) 957 y had been replaced by bottom 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 TABLE 11. Major epiphyte genera (in part after Madison, 1977; Dressler, 1981; Kress, 1986). Genus No. Epiphytic Spp. Total No. Spp. Distribution Pleurothallis 1,520 1,600 Neotropical Bulbophyllum 1,000 1,000 Pantropical endrobiu 900 900 Australasian Epidendrum 720 800+ Neotropical Peperomia 700 1,000 Pantropical Anthurium' 600 850 Neotropical Lepanthes 600 600 Neotropical elis 540 600 Neotropical Ficus 500 800 Pantropical Maxillaria 570 600 Neotropical Eria 500 550 ustralasian Oncidium 475 500 Neotropical splenium 400 650 Pantropical Tillandsia 400 450 Neotropical Grammitis 400 400 Pantropical Philodendron' 350 475 Neotropical Masdevallia 400 400 Neotropical Medinilla 300+ 400+ Paleotropical Liparis 300 350 osmo an Oberonia 300 300 Paleotropical Odontoglossum 285 300 Neotropical Columnea (sensu lato) 262 265 Neotropical Elaphoglos. 250 500 Pantropical Hymenophyllum 250 300 Pantropical Angraecum 206 206 Trop. A Lycopodiu 200 400 Cosmopolitan Polystachya 200 210 Pantropical Ctenopteris 200 200 Pantropical riesia 200 260 Neotropical Phreatia 190 190 Australasian Trichomanes 150 300 Pantropical Polypodiu 140 150 Pantropical Encyclia 130 130 Neotropical Octomeria 130 130 Neotropical Taeniophyllum 120 120 Paleotropical, esp. Afr. uzmania 120 140 Neotropical Dendrochilum 120 120 Australasia Aechmea 120 150 Neotropical Rhododendron 112 850 ostly Asian Coelogyne 100 100 rop. Asia monia 100 110 opical Pyrrosia 100 100 Paleotropical Rhaphidophora 100 100 Paleotropical Appendicula 100 100 Australasia Blakea? 98 100 Neotropical Thrixspermum 90 100 ustralasia Cavendishia? 90 100 Neotropical ! Croat, pers. comm. ? Renner, 1986 and pers. comm. ? Luteyn, pers. comm. 1987] ica and southern Central America. The explosive speciation of relatively few epiphyte genera in this region has been responsible to a large extent for the excess floristic diversity of the Neotropics as compared with the Paleotropics. 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Selbyana 9: 104-111. RICHARDS, P. W. Ecological observations on ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 the rain forest of Mt. Dulit, Sarawak. J. Ecol. 24: Part I: 1-37; Part II: 340-360. 1957. The Tropical Rain Forest. Cambridge Univ. Press (reprint). 1973. Africa, the “Odd Man Out.” Pp. 21- n B. Meggars et al. (editors), Tropical Forest patent mes in Africa and South America: A Com- parative Review. Smithsonian Inst. Press, Wash- ington, RIVEROS, M. & C. RAMÍREZ. 1978. Fitocenosis epi- fitas de la asociación Lapagerio-Aextoxiconetum en el fundo San Martín (Valdivia-Chile). Act. Cient. Botany and Natural History of Panama. Missouri Botanical Garden, St. Louis. ipte A. F. W. 1888. Die Er E Vegeta- on Amerikas. Bot. Mitt. Trop. 2. . 1903. Plant- r b. E a Physiological Basis. Clarendon Press, Oxford. [English transla- tion, revised edition. Scorr, N. J., JR. 1976. The abundance and diversity of the nA of tropical forest litter. Bio- tropica 8: 41-58. ii y DE LA. D Las pteridófitas y el epifitismo el Departamento del Chocó (Colombia). Ann. So oc. Cient. Arg. 194: 245-278. SriLES, G. 1981. RU bi dien aspects of bird-flower coevolution, with special reference to Central America. Ann. Missouri Hot Gard. 68: 323-351. TEMPLETON, A. R. 1980. The theory of speciation via the founder principle. Genetics 94: 1011-1038. VAN DER PIJL, L. & C. DODSON. 1966. Orchid Flowers: Their Pollination and Evolution. Univ. Miami Press, Coral Gables, Florida. 85. Vegetation of the Earth, 3rd edi- tion. Springer-Verlag, Berlin. Tropical Rain Forests of the Far East, 2nd edition. Clarendon Press, Oxford. WIEHLER, H. 1983. A synopsis ofthe neotropical Ges- neriaceae. Selbyana 6: 1-219. ee and the Genetics of Pop- ulations. Volum Experimental Results and Evolutionary Dunt Univ. Chicago Press, Chicago. WUNDERLIN, R. 1982. Guide to the Vascular Plants of Central Florida. Univ. Florida Press, Gaines- ville 1987] GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES 233 APPENDIX I. Changes in data base from epiphyte list in Madison (1977). Additions 450) ,200 species, not 850) Anthurium (600/850) Philodendron (350/475) Cyclanthaceae (125 species, not 31) Dicranopygium (few/44) Evodianthus (1/1) Stelestylis (4/4) Dj coreacea Dioscorea (1/600) (Ecuador, pers. obs.) Liliaceae Rhodocodon (1/8) (Madagascar, pers. obs.) Philesiaceae Philesia (1/1) (Chile, pers. obs.) Luzuriaga (3/3) (Chile, pers. obs.) Alzateaceae Alzatea (1/2) Anacardiaceae iini af 10) (SE Asia, fide Kress) Apocy Mandevilla (1/114) (M. pittieri, Costa Rica) Begon Begonia K 1,000) Bignoniacea Gibs oniothamnus (11/11) Schlegelia (18 ombacaceae Spirotheca (4/4) seraceae MN (1/80) (B. standleyana, Costa Rica) fen eal (ca. 30 epiphytic species, not 3) bum !GSinclairia) (2/90) Mikani desde da (20/24) Nelso po (1/1) Pseudogynoxy. 1) — (Pentacalia) b l p (epiphytic in Neo- s, Madagasca URN ylis (2/2) Crassulaceae Kalanchoé epiphytic in Madagascar, pers. obs. Marcgraviaceae (89 epiphytic species, not 94) Marcgraviastrum (10/15 Marcgravia (ca. 50—55/55) Melastomataceae (567 epiphytic species, not 483) The paleotropical genera Baekeria, Dalenia, Dic- cochaeta, Neodissochaeta, Om phalop us, and Plechiandra each has at least one hemiepiphytic species fide Renner, 1986. yricaceae Myrica (1/35) (SE Asia, fide Kress, pers. comm.) apotaceae xri (Costa Rica, fide Kress, pers. comm.) Saxifrag; H ydrangea (2/80) (epiphytic in Neotropics) Solana fapa (also epiphytic in Neotropics) Deletions Apostasiaceae = Orchidaceae pou = oe Bromelia —Ana Cycla <” pei Ci arian Balsaminaceae — rubr eod Qm epiphytic in Neo- ignoniaceae — Radermachera Gutt viiei -Clusia ari occurs in Neotropics Marcgraviaceae — Car Araliaceae — eade a inr Schefflera Note: Additional occasional or sporadic epiphytes i in- clude s Kress (e.g Cyperu Pseudoeverardia (= Everardia), Arenaria, E ora, Maranta, Pourouma) are also excluded. NITROGEN FIXATION BY EPIPHYLLS IN A TROPICAL RAINFOREST! BARBARA L. BENTLEY? ABSTRACT Bluegreen 5 (Cyanobacteria) growing on the leaf surfaces of understory plants in a tropical rainforest can ospheric n nitrogen. Rates of fixation are strongly influenced by the presence of glucose and pied: nutrients leached from the host leaf, by light intensity as it relates to the pho- tosynthetic rates of the algae, and by desiccation especially as it is influenced by the co-occurrence of epiphyllous bryophytes. A signific ant portion of this newly fixed nitrogen is transferred to the host leaf and may account for 10-25% of the total nitrogen in a leaf. Although the major source of nitrogen for plant growth and reproduction is from decomposition of organic matter, most ecosystems can gain sig- nificant quantities from precipitation and bio- logical fixation of atmospheric nitrogen. In fact, Burns & Hardy (1975) estimated that biological fixation alone may account for 4396 of the nitro- gen transferred worldwide. Subba Rao (1977) suggested that the rates in tropical areas may be even higher and thus fixation could have a major role in the tropics where available nitrogen can be limited. In this paper, I will discuss one aspect of ni- trogen fixation in a tropical ecosystem — that of fixation by epiphylls in a rainforest understory. Although epiphyll fixation can account for up to 2596 of the nitrogen in a host leaf (Bentley & Carpenter, 1980), rates of fixation are quite vari- able in both time and space. Here I document some of the environmental factors that influence fixation rates. What are epiphylls? Epiphylls are epiphytes that are restricted to the surfaces of leaves. In tropical rainforests, most visible epiphylls are leafy liverworts of the family Lejeuneaceae but may also include mosses, lichens and, on some occasions, seedlings of other epiphytes such as bromeliads and orchids. In this study, we focused on the microorganisms, especially the bluegreen algae (Cyanobacteria) growing in association with the visible forms (Fig. 1). The epiphyll community is best developed in regions of high rainfall and low evaporation and is most diverse in tropical rainforests (Richards, ! [ would first like to express my appreciation to Maureen Dunn who gave expert HIIS e in both n Mei obert Bl 80-10676 from the National Science Foundat 1964). Because they grow on photosynthetic sur- faces, epiphylls are often considered to be det- rimental to the host plant by interfering with light penetration to the leaf (Richards, 1964). Indeed, some epiphylls may actually be semiparasitic. For example, the rhizoids of Radula flaccida, an epiphyllous liverwort, penetrate the cuticle and absorb nutrients from their host leaf (Berrie & Eze, 1975). Other workers feel that the presence of a phylls increases the time that leaves remain w and thus contributes to the growth of omens pathogenic bacteria and fungi (Gregory, 1971; Stahl, 1891). Long- term wetting may alse: seduce the rates of and subsequent mineral uptake by the roots (Stahl, 1893; McLean, 1919). As early as 1891, Jungner suggested that rain- forest plants may have adaptations such as leaf “drip tips" to increase the rates of drainage off the leaf and reduce the rate of colonization by epiphylls. Subsequent work has failed to either deny or confirm the adaptive role of these char- acters (Stahl, 1893; Shreve, 1914; Seybold, 1957; and pers. obs.). Nitrogen fixation by epiphylls. Some species Biological Station in Costa Rica, we found that fixation rates are extremely variable, both within and among species (Fig. 2). Interestingly, in con- trast to Ruinen's data where fixation was by free- living bacteria (primarily Beijerinckia), fixation in the La Selva understory was most commonly Black, Young for assistance in the field. = research was supported by grants DEB 78- 12032 and DEB ? State University of New York, Stony Brook. New York 11794, U.S.A. ANN. Missouni Bor. GARD. 74: 234-241. 1987. 1987] BENTLEY — NITROGEN FIXATION BY EPIPHYLLS Photomicrograph of epiphylls. The strand of the nitrogen- rem bluegreen dni is across the lower h. ue of n photograp associated with the presence of bluegreen algae in the genera Scytonema, Stigonema, and Hap- alosiphon. High fixation rates were invariably associated with a dense cover of bryophytes, sug- gesting that the bryophytes provide a good sub- strate for the nitrogen-fixing microorganisms. Most of our work on fixation was done using the acetylene reduction method for determining nitrogenase activity (Bentley & Carpenter, 1980; Prestwich & Bentley, 1981; Burris, 1972). While this is an extremely easy field assay for estimating fixation, it is not a direct measure of nitrogen fixation, and cannot be used to answer the most critical question in our study: does the newly fixed nitrogen get into the host leaf? By using '°N as a tracer, we were able to document that indeed this is the case (Bentley & Carpenter, 1984). There we showed that nitrogen fixation by epiphylls could account for up to 25% of the nitrogen in a host leaf. In other studies with bluegreen algae, Stewart (1963) and Jones & Stewart (1969) found that the extracellular a products are pri- marily a eptides, but less complex compounds, E ammonium nitrite and nitrate, can be present. The pathway for move- ment of the new nitrogen is not through the sto- mata, as might be assumed, but rather to the epidermal cells via threadlike ectodesmata pen- (Franke, phology of the ectodesmata, Franke felt that the movement of soluble materials into and out of the leaf is a normal process, iic de orrelated with foliar absorption of “foreign” substances such as fertilizers and dr Environmental factors affecting fixation. The rates of fixation and transfer that we measured were made under ideal conditions for the activ- ities of microorganisms. Although the high tem- d almost constant moisture in a trop- ical rainforest can permit high fixation rates and concomitant release of nitrogenous products, we have also observed extremely high variance both among and within species at the La Selva Station (Fig. 2). Thus, it becomes important to ask what environmental factors influence fixation rates by epiphylls. Basically, the answer lies in three fac- tors: time, moisture, and nutrients. e o T ng N FIXED/ hr X+S | | PP, SPECIES OF HOST RE 2. Nitrogen fixation rates by epiphylls on t. The host species, from highest to lowest fixation rates, are: Ocotea atirrhensis, Proteum pittieri, Chamaedorea sp., Ficus sp., Swartzea sp., Miconia sp., Piper sp. 1, sp., Costus sp., Syngonium 1 SP., Asterogyne sp., and Geonoma s Time. Colonization by epiphylls is a time phenomenon. Young leaves are epiphyll-free simply because the epiphylls have not hann established. Long-lived leaves have greater probability of having a well- tablished epiphyll community. To document the role of leaf longevity on fixation, we measured fixation on leaves which had been previously marked for up to 24 months (Bentley, 1979). Since we knew the average survivorship of leaves on a plant, we could then test for the correlation between the age of a leaf and the rate of fixation. As can be seen in Figure 3, fixation rates are significantly correlated with leaf longevity. At the same time, we measured fixation by leaves with differing levels of epiphyll biomass. Again, there is a sig- nificant correlation (Fig. 4). Moisture. Interestingly, in the experiments described above, the rates of fixation were cor- related with the biomass of the bryophytes, even ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 though bryophytes per se do not fix nitrogen. On the other hand, bryophytes are correlated with and can influence the levels of moisture on the leaf surface (Richards, 1964). Since moisture can affect fixation rates in other systems (Balandreau et al., 1974; Horne, 1972; Fogg et al., 1973), we set up an experiment to test the effects of des- iccation on fixation by epiphylls. In this case, individual pinnae of Welfia georgii were moist- ened with water 12, 4, 2, or 0 hours before in- cubation. As can be seen in Figure 5, desiccation has a very dramatic effect on fixation, yet the recovery is quite rapid and is almost that of the continuously moist control within four hours. Although moisture conditions in a tropical rainforest are relatively constant compared with most other ecosystems, moisture conditions in the understory do vary, both seasonally and diur- nally (Longman & Jenik, 1974; Ruinen, 1961). For example, water may continue to drip down through the forest canopy long after a rain has ceased, but the leaves of the upper canopy will dry within minutes. Seasonal variation can range p an increased frequency of dry periods in a day days or weeks without any appreciable Rm (Schnell, 1971). Thus, under field con- ditions, moisture can have significant effects. We documented this in another experiment, shown in Figure 6. In this case we found that fixation rates on rainy days y higher than on dry days. Although it is tempting to suggest that the bluegreens and the bryophytes have some close symbiotic relationship, the role of bryophytes is probably no more than to maintain appropriate moisture conditions on the surface of the leaf. Epiphyll-laden leaves dry more slowly, not only because capillary action holds water but also be- cause bryophytes can draw water directly from the interior of the leaf (Berrie & Eze, 1975). In addition, the epiphyll-laden portion ofa leaf sur- face will become wetter faster, again because of capillary action drawing water from surrounding areas. High fixation rates by algae are correlated with the Presente ad bryophytes simply because the bryophytes are appropriate for the growth and activity of microorganisms. 2A Energy sources. Nitrogen fixation consumes energy — free-living microorganisms may require 200 g of glucose for every gram of nitrogen fixed (Mulder, 1975). Unlike other microorganisms, bluegreen can use carbohydrates from photosyn- 1987] BENTLEY — NITROGEN FIXATION BY EPIPHYLLS 237 25- i = 20r ° hom O uj >< TN I5} Z 4 IOF Ix ° " 5r 3 1 1 l l L L l 1 À À J O 20 40 60 80 100 % MARKED LEAVES REMAINING AFTER 24 MONTHS E 3. Nitrogen fixation rates by epiphylls on n with different leaf longevities. Highest rates are on IG Pa unit in which all leaves live longer t months. Low ates were on leaves of Piper (80% est r: last less than two years). The rank correlation of on with longevity is significant (r = 0.95, P < 0.01). thesis as well as exogenous carbon to supply the nitrogen fixation process. In addition, bluegreens may be able to use intermediate products of pho- tosynthesis to provide electrons for nitrogen fix- ation (Mulder, 1975). T T T T T T T T e 80} = ° E o 60 . =] bo. a "ü 4 uj x u 40r " 5 z ° o pe ia € eo ° ° 20+ 4 ° ° 7 L- e° e 4 ee È Er á oY s * ç, * s f 1 1 0.02 0.04 0.06 O. 0.10 ORY WEIGHT OF EPIPHYLLS Th 1 à 4,5 c 1 111 m tion and biomass is significant at the P < 0.05 level (r = 0.39). In the field, nitrogen fixation by bluegreen al- gae is usually light dependent (Henricksson & Simu, 1971; Horne & Viner, 1971; Stewart, 1973; and see Fig. 6) and has the same relationship to light as photosynthesis—decreasing with de- creasing light intensity. However, low rates of fixation can occur in the dark (Forman, 1975; Dugdale & Dugdale, 1962), especially if the or- ganisms are grown on substrates containing glu- cose, fructose, or sucrose (Fay, 1965). Thus, many workers feel that high rates of fixation under nat- ural conditions are a function of both photo- mediated internal processes as well as various forms of exogenous carbohydrates Exogenous carbohydrates may come from a wide variety of sources. The major source for epiphylls is in leachate from the host leaf (Tukey & Morgan, 1962; Tukey, 1971). Since the amount of carbohydrate present in leachate is also a func- tion of the photosynthetic rate of the host leaf (Tukey et al., 1957), the amount of exogenous carbohydrate available to the epiphylls will also be correlated with light intensity in the field. To document the effects of light on the epiphyll system, we set up two series of experiments: the first to establish that levels of light commonly found in the rainforest understory could influ- 238 eor £ N O W x u z e 10h [ —1 0/12 12/0 10/2 8/4 hrs DRY/hrs WET FIGURE 5. The effects of desiccation on nitrogen fixation by epiphylls. Leaflets from the same pinna of georgii were either kept constantly wet (0 hr. dry; 12 hr. wet), or were subjected to varying lengths of drying (12 hr. dry, 0 hr. wet; 10 hr. dry, 2 hr. wet; or 8 hr. dry, 4 hr. wet) ence fixation rates, and the second to differentiate between direct effects of light and effects of ex- ogenous glucose on fixation rates by the blue- greens. In the first set of experiments, samples were incubated at three levels of light set to mim- ic those found in the forest: over epiphyll-laden leaves (12 uEinsteins m °), in a light gap (68 uEinsteins m ’), and in the dark. As can be seen in Figure 7, fixation was highest at the highest light intensity and was still significant even at the usual light intensity for most epiphylls In the second set of experiments, we measured nitrogen fixation by leaf samples that had been dipped in a p 1% glucose solution, a concentra- tion chosen imic concentrations we mea- sured in leaf s (Tukey et al., 1957). Again, we incubated the samples at two light intensities. In this case, we were able to separate out the effects of light from the effects of exogenous glu- cose. Note in Figure 8 that fixation is highest in those samples which were both dipped in the glucose solution and incubated in the light. The lowest rates were for the control samples dipped in water and incubated in the dark. Note, how- ever, that fixation by the samples dipped in the sugar solution but incubated in the dark contin- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 T T cs. EE 425 ° ° 420 ~ N E = ü < x i 2 5 > x Ë * 2 š 2 š 0 z = T ° =] T T Y ME AM6 7 8 9 10 Il 12 | 2 3 4 5 6PM TIME OF DAY T T T T T T T T T T E b " dd 50 # 425 3 bi ` 40} 420 “E = ` = ul à a x B 4 > K 30 5r z $ Ë c yr z 20r K 40 Š x a lor 5 e AM6 7 8 9 10 I| 12 | 2 3 4 5 6PM TIME OF DAY FIGURE 6. The effects of light intensity on nitrogen Natural light intensities were abnormally high because of the lack of cloud cove ued to fix nitrogen at a reasonable rate for at least eight hours. These results document that fixation by epiphylls is influenced by both light directly and by the presence of exogenous glucose. And perhaps as important, they document that fixa- tion can continue at night if exogenous glucose is present on the leaf surface. Mineral nutrients. Growth of nitrogen-fixing microorganisms may also be limited by avail- ability of mineral nutrients—most commonly “upi phosphorus, calcium, molybdenum, and iron (Mulder, 1975; Fogg, 1973; Mague, 977. Deficiencies in any of these can have an 1987] 150r IOOF nMOLES CH, /lO cm* LEAF L 1 L 1 1 L M sub: 1 L... 2 4 6 8 10 l2 I4 16 I8 20 22 24 TIME (hr) FiGuRE 7. Experimental variation of light intensi- ties. Fixation rates are given in terms of nM ethylene to illustrate continuing fixation throughout the 24 hours of the experiment. Ethylene concentrations of samples incubated in the dark do not change after 12 hours. indirect effect on nitrogen fixation because each is involved in general metabolic activities. In addition, Mo and Fe are components in the ni- trogenase molecule. Deficiencies in either of these reduces rates of enzyme synthesis and thus di- rectly affect fixation (Burns & Hardy, 1975). Exogenous combined nitrogen, most notably NH,*, will also inhibit nitrogen fixation (Stew- art, 1973; Burns & Hardy, 1975; Fogg, 1973). Again, the effect is the result of reduced nitroge- nase synthesis. Interestingly, the effects of com- bined nitrogen may explain why fixation rates are low in nongrowing cells: as growth slows down, nitrogenous products accumulate in the cell, which in turn impacts on enzyme synthesis. Thus, deficiencies in any substance required for growth may reduce nitrogenase activity. As with exogenous carbohydrates, the main source of mineral nutrients is through decom- position of organic litter on the forest floor, al- though some may be available in leachate, stem- flow, or throughfall water (Long et al., 1956). The BENTLEY — NITROGEN FIXATION BY EPIPHYLLS pe Lr © T T T T T T T T T 25r =- 20r- 4 £ s i- o I5F 4 o — z o c IOF - 5 E L 1 1 1 L 1 1 L 1 L L L 2 4 6 8 10 I2 14 16 I8 20 22 24 IGURE 8. The influence of light and exogenous glucose on nitrogen fixation by epiphylls. Samples were incubated in the dark (solid symbols) or light (open d water (circles). The light treatments are significantly different after 4 hours (P < 0.05) and all treatments are significantly different after 8 hours (P < 0.01). latter, in fact, might be quite important in rain- forest systems: often a large proportion of de- composition occurs before the dead materials reach the forest floor (Duvigneaud & Denaeyer- de Smet, 1970; UNESCO, 1978). Thus, stem- flow and throughfall water can contain significant quantities of nutrient ions derived from “‘pre- fall" decomposition as well as that leached from living tissue. o document the effects of mineral nutrients under field conditions, we performed two types of experiments. The first was to simply measure the nutrients present in rainfall, throughfall, and water flowing off the surfaces of epiphyll-laden leaves, and the second was to determine if nu- trients in surface waters iier me mation rates. We chose to measure pI simply because these two ions have the most direct effects on fixation. As can be seen in Figure 9, both phosphate and ammonium are present in the environment. Interestingly, the water flow- ing off epiphyll-laden leaves actually has less ni- trogen than the rainfall collected in the open. This suggests that the uptake mechanisms on the leaf surface are extremely efficient. As expected, the concentrations of these two nutrients were T -300 300 L x | f -200+ 4 100 100 F + 50 O T P O T P NH, PO, FIGURE 9. Concentrations of ammonium (ug/ml, left ris and phosphate (ug/liter, right axis) in open- collected (O) rainwater, throughfall (T), and from epi- phyll-laden leaflets of Welfia georgii pinnae (P). Note the Ri iid high variance (error bars) in the through- fall value most variable in water collected as throughfall. Since these nutrients can affect fixation rates (Fig. 10), the concentrations of nutrients in the water washing over the leaf surface can be yet another component adding to the variances of fixation Observed in the field. Summary and conclusion. During the course of this study, we have documented that epi- phyllous microorganisms can fix significant quantities of atmospheric nitrogen. This nitrogen can be absorbed by the host leaf and thereby contributes to the nitrogen economy of the host. However, fixation rates are extremely variable and are strongly influenced by environmenta factors including substrate (leaf) longevity, light and concomitant desiccation, both organic and inorganic nutrients, and by the co-occurrence of bryophytes. Thus, while fixation by epiphylls can account for up to 2596 of the nitrogen present in a host leaf, the contribution of new nitrogen by epiphylls to an ecosystem is probably fairly small. Nevertheless, the very patchiness of the process ANNALS OF THE MISSOURI BOTANICAL GARDEN en 43: —121. [Vor. 74 400r D PO, ó CONTROL + NO, x NH 300|- N N E o — u z u 200} > =x = uj E x c ga 100+ x Zee 1 1 sb 1 6 E 18 24 INCUBATION TIME FIGURE 10. Effects of nutrient ions on nitrogen fix- ation by epiphylls. Leaf samples were dipped into so- lutions containing 176 ug/liter phosphate as NaH,PO, (D), 429 m n as NaNO, (+), 125 ug/liter ammonium X), or deionized water (Ó). Fixation E are d as nM ethylene to document that fixation continued at a constant rate throughout the 24-hour incubation. can free an individual plant from competition with its neighbors, which in turn could allow changes in community interactions. In other words, it is the variance in the system which has made this study so interesting. LITERATURE CITED BALANDREAU, J. C., C. MILLIER & Y. DOMMERGUES. 1974. Diurnal retur of activity in the field. Appl. Microbiol. 27: 662-6 BENTLEY, B. L. 1979, Longevity of individual leaves a tropical eae under-story. Ann. Bot. CARE. 1980. Effects of er washa in a tropical rainforest. Microbiol. Ecol j^ 109-113. & 1984. Direct transfer of newly- fixed nitrogen from free-living epiphyllous DUANE isms to their host plant. eK end BERRIE, G. K. & J. M. O. 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Nitrogen fixation in the phyllosphere. Pp. 85-100 in W. D. P. Stewart (editor), Nitrogen Fixation by Free-living Micro- "von sms. Cam- ridge University Se Cambridge. ies C. E. ndbook for Tropical Biology n Costa Rica: Data, gd s, Illustrations, and De- E ADE Organization for Tropical Studies, San José, Costa Rica. SEYBOLD, A. 1957. Traufelspitzen? Beitr. Biol. Pflan- zen 33: 237-264. SHREVE, F. 1914. The direct effects of rainfall on hygrophilous vegetation. J. Ecol. 2: 82-98. STAHL, E. 1891. Regenfall und Blattgestalt. Ann. Jard. Bot. Buitenzorg 11: 98-189. Ueber bunte Laubblatter, Ein Beitrag zur Pflanzenbiologie Ann. Jard. Bot. Buitenzorg 13: A im D. P. 1963. Liberation of extracellular nitrogen by two nitrogen-fixing blue-green algae. Nature 200: 1020-1021. 1973. Nitrogen fixation by photosynthetic microorganisms. Ann. Rev. Microbiol. 27: 283- 316. SuBBA Rao, N. S. 1977. w a es voci as a world-wide problem. Pp. 3-32 . Hardy & A. H. Gibson (editors), | ole on Dinitrogen Fixation: Agronomy and Ecology. J. Wiley and 1971. Leaching x n io plants. Pp. 67-80 in T. F. Pree C. H. Dick- inson (editors), Ecology of Leaf Surface Mii. organisms. Academic Press, New York & J. V. MORGAN. 1 e occurrence of leaching from above-ground un parts and the nature of the material Gage Proc. Int. Hort. Congr., Gembloux, Belgium 153. H. B. RUE 1957. Leach- ing of carbohydrates from plant foliage as related ce 126: : 120-121 (Natural Resources Research XIV). UNESCO, Paris. ADAPTIVE RADIATION OF SALAMANDERS IN MIDDLE AMERICAN CLOUD FORESTS! DAVID B. WAKE? ABSTRACT Tailed amphibians, or salamanders, occur in the tropics only in the New World, where they are concentrated primarily in Middle America and northwestern South America. All are members of the family Plethodontidae, e lungless salamanders. As recently as 1926 only 30 species of tropical salamanders were known, and all were placed in a single genus. Today 11 genera are recognized. All occur in Middle Pepe and over 140 species have been described. Many local tropical regions are very rich in numbers of species, and as many as 21 species may be present along a single altitudinal gradient. Community organization of species of salamanders in the tropics differs from that in tem- perate regions in that species of tropical salamanders te es, wl y given species restricted to a narrow elevational band. Within ational cie are segregated by major ha in) type, then by microhabitat, body size, and finally trophic and behavioral fi Cloud sa le eleva tions, from about 750 m to 2,500 m, are especially rich in e proportion of species occurring at lower elevations increases com- pared with Mexican and Guatemalan transects. Bromeliads and moss mats in the mid-elevational wet and rain forests are ideal microhabitats for these insectivorous, direct developing amphibians. Bromeliads offer abundant food resources, egg America has featured bo th convergent and parallel evolution. The mid-elevational cloud forests, le their ie iius assem- blages and highly dissected topography, have been of great significance in speciation, morphological and behavioral diversification, geographical ecology, and historical l... of tropical pletho- dontid salamanders. The cloud forests of Middle America are home to a unique group of vertebrates— lungless, climbing salamanders that belong to the family Plethodontidae. By “cloud forest" I refer rather loosely to those forest assemblages which form pending on many local and regional factors ue as temperature of the water of the nearest ocean, topography, rainfall patterns, and wind ir (Grubb & Whitmore, 1966; Myers, 1969). general, cloud forests form between elevations of about 800 and 2,700 m. Both upper and lower boundaries shift with climatic changes, with the lowest occurring along humid slopes in the low rs cu species. Although they do not breed n water, these organisms nevertheless require ici conditions for activity, and the cloud forest ' Research reported in this paper has i£ a collaborative effort involving many "n usaspa especially James whom have given helpful criticisms of the manuscript. Marvalee . Pa H. Wake carefully reviewed the manuscript n: David veia and Nancy Staub also provided useful comments. d à : à y I gratefully acknowledge the cooperation of govern Museum of Vertebrate Zoology. nal agencies in Mexico, Guatemala, and Costa Rica in obtaining permission to See pon in their nee and Douglas Robinson and Pedro Leon for their help and cooperation in Cost I dedicate this paper to ee senna of L. C. Stuart, who generously offered his unparalleled knowledge of the Guatemalan herpetofauna to me, and whose published work remains as an inspiration to future students of Guatemalan and other tropical environments ? Museum of Vertebrate Zoology ep Department of Zoology, University of California, Berkeley, California 985. 94720, U.S.A. Paper received 7 July ANN. MISSOURI BOT. GARD. 74: 242-264. 1987. 1987] environments are effectively buffered from des- iccating conditions as well as from extremes of temperature. In addition, these forests typically support abundant epiphytes that are used exten- sively by salamanders. There are more than 140 species of pletho- dontid salamanders in the New World tropics; about 80% occur in Middle America (Wake & Lynch, 1976; Frost, 1985). What makes them unusual is their great diversity in the New World tropics and their total absence from the Old World tropics. Middle America has been the setting for an extensive, unique adaptive radiation that has remained very localized. The success of these species can be measured against that of salamanders generally. World- wide, there are about 350 species of salamanders divided among nine families (Frost, 1985). Eight families are restricted to North Temperate re- gions. Over 200 of the species are members of the family Plethodontidae, the only group of sal- amanders to radiate in tropical regions. All trop- ical species of salamanders are members of the supergenus Bolitoglossa (Wake, 1966), which contains 11 genera and about 40% of the species of salamanders in the world. The supergenus is exclusively New World in distribution and does not occur north of Mexico. The plethodontids have a curious distribution, with two primary areas of evolutionary diver- sification: North America with concentration in the Appalachian region, and Middle America (Fig. 1). In eastern North America are found three major groups of plethodontids, two of which have life histories involving an aquatic larval stage. The Middle American region contains members of a fourth major group, the tribe Bolitoglossini, members of which have a uniphasic life history featuring direct development without a larval stage (Wake, 1966). The other two supergenera in the Bolitoglossini occur in California and Or- egon (and possibly in Alaska and Mexico), and on Sardinia, the Italian mainland, and a tiny portion of southeastern France. I have argued elsewhere that the absence of an aquatic larval stage facilitated occupancy of the relatively densely crowded, predator-rich tropics (Wake, 1966; Hanken et al., 1980). A unique feeding mechanism and an associated behavioral repertoire (Lombard & Wake, 1977, 1986; Roth Wake, 1985a, 1985b), which could evolve its particular characteristics only in a group lacking aquatic larvae, may have aided in the successful radiation of the tropical species. Those animals WAKE-— ADAPTIVE RADIATION OF SALAMANDERS 243 FIGURE 1. Latitudinal tof diversity in nthe salamander say Plethodontidae. The bers g scribed species for which descriptions are being pre- pared have been included. use a highly specialized, extremely fast tongue projection mechanism to capture moving prey at a considerable distance, and thus they are able to feed on a wide array of invertebrate animals. We still know relatively little about the tropical plethodontids. The authoritative work of Dunn (1926) listed but 30 species and placed them all in a single genus. Taylor (1944) recognized the generic diversity of the group (seven genera), but four additional genera were described as recently as 1983 (Elias & Wake, 1983; Wake & Elias, 1983), and about one-half ofthe 140 species have been described since 1950. Not surprisingly, most published work has dealt with taxonomy and systematics, although there has been some re- search on life history (Vial, 1968; Houck, 1977a, 1977b) and geographical ecology (Schmidt, 1936a; Martin, 1958; Wake & Lynch, 1976; Wake et al., 1987) The present paper attempts to evaluate the role mation that my research collaborators and I have gathered over the past 15 years. In particular, I examine the results of transect studies from Mex- ico to Costa Rica and concentrate on an area that appears to have been of critical importance for salamanders, the mid-elevation cloud forests. EcoLoGICAL GEOGRAPHY AND SYSTEMATICS Salamanders in the tropics have diversified in three major regions, each characterized by a rel- Q ° Nl ë ACTIVE SS saa IGURE 2. Generalized map illustrating ap of major evolutionary diversification within t e family Plethodontidae during Cenozoic times. od major fault t systems are indicated The family is thought to h the old and tectonically relatively adaptiv ve radiation in what is today Middle America, ith ail the three core regions indicated: the southeastern margin of the Mexican Plateau, Nuclear Central America, and Ta- lamancan Cent ral America. Two ' supergenera with e: in a reinvasion of temperate North America through association with land movements and tectonic activity in the extended San Andreas fault sys- tem. ore detailed analysis of geological history in relation to salamander distribution see Hendrickson (1986). atively ancient tectonic core, high topographic diversity, and high tectonic activity along some Central America (Wake & Lynch, 1976; Fig. 2). Each region is characterized by species richness and a high degree of endemism (Savage, 1982). For example, Chiropterotriton, Lineatriton, Thorius, and Parvimolge are endemic to region 1, and Pseudoeurycea occurs mainly in region 1 with only a few species in region 2. Dendrotriton, Bradytriton, and Nyctanolis are endemic to re- gion 2, and the great majority of the species of the beta assemblage of the large genus Bolito- glossa occur there. The few species of Bolito- glossa beta which occur in regions 1 and 3 are members of distinct subgroups (see Papenfuss et al., 1983). The genus Oedipina is centered in re- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FIGURE 3. C kei RAS arboreus, a bromeliad- dwelling salamander from meliad specialist: a long prehensile tail, relatively long legs, large hands and feet with widely spread digits, and frontally directed eyes. The fine divisions on the scale m. gion 3, and only one species extends as far north as Chiapas. The alpha assemblage of Bolitoglossa is centered in region 3 and areas to the south, but a distinct species group extends northwest- ward to the other two regions. The genus Noto- triton is something of a puzzle (see below). It has species in all three regions. Early zoogeographic biases combined with in- adequate collecting led to the perception of trop- ical salamanders as northern invaders that had "trickled" down into the tropics. Dunn (1926) placed all 30 species then recognized in a single genus, and several of his species groups later proved to us polyphyletic. For example, he united p, an nd all small, bromeliad dwellers into another, thereby ob- scuring the extensive parallelism and conver- gence that have occurred. Even after Taylor's (1944) generally progressive division ofthe genus Oedipus into several genera, the bias of recent ku... FiGURE 4. Dendrotriton Vibia a bromeliad- dwelling ian from the p Ovando, Chiapas, Mexico. This bro shares many gross structural s imilarities with species that occupy similar microhabitats (Figs. 3, 5). The scale bar is 25 mm 1987] WAKE— FIGURE 3. dieat . er a bromeliad- dwellin N Santa Cruz, Za- capa, Guatemala, in the Sierra den las Minas. Compare this species with unrelated bromeliad ase in Fig- ures 3 and 4. The scale bar is 25 m penetration from the north persisted. The genus Chiropterotriton as recognized prior to 1983 pro- vides an example. Its species mainly are bro- meliad specialists living in cloud forests, and they are superficially similar in external morphology (Figs. 3-5). Both Rabb (1960) and Wake (1966) recognized that members of the genus differed substantially in osteology, but they chose to in- terpret this as increasing divergence and spe- cialization toward the south. This especially in- structive case is relevant to the main theme o this paper and is developed further below Species once assigned to Chiropterotriton are typically small, slender, ibe tailed, acrobatic forms, that are s of cloud for- ests in Mexico, Nuclear Central America, and T T ER T 105 DU 1400 95* 90 z N \ PM ` P j t 5M : ' ER 4 i t vM 25° ` > 3 aE Pa Chiropterotriton == FIGURE 6. Probable distribution of the genus Chi- ropterotriton. Most species occur in cloud forests, and The gen- a number eralized ranges were derived by grouping known lo- calities into suger contiguous units, based on for- est distribut Se tion gathered by David M. Darda and i autho ADAPTIVE RADIATION OF SALAMANDERS 245 Denarotriton ed to cloud forests. All occupy small geographic ranges, and ; ccurs in more than one of the iso- Re indicated here (Elias, 1984). Costa Rica (Figs. 3-5). Based on their study of comparative osteology, Lynch & Wake (1975, 1978) recognized that the species below the Isth- mus of Tehuantepec formed a cladistically dis- tinct group, which they termed CAiropterotriton beta. The latter group was found to include at Nototriton 1 IGURE 8. Distribution of the genus Nototriton. This genus may not be a monophyletic group. The species morphological and ecological features. All of the species inhabit cloud forests, but the recently discovered and as yet undescribed species. 246 Cerro San Felipe Pine- Oak _Fir Forest GEE 7 »c7/sova/s ENS Punquidentis SSSSS P smithi T. pulmonaris || Y E I maoga P. cochranae BSSSS P belli ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 r t - M ,XPine-Oak Fir a > Š E a [7] Cerro Pelón q. p A A Z S P smithi SSS P juarezi Gl D m d Forest o a o K a 4 P sp.nov 3 Pine-Oak Forest Cloud 2000 5 TW E= Norotriton sp.nov ENS P wer/er/ eed B rufescens Forest SEEN 7 -pE L^ mm C sp nov 4 ° A Š | ` ` ef Tropical Deciduous Forest | a * Oaxaca | a oo] — x l m ý Evergreen Seasonal Forest ` ` Low Deciduous Forest s < SW ^ |00 Kms NE — -northeastward from Ciuda Distribution of plethodontid salamanders along the Northern Oaxacan transect, about 100 km in ort d de O extensive cloud forest which offers habitat to numerous species of salamanders. Updated version of diagram presented by Wake et al. (1987). least two distinct subgroups, an interpretation were given generic status as D drotriton and Nototriton by Wake & Elias (1983), who noted that Nototriton might prove to be paraphyletic. Some species of each of the three genera resulting from the subdivision of Chirop- terotriton include members that are very similar in external morphology. Because the species are very small (frequently «40 mm in head plus body length), the critical differences in mor- phology can be difficult to detect. The species of the three genera typically have small ranges, and the widely disjunct ranges offer little od for symp hese salamanders found mainly in iced cloud forests, and nl of sively allopatric (Figs. 6—8). Chiropterotriton oc- curs as 11 or more discrete geographic isolates in eastern Mexico, north of the Isthmus of Te- huantepec. Dendrotritonis found only in Nuclear Central America, mainly on Pacific slopes, but also in some internal regions of Caribbean drain- age. Nototriton is the southernmost of this group, and it occurs in regions of Caribbean drainage. An undescribed Oaxacan species of Nototriton detailed studies of vertical zonation along a tran- sect (Fig. 9). Our knowledge of the groups prob- ably remains far from complete, even at the alpha taxonomic level, but they clearly offer fascinating opportunities for the study of convergent evo- lution. Lynch & Wake (1978) showed that bromeliad- inhabiting species currently placed in the genera endrotriton an drotriton in external morphology (evaluated by methods of multivariate morphometrics) than to semi-fossorial congeners that live in moss mats on soil banks. These distantly related species have converged so that they share body forms that are especially well suited for life in bromeliads. In- deed, if Nototriton is, as I believe, paraphyletic it is possible that we actually have underesti- mated the true amount of evolutionary conver- gence. 1987] The three genera discussed above are com- monly encountered inhabitants of cloud forests and epiphytes. Other tropical salamander genera also contain cloud forest inhabitants, and many use epiphytes as their main microhabitats. Some of these, such as the Nuclear Central American ered and their habits are very poorly 1984). The monotypic Mexican genera Parvi- molge and Lineatriton (the latter an elongate fos- sorial form that utilizes moss mats to some de- gree; Fig. 11) are relatively rare within their restricted ranges, which lie at the lower margins of cloud forests. Some species of the Mexican genus Thorius occasionally occur in bromeliads; an undescribed species from the northern slopes of the Sierra de Juarez in Oaxaca seems to occur primarily in bromeliads (undescribed species E, fig. 9 in Hanken, 1983). The remaining genera (Pseudoeurycea, Bolitoglossa, and Oedipina) have numerous species that are inhabitants of cloud fore P sn is widespread in Mexican and southwestern Guatemalan cloud forests, but most species are terrestrial and are not often found in epiphytes. The only described species that are bromeliad specialists are P. firscheini (Werler & Smith, 1952; Shannon & Werler, 1955b) and P nigromaculata of Veracruzian cloud forests (un- publ. data, contra Taylor, 1941). An undescribed species from our Monier) Oaxacan transect (Fig. 9) uses arboreal bitats, and an additional undescribed species from our San Marcos tran- sect (Wake & Lynch, 1976: 30) uses bromeliads consistently. Bolitoglossa, with 68 currently recognized species, has by far the greatest geographic range of the tropical salamander genera (from Vera- cruz, Mexico, to Brazil, Bolivia, and Peru). Many of the species in Nuclear Central America are cloud forest specialists, and they frequently occur in bromeliads. Over one-half of the B. engel- hardti encountered during an intensive investi- gation of an elevational transect on the lower slopes of Volcan Tajumulco, western Guate- mala, were found in bromeliads, and B. franklini is also a frequent inhabitant of bromeliads (Wake & Lynch, 1976). Most records for Bolitoglossa in bromeliads refer to members of the beta as- semblage (e.g., Stuart, 1943). Some of these occur north of the Isthmus of Tehuantepec (the north- ern limits of Nuclear Central America), including B. hermosa (Papenfuss et al., 1983), from the Pacific slopes of Guerrero, Mexico, and the wide- WAKE—ADAPTIVE RADIATION OF SALAMANDERS 247 spread Gulf-Caribbean slope species B. rufescens (Taylor & Smith, 1945), which ranges from San Luis Potosi, Mexico, to Honduras. The only two species of the beta assemblage that reach Costa Rica (B. alvaradoi, B. arborescandens) have been taken in bromeliads (Taylor, 1954; unpubl. data Occurrence of members of the large alpha as- semblage of Bolitoglossa in bromeliads is less well documented. The distribution of this group found mainly in the lowlands, and there are no records of the species being found in bromeliads in cloud forests. Two members of the group, B. platydactyla and B. mexicana, have been re- corded from bromeliads, mainly at elevations of <500 m (Taylor & Smith, 1945). There are scat- tered reports of Bolitoglossa alpha in cloud forest bromeliads in Talamancan Central America and regions to the south (e.g., B. borburata near Ran- cho Grande, Venezuela, Trapido, 1942; B. lig- nicolor, Dunn, 1937; B. subpalmata, Robinson, 1977; B. taylori, Wake et al., 1970). But in Costa Rica, where the assemblage is well represented, there are surprisingly few records of its occur- rence in bromeliads (Robinson, 1977), although we now know that some species are common in such microhabitats (see below). Several species of the alpha assemblage of Bo- litoglossa are associated with arboreal microhab- itats in cloud forests. The only known adult of Bolitoglossa diminuta was collected with an egg mass in a mat of liverworts (Robinson, 1976; recent examination of the tiny holotype, which lacks a sublingual fold, suggests that this species should remain in Bolitoglossa, contra Wake & Elias, 1983). Other species associated with moss mats covering tree trunks and branches include B. marmorea of Panama (Wake et al., 1973) and an undescribed species sympatric with B. dimi- nuta. The final genus, Oedipina, is widespread in cloud forest habitats in Costa Rica, the center of its diversity (Brame, 1968). These salamanders are elongate, mainly fossorial species that include some relatively specialized arboreal climbers in lowland forests (e.g., O. parvipes). The species that occur at intermediate elevations in cloud forests typically are found in moss mats covering owned vegetation and soil banks. Information in the above paragraphs makes 248 clear that there has been an extensive adaptive radiation of salamanders in the New World trop- ics, but the age of this radiation remains un- known. Since the initial effort of Dunn (1926), subsequent studies have for the most part sug- gested progressively earlier dates for the entry of salamanders into the region (Martin & Harrell, 1957, is a striking exception), and until recently an Early or Middle Tertiary origin of the group was accepted (Wake & Lynch, 1976). But bio- chemical and immunological studies in the last decade have shown that even within genera there has been very great genic differentiation, which implies relatively great age for the separation of the lineages studied (Hanken, 1983; Hanken & Wake, 1982; Larson, 1983, 1984; Lynch et al., 1983; Maxson & Wake, 1981; Papenfuss et al., 1983; Wake & Lynch, 1982). Progress has been made in defining monophyletic groups, but I be- lieve that we have not yet achieved a robust cla- distic hypothesis for the group (Wake & Elias, 1983), mainly because of the extensive parallel- ism and convergence that have obscured pat- terns. Nevertheless, Hendrickson (1986) has at- tempted to interpret the history of the group by combining what is known about likely cladistic patterns with knowledge of the geological history of the region in a vicariance biogeography study. He suggested that salamanders which gave rise to the tropical radiation first separated from those in the Appalachia area by rifting of an ancient Maya terrane from Appalachia or by a post-Mid- dle Jurassic to Mid-Cretaceous marine transgres- sion. In general he argues for much older times a slater than previous authors, based both nts from earth history and from his belief rois he has not studied these sala- manders directly) that the extensive radiation of the tropical salamanders must have taken a long time. I cannot discuss this provocative study in detail here, but it is important to understand that available evidence suggests that salamanders and habitats have coevolved in areas that became present-day Middle America for a very long time. SALAMANDERS AND EPIPHYTES The epiphytic component of cloud forests of- fers two major classes of microhabitats for sal- angiosperms). These microhabitats, particularly bromeliads, are used on occasion by other ver- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 tebrates: frogs (especially Hyla and Eleuthero- dactylus), lizards (especially Abronia), and snakes (e.g., Bothrops schlegeli). However, with the ex- ception of a few species of frogs whose tadpoles are clearly adapted for life in the water of tank bromeliads, only salamanders rely on epiphytic plants as their main microhabitats, and sala- manders are far more common in bromeliads than are any other vertebrates. The density of salamanders in bromeliads is difficult to document. As many as 34 Dendro- triton xolocalcae have been found in a single bro- meliad in Chiapas, Mexico (Taylor & Smith, 1945), but until recently we have had few quan- titative data to indicate the frequency of occur- rence of salamanders in bromeliads. Although one of the first reports of salamanders living in bromeliads was from Costa Rica (Picado, 1913), the general impression has been that salaman- ders are not common in bromeliads (Robinson, 1977). Bromeliads have been thought to be more important for salamanders in Mexico, Guate- mala, and Honduras. In an early account, Gadow (1908) reported that a Mexican species of Pseu- doeurycea leads a “‘partly arboreal life, their fa- vorite hunting and hiding-places being in the clusters of epiphytic plants, such as tillandsias, orchids and the climbing phyllodendrons.”’ Schmidt (1936a, 1942) described bromeliad- dwelling salamanders as relatively abundant in Guatemala and Honduras, apparently more so than in Costa Rica and Panama (Dunn, 1937). In contrast to the above generalization, sala- manders in Costa Rica use moss mats more com- monly than do salamanders farther to the north and west. The genera Nototriton and Oedipina use moss mats extensively in Costa Rica, but apparently rarely do so in northern Middle America. To the north and west Oedipina is mainly fossorial, and Nototriton is associated mainly with bromeliads (an exception may be a poorly known, undescribed species from Chiapas that has been taken in a moss-covered bank). There is a general morpaoiogy that Character izes most bromeliad igs 3-5). They typically are small animals hei « 50 mm body length) with long, prehensile tails, long limbs with widely spread digits, and fron- tally directed eyes. They are acrobatic climbers and are very adept in a three dimensional en- vironment. Some larger salamanders use bro- meliads on occasion, but the true oe usu- ally approximate the above descriptio Occupants of moss mats are less PH 1987] in morphology. In general they are slender and have relatively short legs. The Appendix con- tains a list of species which have been reported to occur in bromeliads and moss mats. It also contains a few species known by me to have such habits, but which as yet are not reported in the literature. This list does not differentiate between species that specialize on these microhabitats and those that are but casual occupants. COMMUNITY ORGANIZATION For nearly 15 years my colleagues (James F. Lynch and Theodore J. Papenfuss) and I have been engaged in a broadly based survey of geo- graphical ecology and community organization f the salamanders of Mesoamerica (Wake & Lynch, 1976; Wake et al., 1987). We established a series of line transects in Mexico, Guatemala, and Costa Rica and conducted intensive sam- pling over a multi-year period along each of them. r the most intensively studied of these the continental divide, down t of Volcán Tajumulco, to the Pacific coastal plain. Here we have documented the presence of a rich salamander fauna comprised of 15 species, rang- ing from near sea level to nearly 4,000 m. This transect has provided an opportunity to study the ecological organization of this group of species, relative to each other and to various physical and biotic factors. An earlier study (Wake & Lynch, 1976) presented a general overview of our results, and a recent report (Wake et al., 1987) updates the main patterns of distribution. A summary of our primary results and data rele- vant to the use of epiphytes by salamanders is presented here. Schmidt (19362) collected seven species of sal- amanders on the slopes of Volcán Tajumulco and inferred the presence of two additional species. He outlined their main patterns of ver- tical distribution and presented comparisons with the nine species then known from Veracruz, Mexico. His basic conclusion was that zonation was sharper in Guatemala than in Mexico. e (Wake & Lynch, 1976; Wake et al., ysi nized four elevational assemblages of salaman- WAKE—ADAPTIVE RADIATION OF SALAMANDERS 249 ders, including a lower cloud forest (1,600-2,400 m) group of four and an upper cloud forest (2,400— 2,800 m) group of seven species. These ten species (one is present in both elevational belts) are re- stricted not only in elevational zonation, but also in geographic distribution; none of the species occurs beyond the limits of the southwestern Guatemalan volcanoes and the adjacent Sierra Madre of Chiapas, Mexico. The region between 1,600 m and 2,800 m is occupied by Evergreen Cloud Forest (above about 1,900 m) and Mon- tane Rain Forest, using the terminology of Breed- love (1981), who also has characterized these formations floristically Within elevational zones we examined differ- ential use of major habitat types. For example, some species are more common in edge situa- tions, and some favor small clearings and open spaces, while others are found throughout the log-dwelling (within and under the bark of logs and stumps), arboreal (within a leaf-axil micro- habitat, including bromeliads), and fossorial (within subterranean passageways). We found no moss mat specialists and therefore did not rec- ognize this microhabitat category. Within mi- crohabitats we paid special attention to differ- ences in body size and in trophic wampis (morphology of the Jaws and teeth). The primary modes of ecological segregation of de t species are indicated in Table 1. Here a one-sided matrix of potential co-occurrence of the species records our assessment of primary segregation ordered according to decreasing spatial proximity of the segregated species: elevation, habitat, microhab- itat, size, and trophic specialization. Nearly three- fourths of the potential sympatric associations of the 15 species are precluded by differences in elevational distribution, and habitat or micro- habitat differences separate all but nine of the remaining paired associations. Of the nine pairs of species which show elevational, habitat, and microhabitat snp cight differ importantly by size Smari in size and morphology, ekteni that ne species has about halfas many substantially tp er maxillary teeth and enlarged jaw muscles. Six species on the transect commonly occur in arboreal microhabitats (in order of frequency in such microhabitats, from most frequent to least): Bolitoglossa occidentalis, Dendrotriton bromelia- Primary modes of ecological segregation among species that occur along the San Marcos transect. S appearance from high to low elevations. In order of decreasing spatial proximity: E specializations. From Wake & Lynch (1976). TABLE 1. pecies arranged in approximate order of habitat; M 250 — size; T — trophic microhabitat; S elevation; H = P. rex 1. 2. P. sp. (E)M 3. P. brunnata 5. B. rostrata 6. B. resplendens 7. D. bromeliacia ANNALS OF THE MISSOURI BOTANICAL GARDEN (H) M 9. B. franklini 13. B. salvinii 14. O. ignea 15. B. flaviventris [Vor. 74 ad surface of the soil; I = under bark of stumps and stand- ing trees, and inside fallen logs. Open symbol = Pseu- doeurycea; half-closed symbol = Dendrotriton; closed Sy — Bolitogl mbers 2 through 10, and 12, are cloud forest in- habitants E from Wake & Lynch (1976). cia, B. engelhardti, B. franklini, Pseudoeurycea sp., and B. resplendens (Fig. 10). All but the first, which occurs at relatively low elevations, are present in the cloud forest and make extensive use of bromeliads. Bolitoglossa occidentalis is a small, mainly lowland species that occurs in bro- meliads, but it is most commonly found now within the so-called "coffee zone," where it oc- curs in agricultural plantings of bananas. James F. Lynch and I are preparing a detailed ecological account of our work in this transect, and with his permission I present here some of our data concerning use of bromeliads in cloud forests by the above listed species. When we first visited this area in 1969 primary forest extended to roadside and bromeliads were abundant. When we last visited the cloud forest region in 1980 the forest had been removed and pasture occu- pied nearly the entire area between 1,500 m and 00 m. Below 1,500 m traditional coffee plan- tations, which feature large shade trees and ex- tensive plantings of bananas, had given way to a near monoculture of coffee grown in hedgerows without any suitable cover for arboreal salaman- ders. 1987] TABLE 2. Relative abundance of salamanders in bromeliads, San Marcos transect. WAKE—ADAPTIVE RADIATION OF SALAMANDERS Elevation Wet Season! Dry Season? Combined Below 1,750 m — 3/25 = 0.123 3/25 = 0.12 1,750-2,000 m 6/39 = 0.15 41/75 = 0.55 47/114 = 0.41 2,000-2,250 m 48/239 — 0.20 29/64 — 0.45 71/303 — 0.25 2,250-2,500 m 89/121 = 0.74 140/191 — 0.73 229/312 = 0.73 2,500-2,750 m 81/25 = 0.72 27/30 — 0.90 45/55 — 0.82 Above 2,750 m 14/73 = 0.19 13/21 = 0.62 27/94 — 0.29 ! May-September. 2 November- Fe ry. 3 Number of salamanders/number of bromeliads. Salamanders are common inhabitants of bro- meliads along the San Marcos transect (Table 2). We found salamanders in approximately every second bromeliad we opened. These bromeliads, primarily members of the genera Tillandsia and Vriesia, were located relatively low in the trees. Salamanders in southwestern Guatemala are most abundant in bromeliads at elevations be- tween 2,250 and 2,750 m. From these elevations down to approximately 1,700 m, bromeliads re- main relatively common, and there are brome- liads present at elevations up to approximately 3,000 m. All of the bromeliad specialists occur in the cloud forest (roughly 1,500-2,750 m), even though bromeliads are found both above and below that formation. Above the cloud forest those species that use bromeliads at lower ele- vations (e.g., Bolitoglossa rostrata, Pseudoeury- cea rex) shift almost entirely to terrestrial mi- crohabitats. Salamanders are consistently more abundant in bromeliads during the dry season than during the wet season, except in the heart of the cloud forest (2,250-2,500 m), where there is less seasonality than elsewhere. ABLE 3. Distribution of salamanders in brome- liads, San Marcos transect, 16 January 1972, 2,400 m elevation.! Number of Salamanders r Bromeliad Frequency 0 15 l 10 2 8 3 4 1 5 l 6 1 8 1 Martin Feder accompanied us on one trip to our transect and studied the thermal ecology of some of the cloud forest salamanders pies 1982). He found that bromeliads, even in the cloud forest, afford cooler and more tans tem- peratures than microhabitats in immediately surrounding areas. Bromeliad-dwelling salaman- ders appear not to thermoregulate behaviorally or k ario because thermal diversity in their microhabitats is so low as to offer little opportunity for such behavior. As is usual for salamanders, there is a high correlation between body temperatures of salamanders and prevail- ing microenvironmental conditions (Feder & Lynch, 1982), so the more stable the microen- vironment, the less variable will be the temper- ature of the salamander. The tropical salaman- ders contrast sharply with more northern plethodontids in having very limited ability to undergo thermal acclimation (Feder, 1978, 1982). Ibis may be either us Fuse Or ins eflect ar the high fidelity t displayed by many of these species (Feder, 1983). The data and analyses in Tables 3-5 indicate that the distribution of salamanders per bro- TABLE 4. Distribution of salamanders in brome- liads, San Marcos transect, 18 January 1972, 2,300- 2,350 m Number of Salamanders er Bromeliad Frequency 0 12 l 8 2 10 3 4 7 l 5 2 ! These data are for 55 Dendrotriton bromeliacia and three Bolitoglossa franklini taken from a sample o bromeliads (X — 1.45 salamanders per bromeliad). ! These data are for 59 Dendrotriton bromeliacia and four Bolitoglossa franklini collected in a sample of 40 bromeliads (X = 1.58 salamanders per bromeliad). 252 TABLE 5. Test for randomness, combined data from Tables 3 and 4.' Number of Salamanders Expected per Observed Frequency Bromeliad Frequency (Poisson) 0 27 17.5 l 18 26.6 2 18 20.2 3 10 10.3 4 2 3.9 r fH. 2 6 7 0.3 5.5 7 1 > 8 l J Chi-square (goodness of fit) = 8.50 with 4 df 0.05 > P > 0.1 ANNALS OF THE MISSOURI BOTANICAL GARDEN ' Data for 121 salamanders collected from 80 bro- meliads (X = 1.52 salamanders per bromeliad). . meliadisnotsi different from random. — p there is at least a suggestion that there t be an excess of bromeliads that lack sal- de as well as a deficiency of bromeliads containing single salamanders. Thus, there might be a tendency toward clumping under conditions of high salamander abundance romeliads might seem to be a near perfect microhabitat for salamanders, in terms of food availability, thermal stability, and constancy of humidity. But concentration of salamanders in bromeliads might attract predators. Spiders and salamanders are the top resident carnivores with- in bromeliads, and they do not prey on each other very extensively. Some arboreal snakes forage widely and may be important, although infre- quent, predatory visitors to bromeliads; birds might also be important predators. All tropical salamanders have a specialized autotomy zone at the base of the tail, and Shaffer (1978) ex- amined tail loss frequency as a rough index of relative predation pressure on 10 species of sal- amanders from the San Marcos transect (parts of tails may be lost in intraspecific aggressive lation between rates of tail loss and elevation, and we know that snake densities also decrease with elevation. Two of the three cloud forest species commonly found in bromeliads (Den- drotriton bromeliacia, Bolitoglossa franklini) had the second and third highest tail loss percentages of the species studied (the highest was the ar- [Vor. 74 Outlines of the left hind foot of two FIGURE 11. (this ne is 67.5 mm, snout-vent length) with extensive webbing and is capable of generating suction in arboreal ano Bolitoglossa occidentalis is a di- minutive species (this specimen is 38.7 mm, snout- vent length) with feet that superficially appear to be webbed, but in reality are just incompletely developed (paedomorphic) as suggested by the strong gradient in phalangeal structure within each digit. boreal lowland species B. occidentalis). Although exposure to predation may be a relative cost for living in bromeliads, the cost is apparently out- weighed by Sa advantages, such as those men- tioned abov Although Diss dwellers in cloud forest habitats differ greatly in morphology, ecology, and behavior from even the most arboreal North American plethodontids (e.g., Aneides), they are not the most extremely specialized species on the San Marcos transect. At elevations below about 1,400 m the cloud forests, and the lower cloud forest salamander fauna, are absent. A new sal- amander community appears at about 1,000 m composed of three extreme morphological and ecological specialists. This community includes a relatively large and a relatively small arboreal species of the genus Bolitoglossa and an elongate fossorial species of the genus Oedipina. olitoglossa salvinii, a large species, is an ac- tive, climbing animal with a long prehensile tail and large hands and feet with extensive inter- digital webbing (Fig. 11). These animals, which frequent surfaces of Heliconia and other large- leafed plants on moist evenings, are capable of producing suction with their large hands and feet (Alberch, 1981). Bolitoglossa occidentalis, the small species, has small hands and feet that appear to be fully ebbed. In reality the hands and feet manifest incomplete development, a phenomenon known f 1987] as paedomorphosis that affects a number of fea- tures of the organism (Wake, 1966; Wake & Brame, 1969; Alberch et al., 1979; Alberch & Alberch, 1981). Such similarly affected features with digits that show a strong gradient of devel- opment (Fig. 11). Although these animals do not generate suction with their hands and feet (AI- berch, 1981), they are very agile, partly as a result of their small size. They have an extensive ven- tral surface area (body, limb, and tail) relative to their mass, so they "stick" to moist plant surfaces by surface tension. Apparently these salaman- ders are virtually restricted to leaf axil retreats. They especially favor Heliconia and both culti- vated and feral banana plants (Musa spp.). Smith (1945) reported finding Bolitoglossa rufescens ery similar in morphology and ecology to its close relative B. occidentalis) to be abundant in red bananas (he found about 250 per hour) at a Veracruzian site. These animals are adept at climbing small tendrils, stems, and strands of moss. Members of the small species of Bolitoglossa are encountered only rarely in terrestrial sites, and the large species, while occasionally found on the ground (e.g., crossing roads on rainy nights), also are basically arboreal. (An exception may be the very large species B. dofleini, which can be very common in terrestrial situations.) But another group of lowland species, the very elongate, short-legged genus Oedipina, is found only at and beneath the surface of the ground. Species of Oedipina are elongated as a result of the addition of trunk and especially tail verte- brae, and they have bizarre long tails (Fig. 12). In addition to their extremely short legs and tiny hands and feet, they have heads and bodies of very small diameters; all of these features facil- itate use of root channels and underground bur- rows. The San Marcos transect is special because more species of salamanders occur there than in any other area of the Pacific Versant in Middle America. Volcan Tajumulco (4,200 m) is at- tached to the Guatemalan Plateau at about the 3,000 m level, so on both sides of the volcano there are substantial areas of moderate elevation which trap moisture and thereby create favorable salamander habitat. To the south and east the Plateau gradually lowers and rainfall declines. In the vicinity of Guatemala City, Volcan Agua (over WAKE—ADAPTIVE RADIATION OF SALAMANDERS FIGURE 12. eg d bove (top), from near o de las Flores, Ver z, Mexico, and Oedipina Le (bottom), from Fis a Julia, near San Rafael de la Cuesta, San Marcos, Guatemala. The scale bul r is 25 mm. These extremely elongated species are semi-fossorial to fossorial in habit, and have evolved convergently. 3,700 m) towers over the Plateau, to which it is attached along only its northern flank at a low elevation (about 1,000 m). The contrast in sal- amander faunas between Volcán Tajumulco and Volcán Agua is great. Only three species have been collected on the slopes of Agua. There is a single high elevation member of the genus Pseu- doeurycea (P. goebeli), one low elevation species of the genus Bolitoglossa (B. salvinii, a large, fully webbed species), and a middle elevation gener- alized species of the genus Bolitoglossa (B. mo- rio). All three of these species also are present on the San Marcos transect. There is a well-devel- oped cloud forest with abundant bromeliads on Volcán Agua, but the forest is localized and it is isolated from similar habitats to the north and west by low elevations covered by drier vege- tation types. In 1969 we opened about 600 bro- meliads on Volcán Agua, but found only two salamanders! This contrasts sharply with the data areas of the Guatemalan Pl e Marcos transect it is found only at the top of the cloud forest and in drier broadleaf forest above, where the edge of the Plateau contacts Volcan Tajumulco. Here the species occurs occasionally in bromeliads (Fig. 10). On Volcan Agua, al- ough B. morio remains uncommon in bro- ae and apparently has not modified its mi- crohabitat utilization patterns in any dramatic way, its elevational range is 1,300-2,500 m. At this site, where B. morio is the only salamander e a Q oS "mA S T f J 4 = NM J } f nm ghe COSTA RICAN TRANSECTS o Q 25 p 100 i50 eM FiGURE 13. Map of Costa Rica indicating the lo- cation of the Irazü and Tapanti transects illustrated in Figures 14 and 15. present, it occupies an elevational range that accommodates ten species on the San Marcos transect. The patterns of species distribution in the tropics (this example, but see below also) are probably determined by combinations of physiological constraints (in relation to physical factors in the environment) as well as such in- terspecific interactions as predation and com- petition COMPARATIVE ASPECTS OF COMMUNITY ORGANIZATION Until recently most fieldwork by my group has been in Mexico and Guatemala (for general sum- maries see Wake & Lynch, 1976; Wake et al., 1987). In general, the results of our detailed stud- ies of the San Marcos transect have been mir- rored in other areas (cf. Fig. 9). Typically only one or two species occur at elevations above 3,500 m; as one moves lower the number of species present in a given habitat increases dramatically at about 3,000 m and continues to be relatively high until the lower limit of the cloud forest is reached. At elevations below 1,000 m the num- ber of species present typically declines, and at sea level the largest number of species definitely known to be present is four on the Osa Peninsula of Costa Rica, where there are two species o Bolitoglossa and two species of Oedipina. Pos- sibly five species occur together at sea level in the region of Bocas del Toro, Panama (Wake et al., 1973 and unpubl. data), and in northeastern M ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Costa Rica seven species are known from ele- vations below 1,000 m, but not in local sympatry (see below). The proportion of the total sala- mander fauna present in the lowlands increases at lower latitudes within Middle America (see Table 5 in Wake & Lynch, 1976). There is evi- dence of increased *'tropicality" (that is, of closer packing of species in communities and increased numbers of species present locally) as one moves into the deep tropics. Finally, however, in north- western South America this trend stops, perhaps because plethodontids are not thought to have dispersed into South America until late Pliocene times (Wake & Lynch, 1976; Hanken & Wake, 1982). Except in the Chocó and the flanks of the northern Cordilleran regions of Colombia, the number of salamander sp tinlowland sites in South America is not known to exceed two, even in such biotically rich 1m as those of the Río Palenque area of Ecuador. Throughout Middle America, ai cadens are found in cloud forests. In Mexico and in Nuclear Central America, cloud forest salamanders make extensive use of bromeliads as microhabitats, even at the extreme northeastern limits of the range of the supergenus Bolitoglossa in the Gó- mez Farias region of Tamaulipas (where cloud forests also reach their northern limit; Martin, 1958). But the manner in which cloud forest mi- crohabitats are utilized changes in Costa Rica. the tropics, thanks primarily to the work of Tay- ie d 1954). However, very little has been utilization and patterns of co-occurrence of Costa Rican species. Only four species are reported to occur in bro- meliads (Robinson, 1977), although many more species are known to inhabit cloud forests. Recently I have been investigating the system- atics and distribution of Costa Rican salaman- ders in some detail. I have focused attention on two general transects (Figs. 13-15). Results for Costa Rica are preliminary, because major sec- tions of these generalized transects have yet to be searched thoroughly. Nevertheless, certain marked contrasts with more northerly transects are apparent. dominant theme in the history of studies of tropical salamanders has been that the species which are the most conservative ecologically and the most primitive phylogenetically occur in Mexico, and that there is both increased spe- cialization and a decline in the number of species 1987] WAKE—ADAPTIVE RADIATION OF SALAMANDERS Volcan Irazu FR Montane 30004 $ Wet Forest_ Ù m S ° £ ° s > £ Š S Hs 4 = 8 & S 3 ` D e ` ` $ Qr < X 2000 Q Š Š Š Š S Q S S Š SOR 3 S S Q à o Ş ° > S : 9 Š q Š Š | Forest x D t. S S A, = Å 3 Lr 10004 S n € da N X ts f Premontane Rain S 9 : Forest Ss 500- = = EE MAN & o9 o Tropical Wet and ° O COE: = i! | ce ERAN root Premontane Wet Forest O = V ee RA TNE T t o 4 20 Ki O FIGURE 14. Vertical distribution of pula salamanders along the Irazü transect, extending about 55 km from Finca La Selva to Volcán Irazü, Costa Ric to the south (Dunn, 1926; Taylor, 1944, 1952; Brame & Wake, 1963; Wake, 1966; Wake & Lynch, 1976). Even though Wake & Lynch (1976) documented the presence of a large number of species below 10? latitude, we continued to es- pouse the traditional view of relics in the north (e.g., Chiropterotriton priscus of Coahuila and Nuevo León, Mexico) and increasingly derived forms to the south. The recent discovery of the most primitive known genus of tropical pletho- dontid in Guatemala (Elias & Wake, 1983) and a fresh analysis of relationships of new and ex- isting groups (Wake & Elias, 1983) have forced me to re-evaluate my earlier views. It now seems likely that salamanders have been in the tropics of present-day Middle America for a very long time, possibly throughout the Tertiary (see above section on Ecological Geography and System- atics). Middle elevations doubtless have been important areas of both survival and radiation in the group, and we have discovered that pat- terns of microhabitat utilization differ dramati- cally between Mexico and Nuclear Central America on one hand, and Talamancan Central America on the other. Perhaps the most striking difference between lier (Table 2), salamanders are sometime abundant in cloud forests of northern Middle America, where many species utilize bromeliads (Wake & Lynch, 1976). In Veracruz, Mexico, and in Nuclear Central America, elongate, fossorial species of Lineatriton and Oedipina occur only at elevations below the lower limit of cloud for- est. In contrast, elongate members of the genus Oedipina are well represented in Costa Rican and Panamanian cloud forests, extending upwards to elevations in excess of 2,000 m (Figs. 14, 15). Here they utilize moss mats covering soil banks, downed logs, and stumps. Furthermore, living 256 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 E & Š Cerro de Š la Muerte y) Y 2 à E È Ad a X. f Š $ ae 3000 s XS S a s o <, S I Q Š ° q Montane X 3 £5 Rain 4 © ë Š Á Forest E E Qs Um - $Š Saan $ Bs S D š ES Š & Lower 20004 S Montane — ES š y ain i S Š y ^ & S Sk x 3 NEUEN 82 _ 88 3s8h 8 Í Forest š Sei] Š Se Fes ech s Lm š ] $£| Š š Š ° š Š Š ° à p = Š Š š Š SQ Š m Qi £ 2 8 $ S ü SFA 73 Š Qj E] € Premontane 10004 : S a 4 p i 1 BS S ~ 5 Y & Premontane », š : S š S ü Rain Forest Ma | OR SP £ act 5004 $ Š i 8S 9$ Ar Tropical | Moist Forest mo Š y^ Wet Forest S 2 200 e Ü s | a [| — Sines Tropical Moist and Premontane Wet Forest T T T T T 40 60 O 20 FIGURE 15. Vertical distribution of PN aig p ege med a vepres Tapantí transect, which a Ric 6 extends between Siquirres and Cerro de la Muert a. This re EE 0 km long, contains more species of plethodontid salamanders than any bou area in al ann in moss mats in these cloud forests are species is even an undescribed Costa Rican species of Bolitoglossa that has been taken only in moss surrounding twigs on trees and shrubs in cloud forests. This substantial fauna represents an eco- logical component that is rare (e.g., Dendrotriton cuchumatanus occurs in moss mats in Guate- mala, Lynch & Wake, 1975) or is missing in cloud forests of Nuclear Central America. he two Costa Rican transects offer some in- teresting contrasts. Both extend up the Caribbean about 55 km to the peak of Volcán Irazü. Sala- mander distributions along this transect are poorly known except for the region between 1,000 and 2,500 m. For example, Scott et al. (1983) listed two species of Oedipina from Finca La Selva, but study of specimens from the area sug- gests that three occur there. The taxonomy of the group is difficult, but it may be that none of the three are the species listed. (Note that only two species are indicated on Fig. 14.) There are pres- ently 11 species of salamanders (in three genera) known from this transect. As with the more northerly transects, only one species (Bolitoglos- sa subpalmata) is likely to occur at elevations o 7 3,000 m (and I cannot document its presence at that elevation as yet, although I expect it may be found) However, in contrast with more northerly transects (cf. Wake et al., 1987), there are at least five species present at elevations of species of Oedipina), and two Bolitoglossa (B. snags a nee that I have found in moss er microhabitats ds = moe "es sa P p ) and B. alvaradoi use bromeliads: I expect that more species will be found on the ds transect, in part because of the unusually umbers of species present on the Tapantí iue (Fig. 15). The latter transect 1s far more eneralized in its boundaries than the first, and ie essentially a broad (ca. 20 km) swath of ter- 1987] ritory extending about 60 km from the vicinity of Siquirres to the summit of Cerro de la Muerte. The Tapanti transect is not a straight line, but twists somewhat to encompass sites where some of the rarer species are known to occur. There is extensive habitat disturbance along this transect (for example, in the vicinity of Turrialba), and I justify weaving together an indirect transect as an attempt to demonstrate the potential number of species one might reasonably expect to find on a continuous altitudinal transect under pris- tine conditions. The Tapanti transect thus offers a greater diversity of habitats than does the Irazü transect and one might expect more species to be present. To my surprise, the Tapanti transect is the richest that I have found in the tropics, with 21 species. There are only 26 species of plethodontids known from Costa Rica (Scott et al., 1983; their list and mine differ slightly but we obtain the same total number of species), and that about 80% of them occur along this transect attests to its richness The Tapanti anadi has a higher number of species (seven) occurring below 500 m than does the Irazú transect, and in both Panama and Nic- aragua another species (Oedipina collaris) occurs at elevations of «500 m (Brame, 1968), so there is the likelihood that an eighth species eventually will be found. Bolitoglossa alvaradoi apparently occurs at elevations of < 500 m elsewhere in Cos- ta Rica. Thus, as many as nine species might be expected in the lowlands in the area crossed by this transect. Thirteen species occur in the cloud forest of the Tapanti transect, if we accept 750-2,000 m as its elevational bounds. In fact, cloud forest conditions exist almost to 3,000 m, although in general cloud forests are less well defined in Costa Rica and Panama than farther to the north (Myers, 1969). This transect contains the richest cloud forest salamander fauna found anywhere in the tropics. Here, too, I have probably under- estimated the number of species present. At least one more species of Bolitoglossa may be present, and the two poorly known highland species of Oedipina may well extend to lower elevations, since the genus as a whole is strongly concen- trated at lower elevations. Two species of Boli- toglossa (an undescribed species and B. dimi- nuta, Robinson, 1976) use mats of vegetation including mosses and liverworts that surround twigs and branches of trees. Several other species use bromeliads and moss mats, but no quanti- tative data are available. In the forests of Refugio WAKE—ADAPTIVE RADIATION OF SALAMANDERS 257 Tapanti there is an especially rich epiphyte fau- na, and this is the locality where the largest num- ber of species are found. However, in marked contrast to the situation in the cloud forest on our San Marcos transect, the density of individ- ual species is uniformly low on this transect. This situation of high species diversity and low den- sity of individual species is one more indication of the increased “‘tropicality” of the Costa Rican salamander fauna. The contrast between the cloud forest that oc- curs at around 1,000 m and the forest around 3,000 m on this transect is sharp. At high ele- vations the density of Bolitoglossa subpalmata is extraordinarily great, on the order of 9,000/ha (Vial, 1968). Four species occur in sympatry at around 2,500 m; in my experience B. subpalmata is about 100 times more common than B. cer- roensis, about 1,000 times more common than B. sooyorum, and about 10,000 times more com- mon than B. nigrescens! Doubtless there is a col- lecting artifact involved, but the first species is remarkably abundant and the last has been, at the very least, elusive. The high density of a sin- gle species at high elevation is a common theme in tropical salamander biology. Bolitoglossa subpalmata, which can be ex- ceedingly abundant at high elevations in Costa Rica, displays a marked shift in microbabitat tlower 1 utilization and a reduction in elevations. The species is primarily ground- dwelling at high elevation, although it also uses bromeliads. I have collected B. subpalmata from bromeliads 30 m above ground level in an oak tree at 3,000 m elevation. At elevations of «2,000 m the species becomes increasingly common in bromeliads and is encountered only infrequently in terrestrial situations. This species can be ex- traordinarily persistent in the face of even drastic habitat change, so long as bromeliads remain. An anecdote illustrates this point. An old col- lecting locality in the Montes de Aguacate, west of San José, was visited recently. Only tiny frag- ments of forest remain at this site, at about 1,500 m. Salamanders were common residents of bro- meliads in one forest fragment where trees were being felled. We opened 130 bromeliads and found 55 salamanders, including 14 adults and a subadult in a single bromeliad. As many as three adults were found in the axil of a single leaf. This is a graphic demonstration of the suit- ability of bromeliads as microhabitat for sala- anders, especially under conditions of great habitat modification. I have had similar expe- 258 riences with Chiropterotriton lavae in fragments of cloud forest above Jalapa, Veracruz, Mexico, and with Bolitoglossa morio in an area devas- tated by volcanic activity on Volcan Pacaya, Guatemala It is still too early to generalize very extensively from our comparative transect studies.We have come to be suspicious of species with broad el- evational ranges (such as O. uniformis, Figs. 14, 15), and these deserve careful taxonomic re-eval- uation, for most species occur within rather nar- row elevational limits. We also have come to expect few extreme highland or lowland species, but there are more lowland species at low lati- tudes. Cloud forests and salamanders are most abundant at mid-elevations. FACTORS INFLUENCING SALAMANDER SPECIATION AND RADIATION Species of plethodontid salamanders charac- teristically display a high degree of genetic frag- mentation (Larson et al., 1984; Larson, 1984). Larson (1984) has argued that a general pattern for the history of population structure in pleth- odontids is that following origination they ex- pand gradually and contiguously into regions to which they have ecological access. Later, as a result of climatic E which may result from many p causes, their populations become fragmented into ands among which there is little or no genetic exchange. Subsequent climatic changes may lead to re-establishment of ecolog- ical access to areas separating isolated popula- tions. This may lead either to renewed genetic exchange or, depending on the level of genetic divergence and its effect on isolating mechanisms or mate recognition systems, to a variety of in- teractions. There may be a hybrid swarm, a nar- row hybrid zone, a narrow zone of overlap with occasional hybridization, partial sympatry with boundaries set by competitive interactions, or co- existence with different ecological requirements. All of these interactions have been documented among plethodontids. The many studies of pop- ulation structure based on electrophoretic anal- ysis of proteins, together with studies of distri- bution, phylogenetic history, and biogeography (reviewed by Larson, 1984), suggest that geo- graphic or allopatric speciation by subdivision is the common mode in plethodontids. There are similarities with the concept of “taxon cycles” (Wilson, 1961; Ricklefs & Cox, 1972), which usually have involved examples from island ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 chains. These postulated cycles of expansion and contraction of geographic ranges and population densities would establish the setting for the kind of speciational processes I envision having taken place among the tropical salamanders. o primary factors in the history of the tropical salamander fauna probably have been the combination ofadaptation of species to cloud forest environments and the long, complex his- tory of tectonic activity in Middle America. There are some relatively stable areas, such as the an- cient core of Nuclear Central America, where the upper and lower boundaries of the cloud forest have shifted, but where cloud forest in a broad sense probably has been present for much of the Tertiary and Quaternary periods (see discussion in Wake & Lynch, 1976). However, there are many other areas where tectoni ts and associated volcanic activity have been so great and so persistent that cloud forests have been only an ephemeral presence, shifting almost con- stantly. The three foci for salamander radiation in the tropics (Fig. 2) all have stable, ancient tectonic core areas surrounded by regions of great tectonic activity. (For summary of tectonic history, see Hendrickson, 1986.) These areas have been cited by other workers as having phylogenetically dis- tinctive faunal components. For example, Sav- age (1982) stated: I now believe that the distinctive montane her- petofaunas of the southern Sierras of Mexico, Nu- more or less similar sea of widely distributed ancestors. Nuclear Central America (Fig. 16) illustrates the ideas outlined above. That region has as its core the ancient Sierra de los Cuchumatanes, lo- cated on the southern part of the North d plate. The area has been characterized as th Middle American Megathrust by Plafker (1976), because it is the conjunction of three major plates. The Cocos plate, to the southwest, is being sub- ducted near the intersection of the North Amer- ican and Caribbean plates (for evidence of the widespread influence of this phenomenon see Singh et al., 1985). The latter is being forced eastward by the combined plate movements, and a small western tip of the plate is effectively "caught" between the North American and the Cocos plates. The zone between the Caribbean 1987] WAKE—ADAPTIVE RADIATION OF SALAMANDERS 259 Isthmus of | Tehuantepec MEXICO HONDURAS Inspired by and based largely on Plafker (1976). and North American plates is outlined by the Motagua and Polochic fault zone. This has been an area of intense tectonic activity for millions of years and Plafker (1976) has estimated that at least 200 km of lateral movement of plates along this fault zone has occurred since the Miocene. The western tip of the Caribbean plate, trapped between the other plates, is being ripped, or de- coupled (Plafker, 1976). Grabens have formed, with small volcanic cones rising within them. volcanoes are lined up. Volcan Tajumulco and Volcan Tacana lie at the northwestern corner of the Caribbean plate, where the three plates meet. In this topographically complex zone of maximal geological turbulence, the largest number of co- ins of the core region are areas in large part to the fragmentation and reassembly of areas of cloud forest. occurring species of salamanders in Middle America is found, along our San Marcos transect. The relatively stable upland of the Sierra de los Cuchumatanes is in many ways an even more interesting area than the Pacific volcanic belt. We began fieldwork in this area in 1974, at a time when only two species of salamanders were known from the Caribbean slopes of the Cuchu- matanes, despite a number of brief LUIS trips by different herpetologists. Results of our inves- tigations have been summarized by Elias (1984), who found 13 species of salamanders in this re- gion (see also Wake et al., 1987). As elsewhere in Middle America, the cloud forest is of special interest, for six species with narrow elevational ranges occur just above the cloud line, here lo- cated at about 1,300 m. Two new genera of sal- amanders were discovered in this cloud forest 260 (Elias & Wake, 1983; Wake & Elias, 1983), in- cluding the exceptional Nyctanolis, a morpho- logically primitive genus which appears to be the sister group of all other tropical salamanders. Thus, on the one hand, the ephemeral cloud for- ests of the tectonically active and topographically complex margins of the Cuchumatan uplands have contributed to speciation and led to the highly disjunct distributional patterns illustrated previously for Dendrotriton and Nototriton (Figs. ). On the other hand, the more stable cloud forests on the northeastern slopes of the core of the Cuchumatan region have served as refugia for what must be extremely ancient lineages. CONCLUSIONS We still have much to learn about the sala- manders of the New World tropics, and even the best known areas of Middle America have yield- ed many recent surprises. Earlier misconceptions concerning the probable history and ecology of tropical salamanders have led to underestimates of the age and diversity of the group and have contributed to our relative ignorance of the ecol- ogy of the cloud forest and lowland species, es- pecially the arboreal and fossorial forms. New species are being discovered more rapidly than they can be described, for many species are known from small series. Our knowledge of the com- parative osteology and of molecular evolution of this group, while still fragmentary, is sufficient to demonstrate that parallelism and convergence are rampant. This, in turn, implies both that there may be only a limited number of ecological roles avail- able to tropical salamanders, and that there may be functional and developmental-historical con- straints which impose limits on the evolutionary potential of the group. An especially clear case of convergence is the elongation associated with fossorial life in the genera Oedipina (from south and east of the Isthmus of Tehuantepec) and Li- neatriton (from north of the Isthmus) (Fig. 12). The former has become elongate by increasing the numbers of vertebrae; the latter by increasing the length of the individual vertebrae, which are identical in number (in the trunk) to all tropical genera except Oedipina (Tanner, 1950; Wake & Lynch, 1976). Earlier in this paper I highlighted the convergence in the Chiropterotriton-Dendro- triton-Nototriton assemblage. Within Bolitoglos- sa webbing of hands and feet has evolved both convergently (Alberch & Alberch, 1981) and in ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 parallel (Wake, 1966; Wake & Brame, 1969; Lar- son, 1983; Elias, 1984). These phenomena have made phylogenetic analysis very difficult, for any phylogenetic hypothesis requires extensive con- vergence, parallelism, or evolutionary reversal (Wake & Elias, 1983). Epiphytes appear to have been significant fac- tors in the evolution oftropical salamanders. The convergence in morphology and behavior of bromeliad-dwelling salamanders is one indica- tion of this importance, and the extent to which bromeliads and moss mats are utilized as mi- crohabitats is another. It is also significant that salamanders persist in the face of great environ- mental change so long as fragments of forest with the preferred microhabitats remain. The tropical salamanders, which originated from a Laurasian ancestral group (Savage, 1973), are a marked exception to a common pattern of tropical origin and subsequent temperate inva- sion. Late in the history of but nevertheless a very long time ago (perhaps at the beginning of Tertiary), they invaded the area that has become modern Middle America. Compared with salamanders generally, the trop- ical salamanders have been phenomenally suc- cessful. But now their survival and, indeed, the survival of much of the diversity of tropical eco- systems is at risk, for the lowland forests and the middle elevation cloud forests that harbor most tropical salamanders are being cleared at rates that almost defy belief. Not a tree is standing over extensive parts of our San Marcos transect, which was in an almost pristine state as recently as 1969. In a single human lifetime the results of perhaps a hundred million years of evolution will have been dramatically changed, if not ex- tinguished. group, LITERATURE CITED ALBERCH, P. 1981. Convergence and parallelism in foot morphology in the neotropical salamander genus Bolitoglossa. I. Function. Evolution 35: 84- 100 1981. & J. ALBERCH. Heterochronic mecha- Bolitoglossa e VE aa Plethodon- tidae). J. Morphol. 167: SJ. DE, G. F. Osr Arn B. WAKE, 1979. Size and shape in ontogeny and phylogeny. Paleo- biology 5: 296-317 BRAME, A. H., JR. 1963. A new Costa Rican sala- mander (genus Oedipina) with a re-examination of O. collaris and O. serpens. Nat. Hist. Mus. Los Angeles Co.. ` Contrib. Sci. 65: 3-12. 1987] , 1968. i uestrum and evolution (bs Meso- ipina J. Herpetol. 2: 2-64. . B. WAKE. 1963. The salamanders of South America. Nat. Hist Mus. Los Angeles Co., Contrib. Sci. Fé 1-72. New species of salamanders (genus E from Colombia, Ecuador and Panama. Nat. Hist. Mus. Los Angeles Co., Con- trib. Sci. 219: 1-34. BREEDLOVE, D. E. 1981. Introduction to the flora of Chiapas. Pp. 1-35 in Part I, Flora of Chiapas. California Academy of Sciences, San Francisco. DuNN, E. R. The Salamanders of the Family Plethodontidae. Smith College, Northampton, Massachusetts. 1937. The amphibian and reptilian fauna of bromeliads in Costa Rica and Panama. Copeia 1937: 163-167. Erias, P. en Salamanders of the northwestern highlands of Guatemala. Nat. Hist. Mus. Los An- geles Co., Neer Sci. 348: 1-20. ——— & D 1983. Nyctanolis pernix, a new genus nd dedica of plethodontid salamander from aru ci qn and Chiapas, Mex- urs . J. Rhodin & K. Miyata (editors), ie in Su onu. and Evolu- tionary Biology: Essays in Honor of Ernest E. Wil- liams. Mus. Comp. Zool., Cambridge, Massachu- setts. FEDER, M. E. 1978. Environmental variability and thermal acclimation of metabolism in neotropical and temperate zone salamanders. Physiol. Zool. 51: 7-16 1982. Thermal ecology of neotropical lung- less salamanders (Amphibia: Plethodontidae): en- vironmental temperatures and behavioral re- sponses. Ecology 63: 1665-1674 983. Integrating the ecology and physiology of. plethodontid salamanders. Herpetologica 39: p F. LvucH. 1982. Effects of latitude, sea- son, elevation, and microhabitat on field body temperatures of neotropical Ae AONAN zone salamanders. Ecology 63: 1657- Frost, D. R. (editor). 1985. Pine rad Species of the World. Allen Press, Inc. & Assoc. Syst. Collec., Lawrence, Kansas Gapow, H. 1908. Through Southern Mexico. With- erby & Co., London. GRUBB, P. J. & T. C. WHITMORE. 1966. A comparison of montane and lowland rain forest in Ecuador. II. The climate and its effects on the distribution and physiognomy of the forests. J. Ecol. 54: 303- 330. HANKEN, J. 1983. Genetic variation in a dwarfed and evolutionary implications. Copeia 1983: 1051- 1073. WAKE. 1982. Genetic xig ie among plethodontid icr ge (genus Bolito- glossa) in Central and South America: "mpi tions for the South M ian invasion. Herpeto- logica 38: 272-287. WAKE- ADAPTIVE RADIATION OF SALAMANDERS 261 — ——., J.F. LyucH & D. B. WAKE. 1980. Salamander invasion of the tropics. Nat. Hist. 89(12): 46-53. HENDRICKSON, Congruence of bolito- glossine biogeography and phylogeny with geolog- ic history: paleotransport on displaced suspect ter- ranes? Cladistics 2: 113-129. Houck, L. D . Reproductive biology of a neo- tropical salamander, Bolitoglossa rostrata. Copeia 1977: 70-83. . 1977b. Life history patterns and i biology of neotropical salamanders. Pp. 43-72 i . Taylor & S. I. Guttman (editors), The Re. productive Biology of Amphibians. Plenum Press, New Yor LARSON, A. P A molecular phylogenetic per- spective on the origins of a lowlan Neontological inferences of evolution- ary pattern and process in the salamander family Plethodontidae. Pp. 119-217 in M. K. Hecht, B. Wallace & G. T. Prance (editors), Evolutionary ieee Vie 1 Plenum Publ. Co., New York. K. P. YANEvV. 1984. Measuring gene x db among populations having high levels of genetic fragmentation. Genetics 106: 293-308. LOMBARD, R. E. & D. B. WAKE. 1977. Tongue evo- lution in the lungless salamanders, family Pleth- odontidae. II. Function and evolutionary diver- sity. J. Morphol. 153: 39-80. Tongue evolution in the lung- pe s family Plethodontidae. IV. Phy- logeny of plethodontid salamanders and the evo- lution of feeding dynamics. Syst. Zool. 35: 532- 5L 5 LvNcH, J. F. & D. B. WAKE. 1975. The systematics of the C Tn bromeliacia group (Am- phibia: Caudata), w description of two new IAS from «i eh Nat. A Mus. Los An- les Co., Contrib. Sci. 265: 1- i new Ls of Chiropter- iol (Amphibia: Caudata) from Baja Verapaz, mala, with comments on relationships among C md "ayara members of the genus. Nat. Hist. Mus. Los bees Co., Contrib. Sci. 294: 1-22. Y. YANG. 1983. Genic and mor- phol al di the sal- am ander r genus M na inhabiting the 1958. A biogeography of reptiles and amphibians in the Gómez Farias region, Tamau- lipas, Vw Misc. Publ. Mus. Zool. Univ. Mich- igan E. -102. B. E. Taf 1957. tory of temperate biotas in Mexico United States. Ecology 38: 468-4 E The Pleistocene his- and eastern Maxson, L B. E. 19 A bumin evolution and its phylogenetic implications in the pletho- enc anas era Pseudoeuryce dar Chi- l 17 ropterotriton. Herpetologica 37: 9-1 s,C.W. 1969. The ecological geography of cloud forest in Panama. Amer. Mus. Novit. 2396: 1-52. PAPENFUSS, T. J., D. B. WAKE & K. ADLER. 1983. Salamanders of the genus Bolitoglossa from the Sierra Madre del Sur of southern Mexico. J. Her- petol. 17: 295—307. PicApo, C. 1913. Les bromeliacées pus con- ]. Sci. France Belg. 7: 215-360. PLAFKER, G. 1976. Tectonic aspects of the Guate- malan earthquake of 4 February 1976. Science On certain Mexican salamanders of the plethodontid genus Chiropterotriton. Occ. Pap. Mus. Zool. Univ. Michigan 587: 1—37, pls. I-III. 1960. A new salamander of the genus Chi- ropterotriton from Chiapas, Mexico, n notes on related Ade Copeia 1960: 304- idi . E. & G. W. Cox. 1972. m cycles in the us Indian avifauna. Am. Nat. 106: 195- 219. M D. C. 1976. A new dwarf salamander of he ge us Cos- ta Rica. Proc. Biol. Soc. Washington 89: 289-294. 1977. Herpetofauna bromelicola costarri- cense 4 goya de Hyla picadoi Dunn. Pp. 31- D. Gómez (editor), Historia Natural de ia de las Bromeli- tional morphology and sensorimotor control of — behavior in salamanders: an example of the role of internal dynamics in evolution. Acta ia eae 34: 175-192. & 1985b. The structure of the brain- stem and cervical spinal cord in relation to feeding of lungless salamanders, family Plethodontidae. J. Comp. Neurol. 241: 99-110 RUTHVEN, A. G. The amphibians and reptiles of the Sierra Nevada de Santa Marta, Colombia. imi Publ. Mus. Zool. Univ. Michigan 8: 5-69, Is pls. 1-12. SAVAGE, a M. 1973. The geographic s of 5 in J. l enigma of Central AMEN herpetofauna: dispersals or vicariance? Ann. Mis- souri Bot. ae rd. 69: 464—547. SCHMIDT, K. atemalan salamanders of 1936a. Gu the pa Cone Zool. Ser. Field Mus. Nat. Hist. 20: 135- m New D and reptiles from Honduras in the Mus of Comparative Zool- ogy. Proc. Biol. Soc. Washington 49: 43-50. 1942. A cloud forest camp in Honduras. Chi- cago Nat. 5: oem 30. Scorr, N. J., J. M. SAVAGE & D. C. RoBINSON. 1983. Checklist of reptiles and amphibians. Pp. 367-374 in D. H. Janzen (editor), Costa Rican Natural His- ry. Univ. Chicago Press, Chicago. SHAFFER, H. B. 1978. Relative predation pressure on salamanders (Caudata: Plethodontidae) along an altitudinal transect in Guatemala. Copeia 1978: 2- SHANNON, F. A. & J. E. WERLER. 1955a. Notes on amphibians of the Los Tuxtlas Range of Veracruz, Mexico. Trans. Kans. Acad. Sci. 58: 360-386. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 1955b. Report on a small collection of vam from Veracruz, with a description of a new species of Pseudoeurycea. Herpetologica 11: 81-85. SINGH, S. K., G. SUAREZ & T. DOMINGUEZ. 1985. The Oaxaca, Mexico, earthquake of 1931: lithospheric normal faulting in the subducted Cocos Plate. Na- ture 317: 56-58. . 1945. Herpetological collecting in ba- nana fields of Mexico. Ward's Natural Science Bull. 1945: 3-7. SruART, L. C. 1943. Taxonomicand geographic com- ents on Guatemalan salamanders of the genus Oedipus. Misc. Publ. Mus. Zool. Univ. Michigan 56: 1-33, pls. I-II. 1948. The amphibians and reptiles of dein Verapaz, Guatemala. Misc. Publ. Mus. Zool. Uni Michigan 69: 5-107. 1954. Descriptions of some new amphibians and reptiles from iro emala. Proc. Biol. Soc. Washington 67: 159-1 TANNER, W. W. 1950. re new genus of plethodontid a from Mexico. Great Basin Nat. 10: 7-44. n E.H. 1941. New amphibians from the Ho- M. Smith Mexican collections. Univ. Kansas ae all 27: 141-167. 1942. New caudata and salientia from Mex- ico. Univ. Kansas Sci. Bull. 28: 295-323. 1944. The genera of plethodont salamanders in Mexico, Pt. 1. Univ. Kansas Sci. Bull. 30: 189— 232 . 1952. The salamanders and caecilians of Cos- ta Rica. Univ. Kansas Sci. Bull. 34: 695-791. 1954. Additions to the known herpetological fauna RE Costa Rica with comments on other . No. I. Univ. Kansas Sci. Bull. 36: 597- & H. M. SmitH. 1945. Summary of the col- lection of amphibians made in Mexico under the Walter Rathbone Bacon ied Scholarship. Proc. U.S. Natl. Mus. 95: 3. TRAPIDO, H. 1942. A new d oar from Vene- zuela. Bol. Soc. Venezolana Cien. Nat. 8: 297-301. VIAL, J. L. 1968. The ecology of the tropical sala- mander, Wa rs Va ire in Costa Rica. Rev. Biol. Tro l WAKE, D. B. 1966. Co oIBDBIA NE osteology and evo- lution of the lungless salamanders, amit Pleth- odontidae. Mem. Men alif. Acad. Sci. H. BRAME, JR. 1 ys stematics and evolution of n pea salamanders of the Bo- litoglossa helmrichi group. Nat. . Mus. Los ngeles ntri : . ELIAS. genera | and a new species of Central Ameri rican salamanders, with a review of the tropical oe ra (Amphibia, Caudata, eee Nat. Hist. Mus. Los Angeles Co., Contrib. Sci : 1-19. E 1976. The distribution, ecol- Ogy, and evolutionary history of plethodontid sal- amanders in tropical i Nat. Hist. Mus. Los Angeles Co., Sci. Bull. 25: & Pid relationships among Central American salamanders of the Bo- 1987] WAKE—ADAPTIVE RADIATION OF SALAMANDERS 263 litoglossa | Mg ie group, with a description of a om Guatemala. Herpetologica 38: ME & W. E. DUELLMAN. 1973. New species d intende genus Bolitoglossa, from Panama. Nat. Hist. Mus. Los Angeles Co., Con- trib. Sci. 248: 1-1 9. Rs. 1970. Bolitoglossa taylori, a new salamander from cloud forest of the Serrania de Pirre, eastern Panama. Amer. M Novit. 2430: 1-18. — T. J. PAPENFUSS & J. F. LvNcH. 1987. Dis- tribution of salamanders along elevational tran- sec e m ER and Guatemala. Brenesia (Suppl.) (in nd c. F. 1955. A new salamander of the m Pseudoeurycea from Tamaulipas. bs Pap Zool. Univ. WW e 567: 1-8, pl. WERLER, J. E. & H. M. SMITH. vog “Notes ona collection of db and amphibians from Mex- ico, 1951-1952. Texas J. Sci. 4: 551-573. WILSON, E. O. 61. The nature of the taxon cycle in the Melanesian ant fauna. Amer. Nat. 95: 169- 193. APPENDIX I Use of bromeliads and moss mats by neotropical salamanders. All species known to occur = either of these microhabitats are listed below. Literature references do not include all observations for a given species, but either the first or the best documented example. Where no eive Aia Mad field notes or notes from museum collections made by the author are cited. > š B. yucatana Chiropterotriton arboreus C. chiropterus C. chondrostega C. lavae C. multidentatus Dendrotriton bromeliacia D. megarhinus D. rabbi D. xolocalcae Nototriton barbouri Bromeliad Occurrence: roni tase pete DBW notes B. arborescandens Taylor Lay B. Me Trapido (1942) B. cuchumatana Stuart Ev Elias (1984) B. dunni Schmidt (1942 B. engelhardti Schmidt (1936a), Wake & Lynch (1976) B. flavimembris Wn B. franklini Mid (1941), Wake & Lynch (1976) B. hartwegi e & Brame (1969), Elias (1984) B. helmrichi uds t (1936a B. hermosa diae et al. (1983) B. jacksoni Elias (19 ay B. lignicolor Dunn (19 lincolni Wake & im EA. Elias (1984) B. meliana Wake & Lynch (1982) ) Wake & Lynch (1976), Elias (1984) Stuart (1943) Brame & Wake (1963) Shannon & Werler (19552) Stuart (* "probably, " 1948) Schmidt (19362), Taylor & Smith (1945) Ruthven (1922) Dunn (1937) Wake et al. (1970) Brame & Wake (1963) Brame & Wake (1972) DBW notes Rabb (1955) DBW notes Martin (1958) 958 Schmidt (1936a), Wake & Lynch (1976) dei 960) as & ie (1975), Elias (1984) or (194 s EH 264 ANNALS OF THE MISSOURI BOTANICAL GARDEN APPENDIX I. Continued N. nasalis N. picadoi N. veraepacis oeil ea bellii unnata piel firscheini goebeli leprosa . nigromaculata DVT BWR 2 3 . smithi Thorius dubitus Moss Mat Occurrence: Bolitoglossa diminuta ; is dias B. subpalm n cuchumatana, Nototriton ns N. richar Myetancls perni x Oedipina poelzi O. pseudouniformis . uniformis Pseudoeurycea rex P. scandens P. werleri Thorius dubitus Dunn (1926), Schmidt (1942) Picado (1913 Lynch & Mrs (1978) DBW notes from T. J. Papenfus Wake & Seh TUA ie DBW notes Stuart (1954) Werler & Smith (1952), Shannon & Werler Schmidt (1936a), and DBW notes DBW notes DBW notes Walker (1955), Martin (1958) DBW notes from T. J. Papenfuss DBW notes Robinson (1976 — in a liverwort mat) Wake et al. (1973) Taylor (1952) Lynch & Wake (1975) Taylor (1954 notes Elias & Wake (1983) ) 58) Shannon & Werler (19552) Taylor (1941) [VoL. 74 REVISION OF EREMANTHUS (COMPOSITAE: VERNONIEAE)! NANDA F. F. MACLEISH? ABSTRACT Historica d 1 £ dias & T hi d ly I traits of s de dus aggregation of heads into a glomerule), pluriseriate involucres, and achenes with a persistent pappus. Syncephaly i is characteristic of the Lychnophorinae, which is con- In the present treatment Eremanthus includes sidere Vanillosmopsis, but excludes Albertinia, Chresta, Glaziovianthus, Prestelia, Pycnocephalum, Paralych- mopsis taxa) are presented. Recent studies have shown that Eremanthus Less. (Compositae: Vernonieae) includes both Eremanthus Less. (sensu Schultz-Bip.) and Vanil- Josmopsis Schultz-Bip., but excludes Albertinia Sprengel, Chresta Vell. Conc., Glaziovianthus G. Barroso, Prestelia Schultz-Bip., oo (Less.) DC., Sphaerophora Schultz-Bip., and Stachyanthus DC. (MacLeish, eo pv 1984c, 1985a, 1985b, and MacLeish & Schu- macher, 1984). Thus, Eremanthus comprises 18 species of syncephalous trees and shrubs largely restricted to cerrado (sensu Eiten, 1978, 1983) of the arid Central Plateau of Brazil. This is a regibn dominated by woody Compositae; geo- ly rare in the family 4 Heywood et al., 1977). The name Eremanthus is derived from the Greek eremos (solitary) and anthos (flower bear- ing) in reference to the heads, which bear single flowers. Vanillosmopsis refers to the vanillalike odor characteristic of this group of plants. Vanil- losmopsis is locally called candeia, candle, whic refers to its ability to burn luminously. The close relationship between Vanillosmopsis and mem- bers of Eremanthus sensu Schultz-Bip. is fre- De Candolle (1836) is the only other daha to combine Eremanthus and Vanillosmopsis, al- though he mistakenly placed them in Albertinia. He correctly noted that E. incanus and E. elaeag- and Stachyanthus (now Argyrovernon nia). Eremanthus thus com- nus approach Vanillosmopsis in degree of syn- cephaly, number of florets per head, and achene and pappus characters; and that V. erythropappa and V. polycephala resemble Eremanthus in their high degree of syncephaly and, in the latter, re- duced number of florets per head. In the course ofthe present study, numerous specimens of nat- ural hybrids between these two genera were iden- tified. It soon became apparent that the taxo- nomic distinction between Eremanthus and Vanillosmopsis was largely artificial. Host/parasite relationships and terpenoid chemistry support the close relationship of these mopsidia H. S. Jackson & Holway, has been found on E. mattogrossensis and on V. erythropappa. Because rusts (Uredinales) have evolved with their vascular plant hosts, their taxonomy may suggest relationships among vascular plant taxa (Thorne, 1979; McCain & Hennen, 1982). Urban (1973) demonstrated the systematic utility of Puccinia and Vernonia coevolution in determin- ing subgeneric categories. Hence, a close rela- tionship between Eremanthus and Vanillosmop- sis is supported by the presence of a single rust species on members of both genera. In addition, terpenoid profiles appear to be very similar in members of Eremanthus and Vanillosmopsis (H. Schumacher, pers. comm.). Also, many authors have noted the presence of identical terpenoid ! Work at the University of Georgia was supported partially by National Science Foundation Grants DEB t for 81-13522 and ee 79-09831. I am grateful to the Brazilian governmen o members of the Rio de Janeiro Jardim Botánico for generous assistance. In addition, I and, in particular, the opportunity to collect in Brazil thank the E of the ppu herbaria who loaned specimens ae pe project rie ll Baker, Graziela M. Barroso, Christopher Meach Depart m, and Heiko Schumacher, who we especial assis ment of Library Science and Information cee aan [ceni tei. ym Georgia 30322, ANN. Missouni Bor. GARD. 74: 265-290. 1987. 266 yav sisqa in the two groups, one further sup- (Bake ., 1972; Gar cia et al., 1976; Vichnewsid et P 1977; Har- borne & Williams, 1977; Robinson et al., 1980). The systematic utility of terpenoids in members of Vernonieae has been discussed by Abdel-Baset et al. (1971), Mabry et al. (1975), Harborne & Williams (1977), and Robinson et al. (1980). Eremanthus can be distinguished from other syncephalous genera by a combination of char- acteristics (MacLeish, 1984a, 1984c). These in- clude woody habit; indument composed pri- marily of stellate hairs; terminal compound cymes (rarely reduced to a cyme) of glomerules; 1-4 florets per head; 10-ribbed (rarely 20-ribbed) achenes with 3—5-seriate pappus and prominent nectary; and advanced pollen of type A (sensu Keeley & Jones, 1977). Members of Eremanthus have complex glomerules derived from reduc- tion of internodes between subglomerules ar- ranged in a compound cyme. A complete dis- cussion of glomerulescence arrangement and derivation for Eremanthus and related genera can be found in chapter 1 of MacLeish (1984c). The only other syncephalous Vernonieae that ap- proach the combination of characteristics evi- dent in Eremanthus are Paralychnophora MacLeish and Lychnophora Martius (MacLeish, 1984c). However, both of these have a biseriate pappus and glomerules derived from the inter- node reduction of corymbose subglomerules. In addition, Paralychnophora is characterized by axillary, solitary glomerules and angled achenes; and Lychnophora by ericoid leaves and promi- nently twisted inner pappus series. mong nonsyncephalous Vernonieae, Ere- manthus appears to be related to Vernonia sub- sect. Eremosis (DC.) S. B. Jones (MacLeish, 1984c). This subsection also has a woody habit, congested terminal capitulescences, reduced numbers of florets per head, ribbed achenes, and pollen of type A. However, this group is distrib- uted primarily in Central America and northern South America and has a biseriate pappus. If Eremanthus is derived from Eremosis stock, there are two Brazilian refugia that may have served as centers of diversification and dispersal: Vea- deiros and Rondônia (also called “Araguaia” and * Airpuana" by Prance, 1982). Veadeiros is the current site of several locally endemic members of sect. Nectaridium (subg. Vanillosmopsis) that are virtually indistinguishable from many mem- bers of Vernonia subsect. Eremosis. Rondónia contains a single Eremanthus species, E. ron- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 doniensis, which is restricted to that area and is the most unspecialized member of subg. Ere- manthus. TAXONOMIC HISTORY Historically, A/bertinia Sprengel, Chresta Vell. Conc., Glaziovianthus G. Barroso, Pithecoseris DC., Prestelia Schultz-Bip., Pycnocephalum (Less.) DC., Sphaerophora Schultz-Bip., Stach- yanthus DC., and Vanillosmopsis Schultz-Bip. have been placed adjacent to or in synonymy with Eremanthus. These genera are restricted to the arid Central Plateau of Brazil and have not been evaluated comprehensively since Baker’s (1873) treatment of Compositae in Martius’s Flora Brasiliensis. In 1829, Lessing established Eremanthus to include syncephalous trees or shrubs with one floret per head and a paleaceous pappus. At the same time he placed syncephalous trees and shrubs with (presumably) numerous florets per head, filiform pappus, and alveolate receptacles Ibertini d syncephalous h ernonia sect. Pycnocephalum. Also in 1829, two species referable to Eremanthus s. lat. were described by Velloso as Chresta: C. cordata and C. lanceolata. However, the portion of Flora Fluminensis con- taining Chresta was not effectively published un- til 1881 (Carauta, 1973). Thus, de Candolle ac- complished valid publication of Chresta in 1836 when he cited Velloso's figures, which had been published in Florae Fluminensis Icones (Velloso, 1831). Of the two species described by Velloso, only C. cordata was included by de Candolle in Chresta (as C. spherocephala). This distinctive taxon is widespread throughout Brazil. In con- trast, the figure and description of C. /anceolata could represent any one of several species. If one takes into account the itineraries of Velloso and contemporary collectors (Urban, 1906), C. /an- ceolata is either Vanillosmopsis erythropappus, Eremanthus incanus, E. glomerulatus, or E. mattogrossensis. Kuntze (1898) believed this taxon to be synonymous with V. erythropappus but provided no account of his reasoning. In the present treatment, C. /anceolata is excluded be- cause of doubtful identity, and C. cordata is re- tained in Chresta. Two years later, Lessing (1831) dissolved A/- bertinia when he transferred the type species into Vernonia, and a second taxon, correctly consid- ered to have one floret per head, into Ereman- thus. In 1836, de Candolle treated syncephalous taxa as five different genera. Syncephalous trees = 1987] and shrubs were included in Albertinia (he in- correctly assumed that the type species had a single floret per head), and syncephalous herbs were placed variously in Chresta, Pycnocepha- lum, Stachyanthus, and Pithecoseris. Schultz-Bi- pontinus (1861, 1863) recognized the error im- plicit in de Candolle's interpretation of the deeply pitted receptacle A/bertinia as fused heads ho- mologous with those found in many Eremanthus species. In contrast, Schultz-Bipontinus consid- ered the heads of Albertinia to contain many florets, each of which is partially enclosed in a receptacle cavity, and the genus to be monotypic and closely related to Vernonia. However, Schultz-Bipontinus placed taxa with one (rarely three) floret(s) per head, high degrees of synceph- aly, and achenes with a persistent pappus in Er- emanthus or the newly-described Sphaerophora (further delimited by achene surface features), and those with three (to one) florets per head, various degrees of fusion, and achenes with a deciduous pappus in Vanillosmopsis, a second new genus (Schultz-Bipontinus, 1861, 1863). In 1873, Baker followed Schultz-Bipontinus in his delimitation of Vanillosmopsis; however, in his classification, Eremanthus comprised not only the species of Schultz-Bipontinus, but also those of Sphaerophora, Chresta, Pycnocephalum, Pres- telia, and Stachyanthus. Except for Barroso's (1947) transfer of herbs with yellow and red co- rollas to Glaziovianthus, and Robinson's (1980) suggested recombination of all herbaceous taxa into Chresta, the boundaries of Eremanthus and related genera have remained largely as Baker delineated in 1873 Following Baker, curators endeavoring to identify Brazilian specimens have placed most syncephalous Vernonieae with achenes bearing a persistent pappus in Eremanthus. However, syncephalous heads are characteristic of Lych- nophorinae, which is currently considered to be an artificial assemblage (Jones, 1977; MacLeish, 1984c). Traditionally, the distinction between Eremanthus and Vanillosmopsis has been based on the presence or absence of syncephalous heads, one versus three pue r head, and paleaceons and persiste pus. In fact, ihe characteristic of lacking sineephe aly i used w exclude Vanillosmopsis from Ly aced) despite generic descriptions that clearly pee Vanillosmopsis as including several highly syn- cephalous taxa (Baker, 1873; Bentham, 1873). De Candolle is the only author until now to com- MACLEISH — EREMANTHUS REVISION 267 bine Eremanthus and Vanillosmopsis, although erroneously, in Albertinia. Currently, following de Candolle, Eremanthus is considered to in- clude trees and shrubs with reddish-purple co- rollas, compound cymes (rarely reduced to a cyme) of glomerules, 1—4 florets per head, ribbed achenes with persistent or deciduous pappus of 3-5 intergrading series, and advanced pollen of type A (subechinolophate with elongated ger- minal furrows often barely separated at the poles, sensu Keeley & Jones, 1977). e Candolle and Schultz-Bipontinus used the number of florets per head, degree of syncephaly, number of pappus series, and various achene sur- face features to delineate subdivisions of genera. These are good characters that readily indicate natural groups of related species. Thus, subg. Er- emanthus includes sect. Eremanthus and sect. Synglomerulus. Eremanthus sect. Eremanthus is a group of closely related taxa (E. glomerulatus, 7. goyazensis, E. mattogrossensis, and E. ron- doniensis) that may have arisen along an east- west gradient across southern portions of the Central Plateau (Fig. 1). This section is charac- terized by coriaceous leaves, a compound cyme of 10-60 glomerules, 5-100 heads per glomerule, head fusion via the interwoven pubescence of phyllaries, one floret per head, and sericeous achenes with a stramineous, coroniform, persis- tent, paleaceous pappus. Section Synglomerulus comprises Eremanthus argenteus, E. auricula- tus, and E. cinctus. They differ from sect. “Ere. manthus primarily in reduction of the glomer- ulescence to a cyme (not compound) of 2-10 glomerules E omaniku subg. Vanillosmopsis comprises two sections and is characterized by membra- naceous to subcoriaceous leaves, compound cymes of more than 100 glomerules, 1-12 heads per glomerule, 1—4 florets per head, and glabrous achenes with a purple or white (rarely stramin- eous), not coroniform, deciduous, filiform pap- pus. Eremanthus pohlii, E. graciellae, and E. brasiliensis form sect. Nectaridium, which is characterized by near absence of syncephaly (heads solitary or slightly appressed basally in pairs) and is restricted to the northwestern arm of the Central Plateau. Eremanthus capitatus, E. arboreus, and E. uniflorus are members of sect. 'anillosmopsis and have 2-9 heads per glomer- ule, with bases slightly appressed and fused by the interwoven pubescence of the phyllaries. The first two of these species range south to north along the northeastern arm of the Plateau, and 268 À sect. Eremanthus subg. Eremanthus @ sect. Synglomerulus * subg. Isotrichia X subg. P d h A sect. Vanillosmopsis subg. Vanillosmopsis © sect. Nectaridium A FIGURE 1. the last is restricted to the dde sa: arm Eremanthus erythropappus and E. polyc have glomerules with 6-12 heads that are closely appressed and fused by the concrescence of phyl- lary and receptacle tissues. These two species are found only in the southern part of the Central Plateau. In contrast, three species (E. incanus, E. elaeagnus, E. seidelii), historically assigned to Eremanthus s. str., exhibit characteristics inter- mediate to those of both subg. Eremanthus and subg. Vanillosmopsis. Not surprisingly, these species are largely restricted to the southern part of the Plateau, an area where the two major com- plexes are sympatric. This is also the region where a majority of hybrids are found, and it is possible that subg. Pseuderemanthus (E. elaeagnus and E. seidelii) and subg. Isotrichia (E. incanus) rep- resent established hybrids between members of the major complexes. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Distribution of Eremanthus in the Central Plateau of Brazil. TAXONOMIC TREATMENT Eremanthus Less., Linnaea 4: 317. 1829. TYPE: Eremanthus glomerulatus Less. a S e Schultz-Bip., Jahresber. Pollichia 18— 16 61. TYPE: Vanillosmopsis glomerata EU Bip. = Eremanthus erythropappus (DC.) MacLeish. Trees or shrubs to 10 m tall; stems reddish- brown lanate-tomentose or gray lepidote-tomen- tulose, often darkened basally by fire; branches few to many, often tortuous. Leaves bna al- ternate, strongly coriaceous to membranaceous, largely restricted to ends of branches, subsessile to petiolate; blades 2-20 cm long, wide, narrowly elliptic to obovate, the bases acu- minate to obtuse (occasionally auriculate, pan- duriform, or retuse), the apices acuminate to ob- tuse, the margins entire, adaxially glabrate to Jepidare: abaxially silvery-canescent to gray- brown, lanate-tomentulose to lepidote-tomen- 1987] MACLEISH — EREM. tulose; trichomes stellate, the stalk with 1—2 cells; primary venation pinnate and abaxially promi- nent, the midrib furrowed; petioles 1-20 mm long, lanate or lepidote, base slightly expanded. Peduncles slender, to 35 cm tall, terete, reddish- brown to gray, lanate-tomentose to lepidote-to- mentulose, expanded at apex; bracts foliaceous, both intra- and extraglomerular. merules- cence a terminal cyme of 2-20 glomerules or a compound cyme of 8-60(-100) glomerules; mat- uration of glomerules centrifugal. Glomerules 5- 15 mm tall, 4-12 mm diam., hemispherical to spherical; head maturation concurrent to cen- trifugal within a glomerule. Heads homogamous, 1-100 per glomerule, closely to slightly ap- pressed and free, or coherent by pubescence of phyllaries, or connate by concrescence of recep- tacle and phyllary tissues. Involucres obconic or cylindrical, 2-12 mm tall, 0.3-4 mm diam.; phyllaries in 4-7 imbricate, graduated (rarely subequal) series; innermost phyllaries deciduous at maturity: mare ins subscarious, entire to o ten with purple apex, glabrate to lanate-tomentulose REVISION 269 or lepidote-tomentulose, the inner surfaces stra- mineous and vernicose; single vein (when pres- ent) often with mucronate extension. Florets 1— 3(4) per head; corollas purple to white, 5-9 mm tall, the tube and lobes subequal; tube 2-4.5 mm tall; throat 0.5-1 mm tall; lobes 2.5-3.5 mm tall. Anthers 2-4 mm long. Pollen tricolporate with a continuous micropunctate tectum, short echi- nate (Vernonia type A). Style 4-6 mm long, 0. 1 5- 0.3 mm wide, noticeably flattened, the branches short! to mediuer bine l- 2. J mm long. Achenes 1 Lyla J 1-4 mm n tall, glandular, glabrous or sericeous, 10-ribbed (often obscurely or rarely to 20), apex constricted and dark; carpopodium absent or mi- nute; nectary 0.2-0.45 mm tall, persistent or tar- dily deciduous, occasionally pubescent. Pappus 3-5-seriate, of stramineous, white, or purple, strongly coroniform to not coroniform, often ba- sally fused, persistent to promptly deciduous, pa- leaceous to filiform, strigose bristles; outermost series 0.4-3 mm long, innermost series 3-8 mm long. Chromosome number: n = 1 tur binate), KEY TO SPECIES OF EREMANTHUS la. Glomerulescence a compound cyme of 3-60 glomerules or a cyme of 2-20 glomerules; heads (1-)20- E per seen with phyllaries in 3—6 series; florets ericeous with stramineous (rarely white or purple), strongly to subcoroniform, persistent ou cylindri spher 1(2-4) per head; achenes cylindric-turbinate 3a. js om cylindric-turbinate (rarely re 10-ribbed; pappus stramineous (rarely purple), coroniform, persistent, paleaceous; heads onic (rarely cylindric), coherent by interwoven pubescence of phyllaries (subg. Eremanthus = 4a. Glomerulescence a compound cyme of (3-)10—60 glomerules (sect. Eremanthus). 5a. Stem pubescence reddish-brown lanate-to mentose; heads 20-100 per glomerule. re a l4—V» length; leaves S AU Bi Sa rarely E peas -to- lomerulatus 6b. Heads ed entire length; leaves strongly coriaceous, . abaxially esie to- 2. i g mentose . Stem renin gray lepidote- tomentulose; heads 5- -45 per ‘glomerule. E. goyazensis 7a. Heads coherent '^ length; leaf blades 6-16 cm long, 2-10 cm wide uu 7b. Heads coherent Ys length; ‘leaf blade 2-7 cm n long, 0.5-1. 5 cm n wide . .4. E. mattogrossensis “rondoniensis 4b. Glomerulescence a a |cyme (not compound) of 5- -20(-30) glomerules (sect. Synglomerulus. 8a em pubescence gray lepidote-tomentulose; leaf base acute to attenua argenteus 8b. Stem pubescence reddish- brown lanate- tomentose; leaf base variously lobed res acute). 9a. Leaf bases auriculate, the blades 5-8 cm long, appressed- ascending; heads r glomerule, Rai 12 lengt 6. ute, the blades 6-19 long, not appressed-ascending; heads 7. E. 3b. Achenes cylindric, ; papp us off-white to purple, 3 E. auriculatus cinctus subcoroniform, tardily decid- uous, E heads nde connate by concrescence of tissues (subg. /sotrichia) ... 8. E. incanus 270 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 2b. Florets 3-4 per head; heads slightly appressed, free; achenes cylindric; glomerule hemispherical s). (subg. Pseuderemanthu a E E. seidelii phyllaries in 5-7 series; y pene 1-4 adis und cyme of pus than 100 glomerules; heads 1-12 per glomerule, ead; achenes cylindric (rarely cylindric- b pores sei E. elaeagnus with with a purple or white (rarely stramineous pappus; leaves membranaceous to su (rarel ubcoriaceous (subg. Vanlpsmopsis lla. Heads solitary or in pairs slightly appressed basally, cylindric; pappus never twisted (sect. Nec- taridium). 12a. Florets 2-3 per head; involucre with prominent truncate base, phyllaries purple. ad 13a. Florets 3 per he 13b. Florets 2 per he . ll. E. pohlii . 12. E. graciellae Florets 1 per head; 15b. Florets 34 per 16b. n 3-4 per head. a. Heads 2-5 per glomerule |... 17b. Heads 6-9 per glomerule ........... Eremanthus Less. subg. Eremanthus. Ereman- thus subg. Eueremanthus Schultz-Bip., Jah- resber. Pollichia 20-21: 393. 1863 Leaves strongly coriaceous to subcoriaceous. Glomerulescence a compound cyme of 3-60 glomerules or a cyme of 2-20 glomerules. Glom- erules subspherical to hemispherical. Heads (1-) 20-100 per glomerule, obconic; with phyllaries in 3-6 series, closely appressed, coherent by in- terwoven pubescence of phyllaries. Florets | per head. Achenes cylindric-turbinate, 10-ribbed, se- riceous. Pappus stramineous (rarely purple), co- roniform, persistent, paleaceous, strigose bris- tles. Eremanthus Less. sect. Eremanthus d cyme of (3-)10- 60 glomerules. Glomerules 5-10(-15) mm tall, 5-15 mm diam 1 1 (^ mÈ — . Eremanthus glomerulatus Less., Linnaea 4: 317. 1829. Albertinia glomerulata (Less.) DC., Prodr. 5: 82. 1836. TYPE: Brazil: no other data, Sellow s.n. (holotype, P; isotype, TEX) Albertinia rufiseta DC., Prodr. 5: 81. 1836. TYPE: Brazil. a erais: siccis apricis montosis Serro Frio Martius s.n. (holotype, M). a e re DC., Prodr. 5: 81. 1836. Ere- nthus pallidisetus (DC.) Schultz-Bip., Jahres- Bee Pollichia 18-19: 165. 1861. TYPE: Brazil. Mi- So hun campis editis ad Calumbi Praed. Serro artius s.n. (holotype, M). es biis Gardner, London J. Bot. 5: 235. ad l involucre without truncate base, phyllaries brown ...... b 11b. Heads (2-)5-12 per glomerule, cylindric or obconic; pappus often twisted (sect. Fanilosmops 14a. Heads un a Bae and connate by tissue concrescence, 6-12 per glo 15a. Floret r head; heads coherent '^ length r head; heads coherent nearly entire length . 15. 14b. r = iy slightly appressed and coherent by pubescence of phyllaries, 2- 9 per glomerule. l6a. l per head 6. E. brasiliensis l. E polycephalus E. erythropappus 16. E. uniflorus AE ena c | T; E, capitatus "————— ÉOÉ Inr 18. E. arboreus 1846. Eremanthus stellatus (Gardner) Schultz- Bip., Jahresber. Pollichia 18-19: 164. 1861. Er- emanthus stellatus (Gardner) Schultz-Bip. var. pud Schultz-Bip., Mota Pollichia 20- 21: 1863, n das Brazil. "Per- nam dre (now B a Rosa, district of se Rio Preto, Sept. 1839, Canine 2896 (holo- type, BM; isotypes, 2 in F (fragment and photo of W), G, , 2 in NY Eremanthus stellatus (Gardner) Schultz- Bip. var. poh- ultz-Bip., Jahresber. Pollichia rii 394, 1863. TYPE: rs in summitate montiu e . S. Anna, piu d eig W; isotypes, "F (fragment), 2in N Slender to robust tree, 2-10 m tall, to 20 cm diam.; stems reddish-brown lanate-tomentose; branches few to many. Leaves subcoriaceous, sessile to petiolate; petioles to 10 mm long, la- nate-tomentulose; blades 4—14(-20) cm long, 1- 5.5(-7.5) cm wide, elliptic to ovate, the bases acute to obtuse, the apices obtuse (rarely rounded or retuse), margins entire; adaxially tomentulose to glabrate, abaxially gray-brown lanate-tomen- tulose (rarely lepidote-tomentulose). Peduncle slender, to 20 cm tall, ribbed, reddish-brown la- nate-tomentose to gray lepidote-tomentulose. Glomerulescence a compound cyme of 20-60 glomerules. Glomerules 5-15 mm tall, 9215 mm diam., hemispherical. Heads 20-90 per glom- erule, closely appressed basally, fused “4 to !^ length by pubescence of phyllaries. Involucres obconic, 3.5-5.5 mm tall, 1.3-2.5 mm diam.; phyllaries in 4—6 series; outermost phyllaries nar- rowly obtrullate, 1.2-3 mm long, 0.4-1.6 mm wide, the apices acute; innermost phyllaries ob- lanceolate, 3.1-5.2 mm long, 0.6-1.4 mm wide, 1987] 1 200 km 0 100 Elevation:..... 500 meters „1000 MACLEISH — EREMANTHUS REVISION € E. glomerulatus a E. A E. goyazensis mattogrossensis am E. cinctus Figure 2. Distribution of Eremanthus subg. Eremanthus in Minas Gerais. the apices acute; margins entire; abaxial surfaces stramineous with purple apex, glabrate to lanate- tomentulose. Corollas purple to white, 4-7.8 mm tall, the lobes acuminate. Anthers 2-3.4 mm long, the apical appendage acuminate to rounded, the bases obtuse to rounded. Achenes 2.5-4.1 mm tall, glandular, sericeous between the 10 ribs, the apex slightly constricted; nectary 0.15-0.25 mm tall, tardily deciduous. Pappus 4-5-seriate, of strongly coroniform (at maturity) strigose bris- tles; outermost series 1-2.2 mm long; innermost series 4—6.2 mm long. Flowering and fruiting occur from March to October. Eremanthus glomerulatus is distributed throughout Minas Gerais, Goiás, and adjacent regions in Bahia and Sao Paulo (Figs. 2, 3, 6), at altitudes of 700 to 1,500 meters. It is found com- monly in large colonies and may dominate cer- rado and campo rupestre habitats. In Minas Ge- rais, this species often is used for fence posts because of its abundance. Eremanthus glomerulatus is the eastern mem- ber of a group of closely related taxa (sect. Ere- manthus) found throughout the Central Plateau. Eremanthus goyazensis is slightly more robust with a greater fusion of heads and is found in the northcentral branch of the Plateau. In contrast, E. mattogrossensis is less robust and has lepi- dote-tomentose (rather than lanate-tomentose) stems and leaves; it extends throughout the west- ern arm and into the southwestern side. Ere- manthus rondoniensis is closely related to E. mattogrossensis and is restricted to the far west- ern portions of the western arm. Eremanthus glomerulatus appears to be the member of this complex most closely related to the taxa histor- ically considered to compose Vanillosmopsis. The and occasionally purple pappus of E latus are reminiscent of characteristics found in the transitional taxa, E. incanus, E. elaeagnus, and E. seidelii. In addition, it is the only member of this complex that is sympatric with both the transitional taxa and the majority of the Vanil- losmopsis complex. he many epithets ascribed to E. glomerulatus 272 represent variations in pappus color (purple or white) and number of series, branch position (an- gular or not), and leaf shape. Eremanthus glo- merulatus can be found throughout most of the Central Plateau, and the observed variation can be attributed to maturity and habitat differences. These epithets were applied by earlier authors who were attempting to identify Brazilian col- lections in the absence of adequate documenta- tion. Baker (1873) was the first author to rec- ognize E. glomerulatus as a widespread and slightly variable species Paris specimen of E. glomerulatus is clear- ly the holotype. It is annotated appropriately by both Lessing and Schultz-Bipontinus and match- es the description given by Lessing in 1829. Orig- inally both the Paris specimen and the Texas specimen were housed in Berlin because both are stamped “hb. berol." Additional specimens examined. BRAZIL. BAHIA: Rio das Contas, 22 Jul. 1979, King et al. 8099 (US). DISTRITO FEDERAL: Gama, 28 Aug. 1965, Irwin et al. 7926 (MO, NY, US). GorAs: Cristalina, 3 Apr. 1973, Anderson 8004 (F, MO, NY, UB, US). MINAS GERAIS: Nova Lima, 15 Apr. 1945, Williams & Assis 6626 (F, GH, MO, NY, S). Possible hybrids. x E. elaeagnus (C. Martius ex DC.) Schultz-Bip. MINAS GERAIS: Serra do Cipó, : Oct. 1980, ni. et al. 715 , GH MO, NY, S, US), 7 Oct. 1980, et al. 716 (GA, Ra. 30 Jan. 1980, King & Almeda 8354 (US). x E. mattogrossensis Kuntze. MINAS GERAIS: Sac- ramento, 22 Jul. 1980, Schumacher B10 Sea x E. seidelii MacLeish & Schumacher. MINAS GE tween Furnas and Piui, 25 Aug. 1981, Schumacher 1008 (GA). 2. Eremanthus goyazensis (Gardner) Schultz- Bip., Jahresber. Pollichia 18-19: 165. 1861. Albertinia goyazensis Gardner, London J. Bot. 6: 425. 1847. TYPE: Brazil. Goias: hilly campos near Villa de Arrayas, Apr. 1840, Gardner 3804 (holotype, BM; isotypes, 2 in F (fragment and photo of B), 3 in G, GH (photo of B), 3 in NY, TEX (photo of B), W) Eremanthus weddellii Schultz- is Jahresber. ee lichia 18-19: 165. 1861. TYPE: Brazil, Goiás, e in Salinas, May 1844, Weddell 2032 Fiori P). Small tree, 1—3 m tall, the trunk often gnarled; stems reddish-brown to gray-black lanate-to- mentose, to 8 cm diam.; branches few. Leaves strongly coriaceous, sessile to petiolate; petioles to 10 mm long, lanate-tomentulose; blades 7.5— ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 18 cm long, 3.5-10 cm wide, elliptic to ovate, the bases obtuse, the apices obtuse to rounded (rarely retuse), me margins entire; adaxially sparsely lepidote- tomentulose. Peduncle nde. to 20 cm tall, ribbed, gray-brown lepidote-tomentulose. Glomerulescence a compound cyme of 10—40 glomerules. Glomerules 5-15 mm tall, 7215 mm diam., hemispherical. Heads 25-100 per glom- erule, ee appressed, pee entire length y lanose pubescence o yllaries. Involucres one 4.6 mm tall, 0. 4 1.5 mm diam.; phyl- laries in 3—4 series; outermost phyllaries narrow- ly obtrullate, 1.8-3.2 mm long, 0.2-0.8 mm wide, the apices acute to acuminate; innermost phyl- laries oblanceolate, 2.9—4.8 mm long, 0.3-0.8 mm wide, the apices acute; margins entire; abaxial surfaces stramineous, glandular, glabrate to la- nose-tomentose. Corollas pale purple to white basally, 5.2-8.2 mm tall, the lobes acuminate. Anthers 2.4—4.4 mm long, the apical appendage acuminate; bases obtuse to rounded. Achenes 2- 2.8(-3.8) mm tall, glandular, sericeous between 10 ribs, often obscurely ribbed; apex slightly con- stricted; nectary 0.25-0.35 mm tall, tomentose, tardily deciduous. Pappus 4-5-seriate, of strong- ly coroniform (at maturity) strigose bristles; out- ermost series 0.8-2.3 mm long, the innermost series 4.6-6 mm long. Chromosome number: n = 15. Flowering and fruiting occur from January to September. ! ^d si ? Hd E LP e. | d through- out southern Goiás and nearby Minas Gerais (Figs. 2, 3). It is infrequent in cerrado and campo rupestre at altitudes of 700 to 1,600 meters. Eremanthus goyazensis is the northcentral ex- tension of the glomerulatus complex. It is dis- tinguished from E. glomerulatus by its strongly coriaceous leaves with abaxial lepidote-tomen- tose surfaces and heads entirely coherent by in- terwoven pubescence. In addition, it can be easily distinguished from the mattogrossensis-rondon- iensis complex by its lanate stem pubescence, 5-100 heads per glomerule, and high degree of cohesion. Additional specimens examined. BRAZIL. DISTRITO FEDERAL: Chapada da Contagem, 18 Aug. 1964, Irwin & Soderstrom 5287 (NY, TEX, UB). Goiás: Terezina, 18 Mar. 1973, Anderson 7354 (F, MO, NY, UB, US). 3. Eremanthus mattogrossensis Kuntze, Revis. Gen. PI. 3(2): 145. 1898. TYPE: Brazil: “Mat- 1987] 4 5, . argenteus . auriculatus . goyazensis E E E. glomerulatus E E. mattogrossensis E pares . cinctus Elevation: m IT 500 meters 21000 ” a ome LN 2 FIGURE 3. togrosso" (now Mato Grosso): Jul. 1892, Kuntze s.n. (holotype, NY; isotypes, photos of B, F, GH, TEX). Shrub or slender tree, 1-4 m tall; stems gray- brown lepidote-tomentulose; branches few. Leaves coriaceous, sessile to petiolate; petioles to 15 mm long, lepidote; blades 6-16 cm long, 2-10 cm wide, elliptic to page 4 pu the bases acute to acuminate, the ap to rounded, the margins entire; adaxially sparsely lepidote to glabrate, abaxially gray lepidote-to- mentulose. Peduncle slender, to cm tall, ribbed, gray-brown lepidote-tomentulose. Glomerulescence a compound cyme of 6-35 (-80) glomerules. Glomerules 4.5-10 mm tall, 5.5-12 mm diam., subspherical. Heads 10—45 per glomerule, closely appressed basally, coher- ent '2 length by pubescence of phyllaries. Invo- lucres cylindric, 2.3-4 mm tall, 1.1-2.4 mm diam.; phyllaries in 5—6 series; outermost phyl- laries narrowly obtrullate, 1.3-2.3 mm long, 0.2- 0.6 mm wide, the apices acute; innermost phyl- laries oblanceolate, 1.8—3.4 mm long, 0.5—0.9 mm wide, the apices acute; margins entire; abaxial surfaces stramineous, glabrate to lepidote-to- mentulose. Corollas pale purple to white, 4.4— .6 mm tall, the lobes acuminate. Anthers 2.1- 3.4 mm long, the apical appendage acuminate to acute, the bases obtuse to rounded. Achenes 2- 3 mm tall, glandular, sericeous between ribs, 10- MACLEISH — EREMANTHUS REVISION 273 I wy E rida T eo ELS o NEFKEE s OUS i J MS ` FI + phe Lu - / SS d. LO USER eee aes pcr ae 200 kilometers Distribution of Eremanthus subg. Eremanthus in Goias. ribbed (often obscurely), the apex slightly con- stricted; nectary 0.15-0.25 mm tall, tomentose, tardily deciduous. Pappus 3-5-seriate, of strong- ly coroniform (at maturity) strigose bristles; out- ermost series 0.8-3.1 mm long; innermost series 3.3-6 mm long. Flowering and fruiting occur from April to Au- gust. Eremanthus mattogrossensis is widely distrib- uted throughout the western section of the Cen- tral Plateau of Brazil (Figs. 2-4) at elevations of 500 to 1,000 meters in cerrado. It is particularly common in Mato Grosso, where it is called **vel- ludo do cerrado," velvet ofthe cerrado, or **casca freta," cleft bark. Eremanthus mattogrossensis is transitional between b. goyazensi and E. rondoniensis and l location. All three are found within the western arm of the Central Plateau. Eremanthus rondoniensis is restricted to the far western portion of the Serra dos Parecis near the border of Mato Grosso and Rondônia, and E. goyazensis is restricted to eastern portions of the western arm (principally southern Goiás). The type specimen, bearing only the citation “Mattogrosso,” must have been collected in western Mato Grosso, near Rondônia. Zanoni (1980), in his excellent survey of Kuntze’s col- lecting trips, pointed out that Kuntze entered Brazil through Bolivia and explored only a small / mattogrossensis / ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 m E. 57° ` f N æ E. cinctus / -—- 6? ) [4 z3 ` r4 9 E. rondoniensis > e "d \ / [| ) |] z `Y / / “semasa REND / ERE: seed! A. Melasi re Elevation: . 500 meters :. 1000 i FIGURE 4. Distribution of Eremanthus subg. Eremanthus in western Central Plateau. area of Mato Grosso near the border. Zanoni also noted that the bulk of Kuntze's personal her- barium, including many type specimens, is at the w York Botanical Garden. Thus, their spec- imen of E. mattogrossensis, which bears the ap- propriate collecting information in Kuntze's handwriting and which matches his type descrip- tion, is likely the holotype. Additional specimens examined. BRAZIL. DISTRITO FEDERAL: Brasília, 15 Jun. 1981, Heringer et al. 7057 (MO). GorAs: 75 km from Aragarças, near Piranhas, 23 Jun. 1966, Hunt & Ramos 6160 (MO, NY, UB). MATO GROSSO: Barra do Garças, 6 May 1973, Anderson 9801 (G, MO, NY, UB). MINAS GERAIS: Furnas, 5 Jul. 1982, Schumacher 2008, 2009, 2010, 2012, 2014 (GA, je n Sao Felix do Xingu, 12 Jul. 1978, Rosario Y). SAO PAULO: inter Canna Verde et S. José, mdi at 670 (BR, S). Possible hybrids. x E. glomerulatus Less. MINAS GERAIS: Sacramento, 22 Jul. 1980, Schumacher B10 (GA). x E. seidelii MacLeish & Schumacher. MINAS GERAIS: between Furnas and Piuí, 25 Puig 1981, Schu- macher 1003, 1004 (GA). 4. Eremanthus rondoniensis MacLeish & Schu- macher, Syst. Bot. 9: 89. 1984. TYPE: Brazil. Rondônia: Vilhena, 13?16'S, 58*52'W, 18 Apr. 1977, Bantel & Silva s.n. (holotype, RB). Small tree, ca. 1 m tall; stems gray-brown lep- idote-tomentulose, often basally darkened by fire. Leaves coriaceous, sessile to petiolate; petioles to 5 mm long, lepidote; blades 2-7 cm long, 0.5- ces obtuse entire; adaxially deca lepidote to ioc abaxially gray-brown epidote-tomentulose. Peduncle slender, height unknown, terete, gray-brown lepidote-tomen- tulose. Glomerulescence a compound cyme of 3-20 glomerules. Glomerules 5-7 mm tall, 7-10 mm diam., hemispherical. Heads 5-15 per glom- erule, closely appressed, coherent '4 length by pubescence of phyllaries. Involucres obconic, 3.5- 5 mm tall, 1-2.2 mm diam.; phyllaries in 4—6 imbricate graduated series; outermost phyllaries widely trullate, 1.4-2.1 mm long, 0.5-0.7 mm wide, the apices obtuse; innermost phyllaries ob- lanceolate, 2.4-4 mm long, 0.8-1 mm wide, the apices acute; margins entire; abaxial surfaces stramineous with purple apex, glabrate to lepi- dote-tomentulose. Corollas white, 5.1—6 mm tall, the lobes acuminate. Anthers 2.6-3 mm long, the apical appendage acuminate, the bases rounded. Achenes turbinate, 1.9-2.2 mm tall, glandular, obscurely-ribbed, the base sericeous, the apex constricted with prominent dark collar; 1987] nectary 0.2-0.5 mm tall. Pappus 3-5-seriate, of strongly coroniform (at maturity) strigose bris- tles; outermost series 0.6-1.8 mm long; inner- most series 5—6.8 mm long. Flowering and fruiting occur from April to July. Eremanthus rondoniensis is distributed along the Rondónia-Mato Grosso border in the west- ern portion of the Serra dos Parecis (Fig. 4). It is found in periodically burned cerrado. Ere- manthus rondoniensis is closely related to E. mattogrossensis and represents a western exten- sion of the g/omerulatus-goyazensis- mattogros- sensis complex. Additional specimens examin ned. BRAZIL. MATO te Rondônia): RONDÔNIA (latitude and longitude indicate Mato Gros- so): Vilhena, 13?46'S, 59°50'W, 17 Apr. 1977, Bantel & Silva s.n. (RB) Eremanthus sect. Synglomerulus MacLeish, sect. nov. TYPE: Eremanthus argenteus MacLeish & Schumacher. Glomerulescentiae cymosae. Glomerulis 5-10(-30); (5-)10-25 mm longis, 10-25 mm diam WG . Eremanthus argenteus MacLeish & Schu- macher, Syst. Bot. 9: 84. 1984. TYPE: Brazil. Goiás: 33 km N of Alto Paráiso towards Cavalcante, 1,370 m, 14 . 1980, MacLeish, Martinelli, Smith & Stutts 734 (holotype, RB; isotypes, F, G, GA, GH, MO, NY, UB, US). Slender tree, 2-3 m tall; stems ash gray lepi- dote-tomentulose; branches few. Leaves coria- ceous, sessile to petiolate; petioles to 5 mm long, lepidote-tomentulose; blades 7.5-14(-18) cm long, 2.5-6 cm wide, elliptic, the bases acute to attenuate, the apices rounded to obtuse, the mar- gins entire; adaxially silvery lepidote; abaxially silvery lepidote-tomentulose. Peduncle slender, to 30 cm tall, terete, ash gray lepidote-tomen- tulose. Glomerulescence a cyme of 6-20 glom- erules. Glomerules 5-20 mm tall, 10-25 mm diam., subspherical. Heads 10—80 per glomerule, closely appressed, coherent for ?^ their lengths by pubescence of phyllaries. Involucres obconic, 3.5-5 mm tall, 1-2.5 mm diam.; phyllaries in 4— 5 series; outermost phyllaries narrowly obtrul- late, 1.7-2.9 mm long, 0.4 m wide, the apices acute; innermost phyllaries narrowly ob- trullate, 3.6-5.1 mm long, 0.5-1.1 mm wide, the MACLEISH — EREMANTHUS REVISION 245 apices acuminate, margins entire to subfim- briate; abaxial surfaces stramineous with purple apex, glandular, glabrate to tomentose. Corollas purple, 5.2-7.4 mm tall, the lobes acuminate. Anthers 3-3.6 mm long, the apical appendage acuminate to acute, the bases acute to obtuse. Achenes 2.5—4 mm tall, glandular, sericeous, ob- scurely ribbed, the apex slightly constricted; nec- tary 0.2-0.5 mm tall. Pappus 3-5-seriate, of co- roniform (at maturity; strigose bristles; outermost series 1-2.5 mm long; innermost series 5-7 mm long. Flowering and fruiting occur from July to Oc- tober Eremanthus argenteus is found only in the hapada dos Veadeiros of southcentral Goiás (Fig. 3). Although common, it is restricted to the granite outcrops of campo rupestre at high alti- tudes. This species resembles E. cinctus and E. au- riculatus which also show glomerulescence re- duction to a cyme of glomerules from the com- pound cyme, which is more typical of sect. Eremanthus. Eremanthus argenteus is easily separated from both species by its silvery lepi- dote surfaces and absence of auriculate or retuse leaf bases and apiculate phyllaries. At present, it does not appear to be related to any particular species of Eremanthus. Additional specimens examined. BRAZIL. GOIÁS: Brasilia Richtung Campos Belos, 22 km nach Alto Pa- raiso, 1,500 m, 28 Aug. 1981, Schumacher 1035 (GA, M, MB, RB, W) Possible hybrids. x E. uniflorus MacLeish & Schu- macher. GorAs: Alto Paraiso, 28 Aug. 1981, Schu- macher 1038, 1039, 1041 (GA). 6. Eremanthus auriculatus MacLeish & Schu- macher, Syst. Bot. 9: 86. 1984. TYPE: Brazil. Goiás: Brasilia Richtung Campos Belos, 22 km nach Alto Paráiso, 1,500 m, 28 Aug. 1981, Schumacher 1037 (holotype, RB; isotypes, GA, MB). Shrub to small tree, 0.8—1.5 m tall; stems red- Leaves coriaceous, cending-appressed; blades 5-8 cm wide, trullate to ovate, the bases iia the apices acute, the margins entire; adaxially to- mentulose, abaxially gray-brown lanate-tomen- tose. Peduncle slender, to 20 cm tall, ribbed, red- dish-brown lanate-tomentose. Glomerulescence 276 a cyme (rarely compound) of 9-20 glomerules. Glomerules 5-15 mm tall, 15-20 mm diam., subspherical. Heads 30-80 per glomerule, closely appressed basally, coherent !^ length by lanate pubescence of phyllaries. Involucres obconic, 4— 5.5 mm tall, 1.5-2.5 mm diam.; phyllaries in 4— 5 series; outermost phyllaries narrowly obtrul- late, 2.3-4 mm long, 0.4-0 wide, the api- ces acute; innermost phyllaries narrowly obtrul- late, 4.2-4.6 mm long, 0.5-0.8 mm wide, the apices apiculate; margins entire to subfimbriate; abaxial surfaces stramineous with purple apex, often axially green, tomentulose to lanate. Co- rollas purple, 5.5-7 mm tall, the lobes acuminate to acute, sparsely glandular. Anthers 3.3-3.6 sericeous, obscurely-ribbed, apex slightly con- stricted; nectary 0.2-0.5 mm tall. Pappus 3-5- seriate, of coroniform (at maturity) strigose bristles; outermost series 1.2-2.2 mm long; in- nermost series 5.5-7 mm long. Flowering and fruiting occur from July to Sep- tember Eremanthus auriculatus is known only from the Chapada dos Veadeiros in southcentral Goiás (Fig. 3). It occurs on campo rupestre adjacent to the cerrado characteristic of this region. This species superficially resembles E. cinctus; however, E. auriculatus is distinguished by as- represents a further reduction in glomerulescence internodes from a compound cyme to form a cyme of glomerules. It is easily separated from the glomerulatus-complex by the cyme of glom- erules (vs. compound cyme), by auriculate leaf bases (vs. acute to obtuse), and by apiculate phyl- laries (vs. acute to acuminate). Additional aeg examined. BRAZIL. GOIÁS: Alto Paráiso, 4 Jul. 1983, Schumacher 3045 (GA, MB). 7. Eremanthus cinctus Baker in C. Martius, Fl. Bras. 6(2): 162. 1873. TYPE: Brazil. Mato Grosso: *Cuyaba" (now Cuiabá), 1834, Sil- va Manso 55 (holotype, BR). Eremanthus ated piae Baker in C. Martius, Fl. Bras. y! 1873. TYPE: Brazil, no other data, Tam- berlik s.n. ADS. W; photos of W, F, TEX). Shrub or small tree, 0.5—4 m tall; stems red- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 dish-brown lanate-tomentose; branches few. Leaves subcoriaceous, sessile; blades 6-19 cm long, 2-6 cm wide, elliptic, narrowly elliptic, or obovate, the bases retuse to acute, the apices acute to obtuse, the margins entire; adaxially gla- brate, abaxially tomentulose. Peduncle robust, to 15 cm tall, ribbed, reddish-brown lanate-tomen- tose; 3-9 bracts with prominently enlarged bases surrounding each glomerule. Glomerulescence a cyme (rarely compound) of 5-30 glomerules. Glomerules 10-25 mm tall, 10-25 mm diam., hemispherical. Heads 20-50 per glomerule, closely appressed, coherent entire length by la- nate pubescence of phyllaries. Involucres obcon- ic, 3.5-5 mm tall, 0.7-2 mm diam.; phyllaries in 4—5 subequal series; outermost phyllaries oblan- ceolate, 3.3-4.3 mm long, 0.5—0.7 mm wide, api- ces obtuse; innermost phyllari ,3- 4.8 mm long, 0.7-1 mm wide, apices acute; mar- gins entire; abaxial surfaces stramineous with acuminate and glandular. Anthers 4—4.4 mm long, apical appendage acuminate, bases acute. Achenes 2.5-3.5 mm tall, glandular, sericeous between 10 ribs, apex slightly constricted and dark; nectary 0.1-0.2 mm tall. Pappus 3-4-se- riate, of stramineous (occasionally rusty on dried S mens), coroniform b maturity) strigose de outermost series 1.5-2.5 mm long; in- nermost series 5.5-7.5 mm long. Flowering and fruiting occur from April to September. Eremanthus cinctus is known only from three locations: western Minas Gerais, western Goias, and near Cuiabá in central Mato Grosso (Figs. 2-4). Schumacher was able to locate this species in southwestern Goiás, but reported (pers. comm.) that the Minas Gerais locations cited by Good- land on his 1967 collections have been com- pletely logged for charcoal. Baker (1873) separated E. pandurifolius from E. cinctus on the basis of a solitary glomerule versus a cyme of glomerules. This distinction is artifactual — the type specimen of E. pandurifo- lius is fragmented, making the glomerules appear solitary. Eremanthus pandurifolius appears to have been collected in western Minas Gerais, because that is the region Tamberlik is known to have explored (Urban, 1906). Likewise, Silva Manso was known to have been practicing med- icine in Cuiaba during the year recorded on the label ofthe type specimen of E. cinctus. Although 1987] MACLEISH — EREM. E. cinctus and E. pandurifolius were published simultaneously, E. cinctus has been chosen be- cause the type specimen is intact, whereas that of E. pandurifolius is badly fragmented. Until recently E. cinctus appeared to have a disjunct distribution since there were no collections from intervening mountainous regions, particularly Serra do Verdinho, Serra das Araras, and Serra da Sào Jeronimo. Eremanthus cinctus superficially resembles E. auriculatus but is distinguished by subcoriaceous leaves with acute to retuse bases, several prom- inent extraglomerular bracts, and total fusion of heads. By contrast, E. auriculatus has coriaceous leaves with prominent auriculate bases, and heads appressed only at the bases. Eremanthus cinctus, like E. auriculatus, is clearly related to E. glo- merulatus, and represents a further reduction in glomerulescence internodes from a compound cyme to form a cyme of glomerules. Additional pe in examined. BRAZIL. MATO GROSSO: known onl GERAIS: Goodland 3766 (UB). Prata, 29 Sep. 1967, Goodland 1 (MO). Go1Ás: Montividiu towards Amorinopolis, 21 km after Montividiu, 800 m, 28 Apr. 1984, Schu- II 4004 (GA, MB, UNICAMP). subg. Isotrichia (DC.) acLeis . nov. Albertinia Sprengel sect. gn ia DC., Prodr. 5: 82. 1836. Vanillosmopsis Schultz-Bip. subg. [sotrichia (DC.) Schultz-Bip., Jahresber. Pollichia 18- 19: 168. 1861. LECTOTYPE, here designated: E. incanus (Less.) Less. Eremanthus Less. M Leaves coriaceous. Glomerulescence a com- ound cyme of 8-50 glomerules. Glomerules Su pherical Heads per glomerule 30-100, cy- lindric, connate by concrescence of tissues, with phyllaries in 4-6 series. Florets 1 per head. Achenes abe 1 5-20-ribbed, sericeous. Pap- pus off-white to purple, subcoroniform, tardily deciduous, subpaleaceous. 8. Eremanthus incanus (Less.) Less., Linnaea 6: 682. 1831. Albertinia incanus Less., Linnaea 4: 342. 1829. Cacalia incana (Less.) Kuntze, Revis. Gen. Pl. 2: 970. 1891. LECTOTYPE, here designated; Brazil, no other data, Sel- low s.n. (B; isolectotypes, BR, F (fragment from P)) Slender to robust tree, 2-10 m tall, to 15 cm diam.; stems gray-brown lepidote; branches few. Leaves coriaceous, petiolate; petioles 4-17 mm REVISION 277 long, lepidote; blades 5.5214 cm long, 2-6 cm wide, elliptic to ovate, the bases acute, the apices rounded to obtuse, the margins entire; adaxially glabrate, abaxially gray lepidote. Peduncle slen- der, to 27 cm tall, terete, gray lepidote. Glomer- ulescence a compound cyme of 8-50 glomerules. Glomerules 5-15 mm tall, 7-15 mm diam., spherical. Heads 30-100 per glomerule, closely crescence of p dric, 2.4-5 mm tall, 0.3-1.5 m ; phy ries in 4—5 series; outermost phyllaries irianaulét: 1.7-3 mm long, 0.1-0.5 mm wide, the apices acute to obtuse; innermost phyllaries narrowly obtrullate, 2.6-4 mm long, 0.3-0.6 mm wide, the apices acuminate, the margins entire; abaxial surfaces stramineous with purple apex, glabrate to lepidote-tomentulose. Corollas pale purple to white, 4.2—6.6 mm tall, the lobes acuminate. An- thers 2.2-2.7 mm long, the apical appendage acu- minate, the bases acute. Achenes cylindric, 2.2- 2.6 mm tall, glandular, sparsely sericeous, 15- 20-ribbed, the apex slightly constricted; nectary 0.15-0.25 mm tall. Pappus 3—4-seriate, off-white urple, subcoroniform (at maturity), tardily Pied Soiree naqa strigose bristles; out- ermost series 1-1.5 mm long; innermost series 5-5.8 mm ee 2 Flowering and fruiting occur from July to Oc- tober Eremanthus incanus is distributed throughout the southern section and northeastern arm (Figs. 5, 6) of the Central Plateau of Brazil at elevations of 800 to 1,850 meters in cerrado, secondary forest, or caatinga. It is particularly common in Minas Gerais where it is called “pão candeia" Scone and or “‘candeia da serra" (mountain dle), names Mer d attributed to subg. Va- UNE Eremanthus incanus is transitional between gle floret per head, large sericeous achenes, an subpaleaceous pappus. In contrast, in its lepidote tomentum, relatively small glomerules, phyllary tissue fusion, cylindrical achenes, and off-white to purp ] E. pine net of the Vanillosmopsis complex. Lessing’s specimens were deposited at Char- kow. Because Lessing’s original material was un- available, a Sellow collection from B herbarium that bears appropriate annotations by Lessing 4 ED ly e, tardi itapproaches — Elevation jee vane, Oy meters . 1000 T T T 0 100 ( FIGURE 5. was chosen as the lectotype. Historically, the name Albertinia bicolor (now Paralychnophora bicolor) has been confused often with E. incanus due to the prominent spherical glomerules in both taxa. Paralychnophora bicolor is characterized by solitary, relatively large, axillary glomerules, 2- 4 or 8-12 florets per head, and angled, glabrous achenes with a biseriate pappus. Additional specimens examined. BRAZIL. BAHIA: n. de das Contas, Pico das Almas at 18 km e Rio das Contas, 1,600-1,850 m, 24 Jul. 1979 King et al. 8136 (CEPEC, U INAS GERAIS between Paràopeba and Diamantina, 3 Oct. 1980 MacLeish et al. 687 (C, G, , US), between Mendanh a and Diamantina, 4 Oct. 1980, MacLeish et al. 703 (BR, CEPEC, F, GA, M, MO, RB, TEX). Possible hybrids. x E. elaeagnus (C. Martius ex DC.) Schultz-Bip. MINAS GERAIS: between Paraopeba and Diamantina, 3 Oct. 1980, MacLeish et al. 688 (F, G, GA, GH i (DC ) MacLeish. MINAS Barreto 47 x E. u MINAS GERAIS: Itamarandiba, 6 Sep 1116a (GA, MB). .) MacLeish. . 1981, Schumacher Eremanthus Less. subg. Pseuderemanthus Schultz-Bip., Jahresber. Pollichia 20-21: ANNALS OF THE MISSOURI BOTANICAL GARDEN deis: Sere [Vor. 74 n mealet Distribution of Eremanthus subg. Isotrichia and subg. Pseuderemanthus in Minas Gerais. 395. 1863. Eremanthus Less. subg. Pseu- deremanthus Schultz-Bip. “A. Elaeagnus” Schultz-Bip., Jahresber. Pollichia 20-21: 395. 1863. LECTOTYPE: here designated; Er- emanthus elaeagnus (C. Martius ex DC.) Schultz-Bip. Leaves coriaceous. Glomerulescence a com- ispheric g lomerules. rets 3-4 per head. Achenes cylindric, "10- ribbed, sericeous. Pappus stramineous or purple, coro- niform, persistent, subpaleaceous. 9. Eremanthus seidelii MacLeish & Schumach- er, Syst. Bot. 9: 89. 1984. TYPE: Brazil. Minas Gerais: Furnas in Richtung Piui, kurz von Staumauer, 800 m, 25 Aug. 1981, Schu- macher 1006 (holotype, RB; isotypes, 2 in GA, K, M, MB Tree, to 4 m tall; stems gray lepidote-tomen- tulose; branches many. Leaves coriaceous, sessile to petiolate; petioles to 10 mm long, densely lep- idote-tomentose; blades 4-9 cm long, 1.2-3 cm wide, elliptic, the bases acute, the apices acu- 1987] ~ E. arboreus _ . brasiliensis c< 44 \ E. capitatus e Ó © > E. glomerulatus @ E. graciellae * E. incanus O E. pohlii x E. uniflorus rr Elevation: 100 200 km FIGURE 6. oiás minate to acute, the margins entire; adaxially sparsely lepidote, abaxially densely gray lepi- dote-tomentose. Peduncle slender, to 15 cm tall, terete, gray lepidote-tomentulose. Glomerules- cence a compound cyme of 10-60 glomerules. Glomerules 5-10 mm tall, 6-15 mm diam., hemispherical. Heads 1-7 per glomerule, closely ai Bis basally, free. Involucres cylindric, 5.5- 7.2 mm tall, 2.5-4 mm diam.; phyllaries in 4—6 scies: outermost phyllaries trullate, 1.3-4 mm long, 1-1.5 mm wide, the apices acute; innermost phyllaries narrowly rhombic to obtrullate, 4.7— 7 mm long, 0.5-1.2 mm wide, the apices acute; margins entire to subfimbriate; abaxial surfaces stramineous with purple apex, lepidote-tomen- tulose. Florets 3 per head; corollas purple to white, 5.4—6.5 mm tall, the lobes acuminate. Anthers 2.5-3 mm long, the apical appendage acuminate, the bases acuminate. Achenes 2.2-3.1 mm tall, glandular, sparsely sericeous between 10 ribs, apex slightly constricted; nectary 0.1—0.2 mm tall. Pappus 3-4-seriate, of stramineous, subcoroni- form (at maturity) strigose bristles; outermost MACLEISH — EREMANTHUS REVISION 279 LR Nis "f z hus eer s o A i A Ó ` Y $$; Distribution of Eremanthus subg. Vanillosmopsis, E. glomerulatus, and E. incanus in Bahia and series 0.8-1.2 mm long; innermost series 4.5—6.2 mm long. Flowering and fruiting occur from June to Au- gust. Eremanthus seidelii is restricted to the cerrado surrounding the Furnas reservoir in southwest- ern Minas Gerais (Fig. 5). This species is closely related to E. elaeagnus, which is restricted to the Serra da Espinhaco of northeastern Minas Ge- rais. However, E. seidelii is distinguished from E. elaeagnus by having fewer heads per glomer- ule (1-7 vs. 9-20), straw vs. mostly purple pap- pus color, elliptic vs. narrowly elliptic leaves, and flowering June-July vs. August-September Eremanthus seidelii and E. elaeagnus are tran- sitional between subg. Eremanthus and subg. Vanillosmopsis. They approach E. glomerulatus in their coriaceous leaves, only 10-60 glomerules per glomerulescence, large sericeous achenes, and persistent subpaleaceous pappus. In contrast, in their lepidote tomentum, 1-20 heads per glom- erule, relative lack of head fusion, 3-4 florets per 280 head, and occasionally purple pappus they ap- proach many members of subg. Vanillosmopsis. Additional specimens examined. BRA MINAS GERAIS: vor Alpinopolis, 12 Jun. 1977, Seidel s n. (RB); bei Capitolio, 800 m, 25 Aug. 1981, Schumacher n (GA, MB); Furnas in Richtung Capitolio, 18 Oct. 1980, Sc humacher s.n. (GA, MB). Possible hybrids. x E. erythropappus (DC.) MacLeish. MINAS GERAIS: Furnas towards Piui, 25 Aug. 1981, Schumacher 1005 (GA); Alpinopolis, 12 Sep. bel Seidel s.n. (RB). x E. glomerulatus Less. MINAS AIS: between Furnas and Piui, 25 Aug. 1981, Schu- mac che 1008 (GA). x E. mattogrossensis Kuntze. MI- NAS GERAIS: Furnas towards Piuí, 25 Aug. 1981, Schu- Mule 1003, 1004 (GA). 10. Eremanthus elaeagnus (C. Martius ex DC.) Schultz-Bip., Jahresber. Pollichia 20-21: 395. 1863. Albertinia elaeagnus C. Martius ex DC., Prodr. 5: 81. 1836. Vernonia elaeag- nus (C. Martius ex DC.) Schultz-Bip., Jah- resber. Pollichia 18-19: 166. 1861. TYPE Brazil, Minas Gerais, altis lapidosis Serro Frio prope Tejuco (now Diamantina), Mar- tius s.n. (holotype, M). Small tree or shrub, 2-5 m tall; stems gray lepidote-tomentulose; branches many. Leaves coriaceous, sessile to petiolate; petioles to 10 mm long, densely lepidote-tomentose; blades 5-12 cm long, 1.5-3 cm wide, narrowly elliptic, the bases acute to acuminate, the apices acute to rounded, the margins entire; adaxially sparsely lepidote, abaxially densely gray lepidote-tomentose. Pe- duncle slender, to 14 cm tall, terete, gray lepi- dote-tomentulose. Glomerulescence a com- pound cyme of 10-60 glomerules, these 5-10 mm tall, 10-15 mm diam., hemispherical. Heads 9-20 per glomerule, closely appressed basally, free. Involucres cylindric, 5-10 mm tall, 2.5-4 mm diam.; phyllaries in 4—5 series; outermost phyllaries trullate, 1.5-3 mm long, 0.6-1.2 mm wide, the apices acute; innermost phyllaries nar- rowly rhombic to obtrullate, 3.5-6.5 mm long, 0.7-1. ak = wide, fue pieces sente: mangas en- tiret with purple apex, lepidote-tomentulose. pa 3-4 per head; corollas purple to white, 5.5-7 m tali, the lobes acuminate, AE is 5- 3 mm s ate,the bas- es acuminate. Achenes 2.5-3.5 mm tall, glan- dular, sparsely sericeous between 10 ribs, the apex slightly constricted; nectary 0.1-0.15 mm tall. Pappus 3-4-seriate, of purple (rarely stramin- eous), subcoroniform (at maturity), strigose bris- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 tles; outermost series 1-1.5 mm long; innermost series 6-6.5 mm long. Flowering and fruiting occur from August to September. Eremanthus elaeagnus is restricted to the Ser- ra do Espinhaco in northeastern Minas Gerais at elevations of 900 to 1,370 meters (Fig. 5). It is locally common and dominant along with mem- bers of Lychnophora in campo rupestre. Because ofthe gray tomentum covering most of this plant's surface and its general resemblance to members ofthe Vanillosmopsis complex, it is locally known as "candeia branca,” or white candle. This taxon is closely related to E. seidelii and, as noted under that species, both represent intermediates be- tween subgenera Eremanthus and Vanillosmop- Additional Edda examined. BRAZIL. MINAS GERAIS: between Paraopeba and Diamantina. 3 Oct. 1980, MacLeish et al. 689 (F, GA, GH, MO, NY, RB); Possible hybrids. x E. glomerulatus Less. MINAS GERAIS: Serra do Cipó, 7 Oct. rien icr et al. 715 (BR, F, GA, GH, K M MO, NY, RB, S, US), MacLeish et al. 716 (GA, RB), 30 m 1980, King & Almeda seis S). x E. incanus (Less.) Less. MINAS GERAIS: between Paraopeba and Diamantina 3 Oct ee MacLeish et al. 6 , GA MO, NY, US). x E. GERA (DC.) Ma yis MINAS GERAIS: Serra do Cipó. 7 Oct. 1980, MacLeish et al. 718 (GA). Eremanthus Less. subg. Vanillosmopsis (Schultz- Bip.) MacLeish, comb. nov. Vanillosmopsis Schultz-Bip., Jahresber. Pollichia pa 19: 166. 1861. Vanillosmopsis subg. Euvanil- losmopsis Schultz-Bip., Jahresber. Pollichia 20-21: 398. 1863. TYPE: Vanillosmopsis glomerata Schultz-Bip. = Eremanthus erythropappus (DC.) MacLeish. eaves membranaceous to subcoriaceous. Glomerulescence a compound cyme of 100 or more hemispheric glomerules. Heads per glom- erule 1-12, cylindric or obconic, solitary or slightly to closely coherent by interwoven pu- bescence of phyllaries or connate by tissue con- crescence, phyllaries in 5-7 series. Florets per head 1-4. Achenes cylindric (rarely cylindric- turbinate), 10-ribbed, glabrate. Pappus purple or white (rarely stramineous), not coroniform, promptly (rarely tardily) deciduous, filiform. 1987] gii suem Less. sect. Nectaridium (Schultz- MacLeish, comb. nov. Vanillosmopsis Ede: Bip. subg. Nectaridium Schultz-Bip., Jahresber. Pollichia 20-21: 400. 1863. TYPE: Vanillosmopsis brasiliensis (Gardner) Schultz-Bip. = Eremanthus brasiliensis (Gardner) MacLeish. Heads cylindric, solitary or in pairs slightly appressed basally. Pappus never twisted. 11. Eremanthus pohlii (Baker in C. Martius) MacLeish, comb. nov. Vanillosmopsis poh- lii Baker in C. Martius, Fl. Bras. 6(2): 18. 1873. TYPE: Brazil: foz do Viera, Pohl 556 (holotype, K, not seen; isotypes, 2 in F (one a photo of B), 2 in GH (one a photo of B), NY) Small tree, to 3 m tall; stems gray-brown lep- idote; branches many. Leaves subcoriaceous, petiolate; petioles 7-10 mm long, lepidote; blades 6.4—7.6 cm long, 1.8-2.4 cm wide, lanceolate, the bases attenuate, the apices obtuse, the mar- gins entire; adaxially glabrate, abaxially gray lep- idote. Peduncle slender, to 25 cm tall, terete, gray lepidote. Glomerulescence a compound cyme of numerous (> 100) glomerules. Glomerules 6-9 mm tall, 5-12 mm diam., hemispherical. Heads 1-2 per glomerule, slightly w free. In- volucres cylindric, 5-7.5 m 2-3.5 m diam. with prominent truncate ie patera in 6—7 series; outermost phyllaries widely deltate, 0.5-1.2 mm long, 0.5-1.0 mm wide, the apices obtuse; innermost phyllaries lanceolate, 4—6.2 mm long, 0.6-2 mm wide, the apices acute to acuminate; margins entire to subfimbriate; abax- ial surfaces stramineous with purple apex, gla- brate to lepidote-tomentulose. Florets 3 per head; corollas purple, 5-6 mm tall, the lobes acumi- nate. Anthers 2-3 mm long, the apical appendage acuminate, the bases acute to rounded. Achenes 2.5-4 mm tall, sparsely glandular, rarely pilose, 10-ribbed, the apex slightly constricted and dark; nectary 0.2-0.5 mm tall. Pappus 3-4-seriate, of stramineous (rarely purple), tardily deciduous, strigose bristles; outermost series 1-2 mm long; innermost series 5.5-7.5 mm long. Flowering and fruiting occur from July to Sep- tember Eremanthus pohlii is restricted to the Chapada dos Veadeiros, which runs north to south in south central Goiás, and nearby parts of Minas Gerais MACLEISH — EREMANTHUS REVISION 281 (Figs. 6, 7). It occurs in colonies on the cerrado at high elevations (ca. 1,400 m). Eremanthus pohlii is one ofthe more primitive (i.e., approaching Vernonia) members of the subg. Vanillosmopsis. It is closely related to E. gra- ciellae and E. brasiliensis, both of which are re- stricted to the northwestern arm of the Central Plateau. Eremanthus pohlii can be separated from E. graciellae and E. brasiliensis by its three flo- rets per head and relatively long leaves (6.5-7.5 vs. 2.5-6.5 cm long). The only other member of the Vanillosmopsis complex that occurs sym- patrically with these taxa is E. uniflorus. Ere- manthus uniflorus is most closely related to E. capitatus and is distinguished (among other less discriminating characters) by a single floret per head and 3-9 heads per glomerule. Additional specimens examined. BRAZIL. GOIAS: Brasilia towards Campos Belos, 15 km from Alto Par- áiso, 1,400 m, 28 Aug. 1981, Schumacher 1032, 1033 (GA, MB), Schumacher 1034 (MB, NY). MINAS GERAIS: S Pinheiro, 15 Aug. 1967, Heringer 11531 (NY, RB). 12. Eremanthus graciellae MacLeish & Schu- macher, Syst. Bot. 9: 87. 1984. TYPE: Brazil. Bahia: BR 020 Brasília Richtung Barreiras, 15 km weiter in Richtung Barreiras von Fa- zenda Prainha, km 374, 800 m, 28 Aug. 1981, Schumacher 1048 (holotype, RB; iso- types, GA, K, M, MB) Small tree, to 2.5 m tall; stems gray-brown lepidote; branches many. Leaves subcoriaceous, petiolate; petioles 8-15 mm long, lepidote; blades 4.5-6.5 cm long, 1.5-3 cm wide, elliptic to nar- rowly elliptic, the bases acute, the apices acute to obtuse, the margins entire; adaxially glabrate, abaxially gray lepidote. Peduncle slender, to 25 cm tall, terete, gray lepidote. Glomerulescence a compound cyme of numerous (more than a hundred) glomerules. Glomerules 6-9 mm tall, 4-9 mm diam., obconic. Heads 1-2 per glomer- ule, slightly appressed, free. Involucres cylindric, 5.2-7.5 mm tall, 2.1-2.8 mm diam., with prom- inent truncate base; phyllaries in 6-7 series; out- ermost phyllaries widely deltate, 0.6-1.2 mm long, 0.7-1.2 mm wide, the apices acute; inner- most phyllaries lanceolate, 3.8—6. Ë mm long, 0.6— 1.4 mm wide, the apices acute acuminate; margins entire to subfimbriate; iud ee ee stramineous with purple apex, glabrate to lepi- dote-tomentulose. Florets 2 per head; corollas purple to white, 4.2-7 mm tall, the lobes acu- 282 © E. capitatus Sani “ae \ Jes A E. erythropappus 1 ox J e EA O E. pohlii O E. polycephalus Q: x ` ROME y M "X ES p A. Wes ades FIGURE 7. minate. Anthers 2.1-2.8 mm long, the apical ap- pendage acuminate, the bases acuminate. Achenes 2.2-3.5 mm tall, sparsely glandular, 10- ribbed, the apex slightly constricted and dark; nectary 0.2-0.5 mm tall. Pappus 3-5-seriate, of stramineous to white, deciduous, strigose bris- tles; outermost series 1-2 mm long; innermost series 5-7.2 mm long Flowering and fruiting occur from July to Au- gust Eremanthus graciellae is distributed along the Serra Geral de Goiás, which runs north to south along the border of Goiás and Bahia (Fig. 6). It occurs in colonies on the quartzite grasslands characteristic of this plateau. Eremanthus graciellae is also one of the prim- itive (approaching Vernonia) members of Ere- manthus. It can be separated from close relatives, E. graciellae and E. brasiliensis, by its two florets per head and intermediate length leaves. Additional specimens examined. BRAZIL. BAHIA: BR eis Brasília Richtung Barreiras, 15 km weiter in Rich- ung Barreiras von Fazenda Prainha, 800 m, 28 Aug. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Elevation: 500 meters 1000 i Me AME A. o 200 kilometers Distribution of Eremanthus subg. Vanillosmopsis in Minas Gerais. 1981, as 1046 (GH, MB), Schumacher 1047 (GA, MB). 13. Eremanthus brasiliensis (Gardner) Mac- eish, comb. nov. Monosis brasiliensis Gardner, London J. Bot. 5: 230. 1846. Ver- nonia brasiliensis (Gardner) Schultz-Bip., Jahresber. Pollichia 18-19: 161. 1861, nom. illeg. (nom. rej. pro Vernonia brasiliensis (Sprengel) Less., Linnaea 6: 681-682. 1831. = Albertinia brasiliensis). Vanillos osmopsis brasiliensis (Gardner) Schult ia 0. F (fragments from G and P, photos of G and ; ) Small tree, to 4 m tall; stems gray-brown lep- idote; branches many. Leaves subcoriaceous, petiolate; petioles 5-10 mm long, lepidote; blades 2.4-4.2 cm long, 0.8-2.6 cm wide, elliptic to narrowly elliptic, the bases acute to attenuate, the apices obtuse, the margins entire; adaxially 1987] glabrate and glandular, abaxially gray lepidote. Peduncle slender, terete, gray lepidote. Glomer- ulescence a compound cyme of 100 or more glomerules. Glomerules 6-8 mm tall, 4-8 mm diam., obconic. Heads 1-2 per glomerule, slight- ly appressed, ie Involucres cylindric, 5-6.5 m tall, 2-2. iam.; phyllaries in 6-7 series; outermost uires widely deltate, 0.6-1 mm long, 0.5—1 mm wide, the apices acute; innermost phyllaries lanceolate, 3.5-6 mm long, 0.5-1 mm wide, the apices acute to acuminate; margins en- tire to subfimbriate; abaxial surfaces stramin- eous, glabrate to lepidote-tomentulose. Florets 1 er head; corollas not seen. Achenes 2-3.5 mm tall, sparsely glandular, obscurely-ribbed, the apex slightly constricted and dark; nectary 0.3-0.4 mm tall. Pappus 3-4-seriate, of stramineous, tardily deciduous, strigose bristles; outermost series 1— 2 mm long; innermost series 5-7 mm long. Flowering and fruiting probably occur from August to September. Eremanthus brasiliensis is distributed in the northern part of the Serra Geral de Goias (Fig. 6), which runs north to south along the border of Goiás and Bahia. This species is known only from the type specimens. Schumacher (1982) un- successfully attempted to locate it at the same time that he rediscovered E. arboreus in southern Ceará. Like E. graciellae and E. pohlii, E. brasiliensis is one of the more primitive members of the Vanillosmopsis complex. It is distinguished by one floret per head, small leaves (2.5—4.5 vs. 4.5- 7.5 cm long), and heads which lack the promi- nent truncate bases of its closest relatives. Eremanthus Less. sect. Vanillosmopsis (Schultz- Bip.) MacLeish, comb. nov. Vanillosmopsis subg. Euvanillosmopsis Schultz-Bip., Jah- resber. Pollichia 20-21: 398. 1863. LEC- TOTYPE: Eremanthus erythropappus (DC.) MacLeish. Heads cylindric or obconic. Heads per glom- erule (2-)5—12. Pappus often twisted. 14. Eremanthus polycephalus (DC.) MacLeish, comb. nov. Albertinia polycephala DC., Prodr. 5: 82. 1836. Vanillosmopsis poly- cephala (DC.) Schultz-Bip., Jahresber. Pol- lichia 18-19: 168. 1861. TYPE: Brazil. Minas Gerais: planitie alta ad Piedade Villam, Martius s.n. (holotype, M; isotype, M). MACLEISH — EREMANTHUS REVISION 283 Albertinia pcd: C. Martius ex DG., Prodr. 5: 82. 1836. Vanillosmopsis saligna (DC. ) Schultz-Bip., eed iai Pollichia 18-19: 168. 1861. TYPE: Bra- zil. Minas Gerais: in editis siccis rupestribus mon- ium Serro Frio, Martius s.n. (holotype, M). Shrub or small tree, to 3.5 m tall; stems gray- brown lepidote; branches many. Leaves mem- branaceous to subcoriaceous, sessile to petiolate; petioles to 8 mm long, lepidote; blades 4.5-7 cm long, 1-1.8 cm wide, lanceolate, the bases acu- minate to attenuate, the apices acute to obtuse, the margins entire; adaxially glabrate, abaxially gray lepidote. Peduncle slender, to 23 cm tall, terete, gray lepidote. Glomerulescence a com- pound cyme of 100 or more glomerules, these 5—7 mm tall, 4-8 mm diam., hemispherical. Heads 6-12 per glomerule, closely appressed, connate !'^ lengt tissues. Involucres cylindric, 4—5. 2 mm diam.; phyllaries in 6—7 series; outermost phyllaries widely deltate, 0.6-1.2 mm long, 0.7- 1.2 mm wide, the apices obtuse; innermost phyl- laries lanceolate, 3-5 mm long, 0.5—1.2 mm wide, the apices acute as Svar the margins entire purple apex, lepidote-tomentose. Florets 1 per head; corollas purple, 4—6.5 mm tall, the lobes acuminate to acute. Anthers ae : mm long, the apical appendage acuminate, ases acute. Achenes 1.5-2.8 mm tall, ee 10-ribbed, the apex slightly constricted and dark; nectary 0.2-0.4 mm tall. Pappus 3—4-seriate, of stramin- eous, white or purple, tardily deciduous, strigose bristles; outermost series 0.2-1 mm long, the in- nermost series 3-5.5 mm long. Flowering and fruiting occur from June to No- vember Eremanthus polycephalus 1s distributed along the Serra do Espinhaço, which runs north to south through the center of Minas Gerais (Fig. 7). It occurs in colonies on the campo rupestre habitats characteristic of high elevations (700-1,370 me- ters). The common name of “‘candeia,”’ or can- dle, is applied locally. Eremanthus polycephalus is most closely re- lated to E. erythropappus and is sympatric with it in the northernmost portion of its range. Uniquely, in subg. Vanillosmopsis, both of these taxa exhibit concrescence of phyllary and recep- tacle tissues. However, E. polycephalus is easily distinguished by its single floret per head (vs. 3- 4) and partial fusion of heads (vs. nearly total fusion). This species, because of its single floret 284 per head and various characteristics associated with the Vanillosmopsis complex, superficially resembles E. uniflorus and E. brasiliensis, taxa that are restricted to Goiás and nearby Bahia and which are distinguished by their lack of tissue fusion within glomerules. Additional dodge examined. BRAZIL. MINAS GERAIS: 14 km NE o mantina towards Mendanha on Estrada 367, 1,370 A p Oct. 1980, MacLeish et al. 702 (GA, GH, K, LE, NY, RB, UB, US Possible hybrids. x E. elaeagnus (C. Martius ex DC.) Schultz-Bip. MINAS GERAIS: Serra do Cipó, 7 Oct. 35$ MacLeish et al. 718 (GA). x E. incanus (Less.) MINAS GERAIS: Itamarandiba, 6 Sep. 1981, Schu- LU 1116a (GA). 15. Eremanthus erythropappus (DC.) Mac- Leish, comb. nov. Albertinia erythropappa DC., Prodr. 5: 82. 1836. Vanillosmopsis erythropappa (DC.) Schultz-Bip., HUE er. Pollichia 18-19: 167. Minas Gerais: Marianne, Vauthier 334 (ho- lotype, G-DC (as IDC microfiche); isotypes, G, 2 in GH). Albertinia p lay d London J. Bot. 5: 235. l : Brazil. Minas Gerais: near Villa do Principe, nei 1840, Gardner 48 12 (holotype, BM; s, F, 2 in G, 2 in GH, 3 in NY, S). Vanillosmopsis srera Schultz- Bip., Jahresber. Pollichia 18-19: 167. . Vernonia glomerata Schultz-Bip., Bot. Zeitung. (Berlin) 3: 155. 1845. nom. nud syntypes; Brazil. Minas Gerais: Aug.-Apr. 1840, Claussen 2063 (G; isolectotypes, G, 2 in GH, M, MO). Syntypes: Brazil. (Rio de Janeiro): in col- libus siccis calidisque inter Mandioca et Serra Es- trella, Riedel s.n. (577) (not found); no other data, Claussen 863 (853) (not found); no other data, Schucht 75 (W, not found) Shrub or tree, to 10 m tall, to 10 cm diam.; stems gray-brown lepidote; branches many. Leaves membranaceous, petiolate; petioles 3-15 mm long, lepidote; blades 5—10.2 cm long, 1.6— 3.6 cm wide, oblanceolate to elliptic, the bases acute to acuminate, the apices acute to attenuate, the margins entire; adaxially glabrate to lepidote, abaxially gray lepidote. Peduncle slender, to 23 cm tall, terete, gray lepidote. Glomerulescence a compound cyme of 100 or more oe Glomerules 6-15 mm tall, 4-12 m m., per cole closely appressed, connate nearly entire length by con- crescence of phyllary tissues and pubescence. In- Moin obconic, 5-7.2 mm tall, 2.1-4.2 m diam.; phyllaries in 5—6 series; outermost p alice widely deltate, 0.6-1.5 mm long, 0.7-1.3 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 mm wide, the apices acute; innermost phyllaries lanceolate, 3.6-6.8 mm long, 0.5-1.2 mm wide, the apices acute to acuminate; margins entire to subfimbriate; abaxial surfaces stramineous with all, the lobes acu- minate. Anthers 2.1—2.7 mm long, the apical ap- pendage acuminate, the bases acute. Achenes cylindric-turbinate, 1.2—2.5 mm tall, glandular, 10-ribbed, the apex slightly constricted and dark; nectary 0.15-0.25 mm ls Pappus 3-4-seriate, of stramineous, white or purple, often curling, promptly deciduous, ida. bristles; outermost eries 1.2-3 mm long; innermost series 3-6.5 mm long. Flowering and fruiting occur from June to No- vember. remanthus erythropappus is distributed throughout southeastern portions of the Central Plateau (Fig. 7) at 700 to 2,400 meters. It is ex- tremely common in colonies amidst secondary forest of the coastal range and cerrado of the interior plateau. It is often used as a living fence around fields because the high terpenoid content of its leaves deters insects. Also, it is commonly known as “‘candeia,” candle, or “pau de can- deia," candlestick. Because of its extensive range, E. erythropap- manthus erythropappus is further distinguished by its 3-4 florets per head (vs. 1), 6-12 heads per glomerule (vs. 30-100 in E. incanus), and nearly total connation of heads (vs. partial con- nation in E. polycephalus). The epithets assigned by Gardner (1846) and Schultz-Bip. (1861) to this species are indicative of the variation as- cribable to age and habitat differences. Additional specimens examined. BRAZIL. ESPÍRITO SANTO: Mun. de Cachoeus do Itapemirim, Vargem Alta Morro do Sal, 700 m, 3 Aug. 1980, Ferreira & Borges 1882 (GA), 21 es 1948, Brade 19340 (RB). MiNAS GERAIS: Serra rral, Nova Lima, 1,300 m, 6 Aug. 1945, Williams & Assis 8029 (BM, BR, C, F, G, GH. MO, N ). RIO DE JANEIRO: Teresopolis, Serra das B 24 Sep. 1980, MacLeish et al. 670 (C, GA, nid LE, MO, NY, RB, B, W). são PAULO: Serr: A Ae uy: km m Sao José do Barreiro idle k of Bocaina, 1,640 m, 30 Sep. 1980, PAS dei et al 679 (B, BM, CEPEC, GA, RB, S, TEX, W). 1987] Possible hybrids. x E. capitatus (Sprengel) Mac- Leish. MINAS GERAIS: Carai, 6 Sep. 1981, Schumacher 1083, 1085, 1089 (GA). x E. incanus (Less.) Less. MINAS GERAIS: Tiradente, 10 Sep. 1936, Barreto 4792 (F). x E. seidelii MacLeish & Schumacher. MINAS GERAIS: Furnas towards Piui, 25 macher 1005 (GA), Alpinopolis, 12 Sep. 1977, Seidel . (RB). 16. Eremanthus uniflorus MacLeish & Schu- aa Syst. Bot. 9: 93. 1984. TYPE: Brazil. Goiás: 33 km N of Alto Paráiso towards P IE 1,370 m, 14 Oct. 1980, MacLeish, Martinelli, Smith & Stutts 736 (holotype, RB; isotypes, F, G, GA, K, M, NY, P, S, UB, US) Shrubby tree, to 2.5 m tall; stems gray-brown lepidote; branches many. Leaves membrana- ceous to subcoriaceous, petiolate; petioles 5-20 mm long, lepidote; blades 3.8-8 cm long, 1.2-4 cm wide, elliptic, the bases acute, the apices acute, the margins entire; adaxially glabrate, abaxially gray lepidote. Peduncle slender, to 25 cm tall, terete, gray lepidote. Glomerulescence a com- pound cyme of 100 or more glomerules, these 3-7 mm tall, 3-10 mm diam., hemispherical. Heads 3-9 per glomerule, slightly appressed and adherent basally by pubescence of phyllaries. In- volucres cylindric, 4-7 mm tall, 1-2 mm diam.; phyllaries in 5-7 series; outermost phyllaries trullate, 0.7-2 mm long, m wide, the apices acute; innermost phyllaries m 3.5- 5.5 mm long, 0.5-0.9 mm wide, the apices acu- minate; margins entire to subfimbriate; abaxial surfaces stramineous with purple apex and mid- vein, glabrate to lepidote-tomentulose. Florets | per head; corollas purple (to white with age), 5- 7 mm tall, the lobes acuminate. Anthers 2.7—3.8 mm long, the apical appendage acuminate, the bases acuminate. Achenes 2.4—3.2 mm tall, glan- dular, 10-ribbed, the apex slightly constricted and dark, occasionally pilose at base; nectary 0.05- 0.1 mm tall. Pappus 3—5-seriate, of purple to stramineous, deciduous, strigose bristles; outer- most series 0.8-1.8 mm long; innermost series 5.5-7 mm long. Flowering and fruiting occur from August to October. Eremanthus uniflorus is restricted to the Cha- pada dos Veadeiros in central Goiás (Fig. 6). Although locally abundant, it is found only on campo rupestre outcrops A single floret per head. the relatively weak coherence of heads, and 3-9 heads per glomerule MACLEISH — EREMANTHUS REVISION 285 distinguish E. uniflorus from other members of sect. Vanillosmopsis. This taxon is closely related to E. capitatus, which is primarily restricted to the northeast arm of the Central Plateau and differs in that E. capitatus has 3-4 florets per head and 2-5 heads per glomerule. Additional specimens examined. BRAZIL. GOIAS: Brasilia Richtung Alto Paráiso de Goiás, 50 km nach Joà m of Alto Paráiso, 1,220 m, 24 Jan 1980, King & Almeda 8291 (US). Possible hybrids. macher. Goás: Alto Paraiso, 28 macher 1038, 1039, 1041 (GA) x E. argenteus MacLeish & Schu- Aug. 1981, Schu- 17. Eremanthus capitatus (Sprengel) MacLeish, comb. nov. Conyza capitata Sprengel, Syst. Veg. 3: 507.1826. Vernonia capitata (Spren- gel) Less., Linnaea 4: 270. 1829. Albertinia capitata (Sprengel) DC., Prodr. 5: 82. 1836. Vanillosmopsis capitata (Sprengel) Schultz- Bip., Jahresber. Pollichia 18-19: 167. 1861. LECTOTYPE: here designated; Brazil. Bahia: inter Victoria et Bahia (now Salvador), Sel- low s.n. (K; isolectotype, GH). Polypappus discolor DC., Prodr. 7: 281. 1838. Vanil- losm opsis rere (DC. ) Baker in C. Martius, Fl. Bras. 7.1873. TvPE: Brazil. Bahia: Jacobina, Jan. 1843, Bianche el (holotype, G; vile BM, 4 in F (one photo of C), 2 in GH, k albertinioides ppa Bip., Jahresber. Pollichia 18— 168. TYPE: Brazil: no other da s ented Ere B, not found). Shrub or tree, to 6 m tall, to 8 cm diam.; stems gray-brown lepidote; branches many. Leaves membranaceous to subcoriaceous, sessile to pet- iolate; petioles 1-12 mm long, lepidote; blades 4.6-10.5 cm long, 2.4—4.2 cm wide, elliptic to obovate, the bases acute, the apices acute to acu- minate, the margins entire; adaxially glabrate to lepidote, abaxially gray lepidote. Peduncle slen- der, to 30 cm tall, terete, gray lepidote. Glomer- ulescence a compound cyme of 100 or more glomerules. Glomerules 4-10 mm tall, 3-12 mm diam., hemispherical. Heads 2-5 per glomerule, slightly appressed and coherent basally by pu- bescence of phyllaries. Involucres cylindric to obconic with age, 5-8 mm tall, 2-4 mm diam.; phyllaries in 5-6 series; outermost phyllaries widely deltate, 0.5-1.1 mm long. 0 2 mm wide, the apices acute; innermost phyllaries lan- ceolate, 3.4-7.6 mm long, 0.6-1.4 mm wide, the apices acuminate; margins entire to subfim- 286 briate; abaxial surf: t to green, gla- brate to lepidote-tomentulose. Florets 3(-4) per head; corollas purple with white tube and be- coming white with age, 4.2-7.2 mm tall, the lobes acuminate. Anthers 2.2-2.8 mm long, the apical appendage acuminate, the bases acute. Achenes 2-2.5 mm tall, glandular, 10-ribbed, the apex slightly constricted and dark; nectary 0.3-0.4 mm tall. Pappus 3—4-seriate, of purple, white or stra- mineous, often curling, promptly deciduous, stri- gose bristles; outermost series 2.5-3.5 mm long; innermost series 4—6.5 mm long. Flowering and fruiting occur from July to No- vember. Eremanthus capitatus is distributed through- out the northeastern arm of the Central Plateau (Figs. 6, 7) at elevations of 100 to 1,000 meters. It occurs in large colonies in cerrado or on the border of secondary forest and is known as “‘can- deia,” or candle, in Bahia. Eremanthus capitatus is closely related to E. uniflorus, E. arboreus, and E. erythropappus. It is distinguished by its 3—4 florets per head and 2-5 heads per glomerule. Eremanthus uniflorus is likely derived from an isolated population of E. capitatus in the northwestern arm of the Cen- tral Plateau whose number of florets per head has decreased from three to one. In contrast, E. arboreus represents a population found in the extreme northeastern part of the plateau whose number of heads per glomerule has increased from 2-5 to 6-9. Eremanthus capitatus and E. erythropappus have the same number of florets per head, but E. capitatus lacks the fusion of receptacle and phyllary tissues observed in E. erythropappus. Flowering or immature specimens of this tax- on are frequently misidentified as E. pohlii. When young, each head is noticeably cylindric like those of E. pohlii; however, as fruits mature, the in- nermost phyllaries drop off and the outermost spread out so that heads appear to be cylindric- turbinate. Sprengel's types are generally considered to have been deposited at B, BM, BR, or P. Un- fortunately, a holotype was not located in any of these herbaria. However, a specimen was located at K bearing the citation “ex herb. B" and the correct citation in Sprengel's handwriting. There- fore, the Kew specimen has been chosen as lec- totype. Additional specimens examined. BRAZIL. BAHIA: 33 km from BR 101 on road S to Canavieiras, 23 Oct. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 1980, MacLeish & Soares ind 758 (BR, C, CEPEC, F, GA, GH, MO, NY, RB, UC, US). GorAs: 5-10 km N of Veadeiros, Valley of the Rio Paraná, 19 Jul. 1964, Prance & Silva 58251 (F, NY, RB, S, UB). MINAS GERAIS: BR 116 Téofilo Otóni towards Bahia, 2 km from Padre Paraiso, 8 Sep. 1981, 600 m, Schumacher 1082 (GA, MB). Possible hybrids. MacLeish. MINAS GERAIS: Carai, 6 Sep. macher 1083, 1085, 1089 (GA). x E. erythropappus (DC.) 1981, Schu- 18. pisi arboreus (Gardner) MacLeish, comb. Albertinia arborea Gardner, Idea. Bot 5: 236. 1846. Vanillosmopsis arborea (Gardner) Baker in C. Martius, Fl. Bras. 6(2): 16. 1873. TYPE: Brazil. Ceara: Serra de Araripe, Nov. 1838, Gardner 1713 (holotype, BM; isotypes, BR, 4 in F (includ- ing fragment from P and photo of B), 2 in G, 3 in GH (one is photo of B), 2 in NY). Tree, to 6 m tall; stems gray-brown lepidote; branches many. Leaves membranaceous to sub- coriaceous, petiolate; petioles 5-10 mm long, lepidote; blades 5—6.5 cm long, 1.6-2 cm wide, narrowly elliptic to obovate, the bases attenuate, the apices acute to acuminate, the margins entire; adaxially glabrate to lepidote, abaxially gray lep- tall, 8-15 mm diam., hemispherical. Heads 6-9 per glomerule, slightly appressed and coherent basally by pubescence of phyllaries. Involucres obconic, 5-7.5 mm tall, 2-3.5 mm diam.; phyl- laries in 5-6 series; outermost phyllaries widely deltate, 0.6-1 mm long, 0.5-1.2 mm wide, the apices acute; innermost phyllaries lanceolate, 4— 6 mm long, 0.5-1.2 mm wide, the apices acute to acuminate; margins entire to subfimbriate; abaxial surfaces stramineous to green, glabrate to lepidote-tomentulose. Florets 3(-4) per head; corollas white or purple, 4-7 mm tall, the lobes acuminate. Anthers 2-3 mm long, the apical ap- pendage acute, the bases acute. Achenes 1.5-2.5 mm tall, sparsely glandular, 10-ribbed, the apex slightly constricted and dark; nectary 0.25-0.35 mm tall. Pappus 3-5-seriate, of stramineous to off-white (occasionally purple basally), often curling, promptly deciduous, strigose bristles; outermost series 2.5-3 mm long; innermost se- ries 4—6 mm long. debui. and fruiting occur from August to September 1987] Eremanthus arboreus is restricted to the north- eastern slope of the Chapada do Araripe (Fig. 6), near Crato, which runs along the border of Ceara and Pernambuco at elevations of 700 meters. It occurs in colonies on wooded slopes near the top of the plateau which are less dry than the sur- rounding caatinga. Eremanthus arboreus is closely related to E. capitatus and is distinguished by its 6-9 heads per glomerule. This species probably represents an extreme northeastern population of E. capi- tatus whose number of heads per glomerule has increased from 2-5 to 6-9. Until recently, this species was known only from the type collection. However, Schumacher (1982) was able to collect it again from what was probably Gardner’s orig- inal site during several expeditions designed to rediscover members of Eremanthus that were known only from type specimens. Additional specimens examined. BRAZIL. CEARA: Chapada do Araripe, hillside on oe g of Crato, 25 Aug. 1980, Schumacher s.n. (G RB), Crato to- wards Nova Olinda, at slope near Ps ‘of plateau, 700 m, 31 Aug. 1981, Schumacher 1049 (B, GA, K, MB RB). EXCLUDED TAXA 1. Eremanthus Less. sect. Chresta (Vell. Conc. ex DC.) Baker in C. Martius, Fl. Bras. 6(2): 166.1873. Chresta Vell. Conc. ex DC., Prodr. 5:85. 1836. TYPE: (Robinson, 1980): Chresta sphaerocephala DC. = Chresta Vell. Conc. 2. Eremanthus Less. sect. Pycnocephalum (Less.) Baker in C. Martius, Fl. Bras. 6(2): 168. 1873. Vernonia Schreber sect. Pycnocephalum ss., Linnaea 6: 629. 1831. TYPE: (Mac- Leish, 1985a): Pycnocephalum plantagini- Kori (Less.) DC. = Pycnocephalum (Less.) 3. boc Less. sect. Sphaerophora (Schultz-Bip.) Baker in C. Martius, Fl. Bras. 6(2): 165. 1873. Sphaerophora Schultz-Bip., Jahresber. Pollichia 20-21: 402. 1863, nom. illeg. (non Sphaerophora Blume, 1850, Ru- biaceae). Paralychnophora MacLeish, Tax- on 33: 106. 1984. TYPE: Paralychnophora bicolor (DC.) MacLeish = Paralychnophora MacLeish. 4. Eremanthus Less. sect. Stachyanthus (DC.) Baker in C. Martius, Fl. Bras. 6(2): 167. 1873. Stachyanthus DC., Prodr. 5: 84. 1836, nom. rej. (vs. Stachyanthus Engler, 1897, Icaci- naceae). Argyrovernonia MacLeish, Taxon MACLEISH — EREMANTHUS REVISION 5. 6. N oo 9. 12. 287 33: 106-107. 1984. TYPE: Argyrovernonia martii (DC.) MacLeish — Argyrovernonia MacLeish. Eremanthus Less. subg. Pseuderemanthus Schultz-Bip. “B. Jodopappus" Schultz-Bip., Jahresber. Pollichia 20-21: 396. 1863. TYPE: Vernonia crotonoides (DC.) Schultz-Bip. — Vernonia Schreber subg. Vernonia sect. Ver- n Eremanthus angustifolius (Gardner) Baker in C. Martius, Fl. Bras. 6(2): 170. 1873. Chres- ta angustifolia Gardner, London J. Bot. 1: 240, tab. 8. 1842. TYPE: Brazil. Goiás: shady places at Arrayas, Gardner 3802 (holotype, BM); isotypes, 3 in F (includes fragment from P and photo of B), 2 in G, GH (photo of B), NY, S, W) = Pycnocephalum angustifolium (Gardner) MacLeish (19852). . Eremanthus bicolor (DC.) Baker in C. Mar- tius, Fl. Bras. 6(2): 165. 1873. Albertinia bi- color DC., Prodr. 5: 81. 1836. TYPE: Brazil. Minas Gerais: altis, Martius s.n. (holotype, M; isotype, M) = Paralychnophora bicolor (DC.) MacLeish (1984a). . Eremanthus crotonoides (DC.) Schultz-Bip., Jahresber. Pollichia 20-21: 396. 1863. Al- bertinia crotonoides DC., Prodr. 5: 81. 1836. TYPE: Brazil. Minas Gerais: montium sepi- bus, Martius s.n. (holotype, M) = Vernonia crotonoides (DC.) Schultz-Bip. Eremanthus curumbensus Philipson, Kew Bull. 1938: 298. 1938. TYPE: Brazil. Goias: valley of the Rio Curumbo in the plains be- tween the mountains, Glaziou 21645 (ho- lotype, K; isotypes, BR, C, 4 in G, NY) = Glaziovianthus curumbensis (Philipson) MacLeish (1985b). . Eremanthus descampsii Klatt ex De Wild. & T. Durand, Ann. Mus. Congo, Ser. 1, Bot. 1: 99. 1898, non Vernonia descampsii De Wild., Bull. Jard. Bot. État. 5: 97. 1915. TYPE: Rep. of the Congo, Katanga, Vallee de la Liula, 1891, Descamps s.n. (holotype, BR not seen; fragment and drawing of BR, GH) — Vernonia klattii MacLeish (1984b). . Eremanthus eriopus (Schultz-Bip.) Baker in C. Martius, Fl. Bras. 6(2): 169. 1873. Pres- telia eriopus Schultz-Bip., Naturf. Ges. Em- den 1864: 73. 1864. rvPE: Brazil. Minas Gerais: in glareosis S. da Lapa, Riedel 1127 (holotype, LE; isotypes, 2 in F (fragment and photo of P), P, TEX (photo of P)) = Prestelia eriopus Schultz-Bip. Eremanthus exsuccus (DC.) Baker in C. — Ww > — WN — a — - — [re] — o N e ANNALS OF THE MISSOURI BOTANICAL GARDEN Martius, Fl. Bras. 6(2): 166. 1873. Chresta exsucca DC., Prodr. 5: 85. 1836. TYPE: Bra- zil. Minas Gerais: montosis Minarum, Mar- tius s.n. (holotype, M; photos of M, F, TEX) — Chresta exsucca D . Eremanthus harmsianus Taub., Bot. Jahrb. Syst. 21: 453. 1896. TYPE: Brazil. Goiás: Ser- ra dos Pyreneos, Ule 2984 (holotype, HBG, not seen; isotype, P) = Glaziovianthus spe- ciosus (Gardner) MacLeish (1985b). Eremanthus imbricatus G. Barroso, Rodri- guésia 23-24: 6. 1962. TYPE: Brazil. Distrito Federal: Brasilándia, Macedo 4 (holotype, RB; isotype RB) = Chresta exsucca DC . Eremanthus jelskii Hieron., Bot. Jahrb. Syst. 36: 462. 1905, non Vernonia jelskii Hieron., Bot. Jahrb. Syst. 36: 459. 1905. TYPE: Peru, crescit prope Shanyn, Je/skii 776 (lectotype, US (MacLeish, 1984b); isolectotypes, pho- tos of B, F, GH, NY, TEX) - Vernonia shanynensis MacLeish (1984b). . Eremanthus labordeii Glaz., Mem. Soc. Bot. France 3: 380. 1909. TYPE: Brazil. Goiás: entre Paranaua et Rajadinho, 31 Jun. 1895, Glaziou 21675 (lectotype, P (MacLeish, 1985a); isolectotypes, BR, C, 2 in F, 2 in G, GH) = Chresta exsucca DC. Eremanthus leucodendron Mattf., Bot. Gart. Notizbl. 9: 378. 1925. TYPE: Brazil. Bahia: Rio de Contas, Serra das Almas, Carrasco, Luetzelburg 242 (holotype, M; isotypes, F (photo of B), 2 in GH (includes photo of B), M, TEX (photo of B)) — Vernonia leuco- dendron (Mattf.) MacLeish (1984b) . Eremanthus martii (DC.) Baker in C. Mar- tius, Fl. Bras. 6(2): 167. 1873. Stachyanthus martii DC., Prodr. 5: 84. 1836. TYPE: Brazil. Bahia: siccis sylvis aestu aphyllis, ad Ju- azeiro, Martius s.n. (holotype, M; isotypes, 3 in M) - Argyrovernonia martii (DC.) MacLeish (19842). . Eremanthus mollis Schultz-Bip., Jahresber. Pollichia 18-19: 166. 1861, non Vernonia mollis Kunth, Nov. Gen. & Sp. 4: 36. 1820. TYPE: Brazil. Goiás: Montes Claros et Ponte Alto, ante Bomfim, Pohl 171 (lectotype, W (MacLeish, 1984b); isolectotype, 2 in F (fragment and photo of B), GH (photo of B), 2 in NY, TEX (photo of B)) = Vernonia pannosus (Baker in C. Martius) MacLeish (1984b). Eremanthus pabstii G. Barroso, Sellowia 16: 173. 1964. TYPE: Brazil. Goiás: Cristalina, Heringer 9229/1442 (holotype, HB; iso- N 22, N [9v] 2 A N ws 2 nN 2 ~ 2 co t3 e [VoL. 74 types, RB, UB) = Vernonia pabstii (G. Bar- roso) MacLeish (1984b) . Eremanthus pannosus Baker in C. Martius, Fl. Bras. 6(2): 164. 1873. TYPE: Brazil. Goiás: Curralinho, Manso 1 (holotype, BR) = Ver- nonia pannosus (Baker in C. Martius) MacLeish (1984b). Eremanthus pinnatifidus Philipson, Kew Bull. 1938: 299, 1938. TYPE: Brazil. Rio de Janeiro: env. de Rio de Janeiro, Glaziou 14033 (lectotype, C (MacLeish, 1985a)) = Pycnocephalum pinnatifidum (Philipson) MacLeish (19852) . Eremanthus plantaginifolius (Less.) Baker in C. Martius, Fl. Bras. 6(2): 168. 1873. Ver- nonia plantaginifolius Less., Linnaea 4: 251. 1829. TYPE: Brazil: no other data, Se//ow s.n. (lectotype, P (MacLeish, 1985a)) = Pycno- cephalum plantaginifolium (Less.) DC. . Eremanthus purpurascens Glaz. ex Oliver, Hooker's Icon. Pl. 4(3): plate 2282. 1894. TYPE: Brazil. Minas Gerais: Serra do Cipó, Glaziou 19464 (holotype, P; isotypes, BR, C) = Prestelia eriopus Schultz-Bip. . Eremanthus pycnocephalus (DC.) Baker in C. Martius, Fl. Bras. 6(2): 166. 1873. Chres- ta pycnocephala DC., Prodr. 5: 85. 1836. TYPE: Brazil. Minas Gerais: campis deserti inter Min. Nov. et f. S. Francisci, Martius s.n. (holotype, M) = Chresta pycnocephala DC ; Eremanihins reflexo-auriculatus G. Barroso, 6. 1962. TYPE: Brazil. — Paralychnoph- ora reflexoauriculata (G. Barroso) MacLeish ) (1984a . Eremanthus rivularis Taubert, Bot. Jahrb. Syst. 21: 453. 1896. TYPE: Brazil. Goiás: re- giao de Maranhào superior, Ule 26 (2962) (holotype, HBG, not seen; isotypes, 2 in F (fragment and photo of P), GH (photo of P), P) — Pycnocephalum angustifolium (Gard- ner) MacLeish (19852) . Eremanthus scapigerus (Less.) Baker in C. Martius, Fl. Bras. 6(2): 168. 1873. Vernonia scapigera Less., Linnaea 4: 250. 1829. TvPE: Brazil: no other data, Sellow s.n. (lectotype, B (MacLeish, 19852); isolectotypes, K, P) — Chresta scapigera (Less.) Gardner. Eremanthus schwackei Glaz., Bull. Soc. Bot. France 3: 380. 1909. TYPE: Brazil. Minas Gerias: Biribiry, Glaziou 19562 (lectotype, here designated, P; isolectotypes, BR, C, G) 1987] o2 © wə — Ww N [9v] UJ o2 A W CA ww nN o2 N MacLEISH— EREM. = Paralychnophora schwackei (Glaz.) MacLeish (1984a). . Eremanthus speciosus ee Baker in C. Martius, Fl. Bras. 6(2): 169. 1873. Pide. speciosa Gardner, aa J. Bot 240. 1842. TYPE: Brazil. Goiás: dry campos near Villa de Arrayas, Gardner 3801 (holotype, BM; isotype, BM) = Glaziovianthus specio- sus (Gardner) MacLeish (1985b). . Eremanthus sphaerocephalus (DC.) Baker in C. Martius, Fl. Bras. 6(2): 167. 1873. Chres- ta spherocephala DC., Prodr. 5: 85. 1836. TYPE: Brazil. Minas Gerais: Tejuco (now Diamantina), Vauthier 294 (lectotype, G-D (as IDC microfiche, MacLeish, 19852); isolectotypes, G, GH) = Chresta sphaero- cephala D QE nasus sphaerocephalus (DC.) Baker in C. Martius var. intermedia (Gardner) Baker in C. Martius, Fl. Bras. 6(2): 167. 1873. Chresta intermedia Gardner London J. Bot. 5: . 1 . TYPE: Brazil. Minas Gerais: near Formigas (now Montes Claros), Gard- ner 4818 (holotype, BM) = Chresta sphaero- cephala DC . Eremanthus veadeiroensis H. Robinson, Phytologia 45: 94. 1980. TYPE: Brazil. Goiás: Chapada dos Veadeiros, ca. 20 km N of Alto Paraiso, /rwin et al. 32752 (holotype, UB; isotype, US) = Vernonia oo (H. Robinson) MacLeish (1984 . Eremanthus verbascifolius pa Martius ex DC.) Schultz-Bip., Jahresber. Pollichi 20- 21: 397. 1863. TYPE: Brazil. Minas Gerais: ferruginosis Serra do Ant. Pereira, Martius s.n. (holotype, M) = Vernonia crotonoides (DC.) Schultz-Bip. . Vanillosmopsis bicolor (DC.) Schultz-Bip., Jahresber. Pollichia 18-19: 168. 1861. Al- bertinia bicolor DC., Prodr. 5: 81. 1836. TYPE: Brazil. Minas Gerais: altis, Martius s.n. (ho- otype, M; isotype, M) — Paralychnophora bicolor (DC.) ipn (1984a). Vanillosmopsis lanceolata (Vell. Conc.) untze, Re en. Pl. 3(2): 183. ^ dd chius ) PM Vell. Conc., Arq, Nac. Rio de Janeiro 8: 350. 1881. TYPE: isnt zil: mediterraneis transalpinus prope pagum Cunha, Velloso s.n. (lectotype, as figure, Fl. Flum. Ic. 8: tab. 151. 1831 (MacLeish, 1985a)) = Identity doubtful. . Vanillosmopsis syncephala Schultz-Bip., Jahresber. Pollichia 18-19: 168. 1861, non Vernonia syncephala Schultz-Bip. ex Baker REVISION 289 in C. Martius, Fl. Bras. 6(2): 64. 1873. TYPE: Brazil: no other data, Sellow 948 (holotype, B (destroyed); photos of B, F, GH) = nonia crispa (Mattf.) MacLeish (1984b). 354. 19 in via a Sandia ad Chunchusmayo, Weber- bauer 1324 (lectotype, NY, photo of B (MacLeish, 1984b); isolectotypes, photos of B, GH, TEX) = Vernonia ramospatana MacLeish (1984b). LITERATURE CITED ABDEL-BASET, Z. H., L. SouTHWICK, W. G. PADOLINA, YOSHIOKA, T. J. MABRY & S. B. JONES, JR. 1971. Sesquiterpene lactones: a survey of 21 United States taxa from the genus Vernonia (Compositae). Phytochemistry 10: 2201 xj: BAKER, J. G. Compositae. I. Vernonieaceae. In C. Martius (editor), Flora Brasiliensis 6(2): 1- 180. C. C. Fortes, E. G. Fortes, G. C. Lo 1972. Chemoprophylactic agents in schistoso- miasis: eremanthine, costunolide, alpha-cyclocos- tunolide, and bisabolol. J. Pharm. Pharmacol. 24: -857. w G. . Um gënera novo da familia mpositae.” Revista Brasil. Biol. 7: 113-115. SEM G. 1873. In G. Bentham and J. D. Hooker (editors), Genera Plantarum 2: 163-236. CANDOLLE, A. P Vernoniaceae. In Pro- PN romus Systematis Naturalis Regni Vegetabilis 5: —94. ae J. P. P. 1973. The text of Vellozo’s Flora Fluminensis and its effective date of publication. Taxon 22: 281-284. EITEN, G. 1978. Delimitation of th ad ncept uis 36: 169-178. 19 Classificação € IMS do Brasil. Edito A EA. 5 M. B R, B. GIL- J. A. “RABI. 1976. ee E enel of eremanthine, a schistosomicidal sesqui- terpene lactone from Eremanthus elaeagnus. dade iyd 15: 331-332. GARDNER, G. Contributions towards a flora of Brazil. London b: Bot. 5: 235-237. HARBORNE, J. C. A. WILLIAMS. 1977. Verno- nieae — ede review. Pp. 523-537 in V. H. Heywood, J. B. Harborne & B. L. Turner (editors), The Biology and Chemistry of the Compositae. Academic Press, London. Heywoop, V. H., J. B. HARBORNE & B. L. TURNER. 1977. An Do to the Compositae. Pp. 1-19 ood, J. B. Harborne & B. L. Turner (editors), The iS Bisa and Chemistry of the Com- positae. Academic Press, London Jones, S. B., JR. 1977. Ch. 17. Vernonieae—a SyS- 290 tematic eh Pp. 501-519 in V. H. Heywood, .H e & B. L. Turner (e ditor), The Bi ology and Chemistry of the Compositae. Academ ic Press, yes KEELEY, S. C. & NES, JR. 1977. Taxonomic implications ia enn pollen morphology to Vernonia (Compositae) in the West Indies. Amer. J. Bot. 64: 576-584 KUNTZE, C. E. O. 1898. 88. CLIMAT In Revisio Generum Plantarum 3(2): 1 LESSING, C. F. De eek herbarii regii berolinensis dissertationes, I. Vernonieae. Linnaea 4: 240-356 De synanthereis herbarii regii beroli- nensis dissertationes, IV. Vernoniearum mantissa Linnaea 6: 624—721 . ABDEL-BASET, INA & S. 1975. Systematic implications of flavonoids and sesquiterpene lactones in species of Vernonia. Biochem Ecol. 2: 185-192. McCain, J. W. & s the tax- onomy of Berberis and Mahonia (Berberidaceae) supported by their rust pathogens Cumminsiella santa sp. nov. and other Cumminsiella species iip ied n Bot. 7: 48-59. MACLEISH, N. F. F 84a. Argyrovernonia and Para- lychnophora: new names in the tribe Vernonieae (Asteraceae/Compositae). Taxon 33: 105-106. 1984c. positae: ee h.D. Dissertation. Univer- sity jr 9 wasaqa "Revisi n of Chresta and Pycnoceph- Sa. 2 (Compositae: Venice. cms Bot. 10: 459- : 5b. Revision of oo (Com- dip pn Vernonieae). Syst. ds 347-352. hu r. i new species of Ere hus (Vernonieae: Cerise tee) from Bra- zii sei Bot. 9: 85-95. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 PRANCE, G. T. 1982. A review of the pru evidences for Pleistocene climate changes in th Neotropics. Ann. Missouri Bot. Gard. 69: 594— 624 Rosinson, H. 1980. Notes on the Lychnophorine genera Chresta and Eremanthus (Vernonieae: As- teraceae). Phytologia 45: 89-96. F BOHLMANN & R. M. KiNG. 1980. Che- Asteraceae. III. Natural subdivisions of the Vernonieae. Phytologia 46: 21-436. SCHULTZ-BIPONTINUS, C. H. 1861. Me uni- florae. Jahresber. Pollichia 18-19: 0. . 1863. Lychnophora Martius un du nae be- nachbarte Gattungen. Jahresber. Pollichia 20-21: 39. SCHUMACHER, H. 1982. Rediscovery of Vanillos- rig arborea Baker (Compositae). Taxon 31: 801-802. THORNE, R. F. . Parasites and phytophages— pragmatic chemists? Symb. Bot. Upsal. 22: 200- 209. 1906. Vitae itineraque collectorum botani- In C. Martius (editor), Flora Brasiliensis 1(1): 1-154. URBAN, Z. 1973. The autoecious species of Puccinia on Vernonieae in North America. Acta Univ. Car- ol. Biol. 1971: 1-84. VELLOso, J. M. C. 1829. Pp. 1-352 in Florae Flu- minensis. Rio de Janeir 831. Florae fluminensis icones 8: tab. 150- 151. URBAN, I. . 1881. Florae fluminensis. Arq. Mus. Nac. Rio de Janeiro p 349-350. is ipa W., . L. MACHADO, J. A. RAB MURARI & W. Pus 1977. Eregoyazin ind er- o two new guaiangolides from Ereman- thus goyazensis. J. p Chem. 42: dia ZANONI, T. A. o Kuntze, botanist. I. Bi- Ripe ‘bibliography, dd travels. midi 32: 551-57 BIOSYSTEMATICS OF TETRAPLOID EUCHARIS (AMARYLLIDACEAE)! ALAN W. MEEROW? ABSTRACT Eucha rainforest understory from Guatemala to Bolivia. The tw ris is a genus of 16 species of petiolate-leaved, p Amaryllidaceae da lai to . bonpla northernmost species, ii and E. bouchei, are the only two tetraploid (2n = 92) species so far known in ack genus. E ucharis bonplandii entral series for both ‘tetraploid species. On the basis of 17 floral characters, the three varieties of E e tetraploid representatives of Eucharis exhibit a a wide degree of “karyotypic heteromorphism. , E ucharis bouchei var. dressleri i is an unstable tetraploid. ami no ff f polyploidy in some individuals. cm phenotypes of iach bouchei var. bouchei are quite variable and tl F var iety I Lines and a bouchei may b ° ° read tet traploid Asas desp x. The entry of Eucharis Into Central America was probably a “eka recent ons. It is he Colomb of heterozygosity may have been important in facilitating the migration of Eucharis across the Isthmus of Panama. The genus Eucharis Planchon & Linden di pa aged 2 family" Pancratioidinae sensu Traub (1957, 1963)] consists of 16 species ofi rare, petiolate- ms bulbous geophytes in- MAaAVIL Gliatemala to Bolivia (Meerow, 1986). Most of of pr imar y rainfor est from de Eucharis bouchei Woodson & Allen is a highly polymorphic tetraploid (2n = 92) complex of Central America (Fig. 1). The species is concen- trated in Panama (Fig. 2), but has also been re- corded from Costa Pies and Guatemala. Eucha- ris bouchei is th t species of Eucharis and the only one found north of the Darién Gap. It is also the most variable species in the genus, in characteristics that elsewhere justify specific delimitation. Patterns of variation in floral size and tube and limb habit form a complete mosaic throughout the range of E. pen showing little or no geographic consistenc In my recent mono mid of Eucharis (Mee- row, 1986), three varieties are recognized chiefly on the basis of staminal cup morphology (Fig. 3): E. bouchei var. bouchei, var. darienensis Mee- row, and var. dressleri Meerow. Variety bouchei, most common around El Valle de Antón in Coclé Province (Fig. 2), is recognized by its largely edentate staminal cup in which the trapezoidal free filament is not markedly con- stricted distally into a narrow subulate portion Figs. 1C, 3A, B). It is the most variable of the three varieties, both in flower size and staminal cup morphology. The staminal cup of variety ! I thank Robert Dressler, Mark Elliot, Mark Whitten, ce on Botanical Garden for providing living material of tetraploid Eucharis species. Bijan D ehgan p ovided the supportive environment w is work was accomplished. Charles Guy gave freely of his time xm E materials for the electrophoretic analyses. Pete applications. Brent Mishle r Goldblatt, George K. Rogers, and two anonymous reviewers provided Teila B of an earlier version of this paper. S Part of this work was supported by NSF America/World Wildlife Fund Fellowship in pis al B cu Gainesville, Florida 32611, U.S ANN. Missouni Bor. GARD. 74: 291—309. 1987. Doctoral pecu ion P R-8401208 and a Garden Club of 292 — A. Flowers. — _Fic GURE l. Eucharis bouchei. ouchei.—ii. Meerow 1125, FLAS. —iii. |. Mee- FLAS.—B. Tepals, variety bouchei. i, ii. Meerow 1125. —i. Outer tepal.— ii. Inner tepal. iii, iv. Meerow 1157.—iii. Outer tepal.—iv. Inner tepal.—C. Staminal cups, variety bouchei. —i. Meerow 1125.—ii. Meerow 1157. —D. Ovaries, variety bouchei, longitu- dinal section. —i. Meerow 1125.—ii. Meerow 1157. darienensis, found in Panamá and Darién prov- inces, is Dun pene or lobed (Fig. 3D). The tally into a narrow (<2 mm wide) subulate portion. These two va- rieties occur in close proximity in one location near Cerro Campana in Panama Province. The rare var. dresslerii (Fig. 1Ai), with its acutely toothed staminal cup (Fig. 3C) and non-trigo- nous Ovary, occurs close to populations of var. bouchei near El Valle Northwesternmost populations in Panama representing var. bouchei have the most derived androecial morphology (Fig. 3A, B) relative to ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 more southeastern populations (var. darienensis, Fig. 3D). The latter have staminal cups similar to the generalized morphology characteristic of Andean and Amazonian species of subg. Eucha- ris. This may indicate that general movement of E. bouchei in Central America has been away from the Colombian border. The only other naturally occurring tetraploid Eucharis species known is the rare Colombian E. bonplandii (Kunth) Traub. It is separated from E. bouchei by its slightly glaucous leaves (all oth- er Eucharis species have nonglaucous foliage), shorter petioles, and longer pedicels. Eucharis bonplandii is the northernmost species of Eu- charis subg. Eucharis in South America. Results of phenetic, karyotype, and prelimi- nary electrophoretic analyses of E. bouchei and E. bonplandii are presented in this paper. These data offer insight into the evolutionary history of Central American Eucharis and the origins of tetraploidy in the genus, and provide a basis for understanding the enormous degree of pheno- typic variation present within E. bouchei. The use of electrophoretic analyses of isozyme variation in plant systematics has been extensive in recent years. The subject has been reviewed by Gottlieb (1971, 1977, 1981a, 1981b, 1982, 1984) and Crawford (1983, 1985). Unlike many morphological characters, which may be influ- enced by a great deal of environmental or de- velopmental plasticity, the electrophoretic phe- lectrophoretic studies of tropical plants are few (Hamrick & Loveless, 1986; Heywood & Fleming, 1986; Sytsma & Schaal, 1985). Neither have plants of limited or rare distribution been widely investigated (Babbel & Selander, 1974). Members of Eucharis are tropical monocots exclusive to rainforest understory. They are rare and widely dispersed in the wild. Studies of iso- zyme variation of any plant group fitting any one of these characteristics are very limited. Isozyme analyses of polyploid taxa are also not abundant (Crawford, 1985; Soltis & Rieseberg, 1986). Thus, an attempt to explore isozyme variation in nat- ural polyploid Eucharis seemed a worthy avenue of investigation. MATERIALS AND METHODS PHENETIC ANALYSES Principal component and hierarchical cluster analyses of 20 herbarium specimens of Eucharis MEEROW — EUCHARIS 293 1987] '(p Ə[QË L 22s) Ayjeonaioydonosja pozAjeue suorje[ndod 0} 19721 suoreusrsop 191191 "(19sut *erquio[o/)) zpupjduoq *;] pue eureueg ut ;2u2noq Slupyony jo uonnquisqT “Z 380514 i l | I | BL êl 0 18 ü I | | | | — - L + =—— 1 0 Os s 0 \ J40|[SS04p iea [8U2DOQ "3 ($) a 2 sisueuoj18p «^» Jeuonoq '3 D Li O JƏYONOQ ‘180 jegyonoq syseyon”Z e Og zc d D N aga 3 A b L 8 8 3 3 oad A CE \ \ ae = SC ) AV 4 x "E ec \ ``. pee - ^ J X. e `. N N, J `. `< NO - b. a N Q5 `A GMUTÍ ) - e. ¢ be -— -—7 A. ' 2 Y ATNA ( š pi Nur 5 of | VWVNVd zgg3 | td83 | 294 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 3. Staminal cup variation in Eucharis Mieres y B. Variety bouchei. — A. Lewis et al. 2617 (MO). — B. Allen 120 (US).—C. Variety dressleri (Mee- row 1107, FLAS).—D. Variety darienensis (Gentry & Mori 13945, MO). e — 1) were conducted with CLUS- . 2.1 (University of St. Andrews, anne da on eai North Florida Regional Data Center (NERDC) system of the University of Florida. Three-dimensional scattergrams were constructed from P factor scores utilizing PC T, a program written by Bart Schutz- man at the University of Florida. The small number of OTUS (operational taxonomic units) underscores the relative rarity with which Eu- charis is encountered in the field. These 20 spec- imens represent the only specimens from which the full character set could be recorded. A num- ber of additional specimens examined were col- lected in fruit and therefore were useless for these analyses. Twenty-seven characters were used initially. The results suggested that some of these char- acters (e.g., all foliage characters, scape height, ovary length) were unreliable due to environ- mental plasticity, developmental variation, or specimen preparation. Although living material provides additional characters of potential utility (e.g., leaf surface texture, pigmentation pattern of the staminal cup), the inability to consistently determine these characters in dried specimens precluded their inclusion. Where any two char- acters were highly correlated (more than 80% correlation), which can result in data redundancy (Sneath & Sokal, 1973), one of the two was re- moved from the data matrix. In the final anal- yses, 17 floral characters (Tables 2, 3) were se- lected as the basic data set, of which 14 were continuous, quantitative Characters. The re- maining tł e characters were treated by assigning a über cd value for each Sartor state. Since CLUSTAN would treat these values as continuous, the character states were num- bered in a progressive transformation series, such that any two successive numbers would reflect putative character state relationship. These Hansiormaton series were constructed by study of d trends in the genus E ucharis and by comparative study with closely related genera of Amaryllidaceae. Raw data were standardized using the “z-score” method (Sneath & Sokal, 1973) by which initial values for each character were replaced by stan- TABLE 1. Operational taxonomic units sa for multivariate analysis of the Eucharis bouchei complex. CR = Costa Rica, G = Guatemala, P = Pan No Collection and Herbarium Variety Origin l Alston 8727 (BM) dressleri P, Coclé 2 Meerow 1107 (FLAS) dressleri P, Coclé 3 Wendland 207 (GOET) darienensis G 4 Sullivan 718 (M darienensis P, Darién 5 Folsom 4402 (MO) darienensis P, Darién 6 Folsom et al. 6582 (MO) ouchei P, Panamá 7 Allen 5347 (US) bouchei CR 8 Kirkbride & Hayden 305 (MO chei P, Panamá 9 Witherspoon & Witherspoon 8372 (MO) darienensis P, Panamá 10 Duke & Elias 3661 (GH) darienensis P, Darién 11 Gentry & s 13945 (MO) darienensis P, Darién 12 Stern et al. 49 H) darienensis P, Darién 13 Skutch 1585 n ch 14 Seibert 466 (MO) bouchei P, Coclé 15 Lewis 2617 (MO) bouchei P, Coclé 16 Witherspoon & Witherspoon 8736 bouchei P, Coclé 17 Allen 1228 (GH) bouchei P, Coclé 18 Mori & Kallunki 2014 (AAU) bouchei P, Colón 19 Mori et al. 6586 (AAU) bouchei P, Colón 20 Meerow 1158 (FLAS) bouchei P, Colón 1987] dard deviations from the mean value for that character. A distance matrix was then calculated using squared euclidean distance (Cormack, 1971). In addition to PCA, cluster analysis using average linkage (unweighted pair group method (UPGMA) of Sneath & Sokal, 1973) was also applied to the 20 OTUs as a further test of phe- netic relationship. CHROMOSOME CYTOLOGY Root tips were collected from living collec- tions, pretreated for 2-3 hours at room temper- ature in 10 ppm solution of o-isopropyl-N-phen- ylcarbamate (Storey & Mann, 1967), rinsed in distilled water, fixed in 3: 1 mixture of 95% EtOH and chloroform at 18°C for 24 hours, then stored after fixation in 70% EtOH at 18°C. Root tips were hydrolyzed in 1 N HCI at 50°C for 2-3 minutes, squashed, and stained with iron aceto- tuine, Only Min iin slides - were made. Metap d on a Nikon LLabenhot EA with AFX- II camera attachment; haploid idiograms were constructed from photomicrographs. As absolute chromosome length can vary ap- preciably from cell to cell due to differential ef- fects of pretreatment (Tjio & Hagberg, 1951; Schlarbaum & Tsuchiya, 1984), relative length based on a value of 100 for the haploid comple- ment was used to designate size class. Relative size igiene are based on 30% or ' greater corre- lations between al d from Battagla, 1955) and relative leneth (RL) of mi- totic metaphase preparations of various species of Eucharis, Eucrosia, Phaedranassa, and other Amaryllidaceae with 27 — 46, all of which have similar relative length ranges: large, RL = 7.0 ^s gangs length (AL): > 10 um]; moderately ane: 5.0-7.0 (AL: 7-10 um); medium, Ii. (AL: 5-7 um); small, RL < 3.5 (AL: 2- 5 um). For tetraploid karyotypes, diploid RL val- ues were halved to assign size class. Chromo- some morphology, modified from Battaglia (1955), is defined as follows: metacentric, Arm Ratio (AR; long arm/short arm) — 1 0-1.10; near-metacentric, AR = 1.10—1.50; submetacen- tric, AR = 1.50-3.00; subtelocentric, AR = > 3.00. ELECTROPHORETIC ANALYSES Population selection and sample size. Five populations were included in these analyses (Fig. 2: Table 4), representing all living collections of MEEROW — EUCHARIS 295 E2. Characters used for multivariate analysis of E ned species Flower number Limb spread (mm) Length of free filament (mm) Width of free filament (mm) Width of tube at throat (mm) Length of outer tepal (mm) Length of inner tepal (mm) . Width of outer tepal (mm) . Width of inner tepal (mm) . Staminal cup length (mm) LM — E E eel ROOIOTIO9u9tuPtwmr 3: Irregularly toothed 4: Quadrate 5: Lobe 6: Edentate . Cleft of staminal cup: — un 1: <% length of cup 2: 14—!⁄ length of cup 3: Vi—'^ length of cup 4: > length of cup . Relative length of teeth: 0: Edentate « length of filament 2: ^ length of filament 3: = length of filament 4: > length of filament . No. ovules per locule _ o — -l E. bouchei and E. bonplandii in cultivation at the University of Florida. These included three populations of E. bouchei var. bouchei, one from El Valle de Antón in Coclé Province of Panama, and one each from Cerro Brujo and Río Iguanita, respectively, in Colón Province; and one popu- ation of var. dressleri, also from El Valle. The fifth population represented E. bonplandii, a rare species from Colombia, also tetraploid. Sample size was one or three for each population (Table Unfortunately, the small sample size does not allow statistically significant exploration of ge- netic variation among these populations. Most electrophoretic studies in plant systematics have involved taxa of characteristically large popu- lation size in nature (see Crawford, 1983; Gott- lieb, 1981a). Populations of Eucharis, however, are characteristically small. Many species of Eu- 296 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 TABLE 3. Data matrix for PCA and cluster analyses of the Eucharis bouchei complex. Character OTU 1 2 3 4 5 6 7 8 9 10 ll 12 13 14 15 16 17 1 05 43 3.5 18 2.5 330 06.5 262 25.8 05.5 080 094 095 4 2 1 3 2 06 53 42 148 50 410 10.5 320 280 100 135 160 11.45 3 2 2 3 3 06 42 3.5 18 3.7 360 06.5 25.0 230 09.5 127 11.0 120 3 2 1 2 4 04 46 3.1 20 46 25.5 09.5 20.88 200 080 11.1 060 132 6 2 1| 2 5 05 43 2.5 22 43 2363 071 218 210 105 15.5 070 145 6 3 0 2 6 06 55 36 28 352 460 122 314 2300 138 152 106 174 6 3 0 3 7 06 69 35 28 S2 440 12.0 35.0 31.5 155 175 11.0 182 6 3 0 2 8 04 45 28 2.5 49 31.0 10.5 277 255 11.4 145 100 160 5 4 0 2 9 04 38 30 20 43 330 100 200 190 09.5 125 092 140 3 3 1 2 10 05 42 21 17 42 271 06.0 238 220 100 120 090 128 2 2 2 2 11 06 55 20 20 45 43.4 12.0 27.0 250 102 165 140 182 3 2 2 4 12 05 38 18 L8 5.0 30.0 09.8 220 21.0 11.5 150 090 123 2 3 2 2 13 04 41 50 51 51 35.0 10.5 242 21.1 09.8 145 120 154 4 3 0 2 ld 05 45 5.3 4.3 5.3 36.0 08.7 280 260 120 140 110 140 4 3 02 15 07 50 69 5.0 69 43.7 10.8 268 25.0 140 146 167 160 4 3 0 2 16 05 49 45 3.5 45 43.0 10.6 21.0 200 09.0 11.0 080 125 4 3 0 2 17 03 50 65 4.7 65 45.0 120 23.5 220 11.5 15.5 128 145 4 3 02 18 05 34 59 3.7 3.7 34.0 08.3 180 160 09.0 11.5 11.6 11.5 4 3 0 2 19 05 35 37 40 40 33.0 07.0 21.6 19.5 080 100 100 124 4 2 0 2 20 05 40 5.3 38 3.8 40.0 080 260 240 112 137 11.8 097 4 3 0 2 charis frequently occur as single, widely dis- for sample sizes as low as one. Nei stressed, how- persed clumps (Meerow, 1986). Herbarium spec- ever, that with a limited population sample, a imens of Eucharis regularly includ tati large number of loci must be analyzed. The num- indicating the rarity of the plants encountered. ber of enzyme systems assayed in the present Yet, if most Eucharis are primarily visited by study is not sufficient for this purpose. Instead, trap-lining insects (sensu Janzen, 1971) flyinglong individuals were scored for the presence or ab- distances, as may be the case (Meerow, 1986), sence of putatively identical bands (Table 5). population size from the perspective of potential These scores were used to generate distance coef- gene exchange may in fact be greater than oth- ficients by the unweighted pair group method erwise expected from known population densi- (Sneath & Sokal, 1973). Bands showing the same ties. Few workers have addressed the problem mobility, as determined by their position in the of how to apply electrophoretic data to rare plants gel, were considered to be identical. The resulting of characteristically small population size. Nei dendrogram could then be compared with the (1978) presented modified formulas forunbiased results of phenetic analysis of morphological genetic identities and distances that could be used variation, as well as with the data derived from TABLE 4. Eucharis bouchei and E. bonplandii populations examined electrophoretically. Desig- Taxon nation N Collection Information Voucher? Eucharis bouchei var. EBD 3 Panama, Coclé, El Valle de Antón Meerow 1107 dressleri E. bouchei var. bouchei EBBI 3 Panama, Coclé, El Valle de Antón Meerow 1125 E. bouchei var. bouchei EBB2 l Panama, Colón, Río Guanche, Cerro Meerow 1157 Brujo E. bouchei var. bouchei EBB3 l Panama, Colón, Rio Iguanita Meerow E. bonplandii EBN l Colombia, Cundinamarca, vicinity of | Bauml i BUMP Bogotá * All vouchers deposited at FLAS unless otherwise indicated. 1987] MEEROW — EUCHARIS 297 TABLE 5. Presence-absence data matrix for electrophoretic analysis of tetraploid Eucharis. Refer to Table 4 for population designations, Figure 14 for bands. (*) — used only for cladistic analysis. Blank space indicates le data unavailable. Band + wr Population l "19 — — — N — w — + — wa — nN — N 10 *18 ooco--ooooc|u oo--—-—-ooo|o Or Ke Omer eee co—-—-o-—-—--—o|o Ore ee eH ooo] A e@ — — — — ° ° ° ° | % tr! = ux — — O — — O — — = ooooorocoo- —ooooooococ!|o —oooooooo —oooooooo —oooooooo —o—-oco---- —o-oo0---- eee — HK OOO —o-oo-ooo oO-oc--o--- —o Om oe = P= comparative chromosome morphology. A sim- ilar method for analyzing isozyme data was used by Ashton et al. (1984) for Shorea (Dipterocar- paceae), and Chou et al. (1986) for several genera of bambusoid grasses. The data matrix was ad- ditionally subjected to cladistic analysis using PAUP by David Swofford (Illinois Natural His- tory Survey). The “Wagner method” of simple parsimony (Farris, 1970; Kluge & Farris, 1969) was applied in constructing the cladogram, and E. bonplandii was designated as the outgroup for polarization of character states. Of course, without the benefits of formal ge- netic analysis, there is no guarantee of genetic omology between any two bands of seemingly identical mobility. Future studies may allow more precise analysis of genetic variation within and among populations of tetraploid E ucharis. Isozyme ext Crude ex- tracts for isozyme electrophoresis were prepared by grinding ten 5 mm diameter leaf discs in 1 ml of extraction buffer [100 mM Tris-HCl, 10 mM DTT, 20% glycerol, and 1 mM PMSF ad- justed to pH 6.8 (Hames & Rickwood, 1981)]. Extracts were centrifuged twice, for ten minutes and two minutes, and the supernatant was de- canted by pipette after cach inde. ise: BIO-RAD Protean I exlvacrsilunide gel ee Gel rec- ipes were adopted from Hames & Rickwood (1981). Running gels were 0.75 mm thick and 7.5% acrylamide (10 ml 30% acrylamide-bis ac- rylamide, 10 ml 1.5 Tris-HCl at pH 8.8, 19.85 ml H,O, 100 41 10% ammonium persulfate, and 15 wl TEMED). A 2.5% acrylamide stacking gel (1 ml 3096 acrylamide-bis acrylamide, 1.92 ml 0.5 M Tris-HCl at pH 6.8, 9 ml H,O, 20 ul ammonium persulfate, and 7.5 ul TEMED) was employed. Running buffer was 25 mM Tris-gly- cine at pH 8.3 (Hames & Rickwood, 1981). A 20 ul sample of the supernatant was loaded into each stacking gel column. Gels were electro- phoresed at a constant current of 50 mA until a blue indicator line (40 ul of bromophenol blue added to cathodal buffer) migrated off the anodal end of the gel, generally four to five hours Five enzyme systems were assayed: aspartate amino-transferase (AAT), glutathione reductase (GSSGR), malate dehydrogenase (MDH), phos- phoglucoisomerase (PGI), and shikimate dehy- drogenase DH). Staining recipes of Vallejos (1983) were followed for AAT, PGI, and SKDH. The staining system for MDH was that of Shaw & Prasad (1970), and that of Kaplan (1968) was used for GSSGR. Resolution of additional enzyme systems (ga- lactose dehydrogenase, glutamate dehydroge- nase, hexokinase, and isocitrate dehydrogenase) using the same buffer system were unsuccessful. Extracts of Eucharis leaf tissue are characteris- tically mucilaginous, which may impede electro- phoretic separation or contribute to degradation of some enzymes after extraction. Also, cath- odally migrating isozymes cannot be resolved in the same vertical acrylamide gel as anodally mi- grating isozymes. RESULTS PHENETIC ANALYSES Principal Component Analysis (Figs. 4, 5; Ta- ble 6). Cumulative variance of 71.996 across 20 OTUs was resolved in the first three principal components (PCs). Characters 5 (stamen width), 298 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Eucharis bouchel @ var. bouche Æ ver. dressieri X var. darienensis FIGURE 4. PCA scattergram based on variance across 17 floral characters in 20 OTUs representing Eucharis bouchei 6 (tube length), 9 (inner tepal length), and 14 (toothing) contributed the greatest magnitude of variance to PC1, especially character 6. PC2 is largely a measure of outer tepal length (character 8), inner tepal width (11), staminal cup widt (13), and toothing (14). Characters 1, 7, and 14 also substantially contributed to the variance re- flected in PC2. Characters 2 (limb spread), 3 (length of free filament), 13 (staminal cup width), and 15 (toothing) v of variance in PC3. e three varieties of E. bouchei do not clearly resolve into three phenetic groups in Figure 4. Although var. bouchei shows a tendency to as- semble along PC2 (21.196 total variance), this variety is still widely distributed along PCI (38.7% total variance). One OTU each of var. VCIC the most FIGUR across 17 flor Eucharis bouchei, with PC2&3 emphasized. PCA s scattergram based on variance l characters in 20 OTUs representing darienensis (no. 1 1) and var. dressleri (no. 2) form an outlying group, as do 7, and 15 of var. bouchei. Variety darienensis shows a mea- sure of phenetic congruence, but intergrades with var. bouchei. If the scattergram for the E. bouchei complex is rotated so that PC2 and PC3 are visually ac- centuated (Fig. 5), grouping of OTUs becomes largely a measure of androecial Yonne. In this scattergram, the three clearly, particularly var. bouchei. Variety dari- enensis, however, still intergrades with several OTUS of var. bouchei, but one of these OTUs (8) was collected from Cerro Campana in Pan- amá Province, an area of sympatry between these two varieties. The third (no. 7) is a Costa Rican collection. Cluster analysis (Fig. 6). Two major clusters are resolved in the UPMGA dendrogram, each fairly heterogeneous. The first clusters at a dis- tance coefficient (DC) of 1.356. An outlying OTU (one of two representing var. dressleri) fuses with this cluster at DC 1.921. Within this first cluster, two subgroups emerge at DCs 1.207 and 1.213, respectively. The former is made up en- tirely of OTUs representing var. darienensis. The second represents var. bouchei, with the single exception of OTU 5 (var. darienensis). OTU 5 forms together with OTU 8 (var. bouchei) an outlying cluster to this second subgroup. The second major cluster is formed at a DC of 2.502, near where all clusters finally merge (DC 2.779). This smaller cluster is more hetero- d more 1987] TABLE 6. First three principal components for multivariate analysis of the Eucharis bouchei complex. Component Number Character Number l 2 3 l 0.151 0.317 0.120 2 —0.265 —0.189 0.340 3 —0.244 0.112 —0.440 4 —0.197 —0.141 —0.147 5 —0.375 0.129 0.059 6 —0.615 0.099 0.133 7 0.032 —0.302 —0.116 8 —0.100 —0.336 — 0.106 9 — 0.404 0.069 — 0.061 10 0.017 0.168 —0.239 ll —0.030 0.386 0.137 12 0.156 —0.062 —0.045 13 0.013 —0.353 —0.524 14 —0.232 —0.381 0.216 15 —0.184 0.322 —0.436 16 — 0.007 0.056 —0.121 17 — 0.046 —0.195 0.020 Percent of Variance 38.71 21.12 12.03 geneous than the first, but four OTUs of var. bouchei cluster at a DC of 2.112. As in PCA, OTUs 2 and 11 (var. dressleri and darienensis respectively) form a phenetic group. KARYOTYPE ANALYSIS Tetraploidy in Eucharis is known so far to characterize only E. bonplandii and E. bouchei. Karyotypically, the tetraploid Eucharis species are strongly heteromorphic (Figs. 7-12; Table 7). Karyotypes of two geographically isolated and morphologically distinct populations of E. bouchei var. bouchei, from Coclé and Colón provinces of Panama (Figs. 9, 10, 12A, B; Table 7) are quite different. The second largest chro- mosome pair is submetacentric in the Cerro Bru- jo (Colón) population of E. bouchei (Figs. 10, 12B) and also in E. bonplandii (Figs. 11, 12C). Eucharis bouchei var. dressleri is an unstable tet- raploid (Figs. 7, 8, 12D). Fifty percent of all root cells from which metaphase counts were ob- tained had 46 chromosomes. ELECTROPHORETIC ANALYSES Of the five enzyme systems assayed, only SKDH was monomorphic across all populations of E. bouchei and E. bonplandii. Only polymor- MEEROW — EUCHARIS 499 2.635 4 2.108 + 1.580 + DISTANCE 1.317 + 1.053 + -4 0.789 + Ë | " 9 na u i u n 16 ou M, NU ) 1 ", * x | 0 #* x e FicunE 6. Cluster analysis dendrogram based on variance across 17 floral characters in 20 OTUs rep- resenting Eucharis bouchei. Refer to Table 1 for iden- tification of OTUs. u phic loci are discussed below and diagrammed in Figure 14. AAT (Figs. 13A, 14). Two well-separated re- gions of activity were resolved for AAT, one rap- idly migrating anodally (AAT-1) and the other (AAT-2) considerably slower; these probably represent two different loci of this dimeric en- zyme. Electromorphs at both loci were consid- erably more complex than in diploid species of Eucharis (Meerow, 1986). Two putative alleles are inferred from the phenotypes of AAT-1 in the E. bouchei complex. Each “allele” of AAT-1 in all Eucharis characteristically resolves as two very closely spaced bands. This may be the result of breakdown products forming after extraction (see Fig. 1 in Shields et al., 1983). Electromorphs of pollen of diploid (2n = 46) Eucharis (Meerow, unpubl.) also showed this banding pattern. Were each component band of the doublet a distinct allele, pollen would be expected to show only one of the two (Gottlieb, 1982, 1984). Alterna- tively, if the high diploid chromosome number derived (Meerow, 1987a), the doublet banding pattern may reflect duplication of the genome and would show up in pollen. 300 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FiGURES 7-11. tetraploid Eucharis species. 7, 8. E. bouchei Root-tip ce fi id cell. se i cell.—9. E. deren var. bouchei from Colón Province in Pana- . E. bonplandii. Two small chromosomes var. dressleri. i Diplo ma. — 10. ei var. bouchei from Coclé aped in Panama.— 11 are outside rad figure frame. All scales = 10 u Band a was the most common “allele” of AAT-1, found in all individuals analyzed except for two putative homozygotes for "allele" c (the o Brujo population and one individual of the El Valle population of var. bouchei). Variety dressleri and E. bonplandii are homozygous for "allele" a. The Río Iguanito individual of var. bouchei is homozygous for “allele” b, while two individuals of the El Valle population show a putatively heterozygous phenotype with appar- ent heterodimerization Four E. bouchei individuals resolved a four- banded electromorph at the putative AAT-2 lo- cus. Bands f and g were found only in E. bon- 1987] MEEROW — EUCHARIS 5 units 1234526 7 8 9 101 12 134 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 RELATIVE LENGTH ITTE] ) OTE Le D FiGurE 12. Haploid idiograms of tetraploid Eucharis karyotypes.—A. E. bouchei var. bouchei from Coclé Province in Panama.—B. E. bouc. diploid cell.— D. E. bonplandii. plandii. Only two bands were observed in var. dressleri (an unstable tetraploid), representing “alleles” a and b or b and c, and one individual of var. bouchei from EI Valle (*alleles" c and d). All other individuals of E. bouchei resolved a four-banded electromorph for AAT-2. Band e was found in one of the three individuals of var. bouchei from El Valle. As all diploid species of Eucharis species resolve only a two-banded elec- tromorph for this isozyme (Meerow, 1986), it was inferred that the proliferation of bands with- in E. bouchei represented the additive effects of tetraploidy (Crawford, 1983, 1985; Gottlieb, MDH (Figs. 13B, 14). Malate dehydrogenase characteristically forms complex banding pat- terns that require genetic analysis to decipher (Kirkpatrick et al., 1985; Torres & Mau-Lasto- vicka, 1982). Consequently, no attempt is made to infer genotypes in any detail from the banding pattern. However, the phenotype of the most an- odal bands in E. bonplandii suggests the presence of two alleles and their heterodimer [pollen of hei var. bouchei from Colón Province in Panama. —C. E. bouchei var. dressleri, this species resolved only a single band at this locus in a repetitive run (Meerow, unpubl.), sup- porting this interpretation]. The intensity of the bands in the two most cathodal regions of several individuals suggests dosage effects in putative homozygotes (two individuals of EBBI, and Two bands were observed in the single locus resolved for GSSGR, all in- dividuals manifesting one or the other PGI (Fig. 14). Onlyasingle region of activity was resolved for PGI. Two bands were observed, but the more anodal one was found only in the putative heterozygotes (two individuals of E. bouchei var. dressleri, and one of var. bouchei from EI Valle), and in E. bonplandii. Isozyme relationships. The nine individuals for which electrophoretic phenotypes were re- solved were coded for presence or absence of the numbered bands in Figure 14, creating a data set of 19 characters. Phenotypes for PGI were not included in the cluster analysis, as data were not available for all nine individuals. The resulting 302 TABLE 7. otherwise stated. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Karyotype data, Eucharis bouchei and E. bonplandii. All vouchers deposited at FLAS unless Chromo- Chromo- so PRI jii I Chromosome Size^ Chromosome‘ Taxon, Voucher, m Range (relative Groups Morphology & Fig. No Number (um) ngt L ML M S L M M S Eucharis bouchei 46,92 2.4-11.9 2.0-9.7 4 10 14 18 m: 4 2 dressleris nm: 6 10 (Meerow 1107) sm: 4 6 8 Figs. 7, 8, 1 st: 6 E. bouchei v 92 1.6-15.5 0.5-5.3 14 16 14 48 m: 2 bouchei Tom nm: 2 2 18 1157) Figs. 9 sm: 10 6 6 28 12B st: 2 10 6 E. bouchei var. 92 2.0-10.4 0.8-4.5 10 18 l6 48 m: 2 2 4 6 bouchei (Meerow nm: 2 6 28 1125) Figs. 10, sm: 6 8 4 14 12A st: 8 2 E. bonplandii 92 1.9-9.7 1.0-4.9 14 12 24 42 m: 6 (Bauml 686, nm: 6 6 18 HUNT) Figs. 11, sm 2 4 16 18 12D st 6 8 2 * Based on a value of 100 for the ig iio UP ges , ML = moderately long, = , S = e L = lon mall. S “m= metacentric, nm = ki asin dpi sm — submetacentric, st = subtelocentric. * Diploid cell analyzed. UPGMA dendrogram (Fig. 15) and cladogram (Fig. 16) illustrate the isozyme relationships among these individual plants. As might be ex- pected, both trees are similar in topology. All three individuals of E. bouchei var. dressleri (EBD) cluster at a distance coefficient (DC) of only 0.118, indicating their close isozyme rela- tionship. The El Valle (Coclé Province) popu- lation of var. bouchei (EBB1) is rather diverse in its patterns of isozyme variation. Two individ- uals are similar, clustering at a DC of 0.118. This cluster then fuses with the Río Iguanita (Colón Province) pasa of var. bouchei (EBB3) ata DC of 0. . The remaining individual of El Valle var. be EBBI shows greater isozyme relationship to Cerro Brujo (Colón Province) var. bouchei EBB2 than other convarietal individuals from El Valle. This heterogeneous cluster then fuses with var. dressleri (EBD) at a DC of 0.431, followed by the other El Valle individuals (EBB1) and the single Río Iquanita (EBB3) individual (DC 0.443). The single individual representing E. bonplandii (EBN) remains a distant outlyer from all plants of E. bouchei, joining the latter species at a DC of 0.625 The cladogram based on isozyme data was 27 steps long with a consistency index (Kluge & Farris, 1969; CI — total length minus homopla- sies, divided by total length) of 0.704. The clado- gram supports a monophyletic origin of E. bouchei var. dressleri (EBD) from var. bouchei (EBBI) in Coclé Province, but suggests that var. bouchei may be polyphyletic. The Río Iguanita (Colón Province, EBB3) individual of var. bouchei forms a monophyletic group with two individ- uals of El Valle (Coclé province, EBBI) var. bouchei, while the other Colón individual (Cerro Brujo, EBB2) forms a monophyletic group with the third El Valle individual. DISCUSSION The Central American £. bouchei complex does not resolve oe discrete penenie groups. Stam- inal c logy, however te va- rieties to a fair degree (Fig. 5). Floral s size char- acters in this group (Figs. 4, 6) do not succeed as well in resolving phenetic groups. Chromosome number is very stable in Eucha- ris, and polyploidy is infrequent. The origins of the polyploids (i.e., whether auto- or alloploid) EBD EBN EBD EBB) EBD EBB-3 EBB2 EBB) EBB ! 13. Representative gels for borus vois i complex - c — - E re e ru putative Mh discussed in text. Refer to Table 4 for population designations. are inconclusive (attempts to secure meiotic fig- ures have been unsuccessful), but the high levels of morphological diversity in E. bouchei might suggest that they are alloploids. Differences in chromosome morphology among the popula- tions of E. bouchei examined (Table 7) suggest that structural changes in the chromosomes may have been important in interpopulational diver- gence. Eucharis bouchei var. dressleri is an un- stable tetraploid. Somatic cells of b» E tips e both t nte PH iN J TIJ Vu hav Snoad (1955) a karyotype MA a: in as polyploid counts were observed in the cells of the latter species. Polyploid species of Eucharis do not show any marked effects of increased chromosome number beyond an increase in size of root cells and sto- mata, and slight thickening of the leaf blades. Eucharis bonplandii, in addition, develops a glaucous bloom on the leaves in strong light, a novel characteristic for the genus. MEEROW — EUCHARIS | |! EBD EBD EBD EBB-1 EBD EBD EBD — ló — 17 EBB-1 FBB-1 EBB-I EBB-2 EBB-3 EBN EBD EBD EBD i complex. Anoda a à — aspartate transferase, MDH = malate dehydrogenase, GSSGR = glutathione reductase, PGI = phosphoglucoisomerase. Eucharis bouchei var. bouchei, as presently Sea Meera, 1986), is diverse in floral ] nd pat- terns of i isozyme polymorphisms. Xon re- lationships based on the isozyme data indicate that this polymorphic variety may even be poly- gically EBB3 is similar to a O ouchei (Fig. 1Aiii), they are not discontinuous 0.7004 0.600— DISTANCE > [^] ° 1 0.2004 0.100 0.000 EBD EBD E8D FiGURE 15. designations. enough to warrant a clear differentiation of a fourth variety in the species. The two individuals representing different populations from Colón Province each have closer isozyme relationship to Coclé var. bouchei than they have to eac other, while the Coclé population itself appears isozymically diverse. Cladistic relationships based on this data imply that Coclé populations of var. bouchei may be of heterogeneous ancestry. The cladogram (Fig. 16) also points to Colón Prov- ince as the likely origin of those ancestors. Di- vergence between and among the Coclé popu- lations and those in Colón Province, presumably mediated by geographic isolation, may thus an s process. the basis of staminal cup morphology, I i petens that Eucharis bouchei has been ). The Cerro Brujo pop- ulation of E. bouchei var. bouchei may represent an intermediate point in the divergence of a new geographical race of E. bouchei. The best test of this hypothesis would be the results of isozyme analysis of E. bouchei var. darienensis, the one variety for which material is not presently avail- able. Variety darienensis occurs closer to the Co- lombian border than any other population of E. bouchei and has the most generalized staminal cup morphology relative to Eucharis as a whole. If my hypothesis is correct, var. darienensis ANNALS OF THE MISSOURI BOTANICAL GARDEN EBB1 UPGMA dendrogram oftetraploid Eucharis isozyme phenotypes. Refer to Table 4 for population [VoL. 74 EBB2 EBBI EBBI EBB3 EBN should have the lowest genetic identity with pop- ulations of either var. bouchei or var. dressleri from Coclé Province, and higher identity with populations of var. bouchei from Colón Prov- ince. Colón populations of E. bouchei are geograph- ically intermediate between most populations of var. darienensis and the Coclé populations of var. bouchei (Meerow, 1986). The two varieties come into close proximity in the Cerro Campana area in Panamá Province. Colón populations may therefore also be genetically intermediate be- tween the two varieties. Segregating genotypes in such a case could produce populations exhibiting a mosaic of varying genetic identity, some close to Coclé var. bouchei, others perhaps closer to var. darienensis. Further testing of this hypoth- the first step in sympatric speciation. This variety shows greatest isozyme relationship with certain individuals of the Coclé population of var. bouchei. In this regard, the presence of a second band for PGI in certain individuals of var. dres- sleri and El Valle var. bouchei may be significant (Fig. 14), but a larger number of individuals must be assayed to confirm this observation. This va- 1987] EBD MEEROW — EUCHARIS EBBI EBB3 EBBI EBB2 —: reversal æ : homoplasy 16. Cladogram based on isozyme Ee of tetraploid Eucharis. Refer to Table 4 for population P aypuy asi Broken line indicates zero-length branch riety is also an unstable tetraploid. Fifty percent of all chromosome counts of root tip cell mitotic metaphase configurations have 2n = 46, the typ- ical diploid chromosome number in Eucharis. Variety dressleri lacks the additive banding pat- terns or heterozygote phenotypes observed in both loci of AAT in all other populations of E. bo uchei, a factor, perhaps, of this karyotypic in- Eun (CE et al., 1972), Nicotiana (Red- dy & Garber, 1971; Sheen, 1972; Smith et al., 1970), Triticum aestivum (Hart, 1970, 1979; Jaaska, 1978; Torres & Hart, 1976), and Steph- po (Gottlieb, 1973), and are usually in- rpreted as indicative of allopolyploid origins = ach 1983; Gottlieb, 1983; Soltis & Rie- seberg, 1986). Pollen stainability of var. dressleri is 10096 with Alexander's (1969) stain, suggesting that gamete formation is not impaired by the chromosome number instability. Nonetheless, I have not successfully crossed this variety with El Valle populations of var. bouchei The rare Colombian ietraploid E. bonplandii also lacks either heterozygote or additive band- ing patterns for AAT. This may indicate an au- topolyploid origin for this species (Crawford, 1985; Soltis & Rieseberg, 1986), or at least a lower degree of heterozygosity than in E. bouch- ei. The rarity of this species, in relative contrast to E. bouchei, fits Stebbins's (1980) model of the “unsuccessful” autopolyploid. The difficulty in obtaining successful meiotic figures from bulbs pletely inside the bulb) this question. Eucharis bonplandii exhibits large distance on the basis of isozyme phenotypes from all individuals of E. bouchei. The question of whether these two species represent a monophy- letic group on the basis of their tetraploid origin is not conclusive. Increased heterozygosity is an expected con- sequence of allopolyploidy (Crawford, 1983, 1985; Gottlieb, 198 1a; Soltis & Rieseberg, 1986). A certain degree of fixed heterozygosity would also be expected in an allopolyploid (Gottlieb, 198 1a; Soltis & Rieseberg, 1986), due to the pres- ence of two genomes in the allotetraploid. Al- though it is premature to assess the degree of heterozygosity present in tetraploid Eucharis, their morphological diversity and i dim poly- morphisms indicate that it may There is insufficient information on ne breed- ing system and pollination biology of Eucharis to support more than ad hoc hypotheses of the origin of most species in the genus. The char- acteristically small population sizes that are en- countered throughout its range may indicate that ounder effects (Mayr, 1954; Templeton, 1980a, 1980b) have played an important role in the movement of E. bouchei across the Isthmus of Panama, with subsequent isolation restricting gene flow between localized populations. The pu- tatively allotetraploid genotype and karyotypic polymorphy of ouchei would favor the “‘hy- brid recombination" type of "genetic transili- ence," a mode of speciation hypothesized by Templeton (1980a, 1980b). This is consistent with the distribution of E. bouchei, most populations of which are geographically isolated from each other (Meerow, 1986). Additional support comes from the morphological novelties expressed 306 within each geographical variety (or race, in the case of var. bouchei). Templeton’s model of spe- ciation requires a period of successful inbreeding after the initial founding event. Though Eucharis are primarily out-crossing, evidence from green- house studies indicates only partial self-incom- patibility (Meerow, 1986). At least one Ama- zonian species of subg. Eucharis [E. castelanaena (Baillon) Macbride] is autogamous (Meerow, 1986) EUCHARIS IN CENTRAL AMERICA Gentry (1982) suggested that two major op- portunities, widely spaced in time, existed for floristic interchange between Central and South America. The first, occurring during the Late Cretaceous, was limited to a series of volcanic islands (the proto-Antilles; Dengo, 1975; Lille- graven et al., 1979). The degree to which this island arc remained above water is unknown. At the beginning of the Tertiary, however, this link between the continents was disrupted as the pro- to-Antilles began a northward displacement. It was not until the late Tertiary that the second opportunity for floristic interchange began to co- alesce, as formation of the Central American trench and new volcanic activity gave rise to a new series of islands. These islands eventually (1982) concluded that only very well-established Cretaceous taxa would have been able to take advantage of the earlier connection via island- hopping. Entries into Central America dating from this earlier connection would be expected to show strong taxonomic differentiation in Cen- tral America. Gentry (1982) cited tribe Crescen- tieae of the Bignoniaceae as a putative example of early colonization of Central America by is- land-hopping, followed by taxonomic differen- tiation. On the contrary, any migration dating from the Pliocene or Pleistocene would not be expected to show much differentiation at the spe- cific or at the generic level. I have characterized the Eucharis bouchei complex as a semispecies complex of geographically isolated races or va- rieties not yet strongly differentiated. Patterns of isozyme variation, chromosome cytology, and morphological variation in this group suggest that entry of Eucharis into Central America was fairly recent. