VOLUME 72 1985 ANNALS OF THE MISSOURI BOTANICAL GARDEN The ANNALS, published quarterly, contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the Garden will also be ac- cepted. Authors should write the Editor for information concerning arrangements for publishing in the ANNALS. EDITORIAL COMMITTEE Nancy Morin, Editor Missouri Botanical Garden CHERYL R. BAUER, Editorial Assistant Missouri Botanical Garden MARSHALL R. CROSBY Missouri Botanical Garden T DAVIDSE Missouri Botanical Garden Joun D. DWYER Missouri Botanical Garden & St. Louis University PETER GOLDBLATT 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 ANNALS 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. © Missouri Botanical Garden 1985 ISSN 0026-6493 BARRETT, SPENCER C. H. & Jorr S. SHORE. Dimorphic Incompatibility in Turnera hermannioides Camb. (Turneraceae) .. BERNARDELLO, Luis M. (See Eduardo A. Moscone & Luis M. Bernar- dello) BERRY, PAUL E. The Systematics of the Apetalous Fuchsias of South America, Fuchsia Sect. Hemsleyella (Onagraceae) BROOKS, DANIEL R. Historical Ecology: A New Approach to Studying the Evolution of Ecological Associations CARLQUIST, SHERWIN. (See Peter Goldblatt, Hiroshi Tobe, Sherwin Carlquist & Varsha C. Patel) CASTILLO-CAMPOS, GONZALO & DAvID H. LonENCE. Antirhea aromatica (Rubiaceae, Guettardeae), a New Species from Veracruz, Mexico ....... CLARK, LYNN G. Three New Species of Chusquea (Gramineae: Bambu- soideae) COUGHENOUR, MICHAEL B. Graminoid Responses to Grazing by Large Herbivores: Adaptations, Exaptations, and Interacting Processes ......... CRACRAFT, JOEL. (See Vicki A. Funk & Joel Cracraft) CRACRAFT, JOEL. Biological Diversification and Its Causes CRANE, PETER R. Phylogenetic Analysis of Seed Plants and the Origin of Angiosperms CROAT, THOMAS B. Collecting and Preparing Specimens of Araceae .... D’Arcy, W. G. & BARRY HAMMEL. The Plants of ‘Ocoquili’ Island, San Blas Coast, Panama FERGUSON, I. K. (See A. J. Hemsley & I. K. Ferguson) FEUER, SYLVIA M. & Jos Kuyt. Fine Structure of Mistletoe Pollen VI. Small-Flowered Neotropical Loranthaceae Funk, V. A. Phylogenetic Patterns and Hybridization FUNK, VICKI A. & JOEL CRACRAFT. The Implications of Phylogenetic Analysis for Comparative Biology: The Thirtieth Annual Systematics Symposium GOLDBLATT, PETER, HIROSHI TOBE, SHERWIN CARLQUIST & VARSHA C. PATEL. Familial Position of the Cape Genus Empleuridium ........ GOLDBLATT, PETER. Systematics of the Southern African Genus Geisso- rhiza (Iridaceae-Ixioideae GOLDBLATT, PETER. Book Review GRAHAM, ALAN, R. H. STEWART & J. L. STEWART. Studies in Neotropical Paleobotany. III. The Tertiary Communities of Panama—Geology of the Pollen-bearing Deposits GRAHAM, ALAN. Studies in Neotropical Paleobotany. IV. The Eocene Communities of Panama HAMILTON, CLEMENT W. Notes on and Descriptions of Seven New Species of Mesoamerican Clethraceae 504 539 HAMMEL, BARRY. (See W. G. D'Arcy & Barry Hammel) HEMsLEY, A. J. & I. K. FERGUSON. Pollen Morphology of the Genus Erythrina (Leguminosae: Papilionoideae) in Relation to Floral Struc- ture and Pollinators Hiru, KHIDIR W. & THOMAS R. SODERSTROM. Biological Basis for Ad- aptation in Grasses: An Introduction Hurt, MICHAEL J. A New Syngonanthus (Eriocaulaceae) from Southern exico JUDZIEWICZ, EMMET J. Pharus parvifolius subsp. elongatus (Poaceae), a New Subspecies from Tropical America KING, ROBERT M. (See Harold Robinson, A. Michael Powell, Robert M. King & James F. Weedin) KNAPP, SANDRA. New Species of Solanum Section Geminata (G. Don) Walp. (Solanaceae) from South and Central America Kuur, Jos. (See Sylvia M. Feuer & Job Kuijt) LORENCE, DAvID H. A Monograph of the Monimiaceae (Laurales) in the Malagasy Region (Southwest Indian Ocean) LORENCE, DAvID H. (See Gonzalo Castillo-Campos & David H. Lorence) MAYDEN, RICHARD L. (See E. O. Wiley & Richard L. Mayden) -na MoRLEY, THOMAS. Five New Taxa of New World Memecyleae (Mela- stomataceae) MOSCONE, EDUARDO A. & Luis M. BERNARDELLO. Chromosome Studies on Hydromystria laevigata (Hydrocharitaceae) PATEL, VARSHA C. (See Peter Goldblatt, Hiroshi Tobe, Sherwin Carlquist | & Varsha C. Patel) Pour, RICHARD W. Three New Species of Rhipidocladum from Mesoam- erica POWELL, A. MICHAEL. (See Harold Robinson, A. Michael Powell, Robert M. King & James F. Weedin) RAVEN, PETER H. (See Hiroshi Tobe & Peter H. Raven) REDMANN, R. E. Adaptation of Grasses to Water Stress— Leaf Rolling and Stomate Distribution ROBINSON, HAROLD, A. MICHAEL POWELL, ROBERT M. KING & JAMES F. WEEDIN. Chromosome Numbers in Compositae, XV: Liabeae RosEN, DoNN E. Geological Hierarchies and Biogeographic Congruence in the Caribbean |... SCHATZ, GEORGE E. A New Cymbopetalum (Annonaceae) from Costa Rica and Panama with Observations on Natural Hybridization ... SHMIDA, A. Why do Some Compositae Have an Inconsistently Deciduous Pappus? . SHORE, JoEL S. (See Spencer C. H. Barrett & Joel S. Shore) 264 570 451 SODERSTROM, THOMAS R. (See Khidir W. Hilu & Thomas R. Soder- SIME DU mali E M n 823 SOEJARTO, D. D. Saurauia molinae, a New Species of Actinidiaceae from Centa bc d e E Mu D O O 878 STEBBINS, G. LEDYARD. Polyploidy, Hybridization, and the Invasion of New Habitats 824 STEWART, J. L. (See Alan Graham, R. H. Stewart & J. L. stewart) |... 485 STEWART, R. H. (See Alan Graham, R. H. Stewart & J. L. Stewart) ...... 485 THOMASSON, JOSEPH R. Miocene Fossil Grasses: Possible Adaptation in Reproductive Bracts (Lemma and Palea) 843 Tose, HIROSHI. (See Peter Goldblatt, Hiroshi Tobe, Sherwin Carlquist & Varsha C. Patel) 167 TOBE, HIROSHI & PETER H. RAVEN. The Histogenesis and Evolution of Integuments in Onagraceae 451 TopzIA, CAROL A. Campylocentrum tenellum (Orchidaceae), a New Species from Panama 876 WEEDIN, JAMES F. (See Harold Robinson, A. Michael Powell, Robert M. King & James F. Weedin) 469 WHETSTONE, R. DAvip. Burnelia reclinata var. austrofloridensis (Sapo- taceae), a New Variety from South Florida, U.S.A. 544 WILEY, E. O. & RICHARD L. MAYDEN. Species and Speciation in Phylo- genetic Systematics, with Examples from the North American Fish Fauna 596 Volume 72, No. 3, pp. 451—590 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN, was published on 30 August 1985. ANNALS f THE ISSOURI BOTANICAL GARDEN .UME 72 1985 NUMBER 1 CONTENTS A Monograph of the Monimiaceae (Laurales) in the Malagasy Region (South- west Indian Ocean) David H. Lorente c : l VOLUME 72 SPRING 1985 NUMBER 1 ANNALS OF THE | MISSOURI BOTANICAL GARDEN x The ANNALS, published quarterly, contains papers, primarily in | systematic botany, contributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the Garden will also be ac- | cepted. Authors should write the Editor for information concerning 1 arrangements for publishing in the ANNALS. Instructions to Authors | are printed on the inside back cover of this issue. EDITORIAL COMMITTEE NANCY Morin, Editor Missouri Botanical Garden CHERYL R. BAUER, Editorial Assistant Missouri Botanical Garden MARSHALL R. CROSBY Missouri Botanical Garden GERRIT DAVIDSE Missouri Botanical Garden JOHN D. DwvER Missouri Botanical Garden & St. Louis University PETER GOLDBLATT Missouri Botanical Garden For subscription information contact the Business Office of the Annals, P.O. Box 299, St. Louis, MO 63166. Subscription price is $65 per volume U.S., $70 Canada, and Mexico $75 all other countries. Personal subscriptions are available at $30 and $35, respectively. irmail delivery charge, $30 per volume. Four issues per volume. The ANNALS OF THE MISSOURI BOTANICAL GARDEN (ISSN 0026-6493) is pub- lished quarterly by the Missouri Botanical Garden, 2345 Tower Grove Ave., St. Louis, MO ANNALS OF THE MISSOURI BOTANICAL GARDEN VOLUME 72 1985 NUMBER 1 A MONOGRAPH OF THE MONIMIACEAE (LAURALES) IN THE MALAGASY REGION (SOUTHWEST INDIAN OCEAN)!23 DAvID H. LORENCE* ABSTRACT A systematic, ecological, and evolutionary study of the Monimiaceae of the p in the Malagasy floristic region, southwest Indian Ocean (Madagascar and the Mascarene and Com eripe is presented. The Malagasy Monimiaceae are here taken to comprise four genera ad 5 endemic: Monimia (3 spp., "asap Decarydendron (3 spp., Madagascar); Ephippiandra (6 (6 ii. - scar); Tambourissa (43 spp., scar, Mascarenes, Comores). rere a large, ancient continental island, is the regional um ed diversity with three genera a cies. The smaller, younger volcanic islands harbor fewer taxa: Mauritius (2/11); Réunion E E (1/2); Grand Comore u 1); MayoHe ur 1); Mohéli (1/ D; Rodrigues (0). Synopses of! the family’ S taxonomic history, fthe geology, ge eomorphology, and climate of each ee given. Distribution of species according to habitat is discussed for each island and ecological observations are presented. Vegetative and floral morphology and anatomy are described from both the literature and original observations. A synopsis of the palynology is given. Cytological investigations of ten species of Tambourissa revealed all to have n 19. Results were inconclusive for A with n = ca. 44—48. Field studies on floral biology of Tambourissa revealed that variations in floral m het sa color, and odor — for several different pollination syndromes involving Coleoptera and Diptera. Breeding experiment s demonstrated a strong tendency towards outcrossin a self-incompatibi ility in most monoecious species. Population density was fae to pollinator Sabio in the three species studied. ! Based on a Ph.D. dissertation submitted to the Graduate School of Arts and Sciences of Washington University, St. Louis, Missou 2] sincerely thank Alwyn H. Gentry for his continued support, advice, and criticism during the preparation of this monograph. I also thank Peter H. Raven, P. Goldblatt, W. D’Arcy, R. Sussman, B. Schaal, J. Dwyer, O J. Sexton, P. J. Watson, M. Crosby, P. K. Endress, W. R. Philipson, and N. Morin for carefully reviewing and iting various stag, the manuscript or assisting me in various ways. I would like to thank the following individuals whose collaboration facilitated this study: R. E. Vaughan, J. Williams, R. Julien, J. Guého, and G. ier, Mau uga esearch Institute, Mauritius; W ly, ner, and D'Argent, Forestry Dept., Mauritius; J ty, Ministry of Agriculture, Mauritius; T. - entre Univer- sitaire, La Réunion; A. Rakotozafy, Parque Botanique, Tsimbazaza, Madagascar; P. Roulleau, Dept. of Agri- culture, Mayotte; M. J. E. Coode, Royal Botanic Gardens, Kew; J. Bosser, Museum National d'Histoire ae Paris; P. K. Endress, Botanischer Garten und Institut fiir Systematische Botanik der Universitat SC Vinay and V. Zenger, Washington University; N. C. Pant, Commo nwealth Institute of mp London Mrs. Wanling Peng for her excellent illustrations. I also than ck ith specimens: BM, G, K, MAD, MAU, MO, P, REU, TL-R, Z. Finally, I am particularly grateful Ag and ; wife, Ginette, for her assistance, companionship, and endurin support durin: ie s >À 3 This work was supported by an NSF E Dissertation Improvement Grant (DEB 78-19954), a Sigma Xi Grant-in- Pu of Research, and a Washington University Graduate Fellowship, which are all gratefully acknowle * LE de Biología, x Postal 70-233, Universidad Nacional Autónoma de México, C.U., Deleg. Coyoacán, 04510, México, ANN. Missouni Bor. GARD. 72: 1-165. 1985. 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 n the taxonomic treatment, keys to genera and species are eurn Synonymy, descriptions, dis- tribution, habitat, discussions, vernacular names, econom for each species. Hedycaryopsis Dang. is united bis Ephip following new name and combinations: E. dom ses, and herbarium disp ba Maii a Lorence (basionym: H. capuronii ns are given madagascariensis (Cav.) Lorence; E. perrieri ae ees Ephippiandra capuronii Cav. is trans- ferred to Tambourissa as T. beanjadensis Lorence. Finally, Phanerogonocar, Tambourissa Sonn. and the following new names are proposed: T. alaticarpa Lorence (basionym: P. perrieri Cav.); T. longicarpa Lorence (basionym: P. capuronii Cav.). This monograph represents a systematic, eco- logical, and evolutionary study of the Old World tal island of Madagascar and its satellite islands, the Mascarene and Comore archipelagos (Figs. 1, 2). On many of these islands Monimiaceae constitute an important woody element of the wet and cloud forest formations. Monimiaceae were chosen for study because they are a large and diverse but poorly known family belonging to the so-called “woody Ra- nales" that have played a key role in interpre- tation of the morphological and evolutionary patterns of primitive angiosperms. Extensive adaptive radiation into four genera and some 55 species, all endemic, in the islands of this region suggests that the Monimiaceae hold a unique po- sition as a model for island evolution by a relict group of plants. Study of the family is all the more imperative in view of the threat posed by man's destructive activities in the region coupled with the relentless invasive pet from in- troduced plant species. In spite of their importance as floristic com- ponents on many of these islands, the Monimi- aceae have never been the subject of a detailed field study. Not only has there been a dearth of knowledge regarding their ecology and biology, but lack of field studies has also resulted in con- siderable taxonomic confusion because ofthe dif- ficulty in matching androecious and gynoecious flowering material and of understanding mor- phologically variable species. During the course of this work I was able to spend an entire year in the Malagasy region llecting M in their na- tive habitats in the following islands: Mauritius (where the majority of the work was carried out), species in the field and have accumulated a great deal of new land biological that is incorporated into this monograph. TAXONOMIC HISTORY OF THE MONIMIACEAE The Monimiaceae sensu lato are an old and diverse assemblage of about 30 genera and nearly sphere. The family has undergone a great deal of taxonomic flux because of the striking diversity of its members, which display various combi- nations of primitive and advanced characters. morphol- in terms of reduction, both in size and number of parts; in rearrangement of their orientation from spiral to radial and finally to decussate; in closure of the female floral receptacle; and also in a transition from free stalked carpels to an inferior syncarpous gynoecium (Corner, 1976). floral bisexuality to monoecy and finally dioecy. e Monimiaceae have a long and interesting taxonomic history dating back nearly two cen- turies. De Jussieu first founded the family in 1809 (as Monimieae) based on three previously described genera: Monimia Thouars, Atherosperma La- ie and Siparuna Aubl. (as Citrosma Ruiz & av.). Tulasne subsequently monographed the dea in 1855 (18 55b) including seven addi- amily in a much broader sense to include the present day Calycanthaceae and Gomortegaceae, defining five series or tribes: Calycantheae, Hor- tonieae, Tambourisseae, Atherospermeae, and Gomortegeae. Bentham and Hooker (1883) rec- ognized only two tribes (Monimieae and Ath- erospermeae) based on features of the anthers and ovules, adding more genera but excluding the rome ay This system was also fol- lowed by Pax (1891), who raised both tribes to the ee level (Monimioideae and Ath- 1985] ( Grande Comore Mohéli ` 4 Anjouan V Mayotte COMORES oa 6 200 km 46E MADAGASCAR Regional map of study area, showing uius and the Comore archipelago erospermoideae) and created six tribes for the species groups of Bentham and Hooker. n their monograph of the family for “Das Planzenreich " Perkins and Gilg (1901) followed Pax, an ntham and Hooker in recognizing two hcec cim but modified the tribes, creating an additional one under Monimioideae and merging two under Atherospermoideae. Perkins (1911) later updated the earlier revision using the same system. Hutchinson (1964) more or less followed Bentham and Hooker in dividing the Monimiaceae into two subfamilies, but recog- nized only four tribes. As a result of its extreme diversity, many at- tempts have been made to dissolve the Moni- miaceae sensu lato (Brown, 1814; Lindley, 1853; Pichon, 1948; Schodde, 1970; Smith, 1969). The LORENCE— MONIMIACEAE 3 56°E Rodrigues 2 ES f) Mauritius Cy Réunion MASCARENES 200 km IGURE2. Regional map of study area, showing the Mascarene archipelago. truly extraneous elements, however, notably Amborella, Austrobaileya, and Trimeniaceae, have been removed from the family by workers such as Pichon (1948) and particularly Money et al. (1950) in their important study of the mor- phology and relationships of the family. I agree with Thorne (1974) that the Monimiaceae sensu lato comprise a rather large and variable but co- herent group with at least five major evolution- ary lines derived from now extinct common ancestors. Recent morphological and anatomical studies by Sampson (1969a, 1969b), Endress (1980a, 1980b), as well as those by Money et al. (1950), support the rationality of treating these lines as subfamilies belonging to a single, diverse family rather than pursuing the alternative of splitting them into five separate families. Hortonioideae, the smallest subfamily, con- bers pos- sess an elaborate perianth with spirally arranged petaloid and sepaloid series, floral bisexuality, free stalked carpels borne on a discoid receptacle, basal staminal appendages, helical banded pol- len, primitive wood anatomy, and the gametic chromosome number n = 19 (Money et al., 1950; Goldblatt, 1974; Endress, 1980a). Because it has retained the greatest assemblage of primitive representative of the ancestral condition (Money et al., 1950), although recent studies by Endress (1980a) suggest it occupies a position interme- diate between the Atherospermoideae and the Monimiaceae sensu stricto (i.e., Mollinedioideae plus Monimioideae of Thorne Members of the next subfamily, Atherosper- moideae, are linked to the Hortonioideae by the similarly petaloid perianth and floral bisexuality vancements over the whorled arrangement of the parts and reduction 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN in their number, and