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 The species of subg. Eucharis geographically closest to E. bouchei is E. bonplandii, a rare species of central Colombia, and also tetraploid. It is inconclusive whether these two species rep- resent a monophyletic tetraploid group. None- theless, the congruence of phytogeography with chromosome number in these two species sug- gests that this may indeed be the case. It is tempt- ing to wonder if tetraploid Eucharis were at one time more common in northern Colombia, and if E. bu and E. bonplandii represent the remnant populations of a once more widespread, d tetraploid complex. Prance's (1982) most recent distribution of Pleistocene refugia based on phytogeographic patterns includes both a Río Magdalena refuge in northern Colombia (most collections of E. bonplandii are from the Rio Magdalena valley south of Prance's pro- posed refuge), and a Darién refi enensis is most common in the area ofthe Darién refuge and is putatively the least derived variety of the species. The absence of collections of Eu- charis subg. Eucharis from northern Colombia is something of a mystery but may indicate that extinction of intervening populations between E. bonplandii and E. bouchei was widespread in the recent geological past. CONCLUSIONS The Central American E. bouchei complex is a tetraploid, putatively alloploid, possibly highly 1981) still actively evolving. Discrete patterns of isozyme divergence have not yet solidified be- tween morphologically distinct and geographi- cally isolated populations of E. bouchei var. bouchei. Founder effects and geographic isola- tion probably were, and still are, important forces influencing the continued evolution of E. bouch- ei. In one case (E. bouchei var. dressleri) sym- patric divergence may be in process unprecedented degree of variation in E. bouchei is thus likely the result of two main fac- tors: (1) tetraploidy, accompanied or followed by structural rearrangement of chromosomes, and (2) a geologically recent colonization of Central America by this primarily northern Andean and Amazonian genus. The wide variation present in E. bouchei likely represents the segregating phe- notypes of a richly diverse genetic base. On the basis of known distributions, it appears that sub- stantial geographic barriers exist between groups 1987] of populations, probably restricting gene flow be- tween them. Left undisturbed, as is not the case in the Neotropics today, these aggregates could conceivably one day each justify specific recog- nition. The fact that E. bonplandii is the northernmost species of Eucharis subg. Eucharis in South America, and is also tetraploid, lends at least circumstantial credence to the hypothesis that E. bouchei and E. bonplandii diverged from a com- mon tetraploid ancestor. The rare occurrence of polyploidy in Eucharis strengthens this possi- bility. Stebbins (1985), in a recent review of poly- ploidy, found a correlation between high fre- quency of polyploidy and patchy g } (or ecological) distributions, coupled with the oc- currence of secondary contact between these dif- ferentiated populations. Levin (1983) discussed how chromosome doubling may “ ‘propel’ a pop- ulation into a new adaptive sphere.” Though E. bouchei does not exhibit any noticeably novel ecological adaptations, its success in colonizing the Isthmus of Panama may have been aided by its polyploid-related genetic diversity. The het- erogeneous isozyme patterns characteristic of El Valle var. bouchei may indicate either multiple ancestry for this population or that secondary contact has occurred between it and populations in Colón or Panama provinces to the east. Eucharis bouchei offers an excellent opportu- nity for detailed study of the evolution of a trop- ical rainforest organism. Future work should seek to quantify in greater detail the genetic variation present within and among populations of this actively evolving complex. Meiotic pairing fig- ures from dissection of bulbs may help confirm the nature of the polyploid origins of this species. Paleotropical genera of “‘infrafamily”’ Pancra- tioidinae characteristically have 2n — 22 or 20 chromosomes (Ponnamma, 1978; Zaman & Chakraborty, 1974), while almost all neotropical genera have 2n = 46 (Di Fulvio, 1973; Flory 1977; Meerow, 1987a, 1987b; Williams, 1981). The latter number is likely derived through frag- mentation or duplication of a single chromo- some, followed by doubling of the genome (Lakshmi, 1978; Sato, 1938). Increased hetero- zygosity may therefore have accompanied a tet- raploid origin of the neotropical tribes of the Pancratioidinae from an ancestor with 2n = 22 " e somatic number characteristic of Pancra- m L., the largest paleotropical genus of the PEL (Ponnamma, 1978)]. The high generic MEEROW — EUCHARIS 307 diversity of neotropical pancratioids (ca. 15 gen- era) in comparison with the paleotropical taxa (4 genera) itself may be partially a consequence of greater genetic variability. Comparative anal- ysis of isozyme phenotypes between paleotrop- ical and neotropical genera is planned and may provide insight into the evolution of the Pan- cratioidinae. LITERATURE CITED ALEXANDER, M. P. 1969. Differential staining of orted and non-aborted pollen. Stain Technol. 44: 117-122 ASHTON, P. S., YIK-YUEN & F. W. ROBERTSON. 1984, Electrophoretic and morphological com- , G. . K. SE : 1974. Genetic variability in edaphically restricted and wide- spread plant species. Evolution 28: 619—630. BATTAGLIA, E. 5 hromosome morphology and origi Caryologia 8: 178-187. CHERRY, J.P R. M. KATTERMAN & J. E. aedes 1972. nd catalases of species of the genus Gossypium. Theor. Appl. sl 42: 218-226. CHou, C. Y. H. HwANG & S. Y. HwANG. 1986. A Bond aspect of phylogenetic study of Bambusaceae in Taiwan. IV. The genera Arundi- naria, Pseudosasa, Semiarundinaria, Shibataea, Sinobambusa, and Yushania. Bot. Bull. Academia 31. 71. A sides of classification. J. Royal Statist. Soc. A, 134: 321-367. 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Gel electrophoresis: new ap- proach to the study of evolution. BioScience 21: 944, Genetic control of glutamate oxaloac- etate transaminase in the diploid plant Stepha- la exigua and its allotetraploid derivative. Biochem. Genet. 9: 97-107. —— . 1977. Electrophoretic evidence and plant sys- tematics. Ann. Missouri Bot. Gard. 64: 161-180. 1981a. Electrophoretic evidence and plant di errem Prog. Phytochem —46. 81b. Electrophoretic evidence and dun E Pp. 1-46 in L. Reinhold, J. Harborn & T. Swain (editors), Progress in aed Volume 7. Pergamon, New York. . 1982. Conservation and duplication of iso- zymes in plants. Science 216: 373-379. 1984. Isozyme evidence and problem solving in plant systematics. Pp. 342- in W. F. Grant (editor), Plant Biosystematics. Academic Press, Florida. . Plant — 2nd edition. Co- umbia Univ. Press, New D. Rickwoop. T . Gel Electro- phoresis of Proteins. A Practical Approach. IRL Press, Oxford HAMRICK, J. L. & M . D. LovELEss. 1986. Isozyme variation in tropical trees: procedures and prelim- inary results. Biotropica 18: 201-207. HART, G. E. 70. Evidence for pe raure y for alcohol dehydrogenase in ee xaploi at. Proc. Nat. Acad. Sci. USA 66: l 979. Evidence for a nue set of gluco- sephosphate isomerase sag genes in hexa- ploid wheat. Biochem. Genet. 17: 585-598. Heywoop, J. S. & T. H. FLEMING. oe Patterns of allozyme variation in three Costa Rican species of Piper. Biotropica 18: 208-213. . 19 DP-dependent aromatic alcohol 71. Euglossine dna as one: oo pollinators of tropical plants. Science 171: 203- 205 KAPLAN, J. C. 1968. Electrophoretic study of gluta- thione reductase in human erythrocytes and leu- cocytes. Science 217: 256-2 Pliocene closing of the 3 G E: z r pepo complex: C. pepo var. medullosa vs. C. tex- 299. J. S. FARRIS. 1969. Quantitative phy- letics and the evolution of anurans. Syst. Zool. 18: -32. LAKSHMI, N. 1978. Cytological studies in two allo- polyploid T7 of the genus Hymenocallis. Cy- tologia 43: 555 Levin, D. A. 1983. Pu E novelty in flow- ering plants. Amer. Nat. 122: 1-25. LILLEGRAVEN, J. A., M. J. Tapia & T. M. BROWN 1979. Paleogeography of the world of the Me- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 sozoic. Pp. 277-308 in J. A. Lillegraven, Z. Kie- lan-Jaworowska & W. A. Clemens (editors), Me- sozoic Mammals. Univ. California Press, Berke- ey. MARSHALL, L. G., S. D. WEBB, J. J. SEPKOSKI & D. 2 Raup. 1982. Mammalian evolution and the American interchange. Science 215: 1351- 1357. Mayr, E. 1954. Change of genetic environment and evolution. Pp. 157-180 in J. Huxley (editor), du) lution as a M Allen and Unwin, Lon w, A. W. 1986. A Monograph of Eucharis and C Mliohnuria (Amaryllidaceae) Ph.D. Dissertation. University of Florida, Gainesville. l hromosome cytology of Eucharis, C aliphruria and Urceolina (Amaryllidaceae). Amer. J. Bot. 74 (in press). 7b. A monograph of Eucrosia (Amaryl- lidaceae). Syst. Bot. 12 (in press). Nel, M. 1978. Estimation of average Biel and genetic distance from a small n dividuals. Genetics 89: 583-5 PONNAMMA, M. G. 1978. Studies o on bulbous mentals I. Karyomorphology of diploid a "ip loid taxa of Pancratium triflorum Roxb. Cytologi 43: 717-725. MEERO RANCE, G. T. 1982. A review of the phytogeographic evidences for Pleistocene climate changes in Neotropics. Ann. Missouri Bot. Gard. 69: 594- 624. Reppy, M. M. & E. D. GARBER. 1971. Genetic stud- ies on variant enzymes. III. Comparative electro- phoretic studies of esterases and peroxidases for species, hybrids and amphiploids in the genus Ni- cotiana. Bot. Gaz. 132: 158-166. 8. Karyotype evolution and phylogeny. IV. Karyotype in Amaryllidaceae with special ref- erence to SAT chromosomes. Cytologia 9: 203- SCLARBAUM, S. E. & T. TSUCHIYA. onomy and 1984. Cytotax- phylogeny in certain species of Tax- —54. 1970. Starch gel electro- phoresis of enzymes—a compilation of recipes. Bi 7-320. Iso ozymic evidence bearing on the rigin of Nicotiana tabacum. B union 26: 142- 154. SHIELDS, C. R., T. J. edis & C. W. SruBER. 1983. An outline of general resource needs and proce- dures for the iria sues separation of active enzymes from plant tissue. Pp. -468 in S. D. Tanksley & T. J. Orton (editors), Isozymes in Plant Genetics and Breeding, Part A. Elsevier Science Publ. B.V., Am app SMITH, H. H., D. HAMILL, E. WEAVER & K. THOMPSON. 1970. Multiple He isis forms of peroxidases and esterases among Nicotiana species and am- phiploids. Heredity 61: 203-212. SNEATH, P. H. A. & R. R. SOKAL. 1973. Numerical Taxonomy. W. H. Freeman and Co., San Fran- isco. SNOAD, B. 1955. Somatic instability of chromosome number in Hymenocallis calathinum. Heredity 9: 129- = Sorris, D. . H. RiEsEBERG. 1986. Autopoly- hiems in 7 menziesii (Saxifragaceae): ge- 1987] netic on from the enzyme electrophoresis. Amer. J. Bot. 73: 310-318. STEBBINS, G. L. 1980. Polyploidy in plants: unre- solved problems and prospects. Pp. 495-520 in W Lewis (editor), Polyploidy. Plenum Press, New York. 1985. Polyploidy, hybridization, and the in- vasion of new habitats. Ann. Missouri Bot. Gard. 72: 824- Storey, W. B. & J. D. MANN. 1967. Chromosome contraction by o-isopropyl-N-phenylcarbamate (IPC). Stain Technol. 42: 15-18. SYTSMA, K. J. & B. A. SCHAAL. 1985. Genetic vari- ation, differentiation, and evolution in a species complex of tropical shrubs based on isozymic data. e theory of speciation via the founder principle. iine 94: 1011-1138. Modes of speciation and inferences based on Pena distances. Evolution 34: 719- 729 Tyo, J. H. & A. HAGBERG. 1951. Cytological si on some X-ray mutants of barley. An. Estac. Aula Dei 2: 149-167. MEEROW — EUCHARIS 309 Torres, A. M. & G. E. HART. 1976. Developmental specificity and evolution of the acid phosphatase ey soq y Triticum aestivum and its progenitor species. Biochem. Genet. 14: 595—609. . MA -LASTOVICKA. 1982. Citrus isozymes. J. Heredity 73: 335-339. TRAUB, H. P. Classification of the arylli- 3: Am daceae: subfamilies, tribes, and genera. Pl. Life 1 76-81. 1963. Genera of the Amaryllidaceae. Amer- ican Plant Life Society, La Jolla VALLEJOS, C. E. 83. Enzyme staining activity. Pp. 469-516 in S. D. Tanksley & T. J. Orton (editors), Isozymes in Plant Genetics and Breeding, Part Elsevier Science hid B. V. , Amst erdam. WILLIAMS, M. D. nt for Par- amongaia Marie io Velarde. Pl. Life 37: 83- 89. ZAMAN, M. A. & B. N. CHAKRABORTY. 1974. Cyto- genetics of Amaryllidaceae: I. karyomorphology and meiotic behavior ae inversion heterozygote Bangladesh J. Bot 58. THE SHRUBBY GENTIAN GENUS MACROCARPAEA IN PANAMA KENNETH J. SYTSMA! ABSTRACT A single species of the lisianthoid genus Macrocarpaea had been known from Panama. Recent m The shrubby gentians of the Neotropics are some of the most conspicuous elements of higher elevation tropical forests. This complex of about 6 genera comprised Grisebach’s (1838) tribe Lisyantheae. These lisianthoid genera are noto- riously difficult to separate taxonomically with the consequence that they have often been treat- ed as synonyms of Lisianthius (or its orthograph- ic variant "Lisianthus"). The taxonomic diffi- culties are compounded by their remote montane habitat which makes them poorly collected. Only Lisianthius sensu stricto (Weaver, 1972a) and Macrocarpaea (Ewan, 1948) have been ade- quately monographed. Other genera of the tribe Lisyantheae are now being taxonomically re- vised in a multidisciplinary study (Maas et al., 1984; Maas, 1985). Macrocarpaea (Griseb.) Gilg is one of four lis- ianthoid genera in Panama (Elias & Robyns, 1975). Symbolanthus pulcherrimus, Irlbachia alata subsp. alata (formerly Chelonanthus ala- tus), and seven species of Lisianthius (Sytsma, 1987) also occur in Panama. These four genera can be separated in Panama by the following characteristics: la. Main stem terete; stigma capitate; old pla- centae visible as whitish bands along margins of mature capsules; pollen grains as monads . Lisianthius Main stem usually deii dri stigma bi- furcate; old placentae not visible on mature capsules; pollen grains as Min or tetrads. 6-10 cm long; corolla funnel- orm; more or less distinct, membrana- ceous; corona scalelike; pollen grains as tetrads Symbolanthus . Flowers to 5 cm long; corolla usually = N [^ as monads or tetra 3a. Leaves Bee "bracteoles never wallioides, M. anco: and M. subcaudata. Evi- dence is provided to merge the Costa Rican M. jer ll into M. macrophylla. leafy; calyx z mm long; pollen grains as tetra Irlbachia 3b. Leaves petiolate: bracteoles. leafy; calyx 6-20 mm long; pollen grains as monads u. Macrocarpaea The genus Macrocarpaea is centered in the Andes of northern South America but extends into the Amazon Basin and the Guayana High- land. Of the approximately 30—50 species in the genus, only eight are known from Central Amer- ica and adjacent West Indies. Macrocarpaea domingensis Urban & E. Ekman and M. tham- noides (Griseb.) Gilg are restricted to the Do- minican Republic and Jamaica, respectively. Cuba has two endemic species, M. pinetorum Alain and M. pauciflora Alain. Three species have been described from Costa Rica (Weaver, 1972b): m Nilsson, has been known from Panama near the border of Costa Rica (Elias & Robyns, 1975). The type specimen had represented the only col- lection for this apparently epiphytic shrub prior to 1975 Recent explorations of cloud forest habitats in central regions of the Cordillera Talamanca, a ridge extending from the border of Costa Rica to near Panama City, and in the Cordilleras of of the Missouri Botanical Garden through the Flora of Panama Project (funded by NSF grant BSR-8305425). Macrocarpaea sub- caudata, an epiphytic shrub known previously from only two collections in one region of Costa ! Botany Department, University of Wisconsin, Madison, Wisconsin 53706, U.S.A. ANN. MiSSOURI Bor. GARD. 74: 310-313. 1987. 1987] FIGURE 1. square = M. macrophylla; triangle = M. subcaudata Rica, occurs in one cloud forest region in the mountains of central Panama. Recent additional collections of M. browallioides now extend the range of this distinctive epiphytic shrub. Several populations similar to both the Costa Rican M. valerii and the Colombian M. macrophylla (Kunth) Gilg are now known from the Darién. Analysis of morphological variation among the Costa Rican, Panamanian, and Colombian pop- ulations provides a basis for merging M. valerii with the now more widespread M. macrophylla. KEY TO MACROCARPAEA IN PANAMA la. Calyx Ped mm long, lobes ovate-oblong, rounded a 2a. seme viny subshrub, Ment narrow- ly lanceolate, to 18 cm long wide pa . Terrestrial shrub to 4 m, leaves s broadly elliptic, to 45 cm long and 26 wide M. macrophylla . Calyx 15-18 mm long, lobes triangular, a to cuspidate at apex 3. M. pese eee N c ox 1. Macrocarpaea subcaudata Ewan, Contr. U.S. Natl. Herb. 29: 224. 1948. TYPE: Costa Rica: [San José] La Palma, Wercklé 16492 (holo- type, US; isotype, NY). SYTSMA — MACROCARPAEA N Geographical distribution of the genus Macrocarpaea in Panama. Circle = M. browallioides; Epiphytic, viny subshrub. Leaves essentially glabrous except for small scattered hairs, some- what thickened, narrow-lanceolate, to 18 cm long and 5 cm wide; petioles to 23 mm long. Inflo- rescence terminal or axillary from upper nodes, dichasium bi- or tri-ternately compound, often long-stalked. Calyx campanulate, greenish, gla- brescent, 6-9 mm long; lobes slightly unequal, ovate-oblong, rounded or ciliolate at tips, 5-7 mm long, 3-4 mm wide. Corolla greenish yellow, to 3 cm long; tube to 2.3 cm long; lobes slightly recurved (incurved when dried), narrowly tri- angular, to 9 mm long, to 6 mm wide. Stamens inserted near middle of corolla tube; filaments to 17 mm long, just surpassing corolla lobes; anthers yellow, 4-5 mm long. Style just surpass- ing anthers. Capsules woody, 10-12 mm long ex. persistent beak of 3 mm. Flowering period at least mid-April through May. Distribution. 1,500 m in mountains east of San José, Costa Rica and between 1,150-1,260 m near continental divide between Chiriquí and Bocas del Toro provinces in central Panama (Fig. lk Additional idis examined. PANAMA. BOCAS DEL TORO: Con tal divide on carretera del oleo- ducto ca. 1 km x of Quebrada Arena, IRHE Hydro- electric ia s Knapp 5089 (MO), MUR Se (MO). CHIR Hydroelectric bout 10.1 mi. NW of Los "umi x Hornito, Antonio 4190 (MO). Macrocarpaea subcaudata occurs only in cen- tral Costa Rica and northcentral Panama. De- termining whether this disjunction is real or not must await further collecting in as yet inacces- sible cloud forest regions of both countries. Re- lationships of M. subcaudata to other species of Macrocarpaea are unclear. Macrocarpaea bro- wallioides of Panama shares the epiphytic habit with M. subcaudata, but the two are clearly un- related. Ewan (1948) considered M. cerronis Ewan and M. salicifolia Ewan from the tepuis of the Guayana Highland to be the closest relatives of M. subcaudata. The former two species (with related M. arborea (Britton) Ewan, M. quelchii (N.E. Br.) Ewan, and M. tepuiensis (Gleason) Steyerm.), however, differ from all other species of Macrocarpaea in having pollen in tetrads (Nilsson, 1968, 1970). Maas (1985) combined these six Macrocarpaea species with pollen in tetrads into /r/bachia quelchii (N.E. Br.) Maas. Weaver (1972b) cited M. acuminata Weaver and two West Indian species as the closest relatives of M. subcaudata. The Costa Rican M. acumi- nata is almost certainly related to, if not con- specific with, M. macrophylla (including M. va- lerii). Macrocarpaea acuminata and M. macrophylla share with M. subcaudata similar calyx features and might be close relatives of M. subcaudata. 2. M hylla (Kunth) Gilg, Nat. Pflanzenfam. 4(2): 94. 1895. Sap macrophyllus Kunth, Nov. Gen. & S 183. 1819. TYPE: Colombia: trail over = "£ millo to Almaguer between Pansitara and Rio Ruiz, 8,400 ft., Humboldt & Bonpland (Willd. Herb. 3561 fide Grisebach; MO pho- to 37455, of collection in Humboldt Her- barium at Paris). Macrocarpaea valerii Standley, Publ. Field Mus. Bot. 18: 928. 8. TYPE: Costa Rica: La Hondura de San José, Valerio 692 (F). Erect shrub or subshrub to 4 m tall. Leaves essentially glabrous, except for scattered small hairs, venation strongly prominent, broad-ellip- tic, to 45 cm long and to 26 cm wide, blade acute or abruptly acuminate; petioles to 3 cm long. Inflorescence terminal or axillary from upper nodes, dichasium simple or bi- or tri-ternately ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 compound, long-stalked. Calyx campanulate, fleshy, 8-14 mm long; lobes strongly unequal, ovate-oblong, hyaline-margined, 3-6 mm long and wide. Corolla greenish-white or cream, nar- rowly to openly campanulate, to 3.5 cm long; tube 2.5-3.0 cm long; lobes recurved, broadly triangular, 6-8 mm long and wide. Filaments to 18 mm long, included in or surpassing corolla tube; anthers yellow, 3-5 mm long. Style to 15 mm long; stigma bilobed, the lobes 2 mm long. Capsules woody, to 20 mm long, persistent- beaked. Flowering period at least May through July. Distribution. 1,000-1,800 m in mountains of Central Costa Rica; 1,000-1,400 m in Serranía del Sapo and Serranía del Darién of Darién Prov- ince, Panama (Fig. 1); and common at 1,500- 2,500 m in the western and central Cordilleras of Colombia. Additional specimens examined. PANAMA. DARIEN: S o a expedition, Gentry & Mori 13655 (MO), CEU: P ‘al 16975 (MO). Macrocarpaea macrophylla is distinctive as a tall erect shrub with large round leaves and prominent venation. As here defined, M. macro- phylla ranges from Colombia to Costa Rica. Standley originally described the Costa Rican M. valerii based only on the type specimen. Ewan (1948) reiterated the differences between the two species based on only two Costa Rican collec- tions. These differences were confined to size and shape of leaves and to corolla shape. Both sets of characters are subject to sampling error de- pending on how the plants were collected and pressed. Subsequently, additional collections of " M. valerii" in Costa Rica indicated that these features were not consistent, causing Weaver (1972b) to state that “the two are virtually iden- tical, except the calyx of M. valerii is glabrous, while that of M. macrophylla is spiculate." Co- lombian, Panamanian, and Costa Rican speci- mens of these two taxa at MO and WIS showed considerable variation in the degree of surface ornamentation on the calyx. No consistent calyx differences or other foliar and floral differences were seen among specimens described as M. macrophylla and M. valerii. This necessitates the inclusion of these Colombian, Panamanian, and Costa Rican specimens into M. macrophylla (Kunth) Gilg, the name with priority. 1987] 3. Macrocarpaea browallioides (Ewan) Robyns & S. Nilsson, Bull. Jard. Bot. Natl. Belgique 40: 13. 1970. Lisianth(i)us browallioides Ewan, Proc. Biol. Soc. Washington 64: 132. 1951. TYPE: Panama. Bocas del Toro: north- ern slopes of Cerro Horqueta, Robalo Trail, Allen 4932 (MO). Epiphytic shrub to 1.5 m tall. Leaves glabrous, + coriaceous, dark green and black-punctate above, gray-green below, elliptic to ovate-ellip- tic, to 11 cm long and to 5 cm wide; petioles to 25 mm long. Inflorescence a compound dicha- sium, + long-stalked. Calyx campanulate, light green, 15-20 mm long; tube short; lobes ovate- triangular, strongly overlapping, 12-16 mm long, acute to long-acuminate at apex. Corolla fleshy, yellow-green to cream yellow, infundibuliform, 3.0-4.0 cm long; lobes subequal, ovate-trian- gular, acute at apex, to 1.2 cm long. Stamens inserted ca. 6 mm above base of corolla tube, + exserted; anthers yellow, 5-6 mm long. Style just surpassing anthers. Capsule woody, to 14 mm long, excluding persistent beak of 6-8 mm. Flow- ering in February, May, August, and late No- vember. Distribution. 1,690-2,200 m, cloud forests of cerros Horqueta, Pate Macho, and Colorado, provinces of Chiriquí and Bocas del Toro. Addi tional A Li examined. PANAMA. BOCAS st along trail from end of Rio Palo Alto road to Chiriqui/ Bocas del Toro border near peak of Cerro Pate Mac v Hammel 5781 (MO). CHIRIQUI: Cerro Pate Macho, ca. 5 mi. NE of Boquete, trail to continental divide ‘leading to Finca Serrano, Antonio 2654 (MO); Cerro Colorado, along mining road 31. 6 km beyond bridge over Rio San Felix, 10.6 km beyond turnoff to Escopeta, Croat 37155 (MO); Cerro Pate a e of Pate Macho, Hammel 6112 (MO); NW Finca OVIIO ridge no, | eiua et al. 4900 (MO). Macrocarpaea browallioides is easily distin- guished from the other Central American species of Macrocarpaea by its large calyx and long-acu- minate calyx lobes. Like M. subcaudata it is epi- phytic, but unlike M. subcaudata it is erect rather SYTSMA — MACROCARPAEA 313 than viny. Several of the collectors do not men- tion an epiphytic habit. However, the distinction between an epiphytic and terrestrial habit is ten- uous on an extremely wet, peatlike soil as is encountered on Cerro Pate M i phytic as well as free-standing ‘although on mats of living roots or trunks) individuals of M. bro- wallioides were seen on this cloud forest Dead The relationship of this species to other members of Macrocarpaea is unclear. Neither Ewan, who first described the species as a Lisianthius, nor Robyns and Elias, who transferred it to Macro- carpaea, gave any indication as to its likely rel- atives. Only three known species approach M. browallioides with its distinctive short calyx tube and long, overlapping, and acuminate-tipped lobes: M. cochabambensis Gilg of Bolivia; M. glabra (L.f.) Gilg endemic to paramo vegetation above Bogotá, Colombia; and M. guttifera Ewan of Amazonia, Brazil. LITERATURE CITED Euas, T. S. & A. Ropyns. 1975. Family 160. Gen- tianaceae. In Flora of Panama. Ann. Missouri Bot. Gard. 62: 61- EwAN, J. 1948. A revision of Macrocarpaea, a neo- tropical genus of shrubby gentians. Contr. U.S. Natl. Herb. 29: 209-249. GRISEBACH, A. H. R. 1838. Genera et species Gen- tianearum. Stuttgart & Tübingen. MAAS, P. 1985. Nomenclatural notes on neo- tropical Lisyantheae ( — uris Proc. Kon- inkl. Nederl. Akad. Wetensch. C. 88: 405-412. etal. 1984. wd Ves studies in n Pike oup the Lisianthius complex. Acta Bot eerl. 32: 371-374 Pollen morphology in the genus significance. Svensk Bot. Tidskr. 62: 338-364. . 1970. Pollen morphological contributions to t. (Gentiana- 1-43. . Biosystematics and evolution in the Lisianthius skinneri (Gentianaceae) species complex in Panama. Ann. Missouri Bot. Gard. (MS). Weaver, R. E., JR. 1972a. A revision of the neo- tropical genus Lisianthius (Gentianaceae). J. Ar- nold Arbor. 53: 76-100, 234-311. 2b. The genus Macrocarpaea (Gentiana- ceae) i in Costa Rica. J. Arnold Arbor. 53: 553-557. MONOECY AND SEX CHANGES IN FREYCINETIA (PANDANACEAE)! HANS-HELMUT POPPENDIECK2 ABSTRACT In cultivation, did: sae of the s dioecious genus sequentially and MM on dem plans pistillate and s inflore ces "D. 4 A These eg are AM een) E with other cases of occasional deviations from dio According to most textbooks and general ref- erence works the genus Freycinetia is considered to be dioecious (Warburg, 1900; Heywood, 1979; Dahlgren et al., 1985; Stone, 1984), although some exceptions have been noted (Stone, 1972; Cox, 1981; Cox et al., 1984), drawing attention to the significance of these presumably rare ex- ceptions to the dioecious condition. In Freyci- netia reineckei Warb., Cox has observed func- tionally bisexual spikes, and both pistillate and staminate inflorescences on the same branch in Freycinetia scandens Gaudich., and Stone (1972) has noted the same in Freycinetia negrosensis Merr. and F. imbricata Blume The following observations made in the trop- ical greenhouses of the Hamburg Botanical Gar- den confirm and extend these earlier observa- tions and, by adding some new pieces to the puzzle, seem to indicate that sex expression in this genus is to a considerable extent under en- vironmental control. Both Freycinetia funicula- ris (Savigny in Lam.) Merr. and F. cumingiana Gaudich. (synonymous with F. /uzonensis Presl, according to Stone) have been observed to show monoecy. Both of the species mentioned above are easily cultivated from cuttings, and the material in cul- tivation probably represents single clones of each, so that the observations made at different places relate to genetically identical plants. Freycinetia funicularis is comparatively widely distributed in German botanic gardens, but its origin cannot be ascertained; however, it is a native of Indo- nesia, and probably was introduced from the Bo- tanic Gardens in Bogor (formerly Buitenzorg), where it has been in cultivation for perhaps a century and still is common. Freycinetia cumingiana (F. luzonensis) is a Philippine species which came to Hamburg from the Bonn Botanic Garden, and according to the curator there, Dr. K. Kramer (pers. comm .), it was introduced to Europe via Tübingen from a source in the U.S.A., initially under the name "F. cunninghamiana." It has been possible to trace this clone back to its original source. Ap- parently it had been sent by E. D. Merrill from the Philippines to the New York Botanical Gar- den, from there to Puerto Rico, and to the Fair- child Tropical Garden in Miami, Florida, where it caught the attention of Dr. R. A. Howard of Harvard University, who reported his observa- tions to Dr. B. C. Stone (see Table 1). Eventually this species found its way to Europe, perhaps from the Fairchild Tropical Garden OBSERVATIONS ON GROWTH AND SEX EXPRESSION Observations on both species are summarized in Table 1. It is remarkable that despite the pro- ed diffe i hoot hit t i í S ee Fi 1), the two species have produced staminate and pistillate inflorescences, both sequentially and si- multaneously. Freycinetia cumingiana appar- ently produces different kinds of shoots, prede- M shoot diam decreases with age, the initially pistillate shoots convert to the production of staminate inflores- cences. Evidently, shoot diameter and sex expression are correlated in this species. The qualitative “jump” may occur when the shoot diameter decreases to less than 4 mm (Fig. 2; able 2). Perhaps this is also the diameter range wherein mixed i y form. In Fre rey- ' My thanks to Dr. B. C. Stone for encouragement, to Dr. K. Kramer for details on Freycinetia cumingiana, J. Bogner on F. Led gen Dr. M. F our greenhouse sta expert cultiv ? [nst ANN. Missouni Bor. GARD. 74: 314—320. 1987. eis dau Prof. Dr. K. Kubitzki for advice, and Mrs. H. Sch wob-Tonn and .f Alemana. Botanik u. rie Garten, Hesten 10, D 2000 Hamburg 52, West Germany. 1987] POPPENDIECK — FRE YCINETIA 315 TABLE 1. Comparison of growth and flowering of two species of Freycinetia in the Hamburg Botanic Gardens, 1982-1986 Freycinetia funicularis Freycinetia cumingiana Large climber (“rooter”) with horizontal clinging roots and vertical roots; inflores- cences lateral on at least one-year-old, still leafy shoots, often penetrating the leaf-bas- Morphology cei “leaner” with perennial shoots ich may flower in 2nd year; inflores- cences ande in axis or on rather PR lateral shoots; shoots dimorphic, dia mm in staminate shoots and PET mm in pistillate shoots. Flowering particulars 1982 Abundant (ca. 50) inflorescences. Several vegetative shoots; one fertile, with about 12 pistillate inflorescences 1983 Abundant but exclusively pistillate inflores- Only staminate inflorescences observed. ces 1984 Numerous staminate inflorescences plus a Not recorded; allegedly staminate, but pres- single pistillate inflorescence at the base of ence (or not) of pistillate inflorescences un- the plant; viable seeds and offspring ob- certain tained after hand-pollination. 1985 Twelve staminate inflorescences, most pro- Two ramified shoots with staminate inflores- 1986 Additional observations duced later than usual after severe pruning had reduced the shade made by the plant itself. No inflorescences on the old plant. Young plant (4 yr. old) with 12 staminate inflores- cences mostly on vertical axes, 5 riu inflorescences on subapical parts of hori- zontally trained shoots, and one mixed in- Vend on a vertical shoot. pores yields fruit Similar observations by J. Bogner in Munich Bot. Gard. Younger plants from cuttings first al pistillate then both pistillate and staminate inflorescences in the next and unde years. Encke's account (1958) of “Freycinetia insignis" probably refers to F. funicularis. cences which had probably flowered the year before; 3 pistillate shoots, at least 2 flowering for the first time (Fig. 1); several fruits formed (no viable seed). Staminate shoots continue to form staminate apparently pistillate-determined shoots have been formed. New shoot formed in 1985 (diam. 6.2 mm) is pistillate. A somewhat different sequence of flowering y Dr as been observed (Bonn Bot. Gard.): the first inflorescences were staminate; later, also pistillate inflo- res were p the branches. Eventually also mixed inflo- , when I found an inflorescence pro- ducing pollen” (April 1969). cinetia m this shoot differentiation is lacking, s ko seal pistillate inflorescences in 1983 is unexplained. However, it seems to indicate that an adjustment over the whole vegetative body can take place, which may be hormonally (or that the synchronized formation of nutritionally) determined. Observations made in the Munich Botanical Garden by Josef Bogner (pers. comm.) suggest that larger and vigorous plants of F. funicularis may develop inflores- cences of both sexes. It is difficult to assign these Freycinetia species 316 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FI f ah black circles; stami nate a penance circles; vegetative shoots— two species of Bicis Pistillate inflorescenc ces— s. Le laris, an approximately three-year-old | plant raised from a cutting, ribet for de first time; inflorescences, at least one-year-old parts of the main shoots. X—plant t rained oo from this point. * —pistillate inflorescences formed here in the resent year, two pistillate shoots form mingiana, showing (left to right) shoot of p pet for ien first ES a two- dione pistillate shoot, and a (probably) three-year-old staminate shoot. Note acrotonous ramifications, and ‘ influenced bo different yes of shadin to one of the architectural models of Hallé et al., although some species have been so assigned by Castro dos Santos (1981). This is because the “opportunistic” growth of these lianas with eas- ily formed accessory shoots makes it difficult to plagiotropic growth. In F. cumingiana, inflorescences are al- ways terminal on the main shoot as well as on the lateral shoots, so that branching in the floral region is monopodial, except for occasional veg- etative shoots. In F. funicularis, the inflores- cences are lateral on short shoots. When the staminate and pistillate inflores- cences were produced simultaneously, hand-pol- lination of the latter resulted in fruit formation; in F. funicularis seeds from such fruits germi- nated to produce viable seedlings. In F. cumin- giana, attempts to pollinate by hand were un- successful in both Hamburg and Bonn Botanic Gardens. Although the pistillate heads enlarged considerably and were not aborted, fertilization apparently did not occur; at least the pollen was a distribution of inflorescences and vegetative shoots probably able to trigger development. It may be noted that in the related genus Pandanus, no pollen stim- ulus is necessary, at least in some species, and parthenocarpy is normal. It must be remembered that attempts to pollinate tropical plants in tem- perate-zone greenhouses are chancy, for even if the basic aspects of the breeding system and the receptive period of the stigmas are understood, it may be impossible to obtain fertilization. An example of this difficulty is Cinnamomum verum (C. zeylanicum), the true cinnamon, as recounted by Kurz (pers. comm.). On the other hand, if pollination is successful, the possibility of pseu- dogamous agamospermy cannot be ruled out (Gadella, 1983). These observations are casual but may serve as the basis for some speculations, if only for the purpose of stimulating further ex- periments (which cannot be performed with scarce material in cultivation). The evolutionary significance of dioecy has re- : 243-296. 1984) where a survey of the subject and further 1987] TABLE 2. Branch diameter and sex expression in r: wasa cumingian was Visits ca. 3 cm below the inflores- cence d iiie the expansion immediately below it). The pistillate shoot measure is that shown in Figure 2. e respective means are m, = 3.98 + 8 mm for pistillate inflorescences, m, = 3.08 + 0.35 mm for staminate inflorescences on pistillate shoots, and m, — 2.76 + 0.26 mm for staminate inflorescences on sta- minate shoots. Branch diameter for pistillate inflores- cences differs significantly from that for staminate in- florescences. aminate Pistillate Inflorescences Branch Diam. Inflores- 9 3 (in mm) cences Shoots Shoots 1.6-2.0 l 2.1-2.5 l 4 2.6-3.0 l 4 11 3.1-3.5 2 2 3 3.6-4.0 12 2 4.1-4.5 5 l 4.6-5.0 4 references may be found. Discussion centers around the question of whether the so-called benefits of dioecy (Willson, 1983) are EP PPE or ecologically determined, i.e., whether the motion of outcrossing or ecological aera on reproduction are evolutionarily decisive. This matter need not be resolved here, but our con- cern is with occasional departures from dioecy, a phenomenon that has been termed “leaky dioe- cy” by Baker & Cox (1984). Such leaky dioecy may be considered as one prerequisite for suc- cessful colonization of remote islands, since propagation by selfing becomes possible. Apo- mixis is another kind of solution to this problem, as is the parthenogenesis observed in some in- sular animal populations (e.g., Cox in White, 1985; Mau, 8). Rohwer & Kubitzki (1984) recently reported an interesting example of leaky dioecy in the neotropical riverine willow Salix martiana Leyb., which was found to produce regularly mixed cat- kins with staminate, pistillate, and perfect flow- ers. This was attributed to the difficulty of pro- ducing offspring after a probable long-distance dispersal and in an unstable environment. Mixe catkins are not uncommon in Salix (Velenow- sky, 1904; Toepffer, 1925) but the high incidence in a tropical species, certainly much less studied than its temperate congeners, suggests that here POPPENDIECK — FREYCINETIA 317 FIGURE 2. Diagrammatic sketch of sex expression on an initially pistillate shoot in two successive years. oken line marks th note that due to the straggling habit and the necessity of training ui climber, some shoots may have been removed. e further Ting the PIE inflores- ences (open dise formed on the distal parts of the Muse 1986. it is a more or less regular phenomenon. Al- though hermaphroditic flowers are unknown in Freycinetia, the two genera may be taken as par- allel cases for the evolution of monoecy (which may be a reversal of a previous evolutionary trend from monoecy to dioecy). Another aspect of this possible parallelism will be discussed next. Developmental plasticity is one of the char- acteristics of liana species, as exemplified by their *opportunistic" shoot architecture in compari- son with trees (Etifier, 1981). This plasticity is needed to adjust to a temporarily very hetero- eneous habitat, where, for instance, light inten- sity encountered by juvenile plants is drastically different than that encountered by adults. In go ANNALS OF THE MISSOURI BOTANICAL GARDEN \ FIGURE 3. Aspects of growth of two Freycinetia species. a, b. Freycinetia funicularis. —a. Staminate iud rescence on a vertical shoot, ca. 18 cm diam.— b. Pistillate piece at anthesis (spikes 8 cm long). c 1987] Freycinetia, at least in the two species observed for this study, such plasticity extends to sex expression. This is not surprising since environ- mentally determined sex expression has several selective advantages for species living in a “patchy” environment (Charnov & Bull, 1977; Willson, 1983), and sequential hermaphroditism has been reported for many species. The factors invoked include light intensity (e.g., Catasetum; Dodson, 1962), hormonal and nutritional status (e.g., Elaeis guineensis, Williams & Thomas, 1970), or disturbance of vegetative parts (e.g., Arisaema: Schaffner, 1921), all of which may interact variously, and all of which may be ap- plicable to the present cases. Thus, in F. funi- cularis, horizontal training of the branches may have altered the hormonal balance toward the production of pistillate inflorescences; exhaus- tion of nutrient reserves, self-shading of shoots, or pruning may have been responsible for the switch back to staminate inflorescence produc- tion. Experimental proof for this would be dif- ficult to obtain. At least, the : size- -correlated sex expression in plants of F. will permit further study. Cuttings have been made from the differently determined axes and their fate will be observed in coming years. Clearly it should not be too difficult to find plausible explanations for the adaptive values of dioecy, monoecy, or plasticity in sex expression in a given situation. However, the phenomenon of leaky dioecy is still puzzling. Why is it re- stricted to certain genera and absent in others? hy, for instance, are no such cases known in Populus or in Pandanus, which are the sister gen- era of Salix and Freycinetia, respectively? In Populus and Pandanus pollination is believed to be chiefly by wind (Toepffer, 1925; Cox, 1982). A possible explanation is that in anemophilous plants, the staminate and pistillate functions must suit quite different needs and, as a result, tend to evolve divergently. Such divergence seems improbable in zoophilous groups, where a high degree of similarity is advantageous (in overall form, size, color, odor, and perianth details, though not in the details of staminal and gynoeci- al form). The correlation of leaky dioecy with POPPENDIECK — FREYCINETIA 319 change from anemophily to zoophily is a pos- sibility worth investigating. To obtain evidence bearing on these questions, many more observations are needed, preferably on the same specimens over a number of years. The statement by Cox et al. (1984) about the value of living collections for this kind of study may be emphasized again. Phenomena such as these tend to escape the notice of both purely field- and herbarium-oriented workers, but bo- tanic gardens provide an excellent location in which to study such things as “leaky dioecy." VOUCHER MATERIALS Herbarium material of both Freycinetia species discussed is deposited in PH and HBG, and the behavior of F. funicularis is also documented in M (Bogner 1783). Additional prints for photo- graphic documentation are on deposit in PH and HBG LITERATURE CITED BAKER, H. G. & P. Cox. 1984. Further thoughts = dioecism and islands. Ann. Missouri Bot. Gar 71: 230-239. CASTRO DOS SANTOS, A. DE. 1981. L’appareil végétatif des monocotyledons. Thèse, Acad. de Montpel- lier, Univ. des Sciences et Techniques de Langue- doc. CHARNOV, E. L. & J. BULL. 1977. hen is sex en- vironmentally determined? Nature (London) 266: —830. Cox, P. A. 1981. Bisexuality in the Pandanaceae: new findings in the genus Freycinetia. Biotropica 13: 8. . Vertebrate pollination and the main- tenance of dioecism in the genus Freycinetia (Pan- danaceae). — Naturalist 120: 65-80. , B. WALLACE & I. BAKER. 1984. oecism in the genus Fre reycinetia peat an L im 16: 313-314. DAHLGREN, R. T. CLIFFORD & P. F. Yeo. 1985. The Families of the Monocotyledons. Springer- Verlag, Berlin, Heidelberg, aid Dopson, C. H. 19 Pollination ie pa ariation in the subtribe Catasetinae (Onehidaccae). pues Mis- souri Bot. Gard. 49: 35-56. ENcKE, F. 1958. Pareys Blumengartnerei. 2. Auflage. Paul Parey Verlag, Berlin und Hamburg. (See p. 101. ETIFIER, M. E. 1981. Données sur la strategie de — Freycinetia cumingiana. —c. Staminate inflorescence ca. 6 cm diam.— d. Pistillate inflorescence after anthesis, spikes 1.5-2 cm long.—e. Bas shoot with caducous bra thinner). al parts of shoots (from left to right, two old pistillate acts, sais pistillate shoots interspaced with two staminate shoots, the latter much —f. F. funicularis seedlings ca. six months old, their leaves about 2 cm long. shoots, current pistillate 320 croissance de quelques lianes tropicales. Thése, Acad. de Montpellier, Univ. des Sciences et Tech- I 1983. Some notes on the deter- mination of the mode of reproduction in higher plants. Proc. Koninkl. Nederl. Akad. Wetensch. Fowering Plants of the World. Oxford Univ. Press, Oxford, London, Melbourne MAU, K. G. 1978. Nachweis natürlicher Partheno- gense bei Lepi RETR acer (R ig Sauria, Gekkoni- ) mandra ROHWER, KUBITZKI. nus Salix martiana, a K. regularly hermaphrodite willow. Pl. Syst. Evol. 144: HAMM. J. H. 1921. Control of the sexual state in Á š Pea 4 s d 45 A gs Amer. J. Bot. 9: 72-78 SrowE, B. C. 1972. Materials for a monograph of Freycinetia Gaud. (Pandanaceae). XV. The Su- matran species. Federation Mus. J. n.s. (for 1970) 15: 203-207. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 1984. A guide to collecting Pandanaceae (Pandanus, Freycinetia, and b raranga). Ann. Missouri Bot. Gard. 70: 137- TOEPFFER, A. 1925. Salicaceae. = 373- 377 in O. v. Kirchner, E. Loew & C. Schroeter (editors), Le- biu ie EA de Blütenpflanzen Mittel-Euro- pas. Band II, 1. E. Ulmer, Stuttgart. VELENOWSKY, J. Tur Vergleichende Studien über die Salix-Blüte. Beih. Bot. Centralbl. 17: 123-128. WARBURG, O. 1900. Pandanaceae. Pflanzenreich IV.9: 1-99. (Heft III. WHITE, J. (editor). 1985. Plant Demography. Aca- of Pandanus tectorius in Polyne WILLIAMS, C. N . L. THOMAS. 1970. Observa- tions on sex differentiation i in the oil palm, Elaeis guin eue L. Ann. Bot. 34: 957-963 WILLSON, M. . Plant E ass Ecology. John "is & Sons, New York. (See p. 79.) A GUIDE TO COLLECTING LECYTHIDACEAE! Scott A. MORI AND GHILLEAN T. PRANCE? ABSTRACT Because of their arborescent habit and unique flowers and fruits, Lecythidaceae present specialized collection problems. Instructions for the preparation of more informative collections of Lecythidaceae vided. for use in taxonomy are pro The Lecythidaceae sensu lato are a pantropical family of small to very large trees. The family includes four subfamilies: Planchonioideae with 55 species in six caps distributed through trop- ical Asia, Malaysia, northern Australia, and the Pacific Islands; Fie. with five species in a single genus distributed in Madagascar, In- dia, and Malaysia; Napoleonaeoideae, with 11 species in two genera distributed in West Africa and one species in the upper Rio Negro of Ama- zonia; and the Lecythidoideae, with about 206 species in ten genera distributed through tropical merica from Veracruz, Mexico to southern Brazil (Kowal et al., 1977; Prance & Mori, 1979). Because our collecting experience has been most- ly in the Neotropics, this essay emphasizes the Lecythidoideae. However, the methods de- scribed are applicable not only to other subfam- ilies of Lecythidaceae but to tropical trees in gen- ral. General collections of plants provide the data upon which the monographic and floristic treat- ments of plant taxonomists are based. A single collection of a species usually does not provide sufficient data for its taxonomic description. Consequently, collections are needed that rep- resent: (1) all parts of the plant used in classifi- stages of the life cycle of the plant (flowers, fruits, seeds, and seedlings), (3) individuals from tinct habitats in which the plant grows, (3 in- dividuals from throughout the geographic range of the species, and (5) intra- and interindividual variation. Many recent collections of neotropical Lecy- thidaceae have added little to what was already about those features of the plant, such as habit and bark, that are not preserved on the herbar- ium sheet. Careful notes on habitat are usually lacking and even flower color is often not re- corded in a manner that can be interpreted by taxonomists. Worst of all, flowers are often dried under such extreme heat and pressure that they become so carbonized that their structure is im- possible to determine. Even fruits are collected in such a way that their taxonomic features and morphological variation are difficult to interpret. Finally, few collectigns or weir and fruits are from ibe-s same ie. Another problem is the continued collection of herbarium material of the same taxa from the same area by the same collectors. However, some duplication is necessary to voucher ecological studies of tropical forest structure and compo- sition. Collections of some Lecythidaceae, such as Gustavia augusta L. and G. hexapetala (Au- blet) J. E. Smith, continue to late without augmenting the information needed for under- standing the species. These collections do not merit the cost of their processing or the valuable space they occupy in herbaria. This same lack of selectivity in collecting occurs in almost all woody tropical families, for example, the Chrysobala- naceae (Prance & Campbell, in press). If further progress is to be made in the tax- onomy of tropical woody groups, more careful and selective collections by more sophisticated collectors are needed. The purpose of this article is to provide the information needed for making these collections. While we emphasize Lecythi- daceae, similar needs pertain to almost all rain- forest woody plant families. TREE CLIMBING Because of their large size, many species of Lecythidaceae can be collected only by felling or ! We are grateful to Bobbi Angell, Bethia Brehmer, and H. M. Fukuda for the preparation of the original s wks and to Carol Gracie for rearr anging Figure 4 and for her comments on the manuscript. The New York Botanical Garden, Bronx, New York 10458-5126, U.S.A. ANN. Missouni Bor. GARD. 74: 321-330. 1987. 322 40cm FIGURE |. war el pipes pet to collect Lec- ythidaceae. — A. tree im spikes (*griffes"). — B se s En used b. native climbers ("*peconha"). Reprinted with ission fr e m Memoirs of the New York Botanical Garden climbing. As a general rule, trees should be felled earing, struction, etc. Forest clearing provides an excel- lent opportunity for making collespons dom felled trees. Thet from a herbarium specimen often does not war- rant the sacrifice of a tree. This is especially true when collecting near Indian villages or settle- ments where trees may have economic value or cultural significance to natives. Boom (1985) has shown that 82% of the species of trees surround- ing a Chácobo Indian village in Bolivia are used in one way or another by the Indians. We have found climbing to be the most sat- isfactory method of obtaining specimens of Lecy- thidaceae. Although numerous methods are used for gaining access to tropical trees, we have em- ployed (1) native climbers, (2) “Swiss Tree Grip- pers" (Mori, 1984; available from Forestry Sup- pliers, Inc., 205 West Rankin St., P.O. Box 8397, Jackson, Mississippi 39204-0397), and (3) French tree SBEHDEDR spikes (Fig. 1). N becoming i InCI easingly dif: ficult to find. We have employed them most ef- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 fectively to collect Lecythidaceae in Brazil, es- still common because of the custom of gathering edible fruits from Euterpe and Oenocarpus palms. The Brazilian climbers can efficiently scale trees under 35 cm DBH by using an adjustable canvas loop called a “‘peconha” (Fig. 1). A climber can be hired for about $10 per day. For trees from 50 to 72 cm DBH we have used “Swiss Tree Grippers.” To use them, lianas and epiphytes have to be cleared from the trunks before the trees can be ascended, but the trees themselves, in contrast to those scaled with spikes, are not damaged. The weight, bulk, and high price of the grippers often counterbalances the advantage of being able to climb the few addi- tional trees not climbable with spikes. e have found French climbing spikes (Fig. 1) to be the most efficient method for collecting Lecythidaceae. Smaller spikes (24 cm diam.) are used for trees from 10 to 25 cm DBH, whereas larger ones (35 cm diam.) are employed for trees from 26 to 50 cm DBH. The advantage of these spikes over the single spur spike preferred by some climbers is the ease with which trees can be climbed. The climber's weight and safety belt secures him to the tree in such a fashion that his hands are free for using clipper poles, capturing insects for pollination studies, etc. The advan- tage of the single spike apparatus is that trees of all sizes can be ascended with one pair of spikes. The French spikes are available from Ets La- coste, 24160 Excideuil, France for about $50 a pair, not including shippi All climbs should be ads with the climber secured to the tree with a safety belt. We prefer the Klein nylon protective belt with two D rings in combination with the Klein nylon adjustable lanyard (both available from Forestry Suppliers, Inc.). The use of two lanyards insures that the climber is attached to the tree at all times. When the climber reaches a branch, one lanyard re- mains around the trunk below the branch while the other is secured around the trunk above the branch to be passed. The belt around the main trunk beneath the branch can then be released and the climber is free to move above the branch. We have climbed several thousand trees using the French spikes without any falls. Neverthe- less, the climber should be extremely careful be- fore and during any climb. Before climbing, the tree should be carefully inspected to insure that it is in good enough condition to support the weight of the climber, and for the presence of 1987] stinging and biting animals. Special attention should be paid to swarms of bees, as they may be made up of the particularly aggressive African honey bee, which has been responsible for the death of at least one biologist. The bases of trop- ical trees often serve to house creatures that vary from sand flies that carry the protozoan disease of leishmaniasis to snakes that can inflict painful and deadly bites. The tree itself may be occupied by bees, wasps, biting ants, and the deadly snake Bothrops bilineata (an arboreal fer-de-lance). The last is more frequent in fruiting trees, where it lies in wait to prey upon visiting frugivores. Ex- treme care should be used when reaching over branches or when climbing past epiphytes. Near- by trees should also be examined so that wasp and bee nests are not disturbed during manip- ulation of the clipper pole. Finally, the condition of the climbing gear should be constantly mon- itored and any frayed or defective parts should be immediately replaced. The recent fatal fall of a young botanist in Venezuela is a shocking re- minder of the danger involved in making tropical tree collections Collection m specimens from the tree's canopy by a collector on the ground or in the tree itself is greatly facilitated by the use of a tree pruner attached to a series of aluminum poles. We have found the poles developed by botanists of the Missouri Botanical Garden staff to be the most efficient. This system includes three sets of two telescoping aluminum poles attached to one another by a spring activated button that passes from a hole in one pole into that of the next. Each pole is 1.8 m long so that when all six poles are joined nearly 11 meters plus the height ofthe collector can be attained. However, when more than four poles are joined it is very easy to bend the poles. In order to avoid this, it is best to cut with the poles in the most vertical position pos- sible and to wrap the cutting rope around the poles several times in order to insure that the cutting force is directed along the poles. We find the best tree pruner to be the Snap-cut model 33 (available from Forestry Suppliers, Inc.) which fits exactly onto the end of one of the smaller aluminum poles. This head is light enough to allow maneuverability yet strong enough to cut most branches to about 4 cm in diameter. LEAF VARIATION Normal collection for herbaria does not ade- quately sample intra- and interindividual vari- ation in leaf morphology. Collections which in- MORI & PRANCE—LECYTHIDACEAE COLLECTING 323 clude shade and sun leaves, young and old leaves, and extremes in leaf size are invaluable in helping to solve taxonomic problems. In Couroupita, for example, there was much taxonomic confusion due to differences in leaves from the crown ver- sus those that appear on the cauliflorous inflo- rescence. These leaves differ in shape, texture, and pilosity and were sometimes described as different species, though they in fact represent variation within the same species. Consequently, it is important to document leaf variation, even if more than one herbarium sheet is needed to do so. FLOWERS AND FRUITS The remarkable variation in the floral and fruit morphology of Lecythidaceae can often be re- lated to pollinators and seed dispersers. The two principal floral types in neotropical Lecythida- ceae differ in the structure of the androecium. One type is radially symmetrical (actinomor- phic) and the other is bilaterally symmetrical (zy- gomorphic, Fig. 2). The actinomorphic androe- cium consists of a fused basal portion, the staminal ring, which is surmounted by free sta- mens (Fig. 2). In the zygomorphic androecium, the staminal ring is prolonged on one side into a stamen-free area called the ligule, which ter- minates in an appendage-bearing hood (Fig. 2). The hood may be open (with a space between it and the summit of the ovary) or closed and tight- ly appressed to the summit of the ovary. New World genera with the former type of androe- cium are Asteranthos, Gustavia, Grias, and Al- antoma, whereas the latter type is found in Couroupita, Corythophora, Bertholletia, Lecy- this, Eschweilera, and Couratari (Fig. 3). The flowers of Cariniana are intermediate between these two types (Fig. 3 Pollinator d d ] the androecium. In gotiinvanmanephin species Jd reward is always pollen. Moreover, the pollen collected by pollinators from ic onis i flowers is the same kind of pollen that affects fertilization, i.e., no nectar or specialized pollen are produced. In the zygomorphic species either pollen or nectar may serve as the pollinator re- ward oevet, His pellen souaga by polli- — morphologically (e. g., Couroupita guianensis Aublet: Mori & Boeke, 1987; Mori et al., 1980), from “g which affects pollination. This spe- cialized or fodder pollen is found in the anthers ofthe hood or in a group of stamens of the stam- 324 ANTERIOR DISTAL POSTERIOR ANNALS OF THE MISSOURI BOTANICAL GARDEN Staminal Ring Ligule [VoL. 74 PROXIMAL Staminal Ring ANG IS W IGURE 2. Androecial structure of a zygomorphic-flowered New World Lecythidaceae. — A. Side view of flower with petals removed, showing terminology used in describing floral orientation and in describing the androecium. — B. Medial section of the androecium.— C. Artificially opened androecium.— D. Stamen from staminal ring. Reprinted with permission from Flora Neotropica. inal ring adjacent to the ligule. Color differences help to separate the two types of pollen. For ex- ample, in Lecythis pisonis Cambess. the fodder pollen, located in the hood, turns black after 24 hours, whereas the pollen in the staminal ring remains yellow. In L. corrugata Poit. subsp. cor- rugata the fodder pollen, located in a row of stamens on the ligular side of the staminal ring, is yellow in contrast to the white pollen of the remainder of the stamens. Nectar is the principal pollinator reward in those zygomorphic species with coiled androecial hoods. The aforementioned androecial structures as well as other floral characteristics provide fea- tures useful in the classification of Lecythidaceae. Unfortunately, most collections do not ade- 1987] MORI & PRANCE—LECYTHIDACEAE COLLECTING 325 GUSTAVIA HEXAPETALA CARINIANA PAUCIRAMOSA outil FDE LECYTHIS COUROUPITA PISONIS GUIANENSIS = ANANA PRIDE EIE BERR LECYTHIS TUYRANA BERTHOLLETIA EXCELSA ' var I f: : rU ESCHWEILERA ESCHWEILERA COURATAR 1 COLLINA LONGIPES STELLATA FIGURE 3. Variation in androecial structure of New World Lecythidaceae. In all cases the androecium has been removed from the flower and cut in medial section. Gustavia, Grias, and Allantoma are actinomorphic, whereas the remaining genera display varying degrees of zygomorphy. Reprinted with permission from Flora Neotropica. 326 quately preserve or describe these features. The most common error in the collection of flowers of Lecythidaceae is the use of excessive heat and pressure in the drying process. It is useful to dry at least some of the flowers outside the press in a paper bag over low heat and to preserve flowers in F.A.A. or 70% ethanol. Structure of the an- droecial hood can also be recorded by photo- graphing a medial section of it. The photographs should subsequently be attached to the herbar- ium sheet. Flower color and size are also useful features that have not been adequately considered in the classification of Lecythidaceae because they are seldom properly recorded. After drying, it is very difficult to determine flower size, and therefore the maximum diameters of several flowers from the same tree should be measured and stated on the label. It is also important to record the color of all the separate floral parts. It is not sufficient to state that the flowers are white, yellow, or some other color. If indeed they are entirely of a single color, this is unusual and should be clear- ly indicated. In many species of Lecythidaceae, the petals and androecial hood are different colors and this should be noted. In addition, the color and position of differently colored anthers should be recorded, as this indicates whether one or two types of pollen are produced within the same flower. The fruits and seeds of neotropical Lecythi- daceae have evolved a variety of forms in re- sponse to pressures exerted by different seed dis- persal agents, fruit and seed predators, and environmental constraints. Major trends caused by these selective pressures are (1) indehiscence versus dehiscence; (2) retention of the fruit on the tree until the seeds have been released versus drop of fruits with the seeds inside; (3) fleshy versus woody pericarps; (4) development of mu- cilage within the pericarp; (5) lateral versus basal fleshy arils; (6) development of arils that com- pletely surround the seeds; (7) loss of arils; (8) development of membranous, winglike arils; (9) development of plano-convex cotyledons as in Gustavia; (10) development of leaflike cotyle- ons as in Cariniana, Couratari, and Courou- pita; (11) lack of well-developed cotyledons as in Allantoma, Bertholletia, Corythophora, Esch- weilera, Grias, and Lecythis; and (12) terminal versus lateral seed germination. These characters have coevolved with animal and wind seed dis- persers and environmental factors throughout a long history under rainforest conditions (Prance ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 & Mori, 1978, 1979, 1983; Mori & Prance, 1981). The terminology used to describe the fruits and seeds of Lecythidaceae is illustrated in Figure 4. A particularly difficult problem in the taxon- omy of Lecythidaceae has been the determina- tion of the extent of intraspecific fruit variation. Unfortunately, earlier monographers of the fam- ily (Miers, 1874; Knuth, 1939) did not have a concept of intraspecific fruit variation. They sim- ply named all fruit variations as separate species. However, beginning with Dugand (1947), bota- nists have become aware that the fruits of Lecy- thidaceae can display considerable variation within the same species (Mori & Prance, 1981). In Allantoma, fruit morphs representing as many as six of Miers’s species have been found on a single tree (Prance & Mori, 1979). Collectors can help in the description of fruit and seed features of Lecythidaceae. They can also gather material that adequately shows fruit vari- ation within and between individuals. It is es- pecially important that the developmental state of the fruits is recorded. Under heat, young fruits may dehisce and thereby appear to be mature. Because it is often very difficult to open up dried fruits, some should be cut lengthwise through the middle and seeds removed from the fruit with the arils and funicles intact. The features men- tioned earlier should be recorded at the time of collection. It is extremely useful to have flowers and fruits turning to the same trees to gather representative material from all stages of the plant’s life cycle. This is especially critical in Eschweilera, where the similarity of the leaves of many species can lead to falsely relating fruits of one species with flowers of another. Collectors can also contribute to the proper correlation of flowers and fruits by always searching in the canopy and under flow- ering trees for old fruits. Ifthe fruits are too rotten to collect, a photograph should be made and sub- sequently affixed to the herbarium collection. However, care must be taken to avoid attributing fruits from the ground to an incorrect tree. If any doubt exists, the fruits should be given a separate collection number. Local collectors can contribute much to our knowledge of Lecythidaceae by becoming ex- perts on the family in their areas. On-site studies of taxonomy, ecology, pollination biology, pop- ulation dynamics, and ontogenetic development in species-rich areas will add much to our knowl- 1987] MORI & PRANCE—LECYTHIDACEAE COLLECTING proximal end pedicel scar infracalycine zone sepal scor supracalycine zone I . A. line of operculor distal end dehiscence calycine ring operculor iie of pos MA ring aril aril C --— f unicle D E F. 6 G H. L. é FiGurE 4. Fruit, seed, and seedling features of New World Lecythidaceae. — A-C. Lecythis ampla Miers. Note ic basal aril in C. —D. Eschweilera Bonpl. Note that this species lacks an aril.— wing entirely surrounds the seed.—G. Un sp. S em with lateral aril. — Winged seed of C pos stellata A. C. a Note that the ilaterally winged seed of Cariniana micrantha D E. Seed of Bertholletia excelsa Humb. & e.—H. Lateral germination of Eschweilera tenuifolia bad Miers.—I. Apical germination of E. pittieri R. Knuth. Modified with permission from Flora Neotropic edge of the family. These studies are needed be- cause many features that allow recognition of species in the field are not apparent in herbarium specimens. For example, in French Guiana, where the senior author has recently completed a de- tailed study of the 27 species of Lecythidaceae in the proposed national park surrounding Saül (Mori & collaborators, 1987), we were able to resolve a number of taxonomic problems that could not be understood with herbarium mate- rial alone. Similar studies would be especially useful in central and western Amazonia and in the Chocó of Colombia. ADDITIONAL NOTES Collectors generally do not provide enough in- formation on habitat, habit, and bark to aid in the identification and classification of Lecythi- daceae. These features are valuable in separating 328 species of Lecythidaceae and should be recorded by the collector. Species of Lecythidaceae are primarily found in lowland moist forests. Nevertheless, a few species have invaded savanna habitats [e.g., Ca- riniana rubra Gardner ex Miers and Eschweilera nana (Berg) Miers of central Brazil and L. schom- burgkii Berg of Roraima, Brazil], and 14 species of Eschweilera are known to occur at elevations above 1,000 m (Prance & Mori, 1979). Within lowland habitats, some species of Lecythidaceae are restricted to the periodically inundated vár- zea habitat where they may be among the most conspicuous elements of the vegetation [e.g., Al- lantoma lineata (Mart. ex Berg) Miers, Couratari oligantha A. C. Smith, C. tenuicarpa A. C. Smith, Eschweilera ovalifolia (DC.) Niedenzu, E. par- vifolia Mart. ex DC., and E. tenuifolia (Berg) Miers]. However, Lecythidaceae are most di- verse in the nonflooded, or rerra firme, habitat. Even within ferra firme some species appear to prefer ridge tops, others hillsides or valley bot- toms (Mitchell & Mori, 1987). Consequently, it is important that collectors make very careful habitat notes, indicating the vegetation type, al- titude, and slope from which collections are made. It is especially useful to note whether the plant grows near a stream or other body of water, and if the water is white, black, or clear. Although all species of Lecythidaceae are trees (some may occasionally grow as shrubs), the col- lector should be aware of and note differences in habit. Some species (e.g., Gustavia monocaulis Mori) are unbranched pachycauls, others [e.g., G. grandibracteata Croat & Mori and G. superba (Kunth) Berg] are branched pachycauls, and oth- ers [e.g., G. hexapetala (Aublet) J. E. Smith and most other species] are leptocauls. Species of Lecythidaceae are found as under- story, canopy, and emergent trees. In order to communicate this information, the collector should record the height and DBH of collections as well as note the stratum to which this and other individuals of the species belong. Only re- productive individuals should be considered when determining the tree stratum in which the species belongs. For detailed studies of Lecythi- daceae, diameter versus height diagrams should be prepared. The graphs will level out at the stratum in which the species reaches reproduc- tive maturity. In his study of Surinam forests, Schulz (1960) has prepared such graphs for a number of species of Lecythidaceae as well as for species of other families. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 The base ofthe trunk should also be described. Some species (e.g., Lecythis idatimon Aublet) have cylindric trunks to the very base, others (e.g., L. chartacea Berg) have basally swollen trunks, and others (e.g., L. zabucaja Aublet) have well-developed buttresses. The height, width, and thickness of the largest buttress should be mea- sured. An undescribed species from French Guiana has pneumatophores. An important but infrequently described fea- ture of Lecythidaceae is the bark. There are four types of external bark morphology in the family. In the first, the bark is very deeply fissured (e.g., Bertholletia excelsa Humb. & Bonpl., Corytho- phora rimosa W. Rodr., and Lecythis zabucaja Aublet); in the second it is nearly smooth (e.g., Eschweilera collina Eyma); in the third it is more or less smooth, but shallow vertical cracks and lenticels may be present [e.g., Eschweilera ped- icellata (Richard) Mori and L. corrugata Poit.]; and in the last it may be markedly scalloped or dippled [e.g., E. micrantha (Berg) Miers and E. apiculata (Miers) A. C. Smith]. The thickness of the outer and inner barks as well as the color of the inner bark should also be recorded. The bright yellow inner bark of Lecythis poiteaui Berg or the flesh pink inner bark of Eschweilera apiculata are excellent aids in field identification. Heart- wood color should also be recorded when pos- sible. One of the best ways to communicate habit and external bark features is with photographs. The photograph should include a scale and the number of the collection and should eventually be affixed to the herbarium sheet. SPECIALIZED COLLECTIONS Known chromosome counts of Lecythidaceae are summarized in Kowal et al. (1977). Addi- tional counts are needed for Allantoma, Carinia- na, Corythophora, and Couratari of the New World Lecythidoideae, for most of the Old World genera, for Asteranthos brasiliensis Desf. (the only member of Napoleonaeoideae in the New World), and for species of Gustavia to determine the ex- tent of polyploidy. Collections of buds of all sizes should be fixed in either Farmer’s (3 ethanol: | glacial acetic acid, v/v) or Jackson’s (4 ethanol: 2 methanol: 2 chloroform: | proprionic acid: 1 acetone, v/v). The best results have been ob- tained with fixation in Jackson’s solution (Ko- wal, pers. comm.). If possible, the buds should be left on the inflorescence, as this facilitates lo- 1987] cation of buds in the proper stage for counting. In addition, because of their thickness, buds should be slit open to allow penetration of the fixative. After 12 hours of fixation the buds should be transferred to 70% ethanol and stored in a freezer when possible. For anatomical studies, collections of leaves, twigs of several sizes (newly flushing ones are especially useful), and bark should be fixed in F.A.A. (10 ethanol: 7 distilled water: 2 forma- lin: 1 glacial acetic acid, v/v). They may be stored in F.A.A. or in 7096 alcohol. The bark sample should consist of a block ca. 2 cm square and should include a portion of the outermost sap- wood to insure that all layers of the bark are included in the sample. Wood samples may be chisled from standing trees or cut from felled trees. In the former case, the block should be about 10 cm square by 10 cm deep and the bark should be left attached to the wood sample. Smaller blocks can be removed but, because of splitting of the sample upon drying, they are not as desirable. The wound left by the removed block should be painted over with a sealer to minimize infection. In the case of felled trees, larger blocks of wood or segments of trunk that can later be cut into specimens about 10 cm long (parallel to the long axis of the trunk) by 5 cm wide (tangentially) by 2 cm thick (radially) should be collected. Specimens should e taken from breast height and from the bole itself rather than from large branches or from the buttresses. The specimens should be dried over low heat to insure that a minimal amount of cracking occurs. There have been no detailed studies of any aspect of the chemistry of neotropical Lecythi- daceae (Prance & Mori, 1987). If collecting ma- terial for study of flavonoids, the collector should remember that the preservatives used in tropical fieldwork (i.e., aqueous mixtures of formalde- hyde or don Be pentachlorophenol in alcohol, or alcohol) may remove these and other compounds and thus render the specimens useless for chemical study. Therefore, if material is collected for chemical study, it should be kept free of field preservatives and air dried. More- over, all field preserved specimens should have the treatment indicated on the label so that the specimens are not subsequently screened for compounds soluble in the preservatives (Coradin & Giannasi, 1980). It is now known that bees play a major role in the pollination of most neotropical Lecythida- MORI & PRANCE—LECYTHIDACEAE COLLECTING 329 ceae. A recent review of pollination in neotrop- ical members of the family is provided by Mori & Boeke (1987). Nevertheless, in order to un- derstand the complex interactions of bees and Lecythidaceae, many carefully documented col- lections of bees visiting Lecythidaceae are still needed. Especially useful are detailed studies of the pollination of individual species of Lecythi- daceae. Collections should be made at all times of the day when flowers are open. Fortunately, most species of Lecythidaceae open their flowers at daybreak and drop them late in the same after- noon. However, some species either flower en- tirely at night or open their flowers during the night. For example, the bat-pollinated Lecythis poiteaui Berg opens its flowers at dusk and drops its petals and androecia at about 0300, and the bee-pollinated Gustavia augusta L. opens its flowers during the night and drops its petals and androecia by late afternoon the next day. During the day, a French Guianan individual of G. au- gusta was visited mostly by trigonid bees, where- as the night-flying bee Megalopta genalis visited its flowers before daybreak. Consequently, it is necessary to determine floral longevity before a pollination study is begun, and once this is es- tablished, the plant must be observed and pol- linators collected at all times when flowers are open. In addition, the species should be studied and pollinators collected throughout its entire flowering cycle. The pollinators at peak flowering may not be the same as those visiting the plant at the onset or the end of the flowering period. For example, Mori & Boeke (1987) have col- lected completely different species of Trigona dominating the flowers of G. augusta during the day at different times during its flowering cycle. We have collected bees visiting Lecythidaceae by simply climbing into the crown and waiting by flowers until insects enter them. Random cap- ture of all insects that approach flowers of Lecy- thidaceae will often give erroneous ideas of the pollinating species because many species of in- sects, such as wasps, visit flowers of Lecythida- ceae to prey upon other insects, not to collect pollen or nectar. In species with zygomorphic flowers, it is especially important to observe the position and behavior of the bee in the flower, since this will indicate what type of pollinator reward the bee is after. Euglossine bees are best captured after they have completely entered the flower; otherwise, they often are quick to spot the movement of the net and escape We use the chlorocresol method described by 330 Tindale (1962) to preserve our insect collections in the field. This method keeps the insects moist and free of mold and allows them to be pinned directly upon their removal from the storage con- tainer. In this method, a teaspoonful of chloro- cresol is placed on the bottom of a plastic con- tainer. It is held in place by a layer of cotton or tissue followed by a tightly fitting layer of card- board or blotter paper. The bees are then placed in layers separated by tissue paper. Labels with the collection data are inserted in each layer. The plastic container is tightly sealed so that the moisture of the insects’ bodies keeps them sup- ple. If the collections are too dry, a little water may be sprinkled into the container before it is sealed. The insects should be pinned soon after removing from the container as they dry out rel- atively quickly upon removal. If the container is properly sealed, the insects will remain in good condition for months. Chlorocresol can be pur- chased from BioQuip Products, P.O. Box 61, Santa Monica, California 90406. CONCLUSIONS complete understanding of the taxonomy and biology of neotropical Lecythidaceae must be based, to a large extent, on adequately pre- pared and documented herbarium specimens. It is not good practice to simply collect Lecythi- daceae without giving consideration to the prop- er preservation of the structures used in the clas- sification of the family. Moreover, it is essential that more detailed descriptions of structures not preserved on herbarium sheets, such as flower color and bark characteristics, be noted by the collector. Better descriptions of the phenological state of the plant and. its habitat are needed as £ well. Sp the study of chro- mosomes, anatomy, chemistry, and pollination biology will add greatly to our knowledge of Lec- ythidaceae. It is also important that local collec- tors become more selective in the species p a a given locality, en therefore, their continued collection adds no new information to our herbaria and only causes un- due expense and takes up valuable space in her- aria. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 LITERATURE CITED Boom, B. M. 1985. Amazonian Indians and the forest environment. Nature 314: 324. CORADIN, L. & D. E. GIANNASI. 1980. effects of chemical preservatives on plant m edis to be used in chemotaxonomic surveys. Taxon 29: 40 DUGAND, A. 1947. Observaciones taxonómicas sobre M Lecythis del norte de Colombia. Caldasia 4: 1-426 TES R. 1939. Lecythidaceae. Pflanzenreich IV. 219a: 1- jt (Heft 105). Kowa, R. R., S. A. Morr & J. A. KALLUNKI. 1977. Chromosome numbers of Panamanian Lecythi- daceae and their use in subfamilial classification. Brittonia 29: 399-410. Miers, J. 1874. On the Lecythidaceae. Trans. Linn. Soc. London 30: 157-318. S. A. Jep 1987. Chapter X. collaborators, The Le- Use ‘of “Swiss Tree Grippers” for making botanical collections JA tropical trees. Bio- tropica 16: 79-80. . BOEKE. 1987. Chapter XII. Pollination. In S. A. Mori and collaborators, The Lecythida- ceae of a lowland neotropical forest: La Fumée Mountain, French Guiana. Mem. New York Bot. Gard. 44: ea G. T. NCE. 1981. The "sapucaia" group of PEH (Lecythidaceae) Brittonia 33: 70-80. & co The Lecythidaceae ofa eee d SR La Fumée Moun- tain, French Guiana. Mem. New York Bot. Gard. 44 —— J. E. ORCHARD & G. T. Prance. 1980. In- trafloral pollen differentiation in the New World Lecythidaceae, subfamily Lecythidoideae. Science 209: 4 3. PRANCE, G. T. & D. G. CAMPBELL. (In pi The present state of tropical floristics. Tax & S. A. Moni. 1978. salone dn ‘the fru and seed of neotropical hie Daria 30: 21-33. NIMES Part I. Fl. 270. Neotrop. Monogr. 21: & 1983. elec and distribution of Lecythidaceae and Chrysobalanaceae. Sonderb. Naturwiss. Ver. Hamburg 7: 163-186. & . Chapter XIII. Future re- search. In S. A. Mori and collaborators, The Le- cythidaceae of a lowland oo n 2 Fumée Mountain, French Guiana. Mem York Bot. Gard. 44: 156-163. SCHULZ, J. P. 1960. Ecological studies on rain forest : northern Suriname. Meded. Bot. Mus. Herb. Rijks Univ. mie 163: 1-267. ie, N. B. 62. The chlorocresol PTE for field St J. Lepidop. Soc. 15: 195-1 CHROMOSOME CYTOLOGY OF OLDENBURGIA (ASTERACEAE— MUTISIEAE)' PETER GOLDBLATT? ABSTRACT As determined from root tip mitoses, three a the four species of the endemic Cape genus Oldenburgia hav et UIC oc to metacentric ch romosomes. Base . In i thé genus IS MCI to be x = is probably a paleotetraploid relict. It has no close relatives in Africa, a arborescent South American genera of Gochnatiinae, perhaps the most primitive related to a group o subtribe of Mutisieae. Oldenburgia is a distinctive and taxonomically isolated, southern African genus of Asteraceae tribe Mutisieae. It is currently being revised by Pauline Bond, and this cytological study was un- dertaken in conjunction with her work, the genus being unknown cytologically until now. O/den- burgia consists of four species, all of which occur in the Cape Province of South Africa, between Grahamstown in the east and the Tulbagh dis- trict in the west. All species are restricted to the nutrient-poor, sandstone-derived soils of the Cape geological system. All species are woody and O. grandis (syn. O. arbuscula) is a small tree. Mu- tisieae are poorly represented in Africa, but rich- ly developed in South America, where several genera are arborescent. MATERIALS AND METHODS Counts were obtained from root tips harvested from germinating seeds. The root tips were treat- ed in 0.003 M hydroxyquinoline for six hours at refrigerator temperatures, then fixed in 3:1 ab- solute ethanol-glacial acetic acid for two to five minutes, and stored in 7096 ethanol. The roots were squashed in FLP orcein (Jackson, 1973) after a six-minute hydrolysis in 10% HCl at 60°C. Vouchers are cited in Table 1. OBSERVATIONS The three species examined have a diploid number of 21 — 36 and similar karyotypes with relatively small chromosomes, 1.5-3 um long. The chromosomes are too small to be easily char- acterized, but submetacentric pairs predominate and there are a few distinct metacentrics. and a similar karyotype of ag irre aad ric Oldenburgia nd it seems i jm most closely DISCUSSION Oldenburgia is a member of the basal tribe Mutisieae of Asteraceae and is usually regarded as belonging to Gochnatiinae, possibly the most primitive of the four subtribes of Mutisieae (Ca- brera, 1977). The genus is characterized by thick, coriaceous, spirally arranged leaves, large radiate capitula, and bilabiate florets with a well-devel- oped pappus. Its affinities within Gochnatiinae are at present obscure. It appears to have no close relationships to other African members of the subtribe (Bond, in prep.), which include Erythro- cephalum, Achyrothalamus, Pasaccardoa, Di- coma, and Ainsliaea. The last two genera lack ray florets (unlike Oldenburgia), Erythrocepha- lum and Achyrothalamus lack or have a reduced pappus (well-developed in Oldenburgia). Pas- accardoa, which comprises a few annual species, comes closest to Oldenburgia in floral characters but is unlike it in vegetative features. It seems likely that O/denburgia is most closely related to a group of largely arborescent South American genera that includes Chimantaea, Cnicotham nus, Pleiotaxis, Gongylolepis, and Wunderlichia, as well as Gochnatia, which also occurs in south- east Asia. There are no counts for any of these genera. Counts for African genera of Gochna- tiinae include n = 12 and 11 for Ainsliaea (Ar- ano, 1957), and there is only a single count for Dicoma, n = 11, for an Indian member of the genus (Bhandari & Singh, 1977). These genera appear distantly related to O/denburgia on cy- tological as well as morphological grounds. Basic chromosome number for Asteraceae is probably x = 9 (Raven, 1975; Solbrig, 1977) and the same number is most likely basic for Muti- ; P by grant DEB 81-19292 from the United States National Science Foundation. ? B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, USA. ANN. MissouRi Bor. GARD. 74: 331-332. 1987. 332 TABLE 1. Chromosome numbers in Oldenburgia. All counts are original in this paper. Localities are in the Cape Province, South Africa. O. grandis (Thunb.) Baillor (syn. O. arbuscula DC.): 2n = 36; cuit ivated at Kirstenbosch Botanic Gar- dens bat Gal- pin s.n. n. (National Botanic Gardens 984/13 in NBG). O. papionum DC.: 2n = 36; Wangenheim farm, Raw- sonville, rock crevices, Bond 1722 (NBG). O. paradoxa Less.: 2n = 36; Robinsons Pass, Oute- niqua Mts., Bond 1726 (NBG). sieae and Gochnatiinae as well. Oldenburgia ap- pears on present evidence to be a paleotetraploid genus probably allied to a group of South Amer- ican genera, mostly of the Guayana Highlands, and geographically isolated in Africa. Its basic romosome number is possibly the same as for the family and tribe. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 LITERATURE CITED ARANO, H. 1957 taxonomic considerations in the t Japanese J. Genet. 32: 293-299. BHANDARI, M. M. & D. M. SimcH. 1977. In IOBP chromosome number reports LV. Taxon 26: 107- 109 . Karyotype analysis and its karyo- ribe Mutisieae. CABRERA, A. L. 1977. Mutisieae — systematic review. V. H. Heywood, J. B. Harborne & B. L. Turner (editors), The Biology and Chemistry of the Com- positae 2: 1039-1066. JACKSON, R. 1973. Chromosomal evolution in Hap- lopappus gracilis: a centric transposition race. Evolution 27: 243-256. Raven, P. H. 1975. The bases of angiosperm phy- logeny: cytology. Ann. Missouri Bot. Gard. 62: 724-764. SOLBRIG, O. T. 1977. Mar aang ag evo- lution in the family Compositae. /n V. H. He wood, J. B. Harborn rner (editors) The Biology and Chemistry of the Compositae 2 NOTES ON CIPURA (IRIDACEAE) IN SOUTH AND CENTRAL AMERICA, AND A NEW SPECIES FROM VENEZUELA! PETER GOLDBLATT? AND JAMES E. HENRICH? ABSTRACT Cipura is a small genus of Iridaceae — Tigridieae that is widespread in tropical South and Central America, Mexico, and the West Indies. It — of interior Brazil. The basic chromosome number in Cipura is x = 7 ave been established here for C. xanthomelas and C. dolre s both senate e 2n = During preparation of treatments of Iridaceae for three regional floras, Flora Mesoamericana, Flora de Nicaragua, and Flora of the Venezuelan Guayana, the genus Cipura Aublet posed several problems in typification and delimitation of species. As a result we studied this small genus in some detail. Our conclusions relate to the ge- nus as a whole and are presented in the form of a review. A new species, C. rupicola, is described from western Venezuela, and two more unde- scribed species are included in a key to the genus but are not published as new species here for reasons given below. A complete revision of Cip- ura is being prepared by P. Ravenna as part of a treatment of Iridaceae for Flora Neotropica. Cipura is one of a distinctive group of New World Iridaceae comprising tribe Tigridieae (Goldblatt, 1982), which is characterized by a bulbous rootstock, plicate leaves, and a basic chromosome number of x = 7. The first genus in the alliance to be described (Aublet, 1775), it was based on C. paludosa, from what is now French Guiana. Cipura comprises five or prob- ably a few more species, distributed from south- ern Mexico in the north to Bolivia, southern Bra- zil, and Paraguay in the south (Fig. 1). Cipura paludosa occurs over almost the entire range of the genus, but the other species have narrower and sometimes very restricted ranges. Cipura is closely related to the larger genus Cypella Herbert from which it differs in having erect inner tepals that partly conceal the stamens and style-stigma apparatus and a large cauline t compri understood. The type species, C. paludosa Aublet, is often treated as including the similar C. c ises at least five species and is at present poorly am- he differences between the two are detailed. A l hromosome leafinserted at the flowering stem apex just below the single or few and closely set rhipidia (spathe- enclosed inflorescence units). As in Cypella and several other genera of Tigridieae, the fugacious flowers have broadly clawed outer tepals; inner tepals with an adaxial nectariferous area on the limb (usually concealed by a fold in the tepal surface); free stamens with weak filaments; and anthers adhering to the style branches. The style divides above into three thickened branches. In the less specialized species there are one or two pairs of erect appendages (crests) which exceed each transverse stigma lobe. Such elab- orate style branches are probably basic in Cipura and the apparently simpler structure in C. pa- ludosa and C. campanulata, in which the style branches are not developed and the crests are absent or reduced, is derived. The species of Cipura currently recognized are C. xanthomelas Martius ex Klatt, of which C. flava Ravenna is almost certainly a synonym; the new C. rupicola, C. paludosa Aublet; and C. campanulata Ravenna. In addition, there are two undescribed species: Cipura sp. 1, a large and violet-flowered species, some herbarium speci- mens of which have been annotated C. formosa Ravenna by P. Ravenna; and Cipura sp. 2, also with violet flowers, from southwestern Venezue- la and adjacent Colombia. In the review we include the following: 1) typ- ification of Cipura paludosa, which involves the delimitation of C. campanulata, a largely Central American species often confused with C. palu- 1 pu by Grant DEB 81-19292 from the United States National Science Foundatio Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box salee. T 299, St Louis, Missouri 3 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. ANN. Missouni Bor. GARD. 74: 333-340. 1987. 334 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 = LJ 200 400 600 800 1000km O 100 200 300 400 600 800 miles \ 100 Geographical ee of the species of Cipura. The occurrence of C. paludosa in Nicaragua, FIGURE 1. iru El Salvador, Guat we dosa; 2) description of C. rupicola and a discus- sion of its differences from the closely related C. xanthomelas; 3) brief discussion of the two vi- olet-flowered species; 4) key to the species; and 5) synonymy and description of C. paludosa. TYPIFICATION AND DELIMITATION OF IPURA PALUDOSA Cipura paludosa, described by Aublet in 1775 from plants growing near Cayenne, is the most widespread and common species in the genus. It has pale bluish, or sometimes white, flowers with a cylindric-fusiform ovary included in the spathes and non-clawed, broad, closely imbricate inner tepals about half as long as the outer tepals (Fig. 2A). Pen = except from above, the sta- mens, stigma. The style branches are hardly Í but the style is thickened above and weakly three-lobed apically (Fig. 2A). The stigmas are simply expanded lobes located opposite and above the anthers which form a tube enclosing the upper part of the style. Ob- scure crestlike appendages extending above the stigmas can sometimes be seen, but they are often microscopic. The inner tepals of C. paludosa are concave below and have a conspicuous central zone of yellow nectariferous tissue outlined in violet. Identification of the type of Cipura paludosa with living populations is not without problems. The type collection was made by Aublet in wet 1987] savannas at the foot of Mt. Kourou in what is now French Guiana. The illustration accompa- nying the description conforms best with blue- flowered plants that are fairly widespread in the Guianas and Brazil, which generally have short stems, relatively broad leaves, and 2-3 flowers in each rhipidium. The flowers are clearly illus- trated by Aublet as having erect and imbricate inner tepals and a style with short stigmatic lobes (flowers are not present on the specimen that we have designated lectotype in the Rousseau Col- lection in the Paris Herbarium). In the type figure the stigmatic lobes are also most unusual. They are drawn as large, ascending and acute struc- tures, quite unlike those in any living plants of any species of Cipura. We assume that these strange structures were included in error. A species described by Kunth (1816) from eastern Colombia, Cipura graminea, is regarded here as conspecific with C. paludosa. The type specimen lacks flowers but has comparatively narrow leaves and elongate included capsules, of which there are two in some rhipidia. The Bo- livian species Cipura major, described by Rusby (1910), and a plant from Peru, C. goodspeediana (Vargas) R. Foster (1962), based on Cypella goodspeediana Vargas (1945), with pale blue flowers and short flowering stems 7-10 cm long, also conform with C. paludosa. Following Foster (1946), we regard both as synonyms of C. pa- ludosa. Cipura major differs mainly in having very narrow leaves, but similar leaves are known on some specimens of C. paludosa. Similar and closely related is a species com- mon in Mesoamerica which almost always has white flowers. When dried it is difficult to dis- tinguish from the often violet- or pale blue-flow- ered Cipura paludosa, with which it was included or confused in the several local and national flo- ras for Mesoamerica. Living plants that we ex- amined display some important differences not apparent in most dry specimens. The inner tepals are initially more or less erect but soon become semipatent and are held well away from the sta- mens and style-stigma apparatus (Fig. 2B). Also, the inner tepals lack nectariferous tissue and, at least to the unaided eye, have no glands. In ad- dition the Mesoamerican species almost always has only one flower in each rhipidium (two or three in C. paludosa), and the flower emerges laterally from the spathes and is thus somewhat secund. The leaves are often very narrow, and the flowers have comparatively short pedicels 5— 7 mm long. Names in the literature that appear GOLDBLATT & HENRICH — CIPURA FIGURE 2. Dorsal views of the flowers (x2) and detail of the stamens and style from the side (x 6), of Cipura paludosa (A), and C. campanulata (B). to apply to this species are C. campanulata Ra- venna (1964), the type of which is from Yucatán, and C. inornata Ravenna (1984) from Caracas, Venezuela. We have been unable to see the types of either of the species described by Ravenna. Holotypes are in the private herbarium of P. Ravenna who is unwilling to loan them to us. An isotype of C. campanulata, cited as at the Kew Herbarium (Ravenna, 1964), has not yet been received there. From the description, Cipura inornata appears to differ hardly at all from C. campanulata and is reduced to synonymy here. Ravenna distin- guished it, although not expressly, by the absence of style appendages but these are also lacking in C. campanulata. Ravenna (1964) explicitly dis- tinguished C. campanulata from C. paludosa by 336 its cernuous (i.e., secund) rather than erect flow- ers and by the inner tepals being nearly equal to the outer, his measurements being 19 mm long for the inner tepals and 21 mm for the outer. We have four living collections of the white- flowered Mesoamerican Cipura and thus are able to assess, to some extent at least, its floral vari- ation. The size, shape, and orientation of the inner tepals are most like the outer tepals of any of the species of Cipura and are about three- quarters as long (Fig. 2B), thus usually shorter by some 4-5 mm and are reasonably close to the dimensions given by Ravenna for C. campanu- lata. Moreover, the flowers are nearly always dis- placed to one side because they emerge laterally from the spathes rather than apically. Thus, there seems little to distinguish this plant from C. cam- panulata and, despite not having seen the type material, we feel that the name applies to the widespread Mesoamerican species. We must note, however, that several specimens from Yucatan that we have seen are somewhat more robust and have broad, strongly plicate leaves that are dis- tinctly striated when dry. Nevertheless, we prefer not to recognize this form as a distinct taxon. Cipura cubensis Griseb., based on Wright s.n., anno 1865, is also white- flowered (Grisebach, 1866) andi d C. paludosa. We have been unable to locate the type, and it is impossible to tell from the pro- tologue whether C. cubensis corresponds better with C. campanulata or C. paludosa, the pres- ence of only the latter being established in Cuba. The name C. cubensis must for the present be excluded. We note, incidentally, that C. paludosa sensu Grisebach which is based on the cited col- lection, Wright 3256, is a species of Cypella. Cipura paludosa and C. campanulata are probably the most specialized and least repre- sentative species of Cipura. They are distinct in their linear to fusiform ovaries, included in the rhipidia, and inner tepals lacking distinct claws. Both species are autogamous, unlike other mem- bers of Cipura, and have broad inner tepals, style branches represented by short stigmatic lobes, and anthers coherent around the thickened upper part of the style. The thickened part of the style probably represents fused style branches, which are present and free in other species of Cipura. Cipura paludosa is apparently a diploid species (Goldblatt, 1982), 2n — 14, although structural heterozygosity and derived numbers of 13 and 12 have been recorded in plants of horticultural origin. The only count for C. campanulata is das separate from ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 tetraploid, 2” = 28 (reported by Goldblatt in 1982 as C. paludosa). Cipura paludosa is the most widespread species in the genus, extending from northern Bolivia in the south to the Caribbean, Mesoamerica, and the West Indies in the north (Fig. 1). It varies in vegetative features. The stem is often quite short, reaching less than 8 cm from the ground, but in some plants may be up to 15 cm long. The leaves are usually relatively broad, 4-10 mm wide, but they may occasionally be only 1-2 mm wide. Its most distinctive features seem to be the number of flowers (and hence capsules) produced in each rhipidium, usually two or three, rarely more or fewer, and the closely imbricate inner tepals with conspicuous nectar guides. In contrast, Cipura campanulata has pure white or rarely pale blue flowers without yellow nectar guides, and the inner tepals are initially erect (parallel to the axis of the flower) but later be- come somewhat outcurving. The style branches and stamens are similar to those of C. paludosa but always. lack appendages. Cipura campanu- lata seasonally wet or marshy sites from Nayarit and Vera Cruz, Mexico through Meso- america to northern Venezuela and Colombia (Fig. 1). YELLOW-FLOWERED SPECIES A yellow-flowered species of Cipura, C. xan- thomelas, was described by Klatt in 1882, based on collections made by Martius and Regnell in Brazil. It is characterized by globose, exserted capsules and yellow flowers. The relatively large yellow flowers have outer tepals 30-35 mm long. The inner tepals are about half as long as the outer, ca. 14 mm long, and have distinct claws ca. 8 mm long with red striations on the inner surface and dark markings on the knee and apex. The style branches are long and have well-de- veloped crests ca. 2.8 mm long. Cipura flava (Ravenna, 1964) appears to be conspecific and pending examination of the type, deposited in the private herbarium of P. Ravenna, we are provisionally regarding it as a synonym. Bulbs of C. xanthomelas are quite large, usually 2-2.5 cm in diameter, with blackish resinous tunics, the spathes are (2.5-)3-5 cm long, and the nearly globose capsules are exserted and typically 12- 14 mm long. The species is known to us from numerous herbarium specimens mostly with poorly preserved flowers and from a living col- lection grown at the Missouri Botanical Garden. The diploid chromosome number in this collec- 1987] tion (Plowman et al. 9306, MO) is 2n = 28, and the species thus appears to be tetraploid. Details of the karyotype of this and the following species will be reported elsewhere. The flowers of our living collection appeared to differ significantly from other material in having partly united fil- aments, a feature that we have not seen in other specimens. Cipura xanthomelas is centered in the Brazilian state of Goias (Fig. 1) and extends into the neighboring states of Mato Grosso, Mi- nas Gerais, Maranhao, and possibly Piaui, while an unusual variant is recorded from western Per- nambuco, adjacent to Piaui. A similar yellow-flowered species, here de- scribed as Cipura rupicola, has come to our at- tention from Venezuela. It has pale yellow flow- ers with unusual inner tepals having slender claws, between which the anthers can be seen, but the long style branches are partly concealed by the erect tepal limbs (Fig. 3). Cipura rupicola differs from C. xanthomelas in being less robust, in its non-resinous bulbs 1.5-2.5 cm in diameter, and shorter spathes 2.6-3.6 cm long. The flowers are also smaller and lack the markings on the inner tepals and the red striations on the claw char- acteristic of C. xanthomelas. The outer tepals are 25-28 mm long and the inner tepals have claws ca. 9 mm long and limbs ca. 8 mm long. The style branches lack crests. Like C. xantho- melas, C. rupicola has exserted capsules, globose in shape and usually 8-11 mm long, thus some- what smaller than its relative. Chromosome number in C. rupicola is 2n — 28 and it thus appears to be tetraploid like C. xanthomelas. Cipura "n Goldblatt & —L sp. nov. TYPE: Venezuela. T.F.A., Dept. Atures: near Puerto D Davidse 4 Miller 26437 (holotype, MO; isotypes, COL, K, NY, VEN). Figure 3 Plantae 12-40 cm altae, tunicis nigribus non resi- nosis, floribus flavis, tepalis exterioribus 25-28 mm longis, interioribus ca. vies = unguibus angustis, filamentis liberis 3 mm lon ntheris mm longis, ramis styli 5 mm a sine ae capsulis obovoideis 8-11 mm longis exsertis Plants 12-40 cm tall. Bulb 1.5—2.5 cm diam.; tunics dark brown to blackish, brittle-papery but not noticeably resinous. Leaves 3-5, all but one basal, strongly plicate, narrowly lanceolate, about as long to slightly longer than the flowering stem, (6-)9-14 mm wide; subterminal cauline leaf sol- itary (second smaller leaf occasionally present), smaller than the basal, 7-15 cm long, about 12 GOLDBLATT & HENRICH — CIPURA 337 mm wide. Flowering stems 1—3 per plant, erect, lateral to the basal leaves, 8-20(-35) cm long, bearing 2-several crowded rhipidia either sessile or on short branches. Inner spathes 2.6-3.5 cm long; outer V»—^A as long. Flowers bright yellow, eae a more or less distinct perianth tube ca. 2 m long; outer tepals 25-28 mm long, oblong, eios slightly near the base, somewhat distally wisted from horizontal, weakly divided into limb and claw, the claw ascending, plane, the margins becoming transparent, the limb horizontal, chan- neled, ca. 20 mm long, to 10 mm wide; inner tepals ca. 18 mm long, the claws ca. 2.5 mm wide, narrower towards the base, ascending and widely separated from one another and thus forming broad windows making the anthers vis- ible, curving inwards above to become horizon- tal at the apex, the limbs erect, imbricate, the apex rolled abaxially, ca. 8 mm long, ca. 8 mm wide, bearing a small circular yellow zone at the base, running on to the apex of the claw. Fila- ments free, erect, thickened below, 3 mm long; anthers ca. 3.5 mm long, free from the style, latrorse, with a broad connective wider than the anther lobes; pollen yellow. Ovary included in the spathes at anthesis, ca. 4.5 mm long; style ca. 4 mm long, dividing into 3 erect branches, these hidden icai by the inner tepal limbs; style branches 5 mm long, plane, truncate, with- out apical appendages, cspanded apically into bilobed papillose stigmas. d from the spathes, obovoid, 8-11 mm long; seeds an- gular, to 2 mm at the longest axis. Chromosome number 2n = 28. Flowering time June-July. Distribution. Venezuela, T.F.A., Dept. Atures, in the vicinity of Puerto Ayacucho, grow- ing on shallow soils of granite hills and outcrops and on the surrounding dry stony savannas (Fig. e POES Additional specimens examined. VENEZUELA. TERRITORIO FEDERAL AMAZONAS: DEPT. AT VEN); bet de las r o Ayacucho, Williams 13099 (K, US, VEN pris medio de las rocas, Puert Ayacucho, Williams 12967 (F, VEN); medio y alto m N of Puerto Ayacucho, Romero 1254 (MO) VIOLET-FLOWERED SPECIES There are two violet-flowered species known to us. One, Cipura sp. 1, fairly well represented 338 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FIGURE 3. Cipura rupicola. Habit (x0. 5) lateral view of the flower (x ve and detail of the stamens and style ein from the side including a dorsal view of the stigmatic region ( x in some herbaria, is native to Brazil, occurring in Bahia, Goiás, and Mato Grosso. It is easily recognized by the unusually large bulbs with res- inous tunics, relatively broad rigid leaves, a short cauline leaf which is typically shorter than the spathes, and only one or sometimes two rhipidia. The flowers are unusual in their violet color and large size. The outer tepals are ca. 40-45 mm long and 30 mm wide, and the inner tepals are ca. 22 mm long, with a long ascending claw to ca. 18 mm long and a limb that curves forward over the claw and then is rolled outwards distally. The style branches are flattened and petaloid but fairly narrow and have two pairs of crests, the adaxial ca. 5 mm long and the abaxial about half as long. The species is known to us only from 1987] dry specimens and details of the inner tepals and style branches are not completely clear. The second violet-flowered species, Cipura sp. 2, of which we have seen only two specimens, is from southwestern Venezuela (Huber 5711, VEN and adjacent eastern Colombia (Vincelli 1230, MO). It also has resinous bulb tunics, but is taller than C. sp. 1, ca. 45 cm high, and has two to five — GOLDBLATT & HENRICH—CIPURA 339 rhipidia at the stem apex subtended by a cauline leaf about three times as long as the spathes. The flowers are much smaller than those of C. sp. 1 with outer tepals ca. 18 mm long. They are com- parable with those of C. paludosa and C. cam- panulata, and it is probably most closely related to these two species. Floral details are obscure owing to poor preservation. KEY TO THE SPECIES OF CIPURA la. Flowers yellow; capsules globose and exserted from the spathes. 2a. Inner tepals yellow with blackish markings on the knee and at the apex, less than half as long as the outer tepals, to 14 mm long . xanthomelas 2b. Inner tepals entirely pale yellow; somewhat more than half as long as the outer, to 18 mm long C. rupicola . Flowers shades of blue to violet or white; capsules cylindric-fusiform or nearly globose; included in the spathes 3a. Rhipidia solitary or two and subtended by a cauline leaf shorter than or about as long as the C. sp. 1 3b. diem usually more than one and subtended by a cauline leaf usually at least twice as long as the spathes. 4a. "Bulbs 15-20 mm in diameter, with tunics not or only slightly resinous. 5a. Flowers white, rarely pale blue; inner tepals without yellow nectar guides or a zone of nectariferous tissue; dd solitary in each rhipidium; inner tepals about 3⁄4 the length of the outer and not i cat campanulata . Flowers usually pale bluish to violet, rarely white; inner tepals with a yellow Pius guide and a zone of nectariferous tissue, often outlined in violet; flowers (1—)2-3 in each rhi- — c tA I" ; inner tepals closely naa about half as long as the outer „u C. paludosa C. sp. 2 pidium 4b. Bulbs about 30 mm in diameter, with tunics heavily resinous NOMENCLATURE AND DESCRIPTION OF C. PALUDOSA Cipura saa Aublet, Hist. Pl. Guiane 38-39. 1775. TYPE: French Guiana: near Mt. Kou- rou (as a. Aublet s.n. (lectotype, P— Herb. Rousseau, designated here). Figure 2. E a d din Kunth, Nov. Gen. Sp. 1: ene 1816. bem San- Í m del Angostura, Humboldt & Bo npland S.A. atin cad E d DE ted here; isolectotype, P— Herb. Bon Cipura major cenas "Bull. New York Bot. Gard. 6: 493. 1910. TYPE: Bolivia: Tamupasa, 1,800 ft., Neila 546 (lectotype, NY, designated here; Cypella and — Revista Univ. Cuzco 71. 944. TYPE: Peru. Convencion: Hda. Po- trero, Vargas 2509 due ype, CUZ, not seen; iso- ue , GH); Cipura goodspeediana (Vargas) R Foster, Rhodora 64: 31 Cipura paludosa subsp. mexicana Ravenna: Phytolo- 5-1 : Mexico. Sinaloa: joe 278 (holotype, Herb. Ravenna, not seen). Plants 16-27 cm tall. Bulb 12-20 mm diam .; tunics dry and brittle-papery, rarely slightly res- inous. Leaves finely plicate, linear-lanceolate, 1— 3 basal, 16-27 cm long, 2-5 mm wide; subter- minal cauline leaf single, largest, clasping the spathes below, 16-23 cm long. Flowering stem 3.5-8(-15) cm long, bearing several crowded rhipidia, either sessile or on short branches. In- ner spathes 30-35 mm long; outer ca. 73 as long as the inner. Flowers pale to bright blue, or white, the tepals connate into a distinct tube ca. 2 mm long; outer tepals fading to white at the base, oblanceolate, ascending to nearly horizontal, dis- tally twisted ca. 30°, the margins rolled outward, the right more so than the left, 2.5—2.8 cm long, 1.3-1.5 cm wide, with a white nectary in a central zone ca. 1 cm above the base; inner tepals 1.5 cm long, 8-10 mm wide, light blue with a white apex, darkest at the margins, more or less ob- ovate, erect and imbricate, the apex rolled out- wards, strongly concave, with a yellow nectary in a median band extending ca. 2 mm from apex to 2-3 mm from the base, broadening to 4 mm at the widest, surrounding a white to light blue, raised nonglandular area, sometimes with semi- parallel dark blue bands extending at right angles to the nectariferous zone. Filaments 2-3 mm long, thickened and sometimes contiguous at the base for up to 1 mm, free and threadlike above; an- thers adhering to the style, 3.5-4 mm long, la- trorse with a slender connective broadest at the base and tapering to the apex, pollen white. Ovary 340 included in the spathes, 6-10 mm long; style 6— 8 mm long, cylindric, 3-lobed above, the lobes usually emarginate, | mm long, 1 mm wide, dis- tally papillose, with 1—2 obscure (often micro- scopic) erect, crestlike appendages at the base. Capsules oblong-cylindric, 12-18 mm long, in- cluded in the spathes; seeds angular, ca. 1.3-1.8 mm long. Chromosome number 2n = 14. Distribution. Bolivia and Paraguay, through southern Brazil to Central America, southern Mexico, and the West Indies (Fig. 1 LITERATURE CITED AUBLET, F. 1775. Histoire des Plantes de la Guiane ? "Studies i in the flora of Bolivia — r. Gray Herb. 161: 3-19. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 GOLDBLATT, P. 1982. Chromosome cytology in re- lation to suprageneric eae of neotropical ridaceae. Syst. Bot. 7: 1 GRISEBACH, A. 1866. Catalogus Plantarum Cubensis ip ' Erganzungen und Berichtigungen KLATT, F. W. zu Baker’ s Systema Iridacearum. Abh. Naturf. Ges. Halle 15: 337-404. KuNTH, C. S. 1815 [1816]. Voyage m Humboldt et Bonpland. Sixiéme Partie. Botanique. Nova Gen- era et Species Plantarum (= N s Genera e Species Plantarum quas . . .), Volume 1. Libraire Grecque-Latine-Allemande, Paris. Gide Fils, Paris. 1964 a i l . (Bu otes on ligase IV. Phytologia 56: 193-195. RusBv, H. H. 1910. New species from Bolivia col- lected by R. S. Williams— 1. Bull. New York Bot. Gard. 6: 487-517 ITHOMIINAE (LEPIDOPTERA: NYMPHALIDAE): SUMMARY OF KNOWN LARVAL FOOD PLANTS! Boyce A. DRUMMOND III? AND KEITH S. BROWN, JR.* ABSTRACT he known interactions between the larvae of ithomiine butterflies and their host plants (about 400, 90% in the Solanaceae) are described in a table, illustrated, and briefly discussed. The widespread and diversified use of plants in the family Solanaceae by man is reflected in the large number of applied scientific papers pub- lished on these plants (see the taxonomic index of any issue of Biological Abstracts). The alka- loidal nature of most plants in this family has led to their extensive use in folk and proprietary medicine, consciousness expansion, and recently as a source of pharmaceutical intermediates. Other important uses include fodder, fencing, support, insecticide, ornament, and perfume. In tropical America, the most important groups inae. These insects overcome t physical and chemical defenses of these plants and turn them to their own use, at least as rec- ognition cues, if not necessarily as protection against predation (Brown, 1987). Useful Sola- naceae frequently attacked by Ithomiinae in- clude Lycopersicon, Cyphomandra, Solanum sect. x el d and solasodine-producing or tobac- tituting Solanum. Also regularly eaten are E Eun tuberosum, S. melongena and relatives, as well as Capsicum and Physalis. Many orna- mental and medicinal Solanaceae (Brunfelsia, Cestrum, Solandra, Markea s.l., Juanulloa Brugmansia, Acnistus, Solanum pseudocapsicum and Solanum sect. Jasminosolanum) are heavily damaged by ithomiine larvae. Nicotiana and Pe- tunia seem to be immune to these herbivores. This paper lists the known i t ti (to mid- 1985)1 butterflies and their lar- IÇ ! This paper was part of the Second I val food plants, 90% in the Solanaceae (Table 1). It is the data base for papers by Drummond (1985), Brown (1985), and Brown & Drummond (in prep.). The 40 butterfly genera for which food plants are known or inferred are placed in phy- logenetic order in the Table, grouped into tribes as first proposed by Fox (1961), followed basi- cally by Mielke & Brown (1979), and modified by recent studies of early stages leading to a nu- merical phylogeny by Brown (in prep.). Nomen- clature for the butterflies follows Mielke & Brown (1979) except in a few cases in which recent stud- ies, especially of chromosomes, indicate changes in status. Nomenclature of the plants follows the thesis of Mary Fallen (1983, Hamburg) for Apo- cynaceae, recent compendia of the Solanaceae, and the Index Kewensis. We believe that iden- tifications of both insect and plant taxa are ac- curate to the generic level in all cases, to the subgeneric level for Solanum in essentially all cases, and to the species level where given in the vast majority of cases. Many older records, either not confirmed or regarded as unlikely in view of broader recent studies, have been excluded from the list; these include especially those in the ag- ricultural literature of Brazil, compiled in D’Ar- aujo e Silva et al. (1968) and continuing up to the present day [such as a recent report of Me- chanitis lysimnia nesaea Hübner as a pest of Pas- siflora edulis Sims. in northeastern Brazil, clearly a misidentification of Heliconius ethilla narcaea (Godart) or Eueides isabella dianasa (Hübner), common in the area]. Doubtful records are marked with a question mark in parentheses, on the Biology and Systematics of the Solanaceae presented at the Missouri pedum Garden on 3-6 August 1983. ? We are grateful to W. H. H J. R. Trigo, R. F. Monteiro, G. pia mas M., G. Small, W er, J. Vasconcellos-Neto, L. E. Gilbert, S. Knapp, J. Mallet, Renata S. C. Dias, . W. Benson, A. H. Watson, Condorcet Aranha, Lucia F. d'A. Carvalho, F. Fernández Yépez, and W. G. D’ Arey a information on ithomiine host plants, discussion of the patterns revealed here, and identification of organ 3 Pikes Peak Research Station, Colorado Outdoor “upan sN Center, Florissant, Colorado 80816, U.S.A. Please request reprints from this address. 4 Departamento de Zoologia, Instituto de Biología, Universidade Estadual de Campinas, C.P. 6109, Campinas, Sao Paulo 13.100, Brasil. ANN. Missouri Bor. GARD. 74: 341-358. 1987. 342 whereas sure records with tentative (uncon- firmed) plant identifications are indicated by a simple question mark after the name. Localities and sources are coded and given at the end of the table. Numbers or letters in parentheses after a plant name are voucher symbols for that species. The following genera (with number of species in parentheses) of Ithomiinae have yet to be ob- served or suspected as larvae on any plants; from preliminary field abana they are predict- ed to use the sola s genera indicated in each case: Raswelia tit apsicum UM. Pa- tricia (2) (Dunalia), a new genus near Hyposcada (1) (Lycianthes). Paititia (1) (Cyphomandra), Ar- emfoxia (1) and Pagyris (1) (Witheringia, Du- nalia, Brugmansia, and relatives), and Dygoris (1) and Veladyris (1) (Solanum sect. Geminata, Cestrum). The following genera of Solanaceae, with one or more species available to Ithomiinae in their tropical or subtropical moist habitats (genera re- stricted to dry habitats or temperate zones not included), have not yet been seen to be used by any species of Ithomiinae. Solanoideae: Jalto- mata, Athenaea (expected for Epityches and Ith- treae); Nicotlana. Petunia, Fabiana, Nierember- gia, Bouchetia (Nicotianeae); Schwenkia, Protoschwenkia, Melananthus (Schwenkieae); Parabouchetia (Parabouchetieae); Leptoglossis, Browallia, Streptosolen (Salpiglossideae), Het- eranthia (tribal position unclear). While some of these genera may be found to be used by Ith- omiinae with more observation, members of others have been watched for years within large Ithomiinae communities and have not been seen to support the larvae; in a few cases, ovipositions were followed by larval death (e.g., in Campinas, SP, Mechanitis polymnia casabranca Haensch has oviposited on both Capsicum annuum and ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Nicotiana sp., but the larvae died without feeding or developing). The complete picture of the known food plant e of the Ithomiinae, based on the data n the table, is presented in Figure 1, in which Dian: genera (from top to bottom) and butterfly tribes (from left to right) are arranged in phylo- genetic order based on currently accepted evo- lutionary sequences (the vertical position of but- rfly genera represents only convenience in presentation of the figure; details are given in the phylogeny of Brown, in prep.). The complexity of the relationships illustrated in the figure in- dicates that strong ecological influences may out- weigh the presumed evolutionary history of the interactions. See papers by Drummond (1986) and Brown & Drummond (in prep.) for discus- sion of these aspects. c O LITERATURE CITED Brown, K.S., JR. 1987. Chemistry at the Solanaceae/ Ithomiinae Interface. Ann. Missouri Bot. Garden 74: 359-397. Phylogeny of sud Ithomiinae (Lepidoptera: Nymphalidae). (In pre & B. RUMMOND TIT. Biochemical and phy- logenetic coevolution: evaluation of the Solana- Miri one oo (In prep. D'ARAUJO E SILV WG E D. M. GALvAo, A. i L. GONÇALVES, b SILVA & L. SIMONI. 1968, ieee aes ‘dos Insetos que Vivem nas Plantas do Brasil—Seus Parasitos e Predadores. Ministério da Agricultura, Rio de Janeiro, 4 volumes. DRUMMOND, B. A., III. 1986. Coevolution of ith- omiine butterfli d sol pl t Pp. 307- 327 in W. G. D'Arcy (editor), The Solanaceae: Biology and Systematics, Columbia Univ. Press, New ; Fox, R. M. 1961. A checklist of the "ergo " Tribes Tithoreini and Melinaeini. J. Lepid 3. MiELKE, O. H. H. & K. S. BRowN, JR. 1979. Suple- mento ao “Catalogo dos Ithomiidae Americanos" de Romualdo Ferreira D'Almeida (Lepidoptera: Nymphalidae: Ithomiinae). Univ. Federal do Pa- raná, Curitiba. 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VADUD] os niduu2u11M JeW-VA USA JoyIzunyY (u1uny) india Dis24]0n;) “MOH ?9 "ÁpIqg Psspupndi psspupunjdi HA wo Joxyrzung (PUNY) 24india 718241007) Inq P Ng vjoulsnjd n142]22 aa wry (LZEL) ds DisaajonD yosusey p7/714ptup ,90INOS ,AM[?20] satoadg pue (uorjoog IO snuasqns) snusy soroodsqns pue 'saroods 'snuar) ‘QUL ;Sjue[d 1SOH :Əpuuuuoull ‘ponunuod “| #slgV L ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 354 ado SOU (¿) “wey wnijofiaquaásis (uunuowm1s01d2) unuvjos 8x duo "7 wumoisdpoopnesd (voisdpoopnasq) tunup]oç €x ory ¿ds (pipunuay) ununjos ay Ider 'dureo `HƏA 2unanappo (vipuun) tunup]osç (zig) papuawtuy DaDUaWAY DIZMA aq gi (0€€1) ‘ds (DIDUNUAD) wnuvjog (qu»suseg) uətu41 8x Sd Iou]pusgs wWnsO/fixD] (p3punua2)) uunup]oç ppi9ur[V.Gd ?? 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BROWN, JR.? ABSTRACT The secondary chemical constituents of 42 species in sixteen genera of Solanaceae, of five species in three genera of Apocynaceae- Ris amp p of 142 species in 45 genera of Ithomiinae (Lepi- doptera: Nymphalidae) whose larvae feed o ese plants, have been analyzed and compared in a standardized manner. Large orb spiders (Nephila clavipes), which cut field- caught adult Ithomiinae out of their webs, were used to assay fractions present in or derived from larval food plants; the fractions were lied externally to the palatable n uiii butterfly Biblis hyperia. All extracts and fractions from Solanaceous plants were inactive, not defending Biblis against predation by Nephila. The — use defensive compounds of adult Ithomiinae (also found in s and occasionally in Apocynaceae u y some Ithomiinae larvae) are dehydropyrrolizidine alkaloid (PA) monoesters and their N-oxides, stored in up to 2096 of dry weight. These compounds are obtained from a variety of natural sources by the adults after emergence from the pupa (freshly hatched individuals of both sexes in 26 genera were readily eaten by Nephila), especially from de- composing Boraginaceae-Heliotropoideae and flowers of Compositae-Eupatorieae, for which they serve as pollinators. onus Adenostemma, and many Eupatorium flowers were confirmed as rich sources ofthe alkaloid mutualism with the cim pollinators. The highly diversified and variable Solanaceae toxins. seem not to be stored and used for defense by the Ithomiinae, but may be important in mediating larval feeding preferences and oviposition in the butterflies, which show appreciable chemical specificity in host plant preference. Patterns of ithomiine | larval use of oe Solanaceae at the generic level, however, fthe oups. An not p to sequential adaptation by radiating lines of Jibi. to dnm classes of chemical toxins in the already diversified Solanaceae. By obtaining their PAs, necessary for their defense and reproduction (pheromone synthesis), from a source stabilized by mutualism, the Ithomiinae avoid dependence on the Solanaceae chemicals, constantly destabilized by divergent selection in the antagonistic system. Quantitative determination of PA concentrations in parts of different Ithomiinae and We: dian permits the drawing of a diagram for the flow of these substances in natural ecosystems (April 1983), at the 35th and 36th Annual Reunions of the Sociedade Brasileira para o Progresso da Ciéncia (Belém, Para, July 1983 and Sao lees July 1984), and at the first meeting of the International Society of Chemical Ecology (Austin, Texas, June 1984). For a preliminary s see Brown (1984); detailed analysis of the insect side of the interface is presented in Brown (19 85). 2 I am especially grateful to Prof. Joao Vasconcellos-Net o for tł tal biological background of this paper was read and criticized by T. M. Lewinsohn, W. W. Benson, L. E. Gilbert, M. Rothschild, and J. A. Edgar. R. F. D’Almeida, O. H. H. Mielke, and Gerardo Lamas M. stimulated my interest in the Ithomiinae, and special attention to the interface with ree was infused by W. W. Benson, L. E E. Gilbert, and - A. e, e were most helpful with systematic aspects oe the "Solanaceae an nd H. F. Lei tao Fo with. the Compositae. E Aparecida Henriques assisted greatly in extraction and fractionation (Table 3), and José Roberto Trigo and Renata S. C. Dias helped in field study of the Ithomiinae/Solanaceae interaction. T. Eisner provided information and reprints about the Utetheisa/PA/Nephila interaction first studied by him and his students in Florida. R. Forster and D. A. Monteiro facilitated access to field sites in the Fazenda Santa Elisa, Instituto Agronómico de Campinas. A. and C. Kascheres provided important services in NMR and mass spectra at the Instituto de Química, UNICAMP, and I. Valio and L. Sodek permitted extensive use of the Micronal colorimeter in the Departamento de Fisiologia Vegetal, UNICAMP. Chemistry of the Solanaceae was lucidly presented to me by W. Evans, K. Schreiber ; D. Lavie „and J. G. Roddick. I especially thank L x. Brower, J. A. Edgar, and M. Rothschild for helpin nd bioassay techniques. FAPE contributed to the original installation of the Laboratorio de Ecologia Quimica (project 74/0544) and Rhodia NCAMP to its reinstallation. The CNPq provided a research fello 3 Laborat torio de Ecologia Quimica, Departamento de Zoologia, Instituto s COE Universidade Estadual de Campinas, C.P. 6109, Campinas, São Paulo 13.081, Brazil. ANN. MIssouRI Bor. GARD. 74: 359-397. 1987. 360 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 CLAY 07 oO "o A. R*H solamine ReCOOEt solaurethine C " m ed ñ R=COCH=CH(CH2) 12CH3 K. l-acetyl-B- z etaine i carboline-3- carboxylic solapalmitenine G. anatabine* B. cuscohygrine A083 N LA, L—- Nis. C, tigoidine 8 ad ° pesos un N. capsiamide* ^ C P. E C2 hyoscine d ' J D. withasomine E. hopeanine?? B. lupanine M. capsaicin* [ORNITHINE] E bim LALIE j FIGURE |. Representative alkaloids known from the Solanaceae, with probable biosynthetic pathways; see Table 2 for occurrence in genera. Asterisked alkaloid types are not known yet in leaves of natural Ithomiinae host plants. Plants in the cosmopolitan family Solanaceae (Brown, 1979, 1987; Brown & Benson, 1974). are widely known for their toxic qualities derived They even pull in other less abundant and per- from an exceptionally diverse suite of alkaloids haps less protected, but typical aposematic but- (Fig. 1) and steroidal bitter principles, terpenes, terflies such as Heliconiini (Brown, 1972a; Brown saponins, and phenolic glycosides (Fig. 2; Evans, & Benson, 1974), Danainae-Itunini (Brown, 1979; Schreiber, 1979; Harborne & Swain, 1979; 1987), Acraeinae (Brown & Benson, 1974), and Kirson & Glotter, 1982). Such a rich larder of Troidini (which are usually central models of potential poisons could be expected to repel, de- their own mimicry rings), as well as myriad ter, or intoxicate almost all herbivores while Batesian mimics (Satyrinae, Nymphalinae, causing a few to become specialists; the special- Charaxinae, Riodininae, Pierinae, Dismorphiin- ists might be predicted to sequesier the Solana- ae, Hesperiidae, Castniidae, Geometridae, Diop- ceae poisons and use them in their own defense tidae, Arctiinae and Pericopinae— the last two against predators. In fact, the herbivorous insects possibly distasteful Müllerian mimics— and found on Solanaceae leaves in the Neotropicsare members of other insect orders such as Odonata often restricted to the family and some of them and Homoptera). Adult Ithomiinae were in- are brightly colored as ifto suggest unpalatability volved in the original proposals of Batesian and (Table 1). These form ideal systems for the in- Müllerian mimicry and have been shown to be vestigation of biochemical ecology and coevo- rejected by a variety of vertebrate predators lution (Brown, 1980). (Bates, 1862; Belt, 1889; Brower & Brower, 1964; One of the most important groups of Solana- Haber, 1978; Coimbra-Filho, 1981). One known ceae herbivores in the Neotropics is butterfly lar- exception (Brown & Vasconcellos-Neto, 1976) vae of the nymphalid subfamily Ithomiinae involves complex learning behavior by a single (Drummond & Brown, 1987). The brightly col- tanager population (Pipraeidea melanonota), ored adults are regarded as prime distasteful ^ which squeezes out fatty abdominal contents of movers in regional insect mimicry complexes individuals in winter reproductive diapause dur- 1987] STEROIDS Q. BITTER PRINCIPLES Q4 withaphysalin A SAPONIN OH R. hispinin B ! H Ò- Rha-Rha U. solanesol | BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY S. perulactone POLYPRENE 361 TERPENOIDS PHENOLIC GLYCOSIDES . DITERPENES š i i OH ë H W. cestric acid MI raimonol c M 4, I uo didum ad uerci rutinoside ` has TRITERPENE Z. PUNGENT OILS (unknown o T. lanosterol composition) EN H,OH IGURE 2. Examples of non-nitrogenous secondary compounds found in Solanaceae used as larval hosts by Ithomiinae. Carote See Table 2 for distributi on ing the cold of early morning, treating them as it does fruits with a bitter rind. Similar learned behavior has been observed in jays (Corvidae) eating ithomiines in Costa Rica (R. Hagen, pers. comm.), as well as in orioles and grosbeaks at- tacking wintering monarchs (Danaus plexippus) in central Mexico (Calvert et al., 1979; Fink & Brower, 1981). Like most aposematic insects, Ithomiinae have tough and resilient bodies; sur- viving individuals squeezed by the tanager were often captured in flight one or more days after attack. They often remain on the forest floor for weeks after death, avoided by predatory and scavenging ants. Even the giant tropical orb spi- der Nephila clavipes, which often clutters the air- space of ithomiine colonies with broad sticky webs and takes most aposematic butterflies with typical rapaciousness (Vasconcellos-Neto & Lewinsohn, 1982, 1984), cuts out all Itfhomiinae from its web rapidly (10 sec.-2 min.) after con- tact with any part of the body or wings. Other spiders, especially flower-frequenting Thomisi- dae (crab spiders), may take Ithomiinae regular- ly, however (Drummond, 1976; pers. obs.). It has been suggested frequently that the pro- tection of adult Ithomiinae against predators is noids in other pigments are not included; for flavonoids, see Harborne & Swain (1979). due to alkaloids or other toxic chemicals se- questered and stored by the larvae from Sola- naceae and passed on to the adult (Brower & Brower, 1964, p. 154; Young, 1972, p. 291; Drummond, 1976, p. 268, 1981, p. 63; Brown, 1980). A good precedent for this suggestion exists in the storage of cardiac glycosides (Fig. 3A) by larvae of Danainae—sister-group to the Itho- miinae (Ackery & Vane-Wright, 1984)— which are transmitted to adults and help protect them against avian and other predators (Brower et al., 1967; Brower, 1969; Brower & Glazier, 1975). Nevertheless, no evidence has been obtained yet for the presence of any Solanaceae secondary chemical sequestered naturally into the tissues of any herbivore. Indeed, all results reported by Rothschild (1973) showed metabolism and ex- cretion of Solanaceae alkaloids by specialist her- aes a id experiment showed retention of Manduca sexta (tobacco hornworm) (e up et al., 1979), but this herbivore does not normally encounter these insecticidal com- pounds in nature. Since Solanaceae alkaloids and steroids are relatively stable compounds, abun- dantly available to Ithomiinae through the larval food plant and toxic enough to be eminently suit- [VoL. 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN 362 '$210A1q19Q Ə} 1nq 'sjue[d ou YoeNe 1ou op AJY} :2eooeue[os Surl[oned punoj UIO are s10jepoud pruro1eluəq Xue e 10N 1q8rT 9]&Jopo]JA 91e €19U28 [LIIAS sJ9y10 pue 3eprzAÁuloJdy VH3.ldld 9I? 3ulog ÁAvOH 9]&Jopo]JA 1uonboai DIIAT ‘vinvo1dq deploys Ajoley 9]&JopoJA MOT SIOMOY *21e eioues Kue]A oeproqeieog ÁpIey 14311 HLIIPOWN juonbouju] ?louod 9 9?uruoo[) *oeuuopÁuyoeig :aepruornoun;) sok 91€&J9poJA 9]eJ9poJA juonboJju puyoojidy oepi[[2ur220)) 3I? 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VH3.LdOWOH ole IUWOŞ 14311 usrH juonbouju] g13U33 AUREN oeprisur SOX 14317 y3tH 1uonbogju] DIGQIUNY 3&pruiojejuad sox 1ugr] y3 o1 JIBIOPO|P 1uonbouju] sepuoooulLg + oepioeaAT 9I? 1S0]A 93]eJ9poJA Urdu juonboiq snjouojasdA Hy ‘piyiyd IpI 91? Əuros NLIIPOWN y3IH quonbel4 IPUIN Vad ldIWaH A[aiey 9jEJopoJ( «=: O}VIOPOUW! ol MOT 1uənbəsu Sar[rurgj SnOLIEA VHAIdONVSAHL ore AUR ÁAE?9H 91&Jopoul 01 MOT 1uonboiq S19Q10 pue S142D1404Q.) IEUIZ[BWIOY :2PprpuoVv Vaud LdOHLYO ionewosody sjue|d (A27 “ds) Əgəopgup|os gau Áprurgjqng pue Xu. 19pJQ) 199su] 01 o3eure(T uonezi[eroodg uo 29uaLmoo(Q Jo 90139q ((8961) [seg Op sejue[d seu warg onb soləsul sop Zoo OLEND “TE 19 PA[IS 9 OMELIY,P `D `V pue suongAJəsqo [euos1od :I0INOŞ) “[IZeIg ut (pupt1031N ueu 13410) ILILULJOS o1 pəztptoəds S2JOAIQIOY JURLIOdWY '[ 718V L 1987] BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY 363 INSECT DEFENSIVE COMPOUNDS MEC T INSECT MALE PHEROMONES i ead ^ x CARDENOLIDES "iaispencii UR MM lactone" alkaloid monoesters A. Calotropi E. lycopsamine T sclepias "i i . i F 9 CH3 F. (epi-e) intermedine G. (7«-0H) echinatine a OAc š : : H. (7«-OH, epi-«) rinderine I. (epi-«e,8) indicine li. Danaidone H202 9 CHO oleandrose-O (0.15%) “a H I H MeOH u oc glucose 100° 7 , z Ps P n 6 N ¿ B. Urechitoxin (Urechites) n m H Ac.0 : = eroe 3 MN. Án so qo ie J. Echinatine N-oxide acetate : MS NH, 100° K. (epi-7,0,@) Indicine N-oxide ^ APOCYNACEAE TOXIN OO HO CHOR Pon CO 2% BF lal Soe CHZ0H NH 2 CO co (CH) "Hac 3-hydroxy-L- 579. 5 min cH Ehrlich C. kynurenine MATTOCKS- f =a 0. oqui u BINGLEY " pro ins oss) (Ithomiinae) ASSAY N is 562 n nm ——— asa Danaine and ithomiine ED and defensive compounds, including cardiac glycosides (A, able for protection against predators, their ab- sence in Solanaceae-feeding insects was quite a disappointment (Rochschild, 1973). The only known “secondary chemicals" isolated from the Ithomiinae before this work were the amino-acid yellow pigment 3-hydroxy-L-kynurenine (Fig. 3C; Brown, 1967; Brown & Domingues, 1970) and a Sermon lactone related to some es- terfving acids of pyrrolizidine alkaloids (Fig. 3D), as well as the presumed precursors of the latter (Fig. 3E, F) (Edgar et al., 1976). Especially interesting to find in adult Ithomi- inae were these latter compounds (dehydropyr- rolizidine alkaloids, or PAs, Fig. 3E-K), since they have been shown to be very important in both defense and reproduction of the butterflies in the sister group Danainae (Edgar & Culvenor, 1974; Schneider et al., 1975; Schneider, 1977; Rothschild & Marsh, 1978; Boppré, 1978; Edgar et al., 1979; Edgar, 1982). Unlike the rather spo- N), ware pen Men alkaloids (E-I) and their N-oxides (J, assay, and deplete toxins (B, E, radically stored cardenolides (Rothschild et al., 1975; Dixonetal., 1978; Malcolm & Rothschild, 1983; Fig. 3A, B), PAs are universally present in field-captured adult Danainae, which actively seek them out and sequester them (Boppré, 1984) from flower nectar, plant exudates, and even oth- er insects (Bernays et al., 1977). Some danaines (Euploea) also obtain PAs from apocynaceous larval food plants (Parsonsieae: Parsonsia; Ed- gar, 1982). Larvae of a primitive ithomiine iso- lated in the New Guinea region, Tellervo zoilus, feed on these same plants; adults contain PAs presumably derived from Parsonsia (Edgar, 1982), as do adults of a primitive Central Amer- ican ithomiine (Tithorea tarricina) whose larvae feed on Prestonia (Apocynaceae: Parsonsieae) containing the same PA found in the butterflies (Fig. 3E; Edgar, 1982; Edgar & Harvey, in prep.). The purpose of this paper is to examine in detail, by chemical analysis and field bioassay, 364 three basic questions related to the specific in- teraction between Neotropical Ithomiinae and their Solanaceae larval host plants. These ques- tions derive from conventional aspects of insect- plant coevolutionary theory, especially as ap- plied (with much success) to aposematic insects on poisonous plants (Ehrlich & Raven, 1965; Benson et al., 1976; Brown, . (1) To the extent that host plant specificity may exist in the Ithomiinae/Solanaceae relation- ship at generic and specific levels, how does it relate to chemical mediation of cues for ovipo- sition and larval feeding? (2) How might such chemical specificity re- flect mutual interaction of these two groups over long evolutionary time, leading to diversification in both groups as a function of reciprocal selec- tive pressures—that is, what is the evidence for coherent patterns of biochemical or “‘parallel di- versification” coevolution? (3) To what extent do the Ithomiinae use the poisonous secondary chemicals of their Solana- ceae larval hosts for their own defense against predators in any stage of their life cycle? A preliminary phylogenetic (Brown, in prep.) and chemical (Brown, 1984, 1985) survey of the Solanaceae/Ithomiinae interface, accompanied by an efficient spider bioassay for predator-de- fense compounds, has indicated surprisingly that, while the first question seems to merit an affirm- ative response, the other two questions must be answered by “little or no evidence in favor." The Ithomiinae apparently colonized the Solanaceae well after the generic diversification of this plant family in the New World, tolerating and using the diverse secondary chemicals (Figs. 1, 2) for oviposition cues to regulate specificity, but not for defense. Efficient chemical protection of adult Ithomiinae is generally unrelated to toxins en- countered by larvae but not stored; instead, the adults seek out PAs in flower nectar and decom- posing leaves, using them not only in pheromone synthesis (Fig. 3D) but also in defense, exactly as in the Danainae. The results of this survey are plant specificity, and alternative ecological and physiological factors in the rela- ionship MATERIALS AND METHODS STUDY SITES Field observations of interactions between So- lanaceae and their herbivores were undertaken ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 principally in two very different artificial tropical forest systems in the Fazenda Santa Elisa, Insti- tuto Agronómico de Campinas, Sao Paulo (600 m elevation): Monjolinho (a forest garden of na- tive plants) and Amarais (an old eucalyptus for- est with natural under- and middlestory) (see Brown et al., 1981, for photos, maps, and de- scription). Supplementary observations were made in Sumaré, SP (similar to Amarais, 550 m); Martinho Prado, SP (floodable riverine for- est, 400 m); Serra do Japi, Jundiai, SP (native seasonal and montane forest on rocky soils, 850- 1,150 m); Serra Negra, SP (humid montane for- est, 1,000 m); Bertioga, São Vicente and Mon- gagua, coastal Sào Paulo (perhumid tropical for- est, near sea level); Pocos de Caldas, Minas Gerais (montane forest, 1,200 m); coastal Rio de Janeiro and Espírito Santo (disturbed tropical rain forest, sea level to 800 m); Goiânia, Goiás (riverine thicket, 800 m); various parts of Rondónia, SW Brazil (seasonal forest on rich soils, 100—650 m); various parts of NE and S Pará, Brazil (tropical forest, sea level to 800 m); the region north of Manaus, Amazonas, Brazil (disturbed tropical rain forest on poor soils, 50 m); and in northern Venezuela (deciduous and cloud forest, sea level to 1,400 m). Some limited data have also been obtained in Mexico, Panama, Colombia, Ecua- dor, and Peru JUVENILE BIOLOGY Ecological observations in the field were con- ducted with the aid of binoculars and suitable recipients for living adults and larvae (for details see Brown & Benson, 1974, and Brown, 1972b) Identification is secure for adult ithomiines, at least 9096 certain for juveniles, and secure for Solanaceae host plants at the genus or section level but still uncertain in a few cases at the species level. Early stages of ithomiines were brought into the laboratory when necessary and reared to adults in glass or plastic dishes on separated leaves of natural or experimental host plants, kept subhumid. At least one fertile egg could usually be expressed from any field-captured re- productive female ithomiine by careful pressure on the abdomen with thumb and forefinger, working backwards slowly from the fourth seg- ment. Such eggs were kept with expected or po- tential food plants until hatching (3-9 days). A large number of potential food plants could be found in Monjolinho or other areas in Campinas; many others were cultivated as available in my garden. 1987] LIVE INSECTS OR FRESH LEAVES vigorous «d RIS ze with 10 BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY 365 at room temperature volume of MeOH:H20 (8:2 or 9:1) filter and wash with another smaller portion extract solid residue virgini to dpt. volume, under vacuum, homogenize OPTIONAL 40? ving all methanol; acidify with 6% volume of conc. H2SD4 eh 2 x 10x extract 3x with an equal volume of CHC13 extracts lower layers y unite, make basic with conc. NH40H (to pH 10) unite, dry with evaporate chitin or extract with 2 equal volumes of CHC13:Me0H 3:1 anhydrous Na2SD4 under fiber and one larger volume of CHC13 vacuum evaporate under vacuum lower layers E - A aqueous unite, dry with v i I (nonpolar material: mostly fats, layer anhydrous Na2S04 moderate polar acid waxes, carotenes, hydrocarbons) (basic) iris x I or — esl des uec evaporate under vacuum pigme rin steroi many u mE podere extract residue B with MeOH (2 cycles) ( 7 . analysis by TLC on solid extracts moderately polar basic silica gel erige alkaleids and thei 5, inorganic selt vaporate wir glycosides, Aminas) Further fractionation on es) 3504 under vacuum analysis by TLC, columns of silica gel and C silica gelH, 7 fractionation on CHC14-MeOH + (highly polar material: alkaloid N-o pci Te m e nal reducti opt loop at this atage and zwitterions, some salts) Nis Fu deactivated basic alumina or partition columns PAEA by TLC aper chromatography s> Further fres ionation on Sephadex or t -exchange column E4. Mild and rapid standardized fractionation scheme for fresh leaves, flowers or insects, adapted to alkaloid- -containing materials, giving fractions for chemical and ecological analysis and assay. CHEMICAL FRACTIONATION A standard extraction and fractionation was developed for all plant or animal material (Fig. 4). Leaves or flowers of plants observed to be used by Ithomiinae larvae or adults were sepa- rated from petioles or peduncles and maintained fresh until extraction; juvenile and adult Ith- omiinae were kept alive in dishes or envelopes until the moment of extraction. Thus, all ex- tractions were performed on fresh and recently functioning living tissues. The method was adapted to rapidly remove all organic substances of moderate to high polarity and at the same time deactivate enzymes in the living material; stable nonpolar compounds were removed afterward The initial use of heat, aeration, acids, bases, or acetone (which can condense with primary amines) was avoided. Exhaustive extraction was sacrificed in favor of rapidity; the more labile polar compounds (especially glycosides) were immediately and efficiently dissolved by vigor- ous maceration in the first extracting solvent added to fresh material. The leaves, flowers, or insects were divided if necessary into pieces of no more than 2 cm maximum linear dimension by scissors, already under or falling directly into methanol-water 8 : 2 (usual for leaves to extract less chlorophyll) or 9: 1 (for adult insects), using about ten times the volume in milliliters as the fresh weight of material to be extracted. The ” extraction was immediately completed with scis- sors (if less than 40 or with a kitchen homogenizer (Braun **Mini-Pimer" with a two- blade metal propellor) introduced into the sus- pension from above. Within a minute, leaves were reduced to tiny fragments and insects to pieces of chitin, partly coated with a fatty layer. The extract was homogenized for 1-3 minutes and filtered under suction; the insoluble material was washed and stirred with a further amount of aqueous methanol, up to half the original vol- ume. Nonpolar compounds (fats, waxes, caro- tenes, some terpenes, and free flavonoids), 1 soluble in aqueous methanol, were Sewg end extracted from the residue with two portions (each 5 x original fresh weight) of ethyl acetate, giving a “fat fraction" (F) and leaving only polymeric organic fiber or plaque (cellulose, lignin, chitin). The filtered and united aqueous-methanolic extracts were evaporated under vacuum at less than 40°C until all the methanol was removed. The resulting aqueous suspension (at times after filtration, giving a chlorophyll or nonpolar frac- tion F) was acidified with conc. sulfuric acid (roughly 696 of the total volume, making it about = 1 M in acid) and extracted three times Wil: an sanal volume of chloroform. If occa- t this or later stages (especially in some plant extracts) did not sep- arate after 6 hours, they were filtered or broken 366 with MeOH. The organic layers were united, dried with anhydrous granular sodium sulfate, filtered, and evaporated under vacuum to dryness to give a relatively more polar but nonalkaloidal frac- tion (A), often including some chlorophyll from leaves (usually little soluble in aqueous metha- nol) plus terpenoids and steroids (incl cardiac glycosides, withanolides, and saponins), flavonoids, coumarins, and organic acids (Fig. 2). Half or all of the aqueous phase was then made strongly basic (pH above 10) with conc. ammonium hydroxide and extracted with two equal volumes of chloroform-methanol 3: 1 and one to three volumes (1.5 x) of chloroform; this gave effective partition between water-methanol 8:1 and chloroform-methanol 6:1, obviating salting out of polar organic constituents from the upper layer. The organic layers were combined, dried, and evaporated as above to give a mod- erately polar alkaloid fraction (B) containing gly- coalkaloids, tropanes, pyrrolizidines, and sim- pler bases (Figs. 1, 3). The remaining aqueous phase retained alkaloid N-oxides, very polar gly- cosides, more polar acids, and salts and sugars; it was directly evaporated and extracted with methanol (two cycles with filtration) to give a polar extract (C) and leave insoluble organic salts, mostly ammonium sulfate. If the alkaloids cor- responding to the N-oxides were desired, the oth- er half of the acidic solution (after CHCl, ex- traction, 2 N in H,SO,) or fraction c after dissolution in 2 N H,SO, was reduced by stirring for 2 hours at room temperature with an excess of zinc dust (usually in grams equal to one tenth of solution volume), filtered (paper cone, gravity funnel) alcalinized with an excess of conc. NH,OH (until all zinc salts dissolved as the zinc- ammonia complex), and extracted as above with CHCl;-MeOH (2x) and CHCI,. If this gave a much greater weight of alkaloid than in unred- uced extracts, significant amounts of N-oxide were indicated. The complete extraction and fractionation scheme is illustrated in Figure 4. Crude or purified fractions were analyzed by NMR spectra in CDCI, + 1% TMS, on the Var- ian T-60 of the Instituto de Quimica, UNI- CAMP, or eventually by IR spectra (in CHCl, or KBr) on a Perkin-Elmer Infracord. Mass spec- tra were performed by Mrs. Concetta Kascheres on the Varian MAT 311A instrument of the In- stituto de Quimica, at 70 eV, probe temperature 60-105?C. Chromatographic analysis on TLC plates (coated microscope slides) used silica gel H (no binder) and varying amounts of chloro- g most ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 form-methanol or benzene-ethyl acetate, plus 196 NH,OH for alkaloidal fractions; other systems used ethanol, acetone, and amines. Separations on adsorption columns (silica gel or alumina ac- tivity III were usual) followed results of the TLC analysis and used careful gradient elution. AI- kaloids were purified by partition chromatog- raphy with the system ethanol-chloroform-water (usually 1 :9 : 1), stationary (upper) phase 0.75 x (V/W) on Celite 545 and including phenol red as indicator (Brown & Kupchan, 1962) or chlo- roform and buffer solutions. DETERMINATION OF PAS Dehydropyrrolizidine alkaloids (PAs) were quantitatively assayed in fresh plant or insect material (they are destroyed after death or dry- ing), cut up under absolute methanol (at least 10x), and, after at least one day, colorimetric determination on aliquots (usually 1/20, prefer- ably containing no more than 100 ug PA; 40 ug gave initial absorbance off scale > 2.0) and the Beer’s law curve became bove 150 ug) with a modification of the Mattocks (1967, 1968)- Bingley (1968) method. The assay is very sen- sitive as modified (< 0.1 ug is easily seen), very accurate for N-oxides (+ 1%), reasonably accu- rate for total PA (+ 10%), and extremely specific and applicable on total plant or insect extracts with minimal interference by other components; the final product of Ehrlich condensation shows Amax 261.5 nm (in isoamyl acetate-ethanol-ace- tone about |: 1:2 with 2% BF,) with e ~ 57,000 (Fig. 3L). For total PA determination, the aliquot is treated with 0.5 ml of freshly prepared 0.15% methanolic H,O, (from 200: 1 dilution of 30% aqueous H,O, stabilized with 5 mg/ml of Na,P,0,) and heated in a boiling water bath for 30 minutes, followed by drying with hot air for 5 minutes. This procedure destroys about 40 + 5% of the total alkaloid N-oxide formed, but milder methods of oxidation give incomplete re- action or difficulties in the following steps. The resulting completely dry N-oxide (or an aliquot of the original extract air-dried at 80°C) is taken up in 1.0 ml isoamyl acetate, treated with 0.125 ml acetic anhydride, and heated 2 minutes in the boiling water bath. After being cooled to room temperature, the solution is treated with 1.0 ml ofa solution of absolute ethanol : 2096 BF,-meth- anol: p-dimethylaminobenzaldehyde (50:5:1) and heated at 56-59°C for five minutes. Absor- bance is read against a blank which passed through all the reactions, at 561.5 nm, after di- 1987] lution to 3.8 ml with anhydrous acetone; further dilution with acetone gives a linear decrease in absorbance. At room temperature absorbance reaches a maximum several hours after the last reaction, about 5% above the reading immedi- ately after it, and then diminishes; it may be maintained for many days at 4°C. Each micro- gram of PA in the original aliquot contributes to the absorbance 0.05 (N-oxide) or 0.03 (total base) (standard curves with echinatine and lycopsa- mine samples, Fig. 3G or E, provided by J. A. Edgar). Determinations were routinely done in racks of 40 13 x 100 mm glass test tubes, always doing total base and N-oxide determinations in the same batch to equalize the initial aliquot sampling and give an internal control on abnor- mal results. Separate pipets and bulbs should be used for each reagent in the series; water must be rigorously eliminated from the test tubes be- fore the Polonovski reaction (Ac;O-isoamyl ace- tate). PAs with a carbonyl group conjugated with the unsaturated ring (7-ketones, or 9-aldehydes or carboxylates, such as Fig. 3M-O) or with no dou- ble bond in the ring did not give any blue color with this method. Dihydropyrrolizines (with the nucleus of Fig. 3M, N) without carbonyl conju- gation could be directly determined on aliquots to an accuracy of about + 10% using the final reaction in the series. They were usually de- stroyed in the first reaction step (oxidation) if present in the extract. NEPHILA BIOASSAY Fr apne were bioassayed for predator-de- fense compounds, following observations of a and Lewinsohn (1982, 1984), with natural populations of the giant tropical orb spider Nephila clavipes (L.). This predator, com- mon from December through July in Campinas (the same period as that of maximum ithomiine density in the same places) and present in small numbers year round, cuts out of its web, alive and unharmed, any field-captured ithomiine that is introduced naturally or experimentally, after contact with any part of body or wings. If the butterfly struggles excessively (ithomiines gen- erally remain quiet, awaiting liberation), it may be attacked or wrapped in silk, but the spider does not suck it (Vasconcellos-Neto & Lewin- sohn, 1982, 1984). Nephila spiders are among the most abundant, effective, and aggressive po- tential predators of flying insects in Campinas and throughout the more seasonal Neotropics, BROWN—SOLANACEAE/ITHOMIINAE CHEMISTRY 367 and can be safely regarded as very important in the evolution of predator defense mechanisms in the Ithomiinae. Vertebrate predators have not been used yet in the bioassay, but they were shown to reject an itl by Brower & Brower (1964), to reject almost all ithomiines by Haber (1978), to reject two ithomiines by Coimbra-Filho (1981), and to accept only fatty abdominal contents of ithomiines in Brazil by Brown & Vasconcellos- Neto (1 Living panado of the palatable nympha- line butterfly Biblis hyperia (Cr.), readily at- tacked and eaten by the spider, were painted with aqueous solutions, suspensions, or emulsions of fractions or compounds (corresponding to 0.5- 1.5 butterfly, 2 g fresh weight of leaves, or 0.2 g fresh weight of flowers), covering the entire body, legs, antennae, and wing bases to at least half the radius, and let dry. Still alive, they were thrown into part of the web of an adult, non-satiated, and healthy spider (as judged by web structure), from a distance of 0.5-1.5 m on the side opposite the spider, between 1200 hours and 1500 hours on a warm sunny day. The spider normally ad- vanced immediately to attack the butterfly (de- lays of up to 5 minutes can occur if the spider is “dozing” or distracted). If the Biblis was punc- tured, sucked, and then cut out or wrapped in silk by the spider, or (to still its struggles) was bitten, wrapped in silk, cut out, and taken to another part of the web and sucked, the test was regarded as negative. All such tests were repeated at least twice with different spiders and different individuals of Biblis. A positive test consisted of the spider’s drawing back from the animal, at times inspecting various parts with its sensory palps but not biting or sucking, and eventually cutting out the living butterfly by breaking all necessary strands (usually cutting them with the third pair of legs), manipulating the entire insect and letting it drop unharmed. This test, usually repeated at least three times with at least two Biblis, was regarded as strong evidence for effec- tive predator-deterrent compounds in the ex- e - ° t. Biblis is common year round in most parts of the Neotropics, not especially fast- or high-flying, and readily attracted to fermenting baits. It is brightly colored black and red, regarded as an “incipient mimic" by the Browers (1964; Brow- er, 1969; Brower et al., 1971). Its larva feeds upon Tragia, an urticant euphorbiaceous vine, and the adult possesses dorsal scent glands in the abdomen that it displays in evident defensive 368 behavior when handled. Nevertheless, it is readi- ly eaten by Nephila and thus is an ideal substrate for the bioassay of chemical fractions. Alterna- tive organisms used with similar success in the bioassay include some pierids (Eurema) and Heliconius erato, also eaten by the spider (Vas- concellos-Neto & Lewinsohn, 1982, 1984). CHEMICAL RM nena AND COEVOLUTION A SIRE INTERFACE HOST PLANT UTILIZATION PATTERNS In the principal field sites around Campinas, SP, at least 42 species of Solanaceae are used (or potentially should be used, by analogy with other regions) by larval Ithomiinae (Figs. 5-8). A fur- ther four species have been inspected frequently but show no signs of usage: Capsicum praeter- missum Heiser & P. G. Smith, Lycianthes ran- tonnetii Carr. ex Lescuyer, Solanum wendlandii Hook. f. and Solanum americanum Mill. Many additional species occur in nearby field sites (at least 15 more species of Solanum) or in gardens. Six of the 25 species (in 17 genera and 11 of the 13 neotropical tribes) of Ithomiinae known from the Campinas region (Figs. 5, 9) can be regarded as only occasional tors (the two Mel- inaea, Episcada philoclea, Pseudoscada quadri- fasciata, and Hypoleria adasa and goiana), these possibly do not find enough adequate host plants available to establish permanent populations. At least 15 additional species in as many genera occur in healthy populations within 200 km of of the 19 species regularly present feed only on ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Apocynaceae-Parsonsieae (Fig. 5). Thirty-six species of solanaceous food plants have been re- corded for 14 of the remaining 17 species and may be confidently predicted for the rest (Fig. 5); no species is monophagous but only Mechanitis (and possibly Pteronymia) could be called poly- phagous (the rest are best regarded as quite nar- rowly oligophagous, especially in the chemical sense, as will be shown below) This pattern is a microcosm of the general pic- ture of food plant usage by the Ithomiinae (see Drummond & Brown, 1987), comprising about 400 species X species interactions. Monophagy or narrow oligophagy is the rule at the present level of information, not only for local popula- tions but also for whole genera. A summary of all these data (Table 2) with chemical informa- tion added from many sources in the literature (with preliminary verification in this work, see Table 3) shows a reasonable specificity in the interactions between Ithomiinae and Solanaceae, at least at the level of genera of ovipositing fe- males and host plants. Most of the ithomiine genera are known at present from only one or two Solanaceae genera or subgenera of Solanum; even generalists like the common and ubiquitous Mechanitis show a strong epos for only two subgenera of Solanum in most s The records are still I. ices Ith- omiine ovipositions are not commonly ob- served, and 23 of the 52 genera are still repre- sented by three or fewer confirmed food plant records. The possibility cannot be discounted that the specificity patterns shown in Table 2 are pri- marily due to lack of adequate information. Con- fident patterns of food plant specificity are seen only in the well-documented interactions of the — Known and expected food plant relationships of Ithomiinae in the Campinas region, São Paulo. FIGURE 5. Heavier e: have all been verified in interbreeding populations occupyi diameter circle Campinas and Barão ng various habitats within a 5-km Geraldo; note prevalence of oligophagy and some polyphagy e (Mechanitis, Prittwitzia, Pteronymia). Neither Ithomiine tribes nor Solanaceae genera are in evolutionary order (see Table 2 for this), but have been arranged so as to simplify the figure FiGURE 6. Solanaceae cal southeastern Brazil inc iir in Campinas, São Paulo unless otherwise indi- itianum S. s cated): s us m.—A. S. m robustum. —F. S. LUE authors' names, see Figure 5. wartzia E 7. Solanaceae of southeastern Brazil “ee nee otanic o des Janeiro). — ansia suaveolens. — N. Br GUR mansia candida (cultivated; Jardim B Miers (Jardim Botánico, Rio de Janeiro). — m. —C. S. capsicoides. — D. S. aculeatissimum. —E. —G. S. an ei “H S. megalochiton. —]. S. pseudoquina St. Hil. For —J. Solandra grandiflora (cultivated). —K. Brug L. Markea (Dyssochroma) ehe e (Sims) unfelsia australis. —O. Vassobia breviflora. — P. Acnistus arborescens. — Q. micis ipee alum. —R. Cestrum intermedium (Joinvile, Santa ‘Ca a- tarina). 1987] BROWN—SOLANACEAE/ITHOMIINAE CHEMISTRY 369 Known ( m; ) and presumed (——————-—- ) food-plant records for Ithomiinae in the region of Campinas, Sao Paulo, southeast Brazil (23* s, 48° w, 600-1,000 m) ITHOMIINAE FOODPLANT TITHOREINI: TITHOREA harmonia pesudethra Butler PELTASTES peltatus (Vell.) Woodson NADENIA vi Mi (NEW TRIBE): TEMNAD violacea Miers AERIA olena olena Weymer PRESTONIA coalita (Vell.) Woodson APOCYNACEAE PRESTONIA dusenii (Malme) Woodson PRESTONIA acutifolia Schum. METHONINI: METHONA themisto (Hii BRUNFELSIA uniflora (Pohl) D. Don BRUNFELSIA pauciflora (Cham. & Schldl.) Benth. (NEW TRIBE) PLACIDULA euryanassa (Feld. & Feld. )'™ — ku suwa BRUNFELSIA australis Benth. MELINAEINI: SSDS LTT A MELINAEA ethra (Godart) ~ "c ess DATURA stramonium L. NI n. ~~ P MELINAEA ludovica paraiya Reakirt -— “T NEL BRUGMANSIA suaveolens (Willd.) Sweet SSI na T BRUGMANSIA candida Pers. SS ITHOMIINI TM. 7 SOLANDRA grandiflora Sweet ITHOMIA d drymo Hübn TUS iind y» ~ TY ie MARKEA (DYSSOCHROMA) longipes (Sendtner) ~~ Miers ITHOMIA agnosia — Hewit ikani D' Must. nec PHYSALIS neesiana Sendtner ee ACNISTUS arborescens (L.) Schldl. PITYCHES eupompe (Geyer) ———— VASSOBIA breviflora (Sendtner) Hunziker RIPOIHEBIS- eur Lea n Me reisen ee CAPSICUM flexuosum Sendtner HYPOTHYRIS ninonia daeta (Boisduval) NICANDRA physaloides (L.) Gaertner CYPHOMANDRA sciadostylis Sendtner S A CYPHOMANDRA divaricata Sendtner 0-98 ff CYPHOMANDRA fragrans (Hook.) Sendtner MECHANITIN sp T1 cetoides (Ros alb.) pal E lida Godm. & de X hippodamia (Fabr.) CYPHOMANDRA crassicaulis (Ortm.) Kuntze « MECHANITIS polymnia casabranca Hnsch: 4 LYCOPERSICON esculentum Miller E NS SX SOLANUM mauritianum Scop. ie NN SOLANUM granuloso-leprosum Dunal u MECHANITIS lysimnia lysimnia (Fabr.) SOLANUM concinnum Schott ex Sendtner Ds aS AS SOLANUM megalochiton C. Martius ° `< n SOLANUM murinum Sendtner g gioi Jera (thoes Felde SOLANUM swartzianum Roemer & Schultes na ae SOLANUM melongena L. A NN KN. NW ` UN SOLANUM paniculatum L. (NEW TRIBE) E P " PRITTWITZIA hymenaea hymenaea VM S SOLANUM asperolanatum Ruiz & Pavón (Prittw.) SOLANUM robustum Wendl. EPISCADA philoclea (Hewit ) m SOLANUM variabile C. Martius N SOLANUM insidiosum C, Martius EPISCADA carcinia Schaus w \ MN ` SOLANUM brusquense Lyman B. Smith & Downs PTERONYMIA carlia Schaus SOLANUM arcuatum Sendtner SOLANUM viarum Dunal GODYRIDINI: SOLANUM capsicoides All. PSEUDOSCADA quadrifasciata Talbot wmm, SOLANUM atropurpureum Schrank SOLANUM jatrophifolium Dunal PSEUDOSCADA erruca (Hewitson) HYPOLERIA adasa (Hewitson) wa. SOLANUM aculeatissimum Jacq. DIAJ HYPOLERIA plisthenes D HYPOLERIA goiana D'Almeida — Í SOLANUM pseudocapsicum L. SOLANUM caavurana Vell. CESTRUM schlechtendalii G. Don MCCLUNGIA salonina salonina CESTRUM laevigatum Schldl. ewteson) CESTRUM sendtnerianum C. Martius ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 > x E z = = T Q = < z = ° en E u) < = Q < Z < o O N | Z. z ° x m ANNALS OF THE MISSOURI BOTANICAL GARDEN “QS. U iii É P mh AQ; : lite. 4 (4 «6:14 Y Y ` 1 @ M h " C» OCA aN ‘ | ) "3v > = j 1987] genera Tithorea, Elzunia, and Aeria (with Apoc- ynaceae: Parsonsieae), Methona, Melinaea, Thyridia, Mechanitis, Hypothyris, Ithomia, Dir- cenna, Pteronymia, and Greta. At a higher level, almost all members of the last two tribes show collectively a strong specificity to Cestrum and surprisingly also to the distant section Geminata of Solanum. It is probable that more work will provide more evidence of the restriction of Na- peogenes to Lycianthes and confirm the nascent patterns in Placidula, Scada, Epityches, Oleria, and Callithomia, but this is mere speculation at this time. At the species level, Mechanitis females tend to divide up the local Solanum, resulting in min- imal overlapping of oviposition (see Drum- mond, 1976; Haber, 1978; Vasconcellos-Neto, 1980, 1986), but the divisions do not coincide with simple taxonomic lines in the plants. They also vary from area to area and are not correlated with larval choice or survivorship in experi- ments, indicating an unstable ecological deter- mination of the partition. Though the butterflies may be able to recog- nize their food plants, biologists have greater dif- ficulty in Solanaceae identification, even to the genus level, which may result in some spurious patterns in Table 2. In 1969, a top Solanaceae taxonomist identified for me a specimen from near Rio de Janeiro as a Capsicum on the basis of its cleft anthers, but both S. Knapp (pers. comm.) and I regard this plant as Solanum (sect. Geminata) caavurana. The “diagnostic” trait in this case may have been an artifact of the drying BROWN SOLANACEAE/ITHOMIINAE CHEMISTRY 373 method used for the specimen. If such evolu- tionarily distant and chemically distinct taxa could be potentially confusing to an experienced expert, how will they appear to the average field biologist who, having just observed a female ith- omiine oviposit on a (presumably solanaceous) bush with glabrous entire leaves and no flowers, faces the maze of complex and confusing tax- onomy in the family? This plant could be placed preliminarily in over 30 genera, five with over a hundred species and one with over a thousand. Thus the “solid” data base for host plant usage in Table 2, from which a number of completely unlikely records have already been purged, may yet suffer fundamental modifications with fur- ther work. COEVOLUTION AND PHYLOGENETIC DIVERSIFICATION An attempt to relate the phylogeny of the Ith- omiinae with that of their larval food plants, such as was done with reasonable confidence and re- sults for the nymphaline tribe Heliconiini (Ben- son et al., 1976), has met with very little success (Drummond, 1986; Brown & Drummond, prep.; Drummond & Brown, 1987). Only ie broadest pattern can be seen in the progression from Apocynaceae to Solanaceae (with Gesne- riaceae in the middle to provoke the imagina- tion). The most primitive ithomiine genera to use Solanaceae (Athesis, Methona, Olyras, and Melinaea) feed on the genera regarded as highly advanced in this family. Two large tribes (Me- chanitini and Napeogenini) run their food plant — FIGURE 8. K-M, P, R. Second instar larvae, about 4-7 mm long.— S-DD. Fourth or fifth instar larvae, an 20-35 mm long. —EE. Prepupa and p pseudethra a ru sp.), Pirac Napeogenes sulphurina Bates Hans Sp., a vine), Ipojuca, Pernamb — apa.—F, DD P eee eii canaria Brown & D'Alm anjuba, Goiás.— B, U. Aeria olena (Prestonia pé teg Campinas, CC. G m sp., note simple ES poe Para.—E, | Juveniles of Ithomiinae with food plant and locality. A-J, N-O. Eggs, about 0.6-1.0 mm high. — . Adult ovipositing MA AA J. Vasconcellos-Neto). — . A. Tithorea harmonia SP. —C, AA. arsauritis xanthostola n Sendt.), Santa Teresa, Espirito onymia hemixanthe (Feld. & Feld.) (Solanum SUPE Brev vantheru?) Ubata, Bahia.—I, R. Me- i m 102 (Solanu (Solanum asperum), Linhares, Espírito deii originally reported as “Forsteronia sp."), Jar (4 vir mop, anf grandiflora D. Don), Barinitas, Vene Bryk (Prestonia? sp.), Belém, Para.— hona ME reip sus australis), Pinos on E — E. H cidit eucled laphria S ssp. Mone ulei (D m.) Cuatr., e V are from watercolors by Emily Fountaine, uela. d in the British Museum (Natural Histon. Solanologiss are requested to photograph and rear through jn le adult or to preserve in alcohol any eggs (with ridges as arvae of these types found on identifiable plants and send them to B. Drummond or K. Brown for uec Ed and registry in the data bank for Table 2. 374 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 TABLE 2. Approximate number of known interactions (species x species) between Ithomiinae and their larval food plants, with data on plant chemistry (Figs. 1, 2) and on attraction to pyrrolizidine alkaloid sources. Plant Families: Genera*” (no. species) . Solanaceae Attraction to: Solanum (1000) = 8 Yo . § >= = 2 s g Se ERR Y Š a = @ Q2 X d^ =< ii ša Gg Sa NEFS P t Setasacss SzS Š Š sp Se sSssS SISESE ES S S š EQgES Ise, SSS Š g Bese SESS DOE SOQ DOE E Š E š DP 22293923 $53 § Š SEs gre PERSIS SS S $ S goggas ggs i $ Aš us ef83 GSS asa $85 O 5 FSFE aldàd 333 Š š Alkaloids (Fig. 1) Ste roids, a PAs (3E-I) ? === ABFLH ---- BL ABCL BL BC?L C? ABC BC BCP C BC C? LC BCE onstit uents Other C C Fig. 2) Ithomiinae: Cardiac Tribes, Genera!“ glycosides, (no. species) XYZ WY ~ RTUWXYZ -R? Z RWXY YQ? Q QSY QY QYZQR XY XY RWXYZ XY A. Tellervini* Tellervo (Ay l I 5 B ithoreini Elzunia (2) l ? 4 dais dd l 1 16 1? C. (N shi pe 2 3 8 D. (New Tribe) Athesis (1) l l 1 E. Methonini Methona (7)" l l 12 F. (New Tribe Placidula (1) 2 3 1? 6 G linaeini Olyras (2) 2 2 1? 1 ^ utresis (2 2 2 I? 1 Melinaea (10) 2 2 1 9 H nitini Thyridia (1)" 2 3 6 (2) 1 3 l la l 3 l 3 Forbestra ( l 3 l Mechanitis (5)"' l 3 3 1556 1 7 2 1? |! Ll. Oleriini Hyposcada (8) 2 3 3 1 New genus (3) l 1 l Oleria (30)" 2 3 148 3 J. peogenin AKA 2 3 2 l 2 "es a (1) 2 3 1? 1 Napeogenes (18)" 2 3 1? 8 Garsauritis (1) 2 3 4 Hyalyris (16) 2 3 5 Hypothyris (16) 2 3 1 2413 1 K. Ithomiini Miraleria (1) 2 3 l 2 1 Ithomia (21)" 2 3 2 2 11 3 5 L. Dircennini Callithomia (3)^ 3 3 13 I Velamysta ( 3 3 1 New Ge 2 ? 1 Dircenna (7)" l 3 14 17 Hyalenna (5) 3 3 1 M. (New Tribe) Ceratinia (5) 3 3 3 3 Ceratiscada (3) 2 3 2 rittwitzia (1) 2 3 3 1? 5 Episcada (15) 2 3 4 Pteronymia (41)" 3 3 25 3 3 2 1987] BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY 375 TABLE 2. Continued. Plant Families: Genera*” (no. species) Solanaceae Attraction to: Solanum (1000) a 5 doa S — Ithomiinae: š E g is E š Š Rn T 9 a oe e "S cu e Tribes, Genera** s oo g a Ss E 3 > S E S Ss A ago fsa & s (no. species) Ups DugE Bog ESS ES 8 § — 3 "EF s» Vo ss Š s ee be SESS CSR RSs ° vk Ë E 38 BS 8 SS39 Us S sss E 3 esse DID Es -1 20 SSE 2 Š BEE 232353 BID š $ = = = e I Fi = S3 as BESS GSS ase § S55 Š § FSFE AgI S83 Š Š N. Godyridini Godyris (10)^ 3 3 5 Pseudoscada (6)" 2 3 5 Hypomenitis (6) p. 3 2 reta (10 2 3 l l 16 Hypoleria (1 1)" 2 3 1 6 Mcclungia (1) 2 3 3 Heterosais (3)" 3 3 3 a Families are in approximate order of advancement and genera of Solanaceae in order according to Hunziker (1379), from primitive to advanced. ^ In addition to Nicotiana (see note i, below), important Solanaceae genera available to Ithomiinae in tropical America for which no food plant records are yet known include J dicii Athenaea (but nid uc used by Epityches and pos mia), e Saracha, Iochroma, Salpichroa, Jaborosa, Lycium, Grabowskia, Trianaea, essea, Metternichia, Petu Fabiana, Nierembergia, oS Protoschwenkia, Schwenkia, Melananthus, Ppbounedia, oet y Sin za Streptosolen, and Het —weak attraction usually including both sexes, 2—males a attracted, 3—males strongly and females regulary attracte ribes and genera follow the order of Mielke and Brown (1979) as modified by Brown (in prep.), from * os to advanced (including genera within each tribe). * No food plants have yet been recorded for the following genera of Ithomiinae (number of species in paren- theses). * = Andean cari usually of y qd us cloud forests, though Roswellia extends out into the foot- hills.— Tribe D. *Roswellia (1), *Patricia (2).— Tribe G. Athyrtis (1).— Tribe H. Paititia (1).— Tribe I. *a new genus near Hyposcadi i — Tribe J. 7 (1).— Tribe K. *Pagyris (1).— Tribe N. *Dygoris (1) and *Ve- ladyris (1). Total, 9 genera with 10 s f The recent divisions of these ina prod into a number of smaller Es (Hunziker, 1979), most of them used by the same Ithomiinae, are here included under the collective older names. e Restricted to the Old World tropics, regarded by some as closer to the Denials than the Ithomiinae (see Ackery & Vane-Wright, 1984). ^ Food plant patterns are constant from the northern to Edd southern extremes of the range of the genus, usually from Central America to southern sl Aes. of 18 gen ! Ovipositions of Mechanitis polym asabranca were B S ENE in Campinas, São Paulo, on "iaa ad Capsicum annuum L. and feral Nicotiana (rosette only), but the ipe did not grow on these pla j The published structure for *hopeanine" (1E) is biogenetica t with the data presented in the einge phe: (Iyer, 1978); no alkaloids of ` Brunfelsia are lau under investigation. phylogen ds ( d dbutterflies does not lead to a fruitful parallel. The patterns use Por due) veins plants). And although for the Ithomiinae are complex, and the arrange- the most morphologically advanced ithomiine ment of the plant groups in Figure 5 so as to genera concentrate on the relatively advanced produce a maximum parsimony scheme (lowest Cestrum, they use equally well the supposedly number of crossing lines in the middle) has pro- primitive section Geminata of Solanum (Table duced some strange and thought-provoking 2). That the neotropical Ithomiinae representa proximities between genera normally widely sep- widely diversified butterfly group (51 genera in arated; note especially the positions of Brunfelsia 13 tribes) could help explain why a comparison and Cestrum. of their host plant utilization patterns with those While the Solanaceae indeed may have rep- of the 65 rather homogeneous species in no more resented a new field for adaptive radiation of the than ten genera of the single tribe Heliconiini Ithomiinae in the New World, it is probable that 376 the plant family was already very diversified chemically and taxonomically before the butter- fly subfamily began to move onto it. A large number of available genera (at least 24) are still not known to be attacked by Ithomiinae (see Table 2). Furthermore, the genus Brunfelsia, re- garded as a relatively advanced member of the Solanaceae, apparently entered the Caribbean area and diversified greatly (as section Brunfel- sia) before Methona, a relatively primitive ith- omiine genus in a monotypic tribe, could evolve as a specialist herbivore on it (Plowman, 1979, p. 489). It is possible, however, that the extensive speciation verified in Solanum and Cestrum eventually may be related to their heavy use by a variety of Ithomiinae (Table 2). This can be investigated only by careful chemical analysis and bioassay with larvae, Mines in parallel twoi tly developin y lin of species or populations Pie ‘show. specific in- teractions. urther considerations of the incongruities be- tween phylogenies of genera of Solanaceae and Ithomiinae are presented in Brown & Drum- mond (in prep.). 1 1 O n BIOCHEMICAL COEVOLUTION The taxonomic diversification of the Solana- ceae has been accompanied by an impressive diversification in secondary chemicals, leading some to suggest polyphyly for the family as pres- ently constituted. Indeed, no other plant family can challenge the Solanaceae in having alkaloids representing all four major biosynthetic path- ways (lysine-ornithine, phenylalanine-tyrosine, tryptophan, and steroid), an equal number of less important pathways (nicotinic acid, anthranilic acid, glycine, acetate), and a further set of aber- rant or combined sources unique to the family (leading to capsaicins, solanines, withasomnine, and hopeanine) (Fig. 1), not to mention more common amines such as choline, noradrenaline, and hypoxanthine; some species also produce very toxic peptides. As is often the case, these alkaloid-rich plants are singularly poor in lower (volatile) terpenes, but some Datura, Cypho- mandra, and especially Cestrum and Solanum section Geminata share a similar and abundant pungent oil that may include terpenes. Diter- penes and triterpenes abound in some Solana- ceae, and nonalkaloidal steroids are represented as saponins (such as diosgenin glycosides) and withanolides/physalins, unusual steroidal lac- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 tones with marked biological activity (Fig. 2). Flavonoids and coumarins also are documented amply in diverse members of the family (Fig. 2; Harborne & Swain, 1979), and further phenolics and their glycosides (including tannins) are al- most always found when sought in polar extracts. The C,; alcohol solanesol is an unusual unsat- urated linear nonaprenol (Fig. 2). Lacking are reports of cyanogenic glycosides, cardenolides, glucosinolates, nonprotein amino acids, and ir- idoid glycosides, but their absence cannot be de- finitively affirmed because few phytochemists have apecttically sought them in the Solanaceae. hus the p this family are well defended chemically and attacked by rather few insect her- bivores other than Ithomiinae (Table 1). The only ones that seem to promote similar damage in natural systems are generalist grasshoppers and meloid beetles, perhaps molluscs in more cal- cium-rich areas, and specialist chrysomelid bee- tles. The plants also at times are defended ad- mirably against vertebrate chewers by prickles (Symon, 1986) but these have very little effect on smaller invertebrates, who walk and chew around them or build silken pads over them. Other defenses (tough leaves, glandular tri- chomes) are more effective against invertebrates (Vasconcellos-Neto, 1980, 1986), as may be fur- ther quantitative defenses (t amino-acids, and resins) and a host of ecological strategies such as phenology (see Vasconcellos-Neto, 1986) hab- itat, growth form, seed dispersal by chiropte- rochory, and encouragement of spiders and pred- atory pentatomids on the leaves, among others observed in Brazil. When the generic preferences of the butterflies are analyzed in terms of plant chemicals (Table 2), the associations seem to show patterns, though the data are still chemically and biologically very incomplete. There appears to be a certain ten- dency toward chemical specificity, at times with- out reference to taxonomic relationship (as in the similarly potent-smelling Solanum sect. Gemi- nata and Cestrum). It seems probable that female Ithomiinae search for and find specific chemical cues for oviposition on certain genera of Solan- aceae and may be led to place eggs on unrelated plants with similar chemicals. What else would permit a Mechanitis polymnia female in Cam- pinas in May 1981 to recognize a small Nicotiana plant in Monjolinho as a potential Solanaceae host, or to oviposit on Capsicum annuum in my garden, especially when the larvae survived on neither? (The first shares nicotine and the second 1987] steroidal glycoalkaloids with a usual host, So/a- num mauritianum.) Based on the patterns in Ta- ble 2, some still unknown chemical may be pre- dicted to set Solanum subgenera Bassovia, Potatoe, and Lycianthes apart from all other So- lanaceae (see Napeogenes, Oleria, and Calli- thomia) and possibly associate Lycianthes with the withanolide-elaborating genera (see Ithomia) and even with Solanum sect. Geminata (Ptero- nymia uses both). Further support for some degree of chemical mediation at the oviposition sector of the inter- face comes from the broad geographical consis- tency of the more specific relationships, at least from Mexico through Costa Rica and Ecuador to south Brazil for the 18 genera for which food plants are known over this range (Table 2; Drum- mond & Brown, 1987). This surely must have been established over long evolutionary time in developing phyletic lines of the butterflies. The chemical proximity of disparate taxonomic groups of plants, treated as interchangeable by ovipositing females, also supports a chemical mediation at the interface. In order to separate this possible evolutionary component in food plant usage from the ecological noise in the sys- tem, careful comparative tests with free-flying females must be performed under controlled conditions, coupled with larval feeding prefer- ences in multiple-choice tests, and thorough chemical analysis of all potentially active com- pounds in the food plants chosen or rejected. The patterns broadly reflected in Table 2 and supported on a local scale by Figure 5, however, do not support any easily envisioned hypothesis of mutual interaction and fine coevolutionary adjustment between the two groups at a taxo- nomic level. Although some regularity in chem- ical cues for oviposition and chemical specificity is suggested, the data are still very limited and give little support for parallel diversification in the two groups. Much more complete biological and chemical information will be necessary be- fore any claims of biochemical coevolution can be advanced at this plant/herbivore interface. The present information already seems to falsify many attractive and conventional hypotheses. In view ofthe fact that Ithomiinae populations probably are controlled principally by certain limiting adult resources (suitable humid open- understory *pockets" and sources of PAs, see below), they may exert very little, ifany, selective pressure for chemical and phylogenetic diversi- fication on their host plants. In any case, the BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY 377 specific choices seem to vary appreciably with local ecological conditions. Under a moderate degree of Ithomiinae attack, some Solanaceae even respond by vigorous new growth and flow- ering (especially in Brunfelsia and in section Bre- vantherum of Solanum). Under these conditions, it is difficult to establish the most basic prereq- uisites for “classical” coevolution between her- bivores and host plants. AN ALTERNATIVE HYPOTHESIS: SEQUENTIAL COLONIZATION BY CHEMICAL ADAPTATION (TABLE 4) The results of chemical extraction, fractiona- tion, and Biblis/ Nephila bioassay of 48 ithomiine host plants are presented in Table 3. In the cases investigated so far the Solanaceae have shown in their leaves the compounds already reported or presumed for each species, in good quantities. If these chemical characteristics of the host plants are combined with the utilization patterns in Table 2, a most suggestive picture emerges, as summarized in Table 4. Nothing is known of the paleophytochemistry of these plants beyond the global variation in secondary compounds seen in widespread geographical populations of each genus today, but it may be presumed that they were diversified generically before Ithomiinae began attacking them in the New World (since the most advanced genera were attacked first) and probably similarly protected chemically to modern species. It then becomes possible to trace a series of small hypothetical advances in larval toleration of food plant chemicals, each facilitated by pre- adaptations on the existing host plants, which lead to a complete picture of chemical specificity as observed today (Tables 2, 4). The scheme is clearly oversimplified but is attractive as a hy- pothesis for the sequential colonization of plant hosts, already diversified taxonomically and chemically, by progressively adapting herbivores (Futuyma, 1983). It is in excellent agreement with the great diversity and apparent nonstorage by herbivores of the Solanaceae chemicals and the initial dependence of the Ithomiinae stock on PAs (see below). Thus, Apocynaceae-feeding Ithomiinae de- pendent on PAs probably did not adapt sufh- ciently to the great chemical variability in the Parsonsieae (seen today also), which in some cases repelled or poisoned larvae, in others left adults unprotected against predators and deficient in ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 378 - q ‘xq ZAMOOH TOS zt vL (0 I 69 88 SL 99 Ə UNIUOUDAIS DANIDG “TE *— O °g °v “xa AMNOOH 6L O'S 9'6 (0v90 OS 98 Oct LTT Ə SuƏ2Sə40qup SNISIUIP ` [£ *— V ‘xd AMNOOH 68L tr VL (0)r80 L8 98 ZI 06 ©) DsOffiaaag DIQOSSDA “OE *— x4 AMSUOOUV tL 67 vil O 87 88 48 97 oz 3 pupisəəu ‘Iu "ds qd ^67 *— q ‘Vv ‘xd AASHOOHV 7S8 Ct Tt (SOT €ç 98 €6 $9 Ə publsəəu sS1]ps4t/q `gZ LN AMATE vSL TT SEI (S60 YL 8L €9 O71 TS 1əu1puəS AJIQDAIW 'O *LZ _ xq AMYO'ld 799 Tt 891 (ID LI VIL SL ool Or fa 9zjuny :O (pueouojA) npon] 7 `9Z - x3 AMUO'Td 08. €O LI (O SZ OL IL 69 8c f wnsonxajf D `çZ LN AA^MO"IH S69 EZ 8901 (O tI OO! 98 FI OI f (¿) üeuipueS) wnsoyia 7 vc IN AA3O'TOg PLL 91 LLI (0D vO 67 LL Stt £8I ^ RUNY 'O (PUNY) tung wunoisdp?) c - xq ZAM T'l€d 908 Tt v9 (0 SI OL 48 ss zt Ə wnjuajnosa uo21s42d004T "zc IN AM"IHV 98L OF TOL (0Or80 t9 c8 S6 Hs 2 Suv48D4Af 7) `Z - xg ZAM'IO€UV IS8 6€ FS (SI)LTO CS Le SSI 09 NS siisoppios Dapunwoydty “QZ _ v AAT Od 8€6 CI L'€ (0€£VO TI $9 €9 81 D (erugrop) ‘ds 77 *6I IN AM TO «169 89 SLI (0860 ES €8 ZS 0€ o ‘ISAT XƏ IIVO 112440]UTA SaYyIUDIIAT ^8] - x4 AAT Od LES T6 es (ce)8sz70 SI 88 TZ 81 o upuvjpuaa (wmnaaaia]noy `S) "S. 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Adults increasingly exploited PA sources (Boraginaceae leaves, Compositae flowers) and may even have found some substi- tutes for PA precursors in Solanaceae (Brunfel- sia?). Since they were preadapted to bicyclic 8-carbon monoester alkaloids and their N-oxides (both PAs and tropanes fall under this classifi- cation), the generally well-protected, tropane-rich Solanaceae represented an open niche. At least two separate radiations could have moved onto these plants (discussion of the move over onto the Gesneriaceae must await further biological and chemical study of Hyposcada and its food plants; the move onto Brunfelsia, whose alka- loids are still enigmatical, was probably a side- track of one of the radiations). The line that col- onized the tropane-containing Brugmansia (represented by Placidula and Miraleria today) also encountered in these plants the steroidal bit- ter principles, to which they were also preadapted through their experience with cardenolides in the Apocynaceae. Tolerance of these bitter steroids permitted further colonization of the many gen- era defended by them, most of which also contain tropane alkaloids, sometimes with N-oxides. Utilization of Capsicum by this line also gave a gateway, through its steroidal glycoalkaloids and saponins (Table 2), to the immense and under- utilized resource represented by Lycianthes and Solanum, on which the more advanced genera persist today. The second radiation found tropanes initially in Solandra and relatives, and thence in Cy- phomandra, on which it adapted to tolerate the steroidal glycoalkaloids also present there and cada). As a final step, adapta- tion to saponins and strong-smelling oils present in some Solanum (especially section Geminata and relatives) permitted exploitation of the abundant Cestrum in the forest understory. In all cases, tolerance of toxins encountered pre- viously in the evolutionary history of each line could be maintained as new enzymes were de- veloped to detoxify progressively more effective classes of plant defensive chemicals, in many cases not correlated strictly with taxonomy and evolution of the plants. Indeed, some of the more primitive genera— possibly with a broader range of biosynthetic capabilities, according to present concepts of biochemical evolution in plants— seem to have been the last to be colonized by the most advanced Ithomiinae. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Persuasive evidence for this hypothetical sce- nario (Table 4) can come only from extensive studies of female oviposition preferences and lar- val tolerance of food plants or chemical com- pounds. Preliminary data (Table 2) and some initial experiments are encouraging: Placidula cannot eat Solanum, nor sun Aeria accept any Solanaceae, but Mec/ n use tropane- and withanolide-containing iim (Datura, Nican- dra), and some Cestrum-feeders continue to use Solanum and even Lycianthes. Experiments in this area inevitably will encounter problems re- lated to individual and local adaptation, lateral radiation (as onto Brunfelsia), and loss of ances- tral genes but should in the long run help to define the viability of this proposed sequential expan- sion of Ithomiinae through progressive chemical tolerance to Solanaceae food plants. HERBIVORES AS PHYTOCHEMISTS The chemosensory apparatus of the average insect is capable of detecting very small amounts of specific chemicals, either through smell (an- tennae) or taste (tarsal or buccal chemorecep- tors). This highly specialized analytical instru- ment transforms chemical structures into recognizable electrical potentials through me- chanical contact between compound and recep- tor and uses these potentials to stimulate a va- riety of response behavior sequences. It can be fooled at the receptor end by substances of dif- ferent chemical composition but similar confor- mation and spatial relations among active bind- ing sites. It is far more sensitive and accurate, however, than the majority of the instruments in the modern phytochemist’s laboratory. Its specialization is selected over many generations of advantages accrued by those who could rec- ognize a few chemical compounds and thereby compete better for higher quality food, defense, or mates. To the extent that ovipositing female Itho- miinae seem t ificity in host plant recognition (Table 2), which involves use of chemoreceptors on the forelegs ("drumming" on leaves) and antennae, they can serve as precise analytical tools for the phyto- chemist. If some chemical patterns are already evident on a very rough scale in the food plant choices of the Ithomiinae (Table 2), these could be used to suggest analytical methods for still- uninvestigated Solanaceae and predict chemicals in others that have undergone only preliminary analysis. 1987] BROWN—SOLANACEAE/ITHOMIINAE CHEMISTRY 381 E 4. Possible scheme for sequential colonization of Solanaceae by Ithomiinae through progressive TA preadaptive tolerance of secondary chemical classes. Substances Tolerated Step Ithomiines (genera) Plants Colonized l Tellervo, Elzunia, Apocynaceae: Parsonsieae 3E-K: PAs and N-oxides Tithorea, Aeria (Parsonsia, Prestonia, etc.) 3A-B: Cardenolides 2a Athesis, Placidula, Solanaceae: — 1C: Tropane alkaloids and 'raleria ‘apsicum N-oxides 2b Olyras, Melinaea Fearing a 2Q: Bitter steroids Hyposcada Solandra PA precursors? 2c Methona Brunfelsia 3a Epityches, Ithomia Acnistus, Physalis, etc. 1C + much 2Q (varieties) further Capsicum Some 1L: steroid alkaloids 3b Thyridia; New genus Cyphomandra 1C, IL, 2R: saponins 4a Napeogenes, Rhodussa, — Solanum subgen. 1L: glycoalkaloids Oleria Bassovia, Potatoe (sect. 4b Scada, Sais, ieee UM 2R; 1A; possible 1C Callithom 5a Garsauritis, H PIE Solanum (subgenera Leptoste- IL, 2R, 1A; occasional yalyris monum and Brevantherum) nicotine, pungent oils 5b Mechanitis, Dircenna, Pteronymia 6b Ceratinia, Pteronymia, Solanum (sect. Geminata), IL, 2R; 2Z: pungent oils Godyris, Hypoleria, Cestrum Pseudoscada, Greta For example, it could be predicted that the Solandra-Markea-Juanulloa group of genera, which support the danaioid larvae of Melinaea (Fig. 8K, V), might have some protective chem- icals similar to those of Apocynaceae-Parson- to be identified (cardenolides are strongly sus- pected) and should be investigated. Similarly, the strong-smelling essential oil of Solanum sect. Geminata should be chemically very similar to the like-smelling oil of Cestrum; this also seems to have escaped chemical identification. With- eringia could have been predicted to show phy- salins by Ithomia, which also recognized these chemical components with withanolide-struc- tures in Physalis and divided its specificity among c pound; at least one major substance, possibly with this type of structure, is present in the neu- tral fraction (A) of two Capsicum species inves- tigated (Table 3). Do the genera Deprea and Athenaea contain withanolides also? Start by testing some coop- erative females of Ithomia and Epityches. Are any other Solanaceae besides Datura/Brugman- sia especially rich in scopolamine? One cou begin with some Placidula females or perhaps even larvae. If the latter were found on a Sola- num or Cestrum, I would bet on the presence of tropanes in the leaves. Are there glyco-alkaloids in Dunalia? I might start by asking Hypothyris, known only from Solanum, or perhaps even Pteronymia, which specializes on the high-al- kaloid (Bradley et al., 1979; Table 3) Geminata section of Solanum. This analysis could be ex- tended even to individual constituents once ap- propriately biis populations of Ithomi- inae have n identified. It may also give some false UE fee these will hold even more in- terest than the expected ones, pointing out new types of plant components. As a preliminary sur- vey, it should be strongly suggestive of certain types of chemicals. CHEMICAL CONTRIBUTIONS OF THE SOLANACEAE TO PREDATOR DEFENSE IN THE ITHOMIINAE ITHOMIINE CHEMISTRY Chemical investigation of the passage of So- lanaceae toxins to adult Ithomiinae started with 382 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 5. Summary of PA content and distribution, and bioassay of adult Ithomiinae and Danainae. No. of spp. Ana- Tribe Genus In Genus lyzed Typical Species? Source? Host Plants Analyzed ITHOMIINAE® B Tithorea* l harmonia? VB 45 21/16 C Aeria" 3 3 olena: C 43, 44 31/20 elara G 10/15 E tho 7 5 themisto'e C 41, 42 36/27 F Placidula" l l euryanassa* C, J, SL 33 24/17 G ras” 2 l rathisk V 37 4/2 Melinaea” 10 7 ne RO 36 5/4 ludovica* M 36 50/1 menophilus RO 37 15/15 H Thyridia l l psidiï C 20, 21 10/10 Sais 2 l rosalia' G 19 12/12 Scada 6 2 reckia* RO 16 35/13 karschina ES 16 15/4 Mechanitis 5 5 polymnia' C 6-13 44/37 M 4 12/12 lysimnia'9 C 13-15 28/14 I Hyposcada 8 4 egra* M 36 11/1 Oleria 30 13 astrea BJ 12 4/12 RO 12 13/7 J Epityches l l eupom C 25-31 30/33 Rhodussa l l cantobrica RO 8 17/9 Napeogenes 18 8 cyrianassa ES 18 10/8 inachi RO 18 10/11 Garsauritis l l xanthostola M 13-14 48/20 Hypothyris 16 13 ninonia daeta* C 5-14 51/40 euclea’ C 9 22/22 (males attracted) M 10 16/12 V 10 17/14 K Ithomia 21 7 agnosia'* C 30, 31 29/38 L Callithomia" 3 2 lenea xantho* MP 16 18/21 Dircenna 7 4 ero* C 6-13 29/18 M Ceratinia 5 4 neso” BJ, RO 13 24/24 Ceratiscada 3 2 doto BJ 1? 6/7 Prittwitzia 1 l hymenaea" C l 10/12 Episcada 15 5 clausina ES l 17/14 carcinia C 1? 25/29 Hyalenna 5 l pascua“ J 7 1/9 Pteronymiq" 41 10 carlia* € l 31/17 dyris 10 3 zavaleta* RO 38 10/9 Pseudoscada 6 4 ca* C 39 31/28 reta 10 l andromica" V 38 4/4 Mcclungia l 1 salonina* C 39 11/11 H ypoleria 11 10 arzalia G, VB 39 8/4 Heterosais 3 3 nephele VB, RO 40 16/11 giulia V 40 7/4 DANAINAE Danaus (Anosia) 2 2 gilippus' C Oxypetalum 25/16 Danaus (Danaus)^ 1 (S.A.) l plexippus C Asclepias 22/14 Ttuna* l ilione C I 6/5 Lycorea" 2 2 cleobaea C Ficus 12/11 1987] BROWN—SOLANACEAE/ITHOMIINAE CHEMISTRY 383 TABLE 5. Continued. Individual PA Analysis (5/9)? Odes Fractions: Average Maximum Bioas. s Average PA conc. (% av. Ce average dry wi, am Fract. Re- (mg) % N-Ox.) dry wt.) A (neut.) C (polar) F (fat) Tested sults! 60/75 0.3/0.3 (48/45) 0.9/0.6 8.4 8.4 13.9 Ex + 9/13 2.5/1.6 (55/55) 7.1/4.0 3.9/5.2 14.7/15.8 13.9/16.3 Ex + 9/13 1.3/1.1 (61/57) 1.9/1.9 NT 100/120 0.06/0.1 (41/48) 1.6/0.9 3.2/1.9 7.2/7.0 10.8/26.6 Ex + 54/65 0.2/0.2 (48/48) 1.6/0.9 Ex +* 110/110 1.2/0.3 (54/49) 1.8/0.5 NT 55/60 6.3/1.2 (54/49) 10.5/1.5 NT 55/60 0.6/1.3 (57/59) 2.5/— 5.9/3.7 7.2/9.6 15.7/13.9 Ex + 55/60 3.6/1.3 (50/49) 6.5/4.4 NT 64/70 2.7/2.2 (70/68) 5.1/3.9 3.5/4.3 9.3/7.1 12.3/20.7 Ex *h 25/27 3.6/1.6 (59/58) 6.8/3.6 7.0/7.2 12.7/7.4 20.7/21.6 Ex + 6/7 9.7/3.2 (47/54) 20.6/5.3 NT 8/9 13.1/6.7 (61/65) 20.0/10.4 NT 33/45 1.8/2.4 (60/60) 5.8/4.0 8.9/5.8 6.4/6.7 14.1/23.8 Ex, B, C + September 5.6/6.1 5.4/3.0 9.7/18.9 A, F _ 35/45 4.4/2.9 (58/57) 7.6/4.7 6.2 8.3 10.7 NT 25/35 3.8/1.8 (50/55) 8.8/6.4 Ex + 20/22 0.7/0.2 (51/48) 3.8/— NT 13/15 2.0/1.9 (53/58) 4.5/4.0 NT 13/15 4.0/1.8 (50/43) 8.2/3.4 NT 24/26 6.9/3.2 (56/58) 11.4/4.3 2.3 11.4 11.9 Ex + 18/20 3.1/3.3 (48/52) 8.6/6.1 NT 18/20 7.7/3.5 (55/56) 12.9/5.9 NT 13/14 5.4/3.6 (56/54) 11.7/6.2 NT 20/23 0.9/1.0 (53/53) 4.9/3.2 NT 25/27 3.3/2.8 (51/57) 9.2/6.5 3.5 6.3 21.8 Ex + 22/25 2.2/1.5 (48/44) 9.3/5.4 4.2 10.3 12.2 Ex + 20/23 0.8/1.5 (55/46) 2.9/4.5 5.3 10.8 10.4 NT 20/22 5.1/2.8 (58/60) 9.0/5.7 NT 14/16 5.4/2.0 (56/58) 12.9/7.1 1.9 10.6 23.2 Ex + 24/28 0.5/0.4 (52/53) 2.6/1.5 NT 40/45 1.7/0.9 (65/69) 4.3/2.4 4.0 9.9 12.3 Ex + 15/17 2.0/1.0 (55/56) 5.7/3.3 Ex + 14/15 1.1/0.5 (52/48) 1.9/1.2 NT 14/16 2.8/1.7 (57/60) 5.8/3.5 4.6 7.4 19.5 Ex, B, C + 16/17 5.7/5.3 (53/58) 10.0/7.0 A, F NT 17/18 1.6/2.0 (51/58) 6.0/5.2 Ex + 18/20 1.1/2.0 (47/52) —/3.4 NT 14/17 7.8/3.6 (58/62) 13.3/5.8 Ex + 33/33 1.5/1.4 (43/51) 2.9/3.1 NT 15/17 4.9/2.7 (57/54) 9.7/5.8 4.9 14.0 9.6 Ex F 15/17 7.2/5.0 (56/52) 10.6/6.0 NT 16/17 1.7/1.1 (70/82) 3.3/2.2 5.2 7.5 15.5 Ex + 14/15 1.1/0.5 (56/59) 2.3/0.9 NT 21/26 1.6/0.6 (55/53) 4.4/1 NT 20/24 2.7/0.8 (53/43) 5.3/1.7 4.8/— 9.6/— 8.2/— Ex + 80/75 2.1/1.8 (57/52) 4.5/4.5 NT 180/170 0.1/0.1 (50/51) 0.4/0.2 NT 160/140 1.1/1.2 (54/53) 2.5/2.1 NT 130/110 2.7/1.2 (58/55) 4.8/3.2 NT 384 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 5. Continued. ANALYSIS OF PARTS:™ VINEIS Appendag Species & Sex Basal Hairpencil Apical (Legs, esti ) Rest of Head Mechanitis polymnia 1.13/88 4.15/64 0.14/100 0.26/77 0.60/70 casabranca (males) 1.70/100 1.10/100 1.03/65 0.52/92 1.03/72 1.03/58 1.70/58 0.83/51 0.98/97 1.43/75 2.60/85 4.40/100 2.10/48 4.20/77 3.40/75 Ithomia agnosia 4.50/81 — 1.38/78 7.50/68 12.5/69 agnosia (females) 0.69/42 - 0.63/45 1.60/55 2.00/68 0.70/67 — 0.75/48 0.72/100 1.05/44 “pockets” Danaus gilippus (male) 3.10/63 1.00/55 0.73/49 2.00/57 1.75/55 (female) 1.45/50 — 0.47/42 1.20/46 1.50/51 Displaying Male Rest of Insect Spermatophore Mechanitis lysimnia 3.09/61 8.50/54 3.32/59 8.85/55 3.06/52 9.75/49 0.78/54 2.40/64 4.20/52 25.3/55 6.40/56 17.4/54 Mechanitis polymnia 2.23/52 8.60/62 0.26/46 1.05/54 0.92/62 3.92/55 Aeria olena 3.00/50 12.3/67 3.60/69 10.0/72 * Authors’ and subspecies names are in Figure 5 or Brown (1985). > See Table 3, footnote b. Sp din localities: BJ — various parts of Rondónia. VB = Vila Bela, western Bujaru, Para. ES = various parts of Espirito Santo. RO = at ro Numbers refer to the plants aire in Table 3. Italicized numbers indicate probable food plants or species near to that number. * Just-captured butterflies were cut up under 2 ml absolute ese As ml for heavier species) and after at least one day's standing aliquots (usually 1/20, or 1/4 for componen were assayed directly for total PA and N-oxide, following Mattocks (1967, 1968) sien l Bingley (1968); see sol for details. * See fractionation scheme in Figure 4 for oo ' See footnote g of Table 3 for bioassay co * No secondary compounds from larval teh plants have been found stored in adult Ithomiinae. Additional and Hyalyris oulita metella, 13; K Miraleria cymothoe, 31 + 39 and Hypomenitis dercetis, 39 and H. libethris, 39. Rothschild (1973), who obtained only negative results. In August 1978, 3,200 dry bodies of Me- chanitis polymnia (75 g, representing over 300 g fresh weight of insects) were mailed to Dr. Desiré Daloze of the Collectif de Chimie Bio-Organique in Brussels, where ant bioassays were used to follow repellent activity in the fractionation. No alkaloids, cardiac glycosides, or other interesting 33: L Velamysta pupilla, 17; N Dygoris dircenna, active compounds, or even their degradation products, could be found (D. Daloz comm., sults suggested that the protective compounds o adult Ithomiinae might be labile, degraded after death or upon storage, and possibly unrelated to the larval food plant poisons. Total MeOH-H.O extracts of fresh ithomiines 1987] TABLE 5. Continued. BROWN—SOLANACEAE/ITHOMIINAE CHEMISTRY 385 THORAX I ABDOMEN ton Muscles Exoskeleton Fat Reprod. Organs Intestine 0.29/75 0.21/100 0.21/92 0.49/65 0.12/91 0.02/100 NR 1.18/42 0.92/48 1.82/91 0.81/66 2.54/99 0.63/89 1.02/66 0.63/65 0.50/100 0.19/69 0.60/100 0.14/56 4.10/67 4.40/68 6.60/72 3.20/100 4.10/92 4.00/97 Eggs 1.06/79 3.56/50 4.45/65 1.76/65 52.0/45 0.13/91 8.85/78 0.35/45 0.47/20 0.47/38 0.42/54 1.67/40 0.28/14 1.00/30 0.62/48 0.40/33 0.87/100 0.26/18 0.34/100 0.15/60 NR 1.90/57 1.20/50 2.20/56 0.73/65 4.20/58 Hairpencil 3.50/55 1.60/52 1.10/48 1.40/62 3.30/59 0.90/62 ggs 0.90/12 Pairs captured in copula Male: Rest Spermato- Female: Rest of Insect ore of Insect Abdomen Ducts 2.12/52 12.8/65 6.26/52 3.50/59 2.23/64 11.9/54 0.40/74 0.35/57 3.71/54 12.7/66 2.26/62 0.83/100 0.44/56 1.31/78 3.00/62 3.30/60 1.29/50 22.5/60 3.30/51 3.51/65 0.25/54 4.70/48 3.58/63 5.10/62 0.87/52 9.10/61 2.52/48 3.26/58 0.38/48 14.6/72 2.06/72 7.20/88 2.96/59 12.6/72 5.06/61 13.6/63 2.26/51 2.32/57 3.90/58 40.5/64 8.90/62 18.7/77 4.10/54 5.40/59 h Larvae of these species may be considered as aposematic in a and behavior. PAs. ! Eggs of ee subspecies contained up to i Larvae of these species feeding on the indicated plants did al contain any PAs nor did their extracts protect 9% of dry weight a Biblis Medus predation by Nephila; possible exceptions are Methona themisto, Aeria olena and Tithorea har- monia. * Recently emerged adults of both sexes of these species reared from larvae on the indicated food plants, were consumed without hesitation by Nephila, which in most cases had just cut out an adult of the same species and sex gia in the m a are giv field. Notice that almost all S n as percent of dry weight of part/% of PA a -oxide. usually papie pila PA present as A ropy bem DB dd confirmed. Especially a m a an E aga in the food plants. oteworthy values are printed in boldface type. (prepared as in Fig. 4), when applied to the edible Biblis hyperia, were at least as repellent to ur ila spiders as were the live butterflies. re pellency was then located in the alkaloid taa B (Fig. 4) and also in the polar fraction c. Wh these fractions were compared with the corre- sponding fractions of the larval food plants, there appeared to be no compounds in common. Fur- thermore, all food plant extracts and fractions tested in the spider bioassay were negative (Table 3). To put the final nail in the coffin, both sexes of 30 species in 26 genera of Solanaceae-feeding N-oxide value is high, it Ithomiinae, reared from the larvae in the labo- ratory on fresh leaves of natural food plant, upon emergence from the pupa and introduction into Nephila webs were promptly and enthusiastically eaten (Table 5). In most cases, the Nephila had just as efficiently rejected a field-captured adult ithomiine of the same species and sex. In all, 142 species of Ithomiinae in 45 genera and all 14 tribes were examined in parallel with 48 host plants in 16 genera of Solanaceae, three genera of Apocynaceae, and one of Gesneriaceae (see Tables 2, 3, 5), both in chemical analysis 386 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Ithomiinae found in the Campinas region, interior of Sào Paulo state, EE Brazil. Missing are FIGURE 9. *Episcada philoclea (Hewitson) and TPseudoscada quadrifasciata, very similar to U. e to aa ninonia daeta (8%). —O. Dirc thona themisto (s š; on tree-lined city streets and in gardens ian , 7596 in some seasons). PD. Mechanitis lysimnia (1796). — —L. Ithomia agnosia (5. nna dero (upper D. celtina, lo ower D. rhoeo) (0.8%).—G. Hypothyris euclea (laphria x nina, mixed Smal aod (0.4% but can be comm i j i 's common in September). - B. *Melinaea ludovica paraiya. — —E. Mechanitis polymnia n (about 50% of S. Mcclungia salonina 5%).—N. Aeria olena (2. 5%).—P. Prittwitzia . The over 20,000 captures, mostly for marking and recapture population studies. For BM order of genera and species (primitive to advanced), see Tables 2 and 5; for authors’ names, see Figure 5. and bioassay. In terms of the hypothesis of se- questration of defensive chemicals by ithomiine la arvae, these results were thoroughly disappoint- ing; no important compounds were detected that were shared by butterflies and their larval food plants. All butterfly extracts were positive and all plant total extracts and fractions were nega- tive in the Nephila tests. However, all butterflies showed a strong Mayer's test on the acidified total aqueous extract, suggesting the presence of some kind of alkaloids. Since adult ithomiines, like danaines, are known to be strongly attracted to sources of de- hydropyrrolizidine alkaloids (Pliske, 1975a, 1975b) and use these at least for pheromone syn- thesis (Edgar et al., 1976), the extracts were ex- amined for these compounds, using on TLC the iodine/Ehrlich test (de presence of similar moderately polar PAs as the major components of the alkaloid fraction B; in all cases, this fraction was greatly augmented and often became over 9096 of a single compound after the zinc-reduction loop in Figure 4, indi- cating that much of the PAs were present as a single alkaloid and N-oxide in the butterflies, the latter probably responsible for the activity seen in fraction c. Pure pyrrolizidine alkaloid frac- tions accounted for essentially all the activity 387 MLICLAICT DV BROWN-SOLANA( 1987] `[ Aq pawos) si2^0]p 10 spnq 2477081420] wuni401pdn:] WOJ Pepe [e101 paying ^4— (pozij[e1s&ro Aprseo. 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A (0 662 Bet = 0 AN, (suu apuTy) (autmesdooK1) IuaNquvo HnlsOlvdld v 6° ozi 68°) do saol 3 aa Š 30 Su3M014 0 [4 £ ° s A aia e Y + — y » | t € y s 9 maia 4 a a CS SE RER, | | SRL | (auto TPUI) | | An2IGNI | (əutmesdooK1) ort WNIdOULOI14aH | 40 S3AY371 eurpeuisjur VONVUGVSVO rr ree VINHATOd 343043V21NOI38 SIIINVHO3H N RnIsOlvana3 TEO 30 S43M014 v 388 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 E ll. Feeding of dui at PA sources. Bess ree rs, Amarais, m flowers, Colorado, Rondóni seen in the Nephila bioassay, protecting the adults against predation by this spider. he most common species in the Ithomiinae communities in the Campinas area, Mechanitis polymnia casabranca (Brown & Vasconcellos- Neto, 1976; Vasconcellos-Neto, 1980, 1986; Fig. 9), was chosen initially for detailed chemical in- vestigation, using the fractionation scheme shown in Figure 4. e ethyl acetate extract (F) from 455 butterflies captured in Amarais in August 1982 (19.1 g dry weight, 13 of this as insoluble tegument) weighed nearly 5 g and readily solid- ified at 8°C to an off-white crystalline mass, in- dicating nearly pure saturated triglyceride; its ex- act composition is presently under investigation, but it was negative in the Nephila bioassay — probably quite nutritive, in fact. This extensive storage of high-energy fat (2696 of dry weight at the height of the winter dry season) reflects the long (up to six months) reproductive diapause of these species (May-October; Vasconcellos-Neto, 1980) and also helps to explain the advantages of the learned predation behavior of the tanager (Brown & Vasconcellos-Neto, 1976), which squeezes this fat out of the abdomen of the but- terflies. It is interesting that the fat reserves fell to only half of August levels in butterflies ex- tracted in late September; the butterflies are long-lived and have few exogenous resources i ward (here confirmed) and more diffi- culty in capture as the days become warmer Brown & Vasconcellos-Neto, 1976). The total alkaloid extract B, isolated after the zinc reduction loop (which increased its weight from 60 to 260 mg equivalent yield from 19 g butterflies — 1.496 of dry weight) showed on TLC —A. Mechanitis polymnia casabranca on Trichogon s, SP. — B. Mechanitis lysimnia elisa (Guérin) on Eupatorium ape baile hii a single major spot (90%) that gave a positive I,/ Ehrlich test for PAs. The 60-MHz NMR spec- trum of this noncrystallizable fraction (Fig. 10) showed it to be almost exclusively a 60: 40 mix- ture of lycopsamine and intermedine (Fig. 3E, F) compared with a spectrum of pure reference ly- copsamine contributed by J. A. Edgar (Fig. 10C). Very minor impurities due to other dehydro- pyrrolizidines could be seen in the NMR spec- trum. The alkaloid fraction co-chromatographed with authentic lycopsamine (donate J. A Edgar) on TLC in four different systems, giving indications of latent separation of the isomers in some solvent mixtures. he NMR spectrum of the total alkaloid frac- tion showed no signs of the “methylene enve- lope” or quaternary methyl signals typical of ste- roid alkaloids of Solanum, nor of the anomeric protons of glycosides, nor of the N-methyl groups of solanines, nicotine, and tropane alkaloids (Figs. , 10). This supports nonsequestration of al- kaloids from the larval food plant. The spectrum also indicated that most of the butterflies ob- tained their alkaloid from a single source; the structure and the mixture strongly support flow- ers of Trichogonia gardneri, common in Amarais and intensively visited by M. polymnia (Fig. 11A; Table 6). When the zinc reduction was not performed, the much diminished alkaloid fraction showe — he NMR spectrum of the whole fraction was perum complex, indicating several additional components of diverse structures in- cluding saturated PAs, but still showed no signs of the presence of other classes of alkaloids. 1987] In expansion of the work on M. polymnia pre- liminary fractionation gave PAs representing various mixtures of the five isomers of a single structure (Figs. 3, 10) in up to 1396, and in Scada over 2096 of dry weight in some individuals, in all 141 additional Ithomiinae species indicated in Table 5, many captured in the Campinas re- gion but also sought in more distant places for comparison and to verify the generality of the phenomenon. No PAs were seen in the Zn-re- duced extracts of any of the food plants tested (summarized in Table 3). ITHOMIINE DEFENSE That th the prot ti of adult Ith predation by Nephila is due to PAs was con- firmed by feeding 200—400 ug of echinatine N-oxide (Fig. 3J) in dilute honey solution to new- ly emerged adults of Mechanitis lysimnia and Pseudoscada erruca (which normally were eaten by Nephila); within an hour, the butterflies were routinely rejected by the spiders (experiments performed together with J. R. Trigo). Reared but- terflies kept alive for many days did not biosyn- thesize any protective chemicals; protection was lost within a day after death ofa butterfly rejected by the spider, again indicating the instability of the PAs. Very fresh field-captured Ithomiinae often showed no PA and could be consumed by Nephila, but in general the adults seemed to be able to accumulate sufficient defensive com- pound from different sources in their environ- ment within a very short period after emergence from the pupa. The strong dependence of adult ithomiines on PA sources, including for their courtship pher- omones, perhaps made it inevitable that they should also retain the PAs for their defense. The lack of storage of Solanaceae defensive com- pounds, already verified in early investigations, increased the probability of alternate defensive compounds in the Ithomiinae—labile com- pounds not detectable in dead insects. The uni- versal rapid cutting out of Ithomiinae from ephila webs suggested that such alternate de- fense substances did not derive from the widely variable larval foods but from a more homoge- neous adult food source. In the only other re- ported case of a lepidopteran being cut out from spiders’ webs— Utetheisa ornatrix, an aposemat- ic day-flying arctiid whose larvae feed on Cro- talaria and pass sequestered PA diesters on to the adults, where they are also used in phero- mone synthesis— it has been shown that PAs are . BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY 389 baise: for this deterrent activity (Conner et 1981; Eisner, 1982). The Solanaceae poisons have thus been ab- solved, at least for the time being, from partic- ipation in the unpalatability of adult Ithomiinae to their most dangerous predators. It seems pos- sible that exceptions will be found to this, es- pecially as more predators are incorporated into the bioassays and more butterflies and food plants into the chemical analysis. However, the PAs and especially the biologically active (hepato- toxic, tumor-inhibiting) lycopsamine-group monoesters and their N-oxides should be re- garded as the principal, perhaps nearly universal inae tiidae moths (Ctenuchinae, Pericopinae, and Arctiinae), whose adults sometimes inherit them from the larvae but inevitably seek them out at the same sources visited avidly by the Ithomi- inae, and similarly use them for defense and pheromone synthesis. Indeed, lycopsamine and its stereoisomers have been found in wild pop- ulations of essentially every species of Danainae and Ithomiinae investigated (Edgar, 1982; Table 5), though their presence is erratic in the moths. These compounds seem to represent a very ef- fective “ancestral predator defense" that has been retained in diverging phyletic lines up to the pres- ent (Edgar, 1975) in spite ofa variety of habitats, food plants, and behaviors. Adult Ithomiinae sequester their PAs from a variety of sources: decomposing borages and composite flowers (Eupatorieae) especially, but also orchids, Crotalaria, apocynes, and less tra- ditional PA-containing materials. These are inevitably abundant wherever Ithomiinae occur in numbers. The proponderance of PAs in adult Ithomiinae defense could also help to explain the cryptic coloration and behavior observed in most ith- omiine larvae (Fig. 8), notable exceptions being Methona on Brunfelsia (Fig. 8W, X), the feeders on Apocynaceae (Fig. 8T, U), Melinaea with similar larvae (Fig. 8K, V) feeding on Markea, Juanulloa, and related genera, and a few showy larvae scattered on other plants. Newly emerged ithomiine larvae eat their eggshells (which con- tain PA derived from their mother; see below) and immediately move to the underside of the same leaves (if not already there), usually ac- quiring a cryptic coloration. The fact that most ithomiine immatures are cryptic translucent green, closely matching their substrate, and feed [Vor. 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN 390 IN ¿ ville VS CSI (89 OF E'S 69 Ic L9 Ə (so^e9[) IN ¿+#d Ors OS Sit (Ss) r9 61 ZL 9'8 IE Ə (si9AÑojj) `OG Dp40]/lA94q plliojəouuno L LN IH O18 90 ct! 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(09 t'i xs tL 8€ tl 9) qu A EIC) `V 142Up4D8 DIUOBOI(214 | at Ə ‘a xd — 9vL € THI G) $0 RES 89 6L 9c Ə "PIA 21/0f1p402 DIUDAITIA 4(S19:40]) IVAIMOLVdN :3v.lISOdWOO at xq ¿ 978 8E TII (i) Lt'O 6I v9 Lil $9 TS ux OC (JOM) DiJOfJƏDUDII9]DA $21112247] y(SIIMOY) IVANOIDANAS :3V.LISOdWOO sss — ,polso[ iSVd aq J O ALXON% $d V OH Aq usaq MOŞ sƏ910moS poo, 1[npy Paid MM pM AIP JO % SB) suonog14 % 143M -org “QBUTIWOYI] 1[npe Jo s2o1nos pooj jo Áesseoiq pue 'uonnquisip pue 1uəluoo vq Jo Aireurung `'9 318v] 391 BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY 1987] `ç 9|qe[ ‘U 2310u109J 995 wu 'so[nsdeo pəəs Ap pue 'poreurjod Apearye samo uədo ‘png 1940] Iop[O WAS 19A0| ‘SIJALI IIE] :s 1ed 1op[o gurpuodsau1oo sjuasaudo1 so1ngy JO JUI 1240[ JL x opueir) ory ‘IZIV OOd = Vd '"OJmuef op OTA *erenej ap ens = ‘OSL, ouaqoy sof Aq payeuoroey pue paj29[[0)) + (8161 '19qeH :9/61 ‘PuowuNig `qç/61 '€cz6T “Ə3StId) 1e1mqeu sm jo elJoidopidoT Supjoss-yq 1910 pu? aeurruioui] Aq poieumod Aq[eoyroods pue ÁAqpeordÁ? “sISAOJ 114M pews *poie[ost YIM *Aro]s1opun 1sə1oj pruny “Hep əy} JO sjue[d o[rgu1j ae 1un140]Dpdn;] IWOS pue putua]souapy | Tens sem (SƏ19n[OAUI — 'S12|1240]) SIOMOY I} JO IONUT IYI ATUO '(,,) PASN SI YSLI9}se UB UY AA 'soannuenb poo3 UI Syd 3UIMOYs [Ie ‘I sapiozíuoo wnjp428y. pue Od wnprpnbs `: ‘COA ə2pqlpha `T “JE Surureywos Á[qeqoud “qoq sidajojsax wni401pdn;] 319M pouruiexa OS[V « '€ DINBLJ ur suoneungguoo [eoruroq202Jo9]s PUB sa1njonujs 99g , I ?deury ‘uedewy ory 1oddn = `ç 9|qe L Jo 3 pue ə ‘p ‘Dd $310u100]J 29S ssp "'o[neq oes ‘OIJ Ory = OU Th S op dV :Sani[eoo[ [euoriippy `ç pue ç So[qe [ JO q $20u109J 23S 4 “UOTIIVIIXS 10} pai29[[09 219^ si[npe oeuruioui] Aq A[oAtsuo1ur Pasn Sutəq A[[enjoe SIdINOS pooj *ojqtssod 13A9U3Q e 66/660 ES/IL'O 8vP/0t'O Ovr/890 9r/S€'I L£/9€0 tS/£D€ c£€/8L0 | —/S000 €S/c00 (69/91'0 :$1001) (ased 1op[o) CV/L9O ¿b/9U I €€/£€0 vtS/CCl 9S9/PC€ | L9/VCl 09/080 SS/v9C OS/9CI 8S/tCO €tC/tlO 2 (a9quiəsəq) th/6C0 CC/8t€0 cS/81TO LS/CC€ | 6V/I10U1 —/v000 —/0000 (çç/9ç`0 :$1001) (,su1ed 13p10) 19/LU1 OS/S6C 6¢/CL0 = Lv/v9'I 0r/0t'€ CcV/19'O 89/60F — 9v/L9'I 88/L00 | 9v/6S90 ‘IS (190) 2u0foporu012q wini40mDdn 7] LS/L8°0 L1/090 | OL/C8O LI/OTZT O9vt/6L0 0Ot/SCO 0891 tcC/IwO EI/ETO w vS/VCO 2 (ounf) nuvipuaixpua uantaolpdn:] IZ/8€0 L£/08'O 8€/090 ZTS/EO'S O9/0DL 9L/8tC Lv/LOC X 8V/t6óv 8t/98C tUl/£60 6I/c8' 2 (19quioidos) 9t/V0 Tl (6v/£lc L8/P60 À6v/06c CE/OTR 09/0UT* 6L/9V'I LL/OCCc £€L/OL'I 8U/vTCO ¿I/0S € 2 (sun) 142up4p3 Diu0301214] SISAO[4 spass ƏmpnƏoAu[ ued SƏUVJJƏN SoHUueA() Ə2ƏJO9n[OAU]| SIoMO[J BWON[OAU]T səPunpəd SALIT (q991n0S) so1dedg oeouojedn AIq 1odd() SunoA pue swag SunoAa u:/SLHVd LNV1d dO SISATVNV so[nsde;) psag S19^0|, one] spng 21njeuirui 'peanunuo) `9 gv L 392 on the underside of leaves (Fig. 8) may indicate rapid metabolism of the Solanaceae poisons (though cryptic and toxic insects are not rare); at least the poisons are not carried through to the adults, whom they would not protect in any case, and do not seem to be stored in the larvae. Apo- sematic but nontoxic insects are rare (except for obvious Batesian mimics that diverge from their taxonomic relatives), the main case being auto- mimics in the Browerian sense (Brower et al., 1971; Rothschild, 1979). For this reason the strongly aposematic larvae in a few ithomiine genera may incorporate or synthesize some sorts of unpleasant substances that have yet to be iden- tified. In previous work reported without details, a lycopsamine/intermedine mixture was found in Hypomenitis dercetis and Oleria makrena from northern Venezuela (Edgar et al., 1976) and re- garded as primarily precursorial to pheromones (only males were analyzed) rather than as defen- sive. Later work (Edgar, 1982) emphasized the potential defensive role of these compounds, here confirmed SELECTIVE PA STORAGE When male and female Mechanitis polymnia were separately extracted in late September 1982, some interesting differences were observed, in- dicating that careful work on this chemical in- teraction should always maintain the sexes apart in analysis. At least in September, at the begin- ning of the reproductive season, the females con- tained much more fat and PA than males (Table 5). That this difference is closely related to the respective reproductive tasks is supported by the previous pheromone study (Edgar et al., 1976) and by analysis of Mechanitis eggs. A raft of 31 M. polymnia eggs, weighing | 1 mg, was extracted with 2 N H,SO, and directly reduced with zinc, alcalinized, and extracted with CHCl,-MeOH. This showed on TLC the presence of abundant lycopsamine/intermedine mixture, perhaps as much as 1% of the fresh weight or 0.1 mg. All other Ithomiinae eggs analyzed also showed the presence of appreciable PA (Table 5). The bright white Mechanitis eggs, laid in rafts of 5-100 on top of the host plant leaves, with which they contrast vividly (Fig. 8I, N, Q), could be de- scribed as a true collective display of aposematic insects, just ie the adult assemblages in dry season **poc As ei in PENIS 5, male Ithomiinae gen- erally accumulate more PAs than the females; ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 indeed, in many species they are the principal or only sex found on the sources (Table 2; Pliske, 1975a, 1975b). The females of most species seem to get the majority of their PAs in the spermato- phores received from males during mating, which is repetitious (Ehrlich & Ehrlich, 1978); these small sacs often have 20-50x the PA concen- tration as the rest of the male (Brown, 1984, 1985). Both males and females are able to selectively distribute the collected PAs to different tissues, including tegument and wings (possibly by re- gurgitation with fat as a wetting agent) and es- pecially to reproductive parts (pheromone glands, spermatophores, ovaries; Table 5), thence in fe- males to the eggs. Males attracted to Heliotropium baits placed in relatively PA-poor areas and females attracted to displaying males showed lower average and maximum PA content than random samples of the same populations, indicating an "appetite ef- fect" that obviously would be highly adaptive in these organisms. More complete information on PA storage, use, and distribution in Ithomiinae may be found in Brown (1985). SOURCES OF DEHYDROPYRROLIZIDINE LKALOID MONOESTERS In the same paper in which he predicted that PAs would be used for defense in the Ithomiinae, Edgar (1982) predicted that they would be found in the nectar of Eupatorieae, in view of the heavy dependence of Ithomiinae on these plants and their characteristic occurrence in the Boragina- ceae, also frequently visited by Ithomiinae and source of precursor for the pheromone (Fig. 3D) as well as of the most attractant esterifying acids for ithomiine males (Pliske et al., 1976). Indeed, many genera and most females are more strongly attracted to Eupatorieae flowers than to Heli- otropium (Table 2), the difference being espe- cially pronounced in Mechanitis and allies (Fig. Extraction of the flowers of 16 species in four genera of the Eupatorieae actively visited by Ith- omiinae in the field led to the isolation and iden- tification ofa variety of PAs but usually only one isomer and structure in each species (Table 6; see Figs. 3, 10). Alkaloids of this structure were also found in Heliotropium and Tournefortia flowers (Boraginaceae-Heliotropoideae) often visited by Ithomiinae, as well as in the leaves of 1987] Removed by pollinator AN N Concentrated in nectary Translocated to flowering tips & leaves IVA P WA Synthesized in roots FIGURE 12. various Heliotropium species (Table 6), but not in Mikania (Eupatorieae), Cordia (Boraginace- ae), or several other flowers sporadically visited by Ithomiinae. When the plants were analyzed for PAs by parts (Table 6), the highest concen- trations always appeared in the nectaries of still unopened flowers, suggesting that they might be a reward to specific PA-seeking pollinators; the BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY tegument Used for pheromone synthesis Concentrated in reproductive organs, especially in the spermatophore Transferred d to female in mating 393 Pheromones define communities and territories, attract females Spread out over tegument, \ concentrated in UN i N NN reproductive NN \ NN. tissues, and Nec A N placed on ` Nee! eggs. e \ KS VERNY Flow of dehydropyrrolizidine alkaloid monoesters in natural ecosystems. concentration was often 2-4% of dry weight of the whole flower. In open vegetation or poor soil areas with few Itl ii t, Eupatori ] yields of less pure isomers in the flowers and showed dramatically reduced seed set, even though pol- lination was effected by other groups that also depend on PAs (Danainae and Arctiidae). Areas 394 with few E r only very seasonal species showed scattered and transient Ithomiinae pop- ulations, whereas in areas with abundant Ith- omiinae there were always common Eupatorieae in flower throughout the year, especially Tricho- gonia (Fig. 11A; see also Pliske, 1975a). It is evident that this mutualistic relationship, pro- foundly affecting the reproduction and abun- the interaction, in a coevolutionary picture prob- ably much stronger and more stable than that of the Ithomiinae with the Solanaceae. FLOW OF DEHYDROPYRROLIZIDINE ALKALOIDS IN NATURE (FIG. The analysis of PA monoesters in different parts of plants and butterflies in various physiological and reproductive states (Tables 5, 6; Brown, 1984, 1985), made possible by the selectivity and sen- sitivity of the Mattocks-Bingley assay (Fig. 3), permits a diagram to be drawn of the synthesis, Tu, use, and eventual dissipation of these highly (Fig. 12) Thus the PAs are probably synthesized in the roots of Apocynaceae, Boraginaceae, and Com- positae-Eupatorieae (diesters are also elaborated by Compositae-Senecioneae and Leguminosae: Crotalaria), young plants show highest concen- trations in the roots and even mature plants show a bimodal concentration distribution between flower heads and roots. The compounds also may be concentrated in the leaves when this will give important protection against herbivores; in a few cases, these leaves are attacked by specific in- sects, including larvae of some Danainae, Ith- omiinae, Ctenuchinae, Pericopinae, and Arctii- nae among the Lepidoptera, and a variety of Hemiptera and Coleoptera. Many of these spe- cific herbivores store and use the compounds directly for defense or adult reproduction, where- as others excrete them. As the plant comes into flower the alkaloids are translocated to the flowering tips and then into the nectaries, where they guarantee attrac- lies, the first three pantropical (with scattered species in north temperate areas), the fourth neo- tropical and the last cosmopolitan. The alkaloids are also retained in the seeds to deter predation. In the frequent case of selective attraction of ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 male butterflies to the PA sources, these spread the PAs over their tegument for predator defense and channel them into pheromone synthesis glands (Fig. 3), which sometimes are formed only with PA stimulation (Schneider et al., 1982), and other reproductive tissues, especially spermato- phores. The spermatophores are transferred to females during mating and the PAs are similarly spread out over the tegument and channeled into reproductive tissues, eventually being deposited on the eggs for protection. Newly hatched larvae consume the eggshells but quickly lose their PAs unless they find themselves on leaves that con- tain them. This chemico-ecological flow scheme (Fig. 12) opens ample perspectives for the localization, selection, and cultivation of PA-producing plants. Indicine N-oxide (Fig. 3K) and at least one iso- mer (Fig. 3J) now in advanced clinical testing; some E upato- rium in the right ecologico-evolutionary setting ight become important sources for these c pounds (Table 6). The free bases, however, and especially the diesters, are very hepatotoxic and carcinogenic, representing a serious problem in human and veterinary medicine. Knowledge of the flow of PAs in nature should be useful for the control of both plants and PA content in natural and agricultural systems. It may also help when used as herbal teas or fortifying salads, to avoid permanent liver damage to the unsus- pecting consumers. CONCLUSIONS, SYNTHESIS, AND PERSPECTIVES The integrated ecological, phylogenetic, and biochemical investigation of the Solanaceae/Ith- e interface, with the collaboration of Nephila clavipes, a inre spider that is a major LUN predator of the butterflies, has shown m Although there exists a reasonable and geographically widespread specificity in the usage of 19 genera of Solanaceae by Ithomiinae larvae, especially at the level of host plant secondary chemistry, there is very little evidence for parallel phylogenetic diversification of these interacting groups over evolutionary time; in general, ad- vancing phyletic lines of butterflies use ever more primitive hosts. The New World Ithomiinae seem to have colonized the already generically diver- 1987] sified Solanaceae through sequential preadapta- tion to, and encounter and tolerati ion of pro- hei a. y ssively m foc plants. This has undoubtedly affected the distribution, population structure, habit, habitat, phenology, and exceedingly diversified and vari- able chemistry of these plants, but it should not be called **coevolution." (2) The variable and diversified Solanaceae toxins are not stored by larval Ithomiinae and do not protect newly emerged adults against the spider predator, which rejects field-caught indi- viduals. Adult Ithomiinae depend heavily on de- hydropyrrolizidine alkaloid monoesters for de- fense and reproduction. These are sought out and sequestered from decomposing Boraginaceae and especially from a constant source, stabilized by mutualistic interaction: flowers of Compositae- upatorieae, which place in their nectar a single chemical structure (usually as only one of five different stereoisomers) to attract the pollinators that need these alkaloids. This intimate relation- ship has undoubtedly determined many aspects of morphology, physiology, population struc- ture, distribution, abundance, and the highly i aliens chemistry of these plants and their pollinator Eus additional problems that have aris- en during this research, presently under active investigation with similar methodology, include: (1) The special relationship of the Ithomiinae Tithorea and Aeria to Apocynaceae-Parsonsieae, which sometimes contain PAs that may be stored by the aposematic larvae of these genera (re- search under way with J. R. Trigo). (2) The additional aposematic larvae of Ith- omiinae, which feed mostly on tropane-contain- ing plants and Brunfelsia (though some Solanum are also included), and which may be storing effective predator deterrents from the food plant and, in the case of Methona (which contains al- most no PAs in the E possibly passing them on to the adult butterflie (3) Reasons for the qM nonstorage (and perhaps nonstorability) of most Solanaceae tox- ins by herbivores. (4) Possible participation of further com- pounds, volatile or unstable and derived from PAs or similar precursor, in defense of adult Ith- omiinae. (5) Physiological or behavioral mechanisms for the spreading out of PAs on the tegument of the butterflies and their use in synthesis of var- ious pheromones. BROWN -SOLANACEAE/ITHOMIINAE CHEMISTRY 395 (6) The great diversity and variability of tox- ins in both the Apocynaceae and the Solanaceae used by larval Ithomiinae and the importance of these in relation to evolution, oviposition, larval feeding, survivorship, and reproduction in their usual herbivores and in other potential enemies. LITERATURE CITED ACKERY, P. R. & R. V. VANE-WRIGHT. 1984. Milk- weed Butterflies: Their Cladistics and Biology. London the Amazon ‘valley, Lepidoptera: Heliconidae. Trans. Linn. Soc. London 23: 495-566. BELT, T. 1889. The Naturalist in Nicaragua. John Murray, London. Benson, W. W., K. S. BROWN, JR. & L. E. GILBERT. 1976. Coevolution of plants and herbivores: pas- sion flower butterflies. lion 29: 659—680. BERNAYS, E., J. A. E . ROTHSCHILD. 1977. Pyrrolizidine alkaloids sequestered and stored by apos Lagi grasshopper, Zonocerus variega- 5. J. Zool. London 182: 85-87. incu J. 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Evolutionary Ecology of Trop- cal Mimetic Butterflies (Lepidoptera: Ithomi inae). D is. University of Minnesota HARBORNE, 1979. Flavonoids of . B. & T. the pm rd Pp. 257-268 in J. G. Hawkes, R. er & A. D. Skelding (editors), The Biology and 15. of the Solanaceae. Academic Press, don Lon HUNZIKER, A. T. 1979. South American Solanaceae: a synoptic survey. Pp. 49-85 in J. G. Hawkes, R. N. Lester & A. D. Skelding (editors), The Biology and Taxonomy of the Solanaceae. Academic Press, on IYER, R. P. Brunfelsia hopeana — Pharmaco- logic Screening: Isolation and C ie Raie of 1987] Hopeanine. Ph.D. Thesis. University of the Pa- cific, Stockton, — OTT 1982. Recent develop- . S. & M. RorHscHiLD. 1983. A danaid Müllerian mimic, Euploea core amymone (Cra- mer) lacking cardenolides in the pupal and adult stages. J. Linn. Soc. Biol. 19: 27-33. OCKS, A. R. 1967. Spectrophotometric deter- mination of -onn pyrrolizidine alkaloids. Anal Chem. 39: 443-4 pyrrolizidine alkaloi rovements. Anal. Chem. 40: 1749- MiELKE, O. H. H. & K. S. BRowN, JR. 1979. EU mento ao "Catálogo dos Ithomiidae Americano de Romualdo Ferreira D'Almeida Lepidoptera)” (Nymphalidae, pela uate HANE sidade Fe- deral do eie /CNPq, Curi PLISKE, T. E. 75a. Pici of Lepidoptera to plants de par ne alkaloids. Envi- ronm. Entomol. 4: 455-4 1975b. Pollination of pyrrolizidine alkaloid- containing plants by male Lepidoptera. Environm Entomol. 4: 474-479. . J. CULVENOR. edi The chemical basis of attraction of ithomiine butter- flies to plants containing pyrroli izidine 'alkaloids. J. Chem. Ecol. 2: 255-262. PLowMAN, T. 1979. The genus Brunfelsia: a con- pecan 2 the taxonomy and biogeography. Pp. 475-491 in J. G. Hawkes, R. N. Lester & A. D. Seti diem, The Biology and Taxonomy of he Solanaceae. Academic Press, London kina M. 1973. Secondary plant substances and warning colouration in insects. Pp. 59-83 in . van Emden (editor), Insect/Plant Relation- . Soc. London. Black- ships. Symp. No. 8, R. Ent wells, Oxford. 1979. Mimicry, butterflies and plants. Symb. Bot. Upsal. ad —99, Some peculiar aspects of the dan BA: relationship. Entomol. Exp. & Appl. 24: 437-450. , R. APLIN, J. BAKER & N. MARSH. 1979. Tox- icity Aber in the tobacco horn-worm (Manduca BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY 1968. spetophotometric determination of 397 sexta a (Sphingidae, Lepidoptera). Nature 280: 487-488 w, T. REICHSTEIN, D. A. S. SMITH & J. kasq 1975. Cardenolide storage in ipi s irl (L.) with additional notes on D. plex- ppus (L .). Proc. Roy. Entomol. Soc. London B 31. SCHNEIDER, D. 1977. Plant alkaloids as pheromone precursors in danaid butterflies. Pp. 353-355 in V. Labyrie iy Comportement des Insectes et Milieu Trophiq e. C.N.R.S., Paris —— ——-, M. BoPPR É m" SCHNEIDER, W. R. THOM C.J. iani R. L. Petty & J. MEINWALD. 1975. A pheromone precursor and its uptake in male D butterflies. J. Comp. Physiol. 97: 245- 256. MPSON, mes, J. ZWEIG, S. B. Horsey, T. W. BELL, J. MEINWALD, K. pda ciety W. DIEHL. 1982. Scent oa development in Creatonotos moths: regulation by spins sona alkaloids. Science 215: 1979. The steroid alkaloids of Sola- . 193-202 in . Hawkes, R. N. Lester rà D. Skelding (editors), The Biology and Tax- onomy of the Solanaceae. Academic Press, Lon- aen 5m Solanum prickles and marsupial uri Bot. Gard. 73: 745-754. 8 lacdes de Ithomiinae (Lep., Nymphalida S maré, SP. M.Sc. Thesis, Universidade Estadual de Campinas, Sao Paulo 1986. Interactions between Ithomiinae (Lep- idoptera: Nymphalidae) and Solanaceae. Pp. 364- W. G. D'Arcy (editor), Solanaceae, Biology and Systematics. Medie Univ. e New York. T. M. LEWINSOH mportamento de Nephila clavipes (L) (Arancidac) em relação a lepidópteros: discriminação e reje To de presas. Ciénc. Cultura oo 548 Di scrimination and release of unpalatable [niii by Nephila clavipes, a neotropical orb-weaving spider. Ecol. Entom. 9: 37- 44 YOUNG, A. M. 1972. On the life cycle and natural history of Hymenitis nero (Lepidoptera: Ithomi- inae) in Costa Rica. Psyche 79: 284-294. NEW TAXA OF RUBIACEAE FROM VENEZUELA JULIAN A. STEYERMARK! ABSTRACT r new taxa of Rubiaceae are described from Meridie a SOMME apurense, Faramea Ps sei Psychotria buntingii, and Psychotria subimbrica Coccocypselum apurense Steyerm., sp. nov. TYPE: Venezuela. Edo. Apure: Dtto. Pedro Ca- mejo, Cano El Cabello, 16 airline km NW of Mata de Guanabana, between the Río Meta and Río Cinaruco, 6?19'N, 68?19'W, 75 m, 27 Feb. 1979, Gerrit Davidse & Angel Gonzalez 15825 (holotype, MO; isotype, VEN) erba radicans, caulibus 1.5-2.5 mm diam. glabris; vel subobtusis majoribus 6-7 cm longis 3-4 cm latis, adultis supra costa adpresso-pilosa atque marginibus minute adpresso-ciliolatis, pilis 0.1-0.3 mm longis munitis, ceterum glabris, subtus santas Donius ervis latera- libus utroque latere ca. 7 ULLA subtus im- pressis; inflorescentia ca. 12-flora subcapitata pedun- culata; pedunculo 4.5 mm longo glabro; calycis lobis inaequalibus nen lanceolatis 2-2.5 x 7mm; corollis coeruleis 5 mm longis extus glabris. Creeping herb with rooting glabrous stems 1.5- .5 mm diam. Stipules subulate, to 6 mm long, glabrous. Leaves petiolate, appressed-pilosulous on upper margins, otherwise glabrous; leaf blades ovate, shortly acute or subacute at apex, obtuse or subobtuse at base, the larger ones 6-7 cm long, 3-4 cm wide; adult leaf blades appressed-pilose adaxially on midrib, minutely appressed-pilo- sulous on margins with hairs 0.1—0.3 mm long, elsewhere glabrous, abaxially glabrous; young leaf blades pilose abaxially on midrib and lateral nerves; lateral nerves ascending, 6 or 7 each side; petioles 8-13 mm long. Inflorescence ca. 12- flowered, subcapitate, the heads 8 mm wide, 5 mm high, pedunculate; peduncle 4.5 mm long, glabrous. Flowers sessile; bracts subtending flow- ers linear-ligulate, obtuse, 3 mm long, 0.5 mm wide, glabrous; hypanthium campanulate, 1.5 mm long, 1.1 mm wide, glabrous; calyx lobes unequal, linear-lanceolate, acute, the larger pair 2.5 mm long, 0.7 mm wide, the smaller pair 2 mm long, 0.3-0.5 mm wide, sparsely ciliolate to glabrescent; corolla blue, infundibuliform, 5 mm long, glabrous without, tube 2 mm long, 1.2 mm wide, glabrous within, the lobes ovate-oblong, subacute, 2 mm long, 1.5 mm wide, papillate- puberulent within. Anthers slightly exserted, lin- ear, 2 mm long. Style exserted, 6 mm lon This species resembles the newly described C. croatii Steyerm. but is readily distinguished from that taxon by the longer petioles, very short pe- duncles, less conspicuous, shorter hairs of the ciliate-margined leaf blades, shorter stipules, in- florescences with more flowers, and fewer lateral leaf veins that are less arcuate. Faramea cazaderensis Steyerm., Venezuela. Edo. Tachira: Dtto. Lobatera, La pude 2,000 m, 22 Jul. 1983, H. van der Werff & R. Ortiz 5457 (holotype, MO; iso- type, VEN). Sp. nov. TYPE: rutex 4-metralis, ramis juvenilibus quadrangula- vato-oblongis apice abrupte obtuse acutis basi subcordatis vel omen 6.5-13 cm longis 3.5-7 cm latis prominente ne vatis, nervis lateralibus pedicellis 2.5-4 mm longis; calyce hypanthioque 5.5 mm longo, Rd e ein urceolato l mm longo; calyce tubuloso 4.5 m dentibus qeloideis acuminatis 0.3-1 mm longis, deinde uno latere fiss Shrub 4 m tall, glabrous throughout; branches quadrangular. Stipular sheath shallowly and broadly semilunar, 3 mm long, 2 mm wide, end- ing in an awn 2-3 mm long. Leaves subsessile, subamplexicaul, oblong or ovate-oblong, abrupt- ly obtusely acute at apex, subcordate or rounded at base, 6.5-13 cm long, 3.5-7 cm wide, prom- inently nerved; lateral nerves 9-12 each side, nearly horizontai or F widely spreading at an angle of 5-1 5°, ele imp anastomosing 3-6 mm before the margin, ter- adaxially, ' Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. ANN. Missouni Bor. GARD. 74: 398—400. 1987. STEYERMARK— VENEZUELAN RUBIACEAE 399 1987] tiary veins prominently reticulate, marginal nerves somewhat thickened; petioles 2- m long. Inflorescence terminal, umbellately cy- mose, 2-2.5 cm wide, 1.5-1.7 cm high, 12-15- flowered, pedunculate; peduncle erect, 1.7—4 cm long, | mm wide; primary axes 4-5, ascending, 5-7 mm long, 0.7-0.9 mm wide, each bearing 3 flowers. Flowers on pedicels 2.5-4 mm long, 0.5— 0.7 mm wide; bracts subtending axes and pedi- cels deltoid-ovate, acute, 0.5—0.8 mm long. Calyx and hypanthium 5.5 mm long; hypanthium ur- ceolate, | mm long; calyx tubular, 4.5 mm long, shallowly and unequally dentate, teeth broadly deltoid, acute to acuminate, 0.3-1 mm long, eventually splitting on one side. Corolla purple (in bud), abruptly acute, 7 mm long w 3 Paratypes. VENEZUELA. EDO. TACHIRA: Dtto. Lo- batera, La Cazadera, 7?55'N, 72°18'W, 1,600 m, 24 Jul. 1983, van - Werff & Ortiz 5587 (MO, VEN); o, Quebrada Cazadero, 16 km NW of 1981, Liesner & Guariglia 11668 (MO, VEN). This species is well marked by the subam- plexicaul leaves and relatively few-flowered um- bellate inflorescence. It may be distinguished from F. sessilifolia (Kunth) A. DC. by the umbellately disposed, smaller inflorescence of 4 or 5 axes, each of which is only 3-flowered, the obtusely acute leaves with fewer pairs of lateral nerves, and by the tubular, longer calyx which becomes split on one side. Further, the habitat of the new taxon in the cooler montane forests of the Andes of Táchira is in contrast with that of F. sessilifolia in the lowlands of the Orinoco, Rio Negro, and Madeira river basins. Psychotria buntingii Steyerm., sp. nov. TYPE: Venezuela. Edo. Zulia: Dtto. Perijá, alrede- dores de la Estación Hídrológica Aricuaisá- Pie de Monte, 9?35'N, 72?53'55"W, en zona de bosque siempreverde, 100-250 m, 25 Feb.-3 Mar. 1982, G. S. Bunting, G. Pan- apera & H. Lobo 10876 (holotype, MO; iso- types, JBM, VEN). Suffrutex 0. .3- metralis, ramis minute erg ue sti- 5. 5m n- gis; foliis elliptico-lanceolatis apice subfalcato- acumi- M basi acutate angustatis 4. 5- 8.5 cm longis 0. 9-1.9 ute puberulentibus, nervorum lateralium axillis domatiis barbellatis, ceterum. glabris, subtus punctis latis cymoso-umbellatis gracillimis sub anthesi 0.4 cm alto 1.2 cm lato sub fructu 1.2 cm alto 2 cm lato, 12- 17-floris, axibus primariis tribus filiformibus 3-7-flor- is; pedunculo filiformi 1.6-2 cm longo 0.5 mm lato dense puberulo; floribus pedicellatis, pedicellis 0. 5-0. 8 is alyce 1.2 mm lato; corolla (immatura) 3. 5 mm longa. Slender subshrub; branches minutely tomen- tellose with numerous minute cystoliths. Ter- minal stipule and those on next lower node lan- ceolate, acute, up to 5.5 mm long, 1.5 mm wide, glabrous or glabrate except for the ciliate mar- gins, prominently marked with pale cystoliths near margins, nodes below brown-fimbrillate on the stipular scar. as nom elliptic-lanceolate, subfalcately apex, acutely narrowed to the base, 4.5-8.5 cm long, 0.9- 1.9 cm wide, 4'4—4V5 times longer than broad, glabrous adax- ially, minutely puberulent abaxially on midrib and main lateral nerves, otherwise glabrous, the surfac tely brown-dotted, axils of the nerves ae barbellate with domatia, margins sparsely ciliolate; lateral nerves 7-8 each side, ascending at an angle of 45°, faintly anastomos- ing 1.5-2 mm from margin; petiole very slender, 4-7 mm long, moderately tomentellose. Inflo- rescence axillary or terminal, cymosely umbel- late with 3 main axes, 12-17-flowered, 0.4 cm high, 1.2 cm wide in anthesis, 1.2 cm high, 2 cm wide in fruit, pedunculate, the main axes with two additional shorter central axes 1 mm long; main axes filiform, the lateral 3-7-flowered. Pe- duncle 1.6-2 cm long, 0.5 mm wide, densely puberulent and conspicuously marked with cys- toliths. Flowers pedicellate, pedicels filiform, 0.5- 0.8 mm long, 0.3-0.4 mm thick; bract subtend- ing pedicel subulate, 0.2 mm long. Calyx and hypanthium 1.5 mm long, the hypanthium tur- binate, 0.5 mm long, 0.5 mm wide, tomentellose; calyx tube 0.5 mm long, 1 mm wide, lobes gla brous throughout except for ciliolate margins, longer than the tube. Corolla white (in bud) cy- lindric-infundibuliform, 3.5 mm long, glabrous without, sparsely pilose within the tube, sparsely marked with cystoliths, tube 1 mm long, lobes 2 mm long. Fruit (immature) elliptic-oblong, 3 mm long, 2 mm wide, sparsely puberulent. This species resembles the pubescent P. hor- izontalis var. glaucescens (Kunth) Steyerm., from which it is readily distinguished by the minute calyx and hypanthium, shorter, filiform pedun- cles and pedicels, much smaller inflorescence, and narrowly elliptic-lanceolate leaf blades with barbellate domatia in the abaxial nerve axils. 400 It is a pleasure to associate this new taxon with the name of Dr. George S. Bunting, who has contributed greatly to our knowledge of the flora of the state of Zulia in Venezuela. Psychotria subimbricata Steyerm., sp. nov. TYPE: Venezuela. Edo. Miranda: Parque Nacional Guatopo, laderas pendientes pedregosas a lo largo del Rio Santa Cruz de Rio Grande, lado occidental de la carretera, al sureste de Los Alpes, 10?4'45"N, 66?29'30"W, 400 m, 31 Oct. 1981, Julian A. Steyermark & Bruno Manara 125483 (holotype, VEN). Planta subherbacea l.5-metralis; foliorum laminis subtus glabrescentibus, nervio medio iege minute puberulo; inflorescentiae. peduncu lo | 1-6 c m longo, 1 H 1 hel hv ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 petiole 3-8 x 5 mm. Inflorescence axillary in upper leafy axils and in leafless middle axils, fas- ciculate with 1-3 elongated peduncles 1-6 cm ong, glabrous, trichotomously branched at ma- turity with the central longer axis up to 13 mm long and two lateral shorter axes up to 7 mm long, or the peduncle with two lower axes and an upper trichotomously branched portion; 2 bracts subtending the base of the principal axes naviculate, lanceolate to ovate-lanceolate, acute to acuminate, 7-9 mm long, 4-5 mm broad near base, glabrous. Flowers in small compact groups subspicately arranged, each group subtended by imbricate, ovate, obtuse, glabrous bracts 6.5 x 4 mm; flowers numerous in an inflorescence, 3 or 4 of them ene bs a larger, glabrous, group. Calyx lobes ypanthio glabro. Stem subherbaceous, simple, hollow, leafy be- low, 1.5 m tall. Stipule spreading with revolute margins, broadly ovate-oblong, subacute, 6-10 x 6-8 mm. Leaves broadly obovate or elliptic-ob- ovate, eens and shortly acute to shortly acu- minate or obtusely acute at apex, conspicuously narrowed to the decurrent base, the lamina 21- 25 cm long, 10-15 cm wide; lower midrib mi- croscopically appressed-puberulent, the middle and upper portion with two parallel lines of pub- erulence along the margins of the midrib; abaxial surface glabrous to glabrescent; adaxial surface glabrous; lateral nerves 13-16 each side, arcuate- ascending, arising at angle of 15—20?*, strongly sulcate adaxially, elevated abaxially; decurrent 5, narrowly ligulate, unequal, subobtuse, cilio- ate, 3-3.5 x 0.4-0.6 mm; calyx tube and hy- panthium 2 x 1.5 mm, glabrous; | scale-like squamella, oblong-triangular or lanceolate, acute, in the sinus between 2 calyx lobes; corolla tu- bular, 7 mm long, glabrous except for the barbate orifice. Anthers exserted, oblong, 1.2 mm long. tyle 3.5 mm long; style branches filiform, 1.3 mm long, puberulous. — The new taxon differs from P. humensis Stey- erm. and P. decurrens Steyerm. in the glabrous hypanthium, more branched inflorescence with longer peduncules, glabrous, subimbricately ar- ranged bracts and bracteoles, and minutely pu- berulent lower midrib of a glabrescent abaxial leaf surface. SIX NEW SPECIES OF NEOTROPICAL LAURACEAE! HENK VAN DER WERFF? ABSTRACT Among recent collections of Lauraceae and older collections received on loan from various her ba ria de e grandiflorum van der Werff, placements of these new species are discussed. Lauraceae, comprising about 40 predomi- small, inconspicuous flowers. Genera are often hard to recognize and nearly all are in need of modern treatments. During identification of the large number of unidentified Lauraceae in MO and among the various loans I received, several undescribed species were found. Of those, the following six are here described as new AIOUEA AUBLET Aiouea, recently monographed by Renner (1982), comprises about 20 neotropical species and is best represented in Brazil, Venezuela, and the Guianas. Having seen specimens of most species attrib- uted to Aiouea, it seems quite likely to me that Aiouea, as circumscribed by Renner (1982) and accepted here, is polyphyletic. Most of the South a typical aspect, with large, many-flowered inflo- rescences, flowers with rather long pedicels, leaves fal ipia green. ane itwigs with smooth bark. The ral A p quite different but w to Aiouea because of their 1 nine 2-celled an- thers; they do not fit in the other, better defined, neotropical genera with nine 2-celled anthers. it is quite well possible that the Central American Aiouea species are derived from local Ocotea or Nectandra species which have lost two of their four anther cells. Variation in the number of an- ther cells in Aiouea lundelliana Allen, a species excluded from Aiouea by Renner (1982), has been We and P. westphalii van der Werff, are described. Difficulties with the generic reported earlier (van der Werff, 1984). A study of the wood anatomy could probably answer the question whether these Aiouea species share more characters with the South American Aiouea species or with Central American Ocotea or Nec- tandra species. Aiouea inconspicua van der Werff, sp. nov. TYPE: Mexico. Vera Cruz: 0-2 km S del campa- mento Hnos. Cedillo, rumbo a Río Alegre, por la desviacion al E, Hidalgotitlan, 140 m, 22 Apr. 1974, Brigada Dorantes 2929 (ho- lotype, MO; isotypes, MO, BM, UC). Fig- ure 1 Frutex vel arbor parva. Ramuli tenues, teretes, ju- ores minute tomentelli, vetustiores glabrescentes. Folia alterna, lanceolata (8-11 x 2-3 cm), c acuminata es caudata, subtu ac tomentellae vel glabrae, ines » raeter nervorum ax- illas barbellatas glabrae, sicco olivaceae praeter nervos virides. Inflo rescentiae axillares, minute to mentellae, Flore a a 9 mm longa: 6 exteriora introrsa, 3 interiora extrorsa et basi tia. Ovarium label stigmate crasso sessilique. Bacca globos m diametro, basi cupulae planae, in une ae attenuatae insidens, tota exser- ta. Shrub to 7 m tall. Twigs terete, glabrous; young twigs minutely tomentellous. Terminal buds slender, greyish pubescent. Leaves alternate, 8— 11 cm long, 2-3 cm wide; lamina chartaceous, lanceolate, with numerous small gland dots, the base acute, plane, the apex acute or caudate, with a slender tip 1 cm long, the margin thickened ! I thank the curators of A, BM, BR, GH, U, UC, US, and VEN for the often large loans that I received. Dr. J. Dwyer kindly checked the Latin. John Myers made oe eee The designation CORO is used for the herbarium of the Proyecto Flora Falcón in Coro, Venez 2 Department of Botany, Missouri Botanical Garden, P. (5 “Box 299, St. Louis, Missouri, 63166-0299, U.S.A. ANN. MissouRi Bor. GARD. 74: 401-412. 1987. 1.5mm FIGURE |. Aiouea inconspicua.—A. Habit. — B. Detail of twig.—C. Flower. — D. Fruit.—E. Two outer an- thers. — F. Inner anther with basal glands. — G. Ovary and stigma. 1987] and cartilaginous, glabrous below with the ex- ception of small tufts of hair at the bases of the lowermost lateral veins, puberulous above on the midveins of young leaves, otherwise glabrous; lateral veins 4—6 pairs, slightly raised below and lighter than the leaf tissue, not reaching the mar- gins but forming conspicuous arches ca. 3 mm from margin; tertiary venation reticulate, slightly raised on upper and lower surface. Inflorescences in axils of deciduous (rarely persistent) leaves, minutely tomentellous, to 4 cm long, few-flow- red (4-7 flowers per inflorescence in material seen). Flowers greenish white, funnel-shaped, glabrous, with small terminal openings revealing the anthers, the pedicels 2-3 mm long. Tepals 6, equal or the outer 3 slightly shorter than the inner ones, roundish or slightly wider than long (0.9— 1.2 x 1.2-1.4 mm), concave. Fertile stamens 9, ca. 0.9 mm long; anthers 0.3 mm long, with gland dots, all with 2 anther cells, these filling the entire long, as wide as anthers, pubescent, anista of the inner 3 stamens with two basal glands. Ovary 1 mm long, elliptic, glabrous; style lacking, the stigma sessile, large, 0.4 mm wide. No stami- nodia seen. Fruit round, 1.5 cm diam. Cupule a shallow disc, ca. 8 mm diam., gradually nar- rowed into the pedicel. Additional specimens examined. MEXICO. VERA- CRUZ: Hidalgotitlan, ou Mor" entre Hnos. Cedil- . 1974 (fl.), Brigada Vazquez 3 Feb. 1981 (fl, fr), Wendt, L. y» — González ei dedii) (MO). AT PT. ALTA VERAPAZ: najá, 150- 700 m, 1-2 yen "1942 (sterile), poesi 45 578 (A). Aiouea inconspicua represents the first Aiouea species reported trom € It resembles most closely A. g is (Lundell) Renner, known only from Guatemala. dte. most striking differ- ences between the two species are listed in Table 1. In addition, A. guatemalensis has longer, wider leaves than A. inconspicua, and the leaves dry dark green with contrasting lighter venation in A. inconspicua, a feature lacking in A. guate- malensis. An unusual feature of A. inconspicua is the absence of staminodia. In both earlier generic keys for Lauraceae (Kostermans, 1957; Hutch- inson, 1964) and by Renner (1982) presence of staminodia was considered a characteristic fea- ture of Aiouea, although Renner (1982) men- tioned two exceptions. Because other lauraceous I VAN DER WERFF—NEOTROPICAL LAURACEAE 403 TABLE 1. Selected characters of Aiouea inconspicua and A. guatemalensis. inconspicua guatemalensis Young twigs + puberulous glabrous terminal bud Staminodes lacking present (teste Renner, 1982) Axillary tufts of present absent hairs in lower- most veins Leaf texture chartaceous chartaceous-cori- aceous genera (Ocotea, Nectandra) include species with and without staminodia, I do not see the absence of staminodia in A. p.e as an obstacle for its inclusion in Alou Two of the aia, dide were distributed by the Flora Veracruz project and may be present in several additional herbaria: Brigada Dorantes 2929 was distributed as Nectandra salicifolia HBK and Brigada Vazquez 1368 as Nectandra sanguinea Rolander ex Rottboel. The Steyer- mark collection was distributed as Ocotea effusa (Meissner) Hemsle LICARIA AUBLET The genus Licaria, endemic to the Neotropics, was recently monographed by Kurz (1983), who recognized about 40 species ranging from south- ern Florida to southern Brazil. Among the neo- tropical Lauraceae, Licaria can be recognized readily based on its three 2-celled stamens, dou- ble-rimmed cupule, and alternate (rarely oppo- site, never clustered) leaves. Licaria bracteata van der Werff, sp. nov. TYPE: Guatemala. Alta Verapaz: Sacté, large tree in dense humid forest, 900-1,050 m (fl), Z. Kunkel 7 (holotype, BR). Figure 2. rbor magna. Ramuli obtuse angulati, glabri. Folia iptica, 25-40 x magnop mino odia 9, 6 2 late lanceolata, ad 1 mm longa, 3 interiora lanceolata, ad 0.8 mm longa, staminibus 404 ANNALS OF THE MISSOURI BOTANICAL GARDEN FIGURE 2. Licaria bracteata. Habit and flower seen from aside and above. [Vor. 74 1987] alternantia. Ovarium ED, tubo florale dense pu- bescente. Fructus igno Large tree. Twigs roundly angled, glabrous, with small, light-colored lenticels, the terminal bud drying black, glabrous. Leaves alternate, char- taceous, glabrous on both surfaces (only a small part ofthe lower leaf surface visible on specimens seen), elliptic, the apex not seen, the base acute, 25-40 x 11-15 cm, the upper surface opaque, the midvein and lateral veins (11—15 pairs) sunk- en, the tertiary venation slightly raised, the lower surface opaque, with midvein, lateral veins, and tertiary venation raised. Inflorescences in the ax- ils of bracts, these ultimately deciduous but pres- ent in young inflorescences, up to 5 cm long, very ubescent, with 10-20 flowers per in- of the twigs above the leaves or along short, leaf- less spurs in the axils of persistent leaves. Bracts at bases of inflorescences black, glabrous, 1-1.5 cm long. Flowers 3-4 mm long, 2.5-3 mm wide, sparsely grey-puberulous, urn-shaped, abruptly widened at the base and gradually narrowed to- wards the tip. Tepals 6, erect or somewhat in- curved, unequal, the outer 3 triangulate, ca. 1 mm long, touching each other and largely ob- scuring the inner 3 tepals, these ca. 0.6 mm long, 1.1 mm wide. Fertile stamens 3, ca. 1.4 mm long, each with 2 basal glands, 2-celled, the anther cells small, apical, opening towards the centers of the flowers. Staminodia 9, the outer 6 broadly lan- ceolate, ca. 1 mm long, the inner 3 lanceolate, alternating with the 3 fertile stamens, ca. 0.8 mm long, occasionally with minute anther cells. Ovary glabrous, the floral tube densely pubescent in- side. Fruit unknown. Additional specimen examined. GUATEMALA. ALT VERAPAZ: Sacté, large tree in dense humid forest, 900- 1,050 m (fl), r Kunkel 56 (MO). In Kurz’s treatment (1983) L. bracteata keys to the subgenus Guianensis (correctly: subg. Li- caria, because it includes the type species of the genus) and within this subgenus to a small group of three Central American and West Indian have much smaller leaves, smaller flowers, fewer staminodia, and (with the exception of L. urceo- lata) pubescence on leaves and/or twigs. Licaria urceolata, which has glabrous leaves and twigs, lacks the pubescence of L. bracteata on the in- florescence. Licaria bracteata is an unusua species because of its large, rather persistent VAN DER WERFF—NEOTROPICAL LAURACEAE 405 bracts, large flowers, and, uniquely in subg. Li- caria, the presence of nine staminodia. That Mrs. Kunkel-Westphal, who was not trained as a botanist, collected two interesting new species of Lauraceae in Guatemala, is both a compliment to her qualities as a collector and an indication that many surprises await the col- lector of large tropical trees. PHOEBE NEES A good set of characters to separate the genus Ocotea from Phoebe has unfortunately not yet been found. Ocotea is a particularly large and vaguely defined genus which includes various as- semblages of species. Phoebe likewise is poorly understood. In the New World there are two main centers of Phoebe species: one in northern Cen- tral America and one in central-southern Brazil. In the intervening areas Phoebe is very poorly represented, and it is not clear yet whether the Brazilian and Central American species form a monophyletic group. An added difficulty is that the type of Phoebe is an Asian species and that, according to Kostermans (1961), the Asian Phoebe species are not congeneric with the neo- tropical Phoebe species. Kostermans (1961) transferred all neotropical Phoebe species to Cin- namonum, an Asian genus generally not consid- ered native to the Neotropics. He did not discuss the difficulties in separating Ocotea from Phoebe, nor the heterogeneous assemblage of species clas- sified in neotropical Phoebe. Earlier authors relied on two characters to sep- arate Ocotea from Phoebe. In Ocotea the stam- inodes were said to be inconspicuous, in Phoebe conspicuous; in Ocotea the tepals do not persist in the fruiting stage and a cupule is present, while in Phoebe tepals harden and persist in fruit, but no cupule is present. The staminodial character is relative and difficult to interpret. Because flow- ers of Ocotea and Phoebe are small and the stam- inodes almost never exceed | mm, one may rightly ask what makes minute staminodes con- spicuous in some flowers and inconspicuous in others. Besides, the presence of staminodes is not is also a doubtful character. Of the 18 American species attributed to Phoebe for which I have data, only five have persistent tepals in fruit. However, fruits are unknown for several species. Cupule development in Ocotea is very variable and ranges from a widened pedicel with 406 a minute cupule to well-developed cups. Place- ment of new taxa in Ocotea or Phoebe has been subjective and will remain so until Cinnamo- mum, Ocotea, Phoebe, and their satellite groups are studied critically. I use the following characters as indicators for Phoebe: tepals erect in flower and persistent in fruit, flowers with relatively long pedicels, stam- inodes present, and a tendency toward tripli- veined leaves. Because the new species has four of the five characters, I place it in Phoebe. Using these indicator characters for Phoebe, species such as P. helicterifolia (of which Nec- tandra corzoana Lundell is a synonym) and its allies [P. bourgeauviana (of which Ocotea tene- japensis Lundell is a synonym), P. obtusata, P. valeriana, Nectandra capituliforma, and N. lon- gicuspis] would be excluded from Phoebe. These species should all be placed in the same genus, probably in the already variable Ocotea, or, if future study turns up additional characters, in a new genus. Phoebe glabra van der Werff, sp. nov. TYPE: Mex- ico. Oaxaca: Municipio Matias Romero, tree, 13 m, Wendt et al. 4813 (holotype, MO). Figure 3. Arbor, ad 13 m. Ramuli glabri, teretes. Gemma ter- minalis glabra. Folia glabra, alterna, chartacea, basi acuta vel rotundata vata, axillis vorum basalium domatiis cavitatibus m ng Fr uctus ellipsoideu oo `cupula plana mar- gine integra sore incrass Tree, 8-13 m. Twigs terete, slender, glabrous. Terminal bud glabrous. Leaves chartaceous, al- ternate, the apex acuminate, the base acute or somewhat rounded, elliptic, 12-16 x 4-6 cm; lateral veins 3-5 pairs, the lowest pair most strongly developed and with slitlike domatia in the axils, these visible as small lumps on the upper surface; veins and reticulation promi- nently raised on both surfaces; petioles to 1.5 cm long, glabrous. Inflorescences paniculate, gla- brous, 5-10 cm long, in the axils of persistent leaves or deciduous bracts, often appearing ter- minal. Pedicels glabrous, 2-3 mm long. Flowers glabrous, 2.5-3 mm long, funnel-shaped. Tepals 6, equal, ca. 1 mm long, ovate. Stamens 9, all 4-celled, the filaments narrower than the anthers, 1.3-1.4 mm long, glabrous; outer 6 anthers with ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 cells introrse, inner 3 with cells extrorse. Stam- inodia 3, ca. 0.9 mm long. Ovary globose, gla- brous, ca. 1.8 mm long; style short, ca. m long. Fruit an ellipsoid berry, 2 cm long, cupule platelike, the margin entire, 1.4 cm wide, pe- duncle gradually widened towards the fruit. Additional specimens examined. MEXICO. VE- RACRUZ: Municipio Minatitlan, Wendt et al. 3217 (MO); Municipio Hidalgotitlan, Wendt et al. 3141 (MO). OAXACA: Municipio Matias Romero, Wendt et al. 3064 (MO). Phoebe glabra is closely related to Ocotea eu- venosa Lundell. Both species have twigs, leaves, and flowers glabrous, raised reticulation, and slit- like domatia (these were not mentioned by Lun- dell (1965), but the GH isotype of O. euvenosa shows the domatia clearly). Ocotea euvenosa dif- fers in its larger leaves, laxer reticulation (even a small leaf of O. euvenosa, of comparable size to a regular leaf of Phoebe glabra, has a laxer reticulation), and in its pinnately veined leaves, not subtripliveined as in P. glabra. The type of O. euvenosa is in young fruiting stage, which makes comparison difficult. In the fruiting stage the pedicels become much larger, to 2.5 cm long and I therefore attach little taxonomic value to the longer pedicels reported by Lundell (1965) for O. euvenosa. Three of the four collections of P. glabra come from limestone-derived soils. PLEUROTHYRIUM NEES The genus Pleurothyrium is separated from the other neotropical lauraceous genera by a set of characteristics associated with strong enlarge- the staminal glands. Each of the fila- ments of the inner three anthers carries two glands, which, in other genera, are roundish and rather small. There is no doubt these glands pro- duce nectar. In the genus Pleurothyrium these glands become strongly enlarged and grow out- ward between the outer six stamens. In the re- lated genera Ocotea and Nectandra the outer sta- mens usually form a tight ring, with the anthers cells are not possible because the anthers form such a tight ring. However, in Pleurothyrium the stamens are separated from each other by the rather isolated. There are no spatial constraints against lateral cells on the anthers, and indeed one characteristic of Pleurothyrium is that the lower pair of cells on the six outer anthers is laterally positioned. 1987] FiGure 3. Phoebe glabra. — A. Habit VAN DER WERFF—NEOTROPICAL LAURACEAE 407 — B. Leaf base showing bero —C. Flower. — D. Fruit. — E. Two outer stamens. — F. Basally united inner anther, glands, and two stamino A second characteristic of Pleurothyrium is that in many species the staminal glands become fused into a ring at the margin of the flower or even form a cushionlike glandular mass with only the anthers and the pistil protruding. This glandular ass has been called a disc by some authors (Hutchinson, 1964). Sometimes one can still dis- cern that the g nass is the result of fusion of six glands, and Ears that in Pleurothyrium nine stamens have two glands each are erro- neous, as Rohwer & Kubitzki (1985) pointed out. Additional characteristics for Pleurothyrium are the relatively large, warty cupules, distinctly larger than in most Ocotea and Nectandra species and quite like the cupules in Aniba. The leaves have many lateral veins for their size, a character that, in addition to the large cupules, makes iden- tification of fruiting material possible. Several 408 Pleurothyrium species also have hollow twigs, which are frequently inhabited by ants I noticed in several species that the margins of the tepals are thinner than the centers and that in old flowers the margins of the tepals curl downward and become revolute. I have not seen this in other neotropical Lauraceae. Pleurothyrium costanense van der Werff, sp. nov. TYPE: Venezuela. Edo. Falcon: Sierra de San Luis, above La Chapa, 1,200 m, 10 Aug. 1979, van der Werff 3654 (holotype, U; iso- type, CORO). Figure 4. Arbor, 15-20 m. Ramuli eg dense ferrugineo- pubescentes. Folia alterna, chartacea, elliptica vel late Md apice rotundata vel dece acuta, basi obtusa vel acuta, 20-35 x 8-17 cm. Venatio super immersa, Mr elevata reticulatione conspicua. Inflorescentiae axillares, pubescentia ferruginea, 10-15 cm longae. Te- pala 6, aequalia, patentia, elliptica. Stamina 9, 4-lo- cellata, 3 interiora locellis extrorsis, 6 exteriora locellis inferioribus lateralibus, locellis superioribus introrsis. tylum crassum dense pubescens. Glandulae stami- nium in muro humile connatae, staminibus gynoeci- oque in centro exposito. Cupula magna, verrucata. ee, 15-20 m. Twigs more or less terete, densely bisce, pubescent, glabrescent with age. Terminal bud densely ferruginous pubes- cent. Leaves alternate; laminae chartaceous, el- liptic or broadly elliptic, the apex rounded or shortly acute, the base obtuse or acute, 20-35 x 8-17 cm; with upper surface opaque, the vena- tion sunken, glabrous except for some pubes- cence on midvein and main lateral veins; lower surface with raised midvein, lateral veins, and reticulate venation, the midvein and lateral veins with brown pubescence, otherwise with spread- ing hairs in flowering stage, but glabrous when fruiting; lateral veins 10-15 pairs, the veins arch- nected with the superior vein; petioles 2-3 cm long, ferruginous pubescent. Inflorescences in ax- ils of deciduous or persisting leaves, 10-15 cm long, ferruginous-pubescent, rather laxly flow- ere owers white or buff, ca. 1 cm diam. Te- pals 6, equal, spreading at anthesis, the outer 3 ferruginous-pubescent outside, the inner 3 pu- berulous outside except for a narrow, median, pubescent strip, puberulous inside, elliptic, ca. 4 mm long. Stamens 9, all 4-celled, the inner 3 with extrorse cells, the outer 6 with the lower cells lateral, the upper ones introrse. Staminal glands fused into a low wall surrounding the sta- mens and gynoecium. Style short, stout, densely grey pubescent; stigma flat, about as wide as the ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 style, glabrous; ovary glabrous. Flowering in Au- gust. Cupule large, ca. 2 cm high when pressed, conspicuously warty. Additional € examined. |. VENEZUELA. SUCRE: Peninsula de a, Cerro Espejo (fl.), Stevermark & Rabe 96072 (US. VEN) MONAGAS: Y ucucual, E of Ca- ripe (fr.), Lao 10 (MO). Pleurothyrium costanense is known from three collections in the Cordillera de La Costa and the Sierra de San Luis, in northern Venezuela. Di- agnostic characters are the reticulate venation of the leaves, the broad leaves, and the ferruginous pubescence. The only other Pleurothyrium species in northern Venezuela are P. reflexum Lasser and P. zulianense Lasser (these two names represent the same species, for which I have used the name P. zulianense Lasser). Pleurothyrium zulianense differs in the lack of reticulate venation, in having narrower leaves and smaller flowers, and in the absence of dense ferruginous pubescence. The epithet costanense refers to the Cordillera de La Costa, where this species is found. Pleurothyrium grandiflorum van der Werff, sp. nov. TYPE: Colombia. Choco: Rio Mecana, 5-10 m. Tree, 15 m, along river. Flowers yellow, with pleasant but strange fragrance (as in some euglossine bee-pollinated flow- ers), A. Juncosa 1675 (holotype, MO; iso- types to be distributed). Figure or, 15 m. Ramuli teretes dense breviter pubes- centes. Folia elliptica, 15-25 x 4-7 cm, membranacea, glabra, apice acuta, basi argute acuta vel decurrente. Inflorescentiae ad 10 cm longae, pauciflorae, e foliorum interiora angustiora, d ina 9, 4-locellata, 3 interiora locellis extrorsis, 6 ext iora 2 locellis inferioribus a alibus, 2 = ellis PANE loribus laterali-introrsis. Glandulae stam nium in muro connatae, staminibus styloque in nne expositis. Fructus ignoti Tree, 15 m. Twigs more or less terete, slender, with dense, short, brown pubescence, this vel- vety-shiny on the terminal bud. Leaves elliptic, 15-25 x 4-7 cm; laminae membranaceous, gla- brous on both surfaces, but midvein with some pubescence on lower surface, the apex acute, the base sharply acute or decurrent on the petiole; lateral veins 15-20, departing from the midvein at angles of almost 90°, scarcely or not at all decurrent along the midvein, the venation sunk- en above, slightly raised on lower surface; peti- oles 1-1.5 cm long, canaliculate, with same pu- 1987] T JH F VAN DER WERFF—NEOTROPICAL LAURACEAE 409 FIGURE 4. Pleurothyrium costanense. Habit, flower, and cupule. bescence as twigs. Inflorescences to 10 cm long, inserted in axils of deciduous leaves below the leafy apices of the twigs, with same pubescence as twigs, few-flowered (only 2 or 3 flowers present on each inflorescence, but with scars indicating former presence of more buds); peduncles ca. 5 mm long, with same pubescence as flowers. Flowers very large for the family, 1.5-1.7 cm diam., yellow, fragrant, densely short brown pu- bescent, the tube 2-3 mm long. Tepals 6, spread- ing, 6—7 mm long, subequal; the outer 3 broadly elliptic, the inner 3 narrower, especially near the base, all with dense, short pubescence on the inner surface. Fertile stamens 9, all 4-celled, the inner 3 extrorse, the outer 6 with the lower pair of anther cells lateral, the upper pair lateral-in- trorse; anthers glabrous. Staminal glands form- ing a large wall surrounding the stamens or seem- 410 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FIGURE 5. Pleurothyrium grandiflorum. Habit and flower. ingly covering the stamens in very young flowers. Stigma platelike, ca. 0.3 mm diam., of almost same color and texture as the anthers. Fruit un- known. Pleurothyrium grandiflorum is known only from the type collection made in the Chocó area, Colombia, an area rich in endemics. The species is very distinct in its large flowers and few-flow- ered inflorescences. In contrast to most Laura- ceae, the lateral veins are hardly or not at all decurrent along the midrib. d westphalii van der Werff, sp. nov. TYP uatemala. Alta Verapaz: Sacté. Tree in Psp humid forest, 20 Apr. 1976, J. Kunkel 9 (holotype, BR). Figure 6. Arbor, 20 m. Ramwuli glabri, juvenales hinnulei pu- bescentes. Folia alterna, elliptica, apice acuta, basi acu- a, basi acuta vel decurrenti, membranacea, glabra. In- lorcscentiae ad 8 cm longae, parviflorae, pubescentia a,e foliorum m Mem ij Mig patentia sub anthesim, pu- scent tamina 9, 4-locellata, 3 interiora locellis ae 6 exteriora 2 locellis inferioribus la- £ 1987] VAN DER WERFF—NEOTROPICAL LAURACEAE Ficure 6. Pleurothyrium westphalii. Habit and flower with bud. mah É eek superioribus introrsis. Glandulae sta in muro connatae staminibus styloque in odi eR En ignoti Tree, to 20 m. Older twigs glabrous, young twigs with brown, dense, short pubescence. Leaves elliptic, the apex acute, the base sharply acute or somewhat decurrent along petiole, membranaceous, 15-20 x 4-7 cm, glabrous or nearly so above, with some appressed pubes- cence, especially near base, but glabrescent with age below; lateral veins 5-8, fading out near the margin, not looping upward and not connected with other lateral veins; upper leaf surface dark, dull, venation sunken; midvein, secondary, and tertiary venation slightly elevated; petioles to 1 cm long, pubescent as the young twigs. Inflores- cences ca. 8 cm long, densely grey-brown pu- bescent, few-flowered (fewer than 10 flowers per 412 inflorescence), in the axils of deciduous leaves or bracts on the young twigs but attached below the leaves, narrowly pyramidate. Flowers with 6 equal tepals, these spreading or slightly reflexed at an- thesis, densely short tomentose outside, slightly less so inside, the tepals ovate, ca. 4 mm long. Fertile stamens 9, all 4-celled, the inner 3 ex- trorse, the outer 6 with the lower two e lateral, the upper two cells introrse. Anthers glabrous, staminal glands not seg nce! visible, but fused into circular mass ca. 3 mm in diam. with the anthers and pistil iS in e middle. Pistil platelike, gray. Fruit unknown. Additional specimens examined. GUATEMALA. ALT VERAPAZ: Sacté, Kunkel 17 (fl.) (MO), 298 (sterile) (BR). Pleurothyrium westphalii is the northernmost species of its genus and is known only from one locality in Guatemala. Two other Pleurothyrium species, as yet undescribed and under study by W. Burger, are known from Central America. The species represented by Allen 5885 (GH, US) from Costa Rica differs in having leaves with acuminate apices, persistent bracts in the inflo- rescence, leaf bases acute but not decurrent, and the inner surfaces of the tepals papillose-puber- ulous, not pubescent. The leaves of Allen 5885 also have more lateral veins which arch upward and become connected, forming almost a mar- ginal vein, as is often seen in Pleurothyrium ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 species. The second species, represented by Bunt- ing & Licht 872 (NY) from the Rio San Juan area, Nicaragua, differs in having a many-flow- ered inflorescence (20-30 flowers per inflores- cence), darker pubescence, and the staminal glands not fused into a large wall as in P. west- phalii This species is named after the collector, Irene Kunkel, née Westphal, who collected two new and very interesting species of Lauraceae in Gua- temala. LITERATURE CITED HUTCHINSON, J. 1964. The Genera of Flowering Plants. Clarendon Press, Oxford. KOSTERMANS, A.C. J. H. 1957. Lauraceae. Commun. Forest Res. Inst. 57: 1—64. . The New World species of Cinna- momum Trew. (Lauraceae). Reinwardtia 6: 17- 24. Kunz, H. 1983. due dui cu inr eodd Gat- tungen neotropischer Lauraceen und Revision der Gattung d Ph.D. D na a Peu of Hambur LUNDELL, C. L. 1965. Additions to the Lauraceae of Guatemala. Phytologia 12: 243- RENNER, S. 1982. Aiouea. Fl. Neotrop. “Monogr. 31: 85-116. ROHWER, J. & K. Kusirzki. 1985. Entwicklungsli- nien im Ocotea-Komplex (Lauraceae). Bot. Jahrb. Syst. 107: 129- WERFF, H. VAN DER. E Notes on neotropical Lau- raceae. Ann. Missouri Bot. Gard. 71: 1180-1183. A NEW SPECIES OF OCOTEA (LAURACEAE) FROM SOUTHEASTERN MEXICO! ToM WENDT? AND HENK VAN DER WERFF? ABSTRACT The new species Ocotea een is a common riparian tree of the Uxpanapa area of extreme southern Veracruz, Mexico. It appea s to be most closely related to O. eucuneata of Gua temala and Belize. It is yet another species M endemic to the rain forests of the Uxpanapa area. Ocotea uxpanapana Wendt & van der Werff, sp. nov. TYPE: Mexico. Veracruz: Municipio Minatitlán, Zona de Uxpanapa, terracería La Laguna-Uxpanapa, orilla Oeste del Río Oaxaca, un poco al Oeste de Uxpanapa, 17?13'N, 94?13'15"O, 160 m, 14 Feb. 1981 (fü, Wendt, Villalobos & Olmstead 2869 (holotype, MEXU; isotypes, BM, CAS, CHAPA, ENCB, MO, NY, and others). Arbor ad 30 m. Ramuli hornotini sericei cito gla- longiores quam latiores, apice acumin vel cuneata, supra glabratae, ate sibi: oes axillis barbatis. 3 mm longa; stamina externa stipitata. Fructu ellipticus, ad 2.2 cm longus; cupula valde 6-lobata. Tree, to 30 m, to 1 m d.b.h. or often with several trunks from near the base; buttresses small to medium-sized or lacking; bark medium gray- brown to dark chocolate-brown, finely to prom- inently warty, soft; slash of bark aromatic, light brown or yellowish-brown, oxidizing in ca. 1 mi- nute to darker orange- RIO a, sapro pale cream-brown. Shoot portions of stems and new leaves) and axillary buds densely and finely sericeous; twigs soon glabrate, green (drying dark or black). Leaves alternate; blade obovate to elliptic, usually narrowly so, 8-25 cm long, 2.2-8 cm wide, some leaves smaller, mostly 3-4.3 times as long as wide, firmly membrana- ceous, slightly and irregularly conduplicate, me- dium-dark green, finely and densely glandular- punctate, distally acute or rounded to a short- or long-acuminate (to 2.5 cm) apex with a minutely rounded tip, the base acute to usually cuneate; venation slightly raised above, prominently so below, laterals 7-14, diverging at 40—65?, the lowest 1—2 pairs usually more strongly ascending (20-35°), basically eucamptodromous but often more or less brochidodromous distally, the ter- tiary venation more or less transverse between laterals, the fine venation reticulate; adaxial sur- face glabrous, abaxially finely strigose on surface and sides of midvein, at length sometimes sub- glabrate, lowest 1—several pairs of lateral veins prominently barbate in axils; petioles 1-2.3 cm long, canaliculate adaxially, at first finely stri- gose, soon glabrate. Inflorescence complex formed by a group of paniculate cymes, each of these arising from the axil ofa quickly deciduous brac- teate leaf (or rarely a foliage leaf), the apex of each complex a small, temporarily dormant veg- etative shoot, each inflorescence arising either in the axil of a mature leaf or along the apical por- tion of the main stem, several inflorescences oc- curring together to form a large pseudoterminal inflorescence complex, the vegetative apices usu- ally elongating after flowering and producing ma- ture leaves, the fruiting panicles thus more ob- viously lateral; individual inflorescences 5-20 cm long, 2-13 cm broad, cylindrical to pyramidal, with a more or less straight central axis bearing strongly diverging laterals, the latter sometimes similarly rebranched, bearing dichasial groups of flowers, the axes flattened, green to pinkish, light- ly canescent to glabrate in basal parts, to densely canescent distally; flower bracts lance-ovate to deltate, 2-2.5 mm long, densely and finely ca- nescent, very quickly deciduous; pedicels stout, 1.5-3.5 mm long, densely and finely canescent, green. Flowers (4—)4.5-6 mm diam. at anthesis, ! We thank Hans Georg Richter (Hamburg) for bark and wood analysis, and Eduardo Merino, of the Centro de Estudios del Desarrollo Rural of the Colegio de Postgraduados, for the drawing. Fieldw author was jy que by the Centro de Botánica of the Colegio de Postgraduados and th .; thanks are due to all who have participated in that fieldwork, a Ache "Villalobos ork 2 b senior n del Pa- ete ? Centro de Botánica, Colegio de Postgraduados, 56230 Chapingo, Edo. de México, Mexico. A. ? Missouri Botanical Garden, P.O. Box ANN. Missounmi Bor. GARD. 74: 413-415. 1987. 299, St. Louis, Missouri 63166-0299, U.S. 414 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURE |. Ocotea uxpanapana. — A. Branchlet € iota idee inset, lower surface of leaf. — B. Portion of inflorescence, pre- -anthesis. — C. Flower. — D. Parts e flower: above, outer stamen, adaxial view; center, inner stamen, abaxial view; . staminode (frequent absent) right, gynoecium.—E. Fruits with unique lobed calyx. Bar alee A, E = 1 cm; B, C, D = m. Vouchers: A-D, Wendt et al. 2869 (staminode in D, 2865); E, Wendt et al. 2777. 1987] tepals at anthesis ascending 25-45? or more, broadly ovate, 1.923 mm long, 1.5-2.4 mm wide, thick-textured, cream to creamy-green, abaxially canescent with short, thick, gray hairs mostly 0.05-0.15 mm long, adaxially less densely so, the apex broadly acute to acute; hypanthium 0.6-1 mm long. Outer 6 stamens 1.1-1.6 mm long; filaments 0.5—0.9 mm long, pubescent; anthers quadrate-ovate, with truncate to emarginate api- ces, and with the upper pair of thecae more or less above the lower pair; inner 3 stamens 1.5- 2 mm long; filaments 0.7-1.2 mm long, pubes- cent, the basal pair of glands short-stipitate to subsessile; staminodes absent or, when present, less than 1 mm long, linear, pubescent. Ovary 1-1.5 mm long, glabrous; style (0.5—)1-1.4 mm long, shorter than to slightly longer than ovary. Fruit (nearly mature) broadly ellipsoid, to 2.2 cm long, to 1.9 cm thick, green, seated in a very shallow strongly 6-lobed red, fleshy cupule, the lobes 5-7 mm long, 3-5 mm high, 2-3 mm thick; flesh of fruit ca. 1 mm thick, pale green. Testa dark brown; flesh of cotyledons pink. Paratypes. MEXICO. VERACRUZ: Zona de Uxpana- m Mpio. Hidalgotitlán: Campamento La Laguna, along o Las Cuevas, 6 Mar. 1984 (fl.), Taylor 387 (F, MO, n. Campamento Hnos. Cedillo, 18 Jun. 1974 (fr), P. E. Valdivia Q. 808 (MO, XAL); Propiamente en la Escuadra, 7 Aug. 1974 (fr), P. E. Valdivia Q. 1312 (MO, XAL); A 600 m del Campamento Hnos. Cedillo hacia Paso Moral, 8 Aug. 1974 (fr), P. E. Valdivia Q. Brigada. Vázquez 1029 (MEXU, MO); Brecha Hnos. Cedillo-Agustin Melgar, 5 Mar. 1974 (fl), Brigada Vázquez 535 (MO, XAL); La Escuadra-Hnos. Cedillo, por el Río Soloxuchil, 3 May 1974 (fr), Husad Váz- A 729 (MO, XAL); Campamento La Laguna, Rio Las Cuevas, 17?16'30"N, 94*30'W, 130 m, 17 Mar 1982 (fl), Wendt, Villalobos & Navarrete 3725 (CHA- PA, MEXU, MO, and exu Uxpanapa, desde el pobla 17°12'N, 94*10'W, 130 m, y pe 1. 1980 MEUM 2i Wendt, Villalobos & Lara 2564 (CHAPA, and oth a W de La Garganta, 17?13'30"N, 94?15' 30"W. 18 ES Feb. 1981 (fl), Wendt, Villalobos & Olm- sy rp 2865 (CHAPA, MEXU, MO, and others); type locality and date, Wendt, Villalobos & Olmstead 2869A (CHAPA, MEXU, MO, and others). Common names. Aguacatillo, laurel. Ocotea uxpanapana is a common riverside tree of the Uxpanapa area of southern Veracruz, a region of lowland rain forests with high ende- mism (see Wendt et al., 1985, and references there cited); it is interesting that this riparian tree WENDT & VAN DER WERFF—OCOTEA 415 has never been collected farther downstream in the Río Coatzacoalcos basin. In the Uxpanapa area, the new species and Ficus insipida Willd. are the most common large riparian trees. The root system of O. uxpanapana is apparently deep and well adapted to resist flooding; during the major flooding of the Río Oaxaca in October of 1980, all trees (including Ficus insipida) except for many individuals of O. uxpanapana were toppled and carried I Ocotea uxpanapana also occurs scattered sites on deep limestone-derived S It flowers mostly in Feb- ruary and March, fruiting in September and Oc- tober. Ocotea is a large, mostly neotropical genus with a few hundred species. The last comprehensive treatment was by Mez (1889), and since that time many additional species have been described. The Central American species were treated by Allen (1945), but the genus has remained difficult and almost inaccessible to the nonspecialist. Under these circumstances, we were initially hes- itant to describe yet another Ocotea species, but the striking, deeply six-lobed cupule is such a unique feature in the genus, even in the family, that we decided to publish a new species. It ap- pears to be most closely related to O. eucuneata Lundell (type from Belize). In addition to its less strongly lobed cupule (fruiting material collected in the mountains of Guatemala, identified by C. Allen), O. eucuneata differs in its inflorescences borne in the axils of mature leaves (not forming a pseudoterminal inflorescence as in O. uxpan- apana), its smaller flowers, and in having the reticulation on the upper leaf surface immersed (not raised as in O. uxpanapana). Specimens collected by the Brigada Vázquez and P. E. Valdivia Q. are distributed under sev- eral names aaa s loeseneri Mez, N. salici- folia Kunth, Persea americana Miller, and Phoebe gentlei mand Standley & Steyerm.) and are likely filed as such in several additional herbaria. Wood and bark samples of the new species (Wendt et al. 3725) were examined by H Richter. He noted (pers. comm.) that the SM and bark anatomy are typical of Ocotea in gen- eral and are nondescript within the genus. LITERATURE CITED ALLEN, C. K. 1945. Studies in Lauraceae, VI. J. Ar- nold Arbor. 26: 280—434. m i 1889. Sende Se; Americanae. Jahrb. Kónigl. t. Gart. Berlin 5: 1-556. E T. S. A. RI & G. T. PRANCE. 1985. Esch- weilera mexicana (Lecythidaceae): a new family for the flora of Mexico. Brittonia 37: 347-351. FOUR NEW SPECIES OF AXONOPUS (POACEAE: PANICEAE) FROM TROPICAL AMERICA! GERRIT DAVIDSE? ABSTRACT The following species of 4 xonopus sect. jai ipsis are described: A. rupestris, from Goiás, Brazil, A. casiquiarensis from Guainía, Colombia, and Amaz Venezuela, and A. jeanyae from Coclé, Panama. Axonopus is a Aper i pond ican grass genus with a few spec di to the American subtropics i pon Old World tropics. Black (1963) monographed the genus and recognized 109 species. With continued explo- ration it has become apparent that new species remain to be described. On the other hand, it is also probable that some of the species recognized by Black must eventually be synonymized as in- termediate populations become known. Axonopus belongs to the tribe Paniceae and is distinguished by the absence of a lower glume, lower palea, and lower flower, and by its solitary, dorsally compressed spikelets borne inversely (i.e., with the back of the upper glume facing away from the rachis) in two rows on one or usually several to many racemes. This paper reports on recent fieldwork in trop- ical America that has brought to light four un- described species of Axonopus sect. Axonopus. These are published now so that the names will be available for two forthcoming floras, Flora Mesoamericana and Flora of the Venezuelan Guayana. Axonopus rupestris Davidse, sp. nov. TYPE: Bra- zil. Goiás: Mun. Presidente Kennedy, road from highway BR-153 to Itaporá, 12 km W of village of Presidente Kennedy, Fazenda Primavera along Ribeirão Feinho, ca. 3?25'S, 40°37'W, 400—500 m, | Feb. 1980, T. Plow- man, G. Davidse, N. A. Rosa, C. S. Rosário & M. R. dos Santos 8216 (holotype, MG; isotypes, F, MO, NY). Figure 1. xonopus rupestris Davidse, sp. nov. A. triglochi- noides (Mez) Dedecca affinis sed spiculis i ici MARN and NSF in dept It dos Santos, for the stations, Per O. Huber, manuscri nas, Venezuela, A. chimantensis from Bolívar, racemis paucispiculis, glumis minus valde nervis, et anthoeciis glabris differt. Caespitose perennial; culms 10-35 cm tall, the internodes 1-2 mm wide, hollow, glabrous, the nodes l-2, glabrous, the uppermost geniculate. Leaves erect; sheaths to 7.5 cm long, strongly distichous, conduplicate and keeled, apically winged, mostly glabrous, sparsely hirsute near the juncture with the blade; flag leaf sheath to 10 cm long; collar not differentiated, the sheath with a slight marginal constriction apically but oth- erwise merging imperceptibly with the blade; lig- ule a ciliate membrane 0.5-0.6 mm long, the membrane 0.1—0.2 mm long, the cilia 0.4—0.5 mm long; blades erect, mostly 5-10 cm long, folded, mostly 1-2 mm wide as folded, sparsely papillose-hirsute abaxially and adaxially along the midrib to glabrescent, the apex acute. Inflo- rescences 4.5-7 cm long; peduncles 1 or 2, the uppermost to 19 cm long; racemes usually 2, conjugate, sometimes 3 with the lowermost borne 5-10 mm below the upper pair, 4.5-6.5 cm long, widely spreading at maturity, the axil puberulent; rachis ca. 0.6 mm wide, glabrous, 14—16 spikelets per 25 mm; pedicels 0—0.1 mm long, glabrous; spikelets 2.9-3.5 mm long, 0.8-1 mm wide, el- liptic-oblong, obtuse, the upper glume 0.2-0.3 mm longer than the lower lemma, 5-nerved, the lateral nerves marginal, the midnerve weakly de- veloped, prominently pubescent marginally and between the nerves with hairs 0.5-1 mm long, the hairs longest apically, the upper '4 glabrous; idne ally and between the nerves; anthoecium 0.5-0.9 mm shorter than the upper glume, stramineous, ' Fieldwork was conducted with the support of the Projecto Flora Amazónica in Brazil; NSF, CONICIT, R, NGS, Fundación para el Desarrollo de las Ciencias Físicas, Matemáticas y Naturales in Venezuela; . Huber, S. S. Tillett, B. Nelson, T. Plowman, N. A. Rosa, C. S. Rosário, M. R milton, J. s. Miller, and C. Brewer-Carías for help with my personal fieldwork, J. K. Myers J. D. Dwyer, and E. Kellogg for useful suggestions for improving Nie 2 pepa Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. ANN. Missounmi Bor. GARD. 74: 416—423. 1987. 1987] FiGUREl. Isotype of Axonopus rupestris. — A. Habit DAVIDSE—AXONOPUS 417 2.0cm —B. Abax upper glume. — C. Adaxial view of spikelet showing the lower lemma. [After POM et al. 8216 (MO). 1 nearly glabrous at the tip, obtuse; lodicules 2, fleshy, truncate, glabrous; stamens 3, the anthers 1-1.2 mm long, dark yellow; styles 2, separate; stigmas plumose, white; caryopsis not seen. This species is known only from the type col- lection, which was encountered in cracks and shallow, gravelly soil pockets of a black, granitic rock outcrop (Fig. 2). The epithet alludes to this rocky habitat. This species, by virtue of its glabrous rachis, stramineous anthoecia, perennial habit, few ra- cemes, and weakly developed midnerve of the glumes, belongs to sect. Axonopus ser. Axonopu. of Black (1963). However, it does not seem closely related to any of the species grouped by Black in this series. Rather, A. rupestris seems most closely related to A. triglochinoides, a species of the Rio Atabapo and upper Rio Negro drainages, which [97] Black placed in sect. Axonopus ser. Barbigeri subser. Barbigeri because the midnerve of the glume is developed, although not as strongly as in typical members of this series. Axonopus rupestris differs from A. triglochi- noides in the smaller, much more conspicuously pubescent spikelets (2.9-3.5 vs. 3.9-4.8 mm), greater spikelet density on the inflorescence branches (14-16 vs. 6-7 spikelets per 25 mm), less prominently nerved glumes, and lack of a prominent tuft of hairs at the tip of the anthoe- cium. The two species are similar in the general facies of their inflorescences and leaves, espe- cially in the prominently distichous, densely tuft- ed sheaths. Both species are relatively uncom- mon, although local populations may consist of hundreds of individuals. Axonopus triglochi- noides also grows in cracks of granite outcrops and shallow soil pockets. E 2. Axonopus rupestris habit and habitat, type loal. Goiás, Brazil. Axonopus chimantensis Davidse, sp. nov. TYPE: Venezuela. Bolívar: Distr. Piar, Macizo del Chimantá, sector centro-noreste del Chi- mantá-tepui, cabeceras orientales del Caño Chimanta, vegetación litófita y riberena al- rededor del comienzo E del Cañón recto del Río Chimantá superior, ca. 5?18'N, 62°09'W, ca. 2,000 m, 26-29 Jan. 1983, O. Huber & J. A. Steyermark 6931 (holotype, MO; iso- types, K, NY, US, VEN). Figure 3. 4xonopus chimantensis Davidse, sp. nov. A. villosus Swallen affinis sed foliis culmis ligulis et aie mi- noribus differt Caespitose perennial; culms 25—60 cm tall, the internodes 1-2 mm wide, hollow, somewhat flat- tened, glabrous or apically with two lines of pu- bescence along the margins, unbranched, the nodes 1, rarely 2, densely pilose; sheaths 3-9 cm long, very strongly distichous, conduplicate and keeled, apically winged, glabrous except for the upper '4 of the margin and keel where ciliate, the margin membranous; flag leaf sheath 10-21 cm long; collar prominent, densely pilose; ligule a ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 ciliate membrane 0.3-0.5 mm long, the mem- brane 0.1—0.2 mm long, the cilia 0.2-0.4 mm i blades folded, 6-18 cm long, 3.5-5 mm e as folded, divergent, usually prominently Hem on the margins, the keel sparsely ciliate to glabrescent, the tip obtuse, scabrous, adaxially very sparsely pilose to glabrous, abaxially gla- ous. Inflorescence 3-10.5 cm long; axis 5-18 mm long; peduncles 1—2, usually well-exserted; racemes 2-4, 2.5-10 cm long, spreading at acute angles, the axils pubescent, the rachis 0.5-0.8 mm wide, scaberulous or pubescent marginally, 9-13 spikelets per 25 mm; pedicels 0.2-1.1 mm long. Spikelets 3-3.4 mm long, 0.9-1.1 mm wide, oblong-elliptic, obtuse; upper glume as long as the lower lemma or to 0.4 mm longer, 5-nerved, the midnerve developed, inconspicuously and sparsely short-pubescent between the nerves; ower lemma similar to the upper glume; an- thoecium 0.3-0.7 mm shorter than the upper glume, stramineous, apically with a distinct tuft of hairs, obtuse; lodicules 2, fleshy, minutely erose, glabrous; stamens 3, the anthers 1.2-1.5 mm long, purple; styles 2, separate; stigmas plu- mose, purple; caryopsis not seen. Paratypes. "VENEZUELA. BOLIVAR: Distr. Piar, Ma- cizo del Chimanta, sector centro- noreste del Chiman- ta-tepui, cabeceras orientales de ] mantá, altiplanicie en la base me- riodional de los farrallones superiores del Apacará- tepui, sector norte del Macizo, ca. 5?20'N, 62°12’W, ca. 2,200 m, 30 Jan.-1 Feb. 1983, Steyermark et al. 128405 (K, MO, VEN), Huber & Steyermark 7043 (MO, VEN), 7032 (MO, VEN); Chimantá Massif, cen- tral section, island in Río Tirica above Middle Falls e ma central section, swampy savanna along tributary valley > branch of headwaters of Rio Tirica, 2,120 m, 13 Feb. 1955. Steyermark & Wurdack 836 (US, VEN). Axonopus chimantensis belongs to Black's (1963) sect. Axonopus ser. Barbigeri subser. An- cipites because of the perennial habit, non-pa- pillose-pilose rachis and spikelets, stramineous anthoecia, developed midnerve of the glume, and broad, folded blades with obtuse apices. This group of species is best developed in northern South America, especially in the Guayana Re- gion. Axonopus chimantensis seems most closely re- lated to A. villosus and A. steyermarkii Swallen. Both of these are much larger than A. chiman- tensis. In addition A. villosus has slightly larger 1987] DAVIDSE—AXONOPUS 2.0cm FIGURE 3. Holotype of Axonopus chimantensis. — A. Habit. — C. Adaxial view of spikelet showing the lower lemma. [After Huber & Steyermark 6931 (MO glume. and broader spikelets with more prominent nerves, larger ligules, relatively longer anthoecia, and a tendency toward densely pubescent blades and sheaths. Axonopus steyermarkii differs fur- ther in its nearly glabrous foliage, blades notice- ably narrowed at their bases, and usually longer pedicels. Many of the cited specimens of 4. chimanten- Sis have smut-infected spikelets. The rachis of the racemes in A. chimantensis varies from pubescent to scabrous, a character- istic also attributed to A. villosus by Black (1963). Black cited Steyermark & Wurdack 463 and 863 —B. Abaxial view of spikelet showing the upper as A. villosus, but I refer them to A. chimantensis. As far as presently known, A. chimantensis is endemic to the large Chimanta-tepui imn in Bolivar and derives its epithet from t Axonopus steyermarkii is nd to Cake Marahuaca and nearby Cerro Duida in Ama- zonas, Venezuela. Axonopus villosus occurs in the same area as A. steyermarkii and may reach as far north as the Rio Manapiare. S —— eg sp. NOV. TYPE. mazonas: Dpt. Atabapo, Cu- Mine de ptem. ç S rds ofthe middle part 420 FIGURE 4. Holot type of Axonopus casiquiarensis. — A. Habit glume. — C. Adaxial view of spikelets showing the lower lemma. [After Davidse et al. 16907A (MO).] of Caño Caname, 67?22'W, 3?40'N, ca. 100 m, 30 Apr.-1 May 1979, G. Davidse, O. Huber & S. S. Tillett 16907A (holotype, MO, mounted as 2 sheets; isotypes, K, US, VEN). Figure 4. Axonopus casiquiarensis Davidse, sp. mans (Trin.) Kuhlm. et A. camargoanus Hu affini sed stolonibus, basibus culmi gracilioribus et foliis la- tioribus differt Stoloniferous perennial; stolons infrequent, when present well-developed, to 70 cm long, leafy, frequently rebranching and producing tufts of leaves at the nodes, the internodes to 20 cm long; culms 24—90 cm tall, the internodes ca. 1 mm wide, hollow, slightly flattened, glabrous, the nodes 1-2, usually appressed-pubescent, rarely ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 —B. Abaxial view of spikelet showing the upper glabrous. Sheaths mostly 3-12 cm long, rounded on the back, not strongly distichous, not winged, usually glabrous, sometimes pilose at the base, auricular hairs 1-3 mm long usually present; flag leaf sheath to 29 cm long; collar not differen- tiated; ligule a ciliolate membrane 0.1-0.2 mm long, the membrane nearly obsolete to 0.1 mm long, the cilia ca. 0.1 mm long; blades mostly 7— 60 cm long, 1-3 mm wide, flat to convolute, commonly ciliate in the lower !^, otherwise gla- brous, the apex acute. Inflorescence 4-1 5 cm long; axis 2-30 mm long; peduncles 1 or 2, usually well-exserted; racemes 2-3(-5), 4-14.5 cm long, nearly appressed to spreading at acute angles, the axils puberulent; rachis 0.3-0.5 mm wide, sca- berulous, 10-15 spikelets per 25 mm; pedicels 0.1-0.7 mm long. Spikelets (2.5—)2.7—3.7(-4.1) 1987] mm long, 0.8-1.3 mm wide, oblong-elliptic to lanceolate-elliptic, obtuse; lower glume as long as to 0.3 mm shorter than the lower lemma, 5- 7-nerved, the midnerve developed, slightly sul- cate between the nerves, usually sparsely pubes- cent between the nerves, sometimes nearly gla- brous or strongly pubescent; lower lemma similar to the upper glume; anthoecium 0.1-0.7 mm shorter than the upper glume, stramineous, api- cally with a distinct tuft of hairs, obtuse; lodicules 2, fleshy, inconspicuously 3-lobed, glabrous, vas- culated; stamens 3, the anthers 1.5-2.2 mm long, purple; styles 2, separate; stigmas plumose, pur- ple; caryopsis ca. 1.5 mm long and 0.7 mm wide, elliptic-obovate, the hilum elliptic-punctate, the embryo ca. '2 the length of the caryopsis. Paratypes. COLOMBIA. GUIANÍA: Rio Atabapo, ca. 7 km S of San Fernando de Atabapo, 67°43’W, 3*55'N, 28 Apr. 1979, Davidse 16825 (MO, VEN). VENEZUELA. MAZONAS: DPT. ATABAPO: Cano Caname, 67?22'W, 3*40'N, 29 Apr.-4 May 1979, Huber et al. 3643 (MO, VEN), 30 Apr. 1979, Huber et al. 3651 (MO, VEN), 2 May 1979, Davidse et z 17049 (COL, K, MO, TFAV, VEN), 67°13'W, 3°40'N, 3 May 1979, Davidse et al. 17126 (MO, VEN), 5 W, 3°41'N, 2 May 1979, Davidse et al. 17084 p VEN); Cano Yagua, 66?34'W, 3*36'N, Davidse et al. 17421 (MO, VEN), 66*41'W, 3°29' N. 7 May 1979, Davidse et al. 17314 (MO. VEN): 5 km al S de ^a Laguna Yagua, 66?38'W, 3?43'N, 22 Jul. 1980, Huber & Tillett 5476 (MO, VEN); Rio Ata- bapo, 20 km S of San Fernando de Atabapo, 67?39'W, 3°50'N, 29 Apr. 1979, Davidse et al. 16856 (INPA, MG, MO, VEN); Cano Cotuá, cerca Cerro Yapacana, 66°50'O, 3?40'N, 14-28 Feb. 1978, Huber 1605, 1679 (MO, VEN), 66°52'W, 3?*38'N, 6 May 1979, Davidse et al. 17262 (MO, NY, VEN), 17229, 17239 (MO, DPT. CASIQUIARE: medio Río Temi, 67?29'O, 2°57'N, 24 Feb. 1979, Huber 3410 (K, MO, TFAV, ) DPT. RIO NEGRO: lower Río Baria, 66°32'— pag har 1°27'-1°10'N, 22-23 Jul. 1984, pu ie F, XU, MO, NY, TFAV, VEN); Rio 43 Ed Eus W. 1?53'21?27'N, 23-25 Jul. 1984, D vidse 27757 (MO, NY, VEN), 66°32'W, 1?38'N, 9 Feb. 1981, Huber & Medina 5888 (MO, VEN), 66?33'W, 1°35'N, 8 Feb. 1981, Huber & Medina 5858 (VEN); io Siapa, 66?25'W, 2?05'N, 7 Feb. 1981, Huber & Medina 5807 (VEN). By virtue of its few, scaberulous racemes, stra- mineous anthoecia, perennial habit, well-devel- oped midnerve of the glume, and narrow leaves A. casiquiarensis belongs to sect. Axonopus ser. Barbigeri subser. Barbigeri of Black (1963). It is related to A. comans and A. camargoanus of central to southern Brazil both of which lack stolons, and have consistently convolute blades and thickened, usually deeply buried culm bases. Both 4. comans and A. camargoanus inhabit wet savannas, whereas A. casiquiarensis grows in DAVIDSE—AXONOPUS 421 white-sand savannas and river banks, including sandy pockets on granite rock outcrops. This species is distributed in Colombia and Venezuela in the lowland area (elev. 80-125 m) centering around the Departamento Casiquiare from whence it derives its epithet. So far it has been collected from the Río Baria and the Río Siapa in the south to Cerro Yapacana and the Río Atabapo in the north. Approximately one-fifth of the cited collec- tions have some specimens with conspicuous, long stolons. Apparently in this species, as in many of its congeners capable of producing sto- lons, stolon production is negatively correlated with density of plants and in general is highly dependent upon local growing conditions. Those plants (e.g., Davidse et al. 17421, Huber & Medi- na 5858) from completely open exposures in her- baceous, white-sand savannas have very short, erect, stiff leaves, whereas those from more fa- vorable sites, such as sabanetas (e.g., Davidse et al. 16856) and the margins of savannas (e.g., Davidse et al. 16907A), have a more luxurious, lax growth habit. Axonopus jeanyae Davidse, sp. nov. TYPE: Pan- ama. Coclé: area between Cano Blanco del eni Caño Sucio and Chorro del Rio Tife, 80°36'30"—80°38'W, 8?42'19"—8?43'06"N, spray basin of waterfall, 3 Feb. 1983. G. Davidse & C. W. Hamilton 23570 (holotype, MO: isotypes, ISC, PMA, US). Figure 5. Axonopus jeanyae Davidse, sp. nov. A. ciliatifolius Swallen affinis sed nodis et spiculis p et gluma et lemmate anthoecio longiore Densely iue a perennial; culms 50- 75 cm tall, the internodes 1-2 mm wide, solid or nearly so, aa: flattened, especially to- ward the base, the nodes mostly 2, blackish, often noticeably elongated, appressed pubescent with hairs to 1.5 mm long. Sheaths mostly less than 10 cm long, keeled, not strongly USA gla- brous, the ciliate at least when young; flag leaf sheath to 16 cm ww 2s not differentiated; ligule a ciliolate membrane 0.5 mm ong, the membrane 0.2 mm long, the cilia 0.3 mm long; blades to 25 cm long, 2-3 mm wide, folded at the base, flat to folded above, papillose- ciliate towards the base, otherwise glabrous, the apex obtuse, ps Inflorescence 6-12 cm i m long; peduncles pore 2, well-exserted; racemes 2-5, 4-11 cm long, spreading at acute angles, the axils bend to 422 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 ee 5. Holotype of Axonopus jeanyae. — glu — C. Adaxial view of spikelet showing the lower lemma. [After Davidse & Hamilton 23570 (MO).] pubescent, the rachis 0.4—0.6 mm wide, d 10-13 spikelets per 25 mm; pedicels 0.2-0.5 m long, glabrous. Spikelets 1.9-2.5 mm long, 0. E 0.8 mm wide, elliptic-oblong, obtuse; upper glume about as long as the lower lemma, 4.5-nerved, the midnerve not or only weakly developed, with appressed lines of pubescence between the nerves; lower lemma similar to the upper glume; an- thoecium 0.1—0.4 mm shorter than the upper glume, brown, apically minutely papillate, ob- tuse; lodicules 2, minutely 3-lobed, fleshy, vas- culated, glabrous; stamens 3, the anthers 1.0-1.1 —A. Habit. —B. Abaxial view of spikelet showing the upper mm long, purple; styles 2, separate; stigmas plu- mose, purple; caryopsis 1.3 mm long, 0.7 mm wide, oblong, broadly obtuse, the hilum elliptic- punctate, the embryo ca. 2 the length of the cary- opsis. The perennial habit, glabrous racemes, and brown anthoecia place A. jeanyae in sect. Ax- onopus ser. Suffulti of Black (1963). This new species appears to be most closely related to A. ciliatifolius Swallen, a species so far known only from Belize. Axonopus ciliatifolius differs in its 1987] DAVIDSE—AXONOPUS 423 glabrous nodes and more fragile, glabrous, less generated by Missouri Botanical Garden collec- strongly nerved glume and lower lemma, both tors in Mesoamerica. of which are as long as the anthoecium. The epithet recognizes the contributions of my LITERATURE CITED wife, Jeany Vander Neut Davidse, to the Flora Biack, G. A. 1963. Grasses of the genus Axonopus. Mesoamericana project. She is largely respon- Adv. Front. Pl. Sci. 5: 1-186. sible for processing the thousands of collections TWO NEW MESOAMERICAN SPECIES OF CHUSQUEA (POACEAE: BAMBUSOIDEAE)! LYNN G. CLARK? ABSTRACT ew Mesoamerican species of Chusquea are described and ee both from complete Two one ss of vegetative and flowering material. Chusquea grandiflora, a cloud-forest species from nama and northwestern Colombia, has infravaginal branching, long, relatively narrow foliage leaves, open panicles, and rather large spikelets. It is allie e smaller, wider, yellowish-green, and strongly eae WV This species is probably most n. related to C. nelsonii Scribn. & Smith, and C. mueller Munro Duri tion ofa treatment of Chusquea Kunth for Flora Mesoamericana, examination of herbarium material revealed the existence of two more undescribed species of this diverse bamboo genus. Both are represented by complete collections of vegetative and flowering material. For explanation of terminology relating to the vegetative parts, especially buds and branching, see Clark (1985). In the specimen citations, a flowering specimen is indicated by the insertion of (fl) after the collection date. Chusquea grandiflora L. G. Clark, sp. nov. TYPE: Panama. Panamá: along road past Cerro Jefe toward La Eneida, 6 Jan. 1971 (fl), Croat 13070 (holotype, ISC; isotypes, COL, CR, K, MEXU, MO as two sheets, PMA, US). Figure 1A-D. Culmi scandentes, usque ad 18 m longi. Folia cul- morum 16-50 cm longa; vaginae 12- 33 cm longae, 2- 3-plo longiores quam laminae, abaxiales scabrae su- perne; laminae 4-17 cm longae, Abas aias scabrae; cin- glabrae, non tessellatae, ap- icibus brevi-setosis, basibus attenuatis. Inflorescentia aperta, paniculata, 6-11 cm longa; rhachis ron ngulati pubescent "ai 9.7-12.6 mm longae, scaberulae. ae 2 s E .4 mm longa, l-nervis; gluma II 2.2-4. 8 uen a, 1-3-nervis. Lemmata sterilia 2; lemma ste- rile I DES , 5.2-6.7 mm longum, 5-7-nerve; lem- ma sterile II subulatum, 7-9.5 mm longum, 7-nerve. emma fertile subulatum, 8.4-10.9 mm longum, 7-9- nerve. Palea bicarinata, 6.9-10.2 mm longa, 6-8-ner- vis. Culms clambering, viny, to 18 m long. Inter- nodes terete, smooth to scabrous-hirsute just be- low the nodes. Culm leaves 16-50 cm long, the juncture of sheath and blade abaxially indistinct; sheaths 12-33 cm long, 2-3 times as long as the lade, abaxially smooth at the base, scabrous toward the apex; blades erect, usually persistent, 4-17 cm long, abaxially scabrous; girdle well- developed, 2-3 mm wide, glabrous; corky ridge present at the juncture of the sheath and girdle. Nodes with one triangular central bud subtended by numerous (up to 50) smaller, subequal sub- sidiary branches; sheath scar dipping markedly below the bud/branch complement. Branching infravaginal; the central bud developing tardily or sometimes not at all; leafy subsidiary branches not rebranching. Foliage leaves with sheaths gla- brous, the margins ciliate near apex; blades 8- 20 cm long, (0.5—)0.7-1.4 cm wide, L: W = 11- 27, the adaxial surface glabrous, the abaxial sur- face glabrous, not tessellate, the apex short se- tose, the base attenuate; pseudopetiole more or less distinct, 2-4 mm long; outer ligule evident, o 1 mm long; inner ligule truncate, asymmet- rical, 1-1.5(-2) mm long. Inflorescence an open panicle 6-11 cm long, usually not fully exserted from the subtending sheath; rachis triquetrous, pubescent; branches strongly spreading, angular, pubescent, the lower ones short, up to 3 cm long; pedicels 1-2 mm long, angular, pubescent. Spike- lets 9.7-12.6 mm long, scaberulous. Glumes 2; glume I ca. % the spikelet length, 2-2.4 mm long, apically acute, abaxially scaberulous, 1-nerved; glume II ca. '^ the spikelet length, 2.2-4.8 mm long, apically pointed, abaxially scabrous, 1—3- thank Dr. Richard W. Pohl for reviewing this manuscript, and Dr. Gerrit Davidse for editorial and agrostological assistance ? Department of Botany, Iowa State University, Ames, Iowa 50011, U.S.A. ANN. MissounRi Bor. GARD. 74: 424-427. 1987. 1987] FIGURE 1. B. Inflorescence, scale = 1 cm.— C. scale = 1 cm. w^ D based on E 201 14, B, ir cm.—F. Foliage leaf blade, adaxial view, scale = with emerging branches, scale CLARK— CHUSQUEA kelet, scale - 1 mm based on Croat 13070.) E-H. "i per e af 425 bero nin grandiflora and C. aperta. ard C. grandiflora. — A. Culm leaf with emerging branches, m. — D. Foliage leaf beni abaxial view, Culm le orescence, ) e = | cm.—H. Spikelet, anis — | mm. (E based on Soderstrom 2237, F-H ES on pa iae 2239. nerved. Sterile lemmas 2; sterile lemma I ca. 2 the spikelet length, 5.2-6.7 mm long, apically subulate, abaxially scaberulous, 5-7-nerved; sterile lemma II ca. % the spikelet length, 7—9.5 mm long, apically subulate, abaxially scaberu- lous, 7-nerved. Fertile lemma 8.4-10.9 mm long, apically subulate, sence scaberulous, 7—9- nerved. Palea 2-keele te except toward the base, 6.9-10.2 mm long, apiculate, abaxially sca- berulous, 6-8-nerved, the sulcus pubescent. Lod- icules 3; 3—3.5 mm long. Stamens 3; anthers 4.3— 5.5 mm long. Fruit unknown. Additional specimens examined. COLOMBIA. tween Alto del Tigre and 1944 (fl), Core 817 (US). CHOCO: abis. La Sirena, 4 km W of La Mansa 426 at top of Cord. Occidental, Gentry & Renteria 24203 US). PANAMA. CHIRIQUÍ: Vicinity of Gualaca ca. 7.8 mi. from Planes de Hornito, La Fortuna on road to dam site, Antonio 5182 (MO, US); camino hacia la finca Landau, NE del oo e Fortuna (sitio de presa), i Sep. 1976 (fl), Correa et al 2185 (F, MO, NY). coctÉ: El Valle, Soderstrom 2014 (US). PANAMA: 17 Oct. 1977 (fl), Folsom & Page 5944 (MO, PMA, , VEN); La Eneida, 12 km N of Goofy Lake, Sod- erstrom 2006 (US). vERAGUAS: Cerro Tute, trail past agricultural school near Santa Fe, Antonio 1878 (MO). Chusquea grandiflora is characterized by in- fravaginal branching, culm leaves abaxially sca- brous (except basally), open panicles with short, spreading primary branches, pubescent rachis, and scaberulous spikelet with glume II usually twice as long as glume I. This species is known from montane forests at altitudes of 700 to 1,700 m in Panama and northwestern Colombia. Gen- try & Renteria 24203, a vegetative specimen re- ferred to this species, was collected at 2,300- 2,400 m Chusquea grandiflora lacks the leafless, fibril- lar branchlets so characteristic of C. scabra, but otherwise the two species are strikingly similar vegetatively. Chusquea scabra usually exhibits much more rounded leaf bases and consistently scabrous internodes. Chusquea grandiflora is also closely related to C. foliosa L. G. Clark, C. lon- gifolia Swallen, and C. patens L. G. Clark, shar- ing infravaginal branching and long, relatively narrow foliage leaves with these three species. e open panicles with strongly spreading branches of C. grandiflora and C. patens are sim- ilar in some specimens, but C. grandiflora is dis- tinguished by its pubescent rachis, shorter inflo- rescence branches, larger spikelets, and wider foliage leaf blades. Chusquea aperta L. G. Clark, sp. nov. TYPE: Mex- ico. Oaxaca: 107 km SW of Tuxtepec, 2,650 m, 4 Oct. 1977 (fl), Soderstrom 2239 (ho- lotype, US). Figure 1E-H. Culmi usque ad | cm diam., usque ad 1-2 m alti. Folia culmorum: vaginae persistentes, 6.8-8.5 cm lon- alis. Laminae foliorum rigidae, 7-12 cm longae, 1.5 cm e, L: W = 7-11, glabrae, valde a inferne, apicibus brevi-setosis, basibus rotundatis = apic Doe aiden aperta, paniculata, 7-11 c dern triquetra, glabra; rami eius us ue a ongi, angulati, glabri. Spiculae £z: mm longae, Pues adpressae. Glumae 2, squam ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 formes; gluma I 0.3-0.5 mm longa, enervis; gluma II .4-0.8 mm longa, enervis. Lemmata steri sterile I apiculatum, 4-4.6 mm longum, 1-3-nerve; lemma sterile II subulatum, 4.5-5.1 mm longum, sna nerve. Lemma fertile subulatum, 5.7-7.1 mm, S nerv Palea bicarinata, 5.5-6.8 mm longa, 4—6-nerv Culms to 1 cm diam., to 1-2 m tall. /nternodes terete, solid, smooth. Culm leaves with the sheaths persistent, the overlapping margin fused to the sheath to 5 mm above the base, 6.8-8.5 cm long, abaxially smooth; blades evidently not persis- tent, in the one example seen, blade triangular, cm long, abaxially smooth; girdle asymmetri- cally developed, prominent only near the bud complement, to 5 mm wide; inner ligule 2-3 mm long. Nodes swollen, the central bud subtended by up to 25-30 subsidiary branches; supranodal ridge prominent. Branching infravaginal, the central bud often developing; leafy subsidiary branches occasionally rebranching. Foliage pun >= the blades stiff, 7-12 cm long, 0.8-1.5 c e, L:W - 7-11, yellowish-green, the RE si smooth and not tessellate, the abaxial surface smooth and strongly tessellate, the apex short-setose, the base rounded to attenuate; pseudopetiole usually distinct, 2-3 mm long; outer ligule a stiffrim to 0.5 mm long; inner ligule elongate, 2-9 mm long, rounded to acute at the tip. Inflorescence an open panicle 7-11 cm long, completely exserted from the subtending sheath on a peduncle up to 12 cm long; rachis trique- trous, glabrous; branches spreading, angular, gla- brous, the lower branches to 4—6 cm long; ped- icels variable, to 14 mm long, angular, glabrous. Spikelets 6-8 mm long, glabrous, appressed to branches. G/umes 2, scalelike, less than Y,, the spikelet length; glume I 0.3-0.5 mm long, nerve- less; glume II 0.4—0.8 mm long, nerveless. Sterile lemmas 2, ^—h the spikelet length; sterile lemma I 4—4.6 mm long, apiculate, glabrous, 1-3-nerved; sterile lemma II 4.5-5.1 mm long, subulate, gla- brous, 1-5-nerved. Fertile lemma 5.7-7.1 mm long, subulate, glabrous, 7-nerved. Palea 2-keeled, the upper half sulcate, 5.5-6.8 mm long, acute, glabrous, 4—6-nerved. Lodicules 3. Sta- mens 3; anthers 4-5 mm long. Fruit unknown. oe specimens examined. MEXICO. OAXACA: a. 0.5 N i. SW ue 75 to Tuxtepec, 16 Aug. 1975 (fl), Davidse & Davidse 9733 (MO, US); on road between Oaxaca and Tuxtepec, 11 Apr. 1975 (fl), Fisher & Engleman 75-41 (MO); Distrito de Ixtlan, Sierra de Juarez, ruta 1987] 175 Tuxtepec a Oaxaca, ca. 0.5 km al S de Cerro Pelon, 13 Apr. 1982 (fl), Lorence et al. 4004 (US), 108 km SW of Tuxtepec on road to Oaxaca, Soderstrom 2237 (US). According to label data, Chusquea aperta oc- curs in pine-oak cloud forests at elevations from 1,670 to 2,750 m. At present, this species is known only from a relatively restricted area of Oaxaca, Mexico, on the road between Oaxaca and Tux- tepec. Chusquea aperta is characterized by in- fravaginal branching, deciduous culm leaf blades, stiff, yellowish-green foliage leaves with blades strongly tessellate abaxially, and long-exserted, open panicles with spikelets appressed to the spreading branches. Vegetatively, C. aperta re- sembles C. nelsonii Scribn. & Smith, but can be distinguished by its glabrous, abaxially tessellate foliage leaf blades, in contrast to the narrower, abaxially pubescent, non-tessellate leaf blades of the latter species. The inflorescences and spike- lets of C. aperta are similar to those of C. muelleri unro (7 C. carinata Fournier and C. mexicana Hackel), but the inflorescences of C. muelleri are smaller, less open, and not long-exserted. With regard to the spikelets, the glumes of C. aperta CLARK—CHUSQUEA 427 are both small and scalelike, while the glumes of C. muelleri are more developed, with the sec- ond glume usually about paoe as long as the zt The ty tly older, spen T Sod (but still with some spikelets), Fw a few younger, fresh inflorescences derived from regrowth of subsidiary branches. These newer spikelets are just at anthesis, and are slightly larg- er and fuller than the old spikelets produced dur- ing the primary flowering period. The newer spikelets are unusual because they all have a one- or two-keeled palea-like bract in the axil of the second sterile lemma. I observed individual spikelets with two fertile florets subtended by two sterile lemmas in some specimens of a few Chus- quea species, but this is the first indication ofa rudimentary floret in the position of a sterile lemma. No other flowering specimens of C. aper- ta show this condition. The abnormal spikelets were left out of the description. LITERATURE CITED CLARK, L. G. 1985. Three new species of Chusquea (Gramineae: Bambusoideae). Ann. Missouri Bot. Gard. 72: 864-873. NOTES NEW COMBINATIONS IN CHUSQUEA (POACEAE: BAMBUSOIDEAE) The genus Swallenochloa McClure was seg- regated from Chusquea Kunth solely on the basis of differences in certain vegetative features (McClure, 1973). To my knowledge, McClure never collected Swallenochloa, and only limited, incomplete herbarium material of this group of species was available to him. Soderstrom & C. Calderón (1978) applied the terms extravaginal and intravaginal respectively to the branching patterns in Chusquea and Swallenochloa, and discussed other vegetative differences in addition to those listed by McClure. As the result of ex- tensive field and herbarium studies, I determined that the species of Swallenochloa constitute a coherent, natural group that falls within the ge- neric limits of Chusquea. Details of the reduction of Swallenochloa to a section of Chusquea will be presented in a later publication (Clark, in prep.). For the present, this transfer necessitates new combinations for three species. Chusquea angustifolia (Soderstrom & C. Cal- derón) L. G. Clark, comb. nov. Swallenoch- loa angustifolia Soderstrom & C. Calderón, Brittonia 30: 303. 1978. TYPE: Venezuela. Tachira: subparamo y bosque enano, faldas inmediatamente debajo del Paramo de Tama, cerca de la frontera Colombo-Ve- nezolana, Steyermark, Dunsterville & Duns- terville 98615 (holotype, US; isotype, VEN). ANN. MISSOURI Bor. GARD. 74: 428. 1987. Chusquea longiligulata (Soderstrom & C. Cal- erón) L. G. Clark, comb. nov. Swallenoch- loa longiligulata Soderstrom & C. Calderón, Brittonia 30: 305. 1978. TYPE: Costa Rica. San José: along the Carretera Interameri- cana, ca. 5 km SE of Empalme, near Tres de Junio, Poh! & Selva 12842 (holotype, US; isotypes, F, ISC, MO). Chusquea vulcanalis (Soderstrom & C. Calde- rón) L. G. Clark, comb. nov. Swallenochloa vulcanalis Soderstrom & C. Calderón, Brit- tonia 30: 309. 1978. TYPE: Costa Rica. Car- tago: abrupte dominante La Playita, Volcán Irazu, Pittier 14126 (holotype, US; isotypes, ISC, US) LITERATURE CITED MCcCLUuRE, F. > - Genera of bamboos native to the New rld (Gramineae: Bambusoideae). — pens Bot. 9: ix-xii, 1-148. [Edited y T. R. Soderstrom.] SODERSTROM, T.R. & ALDERÓN. 1978. Chus- qu ea and Swallenochloa (Poaceae: Bambuso - species. Brit- tonia 30: 297-312. —Lynn G. Clark, Department of Botany, Iowa State University, Ames, Iowa 50011, U.S.A. NOTES ON CORNUS (CORNACEAE) IN SOUTH AMERICA The family Cornaceae is represented in South America by the genera Cornus L. and Griselinia Forster f. The latter, often segregated as Grise- liniaceae, occurs in Brazil and Chile, and in New Zealand, while the former has been known pre- viously from Bolivia north to Colombia in the ndes. We report here, for the first time, Cornus in Venezuela, ~ viai the first record of the family in Venezu Macbride pene was first to record the genus from South America. Based on fragmentary fruiting material collected from Peru and Boliv- ia, he described two species, C. peruviana and C boliviana, the former based on the collection Macbride 3439 from Cani, Depto. Huánuco, Peru, the latter based on Bang 1799 from Bolivia without exact locality indicated. His decision to separate the two collections as distinct taxa was based on his observation. of the "equally two- armed" trichomes on the abaxial leaf surface of the Peruvian plant as contrasted with the “un- equally branched or pronged" hairs with “one of their two ‘arms’ being much longer than the oth- er" in the Bolivian material. A year later, how- ever, Macbride (1930) changed his mind and concluded that Cornus peruviana was identical with an unpublished species of Viburnum from Peru collected by Ruiz & Pavon which he ex- amined at B, and, accordingly, he transferred it to Viburnum peruvianum (J. Macbr.) J. F. Macbr. At the same time, he decided that the Bolivian material did not belong to Cornus, but, being uncertain as to its proper family position, did not assign it to any genus. Subsequently, Standley (1935) re-examined the specimens described by Macbride and concluded that they did indeed belong to Cornus, and that, moreover, they were conspecific. Standley chose the epithet peruviana for them. In the Flora of Peru Macbride (1959) accepted Standley's con- clusions and placed C. boliviana J. F. Macbr. in synonymy. During an expedition to the Andes of Estado Táchira, Venezuela, near the Colombian border, the authors found a fallen branch from a tree that proved, upon later examination, to belong to the genus Cornus. In determining that this collection pertained to Cornus, and not to Viburnum, the criterion of leaf pubescence, as indicated by So- lereder (1899) and Metcalfe & Chalk (1950), ANN. Missouri Bor. GARD. 74: 429-430. 1987. proved critical. A comparison of the Steyermark & Liesner collection from Venezuela with other South American specimens initiated the present study of the genus. In general, we have observed, after examina- tion of available herbarium material, that the specimens from Venezuela and Colombia show a more appressed type of indument on the calyx, hypanthium, and lower surface of the leaf in con- trast with collections from Ecuador, Peru, and Bolivia, which have a more loosely crisp, spread- ing, and often denser type of pubescence. How- ever, the total number of specimens examined from South America remains inadequate for con- clusions as to the variability present. The ap- parent differences noted between the northern and southern populations may disappear when more abundant material becomes available. For the present, therefore, we do not recognize any segregation of taxa, and conclude that the plants from Venezuela are conspecific (sensu lato) with those from the other parts of South America. In this connection, we are employing the name C. peruviana J. F. Macbr. as selected by Standley (1935) when he concluded that the Peruvian and Bolivian populations were conspecific. ecimens examined (listed ee ssa uth). VEN shale substrata, south of El Reposo, 14 km SE licias, 7°31’N, 72?24'W, 2,150-2,300 m, 22-23 Jul. rmark & Liesner 118420 (MO, V 1935, pes 271 (F). HUAN co: ped NE of Mito, 2,575 m = 1923, Macbride 3439 (type en puaa F); . CAJAMARCA: Prov. Ce sor canyon of the Río Marañon above Balsas, 5 km bel mit of the road to oo 24 deed 1964, Fehon: "i Wright 5336 (US); D AZONAS: Leimebambo, 2,100 m, 23 Dec. 1962, Woytowski 7796 (MO, US); Ambay, 3,100 m, Tus. 1938, Univ. Cuzco s.n. O). BOLIVIA. DEPT. HABAMBA: Ayopaya, Sailapata, 2,800 m, Nov. 1935, Cardenas 3355 (US); Bang que (type of C. boliviana. , The following key indicates the main charac- ters distinguishing Cornus from Viburnum: 1. Abaxial surface of leaves with 2-branched, unicellular, nodose trichomes incrusted with lime carbonate; flowers tetramerous; petals free, separate; l Cornus l. Abaxial surface of leaves not as above, the 430 pubescence, when present, of simple, stellate, peltate, tufted, or glandular hairs; flowers pen- tamerous; corolla HE ovary uniloc- | Nn SOS OS MOSA UPS ks Viburnum LITERATURE CITED CRONQUIST, A. 1981. Pp. 668-670, 1006-1008 in An Integrated System of Classification of Flowering Plants. Columbia Univ. Press, New York. MACBRIDE, J. F. 1929. To a genus new to South merica. Trop. Woods 19: 5. 19 outh NA viburnums incor- rectly desorbed as new species of Cornus. Trop. Woods 24: 29. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 . 1959. Cornaceae. Fl. Peru, Field Mus. Nat. Hist., Bot. Ser. 13(5), no. 1: 44. METCALFE, C. R. & L. CHALK. Pp. 735-737, 752-754 in Anatomy ofthe Dicotyledons, Volume 2. Clarendon Press, Oxf SOLEREDER, H. 1899. Pp. 487 -495 in Systematische Anatomie der vio ace Verlag von Ferdi- nand Enke, Stuttgar STANDLEY, P. C. 1935. "The genus Cornus in South America. Trop. Woods 43: 16-17. —Julian A. Steyermark and Ronald Liesner, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. A NEW COMBINATION IN LUCILIA (COMPOSITAE—INULEAE) Lucilia pusilla (Kunth) Hieron. (1900) is the generally accepted name for a small, caespitose herb from the high Andes of Venezuela, Colom- bia, Ecuador, and Peru. While working on a no- menclator of Conyza (Compositae—Astereae), I found that its basionym Conyza pusilla Kunth (1818) is illegitimate, being a later homonym of C. pusilla Houtt. (1779) from South Africa. De Candolle changed the name of Conyza pusilla Kunth to C. kunthiana DC. (1836). Since under article 55.1 of the 1983 International Code of Botanical Nomenclature the specific epithet of the former correct name must be retained on generic transferences, the correct name for this species in Lucilia is L. kunthiana (DC.) Zardini, comb. nov. iiu 1 yva Y" hu mm $ nivw $.9 ~ x nnn I Wilt ae S S el | Lucilia kunthiana (DC.) Zardini, comb. nov. Based on Conyza kunthiana DC., Prodr. 5: 379. 1836 (new name for C. pusilla Kunth). Gnaphalium kunthianum (DC.) Kuntze, Revis. Gen. Pl. 3(2): 152. 1898. Conyza pu- silla Kunth, Nov. Gen. Sp. 4: 69. 1818, non C. pusilla Houtt. (1779). Lucilia pusilla TYPE: Ecuador: in Andean region, Humboldt & Bonpland s.n. (P holotype; P isotype, B isotype photograph F 15128). I thank Dr. Zijlstra (U) for confirming the ne- cessity of this new combination. — Elsa Zardini, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. FiGURE 1. Holotype of Conyza pusilla Kunth (P, photograph Zardini n. 1036, MO). ANN. Missour!I Bor. GARD. 74: 431. 1987. CHROMOSOME COUNTS OF MISSOURI ASTERACEAE AND POACEAE Few native Missouri plants are chromosomal- ly known from Missouri populations. The list of chromosome counts for composites and grasses in Table 1 is a small contribution in this area. The counts were made from standard anther squashes stained in propionic-carmine and dis- sected from buds fixed in Carnoy's or Newcom- er's fixative. Except for Paspalum laeve, meiosis was normal in all collections, and the observed numbers agree with previously reported num- bers as summarized in Federov (1969), Moore (1973, 1974, 1975), and Goldblatt (1981, 1984, 1985). Vouchers are deposited in MO and KUH. The chromosome number for Paspalum laeve has previously been reported as 2n = 40 (Brown, 1948), 2n — 60 (Burton, 1942, as P. longipilum Nash), and n = 40 (Banks, 1964). Our count of n — 29 apparently represents an aneuploid re- duction from the hexaploid level of n = 30. Mei- osis and pairing were normal. The second pop- ulation sampled had the heptaploid number, n = 35, and the chromosomes were almost complete- ly asynaptic in early meiosis. No more than five bivalents were ever observed at diakinesis or metaphase I (Fig. 1). When chromosomes were paired, the association was very loose. Structures that we believe may be micronucleoli were pres- ent in variable numbers at diakinesis (Fig. 1A). We suspect, based on a similar pattern of asyn- apsis in the tetraploid apomictic cytotype of P. conjugatum Berg. (Fang & Li, 1966; Mehra, 1982), that the plant may have been apomictic. Since only one plant was examined, we do not know whether this condition was isolated or widespread in the population. The species is morphologically variable and further cytotax- y be help- ful in relating some of this variation to ploidy levels Supported by NSF grant INT-8510317. LITERATURE CITED BANKS, D. J. 64. Chromosome counts for Paspa- lum. Rhodora 66: 368-370. BRowN, W. V. 1948. Acytological study in the Gra- mineae. Amer. J. Bot. 35: 382-395. BuRTON, G. W. 1942. A cytological study of some species in the tribe Paniceae. Amer. J. Bot. 29: 355-359. FANG, J. S. & H. W. LI. 1966. Cytological studies in TABLE l. Chromosome numbers of Missouri Asteraceae and Poaceae. Taxon n Voucher? Asteraceae Erigeron strigosus Muhlenb. 18 Vahidy & Davidse 21 Eupatorium coelestinum L. f. 10 Vahidy & Davidse 20 Lactuca floridana (L.) Gaertner 17 Vahidy & Davidse 12 Rudbeckia laciniata L. 27 Davidse & Vahidy 30845 Rudbeckia missouriensis Engelm. 19 Davidse & Vahidy 30838 Solidago nemoralis Aiton 27 Vahidy & Davidse 15 Solidago ulmifolia Muhlenb. 9 Davidse & Vahidy 30835 Poaceae Leersia virginica Willd. ca. 24 Davidse & Vahidy 30840 Panicum capillare L. 9 Vahidy & Davidse 17 Paspalum fluitans (Elliott) Kunth 10 Davidse & Vahidy 30854 Paspalum laeve Michaux var. /aeve 35 Vahidy & Davidse 1 Paspalum laeve Michaux var. pilosum Scribner 29 St. Louis, Tower Grove Park, Da- vidse & Vahidy 30846 Paspalum pubiflorum Rupr. var. glabrum Vasey 30 Vahidy & Davidse 24 Paspalum setaceum Michaux var. muhlenbergii 10 Vahidy & Davidse 14 (Nash) D. Banks Sorghastrum nutans (L.) Nash 20 Vahidy & Davidse 16 “Collected at Shaw Arboretum, Franklin County, Missouri, unless indicated otherwise. ANN. Missouni Bor. GARD. 74: 432-433. 1987. 1987] URE 1. Diakinesis in Paspalum laeve var. laeve, n = NOTES b e? vh. < e ? è A [] ; ~ . y ° ° E b o3 * se e 4 e e ° ; , ° £ r^ dil" . 4 4 ° < e $ e e B * LJ * ° 35.—A. Photomicrograph showing nearly complete FIGURE 1. asynapsis.—B. Camera lucida drawing of A with 5, (marked) + 60, pd a conjugatum Berg. Bot. Bull. Acad. Sin. Feoesov - A. (editor). 1969. Chromosome Num ers of PUE Plants. V. L. Komarov Botanical stitute, Lenin M secre P. (editor). 1981. Index to plant chro- e numbers 1975- xn Monogr. Syst. Bot Missouri Bot. Gard. 5: 3. (editor). ce to plant chromosome numbers 1979-1981. Monogr. Syst. Bot. Missouri Bot. Gard. 8: 1-427. (editor). 1985. Index to plant chromosome numbers 1982- MAS Monogr. Syst. Bot. Missouri Bot. Gard. 13: 1-224. MEHRA, P. N. vus Cytology of East Indian Grasses. P. P. Kapur, New Delhi. Moore, R. J. (editor). 1973. Index to plant chro- mosome numbers 1967-1971. Regnum Veg. 90: (e di tor. 1974. Index to plant chromosome numbers for 1972. Regnum Veg. 91: 1-108. (editor). 1975. Index to plant chromosome numbers for 1973/74. Regnum Veg. 96: 1-257. hsan A. Vahidy, Department of Genetics, c Box 299, St. Louis, Missouri 63166, U.S. Youji Shigenobu, Department of Natural uis Naruto University of Teacher Education, Taka- shima, Naruto-shi 772, Japan. A NEW SPECIES OF THRASYA (POACEAE: PANICOIDEAE) FROM THE MOSQUITIA OF NICARAGUA AND HONDURAS Recent work for the account of the Poaceae for the Flora Mesoamericana has brought to light a previously undescribed species of Thrasya. ups —— Davidse & Burman, sp. nov. TYPE: Nicaragua. Zelaya: Along banks oa pee gallery forest of Rio Likas near Silima Lila, ca. 14?30'N, 83°50’W, 50 m, 5 Mar. 1979, John J. Pipoly 4107 (ho- lotype, MO; isotypes, HNMN, SP, US). Fig- ure | Gramen perenne; culmi 65-110 cm longi; ligula membranacea, 1.5-2.4 mm longa; laminae lineares, -32 papillosum minute; flos superus perfectus. Perennial herb; culms 65-110 cm long, erect, sometimes rooting at the lower nodes, the inter- nodes slightly compressed, hollow, mostly gla- brous, the upper portans ang the nodes ap- uber ulent toward the apex, keeled, the collar appressed pu- bescent; ligule a membrane 1.5- m long; blades 16-32 cm long, 9-16 mm de linear, flat, glabrous or puberulent toward the base be- low, acuminate, the base slightly rounded to gradually narrowed; leaf subtending the inflores- cence distinctly smaller. Racemes terminal and axillary, solitary, 10—16 cm long, arcuate, usually well-exserted from the sheath; peduncles puber- ulent near the tip or entirely glabrous; rachis 2.0— 2.8 mm wide, winged, dorsally and ventrally gla- brous, the margin minutely scabrous, the base with an inconspicuous tuft of hairs. Spikelets 3.4— 4.0 mm long, 1.4-1.8 mm wide, glabrous, ellip- tic, broadly acute, arrayed in one row, oriented back-to-back, paired, the pairs borne on alter- nate sides of the rachis; pedicels puberulent, un- equal, the upper 1.8-2.0 mm long, its lower half adnate to the rachis, the lower 0.2-0.4 mm long; glumes with the bases clasping and slightly swol- len; lower glume dimorphic between the short- and long-pedicellate spikelets, variable in the short-pedicellate spikelets, 0.6-2.5 mm long, tri- angular, membranous to herbaceous, 0-1-nerved, ANN. Missouri Bor. GARD. 74: 434-436. 1987. obtuse to acute or aristate, uniform in the long- pedicellate spikelets, 0.7-1.2 mm long, triangu- lar, membranous, nerveless, acute; upper glume 2.8-3.4 mm long, ca. 3⁄4 as long as the spikelet, herbaceous, 5(-7)-nerved; lower lemma as long as the spikelet, subindurate, deeply sulcate on the back, sometimes splitting at maturity, 6(—7)- nerved, the midnerve often absent, the inner pair of nerves well-developed, slightly keeled and crested toward the tip; lower palea as long as the lower lemma, 2-nerved and 2-keeled; lower flow- er staminate: lodicules 2, the stamens 3, the an- thers 1.3-1.9 mm long, purple; upper lemma 3.0- 3.6 mm long, 1.4-1.6 mm wide, indurate, ellip- tic, minutely papillose, the tip bearing a tuft of minute hairs; upper palea of the same texture as the upper lemma; upper flower perfect: lodicules 2, the stamens 3, the anthers 1.4-1.8 mm long, purple, the styles 2, separate, the stigmas plu- mose, purple; caryopsis (immature) ca. 1.7 mm long, 1.1 mm wide, the embryo nearly '^ as long as the caryopsis, the hilum narrowly elliptic, ca. !^ as long as the caryopsis. Paratypes. NICARAGUA. ZELAYA: Comarco del Cabo, Kornuk Creek, Puente Pozo Azul, 10 Jul. 1972, Sey. mour 5802A (MO), 5803 (MO). HONDURAS. GRA A DIOS: 5 km de Puerto Lempira, 28 Jan. 1984, oe & Cruz 8621 (TEFH). The generic placement of this new species must be considered since the limits of the genus Thra- sya have always been difficult to circumscribe posed of species that have the following features: 1) winged, one-sided, solitary racemes; 2) solitary spikelets arrayed in a single row; 3) back-to-back orientation of the spikelets, i.e., the backs of the lower glume and lower lemma of the short-ped- icellate spikelet face the backs of these same structures of the long-pedicellate spikelet. The problem of ihe outlying Species that show vary- ing i between the core group of Thrasya and the group of Paspalum (sensu Chase, 1929) has been dis- spikelets is superficially lost through the fusion of both the long and short pedicels to the rachis 1987] so that the free portion of all the pedicels is ba- "x the same length. . mosquitiensis the spikelets are arranged ina gibt row in a back-to-back orientation along a winged, one-sided, solitary raceme. Although the spikelets are arranged in a single row, the spikelets are clearly paired and the longer pedicel is only partially fused to the rachis (Fig. 1). The spikelet along the rachis. With respect to | the distinctly in T. mosquitiensis the basal half of the upper pedicel is adnate to the rachis, in contrast to the Decumbentes group where both pedicels of a spikelet pair remain completely free. In addition, TE : aay ea ý in p p g in several rows and the regular back-to-back po- sition of the spikelets is never attained. There- fore, although T. mosquitiensis is intermediate between the core group of Thrasya and the De- cumbentes group of Paspalum, similarities with Thrasya are greater, and this is the reason for its inclusion in that genus. There is a tendency for the spikelets to be di- morphic in the development of the lower glume within single inflorescences. In the short-pedi- cellate spikelets the lower glume may vary from nerveless, membranous, and obtuse to 1-nerved, herbaceous, and aristate. In the long-pedicellate spikelets the lower glume is uniformly nerveless, membranous, and acute. Also, the length of the upper glume relative to the upper lemma is great- er in the short-pedicellate spikelet than in the long-pedicellate spikelet. This is most easily ob- served as the length of the upper lemma that is not covered by the upper glume: (0.1—)0.4—0.5 mm in the short-pedicellate spikelets, 0.5—0.7 mm in the long-pedicellate spikelets. Although such dimorphism is characteristic of Paspalum species of the Decumbentes group, it also occurs in two Thrasya species: T. campylostachya (Hackel) Chase and 7. petrosa (Trin.) Chase. Thrasya mosquitiensis seems to be most closely related to T. campylostachya, the only other Me- soamerican species with glabrous spikelets. Thrasya mosquitiensis is distinguished from the latter by the larger size of all its parts, most no- ticeably the spikelets (3.4-4.0 mm vs. 2.6-3.0 mm long and 1.4-1.8 mm vs. 0.8-1.5 mm wide), racemes (10-16 cm vs. 4-10 cm long), and blades (9-16 mm vs. 3-10 mm wide and 16-32 cm vs. 6—20 cm long). Moreover, the upper glume of the long-pedicellate spikelet is proportionally NOTES 435 = —— a 1 pe teen ee He LLILLVA . `... Tes... mom RR RTI Tm —— LB T xa e L n xx dero on ate RE FIGURE 1. Spikelet m. of Thrasya mosquitiensis Davidse & Burman showing the back-to-back orien- tation ofthe PUN ud rachis (partially removed n one side), and partially oyy maa of the long- pedicellate sakaki. Scale line = longer (34 vs. Y} the length of the spikelet), roader, more clasping, and less papery in tex- ture. Finally, the upper pedicel in 7. mosqui- tiensis tends to be less adnate to the rachis than 436 in 7. campylostachya, although the latter species is somewhat variable in this regard. The specific epithet refers to the area of Nic- aragua and Honduras, dominated by pine sa- vannas, that is commonly referred to as the Mos- quitia or sometimes as the Costa de Miskitos. We thank ken Myers for the illustration, and Peter Lowry an ful comments on oe manuscript. CI 101 USC- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 LITERATURE CITED BURMAN, A, G. 1982. Three iid md of Thrasya (Gramineae) from Brazil. Bri a 34: 458-462. — (In pre Xd Revision of um zo Thrasya. Acta Bot. Vene CHASE, A. 1929. The North American species of Pas- palum. Contr. U.S. Natl. Herb. 28: 1-310. —Gerrit Davidse, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.; and Alasdair G. Burman, Instituto de Botanica, Caixa Postal 4005, 01000 Sao Paulo, Brasil. A NEW SPECIES OF BULBOSTYLIS (CYPERACEAE) FROM TROPICAL AMERICA Bulbostylis, in the sense employed here, in- cludes ca. 100 species centering in the tropics and subtropics of both hemispheres, with best representation in Africa and South bidon It is usually placed in the tribe Scirpeae and is taxonomically (some state Sd ane he related both to Abildgaardia and Fimbristylis. It is dis- tinguishable from both by the sheaths which bear a sparse to dense beard of long, firm, often bristly hairs at the orifice and by the achenes which are surmounted by a persistent tubercle formed from the style base. Many species have weedy tenden- cies, are heliophytes, and thus grow abundantly in open areas that are artificially or naturally disturbed, in soils that are typically acid, sandy, azonal, and seasonally moist or wet. Thus they are much a part of vegetation of savanna, acid rock outcrops, natural or artificial clearings, and fluctuating, sandy-silty shores. Since such habi- tats are common in many areas of Amazonia, Bulbostylis species are an important component of the herbaceous vegetation. One species, fre- quently collected in the last decade along banks and bars of major streams in Amazonian Brazil, Colombia, and Venezuela, represents a new one that we describe here. Bulbostylis fluviatilis Kral & Davidse, sp. nov. TYPE: Venezuela. Amazonas: Dpto. Río Ne- gro, south side of hamlet of Santa Lucia, sandy recently cleared area for heliport, by Río Negro, common, 25 Nov. 1984, R. Kral 71973 (holotype, VEN; isotypes, F, FSU, GH, ISC, MICH, MO, NY, P, U, US, VEN, VDB). Figure 1. Bulbostylis tenuifolia (Rudge) Macbride affine a quo glumis mucronibus excurrentibus et antheris 2, 0.2- 0.3 mm longis differt Annual, densely cespitose, 5-30 cm high. Roots capillary-fibrous. Leaves few per culm, polysti- chous, sub-basal, the lowest ns bladeless or with blades shorter than the sheaths; principal leaves usually '^—'^ as long as the scapes, some- times as long, green; sheaths medially convex or keeled, multicostate, the costas strongly hirtel- lous, the broad, scarious, tan borders hirtellous, narrowing acutely to a long ciliate apex; blades 5-15 cm long, capillary, erect or spreading, proximally involute, 0.2-0.3 mm broad, penta- ANN. Missouni Bor. GARD. 74: 437-439. 1987. costate, distally triquetrous with costas scabrid- ulous, the tips narrow but blunt. Scapes 0.2-0.3 mm thick, capillary, erect, sharply 5—7-costate, smooth or proximally sparsely scabridulous. In- florescence a broad anthella 2-4 cm high, sub- umbellate, diffuse; involucral bracts 3-5, leaflike, subtending prophyllate Tays, ins longest bract equalling or slightly o he anthella, the central spikelet sessile, 8-4: mm "aa its lowest bracts also subtending primary rays; primary rays 0.5—3 cm long, erect to spreading, capillary and costate, 1—3-spicate; secondary rays 5-15 mm long. Spikelets mostly 3-6 mm long, lance-ovoid to lance-cylindric, pedicellate except for the cen- tral spikelet, acute; bracts 1.0-2.0 mm long, loosely spirally imbricate, lance-ovate, navicu- lar, glabrous or hispidulous, 1-nerved, medially carinate, the midnerve green, the apex acute, vil- losulous, the margins sparsely ciliolate, the cusps and mucros 0.1-0.3 mm long, excurvate, sca- bridulous, the broad sides scarious, tan or red- brown, glabrous or hirtellous; lowest bract 2-3 mm long, usually sterile, strongly cuspidate, the cusp to 2 mm long; stamens mostly 2, the fila- ments flat, the anthers 0.2—0.3 mm long, basi- xed, bilocular, laterally dehiscent, ellipsoidal, apiculate; style glabrous, branched medially to 3 slender, papillate stigmas. Achene 0.5-0.7 mm long, broadly obovoid-trigonous, the faces nearly plane, the adaxial face broadest, the surfaces fine- ly papillose-lined longitudinally, transversely ru- gulose, whitish gray to brown; tubercle ca. 0.1 mm long, subglobose to oblate, apiculate, dark brown. Paratypes. COLOMBIA. GUAINÍA: near Coitara, ca. 7 km S of San Fernando, 67?43'W, 3°55'N, Davidse 16827 (COL, MO, VDB, VEN); Finca Buena Vista on the Río Negro, ca. 15 km S of San Felipe and San Carlos de Río Negro, 67°00'W, 1°42'N, Liesner 8780 (COL, MO, Betun along the Río C ; Davidse & González 13121 (MO, VEN); 27 km WSW of Paso de Cinaruco along the Río Cinaruco, 67°45'W, 6°31'N, Davidse & González 12574 (MO, VEN); bank of Río Orinoco on Isla Poyatón, 67°05'W, 7°02'N, vidse & González 122164 (MO, VEN); 9 km W of Paso ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 438 SOD Osco o O6o0cG6o00o090 ad 0005500 009.29 2203 099 O aoa - v. ay ` Se iw ES 1987] de Cinaruco along the Río Cinaruco, 67?35'W, 6?35'N, Davidse & González 12495 (MO, VEN). BOLIVAR: Au- yan- tepui, coche caps n. (VEN). BRAZIL. MATO GROSSO: Rio Aripua Humboldt Campus, 59?21'N, 10°12’S, d et al. 18319 (MO, NY). Among the tropical American species of Bul- bostylis, B. fluviatilis seems most closely related to B. tenuifolia (Rudge) Macbride, and some of the cited paratypes have been distributed under that name. Bulbostylis fluviatilis is similar to B. tenuifolia in its general facies and annual life cycle but differs in having one-nerved glumes with an excurrent mucro and two stamens with anthers 0.2-0.3 mm long. Bulbostylis tenuifolia has three- nerved glumes with the midrib slightly or not at all excurrent and one stamen with anthers 0.4— 0.5 mm long. In superficial appearance B. fluviatilis also re- sembles some species of the Trichelostylis group of Fimbristylis. However, the prominent hairs along the mouth of the sheath and the distinctly tuberculate style base clearly indicate that this species belongs in Bulbostylis. Bulbostylis fluviatilis grows along large rivers (hence the epithet) at elevations of 65-120 m. It grows primarily on the sand bars that become prominently exposed in these rivers during the dry season and on accumulations of sand on ex- posed granite outcrops along the rivers. NOTES 439 We believe that the cited collection of Vareschi from Auyan-tepui is not a reliable record and that it is probable that the wrong label was mounted with the plant. It is perhaps possible that the plant originated along one of the rivers at the base of Auyan-tepui, but it is extremely unlikely that it originated from the highly dif- ferent habitats atop Auyan-tepui. The Brazilian collection differs from the others in having the leaves about as long as the inflorescences. The fieldwork that resulted in some ofthe cited collections was supported by the National Sci- ence Foundation, National Geographic Society, Consejo Nacional de Investigaciones Científicas y Tecnológicas (Venezuela), and Fundación para el Desarrollo de las Ciencias Físicas, Matemá- ticas y Naturales (Caracas). We thank Otto Hu- ber and Charles Brewer-Carías for providing op- portunities for fieldwork in southern Venezuela, and the Herbario Nacional (VEN), INP- ARQUES, for use of their facilities. — Robert Kral, Department of General Biology, Vanderbilt University, Nashville, Tennessee 37203, U.S.A.; and Gerrit Davidse, Missouri Bo- tanical Garden, P.O. Box 299, St. Louis, Mis- souri 63166, U.S.A. — FIGURE 1. sketch. — b. Leaf blade apex.—c. Outer view of the leaf base. — f. Anthella base, showin Bulbostylis fluviatilis Kral & Davidse, drawn from the type Epsum. Kral 71973.—a. Habit id-sector of the leaf blade, dorsal view. —d. I view of the leaf base.— e. t a al and bases of primary ruit rays. —g. Sector of the scape.—h. Fertile bract.—i. Two sorts of spikelets. —j. Floret.—k. A NEW CAREX SECT. OLIGOCARPAE (CYPERACEAE) FROM WESTERN ARKANSAS AND EASTERN OKLAHOMA In late April of 1977, during fieldwork along the summit of Rich Mountain, the highest range of the Ouachita trend in western Arkansas, the senior author discovered on the dry, oak-pine slopes of the southwestern-facing side numerous clones of a Carex similar to C. hitchcockiana Dewey but distinct in habit, indument, and foliar character. On being informed of this plant, Ar- ductive searches for it, not only in the original locality but elsewhere on Rich Mountain (in- cluding Oklahoma) and southward. We are aware now of an abundance of it, often in association with C. hitchcockiana and C. oligocarpa Schk. of sect. Oligocarpae over a substantial range of arenaceous oak-hickory-pine uplands in Arkan- sas (Howard, Polk, Scott counties) as well as in LeFlore County, Oklahoma. This habitat system is so well represented in the Interior Highlands physiographic province, particularly the Ouach- itas, that a sedge thought to be rare and local may indeed be widespread and locally abundant with- in that physiography, as has now been shown in the case of C. /atebracteata Waterf. (Kral, 1983). The new Carex is named after the mountain complex where it first was found. Carex ouachitana Kral, Manhart & Bryson, sp. nov. TYPE: United States. Arkansas: Polk Co.: sandy rocky woods at summit of Rich Mountain, just W of Queen Wilhelmina Park Headquarters, 26 Apr. 1977, R. Kral 59803 (holotype, MO; isotypes, BM, CTB, GA, GH. MICH, MO, US, VDB). Figure 1. Spec. nova e sectione Oligocarpae. Planta perennis, 30-80 cm a glabra, laxe cespitosa, valde squam ate rhizomat s vel paulo brevioribus, Rhizoma repens vel ascendens, plus -5 c nga acuta vel acuminata scopu rpurea, carinata. Culmi dimorphi, lateralibus Stenilibus foliosissimis. Folia principalia 30-70 cm longa. Laminae compressae, 3- 5 mm latae, scabridae, apicem versus attenuatae, sca- bromarginatae; pagina ad medi ae, ae eeu ascendentibus vel erectis, 2-6 mm lon- sis subtendentes, infimae laminis 10-13 cm longis et m erecto rostro ca. 4.5-5.0 mm longa duds concava, pallide brunneola, minute papillosa Perennial 30-80 cm long, smooth, loosely ces- pitose, strongly scaly-rhizomatous, the principal leaves slightly longer or slightly shorter than the culms. Rhizome creeping to ascending, + lig- neous, 2-4 mm thick, branching, the scales ovate, acute, spirally imbricate, multicostate, brown, gradually passing into cataphylls. Cataphylls ob- long, 1-5 cm long, acute or acuminate, red-brown- purple, multicostate (costas often white), cari- nate. Culms dimorphic, spreading, slender, acutely trigonous, scabrid, the lateral ones sterile, more leafy. Principal foliage leaves longer toward culm base and approximate, 30-70 cm long. Leaf sheaths carinate, smooth; ligule erect, a narrow, horseshoe-shaped or acute scale. Blades spread- ing or recurved, flattened, linear, 3-5 mm wide, scabridulous, apically attenuate, triquetrous, the edges harsh, the surfaces at mid-blade above sca- brellous, impressed-nerved, abaxially elevated- nerved, with midcosta sirongly raised, scabrid. Spikes 2-4, narrow, the uppermost all male, nar- GURE l. j. Carex ouachitana. —a. Hab c. quies rare side. — d. Axial (left) and abaxial (right) sides of midsector of leaf blade. — — b. Ventral (left) and lateral-oblique view of leaf sheath apex ssa ~ staminafe flower.—f. Anther and upper part of filam —g. Perigynium and accompanying scale, spike . Peri- gynium and accompanying scale, spike apex, pomi stigmatic branches and m of included fruit. — i. Detail, further anc of perigynium.—j. Fruit. From Kral 59803. ANN. Missouni Bor. GARD. 74: 440-442. 1987. 441 NOTES 1987] “scu ti ESE z5 “ease ass nee 442 rowly densely ellipsoid-cylindric, 2-4 cm long, ca. 3 mm thick, 2-3 times shorter than the pe- duncle. Spikes beneath all female, or androgy- nous with 1-7 male florets at the tip, narrowly linear, few-to-many-flowered, 3-6 cm long, ba- sally interrupted, at apex with the flowers more approximate. Male spike bract without sheath, lanceolate, 10-15 mm long, narrowly acuminate or cuspidate, carinate; bracts of lateral spikes fo- liaceous, erect, subtending erect to ascending pe- duncles 2-6 cm long, the lower with blades 10- 15 cm long and with closed sheaths to 4 cm long. Male scales scarious, oblong-lanceolate, ca. 2.5- 3.0 mm long, acute, carinate, the sides pale brown, the costal area green. Female scales ovate to lan- ceolate, navicular, 5-15 mm long including the cusp, acute to narrowly truncate, the midzone green, unicostate, the sides stramineous to brown, the tips of the lower scales strongly cuspidate. Perigynia obovoid, 4-6 mm long, obscurely tri- gonous, the faces slightly concave to level, im- pressed-nerved, the beak short, excurved. Fruit tightly included, stipitate-obovoid, trigonous, 4.5-5.0 mm long including the short erect beak, the faces concave, pale brown, minutely papil- ate. Additional specimens examined. UNITED STATES. ARKANSAS: Howard Co.: shaley wooded river bluffs, 9.6 mi. E of Wickes, 9 May 1979, Kral 63507; NW of AR Hwy. 4 and Cossatot River on slopes above river under mixed hardwoods, 10 May 1986, Bryson 4332. Polk Co.: Rich Mtn., along Ouachita Trail near Hwy. 88 N 0.3 mi. W of Rich Mtn. tower, 27 Apr. 1981, Rettig & Davis 239; Queen Wilhelmina State Park, Sect. 11, R32W, TIS, below and to N of visitor center and AR Hwy. 88 along trail on N-facing slope under mixed hardwood forest, rich rocky soil, 10 May 1986, Bryson 4289; NW of Rich Mtn. Lookout Tower, N of AR Hwy. 88, SW '4 Sec. 8, R31W, TIS, along Ouachita Trail on N-facing slope under hardwood forest with dense herbaceous undergrowth, 10 May 1986, Bryson 4290; NE of AR Hwy. 246 and Cossatot River crossing NE ‘4 Sec. te R30W, T4S peal pos hardwood forest with few pines, on W-facing slope cky soil with thick leaf E "10 May 1986, Do: 4326. Scott Co.: N of Men slopes of Black Fork a Oklahoma-Arkansas state line near a #1, 30 Aug 1979, Taylor 28122; just W of Sta Line Historical nds on roc hickory forest with few eg toward top, thick brushy and herbaceous unders rich rocky soil in associ- ation with Carex prede € : posses C. hitchcock- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 iana, C. albursina, 10 May 1986, Bryson 4310. Du- plicates are to be distributed later, thus acronyms for recipients are not cited. Staminate terminal spikes, female spike scales with harsh cusps and costae, perigynia with many impressed nerves, and short toothless beaks place this sedge squarely in sect. Oligocarpae, a section first designated by Carey in Gray (1848), and subsequently revised by Kukenthal (1909), Mac- kenzie (1931-1935) and Fernald (1950). In most modern classifications this complex comprises just C. oligocarpa and C. hitchcockiana. Both of these are densely cespitose and erhizomatous; whereas C. ouachitana is loosely cespitose, sometimes forming colonies up to 0.5 m across in open areas and up to 1—2 m across in shaded areas, and produces strong, ligneous, imbricate- scaly rhizomes. The staminate spike is larger, with a longer peduncle, thicker, and bears more florets. The lateral spikes are frequently androg- ynous, while in C. oligocarpa and C. hitchcock- iana they are solely carpellate. The perigynia are much like those of C. hitchcockiana in size and shape but differ in that the beak is shorter, broad- er, and more bent outward. On the other hand, the fruit beak resembles that of C. oligocarpa by being erect or suberect. However the members of this triad are related morphologically, it is significant that, even though they have been ob- served to share habitat in the same localities, no apparent naturally occurring hybrids have thus far been seen. LITERATURE CITED iae M. L. 1950. Gray's eium i Botany, 8th tion. American Book Com , New York, n A Manual of the Botany ofthe North- ern United States. James Munroe & Co., Cam ndon. . Carex eda eii lied Pp. 108- 112 in A report o threatened, or en- dangered forest- ated ae sae of the south. Volume I. USDA Forest Service Technical Publ. R8-TP 2. Atlanta, Georgia. KUKENTHAL, G. 1909. Cyperaceae—Caricoideae. 1931-1935. Cariceae. InN. Amer. Fl. 18(7): 1-478. — Robert Kral, Department of Biology, Vander- bilt University, Nashville, Tennessee 37235, U.S.A.; James Manhart, Division of Biological Sciences, The University of Michigan, Ann Arbor, Michigan 48109, U.S.A.; and Charles T. Bryson, Southern Weed Science Laboratory, P.O. Box 350, Stoneville, Mississippi 38776, U.S.A NOTES ON CHROMOSOME CYTOLOGY OF RUTACEAE— DIOSMEAE The tribe Diosmeae R. Br. emend. Benth. & Hook. f. (1862) of the Rutaceae is strictly African in distribution and consists of the widespread tree Calodendrum capense (L.f.) Thunb. (south- ern Cape to north Kenya) and some 280 species of small to medium-sized shrubs, mostly of er- icoid habit. The tribe is, with the exception of Calodendrum, restricted to southern Africa and largely to the Cape Region of the south and west coast of South Africa. Taxonomic revisions of the tribe have recently been published for all but the large genus Agathosma (Strid, 1972; Wil- liams, 1975, 1978, 1981a, 1981b, 1982) but chromosome cytology of the group is poorly known. Apart from several counts by Strid (1972) for Adenandra, there are only scattered reports for a few genera. The monotypic Calodendrum is a high paleopolyploid with 2n = 54 (Honsell, 1954; Smith-White, 1954). In Agathosma, A. crenulata (L.) Pill. has been reported as having 2n = 45 (as Barosma) (Riley & Hoff, 1961), while Guerra (1984a) found 2n = 26 in A. apiculata E. Meyer (also as Barosma) and A. lanceolata (L.) Engl. There are two counts for Coleonema pulchel- lum I. J. Williams (as C. pulchrum Hook.), Smith- White (1954) reporting 27 — 36 and Guerra (1984a) 2n = 34; Guerra also reported 27n = 34 in the closely related C. album (Thunb.) Bartling & Wendl. Numbers reported in Adenandra are 2n — 28 in two species, 38 in one more, 42 or ca. 42 in two species including A. fragrans, and 2n — 48-50 in another three. In conjunction with continuing systematic studies of Diosmeae a pre- liminary investigation of the chromosomes of some genera was undertaken and this report is the result of the study. All counts were made from mitotic metaphase in root tip squashes of germinating seeds, fol- lowing a method described fully elsewhere (Goldblatt, 1979, 1980). Chromosome number and voucher information are as follows: Diosma subulata Wendl. 2n — 30. South Af- rica, Cape, ex hort. Williams, Williams s.n. (NBG); Wortelgat, Caledon Div., Williams 1721 (NBG). Diosma aristata I. J. Williams 2n = 30. South Africa, Cape, ex hort. Williams, Williams s.n. (NBG) ANN. Missouni Bor. GARD. 74: 443-444. 1987. Diosma oppositifolia L. 2n = 30. South Africa, Cape, Vogelgat, Caledon Div., Williams 2406 (NBG). Euchaetes avisylvana I. J. Williams 2n = 28. South Africa, Cape, Grootvadersbos, Heidelberg Div., Williams 2377 (NBG). Adenandra frag (Sims) Roemer & Schultes 2n = 42. South Africa, Cape, Grootvadersbos, Heidelberg Div., Williams 2378 (NBG). DISCUSSION The count of 2n = 42 for Adenandra fragrans confirms the earlier report by Strid (1972) for this species. Reports here for Euchaetes and Diosma are the first records for these genera. On the basis of four counts for three species of Dios- ma, all 2n = 30, we suggest that x = 15 is basic for the genus. In Euchaetes the single count sug- gests a generic base of x = 14; x = 7 seems un- likely since all other counts in Diosmeae are at paleotetraploid or paleohexaploid levels. Basic chromosome number for Rutaceae is most likely x = 9 as suggested by Smith-White (1954). This hypothesis is founded on the pre- dominance of n = 18 and 36 in the unspecialized Xanthoxyleae and Flindersieae, and Ehrendorfer (1981, 1987) has also pointed out the occurrence of only n = 9 and 18 in Aurantieae. Data for Diosmeae throw no direct light on the possible base number for the family, as they are relatively specialized. However, the number in Ca/oden- drum, n = 27,is most likely paleohexaploid based on x = 9, and the same base number is by ex- tension probable for all Diosmeae, given the bas- al position of Calodendrum in the tribe. Other genera of Diosmeae that have been counted ap- pear fundamentally paleotetraploid, a view en- dorsed by Guerra (1984b) for Coleonema based on a comparison of genome size in Rutaceae. Thus Coleonema, either n = 18 or 17, is probably hypotetraploid based on x = 9. The base of x = 15 in Diosma may represent either further aneu- ploid decrease from an ancestral x — 18, or less likely, allopolyploidy from hypothetical ances- tors with x = 7 and 8. Euchaetes, closely related to Diosma, with n = 14 fits with the apparent trend for decrease from an ancestral tetraploid base. Adenandra appears basically paleotetra- ploid (? x = 14) but some species may be octo- 444 ploid and derived from the secondary hypotet- raploid base. This may best explain the range from n = 25, 24, 21, and 19, as well as n = 14 in two species. Further counts in Diosmeae are needed to re- solve the several questions that this initial study has posed. The variation in chromosome num- ber in the tribe is unexpected and is likely to be of considerable value in future studies of generic limits and of the relationships of genera and species, and ultimately of phylogeny. LITERATURE CITED BENTHAM, G. & J. D. HOOKER. 1862. Rutaceae. In enera denies 1: 278-306. EHRENDORFER, Speciation Mpeg in woody angiosperms roi l origin. Pp. 479—509 in C. Barigozzi (editor), Mechanisms of Speciation. Alan Liss, New York. 87. Affinities of the African dendroflora: suggestions from karyo- and chemosystematics. onogr. Syst. Bot. Missouri Bot. Gard. (in press). GOLDBLATT, P. 1979. Chromosome cytology and hien Pd in Galaxia (Iridaceae). Pl. Syst. Ev 133: 61—69. Redefinition of Homeria and Moraea ue in the light of pb: data, with me nov. Bot. N 5. GUERRA, M. pos S 1984a. New chromosome num- ers in Rutaceae. Pl. Syst. Evol. 146: 13-30. . Cytogenetics of Rutaceae II. Nuclear DNA content. Caryologia 37: 219-226. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 HoNSELL, E. 1954. Osservationi sulla struttura dell'ovulo e sulla cariologio di Calodendron ca- pense Thunb. e Pilocarpus pennatifolius. Ann. Bot. (Rome) 24: 438-447. RirEv, H. P. & V. J. Horr. 1961. Chromosome stud- ies in some South African dicotyledons. Canad. J. Genet. Cytol. 3: 260-271. SMITH- WHITE, S. 1954. Chromosome numbers in the luni mid development of the tribe in rem Austra: ustral. J. Bot. 2: 287-303. . Revision of the genus Adenandra (Ru ta ceae). Opera Bot. 32: 1-112. WILLIAMS, ION. 1975. Studies in the genera of Dios- meae (Rutaceae): 5. J. S. African Bot. 41: 180- i 978. Studies in the genera of Diosmeae (Ru- 23 981a. Studies in the genera of Diosmeae (Rutaceae): 9. A revision of the genus Coleonema. J. S. African Bot. 47: 63-102. 1981b. Studies in the genera of Diosm (Rutaceae): 10. A review of the genus E riii J. S. African Bot. 47: 157-193. 1982. Studies in the genera of Diosmeae (Ru- tacea e): 14. A review of the genus Diosma. J. S. African Bot. 48: 329-407. — Peter Goldblatt, B. A. Krukoff Curator of Af- rican Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.; and Ion Williams, 29 Tenth St., Voelklip, Hermanus 7203, South Africa. THE FRUITS OF DECARYDENDRON (MONIMIACEAE) Decarydendron Danguy is a poorly known ge- nus of Monimiaceae (subfamily Mollinedioi- deae, tribe Hedycareae) comprising three species endemic to Madagascar (Danguy, 1928; Cavaco, 1958a, 1958b; Lorence, 1985). Fruiting material of the genus, hitherto unknown, is described here for the first time based on a specimen of D. per- rieri Cavaco recently collected by L. J. Dorr and L. Barnett in Madagascar. The following descrip- tion is based on FAA-fixed fruits of Dorr & Bar- nett 3203 (Madagascar. Province Tamatave: en- virons d'Andasibe-Périnet, 18?56'S, 48?25'E, 2- 4 Nov. 1984). Voucher specimens have been de- posited at MO and TAN, and the fixed collection is at PTBG. Fruiting receptacle attached at or below ground level, terminal on a finely velutinous peduncle 2.5-3 mm diam., swollen, fleshy, obconical-tur- binate, initially 3 cm diam., during development curling outwards and splitting irregularly into 3 partly everted segments, externally and inter- carpels 12, numerous aborted carpels. Mature carpels sessile to sub- sessile, obpyriform, 2-2.5 cm long including the constricted, curved, beak-like apex 5-7 mm long, 1.3-1.6 cm diam., externally yellowish brown, corky, bearing circular to elliptic corky lenticels S. 1 1cm Ld T faber with attached carpels.— ANN. MissouRi Bor. GARD. 74: 445-446. 1987. over most of the surface; exocarp fleshy, to ] mm thick (thicker at apex), the surface corky, par- enchymatous, the tissue replete with densely ag- gregated brachysclereids, underlain by a vascu- larized zone, the mesocarp parenchymatous with scattered + cuboidal idioblasts; endocarp hard, white, + smooth, discontinuous apically at the micropyle, 0.8-1 mm thick, co densely packed, radially oriented layer of narrow, thick-walled, fusiform columnar sclereids; testa 0.1-0.2 mm thick, brown, composed of 2 layers of tangentially elongated cells with slightly thick- ened walls, underlain by an endotesta many lay- ers thick of short, rounded tracheids with sca- ariform thickenings; tegumen 3-4 layers thick, composed of elongate thin-walled cells. Endo- sperm about 1 cm diam., white, copious, oily, interspersed with crystals. Embryo situated api- cally in the endosperm below the micropyle, cy- lindrical-clavate, 2-3 mm long, | mm diam. dis- tally, the cotyledons erect, + appressed, about half the total embryo length. Figure la, b. DISCUSSION In a recent monographic treatment of the DANIEL nein Fruiting receptacle and carpels of Decarydendron perrieri. —a. b. Carpel, longitudinal section. Based on Dorr & Barnett 3203 (PTB Mature, irregularly split receptacle G). 446 any of the Malagasy genera: both androecious and gynoecious flowers with shallowly concave receptacles bearing 1—2 whorls of large, imbricate tepals; flowers that unfold gradually at anthesis without splitting into segments as in the other three genera (i.e., Ephippiandra Decne., Monim- ia Thouars, and TambourissaS flowers with numerous (ca. 300—1,000) free, sub- sessile, clavate carpels. As Decarydendron closely resembles Hedycarya Forster & Forster f. from Oceania and various Pacific islands in terms of gynoecious floral morphology, it was postulated that mature fruits of the former genus would most likely resemble those of the latter one (Lorence, 1985 ) gynoecious The findings reported here confirm that ma- ture fruits of Decarydendron perrieri do strongly resemble those of other genera in the tribe He- dycareae (Philipson, unpublished) and support its placement there. Members of this tribe are characterized by gynoecious flowers with a non- calyptrate floral cup that encloses few to many free carpels, and fleshy discoidal or cupuliform fruiting receptacles that gradually split or become everted as the carpels mature. Among the other Malagasy genera belonging to the Hedycareae, the free, sessile to subsessile carpels of Decary- dendron are less specialized than those of Ephip- piandra, which are also free but partly immersed in cupules formed by the receptacle. Fruits of both these genera are in turn less specialized than the inferior, syncarpous carpels of Tambourissa, which are completely immersed in and united with the ovary wall (Lorence, 1985). Thus, fruit morphology also supports the suggestion that the ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 three Malagasy genera of Mollinedioideae form part of an increasingly specialized evolutionary series. Among the extra-Malagasy members of the tribe Hedycareae of the subfamily Mollinedioi- deae, Decarydendron is most closely allied to Levieria Becc. from Malesia and Australia, and particularly to Hedycarya, in terms of floral and fruit morphology. Decarydendron differs from both these genera in being monoecious, in having sexually mixed cauliflorous inflorescences, and in having gynoecious flowers with more numer- ous clavate carpels having the stylar canal situ- ated in the basal half of the carpel. I thank L. Dorr of the Missouri Botanical Gar- den for providing fixed material of Decaryden- dron perrieri and Danial Cobian of the Instituto de Biología for the illustration. LITERATURE CITED CAVACO, A. Sur le genre Decarydendron (Monimiacées). ips Soc. Bot. France 105: 38-39 195 espéce nouvelle de Decaryden- ontribution à l'étude des Moni- cées de Madagascar. Bull. Mus. Hist. Nat. (Paris) ae 278-280. Lor 5. A monograph of the Monim- iaceae a in the Malagasy Region segera st Indian Occan). Ann. Missouri Bot. Gard. 7 1-16 PHILIPSON, w. R. A m of the Monimiaceae (unpublished manuscript). — David H. Lorence, Pacific Tropical Botanical Garden, P.O. Box 340, Lawai, Kauai, Hawaii 96765, U.S.A RECONSIDERATION OF THE GENERIC PLACEMENT OF PALICOUREA DOMINGENSIS (RUBIACEAE: PSYCHOTRIEAE) Although the Caribbean species sometimes known as “Palicourea domingensis” has com- monly been treated in Palicourea, it lacks the distinguishing features of this genus and is better placed in Psychotria. Palicourea Aublet (Rubiaceae: Psychotrieae) is distinguished within its tribe by comparatively long (tubes 5—40 mm long), tubular corollas that are -— us elation! gwalen at the base with aringoftri abgve the basal swelling. Characteristically, the ally and bright ly colored, usually red, yellow, blue, or purple. The thyrsiform inflorescences are typically open, with well-developed branches, bracts, and ped- icels. This genus has been loosely circumscribed by many authors, who have used variously one ora few of these features or characters to separate it from Psychotria The species of Psychotrieae now usually called “Palicourea domingensis” (Jacq.) DC. is a shrub or small tree of moist forest on soils derived from limestone. It is distributed through most of the Caribbean islands (Cuba, Jamaica, Hispaniola, Puerto Rico, St. Thomas, St. Croix, Tortola, Antigua, St. Kitts, St. Eustatius, Guadeloupe, Nicaragua, although it has not been previously reported from these areas (Standley & Williams, 1975). This species is characterized by salverform co- rollas with tubes 5-15 mm long. The corollas are predominantly white, although often tinged with rose or purple. Their tubes are glabrous exter- nally and internally, and are not at all swollen or asymmetrical at the base, although the tube may be curved somewhat through its length. The inflorescence bracts are poorly developed or lacking. Thus, “Palicourea domingensis” lacks the dis- tinctive floral features and the common inflo- rescence characters of Palicourea. It has been placed in Palicourea by most authors apparently based on its relatively long, tubular, often rosy corollas, combined with an inconsistent circum- ANN. MissouRi Bor. GARD. 74: 447—448. 1987. scription of Palicourea and a respect for previous practice. This species is better placed in Psychotria subg. Heteropsychotria Steyerm., which is closely re- lated to Palicourea. The floral features of Psy- chotria domingensis are similar to those of other species of Psychotria, and the inflorescence char- acters described above that are rare or lacking in Palicourea are common conditions in this sub- genus. This relationship was recognized previ- ously by Standley when he described this same species from Nicaragua under the synonym Psy- chotria mombachensis Standley. Although the corollas of Psychotria are described in most treat- ments as comparatively short, and typically do have tubes 5 mm long or shorter, long corollas occur in the genus as well, as in Psychotria chia- pensis Standley which has corollas with tubes 30- 50 mm long. In fact, within subg. Heteropsy- chotria, Psychotria domingensis 1s similar in sev- eral characters to Psychotria chiapensis and to Psychotria gardenioides (Scheidw.) Standley: these species share glomerulate, subsessile flow- or rosy corollas with internally glabrous tubes 20 mm long or longer. (The last two species differ from Psychotria domingensis in their longer co- rolla tubes and well-developed inflorescence bracts.) efore, “Palicourea domingensis” seems better classified in Psychotria. In this case, the correct name of this species is Psychotria dom- ingensis Jacq. A complete list of synonymy and citations of specimens from Central America are presented below. NOMENCLATURE Psychotria domingensis Jacq., Enum. Pl. Carib. 60. Palicourea domingensis (Jacq.) DC., Prodr. 4: 529. 1830. TYPE: Santo Do- mingo. Hk pavetta d) Prodr. 45. 1788. Pavetta pen- andra Sw. . Occ. 1: 233. 1797. Palicourea And (Sw) DC. Prodr. 4: 525. 1830. Palicourea pentandra (Sw.) K. Schum., Nat. Pflanzenfam. Bis 115. 1891, nom. illeg.. Art. 63. TYPE: Ja- otographs A, N aica ( NY). m ones tabernaefolia Poir., Encycl. 5: 704. 1804. 448 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FIGURE 1. Palicourea ig sib Lene Ad ) DC., Prodr. 4: 525. 0. TYPE: Santo ingo. Psychotria angustifolia Poir. ps 5: 703. 1804. TYPE: anto Domingo. —— mombachensis Standl., Mus., Bot. Ser. 8: 188. 1930. TYPE: Nicara- ie coffee plantation, Mombacho Volcano, 600- 750 m, Maxon, Harvey & Valentine 7818 (holo- type, F) Specimens examined from Mexico and Central America. co. TABASCO: La Palma, Balancan, Matuda 3236 (NY). GUATEMALA. PETEN: Tikal, Contreras RG 1413 (TEFH); Dolores, Contreras 2503 O); Macanche, Contreras 5847 (DS); Chincila, pede ease 6360 (MO); Isla del Armadillo, Laguna Macanche, Contreras 7 1 A (MO); Sayaxche Road, km 59, Contreras 7247 (M TEFH); Guayacuan, La Pita, Contreras 7477 MOS Tikal, Contreras 7707 (MO); between Paso Caballo and Carmelita, Cox 2835 (EAP); Tikal, Molina 15750 ) Publ. Field Colom- BELIZE. CAYO: vicinity of La Flor at Río de La Flor, 6 mi. S of San Grano de Oro, 1,700-2,700 ft. [520- 820 m], Croat 23739 (MO), Dwyer 10876 (MO); in advanced forest, Lundell 6199 (NY); W of Humming- bird Hwy. 7 mi. S of junction with Western Hwy., N Distribution of Psychotria domingensis Jacq. in Mexico, Central America, and the West Indies. boundary of Roaring River Estate, 80 m, Spellman & 1915 (MO); 1,700 yd. W of Hummingbird Hwy., 7 mi. of junction with Western Hwy., 80 m, Spellman & Newey 2010 (MO); TOLEDO: Dwyer 9863 (MO I thank the curators of the following herbaria, who very kindly made specimens available for study: A, CAS, DUKE, EAP, F, GH, M, MO, NY, TEFH, and US; I thank in particular Dr. James Ackerman of UPRRP, who provided working space. I also thank Orlando Matos Con- cepción for help in preparing the distribution ata LITERATURE CITED STANDLEY, P. C. & L. O. WiLLIAMs. 1975. Rubiaceae. In Flora of Guatemala, Part XI. Fieldiana Bot. 24(11): 1-274 — Charlotte M. Taylor, Department of Biology, Universidad de Puerto Rico— Colegio Universi- tario de Cayey, Cayey, Puerto Rico 00633 CLARIFICATION OF THE TAXONOMY OF LEUCAENA SALVADORENSIS STANDLEY EX BRITTON & ROSE The nomen nudum Leucaena salvadorensis Standley was first published by him in Standley & Calderón (1925) in a list of El Salvador plants, without a complete description or designation of type. In their revision of the genus, Britton & Rose (1928) supplied a description and desig- nated a type, giving the authority as Standley. Later, Standley & pue uem (1946) reduced the species, cited as L. salvadorensis Standley ex Britton & Rose, to synonymy under L. shannonii J. D. Smith. Prewbaker et al. (1972) and Brew- baker in a mimeographed leaflet (1978) placed L. salvadorensis under the synonymy of L. leu- cocephala (Lam.) de Wit, and more recently Brewbaker & Ito (1980) treated it as a subspecies of L. leucocephala (also see Brewbaker’s mim- eographed revision, 1984). Zarate (1982, 1985) concluded that it was the same as the small-leaf- let form of L. shannonii and returned it to syn- onymy with that species. Recently, Colin Hughes, Brian Styles, and An- gela Laguna have collected material in Honduras and Nicaragua (C. Hughes 332, 334, 446 (FHO, MEXU); C. Hughes & B. Styles 37, 102, 129 (FHO, MEXU); A. Laguna 427 (HNMN, MEXU)) referable to this taxon. From study of these specimens it became apparent that it de- serves recognition as a subspecies of L. shan- nonii. The dome-shaped or truncate-conic, oblong petiolar glands and the corolla-to-calyx length ratio of ca. 0.7 in L. shannonii and L. salvador- ensis clearly distinguish these taxa from L. /eu- cocephala with suborbicular or elliptic or ob- ovate thick-bordered and furrowed to concave glands and a corolla-to-calyx length ratio of ca. 0.5. The more numerous pairs of pinnae and leaflets, 6-11 and 23-35, respectively; consis- tently narrow oblong leaflets, ca. 1.5 cm long; florets 4 mm long; and pods strongly stiped, 1 1- 21 cm long and ca. 2.5 cm wide, separate it from the typical element of L. shannonii with (2—)4— ANN. MISSOURI Bor. GARD. 74: 449. 1987. 6(-8) pairs of pinnae, 8—22 leaflet pairs, very variable leaflets (0.8—)1—2 cm long, florets 3-3.5 mm long, and slender-stiped pods ca. 15 cm long and 1.4-2.2 cm wide. Therefore, the following combination is made. Leucaena shannonii J. D. Smith subsp. salva- dorensis (Standley ex Britton & Rose) S. Zá- rate, comb. et stat. nov. Basionym: L. sal- vadorensis Standley ex Britton & Rose, Fl. N. Amer. 23(2): 125. 1928. TYPE: El Sal- vador. Morazán: Jocoro, Calderón 2031 esum NY). LITERATURE CITED BREWBAKER, J. L. 1978. Guide to the systematics of the genus Leucaena (Mimosaceae). C.I.A.T. Cali. dp h. . 1984. Revision ofthe genus Leucaena (Mim- osoideae: Leguminosae). Honolulu. Mimeograph. TO. 1980. Taxonomic studies of the . 1972. Va- rietal variation and yield trials of Leucaena leu- cocephala (Koa pik in Hawaii. Haw. Agric. Exp. Sta. Res. Bull. 166: BRITTON, N. L. & J. N. Rost. N. Amer. 23(2): 12 PE si as PC & S. abu 1925. Lista Preli- minar de Plantas de El Salvador. San Salvador. . STEYERMARK. 1946. Flora of Guatemala. denar Bot. 24(5): 47-48. TE, P. S. 1982. Las especies de Leucaena Benth. de Oaxaca con notas sobre la sistemática del gé- nero para México. Thesis. Facultad de Ciencias, , México. 1984 [1985]. Taxonomic revision of the ge- nus Leucaena Benth. from México. Bull. IGSM 12: 24-34 1928. Mimosaceae. Fl. —Sergio Zárate Pedroche, Instituto de Biología, Herbario Nacional de México (MEXU), U.N.A.M., México. Presently graduate student supported by CONACYT (Reg. No. 49993), De- partment of Biology, Indiana University, Bloom- ington, Indiana 47401, U.S.A. AN UNUSUAL NEW SPECIES OF HELMIOPSIS H. PERRIER (STERCULIACEAE) FROM MADAGASCAR Eight species of He/miopsis H. Perrier (Ster- culiaceae) currently are recognized in the Flore de Madagascar et des Comores (Arénes, 1959). All are characterized by monodelphous androe- cia with staminodes opposite the petals, sessile ovaries, caducous petals, capsular fruits with api- cally winged seeds, and overall vestitures of pel- tate or fimbriate (i.e., peltate with fimbriate mar- gins) scales. All species described to date are restricted to western Madagascar. A new species of Helmiopsis is described here from the north- ern mountains of Madagascar, also part of the phytogeographical Domaine de l'Ouest of the is- land (Humbert, 1965). This species combines several features that are unusual for the genus. For example, its spheroid, rather than conical, capsule morphology is unique and provides the basis for the name He/miopsis sphaerocarpa L. Barnett. The seed wings of H. sphaerocarpa are atypical for the genus in being reduced to only a thin, ventral keel; nnn E angs of Hel- e, to 10 mm in length. The inflorescence i IS ssa in struc- ture, being multibranched and many-flowered, as opposed to the few-flowered cymose inflores- cences more commonly found in the genus. The terminal nature of the inflorescence is also un- usual in this species, although terminal inflores- cences also have been observed in isolated col- lections of H. pseudopopulus (Baillon) Capuron (e.g., Capuron 24661, P). Finally, this new species is unique in bearing glandular tissue both on the calyx lobes and on the petals; in all other Hel- miopsis species with glandular tissue, the tissue is restricted to either the calyx or the corolla, but is never found in both perianth whorls. Helmiopsis sphaerocarpa L. Barnett, sp. nov. TYPE: Madagascar: Centre (Nord) jusqu’aux confins de l'Ouest (Nord), Massif de la Mon- tagne d'Ambre, créte entre les bassins de la riviére des Makis et de la riviére d'Anka- zobe, entre 800 et 600 m d'alt., 26-27 May 1970 (fl), Capuron 29194 SF (holotype, P; isotype, TEF). Figure 1 rbuscula vel arbor, squamarum lepidotarum de- nales, 15-30 flor bis intus Mn duce Petala 5, E glandulosa. Tubus ANN. Missouni Bor. GARD. 74: 450-452. 1987. staminalis | mm altus; staminibus 10, staminodiis 5. Ovarium 5-loculare, loculis 2-ovulatis; stylo glabrato, stigmate leviter 5-lobata. Capsula didi loculici- da. Semen carina ventrale membranace Large shrub or tree to 8-12 m tall, branches terete, finely striate, new growth sparsely peltate- scaled, older twigs glabrate. Leaves alternate, de- ciduous, blades broadly ovate to narrowly ob- ovate, 6.5-12 cm long, 4—9 cm wide, apex acute to acuminate, base shallowly cordate to cordate, palmately 5(-7) nerved, midvein and secondary veins conspicuous, raised below and slightly de- pressed above, discolorous, margins irregularly crenulate, lower surface with scattered peltate and fimbriate scales, becoming glabrate; petioles 1- 6 cm long, with scattered peltate and fimbriate scales. Inflorescences terminal and in the axils of uppermost 3-5 leaves, determinate, branching 3-5 times, bearing 15-30 flowers. Floral buds ovoid, 4-5 mm long, 4-4.5 mm wide. Calyx 5-lobed, lobes connate only at the bases, lanceo- late, ca. 7 mm long, 3 mm wide, outer surfaces densely covered with peltate scales, inner sur- faces glabrate and with an arc of glandular tissue at the base. Petals 5, white (according to label data of Capuron 29194 SF), obovate, asymmet- rical, 7-8 mm long and 7 mm wide, inner surface gland-dotted toward the base. Androecium co- roniform, staminal column ca. 1 mm tall; sta- mens 10, ca. 3 mm long from the base of the column, each of the 5 antisepalous pairs alter- nating with a staminode; filaments 1-1.5 mm long; anthers ca. 2 mm long; staminodes 5, ob- lanceolate, 4-5 mm long, ca. | mm wide. Ovary 5-locular, densely peltate-scaled; ovules 2 per locule, collateral; style 5-6 mm tall, glabrate; stigma obscurely 5-lobed. Fruit capsular, woody, spheroid, dorsally loculicidal, 6-8 mm long, 8- 10 mm wide, subtended by a woody, persistent calyx. Seeds 2 per carpel, asymmetrically ovate and laterally flattened, 5-6 mm long, 3-4 mm wide, each with a narrow, membranous, ventral eel. Additional specimens examined. MADAGASCAR. NORTHERN SECTOR. DISTRICT DIÉGO-SUAREZ: Versant Est du massif de I’ at ati 17 Dec. 1966 (fr), Capuron 27349 SF (P, TE ontagne des Français, 11 e Fla 1952 (fr), Service Forestier 5673 SF (P, TEF); lafa mbodipo-Antsahalalina, 7 June 1956 (fl), po vice Forestier 15962 SF (P, TEF). 1987] NOTES 451 Ficure 1. A-I. Helmiopsis sphaerocarpa. — A. — B. Schematic diagram of an inflorescence axis. — C. p . —D. Inner surface of calyx lobe showing ndun ce gii Inner surface of petal showing glandular —F. Detail of the Werben ies —G. Gynoecium. —H. Fruit. —I. Seed. (A, B. Capuron 27349 SF; C-G. EL... 29194 SF; H, I. Service Forestier 5673 SF. 452 Helmiopsis sphaerocarpa occurs on mountain slopes between 50 and 800 m altitude. It has been reported on black volcanic soils, but its presence on the Montagne des Francais suggests that it also may occur on limestone (Lemoine, 1906). The 5-carpellate gynoecium and reduced stig- mata of Helmiopsis sphaerocarpa place this species in Helmiopsis subg. Helmiopsis. Its broadly ovate leaves, glandular petals, and ten stamens ally it with sect. Glandulipetalae Arénes in subg. Helmiopsis. Helmiopsis pseudopopulus (Baillon) Capuron, a member of the same sec- tion, may be the most closely related species; it also has a many-flowered inflorescence, occa- sionally with a terminal axis. Glandular tissue at the base ofthe calyx, however, has been observed in only one other He/miopsis species, H. inversa H. Perrier (sect. Helmiopsis). This research was supported by NSF Doctoral Dissertation Improvement Grant BSR-8414032. thank the curator of P for the loan of specimens, and of TEF for permission to examine material. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 The illustration was prepared by P. Parker. L. Dorr and B. Stein provided helpful comments on the manuscript. Work in Madagascar was made possible by collaboration between the Mis- souri Botanical Garden and the Parc de Tsim- bazaza, Antananarivo, Madagascar. LITERATURE CITED ARÈNES, J. 1959. 131e Famille—Sterculiacées. Jn H. Humbert (editor), Flore de Ma adagascar et des Co- es. Mus ist. Nat., Ph Notice de la carte, Madagascar. S fique et Technique de l'Institut Francais de Pon- dichéry, Pondichéry, India LEMoINE, P. Etudes géologiques dans le nord de Madagascar. Librairie Scientifique A. Her- mann —Lisa C. Barnett, Department of Botany, Uni- versity of Texas at Austin, Austin, Texas 787 13- 7640, U.S.A WITHERINGIA FOLLICULOIDES (SOLANACEAE): A NEW SPECIES FROM COSTA RICA The genus Witheringia L'Hér. embraces about a dozen species. One species, W. solanacea L'Hér., is weedy in nature and ranges from north- ern Mexico to Bolivia. The other species tend to be more localized, and the genus seems to be best developed in mature wet forests of Costa Rica, Panama, and northern South America. Wither- ingia is a member of subfamily Solanoideae. The species of Witheringia are mostly shrubs or small trees with few- to many-flowered inflorescences of yellow or green flowers held under the leaves, and berries held erect above the foliage. The ca- lyces range from minute and nonaccrescent to arge and, as in the present species, completely enveloping the fruit. Witheringia folliculoides D'Arcy & J. L. Gentry, sp. nov. TYPE: Costa Rica. Puntarenas: moist forest in valley bottom, tropical wet forest with open understory on steep slopes and ridges and areas of secondary vegetation north and west of the air field 5 Rincón de Osa, 50-200 m, J. L. Gentry & Burger 2844 (holotype, MO-2825976; iso- type, F) ° amc Frutex 3-5 m altus, ramis hornotinis dun d dads Folia integra, cupias lanceolatave, saepe o qua, 9.5-23 cm longa lata, venis n: D lanis 14 tiolis gracilibus. nET pauci, 3-4 mm longi. Flores 5-meri, calyce parvo, 2.5-4 mm longo, glabrato, cam- panulato, apice tru Bae in statu fructificanti mag- 2.5-3 mm ee ae obtusis, 3 escentibus, antheris oblongis, haud apiculatis, longitudinaliter dehiscentibus. Acinus subglobosus, 12 olliculo tectus, calyce in folliculum e wA um maturescenti. Small tree 3-5 m tall; young branches slender, glabrate, drying smooth. Leaves entire, drying concolorous, mostly elliptic or lanceolate, some- times oblanceolate, often oblique, 9.5-23 cm long, 2.3-7 cm wide, apically short acuminate, basally mostly cuneate, the veins plane or slightly im- pressed above, sometimes drying somewhat ru- gose, the costs and major lateral veins elevated beneath, the lateral veins arching, 6-14 on each side; minor leaves lanceolate to broadly elliptic or rotund, 2.3-7.8 cm long, 1.4-4.2 cm wide, acute to acuminate or sometimes rounded api- ANN. Missouni Bor. GARD. 74: 453-454. 1987. cally, cuneate to rounded basally; petioles slen- der, 4-9 mm long, those of the minor leaves 2- 3 mm long. Inflorescence with the peduncle mostly subobsolete but sometimes 2-4 mm long and the flowers appearing fasciculate but only one or two appearing at a time; pedicels few, 1— 7 (evidenced by scars), 3-4 mm long, becoming 4—7 mm long, perhaps all but one flower abort- ing. Flowers 5-merous; calyx campanulate, api- cally truncate, 2.5-4 mm Hong, accrescent in fruit, drying dark, minutely y pubescent with short curved simple hairs anoetly on the apical portion, glabrescent, inconspicuously ribbed; co- rolla yellowish, apparently unmarked, glabrous outside, tubular campanulate, the tube 2.5-3 mm long, the throat pubescent with reduced hairs, the limb lobed less than halfway down, the lobes ovate, 3-4 mm long, marginally puberulent; sta- mens with the filaments pubescent at the base, apically glabrous, 1.5 mm long, the anthers yel- low, oblong, 2 mm long, not apiculate; ovary glabrous, the style equalling the stamens, gla- brous, the stigma minute. Fruit a subglobose ?juicy berry 10-12 mm across, loosely and com- pletely enveloped by the strongly accrescent ca- lyx; fruiting calyx rotund, much enlarged and inflated around the berry but open at the apex, apparently unribbed, glabrous, 15-20 mm across, approximately twice the length of the fruit; seeds numerous (?ca. 40), dark brown, flattened, re- niform, 2-3 mm across, the testa wavy retizulate on the face, scalariform on the rim; embryo broadly horseshoe-shaped, terete throughout (lacking any notching along its length). Paratypes. CosTA RICA. PUNTARENAS: Osa Penin- sula, tropical wet forest along dry stream bed parallel- d cón de Osa, 50-200 m, Gentry & Burger 2842 Adi X.S tto airport Rincón de Osa, 20-300 m, . Liesner 1887 (MO). This species is distinct from most other species of Witheringia in its bladdery fruiting calyx, whic loosely and completely envelops the fruit. It oc- curs in southern Costa Rica where a number of other species of the genus occur. It is perhaps most similar to W. exiguiflora D’Arcy and W. morii D'Arcy, which also occur in western Pan- 454 ama and southern Costa Rica, and which both have accrescent calyces, but their calyces do not form the large bladderlike enclosures of W. fol- liculoides. Witheringia folliculoides differs fur- ther from W. exiguiflora in its tubular campan- ulate corolla, in its smaller and much less coriaceous leaves, and in its larger fruiting calyx. From W. morii it differs in its broadly open co- rolla and its few-flowered inflorescences. In hab- itat W. folliculoides also differs from these two species: it occurs near sea level in the seasonally dry Osa Peninsula of Costa Rica, whereas W. exiguiflora and W. morii occur in Am rainy forests. Both W. exiguiflora and W. mo ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 occur in the uplands of 1,100-1,800 m in Costa Rica and Panama, but W. exiguiflora also ranges down to near sea level on the Atlantic slopes of Panama. This work was supported by National Science Foundation grant no. BSR-8305425 (William G. D’Arcy, principal investigator). — William G. D'Arcy, Missouri Botanical Gar- den, P.O. Box 299, St. Louis, Missouri 63166; and Johnnie L. Gentry, Jr., The University Mu- seum, University d Arkansas, Fayetteville, Ar- kansas 72701, U.S ESCHWEILERA COSTARICENSIS (LECY THIDACEAE): A NEW SPECIES FOR THE FLORAS OF COSTA RICA AND NICARAGUA Eschweilera C. Martius ex DC. is the largest (ca. 90 species) and most poorly known genus of neotropical Lecythidaceae. The genus is char- acterized by zygomorphic flowers with coiled an- droecial hoods, absence of anthers and produc- tion of nectar in the androecial hoods, bilocular ovaries with basal placentation, seeds often with lateral arils, and lack of differentiated cotyledons (for illustrations of these features see figs. 9, 10G, 13, and 18 in Prance & Mori, 1979). The Central American species of Eschweilera provide ex- amples of how poorly collected the species of Eschweilera are. Woodson (1958) included only four species of Eschweilera from Panama, where- as today we know ofat least 14 from that country. Further north, species diversity of Eschweilera is markedly reduced. There are only three species in Costa Rica, two or possibly three in Nicaragua, one in Honduras, and one in Mexico. Most of these are known only from one or very few col- lections. The recently described E. mexicana (Wendt et al., 1985) is the first species of Lecythi- daceae recorded from Mexico. It escaped collec- tion until 1983 despite being common locally. Although it might seem more appropriate to describe E. costaricensis in our forthcoming monograph of the zygomorphic-flowered genera of Lecythidaceae (Mori & Prance, manuscript), the need to have a name available for a treatment of the family for the Flora of Nicaragua (Prance & Mori, in press) mandates separate publication. Eschweilera costaricensis Mori, sp. nov. TYPE: Costa Rica. Heredia: Tropical wet forest along Guacimo ridge trail, La Selva Protec- tion Zone, 275 m, 18 Jan. 1983 (fl), Hart- shorn 2555 (holotype, NY; isotypes, BM, CR, F, K, MO, PMA, Ab E. pittieri venis impressis in pagina adaxiali fo- liorum et sine lobis calycis amplificatis in fructibus differt. Understory tree, 5-10 m tall. Leaf-bearing branches 3-5 mm thick. Leaf blades elliptic to widely elliptic, 15-26 x 7-15 cm, glabrous, char- taceous, with 10-11 pairs of adaxially impressed lateral veins; apex long acuminate; base obtuse to rounded; margins entire; petiole 7-8 mm long. ANN. MissounRi Bor. GARD. 74: 455-456. 1987. Inflorescences simple racemes or weakly once- branched, terminal, in axils of uppermost leaves, or from branches, the principal rachis 1.5-3 cm long, pubescent, the pedicels ca. 7 mm long, pu- bescen nt. Flowers 3-4 cm in diam., calyx with 6, d Pini 6- imbricate for '2 length; petals 6, widely ellibtie to orbiculate, cream colored or light yellow. Hood ofandroecium 17 x 17 mm, forming double coil with slight beginning of triple coil; staminal ring with 169—181 stamens; filaments ca. 1 mm long, expanded towards apex; anthers 0.5 mm long. Hypanthium sulcate, pubescent; ovary 2-locular, with 6—9 ovules in each locule, ovules attached to a hemispherically shaped placenta arising from floor of locule; style obconical, oblique, 2 mm long, arising from umbonate ovary summit 1 mm high. Fruits cup-shaped, 2.5 x 4 cm (ex- cluding operculum), the calycine ring inserted near apex of fruit base, supracalycine zone 0.5 cm wide, bearing ie ern calyx lobes and ped- icel when mature, pericarp 2-3 mm thick, with rough, lenticellate koi surface; operculum umbonate, 2 cm high, o 5 mm high. Seeds triangular in cross or. ca. 20 x 15 mm, lat- erally arillate. Distribution. Known only from wet forests in the Caribbean foothills of Costa Rica and Nic- aragua where it is an occasional (0.1—1.0/ha) understory tree (Hartshorn, pers. comm.). Flow- ers have been collected in January and August and mature fruits have been gathered in January. uidi specimens examined. NICARAGUA. RIO o Indio, Caño Negro, 4 Dec. 1982 (im- mature fr), Araquistain baie (M Co dui CA. : La Selva Protection Zone, cimo o Ridge Trail 300 m, 18 Jan. 1983 (fr, ahue eren (NY), 25 Aug. 1979 (fl), Davidson & Don- ahue NY). Eschweilera costaricensis is most closely re- lated to E. pittieri R. Knuth, which is relatively common and distributed from the Pacific coast of western Panama to the Magdalena Valley in the east and to the coastal forest of northern Ecuador in the west. Eschweilera pittieri does not 456 co-occur in the Caribbean forests of Costa Rica and Nicaragua with E. costaricensis. Eschweilera costaricensis differs from E. pit- tieri by having impressed lateral veins on the adaxial leaf surface and fruits without knobby calycine protuberances. I am grateful to Gary Hartshorn who first brought this new species to my attention and to G. T. Prance and G. Hartshorn for reviewing the script. My research on Eschweilera, done in collaboration with G. T. Prance, has been sponsored by National Science Foundation grant DEB-8020920. LITERATURE CITED Moni, S. A. & G. T. Prance. Eschweilera. In S. A. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Mori & G. T. Prance, Lecythidaceae — Part II. Mani act to be submitted for publication in FI. Neotrop. PRANCE, G. T. & S. A. ‘Mori. 1979, Lecythidaceae— Part I. Fl. Neotrop. Monogr. 21: 1-270 n press.) Lecythidaceae. Jn Flora RI & G. T. PRANCE. 1985. Esch- ida aceae): k new family : 347-351. ; ether eld In Flora of Panama. Ann. Missouri Bot. Gard. 45: 115-136. — Scott A. Mori, The New York Botanical Gar- den, Bronx, New York, 10458-5126, U.S.A. A NEW VARIETY OF HEDYOSMUM (CHLORANTHACEAE) FROM NICARAGUA Hedyosmum Swartz is a genus of ca. 40 species of primarily montane neotropical trees and shrubs with one herbaceous species in the Far East. In the New World species are found from southern Mexico to Panama, Venezuela, Colombia, Ec- uador, Peru, Bolivia, Brazil, and Paraguay, as well as in the West Indies. The greatest density of species occurs in the northern wet Andes. In the field the genus is distinguished by its opposite leaves with connate sheathing petiole bases, stipular appendages along the distal margin of the sheath, staminate inflorescences composed of many ebracteate flowers of single stamens, and pistillate inflorescences composed of solitary or clustered, bracteate flowers. Until a recent monograph (Todzia, 1986), the genus has been in a state of taxonomic confusion, in part because in the last hundred years Hed- yosmum had never been studied over its entire geographic range. Several species proposed from Central American material had, in fact, already been described from South America. Such is the case of Hedyosmum goudotianum Solms, a a originally described from Colombia but ow recognized to range from Nicaragua to Peru » bd 1986). This species is distinguished from all other Hedyosmum species by its elongate ra- cemose or paniculate pistillate inflorescences with many-flowered cymules on short peduncles, no- tably coriaceous leaves with impressed second- ary venation and sharp, closely spaced teeth, long leaf sheaths with striate triangular patches of ap- pressed hairs below the stipules, and trichomes (when present) restricted to the abaxial sides of the primary, secondary, and sometimes tertiary eins. Although populations throughout its range are variable with respect to trichome density, leaf size, and inflorescence length, the northernmost Rica (Burger, 1973, 1977). They are unusual in being glabrous and in having shorter internodes with often overlapping leaf bases, shorter leaf sheaths that lack the striate patch beneath the ANN. MissouRi Bor. GARD. 74: 457—459. 1987. stipular processes, and leaves with more widely spaced teeth A key to the varieties is provided below with a full description for Hedyosmum goudotianum var. mombachanum. Hedyosmum goudotianum Solms in A. P. de Candolle, Prodr. 16: 482. 1869. TYPE: Co- lombia: “‘Quindiu, El Inciendial, La pal- milla." Nov.-May 1844 (pist.), Goudot s.n. [lectotype (here designated), P.]. This is the only specimen of Goudot seen from this lo- cality. It 1s clearly designated as type ma- terial in Solms's handwriting. KEY TO VARIETIES OF HEDYOSMUM GOUDOTIANUM 1. Internodes (3-)4-9 cm long; leaves with teeth sheaths well-spaced and not overlapping, (1—) 1.8-3 cm long, with a triangular patch of ap- pressed hairs below stipular processes; Costa Rica, . Venezuela, Colombia, Ecua- dor, a ia oudotianum var. se d Internodes l- Perd cm long; leaves with tee —4 mm distant and with secondary vein flush with surface, these glabrous beneath; leaf sheaths often overlapping, (0.7—)1—1.6 cm long without a striate patch below stipular pro- cesses; Nicaragua . . H. goudotianum var. mombachum — Hedyosmum goudotianum Solms var. goudotian- um ae nada (Solms) O. Kuntze, Revis. Gen. : 566. 189 a monta ion Burger, Phytologia 26: 133. 1973. TYPE: Costa Rica. Heredia: Río Vueltas (up- Nov. 1969 (pist.), Burger & Liesner 6336 aiam type, F; isotypes, BM, COL, CR, GH, MO). Distribution. Costa Rica, Panama, Venezue- la, Colombia, Ecuador, and Peru in montane cloud forest. In Peru this variety has been re- corded in association with Chusquea, while in Costa Rica it is known to occur with Podocarpus, Weinmannia, and Clethra. In Central America flowering is most common May through August, 458 while in South America flowering occurs spo- radically throughout the year. Hedyosmum goudotianum Solms var. momba- chanum Todzia, var. nov. TYPE: Nicaragua. Granada: “En las últimas antenas del Vol- can Mombacho, 1,200-1,220 m, 23 Feb. 1981 (pist.), Moreno & López 7134 (holo- type, MO; isotypes, F, HNMN not seen). Differt a var. goudotianum internodiis 1-4(-5) cm longis, laminis foliorum subtus glabris dentibus 2.5-4 mm distantibus, venis lateralibus subtus non elevatis, vaginae glabrae superpositae Dioecious, aromatic shrubs or small trees, 2— 4 m tall, with prop roots; bark whitish-gray to gray, smooth; young stems quadrate, brittle, usu- ally rugose, sometimes glabrous; large stems te- rete, with tubular leaf bases persisting and be- coming fibrous with age; internodes 1-4(-5) cm long, the nodes slightly swollen. Leaf blades nar- rowly elliptic, elliptic, ovate to obovate, 3.3-13.6 cm long, (1-)2.7-5.3(-7) cm broad, with acu- minate tips 0.2-0.7 cm long, cuneate to obliquely cuneate at base, at margins sharply serrulate with teeth. 2.5-4 mm distant continuing to apex, sometimes revolute, smooth, dull, light green above and beneath when fresh, drying charta- ceous to subcoriaceous, slightly scabrous, gray to brown above and beneath; midveins impressed above, raised beneath, glabrous; lateral veins 6— 8, 7-13 mm distant, arcuate, flush with surface and glabrous beneath; intersecondary veins ex- tending '4 to 2 distance to margin; free portions of petioles smooth to asperous, 0.5-0.8 cm long, narrowly winged; petiolar sheaths smooth to as- perous, glabrous, (0.7—)1-1.6 cm long, 0.6-0.8 cm broad at apex, slightly inflated, terete or quadrangular, with or without 2 raised longitu- dinal ciliate lines extending down length of sheath from stipular appendages, overlapping, persis- tent, becoming gray and fibrous with age, not extending beyond free portions of petioles, at distal margin with 2 caducous, linear to slightly fimbriate stipular appendages ca. 1 mm long. Sta- minate inflorescences terminal or axillary, 1.5— 4 cm long, composed of 1-2 opposing pairs of spikes on a short rachis terminated by a single spike; bracts subtending terminal inflorescence linear to obovate, 0.7-2 cm long, broad; mature spikes 1.5-4.5 cm long, 0.3-0.6 cm broad, with ca. 100 stamens, sessile or borne on short peduncles 1-2 mm long, each subtended by a small, dentate or entire, spatulate to obovate ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 bract 0.2-1 cm long, 1-2 mm broad; stamens congested on axis but becoming 0.5-1.5 mm dis- tant; rachis 1-2.5 mm thick with a thick, irreg- ularly margined, basal annulus; anthers yellow- ish-green, 1.3-1.8 mm long, 0.6-1 mm thick; connectives extended ca. 0.2 mm beyond thecae, acute. Pistillate inflorescences axillary or termi- nal, racemes or panicles (1.3-)2.5—4.4 cm long bearing 8-13 cymules; subtending inflorescence bracts 15-20 mm long, 3-5 mm broad; cymules with (2—)3-4 clustered flowers, 3-4 mm long and broad, borne on short peduncles 1-3(-5) mm long, alternate or opposite on inflorescence axis, 2-6 mm distant; subtending floral bracts green, connate in lower 4 to 34, 2-3 mm long including acuminate tips 1-2 mm long, 2-4 mm broad, ciliate or entire along free margins, enclosing '^ to *⁄4 of flower. Pistillate flowers trigonous, 2-3 mm long, 1.5-2 mm thick with a minute to large pore on each face of the ovary; perianth lobes deltate, acute, 0.2-0.5 mm long, basally connate; stigmas white, 2-3 mm long, irregular in shape, linear to irregularly lobulate, 2- or 3-angled, with long papillae. Fruiting cymules white, irregularly globose, 5-8 mm diam.; seeds ca. 3 mm long, brown, trigonous, minutely papillate. Distribution. Elfin and cloud forest on Vol- cán Mombacho and Volcán Maderas in Nica- ragua in association with Cavendishia, Clusia, and Freziera at elevations of 740-1,220 m. Flowering and fruiting occur sporadically throughout the year. Additional "a is examined. NICARAGUA. GRA- ADA: Volcán mbacho, Atwood 7771 (MO, US); upper slopes of Volcán Mombacho along W shore of Lake Nicaragua, ca. 15 km S of Granada, 1,100-1,200 m, Croat 39092 (F, MO); Volcán Mombacho, Plan de Flores, 740 m, Grijalva 2506 (HNMN, TEX); N slope of Volcán Mombacho, above Finca El Progreso, Neill somewhat above Plan del Flores, 950—1,150 m, Stevens 4332 (F, HNMN, MO, TEX). Rivas: near summit and upper slopes of Volcán Maderas above Balgüe, Isla ae - E 200 m, Nee & Robleto Téllez 28086 (HNM X). I gratefully acknowledge the support ofthe Na- tional Science Foundation (doctoral dissertation improvement grant BSR-8400923) and The University of Texas at Austin (graduate con- tinuing fellowship). I thank B. B. Simpson, J. Henrickson, and B. A. Stein for comments on various drafts of this paper, and M. C. Johnston for help with the Latin diagnosis. The curators 1987] and staff of the following herbaria generously provided loans of specimens: BM, COL, CR, F, GH, MO, MSC, P, TEX, WIS, US. LITERATURE CITED BuRGER, W. C. Notes on the flora of Costa Rica, 2. ages of the Chloranthaceae. Phy- tologia 26: 131- NOTES 459 1977. Chloranthaceae. Jn W. C. Burger (ed- or), Flora Costaricensis. Fieldiana 40: 1-10 Systematics and evolution of the genus Hedyosmum (Chloranthaceae). Unpub- lished Ph.D. Dissertation. The University of Tex- as at Austin, Austin, Texas. — Carol A. Todzia, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 61366, U.S.A. BOOK REVIEW Tomlinson, P. B. 1986. The Botany of Man- groves. Cambridge University Press. ISBN 0-521-25567-8. Price: $69.50. (Initiating a new series: the Cambridge Tropical Biology Series.) Almost any botanist who has visited the trop- ics or subtropics will have encountered man- groves, and this refreshingly written book is the perfect introduction to both their taxonomy and biology. Overall, the book is a pleasing potpourri of selected data from many different fields about many different aspects of mangroves. The bulk of the book (211 pages) is a family-by-family summary of mangrove taxa, including original and very useful keys to the different species and genera of most of the mangrove families. Also included are discussions of various aspects of morphology, architecture, reproductive biology, and biogeography, as well as miscellaneous taxo- nomic notes and elegant illustrations of impor- tant species and genera. Although a key to "strict mangrove" genera is provided in the floristic sec- tion, no attempt is made to key out the different back-mangrove families. Thus the reader must first know the genus or family of the plant in which he is interested in order to take full ad- vantage of Tomlinson's taxonomic section. Nevertheless, from the viewpoint of the system- atic botanist, the gathering together in one place of the available nomenclatural and taxonomic data for mangrove species and genera is a high point of the book, as are the author's personal Observations on identification and biology of the individual species. Throughout, but especially in the photo cap- tions, the author's puckish sense of humor often comes to the fore. A Nypa stem is aptly described as "resembling nothing so much as a series of overlapping cowplats." The fruits of Avicennia alba are said “to resemble a gorged leech.” An “obliging butterfly visitor" in a photo of Lum- nitzera racemosa is contrasted with lack of an obliging bird visitor in L. /ittorea. Another strong point of the book, as might be expected considering the author's research pre- dilection, is the series of chapters devoted to the fascinating morphological and anatomical spe- cializations of mangroves. No fewer than four chapters and 71 pages treat shoot systems, root systems, water relations and salt balance, an seedlings and seeds. In contrast, the five chapters ANN. MissounRi Bor. GARD. 74: 460—462. 1987. devoted to ecology, floristics, biogeography, flowering, and utilization and exploitation amount to only 68 pages I found the treatment of mangrove ecolog disappointing. The author acknowledges this de- ficiency and justifies it by noting that ecology is generally outside his area of expertise. However, tremendous amount of effort has been devoted to this field, and a few extra pages devoted to mangrove zonation and summarizing some of the abundant and often conflicting literature that implicates different toleration of salt concentra- tions vs. soil texture and edaphic conditions vs. rainfall would have been a welcome addition, especially if accompanied by the author's own trenchant evaluations. A minor but bothersome ecological problem is Dr. Tomlinson's adoption of the word **man- gal" to refer to the mangrove community while "mangrove' is reserved for the t plants I find this to serve no useful puis and to be distinctly cumbersome in the same way as the plethora of Braun-Blanquet lene end- ings. Luckily, after emphasizing “mangal” in the first chapter, the author himself largely abandons it in later chapters where traditional terms like "mangroves, mangrove associates," and "mangrove communities" are often used in- stead. In this review I will use exclusively the traditional terms in hopes that “mangal” (as well as Rhizophoretum, etc.) will soon pass into well- deserved oblivion. Another, more significant, ecological problem is the author's attempt to separate mangrove species into "strict mangroves" (subdivided as to major and minor components) and **man- grove associates." While this should be ue a natural way to divide mangrove taxa, in opinion many taxa are wrongly placed in is tables on pages 27-30, and others, even those treated elsewhere in the book, are outright omit- ted. This is especially obvious for the Pacific- American mangrove taxa, and inclusion of some of the omitted full-mangrove species would give rise to very different conclusions. For example, the taxonomic distinctiveness of strict mangrove species is far more often at the level of species than Tomlinson implies. Glaring outright omis- sions from these lists include Tabebuia palustris and Phryganocydia phellosperma, which are strict mangroves, not even **back-mangroves"'; indeed ”°” 66 1987] in the case of Bignoniaceae the back mangroves are correctly listed but the full mangroves are completely ignored! Crenea patentinervis, men- tioned as a potential herbaceous mangrove in the text, is not included in any of the tables of man- groves or mangrove associates on the grounds that as an herb it cannot be a true mangrove. Actually Crenea is a subshrub and is at least as woody as such included genera as Acrostichum, Tuberostylis, Acanthus, or Batis. Even were it an herb, that would be no reason to exclude it, es- pecially as it is a full-mangrove species, not a back-mangrove. Both species of T'uberostylis are mangroves and they are mostly epiphytic on the roots and lower trunks of full-mangroves rather than on back-mangroves. Some major mangrove components are relegated to the “minor” list, e.g., Pelliciera which forms pure mangrove for- ests many km? in extent in the delta of the Rio San Juan in Colombia. Gymnosperms are spe- cifically mentioned as playing no role in man- groves even though Zamia roezli is typically found in mangroves and is restricted to them and to the adjacent coastal fringe. Muellera (p. 263) is stated to be most commonly recorded well away from the sea, but I have seen it only in the back-mangrove regions with a distinct tidal in- fluence and strongly suspect that the entire genus is restricted to this habitat. There are also serious problems with the num- bers of species given for mangrove genera, es- pecially since the data in these tables are used as evidence of the taxonomic distinctiveness of mangrove plants. Thus the single b ofthis 18- -species genus, and Hippomane has four noncoastal species as well as the well-known coastal one. Mangrove epiphytes are implied to be basically plants from nearby terrestrial com- munities that transgress into the mangroves. Yet many epiphytes seem unique to mangroves (such as the entire genus Tuberostylis) and others are certainly more characteristic of mangroves than other habitats. Similarly, Tomlinson states that there are no climbers in mangroves, presumably because climbers have wide vessels subject to extreme water tension, yet there is at least one clear exception. Phryganocydia phellosperma is not a species that roots behind the mangroves and scrambles into them (although its unlisted confamilial Cydista aequinoctialis is and should probably be added to the list of mangrove as- sociates); it is a strict mangrove and roots in (and only in) the mangroves themselves. BOOK REVIEW 461 The biogeographic discussion focuses so much on the dichotomy between the relatively depau- perate western mangroves vs. the more diverse eastern ones that the almost equally striking dif- ference between the richer mangrove flora of the acific Coast of South America as compared with the Atlantic one (possibly related to the rel- atively recent opening of the Atlantic Ocean?) is overlooked. Only eight true mangroves are said to occur in the western hemisphere, ANS a local concentration of “incipient mangroves” western Colombia is acknowledged. While dese species are said to be mere mangrove associates that “lack complete fidelity to mangal” (p. 55), several of them are true and obligate mangroves in the strictest sense (although there are also en- demic back-mangrove species in this region). It may well be that the western Colombian man- groves are fundamentally different from other mangroves in their greater habit diversity and in being mostly individual mangrove species of otherwise nonmangrove genera, but they are not less true mangroves for that. Knowledge of man- groves would be better served by focusing on the unusual aspects of these species rather than by trying to sweep them under the rug. There are a number of inconsistencies, espe- cially in the biogeographic discussions. Rhizoph- ora racemosa is on both Atlantic and Pacific sides of tropical America as reported on p. 334, but in the key (p. 329) it is characterized as being only on the Atlantic coast of South America. There are five Pacific coast collections in the MO herbarium, as well as collections from Honduras, Costa Rica, and Panama, all outside the Vene- zuela and Guianas to West Africa range indicated by Tomlinson (p. 335). Rhizophora harrisonii is rather precisely mapped as having a disjunct population in the middle of the Peruvian coastal desert outside the range of any mangrove, but the numerous records from coastal Ecuador and Colombia were apparently overlooked. On the other hand, the range of R. samoensis is hypo- thetically extended to include the Pacific coast of Sout merica, where it may occur but has not yet been documented; that species does reach the Galapagos, according to R. Horna (pers. comm.). Inevitably a few insignificant errors in spelling of Latin binomials, taxonomic authorities, etc. are unavoidable in a book of this scope. Exam du include *Anaemopegma" (p. 32), " Mouri- ` and **Pachyra" (p. 56) and the authors of AUR species of Phryganocydia (p. 214). More 462 problematic is Lysianthus (presumably = Lys- iana?) supposed to be a mangrove mistletoe (p. 33). Dalbergia amerimnion Benth. (p. 261) has long been regarded as a synonym of D. brownei. Phryganocydia phellosperma is not distinguished from P. corymbosa by a simple tendril, a trait shared with the entire genus. Tabebuia palustris is not deciduous as stated (p. 214), which would have been quite remarkable in a mangrove, but oi like all other mangroves known to me. milar minor errors in biogeographic distri- butions include Tuberostylis rhizophorae, whose aimed “wider distribution in Central America” -consists of a single collection from southernmost Darién, and Pavonia rhizophorae, supposed to be recorded only for Colombia but reported in the Flora of Panama to cross the Panama border in the same part of southern Darién. Obviously, it is difficult to eliminate such mis- cellaneous errors. Whether they are as frequent ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 in other taxa and regions as among the man- groves I happen to know personally (i.e., mostly Bignoniaceae and Pacific American), I do not know. But it seems likely that it would have been useful for the author to run a draft of his manu- script past a few taxonomic specialists or field botanists as well as checking a few more herbaria for distributional data. Such minor imperfec- tions detract very little from the message of the book, except that inasmuch as this is the closest thing to a monograph of many mangrove taxa that we are likely to have in the foreseeable fu- ture, it is a bit of a shame that some of the work that would have gone into an actual monograph was neglected. Overall The Botany of Mangroves succeeds admirably in its purpose of providing a concise and highly readable introduction to the world’s mangroves.— 4/wyn Gentry, Missouri Botanical Garden, P.O. Box 299, St. Louis, Mis- souri 63166, U.S.A. Volume 74, No. 1, pp. 1-182 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN was published on 15 June 987 A 7 ri F P $ | FE j i DUNS. š . š ç | F tí S" r m A ri ETE " d t £ ; I x t. Oligocarpae | ert Kral, Jame I Charlotte M. eH 2223 CONTENTS > Bada S * New Species of Thrasya TL Panicoidaea) bo diy Mow of aise and 52 g avidse & A Cerri t se Adni ir G. Burman dus 63A 43M um Contents c continued on nec Vascular Epiphytism: Taxonomic Participation and kes Diversity David H. Ben- zing 183 Diets and Ships of Neotropical Vascular Epiphytes Alwyn H. Gentry & C. H. Dod. 205 Nus Fixation by Epiphylls in a Tropical Rainforest Barbara L. Bentley -aana 234 Adaptive Radiation of Salamanders in Middle American Cloud Forests David B. Wake 242 Revision of Eremanthus (Compositae: Vernonieae) Nanda F. F. Macebeish 2 265 Biosystematics of Tetraploid Eucharis (Amaryllidaceae) Alan W. Meerow 2 s 291 The Shrubby Gentian Genus Macrocarpaea in Panama Kenneth J. Systm „= 310 . Menoecy and Sex Changes in Freycinetia (Pandaceae) Hans-Helmut Poppendieck .. 314 A Guide to Collecting Lecythidaceae Scott A. Mori & Ghillean T. Prance .cc:2cccco 321 Chromosome Cytology of Aq (Asteraceae- Mutisieae) Peter Goldblatt ......... 331 ` Notes on Cipura (Irid hand Central America, and a New Species from Venezuela Peter Goldblatt & Da E. Henrich . 333 n ‘tenis (Lepidoptera: Nymphalidae): Summary of Known Larval Food Plants Bone A. Peu IE Keith S Brown dro V ens 341 . Chemistry at the Solanaceae/Ithomiinae Interface Keith S. Brown, Jr. 399 : 5 New Taxa of Rubiaceae from Venezuela - Julian A. Steyermark PR euer cic . 398 E New Species of Neot pical | Lauraceae Henk van der Werff... ipii, VOR * A New Species of Ocotea shee from Southeastern Mexico Tom We nš må Henk van der Verf... 2 Mic EL gs Pe E OSEE Four New: Species o of Axonopus (Poterie: Panicese from Tropical America Gerrit ; ! se T | Two Mean ERS : : quea (Poaceae: Bambusoideae) Lynn G. Clark 424 New Caitlin in TRE ea blue Lynn G. Clark ——— M8 Me Ne Cor nus cad in South America Julian A. Steyermark & Ronald ilia | FRESI sian Else Zardini .— u aN Em ; Chromosome Counts of Missouri Asteraceae and Poaceae — Ahsan A. Vahidy, ! Gerrit LO TS — Davidse & Youji . Shige — EE d e W Volume 74 \ Z N 7A N u mber 3 ! Ac N ae i P rd Volume 74, Number 3 Fall 1987 Annals of the Missouri Botanical Garden The Annals, published quarterly, contains papers, primarily in systematic botany, con- tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the Garden will also be accepted. Authors should write the Editor for information concerning arrangements for publishing in the ANNALS. Instructions to Authors are printed on the inside back cover of the last issue of each volume. Editorial Committee George K. Rogers Editor, Missouri Botanical Garden Janice Wilson Editorial Assistant, Missouri Botanical Garden Marshall R. Crosby Missouri Botanical Garden Gerrit Davidse Missouri Botanical Garden John D. Dwyer | Missouri Botanical Garden & Saint Louis University. Peter Goldblatt EE - Missouri Botanical Garden — s Henk van der Werff Mason | Borena ae For s subscription rudis contact Dolu Ey s , P.O. Box 299, St. Louis, MO 63166. Sub- — scription price is $75 per volume U.S., $80 Canada — — Mi and Mexico, $90 all other countries. Airmail deliv- ery saa = pm volume. Four i issues mui vol. xs ume. E: (ISSN 0026-6493) i is published. issouri ` enue, St. Louis, MO 63110. ie; "paid at St. Louis, MO and ac ; mailing o .. PosTMASTER: Send address chang 166. | Bent P.O. Box 299, oe 10 631 The ANNALS OF THE Missouri T Gano r ere. dr Botanical Garden, 2345 Tower 2. postage Ç Volume 74 Number 3 1987 Annals of the Missouri Botanical Garden NZ A REVISION OF PANICUM SUBGENUS PANICUM SECTION RUDGEANA (POACEAE: PANICEAE)! FERNANDO O. ZULOAGA? ABSTRACT Panicum subg. Panicum sect. Rudgeana, herein revised, includes six species: P. cayennense, P. campestre, P. cervicatum, P. ligulare, P. rudgei and P. vinaceum. It is characterized mainly ideis n upper anthecium. The stipe consists of two portions: a membranous portion towards the ve rtion towards its dorsal face. The position of the section within of the spikelet and an indurate po ntral fac subg. Panicum is discussed, as is the relationship of Rudgeana with other sections containing species Rin a stipitate upper anthecium. Hitchcock & Chase (1910) included P. rudgei Roemer & Schultes and P. rotundum A. Hitchc. & Chase within the ungrouped species of the genus and noted their close relationship. In 1915 they repeated this treatment, indicating also that the species they had described was the same as P. campestre Nees ex Trinius. The same year, Hitchcock also placed P. rudgei in an informal group he named Rudgeana. Chase, in unpub- lished manuscripts, later placed both species in the Rudgeana group, which she characterized as “Rather stout much branched perennials with usually harshly pilose or papillose sheaths. Spikelets abruptly pointed, the first glume point- ed, more than half the length of the spikelet, the midnerve scabrous.” The two species mentioned above plus P. cay- ennense Lam., P. ligulare Nees ex Trin., P. vi- naceum Sw., and P. cervicatum Chase share char- acters that allow them to be included in sect. Rudgeana (A. Hitchc.) Zuloaga. Section Rudgeana falls within subg. Panicum, having the following characters in common with the rest of the sections in the subgenus [which are sects. Panicum, Repentia Stapf, Urvilleana (A. Hitchc. & Chase) Pilger, and Dichotomiflora (A. Hitchc. & Chase) Honda]. Species of subg. Panicum are characterized by the presence of the C, photosynthetic pathway of the NAD-me subtype (Brown, 1977) and are distinguished anatomically by having a double sheath around the vascular bundles. The inner ! I wish to thank the Consejo Nacional de Investigaciones Científicas y Técnicas de la Repüblica Argentina (CONICET) for a grant that allowed me to spend 1982 doctoral fellow. I owe appreciation to Cecilia Ezcurra and Em of the manuscript. The line drawings were made by Vladimiro ‘Duda as, and 1983 at the Smithsonian Institution as a post- met J apt sva for help during the preparation am always grateful 2 Instituto de Botanica Darwinion, Casilla de Correo 22, San Isidro, peciit ANN. Missouni Bor. GARD. 74: 463-478. 1987. 464 one is a mestome sheath with thick-walled cells. It is surrounded by a Kranz outer sheath con- taining specialized chloroplasts that are usually disposed centripetally. Between each vascular bundle there are two or three tabular cells ar- ranged radially. The number of secondary vas- cular bundles present between each primary bun- dle varies from two to six. In sect. Rudgeana, as in most sections of subg. Panicum, the plants are cespitose and short rhi- zomatous with erect, few- to many-noded culms. The ligule is membranous at the base and short- to long-ciliate at the upper portion. The leaf blades are lanceolate to linear-lanceolate, with or with- out involute borders. The species are usually found in dry and open places, but some species in sect. Dichotomiflora and in sect. Repentia grow in wet places and have decumbent culms that root at the lower nodes. The inflorescences are pyramidal, lax and dif- fuse, and have ellipsoid to lanceolate spikelets dispersed on the branches. The nervation of the glumes and lemmas and ornamentation of the upper anthecium are dis- tinctive characters that hold together the sections of the subgenus. The upper glume and lower lem- ma are 7- to 9-nerved (1 1- to 15-nerved in species of sects. Rudgeana, Panicum and Urvilleana), with a few exceptions in species of sects. Dichoto- miflora and Repentia, in which these bracts are 5-nerved. The upper anthecium is smooth and shiny over the entire surface, and compound or both compound and simple papillae are present near the apex of the upper palea. Panicum sect. Rudgeana differs from sect. Di- chotomiflora by the length of the lower glume ('⁄4 to 3 the length of the spikelet in sect. Dichotomi- flora) and by the absence of papillae on both surfaces of the leaf epidermis; also, as noted be- fore, species of sect. Dichotomiflora grow in hu- mid places with the culms decumbent and root- ing at the lower nodes. Section Rudgeana is separated from sect. Repentia by the absence of stout rootstocks at the base of the plant. Section Urvilleana is distinguished from sect. Rudgeana by having long macrohairs at the base of the upper lemma and numerous, whitish oa cov- ering both glumes and the lower lem Section Rudgeana can be aoaaa clearly from sect. Panicum and the sections mentioned above by the occurrence of a well-developed stipe at the base of the upper anthecium. Two seg- ments of the stipe can be distinguished: a) a por- tion of membranous tissue towards the palea of ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 the upper anthecium (Figs. la, e, 2d, e, 3b) and ) a portion of indurate, smooth, and shiny tissue towards the lemma of the upper anthecium (Figs. I b, f, 2a, 3c-e). The texture of the indurate por- tion is similar to that of the main portion of the upper anthecium. The membranous portion of the stipe is reduced in P. cayennense and P. cam- pestre (Figs. le, 4g, h) or is larger and more ex- panded in P. rudgei (Fig. 3b) or remarkably no- ticeable in the rest of the species of the section. In P. ligulare and P. vinaceum the membranous portion of the stipe is prominent and may be prolonged into one (Fig. 5h, i) or two wings (Fig. 2e) that cover the base of the upper anthecium. In all cases the membranous portion appears tur- gid when the spikelet is rehydrated, and it is free from the base of the upper anthecium. The indurate portion of the stipe is found be- low the upper anthecium and is appressed to the membranous portion, at least when the spikelet is immature (Figs. la, b, 2a, 3d, e). At maturity, it extends behind the upper anthecium as a mu- cro (Fig. 1f). Size and length of this mucro vary among species of the section, but it usually re- mains on the rachilla when the upper anthecium falls (Fig. 3c). In Australia there exists a group of Panicum species with a structure similar to the stipe found in sect. Rudgeana. These species were trans- ferred from Ichnanthus Beauv. to Panicum by Lazarides (1959), who noted that the appendages found at the base of the upper anthecium are not adnate to the upper lemma (as in Ichnanthus) but rather originate from the apex of a noticeable stipe. Shaw & Webster (1983) supported this concept, emphasizing distinctness of the ap- pendages in /chnanthus from Australian species of Panicum. More recently, Lazarides & Webster (1984) removed these “ichnanthoid” species from Pan- icum, erecting for them the new genus Yakirra. Included in it were four species previously treat- ed in Panicum: Y. muelleri (Hughes) Lazarides & Webster, Y. majuscula (F. Muell. ex Benth.) Lazarides & Webster, Y. australiensis (Domin) Lazarides & Webster, and Y. pauciflora (R. Br.) Lazarides & Webster; also included was a new species, Y. nulla. They provided a table of features separating Yakirra from Ichnanthus and Panicum and stat- ed that there were no conclusive characters to differentiate Yakirra from Panicum besides the presence of a stipe at the base of the upper an- thecium. I regard this as correct, since the other ZULOAGA— PANICUM SUBG. PANICUM SECT. RUDGEANA GURE 1. Scanning electron mt of the upper anthecium of Panicum species. a-d. P. ligulare.—a. Lateral view of the base showing the stipe.—b. Dorsal view of the base showing the indurate portion of the stipe.—c. Apex of the upper anthecium i diit: papillae at the tip of the palea.—d. Detail of the papillae. e, f. P. campestre. —e. Ventral view of the base of the upper anthecium showing the membranous portion of the stipe.—f. Dorsal view showing indurate portion of stipe. a-d, based on Irwin 1 4904; e, f, based on Chase 8645. Scale bars: a—c, f, x 100; d, x 500; e, x150. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURE 2. Scanning electron micrographs of the upper anthecium of Panicum species. a—c. P. cervicatum. — mbranous and indurate portion of the stipe. — b. Apex showing compound papillae i f. P. v pound papillae at the tip of the palea. a—c, based on C ase 7 0737; d-f, based on Steyermark 59173. Scale bars: a, d, x 50; b, x 100; c, x 500; e, x 70; f, x 300. 468 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FIGURE 4. Panicum campestre. —a. Habit.—b. Spikelet, lateral view.—c. Spikelet, ventral view.—d. Spikelet, dorsal view. —e. Lower palea, dorsal view. —f. Lower palea, ventral view.—g. Upper anthecium, dorsal view. — h. Upper anthecium, ventral view.—i. Caryopsis, embryo side.—j. Caryopsis, hilum side. Based on Sendulsky 637. 1987] ZULOAGA — PANICUM SUBG. PANICUM SECT. RUDGEANA 469 FicunE 5. Panicum ligulare.—a. Habit. —b. Spikelet, lateral view.—c. Spikelet, ventral view.— d. Spikelet, dorsal view.—e. Lower palea, ventral view.—f. Lower palea, dorsal view.—g. Upper anthecium, dorsal view.— h. Upper anthecium, ventral view.—i. Upper anthecium, lateral view. Based on Irwin 14904. 470 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE 1. Comparison of sections of Panicum with stipitate upper anthecia. [VoL. 74 Subg. Phanopyrum Sect. Sect. Sect. Characters Stolonifera Parvifolia Phanopyrum Stipe a Homogeneous Homogeneous ' Homogeneous Stipe pre In all species In a few species In the only species Ended: pathway š C, Compound papillae at tip of palea Absent Absent sent Inflorescence type Racemose Paniculate Racemose branches branches branches Upper glume and lower lemma 3- to 5-nerved 3- to 5-nerved 3- to 5-nerved nervation Panicle dimorphism Absent Absent Absent characters listed for Yakirra (habit, life form, ret, and photosynthetic path- way) are common features of different sections of subg. Panicum. Lazarides & Webster (1984) noted the pres- ence ofa stipitate flower in the American P. gym- nocarpon Ell. On this basis they accepted this species as correctly placed by some authors in the monotypic genus Phanopyrum (Raf.) Nash and asserted that “The acceptance of Phanopy- rum as a valid genus makes Yakirra morpho- logically distinct from Panicum, based on the presence or absence of a stipitate flower.” However, stipitate upper florets are also pres- ent in species of sects. Lorea Zuloaga, Agrostoi- dea (A. Hitchc. & Chase) Hsu, Stolonifera (A. Hitchc. & Chase) Pilger, Dichanthelium, and Parvifolia (A. Hitchc. & Chase) Pilger. wean ters distinguishing these taxa are summarize Table 1 Section Rudgeana is similar to Yakirra in de- tails of habit leaf blades, ligules, inflorescences, spikelet compression and length, form and ner- vation of glumes and lower lemma. It differs from the Australian genus mainly by having a heter- ogeneous stipe below the upper anthecium, a lower palea (almost absent in Yakirra) well-de- veloped, and the upper anthecium with com- pound papillae at the tip of the palea only. In species of Yakirra the anthecium has simple pa- pillae in longitudinal rows all over the lemma Therefore, th fal in sect. Rudgeana is a good character for its de- limitation within subg. Panicum, but I judge it to be an insufficient one for removing species from Panicum. The elongation of the rachilla could help in opening the spikelet and posterior dispersal of the caryopsis. Davidse (in press) has pointed out that the stipe below the upper anthecium in P. cervicatum is an elaiosome involved in ant dis- persal of the diaspore and noted that a similar elaiosome might be present in P. vinaceum and P. trinii. Berg (1985) reported a similar elaio- some in the stipe of Panicum australiense Do- min. METHODS AND MATERIALS Classical taxonomic studies have been carried out in this paper, utilizing a Wild M5 dissecting microscope and a Wild M20 microscope. For LE, M, MO, NY, P, R, RB, S, SI, SP, and US. SYSTEMATIC TREATMENT Panicum subg. Panicum sect. Rudgeana (Hitch- up R n 1915. TYPE: Panicum rudgei Roemer & Schultes. Cespitose perennials or occasionally annuals, with erect, more or less branched culms and usu- ally pilose leaves. Ligule membranous, short- to long-ciliate. Leaf blades lanceolate to linear-lan- 1987] ZULOAGA — PANICUM SUBG. PANICUM SECT. RUDGEANA TaBLE l. Continued. Subg. Subg. Subg. Phanopyrum Dichanthelium Agrostoides Subg. Panicum Genus Yakirra Lorea Dichanthelium Agrostoidea Rudgeana Homogeneous Homogeneous Homogeneous Heterogeneous Homogeneous In a few species In a few species In a few species In all species In all species ° C, C4, NADP-me 4 NAD-me C, NAD-me Absent Absent Present Present Paniculate Paniculate Paniculate Paniculate branches Absent Paniculate branche branche 3- to 5-nerved 7- to 9-nerved Absent Absent Present 3- to 5-nerved 7- to 11-nerved 7- to 9-nerved Absent Absent ceolate, flat. Inflorescence a single, terminal and lax panicle or a terminal and several axillary pan- T ple an elongated, compound arrange- pedicels long, flexuous. Spikelets obovoid to van falling from the pedicels, pilose with long, rigid hairs to glabrous, pale to nearly pur- plish; glumes and lower lemma with 5-9(-11) prominent nerves, gaping and exposing the fertile floret at maturity. G/umes unequal, the lower glume '^ to 3⁄4 as long as the spikelet; upper glume and lower lemma a little longer than the anthe- cium, pointed at the apex. Lower palea conspic- uous, membranous, with or without a male flow- er. Upper anthecium stipitate, ovoid, glabrous, smooth and shiny, indurate; stipe membranous ventrally, indurate dorsally; palea with co pound papillae at the apex (papillae occasionally absent in P. cayennense). Stamens 3; stigmas 2, plumose and purple; /odicules 2, membranous, glabrous. Caryopsis with the hilum punctiform. Embryo less than half the length of the caryopsis. Species of sect. Rudgeana grow in open and sunny places, and are common in savannas of Central and South America and in the cerrado of Brazil; they are frequently found in sandy soils from sea level to ca. 1,500 m KEY TO THE SPECIES OF SECTION RUDGEANA . Panicles terminal, lax; axillary m usu- ally absent; spikelets ` ate i ong, 1.2-2.5 mm wide; stipe prominen mm or longer. 2a. Stipe glabrous pam or sheaths pa- pillose-pilose, with glassy hairs; E 8-3.2 mm long; spikelets 4. w long Plu 2b. Stip pil t ll J? l fst th e — £ to grins but without es pe pini m long; spikelets m lon 3a. Spikelets 7-9 mm long, 2. 122. 5 nn m upper anthecium 4-4.5 m P. Bos P 3b. Spikelets 5.9-6.7 mm long, 1.5- m m upper anthecium 3-3.5 eee e 6. P. vinaceum lb. Panicles terminal and axillary, Pormang x 3.5 mm long, 1-1.3 mm wide; stipe 0.5 mm or shorter. 4a. Viri 3-3.5 mm long; leaf sheaths ually with glassy hairs 5. P. rudgei 4b. Spikelets 2.1-2.8 mm long; leaf sheaths hairs. 5a. Plants perennial; spikelets ig sparsely pilose, 2.6-2.8 m g uo us 5b. Plants annual; spikelets d glabrous, 2.1-2.6 mm lon . P. cayennense ]. Panicum campestre Nees ex Trin., Gram. Pan. 197. 1826. TYPE: “V. sp. Brasil (N. ab Esenb.)." Not seen. P. ise A. Hitchc. & Chase, Contr. U.S. Natl. . 15: 139. 1910. TYPE: Brazil. Minas Gerais: TH Widgren s.n. (holotype, US; fragment of holotype, BAA). Perennial, 30-74 cm tall, usually with thick adventitious roots. Cu/ms erect or geniculate at the base, rooting or not at the lower nodes, many- branched; internodes cylindrical or compressed, 3-10 cm long, hirsute with appressed, rigid hairs to glabrescent; nodes densely pilose, with long and whitish hairs. Leaf sheaths 3—7 cm long, usu- ally shorter than the internodes, the lower ones longer, hirsute with long, tuberculate hairs; mar- 472 gins ciliate. Ligule 1.5-2.5 mm long, with hairs on the back towards the base of the blade; collar pilose, stramineous to brownish. Leaf blades lin- ear-lanceolate, 7-25 cm long, 0.5-0.7 cm wide, flat or with involute borders, acuminate apically, rounded or subcordate basally, densely hirsute on both surfaces, with scabrous and ciliate mar- gins, the midnerve manifest. Panicles terminal and axillary, forming an oblong, compound in- florescence, sometimes the terminal panicle dis- tant from the other ones, lax and diffuse; axis sparsely hirsute, at least in the lower portion, longitudinally ridged, flexuous, scabrous, the branches alternate, divaricate, flexuous and sca- brous, the axils of the branches pilose; pedicels long, scabrous. Spikelets ovoid, 2.6-2.8 mm long, 2 wide, sparsely pilose, pale to purple toward the apex to completely purplish. Lower glume 1.8-2.2 mm long, '^—À as long as the spikelet, acuminate to subulate, with rigid hairs toward the apex, 5-9-nerved, the midnerve sca- brous. Upper glume and lower lemma subequal, acuminate, 2.6-2.7 mm long, 7—-9-nerved, sparsely pilose on the inner surface and with or without long and sparse hairs on the outer sur- face. Lower palea elliptic, 1.8-2 mm long, pres- ent or absent. Upper anthecium broadly ellip- soid, 1.8-2 mm long, 1 mm wide, pale; lemma 7-nerved; palea with compound papillae at the apex; stipe with the membranous and indurate portion ca. 0.2 mm long. Caryopsis 1.2-1.3 mm long, 1 mm wide. In flower December-May. Fig- ure 4 Distribution. Brazil, from Para and Bahia to Paraná; 0—1,500 m; growing in sandy or red clay soils in campos or cerrados. Selected specimens examined. BRAZIL. BAHIA: Ser- ra Geral de Caitité, 9.5 km S of Caitité on road to Brejinhos da Ametistas, Harley 21319 (CEPEC). DISTRITO FEDERAL: Universidad de Brasilia, Clayton 4795 (NY, US), 4842 (MO, NY, US); E of Lagoa Par- anoa, Irwin et al. 11181 (F, GH, NY, US); 15 km S of Brasilia, Irwin & Soderstrom 5700 (US); Sobradinho, Clayton 4875 (NY). Goias: 6-7 km E of Alto Paraíso, Anderson 6515 (MO, NY); 26 km NE of Catalào, Irwin et al. 25210 (F, MO, NY, US); 75 km N of Corumbá de Goiás, Irwin et al. 19000 (F, GH, MO, NY, US); 14 km S of le Irwin et al. 34386, 34387 (F, NY, US); 16 km N of São Joao da Aliança, Dawson 14442 (US); between Viannápolis and Ponta Funda, Chase 11315 (US); Serra do Rio Preto, 14 km E of Cabeceiras, Irwin et al. 10354 (US); Corumbá, Macedo 4482 (BAA, US). MATO GROSSO: | km NE of Garapé, 1 Oct. “es "Flee be ee s.n. (US-2642542). MATO G rande, Chase 10790 (GH, RB. "US. poi 9598 (US). MINAS GERAIS: Bar- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 bacena, Serra Mantiqueira, Chase 8645 (F, GH, NY, RB, US); Corinto, Fazenda Diamante, Mexia 5567 (F, : 5 11 km N of Gouveia, Anderson et ine Faria, Serra da Bocaina, Chase m W of Barao de oo Irwin et (F, MO, S US); 7 km W of Campanha, Davidse et al. 10651 (MO, NY); 33 km NE of Francisco Sá, Irwin et al. 23071 (F, MO, NY, SP); el km SW of Gouveia, Anderson et al. 35148 (F, MO, NY); 9 km NE of Estiva, Davidse et al. 10544 (MO); Ouro Preto, Chase 9354 (F, NY, US); 2 km S of Itacolumy, Irwin et al. 29364 (F, MO); Pocos de Caldas, Chase 10637 (US); Piloes, Macedo 4876 (NY, US); lower slopes of Chase 8856 bus pire Chase 8815 (US), Maia 18 (RB); Jardim, Widgren 908 (US); Lagoa Santa, Chase $995 (US); Hermilo Alves, Duarte 6352 (US). PARA: Marajó ate Natal, Schwacke 62 (R); Fazenda d js Rio Aurá, Black 54-16109 (R). PARANA: Jagua iahyva, Dusén 16393 (F, GH, NY, US), 10074 (US), Swallen 8678 (US); 2 km W of Rio Itararé and road PR-11, Davidseet al. 11375 (MO, NY). RIO DEJANEIRO: Monte nin Serra da poss = hase 8358 (F, GH, MO, NY, US); Resende, Hoe US). SAO PAULO: ps N of recon C NY, SP, US); 3 km from Cajurü, Sendulsky 126 (SP, US); 16 km NNE of Padua Sales, Eiten 1669 (NY). Although it was not possible to examine the type specimen of this species, P. campestre is clearly differentiated by the diagnosis given by Trinius and by the illustration of this same au- thor (1829). Nees (1829) published tł same name, the type specimen being completely different from the species described by Trinius. Panicum campestre Nees of 1829 was validly renamed as P. peladoense by Henrard (1940). pecies with the 2. Panicum cayennense Lamarck, Tabl. Encycl. 1: 173. 1791. TYPE: “Cayenne, D. Stoupy” (holotype, P, not seen; fragment of holotype, BAA, US-81397) P. sessilicaule Desv., Opusc. 95. 1831. TYPE: “Habitat in Carolina” (holotype, P; fragment of the holo- A type, BAA). P. floribundum Rich. ex Lam., Encycl. 4: 742. 1798, pr . P. ca nse. P. pedunculare Willd. ex Steudel, Syn. P1. Glum. 1: 77. YPE: “P. cayennense ie Agr. Bras. 195. Brasil” (fragment of the syntype: “America me- ridionale, from Humboldt,” US. 2907507). P. g var. curtatum Doell, in C. Martius, Fl. s. 2(2): 220. 1877. TvPE: "extra fines in via imer Cayenne et Baduel (Yelski, inter plantas a tafinski D mecum communicatas)" ues US-80517 1987] Annual, to 110 cm tall. Culms erect or spread- ing, usually branching at the lower and middle nodes, often zigzag, few-noded; internodes hol- low, compressed, hispid to glabrous; nodes dark, covered with whitish hairs. Leaf sheaths 2-8 cm long, shorter or longer than the internodes, pi- lose, with thick, tuberculate hairs; margins cil- iate. Ligule 0.8—1.6 mm long; collar pilose, pale. Leaf blades linear-lanceolate, 5-28 cm long, 0.4— | cm wide, flat, acuminate apically, rounded at the narrowed base, hispid on both surfaces to glabrescent, the margins scabrous and ciliate, the midnerve manifest. Panicles several, terminal and from the upper leaf axils, forming an elongated compound inflorescence 5-32 cm long, 3-12 cm wide, reaching 73 to almost the entire height of the plant, each panicle included at the base; axis longitudinally ridged, flexuous, scabrous and his- pid towards the base, the branches divaricate, alternate to opposite, sometimes pseudoverticil- late, scabrous and flexuous, the axils of the branches pilose to glabrous; pedicels long, flex- uous and scabrous. Spikelets obovoid, 2.1-2.6 mm long, 1.1-1.3 mm wide, glabrous, ¿lobos pale to purplish. Lower glume 1.2-1.8 mm long, more than half the length of the spikelet, acu- minate apically, pilose on the inner surface, 5-nerved, the midnerve scabrous toward the apex. Upper glume 2.2-2.5 mm long, acute apically, 7-nerved, pilose to glabrous on the inner surface. Lower lemma 2-2.4 mm long, 7-nerved. Lower palea elliptic, 1.6-1.9 mm long, 0.6-1.1 mm wide, membranous, glabrous; male flower absent. Up- per anthecium broadly ovoid, 1.5-1.8 mm long, 0.9-1.2 mm wide, pale; stipe less than 0.3 mm long, the indurate portion prolonged beyond the upper anthecium as a mucro. Caryopsis broadly ovoid, 0.9 mm long, 0.7 mm wide, pale. In flower all year. Distribution. Mesoamerica, West Indies (Cuba, Jamaica and Dominican Republic), and South America, from Venezuela to Bolivia; 0— 1,500 m; occurring in savannas, in sandy or clay soils. Chromosome number. n = Pohl, 1974). 27 (Davidse & Selected Vae aee examined. MEXICO. CHIAPAS: IZABAL: S of Río Dulce, at Shell Station, LeDoux et at ZULOAGA — PANICUM SUBG. PANICUM SECT. RUDGEANA 473 6 (NY). PETEN: Santa Rita, 20 km al S de Santa Elena, 5 (F, ALAJUELA: 601 (F, US), 605 (F). PANAMA. CHIRIQ Boquete, McDaniel 6807 (MO); vicinity of David, Hitchcock 8372 (F, MO, NY, US). P A: Near Ar- raijan, Woodson, Jr. 1402 (MO, NY. US) pia ISLA DE PINOS: Near Nueva Gerona, Curtiss 267 (MO, US); vicinity of San Pedro, Britton et al. 14455 (MO, US); Isla de Pinos, Taylor 34 (MO, US). oRIENTE: Cayo del Rey, Ekman 10028 (US). PINAR DEL RIO: Herradura, Tracy 9073 (US), 9093 (MO), Britton et al. 6520 (US); Sierra de Cabra on Guane road, Britton et al. 7275 (US); Laguna Jovero and vicinity, Shafer 10510 (US). JAMAICA. Halliss Savanna, Upper Clarendon, Harris 12226 (MO). DOMINICAN REPUBLIC. DISTRITO NA- CIONAL: Sierra Prieta, Villa Mella, Liogier 17408 (US). LA VEGA: Vicinity of Pedra Blanca, Allard 16060, 16067 (US); Cordillera Central, Sabana de la Mar, El Valle, Ekman 15694 (US Simba, Fernández 11 (MO). Without department and locality, Mutis 5359, 5378, 5498, 6110 (US). VENEZUELA. AMAZONAS: Puerto Ayacucho, Williams 13085 (F, US); near Capuana, Davidse & Huber 16811 (MO); 23 km NE of Puerto Ayacucho, Davidse & Huber Davidse et al. 4548 (MO). FRENCH GUIANA. Cayenne, Pu hie (MO- ES US- epa 54 ES route de R mbeau, Hoock s. . SURIN n distr Pará, yon 1495 (MO). Cmn . CRUZ: Buena Vista, Steinbach 6935 (BAA, F, GH, LIL, MO, NY US). BRAZIL. AMAPA: Rio Pedra Fróes & Black 17 322 (US); Macapá, Fazendinha, Black & Lobato 50-9659 uu: gente s: 2 km S of HO Prance et al. 8177 (F,G , NY); km 27 of road Humaitá-Porto Vel- p: Prance al. 3517 (MO). BAHIA: Col. Valença, Pinto US). coras: 2 km SW of Araguiana, Eiten 10154 US MATO GROSSO: 20 km S of Garapü, Irwin & Soder- strom 6485 (US). MATO GROSSO DO SUL: 100 km W of Coxim, Bommer 54 (NY, US); Paiaguás, Fazenda Al- vorada, Allem & m 1001 (MO); Xavantina-Cach- imbo road, W of k 3674 te dicia depu a 2- 4 km Eof Mutumparaná, Prance et al. 8831 (F, MO). Panicu m differs from P. campestre mainly by its smaller, glabrous, and obovoi spikelets. It also differs in its annual habit; in P. cayennense the culms are generally short, branched, and bear numerous panicles nearly 474 from the base, the axillary ones aggregating with the apical ones. Nevertheless, there are some specimens with elongated culms in which the terminal panicles are somewhat separated from the axillary ones. This species was included by A. Hitchcock & Chase (1915) in the Capillaria group, along with P. miliaceum, P. capillare, and others, but the presence ofthe characteristic stipe of sect. Rudge- ana clearly separates it from these species. 3. Panicum cervicatum Chase, J. Wash. Acad. Sci. 32: 164, f. 10, 1942. TYPE: Brazil. Mato Grosso do Sul: Tres Lagoas, 4 Feb. 1930, A. Chase 10737 (holotype, US-1500814; iso- types, RB, US-1816795) Perennial, 40-100 cm tall. Culms erect, simple or occasionally branched; internodes 7-23 cm long, terete, glabrous to sparsely pubescent just below the nodes; nodes densely pilose to gla- brous. Leaf sheaths 7-13 cm long, the lower ones overlapping, pale, densely hirsute to glabrous; margins ciliate. Ligu/e 1.5-2 mm long; collar dark brown, short- to long-pilose. Leaf blades lanceo- late, sf, 16-36 cm long, 0. 8-1. 6 cm wide, long- y, flat or the margins involute i in n drying, hispid or strigose to glabrous on both surfaces, the margins scabrous and largely ciliate with papillose-pilose hairs (these hairs caducous), the midnerve prominent. Panicles lax, diffuse, many-flowered, 25-60 cm long, 12-35 cm wide, the spikelets in pairs; axis longitudinally ridged and scabrous, the branches alternate or opposite, scabrous, the axils of the branches pilose and pale; axillary panicles usu- ally absent, when present similar in shape and smaller than the terminal one; pedicels scabrous, 2-20 mm long, the spikelets set obliquely on the pedicels. Spikelets ellipsoid, 7-9 mm long, 2.1— 2.5 mm wide, glabrous, pale to purplish. Lower glume 3.5-3.8 mm long, acuminate, 7-1 1 -nerved, m long, sparsely pilose to gla- brous, long pilose at the base, the inner surface pilose towards the apex, 7-1 1-nerved, the mid- nerve scabrous. Lower lemma glumiform, 6.2— 7.3 mm long, long pilose at the base, the inner surface pilose, purplish, 7—9-nerved. Lower palea elliptic to obovate, 4-5.8 mm long, 1.3-2.2 mm wide, membranous, the borders pilose; male flower absent. Upper anthecium ovoid to ellip- soid, 4—4.5 mm long, 1.8-2.1 mm wide, at ma- turity 2.5 mm wide and dark brown; stipe ca. 1 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 mm long, somewhat fleshy with an expanded summit and a thick, indurate process on the back, prolonged beyond the base of upper anthecium at maturity; rachilla pilose below the stipe. Cary- opsis 2.8-3.2 mm long, 1.5-2.2 mm wide. In flower December-September. Distribution. Bolivia and Brazil; sandy or sandy-clay savannas, campos or open cerrados; 400-1,300 m Additional specimens examined. BOLIVIA. SANTA CRUZ: Santiago de Chiquitos, San pae Cárdenas 4506 (US); Men esp cerca de El Carmen, Cá d as 4503 (US). BRAZIL. BAHIA: Ro De 305 k SW of Barreiras, Irwin B al. 14657 (MO, NY, SP, US): Espigão Mestre, 100 km WSW of Barreiras, Anderson et al. 36654 (F, MO, NY, US); Rio Roda Velha and highway BR-020, Davidse et al. 12084 (MO, NY). DISTRITO FEDERAL: Universidade de Brasilia, Clayton 4809 (SP), 4839 (NY, SP); 20 km E of Brasilia, Jrwin et al. 9213 (F, MO, NY, US); Brasilia, Belém 1970 (CEPEC); 15 km E of Brasilia, Irwin & 121 93 (NY); between Jatahy and Rio Araguaya, Chase 11736 (US); vicinity of Annapolis, Chase 11519 (US); 38 km N of Sao Jose da Alianga, Dawson 14354 (US); 40 km W of Rio Verde, Chase 11713 (US); W of Santa Rita do Araguaya, Chase 11863 (US); between Vian- napolis and Ponta Funda, Chase 11281 (US). MATO Grosso: Rodovia Cuiaba-Santarem, Lemes 4125 (RB); Rondonópolis, Rio Paguba, Rondon 2566 (RB, US); Diamantina, Weddell 3081 (US). MATO GROSSO DO SUL: Xavantina-Cachimbo road, 85 km from Xavantina, Hunt & Ferreira 5739 (NY, SP, US); NW of Sao Lou- m Chase 11959 (US). MINAS GERAIS: Lagoa Santa, 14 Feb. 1864, Warming s.n. (US); 26 km NE of Pa- ire dk et al. 25582 (F, NY, SP); Serra do Cipó, 110k of Belo Horizonte, Chase 9138 (F, GH, MO, m ead between Sucupira and Omega, S of Uberlandia, Chase 11167 (US); 3-4 km de Prata, Sen- dulsky 18 (SP), 37 (SP, US); Frutal, Valls 649 (US); Caldas, Linc TI 1369 (US); Pratinha, Dorsett 189b S). R Pickel 5887 (US); = Sant à n- dulsky 148 (US); 4 raguacu Paulista, Clay- ton 4596 (SP, US); sp E agai Black 51-11072 (B MARANHAO: orda to Grajahü, Swallen 3648 (RB, SP, US). When publishing this species, Chase described and illustrated the frag ment of rachilla below the sai anthecium, show the two constituent rts. Although she indice that she had not due this character in any other species of =e 1987] the genus, she related P. cervicatum to P. oly- roides Kunth, and in unpublished manuscripts placed both species in the **Olyroides" group; recently, Renvoize (1984) also related P. cervi- catum and P. ligulare to P. olyroides. However, in P. olyroides the characteristic stipe of sect. Rudgeana is absent, and there are long, acintate hairs at the base of the upper anthecium on its ventral face (Fig. 4d). The spikelet is frequently obliquely disposed on its pedicel in P. cervicatum, a character pres- ent also in other species of Panicum (e.g., hirtum). Iconsider one ofthe paratypes, Williams 13221 of Venezuela, to belong to P. vinaceum Swallen. Consequently P. cervicatum remains known only rom Brazil and northern Bolivia. 4. Panicum ligulare Nees ex Trinius, Gram. Pan. 206. 1826. TvPE: *V. sp. imperfectum Brasil (N. ab Esenb.)" (lectotype here designated: floriferous part, LE), non Nees, Agrost. Bras.: 196. 1829. TYPE: “Hab. in campis prope Al- meirim provinciae Paraensis" (lectotype here designated: floriferous part of number 3800, M) Perennial, 1.30-2 m tall, with thick adventi- tious roots and lanose cataphylls. Cu/ms erect, many-noded; internodes 8-24 cm long, solid or hollow, pilose to glabrous; nodes dark, pilose to glabrous. Leaf sheaths 8-23 cm long, greenish to purplish, papillose-pilose, the hairs urticant and caducous; margins ciliate. Ligule 1.8-3.2 mm long, with long hairs on the back towards the base of the blade; collar pale, densely villous. Leaf ioa Tac kapuni 8 30-55 cm long, 0.9— 1.9 cm wide, flat, acuminate apically, subcordate basally, panes pilose on both surfaces to gla- brescen lax, diffuse, many-flowered, 47-65 cm 30 cm wide, the branches spreading; axis lon- gitudinally ridged, pilose towards its base, oth- erwise scabrous, the branches alternate or op- posite, sometimes verticillate at the base of the panicle, scabrous, the axils of the branches pi- lose, pale to brown; axillary panicles usually ab- sent, when present similar to the upper one but smaller; pedicels claviform, 2-20 mm long, sca- brous. Spikelets ellipsoid, 4.4-5.7 mm long, 1.2- mm wide, glabrous, greenish to purplish. Lower glume 2.9-3.8 mm long, ⁄2—⁄4 the length of the spikelet, subulate apically, shortly pilose ZULOAGA — PANICUM SUBG. PANICUM SECT. RUDGEANA 475 towards the apex on the inner surface, 7—9-nerved, the midnerve scabrous. Lower lemma glumi- form, 4.1-4.9 mm long, acuminate apically, pi- lose towards the apex in the inner surface, 5-7- nerved. Lower palea elliptic, 3-3.3 mm long, 0.9- 1.5 mm wide, glabrous, whitish, membranous, the margins with or without short hairs; male flower absent. Upper anthecium ovoid, 2.5-3.2 mm long, 1.1-1.5 mm wide, pale; palea with compound papillae at the apex; stipe conspicu- ous, glabrous, with 1 or 2 wings nearly 0.8-1.1 mm long, the indurate portion 0.4—0.7 mm long. Caryopsis 2.4 mm long, 1.3 mm wide. In flower March-October. Figure 5. Brazil, from Maranhào and Ba- rosso; cerrado; 500-1,100 m. Capim elefante. Distribution. hia to Mato Common name. Additional specimens examined. BRAZIL. BAHIA: 150 km SW of Barreiras, Irwin 14904 (F, MO, US). DISTRITO FEDERAL: Chapada de Contagem, ca. 20 km NE of Bra- silia, Irwin & Soderstrom 5166 (US), Irwin et al. 9653 s: 20 km N of Cristalina, Serra 3700 et al. 21525 (F, US): Serra Dourada, Glaziou 22525 (US); vicinity of Goiás, Chase 11460 (F, GH, NY); 26 Ey By S of Goiania, Davidse et al. 12278 (MO). MA AO: Carolina to San Antonio de Balsas, Swal- len 4094 (US); Serra do Penitente, Miranda 128 (RB). Xava ntina, Irwin et al. 16122 Azul, 77 km from Barra do Gargas, Hunt 6075 (NY, US); Serra do Roncador, 86 km N of Xavantina, /rwin F, NY, US); Xavantina-Cachimbo road, 215 km from Xavantina, Hunt & Ferreira Ramos 5606 Y, atter et al. 2090 (NY, RB); Campos Novos, Kuhl- mann 1745 (RB). Trinius (1826), in attributing P. /igulare to Nees, described the species as possessing a lan- ceolate, membranous ligule 6-10 mm long, and used the epithet /igu/are in reference to this char- acter. After examining abundant material of P. ligulare and studying the type of P. /igulare in Leningrad, I discovered that the type sheet con- tains a mixture of material. The panicle of this specimen does correspond to what I consider P. ligulare (which agrees with the description given by Trinius for the floriferous part), but the veg- etative portion (which is separated from the flo- riferous part) is markedly different from the veg- etative parts of the species. The leaf sheaths and leaf blades are completely glabrous, and the membranous ligule is exceptional because of its 476 size. This type of ligule has never been found in any species of Panicum up to now. In 1829, Nees described P. /igulare as Trinius did, mentioning that the type of ligule he ob- served was unique in Panicum. In his descrip- tion, Nees reported the type locality as “Hab. in campis prope Almeirim provinciae Paraensis." On studying the type material in Munich, I found two specimens collected by Martius in that lo- cality, one with the number 3798 (attached to the plant) and the other identified as 3800. In 3800 there is a mixture of material similar to the specimen from Leningrad. Specimen 3800 is un- doubtedly the one Nees used in his diagnosis. In specimen 3798 there is no mixture, and it fits perfectly with what I have described as Panicum ligulare. In this specimen there is a note on which Trinius stated that this material is different in its vegetative parts to the one examined at Lenin- grad. Trinius (1835) and Steudel (1855) treated the species in the same way as Nees and Trinius did before Doell (1877), in Flora Brasiliensis, noted the difference between specimens 3798 and 3800 of Munich. He considered 3800 to be P. ligulare “in sensu strictiore," but erroneously judged 37 98 to be P. virgatum (a completely different North American species I select the floriferous portion ofthe Leningrad material as the lectotype of Panicum ligulare Nees ex Trin., and the floriferous portion of the Mu- nich specimen 3800 as a lectotype of P. ligulare Nees. 5. Panicum rudgei Roemer & Schultes, Syst. Veg. 2: 444. 1817. Based on P. scoparium Rudge, Pl. Guian. 1: 21, pl. 29. 1805, non Lam., 1798. TYPE: “Panicum scoparium Rudge, ex herb. Rudge" (fragment, US-2830540). P. ior e Roth ex BERS ERER Syst. Veg. 4 817. TYPE: “Roth nov. plant Spec. Ms. ... In Essequebo, Mertens” ea at US- 2830939). P. asl var. brasiliense Raddi, , Agrost. Bras. 48. 1823. nse ; P. riiophtlum Mns Syn. Pl. Glum. 1: 1855. s Salzm. Hrbr. Bahia" sss, US-4 w P. cayennense var. divaricatum Doell, in C. Martius, Fl. Bras. 2(2): 220. 1877. TYPE: same as the species. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Perennial, 30-130 cm tall, with short rhizomes and pilose, scaly cataphylls. Cu/ms decumbent or geniculate to erect, often zigzag, rigid, hollow, branching from the lower and upper nodes, many- densely to sparsely pilose; nodes covered wit whitish, appressed hairs to completely glabrous. Leaf sheaths 4-13 cm long, densely papillose with thick, glassy hairs; margins ciliate. Ligule 1.5-2 mm long, with long hairs on the back towards the base of the blade; collar pale, densely to sparsely pilose. Leaf blades eee 20- 40 cm long, 0.6-1.1 cm wide, acuminate apically, narrowed Tai flat or with involute borders, densely hispid to sericeous on both surfaces to glabrescent, the margins scabrous, ciliate or not in the lower portion, the midnerve manifest. rescence !^ or more the length of the plant, 25- 50 cm long, 10-20 cm wide; axis and branches longitudinally ridged, scabrous to pilose, the ax- ils of the branches long-pilose to glabrous, brownish to pale, the branches alternate and di- varicate, somewhat flexuous; pedicels d long-pilose, flexuous. iii ovoid, acum nate, 3-3.5 mm long, I-1. m wide, pale i nearly purplish, sparsely ous with stiff hairs irregularly distributed. Lower glume 2-2.7 mm long, 7^ as long as the spikelet, acuminate api- cally, with stiff, whitish hairs on the upper part to completely pilose, the inner surface pilose, 3- 5-nerved, the midnerve scabrous. Upper glume 2.7-3 mm long, acuminate apically, pilose on the inner surface, 7—9-nerved, the midnerve sca- brous. Lower lemma 2.5-2.9 mm long, acute api- cally, long-pilose to glabrous, 7—9-nerved, the midnerve scabrous. Lower palea elliptic, 1.8-2.3 mm long, 0.5-1.1 mm wide, membranous, the margins shortly pilose; male flower present, the anthers purplish; rachilla with or without whitish hairs. Upper anthecium ellipsoid, 1.8-2.2 mm long, 0.8-1.1 mm wide; palea with compound papillae towards the apex; stipe with the mem- branous portion ca. 0.4 mm long, the indurate portion 0.5 mm long, prolonged beyond the back of the lemma as a mucro. Caryopsis pale, 1.5 mm long, 1 mm wide. In flower all year. Distribution. Mesoamerica, West Indies (Ja- maica, Trinidad) and South America from Co- lombia to Bolivia and Brazil; 0-1,000 m; in open savannas, campos or cerrados, usually in sandy ils 1987] Common names. carricillo (Venezuela). Chromosome number. n= 9 (Davidse & Pohl, 1974, 1978); 2n = 18 (Pohl & Davidse, 1971) makuna-ta (Colombia); Selected specimens examined. MEXICO. TABASCO: Achotal, Matuda 3087 (F, GH, US). GUATEMALA. IZA- 39673 (F). BELIZE. Cabbage Hall, Dwyer et al. 454 (F. MO); Swasey Branch, Monkey River, Gentle 3862 (F, GH, MO, NY, US); Machaca, Gentle 6893, 6923 (F, NY, US). Costa RICA. ALAJUELA: Buenos Aires, León 1184 (US), Tonduz 3679, 4875 (US); Los Palmares, Pittier 10588 (US). PUNTARENAS: Buenos Aires, Molina 2739 73 , MO, US); east of CIA, road to Buenos Aires, Pohl 13116 (F, MO). SAN JOSE: Vicinity of General, Skutch 3065 (GH, MO, Pittier 12064 ). NICARAGUA. ZELAYA: Entr iu a y Limbaikán Seymour 497 : n Mea Jaboga, Killip 4163 s Canal Zone, ne rt Randolph, Standley 285 98 (MO, US); Perlas eee San Jos and, John- 324 (GH, HL sami Clar- endon, Harriss 12845 (NY, US); Halliss Savanna, Up- don, Harriss 12235 (MO, NY, US); Bunkers S. ; Mason River O, NY, US); St. Joseph, Hitchcock abadie, Britton 688 Hitchcock 10083 (M 10181 (US); Piarco d So Pan de Azúcar, Orozco et al. 7 Guainía, Puerto Colombia a et al. 17936 (US) META: 73 km W of Las Gaviotas, Davidse 5390 (MO); 43 km NE of Puerto , Davidse Wilches y au “a km 16 Killip & Smith T (F, GH, MO, NY, US). ToLIMA: El Con W of San Lorenzo, PD 3509 GH, MO. ‘NY. US). VAUPES: hans I Circasia, Eb 7201 (US). VICHADA: 25 k of Cumaribo, Davidse 5325 (MO); 10 km W of Las “sasa Davidse 5367 (COL, MO, NAS: Cerro Duida, Maguire 29424, 29060 (NY); Serrania Parü, Cowan 31486 (NY, US); 20 km S of Puerto Ayacucho, Davidse 2641 (MO); 5 km NE of San Carlos de Río Negro, Liesner 3703 (MO); 25 km S of Samariapo, Gentry & Berry 14600 (MO); Yavita, Williams 13879 (F, US); pie del Cerro Huachacamari, Huber 4990 (MO); El Manguito, 1 km N of Cafio Caname, Davidse et al. 17482 (MO); alrede- dores de Canaripo, Huber 1981 (MO); 8 km S de Puerto Ayacucho, Davidse & Huber 14916 (MO). ANZOATEGUI: Vicinity of Santo Tomé, Chase 12841 (GH, US). APURE: end of the — de Cinaruco, Davidse & González 14667 (MO); nea Fundo El Algarrobo, Davidse & ar den 14217 MC. BARINAS: 16 km SW of the Merida rsa dg just outside of Barinas, Da- vidse 3182 (M raima, Steyermark 59429 (F, US); 0.5 km NE of ZULOAGA — PANICUM SUBG. PANICUM SECT. RUDGEANA 477 ides Steyermark & Wurdack 22 (F, NY, US). ONAGAS: E de Maturin, ca. caserio La Pica, Ariste- iun 4048 (F, MO, NY); Laguna Most, 12 km N de Capirito, Trujillo 14194 (F). ZULIA: 60 km NW of Santa Bárbara-San Carlos del Zulia, near Campamento El Rosario, de Bruijn 1473 (MO, NY, US). GUYANA. Tu- matumari, See os 40 (GH); Waini River, de la Cruz (F, Waramuri Mission, Horuka e Tu Hills Estate, Hitchcock 17191 (F, MO, NY, yea Kaie- St. Laurent ERU. LORETO: , Rio anon, Gentry et al. 29965 (MO); via Nauta- jn E Díaz & Jaramillo 1270 (MO). BOLIVIA. BENI: 15 km de erin, camino a Riberalta, Krapovickas & Bator 35068 (U 9). LA PAZ: San Carlos, Buchtien : polo, Williams 1020 (NY); San Antonio, Buchtien 1 15 9 (US) (CEPEC, MO, NY); Marau, Belém & (CEPEC), Zuloaga et w deg (RB 418, 16 km del cruce A- 01. (CEPEC, NY). MATO GROSSO: Serra Azu Xavantina, Irwin et al. 17302 (F, MO, NY, US); 270 km N of Xavantina, Ratter 2069 (NY s dos pn iion: antina, Irwin et al. 15961 (F, US). M I DO SUL: Tres Lagoas, Pen por (US) P PARA: Santarém, fisv 3721 (US); Soure, Ilha do Marajó E a Acara, Thomé Assu, Mexia 592 io US); 73 km NE se eid FUE "E 17939 (MO. NY); 17 km SE of Vigía, along road Pa-140, Davidse et al. 17610 (MO, NY). PERNAMBUCO: Vicinity of Re- cife, Chase 7675 (F, US), Poazeves, Pickel 3137 (US). DE JANEIRO: Silvestre, Holway et al. 1116 (US); Merity, 20 km N of Rio de Janeiro, Chase 8465 (US). RONDONIA: Porto Velho, Black & Cordeiro 52- 15348 mann 4110 (US); 7 km de São José dos Campos, Eiten & Mimura 3351 (MO, US). 6. Panicum vinaceum Swallen, Fieldiana. Bot. 28(1): 27. 1951. TYPE: Venezuela. Bolívar: Gran Sabana, between Kun and waterfall at Rue-Meru, south of Mount Roraima, elev. 1,065 m, 2 Oct. 1944, J. A. Steyermark 59173 (holotype, US-1911661; isotype, F). Perennial, 40-100 cm tall. Culms erect, few- noded; internodes 6-11 cm long, pilose; nodes 478 pilose. Leaf sheaths 4-14 cm long, covered by long dense hairs or glabrescent; margins ciliate to glabrous. Ligule 0.6-2 mm long; collar pale, pilose. Leaf blades linear-lanceolate, 15-42 cm long, 0.5-1.2 cm wide, flat, acuminate apically, subcordate basally, with appressed hairs on both long, 6-20 cm wide, the branches spreading; axis longitudinally ridged, scabrous, the branches al- ternate to opposite, scabrous, the axils of the branches pilose, pale; axillary panicles usually absent, when present similar to the terminal one but smaller; pedicels scabrous. Spikelets ellip- soid, 5.9-6.7 mm long, 1.5-2 mm wide, globose and glabrous, pale to purplish, the inner surface ofthe glumes and lower lemma densely to sparse- ly pilose. Lower glume 2.9-3.8 mm long, subu- late apically, 5-9-nerved, the midnerve scabrous. Upper glume 5.3-6.4 mm long, acuminate api- cally, 7-1 1-nerved, glabrous. Lower lemma 5.1— 5.5 mm long, acute apically, 7-9-nerved. Lower palea obovate, 3.1-4 mm long, 1-1.5 mm wide, whitish, membranous, the margins pilose; male flower absent. Upper anthecium ovoid, 3-3.5 mm long, 1.4—1.8 mm wide, pale; stipe with the mem- branous portion 0.7-1.2 mm long, with or with- out wings, the indurate portion ca. 0.9 mm long, obtuse; rachilla pilose below the upper anthe- cium. Caryopsis 2.4 mm long, 1.5 mm wide. In flower September-A pril. Distribution. Brazil and Venezuela; 100- ,000 m; savannas. Additional specimens examined. BRAZIL. GOIAS: Rio da Prata, 6 km Posse, Irwin et al. 14509 (US). PARA: Serra do Cachimbo, BR-163 Cuiabá-Santarém, km 823, Prance et al. 24993 (MO, NY); Serra do Ca- Y, US); Estación Bo- livar, en sabanas de Santa P Tamayo 2964 (MO, NY, US). This species is closely related to P. cervicatum, from which it can be separated only by the sizes of the spikelet, lower palea, and upper anthe- cium. The rest ofthe differential characters noted by Swallen (size and pilosity of the plants and size of the panicles) have no value in separating the two species. Irwin 14509 is exceptionally large and differs from Swallen's description. Never- theless, spikelet size (a constant character in the ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 scant material available for this species), shows that Irwin 14509 must be included in P vina- ceum, thereby partially di s con- cept of this species. LITERATURE CITED BERG, R. Y. 1 traliense (Poaceae): taxonom plications. Austral. J. Bot. Brown, W. V. 1977. The Kranz syndrome and its subtypes in grass ee Mem. Torrey Bot. Club 23(3): (ii + 7. DavipsE, G. Fruit pee and seedling iones ment in the Poaceae. Proceedings of the tional Symposium on Grass Systematics ind Evo. lution. Smithsonian Contributions to Botany (in press). 985. Spikelet structure in Panicum aus- c and ecological im- 3: 79-583. W. POHL. 1974. Chromosome numbers, meiotic behavior, and notes on tropical American ine i a Canad. J. Bot. 52: 317-328. . Chromosome numbers of a "nd grasses (Gramineae): 5. Ann. Missouri Bot. Gard. 65: 637-649. DoELL, J. C. 1877. Tribe 3, Paniceae. In C. Martius aa Flora Brasiliensis 2(2): 33-342, pls. 12- eee J. TH. 1940. Notes on the nomenclature of some grasses. II. Blumea 4: 503-507. HirTCHCOCK, A. S. 1915. Panicum. North American Flora 17: 198-289. & A. CHASE. 1910. The North American species of Panicum. Contr. U.S. Natl. Herb. 15: 6 & — 1915. Tropical North American species of Panicum. Contr. U.S. Natl. Herb. 17: 35. LAZARIDES, M. 1959. The Australian species referred to Ichnanthus Beauv. (Gramineae). Austral. J. Bot. 7: 328-346. & R. D. WEBSTER. 1984. Yakirra (Paniceae, Poaceae), a new genus for Australia. Brunonia 7: 289-296. NEES VON ESENBECK, C. G. D. 1829. Agrostologia Brasilien A .... Stuttgart & Tübingen. (This is .l, Gramineae, of Flora Brasiliensis seu oer tara .., edited by C. F. T von Martius.) G. Davipse. 1971. Chromosome numbers of Costa Rican grasses. E 23: 293- 324. RENVOIZE, S. A. 1984. The Grasses of Bahia. Royal Botanical Gardens, Kew, Englan SHAW, R. . D. WEBSTER. 1983. Characteristics ofthe (Poaceae: Pan- iceae). "Bot. jm iced 144: 363-370. e UDEL, E. nopsis plantarum Gram earum. In Synopsis Banta Glumacearum " -47 usns C. B. —-1836. Species Graminum ico- nibus et DUM DER UA 3 volumes, 360 plates. Leningrad, U.S.S.R 1826. De Graminibus Paniceis: Dissertatio botanica altera. 291 pages. Impensis Academiae Imperialis Scientiarum. St Petersburg, U.S.S.R. ARUNDOCLAYTONIA, A NEW GENUS OF THE STEYERMARKOCHLOEAE (POACEAE: ARUNDINOIDEAE) FROM BRAZIL! GERRIT DAVIDSE? AND R. P. ELLIS? ABSTRACT i seni ae i: NE iaa Davidse & Ellis, gen. et sp. nov. is described from Amazonian campinas i a, Bra e second genus | quien asiy lt odis aban it is bor ie b a caespitose growth habit, proliferation. and lignification of the numerous basal culm unisexual iran ag iin inflorescences Haas into a false panicle, 3—9-flow er, 3-flowered female spikelets with only the middle floret ertile and its palea l fusiform caryopsis with an elliptic- -punctate hilum. Anatomically this species is charact isodiametric chlorenchyma, and 2 bundle het we is ternodes, normally developed leaves - spiral phyllotaxy, red male spikelets by the absence of stomata and Edu reduction jajn silica bodies, thick epidermis, and hypodermal ows. Its classification in Steyermarkochloeae is based primarily on the morphology of the aa During 1974 an unusual grass was collected by William R. Anderson and associates in Pará, Brazil. Although it was recognized as an unde- T taxon, the inflorescences were too im- mature to show the exact morphology of the doe In 1979 Cleofé E. Calderón and co- workers collected abundant mature material of a same taxon in Amazonas, Brazil. Our study of both these collections indicates that they rep- resent the second genus ila lan — Š again dse & Ellis, 1984). We are naming the zenux in pid of = W. D. Clayton, eminent agrostologist at the Royal Bo- tanic Gardens, Kew, who has made and contin- ues to make outstanding contributions to agros- tology. The compound generic name at the same time refers to the arundinoid affinity ofthe genus. The specific epithet alludes to the strongly dis- similar male and female inflorescences and spikelets. DESCRIPTION x asi ivan salen aig eae & Ellis, gen. et sp. nov. TYPE: Brazil. Amazonas: Trans- amazon Highway. ca. "53 km W of the Ari- puafia River, abundant dominant plant of the vegetation. Growing in a white sand soil “campina.” This plant grows in large, open areas mixed with shrubs and alternating with narrow strips of islands of low tree forest. Most of the population reduced to burnt bases. These trunks look like big candelabra, some ca. 70 cm or less. From them come up solid stems with thickened bases formed by aerial roots. In many cases from the top of burnt trunks, bunches of leaves start com- ing again. Few plants still blooming. Plants ca. 2-3 m tall when flowering. 28 June 1979, alderón, O. P. Monteiro & J. Guedes R SP. US (mounted as 11 sheets)]. Figures Gramen duced culmi internodiis numerosis in- ferioribus he is; phyllot sam; spicula glumis disarticulates; glumae 2; spiculae masculinae 3- ! We extend our gratitude to the late Dr. Thomas R. Soderstrom, Smithsonian Institution, who made the vii uid Calderón et al. enia and photographs available to us and seful review comments by Dr. uch appreciate the ve still dise with him o do Cuchimbo: ri Botanical Garden who encouraged us in our studies. We Steve Renvoize, but, pending additional m for drawing eid tribal classification of Steyermarkochloeae. We thank Jo r. William R. Anderson, University of Michigan, for information about his collecting itinerary , P.O. Box 299, St. Louis, Missouri 63166, U.S.A. 3 Ry Research Institute, Private Bag X101, Pretoria 0001, South Africa. ANN. Missouni Bor. GARD. 74: 479—490. 1987. 480 9-florae palea 2-carinata; stamina 2; spiculae femineae stigmate 2; caryopsis fusiformis-teres hilo punctato. Perennial 2-3 m tall, erect. Vegetative culms usually densely covered for 2-70 cm to a thick- ness of 1.5-6 cm by aerial roots tightly appressed to the culm and by remnants of leaf sheath bases; internodes numerous, 2-15 mm long, 1-1.5 cm diam., solid, lignified; nodes bearing one prom- ta intravaginal near t asal cluster o Flowering culms to 1 cm diam., consisting of many, often elongated internodes; internodes 1- 16 cm long, glabrous, densely waxy when young, green in the exposed portions when older, hol- low, gradually becoming solid toward the base ofthe plant; nodes glabrous; branching primarily intravaginal, profuse in the upper !^ of the culm to form a false inflorescence. Leaves primarily clustered toward the base, those of the flowering culms fewer and gradually reduced in size toward the tip of the culm. Basal leaves with the sheaths densely overlapping, much longer than the in- ternodes, stramineous, long persistent, turning brown and eventually reduced to fibers in age, rounded and glabrous abaxially, without a dif- ferentiated midrib, the margins glabrous, free to the base, the base pilose at the point of insertion and between the veins or glabrescent, the apex ciliate with hairs 2-4 mm long, wider than the base of the blade, rounded, a collar not clearly differentiated; ligule a ciliate membrane 0.9-2.1 mm long, the membrane 0.3-0.9 mm long, the cilia 0.5-1.2 mm long; blades 45-80 cm long, 8- 16 mm wide, flat with involute margins or en- tirely involute, the upper portion always involute and the apex pungent, the abaxial surface green, glabrous and smooth, the adaxial surface grayish green, densely and minutely scabrous, grooved between the veins, the veins approximately the same size, a midrib not differentiated, the mar- gins ciliate with hairs 2-3 mm long in the lower l^. scaberulous in the upper 7^. Cauline leaves similar to the basal leaves but smaller, the up- permost much reduced with the blade shorter than the sheath and entirely involute. /nflores- cences numerous, borne on axillary, exserted pe- duncles, aggregated into a false panicle, unisex- ual, consisting of hemispherical clusters of 7-20 l or 2 series florescences produced before the female, 9-13 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 mm wide, 6-11 mm high; female inflorescences 20-36 mm wide, 15-23 mm high; peduncles ge- niculate and pilose at the base with hairs 0.5-1.5 mm long, sometimes with a line of pubescence or with a scarious bract 1-3 mm below the cluster of spikelets, always subtended by a sheath, the sheath terminating in a sharp point; peduncles of the male inflorescences usually longer than those of the female inflorescences. Female spike- lets 7-19 mm long at anthesis, sessile or short- pedicellate with pedicels to 0.5 mm long, lan- ceoloid, rounded on the back, disarticulating be- low the glumes, falling as a unit, slightly curved, 3-flowered; glumes 2, unequal, herbaceous, shorter than the lemmas, ovate, broadly acute, pilose at the base, otherwise scaberulous, the nerves free or connected by cross-veinlets, the wer 1.5-2.0 mm long, 1-3-nerved, the upper 2.2-3.5 mm long, 3-5-nerved; lower floret with- out a flower, the lemma 3.0-5.1 mm long, 7-9- nerved, ovate, broadly acute, pilose at the base, otherwise scaberulous, cross-veinlets few, the pa- lea absent or rudimentary, 0.7-1.5 mm lon hyaline when present; middle floret unisexual, ma 5 rved o conspicuous cross-veinlets, ovate, acute, pilose at the base and between the nerves just above the base with hairs 1-1.5 mm long, otherwise scaberulous, the palea conspicuously longer than the lemma, 7.5-17 mm long, 9-13-nerved, slightly curved in the upper half, convolute, shal- lowly grooved on the back, spongy-thickened, smooth and shiny in the lower '2—*4, herbaceous and scaberulous in the upper '4—'/, ciliate on the overlapping margin at the base with hairs 1-1.5 mm long; upper floret rudimentary or consisting ofa single 3-nerved bract, 0.1-4 mm long, ciliate at the base, borne on a prominent rachilla 3.5— 10.5 mm long, the floret and rachilla fitting into the palea groove of the middle floret; lodicules absent; staminodia absent or present as an an- terior pair of rudiments to 0.2 mm long; gynoe- cium cylindrical, the ovary wall free from the ovule, the style one, dividing into 2 inconspic- uously plumose stigmas slightly below the tip of the middle palea, the stigmas 2.5-4 mm long, terminally exserted through an apical, iain orifice formed by the convolute palea; caryopsis fusiform-terete, narrowing apically, 6-7 mm long, 0.8-1.2 mm diam., glabrous, the embryo J49-^,, as long as the caryopsis, the hilum elliptic-punc- tate. Male spikelets 3.5—7.5 mm long, sessile or short-pedicellate with pedicels to 0.5 mm long, rounded on the back, disarticulating below the 1987] glumes, 3—9-flowered, the florets (except the up- permost) bearing flowers, the middle florets slightly larger than those above or below, the uppermost usually rudimentary; glumes 2, un- equal, shorter than the lemmas, membranous, ovate, erose, truncate or obtuse, pilose at the base, usually with cross-veinlets, the lower 1.3— 2.1 mm long, 1—3-nerved, the upper 1.8-2.5 mm long, 3-nerved; lemmas similar to the glumes in pubescence, shape and texture, 2.6-4.1 mm long, shorter than the paleas, 3—9-nerved; paleas 3.2- 5.8 mm long, broadly obtriangular, truncate, sometimes erose, 2-keeled (each keel with a nerve), the base with hairs 1-1.5 mm long, the back sulcate, the keels ciliolate, the margins over- Wine lodicules absent; stamens 2, one situated side of the sulcate palea, terminally Heer through an opening formed by the over- lapping palea margins, the filaments separate, ba- sifixed, the anthers 2.2-2.9 mm long. Paratypes. BRAZIL. PARA: Alto Tapajós, Rio Cu- ruru, northwest edge of Serra do Cachimbo, 25 km by Santos & R. Souza 10950 (MO, NY, UB). MORPHOLOGICAL OBSERVATIONS Arundoclaytonia dissimilis when fully mature and undisturbed by fire has an unusual appear- ance caused by the thick accumulation of leaf sheath bases and adventitious aerial roots (Figs. 1, 3). Such plants in the aspect of their basal parts are more reminiscent of certain species of Vel- lozia. This unusual appearance is accentuated after the plants have been moderately or severely burned (Figs. 2, 4, 5). Although the plant is fundamentally a tussock plant, the dense cluster of leaves, which is nor- mally basal in a tussock grass, is raised up to 70 cm above ground level in older plants of Arun- doclaytonia (Figs. 2-5). These small “trunks” re- sult from the proliferation of numerous, short internodes in the basal portion of the culms. Short basal internodes are typical of grass culms. What is unusual in Arundoclaytonia is their large num- ber, thickness, woodiness, and perennial dura- tion. Annual culms characterize most grasses. In typical caespitose perennial grasses the short basal internodes perennate and bear the buds from which new tillers are produced for the new growing season. It is this region of the culm that DAVIDSE & ELLIS— ARUNDOCLAYTONIA 481 is much elongated by the proliferation of inter- nodes and gives Arundoclaytonia its trunklike es, ei Arundoclaytonia. In contrast, the elongated in- ternodes of the flowering culm produced above the cluster of basal leaves gradually become hol- low, as is common among grasses. When the sheath bases and mass of aerial roots have been removed from the lower portion of the culm (as may happen after severe burning and the subsequent wearing off of the root and sheath remains), it becomes apparent from the position of the axillary buds that the leaf ar- rangement is not distichous. Every sixth node bears a solitary, prophyllate, dormant bud (Fig. 7A) that occurs in the same relative position as the buds five nodes above and below it. Since two complete turns around the culms must be made to attain the same position, phyllotaxy is 2/5. The arrangement of the spikelet bracts ap- pears to be nearly distichous; however, the rel- ative position ofthe bracts is much more difficult to observe because the very short internodes of the spikelet and the broad bases of the glumes and lemmas obscure the exact point of insertion of these spikelet parts. Distichous phyllotaxy is characteristic of the Poaceae (Arber, 1934: 282; Barnard, 1964: 47). Only one other exception has been reported: spiral phyllotaxis in Micraira sub- ulifolia F. Muell., a mosslike plant from Queens- land (Watson & Dallwitz, 1980: 89) Branching occurs near the base of the plant to orm the main culms that constitute the bulk of the tussock (Figs. 4, 5). These branches originate from the buds illustrated in Figure 7A. Branching is very infrequent in the middle portion of the culms but profuse in the upper portion. At each upper node, a smaller axillary branch is produced which itself is rebranched several times (Fig. 8) into branchlets terminating in inflorescences. The branching pattern is the relatively simple one _ 5 era. A prophyllum is the first foliar organ pro- duced at the lowest node of each branch (Fig. 8). Each prophyllum is many-nerved and promi- nently two-keeled with narrow wings on the keels. Buds at subsequent nodes on the branch are sub- ended by leaves with blades reduced and gen- Just below the inflorescences they are reduced to scarious bracts, presumably representing re- duced sheaths only 482 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURES 1-6. Habitat and habit photographs of Arundoclaytonia dissimilis; E W of the Aripuana River, Nerio] Brazil.—1. Unburned campina.— 2. Burned campin t rned flowering plant.— 4. Contrast between a severely burned, killed plant and a moderately burned regenerating plant.—5. Detail of the base of a burned, regenerating plant.—6. Detail of a section of the com inflorescence showing the small, long-peduncled male inflorescences below and above and the uel laa seine short-peduncled female inflorescences in the center. Photographs by Dr. Cleofé E. Calderó IE Ii Mi ; Ill TI | WU R FiGuRE7. Morphology of Arundoclaytonia dissimilis.—A. Basal portion of culm with the sheath bases worn off and showing the approximately identic m Line of the buds every sixth node.—B. Portion of a leaf n ligule. — C. Sq inflorescence. — D. Male spikelet. E-I. Components of ipn spikelet.— —E. Lower glume Upper glume.—G. Lemma.—H. Inner P i palea with the two stamens composing the male flower. E Stamen. — J. Female inflorescence. — K. Female spikelets. L-R. Components of female Pudet —L. Lower iin me.— . Upper glume.—N. Lower lemma.— O. Middle lemma, ventral view.— P. Middle lemma, dorsa Middle palea and rachilla extension with the rudimentary upper floret. — R. T proba bly pollinated and slightly expanded. S-U. Caryopsis in three different views. — —S. Lateral view.— T. Hilum view. es mbryo view. Scales: A, B = 1 cm; C, D, I-K, U = 1 mm; magnification for E-H, K-R, and S-U the sa 48 484 key: v = 7 ANNALS OF THE MISSOURI BOTANICAL GARDEN ff [Vor. 74 foliar bract female inflorescence male inflorescence prophyllum leaf B = meristematic tissue, bud ------- = continuation d illustration of a portion of the inflorescence of Arundoclaytonia dissimilis showing FIGURE 8. the Ein of the internodes, been altered for clarity nre are shown i in two dimensio At any node along the flowering portion of a main culm, male inflorescences are produced be- fore the female inflorescences, and the male in- florescences are borne on longer branches than those of the female inflorescences. Each branch complement along the main culm ultimately ter- minates in a male inflorescence. The proportion of male to female inflorescences varies from 2 male: 1 female in the lowest portion, gradually changing to 1 male:3 female in the uppermost portion. wever, since the male spikelets are functionally 2-8-flowered compared with the functionally one-flowered female spikelets, the total number of male flowers is greater than fe- male flowers. Although the plant is fundamen- tally protandrous, the large number of inflores- leaves, prophylla, nie EEA: The proportions of the structures have cences produced by any mature plant ensures a significant overlap between the flowering of male and female spikelets. No information is available about self-incompatibility or frequency of flow- ering. The male and female spikelets are strongly di- morphic and, besides the difference in flowers and number of anthoecia, differ significantly in the anthoecial morphology. Both the lemma and especially the palea of the functional floret of the female spikelets are convolute and thicker in tex- ture, and the palea is greatly elongated and 9- 13-nerved (Fig. 7K, O-Q). In contrast, the lem- mas and paleas of the male spikelets are mem- branous, the lemmas are 3-9-nerved and round- ed on the back, and the paleas are 2-keeled and 1987] 2-nerved (Fig. 7D, G, H). The relative lengths of lemmas and paleas in the two kinds of spike- lets are similar, the paleas being longer in both kinds, although those of the female spikelets tend to be somewhat longer than those of the male spikelets. In both kinds of inflorescences the outer whorl of spikelets is surrounded by a ring of small ster- ile bracts that we interpret to represent rudi- mentary spikelets (Fig. 7C, J). In some cases these rudimentary spikelets may reach 4 mm in length in the female inflorescence and consist of four or five bracts in the same positions as the normal female spikelet parts. From such rudimentary spikelets there is a gradual diminution and sim- plification to small solitary bracts. Occasionally one of the normally sized female spikelets on the outside of the inflorescence has an extra bract. However, those on the inside of the inflorescence uniformly have the two glumes and three florets. In the several cases where a small extra bract was observed in the inner part of the inflorescence, it clearly originated below the very short pedicel and presumably also represented a rudimentary spikelet. LEAF BLADE ANATOMY ANATOMICAL TECHNIQUES Anatomical studies were carried out on leaves from herbarium specimens and those fixed in the field in FAA. Preparation of the sections fol- lowed the methods outlined by Ellis (1984). The very fibrous nature of the leaf blades frequently caused the sections to tear as they were cut, mak- ing it difficult to obtain completely undamaged sections. LEAF IN TRANSVERSE SECTION Outline: open, expanded with the margins slightly recurved (Fig. 9). Leaf thickness 30 um laterally to 40 um centrally. Ribs and furrows: prominent, flat-topped adaxial ribs with straight, vertical sides present over all the vascular bun- dles (Fig. 10); ribs associated with first-order and third-order vascular bundles of identical size and shape; furrows narrow, cleftlike, penetrating at least half the leaf thickness. Abaxial ribs or fur- rows absent. Median vascular bundle: no midrib or keel developed; median vascular bundle struc- turally indistinguishable from lateral first-order bundles. Vascular 25 first-order bundles with metaxylem vessels 1O1€ than DAVIDSE & ELLIS—ARUNDOCLAYTONIA 485 per leaf section; one third-order bundle without metaxylem vessels between consecutive first-or- der bundles, this alternating pattern occurring across the full width of the blade (Figs. 9, 10). All vascular bundles located slightly closer to the abaxial surface. Vascular bundle structure: first- order bundles round to elliptical in outline (Figs. 9-12); phloem tissue adjoining the inner bundle sheath; protoxylem lacunae present; metaxylem vessel elements wide (+5 um) with a diameter double that of the parenchyma sheath cells, thin- walled and slightly angular (Fig. 12). Third-order bundles elliptical with xylem and phloem tissue distinguishable. Vascular bundle sheaths: first- and third-order bundles completely surrounded by an inner bundle sheath (Fig. 11); mestome sheath cells relatively large, of the same diameter as the parenchyma sheath cells; secondary walls heavily but uniformly thickened, almost exclud- ing the lumen (Figs. 11, 12). Outer bundle sheath round, adaxially and abaxially interrupted by sclerenchyma girders (Figs. 11, 12); bundle sheath extensions absent; cells elliptic, variable in size, thin-walled and lacking chloroplasts. Scleren- chyma: adaxial girders inversely anchor-shaped, following the shape of the adaxial ribs (Figs. 11, 12); abaxial sclerenchyma forming a continuous hypodermal band with projections toward the vascular bundles as well as the bulliform cell groups (Fig. 11). Fibers very thick-walled with lumens almost completely filled; lignified, except those projecting toward the bulliform cells which may have cellulose secondary walls (Fig. 10). Me- sophyll: chlorenchyma not radiately arranged; cells small, isodiametric and tightly packed with- out visible intercellular air spaces (Fig. 11); oc- cupying the sides of the ribs but divided abaxially by the bulliform cells, colorless cells, and abaxial hypodermal sclerenchyma; arm or fusoid cells absent. Colorless, inflated, thin-walled paren- chyma cells linking the bulliform cells to the hypodermal sclerenchyma. Adaxial epidermal cells: bulliform cells at the base of all furrows and occurring in restricted, fan-shaped groups with an inflated central cell. Epidermal cells with a very thick cuticle, even on the sides of the furrows; papillae or macrohairs absent; inter- locking prickles common on the sides of the fur- rows (Figs. 11, 12). Abaxial epidermal cells: bul- liform cells absent; epidermal cells small, with an extremely thick, continuous cuticle equal in thickness to the diameter of the epidermal cells; hairs, papillae, and stomata absent; costal and intercostal zones not differentiated. 486 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURES 9-12. Leaf blade anatomy of Arundoclaytonia dissimilis in transverse section. —9. Outline showing the absence of a keel.— 10. Alternating first- and third-order vascular bundles and buen nent adaxial ribs and cleftlike furrows.—11. Anatomical detail showing double bundle sheath, compact mesophyll, Dance anchor- shaped adaxial sclerenchyma girder, abaxial hypodermal band and reel celis E associated colorless cells. — 12. Scanning electron e adaxial furrows and structure of the metaxylem vessel elements. Scales: 9 — 50 u ; 10 = 20 um; 11, 12 = iD um. " Based on Anderson et al. 10950 (Figs. 9, 10) and Calderón et al. 2706 (Figs. 1 1, 12). ABAXIAL EPIDERMIS rophytic adaptations. This is demonstrated most clearly by the well-developed abaxial hypoder- mal sclerenchyma, extremely thickened abaxial epidermis, and strongly ribbed and furrowed adaxial surface. These xerophytic modifications appear to have led to the consequent loss or re- duction of many epidermal features commonly well developed in most other grasses. The most important reductions are the lack of distinction between the abaxial costal and intercostal zones, the absence of abaxial stomata and microhairs, and the reduction of silica bodies. These xero- phytic features undoubtedly allow a rapid in- rolling of the leaves and may be responsible for the involute margins or completely involute leaves observed on the herbarium specimens. Zonation: costal and intercostal zones indis- tinguishable; entire epidermis composed of uni- form long and short cells (Figs. 13-15). Long cells: elongate rectangular, length 3 x the width, anticlinal ben parallel, end walls vertical (Fig. orizontal and vertical anticlinal walls kaviy Ghee. pitted and deeply sinuous. Long cells usually adjoining one anther but in- frequently separated by cork-silica cell pairs. Sto- mata: lacking on the abaxial surface (Fig. 13). Short cells: tall, with irregular ao associated with silica cell of similar shape (Fig. 14); occur- rence irregular. Papillae: absent. MS ab- sent. Silica bodies: tall and narrow, irregular in outline; scattered throughout the epidermis. + WT CLASSIFICATION AND DISCUSSION ADAXIAL EPIDERMIS As mentioned in the introduction, we consider Arundoclaytonia to belong in the now bigeneric Steyermarkochloeae despite the fact that Arun- doclaytonia differs markedly from Steyermarko- chloa in growth habit, leaf morphology, and in- Sides and tops of the ribs covered with prickle rickles obscuring all other epidermal details of this surface (Fig. 16). ANATOMICAL OBSERVATIONS markable rese bibis ance between the genera be- The anatomy of the leaf blade of Arundoclay- comes evident. Ignoring for the moment a low tonia is highly modified and exhibits many xe- percentage of bisexual spikelets in Steyermarko- heavily thickened, sinuous- walled lon photomicrograph of the adaxial epider DAVIDSE & ELLIS— ARUNDOCLAYTONIA BRRRG G i Aru ann, COUT oye P NMU Y V VENT ^ LUMA CL i TRAIL DOR i m w r^ p A VU" ev t ert try xr a ` um 5 = 10 um; magnification for 14-16 = - 10 um. Based on Anderson et al. 10950 (Figs. 13. 14) and Calderón et al. 2706 (Figs. 15, 16). chloa, the fundamental structure of the unisexual spikelets is identical in the two genera. The male spikelets have two stamens, are cioe and lack lodicules in both genera. The majo difference in the male spikelets is that those of Arundoclaytonia have more florets. This kind of variation is analogous to that between species of Eragrostis, Bromus, and Bambusa, to name just three of the many genera in which this kind of ariation is well known. The female spikelets are in the size and eau aes is SEES These striking and fi et almost certainly not due to: Ameena evolution but indicate a fundamental phylogenetic rela- tionship which is reflected in our classification of Arundoclaytonia in the Steyermarkochloeae The generic status of Arundoclaytonia is jus- tified by the following major differences from Steyermarkochloa: monomorphic vs. dimorphic culms; typical vegetative leaves with open sheath, many-ribbed blade, and ligule vs. highly modi- fied vegetative leaves with stemlike, solid sheath, two-ribbed blade, and ligule absent; many small, hemispherical male and female inflorescences aggregated into a false panicle vs. single, large, terminal, spicate inflorescences bearing male, fe- male, and bisexual spikelets; and in leaf anato- my—vascular bundles at one level vs. different levels; absence vs. presence of lacunae; absence vs. presence of abaxial stomata; and adaxial fur- rows and ribs associated with all vascular bun- dles vs. associated only with the median vascular bundle. Our decision to place Arundoclaytonia in the Steyermarkochloeae necessitates a modification of the description of the tribe (Davidse & Ellis, 1984). Because the leaves of Steyermarkochloa are unique in the family, leaf characters were believed very important i and in differentiating it from others. This i is now gthe tribe 488 shown to be true only when Steyermarkochloa was known. In fact, at the macroscopic level, the tribe now encompasses both “normal” and high- ly modified leaves. In this light the unusual fea- tures of the leaves of Steyermarkochloa, both at the macro- and microscopic levels, must be seen as adaptations to its seasonally inundated habi- tat, just as the strongly xerophytic features of Arundoclaytonia are presumably adaptations that allow it to cope with the nutrient deficiencies, frequent moisture stress, and intense solar ra- diation of the white-sand soils of its campina habitat (Ab'Sáber, 1982; Anderson, 1981). Such white-sand soils are considered to be the most nutrient-deficient soils in South America (Eiten, 1978). Although the campinas are located in high rainfall regions, they dry out rapidly near the surface during periods of low rainfall and never experience the long-sustained inundation of the sabaneta or morichal habitats of Steyermarko- e (Ab'Sáber, 1982; Eiten, 1978; Anderson, ier formal, emended tribal description is the following: Steyermarkochloeae Davidse & Ellis, Ann. Mis- souri Bot. Gard. 71: 994. 