a tendency towards floral nidis The Atherospermoideae are char- acterized by having bisporangiate valvate sta- mens, erect, basal ovules (usually), a single noda trace, advanced wood anatomy, and a basic chro- mosome number of x = 22. These plants also possess 1—2-colpate pollen and show a trend to- wards specialized comose, wind-dispersed car- s. The five genera and 12 species of Ather- ospermoideae have a strongly disjunct austral distribution ranging from Australasia to Chile, emphasizing the subfamily's great antiquity. The Siparunoideae are also characterized by bisporangiate valvate stamens, erect basal ovules, a single nodal trace and a basic chromosome number of x — 22. These features are similar to those found in the Atherospermoideae, and the two groups probably represent lines of parallel evolution from a common ancestral stock (e.g., Schodde, att): bd ean ae panoe lack the b ate wood fibers, and hippocrepiform A j the Ath- erospermoideae. Furthermore, advanced trends such as vessels with simple perforation plates, and closed, reduced floral receptacles with a ve- lum replacing the perianth occur in the Siparu- noideae in addition to other differences such as inaperturate, granulate pollen and bird-dispersed carpels. The subfamily comprises three genera: Siparuna with about 160 species in tropical America, Glossocalyx with three species in west tropical Africa, and Bracteanthus with a single species in Brazil. Thorne (1974) suggested that Bracteanthus may merit tribal or subfamilial sta- tus, and that Glossocalyxoideae be recognized as a subfamily (Thorne, 1983). The largest subfamily, Mollinedioideae, con- tains about 18 genera and over 145 species (Thorne, 1983). Its members are characterized by having apetalous, unisexual flowers display- ing various degrees of reduction and specializa- tion, relatively primitive wood anatomy, and ga- ic chromosome numbers of n = 18, 19, 22, ca. 38, ca. 43, and 57 for various genera (Gold- blatt, 1974). Thorne (1974) united Monimia (3 species, Mascarene Islands) with Peumus (monotypic, Chile) as the sole members of the subfamily Monimioideae. Both genera have paired basal appendages on the stamens; large, spinose pollen, and comparatively advanced wood anatomy. This seems to be the most rational placement of these two genera in light of the available evi- dence [Vor. 72 Although recent subfamilial revisions of the Monimiaceae have been adequate (e.g., Thorne, 1974), all have neglected treatment at the tribal level. As Perkins' (1911, 1925), Perkins and Gilg's (1901), and Hutchinson's (1964) treatments of he subfamilies and tribes are outdated, a critical reassessment of the subfamilies of Monimiaceae at the tribal level is urgently needed. s GEOLOGICAL HISTORY AND REGIONAL SETTING Beginning with a Jurassic Gondwana super- continent, the formation of the Indian Ocean commenced as Africa (with India and Madagas- car) rotated away from South America at the start of the Cretaceous (summarized by Fisher et al., 1967). Atthe same time the Antarctica-Australia mass began drifting southward, whereas the block comprising Africa-India-Madagascar remained intact until the mid-Cretaceous prior to about 90 Ma (Raven, 1979). At that time Madagascar occupied a position against the present Kenya- Somalia coast and was part of a Malagasy-Mas- carene subcontinent joined to India via the now largely submerged Mascarene Plateau (Axelrod & Raven, 1978). Separation of this subcontinent from Africa occurred during the mid- to late Cre- taceous at about 90 Ma (Raven, 1979). Probable separation of India from the Malagasy-Masca- northward tinued until it collided with the Asian landmass at about 45 Ma (Raven, 1979). Formation of the present crescent-shaped Mascarene Plateau began with the separation of a portion of continental crust from Madagascar in the late Cretaceous or early Tertiary as Mad- agascar moved southward (Fisher et al., 1967; Kutina, 1975; Wild, 1975). This fragment, com- sed of Precambrian granites, included the present Seychelles archipelago and possibly the entire western flank of the Mascarene Plateau above the Saya de Malha bank (Fisher et al., 1967). Formation of the remaining portion of the Mascarene Plateau from the Saya de Malha southward to Mauritius was probably accom- plished by late Cretaceous-early Tertiary sub- marine volcanic extrusions (Fisher et al., "d ° Island emerges, is parallel to a fracture zone ex- tending eastward across the Midoceanic ridge complex and was probably formed during the late Tertiary (Fisher et al., 1967). It is a volcanic 1985] entity distinct from the Mascarene Plateau, sep- arating the latter from Mauritius and Réunion (Fig. 2). The Comore archipelago, situated between Madagascar and Africa (Fig. 1), is a product of successive Pliocene-Pleistocene volcanic erup- tions (Esson et al., ). In contrast to Mada- gascar, which is a continental island, the Mas- carenes and Comores are volcanic in origin and therefore strictly oceanic in nature. GEOMORPHOLOGY AND CLIMATE At present Madagascar is separated from the African mainland by the Mozambique channel, some 500 km wide at its narrowest point (Fig. 1). Situated midway between Africa and the northern part of Madagascar and separated on either side by depths of ca. 3,000 m are the four islands comprising the Comore archipelago: Mayotte, Anjouan, Mohéli, and Grande Co- more. Stretching from ca. 800 to 1,600 km due east of Madagascar lies the liceo archipel- ago comprising the islands of Réunion, Mauri- tius, and Rodrigues. The MI are sepa- rated from the nearest landmass, Madagascar, by a stretch of ocean ca. 4,000-5,000 m deep. MADAGASCAR As the world's third largest island with an area of 590,000 km?, Madagascar is truly continental in nature, consisting of a fragment of the original Gondwana landmass. Its large size, insular na- ture, and long isolation have made it an impor- tant refugium for many unique and relictual groups of plants and animals, including the le- murs. Its former connection to Africa and India once provided a route for Cretaceous dinosaurs and the rich angiosperm floras that appeared at that time (Raven & Axelrod, 1978 Geologically, Madagascar is composed of four elements (Brenon, 1972). A much folded and outcropping metamorphic Precambrian base- ment in the form of a peneplain comprises the rugged central plateau which rises above 1,000 m (range 800-1,500 m) and also the highest mountains which are under 3,000 m (2,876 m at the Tsaratanana massif ). This basement drops pd abruptly along the great eastern escarpment wn to a narrow coastal plain. Secondly, a ihi tilted Permo- poses sh continental sedimentary formation belonging to the Karroo system overlies the Precambrian basement and forms a band along the western third of the is- LORENCE—MONIMIACEAE 5 land. Thirdly, basaltic volcanic formations be- longing to upper Cretaceous eruptions form ex- tensive flows in parts of the eastern and western regions and have given rise to certain mountain complexes in the north, center, and south of i pe iue weathering and laterization of the Precambrian basement has resulted in a mantle of ae clays averaging 10-15 m thick overlying the bedrock. This is undergoing mas- sive erosion accelerated by historic removal of much of the island’s vegetation by man. Madagascar’s diverse sana a logy is the re- sult of submersion an ifting, pedogenesis and erosion, and scattered vulcanism during its long history. This edaphic diversity coupled with a variety of climatic types has resulted in the wide spectrum of habitats that exists in Madagascar. Madagascar’s position, considerable size (590,000 km^?), and close juxtaposition of phys- ical features such as lowland coastal plains grad- ually or steeply sloping to areas of medium al- titude, and mountains exceeding 2,500 m elevation, are all responsible for its considerable environment which is most prevalent in the east- em region. Here, wenn creme ; currents mod- erate the t winds and tropical cyclones egit upon the east coast deposit heavy rains along the coast and eastern escarpment, permitting the devel- opment of dense, evergreen wet forest which forms a zone along the eastern third of Mada- gascar. Second, Madagascar is sufficiently long (ca. 1,500 km) and wide (ca. 450 km average) to dis- play many characteristics of a small continent. This continentality IS Tettected in a wide spec- trum of j° lity, pre- E dA and wind regimes. Asa result. of its gitudinal extension, the island possesses sub- Bistum regions in the north (at 12?S) where a monsoon climate and dry season modify the composition of the evergreen forest. In the south, it passes through the Tropic of Capricorn and extends into the subtropics at its southern ex- tremity (25?30'S), giving rise to considerable thermal variation. On a latitudinal basis, both precipitation and temperature decrease consid- erably in the central uplands. Precipitation fur- ther decreases and temperature increases to the west and south as the Trade winds rapidly lose their remaining moisture and velocity due to the rain shadow effect. As a result, the vegetation 5 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE 1. relation to area, elevation, and age of island. [Vor. 72 Distribution of total Monimiaceae and Tambourissa species per island in the Malagasy region in Area Elevation Age Total Species Total Species Island (km?) (m) (Ma) Monimiaceae Tambourissa Madagascar 590,000 3,000 35 26 (as 1sland) Réunion 2,500 3,000 3.0 5 2 Mauritius 1,865 900 7.8 11 10 Grande Comore 1,148 2,400 0.01f 1 1 Anjouan 424 1,600 1.5 2 2 Mayotte 370 700 3.7 1 1 Mohéli 290 800 ca. 1.5 1 1 Rodrigues 110 400 0 0 becomes progressively more xeric in nature to sion, andalsoas! ] In addition the south and west, passing from sclerophyllous evergreen forest to semideciduous forest and fi- nally thorn scrub and desert (Koechelin et al., 1974, give a detailed account of the vegetation). Temperature, wind, and rainfall are further in- fluenced by the island's varied relief, giving rise to a spectrum of bioclimates varying in both tem- perature and precipitation. For example, some of the higher mountains extend into the cloud zone and harbor a montane cloud forest for- mation, above which occurs a low, ericoid scrub characteristic of the high montane zone. his diversity of habitats and extreme ecolog- ical gradients in conjunction with Madagascar's long isolation have resulted in high levels of flo- ristic endemism: up to 4896 of the genera and 9596 of the species are endemic in certain regions (Koechelin et al., 1974; Leroy, 1978). It is there- fore not surprising that Madagascar harbors the greatest constellation of Monimiaceous taxa in the region (three genera and ca. 35 species), Af- rica notwithstanding (Table 1). MASCARENE ARCHIPELAGO The three Mascarenes are volcanic oceanic is- lands that consist ofthe exposed summits of sub- marine mountain chains, i.e., Mauritius and Ro- drigues, and an isolated volcano arising from an abyssal submarine plain, i.e., Réunion (Cadet, All three islands experience a tropical insular climate moderated by the maritime effects of the warm ocean and the steady south-east Trade wind regime. Precipitation is most abundant during the warm and rainy austral summer season last- ing from December to April, occurring as oro- graphic rains resulting from adiabatic air expan- to bringing rain, cyclones, with winds gusting up to 240—320 km/hr, are often destructive to the vegetation. On the average, direct hits by cy- clones occur only once every 12 to 15 years. The relatively drier and cooler austral winter season lasts from May to November. The mean annual precipitation ranges from 1,100 to 1,700 mm in Rodrigues (Lorence, 1976), and from 1,000 to 5,000 mm in Mauritius and Réunion (Lorence, 1978; Cadet, 1977). Meanan- nual temperatures in the Mascarenes are not ex- treme, ranging from 16 to 20?C in the uplands and from 21 to 25°C in the lowlands during the winter, and from 21 to 25°C in the uplands and 27 to 30*C in the lowlands during the summer. Réunion, the largest (2,500 km?) and eastern- most of the Mascarene Islands, is the summit of a volcanic cone rising some 3,000 m above sea level. Its active volcano, Piton de la Fournaise, still erupts frequently, emitting basaltic lavas similar to the Pleistocene flows in Mauritius (Fisher et al., 1967). Quaternary lavas from the Piton des Neiges flows have been dated at 2.1 Ma, although Fisher et al. (1967) suggest that submerged caldera flows may be older, possibly ca. 3 Ma (Cadet, 1977). Some subsidence appears to have taken place during the Quaternary, per- haps resulting in a small reduction of the island's surface area. Coral reef development around Ré- union is virtually absent. In addition to its accentuated relief, Réunion's 3,000 m altitude provides the greatest range of habitats of any of the Mascarenes, ranging from semideciduous dry forest and thicket in the west- ern rain shadow, to lowland wet forest, montane cloud forest and high montane ericoid scrub and grassland. Tambourissa, with only two species here, is less abundant and much less diverse than 1985] in Mauritius (10 species; Table 1). It is signifi- cant, however, that Monimia reaches its greatest diversity and abundance here, with three species often forming locally abundant or dominant ele- ments of the wet and cloud forest formations. Much ofthe upland vegetation in Réunion is still intact, but little remains in the lowlands. Mauritius, the central island, is second largest with an area of 1,8 m? and a maximum al- titude just under 900 m. By far the oldest of the three Mascarenes, its earliest lavas are Pliocene, dating to 7.8 Ma (McDougall & Chamalaun, 1969), and its vulcanism ended about 100,000 years ago during the Pleistocene. During its long history, erosion has moderated the island's relief which consists of a central plateau fringed by a discontinuous mountain range dipping down to coastal plains in all but the southern sector. Ex- cept for its well-developed fringing coral reefs, Mauritius is completely volcanic in origin and composed of alkali olivine basalts deposited in three main strata. The presence of coral deposits at altitudes of up to 100 m above sea level on the mainland suggests past fluctuations in the sea level, although progressive subsidence seems to be the case (Fisher et al., 1967). If the submarine plateau surrounding Mauritius is taken into con- sideration, the island's previous area was prob- ably ca. 2,500 km? during the lowered sea levels of the Quaternary (comparable to Réunion's present area), as opposed to its 1,865 km? today. It seems that earlier in the island's history its relief was more accentuated than today but prob- ably did not exceed 1,000 m in altitude during the last 3.5 Ma (Cadet, 1977). Mauritius supports a fairly diverse array of plant communities, notably semideciduous dry forest in the western rain shadow area, lowland and lower montane evergreen wet forest, small areas of cloud forest on the highest mountains, and patches of heath formation (Lorence, 1978). Next to Madagascar, Mauritius harbors the most species of Tambourissa (10). In contrast, Monimia, which is essentially a genus of cloud Representative areas of most of the island's plant communities are still preserved by way of nature reserves and crown land, but many of these have been irreversibly degraded by invasive exotic plant species. Rodrigues is the smallest and westernmost of the Mascarenes, with an area of only 110 km? and an altitude just under 400 m. A ridge com- LORENCE— MONIMIACEAE 7 posed of a uniform series of alkali olivine basalt flows whose exposed parts do not exceed 1.54 1965), it has apparently mation and probably once occupied nearly ten times its present area (Cadet, 1977). Now only the summit of a large submarine platform re- mains exposed, supporting large, well-developed fringing coral reefs. In view of its high specific level of floristic endemism, equalling the other two Mascarenes, Rodrigues could be older than Réunion but younger than Mauritius (Cadet, 1977), contrary to the suggestion put forth by McDougall et al. (1965). Because of its small size and low elevation, Rodrigues is ecologically the least diverse of the Mascarenes. Most ofthe island was once covered by a low evergreen moist or wet forest, drier on its leeward western side, although sections of uplifted coralline calcarenite support an inter- esting scrub lormalson. Although a species of Monimia no substantiated records of Monimiaceae are known from the island. This is probably because of its isolation and paucity of the proper habitats. Also, nearly all of Rodrigues’ indigenous vege- tation has been destroyed by man. collected in Ro rigues, COMORE ARCHIPELAGO The Comores are a group of four volcanic is- lands located in the Mozambique channel mid- way between Madagascar and Mozambique on the east African coast. Situated on a submarine ridge, lee e are the product of successive volcanic eruption g from southeast to northwest and are no older than the Pliocene-Pleistocene (Guilcher et al., 1965; Hajash & Armstrong, 1972). Analogous to the Mascarenes, the Co- mores differ in being more closely spaced (sep- arated by distances no greater than 65 km; Fig. 1), are closer to major landmasses, and are on the average smaller and younger (Esson et al., — o Mayotte is the most southeasterly, the oldest (ca. 3.65 Ma), and most highly eroded of the four islands. With an area of 370 km? and a maximum altitude of 660 m, it consists of a succession of old lava flows from at least two volcanic erup- tions (basaltic and phonolitic) which gave rise to the bulk of the island. Following periods of rel- ative stability, a final period of vulcanism poured lava across the eastern part of the island. Other indications of Mayotte's relatively advanced age 8 ANNALS OF THE MISSOURI BOTANICAL GARDEN within the archipelago are a highly embayed coastline, development of the finest barrier reef and lagoon in the Indian