1985. Perennial grasses with mono- or dimorphic culms and leaves; leaves solitary or numerous per culm, consisting of a flattened sheath, blade, and ligule, or a solid, cylindrical sheath and flat- tened blade without a ligule, or reduced to blade- less flattened sheaths. Inflorescence spicate, elon- gate and cylindrical or a hemispherical cluster of spikelets, bearing male or female spikelets only, or bearing female spikelets above male and bi- sexual spikelets. Spikelets solitary, usually uni- sexual, dorsally compressed, disarticulation be- m style 1; caryopsis fusiform; male spikelets 2-9-flowered, the paleas 2-keeled; fe- male spikelets 3-flowered, the lowest floret ster- ile, the middle floret fertile, the upper floret ru- dimentary and borne on a prominent rachilla segment; palea of the functional female floret spongy, curved, (5-)7-13-nerved, longer than the lemma When describing the Steyermarkochloeae and including it in primarily on anatomical characters. At the same ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 time, they recognized its uniqueness in the subfamily on the basis of the gross morphology of the leaves, inflorescence, spikelets, and flow- ers. Much of that detailed discussion is appli- cable to Arundoclaytonia as we The discovery of “normal” leaves in the Stey- ermarkochloeae lessens the importance of the unique leaves of Steyermarkochloa vis-a-vis the other genera of the Arundinoideae. In leaf mor- phology we now consider such leaves to be basic in the Steyermarkochloeae and thus well within the norm of the Arundinoideae, and we consider the leaves of Steyermarkochloa to be a later spe- cialization that evolved in its own lineage. n contrast, the 1 flowers of Arundoclaytonia are deri ore special- ized than those of mU Bisexual flowers are universally considered more primi- tive than unisexual flowers among grasses. Their Occurrence, along with the more specialized uni- sexual flowers, in Steyermarkochloa and their absence in A loclaytonia indicates that the lat- ter is more specialized in this respect. Similarly, we consider the occurrence of bisexual and uni- sexual spikelets in one inflorescence in Steyer- markochloa to be less specialized than the seg- regation of the spikelets into strictly male and female inflorescences in Arundoclaytonia. This strict separation of male and female flowers may be considered the ultimate step in the Charles- orth & Charlesworth (1978) model of the evo- lution of monoecism through a gynomonoecious pathway discussed by Davidse & Ellis (1984) for Steyermarkochloa. Only in one respect of spikelet morphology might Arundoclaytonia be considered less spe- cialized than Steyermarkochloa, and this is in the number of florets of the male spikelets. The occurrence of two sterile florets in the female spikelets of both genera suggests a reduction in number of fertile florets. This interpretation is supported by the occurrence of bisexual spikelets in Steyermarkochloa with two bisexual florets and others with a male floret below the bisexual floret. Against this background, the larger num- ber of florets in the male spikelets of Arundo- claytonia might be less specialized. This inter- pretation must be tempered with the fact that floret number may easily change up or down, as is evident in the Arundinoideae, Pooideae, Chloridoideae, and Bambusoideae in general, al- though reduction seems to be prevalent in the — S ily. The number of florets in the Arundinoideae 1987] varies from one to many, but the predominant trend and probably the primitive condition in this trend well in the male spikelets. In the female spikelets both apical and basal reduction are ev- ident. In reduction of the lowest floret to a sterile lemma, Steyermarkochloa and Arundoclaytonia resemble the Panicoideae. However, the termi- nal rudimentary floret plus the numerous florets of the male spikelets of Arundoclaytonia suggest that this similarity is convergent. The rather rounded female spikelets of Arundoclaytonia and Steyermarkochloa are presumably due to reduc- morphology of these two genera is the reduction of the number of spikelets per inflorescence in Arundoclaytonia, but, at the same time, the ag- gregation of the many small unisexual inflores- cences into a large false panicle. This exactly par- allels a trend in other tribes in the family, for example, Saccharum vs. Hyparrhenia in the An- dropogoneae (Clayton, 1969) and Panicum lig- ulare Nees vs. P. rudgei Roem. & Schult. in the Paniceae Anatomically, the absence of abaxial stomata and microhairs and the reduction of silica bodies in Arundoclaytonia complicates the phylogenetic interpretation of the anatomical structure of the leaf blade as many of these features are generally claytonia, therefore, does not exhibit the com plete set to assign grasses to a given subfamily, thus lim- iting our ability to use these characters for de- termining the affinities of this unusual grass. However, by a process of elimination certain possibilities can be discarded. Arundoclaytonia does not possess arm or fu- soid cells and, therefore, cannot be accommo- T in the Bambusoideae. The Chloridoideae is entirely Kranz with only one possible excep- tion ee 1984), and the Panicoideae is pre- dominantly C,. Non-Kranz members of the pan- icoid group all have a semiradiate type of mesophyll and do not have the compact, isodi- ametric emloreschyma a Irunqoerd tona hows no anatomical resemblance with ps panicoid grasses, and phylogenetic relationships with this subfamily appear most unlikely. The type of chlorenchyma found in Arundoclaytonia does, DAVIDSE & ELLIS— ARUNDOCLAYTONIA 489 however, occur in many arundinoid grasses. Some members of the Pooideae also have this type of chlorenchyma, but the pooid grasses typically do not have sinuous long cells or tall, vertical silica bodies as does Arundoclaytonia. The leaf ana- tomical evidence, although somewhat limited, does suggest arundinoid affinities for Arundo- claytonia, and its systematic position does ap- pear to lie with the Arundinoideae. The tribes ofthe Arundinoideae cannot be sep- arated on anatomical criteria and the decision to classify Arundoclaytonia in the Steyermarko- chloeae is based on morphological s. 1. mainly that of the nius This decision is nei- ther confirmed nor refuted by the anatomical evidence. The leaf anatomy of Steyermarkochloa and Arundoclaytonia differs substantially and both appear to have highly advanced and derived leaf anatomy. Because of these great anatomical differences, they undoubtedly cannot be accom- modated in the same genus Clayton & Renvoize (1986) considered the Steyermarkochloeae to be a tribe in the Pani- coideae, noting “an obvious resemblance to Hy- menachne" in features that are not unique. They furthermore believe Steyermarkochloa to fun- damentally differ from C, panicoids only in the lack of microhairs (Renvoize, in litt.). We believe that the additional evidence presented by us for Arundoclaytonia gives further support for our classification of the tribe, although we certainl recognize the isolated position of the Steyer- markochloeae in the Arundinoideae, and rec- ognize that we are adding one more relatively discordant element to the traditional “dumping ground" of the family. Except for the lacunae and stellate cells in the leaves and the gross form of the inflorescence, but not its branching pat- tern, we find it difficult to observe any obvious resemblances between Steyermarkochloa and Hymenachne. We do agree with Clayton & Renvoize (1986) that embryoc would new information for clarifying the duxi cul sition of the tribe. Chromosome information could also be potentially useful. Unfortunately we were unable t obtained from an herbarium specimen af Arun- doclaytonia. In the case of Steyermarkochloa all our meiotic cytological samples were too young. Unlike the typical situation in grasses, inflores- cences of Steyermarkochloa must apparently be well exserted from the sheath before meiosis takes place. crarvnncec 490 The distribution of Steyermarkochloa in the northern Amazon basin and Arundoclaytonia in southcentral Amazonia suggests that other taxa d, since large areas of XEM Bir ie em botanicallv very poorly known. Obviously, the area between the ranges of these two genera would seem to be the most promising in this respect. Any relatively open vegetation on white-sand soils (campinas or Amazonian caatingas) might harbor further taxa ofthis tribe. These vegetation types are found in greatest abundance in the drainages of the Rio Negro and the Rio Branco (Eiten, 1978; Ander- son, 1981). LITERATURE CITED AB'SÁBER, A. N. 1982. oo and paleo- eco logy of Brazilian Amaro a. Pp. Al -39 i n G. cal D Tropics. Columbia 1. pde New York. ANDERSON, A. 81. White-sand — of Brazilian een Biotropica 13: 199- ARBER, A. 1934. The Gramineae. The Du Press, Cambridge ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 BARNARD, C. 1964. Form and structure. Pp. 47-72 in C. Barnard (aeon, Grasses and Grasslands. Macmillan & Co., Ltd., New York. CHARLESWORTH, B. & D. CHARLESWORTH. 1978. ulation genetics of partial male sterility and ike evolution of monoecy and dioecy. Heredity 41: 37- CLAYTON, w. 969. A revision of the genus Hy- parrhenia. yo Bull. Add. Ser. 2: 1-1 & S. A. RENVOIZE. 1986. Genera Graminum. DAVIDSE, G. R. LLIS. 1984 [1985]. Steyer- markochloa unifolia, a a new genus from Venezuela and Colombia (Poaceae: Arundinoideae: Steyer- markochloeae). Ann. Miescün Bot. Gard. 71: 994— 1012. EITEN, G. 1978. Delimitation of the cerrado concept. Vegetatio 36: 169-1 ErLis, R. P. 1984. Eragrostis walteri—a first record of non-Kranz leaf anatomy in the sub-family an : cation of the Gra Watson, L. & M. J. ee Grass em The renee National Univ., Canberr: SIPHOCAMPYLUS OSCITANS (CAMPANULACEAE: LOBELIOIDEAE), A NEW NAME FOR BURMEISTERA WEBERBAUERI FROM PERU! BRUCE A. STEIN? ABSTRACT Burmeistera weberbaueri is transferred to the genus Siphocampylus under the new name Siphocam- pylus oscitans. The species is noteworthy in being one of only three species of Siphocampylus known to have a dilated anther-tube orifice. As part of a reassessment of the Peruvian members of Burmeistera (Stein, 1987), it has be- come apparent that B. weberbaueri Zahlbr. ac- tually belongs to the large Andean genus Sipho- campylus. This paper makes the appropriate generic transfer and proposes a new name to ac- commodate this unusual species. Generic delimitations in Campanulaceae subfamily Lobelioideae have long been recog- nized as problematic (Gleason, 1925; McVaugh, 1940). Traditional systems, such as those of Presl (1836) and Wimmer (1943), rely heavily on fruit morphology for classification at the tribal level. In particular, capsular versus baccate fruit is a fundamental character used to define and align genera. Strict reliance on this dichotomy in the classification of the subfamily has separated close relatives, for example, capsular-fruited Sipho- campylus and baccate-fruited Centropogon. Al- though fruit type in conjunction with other fea- tures can be reliable for clustering related ids of species, probable convergence in fruit c acters suggests caution in its application. Among neotropical Lobelioideae the emphasis on fruit type, and to a lesser degree on the pres- ence or extent of a dorsal slit in the corolla (another seemingly labile character), has yielded genera of convenience. One of the most natural groupings, however, appears to be the genus Bur- meistera, which is characterized by baccate fruits, oblong or linear seeds, mostly nonbracteolate pedicels, entire corolla tubes, and distally open and oblique anther tubes often with little or no apical pubescence. The distally open anther tube of Burmeistera is one of its most distinctive features. In most genera of Lobelioideae the three dorsal anthers are longer than the two ventral ones and curve downward at the apex, effectively closing the mouth of the anther tube. This allows the inter- nally released pollen to build up pressure as the style and stigma elongate, pushing pistonlike through the anther tube. The characteristic tuft of stiff hairs at the tip of the ventral anthers func- tions as a lever that opens the orifice slightly and allows the pressurized pollen. to discharge. Pre- traea by Brantjes (1983) and has been observed in the neotropical genera Centropogon and Sipho- campylus (Stein, in prep.). Since the dorsal an- thers in Burmeistera do not curve downward closing the anther tube, this type of regulated pollen discharge does not occur. This difference in pollen presentation probably explains the cor- related feature of glabrous or only sparsely pilose anther apices in Burmeistera sect. Imberbes F Wimmer: apical hairs there have no function as trip mechanisms. Whether the densely villous tuft at the tip of the ventral anthers in Burmeis- tera sect. Barbatae F. Wimmer has a functional role in pollen discharge is not clear Burmeistera weberbaueri was indludeti in this genus by Zahlbruckner (1906) on the basis of floral features alone, as he had no mature fruit. This decision was probably based upon his ob- servation of a naked and dilated anther-tube ori- fice, the absence of bracteoles, and the somewhat orter P. Low ! I am grateful to Carlos Reynal for locating the type specimen at MOL and to B. E. — for searching B. Dan for type material at B H. Nicolson kindly pro II and Peter Goldblatt made useful suggestions on an earlier draft. Fieldwork i in Peru was densi by National Science Foundation "in Dissertation Improvement Grant BSR84-13912 and by a Garden Club of America Award in Tropical Bot ? Missouri Botanical peA and Washington University. P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A ANN. Missouri Bor. GARD. 74: 491—493. 1987. 492 *burmeisteroid" corolla morphology (a short, straight tube with an abruptly ampliate throat and falcate lobes). The strongly turbinate hypan- thium visible in the Field Museum type photo- graph suggests the mature fruit of this species to be capsular rather than baccate. Furthermore, the coriaceous and rugose texture of the leaves characterizes many members of Siphocampylus but is unknown in Burmeistera. An open anther- tube orifice of the kind described by Zahlbruck- ner is, however, extremely rare in Siphocampy- lus. OBSERVATIONS In order to investigate further the generic placement of this species, I visited the type lo- cality in January 1987 to collect fresh material and to ascertain the fruit type and anther-tube morphology. This represents the first collection since the type was collected by August Weber- bauer 85 years before. The fruits turned out to be capsular with half-superior ovaries. Careful examination of fresh pedicels occasionally re- vealed bracteoles, a feature often difficult to ob- serve in dried material. The presence or absence of these bracteoles was consistent within indi- vidual inflorescences. The most interesting con- firmation is of the dilated anther-tube orifice. Zahlbruckner's description of this feature is true for the species and not based on an artifact of preservation as I had previously assumed. TAXONOMIC TREATMENT Siphocampylus oscitans B. A. Stein, nom. nov. Burmeistera iier A. Zahlbr., Bot. y 451. 1906. Non Siphocam- pylus ee A. Zahlbr., Bot. Jahrb. Syst. 37: 456. 1906. TYPE: Peru. Junin: Prov. Tarma, mountains east of Huacapistana, 3,200 m, Jan. 1902, Weberbauer 2203 (lec- totype here designated, W; isolectotype, G). SYNTYPE: Peru. Junin: Prov. Tarma, moun- tains east of Palca, 3,200-3,600 m, Feb. 1902, Weberbauer 2473 (MOL, photograph of lost B syntype, F neg. 9074). Erect many-branched shrubs to 1.5 m tall, gla- clustered toward branch tips, alternate and spi- rally arranged, rarely appearing subopposite or pseudoverticillate, sessile; blades narrowly ovate ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 to lanceolate, (3—)5—9 cm long, (0.7-)1.5-4 cm wide, apex attenuate, base rounded to obtuse; margins densely and sharply callose-serrulate, 9— 13 teeth per cm; when fresh the lamina fleshy and coriaceous with the tips recurving, the adax- ial surface smooth and dark green, the abaxial surface pale or occasionally tinged purple, drying coriaceous with the adaxial surface nitid and ru- gose, prominently impressed by the 6-8 acutely ascending, almost straight secondary veins. In- florescence subcorymbose and auxotelic, the flowers solitary in congested upper leaf axils, the rachis later elongating and producing normal fo- liage leaves beyond the maturing flowers and fruits; pedicels ascending to spreading, 4-8 cm long, generally exceeding length of subtending leaves, occasionally with sub-basal, subulate bracteoles to 2 mm long. Flowers 35-40 mm panthium widely obconic, the sides erect or slightly spreading, triangular, 8-11 mm long, 3—4 mm wide at base, about twice the length of hypanthium, the margins entire or minutely denticulate; corolla green and maroon, yellowish within, corolla tube straight, 15-18 mm long dor- sally and 10-15 mm long ventrally, ca. 6-8 mm wide at base, narrowing and re-expanding slight- ly above, the lobes lance-oblong with acuminate slightly revolute tips, mostly abruptly decurved, the dorsal lobes falcate, 14-17 mm long, ca. 4 mm wide at base, the lateral and ventral lobes 11-14 mm long; filament tube 27-30 mm long, adnate to corolla ca. 4 mm above base, glabrous and greenish, well exserted from corolla; anthers connate, the upper 7-8.5 mm long, the lower 6.5- 7 mm long, glabrous except for sparse, wispy g capsule dehiscing by two terminal valves, 15 mm long (including valves), 10 mm wide; seeds el- lipsoid, minutely foveate, ca. 0.6 mm long. Distribution. Known only from shrubland at ca. 3,200 m on the eastern slope of the Andes near Palca and Huacapistana in Junín Depart- ment of central Peru. Additional specimens examined. PERU. JUNIN: Vi- toc, 3,600 m, Feb. 1984, Pearce s.n. (BM); Prov. Tar- ma, mountains E of Palca, rd. to Illic, 10-12 km above Culebreo, 11?19'S, 75?33'W, 3,250 m, 13 Jan. 1987, Stein, Kallunki & Diaz 3831 (AAU, B, CAS, F, G, K, MO, NY, P, US, USM 1987] DISCUSSION The presence of capsular fruit in this species excludes it from Burmeistera as currently cir- cumscribed. Among the capsular-fruited genera it is best placed in Siphocampylus based on the entire corolla tube. Since the name Siphocam- pylus weberbaueri has already been used by Zahl- bruckner (1906), I propose the new name Sipho- campylus oscitans for this species. The epithet oscitans derives from the Latin for yawning, in reference to the dilated anther-tube orifice, an unusual feature in Siphocampylus. Siphocam- pylus oscitans is easily recognized among Peru- vian members of this genus by its completely glabrous, stiff, coriaceous, lanceolate leaves, rel- atively short and stout corolla with strongly de- flexed lobes, and erect to spreading sepals that are about twice as long as the strongly obconic hypanthium. open anther tube of the kind found in Sipho- campylus oscitans is known in only two other members of the genus: S. sceptrum Decne. of Venezuela and S. rusbyanus Britton of Bolivia and southern Peru. While S. oscitans occurs nearer geographically to S. Past partas, erana logically it more phases of the highly variable S. eena. In addition to the open and thinly pubescent anther-tube ori- fice, certain collections of S. sceptrum share the following features with S. oscitans: a half-supe- rior ovary and shallow hypanthium; thick, ses- sile, and often lanceolate or narrowly ovate leaves VIUSVI Y pedicels; adnation of the filaments to the corolla tube near its base; and a stout, straight corolla tube. The two species differ most obviously in the length of the corolla and in the extent of fusion of the four upper lobes (the feature that best characterizes S. sceptrum). On the basis of these similarities, S. oscitans seems best placed near S. sceptrum in sect. Brachysiphon F. Wim- er. Zahlbruckner (1906) cited two Weberbauer collections (2203 and 2473) in the protologue of Burmeistera weberbaueri. Gleason (1925) des- ignated Weberbauer 2203 as the “type,” but it is unclear whether this constitutes valid lectotypi- fication. He gave no indication that the type spec- imen was actually examined and may well have STEIN — SIPHOCAMPYLUS OSCITANS 493 been following the then common practice of des- ignating the first collection listed in the original description. The type specimens studied by Zahl- bruckner at Berlin were both apparently de- stroyed during World War II (B. E. Leuenberger, pers. comm.), and my search of that herbarium confirms their absence. Because Zahlbruckner worked at Vienna and annotated the W sheet of Weberbauer 2203, I have designated that spec- imen as the lectotype. Note added in proof: Recent examination of the type of Burmeistera splendens at BM shows this to be a fourth species of Siphocampylus with a dilated anther tube. This Colombian species appears closely related to S. oscitans based on anther tube, hypanthium, sepal, and foliage fea- tures, and provides support for the above dis- cussion linking S. oscitans of Peru with S. scep- trum of Venezuela. I here make the transfer of B. splendens to Siphocampylus, as has already been suggested by Jeppesen's annotation on the type specimen. Siphocampylus splendens (F. Wimmer) J eppesen ex B. A Andes del Norte, July 1857, Tiana 3059/ 23 (holotype, BM). LITERATURE CITED BRANTJES, N. B. MD 83. Regulated VR issue in Isotoma, Campanulaceae, and evo n of sec- onary pollen uit meer Acta Bot. eee 32: 1925. d gem on the flora of northern s Burmeistera. Bull. GusAsoN lg A. h America IV: Torrey Bot. es 52: 93- 104. McVAuGH, R. 40. A revision of “Laurentia” and allied du fe in North America. Bull. Torrey Bot. Club 67: 778-798. . Studies in South American Lobelioi- deae (Campanulaceae) with special reference to Colombian species. Brittonia 6: 450-493. . Prodromus Monographiae Lobeli- . 1987. Synopsis of the genus Burmeistera in Peru. Ann. Missouri Bot. Gard. 74: 494—496. WIMMER, F. . Campanulaceae-Lobelioideae. Pflanzenreich IV. 276b: 1-260 (Heft 106) ZAHLBRUCKNER, A. 06. Campanulaceae andinae. Bot. Jahrb. Syst. 37: 451-463. SYNOPSIS OF THE GENUS BURMEISTERA (CAMPANULACEAE: LOBELIOIDEAE) IN PERU' BRUCE A. STEIN? ABSTRACT The southern distributional limit of peA has long been obscured by the inclusion in that genus of several disharmonious elements from cen southernmost stations known for these two species, and thus the genus as a whole, are at approximately 5°50'S in San Martín Department of northern Peru Burmeistera Triana is a well-delimited neo- tropical genus of Lobelioideae distinguished from other baccate-fruited genera by the combination of oblong or linear seeds, oblique and distally open anther tubes, and mostly nonbracteolate pedicels. The typically distorted, greenish to ma- roon corollas make the genus easily recognizable in the field. However, complex morphological patterns, in particular extreme local differentia- tion, render it taxonomically challenging at the species level. Burmeistera is found primarily in montane wet forests from Chiapas, Mexico south to Venezuela and Peru, with its center of diversity in the Andes of Colombia and Ecuador. The southern distri- butional limits of the genus have been problem- atic, owing to the questionable inclusion of sev- eral central and southern Peruvian taxa. This paper clarifies the status of Burmeistera in Peru and establishes its currently known southern lim- t Six Peruvian species of Burmeistera were in- cluded in the Flora of Peru (Wimmer, 1937) and in Wimmer’s (1943) revision of the genus, the most recent treatment available. These are: Bur- meistera asteriscus F. Wimmer, B. macrocarpa (A. Zahlbr.) F. Wimmer, B. peruviana F. Wim- mer, B. ramosa F. Wimmer, B. tricolorata F. Wimmer, and B. weberbaueri A. Zahlbr. Of these, only B. ramosa is retained in Burmeistera as the genus is presently circumscribed. McVaugh (1949) correctly noted the position of B. macro- carpa, B. peruviana, and B. asteriscus in Centro- pogon and made the necessary transfers for the last two. The two remaining species clearly be- long in Siphocampylus and are discussed below. An Pici species, Burmeistera micro- phylla J. D. Smith, was recently collected in northern ud bringing the number of Peruvian species to two. This collection at approximately 5*50'S in San Martín Department, along with a recent collection of B. ramosa from the same general vicinity, represents the southernmost sta- tions known for the genus The erroneous inclusion of several southern and central Peruvian taxa in Burmeistera has obscured an i The Huancabamba deflection of northern Peru marks the distributional limit for a number of plant and animal groups and appears to have been a significant barrier to north-south migra- tion (Simpson, 1975, 1979; Berry, 1982; Vuil- leumier, 1984). The geographical range of Bur- meistera as defined here provides another example of the Huancabamba deflection com- prising the boundary for a group. Although ap- proximately 30 species are present in Ecuador (Jeppesen, 1981), diversity in the genus falls abruptly in northern Peru, where only two species are known to occur. In Peru, these species have been found only in the area along or near the Río Maranon gap, the lowest elevation depression along the eastern slope of the Central Andes and a major component of the Huancabamba de- flection. Why Burmeistera is absent from apparently suitable habitats further south along the eastern slope of the Peruvian Andes remains a mystery. The relatively low elevation of the Rio Maranon gap (ca. 500 m) does not alone explain the pat- tern, since B. ramosa and other species occur at such elevations. Furthermore, members of the ! I thank James S. densa m Kallunki, and Porter P. Lowry II for helpful comments. Fieldwork in Peru e Foun Xuan n in Tropica dation p iin Dissertation Improvement Grant BSR84-13912 and ub any. ? Missouri Botanical Garden and Washington iM P.O. Box 299, St. Louis, Missouri 63166-0299, A. ANN. Missounmi Bor. GARD. 74: 494—496. 1987. 1987] genus are capable of dispersal past such barriers, as evidenced by their successful establishment in areas across far more formidable low-elevation gaps, in one instance reaching the isolated Cerro de la Neblina massif of the Guayana Highland. The complex patterns of local differentiation and endemism in Burmeistera suggest that dispersal may often be quite limited, even though the var- iously spongy, juicy, or inflated fruits often ap- pear well suited for bird-dispersal. In this regard it may be significant that B. microphylla is one of the widest-ranging members of the genus, ex- tending from Costa Rica to northern Peru. An additional factor may be our insufficient knowl- edge of the middle and upper-elevation forests along the eastern Andean slopes of northern Peru. These inaccessible areas are among the most poorly collected in the Andes. KEY TO THE SPECIES OF BURMEISTERA IN PERU la. Leaves thick and coriaceous when dry; sepals 5-8 mm long, ipd fruit we less thon 10 mm in diamet icrophylla i sepals 2-5 mm long, erect or spreading; fruit inflated, to 25 mm in diameter .. B. ramosa 1. Burmeistera microphylla J. D. Smith, Bot. Gaz. (London) 25: 146. 1898. The single Peruvian collection, as well as col- lections from southern Ecuador treated by Jep- pesen (1981) under the synonym B. aurobarbata F. Wimmer, differ from typical Central Ameri- can B. microphylla in their more uniformly lan- ceolate to narrowly lanceolate leaves and in lack- ing golden external anther trichomes. Further investigation of geographically intermediate populations of this wide-ranging and variable di complex (including B. crassifolia F. Wim- and B. maculata F. Wimmer) may indicate E these southernmost populations warrant treatment as a distinct taxon. Specimen lacius PERU. SAN MARTIN: Prov. W of Rioja on road to Pedro Ruíz, 2 with Amazonas Dept.), 2, na m, 16 Feb. 1985, Stein y Todzia 2198 (MO). 2. Burmeistera ramosa F. Wimmer, Repert. Spec. Nov. Regni Veg. 30: 16. 1932 This species is now known from two collec- tions in Peru. The type, Tessmann 4725, was collected at a relatively low elevation at the mouth STEIN—PERUVIAN BURMEISTERA 495 of the Rio Santiago along the Rio Marañon, not “near Iquitos” as indicated by Wimmer (1937). The second, a fruiting collection, was made in 1983 in the Venceremos region from mid-ele- vation cloud forest. Burmeistera ramosa is ap- parently more common in Ecuador (Jeppesen, 1981) and shows a similar wide range in eleva- tional preference. Since the early botanical ex- plorers Ruiz and Pavón never ventured into northeastern Peru (Steele, 1964) where this species would be expected, one of their collec- tions, formerly thought to originate from Peru (Wimmer, 1943), was probably among the Ecua- dorean collections made by their apprentice J. Tafalla. Specimens examined. PERU. AMAZONAS: Rio Ma- rañon, Pongo de Manseriche, 160 m, Tessmann 4725 (NY; photographs, MO, NY). SAN MARTIN: Rioja Prov., Pedro Ruíz-Moyobamba rd., km 390, Venceremos, 1,770-2,150 m, 5-7 Aug. 1983, Smith & Vasquez 4608 (MO). EXCLUDED SPECIES 1. Burmeistera asteriscus F. Wimmer, Repert. Spec. Nov. Regni Veg. 38: 5. 1935. = Cen- tropogon peruvianus (F. Wimmer) Mc- Vaugh, Brittonia 6: 462. 1949. See discus- sion under Burmeistera peruviana. 2. Burmeistera macrocarpa (A. Zahlbr.) F. Wimmer, Repert. Spec. Nov. Regni Veg. 30: 41. 1935. = Centropogon macrocarpus A. Zahlbr., Bot. Jahrb. Syst. 37: 452. 1906. 3. Burmeistera peruviana F. Wimmer, Repert. Spec. Nov. Regni Veg. 38: 5. 1935. = Cen- tropogon peruvianus (F. Wimmer) Mc- Vaugh, Brittonia 6: 462. 1949 Field studies conducted at the type locality of Centropogon peruvianus and C. asteriscus aroun Pillahuata, Cuzco Department, show that the dif- ferences noted by Wimmer (1935) in describing these two species, primarily leaf width and shape and sepal length, are variable within populations. The type specimens of these two species were collected in the same general vicinity and merely represent upper and lower elevation collections (3,000-3,300 m and 2,200-2,400 m) of a single species. The names were published simulta- neously; however, since McVaugh (1949) select- ed C. peruvianus as the type species for his Cen- tropogon sect. Peruviani, C. peruvianus is the preferred name. 496 4. Burmeistera tricolorata F. Wimmer, Repert. Spec. Nov. Regni Veg. 30: 22. 1932. = Si- phocampylus rusbyanus Britton, Bull. Tor- rey Bot. Club 19: 372. 1892 A fruiting collection of this species (Stein 2505) made recently at the type locality of B. tricolorata The Peruvian collections closely match Siphocampylus rusby- anus, a species recognized previously only from northern Bolivia. 5. Burmeistera weberbaueri A. Zahlbr., Bot. Jahrb. Syst. 37: 451. 1906. c CS oscitans B. A. Stein. A new collection of this species (Stein 3831) has shown the fruit to be capsular, excluding it from Burmeistera. For a discussion of this species, its transfer to Siphocampylus, and the necessary proposal of a new name (the epithet weberbaueri has been used previously in Siphocampylus), see the accompanying paper ii. 1987 Bb weber- . trico mprise Wimmer's (1943) bsc aequilatae of Burmeistera sect. imberbes F. Wimmer. Since both of these species are here excluded from Burmeistera, that strictly Peruvian subsection is no longer recognized. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 LITERATURE CITED BERRY, P. 1982. The systematics and evolution of dern sect. eter iii Ann. Missouri t. Gard. 69: pee S. um 11-170. McVAUuGH, R. 1949. Studies in South American Lo- belioidae (Campanulaceae) with special reference 4 rei EM In Fl. Ecuador 14: s in the flora of the high tropical Andes. EOS 1: 273- 294. 9. Quaternary biogeography of the high s. regions of South America. Pp. 157-1 Siphocampylus oscitans (Cam- panulaceae: Lobelioideae), a new name for Bur- meistera prx rn from Peru. Ann. Missouri Bot. Gard. 7 93. VUILLEUMIER, E pn Zoogeography of Andean birds: two major barriers; and speciation and tax- onomy of the Diglossa caibonaria superspecies. Natl. Geogr. Soc. Res. Rep. 16: 713-731. WIMMER, F. Burmeistera: eine e Pflanzengattung und ihre Arten. Repert. Spe Regni Veg. 30: 1-52. . 1937. Campanulaceae. /n Flora of Peru. Field Mus. Nat. Hist., Bot. Ser. 13: 383-489. ; ; Campanu laceae-Lobelioideae. Pflan- zenreich IV. 276b: 1-260 (Heft 106). NEW SPECIES OF PASSIFLORA SUBGENUS PASSIFLORA ROM ECUADOR L. B. HoLM-NIELSEN? AND J. E. LAWESSON? ABSTRACT new species of Passiflora from Ecuador are described, viz. P. montana, P. palenquensis, P. . and P. pergrandis. A revision of Ecuadorean Passiflora subg. Pas- siflora (subg. Granadilla sensu Killip) for the Flora of Ecuador (Holm-Nielsen et al., in press) has revealed four new species: P. montana, P. palen- quensis, P. deltoifolia, and P. pergrandis, be- longing to “series” Lobatae, Tiliaefoliae, Men- ispermifoliae, and Laurifoliae, respectively. The total number of Ecuadorean subg. Passiflora species is 19. Killip (1938) divided the subgenus into 15 series; although these are not validly pub- lished, we are following Killip for convenience. The “series” occurring in Ecuador are Digitatae, Incarnatae, Kermesianae, a Qua- rangulares, and those mentioned above Passiflora palenquensis Holm-Nielsen & Lawes- son, sp. nov. TYPE: Ecuador. Los Rios: Quevedo—Sto. Domingo road, km 56, Rio Palenque Science Center, 150-200 m, 6 Oct. 1979, C. H. Dodson, A. Gentry & G. Shupp 8854 (holotype, MO). Figure 1. Liana ubique glabra. Caule striato et pins Stipulis oblongo-lanceolatis, acutis, 1 x 0.4 cm, interdum de- p Nem angulo- alatis, 6. 5-9 em ^ longis cum 3-4n ib r apice extremo, pi glandes infra 2- 3 cm; Pedum basi- fixis, late ovatis, ovatis, 14-19 x cordatis, manifeste pinnatinervibus aceis, margine integra. Inflorescentia saltem pedicellis teretis a ess 3-4 cm nfra flore o 3 cm, ad b xe ce lobi calycis oblongi, acuti, 2.5 x 1.3 cm, de- isi et in Vb n mucronem; petalis oblongis, ob- 3 0.6-1 cm. Corona simplice, filamentis dx NE in 2 seriebus ordinatis quae ad apicem ex- tremam tubi calycis locatae sunt; series externa minute liguliformis, apex filiformis, 4 mm longa; serie interna crassisima, 4-angulata, 2 cm longa, apex filiformis, fas- urea et alba; tubus calycis glaber infra co- rme, 0.5 cm lon Jil gynophori limini circumcinctae. Ovario gla Liana, glabrous throughout. Stem terete, striate. Stipules oblong-lanceolate, acute, 1 x Í sometimes deciduous. Petioles angular-winged, 6.5-9 cm long, with 3-4 black, sessile urceolate glands, 1 mm long, one pair at extreme apex, other glands 2-3 cm below; blades basifixed, broad-ovate to ovate, 14-19 x 11—20 cm, deeply cordate, prominently pinnate-veined, lustrous, coriaceous, the margin entire. Inflorescence ses- sile, with at least 2 flowers collateral with the tendril in the axil of the leaf, 5-6 cm diam.; ped- icels terete, striate, 3—4 cm long; bracts verticil- late, united halfway, inserted about 5 mm below ower, oblong-ovate, mucronate, 5 x 2 cm. Hy- panthium tube funnel-shaped, 3 cm long, 1 cm wide at base, 2 cm at apex, outside lavender; calyx lobes oblong, acute, 2.5 x 1.3 cm, ending in a minute mucro; petals oblong, obtuse, 3 x 0.6-1 cm. Corona composed of 2 series situated at the edge of the hypanthium; outer series of short ligulate filaments, 4 mm long, with filiform apices; inner series of very stout, 4-angled, 2 cm long filaments with filiform apices, the inner fil- aments with shifting purple and white, 4 mm broad stripes; hypanthium smooth below coro- na. Operculum situated 1 cm above hypanthium floor, membranaceous, 0.5 cm long, split into filiform segments. Trochlea present at the an- rogynophore opposite the apex of the opercu- lum. Limen surrounding lower 5 mm of andro- ynophore. Gynophore 1 cm longer than the androgynophore present. Ovary glabrous. Fruit ! We thank R. R. Haynes and H. Balslev for iwi the manuscript, K. Tind for making the drawings, and N.-H. Andreasen, who supplied the Latin diagn Fieldwork was su the Danish Natural Science pporte ed by Research kao grant numbers 11-3038, 11- 3689. and 81-4024. This is deris ore No. 53 from the AAU Ecuador Pro ? Botanical PM University of Aarhus, 68 Nordlandsvej, DK-8240 Risskov, Denmark. ? Charles Darwin Rese ANN. Missouni Bor. GARD. 74: 497—504. 1987. arch Station Galapagos, Casilla 5839, Guayaquil, Ecuador 498 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 A \ (€ B 2cm FIGURE 1. Passiflora palenquensis Holm-Nielsen & Lawesson.— A. Flowering stem. — B. Longitudinal section of flower. 1987] ovoid, 7 x 4 cm, green. Seeds ovate, 5 x 3 mm, finely reticulate. Passiflora palenquensis is a member of *'se- ries" Tiliaefoliae, which is characterized by united bracts and entire leaves. Passiflora palenquensis has a distinct gynophore 1 cm longer than the androgynophore and differs from all other mem- bers of Tiliaefoliae by the corona having only two series of filaments. Other species of Tiliae- foliae have at least three series. It is closely re- lated to P. seemannii Griseb. which superficially appears to have two series. However, the two series of filaments are 1-1.2 cm and 2-2.5 cm long and have several rows of tubercles inside. ii iUi palenquensis, on the other hand, has nly two series, these being 4 mm and 2 cm, a Rebus The tubercles are absent. The new species is also related to P. tiliaefolia L., which has more numerous series of filaments and is confined to high altitudes (2,000-3,000 m). Pas- siflora palenquensis is restricted to low eleva- tions. Additional specimens examined. Coro NARINO: railroad Tumaco-Diviso km 86, 27 n: 1952, Castaneda 3326 (AAU). ECUADOR. LOS PI- CHINCHA: Sto. Domingo-Quevedo rd., km 45, Pan s Pilar-24 de Mayo rd., km 12, 600 m, Apr. 1980, Dod- son & Gentry 10337 (MO). Passiflora montana Holm-Nielsen & Lawesson, sp.nov. TYPE: Ecuador. Carchi: Tulcan-Mal- donado rd., km 67, 2,600 m, 78°04'W, pss. 21 May 1973, L. Holm-Nielsen, S. eppesen, B. Lejtnant & B. @llgaard 6200 ei ciuis AAU). Figure 2. Liana, ubique glabra. Caule bp atque tereto. Stip- ulis valde magnis, 4-4.5 x m, latioribus quam lon- gioribus, reniformibus mu incen ad basin obtusis, margi ne glandulare- “serrata, Petioli is 3. 5-4 cm longis, medium. peor ag basifixis, 3-lobatis, interdum re lobatis, 9-11 x ys 5-13 cm, dimidium oon , lobi 3 cm lati, lanceolati, acu Iti; pedato-venatis, integris cum glandibus in uus Bracteae verticillatae, ane portatae 5 cm infra florem, ovatae, acuminatae, ad basin cordatae, integrae, 2.5-3.0 x cm viridibus. Flores solitarii; pedicellis 4.5—6 cm longis, teretis. Flo- res 5-6 cm lati; tubo calycis breve campanulato, 5-6 x 7-8 mm; lobis calycis oblongo-lanceolatis, Meise 1.5 x 0.5 em, d manifeste carinatis, c arista 6 mm longa quae apicem excedit; dein oblon gis, subacutis, pallido- aaa 1-1.5 x 0.3-0.5 c eriebus; externae A se- versus operculum. ENDE uni epum recur- HOLM-NIELSEN & LAWESSON — NEW ECUADOREAN PASSIFLORA SUBG. PASSIFLORA 499 vatum, modice plicatum, 4 mm longum, dun d superioris filiformiter fissum. Limen is n base phori affixum, margine lobata. Ova pr bro. p pallido-virides cum maculis lilacinis. Pom non visi. Liana, glabrous throughout. Stem striate, te- rete. Stipules very large, 4—4.5 x 2 cm, reniform, mucronate, obtuse at base, margin glandular-ser- rate. Petioles 3.5—4 cm long, terete, striate, with two alternate, stipitate glands above middle, 4 mm long; blades basifixed, 3-lobed, occasionally 4-lobed, 9-11 x 11.5-13 cm, lobed halfway; lobes 3 cm wide, lanceolate, acute; pedate-veined, en- acuminate, cordate at base, entire, 2.5-3 x cm. Flowers 5-6 cm wide, solitary, lateral; ped- icels nins cm long, terete; hypanthium short campanulate, 5-6 x 7-8 mm; calyx lobes ob- c sq green, 1.5 x 0.5 cm, obtuse, prominently carinate, with a 6 mm long awn exceeding apex; petals oblong, cr light green, 1-1.5 x .5cm. Coronal filaments of several series; outer three series of filiform elements 12 mm long succeeded by about 5 irregular series, of slender filiform elements, 6 mm long, lilac, extending towards operculum. Operculum mem- branaceous, recurved, slightly plicate, 4 mm long, bu . Nectar ring present, conspicuous. deta absent. Ovary glabrous; styles light green with lilac spots. Fruits not seen. Passiflora montana as member of "series" Lobatae is most clearly related to P. sprucei Mast., from which it differs by not having the leaves divided below the middle, by having the stipules twice as large and having two (vs. three or four) petiolar glands 4 mm long and not sessile, and, especially, by the floral composition. In P. mon- ana the operculum is recurved and has a filiform upper half and the filaments are 3-4 mm long, whereas the operculum in P. sprucei is erect with the lower quarter membraneous and the upper two) outer series of long filiform filaments suc- ceeded by four or five inner series of 4-5 mm (vs. 2-3 mm) long filaments. Passiflora montana is the highest growing member of "series" Loba- tae found in Ecuador at altitudes of 2,000-3,000 m; the related P. sprucei and P. resticulata Mast. & André usually do not occur above 2,000 m. 500 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 2cm FIGURE 2. Passiflora montana Holm-Nielsen & Lawesson.— A. Flowering stem.— B. Longitudinal section of flower. 1987] Additional specimens examined. ECUADOR. PI- CHINCHA: Niebly, Pululagua, Sodiro s.n. (S). T dicas: Holm-Nielsen & Lawes- , Sp. nov. TYPE: Ecuador. Napo: Baeza- Tena rd., DEB 1 800-1, 900 m, 4 Nov. 1980, G. Harling & L. Andersson 16216 (ho- lotype, GB; isotype, AAU). Figure 3. a admodum glabra superficie foliarum. Caule striato, tereto. Stipules reniformibus, 1 x 2 cm, lobus nf obtusus, superior lobus cuspidatus, mucron onga, margine glandulare serataque. Petiolae teretae, 3 cm longae, 4-glan , 1-2 mm gis, al- ternis; lamina basifixa, cordata, integra, cir scrip tione deltoidea, 8 x 9-10 , 5S-7-n a, coriacea Flores solitarii lateralesque; SU. i 5-6 c longis; brac infr t erticilla- tae, liberae, ovatae, acuminateae, , cu margine glandulare serrata. Flores 7-8 b longis, consistunt, tubu secunda usque ad operculum obtegentibus. Operculum membranaceum, erectum, dense plicatum, 8-9 mm lon ngum, MT spathulato-ligulata 3 mm infra operculum. Limen arcte = Sessea “== gynophori, margine lobata. Ovario glabro. Fructu ovoideo. Semina non matura. Liana, stems sparsely puberulent, leaves pu- bescent at lower and glabrous at upper surface. Stem terete, striate. Stipules reniform, 1 x 2 cm, the lower lobe obtuse, the upper lobe cuspidate with a mucro 2 mm long, the margin glandular and serrate. Petioles terete, 3 cm long, with 4 stipitate, alternate glands 1-2 mm long; blades basifixed, cordate, entire, deltoid, 8 x 9-10 cm, 5—7-nerved, coriaceous. Flowers solitary, lateral, 7-8 cm wide; pedicels terete, 5-6 cm long; bracts verticillate, ee inserted 1 cm below flower, ovate, acum , 0.7 x 0.3 cm, margin glan- dub am NM onim short-campanulate, 1 x 2em; calyx lobes and petals ovate to oblong, violet, 3-4 x 1-2 cm; calyx lobes carinate with- out an awn. Coronal filaments, violet, in 5-8 series; outer 2 series of ligulate, 2.5-3 cm long filaments with filiform apices, the second series ongest; innermost 3-6 series irregular, of spat- ulate to ligulate or tuberculate filaments 1-5 mm long, covering the interior of the hypanthium from the second series to the operculum. Oper- culum membranaceous, erect, densely plicate, 8— 9 mm long, the upper part split into spatulate or HOLM-NIELSEN & LAWESSON — NEW ECUADOREAN PASSIFLORA SUBG. PASSIFLORA 501 ligulate segments. Nectar ring a horizontal ridge 3mm below operculum. Limen tightly surround- ing base of gynophore, margin lobulate. Ovary glabrous. Fruit ovoid, 5.5 x 2 cm Passiflora deltoifolia 1s related to P. menisper- mifolia Kunth and P. crassifolia Killip in “series” Menispermifoliae. The new species differs from these two species by having leaves as broad as long and not longer than broad. Passiflora men- ispermifolia has three-lobed leaves, whereas P. deltoifolia and P. crassifolia have unlobed or in- conspicuously iiis leaves. The operculum is plicate and there are no awns at the calyx lobes of P. deltoifolia. The bracts of P. deltoifolia are smaller, 7 mm long and 4 mm broad (vs. 1-2 cm long and 5-8 mm broad) and not cuani hh or long acuminate, as in P. menispermifolia and P. crassifolia. In addition, the new species lacks the dense indumentum characteristic for P. menispermifolia and P. crassifolia. Passiflora pergrandis Holm-Nielsen & Lawes- of Cumbaraza, 900 m, 20 Apr Harling & L. Andersson 13771 (holotype, GB; isotype, AAU). Figure 4. Liana, ubique glabra praeter inflorescentiam atque varium. Caule ,l€ ase; laminibus psi fixis, anguste-ova- tis, SE 15-20 x 9-10 cm, obtuso-truncatis, crasso-coriaceis € — lateralis, ee non-terminalis, usque e ad 30 cm longa, c pluribus 15-16 cm latis floribus aus amet ieee: 11 x 1-4.5 cm. Inflorescentia interdum D modo paucis foribus pedicellis teretis, 3-9 cm longis, puber- nfra florem portatae, erin lide, cm, obscure lacin- ato- lobatae i in parte distale, obtusae in base, margo Ingracum p puberu- lis. Tubo inis campanulato 1x2 cm; lobi calycis aes bs 6 x 3.5—4 cm, obtusi, carinati cum aris- ta folia onga, quae apicem non excedit. Petalis lanceolatis ovatis, 5.5-6 x 2 cm, alba. Fila- a t 3 cm, cruciatim eim colore i um albidocanum tomentosum. Fructus non visi Liana, glabrous throughout except inflores- cence and ovary. Stem terete, striate, older parts 502 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 D A Š 5cm FIGURE 3. Passiflora deltoifolia Holm-Nielsen & Lawesson.—A. Flowering stem.— B. Longitudinal section of flower.—C. Operculum, densely plicate.—D. Fruit. 1987] HOLM-NIELSEN & LAWESSON—NEW ECUADOREAN PASSIFLORA SUBG. PASSIFLORA 503 5cm FiGurE 4. Passiflora pergrandis Holm-Nielsen & Lawesson.— A. Leaf and reduced inflorescence. — B. Old stem. — C. Inflorescence with several flowers and bracts.— D. Longitudinal section of flower with recurved operculum. 504 2 cm diam. Stipules reduced. Petioles 2-3 cm, with one pair of ovate, sessile, black glands 3 x l mm, | cm from base; blades basifixed, narrow- ovate, acuminate, 15-20 x 9-10 cm, obtuse to truncate at base, Laban pinnate-veined, entire, thick-coriaceous. Laminar nectaries ab- sent. A distal bud pe to a short shoot, this forming a conspicuous indeterminate inflo- rescence to 30 x 15-16 cm with several flowers (or sometimes reduced with few flowers), the lowermost flowers developed first, the flowers subtended by 4-11 x 1-4.5 cm reduced leaves; pedicels terete, 3-9 cm long; bracteoles verticil- late, free, inserted 5 mm below flower, ovate to oblanceolate, 5 x 4 cm, obscurely laciniate-lobed at distal part, obtuse at base, margin entire with several stout black glands. Hypanthium cam- panulate, 1 x 2 cm; calyx lobes oblong-ovate, 6 x 3.5—4 cm, obtuse, carinate with a foliaceous awn 4 mm long, not exceeding apex; petals nar- row-ovate, 5.5-6 x 2 cm, white. Corona fila- ments of 3 series; filaments of outer series mi- nutely setaceous, 2 mm long; filaments of second series stout, ligulate, 5 x 0.3 cm, cross-banded with white and dark violet; third series close to operculum, minutely tuberculate 1-2 mm. Oper- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 culum membranaceous, recurved, the margin with short fimbriate filaments. Limen tightly ad- herent to androgynophore between the promi- nent nectar ring and trochlea. Ovary densely whitish-gray tomentose. Mature fruits not seen. Passiflora pergrandis is a member of "series" Laurifoliae and is related to P. ambigua Hems- ley, which is also present in Ecuador. Passiflora pergrandis differs in several aspects from P. am- bigua, these being much larger inflorescence and longer pedicels, larger leaflike bracts, calyx lobes and petals only twice as long as broad (vs. three or four times longer), and much shorter outer coronal filaments. The awn in P. pergrandis does not exceed the apex of the calyx lobe as it does in P. ambigua. Passiflora ambigua is known from Central America. LITERATURE CITED n L. B., P. M. JORGENSEN & J. E. La- ESSON. Passifloraceae. In Harling and Anders- ñoraceae Publ. Field Mus. Nat. Hist., Bot. Ser 19: 1-613. THE GENUS ATTALEA (PALMAE) IN PANAMA! GREGORY C. DE NEVERS? ABSTRACT The two Panamanian species of Attalea (Palmae) are described and illustrated, A. iguadummat for the first time. Attalea (Palmae) is a poorly collected neotrop- ical genus of about 25 species centered in Am- azonian South America and reaching its north- western limit in Panama. When Bailey (1943) treated the palms of Panama the genus was not known from the isthmus. Attalea was last revised by Glassman (1977), at which time one species was recorded in Panama. Recent fieldwork has revealed an undescribed species of Attalea there. A circumscription of the genus in Panama is pro- vided, and the new plant is described and named A. iguadummat. Attalea Kunth, Nov. Gen. et Sp. 1: 309. 1816. TYPE: A. amygdalina Kunth, Nov. Gen. et Sp. 1: 310. 1816. Arborescent or acaulescent, monoecious. Leaves pinnate, the pinnae inequilateral at the tip, clustered in groups or evenly distributed along the rachis. Inflorescences either androgynous or staminate, bearing a pair of bracts, the pedun- cular bract large, woody, sulcate, terminating in a long or short rostrum, enclosing the inflores- cence in bud; staminate inflorescence branched to one order, the rachillae with many flowers, these disposed in 1 or 2 rows, or spirally arranged in dyads. Staminate flowers with 3 short, tri- angular sepals, and 3 valvate, lanceolate-apic- ulate, flat petals; stamens 6-10 (in Panama), shorter than the petals, the anthers straight, de- hiscing longitudinally. Androgynous inflores- cence with sessile or short pedicelled pistillate flowers. Pistillate flowers 2-4 cm long, with 3 sepals and 3 petals, the sepals and petals imbri- cate; stamens reduced to a prominent stamino- dial ring; ovary ovate; stigmas 3, apical. Fruit with exocarp thin, fibrous; mesocarp pulpy and fibrous; endocarp thick, hard, without fibers. Seeds 1-3, irregularly shaped. Attalea is characterized by its large size, sep- arate staminate and androgynous inflorescences, ng, ru Attalea is closely related to Scheelea, Orbignya, and Maximiliana, together comprising the sub- tribe Attaleinae, the genera of which are distin- guished from each other by characters of the an- droecium. Attalea allenii is abundant throughout San Blas and many plants produce fruits annually, yet in- florescences at anthesis are rarely seen. During two years of collecting in San Blas, the androgy- nous inflorescence was seen only once and the staminate inflorescence only three times. When Glassman (1977) prepared his preliminary treat- ment of Attalea, the androgynous inflorescence was unknown. Androgynous inflorescences of A. iguadummat (described below) are equally rare, while the staminate inflorescences, in contrast, The pollination syndrome of the genus is un- known (Henderson, 1986). The staminate inflo- rescences of both Panamanian species produce a strong musky odor at anthesis. KEY TO THE SPECIES OF ATTALEA la. Pinnae evenly ip along the rachis; mid- dle pinnae 6.5-7 ; staminate inflo- rescence 25.155. cm long; rachillae 12.5- ! The first year of fieldwork was made possible by a Smithsonian Fellowship; the second year was conducted under the a uspices of a contract with the Asociación de Empleados K una. Additional support was provided by thi Missouri Botanical Garden through its Flora of Panama project. Bruce Allen translated the Latin diagnosis. Hecho deis assisted with fieldwork. ri Botanical Garden, P.O. Box 9. St. Louis, Missouri 63166, U.S.A. Current address: California iss 29 A pe of Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A. ANN. MissouRi Bor. GARD. 74: 505-510. 1987. 506 18.2 cm long; flowers spirally arranged in dyads; stamens 8-10 iguadummat . Pinnae in groups of 3 - 5 along the rachis; middle pinnae 2-4 cm wide; staminate inflo- rescence 15-25 cm pee rachillae 1-4.5 cm long; flowers two-ranked; stamens 6 ... A. allenii c Attalea allenii H. Moore, Gentes Herb. 8: 191. 949. TvPE: Panama. Colón: Puerto Pilón, 10 m, Allen 4103 (holotype, MO; isotype, BH). Figure 1. Stem solitary, short or subterranean. Leaves 12-15, about 5 m long; petiole 60-80 cm long, 2.5 cm diam. at base, 2 cm diam. at apex; rachis 3.3-3.7 m long; pinnae 85-87 per side, arranged in groups of 3-5, linear, glabrous, inequilateral at tip, the margins ferruginous-lepidote; middle pinnae 75-95 cm long, 2-4 cm wide; apical 9— 10 pinnae regularly spaced, separated basally, coherent at the apex, forming 2 broad lobes about 10 cm wide at the terminus of the rachis. Inflo- rescences interfoliar, produced at ground level, erect, either staminate or androgynous; bracts 2; prophyll 30 cm long, 4 cm wide, the apex short and rounded; peduncular bract 35 cm long with a rostrum 7 cm long. Staminate inflorescence 1 5— 25 cm long; peduncle 8 cm long, 8-10 mm wide, brown furfuraceous at anthesis; rachis 23-26 cm eae demi about 16-28(—50), 1-4.5 cm long, —8 staminate flowers in 2 ranks. Sta- ee flowers with sepals 3, deltoid, 1 mm long; petals 3, valvate, lanceolate-apiculate, glandular, 9-13 mm long, 2-3 mm wide; stamens 6, fila- ments 2-3 mm long, the anthers straight, 4—6 mm long; pistillode minute. Androgynous inflo- rescence unbranched; peduncle 10-24 cm long, 1-1.4 cm wide; rachis 10 cm long; pistillate flow- ers disposed in triads with 2 staminate flowers. Staminate flowers of triads 6-7 mm long, with 3 sepals, these connate for less than 0.5 mm at base, triangular, 2 mm long; petals (5—)6, 4—5 mm long; stamens 6, erect, dorsifixed, the an- thers 1-1.5 mm long; pistillode minute. Pistillate flowers sessile, crowded on the rachis, 3.2 cm long, enveloped at the base by 3 triangular bracts 2 cm long and 1.6 cm wide; sepals 3, broadly imbricate, 2.7-2.9 cm long, 1.5-2 cm wide, ir- regularly lobed at apex; petals 3, imbricate, 2.5- 2.7 cm long, 1.8-2 cm wide, irregularly lobed at apex; staminodial ring 6-7 mm deep, minutely 6-lobed; ovary conical, 2.3-2.5 cm long, 1.2-1.4 cm wide at base, densely brown tomentose; stig- mas 3, arching, 8-11 mm long; ovules 3, basal. Infructescence with 7-24 fruits, these obovoid, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 6-7.5 cm long, 3.5-4 cm wide, the exocarp thin, fibrous, densely ferruginous-lepidote, appearing smooth; mesocarp 1.5-2 mm thick, fibrous; en- docarp 3-5 mm thick, bony. Seeds 1-3, irregu- larly shaped, conforming to the shape of the cav- ity in which they develop. Additional specimens examined. PANA BOCAS DEL TORO: Río San Pedro, Gordon 15c (MO). CANAL AREA: Coco Solo, Gentry 6298 (MO). COLON: Nombre de Diós, July 1911, Pittier 4237 (US); Santa Rita Ridge, Croat 15308 (MO); Santa Rita Ridge, 300-500 m, Gen- try 6556 (MO). PANAMÁ: El Llano-Cartí Rd., 12 Jan. 1981, Read et al. 81-57 (US). COMARCA DE SAN BLAS: El Llano—Carti Road km 16.7, 350 m, 9?19'N, 78?55'W, *igua" (Kuna), “mange” (Spanish), 4 Nov. 1984, de Nevers et al. 4152 (MO, NY); El uon Road km 19, 10 Aug. 1984, de Nevers 3639 (MO, PMA); same locality, 18 June 1986, de Nevers & Herrera 7954 (CAS, MO); same locality, 11 Mar. 1986, de Nevers et al. 7301 (MO); Cangandi, 30-150 m, 9°24’N, 79*8'W, 29 Jan. 1985, de Nevers et al. 4735 (MO); same locality, *igua kaa” or “‘igua sai la let" (Kuna), 10 Feb. 1986, de Nevers & Herrera 7191 (MO); Río Tiwar (R. Acla), 8^48' N, 77°40'W, 25-100 m, Sugden 624 (MO); Playón Hydro Camp 14, 200 m, Duke 11377 (BH); 2 km from Las Animas on rd. to Quibdó, 5?4'N, 76?47'W, King et al. 664 (BH, NY). VALLE: Buenaventura Bay, Agua- dulce Island, Moore et al. 9468 (BH); isi dise dank Moore et al. 9460 (BH); km 14 marker between Bue ventura and Bajo Calima, below 50 m, 3°56'N, 76°59' W, Croat 57552 (MO); Dindo area, Bajo Calima, 100 m, 3°59’N, 76°58'W, Gentry & Monslave 48429 (MO); Bajo Calima, Gentry et al. 40395 (MO); Río Calima, La Trojita, 5-50 m, 19 Feb. 1944, Cuatrecasas 16397 (US); Bahía de Buenaventura, Quebrada de San Joaquin, 0- 10 m, 20 Feb. 1946, Cuatrecasas 19948 (US). BOLÍVAR: Mun. Morales, cgfo. Norosi, camino a Tiquisionuevo, 130-200 m, Cuadros 2194 (MO). Attalea allenii is well known from the original description and many collections. It ranges from Panama to Colombia. In Panama it occurs in tropical wet forest (sensu Holdridge et al., 1971) on the Atlantic slope. The Kuna name is **igua," the leaves are used medicinally, and the imma- ture fruits are eaten. Attalea iguadummat de Nevers, sp. nov. TYPE: n 4 350 79?45'W, 24 Feb. 1986, de Nevers 7197 (ho- lotype, CAS; isotypes, K, MO, PMA). Fig- ure 2 Species nova A. victoriana Dugand similis sed flo- ribus masculis spiratim depositis, staminibus 8-10, fi- lamentis 4-5 mm longis, antherisque 2-3 mm longis 1987] FIGURE 1. Stem solitary, short or subterranean. Juvenile leaves 1.5-2 m long, 20-26 cm wide, undivided, obovate, the margins dentate with triangular teeth 1.5-2 cm long. Mature leaves 9-17, arching; pet- iole 55-82 cm long, 5.7-7.2 cm wide at base, broadly channeled adaxially; rachis 6.75—7.23 m DE NEVERS— ATTALEA IN PANAMA 507 dalli T € P a^ Lh A Infructescence of Attalea allenii, de Nevers 4152. long; pinnae 104-109 per side, evenly spaced, linear, glabrous, inequilateral at tip, the margins ferrugineous-lepidote; middle pinnae 144-148 cm long, 6.5-7 cm wide, with midvein raised adaxially and abaxially; apical pinnae 1.6-1.9 cm wide, 55-59 cm long, free. Inflorescences inter- A 508 foliar, produced at ground level, erect, either sta- minate or androgynous, both inflorescence types produced on the same plant; bracts 2; prophyll 20-25 cm long, encircling base of peduncular bracts; peduncular bract woody, prominently vertically sulcate, enclosing inflorescence in bud, splitting and opening flat at anthesis, erect, 115- 203 cm long, 30 cm wide, with prominent non- splitting rostrum 12-15 cm long. Staminate in- florescence 125-155 cm long; peduncle 90-110 cm long, 1.5 cm wide, covered in a n of fur- furaceous brown tomentum, this soon eroded away; rachis 35-45 cm long: rachillae 40—50, subtended by an acute, triangular bract 1-1.5 cm long, this striate when dry; rachillae with basal sterile portion 2.5-4.2 cm long, fertile portion 10-14 cm long, with scattered groups of minute scales, these bright white in dried specimens. Sta- minate flowers spirally arranged in dyads, each subtended by a minute bract 0.5-1 mm long. Sepals 3, 1-1.5 mm long, striate when dry; petals 3, valvate, 1.3-1.7 cm long, 1 mm wide, flat- tened, curved to recurved at the tip, sometimes straight or “S” curved, striate when dry, but not when fresh; stamens 8-10, about ' the length of the petals; filaments 2-3 mm long, separate and free; adjacent stamens occasionally with the fil- ments connate at the base for 1 mm or the filaments connate completely, divergent just be- low the anthers; tip of filament attached 'A-'^ way up the anther, the thecae united above the point of attachment, separate or occasionally united slightly below it; anthers 2-3 mm long, straight, dehiscing longitudinally; pistillode 1—2 mm long, 3-lobed. Androgynous inflorescence at anthesis not known; peduncular bract as in sta- minate inflorescence; peduncle 60-70 cm long, the rachis 35-45 cm long. Pistillate flowers short- pedicellate or sessile; sepals 3, 22-35 mm long, 15-18 mm wide at base, broadly triangular; pet- als 3, 40-45 mm long, acute; styles 3, apical, exserted from the petals; perianth persistent in fruit, the sepals chartaceous; petals chartaceous, margin thin, undulate; the staminodial ring en- larged, 1.1—1.8 cm deep, with 15-20 lobes, these 2-3 mm deep, 2-4 mm wide, commonly bifid. Infructescence with (3—)45—65 fruits, these 7— 10 x 4.5-6.3 cm, obovate, brown tomentose, with the styles and stigmas persistent; exocarp 1-2 mm thick, tough, fibrous; mesocarp 3-5 mm thick, fibrous; endocarp 2-8 mm thick, bony, smooth, without fibers; seeds 1-3, irregularly shaped; endosperm homogeneous. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 The specific epithet is derived from the Kuna name for the plant, *igua dummat.” It honors the Kuna Indians of San Blas, Panama, who have initiated a self-managed forest reserve and wild- life sanctuary on their tribal lands. Additional specimens examined. |. PANAMA. COLÓN: Santa Rita Ridge, 8 miles from Transisthmica Hwy., in primary forest along small stream, 420 m, 9?25'N, 79°40'W, 2 Feb. 1986, Hammel, McPherson & Merello 14387 (MO); same locality, km 22, in forest on ridges, slopes and in draws, 17 Feb. 1986, Hammel, Mc- Pherson & Roubik 14507 (MO); same locality, 9 Mar. 1968, pid 3405 (US); same PEE km 13.8, 13 May 1986, de Nevers 7734 (BH, MO, PMA). COMARCA DE SAN BLAS: Río Taindi, 9°25’ N, 79°11'W, 5 Apr. 1986, de Nevers & Herrera 7656 (F, MO, PMA). Attalea iguadummat is distinctive in its acau- lescent habit, broad leaflets evenly spaced along the rachis, large inflorescence, and dyads of spi- rally arranged staminate flowers with 8-10 sta- mens. Among acaulescent species of Attalea with pinnae evenly distributed along the rachis, A. iguadummat is similar to A. victoriana Dugand and A. nucifera Karsten. It differs from A. vic- toriana in its spirally arranged staminate flowers (vs. the staminate flowers disposed in 2 rows on one side of the rachilla), shorter staminate rach- illae (12.5-18.2 cm vs. 25 cm), fewer stamens (8-10 vs. 12-15), and sessile (vs. pedicellate) pis- tillate flowers. Attalea iguadummat is distinct from A. nucifera in its longer middle pinnae (144— 148 vs. 93-131 cm) and its glabrous staminate petals (vs. staminate petals reddish glandular). Attalea iguadummat may be most closely related to A. tessmannii Burret, the only other species with staminate flowers spirally arranged in dyads. Attalea tessmannii is an arborescent species from Amazonian Peru which differs from A. iguadum- mat in its branched androgynous inflorescence with dispu saben HOMES: Attalea pias- sabossu Bondar but they are o. in a single row, not spirally arranged, and it is arborescent. Attalea iguadummat is known only from the extremely wet Atlantic slope of the mountains between Colón and the western border of San Blas, Panama. This may be the wettest area in Panama (Myers, 1969; Anonymous, 1975). Fur- ther collecting may reveal additional localities in the Atlantic lowlands to the west in the Provinces of Coclé, Veraguas, or Bocas del Toro, or the palm may be truly endemic to the wet ocean slopes between Colón and the Mandinga River. The Taindi River locality of A. iguadummat 509 DE NEVERS— ATTALEA IN PANAMA 1987] AT Un bu li o. FIGURE 2. Inflorescence of Attalea iguadummat at anthesis, Hammel 14387, photo B. Hammel. lenii, *dummat" means big. When asked where the “big Attalea” could be found the informant mentioned the Iguagandi River, a Taindi tribu- tary. In Kuna “‘-gandi”’ signifies place of, imply- ing the Iguagandi River is the “river of Attalea.” was discovered via a clue from a Kuna Indian. During an ethnobotanical survey in San Blas in- formants were asked to name wild edible plants that they harvest. One man mentioned “igua dummat.” **Igua" is the Kuna name for A. al- 510 Kuna botanists state that the Mandinga River is the eastern limit of A. iguadummat, which has not been found further east in San Blas during two years of intensive palm collecting there. Im- mature fruits of A. iguadummat are eaten oc- casionally by the Kuna. LITERATURE CITED ANONYMOUS. 1975. Atlas Nacional de Panama. In- stituto rein Nacional “Tommy Guardia,” Panamá, Panam ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 BAILEY, L. H. 1943. Palmaceae. In Flora of Panama. nn. Missouri Bot. Gard. 30: 327-396. GLASSMAN, S. F. 1977. Preliminary taxonomic stud- ies in the palm genus Attalea H.B.K. Fieldiana, Bot. 38: 31-61. HENDERSON, A. 1986. A review of pollinanenaty sayas in the Palmae. Bot. Rev. (Lancaster) 52: 22 aes pic E ^ W. C. GRENKE, W. H. HATHEWAY, & J. A. Tost, JR. 1971. Forest Envi- ai in Tropical Life Zones. Pergamon Press, rk. w Yo Myers, C. W. 1969. The ecological geography of cloud forest in Panama. Amer. Mus. Novit. 2396: 1-52. NOVELTIES IN MESOAMERICAN MISTLETOES (LORANTHACEAE AND VISCACEAE)! Jos KUIJT? ABSTRACT ive new species of Loranthaceae (Cladocolea primaria, Psittacanthus angustifolius, P. minor, P. species of Viscaceae (Dendrophthora davidsei, D. (including the var. wurdackii (Rizzini) Kuijt) is proposed for what has usually been called P. calyculatus n in Mesoamerica. The latter name is now restricted to a different species endemic to Mexico The following novelties result from recent studies in connection with the Flora de Nicara- gua and Flora Mesoamericana. 1. Cladocolea primaria Kuijt, sp. nov. TYPE: Panama. Panamá: Cerro Jefe, 2 km along road to Altos de Pacora from junction with road to peak, low cloud forest, 800 m, Syts- ma & Knapp 4797 (holotype, MO; isotype, LEA). Figures 1, 2. Plantae glabrae, pauce ramosae; rami quasi teretes, insignis, petioli ad 15 mm Io bisexuales. I triadis 3- vel 4-paribus, supra pari singulo, tunc pari uno mon m ebracteolatarum et denique flore ter- minali sequentibus; triadae basales ad axillas a - gatae; inflorescentiae paribus nonnullis foliorum squamiformium, c m, m s e. Flo- res 4-partiti; petala dimorpha, 2- e onga; an- therae perpar viora filamentis brevissimis insertae ad ee ovarium 1.5 x 1 mm; stylus rectus, stigma capitatum. Fructus 6 x 4 mm, ruber, obscure violascens, ellipsoideus. Plants sparsely branched, twining, glabrous. Stems terete or slightly 4-ridged, often with con- spicuous lenticels when older, straight and rather apex mostly blunt or slightly apiculate; midrib conspicuous and running into apex; petioles stout, to 15 mm long. Inflorescences subtended by sev- eral pairs of thick, brown scale leaves, solitary in leaf axils, determinate, with 3 or 4 pairs of triads below and a pair above, followed by a pair of ebracteolate monads and a terminal flower, morphic, 2-2.5 mm sessile on the shorter petals and with very short filaments on the longer ones; ovary 1.5 x 1 mm; style more or less straight, the capitate stigma reaching the petal tips. Fruit 6 x 4 mm, red, becoming dark purple, elliptic in outline; calyc- ulus inconspicuous; embryo dicotylous, slender, the haustorial pole scarcely expanded. Additional specimens examined. PANAMA. PANAMA: Cerro Jefe, Clusia forest near radio tower, 900 m, D'Arcy & Hamilton 14817 (LEA, MO); in forest near road to Cerro Jefe near junction with road to Altos de Pacora, Mori & Kallunki 72763 (LEA, MO); Cerro Jefe, 6.6 mi. above Goofy Lake, disturbed cloud forest, 850- 900 m, Sytsma et al. 2839 (LEA, MO). Ait. Cladocolea r ] 7 id bl ficulties in generic assignment. When I mono- graphed Cladocolea (Kuijt, 1975), I proposed the notion that Struthanthus is polyphyletic, at least many species being derived from a number of independent sources within Cladocolea. Thus I spoke of connecting bridges, these in some cases characterized by species pairs, one member of which was placed in Cladocolea, the other in Struthanthus. With some very minor exceptions, this left Cladocolea as a genus with determinate spikes of monads, the flowers 4-, 5-, or 6-partite, and either bisexual or the species dioecious. Stru- panamensis, which has bisexual flowers a i ive di pcs are Sie to Dr. Karel U. Kramer, Zürich, for the Latin descriptions, and to the Natural Sciences and Engineering Res arch Council of Canada for continuing financial suppo ? Department of Biological E. University of Lethbridge, Lethbridge, Alberta TIK 3M4, Canada. ANN. Missounmi Bor. GARD. 74: 511—532. 1987. 512 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FIGURE 1. Cladocolea primaria Kuijt (Sytsma & Knapp 4797). Pendent branch. bracteolate monads), its inflorescence made up of mostly triads. The inflorescence of Struthan- thus is generally indeterminate, but in a few species it also bears two or four monads at the tip, followed by a truly terminal flower. Cladocolea primaria presents a predicament from which there is no completely satisfactory escape, for placement in either genus leads to the need of significant modification of that generic concept. As a species of Struthanthus it would be the first four-partite species, and the second one with bisexual flowers. If placed in Clado- colea, however, it is the first species which is truly triadic. I feel that placement in C/adocolea is more acceptable, although I cannot deny an ele- ment of arbitrariness in this regard. I continue to hold the view that union of the two genera would tend to obscure the complex relationships between them, and that these difficulties concern very few species. In fact, if the evolutionary re- lationships are as I have proposed, such difficult intermediate species would be expected. Viewed in the above context, C. primaria is closely related to C. /enticellata (Diels) Kuijt 1987] 1 ñ KUIJT— MESOAMERICAN MISTLETOES 513 YW b. Flower dissection.—c. Ma ture yt (S & Knapp 4797).—a. Inflorescence, the upper triads removed. — fruit. —d. Embryo. 514 (which it greatly resembles superficially) and C. roraimensis (Steyerm.) Kuijt, while in Struthan- thus it is especially S. leptostachyus (Kunth) G. Don and S. polystachyus (Ruiz & Pavon) Blume that are related. Our species represents the sec- ond Cladocolea reported for Panama and seems to be limited to the Cerro Jefe area. N š ph ney page nios Kuijt, sp. nov. TYPE: Costa Rica. ón: Cordillera de Talaman- ca, Atlantic ue unnamed cordillera be- tween the Río Terbi and the Rio Siní, 2,400- 2,750 m, elfin forest edge, Davidse et al. 28990 (holotype, MO). Figure 3. Planta ad circ. 15 cm alta, monoeca, erecta, olivacea, parvifoliata; cataphylla basalia nulla vel irregularia, ap- pendices basales in plano medio. Folia (ob)lanceolata, succulenta, ad 9 x 2 mm, acuta. Flores feminei ad partes superiores internodium fertilium, plerumque wes d “N 5 cm longae; internodia fertilia 2 vel 3, raro 4; flores uniseriati, ad 20 pro bractea eta Fru ee albus, cllipsoideus ad .5 mm diam., petalis patentibus Plants monoecious, to ca. 15 cm high, erect, olive green, small-leaved; basal cataphylls absent throughout in some individuals, irregularly pres- ent in others; basal appendages oriented in me- dian plane; young parts often with sparse, stiff, white bristlelike hairs. Leaves (ob)lanceolate, succulent, to 9 x 2 mm, acute. Spikes solitary and axillary as well as terminal, to 5 cm long including the peduncle ca. 7 mm long; fertile internodes 2 or 3, rarely 4. Flowers uniseriate, to 20 per fertile bract; female flowers in the upper part of the fertile internodes, not generally out- numbering the male flowers below. Fruit white, ellipsoid to globose, 1.5 mm diam.; petals spreading Additional specimens examined. COSTA RICA. LIMON: Cordillera de Talamanca, Atlantic slope, Valle m, Davidse et al. 28755a (MO). HEREDIA: open road side, Lent 3826 Dendrophthora davidsei is the sixth known ies currently in this paper. The others are D. gua- temalensis Standley, D. mexicana Kuijt, D. squamigera (Benth.) Kuntze, and D. terminalis Kuijt. No serious confusion is possible with those ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 species. The closest relative of D. davidsei, how- ever, is D. paucifolia (Rusby) Kuijt, which ranges from Bolivia to Venezuela. The major difference between D. davidsei and D. paucifolia is that si vegetative branches of the latter invariably a prominent pair of basal cataphylls placed high above the leaf axil (see Fig. 39 in Kuijt, 1986a). This is not seen in any of the ten individuals present in the two Davidse collections, but basal cataphylls are present on some (not all) vegeta- tive laterals of the Lent specimen. The latter looks extremely slender but appears to belong to the present species. A second and apparently con- sistent difference is that leaf size in D. paucifolia indles upwardly until the uppermost lateral te are subtended by leaf scales; even though perhaps a slight diminution takes place in D. davidsei, none of the leaves on the main stem ever reach scale size. The relationship of the two species is certainly very close, but separation ap- pears justified. Terminal inflorescences were not seen in the type but are represented in the otherwise iden- tical collection Davidse et al. 287 55a. In the type, s shown in Figure 3a, the tips of the larger auctio have remained static, giving the impression of buds with crowded small leaves. Such shoot tips undoubtedly will expand into new systems of inflorescences, and perhaps mark a seasonal change. In fact, in the third collection, Lent 3826, there are some branches where such a new expansion seems to have taken place, de- marcated from the older portion by several pairs of very small leaves. 3. r. aC Kuijt, sp. nov. TYPE: Costa a. Limón: Cordillera de Ta- lamanca, od slope, Valle de Silencio, along the Rio Terbi, 0.5-1.5 airline km W of the Costa Rican-Panamanian border, 2,300-2,400 m, Davidse et al. 28755b (ho- lotype, MO). Figure 4. Planta tenuis, monoeca, erecta, ramosissima, circ. 30 cm alta, ad basem squamata; caules inflorescentias gerentes haud ultra 1 mm crassi. Cataphylla basalia ad dispersi et femineis multo pauciores; bacca alba, m diam., ovoidea, sepalis clausis. Plants monoecious, ca. 30 cm high, profusely branched, slender, erect, squamate to the base. Inflorescence-bearing stems | mm or less in 1987] KUIJT — MESOAMERICAN MISTLETOES 515 FIGURE 3. Dendrophthora davidsei Kuijt.—a. Habit (Davidse et al. 28990). —b. Same collection, old inflo- rescence. — c. Terminal portion of compound inflorescence (Davidse et al. 28755a). 516 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURE 4. Dendrophthora talamancana Kuijt (Davidse et al. 28755b). —a. Habit. — b. Fruiting inflorescence. thickness; basal cataphylls absent throughout, the basal appendages in a median orientation. Spikes solitary, axillary only, to 15 mm long, the single fertile internode slightly longer than the pedun- cle, the apex acute. Flowers uniseriate, to 6 flow- ers above each fertile bract; male flowers irreg- ularly distributed and much outnumbered by the female flowers. Berry white, 2 mm diam., ovoid; sepals closed. This distinctive plant represents the third completely squamate Mesoamerican species of Dendrophthora, the others being D. squamigera 1987] (Benth.) Kuntze and D. terminalis Kuijt. It differs from the former in its extremely fine, much- branched stems and short fertile internodes, and from the latter in its size, blunt scale leaves, and lack of terminal spikes. It is conceivable that the unusually slender Friedrichsthal s.n. specimen from Guatemala, as cited in Kuijt (1961), will turn out to be this species. I know of no similar South American plants. A . Phoradendron fasciculatum Kuijt, sp. nov. TYPE: Panama. Chiriquí: Jaramillo Arriba, near Boquete, trail to Rio Palo Alto, 1,100 m, near paved road, hyperparasitic on Pho- radendron undulatum Eichl., in turn para- sitic on Psidium guajava, Churchill & Kuijt 5106 (holotype, MO; isotypes, BM, CR, EAP, MEXU, NY, LEA, PMA). Figure 5. Plantae erectae, glabrae, basi rami 4-6 e pulvino communi orientes; internodia vetustiora bicarinata, novella praecipue statu sicco leviter quadrangularia, ad 10 cm longa; rami laterales paribus cataphyllarum singulis. Rami solum novelli foliosi. Folia carnosa, 8 x 5 mm, apice rotundata, mox decidua. Monoeca; spicae paribus cataphyllorum sterilium vel nullis; internodia fertilia 5 vel 6. Flores masculi 1-3 ad apicem areae floriferae supra bracteam fertilem, feminei usque ad 12 pro bractea et iis suppositae, bi- vel triseriati. Fruc- tus n ovoideus, 3 mm diam.; petalis inconspicuis, claus Plants monoecious, erect, glabrous, fascicled from the base with 4—6 stems from a common cushion. Stems usually lacking basal cataphylls; internodes 2-keeled when older, somewhat quadrangular when young, especially when dry, to 10 cm long, stout; lateral branches with 1 pair of basal cataphylls ca. 4 mm above base, spread- ing when dry. Leaves fleshy, soon deciduous, 8 x 5 mm, the apex rounded, sides parallel, the base clasping. Spikes with 1 pair of sterile cataphylls or without, 5—6 fertile internodes sometimes pro- liferating terminally into a second series of youn- ger fertile internodes. Male flowers 1—3 at the tip of the flower area above each fertile bract, the female flowers to 12 per bract below them, bi- or triseriate. Fruit broadly ovoid, 3 mm diam., the petals very small, inconspicuous, more or less closed. This distinctive species undoubtedly belongs to the P. dipterum group of species, in whic hyperparasitism is the rule. Phoradendron fas- ciculatum is no exception, in that all plants seen of the type collection were growing on P. un- dulatum Eichler. It is impossible to tell at this KUIJT—MESOAMERICAN MISTLETOES 517 time whether this cluster of species is obligately hyperparasitic; plants growing near the base of a primary host may easily be mistaken for being parasitic directly on the host tree. Another fea- ture apparently held in common by these various species is that several stems originate from a bas- al cushion, as illustrated in Figure 5a (arrow). I add a comparable illustration of a small plant of Phoradendron dipterum Eichler from Nicaragua (Fig. 6), which happens to be parasitic on a leaf, the host again being a Phoradendron. A clear basal cushion is visible (arrow). A third example is Phoradendron aequatoris Urban from Ecuador (Kuijt, 1986a, fig. 2), which also has basal sprouting and is parasitic on a Phoradendron. That this is not axillary branching from the nodes ofa much shortened base is demonstrated by the usual lack of basal cataphylls, in contrast to what occurs elsewhere in the plant. The explanation of the basal cushion almost certainly lies in the original haustorial disk of the seedling. It has recently been shown in an un- related species of Viscaceae, Viscum minimum Harvey, that the margin of the haustorial disk regularly produces aerial shoots (Kuijt, 1986b). he same is true for Viscum album L. (Tubeuf, 1923), and for Ixocactus hutchisonii Kuijt of Loranthaceae (Kuijt, 1987). In Phoradendron, this feature would appear to have some taxonomic constancy in the group of species under discus- sion. Additional specimen examined. COLOMBIA. ANTIOQUÍA: highway between Uramita and Canasgor- das, on Phoradendron piperoides (H.B.K.) Trel., Bark- ley & Gutierrez 535457 (US 5. Phoradendron molinae Kuijt, sp. nov. TYPE: Nicaragua. Madriz: cut over cloud forest area on Volcán Somoto, 10 km S of Somoto, 1,400 m, Williams & Molina 20270 (holo- type, US; isotype, F). Figure 7. Internodia compressa carinataque, ad 6 cm longa; rami laterales ; pedunculus statu fructifero circ. 3 s; spica fructifera circ. 3 cm longa, interno- diis fertilibus 2 vel 3, floribus pro bractea sions tribus, uctus ovo- ideus, laevis, 3 x 2 mm, petala clausa. Plants dioecious (only the female seen), stems with compressed, keeled internodes to 6 cm long, basal cataphylls one very low pair on lateral 518 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FiGURE 5. Phoradendron fasciculatum Kuijt (Churchill & Kuijt 5106).—a. Base of young plant, ns sprouting from basal cushion (arrow).— b. Habit of older plant.—c. Older inflorescence, the basal portion fruit, the terminal portion proliferated and in flower 1987] KUIJT—MESOAMERICAN MISTLETOES 519 FiGuRE 6. Phoradendron dipterum Eichler parasitic on Phoradendron sp., Nicaragua (Stevens & Montiel 17931, LEA). The hyperparasite is sprouting from a basal cushion (arrow) attached to a leaf of the primary host. branches. Leaves to 10 x 4.5 cm; blade thin, more or less palmately veined, ovate, the base abruptly contracted into conspicuous, cuneiform petiole to nearly 1.5 cm long. Female inflores- cence solitary, less than 2 cm long, often with a sterile pair of cataphylls; peduncle ca. 3 mm long in fruit; fruiting spike ca. 3 cm long, with 2-3 fertile internodes and 3 flowers per fertile bract just above the middle of the internode. Fruit ovoid, smooth, 3 x 2 mm; petals closed. 6. Phoradendron nitens Kuijt, sp. nov. TYP Costa Rica. Cartago: east side of sip divide between Tres Rios and Cartago, on Euphorbiaceae, Kuijt 2465 (holotype, CR; isotype, UBC). Figure 19 in Kuijt (1964) and Figure 5 in Kuijt (19862). Planta magna, m osa, ramificatione saepe furcata; innovationes gelu cataphyllis basalibus Ramuli novelli aliquantum compressi, que a m dispositi; spica ad 4 cm longa, cataphyllis nullis, pipi m fertilia 3 vel 4, flo- 520 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FiGure 7. Phoradendron molinae Kuijt (Williams & Molina 20270). —a. Habit. — b. Fruiting inflorescence. — c. Basal cataphylls. res pro bractea 6-9, biseriati. — flavus, 3 x 1.5 mm, ellipsoideus; petala claus. Plants fleshy, monoecious, the branching mostly forked, where percurrent perhaps with a pair of intercalary cataphylls. Lateral shoots with a large pair of basal cataphylls just above the axil; stems somewhat compressed when young, becoming terete. Leaves to 15 x 7 cm or larger, thick and shiny when fresh, rigid, obovate to nearly elliptical; base tapering or abruptly con- 1987] tracted; petiole very stout and flat, to 6 mm wide. Spike to 4 cm long, lacking cape fertile internodes 3-4, with flowers 6-9 p act, bi- seriate; male flowers very rare, iu in series. Fruit yellow, 3 x 1.5 mm, ellipsoid; petals closed. Additional specimens examined. NICARAGUA. o desde “Santa Julia" A "La Él 95 1 m, es 680 (HNMN see jos spec- imens listed under ee a. (Presl) Ei- chler in Kuijt, 1964). This species, which in Mesoamerica has been erroneously referred to Phoradendron obliquum (Presl) Eichler, is often extremely difficult to sep- arate from vigorous specimens of P. robustissi- mum Eichler if only herbarium material is avail- able. In the fresh condition, its usually larger robustissimum is also strictly dioecious, but the rarity of male flowers in P. nitens can be very misleading. Phoradendron obliquum is presently placed in synonymy under Dendrophthora obli- qua (Presl) Wiens (Kuijt, 1986a). The species is now known from Panama, Costa Rica (Kuijt, 1964), and Nicaragua, and has been reported recently from Ecuador (Kuijt, 1986a, as Phoradendron #5, fig. 5). We may thus antici- pate that it will be found in Colombia. N . Phoradendron tardispicum Kuijt, sp. nov. TYPE: Panama. Chiriquí: bridge over Río San Felix, on Panama Hwy., 50 m, on Ficus along river directly S of bridge, Churchill & Kuijt 5107 (holotype, MO; isotypes, BM, LEA, MEXU, PMA). Figure 8 Plantae dioeca eae solum visae), vivae o scure virides, glabrae. Caules deinde bs üternódiá recta, 5-9 cm longa. "Ca taphylla basal defoliata aliquot pro nodo, spica quaque par bus cataphyllorum sterilium basalium internodiisque fertilibus circ. ie flores biseriati, ad 10 pro e fer- tili. Fructus ovoideus, luteo-viridis, 3 mm longus, pe- talis clausis KUIJT — MESOAMERICAN MISTLETOES 521 Plants dioecious (only the female seen), bright, S m di slightly flattened when young, soon becoming terete; internodes straight, 5-9 cm long. Basal cataphylls mostly 1 or 2 pairs, very inconspic- uous, if 1 pair present nearly axillary, if 2 pairs present the second pair to 20 mm above axil, rarely to 4 pairs. Intercalary cataphylls present, one pair between successive pairs of foliage leaves but irregularly distributed along the branch, ab- sent from some internodes, always inconspic- uous. Leaves amplexicaul, cordate, shiny when fresh, to 7.5 x 6 cm, rather thin; margin undu- late; venation pinnate but obscure. Inflores- 2 pairs of sterile basal cataphylls less than 5 mm above base, followed by ca. 7 fertile internodes. Flowers biseriate, to 10 per fertile bract, yellow- ish green, each fertile internode with stalk and flowerless tip 2-4 mm long. Fruit ovoid, yellow- ish green, 3 mm long, the petals closed. This is a remarkable species for its irregularly distributed intercalary cataphylls and for late de- velopment of inflorescences. It is difficult to see what known species might be related. As far as I am aware, only in Phoradendron paradoxum Urban from Venezuela do intercalary cataphylls alternate in occurrence, but there this pattern seems to be regular (Trelease, 1916). That species and the closely related P. fendlerianum Eichler, however, have triseriate flowers and long-peti- ut there 0-3 cataphylls may be present on an “internode”; nor does this species seem to be closely related to P. tardispicum. 8. Phoradendron zelayanum Kuijt, sp. nov. TYPE: Nicaragua. Zelaya: N of abandoned airstrip near Alamikamba, along tributary of Caño Alamikamba, 10 m, gallery forest among sa- vanna, on Symphonia globulifera L.f., Ste- vens 21717 (holotype, MO; isotypes, HNMN, LEA). Figure 9. nta monoeca, dichotoma, apice abortivo. Caules menus innovationes laterales cataphyllis € positis binis. Folia late ovata vel orbicularia, la 8 x 8 cm, palmato-venosa; petiolus validus, planus, supra expansus 522 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ficure 8. Phoradendron tardispicum Kuijt (Churchill & Kuijt 5107).—a. Habit. — b. Inflorescence. [VoL. 74 1987] KUIJT—MESOAMERICAN MISTLETOES 523 TS cm FIGURE 9. Phoradendron zelayanum Kuijt (Stevens 21717).—a. Habit.—b. Inflorescence. ternodia fertilia tria; flores 13-15 pro bractea fertili, biseriati; spica 4 cm longa; spicae hermaphroditae. Plants monoecious, forking, the apex aborting, the inflorescences and young shoots dull golden- yellow. Stems terete, stout; internodes to 8 cm long; the lateral shoots with one low pair of cata- phylls. Leaves broadly ovate to orbicular; blade to 8 x 8 cm, with 5 or 7 very conspicuous pal- mate veins running far towards the apex; petiole stout, ca. 8 mm long, flat and expanding distally. Inflorescences bisexual, lacking sterile cata- phylls, the peduncle 3 mm long, this followed by 3 fertile internodes each with 13-15 biseriate flowers per fertile bract, the entire spike 4 cm long 9. Psittacanthus eo Kuijt, sp. nov. TYPE: Nicaragua. Ma .5 km al S de San José de Cusmapa, X m, parasitando en un Pinus, Moreno 24419 (holotype, MO; isotype, HNMN, LEA). Figure 10. Caules acute quadrangulares. Folia bina, anguste fal- cata, tenua, ven natione pinnata; lami sistens. Pedunculi triaderum circ bracteis foliaceis ad 2 cm cm longa, antherae 6 mm longae. Stigma antheras su- perans, capitatum. 524 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ficure 10. Psittacanthus angustifolius (Moreno 24419).—a. Habit.—b. Tip of petal. [Vor. 74 1987] Stems sharply quadrangular. Leaves paired, narrowly falcate, thin; the blade to 17 x 2.5 cm, venation pinnate, the base acute, the apex slen- derly attenuate; petiole to 5 mm long. Inflores- cences terminal, consisting of 4 or 6 triads; triad peduncles ca. 1 cm long, the lowest ones with foliaceous bracts to 2 cm long; pedicels 1.5 cm long, with conspicuous terminal cupule, the ca- lyculus smooth. Bud stout, more or less straight or somewhat curved; base and tip 5 and 4 mm wide, respectively, the latter blunt. Petals orange, 7.5—8 cm long; inner part of flower hairless, the petal base 5 mm wide, without ligule; petal apices 4 mm wide, blunt, each with a fleshy, ligule-like median crest extending inwards. Stamens di- morphic; filaments attached at ca. 2.5 cm above petal base, 5 cm long; anthers 6 mm long. Ovary 5 x 6.5 mm. Stigma placed above anthers, very small, capitate. Fruit unknown. Additional specimen examined. Same data as type, Soza et al. 155 (HNMN, MO). Thi cican t dicti ti , known from what is essentially a single collection. Whether the species is restricted to Pinus, as ap- parently is Psittacanthus pinicola, can only be shown by further fieldwork. The two species can- not be confused, as P. pinicola is dyadic and has irregularly whorled, blunt, more leathery leaves. The type locality suggests that the species may well be present in neighboring Honduras. 10. Psittacanthus minor Kuijt, sp. nov. TYPE: Nicaragua. Matagalpa: SW slopes of Cerro El Pilón and adjacent Laguna Tecomapa, roadside, low thorn scrub and pastures on rocky slopes, on ant acacia, Stevens 9466 (holotype, MO; isotypes, HNMN, LEA). Figure 11 Plantae parva, PE teretibus, foliis binis. Folia ad 5 cm, ovata, gae, series duae vix longa; stylus petalis fere aequilongus; stigma aegre dis- tinguendum. Stems terete, phyllotaxy paired. Leaves thin, the blade to x 3.5 cm, ovate, the apex and base obtuse or nearly so; venation more or less palmate; petiole 3-5 mm long. Inflorescences ter- minal, consisting of 4—6 pairs of triads on pe- duncles ca. 12 mm; pedicels 10 mm long. Buds KUIJT—MESOAMERICAN MISTLETOES 525 straight, not dilated at base. Petals ca. 3.7 cm long, red-orange; ligular area weakly differen- tiated. Stamens dimorphic; the anthers 3 mm long, dorsifixed, the 2 series dpi overlap- ping, the filaments attached ca. 21 mm above the base, some 16 mm long. mel more or less cylindrical, 4 x 2 mm; style nearly as long as the petals; stigma weakly differentiated; calyculus smooth. Fruits ovoid, 1.5 x 1 cm, with con- spicuous calyculus, blackish. Additional specimens examined. NICARAGUA. MA- TAGALPA: Puertas Viejas, 2 km al N sobre la Carretera Panamericana, “San Vicente," 600 m, on Acacia, Mor- din 18288 (HNMN, LEA, MO); same, San Juanillo, 8 al SE de Ciudad Dario, 500 m, pos 2618 (HNMN, LEA de Carretera, quebrada, 460-480 m Moreno 1669. ino o Dari TiO- Presa Santa Dario, on Leguminosae, Grijalva 2693 (HNMN, LEA, MO). Psittacanthus minor is closely related to P. mayanus Standley & Steyerm., which appears to be limited to the Yucatán region, has quadran- gular stems, and fruits which are about half as large as those of P. minor. Psittacanthus ma- yanus is much smaller in general. 11. Psittacanthus pinicola Kuijt, sp. nov. TYPE: Belize. Belize: Manatee Pine Ridge, on pine, 1931-32, Gentle 82 (holotype, GH; isotype, MO). Figures 12, 13. Caules plus minusve teretes; folia symmetrica, ter verticillata, ad 11 x 2.5 cm, anguste elliptica vel lan- ceolata, apice rotundata, basi in petiolum circ. 5 mm longum angustata. Inflorescentiae laterales, ad nodos, umbellulas e gas 2 vel 3 formantes. Petala circ. 4 cm longa, rubra, apice luteo- viridescentia, medio au- uantum curvatum, ad latitudinem circ. 5 mm dilatatum, ad apicem tenuissimum , leviter curvatum, circ. 1.5 mm latum angustatum; beep circ. laevis. Stamina dimorpha; antherae dorsifixa paulo pilosae. Ovarium plus minusve merida 4.5 mm longum; stylus longus, basi aliquantum torsus; stigma ellipsoideum, subtiliter papillosum. Fructus el- lipsoideus, calyculo inconspicuo, 13 x 5 mm, saturate purpureus. Stems more or less terete, becoming coarsely fissured and blackish when old; leaves symmet- rical, in (often somewhat irregular) whorls of 3, to 11 x 2.5 cm, narrowly elliptical to lanceolate; apex rounded; base tapering into petiole ca. 5 mm long. Inflorescences lateral, axillary, often also on older, leafless stems, each being an umbel of 2 or 3 dyads; inflorescence peduncle to 13 mm long; dyad peduncles and floral pedicels 5-7 mm 526 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FiGurE 11. Psittacanthus minor Kuijt (Stevens 9466). —a. Habit, with immature inflorescence. — b. Mature ud.— c. Base of petal. —d. Fruit (Moreno 18288). FiGure 12. Psittacanthus pinicola Kuijt.—a. Habit with immature inflorescences (Pipoly 4013).—b. Same collection, portion of inflorescence with mature buds, a single petal shown separately.—c. Complete inflorescence just after anthesis (Gentle 82).—d. Mature fruit (Stevens 7600). 527 FiGURE 13. Psittacanthus pinicola (Gentle 82).—a. Leafy innovation.— b. Mature flower, the style and anther shown separately to the right.—c. Relative positions of anthers and style. 528 1987] long, the latter scarcely expanded at the tip. Pet- als ca. 4 cm long, red with yellow-green tip, or- ange in the middle, prominently ligulate at the base. Buds somewhat curved above, inflated at the ovary to a width of nearly 5 mm, tapering to a very slender, slightly curved tip ca. 1.5 mm wide. Stamens dimorphic nearly trimorphic (Fig. 130); filaments of the longer type attached 1.5 cm above the base, 1.5 cm long; anthers 3-4 mm long, dorsifixed, sparsely pubescent on back. Ovary more or less cylindrical, green, 4.5 mm long, 2 mm diam. below, expanding slightly above; calyculus smooth. Style 4.6 cm long more or less straight, but the base somewhat twisted; stigma ellipsoid, finely papillate. Fruit ellipsoid, with inconspic- uous calyculus, 13 x 5 mm, “deep purple." Additional specimens examined. BELIZE. BELIZE: western highway, Mile 30, beside track, on pine, White- a 2442 ies same, The de on pine, Whitefoord 562 (MO). DISTR. UNKNOWN: P. Cayo, in roadside SA on ridge area a overlooking l 000 ft. falls, in area ountain Pine Ridge area, Huston ZELAYA: Rio Troncera at junc- gallery RT in savanna, on Pinus caribea, Pipoly 4013 (HNMN, LEA, MO); near Tala Has and Puente Mango (over 1 0—60 m, pine savanna, on Pinus caribea, Stevens 7600 (LEA, MO); Comarca del Cabo, Kornuk Creek above Puente Pozo Azul, old bridge, Robbins 5831 (LEA, MO). NUEVA SEGOVIA: El Jicaro, 7 km sobre la carretera a Murra entrada al Quebracho, 00-620 m, on pine, Moreno 8305 (HNMN, LEA, MO). Other dyadic species north of Panama are P. sonorae (Watson) Kuijt, P. palmeri (Watson) Barlow & Wiens, P. nudus (Molina) Kuyt & Feuer, and P. ramiflorus (DC.) G. Don. Psitta- canthus pinicola is similar to the last species but seems more closely related to P. dichrous (Mar- tius) Martius (see Eichler, 1868, especially Pl. 5). Not only inflorescence structure and general ap- pearance indicate this affinity, but even the pe- culiar curvature of the stylar base is seen in both. 12. Psittacanthus rhynchanthus (Bentham) Kuijt, comb. nov. Loranthus rhynchanthus Bentham, Bot. Voy. Sulphur 102-103. 1845. TYPE: “Dr. Sinclair," Tiger Island (Hondu- ras, Bay of Fonseca) (K). Figure 14d-f. Psittacanthus chrismarii Urban, Bot. Jahrb. 24: 331. 1897. TvPE (here designated): Costa Rica. Foréts de Nicoya, Tonduz 13706 Fir dn. US; iso- lectotypes, CR, GH). Psittacanthus calyculatus auct., non (DC.) G. Don, Gen. KUIJT—MESOAMERICAN MISTLETOES 529 . 1834. TYPE: Mexico. Cuernavaca: UE i 150 (G-DO). An attractive Psittacanthus in which the buds are distinctively curved and beaked occurs throughout Mesoamerica, at low elevations from southern Mexico to Venezuela. In the past, this species has been called P. calyculatus (DC.) G. Don or, earlier, P. chrismarii Urban. After study- ing the types of both Loranthus rhynchanthus Bentham and L. calyculatus DC., I conclude that these are two distinct species. Consequently, the name P. rhynchanthus must be applied to the wide-ranging species mentioned above. True P. calyculatus seems limited to Mexico, the type originating from the area of Cuernavaca, further collections having been seen from Puebla and orelia. Notwithstanding their general similarity, the two species may be consistently separated mostly on the basis of floral features. The mature, un- opened bud of Psittacanthus calyculatus is very nearly straight and has a rather blunt tip; that of P. rhynchanthus shows a distinctive curvature in the distal portion, the apex being sharply acute, and more beaklike. Psittacanthus rhynchanthus has smooth pollen sacs behind which are borne long, conspicuous, reddish stamen hairs; the pol- len sacs of P. calyculatus are distinctly lobed, and stamen hairs are lacking. Furthermore, the stylar base in P. rhynchanthus bears low protuberances, and each adjacent petal base shows a ligule con- sisting of a low, V-shaped ridge; the stylar base in P. calyculatus is smooth, and ligules are ab- sent. Leaves of P. calyculatus tend to be smaller (to 8 x 4 cm), mostly less than twice as long as wide, and approximately symmetrical, while those of P. rhynchanthus are usually larger (to 12 x 4 cm), more than twice as long as wide, and strikingly falcate. In Venezuela, at least some individuals of the latter species have extremely narrow leaves; these plants belong to Psittacanthus rhynchanthus var. wurdackii (Rizzini) Kuijt, comb. nov. (P. calyc- ulatus (DC.) G. Don var. wurdackii Rizzini, Rod- riguésia 41: 15. 1976). I have not yet encountered the species from the Caribbean lowlands of Co- lombia, but it would be surprising if it were not present there. 13. Lione subtilis Kuijt, sp. nov. TYPE: ds nama. Coclé: near continental divide along is. road, 2.2 km beyond sawmill in forest above El Copé, 900 m, Hammel 998 530 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 . w— - — <... ne oe wget: weit ge we — -— RE 14. Psittacanthus calyculatus (DC.) G. Don (a—; Mexico, Puebla, slopes of Popocatepetl, Ugent et i 9, WIS) and P. rhynchanthus (Bentham) Kuijt (d- á MSN. Managua, Cuatro Esquinas, Moreno 4400, LEA). —a, d. Mature buds.— b, e. Petals.— c, f. Stylar ba (holotype, MO; isotypes, LEA, PMA). Fig- lamina ae E ad late ovata, 4-12 cm longa, ure 15. 2-6 ata, ap l mm. Plantae inconspicuae, subtiles; caulis gracilis, plus darum 2 vel constitutentes; pedunculus c minusve teres. Folia tenua; venatio manifesta, pinnata; ad 8 mm longus, triadarum ad 5 mm, uterque 0.5 mm ., 3 oa nd E $ ue TE R Ms E d H H FIGURE 15. Struthanthus subtilis Kuijt.—a. Leafy shoot (Croat 49004) and leaves (Croat 44584).— b. Inflo- rescence (Folsom & Lantz 1894). —c. Same collection, petals and style, male flower. —d. Same collection, petals and style, female flower.— e. Mature fruits (Hammel 998). 531 532 vel minus crassus; flores laterales paces pid 0.5 mm longis; ipsis bracterolaeque m e, caducae; petala 2-2.5 mm longa; stigma cristis indistinct pa- pillosis 6. Siete aurantiacus, subglobos diam., calyculo inconspicuo; pedicelli Puch later- aliu m ad 3 mm elo ongati. Plants inconspicuous, rather delicate, branched. Stems slender, more or less terete; stem roots occasional, thin. Leaves thin, blade narrowly lan- ceolate to broadly ovate, always with caudate tip, 2-6 x cm, the evident venation pinnate; petiole ca. 3 x 1 mm. Inflorescences pale green, solitary in leaf axils, subtended by 4 minute, probably caducous bracts, consisting of a raceme of 2 or 4 triads; inflorescence peduncle to 8 mm long; triad peduncles to 5 mm, both 0.5 mm or less thick. Lateral flowers on pedicels 0.5 mm long at anthesis; bracts and bracteoles minute, caducous. Petals 2-2.5 mm long. Stamens di- orphic; upper portion of sterile stamens pa- pillate; anthers 0.4 mm long; style 2 mm long, the stigma with 6 indistinct papillate crests. Fruits orange, nearly spherical, 5 mm diam.; calyculus inconspicuous; pedicels of lateral fruits elongated to 3 mm. Additional specimens examined. PANAMA. COCLE: near continental divide along lumbering road, 1.5 mi. Copé, Croat 44584 (CR, LEA, MO); El Copé- El Potrosa, Atlantic slope of Alto Calvario, 700-850 m, Folsom & Lantz 1894 (LEA, MO, PMA); along road f à and Coclesito (N of Pintada), 4 mi. N of Llano Grande, 600 m, Antonio 3575 (LEA, MO); trail from Cano Blanco del Norte to continental divide N of El Copé, on Hedyosmum, 400 m, Davidse & Hamilton 23654 (BM, LEA, MEXU, MO, PMA); El Copé, Atlantic side, 1,200 m, Antonio 1153 (LEA, MO); between conti- 2409 (EAP, LEA, MO); El Copé, along gravel road to ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 the right before sawmill, 800 m, Antonio 2207 (LEA, ; Alto Calvario cloud forest, 5.3 km above El Copé, continental divide, above sawmill, 930 m, Antonio 3044 (LE , PMA); above El Potroso sawmill at gare divide, 1,200-1, Ly m, d 1820 (LEA along bora road un beris Agricola Alto P Piedra, aln first major stream, 3 mi. from fork in road at school, 700 m, Croat 49004 (LEA, MO) This species was previously listed as S. aff. dichotrianthus Eichler (Kuijt, 1978). Struthan- aL JS L . L í 4 ç L 1). J F 1 Blume) are indeed related to S. subtilis, as is S. quercicola (Cham. & Schlecht.) Blume of western Panama to Mexico, but the extreme slenderness and small racemes of our plant, the leaf size, and especially the consistently caudate to acuminate leaf apex, leave little doubt that this is a distinct species. Struthanthus subtilis appears to be en- demic to the Coclé-Veraguas region. LITERATURE CITED EICHLER, A. W. 1868. ipid In C. F. P. Mar- tius, Flora Brasiliensis 30 y 1 Kuur, Jos. 1961 revision of Pos NR (Lo- ranthaceae). Wentia 6: 1-145. 964. A revision of qs Loranthaceae of Cos- ta Rica. Bot. Jahrb. 83: 75. The genus C m (Loranthaceae). J. Arnold Arbor. 56: 265-335. . 1978. Co mmentary on the mistletoes of Pan- ama. Ann. Missouri Bot. Gard. 736—763. 1986a. Viscaceae. Jn Flora of hon 24: 6b. Observations on establishment and of Viscum minimum (Vis- caceae). Acta Bot. 6. 987. Mi scellaneous mistletoe notes, 10-19. Brittonia (in press TRELEASE, 19 llinois Press Urba TUBEUF, 23. Vus EE der Mistel. Olden- bourg, Miei ; The, Genus Phoradendron. Univ. A REVISION OF DILODENDRON (SAPINDACEAE)! A. H. GENTRY? AND J. STEYERMARK? ABSTRACT Dilodendron of Brazil and adjacent regions and Dipterodendron of Central America and northern discuss their relationships, present a key to the three species Dilode t costaricense (Radlk.) Gentry & Steyerm. and D. elegans. (Radlk. ) Gentry & Steyerm. Dilodendron Radlk., a monotypic genus of the dry areas of subequatorial South America, is closely related to Cupania L. and Matayba Aubl., from both of which it differs strikingly in having bipinnate leaves. According to Radlkofer (1892- 1900, 1895) the other main differentiating fea- tures of subtribe Cupanineae are that the petals of Cupania and Matayba (as well as closely re- lated monotypic Vouarana Aubl. have two squamae, whereas those of Dilodendron lack squamae. Two other monotypic genera, Brazil- ian Scyphonychium Radlk. and Guianan Pen- tascyphus Radlk., have an intermediate bifid or emarginate petal scale. The final genus of sub- tribe Cupanineae, Tripterodendron Radlk., like- wise monotypic and restricted to Brazil, is unique in having tripinnate leaves and the small sub- cupular calyx and bisquamate petals of Matayba. eneric limits in subtribe Cupanineae are gen- erally not Geary defined, and C “paid and Ma- tayba, the only significant deed the only nonmonotypic genera) recognized by Radlkofer (1892-1900), are notoriously difficult to tell apart. When Radlkofer (1892-1900) published his Flora Brasiliensis treatment, Dilodendron was known from Brazil and Bolivia, and from a single sterile collection from Costa Rica. Dilodendron bipinnatum has also since been collected in Par- aguay and disjunct in the dry part of the Rio Urubamba Valley in Convencion Province of Cuzco Department, Peru (Macbride, 1956), but the Costa qepa collection was subsequently de- termined not to be congeneric. " Dipterodendron Radlk. is a small genus of three ly reported from Costa Rica, Panama, and northwestern Venezuela (Radlkofer, 1933; Steyermark, 1952; Croat, 1976). Dipterodendron was described by Radl- ndron kofer (1914) on the basis of three Costa Rican collections. He recognized two species separated by rather tenuous differences: leaflets smooth and drying bright green in D. costaricense Radlk. vs. leaflets dark green and papillose and appressed puberulous below in D. elegans (Radlk.) Radlk. The sterile Oersted collection now recognized as D. elegans had originally been described as a variety of Dilodendron bipinnatum in the Flora Brasiliensis by Radlkofer. A third species, D. venezuelense Steyerm. was described in 1952 from Merida State in north- western Venezuela, representing the first report of Dipterodendron for South America. Like that of D. elegans before it, the type of D. venezuelense was sterile. The Venezuelan plant was distin- guished from D. elegans by larger, more coarsely toothed leaflets 2.3—4.5 cm long and 0.6-1.5 cm wide, and because of its geographic disjunction. However, recent collections from Costa Rica also have leaflets reaching 4 cm long and 1.5 cm wide. Some Costa Rican collections have leaflets with coarse teeth and others with fine teeth. Coarse teeth appear to reflect juvenile state rather than a consistent specific difference. Moreover, we have recently closed the geographic gap by col- cense and D. elegans. While the vegetative dif- ferences— mainly a more strongly appressed- puberulous leaflet undersurface in D. elegans— might be inadequate to justify maintaining D. costaricense as a species separate from D. ele- gans, there are also previously unreported fruit ! We thank USAID (DAN-5542-G-SS-1086-00 from the Latin America and Caribbean Bureau Office of 2 oe Resources) and the Smithsonian Institution for support of the Peruvian fieldwork that led to this revisio 2 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. ANN. Missounmi Bor. GARD. 74: 533-538. 1987. 534 differences. Thus, we refer all Central American and northern South American material of Dip- terodendron to two rather than three very closely related pas both of which range from Costa Rica to northern Venezuela. The dA between Dipterodendron of northern South America and monotypic Dilo- dendron of subequatorial South American dry forests suggest that even more merging is in or- der. This study was initiated when one of us recently collected a bipinnate-leaved Sapinda- ceae tree as a tree plot voucher at the Tambopata Nature Reserve in geographically intermediate of the genus. However, the equally striking veg- etative similarity between the Peruvian plant and Dilodendron was subsequently discovered in the herbarium when an attempt was made to identify it to species. This led to an examination of the taxonomy ofthe entire complex, which has never een menogrepheg except for a _tecopying of Radlkofer 'searlie I in his posthumous (1933) Pflanzenreich treat- ment. It turns out that Dipterodendron was never adequately differentiated from Dilodendron in the first place. Radlkofer (1914), who had seen no flowers, suggested that Dipterodendron is inter- mediate between Dilodendron and Tripteroden- dron but differentiated it only from the latter (which has tripinnate leaves and a thick oily aril, and lacks saponiferous cells in the embryo) rather than from the former even though he had earlier referred the first Dipterodendron collection to Di- lodendron. Later (1933) he emphasized slight dif- ferences in radicle position. Aristeguieta (1973) questioned the validity of separating Diptero- dendron from Dilodendron, noting that according to the literature Dipterodendron usually lacks petals and has the radicle on the margin of the cotyledon, while Dilodendron has 3-5 petals and the radicle descending down the middle of the dorsal side of the cotyledons. He ultimately as- signed the Venezuelan material to Dipteroden- dron essentially on geographic grounds. Later, Steyermark (in herb.) identified sterile collec- tions from Bolivar State in eastern Venezuela as Dilodendron bipinnatum, which would virtually eliminate the geographic discontinuity. Although Dipterodendron might be retained as distinct on the basis of its apetalous flowers, ves- tigial petals are sometimes present. Moreover, in Dilodendron the very small petals are variable in ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 number, mostly 3-4 but sometimes making up a full complement of 5, and sometimes vestigial (Radlkofer, 1892-1900). In fact, male flowers can lack petals altogether, just as in Dipterodendron. Since number of petals and even their presence or absence is variable in the single species D. bipinnatum, there seems no compelling reason to separate Dipterodendron from Dilodendron on this basis. Th pl in the same genus otherwise similar species s differ in presence or absence of petals, e.g., Swartzia, Licania, or Combretum; moreover, other like Alectryon and Mis- chocarpus have both petaliferous and apetalous species. Although interpretation of floral sexual- ity in Sapindaceae from herbarium specimens is very tricky, in the case of Dipterodendron loss of petals (only in male flowers?) might reflect a shift to full dioecy. We conclude that Dipterodendron should be united with Dilodendron to reflect best their extreme similarity. Indeed we are suspi- cious that monotypic Tripterodendron, of which we have examined only sterile material, might also be congeneric with Dilodendron. At any rate, some collections of Dilodendron have incom- pletely tripinnate leaves or squamellate petals, and the other distinguishing characters of disk margin, aril consistency, and lack of Z cells in the embryo seem weak. pi eceden t for incl uding Dilodendron Radlk., Bees Math.-Phys. l. Konig]. Bayer. Akad. Wiss. München 8: 355. 1878; TYPE: D. nete Dipterodendron Radlk., Smithsonian Misc. Collect. 61(24): 5. 1914. TYPE: D. costaricense. Medium to large dioecious trees. Leaves al- ternate, bipinnate (rarely in part tripinnate), mul- tifoliolate, the leaflets sessile or subsessile, serrate or dentate. Inflorescence a narrow (often almost subspiciform) panicle, usually borne clustered near the end of a branch and flowering preco- ciously or with the newly expanding leaves. Flowers tiny, the sepals 5, the petals smaller than sepals, sometimes absent, when present variable in number and often in part vestigial, the stamens in minute 2-3-lobed stigma. Capsule 2-3-lobed, loculicidally (22)3-valved, the valves woody or subwoody, the 1—2(-3?) seeds ellipsoid, with a thin, shiny, dark brown testa, scarious aril (fide 1987] Radlkofer), and basal hilum. Embryo (fide Radlkofer) subcircinately curved, the thick car- nose cotyledons saponiferous, the radicle dorsal. KEY TO THE SPECIES OF DILODENDRON la. Outer margin of leaflet teeth convex, the leaf- midrib often opposite pinnae; Brazil to southern Peru l. D. bipinnatum . Outer margin ofleaflet teeth dg the leaf- lets not ciliate- -margine ed, the lower midrib and lateral nerves s or strigillose with a few appressed hairs on midrib nerves; flow- ers apetalous; calyx lobes acute; leaves with 8-16, usually ri due pinnae; Costa Rica to Venezuela and P 2a. Fruit trigonal-globose, at dehiscence the or re- c the outer surface dull, puberu ous, and the tertiary venation visibly iig ue EN 2. costaricense 2b. Fruit laterally compressed, at . the valves splitting to the base and r cellate, drying black and shiny, bed scat- (CIC IMINULW visit n the verruculose surface; leaflets densely strigillose beneath, the tertiary venation usually not evident 3. D. elegans 1. Dilodendron bipinnatum Radlk., Sitzungsber. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss München. 8: 355. 1878. svNTYPES: Brazil. Minas Gerais: St. Hilaire 1586, Martius s.n., Riedel 1090, Warming s.n. (US). Tree 8-20 m tall, to 40 cm dbh, the branchlets usually somewhat angled and/or longitudinally ridged, puberulous with both long and short (in part gland-tipped) hairs when young, becoming glabrescent, lenticels absent or minute and in- conspicuous. Leaves bipinnate with 5-10 subop- posite or alternate pinnae, rarely the basal leaflets foliolate, the usually alternate leaflets narrowly ovate or oblong-ovate, obtuse to narrowly acute, 1.5-9 cm long, 0.5-3 cm wide, smaller at base and apex of each pinna, the margin rather ciliate and deeply toothed, the outer tooth margin GENTRY & STEYERMARK—REVISION OF DILODENDRON 535 strongly convex, sometimes with 1 or 2 marginal notches (= doubly toothed), puberulous with erect or suberect (sometimes in part gland-tipped) tri- chomes, above glabrescent except the main veins, elow + persistently pubescent over surface but rtiary tion somewhat prominulous below, the petiole 6-12 cm long. Inflorescence a terminal fascicle of narrow panicles, usually borne in the axils of fallen leaves at apex of a leafless branchlet, 3-36 cm long, tannish puberulous with trichomes of different lengths, the flowers sessile or subsessile, borne singly or in widely spaced, few-flowered clusters along and at tips of the lower branches (only at apex of the much-reduced upper lateral branches), subtended by bracteoles. Flowers greenish to cream or yellowish, 2—3 (male) to 5 (female, fide Radlkofer) mm long; sepals 5, ovate, unequal, sparsely appressed-puberulous and lep- idote, the margins + ciliate; petals reduced, shorter than sepals, 3—4(-5), sometimes com- pletely lacking in male flowers, broadly obovate, contracted to basal claw, puberulous at least on 2 tiny lateral projections (= scales) near apex of basal stalk; disk glabrous pei for tuft of hairs between filament bases; stamens exserted, (6-) 8(-9, fide Radlkofer), idm from center of disk, the filaments ca. 3 mm long, much narrower at apex, the anthers 1-1.5 mm long, reddish; the female flowers (not seen, fide Radlkofer) with puberulous disk, short thick style, and obtuse 3-lobed stigma. Capsule trigonal-obovoid, 1.5-2 cm long, splitting open somewhat unequally to near base, 3-valved, the valves woody, 3-4 mm thick, pubescent inside, glabrous or glabrate out- side, drying black, the surface rugulose. aa so kaa examined. BRAZIL. BAHIA: 10 “aen iras, 500 m, (fr), Irwin et al. 31317 (F, NY, ire RITO FEDERAL: Bacia do Rio Sao Bar- tolomeu, Brasilia, (fl), Heringer et al. 4530 (MO, NY), (fr), Heringer et al. 5224 (NY), (fl), chile d et al. 7011 N of Veadeiros, (fl), Prance & Silva 58264 (MO, NY, X5 Zona ina Ap AEA corrego Maranhão, (fl), Pires et al. 9472 (F). MATO GROSSO: Campinapolis, (st), Haridasan 72 (F); 270 km N of Xavantina, 8 km E of base camp, (fl), Ratter et al. 1874 (NY). MINAS GERAIS: Ituiutaba, (fl), Macedo 761 (MO, US); 15 km de Grão Mogol, estr. Montes Claros- Grão Mogol, (fl), Pirani et al. s.n. (CFCR 880) (NY); Lagoa Santa, (st), Warming s.n. (US). prope Concepcion, (fl), Hassler 7393 BoLiviA. BENI: Lake Rogagua, 300 m, Rusby 1686 (NY). SANTA CRUZ: Río Yapacani, (fl), Kuntze s.n. of June 1892 (NY, US); Velasco, (fl), Kuntze s.n. of July 1892 (NY, US); Provincia del Sara, Montes de Dolores, Cantón Buena Vista, 450 m, (fl), Steinbach 2515 (NY); 536 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 0 80 ^70 : 60 50 m) FIGURE 1. Distribution of Dilodendron. Squares = D. bipinnatum, stars = D. costaricense, circles = D. elegans. Buenavista, Prov. Sara, 450 m, (fl), Steinbach 6479 (F, moniha, pao pobre, farinha secca. Bolivia: cuta. ERU. CUZCO: Santa Ana, 900 m, (fl), Cook & Gilbert Peru: sumbayllo hembra, sumbaillo. 1468 (US), 1617 (US); Potrero, Prov. Convención, 1,500 m, (fl), Vargas 8228 (US); Santa Ana, Prov. Con- 2, Dilodendron costaricense (Radlk.) A. Gentry vención, (fl), Weberbauer 5020 (F), (fr), Weberbauer & Steyerm., comb. nov. Dipterodendron cos- I tari Radlk., Smit! ian Misc. Collect. Vernacular names. Brazil: Maria pobre, ma- 61(24): 7. 1914. TYPE: Costa Rica. Alajuela: 1987] La Balsa de Rio Grande, Pittier 3645 (ho- lotype, US; isotype, US) Tree 14-35 m tall, to 45 cm dbh, with but- tresses and flat spreading crown, the branchlets glabrescently finely puberulous, minutely and in- conspicuously lenticellate. Leaves bipinnate with -14 i n the pinnae 5-32-foliolate (or the leaflets a the uppermost incompletely differentiated), the al- ternate leaflets elliptic to oblong-elliptic, obtuse to acutish when mature, often subacuminate when young, 1-4.5 cm long, 0.3-1.5 cm wide, slightly smaller at base and extreme apex of each pinna (or the terminal leaflets incompletely differen- tiated when young), the margin not ciliate, ser- rate with relatively even teeth (occasionally al- most entire and typically more deeply dentate when young), the outer tooth margin nearly straight, above glandular-punctate, puberulous on midvein, below glabrous or with very few inconspicuous trichomes near base, the tertiary venation visible and often + prominulous below; petiole 2-9 cm long. Inflorescences several per branch, 5-15 cm long, each a few-branched, nar- row panicle arising from axil of fallen leaf, borne below leaves near branch apex in fruit (presum- ably as in D. elegans when in flower), puberulous with crisped and subappressed trichomes. Flow- ers unknown, presumably 8-staminate from the vestiges at base of young fruits (fide Radlkofer). Capsule trigonal-globose (even when only 2 fer- tile seeds), 2-3.4 cm long and diam., splitting open only partially, 3-valved, the valves woody, 2—4 mm thick, densely pilose inside, puberulous and minutely raised lenticellate outside, drying dull brownish with tannish lenticels, the seeds bean-shaped, 1.8-2 cm long with a shiny brown testa and a basal hilum. Additional specimens examined. Co ALAJUELA: La Balsa de Río Grande, h Pittier py (US); El Coyolar, near San Mateo, 100 m, (fr), od 3681 (coll. Werckle) (US); El Coyolar, 240 m Standley 40056 (US); vic. of Capulin, Rio Grande d Tarcoles, 80 m, (st), Standley 40159 (US). GUANA- CASTE: Santa Rosa National Park, 200-300 m, 10*51'N, 85°37'W, (st), Janzen 10683 (MO), Liesner 4236 (MO). PUNTARENAS: Palmar Norte de Osa, 0 m, (fr), Allen 5738 (US ANAMA. CHIRIQUI: Progreso, (st), Cooper & Slater 280 (NY, US); W of San iU Limite, (st), Croat 22159A (MO). DARIEN: Cerr i, Rio Coasi, (seed), Duke 15629 (MO); Yaviza, (fo) pun 6589 (US). PAN- AMA: El Llano, (fr), Duke 5818 (MO); Río Tapia, (st), Kan GENTRY & STEYERMARK—REVISION OF DILODENDRON 537 Standley 28087 (US), 28282 (US); Juan Diaz, (st), Standley 30574 (US); Rio Tapia, (st), Standley 41186 (US). OLOMBIA. BOLIVAR: San Juan Nepomuceno, 200 m, dou 75°10'W, (fr), Cuadros & Gentry 3617 (MO). oco: Municipio de Riosucio, Peye, 60 m, (st), Forero 1781 (COL, MO). Noir BOLIVAR: 48 km NE del caserio Los Ro- s, 17 km de Upata, (st), Blanco 334 (MO, NY, VEN); Altiplanicie de Nuria, ESE of Villa Lola, 315 m, (st), Steyermark 86364 (NY, VEN). MERIDA: El Vigía-Pan- americana, 100-120 m, (st), Bernardi 2093 (NY). ZULIA: Dtto. ColóN, 14—25 km NO de Pto. Chama, (fr), Bunt- ing & Drummond 6324 (VEN); El Toro, 8 km SSO de El Consejo, (fr), Bunting & Alfonzo 7054 (V EN); Misión de Tucuco, 105-250 m, (fr), Ijjasz 88 (NY); La Cocha, Mun. Uribarri, (fr), Trujillo 12211 (F). Vernacular names. Costa Rica: /upinsacca. Panama: guavino. Venezuela: tamarindo de monte, machirio tamirindo. The seeds are said to be edible (Bernardi 2093). 3. Dilodendron elegans (Radlk.) A. Gentry & elegans Radlk., in Mart. Fl. Bras. 13(3): 597 1900. Dipterodendron elegans (Radlk.) Radlk., Smithsonian Misc. Collect. 61(24): 7. 1914. svNTYPES: Costa Rica. Alajuela: prope Alajuela, Oersted 4, 5 (C, not seen). Dipterodendron venezuelense uir. Fieldiana, Bot. 28: 346. 1952. TvPE: Venezuela. Merida: ara Mes Isidro Alto and Sani poses de Mora, 760- 1,800 m, Steyermark 56569 (holotype, F; dH VEN) Tree 8-25 m tall, the branchlets longitudinally striate-ridged or slightly angled, minutely pu- berulous with erect or subappressed trichomes, glabrescent, tl scattered or essentially lacking. Leaves bipinnate with 10-16 frequently opposite or subopposite pinnae, the rachis puberulous with crisped tri- chomes, grooved above; pinnae (5-)9-23-folio- late, the alternate to subopposite leaflets oblong- elliptic, obtuse to acute, 1-6(-7) cm long, 0.3-2 cm wide, smaller at apex and base of each pinna (or the terminal leaflets incompletely differen- tiated), the margin not ciliate, serrate with rela- tively even teeth, the outer tooth margin nearly straight, the upper surface rather glandular and venation usually not evident; petiole long. Inflorescences few-branched, very narrow, subspiciform panicles, typically arising in clus- ters at the end of a leafless branch from the axils 538 of fallen leaves and often accompanying a cluster of unexpanded new leaves, in fruit + clustered at the base of the now fully expanded leaves, 3- 26 cm long, tannish puberulous with + ap- pressed trichomes, the flowers mostly in subses- sile or short-stalked clusters along it. Flowers reddish, apetalous, the sepals 5, ovate, less than 1 mm long, puberulous; disk flat, densely pu- berulous; stamens exserted at anthesis, 6—7(—8?), the expanded filaments ca. 2.5 mm long, the short, thick anthers ca. 1 mm long; female flowers sim- ilar to immature male flowers, with the 1 mm long densely puberulous ovary tapering ap a ong, narrow style and surrounded by ca. 6-8 subsessile sterile stamens, the stigma saucy 2-lobed. Capsule compressed-obovoid, 1-1.8 cm long, 1-2 cm wide, splitting to base at dehiscence with the valves reflexed, 2-valved, the valves subwoody, ca. 2 mm thick, sparsely pilose or + glabrate inside, outside with sparse and incon- spicuous scattered trichomes, drying black (red when fresh) with a minutely wrinkled-verrucose surface, elenticellate; seeds mostly 1 per fruit, flattened ovoid, 1 cm long, with shiny dark brown testa and a tan basal hilum. Additional specimens examined. COSTA RICA. ALAJUELA: Camino de San Ramón, is Brenes 435la (NY); La Palma (San Miguel) de San Ramón, 900- l ,000 m, (fl), Brenes 5351 (F, NY); San Pedro de San JOS n o 675-900 m, (fl), Skutch 4850 (F, MO, NY), (fr), 4876 (MO, NY, US). VENEZUELA. BARINAS: Barinitas, (st), Bernardi 3337 (VEN). BoLIvAR: between Tumeremo and El Dorado, 29 km N of El Dorado, 220 m, (st), Steyermark 86570 (NY, VEN); savanna de los Chacharros, 4 km upstream from Raudal Cotua, É jp ga pad ais 86773 (NY, VEN); 2 km SE os , 30 km S of El deed 365 m, (st), a sma siai 8695 7 (NY, US, VEN). CARABOBO: : carretera Maracay-Magdaleno-Gui- gue, Cuesta de Yuma, 450 TRUJILLO: Cerro Go rdo, sand oil on ridge, 9?45'N, 70?15' m. 1,000 m, (fr), Steyermark & Carreno 111646 (MO, NY, US). zuri4: Dtto. Colón, carretera Ma- chique a Fria entre La Redoma y Placita, (D. Bun- ting & Alfonzo 6930 E PERU. MADRE DE DIO Tambopata, 12*49'S, 69°18'W, 280 m, (st), Gentry et P 46217 (AMAZ, MO, USM). Vernacular names. Costa Rica: lorito, galli- nazo. Venezuela: caro montañero. As thus constituted, Dilodendron is a small genus of three species with one species (D. bi- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 pinnatum) mostly in the subequatorial dry areas of the Brazilian shield and adjacent regions, a Central America, and a third (D. elegans) wide- spread from Costa Rica to Venezuela and Am- azonian Peru. Dilodendron elegans and D. cos- taricense are apparently ecologically separated. In Central America D. elegans occurs in wet for- est, whereas D. costaricense occurs in moist for- ° = B g: 5 Venezuela, D. costaricense occurs mostly in drier forests and D. elegans in wetter ones. Dilodendron provides an excellent example of the importance of sterile collections. Not only was the original Peruvian collection that led to this entire revision sterile, but so are most of the other collections of the former Dipterodendron. Of the 22 collection numbers of D. costaricense, all but six are sterile and six of the ten state records from which it is known, including the only report from Colombia, are based only on sterile collections; the flowers are still unknown. Described from a sterile collection, D. elegans is now known from 17 collection numbers from eight different states in three countries; however, only nine of the collections are fertile and the only records for several states as well as for the country of Peru are based on sterile material. collecting institutions) get over their prejudice against sterile collections. LITERATURE CITED ARISTEGUIETA, L. 1973. Familias y Generos de los Arb de Venezuela. Instituto Botanico, Min- i ra y Cri raca CROAT, T. B. 1976. Sapindaceae. /n Flora of Panama nn. Missouri Bot 19-5 MACBRIDE, J. F pindaceae. Fi Flora of Peru, Fieldiana, Bot. XIIIA): 291- RADLKOFER, L. 1892-1900. SERSA In Martius, mon Bras. a 225-654. Sapindaceae. In A. Engler & K. Prantl, Die ce SA Pfanzenfamilien 3(5): alin 366. New Sapindaceae from Panama an Costa Rica. Smithsonian Misc. Collect. 6104): 1-8. Sapindaceae. In A. Engler, Pflanzen- reich IV. 165: 1019-1274 (Heft 98e). STEYERMARK. J. Contributions to the Flora of Venezuela II. Fieldiana, Bot. 28: 346. UNA NUEVA ESPECIE DEL GENERO DIOSCOREA (DIOSCOREACEAE) DEL ESTADO DE QUERETARO, MEXICO! O. TELLEZ V ? v B. G. SCHUBERT? RESUMEN Se describe Dioscorea matudae, una nueva especie del estado de Queretaro, México. Se discuten sus características en relación al sentido seccional dado por Knuth en 1924. Como parte del proyecto “Desarrollo del Her- bario Nacional" en el Instituto de Biología, .M., se han hecho colecciones por dife- rentes regiones de México, una de éstas es en el estado de Queretaro, en donde fue encontrada una especie del género Dioscorea que después de un análisis, se determinó como nueva especie para la ciencia y la cual se describe a continua- ción. Dioscorea matudae O. Téllez & B. G. Schubert, sp. nov. TIPO: México. Queretaro: 4 km al E del poblado Arroyo Seco, carretera a Jal- pan, alt. 840 m, selva baja caducifolia, 20 Oct. 1982, P. Tenorio L. & C. Romero de T. 2265 (holotipo, MEXU; isotipos, A, ENCB, F, MEXU, MO, XAL) Herba tenella. Caules sinistrorsum volubiles. Folia flores 2-4(-6) quoque Jen pate 6, introrsa. Inflo- me ala abortiva vel vestigialiter semen circundanti. Herbácea trepadora glabra. Tallo sinistrorso 1-1.5 mm, ligeramente angulado a lineado. H jas (2.7-)3.5—5.8. cm de largo, (1.5—)2-4.3 cm de ancho, alternas, ovadas, la base cordada, el ápice agudo a larga y abruptamente acuminado; 7-ner- vias, prominentes en el envés, ocasionalmente escasa y cortamente serruladas, las más externas bifurcadas; peciolo (0.7—)1—2 cm de largo, lige- ramente angulado, cortamente serrulado. Inflo- rescencia estaminada 1 ó 2 racimos de cimas o panículas de cimas de 4-12 cm de largo por axila; t ; flo- res 2- 4(-6) porci cima; pedicelo ca. 1 mm de Lien: rrulado; bráctea ex- terior r1-2 mm de largo, 1 mm de ancho, ovado- lanceolada, acuminada; bráctea interior 1 mm de largo, 0.5-1 mm de ancho, ovada. Perianto 1-1.5 mm, verdoso a amarillento; tépalos 1— 1.3 mm de largo, 0.5 mm de ancho, oblongos a elipticos; estambres 6, 1 mm de largo, insertados en la base de los tépalos; anteras introrsas, las tecas coherentes; pistilodio 0.2-0.4 mm de alto, cónico a triangular, inconspicuo. Inflorescencia pistilada 1 racimo de 2.4—11 cm de largo por axila; raquis angulado, conspicuamente serrula- do; flores solitarias; Delicato 1-2(-3) mm de lar- go; bráctea exterior mm de ancho, ovado-lanceolada, acuminada; bráctea interior 1-1.5 mm de largo, 0.5 mm de ancho, lanceolada. Perianto 1-1.3 mm, verdoso a amarillento, los tépalos 1-1.3 mm de largo, 0.5 mm de ancho, oblongos a elipticos; estami- nodios 6, ca. 0.5 mm de largo, insertados en la base de los tépalos, anteríferos o no; columna estilar 0.8-1 mm de alto, los estilos bifidos, ro- llizos, delgados. Cápsulas 9-12 mm de largo y ancho, con dos de los lóculos abortados, en al- gunos casos incompletamente y no se producen semillas (con los dos lóculos completamente abortados 7-8 x 2-5 mm, o con los lóculos de- sarrollados 9-12 x 6-9 mm), suborbiculares a orbiculares, membranosas; pedicelo acrescente en el fruto 4-6 mm de largo, cortamente serru- lado; semilla 4 mm de largo, 3 mm de ancho, suborbicular, parda, solo una semilla en el lóculo desarrollado, el ala casi completamente abortada u ocasionalmente presente como vestigios cerca al hilo. ! Agradecemos al Biol. Pedro Tenorio por facilitar el material para el pua NONO: A] Dr. Fernando Chiang por su ayuda con la diagnosis latina. A Elvia Esparza por su magnífica ilust š en de Botánica, Instituto de Biologia, U.N.A.M., Apartado Postal ' 70- 367, 04510 México, D.F., Mex 3 Arnold Arboretum, Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. ANN. Missouni Bor. GARD. 74: 539-541. 1987. FIGURA 1. Dioscorea matudae (P. Tenorio L. & C. Romero de T. 2265).—a. Planta estaminada. — b. Cima. — c. Flor estaminada. —d. Planta pistilada.—e. Flor pistilada solitaria. —f. Flor pistilada. — g. Cápsula. —h. Semilla. 540 1987] Especie interesante y a la vez dificil de rela- cionar con las especies cercanas en el sentido seccional dado por Knuth (1924). Un conjunto de caracteristicas permiten distinguirla facil mente de cualquier otra especie. Estas son la pre- sencia de seis estambres introrsos insertados en la base de los tépalos; pistilodio cónico a trian- gular, conspicuo, semejante a los observados en la sección Macrogynodium Uline; el habito her- báceo delicado; los tallos sinistrorsos; las hojas pequenas y glabras, semejantes a las de algunas especies con tres estambres y tres estaminodios encontradas en varias secciones del subgénero Dioscorea Pax.; su cápsula con dos de los lóculos abortados; la semilla con el ala completamente a casi completamente abortada; y el pedicelo acrescente en el fruto. Estos caracteres hasta aho- ra conocidos de D. cyphocarpa Robinson ex R. Knuth y D. tacanensis Lundell de la sección Poly- neuron Uline, pero que difieren de D. matudae por sus tres estambres formando una columna estaminal insertada en el centro del toro erior evidencia con gran punbahilidad la presencia de estructuras homólogas en el gru- TELLEZ V. Y SCHUBERT—NUEVA DIOSCOREA 541 po. Burkill (1960) y Téllez et al. (en prep.) hacen referencia a estas estructuras, interpretándolas como paralelismo en diferentes líneas dentro del género, lo cual probablemente ha dificultado en gran medida determinar las relaciones de sus taxa en una forma objetiva. Con gran probabilidad ésta especie podria dar origen a una nueva sección por si misma o quedar El epiteto específico está dedicado al cud Eizi Matuda, quien contribuyó en forma sustancial al conocimiento de las especies mexicanas de éste género. BIBLIOGRAFIA BURKILL, I. H. 1960. The organography and the evo- luti tion of Dioscoreaceae. The family of the yams. Bot. 56: 319-412. I Dioscoreaceae. Pflanzenreich IV. 43:1-387 (Heft 87). TELLEz, V. O., R. MEDINA L., E. MARTÍNEZ O. & P HIRIART V La Relevancia de la Pali- nología en la Sistemática de Dioscorea (Diosco- reaceae SYSTEMATICS OF THE SOUTHERN AFRICAN GENUS HEXAGLOTTIS (IRIDACEAE—IRIDOIDEAE)' PETER GOLDBLATT? ABSTRACT Hexaglottis is a genus of six species occurring along the west and south coast of southern Africa, an area of predominantly winter rainfall. It is a m btribe Ho ae, whic is characterized by a cormous rootstock and secondarily bifacial leaves. Hexaglottis is defined largely y an unusual flower structure with shortly clawed subeq d completely divi filifo style arms. This revision includes three new species, H. na den sa H. genus Rheome. The ese scr di and history o ottis are discussed, and, owing a detai presentation of taxonomically important characiers includin chromosome cytology, Hexaglottis is analyzed cladistically Sm namaquana is suggeste nc an isolated and primitive relict and the sister species of the remainder of the genus. The so frican genus Homeria is probably the to a group of species of Moraea section Moraea. B while H. nana has x Hexaglottis is a small genus of Iridaceae tribe Irideae restricted to the winter rainfall area of southern Africa (Fig. 1). It has linear, bifacial, and usually channeled leaves; corms of the Mo- raea type composed ofa single swollen internode and apically rooting bud; and umbel-like inflo- ipidia) enclosed in large, op- "3 ° e ace of eee establishes its vite po- sition in the predominantly southern African subtribe Homeriinae OMA pu 1980) of the Old World tribe Irideae. The ñoral structure is distinctive and, although not unique, defines the genus. The subequal tepals have short erect claws and horizontally extended limbs, and the styles are short with branches divided almost to the base into paired filiform arms that extend outwards on either side of the subtending anther. The flowers are yellow and fugacious, lasting only a few hours. Additional features are firm, brown- to blackish- dines corm tunics and a basic chromosome nu Moraea nins (Conn, 1986) has a similar flower structure, but the flowers are blue, romosome n r in Hexaglottis is x = 6, = 10, a number shared with Rheome and basic "y Morae unlike Hexaglottis, and the basic chromosome number is x — 10. A second species, described y Bolus as H. nana, has flowers es- sentially identical to those of other species of Hexaglottis, but it has dark brown, unbroken corm tunics, unusual fasciculate rhipidia, and a base number of x = 10, B is here excluded from Hexaglottis. C phology (detailed below), and vegetative mor- phology indicate that it is related to the small genus Rheome, comprising R. maximiliani and R. umbellata, and probably also to Moraea lin- deri and M. margaretae, which recent unpub- lished investigation has indicated are closely al- lied to Rheome. The peculiar dpa eer type of flower is thus believed to have evolved independently three times. It defines H E only in combination with the vegetative and chromosomal features mentioned above. RELATIONSHIPS Hexaglottis is probably most closely allied, within the Homeriinae, to Homeria, and avail- ! Support for this research from the U.S. National Science wasan grants DEB 78-10655 and DEB 81- fric Compton Herbarium iid woe for the illustrations publishe ff C sch, in the sania of my ment of Enviro n Rourke and his staff, eldwork is acknowledged with gratitude. I also thank urator of African An. ibn Botanical Garden, P.O. Box 299, St. Louis, Missouri S U. Eg A. ANN. Missouni Bor. GARD. 74: 542-569. 1987. 1987] BZ | S GAN Bs AW E 1 H. lewisiae H. virgata H. riparia H. longifolia GOLDBLATT —SOUTHERN AFRICAN HEXAGLOTTIS URNI OE E TELE UG AZ MA S f ON A P D 543 nll y NU QUT 22 s S Ra eak: | <: T u. s N ES AL. D ZZ — — P DATES FIGURE 1. Geography of Hexaglottis. able morphological and cytological data suggest that the two genera have as a common ancestor (Fig. 5) probably a species or group of species of Moraea allied to M. flexuosa (Goldblatt, 1982). The characteristics that Homeria and Hexaglot- tis share include subequal tepals, the claws of which cup the lower part of the filaments, and style branches reduced from the elaborate flat- tened structures basic for Homeriinae (Gold- blatt, 1980, 1986) and probably for the entire tribe Irideae (Goldblatt, in prep.). The two gen- era also share a similar and derived karyotype with x = 6 comprising strongly acrocentric to subacrocentric chromosomes. Genome size (Goldblatt et al., 1984) is similar, 22-29 pg DNA in Homeria and 20.6 pg DNA in Hexaglottis namaquana, the only species of the genus for which this is known. Moraea flexuosa has a com- parable karyotype. Other members of Moraea that appear less closely related include a part of the heterogeneous section Moraea, x — 10, and section Polyanthes, x = 6, the latter distinguished by having blue to violet flowers (a derived con- dition in Moraea). The karyotype in section Polyanthes is also somewhat different in com- prising acrocentric and submetacentric chro- mosomes (Goldblatt, 1980). HISTORY OF HEXAGLOTTIS The taxonomic history of Hexaglottis has been described by G. J. Lewis (1959) in detail, and it is reviewed here briefly. The first of the species now admitted to Hexaglottis was described by Nicholas Jacquin in 1776 as Ixia longifolia. The excellent figure that now serves as the type of the species is unmistakably this Cape Peninsula species, often confused with the more common and widespread H. /ewisiae. Shortly afterward, 544 the younger Linnaeus described a second species as Moraea flexuosa (Linnaeus fil., 1781). Thi name is now regarded as nomenclaturally su- perfluous and illegitimate, and a new name, H. lewisiae, was proposed for the species in 1971 (Goldblatt, 1971a) (see discussion under this species). The very distinctive, late-flowering Hexaglot- tis virgata was described in 1791 by Jacquin, this species also being assigned to Moraea. As with Jacquin's earlier species of Hexaglottis, a fine illustration leaves no doubt about its identity. Thus, all three common southwestern Cape species of the genus were known and described by the beginning of the nineteenth century when E. P. Ventenat erected the genus in 1808. Ven- tenat made no transfers to his new genus, men- tioning only Ixia longifolia Jacq. by name, “Ixia longifolia Jacq. etc.," which leaves one wonder- ing whether he had further un in mind, The acceptanc Nn OPN Ic cnnn oa1mo. were made ‘for H. ut ayay by R. A. Salisbury (1812)and for H. virgata and the illegitimate H. flexuosa by Sweet (1830). One more species was collected in the nineteenth century, H. riparia, discovered by C. F. Ecklon & C. L. Zeyher (their Irid. 30), but E iia was consistently as- signed to H. flex The three PONENS, species of Hexaglottis were regarded as a single taxon by Klatt (1866) under the name Homeria spicata (Ker) Sweet, the type of which is conspecific with the earlier Homeria elegans (Jacq.) Sweet (Goldblatt, 1981). Later, Klatt (1882: 52, 1895: 159) recognized Hexa- glottis with H. longifolia (including H. lewisiae) and H. virgata. Baker’s (1896) definitive nine- teenth century floristic treatment of the Iridaceae in Flora Capensis is identical, but he understood Hexaglottis so inadequately (Lewis, 1959) that is work on the genus must be disregarded. Louisa Bolus added one more species to Hexa- glottis in 1932, the west coast H. nana, which, although common, was apparently only discov- ered in the 1920s. Hexaglottis nana as already outlined differs markedly i in its vegetative mor- Hexaglottis and is now excluded from the genus (for comparison of H. nana with H. lewisiae see Fig. 2). Lewis’s (1959) revision of Hexaglottis admit- ted four species to the genus and two new vari- eties, H. virgata var. lata and H. longifolia var. angustifolia, neither of which is recognized here. Collecting since the publication of Lewis’s re- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 vision, especially in the arid country to the north of the Cape Floristic Region, has substantially expanded the knowledge of Hexaglottis. I dis- covered a new species, H. namaquana, in the Spektakel Mountains of northern Namaqualand in 1974 and extended the range of H. lewisiae into these mountains west of Springbok. The dis- tinctive new H. brevifolia of northern Nama- qualand and the Richtersveld is now recognized, specimens previously having been placed in H. described from the Roggeveld Escarpment in the western Karoo. Lastly, plants treated as H. /on- gifolia var. angustifolia by Lewis (1959) have been re-collected and are regarded as a distinct species described here as H. riparia. The picture in 1959 of Hexaglottis as a small genus essentially of the Cape Floristic Region with minor extensions of H. lewisiae and H. vir- gata to the north into semi-arid Namaqualand has changed fundamentally. Hexaglottis must now be viewed as centered along the interior Cape West Coast with extensions south and east into the Cape Floristic Region. MORPHOLOGY Rootstock. Species of Hexaglottis have a corm ofthe Moraea type (Goldblatt, 1976b: 670, 1981: 428) consisting of a single swollen internode with an apical primordium from which both shoot and roots are produced. This organ is one of the two major specializations defining subtribe Homeriinae. The corm originates from an axil- lary bud near the base of the flowering stem. The corm tunics are basically like those found in H I d several species of M. l L Moraea and consist of a coarse open network of hard, wiry, dark brown to black fibers. In Hexa- glottis the mealy substance between the fibers often persists and clings to the fibers, imparting a lighter color to them. The outer tunic layers are usually paler in color and characteristically medium brown in many collections of H. virgata and H. lewisiae. The two moisture-loving species, H. riparia and H. longifolia, have softer-textured tunics, the outer layers of which become light brown and the fibers are relatively fine Leaves. The leaves are bifacial witha a sheath- ing base and more or less linear and channeled, this being the basic leaf type for Homeriinae. The leaves of Hexaglottis namaquana are the most distinctive, being relatively broad, almost pros- trate, strongly undulate, lightly twisted with the 1987] GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS Habit and flowers of Hexaglottis lewisiae (A) and for comparison the vegetative and floral mo Bises of ‘Hexaglottis’ nana (B). Compare for example, Figures 6 and 11. Habits x 0.5; single flowers full size; separated stamens and style branches x2. margins sometimes somewhat crisped. The leaf condition in H. namaquana must be regarded as derived from the much more common erect lin- ear leaf with straight margins found in the other species of the genus and thus represents one or possibly two autapomorphies. The leaves of H. virgata may be lightly coiled distally, a feature obscure in herbarium material. There are usually two to three foliage leaves per plant, but the num- ber depends on growing conditions, so that fewer leaves are produced in drier seasons. Plants al- ways produce more leaves in the greenhouse than in the wild. Under optimal conditions H. /on- gifolia, H. riparia, and H. virgata subsp. karooica have four or five leaves. The leaves are inserted fairly close together near or slightly below ground level, but in H. lewisiae subsp. secunda, the leaf or leaves may be inserted some distance above the groun Flowering stem, sheathing bract leaves, and 546 branching patterns. The flowering stem is more or less erect, ee so in Hexaglottis vir- gata, but willowy and nodding in H. longifolia and H. riparia. There may be up to three or four major branches, each stalked and bearing a few to several sessile lateral rhipidia (the inflores- cence units). The exception is H. namaquana in which all the rhipidia are stalked, presumably the ancestral condition, and thus terminal on the main or lateral branches. The sessile lateral rhi- pidia are a derived feature, and a synapomorphy separating the main group of species of Hexa- glottis from H. namaquana. A sheathing bract leaf with a closed sheath subtends each branch or lateral rhipidium. In the latter, the sheathing bract leaf resembles the in- florescence spathes, which it may completely conceal. Each sheathing bract leaf generally over- laps the one above, except in H. /ongifolia, the cauline internodes of which are comparans long. Lewis (1959) used this feature as an im portant character for distinguishing H. pee Rhipidia. The inflorescence units are of the basic type for Iridaceae, consisting of compressed cymose umbels enclosed in two large, opposed, sheathing spathes, the inner of which exceeds the outer except in the lateral rhipidia of H. longi- folia, where they are nearly equal, a presumably specialized condition. Individual flowers are pedicellate and subtended by a single membra- nous bract contained within the spathes. The flowers are produced serially, a few days apart, at which time the pedicels elongate to raise the flowers out of the spathes. In Hexaglottis brevi- tuba and H. virgata the pedicels are short and the ovaries are included in the spathes, but the flowers have a perianth tube that serves the same function as the pedicel in extending the flowers beyond the spathes. The spathes are initially her- baceous with dry attenuate apices, but towards the middle of the flowering season they begin to dry out and become light brown and chaffy to- wards the middle. The short pedicels, included ovaries, and perianth tube are important syn- apomorphies separating H. brevituba and H. vir- gata from the rest of the genus. Flower. The flower is almost uniform throughout Hexaglottis, except for the presence of a tube in H. brevituba and H. virgata, and is unusual in Iridaceae in the structure of the style and style branches and the relationship of the latter to the stamens. The shortly clawed tepals are pale to deep yellow and subequal, or those ofthe inner whorl are slightly smaller. The claws, 1-2 mm long, are erect and form a cup around ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 the base ofthe filaments, while the relatively long limbs spread horizontally. The filaments are united below for 1-3 mm into a column, at the apex of which the style divides into three branch- es, each opposite a stamen. The branches divide almost immediately to form two long, filiform, apically stigmatic arms which extend outwards more or less horizontally to either side of the subtending stamen. The flowers are short-lived, opening in the mid to late afternoon and fading about three hours later. The unusual structure of the style of this flower has led to the placement of all species with this character together in a single genus, but data presented in this revision indicate that H. nana, described by Louisa Bolus in 1932, is distantly related to the other species of Hexaglottis and must have acquired its Hexa- glottis-like flower by convergence. A similar flower has also evolved in Moraea hexaglottis (Goldblatt, 1986). The divided style branches and filiform ascending arms are the primary characters separating Hexaglottis from other genera of Homeriinae, to which should perhaps cally free filaments as in the putative relatives Moraea flexuosa and Homeria (Fig. 5). The free part of the filaments allows the long anthers to be displayed prominently and also well separated from the style branches which almost always in Homeriinae divide at the top of the united part of the filaments. As already mentioned in the paragraph dealing ovaries. In other species the pedicels are about as long as the spathes, and at anthesis the ovary is almost always exserted. Fruit. Capsules of Hexaglottis species vary considerably and are important in recognizing species and in assessing phylogenetic relation- ships. The capsules are typically exserted from the spathes and are basically ellipsoid in shape, as in H. namaquana and H. lewisiae subsp. se- cunda. In the latter, the capsule has a short beak, a feature not always evident in populations from dric capsules, while obovoid to clavate capsules distinguish H. riparia and H. longifolia. The cap- sules of H. longifolia are relatively large, 12-16 (7-23) mm long, but only 6-10(-12) mm long in H. riparia, the smaller size presumably basic and consistent with capsule dimensions of other 1987] TABLE 1. GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS 547 Chromosome numbers in Hexaglottis and Rheome. Original counts are marked with an asterisk. Previous counts were reported by Goldblatt (1971b, 1980). Haploid Species Number Collection Data HEXAGLOTTIS H. namaquana Goldbl. 6* Spektakel Pass, west of Springbok, Goldblatt 3059 (MO). H. lewisiae Goldbl. subsp. /ewisiae 6* Tulbagh Cemetery, Goldblatt 5224 (MO); Cape Town, Kir- stenbosch Gardens (wild plants), Goldblatt 5104 (MO). subsp. secunda Goldbl. 6 Loeriesfontein road, north of Nieuwoudtville, Goldblatt 108 (J). 6* Spektakel Pass, west of Springbok, Goldblatt 6513 (MO); gran- ite outcrops SW of Skuinskraal, near Hondeklipbaai, van Berkel 453 (MO); near Nieuwoudtville, Goldblatt 6535 (MO). H. riparia Goldbl. 6* Olifants R. bank at Citrusdal, Goldblatt 6555 (MO). H. longifolia (Jacq.) Sweet 12* Cape Town, Kirstenbosch Gardens (wild plants), Goldblatt 5934 (MO), Malan 120 (NBG). H. brevituba Goldbl. 12* Near the Kosies road, NW of Steinkopf, Goldblatt 5748 (MO). H. virgata (Jacq.) Sweet subsp. virgata 5 Signal Hill, Cape Town, Goldblatt 71 (J). 5” Slopes near parking area, Signal Hill, Cape Town, Goldblatt 6747 (MO); Signal Hill, near Sheik’s tomb, Goldblatt 6768 (MO); lower slopes of Devils Peak, Goldblatt 6717 (MO). 6* Hills west of Riversdale, Goldblatt 5436 (MO); Franskraal, near the coast, Goldblatt 5368 (MO); near Misgund, Gold- blatt 6792 (MO); between Bredasdorp and Napier, Goldblatt 6937 (MO); between Doorn River and Bidouw road, Gold- blatt 5941 (MO) subsp. karooica Goldbl. 7* Roggeveld, near Voelfontein farm, Goldblatt 6336 (MO); Blomfontein farm west of Middelpos, Snijman 765 (MO, NBG) Olifants River valley near Alpha, Goldblatt 5120 (MO); Pak- ‘Hexaglottis’ nana L. Bolus 10* huis Pass, near Soldaat Kop, Goldblatt 5158 (MO). RHEOME R. maximiliani (Schltr.) Goldbl. 10 Brandewyn River near Travellers Rest, Goldblatt 3884 (MO). R. umbellata (Thunb.) Goldbl. 10* East end of Du Toits Pass, Goldblatt 5907 (MO). 15 Paarl Golf Course, Goldblatt 4414 (MO). 15* Piketberg, top of Versveld Pass, Goldblatt 5163 (MO); foot of the Elandskloof Mts. at Elandsberg farm, Goldblatt 5853 (MO) species. The capsules of H. virgata and H. brevi- tuba are included in the spathes and are narrowly fusiform. They remain enclosed by the spathes through ripening and dehisce only in the upper part The seeds are brown and basically angular, but elongate in H. virgata and H. lewisiae subsp. lew- isiae, both of which have narrow capsules (seeds of H. brevituba are not known). Seeds of H. lon- gifolia are unusually large, a feature possibly re- lating to its polyploid state. Hexaglottis lewisiae subsp. secunda can usually be distinguished by the raised and winglike angles of the seeds. The seeds are known from only a few populations of this subspecies, and I hesitate to regard the wing- like angles as characteristic of the taxon until more is known about their occurrence. CHROMOSOME CYTOLOGY The cytology of Hexaglottis was investigated extensively for this study. The method followed here is the same as that outlined for similar in- 548 subsp. /ewisiae.—C. H. lewisiae subsp. kraal, Goldblatt 5368). —F. subsp. Pb us =12 (Doorn 10 (Signal Hill, Goldblatt 674 J. ‘Hexaglottis’ nana, 2n = fed Scale = 10 um vestigations in Iridaceae (Goldblatt, 1979, 1980). The results are presented in Table 1. Base num- ber in Hexaglottis is x = 6, this originally sug- gested on the basis of one count for H. lewisiae (as H. flexuosa), 2n = 12 (Goldblatt, 1971b), while a single count for the specialized H. virgata, 2n — 10, suggested that this species was a derived aneuploid. Several additional counts for Hexa- glottis have confirmed x = 6 as basic. Hexa- ANNALS OF THE MISSOURI BOTANICAL GARDEN FIGURE 3. Mitotic ns cla ne sa of j ge dia species. — unda. — D. H. riparia. — E. H. H. virgata subsp. karooica, 2n = 14 [Vor. 74 C —— N CS < ! L NT —A. H. namaquana.—B. H. lewisiae virgata subsp. virgata, 2n = 12 (Frans- R. BAUN. Goldblatt 5941).—G. subsp. virgata 2n = —I. Rheome umbellata, 2n = 20.— glottis lewisiae, H. namaquana, H. riparia, and three populations of H. virgata all have 2n = 12. The two known populations of H. virgata subsp. karooica both have 2n = 14, while H. longifolia is tetraploid with 2n = 24. The two populations of Hexaglottis nana ex- amined cytologically were found to have a dip- loid number of 2n = 20 and a karyotype exactly like that of the diploid Rheome maximiliani 1987] (Goldblatt, 1980). Basic chromosome number in Rheome, already established (Goldblatt, 1980) as x = 10, has been confirmed here, with addi- tional counts for R. umbellata. This species was originally thought to be triploid, 2n = one diploid population has been discovered, as well as two more triploid populations (Table 1). The chromosomes of Hexaglottis species are fairly large, ranging in size in the basic karyotype from 5 to 9 um with the method used here. The basic karyotype as exemplified in the least spe- cialized species, HE wamaqa, consists Bra eid Ww eo + a satellite on the distal end of the short arm ane the third or fourth longest pair (Fig. 3A). Hexaglottis lewisiae subsp. lewisiae and H. ri- paria (Fig. 3B, D) have a similar karyotype, but the satellite is located on one of the longest chro- mosome pairs. Size differences are relatively small, and the shortest chromosomes are only +35% smaller than the longest (Fig. 3A-C; see also Goldblatt, 197 1b: 364, fig. 14E). Hexaglottis A is tetraploid, 2n = 24, but otherwise has a karyotype comparable to that of G. riparia and G. lewisiae subsp. lewisiae. The Namaqualand populations of Hexaglottis lewisiae subsp. secunda (Fig. 3C) have an ap- parently derived karyotype. The longest chro- mosome pair is metacentric and about 12.5 um long, nearly twice as long as the next in size, an acrocentric pair. The third or fourth pair is sub- metacentric, while the smallest pair has a large satellite (Fig. 3C) and is only 5 um long, about one-third as long as the long metacentric. This karyotype has been found in two widely sepa- rated Namaqualand populations of this poorly sampled subspecies (Table 1), but a population from Nieuwoudtville, well to the south, has a karyotype of acrocentric chromosomes, unusual only in having a satellite on the end of a long arm ofa long chromosome pair. The single plant that I examined was structurally heterozygous, having only one satellite present. In the specialized Hexaglottis virgata, there is unexpected intraspecific variation in the karyo- type. The presumed basic karyotype (Fig. 3E) as found in southern Cape populations of subsp. virgata consists of six pairs of acrocentrics, the first and third of which have a distinctly longer short arm. The satellite is located on the second longest and strongly acrocentric pair. A northern population of H. virgata (Goldblatt 5491) can be distinguished cytologically by having satellites on the distal end of the long arm of the longest GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS 549 and strongly acrocentric pair (Fig. 3F). The northern populations may be a separate cytolog- ical race, but more material needs to be exam- ined. Cape Peninsula populations of H. virgata are aneuploid, 2n = 10 (Goldblatt, 1971b), and have a karyotype exhibiting considerable struc- tural rearrangement. The longest pair (Fig. 3G) is metacentric and about 12 um long, while un- usually large satellites are located on the shortest and acrocentric pair, the satellite being longer than the short arm. Robertsonian fusion of two acrocentric, medium-sized pairs and the trans- location of the satellite to the smallest pair would account for the modified karyotype. Hexaglottis virgata subsp. karooica is unusual in the genus in its diploid number of 2” = 14 (Fig. 3H). Three individuals of both known pop- ulations were examined. The chromosomes are more strongly acrocentric than in the basic 27 = 12 cytotype of subsp. virgata and there are two, rather than a single, small pairs. The origin of the extra small pair is unknown. Satellites in the subspecies are located on the ends of the short arms ora a large Wa dom pair. Kar lottis appears to have ded from a en rather uniform, acrocentric set of chromosomes to increasing mmetry with the development of greater size ieee and, in H. lewisiae subsp. secunda and the Cape Peninsula populations of H. virgata subsp. virgata, the evolution of large metacen- trics, in the latter with a decrease in base number o x = 5. Hexaglottis virgata subsp. karooica seems to be one ofthe rare examples in Iridaceae of an increase in base number, as the karyotype of this specialized and rare taxon is almost cer- tainly derived from ancestors with x — 6. The origin of the extra pair of small chromosomes is problematic. The tetraploid H. /ongifolia may have evolved by amphipolyploidy, and its large size, especially in vegetative and fruit characters, may be a direct result of its polyploidy. Hexa- glottis brevifolia has the same base number as the genus but details of its karyotype were not seen in the poor material available. 2 c HYBRID STUDIES A crossing program involving three species of Hexaglottis, H. nana, and Rheome maximiliani in the spring of 1982 produced results (Fig. 4) that confirm the indications from cytology that H. nana is allied to Rheome rather than to Hexa- glottis. Rheome maximiliani could be crossed readily to emasculated flowers of H. nana, while 550 HEXAGLOTTIS H. virgata RHEOME GURE 4. panies relationships in Hexaglottis, ‘H? nana, and me maximiliani. Heavy lines indicate ncc oe dotted lines indicate crosses were attempted but failed. repeated attempts to cross both of these species with three species of Hexaglottis used in the study failed. Attempts to make interspecific crosses be- tween other species o quana, but all attempts to cross H. virgata with these species failed. The study was not extended to the species of Moraea that are most similar to Rheome, namely M. linderi and M. marga- retae, as neither was available in cultivation. REPRODUCTIVE BIOLOGY AND POLLINATION The Hexaglottis flower is relatively small and times depending on the species or population, in mid to late afternoon and closing in the early evening. All species except H. longifolia are strongly self-incompatible (H. brevifolia un- known). Very rarely a few undersized capsules are produced on plants by autogamy late in the flowering season, but normally flowers do not set seed by their own pollen, even though a small ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 quantity of pollen is usually deposited on the stigmas while the flowers are open. The excep- tion, H. longifolia, is self-compatible and autog- amous. Plants produce numerous full capsules with fertile seed without cross-pollination. The pale to deep yellow stellate flowers of Hex- aglottis are typically pollinated by bees. The small nectaries, located at the base of the outer tepals, marily to the pollen, a large amount of which is produced in the relatively long anthers. Although the flowers of all species are very similar, there is a considerable difference in the response of bees to the flowers ofthe three species, H. lewisiae, H. virgata, and H. lon longifolia is totally unattractive to bees, which ignore open flowers even when they pass close to them. The same bees, however, visit the open- ing flowers of H. lewisiae subsp. lewisiae growing within a few meters of H. longifolia. Bees gather around populations of H. lewisiae in the middle of the afternoon about the time its flowers nor- mally open, and den begin to gather pollen as soon as the flow pen. Hex aglottis bris is similarly attractive to and species of Anthophoridae were observed pollinating Hexaglottis flowers. PHYLOGENY Cladistics affords the most objective and crit- ical method of assessing the phylogeny of a group, and the results of a cladistic analysis of Hexa- glottis and its immediate allies are presented be- low. The cladogram (Fig. 5) was constructed manually following concepts of clustering by shared derived characteristics (synapomorphies) and parsimony established by Hennig (1966) and adapted by several botanists recently (Bremer, 1976; Humphries, 1981; Funk, 1982; Goldblatt, 1985). As discussed in the preceding pages, Hexa- glottis (excluding H. nana) is believed to be a natural (monophyletic) assemblage distin- guished by a number of specialized features, their polarity determined by outgrou by generally accepted trends in Iridaceae. The characters used for the cladistic analysis are pre- sented in Table 2, and most of them are discussed camnari an 1987] D > AC E Qo AM? yt SU VU 2 S ES "Pdl an A c < c S x e e MU. x 2 Cd SP 9? e? Qo? qv < S Ri SU ae ui a? pU um ah < e "i eU e e tuy am SUO M" WT X9 we es eo BB Ww wi RE 5. Cladogram of Hexaglottis and its rela- Homeriinae indicating the possible (par allelisms) are — by double lines and a prob- able reversal by a GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS 551 in more detail in the pages dealing with mor- phology and cytology of Hexaglottis. The basic structure of the Moraea flower and the reduction and SESE 2 Homeria and some species of a hav n discussed at length else- oye (Goldblatt 1980, 1986). Reasons for con- sidering x — e basic chromosome number in Moraea and "s allies have been presented in two studies (Goldblatt, 1971b, 1976a) dealing primarily with chromosome cytology. The immediate sister group of Hexaglottis is probably Homeria, and the genera share a series of derived and reduced floral features as well as the same chromosome number and karyot genera that stand out phenotypically in a variety of unusual features. A study in progress suggests that each of these segregates 1s related to a species or section of Moraea. Homeria and Hexaglottis together are probably related to M. flexuosa of the monotypic section Flexuosa, in turn most likely derived from species at present placed in section Moraea. The sole synapomorphy that LE 2. Characters used in the cladogram (Fig. 5), the derived (apomorphic) state listed first, followed by the presumed ancestral (plesiomorphic) condition. Corm tunics composed of hard, wiry, blackish fibers forming an open reticulum —tunics composed of straw- colored fibers forming a fine reticu um. . Karyotype comprising only acrocentric odo mos Gates comprising submetacentric pairs as weil id. dtu and nectar guides present on inner and outer tepals— nectaries and nectar guides present on outer e branches ned iren arn above the stigma lobe. d 2. Basic chromosome number x = 6— number x 3 as acrocentri 4. Style branches ` narrow and not petaloid—style branches broad and petalo 5. tepals o 6. Filaments united entirely (or free near the apex)— ae united in the lower half. 7. Flowers shades of blue to v UN shades of yello 8. Stem flexuose—stem more mc 9. Tepal claws longer than the limbs- tepal p about as long as or shorter than the limbs. 10. Style branches divi to 11. Armsofthe style branches 1. apically stigmatic 12. Lateral rhipidia sessile and eeding the subtending 13. Hypanthium tube present, E least 1 mm a tepals free from the base. 14. Hypanthium tube at least long—tub m long. 15. Ovary enclosed in the iria es—ova sert om the spath 16. Capsule narrowly obovoid to clavate-truncate — c psule ellipsoid l psule more or less cylindric-trigo patra elli Capsule iro i ellipsoid and iari sed in t u ed— — not bea Plants polyploid (2 Nee 6230.99.54 fo "a a psoid. he s nee uis ellipsoid and exserted. = 24)— ae diploid ei = bns . Leaves spreading and undulate — leaves ascending to erect and more or less straight. 21. Basic c some number x = 7— basi 22. Plants self-compatible and Ben DH ai tneonieedbln 23. Seeds angular-fusiform — seeds broadly angular. 24 25. Leaf margins undulate to lightly crisped —leaf margins straight. 552 unites these species is the corm tunic which con- sists of coarse, dark brown to black, netted fibers. Within the group with coarse black corm tunics, Hexaglottis, Homeria, Moraea section Flexuosa, and Moraea section Polyanthes appear to form a monophyletic alliance that shares the derived basic chromosome number of x = 6 and a karyo- type of predominantly to exclusively acrocentric chromosomes. Section Polyanthes has blue flow- guides on both inner and outer tepals and have style branches reduced from the basic petaloid condition. The species of section Moraea be- longing to this alliance include M. namaqua- montana, M. serpentina, M. tortilis, and their close allies, all except M. namaquamontana being united by having included ovaries and capsules. The detailed relationships of the species in sec- tion Moraea are not dealt with further and will be the subject of a future study. Hexaglottis itself stands out in having spe- cialized style branches and having a possible re- versal in the filaments being free in the upper half (see discussion under Flower in the section dealing with Morphology). Hexaglottis nama- quana stands out in the genus as unspecialized and taxonomically isolated. It is probably close species. It is the sister species to the other species of the genus, which all have sessile lateral rhi- pidia, an important synapomorphy in Hexa- glottis. The included ovary and hypanthium tube are synapomorphies uniting H. brevifolia and H. virgata, the latter distinguished by its longer tube and particularly short pedicel. Hexaglottis vir- gata subsp. karooica has unusually large flowers (probably a specialized condition but not reflect- [o m 4. Hexaglottis riparia and H. longifolia form another species pair, linked by the derived capsule shape. Hexaglottis lon- gifolia stands out here in having large capsules, autogamous reproduction, and in being poly- ploid. The two subspecies of H. lewisiae appar- ently share no synapomorphy, or at least none that I have been able to identify. However, they are too similar morphologically to be regarded as separate species. Further study may throw more light on their relationship and will perhaps in- dicate the presence of specialized features linking them ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 The parallelism shared by Hexaglottis lewisiae subsp. /ewisiae and the H. virgata-H. brevituba clade, narrow fusiform seeds (character 23), sug- gests a possible alternative phylogeny with these taxa forming a single clade. This is an attractive hypothesis, suggesting as it does the derivation of the species with a perianth tube and cylindric, included ovary (i.e., H. virgata and H. brevituba) from an ancestor like subsp. /ewisiae, which at least has an unusually narrow ovary. The sepa- ration of H. lewisiae subsp. secunda from subsp. lewisiae that would result from such an inter- pretation is difficult to accept given their other- wise similar morphology. One significant conclusion that is evident from the cladistic analysis is that Moraea as presently defined is shown to be paraphyletic, consisting of a number of discrete lineages, some of which are treated as distinct genera. The analysis of the relationships of the main species groups in the whole alliance is in progress and there will likely be some changes to the taxonomy of Moraea and its segregates. Such changes may include Hex- aglottis but it seems preferable, nevertheless, to publish the revision according to the present tax- onomy. The species, H. nana, is not reassigned to any genus, as this would be premature given the prevailing uncertainty about the relation- ships of species in Moraea itself. SYSTEMATIC TREATMENT Hexaglottis Ventenat, Decades Generum No- vorum 6, no. 3. 1808. TYPE SPECIES: H. lon- gifolia (Jacq.) Salisb., lectotype, designated by Lewis (1959: 219-222) Important references: Baker, Handbook Irid. 75- 76. 1892 et Flora Cap. 6: 31-32. 1896; Lewis, Flora Cape Peninsula 225. 1950; J. S. African Bot. 25: 215-230. 1959 ? — Herbert, Edwards Bot. Reg. 30: Misc. 89 . TYPE SPECIES: P. flava Herb. [The identity of E flava, treated by Lewis (1959) as conspecific with H. virgata, is uncertain. N known. The protologue seems to ma ae and snae leafed species of Homeria sec- Hom and Conanthera equally we ell. Plantia is iir congeneric wit Hexaglottis.] Plants variable in size, 12-120 cm high. Corm globose, + symmetric, 10-20 mm diam. or larger if surrounded by accumulated tunic layers, the tunics consisting of dark brown to blackish layers of thick vertical ribs connected by fine cross- 1987] fibers, the older layers increasingly fibrous and dissected. Leaves usually 2-3, occasionally only , linear, usually ascending and longer than the stems, often trailing distally, or + prostrate, flat or channeled, usually inserted towards stem base or at some distance above the ground, the mar- gins plane or undulate. Stem erect, straight to slightly flexuose, branched, with only one main axis or with 2-6 diverging secondary axes; lateral o amaqu partly enclosed by the subtending stem bract, this usually as long or longer than spathes. Rhipidia stalked (H. namaquana) or sessile except the ter- minal, 2—several-flowered; spathes herbaceous, or partly to entirely pi and pale at flowering time, attenuate, except in H. namaquana, the outer often d E the sheathing stem bracts and membranous below, !^—/ as long as the inner (subequal in e lateral rhipidia of H. longifolia); in sessile rhipidia the upper part of inner visible above bracts except in H. longifolia. Flowers stel- late, upright or facing sidewards, yellow, scented or not, the nectar guides deeper yellow and usu- ally surrounded by small dark spots, larger on the outer tepals, located near the base of the limbs, the tepals free or united below as a closed tube; perianth tube (when present) cylindric, 1-7 mm long, narrow, usually curving slightly outward, GOLDBLATT —SOUTHERN AFRICAN HEXAGLOTTIS 553 partly enclosed in the spathes; tepals with short erect claws 1-2 mm long, forming a narrow cup enclosing base of filaments or filament column; the outer slightly larger than the inner and with a small nectary on the claw; limbs extended hor- izontally, the outer + ovate to oblong, the inner oblong to cuneate. Filaments 4—6 mm long, mon- adelphous, united for 1-2(-3) mm, weakly di- verging above; anthers 3-9 mm long, linear, ini- tially erect, curling inwards and partly collapsing after anthesis. indric or wider the spathes; style dividing at apex of the united part of the filaments into 3 short branches, each divided almost to the base into 2 filiform (mi- croscopically grooved) arms, extending outwards on either side of the subtending filaments, ciliate and stigmatic only at the apex. Capsule ellipsoid, obovate to clavate, or + cylindric, exserted or included in the spathes, usually only 1 per inflo- rescence in H. virgata; seeds angular, sometimes Ó ud hur basa 2n = 12, 14, Distribution. Winter rainfall parts of south- ern Africa, from Port Elizabeth in the east, to the Cape Peninsula and north throughout Nama- qualand, also locally on the Roggeveld Escarp- ment in the western Karoo. KEY TO HEXAGLOTTIS AND OTHER AFRICAN IRIDACEAE WITH A HEXAGLOTTIS-TYPE FLOWER la. Rhipidia 2-several, arranged in a fascicle, each on a short stalk; outer inflorescence spathes usuall ‘Hi not entirely sheathing, but with a diverging apex exaglottis nana — outer inflorescence spathes entirely sheathing. 2a. Lateral rhipidia stalked. 3a. Flowers blue-violet; style arms 2.5- - mm lon ie mm wide, spreading on the ground nd twisted, the margins undulate and often cri a. 1. Hin 3b. Flowers yellow; style arms about 4 b. Rhipidia many, not fasciculate but in spicate or racemose arrangement, each either sessile or stalked; Moraea bir d dn leaves comparatively broad and short, to 11 namaquana N c 4a. Tepals free to base; point, plants of open a wn c habitats, streamsides, an 6a. ee apsules 12-16(- shorter than outer and concealed by the sheathing bract leaves .. nd ca. 4 mm wide; inner spathe of the lateral rhipidia H. ri ually 6b. Capsules 6—10(-12) mm long an sually longer than outer and not concealed ry Capsules obovoid to clavate, truncate above; 4-8 mm at the widest point; plants of moist . Lateral rhipidia sessile, ed arranged on the main axes. sule partly to well pom from the spathes 5a. Capsules psa dca ellipsoid to | cylindric- trigonous; rarely more than 3 mm at the pus 2. H. lewisiae 23) mm long and 6-8 mm wide; inner spathe of lateral fpa H. longifolia riparia 4b. Tepals united into a tube below the claws; ovary and capsule enclosed within the spathes. 7a. Perianth tube 1-2 mm long; upper part o plants of northern Namaqualand and the Richtersveld 7b. Perianth tube (3-)4-9 mm long; inh entirely included; plants of the northwest, p and 6. southern Cape and western Karo of the ovary often emerging from the spathes; 5. H. brevituba H. virgata 554 — ` Hexaglottis namaquana Goldbl., sp. nov. TYPE: South Africa. Cape: Namaqualand, top of Spektakel Pass, stony clay soil among patches of quartzite, Goldblatt 3059 (holo- type, MO; isotypes, K, NBG). Figure 6. Plantae 15-30 cm altae, foliis 2-3 prostratis undu- latis, marginibus undulatis vel crispis, omnibus rhi- pidiis pedunculatis ex bracteis vaginantibus caulium exsertis, tepalis liberis, ca. 2 cm longis, ovario o 4-5 mm a ellipsoideis ca. 8-10 mm longis. Plants 15-30 cm high. Corm 1-1.5 cm diam., the tunics fibrous, dark brown to black, extend- ing above into short stiff bristles. Leaves 2-3, all + basal, 8-15 cm long, to 11 mm wide, + prostrate, irregularly undulate or twisted, the nches (o bloom) stalked, the branches subtended by dry, sheathing bracts 17-21 mm long. Rhipidia stalked, exserted from the subtending bracts; abov e, acute, the i inner 2.5-3 cm long, the outer about half as long. Flowers yellow, stellate with free, spreading tepals; tepals about 2 cm long with claws about 2 mm long, the outer tepal limbs to 5 mm wide, the inner narrower. Filaments united only at very base (seemingly free), 3—4 mm long; anthers about 3 mm long, straight and suberect before dehis- cence. Ovary 4-5 mm long, usually just exserted from spathes, the style arms spreading, about 4 mm long. Capsules narrowly oblong-ellipsoid, 8— 10 mm long; seeds Ric about 1 mm diam. Chromosome number 2 Flowering time. Lais Sepieribar to October; flowers opening in the mid afternoon, after 3:00 P.M. and fading near sunset. Distribution. Hexaglottis namaquana is known only from the eastern slopes of the Spek- wards the top of Spektakel Pass, in hard, stony, clay soil, sometimes covered with white quartzite pebbles of the Nama System. Outcrops of Nama shales and quartzites are rare in Namaqualand, where granites and granitic sands are the rule. Hexaglottis namaquana may be found in other parts of Namaqualand where there are similar outcrops of the Nama System. The species is sympatric with H. /ewisiae, which on Spektakel Pass is a tall slender, narrow-leafed plant, bloom- ing very late in the day. Its flowers open at about 5:30 P.M. and last about three hours. The flowers ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 of H. namaquana open at about 3:00 P.M. and begin to fade at 5:30. They are further isolated reproductively by a shift in flowering season. Hexaglottis namaquana blooms from late Sep- tember into mid October, whereas H. /ewisiae in this area blooms from mid October to late No- vember. Diagnosis and relationships. Hexaglottis namaquana, discovered only in 1974, is a re- markable member of the genus. It has the small ellow flowers that characterize Hexaglottis, but are borne on long s rather than being sessile. The former feature must be regarded as primitive for the genus and separates H. nama- quana from the other species. Members of this species are fairly short and are unusual also in having the leaves strongly undulate to crisped. The capsule is unspecialized in being well exsert- ed from the spathes and in being ellipsoid. In other species of Hexaglottis the capsule is either elongated and sometimes linear or is shortly stalked and enclosed within the spathes Fendi s specimens s. SOUTH AFRICA. —29.17 (Springbok): Spektakel pec, west of Springbok (DA), Goldblatt 5172 (MO, NBG); rocky eastern the top of Spektakel Pass, Goldblatt 6672 (MO). 2. Hexagon lewisiae Goldbl., J. S. African Bot. : 234. 1971. T the lectotype of the invalid M. /lexuosa)]. Figure 7. Moraea flexuosa L. f., Suppl. Pl. 100. 1782; Ker, Bot. Mag. 19: tab. 695. 1803, nom. illeg. superf. pro Ixia longifolia Jacq. T flexuosum (L. 1825, nom. illeg. 27, nom. illeg. bas. illeg. Hexa- glottis flexuosa (L. f.) Sweet, Hort. Brit. ed. 2: 498. : s, J. S. Afr. Bot. 25: 223. 1959 et FI. Cape mih 225. 1950, nom. illeg. bas. illeg. he as for Ixia longifolia (= Hexaglottis longi- folia ia )Salisb. sensu Baker, Flora Cas, 6: 32. 1896, pro parte (excluding the type of H. longifolia). Homeria spicata (Ker) Sweet sensu Klatt, Linnaea 34: H J 4 25 í Y we ]. s (1866) concept of H. spicata included Hexaglottis virgata, H. longifolia, and H. lewisiae. 1987] GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS 555 E 6. Morphology and distribution of Hexaglottis namaquana. Habit x0.5; flower full size; side and x2. FIGU top view w of the stamens and style branches Plants variable in size, (12-)20-60 cm high. Corm 15-20 mm diam., symmetric, the tunics (pale-)dark brown, fibrous, occasionally pro- duced upwards into a neck. Leaves 1—3(—-4), in- serted towards the base (sometimes shortly above the ground), ascending, linear, channeled, the margins sometimes inrolled or rarely undulate and the leaves rather short, normally much ex- ceeding the stem and trailing above. Stern usually bearing 1—3 secondary branches near the base, often flexuose, the lateral rhipidia sessile, usually overlapping the rhipidium above, subtended by a sheathing stem bract concealing at least the lower part of the spathes, and usually about two- thirds their length. Rhipidia sessile except the terminal; spathes herbaceous, dry above, (2.5-) 3-4.5 cm long, attenuate, the inner slightly longer than the outer. F/ower golden yellow with a strong sweet scent, stellate with free tepals, the outer often feathered brownish on the reverse, the nec- tar guides deeper yellow, usually surrounded by several small dark greenish spots; tepals 19-24 mm x 7-10 mm (subsp. /ewisiae), 24-30 mm x 10-13 mm (subsp. secunda), the claws 1.5-2 556 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 ZZ: UP Sg Qa : p ny ete Hep ur tem L °. = M 1500 7 ALL NM | l Yi PFA , AD ea N Tad as edi uw 4. FIGURE 7. Morphology of Hexaglottis lewisiae subsp. lewisiae and the distribution of both subspecies of H. lewisiae. Habit, corm, and fruiting branch x0.5; flower full size; seeds much enlarged. 1987] mm long, the limbs horizontal, the outer ovate, the inner more or less cuneate, about as long to 2 mm shorter than the outer. Filaments 4-6 mm long, united for 1-1.5 mm; anthers 5-7 mm long before e Ovary (5.5-)6-12 mm long, usually partly to entirely exserted at flowering, rarely eel included (Barker 2558); style about ong, the style arms spreading and 6 mm long. Capsule triangular in section and nar- rowly cylindric, 11—16(-20) mm long (subsp. lewisiae) or terete and ellipsoid, 8-13 (-15) mm long (subsp. secunda), then often shortly beaked, dehiscing in the upper third to half only; seeds angular, sometimes narrowly so and ta- pering at both ends, 1.2-2 mm long, 0.7-1 mm at the widest, winged in northern populations. Chromosome number 12. Flowering time. (September-)October-No- vember; flowers opening between 3:00 and 4:00 P.M. in the south and beginning to fade towards 7:00 P.M., only opening towards sunset in the Namaqualand populations and fading at about Distribution and biology. Hexaglottis lewi- siae is widespread in the southern African winter rainfall area (Fig. 7). It extends from Springbok in the north through the southwestern Cape as far east as the Humansdorp district. It is com- mon only in the western Cape and has a scattered distribution to the north of Piketberg and east of Bredasdorp. It is found in dry and exposed sites, often on stony ground, and occurs on sandy and clay soils. Like most species of Hexaglottis, H. lewisiae is pollinated by bees, but pollination has been studied in most detail in this species, and my observations are summarized here. The rela- tively appear to be very attractive to small bees. When the flowers open in mid afternoon, bees rapidly appear around the plants and begin to gather pollen. They visit the same flowers several times, collecting pollen as soon as it is exposed in the anther sacs, which dehisce slowly from the apex. Pollen is the primary reward and only occasion- ally is some nectar also taken. The small quantity of nectar produced seems to be of minor interest to the pollinators. Fruit and seed production is very successful in H. /ewisiae, and plants develop several full capsules from each of the many in- florescences. Diagnosis and relationships. Hexaglottis lewisiae is a diploid and self-incompatible. Its GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS 337 yellow flowers are almost identical to those of all other species of Hexaglottis, except that unlike H. virgata and H. brevituba, the tepals are free to the base. It shares with all but H. namaquana the similar vegetative feature of sessile lateral rhipidia. It is most easily confused with H. ri- or clavate capsules, thus always broadest in the upper third and markedly flat-topped, unlike the cylindric or ellipsoid, and sometimes beaked, capsules of H. lewisiae. Hexaglottis riparia and H. longifolia grow in moist situations and, pos- sibly as a consequence, have more and longer leaves than is usual in H. /ewisiae. The combi- nation of init three or = leaves; obovate: cla- vate oist habit smaller RR makes it ol that H. riparia and H. longifolia will be confused with H. /ew- isiae. History. Hexaglottis lewisiae has been known since the younger Linnaeus (1782) described the species as Moraea flexuosa based on material collected by Carl Peter Thunberg a few years earlier. The epithet is regarded today as super- fluous and illegitimate since Linnaeus cited as a synonym /xia longifolia Jacq., now H. longifolia. The combination * ee flexuosa was made by Sweet in 1830, and the species, usually in- cluding H. Tonerli, was known by this name for several years. The later nineteenth century botanists generally did not recognize H. flexuosa. F. W. Klatt included it, together with H. virgata and H. longifolia, in Homeria spicata, this a syn- onym of Homeria elegans (Goldblatt, 1981). Lat- er, Klatt (1882) recognized Hexaglottis, includ- ing H. virgata and H. longifolia (presumably but not explicitly including H. lewisiae). J. G. Baker (1896) included H. flexuosa in H. longifolia, and the distinction between the two species was only reestablished by Lewis in 1950. The new name H. lewisiae was proposed in 1971 by the present author for the species that until this time was known by the illegitimate name H. flexuosa. Variation. There appear to be two major forms of Hexaglottis lewisiae, the southern and typical, treated here as subsp. /ewisiae, which extends from the western Cape coast eastwards through the southern Cape to Humansdorp. It has medium-sized flowers with tepals 19-24 mm long and distinctive long slender capsules 1 1-20 mm long that tend to dehisce only in the upper part. The seeds are also comparatively small as a result of the need to be accommodated in the 558 narrow locules. The karyotype in this form ap- pears to be uniform and consists of four strongly acrocentric chromosome pairs and two acro- to submetacentric pairs. Populations to the north of the Olifants River mountains, from Clanwilliam north to Spring- bok, treated as subsp. secunda, comprise plants with larger flowers, the tepals 24-30 mm long, somewhat shorter ellipsoid capsules 8-13(-15) mm long, and larger seeds. The chromosome cy- tology of this series of populations is not as well known, but two Namaqualand populations ex- amined have karyotypes with a large metacentric chromosome pair. This northern form is mor- phologically variable. Namaqualand plants have a single leaf in the wild, but two leaves in cul- tivation, and slightly smaller flowers than those from the northwest Cape. The extensive popu- lations from the western Karoo near Nieuwoudt- ville have capsules with a beaklike apex, a feature n Clanwilliam and Y nrhyn and karyotypes din been determined for one northwest Cape population. The flowers of the Namaqualand populations have a different phe- nology, opening between 5:30 and 6:00 P.M., whereas all other forms of H. /ewisiae open be- tween 3:00 and 4:00 P.M. and fade at about 6:00 P.M KEY TO THE SUBSPECIES OF HEXAGLOTTIS LEWISIAE la. Capsules ellipsoid, 8-13(-15) mm long, often distinctly beaked; outer tepals 24-30 mm long, flowers usually secund ........ lb. pA cylindric or nearly m long, not beaked; outer tepals 19-24 mm pen flowers usually upright aaa. 2B. subsp. /ewisiae 2A. subsp. secunda Goldbl., subsp. nov. TYPE: South Africa. Cape: stony east-facing slopes near the top of Spektakel Pass, west of Springbok, Goldblatt 6673 (holotype, PRE; isotypes, K, MO, NBG, S, US, WAG) Planta 30-60 cm alta, floribus usitate hus te- end 24-30 mm longis 1 rio 5.5-9 m longis, APS ellipsoideis 8- 1315) m mm longis ipia rostratis 3 Plants 30-60 cm high. Flowers usually secund; tepals 24-30 mm long, 10-13 mm wide. Ovary 5.5-9 mm long, usually at least partly exserted at flowering. Capsule terete and ellipsoid, 8-13 (-15) mm long, dehiscing for at least half its ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 length; seeds angular, narrowly winged on the ridges, about 2 mm long, ca. | mm at the widest. Distri bution. Subspecies secunda has a scat- william north through the Olifants River Valley to Vanrhynsdorp, to the Karoo north and east of Nieuwoudtville, and into Namaqualand, where it has been recorded west of Garies, in the Ka- mieskroon district, and on the Spektakelberg, west of Springbok. Plants grow in rocky ground, either in clay soil, as on the Spektakelberg and in the Karoo, or in sand. Additional specimens examined. SOUTH AFRICA. Se 17 (Springbok): stony east-facing pus near p of Spektakel Pass, west of Springbok (DA), Goldratt 6613 (MO). 30.17 (Hondeklipbaai): Klip Vlei, between Kamies- A 9 of Skuinskraal farm, Hondeklipbaai road (DB), van Berkel 453 (M B 31.18 (Vanrhynsdorp): EET byi (K), Barker s665 (NBG), Snijman s G); flats below the erg, 12 km south of in veri Goldblatt 6992 (MO, PRE); slopes of the Olifants River Bridge, south of Klawer, rocky sand- stone soil (DC), M ae (K, MO, NBG, PRE). 31.19 (Calvinia): Loeriesfontein road, ca. 2 mile north of the Waterfall (AC ), Goldblatt 1 08 (J); karroid hills anges of the Klip Koppies, Nieuwoudtville, ndkraal sandveld (D farm, Nieuwoudtville, Goldblatt y; Nieuwoudtville rocks on top of kopies, Galpin 11137 (K 32.18 (Clanwi is north of Clanwilliam, rocky sandstone soil (BB), Goldblatt 6990 (MO, PRE, , US). 2B. Subsp. lewisiae. Plants (12-)20—-60 cm high. Flowers bur upright; tepals 19-24 mm long, 7-10 mm wide. Ovary 8-10 mm long, usually partly to nes exserted at flowering, rarely entirely included. Capsule terete to triangular in section, narrowly cylindric, 11-15(-20) mm long, dehiscing in the upper third to half only; seeds angular-ellipsoid, tapering at both ends, 1.2-1.4 mm long, ca. 0.7 mm at the widest. Distribution. Subspecies /ewisiae extends from the Cape west coast east through the south- ern Cape to Humansdorp. It appears to be com- mon only in the west and records are very scat- tered east of Bredasdorp. Figure 7. Specimens examined. SOUTH AFRICA. CAPE-32.1 (Clanwilliam): NE edge of Verlorenvlei (AD-BC), m 1987] lans 7803 Do De Hoek, Piketberg (DD), Barker 2558 (NBG 33.18 (Ca own): Mamre hills (AD), Compton 9828 (NBG); d (BB), Loubser 466 (NBG); ca. 10 km north of Malmesbury (BC), Goldblatt 6173 (MO, S, US); mountains around Cape Town vy Ecklon & Zeyher s.n. (84) (S); foot of Lions Head, Pappe s.n. (SAM 70674); Oudekraal, Cape Peninsula, Goldblatt 163 (J); hang Lewis 97 1 (SAM); Camps Bay, Moss 13403 (BM); Table Mountain (CD), Ecklon 536 (BM, K, MO, PRE); Wynberg Hill, Ee 10198 (MO, US); Kirstenbosch, Lewis 672 (N SAM); Kirstenbosch, slopes near the herbarium, Lei 5104 (MO); Kir- stenbosch, near kd npud offices, Goldblatt 6634 (BR, M AG); below Pearson House, Kir- stenbosch, i. 6634 (BR, MO, PRE, WAG); near Bishopscourt, Salter 9002 (BOL); behind Groot Schuur, Wolley Dod 360 (BM, BOL, K); Rosebank, H. Bolus 3801 (BOL, K); Tygerberg Nature Reserve (DC), Loub- ser 3059 (MO); Langverwacht, above Kuils River, Oliver 4803 (K, MO, PRE, STE); Stellenbosch Flats (DD), Garside 66 (K); between Klapmuts and Paarl, Acocks 3677 (S); Berg 2 near Paarl, Drége s.n. in 1840 (K, S), Barker 8797 (N 33.19 ocu. Gydo Pass (AB), Wall 705 (S); orcester, beim Wasserfall (AC), Ecklon & Zeyher (LD, MO); near Tulbagh, Leighton 1317 (BOL); Tulbagh plains, Marloth 9575 (PRE); Tulbagh ). 21 (Ladis a 5 mi. west of Ladismith (AC), rat 3030 (N E ot Ruigtevlei, eb C), Fourcade 1525 (BOL). 33.24 T aaa Essenbosch hills (CD), Four- cn 4420 4.18 i i UE Simons Bay (AB), Wright 269 u ene Bay, Wall s.n. (S); s Barker 2728 (BOL, NBG); Bergvliet Farm, Purcell 124 (SAM). 34. 19 (Caledon) Elgin (AA), Dd 1164 iei Dwarsberg-Somerset Sneeukop, Stokoe s.n. (SAM 55728); Genadendal (AB), Prior s.n. (K); between Houw Hoek and Kleinmond (AC), Werdemann & Oberdiec 678 (B, K, PRE); Hemel en Aarde (AD), Gillett 90 (STE); 8 mi. from Stanford on the road to Elim, Gillett 4506 (BOL, K); SEV: Vogelklip, Sapa S. Williams 873 (C, M AG); near Napier (BD), Lei- poldt 3551 (BOL); mb Baardscheerdersbos and Elim (DA), Goldblatt 7107 (MO, PRE). 34.22 (Mossel Bay): ps the river at Great Brak River (AA), Young s.n. (BOL 5541); between (ado and Great Brak River (?AB), y eun 6151 (K). 34.24 (Humansdorp): Oudebosch flats (AA), Four- cade 960 (BO L). Without precise locality: Cape of Good Hope (CBS), near Swart R., Swartz"); rock crevices above forest plantation, Clan- william, Galpin s.n. Introduced: Réunion, Trou aux Cerfs, Vaughan 3255 (SAM), Lorence 15764 (K). 3. Hexaglottis riparia Goldbl., sp. nov. TYPE: South Africa. Cape: along the Olifants River GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS 559 at Citrusdal campsite, after fire, Goldblatt 6706 (holotype, NBG; isotypes, K, MO, PRE, STE). Figure 8 sa aa ee aor var. angustifolia Lewis, J. S. Af- 2. 1959, pro parte (excluding the type ES var. annal e placed in H. longi- n the present pap (Plantia flava Herb., Edwards Bot. R eg. 30: misc. 89. 1844 1. possible synonym. See discussion below der tory.) Planta 45-90 cm alta, foliis (2-)3 linearibus rhipidiis terius sessilibus, bracteis caulis imbricatis lon- gioribus m internodis, spathis onus ide in dudes caulis T tepalis liberis 16-21 mm longis 6-8 mm latis, ovario 5-7 mm longo exserto, capsulis ue do 6-10C1 2) mm longis. Plants 45-90 cm high. Corm 13-20 mm diam., the tunics of fine, light brown fibers. Leaves long as the stem, the upper decreasing in length and width. Stem straight, simple or 1-3-branched from the lower nodes, the lateral rhipidia sessile at each node, subtended by a sheathing stem qas ftan these as ines or sontes hài longer than the inter- node and overlapping the next bract. Rhipidia sessile, except the terminal; spathes herbaceous, attenuate, dry apically, 3.3-3.8 cm long, about as long as the subtending stem bract, the outer shorter than the inner, often hidden. Flowers stel- late with free tepals, deep yellow, strongly scent- ed; tepals 16-21 mm long, 6-8 mm wide, with claws about 2 mm long, the limbs spreading, the inner slightly shorter than but as wide as the outer. Filaments 4-6 mm long, united for 1.5- 2.5 mm; anthers 5-6 mm long. Ovary 5-7 mm long, exserted; style arms 5-6 mm long, extended horizontally. Capsules obovoid-clavate, some- what truncate, 6—10(-12) mm long, about 4 mm wide; seeds angular, 1-1.5 mm long, 1 mm at the widest diam. Chromosome number 2n = 12. Flowering time. October-November; flow- ers open at about 5:00 P.M. and begin to fade after 7:30 P.M. Distribution. Hexaglottis riparia has a lim- south and Clanwilliam in the north (Fig. 9). It appears to be restricted to streambanks and pos- sibly edges of marshes. Such areas are usn) overgrown with tall vegetation, and H. ripar accordingly blooms only after fires or heavy graz- 560 f£ II vias Morphol Habit flowering, and trating branches x0. 5; Fendi flower and eds full siz ing when the habitat has been opened up con- siderably. Diagnosis and relationships. Hexaglottis ri- paria has flowers typical of the genus. It is dis- tinctive largely in its small obovoid to clavate fruits and in its slender, often willowy stems. The capsules are 6-10 mm long, or occasionally in robust plants up to 12 mm. Hexaglottis longifolia has similarly shaped capsules but they are much larger, usually 16-23 mm long. The similarity in ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 capsule shape prompted Lewis (1959) to include the only collection of H. riparia known to her in H. longifolia var. angustifolia. The type of the latter has unusually narrow leaves but in other respects the type corresponds well to typical H. longifolia. This species differs further from H. riparia in having pale yellow, unscented flowers and in being self-compatible, autogamous, and tetraploid with 2n = 24. The flowers of H. riparia are deep yellow and scented, and, like other members ofthe genus, it is self-incompatible and diploid with 2n = 12. The similar capsule shape in H. riparia and H. longifolia, a derived feature, probably indicates that they are closely related. History. This species was apparently collected first by Ecklon & Zeyher in the Tulbagh district in the 1820s, and rarely since then. It was initially assigned to Hexaglottis longifolia and subse- quently cited by Lewis (1959) under H. longifolia var. angustifolia. There is a possibility that the species described as Plantia flava may be the same as H. riparia. Baker (1896) and Lewis (1959) treated the monotypic Plantia as congeneric with Hexaglottis. The type is, however, unknown, and the description is ambiguous and could apply as well to some species of Homeria as to Hexa- glottis. Plantia flava was grown and flowered in England from corms collected by George Synnot, who lived in the Clanwilliam district from 1821 to 1825 (Gunn & Codd, 1981). If Plantia flava is a species of Hexaglottis, it is probably con- specific with H. riparia, the only species of Hexa- glottis in the northwest Cape with capsules that correspond with Herbert’s description of the cap- sules as obovate. Specimens examined. SOUTH AFRICA. CAPE-32.18 (Clanwilliam): river banks, Clanwilliam (BB), Galpin 11483 (BM, K, PRE, UPS); Clanwilliam, Leipoldt 376 (S 32.19 (Wuppertal): along the Olifants river at Cit- rusdal campsite (CA), iege 6555 (K, MO, NBG), P (K, MO, NBG, PRE, STE). 3.19 (Worcester): cu Tulbaghskloof, etc., ; & Zeyher Irid. 30 (77.9) (K, MO). 4. Hexaglottis longifolia (Jacq.) Salisb., Trans. Hort. Soc. 1: 313. 1812; Baker, Flora Cap. 6: 32. 1896, in part excl. H. lewisiae (as H. flexuosa); Lewis, Flora Cape Peninsula 225. 1950 et J. S. African Bot. 25: 223-225. 1959. Ixia e. Jacq., Hort. Vindob. 3: 47 & tab. 90. 1776. Moraea longifolia (Jacq.) Pers., Syn. Pl. 1: 49. 1805. Moraea flexuosa L. f., Suppl. Pl. 100. 1782, nom. illeg. superf. pro 1987] Ixia longifolia Jacq., applied to H. lewisiae. Sisyrinchium flexuosum (L. f.) Spreng., Syst. Veg. 1: 167. 1825, nom. illeg. bas. illeg. Homeria flexuosa (L. f.) Sweet, Hort. Brit. 95. 223. 1959, nom. illeg. bas. illeg., applied to H. lewisiae. TYPE: South Africa. Cape: illus- tration in Hort. Vindob. 3: tab. 90. Figure 9. j ge pone wht es var. angustifolia Lewis, J. S. Af- 9. TYPE: South Africa. Cape: Du pim Rs ‘Pillans 8436 (BOL, holotype), pro parte (including only the type, other speci- mens here treated as H. riparia). Plants 60—150(—200) cm high. Corm 15-20 mm diam., the tunics of fine, light brown fibers. Leaves 3-4(—5), linear, ascending, the lower longest, 50- 100 cm long, the upper decreasing in size, 6-12 (-20) mm wide, flat or channeled, usually bent and trailing distally. Stem straight, simple or 1— 3-branched from the lower nodes; lateral inflo- rescences sessile at each node, subtended by a sheathing stem bract often entirely concealing the spathes, as long, longer, or shorter than the internode. Rhipidia sessile, except the terminal; spathes herbaceous, attenuate, dry at apex, 3.5- 5.5(-7.5) cm long, the inner about as long as the subtending stem bract, the outer shorter, often hidden. Flowers pale yellow, odorless or with a very faint odor, stellate, with free tepals, the nec- tar guides usually only on the outer tepals, in- conspicuous; tepals with claws about 2 mm long, the outer with limbs 19-27 mm long, 8-11 mm wide, the inner 16-23 mm long, 6-9 mm wide. Filaments 4-6 mm long, united for 2-3 mm; anthers 5-6 mm long before anthesis. Ovary 8— 12 mm long, exserted, the style arms compara- tively short, 4-6 mm long, ascending, reaching only to the base of the anthers. Capsules well exserted, obovoid-clavate, truncate, 12-16(-23) mm long, 6-8 mm wide, dehiscing for /2— their length; seeds angular, 2-2.3 mm long and up to 1.5 mmatthe widest. Chromosome number 2n = 24. Flowering time. Mid October-November; flowers opening (2:00—)2:15-2:45 P.M. and be- ginning to fade ca. 6:00 P.M. Distribution and biology. Hexaglottis longi- folia is restricted to the Cape Peninsula and a few valleys in the mountains between Somerset West and Paarl. It grows along streams and in marshes or shady and damp sites (Fig. 9). GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS 561 It is the only polyploid species in the genus and further unusual in Hexaglottis in being self- compatible and normally autogamous. Although the flowers appear to the human eye similar to those of the bee-pollinated H. /ewisiae, H. lon- gifolia is seldom visited by insects. Plants of H. longifolia growing a few feet away from H. lew- isiae are ignored by bees gathering pollen from the latter species. Flowers of H. longifolia placed among those of H. lewisiae are consistently avoided by bees. Clearly to these insects, the odorless flowers of H. longifolia are very different to those of H. lewisiae, which are strongly scent- ed. Possibly there are significant differences in ultraviolet patterning as well. Diagnosis and history. The good illustration of Jacquin's Ixia longifolia accompanying the protologue and now regarded as the type of the species corresponds unmistakably to the tall, pale yellow-flowered Hexaglottis of damp shady sites on the Cape Peninsula. The painting shows clear- ly the characteristic pale yellow flowers, six rath- er short style branches, and the large, obovoid to clavate capsules. Despite this clear illustration, H. longifolia has often been associated with the very different H. lewisiae (as H. flexuosa) (Klatt, 1895; Baker, 1896), which has deep yellow, scented flowers, long style branches, and linear cylindric capsules. Lewis (1959) has explained in detail this historical confusion and has elabo- rated the several differences between these two quite distinct species. No further discussion seems necessary here. Lewis (1959) recognized two varieties of Hex- aglottis longifolia in her revision, the typical, and var. angustifolia, the latter distinguished by nar- rower, stiff, and strongly channeled leaves, short- er spathes, and smaller capsules. Her decision to treat this somewhat smaller form as a variety of H. longifolia was evidently based on similarity of general appearance, including the shape of the capsules, which, although smaller, are obovoid and truncate, and by the habitat: moist places along streams and rivers. The type, from du Toits Kloof, seems to me merely a slightly smaller specimen of H. longifolia and can be matched in size by some collections from the Cape Penin- sula. It does not appear to warrant taxonomic recognition. However, the other collection cited, Galpin 11483, from the Olifants River Valley, is a different species treated here as H. riparia. It has smaller capsules and the flowers, examined live from this area, are unlike those of H. /on- 562 ANNALS OF THE MISSOURI BOTANICAL GARDEN A H. riparia Zm mi Nau | E 9. Morphology of Hexaglottis longifolia and distribution of H. ol ^d H. riparia. Habit, eben and fruiting branches, and corm x0.5; flower + full size; seeds much enlarg gifolia. They are bright yellow and strongly scent- ed, in both characteristics corresponding better to H. lewisiae and in conflict with H. longifolia with its pale yellow, scentless flowers. In addi- tion, H. riparia is diploid, 2n = 12, and self- incompatible, in contrast to the tetraploid and autogamous H. longifolia. Specimens examined. SOUTH AFRICA. CAPE-33.18 (Cape Town): Liesbeek River, below Fernwood (CD), Salter 8973 (NBG), 8775 (NBG, SAM), 8999 (NBG); Kirstenbosch, Lewis 673 (NBG, PRE, SAM), Goldblatt 5934 (MO); Kirstenbosch, below Pearson House, Ma- lan 120 = “a Goldblatt 6635 (K, MO, PRE, WAG); low Fern , Salter 9377 (BM); above Rhodes Drive, Salter 9154 (BM); Table Mt., east base in damp soil, Pillans 10262 (BR, MO); ee Wer- dermann & Oberdieck 722 33.19 (Worcester): du ci ae (CA), Pillans 4836 ou ou Simonstown): Orange Kloof, swamp (AB), Wolle Dod 3479 (BM, BOL, K, PRE); shady roadside ing below Constantia Nek on the road to Groot Ca Goldblatt 6640 (MO, PRE, S). Without precise locality: Thunberg s.n. (S "Herb. Casstrom"). 5. Hexaglottis brevituba Goldbl., sp. nov. TYPE: South Africa. Cape: Richtersveld, Sabiesies, on the a to Cornelsberg, Viviers s.n. in 1983 (holotype, NBG; isotypes, K, MO, PRE). Figure 10. [VoL. 74 1987] Planta 40-55 cm alta, foliis 2-4, rhipidiis : apen exterioribus ex bracteis caulis “ra nalis [aD bonus tubo 1-2 mm longo, ovario 8-1 m longo supra ex bracteis exserto, pedicellis 4-10 mm mg a Plants 40-55 cm high. Corm 12-18 mm diam. or larger if surrounded by accumulated tunic lay- ers, the tunics of coarse, dark brown to blackish fibers. Leaves 2—4, linear, ascending, longer than the stems and trailing distally, seh ie with margins incurved, inserted towards stem base. Stem with 1 main axis or with 23 diverging secondary axes, straight or rarely slightly flex- uose; lateral rhipidia sessile, partly enclosed by the somewhat shorter subtending stem bracts. Rhipidia sessile except the terminal, 3-4-flow- ered; spathes herbaceous, or partly to entirely dry and pale at flowering time, attenuate, (20-)25- 30 mm long, exserted from the stem bracts, often membranous below, the outer half to two-thirds as long as the inner. Flowers stellate, upright or secund, pale yellow, evidently odorless; tepals united below into a short closed tube; perianth tube 1-2 mm long, exserted from the spathes; tepals 16-19 mm long, the claws about 1 mm long, the limbs extended horizontally, 5-8 mm wide, the outer larger, narrowly ovate, the inner smaller. Filaments 3—4 mm long, united for about 1 mm; anthers 7-8 mm long. Ovary 8-11 mm long, cylindric, often curved outwards, the upper 1 mm narrow and sterile, the apex often exserted from the spathes; pedicel 4-10 mm long, the style arms about 6 mm long. Capsule a ep seeds not known. Chromosome number 2n Flowering time. September to mM flow- ers opening mid afternoon and fading after 7:00 P.M. Distribution. Hexaglottis brevituba is known t 10). There is only a single record from the central part of Namaqualand, at Stinkfontein south of Garies, and other collections are from the Spring- bok area and north to the Richtersveld, where the type collection was made. Hexaglottis brevi- tuba is probably more common than the present record indicates. It is inconspicuous except when in bloom, and the flowers are open for only a few hours in the mid afternoon of a few weeks, usu- ally towards the end of spring. Diagnosis and relationships. Hexaglottis brevituba is clearly allied to the widespread H virgata, which has similar vegetative and floral GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS 563 morphology but is readily distinguished by its flowers with a well-developed perianth tube some 4—7 mm long and an entirely included ovary with a pedicel 4-6 mm long. In H. brevituba the peri- anth tube is only 1-2 mm long; the ovary is often apically exserted and the pedicel (4-)7-10 mm ong History. The first record of H lottis brevi- tuba was made by Rudolf Schlechter i in 1897 and the collection was identified only as Hexaglottis. In Lewis's revision of the genus she placed it in H. virgata var. lata, the type and only other col- lection being from the Biedouw Valley. A third collection, made by G. J. Lewis near Springbok, was assigned to H. lewisiae (as H. flexuosa) by Lewis, who did not notice the characteristic short perianth tube in the poorly preserved flowers. The perianth tube can, however, be seen in this collection when buds are examined carefully. The range of the species was substantially extended to the Stinkfontein Mountains in the Richters- veld by Mike Viviers in 1983. This, the only adequate collection of H. brevituba, has been se- lected as the type. Specimens examined. TH AFRICA. CAPE: 28.16 (Vioolsdrif): Richtersveld, Sabiesies, on the road to Cornelsberg (CA), Viviers s.n. (K, MO, NBG, PRE), 1337 (NBG). 29.17 (Springbok): north of Steinkopf, near Kosies road in kloof northeast of Rabas (BA), Goldblatt 5746 ~ 3 Am Nn a - xí 30. g): Stin Doorn R.) (CC), Schlechter 114876 (BOL). 6. Hexaglottis virgata (Jacq.) Sweet, Hort. Brit. ed. 2: 498. 1830; Baker, Flora Cap. 6: 32. 1896, pro parte; Lewis, Flora Cape Penin- sula 225. 1950 et J. S. African Bot. 25: 225. 1959. Moraea virgata Jacq., Ic. Pl. Rar. 2: tab. 228. 1791 et Coll. Bot. 3: 194. 1791. Ixia virgata (Jacq.) Willd., Sp. Pl., 1: 202. 1798. Homeria virgata (Jacq.) Sweet, Hort. Brit. ed. 1: 395. 1827. TYPE: South Africa. Cape: cultivated in Vienna, illustration in Jacq., Ic. Pl. Rar. 2: tab. 228. Figure 11. Plants variable in size, 12-85 cm high. Corm 13-18 mm diam., or larger if surrounded by ac- cumulated tunic layers, the tunics of coarse, dark brown to blackish fibers. Leaves — 2-3, oc- casionally only 1, rarely 4, linear, ascending, longer than the stems and trailing talis. chan- neled with margins curving inward, occasionally loosely coiled above, inserted towards the stem base. Stem with 1 main axis or with 1—3(-6) fairly 564 ANNALS OF THE MISSOURI BOTANICAL GARDEN "X Si [Vor. 74 A B H. brevituba H. virgata subsp. virgata > HER. subsp. karooica JN "r.i 4 D> | — N QUAE id As FIGURE 10. Morphology of Hexaglottis brevituba and the distribution of H. brevituba and H. virgata. Habit x 0.5; flower and corm full size; detail of stamens, ovary, and style branches x 1987] long, strongly diverging secondary branches, straight to slightly flexuose, the lateral rhipidia sessile, each partly to almost entirely enclosed by the subtending stem bract, this usually as long or longer than spathes. Rhipidia sessile except the terminal, 2—3-flowered; spathes herbaceous, or partly to entirely dry and pale at flowering time, attenuate, 22-30(-37) mm long, the outer concealed by a stem bract, often membranous below, !1⁄— as long as the inner, upper part of the inner extending above the bracts. Flowers stellate with the tepals united below into a closed tube, upright or secund, pale yellow, evidently odorless, the nectar guides deeper yellow; peri anth tube cy narrow, usually slightly curv- ing outward, (3-)4—6(-9) mm long, usually at least partly enclosed in the spathes; tepals 14—25 (subsp. virgata), 23-32 (subsp. karooica) mm long, with short claws 1-1.5 mm long, the limbs extended horizontally, 4-8(-11) mm wide; the outer larger, lanceolate to ovate, the inner small- er, lanceolate to cuneate. Filaments 3.3-6 mm long, united for 1-2 mm; anthers initially 6—9 mm long. Ovary 8-16(-20) mm long, the upper in the spathes, the pedicel short, 5-6 mm style arms 5-6 mm long. Capsule narrowly spin- dle-shaped, included in the spathes, 9-13 mm x 2-3 mm (subsp. virgata), or 16-22 mm x 4mm (subsp. karooica), only 1 (rarely 2) developed in each rhipidium; seeds narrowly angular, 0.7-1 mm wide and 1-2 mm long. Chromosome num- ber 2n = 12, 10 (subsp. virgata), or 2n = 14 (subsp. karooica). Flowering time. (Late September-)October- December(—mid January); flowers opening 3:00- :30 P.M. (or up to 4:00 P.M. on cooler days), beginning to fade ca. 6:30 P.M., usually collapsed by 7:00 P.M Distribution. Hexaglottis virgata is wide- spread in the southern African winter rainfall area, extending from the Nieuwoudtville district in the northwest through the western and south- ern Cape to Port Elizabeth in the east (Fig. 10). It is rather scattered in the west of its range, but common from Malmesbury and the Cape Pen- insula eastwards through the southern Cape. It is found more often on heavier soils, particularly shales of the Malmesbury System, but also occurs on granitic substrates. Occasionally H. virgata is found on sandy soils of the Cape System. Sub- species karooica occurs inland on the Roggeveld GOLDBLATT —SOUTHERN AFRICAN HEXAGLOTTIS 565 Escarpment, where it favors sheltered and damp situations. Diagnosis and relationships. Hexaglottis vir- gata is a distinctive species, unmistakable in its slender, relatively long perianth tube some 4-7 mm in length and very straight stems with strongly diverging lateral branches. The historic confusion about the identity of this species has been due to the poor observation that resulted in overlooking the perianth tube or confusing it with the slender ovary (Lewis, 1959). In fact the upper part of the ovary is narrow, sterile, and tubular, but this is clearly different from the peri- anth tube from which it is separated by an ab- scission layer. The only other species of Hexa- glottis with a perianth tube is H. brevituba, described in this paper. Hexaglottis brevituba has a much shorter tube, only 1-2 mm long, com- pared with a tube (3-)4-7 mm long in H. virgata. The two species differ in several other features. Hexaglottis virgata has an ovary entirely includ- ed in the spathes on a short pedicel up to 5 mm long, while H. brevituba has a longer pedicel 7— 10 mm long, and the ovary is curved and usually just exserted from the spathes. Two subspecies of Hexaglottis virgata are rec- ognized here. The typical and most common, subsp. virgata, has relatively small flowers and capsules. It has a wide distribution extending over almost the entire range of the species. Subsp. karooica, known only from two isolated sites on the Roggeveld Escarpment, has unusually large flowers, spathes, and capsules. History. The earliest existing records of Hexa- glottis virgata are those made by the Swedish botanists Carl Peter Thunberg and Anders Sparr- man in the 1770s, but this common southwest- ern Cape species must surely have been collected earlier than this. However, it was Nicholas Jac- quin who first described the species in 1791, based on plants grown in Vienna, and probably sent to him some years earlier by Franz Boos and Georg Schol, the collectors who provided Jacquin with many of the Cape plants that he illustrated and described. The painting that accompanies the de- scription and serves as the type in the absence of preserved material is excellent and leaves no doubt about the identity of what Jacquin called Moraea virgata. After the genera Homeria and Hexaglottis tinct from Mo- raea by E. P. Ventenat in 1808, M. virgata was 566 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 SUBSP. KAROOICA A SUBSP. VIRGATA FIGURE 11. Morphology of sre quia Is —A. Subsp. virgata. —B. _Subsp. karooica. Habits x 0. 25; corms x 0.5; flowers, fruiting branch (su and gynoecium full size; single capsule ale pn es virgata) + x 2: 1987] transferred to Homeria (Sweet, 1827) and shortly afterward to Hexaglottis (Sweet, 1830). J. G. Baker included H. /ewisiae (as H. flexuosa) and H. longifolia in H. virgata in Flora Capensis (1896). This unsatisfactory treatment was cor- rected by G. J. Lewis in 1950. KEY TO THE SUBSPECIES OF HEXAGLOTTIS VIRGATA la. Outer tepals 14-22(-25) mm long x 5-7 (711) mm wide; spathes (at least the terminal) ERU mm long; foliage leaves 1—3, rare- E subsp. virgata 0.5 1b. i dien 23-32 mm long x 7-1 wide; spathes (at least the zn 32-37 mm long; foliage leaves (2— ns subsp. karooica 6A. Subsp. virgata. Figure 11A. Hexaglottis virgata var. lata G. Lewis, J. S. African t. 25: 228-229. 1959. TYPE: South Africa. Cape: Welbedacht, Bidouw Valley, Lewis 2514 (holo- type, SAM 61821; isotypes, BOL, PRE, SAM). Plants 12—50(—70) mm high. Leaves 1—3(-4). Spathes 22-28(-32) mm long. Flowers with peri- anth tube (3—)4-5.5(-7) mm long; tepals 14- 22(-25) mm long. Filaments 3.3-6 mm long; an- thers 5-8 mm long. Ovary 8-12 mm long; cap- sules 9-13 mm long. Chromosome number 2n = 12, 10. Distribution. The distribution of subsp. vir- gata is the same as that for the species, except that it does not occur on the Roggeveld Escarp- ment (Fig. 10) Variation. Ascircumscribed here, subsp. vir- gata includes the variety /ata, described by G. J. Lewis for somewhat larger-flowered plants with comparatively wide tepals, from the Biedouw Valley. When this was described in 1959, no oth- er population of H. virgata was known from the northwestern Cape, and only one was known from Namaqualand, a poorly preserved specimen, which Lewis included in var. /ata despite the distorted flowers that could not be accurately measured. Hexaglottis virgata is now known from several sites in the northwest Cape, from the Oli- a tion in flower dimensions in these jomdan is considerable and covers the whole range from typical rather small-flowered subsp. virgata (te- pals 14-20 x 5-8 mm; spathes 22-28 mm long), which occurs in the Cape Peninsula and southern Cape, to the type of var. lata and very similar plants from Nieuwoudtville (tepals 20-25 x 9- GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS 567 11; spathes 23-30 mm long). The rationale for recognition of the variety has thus disappeared. These northwestern populations do, however, appear to represent a distinct race. Plants from two populations from the northwestern area have a karyotype with strongly acrocentric chromo- somes (Fig. 3F) that contrast with the presence of acrocentrics and submetacentrics in the most common southern cytotype (Fig. 3E Populations examined from the Cape Penin- sula have n = 5, and a very different karyotype from those Si n = 6 (Fig. 3G) (see discussion under Cytology). There seems to be no corre- sponding ipic grid difference in the Cape Peninsula plants. Specimens examined. SOUTH AFRICA. CAPE-31.19 (Calvinia): Grasberg road northwest of Nieuwoudt- ville, renosterveld (AC), Goldblatt 707 1 (K, MO, NBG, PRE, US); 2-3 km from Nieuwoudtville on the north side of hos road to the escarpment, Goldblatt 7411 (MO); 8 km south of Nieuwoudtville on sandstone on oo 7395 (MO 8 (Clanwilliam): 10 km south of Clanwilliam, j a bank (BB), Goldblatt 6705 (MO, PRE); 12 km south of Clanwilliam, stony clay bank, Goldblatt 6989 (K, MO, de PRE, WAG); clay hillside wi south of the Alp rnoff, on the National Road t Citrusdal (BD), pone 3028 (MO). 32.19 (Wuppertal): between Doorn River and Bi- douw Valley turnoff, stony clay (AA), Goldblatt 5941 (MO); Welbedacht, Bidouw Valley, Lewis 2514 (BOL, PRE, SAM 33.18 (Cape Town): Table Mountain (CD), Tyson 2488 (SAM), Bayliss 3054 (UC); kloof between Lions Head and Table Mountain, Burchell 252 (K); slopes of Devils Peak, above de Waal Drive, Cape Town, Gold- blatt 67 17 (MO); near Bishopscourt, Salter 9001 (BOL); Wynberg Hill, Pillans 10819 (MO, UPS), Salter 8978 (SAM), 9545 (BM); Signal Hill, Lewis 665 (SAM), Goldblatt 71 (J), 6747 (MO, S), 6748 (MO), Marloth 7234 (PRE); Camps Bay, Moss 13403 (J); Observatory grounds, Davis s.n. (SAM 61050); Tygerberg Nature Reserve (DC), Loubser 3004 (MO); Stellenbosch (DD), Boucher 3392 (PRE, STE); 4 miles from Faure on the Stellenbosch road, Lewis 2340 T Groot Dra- E und fuss ue POR Drége s.n (Worcester): farm Waterval, near ‘Porterville (AA), Loss 966 (NBG); Tulbagh Cemetery (AC), Goldblat aged K, MO, S); 9 miles along the Leeuwfon ad (AD) Pearson Deg (K); Bains Kloof (CA) uri tie 9106 (BM, BOL B west of (ox saa near the G, PRE); Karoo Gar- orcester, Compton 17849 (NBG), Lewis 5304 DEM Pok i Rawsonville, /e Roux s.n. (PRE); be- tween Worcester and Robertson (DA-DB), Zinn s.n. ice dea 3). 0 (Montagu): pasture pee 10 O’Clock Mt., Svellendam (CD), Wurts 486 (NBG). 3.21 (Ladismith): south dign to Attaquas Kloof m 4 Thompson 1636 (PRE), Attaquaskloof, Mossel Bay, Barker 7692 (NBG). 568 33.23 (Willowmore): hills near Uniondale (CA), H. Bolus 2484 (Ky; Prince Alfreds Pass (CC), Wall 18 (LD); near Misgund (CD), Goldblatt 6792 (MO); be- tween Misgund Net Nieuweplaats Pa Fourcade 5485 (PRE, STE); Kliprivier, Tsitsikamma Park, Bower 625 (PRE); hills hear Jouberti na (DD), Fourcade 2374 BOL, K); Joubertina, B BOL) 33.25 (Port Elizabeth): Van Staadens Hoogte (CC), MacOwan 2055 (BM, K); Greenbushes (CD), Holland 4051 (BOL); Baakens River Valley, Port Elizabeth (DC), Olivier 1739 (WAG); Port Elizabeth, Long 502 (K), —— sub Rogers 2414 (J). 4.18 (Simonstown): Steenberg (AB), Compton 1665 ER Bergvliet, Purcell s.n. (BOL 1638, SAM), 124 (SAM); Somerset West, on stiff clay soil (BB), Parker 4388 (BOL, K, MO, NBG). 34.19 (Caledon): Houw Hoek (AA), Penther 572 (K, S); Greyton-Genadendal (AB), Lindeberg s.n. (S); Na- pier Ruggens (BD), Marloth 10006 (PRE); between Na- pier and Bredasdorp (CA), Goldblatt 6937 (MO); Franskraal, along the coast above the beach (CD), Goldblatt 5368 (MO); sandy soil on slopes near Avoca (DA), Goldblatt 6939 (MO, PRE, S, WAG); Bredas- dorp Poort (DB), Esterhuysen 19580 (BOL). 34.20 (Bredasdorp): Storms Vlei Kloof (AA), Lei- poldt 3549A (BOL); Bontebok Park, Swellendam (AB), Liebenberg 6710 (STE); Zuurbraak (BA), Barker 5029 ( ); Grootvadersbos, paths in wood (BB), Willems shale hills west of Heidelberg, ren 7416 (MO); Potteberg (BC), David s.n. n BG); =a distr., Leipoldt 3550 (B 21 (Riversdale): hill top 5 km west í Riversdale Ps Goldblatt 5436 (MO); Still Bay, limestone hills (AD), Esterhuysen 19538 (BOL, PRE); limestone hills south of Albertinia, Stilbaai road on turnoff to Riet- huiskraal, Goldblatt 7428 (MO); Onverwacht, Alber- tinia (BA), Muir 1207 L). 34.23 (Knysna): Plettenberg Bay (AB), Rogers 28241 (K), 26762 Without precise locality: CBS, A/exander s.n. (BM, K); Sparrman s.n. Iris edulis (S); Thunberg s.n. Iris edulis (S) 6B. Subsp. karooica Goldbl., subsp. nov. TYPE: South Africa. Cape: eastern border of farm Blomfontein, 22 km from Middelpos to- wards de Hoop, Snijman 765 (holotype, NBG; isotypes, K, MO, PRE). Figure 11B. Planta robusta, 30-85 cm alta, ws (2—)4—5, spathis 32-37 mm longis, ind) arag 1 6.5-9 mm longo, tepalis 23-32 mm lon -11 mm S antheris 6—8 mm longis, capsulis TEN mm long Plants robust and 30-85 cm high. Leaves usu- ally 4-5, rarely 2-3. Spathes 32-37 mm long. Flowers with perianth tube 6.5-9 mm long; tepals 23-32 mm long, 7-11 mm wide. Filaments 4—6 mm long; anthers 6-8 mm long. Ovary 15-20 mm long. Capsules 16-22 mm long. Chromo- some number 2n = 14. Distribution. Subsp. karooica is known from two widely separated localities along the Rog- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 geveld Escarpment, near Blomfontein west of Middelpos in the north, and on the slopes of Sneeukrans on the farm Voelfontein northwest of Sutherland, some 50 km to the south. The habitat at both sites is moist, the plants growing among large rocks near seepage zones. Variation. The subspecies is variable in flow- er size. Plants from the Middelpos area have very large tepals 28-32 mm long while those from Voelfontein have smaller flowers with the tepals somewhat narrower and only 23-24 mm long. Despite this variation, other features, including large spathes and capsules, higher leaf number, and the unusual karyotype with 27 = 14, in both populations indicate that subsp. karooica is in- deed a distinct geographical variant of Hexa- glottis virgata. Specimens examined. SOUTH AFRICA. CAPE-31.20 (Williston): Roggeveld Escarpment, 71 km SE of Cal- vinia on Middelpos road via Blomfontein (CC), Gold- blatt 4612 (MO, NBG); eastern border of farm Blom- fontein, 22 km from Middelpos towards de Hoop, Snijman 765 (K, MO, NBG, PRE). 32.20 (Sutherland): Roggeveld, Sneeukrans south of Voelfontein farm, ca. 4,500 ft., in wet site (AD), Gold- blatt 6336 (MO), 7126 (MO). LITERATURE CITED BAKER, J. G. 1892. Handbook of the Irideae. George Bell & Sons, London 1896. Irideae. In W. T. Thiselton-Dyer (ed- itor), dis DRUMS 7-171. Reeve & Co., Ash- d, Ken 1932. Novitates Africanae. Ànn. 9. Opera Bot. 40: 1-85. . The systematics of Montanoa (Asteraceae, Heliantheae). Mem. New York Bot. Gard. 36: 1-133. GOLDBLATT, E 1971a. A new species of Gladiolus and some nomenclatural changes in the Iridaceae. 1 S. African Bot. 37: 229-236. 1971b. Cytological and morphological pud ies in the southern African Iridaceae. J. S. Afric Bot. 37: 317—460. 976a. Evolution, cytology and subgeneric classification in Moraea (Iridaceae). Ann. Mis- souri Bot. Gard. 63: 1-23. 1976b. The genus Moraea (Iridaceae) in the winter rainfall area of í Africa. Ann. Mis ri Bot. Gard. 63: 657- 1979. Chromosome Am and karyotype change in Galaxia (Iridaceae). Pl. Syst. Evol. 133: 61-69. Redefinition of Homeria and Moraea (Iridaceae) in the light of eat sui data, with Rheome gen. nov. Bot. 9. 1981. Systematics aa biology of Homeria pes Ann. Missouri Bot. Gard. 68: 413- 1987] . 1982. A synopsis of Moraea (Iridaceae) with new taxa, transfers and notes. Ann. Missouri Bot. rue 69: 351-369. Systematics of the southern African idaceae — Ixioideae). Ann 77-447 6. Convergent evolution of the Homeria flower type in six new species of Moraea (Irida- ceae — Irideae) 7 eae Africa. Ann. Missouri Bot. Gard. 73: oe E. P ZIMMER. 1984. Estima- tion of genome size (C-value) in Iridaceae by cy- tophotometry. Ann. Missouri Bot. Gard. 71: 176- l , M. & L. E. Copp. 1981. Botanical se of Southern Africa. A. A. Balkema, e wn. 66. Phylogenetic Systematics. Uni Illinois Press, Urbana, Illinois. [English 1 tine tion by D. D. Davis & R. Zangerl.] HUMPHRIES, C. J. 1981. Cytogenetic and cladistic studies in tue are (Compositae: Anthemideae). Nordic J. Bot. 1: 83-96. JACQUIN, N. 1776. Horus Botanicus Vindobonensis 3. Kaliwoda, Vien 91. oe tone virgata. In Icones Plantarum Rariorum 2: tab. 228. Wappler, Vienna. GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS 569 Krarr, F. W. 1866. Revisio Iridearum (Conclusio). Linnaea 34: 537—689. . 1882. Ergánzungen und Berichtigungen zu Baker's Systema Iridacearum. Abh. Naturf. Ges. Halle 15: 337-404 895. Irideae. In Th. Durand & H. Schinz (editors), Conspectus Florae Africanae 5: 143-230. Jardin Botanique de l'État, Bruxelles. Lewis, G. J. Cape Peninsula. Juta & Co., Cape Town. South African Iridaceae. A revision of Hexaglottis. J. S. African Bot. 25: 215-230. LINNAEUS, C. (FiL). 1781 [1782]. Supplementum Plantarum. Orphanotropheus, Braunschweig A. 1812. On the cultivation of rare plants. Trans. Hort. Soc. London -366. 1827. Hortus Britannicus, Ist edition. Ridgway, London pi. _ Hortus Britannicus, 2nd edition. Ridg- y VENTENAT, E P. "1808. Decades Generum Novorum Dufarb, Paris. NOTES ON THE VARIATION AND TAXONOMY OF WATSONIA BORBONICA (W. PYRAMIDATA, W. ARDERNEI) (IRIDACEAE) IN THE SOUTHWESTERN CAPE, SOUTH AFRICA! PETER GOLDBLATT? ABSTRACT Lomenia borbonica Pourret (1788) is an earlier name for the common Cape species formerly called Watsonia pyramidata, and the new combination in Watsonia is made here. Significant either declinate or arcuate, characterizes southern and =e ble. The whi red W. ar uced to synonymy. DUE synonyms of W. borbonica ted. Melee in the stamen and style orientation, rthern races which otherwise seem indist an n albino sport of the northern an t and unusual ite-flowe ernei is considered are discussed and the complete Musicien of the species is presente The name Watsonia borbonica is a new com- bination based on Lomenia borbonica Pourret (1788), only species of Lomenia Pourret. The epithet was given to a plant believed to have been collected on the Isle de Bourbon, now called Ré- union. Lomenia has long been misunderstood and was for a time considered a synonym of Freesia Klatt, a southern African genus. During a study of Freesia (Goldblatt, 1982), I discovered that Lomenia was congeneric with Watsonia. Based on the illustration in the protologue, I con- sidered L. borbonica conspecific with W. mar- ginata (L. f.) Ker, the basionym of which pre- dates Lomenia. Later, I found the type specimen of L. borbonica at the Paris Herbarium (P) in 1985. The specimen is in excellent condition, and there can be no doubt that it is identical with the species presently known as W. pyramidata (Andrews) Klatt, and in the Flora Capensis as W. rosea Ker (Baker, 1896). The type collection is attributed to Philibert Commerson, who col- lected in the Mascarene Islands in 1770-1774. of this and a few other Cape plants attributed to Commerson's collection remains unknown. Nothing could be more inappropriate than for a native southwestern Cape species to have a specific epithet ‘borbonica’. However, there is no alternative but to use the earliest valid and le- gitimate name for a species as mandated by the 1983 Code of Botanical Nomenclature. HABITAT AND VARIATION Watsonia borbonica is a fairly common and well-known species of Watsonia of the south- western Cape, South Africa. It is usually found in mountain habitats and is locally common and conspicuous after fires on lower slopes, but it also occurs on sandy flats and at relatively high al- titudes up to 1,000 m. Its range extends from the Cape Peninsula eastward to Bredasdorp and lo- cally north through the du Toits Kloof moun- tains to the Breede River floodplain near Ro- mans River. Plants flower in the early summer, from October to early December and at high elevations into January. They prefer rocky sand- stone soils but may be found on soils derived from granite. Although Watsonia borbonica is not particu- larly variable, what may be called the northern and southern races differ in one unusual feature, the orientation of the stamens and style. The southern and typical populations have declinate stamens that lie against the lower (anterior) tepal, and when the style is receptive it lies below the stamens. Northern populations from Paarl Mountain, du Toits Kloof, and the Breede River valley have unilateral stamens that lie horizontal to somewhat arched under the upper (posterior) tepal, and the style lies between the stamens and the upper tepal. In all other features the plants appear to be morphologically identical and can- not be distinguished even on close inspection. 1 oe by Grant DEB 81-19292 from the U.S. National Science Foundatio . Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri eee U. S.A. ANN. Missouni Bor. GARD. 74: 570—572. 1987. 1987] The diagnostic features of Watsonia borbonica are its large flowers with subequal, pink tepals 30-36 mm long; a short, flared upper perianth tube 8-12 mm long; robust and branched habit (1-2 m tall); oblong-ovate capsules 20-3 long; and seeds with two wings. Pink-flowered specimens of the northern form have always been included in W. borbonica (as W. pyramidata). However, a white-flowered sport believed to have come from the northern part of its range has been recognized as W. ardernei, a plant that has been in cultivation for close to 100 years and is the best-known cultivated Watsonia species. Wat- sonia ardernei is here reduced to synonymy in W. borbonica. The significance of the variation in stamen and style orientation is uncertain. It is clear that dec- linate stamens and style are unusual in Watsonia, occurring in only six other species. Arcuate sta- mens are more common and are found in all but one of the remaining ca. 45 species of the genus and in many other genera of Ixioideae. Presum- ably the declinate condition, rare in Iridaceae, is derived directly from the more common arcuate state and not independently from an ancestor with symmetrically disposed stamens. Despite the apparently gross morphological difference in stamen and style orientation between the north- ern and southern races of W. borbonica, I believe that they should be treated as a single species. I defer recognition of intraspecific taxa until my monograph of Watsonia, currently in prepara- tion, is completed. OTHER SYNONYMS Among other species I regard as synonyms of Watsonia borbonica is W. cooperi G. Lewis, based on the illegitimate homonym Tritonia cooperi aker. The type is a fragment comprising a lat- eral branch of the spike and three poorly pressed flowers. The leaf mounted with the flowering ma- terial has three conspicuous veins and is clearly misplaced, belonging either to a species of Ana- palina N. E. Br. or Tritoniopsis L. Bolus. How- ever, the flowering material seems to belong to W. borbonica. The tube and tepals are much dis- torted, but the stamens seem declinate, and the dimensions of the tepals and perianth tube ac- cord with W. borbonica. The reasons for treating Watsonia ardernei Sander and W. ardernei Hort. ex Mathews & L. Bolus as conspecific with W. borbonica have al- ready been outlined, but the existence of the GOLDBLATT — WATSONIA BORBONICA 571 homonyms applied to the same species is puz- zling. Possibly Mathews & Bolus believed that the name as described by Sander in a nursery- man’s catalogue was not valid, hence their formal recognition of a name already in use. The white- flowered form, originally collected at Romans River near Wolseley, on which W. ardernei (both of Sander and of Mathews & Bolus) is based is no longer known in the wild. It probably com- prised one or a few individuals that were intro- duced into gardens and then multiplied. The plant named W. iridifolia var. obrienii by N. E. Brown is based on cultivated plants from the same source as the type material of W. ardernei. The complete synonymy of W. borbonica is outlined below. All type specimens cited have been seen. SYNONYMY Watsonia borbonica (Pourret) Goldbl., comb. nov. Lomenia borbonica Pourret, Hist. & Mem. Acad. Roy. Sci. Toulouse (Acta Acad. Sci.) 3: 74 & tab. 5. 1788. TvPE: said to have been collected on Réunion (Ile de France) but clearly from the SW Cape, Commerson s.n. (holotype, P). Gladiolus pyramidalis Lam. egg Meth. 2: 726. 1786. wis : patens Aiton (Lewis et al., fete oo precise locality unknown, collector n (holotype, P—Herb. Lamarc G lado iridifolius varietas speciosa floribus roseis . Pl. tab. 235. 1793. A HAM a (Ker) Ecklon, Top. Verz. 37. 1827, nom. illeg., genus sine descr. Watsonia E (Andrews) Klatt, Durand & S . Africa 5: 194. 1895; Stapf, Bot. Mag. edm pes 0261. 1931; L. Bolus, Fl. Pl. Africa 25: tab. 974. 1946; Lewis, Flora Cape Peninsula 238.1950. Gladiolus pyramidatus Andrews, Bot. Rep. 5: tab. 335. 1803. Watsonia Hae Banks ex Ker, Konig & Sims, Ann. Bot. 1: 230. 1804, Bot. Mag. tab. 1072. 1807; Baker, Handbk. Irid. 158. Africa. Cape: precise locality unknown, 1 = in Bot. Rep. tab. 335 (lectotype here desig- ated). Watsonia striata Klatt, Abh. Nat. Ges. Halle 15: 352. ( elieved Ses hd Identity determined from descriptio — iridifolia var. pom ii N. E. Br., Gard. Chron se 0. 1889; Klatt, Durand & Sch inz, Con- = L l Africae 5: 193. 1895. TYPE: South Africa. Cap t known; authentic material present at Her . Kew (probably originally from Romans River farm, Tulbagh Div., flowered at Kew from plants sent from St. George’s Park, Port Eliza- beth). W. obrienii van Tubergen, Gard. Chron. ser. 3, 01. 1894, nom. nud. (not a valid pee A based on W. iridifolia var. obrienii as suggested by Marais, Kew Bull. 35: 172. 1980). Tritonia cooperi Baker, Handbk. Irid. 192. 1892, Flora Cap 122.1896, hom. illeg., non Klatt (1882), which is Tritonia cooperi (Baker) Klatt. Watsonia iL. Bo Bo : cooperi lus, J 67: 135. 1929; Lewis, J S. African Bot. 7: 55. 1941. Tritonia quinquener- vata Foster, Contr. Gray. H 36, comprises a flowering stem and poorly pressed flowers of W. borbonica and a leaf of a species of Anapalina or rii Watsonia ardernei Sander w Plants for d ie 20 & fig. 18 Watsonia ardernei Hort. ex Mathews & L. Bolus, Ann. Bolus Herb. 4: 25-26. 1925; Fl. Pl. Africa 19: tab. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 750. 1939, hom illeg., non W. ardernei Sander Sane due South Africa. Cape: Romans River farm, Tulbagh Distr. (cult. Kirstenbosch), Tred- midan n. we an atra BOL 17839, here Mcd E isolectotypes, BM (as Arderne s.n.), BOL). LITERATURE CITED BAKER, J. G. 1896. Irideae. /n W. T. Thiselton-Dyer (editor), Flora Capensis 6: 7-171. GOLDBLATT, P. 1982. Systematics of Freesia Klatt (Iridaceae). J. S. African Bot. 48: 39-91. Copp. 1981. Botanical Exploration 1788. Description de deux nouveaux genres de la famille des Liliacées désignés sous le nom de Lomenia & de Lapeirousia. Hist. & Mém. Acad. Roy. Sci. Toulouse (Acta Acad. Sci.) 3: 73- 78. NOTES ON THE FLORAL BIOLOGY, CYTOLOGY, AND EMBRYOLOGY OF CAMPYNEMANTHE (LILIALES: CAMPYNEMATACEAE)! PORTER P. Lowry II,? PETER GOLDBLATT,? AND HIROSHI TOBE‘ ABSTRACT Field observations of the endemic New Caledonian genus Campynemanthe (Liliales: Campynema- Pisin indicate that at least C. neocaledonica and C. viridiflora are strongly p oe and have an he filaments. A chromosome number of n= 11 has been determined for both species from meiosis in 1 pollen mother cells, these being first records for the genus and family. Endosperm formatio putatively related Melanthiaceae. Campynemanthe Baillon comprises three species restricted to the Pacific island of New Campynem a Tabi. and followed Dahlgren & Clifford due and Dahl- gren & Lu (1985) in placing both genera in Cam- pynemataceae. The family, which appears to oc- cupy an unspecialized position in Liliiflorae, is possibly most closely related to Melanthiaceae mall greenish flowers with persistent and ac- crescent tepals, partly inferior ovaries, and free stylodia. The embryology of both genera of Cam- pynemataceae is of the basic type for the mono- cots (Dutt, 1970; Dahlgren & Lu, 1985), except that the endosperm development is now known to be nuclear, a derived condition. We report here some observations on the floral biology of two species of Campynemanthe, in- cluding the occurrence of protandry and an un- usual postanthesis behavior of the persistent fil- am m counts for the genus and family, em we observed nuclear spect ofthe r n in C. neocaledonica is nuclear rather than helobial as in several genera of the embryology of Campynemataceae previously unknown. FLORAL BIOLOGY Recent field observations indicate that Cam- pynemanthe neocaledonica (Rendle) Goldblatt is strongly protandrous. This phenomenon was ini- tially observed in December 1985, in a popula- tion of about 100 individuals covering an area of ca. 20 m? located at 900 m on the Plateau de ers exhibited discrete male and female phases of sexual expression. In the initial, male phase the straight filaments are erect to ascending and are usually twisted slightly counterclockwise so that the oblong to ovate anthers alternate with the tepals (Fig. 1a). During this phase the three small stylodia are erect and closely appressed, formin (Fig. 1b) siderably and recurve to expose the whitish stig- matic surfaces. hile there appears to be no overlap in the sexual phases within a single flower, flowers of ! We thank J. H. Beach for many helpful suggestions on the manuscript, and J. K. Myers for preparing the illustration. Assistance was provided in New Caledonia by L. B. Thien, Ph. Morat, J.-M. Veillon, ORSTOM- ouméa , and the Service des Foréts et du Patrimoine Naturelle. Work by P . Lowry was supported in part by NSF Doctoral Dissertation Improvement Grant BSR83-14691, the Missouri Botanical Garden, the W. purs Jones Foundation, and the Division of Biology and Biomedical Sciences of Washington University, St. Loui ? Missouri Botanical Garden, P.O. Box 299, St. Phanérogamie, ? B. 63166- 0299. Louis, Missouri 63166-0299, U.S.A.; Muséum National d'Histoire Naturelle, 16, rue Buffon, 75005 Paris, France Krukoff Curator of African Botany, Missouri Botanical Garden, P. O. Box 299, St. Louis, Missouri and Laboratoire i * Biological Laboratory, Yoshida College, Kyoto University, Sakyo-ku, Kyoto 606, Japan. ANN. MISSOURI Bor. GARD. 74: 573-576. 1987. 574 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 2mm Flowers of Campynemanthe neocaledonica (Lowry 3945).—a. Male phase.—b. Female phase. FIGURE 1. both phases frequently occur within individual inflorescences. Geitonogamy thus appears pos- sible, assuming at least some self-compatibility. There did not appear to be synchrony of flow- ering among the individuals in the population, although observations were made on only a sin- gle da It has since been possible to confirm protandry in another population of Campynemanthe neo- caledonica from Haute Ouinné (Lowry & Suprin 3698) from photographs taken in the field. Sim- ilarly, two distinct floral phases can be seen in photographs of C. viridiflora Baillon taken at 950 m on the Montagne des Sources (Lowry 3762) and at 1,350 m on Mt. Humboldt (Lowry 3812), indicating that this species is dichogamous and almost certainly protandrous. Unfortunately, we have not been able to determine whether pro- tandry occurs in the third member of the genus, C. parva Goldblatt, or in Campynema lineare anthers, as figured by Dahlgren & Lu (1985) and reported by Goldblatt (1986). Bawa & Beach (1981) and Lloyd & Webb (1986), who have reviewed the evolutionary as- pects of dichogamy in general, and protandry in particular, concluded that dichogamy in self- compatible species permits pollen and stigmas to be positioned close to one another without high levels of self-pollination. They also pointed out that protandry is by far more common than protogyny in biotically pollinated species, which is presumably the case for C. neocaledonica. We did not observe floral visitors on species of Cam- pynemanthe. CHROMOSOME NUMBER A haploid chromosome number of n = 11 was determined from pollen mother cells in flower buds of Campynemanthe neocaledonica and C. viridiflora. Inflorescences of these species were fixed in the field with 3:1 ethanol-acetic acid and squashed with FLP orcein. The relatively large chromosomes were ca. 3 um long at meiotic metaphase but exhibited no distinctive details. Chromosome numbers in Melanthiaceae are diverse. The counts summarized in Table 1 were taken from the /ndex to Plant Chromosome Numbers series (Goldblatt, 1985, et praec.) and unpublished work (Ambrose, 1975; Bodkin, 1978). Tofieldieae apparently have x = 15, the only known base number in the monotypic gen- era Pleea and Harperocallis, and the most com- mon one in Tofieldia. Tofieldia coccinea Rich- ards. has x = 16, a count confirmed by a number of workers, but several other species have x — 15. Counts of n = 14 and n = 15 have been reported for T. calyculata (L.) Wahlenb. Japan- olirion, which perhaps belongs in Tofieldieae (Utech, 1984), has n = 12 In Melanthieae x = 8 is the most frequent base number and the only one in Melanthium, Schoenocaulon, and Veratrum. There are two distinct series in Zigadenus: n = 16 in Z. elegans Pursh, Z. glaucus Nutt., Z. sibiricus (L.) A. Gray ex S. Wats., Z. nuttallii A. Gray ex S. Wats., and Z. volcanicus Benth.; and n = 11 in Z. fontanus 1987] Eastw., Z. fremontii (Torr.) Torr. ex S. Wats., Z. brevibracteatus (M. E. Jones) Hall, and Z. ve- nenosus S. Wats. Amianthium has x = 16 (Utech, 1986), and Stenanthium has x = 8 and a puzzling second base of x = Narthecieae, the largest tribe in the family, ap- pears to be paleotetraploid. Narthecium, Aletris, Nietreria, an x = 13, while Helonias and Heloniopsis, now usually segregat- ed as a distinct tribe (Utech, 1984) or even fam- ily, have x = 17. Lophiola, perhaps allied to Nar- gi Sr P n = 21. The s genus Chionographis, placed in Chionographidea (Dahlgren et al., 1985), has = n C. koidzumiana Ohwi and C. japonica PO) Maxim. var. japonica, but C. japonica var. kurohimensis Ajima & Satomi has n = 22, and subspp. minoensis (Hara) Hara and hisu- achiana (Okuyama) Hara have n = 21. The most parsimonious interpretation here is a base num- ber o = 12, with reduction to x = 11 and polyploidy resulting in n = 22, then reduction to = 21. The taxonomically isolated XeropAyllum es ie has x = Summarizing, the available counts suggest to us a possible ancestral base number of x = 8 for Melanthiaceae, persisting only in Melanthium, Schoenocaulon, Veratrum, and Stenanthium (all Melanthieae). Other genera and tribes are ap- parently paleotetraploid, with Tofieldieae having x = 15 l ° = 17 reported in Helonias and Heloniopsis 1S difficult to reconcile with other members of the Narthecieae. The odd counts of n = 11 in Zi- gadenus, which has several species with x — 16, and n — 20 in Stenanthium, which also has n — 8, remain to be explained as well. Our report of n = 11 for Campynemataceae, which suggests a base number of x = 11, does not fit with any assemblage of Melanthiaceae. sumably being secondary to the higher numbers in the tetraploid series x = 16, 14, 13, 12, 11. EMBRYOLOGY Although it was not possible to pursue the de- tails of early endosperm development in Cam- pynemanthe, the available fixed material of C. LOWRY ET AL.—CAMPYNEMANTHE 575 TABLE 1. Chromosome numbers recorded in Cam- pynemataceae and basic chromosome numbers for genera of Melanthiaceae summarized from the /ndex to Plant Chromosome Numbers (Goldblatt, 1985, et praec.) and supplemented by data in Ambrose (1975), Bodkin (1978), and Utech (1984, 1986). Taxon (species Base Haploid counted/total species) Number Numbers Campynemataceae Campynema (0/1) Uncounted Campynemanthe (2/3) C. neocaledonica' 11 11 C. viridiflora' 11 11 Melanthiaceae Tofieldieae Pleea (1/1) 15 15 Harperocallis 15 15 1/1 Tofieldia (9/20) 15 15, 14, 16, 30 Japanolirion (1/1) 12 12 Melanthieae Veratrum (11/25) 8 8, 16, 40 48 Zigadenus (8/15) 16 16, 11 Schoenocaulon 8 8 /10) Stenanthium (2/2) 8 8, 10 Melanthium 8 Amianthium 16 16 Narthecieae Narthecium (2/5) 13 13, 26 Aletris (7/25) 13 13, 26 Metanarthecium 13 26 (1/2) Nietreria 13 Ypsilandra (0/5) Uncounted Helonias (1/1) 17 17 Heloniopsis (5/5) 17 17 Lophiola (1/1) 21 (or 7) 21 Chionographideae Chionographis 12 12, 21, 22 (2/5) Xerophylleae Xerophyllum (2/2) 15 15 ! Voucher data for original counts: C. neocaledonica, New Caledonia, Plateau de Dogny, 900 m, Lowry 3945 (MO, NOU, P); C. viridiflora, New Caledonia, Mt. Mou, 1,150 m, Lowry 3857 (MO, NOU, P neocaledonica (Lowry 3945), which comprised nearly 100 seeds that had developed past the 64- nucleate stage, indicates that the endosperm of this species is nucleate and not helobial (Stenar, 576 * ax * `; ° r 4 e vo SP Lea apa s Sem í € ahs sz - «M. NEL (c * Q e S š ^ Mt at 4 TASS o* Wise WU Sa BLES Ye FIGURE 2. Longitudinal section of a young seed of Campynemanthe neocaledonica showing free endo- sperm nuclei at the 64-nucleate pedi s, synergid; end, endosperm nucleus. Scale = 50 yu 1949; Eunus, 1951) as in other Melanthiaceae in which this embryological feature is known (in- cluding Tofieldia, Narthecium, Heloniopsis, Metanarthecium, Aletris, Zigadenus, Veratrum, and Amianthium). In Amianthium muscaetoxi- cum Walt. the embryo sac in postfertilization stages forms a micropylar and chalazal endo- sperm chamber characteristic of the helobial type; in an earlier stage the micropylar chamber con- tains 16 nuclei, and the chalazal chamber 8 nuclei (Eunus, 1951, fig. 31). In Campynemanthe, how- ever, there appears to be no such. chalazal cham- ri r, no nuclei at the chalazal end. All of the endosperm nuclei occur within a single cell (i.e., embryo sac), ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 although th bundant toward the cha- lazal end (Fig. 2). The presence of nuclear en- dosperm in C. neocaledonica suggests that this character may be useful in distinguishing Cam- pynemataceae from Melanthiaceae. LITERATURE CITED AMBROSE, J. D. 1975. Comparative Anatomy and Morphology of the Melanthioideae (Liliaceae). Ph.D. Dissertation. Cornell University, Ithaca, New Yor Bawa, K. S. &J. H. BEACH. 1981. Evolution of sexual systems in flowering plants. Ann. Missouri Bot. Gard. 68: 254-2 BopkiN, N. L. 1978. A Revision of North American Melanthium L. (Liliaceae). Ph.D. Dissertation. University of Maryland, College Park, Maryland. DAHLGREN, R. & H T Cu IFFORD. 1982. The Mono- cotyled p dy ic P London. A. M. Lu. 1985. Campynemanthe (Cam- pynemataceae): morphology, microsporogenesis, early ovule ontogeny and relationships. Nordic J. Bot. 5: 321-330. & P. - 1985. The Families of the Monocotyledons: ructure, evolution and tax- onomy. Springer- Verlag. Heidel DUTT, B. S . 1 Hypoxidaceae. Bull. Indian Natl. Sci. Acad. 41: 368-372. UNUS, A. M. 1951. Co "e to the embryology of the Liliaceae. V. Life of Amianthium numbers, 1982-1983. Monogr. Syst. Bot. Mis- sou E Bot. Gard. 13: 1-224. Sy saei and relationships of the d c Pacific family Campynemataceae (Lili- ales). Bull. Mus. Natl. es Nat., Paris, ser. 4, sect. , Adansonia 8: 2. LLOYD, D. G. & C. J. "e 1986. The avoidance of interference between the presentation of pollen and stigmas in angiosperms. I. Dichogamy. New Zea- land J. Bot. 24: 135-162. STENAR, H. 1949. Zur Kenntnis der Embryologie und der Raphiden-Zellen bei Bowiea volubilis Harvey und anderen Liliazeen. Acta Horti Berg. 15: 45- UTECH, F. H. 84. Floral vascular anatomy of Ja- panolirion osense Nakai (Liliaceae) and its tribal relationship. Ann. Carnegie Mus. 53: 447-461. 1986. Floral morphology and vascular anat- omy of Amianthium muscaetoxicum (Walter) A. Gray (Liliaceae-Veratreae) with notes on distri- bution and taxonomy. Ann. Carnegie Mus. 55: 481-504. A REVIEW OF THE NEW WORLD SPECIES OF ORTHROSANTHUS SWEET (IRIDACEAE)! JAMES E. HENRICH? AND PETER GOLDBLATT? ABSTRACT oe a genus of some 9 species of Iridaceae —Sisyrinchieae, occurs in Australia and South This review, arising out of research for floristic treatments for Flora Mesoamericana -— five American species: O. acorifolius and O. Oeerssapunpis are endemic in South America; to Mesoamerica and Mexico. The taxonomic history of Orthrosanthus i in the New World is outlined followed by a key, and the nomenclature, brief descriptions, and distribution ranges for each species are provide The genus Orthrosanthus comprises nine species of Iridaceae, tribe Sisyrinchieae Baker. It has an unusual disjunct distribution, occurring in Australia and central Mexico to South Amer- ica, a pattern shared in the family only with Lib- ertia, although the latter does not extend into Mesoamerica. In the New World the taxonomy of Orthrosanthus has long been confused, with several authors recognizing different numbers of thus is usually regarded as closely allied with Libertia and Sisyrinchium and is distinguished from them by an oblong to cylindric, included, and often sessile to subsessile ovary and capsule. Sisyrinchium and Libertia have globose to sub- spherical seeds without prominent angular ridges. Preparing a treatment of Iridaceae for Flora Mesoamericana and Flora de Nicaragua, we have reviewed the literature dealing with Orthrosan- thus and have examined the ample herbarium material now available. Our conclusions regard- ing the systematics of Orthrosanthus appear to merit the review presented here, since they differ extensively from currently accepted taxonomy, as represented in most major herbaria. Also, in- formation about Orthrosanthus is scattered in the literature and there is no modern summary of the systematics and geography of the genus in the New World. Cooke (1986) treated Orthro- santhus for Flora of Australia, where four species are now admitted. We recognize five New World species (Fig. 1): O. acorifolius (Kunth) Ravenna, O. chimboracensis (Kunth) Baker, O. exsertus short history of Orthrosanthus in the New World is outlined below, followed by a diagnostic key for the species and a review of their systematics and nomenclature. HISTORICAL REVIEW Orthrosanthus was erected by Robert Sweet (1827) for the Australian O. multiflorus Sweet. The first New World species ultimately assigned to Orthrosanthus were described by Kunth (1815) as Moraea (a distantly related African genus, tribe Irideae). Kunth described three species now rec- ognized as Orthr th Moraea chimboracen- sis, M. gladioloides, and M. acorifolia. Baker (1876: 113) was the first to include New World species in Orthrosanthus, recognizing O. chim- boracensis (Kunth) Baker (with M. acorifolia as a synonym and M. gladioloides as a variety) and a second species O. spicatus (Baker) Baker from South America. The latter is a short plant with a winged flowering stem and congested, sessile inflorescence units (rhipidia). The flowers have a short perianth tube, and the globose capsules are carried on slender pedicels above the sub- tending bracts as in Sisyrinchium and Phaio- phleps. Ravenna (1968) placed the species in Phaiophleps as P. brasiliensis, but we doubt that it belongs in this heterogeneous alliance. It should ! Supported by Grant DEB 81-19292 from the United States National Science Foundation. A. 2 Missouri Botanical Garden, P.O. Box 299, St. ouis, Missouri 63166, o U.S. 3 B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. ANN. Missounmi Bor. GARD. 74: 577-582. 1987. 578 be referred back to Sisyrinchium, pending critical study of its affinities. Baker’s (1892) later, more detailed treatment recognized seven species in Orthrosanthus, the same two in the New World and five in Australia. In contrast to Baker’s treatment, Klatt (1861) first placed New World species of Orthrosanthus in Sisyrinchium. He described the distinctive central Andean O. occissapungus and a p f O. chimboracensis, S. moritzianum. Klatt (1882) placed New World Vedi in Solenomelus Miers, recognizing Solenomelus chimboracensis (including M. acorifolia and Sisyrinchium moritzianum) and Solenomelus aet ed (including Sisyrinchium occissapun- , close of the nineteenth century Rus- by dun ed O. nigrorhynchus, not real- izing that Sisyrinchium occissapungum was an earlier name for the plant, and two years later Kuntze described the conspecific O. tunariensis. Orthrosanthus occissapungus has white flowers, characteristic long narrow capsules, and typically single- to few-flowered rhipidia. A review of Orthrosanthus by Steyermark (1948) is consistent with Baker's earlier treat- ment in recognizing only one American species (at least he makes no reference to O. occissapun- gus or its synonyms). Steyermark's contribution is noteworthy for recognizing four varieties and two forms distinguished by capsule size, degree of exsertion of the capsule pe the SAQUE eaf h bracts, flower color, inflorescenc rona and ees distribution. , Ravenna (1965) described Orthro- mo ASN (= O. chimboracensis var. centro-americanus Steyerm.) and made the com- bination O. monadelphus subsp. exsertus (R. Foster) Ravenna, Ravenna (1969) also Pise O. O. chim- boracensis s subsp. tunalieniis (Kuntze) Ravenna. Without mentioning Steyermark's (1948) treat- ment of the genus, in particular O. chimbora- censis var. acorifolius, Ravenna (1977) trans- ferred Moraea acorifolia to Orthrosanthus as O. acorifolius (Kunth) Ravenna. In 1981 he indi- cated that he regarded Orthrosanthus as com- prising four species and one subspecies. TAXONOMY Orthrosanthus Sweet, Flora Australasica, t. 11. l Baker, Handbk. Irid. 117-119. 1892; Cooke, Fl. Australia 46: 10-13. 1986. TYPE: O. multiflorus Sweet. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Elvetria Raf., Fl. Telluriana 4: 30. 1838. TYPE: E. mul- tiflora (Sweet) Raf. (= O. multiflorus Sweet). Evergreen tufted perennials with short, persis- tent, creeping rhizomes. Leaves several, mostly basal, ensiform and equitant, linear to linear- lanceolate, coriaceous, without a midvein, crowded at the apex of the rhizome. Flowering stem erect, branched, the iones ascending, either long or short, term t relatively short; spathes subequal. Flowers acti- nomorphic, subsessile or shortly stalked, blue or white; tepals free (united in a short tube in some Australian species), subequal, spreading hori- zontally from the base; filaments free or united in the lower half, anthers erect; ovary included in the spathes, + sessile, or exserted in O. exser- tus, style short, dividing into 3 relatively long branches extending between the stamens, the branches stigmatic apically. Capsules ellipsoid to cylindric, sometimes pubescent, included or exserted; seeds angular to fusiform. Species: 9; highland areas from Mexico to Pan- ama, the Coastal Cordillera of Venezuela, and through Andean South America to Bolivia and northern Argentina (Fig. 1), and in Australia. KEY TO THE NEW WORLD SPECIES la. ead with pedicels is a 15(-25) m long; capsule glabr laf 2a. capsules slender, Pin e mm long: seeds elongate-angular, 2.5-3 mm E 0.5 mm ua (Peru, Bolivia, and nort ern Arge . $. O. oceissapungus Spathes ira an (2-)3 or more flow capsules oblong to ellipsoid, 10-18 mm long; seeds angular to irregularly globose, 1.2-2 mm long and wi 3a. Mature capsules borne well above the subtending bracts; pedicels 15- 25 mm long (central Mexico: Mex- ico, Puebla, Distrito Federal, Ta- maulipas, Nuevo Leon, Michoacan) . O. exsertus . Mature capsules usually included; pedicels 5-10 mm long (Costa Rica, ode: Colombia, bows dor, EEEE ; imboracensis ] a ide or with short p ne m long; ca ipa sue aT DE se t3 g Uu og c m long; leaves 2-4 mm wide (southern Mexico to northern Pan- ama, except Belize and Nicaragua) ....... 4. O. monadelphus 4b. Capsules 8-11 mm long; leaves 4-12 mm wide (Venezuela) ...... 1. O. acorifolius 1987] HENRICH & GOLDBLATT—NEW WORLD ORTHROSANTHUS SWEET 579 I = | RISIN | | | | ee X | I] = | | | | | | | 1 n | | jt l NM € | | | — | "e | | | | VENT O. EXSERTUS | | | || | 1 O. MONADELPHUS I | | | ——ss,"Nr —— I. | ` x _ (0T | Cc 39 CTI +. Bs: * e. wi £ | O. ACORIFOLIUS or c I O. CHIMBORACENSIS | | | | e pm | | | = | | | | | —r | | | 10h | | | | jen | | — O. OCCISSAPUNGUS | \ \ dl--- A \ t T | 20 \ -k7 \ \ p — \ — \ o 200 400 600 800 1000km \ O 100 200 300 400 500 800 miles \ \ I | 100 90 FiGurE 1. Distribution of the species of Orthrosanthus in the New World. l. Orthrosanthus acorifolius (Kunth) Ravenna, M santhus chimboracensis var. acorifolius (Kunth) Steyerm., Lloydia 11: 19. 1948. Marica acorifolia (Kunth) Martens & Ga- leotti, Bull. Acad. Brux. 10: 110. 1843 (mis- applied, probably to O. monadelphus). TYPE: Venezuela. Distrito Federal: in crepidinibus montis Silla de Caracas, ca. 1,250 m, Bon- pland & Humboldt s.n. (lectotype, P— Gen- barium; the sheet is designated more type material probably existed and may be found. iab erede chimboracensis Mi qiia f. albus Steyerm., Lloydia 11: 19. YPE: Venezuela. Trujillo La Quebrada oe by d line Lara-Trujillo rmark 55339a (holotype, F). Plants to 65 cm tall. Leaves to 50 cm long, (0.4-)0.9-1.2 cm Il broadly linear, gradually stem about as long as the long, pubescent; seeds angular to somew rounded, ca. 1-1.2 mm at the widest diameter. Distribution. Orthrosanthus acorifolius is ap- parently restricted to the Coastal Cordillera and Andes of Venezuela and eastern Colombi This species is usually very robust and has comparatively broad leaves sometimes matched 580 in Orthrosanthus by those of O. chimboracensis. The ovary and young capsules are heavily pu- bescent, as in O. monadelphus, to which O. aco- rifolius is probably most closely related. 2. Orthrosanthus chimboracensis (Kunth) Ba- ker, Gard. Chron. n. ser. 6: 67-68. 1876. Orthrosanthus chimboracensis var. typicus Steyerm., Lloydia 11: 15-16. 1948. Moraea chimboracensis Kunth, Nov. Gen. & Sp. PI. : 322. 18 Tunguragua: Chim- Urcu, ca. 1,640 m, Bonpland & Humboldt s.n. (lectotype, P—Herb. Bonpland, here designated). More type material may be found; the sheet we have designated as lec- totype has the data “Chimborazo” only and the number 3/88. iini: a w ms Gen & Sp. Pl. 1: 322. . Cajamarca: locis frigidis Pe- m et Micuipampam, ca Sisyrinchium moritzianum Klo si x Klatt, Linnaea 78. 1. TYPE: Venezuela (as Colombia). erida: Paramo de la Culata, ae 1204 (iso- type, K). Plants (20—)60—115 cm tall. Leaves 15-70 cm long, 8-12 mm wide, narrowly lanceolate to lin- £e = blue. Capsules 14-18 mm long, glabrous, the pedicels 5-10(-15) mm long; seeds angular, 1.2— 2 mm at the longest axis. Distribution. | Orthrosanthus chimboracensis ranges from Costa Rica and Venezuela south- ward through the Andes from Colombia to Peru. It is a montane species seldom occurring below 2,000 m. Plants are typically less robust than Orthro- santhus acorifolius and can be recognized by a sparsely pubescent to glabrous ovary and com- pletely glabrous capsule with a short pedicel 5- l long 3. Orthrosanthus exsertus (R. Foster) Ravenna, Wrightia 7: 10. 1981. Orthrosanthus chim- boracensis var. exsertus R. Foster, Contr. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Gray Herb. 155: 49. 1945. TYPE: Mexico. Distrito Federal: on the sides of ravines near Eslava, Pringle 8827 (holotype, G; isotypes, C, CAS, F, G, GH, MEXU, MO, NY, O,S, TEX, UC, US). Orthrosanthus chimboracensis var. exsertus f. albus m., Lloydia 11: 17. 1948. TYPE: Mexico. Tamaulipas: Santa Rita, Ranch Tamaulipas, 1,500 m, Runyon 875 (holotype, US; isotype, TEX) Plants 40-70 cm tall. Leaves 40-55 cm long, 3-8 mm wide, narrowly acuminate. Flowerin stem somewhat longer than the leaves; rhipidia several-flowered. Flowers to 2.5 cm diam., blue, or occasionally white; ovary exserted from the spathes shortly after anthesis. Capsules 1-1.7 cm long, glabrous, broadly acuminate apically, well exserted from the spathes on pedicels 15-25 cm long; seeds angular to somewhat rounded, 1.2- 1.5 mm at the widest diameter. Distribution. Orthrosanthus exsertus is en- demic to highland areas of southern and central Mexico (Nuevo León, upon. Puebla, Mi- choacán, Distrito Federal, Méxic The smallest of the New World pam of Or- throsanthus, O. exsertus is distinctive in having comparatively narrow leaves and glabrous cap- sules exserted from the spathes and bracts on pedicels 15-25 mm long. Recognized only in 1981 as a distinct species, and first as a variety in 1948, Orthrosanthus exsertus was recorded as early as 1829 or 1830 by Schiede and a few years later by Liebmann. Their collections are among the three syntypes cited by Klatt for Sisyrinchium occissapungum, now O. occissapungus, and neo- typified here by a collection from Peru. 4. Orthrosanthus monadelphus Ravenna, Bol. Soc. Argentina Bot. 10: 317. 1965. TYPE: Guatemala. Baja Verapaz: cumbre El Chol, ca. 2,200-2,500 m, Ravenna 266 (holotype, Herb. Ravenna, not seen; isotypes, F, HBG). Orthrosanthus chimboracensis var. diis americanus Steyerm all: 19. € 48. T of rae unción Mita, Steyer- mark 31913 fliclotvne. F). Mice chimboracensis or intermedius Stey- Lloydia 11: 19-20. 1948. TYPE: Costa Rica. iei Volcán Irazu, Allen 674 (holotype, F). Plants (15—)30—60 cm tall. Leaves 30-45 cm long, 8-10 mm wide, linear. Flowering stem to 60 cm; rhipidia several-flowered and spaced somewhat irregularly along the stem; spathe 1987] margins narrowly hyaline. Flowers 1.4-3 cm diam., blue. Capsules 10-13 mm long, usually pubescent, sessile or nearly so, only the apices exceeding the spathes; seeds angular to some- what rounded, 1.2-1.5 mm at the widest diam- eter. Distribution. Orthrosanthus monadelphus is common in highland areas of Mesoamerica. It is recorded from southern Mexico to Guatemala and El Salvador, and locally to the south in Costa Rica and northern Panama. This species is probably most closely allied to the Venezuelan Orthrosanthus acorifolius, with which it shares a similar, nearly sessile, and pu- bescent ovary and capsule. The filaments are united in the lower half, not much more so than in the other American species. Although first rec- ognized as a distinct subspecies in 1948, and as a species in 1965, it is interesting to note that the collection cited by Martens & Galeotti (1843) as Marica acorifolia may be the first record for Orthrosanthus monadelphus [Mexico: Oaxaca, rochers gneissiques de Penoles, Misteca alta, ca. 2,150-2,300 m, Galeotti 5368 (?BR, not seen)]. 5. Orthrosanthus occissapungus (as O. ocisa- punga) (Ruiz ex Klatt) Diels, Engler & Prantl, Nat. Pflanzenfam. ed. 2. 15a: 478. 1930. Sisyrinchium occissapungum Ruiz ex Klatt, Linnaea 31: 379. 1861. NEOTYPE: Peru. La Libertad: along the Río Negro 3 km south of Huamachuco, West 8113 (neotype, MO; isoneotypes, GH, UC). The three cited syn- types of S. occissapungum have not been found and are presumed destroyed, hence our designation of a neotype. For complete- ness, the syntypes are cited here as follows: Peru. Huanuco: ad Pillao etc., Ruiz ex Herb. Lambertii (B, not seen); Mexico. Oaxaca: Chinantla, Liebmann 310 (location un- own) (? = O. exsertus); Mexico. Veracruz: Jalapa, Schiede 1029 (location unknown) (? = O. exsertus). Orthrosanthus nigrorhynchus Rusby, Mem. Torrey Bot. C 26. 1896. TYPE: Bolivia. Cochabamba: near Cochabamba, Bang 1074 (lectotype, NY, here designated as the best preserved and most com- plete of three sheets at NY, the institution where Rusby worked; isolectotypes, BM, F, G, GH, K, MO, : Orthrosanthus tunariensis Kuntze, Rev. Gen. Pl. 3: 309. 1898. TYPE: Bolivia. Cocha ba: im Tu- narigebirge, 3,600 m (collecto and location of the type not in the protologue and unknown to us). Orthrosanthus EE siberaceasis subsp. tunariensis HENRICH & GOLDBLATT — NEW WORLD ORTHROSANTHUS SWEET 581 (Kuntze) Ravenna, Revista Inst. Munic. Bot. (Buenos Aires) 2: 30. 1969. Orthrosanthus chimboracensis sensu Rusby, non (Kunth) Baker, in Bull. Torrey Bot. Club 29: 224. 01. Plants to 80 cm tall. Leaves 30-50 cm long, -8 mm wide, narrowly linear, sharply acute. Flowering stem to 80 cm; rhipidia typically sin- gle-flowered. Flowers 2.6-3.4 cm diam., white. Capsules (1 5-)22-30 mm long, glabrous, extend- ing above the spathes on pedicels to 5 mm long; seeds elongate-angular, 2.5-3 mm at the longest axis. Distribution. Orthrosanthus occissapungus is restricted to northwestern Argentina, Bolivia, and Peru, where it grows at elevations above 3,200 m. The most unusual of the New World species of Orthrosanthus, O. occissapungus can be rec- ognized by its one- to few-flowered spathes, long slender capsules, Fei elongate seeds. The flowers are always white in O. occissapungus and only occasionally so in de species in the New World. As indicated in the nomenclature above, we have been unable to locate the original type ma- terial of Sisyrinchium occissapungum. The syn- type from Peru, collected by Ruiz, had the manu- script epithet '"*occissapungum" according to Klatt. It was housed in the Berlin Herbarium and must be presumed destroyed. It was almost cer- tainly conspecific with O. nigrorhynchus de- scribed in 1896. The two other specimens cited in the protologue are from Mexico: Schiede 1029 (also presumed destroyed) from Jalapa in Ve- racruz, and Liebmann 310 from Chinantla, Oa- xaca. The latter specimen is cited as “Herb. mihi" by Klatt, that is, in his own herbarium, which was at one time at Berlin but is now at Stock- holm. No Liebmann collection with this number has, however, been located at either place. Du- plicates of the Liebmann collection at Copen- hagen, where Liebmann’s main collection is pre- served, likewise do not include the missing type number. Other Liebmann specimens at Copen- hagen and at Paris collected at the same locality and time are the Mexican Orthrosanthus exsertus and represent the first collections of that species. e are reg g pung as a valid combination made by Diels (1930: 478) but recognize that it would not be accepted according to the current Code of Botanical No- menclature if made today. Diels did not cite a basionym or indicate in any way that he was making a combination in citing the species as O. ocisapunga (sic) Ruiz. Nevertheless, there can be 582 no doubt that he was placing Sisyrinchium oc- cissapungum Ruiz ex Klatt in Orthrosanthus as the earliest name for the species known until this time as either O. nigrorhynchus or O. tunariensis, neither of which he mentioned in his enumera- tion of the species of Orthrosanthus. The name O. occissapungus was accepted by Macbride (1936) and Vargas (1944) but was attributed by both authors to Diels (as O. ocisapunga Ruiz ex Diels). We do not believe that Diels intended to describe the species, and therefore we cannot ac- cept this treatment. We also note that it was not Diels’s practice to cite the authors of species in later combinations e.g., his listing of O. chim- boracensis Bak. [instead of (Kunth) Baker], and so there is no inconsistency in his not mentioning the authors of the basionym for O. occissapun- gus. LITERATURE CITED BAKER, J. G. 1876. Systema Iridacearum. J. Linn. Soc., Bot. 16: 61-180. 1892. Handbook of the Irideae. George Bell n. 1986. Iridaceae. In Flora of Australia DIELs, L. 1930. Iridaceae. Jn A. Engler & K. Prantl (editors), Die Natiirliche Pflanzenfamilien, ed. 2. 15a: 463-505. Foster, R. C. 945. Studies in the Iridaceae. III. Contr. ed Herb. 155: 3-54. KLATT, F. W. 1861. Berichtigungen und Nachtráge zu der Monographia Generis Sisyrinchium. Lin- naea 31: 370-386. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 . Erganzungen und Berichtigungen zu Baker's Systema Iridacearum. Abh. Naturf. Ges. v — Il 815. ys de nei s Bon- Latine-Allemande, Paris. Gide Fils, Paris. . Iridaceae. In Flora of Peru. Mus. Nat. Hist. Bot. Ser. 13 (1, no. 3): 707- 717. Martens, M. & H. GALEOTTI. 1843. Enumeratio syn- optica plantarum phanerogamicarum ab Henrico Bull Acad. 2 Sci Belgique 10: 110-134. RAVENNA, P. 1965. Notas sobre Iridaceae. II. Bol. Soc. ae Bot. 10: 311-322. l . Notas sobre Iridaceae. III. Bonplandia 2: 27 3-291. . 1969. Notas sobre Iridaceae. IV. Revista Inst. Munic. Bot. (Buenos Aires) 2: 25-38. 1977. Notas sobre Iridaceae. V. Mus. Nac. Hist. Nat. Santiago Notic. Mens. 21: 7-9. . 1981. On the presence of the genus Orthro- (Irid )in the Argentine fl Wrighti 7: 10. RusBv, H. H. 1896. On the collections of Mr. Miguel E in Bolivia, Part III. Mem. Torrey Bot. Club 6: 1-130. ee J. A. 1948. Orthrosanthus chimbora- censis and its varieties (Iridaceae). Lloydia 11: 14— 20. Sweet, R. 1827. Flora Australasica, t. 11. James Ridgway, London. VARGAS, C. C. 1944. Iridaceae Cuzcoensis. Revista Univ. Cuzco 33(87): 167-177. PATCH FORMATION AMONG ISRAELI CRUCIFERS: HOW DO THEY GET AWAY WITH IT?! MICHAEL AUERBACH? AND AVI SHMIDA? ABSTRACT ny m in be ee and desert floras of Israel frequently form large monospecific r . This ted Man patches but do theory of in herbivore loads, a defense against herbivores. In editerr. Papilionaceae, Apiaceae). This is not true for desert year to year. allow Mediterranean patch-forming species to of herbivory. In contrast, desert pa cies ee pins Se rely on pe ropa e phe ovre as wi s. Patc MEA Among . agis PAE patc on-patch formers. In addition, patch for A interactions which predicts that suc nd that herbaceous plants iion rely on unprediethbilid in time and space as a Isr nean crucifers fl of o species-rich families with similar sciri da orms (Lami aceae, se Liliaceae, Solanaceae, Poaceae, tch formers are "ale and have larger pet mation is correlated situation is unexpect from some recent should attract high ower earlier than members of other ucifers We hypothesize that their displaced phenologies id: possession of potent VIDA uce monocultures while not sustaining high lev E as y be reinforced by plant-pollinator tals than do vels of floral ultraviolet ormation m high le ier aed and patterning. These differences may result from perdes: iüerepecific but enhanced intraspecific competition for pollinators among pa tch formers. These patterns are not iud among desert species, although desert patch formers do initiate flowering before other crucifers. The family Brassicaceae (Cruciferae) has fig- ured prominently in the development of theory concerning insect—plant interactions. Responses & Feeny, 1977), as well as considerations of fam- ily-wide attributes (Feeny, 1976, 1977), com- prise much of the empirical and theoretical sup- port for concepts such as ''associational resistance" (Tahvanainen & Root, 1972) and *plant apparency" (Feeny, 1976). Feeny (1977) also cited family-wide attributes of crucifers to support his contention that escape in time and space and allelochemic diversity protect herba- ceous plants from potential herbivores. Several characteristics of crucifers make them particu- larly amenable for studies of herbivore-plant in- teractions. They are well represented in floras of many regions of the world (Hedge, 1976), all species thus far examined contain glucosinolates (Kjaer, 1976), and many species are important cultivated crops. The family contains approximately 400 genera and 3,000 species, most of which are annual herbs (Vaughan et al., 1976). The Irano-Turanian re- gion is home to about 900 crucifer species and was probably the center of origin for at least the Old World taxa (Hedge, 1976). Brassicaceae is a dominant and conspicuous family in terms of ticularly in late winter and early spring (Shmida & Auerbach, 1983). Most of the 111 or more species native to Israel are Irano-Turanian or are descendants of Irano-Turanian stock (Zohary, 1973). Crucifers frequently form enormous mono- r plant interactions which predicts that such re- source concentration should attract high herbi- vore loads, and that herbaceous plants should rely on unpredictability in time and space as a defense against herbivores. Here, we examine the crucifer flora of Mediterranean and desert re- gions of Israel in terms of plant morphology, phenology, and propensity of some species to occur naturally in large monocultures. In partic- ! We thank Nitsa Dagan for assistance with data analysis and Diana Lieberman for her critical review of this manuscript. William Kunin and Micha Boaz provided unpublish ed data. Most analyses presented here were conducted while one of us Sl was a Lady Davis Postdoctoral Fellow in the Department of Botany, Hebrew Evers. The support and e ? Department of Biology, ir. of Nort ANN. MIssourRI Bor. GARD. 74: 583—594. 1987. y akota, Grand For 3 Department of Botany, Hebrew University, Jerusalem, Israel. Davis Trust is gratefully acknowledged. s, North Dakota 58202, U.S.A. 584 ular, we address the question, “how do patch formers avoid colonization by a high density and/ or diversity of phytophages’”? ISRAELI CRUCIFERS We obtained data on plant species from Flora Palaestina (Zohary & Feinbrun-Dothan, 1966), herbarium records of the Hebrew University De- artment of Botany, Hebrew University, field observations from January to June 1983, more than 10 years of vegetation samples taken by A.S., and phenological records from ECOPAS. The last is a computer data base compiled by members of the Department of Botany, Hebrew University, and the Society for the Protection of Nature in Israel. Each month phenological rec- ords are submitted by a network of observers, resulting in a comprehensive record of Israel’s ora. Tables 1 and 2 list species of native crucifers growing in the Mediterranean (76 species) and desert regions (37 species) of Israel (two species are listed in both regions). For each species we show growth form, floral characteristics, pubes- cence, abundance in Israel, and a subjective mea- sure of size of monospecific patches. Within both assemblages we excluded extremely rare species that we never encountered in the field (approx- imately 20 species). We must clarify one entry in Tables 1 and 2. Our index of patch formation ranges from “+++” (largest patches) to ““—” (no patches). Although this categorization is subjective, we classified species as patch formers only if they formed large monospecific associations. A patch in Our usage is not a clump of several individuals of one species; rather, it is an extensive single- species association. Patches range from approx- imately ten meters to several kilometers on a side. For instance, in spring 1983 (highest spring rainfall in 30 years), we measured patches of Eru- caria hispanica L., near Arad, Israel, and Sinapis arvensis L., near Bet She'an, Israel, that extended combined all patch formers for statistical anal- yses. Using these data, we compared phenological attributes of the Brassicaceae with those of seven other species-rich families with similar growth forms in Israel (Table 3). Phenological compar- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 isons were based on peak flowering time (month in which most individuals of a species produce most of their flowers) and on range of flowering time (range of months in which flowers have been observed for each species). For each family we constructed cumulative frequency distributions of flowering range and peak and compared the of association (Kolmogorov-Smirnov 2 sample test, Conover, 1980). We also compared average month of peak flowering between the Brassica- ceae and other families with a parametric test (Welch approximation of the t-test, Remington & Schork, 1970), since these data were approx- imately normally distributed. We limited these comparisons to Mediterranean species because desert species are constrained to flowering over a short, variable period when rains occur. Com- parisons were done on all species and the subset of annual species for each family, except for Lil- iaceae which only has one annual species. e then compared morphological and phe- nological attributes of patch- and non-patch- forming crucifers. These analyses were conduct- ed separately for Mediterranean and desert species. We compared phenologies based on cu- mulative frequency distributions of flowering range and flowering peak and average month of peak flowering. In addition, we compared the average month of start of flowering using a para- metric test (Welch approximation of t-test). Av- erage plant height and petal size from ECOPAS were compared using f-tests, whereas analyses of associations of pubescence and ultraviolet pat- terns with patch formation were based on two- by-two contingency tables. RESULTS INTERFAMILY DIFFERENCES Comparison of the phenology of Mediterra- nean crucifers with members of other families indicates that in general crucifers flower earlier in the year, both in terms of flowering range and flowering peak (Table 3). The single exception is no significant difference between the peak flow- ering time of the Brassicaceae and Liliaceae (Ta- ble 3). The average duration of flowering of all crucifers differs from that of all Lamiaceae, As- teraceae, Solanaceae, and Poaceae (Table 3). Among all families we examined, medians of flowering range and peak occur earlier for annual species than for perennials except for the Solana- ceae, in which the situation is reversed. Peren- 585 AUERBACH & SHMIDA—ISRAELI CRUCIFER PATCH FORMATION 1987] +k + Ə SI A £-l - Ot v ‘Od CT) sapioonaa sixvjojdiq = O t 9 t-z + Sz V pueig xo qqoA (7T) vIydos viuipanosoq * . t ^ v-t * $9 V "] 2oruvdsij 2quiva?) + XI [4 ^ t-z = OI V "osy (Texss1oJA) sn;bpuvnbs sndouosoD = O OI A r _ os V "zipuy (7T) sijn1u2140 713u14407) = aa T Ə £-I ++ OI V 'ssiog 222nddvj `) ú OO [4 9 £-I ++ 8 V 7] idsppuiuof 2 = u Ç e) €I + OI V [Lun] (n210) vads vjoaddjD + aad 8I d r-t + SC v 8143 (‘JOS 19 squeg) suaospnandand puodsiiot/Ə E OT 6 dO r-t ++ SI V ‘Jd CT pnuup v421214407) PET JI t A r-t + LC H "ASIQ (7T) 9?qpap DUDpADD = aa C ^ v-r = 0c V "1 Dinsaly 2aunupav?) = O [4 ^ T-I = SI V Janay DjJ2qna `J Zi OO [4 AA r_I = 8I V snxrpojN (7T) $24078pd-pbsanq vjjasdp.) = e t ^ £-I — 8I V ‘OyL (Ossy) si4Dpn82441 puidə]poO + 99 6 d TI-E = O€ V 'doos pupu 2]D/22) a 9 L Ə b-t = 08 V uenor) /740/2u4n0] 'q +++ OT OI X 9-v = OcI V yooy ("D patu `g = O cI Ə c+ = 09 NHO ‘SSN 2217242 DIISSDAG = OO t AO L2! t Lg V “| puuápip vjjmosig = e L ^ r + SI V ug ^W CT 2u424 ^F = aa II 4 9-+ ++ 09 H "1 bnan `V = O r M £ = cl V SJEA DAOU "V = O £I A s-t + oz H NYOS VASVINDI "p = A ç M LaK + LI V SSIOg 142122 siqpiy x: aa [4 A^ t^c = SI V Inyos pupijntj] `V = O [4 Ə t^c = €I V qosng (PIM xo ueudois) vjnund sisdopiqpay = 99) [4 e, £-I t Ol V ‘uyjoy snur y a Ə [4 Ə r + 8 V TILED 19 'sstog tunuəoSSsptupp wnssá]y = Ə [4 dd £ = cI V Ker) und4p2042j21 `V = ada [4 dd £ E ZI V "yosipay `g (IOS 19 sxueg) wnauavo vwuauorjar ssourqojeq oouep (ww) 1O[OO KSo[ousud 95uo2 (wd) UNO. sor»odg -unqy yuy] — JaMOLY Jo9MO[J] -saqng 14310 ymo ANLA Wd “UOWIWIOD AIDA = OO :uouliuloo Á[Te90[ = QT ‘UOWWIOS = O ‘orel = Y ‘orel ÁIIA = YY :S92uanmooo €-I = O :aouepunge əanepy ‘yuld sed = qO ‘yurd = dd ‘Adnd = q YSA = 4 WL = O MoA Jed = AO ‘mood = A :9iuM^ = M H009 ISA0[4 '(919 *qo4 = z *uef = I) siSAo] sarooads e YOTYM ur syJUOW = AZO[OusYyd 1940[4 'ə1Kudəgureuo = HO Áydəeweyo pue aAydoydAnrway u33419q = NHO *eYAuqdoijdÁdorursu = H '[enuue = y WIO} YIMOID JOLIS] JO e1op ULIULLIYIPAN aui ur soroods 13jronao JO sojnquaj? [eorSo[ooo pu? [eordoloudIo]Nq `I 1189Y L [Vor. 74 Q AMOUMOUMOOUMOHOYL Q ANNALS OF THE MISSOURI BOTANICAL GARDEN 6326 zz 3 Ve UR O mm hm > Z Z Z n. Z n. Z n. R > > Z > OÓ ra! Z, se qeqeeeretCdCort << etree << << < <<< > Sisymbrium damascenum Boiss. et Gaill. S. officinale (L.) Scop. arnn >Z z z Thlaspi arvense L. Turritis laxa (Sibth. et Smith) Hayek nials also have longer average durations of flow- ering than annuals, except in the Solanaceae. INTRAFAMILY COMPARISONS Tables 1 and 2 show that most crucifer species In Israel, as in other areas, are annuals. As men- tioned above, the few perennial species in both the Mediterranean and desert floras generally flowerlaterandlongerthantheannuals,although exceptions do exist (e.g., Fibigia clypeata L.). Ad- ditionally, a distinct guild can be recognized by small plants (5-15 cm), with small, generally white or cream-colored flowers that appear early in the year (mid January through February). Within the two floral associations, we conducted all statistical analyses on crucifers four times: all species, annuals only, pygmy guild excluded, and perennials and pygmies excluded For the Mediterranean crucifers, patch-form- ing S species (Table formers generally paralleled our rank of patch formation (+++ > ++ > + > —,cases 3 and 4,++>+++ > + > —, cases | and 2, Table 4). In three of four comparisons, patch formers also had significantly larger petals than did non- patch formers (Table 4), with patch rank gen- erally paralleling petal size (++ > +++ > —, all cases). Few systematic differences exist among phenologies; patch formers gener- ally do not flower earlier or longer than non- patch formers, although there are a few excep- tions. There is no dominant flower color among Mediterranean patch- forming species; oe white, cream, and p comm Pubescence is significantly associated with non- patch-forming species in all analyses. Unlike Mediterranean species, the desert patch- forming species are neither taller nor have larger durations of 4 generally do not differ be- tween the grou Patch sans in the desert, with the exception of pygmy species, generally have purple flowers (8 of 12 species). Once again, pubescence is sig- nificantly ree aa with non-patch-forming species (Table 4). Horovitz & Cohen (1972) analyzed ultraviolet [VoL. 74 4 T T x < LI d b-t = o£ v ZINYIE `O (191no 19 `sstog) pipapj2 pluponaəphnəsq + 99) [4 9 £€-I + ç V PIEJ (oS) 244021Q $0422010N +++ OO t A rc = SI V `sstog] D21qD4D Si$dO14n1$DN = aa cl d r = 09 HO 'Sstog ('sstog) DIIDUIS Jy F 39 8I d cI-I = ct HO oneneg "5 19 pug&un(T "V `d (AIA) Suəllu plpup3li0 w = WI r AA OI-Z +++ Oc HO `sstog puo]fiuupd "py = TI 8 AA Or-c +++ tc HO `SSIOg SUJISAUDI 7111240]A E ad 8 d r€ ++ LI V ‘Ig “Y (90qsnouyos) paojfiaapd Jy = - 9) ZI 4 t-z ++ £€ V ‘OC CPA) vp] Jv ^ Tet OT €I d p= ++ oe V `sstog p.ədsp `w x = AA cl d $-t ++ Or H “SSIOG DIIGDID DjIOIYIID 5 — t:l 6 d t-t — L V ZINYIS :O (PA) vanwusdd vison W z, = u 8 d t-z + 9I V 1g ^W (TD) Puroufp pnujoo]pppy - _ u r ^ £l * SI V "yog 19 qqƏAA CAIA) p54(q1] 77 = = OT r AA t-I = ZI V Io[qosn]jN ('sstog) Dorqpap DIADjNGOT 5 ++ D L dd £-I - t V OG CPIILA) wmiofinf unajojidaT PA _ da E A rs = 0c V SSIOg Xo ABD pdav20421tu SIDS] z = 99) cI 4 II-I + SE HƏ enn] 22D1dÁ32D pt1əsap T S ++ a II d r-t = ct V ‘JUIBMYIS 19 "uosy ('sstog) pipoun ^q 2 k 90 01 d — - sE M 5507) DUD240Q DUDINAT < m Ə 9 dd T + SI d "SSIOG X9 “JUIOMYOS = 19 "Qosy (po3uouds) wnonydd3an wniqowasyq E Ë OT cI . t-I + oE V 'sstog 57tppna3unajs sndavooau14pu] o = 99 6 A 9-7 + SE V 'sstog ([exss1oA) Dru `Q 3 + 2 OI d La: = ce V 'sstog ([Pxss10) $242» sixpjojdiq z = O 8 2) p + LE V Jsoq snipouniw sndav20]p407) Z = JI 6 Ə r-t ++ SI V ‘Od CT pnuup 421421407) +E ud € ^ t-t + ¢ V U] 02uunu204214 7210D1$Du Y = Ə [4 Ə t-c + OI V "uno (T) Muru y = a T 2 t-r + Ol V “SSIOG XƏ |Əpnə1S wnjpuis4pu `V = a [4 AA £-I = OI V PIIM xe ueudoig wunijofiui `V = da t Ə + + OI V ‘PIM x? ueudoigs wundarotspp wnss4]p ssoulqojed oouep (ww) IO[OD K8o[ouaud Ə9ouəosəqnqdq (wd) W404 so1seds -unqy yWwuey — 19^0pLI I9^O[l 1431H ymo IANLPYA əd ure 588 *suongrA31qqe 10j | 9[QE.] JO PUIA IF ‘[OBIS] JO LIOY HSP ay} UT səroəds IIIN JO soINQUIIe [eoISojoIe pue [eordo[oudiojq ‘Z «ATAV_L 1987] AUERBACH & SHMIDA—ISRAELI CRUCIFER PATCH FORMATION 589 characteristics of the flowers of 22 species of cru- cifers native to Israel (17 Mediterranean, 5 des- Š " ert). Among these, high ultraviolet reflectance a H sÉ : H i p and/or patterning Is positively associated with = + patch formation (G,,; = 12.3, P < 0.001). The * same is true for the subset of Mediterranean °, species (Gaa; = 6.59, P < 0.025), but not the 8B Š ou e desert ones (G,4; = 0.5, P > 0.05). Oz ZlpOARonY = DISCUSSION E SE ——— WHY NOT FORM PATCHES? x E Several lines of evidence suggest that patch formation should increase the susceptibility of a Š 5 plant species to insect attack. Many herbivores, e Ç: Qm > > Z n. particularly those with narrow host ranges, are more likely to find, to remain on, and to repro- uce on hosts that are concentrated in space (Root, 1973; Risch, 1981). This is one reason pod ood (d why monocultures of herbaceous plants fre- quently support greater densities of insect pests than do polycultures (Pimentel, 1961; Root, 1973; Feeny, 1976; Altieri et al., 1977; Risch, 1981). The diversity and abundance of phytophagous ll l gr + insects attacking a plant species in a given area are also influenced by the botanical diversity of the area (Strong et al., 1984). Members ofa mixed- species plant assemblage often escape attack be- CQ C QN 6 ae cause increased species diversity reduces the sus- ceptibility of each plant species to discovery. This phenomenon has been referred to as “‘associa- tional resistance" (Tahvanainen & Root, 1972) < < < < < pn. or “plant defense guilds” (Atsatt & O'Dowd, 1976). Considerations of alternative resource availability for phytophagous insects as well as their frequent use of visual or chemosensory cues in host-plant location suggest that this form of escape should increase in effectiveness as phy- tophage dietary specialization increases (Root, 1973; Atsatt & O'Dowd, 1976). Rhoades & Cates’s (1976) and Feeny’s (1976) general theories of plant defensive chemistry ex- pand the concept of plant escape from enemies in ecological time into evolutionary arguments concerning selection for different classes of al- lelochemicals. Both postulate that as the occur- rence of a plant species or tissue becomes more predictable (apparent) in space and time there should be greater selection for generalized, quan- titative (dosage-dependent) defenses as opposed to specialized, qualitative defenses (toxins). Quantitative defenses are presumed to be more metabolically costly but harder for a herbivore to circumvent than qualitative ones. Thus, it is Phenology 4 4 4 5 4 9 Pubescence Mean Height (cm) 3 0 5 0 5 5 Growth Form Species TABLE 2. Continued. Reboudia pinnata (Viv.) O. Schulz Savignya parviflora (Del.) Webb Schimpera arabica Hochst. et Steud. ex Boiss. Sisymbrium erysimoides Desf. Zilla spinosa (L.) Prantl 590 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 BLE 3. Comparison of phenologies of - Brassicaceae with other families in the Mediterranean flora of Feb. = 2, etc. ae] Months were converted to integers (Jan. = 1, or flowering range and peak analyses. Inequality signs indicate earlier (<) and later ue in the growing season. Compared with Brassicaceae Flowering Flowering Flowering Range Peak Duration Flowering Flowering Number of (median (median (mean number Range Peak Family Species month) month) months (s.d.)) (distribution) (distribution) Brassicaceae 76 3.71 3.61 2.94 (1.47) Annual 64 3.46 3.52 2.88 (1.20) Perennial 12 5.50 4.75 -— Lamiaceae 80 5.79 5.27 3.71 (1.49)* >+ >+ nnual 17 4.86 4.42 3.35 (1.41) >++ >++ Perennial 63 6.07 5.50 — Asteraceae 169 5.74 4.88 3.43 (1.74)* >+ >+ Annual 105 5.17 4.54 3.25 (1.69) >++ >++ Perennial 64 6.63 5.83 — — — Liliaceae 66 4.33 3.66 2.91 (1.34) >+ = Perennial 65 4.33 3.67 — — — Solanaceae 15 6.96 6.50 5.13 (2.64)* >+ >+ 6 7.60 7.00 5.33 (0.82)** >++ >++ Perennial 9 6.50 6.17 — — Poaceae 168 5.67 4.86 3.77 (1.95)* >+ >+ Annual 111 5.18 4.59 3.40 (1.61)** >++ >++ Perennial 57 6.65 5.68 — — Papilionaceae 206 4.52 4.19 3.13 (1.10) >+ >+ Annual 173 4.33 4.08 2.92 (0.67) >++ >++ Perennial 33 5.70 5.21 — Apiaceae 87 5.35 4.83 3.03 (1.24) >+ >+ Annua 48 4.93 4.55 2.77 (1.10) >++ >++ Perennial 39 5.79 5.39 — * = * Significantly different from all, or annual (**) crucifer species (t-test, Welsh approximation, P < 0.05). + = Significantly different from all, or annual (+ +) crucifer species (Kolmogorov-Smirnov 2 sample test, P « 0.05). argued that a plant that is apparent by virtue of its growth form, abundance, and longevity, such as a common tree species, must invest a consid- erable amount of energy into quantitative defen- ses, whereas an annual herbaceous plant can in- vest less energy in qualitative defenses, provided it also remains unapparent in time and/or space. By definition, unapparent plants should have small patch sizes and low p density (Fox, 1981 Escape in time and space should be effective against specialized herbivores in ecological time and against nonspecialized herbivores over evo- lutionary time. Specialized herbivores that have evolved a means of circumventing an allelo- chemic defense frequently respond to the allelo- es for host location and phagostimulation; at herbivores have, over evolutionary time, turned an unapparent re- source into an apparent one (see Courtney, 1985, for various interpretations of apparency). Once this occurs the plant benefits from anything that reduces its apparency to adapted enemies, such as having an irregular phenology or growing in multispecies assemblages. Against nonspecial- ized herbivores, escape in space and/or time re- duces the frequency of encounters between po- tential enemy and plant tissue, thereby decreasing the likelihood of an herbivore evolving a detox- ification mechanism. Since most crucifers are short-lived herbs and contain qualitative allelochemicals, one might expect them to rely quite heavily on escape in time and space from phytophagous insects. In fact, Feeny ar regarded crucifers as typical unapparent plants and much of his theory of plant apparency is based on crucifers (see ref- erences in Feeny, 1976). Most Israeli crucifers 1987] AUERBACH & SHMIDA—ISRAELI CRUCIFER PATCH FORMATION do rely on escape on one time scale by nature of their annual growth form with concomitant short- term availability each year of leaves. However, the propensity of many species to form large monotypic patches confers a high degree of spa- tial predictability and resource concentration on these species. In addition, location of patches of Mediterranean patch-forming species frequently does not change from year to year. This situation would appear to be disadvantageous in light of the theory discussed above. HOW DO THEY GET AWAY WITH IT? Mediterranean patch-forming crucifers in Is- rael suffer little herbivore damage despite their short-term spatial predictability. During a season of sampling (1983), we never observed high den- sities or diversities of phytophages on patch- forming species. Neither high numbers of sup- ported insect herbivores nor large amounts of leaf area removal were detected with spot cen- susing during ten years of vegetation sampling. We suggest that Mediterranean patch-forming crucifers are able to grow and persist in dense monospecific associations owing to possession of glucosinolates (and other allelochemicals in some species), and because they flower earlier than oth- er local species. Both displaced phenologies and glucosinolates prevent seasonal tracking and use of crucifers over evolutionary time by presently non-crucifer-adapted herbivores. By growing earlier than most other annual species, these cru- cifers have reduced the number of chance en- dietary shifts in phytophages. If patch formation occurred later in the growing season, more en- counters between pest and potential host would ensue, and over evolutionary time one would expect an increase in diversity of insect species adapting to patch-forming crucifer species. Many crucifers germinate, grow, and flower before the annual peak of insect biomass. Indeed, many of the patch-forming species flower before the annual appearance of crucifer-adapted her- mida, unpubl. data). Occasionally, these adapted herbivores do appear on patch formers during the late stages of flowering or seed set, Groups NP > P** NP > P* NP > P* Pubescence Start of Flowering P < NP* P < NP** NP NP NP Flowering Range (distribution) P=N P= P= P=N P=N P=N Flowering Peak (mean) P=N P > NP* P P = NP P = NP P=N P=N = NP -forming (P) and non-patch-forming (NP) crucifer species in the (A) Medi- P < NP P=N = N P = (B) DESERT P=N P=N P=N (A) MEDITERRANEAN (distribution) Duration P > NP* P P=N NP P=N P=N P=N pygmy guild excluded; (4) perennials and pygmies excluded. P s Petal Size P > NP** P > NP*** N P > NP* P=N P=N P=N P=N Plant Height P > NP*** P > NP*** P=N TABLE 4. Comparisons of morphological and phenological attributes of patch terranean and (B) desert floras of Israel. In reference to phenological traits, the inequality signs indicate earlier (<) and later (>) in the growing season. are defined as: (1) = all species; (2) = annuals; (3) * = Significant difference, P < 0.05; ** = P < 0.01; *** = P < 0.001. Group 592 when herbivory has a minimal effect on the fit- ness of n Germination pane in the desert is far less ‘redictable than in Mediterranean regions be- cause of var A eae a early-spring rains (Sbmida et al., 1985). Thus, desert crucifers are far more unpredictable in time than their Med- iterranean counterparts. Year-to-year variation in composition, location, and size of patches is often tremendous. Despite these differences, flowering is still well under way by the time most herbivores, even crucifer specialists, arrive. In- terestingly, pubescence, a characteristic fre- quently interpreted as a possible herbivore de- fense (Coley, 1983) as well as an adaptation that reduces rates of evapotranspiration (Lieberman & Lieberman, 1984), is more common among non-patch- than patch-forming species in both floristic regions. e can only speculate why crucifer-adapted herbivores have not undergone phenological dis- placement to exploit early-season crucifers. Se- m for Synenrony may li countered by phys- early-season cold temperatures. The exsul: variable phe- aAlonoical ° ony. As Feeny (1977) noted, *Short growth season, shifting pattern of geo- graphic distribution, and association with harsh and somewhat unpredictable climatic conditions are all characteristics which are likely to favor escape by plants from their adapted enemies"; this has been echoed elsewhere (Janzen, 1970; Rhoades & Cates, 1976; Feeny, 1976). In addi- tion, selective pressure for phenological changes among Mediterranean crucifer-feeding insects may not be very great, since cruciferous hosts are available later in the season. There also may e phytochemical barriers that function with phenology in much the same manner as we en- visage for nonadapted herbivores. Many cruci- fers contain diverse glucosinolates as well as oth- er allelochemicals, and the performance of crucifer feeders frequently varies greatly among host plants (Root & Olson, 1969; Feeny, 1976; Rodman & Chew, 1980). HOW ARE PATCHES FORMED? The ability to form patches clearly involves more than early phenologies, since we found few significant phenological differences between editerranean patch- and non-patch-forming crucifers (Table 4). In Mediterranean and desert ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 regions of Israel, late-winter rains initiate ger- mination of seeds of crucifers and other annuals. By early spring, patches of various crucifers dom- inate the landscape of much of Israel (Shmida & Auerbach, 1983). Bell & Muller (1973) described similar patch formation by an introduced cru- cifer, Brassica nigra, in some annual California grasslands. There, B. nigra produces monospe- cific patches throughout the grasslands despite synchronous germination of all annual species following late-autumn rains. Bell & Muller (1973) found that patch formation by B. nigra ips production of potent allelopathic chemicals. do not know if allelopathy is important in nth formation by Israeli crucifers. Growth rate ap- pears to have a role in patch formation. Many crucifers in Israel and elsewhere (Feeny, 1977) mature and set seed extremely rapidly, a trait associated with their occurrence in environmen- tally harsh or disturbed sites. In addition, patch formation may involve differential responses to microsite variability in edaphic parameters and grazing pressure. WHY FORM PATCHES? Patch formation may be reinforced by com- petition for pollinators and increased pollination ficiency. Most crucifers, including Israeli pecies, are either facultative or obligate out- crossers (Fryxell, 1957; Crisp, 1976; D. Zohary, pers. comm.). Seasonally advanced flowering among Mediterranean crucifers may reduce in- terspecific competition for pollinators, provided flowering does not occur before pollinators are available. Patch formation among these species may also increase pollination success over that of solitary individuals if pollinators are limited. Thus, patch formation could be reinforced by pollination success. While patch formation may concentrate re- sources and produce attractive displays for pol- linators (Handel, 1983; Rathcke, 1983), intra- specific competition for pollinators among members of a patch may increase if patch size becomes sufficiently large. For example, Rathcke's (1983) density-visitation model en- visages a facilitation in pollination as patch size increases up to the point where pollinators are V O advanced phenologies result in flowering before the annual peak in pollinator availability 1987] (Schemske et al., 1978). As intraspecific com- petition intensifies, selection for enhanced at- tractiveness may result in differences between patch- and non-patch-forming species in polli- nation-associated traits. In our analyses, two morphological attributes frequently associated with pollination, plant height and petal size (Faegri & van der Pijl, 1979; Waddington, 1979; Waser, 1983a), did vary be- tween non-patch- and patch-forming species. Among Mediterranean species, patch formers are taller and they generally have larger petals than do non-patch-forming species. Although we do not know the intensity of intraspecific competi- tion for pollinators in patches, increases in height and petal size were also positively associated with our measure of patch size. In addition, plant height, petal size, and indices of patch size were positively correlated with propensity for out- crossing as measured by pollen/ovule ratios (Ku- nin, unpubl. ms.). Thus, taller patch formers ap- pear to have a greater tendency towards outcrossing than do smaller patch formers, and species that do not form patches have the lowest tendency to outcross. That high levels of ultra- violet reflectance and patterning is positively correlated with patch formation found among Mediterranean species also suggests that patch formation may result in increased intraspecific competition for pollinators. Differences in height and petal size do not oc- cur among desert species; however, desert patch- forming species do begin flowering before non- patch formers, a difference observed among Mediterranean species only when the early-flow- ering pygmy guild is excluded from analyses. Ad- vanced flowering among the desert patch formers may reduce interspecific competition for polli- nators (Waser, 1983b) and possibly permit pol- linator specialization. Some pollinators in other deserts are capable of synchronizing with the ephemeral phenologies of their hosts (Baker & Hurd, 1968; Crawford, 1981). With the excep- tion of the pygmy guild, a high proportion of desert patch formers have purple flowers (67% patch vs. 27% nonpatch), which suggests pos- sible pollinator specialization or segregation be- tween patch- and non-patch-forming species. In summary, we suggest that growing in patches aids in pollinator attraction, especially in con- junction with atypical phenologies. Displaced phenologies also may function to segregate pol- linator and herbivore guilds temporally. If pol- linator efficiency is high and seed dispersal low, AUERBACH & SHMIDA—ISRAELI CRUCIFER PATCH FORMATION 393 patch formation may be self-perpetuating, be- cause of concentration of seeds into localized seed banks LITERATURE CITED ALTIERI, M., A. VON SCHOONHOVEN & J. DOLL. The — role of weeds in insect 1977. agement systems: a review illustrated by bean (Phaseolus vulgaris) cropping systems. Pest Arti- cles pn . 23: 195-205. ATSATT, P. D. O'Dowp. 1976. Plant defense guilds. "uic 193: 24-29. — H. G. & P. D. Hurd. 1968. Intrafloral ecol- y. Ann. Rev. Entomol. 13: 385-414. Bar. D. T. & C. H. MuLLER. 1973. Dominance the California annual p by Brassica uds Amer. Midl. Naturalist 90: 277-299. Co_ey, P. D. Herbivory and defensive char- acteristics of tree species in a lowland gd for- est. Ecol. Monogr. 3: 209-233. Conover, W. J. 1980. Practical Nonparametric Sta- tistics. John Wiley, New York. CourTney, S. P. 1985. Apparency in coevolving re- lationships. Oikos 44: 91-98. CRAWFORD, C. S. 1981. Biology of Desert Inverte- brates. Springer-Verlag, Berlin. Crisp, P.C. 1976. Trends in breeding and cultivation of cruciferous crops. Pp. 69-118 in J. G. Vaughan, A. J. MacLeod & B. M. G. Jones (editors), The Biology and Chemistry of the Cruciferae. Academ- ic Press, New York. FAEGRI, K. & L. VAN DER Pur. 1979. The Principles of Pa qur Ecology. Pergamon Press, Oxford. 19 Plant apparency and chemical S 1-40 in J. W. Wallace & R. L. Mansell (editors), Biochemical Interactions Between AA and Insects. Recent Adv. Phytochem., Volume Press, New York. Defensive ecology of the Cruciferae. Ann. Missouri Bot. Gard. 64: 221-234. Defense and dynamics in plant- 853-864 FURTH, D. G. 1979. Zoogeogra and host plant ecology of the Alticinae =p p especially Phyl- lotreta; with descriptions of three new species (Co- leoptera: Chrysomelidae). Israel J. Zool. 28: 1-37. HANDEL, S. N. 83. Po quen eR. plant pop- ulation structure Pp. 163-211 in and w. L. Real (editor), En Biology. yee ere Press, Orlando, Florida. HEDGE, I. C. 197 6. A systematic and o survey of the Old . Cruciferae. Pp. 1-45 ; G. Vaughan, A. J. M od M. G. Jana (editors), The Biology ate Chemistry ae the Cru- ciferae. Academic Press, New York. Horovitz, A. & Y. COHEN. 1972. Ultraviolet reflec- tance characteristics in flowers of crucifers. Amer. J. Bot. 59: 706-713. JANZEN, D. H. 1970. Herbivores and the number of tree species in tropical forests. Amer. Naturalist 104: 501-528. KJAER, A. 1976. Glucosinolates in the Cruciferae. Pp. 594 207-219 in J. G. Vaughan, A. J. MacLeod & B. M. G. Jones (editors), The Biology and Chemistry of the Cruciferae. Academic Press, New York. LIEBERMAN, D. & M. LIEBERMAN. 1984. The causes a dry tropical forest. Biotropica 16: 193-201. PIMENTEL, D. 1961. The influence of plant pm patterns on insect populations. Ann. Entomol. Soc Amer. 54: 61—69. RATHCKE, B. 1983. Competition and irpoc among plants for pollination. Pp. 305-329 i Real (editor), Pollination Biology. Academic ES Orlando, Florida. REMINGTON, R. & M. ScHoRK. 1970. Statistics with Application to the aea and Health Sciences. Prentice- Pup ia Jers D. RHOADES, e 1976. Toward a general es of plant antiherbivore chemistry. Pp. 168-213 in J. . Wa llace & R. L. i Rude (editors) BL ca Pla and Insects. Recent Adv. Phytochem., Volume 10. ; l Inse t herbivore abundance in tropical scales and polycultures: an exper- imental test of two hypotheses. Ecology 62: 1325- RODMA N, J. & F. d nie Phytochemical cor- sens of herbivo ommunity of native and naturalized Crucifome, onem Syst. Ecol. 8: 43- Roor, R. B. 4973. Organization ofa plant-arthropod Ne fau- na of collards (Brassica oleracea). Ecol. Monogr. 43: 95-124. OLSON. 1969. Population increase of the cabbage aphid, Brevicoryne brassicae, on different host plants. Canad. Entomol. 101: 768- 773. SCHEMSKE, D., M. WILLSON, M. MELAMPY, L. MILLER, L. VERNER, K. SCHEMSKE & L. BEST. 1978. Flow- ering ecology of some spring woodland herbs. Ecology 59: 351-366. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 SHMIDA, A. & M. AUERBACH. 1983. The strange mus- tard smell of the crucifers. Israel— Land and Na- ture 9: 61-66. , M. EvENARI & I. Nov-MEIR. 1985. Hot desert ecosystems: an integrated view. /n M. Evenari & I. Noy-Meir (editors), Hot Desert Ecosystems of the World. psi New York. SLANSKY, F. & P 1977. Stabilization of ied rate of nitrogen is by larvae of the c bage butterfly on wild and cultivated food ade. 9-22 D. R., J. H. LAwTON & T. R. E. SOUTHWOOD. 984. In sects on Plants. Blackwell Scientific Pub- lications, Oxford. mE N, J. O. & R. B. Roor. 1972. The influ- ence of vegetational diversity on the population eee of a specialized herbivore, Phyllotreta cru- ciferae (Coleoptera: Chrysomelidae). Oecologia 10: VAUGHAN, J. G., A. J. MACLEoD & B. M. G. JONES 1976. The Biology and Chemistry of the Cruciferae. Academic Press, New York. NGT . 1979. Divergence in inflores- cence height: an evolutionary response to polli- nator fidelity. Oecologia 40: 43-50. Waser, N. M. The adaptive nature of floral traits: ideas and evidence. Pp. 242-285 in L. Real (editor), Pollination Biology. Academic Press, Or- lando i . Competition for pollination and floral character differences among n ; E. Jones & R. J. Little (editors), Handbook of Ex- perimental Pollination Biology. Scientific and Ac- ademic rr We New York. ZOHARY, M. 19 Geobotanical Foundations of the Middle Ei 2 volumes. Gustav Fischer Verlag, Stuttgart. RUN-DOTHA 1966. Flora Pa- laestina. ‘Israel don of : Sciences, Jerusalem, Israel. FLAVONOID SYSTEMATICS OF SEVEN SECTIONS OF LUDWIGIA (ONAGRACEAE)! JOHN E. AvERETT,? PETER H. RAVEN,? AND ELSA ZARDINP ABSTRACT Da ita are presented for the flavonoids of 24 species vm seven sections of Ludwigia, all of which sections reported suggests that these sections are advanced in this fe al of eight flavonoids, three glycoflavones, ature, as does a EUM reduction of structural types and a reduction in the number of glycosidic substitutions in them As part of a comprehensive study of the fla- vonoids of Onagraceae, we are herein reporting results from seven sections of Ludwigia. As ex- plained in earlier papers (Averett et al., 1978, 1979), the objectives of the overall study on On- agraceae are to provide an analysis of the fla- vonoids at the generic level for the entire family and to gain insight into the evolution of flavo- noid compounds by correlating substitutional and structural changes with a phylogeny based on other systematic data. This is the first of several papers in which we shall present the results of flavonoid analyses of Ludwigia. The only pre- vious report on flavonoids in Ludwigia is a brief summary of data for the whole genus (including that reported here) in Averett & Raven (1984). udwigia is the only genus of the tribe Jus- siaeeae and comprises approximately 82 species found in wet habitats in both temperate and trop- ical regions worldwide (Raven, 1963). Ludwigia appears to represent a branch of the family dis- tinct from all other members (Eyde, 1977, 1978, 1981; Raven & Tai, 1979). It is therefore of par- ticular interest to consider the evolution of fea- tures in this isolated evolutionary line. This report deals with the species included in the original broad circumscription by Munz (1942; see also Raven, 1963) of sect. Myrtocar- pus, a group of some 23 species centered in trop- ical and subtropical South America that ap- peared to be “phylogenetically central" in the genus (Ravon, 1963). Based on his studies of the and , cyto logy of this complex, Ramamoorthy (1979) divided sect. Myrtocarpus into seven sections, com- menting that the species had been grouped pri- marily on the basis of shared primitive charac- ters. Subsequent work has suggested that his sect. Michelia is not distinct from sect. Myrtocarpus sensu stricto, and that Ludwigia mexiae (Munz) Hara is sufficiently distinct from other members of sect. Pterocaulon that it is best treated as the monotypic sect. Cinerascentes (Ramamoorthy & Zardini, 1987). Thus we now recognize seven sections in this group, delimited as follows: sect. ocarpus with 20 species (14 examined herein), including some with the most primitive assemblages of characters in the genus; sect. Pterocaulon with five species that comprise a well- ve monotypic Amazonia, Heterophylla, Tectiflora, Humboldt- ia, and Cinerascentes—that are each specialized relative to sect. Myrtocarpus (Ramamoorthy & Zardini, 1987). MATERIALS AND METHODS Dried leaf materials from 24 species of seven sections of Ludwigia were examined for flavo- noids. Approximately 120 populations in total ! We gratefully acknowledge support from the U.S. National Science Foundation through individual grants to Averett and Raven. Plant material was received from . P. Ramamoorthy to whom we are especially indebted for collecting in Brazil most : the species analyzed for this study. W. D. Stevens also collected material, and his assistance is greatly appre ment of Biolo 3 Missouri Botanical Ga , P.O. Box ANN. MIssouRI Bor. GARD. 74: 595-599. 1987. ate ogy, aes of Missouri, St. Louis, Missouri 63121, U.S.A. rden 299, St. Louis, Missouri 63166, U.S.A. 596 and as many as 25 populations of some of the more variable species were sampled. The sam- ples included most chromosomal races of the species concerned. Voucher specimens are listed in el. The leaf material was extracted overnight in 85% methanol and the resulting extract was ex- amined by two-dimensional paper chromatog- raphy. Certain of the extracts were analyzed us- ing TLC (polyamide and cellulose) as ~~ In some cases, the flavonoids were crudely rated on Sephadex LH 20 witha iese hana system as described by Hiermann et al. (1978). For structural elucidation, replicate chromato- grams were run and the isolated compounds cut from the paper for further purification and anal- ysis. The quantity of leaf material varied ac- cording to usage, but approximate amounts were 0.5-1.0 g for general screening, 5-10 g for rep- licate chromatograms, and 20-30 g for column hromatography. Identifications of the glyco- sides, their aglycones, and sugars were made as previously described (Averett et al., 1978, 1979) and were compared with standard Rf values and absorption maxima (Averett, 1977). In addition, most of the aglycones and sugars were run, along with authentic reference compounds, by circular thin-layer chromatography as described by Ex- ner et al. (1977). Base hydrolysis was employed to determine acylation but the unction was not determined. RESULTS AND DISCUSSION Eight flavonoids were found among the species sampled (Table 2): orientin (1), isoorientin (2), orientin-O-acylate (3), quercetin 3-O-rhamno- side (4), quercetin 3-O-arabinoside (5), quercetin 3-O-glucoside (6), quercetin 3-O-diglucoside (7), and quercetin 3-O-rutinoside (8). Compounds 1- 3 are glycoflavones and compounds 4-8 are fla- vonol glycosides. All of the compounds are based on structures having two hydroxyl substituents in the B-ring. Some infraspecific variation was found, es- pecially in such variable species as Ludwigia pe- ruviana, L. elegans, and L. laruotteana. Al- though variation was present, no flavonoid unique to any species was found in a single population. We also detected differences in concentrations of compounds between populations of some species, but made no attempt to document these differ- ences. Interspecific variation within the larger sections is apparent but, like populational vari- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 ation, is largely confined to the presence or ab- sence of certain compounds. The more interest- ing variation is found in intersectional comparisons. Because of this, samples are ar- ranged by species and grouped in their respective sections in Table 2 Compound 3, the acylated glycoflavone, is found in only five species in three different sec- tions, including the closely related sections Cine- rascentes and Pterocaulon. Compounds 5 and 7 are the next most frequently absent, occurring in only six and seven species, respectively, and typ- ically in low concentration when present. Com- pounds 1, 2, and 8 are the least variable and are found in 18, 20, and 19 of the species, respec- tively. Compound 2 is present in all species in which glycoflavones occur, and compound 8 is present in all species in which flavonols are found. Compounds 4 and 6 occur in 13 and 14 of the species sampled. Glycoflavones are found in 20 of the 24 species, flavonols in 19, both glycofla- vones and flavonols in 15, only glycoflavones in five, and only flavonols in four All ofthe species have at least two compounds and none has more than seven. The average number of compounds per species is 5.37 for the entire group. The comparison of numbers of compounds at the sectional level is especially interesting. Species of sect. Myrtocarpus, which has relatively generalized features, and sect. (Piero aalon. 3.0 (Humboldtia and Heterophyl- la), and 2.0 (Amazonia and Tectiflora). Flavonols are present throughout Onagraceae and glycoflavones are present in all tribes except Fuchsieae and Epilobieae (Averett & Raven, 1984). Glycoflavones are especially well repre- sented in Circaea (Boufford et al., 1978; Averett & Boufford, 1985). The presence of glycoflavones is considered primitive relative to the presence of other groups of flavonoids (Harborne, 1977), and the distribution of these compounds within Onagraceae does not contradict that contention. Except for Ludwigia sericea, which has only flavonols, all species examined of sect. Myrto- carpus and the single species, L. mexiae, of sect. Cinerascentes have both glycoflavones and fla- onols. The remaining monotypic sections ex- hibit either glycoflavones only—sections Heter- ophylla and Amazonia—or flavonols only— sections Humboldtia and Tectiflora. Collective- ly, the species of sect. Pterocaulon exhibit both ABLE 1. Voucher specimens of Ludwigia used for flavonoid analysis in this study. Specimens of all material deposited at MO, unless otherwise indicated. Ludwigia section Amazonia Ramamoorthy Ludwigia densiflora (Micheli) Hara. BRAZIL. RONDONIA: Duarte 7329 (MO, RB). Ludwigia section Cinerascentes Ramamoorthy & Zardini Ludwigia mexiae (Munz) Hara. BRAZIL. PARA: Ramamoorthy 652. Ludwigia section Heterophylla Ramamoorth Ludwigia inclinata (L. f.) Gómez. MEXICO. OAXACA: Breedlove & Raven 13686. CosrA RICA. PUNTARENAS: Stork & Horton 8912 (US). BRAZIL. AMAPA: Froes & Black 27732 (IAN). Ludwigia section Humboldtia Ludwigia sedoides (H. & B.) Hara. PANAMA. CANAL ZONE: D'Arcy 12350. BRAZIL. PARA: Archer 8411 (RSA). Ludwigia section Myrtocarpus (Munz) Hara Ludwigia albiflora Ramamoorthy. BRAZIL. GoIAS: Ramamoorthy 545. MINAS GERAIS: Ramamoorthy 427 MO, SP). me d (Hassler) Hara. BRAZIL. MATO GROSSO Do suL: Ramamoorthy 610 (MO, SP), Ramamoor- thy & Ludwigia jou (Camb.) Hara. BRAZIL. Goras: Ramamoorthy 532, 560, 564. MINAS GERAIS: Ramamoorthy 403,4 4; Ramamoorthy & Vital 140; Ramamoorthy et al. 148, 153, 161, 168, 176, 178, 179, 181 (MO, SP), 184, 195, 299, 301, 308, 309, 319. RIO DE JANEIRO: Ramamoorthy et al. 291. SAO PAULO: Ramamoorthy 379, 384, 395; Ramamoorthy & Vital 112; Ramamoorthy et al. 196 Ludwigia hassleriana (Chodat) Hassler. BRAZIL. MATO GROSSE DO SUL: Ramamoorthy 629, Ramamoorthy et b Ludwigia irwinii Ramamoorthy. BRAZIL. MINAS GERAIS: Ramamoorthy et al. 142. SAO PAULO: Ramamoorthy 80; Munz 15406 (NY, POM, US). Ludwigia laruotteana (Camb.) Hara. BRAZIL. Goras: Ramamoorthy 419, 420. MINAS GERAIS: alga 101; Ramamoorthy & Vital 90; RE et al. 143, 147, 160, 172, 188, 311. SAO PAULO: Ram moorthy 69. Ludwigia martii (Micheli) Ramamoorthy. BRAZIL. MINAS GERAIS: G/aziou : ” (B, C, F, P, R). Ludwigia myrtifolia (Camb.) Hara. BRAZIL. MINAS GERAIS: Ramamoorthy Ludwigia nervosa (Poir.) Hara. M ZELAYA: maka 8275. onl BAHIA: ti eoa be et HR 328. DISTRITO FEDERAL: Ramamoorthy 526. goras: Ramamoorthy 561; Ramamoorthy & Vital 544 rosso: Ramamoorthy 571. MATO GROSSO DO SUL: Ramamoorthy 605, 607. MINAS GERAIS: cede et al. 170. sao PAULO: Ramamoorthy 393 (MO, SP); Ramamoorthy & Vital 78 (MO, SP). Ludwigia peruviana (L.) Hara. BRAZIL. MINAS GERAIS: Ramamoorthy 366. PARANA: Ramamoorthy 207, 275, 281. Ludwigia pseudo-narcissus (Chodat) Ramamoorthy. BRAZIL. PARANA: Ramamoorthy et al. 283. Ludwigia rigida (Miq.) jvc SURINAM: Pulle 475. VENEZUELA. COJEDES: Pittier 11711 (B, US, VEN). P sericea (Camb.) H RAZIL. MINAS GERAIS: Ramamoorthy 68; Ramamoorthy et al. 157, 158, s PARANA: Ramamoorthy et al. 215, 216, 288, 289. SANTA CATARINA: Ramamoorthy et al. 240. SAO PAULO: Ramamoorthy 44 Ludwigia pistas (Camb.) Hara. BRAZIL. BAHIA: Ramamoorthy et al. 336. DISTRITO FEDERAL: Ram moorthy 513; Ramamoorthy et al. 349, 351. Goras: Ramamoorthy et al. 342, 344, 345, 506. MATO Grosso: Ramamoorthy & Vital 579. MINAS GERAIS: Ramamoorthy 163, 164, 165, 405; Ramamoorthy et al. 187. Ludwigia section Pterocaulon Ramamoorth Ludwigia decurrens Walt. NICARAGUA. ZELAYA: Stevens 4916. BRAZIL. MINAS GERAIS: Ramamoorthy et al. 303, 306. SANTA CATARINA: Ramamoorthy et al. 258, 259. po erecta (L.) Hara. MEXICO. OAXACA: Breedlove & Raven 13669 (DS, MO). NICARAGUA. ZELAYA: Stevens 8274. CUBA. ORIENTE: Ekman 6537 (S). COLOMBIA. HUILA: Smith 1204 (GH, UC, US). hrai RHODESIA. NDANGA: Goodier 977. Ludwigia filiformis (Micheli) Ramamoorthy. BRAZIL. GOIAS: Ramamoorthy & Vital 555. SAO PAULO: Rama- moorthy 73: pu longifolia (DC.) Hara. BRAZIL. MINAS GERAIS: Ramamoorthy & Vital 96; Ramamoorthy et al. 150. A CATARINA: Ramamoorthy et al. 231, 233, 237. a major (Micheli) u n BRAZIL. RIO GRANDE DO SUL: Ramamoorthy et al. 245. Ludwigia section Tectiflora Ramamoorthy i latifolia (Benth.) Hara. NICARAGUA. RIO SAN JUAN: Neill 3361. GUYANA. WEST DEMARARA: Ma- e & Fanshave 22951 (NY, U, US). PERU. SAN MARTIN: Ferreyra 18506, Williams 7153 (F, US). 597 598 fA 13 A ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 f Ludwigia. + = flavonoid detected; dm flavonoid TABLE2. Distribution not detected. Key: 1 = orien n, 2 = isoori rientin, 3 — orientin-O-acylate, 4 = quercetin-3-O-rhamnoside, 5 = quercetin-3-O-arabinoside, P. — quercetin-3-O-glucoside, 7 — quercetin- -3- O-diglucoside, and 8 = pene nes oside. O-rutin Glycoflavones Flavonols l 2 3 4 5 6 7 8 Sect. Amazonia L. densiflora + + 0 0 0 0 0 0 Sect. Cinerascentes L. mexiae + + + 0 + 0 0 + Sect. Heterophylla L. inclinata + + + 0 0 0 0 0 Sect. Humboldtia L. sedoides 0 0 0 + 0 + 0 + Sect. Myrtocarpus L. albiflora + + 0 + 0 + 0 + L. bullata + + 0 + 0 + 0 + L. elegans + + 0 + + + + + L. hassleriana 0 + 0 + 0 0 + + L. irwinii + + 0 + + + + + L. laruotteana + + 0 + + E E ~ L. martii 0 + 0 0 0 0 + + L. myrtifolia + + 0 + 0 + 0 + L. nervosa + + 0 + 0 + 0 + L. peruviana + + 0 t + + + + L. pseudo-narcissus + + 0 0 0 0 0 + L. rigida + + 0 0 0 0 0 + L. sericea 0 0 0 + 0 + 0 + L. tomentosa + + 0 + + + 0 4 Sect. Pterocaulon L. decurrens + + 0 0 0 + + + L. + + + 0 0 0 0 0 L. filiformis 0 0 0 + 0 + 0 + L. longifolia + + + 0 0 0 0 0 L. major + + + 0 0 0 0 0 Sect. Tectiflora L. latifolia 0 0 0 0 0 + 0 + glycoflavones and flavonols, but only one species, L. decurrens, has both classes of compounds. One species, L. filiformis, has only flavonols, and the remaining three species have only glycoflavones. Thus, if sect. Pterocaulon is a monophyletic group derived from sect. Myrtocarpus, as is indicated from morphological studies, then within this sec- tion of five species one has lost the ability to produce glycoflavones and three to produce fla- vonols; that, at sd would be the most parsi- monious explanat The five denied sections we are consid- ering here do not appear to be more closely re- lated to sect. Myrtocarpus, on the basis of their overall characteristics, than they do to any other dwigia t and there is no evidence of a direct relationship between any two. The overall similarity of fla- vonoids between sections ee and Hum- boldtia and between sectio d Het- erophylla could not, en. be taken as an indication of relationship between those taxa. Rather, the similarity of flavonoids between these 599 AVERETT ET AL.—FLAVONOID 1987] groups seems to reflect parallel and independent loss of particular classes of flavonoids and/or in- dividual compounds, a trend that has character- ized the evolution of the genus overall. A further evaluation of their relationships, which must be multidimensional, would need to take into ac- count the remainder of the genus. It does appear, in terms of admittedly largely plesiomorphic characteristics, that sections Pterocaulon an Cinerascentes are more clearly related to sect. Myrtocarpus than are the others. n summary, our analysis has revealed a pat- tern of loss of individual flavonoids and groups of flavonoids in the seven sections of Ludwigia that we have considered in this paper. Further resolution of the relationships of these species must await more detailed study. LITERATURE CITED AVERETT, J. E. 1977. Absorption maxima and R values as an aid to the identification of selected flavonoids. Phytochem. Bull. 1 0-26. FFORD. 1985. The flavonoids and flavo noid ps of Circaea (Circaeeae, On- agracea ux Sy y pa 3-373. 984. Flavonoids of Ona- graceae. redu Missouri Bot. Gard. 71: , B. KERR & . RAVEN. 1978. Flavon oids of Onagraceae: ppm sect. Epilobium. Amer J. Bot. 65: IRAE: N & H. BECKER. 1979. Flavonoids i oe Epilobieae. Amer. J. Bot. 66: 1151- 155. TICS OF LUDWIGIA BourroRD, D. E., P. H. RAVEN & J. E. AVERETT. 197 Glycoflavones i x cuis (Onagraceae). Biochem. Syst. & Ecol. 6: 59-60. EXNE E Lei KER. 1977. Circular chroma a co hy- o lution in Ludwigia (Onagraceae). I. Androecium, placentation, merism. Ann. Missouri Bot. Gard. 64: 644-655 . 1978. Re Mdh pi structures and evolution in Ludwigia (Onagrace and seed. Ann . II. Fruit Missouri Bot. Gard. 65: 6—675. 1981. Reproductive structures and evolution in Ludwigia (Onagraceae). III. Vasculature, nec- taries, conclusions. Ann. Missouri Bot. Gard. 68: 379-412. HARBORNE, J. B. 1977. Flavonoids and the evolution of angiosperms. Biochem. Syst. Eco 2 HIERMANN, A., J. ExNER, H. BECKER & J. E. “AV VERETT. 1978. Gel filtration of flavonoids. Phytochem. 1942. Studies in Onagraceae XII. A revision of the New World species of Jussiaea. Darwiniana 4: 179-284. 1979. A sectional revision of Ann. Missouri Bot. Gard. 66: 8 . M. ZARDINI. 1987. The systematics and evolution of Ludwigia sect. Myrtocarpus sensu lato Die prends Monogr. Syst. Bot. Missouri Bot. ard. 19: 1-120. RAVEN, P. H. TA The Old World species of Lud- wigia (including Jussiaea), with a synopsis for the genus (Onagraceae). Reinwardtia 6: 327-427. & W. Observations of chromo- somes in Ludwigia —ewaspupa Ann. Missouri Bot. Gard. 66: 862-879. A CHEMOTAXONOMIC CLASSIFICATION OF THE SOLANACEAE! PETER TETENYI’ ABSTRACT Alkaloids and steroids in the Solanaceae are reported extensively in the baa nse By examining the in deen pai leading to pape alkaloids, the pathways can be v which th . Arrangem c us compounds can be der sualized as a spi ment of the genera of Solanacea their Padus contents in mesi em to pus igh — traditional classifications of the family, w subfamilie t unalia must be allied with Jaborosa in tribe olas Although solanaceous species are well known to afford an array of alkaloids and steroids, the family has not been arranged yet according to chemical features. Indeed, it is difficult to find reports of most features because, except for the occurrence of calcium oxalate crystals, only their absence is recorded. Thus Philipson (1977) con- cluded that a lack of iridioids characterizes the Solanaceae within the Unitegminae. Dahlgren (1975, 1980) came to similar conclusions, adding the deficiency of polyacetylenes as a character- istic. Sporne (1980) believed that absence of leu- coanthocyanins and ellagitanins are chemical characteristics of the family and indicated that whether or not the seeds contain endosperm may or may not be significant. In an investigation of the cytochrome-C and plastocyanin amino acid sequence, Boulter et al. (1979) placed the Solanaceae far from the As- teraceae in their family tree, and although they related it closely to the Scrophulariaceae, they & Emberger (1960) that the Solanaceae and Cap- rifoliaceae are closely related based on embry- ological characters. Nevertheless, mature sola- naceous plants are typical alkaloid-accumulators, whereas mature caprifoliaceous plants have phenol-glycosides (Hegnauer, 1973). In the same publication Boulter et al. (1979) put the tomato alongside the potato and separated tobacco and the woody Solanum crispum Ruiz & Pavón, and they held Capsicum frutescens L. to be signifi- 1c ! This paper was part of the Second I s due to their e must be a pei ed as separate and exclusive uod synthesis. Acnistus and cantly different from the preceding species based on the amino acid complement. The use of active principles found in different Solanaceae to construct systematic schemes can be accepted only when it can be demonstrated that the biosynthetic routes leading to these chemical structures are homologous (Tétényi, 1973). The valid chemical patterns are in the various biosynthetic pathways and not in the thetic pathways in alkaloid production in the So- lanaceae in Table | and Figure 1. Numbers in the following paragraphs correspond to those of Table 1 and Figure 1. The first evidence supporting this scheme lies in the well-known, genetically determined chem- ical differentiation in infraspecific chemotaxa of Duboisia myoporoides R. Br. (Tétényi, 1970). The characteristic active alkaloid ingredient, nicotine (1), in one chemotaxon ofthis species is the result of the synthesis of ornithine and tryptophan to an alkaloid. Another chemotaxon of D. myopo- roides is characterized by a splicing of the as- partate pattern (lysine) and acetyl-CoA to the alkaloid biosynthesis, and the main alkaloids are then anabasine (A) and isopelletierine (B). In a third infraspecific chemotaxon, an entirely dif- ferent pattern occurs: an a-face nucleophilic at- tack instead of the 8-face one of the N-methyl- A'-pyrrolinium salt (Fig. 2; Leete, 1979) leads to the pathways indicated by the solalkoid spiral (Fig. 1), that is, the linking of ornithine and ace- tyl-CoA to hygrine (2) and then the development on the Biology and Systematics of the Solanaceae presented at the Missouri Botanical Garden on 3-6 August t 1983. ? Research Institute for Medicinal Plants, P.O. Box 11, 2011 Budakalász, Hungary. ANN. MISSOURI Bor. GARD. 74: 600—608. 1987. 1987] TABLE l. TÉTÉNYI—CHEMOTAXONOMIC CLASSIFICATION OF SOLANACEAE Alkaloids of Solanaceae included in and arranged conforming to the solalkoid spiral. Nicotine and its derivatives ug (1) HN; p 11-5] Tropane alkaloids hygrine (2) C,H,;N;O, [496-49-1] cuscohygrine (4) C. H, NO [454-14-8] tropinone (5) C,H,,NO [532-24-1] isovaleryloxytropane (6) 13H23 y. [490-96-0] tigloidine (7) C.H; NO; [495-83-0] scopolamine (8) Other alkaloids fabianine (—) O anabasine (A) C l oH ThE [494-52-0] hygroline (2) aH; [1617-83-0] physoperuvine (5) 8 15 [60723-27-5] tigloyloxytropane (7) 134421 3 [55727-41-8] hyoscyamine (8) iH; NO; [101-31-5] 8-carboline (C) isopelletierine (B) C,H;,NO [539-00-4] withasomnine (3) 12 15 a [10183-74-1] belladonnine (4) C,,H4;N; 4 [6696-63-5] tropine (5) C,H,;NO [120-29-6] senecioyloxytropane (6) 134421 O, [77101-57-6] valtropine (7) physochlaine (8) C,,H2;NO, [54357-41-4] capsaicine (D) CHa, N n N.O 184427 3 [6871-51-8] [244-63-3] [54357-41-4] betaine (E) Pictur (E) C. Hi NO, C;H;; NO, [107-43-7] [62- 49- 7] Explanation: common name; symbols in parentheses refer to Figure 1 formula [registry number] of other simple and ester tropane alkaloids. Thus pyruvate-leucine yields valeroidine (6) with iso- valeric acid; aspartate-isoleucine yields tiglo- idine (7) with tiglic acid; and om phe- nylalanine and tropic acid, becomes the characteristic alkaloid of the third chemotaxon. The genus Duboisia R. Br. provides further evidence of this kind of chemical differentiation separating the infraspecific taxa of D. myopo- roides from one another. In D. hopwoodii F. Muell., nicotine (1) and its derivatives are syn- thesized by the 8-face attack of ornithine, where- a this alkaloid is accumulated through the con- verse formation. Thus the scheme in Figure 1 is true for Duboisia as a genus as well as for its components. Two other genera examined in subfamily An- thocercidoideae (described on p. 607)— which includes Duboisia —show the same diversity in asine (A), cus e (4) opine (5), va- leroidine (6), tigloidine (7), and scopolamine (8) have been detected. Alkaloid synthesis in subfamily Cestroideae Schldl. also proceeds according to Figure 1, but its derivatives, a somewhat different picture from the Anthocercidoideae. Thus nicotine (1) is the sole product in the genus Cestrum L., whereas fabianine, a tetrahydroquinoline alkaloid arising from a biosynthesis preceding the pyridine-nu- cleotide cycle, is accumulated as well as nicotine in Fabiana Ruiz & Pavón. The aspartate-lysine path is switched in Streptosolen Miers, and an- abasine is accumulated in addition to nicotine. The alkaloid spectrum of Nicotiana L. and Sal- piglossis Ruiz & Pavón by intervention of acetyl- CoA, accumulates isopelletierine (B) in addition 602 SOLALKOID SPIRAL P-hydroxypyruvate serine Pere oe tryptophan aspartate ae ornithin phenylalanine DOSE DN (0 nicotine (2) hygrine (3) withasomnine (4) cuscohygrine (5) tropinone-physoperuvine isovaleryl (8) sonecioyl d ED) tigloyl (8) scopolamine-hyoscyamine tropane FIGURE 1. from plants belongi ng t e Solan Am cids alkaloid biosynthesis in “issuu are e clarified ir in rn to the above substances. Nicotiana is even en- kaloid of Vestia Willd., which contains a 8-car- boline skeleton. My scheme might not be tenable if the Ces- troideae were not differentiated chemically also. In fact, the alkaloid biogenesis of hygrine deriv- atives is found in this subfamily too, although only Brunfelsia L. accumulates just cuscohygrine (4) and pyrrole-3-carboximidine, which was also detected in Nierembergia Ruiz & Pavón, whereas 4 th z A in v. L: L ANNALS OF THE MISSOURI BOTANICAL GARDEN > A B isopelletierine c D E realt [Vor. 74 pyruvate leucine-valine anabasine B-carboline capsaicine choline-betaine Solalkoid m Psp on ey ea Piet load patterns of most important alkaloids isolated of d d starting and combined points. Patterns Ruiz & Pavón. However, the alkaloid spectrum in Schizanthus is unique in the whole family with its hygroline (occurring also in the Erythroxy- laceae) and because the sencioyl and angeloyl tropane esters (6) are formed from pyruvate-leu- cine and ornithine. Formation of the alkaloid valeroidine in Schizanthus by this biosynthetic pathway is characteristic of the family Solana- ceae as well as of Anthocercis Labill., Cyphanth- era Miers, and Duboisia. However, taking into account the two kinds of hygrine synthesis, the predominance of nicotine in its various deriva- tives (A, B, C), and the primary substances brun- felsamine and fabianine, we can assert that the 1987] TETENYI—CHEMOTAXONOMIC CLASSIFICATION OF SOLANACEAE (n- (2R) - Hygrine FIGURE 2. chemical differentiation of the Cestroideae, in having an alkaloid synthesis with several simi- larities to that ofthe Erythroxylaceae, Rubiaceae, Elaeagnaceae, and Malpighiaceae, significantly surpasses that of the Anthocercidoideae. Chemical differentiation in subfamily Sola- noideae is of a different character than in the Cestroideae, for here the hygrine derivatives are always preponderant, and the differences are in the ever-increasing complexity of the molecular entities synthesized by the different genera as if overcoming a barrier. Similar to the case of Brun- (4) Adans. starts from hygrine (2) and, like CypAo- mandra Sendtner, reaches the formation of tro- pinone (5). The alkaloid physoperuvine (5) of Physalis L. is equivalent to this, although tiglo- idine (7), an alkaloid of this genus, is an ester but not with tropic acid. In Withania Pauquy aspartate-isoleucine and ornithine are synthe- sized to 3'-tigloyloxytropane without the pres- (-)- (25)- Nicotine Nucleophilic attack of N-methyl-A'-pyrrolinium salt from the a- and 6-face (Leete, 1979). ence of any tropic ester, but the alkaloid spec- trum of this genus is very broad; thus withasomnine (3) is formed by coupling orni- thine to phenylalanine. Isopelletierine (B) from lysine and acetyl-CoA and even the alkaloid cho- line (E) arise from the hydroxypyruvate synthesis pathway. These are only the main types repre- sented. In the genera Leucophysalis Rydb. (Phys- aliastrum japonicum (Franchet & Savat.) Hon- da) and Lycium L. of this subfamily, the unique alkaloid detected so far is betaine (E), formed from the serine cycle in the same way as choline. synthetic routes from nicotine (1) to scopolamine (8), while on the other hand, they also form and accumulate the alkaloid valeroidine (6) like An- thocercis and Duboisia. In the genus Solandra Sw., valtropine (7), originating from the coupling of isoleucine with ornithine, follows a path of alkaloid biosynthesis homologous to that in Cy- phanthera and Grammosolen Haegi. However, 604 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Differences of t um derivative alkaloids in subfamily Atropoideae. Data from TABLE 2. Romeike (1978) except where noted otherw Genus Man- Hyoscy- Scopo- Whit- dra- Prze- Atro- Physo- Alkaloids Atropa amus lia leya gora walskia | panthe | chlaina Physochlaine + Hyoscyamine + + + + + + + + 3'-tigloyloxytropane + + + Belladonnine + + Tropine + + + + Tetramethylputrescine + A-N-methylornithine (Hedges & Herbert, 1981) + a related feature in the genus Anthotroche Endl. may be that hyoscyamine (8) predominates in the spectrum. The hygrine (2) of Salpichroa Miers also shows relationship to Duboisia, bu e hyoscyamine (8) is the characteristic alkaloid, similar to the content of tigloidine (7) in Acnistus Schott and the small amount of scopolamine (8) E Latua Phn, ds combination of pyruvate-va- (D) supports the curious alkaloid ere and the isola- tion of Capsicum L. as stated by Boulter et al. (1979) The system shown in Figure | is also supported by the alkaloid synthesis in subfamily Atropo- ideae (described on p. 607). In contrast to the Solanoideae, the simple alkaloid synthesis of the Atropoideae is subordinate to that of tropane esters. This is illustrated by the data in Table 2, for initially nicotine formation is only inhibit- ed—as exemplified by Atropa L. and Hyo- scyamus L.—and the genera Scopolia Jacq. and Whitleya Sweet (Anisodus Link) are the most advanced cases in which a simple tropane base could be detected. Occurrence of tigloyl ester (7) can be verified in Mandragora L. only by the presence of cuscohygrine (4) and a little scopol- amine (8). Przewalskia Maxim. and Atropanthe Pascher accumulate mainly hyoscyamine (8), which is characteristic in the whole family and occurs in each genus. Although the result of the synthesis in Physochlaina G. Don is also pre- dominantly hyoscyamine, its peculiar alkaloid, physochlaine (8), is the tropane ester of 4 si beastacsticacid: nol "Piu oM" = = Q € r rivative. The alkaloid-forming distinctions of solana- ceous taxa at various levels have been arranged according to Figure 1. The requirements were satisfied by this scheme, which we have termed the solalkoid spiral. One objection may be that I have not mentioned the connection between terpenoid and alkaloid biosynthesis. This is be- cause the steroid synthesis that is characteristic of the family is exclusive, alternative, and with- out transition, in contrast to the process of al- kaloid synthesis, which is gradual and proceeds y inhibition "This is poai by the omg * infraspecific chemotaxa carried out with anolides on Withania somnifera Dunal UE 1973) as well as our own analysis of steroidal alkaloid taxa of Solanum dulcamara L. (Tétényi et al., 1977). No instance of a steroidal alkaloid in a withanolide- containing species or the reverse is known, al- though this may be due to deficiency in our equipment or methods. It seems that these two routes of steroid syn- thesis — steroidal alkaloids versus lactones — rep- resent a phylogenetic branching alternative with cholesterol as its starting point. Oxidation leads to withanolides compared with partial etherifi- cation to neutral saponins, followed by cycliza- tion to steroidal alkaloids. Duboisia —subfamily Anthocercidoideae — synthesizes only neutral sapogenins, and Cy- phanthera, Anthotroche, Crenidium Haegi ur- solic acid only, a further feature pointing to their primordial endemism. Simple sapogenins were detected in subfamily Cestroideae: in Combera Sandw., Fabiana, Nierembergia, Browallia L., 1987] TETENYI—CHEMOTAXONOMIC CLASSIFICATION OF SOLANACEAE 605 FIGURE 3. Chemotaxonomic syste t fg in four subfamilies containing 16 tribes and i Duro Dahlgren o Axes separate most importan ] isopelletierine type [N-I], O = LE -carb [H-T], © = capsaicine. Anthoc 3. Nicotianeae = Don. (8). etieae A. T. Hunz. (1). Solanoideae.—7. Juannullo Nicandreae Wettst. (1).—14. Lycieae A. T. Hun des L.A 24 Salpislosolid|eae Benth. (7 oeae A. T. Hunz. (8). neae-Phy idem (9). —10. Jaboroscae Miers (7).—11. Datureae Reichb. (2).—12 z. (3). Atropoideae.— 15. Discopodiineae Baehni (2).— 16. Atro- kaloid [ALO = lactone [L]. aru ids (A): V = ae. oline-betaine, A = hygrine-tropane type 5 hocerceae G. Don. (7). Cestroideae. E bid aaa Don ).— ).— 5. Schwenckieae A. T z. (3).—6. Parabouch- —8. in Reh G (15).—9. Sola- . Solandreae Miers (2). —13. peae Reichb. (9). Numbers in ei mide numbers of genera Salpiglossis, Vestia, and Streptosolen, as well as in Cestrum and Nicotiana, but these two have steroidal alkaloids. Subfamily Solanoideae, apart from Exodeco- nus Raf., shows a prevalence of steroidal alka- loids when the genera Solanum, Cyphomandra, Capsicum, and Lycopersicon Miller are consid- pogenin-containing 7rechonaetes Miers, should be evaluated as quite distinct. This differentiated steroid synthesis divides subfamily Solanoideae. The connection between alkaloid and steroid biosynthesis is shown by the fact that Solanum, arrested at cuscohygrine, includes species accu- mulating steroidal alkaloids building in arginine released from the ornithine cycle in their genesis. The genera Physochlaina and Scopolia o subfamily bd ones synthesize kei yoscyam thus the rss Is nearer to the subtribe of the Solanoi- deae characterized by formation of the same ste- roids. If we base a chemical classification of the So- lanaceae on the facts mentioned above, we can- not accept the analysis of Evans (1979) or Romeike (1978), who considered only alkaloid properties in evaluating Wettstein’s century-old system. I have developed in Figure 3 a chemo- taxonomic classification of the Solanaceae. It "- 606 TABLE 3. Characteristic alkaloids and steroids for the chemotaxonomic system of the Solanaceae (corre- sponding to Fig. 3). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Tribe Genus Alkaloid Steroid 1 Anthocercis oe, 10 scopolamine”? t Anthotroche hyoscyam Crenidium ee anabasine'* T Cyphanthera scopolamine,'* nicotine!* Duboisia SEROREN isopelletierine! sapogenin? Grammosolen scopolamine!? Symonanthus tigloyl wont 2 Cestrum nicotine? solasodin? Vestia B-carboline'? diosgenin? 3 Combera — sapogenin? abiana "en send sapogenin? Nicotiana betaine,’ isopellet solasodin? Nierembergia B- nea paces + EE E E sapogenin‘* 4 Browallia sapogenin? Brunfelsia cu uscohyrine" + brunfelsamine!* — Salpiglossis isopelletieri sapogenin? Schizanth a LUN hygrolines' — Streptosolen nabasine? sapogenin? 8 sicum capsaicine? solanidin? Cyphomandra tropinone! solasodin! Exodeconus — neotigogenin? Jaltomata cuscohygrine? — Lycopersicon — tomatin’ nu cuscohygrine (solamine)* solasodin? 9 Leucophysalis etaine? — Margaranthus cuscohygrine* — Physalis tigloidine,' physoperuvine' physalin? Withania choline, isopelletierine, 3'-tigloyloxytropane! withanolide? Witheringia physalin? 10 Acnistus — 19 withanolide!! Dunalia — withanolide!! Jaborosa — steroidlactone!! Latua hyoscyamine! — Salpichr hyoscyamine! — Trechonaetes — sapogenin? 11 Brugmansia nicotine, scopolamine! — Datura nicotine, scopolamine! withanolide? 12 Solandra hyoscyamine! — 13 Nicandra tropinone! withanolide? 14 Lycium betaine? withanolide? 16 ropa choline, '? Mio — Atropanthe hyoscya — yoscyamus choline? hyoscyamine’ sapogenin? Mandragora hyoscyamine — Physochlaina physochlaine! solanidin? Przewalskia hyoscyamine!' — Scopolia choline,'? hyoscyamine! solanidin? Whitleya choline,'? hyoscyamine! — T Ursolic acids present.'^ ! Romeike, 1978; ? Gibbs, 1974; ? Hegnauer, 1973; * Lorenti et al., 1981; * Evans, 1979; * San Martin et al., 1980; ? Paris & Moyse; 1971: 8 Evans & Somanabandhu, 1980; ° Antoun et al., 1981; '? Evans & Ramsey, 1981; u D'Arcy, 1979; '? Gessner, 1977; ? Evans & Ramsey, 1983; '* El Imam & Evans, 1984; 15 Buschi & Pomilio, 1986; '^ Lloyd et al., 1985; 1 Faini et al., 1980. 1987] considers the chemosyndrome of the Solanaceae, has reference to the cross-sectional presentation of Dahlgren (1975, 1980), and draws upon the systematic data of D’Arcy (1979), Haegi (1979, 1981), and Hunziker (1979). The surface dimensions of the four subfamilies correspond to the number of included genera, while the curves involving the subfamilies label the affinity points of the homologous chemical qualities. The two main trends of alkaloid syn- thesis in the family— accumulation of nicotine- steroidal alkaloid taxa. Tribes of the subfamilies are shown by numbers and dotted lines. I have divided the tribe Solaneae Reichb. into subtribes Solaninae (Solanineae Dunal) and Physalinae (Physalidineae Reichb.) on the basis of different and exclusive types of steroid synthesis and con- sidering the data of Baehni (1 Acnistus and Dunalia are placed in tribe Jabo- roseae Miers in accordance with Baehni on the basis of their steroidal lactone content. Latua was similarly treated—its bent embryo agrees with this placement and its alkaloid chemistry precludes assigning it to subfamily Cestroideae. e characteristic alkaloid or steroidal data for the genera shown in Figure 3 are also shown in able 3. I have not found reliable recent data on the alkaloid or steroid active ingredient of four tribes and 43 genera; however, I feel justified in presenting this review, a novel chemotaxonomic evaluation of facts known to others, and a clas- sification system for the Solanaceae. This new system consists of four subfamilies. We must separate the Anthocercidoideae from the Cestroideae, and the Atropoideae from the Solanoideae because of differences in area of or- igin, morphology, Bowers; ane enue oe and are endemic to Australia. They differ mor- phological from the Cestroideae by having long, d corolla tubes, grains. pm chemosyndrome combines predominantly hy- grine derivatives— characteri Such pude. amine — with neutral sapogenins, while the Ces- troideae have chiefly nicotine derivatives together with simple and complex steroidal alkaloids, and only one tropane ester alkaloid in common with the Anthocercidoideae The Atopadas; which have a cistmcuive al- kaloi differ HRALVIUNL pa J F TÉTÉNYI—CHEMOTAXONOMIC CLASSIFICATION OF SOLANACEAE 607 also in having dispersed from their l cen- ter. They are adapted to withstand cold seasons in Eurasia and have become isolated on islands mountains in Africa. The Atropoideae having imb noideae have valvate—sometimes twofold val- vate—aestivation. The Atropoideae chemosyn- drome combines highly derived tropane ester alkaloids with steroidal alkaloids, while the So- lanoideae possess one of these biosyntheses but have the other only in inhibited form as simple tropanes or as steroidal lactones. These evolu- tionary patterns are sufficient to warrant recog- nition of the taxa as subfamilies. Subfamily ANTHOCERCIDOIDEAE Tété- nyi, subfam. nov Plantae frutescentes caulibus lignosis in Australia habitantes. Aestivatio valvato-aperta. Co Mid profundae didynama v extrorsae longitudinaliter dehiscentes. die capsu- lares aut baccatus. Embryo parum curvus. Plantae praecipue alcaloidam *Scopolamin" ds eoi -sapo- ninem continentes. Woody shrubs inhabiting Australia. Corolla with rolled, inflexed-valvate aestivation of the 5 lobes, these striated along main veins, varying in length but never as long as the tube. Stamens didynamous or equal, epipetalous low in corolla tube, the anthers dehiscing extrorsely by longi- tudinal slits. Fruit capsular or baccate. Embryo only slightly curved. Plants containing predom- inantly the alkaloid scopolamine and steroid sa- ponins. Type: Anthocercis Labill. Subfamily ATROPOIDEAE Tétényi, subfam. nov. Plantae ex orbe antiquo oriundae. Herbae locis cal- regionis ‘montium excelsium vel insularum. Corolla tus. Plantae praecipue alcaloidam “‘Hyoscyamin” steroidalcaloidam continentes Herbaceous plants of the Old World, occurring exceptionally as endemic woody shrubs on mountains or islands in warmer regions. Corolla tubular or campanulate, the aestivation of lobes imbricate. Stamens 5, equal. Fruit baccate or capsular. Embryo curved circularly or in a spiral. 608 Plants containing pre onu nanu y the alkaloid lalkaloids. Type: Atro- pa b LITERATURE CITED ANTOUN, M. D., D. ABRAHAMSON, L. R. TvsoN, C. M McLAUGHLIN, G. PECK & J. . Potential antitumor agents md Physalin B and 25, 26-epidihydrophysalin C from Witheringia e Lloydia 44: 579—585. BAEHNI, C. uverture du bouton chez les fleurs de Solanées. eile 10: 399-492. BOULTER, D., EACOCK, A. GUISE, J. T. GLEAVES & G. ESTABROOK. 1979. piede ts between partial amino acid sequences. Phytochemistry 18: —608 BuscHi, C. A. & A. B. PoMiLio. 1986. Identificación del compuesto letal de Nierembergia hippomani- ca. II. Latinamerican Phytochemical Congress, La Plata, Argentina. Abstract CHADEFAUD, M. & EMBERGER. 1960. Traité de Botanique Systématique, Volume 2. Masson, Paris. DAHLGREN, R. A system of classification of the apom Bot. Not. 128: 119-147 1980. A revised system of classification of L^ e o wies J. Linn. Soc., Bot. 4. v, W. G. 1979. Classification of Solanaceae. . G. Hawkes, R. N. Lester & A. D. Skelding META The Biology and Taxonomy of the So- Academic Press, London. W. C. Evans. 1984. Tropane alkaloids of species of Anthocercis, Cyphanthera a and Crenidium. Pl. Med. 50: 86-87. Evans, W. C 79. Tropane alkaloids of the Sola- nac . G. Hawkes, R. N. Lester & A. D. Skelding (editors), The Biology and Taxonomy of the Solanaceae. Academic Press, London. . A. RAMSEY. 1981. Tropane alkaloids from Anthocercis and Anthotroche. Phytochem- istry 20: 497- Alkaloids of the Solanaceae tribe Anthocercideae. Phytochemistry 22: 2219- 2225. & A. SOMANABANDHU. 1980. Nitrogen-con- taining non-steroidal secondary metabolites of Solanum, Cyphomandra, Lycianthes and Mar- garanthus. Phytochemistry 19: 2351-2356. FAINI, F., R. TORR carboxy-8-carbolin cid a rss An Vestia ia. PI. Med. 38: mi 132. GESSNER, O. 1977. Gift- und Arzneipflanzen von Mitteleuropa. Carl Winter Universitäts Vlg., Hei- delberg. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 GiBBs, R. D. 1974. Chemotaxonomy of Plants. McGill-Queen's, Montreal & London HAEGI, L. 1979. Australian genera of the Solanaceae. J. s, R. N. Lester & A. D. Skelding (editors), The Biology and Taxonomy of the So- lanaceae. Academic Press, London. A conspectus of Solanaceae tribe An- thocercideae. Telopea 2: 173- HEDGES, S. H. & . HERBERT. 1981. 6-N-meth- ylornithine: a n natural constituent of Atropa bel- ladonna. Phytochemistry 20: 2064-2065. 1973. Chemotaxonomie der Pflanzen, asel. ae: A. T. a synoptic surv n J. & A. D. Sk elding pens The | Biology and Tax- onomy of the Solanaceae. Academic Press, Lon- Se. uth n Solanaceae: , R. N. Lester LaviE, E. 1973. Applying chemistry to genetics in certain Solanaceae. Pp. 181-188 in m n in Botanical Classification. Nobel Symp. 25. Aca- demic Press, London & Ne LEETE, E. 1979. VAL and metabolism of the tropane alkaloids. Pl. Med. 36: 97- Be LLovp, H. A., H. M. FALEs, M. E. GOLDMAN, D. M JERINA, T. PLOWMAN p R. E. ume 1985. Brunfelsamine: a novel convulsant. Tetrahedron Lett. 26: 2623-2624. LonENTI, A. E., A. A. VITALE, C. A. Bus ZALEZ, C. D. “pass si . M. IRRIBAREN & A. B. PoMILIO. 1981. Antimicrobial activity of some Argentine higher plants. Fitoterapia 52: 81-85. Paris, R. R. & H. Moyse. 1971. médicale. Masson, Paris. PHILIPSON, W. B 77. Ovular morphology and the CHI, M. D. Précis de matiére classification of the dicotyledons. Pl. Syst. Evol. Suppl. 1: ROMEIKE, A. 1978. Tropane alkaloids — occurrence and systematic importance in angiosperms. Bot Not. 131: 85 rm SAN MARTIN, A., J. R A, O. V. GAMBARO & M CASTILLO. i980. noe alkaloids fom Schi- zanthus hookeri. Phytochemistry 19: 20 00 SPORNE, K. 19 re-investigation of character ati correlations among dicotyledonous plants. New Phytol. 85: 419-449, TÉTÉNYI, P. 19 Infraspecific Chemical Taxa of Medicinal Plants. Chemical Publishing Co., New de Homology of biosynthetic bacs the base i chemotax l n Chem- Istry in eo a Classification. Nobel Symp. 25. agg tee iin London & New York. H & N. Vo-Honc. 1977. Steroid BE in ` dulcamara populations. Herba. Hung. 16: 55-60. FLORA OF THE VENEZUELAN GUAYANA — III! JULIAN A. STEYERMARK2 ABSTRACT Continued pote of various families of the flora of the Venezuelan Guayana have resulted in the hu xa: Stegolepis albiflora, S. beri, S. humilis (Rapateaceae); Panopsis cuaensis, i chae, G. bolivarensis, G sancarlosiana sip na, Neea amaruayensis, d nardi, N. pisite = ode acaba N. cedenensis, N. clarkii, N. davidsei, N. guaiquinimae, cola, N. lie. neri, N. mapourioides, N. . marahuacae, N. parimensis, N. robusta, " sebastianii, N. A redis N. pies (Nyctaginaceae); Brunellia neblinensis (Brunelliaceae): Matayba ptariana subsp. guaiquinimae (Sapin sutula, C. Bonnetia bolivarensis, B. euryant daceae); Catostemma cla C. marahuacensis, C. rie tyla, C. sancarlosiana, Scleronema si spe (Bombacaceae); . guaiquinimae, B. ptariensis, B. trist la placea fruticosa var. chimantae (Theaceae). Daphnopsis guaiquinimae, D. monocephala (Rubiaceae), a total of 50 spe or the Venezuelan Gua piel zur of Proteaceae, rkii, C. ebracteolata, C. hir- variety. uapira, Neea, Catostemma, Symplocos, and newly described taxa of Bonnet RAPATEACEAE STEGOLEPIS Stegolepis albiflora Steyermark, sp. nov. TYPE Venezuela. Bolívar: Meseta de Jaua; Cerro Sarisarinama, northeastern part, 4?41'40"N, 64*13'20"W, 1,410 m, 10 Feb. 1974, Stey- ermark et al. 108873 (holotype, VEN; iso- type, NY). erbae perennes usque 1.5 m; vaginis prope apicem Peroni tenuiterque multinervatis eligulatis; pedun- culis 7-18 axillaribus 4.5-7 dm longis 0.8-1.5 mm diam.; capitulis globosis 1.8-2.5 cm diam., spiculis 6— 14 maturis lanceolatis acutis 7210 mm longis 2-3 mm latis; petalis albidis. Herbaceous perennials up to 1.5 m tall. Sheaths eligulate, finely and many-nerved near the apex or along one side, 20-25 cm long, 4-6 cm wide. Leaf blades rich green both sides, 58-70 cm long, 3.5-5 cm wide, the nerves and midrib on lower side somewhat more prominent than on upper side. Peduncles numerous, 7-18, 45-70 cm long, 0.8-1.5 mm diam., many-sulcate. Heads glo- bose, tan or tawny, 1.8-2.5 cm diam. Spikelets spreading in various €— "sasawa acute, 7-10 mm long, 2-3 mm e. Bracteoles 14-15, graduate, the lower outer ones ien be ular, 222.5 x 2-2.5 mm, the middle ones obtuse, the upper ovate-lanceolate, obtuse to subacute, 7 x 4 mm. Sepals lanceolate, subacute, 6—7 mm long, 2.5-3 mm wide. Petals white, oblanceolate or lanceolate, acute, recurved, 5.5-7 mm long, 1.5-2 mm wide toward the middle. Anthers lin- ear, 3.5 x 0.4 mm. Ovary depressed-globose, | m high. Style 1 mm long. Seeds ire barrel- ee rounded at both ends, 1.5 x 1 mm Paratypes. VENEZUELA. BOLÍVAR: Cerro Sarisari- ñama, summit, W-central part, 4?45'N, 64?26'W, 1,922— 2,100 m, 22-27 Feb. 1967, Steyermark 97839 (VEN); Meseta de Jaua, Cerro Jaua, summit, SW forest, 64?34'10"W, 1,750-1,800 m, 22-28 Feb. 1974, Stey- ermark, Carreño & NE n (VEN); eseta de Jaua, o Jaua, summit, SW part, 4*47'22"N, 64?33' 35"W, 2 228-2,250 m, 2j Feb. 1974, Steyermark, Carrefio & Brewer-Carias 109581 (VEN); Meseta de Jaua, Cerro Jaua, summit, E-central part, 4°35'N, 64?15'W, 14 Feb. 1981, Steyermark, Brewer- Carias & Liesner 124320 (VEN), 124321 (VEN); trail to Sima menor, Cerro Sarisarifiama, Ravelo 17 (MY). This species is remarkable for having white petals and is the only white-flowered member of the otherwise yellow-petaled genus Stegolepis. Maguire (1976) identified all the from the Meseta Jaua (Cerro Jaua and Cerro Sarisa- rinama) as S. choripetala, a species originally de- scribed from Cerro Sipapo and not known else- where. The new taxon differs from S. choripetala not only in the white petals, but also in the small- er mature spikes, the greater number of more slender, shorter peduncles, and in the finely nerved sheath in the apical portion. The collections cited were obtained from var- ious parts of the huge Meseta de Jaua, indicating that the species is well distributed over the whole ! Flora ofthe V 1 y : e 4 under NI a 1¢ T 4 Fal ? Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ANN. MissouRi Bor. GARD. 74: 609-658. 1987. «/(DCDiQ&1&^5nQo« J ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Stegolepis iid — A. Habit —G. Stamen. Based on holoty FIGURE |. natural position. — F. Pet summit and is autochthonous to that table mountain. Stegolepis huberi Steyermark, sp. nov. TYPE: ela. Bolívar: Kukenán tepui, 5?16'N, 60?48'W, 2,500 m, 28 Apr. 1984, Otto Hu- ber 9467 (holotype, VEN; isotype, MO). erbae perennes humiles 23-35 cm; vaginis 7-10 m latis omnino nervatis ligulatis, au- m latis; laminis coriaceis ligulato-lanceolatis acutis < 20 cm longis 1.8-2.5 cm latis, costa media haud m nifesta subtus tenuiter aden puse nervis ne soletis; pedunculis 17-25 cm longis v mm latis infra capitu tulum 4-7 mm dilatatis; capitu lis ompressis 4—6-floris 2—3.5 cm latis; spiculis elliptico- dcos . — B, C. Base A leaf with auricle of sheath. — D. Spikelet. — E. Flower, sub fructu 15-18 mm longis; bracteolis oblongo-lan- d vel lanceolatis acutis vel acuminatis 4-10 mm longis 1.1-3 mm latis; petalis flabellato-rhomboideis 10 mm ns 7-8 mm latis. Dwarf perennials 23-35 cm tall. Sheaths mem- branous, subscarious, 7-10 cm long, 3-3.5 cm wide, venose throughout, the auricles rounded at apex, 10-15 mm long, 17-22 mm wide. Leaf blades coriaceous, galate: -lanceolate, 15-20 cm m wide, acute, finely nerved on midrib obsolete on both sides. Peduncles com- pressed, somewhat alate, 3—6-costate, 17-25 cm long, 2-3.5 mm wide except 4-7 mm where di- lated below inflorescence. Heads compressed, 4— 1987] 6-flowered, 2-3.5 cm wide, 1—1.7 cm high. Spike- lets (in fruit) elliptic-lanceolate, 15-18 mm long, 5-seriate. Bracteoles 18-24, oblong-lanceolate or lanceolate, acute to acuminate, lower ones 4-5 mm long, 1.1-1.2 mm wide, the others 7-10 mm long, 2-3 mm wide. Sepals (in fruit) oblanceolate, acute, 13 mm long, 5 mm wide. Petals flabellate- rhomboid, 10 mm long, 7-8 mm w This species is the shortest known in the genus. All the plants seen (Huber, pers. comm.) in a large colony were similarly small. The member of the genus to which it shows closest affinity is S. ptaritepuiensis Steyerm., from which it differs in the shorter peduncles and leaves, membra- nous, striate-nerved sheaths, smaller heads with shorter spikelets, l lets, midri not perceptible on either side, and a longer au- ricle of the sheath. Stegolepis humilis Steyermark, sp. nov. TYPE: Venezuela. Bolivar: Camarcaibarai tepui, SW-facing shoulder, 5°52'N, 62?1'W, 1,800- 1,825 m, 22-24 May 1986, Julian A. Stey- ermark, R. Liesner & B. Holst 132006 (ho- lotype, MO; isotype, VEN). Figure 1. erbae perennes humiles 27-52 cm, vaginis mani- feste nervatis 12 cm longis ligulatis, auriculis apice rotundatis 5-7 mm longis; rpm nis linearibus apice acutis 27-45 cm longis 1.3-2. i subtus prominenti, nervis subt subtiliter prominentibus; pedunculi s lanceo- lanceolatis acutis 15 mm longis 5-6 mm latis; petalis late flabellati i bt to-rotundatis 12-13 mm longis 18 mm latis. Dwarf perennials 27-52 cm tall, the caudex flattened, 11-12 cm long, 7-9 cm wide. Leaf sheaths conduplicate, firmly membranous, con- spicuously nerved, 12 cm long, 7 cm wide, lig- ulate, the auricles rounded, these 5-7 mm long, STEYERMARK— VENEZUELAN GUAYANA FLORA- III 611 12-13 mm wide. Leaf blades silvery green below, linear, symmetrical to slightly subfalcate at the acute apex, 27-45 cm long, 1.3-2.5 cm wide, finely nerved above, prominently nerved below, midrib prominent below, 1 mm wide. Peduncles 2-3, 5-costate with rounded ribs, 27-52 cm long, 1-1.5 mm diam. except below the inflorescence where flattened and dilated to 4-5 mm wide. mainly 1, sometimes 2, compressed. Spikelets d fusiform, 15-18 mm long, e. Bracteoles dark mahogany or chestnut is indurated, 19-21, the lowest suborbicular-ovate, broadly acute, 4.5-6 mm long, 4-5 mm wide at base, the upper ones lan- ceolate, acuminate, 12 mm long, 5 mm wide at base. Sepals broadly lanceolate, indurated above the middle, sharply slenderly acute, 15 mm long, mm wide. Petals with a broadly rhombic- flabellate blade, broadly subtruncate-rounded at apex with a mucronate center, 12-13 mm long, 18 mm wide, unguiculate 11 mm. Anthers 10 mm long; filaments 9.5 mm long. Style subulate, 12 mm long. Paratypes. "VENEZUELA. BOLÍVAR: Murisipán tepui summit, 5°52 Steyermark & eee re (MO, tepui, summit °52'N 2 : rd 1986, Liesner. y odds & Ha 21 075 (MO, N). "N This taxon is related to Stegolepis terrama- rensis Steyerm. from Cerro Marahuaca, from which it differs in the solitary, or rarely two spikes, acute leaf apex, fewer and more slender pedun- cles, smaller size, shorter auricles of the nerved, nonindurated ligulate sheath, and more con- spicuous midrib and secondary nerves. It differs from the other species with one to few spikelets in size and details of spikelets, bracteoles, leaf nervation, and peduncle LITERATURE CITED MaGuinE, B. 1976. Rapateaceae. In J. A. Steyermark & C. Brewer-Carias, La Vegetación de la Cima del Macizo de Jaua. Bol. Soc. Venez. Ci. Nat. 32(132- 133): 279. PROTEACEAE PANOPSIS KEY TO THE SPECIES OF PANOPSIS la. Leaves sessile or subsessile; reticulation coarsely areolate pod areoles 1-5 mm diam.; leaves crowded on the stem, pseudoverticillate; fruit subglobose, 4-5 cm P. sessilifolia lb. Leaves petiolate; reticulation generally more minutely AR with areoles 0.5-1 mm diam.; leaves 612 lam ANNALS OF THE MISSOURI BOTANICAL GARDEN scattered on the stem, not pseudoverticillate; fruit fusiform or longer than broad or less than 4 cm in di [Vor. 74 2a. Stems, leaf blades, and petioles glabrous or d so 3a Leaves abruptly acuminate-cuspidate wh pex, 8.5-11.5 cm long; reticulation minute and ics on both sides; inflorescence m ^ cm long with axes 0.7-1 cm; below 150 m elev., Terr. Fed. Amazonas P. cuaensis 3b. Leaves rounded at apex, 3.5-6 cm long; reticulation elevated and manifest on both sides; N c bana, Edo. Bolív . Young stems, portions of the leaf blades, and icr oles l cm with axes 2-3 cm long; plants at elev. of P. ptariana pubesce 4a. oo mainly 8-20 cm long, 3-7 cm wide; inflorescence 15-30 cm long; plants of d elev rubescens 4b. pati 2-15 cm long, 2.5-4 cm wide; inflorescence 5-8 cm long; plants of 1,150-2, cri l eiev. 5a. Trees 19—22 m tall; leaves 8-15 cm lon 5b. Small shrubs 1-1.5 m tall; leaves 2-9 cm lon g P. tepuiana 6a. Flowers sessile to 2 mm pedicellate; style — EPA E of lower leaf surface with larger areoles than on upper surface, subeleva 9 cm long; plants of the Sierra Parima, Terr. Fed. jean a c . Flowers on pedicels 2.5-7 m n upper surface; leaves ai rimensis mm long; style strigillose below the middle and iud the base; reticulation subelevated on lower surface, scarcely evident or obscure on upper se ly leaves 2-7 cm long; plants of sandstone table mountains of eastern F: Edo. Boliv ornatinervia Panopsis cuaensis Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: Río Cuao, Río Orinoco, 125 m, 17 Jan. 1949. Bassett Maguire & Louis Politi 28409 (ho- lotype, NY). Arbuscula, ramis glabris; foliis petiolatis, petiolis 5— 8 mm longis glabris, laminis discoloribus subtus mar- ronino-brunneis oblongo-ellipticis apice acuminatis vel subcuspidatis basi subacutis vel subobtusis 8.5-11.5 cm longis 3-4 cm latis ubique glabris, nervis lateralibus vix bidon utroque latere 9, venulis tertiariis ubique ICHU visis; Vinftuclescentiae rhachidi terminali 15-17 cm longa adpresso-pubescenti pilis pallidis instructa, axibus tri- bus vel quattuor 7-10 mm longis; fructu fusiformi ex- tremitatibus rotundatis 2.2-2.5 cm longo 1 cm lato dense brunneo-velutino. Small tree with glabrous branches. Petioles 5- 8 mm long; leaf blades discolored, dull olive green above, maroon brown beneath, oblong- elliptic, 1 "s bruptiy ac t apex, sub- acute to subobtuse at base, 8. a 11.5 cm long, 3— 4c e side, divaricate at approximately 10-15°, ees and tertiary vena- tion immersed, the tertiary venation finely and minutely reticulate-subimpressed, the midrib el- evated below, shallowly depressed above. Flow- ers not seen. Fruiting rachis terminal, 15-17 cm long, the 3-4 lateral axes 7-10 mm long, the rachis and axes pale appressed-pubescent. Fruit fusiform, rounded at both ends, 2.2-2.5 cm long, 1 cm wide, densely brown velutinous. This species differs from Panopsis rubescens (Pohl) Pittier in its completely glabrous leaves which terminate abruptly in a shortly acuminate or cuspidate apex, fewer and less distinct lateral foliar nerves, glabrous stems, and smaller, short- er fruits rounded at each end. Panopsis parimensis Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: Departamento Atabapo, helechales y for- maciones secundarias en la i 25 km NNE de Parima “B,” cabeceras del Río Ocamo, 3?3'N, 64*13'W, 1,150 m, 12 Jun. 1981, Otto Huber 6136 (holotype, VEN; isotypes, MO, NY). Fr .5 m, ramis novellis dense ferrugineo-to- monies foliis Min ae petiolis 2-8 mm longis fer- rugineo-tomentellis; foliorum laminis elliptico-obova- tis vel elliptico-oblongis apice subacute obtusis basi subacutis vel acutis 4.5—9 cm longis 2-4 cm latis supra floribus sessilibus vel sieur usque . 2 mm longis; perianthio 4-4. 5m o, tepali co ong instructis; stylo 2.3 mm longo glabro. Shrub, 1.5 m tall; young leafy stems densely ferruginous tomentose; mature branches dark gray, glabrous. Petioles 2-8 mm long, ferrugi- nous tomentose; leaf blades elliptic-obovate or elliptic-oblong, subacute-obtuse at apex, sub- acute to acute at base, 4.5-9 cm long, 2-4 cm 1987] wide, finely strigose above with pale hairs, more densely strigose along upper midrib, more dense- ly appressed below, especially along the midrib, with shorter ferruginous hairs; lateral nerves ca. 9 each side, not prominent but more manifest than the tertiary venation; tertiary venation of upper surface minutely and finely reticulate, sub- elevated, impressed on lower surface. Inflores- cences terminal, paniculate, densely ferruginous tomentose, 4-8 cm long (including the peduncle), 5-7 cm wide, with 4 divaricately spreading branches up to 4 cm long, and 1 mm diam. Pe- duncle 8 mm long. Bracts subtending the branch- es of the inflorescence subulate, 4 mm long, densely ferruginous tomentose. Flowers irregu- larly crowded on the axes, solitary or 2-3-fas- ciculate, sessile to 2 mm pedicellate. Perianth 4— 4.5 mm long, the segments densely appressed STEYERMARK — VENEZUELAN GUAYANA FLORA- III 613 pubescent without. Hypogynous disk slightly an- gulate. Ovary ferruginous setose. Style 2-3 mm long, glabrous. This species differs from Panopsis ornatinervia Steyerm. of eastern Venezuelan Guayana in the upper leaf surface having a minute subelevated reticulation and the lower surface a larger areo- lation of impressed veinlets. In P. ornatinervia, the upper surface has a pebbly rugulose, but not reticulate, pattern, whereas the lower surface has a subelevated and finer reticulation. Moreover, in P. ornatinervia the leaves are rounded at the apex, whereas those of P. parimensis are sub- acutely obtuse and usually larger. Finally, the flowers of P. parimensis have shorter pedicels than those of P. ornatinervia and have a glabrous style. ROUPALA KEY TO SPECIES OF ROUPALA la. edi ciel less than 2 m tall; leaves 1.3-3 cm long, 0.7-2.5 cm wide; petioles 1-2 mm long; ovary R. glabro minima lb. Shrub ç or sues 3-20 m tall; leaves 3.5-17 cm long, (2-)2.5-10 cm wide; petioles 10-50 mm long; ovary pubescent 2a. prem rounded or obtuse at apex 3 3a. vigi and rachis of inflorescence glabrous; leaves glabrous below ioles 3-5 mm long; leaf blades 3.5-7.5 c Hs ds 12-15(-30) mm long; leaf blades (5-)7-11(-17) cm long, acute at base .. par pie m long, obtuse at base R. R. obtusa 3b. ee tomentose or furfuraceous-puberulent; rachis minutely ferruginous puberulent or castane 5a Fern the base, 2.5-7 cm ves minutely puberulent or furfuraceous us-furfuraceous or puberulent; le th 7-9 mm lon ne ovary hirsutulous; leaf blades elliptic or ovate-oblong, obtuse at R. sororopana 5b. Perianth 12-13 mm resa ovary shortly appressed-pubescent; lage blades ovate to sub- orbicular-ovate, truncate or broadly rounded at base, 7.5-10 c N leg . Leaves acute to acuminate at a wide ............ R. chimantensis 6 to glabrescent toward apex apex 6a. Stems, petioles, and lower surface of leaf blades glabrous or glabrescent; perianth strigillose R. montana 6b. itn ps and lower surface of leaf blades manifestly pubescent; perianth tomentellose villou 78. Perianth densely pale brown and villous with spreading hairs; rachis of inflorescence t ntose; fruit minutely tomentose 7b. Perianth tomentose; rachis of inflorescence yellow tomentose; fruit glabrous ... Roupala paruensis sd sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: Cerro Parú, be south to southeast to edge descent to tributary of Cano Asisa, rocky sabanita, open Río Ventuari, 2,000 m, 10 Feb. 1951, R. S. Cowan & John J. Wurdack 31378 (holotype, NY). Arbor 7 m, ramis glabris; foliis petiolatis, petiolis 3- a aminis ovato-oblongis apice ob- datis -— obtusis 3.5-7.5 cm longis 2- 3.8(-4) cm glabris; nervis eval supra haud manifestis vel Obsoletis subtus subimpressis vel . griotii R. suaveolens subelevatis utroque latere 4—5; floribus uy ctes rua > cm lo 2-4 mm y ees fructu o c TI 2 lato glabro; sem O aios ORE 1.2-1.4 cm longis 0.8-1 cm latis non vISIS; 1N- Tree 7 m, with glabrous branches. Petioles 3— 5 mm long, glabrous; leaf blades alternate, co- riaceous, ovate-oblong, obtuse or rounded at apex, obtuse at base, ETP m long, 2-3.8(-4) cm wide, glabrous both sides, narrowly subrev- olute, lateral nerves 4—5 each side, obscure or 614 obsolete above, subimpressed or subelevated be- low. Inflorescence 18 cm long, rachis glabrous; pedicels 2-4 mm long, glabrous. Fruit obliquely obovoid, 1.5-2.5 cm long, 1-1.2 cm wide, gla- brous; seeds brown, ovate, rounded at each end, 1.2-1.4 cm long, 0.8-1 cm wide. This species i dily distinguished from Rou- pala obtusata Kl., its closest relative, by the smaller, basally obtuse leaf blades with shorter petioles. NYCTAGINACEAE The genera Guapira and Neea are represented in the Venezuelan Guayana by a large number of taxa. Schmidt (1872) treated the known species of these genera at that time as they occurred in Brazil, Guapira then included within the genus Pisonia. Later, Heimerl (1896) published addi- tions for an account of the West Indian taxa and (1897, 1914, 1932) described various new species from tropical America. Huber (1909) also de- scribed several species of these genera from Am- azonian Brazil. Standley (1931) took up the fam- ily for northwestern South America, recognizing the genus Torrubia of Vellozo with a dozen species previously assigned to Pisonia. Lundell (1968) transferred the taxa formerly assigned to Tor- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 rubia, as well as some others which had been placed under Pisonia, to the genus Guapira Au- blet (1775), an earlier legitimate name. Little (1968) transferred d additional species from Torrubia to Gua Unfortunately, s has been no recent study published for the taxa occurring in the Venezue- lan Guayana. During the many years that have elapsed since the publication of works by Schmidt, Heimerl, and Standley, many unnamed or mis- identified collections have accumulated in her- baria. S large number of the taxa represented by these separated, while oth- ers appear to be closely related and differentiated on characters relating to indument, peduncles, leaf shape, size, venation, branching of inflores- cence, and cauliflory. In some cases it is difficult to be certain of the generic distinction where only pistillate flowers are present. In general, how- ever, Guapira and Neea may be separated using staminate material. Yet Burger (1983) suggested that the two genera may have to be united under Guapira. Further studies will be necessary to judge the merits of generic separation. The present study of the taxa of the Venezue- lan Guayana has resulted in the following 26 new taxa, with keys provided for the species of Gua- pira and Neea. GUAPIRA KEY TO THE SPECIES OF GUAPIRA OF THE VENEZUELAN GUAYANA la. Leaves 1-4 cm long, 0.8-1.8 cm wide, rounded or manifestly obtuse at apex G. oe ylla do lb. Leaves larger than 4 cm long and 1.8 cm wide, mainly acute to acuminate at apex, or, if rounde obtuse, the leaves larger 2a. Leaves 25-30 cm long, 15 cm wide 2 G. sipapoana 3 2b. Leaves 5-15 cm long, 2-15 cm wide 3 a. Lower and/or upper surface of leaves, or lower midrib, pilosulous with lax, spreading, or I divaricate 4a. Up densely per je n pide and shining; inflorescence subglobose or subhemispheric, red and c ted G. davidsei + c cymosely or widely spreading . Upper leaf s Miu Or a d upper midrib, pubescent, and not shining; inflorescence Sa. Petiole 0.5 cm or less long; peduncle 1.5-2.5 cm long 5b. Petiole 1-2 cm long; peduncle (1—)3-9.5 cm lon G. pubescens 6 6a. Peduncle 7-9.5 cm long; petiole densely pubescent with spreading hairs 0.2—0.5 mm long; young stems densely pubescent with spreading hairs 0.2-0.5 mm lon G.m marcano-bertii 6b. eae (1-)3.5-5 cm long; petiole and young stems with hairs less than 0. usbyana w S G.r . Lower —Ó of en including midrib, glabrous or pubescent, but the indument not ate spreading or divaric 7a. Lower surface of leaves glabrous or essentially so, the midrib or nerves with scattered ntu microscopic tom uncle "E petiole densely ferruginous tomentose 8b. Peduncle and petiole glabrous or sparsely cle and/or axes of in reais ae to moderately puberulen Oa. Leaves often broadest above the middle, conspicuously venose; Tu 9a. Pedun l G. amacurensis pubescent 1987] STEYERMARK — VENEZUELAN GUAYANA FLORA —III 615 nerves conspicuous, 9-11 each side, subelevated or impressed on both sides, conspicuously anastomosing with t he tertiary veinlets, ascending at an angle of 45? or more; tertiary veinlets forming a prominent network e G. fragrans 10b. Leaves often broadest near the middle, not venose, opaque; lateral nerves Rr dar 5—6 each side, impressed, divarica seed spreading at an angle G. 15-30°; tertiary veinlets obsolete or E cu guianensis 11 9b. gian: and/or axes of inflorescence glabrous .... us lla. Tertiary veinlets Me and finely pier ee on both sides of leaf blades; eastern Edo. Boliva - G. bolivarensis llb. Tertiary veinlets as i p d Te mazonas . 12 12a. ` Leaves ss to es hee -lanceolate; staminate — cylin- dric-tubular, 1.8-2 mm wide; stamens 8 glabriflora 12b. Leaves ovate or elliptic- jee staminate perianth ere rd mm wide; stamens 10 G. neblinensis 7b. Lower surface of leaves, midrib, or nerves with a minute tomentum of nonspreading S 13a. Tertiary venation on upper and lower leaf surface very conspicuous or elevated ... G. 13b. Tertiary venation either not evident, inconspicuous, or not elevated 14a. Pri ncipal lateral nerves 8-10 e f us to Du ions on biis branches generally cuspidate at G.c apex; dry frui gam — + c the sparsely rufous m sa iesu NT PNE EE 14 ach side; lower leaf surface usually with a ong, 3-4 m uspidata . Principal lateral nerves pi : each side; eae leaf surface glabrous except for idrib a nd sometimes sparsely tomentose or glabrescent secondary nerves; leaves on fertile branches rounded, subtruncate, or abruptly shortly acute at apex; dry fruit 8-9 mm long, 5-5.5 mm wi ide Guapira amacurensis Steyermark, sp. nov. TYPE: Venezuela. pare bosque pluvial, E of Rio El Palmar, near limits of 1965, Luis Marcano-Berti 551 (holotype, VEN; isotypes, MO, NY 5-25 m, ramulis dense-ferrugineo-tomen- tosis; foliis ovatis, elliptico- ovatis vel lanceolato- ellip- costa media subtus sparsim ubique glabris; inflorescentiis dense ferrugi tellis, pedunculis ferrugineo-tomentosis pilis 0.2-0.5 S mentoso; staminibus 7, nd foemineo su extus dense ferrugineo-tomentoso Tree 15-25 m tall, the younger branches densely ferruginous tomentose. Petioles 1-3.5 cm long, densely ferruginous tomentose; leaf blades turning black or dark brown, coriaceous, elliptic- ovate, lance-elliptic, or ovate, acute, obtusely acute to acuminate at the apex, acute to cuneate at the generally inequilateral base, 6.5-14 cm long, 3-6.3 cm wide, glabrous both sides except the midrib on lower side sparsely rufous tomen- tellous or glabrous; lateral nerves 6-11 each side, obsolescent above, slightly more evident below. Inflorescence of staminate plant 2-3 cm high, 4— 5 cm wide, ferruginous tomentose; peduncle 1.7— DENS G. ayacuchae 4 cm long, 1-1.5 mm wide, not enlarged at junc- tion with the lowest inflorescence axes, the fer- ruginous hairs somewhat lax. Staminate flowers sessile to 1 mm pedicellate; bracts 0.5-1 mm long, densely ferruginous tomentose. Staminate perianth infundibuliform, 7 mm long, 3 mm wide bove; stamens 7, the filaments exserted 3-5 mm beyond the orifice. Pistillate perianth tubular, up to 7 mm long, densely ferruginous tomentellous without. Paratype. VENEZUELA. BOLÍVAR: Represa Guri, 55 km NE of Ciudad Piar, 7?35'N, 63?7'W, 200-300 m, 4-5 Apr. 1981, Liesner & Gonzalez 11174. This species differs from Guapira bolivarensis, described below, in the larger staminate perianth, generally inequilateral leaf base, and the dark brown to blackish leaves upon drying. Guapira ayacuchae Steyermark, sp. nov. TYPE: Steyermark & Otto Huber 113856 (holo- type, VEN; isotype, MO) Arbuscula 3-4 m, foliis late ovatis ramulorum tilium. apice late rotundatis subtruncat tis vel ed SE 5- 15 cm | longis 4—9 cm latis, supra costa dies lan lateralibusque ferrugineo- -pubescentibus aliter glabris vel glabrescentibus subtus costa nervis lateralibusque 616 s ferrugineo-pubescentibus inter nervos magis ilabresc en ntibus; nervis iiri adn. utroque latere 6-7; rcte rrugineo-tomentellis, pe- m longo ext rrugineo-tomentello, perianthio PUES tubuloso 2.5-3 mm longo extus ferrugineo- tomentello; staminibus 5; fructibus in sicco ellipso- ideo-oblongis 8-9 x 5-5.5 mm in vivo ovoideis 10 x 9 mm; pedicellis fructiferis 4-5 mm longis Small tree 3-4 m tall. Petioles 1-2 cm long; leaf blades membranous, broadly ovate, on the fertile branches broadly rounded or subtruncate to shortly and abruptly acute at apex, rounded, truncate, or cuneate at base, 5-15 cm long, 4—9 cm wide, the upper surface glabrescent or re- motely and minutely puberulent, the midrib and lateral nerves rather densely rufous-ferruginous tomentose, the lower surface more glabrescent, here the midrib less densely rufous tomentose and the secondary nerves only sparsely tomen- tose to glabrescent; lateral nerves 6-7 each side, Staminate inflorescence umbel- lately 4-branched, the 4 primary axes 8 mm long; peduncle 2 cm long, 0.9-1 mm wide, this and axes of the inflorescence minutely densely ru- fous-ferruginous tomentellose. Flowers on ped- icels 1-1.8 mm long. Staminate perianth infun- dibuliform, 3.5-4 mm long, 2-3 mm wide at summit, moderately rufous-ferruginous without. Stamens 5, filaments 6-7 mm long, exserted 2- 3 mm. Pistillate inflorescence umbellately 4-branched, the primary axes 10-12 mm long, 1 mm wide; peduncle 5 cm long, this and the axes of the inflorescence closely rufous-ferruginous tomentellose. Flowers on pedicels 1-2 mm long, 4-5 mm long in fruit. scene perianth tubular, 2.5-3 mm long, 1-1.5 m , ferruginous to- mentellous without. Fruiting | primary axes 1-2.5 cm long, 1-1.5 mm wide, the secondary axes 8- 17 mm long. Anthocarp black, broadly ellipsoid- oblong in dried state, ovoid in living state, sub- sulcate, glabrous, 8-9 mm long, 5-5.5 mm broad in dried state, when fresh 10 mm long, 9 mm wide. Paratypes. VENEZUELA. TERRITORIO FEDERAL AMAZONAS: alrededores del aeropuerto Puerto Ayacu- cho, 120 m, Morillo 3142 (VEN); Tobogán de la Selva, 5°22'N, 67?33'W, 150 m, 14 May 1980, Steyermark, Davidse & Guanchez 122545 (MO, VEN); 6 km N of ; Estación de Piscicultura, erto Ayacucho 5°37'N, 67?36' , Huber 618, 662 (VEN); Ori- popos, 7 km N of Puerto Ayacucho, Miller 1618 (MO, VEN). COLOMBIA. DEPARTAMENTO VICHADA: Casuarito, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 across from Puerto Ayacucho, 5?40'N, 67°40’W, 100 m, Gentry & Stein 46330 (MO, VEN). This species has been confused with Guapira cuspidata (Heim.) Lundell of northeastern Ven- ezuela. It differs from that species in the larger, glabrous fruit with longer pedicels, 5 instead of 7 stamens in the staminate flowers, generally rounded to subtruncate or abruptly shortly acute apex of the leaves on the fertile branches, fewer and more distantly separated pairs of lateral nerves on the leaf blades, glabrous or glabrescent lower surface of the leaves between the latera nerves, more sparsely puberulent upper leaf sur- face with more conspicuously rufous puberulent midrib and lateral nerves, and fewer ultimate axes of the pistillate infructescences. Guapira bolivarensis um. sp. nov. TYPE: enezuela. Bolívar: Departamento Piar, summit of Amaruay-tepui, south side, east- ern half, 5°55’N, 62°13'W, 950-1,100 m, 11 May 1986, Ronald Liesner & Bruce Holst 20800 (holotype, MO; isotype, VEN). 6 m, ramulis sparsim puberulis pilis dnd neo-glanduliferis munitis; foliis peat ova ice acutis vel acuminatis basi a -] gis ia 8 cm latis utrinque glabris, nervis s Tateralibus utroque i el ulum ad- m bel glabro; perianthio mascu- o 1.8-3.3 cm longo; peri- Mert ‘foemineae peduncu anthi 5 mm longo superne 1.1 mm lato extus glabro. Tree 6 m tall, the young branches sparsely pu- berulent with appressed-ascending, ferruginous- glandular trichomes. Leaves alternate or oppo- site. Petiole 0.7-2 cm long, glabrous to sparsely puberulent; leaf blades elliptic to ovate, acute to acuminate at apex, acute at base, minutely dark- dotted beneath, 6-17 cm long, 3-8 cm wide, gla- brous both sides; principal lateral nerves 8-10 each side, irregularly spaced, subhorizontal or ascending at an angle of 10—25*; tertiary venation finely reticulate both sides, the veinlets promi- nulous. Staminate inflorescence somewhat broader than long, 4-flowered, 2-2.5 cm high, 2.5-4 cm wide, subumbellate to broadly panic- ulate with 3-8 axes, the lower axes larger, gla- brate or sparsely puberulent near the apices; pe- duncle 4-7.8 cm long, 1-1.3 mm broad, glabrous. Staminate perianth narrowly infundibuliform, 4 mm long, 1.5 mm wide at summit, glabrous 1987] without except for a few minute hairs at base. Stamens 10, the filaments unequally exserted up to 4 mm above orifice. Pistillate inflorescence terminal, 3-5 x 3.5 cm, sparsely 3-5-flowered, the 2-3 axes sparsely puberulent; peduncle in anthesis 1.8-3.3 cm long, sparsely puberulent. Pistillate perianth tubular-cylindric, 3-3.5 mm long, 1.1 mm near orifice, glabrous without. Fruiting axes 2.5-5 mm long. Young fruit nar- rowly fusiform, 11-12 mm long, 3 mm wide at middle. Paratype. VENEZUELA. BOLÍVAR: Amaruay-tepui, steep slopes on W side, 5*55'N, 62?15'W, 550-800 m, 20 May 1986, Liesner & Holst 20932 (MO, VEN). This taxon resembles Guapira glabra (Hei- merl) Steyerm., comb. nov. (Pisonia glabra Hei- merl, Kew Bull. 1932: 220. 1932) in the glabrity of the leaves and floral parts, prominently retic- ulate tertiary venation on both leaf surfaces, and subhorizontal to shallowly ascending lateral nerves but differs in having 10 instead of 8 (rarely 9) stamens, short and narrower staminate peri- anth, and shorter staminate inflorescence with shorter axes. Guapira davidsei Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Delta Ama- mento Tucupita, 5-14 km ESE 1979, Gerrit Davidse & Angel González 16382 (holotype, VEN; isotype, MO) Arbor 8 m, foliis oppositis. vel ternatis lanceolato- vel oblongo 12cm lon- C1 OUIOIIBU 1 presso-puberulis costa media pilis patentibus 0.1 mm culo 1-2.5 cm longo dense ferrugineo-adpresso-pubes- centi; floribus 10-35 sessilibus; perianthio masculo p a 4—4.5 mm longo supra medium 2.5- m lato extus adpresso-pubescenti pilis glanduli- feris r mixtis munito. Tree 8 m tall, the younger branches appressed puberulent, the older ones sparsely so. Leaves opposite or 3 at a node. Petioles 3-10 mm long, densely appressed pale puberulent; leaf blades lance-elliptic or oblong-elliptic, acute at apex, acute at base, 6-12 cm long, 2-5 cm wide, gla- brous and shining above, minutely subappressed puberulent below with pale hairs 0.1 mm long, the midrib below elevated, with minute, spread- ing, slightly rigid hairs; lateral nerves 7-9 each STEYERMARK — VENEZUELAN GUAYANA FLORA —III 617 side, inconspicuous, impressed. Staminate inflo- rescence terminal, densely congested, globose or subhemispheric, 8-14 mm high, - mm broad, the primary axes suppressed. Peduncles 1-3, 1-2.5 cm long, with densely ferruginous, subappressed hairs. Flowers 10—35, sessile. Bracts lanceolate-oblong, subobtuse or rounded, 1 mm long, densely t without. Sta- minate perianth infundibuliform, 4—4.5 mm long, 2.5-2.7 mm above middle, densely ferruginous appressed-pubescent without, mixed with glan- dular Aun Stamens 7-8, exserted 2-2.2 mm beyond ori Guapira davidsei is well marked by the con- gested globose inflorescence with dense ferrugi- nous pubescence. From G. ferruginea (Klotzsch ex Choisy) Lundell it is distinguished by having much larger, acutely pointed leaves and numer- ous glandular hairs intermixed with the pubes- cence on the outer surface of the staminate peri- anth. Guapira glabriflora Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: between San Carlos and El Solano, Depar- tamento Río Negro, 11-17 Mar. 1979, Luis Marcano-Berti & P. Salcedo 54-979 (holo- type, MER). rbuscula, ramulis glabris; petiolis 6-10 mm longis; 5 mm ‘longo (in sicco 2.2 mm ) apicem versus 1.8-2 berulo pilis glanduliferis atque apice dense papillato- puberulo aliter glabro; staminibus 8 Small tree. Petioles 6-10 mm long, glabrous; leaves opposite, the blades drying fuscous, ellip- tic-lanceolate to oblanceolate, obtusely acute at apex, cuneately acute at base, 6.5-12.5 cm long, .5-4.7 cm wide, glabrous both sides, the lateral nerves 6-9 each side, inconspicuous, impressed. Staminate inflorescence 10-15-flowered, the pri- mary axes 2-3, 3-15 mm long, bearing the flow- ers directly or with short secondary axes up to 5 mm long bearing a few sessile flowers, the axes minutely and sparsely ferruginous puberulent. Peduncle 1.5-4.5 cm long, glabrous or glabres- cent or with sparse ferruginous glandular or non- 618 glandular hairs. Bracts minute, ovate-deltoid, subacute, 0.3-0.4 mm long, puberulent. Perianth tubular, slightly and inconspicuously enlarged above the middle, 4.5—5 mm long (2.2 mm dried), 1.8-2 mm wide at summit, 1 mm wide (0.6 mm dried) in lower 3, nearly glabrous without except for a few sparse, ferruginous papillate hairs in basal 1 mm and densely papillate-puberulent apically bordering and between lobes. Stamens , the filaments exserted 4 mm beyond orifice. This species is distinguished by its glabrous leaves and nearly glabrous peduncle and stami- nate perianth. It differs from Guapira neblinensis Mag. & Steyerm. in the narrower oblanceolate to elliptic-lanceolate leaves and narrower sta- minate perianth. Guapira marcano-bertii Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Delta Amacuro: east of Rio Grande, ENE of El Palmar, near limits of Estado Bolívar, 26 May 1964, Luis Marcano-Berti 209 (holo- type, VEN; isotype, MO). LE Eg Arb Us amaka 0.2-0.5 mm es munitis; pe- tiolis 1-2 cm -8 incon escencia oo ae A vejer axibus principalibus 2-4, 12- gis 1.5 ia pe edunculo fructifero a 5cm s. 2-2.5 mm s pis e tomentoso pilis patentibus 0.2—0.4 mm longis ne anthocarpio anguste ellipsoideo 9-10 mm longo 3.5—4 mm lato sparsim puberulo 10-costato. Tree with young stems densely villosulous with spreading ferruginous hairs 0.2-0.5 mm long. e 1-2 cm long, densely ferruginous villo- sulous with spreading hairs 0.2-0.5 mm long; leaf blades membranous, broadly elliptic-ovate, shortly and abruptly acute to short acuminate at apex, obtuse to acute at base, 7-12.5 cm long, 3-7 cm wide, the midrib above with spreading villous hairs 0.5-0.7 mm long, the upper surface elsewhere with shorter and longer pale hairs 0.5- 0.7 mm long, the lower surface densely soft-vil- losulous with crisp hairs 0.5 mm or more long, the midrib with divaricate hairs up to 1 mm long. Infructescence subumbellate to irregularly panic- ulately branched, the 2-4 p l mm long, 1.5 mm diam., dud ferruginous villosulous with spreading hairs to 0.2 mm lon Fruiting peduncle 7-9.5 cm long, 2-2.5 mm ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 diam., densely villosulous with spreading hairs 0.2-0.4 mm long. Fruiting bracts lanceolate, sub- acute or obtuse, densely tomentose without. Fruiting pedicels 1-3 mm long. Anthocarps nar- rowly ellipsoid, 9-10 mm long, 3.5—4 mm wide, sparsely puberulent, longitudinally 10-costate. This taxon differs from Guapira pubescens (HBK) Lundell in the much longer petioles and the longer and more abundant pubescence on all parts. From G. cuspidata (Heim.) Lundell it dif- fers in the longer peduncles, longer fruits, and the longer spreading pubescence throughout. Guapira sancarlosiana Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: 4.3 km NNE from San Carlos on Solano Road, IVIC study site, 1°56'N, 67°3'W, 119 m, 8 Jan. 1981, H. L. Clark & Pedro Maquirino 7794 (holotype, MO). Figure 2. Arbor 7-12 m, ramulis juvenilibus dense — seque rufo- ferrugineo- tomentosis; foliorum lamini subtus arcte rufo-tomentosis pilis dense adpressi is om- nino obtectis ovato-ellipticis vel sublanceolato-ellip- ticis apice breviter abrupteque acutis vel breviter acu- minatis interdum rotundatis basi cuneatim acutis 7.5- 14.5 cm longis 4.5-7.5 cm latis; nervis lateralibus ut- roque latere 9-11 subtus elevatis, venulis tertiariis su- pra subtiliter conspicueque elevatis reticulatis subtus mosissima, axibus primariis 0.8—3 cm re pedun- culo terminali 0.2-3 cm longo 1.5-2 mm m. dense 4.8 mm longo sup m 3.8-4 mm lato extus dense rufo-tomentoso; bs 5-6. Tree 7-12 m tall, the young stems densely ap- pressed rufous-ferruginous tomentose. Petiole 1.5-3.5 cm long, densely and minutely appressed rufous tomentose; leaf blades coriaceous, glau- cous above, ovate- to sublanceolate-elliptic, shortly abruptly acute to shortly acuminate at apex, sometimes rounded, cuneately acute and often asymmetric at base, 7.5—14.5 cm long, 4.5- 7.5 cm wide, glabrous or moderately strigillose ulate and elevated. Staminate inflorescence many and closely flowered, much branched, 1.5-3.5 cm high, 3-7 cm wide, the primary axes 0.8-3 cm STEYERMARK 1987] FIGURE 2. Guapira sancarlosiana. long, paniculately branched with 5—6 short axes, earing small clusters of scattered flowers along the length of the axes. Flowers sessile. Perianth shortly infundibuliform to subcampanulate, 4— 4.8 mm long, 3.8-4 mm above middle, densely rufous tomentose without. Stamens 5—6, 3 ofthe filaments exserted 2.2 mm beyond the perianth. VENEZUELA. TERRITORIO FEDERA AMAZONAS: same data as type, 6909 (NY), 7031 (NY), Clark 7251 (MO); Clark & Maquirino 7776 (MO); be- tween San Carlos and Solano, Marcano-Berti & Sal- Es —VENEZUELAN GUAYANA FLORA —III s —A. Flowering hei staminate plant. — rescence.—C. Staminate flower opened. Based on holotyp 619 B. Portion of staminate inflo- mre 126-979 (MER); supra ostium fluminis Casi- uiare, 1854, Spruce 3751 (NY). This species is well characterized by the co- riaceous leaves which are glaucous above and bear densely appressed rufous puberulence be- neath, by the upper and lower leaf surfaces with elevated reticulate tertiary venation, and by the densely branched staminate inflorescences bear- ing numerous flowers with five stamens, of which three are exserted. Spruce 3751 was labeled as 620 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURE 3. Guapira sipapoana. — A. Fruiting branch.—B. Fruit.—C. Apical end of fruit. Based on holotype. an unpublished new species doubtfully referred to Neea (as “Neea? rubiginosa”). Guapira sipapoana Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: eserva Forestal Sipapo, left margin of Rio Sipapo, Bloque 1, May 1971, Carlos Blanco 1158 (holotype, VEN). Figure 3. Arbuscula, ramulis juvenilibus dense rufo-tomen- tosis; foliis elliptico- ovatis apice acuminatis basi ro- tundatis 25-29 cm longis 14.5-15 cm latis supra costa media puberula aliter gla- bris subtus puberulis pilis erectis 0.1-0.2 mm longis praeditis, venulis tertiariis reticulatis subtus promi- ene reticulatis; infructescencia axillari, axibus pri- mariis 1.5-2 cm longis; pedunculo 3.8 cm longo 3 mm lato denos rufo-tomentoso; anthocarpio lineari-ellip- 1987] STEYERMARK— VENEZUELAN GUAYANA FLORA-~III 621 soideo 1.5-2 cm longo 4-5 mm lato rufo-tomentoso nently so with large areoles on lower surface. conspicue 10-costato. Infructescence in the upper axil, the primary axes 1.5-2 cm long, the secondary axes subumbellate, 2.5-3 cm long, branched above into shorter axes 1.5-2 cm long. Anthocarp linear-ellipsoid, 1.5- 2 cm long, 4-5 mm wide, rufous tomentose, 10- costate. Small tree with young branches densely rufous tomentose. Petiole 3.8 cm long, 3 mm wide, densely rufous tomentose; leaf blades very large, elliptic-ovate, shortly acuminate at apex, round- ed or obtuse at the unequal base, 25-29 cm long, 14.5-15 cm wide, upper surface glabrous except This taxon is distinctive in having very large, for the puberulous midrib; lower surface, in- prominently reticulate leaves with loose pubes- cluding midrib, lateral nerves, and tertiary vein- cence on the lower surface and in having long, lets, puberulous with erect, slender hairs 0.1-0.2 pubescent fruits. mm long; tertiary veinlets reticulate, promi- NEEA KEY TO THE SPECIES OF NEEA la. Leaves sessile or nearly so, usually obtuse at base, the petiole to 1 mm long; young stem and petiole hirtellous with subspreading rufous-brown hairs; inflorescence both axillary and terminal on the stem N. ignicola . Leaves petiolate, generally acute to acuminate at base, the petiole 2-35 mm long; young stem and petiole glabrous or pubescent with appressed hairs; inflorescence either cauliflorous on the old stem = 2a. Inflorescence cauliflorous on the old stem .... 3 3a. Leaves oblanceolate-elliptic or je elliptic, 8-12.5(-15) cm long, 2.5-4.5(-5.5)cm ~ UN 3b. Leaves mainly obovate, oblong, or oblong-ovate, 10-30 cm long, 5-16 cm wide ....... 4a. Principal secondary leaf nerves 6-8 each side, slightly elevated on lower surface N. e 5 4b. Principal secondary leaf nerves 8-12 each side, conspicuously elevated on lower surfac Sa. Lower leaf surface and nerves completely glabrous; fruiting peduncle 1-3.5 cm long; flowers sessile; inflorescence dichotomous or with short axes on an elongated rachis wid avidsei 5b. Lower leaf surface glabrous, but midrib and secondary nerves with minute spreading s; flowering peduncle 0.5-0.6 cm long; flowers with pedicels 2-2.5 mm long; inflorescence much branched divaricately N. liesneri 2b. Inflorescence Wigs ducas the stem or its branches Leaves broadly rounded at the apex, obovate or elliptic-obovate N. obovata 6b. Leaves mainly acute to acuminate at the apex, of other shapes than 7 abov 7a. Main secondary nerves of leaf blades 15-25 each side, subhorizontal: or ascending at an angle less than 20°, relatively close together, 3-4 mm ovalifolia 7b. Main secondary nerves of leaf blades generally amis ee 12 each side (fainter inter- mediate nerves may be present), ascending at an angle usually greater than el or if m 4 es 12 nerves or at a smaller angle, then the nerves more than 4 mm apart „uu 8a. Peduncle 6-11 cm lon 9 9a. Stem, peduncle, and axes of inflorescence densely ferruginous tomentose, peri- anth 3 x 1.5 mm, densely ferruginous tomentose N. bernardii 9b. Stem glabrous, peduncle and axes of i nflorescence minutely year Te or PE rescent; perianth 8-9 x 3 mm, glabro grandis 8b. Peduncle 1-5 cm 10 10a. apa and axes of inflorescence glabrous throughout or essentially so ......... 11 —— perianth slightly or moderately ferruginous pubescent toward N. clarkii 11b. Staminate perianth glabrous i p ly put l 12a. Staminate perianth 6-6.5 x 3.5-4 mm; inflorescence paniculately and irregularly Rory oe and a with elongated ra- chis and axes, 6-15 cm long, 5-10 cm wide; peduncle 3-5 mm in diameter; principal se d = nerves 19-12 each side; tertiary vein- lets conspicuously ie and subelevated on upper leaf ay tose robusta 622 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 12b. Staminate perianth 2 x 0.8 mm; inflorescence umbellately or tri- mously branched, 1-2 cm long (or high), 2.5-4 cm wide; 0 diameter; principal ges nerves 5- obsolete er ace r leaf surface lustrous; lower leaf decies with subele- i rl vated, subreticulate tertiary venation more or less manifest; leaves ira or rarely rounded at apex; Jima (pistillate) 1-1 n diameter; leaves ovate to subovate ... N. tepuiensis 13b. Upper kar eres dull; lower leaf surface with obsolescent ertiary venation; leaves obtusely acute to acute at apex; pe- duncle ai 0.8-1 mm in diameter; leaves prua late N. subglabrata 10b. Peduncle and axes of inflorescence i to densely ferruginous or puber- T% sometimes with glandular trichome . Junction of lowest axes of inflorescence with summit of peduncle not ug enlarged, 0.8-2 mm w 15a. Peduncle usually with pai ired ferruginous bra ri cts ry axes of inflorescence 4-7, umbellate; pistillate peri- anth ferruginous tom E main lateral nerves of leaf blades sharply ascending at 50—60? N. bracteosa 16b. Primary axes of inflorescence 3, not umbellate; staminate perianth glabrous except for papillate exterior of lobes; main lateral nerves of leaf blades shallowly ascending at 15-25°. 15b. ea ebracteat N. amaruayensis 17 anth a or glabrescent; bracts mainly pM ex- ou “for the puberulent margins marahuacae N ° . Perianth variously pubescent; bracts ferruginous dorsally and on margins 18a. Ultimate axes of inflorescence racemose with alter- 18b. nately arranged flowers; upper and lower Sei of the leaf blade with minut e, ferruginous indumen 9a. Lower leaf th Z . IM indume eol alc, hioen e much- branched; periant nth and inflo- rescence axes densely ferruginous tomentose with some glandular ie petal to 1.2 cm long .... N. parimensis 19b. Lower leaf surface glabrous or glabrescent; peri- sparsely branched; perianth and inflorescence axes rather sparsely ferruginous tomentose without ong ........... N. tristis Ultimate axes of inflorescence cymose or corymbiform, or with fasciculately arranged flowers; upper midrib of leaf blades glabrous, the lower midrib glabrous or the basal part sparsely puberulent 20 20a. Petiole and upper part of stem with pale, minute, spreading papillalike trichomes; perianth with edu florescence densely ferruginous ba sat fruit 3.5-5 mm sh N. clarkii 21b. Tertiary venation grossly reticulate, subel- evated andm smile on both sides of leaves; axes and peduncle of inflorescence aie rsely to moderately ge scent; antho- carp 6-7 mm wi .N.gi guaiquinimae 14b. edes of lowest axes of inflorescences with summit of peduncle le wide , 2.5-4 mm 1987] STEYERMARK— VENEZUELAN GUAYANA FLORA-III 623 22a. Perianth 3-4.5 mm long, 1.5 mm wide; petiole and young stem N. sparsely puberulent Perianth 6-8 mm mg 2-3.5 mm wide; petiole and young stem 22b. og . sebastianii glabrous or glabresc 23 23a. Tertiary ene manifest, elevated on lower surface, im- pressed on upper surface; flowers in small clusters at or near the end of the ultimate axes N. neblinensis 23b. Tertiary venation obsolete or scarcely manifest; e if 2 mainly scattered along the length of the ultimate m 24 oe Pistillate perianth glabrous without; leaves drying blackish; principal secondary nerves 9-12 on each side N. D NT 24b. Staminate perianth with minute, sparse to modera erruginous indument; leaves drying deri “sasa principal secondary nerves 6—8 on each s "N. mapourioides Neea amaruayensis Steyermark, sp. nov. TYPE: Venezuela. Bolívar: Amaruay-tepui, west side, steep slopes, 5?55'N, 62°15'W 0- 740 m, 2 May 1986, Ronald Liesner & Bruce Holst 20514 (holotype, MO; isotype, VEN). Frutex vel arbor 2.5-5 m; foliis prs dini piee vel lanceolato- sey apice breviter obtuse s 6.5- 15 cm longis (2.5-)3.5-7.5 cm latis ne ruis nervis Enea principalibus utroque 8-10 ad an- gulum 15-25* adscendentibus, venulis tertiariis reti- lati infl iin masculina 12- 21- flora 0. 9-1. 4 sha longa 1.3-2 cm 8 mm longis 0.5 mm eis sparsim minute ferrugineo- ubera inaequaliter insertis; pedu nculo l- 3 cm Jonga sparsim m vel modice ferrugineo-p munito, bibrac- teato, bracteis oppositis oblongo- ovatis vel oblongis- 1 mm dense ferrugineo-puberulis 4 mm spositis; perianthio masculino cylindri- i i 3-3.5 mm i medio 1.5 mm lato extus glabra; staminibus 8 inclusis Shrub or tree 2.5-5 m tall. Leaves mainly op- posite; petiole 7-20 mm long, glabrous; leaf blades elliptic-ovate, oblong- or lanceolate-elliptic, abruptly shortly and ens subobliquely and ob- tusely acute, 6.5-15 cm long, (2.5-)3.5-7.5 cm wide, with dark dots beneath, glabrous both sides; principal lateral nerves 8-10 each side with ad- ditional shorter intermediate ones, anastomos- ing 2-5 mm from margin; tertiary venation re- ticulate, more conspicuous on upper than lower side. Staminate inflorescence relatively small, 0.9-1.4 cm long, 1.3-2 cm wide, 12-21-flowered, the axes unequally inserted, the 3 main ones 5- 8 mm long, 0.5 mm wide, sparsely minutely fer- ruginous puberulent, each axis terminating in 3 shorter axes 0.7-2 mm long, these each sup- porting 2-3 sessile flowers. Peduncle 1—3 cm long, sparsely to moderately ferruginous puberulent, bibracteate with a pair of oblong-ovate or ob- bs obtuse, densely ferruginous bracts 1-2 mm ong and 1 mm wide situated 4 mm above base. Staminate perianth subinfundibuliform-cylin- dric, 3-3.5 mm long, 1.5 mm wide at middle, 1— 1.2 mm wide at summit, glabrous without. Sta- mens 8, included; filaments 0.5-1.5 mm long; anthers suborbicular, 0.6-7 mm long, 0.5-0.6 = Paratype. VENEZUELA. BOLÍVAR: Amaruay-tepul, southwest slope, 750-900 m, 24 Apr. 1986, Holst & Liesner 2702 (MO, VEN). This taxon possesses paired bracts on the pe- duncle, as in Neea bracteata Steyerm., but differs in the perianth of the staminate flower being completely glabrous except for the papillate mar- gins and exterior surface of the shallow perianth lobes. It differs further in the longer petioles, the unequally inserted lower nonumbellate axes of the staminate inflorescences, and the shallowly ascending lateral nerves of the leaves at an angle of 15-25? instead of the sharply ascending ones of N. bracteata arising at 50—60? Neea bernardii Steyermark, sp. nov. TYPE: Ven- ezuela. Bolívar: de Santa Elena de Uairen al sur-este, 800-900 m, 23 Apr. 1957, L. Ber- nardi 6746 (holotype, MER; isotypes, MO, NY Arbuscula vel arbor, m dense ferrugineo-tomen tosis; foliis in sicco brunneis oppositis late ovatis d liptico-ovatis vel oblanceolato- ellipticis apice ke y acutis vel breviter 11.5 cm longis (2.7- )3-6. 5 cm latis subtus costa ui ER ia tere 6-10 ee i cula subcorymbos ae 5c ta chotome ramosa den zou tua rruginea; axibus pri- mariis duobus, (0. 421-3 cm longis 1.5 mm diam. dense 624 adpresso-ferrugineo-t tosi d l -)8-10 cm longo 1.5-2 mm diam. dense. adpresso- EU. tomentoso apice haud manifeste ee perianthio masculino late cylindrico 3-4 mm o 1.5 mm lato extus dense ferrugineo tomentoso; UE 8. Small or large tree to 40 m tall, with densely ferruginous posite; petiole (0.8—)1.5-3 cm long, appressed- ferruginous tomentose; leaf blades drying dull brown to rufous-brown, broadly ovate, elliptic- ovate, or oblanceolate-elliptic, obtusely acute to shortly acuminate at apex, cuneately acute at base, 5.5-11.5 cm long, (2.7-)3-6.5 cm wide, glabrous on both sides except strigose beneath on midrib; lateral nerves 6-10 each side, impressed and in- conspicuous above, faintly manifest below; ter- tiary venation scarcely manifest above, incon- spicuously finely reticulate beneath. Staminate inflorescence terminal, subcorymbose, 1.5-5 cm high, 2.5-10 cm broad, dichotomously me with 2 primary axes (0.4—)1—3 cm long, 1.5 m diam., these branched into 3 Seana sec- ondary axes (3-)12-16 mm long, each with 2 branches bearing 3-7 clusters of sessile or sub- sessile flowers, the axes with densely appressed ferruginous tomentum. Peduncle (2.5-)8-10 cm long, 1.5-2 mm diam., densely appressed fer- ruginous tomentose, its summit 2-3 mm broad, not enlarged at junction with lowest areas. Bracts ovate, acute, | mm long, densely ferruginous to- entose. Perianth broadly cylindric or tubular- subinfundibuliform, 3—4 mm long, 1.5 mm wide, densely ferruginous tomentose. Stamens 8, oc- cupying 7^ of the length of the tube. Fruit oblong, 14 mm long, 7 mm wide, glabrous. iei i VENEZUELA. BOLÍVAR: El dare in margin of Río Grande, 300 m, 20 Feb. 1959, Ber- sees 7156 (VEN); 40 km SE of Tumeremo, nea ath Botanamo, 100 m, 7 May 1960, Little 17597 (US, VEN) Neea bernardii is distinguished by the greatly elongated peduncle. Neea bracteosa Steyermark, sp. nov. TYPE: Ven- ezuela. Bolivar: Represa Guri, 55 km NE of Ciudad Piar, 7?35'N, 63°7'W, 250-300 m, 4-5 Apr. 1981, R. Liesner & A. González 11277 (holotype, VEN; isotype, MO). r8m, ferrugineo-pubescen- tibus; petiolis 3-10 m m longis; foliorum laminis op- positis. vel ad nodos superiores ternatis coriaceis ellip- basi cuneatim acutis 6- 9 cm longis 2-3.5 cm latis, nervis lateralibus utroque latere 8-10 inconspicuis; in- florescentia foeminea umbellatim ramosa 1.5-2 cm alta ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 2-3.5 cm lata, axibus Aia ee 4-7 tenuissimis E 11 mm longis 0.5 mm latis. Pedunculo tenui 1.5- m longo modice ferrugineo- hirtello pilis laxis E g floribus (immaturis) congestis; perianthio (immaturo) modice vel sparsim ferrugineo-adpresso tomentoso. Tree 8 m with young branchlets ferruginous pubescent. Leaves opposite or sometimes ternate at the upper nodes; petiole 3-10 mm long, gla- brous or moderately to sparsely ferruginous pu- bescent at the base; leaf blades coriaceous, dull brown upon drying, elliptic or lance-elliptic, acute to obtusely acute at apex, cuneately acute at the mainly equilateral base, 6-9 cm long, 2-3.5 cm wide; lateral nerves 8-10, fine and slightly evi- dent below, scarcely evident above, arising at an angle of 50-60°. Pistillate inflorescence 1.5-2 cm high, 2-3.5 cm broad, with 4-7 umbellately ies very ne axes 6-11 mm long, 0.5 mm wide, moderately fe t with lax bi hairs 0.1-0.2 mm long; sec- ondary axes 2-5, also umbellate, 3-5 mm long, bearing several clustered flowers at the apices. Peduncle slender, 1.5-3.2 cm long, | mm diam., moderately ferruginous with lax subspreading hairs 0.1—0.2 mm long, bibracteate, not enlarged at junction of summit of peduncle with lowest ally present between '4-'^ the length of the pe- duncle. Flowers (immature) congested, 85-100. Perianth sparsely to moderately ferruginous pu- bescent This taxon may be differentiated by the bi- bracteate, very slender peduncles with slender umbellate axes. Neea b ] l St , Sp. nov. TYPE: Venez uela. Territorio Federal Amazonas: etween Paso El Diablo and Cano de Cu- lebra,25-30 km SE of Puerto Ayacucho, 100 m, 12 May 1980, Julian A. Steyermark, Ger- rit Davidse & Francisco Guanchez 122366 (holotype, pistillate plant, VEN; isotype, MO). Figure 4. AT 6-20 m, ramis glabris, foliis coriaceis opacis obovatis late oblanceolatis vel elliptico-oblongis apice acutis vel incrassatos 1-5 inflorescentiis pedunculatis praedita; pedunculis brevibus 1-1.5 cm longis minute puberulis, 1987] omnibus ramosis cum 3-4 axibus divaricatis subfas- ciculatis vel subumbellatis 9-20 mm longis minute adpressoque puberulis; abend foemineo infundi- buliformi 3. 5 mm longo superne - mm lato extus t .7 mm longo 1. 5 mm lato extus glabro; staminibus 7 inclusis. Tree 6-20 m tall, with glabrous branchlets. Petioles 1-2.5 cm long, glabrous; leaf blades co- riaceous, opaque, obovate, broadly oblanceolate, or elliptic-oblong, abruptly and obtusely acute to rarely rounded at apex, cuneately acute to obtuse at base, 10-22 cm long, 5-12.5 cm wide; lateral nerves 6-8 each side, slightly elevated below, obsolescent above; tertiary veinlets obscure above, slightly evident below, forming large ar- eoles. Pistillate inflorescence cauliflorous, 1.5-2 cm high, 2-4 cm wide, with 1-5 short, many- flowered, pedunculate inflorescences arising from the old wood at the usually thickened nodes. Pe- duncles 1-1.5 cm long, minutely puberulent, each branched into 3-4 widely divaricate, subfascic- ulate or subumbellate primary axes 9-20 mm long, these branched above into short, alternate secondary axes 2-7 mm long bearing the flowers, minutely appressed pale tomentellose. Pistillate perianth infundibuliform, 3.5 mm long, 2 wide, externally minutely puberulent; pistil 3 mm long (ovary 1 mm long; style 1 mm long with prominently penicillate stigmas 1 mm long); ster- ile filaments 0.5-1. long. Staminate flowers in sessile alternately arranged clusters along the short axes of the inflorescence. Staminate peri- anth tubular, somewhat narrowed at base, 3.7 mm long, 1.5 mm wide, glabrous. Stamens 7, included within perianth; filaments 0.5-2 mm long. Paratype. | VENEZUELA. at edge of bana (lower Amaz inga), 67*3'W, 119 m, H. Clark 6605 panna plant, MO). This taxon has been confused with Neea flo- ribunda Poeppig & Endl. but differs in the shorter peduncles with more closely crowded, smaller flowers of a different form and in the glabrous staminate perianth. Neea cedenensis Steyermark, sp. nov. TYPE: Ven- ezuela. Bolivar: Departamento Cedeno: bosque de galeria del Caño Chaviripa del drenaje del Escudo Guayanes, carretera Cai- cara—El Burro, 16 Apr. 1984, B. Stergios, D. Tephorn, L. Nico & C. Gilbert 8611 (holo- type, MO; isotype, PORT) Frutex, ramulis modice puberulis trichomatibus STEYERMARK — VENEZUELAN GUAYANA FLORA-III TERRITORIO FEDERAL 625 brevibus patentibus instructis; laminarum foliis lan- ceolatis vel lanceolato-ellipticis apice anguste subob- tusis basi acutis 8.5-11.5 cm longis 2.2-3.5 cm latis utrinque glabris costa media inferiore prope basem tri umbellata ca. 27-flora inaequaliter ramosa, axibus quinque dense ferrugineo-tomentellis; pedunculo 12 m ferrugineo-puberulo; perianthio mas- m longo dens culino WE Reed 5 Cs 3-4 mm longo superne 1-1.2 mm lato indumento ferrugineo pilis ver- miformibus munito; staminibus 8. Shrub having stems moderately yis with pale, short, spreading trichom .05-0.1 mm long. Petiole with minute, IE short papillalike trichomes; leaf blades mainly oppo- site, lanceolate or lance-elliptic, narrowed to a subobtuse apex, acute at base, 8.5-11.5 cm long, 2.2-3.5 cm wide, 3-3'^ times longer than broad, glabrous on both sides except for scattered pale- erruginous trichomes at base of lower midrib, the upper surface punctate; lateral nerves about 10 and inconspicuous both sides; tertiary vena- tion inconspicuously reticulate. Staminate inflo- rescence terminal, 0.8 cm long, 2 cm wide, ca. 27-flowered, subumbellate with 5 main axes un- equally branched 1-2.5 mm above the base, densely ferruginous puberulent, each axis un- equally divided into 4 branches 1.5-2 mm long, these 1-2-flowered. Flowers sessile. Bracts sub- tending flowers lanceolate, acute, 0.8-1 mm long, unequal, densely ferruginous puberulent. Sta- minate perianth cylindric-subinfundibuliform, 3-4 mm long, 1-1.2 mm wide at summit, abun- dantly streaked with appressed, ferruginous, ver- miform indument without. Stamens 8; filaments 1.5-2 mm long; anthers suborbicular, 0.6 mm long, 0.8 mm wide. This taxon is characterized by short, spreading trichomes on stems and petioles, vermiform fer- ruginous indument on the staminate perianth, small inflorescences, and inconspicuous tertiary venation. Neea clarkii Steyermark, sp. nov. TYPE: Vene- m, Howard L. Clark 6980 (holotype, NY). tex 3 m, ramulis glabris; petiolis 8-15 mm longis glabris; foliorum laminis coriaceis lanceolato- pace dex vel oblanceolato- ellipticis apice acutis vel acuminatis —15) cm longis 25-45 illari trichotome ramosa, axibus dense r ar hasa to- 626 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 RE 4. Neea brevipedunculata. — A. Flowering branch, “seo: plant. view. .—C. Pistillate perianth, mens. mentosis; pedunculo ferrugineo- tomentoso ? apice haud dilatato; p longo supra medium 1.5 mm lato extus minute spar- simque adpresso-puberulo pilis pallidis munito; sta- minibus 8, anthocarpio ellipsoideo 8.5-10 mm longo, 3.5-5 mm lato glabro Shrub 3 m tall with grayish-white, glabrous branches. Petioles 8-15 mm long, glabrous; leaf blades coriaceous, fuscous, lanceolate-elliptic or apical view showing thicken —E. Portion of staminate inflorescence. Based on ho — B. Pistillate perianth, exterior ed perianth lobes. — D. Pistil with rudiments of sta- lot UD. oblanceolate-elliptic, acute to shortly acuminate at apex, cuneately acute at the equilateral base, 8-12.5(-15) cm long, 2.5-5.5 cm wide; glabrous both sides, densely dark punctate below; lateral nerves 8-11 each side, finely impressed or ob- solescent above, faint or finely elevated below; tertiary venation inconspicuous. Staminate in- florescence terminal or axillary (immature?), tri- chotomously branched, 10 mm high, 1 STEYERMARK 1987] — VENEZUELAN GUAYANA FLORA-III 627 wide, shortly pedunculate; 3 primary axes 2 mm long (immature), ferruginous tomentose, termi- nating in groups of 3 or more sessile flowers. Bracts ovate-deltoid, subacute, 0.3-0.4 mm long, densely ferruginous tomentose. Peduncle 4-15 (-30) mm long, 1-1.5 mm wide, ferruginous to- mentose, not enlarged at junction of peduncle with lowest axes of inflorescence. Staminate peri- anth subinfundibuliform, 4 mm long, 1.5 mm wide above the middle, minutely sparsely pu- bescent with pale hairs without. Stamens 8, in- cluded, unequal; filaments 1.3-2.2 mm long. Fruit ellipsoid, 8.5210 mm long, 3.5-5 mm wide, gla- brous. Paratypes. "VENEZUELA. TERRITORIO FEDERAL AMAZONAS: San Carlos, 21 Mar.-17 Apr. 1981, Delas- cio, Christensen & Broome 9496 (VEN); 4 km NE of San Carlos, Liesner 6422 (MO, VEN); prope San Car- los and Río Negro, 1853, Spruce 3766 (NY); San Carlos de Río Negro, 16 Feb. 1981, Otto Huber 6067 (VEN). This species is related to Neea guaiquinimae, from which it differs in the absence of glandu- larity on the peduncles and axes of the inflores- cence, the lack of tertiary reticulation on the leaves, and in a denser indument on the peduncle and inflorescence axes. Huber's specimen matches the photographs of two unpublished names by Heimerl of a Spruce 3766 collection from **prope San Carlos, ad Rio Negro." The photograph from the Vienna her- barium has a note by Heimerl pertaining to a pistillate plant, *Insbesonders durch die Kráftig berandeten Blatter u. wahrscheinlich auch durch Cauliflori ausgezeichnet." This photo shows a cauliflorous, dense, short inflorescence only 10 mm high and 8 mm broad. The other photo from the Berlin herbarium does not show any inflo- rescence. The specimen of Huber 6067 also shows a small inflorescence 10 mm high and 8-12 mm broad. Of the three small inflorescences present, one appears at the apex ofa leafless short branch, whereas the other two originate 5 mm below the apex of a leafless stem. The lower leaf surface of the Huber 6067 col- lection is densely punctate, but the other collec- tions assigned to Neea clarkii do not show punc- tation. Neea davidsei Steyermark, sp. nov. TYPE: Ven- ezuela. Territorio Federal Amacuro: Depar- tamento Tucupita, mountain area, ca. 13 km by road ESE of town of Sierra Imata, 8?23'N, 62°23'W, 400 m, 4-6 Apr. 1979, Gerrit Da- vidse & A. González 16558 (holotype, VEN; isotype, VEN) bor 8 m, ramulis glabris; petiolis 1-2 cm longis Ar glabris; foliorum laminis oppositis late obovatis vel vel acutis lo ong 1AW DUIUI ull pedunculata, pu es pilis adpressis hivis munitis, primariis 3 cm longis superne ue 5mm longis pss lo vetere fructifero 1-3.5 cm es 3m .; bracteis ovatis subacutis 0.5 mm longis extus sparsim adpresso puberulis; fl 1.2 cm longo 0. 8-1 cm lato haud costato. Tree 8 m, the branches glabrous. Leaves op- posite; petiole 1-2 cm long, glabrous; leaf blades broadly obovate or broadly oblong-elliptic, nar- rowed to a subacutely obtuse or acute apex, cu- neately acute at base, 11-24(-30) cm long, 6.5- 1(-16) cm wide, wholly glabrous; lateral nerves 8-10 each side, ascending at an angle of 35—45*, indistinctly anastomosing 10-15 mm from mar- gins, elevated and somewhat prominent below, obsolescent or faintly impressed above; tertiary veinlets inconspicuous, grossly reticulate below, obsolescent above. Pistillate inflorescence axil- lary, cauliflorous on the old wood, pedunculate or epedunculate. Flowers sessile. Old fruiting pe- duncle 1-3.5 cm long, 3 mm diam. Primary axis on old inflorescence 3 cm long, branched in up- per !^ with alternate short axes 5 mm long, all axes minutely and sparsely puberulent with pale appressed hairs. Anthocarp 1.2 cm long, 0.8-1 cm wide, not ribbed. Paratypes. VENEZUELA. BOLIVAR: between km 11 and 18.5, south of El Dorado, 215 m, 23 July 1960, Stevermark 86585 (NY, VEN). TERRITORIO FEDERAL TA AMACURO: Orocoima, Rio Toro, Bernardi 7573, pe 3734 (NY). The axillary, cauliflorous infructescence re- lates this species to Neea liesneri of Cerro de La Neblina, but the former differs in the longer, stouter peduncle, sessile flowers, and completely glabrous lower leaf surface. The fruiting paratype has the leaves broadly oblong-elliptic and acute, whereas in the holo- type the leaf blades are broadly obovate nar- rowed to a subacutely obtuse apex. Neea guaiquinimae Steyermark, sp. nov. TYPE: Venezuela. Bolívar: Cerro Guaiquinima, summit, NE sector, 6?00'N, 63?28'W, 1,650 m, 9 Apr. 1979, Julian A. Steyermark, G. 628 C. K. & E. Dunsterville 117971 (holotype, VEN; isotype, MO). Arbuscula vel frutex 2-3 m, ramis glabris; petiolis 5-20 mm longis. minute eine vel glabris; fo- liorum a basi subacutis vel acutis 5-12. 5c cm longis 2-6 cm latis supra glabris subtus glanduloso-punctatis, nervis lateralibus utroque latere venulis tertiariis ubique manifeste gros- seque reticulatis; inflorescentia foeminea trichotome ramosa | cm alta 1.8 cm lata, axibus principalibus quattuor 4-5 mm longis 0.8 mm diam., modice glan- jene ae sub anthesi 2.5 cm longo | mm dia b fructu 5-7 cm longo 1.5-2 mm lato, sparsim glandulifero: “s 9 ea tubuloso 4 mm longo 1.2 mm lato extus dense glandulifero. Anthocarpio oblongo- ellipsoideo ina ovato-oblongo 12-13 mm longo 6-7 mm lato obtuse 10-costato glabro. vv UU lU SK ALULIS Small tree or shrub 2—3 m tall, the branchlets glabrous. Petioles 5-20 mm long, glanduliferous or the glands deciduous; leaf blades coriaceous, elliptic-ovate, obtusely acute at apex, subacute to cuneately acute at base, 5-12.5 cm long, 2-6 cm wide, glabrous above, densely dark gland- dotted below, slightly revolute on margins; lat- eral nerves 6-10 each side, slightly impressed or obsolescent above, slightly elevated below; ter- tiary venation non both sides subelevated, grossly reticulate. P branched, 1 cm high, 1.8 cm broad, with 4 pri- mary axes 4-5 mm long, 0.8 mm diam., mod- erately glanduliferous, the primary axes shortly branched into 4 shorter, moderately glandulifer- ous secondary axes 1-2 mm long terminating in shortly pedicellate extensions, each bearing 2-3 sessile flowers. Peduncle in anthesis 2.5 cm long, | mm diam., sparsely glanduliferous in fruit, 5— 7 cm long, 1.5-2 mm wide and glabrous, pendent in fruit, the junction of peduncle with lowest axes not conspicuously enlarged. Bracts narrowly lan- ceolate, 1 mm long, densely glanduliferous. Pis- tillate perianth tubular, 4 mm long, 1.2 mm wide, densely glandular without; ovary ellipsoid, 1.3 mm long, sessile; style 1 mm long; stigmas 0.4 mm long, fimbrillate. Anthocarp oblong-ellip- soid to ovate-oblong, 12-13 mm long, 6-7 mm wide, obtusely 10-costate. Paratype. | VENEZUELA. BOLIVAR: Cerro Guaiquini- ma, W sector near forested border, 5?45'N, 63°43' W, 1,540 m, Steyermark, Berry, G. C. K. & E. Dunsterville 117499 (MO, VEN). This species is distinguished by the sparsely to moderately glandular pubescence of the peduncle and inflorescence axes and by the manifestly sub- elevated, grossly reticulate tertiary venation on both leaf surfaces. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Neea huachamacarae Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: Río Cunucunuma, base of trail to Cerro Huachamacari, 3?39'N, 65?41'W, 220 m, 7 Mar. 1985, Ronald Liesner 18403 (holotype VEN; isotype, MO). r 10 m, ramis glabris; petiolis 1-2.5 cm longis Mes "foliorum laminis oppositis oblanceolatis vel oblongo-ellipticis apice acutis obtusis vel rotundatis basi cuneatim acutis 6-15.5 cm longis 3-5.5 cm latis glabris; nervis lateralibus utroque latere 9-12 incon- spicuis; inflorescentia foeminea terminali, axibus dense ferrugineo-adpresso-pubescentibus, pedunculo 1-2.5 cm longo 2 mm lato autem i durato-tubuloso 7.5-8 mm longo 3-3.5 mm lato extus glabro Tree 10 m tall with glabrous branches. Leaves opposite; petiole 1-2.5 cm long, glabrous; leaf blades opaque, drying blackish, oblanceolate or oblong-elliptic, shortly acute, obtuse, or rounded at apex, cuneately acute at base, 6-15.5 cm long, 3-5.5 cm wide; lateral nerves 9-12 each side, not elevated, i spicuous. Pistillate e terminal, the usually 4 primary axes 5-13 mm long, branched into 10-15 secondary axes, the solitary flowers or groups of flowers borne along the length of the axes, not congested in groups at the ends of the axes, the axes densely ferruginous appressed-pu- bescent. Peduncle 1-2.5 cm long, 2 mm wide except at the junction of the lowest axes of the inflorescence with the top of the peduncle where enlarged to 3-4 mm wide. Perianth subpandu- rate-tubular, 7.5-8 mm long, 3-3.5 mm wide, glabrous without. Ovary 2-3 mm long; style 5- 6 mm long, exserted 2 mm; sterile stamens 6 with filaments 3 mm long CI tiar y venation incon- This species is characterized by having the flowers inserted along the length of the axes of the inflorescence rather than in congested groups at the ends of pedicellate branches. In this respect it resembles Neea mapourioides but differs in the glabrous pistillate perianth and the greater num- ber of principal secondary nerves of the leaf blades. Neea ignicola Steyermark, sp. nov. TYPE: Ven- ezuela. Territorio Federal Amazonas: 11 km N of Puerto Ayacucho toward El Burro, 5?44'N, 67°30'W, 90 m, 26 Jan. 1978, Otto Huber & Julio Cerda 1456 (holotype, VEN). Frutex 2-3 m, ramulis dense hirtellis pilis patentibus 1987] rufo- brunneis munitis; ed vix dici usque | mm eis munito; foliorum la- lateralibus utroque later RÀ eii reticulatis; inforeseentia mascula terminali axillarique 5-13 mm alta 7-10 mm la axibus dun ae alternatim axibus lanceolatis 1-1.5 mm longis; pedunculo m longo 0.4-0.5 mm lato hirtello pilis rufo- brunneis patentibus munito; perianthio masculo (im- maturo) tubuloso extus sparsim hirtello pilis ferrugi- neis munito; staminibus 6-7 Shrub 2-3 m tall; branches slender, 1-1.5 mm diam., densely reddish-brown hirtellous with spreading hairs 0.5 mm long. Petiole scarcely developed, up to 1 mm long, hirtellous with lax, subspreading hairs; leaf blades subsessile, mem- branous, ovate-lanceolate, usually narrowed to an acuminate apex, obtuse or rarely rounded at base, 3.5-6 cm long, 1.3-2.5 cm wide, glabrous both sides; principal lateral nerves 10 or some- times more on both sides, elevated above, less conspicuous beneath; tertiary venation reticulate both sides, less so beneath. Staminate inflores- cence axillary and terminal, 5-13 mm high, 7- 10 mm broad, densely hirtellous with rufous- brown spreading hairs; the 4 primary axes alter- nately and racemosely branched with flowers solitary along the length of the axes, but with terminal flowers in 2s or 3s, the secondary axes scarcely or not developed. Peduncle filiform, 8— 14 mm long, 0.4-0.5 mm diam., hirtellous with rufous-brown spreading hairs, not enlarged at apex where joining the lowest axes of the inflo- rescence. Bracts subtending base of primary axes lanceolate, subacute, 1-1.5 mm long. Staminate perianth (immature) tubular, sparsely hirtellous with ferruginous hairs. Stamens 6-7 This taxon is readily differentiated by its sessile or subsessile leaves and spreading pubescence. Neea liesneri Steyermark, sp. nov. TYPE: Vene- zuela. Territorio Federal Amazonas: De- partamento Río Negro: Cerro de La Neblina, 0.4 km W of base camp on Río Mawari- numa, 0?50'N, 66°10’W, 140 m, 9 Mar. 1984, Liesner 16502 (holotype, VEN; isotype, MO). Figure 5. 5 m; petiolis 1-2 cm longis; foliorum laminis late obovatis vel lato oblongo-ovatis 2d subacutis obtusis vel rotundatis basi acutis majo asymme- tricis 16-30 cm longis 9-16 cm latis Men vila sub- STEYERMARK — VENEZUELAN GUAYANA FLORA —III 629 tus costa media nervis lateralibusque elevatis promi- tantibus 12-27 m pressis munitis; peduncu dense pu- berulo pilis adpressis munito; floribus pedicellatis, pe- dicellis 2-2.5 mm longis dense puberulis pilis adpressis obtectis; perianthio urceolato 6 mm longo basi 4 mm lato apicem versus 1.5-2 mm lato extus dense puberulo pilis adpressis obtecto; staminibus 7-9; anthocarpio oblongo-ellipsoideo 20 mm longo 8 mm lato minute pubescenti pilis subferrugineis adpressis obtecto. Tree 5 m tall. Petioles 1—2 cm long; leaf blades broadly obovate or broadly oblong-ovate, sub- acute to obtuse or rounded at apex, acute at base, the larger leaves conspicuously asymmetrical, the smaller leaves nearly symmetrical, 16—30 cm long, 9-16 cm wide, glabrous above, the midrib and lateral nerves beneath prominent and elevated with minute spreading pubescence, elsewhere glabrous; lateral nerves 9-12 each side, ascending at an angle of 35-45?, anastomosing 5-13 mm from the margin, regularly 12-27 mm equidis- tant or irregularly spaced, conspicuously elevat- ed below, inconspicuously impressed above; in- termediate and tertiary venation below inconspicuously grossly reticulate, inconspic- uous above. Staminate inflorescences cauliflo- rous on the old wood, broader than high, 2.5 cm long, 5 cm broad, much branched, many-flow- ered, the lateral primary axes divaricate, 1-2 cm long, all parts densely puberulent with pale-ful- vous, appressed hairs. Peduncle 5-6 mm long. Staminate flowers on pedicels 2-2.5 mm long and covered with a pale-fulvous, appressed pu- berulence. Staminate perianth urceolate, 6 mm long, 4 mm wide at base, 1.5-2 mm wide near summit, densely puberulent with pale-fulvous, appressed hairs. Stamens 7-9, the anthers 1.3-2 mm Ros apiculate or exapiculate, the shorter filaments 1-2 mm long, the longer ones 3-4 mm grad Pstillod 4—5 mm long. Anthocarp oblong- elli , 20 mm long, 8 mm wide, minutely Benin with appressed, subferruginous hairs. Paratype. BRAZIL. AMAZONAS: Between Maloca and Rio md south of Cerro de La Neblina, 50-100 m, Nilo T. Silva & Umbelino Brazáo 60782 (MO, NY). This remarkably distinct species is character- ized by the cauliflorous, many-branched, cymose inflorescence developed on the old wood; large, obovate, conspicuously nerved leaves; and large, pedicellate, urceolate staminate flowers with strongly developed asymmetrical bases. 630 FIG Neea liesneri.—A. Portion d ees branch of staminate plant. Pistillodium with stamens. Based on holoty Neea mapourioides Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: .5-1.5 km east of San Carlos, 1?55'N, 67°5'W, 120 m, 1 Dec. 1977, Ronald Liesner 4096 (holotype, VEN; isotype, MO) ex 3 m, ramulis glabris; petiolis 5-15 mm longis glabris; foliorum laminis oppositis elliptico- Medio 3.5— 7.5 cm longis 4—7 cm latis ubique glabris; nervis la- cate principalibus utroque latere 6-8; inflorescen- tia mascula terminali 2.5-4.5 cm alta 3.5-7 cm lata, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 — B. Staminate flower. — C. axibus late divaricati il 4, dense fer- rugineo o-tomentosis, axium ultimorum fa alter- -2 o 1.5-2.5 cm lon angustato; staminibus 7 ea TR Shrub 3 m tall; branches glabrous. Leaves op- posite; petiole 5-15 mm long, glabrous; leaf blades 1987] coriaceous, opaque, fuscous, elliptic-obovate, abruptly acuminate at apex, cuneately acuminate at base, 5.5-17.5 cm long, 4-7 cm wide; lateral nerves 6-8 each side, anastomosing 4-6 mm from margin, impressed and slightly manifest below, but essentially obsolescent; tertiary venation ob- solete above, scarcely impressed beneath. Sta- minate inflorescence terminal, 2.5-4.5 cm high, 3.5-7 cm wide, the axes widely spica dense- ly ferruginous tomentose, the 3-4 primary axes 1.5-3.5 cm long with the diea part bearing flowers, the ultimate branches bearing sessile flower clusters scattered alternately ita the length; peduncle 1.5-2.5 cm long, 1.5-2 mm wide, but at the summit with the lowest axes s of the inflorescence enlarged to 2.5 mm wide, sparsely to moderately minutely brown puberulent. Bracts of base of primary axes 1-1.5 mm long, ferru- ginous tomentose, those subtending the flowers ovate-deltoid, subacute, 0.5-0.8 mm long, fer- ruginous tomentose. Staminate perianth narrow- ly ellipsoid, 6.5-7 mm long, 2.8-3 mm wide, narrower at summit, mie to moderately mi- nutely ferruginous puberulent. Stamens 7, in- cluded within the lower '4—'/ of the perianth tube, unequal; anthers 1.5 x 0.5-1 mm; filaments 1— 3 mm long. Stylopodium 4 mm long. This species possesses alternately arranged flowers on the axes of the inflorescence as in Neea huachamacarae but differs in the fewer lateral nerves, the fuscous-brown instead of blackish leaves when dried, and in having pubescence on the staminate perianth, as opposed to the gla- brous pistillate perianth in N. huachamacarae. Neea marahuacae Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: Cerro Marahuaca, forested slopes 1-2 km N of Sima Camp, 3?43'N, 65?31'W, 1,100 m, 8—9 Mar. 1985, Ronald Liesner 18428 (ho- lotype, VEN; isotype, MO). x2m, ramulis glabris; petiolis t= 20 mm longis; ferrugineo aarin erigi tenui 1.5-2 cm longo .5-0.7 m sim puberulo pilis adpressis munito; oot ok ae tubuloso 1.5 mm longo mm lato extus glabro vel glabrescenti; staminibus 5-6 inaequalibus inclusis. Shrub 2 m. Young stems glabrous. Petioles 7— 20 mm long; leaf blades opaque, elliptic-oblan- STEYERMARK— VENEZUELAN GUAYANA FLORA- III 631 ceolate, acuminate at apex, acute to acuminate at base, 7.5-10.5 cm long, 2.5-4.5 cm wide, gla- brous; lateral nerves 6-8 each side, inconspic- uous; tertiary venation obsolete or inconspic- uous. Peduncle slender, 1.5-2 cm long, 0.5-0.7 mm diam., simple or forked near base, sparsely appressed-pubescent with reddish-brown pubes- cence, not enlarged at junction of summit with lowest axes of inflorescence. Staminate inflores- cence terminal, subumbellately 5-branched, 5-8 mm high, 10-15 mm wide, sparsely rufous-brown appressed-puberulent; primary axes 2-3 mm long; secondary axes 0.5-1 mm long, each terminating in branched congested clusters, each of these bearing 4—5 flowers. Bracts at base of flower clus- ters ligulate-lanceolate, acute, 1.2 mm long, 0.3- .4 mm wide, glabrous except for the minutely puberulous margins near the apex. Staminate perianth tubular, 1.5 mm long, 0.7 mm wide, glabrous or glabrescent without. Stamens 5-6, unequal, included; anthers suborbicular, 0.1 mm long (immature). Paratype. VENEZUELA. TERRITORIO FEDERAL AMAZONAS: Cerro Marahuaca, Sima Camp, S-Central portion of forested slopes along east branch of Cano Negro, 3*43'N, 65°31'W, 1,140 m, 21-22, 24 Feb. 1985, Steyermark & Holst 130545 (MO). This montane species of Cerro Marahuaca dif- fers from related species in the glabrous or gla- brescent staminate perianth and in the mainly glabrous bracts, except for the apical margins. The paratype differs from the type collection in having ovate-elliptic to elliptic-obovate leaves, which are larger, indistinctly nerved, and sub- acutely obtuse at the apex. Neea parimensis Steyermark, sp. nov. TYPE: Ven- ezuela. Territorio Federal Amazonas: Sierra arima, Simarawochi, Río Matacuni, one 64°36’W, 6-7 km west of Venezue- iom ngu 795-830 m, 18-23 ; an A. Steyermark 106982 dudas E isotype, NY). Arbor 10 m, ramulis dense adpresso- -pubescentibus; rugineo-tomentosis cum pilis glandulosis interspersis; pedunculo usque 12 mm longis vel nullo dense fer- rugineo-tomentoso cum pilis glandulosis interspersis; 632 bracteis cp pn quattuor cupulam facientibus a ovatis ongis extus dense ferrugineo-tomen- tosis; rans masculo urceolato 4 mm longo > 5 mm lato extus dense ferrugineo cum pilis denials interspersis; staminibus 7 inclusis, filamentis 1.2 mm longis. Tree 10 m tall with densely appressed-pubes- cent stems. Petioles 5-8 mm long, densely ap- pressed-pubescent; leaf blades membranous, ob- lanceolate, acute at apex, narrowed to an acute base, 4-7 cm long, 1.5-2.5 cm wide, the upper surface glabrous, the lower surface densely ap- pressed-puberulent with minute scalelike hairs; lateral nerves about 8 each side, scarcely evident. Staminate inflorescence terminal, 4 cm high, 5 cm broad, much branched, the flowers alter- nately arranged on ina numerous axes, the 4-8 principal axes unbranched in the basal 8-13 mm, then forked into alternately or fasciculately dis- posed secondary axes, densely ferruginous to- mentose mixed with glandular hairs. Peduncle none or up to 12 mm long, densely ferruginous tomentose with intermixed glandular hairs. Bracts at base of flowers 4, forming a cupule, broa ovate, subacute, 0.7 mm long, densely ferrugi- nous tomentose without. Staminate perianth ur- ceolate, 4 mm long, 2.5 mm wide at the middle, constricted below the summit, densely ferrugi- nous tomentose without, mixed with short glan- dular hairs. Stamens 7, included within the lower half of the perianth tube; filaments 1.2 mm long. This species is characterized by having a broadly urceolate staminate perianth, much- branched inflorescence, lower leaf surface with a dense appressed indument, and a short peduncle. Glandular hairs are scattered on the perianth, peduncle, and axes of the inflorescence. Neea robusta Steyermark, Sp. nov. TYPE: Vene- 0 m, 14-28 Feb. Otto Huber 1693 (holotype, VEN). 1978, Arbor 3 m, Mona: glabris; petiolis 1-3.5 cm longis glabris; foliorum laminis obovatis elliptico- a elliptico- oblongis mus aiu ui QAI apice pleri que brevit basi cuneatim acutis 8-24 cm e glabri 5. 4.5-9.5 cm latis cm longa 5-10 cm lata oppositis vel alternis, axis parte apicali minute ad- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 presso- -ferrugineo puberula axibus aliter glabris, flori- extus glabro; staminibus 9-10 inclusis; pistillodio 3 mm longo; infructescentia 6-15 cm longo 7-15 cm lato; pedunculo fructifero 1-6 cm longo 3-5 mm lato ad apicem dilatato; anthocarpio in sicco 1.2 x 0.5 cm in vivo 2 x 1 cm. Tree 3 m tall, the branches glabrous. Petioles 1-3.5 cm long, glabrous; leaf blades coriaceous, obovate, elliptic-lanceolate, elliptic-oblong, or oblong-elliptic, mainly shortly and obtusely acute or obtuse at apex, sometimes rounded, cuneately acute at the base, 8-24 cm long, 4.5—9.5 cm wide, glabrous both sides; lateral nerves 9-12 each side, slightly elevated on upper surface, less conspic- uous and impressed on lower surface, anasto- mosing 5-10 mm from the margin; tertiary ve- nation subelevated and conspicuously reticulate on upper surface, less conspicuously reticulate on lower surface. Staminate inflorescence elon- gated, subpyramidal, paniculately branched, 6- 15 cm long, 5-10 cm wide, the upper axis longer than the lateral ones, 6-9 cm long, 6 mm diam., the 4-8 lateral axes divaricately spreading, 0.3- 2 cm long, 2-3 mm diam., the upper ones gen- erally shorter and alternate, the lower axes long- er, opposite or alternate, glabrous except mi- nutely appressed ferruginous puberulent apically. Peduncles in anthesis 1-1.2 cm long, in fruiting stage 1-6 cm long, 3-5 mm diam. Flowers sessile and scattered but more congested apically. Bracts subtending flowers deltoid, obtuse, 0.2 mm lon Staminate perianth carnose, suburceolate, nar- rowed and contracted below summit, rounded at base, 6-6.5 mm long, 3.5-4 mm wide, narrower at summit. Stamens 9-10, included, unequal; an- thers broadly rhomboid; filaments 2-4.5 mm long. Pistillode 3 mm long. Infructescence 6-15 cm long, 7-15 cm wide. Anthocarp 1.2 x 0.5 cm in dried state, 2 x 1 cm fresh. Paratypes. | VENEZUELA. TERRITORIO FEDERAL AMAZONAS: Cano Yagua at Cucurital de Yagua, 3?36'N, UEM, 120 m, Davidse et al. 17362 (MO, Mp año Yagua, Chipital, 3°29'N, 66°41'W, 120 m, vide et al. 17320 (MO, VEN); Cucurital de eon ño Caname, 3?40'N, 67°22'W, 100 m, Davidse et al. 16880 (MO, VEN); 1-2 km SE and E of San Carlos de Yos Lo 1°56’N, 67°3'W, 120 m, Liesner 6868 (MO, N); m NE of San Carlos on Solano Road, Mm un (MO, NY, VEN); 4.3 km NE of San Carlos on Solano Road, Clark 7038, 7164 (NY); S and SW of San Carlos de Río Negro, Liesner 6732 (MO, VEN); Huber Pn al. 374 (VEN); NE and E base of Cerro 1987] Cucurito, 120 m, Huber & Tillett 2973 (VEN); 12-15 km NE of San Carlos de Rio Negro, road to Solano, 100 m, Morillo et al. 4170 (VEN). BRAZIL. AMAZONAS: Serra de Neblina, between Maloca and Rio Cauaburi, 50-100 m, Silva & Brazáo 60765 (MO, NY) This species is characterized by the large, ur- ceolate, glabrous staminate perianth, the irreg- ularly paniculately branched, large inflorescence and general glabrity of parts. The more numerous secondary nerves, thick peduncle, and conspic- uously reticulate tertiary venation of the upper leaf surface distinguish it from related species of the Venezuelan Guayana. Neea sebastianii Steyermark, sp. nov. TYPE: Ven- ezuela. Territorio Federal Amazonas: high rebalsa (seasonally flooded forest), Isla Se- bastian, Rio Casiquiare above Chapezon, between Boca and Solano, 1?58'N, 67?3'W, 120 m, 31 Jan. 1980, Ronald Liesner & Howard Clark 8959 (holotype, VEN; iso- type, ) rbor 8 m, ramulis juvenilibus dense miniteque ad- MSS -puberulis pilis ferrugineis antes petiolis 3- foliorum laminis ova- to- ellipticis apice obtuse acutis vel ndis basi cu- neatim acutis 7.5-14 cm longis 3.5-6.5 cm latis supra glabris subtus praesertim costa media nervisque mi- nute puberulis aliter pagina inferiore minute puncti- culatis atque glandulis sessilibus obtectis; nervis late- ralibus utroque latere 10-12; inflorescentia mascula 1— 2.5 cm alta 2-2.5 cm lata, axibus 4—5 dense breviter ferrugineo-tomentosis 3-5 mm pedunculo 1. m 2.5 cm longo 1-2 mm m 2.5-4 m maturo) dense ferrugineo-tomentoso; staminibus 6-8 inclusis. Tree 8 m, the younger stems with a dense, minutely appressed ferruginous puberulence. Petiole 3-15 mm long, sparsely puberulent; leaf blades subcoriaceous, ovate-elliptic, obtusely acute, or rarely obtuse or rounded at apex, cu- neately acute at the slightly asymmetric base, 7.5-14 cm long, 3.5—6.5 cm wide, glabrous above, below minutely puberulent, especially the midrib and nerves, with erect hairs and also with sparse sessile glands on the leaf surface; lateral nerves 10-12 each side, impressed below. Staminate in- florescence terminal, 1-2.5 cm long, 2-2.5 cm wide with 4—5 axes 3-5 mm long, densely fer- ruginous tomentellose. Bracts subtending flowers ovate, 0.5-1 mm long, densely ferruginous to- mentose. Peduncle 1.5-2.5 cm long, 1- m diam., enlarged to 2.5-4 mm wide at the junction above with the lowest primary axes. Perianth STEYERMARK — VENEZUELAN GUAYANA FLORA-III 633 suburceolate, 3-4.5 mm long, 1.5 mm wide (im- mature), densely ferruginous tomentose. Sta- mens 6-8, included; anthers rhomboid, 0.8-1 mm long This species is related to the other newly de- scribed species, Neea neblinensis Maguire & Steyerm., N. huachamacarae Steyerm., and N. mapourioides Steyerm., but has smaller peri- anths and has sparsely puberulent petioles and young stems. Neea iia . sp. nov Venezuela. Bolivar: between Betania iid ae = de ed 40 km west of Santa Elena, 4?35'N, 61?28'W, 900 m, 15 Dec. 1978, Julian A. Steyermark et al. 117615 (holotype, VEN; isotype, NY or 15 m, ramulis juvenilibus superne sparsim puberulis pilis laxis 0.1 mm praeditis; petiolis 3-15 mm longis glabris vel sparse puberulis; foliorum la- minis oe apice acutis vel obtuse acutis basi cuneatim angustatis 4—9.5 cm longis 2.5-4 cm latis ubique ga bris, nervis lateralibus utroque latere 6-7 eben venulis terti la we ` oo ferentibus; pedunculo ten m lato apice haud dilatato; E sub floribus lanceo- latis 0.8-1.2 mm longis extus parce puberulo-ciliatis; perianthio mascula infundibuliformi 3-3.5 mm longo superne 1-1.6 mm lato basi 0.3 mm lato extus inferne pilis laxis paucis praeditis atque loborum ee papillatis aliter glabris; staminibus 6 inclusis Tree 15 m, the young branches near tip sparse- ly laxly pubescent, elsewhere glabrous. Petioles 3-15 mm long, glabrous or with a few sparse, lax hairs; leaf blades opaque, drying dull brown, ob- anceolate, acute or abruptly obtusely acute at apex, cuneately narrowed at the equal to slightly asymmetric base, 4—9.5 cm long, 2.5-4 cm wide, glabrous both sides; lateral nerves 6-7 each side, m hina the margin, the ter- uous. Staminate inflo- rescence subhemisphe cm wide, with 4—6 umbellately disposed primary axes 5-17 mm long, glabrous or sparsely laxly pubescent; secondary axes 2-5, filiform, umbel- lately disposed, 3-5 mm long, glabrous or laxly sparsely pubescent, these branched into pedicel- late axes 0.5-2 mm long, each bearing 1-3 flow- ers subtended by lanceolate, acute bracts 0.8-1.2 mm long, sparsely ciliate-puberulent. Peduncle slender, 2—3.8 cm long, 0.8-1 mm wide, gla- 634 brous, not manifestly enlarged at its junction, 1.5 mm wide where united with the lowest primary axes of the inflorescence. Staminate perianth in- fundibuliform, 3-3.5 mm long, 1-1.6 mm wide at summit, 0.3 mm wide at base, glabrous except for a few sparse hairs near base and papillate- margined lobes. Stamens 6, included; anthers broadly rhomboid, 0.6 x 0.3 mm; filaments 1.5- 2 mm long. Paratype. VENEZUELA. BOLIVAR: Río Aponguao 2, 151-152 km S of El _ sA et al. 10524 (NY, VEN); Uei-tepui, between Luepa and Cerro Venamo, 1,100-1,300 m, EDEN 318 (NY. VEN). Neea tepuiensis Steyermark, sp. nov. TYPE: Ven- ezuela. Bolivar: Chimantá Massif, Torono tepui, summit, south-facing forested slopes above valley of South Cano, 1,955-2, m, 23 Feb. 1955, Julian A. Steyermark & John Wurdack 1104 (holotype, VEN; isotypes, F, MO, NY). rbor 4-5 m praeter gemmas terminales ferrugineas omnino glaberrima; petiolis 2-10 mm longis; foliorum ovatis vel obovato-ellipticis vel foliis parvis interdum suborbicularibus apice anguste obtusis vel rotundatis basi praeter folia parva rotundata vulgo subacutis vel anguste acutis (2.7-4.3)7-9.5 cm longis (1.5-3)3.5—5.5 cm latis, nervis lateralibus principalibus utroque latere 5-6; inflorescentia foeminea 1 cm alta s cm iura im- longis s 2-3 mm sae gerentibus cula 2 mm iris 0.8 mm la pice haud manifeste dilatato; anthocarpio anguste el- idee oblongo 9 mm longo 4 mm lato glabro. Tree 4-5 m tall, glabrous throughout, only the terminal bud glandular-ferruginous pubescent. Leaves opposite or subopposite; petiole 2-10 mm long; leaf blades shining above, elliptic-ovate to obovate-elliptic, or some of the smaller leaves suborbicular, obtuse to rounded at apex, mainly subacute to acutely narrowed at base, or some of the smaller leaves rounded at base, the small leaves 2.7-4.3 cm long and 1.5-4.5 cm wide, but leaves mostly 7-9.5 cm long and 3.5-5.5 cm wide, finely impressed-nerved above, finely and slight- ly elevated-nerved below, secondary nerves 5-6 each side, faintly anastomosing 5-13 mm from margin, ascending to an angle of 15—20°; tertiary venation slightly evident and reticulate beneath. Pistillate inflorescence 1 cm high, 1.7 cm wide (immature?) with 4 primary axes 6-7 mm long, .6 mm wide, each branched into 4 smaller sec- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 ondary axes 2-3 mm long ending in 3 axes bear- ing the flowers. Flowers sessile. Pistillate peri- anth 6 mm long, 2 mm wide. Staminate perianth tubular, glabrous except for the papillate-puber- ulent npes at the summit, 2 mm long, 0.8 mm wide. B th lanceolate, sub- acute, 0.8-1 mm long, | glandular-pubescent. In- fructescence terminal, bearing 3-4 main axes 7- 11 mm long and 1 mm wide, these suns into 4 secondary, subumbellate axes 4-7 m long, these eventually terminating in 4 idle or slightly branched pedicels 1.5-2 mm long bear- ing the fruits. Anthocarp narrowly ellipsoid-ob- long, 9 mm long, 4 mm wide, glabrous, slightly striate. Paratype. VENEZUELA. BOLIVAR: Chimanta Massif, Agparaman tepui, forested middle slopes near Río Ti- rica, 1,365 m, 5 Mar. 1955, Steyermark & Wurdack 1255 (F, NY, VEN). This species attains the highest altitude on the sandstone tabular mountains of the Venezuelan Guayana, where it grows at an altitude of 1,365- 2,090 m. It is characterized by the glabrity of all parts and is distinguished from N. robusta n. erm. byi its smaller flowers and by the obsolete venation of the tertiary veinlets. From N. subglabrata Steyerm. it differs in leaf shape, greater diameter of the peduncle, usually subobtuse to rounded leaf apices, lustrous upper surface of leaves, and subelevated, subreticulate tertiary venation of the lower leaf surface. LITERATURE CITED BURGER, W. 1983 . Nyctaginaceae. Jn Flora Costa- ricensis. Fieldiana, Bot. N.S. 13: 190-191, 196 193- HEIMERL, A. 1896. Additamenta ad cognitionem flo- e Indiae occidentalis. III. Bot. Jahrb. Syst. 21: 615-636. 897. Beitráge zur Systematik der Nyctagi- naceen. Jahresber. Staats-Oberreal. Wien XXIII. —39. 1914. a In R. Pilger a Pl antae Uleanae novae vel minus cognitae. No- tizbl. Kónigl. Bot. "Ga rt. Berin 6: 126- 1932. Nyctaginaceae. Jn N. Y. Sandwith (ed- itor), Contr. to the Flora of Tropical Am. XII. Kew Bull. 1932: 220-221. Huser, H. 1909. Materiaes para a Flora Amazonica. VII. Plantae Duckeanae austro-guyanenses. Bol. Belém Mus. Paran. Emilio Goeldi 5: 347-351. LITTLE, E. L. 1968. Transfers to Guapira from Tor- rubia (Nyctaginaceae). Phytologia 17: 367-368. LUNDELL, C. L. 19 tudies of tropical American plants—V. Wrightia 4: 72-84. SCHMIDT, J. A. Nyctagineae. /n Martius, Flora Brasiliensis 14(2): 351-369. STEYERMARK 1987] — VENEZUELAN GUAYANA FLORA~III 635 STANDLEY, P. C. 1931. western South America. Fie Bot. Ser. 11: 73-90. The Nyctaginaceae of North- ld Mus. Nat. Hist BRUNELLIACEAE BRUNELLIA Brunellia nebli is St k & Cuatrecasas, nov. TYPE: Venezuela-Bexzil border. za de La Neblina, en bosque alto, 1,500 m, 29 Apr. 1964, J. Ewel 209 (holotype, NY). Arbor, kes sad a imparipinnatis 6-jugatis; fo- liolorum laminis ha ud discoloribus chartaceis ellipti- nervis ipe eas 16-17 utroque latere distantibus 8- ; foliolis inferioribus 1-2 m lobato 3.5-4.2 triangulari-lanceolatis acutis 1.5-1.8 mm longis 1-1.2 mm latis; folliculis subrotundatis 3 mm longis 2.5 mm latis dense hirtellis basi pilis hispidis munitis; enl nibus obconico-subglobosis subcompressis 3 mm 1 is 2.5 mm latis. Tree with opposite imparipinnate leaves. Leaf rachis minutely puberulent. Leaflets in 6 pairs, not discolored, chartaceous, elliptic-oblong, acute at apex, asymmetrically acute to subobtuse at base, 15-18.5 cm long, 4.5-6 cm wide, crenate- serrate, the upper leaflets sessile, the lower ones 1-2 mm, shortly petiolate; lower surface gla- brous, except minutely puberulent with short spreading hairs on midrib, lateral nerves, and tertiary veinlets; upper surface glabrous. Infruc- tescence densely tomentellose, the branches 1 mm diam. Calyx 4—5-lobed, 3.5-4.2 mm in fruit, densely hirtellous, the lobes triangular-lanceo- late, acute, 1.5-1.8 mm long, 1-1.2 mm wide. Follicles subrotund, 3 mm long, 2.5 mm wide, densely hirtellous with additional hispid hairs at ase. Seeds shining, obconic-subglobose, nar- rowed toward one end, 3 mm long, 2.5 mm wide. This taxon differs from Brunellia comocladi- folia Humb. & Bonpl. and especially the subsp. ptariana (Steyerm.) Cuatrec. in the smaller fol- licles, shorter calyx with slightly narrower calyx lobes, larger seeds, and the glabrous lower leaf surfaces, except for the minutely tomentellose nerves. From B. hygrothermica Cuatrec. it may be distinguished by the concolorous leaves with glabrous lower leaf surface, more numerous pairs of lateral nerves, more depressed and less con- spicuous serrations, and shorter calyx. From B. gentryi Cuatrec. it is differentiated by the slightly larger calyx diameter, larger calyx lobes, longer and numerous hispid hairs of the follicles, and concolorous leaves not silvery green beneath. SAPINDACEAE MATAYBA Matayba ptariana Steyermark subsp. guaiqui- nimae Steyermark, subsp. nov. TYPE: Ven- ezuela. Bolívar: Departamento Heres, Me- seta de Guaiquinima, sector NE de la meseta, cerca de la cumbre, 5?58'N, 63?29'W, 1,400 m, 27 Mar. 1985, Otto Huber 10388 (ho- lotype, MO; isotypes, NY, VEN) A subsp. ptariana foliolis minoribus apice rotun- datis, venulis tertiariis haud elevatis, calycis lobis sub- iai predic dense — pubescentibus, petalo- mis interioribus longitudinem petali nnii en recedit. Shrub, 2-3 m tall. Leaves bifoliate, glabrous, the leaflets oblong, rounded at the apex, subob- tuse or subacute at the base, 2-4 cm long, 1-2.2 cm wide. Inflorescence axillary, 4.5 cm long, the flowering part 2.5 cm long, 6-7 mm wide, sub- racemose, 23-flowered. Peduncle 2 cm long, 1 mm wide, solitary, minutely appressed-pubes- cent; bracts ovate-oblong, subacute, 0.7-1 mm long, 0.5 mm wide. Pedicels 2 mm long, densely pubescent with ascending, appressed, pale hairs. Calyx lobes 5, broadly suborbicular-ovate rounded to obtuse, appressed ne eee withbut. 1x 1 mm, sericeous within. Petals pilose-ciliate, subalbiciine broadly rounded above, gee de id short-unguiculate at base, 1.3 mm long, broad; 2 inner segments oblong, rounded, b mm long, 0.8 mm wide, densely pilose both sides. Stamens 8, exserted; anthers suborbicular, 0.8 mm long; filaments 1.7-1.8 mm long, pilose be- low, glabrous above the middle; disk cupulate, glabrous, 0.5 mm high. At first, this collection appeared to be distinct from Matayba ptariana. However, a second col- lection (Huber 10339), assigned to subsp. pta- riana, in fruit from a lower altitude at 480 m (“Rio Trueno, 35 km west of caserio of Chig- uao”), although showing the rounded leaf apices together with inconspicuous impressed veinlets on the lower surface of the new subspecies, has much larger leaflets and longer petioles similar 636 to typical M. ptariana subsp. ptariana. Until ad- ditional material is forthcoming, it seems best to retain a subspecific rank for the Huber collection of the higher altitude. BOMBACACEAE NOTES ON CATOSTEMMA AND SCLERONEMA (BOMBACACEAE) In the course of preparation of the genus Ca- tostemma for the Flora of the Venezuelan Gua- yana, it was found that all the Venezuelan ma- terial had been identified either as C. commune Sandw. or had remained unidentified. An ex- amination of available material in the herbaria of F, K, MO, NY, US, and VEN indicated that more than one species of Catostemma was rep- resented in Venezuela and revealed further the presence of the genus Scleronema. The present study includes an attempt to delimit the genus Catostemma in Venezuela, together with a de- scription of a new species in Scleronema. Publications by Sandwith (1931, 1948), Ducke (1937), and Paula (1969) have greatly enhanced our knowledge of both genera with the result that the characters differentiating Catostemma from Scleronema can now be more clearly defined, involving chiefly stamen differences. In Sclero- nema the filaments are dilated and thickened apically with the small, strongly sessile anther appressed across the summit of the filament, whereas in Catostemma the uniformly slender filaments bear a larger incumbent anther folded inward over the summit. Additional differences ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 are the longer and more numerous filaments and longer style branches in Catostemma, also from available fruiting material, the elongated, ellip- soid, oblong, or obovoid, tardily dehiscent fruit of Catostemma as opposed to the globose or ee ds indehiscent fruit of Scleronema. Sandwith (1931) was able to differentiate two species of Catostemma from Guyana, C. com- mune Sandw. and C. fragrans Benth., on the ba- sis of vegetative characters found in the seedling leaves and in the venation, especially ofthe lower leaf surface. In the present study vegetative dif- ferences have been noted and employed to dis- tinguish several taxa. Unfortunately, leaves of seedling plants are still unknown from most of the taxa recognized. In addition to the type of tertiary venation of the lower leaf surface, useful characters have been found in the glabrity or stellate tomentum of the buds and petioles. Moreover, the absence or presence of stellate to- mentum and its distribution on the style and style branches, and the presence of bracteoles and their position on the pedicels serve to delimit various taxa within the complex. Much of the material in herbaria is sterile or lacks flowers, and this has made delimitation of the taxa dif- cult. The author wishes to acknowledge to the cu- rators of F, K, MO, NY, US, and VEN his great appreciation for the loan of herbarium material. Based on the material examined, a description is provided for the taxa of Catostemma, together with a citation of specimens, followed by a key to the species. CATOSTEMMA KEY TO THE SPECIES OF CATOSTEMMA la. Adult leaves compound lb. Adult leaves simple C. digitatum 2 2a. Petioles and young stems hirsutulous with spreading to ascending stiffish hairs 0.2-0.4 mm long 2b. ape and young stems glabrous or with minute or closely appressed tomentum ioles and New stems stellate- tomentose C. hirsutulum 3 4 a. Petioles tion E di wer ‘leaf surface irregular or scarcely et Pe leaf surface glabrous; ejon nt from pedicels; en nerves of leaf blades 5-6 each side; style and style branches C. clarkii ox ste late pubesce . Petioles and y venatio a oflo ower leaf su + c rfacem oung stems n nope with larger, less crowded tomentum; tertiary r less regular, subparallel and subhorizontal; lower leaf surface minutely stellate- SU to glabrous; bracteoles Arai on pedicels; lateral af blades nerves of le 8-12 each side; style glabrous except at bas . C. fragrans 5 3b. dg pa young stems glabro Sa. veins of lower leaf ipd see in more or less subparallel lines obliquely connecting C. m "a dM nerves arahuacense 1987] STEYERMARK— VENEZUELAN GUAYANA FLORA —III 637 5b. Tertiary m Ws lower leaf surface irregularly reticulate and anastomosing, not regularly obliquely par 6a. Style stellate- n throughout; di branches stellate-pubescent; us scales wit ssed, stellate pubes pubistylum . Style Mes ee at ol or lowest ^; stylar | or the innermost ones pilos 7a. Bracteoles absent fent een calyx lobes densely strigose within; innermost bud scales pilose apically ebracteolatum 7b. Bracteoles present, either on the pedicels or immediately subtending the base of the calyx; calyx lobes glabrous within or partly strigose near apex; bud scales glabrous 8 8a. itl mediately subtending the calyx or on the uppermost 1- sel: leaves without revolute ier kane 9 the scales more or less con- tiguous or overlapping with numerous, relatively elongated rays; leaf base mainly acute to cuneate, more rarely obtuse „u... C. commune 9b. Pedicels with moderately stellate ironies | the scales minute, scat- dd. and more separated with fewer and shorter rays; leaf 2 mainly unded, or sometimes subcordate sclerophyllum Í Ma at least some of them, toward or below the middle 2 T pedicel; leaves usually with revolute margins 10a. Pedicels 4-10 cm long; principal lateral nerves of the leaves 8-12 each ies leaf blades up to 30 cm long and 11.5 cm wide; petals 17-23 m long; tree 10-45 m tall C. altsonii 10b. Pedicels 2.5-3 cm long; principal lateral nerves of the pis "uad each side; leaf blades up to 15 cm long and 4.5 cm wide; petals 15 mm long; shrub 2-4 m tall a sancarlosiana 1 lat hiv o n = oo c Catostemma altsonii Sandw., Kew Bull. 1928: 66. 1928. TYPE: Guyana. Macreba Falls, Kurupung River, Sept. 1925, Altson 391 (holotype, K). Figure 6A, a. Tree, 10-45 m tall, the branchlets glabrous; bud scales glabrous. Seedling leaves 3-foliate. ture leaves with petioles 0.8—6 cm rounded and mucronate at apex, narrowed to a cuneate or obtuse base, 8—30 cm long, 3-11.5 cm wide, glabrous, the margins revolute; primary lateral nerves 8-12 each side, impressed above, slightly elevated below, at irregular angles and not uniformly ascending, branching or anasto- imo ass reaching the margins, 7-25 mm apart; tertiary venation of the lower leaf surface So reticulate, usually prominulous, usu- ally impressed above on sterile branchlets. Ped- icels in anthesis 4—10 cm long, densely stellate- 3 e. acute, situated alter- nately and mend boxer below and above the middle. Calyx densely stellate-tomentose with- out, 13-15 mm long, the tube 3-5 mm long, the lobes ovate-oblong, obtuse, 10 mm long, gla- brous within. Petals obovate-oblong, rounded, 17-23 mm long, 7-8 mm wide. Stamens ca. 35- 40; filaments 10-14 mm long; anthers 1-1.3 mm long. Style 10 mm long, glabrous except in low- ermost 4-6 mm, there stellulate; style branches 2-3.5 mm long, glabrous. Mature fruit ellipsoid, 15-20 cm long, 3-5 cm diam.; immature fruit obovoid-oblong, ca. 5 cm long, 2 cm diam., mi- nutely ferruginous furfuraceous, fruiting pedicsl l Distribution. Wallaba forest at 50-125 m el- evation; known only from Guyana Additional specimens examined. GUYANA: Macre- ba Falls, Kurupung River, Mazaruni tributary, A/tson 391 (K, ie Essequibo-Mazaruni Divide, about 25 mi. south of Bartica, Forest Dept. Record No. 2316 (Field ios D323) (K); Membaru creek, u uppe er Mazaruni Bartica-Potaro road, Forest Dept. Record No. F1520 DM 4256) (K); 1'2 mi. Bartica-Potaro road, Forest D t. Rec ord No. 6906( Field No. CAP149) Forest Dept. Record No. 2315 (Field No. D Potaro River below Tukeit, Maguire & Fanshawe 23486 (NY); Bartica-Potaro road, 24 mi. south of Bartica at Forest Service's “24 mile Camp," 50-125 m, Mori et al. 8902 (NY); Membaru Creek, Upper Mazaruni Riv- er, Pinkus 238 (F, NY, US); Bartica-Potaro road, For- est Dept. No. 107 (Field No. F 1520, Record No. 4256) (NY). 638 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FiGURE 6.—A. Catostemma altsonii, leaf. a. Detail of lower leaf surface n irregularly reticulate tertiary paqa —B. i e es fragrans, leaf. b. Deta rtiary venatio siut iun lattiat ascent Based on holoty Two other species of Catostemma are known from Guyana. The present taxon may be distin- guished from C. fragrans Benth. by the glabrous uds, petioles, and young stems, by the irregu- larly reticulate tertiary venation on the lower leaf surface, and by the irregular branching of the main lateral nerves. From C une it can be e es longer pedicels, longer petals and calyx lobes, and proportionately longer leaf blades Catostemma clarkii Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: il of lower tostemma haha isis leaf. c leaf surface showing subparallel, subhorizontal gael of eae leaf surface showing subparallel, Mari’s bana (low Amazon caatinga), 10.8 km NE of San Carlos on road to Solano, 1°56’N, 67°03'W, 119 m, 19 Aug. 1981, Howard L. Clark 8126 (holotype, MO). Arbuscula vel € gemmis ramulis juvenili- busque minute confertimque cano-stellato-tomentel- lis; petiolis minute cano- alate foliorum laminis sub- di inute stellat laminarum tus praete juvenilium glabris anguste obovatis, nervis lateralibus stellatis bracteolis desunt; calyce extus dense stellato- omentello intus lobis glabris; petalis 20-22 mm longis; ae inm stellato-tomentello ramis jure mm longis leviter stellatis STEYERMARK 1987] — VENEZUELAN GUAYANA FLORA —III 639 Shrub or small tree to 12 m; young stems densely and minutely gray stellate-tomentellose; older branches glabrous. Petioles 4-25 mm long, minutely and densely gray stellate-tomentellose on young leaves, minutely gray stellate to gla- brescent on mature leaves; leaf blades narrowly obovate, rounded or retuse at apex, mucronulate in the sinus, narrowed to a rounded or subcor- date base; mature leaf blades rigid-coriaceous, 4.5-10.5 cm long, 2-4.5 cm wide, glabrous throughout except for the Ciel stellate basal portion of the lower midrib; young leaves mi- nutely gray stellate- utei tm on the lower midrib; main lateral nerves 5-6 each side, scarce- ly manifest, lightly impressed on the upper side, branching t g the margins, 8-15 mm distant; tertiary venation beneath inconspic- uously anastomosing. Pedicels ebracteolate, ca. 3.2 cm long, densely and minutely gray stellate- tomentellose. Petals ligulate-oblong to subspat- ulate, 20-22 mm long, 6-9 mm wide. Calyx densely stellate without, the tube campanulate, m long, the lobes ovate, obtuse, 11 mm long, i ae ar Stamens numerous, about 40; filaments 12—13 mm long; anthers 0.8-1 mm long. Ovary conic, 7 mm long, 3 mm diam., tapering to an elongate beak 2.7 mm long. Style 9 mm long, sparsely stellate-pubescent from base to apex, the branches 4-5 mm long, lightly stellate- pubescent. Fruit ellipsoid, 6-6.5 cm long, 4.5 cm diam., fruiting pedicel 4.5 cm long. r 1 VENEZUELA atype. lity as type, | Jan. Ti Clark 6919 (MO). This species is characterized by the minute, pale gray, closely stellate tomentum of the veg- etative buds, young petioles, and lower midrib of the early leaves, by the sparsely stellate pu- bescence on style and branches, and by the scarcely manifest tertiary venation of the lower leaf surface. The stellate tomentum of this species is the most minute of the genus. Catostemma commune Sandw., Kew Bull. 1931: 1. 1931; Oliver in Hook. Icon. Pl. 1986. 1891, as to flowers and leaf, but not to fruit. TYPE: Guyana. Moraballi Creek, Essequibo River, Aug. 1929, Sandwith 435 (holotype, K) Tree to 45 m. Young branches and stems and bud scales glabrous. Seedling leaves usually 3-foliate (rarely 2- or 4—5-foliate), the leaflets ob- lanceolate, obliquely acuminate at the apex, the acumen triangular-lanceolate, 1-1.8 cm long, 3- 6 mm wide above the middle, obtuse, but the midrib excurrent, cuneately attenuate at base ex- cept for the rounded lower side of the outer leaf- lets, 9-32 cm long, 3-8 cm wide, glabrous both sides. Mature leaves with petioles 1—9 cm long, glabrous. Mature leaf blades stiffly chartaceous or subcoriaceous, elliptic o or obovate, pounce’ or retuse at apex, rrent costa, cuneate to rarely obtuse at base, 4-19 cm long, 2-9 cm wide, glabrous on both sides; main lateral nerves 8-12 each side, impressed above, slightly elevated below, ascending at an angle of 40-50°, branching and anastomosing just before reaching the margins, 8-15 mm distant; tertiary venation faintly impressed-reticulate above, irregularly re- ticulate with slightly elevated veins below. Ped- icels 0.5-2 or 3 cm long, densely stellate-tomen- tose, bracteolate. Bracteoles paired, approximate, overlapping, alternate, squamiform or rarely conspicuous, situated in the upper part of the pedicel or immediately subtending the calyx, ovate or suborbicular, obtuse to subacute, 1.5- 8(-6) mm long, 1-1.5(-4.5) mm wide. Calyx densely stellate-tomentose without, the tube campanulate, 4 mm long, 3-4 mm wide; lobes ovate, 7 mm long, 6 mm wide. Petals d pad spatulate, obtuse, 12 mm long, 4-5 mm w Stamens numerous, 40-50; filaments 8-10 mm long; anthers 0.8-0.9 mm long. Style 13 mm long, stellate-tomentose in lower !^5, elsewhere gla- brous; style branches 0.75-1.5 mm long. Fruit oblong-ellipsoid, 7-10 cm long, 3-4 cm diam., densely ferruginous tomentellous. Common names. Baromalli, common ba- romalli (Guyana), baramán (Venezuela). Distribution. In mixed tall forest of green- heart, mora, and morakubea of Guyana, prin- cipally in the Essequibo and Cuyuni river basins, and in evergreen mixed forests, often with Eper- ua, in Estado Bolívar and Territorio Federal Del- ta Amacuro of eastern Venezuelan Guayana, at altitudes of 80-850 m Additional erue examined. WEST INDIES: cul- tivated in Old Botanic Gardens, St. Vincent, April 1891, H. Powell (K). GUYANA. Moraballi Creek, Essequibo No. D402) (K); Upper Mazaruni Kit Leng 416 (seed- ling leaves, NY); Matt Ridge, ma River, Northwest Territory, 80 m, Cowan 39361 (F, NY), 39361-A (fruit, NY); P Forest Dept. Record No. 7712 (K Tutin 338 (K, US). VENEZUELA: TER d Cuyuni River, ORIO RAL DELTA AMACURO: este de Rio Grande E on de 640 El Palmar cerca de xp "hen del Estado Eae Nov. 1965, Carlos Blanco 495 (MO, US, VEN); La Paloma, Río Cuyubini, T 100-200 m, pppn rk 87583 (F, MO, NY, US, VEN); El Morro, Atabuina, Caño Arature, base of Sierra Imataca, 750 m, Bernardi 7529 (K, NY, VEN); same locality and date, Buza 329 (K, NY, VEN); near Río Grande, east of Upata, border between Estado Bolívar and Territorio Federal Delta Amacuro, 8?14'N, 61°4'W, 300 m, de Bruijn 1631 (F, € I US, VEN), 1677 (US, VEN), 1623 (F, M ; same locality, Breteler 3841 Dee tie leaves, NY, US), 4958 (NY, US), 4972 (NY); ENE de El Palmar, cerca de los limites del Estado Bolívar, Zabala 96 (VEN), 163 (VEN). BOLÍVAR: base of Cerro Pauji, Quebrada 94, km 94 south of El Do- rado, 250 m, Steyermark 867 11 (NY, US, VEN), 86712 (VEN), 86715 (F, US, VEN); La Isabel a Rio Grande, El Palmar, Conejos 36 (MO, VEN); Cerro La Reforma, above junction of Río Reforma with Río Toro (Río Grande), Sierra Imataca, 200-250 m, Steyermark 88113 La Lira trail, km 27 S of El Dorado, o, ESE of Villa Lola, Altiplanicie de dr. 315m, Foi rk 86049 (seedling leaves, y Río Chirca, Bernardi 898 (3-foliolate leaves, Se east of El Palmar, Marcano-Berti 319 (VEN); Río ree between mouth of Rio Aparurén and Uriman, 40 Steyermark 76077 (F, VEN); región de las eens del Rio Hacha, 450-850 m, Bernardi 2897 (NY); Rio Caura, arriba del Salto Para, 2-3 km arriba del cam- pamento “Las Pavas,” 250—300 m, J. Steyermark, G. . K. & E. Dunsterville 112934 (MO, VEN); same lo- cality, Morillo & Liesner 8944 (MO, VEN); 30 km S of El Dorado, 140 m, Bernardi 2097 (NY, VEN); Río Venamo, afluente del Ikabaru, Cardona 1706 (VEN); Río Curutü, upper Paragua, 550 m, Cardona 2484 (VEN); Salto Ichun, Río Ichun, tributary of Rio Par- agua, 4?46'N, 63?*18'W, 500 m, Steyermark 90371 (VEN); El Abismo, Río Samay, 4?27'N, 61?34'W, 550- 600 m, Holst & Liesner 2422 (MO, VEN). In Sandwith's excellent discussion of the ba- romallis (Catostemma) of Guyana (1931), a number of characters were noted by which C. commune Sandw., or common baromalli, could be distinguished from C. fragrans Benth., or sand baromalli, the latter being the type species of the genus based on a collection by Robert Schom- burgk 280 in 1837 from Berbice, Guyana. One of the differentiating | characters mentioned by C. commune was the " bracteoles. Sandwith grans (plate 1986 of Hook. Icon. Pl., 1891). Bracts conforming to these dimensions are evidenced by a specimen at K which had been sent from the Old Botanic Garden of St. Vincent, collected by H. Powell in April 1891, where it had been cultivated for many years. This specimen was cited by Sandwith as a paratype. However, the ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 actual type specimen at K (Sandwith 116) does not have either large or conspicuous bracts. Thus, the only flowering material available to Sandwith was his own collection (the type) and that of Powell from a cultivated specimen Three subsequently collected specimens from the wild state show small squamiform bracts only, which do not exceed 2.8 mm in length and 1.5 mm in width. These bracts or bracteoles subtend the very base of the bud, as in Tutin 338 and Forest Dept. Record No. 7712 (both at K), or occur on the uppermost 1-6 mm of the pedicel below the base of the calyx, as in Forest Dept. Record No. 2398, Davis 402 (K). All these spec- imens were annotated by Sandwith as C. com- mune Sandw. Similarly, flowering collections of this species from adjacent eastern Venezuela show only squamiform bracts va on the upper- most 1-6 mm of the pe Thus, the description s de bracts in C. com- mune should be emended as follows: “bracteis squamiformibus late ovatis vel suborbicularibus obtusis vel subacutis 1.5-2.8(5—6) mm longis 1— 1.5(-4.5) mm latis binis vel arcte alternantibus." Catostemma digitatum Shepherd & Alverson, Brittonia 33: 587, fig. 1. 1981. TYPE: Colom- bia. Antioquia: confluence of Quebrada La Tirana and Río Anori, 3 km upriver (SW) from Planta Waaa ca. 28 km SW of Zaragota, 7°13'N, 75°3'W, 500 m, 9 Apr. 1977, W. S. Piu S. White & E. J. D. Shepherd 397 (holotype, COL). Tree to 30 m; young branches or stems gla- brous. Buds sericeous-tomentose. Mature leaves 3-5-palmately compound. Petioles 6-21 cm long, glabrous. Leaf blades chartaceous, lanceolate, long-caudate in seedling leaves, subacute or ob- tuse and mucronulate in mature leaflets, acute at base, mainly glabrous; principal lateral nerves 10-16 each side, arcuately ascending near the margins; tertiary venation below irregularly laxly reticulate, scarcely prominulous. Flowers un- known. Fruit obovoid, 9-12 cm long, 5.5-6 cm diam.; seed 5-7 cm long, 2-4 cm wide. Distribution. Known only from the depart- ments of Antioquia and Santander, Colombia. Additional specimens examined. COLOMBIA. ANTIOQUIA: confluence of Quebrada La Tirana and Río Anori, 28 km SW of Zaragota, 7?13'N, 75°3'W, 500 m, Alverson, White & Shepherd 397 (holotype, COL; isotypes, HUA, MO, NY, WIS); slope facing Río An- ori, near Planta Providencia, Denslow 1404 (seedling, COL, HUA, MO, NY, WIS); confluence of Quebrada 1987] La Tirana and Rio Anori, 3 km upriver (SW) from Planta Providencia, ca. 28 km SW of Zaragota, 7?13'N, 75°3'W, 500 m, 9 Apr. 1977, W. S. Alverson, S. White, & E. J. D. Shepherd 397 DE COL). SANTANDER: Magdalena Valley, Cam E of Carare, 300 m, Gentry & Renteria 20065 er MO, UIS), 20106 (COL, MO, UIS). This is the only species thus far known in the genus with the mature leaves compound. Catostemma ebracteolatum oe sp. nov. TYPE: Venezuela. Territorio Federal Ama- zonas: Cerro Sipapo aa ga water course above Intermediate Camp, 2 Feb. 1949, Bassett Maguire & Louis Politi 28741 (ho- lotype, MO; isotype, NY) E on 15-30 m, ramulis juvenilibus glabris; gem- e squamis intimis acuminatis pilosis extimis cau- datis petiolis 3- -35 mm longis glabris; foliorum laminis asi acutis vel cuneatis majoribus 6-10 cm longis 2.5-6.5 cm latis ubiq ue glabris, nervis lateralibus utroque latere 7-9 supra impressis subtus leviter elevatis, venatione tertiaria subtus leviter irregulariterque reticulatis; ped- icellis 1.5-4 cm longis pilis dimorphis munitis, pilis brunneis minoribus t pallidis lon- gioribus patulis plus minusve seriali bus alternantibus so, tubo 4 ete] lato-to longis tabes dense strigosis; petalis ligulato- oblongis 15 mm longis; filamentis ad 9 mm longis; stylo parte in- feriore » ce 2 stellato-tomentoso ceterum glabro, ramis ad 4 mm longis glabris. Tree 15-30 m tall, the young stems glabrous; innermost bud scales acuminate, pilosulous, the others glabrous, the outermost caudate. Seedling leaves trifoliolate. Petioles 3-35 mm long, gla- brous; leaf blades coriaceous, rich green above, paler below, obovate, rounded and retuse at apex, mucronulate, narrowed to an acute or cuneate base, 6-10 cm long, 2.5-6.5 cm wide, glabrous; lateral nerves 7-9 each side, spreading at first from an angle of 10—20°, then ascending, branch- ing before reaching the margins, impressed above, lightly and narrowly elevated on the lower side; tertiary venation irregularly and inconspicuously reticulate below. Pedicels costate, especially to- ward apex, 1.5-4 cm long, ebracteolate, with 2-tiered pubescence, the shorter brownish, ap- pressed, stellate-tomentose hairs between paler longer hairs + prominently spreading from pseu- dovertical rows. Calyx in pre-anthesis 12.5 mm long, split above the side from one portion, densely stellate without, 4 mm long, 3-5 mm wide above; lobes longer than the tube, 7-8 mm long, suborbicular-ovate, obtuse, densely stri- gose within. Petals ligulate-oblong, rounded, 15 STEYERMARK — VENEZUELAN GUAYANA FLORA—III 641 mm long, 5-6 mm wide. Stamens 25-30; fila- ments up to 9 mm long; anthers 0.8-0.9 mm long. Style pubescent for 2.5-4.5 mm from the base up to !^-'^ of the length, glabrous above; style branches up to 4 mm long, glabrous. Fruit orange, rounded at apex, manifestly narrowed to the base, obtusely 3-costate, 7 cm long, 4 cm iam., minutely SONOMA fruiting pedicel 4 cm long, 5 cm dia Paratypes. "VENEZUELA. BOLÍVAR: Sierra Ichün, cer- canías del Salto María Espuma (Salto Ichün), base of la Sierra Ichün, tributary of Río Paragua, 4?46'N, 68?18'W, 500 m, = a 1961, Steyermark 90371 (F, NY, US, VEN). T ORIO FEDERAL AMAZONAS: 4.3 km NNE of San nue on Solano road, 1?56'N, 67°3' W, 100-300 m, Steyermark, Davidse & Guanchez 122136 (MO, VEN), 172386 (MO, VEN), 122402 (MO, VEN); Sierra Parima, headwaters of Rio Siapa and Rio Pa- dauiri, 1°22’N, 64°38’W, 1,260 m, Cardona 1507 (US, VEN); Yavita, Lizot 1972-4 (US, VEN). This taxon differs from other members of the genus in its two-tiered pub on the pedicel, with longer, pale, spreading hairs in more or less separate vertical rows or lines rather prominently differentiated from the more abundant, smaller, brownish, more appressed stellate tomentum in between. In the other species of the genus the stellate tomentum is more uniformly appressed, pale to dark brown, and uninterrupted by longer hairs. Catostemma fragrans Benth. in Hook. London Bot. 2: 365. 1843; Baker in Hook. Icon. Pl. 1793. 1888; Oliver in Hook. Icon. Pl. 1986. 1891, as to fruit. TYPE: Guyana: Ber- bice, 1837, Rob. Schomburgk 280 (holotype, K). Figure 6B, b. um € Sagot ex vires Bull. Mus. Hist. Nat. (Paris) 25: 387. 19. TYPE: French nndis Acarouany, Sagot s.n. (holotype, P). Tree (4-)1 5-30 m tall. Young branches or stem stellate-tomentose. Buds stellate-tomentose. Seedling leaves simple, 12-38 cm long, 3-7 cm wide, obovate-oblong or obovate-elliptic, nar- rowly elongated, cuspidate at apex, the cuspidate portion linear-subulate, this 1-5 cm long, 0.7- 2.5 mm wide, narrowed to a subobtuse to sub- rotundate base. Mature leaves with petioles 0.5— 5 cm long, stellate-tomentose; mature leaf blades coriaceous or subcoriaceous, obovate-oblong, obovate-elliptic, or oblong-elliptic, rounded or retuse, sometimes cuspidate at apex, rounded at base, the larger ones 17-22 cm long, 3-8 cm 642 wide, glabrous above, glabrous to sparsely stel- late-puberulent below, sparsely to moderately stellate-pubescent on the lower midrib and main lateral nerves; lateral nerves commonly 8-12 each side, impressed above, elevated below, + regu- larly ascending at an angle of 45-60° and sub- parallel nearly to the margins, 5-20 mm distant; tertiary venation inconspicuous above, below conspicuous and elevated with the veins sub- parallel and subhorizontal. Pedicels 1.5-4 cm long, densely stellate-tomentose, bracteolate. Bracteoles 2 or 3, situated near the middle or in the lower or upper part of the pedicels, some- times only 2-3 mm below the base of calyx, paired or usually proximate alternately, ovate, obtuse, 1-2.5 mm long, 1 mm wide. Calyx densely stel- peti cna without, the tube 3-4 mm long, m wide, the lobes broadly ovate or ovate- pais obtuse, 9 mm lon 7 wide, gla- brous within. Petals spathulate- abus obtuse, 13-15 mm long, 4-7 mm wide. Stamens about 35-40; filaments 7-8 mm long; anthers 1-1.2 mm long. Style 11-15 mm long, glabrous except stellulate at the very base, the style branches 1.5- 2 mm long, glabrous. Fruit oblong-ellipsoid, 4— 8 cm long, 3-3.5 cm diam., densely and minutely ferruginous tomentellous; fruiting pedicel 2 cm long. Common names. Sand baromalli, baromalli or wallaba; baromalli (Guyana); kajoewaballi barmani (Suriname). Distribution. Sandy soils of wallaba forests at low elevations in Guyana, Suriname, and French Guiana. Gegen specimens examined. GUYANA: near Bartica, Moraballi Creek, Essequibo River, near sea ede Sandwith 485 (K, NY); Coverden, Persaud 134 (F, NY); basin of Essequibo River, near mouth of Ono- ro Creek, l cans , A. C. Smith 2739 (F, MO, NY, US); a Riv . No rthwest pops bso 59*50'W, ,NY,U ica—Potaro road, Creek, Rewa River, 35 mi. SSE of mouth, Forest Dept. Record No. 2087 (Field No. D96) (K); lower Demarara River, Demarara River, Hohenkerk 704-C (K). SURI- NAME: km 6, Wijneweg, “B.B.S. 174" (K, MO); Wa- yombo, Wood Herbarium 352 (K, NY); Maratakka River, Snake Creek, Maas 10799 (K, NY, U); Cor- antynes B.B.S. 901 (K); Tafelberg, Maguire Foengoe Island, Nat. Park Raleigh Falls, Roberts 14764 (NY); Christianburg, Demarara River, Anderson 271 (K); Wilhelmina Gebergte, 2 km below affluence of Oost River, 225 m, Maguire et al. 54098 (MO, NY). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FRENCH GUIANA: south of Crique Gregoire, Sinnamary River, Oldeman 1600 (NY); Sagot & Melinon (photo of Guenetia macrosperma, F, MO); Godebert, Wach- enheim s.n. (K) (as Guenetia macrosperma). This species is the most widely distributed of the species of Catostemma, being found 1n all the Guianas, where it is common, especially in the wallaba forests of Guyana. It differs from the common taller forest species C. commune Sandw. in it stellate-pubescent bud scales, petioles, and young stems; simple, long-cuspidate seedling leaves; completely different venation ofthe lower surface of both seedlings and mature leaves; style glabrous throughout, except at very base; and somewhat shorter, usually alternately Henn bracteoles on the pedicels. Catostemma hirsutulum Steyermark, sp. nov. TYPE: Venezuela. Bolívar: Chimantá Massif, rich rain forest on northwestern slopes of Abácapa-tepui, vicinity of camp | along Río Abácapa, 420 m, 30-31 Mar. 1953, Julian A. Steyermark 74781 (holotype, MO; iso- types, F, MO, VEN). r 20-25 m; gemmis petiolis ramulis juvenilib- pat visis; fructu obovoideo-oblongo 8-9 cm longo 4 cm lato. Tree 20-25 m tall. Vegetative buds, petioles, and young stems densely hirsute with spreading- ascending hairs 0.2-0.4 mm long. Petioles 3-9 mm long (on young shoots 35 mm); leaf blades crowded, becoming subverticillate at the ends of branches, 3-5 cm long, 0.9-3 cm wide, coria- ceous, shining and deep green above, paler green below, oblong-obovate, rounded and minutely mucronulate at the retuse apex, narrowed to a subacute to subobtuse base, on sterile shoots to 10-11 x 3-6 cm; lateral nerves 5-7 each side, ascending at an angle of 50-60°, branching before reaching the margins, the principal lateral nerves faintly impressed above, slightly elevated below, 2-6 mm distant or on sterile shoots 10-12 mm; tertiary venation beneath faintly to moderately prominent with an irregularly reticulate pattern. ies pendent, i eas obovoid-oblong, 8- m long, 4 c iam., minutely stellulate-ve- Saa `D STEYERMARK— VENEZUELAN GUAYANA FLORA —III 643 1987] Paratypes. VENEZUELA. BOLÍVAR: Quebrada Los Brasileros, 4.5 km al SW de Icabarü, 4?20'N, 61?48'W, 0 m, 16 Dec. 1978, Weed Carreño Espinosa & Dunsterville 117778 (MO, VEN). TERRITORIO FE- DERAL AMAZONAS: Cerro Neblina, south of Base Camp, Río Mawarinuma, 0*50'N, 66°11’W, 150—350 m, Gen- try & Stein 47115 (MO). This taxon differs from other species of the genus in having the pubescence of the buds, pet- ioles, and young stems of elongate, spreading- ascending stiffish, hirsutulous hairs rather than the densely appressed, short-stellate tomentum or glabrity of the other members of the genus. The paratypes differ from the type collection in having larger leaves widest near, instead of above, e middle, but possesses identical elongated, hirsute pubescence. rr arl Catostemma ,Sp.nov TYPE: Venezuela. Territorio Federal ini. zonas: Departamento Atabapo, Cerro Ma- rahuaca, “Sima Camp,” south-central por- tion of forested slopes along eastern branch of Cafio Negro, 3°43’N, 65°31'W, 1,140 m, 28 Feb.-1 Mar. 1985, Julian A. Steyermark & Bruce Holst 130878 (holotype, MO). Fig- ure 6C, c. Arbor 20-30 m, ramulis Bonis is glabris; gemmis glabris; petiolis 0.7-3.2 cm longis glabris; foliorum laminis elliptico-ovatis vl fe ellipticis ad me- dium latioribus apice rotundatis obtusis vel subacutis interdum mucronulatis basi obtusis vel subrotundatis 8-14 cm longis 3.5-8 cm latis utrinque glabris, nervis lateralibus utroque latere 9-12 plus minusve fere ad venulis tertiariis subtus tenuibus nec elevatis plus mi- nusve subparalle lis atque subhorizontalibus; floribus non visis; fructu maturo ellipsoideo 2-3- obtuse costato vel subangulat 2-sper- mo fide Holst & Liesner) 8-9 cm longo 4-4. 5 cm diam. (fructu immaturo magis elongato angustiorique stria- tulo indumento minute olivaceo-tomentello munito). Tree 20-30 m tall, the branches glabrous. Bud scales glabrous. Petiole 0.7-3.2 cm long, gla- brous; leaf blades chartaceous or subcoriaceous, brittle, elliptic-ovate, oblong- or lanceolate-ellip- tic, wiss: at the middle, pounced, boss or sub- ute at pex, obtuse Or T cba at the base, 8-14 cm long; 3.5-8 cm wide, glabrous both sides, the midrib im- pressed above, subelevated below; lateral nerves slender, 9-12 each side, impressed above, uni- formly ascending at an angle of 45-50°, subpar- allel and unbranched nearly to the margins, 3- 15 mm distant; tertiary venation subimpressed above, faintly subimpressed below and rather in- conspicuous, not elevated, forming a + subpar- allel and subhorizontal pattern of veinlets. Ma- ture fruit ellipsoid, 2-3-obtusely costate or subangled, unilocular, 1-(2-testa Holst & Lies- ner)-seeded, 8-9 cm long, 4—4.5 cm diam. (im- mature fruit longer and narrower after shrink- ing), striatulate with minute olivaceous indument. Paratype. VENEZUELA. TERRITORIO FEDE AMAZONAS: Cerro Marahuaca, same locality as type, as Feb. 1985, Steyermark & Holst 130709 (MO). The venation of the lower surface in this species resembles that found in Catostemma fragrans Benth. of Guyana, but C. marahuacenses differs in having glabrous bud scales and less prominent tertiary venation on the lower leaf surface. The tertiary veinlets resemble those found in some species of Scleronema, such as S. neblinense, but C. marahuacenses differs in its elongate ellipsoid, instead of globose, fruit. Catostemma pubistylum Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Ama- zonas: IVIC main study site, 4.3 km NE of San Carlos de Rio Negro, 1°56'N, 67°03' W, 119 m, 3 Aug. 1978, Howard L. Clark & P. Maquirino 6742 (holotype, NY). Arbor 4—5.5(-8) m; gemmis minute adpressoque stellato- 5. ramulis juvenilibus glabris; petiolis (5-)12-15 mm longis glabris; foliorum laminis obov- ato- pane apice rotundatis retusisque basi rotun- datis vel leviter subcordatis majoribus 9-16 em longis roque e latere 5 irregulariter reticulatis; pedicellis 2-2.5 cm longis mi- que adpresso-stellato- tomentoso ebrac- rte apicali strigosa ceteru ongo om nino stellato-tomentello, ramis 4 mm longis stellatis. Tree, 4-5.5(-8) m tall; buds minutely ap- pressed stellate-tomentose; young branchlets gla- brous. Petioles (5—)12—15 mm long, glabrous; leaf blades thick-coriaceous with revolute margins, shining above, oblong-obovate, rounded, retuse and mucronulate at apex, narrowed to a rounded or slightly subcordate base, the larger ones 9-16 cm long and 4.5—6.8 cm wide, glabrous both sides; lateral nerves 5—6 each side, ascending at an angle of 30-45*, prominently elevated below, 10-25 mm distant on the larger leaves, branching before slightly prominulous veinlets. Pedice long, minutely stellulate-tomentose, ebracteo- late. Calyx densely stellate-tomentose without, 644 the tube 4 mm long, 4-5 mm above, the lobes ovate, obtuse, or rounded, 6 mm long, 3.5-4 mm wide, glabrous within except for the strigose api- cal portion. Petals ligulate-oblong, rounded at apex, 16 mm long, 6.5 mm wide. Stamens 45- 55; filaments up to 9 mm long; anthers 0.5-1 mm long. Ovary suborbicular-ovoid, 3 mm long, 3 mm broad at base. Style 6 mm long, stellate- tomentellose throughout; style branches 4 mm long, stellate-pubescent; fruit unknown. bibi (Ad VENEZUELA. TERRITORIO FEDERAL : same locality as type collection, Clark & l. 8115 (MO). This species Is characterized by a combination of completely stellate style and stylar branches; minutely appressed-stellate bud scales; glabrous petioles, stems, and leaf blades, the latter strongly revolute; and strigose apical inner portion of the calyx lobes. Catost losi Stey k, sp. nov. TYPE: Venezuela. Territorio Federal Ama- zonas: Mari's bana (low Amazon caatinga), 10.8 km NE of San Carlos on Solano Road, 1°56'N, 67*03'W, 119 m, 16 Aug. 1981, H. L. & K. Clark 8117 (holotype, MO). Frutex 2-4 m, ramulis praecipue ad vel partim microscopico stellato- -pu ube rulis; g eci glabris vel a exudato m petiolis 2- 5; mm longis plerumque glabris vel partim microscopico stellato- puberulis; foliorum laminis ma- disnositit calyce extus dense stellato-tomentoso, tubo 3 mm longo 4 mm lato, lobis late ovato-oblongis ob- tuse acutis 10-12 mm longis 5-6 mm latis intus glabris; petalis 15 mm longis 6-8 mm latis; stylo 9-11 mm longo glabro, ramis ^ mm longis glabris vel pilis stel- latis minutis munit Shrub 2-4 m tall, the branches mainly gla- brous or partly stellulate-puberulent with micro- scopic indument. Buds mainly glabrous or some- times furnished with a microscopic pale exudate. Petioles 2215 mm long, mainly glabrous or partly stellate-puberulent with microscopic indument; mature leaves thick, rigid, coriaceous, strongly ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 revolute, obovate, rounded and retuse at the usu- ally mucronulate apex, 4.5-15 cm long, 2-4.5 cm wide, glabrous both sides; main lateral nerves 4—6 each side, scarcely evident and impressed or subsulcate above, usually elevated and conspic- uous below, 5-20 mm distant; tertiary venation above with a minutely cellular impressed retic- ulum surrounded by a slightly more impressed network, reticulate below with mainly finely im- pressed veinlets, some of these obliquely trans- verse between the main lateral nerves. Flowers few, (1-)2-7-fasciculate; pedicels 2.5-3 cm long, densely stellate-tomentose, bracteolate, the 2 bracteoles alternately situated at or below the middle, deltoid-ovate, obtuse, 0.8-1 mm long. Calyx densely stellate-tomentose without, 3 mm ong, 4 mm wide above, the lobes broadly ovate- oblong, obtusely acute, 10-12 mm long, 5-6 mm wide, glabrous within. Petals ligulate-oblong or ligulate-spatulate, rounded, 15 mm long, 6-8 mm wide. Stamens 35-40; filaments 9-12 mm long; anthers 0.8-0.9 mm long. Style 9-11 mm long, glabrous except stellate in basal 1.5 mm, glabrous or partly minutely stellulate, with style branches 2 mm long. This species differs from Catostemma altsonii in its shrubby habit and low stature and in its thick coriaceous and strongly revolute leaf blades with only 4-6 main lateral nerves on each side. — Catostemma sclerophyllum Ducke, Trop. Woods 50: 39. 1937. TYPE: Brazil. Amazonas: near Manaos, 20 May 1936, A. Ducke (holotype, RB no. 29040). Medium-sized tree. Bud scales and branches glabrous. Petioles 2-5 cm long, glabrous; leaf blades congested at the apex of the branches, oblong-elliptic, rounded and retuse at apex, sub- cordate or rounded at the base, 6-10 cm long, 4—7.5 cm wide, glabrous both sides, the main lateral nerves 6-10 each side, branching before reaching the margins; tertiary venation irregu- larly reticulate between the main secondary lat- eral nerves, less conspicuously elevated below than the main lateral nerves. Pedicels 2.5-5 cm long, moderately stellate-tomentose, bracteolate; bracteoles alternate, immediately subtending the calyx, 1-1.5 mm long, | mm wide, broadly ovate, obtuse. Calyx stellate-tomentose without, 10-12 mm long, the tube 4-5 mm long, the lobes 5—6 mm long, glabrous within. Petals obovate-ob- long, 12-15 mm long. Stamens 30-35; filaments 7-9 mm long; anthers 0.5-0.7 mm long. Style 6- 7 mm long, glabrous except stellate-tomentose 1987] in the basal 2 mm portion; style branches 2-2.5 mm long, glabrous. Fruit ellipsoid, 6-7 cm long. Distribution. Amazonian Brazil. re gundihciy examined. BRAZIL. AMAZONAS: Ducke 20-5-1936, Herb. Jard. Bot. Rio de Janeiro M (holotype, RB; beri MO); Man- A do Aleixo, Ducke 490 (F, US); Manáos, minis oni Ducke 749 (F, US); 1469 (F); same locality, Steward et al. (NY). Seen also near Santa Izabel on Rio Negro by Ducke. This species resembles C. commune but differs in having the bracteoles on the pedicels imme- diately subtending the calyx and alternately ar- ranged; longer, less stellate-tomentose pedicels; somewhat longer petals; and more rigid and thicker leaves, subcordate to rounded at the base. From C. fragrans it may be distinguished by hav- ing glabrous buds and young stems and by having completely different venation of the lower leaf surface. SCLERONEMA The genus Sc/ has hitherto been known only from Brazil and Guyana, the most recently described species being S. guianense Sandw. (1948). The following species is newly described as the first one known from Venezuela. Scleronema nebli St k, sp. nov. TYPE: Brazil. Amazonas: Serra de Neblina, vicin- ity of Base Camp, Cano Tucano, Rio Cau- aburi, 100 m, 15 Nov. 1965, Bassett Ma- guire, Julian A. Steyermark & Celia K. Maguire 60181 (holotype, MO; isotype, flowers, NY). Arbor 20-40 m, ramulis juvenilibus stellato-tomen- tellis; gemmis dense stellato-tomentosis; petiolis 1-4 cm longis stellato-tomentosis; foliorum laminis plan- tarum incipientium simplicibus, elliptico- Mrd subito poe cuspide 1.5-3 cm longo, 14-24 cm longis, m latis utrinque see iliorum laminis maturis late ° oblongis vel ae apice ro- l4 xps 3.5-7.5 cm latis subtus praeter costam mediam nervis lateralibusque sparsim stellatis sabre. 1 ne libus EE latere 7- I: subtus prominenti vatis plus m argines uniformiter ad- scendentibus subparallelisque; venulis tertiariis subtus talibus; „pedicellis (1.3-)1. 8- 2.5 cm longis stellato-to- s; calyce extus dense stellato-tomentoso lobis tribus intus moderatim pubescentibus pilis hirsutulo- strigosis praeditis; staminibus 14-16, filamentis apice dilatatis 7-8 mm longis, antheris sessilibus 0.2-0.3 mm longis; stylo 13.5 mm longo, parte basali 4 mm longa STEYERMARK — VENEZUELAN GUAYANA FLORA-—III 645 stellata, ceterum glabro; fructu globoso 7 x 7 cm sub- ruguloso subglabrescente. Tree 20-40 m tall, the young branches stellate- tomentose. Buds densely stellate-tomentose. Seedling leaves simple, elliptic-oblong, abruptly long cuspidate, 14—24 cm long, 5.5-7.5 cm wide, the cuspid part 1.5-3 cm long, glabrous both sides. Petioles 1-4 cm long, stellate-tomentose; mature leaf blades chartaceous, oblong or ob- ovate, rounded and sometimes mucronate at apex, rounded to subacute at base, 5.5-14 cm long, 3.5-7.5 cm wide, glabrous above (a few stellate hairs sometimes on midrib), the lower surface glabrous; lateral nerves 7-13 each side, impressed above, prominently elevated below, 5-15 mm distant, + uniformly ascending at an angle of 45-55?, + subparallel and unbranched nearly to the margins; tertiary venation promi- nent and elevated below with the veins + reg- ularly subparallel and subhorizontal. Pedicels (1.3 —2.5 cm long, ebracteolate, densely stel- late-tomentose with shorter and longer hairs in- termixed, the shorter brownish hairs predomi- nating. Calyx minutely densely stellate-tomentose without, tube campanulate, 4 mm long, 4 mm broad at summit, the 3 lobes 4.5 mm long, 4 mm wide, moderately hirsute-strigose within with hairs 1 mm long. Petals lance-oblong, obtuse, 13 mm long, 4 mm wide. Stamens 14-16; filaments 7-8 mm long; anthers 0.2-0.3 mm long. Style 13.5 mm long, stellate-tomentose in the basal 4 mm. Fruit globose, 7 x 7 cm, faintly rugulose, subglabrescent. Paratype. M TERRITORIO FEDERAL á Stein 47148 (fruiting material with one flower pres- ent, MO), 47152 (seedling leaves, MO); same locality, 11 Mar. 1984, Liesner 16539 (MO). This taxon possesses small sessile anthers ter- minating the relatively few dilated ñlaments and indehiscent, globose fruits. The morphologies of the anthers and the dilated filaments are typical of Scleronema, and globose fruits are known from other species of the genus, in contrast to the more elongated ellipsoid or oblong fruits of Catostem- ma. Others features of S. neblinense shared by the known species of the genus are the relatively few stamens, the relatively few flowers of each axillary fascicle, and the more or less character- istic tertiary venation and subparallel nen nerves. While the calyx lobes of Catostemma are regularly three, those of Scleronema vary from three to five. The three calyx lobes of S. nebli- 646 nense are also found in S. micranthum Ducke, to which it is most closely related. The former differs in the rounded apex of the perfectly glo- bose fruit and in the greater number of lateral nerves of the leaf blades. LITERATURE CITED BAKER, J. G. 1888. Catostemma fragrans Benth. Hooker's Icon. Pl., pl. 1793. BENTHAM, G. 1843. Contributions toward a flora of South America —enumeration of plants collected by Mr. Schomburgk, in British Guiana. London 5. 0. Plantes bir beg ou peu connues e la region amazonienne. Arch. Jard. Bot. Rio de Janeiro 5: 163-165. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 —. 1937. New forest trees of the Brazilian Am- n. Trop. Woods 50: 37-39. one D. 1891. Catostemma fragrans. Hooker’s Icon. Pl., pl. 1 PAULA, J. E. 1969. Estudios sobre Bombacaceae — I. Ci. e Cultura 21: 19. SANDWITH, l N British Guiana. Bull. Misc. Inform. 1928, no. 9: 365-379. 1931. Contributions to the flora of Tropical America. IV and V. The baromallis of British Guiana. Kew Bull. 1931: 46-54. 1948. Contributions to the flora of Tropical America. XLVIII. Kew Bull. 1948: 304—306. SHEPHERD, J. D. & W. S. ALvERSON. 1981. A new Catostemma (Bombacaceae) from Colombia. Brittonia 33: 588-590. THEACEAE BONNETIA KEY TO RECENTLY DESCRIBED TAXA OF BONNETIA la. Anthers broader than long or as broad as long, subreniform; ovary incompletely trilocular lb. Anthers longer than broad, oblong or suborbicular; ovary completely trilocular 2 Style undivided . euryanthera 2 3 Lateral nerves elevated on upper surface, impressed on lower surface; leaf blades 2-4 x 1.5- 5 cm . tepuiensis ve peia blades 1-2 x 3b. Lateral nerves ewe sia on upper surface, mainly not evident on lower surfac 4c N c 4a. Style shallowly 3-lobed i Ps divided nearly or all x length into 3 branches, or merely shallowly 3- lobed at the tepuiensis isi minor eps Emo quinimae 4b. vid deeply parted into 3 branches, divided nearly all the way to the base or at least V "0 from top P Petals white or p 6a. Uppe oui send or 3-parted to a. Petals 9-9.5 x ink r leaf ms e sepals subacute or obtusely acute; styles divided ?^ of ba 4-6 mm; leaves 1.2-2.7 x 0.5-0.8 cm; sepals 9-9. 1 mm lon B. chimantensis 7b. Petals 21-22 x 14-16 mm at the summit; leaves (2—)2.5—5 x 0.8-1. 7 cm; sepals 11-12.5 mm long B. boli varensis 6b. Upper leaf Puce with impressed n above; sepals obtuse or ee at apex; styles 3-parted '4 distance from the 5b. Petals yellow B. toroniensis 8 a. Peduncle elongate, 3—6.5 cm long, often surpassing the leaves, ebracteolate cm mu 2.7-3 cm wide; lateral nerves o 9a. Petals 3.5-4.2 c leaves scarcely elev sa surfaces of th tristyla subsp. tristyla ted 9b. Petals smaller; ues nerves of both leaf surfaces prominently elevated tristyla subsp. nervosa 8b. digi lacking or at most 1.6 cm long, usually concealed by and much short than e leaves, verticillate-bracte = oO 10a. pend linear-oblanceolate, > 5-7 mm wide; petals 8 mm long; — es 8- 16 mm long B. huberiana 10b. mig broadly lanceolate, obovate, or oblong-lanceolate, 4-15 mm wide (leaves n vegetative shoots often wider); petals 9-16 mm long; peduncles 3-16 mm ong 11a. Leaves lanceolate, acute at the apex, only Ec» ipli ug at ag base, width for most of len nearly the same w th, 3.5-4 aed impressed-nerved beneath, not pale punctate there, ihe Ph aum B ifest; sepals 12-13 m Leaves oblanceolate or obovate, obtuse, rounded t m long ptariensis tthea conspicuously narrowed : the base, broadest above the oet 1-3. S x Is] tomata yen manifest; sepals 9-10 mm ines e wurdackii 1987] In the first installment of the Flora of the Ven- ezuelan Guayana (Steyermark, 1984), four new taxa of Bonnetia were described. Continued ex- ploration of the summit floras of the Guayana Highland has yielded the following five new bon- netias. Bonnetia bolivarensis Steyermark, sp. nov. TYPE: Venezuela. Bolivar; Ptari-tepui, cumbre, 5°47'N, 61°47'W, 2,400 m, 19 Nov. 1984, Otto Huber 9818 (holotype, VEN; isotype, MO) Frutex 1 m; foliorum laminis dense rosulatis sub- sessilibus ber Vier dr apice obtuse acutis basi obtusis (2-)2.5-5 .7 cm; sepalis 11-12.5 x 5 mm; petalis 2122 x 14-16 mm; lag tribus 3-3.5 mm longis fere usque ad basem divisis Shrub 1 m tall. Leaves crowded at summit of branches, oblong-lanceolate, subsessile, nar- rowed to an obtusely acute apex, obtuse at base, (2-)2.5-5 x 0.8-1.7 cm, faintly impressed-nerved both sides (the midrib slightly elevated below) or the lateral nerves not evident. Flowers soli- tary, sessile or subsessile; bracts immediately subtending flower oblong-lanceolate, acute to obtuse, 11-12 x 4.5 mm, dorsally carinate, se- tulose marginally with dark setae 1 mm long. Sepals lance-oblong, subacute, 11-12.5 x 4.5 mm, obtusely dorsally keeled basally and api- cally; petals white, subcuneately obovate, sub- truncate apically with unequally rounded sides, narrowed to the base, 21-22 mm long, 14-16 mm wide at summit, 4 mm wide at base. Sta- mens numerous, multiseriate; filaments 5 mm or less long; anthers .6 mm. Pistil 9 mm long, the styles 3, 3-3.5 mm long, divided about 25 of their length. The larger flowers and larger subacute leaves differentiate this taxon from B. chimantensis Steyerm., B. tepuiensis Kobuski & Steyerm., and B. toronoensis Steyerm. In its deeply 3-parted style it further differs from B. tepuiensis. Bonnetia euryanthera Steyermark, sp. nov. TYPE: Venezuela. Bolivar: meseta de Jaua, cumbre, sección oriental-central, afloramientos de piedra arenisca en sitios expuestos con ve- getación herbácea y arbustos achaparrados, 4°35'N, 64°15’W, 2,020 m, 14 Feb. 1981, Julian A. Steyermark, Charles Brewer-Ca- rias & Ron Liesner 124311 (holotype, NY; isotypes, MO, VEN). Figure 7 Subfrutex 1 m, ramulis parum ramosis; cicatricibus STEYERMARK— VENEZUELAN GUAYANA FLORA III 647 valde confertis; foliis alternis ad apicem confertis ob- m latis, nervis lateralibus vix manifestis; inflorescen- tia terminali conferta dense cymosa, 12-15 flora, ax- S datis 3-4.5 mm longis 1.5-2 mm latis marginibus ver- vel subsessilibus; sepalis 5, inaequalibus exterioribus ovalibus vel suboblongis rotundatis 5-6 mm longis 3- is mm latis, interioribus majoribus ovatis obtusis 6-7 m longis 4 mm latis dorsalibus carinatis; petalis 5 suborbiculari, obovatis apice rotundato-subtruncatis 10-11 mm longis apicem versus 6-8 mm latis; fila- mentis numerosis 2-seriatis; antheris subreniformibus latioribus quam longioribus 0.5 mm altis 0.6-0.7 mm s " thecatis; ovario a mad imperfecte 3- locu- ietali placentis ;Sty- lis trifidis, ramis 1- ‘I. 25 mm lonis Dwarf shrub 1 m tall, sparsely branched with the leaf scars closely crowded, about 2 mm dis- tant. Leaves alternate, closely crowded near the summit, coriaceous, oblong to obovate-oblong, rounded or obtuse at apex, sessile or subsessile, 3-4.5 cm long, 1-2 cm wide; lateral nerves scarcely evident on either side. Inflorescence ter- minal, compact, densely cymosely subpanicu- late, 2 cm high, cm wide with 2-3 short axes 3-5 mm long, 1.5 mm wide, 12-15-flowered, on a short peduncle 2 mm long, 1.5 mm wide. Brac- teoles ligulate-oblong, rounded, 3-4.5 mm long 1.5-2 mm wide, with minutely setulose margins especially manifest in lower half. Flowers sessile, crowded. Sepals 5, unequal, the outer oval or oblong, rounded, 5-6 mm long, 3-4 mm wide, the inner larger, ovate, obtuse, 6-7 mm long, 4 mm wide, dorsally carinate. Petals 5, pink, rose and white, suborbicular-obovate, subtruncate- rounded at apex, 10-11 mm long, 6-8 mm wide near apex, 2 mm wide at base. Stamens numer- ous; filaments filiform, 3-5 mm long, 2-seriate, attached at base to a thickened tissue; anthers 4-celled, subreniform, broader than long, 0.5 mm high, 0.6-0.7 mm wide, the lop d Sery l or subelliptic. Pistil 5.5 mm long; o long, l-celled (incompletely 3- elled), the he. centae parietal and conspicuously intruding to- ward center; ovules numerous. Style 1 mm long, trifid, the style branches 1-1.25 mm long; stig- mas terminal. Paratype. VENEZUELA. Same locality and date as type, E Brewer-Carias & Liesner 124322 (NY, VEN). This species is remarkably distinct in the shape of the anthers, which are broader than long or 648 20mm G 0.7mm ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 4.5cm 5.5mm lace euryanthera. — A. Fertile branch in bud stage. — B. Portion of iioi. branch enlarged. — . Flower, lateral view. gus Petal. tu dinal aces oe howing placentae and o —E. Anther, poe view. — F. Anther, dorsal view.