Ocean, and occurrence of deep, red laterized soils (Guilcher et al., 1965). Both are relatively old structures with eroded but still recognizable craters and cones. Anjouan is larger, attaining 1,595 m, and has an area of 424 km? with an accentuated relief and narrow but fringing coral reefs emcompassing about two- thirds of the papi Its oldest lavas have been dated at 1.52 Ma (Hajash & Armstrong,1972). Mohéli is the nd or the Comores, rising to 790 m altitude with an area of only 290 km". Its relief is less rugged but its reefs are also wide and well developed, although its lavas have not been dated. Mayotte, Anjouan, and Mohéli are all vol- canically extinct. Grande Comore, largest (1,148 km?) and youngest of the four islands, has an active vol- cano whose summit attains 2,360 m, and is es- sentially a mass of scarcely eroded lava lacking permanent rivers and streams. The only dated lavas from Grande Comore are recent (0.01 Ma), although underlying flows are thought to be older (Hajash & Armstrong, 1972). Coral reef devel- opment around the island is sparse and embry- onic, also reflecting its relative youth. The Comores lie in the south-east Trade wind belt, although they are somewhat sheltered by Madagascar, and cyclones are rare. The drier and cooler winter season extends from May to Oc- tober, the majority of the rain falling during the hot austral summer season from November to April (Manicacci, 1939). In 1935, the following annual rainfalls were recorded: Grande Comore, 4,254 mm; Anjouan, 1,411 mm; Mayotte, 869 mm (at Dzaoudzi, but probably double inland); Mohéli, 478 mm (Manicacci, 1939). Tempera- tures range from 25 to 35?C in the summer and from 18 to 25°C in the winter, depending on the elevation. The Comores generally support evergreen moist or wet forest on their lower windward and l , and dry for- est in the low altitude regions of the rain shadow, Anjouan and Grande Comore have zones of cloud forest above 900 or 1,000 m, and this gives way to Philippia heath formation on the Karthala vol- cano massif on Grande Comore. Each island has but a single species of Tambourissa, except for Anjouan which has two (Table 1). Four of the five Comorean species of Tambourissa appear to [Vor. 72 be closely related and may be derived from a common ancestor. BIOGEOGRAPHY AND DISTRIBUTION Biogeographically, it appears that direct mi- gration of the Monimiaceae between Africa and South America occurred by the Eocene, after which widespread extinction took place in Africa (Raven & Axelrod, 1974; Axelrod & Raven, 1978). Fossils that have been considered Mo- nimiaceous are known from the late Cretaceous (ca. 70—80 Ma) in Europe, Africa, and Argentina. In contrast to the widespread floral and faunal impoverishment that took place in continental tropical Africa (Richards, 1973, 1979), the Mal- agasy region provided an important refuge for many elements of the area's biota; for example, the lemurs in Madagascar and the Comores, and the now extinct flightless birds such as the dodo and solitaire in the Mascarenes (Carlquist, 1965). The Monimiaceae may be thought of as bo- tanical counterparts of the lemurs and are rep- resented by four genera and about 55 species in the Malagasy region, all endemic. The isolation and habitat diversity offered by islands are well known to be conducive to rapid adaptive radia- tion (e.g., Carlquist, 1965, 1974), which is ex- emplified by the Malagasy area Monimiaceae. In marked contrast, only two genera of Monimiace- ae have persisted in continental Africa—i.e., Glossocalyx and Xymalos— comprising a total of four species. This is primarily because the Mo- nimiaceae attain their greatest diversity in pre- montane wet and montane cloud forest forma- tions, habitats that are abundant in the Malagasy region but poorly represented in Africa. Fur- thermore, most cannot tolerate the ecologically significant dry periods that are characteristic of even the wet forest in Africa (Richards, 1973), although their absence from the cloud forests of the east African mountains is puzzling. Perhaps these forests were greatly reduced in extent dur- ing the dry periods of the Pleistocene, and Mo- nimiaceae were unable to recolonize them from their more equable refuge areas on Madagascar and the Mascarenes. Madagascar harbors three closely allied genera of trees and shrubs belonging to the subfamily Mollinedioideae. Of these, Ephippiandra (6 spp.) and Decarydendron (3 spp.), are endemic. Their inability to spread to the other islands may be a reflection of their poor dispersibility or seed vi- ability. Alternatively, suitable habitats may be 1985] lacking on the satellite islands. This does not appear to be the case, however, because relatively large and elevated islands such as Réunion and Mauritius offer ed of the same habitats as Madagascar (Table Tambourissa, oe has reached six of the seven Satellite islands where it has become es- tablished and undergone successful adaptive ra- diation (Table 1). Reports that Tambourissa oc- curs in Java (e.g., Cavaco, 1965) appear to be erroneous (see discussion under 7. ficus), and the genus appears to be restricted to the Malagasy floristic region. Possible reasons for its success most likely include its bright red-orange drupes, which are ostensibly more attractive to birds (those of Ephippiandra ripen black) resulting in greater dispersibility, a greater seed viability (drupes of Tambourissa may take two or three months to germinate; D. Lorence, pers. observ.), and enhanced ability to compete and evolve rap- idly. In any case, Tambourissa provides an ex- cellent model for island evolution in a family of primitive angiosperms. onimia (Monimioideae) consists of three closely related species endemic to the Mascarene Islands of Mauritius and Réunion. Its absence from Madagascar is puzzling in view of its bird- dispersed carpels. Peumus in Chile appears to be its closest relative, although it also shares a num- ber of features with Palmeria from Oceania (see discussion under Monimia). HABITAT AND ECOLOGY The majority of ecological and floristic studies or surveys conducted for the islands of the Mal- agasy region demonstrate or at least mention the importance of the Monimiaceae as woody ele- ments of the vegetation (Vaughan & Wiehe, 1937, 1941; Humbert, 1965; Legris, 1969; Koechlin, 1972; Koechlin et al., 1974; Cadet, 1977). In addition, data from specimens provide much in- formation. MADAGASCAR Although I am unaware of any specific eco- logical studies for Madagascar involving Mo- nimiaceae, the broad vegetational surveys of Humbert (1965) and Koechlin et al. (1974) cou- pled with data from specimens which I have ex- amined and those cited by Cavaco (1959) have enabled the habitat and distribution of most species to be established. A significant number of taxa, however, are known from only one or LORENCE— MONIMIACEAE 9 several collections, possibly a result of under- collecting, which may therefore inaccurately por- tray their actual distributions. Of the Imperfectly Known Tambourissa species, only numbers 40 (T. sp. A) and 42 (T. sp. C), are included in this discussion, because collection localities for the others are either too vague or were not located. Work done by Humbert (1965) and Koechlin et al. (1974) indicates that in Madagascar species of Tambourissa, along with Weinmannia (Cu noniaceae), are among the most frequent ba characteristic components of the canopy in the montane wet/cloud forest zone. Tambourissa at- tains its greatest diversity here, with at least 14 of the island's 26 species occurring in this zone (Table 2) which forms a belt along the eastern part of the island from ca. 800 m and extends upward to the mountains of the central plateau to between 1,300 and 1,800 m. Annual precip- itation here ranges from ca. 1,000 to 2,000 mm, with a short dry season. In addition to Tambourissa, two endemic Madagascan genera also occur in this zone, De- carydendron (2 spp.) and Ephippiandra (3 spp.), bringing the total number of Monimiaceae to 19 (Table 2). The Tambourissa/Weinmannia zone is clearly Madagascar's richest habitat in terms of Monimiaceae species and genera. In the east, this ae zone is subtended by: a lowland wet forest o udi receiving an annual waspa sb. Bul ca. 1, —3,500 mm and having no significant dis season. In addition to sharing about three species with the former zone, including Ephip- piandra madagascariensis and two species of Tambourissa, this zone possesses a number o its own species, most notably Decarydendron la- mii and five species of Tambourissa: T. alati- carpa, T. castri-delphinii, T. decaryana, T. lon- gicarpa, and T. sp. C. Besides T. castri-delphinii, the wide-ranging 7. purpurea and T. religiosa are the only Monimiaceae known to inhabit the coastal forests on sand dunes The lowland wet forest also extends to the small Sambirano region in the northwest, including Nosy Be Island, where it is further characterized by the presence of many Chlaenaceae, an endem- ic family. In addition to harboring such wide- sis and Tambourissa purpurea, the Sambirano area has two endemics: 7. nitida and T. nosy- ensis. At altitudes from ca. 1,800 to 2,000 m in the cloud zone on the mountains of the central re- 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE 2. Distribution of Monimiaceae according to baia in Madagascar. . Lowland wet forest zone of Myristicaceae and An- thostema (ca. 0—800 m), including littoral forest and Sambirano s Decarydendron Ephippiandra Mad ys, Tambourissa as rpa T. castri- un decaryan T. P T. nitida T. nosybensis T. perrieri MN Medium altitude wet/cloud forest zone of Tam bourissa and Weinmannia [ca. 800—1,300(-1,800) m]. Decarydendron helenae perrieri Enkinpiandra domatiata E. madagascariensis E. microphylla d beanjadensis capuronii d flricostata T. perrie . Sclerophyllous montane lichen forest zone (central mountains, 1,300-2,000 m). t myrtoidea E. perrie. E. tsaratanensis 5 gracilis T. parvifolia High montane Philippia idque heath formation (central mountains, above ). pisci ne aj E. perri RU USA parvifolia . Sclerophyllous nont oo zone id siasa bojeri and Chl pes of central domain ; up to ca. 800-900 m). Tambourissa hildebrandtii T. purpurea Uo i wr [VoL. 72 TABLE 2. Continued. 6. Semideciduous dry forest zone of western domain (below ca. 700-800 m) Tambourissa bathiei (probably riverine) T. perrieri (probably riverine) gion, the 7ambourissa/ Weinmannia belt is re- placed by a dense sclerophyllous forest with an abundance of epiphytic lichens. Perhaps because of the lower temperatures, the Monimiaceae drop out rapidly here (Table 2). Ephippiandra micro- phylla and Tambourissa gracilis are virtually the only species shared with the zone below. The only species characteristic of this zone and, to a lesser extent, of the high montane ericoid thicket superseding it beyond the cloud zone at above ca. 2,000 m are Ephippiandra myrtoidea, E. per- rieri, E. tsaratanensis, and Tambourissa parvi- folia. The western slopes of the central plateau at and below ca. 800-900 m are characterized by a hotter, drier climate with more extreme tem- peratures. This area supports sclerophyllous for- est of Uapaca bojeri (Euphorbiaceae) and Chlaenaceae, with only Tambourissa hilde- brandtii and T. purpurea, two of the most wide- spread species, able to persist here. In the dry western regions that extend to the coast, the sclerophyllous forest is gradually re- placed by a semideciduous dry forest of Dalber- gia, Commiphora, and Hildegardia. The only Monimiaceae found here are Tambourissa bath- iei and T. perrieri, which apparently occur in riverine forests. It should be emphasized that the vegetation of Madagascar, like most of the other islands, has undergone and continues to undergo severe deg- radation and destruction under the influence of man (Rauh, 1979). Because of its size, however, remnants of most and even fairly ext areas of some vegetation types still remain (Rakoto- zafy, pers. comm.), but the need for their con- servation is essential. MASCARENE ISLANDS On the average larger and older than the Co- mores, the Mascarene Islands also harbor more genera (two) and species (15) of Monimiaceae despite their greater isolation. In addition, the Mascarenes have been comparatively well stud- 1985] LORENCE-— MONIMIACEAE T ied floristically (Baker, 1877; Balfour, 1879; Cor- demoy, 1895; “Flore des Mascareignes," in pro- gress) and — (Vaughan & Wiehe, 1937, 1939, 1941; Wiehe, 1949; Rivals, 1952; Cadet, 1977), thus Meise a more definitive assess- ment of the family's status there. RÉUNION Réunion is analogous to Grande Comore in being the only volcanically active member of its archipelago but is larger and older, supporting two genera and five species of Monimiaceae in contrast to the latter's unique species. Cadet's (1977) comprehensive study of the island's vege- tation emphasizes the importance of the family as woody components ofthe flora. Although har- boring the greatest expanse of indigenous forest of the three Mascarenes, many areas of Réunion are undergoing a dramatic invasion by vigorous exotic species such as Psidium, Rubus, and Fur- craea, similar to that occurring in the other two islands. Much of Réunion's lowland forest has disap- peared, particularly in the dry western sector. Fairly extensive areas of lowland wet forest (‘‘fo- rét mégatherme hygrophile" sensu Cadet, 1977) still remain above ca. however, on the southern slopes of the active volcano, Piton de la Fournaise (at Mare Longue, Brülé de Baril and other areas). Here a lush wet forest dominated y Sapotaceae (Labourdonnaisia, Mimusops, and Sideroxylon) has become established on the mo- saic of lava flows where the rainfall averages 4,000-5,000 mm per year. Tambourissa elliptica subsp. micrantha appears here at ca. 300 m asa local, occasional to common component of the middle and upper strata and ranges to the mid or rarely upper limits of the wet forest zone (ca. 800-1,000 m) where it is generally replaced by T. elliptica subsp. elliptica. The latter is char- acteristically a smaller understory tree or treelet of occasional to common occurrence in both the wet forest zone below 1,100 m and the cloud forest zones (ca. 1,100-2,000 m) (Fig. 31). he only other species to occur in the lowland wet forest zone of Réunion is Monimia ovali- folia, which appears at ca. 500 m and extends to 1,700 m in the cloud forest zone. It typically forms local populations on sheltered slopes and in valleys and may become a canopy tree. The lower limits of the cloud belt lie at ca. 800-1,100 m and mark the transition from wet to cloud forest (“forêt mésotherme hygrophile" of Cadet), a zone characterized by lower tem- peratures and relatively constant high humidity. Itisin the cloud forests of Réunion that Monimia attains its greatest abundance and diversity. The climax is considered to be the Monimia/Dom- beya association (“Bois de couleurs") although Acacia heterophylla Willd. is dominant in some areas (Cadet, 1977). Ofthe three Monimia species occurring in this zone, M. rotundifolia is by far the most abundant and widespread, codominant in many areas with species of Dombeya (Sterculiaceae) and other genera, e.g., at Col de Bellevue and Bébour forest (Fig. 22). Mature individuals 8-16 m high with dense, spreading crowns are often major com- ponents of the mid stratum and canopy. It occurs less frequently in the Acacia/Nastus forest and occasionally in Philippia heath up to ca. 2,000 m, with its lower limit extending down to ca. 1,000-1,100 m at the interface with lower alti- tude wet forest. Monimia amplexicaulis is more restricted in distribution than M. rotundifolia and is generally confined to the mid and upper limits of the cloud forest, occasionally with Acacia. It characteris- tically forms discrete populations on the shel- tered, leeward Scarpin | of the “cirques” or ancient d is known primarily from the following localities where it usually re- places M. rotundifolia: Hauts du Tévelave, Plaine des Cafres, and Cirque de Cilaos (Fig. 22). An example of ecological differentiation between the two species was studied at Hauts du Tévelave, where Monimia amplexicaulis begins to replace M. rotundifolia, which occurs alone below ca. 1,700 m. Here, both species occur sympatrically at ca. 1,700 m along the Domanial line (Ligne Dominicale); neither intermediates nor hybrids were observed among dozens of individuals ex- amined. The two species are easily recognized in the field and appear to maintain their integrity by flowering at different times. Monimia ovalifolia is an occasional tree up to ca. 1,700 m altitude but is rarely abundant. It occurs sympatrically with M. rotundifolia in many localities, e.g., at Bébour forest where both species remain distinct and have different flowering times (Fig. 22). Only two species of Tambourissa occur in the cloud zone: 7. crassa and the macrophyllous up- land form of T. elliptica subsp. elliptica. Tam- bourissa crassa is a light-demanding small tree which occupies the upper stratum of low forest or occurs in relatively open mixed heath at 1,200- 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE 3. nities. [VoL. 72 Occurrence of Monimiaceae species according to habitat in Mauritius: indigenous plant commu- Sideroxylon Cloud Semide- Philippia Wet Forest: High Forest: Forest: ciduous Heute Pieter el Brise Mt. Dry Forest: Pétrin; Both; Ombre; Macabé; er; Cocotte; Yemen; 660 m, 550 m, 300 m 660 m 600 m, 76 200 m, pptn. pptn. pptn. pptn. pptn. pptn. pptn. 4,000 mm 2,400 mm 2,400 mm 3,200 mm 2,400 mm 5,000 mm 1,600 mm Monimia ovalifolia X Tambourissa amplifolia X X T. cocottensis X T. cordifolia X T. ficus X X X X T. pedicellata X X T. peltata X X X X X T. quadrifida X T. sieberi X X T. tau X X X X T. tetragona X 2,000 m. Tambourissa elliptica subsp. elliptica is a slender, sciaphilous understory forest treelet (1,100-2,000 m). Both species are occasional woody elements of the cloud forest (Fig. 31). On the tops of exposed crests and at altitudes above ca. 1,700-2,000 