—G. Pistil.—H. Longi- Cross eis through ovary showing single stil s : locule occupied by sea parietal placentae with ss ie pw on holoty as broad as long and subreniform to suborbic- ular. Other distinctive features are found in the incompletely trilocular ovary with parietal pla- centae extending toward and almost fusing at the center, their ovules crowded and touching the ends of the placenta dissepiment, in this respect simulating the drawing of the ovary of Bonnetia celiae Maguire (Maguire, 1972, fig. 22k), which is quite different from the completely trilocular ovary of B. neblinae Maguire (Maguire, 1972, fig. 23g). Several dissections of the ovary verify observations which were also confirmed by bo- tanical colleagues at the Missouri Botanical Gar- den. A longitudinal section of the ovary shows that the placental partitions are not united. They nearly come together, but a slight space develops angustifolia Maguire (A, b-c) nifolium (B, b—c), Bonnetia neblinensis Maguire (D, b=), and B. steyermarkii Kobuski (F, b-c). In this respect they differ from the linear-oblong anthers of Bonnetia jauaensis Maguire (Maguire, 1976) collected on another part of the summit of Cerro Jaua of the Meseta de Jaua. Cronquist (1981) stated that the gynoecium of all four subfamilies of the Theaceae has axile placentation, **(2-2)3-5(-10) carpels," and forms a compound ovary with equal ovule and carpel 1987] number, except for Piquetia, which has the car- pels united only at the base. In his treatment of the Bonnetiaceae, Maguire (1972) stated that the ovary is **5-3-locular" and that the placentation is *axial or a permutation thereof." In our species, it would be more accurate to state that the ovary is incompletely trilocular, or actually unilocular with three parietal placentae extending inward but not fusing or unite Bonnetia guaiquinimae Steyerm., sp. nov. TYPE: Venezuela. Bolívar: Cerro Guaiquinima, cumbre, sector SE, 5?40'N, 63?26'W, 1,250 m, 26 May 1978, Julian A. Steyermark, Paul Berry, G. C. K. & E. Dunsterville 117421 (holotype, MO). Frutex 1.5 m; foliorum laminis subpetiolatis oblon- vatis integerrimis; sepalis minute mucronatis 10-12 x m —20 mm; filamentis 3.5-7 mm locis stylo subulato apice leviter 3-lobato. Shrub 1.5 m tall. Petiole 1-2 mm long; leaves coriaceous, entire, oblong-lanceolate to oblan- ceolate, acute at apex, gradually narrowed to a subacute or subobtuse base, 5-6.5 x 1.3-1.6 cm, enervate below, the midrib subimpressed below, the lateral nerves elevated above. Sepals coria- ceous, suborbicular-obovate, rounded at a short- ly cuspidate apex, 10-12 mm long, 6-8 mm wide above the middle, 3-4 mm wide at base. Petals white, obovate, narrowed to a niin mid base, 20 mm long, 15-20 mm wide at the sum- mit, 2-3 mm wide at base. Filaments Mi une 3.5-7 mm long; anthers 1.5-1.8 x 0.7 mm. Pistil 9 mm long; style united, merely 3-lobed at apex. This taxon is characterized by the shallowly 3-lobed style and the entire, oblong-lanceolate, acute leaves, which are enervate beneath. It may be distinguished from B. chimantensis Steyerm. by the larger petals, shallowly trilobed style, and larger leaves enervate beneath. It differs from B. toronoensis Steyerm. in the larger sepals and pet- als, and larger, entire leaves enervate below, while it is differentiated from B. tepuiensis and subsp. minor Steyerm. by the larger, minutely mucro- nate sepals, longer filaments, and completely en- tire leaves. Bonnetia ptariensis Steyerm., sp. nov. TYPE: Venezuela. Bolívar: Ptari-tepui, cumbre, 5?45'N, 61?45'W, 2,360-2,420 m, 23 Feb. 1978, Julian Steyermark, V. Carrefio E., R. STEYERMARK — VENEZUELAN GUAYANA FLORA III 649 McDiarmid & C. Brewer-Carias 115645 (holotype, MO). Frutex 2.5 m; foliis sessilibus lanceolatis apice acutis majoribus 3.5-4.5 x 1-1.5 cm minute obscureque ser- rulatis; floribus solitariis minute ae pedunculatis, pedunculis teretibus vel subteretibus 4—6 cm longis; nes lanceolatis vel suboblanceolatis acutis 12-13 x m; petalis luteis obovatis RE rotundatis 16- 18 x T mm; antheris 0.8 x 0.7 mm; stylis tribus 6 mm longis fere usque ad basim divisis. Subshrubs 2.5 m tall. Leaves coriaceous, lan- ceolate, acute at apex, slightly narrowed to the base, the larger ones 3.5-4.5 x 1-1.5 cm and obsoletely pinnately nerved, the midrib slightly elevated below, microscopically and obscurely serrulate. Flowers solitary on (sub-)terete pedun- cle 4-6 mm long; bracts immediately subtending flower narrowly oblanceolate, acute, 12-13 x 3- 4 mm. Sepals subconvex, lanceolate or sublan- ceolate, acute, 12-13 x 3-5 mm, 10-striate, mi- nutely ciliolate, Mt ud carinate; Vine yellow, obovate, rounded above, 16-18 mm long (pre- anthesis), 12 mm wien near apex, - mm wide at base; filaments 1.5—3.5 mm long (pre-anthesis); anthers 0.8 x 0.7 mm long. Pistil 10 mm long; styles 3, 6 mm long, free nearly to the base. This species differs from the yellow-flowered B. wurdackii Maguire in the larger, lanceolate, and acute leaves, longer sepals, larger petals (even in bud), longer filaments, and longer style branches. Furthermore, the leaves do not show the pale-punctate stomata, which are clearly vis- ible in B. wurdackii. It is easily distinguished from B. tristyla by the short pedicels, smaller floral parts, and smaller leaves, and is well dif- ferentiated from B. huberiana by the larger lan- ceolate leaves, shorter pedicels, and larger floral parts. Bonnetia tristyla Gleason subsp. nervosa "Don ermark subsp. nov. TYPE: Venezuela. Ter- ritorio Federal Amazonas: Cerro X. 2 Jan. 1951, Maguire, Cowan & Wurdack 30632 (holotype, VEN; isotype, NY). subsp. B. tristyla foliis supra costa media necnon nervis lateralibus nonnullis (6-8) conspicue manifes- teque elevatis, nervis lateralibus supra inaequaliter prominentibus; petalis minoribus 25-38 mm longis re- cedit. Petiole absent or 1-2 mm long. Leaves subob- long, oblong-oblanceolate, or oblong-obovate, obtuse to rounded at apex, narrowed to a sub- obtuse or subacute base, 4-8 cm long, 1.5-3.5 cm wide, the midrib on upper surface and 6-8 650 pairs of lateral nerves conspicuous and elevated, the other intermediate pairs of lateral nerves less conspicuously and lightly impressed, nerves on lower surface subequal and lightly impressed. Pe- duncle 3-4.5(-6.5) cm long. Sepals 12-20 mm long, the outer 12-15 mm long, the inner ones 15-20 mm long. Petals 25-38 x 15-22 mm. Paratypes. VENEZ ELA. TERRITORIO FEDERA acana, summit, 3?45'N, 66?45'W, 1,000-2,000 m, ena & Bunting 103103; Yapacana, 1,200 m, Maguire et al. 30665; Cerro Avispa, Río Siapa, sum- mit, 1?30'N, 65*51'W, 1,510 m, Cardona 3098; Neb- lina, summit, Canon Grande slopes E of summit camp “1,200-2,200” m, Maguire et al. 42179; Neblina, Cañ- on Grande SSW of summit camp, 1,050-1,100 m, Ma- guire et al. 42498; Yapacana, Maguire et al. 30632; Neblina, Cañon Grande, slopes E of summit camp, 1,200-1,300 m, co al. 42235; vicinity of Cerro Vi nilla, 30 km Ocamo, caños affluent to Río Huber 6168; altiplanicie de arenisca, E side of Río Siapa or Matapire, slightly above leaving the Macizo of Aracumuni, 1?36'N, 65?41'W, 600 m, Huber 6006 This taxon differs from typical Bonnetia tri- styla in having smaller petals, prominently ele- vated upper midrib, and unequally prominent lateral nerves on the upper surface of the leaf, with 6-8 conspicuously elevated pairs alternat- ing with fine, lightly impressed alternate nerves. There is also a tendency for the base of the leaf blade in subsp. tristyla to be cuneately narrowed, whereas in subsp. nervosa, the base of the leaf blade is usually slightly obtusely curved or rounded above its junction with the petiole. LITERATURE CITED CRONQUIST, A. 1981. An Integrated System of the C sini w Flowering Plants. Columbia Univ. Press, New ce B. 1972. The as of the Guayana High- m. New York Bot. Gard. D, b-c; F, b-«; 142, Theaceae. In J. A. Steyermark & C. Brewer- Carias (editors), La Vegetación de la Cima del Macizo de Jaua. Bol. Soc. Venez. Ci. Nat. 1984. Flora of the Venezuelan LI. Ann. Missouri Bot. Gard. 71: 326- NOTES ON LAPLACEA The last revision of the genus Laplacea, by Kobuski (1950), recognized nine species. Two of ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 these, L. acutifolia (Wawra) Kobuski and L. ob- ovata (Wawra) Kobuski, were considered by Wa- wra (1886) as varieties of Laplacea semiserrata, or of Wikstroemia fruticosa by Blake (1918). Kobuski noted that great variation existed in size of the flowers and fruits, even within a single species, and that the keys to the species had been “based almost entirely on leaf characters and pu- bescence." The species of Laplacea can be divided into two groups, those with asymmetrical leaves and those with symmetrical leaves, both of which occur in the Venezuelan Guayana, the common asymmetrical type represented by L. fruticosa (Schrader) Kobuski, the symmetrical type by L. pubescens Planchon & Linden. Variations occur in each species (Fig. 8B, b, 8C, c). Specimens from the Territorio Federal Amazonas (Maroa, Duida, Huachamacari, Yutaje, and Cerro de La Neblina), as well as a specimen from Cerro Sa- risarihama of the state of Bolivar (Cardona 380) (Fig. 8A, a) may be referred to typical L. fruti- cosa. However, two specimens from the summit of the Chimanta Massif (Steyermark & Wurdack 879 and 1126) (Fig. 8D, d) from the state of Bolivar, differ conspicuously from other Gua- yanan material, as well as from specimens out- side the Venezuelan Rind conforming to L. fruticosa. TI with larger leaf sc scars; thicker, larger leaves, en- ervate beneath, broader at the subsessile base as to appear nonpetiolate; rather conspicuous hairy tufts along the leaf margins in the sinuses of the marginal teeth; longer, stouter pedicels; and larg- er sepals and petals. The isolation of the Chi- mantá specimens from the upper slopes of this sandstone mountain, well known for its endemic flora, may help explain the evolutionary trend here noted toward the segregation of morpho- logical characters at variance from typical L. fru- ticosa. A specific or subspecific category for this population might be envisioned. However, in view of the degree of variation shown by other taxa of this species, varietal rank is here pro- posed. br anches Laplacea fruticosa (Schrader) Kobuski var. chi- mantae Steyermark, var. nov. TYPE: Vene- zuela. Bolivar: Chimantá Massif, central section, along west branch of headwaters of Rio Tirica above Upper Falls, 2,090 m, 17 Feb. 1955, Julian A. Steyermark & John J. Wurdack 879 (holotype, VEN; isotypes, F, NY). Figure 8D, d. VENEZUELAN GUAYANA FLORA—III 1987] STEYERMARK— URE 8. of a sinus (1).—B. Leaf of Maguire et al. 37153 Steyermark et al. 120086. c. portion of margin with detail of a sinus (3). A-C. Laplacea fruticosa var. fruticosa.—A. Leaf of Cardona 380. a. portion of margin Mas detail 3. b. portion of margin with detail of a sinus (2). — C. Leaf of — D. Laplacea fruticosa var. chimantae, 12 leaf of Steyermark & Wurdack 879. d. portion of margin with detail of a sinus showing tufts of hairs (4). A var. fruticosa a sessilibus vel subsessilibus ap- etiolum nihil angustatis basi 5— i marginalibus p o ab ids abus urs I. robustis 2—3 c s 2.5-3 crassis, sepalis petalisque majoribus . Tree 5—20 m tall, the leafy branches 5-10 mm thick, the leaf scars conspicuous, orbisilat, 2.5- 4 mm wide. Leaves sessile to abruptly contracted into a petiolar portion 1 -21 mm long; leaf blades thick-subcoriaceous, asymmetric, ob- lanceolate-oblong, rounded at the slightly emar- ginate apex, gradually narrowed to a broad base, 9.5-12.5 cm long, 3—4.8 cm wide, 5-7 mm wide at base, the midrib bordered on one side by foliar tissue 3-4 mm wide, on other side 2-3 mm wide, the lower surface enervate, mainly glabrous but with scattered, pale, minute, appressed, simple hairs over a pustulate surface, inconspicuously appressed-subserrulate from about % distance from base to apex with dark acicular teeth 1 mm long, these bearing at their base a tuft of silky hairs 1 mm long (seen best from lower margin); cocc1l upper surface enervate, glabrous. Peduncle 2-3 cm long, 2.5-3 mm thick, glabrous. Flower bud just before anthesis 2-2.5 cm long, 1.5-2.5 cm wide. Sepals broadly suborbicular, the two out- ermost 20 mm long, 15 mm wide, the others 17— 25 mm long, 13-22 mm wide, minutely gray sericeous most of the length without, glabrous within. Petals broadly oblong-obovate, broa rounded and emarginate at apex, mm long, 15-22 mm wide, minutely gray sericeous without in the central 5-9 mm portion, glabrous on the marginal 4-5 mm, glabrous within. Filaments 5- 6 mm long, glabrous; anthers up (e oblong, broadly rounded at the m long, 1.5 mm wide. Ovary barrel-shaped, 8 mm long, 5 mm wide, densely sericeous. Paratype. VENEZUELA. BOLIVAR: Chimantá Massif, Agparaman tepui, southeast-facing forested slopes be- low escarpment, 1,880-1,955 m, 26 Feb. 1955, Stey- ermark & Wurdack 1126 (F, NY, VEN). The broad, nonpetiolate leaf bases of var. chi- 652 mantae differ markedly from the generally nar- rowed, subpetiolar ones of typical Laplacea fru- ticosa, in which the leaf base is conspicuously narrowed toward the base into a subpetiolar por- tion 5-8 mm long and 1-2 mm wide with the midrib bordered on either side by a very narrow strip of tissue only 0.5-1 mm wide on each side. In L. fruticosa var. pulcherrima (Melchior) Ko- buski, the leaf base is broader than in typical L. fruticosa, but the apex is gradually obtusely nar- rowed and is not broadly rounded as in var. chi- mantae. The tufts of hairs at the base of the marginal teeth are absent from most specimens examined but are present to a lesser degree in Dusen 15451 from Brazil and Maguire et al. 37259 from Cerro de La Neblina in Venezuela. The enervate lower leaf surface may also occur in other specimens of L. fruticosa, but more fre- quently the lower surface shows venation. The second species of Laplacea in the Gua- yana Highland, L. pubescens Planchon & Linden ex Triana & Planchon, is represented by two col- lections, Maguire, Wurdack & Bunting 37272 from the summit of Cerro de La Neblina, and Maguire, Cowan & Wurdack 30285 from the summit of Cerro Huachamacari, both from Ter- ritorio Federal Amazonas. Elsewhere, the species ranges in the Andes from Venezuela south to Peru and Bolivia. On Cerro de La Neblina it varies slightly from typical L. pubescens of the Andes, whereas on Cerro Huachamacari it has developed more reduced leaves, and in both Guayanan collections the lower surface of the leaves is enervate, as contrasted with the more evident venation shown in most ofthe specimens of L. pubescens. Laplacea fruticosa, the more common species, also occurs on Cerro Huacha- macari, but at a lower altitude of 1,100 m of forested talus slopes, as well as on Cerro de La Neblina. Laplacea pubescens Planchon & Linden ex Triana Planchon : aguire, R. Cowan & J. Wurdack 30285 putida NY). var. pubescens foliorum laminis minoribus rece- .5-4.5(-9) cm E 2(-3) cm latis oe atis; pedicellis 6-8 m iis i sepalis 5-11 m pues is; petalis cuneiformi- obovat 6-21 mm longi supra medium 14-15 mm latis. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Leaf blades symmetrical, oblanceolate, nar- rowly obtuse at the apex, strongly narrowed at the base, 2.5-4.5(-9) cm long, 1-2(-3) cm wide, closely crenulate above the middle, enervate be- neath. Pedicels 6-8 mm long. Sepals 5-11 mm long. Petals cuneiform-obovate, 16-21 mm long, 14-15 mm wide above the middle. Paratype. "VENEZUELA. TERRITORIO FEDERAL AMAZONAS: Cerro de La Neblina, savanna edge, 3 km N of Cumbre Camp, 1,800 m, 12 Jan. 1953, Maguire, Wurdack & Bunting 37272 (NY). Both collections have the leaf surface enervate beneath. However, the paratype (Maguire et al. 37272) has the larger leaves typical of L. pubes- cens var. pubescens. In both collections the in- dument of the lower surface is sparse compared with specimens from the Andean portion of the range of the species. LITERATURE CITED BLAKE, S. F. 1918. New isting VL i Ed in Venezuela and Curacao by Messrs. Curran Haman. Contr. Gray Herb. n.s. 53: 39. KoBuski, C. E. 1950. Studies in the Theaceae, XX. Mese S Plantae olu mbianae, Kew Bull. Misc. Sarma te 1926: 43-44. Wawra, H. 86. Ternstroemiaceae. /n C. F. Mar- tius, Flora Brasiliensis 12(1): 289. THYMELEACEAE DAPHNOPSIS Daphnopsis guaiquinimae ipsun. sp. nov. TYPE: Venezuela. Bolív : Cerro Guaiqui- nima, INE la extensión al del cerro en el sector noreste, la parte pedregosa, abierta y plana con vegetación baja, 6°0'N, 63?28'W, 1,650 m, 9 Apr. 1979, Julian A. Steyermark, G. C. K. & E. Duns- terville 117977 (holotype, VEN; isotype, MO). Figure 9. x 1.5 m; ramulis juvenilibus sericeo-strigosis; laa laminis elliptico- oblongis vel lance A eg iod lipticis apice p tusis vel subacutis 3-6 cm longis 1-3 cm we supra glabris subtus minute strigillosis costa media subtus dense strigillosis venulis tertiariis pagina superiori ir- regulariter contortis; pedunculis filiformibus 10-20 mm oblongis subobtusis utrinque sericeo-hirtellis, orificio glabro; staminibus 8, 2-seriatis; squamis pa 7- 8 subulatis glabris fere liberis 1.7-2 mm lon 1987] Shrub 1.5 m tall; juvenile stems 2-3 mm diam., sericeous-strigose. Petiole 2-3 mm long, strigil- lose, leaf blades subcoriaceous, gray-white be- low, elliptic-oblong or lance-elliptic, mostly ob- tuse to rounded at apex, rarely subacute, obtuse to subacute at base, 3-6 cm long, 1-3 cm wide, glabrous above, minutely strigillose below, slightly more densely on the lower midrib; prin- cipal lateral nerves 7-10 each side; tertiary ve- nation above irregularly contorted and twisted, below more regularly reticulate. Inflorescence terminal or axillary, the peduncles filiform, 10— 20 mm long, 0.5 mm wide, sericeous-strigillose. Calyx subcylindric-obovoid, 7 mm long, 2 mm wide at and above middle, 1 mm wide at base, antrorsely sericeous without, densely antrorsely sericeous within in the lower 4 mm; lobes 4, unequal, the outer ligulate-oblong, obtuse, 1.2- 1.5 x 0.5 mm, theinner 1 x 0.08 mm, sericeous- hirtellous on both surfaces, the orifice glabrous. Stamens 8 in 2 series; anthers sessile, suborbic- ular-oblong, 0.6 mm long, the upper 4 opposite the calyx lobes, the lower 4 alternate with the calyx lobes; style terminal, 0.3 mm long; stigma clavate, 0.3 mm long. Hypogynous scales 7-8, subulate, glabrous, 1.7-2 mm long, cleft nearly or all the way to the base, free or nearly so. The type and only specimen of this species was indicated by Nevling (1967) as a new species of Daphnopsis related to D. longipedunculata Gilg ex Domke, described originally from Mount Ro- raima. she chief differences possessed by the idi number Oo f1 (8 vs. 4 as in thé Roraima specimen), digne. 1-2 cm long contrasted with 2.5-9 cm on the Roraima specimen, and the shorter aie Domke (1935) stated that the umbels in D. longipedun- culata were on axillary peduncles but added (“atque terminali?"), apparently suspecting that they may also be terminal, as is the case in the Guaiquinima specimens. He also stated that sta- minate as well as pistillate flowers occur. This indicates that the specimen Ule 8739 from Ro- raima studied by Domke was monoecious. In the Guaiquinima specimens, there is also an in- dication of monoecism, since the anthers are well developed and suborbicular-oblong on some flowers but poorly developed and linear-oblong on others, even on the same plant. The hypogynous scales in the Guaiquinima plant are regularly disposed as eight filiform ap- pendages. Although they appear to be free, as in Funifera utilis Leandro, they are connected at their very base by a slight membrane on the disk, STEYERMARK— VENEZUELAN GUAYANA FLORA —III 653 which appears to be adnate to the actual base of the calyx tube, thus showing the relationship with Daphnopsis. Conversely, they could be inter- preted as completely free as in the genus Funi- fera. Nevling (1967) at first interpreted the style to be lateral (eccentric) in Daphnopsis longipe- dunculata, but later (pers. comm., 1986) con- firmed my observation that the style is terminal. Daphnopsis nevlingiana Steyermark, sp. nov. TYPE: Venezuela. Bolivar: Cerro Sarisari- ñama, Meseta de Jaua, summit, 4?41'40"N, 64?13'20"W, 1,400 m, 16-18 Feb. 1974, Ju- lian A. Steyermark, V. Carreño E. & Charles Brewer-Carias 109199 (holotype, VEN; iso- type, ). A ical longipedunculata Gilg ex Domke fo- liorum paginis superio oribus venatione tertiaria irre- gulariter contorta calycis tubo tys practer infra medium sparse pier po apice | Gain ti tube seu eie cedit. Slender tree 3 m tall with bark difficult to tear, the young branches strigose. Petiole 3-4 mm long, moderately strigose. Leaf blades elliptic-oblong or ovate-oblong, rounded at apex, acute at base, 7.5 cm long, 2-4.5 cm wide; upper surface mainly glabrous except strigose along the de- pressed midrib; lower surface uniformly short- strigose with the lower midrib more densely stri- gose with longer hairs; lateral nerves 7-9 each side, faint above, slender and slightly manifest below; tertiary veins of upper surface irregularly contorted, loosely and inconspicuously reticu- late. Pistillate calyx dull red with whitish tips, cylindric, 5 mm long, 4-lobed; lobes unequal, puberulous in upper half, glabrous below, the larger ones 1 mm long, 0.8 mm wide, pubescent over a larger area with longer hairs toward the base; the smaller lobes 0.7 mm long, 0.7 mm wide, ovate-oblong; calyx tube glabrous within except for small sericeous patches below the mid- dle. Hypogynous scales 4, linear-lanceolate, 1.5— 2 mm long, 0.2 mm wide in lower part, dull yellow, dull reddish near the tip. Pistil 4 mm long; ovary ellipsoid, 2.4 mm long, densely se- riceous; style 1.5 mm long, glabrous; stigma subglobose, 0.5 mm long. This species differs from Daphnopsis longi- pedunculata Gilg ex Domke by having peculiarly contorted, irregular tertiary venation on the up- per leaf surface; the calyx reddish white with white tips and with glabrous interior except for sparse 654 00 0. a UN OL / 7mm FiGurE 9. Daphnopsis guaiquinimae.— scales. — D. Peduncle with flowers. — E. Gro pubescence below the middle; and hypogynous scales thickened and dull yellow with reddish near the obtuse tips. SCHOENOBIBLUS Schoenobiblus amazonicus Steyermark, sp. nov. TYPE: Venezuela. Territorio Federal Ama- zonas: 25 km S of Puerto Ayacucho, 5?30'N, 67?35'W, 5 Aug. 1967, Wessels- Boer 1953 (holotype, MER; isotype, MY). Figure 10. Arbor 6 m; foliorum laminis oblanceolato-obovatis apice acuminatis base acutis 36 cm longis s cm latis subtus praesertim costa media necnon nerv teralibus puberulis; petiolis subtus dense strigillosis calycis tubo lineari 17-18 m extus sericeo intus supra e retrorse sparsimque strigoso ceterum glabro, lobis quattuor linearibus apice ANNALS OF THE MISSOURI BOTANICAL GARDEN — A. Flo a branch.—B. oup of flower [VoL. 74 Flower, interior view.—C. Hypogynous rotundatis 6 mm longis 1.5 mm latis extus sericeo- i š lanuginosis intus villosis; stylo filiformi 18 mm longo Tree 6 m tall; buds fusiform, subobtuse, 6.5 mm long, 2.5 mm wide. Petiole 6-9 mm long, 2.5-3 mm wide, strigillose beneath; leaf blades oblanceolate-obovate, acutely acuminate at apex, acute at base, 36 cm long, 12.5-14 cm wide, glabrous above, moderately strigillose on midrib and lateral nerves below, these sparsely puber- ulous on leaf surface, with the midrib elevated, r ceous axes, one of them solitary, the other forking into 2-3 shorter secondary axes 2-4 cm lon Flowers 10-20-umbellate; pedicels 10-18 mm 1987] 0. popa E amazonicus. — A. Flowering branch. — B. Flower. FIGURE 1 E Sent long. Calyx tube linear, 17-18 mm long, 0.6-0.8 mm wide, short-sericeous without, retrorsely sparsely strigose within in the upper half, gla- brous in the lower half; lobes 4, spreading, rev- olute at margins and at apex, linear, rounded at apex, 6 mm long, 1.5 mm wide, sericeous- lanuginose without, villosulous within. Stamens 4; anthers suborbicular or broadly oblong, 1.1 mm long, mm b exserted, erect-ascending, 5-5.5 mm 1 brous. Style filiform, 18 mm long, glabrous, at- taining the orifice; stigma ellipsoid, 1.2 mm long. STEYERMARK — VENEZUELAN GUAYANA FLORA —III 655 AN F DOM EP APP EE Dr ER eee See nga Sart ere —C. Stamen, dorsal view.—D. The new taxon differs from Schoenobiblus daphnoides Martius of Brazil in the much broad- er leaves and less-branched inflorescence, while from other Venezuelan material identified as S. daphnoides it is differentiated by the longer, more slender calyx tube, narrower calyx lobes with shorter, gray sericeous pubescence, and much longer peduncle LITERATURE CITED , W. 1935. Neue Arten und Varietäten der Nun en Daphnopsis Mart. et Zucc. und Fu- 656 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 74 dala (Leandro ex) C. A. Mey. aus Mittel- und merika. Notizbl. Bot. Gart. Berlin-Dahlem 12: 723-1 24. MEISNER, C. F. Thymeleaceae. Jn C. F. Mar- tius, Flora Brasiliensis 5(1): 65. t. 26 (1). NEVLING, L. 1959. Revision ofthe genus Daphnopsis. p 352. 67. n J. A. Steyermark & B. i A (editors), eme of the Chimantá Massif. II. Mem. New York Bot. Gard. 17: 454. SYMPLOCACEAE SYMPLOCOS KEY TO THE SPECIES OF SYMPLOCOS OF THE VENEZUELAN GUAYANA la. Ovary 4-celled; pubescence of Sd or iiis xk at least on the upper part of the , petioles and lower leaf surface, and/or on the low ulei lb. Ovary 3- or 5-celled; sre eal absent or of abba or sericeous hairs 2 2a. Calyx lobes acute to acuminate at apex; style pubescent 3a. het E shorter than the petiole, sessile or nearly so, 2-3- hielo calyx lobes 1.8-2 m long; corolla 7 mm long; leaves minutely and distantly crenulate U... . acananensis 3b. ne equaling or longer than the petiole, shortly pedunculate, 5-8- flowered: calyx .5-3 mm long; corolla 8-9 mm long; leaves lo to subentire 0. ibaa 2b. Calyx dies obtuse to rounded at apex; style glabrous 4a. Ova celled 5a. los glabrous beneath, acuminate at apex, acute at base; petiole glabrous throughout UM 5b. Leaves sparsely pubescent beneath on the nerves, obtuse at apex, rounded or ue base; ae strigillose adaxially 5S. V rid NS 4b. Ovary 3-cell 6 6a. Calyx is without, except for the ciliate lobes; filaments not connate, free or nearly so to th bes; e below; leaves with at least the lower midrib appressed-pubescent; petioles bey iren. Or seri 7a. Calyx lobes longer than the calyx tube; leaves rounded to obtuse at apex, rounded or obtuse at cm lon . Jauensis . Calyx lobes shorter than or equaling the calyx tube; leaves acuminate at apex, acute at base, 7-10 cm long 8a. Fruit densely strigillose, 5 mm long; stamens 25-40; lower leaf n sparsely appressed-pubescent to glabrescent; bracts suborbicular, rounded or obtuse; co- rolla 3 mm lon $ yapacanensis . Fruit glabrate, 6-8 mm long; stamens 20-25; lower leaf surface glabrous; bracts acute; corolla 4-4.5 mm long S. neblinae oo c Symplocos acananensis Steyermark, sp. nov. TYPE: Venezuela. Bolivar: gallery forest bor- dering savanna, vicinity of Guadequen, Rio Acanan, affluent of Rio Carrao, west of Cer- ro Las Hermanas, 5?56'N, 62°17'W, 470 m, 1-3, 20 May 1986, Julian A. Steyermark, R. Liesner & B. Holst 131868 (holotype, VEN; isotype, MO) superne libris inferne in tubum 3.5 mm longum coa- lescentibus glabris 1.5-3.5 mm longis; stylo 5 mm lon- o in dimidio inferiore piloso. Shrub 1.5 m tall with mainly glabrous branch- es. Petioles 1.2-5 mm long, glabrous; leaf blades chartaceous, elliptic-oblong, abruptly short-acu- minate at apex, obtuse t F 1.5 m, ramis plerumque glabris; foliis pe- tiolatis, "petiolis 1.2-5 mm lo a í tiis axillaribus sessilibus vel fere 2-3 floris; bracteis sub lycis tubo cupulato 1.2-1.5 mm longo glabro, lobis 5 late lanceolatis acuminatis 2 mm longis 0.8-0.9 mm latis praeter margines ciliatos glabris; corolla 7 mm longo, tubo 2.5 mm longo, lobis 5 oblongis apice ro tundatis 4.5 mm yobis staminibus ca. 20, ments sides, appearing subentire but the margins with microscopically obtuse serrulations 0. 1—0.2 mm long, 2-4 of these dispersed in 1 cm of mar- gin; lateral nerves 4-5 each side, faintly anas- tomosing with the tertiary veinlets 4-8 mm from the margin, obsolete above, slightly manifest be- low; tertiary veinlets slightly reticulate below, scarcely or not manifest above. Inflorescence ax- illary, 2-3-flowered, sessile or nearly so, shorter than the petioles; flowers fasciculate, sessile. Bracts subtending flowers lanceolate, subacute 1 x ] mm, pubescent without, strongly ciliate, STEYERMARK 1987] —VENEZUELAN GUAYANA FLORA —III 657 with caducous black glands. Corolla 7 mm long, the tube 2.5 mm long, 1.5 mm wide; lobes 5, oblong, rounded at summit, 4.5 mm long, 2.1 mm wide. Calyx tube cupulate, 1.2-1.5 mm long, 1.5-2 mm wide above, glabrous; sa lobes 5, broadly lanceolate, acuminate, 2 mm long, 0.8- 0.9 mm wide, glabrous except for the ciliate mar- gins. Corolla 7 mm long, the tube 2.5 mm long, the 5 lobes oblong, rounded at the apex, 4.5 mm long. Stamens about 20, ca. 4-seriate; filaments free above, forming a tube 3.5 mm long, the longer filaments 1.5-3.5 mm long, the shorter ones | mm long, all glabrous; anthers broadly oblong, 0.2 mm long. Style 5 mm long, pilose in the lower half. Fruit not seen. This species is closely related to the group of Symplocos schomburgkii Klotzsch ex Schomb., S. guianensis (Aublet) Giirke, and S. pilosiuscula Brand. It is distinguished from S. guianensis by the glabrous branches, from S. schomburgkii by the sessile or nearly sessile inflorescence which is fewer-flowered and shorter than the petiole, the shorter corolla and calyx lobes, and minutely crenulate leaf margins. The new species differs from S. pilosiuscula in the acute to acuminate calyx lobes. RUBIACEAE Chomelia glabricalyx Steyermark, sp. nov. TYPE: Venezuela. Bolivar: primary forest and river edge, Rio Caura, 5-10 km S of Las Pavas (Salto Para), 6°12'N, 64?28'W, 240 m, May — Gilberto Morillo 6811 (holotype, N). x 1 m, ramis glabris spinosis; foliorum laminis ovato- vel lanceolato-ellipticis apice acutis vel sub- rum atque int glabris; inflorescentia. 6- flora, pedunculo filiformi 14- E Poir m longo glabr o; 4. s ° 1 axils of the lateral nerves and sometimes with few sparse ciliate hairs at the base; lateral nerves 5-6 each side, scarcely evident above, slightly impressed below. Stipules subulate, projecting 0.5 mm long. Inflorescence 6-flowered; peduncle filiform, 14-20 mm long, glabrous; flowers not involucrate at base, sessile, or in fruit 0.5-1 mm pedicellate. Calyx and hypanthium glabrous; hy- panthium tubular, 1.5 mm long, | mm wide, glabrous; calyx E unequal, ligulate- oblong, obtuse, 0.5-1 m wide, gla brous. Corolla bei, 4 mm long, 1.2 mm wide; tube 2.5-3 mm long, glabrous without; lobes oblong, obtuse, 1.5 mm long, glabrous below, sparsely iis mas upward. Fruit narrow-cylin- dric, 10 m .5 mm wide, 0.5-1 mm long eae e rq This taxon is related to the recently described Chomelia delascioi Steyerm. and C. stergiosii Steyerm e present taxon differs from C. de- lascioi in the larger, barbellate leaves, shorter corollas, and more numerously flowered inflo- rescence, and from C. stergoisii in the shorter corolla and calyx lobes and the completely gla- brous hypanthium and calyx lobes. ae huberi Steyermark, sp. nov. TYPE: Venezuela. Bolívar: Distrito Roscio, Serra- nia es Caco, 25 km NW of San Ignacio de Yuruani, 2.5 km ESE of Wanaru, 5?12'N, 61?15'W, 1,150-1,200 m, 1 Mar. 1984, Otto Huber 9123 (holotype, VEN; isotype, MO). eta acel Herba radicans, caulibus fepenusus 0. ie mm at s 1.2 mm lon- gis sparsim | strigillosis; foliis petiolatis, ie sare mm longis dense strigosis; foliorum laminis ellipticis anguste oblongo-ovatis vel elliptico-ovatis apice $ed acutis vel rsen basi acutis vel obtusis 2-2.8 c longis 1-1.5 cm latis utrinque breviter strigillosis, ner- vis gi d utroque latere 5-7; inflorescentia 1 -flor dunculo sub anthesi 1.5 mm longo sub fructu 4.5 mm longo dense strigilloso; hypanthio dense strigoso; calycis ] ceolatis acutis 1.8—2.5 mm 5-1 it wa thioque glabro, semen 1.5 mm longo 1 mm lato, lobis inaequalibus ligulato-oblongis obtusis 0.5- 1 mm longis glabris; corolla 4 mm longa tubo extus glabro, lobis subtus glabris apicem versus sparsim stri- gillosis; fructu anguste cylindrico 10 mm longo 3.5 mm lato 0.5-1 mm pedicellato. rub ] m tall; branches glabrous, spinose. o elliptic, acute to subacuminate at apex, cuneately acute at base, 4.5—9.5 cm long, 1.5-4 cm wide, glabrous both sides except for barbellate lower longis 0.4-0.5 mm latis ieee sparsim strigillosis ad gillosa; bacca (immatura) in statu vivo 1-1.5 cm diam., in sicco 5 mm longo 4.5 mm lato modice strigilloso. Creeping herb with moderately strigillose stems 0.8-1 mm diam.; stipules linear-setaceous, 1.2 mm long, sparsely strigillose. Petioles ofthe larg- er leaves 10-13 mm long, of the smaller ones 3- 5 mm long, densely strigose; leaf blades elliptic, narrowly oblong-ovate, or elliptic-ovate, sub- acute to subobtuse at apex, acute to obtuse at 658 eidec more base, moderat abundantly on nerves d Beneath: 2-2.8 cm long, 1-1.5 cm wide; lateral nerves 5-7 each side, faint on lower side. Flowers solitary, the peduncle 1.5 mm long in anthesis, 4-5 mm long in fruit, densely strigillose. Calyx 3.5 mm long, the tube subglo- bose, 1.3 x 1.3 mm, densely strigose, the lobes unequal, lanceolate, acute, 1.8—2.5 mm long, 0.4— 0.5 mm wide, sparsely strigillose on both sides, at base within setose with erect hairs, each sinus at the base furnished with 1 squamella. Corolla white, infundibuliform, 4.8-5 mm long, sparsely strigillose without, the tube 2.8-3 mm long, the lobes broadly lanceolate, 2 mm lon 8 mm wide. Stamens not exserted, attached Uh distance of the tube length; anthers oblong, obtuse, 1.1 mm long. Style 3 mm long, glabrous. Immature fruit urceolate-globose, 5 mm long, 4.5 mm wide (dried), 1-1.5 cm diam. (living). The present taxon approaches the endemic Ja- maican species, Coccocypselum pseudotontanea Griseb. Both possess white corollas and small leaf blades, but the Venezuelan species differs in having shorter stipules, shorter calyx lobes and corolla, shorter and 1-flowered peduncles, one squamella at the base of each side of the inner part of the calyx lobe, larger fruits with appressed pubescence, and the freely rooting habit. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Perama dichotoma Poepp. & Endl. var. mono- cephala Steyerm., var. nov. TYPE: Venezue- la. Territorio Federal Amazonas: Departa- mento Atabapo, Cerro Huachamacari, ladera sur-este 175-180 m, 12-14 Oct. 1984, Fran- cisco Delascio 12358; hierba pilosa; hojas arrosadas verde oscura por el haz, verde pal- ido por el envés; pedünculo floral erecto, violaceo-morado en su base; flores blancas con el apice de los pétalos lila) (holotype, VEN; isotype, MO). A P. dichotoma var. dichotoma et var. scaposa c libus simplicibus ipei pa recedit; capitulis so so- litariis depressis 2-4 mm altis 5-6 mm latis. Herb, 4-5 cm tall. Leaves rosulate, ovate or elliptic, 7-10 mm long, 3-6 mm wide, densely pilose above, pilose on main nerves below. Heads solitary, ca. 10-flowered, depressed-hemispher- ic, 2-4 mm high, 5-6 mm wide. Corolla laven- der. Perama dichotoma has the inflorescence usu- ally consisting of several to numerous heads on filiform forking branches. The new variety has only a solitary head of flowers terminating a sim- ple peduncle. NOTES NEW COMBINATIONS IN CENTRAL AMERICAN ARACEAE Work in recent years with the Araceae of Cen- tral America necessitates new combinations in- volving species in the genera Monstera and Philodendron which were erroneously published as belonging to Rhodospatha and Syngonium re- spectively. Monstera costaricensis (Engl. & Krause) Croat rayum, comb. nov. Rhodospatha cos- taricensis Engl. & Krause, Das Pflanzenr. 4. 23B (Heft 37): 95. 1908. TYPE: Costa Rica. Limon: Ferme de Boston, Atlantic wa- tershed, 30 m elev. 10?01'N, 83?15'30"W, Tonduz 14628 (holotype, B). We had concluded that this distinctive species of Monstera, occurring in primary forest at Finca La Selva, was undescribed and had provisionally assigned it the name “Monstera undulata.” It now turns out that this species was previously described by Engler & Krause (1908) in the genus Rhodospatha; the transfer to Monstera is here- with effected. Monstera costaricensis 1s distin- guished from other species by its strongly un- dulate petiole sheaths, tuberculate petiole bases and pistils shaped like bowling pins. The lamina may be either entire (as on the holotype) or per- forate. Monstera costaricensis is now known in the Atlantic lowlands of Costa Rica from La Sel- va to Punta Mona. It probably occurs in Nica- ragua and Panama as well. Monstera dissecta (Schott) Croat & Grayum, comb. nov. Tornelia dissecta Schott, Oes- terr. Bot. Z. 8: 179. 1858. TYPE: Costa Rica. Cartago: Volcán de Turrialba, Wendland 500 (holotype, GOET). Although the name Monstera dilacerata is well known and firmly entrenched in both taxonomic and horticultural aroid literature, it has been ap- plied in a highly indiscriminate manner and, in- deed, may be impossible to confidently assign to any real biological entity. As described by Mad- ison (1977), “the type of M. dilacerata is an in- ferior specimen consisting of a few leaves from an immature cultivated plant of uncertain geo- graphic origin." Such a specimen, even if it could be located, would be virtually impossible to iden- tify with any biological species in a genus as phe- ANN. MissouRi Bor. GARD. 74: 659—660. 1987. notypically plastic as Monstera; but the type of Monstera dilacerata is apparently not extant, having been destroyed in the Berlin herbarium during World War II, and we know it only from photos. Madison (1977), determined to salvage this well-known name by any means, adopted the concept of Engler & Krause (1908). Although the latter authors applied the name M. dilacerata rather consistently (i.e., to the species here treat- ed as M. dissecta), their concept would appear to be completely irrelevant; what matters is the interpretation of the type material. It could be argued that Engler & Krause might have seen living (perhaps even fertile) material from the original collection, but they give no such indi- cation and the point is moot. Madison (1977) himself made matters signif- icantly worse by applying the name Monstera dilacerata with reckless abandon. Based on his specimen citations, herbarium annotations, and the use of his keys, Madison has employed the name M. dilacerata for no fewer than four dis- tinctly different species in Costa Rica alone (Croat & Grayum, unpubl. data). Given the expansive geographic distribution of Monstera dilacerata sensu Madison (from Guatemala to Amazonian Brazil), the application of the latter name to any one of the four Costa Rican species would have to be entirely arbitrary. Indeed, the type might just as well have come from Brazil, and may represent a different species altogether. The only alternative seems to be the relegation of the much-abused name Monstera dilacerata Schott to the limbo of nomen dubium status, where it will probably lie forever unless someone can come up with a way (epidermal anatomy?) to unequivocally interpret the type specimen, in the unlikely event that it ever turns up. Three of the four Costa Rican species in the Monstera dilacerata complex have apparently never been described (they will be described elsewhere). The fourth clearly corresponds to Tornelia dissecta Schott, the type of which is extant at GOET and has been studied by the present authors (Madi- son was apparently unaware of its existence). The specimen is a good one and agrees in all details with a species we know well from throughout Costa Rica (but mostly from the Atlantic slope) 660 at elevations of ca. 100-1,800 m. Contemporary collections from the general vicinity of the type locality include: Grayum & Sleeper 3304 (CR, MO); Grayum et al. 3490 (CR, MO); and Gra- yum & Hammel 5739 (MO). The appropriate new combination in Monstera is made above. Philodendron rothschuhianum (Engl. & Krause) Grayum, comb. nov. Syngonium rothschuhianum Engl. & Krause, Das Pflan- zenr. 4. 23E (Heft 71): 124. 1920. TYPE: Nic- aragua. Matagalpa: Matagalpa, 1,000 m, Rothschuh 229 (holotype, B). This species has long been excluded from Syn- gonium, perhaps having been first so treated by Birdsey (1955) in an unpublished thesis. Croat (1981) erroneously placed the species with P. an- isotomum Schott. There is no longer any doubt that it is a distinct species, distinguished from the latter by having the posterior lobes more ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 broadly confluent with the anterior lobe and by details of the inflorescence. The species ranges from Nicaragua to western Panama (Veraguas) from sea level to 1,000 m LITERATURE CITED BinpsEv, M. 1955. The Morphology and Taxonomy of the Genus Syngonium (Araceae). Ph.D. Dis- sertation. University of California, Berkeley, Cal- ifornia. Croat, T. B. 1981. A revision of Syngonium (Ara- ceae). Ann. Missouri Bot. Gar : 649. ENGLER, A. & K. KRAUSE. 1908. Ara ceae— Monst ster- oideae. Pflanzenreich IV. 23B (Heft 37): 4-139. MADISON, M. 1977. A revision of Monstera (Ara- ceae). Contr. Gray Herb. Harv. Univ. 207: 3-100. — Thomas B. Croat, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.; and Michael H. Grayum, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. TWO CUBAN NOVELTIES IN XYRIS Among the X: de kindly loaned by the curators of F, NY, MO, US for a study of North American Sechs were materials of two species of undescribed Yyris, both from the Isle of Pines. From descriptions of the white sand savannas of that island, once so frequently vis- ited by such collectors as N. L. Britton, E. P Killip, and P. Wilson, it is easily seen that it 1s ideal Xyris habitat, having nearly all of the Xyris species endemic to Cuba as well as most of those found both in southern Florida and along the Mosquito Coast and Belize. It is seen that west- ern Cuba is perhaps the most significant floristic connector between the Florida peninsula and esoamerica for not only Xyridaceae but for Eriocaulaceae and many genera of sedges as well. Thus these two novelties are presented as but a small part of what promises to be a considerable updating as the new Flora de Cuba progresses and as Cuban-based botanists continue the work. Xyris curassavica Kral & Urquiola, sp. nov. TYPE: Cuba. Isle of Pines: vicinity of Los Indios, 13 Feb. 1916, N. L. Britton, E. G. Britton & P. Wilson 14219 (holotype, NY; isotypes, CM, F, MO, US). Figure 1 Planta perennis, densicespitosa, laxa, 1.5-4 dm alta, caulibus contractis, radicibus gracilibus fibrosis. Folia bud erecta vel leviter expansa, torta, (5-)10-20(-25) vaginae integrae, laminis 4—5-plo breviores, PE a ë Te a vel roseolae, marginibus in laminas paume m ad apicem pii m (us- lam- inae wei RE anguste dete pallide luteo- virides, 0.5-2 mm latae, ad apicem subulatae, raro paucitrichomatiferae, marginibus integris leviter in- crassatis, raro distante papillosis, paginis glabris, lon- gitudine valde 1—3-nervatis. Vaginae scaporum prox- imale bras glandaceae, folia principalia superantia, pallide 3, fertilibus E bractea Siri bpt ovatae, 4.5-5 mm longae, anguste rotundatae arginem tenues, integri, tum erosae. Sevals teas libera, sub- aequilatera, elliptica, curvata, 4—4.5 ga; [v rinalis et firm 1 apice abrido-papil- losa. Laminae petalorum late obovatae, luteolae, ca .5 mm longae, apice anguste rotundatae, erosae. An therae ee ca. 1.5 m pon vald bifidae et sagittatae, filamentis ca. 0.5 m ngis. ANN. ait Bor. GARD. 74: DEUS 1987. Staminodia bibrachiata, brachiis apice sparsim peni- cillatis. Capsula SOMME ca. 3m onga; placenta ma "n nalis. Sem elc rtocylindra- aie nga, succinea, , trans lucida, apicu- ha dongiudine e valde et irregulariter anastomosocos- Densely cespitose, soft-based perennial 1.5—4 dm high, the stems contracted, the roots slender- fibrous. Leaves mostly erect to slightly spreading, twisted, (5-)10-20(-25) cm long; sheaths '4-!5 as long as blades, entire, a lustrous tan or red- brown, tapering gradually from the clasping base, multicostate and carinate to the blade, there with a scarious, rounded ligule to 2 mm long; blades strongly flattened, narrowly linear, pale yellow- green, 0.5- m wide, apically subulate and sometimes with a apa coarse trichomes, the edges entire and slightly thickened, sometimes re- motely papillate, the surfaces longitudinally strongly 1—3-nerved, often papillate or rugulose- papillate proximally. Scape sheaths lustrous tan proximally, open and short-bladed distally, shorter than principal leaves. Scapes linear-fili- form, flexuous, twisted, distally subterete or oval in cross section, 1—3(—4)-costate, ca. 0.5 mm thick, the costas smooth or distantly scabro-papillose. Spikes broadly ovoid to subglobose, 5-7(-10) mm long, the bracts few, loosely spirally imbricate, lustrous pale red-brown with distinct or indis- tinct ovate pale green (when young) dorsal areas, the backs convex, toward apex low-carinate; ster- ile bracts 2-3, smaller than the fertile bracts and the broad keel firm, a base to acute apex. Petal blades broadly obovate, yellow, ca. 4.5 mm long, the narrowly rounde apex erose. Anthers lance-oblong, ca. 1.5 mm long, deeply bifid and sagittate, on stout fila- ments ca. 0.5 mm long. Staminodia bibra- chiate, the flat, narrow branches sparsely pen- llac, Capsule ellipsoid, ca. 3 mm long, the placentation marginal with placentas extending short-cylindric, 0.5-0.7 mm long, apicula e red-brown, translucent, longitudinally distinctly but irregularly anastomosing-ribbe Frequent in sand savannas, Isle of Pines. Cuba. FIGURE l. Xyris curassavica (Alain & Killip 2194).—a. Habit sketch. — b. Leaf apex. —c. Sector of mid-blade. — d. Leaf blade-sheath junction.—e. Apex of leaf sheath.—f. Leaf base.—g. Spike.—h. Fertile bract.—i. Lateral sepal.—j. Petal blade and stamen.—k. Staminode.—1. Stylar apex.—m. Capsule, one valve removed, showing placentation.—n. Seed. 662 1987] iy yum FIGURE 2. Xyris paleacea (Leon & Victorin 17823).—a. Habit sketch.—b. Lea f. Spike. —g. Fertile bract. mid-blade.—d. Leaf blade-sheath junction. —e. Leaf. — valve with placenta. —j. Seed. es aie specimens examined. CUBA. ISLE OF PINES ios, white sand sabanas, 27 Dec. 1951, Alain & Crus 2194 (US); Los Indios, pir rate 4 Feb. 22 Feb. 1953, Killip 42873 (CAS, F, GH, NY, U, US). NOTES f apex.—c. Sector of leaf at —h. Cen sepal.—i. Capsule This species most resembles X. bicarinata Griseb., another Cuban endemic, in general hab- it and somewhat in leaf but has thinner, paler leaf bases; flatter leaf blades; more slender scapes; and broader spikes with thinner fertile bracts less 664 tending to spread and recurve. The aspect is less robust and smoother. Xyris paleacea Kral & Urquiola, sp. nov. TYPE: Cuba. ixi del Río: Laguna de Sta. Maria, W of Sn. Luis, May 1940, H. Leon & M. Victorin 1 mE (hope US). Figure 2. Planta solitaria vel parum LR annua vel pe- rennis,usque ad 3 dm alta, caulibus contractis, radi- ld gracilibus fibrosis. Folia leviter ie expan- , 4-6 cm longa; vaginae in tegrae, laminis 2-3-plo dug carinatae, lampro-ferrugineae, marginibus ientibus, aut eligulatis; lam- inae compressae, mu ies eae 0.5-1.5 mm latae, ad apicem anguste acutae, leviter incrassatae, marginibus pallidis incrassatis, papillatis vel mein Twana s. Vaginae sca- porum brunneolae, nitidae, klani inio folia prin- cipalia superantes, brevilaminae. Scapi filiformes, ap- icem versus leviter compressi, ca. 0.5 mm lati, distincte bicostales, costis pallidis glabris aut papillosis. Spicae rtilibus breviores; bracteae fer- m longae, apicem versus anguste c piste, valde carinatae. Sepala hteralia libera, ' Subaequilatera, pallide fusca, valde cur- -5 cinea, apiculata, translucida, longitudine valde et ir- regulariter anastomoso-costata Solitary or in small tufts, annual or short-lived perennial to 3 dm high, the stems contracted, the roots filiform-fibrous. Leaves flabellately spread- ing-ascending, 4—6 cm long; sheaths entire, ^-^ as long as the blades, carinate, lustrous red-brown, narrowing gradually from the dilated, clasping base to the blade, there either with a short, scar- ious, rounded ligule or merging with blade base; blades green-brown or maroon, flattened, slightly twisted, 0.5-1.5 mm wide, the apex incurved, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 narrowly acute, slightly thickened, papillate or tuberculate-edged, the margins pale, incrassate, smooth or papillose, the surfaces sm on- gitudinally striate-nerved. Scape sheaths shorter than principal leaves, at base tubular, lustrous brown, multicostate, opening distally, short- bladed. Scapes filiform, distally slightly com- pressed, ca. m wide, distinctly bicostate, the costas pale, smooth or papillate. Spikes lance- ovoid, ca. 1 cm long, acute, of a few, loosely spirally imbricate, red-brown, lacerate bracts with pale green, papillose, subapical, lanceolate dorsal areas; sterile bracts usually 2, oblong, strongly carinate, 3.5-4 mm long; fertile bracts lance- ovate, 5-6 mm long, toward the narrow, sub- cucullate tip strongly carinate, the borders thin, becoming strongly lacerate. Lateral sepals 4.5-5 mm long, free, subequilateral, strongly curved, acute, the thick, narrow keel scabrid from near base to tip. Flowers and stamens not seen. Cap- sule ellipsoid, ca. 3 mm long, the placentation marginal, the placentas extending from base to near tip. Seeds ellipsoid, ca. 0.6 mm long, amber, apiculate, longitudinally strongly but irregularly anastomosing-ribbed. Endemic to pine savanna, Pinar del Río, Cuba. This species, thus far known only from the type specimen, appears allied to X. brevifolia Michaux and X. flabelliformis Chapman and is distin- guished from them by a combination of (often) longer, pale-incrassate-bordered leaves; overall narrower spike outline with narrower, lacerate- bordered fertile bracts; and slightly longer and differently sculptured seeds. — Robert Kral, Department of General Biology, Vanderbilt x 1705, Station B, do Urquiola, Sta. final, Edif. #52, Apto. C-4, Rpto. Hermanos Cruz, Pinar del Río, Cuba. A NEW “VIORNA” CLEMATIS FROM NORTHERN ALABAMA In 1983, while examining specimens from the Huntsville, Alabama, area donated to VDB by the collector, Mr. James D. Morefield, I was par- ticularly interested in a distinctive ““Viorna” that he had collected previously from the lower west- ern slopes of Round Top Mountain (summit el- evation ca. 1,600 feet). This mountain is a dis- tinctive westward lobe of Huntsville Mountain and is south-southwest of Monte Sano Mountain in the same trend, the whole massif a part of the Cumberland Plateau. The Viornae subsection of Clematis is distinguished primarily by the thick- ened, bevelled-edged sepals which in the live flower form (usually) an urceolate design, the flowers themselves being arranged singly or in few-flowered cymes in leaf axils with the primary peduncle short to elongated but always having leafy bracts. The subsection is confined to North America and centers in the southeastern United States; it is notable for its narrow endemics. I visited the locality and, thanks to the accu- racy of Morefield’s label information, was able to locate the population quickly and to find an abundance of plants in flower and early fruit. Further exploration on the same mountain trend resulted in discovery of yet another thriving pop- ulation of this new Clematis. Named in honor of its perceptive first collector, it is described as follows: Clematis morefieldii Kral, sp. nov. TYPE: United States. Alabama: Madison Co., SE Hunts- ville, along upslope side of eastward un- paved extension of Deborah Avenue, 0.65 mi. SSW of Round Top Mtn., limerocky face of slope, clay soil, in Cotinus-Quercus. Vines —5 m, sprawling on boulders or shrubs and forest reproduction; calyx greenish with rose tints, 17 June 1983, R. Kral 70176 (fruiting material from same locality, 27 June 1986, R. Kral with J. R. Carter 73540) (holotype, MO; isotypes, ALU, AUA, BM, CM, DOV, DUR, EKY, F, FLAS, FSU, GA, GH, ILL, ISC, K, LAF, MICH, MISSA, MO, NCU, NY, OS, PAC, RSA, SMU, TENN, TEX, UC, US, VDB, VSC, WAT, WILLI). Figures L. 2 Planta perennis, scandens, usque ad 5 m E cir- rhos efferens. er valde ides subteretes, 2-3 mm crassi, valde cos obrunnei, parce vel jc albo- villosi vel pilosi de araneosi. Foliola principalia re- ANN. Missouni Bor. GARD. 74: 665-669. 1987. ota, expansa, imparipinnata, usque ad 2 dm longa, rhachidi breviter pilosa vel villosa, flexuosa; foliola mucronata. tenuia, integra vel bi- vel-triloba, elon pilosis, 4-15 mm longis; pagina er L^ a fla- vovirens; pagina T sericea aut pilos illares, solitares aut pau in cyma d v expansi mm var. ad — bibracteolis. Se- oy um, acuminatum, compre r ma gine Tedania pons 30:35 mm longo, tmb moso. Perennial scandent vine to 5 m long. Stems flexuous, copiously villous and/or arachnoid with white hairs. Principal leaves imparipinnate, to 2 dm long, spreading, the rachis base shorter than the lowest leaflets, the rachis axis flexuous, pi- losulous or villous; leaflets paired, 9-11, spread- ing or erect, reduced distally on rachis, the upper 1-3 forming tendrils, the lowest broadly to nar- rowly ovate, 5-10 cm long, acute to acuminate, mucronulate, thin, entire to 2- or 3-lobate, on pilose petiolules 4-15 mm long; upper surface smooth, yellow-green; lower surface sericeous or pilose. Flowers axillary, solitary or (more often) 1—3(—5) in sessile cymes, the peduncles at anthe- sis densely white-villous, erect or spreading, 15- 25 mm long, with 2 bracteoles at base. Sepals oblong-lanceolate, 20-25 mm long, erect, the tips acuminate, with narrow white borders, slightly spreading to short-reflexed, the backs pink or pale green-and-red, albosericeous, the edges thick, white-tomentulose, the inner surface smooth, longitudinally inconspicuously few-nerved. Sta- mens linear, 12-20 mm long, the filaments flat- tened, pilose from middle to apex, the anthers including apiculus 3—3.5 mm long, pilose. Fruit omboidal-ovate, 7-9 mm long, acumi- nate, compressed, marginally thickened, seri- ceous, the style 30-35 mm long, with a brown, plumose coma. Additional specimens examined. UNITED STATES. ALABAMA: Madison Co., SE Huntsville, along upslope ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 74 666 vine.—c. Flow- FIGURE |. Clematis morefieldii. —a. Habit sketch, lower node.—b. Sketch of node from mid ering node, largest leaves removed. —d. Sector of mid-stem. (Drawn from Kral 70176.) 1987] NOTES 667 ane Ws M SS FiGuRE 2. Clematis morefieldii. —a. Upper flowering node.—b. Flower.—c. Dorsal (left) and ventral (right) sides of sepal.—d. Three stamens. —e. Anther, enlarged. —f. Carpel.—g. Fruit with long persistent style. (Drawn from Kral 73540.) 668 side of eastward dirt extension of Deborah Avenue, 0.65 mi. SSW of Round T tn., margin of mixed woods on rocky limestone slope, locally common, elev 920 ft., 31 May 1982, J. D. Morefield 629 [JDM (More- field Herbarium), VDB]; vine of loam pockets in ju- niper-Cotinus-mixed A rather dry area, up- per end of Drake Avenue, W face of mountian at E side of Huntsville; calyx pinkish, 17 June 1983, R. Kral 70216 (VDB, and to be distributed) This viorna, so far found only in the limestone uplands around Huntsville, Alabama, is closely related to the variable Clematis viorna L. Dr. Carl S. Keener (1975: 45), an pee on w genus, considers such variants Small, C. flaccida Small, and C. p an Erickson as part of that species, an opinion sup- ported by the biosystematic study of the complex by Dr. W. M. Dennis (1976). However, Keener suggested (loc. cit.), “Nevertheless, critical pop- ulation studies of C. viorna, especially in central Tennessee and adjacent Kentucky, would be in- structive and might reveal more precise taxo- nomically definable topogamodemes." It seems that this species, nested well inside an area of much of the diversity in C. viorna, is indeed distinctive. Of particular interest is the character combination of villous and arachnoid tomentum on the shoot, velvety lower leaflet sur- face, and stouter, usually shorter, peduncles which bear bracts only at the very base (this of partic- ular significance as a character state in the vior- nas). The inflorescence is curious,the lower flow- ering nodes often producing sessile dichasia in the axils, or the bracteoles of these dichasia sup- porting more flower buds in their axils. The up- per nodes are unifoliar and frequently display a wandlike length of progressively reduced leaves with the flowers paired in the axils, while the ultimate and penultimate nodes are often single- flowered; thus the o ct is one ofa narrow raceme with well-spaced nodes. The plants are found in consistent habitat, namely the limestone measures that outcrop be- low the sandstone caprock of the Huntsville Mountain chain ofthe Cumberland Plateau which borders the Highland Rim on the east side of Huntsville. The vines root in a basic clay-loam amongst boulders of massive limestone, often sprawling over the rock itself. The forest type is an open to dense mixture of Juniperus with hard- woods typical of basic substrate, the more dom- inant being Carya carolinae-septentrionalis, C. ovata, Quercus shumardii, Q. muehlenbergii, Q. alba, Q. stellata, Ulmus, Celtis, Acer saccharum, and Fraxinus americana. However, the most no- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 table indicator is the Smoketree, Cotinus obova- tus. The shrub layer is marked particularly by Rhus aromatica, Symphoricarpos, Hypericum Jrondosum, Forestiera ligustrina, and Viburnum rufidulum. Associated and prevalent herbaceous markers include Anemone virginiana, Delphin- ium carolini m, Euphorbia corollata (var.), aptera, Thaspium pinnatifidum, Spigelia ARE Scutellaria ovata, Pycnanthemum incanum (complex), Blephilia hirsuta, Dasysto- ma macrophylla, Hedyotis purpurea, and many composites — particularly Aster, Solidago, and the handsome white-liguled Polymnia canadensis var. ligulata. Most notable here are, however, Silphium brachiatum and Solidago auriculata, these always in close association with Cotinus and the new Clematis. The only other Clematis so far found in the same area is C. virginiana, which is part of a different section. Plants of this entire assemblage grow well on the clays and cherts derived from surrounding limestones and are markedly and abruptly different from the as- sociations of the overlying shales and sandstones of this region. Further field exploration of the same geologic and floristic system is being conducted with an eye toward occurrence of the Cotinus associa- tion, this now known to extend north into Frank- lin and Marion counties in Tennessee and south into Morgan County in Alabama. Where this tree occurs there is likely habitat for Clematis more- fieldii. Mr. J. D. Morefield, a careful and perceptive student of botany, is gratefully acknowledged. During the few years of his adult residence in Huntsville he developed an excellent personal herbarium which adds much to our information about the flora of northern Alabama, his Clem- atis being but one of many interesting discov- eries. Today he is a graduate resident at the Ran- cho Santa Ana Botanic Garden and is involved in revisional and floristic studies centering in the Basin and Range physiography. The Clematis is therefore named in his honor and as a reminder that he is missed back east. LITERATURE CITED PEDE W.M. 1976. A Biosystematic Study of Clem- s, Sect. Viorna, Subsection Viornae. Ph.D. Dis- wnatio on. The University of Tennessee, Knoxville, ssee E R. O. 1943. Taxonomy of Clematis sec- tion Viorna. Ann. Missouri Bot. Gard. 30: 1-60. 1987] NOTES 669 KEENER, C. S. 1975. Studies in the Ranunculaceae of — Robert Kral, Department of Biology, Vander- the southeastern United States III. Clematis L. pilt University, Nashville, Tennessee 37235 Sida 6: 33-47. , : i SMALL, J. K. 1933. Viorna Reichenb. Pp. 526-528 in U Manual of the Southeastern Flora. Chapel Hill ition. TWO NEW SPECIES OF CORDIA (BORAGINACEAE) FROM CENTRAL AMERICA Increased collecting efforts in Central America associated with various floristic projects have provided a wealth of new material. A large num- ber of new species have been reported, and many others, previously poorly known, are now rep- resented by a sufficient number of collections to be understood more adequately. Since 1970, ef- forts in southern Central America have rendered earlier works incomplete in terms of taxa in- cluded and distributional data. Recent studies of Cordia in Mexico and Central America (Miller, 1985), and preparations of treatments for floras of Nicaragua (Miller, in press) and Panama (Mil- ler, in press) have uncovered two new species rom Central America. Cordia liesneri James S. Miller, sp. nov. TYPE: L. Liesner 1976 (holotype, MO 2664908; isotypes, AAU, US). Figure 1. Frutex vel arbor parva 3-6(-8) m alta, ramunculis glabris. Folia persistentia, petiolis 6-1 1(-18) mm lon- gis; laminae anguste elliptico-ovate, (11—)14.5-30(-40) cm longae, (3.8—)6.5-9.5(-13.5) cm latae, glabrae, apice longi-acuminatis, A rotundatis ad obtusis. Inflores- centiae axillaris, (2.5—)6—8.5 cm latis. Flores sessiles, bisexualis; calyx tubulari-campanulatus, 5-6.8 mm longus, 3-lobatus; corolla alba, tubularis, 8.4-11 mm longa, 5-lobata, lobis oblongis, reflexis; stamina 5, filis 6-10 mm longis, puberulis, antheris oblongis, raliter ovoideo, 9-18 mm longo, 9-12 mm lato, ros- trato ad apiceum Shrub or small, slender-trunked tree 3-6(-8) m tall, the twigs glabrous. Leaves persistent, co- riaceous; petioles 6-11(-18) mm long, dl canaliculate adaxially, glabro —9.5(-13.5) cm wide, the apex long-acuminate, the acumen (2.3-)2.8-3.5(-4.6) cm long, the base rounded to obtuse or rarely acute, the margin entire, the adaxial surface gla- brous or rarely minutely strigillose, the abaxial surface glabrous or minutely strigillose. Inflores- cence axillary or sometimes internodal, pendu- lous, cymose, (2.5-)6—8.5 cm broad, the branches shortly brown-canescent. Flowers sessile, bisex- ual, monomorphic; calyx tubular-campanulate, 5-6.8 mm long, 3-4.7 mm wide at the mouth, ANN. Missouni Bor. GARD. 74: 670—673. 1987. lacking ribs, sparsely and minutely brown-strig- illose, 3-lobed, the lobes often somewhat un- equal, deltate to ovate, 0.6-1.1 mm long; corolla white, tubular with reflexed lobes, 8.4211 mm long, 5-merous, the lobes oblong, 2.4—4 mm long, 1.8-2.5 mm wide, the tube 4.4-7.2 mm long; stamens 5, the filaments 7.6-10 mm long, the upper 2.1—3.2 mm free, pubescent on the lower free portion and at the point of insertion, the anthers oblong, 1.3-2 mm long; ovary ovoid to broadly ovoid, 1.3-2 mm long, 1-1.3 mm broad; disc crateriform, 0.6-0.7 mm tall, 1-1.1 mm broad, glabrous; style 5.4-9 mm long, the stylar branches 2-3(-6) mm long, the stigma lobes cla- vate to nearly discoid. Fruits borne in the cup- shaped calyx, red or orange at maturity, drupa- ceous, the stone inequilaterally ovoid, rostrate at the apex, 9-18 mm long, 9-12 mm broad, the surface with low ridges, the endocarp bony. Distribution. Cordia liesneri is known only from the Golfo Dulce region of Puntarenas, Cos- ta Rica, where it occurs in wet forests below 200 m in elevation. Among the Central American members of sect. Myxa (Endl.) DC., Cordia liesneri is distinctive in its oblong-ovate leaves longer than those of related species, axillary inflorescences, and bright red fruits containing large and prominently ros- trate stones. This species is perhaps most closely related to Cordia lucidula I. M. Johnston with which it shares a similar habit of growth, rela- tively large, glabrous leaves, a 3-lobed calyx, and red drupacous fruits. However, it differs from C. lucidula by bui axillary inflorescences and rostrate endoc Collections of C oua liesneri have existed since Skutch and Allen collected in the Golfo Dulce region in 1947 and 1951 respectively. However, they have been identified incorrectly as Cordia protracta I. M. Johnston, which differs by its terminal inflorescences and pentamerous calyx, and as Cordia eriostigma Pittier, which differs in having terminal inflorescences and a campan- ulate corolla. Allen (1956) briefly described C. liesneri under the name C. protracta and stated that flowering occurs in December and that fruits mature in January Additional specimens examined. |. CosrA RICA. PUNTARENAS: region between Esquinas and Pas Sur 1987] NOTES 671 FicurE 1. Cordia liesneri.—A. Fruiting branch, after Liesner 1976 (MO).—B. Open corolla, after Burger & Liesner 7224 (NY). de Osa, elev. 30 m, Allen 5772 [DS, F (2), GH, US]; ner 7224 (CR, F, NY); slopes adjacent to airport, Rin- region between Equinas and Palmar Sur de Osa, elev. cón de Osa, Liesner 1858 (AAU, CR, MO, US); Golfo 75 m, Allen 5827 (DS, P); about 5 km W of Rincón Dulce and Rio Terraba, elev. 30 m, Skutch 5303 (F, de Osa, Osa Peninsula, elev. 50-200 m, Burger & Lies- | MICH). FIGURE 2. Cordia cardenasiana. Pip intact striate calyx. All after Contreras 6835 (holotype). Cordia cardenasiana James S. Miller, sp. nov. TYPE: Guatemala. Petén: Cardenas, on rocky hill, 24 Mar. 1967, Elias Contreras 6835 (holotype, LL 279752; isotypes, DS, F, LL, US). Figure 2. Arbor ad 15 m, ramunculis glabris. Folia deciduis, laminis ellipticis, 4.4—1 1.4 cm longis, 2.4—4.5 cm latis glabris. Inflorescentia cymo-paniculatis. Flores hetero- ` ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 —A. Flowering branch.—B. Open flower.—C. Apex of a flowering branch styli; calyx tubularis, 5—6 mm longus; corolla tubularis 4 ; m mm longum; discus annularis. Fructus non visi. Tree 15 m tall, the twigs glabrous. Leaves de- ciduous; petioles 8-30 mm long, Pep en ipe glabrous; blades elliptic, 4.4-11.4 c long, 2.4-4.5 cm wide, the apex acute to acu- 1987] minate, the base acute, the margin entire, the adaxial surface glabrous but evenly papillose, the abaxial surface glabrous. Flowers on short ped- icels to 1.5 mm long, distylous, borne with the leaves; calyx tubular, 5-6 mm long, 3-3.8 mm wide at the mouth, unequally lobed and tearing upon dehiscence or dehiscing circumscissily, striate or faintly costate, inconspicuously short- puberulent; corolla white, tubular with some- what spreading lobes, 14.2-16.8 mm long, 5-merous, the lobes oblong, 5.2-6.2 mm long, 4.8-6 mm wide, the tube 4.7-5 mm long; sta- mens 5, the filaments 9.8-11.5 mm long, t upper 7-7.6 mm free, glabrous, the anthers ob- long, 2.3-3 mm long; ovary obloid, 1.2-1.4 mm long, 1.3-2.5 mm broad; disc annular, 1.4 mm tall, 1.3 mm broad, glabrous; style 6.8-7 mm long, the stylar branches 1.4-1.6 mm long, the stigma lobes clavate. Fruits unknown. Distribution. Cordia cardenasiana is known only from the type collection made in Petén, Guatemala. Cordia cardenasiana is a small tree that must be quite attractive in flower. Like related species, it presumably produces all of its flowers in a short period during the dry sesaon. It is assigned pro- visionally to sect. Gerascanthus (P. Br.) G. Don, but a definite sectional placement cannot be made until fruits are known. Members of sect. Ger- ascanthus have ellipsoid fruits with a fibrous wall, and the base of the style remains attached. Mem- bers of sect. Rhabdocalyx A. DC. have fruits with a bony endocarp; some species are otherwise in- distinguishable from sect. Gerascanthus. Al- though Cordia is diverse in pollen morphology, the two sections share pollen grains of the same type (Nowicke & Ridgway, 1973), and this char- acter cannot zagi the sectional placement of C. cardenasia Cordia ae en is probably most closely related to C. gerascanthus L., with which it shares glabrous leaves and a terminal, cymose-panic- ulate inflorescence. Although most populations of C. gerascanthus have pubescent staminal fil- aments, populations with glabrous filaments are known from southeastern Mexico (Johnston, 1950; Miller, 1985); filaments of C. cardenasiana NOTES 673 are glabrous. Among the Mexican and Central American species of Cordia, C. cardenasiana is distinctive in having parallel-sided corolla lobes, a character that it shares only with C. alliodora (Ruiz & Pavón) Oken, C. gerascanthus, and C globulifera 1. M. Johnston. A key is provided below, and a key to the remaining species of the section from this region is in Miller (1986). la. n with ant domatia; indument of stellate hai C. alliodora Ib. Pant lacking ant domatia; indument of sim- ple hairs. 2a. Inflorescence cymose-paniculate. Calyx striate or faintly costate, gla- brous; corolla less than 18 mm long RSV RAYS OAE en rune C. tals Mita 3b. Calyx costate, pubescent; corolla mo th m lon C. Md 2b. Inflorescence pat AER nearly umbel- Vet x C. globulifera I thank the curators of the following herbaria for lending collections: AAU, CR, DS, F, GH, LL, MICH, MO, US. I also thank my wife, Leslie Miller, for support and the illustration of Cordia liesneri, John Myers for the illustration of Cordia cardenasiana, and G. Schatz for helpful com- ments on the manuscript. LITERATURE CITED ALLEN, P. H. 1956. The Rain Forests of Golfo Dulce. Univ. Florida Press, Gainesville, Florida JoHNsTON, I. M. 1950. Studies in the Boraginaceae XIX. Cordia section Gerascanthus in Mexico an . Arnold Arbor. 31: 179-187. MILLER, J. S. _ Systematics of the Genus Cordia (Boraginaceae) in Mexico and Central America. h.D. Dissertation. St. “Loui is Univ., St. Louis, Missouri 1986. Cordia macvaughii, a new species of Boraginaceae from western Mexico. Syst. Bot. 11: 179-187 Central America — T Boraginaceae. In W. D. Stevens (editor), Flora de ein dr (in press). A revised treatment of Boraginaceae for Pan- . Ann. - Miles uri Bot. Gard. (in press). NowICKE, J. W. & J. E. RipGwAv. 1973. Pollen stud- ies in the ur Cordia (Boraginaceae). Amer. J. Bot. 60: 584-5 —James S. Miller, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. SARACHA SPINOSA—A NEW COMBINATION IN PERUVIAN SOLANACEAE The name Saracha R. & P. was long misap- plied to the group of herbaceous neotropical plants now correctly known as Jaltomata Schldl. (Gentry, 1973). With reestablishment of the name Jaltomata for the herbaceous plants, the need for reinstatement of Saracha to the group of shrubs of upland South America that had long been er- roneously known as Poecilochroma Miers be- came evident. The redefinition of Ja/tomata was followed by D'Arcy (1973: 626, 1986: 28), Gen- try & Standley (1974: 42), Hunziker (1979: 53), and Nee (1986: 76) and the reinstatement of Saracha by Hunziker (1979: 56). Both genera are in need of revision, and not all combinations have been made to recognize the restored generic names. However, determination of a collection made in the course of preparing a flora for Huas- carán National Park in the uplands of the Cor- dillera Blanca in northcentral Peru (Smith, in prep.) requires the following new combination: Saracha spinosa (Dammer) D'Arcy & D. N. Smith, comb. nov. Poecilochroma spinosa Dammer, Bot. Jahrb. Syst. 37: 637. 1906. TYPE: Peru. Weberbauer 2907 (B, not seen, destroyed; photo, MO). Copiously armed shrub with long, trailing or weeping branches; twigs glabrate, sometimes with a simple Pp, paia ed many short d lea A Bray- ish, acicular spines 3-25; mm long. Leaves entire; elliptical, obtuse or rounded at each end, mostly 12-20 mm long, coriaceous, glabrous or with minute simple trichomes, shiny above, the mar- gins subrevolute; petioles 2-5 mm long. Inflo- rescences consisting of solitary terminal or ax- illary flowers; peduncles wanting; pedicels slender but broadening upwards, 10-15 mm long. Flow- ers pendant, the calyx cupular, ribbed, dentate, sometimes irregular, 6-8 mm long, sometimes splitting; corolla apically plicate in bud, narrowly campanulate, ca. 20-25 mm long, ca. 15 mm wide at the mouth, the apex truncate with 5 del- toid teeth 2 mm long; yellow and evenly mi- nutely puberulent outside, inside basally uni- formly dark violet, this reduced upwards to a field of lobes and spots and pure yellow at the apex; filaments ca. 15 mm long, inserted at the base of corolla tube, glabrous and unappendaged, ANN. Missouri Bor. GARD. 74: 674—675. 1987. the anthers oblong, apiculate, basally cordate, 4— 5 mm long, yellow, included; ovary obconical, 4 mm long, the basal third enveloped in nectary, glabrous, the style glabrous, exserted beyond the anthers and nearly equalling the corolla. Fruit (not preserved) a leathery berry about the size of a gooseberry, perhaps 7 mm across. Saracha spinosa is a weak, clambering shrub that assumes a down-curving, weeping habit when it lacks support. The branches can reach several meters in length. The corollas are not so broadly campanulate as in other members of the genus. The plicate folds in the corolla in bud were not found by Hunziker (1979: 58) i in the species he discussed. Saracha distinct from other members of the genus in n having formidable slender spines which appar erive from short leafy shoots in the manner of Dunalia spinosa, Iochroma horrida, and some other related So- lanaceae. The spines on the specimen at hand are copious, ascending, 8-25 mm long, and needlelike. Fruit was not available for dissection to ascertain the details referred to by Hunziker (1979: 58) in his notes on three other species of Saracha. In his discussion under Poecilochroma in the Flora of Peru, Macbride (1962: 58) suggested that this species is similar to P. lindeniana Miers and to P. lobbiana Miers, but both of these are un- armed and have more open campanulate corol- las, as is to be seen in the type illustration and photos examined of these two species The collection made in this study bears a re- markable similarity to a specimen of Dunalia collected by López & Sagástegui which we ten- tatively refer to D. spinosa (Meyen) Dammer. The leaves of the López & Sagástegui collection are small and elliptical like those of our collection of Saracha and unlike the larger oblanceolate leaves of other specimens of D. spinosa, includ- ing the type collection (Meyer, BM, destroyed, photo MO). The flowers, too, are superficially similar but narrower, and of course the filaments have the basal appendages characteristic of the genus Dunalia (Hunziker, 1959-60: 212). Had we not examined the interior of the flowers, we would have thought these the same species, not different genera. The great similarity between Saracha spinosa 1987] and the neighboring Dunalia within adjacent parts of Peru argues a need for unusual care in deter- mining specimens of these plants and caution in accepting past determinations. Under his treat- ment of Saracha (Poecilochroma) spinosa Mac- bride cited the following four specimens, which appear to be at least geographically consistent: Dept. La Libertad: Prov. Bolivar, Ferreyra 1254; Prov. Santiago de Chuco, toward Angasmarca, West 8162. Dept. Ancash: Prov. Huari, 3,600 m, Weberbauer 7014. Dept. Ayacucho: Prov. Huamanga, above Quinua, Weberbauer 5542. Saracha spinosa is apparently restricted to in- terandean valleys of the western and central chains of the Peruvian Andes. It has an extended range of about 700 km, within which, at least according to Weberbauer (1945: 420), the species is sometimes locally common. Collection sites known to us show that it occurs in shrubland ranging from about 78°05’ W to 74*08'W and from about 7?22'S to 8?05'S. The species has an ap- parent elevation range from 3,300 to 3,700 m. The type locality, which was imprecise, is near or within Huascarán National Park. The Park is in the Ancash Department of cen- tral Peru, 300 km (air distance) north of Lima. The reserve occupies nearly all the Cordillera Blanca, which is the world's highest tropical mountain range. The cordillera is located be- tween 8?50'S and 10?00'S latitude and between 77°05'W and 77?49"W longitude with a north- south length of 158 km and an area of 340,000 ha (131 sq. mi.). The elevation range is from 3,240 m to the 6,770 m summit of Nevado Huas- carán Sur, the bulk of the Park above 3,500 m. The outcrop is a mixture of igneous and sedi- mentary rocks. The cordillera was extensively glaciated and still has many glaciers and icefields. Its topography is complex and supports a mosaic of vegetation types. The most diverse and den- sest shrub communities are found in the valleys reaching lower elevations (3,500—3,800 m), where the microclimate is warmer and moister. Al- though valleys with these conditions are found on both sides of the cordillera, the greatest num- ber are on the eastern side. In over a year of field collecting throughout NOTES 675 the entire park, Saracha spinosa was located only once, in Quebrada Rurichinchay near the valley bottom in a community dominated by Miconia salicifolia and with Alnus acuminata, Myrica pu- bescens, Vallea stipularis, and Weinmannia aff. laxiflora SPECIMENS EXAMINED Saracha spinosa. PERU. DEPT. ANCASH: Prov. Huari, Huascarán National Park, Quebrada Rurichinchay be- tween boundary and Quebrada Pachachaca. 3,600- 3,700 m, D. N. Smith 1 2475 (CPUN, HUT, MO, USM, dupla). DEPT. AYACUCHO: Prov. Huamanga, road from La Quinua to Abra Apacheta de Tambo, 12,000 ft., Plowman & Davis 4651 Dunalia aff. spinosa. PERU. DEPT. LA LIBERTAD: Prov. Bolivar, Laguna de Los Ichus, al pié de rocas, 3,600 m, López & Sagástequi 3241 (MO). This study was supported by National Science Foundation Grant BSR-8305425 LITERATURE CITED D’Arcy, W. G. 1973 [1974]. Solanaceae. In Flora of Panama. Ann. Missouri Bot. Gard. 60: 573- 780 1986. The genera of Solanaceae and their t ypes. Solanaceae Newsletter 2(4): 10-3 GENTRY, J. L Restoration of the genus Jal- tomata (Solanaceae), Phytologia 27: 286-288. TANDLEY. 1974. Solanaceae. Jn Flora ofG Field Mus. Bot. 24(10-1&2): 1-151. HUNZIKER, A. J. 1959-60 [1960]. Studios Lai So- lanaceae. II. Sinopsis taxonómica del género Du- nalia H.B.K. Bol. Acad. Nac. Ci. Córdoba (Ar- gentina) 41: 211-244 — oo. American Solanaceae: a syn- optic s . Pp. 49-85 in J. G. Hawkes et al. (editors) The Biology and ERU of the So- cademic Press, Lon Mich n 1962. Solanaceae. Jn Flora of Peru. eld Mus. Bot. 13(5B-1): 3-269. NEE, M. 1986. Solanaceae I. Jn Flora de Veracruz, Fasc. 49. Inst. Nac. Invest. Kcu Bióticas, Xa- lapa, Veracruz, Mexico. WEBERBAUER, A. 1945. El digo PME de los An- des Peruanos. Min. Agric., — William G. D'Arcy, Missouri Botanical Gar- den, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.; and David N. Smith, Missouri Botanical Garden and Department of Botany, Iowa State University, Ames, Iowa 50011, U.S.A. YUTAJEA, ANOTHER NEW GENUS OF RUBIACEAE FROM THE GUAYANA HIGHLAND Of the genera of Rubiaceae endemic to the region of the Guayana Highland, none had pre- viously been described from the Serrania de Yu- taje in the Territorio Federal Amazonas of Ven- ezuela. A recent expedition to this region collected material of a rubiaceous tree, here described as a new genus. It is fitting to name the genus for Yutaje, a sandstone mountainous area where nu- merous endemic species are already known. Yutajea Steyermark, gen. nov. TYPE: Y. /iesneri Steyermark. Tribe Isertieae. Arbor. Stipulae interpetiolares persistentes late lan- Inflorescentia thyrsiformi-paniculata alyx eases tandem in k S 5111 Adv sape DEN ° E ovulis numerosis. Yutajea liesneri Steyermark, sp. nov. TYPE: Ven- ezuela. Territorio Federal Amazonas: Dep- to. Atures, 5-8 km NW of Yutaje settle- ment, along stream flowing south from east side of unnamed peak, 3 km west of Río Coro Coro, west of Serranía de Yutaje, 05?40'N, 66°9’'W, 700-1,000 m, 10 Mar. 1987, Ronald Liesner & Bruce Holst 21826 (holotype, MO; isotype, VEN). _ Arbor € 6- metralis li ib d r 4 mm latis; foliis oblanceolatis vel pera ipii apice acumina rra iie cutis 13.5-20 cm longis 5.5-8.5 cm latis e. thyrsiformi- sein late ovoidea mul crag saratapas , axibus p 2-16 modice vel dense hirtellis in verticillos dinatos ema in dichasia composita desinsatibus. ca- lyce hypanthioque 4-6 mm longo subadpresso-pubes- centi; calyce ante anthesin truncato vel paullo undulato sub anthesi in lobos 3-4 irregulariter rumpenti, lobis suborbicularibus vel suborbiculari-ovatis apice rotun- datis, marginibus minute ciliolatis; pec rosea late cylindrica vel subinfundibuliformi 8-9 mm longa 4-7 mm lata, tubo extus glabro intus seis partem ba- silarem 1.5 mm glabrum adpresso-pubescenti atque prope orificium barbato pilis longioribus instructo; lobis 6 paullo inaequalibus suborbiculari-ovatis vel obova- to-oblongis apice obtusis vel rotundatis ubique glabris marginibus minute papillatis; staminibus 6, antheris linearibus ANN. MISSGURI Bor. GARD. 74: 676—678. 1987. Tree 6 m tall, the twigs appressed-pubescent distally. Stipules broadly lanceolate, acute, 8 mm long, 4 mm wide, strigose. vid oblanceolate to lance-elliptic pex, acute at base, 13.5-20 cm long, 5.5-8.5 cm wide, the up surface glabrous, the lower surface pilosulous on midrib and lateral nerves, sparsely pubescent on some tertiary veins, glabrous on surface between the tertiary veins; lateral nerves 12-17 each side, anastomosing near margin, elevated below; mid- rib sulcate above and elevated below; tertiary veinlets inconspicuous above, finely reticulate and impressed below. Inflorescence thyrsiform-pan- iculate, broadly ovoid, 35-50-flowered, 6-11 cm long excluding the peduncle, 4.5-7 cm wide, the main axes 12-16 in 3-5 verticils ending in often compound dichasia, moderately to densely hir- tellous; lowest axes 12-25 mm long, the others 7-15 mm long. Bracts subtending axes ovate, acute, 2.5-4 mm long, m wide, minutely hirtellous. Peduncle terminal, 8-12 cm long, 2.5- 3 mm diam., moderately pubescent. Calyx and hypanthium 4-6 mm long, subappressed-pubes- cent; hypanthium obconic, 3-5 x 3-5 mm. Calyx truncate or slightly undulate before anthesis, splitting irregularly during anthesis into 3-4 thickened, suborbicular or suborbicular-ovate obes, these rounded at apex, 3-3.5 mm long, 4- 8 mm wide, minutely ciliolate at the margins. Corolla pink or roseate, fleshy-thickened, broad- ly cylindric or subinfundibuliform, 8-9 mm long, 4—7 mm wide, the tube 5 mm long, 4-7 mm wide, glabrous without, within densely appressed-pu- bescent except for the glabrous basal 1.5 mm, and with longer barbate pubescence at the orifice; lobes 6, slightly unequal, suborbicular-ovate or obovate-oblong, obtuse or rounded at apex, 4-5 mm long, 2.5-3 mm wide, glabrous both sides, thers linear, 3.5-5 mm long, 0.8-1. the thecae transversely rugulose, terminating in an oblong, rounded connective 0.5-1 mm long, 0.5 mm wide; filaments ligulate-linear, thick- ened, 1.5 mm long, 0.7 mm wide, inserted 1.5- 2.5 mm above the base ofthe corolla tube, broad- er than the base of the thecae. Style 4-6 mm long, papillate-verrucose; stigmas 4—6, ligulate, 0.5 mm long. Disk annular, the margin undulate, 1 mm long, 4 mm diam. Ovary 6-celled, ovules ca. 8 in each cell. 1987] NOTES 677 LECT FiGURE 1. Yutajea liesneri.—A. Habit.—B. Portion of inflorescence.—C. Calyx and hypanthium before anthesis. — D. Calyx and hypanthium during anthesis. — E. Stamen (lateral view). — F. Stamen (ventral view). — G. Corolla, interior view. — H. Transverse section through ovary, semidiagrammatic. —I. Corolla, subinfundi- buliform type.—J. Disk, from above, in depression at base of calyx tube. — K. Floral bract, exterior view.— Detail of lower leaf surface. — M. Style and stigmas. — N. Flower with broadly cylindrical corolla type with slightly unequal lobes. Yutajea is a member of the tribe Isertieae as stamens inserted near the base of the corolla tube circumscribed by Kirkbride (1979) and may be and in having sub-basifixed anthers. The slightly placed next to its closest related genus, /sertia. ^ unequal corolla lobes and the uniformly pubes- From I/sertia it differs especially in having the cent interior of the corolla tube are additional 678 characers at variance with /sertia. The corolla of Yutajea is short-cylindric or subinfundibuliform with imbricate lobes. Boom (1984) described the corolla tube in /sertia as “cylindrical, short or elongate" and the lobes as **valvate or imbricate in bud.” In her abstract of characters delimiting the tribe Isertieae, Kirkbride (1979) character- ized the aestivation of the corolla as **valvate" but later (p. 315) gave the aestivation as “‘valvate or valvate-imbricate." As Boom indicated, Zs- ertia may have either valvate or imbricate aes- tivation. In /sertia the stamens are inserted near the mouth of the corolla tube, the anthers are dorsifixed, and the orifice of the corolla is usually villous or barbate, except in J. scorpioides Boom, while the remainder of the interior of the corolla tube is glabrous in all the species with the ex- ception of I. longifolia (Hoffsg. ex Roemer & Schultes) Schumann. Yutajea adds another endemic genus of the ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Rubiaceae to the list of twelve previously de- scribed from the Guayana Highland (Steyer- mark, 6). The collections cited in this paper were made under Grant No. 3264-86 of The National Geo- graphic Society. LITERATURE CITED Boom, B 84. A revision of /sertia (Isertieae: Ru- biaceae). Brittonia 36: 425-454. KIRKBRIDE, M. CRISTINA GARCÍA. 1979. Review of } š 1T : Danhi \ B : : rittonia 31: STEYERMARK, J. 1986. Holstianthus, a new genus of Rubiaceae from the Guayana Highland. Ann. Mis- souri Bot. Gard. 73: 495-497. —Julian A. Steyermark, Missouri Botanical Gar- den, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. BOOK REVIEW Duke, James A. 1986. Isthmian ee ical Dictionary, Third edition. Pawan mar Scientific Publishers, Jodhpur odis ISBN 81-85046-35-2. Price $60 The Isthmian Ethnobotanical Dictionary, con- cerned with useful plants of Panama, is 205 pages in length, each page measuring approximately 81^ x 11 inches. It is in its third edition, the first appearing in 1972, the second in 1978. The latest contains 325 black-and-white line drawings of flowering plants and a few ferns The author explains that the Dictionary “° contains herbal folklore in tropical plants which complements the voluminous folklore on tem- perate plants." In his introduction he indexes “some frequent ailments and the more common herbs used to treat them." Actually, many of the plants listed are trees and shrubs. The diseases range from abscesses to yellow fever. For ex- ample, under *'indigestion" he lists by common betical and composite fashion, he lists the com- mon names ofthe plants without discussing them; secondly, the diseases themselves (in full capi- tals), including a brief layman's definition of the isease and the genera of plants used therapeu tically; lastly, the binomial name (in full Mida and a discussion of how the plants are used by z natives. While emphasis is placed on medic- inal plants, the discussion may include such uses of the plants as food, in construction, in the fash- ioning of fishhooks, etc. Occasionally, the treat- ments extend extra Panama, e.g., under bamboo there is an elaborate discussion of about 1,500 words, a goodly segment of Hsing bou to the uses of the grass in India and Ben y the use of abbreviations in uma names are used in 5 Panama, “Darien Spanish (D)"; “English yao etc. In the eyes of a botanist, apart from the wealth of ethnobotanical data, the three most striking features of the Dictionary are: 1) the complete- ness of the list of ethnobotanical plants; 2) the accuracy in the spelling of binomial names and the authorities; and 3) how easily one familiar ANN. MissouRni Bor. GARD. 74: 679-680. 1987. with the vegetation of Panama can recognize the individual species from the line drawings. here are several items which I find disturbing, the prime being a lack of discussion in the In- troduction concerning the various ethnic groups in Panama, plus the lack of a map of their lo- cations in the Republic. The author sometimes leaves the impression that the Kuna Indians are the only aboriginals in Panama worth consid- ering ethnobotanically. Why not more consid- eration of the Chocó Indians, granted that they are not as sedentary in Panama as are the Kunas? After all, Duke tells us (p. 26): “I am called ‘Bo- rojo' among the Darién Chocó because, for a while I was distributing ‘borojo’ seedlings like Johnny Appleseed." The Guyami Indians of western Panama are hardly given a nod. The obvious answer is that Duke spent a considerable portion of his early scientific life among the Ku- as. From the viewpoint of a print job, the book leaves something to be desired, considering that the price of the book is $60.00 (U.S.). The paper is of questionable quality, occasionally the ink is smudged; all too often one encounters a blank space in a sentence, a word or words having been deleted; occasionally the alignment of words in a column or sentence is poor. An important point: books whose titles con- tain the word “dictionary” are usually segregated on library shelves under “for reference only." Such a restriction may limit the uses of the book and have an impact on its popularity. I suggest that in the next edition a small su- perscript number be appended after each bino- mial to indicate the plant family; this could be checked against a list of numbered families in an appendix. In the introduction, Duke in a cursory, but = its vegetation. His concern for the future disap- pearance of the magnificent jungles of Darién strikes a familiar note; his defense of folklore medicine is convincing: . I could spend a month with the herbals and dic medical texts and come up with hundreds of examples of folk medicine that have been vindicated or justified by subsequent scientific research." As far as Pan- ama's herb lore is concerned, he has already done 680 the basic spadework in admirable fashion. One must not forget that James Duke collected more than 6,000 “numbers” (herbarium specimens, each number usually in duplicate or triplicate) in the Republic of Panama (cf. Dwyer, in the ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Botany and Natural History of Panama, Mis- souri Botanical Garden, 1985), a memorial to his assiduity and a base for his ethnobotany.— John . Dwyer, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A ERRATUM The incorrect author citation was used in the intro- duction to the paper titled “Notes on the Breeding Systems of Sacoila lanceolata (Aublet) Garay (Orchi- daceae),” by Paul M. Catling [74(1): 58-68. 1987]. In- stead of “var. paludicola Luer” it should be “var. pal- udicola (Luer) eroi Wunderlin, et Hansen, Phytologia 56: 308. Volume 74, No. 2, pp. 183-462 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN was published on 29 September 1987. THE GENERA OF EUPATORIEAE (ASTERACEAE) R. M. King & H. Robinson Smithsonian Institution Starting in 1966 the authors have provided a series of partial revisions of the Eupatorieae, publishing over 200 papers on the group. The Genera provides a synthesis of their contributions to our understanding of this species-rich group: one in every 150 species of flowering plants belongs to the Eupatorieae. This new work treats all 180 genera of the tribe with a description, full-page illustration, discussion, and listing of recognized species and their distributions. Keys to all the subtribes and genera and regional keys for portions of tropical America are included. A nomenclator of the over 6,000 specific and infraspecific names that have been placed in the tribe at one time or another indicates their present disposition. Published as Monographs in Systematic Botany from the Missouri Botanical Garden, volume 22, 581 pages, 81⁄2 by 11 inches, color frontispiece, hardbound. October 1987. Price $70.00 For U.S. shipments: add $1.50 for one copy and $.50 for each additional copy. Orders should be pre-paid; a $1.00 fee will be added to orders requiring invoices. No shipments are made until payment is received. Mail order with your check or money order, payable to Missouri Botanical Garden to: Department Eleven Missouri Botanical Garden P.O. Box 299 St. Louis, MO 63166-0299, U.S.A. Please send copy(ies) of The Genera of Eupatorieae to: O Payment enclosed. Name : : — ———— TEL Send invoice Address | ec Sg ($1.00 will be added to total) iT 743) CONTENTS A Revision of Panicum Subgenus Poulin Section Rudgeana (Poaceae: Paniceae) nando O. Zuloaga Arundoclaytonia, a New Genus of the Steyermarkochloeae (Poaceae: Arundinoideae) from Brazil Gerrit Davidse & R. P. Ellis Siphocampylus oscitans (Campanulaceae: Lobelioideae), a New Name for Burmeistera weberbaueri from Peru Bruce A. Stein Synopsis of the Genus antera (Campanulaceae: nd in Peru Stein New Species of Possiflora Subgenus sovr from Ecuador J. E. Lawesson E The Genus Attalea C Palenie) i in Panama X4 : Ç Novelties In i Fer- Bruce A. L. B. Holm-Nielsen & eror C. de Nevers LY HE (Loranthaceae and Viscaceae) Job Kuip E A Revision of Pisi (Sapindaceae) O. Téllez V. & B. G. Schubert .... z : Systematics of the Southern African Genus cea (Iridaceae Iridoideae) Goldblatt . dd. Peter Goldblatt wars od ç (Iridaceae) i in the Southwestern Cape, South Africa ae x A. H. Gentry & J. Steyermark -n -— | cte Una Nueva Especie del Genero Dioscorea (Dioscoreaceae) del Estado de Queretaro, Mexico - Peter ` es on the ME id Tenn; of Watsonie Giboni (IV. pyramidata, W. -ardernei) k , Cytology, and. Embryology of Campynemanthe (Liliales: ae AL "Porter P. pe I, r Goldblatt * Hiroshi [PC icd ; a Annals of the . Missouri — Botanical US Volume 74, Number 4 Winter 1987 Annals of the Missouri Botanical Garden The Annals, published quarterly, contains papers, primarily in systematic botany, con- tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the Garden will also be accepted. Authors should write the Editor for information concerning arrangements for publishing in the ANNALS. Instructions to Authors are printed on the inside back cover of the last issue of each volume. Editorial Committee George K. Rogers : Editor, Missouri Botanical Garden Janice Wilson Editorial Assistant, Missouri Botanical Garden Marshall R. Crosby Missouri Botanical Garden Gerrit Davidse Missouri Botanical Garden John D. Dwyer Missouri Botanical Garden & Saint Louis University Peter Goldblatt Missouri Botanical Garden Henk van der Werff Missouri Botanical Garden For subscription information contact Departmen t Eleven, P.O. Box 299, St. Louis, MO 63 166. Sub- scription price is $75 per volume U.S., $80 Canada x and Mexico, $90 all other countries. Airmail deliv- ~ ery charge, $35 per volume. Four issues per vol- Eleven, P.O. Box 299, St. Louis, MO 63166. A be ANNALS OF THE Missouri BOTANICAL pape (ISSN 0026-6493) is published quarterly by the - Missouri Botanical Garden, 2345 Tower Grove Av- . enue, St. Louis, MO 63110. Second class postage — id at St. Louis, MO and additional mailing offices. 2 POSTMASTER: Send address changes to Department — Volume 74 Number 4 1987 Annals of the Missouri Dotanical Garden NZ CONTRIBUTIONS TO A SYMPOSIUM ON THE EVOLUTION OF THE MODERN FLORA OF THE NORTHERN ROCKY MOUNTAINS: INTRODUCTORY REMARKS The following nine papers represent contri- butions from a symposium presented on June 11, 1985, at the University of Montana. The ymposium was sponsored jointly by the Paleo- botanical Section and the Pacific Section of the Botanical Society of America and was held dur- ing the annual meetings of the Pacific Division of the American Association for the Advance- ment of Science. The main purpose of the symposium was to bring together current information about the life and earth sciences, and they have constructed their written reports accordingly. A frequent problem for nonpaleobotanists in- terested in learning about the past vegetation in a given region is that it is difficult to locate this information. Reports about past vegetation are by no means rare, but they are scattered, and we have lacked good syntheses. The following pa- pers attempt to solve that problem for the north- ern Rocky Mountain region. Each is a succinct yet authoritative summary of a particular body of information authored by leading specialists in ANN. MISSOURI Bor. GARD. 74: 681-682. 1987. that field. A number ofauthors have gone beyond presenting reviews and have included informa- tion and ideas in their treatments that have not been published elsewhere. Thus, each report is intended to be a definitive work on a particular duration of geologic time. Readers will note what may appear to be two major inconsistencies from paper to paper. One has to do with geologic time and the application of epoch names, while the other is the use of the terms “flora” and “vegetation.” Paleobotanists generally agree on the absolute ages of the various plant-bearing strata, but they may disagree on the absolute age of the boundary between epochs. Thus, the same deposit may be regarded as latest Eocene by one worker and ear- liest Oligocene by another. Where this might cause confusion in this series of papers, authors have been asked to provide the absolute age as well as the epoch. Workers dealing with modern plants and with those from the not too distant past use the term "flora" to indicate a list of taxa present and “vegetation” to include all aspects of those plants. Historically, fossil “floras” were little more than lists of species identified from a deposit. Later, such lists expanded to include taxonomic de- scriptions, comparisons, and, more recently, in- 682 formation about frequency of occurrence of re- ins and environment of deposition. Interpretations of community structure, mean annual temperature, mean annual range of tem- perature, seasonality, and amount and seasonal distribution of rainfall are often included in mod- ern treatments as well. ile these are clearly aspects of “vegetation,” the term “flora” has per- sisted and is often used instead. Thus, those working with ancient plants may not iudi between these two terms and may use "flora when “vegetation” is appropriate. A second purpose of the symposium was to honor four workers who pioneered the study of the past vegetation of the northern Rocky Moun- tain region. These are: Herman F. Becker, Ro- land W. Brown, Edu Dorf, and Harry D. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 MacGinitie. Their contributions have been sum- marized by Jack Wolfe, and the summary is in- cluded as part of this publication. Drs. Becker and MacGinitie passed away after the sympo- sium was held and their bibliographies are in- cluded here. The accomplishments of these sci- entists provided a solid foundation on which modern work rests. Even more importantly, they provided leadership and encouragement to the current generation of paleobotanists, and the pa- pers published as a part of this symposium owe much to their efforts. — Charles N. Miller, Jr., Department of Botany, University of Montana, Missoula, Montana 59812, U.S.A. DEDICATION This symposium is dedicated to four col- leagues, all unfortunately deceased, who have contributed significantly to the paleobotany of the northern Rocky Mountains: Herman F. Becker, Roland W. Brown, Erling Dorf, and Har- ry D. MacGinitie. The work of all four men has provided a foundation of collections and knowl- edge on which this symposium is partially based. Two ofthese valued colleagues were, in fact, alive at the time (June 1985) the symposium was held. We had intended to honor Herman Becker dur- ing the course of the meetings at which the sym- posium was held, but he was at that time in the throes of his terminal illness and was unable to attend. Also regrettable is the death in January 1987 of Harry MacGinitie. Early in his paleo- botanical career, Becker solicited and received much advice from Brown, Dorf, and MacGinitie, colleagues whom Becker greatly respected; thus, dedication of this symposium to all four men is particularly appropriate. As Dilcher (1987) notes, Herman F. Becker (1907-1985) entered paleobotany late in life, and thus his paleobotanical career, mostly on the staff of the New York Botanical Garden, covered less than two decades. In that time, however, Becker monographed several latest Eocene (Oligocene in some chronologies) floras from southwestern Montana and discussed in shorter contributions some enigmatic plant fossils. The Montana floras contain early records of many microthermal taxa, and Becker's extensive collections of these floras have been a major basis for expanding knowl- edge of the development of microthermal vege- tation and the evolution of microthermal taxa. Roland W. Brown (1893-1961) was for 30 years the Mesozoic and Cenozoic paleobotanist of the U.S. Geological Survey (see Mamay, 1963) and was particularly concerned with the Cretaceous and Tertiary floras of the Rocky Mountains and adjacent High Plains. His contributions included numerous short, largely taxonomic papers; Brown's sharp eye for details kept his contem- poraries “on their toes" in regard to determi- nations. However, Brown's major contribution was the monograph “Paleocene floras of the Rocky Mountains and Great Plains," which rep- resents the accumulation of a massive amount opment of the flora and vegetation in this region, ANN. Missouni Bor. GARD. 74: 683. 1987. or the general distribution of Paleocene vegeta- tion mad on Brown’s monograph. For many years, Erling Dorf (1905-1985) was a highly respected professor in Geology at Princeton cea His paleobotanical con- tributions "ass from studies of Devonian plants from Beart Butte, Wyoming, to studies of the 1. iin ci floras of California to taphonomic observations following the eruption of Paricutin in Mexico. Dorf was also concerned with general patterns of climatic change as evi- denced by land floras. Of particular relevance to this symposium, however, is Dorf’s fieldwork on rocks and their floras near the Cretaceous—Ter- tiary boundary in the northern Rocky Mountain region and particularly his monographic treat- ment of the latest Cretaceous Lance and Medi- cine Bow floras; this monograph remains the most recent and significant treatment of latest Creta- ceous plants in the northern Rockies. Harry D. MacGinitie (1896-1987) was a source of encouragement and advice to all concerned with the Tertiary floras of western North Amer- ica (Wolfe, 1987). His published contributions span 42 years (1933-1974) and include several major monographs of both Paleogene and Neo- gene floras, including those of the Rocky Moun- tains. MacGinitie's systematic work was always of high quality. The areas of historical plant ge- osraphy, paledecolce on analyses of fossil plant ange were, however, of major ‘concern to him, and embedded in his floral monographs are thoughtful and extensive discussions of the development of the vegetation and flora of western North America. Many now generally accepted concepts regarding the floris- tic and vegetational history of western North America during the Tertiary have their origins in MacGinitie’s work. LITERATURE CITED DircHER, D. L. 1987 [1988]. Memorial to Herman F. Becker (1907-1985). Ann. Missouri Bot. Gard. he P gta MAY, S. H. 1963. Memorial to Roland W. Brown ee: isc Geol. Soc. Am. Bull. 74: 79- Wo tre, J. A. 1987 [1988]. Memorial to Harry MacGinitie (1896-1987). Ann. Missouri Bot. he 74: 684—688. — Jack A. ie Paleontology and Stratigraphy Branch, MS-919, U.S. Geological Survey, Fed- eral Center, nr Colorado 80225, U.S.A Harry D. MACGINITIE MEMORIAL TO HARRY D. Back in the early 1950s, a bus trip from Port- land, Oregon, to Boston would have been a long one, especially for a kid just entering college. But not for this freshman, because I had along Harry D. MacGinitie's recently published “Fossil Plants of the Florissant Beds, Colorado." The copy is now well worn, the binding tattered. Why a bud- ding paleobotanist could and should have read this book is part of the story that follows. In December 1952, I visited Berkeley—then the Mecca of Tertiary paleobotany —and met with R. W. Chaney, the widely accepted leader of this field. Chaney assigned Marie Pabst the task of entertaining this possible novitiate. Among the many questions asked of Pabst were: who pro- duced good work, who was a model? Unhesitat- ingly, Pabst reached for the bookshelf and pulled out MacGinitie’s (1941) “A Middle Eocene Flora from the Central Sierra Nevada." “This is the best work that has been done in western Tertiary paleobotany," she declared. “This is the work to emulate.” Mac MacGinitie was a paleobotanist’s paleo- botanist to Pabst, as well as to his other col- leagues. He considered all evidence that bore on a scientific problem, and his conclusions and hy- potheses were written logically and clearly, es- chewing jargon whenever possible. He consid- ered all reasonable alternatives and stated basic assumptions. If he changed his mind on a subject, he clearly stated this and the evidence that re- sulted in the change. His knowledge of leaf ar- chitecture of the angiosperms was immense, and ANN. MISSOURI Bor. GARD. 74: 684—688. 1987. MACGINITIE (1896-1987) the great bulk of his determinations of fossils are still valid. Most importantly, Mac shared his knowledge continually by discussions with his colleagues and would even spend his time in the herbarium helping with identifications. In both personal contact and his publications, Mac has had a profound me in Tertiary paleobotany in North Amer By modern * oubli or perish" standards, Mac's publications were few. Each, however, contains a few to several essays that could have readily been extracted as separate papers. And, each 5 UN new approaches to Ter- tiary paleobot Mac's first uo NA in 1933 was the first comprehensive account of an upland Miocene flora from the Columbia Plateaus. In this paper, Mac's discussion of the climatic implications of the flora is the first to attempt to assign numbers to climatic parameters and clearly reveals his interest in, above all, paleoclimate. I recall one discussion with Mac in which he stated that he entered paleobotany largely because he thought that fossil plants were the best indicators of pa- leoclimates. He was well aware ofthe significance of altitude, and his assignment of about 2,500 feet to the Trout Creek has not been significantly altered by newer (and supposedly more reliable) techniques. The Weaverville flora, published in 1937, was Mac's Ph.D. thesis, which was completed under R. W. Chaney in 1935. This year also saw another major event in Mac's life; Beatrice MacGinitie 1987] WOLFE— (nee Hess), who became his wife on February 2, 1935, was a constant source of encouragement and emotional support to Mac for the rest of his life. The Weaverville flora reveals Mac’s increasing involvement with both the detailed climatic im- plications of fossil-plant assemblages and the possible causes of climatic change. Further, the detailed discussion of the geologic occurrence of the fossils and of basinal geology placed the Weaverville flora in a taphonomic context. Mac attributed the Weaverville flora to the early Oli- gocene, recognizing that the flora contained ele- ments that indicated an age younger than the Goshen flora. The monograph of the Eocene Chalk Bluffs flora, published in 1941, shows a maturation of many concepts. As in his earlier works, Mac con- sidered the paleoecological significance of the fossil taxa in a depositional framework. He pre- sented solid evidence for the co-occurrence of taxa whose modern counterparts have markedly different climatic tolerances and suggested that at least some tolerances have changed through time. Mac documented that the neotropical ele- ment in the Chalk Bluffs was today largely found in the Mexican uplands and that the flora also contained a large paleotropical element. This monograph also contains the first significant dis- cussion of the plant biogeography of the North American Eocene. Mac laid the cornerstones for future discussions of the historical biogeography of the western North American Tertiary. The taxonomy in the Chalk Bluffs paper has largely withstood the test of time. Mac, using solid leaf-architectural criteria, demonstrated that the many leaves of “Aralia,” a major element in western American Paleogene floras, represented a platanaceous genus. He referred the leaves to Platanophyllum and discussed the phyletic re- lationships of the species; recently they have been reassigned to the new genus Macginitiea, named eo climatic inferences still remain valid is revealed in the Chalk Bluffs monograph: “In no case were ecological considerations given weight in the choice between two living forms as modern rep- resentatives of fossil plants... . If identifications are based on ecology, and then ecology is de- duced from identified species, the result is a kind of cyclic reasoning which may lead to consid- erable error" (MacGinitie, 1941: 96). While teaching briefly at the University of Col- NORTHERN ROCKY MOUNTAIN SYMPOSIUM 685 orado, Mac was associated with T. D. A. Cock- erell, the paleoentomologist who had worked ex- tensively with insects from the famous lake beds at Florissant, Colorado. No comprehensive treatment—systematic or ecologic—existed of the Florissant flora, and, at the urging of Cockerell and the vertebrate paleontologist Childs Frick, Mac began work on the Florissant flora in 1936, work that culminated in his 1953 monograph of the flora. Mac’s research was, however, interrupted by World War II. Although Mac was 45 when the United States entered the war, he still decided to serve. Because of Mac’s expertise in the the- oretical aspects of climatology and meteorology, he enrolled as an instructor in the U.S. Army Air orps. Lt. MacGinitie’s primary responsibility was teaching crewmen the significance of, and to recognize, different cloud formations. After the war, Mac returned to Humboldt State and to his work on the Florissant. Above all, the Florissant work involved care- fully detailed taxonomic work. Mac reduced the 258 species previously attributed to the flora to less than half that number, but still the Florissant is the largest flora yet described from the Tertiary of western North America. One common prac- tice in Tertiary paleobotany was the erection of a “fossil” species for fossils that resembled a giv- en living species. Mac, however, challenged that practice: “Fossil plants in two different floras may be likened to the same /iving species without in any way implying identity ofthe two fossil forms. Their differences may be great enough in oppo- site directions to place them in different species ." (MacGinitie, 1953: 79). As Mac noted, the usual practice led to erroneous concepts of age and floristic relationships. In the Florissant monograph, Mac introduced and elaborated on a number of new ideas and Observations that have subsequently been well substantiated. Particularly significant is the idea that given lineages may persist in a region by adapting to changing environmental conditions; he was led to this conclusion because of the shar- ing of numerous lineages between the older Green River flora and the Florissant. Mac also clearly stated (p. 46) that fossil leaf are biased towards streamside or lakeside vege- tation and, hence, may not adequately reflect the regional flora and vegetation; this point has been increasingly made apparent by many tapho- nomic studies. In discussing the paleoclimatic significance of floras then placed in the Oligo- 686 cene, Mac noted: “The point emphasized by this discussion is that if we accept a Middle Oligocene age for the Goshen flora, the remainder of that period must have witnessed an almost complete revolution in the flora of the region, and more critical events must be crowded into the Upper Oligocene than paleontologists have hitherto been willing to concede” (MacGinitie, 1953: 67); Mac was the first to recognize the major and rapid climatic change that is now placed near the end of the Eocene. Mac completed the Florissant study in 1951. Probably because of administrative duties as Chairman of Natural Sciences at Humboldt State, Mac did not immediately start on a new project. In 1954, James Bump sent Mac some leaves from a new site in Nebraska, and encouragement from Bump and Harold J. Cook enticed Mac to start fieldwork on the Kilgore flora, which was com- pleted in 1958. Following his retirement from teaching in 1960, Mac finished the systematics of this assemblage. He had become increasingly aware of the value of palynology in paleoecolog- ical interpretations, and, despite no prior paly- nological experience, he became knowledgeable of basic pollen morphology with the help of Es- tella Leopold. The Kilgore paper was the first in North American Tertiary paleobotany to illus- trate both megafossils and microfossils and ex- tract from both types of fossils paleoecological data. The climatic discussion in the Kilgore pa- per was based on a wide range of data (paleo- botanical, sedimentological, vertebrate, and molluscan) and probably remains the most com- prehensive discussion of Neogene climates in the Plains region of North America. The Kilgore paper also contains an expansion of the thesis, first expounded in the Florissant paper, that many plant lineages can persist in a given region despite significant climatic change. If, as Mac demonstrated, the bulk of the Kilgore species were derived from older species in the . it should be questioned ry. dro-Tertiary,’ and the like imply esten els use- ful concepts if we do not think of these terms as representing areas or centers from which mass migrations occurred. They picture to us in a gen- eral way the vegetation occupying an area, al- though the particular type of vegetation was slowly changing" (MacGinitie, 1962: 87). This statement by Mac represented a clear-cut depar- ture from the geofloral hypotheses then so widely ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 accepted in North American Tertiary paleobot- any and historical plant geography. However, as originally submitted to the Board of Editors of the University of California Publications in Geo- logical Sciences, Mac's manuscript was unchar- acteristically vague about this point. Clyde Wahrhaftig, one of the editors, asked me what Mac meant. From numerous discussions with Mac, I knew what he was attempting to state, and at Wahrhaftig's request, Mac rewrote his manuscript to the statement quoted above. On retiring from Humboldt State, the MacGinities moved from Arcata to Napa, a move that was beneficial to Mac and especially to many students and colleagues. He was made a Research Associate at the University of Cali- fornia Museum of Paleontology and had read access to extensive paleobotanical collections (many of which he had made) and the University of California Herbarium and libraries. He was in an area active in the geological and botanical sciences, and Mac was often to ve found in deep discussions gues, to whom he freely gave of his time. Every thesis in paleo- botany completed at Berkeley from the 1960s through the 1980s acknowledges Mac’s assis- — £2 nce. With typical enthusiasm, Mac then began work on the Eocene floras of the Rocky Mountains to understand better the floras and vegetation that preceded the Florissant. The first of these floras, that of the upper part of the Green River For- mation in Utah and adjacent Colorado, was, like the Florissant, a classic flora. The last significant descriptive work, that by Roland Brown, was more than 30 years old, and the paleoecological significance of the Green River flora had never Eocene localities in Wyoming. In 1968 he com- pleted work on the Green River flora (Mac- Ginitie, 1969) and in early 1972 on the Kisinger Lakes flora (MacGinitie, 1974). He continued work on other Wyoming Eocene floras, but, un- fortunately, none were completed for publica- tion. A major hindrance to completion of these works was a deterioration of leg joints, which required surgery and made fieldwork difficult and painful. In Mac’s last two works on the Green River and Kisinger Lakes floras are the same type of careful and detailed discussions of paleoecology and paleoclimatology that are present in his ear- lier publications. Certain parts of these mono- 1987] graphs should be carefully read by any paleobot- anist or student in paleobotany, e.g., the chapters "Certain Aspects of Floristic Evolution" and "Paleobotanical and Botanical Species" (MacGinitie, 1969: 68-70, 81-86), and “Distri- bution of Correlative Living Species” (Mac- Ginitie, 1974: 29-34). Indeed, Tom Taylor and Edie Smoot selected the second-listed chapter as one of eight papers on Tertiary paleobotany in the “Benchmark Papers in Systematic and Evo- lutionary Biology Series" (MacGinitie, 1984). Mac’s discussions of systematic determinations also had become more sophisticated; he contin- ually discussed the exact reasons certain generic determinations were made (not that a fossil tax- on simply “looked like” an extant taxon) and illustrated mu ofthe material in detail. Written with Estella Leopold, who also contributed a chapter to the Kisinger Lakes paper, the sum- mary of floristic and vegetational development in the Tertiary ofthe Rocky Mountains (Leopol & MacGinitie, 1972) remains a most useful and concise summary on that topic. The symposium held in honor of MacGinitie at the 1983 annual meeting of the American As- sociation of Stratigraphic Palynologists was a well-deserved tribute. Palynologists recognized both the wide scope of Mac's contributions throughout his career and his bringing together of palynology and megafossil research in the gore and later papers. During a tribute listing his many accomplishments, Mac, sitting in the front row, nudged Estella Leopold and whispered, “I never realized that I was that good." Mac had generally lived in the shadow of R. W. Chaney, and even when disagreeing with Chaney or other colleagues, was careful not to cite by name the originators of hypotheses; instead, Mac would simply and clearly, through data and logic, refute the hypotheses. One of the rare instances when Mac named names was, in fact, at the 1983 AASP symposium when he stated of a colleague, “He was a nice man but a terrible paleobotanist," much to the amusement of the audience. One of Mac's major attributes was his ability to grow intellectually, to accept new concepts and weld them into his already considerable frame- work of floristic evolution and vegetational/cli- matic change in the Tertiary of western North H '1 merica. If new and/or refined techniques of analysis (e.g., palynology and leaf architecture) appeared, he s among the first to use these techniques. Mac kept up with advances in pa- leobotany even after he was no longer an active WOLFE—NORTHERN ROCKY MOUNTAIN SYMPOSIUM 687 researcher in the mid 1980s. Above all, Mac was always ready to discuss with colleagues any prob- lem of mutual interest and to share with them his vast wealth of knowledge and experience. Fortunately, part of Mac’s legacy to us—his en- quiring mind and thoughtful statements—will endure in his publications. When Mac ceased being an active researcher, he, in typical generos- ity, brought down to Berkeley his entire profes- sional library, leaving for present and future stu- dents an additional legacy. Thanks are due to Beatrice Ann Minkler (nee MacGinitie) for supplying much pertinent bio- graphical material. Howard E. Schorn supplied the accompanying photograph and contributed to some of the content of this memorial. Patrick F. Fields supplied the list of MacGinitie’s pub- lications. BIOGRAPHICAL DATA Harry Dunlap MacGinitie was born in Lynch, Ne- braska, on March 29, 1896. After graduating from high school in Sturgis, South Dakota, he moved to Califor- nia. He attended and received an A.B. from Fresno State College in 1926, after which he attended Stanford University for a year. In 1926-1 1928, Humboldt State College (now University) in Arcata, California, where he taught until 1960, except for a year (1932-1933) when he attended Berkeley full time, a year (1936-1937) when he taught at the University of Colorado (Boulder), and two years (1943-1945) when Paleontology. He was a Fellow of emy of Sciences and the Geological Society of America and a member of the Paleontological Society. He died on January 31, 1987, in Napa, California. PUBLISHED CONTRIBUTIONS OF Harry D. MACGINITIE 1933. Redwoods and frost. Science Ser. 2, 78(2018): 90. The Trout Creek flora of southeastern Oregon. Publ. Carnegie Inst. Wash. 416: 21-68, 16 pls. Ecological aspects of the floras of the auriferous gravels. Geol. Soc. Amer. Proc. for 1933, Ab- stracts, p. 356. [Abstract.] Tertic floras of Trinity County, California. Pan- Amer. Geol., Abstracts, 62: 75-76. [Abstract.] Tertiary floras of Trinity County, California. 1933. 1934. 1934. 1935. 688 1937. 1937. 1938. 1938. 1941. 1943. 1953. 1958. i A 1961. 1962. ANNALS OF THE MISSOURI BOTANICAL GARDEN Geol. Soc. Amer. Proc. for 1934, Abstracts, p. 390. [Abstract.] Stratigraphy and flora of the Florissant beds. Geol. Soc. Amer. Proc. for 1936, Abstracts, pp. 362-363. [Abstract.] The flora of the Weaverville beds of Trinity County, California. oo Carnegie Inst. Wash. 465: 83-151, Stratigraphic — in the Tertiary gravels of the Sierr vada. Geol. Soc. Amer. for 1937, aN pp. 246-247. [Abstract.] Geologic relations along the southwest border of the Klamath Mountains. Geol. Soc. Amer. Proc. for 1937, Abstracts, p. 247. [Abstract.] A middle Eocene flora from the central Sierra Nevada. Publ. Carnegie Inst. Wash. 534: 1-178, 47 pls. Central and southern Humboldt County. Calif. Div. Mines Bull. 118: 633-635. Fossil plants of the Florissant beds, Colorado. : 1-198, 75 pls. Late Cretaceous. Pp. 61-79 bs (editor), pes raphy. Amer. gocene plants from the upper Ruby River Basin, southwestern Montana" by H. F. Becker). Ecol- ogy 42: 853-854. The Kilgore flora, a late Miocene flora fro northern Nebraska. Univ. Calif. Publ. Geol. Sci. 35: 67-157 1968. 1969. 1972. 1972. 1974. 1984. [VoL. 74 Some vegetation types in the Eocene of l. mid- dle Rocky Mountains. Geol. Soc. Amer. Spec. Pap. 101: 321, 408-409. [Abstract.] The Eocene Green River flora of northwestern Colorado and iesu Utah. Univ. Calif. Publ. Geol. Sci. 83: 3, (Wi th E. B. ay Development and affin- and Paleofloristics of Asia and Eastern Nort America. Elsevier Publ. Co., Amsterdam (With R. A. Scott, P. L. Williams, L. C. Cra age of the palm woods and roots. Amer. J. Bot. 59: 886-896. An early middle Eocene flora from the Yellow- stone-Absaroka volcanic pon OV ince, northwest- ern Wind River AR Wyoming. Univ. Calif. Publ. Geol. Sci. 108: 210» The Eocene Green E flora of northwestern Colorado and northeastern Utah (Systematic considerations. Paleobotanical and botanical species). Pp. 262-267 in T. N. Taylor & E. L. Smoot (editors), Paleobotany, Part II. Triassic through Pliocene. Van Nostrand Reinhold Co., New York —Jack A. Wolfe, Paleontology and Stratigraphy Branch, MS-91 9, U.S. Geological Survey, Fed- eral Center, Denver, Colorado 80225, U.S.A HERMAN F. BECKER MEMORIAL TO HERMAN F. BECKER (1907-1985) Herman F. Becker was born in Duesseldorf, Germany, on January 10, 1907. After complet- ing his initial education at the Realgymnasium in Neunkirchen (1923), he undertook the study of botany in Dahlem (Berlin) at the Botanical Gardens with the encouragement of his father and grandfather. There he completed studies as a horticulturist. He met his wife, Ruth, in Berlin and they were married in Frankfurt, Germany in 1928. He saw no possibility of continuing his study of botany in Germany so he decided to go to the United States and arrived in New York in March 1930. His wife and young son followed about one year later after he had obtained work as a Horticultural Assistant (1930-1939) at the Brooklyn Botanical Garden. Twin sons were born in New Beginning in 1939 he taught at Brooklyn Col- lege as a Lecturer in Geology while he undertook his B.A. degree, which he completed in 1947. He continued to teach at Brooklyn College while earning an M.A. degree from Columbia Univer- sity in 1952. ANN. MISSOURI Bor. GARD. 74: 689-691. 1987. During this time he took part in a geology field trip to the West with several geology students. While on this trip he was told, “Becker, you look for plant fossils," because everyone knew of his interest in plants. He came back to New York excited about a shoe box full of fossil plants that he had collected. Becker wanted to continue in research but had little opportunity to do so while teaching a heavy course load as a Lecturer and later Instructor at Brooklyn College (1939-1955). However, in the summer of 1947 he began field studies under the direction of Professor A. J. Eardley, director of geologic work at the Rocky Mountain Field Sta- tion of the University of Michigan. Soon he was directed to Professor Chester Arnold. He con- tinued to maintain contact with Arnold and col- lected fossils in the Ruby River Basin area of southwestern Montana during the summer of 1949. He then enrolled in a Ph.D. program at the University of Michigan to work in Paleo- botany with Professor Arnold. At first he worked on his degree in the summers. But, after com- pleting additional fieldwork on the paper shales 690 of the Ruby River Basin in the summer of 1955, he remained at Michigan, completing his Ph.D. degree in 1956. He was awarded the Ermine Cowles Case Award in Geology for his outstand- ing Ph.D. dissertation. Herman Becker attempted to teach again at wasa Cie but the heavy course load gave him no time for his research on the fossil plants of the Ruby River Basin to which he was by this time very committed. Thus, in 1958 he joined the New York Botanical Garden, first as a Re- search Associate and later as a Curator of Paleo- botany. Between 1957 and 1968 he made seven more major collecting trips to southwestern Montana, which provided him with the research material that formed the basis of many of his publications. The first two major floristic papers he pub- lished were on the paper shale beds of the Ruby River Basin flora (middle Late Oligocene) (196 1a) and the Mormon Creek flora (upper Middle Oli- gocene) (1960c). With continued extensive sum- mer fieldwork in southwestern Montana, he pub- lished monographic papers on the Beaverhead Basin (lower Middle Paleocene) (1969c), the Metzel Ranch flora in the Ruby River Basin (lower Late Oligocene) (1972b), and the York Ranch flora of the Ruby River Basin (Upper Oli- gocene—Lower Miocene) (1973b). In these floris- tic papers he not only presented the fossil flora from the specific fossil beds he collected, but he applied the resulting data to questions of the cor- relation of their floristic elements to other North American Tertiary floras, taxonomic age rela- tionships of the specific floras, and the phyto- geographical distribution of modern equivalents for the fossil plants described. Herman Becker worked at a time when the questions being asked in the field of paleobotany were concerned primarily with paleoecological and phytogeographical issues. However, perhaps because of his strong botanical background, he developed a special interest in the relationship between fossil and extant plants and the evolu- tion of plants through time. In his paper on the Beaverhead flora he wrote: “The writer does not agree to calling Tertiary fossil species ‘identical’ with any extant species, and applying to it the modern specific epithet, especially where Mio- cene-Oligocene, or older plants are concerned . In a strict sense, no individual of a modern species is morphologically or genetically identi- cal with another, a fact expressed in major or minor variability as a phenotype. Genetic, phys- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 iological, and many morphological characters are usually not available to the paleobotanist to pro- nounce a specimen identical or NA equivalent to a modern form" (1969c: 10). The importance of his research was pa M by several honors he received after his retire- ment. In 1977 he was awarded a special Distin- guished Service Award for his research in paleo- botany by the Paleobotany Section of the Botanical Society of America, and in June 1985 a special symposium was held in his honor. This symposium was sponsored by the Pacific Sec- tions of the American Association for the Ad- vancement of Science and the Botanical Society of America and was titled “Evolution of the Modern Vegetation of the Northern Rocky Mountains." Several well-known scientists pre- sented papers at this symposium and a special plaque was presented to Dr. Becker. In 1981 he received a Distinguished Achievement Award from the Brooklyn College Alumni Association. Becker was always characterized by an out- going friendly nature, an interest in people, and an enthusiasm for plants, living and fossil. His extensive research papers, which students of pa- leobotany continue to study and cite, stand as a record of his persistent hard work in the field and the laboratory. His research collections provide an exceptional documentation of the history of ancient plants that once lived in the area of pres- ent-day southwestern Montana. To paleobota- nists of now and of the future, Herman Becker showed pages of new chapters in plant history when he dug out and pried open the “books of paper shale" at the Ruby River Basin. He will be remembered as a friend, a good colleague, and a research scholar. PALEOBOTANICAL PUBLICATIONS OF HERMAN F. BECKER Coauthor: W. Donn occurrence and description of the fossil d noA UE Science 115(2982): 214-215. 1955. A new method of! dli Ë di ving soft matrix fossils. Asa Gray Bull., N.S. 3: 57-58. An Oligocene Flora from the Ruby River Basin in Southwestern Montana. Ph.D. Dissertation. University of Michigan, Ann Arbor, Michigan. A new species of Mahonia from the Oligocene Ruby flora of southwestern Montana. Contr. 1. Paleontol. Univ. "Michigan 15(3): 33-38, 1952. 1956. 1959. 1960a. ES to Eopuntia one Cact. Succ. J. (Los Angeles) 32: 28-2 The Ruby flora of u SS Montana. Bil- lings Geological Society, Eleventh Annual Field Conference Guidebook. September 7-10, 1960. 1960b. 1987] 1960c. 1960d. 196la. 196 1b. 1962a. 1962b. 1962c. 1962d. 1962e. 1962f. 1962g. 1964a. 1964b. 1965a. 1965b. 1966. DILCHER — NORTHERN ROCKY MOUNTAIN SYMPOSIUM The Tertiary Mormon Creek flora from the Upper Ruby River Basin. Palaeontographica, Abt. B, Paláophytol. 107(406): 83-126, pls. 18- Additions to the Tertiary Ruby paper shale flora of southwestern Montana. Bull. Torrey Bot. Club 87: 386-396. Oligocene plants from the Upper Ruby River Basin in southwestern Montana. Mem. Geol. Soc. Amer. 82: 1-127, 32 pls. esas ay record of solar change. Ann. New York Acad. Sci. 95: 684-687. The resurrection of a landscape. Gard. J. New York Bot. Gard. 12: 8-1 Roland W. Brown, 1893-1961. Gard. J. New York Bot. Gard. 12: 71-72. Two new species of es from the Grant- Roses of the past. Gard. J. New York Bot. Gard. 13: 103-105. Roland W. Brown, i, Bull. Torrey Bot. Club 89: 260-26 ee 2 P to C Vader det di erus, comb. . Bull. Torrey Bot. Clu 301-307. nls a Pp. 150-153 in American ord Jr. peepee eg ‘additions to the Oligocene ora of southwestern Montana. Bull. Torrey Bot. Club 91: 206-213 Paleobotanical exploits in Colorado and Kan- sas. Gard. J. New York Bot. Gard. 14: 231- 233. Stir abi and evolution. Nat. Hist. 74(2): 3 Rare ES "iom Montana (insects). Animals (England) 8: 9 Additions to 2 revision of the Oligocene Ruby paper shale flora of southwestern Mon- tana. Contr. Mus. Paleontol. Univ. Michigan 20(5): 89-119, 6 pls. 1967a. 1967b. 1968. 1969a. 1969b. 1969c. 1971a. 1971b. 1972a. 1972b. 1973a. 1973b. 1976. 691 On western fossils and ss plants. Gard. J. w York Bot. Gard. 17: 30-33. Flowers, insects, and irure (1965). R printed as pp. 392-399 in W. Knobloch (edi- tor), Readings in Biological Science, 2nd edi- tion. Appleton, New Yor A hexasepalous calyx of the fossil Astronium truncatum (Lesquereux) MacGinitie. Bull. Torrey Bot. Club 95: 262-263, 1 pl. ea pis cc forests. Amer. Forests 75(3): 1 Some Mtis Dad fossils. Amer. Forests 75(7): Fossil plants ofthe Tertiary Beaverhead basins in southwestern Montana. Palaeontographica, Abt. B, Palàophytol. 167: 1-144, pls. 1-44 Fossil collecting. Gard. J. New York Bot. Gard. 21: sh Digging in Montana’s evolutionary past. Amer. ineft TIO 44-47, 57- Kun Sanmiguelia, an enigma ounded. Palae- Tire Abt. B, Paldophytol. 138: 181- 185, p The md pw flora of the Upper Ruby River Basin, southwestern Montana. Palae- ontographica, Abt. B, Paláophytol. 141: 1—61, pls. 1- yet Tertiary gramineous fossil. Bull. Torrey Bot. Club 100: 318-320. The York Ranch flora of the Upper Ruby Riv- er Basin, southwestern Montana. Palaeonto- graphica, Abt. B, Palàophytol. 143: 18-93, pls. 13-40. Like a phoenix from the ashes, Oregon's long- vanished forests are revealed in fossils. Gard. J. New York Bot. Gard. 26: 161-165 — David Dilcher, Department of Biology, Indiana University, Bloomington, Indiana 47405, U.S.A. LAND PLANTS OF THE NORTHERN ROCKY MOUNTAINS BEFORE THE APPEARANCE OF FLOWERING PLANTS CHARLES N. MILLER, JR.! ABSTRACT Bana £1 4 $ £, +h flowering plants near the end of the Early Cretaceous is spotty. Seas covered much of the land during the Paleozoic and ea rly Mesozoic, but certain areas were exposed, perhaps as islands. Evidence of Devonian vegetation from Beartooth Butte, Wyoming includes rhyniophytes, iens zosterophylls, and lycopsids. Fossils from several different de- posits show Late Mississippian to The floras include lepidodendrids, Eus mites, fer nnsylvanian e vegetation like that of coal swamps to the east. , seed fern d PUn dne conifers present at nder a generally arid but seasonally wet frost-free climate. It was into this environment that the first u flowering plants of the region migrate The object of this paper is to give an overview of the types of vegetation that occurred in the northern Rocky Mountain region before the ap- pearance of flowering plants there. This region extends along the Rocky Mountains from central Colorado and Utah north into southern Alberta and British Columbia. The time span involved extends from the Early Devonian Siegenian, about 385 million years ago, to the Early Cretaceous Aptian, about 110 million years ago (Fig. 1). During the intervening 275 million years both the vegetation and the land it occupied changed radically, keeping pace with similar vegetational and land mass changes elsewhere in the world. In fact, evidence of vegetation within the pres- ent northern Rocky Mountain region is spotty (Fig. 2). The best records of Paleozoic vegetation come from deposits in Europe and eastern and midwestern North America. Similarly, early Me- sozoic, i.e., Triassic and Jurassic, floras are more abundant and better preserved in Europe and the southwestern United States than in the northern Rocky Mountains. It isn't until the Late Jurassic and Early Cretaceous that records within the re- gion are as good as those found elsewhere. The Paleozoic and Mesozoic sites that are known in the northern Rocky Mountains, however spotty, show that the types of plants in the region are similar to those elsewhere and that they probably were organized in similar communities which occupied similar habitats. Thus, while the de- posits within the region fail to provide conclusive evidence, they nonetheless permit inference based on our knowledge of nearby vegetation outside the area. DEVONIAN During the Early Devonian, the only part of the period for which we have evidence, the pres- ent northern Rocky Mountains was an area of lowlands and shallow seas along the western margin of the early land mass Laurussia (Bam- bach et al., 1980). This continent lay across the equator, extending from about 10? south latitude to about 40? north latitude, with the present northern Rocky Mountains situated at about 12? north latitude (Ziegler et al., 1981, fig. 7.5). Be- cause of its position, the area had a tropical cli- a rm t rainfall throughout the year. This, combined with the location of the northern Rocky Mountains in the Devonian along the western coast of the continent where sharp changes in climate were moderated by ocean currents, resulted in what must have been ideal growing conditions. Two deposits of Early Devonian plant fossils have been recorded in the Beartooth Butte For- mation in the northern Rocky Mountains. The classic site at Beartooth Butte, Wyoming (Fig. 1 A) has been known for over 50 years. Five taxa were described by Dorf (1933, 1934a, 1934b) ! Department of Botany, University of Montana, Missoula, Montana 59812, U.S.A. ANN. MissouRi Bor. GARD. 74: 692-706. 1987. 1987] MILLER—LAND PLANTS OF NORTHERN ROCKY MOUNTAINS 693 O 110 D E ° mo) ra RSTU NN 135m = MNOPQ | | | = rm co [j=] N =! Pis 5 B D ° > ay e w | x | | | — n: A uewad X S osseu| 3 Olsseunr _ z r è o ka om xr 9 - | 7n - e “8 g $5 S uueg Š 310Z031Vd > [vv] upiuoAə | E = a =] FIGURE 1. Index map and P time nia om showing the approximate locations and ages of m D Early De vonian.— B. Cottonwood Gulch, Ea anning Canyon Shale, Early Pennsylvanian. — H r Formation, Early Pennsylvanian.—I Coy Formation, Late Penn- sylvanian.— J. Spotted Ridge Flora, Pennsylvanian. — K. Cutler Formation, Permian. Wyoming Red Beds, Triassic. — M. Morrison Flora, Utah, Late Jurassic. — N. Morrison Flora, Wyoming, Late Jurassic. — O. Morrison T oe Late Jurassic.—P. Kootenay Formation, Late Jurassic.—Q. Douglas County, ei Late Jurassic. — R. Kootenai Formation, Early Cretaceous. — S. Lower Blairmore, Early Cretaceous. — T. Shasta Flora, Early yc —U. Burro C anyon Formation and Cedar Mountain Formation, Early Cretaceous. based on imprints of plant fragments. More re- are not isolated occurrences and that further cently, Hueber (1972) and Tanner (1982, 1983) described additional remains from the locality. he deposit is regarded as Emsian in age, about 375 million years old. Tanner (1982, 1983) described fossils from the new Beartooth Butte Formation locality near Cottonwood Gulch (Fig. 1 B) in northcentral Wy- oming, about 50 km east of the original site (Fig. 1A). It is believed to be Siegenian in age, about 385 million years old, and thus somewhat older than the Beartooth Butte locality. Locating a sec- ond deposit in the region raises hope that these search will turn up more deposits. Five taxa were originally described by Dorf (1933) from the Beartooth Butte locality. These include Psilophyton wyomingense Dorf for flat- tened branch systems bearing numerous spines (Fig. 2A, B), (?) Psilophyton sp. for an elongated bearing numerous lateral sporangia (Fig. 2C, D), Hostimella sp. for flattened branch systems lack- ing spines, and (?) Broggeria strobiformis for a cylindrical strobiluslike structure. Hueber (1970) ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FIGURE 2. Examples of plant fossils from the Early Devonian d Butte Formation. A, B. Psilophyton wyomingense Dorf. — A. Holot 5), x 2.5. —C. Rebuchia ovata (Dorf) Hueber, USNM 1689 C, D from photographs courtesy of F. M. Hue changed the name of Bucheria ovata to Rebuchia ovata and later (1972) described its vegetative axes linked with the fertile stems (Fig. 2C, D). These clearly show the affinity of the plant with the Zosterophyllophytina. More recently Tanner (1982, 1983) described additional taxa from both sites of the Beartooth Butte Formation, bringing the total known to 14. Six of these are based on vegetative axes with attached fertile parts, five are known from veg- etative axes alone, while one taxon is based on a sporangial pair. Present are rhyniophytes, tri- merophytes, zosterophylls, and the early lycop- sids Drepanophycus and Leclercquia (Tanner, 1983). There are also two taxa of undetermined botanical affinities. Discussion. Even though the two localities occur in the same formation, palynological evi- otype, from Dorf (1933, plate V, fig. 3), Hueber, USNM 168991B counterpart, 92, x1.3. (Figs. A, B reproduced by permission of the University of Chicago Press; Figs. er.) B. from Dorf (1933, plate V, fig. s 3.—D. Rebuchia ovata (Dorf) dence indicates that they may be separated by as much as 20 million years. Such a gap is not ap- parent in the megafossils or the fish fauna pre- served at both, although the Cottonwood Creek assemblage lacks definitive marker species (Tan- ner, 1983). n comparison with other Early Devonian as- posed by Gierlowski (Ziegler et al., 1981) in which the western North America zone is characterized by relatively low generic diversity of two to six taxa per assemblage and a lack or restriction of zosterophylls. Including the unnamed sporangial 1987] pair and two morphotypes of sterile axes, the Wyoming assemblage has ten genera, three of which belong to the Zosterophyllaceae. Early Devonian plants share an architectural plan that with few exceptions consists of rela- tively naked aerial branches up to a meter tall, which branch from rhizomes that bear adven- titious roots or rhizoids (Gensel & Andrews, 1984). This is often interpreted as implying that these plants grew in marshy habitat. Since many of the known Early Devonian plant assemblages, and particularly the Wyoming sites, are from near equatorial paleolatitudes with equable temper- atures and moisture, transpiration stress may not have been enough to require actual growth in standing water or even saturated soil as would be found in coastal marshes. Occurrence at more inland and upland sites may have been possible. MISSISSIPPIAN While movement of other continental plates between the Early Devonian and Mississippian was significant, the position of Laurussia, and thus the present northern Rocky Mountains, was relatively unchanged. There was an apparent in- crease in mountain building activity west of Lau- russia and an increase in the amount of shallow seas and evaporite formation (Bambach et al., 1980, fig. 11). Much of the northern Rocky Mountains was an evaporitic basin in which limestone was deposited. However, some land was exposed within this zone of shallow seas, for plant remains are recorded as petrifactions in northwestern Utah (Fig. 1C; Tidwell & Jennings, 1986), as impressions and as pollen and spores in northwestern Utah (Fig. 1D, E; Arnold & Sad- lick, 1962; Schemel, 1950), and as pollen and spores associated with exceptionally well-pre- served fish and invertebrates at Bear Gulch (Fig. 1F) near Lewistown, Montana (Cox, 1986). Deseret Limestone petrifactions. Tidwell & Jennings (1986) described a new lycopod, Stans- burya, from the Early Mississippian Deseret Limestone near Grantsville, Utah, south of Great Salt Lake. Six stem specimens were included in a single carbonate concretion. The stems are about 1.4 cm in diameter and are dichotomously branched. Scalelike leaves are tightly appressed helically around the stems. None of the axes con- tain secondary tissues, and this led the authors to interpret the fossils as representing an her- baceous plant. Much remains to be learned about this new genus, but discovery of these fossils in MILLER—LAND PLANTS OF NORTHERN ROCKY MOUNTAINS 695 the Deseret Li t t further search may be rewarding. Daggett County, Utah, Palynoflora. Schemel (1950) reported several different types of pollen and spores isolated from a coal sample from Late Mississippian sediments in northwestern Utah (Fig. 1E). Over 90% of the palynomorphs belong to Densosporites. Similar spores have been found in Pennsylvanian age sediments both dispersed and in strobili of presumed arborescent lycopods (Taylor, 1981). Several other types of spores oc- cur in the Utah coal in small amounts and suggest that sphenopsids, ferns, pteridosperms, and cor- daitaleans grew in the vicinity. Uinta Mountains Flora. The megafossil as- semblage reported from a locality in the Uinta Mountains in northeastern Utah (Fig. 1D) is a small one based on poorly preserved imprints of vegetative remains (Arnold & Sadlick, 1962). Ar- chaeocalamites was identified from pith casts. These show ribs continuous across the nodes rather than alternate as in Calamites. The oc- currence of Lepidodendron is patina by im- prints of characteristic stems, leafy twigs, and detached leaves. Defoliated axes, probably of seed fern fronds, identified as Cau/opsis, are present, as are detached fern or seed fern pinnules of Fryopsis (formerly Cardiopteris). Lastly, Rhodea, frond portions of a seed fern characteristic of the Late Mississippian and Early Pennsylvanian, has also been identified. Arnold & Sadlick (1962) interpreted the plant remains as representing streamside vegetation with fragments washed into an estuary or bay by river currents. The plant fragments are small and are imprints with occasional flecks of carbon. However, the siltstone matrix contains numer- ous flecks of black material that presumably rep- resent macerated plant debris (Arnold & Sadlick, 1962). The assemblage is regarded as Chesterian in age based on associated invertebrates (Arnold & Sadlick, 1962) and is correlated with the Late Visean or Early Namurian stages in Europe. The plant assemblage compares well with floras of similar age in the Appalachian trough. Bear Gulch palynoflora. Palynomorphs are also preserved in rocks that have given up ex- ceptionally well-preserved vertebrate and inver- tebrate fossils at Bear Gulch, near Lewistown, Montana (Fig. 1F; Melton, 1971). The palyno- morphs (Cox, 1986) include: Acanthotriletes, Anapiculatisporites, Chaetosphaerites, Convolu- tispora, Cyclogranulatisporites, Denosporites, 696 Endosporites, Lycospora, Procoronospora, Punc- tatisporites, Raistrickia, Rotaspora, Savitrispo- rites, Tripartites, and Verrucosisporites. The as- semblage indicates a Chesterian age for the deposit and correlates with the Namurian A stage of Europe. Furthermore, the palynoflora indicates that a diverse assemblage of ancient vascular plants in- habited the land that then surrounded the bay in which the Bear Gulch sediments were depos- ited. Although the correlation of the pollen and spore species found here with the plants that pro- duced them remains uncertain, other species of some of these genera have been found in spo- rangia. By analogy the following megafossil forms may have been present: lepidodendrids (Denos- porites and Lycospora), zygopterid ferns (Con- volutispora), progymnosperms ee calean fern (Raistrickia), marattialean ferns ae rites), and seed ferns (Punctatisporites and Schul- zospora) (Kosanke, 1950; Millay & Taylor, 1979; Taylor, 1981) Discussion. The two palynofloras and the Uinta Mountain megaflora combine to give solid evidence that vegetation similar to that occurring in eastern North America and Europe was pres- ent in the northern Rocky Mountains during the Late Mississippian. rites), PENNSYLVANIAN By Pennsylvanian time Laurussia was con- nected to Gondwana in the process of forming Pangaea, but the present northern Rocky Moun- tains remained at about 10° north latitude (Bam- bach et al., 1980; Ziegler et al., 1981). More of the present land surface was covered by shallow seas than earlier, but sufficient land was exposed to support vegetation characteristic of the period. Early Pennsylvanian vegetation is well in evi- dence in the Manning Canyon Shale flora from northern Utah (Fig. 1G). Similar remains, though not as extensive, are known from the Weber For- mation in central Colorado (Fig. 1H). Late Penn- sylvanian plant remains are known from the McCoy Formation in central and northern Col- orado (Fig. 11), and additional fossils have been reported from a locality in central Oregon (Fig. 1J) of uncertain stratigraphic placement within the Pennsylvanian. Manning Canyon Shale. Plant fossils re- covered from the Manning Canyon Shale in northern Utah (Fig. 1G) are well-preserved im- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 prints that show considerable variety of plant form (Tidwell, 1962, 1967; Tidwell et al., 1974). While the strata were once thought to straddle the Mississippian-Pennsylvanian boundary, the flora best correlates with Early Pennsylvanian Namurian B or possibly late Namurian A stages of Europe. The plants are like those of coal swamps to the east. Arborescent lycopods were numerous (Tid- well et al., 1974). Lepidodendron (Fig. 3D), Lep- idophloios, and Si nown to have been present based on stem imprints. In addition, re- mains of the presumed reproductive organs of these plants are represented by Lepidostrobus, Lepidocarpon, and Sigillariostrobus, with Stig- maria (Fig. 3C) representing the underground parts and Lepidophyllum and Cyperites the fo- lia age. Both Archaeocalamites and Calamites (Fig. 3A) were present, as were three species of calamita- lean foliage of the Asterophyllites type (Fig. 3B). A poorly preserved Calamostachys has been found, but details of its construction ane the type of spores it produced remain unknow Ferns and seed ferns are Miei by 11 species of foliage assigned to the genera Adian- tites, Cornypteris, Crossopteris (Fig. 3F), Diploth- mema, Mariopteris, Neuropteris, Rhodea, Sphenopteridium, Sphenopteris (Fig. 3E), and Zeilleria (Fig. 3G). In addition, the pollen organs Aulacotheca and Telangium are present, as well as seeds of Cornucarpus, Holcospermum, La- genospermum, Rigbycarpus, and Trigonocarpus. Cordaitales are represented by three species of foliage, a Cordaianthus (Fig. 3H), and six species of seeds Thus, the Manning Canyon Shale flora rep- resents vegetation similar to that of coal swamps in the midwestern and eastern United States. The setting is interpreted as a coastal swamp com- munity at the margin ofa fresh or brackish water embayment that underwent several transgres- sive-regressive cycles (Tidwell, 1967). Some of the taxa have stratigraphic ranges suggesting a Mississippian age for the flora, but most indicate a Pennsylvanian age correlating with the Na- murian B stage in Weber Formation, C. blado: Scattered plant remains are known from the Weber Formation in Colorado (Fig. 1H), which Read & Mamay (1964) treated as Early Pennsylvanian Zone 6. Lepidodendron johnsonii Arnold (1940) repre- sents the westward extension of typical coal swamp vegetation into central Colorado. The Hd ° ge] 1987] MILLER—LAND PLANTS OF NORTHERN ROCKY MOUNTAINS 697 A! NOR "ATTE, d í FIGURE3. Example Pennsylvanian plant ye from the northern Rocky Mountains. — A. Calamites, x 0.6.— B. Asterophylites x1.0.—C. Stigmaria, x0.5.— vio d aire x0.95.—E. Sphenopteris, x0.7.—F. Cros- sopteris, x 0.6.—G. Zeilleria, x 1.1. —H. Pais naira x0.9.—I. Walchia sp., UMMP 22131, x1.0.—J. Lecrosia wouldii Arnold, UMMP 21005, x1.0. (Figs. A-H from “wm ss courtesy of W. D. Tidwell.) 698 small amount of coal in the Weber Formation suggests, however, that the coal swamp was of limited development and of short duration (Ar- nold, 1940). Read (1933) reported imprints of Trichopitys whitei from the Weber Formation and in 1934 added Lepidostrobus, Stigmaria, Calamites, Asterophyllites, Adiantites, Dactylo- phyllum, Neuropteris, Sphenopteris, Diplomema, Cordaites, and Cordaianthus. McCoy Formation. Several localities within the McCoy Formation in Colorado (Fig. 1I) have yielded imprints of plant remains (Arnold, 1941). Examples are: Calamites gigas Brongn., Odon- topteris mccoyensis Arnold, Cordaites anguloso- striatus Grand'Eury, Samaropsis hesperius Ar- nold, Lecrosia gouldii Arnold (Fig. 3J), Walchia stricta Florin, and Walchia sp. (Fig. 3I; Arnold, 1941). Arnold commented on the arid environ- ment evidenced by the plants and by desiccation features in the rocks. Thus, the fossils may rep- resent forms that grew on drier sites than the coal swamp represented by the plants from the Weber Formation. The McCoy Formation is regarded as Late Pennsylvanian Zone 10 by Read & Ma- may (1964), and the drier setting may correlate with similar events in coal swamps to the east (Phillips et al., 1985) or may reflect some early uplift of the ancestral Rocky Mountains (Pfef- ferkorn & Gillespie, 1980). Spotted Ridge flora, Oregon. The position of this plant megafossil assemblage (Fig. 1J) within the Pennsylvanian has not been determined; none ofthe fossils found to date in this flora are reliable stratigraphic indicators. The flora has the dis- tinction of being the westernmost deposit of Pa- leozoic plants in the conterminous United States and is thought to have grown on an island arc or microcontinent that was some distance west ofthe North American craton (Pfefferkorn & Gil- lespie, 1980). The following taxa have been re- ported: an unnamed lepidodendrid branchlet, Mesocalamites hesperius (Arnold) Mamay & ead, M. crookensis Mamay & Read, Mesocal- amites sp., Phyllotheca paulinensis Mamay & Read, cf. Asterophyllites equisetiformis (Schlot- Schizopteris trichomanoides Goeppert (Arnold, 1953; Mamay & Read, 1956; Read & Merriam, 1940). Of these, Mesocalamites hesperius, Phyl- lotheca paulinensis, Pecopteris oregonensis, and Dicranophyllum rigidum are the most common, with the others represented in the collection by ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 one to a very few specimens (Mamay & Read, 1956). The large size of the collection rules out sampling bias, and the numbers of specimens are thought to reflect plant abundances with reason- able accuracy (Mamay & Read, 1956). Mamay & Read (1956) commented that the plant remains occur in sandstone and mudstone and show little evidence of transport. In fact, some Calamites stems appear to have been pre- served in growth position. Most ofthe sediments are terrestrial in origin or some possibly origi- nated in a marine or brackish water estuary. Thus, the vegetation represented does not appear to be directly comparable with any of the coal swamp communities of the eastern and midwestern United States. While the plants probably grew ina moist habitat, the latter was not a coal swamp. iscussion. In addition to the fossil assem- blages described above, fragmentary plant re- mains are known from subsurface samples in Montana (Pfefferkorn & Gillespie, 1980), and a small collection of fossils has been reported from ee sediments in Wyoming (Sando et , 1975). These indicate that the type of vege- n present in eastern AE AS PUE d land e Penns yivanl surfaces in the HUND northern mi Rocky! Mountain region. Nonetheless, Pfefferkorn & Gillespie (1980) recognized the vegetation of the region as belonging to a distinct floral province, the Cor- dilleran, of which Crossopteris (Fig. 3F) is a characteristic endemic. PERMIAN The Permian was a pivotal period of geologic time from the standpoint of plant evolution and from the types of vegetation that occurred. To- ward the end of the Pennsylvanian, eastern and midwestern coal ta change in com munity structure from domination by arbores- cent lycopods to domination by marattiaceous tree ferns (Phillips et al., 1985). This perhaps foretells of changes to come with breakup of Pan- gaea and the movement of continents poleward during the early Mesozoic. Deposits of Permian age plant remains in the region are few, and none that are known provide the quantity of well-preserved fossils typical of earlier periods. One occurrence of Permian age plant remains has been reported by Mamay & Breed (1970) from the southern periphery of the present northern Rocky Mountains. They re- ported eight specimens from the Cutler Forma- 1987] tion in southern Utah (Fig. 1K). From these are identified Taeniopteris sp., Supaia rigida White, Protoblechnum bradyi Mamay & Breed, and Cal- lipteris sp. This represents the northernmost oc- currence of Supaia. Protoblechnum bradyi is most similar to a species from eastern Asia and sug- gests the possible affinities of the Utah assem- blage with the Asian flora (Mamay & Breed, 1970). Similar remains are found in deposits in Texas, New Mexico, and Arizona, and we can only spec- ulate that exposed land surfaces in the present northern Rocky Mountains supported similar plants. TRIASSIC The only plant fossils reported from the Trias- sic in our region are an Equisetum, two species of Pterophyllum, two of Zamites and one Po- dozamites (Berry, 1924). These come from the so-called “red beds" in westcentral Wyoming (Fig. 1L). These fossils are similar to those found in the Chinle Formation in the Southwest (Ash, 1972; Daugherty, 1941) and suggest that a sim- ilar vegetation occurred in the northern Rocky Mountains. JURASSIC During the Early and Middle Jurassic, the pres- ent northern Rocky Mountain region was cov- ered by an inland sea in which marine limestone and sandstones were deposited (Imlay, 1984; Sil- verman & Harris, 1967). Uplift in the Nevadan orogenic belt south and west of the present north- ern Rocky Mountains resulted in a broad interior basin of low relief that was poorly drained by rivers northward and received sediments from highlands to the southwest and from the Cana- dian shield to the east (Brenner, 1983; Walker, 1974). The large size of the interior basin is re- flected by the widespread deposition of shales and sands of the Late Jurassic Morrison For- mation and its equivalents, which are recognized from southern Utah and Colorado north into southern Canada (Walker, 1974). Geologic studies suggest that a somewhat arid but seasonally wet climate prevailed during the Late Jurassic (Walker, 1974), supporting a sa- vannalike vegetation with many shallow lakes and swamps. There were probably dry periods during which water tables dropped and exposed accumulated debris to oxidation and decay. This MILLER—LAND PLANTS OF NORTHERN ROCKY MOUNTAINS 699 may explain why plant fossils are not more abun- dant in the region (Walker, 1974). Plant fossils are known from scattered locali- ties throughout the region (Delevoryas, 1969; Tidwell, 1975), but large assemblages of remains that reflect vegetation are known only from cen- tral Montana (Fig. 1O) and southern Canada (Fig. | P). South of the northern Rocky Mountain re- gion are a number of occurrences of silicified remains from the Late Jurassic Morrison For- mation (Fig. 1M). Some noteworthy examples are: Hermanophyton kirkbyorum Arnold (1962), a Rhexoxylon-like log, Osmundacaulis wadei Tidwell & Rushforth (1970), the only species of petrified osmundaceous rhizome known from the Jurassic of North America, as well as several different coniferous woods (Medlyn & Tidwell, 1975, 1979). Within the region there are reports of foliage imprints of the two cycadophytes Nilssonia ni- gricollensis Wieland and Zamites arcticus Goep- pert (Knowlton, 1916) and of petrified cycadeoid trunks (Delevoryas, 1960; Ward, 1905), both from Wyoming (Fig. 1N). The most extensive Late Jurassic flora (Table 1) in the northern Rocky Mountains is that pre- served in shales of the Morrison Formation in central Montana (Silverman & Harris, 1967; Brown, 1972). Plant remains have been found at six localities, one near Lewistown and the rest south of Great Falls near the town of Belt (Fig. 10). While three of the six localities in central Mon- tana are close to one another, the most distant sites are over 170 km apart. Zamites arcticus (Fig. 4A), Nilssonia cf. compacta (Fig. 4D), Po- dozamites lanceolatus (Fig. 4G), and Pityophyl- lum lindstromii (Fig. 4F) occur at all six localities; and these taxa probably represent the dominants ofthe regional vegetation. Pagiophyllum sp. (Fig. 4F) and Cladophlebis virginiensis (Fig. 4C) occur at five of the sites and were thus also widespread. Sagenopteris elliptica (Fig. 4E), leaflets of a cay- tonialean seed fern, and Coniopteris hymeno- phylloides (Fig. 4B), foliage of a dicksoniaceous fern, are locally abundant at certain localities. Nowhere are remains of conifers as abundant as those of cycadophytes. Ginkgophytes (Brown, 1975) are present but rare. Thus, the vegetation appears to have been a relatively open one with scattered conifers, more closely spaced cycado- phytes and tree ferns, occasional ginkgophytes, and an understory of ferns. One of the localities clearly represents material [Vor. 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN —— LÀ. | mi P" fm "er m. 7 c= n. k^ * e FIGURE 4. Typical apt pa from the Late Jurassic Morrison Aeae cer —A. ud eel —B. C Mp iý Shap e (Brongn. ard, UMPM 1092, —D. Nilssonia cf. NE (Phill.) seen DUM G Cladophlebis virginiensis a. UMPM 1987] washed in from surrounding areas, and others provide evidence of an extensive swamp and swamp margin community stable enough to re- sult in a thin but mineable seam of low grade coal (Silverman & Harris, 1967). While coal for- mation indicates deposition of plant debris in water having oxygen low enough to retard deg- radation of remains by aerobic bacteria, some areas of the swamp were above water level and supported growth of marchantioid liverworts. A similar vegetation extended northward into Canada (Fig. 1P) where megafossils are pre- served in sediments of the Kootenay Formation (Bell, 1956), and microfossils are known from the Upper Vanguard Formation (Pocock, 1962). However, there is no reason to assume that a given swamp extended over that distance. Rath- er, there were probably many swamps that formed and disappeared over the several million years represented by the Late Morrison Formation and the Kootenay and Upper Vanguard formations in Canada. Thus, it is doubtful that direct cor- relation of these strata is possible. Furthermore, the Canadian flora appears more diverse than that in Montana (Bell, 1956). While some of this apparent diversity is simply more species of the same genera and is probably due to taxonomic splitting, several genera occur in the Canadian flora that are absent in Montana. These are the fern K/ukia, the seed fern Czeka- nowskia, the cycadophytes Ctenis, Pseudoctenis, and Prilophyllum, and the ginkgophyte Baieria (Bell, 1956). There is another Late Jurassic flora close to but outside the northern Rocky Mountain region (Fig. 1Q). Over 60 taxa have been reported from 20 localities in Douglas County, Oregon (Ward, 1905), and essentially no research has been done on this assemblage since it was treated in the Status of the Mesozoic floras of the United States, 2nd paper over 80 years ago. A modern study of these fossils has great potential to advance sub- stantially our knowledge of Jurassic vegetation. EARLY CRETACEOUS The broad interior basin in which sediments of the Late Jurassic Morrison Formation were deposited persisted into the Early Cretaceous. MILLER —LAND PLANTS OF NORTHERN ROCKY MOUNTAINS 701 BLE l. Fossil plants from the Morrison Forma- tion of central Montana Bryophytes Marchantiolites sp. Ferns and fern allies Equisetum laterale Phillips Hausmannia fisheri (Knowlton) Oishi & Yamasita Coniopteris Voies oeil (Brongn.) Seward Adiantites montanensis aah Brown Cladophlebis albenia ies Bel C. heterophylla Fon C. virginiensis ade waq Cycadophytes Nilssonia cf. compacta (Phillips) Bronn. Zamites arcticus Goeppert Cycadolepis spp. Weltrichia sp. Anomozamites Sagenopteris dd Fontaine Ginkgophytes Ginkgoites cascadensis Brown G. pluripartita (Schimper) Seward Conifers Pagiophyllum sp. Podozamites lanceolatus (Lindley & n Braun e lindstromii (Heer) Nathors Pityocladus E sp. Sedimentation was not continuous, however. It ceased for a time and then resumed. This ces- sation resulted in a disconformity inpia by a layer of coarse sandstone which occurs through- out the region (Walker, 1974). The Teone is regarded as the boundary between the Jurassic and the Cretaceous (Brown, 1956). Sometime after the resumption of deposition, plant remains were deposited. In central Mon- tana this was not until what is believed to be the Aptian stage of the Early Cretaceous (LaPasha & Miller, 1984) even though invertebrates in the basal sandstone of the Kootenai Formation in- dicate a Neocomian age for those strata (Walker, The plant-bearing layers of the Kootenai For- — 1093, —E. Sagenopteris elliptica Fontaine, UMPM 1095, x2.7.—F. Pagiophyllum sp. (Pg) and Pityo- prar de seas evi Nathorst (Pi), UMPM 1096, x1.5.—G. Podozamites lanceolatus (Lindley & Hutton) Braun, UMPM 1097, x1 702 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 B d FIGURE 5. Typical plant fossils from the Early apr ep PIER Formation. — A. Klukia canadensis Bell (K) and Sphenopteris latiloba Fontaine (S), UMP 1.6.—B. Sagenopteris williamsii (Newberry) Bell, UMPM 1099, x1.3.—C. Cladophlebis nA Fontaine UMPM 1100, x3.5.—D. Sagenopteris mclearnii 1987] mation represent deposition of sediments and plant remains in poorly drained swamps that formed from infilling of lakes. Evidence suggests that the swamps contained standing water throughout the year. Some areas were above water level as indicated by the occurrence of liverworts, which were probably preserved in place (Brown & Robison, 1974, 1976; LaPasha & Miller, 1984, 1985). However, the water table at these sites was high enough to keep the soil saturated. Certain plants (Table 2) probably grew in the swamps in which their remains were deposited. This is indicated by the occurrence of large, lo- cally abundant remains in low diversity assem- blages in a wide variety of sediment size classes (LaPasha & Miller, 1984). Fossils that fit this category are Marchantiolites, Diettertia, Coniop- teris simplex (Fig. 5F), Acrostichum, Sagenop- teris williamsii (Fig. 5B), and Athrotaxites berryi (Fig. SE). Remains of other plants are generally found as abundant, small fossils occurring in high diversity assemblages and in narrow ranges of sediment size classes, particularly coarse sedi- ments suggesting slight transport. Examples of these are Coniopteris, Cladophlebis (Fig. 5C), Sphenopteris (Fig. 5A), Sagenopteris elliptica, S. mclearnii (Fig. 5D), Ginkgo (Brown, 1975), Ela- tides (Fig. SH), and Elatocladus (Fig. 5G). The same reasoning, combined with the relative scar- city of fossils, suggests that remains of Equise- tum, Klukia (Fig. 5A), Hausmannia, Zamites, and other bennettitaleans were transported some distance, but exactly how far is unknown While certain species are common to both the Morrison flora and the Kootenai flora (Tables 1, 2), the two represent different types of mm The Kootenai flora appears to have been dom inated by conifers, especially Athrotaxites, vdih bennettitaleans relatively rare iler & 84) certain localities, the stratum of vegetation it oc- cupied remains unknow The Kootenai Purus is widespread in Montana and has equivalents in adjacent states, but plant fossils are known in abundance only near Great Falls. Permineralized cycadeoid trunks have been reported from the Burro Canyon For- MILLER — LAND PLANTS OF NORTHERN ROCKY MOUNTAINS 703 ABLE 2. Fossil plants from the Kootenai Forma- tion of central Montana. Bryophytes Ar. REX rf. FU y) Brown & Rob- À ison Megzgeriites montanensis LaPasha & Miller Diettertia montanensis Brown & Robison Sphenopsids Equisetum montanensis LaPasha & Miller E. cascadensis LaPasha & Miller Lycopsids Minerisporites sp. A Minerisporites sp. B Pteropsids Coniopteris keri gr n ) Seward C. simplex (Lindley & Hut Klukia canadensis Bell Hausmannia montanensis LaPasha & Miller Acrostichum longipennis Fonta Cladophlebis constricta Aes C. inclinata Fontaine C. oblongifolia Fontaine C. oerstedii (Heer) Seward C. virginiensis Fontaine papell dining Bell S. latiloba F S. mclearnii m S. JP e Dunker Arcellite. I cascadensis LaPasha & Miller Cycadophytes Sagenopteris elliptica Fontaine i Be S. williamsii aput] Bell Zamites arcticus Goepp Pseudocycas douglasii Linda & Miller Ginkgophytes Ginkgo pluripartita (Schimper) Heer Conifers Athrotaxites berryi Bell Elatides curvifolia (Dunker) Nathorst Elatocladus dunnii Miller & LaPasha . montanensis Miller & LaPa sis Masculostrobus montanensis Miller & LaPasha Conites sp. — Berry, UMPM 1101, x2.3.—E. Athrotaxites berryi Bell, UMPM 894a, x1.8.—F. Coniopteris simplex PU & Hutton) Harris, UMPM 1102, x1.3.— Elatocladus pres Miller & LaPasha, holotype UMPM 914 x 1.7.—H. Elatides curvifolia (Dunker) Nathorst, UMPM 907, 704 mation (Fig. 1U) in Colorado (Brown, 1950). Frenelopsis varians and Tempskya are known from the Cedar Mountain Formation (Fig. 1U) in Utah (Tidwell et al., 1976), although the upper parts of both of these formations are regarded by some as younger than the Kootenai Forma- tion (Ash & Read, 1976; Tschudy et al., 1984). An extensive assemblage of plant remains (Fig. 1S) is known from the Lower Blairmore For- mation of Alberta (Bell, 1956), and the Kootenai flora compares best with this assemblage. The Lower Blairmore flora is of special interest be- cause it contains the earliest convincing flower- ing plant megafossils in the region, leaf imprints identified as Sapindopsis angusta (Heer) Seward & Conway (Bell, 1956). While it is possible that the sites these leaves come from are in fact some- what younger than the Kootenai flora, it is also possible that the leaves indeed represent some of the earliest flowering plant migrants into the region. Another extensive Early Cretaceous assem- blage is known from outside the northern Rocky Mountain region, but it is nonetheless of special interest. Like the Jurassic plants from Douglas County, Oregon, those from the Shasta Series in northern California (Fig. 1T) represent a large and diverse assemblage that has received no fur- ther work since the flora was described over 80 years ago (Ward, 1905). These fossils should be the subject of a modern study. SUMMARY This report deals with the record of terrestrial vegetation in the northern Rocky Mountains prior to the appearance of flowering plants. The span of time involved covers the Early Devonian 385 million years ago to the Early Cretaceous about inal and a second collecting site in the Early De- vonian Beartooth Butte Formation of northern Wyoming has added considerably to the forms known. Represented are rhyniophytes, trimero- phytes, zosterophylls, and early lycopsids. These compare well with plants of similar age known from eastern North America and western Eu- rope, making the Beartooth Butte assemblage one of the better known Early Devonian floras of the world. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Several deposits of Late Mississippian and Early Pennsylvanian fossils show that vegetation like that of coal swamps of Appalachia and the North American interior occurred in the north- ern Rocky Mountain region as well. The latter vegetation was composed of arborescent lycop- sids, arborescent sphenopsids, tree ferns, seed ferns, and cordaites like that of the Interior-Ap- palachian Province to the east, but the presence of unique forms is grounds for treating the west- ern vegetation in the distinct Cordilleran Prov- ince. Similarly, because of species unique to it, the Spotted Ridge flora of Oregon is treated as representing the Oregonian Province. ur record of vegetation is spotty in sediments of the Permian, Triassic, and Early and Middle Jurassic in the northern Rocky Mountains. By comparison, evidence of Late Jurassic and Early Cretaceous vegetation is relatively abundant and informative. Much of the present northern Rocky Mountains was then a broad interior basin of low relief that supported an open savanna type of vegetation interspersed with shallow lakes, swamps, and rivers. During the Late Jurassic co- nifers appear to have been widely scattered, cy- cadophytes and ferns were more closely spaced, and ginkgophytes and caytonialean seed ferns were more rare and occasional. This vegetation were equable, and there is no evidence of freezing conditions. While the interior basin persisted into the Ear- ly Cretaceous and was the site of deposition of plant fossils, the vegetation represented had an entirely different aspect. Conifers were abundant and dominated the vegetation, with cycado- phytes rare and widely scattered. Ferns formed the understory and caytonialean seed ferns were locally abundant. Rainfall and temperatures were like those of the Late Jurassic. There is no evidence of flowering plants in the flora of the Early Cretaceous Kootenai Forma- tion in Montana, which is regarded as Aptian in age. However, imprints of flowering plant leaves are present but rare in sediments of presumably leaves are somewhat younger than Aptian, as some believe, these imprints represent some of the first flow- ering plants in the northern Rocky Mountain 3 * = © aN ° B oO 3 Lond n [e] ° 5 e ie 3 = ge [ad = [4] Thus, de their in the northern Rocky Mountains toward the end 1987] of the Early Cretaceous, and it is important to know about the vegetation that occurred on the land as they migrated into the region. Unlike problems confronting the first land plants in the Early Devonian that colonized relatively barren land, the establishment of flowering plants re- quired their successful competition with vege- tation that was already in place. This is indeed a remarkable feat; however, this survey shows that replacement of one form of vegetation by another has been the rule throughout the 275 million years discussed in this paper. LITERATURE CITED ARNOLD, C. A. 1940. Lepidodendron johnsonii, sp. nov. from the Lower Pennsylvanian of central Col- orado. Contr. Univ. Michigan Mus. Paleontol. 6: -52. 1941. Some Paleozoic plants. from central Contr. Univ. Michigan Mus. Paleontol. 6: 59-7 . 1953. Fossil plants of Early Pennsylvanian 1 Oregon g Abt B, Paliophytol 93: 61 —68. ————. 1962. A Rhexoxylon-like stem from the Mor- rison Formation of Utah. Amer. J. Bot. 49: 883- 88 W. SADLICK. 1962. A Mississippian flora from northeastern Utah and its faunal and strati- graphic e Contr. Univ. Michigan Mus. Pa- leontol. 17: AsH, S. R. 1972. Plant megafossils of the Chinle For- mation. C. S. Breed Breed (editors), In- vestigations in ihe Triassic Chinle Formation. Mus N. Arizona Bull. 47: 23-44 & C. B. READ. 1976. North American species of Tempskya and their stratigraphic Significance. U.S. Geol. Surv. Profess. Pap. 874: . R. Sc COTESE & A. M. ZIEGLER. of the Pa- BAMBACH, R. K., 1980. Before Pangaea: i phi leozoic world. Amer. Sci. 68: 26-38. 1956. Lower Cretaceous floras of western 83. Late Jurassic roig setting and paleogeography of Western Interio .119- 132 in M. W. Reynolds & E. D. Do olly (editors) nited BRowN, J. T. 1972. mation (Upper Jurassic) of Central Montana. Ph.D. Dissertation. University of Montana, Missoula, Montana. 9 Upper Jurassic and Lower Cretaceous ginkgophytes from Montana. J. Paleontol. 49: 724- 7 R. Rosison. 1974. Diettertia monta- tana. Bot. Gaz. (Crawfordsville) 135: 170-173. MILLER —LAND PLANTS OF NORTHERN ROCKY MOUNTAINS 705 976. Observations on the structure of posee cde blairmorensis (Berry) n. comb. from the Lower Cretaceous of Montana, U.S.A. J. Paleontol. 50: 309-311 BRowN, R. W. 1950. Cretaceous plants from south- western Colorado. U.S. Geol. Surv. Profess. Pap. 221-D: 45-66. Fossil plants and the Jurassic-Creta- ontana and Alberta. Amer. e Upper Triassic flora of Arizona. Publ. Poe Inst. Wash. 526: 1- 108. DELEVORYAS, T. 1960. Investigations of North Amer- ican cycadeoids: trunks from Wyoming. Amer. J. Bot. 47: —786. . 1969. Biotic provinces and the dees Sieg taceous floral transition. Proc. N. Am r. Paleon- tol. Conv. II: 1660-1674 Dorr, E. The occurrence of the oldest known rom Beartooth Butte, Wy- i ie M po oming. Bot. Gaz. (Crawfordsville) 95: 240-256. s ped Devonian flora from Beartooth Butte, Wyoming. Bull. Geol. Soc. Amer. 45: 425- 440. . 1934b. Stratigraphy and paleontology of a new Devonian formation at Beartooth Butte, Wy- oming. J. Geol. 42: 720-737. GENSEL, P. G. & H. N. ANDREws. 1984. Plant Life in the Devonian. Praeger Publishers, New York 70. Rebuchia: a new name for Bu- Rebuchia ovata, its vegetative mor- phology and classification with the Zosterophyl- lo dew Rev. Palaeobot. Palynol. 14: 113-127. 1984 IMLAY, R. I Jurassic ammonite successions in e Pp. 1-12 inG. E.G. Westerman (editor), Jurassic- North America. Geol. Assoc. Canada Spec. Pap. 27. KNOWLTON, F. H. 1916. Note on a recent discovery of fossil plants i in the oo on Formation. J. Wash. Acad. Sci n 180- KOSANKE, R. 50. I nsylvanian spores and their use in ipie: LAE, Illinois State Geol. Surv. : 1-128. . N. MILLER. 1984. Flora of the Early Cretaceous. Kootenai ee = Mon- paleoecolo grap t. B, Pa- aca 194 109-1 30. & 985. Flora of the Early Cretaceous Kootenai | in Montana, bryophytes and tracheophytes excluding conifers. Palaeonto- graphica, Abt. B, Paláophytol. 196: 111-145. MAMAY . & N. J. BREED. 1970. Bp Permian plants from the Cutler Formation in Monument Valley, Utah. U.S. Geol. Surv. Profess. Pap. 700- i —117. EAD. 1956. Additions to the flora ofthe Spotted Ridge Formation in central Oregon. . Geol. Surv. Profess. Pap. 274-I: 211-226. MEDLYN, D. A. & W. D. TIpwELL. 1975. Conifer 706 wood from the Upper Jurassic of Utah Part I: Xenoxylon morrisonense sp. nov. Amer. J. Bot. : 203-208. 979. A review of the genus Pro- topiceoxylon with emphasis on North American anad. J. Bot. 57: 1451-1463. MELTON, W. G. 1971. The Bear Gulch fauna from N. TAYLOR. 1979. Paleozoic seed fern pollen organs. Bot. Rev. (Lancaster) 45: 301-375. MILLER, C. N. & C. A. LAPASHA. 1983. Structure and affinities of Athrotaxites berryi Bell, an eles Cre- taceous conifer. Amer. J. Bot. 70: 772-77 1984. Flora of the Early Cretaceous Ko MEET Formation i in Montana, conifers. Palae- ontographica, Abt. B, Paláophytol. 193: PFEFFERKORN, H. F. & W. H. GILLESPIE stratigraphy and biogeography of plant compres- sio n fossils in the Penn LP ie of North Amer- a. Pp. 93-118 in D er & T. N. Taylor esie fee itd d rue Plants. Dow- on & Ross, Inc., Stroudsburg, Penn- . A. PEPPERS & W. A. DI MICHELE. C. B. Cecil (editors), ` Paleoclimatic Controls on Coal Resources of the Pennsylvanian System of North America. Int. J. Coal Geol. 5. Pocock, S. A. J. 1962. Microfloral analysis and age determination of strata at the J ern uo S s. Palae- —95. o READ, boniferous of Colorado. J. Wash. Acad. Sci. 23: 461—463. . 1934. A flora of Pottsville age from the Mos- quito pena Colorado. U.S. Geol. Surv. Profess. Pap. 185D: 7 1947. Pannevivaiian floral zones and floral provinces. J. Geol. 55: 271-279. . MAMAY. 1964. Upper Paleozoic floral zones and floral provinces of the E States. U.S. Geol. Surv. Profess. Pap. 454K: . MERRIAM. 1940. A Uer NN flora from central Oregon. Amer. J. Sci. 238: 107- 111. SANDO, W. J., G. MACKENZIE, JR. & J. T. DUTRO. 1975. Stratigraphic and geologic history of the Amsden Formation (Mississippian and Pennsyl- ary of NE g. U.S. Geol. Surv. Profess. p. 848-A: 1-83. A M. P. 1950. Carboniferous plant spores from Daggett County, Utah. J. Paleontol. 24: 232- SILVERMAN, A. J. & W. L. HARRIS. 1967. ag mid and economic geology of the Great Falls-Lewis ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 town coal field, central Montana. Montana Bureau of Mines and Lowe y, Bull. 56: 1- TANNER, W. R. new species of Gosslingia d ME from the Lower Devonian artooth Butte Formation of northern Wyoming. 3rd N. Amer. Paleontol. Conf. Proc. 2: 541-546. . 1983. A Fossil Flora from the Beartooth es Formation of Northern Wyoming. Ph.D. D n Southern Illinois University, en Illino TAYLOR, T. N. 1981. Paleobotany, an Introduction to Fossil Plant Biology. McGraw-Hill, Inc., New York ork. dige. W. D. 1962. An Early Pennsylvanian flora m the Manning Canyon Shale, Utah. Brigham Young Univ. Geol. Stud. 9: 83- 102. 1967. Flora of Manning Canyon Shale, Part I:a | Lowermost Pennsylvanian flora from the Man- ning Canyon Shale, Utah, and its stratigraphic " nificance. Brigham Young Univ. Geol. Stud. 14: 3-66. l Common Fossil Plants of Western North America. Brigham Young Univ. Press, Pro- vo, Utah. — & J. R. JENNINGS. 1986. Stansburya petersenii gen. et sp. nov., an anatomically preserved lyco- pod from the Deseret Limestone (Mississippian) of Utah. Palaeontographica, Abt. B, Paláophytol. 198: 1-11. S. R. RUSHFORTH. l 970. “aasan wadei, a rison Formabon (Jurassic) of Utah. Bull i umaq Bot. Club 97: 137-144. -c D. . MEDLYN & A. D. SiMPER. 1974. Flora of the Manning Canyon Shale, Part II: Lepido- dendrales. Brigham Young Univ. Geol. Stud. 21: 119-146. N & J. L. ROTH. 1976. Cretaceous and Early Tertiary floras of the eire: area Brigham Young Univ. Geol. Stud. 22: 7 TscHuDY, R. H., . TSCHUDY & L. C. d m 1984. Palynological evaluation of Cedar Mountain and Burro Canyon Formations, Colorado Plateau. U.S. Geol. Surv. Profess. Pap. 128I: 1-24. WALKER, T. F. 1974. Stratigraphy and Depositional Environments of the Morrison and Kootenai For- mations in the Great Falls Area, Central Montana. Ph.D. inm University of Montana, Mis- soula, Monta WARD, L. F. 1905. Status of the Mesozoic floras of the United States, 2nd paper. U.S. Geol. Surv. 6. R. K. BAMBACH, J. T. PARRISH, S. F. RRETT, E. H. GIERLOWSKI, W. C. PARKER, A. RE: J. SEPKOSKI. 1981. Paleozoic bio- and climatology. Pp. 231-266 in K. Niklas (editor), Paleobotany, Paleoecology, and Evolution, Volume II. Praeger Publishers, New York. ANGIOSPERMS OF THE NORTHERN ROCKY MOUNTAINS: ALBIAN TO CAMPANIAN (CRETACEOUS) MEGAFOSSIL FLORAS'! DAVID R. CRABTREE? ABSTRACT Synchronous first occurrences of pollen and leaves indicate that angiosperms entered the Northern Rocky Mountain (NRM) region during the Middle Albian, approximately 8 Ma later than such shifts in a from southern Laurasia. The earliest angiosperm pollen and leaf flora Mid s to the P ac sub-Zone IIB in that it poda a pretricolporate palynoflora and a comparable grade of e ee based on leaf rank. The Albia un tanophyll, Protophyll, and Pentalobaphyll mo rphotypes. Sapindophylls are common an flora contains early North American occurrences of the widespread Upper Cretaceous leaf form genera Trochodendroides and Cineman Pentalobaphylls (Araliaephylls) appear early and assume a numerical dominants. Pinnate palms are present by the early Campanian. Higher-level taxa present in the ji cdi in the Ear ly y Campanian include Magnoliales, Laurales, Chloranthales, Nymphaeales, Menisperm Platanaceae, H C agales, Rosidae and sine Dilleniidae. Mountain (NRM) region based on the megafloral record. The refinement of descriptive terminol- ogy for leaf architecture (Hickey, 979; Dilcher, 1974) and elucidation of the phyloge- netic significance of leaf morphology (Hickey & Wolfe, 1975), along with the U.S. National In 1874 Leo Lesquereux stated: he plants of the Dakota group, as known mostl of our time ... and the evident likeness of their : . facies . . . strikes the paleontologist and may lead Cleared Leaf Collection and the U.S. Geological him into error... for, really, when we enter into Survey Cleared Leaf Collection, have provided the framework for the present stu a more detailed analysis of these Cretaceous leaves, y. The rich Cretaceous leaf fossil deposits of the we are by and by forcibly impressed by t strangeness of the characters ... which seem at riance With any of those recognized anywhere western interior of North America have long been in the floras this flora recognized for their importance to angiosper paleobotany. of o does not leave see pew ay tie E to my ind. (during the late 1800s and early 1900s) mistak- In this paper I summarize and evaluate the evidence relevant to the paleoecology and evo- lution ofearly angiosperms in the Northern Rocky enly placed many leaf species into Recent genera. Even investigators who realized their predica- ment (see Lesquereux quote above) were ham- !T xc L. J. Hickey, J. U. McClammer, J. A. Wolfe, S. L. Wing, G. R. Upchurch, L. S. Pierce, B. H. Tiffne and K. R. Johnson for their interest in and Pss of early angiosperms. C. N. Miller provided ie. ic for the photography and technical detail, as well as much encouragement. The Department of Botany, University of Montana, funded the preparation of parts of the manuscript. NSF Doctoral Dissertation Improvement Grant BSR 8400303 funded visits to museums. F. Heuber, S. Wing, and J. Ferrigno made the collections of the U.S. National Museum available for study. T. Bolton assisted me at the Geological Survey of Canada, Ottawa, and I. Birker helped at the Redpath Museum, Montreal. C. Blount, Department of Geology, Idaho State University, loaned me a collection of fossils from the Wayan Formation. C. A. LaPasha made many of the thin sections of wood from the Vaughn Member. I thank H. Chambers for assistance with Tables 1, 2, and 3 and M. Crabtree for assistance with some of the figures. Fieldwork i in 1982 was supported in part by a Sigma Xi Grants-in-Aid of Research to the Author ? Department of Botany, University of Montana, Missoula, Montana 59812, U.S.A. ANN. Missouni Bor. GARD. 74: 707-747. 1987. 708 pered in their attempts to interpret these leaves by the absence of appropriate operational para- digms. Throughout this report, genera of doubt- ful occurrence in the Cretaceous are marked with quotation marks. Where possible I have provid- ed reappraisals of the affinities of these leaves in my discussions of Cretaceous leaf morphotypes and individual floras. I cover the period of about 30 million years from the apparent advent of angiosperms during the Middle Albian to the middle of the Cam- panian. This includes the latter part of stage one of angiosperm evolution (Vakhrameyev, 1982), during which the group became established in restricted areas of a predominantly gymno- sperm-dominated vegetation; and it includes most of stage two, in which angiosperms radiated explosively and displaced gymnosperms from many parts of the wor The floras reported here come from British Columbia, Washington, Alberta, Montana, Wy- oming, and Idaho (Figs. 1, 2). Coniferous communities were the dominant form of vegetation in the world, at least through the Early Cretaceous (Hughes, 1969, 1976; Kras- silov, 1981; Miller, 1977; Penny, 1969). Lower Cretaceous conifers from the NRM region are regarded as belonging to the Taxodiaceae, Ar- aucariaceae, Podocarpaceae, Pinaceae, and Chei- rolepidiaceae (Bell, 1956; Miller & LaPasha, 1984; Singh, 1964, 1971). With the exception of the Cheirolepidiaceae, which declined during the Albian, and the addition of the Cupressaceae which appear to have diversified during the mid- dle and Late Cretaceous, these groups form the nucleus for the development in the Late Creta- ceous of the North Pacific refugium (Vakhra- meyev, 1982). Compressions and impressions of foliage and cones of Taxodiaceae, Cupressaceae, and Araucariaceae occur abundantly in coarser sediments throughout the region in the Upper Cretaceous (Berry, 1929a; Knowlton, 1905). Ginkgophytes decrease in the region during the Albian, the latest flora in which they form an important component being the Lower Blair- more (Dawson, 1886; Bell, 1956), although fo- liage is reported from the Cenomanian Dunve- gan Formation in northern Alberta (Bell, 1963), and Salisburia (cf. Ginkgo) seeds are reported as late as the Campanian Belly River flora (Dawson, 1886). The Ginkgophyte decline is delayed in the extreme northern latitudes. Although northslope Alaskan floras exhibit a notable decline in the Late Albian (Scott & Smiley, 1979), the group ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 remains important at least into the early Se- nonian in the Atane beds of Greenland (Seward & Conway, 1935), and into the Campanian in the Chignik Formation of central Alaska (Hollick & Martin, 1930). Ginkgo persists into the Paleo- gene in the Ft. Union Formation of Montana and the Willwood Formation of Wyoming (pers. obs.). Fern communities of the Recent families Glei- cheniaceae, Schizaeaceae, Dicksoniaceae, and mundaceae, and the extinct family Temp- skyaceae occur abundantly in certain facies, a characteristic feature of the middle Cretaceous that Krassilov (1981) interpreted as evidence for extensive fern marshes. Pteridophytic commu- nities colonized the widespread upper Albian and Cenomanian volcanic ash flats, occasionally being preserved in situ as in the Albino Member of the Mowry Shale in southwestern Montana (Crab- tree, 1983; Vuke, 1982). Ferns such as Glei- chenia, Anemia, Sphenopteris, Cladophlebis, Tempskya, Coniopteris, and Onychiopsis (but see Skog, 1985) are found throughout the region in the Albian and Cenomanian (Andrews, 1948; Andrews & Kern, 1947; Andrews & Pearsall, 1941: Ash & Read, 1976; Bell, 1956; Knowlton, 1917; Read & Brown, 1937; Seward, 1924). Post- Cenomanian floras show a decreasing represen- tation of these genera, concomitant with increas- ing Polypodiaceae s.l. (pers. obs.). Cycadophytes such as Nilssonia, Zamites, Pseudoctenis, Ctenis, and Otozamites decline rapidly after the Aptian. However, certain species persist in the Cenomanian Dunvegan flora (Bell, 1963) and the Turonian upper Frontier flora (Berry, 1929d), and Zamites albertensis Berry is abundant in the Campanian Allison flora of Al- berta (Berry, 19292). Cycadophytes also persist in the Late Cretaceous of Alaska (J. Wolfe, pers. omm.). Sagenopteris, foliage of pteridosperms of the Caytoniales, is abundant during the Aptian and early Albian in the region (Bell, 1956; LaPasha & Miller, 1985) but is only rarely reported from the later Albian or from the Upper Cretaceous (see Winthrop flora in this report). The relative abundance of major plant groups, and the entry of angiosperms into the region dur- ing the Middle and Upper Albian, can be seen in the histograms depicting the changing com- position of the flora during the Middle and Up- per Albian (Fig. 3). One histogram (Fig. 3A) is based on published megafossil floras and my own observations of unpublished floras. The other e 1987] CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 709 x = ° % = - 60°N BRITISH COLUMBIA ALBERTA A , O D «on MONTANA WASHINGTON 459N 250 km Qn WYOMING IGURE l. Geographic locations of plant megafossil collections in the northern Rocky Mountain region. Collection sites are numbered in alphabetical sequence. Names of sites appear in Figure 2. Appendix I contains additional information on locations. (Fig. 3B) is based on published palynofloras. Per- centages shown in the histograms were calculated by counting species of each major plant group within individual floras. These raw count totals were summed and a percentage calculated for each chronostratigraphic subdivision of the AI- bian. Megafossil floras show a bias towards pres- ervation of vegetation adjacent to the deposi- 710 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 o NT oF u PALEOBOTANICAL ON < STAGE COLLECTION a 73 = " 1 ALBINO c 2 ASPEN - 3 BADHEART P (13) 4 BEAVER MINES ae. 5 BLACK EAGLE I 6 COMMOTIO _ A 6a LOWER PART ©) ae MEMBER B (21) 6b UPPER PART wp (15) GATES MEMBER GS) 6c BOULDER CREEK — SANTONIAN MEMBER (3) 7 CROWSNEST 875 — 8 DUNVEGAN 885 CQNIACIAN 9 AGLE = © 10 FALL RIVER — TURONIAN 11 FRONTIER — lla LOWER PART = UR FRONTIER _ M FORMATION mo — M 11b UPPER PART 4 FRONTIER - — i ORMAT I on (23) (2) 12 JACKASS MOUNTAIN = 13 JUDITH RIVER - "c — C1) (22)? LOMA = UPPER (10) 14 KINGSVALE -—— 15 MILK RIVER E 16 MILL CREEK _ A DEK Q9) 17 PASAYTEN "B s. 18 — PRE-MUDDY WE. - D (16) 19 SUMMIT ín (c) 20 SUN RIVER E AOOO) 21 TWO MEDICINE _ " (Ba) Q 22 + WAYAN 7 23 WINTHROP LOWER 13 — FiGuRE 2. Chronostratigraphic positions of plant Wicow collections in the sels Rocky Mountain region. Stage boundary dates according to Harland et al. (1982). Hickey & Doyle (1977). chy aan Voie of aida is bow on my in weste h American chronostra Substage boundaries are base nterpretation of ye otomac Zones from terpretation of the literature. tigraphy. Collections Ages are in million of uncertain age are indicated n denis marks (see text ud rises I for discussion). tionalenvironment; thus Figure 3A approximates the components of the lowland vegetation. Pal- ynofloras used in Figure 3B were recovered most- ly from marine and marginal marine environ- ments, and thus more closely reflect the regional flora. From Figure 3A it is apparent that dra- matic changes occurred in the lowland flora dur- ing the Middle and Upper Albian. In these en- vironments angiosperms increased at the expense of ferns. Because this trend is not evident in Fig- ure 3B, it is apparent that this early angiosperm radiation had its greatest impact on lowland vegetation. PALEOGEOGRAPHIC AND GEOLOGIC SETTING The northern Rocky Mountain region during most of the Cretaceous occupied a strip of land some several hundred kilometers wide, situated between the Pacific geosyncline/island arc to the west and the mid-continent epeiric sea to the east (McGookey et al., 1972; Williams & Stelck, 1975). Paleolatitudes may have varied up to ten degrees from the present range of 41?N in southern Wy- oming to 60°N in northern Alberta and British Colombia. The paleocontinent positions of Smith et al. (1981) indicate that northern Alberta has 1987] rotated 8—10° to the south since the Cretaceous, and southern parts of the region are presently approximately 2—5? farther south. The first major expansion of the Cretaceous epeiric sea began in the Late Aptian and contin- ued into the Albian, attaining its maximum ex- tent in the late Middle Albian when it reached unbroken from the Arctic Ocean to the Gulf of Mexico (McGookey et al., 1972; Vuke, 1984; Williams & Stelck, 1975). The fluctuations of the sea initiated a famous series of transgressive— regressive clastic cycles of deposition that con- tinued until the end of the Cretaceous (Waage, 1975). Geological formations in the region tend to parallel the north-south trending western shoreline of the seaway. Cretaceous sediments accumulated in terrestrial and marine environ- ments to great thicknesses: up to 3,500 m in southwest Wyoming (Rubey et al., 1975) and 2,134 m in northwest Montana (Rice & Cobban, 1977). The source areas in present British Co- lumbia, western Alberta, Idaho, Washington, Oregon, western Montana, and western Wyo- ming are inferred to have had considerable relief n order to account for the great thickness of accumulated sediments. Orogenic activity in the region was of long duration beginning in the Late Jurassic and Early Cretaceous with the uplift of the Nevadan Orogeny. The main thrusting of the Sevier Orogeny occurred throughout the Creta- ceous, and the uplift of the Laramide Orogeny took place during and subsequent to the Cam- panian (Gilluly, 1963; Nichols et al., 1985). PALEOCLIMATE Generalized interpretations of paleoclimate, which include the North American Cretaceous, have been published by Frakes (1979), Habicht (1979), Lamb (1977), Schwarzbach (1974), and Vakhrameyev (1978). They recognized a general humidification in Laurasian climate from the Ju- rassic into the Cretaceous. This can be correlated with the widespread inundation of continental crust, a worldwide phenomenon in the Creta- ceous, and the opening of the Atlantic Ocean. Assessment of precipitation patterns and amounts during the Cretaceous in the NRM re- gion is difficult with our present knowledge. Par- rish et al. (1982) mapped the Cretaceous precip- itation in the region as low to moderately low largely on account of the presumed orographic effect of the Rocky Mountains. However, they noted that the presence of important high lati- tude coals in the region, especially in the Late CRABTREE— NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 711 Cretaceous, may argue for heavy seasonal pre- cipitation, perhaps a monsoonal effect Growth rings in fossil wood can be used to assess environmental influences on plant de- velopment (Creber, 1977; Creber & Chaloner, 1984, 1985; Fritts, 1976). Presence and degree of development of latewood and growth rings is generally correlated with available moisture. However, there are many examples of growth rings in trees living in regions of high rainfall. For instance, Agathis from Fiji, where annual rainfall is 2,000-6,000 mm, have well-developed growth rings (Ash, 1985). Nonetheless, I know of no exception to the — of growth rings in the wood of trees gro climates, and conversely, the absence of growth rings al- pt per- haps in certain warm-climate eun commu- nities). I examined 27 specimens of six species of co- niferous woods from the Late Albian Vaughn Member of the Blackleaf Formation in northwest Montana in thin section for the presence and configuration of growth rings. Transverse sec- tions could be grouped into three broad cate- gories: Category A. Woodsexhibiting no growth rings. Category B. Woods exhibiting broad (3-5 mm) and consistent growth rings with nsiderable latewood. Woods exhibiting rings of variable width, including broad rings (up to m), pseudorings, and consid- erable latewood. seacnnyal Category C. The Vaughn Member has been interpreted as a deltaic swamp deposit (Cannon, 1966) and was situated at about 55°N paleolatitude (Couillard & Irving, 1975). Cold-seasonality did not exist during the Albian at this paleolatitude. Season- ality as the result of low light during winter months may be a factor at 50°N, although I know of no literature on the subject. Broad-leaved ev- ergreen forests are known to have extended to 65°N during the Cretaceous (Wolfe & Upchurch, 1986). Incomplete seasonal leaf-drop, formation of latewood, and cambial quiescence might well e ] with the northern extensions of this forest. Among several possible explanations for the mixed assemblage of woods, the following is suggested as the most plausible: the trees were growing in a warm, seasonally dry climate, with some species (Category A) living in areas of year- round groundwater. The pronounced develop- 712 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 $ 4 «e ev? «e ` P o0 Y "d "Y e «e e r of e N eo ue ot Nt ANTHOPHYTES ce «e "d CONIFEROPHYTES lower pp Albian m ° © = a & LJ E Lun Lower 0 10 20 30 WA Albian % Species e Cd é s Fà # z? ° o o ge ` Pteridophytes ¥ ? "à Coniferales upper CM Upper Albian lower Upper Albian 104 upper Middle Albian 106| middle Middle Albian 108 lower Middle and Lower Albian NO DES N as N ` * non-angiospermous 9 10 20 30 % species FIGURE 3. Relative abundance of major plant groups during the Albian of pe d Rocky Mountain region. A in million years before present rcentage occurrence of major plant groups based on megafossils. Occurrences of species for each plant group were tabulated for individual ne These counts were then sum a percentage calculated for each time interval as list elow. Total species counted for each interval are 1 e parentheses. Aptian and Lower Albian (119): Kootenai (Miller & LaPasha, 1984 indicated in ntheses. : : LaPasha & Miller, 1985); Bullhead, Gething, and Luscar (Bell, 1956). Lower Middle Albian (150): Beaver Mines (Berry, 1929b; Bell, 1956; Mellon, 1967); Lower Gates Member (Stott, 1963). Middle Middle Albian (130): Upper Gates Member (Mellon et al., 1963); Pasayten (Penhallow, 1907; Bell, 1956); Kingsvale and Jackass Mountain (Bell, 1956); Fall River and i bw aep Obs.); rd Creek (Bell, 1956; Stott, 1963). Upper Middle Albian (150): Mill Creek (Dawson, 1886; B , 1929c; Bell, 1956; Mellon, 1967). Lower Upper Albian (89): Crowsnest (Bell, 1956; Mellon, 1967) ur "Su n River, and Black Eagle (pers. obs.). B. Percentage oc- currence of major plant groups based on microfossils. Percentages calculated as above. Assignment of spores 1987] ment of latewood in Categories B and C suggests seasonality in precipitation. An alternative ex- planation is that low light availability in winter induced latewood formation in some species but not others. Late Albian or Cenomanian woods of Cu- pressinoxylon sp. (Andrews & Kern, 1947) and Paraphyllanthroxylon idahoense Spackman (1948) from the Wayan Formation in south- eastern Idaho have no prominent growth rings and can be seen as evidence for equable year- round temperatures. These woods suggest either the absence of seasonality in precipitation or that the trees grew in swampy or other well-watered habitats. Later in the Cretaceous there is further evi- dence for seasonal or low precipitation in the region, possibly related to the mountain building of the Laramide Orogeny. The Campanian Two Medicine Formation in northwestern Montana includes caliches, desiccated carbonate nodules, and sandstone bodies of episodic rivers that in- dicate a seasonal wet-dry climate, with the dry season longer (Lorenz, 1981). The abundance of large (mesophyll and megaphyll size), apparently deciduous leaves in the Two Medicine flora sup- ports such a climatic seasonality, although the leaves can be interpreted alternatively as indic- ative of successional communities. Low or sea- sonal precipitation is further indicated by a sub- stantial notophyllous evergreen component of coriaceous leaves typically without drip tips. Co- niferous woods from the lower Two Medicine exhibit pronounced growth rings of highly vari- able thickness (Crabtree, pers. obs). Oxygen isotope ratios are used most accurately to estimate maximum paleotemperature and to establish temperature trends of ocean waters (Frakes, 1979). Oceanic temperatures can be used to estimate temperatures on nearby land masses. Since the NRM region during much of the Cre- taceous was a relatively narrow land mass po- sitioned between the Pacific Ocean on the west and the epeiric seaway on the east, it is likely that oceanic temperatures are significant for ap- proximation of the land temperatures. Isotopic ratios from the continental platform CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 713 off the Soviet Union (Douglas & Savin, 1975) indicate a Cretaceous temperature maximum in the Albian, followed by cooling in the Late AI- bian and Cenomanian. Subsequently, a rewarm- ing occurred in the early Senonian before a Maas- trichtian cooling (but see also Boersma, 1984). The Albian maximum is probably the warmest period that the world has experienced since Pa- leozoic time (Frakes, 1979). Latitudinal gradi- ents in surface water temperatures were less steep during the Cretaceous, and the average temper- atures of deep water appear to have been signif- icantly higher than at present (Schopf, 1980). Mid and Late Jurassic oxygen isotope ratios from Montana and Wyoming indicate es the surface water had a maximum of 20°C ( n & Shaw, 1977). Polar ice was unknown das the Cre- taceous, and Frakes (1979) hypothesized that north polar surface water was no cooler than 7— 19°C and may have been at the high end of this range based on isotopic paleotemperatures of about 15?C for putative North Pacific Deep Water from the equatorial Pacific (Stevens, 1971). A computerized model of surface water tempera- tures for an ice-free Arctic predicts a temperature increase of 7-10?C over the present 0—-5°C tem- perature range (Sellers, 1969) Hermatypic coral reefs occur in tropical and subtropical oceans. Because modern reefs form only when minimum water temperatures exceed 18°C, their fossil distribution can be used to es- Mesozoic distributions of reefs (Newell, Beauvais, 1973) indicate that oceanic currents in temperate latitudes were significantly warmer than at present. A reef from the Jurassic of Wy- oming (Beauvais, 1973) indicates that they formed as far north as 50?N paleolatitude, but there is doubt that the Wyoming reef is herma- Jurassic and Cretaceous at latitudes 10-20? north of their present distribution. Several papers discuss paleofloristic climatic zonation during the Cretaceous based on the composition of eastern Asian megafloras (Kras- silov, 1973a, 1975, 1978) and microfloras — and pollen to major plant groups follows the Wisi pans ag E of Singh (1971) and Norris (1967). Numbers in mber of s parentheses after age below indicate total num species counted for that interval. Lower Albian and lower Middle Albian (179): MeMurray-Clearwater(Vagvolgy & Hills 1969); Loon River (Singh, 1971). Middle Middle Albian (206): Harmon (Singh, 1971); per Grand Rap (Singh, 1971); Joli Fou (Norris, 1967). L s (Norris, 1967). Upper Middle Albian (200): Cadotte ower Upper Albian (184): Paddy rer 1971); Viking (Norris, 1967). ). Upper Upper Albian (213): Shaftsbury (Singh, 1971); Upper shale (Norris, 1 714 "c innamomo: phyll DD kJ x Thy Aud ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Cornophyll A Rhamnophyll | B FIGURE 4. Cretaceous dicotyledonous leaf morphotypes. Sources of previously published drawings are in- dicated. SAPINDOPHYLL M erui obtusilobum Mrs eite Format on) Bell.— 12, figs. 6, 7 in Lesquereux, 1892).— Heer, Raritan Formation, Marylan medium Newberry (pl. 29, fig. 7 in Hollick, 1906). (Chlonova, 1980). Krassilov's papers propose four intuitively based, latitudinal climatic zones: boreal, temperate, warm-temperate, and sub- tropical. Relative to these zones, the NRM re- gion is closest to the warm- temperate zone based the presence of palms and large-leaved deciduous dicots. A position in the warm-temperate zone is consistent with Campanian floras in Sakhalin, Amur, Altai, western Greenland, and Vancouver Island (Krassilov, 1981). Vakhrameyev (1978) placed the NRM region straddling the boundary between hist te-warm | hu- mid zones for the Early Cretaceous. This place- “Ficus” beckwithii Lesquereux (Crabtree), Dakota Formation, Golden, (Peabody Museum Poleibolanite! Collection no. oe —B Virginia (fig. 17 in Doyle & Hickey, us — Sapindopsis magnifolia Fontaine, Patapsco For- y "o t : ] B. Generalized leaf. CORNOPHYLL: “C ornus" forchhammeri nd (pl. 82, fig. 1 in Berry, 1916). RHAMNOPHYLL: — B. Generalized leaf. — A. “Cinnamomum” inter- ment is based on the presence of Tempskya and Cycadeoidea in Montana and South Dakota. The sarge shifts approximately 100 km to the ort his Late Cretaceous reconstruction ae it just north of the present 49th parallel (about 55°N paleolatitude). The Late Cretaceous temperate-warm humid zone is based on the oc- currence of Ni/ssonia and Pseudoprotophyllum, and the subtropical humid zone is based on palms and Dewalquea. Smiley (1967), basing his arguments primarily on North American high-latitude floras, indi- cated that the climate gradually warmed from the Jurassic into the Albian, after which a general cooling occurred through the Late Cretaceous (at the same time as Alaska was rotating to the south). BE. 1987] Scott & Smiley (1979) reported on the micro- and megafossil flora from the north slope of Alas- ka. Their treatment indicates that northern floras emain predominantly gymnospermous-pteri- dophytic until the end of the Late Albian, that they do not exhibit Classopollis pollen in any significant quantity (but see Herngreen & Chlo- nova, 1981), and that they are allied to more southerly Albian floras by abundant spores of Gleicheniaceae and Schizaeaceae. Vakhrameyev (1982) recognized the Early Cretaceous as a time of increasing aridity followed by “humidification sensitive ferns throughout the Lower Cretaceous at high latitudes indicates, and did not continue, since palms in the Senonian indicate a generally equable year-round temperature. Krassilov (1973a, 1975, 1981) stated that Late Cretaceous climates in Asia warm into the Campanian and subsequently cool through the end of the Cre- taceous. THE EARLY ANGIOSPERM FLORA Hickey & Wolfe’s (1975) synthesis of the sys- tematic implications of leaf morphology among extant dicots enables paleobotanists to assess the relationship of fossil taxa by extrapolation. Kras- silov (1977) and Hickey (1984) used a series of informal descriptive names for early angiosperm leaf fossils that attempt to unify fossil species with similar morphology. Such leaf morphotypes are especially appropriate for Early Cretaceous dicot leaves that have been placed mistakenly in extant genera. Morphotypes are best viewed as serving as a descriptive terminology in lieu of revision. Most important morphotypes from western North American mid-Cretaceous floras are il- lustrated in Figure 4 (see also Hickey, 1984). The drawings are based in some cases on individual taxa and in others on an idealized composite of several different taxa (see figure legend). In all cases the drawings represent the general form or range of forms present in the morphotype. Table 1 lists the morphotypes along with the genera that have been used in the past for species be- longing to each. Extant genera, such as Populus, which has been mistakenly regarded as embrac- ing a large number of disparate fossil taxa, may be listed under more than one morphotype. Ex- tant genera cannot be demonstrated from the mid Cretaceous, with rare exceptions (Doyle, CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 715 1969; Walker & Walker, 1984). Extinct genera for which the type species is of Cretaceous age are included under the appropriate morphotype according to the type description. Fossil genera for which the type is younger than Late Creta- ceous have not been considered for inclusion, seem the inten focus on early angio- perm hus, Laurophyllum Saporta is named from ae material; hence, I consider it in- appropriate as a generic placement for Early Cre- taceous angiosperms. Morphotypes, since they are not formal taxonomic classifications, have no types. I have chosen to discuss the early angiosperm flora of the NRM region using the leaf morpho- types as a descriptive base. For each morphotype I provide a brief description using the leaf ar- chitectural terminology of Hickey (1979). In ad- dition, I provide particulars relevant to the dis- tribution in time and space for each morphotype along with observations on relative abundance and diversity. Possible relationships for each morphotype are discussed individually and sum- marized in Table The occurrence and distribution of early an- giosperms in the NRM region is presented in Table 2, which encompasses the first five million years of angiosperm history in the region. Where possible, species are grouped according to mor- photype. All Albian and major floras from the region are included, in addition to several pre- viously unreported collections. Names of taxa appear as originally published except for floras examined by myself, for which I have provided identifications. Ages and locations of floras ap- pear in Figures | and 2 and in Appendix I. o Pentalobaphyll (Araliaephyll). Leaves sim- ple, orbicular, 3-5-lobed. Margin entire. Base + cuneate. Primary venation palinactinodromous, with 3 primary veins diverging from above top of petiole, and 2 subprimary veins branching from lateral primaries just above base. Secondary ve- nation eucamptodromous, rather weakly devel- oped. Tertiary venation reticulate to transverse, AR to AO. Higher order venation and cuticle not observed. This very distinctive group is first recognized in the region during the Middle Albian. Penta- lobaphylls are lobate leaves with five principal veins (Fig. 4). Araliaephyllum obtusilobum Fon- taine is included in this group on the basis of the 716 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 1. Botanical affinity and important genera for Cretaceous leaf morphotypes. Morphotypes Important genera! Botanical affinity Sapindophyll Rosidae Pentalobaphyll Fontainea Sade! CM die pure “Androm- eda," “Diosp ” “Ficus,” “Rhus ‘Salix,” 5 it a. a” Araliaephyllum Fontaine, Sterculites Dawson, ‘‘Ara- lia,’ "H edera,” “Liquidambar,” “Sassafras,” "Ster- Unknown; possibly Rosidae or Magnoliidae (Laurales) culia’ Platanophyll * Araliopsis," Araliopsoides Berry, Aspidophyllum Les- | Hamamelididae-Platanales quereux, nari phyllum ae ioe spido- phyllum Hollick, “Aralia,” “Plat " “Sassafras” Protophyll Cissites Debey, Credneri Ae ee Polyphyletic within Hamamel- Richter, Protophyllum Lesquereux, Pseudoproto- ididae — probably Platanales phyllum Hollick, Viburnites Lesquereux, “Alnus,” and Hamamelidales * Betula," “Cissus,” *Parrotia," “Platanus,” “ Popu- lus,” “ Trochodendrophyll “Cercidiphyllum,” ** Coccu- dicus sa Berry, lus " “Paliurus,” “Populus,” “Smilax,” o Menispermaphyll is," “Cocculus,” “Heder Nymphaeaphyll geai tes Lesquereux, oe. pelop IE Hollick, Hederaephyllum Fontaine, Nelum- Trochodendrales and Crecidi- phyllales 99 66 Cissam- Unknown; possibly Ranuncu- ii i Magnoliid-Nymphaeales bites Berry, Paleonuphar Hollick, Populophyllum Fontaine, , Proteaephyllum Fontaine, “Castalia Hedera Magnoliaephyll Liriodendropsis Newberry, Liriophyllum Li je acy i some Magnoli- gnolia Plein (Krasser) Seward, “Bauhin idae, p y Laurales, *Ficus," "Laurus," “Liriodendron,” u Magnolia," TE * Persea," mesas S ra Cinnamomophyll Cinnamomoides Seward, “*Benzoin,” Cinnamo Polyphyletic; some probably mum," “Cocculus,” “Litsea,” *Oreodaphne," “Pal- Laurales iurus,” Ene ud Cornophyll C Eis ipie ewberry, ° <. ” “Cornus,” Polyphyletic; some possibly “Dio. i pos cus" “Rham Rosidae Rhamnophyll Polyphyletic; some possibly >“ Rhamnus,” us" Macclintockia Heer, PM " “Ficus,” “Pal- rus," “Piper, “Smilax,” “Zizy- palmate Dilleniidae ' See text for criteria used for inclusion of gener. ? Quotation marks indicate extant genera Loma identified in the Cretaceous flora. characteristic venation, although this and several other species may have three-lobed leaves. Doyle & Hickey (1976) include this species in their Plat- anoid line, based primarily on the palinactino- dromous primary venation. The palmate lobing and palinactinodromous venation of the Penta- lobaphylls is here considered to be insufficient evidence to establish relationship with the Plat- anoid line when viewed along with the balance of leaf-morphological characters. Pentaloba- phylls show leaf-morphological characters, in- cluding entire margins, eucamptodromous sec- ondary venation, and weak tertiary venation, which serve to distinguish the group from Plat- anophylls. The tendency towards orthogonal branching of tertiary and quaternary veins so characteristic of Platanophylls is absent in Pen- talobaphylls. Certain palmate leaves from the Albian flora cannot be accommodated under either the Pen- talobaphyll or Platanophyll morphotype. These include “Liquidambar” fontanella Brown, with glandular-toothed margins, and “Sassafras” bradleyi Brown, which has smooth margins but lacks the suprabasal lateral branches character- istic of Pentalobaphylls. Fritel (1914) recognized seven species of Pen- Cenomanian. He characterized the group based 1987] primarily on the three basal primary veins, the lateral two of which give off prominent lateral veins just above the point of radius, and on the camptodromous secondary venation. However, " included dnas E serrate margins, highe order venation iens as piae saportanea Les- quereux, which I assign to the Platanophylls. Pentalobaphylls are perhaps the most abun- dant dicots from Albian floras of the NRM re- gion. Albian Pentalobaphylls from Wyoming (Fig. 5), Montana (Fig. 6), and Alberta (Figs. 7-9) are representative of the group. Fritel (1914) placed the Alberta species Araliaephyllum westonii (Dawson) Bell and “Aralia” rotundata Dawson into synonymy with Araliaephyllum kowalews- kiana (Saporta & Marion) Fritel from Europe. A more thorough treatment of the group is needed before such conclusions are substantiated. Cu- ticular detail for the group is unknown with the exception of the Senonian Araliaephyllum po- levoi (Krystofovich) Krassilov (1973b) from the eastern Soviet Union, for which a similarity to Sassafras and Lindera is suggested. G. Upchurch (pers. comm.) has suggested a lauralean affinity for leaves of the A. polevoi group based on cu- ticular detail, sinus bracing, mesophyll secretory glands, and possible basilaminar secondary veins. The secondary and tertiary venation (Figs. 4, 8) in this group suggests rosid affinity. "t Platanophyll. Leaves simple. Margins entire or serrate, lobed or unlobed. Primary veins usu- ally 3, palinactinodromous with several pectinal secondaries on laterals. Secondary veins straight, forking or exmedially branched, craspedodro- mous to teeth, brochidodromous if margin en- tire. Tertiaries and quaternaries forming an or- thogonal network. Teeth platanoid, glandular processes often nipple-shaped. Cuticular struc- ture in Upchurch (1984b), Krassilov (1973b), Kvacek (1983), Bůžek et al. (1967), Némejc & Kvacek (1975). Members of this widespread Laurasian mor- photype date from the Middle Albian in North America. The earliest occurrence from the NRM region is “Platanus” sp. (Fig. 10) from the upper Middle Albian of Alberta (Bell, 1956). Plata- nophylls of Middle and Late Albian age occur in several floras from the NRM region (Table 2). A particularly rich assemblage of Platanophylls oc- curs in the Late Albian Blackleaf Formation in Montana (Figs. 1 1-14). Several poorly preserved specimens representing the group are reported CRABTREE— NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 717 from the Middle and Upper Albian of Alberta (Bell, 1956). As in the Albian flora of Alaska (Hollick & Martin, 1930) and eastern North America (Doyle & Hickey, 1976), the group is most Hio quad encountered in coarse fluvial sedim pem cretacea (Lesquereux) Berry is an early member of this complex and is reported from the Late Albian in the Patapsco Formation in eastern North America (Hickey Doyle, 77). Arali d f is al ted from No the Cenomanian of Texas (Berry, 1922b), Kansas (Lesquereux, 1874), and eastern North America (Berry, 1916; Hickey & Doyle, 1977). In the NRM region A. cf. cretacea is known from the Late Albian Summit locality in the Blackleaf For- mation (Table 2, Fig. 14). Platanophylls are typically more abundant in higher latitude floras (above approximately pa- leolatitude 45°N), and the group may appear in the NRM region before eastern North America. Little debate exists over the platanoid affinities of Platanophylls, but the precise timing of the origin of the Recent order and family remains unresolved. Recent Platanus is reported as being of Senonian age on the basis of pollen mor- phology (Pactlova, 1978), with pollen of platan- aceous character recorded from the Albian (Pact- lova, 1982). Némejc & Kvacek (1975) proposed on the basis of cuticular analysis that Cenoma- nian Credneria spp. (here considered Proto- phylls) from Bohemia are related to the platan- aceous line. Early Platanophylls are related to, and convergent with, other groups of the emerg- ing Hamamelididae, especially the Protophylls. Fructifications regarded as of platanoid affinity are widespread in the Late Albian and Ceno- manian of North America. Notable among these are records of “Sparganium” from Washington (Fig. 35) and Wyoming (Fig. 41; Brown, 1933a), and “Platanus” from Greenland (Seward & Con- way, 1935) and Kansas (Lesquereux, 1892). For further discussion of Cretaceous fructifications of platanoid affinity see Dilcher (1979) and Crane et al. (1986). Platanophylls are allied to Protophylls on the basis of well-defined, orthogonal-reticulate, ter- tiary and quaternary venation. Palinactinodro- mous venation and a greater tendency towards lobation distinguishes the Platanophylls from the Protophylls, which have pinnate organization (Fig. 4). I have chosen to segregate these leaves in the hope that the Cenomanian Cissites-Bet- ulites-Alnites complex (Protophylls) might be 718 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 TABLE 2. Early angiosperm flora from the northern Rocky Mountain region. 2? Boul- der Pasay- Cree ten (6c) (17) Beaver Mines (4) A BC Mill Creek (16) Lower Gates (6a) Upper Gates (6b) — © — ~ — o > x = = Kingsvale (14) Pre-Muddy (18) Jackass (12) Summit (19) Black Eagle (5) 3 g A B C D > = > = Sapindophyll Celastrophyllum acuti- i X X X X Celastrophyllum sp. x cf. Celastrophyllum sp. x Daphnophyllum dako- te Fontainea grandifolia Newberry x X X X rues E F. grandi- folia berry X . ed sp. X “Laurus” crassinervis D awson “Myrica” serrata Pen- hallow Proteoides daphnoge- noides Heer X “Quercus” flexuosa? Newberry “Salix” inaequalix? Newberry cf. “Rhus”? powelliana Lesquereux X Sapindopsis angusta (Heer) Seward & Conway Sapindopsis cf. angusta X Sapindopsis sp. aff. an- gusta yia belvideren- erry Sapindopsi brevifolia? Fon ontaine X Sapindopsis cf. brevifo- li ia Sapindopsis magnifolia Sapindopsis cf. magni- folia Sapindopsis sp. X Sapindopsis sp. X cf. Sapindopsis sp. X cf. Sapindopsis sp. X Pentalobaphyll “Aralia” rotundata awson “Aralia” westonii Daw- 1987] CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS TABLE 2. Continued. Beaver Mines (4) ABC Lower Gates (6a) Upper Gates (6b) Pre-Muddy (18) (6c) (17) AB A B Kingsvale (14) Jackass (12) ABC Mill Creek (16) Summit (19) g Sun River (20) Black Eagle (5) > w Araliaephyllum westonii (Dawson) Bell Araliaephyllum? sp. raliaephyllum sp. i ar" inte grifolium Lesquereux “Sassafras” mudgii? Lesquereux cf. “Sterculia? mucro- nata Lesquereux “Sterculia” vetustula Dawson “Sterculia” sp. Protophyll Alnites insignis? Daw- Cissites? sp. cf. Cissites sp. “Platanus” affinis Les- quereux “Platanus” affinis var. ampla Dawson Populites dawsonii Bell Populites cf. dawsonii Populites sp. cf. Protophyllum sp. Platanophyll “Aralia” wellingtoniana Lesquereux cf. “Aralia” wellington- Araliopsoides cf. creta- cea (Newberry) Berry cf. Aspidophyllum trilo- atum Lesquereux Platanophyllum sp. “Platanus” heeri Les- “Sassafras” cretaceum Newberry Trochodendrophyll “Cercidiphyllum” sp. X X X XX XX X x< 720 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 2. Continued. Boul- der Pasay- reek te (6c) (17) A B A B Beaver i Mill Creek (16) Lower Gates (6a) Pre-Muddy (18) 4 A B C ABCD Kingsvale (14) Sun River (20) Black Eagle (5) Jackass (12) Summit (19) jpe potoma- x Tr serena am (“Cercidiphyllum”) potomacensis (Ward) Nymphaeaphyll “Hedera” ovalis? Les- quereux Menispermites renifor- nis Dawson Menispermites sp. X Nelumbites sp. X Nymphaeites sp. X Magnoliaephyll “Magnolia” alternans eer "Magnolia" magnifica Dawson Cornophyll “Ficus” fontainii? Berry Laurophyllum debile Dawson Rhamnites sp. X “Rhamnus” sp. x Cinnamomophyll Cinnamomoides ovalis (Dawson) Bell x x x X x Cinnamomoides cf. ovalis (Dawson) Bell x Cinnamomoides sp. x Cinnamomoides sp. x Cinnamomoides sp. x Macclintockia cretacea Heer “Paliurus” ovalis Daw- son X X Uncertain Capparites? sp. X cf. “Ficus” ovatifolia 1987] TABLE 2. Continued. CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS Beaver Mines (4) A B C Lower Gates (6a) Upper Gates (6b) Fall River (10) (18) Boul- der reek (60 - “ay AB AB ae) zg ° "< Mill Creek (16) (14) Jackass (12) Summit (19) Black Eagle (5) A B C D Juglandites cretacea Dawson * Magnolia"? sp. Menispermites? sp. Myrtophyllum Pu Seward & Conwa * Populus" cf. berggrenii eer cod cyclophylla Quercophyllun? sp. "Quercus" coriacea Ne “Salix”? sp. "Salix" s “Sassafras” parvifolium Fontain x X mK mK ! Numbers in parentheses refer to geographic and stratigraphic position of floras as indicated in Figures | a 2 All species are included as originally cited in the references given below. I have identified the species from the Summit, Sun ? Vertical rows indicate Penhallow, 19 1929c. (16)C— aon 1956. (16)D— Mellon, 1967. (7)A— and (5)— Cra icis Canada, provided tl 07. (17)B — Bell, 1956. (14) — Bell, 1956. (12)— Bell, 1956. (16)A Denion: 1886. (16 ee, this report. It should be noted that W. A. Bell, in his capac ty | as baleobotan River, Black Eagle, Fall River, and Pre-M uddy collections. oe Berry, 1929b. o> Bell, 1956. (4)C— Bell, 1956. (7)B— Mellon, 1967. (10), (18), (19), (20), nist with the t (1963), Mellon (1967), Geological ^m and Mellon et al. (1963). allasa, p better understood. I suggest that Protophylls are more similar to mainline Hamamelidales on the basis of leaf architecture than are Platanophylls, which are closer to the platanoid line of Ham- amelids. Protophyll. Leaves simple. Margin entire to entate, occasionally lobed. Venation pinnate, the basal secondaries sometimes strengthened phylls. Cuticular details in Némejc & Kvacek (1975) and Krassilov (1973b) Richter (1905) monographed Credneria Zen- ker and Paracredneria Richter. Protophylls can usually be distinguished from Platanophylls by the basically pinnate venation and by the more consistent craspedodromy of the secondary veins (Fig. 4). However, in many cases the two morphotypes intergrade to the ex- tent that recognition is difficult. Protophylls are common in the NRM region but do not exhibit the diversity or dominance found in the coarse facies of the Cenomanian Dakota flora of Ne- braska and Kansas (Lesquereux, 1892). side species from the Albian floras of Alberta are placed in this morphotype (Table 2). Seit pen are present in the Blackleaf flora (Fig. 15). Basal venation in Platanophylls and Protophylls is compared in Figures 16 and 17 x 3 ng EET = w Z. WW a a < c < = Z. < = ° x 2 e N na = Ln x = u. ° N — < Z Z, < 1987] Sapindophyll. Leaves even to odd pinnatifid or pinnately compound. Leaflets opposite or al- ternate, sometimes decurrent on rachis. Margins entire to serrate. Teeth apparently cunonioid. Petiole stout. Petiolule stout. Primary vein stout. Secondary veins eucamptodromous or semicraspedodromous, arched-ascending, often numerous, diverging at + uniform angle of 35- 50*. Intersecondary veins typically present. Ter- tiaries weakly percurrent, transverse, AR but highly variable. Quaternaries irregularly reticu- late. Cuticle and tooth morphology in Upchurch (1984b). This unique Lower and mid-Cretaceous mor- photype encompasses fossils with pinnatifid or pinnately compound leaves (Fig. 4). Rachises with attached leaflets have been reported as Sapin- dopsis Fontaine and Fontainea Newberry. The widespread Laurasian species “Andromeda” a Heer, “Ficus” daphnogenoides (Heer) a F> Rus Lesquereux, ' *Sapindus morrisonii Lesquereux, and “Rhus” powelliana Lesquereux represent i pene leaflets of Sap- indophylls. Dissected leaves with palmate or- ganization, cf. Halyserites Sternberg and Cus- soniophyllum Velenovsky, are excluded from the Sapindophylls "i have not been observed in the NRM regio Arahe are an important component of many North American Albian and Cenomanian floras (Berry, 1922a, 1922b; Fontaine, 1889; Hickey & Doyle, 1977; Lesquereux, 1874, 1883, 1892; MacNeal, 1958; Upchurch, 1984b). The morphotype is generally considered to be more abundant in floras from regions to the south of the NRM region and is sometimes regarded as representing thermophilous plants. Nonetheless, CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS T23 the morphotype is common in mid-latitude flo- ras from the NRM region and Greenland (Sew- ard & Conway, 1935), and occurs in southern Alaska (*Ficus" daphnogenoides in Hollick & Martin, 1930) and the north slope (pers. obs.). This morphotype is represented by several species in the NRM region (Table 2). A probable new species of Sapindopsis of Middle Albian age iS ostate from wicks equivalent to the Ther- Montana (Fig. 18). It aoan to be related to the Cenomanian species “Sapindus” morrisonii and “Ficus” beckwithii. A Sapindophyll similar to Sapindopsis variabilis Fontaine is present in the Blackleaf Formation of Montana (Fig. 19). Sapindophylls are present in the Albian of Alberta (Table 2, Fig. 20), British Columbia (Fig. 27), and Washington (Figs. 28- 32). A putative Sapindophyll (cf. Sapindopsis sp.) from the Upper Albian of Idaho is tentatively assigned to this morphotype (Figs. 21-23). This species is allied to the morphotype by admedial orientation of the tertiary veins (Fig. 21) and dissected leaves with decurrent leaflets (Figs. 22, 23), but shows the irregular laminar furcations of Halyserites. Sapindophylls may be most closely allied to Recent basal Saxifragales and to other rosid groups such as the Sapindales and Rutales. This relationship is based on the pinnately compound leaves, eucamptodromous venation, admedia orientation ofthe transverse tertiary veins (Hick- ey & Doyle, 1977), cunonioid teeth in some species (Upchurch, 1984b), and cuticular fea- tures (Upchurch, 1984b). Crane et al. (1986) dis- cussed possible relationships between Sapindo- duced tricolpate pollen (viz. Hamamelididae and — FiGures 5-58. Representative dicotyledonous leaves from the Albian through middle Campanian of the NRM region. All figures with centimeter scales. Specimens from t he U.S. Nation m are provided with al haad field locality numbers, and those from the Geological Survey of Canada are provided with GSC locality mbers. Types and other cataloged specimens are indicated where appropriate, along wit National akua (USNM) and Geological Survey of Canada d accession numbe ames applied to illustrated specimens are as originally published or as identified in paleobotanical collections at the US and GSC. I have provided names edd for species from the Fall River, Blackleaf, Pre-Muddy, Pasayten, MES Winthrop, and Two Medicine floras FIGURES 5-12. Albian age Senet Nps leaves of the Pentalobaphyll and Platanophyll morphotypes. 5- 9. Pentalobaphylls.— 5. cf. “Sterculia” mucronata Lesquereux, Fall River Formation, USGS locality 7404. 6- 8. Araliaephyllum westonii (Dawson) Bell.—6. Blackleaf Formation, USGS locality 9439. 7, 8. Mill Creek Formation. — locality 1815, GSC 5117, holotype.—8. GSC locality 3066, GSC 5874. —9. “Aralia” rotundata Dawson, Mill Creek Formation, GSC locality 1815, GSC 5116, holotype. 10-12. Platanophylls.— 10. “Platanus” sp., Mill Creek Formation, GSC locality 1815, GSC 5106.—11. Platanophyllum sp., Blackleaf Formation, USGS locality 9437.—12. cf. Aspidophyllum trilobatum Lesquereux, Blackleaf Formation, USGS locality 9437. 724 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIG an Beg age e dicotyledonous leaves ih Platanophyll and Protophyll morphotypes. 13, 14. Waaa — 13. cf. “Aralia” wellingtoniana Lesquereux, Blackleaf Form U locality l leas Na cf. mae a (Newberry a erry, Blackle cea 15. Protophyll, cf. Protophyllum sp., Blackleaf JSGS locality 5986. . Comparison of basal venation of PI land l Sub Platanophyllum sp., dubi cue USGS locality 5939. — 17. Protophyll, Protophyllum sp., Blackleaf Formation, USGS locality 5986. Rosidae). Friis & Skarby (1982) and Basinger & Dilcher (1984) discussed Cretaceous flowers and pollen of early Rosidae. Sapindophylls are restricted to Albian and Cenomanian floras in the NRM region. I am not familiar with their Late Cretaceous range else- where. The rosid affinities of Sapindophylls need to be confirmed by further study, as does the relationship between the group and the putative Upper Cretaceous and Paleocene derivative Av- erhoites. Trochodendrophyll. Leaves simple, ovate to reniform. Margin crenate. Teeth convex-convex (A-1), chloranthoid. Venation actinodromous. Primary veins usually 3 or 5, sometimes more, arched apically. Secondary venation semicras- pedodromous with + prominent brochidodro- 1987] mousarches present apically. Tertiary and higher order venation not well known, appearing retic- ulate in some with tendency towards orthogonal branching This oi is characterized by actinodromous rimary venation and crenate margins bearing d teeth (Fig. 4). The latter character distinguishes the group from those Menisper- maphylls with actinodromous venation. Tro- chodendrophylls are reported in low frequencies during the Albian, and are represented in the NRM region primarily from Alberta and British Columbia (Table 2). Trochodendroides ('* Cerci- diphyllum”) potomacensis (Ward) Bell conforms most closely to the concept of the morphotype with five primary veins, brochidodromous sec- ondary veins, and crenate margin (Figs. 24, 25). Trochodendrophylls have been assigned to rather unlikely genera (Table 1). Cuticular stud- d. of Menispermites potomacensis Berry from Lower Cretaceous Potomac Group (Up- Y adio l ; pers. comm.) indicate that the stomatal apace of this species does not con- form to that of extant Cercidiphyllales and Trochodendrales. Probable members of the lin- eage are reported in high frequencies from Seno- nian and latest Cretaceous deposits, but these are more advanced in morphology and in some cases can be appropriately assigned to the Recent Trochodendraceae and rcidiphyllaceae (see Krassilov, 1973b). Trochodendrophylls, along with a series of pinnately veined Lower Creta- ceous leaves with chloranthoid teeth and open craspedodromous secondaries (see Upchurch, 1984b; Fig. 36, this report), may represent an ancestral plexus from which Recent Trochoden- drales and Cercidiphyllales evolved during the Cretaceous (but see Wolfe, 1973). The Campani- n Two Medicine flora (see below) includes four leaf taxa which have puni reds teeth, but which and higher order suggest mosa d elopment ofleaf characters in Cercidiphyllales, Trochodendrales, and Chloranthales. Menispermaphyll. Leaves apparently sim- ple, broadly deltoid and + trilobed to circular or ovate-elliptic. Base cordate to obtuse. Margins entire or lobate forms occasionally with teeth. Venation acrodromous or actinodromous with 3-9 primary veins. Primary veins craspedo- drome or becoming camptodrome just before margin. Secondary veins irregularly spaced, camptodrome in many but craspedodrome to the CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 725 teeth in nonentire forms. Tertiary veins ran- dom-reticulate to transverse. Higher order ve- nation not observed. Cuticle unknown. This group (Fig. 4) superficially resembles Nymphaeaphylls but is never peltate and never has an expanded, multistranded petiole. Cor- date leaves with major veins unbranched to the margin are best placed in Menispermaphylls, whereas such leaves with highly branched pri- mary and secondary veins are more appropri- ately placed in Nymphaeaphylls. Trochodendro- phylls may be distinguished by their crenate margin. Menispermaphylls are morphologically con- vergent with extant Menispermaceae (Ranun- culiidae). The modern family includes both sim- ple-leaved and ternately compound members. The leaves and leaflets are deltoid to ovate with entire or sparingly toothed margins. Tooth mor- phology is a modification of the chloranthoid type (Hickey & Wolfe, 1975). Primary venation is actinodromous or acrodromous, and major veins terminate in a fimbrial vein, thus appearing craspedodromous. Many Recent species exhibit a mucronate leaf apex; simple, unbranched tri- homes; and well-developed, orthogonal-retic- ulate, higher order venation. Menispermaphylls resemble Recent Meni- spermaceae in shape, margin, and venation. However, before relationship to the family can be established it is necessary to demonstrate the presence in the fossils of such characters as fim- brial veins and simple, unbranched trichomes. Nevertheless, the unusual combination of entire margin and craspedodrome primary veins is an important point of agreement between the fossils and Recent Menispermaceae. Menispermaphylls show their greatest diver- sity in the Dakota flora of Kansas and Nebraska (Lesquereux, 1892). However, here, as in the NRM region (Table 2), they are a relatively mi- nor component ofthe flora. The Campanian Two Medicine flora (see below) includes a species that is entirely consistent with the leaf morphology of extant Menispermaceae. Magnoliaephyll. Leaves simple, ovate to el- liptical. Margin entire or 2- or 3-lobed. Primary venation pinnate in entire forms, with 3 primary veins from base in lobed forms. Secondary ve- nation brochidodromous or festooned brochi- odromous. Higher order venation variable. This morphotype (Table 1) comprises a large number of problematic taxa that form an im- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 1987] portant part of all but extreme northern Laura- sian floras. The group is erected primarily for Te due to the enigmatic positions of most of its members. Some Potomac Grou a for which cuticular evidence is available (Upchurch, 1984a, 1984b) appear to be related to Laurales and Illiciales. “Magnolia” magnifica Dawson is illustrated as an example of the morphotype (Fig. 26). Early members of the morphotype from the NRM region are listed in Table Putative magnoliid infructescences are known from the mid Cretaceous of central North Amer- ica (Dilcher, 1979), Japan (Nishida, 1985), and Greenland (Seward & Conway, 1935). Walker et al. (1983) reported pollen of Winteraceae from the Lower Cretaceous of Israel. Cinnamomophyll. Leaves simple. Margin entire. Venation pinnate. Prominent suprabasal secondary veins arching apically. Secondary ve- nation brochidodromous. Basilaminar second- ary veins sometimes present. Tertiary veins transverse, regular or somewhat irregular in spacing and course. Quaternary veins weak, ran- dom-reticulate. Cuticle unknown. This morphotype (Fig. 4) is widely reported from NRM region floras and is an important component of the angiosperm vegetation during the Albian (Table 2). Although not present in all Cinnamomophylls, basilaminar secondary veins indicate that some may be related to extant Lau- rales. Cinnamomophylls are most important in floras from middle latitudes in Laurasia where they are consistently represented in the Albian floras Nymphaeaphyll. Leaves simple, orbicular to if Margin entire. Petiole often multi- venation forming weak brochidodromous arch- CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS TET es, craspedodromous in “Castalia” and Casta- liites. Cuticle unknown. his morphotype converges with Trocho- E M from which it is distinguished by its entire margin, and with Menispermaphylls (see discussion of Menispermaphylls and Fig. 4). Several species of dique ee exhibit leaf architecture consistent with those of extant Nymphaeales and "URN iae ius from cuticles and fruits is needed to confirm the re- lationship. Cordate and peltate leaves with actinodro- mous venation and entire margins are abundant and diverse in the North American mid Creta- ceous, but they are not well represented in the NRM flora (Table 2). Nymphaeaphylls may rep- resent an adaptational syndrome for aquatic hab- itats as suggested by Samylina (1968), Axelrod (1970), and Krassilov (1977). Hughes (1976) sug- gested that this leaf morphology may represent simply an early “experimental” shape that is not correlated with aquatic habitats. Rhamnophyll. Leaves simple. Margin entire or crenate-dentate. Base acute to truncate. Ve- nation acrodromous. Primary veins 3-5, arising from within the petiole and arching apically. Sec- ondary venation brochidodromous or semicras- pedodromous to teeth, often with several M ipn and prominent secondary veins originating on outermost primary veins and arching apically along margins. Tertiary veins transverse, regu- larly spaced and usually numerous. Higher order venation not observed. Cuticle unknown. Rhamnophylls are rare from floras of Albian age in the NRM region (Table 2). The group occurs with greater frequency in Late Cretaceous tanical affinities of the former remain unknown. — FIGURES 18-26. phyll morphotypes. 18-23. Sapindophylls. Albian age dicotyledonous leaves 2 ps CC aesan Hoe cp de and Magnoliae- 3. —18. cf. * wellia a Lesqu , Pre-Muddy collection. — 19. cf. Sapindopsis Sp., Blackleaf Formation, USGS Mat = 6007. ri “Ko nines "erandifolia Newberry, Com- (Ward) Bell, Kingsvale Form 26. Magnoliaephylls. cress S=. C collec GSC 6639. 21-23. cf. Sap- ns), 5. Trochodendrophylls, S ucdendroidis CC ercidiphyllum’’) potomacensis 4. ocality 3449, GSC 6654.— mas Dawson, GSC locality 290, GSC 5133, holoty . GSC locality lt GSC 5907.— 728 Cornophyll. Leaves simple. Margin entire. Primary venation pinnate. Secondary venation camptodromous, arching apically. Tertiary veins transverse, regularly spaced, more or less nu- merous. Higher order venation and cuticle un- known Cornophylls occur in limited diversity and abundance in the Albian floras of the region (Ta- ble 2). The group assumes a more important po- sition in later Cretaceous floras. The botanical affinity of the group is unknown, but some mem- bers appear to exhibit morphological similarities to Rosidae. Dryophyll. Leaves palmately compound. Leaflets ovate to lanceolate. Margin serrate. Pri- mary venation pinnate. Secondary venation craspedodromous. Higher order venation not observed. Cuticle in Krassilov (1973b) and Né- mejc & Kvaéek (1975). This group of palmately compound leaves with serrate margins is not present in the early angio- sperm floras. They appear first in the uppermost Albian and Cenomanian. Serrate forms with craspedodromous venation and lanceolate leaf- lets (cf. Dryophyllum Debey) can be shown to with Fagales, al- though this Is a a tenuous association for most Cretaceous forms (Wolfe, 1973). FLORISTIC PROVINCIALITY OF THE NORTHERN ROCKY MOUNTAIN REGION DURING THE ALBIAN Discussion of floristic provinciality during the Cretaceous ultimately depends on accurate cor- relation of stage and substage boundaries be- tween the proposed provinces. The Upper-Low- er Cretaceous boundary and the position of the Albian substages in the NRM region are placed based on ammonites in the United States (Cob- ban & Reeside, 1951, 1952a) and on a varied molluscan fauna in Canada (Jeletsky, 1968; Kauffman, 1975). Geologists have generally placed the boundary between the Upper and Lower Cretaceous in the NRM region at the top of the Mowry Shale. This boundary coincides with the uppermost widespread occurrence of Neogastroplites (Cobban & Reeside, 1951). It also coincides with the first occurrence of tricolporate pollen in the region (Nichols et al., 1983; Norris et al., 1975). Nonetheless, uncertainty exists over the correlation of the regional boundary to the European standard. The boundary in the western ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 interior is radiometrically dated as 94 Ma (Obradovich & Cobban, 1975; but also see Fouch et al., 1983), whereas in Europe it is dated as 97.5 Ma (Harland et al., 1982) Brenner (1976) recognized a Northern Lau- rasian Province and a Southern Laurasian Prov- ince based on the palynology of Lower Creta- ceous deposits. The northern province is recognized by a coniferous palynoflora less di- verse and more heavily bisaccate than the south- ern province. The NRM region lies on the boundary between provinces. Such provinciality is supported by Srivastava (1981) and by Hickey & Doyle (1977) and rejected by Herngreen & Chlonova (1981). Angiosperm pollen has been used for the recognition of Late Cretaceous pro- vinciality (Srivastava, 1981). The extension of angiosperm palynofloral provinces into the Early Cretaceous seems reasonable in light of the elab- oration of the timing of the migration of angio- sperms provided by Brenner (1976) and by Hick- ey & Doyle (1977). The poleward migration of early angiosperms was first recognized from megafossils (Axelrod, 1959) and subsequently refined through paly- nological studies (Brenner, 1976; Hickey & Doyle, 1977). Early angiosperms appear to have arisen in northern Gondwana and to have crossed the Tethys into Laurasia in the Late Barremian and Aptian (Hickey & Doyle, 1977). aby of mono- sulcate-prod rthern lat- itudes of Laurasia was is delayed by several million years. This may be attributed to climatic or other environmental barriers, inadequate dispersal mechanisms, or merely to stalwart pre-existing vegetation. Tricolpate taxa exhibit a far shorter lag period. The first major radiation of the tri- colpate group occurs on a worldwide scale during the late Middle Albian (excepting at the very highest latitudes). Considerations of floral provinciality of early o common prob- Sw hudy & Tschudy, 1986), es- pecially in the Paleozoic and Mesozoic, and 2) spurious conspecificity, 1.e., the tendency to as- sign pollen grains to existing taxa. Both problems obscure provincialism at lower taxonomic levels. Penny (1969) commented on the impoverished appearance of Cenomanian pollen floras relative to leaf floras of that age. A related problem, which is more or less confined to the early angiosperm 1987] pollen floras, is the general absence of differen- tiation of grains with respect to ornamentation, shape, and gross morphology. In addition, Me- sozoic palynologists do not normally elaborate species frequency as is common for Holocene workers, a situation that makes it difficult to re- view existing literature for data which could be used to determine provinciality based on dom- inance and importance of species. On the other hand, a common problem with megafloral tax- onomy has been overspeciation due to failure to recognize physiognomic variation in leaf mor- phology within natural species. Both megafloral and microfloral treatments are restricted by the taphonomic biases implicit in any depositional environment. The first angiosperms known from the lower Middle Albian (Potomac subzone IIB) of the NRM region are two grains of Tricolpites micro- munus (Groot & Penny) Singh from the upper Loon River Formation in northwest Alberta (Singh, 1971) and monosulcate grains from the Cloverly Formation in Wyoming (Davis, 1963). Pollen sz megafossils are known throughout the NRM region with certainty from the middle Middle hae (Bell, 1956; Davis, 1963; Nichols & Jacobson, 1982; Norris, 1967; Singh, 1975; Roberts, 1972). Singh (1971) reported nine species of middle Middle Albian angiosperm pollen from northwest Alberta. By upper Middle Albian (Po- tomac subzone IIB), angiosperm pollen becomes a persistent element of moderate frequency in the microflora. This upper Middle Albian surge IIB). Localities in British Columbia of equal age yield Sapindopsis and a limited diversity of other dicot leaves (Table 2). The first diverse assem- blages of megafossils are of middle Middle and upper Middle Albian age and include Sapindo- phylls, Pentalobaphylls, Platanophylls, Cinna- momoides, Trochodendroides, and Menisper- mites in order of frequency in collections. The components of this megaflora continue into the lower Upper Albian with little apparent change. The early angiosperm flora of the region is dis- tinct from more southerly floras in its high rep- resentation of Pentalobaphylls, Cinnamomo- phylls, and to a lesser degree Led and several endemic species are prese I compared the Lower Cretaceous Ed codon CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 729 of the Patapsco Formation of Maryland (Bren- ner, 1963) with that of the Loon River and Peace River formations of northwest Alberta (Singh, 1971) to contrast composition, abundance, and diversity of Middle and Late Albian floras of eastern North America with those of the NRM region. More recent treatments of the Patapsco palynoflora exist (Doyle, 1969; Doyle & Hickey, 1976; Hickey & Doyle, 1977), but Brenner's work remains the standard for taxonomic comparison is likely that some of his taxa represent species complexes or genera. Monosulcates first occur in the latest Barre- mian or early Aptian in eastern floras, consid- erably antedating their lower Middle Albian oc- currence in the NRM region. Seven ofthe twelve Patapsco species of angiosperm pollen are tri- colpate. The tricolpates appear during the Lower and Middle Albian and continue upsection to the top of the formation (middle Upper Albian) with no addition of new species (Brenner, 1963; but also see Doyle & Hickey, 1976). Of these seven species, Tricolpites micromunus occurs in the lower Middle Albian Loon River Formation, and six of the seven tricolpates occur in the mid- dle Middle Albian Harmon Member ofthe Peace River Formation. Such palynological data indi- cate close similarity between lower and middle Middle Albian dicots in the eastern and western floras of North America, at least at the generic level. An endemic flora in the region is indicated by angiosperm pollen recovered from the Peace River Group (Singh, 1971), which was deposited tricolpates, are reported fro a high degree ofsimilarity, es strikingly less than that of the earlier flora. Furthermore, five Ca- dotte taxa have not been reported from Creta- ceous palynofloras outside the NRM region. Eastern groups apparently moved north and east into the NRM region during the Middle Albian and persisted in the western flora; however, by the upper Middle Albian, tricolpate and mono- Liliacidites trichotomosulcatus Singh and the tri- colpates Retitricolpites maximus Singh, Tricol- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 1987] pites sagax Norris, T. parvus Stanley, and Frax- inaipollenites venustus Singh ndemic character of the flora appears to result largely from separation of the region from eastern North America by the Skull Creek Sea- way for approximately two million years during the upper Middle and lower Upper Albian. Thus, shortly after angiosperms entered the region, they were cut off from their presumed ancestral stock to the south and east in an area some 20? further north and potentially subject to substantially dif- ferent selective pressures. However, it should be noted that available palynofloras make it difficult to resolve an east-to-west adaptive radiation within groups producing small, reticulate, tri- colpate pollen, since the equivalence of named species to biological species is debatable. In con- trast to the simple tricolpate pollen, leaf archi- tectures and cuticles of Lower Cretaceous angio- sperms provide an expanded suite of characters from which biological species can be circum- scribed. A detailed comparison of North Amer- ican floras of Albian age is beyond the scope of this paper. WINTHROP FLORA A large collection of fossil foliage from the type locality of the Late Albian or Early Cenomanian (Barksdale, 1975) Winthrop Formation in cen- tral Washington (Figs. 1, 2) appears to be the latest occurring **archaic" flora in the region. The flora contains abundant Sapindophylls and Pen- talobaphylls that are closely related to but not conspecific i those ofearlier Albian floras from the NRM regio Several small p of plant fossils from the Winthrop have appeared as floral lists in geo- logical papers (see Barksdale, 1975). Apparently the collections were sent to the U.S. National Museum and examined by F. Knowlton and R. Brown. Reevaluation of these collections in 1986 by S. Wing and J. Wolfe (pers. comm.) indicated a Paleocene age for the flora, and hence their origin in the Winthrop Formation must be ques- tioned. The present report includes only those CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 731 specimens collected from the type locality (Ap- pendix I) by R. Rau, J. Robison, and myself. The Winthrop flora is a moderately diverse assemblage consisting of about 20 species of ferns, conifers, dicotyledons, and Sagenopteris (Cay- toniales). Dicotyledonous species include Sap- indopsis sp. (Figs. 28-32), Magnoliaephyllum sp. (Fig. 33), Araliaephyllum sp. (Fig. 34), Araliae- phyllum westonii, “Sparganium” sp. (Fig. 35), an unidentified dicot with OE r sec- ondary venation and chloranthoid teeth (Fig. 36), Nelumbites sp. (Fig. 37), “Ficus” ovalifolia, Men- ispermites sp., Eucalyptophyllum sp., and several poorly preserved species ee UPPERMOST ALBIAN, CENOMANIAN, AND TURONIAN FLORAS Three well-developed floras, the Aspen, Fron- tier, and Dunvegan (Table 3), provide much of the megafossil evidence for the angiosperm flora of the NRM region during the uppermost Albian and Cenomanian (Fig. 2). The Dunvegan flora is near the northern limit of the region; the Aspen and Frontier are at the southern limit (Fig. 1). Collections from the Albino Member of the Mowry Formation and the Wayan Formation (Figs. 1, 2) provide a few additional species. The Albian angiosperm flora of the region shows an underlying unity due to the occurrence in most assemblages of Sapindophylls, Penta- lobaphylls, Platanophylls, Protophylls, and Cin namomophylls, which give the flora an “archaic” appearance when contrasted to the flora of the post-Albian. The post-Albian flora shows the de- velopment of north-south provinciality in the region. However, because of the small number of floras known from this period (Fig. 2), and the restricted volcaniclastic facies of the Aspen and Frontier floras, such provinciality is at this time al deposits in the northern half of the region, there is uncertainty over how widespread and uniform the early flora was The development of intraregional provincial- — FIGURES 27-37. Sapindopsis sp. 28-37. Winthrop Formation. 28-32. Enlarged leaflet of p te in Figure 28.— joined upper three leaflets. — 33. Magnoliaephyllum sp.— Dicotyledonous leaves from the Pasayten and Winthrop formations. 27. Pasayten Formation, Sapindopsis sp.— 28. 9. . Upper part of compound leaf. — 31. Base of leaf. — 32. Base of 36. Upper part of compound leaf.— —34. Araliaephyllum sp.—35. “Sparganium Unidentified leaf with iuum secondaries and chloranthoid teeth. —37. Nelumbites sp. 732 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 3. eo flora from the uppermost Albian, Cenomanian and Turonian of the northern Rocky Mom region.! Dunvegan (8) Lower Upper Aspen Frontier Frontier (2) (lla) A B (11b) Sapindophyll “Andromeda”? spatula Bel X “Ficus” eiii irai (Heen Berry X “Myrica” nervosa K X Sapindopsis huts it X Rosidlike Leaves “Diospyros” nitida Dawson X “Salix” cumberlandensis Knowlton “Staphylea”? fremontii Knowlton x XX Platanophyll Ampelophyllites attenuatus (Lesquereux) Knowlton “Aralia” cf. parvidens Hollick “Aralia” veatchii Knowlton X Aspidophyllum dentatum? Lesquereux "H ereu Xx B Q [e] = S S 3 & 3 ë r* 6 ld z Š Ë > x X X x stantonii x omm aspensis dig X sins a Mp ulum Hollick "Que Protophyll Credneria macrophylla Heer Credneria truncatodenticulata Bell “Platanus” affinis Lesquereux “Platanus” williamsii Bell Populites cyclophylla Dawson Protophyllum boreale Dawson x Protophyllum leconteanum? Dawson vr multinerve? peni X tophyllum rugosum Lesquereux X ED DM bipes ciem Hollick x XX x x< Pentalobaphyll Araliaephyllum rotundiloba (Newberry) Fritel Araliaephyllum groenlandica? Heer “Sterculia” towneri (Lesquereux) Berry X Xx Dryophyll Dewalquea pulchella Knowlton ey Dryophyllum fesses (Knowlton) Berry x x Trochodendrophyll Castaliites cf. cordatus Hollick (in part) x Trochodendroides (“Cercidiphyllum”) potomacensis (Ward) Berry 1987] CRABTREE— NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 733 TABLE 3. Continued. Lowe Aspen Frontier Dunvegan (5) (2) (11a) A B Upper Frontier (11b) Cinnamomophyll “Cinnamomum” heeri Lesquereux S r. ` hesperium Knowlton “Cinnamom sp Phyllites sp. Nymphaeaphyll Castaliites cf. cordatus (in part) Menispermites reniformis Dawson n Nymphaeites exemplaris Hollick Paleonuphar nordenskioldii (Heer) Bell Magnoliaephyll “Ficus” maxima Dawson “Liriodendron” giganteum Lesquereux * Magnolia" cf. adu Newberry * Magnolia" tenuifolia Dawson “Sassafras” bradleyi Brown Cornophyll sni Aba lesquereuxii Knowlton & Cockerell Knowlton Laurophyllum debile Dawson * Magnolia" rhamnoides Bell Monocots Sabalites sp. Uncertain * Bauhinia”? cretacea? Newberry “Fa 3" Daane Dawson PE ied glascoeana Lesquereux “Ficus” inaequalis Lesquereux "Ficus"? s - “Ficus” s E ca flexuosum oe Bell Leguminosites spatulatus B X »X X Xx XX X KK < x KKK X ! Numbers in parentheses refer to geographic and stratigraphic position of floras as indicated in Figures 1 an š ` All species are included as originally cited in the references given belo vn, 1933a. (11a) —Knowlton, 1917. (8)A— Diwson, 1883. (8)B—Bell, 1963. (11b)—Berry, 1929d. ity in the uppermost Albian and Cenomanian is correlated with the ebb of the Albian epeiric sea- way. As previously discussed, the seaway may have influenced the development of endemism in the flora by restricting movement of eastern species into the Site rae much of the Upper bian. / ithdr at the end of the Albian, venden as from east- 734 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 1987] ern and southern regions may have altered the floristics of the southern part of the NRM region. Although this scenario is hypothetical, the up- permost Albian and Cenomanian floras of Wy- oming contain Dewalquea, a genus characteristic of floras to the south and east of the region. A large collection of fossil leaves from the Wayan Formation in southeast Idaho (Figs. 1, 2) is dominated by pteridophytes (Crabtree, 1983) that are undescribed but appear to be similar to those from the Frontier Formation in southwest Wong (Knowlton, a The Wayan con- p. (Figs. 21-23), which appears to iro a new species. An- drews (1948) and Andrews & Kern (1947) de- scribed a large collection of pteridophytic petri- factions (Tempskya) from the Wayan (Appendix Another undescribed megaflora is known from several localities in the Albino Member of the M ormation in southern Montana (Figs. 1, 2, Appendix I). An extensive fern community is preserved in the deltaic deposits of the Albino (Vuke, 1982; Crabtree, 1983). Dicot leaf mats have been excavated at one locality in the Albino (Appendix I). Unfortunately, work on these fos- sils is incomplete and the floristics of this site remain poorly known. Figure 38 shows a spec- imen from the Albino that appears to be a pin- nately compound leaf, possibly of rosid affinity, but the venation is too poorly preserved to con- firm such a determination. The uppermost Albian Aspen.flora (Brown, 1933a, 1933b) from southwest Wyoming (Figs. l, 2, Table 3) comprises nine species of angio- sperms. No species from the Aspen are known from the older floras ofthe region, although some are recorded from the nearby and slightly youn- ger Frontier Formation (Table 3). Notable in the flora is “Liquidambar” fontanella Brown (Fig. 39), which possesses glandular teeth similar to those of Recent Liquidambar and Altingia. ''Sas- safras" bradleyi Brown (Fig. 40) is preserved in a semicoalified state, which makes higher order ° < CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 735 venation difficult to distinguish. This species shows smooth-margined, palinactinodromous leaves with three lobes such as in extant Lau- raceae and certain palmate Dilleniid families. “Sparganium” aspensis Brown (Fig. 41) is be- lieved to be a Hamamelid fructification. Such fossils are most commonly attributed to Plata- noids; however, associated leaves of this form are not present in the Aspen. It is possible that the fructification was produced by Liquidambar fontanella. An advanced Sapindophyll, Sapin- dopsis schultzii Brown, is present in the Aspen, as well as several other species of putative rosid affinity (Table 3). The Cenomanian flora from the Frontier For- mation of southwest Wyoming (Hall, 1845; Knowlton, 1917; Andrews & Pearsall, 1941) is a mixed pteridophyte-angiosperm assemblage preserved in volcanic sediments similar to those which yield the Aspen and Albino floras. The flora is from the lowermost part of the Frontier Formation and is referred to as the Lower Fron- tier flora in Figures 1 and 2 and in Table 3. “Cinnamomum” hesperium Knowlton (Fig. 42) is very similar to Cinnamomophylls from the Middle and Late Albian, and as such is the only archaic element in this flora. Platanophylls are represented by “Aralia” veatchii Knowlton (Fig. 43), which is advanced over earlier mem- bers of the morphotype by its deeply divided lamina. Dewalquea pulchella Knowlton (Fig. 44) is a toothed, palmately compound leaf of uncer- tain affinity. The entire-margined, palmately compound leaf of “Staphylea”? fremontii Knowlton (Fig. 45), which also occurs in the As- pen flora (Table 3), shows well-developed, eu- camptodromous secondary venation such as characterized ma pound leaves are restricted to extant Ranuncu- liidae, Rosidae, and palmate Dilleniidae. Species with and without entire margins can be found in each of these groups. More detailed analysis of the venation and tooth morphology of the pal- mately compound leaves from the mid Creta- — FIGURES 38-47. Pinnately compound leaf, Mowry Form ambar” fontanella Brown, USNM 39146, holotype. — 40. “Sparganium” aspensis Brown, USNM 39140A, holotype. 42-47. F type.—41. Dicotyledonous leaves from the Mowry, Aspen, dd and Dunvegan formations. 38. ation. 39-41. Aspen Formati , USGS locality 8169.—39. “ Liquid- “Sassafras” ‘bradle yi Bue USNM 39144, holo- ontier Formation. —42. “Cin namomum” hesperium Knowlton, USGS locality 6527, USNM re e 43-45. USGS locality 5049.— 43. “Aralia” veatchii Knowlton, portion Seir ie eaf, USN t pulchella Knowlton, m 35255, holot 46, 47. Dunvegan Form boreale Dawson, GSC i y 4205, GSC 5398, holoty d holotype. —44. Dewalquea wlton, USNM 35230, holotype. UN 46. Pseudoaspidonhllum. tilia Hollick, GSC locality 4197. —47. Protophyllum 736 ceous of the region ii ida pulchella, ** Dios- = *Hymenaea" agde Berry, Fauni ie gracile Debey, D. lanceo- latum (Knowlton) Berry, and m fre- montii) is necessary before further conclusions are possible. Far to the north of the floras just mentioned, the Dunvegan flora (Figs. 1, 2; Dawson, 1883; Bell, 1963, 1965) was recovered from coarse flu- vial deposits in marked contrast to the volcan- iclastic southern deposits. The Dunvegan flora, unlike the more southerly floras, contains several leaf genera that indicate considerable affinity with early angiosperm floras (Araliaephyllum, Tro- chodendroides, and “Cinnamomum” ), as well as new forms. Platanophylls (Fig. 46) and Proto- phylls (Fig. 47) are numerically dominant in the Dunvegan assemblage. Entire-margined, pal- mately compound leaves and Dryophylls relate this flora to the Frontier and Aspen floras. The Dunvegan flora contains a suite of large, entire- margined leaves of the Magnoliaephyll morpho- type (Table 3). The Turonian flora (Berry, 1929d) from the upper part of the Frontier Formation (Table 3, Appendix I) comprises cycadophytes, taxodia- ceous conifers, Cornophylls, Pentalobaphylls, and fragments of the laminae of a putative palm to which Berry (1929d) applied the name Sabalites, although the specimens lack evidence of costa- palmate morphology. In addition, several ro- sidlike leaves are reported from the upper Fron- tier (Table 3). SANTONIAN FLORAS The lower Santonian Badheart flora from northeast British Columbia is notable for an abundance of 0 2 a. droides arctica (Heer) B yphus" mcgregorii (Bell)). These occur in elis with the common Upper Cretaceous araucarian co- nifer Geinitzia formosa Heer (Bell, 1963). LOWER CAMPANIAN FLORAS The floristic record from this time rivals that for the Albian. Near equivalent floras are known from the Milk River, Eagle, and Two Medicine formations in Alberta and Montana (Figs. 1, 2). Floristics of the individual floras vary within a context of dominant Hamamelididae. Also im- portant are Cornophylls and Rhamnophylls. Some collections contain an abundance of ar- aucarian, taxodiaceous, and cupressaceous fo- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 liage. The small-leaved aquatic dicot Quereuxia (cf. "Trapa"? microphylla Lesquereux) is found in the Two Medicine. The earliest pinnate palm from the region is present in the Two Medicine. The woody dicotyledonous flora (Bell, 1963, 1965) of the Milk River Formation in southern Alberta (Figs. 1, 2) is preserved in coarse fluvial sandstones and associated sediments. Unusual is the complete absence of Platanophylls from the flora. Other than Trochodendrophylls, there is an overall dearth of Hamamelidids. Trocho- dendrophylls (Trochodendroides arctica and T. dorfii Bell), Cornophylls (** Magnolia"? coalvil- ensis Knowlton and Celastrinites sp.), and Rhamnophylls (** Ficus" trinervis Knowlton) are the most abundant dicotyledonous leaf fossils. The Eagle Formation extends through most of central Montana and northern Wyoming. A number of fossil localities are known, and some collecting has been done (see Appendix I), but our knowledge of the Eagle flora is based at pres- ent on the work of Knowlton (1900) in central Montana, and Bell (1963) in northcentral Mon- tana. I was not able to locate the original collec- tions for either ofthese treatments (see Appendix I), so my discussion is based on the published nts. Bell (1963) recognized five dicotyledons from the Eagle. The composition of the flora is very similar to that of the Milk River with Trocho- dendroides dorfii, Cornophylls, and Rhamno- phylls. In addition, the flora contains a magno- liid-grade leaf “Ficus?” missouriensis Knowlton, which was first described from the Eagle in cen- tral Montana (Knowlton, 1900). Knowlton (1900) described a small, poorly preserved collection from the banks of the Mis- souri River (Appendix I). The flora contains a Platanophyll (*P/latanus"? wardii Knowlton), which seems to be reliable to the family level. A Rhamnophyll (** Ficus” trinervis) and a rosid leaf veins that terminate near the sinuses ofthe teeth, margins with rosid teeth and rosidlike sinus brac- ing, and percurrent, transverse tertiary venation. “Liriodendron” alatum Newberry, a large leaf of probable magnolialean affinity, was also reported by Knowlton (1900). Knowlton's specimen shows little more than secondary venation, and identity with the type from the Amboy Clays (Newberry, 1895) must be considered tentative. A more extensive, well-preserved assemblage 1987] from the lower Campanian Two Medicine For- mation in northern Montana (Figs. 1, 2) is cur- rently under investigation as part of my doctoral program. The flora consists of about 30 species of angiosperms, two conifers, and six ferns, and was recovered in Oilfield Coulee (Appendix I) from a single fluvial system extending in outcrop for two kilometers. It records the vegetation along a coastal plain river. Pinnate palm fronds (Fig. 37) dominate the channel margin facies throughout the outcrop. The fronds are up to two meters long and are incompletely divided into induplicate segments. The fossil palm could be assigned to the form genus Phoenicites (sensu Read & Hickey, 1972) were it not for the absence of spines at the base of the frond. Occasional interbedded paludal fa- cies contain “Trapa” (Quereuxia). Taxodiaceous and cupressaceous conifers are found in the basal lag deposits of paludal facies. Nymphaeites sp. foliage occurs in restricted parts of the outcrop. Levee and other near-channel facies contain a diverse dicotyledonous flora that is dominated by leaves of Hamamelididae. Nonhamamelidid dicot leaves typically comprise about 10% of the nego tonsils: Dicot 43 (Fig. 49) is characterized and weakly percurrent tertiary venation irregularly spaced, eucamptodromous secondary veins that origi- nate almost perpendicular to the midvein and turn upward rather abruptly near the margin. The leaf morphology indicates closest relation- ship to extant Rutales but should be considered tentative since Rutales have pinnately com- flora have affinities to Dilleniidae on the basis of tooth morphology. Two species (Dicots 57 and 19, unfigured) of entire-margined, imperfectly actinodromous leaves with thin, percurrent, ter- tiary veins arranged in concentric series relative to the base of the leaf indicate palmate dilleniids. A complex of four species (three of which are illustrated in Figs. 51-53) are related by their c oranthoid teeth and form an important and verse-percurrent. The leaf shows closest affinity with Trochodendrales but cannot be accom- CRABTREE— NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 737 modated in either of the extant genera Trocho- dendron or Tetracentron. Another leaf in this complex (Dicot 12, not figured) more closely re- sembles extant Trochodendron, especially in de- tails of tertiary and quaternary venation. Dicot 53 (Fig. 52) is a small, broadly elliptic leaf with pinnate venation and with secondary veins clus- tered at the base. Dicot 53 shares with extant Cercidiphyllum such features as shape, orienta- tion and behavior of tertiary and higher order venation, and chloranthoid teeth with a tripod- like arrangement of accessory veins. Cercidi- phyllum differs from the fossil in its well-devel- oped actinodromous venation and greater tendency for tertiary veins to bifurcate. Dicot 53 is sufficiently like extant Cercidiphyllum that it can be placed into Cercidiphyllales. Dicot 50 (Fig. 53) has pinnate venation, craspedodromous sec- ondary veins with the basal pair strengthened, and acute chloranthoid teeth. Tooth morphology most closely approaches extant Atherosperma- taceae; however, the craspedodromous second- aries and strong basal apd veins of eke 50 make it a problematic species intermediat in morphology between ae cnn and Trochodendrales Dicot 25 (Fig. 54) has a morphology close to some extant Hamamelidaceae, notably Recent Fortunearia. The fossil has strong craspedo- dromous secondary veins that end in scalloped teeth with spinose tips; the tertiary and quater- nary venation is strong and branched orthogo- nally. Dicot 25 is the most abundant and wide- nis of the To Memomie dicot taxa and ob abl 1 4 : 41 Ec leaves. Another hamamelid (Dicot 36, not figured) shows leaf morphological fea- tures convergent with extant Hamamelis and Fothergilla. Second in overall abundance at Oil- field Coulee are two species of Platanaceae (Figs. 56, 57). Both are large, rhombic leaves with pal- inactinodromous venation, pectinal veins, nip- ple-shaped teeth, and orthogonal tertiary and quaternary venation Much of the diverse Oilfield Coulee flora can- not be treated fully here. One leaf species (Dicot 17, not figured) is entirely consistent with the leaf morphology of extant Menispermaceae. Sev- eral taxa of magnoliid-grade leaf morphology are present (e.g., Fig. 55). Other poorly known collections from the Two Medicine exist (Appendix I). Because the Two Medicine Formation extends upsection to the top of the Campanian, and I have not revisited [VoL. 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1987] the collecting localities, it is difficult to assess the age of these collections. One specimen is illus- trated (Fig. 47) from these localities. The leaf fits the Cinnamomophyll morphotype and may rep- resent Laurales based on the presence of basi- laminar secondary veins, basal secondaries at a lower angle than superadjacent secondaries, ad- medial orientation of tertiary venation, subpar- allel and irregularly branched quaternary vena- tion, and possible mesophyll secretory bodies. MIDDLE CAMPANIAN FLORAS Floras are known from the Judith River For- mation in central Montana (Knowlton, 1905) and the Oldman Formation in southern Alberta (Penhallow, 1908; Bell, 1963, 1965). The two floras were recovered from fluvial sandstones and exhibit a similar composition. Simple leaves with errate margins and pinnate, craspedodromous venation allied to Hamamelidaceae predomi- nate. There is some evidence for the presence of Trochodendrophylls (Hickey, 1984). Diverse Platanophylls, including “Platanus” affinis Les- quereux, are found in the Oldman flora. “Arto- carpus" sp. (Bell, 1965) from the Oldman is a pinnately compound leaf, the leaflets of which are pinnately veined with eucamptodromous secondary veins and many intersecondaries. This morphology relates it to rosid groups, including Sapindales, Rutales, and Rosales. Dawson (1886, 1887) described plants from the Belly River se- ries, which is equivalent to the Judith River- Oldman floras. His species appear unusually ad- vanced for the Campanian, an attribute Knowl- ton (1900) ascribed to stratigraphic mixing of the collections. Ramanujan (1972) reported on per- 1977), and the Livingston flora (Knowlton, 1893) CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 739 from southcentral Montana, the Adaville flora from southwest Wyoming (Dorf, 1955), and the Point of Rocks flora from southcentral Wyoming (Brown, 1956; Dorf, 1955; Knowlton, 1900; Les- quereux, 1878; Newberry, 1898; Ward, 1885, 1887) SUMMARY During an approximate three million year pe- riod of the Middle and Late Albian, angiosperms displaced gymnosperms as the dominant low- land vegetation of the NRM region. This oc- curred shortly after the entry of angiosperms into the region. Upland vegetation retained a pre- dominantly gymnospermous composition. Pte- ridophytes remained important through this period, although composition changed. Gink- gophytes, nm and Pteridosperms de- creased dramatically. Although "m pec Um of angiosperms can be seen as the essential agent of change, several en- vironmental factors correlate with the floristic changes in the Albian. The Cretaceous epeiric seaway expanded across the mid-continent for the first time during the Albian. Climatic sea- sonality may have increased due to increased orogenic ee in the Cordillera and increased temperatur arly angiosperms with low rank leaves such as Rogersia and Ficophyllum are not recognized in the NRM region. The Middle Albian flora is closely allied to time equivalent floras at lower latitudes in North America. The similarity is most pronounced at the level of genus, although a few species appear to have ranged across North America during this time. As in the Potomac flora, Sapindophylls and Platanophylls form a conspicuous element. Other groups, such as the Cinnamomophylls and Pentalobaphylls, appear to have achieved prominence earlier in the NR region than elsewhere in North America and im- part a uniqueness to the regional flora. Leaf morphologies indicate that early mem- , FIGURES 48-58. Angiosperm megafossils from the Two Medicine Formation. Specimen numbers correspond to the research collection housed at the Department of Botany, University of Montana. 48-57. “Oilfield Coulee” locality.— 48. Pinnate palm, 23A/M1/003.— 49. Rutales?, Dicot 43, 564/43/001.— 50. Dilleniidae, Dicot 1, 40A/01/001.—51. Trochodendrales, Dicot 11, 50A/11/001.— 52. Cercidiphyllales, Dicot 53, 57A/53/001.— 53. Indeterminate leaf with craspedodrome secondaries, strong basal secondary veins, and chloranthoid teeth, Dicot 50, 55A/50/001.— 54. Hamamelidaceae, Dicot 25, S174A/25/010.— 55. Magnoliid-grade leaf of uncertain af- finity, Dicot 27, 39B/27/002. 56, 57. Platanaceae. — 56. Dicot 32, HPC/32/022.— 57. Dicot 6, 24A/06/021.— 58. Laurales, USGS locality 6015. 740 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 HAMAMELIDIDAE PLATANALES CONIACIAN (PLATANALES) COMPLEX ">>> Pentolobophylls Gianna) LATE MIDDLE ALBIAN UPPERMOST ALBIAN m a (HAMAMELIOALES 7 GAPINDALES) K GRADE COMPLEX EUPHORBIALE PALMATE DILLENIIDAE @| T CAMPANIAN FIGURES 59-62. Changing diversity and floristic affinity of the pre-M a: e northern Rocky Mountain region. Occurrences of taxa are based on interpretation of leaf biduo ex iod in the following cases: Chloranthales (Clavatipollenites), Hippuridales (Retitricolpites microreticulatus Brenner), Fagales (Tricolporopollenites sp.), and Euphorbiales (Erdtmanipollis). Muller (1981) accepted these occurrences ). as indicative of the presence of the taxa (see also Norris et al., bers of several Recent higher taxonomic cate- gories of angiosperms existed in the Albian flora (Figs. 59, 60). Sapindophylls appear to be allied to primitive Rosidae. Hamamelididae is well represented by Platanophylls, Protophylls, and Trochodendrophylls. Magnoliidae allied to Laurales, Magnoliales, Chloranthales, and Nym- phaeales are probable. Monocotyledons are not apparent in the megaflora of the NRM region until the Late Cretaceous. The early angiosperm flora includes several prominent groups with no clear relationship to Recent taxa (Table 1). Pentalobaphylls exhibit Cornophyll, and Magnoliaephyll are informal categories based on leaf morphology. The names do not imply relationship to any modern group, merely convergence in leaf morphology with 1975; Rouse et al., modern forms. Much of the diversity of ined dicotyledons is manifest in these morphotype New higher level taxa will probably ve s to allow for their adequate classificat Cenomanian floras in the region retain some older elements (Platanophylls, Protophylls, Cin- namomophylls, and Trochodendrophylls are worthy of mention), but on the whole are more modern in aspect, contain higher rank leaves, and are more diverse than Albian floras. Latest Albian and Cenomanian floras from lower lati- tudes within the region contain Dryophylls and Dewalquea. Post-Cenomanian floras continue to diversify, attaining levels of advancement consistent with d gales), palmate Dilleniidae, and Rosidae are a 1987] persistent part of the flora. Magnoliidae (Lau- rales, Chloranthales, Nymphaeales, and Mag- noliales) appear as members of individual floras. Ranunculiidae (Menispermaceae) is recognized from the lower Campanian in Montana. Pinnate palms date from the Lower Campanian. The group is historically associated with stream mar- gin environments LITERATURE CITED ANDREWS, H. N. 1948. Fossil tree ferns of Idaho. Archaeology 1: 190-195. E KERN. 1947. The Idaho tempskyas and associated fossils. Ann. Missouri Bot. Gard. 34: 119-183. . S. PEARsALL. 1941. On the flora of the Frontier Formation of southwestern Wyoming. Ann. Missouri Bot. Gard. 28: 165-193. AsH, J. 1985. Growth rings and PL adiu in Agathis vitiensis (Seemann) Benth. and Hook. f. ex Drake in Fiji. Austral. J. Bot. 33: 81-88. AsH, S. R. & C. B. READ. 1976. North American species of Tempskya and their stratigraphic sig- nificance. Profess. Pap. U.S. Geol. Surv. 874: 1- AXELROD, D. I. 1959. Poleward migration of early angiosperm flora. Science 130: 203-207. 1970. Mesozoic paleogeography and early angiosperm history. Bot. Rev. 36: 277-31 BARKSDALE, J. D. 1975. Geology of the Methow ley, Okanogan County, tos dr Bull. Was State Div. Geol. Earth Sci. 68: BASINGER, J. F. & D. L. 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Numbers i in paren- used in Figures 2 and 3 and in Tables 2 and 3. Chrono- stratigraphic references are placed in parent theses i im- cality information absent or sketchy. All others are referenced to the publications that provide locality specifications. Institutions where collections are housed are indicated. ALBINO (1). Upper Upper Albian (Roberts, 1972). I made a small collection from leaf mats in the Albino Member of the Mowry Formation along Beaver Creek, Gallatin County, Montana. This locality is in pink and white volcaniclastic siltstones of the Albino Member of the — = rmation in the NW bs od 1⁄4, sec. 18, T.7S,R.4 Department of Botany, G of Montana. jio extensive fern ora is also known from the Albino Member (Crabtree, 1983) N (2). Upper Upper Albian (Cobban & Reeside, 19522; Nichols & Jacobson, 1982). This e is from the predominantly marine Aspen gon tion at USGS locality 8168 located in the NW 4, sec. 6, T 24 N, R. 115 b n County, Wyom here, and the SUE occur 125 feet from the top (Brown, ines Collections are housed at the U.S. National um. Lower Santonian Pa 1961, 1963; Bell, 1963). Collections are from the Badheart For- mation in the vicinity of Belcourt m eastcentral British Columbia. Collections are housed at the Geo logical Survey of Canada BEAVER MINES (4). Lower Middle Albian (Bell, 1956; Mellon, 1967; Mellon & Wall, 1963; Stott, 1968; Singh, 1971). pix are from the Beaver Mines For- mation in the vicinity of Crowsnest Pass, southwest Alberta. Collections are housed at the Geological Sur- vey of Canada BADHEART (3). BLACKLEAF (5, 19, 20). Lower Upper Albian (Cannon, 1966; Rice & Cobban, 1977). Collections come from several USGS localities in the Vaughn Member of the Blackleaf Formation in northwestern Montana. Col- 746 lections are described under sh headings which cor- The single locality i is in the NW Ys, SE 4, NE 4, sec. 13, T about 5 km north of Great Falls, TM road Montana. Col- lections are “itt at the Department of Botany, Uni- versity of Montan Summit (19). Collecting Ege are scattered for ab s Pass alon ties 5984, 5985, 5986, 6007, and 9439 in Glacier Coun- ty, Montana. A separate area exists on the west side of the pass in the S '2, sec. 1, T. 29 N, R. 14 W Flathead County, Montana. Collections are housed at the U.S. National Museum with the exception of a small col- lection from '2 mile west of Marias . which is at n rsity of Montana. USGS locality 9437 from the SW 1⁄4, SE “4, NE 4, sec. 36, T. 22 N, R. 13 W on the north side of the Sun River, Teton County, Montana. Col- lections are housed at the U.S. National Museum COMMOTION (6a, 6b, 6c). Collections are from three members of the Commotion Formation and are housed at the Redpath Museum and the Geological Survey of Canada. Each member is treated separately below. Lower Gates (6a). Lower Middle Albian (Bell, 1956; Mellon et al., 1963; Stott, 1960, 1968; Singh, 1971). Collections are from the lower Gates Member in the vicinity of Pine River, northeast British Columbia. Upper Gates (6b). Middle Middle Albian es 1956; Mellon et al., 1963; Stott, 1960, 1968; Singh, 1971). Collections are from the upper part of the Gates Member along Belcourt Ridge, eastcentral British Co- lumbia. Boulder Creek (6c). Middle Middle Albian (Bell, 1956; Stott, 1968; Singh, 1971). Collections are from the Boulder Creek Member in the vicinity of Pine Riv- er, northeast British Columbia veteri (7. Lower Upper Albian (Bell, 1956; Mellon & Wall, 1963; "Mellon: 1967). C ollections are from the Mill Creek Formation in the vi ange in southwest Alberta. ee are housed at the Geological Survey of Cana DUNVEGAN Cenomanian (Bell, 1963; Stott, 1960, housed at the Redpath Museum and the Geological Survey of Canada EAGLE (9). Lower Campanian (Cobban & Reeside, 1952a; Russell, 1970). The Eagle flora reported by Knowlton (1900) was collected by Lester Ward in 1883. Knowlton’s ipe du several of them types, are miss- ing from the U.S. National Museum I attem wee to relocate Ward's site and found a small assemblage from along the Missouri River below Rat- tlesnake Coulee about 12 km south of nie Banks, Chouteau County, Montana, in section 24, T. 26 N, R. 12 E. This paca is at the Department A en University of M Bell (1963) E a flora from Buckley and Red ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Coulees, Toole County, Montana, in the vicinity of the International Boundary south of the town of Milk Riv- er, Alberta (see Evans, 1930). I was unable to locate this —— at the Geological Survey of Canada. In recent years, collecting has been carried out at USGS “locality 7633 from the Eagle Formation by L. W. His eve are housed at the U. S. National Museu FALL RIVER (10). Upper Albian Reds pers. obs.). Whileat the U.S. National Museum I examined several specimens from USGS locality 7312 i in the general collection. The ledger of field collection sites indicates that these plants were found in the Fall River Sand- stone near Pine Creek, Wyoming. This is probably the Pine Creek mentioned by Ward (1899). As best I can Judae from Ward’s descriptions, the fossils are from R. 62 W, T. 55 N in Crook County, Wyoming, from a level 60 feet below the Dakota ia This locality could be as old as Middle Albia FRONTIER (lla, 11b). Two main gaa levels in the Frontier Formation have yielded plant mega- fossils. These are treated separately below. Lower Frontier(1la) Lower omanian (C fus & Reeside, 1952b; Nichols & Jacobson, 1982). M — were made from nis lower Frontier For. matio eldwork Hall (1845) seco on the earliest collections from the Fr Char emo County, Wyoming, in the S '4, sec. 29, T. 19 N, R. 116 .S. Ge ological Survey UM from this area and the adjacent sec 116 W include USGS field localities 8676, n i and 5049. Knowlton (1917) reported on collections primarily from locality 5049. Subsequent collections from this and nearby localities were reported by Andrews & Pearsall (1941). All of these localities are from the same strati- graphic level about 1,200 feet below the Oyster Ridge Sandstone and are considered to be of lower Ceno- manian age. Hall's original collection no longer exists. Knowlton's published collections are housed at the U.S. National Museum versity of Connecticut. Scania collections are at the U.S. National Museu Upper Frontier (1 Ib). Turonian (Nichols & Jacob- son, 1982). Collections from the Frontier near Lander, Wyoming (Berry, 1929d) are from well above the Oys- ter Ridge Sandstone. These collections are housed at the U.S. National Museum. JACKASS MOUNTAIN (12). Middle Ped eg ee 1956). Collections are from localities th Mountain Formation in southcentral British ciens and are housed at the Geological Survey of Canada (= £e JUDITH RIVER-OLDMAN (13). Middle Campanian (Cobban & Reeside, 1952a; Russell, 1970). The col- lections from the Judith River Formation described by Knowlton (1905) were collected at USGS locality 3272 alon 4 km east of old Fort a locality information for the Oldman Formation i southern Alberta, but see Ramanujan (1972) and Bell 1987] (1965). Judith iia collections are housed at the U.S. National Muse KINGSVALE (14). Middle pipe debe (Bell, 1956). From localities in the Kin mation in south- central British Columbia. C ollections are housed at the Geological A of Cana ir IVER (15). pedido vene (Bell, 1963; Rus- l, 1970). Localities = in the iver Formation ie Milk River, s Alberta (Bell 1963). Collec- Gols are housed at e Geological Survey of Canada. MILL CREEK (16). Lower Upper Albian (Mellon & Wall, 1963; Mellon, e Collections are from the Mill Creek Forma ear Crowsnest Pass and north along tion the foothills in Glad Alberta. They are housed at the Geological Survey of Canada Middle Upper Albian (Barksdale, the Pasayten Fo tion in PASAYTEN (17 at the Geological Survey of Canada. specim d. A - is at the Department of Botany, University of Montana PRE-MUDDY (18). Middle Middle Albian (Roberts, 1972). S. Vuke made this collection from beds below the Muddy Sandstone V Buck Creek, Gallatin County, Montana, in T. 8 S, R. 3 E. It is housed at the Department of Botany, University of Montana. TWO MEDICINE (21). Lower Campanian (Russell, 1970; Lorenz, 1981). Collections are from the lower the Two Medicine Formation in “Oilfield Coulee” about 20 m above the contact with the Virgelle Sandstone in CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS art of 747 the SW '4, sec. 18, T. 32 N, R. 5 W about 6 km south of Cut Bank, Glacier County, Montana. These are housed at the Department of Botany, University of Montana. Unstudied collections from USGS localities 6009, 6010, 6011, 6013, 6015, 6129, 6388, 6390, 8574, and 9449 in Glacier County, Montana, are housed at the U.S. National Museum. WAYAN (22). Upper Upper Albian or Lower Ceno- manian (Cobban & Reeside, 1952a; Moritz, 1953). During the course of many years, an amateur collector named Thomas Henry made a very large collection of in the vicinity of Wayan, Caribou County, Idaho. The exact locality (-ies) is not known, but appears to have n in sec. 24, T. 5 S, R. 43 E and secs. 19 and 20, 58, » x E. I could not . the sites during my own visits to the area in 1983. Most of the Wayan in AURA 1S ae at the Department "e Geology, Ida- ho State University. The specimens figured in this re- port (Figs. 21—23) are Dr in the Department of Botany, University of Mont WINT P (23). Upper Albian or Cenomanian (pend e 1975; R. Rau, pers. = Collections housed at the U.S. National Museum and reputed to be from the hippies Formation are probably Paleo- cene in age (J. Wolfe, pers. comm.). Material in T report was collected by myself, R. Rau, and J. Robis from the type locality of the Winthrop Formation near the Boesel Ranch along the Methow River road in the NW 4, NE 4, SE %4, sec. 14, T. 35 N, R. 20 E, Okanogan County, Washington. This collection is housed at the Department of Botany, University of Montana. EOCENE AND OLIGOCENE FLORAS AND VEGETATION OF THE ROCKY MOUNTAINS! Scott L. WING? ABSTRACT 1 T ^A (1); Yes of major evolutionary, biogeographic, and vegetational UE ti changes in Sepie plants. From a floristic perspective three major trends characterize this period cky Mo in the Ro ntain region: 1) development of more distinct phytogeographic provinces from a relatively Cana s Paleocene holarctic flora, 2) the early diversification in Eocene upland areas d milies, an ) the first appearances of many extant angiosperm Pale evergreen forest to high northern latitudes (60°N), possibly ` the greatest geographic coverage suc ch vegetation everachieved. Thi more open vegetation of lower stature in many areas. These floristic and vegetational trends ` were of great conseq uence for angiosperm evolution and biogeography. They were also significant in creating the ecological milieu for the evolution of early and mid Tertiary animals. The Paleogene rocks and fossils of the Rocky Mountains and adjacent areas provide an excep- tional window on the flora and vegetation of the time and place. Eocene and Oligocene rocks cov- er some 40% of the surface of the state of Wy- oming (106,000 km?), and their total outcrop area in the Rocky Mountain region is in the range of 300,000? km (Fig. 1). Over much of this vast outcrop, weathering has exposed the strata, mak- ing them available for paleontological and geo- logical study. Eocene-Oligocene deposits from a variety of fluvial, lacustrine, and volcanically in- fluenced settings have produced tens of thou- sands of fossils collected at hundreds of localities. These Paleogene sequences have the potential to yield one of the most complete and geographi- cally extensive records of evolutionary and eco- logical change in a terrestrial biota. A century or more of paleontological and geological research has only begun to explore this potential, in part because of the vast area and great thickness of the sedimentary sequences. This paper is intended to serve three goals. First, to set out the basic data on the location, stratigraphic positions, and relative ages of the fossil floras. Second, to summarize published work on Eocene and Oligocene flora and vege- tation of the region and discuss previous hy- potheses in light of current data. Third, to suggest new directions for future research aimed at a regional understanding of Paleogene floras that takes full advantage of the potential of such a widespread and prolific fossil record. GEOLOGIC SETTING The Rocky Mountain region is geologically di- verse, and its Cenozoic history has been reviewed in a number of recent volumes (e.g., Robinson, 1972; Curtis, 1975; Flores & ise 1985). Eocene and Oligocene plant fossils are best known from the large intermontane basin fill deposits of the eastern Rocky Mountain area and from the primarily volcanic deposits of western Mon- tana, northwestern Wyoming, Idaho, Oregon, and Washington. The separate depositional histories of these two areas are important for interpreting historical change in their floras. The intermontane basins of the eastern Rocky Mountain area (Fig. 1) began to develop in the way. Uplift of local, initially east-west trending, mountain ranges during the latest Cretaceous- Paleocene began to divide the foreland basin into structurally, topographically, and sedimentolog- ically discrete units. Consequently, each of the present intermontane basins has a separate Ce- nozoic history, yet some general trends have af- fected most or all of the basins. Fluvial to paludal environments were preva- l inen to Jack A. Wolfe, Ralph Taggart, and George Rogers for review, and to Mary Parrish for drafting the figur : on PEN of Paleobiology, MRC 164, Natural History Building, Smithsonian Institution, Washington, S.A. D.C. 20560, U.S ANN. MISSOURI Bor. GARD. 74: 748-784. 1987. 1987] 120 * `< x CARN P VOLCANIC FIELD E Eocene BASIN FILL OLIGOCENE ll EOCENE/OLIGOCENE VOLCANICS FIGURE 1. lent throughout most of the area in the Paleocene and were environments of deposition for coals in the Powder River, Bighorn, Green River, and other basins. Fresh or brackish water lakes formed in the Uinta, Wind River, and Bighorn basins during the latter half of the Paleocene (Johnson, 1985; Keefer, 1965; Yuretich et al., 1984). Be- ginning in the Clarkforkian? and continuing into 3 Through much of this paper, North American pro- vincial land m are known to be door VA i ede ar. out western North America (Flynn 84) and because it is easier to Pieces floras S. m mam- malian sites than with the type age/stages of Europe. Correlation of NALMA with epochal boundaries and time are given in Figure 2. WING —EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS BEARPAW VOLCANIC* YELLOWSTONE/ABSAROKA VOLCANIC FIELD BIGHORN 234,05. ASIN - L POWDER RIVER BASINI e WIND RIVER GREAT DIVIDE € Il BASIN Xe E ANDwASH BASIN u — C Ma PARK yea PICEANCE CREEK BASIN) to t... -NINE MILE eg VOLCANIC FIELD SAN JUAN BASIN ok] NM N Outcrop area of Eocene and Oligocene rocks in the greater Rocky Mountain area the Wasatchian, paludal deposition of organic- rich sediment waned in most of the basins, and alluvial floodplain sediments began to be mod- ified by oxidizing pedogenic processes (Bown 1979; Bown & Kraus, 1981). The onset of this kind of pedogenic modification was probably controlled by a combination oflocal tectonic and physiographic factors, as well as by regional cli- matic change (Wing & Bown, 1985 In the later Wasatchian and Bridgerian, large lakes again developed in many ofthe basins (Big- horn, Green River, Uinta-Piceance). In at least some cases this appears to reflect hydraulic clo- sure of the basin rather than climatic change (Johnson, 1985), and the lakes of the Green Riv- er and Uinta-Piceance Basins are known to have been saline during much of their later histories. 749 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 RRENEEEESEEZSEXEEREEXEEXGEEEM. Ts * Ë = O C E N E ‘OLIGOCENE Ë [ EARLY MID AT EARL | . ^ TRETA | srIoceRIAN | UINTAN | | DUCHESNEAN Ë E ~ ARIKAREEAN | E 2. Correspondence of North American Land Mammal Ages (NALMA) with epochal and stage ). FIG efie pr (after Berggren et al., 198 Much ofthe sediment filling these middle Eocene lakes and forming coeval fluvial deposits was derived from newly active volcanic fields in west- ern Wyoming and Idaho During the later Bridgerian and Uintan most of the intermontane basins began to fill with sed- iment, and during the Duschesnean and Chad- ronian these sediments began to lap outward onto the tops of the bordering ranges. During the Oli- gocene, volcanically derived sediment continued accumulating at relatively slower rates in the eastern Rocky Mountains and Great Plains. Oli- gocene formations over much of the northern Rocky Mountains and Great Plains are typically thin, widespread units that were probably de- posited on a surface of low relief. The history of the volcanic centers in the west- ern part of the northern Rocky Mountains is largely independent of that of the intermontane the Elkhorn volcanic field) beginning in the latest Cretaceous and throughout the Paleocene (Roberts, 1972). How- ever, the greatest pulse of volcanic activity began in the late Wasatchian to Bridgerian and contin- ued through to the end of the Eocene From about 53 to 38 Ma there was volcanism across much of western Montana (Bearpaw, Gar- net Range, Lowland Creek, Gallatin, and Bea- verhead Canyon fields), northwestern Wyoming (Yellowstone—Absaroka Field), and Idaho (Chal- lis Field), and in the Clarno area of Oregon, the Republic graben of northern Washington, and southern British Columbia (Chadwick, 1985). Although the intrusive and sedimentary rocks m, it is likely that most of their original extent has been removed by erosion. Armstrong (1974) suggested that the Challis volcanic field once covered much of Ida- ho The extent and thickness of these broadly co- eval volcanic deposits in the northern Rocky Mountain area indicate the region may have been a rugged, but more or less connected, volcanic highland during much of the Eocene (Axelrod, 1968; Fritz & Harrison, 1985). The prevalence of volcanic conglomerates and mudflows, and analogy with present day volcanic terrains, sug- gest strong topographic relief, which Fritz & Har- rison (1985) estimated at 1,000-2,500 m differ- ence between adjacent valleys and peaks. Following the end of the Eocene, heavy vol- canic activity shifted southward to southern Col- orado, exico, Arizona, and west Texas, and extensional tectonics began to affect the northern Rocky Mountains (Chadwick, 1985). A number of small, fault-bounded basins devel- oped in western Montana during the Late Eocene and Oligocene. These basins filled with primarily fluvial and lacustrine deposits that were derived from both local sources and volcanics as far away as the Cascade Range of Oregon (Fields et al., 1985) The geological events summarized above are highly relevant to the floristic and vegetational history of western North America. The devel- opment of a volcanic highland region in the northern Rocky Mountain region during the Eocene had several major effects on plant life, including: 1) dividing lowland areas that for- merly had been continuous from the West Coast to the eastern Rocky Mountain region, 2) reduc- ing the flow of Pacific-influenced air into the in- terior of North America, and 3) changing soil and groundwater conditions by the influx of air- borne volcanic ash. Prior to and during the onset of these effects the low-lying area in the eastern Rocky Mountains was itself broken into a series of topographically isolated and somewhat cli- matically different basins. Thus the overall effect of geological events during the Eocene and Oli- gocene was fragmentation of a relatively ho- g both north-south and mogeneous ] Fr east—west axes. DISTRIBUTION OF FOSSIL FLORAS NI Tm a 1 : A n of Tertiary plant localities in western North America. The list of localities given here (Ap- pendix I) represents an attempt to bring together 1987] the bulk of Eocene and Oligocene localities of the greater Rocky Mountain m I estimate that this list is 60-80% compl The locality information was m from a number of published and unpublished sources. Previous works listing a significant number of localities include those by Axelrod (1966a, 1966b, 1968), Axelrod & Raven (1985), Brown (1937), Hickey (1977), Leopold & MacGinitie (1972), MacGinitie (1941, 1953, 1969, 1974), and Wolfe (1981, 1985). Unpublished locality information was derived from collections at the U.S. National Museum of Natural History (including those of the USNM, the U.S. Geological Survey, and some formerly belonging to Princeton University), and from collections at the Museum of Paleontology (University of California, Berkeley) and the Yale Peabody Museum. Not all of the 240 “localities” compiled here are strictly equivalent. Some floras commonly considered as units actually consist ofthe summed floral lists of a number of separate quarry sites (e.g., the Ruby paper shale flora of Becker, 1961, was collected at 1 5 quarries), whereas others were derived from a single excavation (e.g., Copper Basin flora of Axelrod, 1966a). Most of the listed localities are individual quarry sites, but for some floras listed in the literature it was not possible to determine how many sites contributed to the flora. Criteria used in correlating and determining the ages of floral localities included mammalian biostratigraphy, lithostratigraphic equivalence, magnetic polarity stratigraphy, and radiometric dates. In order to avoid circularity, floras were not correlated on the basis of their taxonomic composition or physiognomic characteristics. The Tertiary time scale used here is that of Berggren et al. (1985). Chronological boundaries for North American Land Mammal Ages are from West et al. (in press), and Prothero (1985a, 1985b). There is controversy over the precise age of the Eocene/ Oligocene boundary. The date used here (36.6 Ma) is the most recent opinion, but many authors (e.g., Wolfe, this volume) use a date of about 34 Ma. Radiometric dates from older publications have been corrected for new constants using the tables published by Dalrymple (1979). Fossil floras of Eocene and Oligocene age have a highly uneven geographic and stratigraphic dis- tribution. Stratigraphically, the Wasatchian and Bridgerian are by far the best-represented time intervals, with the number of floras diminishing sharply through the Uintan and Duschesnean, WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS 751 and into the Oligocene (Fig. 3). Geographically, the northern Rocky Mountain region is far better sampled than the southern region (Fig. 4). These inequities in distribution result from at least four factors. 1) Bias of the author: my work has cen- tered on the lower Eocene of Wyoming, so I am particularly aware of these localities. 2) Wa- satchian, Bridgerian, and early Uintan sections are thicker and more widely exposed than later Eocene and Oligocene rocks. 3) Later Eocene and Oligocene rocks are generally less fossiliferous than early Eocene rocks, probably reflecting less favorable conditions for plant preservation as- sociated with a drying climate in the Western Interior. 4) Collecting has been more intense in some areas than in others. In addition to the uneven distribution of floras in space and time, there are also changes through time in environments of deposition. Nearly all Wasatchian plant assemblages were recovered from rocks deposited in fluvial or fluvial/paludal settings. Although some Bridgerian and later flo- ras are from fluvial deposits, the best known (e.g., Green River, Florissant) are from lacustrine sed- iments. Floral samples drawn from fluvial en- vironments represent the flora differently than do those from lacustrine environments. Fluvial assemblages, particularly those depos- ited on low-energy flood basins, are largely de- rived from local vegetation (Scheihing & Pfef- ferkorn, 1984; Spicer & Greer, 1986; Burnham, in press). This results in considerable spatial het- erogeneity in the fossil assemblage that probably reflects original variation in the local vegetation (Hickey, 1980; Wing, 1980, 1984) By contrast, lakes receive input from sur- rounding lakeshore vegetation and from inflow- ing rivers (Spicer, 1981). These different ele- ments may be mixed to various degrees and scattered across the lake bottom. Although la- custrine assemblages are overwhelmingly dom- inated by plant parts derived from nearby vege- tation (Drake & Burrows, 1980; Spicer & Wolfe, 1987), they tend not to preserve in situ variation in vegetation. They may, however, preserve more of the species that were present in the regional flora. As a result of being derived from a larger source area and of predepositional irai and mixing, l diverse than fluvial assemblages, udi specimens are more evenly distributed among the taxa pres- ent. As a consequence of the inequities in distri- bution of fossil floras and nonequivalence in their bly more ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 || N Yellowstone- | Western Wind Northern & Eastern Williston Basin, | Western Powder | Northern Bighorn ee Hor "^ dua Absaroka River Basin/ Wind River W. WY Y one Basin, WY OLIGOCENE | w = DU CHES- N NEAN N ICHADRONIAN =<.) | ARIKAREEAN | WHITN EYAN ao D o — UINTAN | Lostcasinean [S : LYSI- TEAN WASATCHIAN a 5 m GRAYBULLIAN T North Dakota | River Basin, Volcanics, WY |S. Absarokas, WY T T | | | | | | | | a —— 0? ————2——. 2 — t t 38-42% | | AYCROSS FM. J oY MBR. | e 9 ILV E R 32 w LYSITE "Wen. 9 t | CAMELS WASATCH ©7277 M o @17,18 RY] - 78-109 «mrr»«zmoro 93-8,10-15 PALEOCENE TIFFANIAN| es 016 OW EU oci | | F BEAR DEN — *110-111 i | J 7-7? F + TONGUE RIVER det | | M. MBR. : i : | M MA Vou MBR. A _ ç M t) | ? | E] + | o š | "d | | | \ ^ O "| I Flora discussed á f f i by Hickey, 1980 A | | | 9 | | | i | | | | | FIGURE 3. Correlation chart of Eocene and Oligocene floras from the Rocky Mountains and surrounding areas. Numbers indicate floras listed in Appendix I; letters (A-FF) indicate radiometric dates listed in Table 1. 1987] WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS Green River Washakie Na mam & Great Divide Colorado Utah/Nevada/CA taanoa iir i - — noce Basins, WY Basin, WY & CO SW Montana Washington British Columbia DUM. P| 4 A 1 » S CREEDE S = ; FM. A x 1 @242 | i > O m R | I u di | | | | |] |>| | z | IR M G) B 1374 B ni 1 } 1470 R HE Q z —?—?— To 1 1 ] )UNBAR 1 > r 148 "m ; ae m ^ O aa MEE | om > C ood 4 ‘A L ] ] 2 i| I > JF 1 Oo M rm m M B 4 ° 158-160 1 z : — & €. I <= | 34-Mile Volcanic lower] B U | e03, N 4 > aes ae ue ees] P | iQ | 200-201 è A|~_4 Z | L u } 197-199 e || R š Z | R ly RI|H | 4 | DUCHESNE Y À o c ze RIVER | os | 29 L L zi MIIL m le v1 @ N x ^ | RJIL + 175 4 T P. 166 TT ALLL s — ! R 1 m S 1740 c 23 4 4 = 162-164 e ios] =o v !67 180 ] 4 O AA > =a ALLENBY FM. | z O 1650 ccc | I -b ed L 2 | n =] A IM. 172 po occ. sm AZ m N 1 + > 173 | JFF [5| Z | : I 12? volcanics 1 ° z J 6 L ] š = - WASATCH FM 2 > c 4 12 o S LE —9?9—?--7?-—7 Es > m R E MAIN BODY RDE e] mat ^ DESERT Wiss 22 S. WASATCH FM. | COAL MONT irs F TONGUE, + ` > P EM. x | > WASATCH Jel z e209-223 = ] oe Ed 57.5 : ? r z | p er ail | HË 1 aim 7 O | > | m z |z J 4 > m | Zh FIGURE 3. Continued. 754 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE |. Radiometric dates of Tertiary floras in Figure 3. Age Stratigraphic Unit Publication A, 49.2 + 0.7 Trout Peak Trachyandesite Smedes & Prostka, 1972 B, 50.5 + 1.55 Sepulcher Formation Smedes & Prostka, 1972 C, 43.1 iggins Formation Bown, 1 D, 44.6 Wiggins Formation Bown, 1982 E, 46.7 Wiggins Formation Bown, 1982 F, 47.1 Wiggins Formation Bown, 1982 G, 47.9 + 0.5 Wiggins Formation Bown, 1982 H, 48.5 Wiggins Formation Bown, 1 L 50.4 Aycross Formation MacGinitie, 1974 J, 50.6 Aycross Formation MacGinitie, 1974 K, 42.3 + 1.4 Henry Ranch Member, Wagon Bed Formation Black, 1969 L, 46.2 Wagon Bed Formation, near Badwater, Wyoming Evernden et al., 1964 M, 50.5 Halfway Draw Tuff, Wind River Formation Evernden et al., 1964 N, 50.2 Little Mountain a beu Peak Member, Green River Form Mauger, 1977 O, 46.7 + 0.9 Washakie Formation, pu 664 of Roehler Mauger, 1977 P, 462 several hundred feet above Mahogany Ledge, Parachute e Member, Green River Formation MacGinitie, 1974 Q, 26.5 Creede Formation Axelrod, 1987 R, 34 Antero ign Epis & Chapin, 1975 S, 35 Florissant Formation Epis & Chapin, 1975 T, 36-37 Wall Mountain Tuff Epis & Chapin, 1975 U, 36 "Chicken Creek Formation," 5 feet above highest floral locality Axelrod, 1966b V, 41+1 Deadhorse Tuff Axelrod, 1966b W, 43 “Frost Creek m 1,500 feet below lowest floral loc Axelrod, 1966b X, 31.1 Williams ced ai Fields et al., 1985 Y, 41.1 + 1.6 Salmon ar Fritz & Harrison, 1985 Z, 44.2 + 1.7 almon area t Fritz & Harrison, 1985 AA, 46.3 + 1.0 rhyolite below “Dewey Beds" Fritz & Harrison, 1985 BB, 47.2 + 1.8 basalt above Germer Tuffaceous member Edelman, 1975 CC, 48.0 + 1.0 Latite-andesite Member, Challis Volcanics Edelman, 1975 DD, 47.0 + 1.8 Klondike Mountain Formation Wolfe & Wehr, 1987 EE, 48.2 + 1.6 Klondike Mountain Formation Wolfe & Wehr, 1987 FF, 50-51 Sanpoil Volcanics Wolfe & Wehr, 1987 depositional environments, there are obvious dif- ficulties in interpreting patterns of vegetational and floristic change through time. In spite of these difficulties, a number of durable generalizations have emerged concerning the early Tertiary de- velopment of the Rocky Mountain flora. FLORISTIC AND VEGETATIONAL HISTORY Three major floristic trends are indicated by Eocene-Oligocene fossil assemblages from the Rocky Mountain area: the modernization of the angiosperm flora at the generic level, the breakup of the Paleocene North American province into distinct phytogeographic subregions, and the major diversification of many present-day mi- crothermal lineages (e.g., Rosaceae, Betulaceae, Aceraceae). There were also three major vege- tational periods during the Eocene and Oligo- cene. During the Wasatchian and Bridgerian there appears to have been a major poleward expan- sion of subtropical and paratropical forest types in response to global climatic warming (Wolfe, 1985). Perhaps beginning as early as the Clark- forkian in some areas, and of increasing impor- tance during Bridgerian and later time, there is MacGinitie, 1969). Finally, during the latest 1987] @Fossn @172-181 $19 ekom ° WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS 7-147 151-152 $ 148-150 .. 153-157 1 HAY 119-136 € Havre em s Pe 49 104-111 ° 7), Na 103 71 OR e! LAND * ROCK 30e ie 28 SPRINGS ias 224-229@ gi COLORADO SPRINGS JUNCTION Map position of Eocene and Oligocene floras mentioned in the text and in Figure 3. Numbers dix I. "us ^ n listed in Appen Eocene and Oligocene, vegetation of the Rocky Mountain region became dominated by a mix- ture of conifers and broad-leaved deciduous forms. MODERNIZATION Paleobotanical systematics of the last century and early in this century was based largely on superficial characteristics of leaves. This led to exaggerated estimates of the similiarities be- tween extinct and modern floras. More recent systematic work has focused on detailed com- parisons of leaf venation (e.g., Hickey, 1977) and on greater use of multiple organs (e.g., Man- chester, 1986). One result of such work has been the realization that many Paleocene angiosperms represent extinct genera or intermediates be- tween several related living genera. In contrast, many, if not most, angiosperm remains younger than Late Eocene can be as- signed to extant genera with little ambiguity, al- though sectional or other subgeneric-level affin- ities may be unclear (see Manchester & Crane, 1983 and Wolfe & Tanai, 1987 for well-docu- mented exceptions). The temporal pattern of ap- pearance of extant genera has not been quanti- fied, nor has it been determined the extent to which this modernization reflects evolution within lineages as opposed to extinction of ar- chaic lines and replacement by modern ones. Al- 756 though this generic modernization of angio- sperms is a striking systematic pattern, little has been said about its biological significance. Mod- ernization at the generic level may reflect a true radiation of angiosperms, perhaps in response to changing climatic and topographic conditions in western North America, or it may be an artifact of our retrospective view. That is, the appearance of many modern genera in the Eocene occurred because angiosperms in a number of independent lineages accumulated enough recognizable ge- neric characters to be pigeonholed easily in pres- ent-day categories. This question could be re- solved by measuring and comparing rates of morphological change during the Paleocene and Eocene. PROVINCIALITY The Eocene divergence of Rocky Mountain and West Coast floras and its relationship to East Asian-North American disjunct distributions in a number of living plant groups is probably the most discussed aspect of the Tertiary paleobo- tanical record in western North America (e.g., MacGinitie, 1941; Leopold & MacGinitie, 1972; Wolfe, 1972; Hickey, 1977). Consequently, I will summarize the pattern only briefly. Paleocene floras from across northern North America and to some extent of Europe and Si- beria are relatively homogeneous (Wolfe, 1966). Leaf assemblages typically are of low diversity and dominated by a group of taxa including Ginkgo, Metasequoia, G sb tiu Macgini- tiea na die s") nobilis, “Carya” antiquorum, “Ampelopsis” "acea. and ains. of the Cercidiphyllum complex, among others. Such floras have been reported from Alaska (Wolfe, 1972), many areas of the northern conterminous U.S. (Brown, 1962), the Canadian high Arctic (Hickey et al., 1983), and Greenland (Koch, 1963) Because some of these Holarctic Paleocene taxa been described as having “East Asian" affinities (although they are only East Asian in a modern context). During the Eocene many of these Pa- leocene (“East Asian") forms were eliminated from floras in the eastern Rocky Mountains, pre- sumably as a result of their inability to withstand seasonal dryness (Leopold & MacGinitie, 1972). In contrast, some of the same lineages survived into the Neogene along the West Coast and in ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 the northern intermountain region of North America. At the same time that the “East Asian” forms were being eliminated in the eastern Rocky Mountains, some of the new genera appearing there were "Central American"; that is, their closest living relatives are in the seasonally dry subtropics of Mexico and Central America (MacGinitie, 1969; Leopold & MacGinitie, 1972). This general pattern has been confirmed by ad- ditional work in the intermontane basins of the eastern Rocky Mountains, but three modifica- tions should be noted. Paleocene megafloral samples may be more uniform than actual Paleocene floras because the fossil assemblages have been collected from a limited and similar array of paleoenvironments that tend to have low floral diversity. This would accentuate the impression that a homogeneous floral province was being broken up in the Eocene. The regional extinction oftaxa with East Asian affinities and the appearance oftaxa with Central American affinities began in the Clarkforkian and was well under way in some areas by the mid Wasatchian, considerably earlier than was rec- ognized by L Id & MacGinitie (1972), who lacked floras from the Paleocene-Eocene tran- sition period. Clarkforkian taxa with closest liv- ing relatives in Central America include Chae- toptelea microphylla, Woodwardia gravida, and species of Populus sect. Abaso. By late Graybul- lian or Lysitean time, some taxa with closest liv- ing relatives in East Asia, such as Metasequoia and some members of the Cercidiphyllum com- plex, were already regionally extinct in basinal floras of the eastern Rocky Mountains. The shift from East Asian to Central American affinities in the flora of the eastern Rocky Moun- tains was not a uniform process, and some sig- nificant taxa do not follow the pattern. For in- stance, the genus P/atycarya, now confined to East Asia, did not appear in western North America until the early Wasatchian and achieved maximum abundance throughout the region in the latest Wasatchian and early Bridgerian before going regionally extinct (Leopold & MacGinitie, 1972; Wing & Hickey, 1984). This pattern also appears to hold true for several undescribed species in the Icacinaceae, Flacourtiaceae, and Menispermaceae. Ailanthus, also now endemic to East Asia, appeared in North America during the early to mid Eocene (Chalk Bluffs flora, Green River flora, Rate Homestead flora, MacGinitie, 1941, 1969), and was abundant in Oligocene flo- ras from southwestern Montana (Becker, 1961, 1987] 1969) before going regionally extinct in the later Tertiary. Floristic segregation of northern and southern areas was probably occurring at the same time as the better-documented and better-discussed east-west divergence (Axelrod & Raven, 1985). Unfortunately, few Wasatchian-Bridgerian flo- ras have been reported from the southern Rocky Mountains (Fig. 4). One small Lysitean flora from the San Juan Basin (Tidwell et al., 1981) shows that two taxa common in the Gardnerbuttean and Bridgerian of Wyoming (““Sapindus” den- tonii and Eugenia americana) appeared two to three million years earlier in New Mexico. This suggests the possibility that some taxa were mi- grating northward during the climatic warming of the early Eocene. A similar geographic pattern has been observed in some mammalian species (Beard, pers. comm., 1986). DIVERSIFICATION OF MICROTHERMAL LINEAGES The Wasatchian and Bridgerian floras from the eastern Rocky Mountain region are a mixture of taxa now associated with temperate and sub- tropical to paratropical climates. For instance, mesothermal to microthermal groups like the Betulaceae (A/nus, Paleocarpinus), Cercidiphyl- laceae, and Hamamelidaceae amamelidoi- deae) are frequently associated with members of megathermal groups such as the Icacinaceae (Pa- leophytocrene), Lauraceae (Phoebe), Palmae, and Cyatheaceae (Cnemidaria). In most of these flo- ras the ipsae: elements are not diverse, y be important in terms of 7). younger age from the volcanic areas farther west may be strongly dominated in both abundance and diversity by microthermal taxa. This has been attributed to the relatively high paleoele- vation of these floras (Axelrod, 1966b, 1968; Wolfe & Wehr, 1987). Currently the best known of these floras is from Republic, Washington (Fig. 4; Wolfe & Wehr, 1987, and unpubl.). The vege- tation at Republic is inferred to have been a Mixed Coniferous forest, but the diversity of microther- al angiosperm groups is striking: Hamamel- idoideae, 4 species; Fagaceae, 4 species; Betu- laceae, 5 species; Rosaceae, 19 species; Aceraceae, 7 species (Wolfe & Wehr, 1987; pers. comm., 1987). Although the Copper Basin flora of north- ern Nevada (Axelrod, 1966a) is younger and less diverse, it shows a similar domination by mi- WING —EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS 757 crothermal taxa. Alnus is represented by 2 species, one of which accounts for 85% of the specimens, and there are 3 species of Mahonia, 6 of Rosa- ceae, 3 of Acer, and 3 of Ericaceae. Although upland floras of Paleocene or greater age are not known, it seems possible that the first major diversification of many present-day mi- crothermal lineages took place during the Eocene in upland areas like the volcanic highland of the northwestern United States (Wolfe, 1986, this volume). Some of these groups were represented by relatively generalized species in later Paleo- cene or early Eocene lowland floras. Thus the adjustment of basically megathermal or meso- thermal lines to cooler climates initially may have pers. comm., matic cooling, these upland lineages spread and diversified over much of the Northern Hemi- sphere, whereas their megathermal or meso- thermal sister taxa now have relictual southerly distributions. One particularly good example of this pattern is seen in the genus Populus. Species of the prim- itive section Abaso are common in Clarkforkian, Wasatchian, and Bridgerian floras over much of North America (Eckenwalder, 1977; Wing, 1981), usually in lowland settings, although Populus adamantea does occur in probable Oligocene up- land floras described by Becker (1960, 1972, 1973). The extant species of this section, P. mex- icana, is distributed only in limited parts of northeastern and northwestern Mexico. The ear- liest record of the more advanced sections that account for most of the present-day diversity and distribution of the genus is in the late Eocene Bull Run flora (Axelrod, 1966a). A number of species belonging to the more advanced sections are common in later Tertiary floras, but there is no known fossil record of sect. Abaso poplars ane the early Oligocene. PATTERNS OF VEGETATIONAL CHANGE The general pattern of vegetational change during the Eocene and Oligocene in western North America has been outlined in a number of pub- lications (e.g., Axelrod, 1958, 1968; Axelrod & Raven, 1985; MacGinitie, 1941, 1969; Wolfe, 1971, 1975, 1985). Despite considerable dis- agreement on details, there is overall agreement about the large-scale trends. Broad-leaved ev- ergreen forests were dominant over most of the area during most of the Eocene, with two main 758 exceptions. Some areas in the eastern Rocky Mountains may have been dry enough to create more open, partially deciduous vegetation. Vol- canic activity in the western Rockies during the late Wasatchian, Bridgerian, and Uintan gener- ated uplands where conifers and broad-leaved deciduous taxa became important components of the vegetation. Climatic cooling and drying during the late Eocene and Oligocene brought about increasing dominance of mixed coniferous and broad-leaved deciduous forest. Late Paleocene vegetation in most areas of western North America was broad-leaved ever- green forest with an admixture of deciduous ele- ments (e.g., Golden Valley flora of Hickey, 1977). During the latest Paleocene and early Eocene (Wasatchian) world climates were warming (Sa- vin, 1977; Wolfe, 1971, 1978; Wolfe & Poore, 1982), possibly as a result of elevated levels of atmospheric CO, and associated effects on cir- culation patterns (Owen & Rea, 1985; Rea et al., 1985). At this time evergreen broad-leaved taxa increased in importance within local floras. Floras from the Willwood Formation of north- western Wyoming span most of the Wasatchian and show an upward increase in the number of entire-margined (41% to 52%) and thick-tex- tured leaves, although there are also changes in sedimentary environment that may be causally related to this shift (Wing, 1981). Using data on living vegetation presented by Wolfe (1979), these leaf-margin percentages correspond with Micro- phyllous or Notophyllous Broad-leaved Ever- green Forest. Wolfe (1985) interpreted slightly older (Clarkforkian) floras from the of streamside taxa in Bighorn Basin floras from Fort Union and Willwood formations, or a relatively higher elevation, has given them a “cooler” aspect. I think it more likely that the less entire-margined, presumably more decidu- ous, floras of the Bighorn Basin were under the influence of a seasonally dry climate as early as the middle Clarkforkian. This is also suggested by the development of oxidized soil horizons in floodplain sediments of this age in the Bighorn Basin (Gingerich et al., 1980). A resolution of this apparent conflict would involve detailed comparison of depositional settings of the floras or independent evidence for paleoelevation. Re- gardless of these somewhat different interpreta- tons of vegetation in northern Wyoming, it is apparent that some form of broad-leaved ever- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 green forest extended as far north as 65? during the Wasatchian (Wolfe, 1985). A great many Lostcabinian-aged (late Wa- satchian) fossil localities are known from all over the state of Wyoming and from the Golden Val- ley Formation of western North Dakota (e.g., numbers 1, 2, 9, 19, 27-30, 32-33, 50-71; Fig. 3, Appendix I). These floras are mostly uniform in composition and dominated by one of two species of Platycarya, with important subdom- inants being Alnus, *Dombeya" novi-mundi, “Dalbergia,” Zingiberopsis isonervosa, an species of Icacinaceae, Lauraceae, Magnoliales, and Palmae. Common pteridophytes are Cne- midaria magna, Lygodium kaulfussii, Thelyp- teris weedii, T. iddingsii, and a large-statured species of Equisetum. The similarity in these flo- ras may in part reflect similar environments of deposition occurring ina à number of intermon- tane basins at app en, however, that some range of depositional en- vironments is spanned by these floras, it is likely that the successional vegetation most likely to be preserved in fluvial sediments was truly similar over this large region. This in turn implies the existence of few sharp climatic differences across the area, which is in distinct contrast with the early Bridgerian floras discussed in the next sec- tion. the Sa Tit time. Giv- Vegetation during early Bridgerian time (about 50 Ma). Although the latest Wasatchian and Bridgerian were the times of maximum poleward extent of broad-leaved evergreen forests, vege- tation of the Rocky Mountain region began to differentiate more strongly during this time in- terval. This differentiation is illustrated by com- paring four floras: the Little Mountain flora from the upper Wilkins Peak Member of the Green River Formation in southern Wyoming (23 in Figs. 3 & 4, Appendix I), the Boysen flora from the upper part of the Wind River Formation in the northcentral Wind River Basin (36), the Kis- inger Lakes-Tipperary flora from the Aycross Formation in the western Wind River Basin (38, 42), and the flora of the lower Sepulcher and Lamar River formations in Yellowstone Nation- al Park in northwestern Wyoming (112-118). All four of these floras are approximately 50-51 Ma and correlate with Gardnerbuttean (early Bridg- erian) mammalian faunas Based on floristic affinities, foliar physiogno- my, and sedimentological data, MacGinitie (1969) inferred that the Little Mountain flora was 1987] derived from open woodland vegetation. (MacGinitie used the term “savanna woodlan although he pointed out this might vilsieadinsl imply that grasses played an important role in the vegetation.) More recently, Wolfe (1985) stated that leaf size in the Green River floras is too large for scrub or savanna vegetation and is more consistent with semideciduous subtropical to paratropical forest. The 22 species of dicoty- ledonous leaves from the Little Mountain flora are generally small and thick textured, and a number of the species belong to families or gen- era typical of seasonally dry subtropical vege- tation (e.g., Alchornea, Cardiospermum, Populus sect. Abaso, and a number of microphyllous Le- guminosae; MacGinitie, 1969: 67-68). Recent sedimentological work on the Wilkins Peak Member suggests that deposition took place in and around the margins of a playa lake that lay in an orographic desert basin (Smoot, 1983). The upper part of the Wilkins Peak Member appar- ently represents the maximum transgression of the lake and hence the wettest period during the deposition of this part of the Green River For- mation (Smoot, 1983). The Boysen flora is largely undescribed, al- though it was referred to in a treatment of the Green River flora (MacGinitie, 1969). This flora occurs in fluvially deposited, irregularly fissile, tuffaceous mudstones at the southern edge of the Owl Creek Mountains. The assemblage has ap- proximately 15 species and is heavily dominated y palm leaves, Lygodium kaulfussii, and an en- tire-margined dicot leaf resembling Sapindus spp. Other common elements include "Populus" wyomingiana, Canavalia diuturna, cf. Typha, Zingiberopsis isonervosa, and Musophyllum complicatum. The importance of palms, herba- ceous monocots, and vines suggests low-stature, perhaps relatively open, floodplain vegetation. Somewhat similar vegetation may have been re- sponsible for forming the Lostcabinian-aged Vermillion pa coal in southern Wyoming (Nichols, in pre The Kissinger pm Tipperary flora consists of 5 ferns, 1 horsetail, 2 conifers (G/yptostrobus and Chamaecyparis), and 44 angiosperms (MacGinitie, 1974). Of the 36 well-defined di- cotyledonous leaf types, 5596 have nonentire margins, and judging by their closest living rel- atives, some 60% were deciduous. A number of the genera in the fossil assemblage are presently restricted to subtropical or tropical climates (e.g., Acrostichum, Apeiba, Canavalia, Dendropanax, WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS 759 ini octal Based on foliar physiognomy and e distribution of living relatives, MacGinitie MA inferred that the Kisinger Lakes flora was derived from a subtropical to tropical, semide- ciduous forest resembling those native to the southwest coast of Mexico at elevations of about 1,000 m. These inferences are consistent with the diverse palynoflora, which in most samples iS ET by angiosperms and ferns (Leo- pold, 1 The FU floras of Yellowstone National Park were first described by Knowlton (1899) and have not been subjected to a general revision since. Therefore, most of the published identifications are probably incorrect. Furthermore, much of the megafossil material comes from sites that are imprecisely located and of unknown stratigraph- ic relationship. It may be that significant tem- has remained undetected as a result. In spite of these problems, most of the Yellowstone assemblages were derived from vegetation that was quite dif- ferent from the kind inferred for the assemblages discussed above our genera of conifers are known from mega- fossil remains (Sequoia, Glyptostrobus, Pinus, and ?Podocarpus), and the first three are abundant at many localities (Dorf, 1960; Aguirre, 1977). As might be expected, the importance of conifers is even more strongly indicated by the palynoflora, where they are diverse (12 genera) and consis- tently make up about half ofthe noie (Fisk, 1976). Ferns are also highly diverse and abundant at many localities in the Sepulcher Formation. Fisk from the Yellowstone Park palynoflora, and there are 10-15 named species from the macroflora. The local abundance of ferns at some localities (particularly Thelypteris weedii and Allantoidiop- sis erosa) may be related to the frequency of dis- turbance by volcanic events. A similar domi- nance of ferns in the colonizing vegetation has been noted following eruption and deposition in the vicinity of EI Chichón volcano in Mexico (Spicer et al., 5 The foliar physiognomy of the Yellowstone National Park floras has not been studied in de- tail, but dicot leaves in the collectionsat the U.S. National Museum of Natural History are mostly in the notophyll and mesophyll size categories. Drip tips are present on a few taxa, and roughly half of the species have entire-margined leaves. Although the percentage of entire-margined species is similar at Yellowstone (approximately No oo 760 50%) and Kisinger Lakes (54%), leaf size in the Kisinger Lakes flora is generally in the micro- phyll to notophyll range (Wing, unpubl. data; Wolfe, pers. comm., 1987). These PER aie aspects of the Yellowstone assemblages, in ad- dition to the importance of conifers, d that the vegetation grew under a somewhat cooler and perhaps less seasonally dry climate. The forest from which the Yellowstone flora was derived appears to have been a variety of broad-leaved evergreen forest that included a substantial ele- ment of conifers in some local environments. The question of taphonomic mixing of floral assemblages is ever present but has been a par- ticular focus of debate with regard to fossil floras from Yellowstone National Park (Fisk, 1976; Fritz & Fisk, 1978; Fritz, 1980a, 1980b, 198 La, 1981b; Retallack, 1981; Yuretich, 1984; Karowe & Jefferson, 1987). The mixture of "tropical" and "temperate" elements (e.g., Thuja and pre- sumed evergreen members of the Lauraceae) in the Yellowstone floras has been attributed to transport of plant remains derived from vege- tation growing at a range ofelevations (Fisk, 1976; Fritz, 1980a, 1986), and the upright stumps and autochthonous “fossil forests" described by Knowlton (1899) and Dorf (1964) have been ex- plained in part as the consequence of transport by mudflows associated with volcanic activity (Coffin, 1976; Fritz, 1980a, 1980b, 1986; Fritz & Harrison, 1985). Observations of present-day volcanic systems have shown that high-energy mudflows can transport upright stumps from higher to lower elevations, and that stumps weighted by soil trapped in their roots may float upright for a time in lacustrine situations (Fritz, 1980a, 1980b, 1986; Coffin, 1983). Furthermore, sedimento- logical and stratigraphic studies of the Sepulcher and Lamar River formations and other Eocene volcanic units have highlighted the importance of high-energy deposits that are presumably in- dicators of steep paleotopography (Fritz, 1980c; Fritz & Harrison, 1985). In spite of these im- portant observations, several lines of evidence suggest that the Yellowstone megafossil assem- blages are not highly allochthonous First, although many fossil trees in the Spec- imen Ridge section may be prone rather than upright, this is not always the case. The presence of paleosols around some of the upright stumps is evidence that these were fossilized in place rather than transported upright to the site of buri- al and preservation (Retallack, 1981). Yuretich ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 (1984) and Karowe & Jefferson (1987) have pre- sented petrographic and sedimentological evi- dence that the upright stumps are generally in situ. 1) Tree stumps are rooted in fine-grained sediments, not in conglomerates. 2) Some con- glomerates have structures showing that they flowed around trunks in place. 3) Upper parts of inclined axes are abraded, but parts contained in finer-grained rock are not. 4) Petrographic sec- tions of sediment containing fossil tree roots show no signs of extensive current bedding but do show indications of pedogenesis. Many of the leaf EGER assemblages, which also show mixed "tempera and "tropical" forms, are derived from ue fluvial sediments and thus are unlikely to be highly transported (Wing, unpubl. data). Some fine- to medium-grained, airfall tuffs have the potential to entomb plant assemblages that quite accurately reflect local vegetation (Burnham & Spicer, 1986). Second, the presence in the same sedimentary units of trees showing distinct seasonal growth with trees lacking distinct growth rings (Wheeler et al., 1977, 1978) has been cited as evidence that plants that grew under more than one cli- matic regime are present (Fritz & Fisk, 1978). This interpretation is not justified. Tree species have varying genetic capacities for seasonal growth, and individual trees are variably influ- enced by microclimatic and edaphic factors. The result is that different trees in the same region may show different patterns of growth (e.g., Tomlinson & Craighead, 1972). Fossil log as- semblages containing specimens with both sea- sonal and aseasonal growth have been observed in sedimentary environments that could not have produced long-distance transport (Bown et al., 1982). Third, most of the extreme examples of dis- sonance in the climatic tolerances of elements in the flora are generated by comparing the paly- noflora with the megaflora. Pollen of Abies and Larix were less than 196 of the assemblages in which they occurred, which in turn were only a few of the 20 samples taken by Fisk (1976). Ob- viously, these could be highly allochthonous pol- len grains derived from vegetation growing at higher elevations than the site of deposition. Pic- ea occurred as 1—3 grains in 12 of the 20 samples, and reached 296—696 of the flora in three samples (Fisk, 1976). Cross & Taggart (1982) have noted that abundance of Picea pollen is generally in rough proportion to its importance in the source vegetation, implying that spruce could have been 1987] a minor part of local vegetation, probably higher elevations in the watershed containing the site of deposition. Fourth, almost all early Eocene floras from western North America contain some taxa that presently have mutually exclusive climatic re- quirements (MacGinitie, 19 74; Wolfe, 1980). In many floras the depositional setting argues strongly against explaining the presence of these dissonant elements by transport. For instance, backswamp compression fossil assem- blages from the Willwood Formation, perhaps one million years older than the Yellowstone flora, have abundant palms, Cnemidaria (cy- atheaceous tree fern), and A/nus (Wing, 1981). The Willwood Formation is a mostly fine-grained, basin-fill formation deposited by low-energy streams (Bown, 1979); low paleotopography is further indicated by thin, laterally extensive sheet sands (Kraus, 1980) and paleosol horizons that are traceable over many manis n & Kraus, 1981). Many of the collecting sites for the Willwood flora were near the center gn e basin, at least 100 km from the uplifted basin margins. The assemblages were derived from fine-grained mudstones (Wing, 1984), and similar assem- blages are characteristic of this lithology over a large part of Wyoming (Wing, unpubl. data). Given this depositional environment, the poten- tial for long-distance transport of leaves is min- iscule, and the assemblages must be autochtho- nous or transported only a short distance. Therefore there has been change through time in the climatic requirements of the taxa involved and/or early Tertiary climates permitted the co- existence of genera that at present have largely nonoverlapping ranges In conclusion, recent stratigraphic and sedi- mentological studies of volcanic strata in Yel- lowstone National Park and other Paleogene vol- canic sequences have demonstrated the importance of high-energy mudflow deposition and have made it clear that such deposits are characterized by a high degree of lateral vari- ability. However, these studies have failed to demonstrate that the majority of the megafossil assemblages from such deposits are allochtho- nous. Currently, prevailing evidence suggests that plant megafossils, especially compression assem- blages of leaves, generally accumulated from lo- cal sources during quiet periods between violent, localized, mudflow deposition. Although the Republic flora (168-170; Figs. 3, 4) is approximately 2 Ma younger than the four WING —EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS 761 floras discussed above, it is similar to the Yel- lowstone flora in having abundant conifers in both the megaflora and palynoflora. Wolfe & Wehr (1987) argued that the Republic paleoveg- etation was a mixed coniferous forest dominated by conifers because: 1) Chamaecyparis and Pinus are abundant at the Graphite Creek locality, 2) bisaccate pollen dominates the palynoflora, and 3) conifers are quite diverse (including Thuja, Abies, Picea, Tsuga, and Pseudolarix in the megaflora). Deciduous angiosperms are the most abundant and diverse group at the other two lo- calities, but broad-leaved evergreens are present, including Phoebe, Photinia, Ternstroemites spp., and a new extinct genus. Based on the physiog- nomy of the assemblage, Wolfe & Wehr (1987) inferred a mean annual temperature of 12—13°C and a mean annual range of temperature of 5— 6°C. By comparing these temperature estimates with those inferred for approximately coeval flo- ras from the Puget Group, Wolfe & Wehr (1987) estimated the paleoelevation of the Republic flora as 725-910 m. The greater importance of coni- fers and deciduous broad-leaved plants at Re- public than in the Yellowstone flora probably does not reflect a higher paleoelevation if esti- mates for the two areas are even approximately correct. Therefore, the compositional and phys- iognomic differences between the two floras probably result from differing taphonomic pro- cesses, climatic cooling during Bridgerian time, or both. Vegetation during the Chadronian (38-33 Ma). Comparison of the approximately con- temporaneous Little Mountain, Boysen, Kisin- ger Lakes, and Yellowstone floras illustrates the kinds of vegetational and floristic d t existed over one part of the northern Rockies at about 50 Ma. The regional importance of conif- erous and broad-leaved deciduous elements seems to have increased during the later Eocene and Oligocene, with dominantly coniferous for- ests probably becoming established at higher el- evations. However, later Eocene and Oligocene floras provide less evidence of the kinds of vege- tational boundaries implied by the contrasts be- tween the four floras discussed above. This may reflect in part a less adequate sample of coeval floras. One time interval in the remainder of the Eocene and Oligocene contains enough fossil flo- ras to permit an attempt at analyzing vegeta- tional variability in the Rocky Mountain region. 762 Ten floras in Appendix I are from rocks of prob- able Chadronian age: Missoula, from westcentral Montana (245-246; Jennings, 1920); Christen- sen Ranch, Horse Prairie, and Medicine Lodge from southwestern Montana (151-152, 153-157, 159-160; the Beaverhead Basins floras of Becker, 1969); uppermost Bull Run from northeastern Nevada (202; Axelrod, 1966a, 1966b); Florissant from central Colorado (224—229; MacGinitie, 1953); Red Rock Ranch, Hermosa, and Hills- boro from the Rio Grande Rift of New Mexico (232, 233, 234; Axelrod & Bailey, 1976; Meyer, 1986); and a small flora from the White River Group in the Flagstaff Rim area of central Wy- oming (2 The Missoula flora was derived from a se- uence of coal and lacustrine ash beds that Jen- nings (1920) believed were correlative with the lower part of the White River Formation in South Dakota. The flora has not been studied since 1920, and many of Jennings's identifications are questionable. However the flora appears to be dominated by Metasequoia, Sequoia, and one or more species of Betulaceae. Equisetum, ?Thuites, ?Populus sect. Abaso, Acer, and ?Cercidiphyllum are also present. It is difficult to reach any con- clusion about vegetation based on such a small assemblage of relatively ubiquitous taxa. Becker (1969) considered the Beaverhead Ba- sins floras to be of latest Oligocene to Miocene age, but this was based on the thickness of the local stratigraphic sequence and on floral corre- lation; the latter was in turn based on an incom- plete understanding of the taxa involved. More recent geological and paleontological work sug- gests that the sections are thinner than originally thought, and that the “Medicine Lodge Beds,” from which the floras were collected, may cor- relate with sediments producing Chadronian vertebrates (Fields et al., 1985). All three of the Beaverhead Basins floras represent mixed conif- erous and deciduous broad-leaved forest, which Becker (1969) felt reflected a subhumid climate. The subhumid elements include such taxa as Mahonia, Juniperus, and various nanophyllous Leguminosae. Common broad-leaved deciduous taxa are Fagopsis longifolia, Cercidiphyllum, Populus, Sassafras, and various Betulaceae and Ulmoideae. Upsection changes in floral com- position are relatively modest, although the up- per two floras have 14 (Medicine Lodge) and 16 (Horse Prairie) species of conifers to the 10 species found in the lowest flora. There is a pronounced physiognomic change between the Christensen ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Ranch flora and the upper two in that the per- centage of species with entire-margined leaves increases from 18% to 34% and 37%. The sig- nificance of this shift is cast into some doubt by concomitant changes in depositional environ- ment and floral diversity, and by the lower re- liability of leaf margin data from coniferous vegetation (Wolfe, 1979), but taken at face value the increase in percentage of species with entire- margined leaves would indicate an increase in mean annual temperature from 7? to 12°C. Axelrod (1966a, 1966b) stated that the upper Bull Run floras represented montane forest al- most totally dominated by conifers, including Abies, Picea, Pinus, Pseudotsuga, Tsuga, Cham- aecyparis, and Thuja. The angiosperm compo- nents are small-leaved and, with the exception of Mahonia, deciduous genera such as Alnus, Betula, Ribes, and Zelkova, This low diversity, igly conifer markedio “with the Beaverhead Basins floras, and although the upper Bull Run floras may be slight- ly older than those of the Beaverhead Basins (the highest locality is five feet below a tuff dated at 36 Ma; Axelrod, 1966a; Table 1), it seems prob- able that a difference in paleoelevation is also involved. The small flora from the Chadronian of the Flagstaff Rim area was ted from clastic dike fillings that also preserve vertebrate skeletons (Emry, pers. comm., 1986). This peculiar mode of deposition makes comparisons between this flora and others questionable, but the low di- versity (about six forms) and the small leaf size (microphylls or nanophylls) probably indicate relatively dry conditions. The identifiable taxa are Mahonia, ?Ribes, ?UImaceae, Leguminosae, and an undetermined conifer. This small flora may represent interfluve vegetation better than typical collections obtained from rocks deposited in paludal or lacustrine settings. The Florissant flora is the most diverse from the Oligocene of the Rocky Mountain area. Based on a combination of floristic and physiognomic criteria, MacGinitie (1953) inferred that the Florissant fossil d two main types of vegetation: a mesic, broad-leaved de- ciduous forest along Ram and lakeshores and on slopes and interfluve areas. The ten most common species in the flora comprise 6096 of the speci- mens je Fagopsis longifolia, Zelkova dry- eja, Chamaecyparis, Typha, Populus crassa, Bhus ae Sequoia affinis, Cercocarpus a drier Scrub 1987] myricaefolius, Staphylea acuminata, and Ath- yana haydenii. The broad-leaved evergreen com- ponent of the Florissant is not dominant, but the diversity and abundance of conifers is much less than in either the Beaverhead Basins or the upper Bull Run floras. Chadronian floras from New Mexico (Axelrod & Bailey, 1976; Meyer, 1986) appear to represent two different types of vegetation. The Red Rock Ranch flora is numerically dominated by spec- imens of Pinus subsection Balfourianae and con- tains species of Picea, Abies, Zelkova, and pos- sibly Salix and Rosa (Meyer, 1986). Based on a list by Farkas (1969), Axelrod & Bailey (1976) reported several additional taxa including Fa- gopsis, Halesia, Mahonia, Rhus, and Sapindus, but, with the exception of Mahonia, these were not confirmed by Meyer (1986). Although the Red Rock Ranch flora is small, on the basis of relatively high conifer diversity and abundance and low broad-leaf diversity, Meyer (1986) con- cluded that it most likely represents a mixed co- niferous forest. This flora was correlated with Florissant (35 Ma) by Axelrod & Bailey (1976), but the more recent and direct radiometric date obtained by Meyer (1986) indicates the flora is no younger ool > : + 1.1 Ma, or about 2 Ma older than Flori The Hermosa nd fiios floras are derived from sediments associated with the infilling of the moat of the Emory caldera, and both were dated at about 32 Ma by Axelrod & Bailey (1976). New radiometric dates reported by Meyer (1986) indicate the Hillsboro flora is probably 28.1—30.6 Ma (Whitneyan or early Arikareean) and that the Hermosa flora is about 33.6 + 1.0 Ma (Chad- ronian). The floras are similar in composition, with an overwhelming dominance of specimens of Pinus subsection Balfourianae, along with a few small leaves of Mahonia, and possibly Picea and Crataegus. These floristic and physiognomic attributes indicate a taiga-type forest growing un- der a cold temperate climate (Meyer, 1986). Ax- elrod & Bailey (1976) argued that the difference between the Red Rock Ranch and Hermosa/ Hillsboro floras was a response to higher pa- leoelevation of the latter floras; however, Meyer (1986) pointed out that these floras may bracket the major decrease in mean annual temperature and increase in mean annual range of tempera- ture that occurred at about 33 Ma (Wolfe & Hop- kins, 1967; Wolfe, 1986). The pattern of geographic variation in Chad- ronian vegetation is less obvious than that of the WING —EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS 763 earliest Bridgerian, and this can be attributed to several causes. First, the Chadronian floras are less tightly correlated and probably occurred over a longer interval of time, perhaps 5 Ma. Second, several of the floras are of low diversity or are not completely described, with the result that they are poorer samples of regional vegetation. Third, there may be a greater altitudinal range represented by the floras. Fourth, the youngest floras in the set (Hermosa/Hillsboro) may lie on the opposite side of a major temperature de- crease from the other floras. In spite of these problems, the three floras of presumed inter- mediate elevation (Beaverhead Basins, Floris- Rock Ranch) seem to have broadly similar compositions and to represent mixed co- niferous and broad-leaved deciduous forest growing under a seasonally dry climate. The higher diversity of conifers and mesic taxa in the Montana floras may indicate higher rainfall and/ or lower rates of evapotranspiration in the north- ern part of the Rocky Mountains. The subhumid aspect of all of these floras when compared with those of similar age from the Pacific Northwest demonstrates the continuation of the pattern of regional climatic variation that began during the early Eocene/latest Paleocene. The mid Tertiary climatic deteriora- tion. Much of the evidence for the major de- crease in mean annual temperature and increase in mean annual range of temperature that oc- curred at approximately 33 Ma is derived from lowland floras from the Pacific Coast of North America (Wolfe & Hopkins, 1967; Wolfe, 1971, 1986). Isthere unambiguous evidence ofthis ma- jor climatological change in floras from the Rocky Mountain re presented in Figure 3, there are six floras that closely follow after the 34 Ma date: Mormon Creek, Metzel Ranch, York Ranch, Ruby paper shales, Hermosa, and possibly Hillsboro. The dating of the Montana floras has been uncertain since their initial descriptions (Becker, 1960, 1961, 1972, 1973), and opinions on the age of the Mormon Creek flora have embraced some 20 Ma. Recent geological and mammalian bio- stratigraphic correlations suggest the Mormon Creek, Metzel Ranch, and York Ranch floras are of Orellan age (32.2-30.8 Ma) and that the Ruby paper shale flora is Whitneyan (30.8-29.2 Ma). These floras were judged by Becker (1960, 1961, 1972, 1973) to represent mixed coniferous and 764 broad-leaved deciduous forest and shrubland growing under temperate to dry-temperate cli- mates. Although the Ruby paper shale flora is inferred to have been somewhat dryer than the other three, they all bear substantial resemblance to one another and to the older (Chadronian) Beaverhead Basins floras. As noted above, the New Mexican floras are derived from a setting of some paleotopographic and structural com- plexity, so that it is difficult to separate potential effects of elevation and regional climatic change. Thus the only floras of suitable age do not provide good evidence of a major temperature decrease at 33 Ma. This absence of evidence may result from the confounding effects of changing elevations and dryer climates in the Rocky Mountains, or it may simply be a problem of insufficient data and poor stratigraphic control. PRESENT METHODS AND FUTURE STUDY Perhaps the most useful result of summarizing current knowledge and opinions in a field of study is that this activity reveals gaps in the data base and reveals possible directions for future re- search. At present, research in Tertiary paleo- botany follows two main themes: the systematic/ evolutionary approach is concerned primarily with describing new fossil forms and understand- ing their implications for the evolutionary his- tory of lineages and relationships among living groups of plants; the paleoecological/environ- mental approach focuses largely on understand- ing habits of extinct species, structure of extinct vegetation, and patterns of ancient climates. His- torically these approaches often were combined in the treatment of a single fossil flora. More recently, as standards have become higher and techniques more sophisticated in both ap- proaches, workers have specialized on one or the other. This probably reflects a more general sep- aration of ecology from systematics, but the dis- junction in viewpoints should not be accepted as logical data provide the context for understand- ing the genealogical changes that are inferred from systematic studies. Furthermore, in paleobotany both approaches are united at a practical level by the study of the same sites and specimens. FIELD DATA As noted in the section on the data base for Eocene and Oligocene floras, the stratigraphic ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 and geographic distribution of sites is very un- even. The total number of sites from which col- lections have been made is also small, consid- ering the commonness of plant fossils and the large outcrop area. The meagerness of the data set results from there being few paleobotanists, rom a tendency for the same sites to be collecte repeatedly, and from relatively small efforts to- ward finding new sites. The largest “holes” in the record would be filled by: 1) more late Eocene and Oligocene floras from the eastern Rocky Mountain area; 2) more Paleogene floras from the southern Rocky Mountains; and 3) more flu- vially deposited late Eocene or Oligocene assem- blages (or conversely, more Paleocene and early Eocene lacustrine assemblages). Field data that would Bien reports of fossil plants include: number of quarry sites collected; size of quarry; precise locality information; stratigraphic unit; detailed lithological descrip- tion; abundance of fossils and relative abundance of species; and, where available, biostratigraphic correlation, radiometric age, and paleomagnetic correlation. If such data were available, even as preliminary approximations, the published rec- ord of Tertiary fossils would be more useful for interpreting paleovege no paleoclimate, and possible associations of dispersed organs SYSTEMATICS During the last 15 to 20 years several new methods have brought increased resolution and rigor to systematic studies of Tertiary angio- sperms. Comparative studies of the leaf archi- tecture of living dicotyledons have created a much firmer basis for interpreting the systematic affin- ities of fossil leaves (Hickey & Wolfe, 1975). The structure of fossil pollen (e.g., Crepet et al., 1980), and from more detailed analyses of fossil flowers (e.g., Crepet & Daghlian, 1981, 1983). At the same time, more studies have come to base their systematic conclusions on multiple fossil organs belonging to the same species (e.g., Manchester & Crane, 1983; Wing & Hickey, 1984; Man- chester, 1986). Finally, in many areas of paleo- botany, refinements in methods of systematic the reasoning behind systematic decisions more explicit (e.g., Stein et al., 4 Although major advances have been made in 1987] methodology, the vast majority of fossil angio- sperms from western North America are as yet very poorly understood. For most times and places the floras either have not been described or the only descriptions are those of late-19th century workers whose goals were more bio- stratigraphic than systematic. New methods will have to be applied repeatedly before the botan- ical relationships of any significant number of Te estiary fossils will be MASSE ON the quantification of variability. Comsemtwe y architecture was an important advance in ana- lyzing higher-level systematic affinities but has been much less useful at the level of species. With few exceptions (e.g., Dolph, 1975; Burnham, 1986a, 1986b; Wing & Eckenwalder, 1987), pa- leobotanists have been little concerned with quantifying the variability of their taxa. Yet in- dividuating taxa in a paleobotanical sample is the initial step in subsequent systematic, bio- stratigraphic, and paleoecological syntheses. De- tection and quantification of low-level morpho- logical variability is also a key to uncovering patterns of temporal change in closely spaced stratigraphic samples. PALEOECOLOGY Traditional paleoecological interpretation of fossil angiosperm floras has been based on flo- ristic analogy and on leaf physiognomic analysis. The floristic method assumes that the ecological/ climatic requirements of the fossil taxa were sim- ilar to those of their closest living relatives. This kind of direct analogy suffers from several defects (see Wolfe, 1979). First, it assumes that the bo- tanical relationships of the fossils have been de- termined correctly. Second, it assumes that little evolutionary change has occurred in the climatic or ecological preferences of the lineages under study. Justification for both of these assumptions diminishes as one considers older floras, because evolution is more likely to have occurred in the intervening time, making it more difficult to de- termine close living relatives of older fossil plants. A third problem with floristic inference is that it implies that the present-day distribution ofa tax- on accurately reflects even its present climatic tolerances. Given the rapid climatic fluctuations typical of the last two million years, it may be that the current distributions of many taxa are strongly influenced by migration rate, plant com- petitors, or other nonclimatic factors (Davis, 1976 The second commonly used method of pa- WING —EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS 765 leovegetational/paleoclimatic inference is anal- ysis of leaf physiognomy. Physiognomic analysis primarily relies on a relationship observed in living floras; the percent of species in a local flora that have entire-margined leaves rises with the mean annual temperature of the site (Bailey & Sinnott, 1915, 1916). Thus tropical floras have nearly 10096 species with entire leaves, whereas temperate floras are dominated by species with toothed or lobed leaves. This relationship has been worked out with some precision based on the humid floras of East Asia (Wolfe, 1979). Studies based on smaller regions have been used to question the resolution of leaf-margin infer- ences (Dolph, 1976, 1978a, 1978b, 1979; Dolph & Dilcher, 1979), but the basic pattern of climate change as inferred from Tertiary floras agrees with data from a variety of other sources (e.g., Savin, 1977; Wolfe & Poore, 1982; Hutchison, 1982; Owen & Rea, 1985; Rea et al., 1985). Phys- iognomic analysis also considers the average size of leaves in a fossil assemblage, their apparent number of | number of species that are probable lianes (those with cordate-based leaves). Generally these at- tributes increase with increasing tropicality of vegetation. Although physiognomic analysis offers signif- icant improvement on the floristic method, it has defects. In addition to their correlation with mean annual temperature, physiognomic characteris- tics of leaves are also correlated with water avail- ability, intensity and angle of incident radiation, and a variety of other factors. Consequently, changes in the leaf physiognomy of fossil assem- blages cannot always be read unambiguously as changes in mean annual temperature. An in- daphically Induced water stress could produce Xn with small, thick, entire-margined leaves and few lianes. Greater representation of canopy species in a fossil flora would produce an assemblage with smaller leaves (Roth & Dilcher, 1978). This is because canopy leaves tend to be smaller than interior leaves in order to radiate heat more efficiently and main- tain an optimal photosynthetic rate. Perhaps the most serious factor biasing leaf physiognomic analysis of fossil assemblages is the probable overrepresentation of early successional and stream ges up which grow close to sites of dep in fluvial and volcanic settings Maec 1969). Successional and riparian vegetation in most climatic zones is dominated by species with lobed, toothed, or compound 766 leaves, probably because these species hold in- dividual leaves for only a short time, and these leaf shapes provide a large photosynthetic sur- face at a small cost of support tissue (Givnish, 1978). Thus a change in the frequency with which fossil vegetation was disturbed might produce a change in leaf physiognomy that might be inter- preted as a change in mean annual temperature. In spite of their defects, both the floristic and leaf physiognomic methods produce inferences about paleovegetation that are generally consis- tent with paleoclimatic reconstructions based on other, independent data sets. Furthermore, they generally agree with one another (e.g., Mac- Ginitie, 1974; Hickey, 1977). The problem with these methods is not that they produce grossly incorrect interpretations of past vegetational structure, but rather that the inferences lack de- tail, frustrating the most interesting comparisons that might be made between extinct and living forests. For instance, because of strong seasonality of light and a low angle of incident radiation, it is likely that the structure of high latitude, broad- leaved, evergreen forests in the early Tertiary was significantly different from that of living broad- leaved evergreen forests, even though the two types of vegetation are similar in leaf physiog- nomy and floristic composition. This hypothesis can only be examined by finding more ways to compare fossil and living vegetation. These new methods of comparison will probably require collecting data on the distribution of fossils in the sediment. These distributional data (e.g., al- pha and beta diversity, relative abundance, spa- tial heterogeneity) may reflect actual synecolog- ical characteristics of the vegetation that produced a fossil assemblage; the difficulty in interpreta- tion arises from the probability that taphonomic processes have also influenced the distribution of fossils. In spite of recent work on the taphon- omy of fossil plants (Spicer, 1981; Scheihing & Pfefferkorn, 1984; Spicer & Greer, 1986; Fer- guson, 1985; Gastaldo, 1986; Burnham & Spicer, 1986; Spicer & Wolfe, 1987), there are as yet no general recommendations for how leaf assem- blages can be sampled to reflect best given char- acteristics of the former vegetation. 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Additions to and revision of the Oli- gocene Ruby paper shale flora of southwestern Montana. Contr. Mus. Paleontol Univ. Michigan 20: 89-119. 1968. A hexasepalous calyx of the fossil As- tronium truncatum (Lesquereux) MacGinitie. Bull. Torrey Bot. Club 95: 262-263. 1973. A new Tertiary gramineous fossil. Bull. Torrey B Bot. Club 100: 318-319 BERRY, E. 924a. An early Eocene florule from nd Texas. U.S. Geol. Surv. Prof. Paper 132-E: 7-92 1924b. A Sparganium from the middle Eocene of Wyoming. Bot. Gaz. (Crawfordsville) 78: 342- “ë 8. 19 A new Salvinia from the Eocene (Wy- oming a Tennessee). Torreya 25: 116-118. 1926. Tertiary floras from British Columbia. Canad. Dept. Mines, Geol. Surv. Branch Publ. 42: 91-116. ANNALS OF THE MISSOURI BOTANICAL GARDEN — ə 1937b. [VoL. 74 APPENDIX II. Continued. . 1930. A flora jae aiie River age in the Wind River Basin of Wyoming. U.S. Geol. Surv. Prof. ap 165-B: 55-81. — 1932a. poi plants from Wyoming. Amer. Bo Novit. 527: 1-13. 32b. 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Pinus driftwoodensis sp. n. from the Early Tertiary of British Columbia. Bot. Gaz. (Crawfordsville) 144: 148-156. 9 Middle Eocene Pinus remains fi British Columbia. Bot. Gaz. (Crawfordsville) 145: 252-274. & J. F. BASINGER. 1980. An Eocene pinaceous Misc. Ser. Publ. 158: 112. [Abstract.] apnd = L. 1978. Transported fossil biota of the n River Formation, Utah. Palaeogeogr. Pa- pe ones Palaeoecol. 25: 353-364. TIDWELL, W. D., D. A. MEDLYN & G. F. THAYN. 1973. Three new species of Palmoxylon from the Eocene Green River PNE Wyoming. Great Basin Naturalist 33: 61- E. M. V. Digicam & A. D. SiMPER. 1977. A new Palmoxylon species from the Eocene Gold- en Ranch Formation, Utah. Bot. Soc. Amer. Misc. Ser. Publ. 154: 44. [Abstract.] L. R. PARKER & N. P. HEBBERT. 1980. A new arborescent member of the Osmundaceae n the Union Formation (Paleocene), Wyoming. Bot. Soc. Amer. Misc. Ser. Publ. 158: 118. T) F. THAYN & J. L. RorH. 1976. m and early b ge floras ofthe agama g Univ. Geol. Stud. 22: 7 Re-evaluation of oe flo- rissantensis (Oligocene, North America). Trans. British Mycol. Soc. 76: 493-496. . 1985. The Eocene North spe land bridge: its importance in Tertia modern phyto- geography of the northern Sees eae J. Arnold Arbor. 66: 243-273. TiNG, W. S. 1968. Fossil pan grains of Coniferales from Early Tertiary of Idaho, Nevada and Colo- rado E Pollen & Spores 10: £1» 598. . Eocene pollen assemblage from Del Mar, southern California. Geosci. & Man 11: 159. [Abstrac TRIPLEHORN, m M., D. L. TURNER & C. W. N Radiometric age of the Chickaloon "For. mation of south-central Alaska: location of the 784 APPENDIX II. Continued. Paleocene-Eocene boundary. Bull. Geol. Soc. Amer. 95: 74 TscHupy, R. 1961. Palynomorphs as indicators of facies environments in Upper C ines n Lower Tertiary strata, Colorado and Wyoming. Wyoming Geol. Assoc. Guidebook, 16th pees Field Conf., pp. 53-59 1976. Pollen changes near the Fort Union- Wasatch boundary, Powder River Basin. 7n R. B. Laudon (editor), Geology and Energy Resources of the Powder River. Guidebook of Annual Field ESE 3. Ra partings in coal of the Eocene Puget Group, Wash- e- pits li: 527-531. UNDERWOOD, J.C. 1977. A new Pinus from the Oli- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 APPENDIX II. Continued. gocene of western Washington. Geol. Soc. Amer. Abstr. with Programs 9: 770. [Abstract.] Mir DR. JR. 1980. Pinus buchananii, a new species based on a petrified cone from the a of Washington. Amer. J. Bot. 67: 1132- 135. A E. A. & R. A. Scorr. 1982. Fossil woods from the Eocene Clarno Formation. IAWA Bull. 3: 135-154. WiNG, S. L. 1984. A new basis for recognizing the Paleocene/Eocene boundary 1 estern interior North America. Science 226: 439-441 WODEHOUSE, R. P 3. Tertiary pollen. II. Pollen of the Green River oil shales. Bull. Torrey Bot. Club 60: 479-524. Wo re, J. A. & E. S. BARGHOORN. 1960. Generic change in Tertiary floras in relation to age. Amer. J. Sci. 258: 388-399. AN OVERVIEW OF THE ORIGINS OF THE MODERN VEGETATION AND FLORA OF THE NORTHERN ROCKY MOUNTAINS! JACK A. WOLFE? ABSTRACT The present flora of the northern Rocky Mountains has diverse origins. The Late Cretaceous meso- thermal to megathermal evergreen vegetation of this region had few taxa, even at the familial level, that live in the northern Ro ok i i leaved deciduous taxa occupyi they un nderwent major diversification. Bes early Eocene thermal maximum severely restricted the areas of l lineages By the early middle Eocene, volcanic sehen that supported micrathera coniferous forests of Pinaceae and Cupressaceae had developed in s of the northern Rocky Mountain region. These ades into newly created T climate. During Eocene by way of Beringia. Some of the Eocene microthermal lineages s in the northern Rocky Mountains today, and other extant lineages, although ultimately derived from taxa in the Eocene uplands, represent morphological types S — ated and diverged in Eurasia, arriving in North America by migration. Oligocene and Neo northern Rocky Mountains, although the Columbia Plateaus [s oe t contain m of these ages. Th l broad-lea . deciduous or coniferous forests, t lineages that were derived fro the terminal Eocene temperature deterioration. With the presence of many extant northern Rocky the mid-Miocen exchanged between western North sampa, me and Eura x lineages are known during the Miocene only in areas ne as Alaska and probably represent migrants into the northern Rocky Mountains during the late Neogene. The present flora of the northern Rocky Mountains therefore clearly represents a complex overlay of numerous historical biogeographic pat- terns. The modern vegetation of the northern Rocky Mountain region is primarily steppe at low al- titudes and coniferous forest at higher altitudes (Habeck, 1987). Temperatures are entirely mi- crothermal.? The lower and drier part of the co- niferous forest belongs to the Pinus ponderosa zone, whereas most of the mesic coniferous forest at higher altitudes belongs to the Abies grandis nklin & Dyrness, 1969). The vegeta- northern Rocky Mountain region are docu- mented by successive microfossil and megafossil plant assemblages. In some instances, lack of as- semblages that represent particular intervals makes it necessary to infer climate and vegeta- tion from the fossil record of adjacent regions. ic) types that have occupied Mountain region in the past and the history of the lineages that comprise the modern flora. Throughout, the focus is primarily on the woody flora. ! For many helpful discussions, I thank H. E. Schorn, G. R. Upchurch, = and Wesley Wehr. Schorn has also supplied data from his unpublished studies on the p ú iun . R. Crane and B. H. Tiffney offered useful comments o ern American Tertiary conifers. ? Paleontology and Stratigraphy Branch, MS-919, U.S. s EE “sassa Federal Center, Denver, Colorado 80225, U.S.A. 3 Microthermal = mean annual temperature <13°C, mesothermal = °C megathermal = mean annual temperature > 2 ANN. MISSOURI Bor. GARD. 74: 785-803. 1987. mean annual temperature 13-20?C, 786 The extant woody flora is composed of 10 gen- era and 19 species of conifers and 30 genera and 80 species of woody dicotyledons. The dominant conifers are Abies grandis, Larix occidentalis, and other members of Pinaceae, as well as a fe members of the C upressaceae, The dicotyledons arc pri Imar ily tor form streamside communities. Conspicuous dicoty- ledonous families, as elsewhere in Northern emisphere vegetation, include Betulaceae, Er- icaceae, Salicaceae, Rosaceae, and Caprifoli- aceae, with a few Aceraceae. The origins of the flora of a region such as the northern Rocky Mountains are, of course, highly complex. Each component species, genus, and Family hasa unique history, REDE some his- ories pace and time. indeed ‘the search for patterns to which numer- ous histories approximately conform is a pri- mary concern of historical plant geography. Such patterns might be indicative of similar re- sponses to specific historical factors, particularly responses to environmental changes, including various parameters of climate and topographic changes resulting from orogenic factors. How- ever, because two taxa a are now in association or have similar p timply similar histories. The classical methodology for determining the histories of biogeographic patterns involves re- construction of phylogenies from phenetic sim- ilarities of extant and fossil taxa, observing dis- tribution patterns of extant taxa, and determining the patterns of distribution of fossils in time and space. The relative completeness of the fossil rec- ord is the major problem in using it in biogeo- graphic analyses. The problem is underscored in the present case by the almost total absence of Oligocene and Neogene assemblages of plant inet in tae northern Rocky Mountains. ges fill this gap, the typical taxonomic resolution of palyno- logical morphology is typically at a generic or higher level (Muller, 1970) and is of limited as- sistance in determining histories of lineages on the scale desirable in biogeographic analyses. Further, the upland Eocene megafossil assem- blages from the northern Rocky Mountain region have been actively collected and studied for only the last two decades. These assemblages are crit- ical because they contain the records of diver- sification of many microthermal groups (see be- low), and additional collecting and study are needed. In contrast, the Oligocene and Neogene E ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 plant-megafossil assemblages from adjoining re- gions such as the Columbia Plateaus and from possible source areas such as Beringia are well known. Estimates based on the geographic and climatic distribution of fossil and extant taxa in- dicate that, at least in the Beringian Neogene, the lineages represented in the fossil record are thought to be more than 9596 complete for woody plants. Inferences also can be made regarding histories of lineages if a cladistic analysis of a group is combined with detailed knowledge of history of the physical environment. This approach, termed og ick & Nel- son, 1978) has considerable merit for groups that have poor or no fossil records. However, it rests on major assumptions and requirements: (1) Cladistic analyses inherently rest on the as- sumption of parsimony in evolution of char- cters. (2) Cladistic analyses must be based on valid determination of polarities of character states. (3) idi of the physical environments in all areas m known in detail. (4) The iid biogeographic analysis also rests on the assumption of parsimony in dispersals. In most recent applications, vicariance biogeog- raphers have either ignored the fossil record or, at best, have incorporated fossil taxa in cladistic analyses without reference to stratigraphy, al- though Grande (1985) emphasized the signifi- cance of the age relations of fossil taxa, which provide additional geographic data on distribu- tions. Another major tendency has been to con- sider the history of the physical environment solely in terms of plate tectonics, thus ignoring particularly climatic changes. Because climates of a given area generally determine vegetational types and, in turn, vegetation organizes the en- vironment into a variety of microenvironments (Upchurch & Wolfe, 1987) that determine whether a particular organism can live in a given area, the history of vegetation assumes para- mount importance for interpreting the biogeo- graphic histories of land organisms. The fossil record can be of major significance in any analysis in vicariance biogeography, as Grande (1985) emphasized. Howe b from the standpoint of icthyology, some contri- butions of paleontology to vicariance biogeog- raphy were not stressed or were overlooked: 1987] (1) The stratigraphic sequence of grades, if cau- tiously interpreted, can determine polarity states (Hennig, 1966). The fossil record can supply evidence of en- vironments in particular areas at particular times. (3) The fossil record can supply evidence of the existence of dispersal barriers and routes oth- er than those related to plate tectonics. The fossil record can supply minimal times cladogram, and these times can then be placed in the framework of the then-existing environments to determine probabilities of dipersals of the resulting, di- verging lineages. The fossil record can independently test the assumptions of parsimony in both evolution and dispersal. ~ N — ~ A — h, wa — Unfortunately, almost no cladistic analyses of groups of concern in the northern Rocky Moun- tains have been carried out. However, the results ` of a cladistic analysis of Acer (Wolfe & Tanai, 1987) when compared with the known fossil re- cord of Acer (especially in North America) in- dicate: (1) The known relative times of appearances of sections of Acer generally correspond to those predicted from the cladogram; those few that do not correspond probably resulted from the absence of collections from the appro- priate environment or areas Phylogeny reconstructed on a phenetic pa- leobotanical basis compares well with phy- logeny reconstructed on a cladistic basis. Dispersals have not been as parsimonious as would be predicted from vicariance bio- geography. The present distributions of species and sec- tions of Acer are complex and have resulted from a variety ofenvironmental, particularly climatic, factors. ~ N — ~ Ww wa ~ +. — As more cladistic analyses of extant groups are completed, they will provide a framework for evaluating phenetic paleobotanical phylogenies, but cladistic analyses, because of biogeographic “noise,” can provide only highly generalized concepts of histories of biogeographic patterns, some of which are much more complex than indicated by cladistics alone and some of which are invalid (Wolfe, 1981b). In essence, cladistic analyses test phenetic phylogenies based on the fossil record, whereas the fossil record tests WOLFE—ORIGINS OF NORTHERN ROCKY MODERN VEGETATION 787 models developed from vicariance biogeogra- h The divergence that apparently exists between *vicariance" and “‘dispersalist’”’ schools of bio- geography has resulted from psychological and sociological factors. Each school typically has re- jected the totality of the other’s methods and results. Both have inherent problems that can be resolved by reference to the other’s methods and/ or by using the other’s conclusions. Indeed, Grande’s (1985) discussion of vicariance bio- geography basically incorporates some dispers- alist concepts and uses some dispersalist data that support his conclusions; he argued for sys- tematic/biogeographic analyses on successive time planes to attempt to filter out biogeographic "noise" introduced by subsequent dispersal events. The approach used in the present report similarly will involve both cladistic and dispers- alist concepts and data. CRETACEOUS The northern Rocky Mountain region was oc- cupied by a mesothermal, broad-leaved ever- green forest during the Late Cretaceous (Dorf, 1942). Some conifers (e.g., Araucariaceae, ev- ergreen Taxodiaceae) probably were emergents in this vegetation (Wolfe & Upchurch, 1987b), and broad-leaved deciduous plants were restrict- ed largely to successional or disturbed vegeta- tion, especially along streams. General absence of drip-tips and a somewhat small leaf size in- dicate slightly subhumid conditions (Wolfe & Upchurch, 1986, 1987a). Analyses of woods from the North American mesothermal region suggest little seasonality of either temperature or precip- itation (Wolfe & Upchurch, 1987b). Presence of extensive marine rocks of Late Cretaceous age throughout much of the Western Interior indi- cates generally low altitudes. Miller (1987) emphasized the great floristic disparity between known Early Cretaceous flo- ras, including those of the northern Rocky Mountain region, and the modern flora of this region. Even with the rise to dominance of the angiosperms in the early Late Cretaceous, the zed ber dae hese. 1987). Whether any of these ul- timately gave rise to taxa that now occur in the northern Rocky Mountains cannot now be de- termined. By the later part of the Late Creta- ceous, some families that today have microther- mal members become recognizable. Aceraceae, 788 for example, occur in the latest Cretaceous of central Alberta, represented by an extinct genus interpreted as a sister group to Acer (Wolfe & Tanai, 1987). Even at high paleolatitudes, the flora was composed largely of trochodendroids, hamamelidaleans, and deciduous Taxodiaceae (Spicer et al., 1987); this high-latitude flora has some floristic similarities to that of the early Ter- tiary of the northern Rocky Mountains but no similarity to the extant flora of this region. PALEOCENE In areas such as eastern Montana, the Creta- ceous-Tertiary boundary is marked by a pre- sumed fallout layer that has anomalously high amounts of iridium and shocked minerals (Boh- or et al., 1984); these are considered to be evi- dence of an impact by an extraterrestrial body (Alvarez et al., 1984). This event at the end of the Cretaceous had a profound effect on the flora and vegetation of the northern Rocky Mountain region (Wolfe & Upchurch, 1986). At least 50- 60% of the latest Cretaceous lineages became ex- tinct (Hickey, 1981). Most extinctions were in the previously dominant broad-leaved evergreen element; in addition, araucarians and many oth- er evergreen conifers became extinct in this re- gion. Immediately above the presumed fallout horizon, palynofloras contain almost exclusively fern spores (Hotton, 1984; Tschudy & Tschudy, 1986). Within a few centimeters above this ho- rizon, angiosperm pollen occurs with the fern spores; the associated leaf flora contains both fern pinnae and a compound-leaved rosid, which is thin-leaved and probably deciduous. About 10 meters higher in the section, the leaf assemblage of about 15 species consists almost entirely of deciduous angiosperms (trochodendroids, ha- mamelidaleans, Tiliaceae) and deciduous coni- fers (Taxodiaceae). The eastern Montana sequence immediately above the Cretaceous- Tertiary boundary resem- bles normal secondary succession following a volcanic eruption (Richards, 1952); in mesother- mal, broad-leaved evergreen regions, deciduous plants are typically dominant in secondary successions (Wang, 1961). However, in the northern Rocky Mountain region, broad-leaved deciduous plants dominated north of the Colo- rado- Wyoming border (Brown, 1962), and the broad-leaved evergreen element did not return to dominance until almost the end of the Paleo- cene, about 10 Ma following the Cretaceous. Hickey (1981, 1984) attributed the deciduous- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 ness of this Paleocene vegetation to an overall decline in temperature that persisted through most of the Paleocene. The occurrence of large (and presumably ectothermic) reptiles (crocodil- ians, champsosaurs, large turtles) with this de- ciduous vegetation, however, indicates the prob- able absence of low winter temperatures that would select for deciduousness. That is, the fau- nal data indicate strongly that the deciduousness of the vegetation is anomalous; just as signifi- h vegetation persisted in the early Paleocene at about the Colorado- Wyoming border. Wolfe & Upchurch (1986) suggested instead that the terminal Cretaceous event resulted in a brief low-temperature excursion that devastated the vegetation of both mesothermal and mega- thermal regions of North America. Some mega- thermal, broad-leaved evergreen lineages sur- vived (perhaps in refugia; Tschudy et al., 1984), and megathermal vegetation continued to be dominantly evergreen. The megathermal vege- tation underwent physiognomic change and gradually increasing diversity during the Paleo- cene, a phenomenon sess short-term sec- ondary succession and termed “quasisucces- Wolfe & Upchurch (1986, 1987a). broad-leaved evergreen lineages survived, and replenishment of them would have had to occur y adaptation of some megathermal lineages to mesothermal climate on an evolutionary (not successional) time scale. The anomalously deciduous character of Pa- leocene mesothermal vegetation throughout North America (and in Eurasia) gave deciduous angiosperms a unique opportunity. These decid- uous lineages were derived primarily from ele- ments that were uncommon in Late Cretaceous mesothermal evergreen vegetation and/or dom- inant in Late Cretaceous, high-latitude decidu- ous vegetation (Wolfe & Upchurch, 1986). In mesothermal regions, dominantly deciduous taxa (e.g., Hamamelidaceae, Fagaceae, Betulaceae, Ulmoideae, Juglandaceae) diversified during the Paleocene (e.g., Nichols & Ott, 1978; Manches- ter, 1987). By the end of the Paleocene, the Northern Hemisphere had a considerable diver- sity of broad-leaved deciduous plants, and many families of woody angiosperms that now occur in the Rocky Mountain region were extant. major increase in precipitation occurred at the Cretaceous—Tertiary boundary. In the north- ern Rocky Mountain region this is evidenced by 1987] a substantial increase in leaf size and by the ini- tiation of wide-scale peat deposition that typi- cally marks the early Paleocene in this region (Wolfe & Upchurch, 1986). Not only would in- creased precipitation result in an increase in bio- mass, but water tables also would be raised and more swamps developed. Pre-Eocene microthermal vegetation contrasts markedly with Eocene and later microthermal vegetation in floristic composition. Pre-Eocene microthermal vegetation was restricted to high latitudes and, probably due to low winter light levels, was almost entirely deciduous (Wolfe, iS a n - O o = Re a) m P Ne) ç ON = E ° = o = e "3 - ' ciduous Pinaceae (Pseudolarix), and deciduous dicotyledons, the great majority of which rep- resents a few clades of trochodendraleans, plat- anoids,and other hamamelidaleans (Spicer et al., 1987; Wolfe, 1987). Although these clades later contributed some lineages first to the meso- thermal deciduous vegetation of the Paleocene and then to the Eocene upland microthermal vegetation, they comprise a small fraction of the Eocene and later microthermal flora. The mi- crothermal vegetation and flora of the Late Cre- taceous and the Paleocene have few similiarities to Eocene and later vegetation and flora. EOCENE A warming initiated during the latest Paleo- cene culminated in the early Eocene thermal maximum (Savin, 1977; Wolfe & Poore, 1982; Wolfe, 1985). Vegetation in areas such as north- western Wyoming was dominantly broad-leaved evergreen and represented warm mesothermal temperatures (Wing, 1981, 1987). At somewhat lower altitudes (and particularly in the lowland acific Northwest), vegetation represented , 1985). region was still generally of low altitudes is in- dicated by the persistence of large lakes from the early into late Eocene in areas such as south- western Wyoming and adjacent Utah. The early Eocene vegetation indicates abundant precipi- tation, but later Eocene vegetation indicates increased seasonality of precipitation and de- velopment of subhumid, seasonal climate (MacGinitie, 1969; Leopold & MacGinitie, 1972). e warm mesothermal to megathermal vege- tation of the early and middle Eocene has few floristic similarities to the extant flora of the northern Rocky Mountains. A few extant north- WOLFE-ORIGINS OF NORTHERN ROCKY MODERN VEGETATION 789 ern Rocky Mountain genera (e.g., A/nus, Populus, and Acer) are present in this vegetation, but the species represented are not closely related to ex- tant species in this regio Extensive vulcanism and associated tectonism resulted in a major upland region that extended from northern Nevada and central Idaho north into British Columbia during the middle Eocene (Axelrod, 1966a, 1966b); this upland may have actually been a series of upland volcanic centers. Known upland assemblages of early middle Eocene age (Fig. 1) occur from northeastern Washington (Republic and associated floras) northward into central British Columbia (Prince- ton, Chu Chua Creek, and coeval floras). Exten- sive vulcanism of early Eocene age became less intense during the early middle Eocene, and tec- tonism resulted in a series of grabens, in which the plant-bearing lacustrine sediments were de- posited (Pearson & Obradovich, 1977). Altitude of the Republic basin of deposition is estimated to have been about 900 m (Wolfe & Wehr, 1987). Later in the middle Eocene and in the early part of the late Eocene, major volcanic centers occurred in central Idaho (Challis Volcanics), central Oregon (Clarno Formation), and north- eastern Nevada (rocks containing the Copper Ba- sin and Bull Run floras of Axelrod, 1966b). Most Clarno assemblages represent only low altitudes, but a newly collected assemblage (Sheep Rock Creek) from the Crooked River Basin is probably microthermal and indicates the presence of higher altitudes. By the late Eocene, tectonism in the central Idaho region resulted in the formation of grabens and associated lakes on the post-Challis surface; these lake beds contain floras such as the Salmon. A similar depositional regime occurred in southwestern Montana (the numerous floras described by Becker); to the north in western Montana are plant-bearing lacustrine beds that have been collected recently by C. N. Miller and associates. The development of this upland region, con- comitant with a general decline of temperature following the early Eocene thermal maximum, tation, such as that at Republic (Wolfe & Wehr, 1987), represents the oldest known coniferous forest dominated by Pinaceae (Abies, Picea, Pi- nus, Pseudolarix, and Tsuga) and Cupressaceae (Chamaecyparis and Thuja). These genera were of low diversity in the early middle Eocene, but had increased in diversity by the end of the 790 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 p. S 7A S %Go fg» S $2» $ $2» Jgo € / D S x \ I . à j U f | 40* Ibe I JTA S r / Í 0 100 200 300 MILES ANA i s 2 0 — 100 200 300 400 — 500 KILOMETERS L Lf i : M TER j 38 L L — dll iz {- l $ $ è Š £ 3 A Š Š l I estern conterminous United FiGurRE 1. Map showing location of E i States por adjacent Canada. For each prei o i or group of PEREA sapi ig nae references or repositories of largely unpublished assemblages are cited. These repositories are: C (Geological Survey of Canada); OMSI (Oregon Museum of Science and Industry); TMP (Tyrrell M e 5. UAPC (University of Alberta Paleobotany Collections); UBC (University of British Columbia); UCMP (University of California Museum of Paleontology); UMPC (University of Montana Paleobotany Collections); USGS Geological Survey); UWBM (University of Washington Burke Museum). Early middle Eocene: 1. Smithers (UWBM); 2. Chu Chua Creek (Berry, 1926); 3. Kamloops assemblages (TMP); 4. Princeton assemblages (GSC, UAPC, UWBM); 5. Republic assemblages (Wolfe & Wehr, 1987; UWBM). Late middle to early late Eocene: 6. Sheep Rock Creek (OMSI); 7. Alvord Creek (Axelrod, 1944b); 8. Elko assemblages (UCMP, USGS); 9. Bull ( ); 16. Beaverhead, Metzel Ranch, Mormon Creek, Ruby, and York Ranch (Becker, 1960, 1961, 1969, 1972, 1973). 1987] Eocene. As discussed later, major diversification also occurred in numerous woody dic direi ous families. This probably resulted from (1) th areal restriction of microthermal climate Pian the early Eocene and (2) the opportunistic ex- pansion of mesothermal lineages into newly cre- ated microthermal climate in the uplan reas of microthermal climate were des highly restricted during the early Eocene thermal maximum. Indeed, I know of no latest Paleocene or early Eocene Northern Hemisphere plant as- semblage that represents microthermal vegeta- tion; possibly only on mountains in polar lati- tudes could such vegetation have been present (Wolfe, 1985, fig. 5). Because of the absence or extreme restriction of microthermal climates during the early Eocene, probably few pre-Eocene microthermal taxa were able to survive this cli- matic-geographic bottleneck, leaving the newly expanding microthermal regions available for l lineages. Further, to- bokranhie and edaphic diversity, Juxtaposition of many microclimates, and relative isolation of one volcanic center from another would all lead to rapid diversification of lineages in the uplands. Geographic proximity to lowland mesothermal vegetation would allow a continuing supply of new clades. robable instability of community composi- tion during the middle Eocene is indicated by two lines of evidence. First, from one approxi- mately coeval depositional site to another, species composition varies markedly; almost all the lo- calities are in lacustrine shales and presumably represent similar ecologic settings. Spatial het- erogeneity more resembled that of mesothermal or even megathermal vegetation than that of mi- crothermal vegetation. Second, floristic compar- isons of early middle Eocene lacustrine assem- blages with late middle to early late Eocene assemblages indicate large changes in composi- tion, some related to evolution of lineages within microthermal vegetation and some related to ad- ditions from mesothermal vegetation, as well as to extinction. Numerous mesothermal (or mesothermal/ megathermal) evergreen dicotyledonous clades had adapted to the microthermal climate by the early middle Eocene, although these typically are phytocrene (Icacinaceae), and Schoepfia (Olaca- WOLFE-— ORIGINS OF NORTHERN ROCKY MODERN VEGETATION 791 ceae). None of these genera are known to have survivedi l vegetation of the late norm and they can best be considered as un- S sf although other evergreen dicotyledons successfully adapted to the mi- crothermal climate during the Eocene. The pres- ence of evergreen dicotyledons in microthermal coniferous forests during the Eocene is consistent with the low (ca. 5°C) mean annual range of tem- perature inferred for coeval lowland assemblages (Wolfe, 1978). Modern microthermal coniferous vegetation of low-latitude, upland areas (e.g., Taiwan, Himalayas) also has many taxa of ev- ergreen dicotyledons. Also included in the upland early middle Eocene vegetation are numerous extant genera of trees and shrubs that contributed to broad- leaved deciduous and coniferous forests of the Neogene and Holocene, although not present in the northern Rocky Mountain region today. In- cluded are: Sassafras, Cercidiphyllum, Corylop- sis, Comptonia, Castanea, Fagus, Tilia, Ulmus, a, Photinia, Decodon, Rhus, and Aesculus. Overall, diversity among woody angiosperms appears to have been higher in the initial phase (early middle Eocene) of development of the mi- crothermal coniferous forests than later in the Eocene. At least 95 genera and 140 species of woody angiosperms are known in the Republic and Princeton floras, which are still not thor- oughly collected. In contrast, about 70 genera and 110 species of woody angiosperms are known in the extensively collected and described latest Eocene coniferous forests from western Mon- tana (Becker, 1960, 1961, 1969, 1972, 1973). This general decrease is particularly notable in presumed large trees, which were perhaps grad- ually replaced by newly evolved species of co- nifers. Very few of the extant northern Rocky Moun- tain lineages are recognizable by the early middle Eocene. The oldest known Betula clearly allied to the B. papyrifera—B. occidentalis complex oc- curs at Republic, as does an Acer allied to A negundo. However, the Acer, although possibly ancestral to A. negundo, is assignable to an extinct section (Wolfe & Tanai, 1987). Many other ex- tant native microthermal genera were also par- ticipants in this early middle Eocene vegetation (e.g., Alnus, Corylus, Ribes, Rubus, Spiraea, Cra- taegus, Prunus, and Cornus), but the species can- not be placed directly in the ancestry of extant northern Rocky Mountain lineages. The early middle Eocene microthermal flora 792 of western North America had numerous archaic elements in terms of Neogene or Holocene mi- crothermal vegetation. Other than the bond cessful tl ae ed, the vegetation had extinct genera of Trochodendraceae, Cercidiphyllaceae, aie a- melidaceae, Platanaceae, Fagaceae, Betulaceae, Rosaceae, and other families; about 40% of the genera are totally extinct (Wolfe & Wehr, 1987, unpubl. data By the late middle to late Eocene, upland mi- crothermal vegetation had been floristically al- tered as the result of: (1) Extinction, e.g., the unsuccessful thermo- philic experiments. (2) Gradual evolution in phylads, e.g., in a phy- lad ultimately leading to Chamaebatiaria (Rosaceae), the early middle and late Eocene members represent distinct genera. (3) Major diversification of early middle Eocene microthermal clades, e.g., whereas early middle Eocene Acer comprised 10 species and 3 extinct sections, late middle to late Eocene Acer comprised 35 species and 17 sections, 14 of which are extant (including the first members of sects. Negundo and Ma- crophylla). Adaptation of members of previously meso- thermal clades to microthermal climates, e.g., invasion of newly evolved species of Quer- cus, Mahonia, Salicaceae, and Leguminosae. — A ~x Rosaceae and Aceraceae underwent major di- versification in Eocene upland microthermal vegetation. Both are today primarily bee-polli- nated; entomophily would have been advanta- angiosperms that are abundantly represented in the fossil assemblages and that were presumably of fluviatile habitats: trochodendroids, hama- melidaleans, Fagopsis, most Betulaceae, Comp- tonia, and Ulmoideae. However, the fluviatile habitat and concomitant anemophily/abiotic dispersal syndrome did not lead to more than species-level diversification in most of these groups. The great majority of extant genera of woody microthermal angiosperms had evolved by the end of the Eocene, yet diversification in some genera and perhaps families had yet to occur: ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 (1) In Salix, all known Eocene species are re- ferable to subg. Salix. (2) Ericaceae were of low generic and specific diversity in the Eocene; thus far only Rho- dodendron (subg. Rhododendron) has been found in the upland Eocene vegetation, al- though several small Eocene leaves that lack diagnostic characters have been referred to Vaccinium. Only a few scattered Eocene palynological records of Caprifoliaceae s. str. are known, e.g., Diervilla in the latest Eocene of Alaska (E. B. Leopold, written comm., 1970). A few nondiagnostic, microphyllous leaves have been referred to Symphoricarpos. ~ w — OLIGOCENE AND NEOGENE Wide-scale deposition of lacustrine sediments apparently ended in the northern Rocky Moun- tains by the end of the Eocene (ca. 33 Ma)*. This termination presumably resulted from consid- erable uplift, leading to downcutting and erosion. At approximately the same time, a worldwide major climatic deterioration occurred (Wolfe, 1978), resulting in development of microther- mal, broad-leaved, deciduous forest at low alti- tudes of middle latitudes and broad-leaved de- ciduous and coniferous forests at high latitudes (Wolfe, 1985). This deterioration involved a de- cline in mean annual temperature and a major increase in mean annual range of temperature (Wolfe, 1978). Megafloras of Oligocene and Neogene age are not known from the northern Rocky Mountains. Wing (1987) and Leopold & Denton (1987) have therefore largely concentrated on well-known as- diee from the adjacent Columbia Plateaus Fig. 2). From the Oligocene into the middle Miocene (ca. 13-14 Ma), the Columbia Plateaus were occupied largely by Mixed Mesophytic for- est; during the middle Miocene, this forest was replaced by Mixed Coniferous forest (Wolfe, 1981a). That Miocene basalts of the Columbia River Group generally lap onto highlands to the east and north indicates that certainly the region of the northern Rocky Mountains was higher than = 4 Many workers place the Eocene-Oligocene bound- ary at about 37 Ma and would include the latest Eocene Miss discussed above in the early Oligocene. Which- ver placeme trelative or actual ee of the on or paleoclimatic changes. 1987] the Columbia Plateaus. Based on the tempera- ture relations of extant vegetational types (Wolfe, 1979), I infer that during the Oligocene through middle Miocene, the northern Rocky Mountains had a lower altitudinal belt of Mixed Northern Hardwood forest (Fig. 3) and an upper belt of igh Montane Mixed Coniferous forest (the vegetational type that generally occupies mesic sites within the present northern Rocky Moun- tains). The terminal Eocene temperature deteriora- tion resulted both in numerous microthermal lineages migrating downslope from Eocene up- lands into Oligocene lowlands and in extinction of many microthermal lineages. For example, the relationships of most Bridge Creek (early Oli- gocene) taxa are with Eocene upland taxa (MacGinitie, 1953) and not with Eocene high- latitude taxa. Among the Oligocene taxa that il- lustrate this relationship are: Cercidiphyllum crenatum, Castanea orientalis, Fagus pacifica, Quercus clarnensis, Alnus “‘carpinoides,” Betula aff. papyrifera, “Carpinus grandis," Mahonia simplex, Plafkeria sp., * Ulmus” brownelli, Ame- lanchier sp., Crataegus newberryi, Acer (Negun- do) sp., A. (Macrophylla) osmonti, A. (Lithocar- pa) sp., Bohlenia sp. (an extinct sapindaceous genus that = Dipteronia auct.), and * Holmskiol- dia" speirii. Some lowland Oligocene taxa were probably derived from mesothermal to mega- thermal taxa of the Eocene lowlands: Liqui- dambar, Platanus, Engelhardtia, and Paleophy- tocrene. Although such taxa are few, both a and Platanus were vegetationally mportant during the Neogene. None of the Bridge os lineages appear to be derived from a north- ern source. Although the Oligocene data base is not as Eocene include Fagopsis, some genera of ceae, and the sister genus of Acer (" Acer" arcti- cum group). Some Eocene microthermal lineages that participated in Oligocene vegetation of the Columbia Plateaus did not survive into the Mio- cene (e.g., “Zelkova,” Plafkeria, Bohlenia, * Holmskioldia"). Clearly, extinction continued to play a significant role. The Oligocene vegetation in high-latitude re- gions such as Alaska was of low diversity (Wolfe, 1972, 1985) and provided a limited reservoir for mid-latitude lineages. Only near the end of the Oligocene did taxonomic diversity increase in WOLFE—ORIGINS OF NORTHERN ROCKY MODERN VEGETATION 793 high-latitude regions, largely from migration from lower latitudes and some diversification of im- migrant clades; the influence at mid latitudes of the high-latitude flora was generally not felt until the middle and late Miocene. By the early to middle Miocene, the flora of the Columbia Plateaus had increased in diversity relative to the Oligocene. This diversity increase resulted from three major factors (Wolfe, 1972, 1977): (1) Adaptation of mesothermal lineages to mi- crothermal climates (e.g., Magnolia, some Lauraceae, and Exbucklandia). (2) Diversification of Oligocene microthermal pee (e.g., Fagus, Alnus, Ulmus, Carya, and Ace ( o ~x RE migration of Asian mid-latitude lineages into Beringia and subsequent south- ward migration onto the Columbia Plateaus (e.g., four lineages of Acer belonging to sec- tions Macrantha, Platanoidea, and Parvi- ora) Maximal diversity on the Columbia Plateaus and adjacent areas was reached during the middle Miocene (ca. 13-16 Ma) Notable are the few species shared between middle Miocene floras on the Columbia Plateaus and the Kilgore flora of Nebraska (MacGinitie, 1962). Even a putative sh erodentatum,” in sect. Negundo) is represented in either region by distinct subspecies. Further, most Columbia Plateaus species have only a dis- ant to ofeastern North America. Divergences between most western and eastern American lineages had probably oc- curred during, or at the end of, the Eocene, which led to a distinctive western American element (Wolfe & Tanai, 1980: 16-18). By the Miocene (if not the Oligocene), the Rocky Mountain re- gion must have formed an effective climatic bar- rier to migrations of most warm microthermal pianis (Laopoid & Pentan, R d species (Acer “ snecies cer = p ( het -— m j f Acer, which was apparently able to disperse from east- ern into western North America. Members of this series appear suddenly at middle latitudes of western North America during the early Mio- cene and have no Beringian record. Cladistic re- lationships and the fossil record both suggest dis- persal across the North Atlantic during the Miocene (Wolfe, 1981b; Wolfe & Tanai, 1987), and the two lineages of series Saccharodendron present in the western American Neogene are ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 & g ve, ç, o h ae AS +< thy, SOS. * S ii fi me N "d / \ MONTANA $ BIN ay ° 0 100 200 300 400 500 MILES Se L L L 1 L J < rey oe 0 100 200 300 400 500 KILOMETERS Wo L / yi L l 1 132° 130° 128° 126° 124° 122° 1987] WOLFE—ORIGINS OF NORTHERN ROCKY MODERN VEGETATION 795 Notophyllous broad-leaved evergreen forest Mixed mesophytic forest e temperatures z peli ateaus o Low montane - mixed coniferous forest Mixed northern hardwood forest Inferred early in nort Ocky jos MEAN ANNUAL TEMPERATURE *C High montane _ ene sa fore bibi early ne temperatures in in alt cs 1 20 25 MEAN ANNUAL RANGE OF TEMPERATURE °C FIGURE 3. gram showing inferred temperatures for the northern Rocky Mountains during the early Neogene. C Nora) and temperature relations of forests adapted from Wolfe (1979). Inferred temperatures bi for Columbia Plateaus and Beringian assemblages adapted from Wolfe & Tanai (1980) and Wolfe (198 1a). Arrows indicate direction of Neogene temperature trends. E 2. Map showing location of some Oligocene and early Neogene (> 13 Ma) plant assemblages in e eetnuk River assemblages (USGS); 2. Colorado Creek (USGS); 3. Eagle River (Wolfe et al., 1966); 4. Harriet Point, Harriet Creek, and Redoubt Point Pe 5. Douglas Mountain (USGS); 6. Sitkinak Island (USGS); 7. Kukak Bay assemblages (Knowlton, 1904; U nee 8. re i pcd (Wolfe in Lathram et al., 1965); 9. Sooke (LaMotte, 1935; GSC); 10. Gu n untain (UCMP); 11. Cascade Locks (USGS); 12. Sandstone Creek (USGS); 13. Cascadia (UCMP); 14. end: M 1973); 15. eee (UCMP); 16. Rujada (Lakhanpal, 1958); 17. Yaquina (McClammer, 1978); 18. ap City (USGS); 19. Bridge Creek assemblages (Chaney, 1927; OMSI, UCMP USGS). Early Neogene: 20. Nenana coal field assemblages (Wahrhaftig et al., 1969); 21. Lower Cache Creek (Wolfe et al., 1966); 22. Houston (Wolfe et al., 1966); 23. Capps Glacier (Wolfe, 1966; USGS); i i 25. € : : ; 33. Clarki (Smiley et al., 1975); 34. Orofino Creek (Brown, 1940); 35. Whitebird (Berry, 1934); 36. Eagle Creek assemblages (Chaney, 1920); 37. Collawash and Fish Creek (USGS); 38. Little Butte Creek (USGS); 39. Mascall assemblages (Chaney & Axelrod, 1959); 40. Baker (USGS); 41. Skull Spring (USGS); 42. Succor Creek assemblages (Graham, 1964, in part; cf. Fields, 1983); 43. '49 Camp (LaMotte, 1936); 44. Rabbit Hole (USGS); 45. Eastgate (Axelrod, 1985); 46. Middlegate (Axelrod, 1985); 47. Goldyke (UCMP); 48. Fingerrock (Wolfe, 1964); 49. San Antonio (UCMP); 50. Thurston Ranch (UCMP). 796 inferred to have crossed the Rocky Mountains; consistent with such crossings are the present distributions of the related 4. brachypterum and the descendant A. grandidentatum, both of which live in the Rocky Mountains today. Thus some probably very limited floristic interchange be- tween eastern and western North America oc- curred during the Neogene. In the absence of actual assemblages, the flora of the northern Rocky Mountain region during the post-Eocene can be inferred only from as- semblages outside the region. Except for floras to the west and southwest on the Columbia Pla- teaus, however, floras from adjacent regions are few. To the north in central British Columbia, the probable early Miocene assemblage from near Hanceville on the Chilcotin River appears to rep- resent a High Montane Mixed Coniferous forest. In southern Colorado, the late Oligocene Creede assemblage represents subalpine coniferous vegetation. In Nebraska, the middle Miocene Kilgore assemblages largely represent a meso- thermal gallery forest (MacGinitie, 1962) Taxa that occurred both in Mixed Northern Hardwood forest of Alaska and in Mixed Me- sophytic forest of the Columbia Plateaus repre- sent taxa that (1) could exist under temperatures inferred for the northern Rocky Mountain region and (2) are known to have occurred adjacent to the northern Rocky Mountains. Such taxa thus can be reasonably inferred to have been in the northern Rocky Mountain region. Included in this category are lineages that are extant in the northern Rocky Mountains: Alnus aff. incana, A. aff. viridis, Betula aff. papyrifera, Populus aff. trichocarpa, Salix aff. lasiandra, Prunus aff. vir- giniana, Acer aff. negundo, and A. aff. macro- phyllum. Also included are species of now extinct lineages: Nordenskioldia ("Cocculus" auricula- Carya bendirei, Pterocarya nigella, Acer scottiae, A. septilobatum, A. tigilense, and Nyssa knowl- tonii. Both lists contain primarily arcto-tertiary genera, genera that are now disjunct between eastern Asia and eastern North America or gen- era that are now characteristic of forests of north temperate regions (Engler, 1879). These genera (and many subgeneric groups), however, were represented in the Eocene upland microthermal vegetation; their inferred presence in the north- rn Rocky Mountain region is not necessarily the ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 result of southward migration from high lati- = (= es. Composition of the northern Rocky Mountain coniferous element during the Oligocene and Miocene must also be inferred. In the middle Miocene of Nevada, the conifers are primarily those that now inhabit Low Montane Mixed Co- niferous forest (e.g., Abies aff. concolor, Picea aff. breweriana, Pinus aff. monticola, P. aff. ponder- osa, Tsuga aff. heterophylla, and Chamaecyparis aff. nootkatensis). Most of these conifers were also dominants of the Columbia Plateaus vege- tation between 10 and 13 Ma (Fig. 4). In at least one instance, the fossil species is a sister species to the lineage that gave rise to the extant species (Edwards, 1983), suggesting that the fossil may have been adapted to a different environment than the extant species. In the Beringian Neogene, most coniferous lin- eages were distinct from mid-latitude lineages. The Abies belong to a group of which the extant A. sibirica and A. grandis are members. Larix was a diverse and major element in the Beringian Neogene; no valid Larix is known during the Miocene at middle latitudes of western North America (H. E. Schorn, pers. comm., 1984). The Beringian Picea either are related to extant Asian species or represent the P. glauca group (includ- ing an extinct, large-coned species that survived into the Wisconsin glacial of southeastern North America according to Critchfield, 1984). Thus, the conifers (Abies grandis, Larix, and Picea glauca) that are now the most distinctive element of the northern Rocky Mountain region relative to other regions of the western United States ap- pear to be derived from a high-latitude source. Precisely when the Beringian coniferous ele- ment arrived in the northern Rocky Mountains is unknown. I suggest, however, that the arrival was probably post-Miocene. Coniferous lineages allied to taxa that are now restricted to Low Mon- tane Mixed Coniferous forest occupied the Co- lumbia Plateaus and Nevada during the Neogene and presumably represent ecotypes distinct from extant relatives (Wolfe, 1964); such extinct eco- Neogene, many of these northern tain conifers possibly became extinct as the Be- ringian lineages migrated south. Perhaps signif- icant is that the first record of the coastal ecotype of Abies grandis at middle latitudes is in the Plio- cene-Pleistocene Sonoma assemblage of Cali- fornia (Axelrod, 1944). 1987] Various dicotyledons also may have penetrat- ed southward from high latitudes during the Neogene. All North American (including Be- ringian) Eocene Salix represent subg. Salix. In Beringia, subg. Vetrix appears in the late Oli- gocene, approximately coincidental with the first appearance of other taxa of Asian affinities; members of sect. Glauceae appear in Beringia by the early Miocene. Sect. Retusae of subg. Cham- aetia appear by the end of the Miocene. Skvort- sov (1968) suggested that subg. Vetrix and subg. Chamaetia were of Asian origin, based on the present distribution of species that have inferred primitive morphologies. Although Asian fossil data are lacking, certainly the Beringian data are consistent with Skvortsov's interpretations. Dur- ing the Neogene, various members of subg. Ve- trix and sect. Glauceae underwent diversification at high latitudes. A few lineages penetrated southward in North America by the late early Miocene and middle Miocene, but most lineages of subg. Vetrix and subg. Chamaetia now extant at middle latitudes of western North America are probably late Neogene immigrants. Representatives of Caprifoliaceae such as Diervilla, Lonicera, and significant and diverse elements i in the Beringian Miocene, as were Ericaceae such as Rhododen- dron (subg. Anthodendron — Azalea) and Vac- cinium. These are generally absent in Neogene floras at middle latitudes, although Symphori- carpos is known in the late middle Miocene of Nevada (Axelrod, 1956) and the late Miocene of Idaho (Chaney & Axelrod, 1959). Presumably most Caprifoliaceae and Ericaceae also represent late Neogene immigrants into the northern Rocky Mountain region. Some extant northern Rocky Mountain Ro- saceae (e.g., Amelanchier) appear to be derived from lineages that have lived in or near the north- ern Rocky Mountain region since the Eocene. Other Rosaceae, however, probably belong to the late Neogene Beringian element. Although Ru- bus was represented in the Eocene upland mi- crothermal vegetation, the extant R. idaeus has a probable ancestor in the Beringian Miocene. Similarly, Prunus subg. Padus has a continuous record in western North America during the Eocene and later, but subg. Prunophora first ap- pears in western North America in Beringia dur- ing the Miocene, and the extant Pacific North- west Prunus subcordata appears to represent a late Neogene immigrant from Beringia. A pattern somewhat similar to that of Prunus wUIC also WOLFE—ORIGINS OF NORTHERN ROCKY MODERN VEGETATION 797 is indicated for A/nus. Members of subg. A/nus were present in the northern Rocky Mountain region as early as late Paleocene, and all the up- land Eocene A/nus represent this subgenus. The mid-latitude members of the subgenus differ- entiated by the Oligocene into a lineage leading to the extant A. oregona, and the lineage is clearly recognizable in the early Miocene of Oregon. High-latitude members of the subgenus are rec- ognizable as belonging to the A. crispa lineage by the Oligocene, and this lineage had penetrated into middle latitudes by the middle Miocene. Members of subg. A/nobetula are high-latitude in distribution through the Miocene, and the en- try of the extant A. viridis into middle latitudes probably occurred during the late Neogene. DISCUSSION The terminal Cretaceous event selected for de- ciduousness and created a mesothermal region in the Northern Hemisphere almost devoid of broad-leaved evergreens, allowing by default oc- cupation of this region by many deciduous groups. These groups then underwent major generic-level diversification. The elimination (or nearly so) of microthermal climates by the early Eocene ther- mal maximum and the following creation of mi- crothermal, mid-latitude uplands of the later Eocene provided the opportunity for adaptive radiation of these denuo mesothermal clades into the new opportunity for continuing diversification. Fun- damentally, microthermal ecosystems became extinct during the early Eocene thermal maxi- mum and arose again de novo following the ther- mal maximum. The continuing and major al- terations in floristic composition during the Eocene, the apparent high degree of community instability during at least the early middle Eocene, and the rapid diversification of groups such as Rosaceae and Aceraceae during the Eocene can be viewed as symptomatic of the evolution of this new ecosystem. Insofar as known, the west- ern North American volcanic uplands made up the only major, mid-latitude upland region in the Northern Hemisphere during the Eocene. These volcanic uplands thus occupy a central place in the diversification of many (if not most) microthermal clades of arcto-tertiary type. Primarily during the middle and late Eocene, many microthermal, arcto-tertiary lineages were able to disperse readily from North America into Eurasia as adjuncts of a continuous expanse of coniferous forests (Wolfe, 1985, fig. 10). Partic- 798 ANNALS OF THE MISSOURI BOTANICAL GARDEN De FS To we Sy, Sy District NORT of M *6kenjj, ~ [Vor. 74 : é 2 eal! E $ 4 5 KA ud / AN UN 1 Tee ol / EN el ` P ° e / 3 MONTA N4 rd 1840. 2»?! i d ( $ = MT x Mrs ri P € 23 EN end \ i f WYOMING ay J À ez. : o ow m x « s MILES Í | / n d. S + T. T U š J g NEVADA 0 100 200 300 400 500 KILOMETERS N Ó Jo} I! EY UTAH j — — . x / | ° 7 j % l: fi: COL ORA g L % \ L 4 do d Jo gd Ip. È Í Ë Sivan aN 132° 130° ne ne me ur — 120% — "gf — ne 1 — no one 108° 1987] ularly with the general cooling that characterized the later Eocene (Savin, 1977), microthermal cli- mates were found at increasingly lower latitudes and produced a greater area available for occu- pation by microthermal vegetation. As migrating lineages elaborated over this area, diversification would continue and some new divergences that occurred in other regions would probably, in turn, migrate into North America. Thus, even in the Eocene, complexities were probably introduced into historical patterns of a given lineage. Histories of the component lineages now in the flora of the northern Rocky Mountains are varied. (1) Some lineages can be traced back, more or less continuously, into microthermal vege- tation of the Eocene uplands (e.g., Betula pa- pyrifera complex). Some lineages have generic representation in this upland vegetation, but divergences that led to the extant species probably occurred in the Oligocene or Neogene at middle lati- tudes (e.g., A/nus oregona) and some at high latitudes (e.g., A. viridis). Some lineages represented in the Eocene up- lands probably dispersed into Asia, under- went major divergences into new subgenera, ~ N — ~ o2 — en- tered the northern Rocky Mountain region in the late Neogene (e.g., species of Salix subg. Vetrix). Some lineages represented in the Eocene up- lands dispersed into Eurasia, underwent di- versification at the sectional level and re- turned to North America via long-distance dispersal from Europe (e.g., Acer series Sac- charodendron) ~ + — WOLFE—ORIGINS OF NORTHERN ROCKY MODERN VEGETATION 799 (5) Some lineages have long Beringian histories and entered the northern Rocky Mountain region during the late Neogene (e.g., Lonic- era (6) Another major pattern must be inferred for many alpine tundra plants; Arctic tundra is not recognizable until about the Pliocene- Pleistocene boundary (Wolfe, 1985), and ele- ments shared between Arctic tundra and al- pine tundra ofthe northern Rocky Mountain region may have entered the northern Rocky Mountains during the Quaternary (7) Another pattern must be inferred for plants of subhumid to xeric, southern affinities (Barnosky, 1984; Leopold & Denton, 1987); some of these belong to families that did not originate until about 25 Ma (Muller, 1981). he Neogene spread of Compositae was rap- id, probably in relation to the high degree of dispersibility of their diaspores; however, the spread appears to be associated also with the development of dry climates. Pollen indicates that Sarcobatus (Chenopo- iaceae) was a member of mesic coniferous forest during the later Miocene in areas such as southern Idaho (Wolfe, 1969); adaptation of Sarcobatus to xeric climate may be a late Neogene phenomenon. ~ oo — No single pattern explains the origin of the majority of the modern flora of the northern Rocky Mountain region or even the majority of the flora of a given vegetational type or zone in the region. Certainly late Neogene cooling Aprap- ably along with mates in the northem Rocky Mountains fy able to Beringian taxa, but these are intermixed with taxa whose ancestry goes back to the Eocene in the northern Rocky Mountains. Further, Be- — FIGURE 4. Map showing location of some late Neogene (<13 Ma) plant assemblages in western North America. For each assemblage or group of assemblages, either published references or repositories of largely L. assemblages are cited; see Figure 1 for explanation of reposito ., 1971); 2. McCallum Creek (USGS); 3. Grubstake (Wahrhaftig et al., ., 1966); 5. Chuitna River (Wolfe, 1966); 6. Tyonek (Wolfe et al., 1966); 7. Assemblages of type and referred : " W. . ries. 1. Hoogendoorn 1969); 4. : gue Bay (USGS); 18. Hidden Lake (USG nem Tipton id Vinegar Creek (Chaney, 1959; Chaney cf. Fields, 1983): 27. Thorn Creek (Smith, 1941); 28. Trapper Creek (Axelrod, 1964); 29. Cache Valley ‘ark 1949): 30. Pit River (LaMotte, 1936). 800 ringian taxa entered at different times during the Neogene and even Quaterna Despite considerable resemblance at the ge- neric level to the modern flora of the northern Rocky Mountains, the microthermal upland vegetation of the Eocene had many genera that are now extinct and many genera that are now exotic to the region. Microthermal vegetation of western North America experienced major flo- ristic alteration during the Eocene and later in response to various extrinsic climatic factors and intrinsic biotic factors such as competition. The primary floristic result of the terminal Eocene temperature deterioration for micro- thermal vegetation was the elimination of many dicotyledonous genera, particularly those con- taining evergreens. Many deciduous dicotyle- dons, however, also suffered extinction. For ex- ample, numerous lineages in Acer became extinct, including all North American members of sec- tions Acer, Platanoidea, C ampestria, and Mac- rantha (all of which survived in Eurasia). Pat- terns of survival among decus taxa differed markedly between Eurasia and North America, probably due to intrinsic factors. The relatively depauperate flora of the early for radiation and diversification, with : a general increase in diver- sity and blurring of the regional Oligocene ge- neric and subgeneric endemism, especially be- tween western North America and eastern Asia; for example, both Platanoidea and Macrantha reappeared in western North America during the Neogene. Migrations across Beringia were en- hanced by the mid-Miocene warm interval, when broad-leaved deciduous forests were probably continuous from the Pacific Northwest across Beringia and into P latitudes of eastern Asia (Wolfe, 1985, fig. 1 Cooling at about n Ma (late middle Miocene), concomitant with a general Neogene decline in summer temperatures and decline in mean an- nual ranges of temperature (Wolfe, 198 1a), elim- inated many broad-leaved taxa of trees and shrubs from the Columbia Plateaus, Beringia, and pre- sumably the northern Rocky Mountains. Conif- erous forests probably occupied a continuous belt from the Columbia Plateaus and the northern Rocky Mountains north into Beringia. Mixed Northern Hardwood forest would have been to- tally eliminated from western North America. Climates in the northern Rocky Mountain region during the Neogene may have initially exhibited the trend toward lower mean annual ranges of ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 temperature. However, by the late Miocene, in- crease in altitudes of mountains along the Pacific Coast would have countered this trend, allowing incursions of Arctic air masses; these masses are inferred to have influenced eastern North Amer- ica after 13 Ma (Wolfe, 1985). During the Neogene a summer-dry precipita- tion regime gradually encroached from the south (Wolfe, 1978), and during the late Miocene (ca. 5-11 Ma) increase in altitude of the Cascade Range created a significant rain shadow east of the range (Chaney, 1944c; Smiley, 1963). At low- er altitudes on the Columbia Plateaus, forests were replaced by steppe vegetation during the late Miocene (Leopold & Denton, 1987; Wolfe, 1985). An analogous replacement was occurring in the northern Rocky Mountain region (Bar- nosky, 1984). Considering the complexities of climatic changes and the complexities of orogenic activity in the northern Rocky Mountain and nearby re- ions, determination of historical biogeographic patterns, without recourse to the fossil record of extant lineages and analyses of successive fossil biotas, can be highly problematic. As Grande (1285) emphasized, vicanance methodology as typically applied complexities; several patterns can ine artis on one another in such a region. Numerous pat- terns have been detected in the modern flora of the northern Rocky Mountains by recourse to the fossil record, and different species within the same genus can represent different patterns. Lin- eages, some closely related, can be subjected to the same extrinsic environmental factors and can respond in different ways, a fact that further in- creases the general complexity of the biogeo- graphic tapestry in a region such as the northern Rocky Mountains. LITERATURE CITED ALVAREZ, W., L. W. ALVAREZ, S drcum & H. V. Mı- CHEL. 1984. The end o retaceous: sharp boundary or gradual iiie ° Science 223: 1183- 1186 AXELROD, D. p 1944a. The Sonoma flora. Pp. 167- 206 in R. W. Chaney (editor), Pliocene Floras of California and Oregon. Publ. Carnegie Inst. Wash. 553 š 1944b. The Alvord Creek flora. Pp. 225-262 in R. W. Chaney (editor), Pliocene Floras of Cal- ifornia and Oregon. Publ. Carnegie Inst. Wash. 553. — 6. Mio-Pliocene floras from west-central Nevada. Univ. Calif. Publ. Geol. Sci. 33: 1-322. —— 1964. The Miocene Trapper Creek flora of 1987] southern Idaho. Univ. Calif. Publ. Geol. Sci. 51: 1-181 1966a. The Eocene Copper Basin flora of northeastern Nevada. Univ. Calif. Publ. Geol. Sci. 9: 1-125. 1966b. A method for determining the a tudes of Tertiary floras. Palaeobotanist 14: 14 171. 1985. Miocene floras from the Middlegate Basin, me? central Nevada. Univ. Calif. Publ. Geol. Sci. 79. BARNOSKY, C. " 1984. Late Miocene vegetational and climatic variations inferred from a pollen rec- ord in northwest W BECKER, H. F. ins from the upper Ruby R ern Montana. l s mites Abt. B, Pa- Misit 107: 83- 61. Oligocene plants from the upper Ruby River Basin, southwestern Montana. Mem. Geol. mer. 82: 1-1 7 Palaeon- Ruby River Basin, southwestern e Palae- ontographica, Abt. B, Palaophytol. 141: 1-61. 1973. The York Ranch flora of the upper R uby River Basin, southwestern Montana. Palae- ontographica, Abt. B, Prise viel 143: 18-93. BERRY, E. W. 1926. Tertia ry floras from eee Co- a 42: 91-116. Formation. U.S. Geol. Surv. Prof. Pap. 154-H: 225-265 . A Miocene flora from Grand Coulee, Nari U.S. Geol. Surv. Prof. Pap. 170-C: 31-4 TM Miocene plants from Idaho. U.S. Geol. Surv. Prof. EE 185-E: 97-125. Bouor, B. F., E. E. FooRp, P. J. MODRESKI & D. ai TRIPLEHORN. 1984. M dence impact event at the ossis a M s Bonds ary. Science 224: 867-869. BRowN, R. W. 1937. 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ABSTRACT The present-day northern Rocky Mountain vegetation 1s the product of a long history of geologic ith and climatic events that have interacted w General concepts relating to the organization, classification, an the species populations composing the regional flora. nd dynamic nature of vegetation are reviewed. Distributional and structural features of the vegetation cover between the Colorado Rockies and the Southern Canadian Rockies are discussed. Alpine, upper timberline, subalpine, montane, lower timberline, and grassland/steppe zones are treated. Climatic, physiographic, edaphic, and geo- Ulltaill vegetation. Iti is likely that members of the modern Rocky Mountain flora are not in Fani br Qui with o tion. Present etati t pla re altere structures and compositions that may represent new ecosystem equilibria acs could be irreversible under present-day climates. Vegetation ecology encompasses the descrip- tion and interpretation of distribution patterns exhibited by plants and plant communities oc- cupying a given geographical area (Miles, 1979; McIntosh, 1985; Walter, 1985). The objective of this symposium is centered on an interpretation ofthe evolutionary development ofthe Northern Rocky Mountain (NRM) flora and vegetation, including the earliest origins and ages evidenced. This effort closely complements the reviews of the origins of the cordilleran flora (Great Basin and southern Rocky Mountains) provided by Reveal's (1979) and Axelrod & Raven's (1985) biogeographic treatments of the western inter- mountain region, and Peet's (1988) detailed analysis of structure, distribution, and produc- tion of forests from throughout the Rocky Moun- tains. My description of NRM vegetation serves to orient readers to other symposium contribu- tions that consider the roles of past events in shaping this region's flora and vegetation. THE NATURE OF VEGETATION Literature on vegetation science reminds us of the dynamic nature of floras and plant com- munities (Knapp, 1974; Mueller-Dombois & El- lensberg, 1974; Miles, 1979; Chabot & Mooney, 1985; McIntosh, 1985). Many botanists engaged in the study of floras and plant communities agree that vegetation units associated with any land- scape arise over time as chance combinations or assemblages of plant populations, with each species following more or less independent or individualistic ecological responses and migra- tional pathways, as Gleason (1926) hypothe- sized. Such ideas have been substantiated by Whittaker (1953), Curtis (1959), McIntosh (1985), and others over the past half century. In his recent remarks addressing vegetation history in the northern Great Basin and Pacific Northwest, Mehringer (1985: 168) stated, “Plants responded as vagaries of climate, dispersal po- tential, competition, selection, soils, topography, volcanic eruptions, fire, man and chance dictat- ed. A hazy outline of these responses has been traced through pictures painted with pollen ex- tracted from yesterday’s mud. Sketchy as the im- ages may be, they prove the dynamic nature of this area’s vegetation.” Floristic changes over time, whether gradual or abrupt, underlie the dynamic nature of the NRM vegetation. Post-Pleistocene pollen se- quences in this region reveal “unquestionable fluctuations in vegetation wrought by short, sharp climatic episodes, by fire and by volcanic erup- tions” (Mehringer, 1985: 185). Wells (1983), analyzing wood rat middens throughout western ! Drs. Stephen Arno, Stephen Cooper and Robert Peet provided useful si bad pilus inam technical vided review of early stages of the manuscript; valuable suggestions were also provi by tw onymous reviewers d. The contributions of Dr. Charles N. Miller, University of Montana, are also Rios ? Department of Botany, University of Montana, Missoula, Montana 59812, U.S.A. ANN. MISSOURI Bor. GARD. 74: 804-840. 1987. 1987] North America, provided descriptions and his- torical documentations of the ee ic shifts of western vegetation Davis (1976, 1981) and mimi ne in their analyses of forest vegetation stability during the Quaternary, also addressed postglacial vegeta- tion dynamics and suggested that eastern North American plant populations are displaying dif- ferential rates of range reoccupancy following the Wisconsin glacial phase. Both Davis and Webb urged us not to assume that an equilibrium exists between modern plant distributions and the ir climate. The NRM assuredly experi- enced its own Pleistocene stresses and phytogeo- graphic rearrangements, and it is likely that the modern flora and vegetation in this area are not in equilibrium with modern climate. merous environmental variables have in- fluenced the NRM flora and vegetation over time. Lassoie et al. (1985), Smith (1985), and Peet (1988) provided summary interpretations of the ecophysiology of Pacific Northwest and Rocky Mountain conifer forests. Factors such as moun- tain building, differences in soil chemistry and physical structure, precipitation and temperature gradients, seasonal surpluses and water deficits, evaporation potentials, and wildfire, all inter- acting with each other and locally conditioned by physiographic factors, create a multitude of specific habitats for plant occupancy. Over time, plant populations sort themselves out differen- tially among available habitats. The spatial or geographic pattern (local and regional range lim- its) of each species is at least partially based on adaptive features. The physiological studies cited above emphasize how well the Rocky Mountain conifer species are adapted and adjusted to their environment. Within the context of her eastern North Amer- ican studies, Davis (1981) supported the view that species with long histories of co-occurrences in similar plant communities or habitats have co-evolved adaptive adjustments to a. in- terspecific competition. Davis warned, however, that it is important for ecologists to AA assess as to whether sufficient time has elapsed for coevolution to have happened among closely associated plant species. Often these sorts of as- sessments represent research that must still be undertaken on our western vegetation types. Each plant population is unique in its degree ofadaptive expression or amplitude of tolerance. This uniqueness, founded in the genetic consti- tution ofa species, is reflected in its rates of range HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 805 extension or retreat, its present ecologic range limits, and its specific habitat. Differential rates of migration will affect vegetation distribution patterns, as well as the presence or absence of a species in a given vegetation association. The dynamic nature of plant populations and plant communities requires that ecologists recognize that they are dealing with a vegetational phe- nomenon that exhibits short-term stability but lacks long-term permanency. Rocky MOUNTAIN VEGETATION STUDIES The earliest efforts to provide descriptive sum- maries of Rocky Mountain vegetation date back to the turn of the century. C. Hart Merriam (1889), a prominent figure in the history of North Amer- ican vegetation ecology, developed a system of life zones that stressed the importance of tem- perature in explaining vegetation patterns. He popularized such life zone terms as Boreal, Tran- sitional, Canadian, and Hudsonian. ui of Merriam's scheme is used today. P Rydberg, an early botanist associated with the New York Botanical Garden, made major contributions to our understanding of the phytogeography of the Roc Bulletin of the Torre 1913 and 1917). Rydberg's typical reporting for- mat involved listing the flora of each mountain zone with its geographic origin and limits. J. irkwood, pioneer botanist at the Uni- duit of Montana, published (1918, 1922) some of the first in-depth ecological interpretations of northern Rocky Mountain forest compositions and distributions, applying to this region much of the basic ecological insight promulgated by Schimper (1903). Kirkwood provided a detailed literature review of NRM vegetation studies available at that time. He clearly emphasized that vegetation distribution patterns would be interpreted best when the “‘individualities” of the “constituent species" were understood, an idea that Gleason (1926) also advocated. Kirkwood stated that “the complex interaction of climatic, edaphic, biotic and other influences that guide and control the distribution of species is of keen interest to the student of phytogeography, and no treatment of the subject is fair that does not seek to give full weight to the many factors in- volved and especially to the nature and require- ments of the individual species" (1922: 15). He also pointed out that Merriam's life zones often ANNALS OF THE MISSOURI BOTANICAL GARDEN 110° A ae A “ey ake ZA 2 gs “GE XZ V Z A Z í > wasife ye # & ORE Z Z 1! 22 SES r T AT S =, © SS ws Z Se sss 45^. 27 SS MT A Ë A T 245? IDA WYO NE UTAH CO 110° + SES C $/ A/S —————À NADA A 3 E USA C Wo) WASH SX ORE ST 1594. MT 445° IDA WYO NEV [UTAH : CO FIGURE l. ug cedar-hemlock-pine reste ae plieaie. Tsuga heterophy a and Pinus monticola).— il). [Vor. 74 am o y YY 4 777 72) 4 yj ⁄ 77, (T, / ft 7 Tp A Mo CM e, "n" CIM diy fC UA LZA p^ (9 ay, fi, 7 jji ⁄Z wo AG 4 4 P g 7 OL, ü M ^ A 4 4, 7 yh 4 yh IDA 7 // 4, 7/4/72 / ZZ 77 Saker ZZ 4 "n (A A Mf Wy ", V up lt My 4 uid! + 45° + 110° 110° + 72 M IP © why ys iii pes iff! 4 4 My ” 27%, ^ ros Kuchler’s K) n natural vegetation types shee with ionem, ois listed. — A. K-1 western ponderosa yes d astern ponder ponderosa var. scopul rum).— s ponderosa d ou Douglas-fir (Abies grandis and Pseudotsuga menziesii lacked spatial distinction (discontinuity) because of elevational intermingling of the conifer dom- inants in the nort The many contributions made by R. F. Dau- benmire to our knowledge of northern Rocky Mountain vegetation must also be recognized. In a career spanning nearly a half century, Dauben- mire provided historical interpretations of the origins of the Rocky Mountain flora, studied the ecophysiological features of NRM plants, and was instrumental in developing the habitat type ern Rockies. var. ponderos las-fir forest usine uga menziesii bm pou —C. D. K-14 = grand fir & ; K-16 = sa pine forest n" K-13 = classification system widely employed in the Rockies (1943, 1966, 1968, 1969, 1970, 1975, 1978, 1980, 1981). Daubenmire’s classic review paper, “Veget ation Zonation in the Rocky Mountains’* (1943), summarized 175 ecological studies that bear on the extant vegetation of the Rocky Mountains. He reviewed floristic origins and provided interpretations of relationships be- tween vegetation zonation and mountain cli- ates. Following these pioneers, many recent efforts 1987] HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 807 110° 110° u + B CANA N š DA ES P UNE USA a DUM. e EN > WN m ad X RS (60) S š $` xš SS = “S Z S Sc wasi z Pu SUE fo) Z= £ NES = J X S NES RRS ¿ SsX a p^ XE PF S s SER ° Aw SS 459. ` 45 + as SS RC - MT x e> Ss EN ~ Yas ` SR IDA S WYO NEV JUTAH A NEV JUTAH , CO 1109 1109 1109 u C CANADA N SA oO XN 7 EO M 2 * VEZ x & š Z eS. 3 x. " E NSS S 45° SS SN NN M < ia NAGI Trg SIX Ss ° X N SX WW . * & PRENSA s Faso E S /S | @ ° IDA ~ Su = S SHE WE SS = ES Š SS ` l A š = = WYO ENSE ES NEV JUTAH n CO Ficure 2. Kuchler vegetation types.—A. K-15 = western spruce-fir forest (Picea engelmannii and Abies lasiocarpa). — B. K-50 = fescue-wheatg (Agrop picat d Festuca idahoensis); K-63 = foothills prairie (Agropyron spicatum, Festuca idahoensis, F. scabrella, and Stipa comata).—C. K-51 = wheatgrass-bluegrass (Agropyron spicatum, Festuca idahoensis, and Artemisia tridentata), K-55 = sagebrush steppe (Agropyron spi- catum, Artemisia tridentata). — D. K-56 = wheatgrass-needlegrass-shrub steppe (Agropyron smithii, Artemisia tridentata, and Stipa comata); K-64 = grama-needlegrass-wheatgrass (Agropyron smithii, Bouteloua gracilis, and ) Stipa comata). have been made by resource managers, plant community types most likely to express them- ecologists, and geograr to describe, interpret, selves in the absence of human influence. Kuch- map, and classify parts or all of the Rocky Moun- ler realized that the presence or absence of wild- tain vegetation (Kuchler, 1964; Daubenmire & fire and/or grazing alters the specific nature of Daubenmire, 1968; Pfisteretal., 1977; Peet, 1981, vegetation cover; this complicated his mapping 1 ; Arno & Hammerly, 1984; Smith, 1985). efforts. Figures 1, 2, and 3 show the general dis- Kuchler (1964) published a map of what he tributions of Kuchler’s northern Rocky Moun- termed the “potential natural vegetation" of the tain vegetation types. coterminous United States. His map portrayed Pfister et al. (1977) and Pfister (1984) provided the present-day vegetation in terms of those detailed literature reviews of the vegetation clas- 1 [Vor. 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN 777777 NN N BZ Z 110° u C CA N NADA c u C WASH ORE 459J. MT 4t T2459 IDA 3 8 R3 S b. = * SS š š ois 3” RE > = p> SERES Il > > a £ S s P) — STE wyo RON > = ` a SENS šs* S SS .S8 AHg oS E S X SO MSS = AS SS ju | s ENS S SA? o rer & ` IAJN xs T S NE V UTAHSS a> SVS SS cC 110 Kuchler vegeta vermiculatus). — B. Greatbasin sagebrush (Arte osteosperma, Pinus edulis (e sification efforts in the R misia tridentat tion types.— A n ocky Mountains. Clas- sifications are often individualized by each au- thor’s philosophy and objectives, and the absence of universally accepted concepts and principles tion approach. Many R tion classifications have source management, map storage, and vegetation preservation. These ap- proaches often assume the existence of repeating combinations or assemblages of plant popula- been developed for re- ping, information . K-40 = saltbush-greasewood (Atriplex confertifoli erosa pine-Douglas-fir (Pinus ponderosa and Pseudotsuga menziesii), K-38 = a).—C. K-23 = juniper-pinyon woodland (Juniperus monosperma, ast), P. monophylla (west), and Quercus spp.).—D. K-3 oak scrub (Cercocarpus ledifolius, Quercus gambelii, and Acer gran : a and Sarcobatu -37 = mountain mahogany- ss didentatum) tions, in association with more or less specific site qualities. Pfister (1984) supplied a recent concise sum- ese hav categories called “habitat types": physical set- tings definable in terms of the potential climax vegetation they are capable of supporting (Dau- benmire, 1966). Habitat type classifications cov- er large parts of the Rocky Mountains. However, 1987] th 1 245 ti affnrtch licited some crit- ical commentary from other ecologists who have studied in the same regions (Antos & Habeck, 1981; McCune & Antos, 1981; Baker, 1984; Cloonan & Habeck, 1985; Crawford & Johnson, 1985; McCune & Allen, 1985a, 1985b). Baker (1984), working with the Colorado Nat- ural Heritage Inventory, maintained that vege- tation classification in the Rocky Mountains is handicapped by the absence of a standardized nomenclature. He suggested, however, that no unique classification may be possible because of the vegetational variability, as noted above. Ba- ker provided his own classification of Colorado’s natural vegetation, a hierarchical listing of plant community types. Peet (1981) successfully em- ployed gradient analysis and ordination tech- niques to the Colorado Front Range vegetation and derived a workable classification of forests in that area. An outline of Peet’s results will be presented later. My literature evaluation revealed that regional climatic patterns, physiographic provinces, ele- vation zonations, soil features, topographic- moisture gradients, habitat types (potential cli- max), seasonal moisture regimes, drought stress categories, plant life forms, community life forms, ecological interactions between dominants, as well as human intuition, have all been used to design classifications of Rocky Mountain vege- tation. Furthermore, since the natural ecologic role of wildfire in the Rockies has become better understood (Habeck & Mutch, 1973; Arno, 1980; 1983; Habeck, 1995), some recent T disturbed plant community types (Arno et 1., 1985), as well as the multiplicity of seral vege- ee types created by human activities. Suc- cessful fire suppression during a century of graz- ing and crop production has significantly altered the natural plant cover in the NRM. Fire-de- pendent vegetation types in the Rockies, espe- cially, have changed drastically in the past cen- tury (Gruell, 1983). Weaver & Dale (1974), Pfister et al. (1977), Arno (1979), Weaver (1979, 1980), Johnson & Pfister (1981, 1982), Peet (1981, 1988), Steele et al. (1981, 1983), and Arno & Hammerly (1984) contributed recent summaries of the climatic and physiographic features of the Rocky Mountains and detailed descriptive interpretations of the existing vegetation covering this region. These publications serve as my primary references; they are based on extensive field studies and in-depth HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 809 literature reviews covering eastern Washington and northern Idaho, northeastern Oregon, west- ern and central Montana, central and eastern Idaho, Wyoming, the Yellowstone Park area, and parts of Utah and Colorado. rno & Hammerly (1984) described and in- terpreted the lower (warm) timberline and upper mountainous regions of western No with supplementary comments on vegetation types adjacent to or between timberlines. John- son & Pfister's studies (1981, 1982) centered on identification and description of naturally oc- curring geologic and ecologic units in the middle and northern Rocky Mountains. Smith (1985) made an effort to provide an ecophysiological explanation for the distribution patterns within western montane forests. Var- ious factors—including moisture, solar radia- tion, and temperature — exhibiting seasonal vari- ations within the western plex and interactive with soil structure and nutrient con- tent, form the environmental complex that de- fines plant community distributions and succes- sional patterns. Smith emphasized the importance of “biophysical coupling" of site factors, fire, and various disturbances in dictating successional mountain needed before more com vegetation patterns can be made (Peet, 1988). Future studies should also emphasize the long- term importance of human disturbances. THE ROCKY MOUNTAIN SETTING: PHYTOGEOGRAPHIC SUBREGIONS The region typically designated as the northern Rocky Mountains extends from the Snake River Plain in southern Idaho to the international bor- der, joining with the southern Soe ee Rockies in British Columbia and Alberta. This region averages 500 km in width, Denice from near Yellowstone Park (northwestern Wyoming) westward to Hell’s Canyon on the Ida ho Olan state lines. Northward, the principal region ex- tends from central Montana westward to north- eastern Washington. Peet's (1988) description and interpretation of the forest vegetation of the entire Rocky Moun- tain system recognized the existence of four flo- ristic regions: far-northern, northern, southern, and madrean. My review encompasses all of the northern and part of the southern regions. The latter closely coincides with Daubenmire’s (1943) 810 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 YP »\® aa ae C (25) s wo FIGURE 4. Northern Rocky Mountains with oe P regions (after Arno & Hammerly, 1984; see text): A—Southern Continental Ranges; B— Northern Continental Ranges; C — Intermountain Ran ges; D— Inland Maritime Ranges; E—Southern Canadian deris Mountains; F — Middle Rocky Mountains. The Con- tinental Divide is shown. as a dashed line. States, provinces, Ves national parks, m features are shown by number and letter symbols keyed as follows: 1 — Canadian “Great Divide" (BC, ALB); 2 — Purcell Mountains (BC, MT); 3—Selkirk Mountains (BC, I MU gie inicium Mountains (BC); 5 — Whitefish Í A, WYO, ; 8—Swan Range (MT); 9—Clearwater Mountains (IDA); 10— Bitterroot Mountains (MT, IDA); 11 — Sapphire Mountains (MT); 12— Anaconda-Pintler Range (MT); 13— Little Belt and Big Belt Mountains (MT); 14— a dig Range (MT); 15— Madison Range (MT); 16— Blue Mountains (ORE); 17— Wallowa Mountains (ORE); 18— Idaho Batholith a : S ; d Teton Park (WYO); 27— Wind River Range (WYO); 28— Salt River & Wyoming Ranges ); 29— Wyoming Basin & Great Divide Basin (WYO); 30— Wasatch Mountains (UTAH); 31 — Uinta Mountains (UTAH); 32— 1987] HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 811 NIVAL ZONE Glaciers and S owfie Rock Outcrops & jode m ALPINE ZONE Alpine Tundra; Meadow ens (Cold) relifiends, ret bgt ps des ges E edge Communi s (Treeless) 2700m « UPPER TIMBERLINE Ipine 1 ; Whitebark pine - Pinus albicaulis Mountain hemlock - Tsuga mertenstana - IM b Lodgepole pine (seral) n d Pinus contorta SUBALP INE ZONE annii (Cool, Molst) Limber pine - Pinus flexilt 1800-2700m Bougies tir - Pseudotsuga menziesii Western larch (seral) ah Lariz oc and iud - Abies grandis - ctidentalis MONTANE ZONE ee redcedar - Thuja pl Mice hemlock - Tsuga heterophylla - IM Western yew - Taxus brevifolia - (Temperate-Mesophytic) are apen - Populus tremulo aida vestern seine. = IM P : r blrch - Betula papyrifen ($05 BO inus monticola (seral) Ponderosa pine - Pinus pondero + varieties) DRY CONIFER FOREST Limber pine - Pinus flexilis oe Hel vpe) reri pele Juniper - Juniperus WOODLAND ZONE Bitterbrush - Purshia tridenta ta « Fidi poids - Cen Ledifoli LOWER TIMBERLINE (Temperate-Xerophytic) i iia lanl iia 800-1500m _ Bluebunch wheatgrass - Agropyron spic SEMI-ARID STEPPE. AND di^ e - Festuca s and ae pene GRASSLAND ZONE Needlegrass - Sttpa coma Big sagebrush - Artemisia VT REN (+ varieties) (Warm, Dry) 1000m RIPARIAN/FLOODPLAIN Black cottonwood - Populus Dee (* other specles) QuakIng aspen - Populus tremuloides Ist) pe ied cae - Betula oaotdentatiai Paper birch - B. papyrifera (Warm, Molst Alde Willow specie Alnus spp & Salix spp FiGURE 5. Vegetation zones in the northern Rocky Mountains, typical of Montana, northern Idaho, and adjacent parts of British Columbia and Alberta. Approximate elevational limits of these zones are given, as are average positions of the lower and upper timberlines. IM = restricted to Inland Maritime region "central Rockies," and in my review will be re- as the Middle Rocky Mountains (MRM) is lo- ferred to as the “middle Rockies,” after Arno & cated south of the Yellowstone River in Montana Hammerly (1984). Figure 4 shows the geographic and extends southward through Wyoming to parts units employed in this review of Utah and Colorado; it includes the Uinta and The Southern Canadian Rocky Mountains Wasatch mountains, Medicine Bow Mountains, (SCRM), which extend from the international and the Colorado Front Range (Fig. 4). Both the boundary northward, are closely related physio- SCRM and MRM will be discussed, but empha- graphically and floristically to the NRM in the sis will be placed on the traditional NRM prov- United States, as defined above (Rowe, 1959; ince. Arno & Hammerly, 1984). The region designated The importance of elevation and topographic- < Medicine Bow Range (WYO, CO); 33—Park Range (CO); 34—Snake ua Plain (IDA); 35 — Steens Mountains (ORE); 36— Northeastern Great Basin (NEV); 37 — Great Basin (UTAH); 38— Juniper-Pinyon Woodland (UTAH); 39— Wyoming Basin Sagebrush- oe (WYO); 40— Plains Grasslands on 4] — Palouse Prairie (WASH, IDA); 42—Sagebrush Steppe (WASH). N-BC— Nelson, British Columbia; K-BC— Kimberley, British Columbia; J-ALB— R-IDA— Riggins, Idaho; S-IDA —Salmon, Idaho; B-IDA — Boise, Idaho; IF-IDA — Idaho Falls, Idaho; P-IDA— Pocatello, Idaho; TF-IDA — Twin Falls, Idaho; GP-MT —Glacier National Park, Montana; K-MT — Kalispell, Montana; M-MT-— Missoula, Montana; G-MT—Great Falls, Montana; H-MT—Helena, Montana; B-MT— (west) — Butte, Montana; B-MT (east)— Billings, Montana; D-MT —Dillon, Montana; YP- WYO- Yellowstone National Park, Wyoming; C-WYO (north)— Cody, Wyoming; C-WYO (south)— Casper, Wyoming; T-WYO— Thermopolis, Wyoming; RS-WYO-— Rock E Wyoming; R-WYO— Rawlins, Wyoming; L-WYO-— Lara- mie, Wyoming; E-NEV —Elko, Nevada; O-UTAH —Ogden, Utah; SLC-UTAH - Salt Lake City, Utah; V-UTAH — Vernal, Utah; C-CO — Craig, Colorado; en Rocky Mountain National Park, Colorado; B-CO— Boulder, Colorado. 812 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 NIVAL ZONE Ice, Snow, Rocks > 4200m Sedge/Gra ALPINE ZONE (Cold) lmann spruce - SUBALP INE ristlecone pine (Cool/Mesic) Wnitebark pine - Pinus albicaulis (N) - Pinus arista s Meadows, Fellfields, De A Tussock, Dl a cla Fields) edge marshes > 3500m Q@urrer TIMBERLINE a engelmannii bien Lice saya a (s) 2800m(N)/3500m(S) rosa pine MONTANE FOREST ZONE (Warm, Mesic) - Pinus ponderosa Blue spruce - Picea pungena : (E E) Open Parklands w) t eei bal te Moist sites/Canyo 22 pene Sonar cles oak - nipers - FOOTHILLS/STEPPE- Pinyon SHRUBLAND ZONE (Warm, Dry) “urus gambelii unipe rus osteosperma, communis, pines - Pinus edulis (E); g untain Maho ig - age iai us ledt foLiu tus; ea Greasewood - Sar «^n J. monosperma, J. TIMBERLINE J. horizontalis, J. scopulorum Pinus monophylla (w) tum: Lower slopes/Canyons - Artemista greeny Shadscale - Atriplex cobatus vermiculatus virginiana; Sagebr 1600m-2100m Grama/Buffalo gra PLAINS/GRASSLAND ZONE (Hot, Dry) grass - Bluebunch wheatgrass - Agropyron pis uM Bluegrasses - Poa species «1600 - Bouteloua i dactyloides Cottonwoods - Popu Bi rches RIPARIAN/FLOODPLAIN ZONE (Warm, Moist) ulus species; Mill - Betula species; Alders - Aad i spec - Saliz species cles FiGuRE 6. Vegetation zones typical of the middle Rocky Mountains. Major dominants in each life zone are iste moisture gradients in determining Rocky Moun- tain forest distribution patterns was emphasized by Peet (1988). He provided a series of “gradient mosaic diagrams” which portray the latitudinal shifts in forest composition along the entire length of the Rocky Mountains. Soil constitutes a third environmental variable important in interpret- ing Rocky Mountain vegetation patterns (Weav- er, 1978; Peet, 1 Figures 5 and 6 show generalized vegetation zonations in the northern and middle Rockies. Daubenmire (1980) also documented that NRM vegetation closely follows environmental gradi- ents from warm-and-dry to cold-and-wet, re- gardless of altitude. He stated that elevation above sea level is of reduced significance in the NRM because of the interdigitation of vegetation types within the mountain topography. Within the NRM, there are significant climatic gradients which play major roles in modern vege- tation distribution patterns. In the northwestern sector of this region (which includ theast Washington, northern Idaho, and northwestern Montana to the west slope of the Continental Divide in Glacier Park), there exists a moist in- land-maritime zone that features a well-devel- oped oceanic influence (Kirkwood, 1922; Larsen, 1930; Daubenmire, 1943; McMinn, 1952; Weaver, 1979, 1980; Lassoie et al., 1985; Peet, 1988). Here the NRM exhibits nearly continuous forest cover composed of several conifers with Pacific coastal affinities. These include Thuja pli- cata, Tsuga heterophylla, T. mertensiana. Taxus brevifolia, Abies grandis, and Pinus monticola, as well as coastal shrubs, herbs, and nonvascular plants (McCune, 1984). The lower slopes and valleys are also heavily forested, without a well- defined lower timberline. As the maritime influence diminishes, much of the remaining NRM region experiences a sig- nificantly colder and drier continental climate. The mountain peaks of this region are also much higher than those of the inland maritime region, vegetation types are caused by the much warmer and drier continental conditions. Here Pseudotsuga men- ziesii var. glauca, Pinus ponderosa var. ponder- 1987] osa, P. ponderosa var. scopulorum, Picea engel- mannii (and hybrids between P. enge/mannii and P. glauca), Abies lasiocarpa, Pinus contorta var. latifolia, P. flexilis, and P. albicaulis are the char- acteristic tree species. rno & Hammerly (1984) established several NRM subregions based on geographic and bio- climatic criteria. Each of these includes several mountain ranges (Fig. 4): A. Southern Continen- tal ranges, B. Northern Continental ranges, C. Intermountain ranges, and D. Inland Maritime ranges. The SCRM and MRM ranges were dis- cussed separately. In their reviews, Johnson & Pfister (1981, 1982) made use of ‘ ‘physiographic provinces," as well as forest “climax series" sim- ilar to those used in the habitat type classifica- tions cited above A very comprehensive biophysical land clas- sification for the southern Canadian Rockies, in- cluding an in-depth treatment of the be a is provided by Corns & Achuff (1982). T study focused on Banff and Jasper National en but has general usefulness for the SCRM. They established a series of physiognomic classes: A. Closed Forest, B. Open Forest, C. Shrub, D. Low Shrub-Herb, and E. Herb Dwarf-Shrub. Corns & Achuff used these vegetation types, combined with climatic and soil factors, to define “‘ecore- gions” in Banff and Jasper National Parks. VEGETATION OF THE NORTHERN Rocky MOUNTAINS The following description of the present-day NRM vegetation will make use of the geograph- ic-biophysical subregions established by Arno & Hammerly (1984). The information will empha- size the forested zones as depicted in Figure 5, with supplementary material on associated non- forest types. SOUTHERN CONTINENTAL MOUNTAINS The southern continental mountains include those found in eastcentral Idaho and southwest- ern Montana (Fig. 4, Region A). The Sawtooth Mountains, Lost River Range, Lemhi Range, Beaverhead Range, Anaconda-Pintler Range, Gravelly Range, and Madison Range are includ- ed. This region is dry and cool; the low moisture (20-40 cm annually) is due to a rainshadow effect caused by Oregon's Blue and Wallowa moun- tains and other Idaho ranges. Steppe and grass- land types are present on lower slopes and val- HABECK — NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 813 leys, dominated by several species of Artemisia and bunchgrasses (mostly Agropyron spicatum and Festuca idahoensis). The forest cover is con- fined to a relatively narrow zone between the lower timberline at 2,000 m and the upper tim- berline at about 2,700 m Pinus ponderosa is not well represented in the lowest forested zones of this part of the NRM because of the cold climate and short growing season at lower timberline (Arno & Hammerly, 1984). Instead, Pseudotsuga menziesii is the more common dominant tree adjacent to the steppe zone, sometimes mixing with Pinus flexilis at or near the lower timberline (1,800-2,300 m). In dry mountain ranges (Lost River Range), krummholz forms of Pinus flexilis and Picea en- gelmannii define the upper free line at ^ diis m, while on r tooth Range, Beavethead Range, and Madison Range) Abies lasiocarpa and Pinus albicaulis join Picea (Arno & Hammerly, 1984). Some southern continental mountain ranges have such severe moisture shortages that sagebrush-grassland vegetation extends up to and through the sub- alpine zone. Forest understories in the southern continental region are low in species richness. Semiarid steppe and grasslands dominated by Festuca idahoensis, Agropyron spicatum, Stipa comata, and Arte- misia tridentata form a mosaic among the Pseu- dotsuga menziesii forests. At higher elevations, Vaccinium globulare, Arnica cordifolia, sonia betulifolia, and Acer glabrum are common understory associates. Some of these shrubs and herbs extend into the subalpine zone where they are joined by Vaccinium scoparium, Linnaea bo- realis, Smilacina stellata, and Arnica latifolia. According to Dunwiddie (1977), Young & Ev- ans (1981), Arno & Gruell (1983), Arno & Ham- merly (1984), and Butler (1986), fire suppression since 1900, climatic changes, and livestock graz- ing have contributed to conifer invasions into the steppe-grassland zones at lower and middle elevations throughout the Rocky Mountains. Be- fore 1900, fires swept through these mountain ranges at intervals of 20-30 years, killing many young conifers that were establishing themselves in grasslands and shrublands. Intensive and ex- tensive grazing removes or reduces competing grasses, further facilitating spatial shifts in the forest-grassland boundaries. The southern continental region also supports 814 woodlands composed mostly of Juniperus scopu- orum. This vegetation type is found on dry, rocky sites, including outcroppings, generally below the limits of Pseudotsuga menziesii. A common as- sociate of juniper is Cercocarpus ledifolius. Pres- ent but much less common is Juniperus osteo- sperma, which occupies very severe sites and may be found mixed with Pinus flexilis. Sagebrush- steppe, dominated by Agropyron spicatum, Ar- temisia tridentata, Festuca idahoensis, and Pur- shia tridentata, occurs throughout Region A in Idaho and Montana (Kuchler, 1964). NORTHERN CONTINENTAL MOUNTAINS The northern continental mountains consist of the central Montana mountain ranges east of the Continental Divide (including the Lewis Range and “Rocky Mountain Front”), from the vicinity of Helena and Harlowton (Big Belt Mountains, Crazy Mountains, and Little Belt Mountains) northward to the southern Canadian Rockies in Alberta (Fig. 4, Region B). The isolated Bear Paw Mountains and Sweetgrass Hills in northcentral Montana and the Cypress Hills in southern Al- berta can be included in this geographic subunit, although they are located 150 km or more east of the cordilleran Rocky Mountains. The valley elevations here are lower than in the southern continental mountains but the northern continental ranges experience some- what less moisture stress (Arno & Hammerly, 1984). This subregion receives 25-30 cm of pre- cipitation annually, causing the lower timberline zone to be expressed at 1,200-1,500 m; contin- uous forest cover occupies a broader elevational zone. Pinus ponderosa forms the lower timber- line in parts of central Montana ranges (Arno, 1979) but plays a lesser role north of 47°. Pacific coastal understory species (see Inland Maritime Region) are uncommon in central Montana for- ests but Great Plains species are present, includ- ing Bouteloua gracilis, B. curtipendula, Andro- pogon spp., Opuntia fragilis, and Yucca glauca. Pinus ponderosa is replaced by combinations of Populus tremuloides, Pseudotsuga menziesii, Pinus flexilis, and Pinus contorta north of Great Falls, Montana. Cold winter climates with fluc- tuating temperatures and 115 kph winds seem to exceed the tolerance of Pinus ponderosa. Pop- ulus tremuloides and associated conifers that oc- cupy this area are often deformed and/or stunt- ed by the cold winter winds. Good examples of these effects can be seen on the east side of Gla- cier National Park. With the reduction of fire, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Pinus flexilis woodlands have extended into the prairie communities in this same region. The montane zone, occupying the middle el- evations of these central Montana mountains (1,500-1,800 m), is dominated by Pseudotsuga menziesii and Pinus contorta in varying amounts. Picea engelmannii is a dominant in areas tran- sitional to the subalpine zone, where it is codom- inant with Abies lasiocarpa and Pinus albicaulis. The upper timberline (tree limit) occurs between 2,300-2,600 m, with some terrain above 2,400 m supporting alpine tundra. Associated undergrowth species near the lower timberline in the northern continental moun- tains include Agropyron spicatum, Festuca ida- 2 F. oe Bouteloua gracilis, and Sti- a richardsonii. Symphoricarpos albus, Spiraea pins pt stellata, Linnaea borealis, Galium triflorum, Menziesia ferruginea, Vaccin- ium globulare, and V. scoparium dominate the understories of higher forest zones The disjunct Sweetgrass Hills (2,100 m) in northcentral Montana, although completely sur- rounded by semiarid Great Plains grasslands and foothills prairie, support forests composed of many of the montane and subalpine conifer species listed above, as well as the hybrid spruce Picea glauca x engelmannii (Habeck & Weaver, 1969). According to Thompson & Kuijt (1976), the Sweetgrass Hills were surrounded but not covered by the Laurentide glaciers; the hill- top nunataks may have supported tundra vege- tation. Later, the Hills were part of a more con- tinuously forested region in early post-Wisconsin, but climatic changes isolated the forests from the Rockies. The post-Hypsithermal shift towards increased warmth and dryness appears to dis- favor maintenance of the present-day subalpine understory species in the Sweetgrass Hills. INTERMOUNTAIN RANGES The Intermountain Ranges are found in north- eastern Oregon, central Idaho, and westcentral Montana (Fig. 4, Region C). The Blue and Wal- owa mountains are in Oregon; the Clearwater Mountains, Salmon River Mountains, and the western edge of the Bitterroot Mountains are in Idaho; the Sapphire Range, Anaconda-Pintler Range (border between Regions A and C), Flint Creek Range, and the southern parts of the Mis- sion and Swan ranges are located in westcentral Montana. The climate is partially influenced by moist maritime air masses that pass nearby, but this region receives less total moisture than does HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 815 FIGURE 7. the more western Inland Maritime region. On the other hand, the more severe continental cli- mate that features temperature extremes, cold dry winter, and stressful chinook winds is not as common in this region (Arno & Hammerly, 1984). The Intermountain Ranges have well-defined lower and upper timberlines, and the valley bot- toms (900-1,050 m) and southern aspects sup- port bunchgrass prairies (Agropyron and Festuca shrub types were originally interfacing or inter- spersed with Pinus ponderosa, forming the low- est forest zone. In presettlement times, the exact position of the lower timberline was partially conditioned by wildfires caused by lightning and aboriginal man (Gruell, 1983). Frequent low intensity ground fires perpetu- ated open Pinus ponderosa savannas (Fig. 7), even Pinus ponderosa forest in Intermountain region, western Montana. Located near Darby, Montana, at 1,050 m within the Pseudotsuga menziesii zone. This forest type historically was maintained as open savannas by ground fires. USDA Forest Servic on mesic sites where Pseudotsuga menziesii and Abies g Mutch, 1973; Thak. 1976, 1985; Barrett & Arno, 1982; Freedman & Habeck, 1984). My own analysis of fire scars on 300- to 350-year-old Pinus ponder- osa specimens occupying a drainage near Mis- d 1700s, fro 1750 to 1870 the fire frequency doubled! That is, intervals between fires were reduced to an average of five years. Indians practiced routine burning throughout the Rocky Mountains before their encounter with Euroamerican influences (Barrett & Arno, 1982; Gruell, 1985; Arno, 1985; Lewis, 1985) but may have started more fires following introduction of horses into western Montana in the 1730s. Such frequent burning maintained even moist Pseudotsuga forest sites as grassland savannas. Remnants of the savanna bunchgrass, Festuca scabrella, still exist beneath the Douglas-fir forest canopies that have devel- oped without fire since 1900. Above the Pinus ponderosa zone lies the Pseu- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 < p ae E- ` Ses oar Old- ae Larix occidentalis-dominated forest in Inland Maritime region in Flathead National 00 m). Established M a wildfire, the Larix 1s being replaced by the climax species, For a (1,1 Palas menziesii. USDA Forest Servic 1987] > "g^ " gn gef ^ t Yy t w E9. Subalpine zone exhibiting unev HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 817 -distributed Abies lasiocarpa. Pinus albicaulis and P. contorta near ‘Wahoo Pass (2,000 m) in Susa Bitter Wilderness, Montana/Idaho state line. USDA Forest Service. dotsuga menziesii forest zone, extending from 1,200 to 1,800 m. Within this zone, Larix oc- cidentalis prevails as a major pioneer species (Fig. 8). This imi dependent species may achieve diameters of 1—1.5 m (thick basal bark layers that sauer fire protection contribute to large girths) and heights over 40 m. Another common but short-lived associate in the upper part of the Pseudotsuga menziesii zone is Pinus contorta; it is also dependent in the NRM on periodic fires or other site disturbances (Lotan & Perry, 1983; Muir & Lotan, 198 Common understory species in the Douglas- fir/western larch forests include Physocarpus malvaceus, Acer glabrum, Vaccinium globulare, V. caespitosum, Calamagrostis rubescens, Carex geyeri, Spiraea betulifolia, Arctostaphylos uva- rsi, Mahonia repens, Linnaea borealis, and Symphoricarpos albus. t from 1,800 m to treeline near 2,700 m (Fig. 9). Pinus con- torta extends into this zone and mixes with Pinus albicaulis and Picea engelmannii. Again, fire his- torically occurred in these higher forest zones (Fig. 10) but at long intervals, usually more than 100 years and sometimes at 200- to 300-year intervals (Habeck, 1985) Dominant understory species associated with Intermountain subalpine forests include Men- ziesia ferruginea, Vaccinium globulare, V. sco- parium, Xerophyllum tenax, Carex geyeri, Ar- nica cordifolia, A. latifolia, Calamagrostis rubescens, C. canadensis, Luzula hitchcockii, Clintonia uniflora, Tiarella trifoliata, Anemone piperi, and Linnaea borealis. Insects and other pathogens are known to have caused periodic ecosystem disruptions in the In- termountain forests. McCune (1983), however, has suggested that man’s reduction of wildfires Montana has predisposed the forests 1982; Anderson, 1985). Their data suggest that ANNALS OF THE MISSOURI BOTANICAL GARDEN x ee aR ; he $ | uy. Pn ^ A. ^. Wed “`` suce à UNS p A EE AR pote Es. Sent wd. ` - stel. 2 LE TL ju det S — * % E NE K (Aes xt Y os i Z a af xx à *j de ee uc M ox, Mors MEL ge vere NN UM M. à * n « = ] 4 Melo FIGUR fire suppression and logging practices in the Pseu- dotsuga menziesii forests in western Montana create conditions that encourage outbreaks of western spruce budworm (Choristoneura occi- dentalis) of greater severity and longevity than those experienced prior to fire control. Without fire, forest cover has become more structurally te, "Tr A "E el y ` 4 - 5 - = « A rdi ° Be n 224. " y " Doi 3 [Vor. 74 on €" URE 10. Fire-generated, even-aged Pinus contorta in the subalpine zone (2,500 m) in northern Rocky Mountains region. USDA Forest Service. uniform, which facilitates insect spread over large reas. The upper timberline in the Intermountain Ranges forms at about 2,700 m and is dominated by Pinus albicaulis, Abies lasiocarpa, Picea en- gelmannii and, not infrequently, Larix lyallii. Each of these exhibits a wind timber develop- 1987] HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION Ta Ze. FIGURE 11. USDA Forest Service. ment (dwarfed in size and shape) as each reaches its individual upper limit of growth. Larix lyallii is confined to relatively harsh north slopes over 2,100 m and frequently occupies talus slides and boulder fields. Only occasionally does Larix lyal- lii grow on level sites with developed soil (Arno & Habeck, 1972). Near timberline, Pinus albicaulis, Picea en- gelmannii, and Larix lyallii form open-grown, parklike stands. Topography, wind, and snow deposition paths cause a variety of interesting spatial arrangements in the timberline zone, such s "ribbon-forests" and krummholz mats (Bil- lings, 1969; Fig. 11). Some individual timberline specimens of Pinus albicaulis, Larix lyallii, and Picea engelmannii are relatively large (1.0-1. m in diameter) and old (300 to 500 years). PAyl- lodoce empetriformis, P. glanduliflora, Cassiope mertensiana, Vaccinium scoparium, Xerophyl- lum tenax, and various sedges (Carex spp.) and rushes (Luzula spp.) are the understory domi- nants in these timberline forests. Some bunchgrass communities (Agropyron Wind-shaped Pinus albicaulis within upper ti zone (2,500 m) in southcentral Montana. spicatum, Festuca scabrella, and F. idahoensis), compositionally similar to lower timberline grasslands, exist islandlike within the Inter- mountain subalpine forest zone well above 1,800 m. They are believed to be the results of hyp- sithermal displacements (Daubenmire, 1968, 1975, 1981; Root & Habeck, 1972) and are pres- ently maintained by localized xeric conditions induced by slope, aspect, shallow soils, or un- weathered parent rock substrates. Although sub- ject to burning, there is no evidence that fire is required to perpetuate them. Other grasslands, or *grass balds," featuring Festuca viridula com- munities may dominate the subalpine zone in central Idaho and northeastern Oregon, extend- ing to over 2,40 Detailed donodan and classifications of the lower zone Intermountain grasslands (Fig. 12) have been presented by Mueggler & Stewart (1980). Some of their 30 grassland and shrubland habitat types are similar to those documented by Daubenmire (1970) but others are distinctive for the east front of the Montana Rockies. Major 820 E 12. Bun ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 PRES A (Noe i nchgrass prairie community on the Deerlodge National Forest (1, 750 m), Montana, Inter- DA ou ina EE Pseudotsuga menziesii with groves of Populus tremuloides occupying lower elevations. USD Forest Servic Intermountain grassland dominants include Stipa comata, Agropyron spicatum, Festuca ida- hoensis, F. scabrella, Deschampsia caespitosum, Artemisia tridentata, A. arbuscula, Potentilla fru- ticosa, Purshia tridentata, Cercocarpus ledifolius, Rhus trilobata, and Sarcobatus vermiculatus. Other NRM grassland studies have focused on local expressions of prairies on sites surrounded by continuous forests. In many instances, dis- junct grasslands are the result of local mountain rainshadows, but soil texture (e.g., coarse glacial till) and past fires have also favored grassland development (Blinn & Habeck, 1967; Koterba & Habeck, 1971; Root & Habeck, 1972). Wright & Wright (1948) have described the grasslands in southcentral Montana; their five types are list- ed by community dominants (from most me- sophytic to more xerophytic): a) Festuca ida- hoensis, b) Agropyron spicatum, c) Agropyron spicatum/Carex filifolia/Bouteloua gracilis, Bouteloua gracilis/Stipa comata/Koeleria cris- tata, and e) Bouteloua gracilis/Stipa comata. e. — Representative shrubland types in the Inter- sen region € 13) were described by Mueggler & Stewart (1980) and by Kuchler pica "C ercocarpus ledifolius, Purshia triden- tata, Symphoricarpos oreophilus, and Artemisia tridentata all form shrub-bunchgrass commu- nities below lower timberline, occupying dry rocky sites or river floodplains, although these shrubland communities may also be found in- terspersed within the timbered zones. The grass- es are the same as those that form the prairie vegetation types in the valleys and lower slopes, that is, Agropyron spicatum, Festuca idahoensis, Stipa comata, and Poa sandbergii. INLAND MARITIME REGION The Inland Maritime region spans northeast- n Idaho, and north- (Fig. 4, Region D). The Selkirk Mountains of northern Idaho and adjacent parts of Washington and British Columbia, the north- 1987] HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 821 FIGURE 13. ern Bitterroot Range, the Cabinet Mountains, the Whitefish Range, the northern Swan Range, and the west slope of the Continental Divide (in Gla- cier Park) collectively comprise this region. s its name suggests, this region is relatively stations in localized rainshadows may record 40 cm or less. Warm, dry weather usually prevails during July and August. Grassland and Pinus ponderosa communities, which exhibit drought tolerance (Minore, 1979; Lassoie et al., 1985), occur locally in some of the driest rainshadow areas. The reduced summer moisture sets the stage for wildfires. Historic fires are believed to have perpetuated xeric vegetation types in cli- matically moist parts of this geographic subunit (Habeck & Mutch, 1973). Typically, the forest zones in the Inland Mar- itime region contain combinations of the follow- Sagebrush shrubland and rangeland vegetation types in upper Big Hole drainage (2,000 m), in southern continental ranges, Beaverhead National Forest, Montana. Pseudotsuga menziesii forest in distance. USDA Forest Service. ing dominants, ranked from highest to lowest in drought resistance (Minore, 1979): Pinus pon- derosa, P. contorta, Pseudotsuga menziesii, Picea engelmannii, Abies grandis, Larix occidentalis, Abies lasiocarpa, Thuja plicata, Taxus brevifolia, Pinus monticola, Tsuga heterophylla, and Tsuga mertensiana (Figs. 14, 15). Physiological adaptive features of these coastal conifers were reviewed in detail by Lassoie et al. (1985). The mesophytic forests dominated by Thuja and Tsuga resemble in many ways the forests found on the western slopes of the Coastal nd Cascade mountain ranges in Washington and British e E (Daubenmire & Daubenmire, 1968; Habeck, 1978; Williams & Lillybridge, 1983; Cooper et al., 1988) With increased elevation, even greater amounts of precipitation occur, often reaching and ex- ceeding 200 cm annually. A large portion (75- 8596) of this precipitation falls as snow between September and March. Pseudotsuga menziesii, £ Abies lasiocarpa, and Picea engelmannii occur ANNALS OF THE MISSOURI BOTANICAL GARDEN rae — T mtm — JP ` a wem d LI $ — d 1^ d FIGURE 14. Mature forest in the Inland Maritime region; Deception Creek, Coeur d’Alene National Forest, Idaho (1,000 m). Tsuga heterophylla, Abies grandis, and Pinus monticola dominate. Note fire-scarred snag on left. USDA Forest Service. in abundance on the higher inland Maritime mountain slopes. Pinus albicaulis is locally abun- dant on warm aspects that may experience some July-August moisture shortages. Between 1,500 and 1,800 m, near the Idaho- Montana state line, Tsuga mertensiana joins Abies lasiocarpa in forming the highest closed-canopy forest type (Habeck, 1967). Tsuga mertensiana is very intolerant of summer drought and heat (Minore, 1979) and severe continental winters, but survives very well in parts of the Inland Mar- itime Region where suitably mild conditions pre- vail. Tsuga mertensiana is an abundant timber- linespeciesin the Coastal and Cascade mountains of western Oregon, western Washington, British Columbia, and southeastern; Alaska (Arno & Hammerly, 1984). The Inland Maritime region's timberline, formed at 2,000-2,300 m (Fig. 16), features krummholz of Abies lasiocarpa and Pi- nus albicaulis; erect or partly wind-shaped Larix lyallii is also present at times (Habeck, 1969; Arno & Habeck, 1972). On glacially scoured parts of the Selkirk Mountains in northern Idaho, Tsu- ga mertensiana and Larix lyallii are rare or ab- sent. Each of the Inland Maritime trees has an in- dividualistic distribution. They can be ordered from most restricted to most widely distributed within this region: A/nus rubra, Tsuga merten- siana, T. heterophylla, Pinus monticola, Thuja plicata, Taxus brevifolia, and Abies grandis. It has been noted that the individuals or popula- tions occurring at the range limits of these mar- itime trees, such as Thuja plicata near Missoula, Montana, are confined to isolated higher (1,400 m) ravine sites (Habeck, 1978; McCune & Allen, 1985b). Summer moisture deficiencies encoun- tered in westcentral Montana may be counter- balanced by the cooler temperatures in these middle elevation ravine sites. The undergrowth in the mesic and wet-mesic Inland Maritime region forests is floristically 340636 PAG n aa, fing KUE m, x ym NS "€ P TSS . vum . 1 è e ` AS ~~ a v dI j r T ik n. N - * = À E-' M^ DA Jac. Y MN i N as ¿š : b di > š yy n x TF DEC ue $ , ; &. 4 ` xa) $ P m ku 22 88 ch š RUN I Vir pe VEN [LS zs e T z CE — — ee Ge, : pO -BwA "G E y HABECK — NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION E ° [^2] o z zr [^2] e S "n 3 g: o = S z 9 [2] el A E o wv ™ — = O = x g: © [^2] Q E © f = o = — g: © — wu O — & 9 S e wass & S = X = e - -— z FIGURE 15. Montana. USDA Forest Service. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 FIGURE 16. richer than in any of the other subunits. It in- cludes Clintonia uniflora, Adenocaulon bicolor, Disporum hookeri, Tiarella trifoliata, Cornus canadensis, Aralia nudicaulis, Rubus parviflorus, Athyrium filix -femina, G iymnocarpium dryopte- many ed species. dient exists, west to east, between S Washington (38 cm/yr., pine-grassland savanna) and nearby (40 km east) Coeur d'Alene, Idaho (75 cm/yr., cedar-hemlock-western white pine forests). Fires occur at lower frequencies (100-year in- tervals or longer) in this geographic region (Ha- beck, 1985). When wildfires do occur, however, they are often of much higher intensity (“stand “Snow ghost" forest in the subalpine zone of northern Idaho's Bald Mountain (2,300 m). Snow- caked trees are Abies lasiocarpa. USDA Forest Service replacement" fires), due to the greater organic matter (“fuel”) accumulations in these moist for- ests. When the right combination of midsummer "fire weather" occurs—characterized by high paige gusty afternoon winds, dry ground potential climax species to make an initial entry at the same time. Examples of short interval (less than 10 years) “double” and “triple” burns exist, and forest recovery on such sites may be retarded for many decades. Deep snow accumulations in the Inland Mar- itime mountains (Fig. 17) cause frequent ava- often dominated by such woody angiosperms as 1987] ; ` t N _ PL >= 22 HABECK — NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION FiGuRE 17. Snow cornice above Picea engelmannii and Abies lasiocarpa forest at 2,100 m in Panhandle National Forest, coal Idaho. USDA Forest Service. ] Alnus sinuata alnifolia, Sorbus spp., Acer glabrum, Populus trichocarpa, P. tremu- loides, Salix spp., and a rich assortment of pe- rennial herbs that seem well adapted to frequent avalanches. VEGETATION OF THE SOUTHERN CANADIAN ROCKIES The southern Canadian Rockies are composed of high, rugged terrain with a long history of glaciation. The mountain ranges often reach or exceed 2,400 m, with peaks over 3,000 m. The southern Canadian Rockies embrace several ma- jor mountain ranges near the United States/Can- g Frigid polar air masses characterize the con- tinental climate east of the Great Divide in Al- berta. The dessicating winter winds are similar to those described for the east front of the Mon- tana Rockies; these winds are responsible for “red belt” mortality among the conifer communities located here. Just west of the Great Divide, the climate is more like that of the Inland Maritime 826 region in northern Idaho and northwestern Mon- tana; mild and wet weather prevails as Pacific coastal storm tracks penetrate eastward (Arno & Hammerly, 1984). Southeastern British Columbia, parts of which lie in the Cascade Mountains rainshadow, has a warm and dry lower timberline in valley bottoms where only 25-40 cm of precipitation occurs an- nually. Open Pinus ponderosa woodlands, bunchgrass prairies (Agropyron spicatum and Festuca idahoensis), and sagebrush steppe (4r- temisia tridentata) form the major plant com- munities in the semiarid valleys (Tisdale & McLean, 1957; McLean, 1970; Tisdale, 1974). Some of the upper slopes of nearby mountains also have low annual precipitation, and at mid- elevation slopes a forest zone dominated by Pseudotsuga menziesii occurs. Above this is the subalpine zone dominated by Abies lasiocarpa, Picea engelmannii, and Pinus albicaulis, which ultimately form a krummholz on upper slopes above 2,100 m. Past fires in southeastern British Columbia perpetuated extensive stands of Pinus contorta at middle and upper elevations (LaRoi & Hnatiuk, 1980; Arno & Hammerly, 1984). Those parts of southeastern British Columbia that receive greater coastal moisture display for- est vegetation similar to that of the Inland Mar- itime region in the United States. Warm, dry, lower timberlines occur only rarely in this part of the Canadian Rockies, where dominants in- clude Thuja plicata, Tsuga heterophylla, Abies grandis, Pseudotsuga menziesii, Picea engelman- nii, and P. engelmannii x P. glauca hybrids. Pinus contorta, Abies lasiocarpa, Tsuga merten- siana, Pinus albicaulis, and Larix lyallii are the major timberline dominants in the Selkirk and Purcell mountains. Timberline in these ranges typically occurs at elevations below 2,100 m; however, it may occur below 1,500 m in areas of deep snow accumulations. The highest moun- tain peaks (over 3,000 m) support permanent snowfields and glaciers (Shaw, 1916) The Alberta Rockies consist of the mountains forming the Great Divide. From Waterton Park northward to Banff and Jasper National Parks, the Divide peaks reach from 2,800 m to nearly 3,650 m in height. The continental climate in Alberta disfavors the occurrence of tree species adapted to milder maritime conditions. Annual precipitation is near 150 cm at the higher ele- vations but less than 50 cm on the east front at Calgary. The subalpine and montane forest zones are dominated by the same species listed above ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 for British Columbia, except that maritime trees like Thuja and Tsuga are missing. At timberline, Abies lasiocarpa, Picea engelmannii, Larix lyal- lii, and Pinus albicaulis form the upper treeline communities, including the krummholz zone (Moss, 1955; Ogilvie, 1963, 1976; Day, 1972; Corns & Achuff, 1982). The lower forest are mixtures of Pseudotsuga menziesii, Picea glauca (and hybrids P. glauca x P. engelmannii), and Populus tremuloides. Local moisture-stressed sites support Pinus flexilis (Moss, 1944; Rowe, 1959; Stringer & LaRoi, 1970). Pinus ponderosa is absent, according to Arno & Hammerly (1984), seemingly unable to tolerate winter desiccation and rapid temperature changes. As was described for other areas of the NRM, frequent high in- tensity wildfires helped develop and perpetuate extensive stands of Pinus contorta throughout much of the Alberta Rockies, from the lower to the upper timberline (Tande, 1979). The natural grasslands in southern Alberta east of the Great Divide occur where precipitation becomes lower, temperatures are warmer, Chi- nook winds prevail, and evaporation rates are high (Moss, 1944; Moss & Campbell, 1947). Moss (1944) described three prairie community types: Bouteloua gracilis/Stipa comata, Agropyron/Sti- pa/Carex, and Festuca scabrella/F. idahoensis/ Danthonia intermedia/D. parryi. Much of the native prairie no longer exists because of modern levels of grazing and mowing. +h 1 A VEGETATION OF THE MIDDLE Rocky MOUNTAINS The last part of the Rocky Mountains to be treated here lies north of Provo and Salt Lake City, Utah, in the vicinity of 40°N, and includes the adjacent parts of northwestern Colorado, the Medicine Bow Mountains in southeastern Wy- oming, the remainder of Wyoming west and north of the Wyoming Basin desert, north to south- central Montana (Beartooth Plateau), and west to the Snake River Plain in southern Idaho (Fig. 4, Region F). This region coincides with the "Middle Rockies" as discussed by Oosting & Reed (1952), Johnson & Pfister (1982), Arno & Hammerly (1984), and Mutel & Emerick (1984), and includes the northeastern part of the Rockies that Reveal (1979) designated “the Intermoun- tain Region.” Figure 6 shows the typical vege- tation zones in this part of the Rocky Mountains. The Utah portions of the middle Rocky Moun- 1987] tains support a flora that contains many elements that are geographically centered further to the south and southwest, characteristic of the pin- yon-juniper, chaparral, and Great Basin desert vegetation types. In contrast, western Wyoming ranges are without these floristic inclusions and display more elements common to the East Front of the Rockies in central Montana (Arno & Ham- merly, 1984). The forest vegetation of this region is usually dependent on melting of deep winter snow packs (from Pacific storms) to supply need- ed summer moisture. The inland continental cli- mate lacks a maritime component of the sort described earlier (Baker, 1944). Arno & Hammerly (1984) described the mid- dle Rockies as an area of well-separated ranges diverse in geologic structure and rock types. Johnson & Pfister (1982) provided detailed in- terpretive summaries of the geology of this re- gion. Three geologic mountain types exist (see Fig. 4 for locations): 1) anticlinal mountains (Beartooth Plateau, Big Horn—Pryor Mountains, Uinta Mountains, and Wind River Range); 2) mountains of the overthrust belt (Tetons, Wy- oming, and Wasatch ranges); and 3) volcanic mountains (Yellowstone Plateau and Absaroka Range). The highest points in the middle Rocky Mountains reach up to approximately 4,200 m (Gannett Peak and Grand Teton), although more typical heights range between 3,000 and 3,600 m. The Wasatch and Uinta ranges have broken patches and stringers of timber on their upper mountain slopes, a result of frequent avalanches clearing wide tracks through the timber. Several tree species achieve dominance here, including Pseudotsuga menziesii, Pinus flexilis, Picea pun- gens, Abies concolor, and Populus tremuloides. Pinus contorta is present in the northern part of the Wasatch Range but Pinus ponderosa is scarce. The lower Wasatch slopes are covered with Quercus gambelii, Cercocarpus ledifolius, and Acer grandidentatum. This mountain shrub or chaparral type replaces the pinyon-juniper vege- tation zone found further westward in Nevada (Hayward, 1945, 1948; Arno & Hammerly, 1984). The Uinta Mountains, which have an east- per subalpine zones are stands of Picea engel- mannii, Abies lasiocarpa, and Pinus flexilis; this region is beyond the distribution of Pinus albi- caulis. Extending down to 2,700 m below these HABECK — NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 827 zones are Pinus contorta forests. A wide band of Populus tremuloides, serving as a major seral species in the middle Rockies, occurs below the Pinus contorta zone. Pinus ponderosa is scattered on south slopes and found more extensively on north ones at the lower Uinta timberline down to 2,100 m. Mid elevations support extensive stands of Pseudotsuga menziesii; moist drainages (riparian sites) contain Picea pungens commu- nities Arno & Hammerly (1984) and Ellison (1954) provided additional descriptions of the contrast- ing climatic and vegetational features between the Wasatch and nearby Uinta ranges. In contrast to the Wasatch Range, the Uintas lack the oak- maple chaparral but support a pinyon-juniper type. In both areas, the lowest and driest shrub zones are dominated by a dozen different species and varieties of sagebrush (Artemisia spp.) and Atriplex canescens, A. confertifolia, and Cerco- carpus ledifolius. Variations in summer rainfall, geologic and topographic features, and contract- ing patterns of past plant migrations have all contributed to these differences. The modern forest vegetation occurring in Gordo! s Front Range (Fig. 4, vicinity of Rocky Mountain National Park) has been the subject of many studies dating back 80 or more years. The most recent and comprehensive among these is the detailed treatment provided by Peet (1981), who reviewed muc the previous vegetation literature for this ad of the middle Rocky Mountains. Employing modified gradient anal- ysis techniques, he derived a classification of the Front Range vegetation that accommodated both developmental (successional) and mature (late seral/climax) community types, relating them to a moisture-elevational complex. Peet classified the Front Range vegetation (1981, his fig. 5) into community series as follows: A. Pinus ponderosa woodland series; B. Pinus ponderosa/Pseudotsu- ga menziesii forest series; C. Mesic montane for- est series (a heterogeneous group that includes Picea, Abies, Pseudotsuga, Populus, Betula, and alnus); D. Pinus contorta forest series; E. Picea/ Abies forest series; F. Pinus flexilis forest series; Alpine transition (krummholz) series. His classification also delimits community type sub- units within these series. Subsequently, Green- land et al. (1985) described a methodology for defining bioclimatic zones in the Colorado Front Range utilizing a one-dimensional version of Peet's vegetation ordination model The importance of 19th century anthropogenic 828 disturbances within the montane forests of the Front Range have been investigated by Veblen & Lorenz (1986). Patterns of forest recovery fol- lowing severe logging and burning disturbances inflicted during the Colorado g booms were studied. Their findings clarify the successional pathways that forest recovery has followed dur- ing the last century in this part of the Rocky Mountains. The upper-elevation forests in a representative watershed within the Medicine Bow Mountains in central Wyoming were discussed in detail by Romme & Knight (1981), who investigated and modeled the interactions of fire frequency and topographic position on vegetation dynamics in this part of the middle Rockies. Although much of the high country in the Medicine Bows has a potential for supporting climax forests dominat- ed by Picea engelmannii and Abies lasiocarpa, such expressions of spruce-fir forests are pri- marily confined at this time to moist sites such as ravines and valley bottoms. Past occurrences of short-interval (less than 100 years) wildfires on open slopes and ridgetops favored develop- ment and maintenance of Pinus contorta forests (Lotan & Perry, 1983). Even though fire suppres- sion has been in operation for more than a cen- tury, the slow rates of successional processes in these mountains have tended to impede the re- turn of spruce-fir forests. Romme & Knight (1981) included within their watershed study area a two-dimensional ordi- nation of all major plant community types oc- curring between 2,250 and 3,000 m on sites rang- ing from moist valley bottoms to south slope ridgetops. Salix, Alnus, and Populus mixtures occur in the lower, moist habitats; Pseudotsuga menziesii forests occur on the mesic, lower north slopes; Artemisia communities occur on most xeric foothill elevations. At intermediate eleva- tions of 2,500 to 2,700 m, combinations of Pseu- dotsuga menziesii, Pinus contorta, and Populus tremuloides prevail. Only moist sites above 2,700 m currently support mature spruce-fir forests. The mountains in western Wyoming (Wind River, Salt River, and Wyoming ranges, as shown nii and Abies lasiocarpa (Loope, 1976; Steele et al., 1983; Arno & Hammerly, 1984). A krummholz zone composed of these conifers is found at elevations near 3,050 m throughout these mountains. Much of the sub- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 alpine forest zone, however, is dominated by ex- pansive stands of Pinus contorta except where limestone substrate prevails. This pine, together with Pseudotsuga menziesii, often forms the up- per timberline forests at elevations above 2,700 m and then extends downward to the lower zones at or below 2,100 m (Moir, 1969; Arno & Ham- merly, 1984). Loope & Gruell (1973) have de- scribed the natural role of fire in these north- western Wyoming forests. Near the lower timberline, Populus tremu- loides groves comprise a broken belt separating Pinus contorta forests from the sagebrush com- munities in the lower valleys (Reed, 1971; Youngblood & Mueggler, 1981; Arno & Ham- merly, 1984; Fig. 18). Although Pseudotsuga menziesii can be found in the western Wyoming mountain ranges, it does not form a distinctive forest zone as it does elsewhere in the northern Rockies, such as in western Montana (Arno, 1979). This is due to the importance of Pinus contorta as a seral species in zones where Pseu- dotsuga menziesii is the potential climax domi- nant. Many ofthe subalpine conifers can be found at lower timberline, often along mountain stream courses (Arno & Hammerly, 1984). The valleys in the western Wyoming mountain ranges are much cooler than comparable sites elsewhere in the northern Rockies; for this reason, Populus tremuloides and mesophytic conifers occupy the lower timberline zone (Steele et al., 1983). Pin- yon-juniper and mountain chaparral are missing from the cold western Wyoming mountains (Arno & Hammerly, 1984). The Absaroka Range and adjacent Beartooth Plateau (Montana-Wyoming border; Fig. 4) dis- play an extensive alpine tundra zone with many ice fields at elevations over 3,000 m. Timberline parkland (open-canopied groves and scattered individuals) is also present, dominated by Pinus albicaulis, Picea engelmannii, and Abies lasio- carpa, similar to those in the Wind River and Teton ranges. The Big Horn Mountains, spatially separated from the other middle Rocky Mountain units, are located east of the Big Horn Basin in west- central Wyoming and extend into adjacent parts reaching about 60 cm in the subalpine forest zone at 2,700 m. Soils derived from granite, shale, 1987] HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 829 FiGuRE 18. Old growth Populus tremuloides forest near Scout Lake (3,100 m), Utah. Note conifer invasion in understory. USDA Forest Service. 