m the cloud forest gives way to a low ericoid heath formation dominated by Philippia, as in Madagascar and Grande Co- more. Apart from the occasional occurrence of Monimia amplexicaulis, M. rotundifolia, Tam- bourissa crassa, and T. elliptica subsp. elliptica, whose upper ranges end here, no species are spe- cially adapted to the heath formation. Likewise, no species are known to be adapted to the semi- deciduous dry forest (“forêt mégatherme" of Ca- det) in the island's dry, leeward western sector (annual precipitation 800-1,500 mm n essence, Réunion's extensive cloud forests have favored adaptive radiation in the genus Monimia, which comprises three of its five Mo- nimiaceae species. Monimia rotundifolia is the most widespread and abundant species, consti- tuting a woody element of major importance in the cloud forest zone. Tambourissa, with two species, has not undergone the extent of adaptive radiation here that has occurred in Mauritius and Madagascar, although seemingly suitable habi- tats exist. This may be attributable to its more recent arrival, or to the island" Syounger age. That in — Ti ambour Bionic IS ident by t the presence oftwo closely related species, one with two subspecies occu- pying different ecological zones. MAURITIUS Geologically the oldest of the Mascarene Is- lands, Mauritius has probably undergone con- siderable reduction by erosion since its incep- tion, being slightly smaller and only a third as high as Réunion. Since the island's colonization by man almost 400 years ago, its original vege- tation has been removed from nearly every ac- cessible surface and has also undergone extreme degradation by introduced biota. Tracts repre- senting most of the island's original vegetion still remain, however, in the form of nature reserves and crown land which comprise ca. 5-6% of its superficies (Lorence, 1978). Unfortunately, vir- tually all have been invaded by introduced plants such as Furcraea, Ligustrum, Psidium, Rubus, an many others, not to mention introduced animals (monkeys, stag, pigs) and birds (sparrows, my- nahs, bulbuls), which help spread seeds of the 1975; Jolly, 1982; Pasquier & Jones, 1982). The majority of forested areas remaining in the uplands (precipitation 2,000-5,000 mm) are occupied by floristically rich wet forest of vari- 1985] able composition but usually characterized by the presence of Sideroxylon cinereum Lam., puberulum A.DC., and other Sapotaceae such as Labourdonnaisia (the “Sideroxylon thicket” of Vaughan & Wiehe). Remnants of this formation extend down to ca. 200 m in some areas (e.g., at Bel Ombre) where up to three species of Tam- bourissa occur: T. amplifolia, T. peltata, and T. tau (Table 3; Figs. 38, 40) In addition to the above, species such as Tam- bourissa ficus and T. pedicellata appear at higher altitudes, e.g., at Pieter Both Mt. (Table 3). Vaughan and Wiehe's (1941) detailed study of the Sideroxylon formation at Perrier Nature Re- serve revealed that two individuals of Tam- bourissa peltata and 16 of T. tau occurred in a 1,000 m? plot (Table 4), and family dominance at Perrier was only 0.4796, lowest of the three sites (Table 4). My survey of populations of these two species in the same area shows 26 individ- uals of T. peltata per 12,700 m? (= 21/ha) and 57 individuals of T. tau per 512 m? (= 1,1137 ha) (Fig. 3). The higher figure for the latter species, an understory treelet, suggests aggregation into much denser populations than T. peltata, a more highly dispersed canopy tree. Population density of these two species appears to correspond to their pollinator strategies (see discussion under Floral Biology). arts of the upland plateau and in some sheltered valleys the Sideroxylon formation passes into a high forest (“climax forest" of Vaughan & Wiehe). Vaughan and Wiehe's (1939, 1941) studies of the high forest at Macabé re- vealed the presence of 41 individuals 10 cm or more DBH belonging to three species of Tam- bourissa in a one hectare plot. In another 1,000 m? sample plot at Macabé, the genus was found to comprise 2.6796 of the total number of indi- viduals 10 cm or more DBH, with the following breakdown: 7. sieberi, 13 (upper stratum tree); T. ficus (as T. elliptica), four (mid stratum tree); T. pedicellata, 28 (mid stratum tree; my survey in 1979 failed to show this species at Macabé); T. tau (as T. amplifolia), one (understory treelet). In addition, I found 7. amplifolia to be a rare understory treelet or small tree at Macabé in 1979 (Table 3). A comparable survey was carried out in some- what drier high forest at Brise Fer (Table 3) sev- eral kilometers to the west of Macabé (Lorence et al., in prep.). Only two species of Tambourissa were found here, T. ficus and T. peltata, presum- ably due to the lower rainfall. Both showed up LORENCE— MONIMIACEAE 13 TABLE 4. Comparative frequency of Tambourissa species in three uplant plant formations in Mauritius (adapted from Vaughan & Wiehe, 1941). Number Individuals per 1,000 m? Philippia Sideroxylon | Macabé eath Wet Forest High Species (Pétrin) (Perrier) Forest T. cordifolia 44 — — T. ficus — — 4 T. pedicellata — — 28 T. peltata 1 2 — T. sieberi — — 13 T. tau — 16 1 Total 45 18 46 % Family dominance 2.04 0.47 2.67 in our study at the rate of two individuals 25 mm or more DBH per 1,000 m°. Because of the island's low altitude, cloud for- est only occurs on the flanks and summits of its highest mountains in areas of high rainfall (ca. 4,500-5,000 mm). In their study of the cloud forest (as “mossy forest") on Mt. Cocotte, Vaughan and Wiehe (1937) recorded only two species of Tambourissain a 40 m? transect: Tam- bourissa sieberi, three individuals; T. tau (as T. amplifolia), two individuals. My survey of the summit of Mt. Cocotte in 1979 (unpubl. data) revealed the presence of five additional species: Monimia ovalifolia, Tambourissa cocottensis (endemic to the mountain), 7. ficus, T. peltata, and T. tetragona (also endemic to the mountain) (Table 3). Mt. Cocotte harbors the highest con- centration of Monimiaceae in the entire island, with seven of its 11 species found here. À survey of the cloud forest on Mt. Lagrave revealed a comparable situation, with six species present. Another consequence of the island's low alti- tude is the paucity of Philippia heath formation in Mauritius, which occurs in small patches at only a few edaphically suitable areas such as Pé- trin and Mt. Laselle. Formations of this type are restricted to areas of unweathered, highly fer- Although receiving a high rainfall (4,000 mm at Pétrin), the laterite is nearly devoid of true soil, and the heath formation is exposed to constant drying winds. As a result, most woody species possess xeromorphic leaves (microphyllous, sclerophyllous, or variously pubescent). 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN % of Total 200 400 % of Total UJ ° 200 400 600 > % of Total 0 200 400 600 800 1000 [Vor. 72 1500 2500 3000 Nearest Neighbor Distance (cm) Nearest neighbor distance as percent of population for three species of dicia studied in 700 m — 26 indiv FIG š Maine 1978-1979. estimated frequency per hectare = 21.— dividuals, estimated frequency per ma = 1,11 —A. T. peltata, us Nature Reserve, sample area — 12, . T. tau, Perrier 13.—C. T. m Pétrin Nature Reserve, sample area — piene Nature Reserve, sample area — diim m?, N = 571 410 m?, N = 69 individuals, estimated frequency per hectare = 1,684 Only a single species of Monimiaceae has suc- cessfully adapted to this environment: Tam- bourissa cordifolia, a low dioecious shrub with small, sessile coriaceous leaves. It is peculiar in being the only truly shrubby member of the genus known to me. In their study of a 1,000 m? tran- sect at Pétrin, Vaughan and Wiehe (1941) found 44 individuals of T. cordifolia. My study of a population of the species at the same locality gave a much higher dias (Fig. 3), 69 individ- uals per 410 m? (= 4/ha), suggesting the species is aggregated s dense, local popula- 1985] J;4-—dH 1 afew tions. aber regions with similar edaphic conditions, e.g., Mt. Laselle, Les Mares, and Mare Longue Plateau (Fig. 40). Although Tambourissa peltata, and to a lesser degree 7. tau, also occur in the heath formation at Pétrin, , they are only marginally successful and OCcur a te, stunted individuals. In their 1,000 m? sample, Vaughan and Wiehe (1941) only found a single individual of Tambourissa peltata, although I found it to be more abundant = ;. areas. Family dominance for Mo- aceae in this habitat was 2.04% of the total a DU (Table 4), somewhat less than at Ma- cabé but still a significant element of the woody flora (Vaughan & Wiehe, 1941) Areas lying to the west of the central plateau, i.e., the western escarpment and lowland areas in the rain shadow (1,000-1,600 mm precipita- tion), experience a distinct dry season and sup- port a semideciduous dry forest of Mauritius Ebony (Diospyros tessellaria Poir.), Mimusops, and Elaeodendron (Celastraceae). Of the island's ten species of Tambourissa, only T. quadrifida occupies this habitat to which it is restricted, e.g., at Yemen Valley (Table 3). The species occurs here as a mid stratum or canopy tree often form- ing small, localized populations. Ithough many Monimiaceae readily sprout sucker shoots from cut stumps, no species were even in genous forest highly invaded by vig- hould be considered as threatened species. For exam- ple, on Mt. Cocotte, which is heavily invaded by numerous exotics, only limited regeneration was observed in Tambourissa ficus and T. peltata, two of the commonest species. No signs of recent regeneration, however, were apparent in the rare 4 many S bourissa peltata and T. tau was observed at Per- rier Nature Reserve, which has been kept free of exotics for the last decade by a stringent weeding program carried out by the Forestry Department. In summary, on one hand, Mauritius has fa- vored the greatest adaptive radiation in Tam- bourissa of any of the satellite islands around Madagascar, probably as a result of its compar- atively greater age coupled with the availability of a range of suitable habitats. Monimia, on the other hand, is represented by a single species confined to the island's few patches of cloud for- LORENCE— MONIMIACEAE 15 est. Its poor representation may be attributable to a lack of suitable habitats or a relatively recent introduction. RODRIGUES Rodrigues, the smallest, most isolated, and possibly youngest of the Mascarene islands, only attains ca. 400 m maximum elevation and is the least diverse ecologically. As a result of its ex- treme isolation and paucity of suitable habitats, Monimiaceae appear to be absent from the island (e.g., Balfour, 1879; Wiehe, 1949; J. Guého, pers. comm.). A single collection of Monimia ovali- folia supposedly from Rodrigues is probably wrongly labelled (see discussion under the genus Monimia) COMORE ARCHIPELAGO The Comores are poorly known botanically but appear to harbor predominantly African and Madagascan floristic elements (Voeltzkow, 1917). Of the four islands, information is available only for Grande Comore (Legris, 1969) in the form of a survey of vegetation types and enumeration of their most common and conspicuous ele- ments GRANDE COMORE Grande Comore, youngest of the islands, has virtually no intact lowland forest remaining. Areas of well-developed lower montane wet and montane cloud forest still remain on the lower b ere and mid altitude slopes beginning at . 500-800 m, bou notably on the Kar- a vol rille massifs where annual precipitation ranges gs ca. 3,000 to 5,000 mm (Legris, 1969). Of recent volcanic origin, the La Grille massif supports mature wet forest domi- nated by large individuals of Weinntannia, An- thocleista, Eugenia, and M. 0-30 m, and also harbors the island’s only e S8 constitutes an important element of the forest, conspicuous because of its numerous large, brown cupuliform fruits hanging from the trunk and major branches (Fig. 1 Wet forest on the Karthala volcano massif is dominated by Ocotea, Nuxia, Olea, and Aphloia, but no Monimiaceae have been noted or col- lected there to my knowledge. A cloud zone often 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN forms above ca. 1,000 m. At ca. 1,800-2,000 m the wet/cloud forest zone on Karthala gives way to an ericoid heath formation of Philippia similar to that in Madagascar and Réunion; no Monimi- aceae are known from this zone. The leeward western region of the island prob- ably once supported dry forest, of which huge baobabs (Adansonia digitata L.) and other scat- tered trees are the only remnants. No Monimia- ceae are known from this zone either. ANJOUAN Unfortunately, I was unable to visit Anjouan, which is extremely poorly known botanically. Most of its lowland forest has been destroyed, although the mountainous central massifs (e.g., N'Tingui) support fairly extensive areas of wet and cloud forest due to their rugged relief (I. Tattersall, pers. comm.). Two species of Tam- bourissa, T. kirkii and T. paradoxa (syn. T. jo- hannae) are known from Anjouan, but little is known of their ecology and distribution. Both apparently occur in the wet and cloud forest zones (Fig. 35). Although Anjouan only attains ca. 1,600 m, it may harbor small amounts of heath vege- tation on its highest peaks. Dry forest probably occurred on its dry, lower leeward slopes as is the case for Mayotte and Grande Comore. MOHELI Mohéli, smallest of the Comores, is also vir- tually unknown botanically. Although I was un- able to visit it, I. Tattersall (pers. comm.) stated that several extensive areas of montane wet for- est still remain on its central ridge at Mt. St. Antonio, and between Fomboni and Drondoni (ca. 500-700 m elevation). Collections from these areas made by L. Bernardi in 1967 have revealed the presence of a new endemic species of Tam- bourissa, T. moheliensis, and apparently the only one to occur there. Little is known of its distri- bution and ecology (Fig. 35). Most of Mohéli's lowland vegetation is now gone. Presumably wet forest occurred on the windward slopes and dry forest on the lower slopes in the rain shadow, as is the case for Mayotte and Grande Comore. MAYOTTE Oldest and lowest of the Comores, Mayotte supports only a few small areas of indigenous forest, the rest being cultivated or secondary in [Vor. 72 nature. Several of the island's highest mountain massifs, i.e., Benara (650 m) and Hachiroungou (500 m), support wet forest dominated by Nuxia, Aphloia, and Macaranga similar to that found he miaceae species, d diei leptophylla, still occur here (Fig 35) An r of large mature individuals and signs of puse regeneration were observed on Benara. Tambourissa leptophylla also occurs sporadically as a remnant in wet lowland sec- ondary forest at Barakani and Sada. In the dry, western leeward region remnants of dry forest with Adansonia digitata occur on isolated peaks and offshore islets. No Monimiaceae were found to occur there. In essence, although each of the Comores sup- ports its own single endemic species of Tam- bourissa (two in Anjouan), adaptive radiation of the genus here has been less extensive than in the Mascarenes or Madagascar. As habitat di- versity in the Comores is roughly comparable to that in the Mascarenes in spite of their smaller average size, the low Tambourissa species di- versity may be due to the islands' relative youth or possibly a more recent date of introduction of the genus. That four ofthe five Comorean species appear to be closely related, and that all species for which data are available are restricted to the wet/cloud forest zone (none are adapted to either dry forest or heath formation), suggests a rela- tively short history for Tambourissa in the Co- mores This review of the ecology and distribution of the Monimiaceae in the Malagasy region dem- onstrates that members of the family are floristi- cally important constituents of the islands' plant formations. Furthermore, the various species have different life forms ranging from shrubs or understory treelets to nds Brace Or even can- opy trees. The family test diversity and abundance in the lower montane wet and onn cloud forest zones, although a signifi- cant number of species are adapted to the low- land wet forest zone. Only a few species are adapted to the high altitude heath formation and the semideciduous dry forest, however. Al- though many species of Tambourissa have the ability to form coppice shoots from cut trunks, they are virtually absent from secondary vege- tation and exhibit a poor ability to regenerate when competing with vigorous, invasive exotic species 1985] MORPHOLOGY AND ANATOMY Because they are generally considered to be a key family for the critical interpretation of a Lauraceae), the Monimiaceae have tively well studied morphologically and anatom- ically (Baillon, 1871; Hobein, 1889; Perkins, 1898, 1925; Perkins & Gilg, 1901; Metay, 1921; Garratt, 1934; Money et al., 1950; Lemesle & Prichard, 1954; Sampson, 1969a, 1969b, 1969c; Endress, 1972, 1980a, 1980b). However, apart from a few species in the genera Monimia, Tam- bourissa, and to a lesser degree Decarydendron, the species in the Malagasy area have not been well studied morphologically or anatomically. METHODS AND MATERIALS Gross morphological observations were made from herbarium specimens and collections pre- served in alcohol. For venation studies, dried leaves were rehydrated, cleared in KOH, bleached, stained in safranin, dehydrated, and mounted in Permount resin between glass plates according to the method of Dilcher (1974). Prints of venation patterns were made using a photo- graphic enlarger and standard photographic print paper (f3, f4, or f5). Leaf trichomes were pre- pared for the SEM by Au-Pd sputter-coating and photographs were taken on a Cambridge Stereo- scan Mark II. Material for floral anatomical stud- ies was fixed in alcohol or FAA (Monimia, most piandra) after which it was dehydrated in a TBA series, embedded in Paraplast, sectioned on a rotary microtome, dewaxed and stained with saf- ranin and fast green, and mounted according to methods in Jensen (1962). In some cases, carpels were cleared in KOH, stained, dehydrated, and mounted in Permount to observe the vascula- tion. HABIT AND BARK Monimiaceae in the Malagasy region are most- ly understory to mid stratum treelets or small trees, or large canopy trees. I know of no species that occur as emergents. The shrubby habit is quite rare, and no true vines are known from the region. The boles of all species I have observed were cylindric, often with knobby meristematic swellings in the cauliflorous species. The trunk LORENCE—MONIMIACEAE 1 7 is basally swollen in Tambourissa ficus, Tam- bourissa longicarpa, and Decarydendron perrieri, where the flowers are generally produced. None of the species are known to have buttresses or prop roots. Species of Monimia are small- to moderate- sized mid stratum or canopy trees, often shrubby hen growing in exposed habitats. Both M. am- plexicaulis and M. rotundifolia reach ca. 10 m tall, whereas M. ovalifolia is known to attain 15 m. Trunks of old individuals of all three species are often somewhat reclining and may reach di- ameters of 30-40 cm in M. amplexicaulis and M. ovalifolia, and up to 60 cm in M. rotundifolia. The bark is generally pale brown or reddis brown, soft, smooth, and flakes off in patches. Because the species are confined to very humid habitats, the bark is often overgrown with bryo- phytes and other epiphytes. Of the Madagascan endemic genera, Decary- lend d Ephippiandi , treelets or trees to 10-25 m tall (e.g., E. madagascariensis and E. tsaratanensis). No information is avail- able on the boles or bark of these two genera. Tambourissa includes species with a wide range of woody life forms ranging from shrubs rarely exceeding a meter (7. cordifolia) to large canopy trees 10-20 m tall (e.g., 7. comorensis, T. sie- beri) The majority of species, however, are understory treelets (e.g., T. amplifolia, T. tau) or small mid stratum trees (e.g., 7. ficus, T. pur- purea). The only reportedly vining species is 7. madagascariensis, a “lianescent shrub" (Raba- boto sub RN 6028, P), which may also be a large tree 15 m tall (Anon. sub herb. Alaotra 2126, MAD). Except for the shrubby 7. cordifolia, most species are monocaulescent, although many are capable of forming multiple stems from coppice shoots (e.g., T. cocottensis, T. tau, T. quadrifida), after death of the main stem. Although often licheniferous or covered with bryophytes, the bark of most species of Tam- bourissa is pale grayish or brownish, soft and corky, smooth to longitudinally fissured, and flakes or peels off in strips or patches. For the 16 species examined, characteristics of the slash (an oblique cut through the outer and inner bark) include a beige or pale pink to salmon-colored inner bark (phloem), and whitish or straw col- ored wood (xylem), both showing the conspic- uous wide rays characteristic of many Monimi- aceae. No sap or exudate was apparent. In summary, the Malagasy area Monimiaceae 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN are treelets or trees most abundant in the canopy and lower strata of tropical forests. Only a small number of species have shrubby or lianescent habits. The bark of Monimia and many species of Tambourissa is relatively uniform, but needs further study in Decarydendron and Ephippian- ra. WOOD ANATOMY Anatomical studies of the wood of represen- tative genera and subfamilies of Monimiaceae conducted by Garratt (1934), Money et al. (1950), and Lemesle and Prichard (1954) reveal a num- ber of trends of specialization. Most striking are progressive changes in the vessel members which include a shortening and broadening, a decrease in the degree of end wall overlapping, and a tran- sition from scalariform to simple perforation plates. Of the four genera examined from the Mala- gasy area (Money et al., 1950), Decarydendron, Ephippiandra (as Hedycaryopsis), and Tam- bourissa (all Mollinedioideae sensu Thorne) pos- Ing relatively unspecialized within the family. These include long, thin-walled vessel members with angular, extensively overlapping end walls, and in T. ly genus for which mature wood was available) salan om perforation plates (Garratt, 1934; Money et al., 1950). The xylem of Monimia (Monimioideae sensu Thorne) exhibits a number of trends that have been interpreted as advanced within the family Money et al., 1950). Vessel members of M. ovalifolia are relatively short with somewhat ob- tuse end walls that do not overlap extensively and have perforation plates transitional between scalariform and simple porous. In this respect, specialization in the xylem of Monimia (and also Palmeria — Mollinedioideae) has attained a level comparable to that of Peumus (also Monimioi- deae), eid with simple perforation plates. ommon wood anatomical features shared by both the Monimioideae and the Mollinedioideae are: scanty wood parenchyma; rather broad, multiseriate rays; presence of septate fibers; pres- ence of fibers and hippocrepiform sclerids in the stem pericycle (Money et al., 1950). STEMS AND PHYLLOTAXY Adult phyllotaxy in Monimia, Decaryden- dron, Ephippiandra, and most species of Tam- [Vor. 72 bourissa is opposite to subopposite and decus- e The three Tambourissa species in species Group 8 have ternate to subalternate adult leaves (Table 1), although seedlings of two, 7. ficus and T. peltata, have opposite decussate leaves. On the other hand, adult leaves of species which are normally oa are sometimes ternate on vig- orous sucker shoots (e.g., Monimia ovalifolia, Tambourissa purpurea, and T. pedicellata). Al- though opposite and decussate, the adult leaves of T. tau (Fig. 39A) are unique in being clustered into pseudoverticels of two to four pairs, sepa- rated by long “internodes.” Leafy stems of most species of all four genera are more or less terete with flattened, somewhat dialated nodes. Tambourissa tetragona is unique in having winged, quadrangular stems, whereas those of the ternate S species (Group 8) are somewhat trigono Stems of DAD ERROR Ephippiandra, and most Tambourissa species bear sparse to dense, straight or undulate, simple or rarely fasciculate e.g., T. comorensis, T. mohelensis) lignified tri- chomes. In a few species even the new growth is virtually glabrous (e.g., 7. cocottensis, T. cordi- folia, and T. tetragona). In Monimia, the stems are scabrous to velutinous with an indument of stellate, fasciculate and simple trichomes. Len- ticels often occur on the stems of certain species (e.g., T. ficus, T. peltata, E. perrieri, E. domatia- ta, and E. myrtoidea) but are absent in most. Stem characters thus provide a number of species- specific, taxonomically useful features. Monimia, Decarydendron, Ephippiandra, and Tambourissa all characteristically possess mul- tiple axillary buds of which I found no mention in the literature. In all three species of Monimia (as in Peumus), three horizontally oriented buds are usually present in each leaf axil, the central one being largest and developing first (if at all). The vegetative buds have small, pubescent scales, whereas those enclosing the inflorescence are large, brown, and subglabrous. In the remaining genera, all Mollinedioideae, the buds are orient- ed longitudinally in the leaf axil, the central one usually being largest and the first to develop (if at all). On flowering branches of the ramiflorous species E. madagascariensis and E. tsaratanen- sis there are three to five buds, the central one usually giving rise to the inflorescence and the lateral ones to vegetative shoots. In D. helenae var. stenophyllum the buds occur in groups of three, whereas in E. microphylla and E. myrtoid- 1985] LORENCE—MONIMIACEAE 19 ea they are solitary or in pairs. Three, less com- monly two, axillary buds are present in nearly all species of Tambourissa examined, regardless of whether the inflorescence is terminal, rami- florous or cauliflorous. On flowering stems of ramiflorous species such as 7. purpurea, the large central bud frequently but not invariably gives rise to the inflorescence. On vegetative stems, the central bud usually develops first, although two buds may occasionally develop into vege- tative shoots simultaneously. Tambourissa te- tragona is exceptional in that only one bud is obvious. However, vigorous sucker shoots of 7. ficus may produce up to six buds extending in a row for several centimeters above the leaf axil (Lorence 2235, MO) LEAVES The terminology used to describe leaves in this monograph follows Hickey (1973, 1977), Hickey and Wolfe (1975), and to an extent Lawrence (1951). Leaves of all Monimiaceae are exstipu- late, considered by Hickey and Wolfe (1975) to be an advanced condition within the Laurales. Characters of the petiole and lamina, including size and shape, margin, venation, and pubes- cence, provide a id of taxonomically useful features in many t Petiole. The duis, of most genera and species have a well-differentiated lamina and petiole, but Monimia eee Ate eee (Fig. 4A) and Tam- g. 8B) are unique in having sessile cunis apiid leaves. Petioles of most other species are slender to stout, of vari- able length but much shorter than the lamina, and are adaxially flattened or canaliculate. They are either glabrous or bear an indument of tri- chomes similar in type and density to that of the stem internodes. In all three species of Monimia (M. amplexicaulis, Lorence 2503, MO; M. ovali- folia, Lorence 1857, MO; M. rotundifolia, Lo- rence 2472, MO), the vascular strands of the pet- iole are united into an adaxially flattened cylinder (see also Money et al., 1950). In the other genera which I examined (Ephippiandra myrtoidea, Baron 1355, K; E. madagascariensis, Bernardi 11977, MO), and six species of Tambourissa (T. cocottensis, Lorence 2287, MO; T. crassa, Lo- , MO; T. elliptica subsp. micrantha, ? ; T. ficus, Lorence 2162, MO; T. pedicellata, Lorence 2385, MO; and T. tetra- gona, Lorence 2286, MO), the petiolar vascular strands are aggregated into a shallow, U-shaped hmuricecn arc. This corresponds with Metay's (1921) find- ings for 7. floricostata (as Schrameckia mada- gascariensis) and T. thouvenotii. Lamina. Leaves of Monimia are subcoria- ceous to coriaceous, usually ovate to elliptic or suborbiculate, and their size and shape are gen- erally much more polymorphic in M. ovalifolia and M. rotundifolia than in M. amplexicaulis. Although the extreme forms appear to be very distinct (Fig. 4), leaf size and shape in Monimia do not provide stable taxonomic features. Like- wise, shape of the apex is quite variable (acu- minate, acute, or obtuse) and of little taxonomic importance in all three species. Characters of the leaf base, however, are useful in distinguishing . amplexicaulis, where it is deeply cordate and amplexicaul, from the other two species where it ranges from shallowly cordate to rounded, ob- tuse, acute or decurrent (Fig. Leaves of the three species of Decarydendron do not differ markedly from each other, ranging from elliptic to oblong or obovate. The texture is chartaceous to subcoriaceous, the apex acute or acuminate, and the base cuneate. Leaves of the six species of Ephippiandra are small- to medium-sized and chartaceous to co- riaceous in texture. Laminar shape is generally characteristic for each species: ovate or elliptic with acute or acuminate apices and cuneate to rounded bases in E. microphylla and E. myr- toidea (Fig. 6A); elliptic with a retuse apex folded downward into a flap in E. perrieri (Fig. 6B); obovate-elliptic to rhombic and apiculate in E. domatiata; orbicular with an obtuse to truncate d gascariensis and E. tsaratanensis. (Fig. 6E, F). The 43 species of Tambourissa display a wide array of leaf morphologies which are relatively constant and species-specific. In widespread species with a broad ecological amplitude (e.g., T. peltata, T. purpurea), considerable intraspe- cific variation occurs in size and shape, generally between populations growing in different habi- ats. Texture ranges from membranaceous in T. leptophylla to coriaceous in T. parvifolia and T. religiosa. Size varies from small in T. parvifolia (attaining only 47 mm by 22 mm) which grows in high altitude montane forest, to very large in T. amplifolia (attaining 500 mm by 240 mm) which grows in lowland wet forest. Leaf shape varies considerably within many species but in others it is of considerable taxonomic signifi- cance, e.g., in distinguishing the two subspecies of T. elliptica. Leaf shape in the genus ranges ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURE 4. Leaf clearings of Monimia. — A. M. amplexicaulis, Rivals s.n. (TL-R). — B. M. rotundifolia, Rivals s.n. in June 1943 (TL-R).—C. M. rotundifolia, Bosser 21332 (P).—D. M. ovalifolia, Cordemoy s.n. (TL-R).— E. M. ovalifolia, Guého sub MAU 14130 (MAU). Bar equals 25 mm. 1985] from cordate or ovate to elliptic, oblong, or ob- ov ate. Studies ofleafanatomy in Monimia and Tam- bourissa (Hobein, 1889; Metay, 1921) show their laminar structure to be similar to that of many bifacial dicot leaves. The two species of Monimia and six species of Tambourissa examined by them all have a cuticle, an upper epidermis, a hypo- dermis consisting of one to six cell layers, a me- sophyll differentiated into a palisade layer one or two cells thick and a spongy layer many cells thick, and a lower epidermis also covered by a cuticle. Stomata are restricted to the lower epi- dermis in all species except 7. thouvenotii, which also has stomata in the upper epidermis (Metay, 1921 Leaves of Monimiaceae are characterized by the presence of spheroid laminar glands or se- cretory cells containing so-called “ethereal” oils which impart a particular odor to many species. In most species of Tambourissa these are con- centrated in the mesophyll, but lesser numbers also occur in the lower epidermis and hypoder- mis (Hobein, 1889; Metay, 1921). Hobein (1889) stated that both Monimia ovalifolia and M. ro- tundifolia have secretory cells restricted to the hypodermis and not in the mesophyll as in Tam- bourissa and the other genera he studied. M investigations of hand sectioned, alcohol fixed leaves show that secretory cells are restricted to the hypodermis only in M. ovalifolia, both in plants from Mauritius (Lorence 1857, 2896, MO) and from Réunion (Lorence 2757, MO). In M. amplexicaulis (Lorence 2503, MO) and M. ro- tundifolia (Lorence 2472, MO), secretory cells were found to occur not only in the hypodermis, but also in the interface between the palisades and spongy mesophyll. An examination of sec- tions of restored leaves of Decarydendron hele- nae var. stenophyllum ( Humbert 6613, K), and E. myrtoidea (Baron 1263, K) shows that they are fundamentally similar to species of Tambourissa anatomically. Venation and margin. The Monimiaceae ossess several distinctive trends which Hickey 9 an indurated cap, and secondary veins originat- ing at uniform angles to the costa. All three species of Monimia possess stoutly brochidodromous venation ofthe festooned type characteristic of the family. Three to four pairs LORENCE— MONIMIACEAE 21 FiGuRE 5. Camera lucida drawing of ultimate ve- nation in cleared leaf of Monimia ovalifolia, Bijoux sub MAU 1310 (MAU). 2° = secondary venation, 3? = tertiary venation. of secondary veins originate at comparatively uniform angles, are bowed to geniculate, follow a regular, slightly sinuous course and delimit rel- atively regular and uniform intercostal areas (Fig. 4A-E). Consequently, all three species fall into Hickey's (1977) second rank leaf category. Dis- tally the secondaries continue to form another, higher order of loops characteristic of the fes- tooned brochidodromous venation pattern (Hickey & Wolfe, 1975). The numerous inter- marginal loops of the last order unite to form a weak, fimbriate intramarginal vein generally lacking in the other genera. Intercostally, the tertiary veins originate at right angles from the secondaries and their arches and unite to form a series of strong, transverse ter- tiaries which in turn ramify at right angles and then anastomose admedially to form composite thogonal (rectangular or pentagonal) and much more regular than in the other genera (Figs. 4, 5). The quaternaries arise from the tertiaries at nearly right angles to form fairly regular rect- angular areoles. Vein orders above the quater- 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN nary are generally indistinct and ramify to form a reticulum of irregular areoles with included veinlets that anastomose or remain free (Fig. 5). Terminal idioblasts appear to be absent from the veinlets. The number and pattern of veins in Monimia provide few characters useful in sep- arating the species, although they are character- istic of the genus as a whole Adult leaves of all three Monimia species have entire, generally slightly revolute margins. Seed- ling leaves of M. ovalifolia and M. rotundifolia (unknown in M. amplexicaulis) are also entire, as opposed to those of Tambourissa (D. Lorence, pers. observ.) and Hortonia (Endress, 1980a) which are generally dentate. Although the Mo- nimioid tooth is, ironically, absent i in Monimia, sibly have undergone loss of teeth secondarily, as appears to be the case for Tambourissa. Venation in all three species of Decarydendron is festooned brochidodromous. The six to 12 pairs of secondaries arise at quite constant angles, de- limit regular intercostal areas, and continue to to descend obliquely across the intercostal area. The quaternaries arise from the tertiaries at irregular angles and delimit irregular areoles. Vein orders above the quaternary are indistinct, and proba- bly ramify to form free, included veinlets. Adult leaves of all three species of Decarydendron are serrate-dentate along the greater portion of their plane margins, each tooth ending in an indurated cap. An unbranched distal excurrent vein arising from the outer series of brochidodromous arches runs directly into each tooth. In this respect the venation is not truly