830 limestone, and dolomite rocks play a major role in plant distribution patterns in many of the mid- dle and northern Rockies east of the Continental Divide, including the Big Horns. A series of vege- tation studies has been completed in the Big Horns (Hurd, 1961; Despain, 1973; Hoffman & Alexander, 1977). Big Horn Mountain Pinus contorta forests oc- re g the coarse and nutrient-poor granitic soils. Grasslands and shrublands gen- erally occupy soils from shale and limestone, ex- cept where forests are supported in areas where moisture stress is lessened due to slope aspect (Arno & Hammerly, 1984). Picea engelmannii and Abies lasiocarpa occur in the subalpine-tim- berline zone of the Big Horn Range, but Pinus albicaulis is absent from this area. Pinus contorta is dominant on granitic soils at elevations below the subalpine zone which extends from 2,700 m to 2,100 m. The lower timberline is formed by Pseudotsuga menziesii near 1,800 m. Juniper woodlands (Juniperus scopulorum and J. osteo- sperma) are closely associated with the Pseudo- tsuga menziesii zone on the west slopes; Pinus edulis is replaced by Pinus flexilis in this juniper community type. Pinus ponderosa var. scopu- lorum becomes important only in this part of the middle Rocky Mountains (east face of Wyoming Big Horns and northward into Montana) but does not display the large growth forms seen among the P. ponderosa var. ponderosa occurring fur- ther west. Geographically, P. ponderosa var. scopulorum occurs eastward into the Black Hills of South Dakota, to the exclusion of Rocky Mountain conifers (Johnson & Pfister, 1982). The middle Rockies support widespread sage- brush-grass; Tisdale & Hironaka (1981), Blais- dell et al. (1982), oe < al, (1983), and West (1983) provided in-dep t view of the major community ae in the nonforested areas of this region. Where the Wyoming Basin and Great Plains meet, there exist sod-forming grasslands dominated by Bouteloua gracilis and Agropyron smithii intermixed with Artemisia tri- dentata and other sagebrush taxa. These com- munities are the products of the continental cli- mate, with growth moisture arriving in early summer rainfall. Elsewhere in southern and cen- tral Idaho and southwestern Montana, bunch- grass prairies are present, dominated by Agro- pyron spicatum, Festuca idahoensis, Stipa comata, and Poa sandbergii. In these sagebrush- bunchgrass communities, moisture comes pri- marily in winter and spring, with droughty sum- mers. citec [^2] ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 The middle Rockies sagebrush-bunchgrass community types recognized by Hurd (1961), Beetle & Lud (1982), Johnson & Pfister (1982), and Hironaka et al. (1983) include the following, each named after the major dominant: a) Festuca idahoensis, b) Agropyron spicatum, c) Bouteloua gracilis, d) Purshia tridentata, e) Sym- phoricarpos oreophilus, f) Artemisia tridentata, g) A. tripartita, h) A. nova, i) A. arbuscula, j) A. longiloba, k) A. cana, 1) A. rigida, in, Atriplex canescens/A. confer. AUAA. nutta and n) arcobatus vermiculatus. Shrubland ane types in the same region include a) Quercus gam- belii, b) Acer grandidentatum, and c) Cercocar- pus ledifolius. Modern (post-1900) vegetation shifts in mid- dle Rocky Mountain 55) believed to be induced by fir (Sauer, 1950; Humphrey, 1962; Steele et al., : 1981, 1983), also involve extensive invasion by Junip- erus occidentalis into grassland and sagebrush communities. Tisdale & Hironaka (1981), point- ing to the sensitivity of Artemisia to fire, sug- gested that fires in presettlement times must have been infrequent (long interval) in some parts of the sagebrush region, such as drier Artemisia Barney & Frischnecht (1974) and Young & Evans (1981). Problems attending these changes were described in a lengthy juniper man- agement symposium (Martin et al., 1978). Through wood rat (Neotoma) midden and lake sediment analysis, Mehringer & Wigand (1986) documented that western juniper has had a long history of rapid geographic shifts in response to climate changes during the past 4,000 years. They stated, “. . . the spectacular and persistent ex- pansion of western juniper over the last hundred years — despite chaining, bulldozing, cutting, poi- soning and burning—i is not an unusual event nec- essarily req g explanations unique to the his- toric period. In fact, the rate and degree of change in the comings and goings of western juniper over ant communities historically used as sheep and cattle range throughout the middle Rockies have been severely altered as a result of abusive levels of grazing. Employing paired-stand anal- ysis techniques, range ecologists have provided extensive documentation of compositional changes attributed to grazing. Native species of 1987] HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION FiGuRE 19. Abies mes (center) and Pinus flexilis (left & right) on King's Hill (2,200 m) in northern aa ranges, central Montana; Lewis and Clark National Forest, Montana. USDA Forest Service. Agropyron, Festuca, Poa, and Stipa are classed as "decreasers" which are reduced in abundance. In their place, introduced grasses and forbs in- vaded and, for all practical purposes, have cre- ated new vegetational equilibria on the western rangelands. As an example, cheatgrass (Bromus tectorum), an introduced annual grass, has prov- en to be a superior competitor in western bunch- grass communities (Harris, 1967; Mueggler & Harris, 1969; Daubenmire, 1970; Mueggler & Stewart, 1980) and has come to dominate large areas in the middle and northern Rockies. Other invaders that have become widespread maculosa, Cirsium vulgare, Tra on dubius, Euphorbia esula, ypericum perforatum any of these species, along wit dbeckia oc- nifer forests severely disturbed by logging and/ or domestic grazing. Man's present-day penchant for working land on higher and steeper slopes is contributing to an acceleration of vegetation change. Furthermore, conflicts between agricul- ture and livestock, and winter foraging by big game, frustrate efforts to reverse the loss ofnative plant components. OTHER ROCKY MOUNTAIN VEGETATION TYPES Other vegetation types occurring within the northern Rockies are described below. Some of these are distributed throughout the northern Rocky Mountains, i.e., timberline/alpine tundra and wetland/riparian types, while others are con- fined to single mountain ranges or unique local- ized habitats. TIMBERLINE AND ALPINE TUNDRA Upper treeline reaches 2,275 m in southern Alberta and nearly ing summer. Sharp temperature fluctuations also create severe winter stresses. In contrast, inland maritime timberline/alpine tundra is more moist ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FIGURE 20. nes and nival life zones in vicinity Montana. USDA Forest Service. than the continental areas and experiences less extreme winter temperatur -ommon northern ipis Mauntaln umber- line tree species have been identified earlier; these intermix islandlike within the lower alpine tun- dra zone. Slope aspect, microtopography, pat- terned ground (resulting from cryopedogenesis), soil chemistry, soil water potential, and wind ex- posure combine to form an environmental com- plex (Fig. 20) that dictates the arrays oflow shrubs, perennial herbs, and graminoids forming the northern Rockies timberline/tundra ecosystems (Griggs, 1938; Choate & Habeck, 1967; Bamberg & Major, 1968; Habeck, 1969; Billings, 1969, 1978; Smith, 1969; Johnson & Billings, 1962; Arno & Hammerly, 1984; Bliss, 1985; Spence, 1985; Peet, 1987). Alpine permafrost occurs in some northern Rocky Mountain ranges, where mean annual air temperature is at or below 0°C and may, accord- ing to Pewe (1983), date back to Wisconsin time when such frozen soil was widespread 1,000 m of Beartooth Pass (3,300 m), Custer National Forest, below present-day elevations. Such permafrost areas coincide with modern treeless areas where alpine vegetation may favor maintenance of the frozen soil conditions. Vegetation dynamics on sites that have experienced recent ice retreat have been described in detail by Spence (1985) for parts of Wyoming's Teton Range. He investi- gated plant invasions on moraines fronting gla- ciers. Continual disturbances take place on the moraines, and compositional differences among his study areas do not seem om related to a successional chronosequenc Bliss (1985) reviewed His clivsiolorical ecol- ogy of alpine and timberline plant communities in North America, adding to earlier efforts that emphasized floristic phytogeography. He stated that complexities of community structure and composition in the alpine zone, including spatial discontinuities between mountain ranges, have permitted only generalized classification units. Growth and survival in the alpine are closely related to growth forms. Bliss (1985) listed and 1987] discussed the following major growth form cat- egories: 1) annuals (uncommon), 2) cushion and rosette-forming species occurring on exposed ridges, 3) graminoids found in all alpine habitats, 4) forbs, and 5) deciduous and evergreen shrubs, many of which are low mat-forming types. Based on an extensive literature review, John- son & Pfister (1981, 1982) listed the following alpine tundra community types typical of the Rocky Mountains. The common dominants are listed for each type: 1. Shrub Communities: Salix rotundifolia, S. re- ticulata, S. arctica, Betula glandulosa, Dry octopetala, Kalmia microphylla, Phy llodoce empetriformis, P. glanduliflora, and Vaccin- ium scoparium. . Meadow and Turf Communities: Geum ros- sii, Deschampsia caespitosum, Carex tolmiei, C. nigricans, Juncus s Poa alpina, Phleum alpinum, Polygonum bistortoides, and Oxyria digyna. Bog and Fen Communities: Sphagnum spp., Carex aquatilis, C. rostrata, C. simulata, Er- iophorum spp., Juncus spp., Pedicularis groenlandica, P. contorta, Kalmia micro- phylla, Ranunculus spp., Calamagrostis can- adensis, and Eleocharis spp. Cushion Plant and Fellfield Communities: Si- lene acaulis, Dryas octopetala, Trifolium nan- um, Luzula spicata, and Selaginella densa. . Boulder Field Communities: Geum rossii, Mertensia spp., Ribes cereum, Polemonium spp., Aquilegia spp., Penstemon fruticosus, and Sibbaldia procumbens. Snowpack Communities: Carex nigricans, Juncus drummondii, Erythronium grandiflo- rum, Valeriana sitchensis, Luzula glabrata, and Senecio triangularis N Ww A un on The nival zones of the Rockies (Figs. 5, 6, and 20) exhibit snowfields that support cold-adapted floras composed predominantly of algae, but not vascular plants, although some fungi species may also be present (Garric, 1965; Vinyard & Whar- ton, 1978). Snow algae occur on the surfaces of glaciers and snowfields and may give the snow a reddish color (or even yellow, green, blue, black, or purple), depending on the species involved. These algae have enzyme systems that catalyze reactions most efficiently at lower temperatures (Hoham, 1975). In the Rockies, and elsewhere in western North America, one of the most com- mon species is Chlamydomonas nivalis, which is known to cause the pink-red “watermelon HABECK —NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 833 snow." Red coloration on the snow surface in summer is caused by an accessory photosyn- thetic pigment called astaxanthin, which in- creases in abundance in intense sunlight. The cryophilic algae encyst during the four to eight months of winter and break dormancy in sum- mer. Other snow algae listed by Garric (1965) in- clude Scotiella nivalis, S. cryophila, Chodatella brevispina, C. granulosa, Raphidonema nivale, R. tatrae, and Romeria elegans var. nivicola. Snow fungi that may be saprophytic on snow algae include Chionaster nivalis and Selenotila nivalis, while another group of fungi, the Chy- tridiales, has been reported to be parasitic on snow algae (Vinyard & Wharton, 1978). The snowfield ecosystems also support ciliated protozoans, ro- tifers, springtails, and snow worms. WETLAND TYPES Wetland ecosystems in the Rocky Mountains occupy landscape units such as marshes, swamps, bogs, fens, and other topographic depressions with high water tables. Water may cover the sites ephemerally or intermittently and may be shal- low or deep (under or over 2 meters). The term “wetland”? also embraces wet meadows, pot- holes, sloughs, riparian zones (along stream ses), and river-overflow areas. Permanently- filled, shallow lakes and ponds less than 2 meters in depth, supporting emergent vegetation, are also included in this term. Wetland site definitions and classifications have been the subject of thor- ough attention by Windell et al. (1986). Their review of Rocky Mountain wetlands represents a comprehensive treatment of the subject and is the primary source for my synthesis of Rocky Mountain vegetation. Rocky Mountain wetlands do not exist as re- gional climaxes, as in some high latitude arctic areas, because of the continental climate which prevails over much of the region. Snowmelt run- off is rapid, much of the northern Rockies do not receive high amounts of precipitation, and evap- oration rates are high in many areas. Thus, wet- land communities are formed only in certain to- pographic settings where an abundance of water occurs seasonally or permanently. In their ecology of Rocky Mountain wetlands, Windell et al. (1986) provide detailed informa- tion on community structure and a general clas- sification of these ecosystems. Their literature review covers the entire Rocky Mountains. An 834 overview of their findings is given below, sup- plemented with information taken from other regional studies (Pfister & Batchelor, Youngb 1984; ood et al., 1985). An outline of wetland community types will be presented, together with a listing of plant dominants for each. l. N 3. Permanent Shallow Standing Water (less than -4 m): Floating communities: open water surface dominated by Lemna spp. b. Rooted submergent communities: domi- nated by species of /soetes, Nitella, Pota- mogeton, Najas, and Myriophyllum. ooted floating-leaved communities: dominated by species of Nuphar, Nym- phaea, Sparganium, and Potamogeton. . Rooted emergent communities: dominat- ed by species of Carex, Eleocharis, Juncus, Glyceria, Phragmites, Sagittaria, Scripus, Sparangium, Typha, Menyanthes, and Petasites. Ë tƏ ° e. . Seasonal or Permanent High Water Tables, Without Permanent Standing Water: Herbaceous wetlands: floating mats dom- inated by species of Carex and Sphagnum. On mineral soils: marshes or wet meadows dominated by grasses, sedges and rushes, P . Fe erbaceous wetlands on organic soils and dominated by sedges (Carex spp.) and other graminoids such as species of Juncus, Eleocharis, Deschamp- sia, and Calamagrostis. Bog communities: Northern Hemisphere bogs are dominated floristically by a mat of Sphagnum spp. and members of the in the mat. Rocky Mountain regional bogs include species of almia, Drosera, Menyanthes, Eriopho- rum, Potentilla, and Gaultheria. . Marsh/wet meadow communities with fresh water: occur on mineral soils and are dominated by herbaceous species of the following genera: Carex, Deschampsia, Danthonia, Juncus, Penstemon, Erigeron, and Calamagrostis. Marsh/wet meadow communities with sa- line water: alkaline sites with sodium con- centrations over 15%, supporting onl salt-tolerant species such as Distichlis, a Puccinellia, Tr. ie lochin, and Salicor Forested Wetlands: Rivers and streams in the = 5 e ° B B £ 5 = © e e. ° ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Rocky Mountains that have well-developed floodplains supporting broadleaf deciduous forests and woodlands. The term “riparian wetlands” refers to plant communities oc- curring adjacent to running water. Rocky Mountain riparian forests are dominated by Populus (P. trichocarpa, P. deltoides, P. an- gustifolia, P. balsamifera, and P. tremu- loides), as well as Betula papyrifera, Fraxinus pennsylvanica, Ulmus americana, and Acer negundo. Needle-leaved evergreens such as icea engelmannii, Pinus ponderosa, Pseu- dotsuga menziesii. Thuja plicata, and Abies grandis also exhibit local dominance on ri- isa sites extending through elevational zones. Mixtures of Populus spp., Juniperus ny pa and/or Pinus ponderosa are also ound on floodplains. Such sites are subject to frequent disruptions by flooding and do- mestic grazing (and historically by fire), and communities are often in various stages of successional recover Shrub-dominated Wetlands (Carrs): Some wetlands are dominated by shrub species rather than by forbs, grasses, or trees. Shrub species listed by Pfister & Batchelor (1984) as dominants in the northern Rockies are Salix bebbiana, Cornus stolonifera, Alnus sinuata, A. tenuifolia, Betula occidentalis, B. glandu- losa, Prunus virginiana, Sambucus melano- spp., Symphoricarpos spp., Rhamnus alni- folia, and Acer glabrum. . Herbaceous Wetlands along Streams: Along fast-moving, steep-gradient streams are ri- parian communities that are dominated by mosses and/or a variety of herbaceous vas- elyma, Fontinalis, Funaria, Hygrophynum, Philonitis, and Oncophorus. Streamside herb- dominated communities feature species of Mertensia, Senecio, Mimulus, Heracleum, Delphinium, Aconitum, Primula, Saxifraga, Veratrum, and Athyrium ASPEN COMMUNITIES Trembling aspen (Populus tremuloides) forms clonal stands or continuous forests under various site conditions in all mountain vegetation zones 1987] from 900 m to 3,600 m throughout the Rocky Mountains. It intermixes with semiarid shrub- ands and with wet spruce-fir forests. It is the most widely distributed native North American tree; aspenlike trees appear to have been present in western North America since middle Miocene times (Harper et al., 1985). Aspen populations are variable, but no subspecies are recognized. The most expansive display of aspen in the Rocky Mountains occurs in the middle Rockies (Reed, 1971; Smith, 1985; Mueggler & Campbell, 1986), and it forms scattered grovelands in the northern Rockies (Lynch, 1955; Steele et al., 1981; Arno & Hammerly, 1984). Biological and ecological attributes of aspen in the western United States have been recorded by DeByle & Winokur (1985). Aspens reproduce profusely by root shoots (“suckers”), especially following fire treatment. Establishment from seed is rare and may occur only during unusually wet years. As shown in Figure 18, aspen communities are often fire-induced pioneer or seral stages that become invaded and replaced by conifers (Lynch, 1955), while under other conditions— beyond conifer’s limits—the groves may perpetuate themselves as local uneven-aged climax stands (Steele et al., 1983; DeByle & Winokur, 1985). Aspen communities do not readily burn, but aspen’s thin bark makes it extremely sensitive to fire. Although it is considered a fire-dependent community type, aspen stands have low flamma- bility characteristics (Mutch, 1970). Modern fire suppression has led to an increase in mature as- pen; young stands are not common (DeByle & Winokur, 198 Populus tremuloides communities have been subject to heavy use by domestic stock as well as by elk and moose, leading to replacement of un — arvense, Rudbeckia occidentale, Poa pratensis, and Helianthella spp. (Steele et al., 1983). Pre- vious severe browsing of aspen shoots by elk and moose on the Yellowstone Park winter range, together with fungi and insect damage, led to the demise of aspen (Krebill, 1972). Recent browse pressure reduction has allowed vigorous aspen regrowth (Krebill, 1985, pers. comm.). MINOR COMMUNITY TYPES Pinus flexilis has been mentioned as a member of both the lower and upper timberlines within HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 835 parts of the southern Canadian Rockies, north- ern Rockies (confined primarily to the East Fronts in Alberta and Montana), and middle Rockies. It also deserves mention as a dominant pioneer- ing tree occupying volcanic deposits (cinder cones and lava flows) in and near the Craters of the Moon National Monument (Fig. 4, Area 22) in a Idaho (Eggler, 1941). Some Pseudo- tsuga menziesii is also found with Pinus flexilis on these extremely xeric sites, as are Artemisia tridentata, Chrysothamnus nauseosus, Purshia tridentata, and Ribes cereum. communities occupying localized mi- crosites, such as talus/scree, ponds, bogs, snow glades, rock outcrops, rock crevices, vernal pools, and sand dunes have been described by Chad- wick & Dalke (1965), Daubenmire (1970, 1975, 1978, 1980), and by others. Throughout the Rocky Mountains, there are numerous special- ized habitats that support minor communities often composed of rather unique combinations of plants. Some of these species have been iden- tified by Rocky Mountain botanists as rare and/ or endangered. Furthermore, some parts of the northern Rocky Mountain region support nu- merous geographic disjuncts and endemics, such as coastal Cornus nuttallii and Alnus rubra pop- ulations; these occur in the intermountain ranges in northern Idaho (Daubenmire, 1943, 1975; Johnson, 1968; Johnson & Steele, 1974). CONCLUDING REMARKS The Rocky Mountain vegetation exhibits vari- ations in its local and regional distribution pat- terns as a result of the diversity associated with this region’s physical setting and a variety of his- namic nature of vegetation is the basis of its re- sponses to spanning mil- lions of years. Any modern description of the Rocky Mountain flora and plant communities is really only capturing a “moment” in an ever- changing phenomenon. 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Western intermountain sagebrush ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 steppe. Pp. 351-373 in N. E. West (editor), Tem- perate Deserts and Semi-deserts. Ecosystems of the World, Part 5. Elsevier Sci. Publ. Co., New WHITTAKER, R. H. 1953. A consideration of the cli- max theory: the climax as a population and pat- tern. Ecol. Monogr. 23: 41-78. T. R. LILLYBRIDGE. 1983. For- ested plant associations of the Okanogan National Forest. USDA Forest ande Pacific Northw. Reg., Publ. R6-ECOL-132b: Cooper, S. Q. : ,L. : 1986. Si ecological characterization of ocky Mou and subalpine wet- lands. "USDI Fish 4 Wildlife Ser. Biol. Rep. 86(11): 1-298. WRIGHT, J. C. & E. A. WRIGHT. 1948. Grassland types of south-central Montana. Ecology 29: 449- 460 1981. YOUNG, J. & R. EVANS. Demography and fire history of a A a stand. J. Range Man- agem. 34: -506. YOUNGBLOOD, A. 5 & W. F. MUEGGLER. 1981. As- pen community types on the Bridger-Teton Na- tional Forest in western Wyoming. USDA Forest Service. Inter. Forest. & Range Exp. Sta., Res. Pap. INT-27 — ADGETT & A. H. WINWARD. 1985. Riparian community ty cl astern Idaho-western Wyoming. USDA Forest Service, Inter. Reg. Publ. R4-ECOL-85-01: 1-78. COMPARATIVE AGE OF GRASSLAND AND STEPPE EAST AND WEST OF THE NORTHERN ROCKY MOUNTAINS! E. B. LEOPOLD? AND M. F. DENTON? ABSTRACT Given the taxonomic and biogeographic differences between dominant species in grasslands of the Great Plains and the palouse grasslands and steppe of the two biomes have had separate origi l of the Rocky Mountain cordillera and pollen data from the cordillera area suggest that three distinct floristic provinces were in existence by the beginning of mid-Miocene (Barstovian) time. Montane conifer forest poor in genera typified the Rocky Mountain a and nearby pesi while mixed conifer-deciduous hardwood forest and Taxodium swamps rich in woody genera occurred in the Columbia Basin. On the Great Plains the Kilgore flora (Barstovian of Nebraska) iugo deciduous open forest and prairie dominated by species of eastern and southern affinities. Younger Neogene floras in the Pacific Northwest suggest that steppe and local grassland were beginning to be important in the Pliocene of the Pacific Northwest, while grasslands and open forest became widespread on the Great Plains somewhat earlier (late Miocene or Clarendonian-Hemphillian time). The persistence of mixed conifer and broad-leaved forest on the Columbia Plateaus suggests that this area was open to the west through about 8 Ma. Due to the increasing height of the Cascade Rangea nd/or regional up- According to the regional pollen record, terrestrial herbs become more diverse in the late Neogene. Recent fossil evidence has helped substantiate that the grasslands and steppe west of the tinental Divide in the Columbia Basin area are about three million years (Ma) old, about ten million years younger than similar vegetation types east of the Divide in the Rocky Mountain foothills and in the Great Plains. Biogeographic differences between the two areas are reflected in species composition and in the dominant habits of grasses. Some climatic differences between ‘east’ and ‘west’ exist, but they do not seem suf- ficient to account for the vegetation contrasts. We propose that an examination of the Late Cenozoic history (Miocene to present) of the northern Rocky Mountains may help illuminate the nature of origins of the grassland and steppe east and west of the Divide. The region to the America, while the region to the east, the Rocky Mountain foothills and the Great Plains, have Mio-Pliocene deposits for which pollen, leaf, and seed data are now available. To understand the history of grassland and steppe development, be- cause they hinge on the present east/west con- trasts, the following questions need to be an- swered: 1) When did grassland and/or steppe first develop in the northern Rocky Mountains area? 2) What were the regional patterns of climate during the time the grasslands developed? The Pliocene) vegetation inferred from pollen or leaf data. Species identifications from the leaf and seed floras give an index of floristic patterns that can be compared with the perceived regimes. It is our thesis that three distinct vegetation provinces had developed in the northern Rocky Mountain region by early mid-Miocene time (ca. 16 Ma): 1) a rich deciduous hardwood and mon- tane conifer forest west of the Continental Divide mbi with steppe elements in the Rocky Mountain foothills; and 3) a deciduous forest with ancestral grassland elements to the east and in the Great Plains. If vegetation types east and west of the cor- dillera were basically different during the Mio- ! We thank Daniel Axelrod, H. D. MacGinitie, seal s“: Malde, Jack A. Wolfe, and Robyn Burnham for their comments, and M. Kay Suiter for typing the ma 2 Department of B 3 “Columbia Plateaus” ANN. MISSOURI Bor. GARD. 74: 841—867. 1987. Botany, University of Washington, corns “Washington 98195, U.S.A. is the official physiographic usage by the U.S. Geological Survey (1981). 842 ANNALS OF THE MISSOURI BOTANICAL GARDEN FIGURE 1. Present distribution of the chief grass- and vincesin North America according to dominant a io om Daubenmire, 1978).—A, B. Andropo. gon scoparius province. — C. Festuca scabrella province. — D. Bouteloua gracilis province. — E. Agro- pyron spicatum province cene, the importance of this massif as a floristic barrier can be evaluated. Did the same species occur east and west of the divide or were they different? How do these vegetation patterns re- late to the Neogene history of the region and to the development of grassland and steppe biomes? PRESENT VEGETATIONAL PROVINCES Figure 1 illustrates the outlines of ied grassland and steppe types in the U. each side of the Rocky Mountain cordillera mus grass- lands of different character. (1) At the eastern margin of the mid-continent grassland, there is the tall-grass prairie, a sod-forming grass asso- ciation with rhizomatous root habit. The affin- ities of the dominant taxa, Andropogon scopar- ius, Andropogon gerardii (little and big bluestem), Panicum spp., and Sorghastrum nutans (Indian grass), lie to the south, in Central America, Mex- ico, and even South America. (2) The short-grass prairie immediately east of the cordillera, char- acterized by the dominance of Bouteloua gracilis (buffalo grass), Aristida spp., and others, is a province wit izomatous and ESAE grasses. Most of the dominant taxa have t floristic affinities with the intermontane dn of the Rocky Mountain region. (3) To the west exists the palouse grassland and steppe of Idaho, [Vor. 74 X — Ns AT T Sa. ( mM a c 150 Y X \ — ——— 5 I N J j c Agropyron spicatum | FIGURE 2. catum (after Hultén, 1968; Hitchcock et al., 1972). Present distribution of Agropyron spi- 1969; Voss, eastern Washington, and eastern Oregon (Fig. 1), characterized by bunch grasses and steppic ele- ments such as Atriplex, Artemisia, Sarcobatus, and other diverse desert-scrub genera. The bio- geographic affinities of taxa in the palouse grass- land and steppe are with areas to the north; for example, Agropyron spicatum ranges northward to Alaska and boreal regions (Fig. 2). Festuca idahoensis has its nearest relatives, the F. ovina omplex, in the arctic and steppes of North America, Asia, and northern Europe (Fig. 3). Ar- temisia cana (Fig. 4) and other members of the A. tridentata group are mainly arctic or boreal. Climates east and west of the Continental Di- vide differ significantly, chiefly with respect to the amount of summer precipitation. On the Great Plains summer rain emanates from trop- ical air masses moving northward from the Ca- ribbean across this region during June and early July (Bryson & Hare, 1974). Along the southern cordillera summer rainfall occurs, especially dur- ing warm years, along the north-south path of the “Arizona Monsoon" (Neilson, 1986; Neilson & Wullstein, 1983). To the west, the palouse grassland and steppe region is characterized by l e area is dominated by the Pacific air in summer and is in the rain shadow 1987] FIGURE 3. Hultén, 1968; Hitchcock et al., 1969 of the Cascade Mountains; here, the distribution of grassland and steppe is limited to areas re- ceiving less than 15-20 inches (380-500 mm) of annual precipitation, most of which falls in win- ter (Fig. 5). The arand provinces east and west of the Rockies are now dominated by separate taxa dif- fering in geographic affinity and in growth habit (Table 1). These grasslands differ in distribution of C, (warm growing season) and C, (cool grow- ing season) grasses. C, grasses are always of low frequency (less than 1896) west of the Rockies t may dominate in certain areas east of the Rockies (Mack & Thompson, 1982; Teeri & Stowe, 1976). In addition, there are fundamental differences between these two grassland prov- inces with respect to their carrying capacities for large, grazing ungulates; for example, the carry- ing capacity of the palouse grassland dominated by bunch grasses sensitive to trampling is much lower than that of grasslands east ofthe cordillera (Mack & Thompson, 1982). The biogeographic, physical, and climatic contrasts of these regions imply that thei t must have been very different. This is the subject of the discussion that follows. DATA BASE Because our study concerns events after the early Miocene, we discuss data from sites youn- ger than 18 Ma (Hemingfordian and younger stages). The sites are mapped in Figures 6 and 7 LEOPOLD & DENTON —GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 843 Present distribution of Festuca idahoensis and the closely related species Festuca ovina (after ). and are arranged according to geologic ages in Table 2 West of the Rocky Mountain cordillera, the area of the Columbia Plateaus contains a wealt of well-documented fossil plant sites. We have studied data from 15 of these, of which four have yielded pollen data (Fig. 6). Dating is established by K/Ar isotope ratios for a few of these and by land-mammal evidence for others. 150 Artemisia cana E 4. Present distribution of Artemisia cana (after Hultén, 1968; Hitchcock et al., 1955 844 ANNALS OF THE MISSOURI BOTANICAL GARDEN 20“ ISOHYET -——-— 15“ ISOHYET ------. == 80 0 SCALE IN MILES FOREST OR WOODLAND AREAS øm [Vor. 74 Relationship of the forest/steppe border to the 15- and 20-inch (380 and 500 n isohyets of "en ETE in the Pacific Northwest (after Sherman, 1947, and U.S. Weather Bureau data In the foothills of the northern and e Ps Mountains, we have four localities (Fi 11 which the main documentation is His fossil NS although some megafossil dataexist data. Ln of taxa is on the generic level only). y scattered, but their eoi ages were ascertained by land-mammal 120 110 100 | i L ` i \ ^ I ` l ' \ i ! RO ee 48 vy Gee E i E LL EE 4 ae aie NOR : SP) WASHINGTON r3 aa i ` ; | ' E 1 Á \ aen | Y | NORTH DAKOTA ) 46 . Ellensburg @ ; ` | N l \ Hanford Clarkia 7 MSNIANA ! n —2 Cone ; | sme S G | Deschute Ve i eschutes g Mascall IDAHO N m 4 a permet eee err MM | ii UNS | SOUTH DAKOTA lo SS WYOMING | Stinking Water Í W Horseshoe Bend e Teewinot i S [e k ^l GI F | cs cu de Uccor reer: ; enns Ferry — 42 | | OREGON Trout Creek Moonstone s Split Rock | ul . - 1 oper LP! \ pesi eas ONO _ i NEBRASKA i T = Saratoga | ! e i ! I “SZ = E a 40 . ! —— nem I i | Troublesome v7 | | e ! : I i NEVADA ! UTAH i i | i COLORADO \ KANSAS ES ! : : 38 E i : E : | FiGuRE6. Locations of Miocene and Pliocene fossil flora sites in the northern and central Rocky Mountains and Columbia Plateaus. Bi = le ollen af localities; @ = po sites; A indicate leaf and pollen data from the same site. Coal Mine Gulch (not shown) is 50 miles north of Succor Cree 1987] LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS In the region east of the cordillera, there are several megafossil localities (Fig. 7) containing small leaf floras and there are abundant fruit and seed localities (Elias, 1942); recent work with pollen analysis and electron microscopy of fruit- ing parts, grass anthoecia, epidermal patterns, and phytoliths revealed the composition of many of these assemblages. MIOCENE VEGETATION PROVINCES OF THE NORTHERN RocKY MOUNTAIN REGION Three vegetation provinces had developed by the mid Miocene in the northwestern and north- central mid-continent. I. WEST OF THE ROCKY MOUNTAINS — THE COLUMBIA BASIN A. MIOCENE FLORAS Fifteen leaf floras of mid and late Miocene age demonstrate that mesic forest vegetation under a warm-temperate summer-wet climate existed in the Columbia Basin by about 18 Ma until Hemphillian time (about 8-4.5 Ma). Difference in warm- and cold-season average monthly tem- perature has been estimated at about 20°C Mae 1978). Even though important changes in cli- mate and vegetation occurred through the main sweep of Miocene time, the major elements of the flora were not eliminated. Some chieffeatures of the vegetation derived from the leaf flora are: (1) Dominant vegetation (Trapper Creek? of Clarendonian age is an example; summary in Table 3) was deciduous hardwood forest and mixed montane conifer-deciduous forest, with some broad-leaved evergreen elements and di- verse (8-26 genera) woody dicots. Shrubs were important (up to 3096), while herbaceous groups were few (only four taxa and these were generally rare aquatics). (2) Species showed close relationships with of summer-wet areas in eastern about equal percentages. For example, at Trap- per Creek modern affinities seem split between * Axelrod (1964) considered the age of the Trapper Creek flora as 15-16 Ma. More recent evidence from K/Ar dating (Armstrong et al., 1975; Fields, 1983) sug- gests an age of 10.5-12 Ma. Chief characteristics of modern grassland types in the U.S.A. TABLE l. EAST Rockv MOUNTAINS Tall-grass prairie Short-grass prairie Grassy steppe Agropyron spicatum P KEY SPECIES: Andropogon scoparius Bouteloua gracilis a O Festuca idahoensis Panicum virgatum Sorghastrum nutans Sporobolus (southern affinities) (intermontane basin affinities) (north temperate & boreal affinities) DOMINATED MAINLY BY: rhizomatous/stoloniferous grasses caespitose (bunch) grasses summer moist summer dry CLIMATE: 845 846 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 : nm (e mmm (c) Valley forest, including some of the taxa DAK 1 . : ar TIT 3 en s : mentioned above and a wide range of woody | oshin Drank n deciduous groups such as Alnus, Amelanchier, » IOWA : : WYO, | Ash Hollow fm2 9 i runus, Parthenocissus, Cornus, Fraxinus, Ul- BENE € ) mus, Pterocarya, Carya, and Sophora as well as — NN. ! y uu e ------ -| an important component of broad-leaved ever- eWray j S 404 greens, such as Quercus (cf. Q. chrysolepis), Sas- E s. safras, Berberis, Ilex, and diverse conifers such ogan Co. ' š ; ; COLO, |@Wallaceco@ Ellis Co. T as Keteleeria, Picea (cf. P. breweriana), Pseudo- tsuga, Tsuga, Abies, and Sequoiadendron. (d) Mountain-slope forest, including many of the conifers and hardwoods mentioned in the ae. 9 Beaver I------1 valley forest, also Garrya (G. cf. fremontii), Rhus, Ghian x Ungnadia, Abies (A. cf. delavayi or A. recurvata), ° | ! Abies concoloroides, Calocedrus (C. cf. decur- | i ARK rens), and most importantly, Pinus (P. cf. pon- | derosa, P. cf. monticola), and Cedrus. Some of the most frost-sensitive forms (Liq- ane Vi, | Uidambar, Cedrela) seem to disappear during the ' mid Miocene, especially in regions close to the Location of Miocene and Pliocene fossil Rocky Mountains. However, the occurrence of flora sites on the Great Plains (High Plains of Axelrod, warm temperate Taxodium swamp vegetation 1979). persisted in eastern Washington until ca. 8 Ma and in Idaho until ca. 12 Ma. LJ Clarendon NEW MEX. M ee s FIGURE 7. Although Miocene communities typically in- these three areas (Axelrod, 1964). There were cluded species of eastern Asian and eastern only minor temporal changes in the role of these American affinities, western American elements geographic elements through mid-Miocene time seem to dominate in the montane slope forest (Table 4). communities. _ GB) Vegetation was as Chiefly woody. Character- Chaney (see Chaney & Axelrod, 1959: 53) dis- forthe mid- cussed the possibility that open savanna or prai- Miocene floras of southern Idaho included: rie vegetation existed at some mid-Miocene sites (a) Swamp forest, particularly Taxodium where pollen and leaf data are available, e.g., swamp with associated Nyssa, Liquidambar, | Mascall and Stinking Water floras of eastern Or- Persea, Salix, and Alnus. egon. Part of his rationale was based on the di- e-border woodland, with Quercus sim- verse fossil mammals whose modern relatives ulata and species of Acer, Betula, and Popu- livein savanna habitats today ieee horses, lus. rodents, lagomorphs, camels, and most abun- E 2. Stratigraphic ages of Late Cenozoic floral localities of the Columbia Plateaus, northern Rocky Mountains, and Great Plains. EASTERN | EASTERN SOUTHERN|NORTHERN| WYOMING | NORTHERN Ma OREGON WASH. IDAHO IDAHO COLORADO| NEBRASKA uaternary | Bruneau Pliocene ` |Blancan Glenns Ferry ain Deschutes è Hemphillian Chalk Hills E] _ Hanford Banbury Basalt ering " Clarendonian | Ellensburg | rappar Creek am Hollow e 12 Stinking Water oison Creek Š Trout Vantage © 3 Barstovian nad Payette Saratoga Kilgore s _ Jj _ Clarkia _ _ | Sheep Creek 16 > Hemingfordian ist OAE Ww 18 1987] Dominant vegetation types in Late Cenozoic floras of the Columbia P LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 847 t (from Axelrod, 1964; TABLE 3. Chaney & Axelrod, 1959; Graham, 1963; Leopold & Wright, 1985; MacGinitie, 1933; Smiley & Rember, 1985; Taggart et al., 1982) Barstovian Clarendonian Hemphillian LJ ° v S 5 E S a D S x " ° x x e OQ 8 a Š O E > o 9 m m z 905 š g D Ble 5 8 S o & w d uc pt y Jat Els O ` S @ €t Ss qd Bou 9 £ afla Els = o ` oO S Š ‘9 o A < © o o|8 gje = Ó GY É uü ^? > à 3 £ 285 SS e 9 8 6 8 8 S E S S s os Sie B à D$ > OE 4d doe OF «BO Om Mixed deciduous evergreen hardwood forest X Deciduous hardwood forest X X X X X cotone X X X Montane conifer-deciduous hardwood forest X X X X X Ecotone X Montane conifer forest X Grassland and conifer forest X Steppe and conifer forest X X Steppe X Taxodium swamp forest X X X x x x dant, the oreodonts). Direct evidence, however, is limited. Herbaceous plants are an important elementin modern savanna. Aside from aquatics and ferns, herbaceous groups are rare in the Mio- cene leaf record of the Columbia Plateaus. Even pollen evidence indicates that herbaceous groups were limited in diversity and abundance (Ap- pendix I); in the Mascall only two nonaquatic herb types were identified by Jane Gray (in Cha- ney & Axelrod, 1959: 43): “Pollen of Gramineae h were found, but these, as in the leaf flora, were all from woody taxa. The floristic role of herbs in mid Miocene of the region is illustrated in the pollen lists from the Succor Creek Formation (14-12 Ma at the type section; Fields, 1983) in southeastern Ore- gon (Taggart & Cross, 1980). Six probably terrestrial herbaceous groups are recorded: Pachysandra, Ambrosia, Onagraceae, Amaran- thaceae, Gramineae, and Umbelliferae (Appen- dix I). Because the pollen diagrams summarize their abundance according to broad ecological groupings, the relative importance of herbs is not documented. In their discussion Taggart & Cross made it clear, however, that grass and Compos- itae pollen are sporadically abundant as part of a successional cycle (see below). TABLE 4. Percentages of element representation in mid-Miocene floras and age groups, Columbia Plateaus region (from Chaney & Axelrod, 1959, table 32; Axelrod, 1964, table 5). Barstovian Clarendonian uccor Trout Stinking Trappe Lower Geographic Elements Mascall Creek Creek Water poe Ellensburg Eastern American 65 61 50 58 58 70 Eastern Asian 50 50 50 53 69 45 Western American 37 37 56 60 62 58 Total taxa 64 46 46 38 61 32 848 A. SUCCOR CREEK EASTERN OREGON % total pollen measured er m ection section === E3 Locustrine F -— ANNALS OF THE MISSOURI BOTANICAL GARDEN 14-12 Mo [VoL. 74 B. SUCCOR CREEK MODEL PALEOASSOCIATIONS Generalized pollen diagram indicating vegetation phases in a 200—meter section at the Succor of E 8. C 2 "fora type locality shown on right (from Cross & Taggart, 1982 (4) Vertical arrangements of plant commu- nities were suggested by pollen data. Many leaf floras do not demonstrate clearly that an altitu- dinal zonation occurred in a the Miocene floras, yet pollen-st | indeed. At Succor Creek, Oregon, Taggart & Cross (1980) and Taggart et al. (1982) showed a re- peated successional cycle of montane conifer for- est, bottomland/slope associations, and xeric shrub with steppic elements followed by Pinus spp.; the sequence then reverts to swamp, bot- tomland/slope, and then to montane conifers. A single cycle in a 200-meter section is shown in Figure 8. The dramatic oscillations can readily be interpreted as elements from various altitu- dinal in an altitudinal tcertain succession caused by climatic changes. Taggart et al. (1982) believed the successional changes are related to disturbance episodes of volcanism, but it is also possible that these are forced by small oscillations of climate, or both. Ata much younger (ca. 8.5 Ma) site at Hanford in eastern Washington, floodplain sediments (lowest Ringold formation) of the ancestral Co- lumbia River record a somewhat similar succes- sion (Fig. 9a, b; Leopold & Nickmann, unpubl.). The sediments overlie the Upper Columbia Riv- er Basalts (10.5 Ma) and an unconformity dated at 8.5 Ma (Tallman et al., 1981; DOE, 1986: 3- 40). The lithology indicates that local deposi- tional environments were changing. In a series mid Miocene age, eastern Oregon (A); a model of the successional sequence (B) is ). of cycles, fine-grained swamp deposits grade up- ward to increasingly coarse riparian sediments. Two pollen diagrams (core holes DC-3 and DC- 7/8), each beginning in fine-grained sediments, register a rich Taxodiaceae-type (cf. Taxodium) swamp association. (We infer this identification because Taxodium is abundant in the underlying Ellensburg Formation.) This phase is followed i int tt l d/sl I 1 d and 1 by p ; finally Cedrus and Pinus dominate. Herbs and xeric elements are always present in trace amounts. Presumably in a floodplain area such as Hanford, the vegetation changes record either shifting riparian environments (edaphic factors) or changes wrought by climate. While the ele- vational relief of each of these forest types oc- cupied is not clear at either site, it is possible that as much as 500-1,000 feet of relief existed at Succor Creek. The Cascade Range was probably rising during Miocene time (McKee, 1972; Smiley, 1963), and its rain shadow eastward eventually changed the ici of the vegetation from mesic and sum- r-wet to xeric and summer-dry. The lower Ellensburg and the Hanford floras suggest that eastern Washington was open to the west through ca. 8 Ma. This meant that the Cascades were not significant enough to block moisture from the westerlies until some time after the Clarendon- ian. In part this helps explain the general simi- 1987] larity of Pacific Northwest floras during the early and mid Miocene. B. IMPLICATIONS FROM PLIO-PLEISTOCENE FLORAS Given the data above concerning the charac- teristics of the Miocene forests of the Columbia Plateaus, when did forest vegetation diminish, allowing the development of grassland and steppe? West of the Rocky Mountains, pollen data from Idaho demonstrate the decline and impoverish- ment of the Miocene forests and the develop- ment of local grassland and steppe. These changes that occurred from late Pliocene to Quaternary were surprisingly late. In lake and stream de- posits of southwestern Idaho a unique and well- dated composite pollen sequence embraces parts of the last 11 million years, after Trapper Creek time through the early Quaternary (Fig. 10; Leo- pold & Wright, 1985). Fossil pollen in these de- posits tends to be scarce, and pollen-bearing beds are hard to find (90% of our collections were barren). Some megafossils have been identified at certain sites. In the sequence (Idaho Group) the lower sed- iment units are from the Poison Creek, Jenny Creek, and Chalk Hills formations and Banbury Basalt of mid and late Miocene age (Armstrong et al., 1975; Fields, 1983; Leopold & Wright, 1985; Appendix II). The floras record mixed de- ciduous and conifer forest with declining hard- woods; these were mainly U/mus but also in- cluded Pterocarya, Carya, and Juglans. A holly-leaved oak (leaf evidence from the Poison Creek Formation) is reminiscent of that recorded at Trapper Creek (ca. 11 Ma). Wood from the Chalk Hills Formation records diverse hard- woods. Younger sediments of the Glenns Ferry Formation (ca. 3-2 Ma) containing the Hager- man lake beds record an impoverished pine and mixed conifer assemblage with rare pollen of ex- otic hardwoods. Steppe elements (Sarcobatus and other Chenopodiaceae, Artemisia and other Compositae) are consistently present and in- crease sporadically upward in this section. Peaks (up to 60%) of grass pollen are associated with clines. Taxa of terrestrial herbs are more diverse than in the Miocene (Appendix I). LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 849 Miscellaneous Pleistocene samples from the area suggest that Artemisia and Chenopodiaceae are important, if not the dominant, pollen forms. Above an unconformity (see top of Fig. 10) Ir- vingtonian g Equus plus K/Ar evidence date the Bruneau Formation as middle Pleistocene; a Bruneau pollen sample (D1694) shows Artemisia to be 50% of the count, sug- gesting a true Artemisia steppe had developed. Two other samples rich in Artemisia pollen (D1120and D1698; Fig. 10), previously reported as Bruneau (Leopold & Wright, 1985), are now classified as Yahoo Clay of late Pleistocene age (Malde, 1982). The top fossil sample (D1697), showing 80% Artemisia and other Compositae pollen, is from a late Pleistocene soil above the King Hill Basalt. In northern Oregon the Deschutes flora (ca. 4— 5 Ma) suggests low-diversity riparian vegetation typical of unforested regions and is consistent with a decrease in summer precipitation in late Miocene time (Appendix III; Chaney, 1938). Taken as a whole, the data indicate that steppe in the Columbia Basin probably did not develop as a major vegetation unit until after the Hemp- hillian (4.5 Ma). The Snake River Plain section places the change from rich (deciduous and) co- niferous forests to montane conifer forest be- tween 10 and 3 Ma. In this region steppe and palouse grassland probably became widespread for the first time in the Quaternary. II. THE ROCKY MOUNTAIN FOOTHILLS The lack of megafossil evidence for northern Rocky Mountain Neogene sites is unfortunate (the well-documented Clarkia flora of middle Miocene age lies in the Columbia Basin floristic province). Pollen records from the Rocky Moun- tains and eastern foothills demonstrate the com- paratively modern aspect of plant communities there during the Miocene. At Jackson Hole, Wyoming, the Teewinot lake deposits (predating the Teton Range) provide a long record of montane conifer forest with oc- casional bursts of lowland steppe and riparian types (Fig. 11; Barnosky, 1984). Presumed plant communities include: (1) Saline basins. High percentages of Sar- cobatus pollen accompany other Chenopodi- S These are char- acterized by pollen of probable phreatophytes, ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 850 HANFORD RESERVATION SHINGTON ate Miocene £ $ oF Oe RAE ES 7 TOC 9 kA Y sd! JI E E | | iM XE ! i na i L P IE s E ES ud pe de Jn DE IT tei cold :. KASS SPS Da a E" == Walli SILT ee SILTY - == ae CLAY SAND SPORES- ra tlla Lbs from lowermost Ringold Formation, % total pollen (Leopold & Nickman, unpubl. data). Stratigraphic data are from Department of Energy (1986: 3-16) drill holes DC-3 and DC-78, which are correlative in age and about 12 miles apart. Depths below surface are given in feet. The sediments are dated at 5-10 Ma (probably 8.5 Ma). FIGURE 9A. Late Miocene pollen sequence, Hanford, Benton County, Washington. Two pollen diagrams h 1987] HANFORD FOREST ASSOCIATIONS 8.5 Ma E. Washington Taxodium Bottomland / Swamp lope Cedrus Shrubs Forest Hardwoods & Herbs + 4 + 3 + 2 + + 1 + + FIGURE9B. Schematic phases of vegetation shown by Figure 9A pollen diagrams. Carya, Ulmus-Zelkova, Pterocarya, Sapinda- ceae, and Salix. _ (3) Montane slope forest. This assemblage is dominated by Pinus spp. with important amounts of Abies, Picea, and Cupressaceae pollen. Tsuga, LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 851 Quercus, Artemisia, grasses, Onagraceae, and ad- ditional herbs may have been locally abundant (Leopold & MacGinitie, 1972: 198) The proportion of broad-leaved Tertiary relict genera that are now exotic (eastern Asian and eastern American genera such as Pterocarya, Carya, and Ulmus-Zelkova) obviously is low compared with those in pollen and leaf assem- blages of the same time period west of the Rock- ies (e.g., Hanford, Poison Creek, and Trapper Creek). Except for the “riparian” hardwoods, the flora has a modern aspect indeed, as it compares well with modern pollen rain (see top of Fig. 11). Four other pollen sites from widely different times in the Miocene are in basins along foothills and in the eastern Rockies of Colorado and Wy- oming (Fig. 12; localities in Appendix IV; se- lected pollen counts in Appendices V, VI; Leo- pold & MacGinitie, 1972). Early Miocene (Hemingfordian) sites are the Troublesome For- mation from Grand County, Colorado (Izett, MODERN: D mw 8) SAMPLE NO. CONIFERS TERT PEHE! eels ae SHRUBS 01097 ob ET! + ui D1694 r: D-1695-2 £3 s = 11698 aa Xe D1120 T es CJ 0 Fog D1671 [ i 11695-4 32a E 2 + | um rm Lan m — Te . 1 SAGE & | SALT * 01710 L — | iR Q |t [t | WALNUT, |COMPOS- BUSH z la +O *0* ^u +x HICKORY, GRASS - E 2 D1715 AAR > € A = +a Š «c |+Ə | LING NUT, o D1712 ae x + ha a = | OAK E 1: uj + + uj ore MAPLE 3 D1711 u. + |+ h + q D)713-1 2 ' + + Je z D1692-5 u + + Di692-4 e + D169) r + + D1690-1 + x od | INANA q P nisse > g =) + 2 PINE £ uj + + I| + n1208 FX + © pi OL———3À * D1699 OY — ——i (0) 80% 015 o 30 O 80 (0) TO O IOO % FIGURE 10. C omposite pollen diagram, late Miocene to late Pleistocene, Hagerman area, southwestern Idaho. old ght, n is shown Note that Pleistocene samples D1120 and D1698 are pu EAE fied as from Sample D1695-2 is Glenns Ferry Formation (Malde, at top _ a S & Wri 1985). Fossil wood indicated along is Quercus, white oak type (Malde & Powers, 1962). Yahoo Clay of late Pleistocene age. 982). Sample D1697 is from a Late Pleistocene buried soil on the King Hill Basalt. Sample D1694 is of the ed Formation of middle Pleistocene age. Celtis fruits (see symbol) are of Late Pleistocene age. 852 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Lower slopes Valley Basin floor Samples =] 1 m S $ š ° E 8 © g Š š Fle- š - o š dE E E v = > - = m m m io Total pollen (99 FIGURE 11. Pollen diagram, showing selected types and inferred habitat, upper Teewinot lake beds, Jackson Hole, Wyoming (% total pollen). K/Ar age is 9 Ma or late Miocene (from Barnosky, 1984). 1968), and the Split Rock Formation from the est, or whether most of these were restricted to Sweetwater Basin, Natrona County, Wyoming riparian environments cannot be deduced from (Love, 1961). A mid-Miocene (Barstovian) site the available information, though the latter is is from the Saratoga Valley, Carbon County, Wy- more proba oming. The youngest site, the Moonstone For- mation in the Sweetwater Basin, Wyoming, may be of Barstovian or younger age (Robinson, 1971; Love 1961). All are old lake beds near low rolling granite or bedrock hills that probably existed in the Miocene. Pollen is moderately abundant to rare and demonstrates impoverished floras sim- ilar to that evident at the Jackson Hole, Wyo- ming site. Only two types of communities can be conjectured from the data: Implications. The consistent presence of xe- ric and desert-scrub taxa that are sometimes abundant suggests the appearance of steppic vegetation with a diverse herb flora during widely separate times during the Miocene. Grass pollen grains are usually present but never abundant. They are probably associated with depauperate conifer forest or woodland. Several other basin sites in the central and southern Rockies are con- sistent with the modern aspects of these Miocene floras (Meyer, 1986; Leopold & MacGinitie, 1972) The Rocky Mountain data stand in stark con- trast to the Miocene basin sequences from Idaho westward to eastern Washington where pollen and leaf data alike point to rich forest vegetation containing abundant and diverse deciduous ardwoods. Broad-leaved evergreen trees are ap- parently ME in the Neogene of the Rocky Mountain (1) Open basin and lakeside environments in which Artemisia, Sarcobatus and other Cheno- podiineae, Ephedra, and Eriogonum suggest steppic and halophytic environments and in which Salix, Betulaceae, and aquatics imply lake- margin or riparian environments. Terrestrial herbs of the Polemoniaceae, Compositae, Ona- graceae, Umbelliferae, and Polygonaceae may have grown in these basin environments (Ap- pendix VII). (2) Mountain slopes dominated by Pinus with lesser amounts of Picea, Abies, and Juniperus. Whether Juglans, Carya, Quercus, and Ulmus- Megafossil floras from the Great Plains region Zelkova were associated with woodland or for- (Fig. 7) suggest that prairie elements were present HI. GREAT PLAINS LOCALITIES 1987] S q` E z ° = $ ° wawam m+ | o: + + ° < ^ ww mj bos i ae a < o e E ERIOGONUM x + e [5] ARCEUTHOB. ° L ac š pe o T ERIOGONUM É š i POLEMON s ° gi ae la a mm FETI r? mM = O 50 0 30 FIGURE 12. Rock Formation, Wyoming, lower Mi Wyoming, of younger Neogene age (Appendix V, VI). and that grassland developed to various degrees during the Miocene (Elias, 1942; Chaney & Elias, 1938). An excellent summary by Axelrod (1985) portrays a sequence indicating a generally de- creasing Lodge Saga regime from about 16 Ma onw older sites are deciduous hardwood forest se prairie elements (Fig. 13). According to Axelrod, the younger sites suggest woodland or riparian border hardwoods with more exten- sive grassland (Fig. 13) From the Neogene sediments of the upper Arikaree and Ogalalla groups on the Great Plains and High Plains, Elias (1932, 1935, 1942) made systematic fruit/seed and leaf collections at al- most 100 localities from South Dakota to north- ern Texas. The widespread sediment layers with occasional volcanic ashes and vertebrate fossils provided stratigraphy. Elias undertook to define a sequence of fossil seed zones. Modern dating indicates that the main part of this sequence ranges from Hemingfordian through Hemphil- LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 853 —9$—€9— — —e- —e—e- E End =] I] Pl of PE Eos r trr m rr? rn FT T T T | 0 40 o 100 Pollen spectra from four Rocky Mountain sites: Troublesome Formation, Colorado, and Split iocene; Saratoga Valley, Wyoming, mid Miocene; Moonstone Formation, 3? e 1000 SOS ea Sheep * NE eC "I Neb. s> Ee ow e S Kilgore, Neb. iJ al ss” e Eu Beaver Com ~~ Q o Oki. ` rA V Clarendon; anning}, Tex." al Tx. UY N o Annual Precipitation Logan Co., Toda Kan. day HIGH PLAINS 10 5 million years B.P. FIGURE 13. Fossil floras from the Great Plains (High Plains of Axelrod, 1979) suggest a gradual reduction in precipitation during the late Tertiary 854 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 TABLE 5. Geographic affinities of the Kilgore flora, Nebraska (15-12 Ma, mid Miocene), showing the rep- resentation of its species in Miocene floras of the Columbia Plateaus (after MacGinitie, 1962). Modern Distribution Groups Number of Species in Kilgore Number of These Flora by Species oe East of Rocky Mountains only 10 l Mexico and southern Rocky Mountains 7 l Rocky Mountains only 3 0 West of Rocky Mountains 2 0 Split affinities between distinctly eastern and distinctly western species 3 1? Cosmopolitan 1? 1? Eastern Asia 2 2 Total number of species: 27-28 4 6 or ca. 18-21% lian (ca. 18-4 Ma). At most of his sites in the Ogalalla Group (Kilgore and younger age) fruit- ing parts of grasses and prairie herbs were the most common and in most cases the only fossils, although arboreal leaves were present at a few sites (most ofthe latter were summarized by Cha- ney & Elias, 1938, and by Axelrod, 1979, 1985). Elias's identifications, corroborated in large part by the recent work of Thomasson, demon- strate that ancestral Stipeae appear in the earliest Hemingfordian strata or earlier, and that Borag- inaceae (Biorbia, Krynitzka) and Paniceae ap- Hollow Formation) sites. Diversity of prairie taxa increases up section erhaps the most pang feature is the widespread nature of Elias’s fossil fruit and seed zones. In Nebraska the following zones range from Kilgore age to Ash Hollow and younger units (Kimball Formation) of the Ogalalla Group (in stratigraphic order; Elias, 1942): VI. Kimball Formation (calliche). . Biorbia fossilia with Krynitzka coroniformis and Stipidium grande (Ash Hollow For- < mation). Krynitzka coroniformis zone with Stipidium Tune (Valentine Formation). Elias found the same dominant species at the same zones in the Ogalalla at Wray, Yuma Coun- ty, and at many other points in eastern Colorado. He found his Krynitzka zone underlying the Biorbia zone at Wallace County, Kansas (asso- ciated with fossil rhinoceros) and at other sites in Kansas. Biorbia subzones (not described here) were found in Beaver and Ellis counties, Okla- homa. The presence of prairie elements during the Miocene has been documented by Thomasson using scanning electron microscopy to identify eaves, anthoecia (lemmas and paleas of grass LE 6. Kilgore species occurring in Miocene floras of the Columbia Piatssi (from MacGinitie, 1962; TAB Edwards, 1983). Q certain P , but affinity Species Related to Populus washoensis? Pterocarya oregoniana Mahonia MIR C — train P. grandidentata (E U.S.A., SE Canada) P. insignis (E Asia) M. bealii (China) C. mexicana (NW Mexico Ace me A. negundo (W & 7 U.S.A. & S Canada) eas coulteri? F. oregona (W U F. americana (E ^ cent U.S.A., S Canada) F. pennsylvanica (E & central U.S.A., S Canada) 1987] spikelets), fruits, and seeds (Thomasson, 1978a, 1978b, 1979, 1980a, 1980b, 1980c, 1983, 1984, 1985; Voorhies & Thomasson, 1979). For as- signment of fossil grasses, epidermal patterns and silica bodies (phytoliths) were particularly di- agnostic. The plant fossils occurred frequently with vertebrate fossils (rhinoceros, elephantids, horse, and camel). The Ash Hollow Formation (Ogallala Group) in Nebraska, Clarendonian and Hemphillian in age, contains a rich assemblage of prairie plants and aquatics, including repre- sentatives of eight plant families (Chara sp., Characeae; Equisetum sp., Equisetaceae; Pota- mogeton sp., Potamogetonaceae; Carex gracei and two other Carex spp., Cyperocarpus pul- cherrima, and C. terrestris, Cyperaceae; Archae- oleersia nebraskensis, 12 species of Berriochloa, Nassella sp., Oryzopsis sp., Paleoeriocoma hitch- cockii, and Panicum elegans, Gramineae; Bior- bia, Cryptantha spp., Prolappula sp., Boragina- ceae; Celtis willistonii, Ulmaceae; and Polygonum sp., Polygonaceae). In addition, representatives of the Gramineae (species of Berriochloa) and Cyperaceae (Cyperocarpus eliasii) have been re- ported from the Sheep Creek Formation in Ne- braska (late Hemingfordian), and new species of the Gramineae (Berriochloa spp.) and Boragi- naceae (Biorbia sp., Cryptantha spp., and Elia- siana sp.) have been described from the Keller site in Ellis County, Kansas (Hemphillian or Clarendonian). Although the affinities of most of the fossil species are speculative, those of the grasses ap- pear to be rather straightforward. For example, the fossil grasses are classified in tribes Oryzeae, Stipeae, and Paniceae. Archaeoleersia appears to be the forerunner of Leersia (tribe Oryzeae) and is most similar to living Leersia ligularis and L. monandra of North, Central, and South America and to L. triandra of Africa (Thomassan, 1980b). Berriochloa, Nassella, Oryzopsis, and Paleoerio- coma are all classified in Stipeae. Berriochloa shows features that suggest it is ancestral in th evolutionary series: Berriochloa-Piptochaetium- Stipa (sect. Hesperostipa) (Thomasson, 1978a) Paleoeriocoma, likewise, belongs to an evolu- tionary series: Nassella—Oryzopsis—Stipa; this ge- nus is found with Nassella in deposits and ap- pears to be ancestral to species of Oryzopsis (sects. Eriocoma and Oryzopsis) and possibly to some species of Stipa (Thomasson, 1980c). Panicum elegans is most similar to extant species of Di- chanthelium but has not been transferred to that genus due to inadequate sampling of the micro- oO LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 855 morphological characters of the many species of Panicum (Thomasson, 1980b). Tribes Oryzeae and Paniceae, especially, contain grasses of southern affinities; however, the relationships of tribe Stipeae remain uncertain (Barkworth, 1981) even though many extant species have southern affinities. The other families represented in the fossil deposits (Boraginaceae, Cyperaceae, and Ulmaceae) generally are best represented in sub- tropical or warm regions and, predictably, would have southern affinities. he Kilgore flora of Nebraska is central to our data base since the locality occurs at about the same latitude as the Columbia Plateaus floras. It is of mid-Miocene (Barstovian) age and was well documented by MacGinitie (1962), who provid- ed leaf and pollen data. Hence it can be compared with sites of that age on the Columbia Plateaus. The 28 species MacGinitie identified show the strongest affinity (57%) with modern taxa that grow chiefly east of the Rocky Mountains; sec- ondary affinity is with southerly taxa that now occur in Mexico and the southern Rocky Moun- tains (2096). Other relationships are minor and include a few species that occur west of the Rocky Mountains: two in eastern Asia, and one cos- mopolitan group. Importantly, at least three species are intermediate between distinctly east- ern and distinctly western North American taxa: Fraxinus coulteri, Populus gallowayi, and Celtis kansana (Table 5) Only six Kilgore species (21%) occur as fossils in the Miocene floras of the Columbia Plateaus (Table 6). These include a mixture of geographic elements, and small east-west differences may gundo var. negundo, but the Columbia Plateaus species Acer negundoides is morphologically close to A. negundo var. californicum (J. A. Wolfe, pers. comm., 1986). In summary, the floristic relationship of the Kilgore with the Columbia Plateaus floras is slight. I mplications. The mid- Miocene -a (e.g., o those of the present Great Plains grasslands, many with southern affinity. Woody elements (e.g., Kilgore flora) are chiefly related to living taxa of the east- ern U.S. The combined data suggest that the Great Plains flora of the mid Miocene was floristically distinct and had little in common with the Co- lumbia Plateaus Miocene floristic province. The Kilgore flora of the Great Plains bears a stronger relationship to Miocene floras of the eastern sea- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 856 BurmoAM “YOOU LI IdS BYSeIGON ‘FUOOTM opelojoD “AWOSATANOAL VPV TO TIVOSVW eN LI-TI duwems dureAs oureJd sduruado oddajs WUNIPOXD [ ?g 15310} UNIPOXD L ? 18310} uerp1ojduruaH 29 19210] uodo snonpidaq YIM 1SƏ1OJ 19jruoo auejuoJq poo^paeu snonpisaq poompey snonproo(] 9187 /URIAOISIEg UXSCJq9N “MOTIOH HSV Maga AdddVaAL ?ulogepiO “OO HAV *333ID NOSIOd DOANASNATTA AO T Suruo£A “LONIMAAL IIVSV8 AYNdNva G3HOJNVH BW CT1-8 dureAs winipoxn I pue[sse18 2 surejd sduruodo əoddə1s 1sə1oJj snonptoop *]s210J snonproop *]$210J Aa3[[E£A snonpiooq U1IA 1S310J 19j1uOO 2u£]uoJA % 19j1uOO əupluo], 7 1ƏJtuoo əupluo], ueruopuaie[.) Sure[d 18210) s[Iiy0ooJ urejunoJA AyDOY oyep] uo8əi!O u1ojseq *uojgurQuse A, ulo1se4 ‘(2194 UMBIP 10U st parin2o0 Ajqeqoid yorym jjridn jeuordes) a10joq uey} ueruopuoie[) Suunp JoYysIY eYMOWOS oq 0} pouinsoud oe soSuei urejunoui oneulouos -pozi[elideo s1e seiop jo sojdurex3 UNON I} Suunp s[eA191UI SUIT] OM) 10J eJo[[rp102 urejunoJA A20 I} sso1oe sadi UONLIIBIA JULUTWOP JO Sj2O9sue1] js2-]S?] `L d18Vv] 1987] board, e.g., with Calvert Cliffs, than with floras of the Columbia Plateaus (J. A. Wolfe, pers. comm., 1987). Kilgore vegetation was in Axelrod’s (1985) view probably “a wooded grassland with semi-open grassy forests and woodlands on the interfluves as well as patchy grasslands.” The fauna suggests a frost-free climate. This is consistent with MacGinitie’s (1962) analysis. In younger Miocene floras (Clarendonian and Hemphillian, e.g., from Ash Hollow, Nebraska and Beaver County, Oklahoma), seeds of prairie elements are much more abundant and more widespread. Leaf floras suggest that woody vege- tation was confined chiefly to valley-bottom and riparian-border habitats. Together these data suggest “either parklands or grasslands” on the interfluves (Axelrod, 1985; 171). Pliocene floras indicate extensive grassland on the Great Plains, although, as Axelrod and MacGinitie pointed out, the evidence is not adequate to establish that pure grasslands occurred before the postglacial. SYNTHESIS— CONTRASTS EAST AND WEST OF CORDILLERA From the data presented, dominant vegetation types east and west of the cordillera can be com- pared for two times during the Miocene. Table 7 summarizes the general picture for the Barsto- vian/late Hemingfordian (ca. 17-12 Ma) and the Clarendonian (ca. 12-8 Ma). During these two intervals, the Columbia Plateaus west of the Continental Divide consistently maintained de- ciduous hardwood and montane conifer forests, both rich in woody genera, while evidence from the Great Plains east of the Rockies clearly in- dicated the existence of deciduous valley forest with grassland elements. Floristically the eastern floras bear little relation to those of the western area. During both of these times the eastern de- ciduous forest and g ted from forests of the west (1) by the continued existence z the Rocky Mountain massif, which may have ecome more elevated during the Miocene A 1980), and (2) by the continued pres- ence of an impoverished montane conifer forest with steppe elements in the cordilleran area. Through most of the Miocene there appear to have been floristic provinces distinct from each other: (1) The Great Plains province had its primary affinity with species of the eastern U.S.A. and LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 857 was floristically distinct from floras of the Co- lumbia Plateaus. It contained elements that later became important forest species of riparian val- lies (e.g., American elm) as well as extinct genera ancestral to modern grassland elements (Ber- riochloa and Archaeoleersia). (2) The Rocky Mountain province was dom- inated by depauperate conifer forest or woodland with steppe. Except for infrequent pollen of Ju- glandaceae, Ulmaceae, and Tsuga, the spectra resemble modern pollen rain. The presence of Tsuga suggests that the “maritime conifer forest” (of Habeck, 1987) now found near the Canadian border may then have been in a more southerly position. We do not know what species may have been involved, but the impoverished generic list of Pinaceae alone places it in stark contrast with the rich conifer assemblages of the Columbia Pla- teaus region during the Miocene. (3) The Columbia Plateaus province did not take on a relatively modern character on a ge- neric basis until at least late Pliocene (Blancan) time or possibly even Pleistocene time. Although grassland and steppic elements (e.g., Artemisia, Sarcobatus) existed in this bewa d the Miocene, they were unimpor Plain data, grasses became sporadically abun- dant in Pliocene (Blancan) time— fully 10 Ma after grassland may have developed in the Great Plains region. The late appearance of grassland west of the Rocky Mountains may mean that eastern and western grassland provinces are of different origins. Support for this tentative conclusion is derived from the following statement based on data presented earlier in the paper: (1) Great Plains grassland taxa with southern affinities, now considered ancestral forms, ap- peared during the Miocene. (2) The Great Plains floristic province was separated from the Columbia Plateaus during the Miocene by the orographic barrier and by the existence of an impoverished montane conifer forest with steppe on the Rocky Mountain cor- illera. (3) Grasses of northern affinities adapted to summer-dry conditions may have moved into the Columbia Plateaus from the north after the major shift from summer-wet to summer-dry climate about 6 Ma (Hemphillian)— during the demise of the rich Columbia Plateaus hardwood- and certain conifer-forest elements. Floristic evi- 858 dence confirms the strength of these barriers since Barstovian (mid Miocene) time. BIOGEOGRAPHIC IMPLICATIONS The existence of montane conifer forest and steppe on the Rocky Mountain massif through- out the Miocene means that biogeographic con- nections between the deciduous forests of the Columbia Plateaus and those of eastern U.S.A were cut off since at least the early Miocene. Eastern hardwood and swamp elements co- existed on each side of the divide in the Miocene until they were eliminated in the west by the ae deterioration during Pliocene time. mportant shift to a summer-dry climate came etd rapidly to the Columbia Pla- teaus, diminishing a vast reservoir of genetic re- sources; this included many deciduous and co- niferous taxa with affinities to modern taxa of eastern Asia and eastern U.S.A. that show ad- aptations to a summer-wet climatic regime. This basic change reduced the diversity of the mon- tane coniferous forests of the interior in the Pa- cific Northwest except along the Pacific coast. It further established conditions in which steppic elements and grassland of northern distribution probably became established and prominent on the Columbia Plateaus. LITERATURE CITED ARMSTRONG, R. K., . LEEMAN & H. E. MALDE. 1975. K-Ar dating, Quaterna ary and Neogene vol- canic rocks of the Snake River Plain, Idaho. Amer. J. Sci. 275: 225-251. AXELROD, D. I. 1964. The Miocene Trapper Creek flora of southern Idaho. Univ. Calif. Publ. Geol. Sci. Ser : 1-148. 1979. aes Pana its age and origin. Pp. 1-72 in J. n& D er Arid pi pope Res : Arid Lan onf. Pl. Resources. Texas Tech Univ., d Texas. 985. Rise of Es haere biome, central America. Bot. ane —-201. 81. pe A and tax- onomy of North American Stipeae (Gramineae). Syst. Bot. 6: 136-152. BARNOSKy, C. W. 1984. Late Miocene vegetational and climatic variations inferred from a pollen rec- ord in northwest Wyoming. Science 393. 49-5]. Bryson, R. A. & F. K. Hare. 1974. P of North America. Volume 11 in H. E. 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Tertiary dme and pa- ie ec of south-central Wyo Pp. 54-57 Wyoming Geological peices Guidebook, 6th Annual Field Conference, 1951. MCKEE, E. H. 72. Cascadia, the Geologic Evolu- tion of the Pacific Northwest. McGraw-Hill, Inc., New Yor Mack, R. N. & J. N. THOMPSON. 1982. Evolution in steppe with a few large, hooved mammals. Amer Naturalist 119: 757-773. MALDE, H. E. 1982. The Yahoo Clay, a lacustrine unit impounded by the McKinney Basalt in the Snake River Canyon near Bliss, Idaho. In W. Bon- nichsen & R. M. Breckenridge (editors), Cenozoic Geology of Idaho: Idaho Bur. Mines & Geol. Bull. 8. & H. A. Powers. 1962. Upper Cenozoic Stra- bb of pnis = Say Plain, Idaho. Bull. . Soc. Amer. 73: 1197-1220. Y Mn ^H. W 1986. ju J of the Method for Estimating Ako ayapa Ein Tertiary Floras from the Rio Grande R w Mexico and Col- orado. Ph.D. P (Paleontology) Univ. California, Berkeley, Califor NEILSON, R. P. 1986. High- aa oH climatic anal- i d } bi graphy. Science 232: 27- o 1983. Biogeography of to at- L. H. WULLSTEIN. eus southwest American oaks in relation ospheric dynamics. J. Biogeogr. 10: 275. RAS P. 1971. A Geologic Atlas of the Rocky Mountain Region — Tertiary Rocks. Rocky Moun- tain Assoc. Geol. Publ. Rocers, K. L., C. A. REPENNING, R. FORESTER, a LARSON. S. A. HALL, G. R. SMITH, E. ANDE T. J. BRowv. 1985. Middle PB toa: (ee LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 859 Irvingtonian: Nebraskan) climatic changes in south-central Colorado. Natl. Geogr. Res. 1: 535- SHERMAN, J. C. 1947. Precipitation of Eastern Wash- ington. Ph.D. Dissertation. Univ. Washington, Se- attle, Washington SMILEY, C. J. 1963. The Ellensburg flora of Wash- ington. Publ. Univ. Calif. Geol. Sci. 35: 159-276. W. C. REMBER. 1985. Composition of the Miocene Clarkia flora. Pp. 95-112 in C. J. Smiley (editor), Late Cenozoic History of the P Northwest. A. A.A.S. P. n i TAGGART, R. E A Vegetation change in the Miocene Succor Creek flora of Or- egon and Idaho: a case study in paleosuccession. Pp. 185-210 in D. L. Dilcher & T. M. Taylor (editors), Biostratigraphy of Fossil Plants— Successional and Paleoecological Analyses. Dow- den, Hutchinson & Ross, Inc., Stroudsburg, Penn- sylvania . Cross & L. SATCHELL. 1982. Effects of periodic volcanism on Miocene vegetation distri- bution in eastern Oregon and western Idaho. 3rd 54 Supra- basalt: sediments of the Cold Creek Syncline area. In W. C. Myers & S. M. Price (editors), RHO- BW-DO-C-60, Rockwell Hanford Operations, Richland, Washington. TEERI, J. A. & L. G. Stowe. 1976. Climatic patterns and the distribution of C, grasses in North Amer- ica. Oecologia 23: 1-12. THomasson, J. R. 1978a. Epidermal patterns of the mas in some fossil and living grasses and their phylogenetic significance. Science 199: 975-977. 978b bservations on the characteristics of the lemma and palea of the late Cenozoic grass Panicum elegans. Amer. J. Bot. 65: 34-39. 1 Angiosperms from the late Tertiary Keller local fauna of Ellis County, Kansas. Contr. Geol. Univ. W g 17: eee p. (family 198 taceae, Subgenus H. LUE E the late T Hollow Formation of Nebraska. pes 7. 1980b. Archaeoleersia nebraskensis gen. et sp. nov. (Gramineae-Oryzeae), a new fossil grass from the late Tertiary of Nebraska. Amer. J. Bot. i Paleoeriocoma (Gramineae, Stipeae) from the Miocene of Nebraska: taxonomic and phylogenetic significance, Syst. Bot. 5: 233-240. l Ssil grass anthoecia and other plant fossils from hae d burrows in the Miocene of western Nebraska. J. Paleontol. 56: 1011-1017. 1983. Carex graceii sp. n., C Pre elia- sii sp. n., Cyperocarpus terrestris sp. n., Cyper- ocarpus pulcherrima sp. n. (opens from the Miocene of Nebra ska. Amer. J. Bot. 70: 435-449. I . . n u noideae) leaves showing external micromo logical and iier nal anatomical features. Bot. Gaz Epl 145: 2 ————. 1985. pen fossil plants found in Ne- bra ska. Pu Geogr. Soc. Res . 19: 553-564. TRIMBLE, D. OZOIC ieee history of en the u) Plains contrasted with that of the south- 860 ern Rocky Mountains: a synthesis. Mountain Ge- grass anthoecia within Miocene rhinoceros skel- etons: diet in an extinct species. Science 206: 331- 333. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Voss, E. G. 1972. Michigan Flora, Part 1. Cranbrook Inst. Sci., Bloomfield Hills, Michigan. Wo re, J. A. 1969. Neogene floristic and vegetational history of the Pacific Northwest. Madrono 20: 83- l . 1978. A paleobotanical interpretation of Ter- tiary climates in the northern hemisphere. Amer. Sci. 66: 694-703. APPENDIX I. Pollen records showing occurrence of shrubs, terrestrial herbs?, and herbaceous aquatics in Late Cenozoic samples from Idaho, eastern Oregon, and eastern Washington. Numbers show maximum values in pollen/spore counts; + indicates presence. MIOCENE Middle Late E aq £ ` E - _ = © o E x 9 S S = x © 8 E > Š 5 š š G ó Š Y. 3 ER š = Š Š s » £ š ç 9 Š t c st Ze š S z 5 E B ° 5 & E C 2 £ ug ] 9» Š š š š P 5 3 F š P Sio EZ ë 2 O à X &k F & $ m= A Wo Bla 3 SHRUBS Ephedra + 1 cf. nevadensis T + 2 4 + cf. torreyana + + + Inus + + + 5 + + 4 + l l Chenopodiineae + + 1 +? + 7 20 20 3 Sarcobatus + +? + 2 4 3 1 7 l Compositae Artemisia + + + 14 30 52 36 Elaeagnus + + Shepherdia + + + saceae + 3 Cercocarpus + Ilex + + + + 4 Rhus 4 Corylus + l + + rica + + Malvaceae + + + Myrtaceae + Yucca type + Ericales + Vaccinium ? ? + + Caprifoliaceae + Viburnum + Symphoricarpos + Sapindaceae + Mahonia + Cornus + TERRESTRIAL HERBS? Compositae undet. + + 9 11 2 49 Liguliflorae + + l Tubuliflorae 3 6 Ambrosia + + 4 + 1987] APPENDIX I. Continued. LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 861 MIOCENE Middle Mascall' Blue Mountain! Succor Creek? Clarkia? Stinking Water! Miocene, Oregon? Pliocene, Oregon? Trapper Creek? Poison Creek? Jenny Creek‘ Hanford‘ Banbury Basalt‘ Glenns Ferry” PLIOCENE QUATERNARY Bruneau‘ Late Pleistocene, Idaho‘ Cruciferae Acalypha type Onagraceae Polemoniaceae Gilia t Navarretia Caryophyllaceae Stellaria Gramineae Polygonum cf. californicum Eriogonum Anemone Aizoaceae type Pachysandra Umbelliferae Ferns AQUATIC HERBS Myriophyllum +? + 4 + + +? + ++ ++ + ++ + + + De ee ee ee — — - ! Jane Gray in Chaney & Axelrod, 1959. z — & ? Gray, 1985. * Gray, | 1964. Š oc jud u MacGinitie, 1972. * This T enber " Wright, 1985. Cross, 1980; Graham, 1963. 862 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 APPENDIX II. Selected pollen counts, Snake River Plain, Idaho (Fig. 10). Localities given in Leopold & Wright, 1985; the Jenny Creek Formation locality was previously considered as “Salt Lake Formation." Poison Creek Jenny Creek Formation* Formation Banbury Basalt USGS Paleobot. Loc.: D1208-3 D1696 Pinus 84.5 30.2 7.2 Picea 1.0 6.0 Abies cf. grandis 5.4 3.6 A. cf. lasiocarpa 1.0 + Pseudotsuga 1.2 niperus 1.0 4.2 opulus 1.8 Betula 1.8 0.8 Quercus 4.8 us Subtotal: 92.9 53.6 8.0 Cedrus-type 0.5 Carya 0.5 1.8 Juglans 1.8 + Pterocarya 0.6 Ulmus- Zelkova 10.1 0.8 Subtotal Tertiary relicts: 1.0 14.3 0.8 Acer 0.4 Ephedra cf. torreyana 0.6 0.8 Ephedra i R. type) 0.5 Sarcobat 2.0 4.2 0.8 Other Chenopodiincae 6.8 Artem 14.0 Other C aie 9.3 Cercocarpus type 0.6 Elaeagnus + Subtotal shrubs: 2.5 5.4 32.1 udis 0.5 Gramineae 0.6 1.3 Mosecon 1.0 4.8 Cruciferae 0.4 Dicots 2.0 11.9 0.8 Potamogeton 0.6 T C3P3 56.4 Lycopodium + Onagraceae + cf. Parthenocissus + Fern spores 1.0 8.9 1.0 Subtotal NAP: 4.5 26.8 59.9 Total percent 100.9 100.1 100.8 Total tally (202) (336) (236) Cysts 0.5 1.2 Botryococcus 12.0 * Fossil leaves from the Poison Creek Formation identified by J. A. Wolfe (pers. comm., 1987) include Quercus prelobata Condit. and Q. cf. chrysolepis Liebm 1987] LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 863 APPENDIX III. Revised list of the Deschutes flora, northern Oregon (after Chaney, 1938, with additions int. A. Wolfe, pers. comm., 1987; Wolfe includes certain collections made by R. W. Brown from Cha- ney’s floral horizon but a few miles away). Acer negundoides Quercus prelobata us washoensis Populus alexanderi (P. aff. trichocarpa) Prunus irvingii (P. aff. emarginata) Salix sp. (S. cf. xta of Chaney) Salix aff. caudat Ulmus affinis Tod & Wolfe APPENDIX IV. Localities of selected pollen samples from Wyoming and Colorado (Figs. 6, 12). USGS Paleo- botan Formation Locality Site Description Split Rock D1309 Type section: SE '4 NE !⁄4 Sec. 25, T. 29 N, R. 89 W, Natrona Co., Wyoming. Limestone at top of pumicite, from unit 21 of J. D. Love (1961). he from the Split Rock type area are of Hemingfordian age. Split Rock D3319 Upper part of 2,650 ft. EWL 600 ft. SNL Sec. 36, T. 27 N, R. 85 W, Carbon Co., Wyoming. J. D. Love measured section L63-11. Sample B & Ca 6 and 12 ft. Aus base of his unit 3. Sample D/E from his unit 4. Samples ikher in this unit contain abundant reworked Cretaceous pollen Saratoga unit D1540 os Valley, — oil shale. SW !⁄4 NW '% Sec. 26, T. 18 N, R. 84 W, . D. Love locality L59-49. Associated with vertebrates of aiden age ponia to McGrew (1951). Moonstone D1308 Type section; white tuff and white laminated shale. Sec. 17, T. 30 N, R. 89 W, Natrona Co., Wyoming. Samples | and 2 are from Love's (1961) unit 16 and 17, respectively. Troublesome D3473 Rock Creek facies. NW !⁄4 SW 4 Sec. 32, T. 2 N, R. 79 W, Kremmling quadrangle, Grand Co., Colorado. Glen Izett loc. G-63-303 and -304. Sample A is 18 ft. above sample B. Vertebrates from the lower and upper parts of the formation correlate respectively with the lower and upper Hemingfordian Group of Ne- braska (Izett, 1968). Troublesome D1905 SW INE IA Sec. 29, T. 2 N, R. 79 W, Kremmling quadrangle, Grand Co., Colorado. Carbonaceous bed interbedded with silt lake facies containing mid Miocene ver- tebrates. Izett coll. ra 0153- A Troublesome D3493 NE % SE % Sec. 28, T. 3 N, R. 80 W, Gunsight Pass, Twin Peaks, Kremmling quadrangle, Grand i es Izett loc. G-63-305. 864 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 APPENDIX V. Pollen counts, Troublesome Formation, Grand County, Colorado (Figs. 6, 12). Percent pollen and spores. Small counts (shown in parentheses) indicate comparative abundance; + records taxa seen but not in tally USGS Paleobotany Locality: D3493 DINA -b -a Pinus 18.0 72.0 (105) (62) Abies sp. 1.3 1.3 (1) A. cf. lasiocarpa 0.7 A. cf. grandis + (1) Picea sp. 0.7 1.3 (6) (12) P. cf. lal (27) Juniperu. e 32.4 0.7 Ephedra P nevadensis 8.0 2.7 (1) Ephedra cf. torreyana * 0.7 (1) Ephedra (short axis type) 4 *Juglans + 0.7 *Pterocarya 1.3 Carya + 0.3 *Ulmus-Zelkova 4-5 pored 0.7 1.1 (1) 6 pored + Acer + cf. Populus 4 Betula E _0.7 Subtotal trees: 62.4 82.2 Elaeagnus 0.7 Alnus 1.3 3.2 alix 1.3 Sphaeralcea 3 0.7 Caprifoliaceae onicera 0.3 Symphoricarpos 0.3 0.7 Chenopodiaceae 4.6 PA, (3) (1) arcobatus cf. vermiculatus 0.7 (3) Compositae + (1) temisia + 0.7 (3) Xanthium type (1) Gramineae 1.3 1.1 (1) Umbelliferae 7 Polemoniaceae 0.3 Dipsacaceae? + Labiatae 0.3 Arceuthobium 0.3 Apocynaceae + Claytonia type + 0.7 Sparganium 1.1 icots 3.4 2.0 Monocots 16.6 (4) Botrychium type (4) Riccia ty + Trilete spores 7.4 2.7 _ (2) — Total percent 100.9 99.1 Total tally (300) (300) (130) (110) * Genera now exotic to Colorado. 1987] LEOPOLD & DENTON — GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 865 APPENDIX VI. Selected pollen counts, Natrona and Carbon counties, Wyoming (Figs. 6, 12). Percent pollen and spores: + records taxa seen but not in tally. Stratigraphic Unit: Split Rock Saratoga Moonstone USGS Paleobotany D1309 D3319 D1540 D1308 ocality: -B -C -D/E -1 -2 Pinus 8.3 39.5 35.3 51.5 80.0 53.0 72.0 Abies spp. 0.8 cf. concolor 2.6 cf. lasiocarpa 2.0 cf. grandis 0.6 1.3 Picea spp. 0.5 1.2 1.1 cf. engelmannii 3.6 5.2 cf. pungens 7.6 4.6 *Tsuga cf. mertensiana 0.4 cf. heterophylla 0.4 Pseudotsuga cf. menziesii 0.8 1.2 1.1 0.4 Juniperus type 0.4 20.5 20.7 7.4 1.8 Quercus 1.2 1.0 0.4 0.4? Populus + 0.5 0.6 *Carya 1.0 0.4? *Juglans 0.5 0.4 *cf. Liquidambar 1.2 *Ulmus-Zelkova 0.4 1.0 1.6 0.9 12.5 3.1 Betula 0.4 3.2 Salix 0.5 0.4 *Ostrya-Carpinus 0.5 3.2 Subtotal trees: 11.5 65.0 58.4 63.3 95.6 71.4 95.9 Alnus 0.4 0.4 1.1 0.8 Chenopodiineae 5.0 1.0 9.5 4.7 8.5 1.8 Chenopodium type 0.4 Sar t 2.1 6.0 1.8 0.9 7.4 1.2 Ephedra cf. torreyana + 1.2 0.8 0.4 0.4 cf. nevadensis 0.4 1.5 4.1 Compositae 7.5 0.5 1.2 Liguliflorae 0.5 0.6 Artemisia 42.4 1.2 1.1 mbrosia 0.8 0.5 Sapindaceae 0.5 Ribes 1.0 cf. Ptelea 0.4 Arceuthobium 0.4 Caprifoliaceae type 0.6 Eriogonum 0.4 0.4 Gramineae 0.4 1.5 1.8 0.4 0.4 Umbelliferae 0.5 Cruciferae 0.4 Sparganium 1.5 0.6 0.8 Cyperaceae + 0.5 1.8 0.8 Scirpus 2.4 GP; 4.5 Ç, 15.4 2.5 5.1 Monocots 9.0 13.0 6.6 1.1 866 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 APPENDIX VI. Continued. Stratigraphic Unit: Split Rock Saratoga Moonstone USGS Paleobotany D1309 D3319 D1540 D1308 ocality: -B -C -D/E -1 -2 Dicots 3.3 4.0 0.4 5.3 Indeterminate 2.9 4.5 0.6 5.7 3.1 3.2 2.4 Fern spores 1.8 3.0 2.4 7.8 1.1 Total percent 99.6 103.0 98.8 100.4 100.0 100.2 102.9 Total tally (245) (208) (169) (257) (223) (188) (251) Reworked pollen 21 3 28 Botryococcus ++ * Genera now exotic to Wyoming. 1987] LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS APPENDIX VII. Pollen records showing occurrence of shrubs, terrestrial herbs?, and herbaceous aquatics in pollen samples from the Mio-Pliocene of the central Rocky Mountains. Maximum values, % pollen and spores. + indicates presence or less than 1%. O E 4 š - O ° ° S i | mm 8 B 2 2 = 2 v 9 € a 8 d <Š E N E N 2 SHRUBS Ephedra cf. nevadensis 8 4 3 cf. torreyana 1 1 1 l l Alnus 3 + 2 l Sphaeralcea l henopodiaceae 5 10 3 9 Sarcobatus l 6 70 l 7 Compositae ` Artemisia l 42 5 l Elaeagnus l + Caprifoliaceae l onicera Symphoricarpos l ibes l Ptelea type + Yucca type 3 TERRESTRIAL HERBS? Composita Liguliflorae 1 4 ubuliflorae l 8 15 osia 1 + Xanthium type l Polemoniaceae l ilia ty 11 Cruciferae + Claytonia type 1 ace + Polygonaceae "P iog l + Umbelliferae l 1 Apocynaceae + Labiatae + Dipsacaceae + ramineae l 2 15 + 7 3 l HERBACEOUS AQUATICS Cyperaceae 2 + Sparganium l 2 Scirpus 2 + 867 FOSSIL POLLEN OF SABICEA (RUBIACEAE) FROM THE LOWER MIOCENE CULEBRA FORMATION OF PANAMA! ALAN GRAHAM? ABSTRACT Fossil bon of Sabicea (Rubiaceae) has been recovered from the lower Miocene Culebra Formation of Pana well represented in E tropical moist it was part - to mo ada north u lombia. Its association with other m enus is presently widely distributed in vg lees northern South A and premontane wet for erate-altitude insular vegetation p the landscape between merica; it is of Panama. In the lower Miocene embers of the Culebra assemblage h ern act tropical paleoclimates similar to those of the present. The genus has not been reported previously in the fossil record. During studies on Tertiary vegetational his- tory of the Gulf/Caribbean region, pollen and spores are frequently encountered representing genera with no previous fossil record, or whose stratigraphic and/or geographic range is consid- erably extended by the new records. Examples include Pelliceria (Theaceae/Pelliceriaceae; Gra- ham, 1977), Mortoniodendron (Tiliaceae) and Sphaeropteris/Trichipteris (Cyatheaceae; Gra- ham, 1979), Micractinium ee Gra- ham, 1981), and Lisianthiu Gra- ham, 1984). Fossil pollen of Sis (Rubiaceae; Figs. 1-4) has recently been recovered from the lower Miocene Culebra formation of Panama, representing its first known occurrence in the geologic record. ' The author gratefully acknowledges comments provided by John Dwyer and pollen material provided by curators at MO. The research was supported by NSF grants BSR-8500850 and BSR-8619203. > Department of Biological Sciences, Kent State University, Kent, Ohio 44242, U.S.A. * iy ev P $ noe p's SS) PRA eh: ay - FiGuRES 1-6. Fossil and modern pollen of e (Rubiaceae). — 1, 2. Fossil pollen, Pan core 456, slide 1, ESF os D-17.—3, 4. Fossil pollen, Pan pollen of S. colombiana. All taken at 400 x cine : ue 3a, ESF Werl U-31,3.— 5, 6. Modern given in text. Fossil Vg and modern ctua ay reference material are deposited in the eae nd silica. Kent State University, Kent, ANN. Missouni Bor. GARD. 74: 868-870. 1987. 1987] GRAHAM —FOSSIL POLLEN OF SABICEA 869 TABLE l. Modern Rubiaceae pollen examined. Herbar- ium where Voucher Taxon Country Voucher Collection Deposited Amphidaysa ambigua (Standley) Standley Panama Busey 385 MO Coccocypselum guianense (Aublet) Schum. Honduras “ew & Romero 4263 MO C. herbaceum Lam nama D'Arcy & D'Arcy 6731 MO C. lanceolatum (Ruiz & Pavón) Pers. Panama Antonio 1425 MO Didymochlamys connellii N. E. Br Guyana Maguire et al. 32362 MO Gonzalagunia brenesii Standley Costa Rica Croat 2659 MO G. bunchosioides Standley Ferreyra 1660 US G. panamensis (Cav.) Schum. Panama Johnston 76 GH Honduras N. Mex. exch MO G. rosea Standley Panama White 7 GH Hippotis mollis Standley Colombia Lawrence 505 MO H. tubiflora Spr Peru Klug 3084 MO Isertia deamii Sw Guatemala Deam 6016 MO I. haenkeana A. D rvard exc GH I. hypoleuca Benth. Guyana Shell Oil exch Costa Rica Jiménez 4127 MO na Stimson 5062 MO I. pittieri (Standley) Standle Colombia St. George Exped. 337 US Pentagonia brachyotis Cds Standley Panama Dwyer 1385 MO P. macrophylla Benth. Panama STRI exch. MO Panama Croat 4646 MO P. pubescens Standley Panama Croat 4685 MO P. wendlandii Hook. Panama von Wedel 2018 GH Raritebe palicoureoides Wernham subsp. Panama Mori et al. 6617 MO dwyerianum Kirkb Sabicea brasiliensis Wernham Brazil Irwin et al. 24943 MO S. colombiana Wernham Colombia Uribe 3041 US Colombia Gentry et al. 47975 MO S. panamensis Wernham Panama Dwyer MO S. paranensis (Schum.) Wernham eru Schunke V. 10548 MO S. villosa var. adpressa (Wernham) Standley Harvard exch. GH S. villosa Rose & Standley var. villosa Panama Luteyn et al. 1798 MO Panama von Wedel 2889 GH Panama Tyson 3437A MO Schradera blumii Dwyer & Hayden Panama Mori et al. 6625 MO Sommera grandis (Bartlett) Standley Panama Allen 1575 GH Panama Gentry et al. 13581 MO THE COLLECTING LOCALITY In 1958 the Engineering and Construction Bu- reau of the Panama Canal Commission drilled a well through the Culebra Formation in front of Gold Hill on the west side of the Canal at latitude 9°02'N, longitude 79°38'W (Hole No. GH-9). Fifty-seven samples were taken from along the 124-meter core, and 21 contained well-preserved pollen and spores. The specimens of Sabicea were isolated from samples at the 456- and 470.6-foot depths. Other details on the Culebra Formation are provided by Graham et al. (1985) and Stew- art & Stewart (1980). The materials and methods were as described in Graham (1985). DESCRIPTION Pollen oblate, amb oval-triangular to nearly circular; tricolpate/porate (apertures short, slit- like, ca. 2: 1 length:width ratio), 4-6 x 2-3 um, equatorially arranged, meridionally elongated, equidistant, inner margin faintly dentate (due to overlying sculpture elements), faint costae colpi; 870 wall 2-3 um thick, tectate-perforate; finely retic- ulate, muri smooth, width about equal to di- ameter of lumen (ca. 0.5 um); 32-36 um. DISCUSSION Sabicea is a genus of about 125 species of climbing shrubs and vines widely distributed in tropical America and in Africa and Madagascar (Dwyer, 1980a, 1980b). In Panama it is repre- sented by three species: S. panamensis (Guate- mala to Colombia); S. vi/losa (with var. villosa widely distributed in Central and northern South America and with var. adpressa known from Panama, Colombia, and Peru); and S. stellaris (Panama). Sabicea villosa var. adpressa grows on Barro Colorado Island where it is “occasional in older clearings, on trails and at the margin of the forest along the lake; less commonly climbing to the top of the forest canopy and sometimes root- ing in water," and it “is known from tropical moist forest in the Canal Zone, Bocas del Toro, San Blas, Panamá, and Darién, from premontane wet forest in the Canal Zone, and from premon- tane wet forest in Panamá and Darién" (Croat, 1978: 827) Dwyer (1980a: 7) placed Sabicea in the tribe Mussaendeae with Pentagonia, Sommera, Hip- potis, Schradera, Amphidaysa, Gonzalagunia, these (Table 1), and Sabicea can be distinguished on the basis of pollen characters. For example, grains of Raritebe examined were smaller (ca. 25 um), thicker-walled, and tetra- to stephanocol- porate. The pollen of Coccocypselum was con- siderably larger (45-50 um), oblate-spheroidal, and scabrate to finely verrucate. hree species of Sabicea were examined (Ta- ble 1), including all species and varieties reported from Panama except S. stellaris (holotype, MO, not sampled). The pollen showed only minor dif- ferences between the species (e.g., minute vari- ations in wall thickness). The reticulum of the fossil specimens appears slightly more distinct than in the modern pollen examined. The mod- ern forms (Figs. 5, 6) having slightly thicker walls darken more than those with thinner walls during acetolysis and more closely resemble the speci- mens in this respect. These are not inherent mor- phological differences or consistent characteris- tics of the pollen, however, and the specimens cannot be referred to any one modern species. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 The specimens were part of a fossil assemblage that includes bs unos associates (Graham, in prep.; pre ry identifications): Lycopo- dium, HUN person yathea, Pteris, Ly- godium, cf. Antrophyum, Danaea, Gramineae, Palmae, //ex, Chenopodiaceae/Amaran tha aceae, pighiaceae, Hampea/Hibiscus, Eugenia/Myrcia, Rhizophora, Allophyllus, Cupania, Matayba, Sideroxylon, and cf. Guazuma. The landscape of present-day Central America consisted of low- lying volcanic islands at the time the Culebra assemblage was being deposited during the lower Miocene, ca. 25 Ma (e.g., Stehli & Webb, 1985). The association indicates that Sabicea grew in this physiographic setting and under paleocli- mates not greatly different from those ofthe pres- ent. LITERATURE CITED Croat, T. B. 1978. Flora of Barro tbe Island. i iforn n Flora of DE GRAHAM, A. . New records of Pelliceria (Thea- ceae/Pelliceriaceae) a the Tertiary of the Carib- bean. Biotropica -52. orton UU (Tiliaceae) and Sphaeropteris/Trichipteris (Cyatheaceae) in Ce- nozoic deposits of the Gulf-Caribbean region. Ann. ———. 1985. Studies in neotropical paleobotany. IV. The Eocene communities of Panama. Ann. Mis- 72: 504-534 souri Bot. Gard. ; TEWART. 1985. Stud- bearing sediments. Ann. Missouri Bot. Gard. 485-503. SrEHLI, F. G. & S. D. Wess (editors). 1985. Great American Biotic Interchange. Plenum, New York. STEWART, R. H. & J. L. STEWART (with the collabo- ration of W. P. Woodring). 1980. Geologic Map of the Panama Canal and Vicinity, Republic of Panama. Scale: 1 : 100,000. U.S. Geol. Surv. Misc. Invest. Map I-1232 SYSTEMATICS OF THE AMPHI-ATLANTIC BAMBUSOID GENUS STREPTOGYNA (POACEAE)! THOMAS R. SODERSTROM? AND EMMET J. JUDZIEWICZ? ABSTRACT Streptogyna i is the only herbaceous bamboo genus with an amphi-Atlantic distribution. Streptogyna Sri Lanka, and India, and S. americana is found throughout the suggest segregation at the generic level, they are here retained in a single genus because they are quite similar in spikelet morphology and leaf anatomy. Multicellular microhairs are present on the lodicules primitive feature in the family as they are com ffinities in its ligule and leaf anatomy, spikelet structure, caryop- demb ut differs from the core group ofthe subfamily in its seedling sal i S and lack of epidermal so "oq en in the two species suggest of it 1 that neither could have been derived directly from the oth The grass genus Streptogyna was first brought to the attention of Western botanists in the late 18th century, when British and Swedish collec- tors brought back specimens of “rat-catching grass" from the forests of West Africa. A gath- ering from Nigeria by Palisot de Beauvois was used as the basis for Streptogyna in 1812, which he based on the only species known to him, S. crinita. The narrow leaves and many-flowered spikelets of Streptogyna were long taken as in- Thus, Bentham (1883), Hackel (1887), a ar (1936) all considered that the proper disposition of this genus from the tropical rainforest lay with this large, temperate-region grass group. But there were dissenters, and Nees von Esenbeck (1835) and Steudel (1855), for example, suspected the bambusoid affinities of the genus. Streptogyna was briefly revised by Hubbard (1956), who in- dicated that the group deserved tribal status, but it was not until Tateoka (1958a) and Metcalfe (1960) examined its leaf anatomy that the bam- busoid affinities of Streptogyna became clear. Recent workers agree that Streptogyna should be placed in its own tribe in the Bambusoideae (Calderón & Soderstrom, 1980; Clayton & Ren- voize, 1986). In a treatment of the herbaceous bamboos of Sri Lanka, Soderstrom et al. (1987) offered a detailed descriptive account of the leaf- blade anatomy in the two taxa. The present study provides a revision of the genus and attempts to € the relationships of the two species by examining characters that have not yet been es in detail, such as morphology of lodi- cules, starch grains, and embryos. MATERIALS AND METHODS Specimens of Streptogyna were examined from the following herbaria: AAU, B, CAY, CEPEC, F, G, ISC, K, M, MO, NA, NY, P, PDA, RB, S, US, W, and WIS. For anatomical studies, spikelets, leaves, and embryos (Table 1) were dehydrated in dimethoxypropane, infiltrated with tertiary butanol, embedded in wax, sectioned us- ing a rotary microtome, and stained in chlorazol black E. Lodicules were rehydrated with Aerosol- OT before examination. Starch grains from cary- opses were cut on a freezing microtome and stained with I,KI. Observations of living plants of Streptogyna were made by Soderstrom in Bra- zil (March 1972, and May 1976) and by Jud- ! We are grateful to Alice R. Tangerini for the illustrations, to Stanley Yankowski for the floret sections, to C the curators of the herbaria that lent us specimens for study, an and to Gerrit Davidse, Lynn G. Clark, and Ma Sangrey. Part of the research Ris to this paper was performed while the second author was a predoctoral fellow at the Smithsonian Institutio ? Department of Botany, nds Institution, Washington, D.C. 20560, U.S.A. Died on September 1, 987. ? Department of Botany, Smithsonian Institution, Washington, D.C. 20560, U.S.A. ANN. MISSOURI Bor. GARD. 74: 871-888. 1987. 872 TABLE 1. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Specimens of Streptogyna for which D (e), floret bases (using the scanning electron micro- scope) (f), lodicules (1), and starch grains (s) were examine S. crinita CAMEROON: Buesgen 530 (1) (US). GABON: without collector, Limbareni, May 1875 (1) (US). GUINEA-BISSAU: Espirito Santo Expedition 3735 (1) (US). INDIA: Wight 2362 (1, s) (PDA). Ivory Coast: Bamps 2175 (e, s) (BR). LiBERIA: Baldwin 6305 (1) (US). SIERRA LEONE: Fairchild s.n., Jan. 1927 (e, 1, s) (US). SRI LANKA: without collector, 3 Jan. 1881, Henaratgoda (1) ( PDA), Senaratna 2700 (1 ) (PDA); Gardner s.n., Dec. 1846 (1) (PDA). ZAIRE: Gilbert 14213 (s) (BR); Lebrun 605 (e, 1, s) (P); Mullenders 1226 (s) (BR); Vanderyst 993 (1) (P). S. americana BRAZIL: Eiten & Eiten 8903 (1) (US); Prance et al. 6454 (l, e) (US); Prance et al. 6527 (1, s) (US); Swallen 5089 (1, s) (US); Soderstrom 2193 (e, 1, s) (US). COLOMBIA: Blydenstein 1687 (1) (US); Idrobo & Schultes 608 (1, s) (US). FRENCH Guiana: Broadway 77 1 (1) (US). GUATEMALA : Weatherwax 104 (e, s) (US). SURINAM: Maguire 54093 (1) (US). TRINIDAD: Hitchcock 10122 (1) (US). VENEZUELA: Steyermark 86760 (1) (US). ziewicz in Panama (March 1983). A Cambridge 35 scanning electron microscope was used to ex- amine the lemmas of Streptogyna species. RESULTS Vegetative morphology. Streptogyna species are herbs of the rainforest understory and are less than one meter in height. Streptogyna crinita has long sympodial rhizomes that are densely covered with striate scale leaves, and the erect culms are leafy, bearing lanceolate leaves along most of their length (see figure in Soderstrom et al., 1987). Streptogyna americana is densely ces- pitose with the culms produced from short, non- scaly, sympodial rhizomes, and the linear leaves are all borne near the base of the plant (Fig. 6A). The leaves of S. crinita have glabrous sheaths while those of S. americana are hispid near the summit (Fig. 6B). At the summit of the sheath are borne membranous flanges that have been called eal appendages (Tran Van Nam, 1972), as they appear to arise from a meristem inde- pendent of the sheath. These and the oral setae are moderately well developed in S. crinita but inconspicuous or absent in S. americana. Both species also have an indurate abaxial rim at the summit of the sheath (external ligule), and the leaf blades are deciduous aboye this structure. A entin both taxa. The narrow leaf blades of S. amemus are strictly glabrous, whereas the broader blades of S. crinita may have a scattering of long, delicate short E > ie] = ° = Ë = [2 o 3 E = oO £e e. £5 x E oO E e. oO = panicle of Sirepiasvna species Is spikelike: but in both species individual plants have been seen in which an additional floriferous branch is borne at the base of the main axis of the inflorescence. The inflorescence of S. crinita 1s generally shorter than that of S. amer- icana and appears to be more densely flowered; this appearance is due in part to the larger glumes in S. crinita. Both species have loosely several- flowered spikelets (Fig. 6D) with the uppermost florets successively smaller and sterile and with a peculiar downward prolongation of the base of each floret, which, however, does not appear to be an elaisome, as no oil was detected within it. The glumes of S. crinita are broad, elliptic, and have many nerves, while those of S. americana have few nerves and are linear to lanceolate (Fig. 6E, F). The florets of both species are indurate and fall attached to a pointed, persistent rachilla internode (Fig. 6G) that presumably aids in ex- ternal animal dispersal. In S. crinita the base of the floret is pilose and in the scanning electron microscope (SEM) the bulk of the epidermis, which appears granular through the light micro- scope, is composed of short prickles alternating with square to rectangular cells in a sharply de- fined pattern (Fig. la, b). The rectangular cells were observed to deflate when placed under the electron beam of the SEM. The lemmas of SS. americana are glabrous and the epidermis is ap- parently covered by a thick cuticle that obscures the rounded p and intercalated rectangular cells (Fig. 1c, d). The lemmas of both species have long, antrorsely scabrous awns. The paleas are strongly bikeeled (Fig. 2a). Flowers. The flowers of both species have three relatively large lodicules (Fig. 3). Those of S. crinita are spatulate, of firm texture throughout, and are strongly nerved until near the summit (Fig. 3A). Microscopically the epidermis of the upper portion of each lodicule consists of a uni- form network of polygonal cells with moderately on Sierra Leone, 2 Idrob Soderstrom et al. 2193 (US). Scale bar in a = thickened walls (Fig. 3B). The apices of the lod- icules are fringed with about 5-15 thin-walled microhairs, and each hair contains 4-8 cylindri- cal cells. The basal cell is slightly longer than the apical cells and does not collapse when dried as do the apical cells. The longer lodicules of S. americana are narrowly lanceolate or linear and SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA I : as 3 (US); o & Schultes 608 (US); e based on Sierra Leone, Afzelius and wasiwi. s.n. (BM); f based on Brazil, 00 u f.) m for a, c; 20 um for b, d- often are widest just below the middle (Fig. 3C). In their lower portions they are strongly nerved and of firm texture, but at about the middle of their length the vascular bundles end and the lodicule tapers gradually to a very delicate, hya- line apex. Most specimens have the apices and uppermost margins of the lodicules fringed with 874 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 *005 e s A (YYY °) \ " p v % ote T M o 5 FiGURE 2. Floret anatomy of Streptogyna americana.—a. Transverse section through lower part showing lemma, palea, three lodicules (1), two stamens (s), and gynoecium (gy) with three stigmatic vascular bundles 1987] EA i iy "ull EEF iiid HE H ul FiGure 3. Lodicules of Streptogyna crinita (A, B) a apex, showing multicellular microhairs Lodicules. — D. Detail of apex short cells interspersed with long cells on surface. (Streptogyna crinita based SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA — B. Detail of — A. Lodicules. and undifferentiated, thick-walled parenchyma cells on surface. — C. —E. Detail of upper portion of lodicule, showing prickles (mainly marginal) and nd S. americana (C—E). meroon, Buesgen 530 (US); ) S. americana based on Trinidad, Hitchcock 10122 (US). Scale bar = 1 mm for A, C; 0.2 mm for B, D, E abundant prickles that occasionally grade into short cilia. The epidermis of the upper portion consists of alternating, thin-walled long and short cells, some of the latter modified into prickles americana. The two stamens of both species are lateral anterior in position and are free to their bases (Figure 2a); the anthers are linear (Fig. 6M). The gynecium of Streptogyna crinita bears a sin- gle style branching into two stigmas, and the ovary is pilose near the summit. The stigmas, which are supplied by each of two lateral posterior vas- cular bundles in the ovary, are unusual in that they continue to grow after anthesis, elongating and producing stout retrorse barbs on their adax- ial surfaces. Streptogyna americana has three stigmas (Fig. 6L) produced from three en bundles within the glabrous ovary (Fig. 2a). T i grow after ee g short, papillate pro- cesses from a meristematic layer near the adaxial surface (Figs. 2b, 6N Fruit. Both species have a cylindrical, linear caryopsis with a linear hilum that extends the full length of the fruit (Fig. 60, P). The mature endosperm starch granules of S. crinita are weak- ly to moderately coherent into masses of 3-6 that form individual grains 6-10 (rarely 15) um in diameter (Fig. 1e). The starch grains of S. amer- icana are nearly round, 10—30 um in diameter, and are simple or rarely compounded into small masses (Fig. 1f). There is a prominent lacuna in the center of many of the grains. The embryo of Streptogyna species is small, basal, and when dissected out of the caryopsis is observed to be about one-half again as tall as it is wide. In me- dian sagittal sections the embryos of S. ameri- cana (Fig. 4b) sampled had a prominent epiblast, no mesocotyledonary internode, and a small cleft between the scutellum and coleorhiza. In trans- verse sections the coleoptile had two lateral nerves and fused margins, the first embryonic leaf had five nerves and strongly overlapping margins, and the scutellum had three vascular traces (Fig. 4d). Streptogyna crinita had a similar embryonic d. — b. Transverse section through upper part showing three stigmas with papillae arising from meriste- ws). matic regions on adaxial surfaces. (Based on Brazil, Soderstrom et al. 2193 (US). Scale bar: a — 25 um.) 100 um; b = 876 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 transverse section through plumule show verlapping margins of first embryonic leaf; plumule is attached to scutellum on bottom. (a based on Zaire, Lae 605 (US); b, d based on Brazil, i a eae et al. 2193 (US); c 1987] SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA TABLE 2. Comparison of Streptogyna crinita with S. americana. Character S. crinita S. americana Distribution Tropical Africa, Sri Lanka, India Tropical America H Rhizomatous; leaves spaced along Cespitose from short, knotty rhi- Ligular area Morphology of leaf blade Upper surface of leaf blade Inflorescence Glumes Lemma Lodicules Ovary Stigmas Starch grains Embryo Sheath summit glabrous; sheath au- ricles and oral setae conspicuous en x 29(—40) cm long, 1- 2.5c Nerves say, raised; glabrous or aringly l j cm lappin Large; first glume 6-8 mm long, the second 17-26 mm long, 11-17- id ne spikelets over- nerve Base pilose; texture granular, the epidermal cells distinct in SE Oblanceolate, 1.5-3.5 mm long; multicellular microhairs present; prickles rare or absent; short cells absent; texture near apex firm Pilose near s 2, retrorsely a rbed ompound, individual granules 6— va s 5) um in diameter Cleft between scutellum and coleo- zomes; leaves clustered at plant base Sheath summit hispid; sheath auricles and oral setae inconspicuous Linear, 50-78 cm long, 0.8-1.6(-2.4) cm wi e Nerves not raised; glabrous 25—40(—67) cm long, the spikelets not overlapping Small; first glume 3-12 mm long, the seco EE 12-16 mm long, 7-9- en pom texture smooth, the rmal cells obscure in SEM m long; a valine i absent; pri dant; short cells sain Ld near apex hyaline Glabrous 3, with soft, short hairs Simple, grains 10—30 um in diameter Cleft between scutellum and coleo- rhiza not conspicuous Seedlin g Unknown Chromosome number = 12 rhiza distinct First leaf linear, erect Unknown structure except that the cleft between the scu- tellum and coleorhiza was less conspicuous, often present merely as an embayment on the lowe side of the embryo (Fig. 4a). DISCUSSION Despite the statement of Jacques-Félix (1962) that there is little difference between the species, the Afro-Asian Streptogyna crinita and Ameri- can S. americana are quite distinct from each other in a number of characters (Table 2). The long, scaly, flagelliform sympodial rhizomes of S. crinita are rare in herbaceous bamboos, a sim- ilar type being found only in some olyroids such as Pariana parvispica. The divergence in lodicule morphology (Fig. 3) between the species is great and in most grass taxa would imply separation at the genus level. Microhairs are not found on the foliage of either species (Tateoka, 1958a; Metcalfe, 1960; Jacques-Félix, 1962; Renvoize, 1985; Soderstrom et al., 1987), but this study. has bau those found on the foliage (Metcalfe, 1960; Jacques-Félix, 1962) and lodicules (pers. Obs.) of the African herbaceous bamboo genus Guaduella (Guaduellae). Multicellular micro- hairs are also found on the foliage (Smithson, 1956) and floral bracts (pers. obs.) of species of Joinvillea (Joinvilleaceae), a group often consid- ered to be among the closest relatives ofthe grass- es (Stebbins, 1982; Dahlgren et al., 1985; Camp- bell & Kellogg, in press). Multicellular microhairs, consisting of three or more thin-walled cells, and with a blunt distal cell, have also been noted in the olyroid bamboos Maclurolyra tecta (Calde- rón & Soderstrom, 1973), Diandrolyra tatianae ed on Ivory Coast, Bamps 2175 (BR). Abbreviations: io coleoptile; cr, coleorhiza; ep, epiblast; If, first um.) embryonic leaf; ra, radicle; sc, scutellum. Scale bar: a, b = um; c, d = 878 (pers. obs.), the woody bamboo Arundinaria va- gans (Metcalfe, 1960), as well as in several other woody bamboo genera such as Pleioblastus and in several species of the arundinoid genus Dan- thonia (Tateoka & Takagi, 1967). The presence of microhairs on the lodicules but not on the foliage of S. crinita provides an exception to the generalization of these authors that only grasses with microhairs on their leaves also exhibit them on their lodicules. Tateoka & Takagi also illus- trated a typical woody bamboo (Sasa species) lodicule with cilia, prickles, short and long cells, and bicellular microhairs that in many ways rep- resents a composite of the features found in the two Streptogyna species. The lodicules of S. americana possess abundant marginal prickle hairs (which occasionally grade into short cilia) and have surfaces that are differentiated into long and short cells in the upper portions. In S. crinita, prickles are rare and cilia and short cells are ab- sent on the lodicules, but microhairs are present. The two species differ in stigma number and morphology. Streptogyna americana possesses three merely papillose stigmas, whereas S. crinita has two stigmas that are coarsely armed with retrorse barbs. However, in both taxa the stigmas elongate after anthesis, becoming entangled with each other and with the lemma awns. Reports of three stamens in at least some florets of both Streptogyna species (Doell, 1880; Bentham, 1883; Jacques-Félix, 1962) could not be confirmed in this stu Streptogyna crinita consistently has small, weakly to moderately compound starch grains (Fig. le). This agrees with the descriptions of Jacques-Félix (1962) and also Yakovlev (1950), neither of whom cite specimens. The starch grains of S. americana are larger and essentially simple (Fig. 1f). Starch grain characters are not as reli- able as anatomical or cytological data, but they have been considered to be taxonomically useful at the tribal level (Tateoka, 1962). Most bam- busoids, as well as Joinvillea (pers. obs.), possess compound starch grains. The two species of Streptogyna are similar in embryo structure (Fig. 4). Both possess typically bambusoid embryos with a formula of F+ PP (see Reeder, 1962); that is, no internode is pres- ent between the divergence of the scutellar and coleoptilar vascular traces (F), an epiblast is pres- ent (+), a cleft between the lower part of the scutellum and the coleoptile is present (P), and the margins of the first embryonic leaf overlap (P). Jacques-Félix (1962) illustrated an embryo ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 of S. crinita with no internode between the scu- tellar traces (F), an epiblast present (+), and lack- ing a cleft between the scutellum and coleorhiza (F), but in all individual embryos of the speci- mens that we examined a small embayment was present at the base of the embryo that could be interpreted as a cleft, although it was not as prominent as in S. americana. In transverse sec- tions the embryos of both species had a first em- bryonic leaf with strongly overlapping margins (P). The embryo of both species is unusual among the Bambusoideae examined in that it is dis- tinctly taller than it is wide. CONCLUSIONS Many grass genera occur in both the Old and the New World, but only very few tropical-forest Poaceae have this type of bihemispheric distri- bution pattern. Among the woody bamboos, Arundinaria and Bambusa (both Bambuseae), taken in their widest sense, have been considered to range across the Atlantic Ocean. However, studies of these groups by Soderstrom, Ellis, and collaborators are revealing them to be hetero- geneous assemblages, and it appears Bambusa cannot be considered to exhibit a true amphi- Atlantic distribution. Among the herbaceous bamboos, only Olyra and Streptogyna occur both in South America and in the Old World, but Olyra latifolia is a widespread, weedy species that was almost certainly introduced by humans into Africa from South America quite early. Therefore the only bamboo genera with amphi- Atlantic distributions are Streptogyna and pos- sibly Arundinaria. Streptogyna differs from the bambusoid core group (Soderstrom & Ellis, in press) principally in details of seedling morphology (the first ex- panded blade of S. americana is narrow and ver- tical, not broad and horizontal as in all other bambusoids; Soderstrom, 1981) and lack of epi- dermal papillae, microhairs, and well-developed arm cells. Of these divergences, the distinctive seedling morphology is perhaps the most signif- icant (see Kuwabara, 1960), but the seedling of S. crinita has not yet been examined. Strepto- gyna resembles the woody bamboos in possess- ing many-flowered spikelets, strongly developed fusoid cells, an adaxially projecting leaf blade midrib with complex vasculature, multicellular microhairs (as noted, present in Guaduella and several woody bambusoid genera), lateral ap- pendages (Tran Van Nam, 1972), oral setae, and 1987] especially an external ligule and deciduous leaf blades. It may be noted that the multicellular microhairs of Guaduella species occur on bot the leaf blades (throughout) and lodicules (on apices only) and that the basal cell is very long, about half the total length of the hair. Strepto- gyna crinita, in contrast, has uniseriate micro- hairs in which the basal cells are not conspicu- ously longer than the upper cells. Cytologically, Streptogyna crinita is similar to the — uq and has a basic chromosome number of n = 12, as reported by Veyret (1958), REA (BAS 1965), Kammacher et al. (1973), and Dujardin (1978), who studied material from Sri Lanka, Uganda, the Ivory Coast, and Zaire, respectively. Streptogyna americana has not been studied cy- tologically. The generalization that microcharacters in the grass family are often more useful in generic, tribal, and subfamilial delimitations than are spikelet phology be- cause microcharacters are under less intense se- lective pressure thus finds an exception in the case of Streptogyna. In this genus, floret mor- phology, with its special adaptations to external animal dispersal (Ridley, 1930; van der Pijl, 1982; Soderstrom et al., 1987) 1s clearly a conservative feature. The leaf anatomy of the two species is quite similar, and one of the only consistent dif- ferences between the two is the presence of more prominent adaxial ribs over the veins in S. c nita (Soderstrom et al., 1987) and the ape presence of adaxial ciliate macrohairs in the lat- ter species. We have shown that in the habit, lemma tex- ture, starch grain morphology, and especially the lodicule structure the two Streptogyna species are quite different from each other, to the extent that segregation at the generic level might be seriously considered. Based on a comparison with Join- are probably derived from three, which S icana retains. It seems likely that both species evolved from an extinct ancestor and that their separation is ancient. However, barring strong differences in leaf anatomy and cytology, grasses are traditionally segregated into genera on the basis of gross spikelet morphology, and the spikelet and floret structures of the two species of Streptogyna are quite similar, as is the leaf SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA 879 anatomy. It will be necessary to study the seed- lings of S. crinita and the cytology of S. ameri- cana before a final decision can be made on the taxonomic level at which these two sibling species should be recognized. TAXONOMIC TREATMENT Streptogyna Palisot de Beauvois, Essai Agrost. 80. 1812. TYPE SPECIES: S. crinita P. Beauv. Streptia Rich. ex Doell in Mart., Fl. Bras. 2(3): 171. 1880. ; Nomen nudum Perennial forest grasses; culms solid, un- branched above the base; leaf sheaths strongly ribbed, extending upward along both sides of the pseudopetiole and contiguous with the inner lig- ule; outer ligule present as a short, indurate rim; inner ligule short, membranous; lateral append- ages and oral setae present at summit of leaf sheath; leaf blades deciduous, linear to lanceo- late, narrowed below into a short pseudopetiole, the nerves parallel or very slightly oblique from the midvein at its base; midvein and prima nerves manifest only on the abaxial (lower) sur- face, the secondary lateral nerves and transverse veinlets inconspicuous; leaf margins antrorsely scabrous. Inflorescence pedunculate, a spikelike panicle, unbranched or occasionally with a spike- like branch at the base; rachis 3-angled, one side convex and other sides concave and alter- ately bearing the spikelet pedicels. T Bos pedi ET greenish, several-flowered, the lower florets well developed, Hio wet, laterally compressed, hermaphrodite, the upper ones pro- gressively smaller and sterile, disarticulation oc- curring between the fertile florets, each of these falling attached to the extended curved rachilla segment above it; glumes 2, membranous, per- sistent, many-nerved, the first shorter than the second and attached to the side of the thickened 2-keeled, sulcate between the keels; elongate, strongly nerved; stamens 2, lateral an- terior, the anthers basifixed; ovary with a long style; stigmas 2 or 3, becoming hardened and persistent, intertwined at maturity with the stig- mas of other florets in the same spikelet and 880 inflorescence; fruit a linear caryopsis, the hilum narrow, extending nearly the entire length of fruit, the embryo small, basal. Basic chromosome number, n = 12. KEY TO THE SPECIES OF STREPTOGYNA la. Plant with long, scaly rhizomes; stigmas 2, retrorsely barbed; second glume 17-26 mm long; base of lemma pilose; leaf blades 10- 25(-40) mm wide; — ERE Stre ii iin crinita lb. Plant cespitose from short, Seid dip es; stigmas 3, subglabrous; second glum 12-16 mm long; base of lemma slabrous leaf blades 8-16(-24) mm wide; UE pical ue [doppie americana 1. Streptogyna crinita P. Beauv., Essai Agrost. 80 + plate 16. 1812. TYPE: Nigeria: [“ prob- ably gathered in the forests of Oware or Be- nin" (Hubbard, 1956)] anno 1786-1788, Palisot de Beauvois s.n. (holotype, P, not seen). Streptia crinita Rich., a herbarium name given as a nonym of Streptogyna crinita by Doell in Mart., Fl. Bras. 2(3): 172. 1880. Streptia secunda Rich., a herbarium name given as a synonym of “yasa qapya crinita by Doell in Mart., Fl. Bras. 2(3): 172. 1880. — e es Hook. f. in Trimen, Spars Fl. n: 301—302. 1900. TYPE: Sri Lanka, w out ii or collector, C. P. [Ceylon Plants] 922 (holot 1). Culms 55-100 cm tall, each representing the aerial extension of an upturned sympodial rhi- zome, the culm itself producing at its base 1-3 additional rhizomes to 25 cm long, these with short internodes about 1 cm long covered by the overlapping sheaths; sheaths bladeless, ovate- lanceolate, strongly striate, 7-10 mm long. Leaves evenly distributed on culm, not overlapping; sheaths finely ciliate on the margins, otherwise glabrous; outer ligule 0.4—0.9 mm long, tipped by a fringe of ciliate hairs ca. 1 mm long; inner ligule 1-2 mm long, the upper margin erose, cil- iate on the abaxial (outer) surface, glabrous on the adaxial surface; sheath auricles 1-2 mm long; lateral appendages 0.3-1.2 mm long; oral setae sparse, delicate, less than 1 mm long; pseudo- petioles 5-15(-25) mm long; leaf blades narrowly to broadly lanceolate, 18-29(-40) cm long, 1- 2.5(-4) cm wide, acute at the tip, c) below to the pseudopetiole, primary lateral veins 4-8 on each side of the midrib; upper blade surface pale green, glabrous or occasionally with scat- tered spinelike hairs; lower surface lighter green ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 than the upper surface, glabrous. Inflorescence 11-29 cm long, borne on a peduncle 2-10(-40) cm long, erect at first, becoming strongly pendent with age; spikelets (11—)20—30(-42), the glumes of adjacent spikelets strongly overlapping; rachis appressed-pubescent. Spikelets on pedicels 0.8— 1.7 mm long, 4—5(-7)-flowered; first glume 6.5- 10 mm long, linear-lanceolate, glabrous, 1—3- nerved, transversely veined; second glume (16-)20-23 mm long, 1.7-2.5 mm wide, ellip- tic-lanceolate, glabrous, 1 1-17-nerved, the nerves of various thicknesses, not all extending the length of the glume, with numerous transverse veinlets, the terminal awn up to 2 mm long, or occasion- ally the apex emarginate; lemma of lowest floret 18-25 mm long, lanceolate, indurate except for the broad, scarious margins, 7(—9)-nerved, the callus 1.5-2.5 mm long, pilose, the terminal awn (12-)15-26 mm long; rachilla internode attached to base of lowest floret 3-4 mm long; palea about as long as the lemma, not conspicuously pro- truding from it; lodicules firmly membranous, narrowly obovate or spatulate, with a fringe of multicellular microhairs at the apex (rarely gla- brous), the anterior pair 1.5-3.5 mm long, 1-5- nerved, the posterior pair often slightly longer, narrowly oblanceolate, 1—3-nerved; filaments weak and ribbonlike, the anthers pale yellow, about 4 mm long; ovary fusiform, long-ciliate on the upper third and lower part of the style; style long, flattened, with sparse appressed hairs; stig- mas 2, strongly retrorsely barbed above, the barbed portions elongating and coiling with age; caryopsis 12-16 mm long, 0.8-1.2 mm wide, slightly tapering to a persistent, ciliate beak at its summit. Additional d die examined. ANGOLA. CA- BINDA (MAIO OMBE o Zau "Gossweiler 6557 (BM, K). BENIN: Djou ` Chevalier 23891 (P.). CAMEROON: Without aa ras anno 1908-1909, Buesgen 530 (US); Mbamkin, Letouzey 2652 (BR); Dimako, rive ine de la riviére Mbonda, Letouzey 2682 (BR); 38 m ESE Djoum, pres Akoafim, Letouzey 8400 (B, K); x de Yokadouma, Meijer 15146 (MO); N'Kolbrisson, ca. 8 km W of Yaoundé, de Wilde 1205 (B, BR, K, MO); Yaoundé, Zenker 533 (U , Chevalier 10552 (P); oa Gounda- St. Floris National Park, 8 km S of Cam Koumbala at confluence of Mbingou and Koumbala, 8°26'N, 21?15'E, Fay 4087 (K); Mbaiki, le Testu 3446 (BM), Tisserant 3446 (K). CONGo: Brazaville a St. Jo- seph, Chevalier 27341 (K). EQUATORIAL GUINEA. CAMPO DISTRICT: Bebady, route from Anio, Tessmann 658 (K). NDO Po: anno 1859, Mann 108 (K, W). ETHIOPIA. ILLUBABOR: E of Abobo, 7?48'N, 34°37'E, Chaffey 908 (K). GABON: without locality, Griffon du Bellay s.n. (K); 1987] Limbareni, May 1875, collector’s name wee. (US); Tchibanga, /e . TEN M). GHANA: 6 chs. from Akudum, Ankra 5 (US); Amuni, Chipp 53 (K); Ashanti, C ummins r 164; Pra Suhien E Reserve, Deaw 363 (F, MO); Kade, Agricultural Research Sta- tion, Enti s.n., GC-42033 (MO), Ankrah s.n., Ghana Herbarium No. 20190 (K); Sekodumasi, Kitson 8 (BM); E of Anyaboni at S edge of Afram Plains, Morton 6095 (K); Dawo Mato Kola, Thomas D-28 (K); Akatri, Tho- mas D-90 (K); Atuna, Vigne 3522 (BR). GUINEA: Ba- fing, Adam 13795 (MO); 50 km de Kindia vers Mansu, Roberty 10682 (G); Télimélé nord, Roberty 10775 (G). GUINEA-BISSAU: entre Sedengal e Ingone, Espirito San- to Explorações Botánicas 3735 (US); Bedanda, Pereira & Correia Eo K). INDIA: without exact locality, Wight ag (G DA). KERALA: South Travancore, 2,000 yid P (BM, K); Vetillapara forest, Cochin, jmd 12 (K); Courtallum, Tinnevelly District, n 1353 (K, W). Ivory Coast: Um Adam 8 (MO); Duékoué-Buyo, fôret N du o, Bamps ad (BR); near Issia, Daloa-Abidjan uL poA 13609 (K); Bingerville, = hevalier 16036 (P); Amatioré Forest Reserve, 10 km W of Tiassalé, Fosberg 40539 (MO, US); Yalé, near M Nimba, Geerlin g & Bokdam 1828 (BR, MO); 5 km N of Sassandra, Pu 2261 (BR, MO); N of Sékré, ca. Í ee š Oldeman 604 (BR, K, MO); 10 k Oldeman 701 (BR, MO); N’Zidah, sarin 13683 (G, MO); Dabou, Roberty 15528 (G); N'Zo, Roberty 16049 (G); Adiopodoumé, de Wilde 861 (BR). LiBER1IA: Die- bla, Webo District, Baldwin 6305 (K, MO, NY, US); Gbawia, Baldwin 67 14 (MO, NY, US); Gretown, Tchien District, Baldwin 6924 (MO); Sanokwele, Baldwin 9550 d ina Kle, Boporo District, Baldwin 10570 (K, MO); Wohmen, Vonjama District, Baldwin 12027 (MO); hinterland of Monrovia, Dinklage 3375 (BR); Ganta, Sanokwele District, Harley 959 (MO); Peahtah, Be- quaert in Hb. Linder 1012 (K, US). NIGERIA: Itu, Cross River State, Ariwaodo 83 (MO); Onitsha, Barter 1814 (K, W); Obom Itiat, um to Atam Eki, Calabar Prov- ince, Jones in F.H.I. 6870 (K); Olokemeji Forest Re- serve, Abeokula Province, Jones et al. in F.H.I. 14564 y; aga Ndoro Di gato Baissa po : Fore est Herbarium Ibadan 61439 (K); Ilaro Forest Reserve, 45 Ga SW of pees S.N. Oban, Talbot 856 (BM); valier 2390 (P); Bignona, Roberty 6424 (P). SIERRA LEONE: without exact locality, ee & Smeathman s.n., anno 1792-1796 (BM, S), J. E. Smith s.n., anno 1791 [*Guinea"] (BM). Mt. Loma, Adam 22422 (MO); Njala, Dalziel 8420 (US); Kennema, Deighton 397 (K), Thomas 7822, 7903 (K); Kambia, Deighton 838 (K); Heddle’s Farm, Elliot 3939 (BM, K, US); near Kambia, on Scarcies River, Elliot 4389 (BM); Jola, 20 Jan. 1927, Fairchild s.n. (US); Zimi DM DE Fisher 1 (K); Ma- T Glanville 58 (K), K. ai Reserve, Lane-Poole 4 (K); Kuntaia, ons 5 (K); Yakala, Thomas 24 (K); Jigaya, Thomas 2719 (K); Kanya, Thomas 2983 (K). SRI LANKA: without definite locality, C.P. [^T hwaites"] 922 (BM, BR, G, K, W); Henaratgoda, 3 Jan. 1881, Ferguson s.n. (PDA, W); Matale, Dec. 1846, Gardner s.n., C. P. 922 (PDA), Mar. 1883, Lawson s.n. (K); Buttala to Sirigala, 3 Mar. 1907, Ro ck s. n. (PDA); Dolukanda, Senaratna 2700 (PDA); Dulva Kanda, Ta- SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA 881 teoka 599 (B). SUDAN. EQUATORIA: Talanga, Imatong Mountains, 4°01'N, 32*45'W, Friis & Vollesen 484 (BR, K); Lotti forest, Myers 9655 (BM); Sakure, Zande Land, Wyld 334 (BM š sawasawa, Haerdi 624 (BR, G). ToGo: Tomegbé, Bru- nel & Heitz 5837 (B); Cascade de Tomegbé, S of Badou, Ern 2096 (B); Plateau de Danyi, zwischen Adéta und Ndigbe-Apédomé, Ern 2703 (B, K). UGANDA: Damba Island, Kyagwe County, Dawkins 459 (BM, K), Mait- land 801 (K); Gulu, Zoka forest, Acholi District, Thomas 4031 (K). ZAIRE: Litendale, Achten 485-A (BR); Barumbu, Bequaert 969 (BR); Avakubi, Bequaert 1726 (BR); Mayumbe N’Benga, Bittremieux 106 (BR); Men- kao, Breyne 918 (BR): Mabana, Maluku, Breyne 3292 MO); Mayombi, Brishe 31 (BR); Luni, Brishe s.n. Delhaye 24 (BR); Nkolo, SNE Devr 54 (BR); ukolela, Dewevre 544 (BR); B eut Dewulf 331 (BR); INEAC, Luki, Du Bois 333 (BR, K), 334 (BR) kamba, Bulungu, Dujardin 491 (BR); Bingila, Dupuis s.n. (BR); Boyabokuda-Bogula (Badangabo), Evrard 310 (BR); Djoa, Bolombo, Evrard 4952 (BR); Tukpwo, Gerard 2182, 4298 (BR); Ile Esali, Yangambe, Ger- main 384 (BR, K), Louis 6948 (B, BR), 7900, 13072 (BR); Panza, Inongo, Gilbert 14213 (BR); Kisantu, anno 1900, Gillet s.n. (BR); Yambata, de Giorgi 1669 (BR); Bolobo, env. Eala, Goosens 2448 (BR); Karawa, Uban- gi River, Goosens 4123 (BR); Gatanga, de Graer 294 ville, 15 Jan. 1904, Laurent s.n. (BR); Gimbi, Laurent 600 (BR); Bolombo, Lebrun 605 (BR, US); entre Bu- singa et Banzyville, Lebrun 2040 (BR, US); Moburasa, p ps 193 (BR); Tambwe-Mvwenza, Dibaya, Liben 4 (BR); Tuzule, riv. Lubi, Liben 2976 (BR); Mu- s Luluabourg, Liben 3507 (BM, BR); bord de la cM 40 km N of Kisangani, Lisowski 16480 (BR); panga, bord de la Mobi, 34 km SE : Kisangani, Lisowski * 7219 (BR); Lovanium, Kinshasa, Lisowski 18349 (BR, K); 8 km N of Yakusu, Lisowski 86435 (BR); BR Booke wa Mbole, Yangambi, Louis 10780 (BR, NY); entre Ngazi et Aruwimi, Louis 12181 (BR); dango 3050 (BR); Loata, Meurillon 23, 224 (BR); Yam- bata, Montchal 136 (BR); Dundusana, Mortehan 634 (BR); Kaniana-Haut Lomami, Mullenders 472, 1226 maka, Luki, Nsimundele 57 (BR); Pauwels 2360, 2388 (BR); Campus UNAZA (Lemba), Kinshasa, Pauwels 6378 (BR); Epulu, Putnam 50 (BR); s.n. (BR); Boguge, prés Mobw y Luki, be de la Minkudu, Toussaint 2273 (BM, BR, M); Sonso, Kwango, Vanderyst B-48 (BR); Kiala, Mar. 1907, T wm s.n. (BR); et Wemba, = r Sacre-coeur, BR); Yindu, Puede 93 (BR); Kimuingu, Vanderyst 882 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 AFRICA = C sn i] L P pz E "ry. c.l (1000 km Y f M fr A l | ad \ VTACN 30 L Y) Y " U I ea UN 800 MILES ° 400 800 1200 KILOMETERS | SINUSOIDAL PROJECTION | \ | | EL \ a \ l l 1 E l | J Lo 20 O WEST LONGITUDE FAST NGITUDE ( 20 30 _ 40 50 | 60 GOODE BASE MAP SERIES Prepared by Henry M. Leppard DEPARTMENT OF GEOGRAPHY © 1964 by The University of Chicago Of THE UNIVERSITY OF CHICAGO HENRY M. LEPPARD, EDITOR FIGURE 5. Distribution of Streptogyna crinita in Africa; inset, distribution in Sri Lanka and southern India. 1987] 313 (BR); Dima, Vanderyst 862 (BR); Iles du Kasai, Vanderyst 993 (BR, US); Mokaba, Vanderyst 1693, 3606 (BR); Kikwit, Vanderyst 2783 (BR), 2913 (BR, MO), 9235 (BR); Mukulu, Mee hae 91 (BR); Chen- al, dei 4502 (BR); Yanga, Apr. 1915, Vanderyst .H. mpako, Vanderyst 5429 (BR); Kisantu, Vanderyst 5024, 20399, 29971, 29979, 32082 (BR); (BR); = 25681 (BR); Tsanga, Vanderyst 26986 (BR); u, Kipako, Vanderyst 30696 (BR); Bokoimkori, E. 32624 (BR); Lemfu, Apr. 1907, Vantolborg o name illegible] 21 6 (BR). Streptogyna crinita is widely distributed in wet to seasonally dry forests from sea level to 1,000 meters elevation in tropical Africa (Senegal to southwestern Ethiopia south to Fernando Po, northern Angola, and central Tanzania), south- ern India (Kerala), and Sri Lanka (Fig. 5). Based on abundant collection data from Africa, S. cri- nita appears to flower all year, with a maximum during October through January and slight min- ima (as denoted by a small decrease in the num- ber of flowering collections and a considerable increase in the number of sterile gatherings) dur- ing August and September and to a lesser extent February and March, although Hens 156 states that the species flowers "toute l'année" in Zaire. Several collectors note that S. crinita may be locally dominant, covering the forest floor in large rhizomatous clones. The species is used in west- ern Africa to catch mice and rats, the animals becoming entangled in inflorescences that are placed outside their holes (Hubbard, 1956). zelius & Smeathman s.n., anno 1792-1796, gave the herbarium name of “‘Aristidoides mu- ricida" [*awned mouse-killer"] to the plant. N . Streptogyna americana C. E. Hubb., Hook. Icon. Plant. 36(6): 1—6, tab. 3572. 1956. TYPE. Suriname: trail to Coppename River, rear of village of Paka-Paka, Maguire 23975 (ho- lotype, K, not seen; isotypes, F, MO, NY). Figure 6 Plant cespitose or rarely from a series of short, knotty, horizontal sympodial rhizomes up to 2.5 cm long, the erect culms representing aerial ex- tensions of very short sympodial rhizomes, sometimes becoming decumbent and rooting at the lower nodes. Leaves clustered at base of plant, strongly overlapping, usually concealing all the nodes, often displayed in a fan-shaped arrange- SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA 883 ment; leaf sheaths glabrous below, ciliate on the margins, hispid at the summit; outer ligule 0.6- 1.1 mm long, erose or with a smaller apical fringe of cilia; inner ligule 1.1-2.7 mm long; sheath auricles not developed; lateral appendages usu- ally inconspicuous; oral setae not evident, 1-2 mm long; pseudopetiole not well differentiated from remainder of blade; leaf blades linear, 50- 78 cm long, 0.8-1.6(-2.4) cm wide, glabrous, oblique, the midrib noticeably excentric, flat but often becoming inrolled; primary lateral veins 3— 5 on each side of the midrib; upper (adaxial) blade surface dark green, the lower surface lighter green; cross-veins inconspicuous. Inflorescence 25—40(-67) cm long, borne on a peduncle 1-8- (740) cm long; spikelets 14—25(—49), the glumes of adjacent spikelets not or only slightly over- lapping; rachis subglabrous below, appressed- pubescent above. Spikelets 4—6 flowered, borne on pedicels 1-3 mm long; first glume 3-12 mm long, linear to lanceolate, a 1-3(-5)- nerved; second glume 10- m long, 1.1-1.7 mm wide, ovate-lanceolate, aoe 9-nerved, with scattered inconspicuous transverse veinlets and an awn up to 3 mm long; lemma of lowest floret 19-24 mm long, narrowly lanceolate, completely glabrous, the (5—)7(—9)-nerves green, evident on the inner surface but not visible on the granular outer surface except near the summit ofthe body; calluslike prolongation of lemma 1-2 mm long, the terminal awn 12-21 mm long, arising from between 2 inconspicuous teeth at the summit of the body of the lemma; rachilla internode at- tached to lowest floret, persistent, 4-6 mm long; palea slightly longer than the lemma, usually pro- truding from it by 0.5-3 mm; lodicules narrowly lanceolate, firmly membranous and strongly nerved below, often abruptly widening about '4 of the way from the base, then tapering in the upper 75 to an attentuate, nerveless, hyaline apex, this usually fringed with prickles or occasionally with a few cilia, microhairs absent; anterior lod- icules 3.2-6 mm long, 0.5-1 mm wide, 1—3(-5)- nerved, the posterior lodicule often slightly shorter and narrower, | (—3)-nerved; anthers 2.5- 3.5 mm long, narrowly linear; ovary glabrous; stigmas 3, lacking coarse barbs, glabrous below, at maturity hispidulous-papillose adaxially near the summits; caryopsis 14-17 mm long, 1-1.2 mm wide, glabrous. _ Addit nd specimens examined. BELIZE: Cohune n road, Gentle 8121 (BM, F, G, NY, S, US); Cohune riae. ill slope, Hummingbird Highway, Genile 8682 (BM, F, G, NY, S, US); 40 miles section 884 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 A 4 — = POP f FIGURE 6. Streptogyna americana. — A. Habit of plant, showing deciduous leaf blades.—B. Ligular region, showing hispid sheath summit, outer ligule (left), and inner ligule and oral setae (right). — C. Section of abaxial surface of leaf blade, showing midrib (left) and absence of transverse veinlets. — D. Spikelet.—E. First glume. — F. Second glume. — G. Base of floret, showing basal prolongation of lemma and persistent rachilla internode.— . Lemma, unrolled, showing inner surface. — I. Palea, ventral surface. — J. Palea, profile. — K. Lodicules, with posterior member in center. — L. Androecium of two stamens and gynoecium. — M. Stamen. — N. Stigma, showing 1987] Hummingbird Highway, Gentle 9005 (S, US). BOLIVIA. BENI: Alto Ivon, 11?45'S, 66°02'W, Boom 4792 (NY), 4827 (MO, NY, US); 18 km E of Riberalta, 11°05’S, 65°50'W, Solomon 6169, 7801 (MO). BRAZIL. ACRE: 125 km from Rio Branco on road to Pórto Velho, 9°45'S, 66°20'W, Calderón & Soderstrom 2301 (US). AMAPÁ: Rio Jari, near Cachoeiras das Guaribas, 0?24'N, 53°07'W, Egler & Irwin 46417 (NY, US); road to Ama- pá, vicinity of km 108, Rio Pedreira, Pires & Caval- , 2.5 km sommet, Sastre 1656 (CAY nte Pascoal, p S, 39°24’ W, 2410 (B, NY o 101 at point 13 km N of Itama arajú, Soderstrom et al. 2193 (CEPEC, US). E : Reserva Florestal de Linhares, 19°24’ S. 40°04 W, Martinelli & Soder- strom 9757 (RB), Soderstrom & Sucre 1883 (CEPEC, US). MARANHÃO: Fazenda Bacaba, Doctor Haroldo, 5 km S of MA-119 from entrance 3 km NW of Lago do Junco, 4?26'S, 44°58’ W, Daly et al. 469 (MO, NY); Rio Pindaré, Monção, Fróes 20312 (US); Caxias to Barra do Corda, before Curador [Pres. Dutra], Swallen 3583 (US). MATO GRosso: Fazenda Az de Ouro, 14?13'S, 57*02'W, Amaral 9 (RB); 260 km along road NNE of Xavantina, a few miles W of base camp at 12°51’S, 51?45'W, Eiten & Eiten 8903 (US); 5 km NW of base camp, 12?49'S, 51?46'W, Harley & Souza 10271 (US); 8 km NE of base camp, Ratter et al. 974 (NY, US); 4 km N of base camp, Ratter et al. 1820 (K, NY, US); Serra do Itapirapuan, Alfonso, Lindman A-3347; Sar- aré, 15?05'S, 59°50'W, Pires & Santos 16316 (MO, NY). PARÁ: Belterra, Black 47-936 (NY); inter Sào Joao et Santa Anna, Burchell 9201 (BR, US); Rio Part de Oeste (TIRIOS), Cavalcante 824 (US); Gorotire vil- lage at Rio Fresco, 7°47’S, 51°07'W, tidie QA ad - sey 17-22183, 32-24183 (MO); Curuá Al r, Kuhlmann yp iud S); Maicuru, s Francisco, Pires Y, US); Serra Buritirama entre B-2 e Marabá, ese 12320-A (US); range of low hills ca. 20 km of Rendencao, near Córrego pad João eresa, 8°03’S, 50°10'W, Rendencáo on road to Barreiras dos Campos: Fazenda Inajapora between Rio Inajazinho and Rio Inajá, ca. 8945'S, 50°25’W, Plowman et al. 8883 (F, MO, NY, US); Belém-Brasilia Highway 17 km S of Ligacao do Para, near km 1,509, ca. 4°17'S, 47°32'W, Plowman et al. 9409 (F, MO, NY, US, WIS); 6 km N of Ligação do Para, near km 1,532, 4°05'S, 47°32'W, Plowman et al. 9522 (MO); 12 km E of Représa Tucurui (Rio To- cantins), 3°45’S, 49°40'W, Plowman et al. 9799 (NY); Jari, estrada do Munguba, km 14, Si/va 2198 (MO, Y); Lageira, airstrip on Rio Maicurt, 0°55'S, 54°26' W, Strudwick & Sobel 3109 (F, MO, NY); Sete Varas air- strip, Rio Curua, 0°59’S, 54?29'W, Strudwick & Sobel 4289, 4313 (F, MO, NY); Santarém, Swallen 3281 SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA 885 (US); Japanese concession 35 km N of Monte Alegre, Swallen 3411 (US); Obidos, Swallen 5089 (US ). Cordeiro 924 (MO); 1 km N of Riberáo, road Abuná- Guajará-Mirim, Prance et al. 6454 (MO, NY, US), 6527 (F, MO, NY, S, US); in sylvis umbrosis ad flumen Guaporé, Riedel 1248 (G, NY); Mineração Mibrasa, setor Alto Candeias, km 128, 10?35'S, 63°35'W, Teix- eira et al. 625 (MO). RORAIMA: Conceição, Rio Blanco, Luetzelburg 21383 (M). COLOMBIA. CAQUETÁ: Entre 3629 (F). META: E artin, del Cano Camoa, í. 1687 ); Monte de Caño oa, Herman S); sabanas de Sa Juan de Arama, margen (Ee. ] iiejar, ater- rizaje “Los Micos,” Idrobo & Schultes 608, 1217 (US); margen izquierda del Rio Sansa, Sierra de la Macarena, Idrobo 2160 (NY); Cano Ciervo, Sierra de la Macarena, Philipson et al. 2023 (BM, S, US); margen derecha del Rio Guayabero, Raudal de la Macarena (Angostura 1), Pinto & Bischler 334 (US). VICHADA: ca. 35 km from Las Gaviotas on road to Santa Rita, Davidse & Llanos 5211 (MO); Gualandayas, ca. 100 km E of Gaviotas, Wood 4220 (K). CosrA RICA. PUNTARENAS: Finca Los Helechales, between Buenos Aires and Cerro Pittier, Hatheway 1686 (US); Los Tejares de Buenos Aires, Pittier 10602 (BR, M, US, W); entre le Rio del Con- vento et Buenos Aires, Tonduz 3643 (BR, W). FRENCH Cremers 4546 (CA pann-Mitaraka (frontiére) P.K. 7.5, Granville 1139, 1417 (CAY, US); versant N des Monts Galbao, 10 km WSW de Saül, 400 m, Granville 1621 (CAY); 14 km de Dégrad Claude, Granville 2267 (CAY); Sommet Ta- bulaire, ca. 50 km SE Saül, Granville 3586 (CAY, MO, : min des Emérillons, 1 km de Saut Verdun, Granville B- 5037 (CA Y); Cayenne, Chemin du Moulin Vidal, 13 July 1955, Hoock 1187 (K, P); Saül, 30 June 1956, Hoock s.n. (NY); Cayenne, anno 1839, Leprieur s.n. (G). anno 1866, Jelski s.n. (W); Montabo, Herb. L. C. Richard s.n. (Wy; Karouany, Sagot 1076 (BM, W). GUATEMALA. ALTA VERAPAZ: ca. 6 km E of Sebol on San Luis Road to Achiote, Contreras 4486 (US). MEXICO. CHIAPAS: Javalinero, Palenque, Matuda 3637 , US). VERACRUZ: Sanborn [ca. 17?34'N, 95?07' w], Orcutt 2933 (K, MO, US). NICARAGUA. ZELAYA: region of Braggman's Bluff, E nglesing 218 (F); Miguel Bikan, aspám, P. Lake, Miller 2045 (US yh y 27480 (MO, US); forest eee telephone cable trail — adaxial papillae near tip.—O. Caryopsis, ventral surface showing linear hilum.—P. Caryopsis, dorsal pid showing small, basal embryo. (Based on uie for CF, L J, L, O, P; 2 mm for B, G, H; zil, Soderstrom et al. 2193 xa Scale bar = 24 mm for A; 4 mm for K, M; an N.) nd 0.5 mm ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Ratte d O 100 200 300 400 500 600 mies Prepared by Hendrik R Rypkema \ 30 | FiGuRE 7. Distribution of Streptogyna americana. between S-16 and S-49, Rio Indio, Steyermark & Allen 17459 (BR, G, MO, S, US). DARIEN: Cerro Pirre, Gentry & Clewell 7147 (MO), Mori & Kallunki 5374 (MO, ; 0.5-2.5 km NE of Manené, Hartman 12175 (MO). PANAMÁ: San José Island, Johnston 433 (BM, US), 728, 1115 (US). PERU. MADRE DE pios: Rio Tambopata, Lago 3 Chimbadas, ca. 65-70 river km SSW of Puerto Maldonado, ca. 10-15 km air SW eff. Río La Torre, 12°49 9°17'W, P 5762 2: SURINAM: without locality, Weigelt s.n. (W); Zuid River, 3 km above confluence with Lucie River, /rwin et - 55900 (B, MO, NY, US); ab Wia wia-bank e Zwie- belzwamp, Lanjouw & a 1153 (NY) ad prope Moengo, Lindeman F); Brownsberg, near Is fall, Lindeman ATES (K, ih Lucie River, ca. 2 km below affluence of Oost River, Maguire et al. 54093 (NY, US); Avanavero oever, Stahel & Boon- acker 4579 (US). TRINIDAD: without locality, Botanical Garden Herbarium No. 3367 (US), Crueger 362 (US fragment), Finlay 3 (K, W); Caparo forests, Broadway 4932 (US); Tabaquite, edge of High Woods, Hitchcock 10122 (BM, US). VENEZUELA: Alto Orinoco, Rusby & Squires 349 (K, NY). AMAZONAS: vicinity of Culebra, Río Cunucunuma, 3?40'N, 65?45'W, Steyermark & Delascio 129185 (MO); Mavaca, Alto Orinoco, Aris- teguieta & Lizot 7380 (NY). BOLIVAR: alrededores km 88, carretera El Dorado, Aristeguieta 3713 (MO); 17 km W of Río Caura on road between Caicara and Ciudad Bolivar, Davidse 4443 (MO, WIS); 10 km SW of Río Aro on road between Caicara and Ciudad Bo- livar, Davidse 4476 (MO); km 28, S of El Dorado, Davidse 4966 (MO); 20-35 km SE of Monteco on road to San Pedro de las Dos Bocas, 7?10'N, 62*55'W, Lies- ner & Gonzalez 5851 (MO); along pica 105, 40 km S of Tumeremo, 29 km N of El Dorado, Steyermark 86572 (NY); Pica La Lira, at km 27 S of El Dorado, l m E of hi ave ages 86638 (NY); woods bordering savannah by Rio S : hee Par ke Steyermark 86760 (U AMACURO: 3 k Piacoa [p E in the state of Delta yen ancl Steermark tag : Rio Cuyubini, Cer a, (NY). SUCRE: S ies n in Imp Cedeno and Boca del Tataricual, along Quebrada Im- posible, Steyermark 62845 (F, US). a, above raudal Cotua, S). DELTA Streptogyna americana is found in shaded, well-drained sites in moist forests below 500 1987] (-800) meters from Veracruz, Mexico, and Trinidad south to northern Bolivia and Espirito Santo, Brazil (Fig. 7). Most common on the mar- gins of the Guyana Highlands and in easternmost and southernmost Amazonia, this species is rare or absent in the central portion of the Amazon Basin. Collectors in Panama (Judziewicz 4440), Venezuela (Davidse 5211), and Surinam (Irwin 55900) noted that the leaf blades become inrolled during hot dry weather or soon after collection. Most Central American collections were made November through April, indicating a dry season peak of bloom, while Guyanan and eastern Am- azonian collections have been made principally June through August; the five Atlantic Brazilian gatherings were made from March to May. Com- of S. americana include “barba de azil) and "barba- tigre" (Cabrera 3629, CE. suggesting ex- ternal animal dispersal. LITERATURE CITED BENTHAM, G. 1883. Gramineae. In G. Bentham & J. . Hooker, Genera Plantarum, Volume 3, Part 2. R. SODERSTROM. 1973. Mor- phological and carpets considerations of the grass subfamily Bambusoideae based on the new genus Maclurolyra. Smithsonian Contr. Bot. 11: i-i, 1— 1980. The genera of the Bambu- soideae (Poaceae) of the American continent: keys nd comments. Smithsonian Contr. Bot. 44: 1 CAMPBELL, C. S. & E. A. KELLOGG. c: press. Sister group EENE om the Poaceae. /n T. R. So- l. (ed tors), Una cedet s and Wash- Symposium on Grass Systematics and Evolution. Washington, D.C., 27-31 July 1986.] CLAYTON, W. D. & S. A. RENVOIZE. 1986. Genera Graminum. Her Majesty's Stationary Office, Lon- don DAHLGREN, R. M. T., H. T. CLIFFORD & P 1985. The Families of the vetula ua Springer-Verlag, Berlin. DoELL, J. C. 80. Tribe 11, Bambusaceae. 7n C. F P. von Martius (editor), Flora Brasiliensis 2(3): 161—220, pls. 44—56. [Fascicle 83.] DuJARDIN, M. 1978. Chromosome numbers of some tropical 'African grasses from western Zaire. Ca- nad. - Bot. 56: patient ERR 1887. Gra Peer Die a in. spa] sa ea pex E 9 HOOKER, : 1900. In H. a teri Hand- book an Flora of Ceylon, V e HUBBARD, pi E. 1936. Pp. 475- 602 in} Hutchinson SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA 887 & J. M. Dalziel (editors), Flora of Tropical West Africa. Crown Agents for the Colonies, London. 1956. Streptogyna crinita Beauv., Gramin- eae, Tribus Streptogyneae. Jn Hook. Bot. Icon. 36(6): 1-6, tab. 3572. É 62. Les graminées d'Afrique cale. I. Généralités, classification, descrip- Viviéres, Bull. Sci. (Paris) 8: xi + 1-345. . ANOMA, E. ADJANOHOUN & L. AKE 3. Nombres chromosomiques de Gra- ées de Cóte d'Ivoire. Candollea 28: 191-217. SERE Y. 1960. The first FUE, leaf in grass systematics. J. Jap. Bot. 35: 5. METCALFE, C. R. 1960. . of the Monocoty- ledons. I. Gramineae. Clarendon Press, Oxford. NEES VON ESENBECK, C. G. D 35. Bambuseae Bra- silienses. Recensuit, et alias i in India orientali pro- 4. . 1812. Es- une Nouvelle Agrostographie: ou Nouveaux Gen nres des Graminées; avec Figures Sali in i: d de Tous les Genres. Fai PIL, 1982. Principles of olde in te "Plane 2nd edition. Springer-Verlag, Ber- lin. REEDER, J. C. 1962. The bambusoid embryo: a reap- praisal. k J. Bot. 49: 639-641. RENVOIZE, S. A. 85. A survey of leaf-blade anat- omy in — V. The bamboo allies. Kew Bull. 40: 509- cy RIDLEY, H. 1930. The Dispersal of Plants =a i the World. L. Reeve, Ashford, Kent, England. nglan SMITHSON, E. 1956. The comparative anatomy of the Flagellariaceae. Kew Bull. 3: 491-501. SODERSTROM, T. R. Some evolutionary trends in the Bambusoideae (Poaceae). Ann. Missouri Bot. Gard. 15-47. R. P. ELLIS. In press. The position of bam- and Evolution. Washington, D.C., 27-31 July 86.] E. J. JUDZIEWICZ. ded pes Phar- eae and Streptogyneae of Sri orpho- logical-anatomical study. Hu meu Cae Bot. 65: 1-27. STEBBINS, G. L. 1982. Major trends of evolution in the Poaceae and their possible significance. Pp. 3- 36 in J. R. Estes et al. (editors), Grasses and Grass- lands. Univ. Oklahoma Press, Norman, Okla- homa. SrEUDEL, E. G. 1853 [1855]. ped: ii T Glumacearum, Part 1. J. B. Metzler, Stu n the genus [oed (Po- : 364-366. omatic chromosomes of Leptaspis and Streptogyna (Poaceaz). Nature 182: 1619- Starch grains of endosperm in grass systematics. Bot. Mag. (Tokyo) 75(892): 377-383. 888 1965. Chromosome numbers of some East African grasses. Amer. J. Bot. 52: 864-869. —— & T. TAKAGI. 1967. Notes on some grasses. XIX: systematic significance of microhairs of lod- icule epidermis. Bot. Mag. (Tokyo) 80(952): 394— 4 TRAN VAN Nam, MME. (TRAN THI TUYET Hoa). 1972. Les "oreillettes" des Graminées. Bull. Soc. Bot. France 119: 441—462. VEYRET, Y. 1958. Observations caryologiques chez quelques Graminées tropicales. J. Agric. Trop. Bot. Appl. 5: 308-310. YAKOVLEV, M. S. 1950. Struktura endosperma 1 za- rodysha zlakov kak sistematicheskii priznak. Moskva , Lenin ngrad: Akad seriya orfologiya i Anatomiya Rastenil) 1: 122-218. [Endosperm and embryo structure of grasses as a taxonomic feature. Moscow, Lenin- grad: Transactions ofthe V. L. Komarov Botanical Institute of the Academy of Sciences of the USSR, series 7 (Morphology and Anatomy of Plants) 1: 121-218. Unpublished English translation by the Indian National Scientific Documentation Centre, prepared in 1975.] ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 APPENDIX I. LIST OF TAXA CITED Arundinaria vagans Gamble am h Joinvilleaceae Tomlinson Maclurolyra tecta Calderón & Soderstrom Feds latifolia L na parvispica R. Pohl Pleiobla stus Nakai Sasa Makino & Shibata Streptia crinita Rich. ex Doell Streptia secunda Rich. ex Doell Streptogyna P. Beauv S. americana C. s Hubb. S. crinita P. Bea S. gerontogaea Hook. f. in Trim Streptogyneae C. E. Hubb. ex ^p E & Calderón WOOD ANATOMY OF NOTEWORTHY SPECIES OF LUDWIGIA (ONAGRACEAE) WITH RELATION TO ECOLOGY AND SYSTEMATICS! SHERWIN CARLQUIST? ABSTRACT Ludwigia anastomosans, a tree to 10 m, is studied because it is unusual in the genus in its arborescent habit. It proves to vods a = vessels; it also has interxylary phloem, hitherto reported for only one species of the they may be more a are the first observed in Onagrac hos . Wood anatomy of Ludwigia peduncularis and L. torulosa shows that eae uc Bud don so far indicated. Vestigial bars on perforation plates of L. aceae and are believed to represent an instance of paedomor- phosis, but also retention of a primitive feature. Other indicators of paedomorphosis in Ludwigia are abundance grouping is related to ecology in taxa having fi of erect ray cells and — long vessel elements. The hypothesis that degree of vessel ber -tracheids Or libriform fibers i is validated by Ludwigia, which has the lowest degree of the y, combined in ‘the Mesomorphy ratio, present a not dissimilar pattern features reflective of ecolog 1dilill c. Other eyes e that can be integrated with hn given by HF rates and temperature regimes as well as water availability. Data on wood anatomy of eight species of Lud- wigia have been presented earlier (Carlquist, 1975, 1982a). That number seems small unless one takes into account the fact that Ludwigia is predominantly herbaceous; the most familiar species are nonwoody herbs of very wet habitats such as ponds, ditches, and streams. One of the species in the present study is a notable excep- tion: L. anastomosans (DC.) Hara is a tree. The data on the collection studied here describe it as a tree 10 m tall with a trunk 15 cm dbh. Wood anatomy is of special interest because of this hab- it. In fact, the results obtained below demon- strate that the wood of L. anastomosans differs appreciably from that of other Ludwigia species. Dr. Elsa Zardini, who collected the material see if wood anatomy demonstrated the degree of relationship between them. Dr. Zardini has con- templated the idea that L. peduncularis (Griseb.) Gómez may be closely related to L. torulosa (Arn.) ara. The wood anatomy of Ludwigia is of consid- erable interest with respect to ecology because Ludwigia characteristically grows in very wet places. Ludwigia anastomosans was collected in bamboo clumps by a blackwater stream in the Parque Natural de Caraca, Minas Geraes, Brazil. The material of L. peduncularis came from ditch- es near Havana, Cuba. The L. torulosa specimen was collected in a natural pond 17 km south of Tumeremo, Distrito Roscia, Estado Bolívar, Venezuela. In taxa with libriform fibers such as Onagraceae, Carlquist (19842) hypothesized de- gree of vessel grouping to be in direct proportion to adaptation to dry conditions. In this case Lud- wigia species ought to exhibit a low degree of vessel grouping. Although figures for vessels per group were developed in the earlier survey o woods of the family (Carlquist, 1975), no com- parisons were made between those figures and ecological regimes occupied by the various species. Ludwigia is a group of interest with relation to paedomorphosis in wood anatomy. This is, in turn, related to habit and ecology. The species in the present study were examined in this con- text to see if woodiness in Ludwigia is primary or secondary. Although a study on wood anatomy can be expected to reveal new records for anatomical features, two in the present study proved of es- pecial interest and worthy of discussion: occur- rence of interxylary phloem and presence of ves- tigial bars on some perforation plates. ' Grants from the National Science Foundation to Peter H. Raven made possible the collection and selection of the material for this researc! ? Rancho Santa Ana Botan Pomona College, Claremont, California 91711, ANN. Missouni Bor. GARD. 74: 889-896. 1987. c Garden, Claremont, California 91711, U.S.A., and Department of Biology, U.S.A. 890 MATERIALS AND METHODS Voucher specimens are located at the Missouri Botanical Garden. Appreciation is expressed to Dr. Elsa Zardini for providing dried wood sam- ples suitable for study. For L. peduncularis and L. torulosa, samples represented basal portions, but diameter was small (2 and 4 mm, respec- tively. The material of L. anastomosans was from a branch about 1.5 cm in diameter. Al- though this is much less than the 15 cm diameter reported for trunks of this species, the branch material is considered here to represent an es- sentially mature pattern. Woods were boiled in water, stored in 5096 ethyl alcohol, and sectioned on a sliding micro- tome. Sections prepared in this way were, in part, satisfactory, but cell collapse on account of thin- ness of wood cells was excessive in some in- stances. Therefore an alternative method in- volving further softening, embedding in paraffin, and sectioning on a rotary microtome (Carlquist, 1982b) was employed. Sections prepared by both techniques were stained in a safranin-fast green combination. Macerations were prepared with Jeffrey's Fluid and stained with safranin. Means are based upon 25 measurements (few- er if feature is scarce) except for vessel wall thick- ness, libriform fiber diameter, and libriform fiber wall thickness, in which a few typical cells were measured. Vessel diameter includes the wall, al- though lumen diameter may be preferable for some purposes and may be calculated by sub- tracting wall thickness from the data presented here. Mean values for vessel grouping are ob- tained on the following basis: a solitary vessel — 1.0, a pair of vessels in contact — 2.0, etc. Bark was not observed specifically in the present study, although sections of the stem of L. torulosa in- cluded portions of the spongy stem covering that proves to be aerenchyma like that figured for “Jussiaea repens" L. (now a Ludwigia) by Met- calfe & Chalk (1950), and studied in detail by Ellmore (1981) for L. peploides (HBK) Raven. ANATOMICAL DESCRIPTIONS Ludwigia anastomosans (DC.) Hara, Zardini & Gentry 2175 (Figs. 1—6). Growth rings inconspicuous, and probably re- lated to water level of the riparian habitat (Fig. 1). Mean number of vessels per mm?, 51. Mean number of vessels per group, 1.24; vessels tend- ing to be grouped into radial multiples (Figs. 1, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 3). Mean vessel diam., 109 um. Mean vessel wall thickness, 2.5 um. Mean vessel element length, 458 um. Perforation plates simple (Figs. 2, Lateral wall pitting of vessels alternate, pits crowded and circular to polygonal in outline, about 12 um diam. on vessel-vessel interfaces (Figs. 4, 5). Vessel-axial parenchyma and vessel- ray pitting alternate to scalariform, pit apertures long (sometimes scalariform), pit apertures wide (“gaping”). Pitting with relatively conspicuous vesturing on vessel-vessel pits (Fig. 5, upper right), vesturing less pronounced on vessel-axial parenchyma and vessel-ray pits. Occasional ves- loses abundant (Figs. 1, 3). Imperforate (cheap elements all libriform fibers because pits appar- ently simple, although a few exceptional pits with small borders also observed. Many libriform fi- bers septate. Mean libriform fiber diam., 28 um. Mean libriform fiber wall thickness, 2.3 um. Mean libriform fiber length, 588 um. Numerous libri- form fibers with gelatinous walls (Fig. 6) and therefore probably reaction wood. Axial paren- chyma vasicentric scanty. Bands of phloem-con- taining axial parenchyma present in marginal po- sitions (end of growth rings) or scattered without any relation to growth rings (Figs. 1, 3). Rays multiseriate and uniseriate, the former slightly more frequent (indicated by relative heights in Fig. 2). Ray cells predominantly erect and square (Fig. 2), a few procumbent cells present in mul- tiseriate portions of multiseriate rays. Mean mul- tiseriate ray height, 2,707 um. Mean uniseriate ray height, 349 um. Mean width multiseriate rays at widest point, 3.95 cells. Ray cell walls mod- erately thin, lignified. Wood nonstoried. Raph- ides present in phloem-containing axial paren- chyma strands. Ludwigia peduncularis (Griseb.) Gomez, Ekman 13416 (Figs. 7, 8). Growth rings absent (portion studied probably less than one year's accumulation). Mean num- ber of vessels per mm’, 62. Mean number of vessels per group, 1.30. Mean vessel diam., 50 um. Mean vessel wall thickness, 1.8 um. Mean vessel element length, 484 um. Perforation plates simple. Lateral walls of vessels with crowded al- ternate elliptical pits about 4 x A. Emo with pointed ends l-vessel contacts). ensis pi mE and vessel- ray pitting similar but with pits longer (appearing pseudoscalariform) and wider, and with wider 1987] CARLQUIST—WOOD ANATOMY OF LUDWIGIA 891 4 Wi "ye NUM M oe AN DW ANGE UN Mig M ssh ane a DT 4 ue ^ dil d we UTS A 22 | ki rara p> I " tasbe com 9 Mer 4 z ZU III > zz -. - e P GI — D — — la e 2mos 1@eettan h “S Sas L LLL - a — uS " | | ASE = xy 9 gp» a. argo Adi 47 45 06 VN pean sib B LIN A sana e . e, CI - 2 Pf i. P rg - ae ed -— ALI yy FiGURES 1-4. Wood one of Ludwigia anastomosans, Zardini & Gentry 2175.—1. Transection; termi- nation of growth ring occurs in the middle of the photograph. —2. Tangential section; multiseriate rays are wide, more numerous than uniseriate rays.— 3. Portion of transection, showing strands of interxylary phloem (arrows) and vessels in radial multiples.—4. Portion of vessel wall from tangential section; intervascular pitting and A dopa: plate rim, below; axial parenchyma cells, upper left. Figures 1, 2, i5) r siapa scale above Figure finest divisions = 10 um); Figure 3, divisions = 10 um; Figure 4, divisions = — JOu 892 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 * TT e°, 99 ion "Vac . S. ! i^ $ NA FIGURES 5-8. Wood sections of Ludwigia. 5, 6. L. anastomosans, Zardini & Gentry 2175. —5. Intervascular pitting from tangential section, showing transition between circular and polygonal pit shapes; vesturing evident in pits at upper right. —6. Portion of vessel and associated cells from tangential section; note aberrant pit and pit aperture configurations and splits in gelatinous fiber walls (right). 7, 8. L. peduncularis, Ekman 13416.— 7. Transection; vessels solitary; dark-colored deposits in ray cells. — 8. Tangential section; all rays are uniseriate. Figures 5, 6, magnification scale above Figure 4; Figures 7, 8, magnification scale above Figure 3. 1987] CARLQUIST—WOOD ANATOMY OF LUDWIGIA 893 AT un - tw — 'I i A i y - Ñ 6 t i ` “ ` e D £ ? H "4 Y WU SH IM Fa SH N n A: |s See Hh 4, ass ` i Do | ! aq Top ; f '. t * ot TRA ow: "C . | | | / Tn i h | wb ' t | | Rat Ti Cy | e hielo 9; at in t MA 3t x 187 Meh 4 ite ‘eR :. , | AM YEAR TERTEL 3 | HH WE "e: oat } 1 EAT MES HR < | P w. HTH lust a 8 i AN tt BR ALAA | | MANES HH. ORE Ñ \ <1 AS “lee. N POUR St aa H OSIM EEIE ear TH if Nus Wi spl e WH h NS EIU Ai RC PACER |l: Vr deltas (Sige aes area "| Nuptiis gio dvo HHI / As bs E FEET a st RETE EHI X j AUR BL ERO HEU BL DHRC Sa EA | mE, E FIGURES 9-13. Wood sections of Ludwigia torulosa, Holst, Steyermark & Manara 2257.—9. Transection; libriform fibers are thin-walled.— 10. Tangential secti Itiseriat „l juent tł iseriat rays.— 11. Portion of radial section showing vestigial bars on a perforation plate.— 12. Portion of a radial section showing vessel-ray pitting.— 13. Portion of a radial section showing rodlike crystals in two ray cells. Figures 9, 10, magnification scale above Figure 1; Figures 11-13, scale above Figure 4. 894 apertures. Pits of vessels vestured, less so in ves- sel-axial parenchyma and vessel-ray contacts than on vessel-vessel pitting. Imperforate tra- cheary elements all libriform fibers, the pits mi- nute, apparently simple. Libriform fibers mostly septate. Mean libriform fiber diam., 18 um. Mean libriform fiber wall thickness, 1.2 um. Mean li- briform fiber length, 778 um. About half of the wood consisting of gelatinous fibers (Fig. 7), therefore apparently reaction wood. Axial paren- chyma vasicentric scanty, very few strands ad- jacent to each vessel (or some vessels with no adjacent axial parenchyma). Interxylary phloem absent. Rays almost all uniseriate (Fig. 8). Ray cells all erect. Mean uniseriate ray height, 1,566 um. Ray cells moderately thin-walled but ligni- fied (Fig. 8). Wood nonstoried. No crystals ob- served. Axial parenchyma and ray cells contain- ing droplets or massive accumulations of a brownish substance (Figs. 7, 8). Ludwigia torulosa (Arn.) Hara, Holst, Steyer- mark & Manara 2257 (Figs. 9-13) Growth rings absent (portion studied probably less than a year's accumulation). Mean number of vessels per mm?, 62. Mean number of vessels per group, 1.55. Mean vessel diam., 55 um. Mean vessel wall thickness, 1.6 um. Mean vessel ele- ment length, 374 um. Perforation plates mostly simple, but some with modified or vestigial bars (Fig. 11). Lateral walls of vessels with alternate polygonal pits about 5 um diam. on vessel-vessel contacts. Vessel-axial parenchyma and vessel- ray pits alternate, opposite, or scalariform, larger and with wider apertures than the vessel-vessel pits (Fig. 12). Vestures dense within cavities of vessel-vessel pits, somewhat less abundant on vessel-axial parenchyma and vessel-ray pits. Imperforate tracheary elements all libriform fi- bers, because the pits apparently simple. Libri- form fibers nonseptate, but starch present in them. Mean libriform fiber diam., 25 um. Mean libri- form fiber wall thickness, 1.5 um. Mean libriform fiber length, 538 um. Libriform fiber walls some- what gelatinous. Axial parenchyma vasicentric scanty, often only one strand adjacent to a vessel. Rays multiseriate and uniseriate, uniseriates more abundant (Fig. 10). Ray cells mostly erect, a few square cells present. Mean multiseriate ray height, 1,188 um. Mean uniseriate ray height, 270 um. Mean multiseriate ray width at widest point, 2.2 cells. Cell walls moderately thin but lignified. Wood nonstoried. Large rodlike crystals (one tip ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 concave, the other convex, suggesting paired crystals) present in a few ray cells (Fig. 13). CONCLUSIONS HABIT AND ECOLOGY In a given floristic area, wider vessels char- acterize trees as compared with shrubs and sub- shrubs (Carlquist & Hoekman, 1985). In a genus or family that ranges into diverse habitats, trop- ical tree species tend to have the widest vessels (e.g., Vliet, 1979). Ludwigia anastomosans has notably wide vessels for the family Onagraceae. Mean vessel diameter in this species equals the widest mean diameter reported in the earlier sur- vey (Carlquist, 1975), a fact very likely related both to its arboreal nature and tropical habitat. The figure recorded for vessel diameter in L. an- astomosans may be conservative, because the EE studied is relatively small compared with e large diameter of trunks of these trees. In trees, vessel diameter is greater at the ica of older stems (e.g., Carlquist, 1984 The abundance oferect ray cells in comparison with procumbent ray cells in Ludwigia, even in L. anastomosans, suggests that Ludwigia may represent some degree of secondary woodiness. his idea was entertained for Ludwigia earlier (Carlquist, 1975). The presence of a few vestigial bars on perforation plates in L. torulosa (Fig. 11) is pertinent in this regard. Presence of a few such plates does suggest retention of primitive fea- tures in the primary xylem (the "refugium" the- ory of Bailey, 1944), but it also suggests that this feature is carried forward into the secondary xy- lem by virtue of paedomorphosis. Paedomor- phosis is indicated by erect ray cells and the rel- atively long vessel elements (Carlquist, 1962), two features well displayed in Ludwigia. Pres- ence of occasional scalariform perforation plates of the Asteraceae (Carlquist, 1983a), or Patrinia of the Valerianaceae (Carlquist, 1983b). The presence of scalariform perforation plates is, however, more primitive than presence of simple plates exclusively. In an earlier paper (Carlquist, 1984a), vessel grouping in taxa with libriform fibers (as in On- agraceae) or fiber-tracheids was held to be pro- portional to adaptation to dry conditions. This hypothesis is worth testing in Onagraceae, be- 1987] cause of the wide range of ecological circum- stances occupied by its members. Data on vessel grouping were prepared earlier for the family (Carlquist, 1975), and indices designed to enable comparison with ecological conditions were de- vised (Carlquist, 1977). These data can now be analyzed in the light of the vessel grouping hy- pothesis. Vessel grouping figures were not offered for six Ludwigia species studied earlier (Carl- quist, 1982a), and so are presented here: L. bul- lata rers! Hara, 1.08; L. elegans saad Hara, 1.36; L. peruviana (L.) Hara, 1.10; L. s icea rae Hara, 1.12; L. sonémtosà s bess.) Hara, 1.04; L. sp. (aff. L. longifolia), 1.23. The mean value for vessel grouping in the 12 collections of Ludwigia, sole genus of tribe Jus- sieeae, is e next lowest value, by tribe, occurs in Hauyeae (1.67), followed by Fuchsieae (1.80) and Epilobieae (1.85); data for Lopezieae and Onagreae were subdivided into habit cate- gories. If one considers that the aquatic habitats characteristic of Ludwigia species represent the most mesic ecological situations of the family, vessel grouping is a more accurate indicator of ecology within Onagraceae than the indices morphy figure for Hauyeae (1,242), which seems high compared with that for 12 collections of Ludwigia (415), is related to the wide vessels in that tribe, which in turn is doubtless character- istic for a species transpiring large volumes of water in a warm, moist tropical forest. Both of these figures are much higher than for groups in the family occupying drier habitats, such as an- nuals of tribe Onagreae (161) or caudex peren- nials of Onagreae (48). SYSTEMATICS The data provided are consonant with the idea that L. peduncularis and L. torulosa may be closely related. These species have an unusual feature in common, the presence of rays exclu- sively or nearly aerial at the outset of sec L. torulosa is not considered decisive. Such crys- tals have been reported in L. bullata, L. octo- valvis, and L. peruviana (Carlquist, 1975, 1982a). In turn, these crystals resemble the large, sty- loidlike crystals of Hauya; these latter crystals occur in axial parenchyma rather than in rays, as in Ludwigia. Data from wood anatomy should CARLQUIST—WOOD ANATOMY OF LUDWIGIA 895 not be interpreted in too detailed a way here: variability within species is not known and not all species of all sections of Ludwigia have been studied. Wood anatomy is not often a decisive indicator of relationships at species and section (subgenus) levels e occurrence of interxylary phloem in L. anastomosans is noteworthy. Interxylary phloem has been reported previously for the genus only in L. sericea (Carlquist, 1982a), although it oc- curs elsewhere in the family (Carlquist, 1975). The significance of interxylary phloem seems to lie with seasonality in translocation of photo- synthates, as noted earlier (Carlquist, 1975). Be- cause L. anastomosans sa tree, presumably with an annual flowering season, production of in- terxylary phloem would be adaptive, whereas in an annual or short-lived perennial Ludwigia plant, the phloem from a single year, undiminished b crushing at the end ofa season as in longer-lived plants, would probably suffice to channel pho- tosynthates into flowers and developing fruits. LITERATURE CITED BAILEY, I. W. 1944. The development of vessels in angiosperms and its significance in morphological research. Amer. J. Bot. 31: 421-428. CARLQUIST, S. 1962. A theory of paedomorphosis in dicotyledonous woods. Phytomorphology 12: 30- 1975. Wood anatomy of Onagraceae, with notes on alternative modes of photosynthate movement in dicotyledon woods. Ann. Missouri Bot. Gard. 62: 386-424. 1977. Wood anatomy of Onagraceae: addi- tional rd and concepts. Ann. Missouri Bot. Gard. 64: 627-637. . 1982a. Wood anatomy of Onagraceae: fur- ther species; root anatomy; significance of ves- tured pits and allied structures in b E ai Ann. Missouri Bot. Gard. 69: 755-769. 2b. The use of eibvlenediantinb] in soft- ening hard plant structures for paraffin sectioning. Stain Techn. 57: 311-317. 83a. Observations on the vegetative anat- omy of Crepidiastrum and Dendrocacalia (Aster- aceae). Aliso 10: 383-395. 1983b. Wood anatomy of Calyceraceae and Valerianaceae, with comments on aberrant per- of dicotyledons. Aliso 10: 413-425. j ssel grouping in dap adus ue wood: s mperforate tra- i anatomy of Tiimenitiosds Plant Syst. Evol. 144: & bg 1985. Ecological wood anatomy of the s Aa southern Californian flora. IAWA Bull. 6: 319-347 896 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 ELLMORE, G. S. 1981. Root dimorphism in Ludwigia "napua sere stem in = family ipi p arcis eploides Onagraceae): structure and gas content S. str. ation to the evolution of life for aa roots. Amer. J. Bot. 68: 557-568. Bot. Z dier ics te dean a 65: 627-639. METCALFE, CR .& L. CHALK. 1950. Anatomy of the [In Peka Dicotyledons. Clarendon Press, Oxford. VLIET, G. J. C. M. vAN. 1979. Wood anatomy of the SHULKINA, T. V. & S. E. Zikov. 1980. The anatomical Combretaceae. Blumea 25: 141-223. THE ORIGAMI OF BOTANY: A GUIDE TO COLLECTING AND MOUNTING SPECIMENS OF CYCLANTHACEAE! BARRY E. HAMMEL? ABSTRACT Guidelines for preparing and mounting specimens of Cyclanthaceae, as well as a key to the genera, are presented in order to facilitate collecting of a difficult and often neglected family. In general, specimens should be prepared so the depth of the leaf division is apparent and the lower surface of the basal part of the leaf is visible or accessible. The family Cyclanthaceae has far fewer species than the other related families oflarge intractable monocots for which guides to the preparation of herbarium specimens have recently been pub- lished (Croat, 1985; Dransfield, 1986; Stone, 1983). Neverthel in much ofthe wet lowlands of the Neotropics the cyclanths form a very con- spicuous element of the understory and epiphyte flora (Figs. 1, 2). Careful collecting in almost any area of wet primary forest often yields new species because the family is poorly represented in her- baria and many species are narrowly endemic This paper is a call for more collections as well as a guide to help insure that they be properly prepared. SPECIMEN SELECTION AND PREPARATION How to find them. Cyclanths are often left uncollected due to their large size, epiphytic hab- it, ephemeral and seasonal flowering, and green fruits. Moreover, the fertile structures are born low on the plant and are overtopped by obscuring leaves (Fig. 3). However, even in a population that is out of season, examination of numerous plants will often reveal fertile structures. This effort is important. Both staminate and mature fruiting materials are essential for describing new species and often for identifying known species. Dramatic—though nearly microscopic — dif- ferences in floral and fruit structure between veg- etatively similar species are easily overlooked by the nonspecialist collector who may tend to “see” (and collect) only one common species of a genus in an area where three or four occur. Different species may occupy the same habitat on adjacent ridges or along different branches of a single stream. Differences in depth of lamina division, presence/absence and position of lateral costae, phyllotaxy, habit, and even subtle differences in lamina and petiole texture and color help reveal different taxa. What to do to them. The artful folding (ori- gami) of whole plants of large-leaved monocots may in itself be reward enough for a specialist in the particular family, but one can hardly ex- pect specialists in other groups or even generalist collectors to spend so much time on one gath- ering. The final specimen is usually better for data recovery when redundant material has been removed and the essential properly folded. Thus, the first need in collecting large-leaved monocots is to know what parts ofthe plant to collect. Most of the following comments pertain to vegetative parts because most of the difficulties stem from PES = em. Since the leaves of cyclanths are bilaterally symmetrical, they can be split down the middle (Fig. 4). In general, the whole petiole and a picce of stem with attached inflorescence or infructes- cence should be included. Even when whole or half leaves are collected, two sources of infor- mation are often obscured or lost: the depth of division of the leaf and an abaxial view of the blade. It is not sufficient simply to press leaves so that the division is visible, since it often splits deeper on drying. On large leaves, which must be split to fit the press, the point of division can be indicated by cutting a notch at this point and folding the leaf so that the notch shows (Fig. 5). For leaves small enough to press entire, the best way to avoid ambiguity is to cut off one leaf lobe at the point of division (Fig. 6). The label should briefly record this notching and lopping of lobes, ' Supported by so BSR-8508463 from the United States National Science Foundation. Many thanks also to John Myers for draft of the manu ne drawings and to G. Wilder and an anonymous reviewer for helpful comments on a ? Missouri l. Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. ANN. Missouni Bor. GARD. 74: 897-902. 1987. FIGURES |, 2.— ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 | p H / ba h | "ç I 9 , M j 1. Dense understory of Asplundia uncinata Harl. in the wet lowlands of northeastern Costa Rica. Plants ca. 1.5 m tall.—2. Evodianthus funifer (Poit.) Lindman, a very widespread root-climbing epiphyte. Leaves ca. 50 cm long. e.g., “notch marks point of division,” or “lobe cut off at point of division.” The presence/absence and position of lateral costae is taxonomically meaningful at species and higher levels. Although these costae are very con- spicuous on the lower surface, they are not visible at all from above. Thus, the blade should be folded to show at least the basal portion of the abaxial surface. This is true in all genera except Carludovica, and a few species of Asplundia, GURE 3. The F most ice auris fertile mace: when the staminodia are exserted, is ephemeral and hidden among the leaves. Leaf blades ca. 65 cm long. where features on the upper surface become im- portant. SPECIFIC (AND GENERIC) CONSIDERATIONS The following key to the 10 genera of Cyclan- thaceae is presented to facilitate discussion of generic characters and the requirements for col- lecting and mounting each genus. Pseudoludovia, no longer accepted, is not included in the key, nor are two new genera (R. Erikson, pers. comm.; Hammel & Wilder, in prep.), which require no special techniques. ere 4. Asplundia aii n ready for press- g.—A. Section to be discarded. — B. Notch to mar Fn of division. — C. Lateral costae. 1987] HAMMEL—COLLECTING AND MOUNTING CYCLANTHACEAE 899 KEY TO THE GENERA OF CYCLANTHACEAE WITH EMPHASIS ON VEGETATIVE CHARACTERS la. Leaf blades with the 2 major lateral costae running nearly the entire length of the blade, usually deeply = bifid nearly to the base; spadix a cylinder of rings or TE or both formed by alternating staminate and pistillate units; plants terrestrial, usually short-stemm Cyclanthus Leaf blades with the 2 lateral costae (if present) always w era well below the tip of the blade, entire to deeply divided; spadix a cylinder or sphere of tightly packed and variously connate to free, oh more typical staminate and pistillate flowers; plants epiphytic, lithophytic, or terrestrial, often long- em . Petioles mostly 1.5-3 m ied - blades palmately divided into 4 segments, the segments deeply toothed, the lateral costae and far removed from the margin; plants terrestrial; surface of € mature spadix Brrr i to reveal the pubes orange seed pulp and rachis; seeds Carludovica I Petioles mostly less than | m long; leaf € piem entire or bifid, the segments rarely toothed but then the lateral costae long and running in t N c argin; plants terrestrial, epiphytic, or litho- ete. phytic; surface of mature spadix not splitting irregularly, seeds flat or ter 3a. Leaves with 2 ta (oft Asplundia rarely in Dicranopygium, always in the monotypic Schultesiophytum). 4a. e thes mostly dispersed along the peduncle; fruits dehiscent by apical caps, fused at the bas Asplundia 4b. pem ena ne just below the spadix; fruits indehiscent. bruising and drying black; fruits completely separate U. WADE 5b. eiie remaining green; fruits connate in basal !⁄ or more u... Dicranopygiu w q . Leaves without conspicuous lateral costae. 6a. Pg clustered immediately below the sp ds nts lo 6b. Spathes stig rinse along the peduncle. ichous. 9a. ng-stemmed and openly eT usually climbing; stems and € leaves rous; fruits completely separate; seeds fla 7b. Plants zd short-stemmed and g on rocks along streams; stems and dry leaves smooth; fruits connate in basal er Evodianthus clumped, rarely climbing a short distance, usually Dicranopygium Leaf ae crenate at most, neve r bifid; fruiting spadix nearly smooth, the pistillate flowers connate throughout; plants lianas or short-stemmed epiphytes Ludovia 9b. Leaf blades of mature individuals bifid; fruiting spadix not smooth, the pistillate flowers completely or partly free; plants terrestrial or epiphytic, usually short- stemme 10a. Seeds mostly rounded at both ends Seeds bile oe appendages at both ‘ends 8b. ed spirally arra Sphaeradenia Stelestylis la. Petioles urs jie broadly channeled false petioles (sheaths) extending to the TO, leaves of climbing stems a undivided; mature fruits tan; s plants lianalike canopy clim spathes 8- Tho oracocarpus 11b. petioles present i in most species, the. false petioles narrowly channeled when pre debui 2-8; plants low trunk climbers or terrestrial A J. Ay a Fis p pyg t for over half the species in the family, and it is within the lower leaf surface applies especially to these genera. For the small-leaved species of Asplundia it is important to insure that more than one leaf is preserved and that both surfaces of the basal part of the leaf are visible. This is necessary in order to see adaxial scars at the distal end of the false petioles, characteristic of several species in subgenus Chaonopsis. In Asplundia and Dicrano- pygium, whether or not the sheath splits up into OF pte And ly orange fibers and the color and quality (dull vs. shiny) of its surface are necessary data points for iden- tifying or characterizing species. Here, specimens should preserve a portion of the stem together with the leaf at its point of attachment. Although Sphaeradenia consistently has only the median and no lateral costae, nothing is lost by preparing specimens of this genus (and of most others) as for Asplundia and Dicranopygium. In any case, for Ludovia, Sphaeradenia, and Ste- lestylis one should preserve part of the stem with attached sheaths in order to verify the distichous arrangement of the leaves and to show internode length (Fig. 6). Asplundia, Dicranopygium, and 900 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 FiGuRE 7. Carludovica specimen ready for press- e 2 sections to be discarded cut at point of div Sphaeradenia account for most of the species in the family so that the general rules for collecting these genera apply to most of the species one encounters. Carludovica needs special attention in con- nection with the very large size and unusual shape FIGURE 5. Asplundia specimen properly folded and of its mature leaves. In order to represent the mounted with string. essential features (the depth of the teeth and the FIGURE 6. Sphaeradenia specimen showing blade FIGURE 8. Carludovica specimen properly folded cut off at point of division (A) and distichous phyllo- and i ga with string, showing hastulae (A) on up- taxy (B). per surfac 1987] depth of the divisions) only one of the four leaf segments need be preserved. The best approach is to cut off the two lateral segments and one of the central segments at the point of division and then fold the blade so that the teeth and the adaxial surface of the basal portion of the leaf are visible (Figs. 7, 8), thus showing the presence/ absence of hastulae (Wilder, 1976). Sometimes even this reduced leaf must be split to fit on one sheet. Variation in petiole and peduncle length appears to be of little use for et arn species of Carludovica, but the length of these structures should at least be recorded in the field notes. Finally, for Carludovica, variation between species in shape of juvenile and immature foliage makes leaves from young plants worth preserv- ing. FIELD NOTES abit. Approximately 60% of the species of Cyclanthaceae are either strict epiphytes or root- climbers, which may eventually lose connection with the ground. Habit is quite diverse (and sometimes variable within species) in Asplundia and Dicranopygium. The longest-stemmed species are usually found only as climbers, some with shorter stems climb or grow free-standing, and the shortest-stemmed species never climb. Strict epiphytes occur only in Ludovia, Sphaera- denia, and Stelestylis, and terrestrial species oc- cur in all of these genera. Carludovica is always terrestrial; Cyclanthus rarely climbs. Only species of Dicranopygium grow on rocks along streams. For these reasons one should always make care- ful note of the habit: epiphytic, lithophytic, ter- restrial, or root-climbing. If the collector 1s cer- tain that a species is both terrestrial and climbing, it is very useful to write that in the notes, e.g., “apparently same species also climber here but not fertile." If the stem is not preserved, the leaf arrangement should be noted. The approximate length of the stem or height above ground for climbers (e.g., “climbing to 10 m") should also be recorded. Other d rini characters. Even though care has been taken to record on the specimen the depth of the us du it is also a good idea to record in the field notes the approximate range of this cl based on the plant or population collected, e.g., “leaves divided from !^—/." The leaf segments of all species of Carludovica and a few species of Asplundia develop teeth in the unopened leaf. Unfortunately, with age, drying, HAMMEL- COLLECTING AND MOUNTING CYCLANTHACEAE 901 or other trauma some large Asplundia leaves can split into teeth, which on herbarium sheets re- semble those arising developmentally. Thus, for Asplundia one should indicate whether such teeth originate developmentally or by trauma. As noted above, subtle differences in leaf and petiole surface textures and colors often separate species and should be recorded. In a few species of Asplundia and Dicranopygium the leaf folds are not adaxially keeled. Because all leaves on herbarium specimens appear to have variously keeled folds, it is best to record exceptions in the field notes. The cross-sectional petiole shape is less vari- able in the Cyclanthaceae than in certain genera of Araceae (Croat, 1985), but again this feature is difficult to assess using dried herbarium ma- terial and should be described for unusual cases. Most petioles are basically terete and barely to obviously flattened above (D-shaped) and may have a small median adaxial groove. In Thora- cocarpus and a few species of Asplundia the sheath reaches all the way to the base of the blade so the leaves lack a petiole. The petioles of ScAul- tesiophytum are asymmetrically D-shaped with one margin rounded and the other sharply an- gled. The color of the stem cross section may also be taxonomically significant; in certain species of Asplundia and Dicranopygium cut stems rapidly turn reddish brown om in- dicating high concentrations of tan Fertile structures. Rui n of cy- clanths are monoecious and protogynous. Their most conspicuous stage— when the staminodia are exserted—is also the most ephemeral. Vary- ing in size and color, the vermicellilike stami- nodia may harbor taxonomic structure but have not been used because they are so seldom seen and poorly did on drying. When an inflo- rescence with dia is collected, field notes should incline color and approximate length of staminodia. The number of spathes serves to distinguish taxa but is often difficult to count on herbarium material without damaging it. Field notes should mention the number, color, and texture of spathes. Most of these time-con- suming measurements can be delayed (and many more characters accurately preserved) by pre serving inflorescences in liquid. Dried pu of these extremely succulent parts so misrepre- sents the living condition that, except for well- known species, collectors are strongly urged to preserve inflorescences or parts thereof in 70% Et 902 ae ome | A ak 1d The color of be noted. They may be bite: red, yellow, or green at maturity. In Sphaeradenia the color of mature seeds is also variable and should be re- corded. MOUNTING THE SPECIMENS All of the foregoing discussion, especially on artful folding, is to no avail if the dried material is mounted upside down. Collectors always as- sume that their material will be mounted the way they pressed it, with the numbered side of the newsprint equal to the up side of the specimen. However, once the connection between news- paper and plant is broken, as it can easily be with so many steps between collecting and mounting, and as it always is in the end, the decision must be made: which side is up? Responsibility for this final important step rests upon the mounter. In the end, a liberal use of string and judicious use of glue can obviate even this decision. Gen- erally, one need glue down only the first layer of leaf fold and then tie the stem and petiole(s) at appropriate places (see Figs. 5, 8). In this way ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 the specimen can be unfolded partly or com- pletely, and both sides of the leaf can be easily examined Mounters, of course, will continue to receive material gathered by those who have missed, ig- nored, or forgotten this guide, or who are not artists. If parts are missing, or no mark records the point of division, nothing can be done. But when the leaves have been folded improperly, this guide is especially for the mounters, who with a little care can salvage the specimen and enjoy the art. LITERATURE CITED CROAT, T. B. 198 mens of Araceae. 252-258. DRANSFIELD, J. 1986 5. Collecting and preparing dien Ann. Missouri Bot. Gard. A guide to collecting palms. : 166-176. A guide to collecting Pandanaceae (Pandanus, Freycinetia, and Sararanga). Ann. Missouri Bot. Gard. 70: 137-1 WILDER, he ir 1976. Structure and development of leav 7 reference to other Cyclanthaceae and Palmae. Amer. J. Bot. 63: 1237-1256. MESOAMERICAN SISYRINCHIUM (IRIDACEAE): NEW SPECIES AND RECORDS, D NOTES ON TYPIFICATION' JAMES E. HENRICH? AND PETER GOLDBLATT? ABSTRACT This paper supplements our treatment of Sisyrinchium for Flora Mesoamericana in which 13 species are recognized. One new species is described, S. su and Costa Rica. Sisyrinchium trinerve, previously known from T alpinum, from Mexico (only Chiapas), Guatemala, from Andean South America, S. dimorphum Mexico, S. longispathum from Oaxaca, Mexico, and S. subcernuum from northern Mexico are reported in Mesoamerica for the first time. The type material for S. convolutum, S. tenuifolium, S. designated for each species. In preparation for a treatment of Iridaceae for Flora Mesoamericana (Henrich & Goldblatt, in press), it has become clear that previous re- gional treatments of Sisyrinchium are to varying degrees inadequate or erroneous. The same species are treated under different names in some floras and few of the species have been matched with their types. Notes are thus presented here on the typification of several widely recognized species to supplement the formal treatment. There is also apparently one new species in Mesoamer- ica, S. subalpinum, a yellow-flowered species of section Echthronema Bentham. This is a dwarf, high-altitude relative of the common S. tincto- rium. We also report for the first time the oc- TABLE 1. mandonii, S. tinctorium, and S. bogotense has been examined and a lectotype currence in Mesoamerica of the Andean S. tri- nerve, the northern Mexican S. subcernuum, the southern Mexican S. /ongispathum, and the pre- dominantly Texan S. dimorphum. Finally, the species previously identified as S. iridifolium Kunth in Guatemala and Honduras corresponds closely to the Oaxacan S. exalatum, a tall, branched species with narrow leaves and thick- ened roots sometimes swollen terminally. We recognize 13 species (Table 1) of Sisyrin- chium in Mesoamerica, which is regarded for the Flora as including all the territory from the southern Mexican states of Yucatán, Campeche, Tabasco, Quintana Roo, and Chiapas south ward to Panama. The species are numbered as they The species of Sisyrinchium recorded from Mesoamerica with their general distributions. S. chiricanum Woodson S. convolutum Nocca Mexico Pisas Guatemala, El Salvador, Costa Rica, Panam Chiapas, Mexico to Panama, possibly the north coast of South Ameri k p a kiya dimorphum R. Oliver exalatum Robinson & Greenman . Johnstonii Standley longispathum Conzatti mandonii und micranthum subalpinum See & Goldbl. subcernuum Pon Henrich & Goldbl. mb. & tenuifolium H tinctorium on trinerve Baker Bonpl. ex Willd. U.S.A. a Sy T" Guatemala southern m (Oaxaca, Chiapas), Guatemala, Hondur: Mexico POM Guatemala, Costa Rica Mexico (Oaxaca, Chiapas) Guatemala, Costa Rica, Panama, Peru, Bolivia Mexico to South America, West Indies Mexico (Chiapas), Guatemala, Costa Rica Mexico, Belize Mexico, Guatemala, Panam southern Mexico to ind and Venezuela Costa Rica, Colombia, Venezuela, Ecuador to Bolivia iS ! Support for this research from U.S. National Science Foundation grants DEB 81-19292 and BSR 85-00148 quad acknowledge ouri Botanical Garden ? B. AK , P.O. Box 299, St. Louis, Missouri 63166, U.S.A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. ANN. Missouni Bor. GARD. 74: 903-910. 1987. 904 appear in the Flora treatment. Species 1—3 belong to section Sisyrinchium (= section Bermudiana ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Bentham) and 4-12 to section Echthronema (Bentham, 1883). KEY TO SISYRINCHIUM IN MESOAMERICA la. Flowering stems unbranched, consisting of 1 long internode without cauline leaves, but sometimes with a terminal bract subtending the opposed spathes of the 1-3 apical inflorescence units (rhipidia) Apos rhipidia with a short stalk, more often sessile). s filiform wide; inflorescence units 1—3 T nie a to lanceolate, at least 1.5 m tuberous, fleshy (Fig. 1A); flowers blue to w .. 8. S. longispathum mm wide; Aue ha units ] or 2. cR 3. S. johnstonii 3b. Root ots pois rous OT nerd. but not tuberous (Fig. 2B. "DX flowers yellow. 4a. Flowering stems subterete and not winged; ovary included in the upper part of the spathes 7. S. trinerve 4b. Flowering stems flattened and winged; ovary exse 5a. Spathes (20-)25-50 mm long, the outer euh EE twice) than the inner 11. S. tinctorium 5b. Spathes 11-16 mm long, usually subequal. 6a. Plants 5-12 cm tall; roots slender but somewhat thickened (Fig. 3) .. 12. E subalpinum 6b. Plants 15-25 cm tall; roots fibrous (Fig. 2D) . Flowering stems EA of more than | internode, branched or at least with | or more cauline le c eav 7a. Roots fibrous or thickened and cylindric, sometimes with persistent hairs (Fig. 1B, 2. Is 5-15(-25) cm tall; capsules 3 mm long, globose 8a. Annua S. subcernuum 9: eig dé 8b. Perennials (15-)20-60 cm tall; capsules 4-5 mm long and globose, or 8-11 mm long a ob ovoid. 9a. Flowers blue; spathes 14-18 mm long ..... 9b. Flowers yellow, spathes 26-32 m . S. dimorphum Oa. Roots with persistent hairs throughout or at least near the base (Fig. 1B) 5. 10b. Roots smooth, not bearing persistent hairs 1C) 7b. Some roots tuberous and fleshy, either close to o lla. Basal leaves 3-8 mm ^u chiricanum OCURRE: S. exalatum m the p i (Fig. Y C). falca wide, narrowly to broadly eerie to 2a. Plants usually less than 30 cm high and with falcate leaves; ue strongly 3-lobed 6. sually with swollen fusiform us thickened from the base (Fig. 2A ore m high and with lanceolate elio capsules hae d; roots ceri if at "all. distant from the base (Fi js wide, linear; capsules not strongly 3- ne De 5-7 mm long .... 13b. Capsules oblong, 12-16 mm long . lobe llb. Basal Testa 0.5-2 m 13a. Capsules glo NOTES ON SELECTED SPECIES 1. Sisyrinchium dimorphum R. Oliver, Ann. Missouri Bot. Gard. 55: 397. 1969. TYPE: United States. Texas: Val Verde Co., San Felipe Springs, Del Rio, Warnock & Cam- eron 9894 (holotype, SMU). A species of Sisyrinchium from Chiapas and Guatemala sometimes identified as S. scabrum ate size with relatively short leaves 1-2 mm wide, a branched stem, and persistent fibrous leaf bas- es, distinctive in Sisyrinchium. It clearly belongs in section Sisyrinchium and has the blue (to white) campanulate perianth and connate (or nearly connate) filaments that characterize the group. The Mexican S. scabrum, typified by Schiede S. patinen exalatum —-— . 9. S. tenuifolium 10. S. mandonii 1020 (HAL), is similar in general morphology yet it is smaller in stature, has a smaller flower with an unusually small, sparsely pubescent ovary less than 1 mm long, and globose capsules ca. 2 mm in diameter, whereas S. dimorphum has a glabrous ovary ca. 2 mm long and larger, some- what obovoid capsules 4-7 mm long. 4. Sisyrinchium exalatum Robinson & Green- man, Amer. J. Sci. 50: 166. 1895. TYPE: Mexico. Oaxaca: Cuilapan Mountains, 7,000 ft., Smith 52 (not seen). Figure 1C When reported from Guatemala and Hondu- ras (Standley & Steyermark, 1952; Molina, 1975), Sisyrinchium exalatum was treated as S. iridi- folium Kunth. We concur in regarding it as dif- ferent from all other Mesoamerican species but it does not correspond with S. iridifolium. This 1987] FIGURE 1. Root systems n stem bases. Scale bars = 2 cm; top bar fo B. is a South American species, the type from Car- acas, Venezuela, which is similar to S. exalatum in leaf and stem morphology, but the roots are fibrous and not thickened as in the latter and the capsules are relatively small and nearly globose, rather than obovoid-ellipsoid and somewhat trigonous. In Mesoamerica Sisyrinchium exalatum oc- curs in Mexico (Chiapas), Guatemala, El Sal- vador, and Honduras. It grows on grassy to light- ly wooded slopes at elevations of 1,700-3,700 m Additional specimens examined. XICO. CHIAPA north end of San Cristóbal Las ae sap wa 6045 (F). EL SALVADOR. MORAZAN: Slopes of La Montañita, HENRICH & GOLDBLATT—MESOAMERICAN SISYRINCHIUM 905 —A. Sisyrinchium johnstonii. — B. S. chiricanum. —C. S. exalatum. 1,700 m, Williams & Molina R. 10429 (F, MEXU, ICH, MO). aar EMALA. HUEHUETENANGO: between Tojqu iá and Caxín bluff, Steyermark 5014 1 (F); Sierra Cuchumatanes, Sur 1243 (F) anto Tomás, COMAYAGUA: abierto de La Piramide, Molina R. 14357 F). 6. Sisyrinchium convolutum Nocca, Ticin, Hort. l . TYPE: Guiana “Cap. B. Spei," cultivated in “Hort. Parolin,” collector un- known (lectotype, BASSA, here designated). Figure 2A Sisyrinchium vg crois (Baker) Standley & Stey- eld Mus. Nat. Hist., Bot. Ser. 23: 39. 1944: s of Guatemala, Fieldiana, Bot. 24: 906 73. 1952. S. alatum var. guatemalense Baker, andbk. Irideae 180. 1892. TYPE: none cited nor locaied but authentic material at K, annotated by Baker Sisyrinchium convolutum is one of the more common Mesoamerican species of Sisyrinchium. It can generally be recognized by its moderate height, more or less falcate leaves, winged and branched stems, and large, inflated, trigonous capsules 8-10 mm in diameter. The capsules are glabrous, in contrast to the related and some- times similar S. tenuifolium, in which they are sparsely pubescent. The flowers are yellow and relatively large with tepals to 15 mm long, and the spathes of the inflorescence are distinctively broad, giving it a somewhat inflated appearance. The roots are fleshy and tuberous (Fig. 2A Typification of Sisyrinchium convolutum has posed some difficulty. The illustration in the pro- tologue matches the species to which the name has been applied here and elsewhere, except for the swollen, almost bulbous base and the roots figured as forming a fibrous mass. A specimen in the herbarium at Bassano del Grappa, Vicen- za, where Nocca’s types are believed to be housed, matches closely the illustration but lacks a base and roots. Given the stylized depiction of these organs, it seems likely that the artist drew them from imagination. We have chosen the specimen rather than the illustration as lectotype. Sisyrinchium convolutum was confused with S. alatum, a strictly South American species, by Baker (1892) amongst others. Baker used the name S. alatum var. guatemalense for the plant, as did Standley & Steyermark (1952) in the Flora of Guatemala and Molina (1975) in an enumer- ation of the plants of Honduras. Sisyrinchium alatum is probably not closely related to S. con- volutum and does not have fleshy tuberous roots as do the latter and related species. 7. Sisyrinchium trinerve Baker, J. Bot. 14: 267. 1876. Sisyrinchium bakeri Klatt, Abh. Na- turf. Ges. Halle 15: 378. 1882, nom. superf. pro S. trinerve. TYPE: Bolivia. La Paz: vicin- ity of Sorata, Mandon 1218 (lectotype, K, here designated); syntypes: Mandon 1220 (K); Pearce 87 (K). Figure 2B Known until recently only from the high An- des of Bolivia, Peru, Ecuador, Colombia, and Venezuela, Sisyrinchium trinerve has now been found in the páramo of Costa Rica above 3,000 m. It is readily recognized by its long, un- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 branched, nearly terete stem; rigid, narrow, pre- dominantly three-veined leaves; yellow perianth; and ovary at least partly included in the spathes. A lectotype has been designated, since the pro- tologue includes three collections, none desig- nated as a type (Baker, 1876), and no herbarium is mentioned, although the Kew Herbarium is assumed to house the specimens seen by Baker. The collection Mandon 1218 was chosen, since it comprises the best-preserved and most com- plete material. Additional specimens examined. CosTA RICA. LIMON: Cordillera de Talamanca, Kamuk massif, 3,000- 3,300 m, a & Herrera Ch. 29375 (MO). CAR- AGO-SAN JOSE: Cerro de la Muerte, paramo, along stream, Heithaus 239 (MO). 8. Sisyrinchium longispathum Conzatti, Flora Taxonomica Mexicana 2, 2: 124. 1947. TYPE: Mexico. Oaxaca: Conzatti 4203 (not seen). Known from only a handful of collections, all from mountainous areas of the southern Mexi- can state of Oaxaca, Sisyrinchium longispathum has been recorded from a single site in Chiapas, Breedlove 27156, 22 km north of Tuxtla Gut- tierrez. The species appears to be restricted to limestone outcrops and cliffs. Sisyrinchium lon- gispathum is distinctive among the Mesoamer- ican species of Sisyrinchium in having filiform leaves and 2-3 inflorescence units per flowering stem, all crowded at the apex and each subtended by a short leaf. The yellow flowers are typical of section Echthronema. Sisyrinchium longispa- thum has no close relatives among the Mesoam- erican species of Sisyrinchium. 9. Sisyrinchium tenuifolium Humb. & Bonpl. ex Willd., Enum. Pl. Hort. Berol. 2: 691. 1809, Hort. Berol. 2: 691. 1809. TYPE: Mexico. Without precise locality, Humboldt & Bon- pland s.n. (lectotype here designated, B— Herb. Willdenow — only microfiche seen). Sisyrinchium tenuifolium is one of the smaller species of the genus and can usually be recog- nized by its branched stems, narrow basal leaves 0.5-2 mm wide, and swollen tuberous roots. The bright yellow flowers are more or less typical for section Echthronema. It is particularly variable, with plants ranging from slender and unbranched with very narrow leaves to robust with broad leaves and many-branched stems. The latter forms resemble S. convolutum fairly closely and 1987] FiGurE 2. Root systems and pa bases. D. S. per ee Scale bar = 2c can be distinguished by their minutely pubescent ovary and smaller capsule. The specimen in the Willdenow Herbarium at Berlin is chosen as the lectotype. Other Hum- boldt and Bonpland specimens of this species may be duplicates of the type collection and thus isolectotyes, as is the illustration in the Hortus Berolinensis. 10. Sisyrinchium mandonii Baker, J. Bot. 14: 269. 1876. TYPE: Bolivia. La Paz: vicinity of Sora- ta, Rancha de Cochipata, Mandon 1217 (lec- totype, K, here designated; isolectotypes, B, not seen, MO photo); syntypes: New Gran- HENRICH & GOLDBLATT —MESOAMERICAN S/SYRINCHIUM —A. Sisyrinchium convolutum. —B. S. trinerve. — C. S. mandonii. — ada (Brazil), Jurgensen id o New Gran- ada, Purdie s.n. (K). Fig Sisyrinchium mandonii has distinctive small, swollen, tuberous roots (Fig. 2C); branched stems; relatively narrow leaves 1-2 mm wide; and un- usual large, ellipsoid capsules. It is predomi- nantly Andean, represented by a few collections from Mesoamerica (Steyermark 30513, Guate- mala; Allen 698, Costa Rica; Woodson & Schery 427, Panama), all from relatively high altitudes. It was first reported for Mesoamerica by Wood- son (1945) from Panama. Of the three collections cited by Baker in the protologue, Mandon 1217 is selected as the lec- 908 totype as it is in good condition and Baker an- notated it as the "Type." There is no indication whether Baker saw any duplicates of this collec- tion but this 1s certainly possible. The two other collections cited, both at Kew, appear to be the same species, although they are from New Gra- nada in eastern Brazil where Sisyrinchium man- donii does not grow. Mislabeling seems likely here. 11. Sisyrinchium tinctorium Kunth, Nov. Gen. Sp. 1: 324. 1815. TYPE: Venezuela. T.F. Amazonas: banks of the Orinoco near Es- meralda and the confluence of the Sodo- monis, Humboldt & Bonpland s.n. (holo- type, su Lag id Kunth, Nov. Gen. Sp. 1: 323. . TYPE: Colombia. Cundinamarca: Bogotá, En. Sins between Suba and Suacha, Humboldi & Bonpland s.n. (holotype, P). — Sisyrinchium tinctorium is a common and well- known species in Mesoamerica. It also occurs in northern South America but is apparently less common there. It is distinguished by unbranched and often broadly winged flowering stems with terminal spathes, pale yellow flowers, and pen- dent, more or less pyriform capsules. Specimens in fruit are collected most often. The leaves have a characteristic thin, almost membranous tex- ture when dry, and the roots are fibrous. Me- soamerican collections match the type of a sec- ond species, S. bogotense, particularly well. Our examination of the types of this and the Vene- zuelan S. tinctorium showed the two to be con- specific. The type specimens of S. tinctorium have comparatively narrowly winged stems and cap- sules smaller than most collections ofthe species. However, a few similar specimens have been col- lected in Mesoamerica, and variation is contin- uous from typical S. tinctorium to typical S. bo- gotense. A diminutive species clearly allied to Sisyrin- chium tinctorium is recognized here as S. sub- alpinum, which is shorter and has narrower leaves and smaller capsules than those of S. tinctorium. 12. Sisyrinchium subalpinum Henrich & Gold- blatt, sp. nov. TYPE: Costa Rica. Limón: Cor- dillera de Talamanca, Atlantic slope, Ká- ,30 d Herrera Ch. 29294 (holotype, MO). Fig- e 3. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 Plantae 5- 12 cm altae, radicibus aliquantum in- m longis iai pr filamentis 1.5 mm longi infra liberis antheris ca. 2 mm longis, ramis styli ca. 1.5 mm longis, capsulis nutantibus glabris. Plants 5-12 cm tall; the roots slender, some- what thickened but cylindric, not fleshy or tu- berous. Leaves several, all basal, '^ to as long as the flowering stems, 1-2 mm wide. Flowering stems less than | mm wide, unbranched, com- prising | internode, flattened but barely winged; spathes 11-16 mm, subequal or the outer to !^ longer than the inner. Flowers small, yellow, stel- late; tepals 4-5 mm long, subequal; filaments 1.5 mm long, free at least above (not seen below); anthers ca. 2 mm long. Ovary about 3 mm long; style branches ca. 1.5 mm long, slender, extend- Ing between the stamens. Capsules nodding, ma- t seen, these probably somewhat pyr- ton and glabrous; seeds unknown. Sisyrinchium subalpinum has been recorded at high elevations in Mexico (Chiapas) and in Guatemala and Costa Rica. It 1s restricted to paramos and moist, high, pine forests. Additional spiel examined. MEXICO. CHIAPAS: . Male near Porvenir, 3,200 m, Matuda 4629 (MO). GUATEMALA. SAN MARCOS: along quebrada Canjula be- tween Sibinal and Canjula, Volcán oq 2,200- 2,500, Steyermark 35970 (F). Costa RICA. : within 200 m of the summit of Cerro Chirriposillo. 3,400 m, Weston 1574 (MO 13. Sisyrinchium M (Bickn.) Henrich & Goldblatt, comb. nov. Hydastylis sub- cernuus 2 Bull. Torrey Bot. Club 27: 385. 1900. TYPE: Mexico. Baja California Sur: Sierra i Laguna, Brandegee s.n. (ho- lotype, CAS). Figure 2D. Sisyrinchium subcernuum belongs to what may be called the S. tinctorium complex, the species of which share similarly textured leaves, un- branched and winged flowering stems, and drooping capsules. The type is from Baja Cali- fornia Sur, and several more collections are known from higher elevations in Mexico. A few ofthe collections from Belize match the northern Mexico plants closely and must be treated as the tepals about 6 mm long, and small capsules 6 mm long and 3 mm wide. The outer spathe of 1987] FIGURE 3. HENRICH & GOLDBLATT — MESOAMERICAN SISYRINCHIUM Type specimen of Sisyrinchium subalpinum (scale 909 MISSOURI BOTANICAL GARDEN HERBARIUM N° 3212055 olo! o IS rin ehiu. ubo pin | — r Sisyrinehui gu end Goldblatt Ref Sisyrinchium subalpinum Henrich ¢ Goldblatt qc ang by jJ. E. Henrich & P. Goldblatt 1986 un Botanical Garden COSTA RICA IRIDACEAE Sisyrinchyum LIMON: Cordillera de Talamanca, Atlanti slope nen uk massifs í paramo north east the main Kamuk peak; elev 3000- $300 m; 9°16'-9°17'Ne 83*00'-03*02'W. Blechnum-shrub association in Chusquea- Hyper icum pårano. Flowers yellow aoe 19 Ser 1984 G. Davidse & G. Herrera 4 (MO) MISSOURI BOTANICAL. GARDEN HERBAR TUM — 5 cm). 910 the inflorescence is longer than the inner, some- times as much as twice as long. Additional specimens eS BELIZE. CAYO: near Millionario, , roadside, Gentry 7694 (MO); between Millionario and ren Dwyer 10821 (MO); vicinity of Cuevas, south of Millionario, ca. 1,900 m, Croat 23602 (MO LITERATURE CITED BAKER, J. G. 1876. New Aristeae and Sisyrinchia. J. B 69. . Handbook of the Irideae. George Bell Sons, London. BENTHAM, G. 1883. Irideae. /n G. red & J. D. ooker, Genera Plantarum 3: 681-710. Bei E. P. 1900. Studies in ele hium VIII. Sisyrinchium californicum and related species of the neglected genus H ydast ylus. Bull. Torrey Bot. Club 27: 373-387. HENRICH, J. E. & P. GOLDBLATT. Iridaceae. Jn G. Da- lees et al. (editors), Flora Mesoamericana, Vol- me 1 (in press). ia F.W. 1882. Ergänzungen und Berichtigungen ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 zu Baker's Systema Iridearum. Abh. Naturf. Ges. Halle 15: 337-404. KuNTH, C. S. 1815. Voyage de Humboldt et Bon- pland. Sixiéme Partie. Botanique. Nova Genera et Species Plantarum (= Novae Genera et Species Plantarum quas. Latine-Allemande, Paris. Gide Fils, MOLINA, R. A. 75. Enumeración de las sies de Honduras. Ceiba 19: 1- NOCCA, 1800. Ticinensis Horti Academici Plantae T I . Sisyrinchium dimorphum pis rege a new species from Texas and Mexi . Missouri Bot. Gard. 55: 397. eon B. L. & J. M. GREENMAN. 1985. New and noteworthy plants chiefly from Oaxaca collected by Messrs. C. G. Pringle, L. C. Smith and E. W Nelson. Amer. J. Sci. 50(296): 150-168. STANDLEY, P. C. & J. A. STEYERMARK. 1944. Studies of Central png iet plants. M Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 31-109. 1952. aoe In Flora of Gua- temala. _Fieldiana, Bot. 24: 159-178. WOODSON 45. Iridaceae. /n Flora of Panama. Ann. Ve Bot. Gard. 32: 34-42. NOTES THE MESOAMERICAN NEOMARICA (IRIDACEAE), N. VARIEGATA HENRICH & GOLDBLATT, COMB. NOV. The genus Neomarica (Sprague, 1928) com- prises some 15 species of evergreen herbs of trop- ical forests and woodlands. Ravenna (1976) ap- parently merged it with Trimezia Herbert, a treatment we do not accept. The genus is dis- tinctive in having a broadly winged flowering stem, the branches ofthe inflorescence often sub- sessile and crowded at the stem apices, and a single large cauline leaf subtending and extending well beyond the inflorescence. Neomarica is cen- tered in eastern Brazil but occurs widely in South America, with a single species extending through Mesoamerica into southern Mexico. In several regional floristic treatments for countries in Me- soamerica the native Neomarica has been treated as N. gracilis (Herbert) Sprague. The type of N. gracilis, a figure in Curtis' Botanical Magazine (66: tab. 3713. 1839), is believed to have been found in Brazil and it matches an eastern Bra- zilian species. It does not correspond with the Mesoamerican species in morphology of inflo- rescence, flower, or seed. The single Mesoamerican species of Neomar- ica was validly described in 1843 by Martens & , Galeotti as Marica variegata, based on Galeotti 5370, collected in Veracruz, Mexico. The type at the Herbier du Jardin Botanique National de Belgique (BR) matches other collections of Neo- marica from Mesoamerica in their subsessile in- ANN. MissouRi Bor. GARD. 74: 911. 1987. florescence spathes and flower morphology. Var- iegata 1s the only epithet available for the Mesoamerican Neomarica and the combination is made below. Neomarica variegata (Martens & Galeotti) Hen- rich & Goldblatt, comb. nov. Marica var- iegata Martens & Galeotti, Bull. Acad. Roy. Sci. Belgique 10: 112. 1843. TYPE: Mexico. Veracruz: Zacuapan, Galeotti 5370 (BR, ho- lotype). LITERATURE CITED Martens, M. & H. G. GALEoTTI. 1843. Enumeratio synoptica plantarum phanerogamicarum ab H Galeotti in regionibus Mexicanis collectarum. Bull. cad. Roy. Sci. Belgique 10: 110-134. RAVENNA, P. 76. Neotropical species threatened and endangered by human activity in the Irida- Extinction is Forever. New York Botanical Gar- , New York. SPRAGUE, T. A. 1928. Marica and Neomarica. Kew Bull. 1928: 278-281. —James E. Henrich and Peter Goldblatt, B. A. Krukoff Curator of African Botany, both at Mis- souri Botanical Garden, St. Louis, Missouri 63166, U.S.A TWO NEW XYRIS (XYRIDACEAE) FROM THE AMAZON BASIN OF BRAZIL Among undetermined Xyridaceae sent to the senior author for definitive treatment during 1980-1982 are two from northern Brazil that we agree are new species to be added to the flora of Xyris pectinata Kral, Smith & Wanderley, sp. nov. TYPE: Brazil. Amazonas: Estrada Transamazónica, campina aberta, terreno arenoso, Proj. RADAM, | June 1976, T. Bahia 35 (holotype, INPA; isotypes, US, VDB). Figure 1. erba perennis densicaespitosa, tenella, glabra; ra- dices graciles. Folia line nearia, 4-6 cm longa, erecta vel sc lora inae satis, lateribus valde longitudine ideo n vel Intense ferrugineae, me ibus in lamai na ne ada longam fascientes, infime as dilatatae. ane retes, filiformes, r ter torti, i 5 mm crassi, olivacei, distaliter iu bicostati, costis laevibus. ipie subglobosae is late obovoideae, 4.5-6 mm longae, e, pluriflorae, d at- n tae, la t atae, suborbiculatae, aut reni- formes ae, convexae et leviter carinatae, obtusae vel subt fir ad apicem utrinque erosae et ciliatae, scariosae, minute tuberculato-rugulosae, a medio ad basim lto crassiores, nitidae, brunneolae, marginibus effuse et pectinate rigidofimbriatis; arca dorsalis ovata, ca. 2-2. , glau la la- teralia ca. ^ connata, ca. 2 lobis acutis scariosis, ala carinali angusta, integra Laminae petalorum bov , Ca. mm lon- gae, ad apicem rotundatae, laceratae, luteolae. Stami- m longae, loculis parallelis dis- tinctis; filamenta ca. 0. 5 mm longa. Capsula dorsali- ventraliter compressa, oblongo-cylindrica, tenuissima, 1.2-1.3 mm longa; placenta basalis. Semen solitarium, lenticulariter oblongo-ellipsoideum, 1-1.2 mm lon- gum, translucidum, pallide luteo- brunneolum, longi- tudine subtiliter striatum. Delicate, smooth, cespitose perennial; roots slender. Leaves linear, 4-6 cm long, erect or ANN. Missouni Bor. GARD. 74: 912-916. 1987. somewhat spreading, longer than the scape sheaths; blades 3-5 times longer than the sheaths, plane or slightly twisted, 0.9-1.2 mm wide, lon- gitudinally distinctly multinerved, strongly flat- tened, ferrugineous to olive-green; apices con- tracted, incurved-acute; margins thickened, minutely ciliate; sheaths carinate, with carinas minutely red ciliate, incrassate, the sides strongly longitudinally nerved, pale to deep red-brown, the margins gradually converging into the blade, at apex producing an acute ligule 0.5 mm long, below gradually dilating. Sheaths of scapes lax, mostly open, twisted, shining toward the base, carinate at the middle, with blades either similar to those of principal leaves or shorter. Scapes subterete, filiform, + spirally twisted, 1.2-2 dm high, ca. 0.4—0.5 mm thick, olivaceous, distally with sharp, smooth costa. Spikes subglobose to broadly obovoid or short-cylindric, 4.5-6 mm long, several-flowered, short-attenuate; sterile bracts 2(-4), 2-2.5 mm long, with dorsal areas linear and as long as bract, the lowest pair oblong; fertile bracts tightly spirally imbricate, broadly ovate, obovate, suborbicular or reniform, ca. 3 mm long, convex and slightly carinate, obtuse to subtruncate at apex on either side, erose, scari- ous, minutely rugulose-tuberculate, much thick- er from the middle to the base, shining, brown- ish, with margins effusely, dini and rigidly fimbriate; dorsal area ovate, ca. 2-2.5 mm long, gray-green. Lateral sepals ca. p connate, ca. 2 mm long, inequilateral, the lobes acute, scarious, the carinal keel narrow, entire. Petal blades nar- rowly obovate, ca. 1.5 mm long, apically round- ed, lacerate, yellow. Staminodia somewhat re- duced, bibrachiate, the branches at apex short-plumose with moniliform hairs. Anthers oblong, ca. 0.3-0.4 mm long, the locules parallel, distinct; filaments ca. 0.5 mm 1 ids: lenticularly pres ellipsoid, long, filling capsule, translucent, pale yellow- brown, finely longitudinally striate. Additional material examined. BRAZIL. AMAZONAS: Transamazonas às anes y,53km W ee River; bus na" regio MARE in open campina of white 27 june 1979, Cleofé E. Calderón, O. P. a & J. Guedes 2696 (INPA, US, VDB); Muni- cipio de Borba, acima de Terra Preta, campina do rio 1987] Surubim, afluente do rio Abacaxis, dos S, 58?*33'W. Campina aberta, areia branca. Erva de 10 cm de altura; flores SAN 4 July 1983, C. A. d 4026 (INPA, NY, VDB). This species is easily distinguished by its fringe of strong though slenderly tapering rigid bristles on the margins of the fertile bracts, nearly sep- arate anther locules, and reduced staminodial condition. It is unusual in its particularly small and thin-walled capsule, this tightly filled by a single large seed. Xyris calderonii Kral, Smith & Wanderley, sp. nov. TYPE: Brazil. Amazonas: Transama- zonas Highway, 9 km W of Rio dos Pombos, ca. 1.5 km E of Igarapé dos Pombos, and ca. 64 km E of the Aripuanà. Common in white sand campina, flowers yellow, 18 June 1979, Cleofé E. Calderón, O. P. Monteiro & J. Guedes 2549 (holotype, INPA; isotypes, US, VDB). Figure 2. Planta humilis, annua, praeter inflorescentiam gla- bra. Radices filiformes. Folia linearia, solum basal à carinatae, porphyreae, pluricostatae, alesis um sca- osae £ aminas gra- curtam latam fascientibus, infime pean ee expansae, apertae, foliorum aut brevibus. Scapi che ree plus minusve spiraliter torti, 5-10 cm m cras- si, distaliter leviter multicostati, ied levibus Spicae subglobosae vel late ovoideae, 3-5 mm longae, pluri- florae, obtusae, involucratae. Bracteae steriles 2-4, m "0 c vel acuminatum, profunde Ideen: areis dorsalibus valde papillosis, vulgo sine laminis. Bracteae fertiles late ovatae vel subor- biculatae, 2.5-3 mm longae, valde rotundato-convex- ae, villosociliati, areis dorsalibus ovatis, valde granu- NOTES 913 lato-papillosis. Sepala lateralia libera, oblonga vel ovata, 2-2.5 mm longa, valde inaequilatera, ala carinali lata, a integ d istante ciliata, picem versus lacerata. Laminae petalor late ob atae, ca ngae, luteolae, ad api late ro- MAR don ca. 0.5 mm lon longa, pallide brunneola, plus minusve reticulata. Low annual, smooth except for the inflores- cence. Roots filiform. Leaves linear, strictly bas- al, 3-7 cm long, spreading flabellately, com- monly longer than the scape sheaths. Leaf blades 2-4 times longer than sheaths, 0.5-1 mm wide, flat, straight, longitudinally few-costate, flattened from base to apex, brown to yellow-green, the tips abruptly narrowed, incurved-acute, the mar- gins entire, not thickened; sheaths carinate, brown, many-ribbed, scarious except for the ribs, with the thin edges stramineous, gradually nar- rowing into the blades or apically producing a m scarious, broad ligule, gradually dilating base, the edges entire. Sheaths of scape lax, Sici open, straight, carinate, with blades similar to those of foliage leaves or shorter. Scapes subterete, filiform, + spirally twisted, 5-10 cm high, 0.4-0.5 mm thick, distally with many low, smooth costae. Spikes subglobose to broadly ovoid, 3-5 mm long, several-flowered, obtuse, involucrate. Sterile bracts 2-4, xubdecussite: vil- lous-ciliate, the lowermost pair foliaceous, rigid, 2-5 times longer than the spike, lanceolate to oblong, 2-3 mm long, carinate, the dorsal areas linear, green, with blades similar to those of fo- liage leaves but triangulate; inner pair (if present) mm long, strongly rounded-convex, villous-ciliate, the dorsal areas ovate, strongly granular-papillose. Lateral sepals free, oblong to ovate, 2-2.5 mm long, very inequilateral, with keel broad, proximally entire, at middle distantly ciliate, toward apex lacerate. Petal blades broadly FIGURE 1. Blade and sheath junction.— e. blade, stamen, staminodial min j. See FIGURE 2. mid bla bract.—f. Petal blade and stamen.—g. Stylar Xyris pectinata i R. Bahia 35).—a. Habit sketch.—b. Leaf tip.—c. Sector of mid blade. eaf base. — f. Spike. — g. Fertile bract. d. —d. — h. Lateral sepals.—i. Stylar apex, petal Xyris calderonii Kral, Smith & Wanderley (Calderon et al. 2549).—a. Habit sketch. —b. Leaf at de.— c. Leaf blade-sheath junction, side (left) and ventral (right) views. — — d. Mature spike.— e. Fertile r apex, staminode.—h. Lateral sepal, capsule. — i. Seed. 914 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 1987] NOTES 915 916 obovate, ca. 3 mm long, yellowish, broadly rounded and strongly erose at apex. Staminodia bibrachiate, the branches sparsely long-penicil- late. Anthers oblong, adi ca. 0.5 mm long; filaments ca. 0.8-1 mm long. Mature capsule broadly obovoid, a, ca. 1 mm long, the placenta basal. Seed broadly ellipsoid, ca. 0.3 mm long, pale brown, + reticulate. There is no question that the affinities of this little plant are with X. u/eana Malme; in the production of narrow leaf blades it is most sim- ilar to var. angustifolia Lanj., which also some- times produces long-tipped basal bracts. How- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 ever, the lateral sepals are smaller and show a somewhat different keel configuration; the dorsal areas are consistently long excurrent to produce acicular blades several times longer than the sub- tended spike. The scapes are uniformly terete. — Robert Kral, Herbarium, Department of Biol- ogy, Vanderbilt University, Box 1705, Station B, Nashville, Tennessee 27235, U.S.A.; L. B. Smith, Department of Botany, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; and Sra. Dra. Maria das Graças de Lapa Wanderley, Instituto Botánica, Caixa Postal 4005, 01000 Sao Paulo, razil A NEW SPECIES OF PIPER (PIPERACEAE) FROM CENTRAL AMERICA Piper calcariformis M. Tebbs, sp. nov. TYPE: Costa Rica. Alajuela: Finca Los Ensayos, NW of Zarcero, 850 m, Croat 43546 (ho- lotype, MO; isotype, BM). Figure 1. , Frutex I- 2 m altus. Folia oblongo- ‘ovata apice acuta Petiolus va- ginatus spica pendula, bracteis triangularibus, basin versus longe calcariformibus Shrub 1-2 m high, the stems shortly pubescent. Leaves 16-24(-28) cm long, 10-17(-21) cm wide, ovate-oblong, glabrous or with short sparse hairs on upper surface, pilose on veins beneath, the apex acute, the base cordate and sometimes with one lobe slightly longer than the other; secondary veins 4—6, arcuate-ascending towards apex; pet- iole sheathing, pubescent, 4—9 cm long. Inflores- cence pendulous, 10-18 cm long, 7-10 mm wide in fruit, the peduncles 2-4 cm long. Floral bracts 2 mm long, triangular, sparsely to densely pu- bescent with lower part elongated into a long spurlike process. Anthers 0.8-1 mm long, de- hiscing laterally; filaments 1-1.2 mm long. Style 3-4 mm long; stigmas 3, recurved. Fruit 1-1.5 mm long, round-oblong Additional specimens examined. | PANAMA. BOCAS DEL TORO: McPherson 7362, 8658 (MO). CHIRIQUÍ: ANN. MissouRi Bor. GARD. 74: 917-918. 1987. aiias ge Vodicka 5519 (MO); Correa et al. 2113 (MO). VERAGUAS: Croat 27695, 27726 (MO); Mori & Kallunki 2586, 3891, 3896 (MO). Distribution. In moist forest from 400 to 1,200 m in the Cordillera Central in Costa Rica and in the Serranía de Tabasara in Panama. Only one collection from Costa Rica, the majority of specimens coming from Panama. Piper calcariformis is most closely related to P. sagittifolium C. DC. and shares similarities of inflorescence, bract and fruit shape. However, it can be separated easily from the latter by its ovate- oblong as opposed to sagittate leaves, its much longer, pendulous inflorescence, and by its styles with three stigmas rather than two as in P. sagitti- folium. Both of these species can be separated from the rest of Piper by their distinctive inflo- rescences with unusually shaped bracts. They are related to the large-leaved pipers with long pen- dulous inflorescences of section Macrostachys Miquel. — Margaret Tebbs, Department of Botany, Brit- ish Museum (Natural History), Cromwell Road, London, England. FIGURE 1. ANNALS OF THE MISSOURI BOTANICAL GARDEN MT. Lu Piper calcariformis (Croat 43546). — A. Habit. — B. Bract. — C. Fruit. — D. Stamen. [Vor. 74 A NEW SUBSPECIES OF QUARARIBEA FUNEBRIS (BOMBACACEAE) FROM NICARAGUA Recent monographic work has clarified the re- lationships among members of Quararibea Au- blet and Matisia Humb. & Bonpl. (Bombaca- ceae) in the northern Neotropics (Alverson, 1986). Taken together, these two genera consist of at least 60 species of trees found in wet or moist, primary forests from central Mexico to Brazil. Fieldwork in Mexico, Nicaragua, and Costa Rica has resolved the taxonomy of some of the more problematic species belonging to Quararibea sensu stricto (subgenera Archiquararibea and Lexarza of Vischer, 1920), including Quararibea funebris (Llave) Vischer. This widespread species, named from Mexican material (La Llave & Le- xarza, 1825; Standley, 1923), includes two sub- species in Nicaragua, subsp. funebris and subsp. nicaraguensis. The typical subspecies occurs in moist to wet lowland and mid-altitude forests from central Mexico to northwestern Costa Rica, including forests at altitudes of 300-800 m with- in Nicaragua. The diminutive a first described here, occurs above 1,200 m in high- land forests of a restricted area of s ia A Nicaragua. Quararibea funebris (Llave) Vischer subsp. ni- caraguensis Alverson, subsp. nov. TYPE: Nicaragua. Matagalpa: Pon "gel María de Ostuma, Cordillera Central de Nicara- gua, 1,400 m, 18 Jan. 1965 (fr), L. O. Wil- liams, A. Molina R., T. P. Williams, D. N. Gibson & C. Laskowski 27978 (holotype, WIS; isotypes, F, NY, US). Figure 1. rbor, 5-25 m alta. Folia obovato-elliptica, 7-22 cm longa, 2-10 cm lata, pilorum caespitibus densis infra in venarum secundarium axillis exceptis glabrata 16-22 mm longa, 24(-30?) thecis in apice fasciculatis. Stylus 17-26 mm longus. Fructus drupaceus, subglo- bosus, in sicco 20-25 mm longus et 14-18 mm dia- metro, calyce persistenti accrescenti in sicco rugoso et 11-13 mm longo per 3—2 longitudinis inclusus. Sub- species typicae similis sed praeter pedicellos saepius longiores in omnes partes minor. Tree, 5—25 m tall, 25-30 cm dbh, monopo- dial; trunk smooth and slightly fluted; branches verticillate, diverging horizontally from trunk and ANN. Missouni Bor. GARD. 74: 919-922. 1987. drooping along distal half; bark pale and rela- tively smooth, not conspicuously peeling; stip- ules decid narrowly triangular, 2-5 mm long, 1 mm wide, densely lepidote with fimbriate-pel- tate trichomes. Leaves simple (unifoliolate?), al- ternate, entire; petioles 7-18 mm long, including the inconspicuous proximal and distal pulvini, green, densely lepidote; b/ades entire, obovate to elliptic, the apex acute to acuminate, the base acute to rounded, 7-22 cm long, 2-10 cm wide, dark green, lustrous and glabrate above, medium green, duller, and glabrous to sparsely pubescent below, firm-chartaceous; veins prominent below; secondary veins 5—7 per side, pinnate, arched, loosely brochidodromous, bearing conspicuous domatia in their axils; tertiary veins reticulate. Flowers solitary or few together; pedicels 11-15 mm long, densely puberulent with pale yellow- ish-brown echinate-stellate trichomes, bearing 3 bracteoles, these broadly triangular, ca. 1.5 mm ong, 1.5 mm broad at base; calyx infundibuli- form, irregularly lobed at summit, 9-11 mm long, pale green, densely pubescent with pale stellate- echinate trichomes without, densely sericeous with long, pale, ascending trichomes within, the apical lobes bluntly triangular, to 3 mm long; petals spathulate, recurved at anthesis, 15-16 mm ong, 5-6 mm wide, white when fresh, usually becoming sepia-brown when dried, moderately to densely pubescent with pale, lax, stellate tri- chomes on both sides; staminal column cylin- dric, dilated towards the toothed summit, 16-22 mm long, 1.5 mm diam., densely pubescent with pale, stellate trichomes for most of its length, becoming puberulent towards apex, bearing 24(- 30?) thecae, the 5 apical teeth inconspicuous, ca. 0.5 mm long; thecae distinctly paired, ca. 2 mm long, 1 mm wide; style filiform, 17-26 mm long, exceeding the staminal column by 1-2 mm, to- mentose; stigma capitate, ca. 2 mm diam. Fruits drupaceous, broadly elliptic to slightly obovate, bluntly mammillate and briefly apiculate with persistent style base, 20-25 mm long, 14-18 mm — brown color when dried; mesocarp fibrous-fleshy, without conspicuous taste or odor in fresh, un- ripe fruits; endocarp bony-fibrous; pedicels of 920 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74 4 a— ee EE ee eS i Ñ ha i aj uararibea funebris (Llave) Mos T m | ssp. nicaraguensis Alver ç y [ HOLOTYPE! - | ^ | Det.: W.S. ace peach 3 1986 | L L Her ba rium, Un jf Wisconsin | (WIS) lw, g. PME; e [fo N Chicago Natural History Museum Esc uela Agricola Panamericana Quararibea funebris (Llave) Vischer det. AmolinaR. 1967 Mente - fores or cloud forest, Finca Sta. Marta de Ostuma, Cor- HERBARIUM alt. 1400 m January 18, 1965. To MADISON (WIS) a e. wa "- Melina R., Terus P. Derothy N. Git — V — and Chester Laskowski, FIGURE |. Holotype of Quararibea funebris subsp. nicaraguensis. 1987] fruits slender, 10-19 mm long, bearing 2-3 per- sistent bracteoles near middle; ca/yx persistent, accrescent, cupulate, enclosing fruits for ^-^ their length, longitudinally finely rugose when dry, wingless, green, with vestiture as in flowering condition, the summit erose or irregularly toothed. Seeds 2, or 1 by abortion, ca. 13 mm long, 8 mm diam., consisting mostly of pale cot- yledonal tissue; testa thin, dark brown. Seedlings unknown when young, probably hypogeal and cryptocotylar, when older bearing leaves smaller and more than those of mature plants. Paratypes. NICARAGUA. ESTELÍ: El Zacatón, “El De- lirio” camino a la laguna de Miraflor, 13?13'N, 86°14' W 1,400 m, 11 Jan. 1984 (fr), Moreno 22676 (MO, WIS). JINOTEGA: entre Santa Lastenia y entrada a Aranjuez, 13°02'N, 85?55'W, 1,200-1,250 m, 19 Jan. 1984 (fr), Sandino 4704 (MO); Finca Aventina, in sierra east of Jinotega, 1,400-1,500 m, 23 June 1947 (st), Standley 10002 (F); region of Las Mercedes, sierra east of Jino- tega, 1,200-1,500 m, 3 July 1947 (st), Standley 10746 (F); vicinity of Finca San Roque, sierra east of Jinotega, 300-1,500 m, 5 July 1947 (fr), Standley 10856 (F, US). MADRIz: Cerro Volcán Somoto (Volcán Tepeso- moto), 13?26'N, 86?35'W, 25 Sep. 1980 (fl), Moreno 2892 (MO). MATAGALPA: vicinity Santa María de Os- tuma, on flanks of Cerro El Picacho, immediately east of Hwy. 3, ca. 9 km north (by air) of Matagalpa, 1 3300'N 85°55'W, ca. 1,500 m, : ies 1982 (fl buds, fr). Alver- son & Moreno 1982 , CR, F, MEXU, MO, NY, US, WIS); obice 1984 (MO, NY, WIS); (st), 420 m, 5 Jan. 1984 (fr), Gentry, Stevens O aL rarer carretera a Jinotega, 8 1,400-1,450 m, 7 Oct. 1980 (fr), MEE 3381 (MO, WIS); 13°00'N, 85°55'W, 1,400-1,480 m, 1 Dec. 1982 (fr), 18951 (MO) (same locality as . Alverson & Moreno : 982); Cordillera cards Santa M m N of Matagalpa, 1, m, I Ness (MO); 26 Nov. 1977 (fr), 3005 (BM, ed MO); 1,500 m, 15 Jan. 1 (fr), b nti A Ub ndn 23952 (F, G, us en Hda. 95°53" m 1,200-1,300 m, 22574 (MO). B Dec. 1983 (fr), Stevens Quararibea funebris subsp. nicaraguensis is known only from the highlands of northwestern Nicaragua in the departments of Estelí, Jinotega, Matagalpa, and Madriz. It is common to loca ally between 1,200 and 1,600 m. Flowering occurs during August and September, and fruiting takes place October through January, indicating a phe- nology similar to that of the typical lowland sub- species. Although very similar, this new subspecies is NOTES 921 BLE l. Comparison of the subspecies of Quara- ribea funebris in Nicaragua. Material is from Nicaragua with the exception of flowers of subsp. funebris, which are from northwestern Costa Rica Subspecies nicara- Character guensis funebris Leaf length 71-221 145-289 Leaf width 27-104 62-125 Petiole length 7-18 15-27 Floral pedicel length 11-15 13-14 Floral calyx length 9-11 18-20 Petal length 15-16 33-40 Staminal column length 16-22 33-34 Style length 17-26 35-36 Fruiting pedicel length 10-19 9-17 Fruiting edi length 11-13 18-26 Fruit leng 20-25 25-30 Measurements in mm. morphologically distinct from Quararibea fu- nebris subsp. funebris, being smaller in nearly all respects (Table 1). Its fruiting pedicels, however, tend to be longer than those of subsp. funebris. The two subspecies can be easily distinguished by their floral measurements. When in fruiting condition, the calyces provide the most obvious key character; they cover half or less of the ma- ture fruit of subsp. nicaraguensis, as opposed to more than half in subsp. funebris. Thanks are due to D. A. Kolterman for help with the Latin description, to C. Lipke for the photograph, to A. Pool for providing specimen data, and to H. H. Iltis, A. H. Gentry, and W D. Stevens for comments on the manuscript. I gratefully acknowledge the O. N. and E. K. Allen Fund : i , Wisconsin for financial support of field research in 1982 LITERATURE CITED ALVERSON, W. S A emiri aes Aubl. s.l. (Bom- bacacea - America and the . 12 in Novorum Vegetabilium Descriptiones, Volume 2. Mex STANDLEY, P. C. 1923. Pp. 787—788 in Trees and 922 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74 Shrubs of Mexico, Part 3. Contr. U.S. Natl. Herb. — William S. Alverson, Herbarium, Botany De- 23(3): 517-848. artment, University of Wisconsin, Madison VISCHER, W. 1920. Surles Quararibea Aubl., un genre a ds 53706 ey ie Discs de Bombacées à ovaire infére. Bull. Soc. Bot. Ge- COMAN ae néve 11: 199-210. ANNALS OF THE MISSOURI BOTANICAL GARDEN VOLUME 74 1987 The Annais, published quarterly, contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden, St. Louis. Papers origi- nating outside the Garden will also be accepted. Authors should write the Editor for information concerning arrangements for publishing in the ANNALS. EDITORIAL COMMITTEE GEoncE K. Rocers, Editor Missouri Botanical Garden MARsHALL R. CROSBY Missouri Botanical Garden GERRIT DAVIDSE Missouri Botanical Garden Jonn D. DWYER Missouri Botanical Garden & St. Louis University PETER GOLDBLATT Missouri Botanical Garden HENK VAN DER WERFF Missouri Botanical Garden Colophon This volume of the ANNALS of the Missouri Botanical Garden has been set in APS Times Roman. The text is set in 9 point type while the figure legends and literature cited sections are set in 8 point type. The volume has been printed on 70# Centura Gloss, an acid-free paper designed to have a shelf-life of over 100 years. Centura Gloss is manufactured by the Consolidated Paper Company. Photographs used in the ANNALS are reproduced using 300 line screen halftones. The binding used in the production of the Annars is a proprietary method known as Permanent Binding. The ANNALS is printed and distributed by Allen Press, Inc. of Lawrence, Kansas 66044, U.S.A. (O Missouri Botanical Garden 1988 ISSN 0026-6493 VOLUME 74 ALVERSON, WILLIAM S. A New Subspecies of Quararibea funebris (Bom- bacaceae) from Nicaragua AUERBACH, MICHAEL & Avi SHMIDA. Patch Formation among Israeli Cru- cifers: How Do They Get Away with It? AVERETT, JOHN E., PETER H. RAVEN & ELSA ZARDINI. Flavonoid Systematics of Seven Sections of Ludwigia (Onagraceae) BARNETT, Lisa C. An Unusual New Species of Helmiopsis H. Perrier (Ster- culiaceae) from Madagascar BENTLEY, BARBARA L. Nitrogen Fixation by Epiphylls in a Tropical Rain- forest Benzinc, Davip H. Vascular Epiphytism: Taxonomic Participation and Adaptive Diversity BERNHARDT, PETER. A Comparison of the Diversity, Density, and Foraging Behavior of Bees and Wasps on Australian Acacia Brown, KEITH S., JR. Chemistry at the Solanaceae/Ithomiinae Interface Brown, KEITH S., JR. (See Boyce A. Drummond III & Keith S. Brown, J r. BRYSON, CHARLES T. (See Robert Kral, James Manhart & Charles T. Bry- son) BURMAN, ALASDAIR G. (See Gerrit Davidse & Alasdair G. Burman) ............... CARLQUIST, SHERWIN. Wood Anatomy of Noteworthy Species of Ludwigia (Onagraceae) with Relation to Ecology and Systematics CARR, GERALD D. (See Elisabeth Rabakonandrianina & Gerald D. Carr) CaTLING, PauL M. Notes on the Breeding Systems of Sacoila lanceolata (Aublet) Garay (Orchidaceae) CLARK, LYNN G. Two New Mesoamerican Species of Chusquea (Poaceae: Bambusoideae) CLARK, LYNN G. New Combinations in Chusquea (Poaceae: Babusoideae) CRABTREE, Davip R. Angiosperms of the Northern Rocky Mountains: Albian to Campanian (Cretaceous) Megafossil Floras Croat, THoMAS B. & MICHAEL H. GRAYUM. New Combinations in Central American Araceae D'Arcy, WILLIAM C. & JOHNNIE L. GENTRY, JR. Witheringia folliculoides (Solanaceae): A New Species from Costa Rica D'Arcy, WiLLIAM G. & Davip N. SMITH. Saracha spinosa—A New Com- bination in Peruvian Solanaceae DAVIDSE, GERRIT. Four New Species of Axonopus (Poaceae: Paniceae) from Tropical America 1987 DAVIDSE, GERRIT. (See Robert Kral & Gerrit Davidse) DAVIDSE, GERRIT. (See Ahsan A. Vahidy, Gerrit Davidse & Youji Shige- nobu) DAVIDSE, Gerrit & ALASDAIR G. BURMAN. A New Species of Thrasya (Poaceae: Panicoideae) from the Mosquitia of Nicaragua and Hon- duras DavipsE, GERRIT & R. P. Erlis. drundoclaytonia, a New Genus of the teyermarkochloeae (Poaceae: Arundinoideae) from Brazil 0... DE Nevers, GREGORY C. The Genus Attalea (Palmae) in Panama ........... Denton, M. F. (See E. B. Leopold & M. F. Denton) DIETRICH, WERNER & WARREN L. WAGNER. New Taxa of Oenothera L. Sect. Oenothera (Onagraceae) DIETRICH, WERNER & WARREN L. WAGNER. A New Combination and New Subspecies in Oenothera elata Kunth (Onagraceae) DiLcHER, Davip. Memorial to Herman F. Becker (1907-1985) oo. DopsoN, C. H. (See Alwyn H. Gentry & C. H. Dodson) DRUMMOND, Boyce A., II] & KrrrH S. Brown, JR. Ithomiinae (Lepidoptera: Nymphalidae): Summary of Known Larval Food Plants Dwyer, JoHN D. Book Review Erus, R. P. (See Gerrit Davidse & R. P. Ellis) FADEN, RoBERT B. & D. R. Hunt. Reunion of Phaeosphaerion and Com- melinopsis with Commelina (Commelinaceae) GENTRY, ALWYN. Book Review GENTRY, ALWYN H. & C. H. Dopson. Diversity and Biogeography of Neo- tropical Vascular Epiphytes GENTRY, A. H. & J. STEYERMARK. A Revision of Dilodendron (Sapindaceae) GENTRY, JOHNNIE L., JR. (See William G. D'Arcy & Johnnie L. Gentry) GOLDBLATT, PETER. Chromosome Cytology of Oldenburgia (Asteraceae— Mutisieae) GOLDBLATT, PETER. Systematics of the Southern African Genus Hexaglottis (Iridaceae—Iridoideae GOLDBLATT, PETER. Notes on the Variation and Taxonomy of Watsonia borbonica (W. pyramidata, W. ardernei) (Iridaceae) in the South- western Cape, South Africa GOLDBLATT, PETER. (See James E. Henrich & Peter Goldblatt) u. GOLDBLATT, PETER. (See James F. Henrich & Peter Goldblatt) ..... GOLDBLATT, PETER. (See James E. Henrich & Peter Goldblatt) 20000000... GOLDBLATT, PETER. (See Porter P. Lowry II, Peter Goldblatt & Hiroshi Tobe) GOLDBLATT, PETER & JAMES E. HENRICH. Notes on Cipura (Iridaceae) in South and Central America, and a New Species from Venezuela ............. 121 GoLDBLATT, PETER & Jon WILLIAMS. Notes on Chromosome Cytology of Rutaceae—Diosmeae GRAHAM, ALAN. Fossil Pollen of Sabicea (Rubiaceae) from the Lower Mio- cene Culebra Formation of Panama GRAYUM, MicHAEL H. (See Thomas B. Croat & Michael H. Grayum) ....... HaBECK, JAMES R. Present-day Vegetation in the Northern Rocky Moun- HAMMEL, Barry E. The Origami of Botany: A Guide to Collecting and Mounting Specimens of Cyclanthaceae HENRICH, JAMES E. (See Peter Goldblatt & James E. Henrich) U... Henrico, JAMES E. & PETER GorpBLATT. A Review of the New World Species of Orthrosanthus Sweet (Iridaceae) HENRICH, JAMES E. & PETER GorpBLATT. Mesoamerican Sisyrinchium (Iri- daceae) New Species and Records, and Notes on Typification ............. HENRICH, JAMES E. & PETER GorpBLATT. The Mesoamerican Neomarica (Iridaceae), N. variegata Henrich & Goldblatt, Comb. Nov. .. u... HERRERA, JAVIER. Flower and Fruit Biology in Southern Spanish Mediter- ranean Shrublands Hoim-NIELSEN, L. B. & J. E. LawEssoN. New Species of Passiflora Sub- genus Passiflora from Ecuador Hunt, D. R. (See Robert B. Faden & D. R. Hunt) Jupziewicz, EMMET J. (See Thomas R. Soderstrom & Emmet J. Judziewicz) KRAL, ROBERT. A new *Viorna" Clematis from Northern Alabama .......... KRAL, Robert & Gerrit Davipse. A New Species of Bulbostylis (Cyper- aceae) from Tropical America .. KRAL, ROBERT & ARMANDO URQUIOLA. Two Cuban Novelties in Xyris ...... KRAL, ROBERT, JAMES MANHART & CHARLES T. Bryson. A New Carex Sect. Oligocarpae (Cyperaceae) from Western Arkansas and Eastern Okla- homa ...... KRAL, ROBERT, L. B. SMITH & MARIA DAS GRAÇAS DE Lapa WANDERLEY. Two New Xyris (Xyridaceae) from the Amazon Basin of Brazil ecco Kuyt, Joa. Novelties in Mesoamerican Mistletoes (Loranthaceae and Vis- caceae) . Lawesson, J. E. (See L. B. Holm-Nielsen & J. E. Lawesson) ..... Leopo.p, E. B. & M. F. Denton. Comparative Age of Grassland and Steppe East and West of the Northern Rocky Mountains LIESNER, RONALD. (See Julian A. Steyermark & Ronald Liesner) U. LITTLETON, JAN. (See C. J. Webb & Jan Littleton) LonENCE, Davip H. The Fruits of Decarydendron (Monimiaceae) .............. Lowry, Porter P., II, PETER GorpBLATT & HiRosui Tose. Notes on the Floral Biology, Cytology, and Embryology of Campynemanthe (Liliales: Campynemataceae) " MAcLEISH, NANDA F. F. Revision of Eremanthus (Compositae: Vernonieae) MANHART, JAMES. (See Robert Kral, James Manhart & Charles T. Bryson) MEEROW, ALAN W. Biosystematics of Tetraploid Eucharis (Amaryllidaceae) MiLLER, CHARLES N., JR. Contributions to a Symposium on the Evolution of the Modern Flora of the Northern Rocky Mountains: Introductory Remarks MiLLER, CHARLES N., JR. Land Plants of the Northern Rocky Mountains before the Appearance of Flowering Plants MILLER, JAMES S. Two New Species of Cordia (Boraginaceae) from Central merica Mort, Scorr A. Eschweilera costaricensis (Lecythidaceae): A New Species for the Floras of Costa Rica and Nicaragua Mort, Scorr A. & GHILLEAN T. PRANCE. A Guide to Collecting Lecythidaceae NEILL, Davip A. Trapliners in the Trees: Hummingbird Pollination of Fr- ythrina Sect. Erythrina (Leguminosae: Papilionoideae) ORNDUFF, RoBERT. Reproductive Systems and Chromosome Races of Oxalis pes-caprae L. and Their Bearing on the Genesis of a Noxious Weed OYEWOLE, S. O. Cytotaxonomic Studies in the Genus Urginea Stein in West Africa. II. Karyotype Evolution in Urginea altisima (L.) Baker ............. OYEWOLE, S. O. Cytotaxonomic Studies in the Genus Urginea Stein in West Africa. III. The Case of Urginea indica (Roxb.) Kunth in Nigeria ...... OYEWOLE, S. O. Cytotaxonomic Studies in the Genus Urginea Stein in West Africa. IV. Population Differentiation and Karyotype Variation in Ur- ginea indica (Roxb.) Kunth PorPENDIECK, HaNs-HELMUT. Monoecy and Sex Changes in Freycinetia (Pandaceae) PRANCE, GHILLEAN T. (See Scott A. Mori & Ghillean T. Prance) l. RABAKONANDRIANINA, ELISABETH & GERALD D. CARR. Chromosome Numbers of Madagascar Plants RAVEN, PETER H. (See Hiroshi Tobe & Peter H. Raven) RAVEN, PETER H. (See John E. Averett, Peter H. Raven & Elsa Zardini). SCHUBERT, B. G. (See O. Téllez V. & B. G. Schubert) SHIGENOBU, YouJI. (See Ahsan A. Vahidy, Gerrit Davidse & Youji Shige- nobu) SHMIDA, AVI. (See Michael Auerbach & Avi Shmida) SMITH, Davip N. (See William C. D'Arcy & David N. Smith)... SMITH, L. B. (See Robert Kral, L. B. Smith & Maria das Gracas de Lapa Wanderley) .. SODERSTROM, THOMAS R. & Emmet J. Jupziewicz. Systematics of the Amphi- Atlantic Bambusoid Genus Streptogyna (Poaceae 871 STEIN, BRUCE A. Siphocampylus oscitans (Campanulaceae: Lobelioideae), a New Name for Burmeistera weberbaueri from Peru STEIN, BRUCE A. Synopsis of the Genus Burmeistera (Campanulaceae: Lo- belioideae) in Peru STEYERMARK, JULIAN A. Flora of the Venezuelan Guayana—ll 2... STEYERMARK, JULIAN A. New Taxa of Rubiaceae from Venezuela ......... STEYERMARK, JULIAN A. Flora of the Venezuelan Guayana— III U. STEYERMARK, JULIAN A. Yutajea, Another New Genus of Rubiaceae from the Guayana Highland STEYERMARK, J. (See A. H. Gentry & J. Steyermark) STEYERMARK, JULIAN A. & RONALD LIESNER. Notes on Cornus (Cornaceae) in South America SYTSMA, KENNETH J. The Shrubby Gentian Genus Macrocarpaea in Panama TAYLOR, CHARLOTTE M. Reconsideration of the Generic Placement of Pal- icourea domingensis (Rubiaceae: Psychotrieae) TEBBS, MARGARET. A New Species of Piper (Piperaceae) from Central Amer- ica TÉLLEZ V., O. & B. G. SCHUBERT. Una Nueva Especie del Género Dioscorea (Dioscoreaceae) del Estado de Queretaro, México TETENYI, PETER. A Chemotaxonomic Classification of the Solanaceae ....... Tope, Hinosur. (See Porter P. Lowry II, Peter Goldblatt & Hiroshi Tobe) Tose, HiRosHi & PETER H. RAVEN. Systematic Embryology of the Aniso- phylleaceae Topzia, CanoL A. A New Variety of Hedyosmum (Chloranthaceae) from Nicaragua URQUIOLA, ARMANDO. (See Robert Kral & Armando Urquiola) U. VaAHIDY, AHSAN A., GERRIT DAVIDSE & YoujJI SHIGENOBU. Chromosome Counts of Missouri Asteraceae and Poaceae WAGNER, WARREN L. (See Werner Dietrich & Warren L. Wagner) WAGNER, WARREN L. (See Werner Dietrich & Warren L. Wagner) _... Wake, Davip B. Adaptive Radiation of Salamanders in Middle American Cloud Forests WANDERLEY, MARIA DAS GRACAS DE LaPa. (See Robert Kral, L. B. Smith & Maria das Gragas de Lapa Wanderley) Wess, C. J. & JAN LITTLETON. Flower Longevity and Protandry in Two Species of Gentiana (Gentianaceae) WEBSTER, GRADY L. A New Species of Jatropha (Euphorbiaceae) from Nicaragua WENDT, Tom & HENK VAN DER WERFF. A New Species of Ocotea (Lau- raceae) from Southeastern Mexico WERFF, HENK VAN DER. A Revision of Mezilaurus (Lauraceae) „u WERFF, HENK VAN DER. Six New Species of Neotropical Lauraceae WERFF, HENK VAN DER. (See Tom Wendt & Henk van der Werff) WILLIAMS, Ion. (See Peter Goldblatt & Ion Williams) Winc, Scorr L. Eocene and Oligocene Floras and Vegetation of the Rocky Mountains Wotre, Jack A. Contributions to a Symposium on the Evolution of the Modern Flora of the Northern Rocky Mountains: Dedication Wo re, Jack A. Memorial to Harry D. MacGinitie (1896-1987) WOLFE, Jack A. An Overview of the Origins of the Modern Vegetation and Flora of the Northern Rocky Mountains ZARATE PEDROCHE, SERGIO. Clarification of the Taxonomy of Leucaena salvadorensis Standley ex Britton & Rose ZARDINI, Elsa. A New Combination in Lucilia (Compositae—Inuleae) ......... ZARDINI, Ersa. (See John E. Averett, Peter H. Raven & Elsa Zardini) ZULOAGA, FERNANDO O. A Revision of Panicum Subgenus Panicum Section Rudgeana (Poaceae: Paniceae) 413 Volume 74, No. 3, pp. 463-680 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN, was published on December 22, 1987. Instructions for Authors, Annals of the Missouri Botanical Garden The Annals publishes oes ewes in systematic botany and related fields. Tex n English or Spanish. When in Spanish, an English Pedes in addition to a Spanish abstract is required. Short papers, especially Authors are urged to contact the editor prior to submitting manuscripts. 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OAXACA: Sierra o Nolesco, Talea, 3 Feb. 1987 (fl), Jergensen - 865 (BM, G, K, US). Dates and 22325 status are tional. - ^ page width (514 inches or ca. 140 mm). Figures are numbered i numerals in | the order of their citation | letters. CONTENTS Contributions to a Symposium on the Evolution of the Modern Flora of the Northern Rocky Mountains: Introductory Remarks Charles N. Miller, Jr. Dedication Jack A. Wolfe Memorial to Harry D. MacGinitie (1896-1987) Jack A. Wolfe Memorial to Herman F. Becker (1907-1985) David Dilcher Land Plants of the Northern Rocky Mountains before the Appearance of Flowering Plants Charles N. Miller, Jr. Angiosperms of the Northern Rocky Mountains: Albian to Campanian (Cretaceous) Megafossil Floras David R. Crabtree Eocene and Oligocene Floras and Vegetation of the Rocky Mountains Scott L. Wing An Overview of the Origins of the Modern Vegetation and Flora of the Northern Rocky Mountains Jack A. Wolfe Present-day Vegetation in the Northern Rocky Mountains James R. Habeck .... Comparative Age of Grassland and Steppe East and West of the Northern Rocky | Mountains E. B. Leopold & M. F. Denton AEN Fossil Pollen of Sabicea (Rubiaceae) from the Lower Miocene Culebra Formation of ; Panama Alan Graham ee ‘Systematics of the Amphi-Atlantic Bambusoid Genus Streptogyna quiin Thomas R. Soderstrom & Emmet J. Judziewiez Wood Anatomy of Noteworthy Species of Ludwigia (Onagraceae) with Relation to Ecalogy and Systematics . Sheruin Carlquist The Origami of Botany: a Guide to Collecting and Mounting Specimens of Cyclanthaceae Barry E. Hamme | . Mesoamerican Sisyrinchium | (Iridaceae): New Species suit Records, ind Notes on "t Typification Ms pue E. Henrich & Peter Caen. 681 683 684 689 692 707 William š.