semicraspedodromous ac- cording to Hickey’s (1973) definition. Species of the genus Ephippiandra (including Hedycaryopsis) display a distinctive trend from festooned brochidodromous to specialized and highly organized semicraspedodromous and craspedodromous patterns, the latter unknown elsewhere in the family. Leaves of E. micro- phylla, E. myrtoidea, and E. perrieri are small with only four to six pairs of secondary veins s or somewhat obliquely across the in- tercostal areas, often ramifying into strongly [Vor. 72 admedial branches as they approach the costa. The tertiaries delimit irregular intercostal areas in E. myrtoidea, but these are more regular and orthogonal in E. perrieri. Higher vein orders above the tertiary are not very distinct and gen- erally form a more or less random reticulum of irregular areoles, ultimately terminating in free, dendroid veinlets lacking idioblasts. Margins are entire and somewhat revolute in E. myrtoidea and E. perrieri, but those of E. microphylla gen- erally have a single pair of small teeth in the apical portion arising from exmedial branches of the secondary arches Leaves of E. domatiata (Fig. 6C, D) are like- wise small with two to hoa pairs of secondaries, but show certain di craspedodromous pattern. Although ihe weak basal pair of secondary veins forms a series o festooned loops in the customary brochidodro- the next pair above. On each side of the lamina one of the secondary branches runs directly into the margin, terminating in a small Hon The tertiaries d l although the quaternaries form more regular ar- eoles with free, branching veinlets. The margin is moderately revolute, and it is the only species known to have barbate domatia in the basal sec- ondary vein axils, hence the specific epithet. phippiandra tsaratanensis (Fig. 6E) has much larger, broader leaves which show a weakening of the brochidodromous pattern and the for- mation of sunken marginal teeth. As each of the four to seven pairs of secondary veins bifurcates, each branch generally unites with an adjacent branch to form a weak arch whose exmedial por- tion in turn gives rise to one or more distal branches. Each of these runs to the margin e it terminates in a rounded, centrally depressed tooth. The intersecondary webbing pattern of the tertiaries is stronger than in the preceding species, with more regular orthogonal, often quadran- gular, tertiary areoles frequently bisected by zig- zagging admedial branches. The quaternaries form generally small, more or less orthogonal areoles with free, included branched veinlets. Leaves of the sixth species, Ephippiandra madagascariensis (Fig. 6F) are the most highly specialized of all, being almost orbicular with four to five pairs of secondary veins forming an open, craspedodromous pattern due to a further weakening of the secondary arches. Each sec- ondary departs from the costa at a uniform angle t fair ly ir I egular LORENCE— MONIMIACEAE FiGuRE 6. Leaf clearing of Ephippiandra. — A. E. myrtoidea, Baron 1355 (K).—B. E. perrieri, Morat 2332 (MAD).—C, D. E. domatiata, Capuron 8828-SF (P). —E. E. tsaratanensis, Gentry 11605 (MO).—F. E. mad- agascariensis, Bernardi 11977 (MO). Bars equal 10 mm. 24 and generally bifurcates once or rarely twice be- fore the (usually basal) branch terminates in a single, depressed tooth as in the previous species. Occasionally, an apical sub-branch unites with a superadjacent secondary. As in the latter species, the tertiaries form a strong intersecondary cros webbing pattern delimiting regular intercostal areas. The lertisries nearest the costa frequently display a strongly tern, whereas the quaternaries s generally form i ir- regular areoles with highly branched, free in- cluded veinlets as in the other species. The margin of E. madagascariensis is sinuate-dentate with seven to nine pairs of sunken teeth, including one at the apex, as in the preceding species. These teeth contain terminal tracheary elements and appear to be hydathodal. A situation analogous to that in Ephippiandra occurs in Hibbertia (Dilleniaceae). Rury and Dickison (1977) explain the ontogenetic transi- tions in Hibbertia as follows: “The craspedodromous venation pattern appears ri rn. iran epa gwa venation apparently tion and u th v - ochidodromous arch in leaves of Hibbertia SUEDE etc. This same transition is apparent in the six species of Ephippiandra: a weakening and open- ing of the brochidodromous pattern with sub- sequent vascularization of the teeth from the sec- ondary branches. This transition from wea brochidodromy to simple craspedodromy could result from a differential thickening of the por- tion which connects the basal part of the sec- ondary arch with the marginal tooth, which can be seen in the basal veins of E. madagascariensis. According to Rury and Dickison (1977), Hickey (pers. comm.) stated that the ontogenetic tran- sition from semicraspedodromy to simple cras- pedodromy has resulted from such a shift in vein development in many dicot leaves. Tambourissa species (including Phanerogono- carpus) have venation patterns based on the fes- tooned brochidodromous theme. Number of sec- ondary veins ranges from three or four pairs (e.g., T. pariijalia, Fig. 8C; T. elliptica subsp. elliptica, ), to six to 12 a subsp. micrantha (Fig. 7E). The Ord are stout, straight or sinuate, and delimit intercostal areas which may be ir- >= ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 regular (T. thouvenotii, Fig. 7A), or quite regular (T. elliptica subsp. micrantha, Fig. 7E; T. cor- difolia, Fig. 8B). Species with fewer secondary veins frequently have less regular intercostal areas; some of these approach Hickey's (1977) “first rank" leaf category in this respect (e.g., 7. thouvenotii, Fig. 7A). The majority of species, however, fall into his “second rank" category. In these, the secondaries from a fairly strong series and continue to loop or festoon for several orders. The ultimate ex- medial series of loops is often irregular and dis- continuous (e.g., T. leptophylla, Fig. 8E), and does not form a true marginal fimbrial vein, although the strong intramarginal loops of 7. religiosa re- semble a fimbrial vein (Fig. 8D). The tertiary veins arise at wide acute to slightly obtuse angles from the secondaries and their arches, and are often admedially and obliquely ramified, form- ing strong, composite EL (e.g., T. elliptica subsp. micrantha, Fig. 7E). Tertiary ar- eoles are generally irregular, sigue orthogonal areoles may be delimited by transverse tertiaries in some species with more highly organized ve- nation (e.g., 7. peltata, Fig. 8A; T. elliptica subsp. elliptica, Fig. 7D). Higher order veins are gen- erally indistinct above the quaternar d some- times tertiary levels. The quatemaries g generally ramify and anastomose to form gular areoles Bode: ic include ° = he margins of adult leaves of the majority of Tambourissa species are entire and plane to rev- olute. Exceptional are 7. thouvenotii, T. tricho- phylla, and the two species formerly included in Phanerogonocarpus, i.e., T. alaticarpa and T. longicarpa. The former two possess half-toothed leaves with one to eight pairs of Monimioid teeth in the apical portion of the lamina. In 7. tricho- phylla (Fig. 7C), the secondary arches are strong and delimit low, broad intercostal areas, with an therefore semicraspe notii (Fig. 7A), the secondaries delimit high, nar- row intercostal areas and the arches are much weaker, approaching the semicraspedodromous pattern of Ephippiandra. In some leaves of T. thouvenotii the main, apical secondary branch runs directly into the tooth, and any festooning is very weak. In others, it more closely resembles the pattern in 7. trichophylla. Leaves of the two species formerly included in Phanerogonocarpus (T. alaticarpa and T. lon- LORENCE—MONIMIACEAE Fi y . Leaf clearings of Tambourissa.—A. T. thouvenotii, Thouvenot s.n. in 1919 (BM).—B. T. ficus juven om sucker shoot, Richardson 4042 (K).—C. T. trichophylla, Lorence 1879 (MO).—D. z elliptica ipp. as Friedmann 2299 (P). —E. T. elliptica subsp. micrantha, Coode 4956 (K). Bar equals 25 m ANNALS OF THE MISSOURI BOTANICAL GARDEN RE 8. Leaf clearings of Tambourissa. —A. T. peltata, Guého sub MAU 12543 (MAU).— B. T. cordifolia, Fi Lorence M 144 (MO).—C. T. A ata Baron 4204 (K).— D. T. religiosa, Decary 6554 (P). —E. T. leptophylla, Tattersall s.n. (MO). Bar equals 25 m 1985] gicarpa) have five to eight pairs of secondary veins arising at quite uniform angles which de- limit fairly iig intercostal areas and unite in a festooned brochidodromous pattern. In 7. a/a- ticarpa, the apical 25A of the margin is dentate with eight to ten pairs of slender, antrorse teeth (Fig. 17A). That of T. longicarpa is broadly den- tate with two or three pairs of large teeth in the apical !^—'2 of the lamina. In both, the teeth arise from exmedial branches of the secondary or ter- tiary arches, thus rendering the venation semi- craspedodromous. In the context of Rury and Dickison's (1977) findings for Hibbertia where teeth develop in the adult leaves, it would seem that the dentate species of virtually all species of Tambourissa, whic have seen, produce apically dentate leaves. This condition frequently also occurs on sucker shoots (e.g., T. ficus, Fig. 7B) where they are gradually lost in the ontogenetic transition to entire adult leaves. These Tambourissa species therefore ap- pear to lose their teeth secondarily, as also occurs in Hortonia (Endress, 19802). In this context, the four dentate species may represent cases of neo- teny. This does not, however, contradict Hickey and Wolfe's (1975) suggestion that the Monim- oid tooth is an advanced feature within the fam- TRICHOMES Apart from the three virtually glabrous Tam- bourissa species mentioned above, stems and leaves of most species in all four genera bear at least some trichomes. Leaf pubescence ranges from a few scattered hairs along the petiole and major veins, to the entire lamina being canescent or velutinous-tomentose. Stem pubescence is comparable to that of the leaf. Trichomes of all the genera examined are nonglandular, lignified (e.g., stain pink or red with phloroglucinol; Ho- bein, 1889), and are simple and unicellular (ex- cept in Monimia). In a few Tambourissa species they are fasciculate, i.e., grouped into clusters of two or three. Three types of trichomes occur in Monimia: long, unicellular; multicellular, fasciculate con- sisting of two to eight long, ascendent, unicellular arms united basally to form the stalk; short, ap- pressed, multicellular stellate or stellate-peltate composed of six to 16 horizontal arms radiating from a common center and joined basally to form LORENCE—MONIMIACEAE 27 the stalk. Trichome density is greatest on the abaxial laminar surface where they form a thick, whitish indument completely obscuring the leaf surface (Fig. 9E), whereas they are scattered and more easily distinguished on the adaxial surface. Trichomes play a key role in distinguishing the three species of Monimia. Monimia ovalifolia is distinct in having small, appressed, stellate-pel- tate trichomes with six to 16 short arms united basally for 4-2 their length (Fig. 9D, E), a char- acter shared by populations both from Mauritius and Réunion. In addition to stellate trichomes with fewer long, free arms, M. amplexicaulis and M. rotundifolia also have simple and fasciculate trichomes (Fig. 9A—C). Although both the latter species may display considerable variation in tri- chome composition, even a a single pop- ulation, they are modally distinct. n the lin po irae. E n) doma- tiata, E. modica 8 iensis, E. microphylla, E. myrtoidea, and E. perrieri are all sparsely pu- bescent or subglabrous with short, pale simple trichomes, whereas E. tsaratanensis is easily dis- tinguished by its long, dense simple reddish tri- chomes. Trichomes of Decarydendron are sim- ple, yellowish, and generally longer than in Ephippiandra. Leaf pubescence in Tambourissa ranges from completely glabrous in a few species (e.g., T. cocottensis, T. tetragona, species Group 5), to subglabrous (e.g., T. tau, species Group 5), canescent or villous in many (e.g., T. nosybensis, T. tonii species Group 1B), or even densely velutinous-tomentose in a few (e.g., T. thouvenotii, T. trichophylla, species Group 1A; ongicarpa, species Group 8). Trichomes isture constitute a b useful char- acter in the Monimiaceae. INFLORESCENCE Inflorescence structure in the four genera is quite variable: thyrsic, cymose (pleiochasial, di- chasial, or monochasial), fasciculate, or solitary. Although more than one inflorescence type fre- quently occurs within a given species, and often on a single individual, the inflorescence generally provides a taxonomically useful suite of char- acters. Flowers of all the genera and species in this study are unisexual, and sexual makeup of the inflorescence is often stable and character- varies greatly within the genera, but is frequently 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURE 9. Scanning electron micrographs of Monimia leaf trichomes.— A. M. amplexicaulis, adaxial leaf surface, Capuron SF-28210 (P).—B. M. rotundifolia, adaxial leaf surface, Friedmann 781 (P).—C. M. rotun- difolia, adaxial leaf surface, Bosser 21332 (P).—D. M. ovalifolia, adaxial leaf surface, Guého sub MAU 14097 (MAU).—E. M. ovalifolia, abaxial leaf surface, Friedmann 1884 (P). Bars equal 0.5 mm. 1985] LORENCE-— MONIMIACEAE 29 FicunE 10. Monimia inflorescence types, hypothetical reduction series showing derivation of different types. A.I Flowers are depicted as being in one plane, whereas they are actually decussately arrang ed. — A. Indeterminate 0 leafy shoot with axillary thyrses. — B, C. Determinate leafy thyrses. — D. Thyrse. Bar equals ca. 50 mm. modally distinct for given species. A brief review of the inflorescence characters will be given for each genus. Observations on numerous herbarium speci- mens and living plants reveal that all three species of Monimia are dioecious. Inflorescences of all th i t l l hi d have been variously described as panicles, umbels, ra- cemes (e.g., Baker, 1877), and cymes (Hutchin- son, 1964). The ultimate determinate floral unit consists of from two to eight, but usually three, flowers on subequal pedicels attached to a com- mon axis at the same point. As all the flowers in the unit are of the same sex and size, and fur- thermore all open at about the same time, it does not represent a cyme or pleiochasium in the usual sense (e.g., Lawrence, 1951), but is truly umbel- loid. Groups of two to four umbelloid units may be further aggregated or dispersed along a com- mon axis in a thyrse-like manner. The resulting units are borne decussately on a central axis in various ways; up to four different inflorescence types may occur on a given individual, all ram- iflorous. These appear to represent a reduction series from an indeterminate leafy shoot with simple axillary thyrses to a leafless, determinate thyrse (Fig. 10A-D). A) The discrete umbelloid or thyrsic floral units are axillary and occasionally subfoliar on leafless nodes near the basal portion ofthe actively grow- ing, indeterminate leafy shoot. Each subfoliar unit is subtended by a single caducous, brown bract presumably representing a reduced leaf. B) The next stage involves the addition of a terminal floral unit. Addition of further lateral units with a concomitant reduction in the num- ber of leaves, which are replaced by bracts, re- sults in the entire structure becoming a deter- minate leafy thyrse. C) Further suppression of leaves and short- ening of the floral axis results in a more con- tracted, but still leafy, thyrse. D) Finally, suppression of all the leaves and heir replacement by bracts and still further shortening of the floral axis results in a true thyrse. A similar reduction series occurs in Tambouris- sa, ultimately resulting in a monochasium. In Monimia ovalifolia, the ultimate floral units are generally quite contracted, although the sec- ondary axes may be elongated, with up to 87 flowers per inflorescence. The inflorescence of M. amplexicaulis is invariably short and contracted, consisting of not more than 37 flowers in spec- imens I have examined. Inflorescence structure is much more variable in M. rotundifolia, rang- ing from short and contracted to lax and elon- gated with up to 76 flowers. Plants of the latter PS Such variation may occur with ulation, e.g., at Col de Bellevue, Réunion (Lor- ence 2471-2474; 2773-2775, all MO). In view of the variability of inflorescence structure in Monimia, it is nota particularly useful taxonom- ic character. The basic inflorescence unit in the three species of Decarydendron (all monoecious) is a sexually mixed or unisexual pleiochasium (termed “‘spi- ciform” by Danguy, 1928). It consists of an elon- gated floral axis bearing two to four decussate or subalternate pairs of pedicellate androecious [VoL. 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN “HOS "Xosiun “pəxrtu “seyorp *umrseqooro[d — "uio VPXL “Tue əmu 911soddo snoJqe[2 pew r naundand `I "Xosiun umniseuooro[d "IUIEI 91nuo 911soddo snoJqv[3 pew r S1H2DA8 `I xəsrun uniseuooro[d “osiÁu1 "Iure1 “INE 91rjuo 911soddo "qe[aqns uo] £ SISUANJAYOU `I (pexrur) "Xosiun uniseuooro[d "I[ne5 əinuə 911soddo snoJqe[3 pew € nuo4ndpo ` | `xəsrun "os *umnriseuooro[d ‘PIXE *rurei Iu 911soddo snoJqev[8 fuv gz Dxoppbavd `I (¿) `xəsrun umrsguoorəd Š əmuə 91rsoddo 9jnsJr ‘fuy gz 1413 L `xəsrun umtirsguoorm[d "IUIEJ əmuə 911soddo qe[aqns PEN VC SISU214D2SD3DpUu `I, (pexrui) ‘MOS TIixe `xəsrun *umrseuooro[d ‘asAry} “Tel “ined oua 91soddo ‘osoqnd A[ouy A vc njy(doidaj `) ( xos "H[os ‘xe -tun) pəxttu *unriseuooro[d *2rosej *TUurel1 “Tynes əinmuə 911soddo "qep[aqns uny Vv poudij `I `xəsiun "nos "uio 9Jnuo 911soddo "qep[aqns "unay vc DSSU42 ` | aJou poxrui urmniseuoorojd ‘Tynes *rureJ 9Jrjuo 911soddo "qe[Sqns -0) pO WZ $1$42401402 `I "xosiun uniseuooro[d ‘TWEJ əmuə 911soddo snoJqe[3 PEN VC sisuapp(uvaq `] "Xosiun umniseuooro[d ‘TWEJ ərnuə ə1tsoddo `qe[8qns PEA OI ptjoñaupd ` | `xəsrun yos *umrseuooro[d 'JIxe “rues əmmuə ə1rsoddo `qe[8qns PEN OI D1DJS02140]f `] `xəsrun umiseuooro[d “Tynes əmuə əsoddo əsojıd PEA gi sisuaqdsou `I poxrui yos *umrseuooro[d '[Ixe 91rnuoa 911soddo oso[mid PPA qI ppu `I `xəsrun umniseuooro[d ‘or1osey “Tynes 91nuo 911isoddo 9jnsJm PPW GI mn4aquany `] poxrui ‘jos *umiseuooro[d [xe “urpi Əəinuə 911soddo SNO][IA PPA gi npuvaqopjn `L `xəsrun umniseuooro[d “Tne əimuə 911soddo 9jnsurmq-oso[rd PEN gi pupá4uapoəəp `I `xəsrun uniseuooro[d “TUPI əmmuə ə1tsoddo `uqn[əA PEN VI pijofiopndpn `I pəxriur unriseuooro[d ‘IXE *rurea pəu1loo1 9eysoddo əso}uəwo}-əsojıd PEN VI Djyjdydoyrta} `I poxrui ‘uos *umniseuooro[d ‘IXE “Tues P2u1ooi 911soddo `unn|əA PPW VI 1llOuƏ4nO1/] ` |). usoduioo ƏədÁ I 'IUuI `usod `I1UuI ‘ZIEN jer] SIxplol[|Aud `9səqnd 181989 A 'nsıq dnoip səroəds '`1Uul "dno 0} 3urp1oooe soroods psstunoquy I jo SIM} [eor8gol[oudioui sso18 pA `ç 318v LORENCE— MONIMIACEAE 1985] "Xosiun "oIosej *11[os ( ne) *rure1 əmuə NLU snoJqe[3 “Mey 6 DiUjad `I (pəxru) "Xosrun '0IOSej "'11[oS "Teo əmuə Lu oSOjUO2UIOj-OSO[Id ‘new 6 Sn `L "Xosrun "omosej *umirseuooroa[d "IUIEI oua NLU snoJqe[3 “pew 6 nunjdjap-1418v2 `I (`xəs -tum) poxrui unrseuoS:orəd “Tynes poy}00} aytsoddo “UIINIOA ‘PEN 8 vd4r218u0] `I ("xas -Iun) poxrui umniseuoorojd ‘Ineo pəy100} asoddo sno|l[IA "PEN 8 DdADI1IDID `I (pexrur) `xəsrun umniseuooro[d ‘astAy} ( rure1) ‘Tynes əmuə 911soddo snoJqe[3 "meN L ppifiuppnb `I pəxruu *xosiun umniseuoorojd "Tynes əinuə ə1tsoddo `qe[8qns "PEN L 1421442d `I `xəsiun winiseysorsyd “əsiÁu1 neo 91ruo ousoddo yuswo- Unna "meN 9 14Əqəts ` |. "xosiun ƏsiÁu) *umniseuooro[d neo ərnuə ə1tsoddo 9jnsJtq “pew 9 13231410Q `] `xəsiun tuiniseuoScormd nes əinuə a1tsoddo snoJqe[3 "une ç puo8n4121 `I (sjson ‘xe -19Aopnəsd poxrui “xəsıun ‘MIOS “SLY *seuorp “TWEJ CIMED o1nuo ut) 3j31soddo "qe[aqns “mney ç np `L (poxrur) "Xosiun 'Seqorp *umrseuooro[d "I[n£5 “rugi əmuə 9eysoddo "qeraqus "Ine ç pn]jJə521pəd ` |. "Xosiun “OTOSBy “IOS Tynes “wed əimuə 9yrsoddo snoJqe[3 meN ç Duofip4o2 `). `xəsiun “MOS ““OTOSRy “meo onus 911soddo snoJqe[3 ‘MEN ç $18UI]JOIOD `I "HOS 'xəsrun 'poxrui *orosej *umrseuooro[d ( rure1) "ineo o1nuo 91isoddo "qej[aqns "IneJA ç ptjoñi)du `I IOS "Xostun 'poxrui *seuorp tumrseuqooro[d ‘WII *[rxe *rurei əimuə 9yisoddo snoJqe[3 "PENA r DSO181]Ə4 `I "usoduio;) ƏədÁ I '1gu `usod `1!Jul “Zep Jer] sixejo[IAud `9səqnd `181989 A 'nsıq dnog sar»odg "1gu "penupguo) `ç digdv[ 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN _ 9.0.9. QUO | A LB FIGURE 11. and elaboration (C, G, H) series. — BS chasium. — G. Pleiochasium.—H. T decussately arranged. Not to a flowers terminated by a single gynoecious or an- droecious flower. In all three species the floral axis and pedicels are velutinous or puberulent and the pedicels are generally minutely bracteo- late (Fig. 14E). The inflorescence is usually ba- sally cauliflorous in D. perrieri, and cauli- or ramiflorous in D. lamii and probably also in D. helenae. In Ephippiandra (all monoecious), the in- florescence is a sparsely to densely pubescent dichasium or fascicle, or the flowers may solitary. In E. capuronii, E. domatiata, E. mad- agascariensis, and E. tsaratanensis it is an axil- lary three- or five-flowered sexually mixed di- chasium produced on the new growth (Fig. 14F, G). The terminal, central gynoecious flower ap- pears to mature first in the dichasium, after which the lateral androecious flowers come into anthe- sis. In E. myrtoidea, the androecious flowers are axillary, either solitary or in three-flowered fas- cicles or dichasia, whereas the gynoecious flower is terminal, either solitary or in a three- or five- iain dichasium with lateral androecious wers. A similar pattern occurs in micro- phyla although the flowers are d with the id the gynoe- cious flowers terminal on the pim In all the species, the short to long floral pedicels are sub- ° [VoL. 72 " PM i Schematic diagrams of gig as erence types showing i gee deaur (A-F) Leafy pleiochasia.—C. Pleiochasium.— l yrse. a are ded as being i in one plane, whereas nc are actually Mono- tended by a pair of acute bracteoles and the pe- duncles may also be bracteolate. Tambourissa displays a diverse array of inflo- rescence types, ranging from elaborate multi- flowered thyrses on the one hand to extremely reduced, single-flowered monochasia on the oth- er (Table 5). Pubescence of the floral axes and pedicels varies greatly, from totally glabrous to densely velutinous-tomentose. In all species the pedicel is subtended by one or rarely several mi- nute bracteoles, and the floral axis also bears several p les basally. Ped- icel length i is relatively constant for a given species. A short, determinate pleiochasium is the most widespread inflorescence type in the genus, oc- curring occasionally or consistently in 32 of the 43 species. This inflorescence type | consists of a number along a short to elongated. floral axis ; (Fig. 1 1C). Although pleiochasia of this type are unisexual in Py usss (e.g., T. tetragona), in others the flowers may be randomly mixed sexually (e.g., T. med els. and in yet others they are fre- quently terminated by a single gynoecious flower with lateral androecious flowers (e.g., T. hilde- brandtii, T. longicarpa). In this case the flowers of one sex (usually gynoecious) generally open first and it is therefore a cyme in the classical 1985] sense (e.g., Lawrence, 1951). The pleiochasia may be axillary, ramiflorous, or rarely cauliflorous. The pleiochasial inflorescence may be derived from a leafy stem with solitary, axillary, and ter- minal flowers by gradual reduction of the leaves into bracteoles, and by a concomitant shortening of the main axis (Fig. 11 A-C) similar to that described for Monimia. Single individuals of species such as T. elliptica (Lorence 2528, MO and T. pedicellata (Lorence 2584-2586, all MO) often show the entire range of stages in the re- duction series. As Tambourissa pedicellata is one of the least jp en species in terms of floral morphology, it may have possibly retained an unspecialized prece type as well. Gotts- berger (1974) considers lateral flowers or few- flowered inflorescences to be unspecialized in primitive angiosperms, from which other types have been derived. urther shortening and reduction of the pleio- chasium ultimately results in a dichasium of three flowers supported by a short peduncle (Fig. 11D, E), which may either be ramiflorous as in Tam- bourissa castri-delphinii, a dioecious species, or terminal and axillary as in 7. purpurea and T. religiosa, which have sexually mixed dichasia or pleiochasia. Suppression of both lateral flowers thus results in a single-flowered monochasium (Fig. 11F), as in T. ficus or T. tau, which are cauli- and ramiflorous respectively, and in T. crassa, which has terminal flowers. Many of the cauliflorous species are subdioecious or dioe- cious and produce clusters or fascicles of mono- chasia from knobby meristematic swellings on the trunk. That the pedicel is jointed to a distinct peduncle, where it often bears one of more pairs of decussate bracteoles, suggests that it is a re- duced inflorescence and not merely a solitary flower per se. Ontogenetic studies are required to confirm this, however. At the opposite extreme, a number of species have compound inflorescences with numerous flowers that may be derived by an elaboration of the pleiochasium. In species such as Tam- bourissa quadrifida (Group 7), there has been a multiplication of the floral pairs into cymose or umbelloid units of three to five flowers (Figs. 11G, 20B) The two species in Group 8 (T. alaticarpa and T. longicarpa) also possess pleiochasia with the androecious flowers produced in lateral umbel- loid or cymose groups of one to three, whereas one to three gynoecious flowers are segregated in a terminal unit (Fig. 17A). The terms cymose — LORENCE-— MONIMIACEAE 33 and umbelloid are used loosely here, as the flow- ers of a group may either mature simultaneously or in a staggered, random fashion. Pleiochasia witht flowers also occur in 7. perrieri (Group 7) and T. hildebrandtii (Group 1B). Floral units in other species such as T. sieberi and T. bathiei (Group 6) are produced on distinct peduncles and the resulting inflorescence is thyr- sic (Fig. 11H). In these species, the inflorescences tend to be cauliflorous and predominantly uni- sexual, although flowers of the opposite sex are occasionally intermixed. n summary, although inflorescence type and position may vary considerably in the four gen- era discussed above, they are frequently modally distinct for given species and often provide a number of taxonomically important features. FLORAL MORPHOLOGY AND ANATOMY In terms of floral morphology, the Monimi- aceae are an extremely diverse, and at the same time, coherent family in which many of the gen- era exhibit trends towards floral specialization, often via reduction (Perkins & Gilg, 1901; Per- kins, 1911, 1925; Money et al., 1950). The fol- lowing general evolutionary trends in the Mon- imiaceae sensu lato have been summarized by Corner (1976) and Endress (1980b): 1) Transition from floral bisexuality to unisex- uality. 2) Reduction in size and number of parts. 3) Transition from spiral, to radial, to decussate arrangement of parts. 4) Massive development of the receptacle (floral cup) which assumes the functions of a peri- anth and ovary wall. Closure of the receptacle, involving: ) Enclosure of the inner tepals by the re- ceptacle. Enclosure of the carpels (ovaries) by the female receptacle. C) Permanent closure of the gynoecious re- ceptacle, which dehisces only in fruit. Enclosure of the stamens by the androe- cious receptacle, which splits open flat by lateral fissures at anthesis, or remains CA — B — g partly closed. 6) Transition from free, stalked, or sessile car- pels borne on the flattened or shallow recep- tacle, to inferior syncarpous carpels im- mersed in the receptacle wall. 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN 7) Production ofa mucilaginous compitum, and nally a *hyperstigma" which assumes a sty- lar function. 8) Apical (or rarely basal) fusion of the anther loculi into a single loculus with a progressive shortening and broadening of the filament. Although many of these trends are most ap- parent at the subfamilial or tribal levels (see dis- cussion under Taxonomic History of the Mo- nimiaceae), a significant number also occur within the Mollinedioideae (sensu Thorne) and occa- sionally within a single genus (e.g., Tambouris- sa). The four genera belonging to the two subfam- ilies discussed here are all relatively specialized, i.e., have unisexual flowers lacking an elaborate petaloid perianth. Although Monimia displays a number of the same features as the other three genera, it will be discussed first as it belongs to a separate subfamily, the Monimioideae. Be- cause the other three genera, all Mollinedioideae, display obvious trends involving three levels of rm a natural coherent series, they will be discussed as such. Floral construction in Monimia is reduced, highly specialized, and very similar in all three species. Flowers of both sexes have closed, glob- ular receptacles in bud surmounted by three to six minute, decussate, scale-like tepals (Perkins & Gilg, 1901; Money et al., 1950). In Monimia, the receptacle has assumed the function of the perianth and in the gynoecious flowers turns or- ange or on at anthesis. The globose, internally pubesce free, RE carpels, each with a single pen- ulous, anatropous ovule. At anthesis, the re- ceptacle opens by means of a small, three to six lobed apical pore through which the glistening, slightly papillose white styles are exserted (Figs. 12.1A, 14B At anthesis, the androecious flower splits open by lateral fissures into four to six valvate, ulti- mately recurved lobes with numerous (ca. 25- 150) irregularly disposed, white bisporangiate stamens. Each stamen bears a pair of white, ear- like appendages at or near its base (Fig. 12.1C, D). Anther dehiscence is lateral along the inner, adaxial margins of the monosporangiate loculi, which then open outward like flaps (Figs. 12.1D, 13A) releasing the slightly coalescent pollen. Staminal dehiscence in Monimia has been de- scribed as subintrorse by Hutchinson (1964), but is more nearly introrse (De Candolle, 1868; Per- kins & Gilg, 1901). Observations on living flow- ers of all three species revealed no apparent nec- [Vor. 72 tar production by the stamens as in Atherospermoideae (Sampson, 19692), lea sweet odors are emitted by flowers of both sexes. Obvious floral morphological differences in the species are expressed in terms of receptacle size (smallest in Monimia ovalifolia), number of sta- mens (fewest in M. ovalifolia, most numerous in M. amplexicaulis), and in stamen length (short- est in M. ovalifolia, longest in M. rotundifolia). Flowers of both sexes are externally pubescent, and trichomes furnish additional useful specific characters. A more detailed study of the ontog- eny and floral construction in Monimia is forth- coming (Endress & Lorence, in prep.). The three genera belonging to the Molline- dioideae form a natural series in which three evolutionary levels of floral morphological spe- cialization are obvious, primarily in the gynoe- cious waits The least specialized genus is De- carydendron, and the most specialized is Tam- bourissa (Fi 12.2-12.4). 1) Decarydendron: receptacles of both sexes are shallowly cupuliform and open gradually without splitting at anthesis; tepals are large, nu- merous, in radial series; carpels are free, numer- ous, long and clavate, subsessile; fruiting carpels are presumably exposed (?); stamens have a dis- tinct filament and separate loculi (Fig. 12.2A- 2) Ephippiandra (including Hedycaryopsis): gynoecious receptacles are discoid and develop gradually without splitting at anthesis; tepals are few, small, decussate; carpels are free, fewer and shorter, columnar, sessile; fruiting carpels are ba- sally immersed in the receptacle; androecious re- ceptacle is closed in bud, splitting open by lon- gitudinal fissures at anthesis; stamens are subsessile, with short, broad filaments, loculi are free or confluent (Fig. 12.3A-D). 3) Tambourissa (including Phanerogonocar- us): gynoecious receptacle is closed, globular to P cylindrical, the apex often splitting partly open at anthesis, tepals are few, minute, decussate; carpels are inferior, syncarpous; fruiting carpels are enclosed; fruiting receptacle splits open at maturity; androecious receptacle is closed in bud, splitting open by longitudinal fissures at anthesis; stamens have distinct filaments or are short and sessile, loculi are free or confluent (Fig. 12.4A- D Both androecious and gynoecious flowers of Decarydendron are relatively monomorphic, somewhat subglobose in bud, and have seven to 15 large, thick, obtuse tepals in two or three 1985] LORENCE— MONIMIACEAE 35 A B C D FIGURE 12. Schematic drawings of flowers of the four genera of Monimiaceae in the Malagasy region. 1. Monimia; 2. Decarydendron; 3. Ephippiandra; 4. Tambourissa. — A. Gynoecious flower, longitudinal section. — B. Individual carpel, longitudinal section. — C. Androecious flower at anthesis. — D. Stamen. Not to scale. [VoL. 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1985] LORENCE— whorled series. The tepals are infolded in bud and the flowers consequently do not split, but open gradually at anthesis as the tepals become erect or everted. The external receptacle surface dendron is hemispherical or obconical with thick walls and straight or slightly incurved tepals. Nu- merous free, clavate carpels line the inner recep- tacle surface (ca. 300 in D. helenae) and are in- terspersed with short, densely tufted simple lignified trichomes. In D. helenae var. steno- phyllum (Humbert 6613, K), each carpel is cla- vate or cylindrical, ca. 1.5 mm long, slightly con- stricted basally and obtuse apically (Fig. 13B). A single, pendulous anatropous bitegmic ovule is contained in a small loculus which is confined to the base of the carpel and from which the oblique stylar canal runs upward and opens at the side of the carpel in its basal third (Fig. 13B). The carpellary ground tissue contains scattered oil cells above the ovary and densely aggregated stone cells in the terminal portion (Fig. 13B). Stone cells also occur in the receptacular ground tissue, but no tanniferous idioblasts were ob- served. In cleared carpels, a single vascular bun- dle enters from the base, supplies the ovule, and extends upward almost to the apex, terminating in tracheoids. The exterior surface of the carpel is possibly secretory in nature as in many Mo- nimiaceae (Endress, 1980b), but no traces of mu- cilage were observe In the züdecscian flower, from 16 to 60 sta- mens are irregularly dispersed over the glabrous to pubescent inner surface of the shallowly con- cave or cupuliform receptacle (Figs. 12.2C, 1 4E). Stamens of all three species are ieee unspecialized with a distinct, narrow filam unprolonged connective, and separate, eee dehiscent, tetrasporangiate anthers (Fig. 12.2D). Useful taxonomic features of Decr. flowers include: number and size of tepals and characters of their margins (crenulate or not); number of stamens; size of the receptacle and presence or absence of lateral ridges (Cavaco, = MONIMIACEAE 37 1959). The genus is clearly the least specialized florally of the Mollinedioideae in the Malagasy region Floral construction in adele iene nd ing Hedyca ding a strong mate male floral DNE b: ig 12.3A-D, 14F, G). Gynoecious flowers are somewhat globular and concave in bud, but open gradually and become discoid during ontogenetic development. In Ephippiandra madagascarien- sis (Bernardi 11977, K, MO), the receptacular rim initially bears four small, decussate, deltoid tepals which apparently divide once by lateral doubling, resulting in eight tepals in the mature flower. Most species also appear to have four, eight, or Far 16 minute, decussate tepals, and pubescent with short to long simple hairs. At anthesis the receptacle is flat and discoid, crowded with 20-150 free, A sessile, short tly columnar with clusters of simple, lignified trichomes (Fig. 12.3A, B). Each carpel is four to six sided, ca. 1-1.5 mm long, with a flat top, straight to inclined sides, and a slightly constricted base (Fig. 13C, D). The solitary ovule is anatropous or amphitropous and bitegmic, situated in a small loculus near the base ofthe carpel, with a stylar canal leading obliquely upward and opening at the side near the middle of the carpel (Fig. 13C, D). The entire carpel is covered by an epidermal layer which appears to be secretory and produce mucilage at least around a central zone. Occasional short, simple hairs occur on the carpellary surface in E. madagas- cariensis. In cleared carpels, two vascular bun- dles enter from the base on either side of the ovule in E. myrtoidea, E. madagascariensis, and E. tsaratanensis, and fan out into a zone with merous, terminal tracheoids near the apex. iid (1980a) also found tracheoids in other genera and suggested that they might function as water storage cells. The ventral bundle gives rise to a weak, arcuate secondary branch supplying the ovule. The floral ground tissue of E. myr- toidea contains oil cells, stone cells, and large numbers of tanniferous idioblasts. That of E. — FIGURE 13. Photomicrographs of a parts.—A. Stam dehiscence and basal glands, whole mount, Lorence stenophyllum, longitudinal section, eee rt 6613 (K).— rpel of E. myrtoidea, longitudinal section, dai tudinal section, Bernardi 11977 (M O). —D. Ca fT. . Annimigq en of A ntrorse 2444 (MO).—B. Carpels of ptr pides ae var. —C. Carpel of Ephippiandra madagascariensis, longi- n. (K).— = Rm leading Lorence 2630 (MO). Bars equal 0.5 mm Carpel o to ovule; receptacular ground tissue; s — style; t — trichome. f = flap of anther loculus; g= glands at base of stamen; m = mucilaginous usa at o = ovule; 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 ap pM, E E | ° — -* III 1985] LORENCE— MONIMIACEAE 39 iarafanensis contains both oil cells and numer- ous ta e ells, whereas only oil cells occur in E. madagascariensis. Receptacular ground tis- sue of these three species contains idioblasts comparable to those found in the carpels. The fewer and shorter, sessile carpels and reduced, decussate tepals of Ephippiandra represent spe- cializations in comparison with Decarydendron. Androecious flowers of Ephippiandra are glob- ular and closed in bud (Fig. 14F, G), externally pubescent with short to long simple hairs, and generally bear four pairs of small to minute, acute to deltoid, decussate tepals at or near the apex (Fig. 12.3C, D). At anthesis, the floral cup splits by lateral fissures into four subequal, valvate seg- ments exposing the 9-50 stamens (Fig. 12.3C, D) scattered or situated in several series on the glabrous to pubescent inner receptacle surface. The apex of each lobe usually bears one or rarely two obtuse, scale-like tepals in all but E. myr- toidea. The tetrasporangiate stamens range from ligulate and incurved to short, broad, and ob- tusely deltoid. Those of E. domatiata, E. mad- agascariensis, E. perrieri, and E. tsaratanensis are somewhat longer with more distinct fila- ments, separate loculi, and slightly prolonged or apiculate connectives, whereas those of E. mi- crophylla and E. myrtoidea are generally shorter and sessile or subsessile with the loculi apically connivent or confluent into a single, crescenti- form loculus. Staminal types within the genus intergrade, however, and a comparable range of variation occurs in Tambourissa. Dehiscence is lateral and longitudinal, but is continuous in sta- mens with a single, crescentiform loculus (both separate and confluent loculi may occur in a sin- gle flower, e.g., E. myrtoidea). Ephippiandra mi- crophylla and E. myrtoidea have few (10-18), highly reduced stamens with connivent or con- fluent loculi; these represent specialized features. Taxonomically useful characters in Ephip piandra flowers include: size of the receptacle; number of carpels, stamens, or tepals; presence or absence of tepals on the inner receptacle lobes; size and shape of tepals; size and shape of sta- mens; condition of the loculi (free or confluent). Floral reduction in the genus shows specializa- tion over Decarydendron. Floral construction in Tambourissa (including Phanerogonocarpus) displays a number of still more specialized trends, one being a strong an- droecious/gynoecious floral dimorphism (Fig. 12.4A-D), except in T. pedicellata, T. purpurea, anda few other species. Perhaps the most striking and important trend is apparent in the gynoe- cious flowers, which have their carpels immersed in and fused with the receptacular wall and are therefore inferior an rpous is (Endress, 1979, 1980b), t over Decarydendron and Ephippiandra. Al- though androecious flowers of some Tambour- issa species are comparable to those of Ephip- iandra, those of other species are even more specialized. Because a more detailed study of the ontogeny and floral construction of Tambourissa has been published elsewhere (Endress & Lor- ence, 1983), only a brief account is given here. Gynoecious flowers of Tambourissa consist of a massively developed receptacle which is gen- erally closed in bud. Considerable variation oc- curs in receptacle size and morphology, the main types of which are shown in Figure 15. The re- ceptacle is more or less napiform or cup-like in the majority of species (e.g., T. cordifolia, T. fi- cus, T. pedicellata; Fig. 15B, C, J), but ranges from flat and discoid in T. peltata (Fig. 15A), to closed and spherical in T. sieberi, or even cylin- drical in T. alaticarpa and T. longicarpa (Fig. 1 5E, H). Both the flat, open and the closed, glo- bose or cylindrical types appear to be highly spe- cialized and derived from a more generalized cupuliform type like that found in T. pedicellata. The receptacles of T. alaticarpa and T. longi- carpa, although extremely elongated and later- ally winged, still fall within the range of variation displayed by certain other species. For example, RE 14. (MO).—B. M. ovalifolia, gynoecious inflorescence M. niis ripe fruiting receptacles splitting to expose carpels, Coode 4954 (K).— ( Flowers and fruits. — A. Monimia d androecious inflorescence at an A anthesis, note exserted styles (s), Lorence 2757 (MO).—C. thesis, Lorence 2896 —D. As for C Lorence 2350 MO).—E. Decarydendron lamii, showing stamens and large tepals, Lam T Meeuse 5835 P.— -F Ephippiandra madagascariensis, inflorescences , Ursch 66 (P). —G. E. perrieri, flo owering y stem showing glo bose androecious ind in bud ind discoid central ] gynoecious flower, Perrier de la Báthie 16249 (P).—H. E. tsaratanensis, old with three green carpels seated in n cupules (right), Gentry 11 605 (MO). Bars equal 10 mm. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 10 mm S 2 > [2 o o S = ° ° c e S o e S o P 9 J k o 3 S s o 3S ° o o 3 o ° Z > S = - 2 Ó 9 o Š o Ó Q o s 7000 TN 2 S 00000 00 A S 2, 7009 o oo 00 o? * g E FIGURE 15. Schematic longitudinal sections throngh pga floral a of some Tambourissa species.—A. T. peltata.—B. T. cordifolia.—C. T. eri.—F. T. orensis. — G. T. elliptica. —H. T. alaticarpa. —I. T. quadrifida. —J. T. eu es ‘Stippled areas represent mucilaginous com- pitum. Drawn to approximate scale. 1985] LORENCE-— MONIMIACEAE 41 gynoecious receptacles of 7. quadrifida are cy- lindrical-ellipsoid, and those of T. floricostata bear four lateral, longitudinal ridges. The exter- nal receptacular surface in the genus ranges from glabrous (e.g., T. tetragona) to ee velutinous (e.g., T. sieberi) or corky (e.g., T. tau). The gynoecious bud apex in Tambourissa gen- erally bears several pairs of minute, decussate obtuse or deltoid, scale-like tepals. Two pairs are usually visible externally in most species, and these are sometimes fused into an apicule (e.g., in T. sieberi, T. tau). In T. purpurea there are usually five pairs; the innermost are enclosed in the floral cup and intergrade into the carpels (En- dress, 1980b; as T. religiosa). In several of the Comorean species in Group 2 (T. comorensis, Figs. 15F, 17B; T. kirkii; and T. leptophylla) where the gynoecious receptacle remains open during the entire floral development, the tepals are apparent only during the very young stages. This is an unusual feature within the genus, and could represent either a highly advanced char- acter or, alternatively, a transitional stage be- tween the open, discoid receptacles of Ephip- piandra and the closed buds of most Tambourissa species. At floral anthesis, the receptacles of most species generally split open by shallow (e.g., T. purpurea) to deep (e.g., 7. peltata) longitudinal fissures between the tepals, forming a number of subequal lobes ranging from four in most species (e.g., T. cordifolia, Fig. 42G, H) to as many as eight or ten in T. ficus (Fig. 15C) and T. tau (Fig. 39E). The sterile internal surface of the lobes may be pubescent (e.g., 7. moheliensis, bbe 36E) or glabrous (7. tau). Inner lobes of 7. peltata, and to a lesser degree 7. cordifolia, are sculptured with longitudinal ridges or lines, somewhat ver- rucose in the former species and bearing stomata in both species (Endress & Lorence, 1983). Po- sition of the lobes at anthesis ranges from in- curved (e.g., 7. pedicellata, Fig. 15J), or erect (e.g., T. hildebrandtii), to spreading-reflexed (e.g., T. peltata; Figs. 15A, 17E). The colorful, everted lobes of T. peltata have assumed the function of the perianth. The carpels of Tambourissa are immersed in and united with the receptacle wall and are there- fore inferior and syncarpous in a morphological sense (Endress, 1979, 1980b, 1982), not embed- ded in sockets as stated by Baillon (1871) and Corner (1976). Each carpel is unilocular with a solitary, bitegmic, anatropous ovule (amphitro- pous in 7. decaryana, T. nosybensis). The two species formerly included in Phanerogonocarpus (T. alaticarpa and T. longicarpa) also possess this type of carpel (Fig. 16A). The inner recep- tacle surface may be glabrous (e.g., T. tau; Fig. 16C) or interspersed with dense tufts of short, simple lignified trichomes (e.g., 7. sieberi; Fig. l The number of carpels varies greatly between species, is proportional to the size of the recep- tacle, and provides a good, species-specific char- acter (Table 6). For example, fresh flowers of T. purpurea are only 5-6 mm diam. and have 35- 70 carpels, whereas those of T. ficus are 35—40 mm diam. and have ca. 1,000-2,000 carpels. Size and shape of the styles are constant and often species-specific features, varying greatly within the genus. Styles of T. peltata (Fig. 16B) are short (ca. 0.5 mm long), crowded, and obtusely colum- nar. Those of T. tau (Fig. 16C) are longer (ca. 2- 3 mm), free and setose, whereas in T. sieberi (Fig. 16D) they are 3-4 mm long, setose and coales- cent. A canal was found to run obliquely upward from the ovule before opening at the base of the style in all 31 species examined in this study. This canal has been termed the “ventral slit" by Endress (1980b) and allows passage of the pollen tube for fertilization. Pollen tube transmission in Tambourissa does not occur through the stylar tissue; rather, the site of transmission has shifted to the exterior of the style as in many other genera in the Monimia- ceae (Endress, 1979, 1980b). In many species the entire external stylar surface is secretory, pro- ducing a mucilaginous exudate through which the pollen tubes grows until encountering the sty- lar canal and passing through it directly to the ovule. Traces of this mucilage can be seen in Figures 13E and 16C. A further specialization is a massive mucilage production forming a thick layer lining the floral e.g., T. peltata; Figs. 15A, 17E). This “ex- tragynoecial compitum" functionally links and interconnects all the carpels (Endress, 1980b, 1982), enabling pollen grains deposited in one place to cross over and grow to any other carpel, thereby maximizing efficiency of the pollen. It is analogous to the usual intragynoecial compitum (transmitting tissue) in syncarpous gynoecia (Carr & Carr, 1961). The mucilage may also have a nectar function (see chapter on Floral Biology). n Tambourissa purpurea, the extra-carpel- lary, stigmati tory epid i tonly the surface of the carpels but also the entire inner orifice of the floral cup including the inner tepals. Termed a “hyperstigma” by Endress (1979, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 42 1985] LORENCE— 1980b; as T. religiosa), this unique floral con- struction is the culmination of floral specializa- tion in Monimiaceae. In addition to 7. purpurea it also occurs in 7. elliptica and T. crassa (Lo- rence, pers. observ.), as well as Wilkiea, Kibara, and Hennecartia. (Endress, 1980b). In T. pur- purea, a mucilaginous plug completely occludes the floral pore and forms a continuous layer ex- tending down to the carpels and functions as the pollen reception and transmission site. In crassa and T. elliptica, the orifice is only osiüd- ed early in anthesis, but the mucilage otherwise completely lines the floral cup. À conspicuous mucilaginous exudate was also observed in fresh or rehydrated flowers of 7. decaryana, T. co- morensis, T. longicarpa, T. madagascariensis, T. nosybensis, T. peltata, and T. beanjadensis (Ta- ble 7). Each Tambourissa carpel is supplied by a sin- gle vascular bundle dorsal to the ovule which runs into the style for much of its length. The carpels and receptacular ground tissue of each species characteristically contain varying pro- portions of one or more types of the following idioblasts: oil cells; tanniferous cells; stone cells. Amount and type of each provide useful, species- specific characters (Table 7). Androecious flowers of all species of Tam- bourissa are globose to subglobose and closed in bud, usually bearing apically two external pairs of small to minute, free to fused tepals, and one or more additional pairs enclosed in the recep- tacle. The presence of tepals (as “‘staminodes’’) on inner receptacle lobes of the species formerly included in Phanerogonocarpus occurs in many species of Tambourissa and does not serve to distinguish the two genera as stated by Cavaco (1957c). The external receptacle surface is com- parable to that of the gynoecious flower, ranging from gahreus to SDN pubescent or corky. Male flora is often eap to that of the gynoecious receptacle. Two of the Comorean species, 7. kir- kii and T. moheliensis (Figs. 18D, 36D, E) have androecious receptacles that are much smaller than the gynoecious ones, however. The number of stamens is likewise proportional to receptacle size, and is generally somewhat less than the number of carpels in a given species (Table 6 variation, but MONIMIACEAE 43 To cite extremes, fresh apes pim buds of T. purpurea are 5-6 mm diam. with 20—32 stamens, whereas those of T. ficus are "a 50 mm diam. and contain ca. 900-1,800 stamens, making it the largest androecious flower in the family! At anthesis, androecious flowers of most Tam- bourissa species split open by longitudinal fis- sures between the tepals into generally four or sometimes five (range three to seven) subequal, valvate lobes. The number of lobes and degree to which they split and reflex provide useful, species-specific characters. For example, lobes of T. pedicellata (Figs. 17C, 18F) are shallowly fis- sured, incurved, and scarcely open more than halfway. Those of T. amplifolia (Fig. 18B) split about halfway and open flat, whereas those of T. quadrifida (Figs. 181, 19B) split deeply to the base and recurve completely. In some, including T. purpurea (Fig. 18E) and Comorean species such as T. moheliensis (Fig. 18D), the floral cup mere- ly splits by shallow fissures and remains rela- tively closed and globose. An extreme form oc- curs in T. gracilis and T. paradoxa, which do not split at all; a small pore flanked by tepals provides the only access to the stamens. Such minute, closed flowers and very large, open flow- ers probably represent highly specialized, ex- treme forms derived from a more generalized type, perhaps similar to that of T. pedicellata. Stamens of many Tambourissa species (e.g., T. pedicellata; Fig. 18F) consist of a distinct, broad filament and two lateral, bisporangiate, longitudinally dehiscent, extrorse loculi beyon which the connective is often slightly prolonged. j 9 EM type resembles those found in Decarydendron and some species of Ephippiandra. Other species of Tambourissa display a number of extreme and highly spe- cialized trends. One trend is towards extreme staminal elongation and prolongation of the con- nective, e.g., in 7. capuronii (Fig. 18C) with sta- mens 6-8 mm long. In 7. tau (Fig. 18H), the loculi occur on extended lateral arms of the con- nective in the form “T.” The loculi of T. ficus (Fig. 18G) have shifted to an abaxial posi- tion with the filament and prolonged connective thickened. In other species such as T. amplifolia (Fig. 18B) and 7. peltata (Figs. 17D, 18A), the loculi are often apically confluent and dehiscence — FIGURE 16. AE and carpels of some Tambourissa — longitudinal sections. — —B. T. peltata, Lorence 2555 (MO).— 2953 (MO). — leading to ovule; m = mucilaginous exudate; o = ovule; r = receptacular ground tissue; s = style; t = trichom A. T. longicarpa, Ca- T. tau, Lorence 2953 —D. 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In T. purpurea (Fig. 18E), locular confluence is accompanied by an extreme stami- nal shortening and broadening, the resulting sta- men being sessile or subsessile (0.7-1.2 mm long) with a single, crescentiform loculus which de- hisces apically (Endress, 1980b; as 7. religiosa) as in some species of Ephippiandra. However, basal fusion of the loculi occurs only in T. mohe- liensis (Fig. 18D), and also in some (but not all) stamens of T. cordifolia (Fig. 42C- Anatomically, the androecious n — IS similar to that of the gynoecious one. À single vascular bundle supplies each stamen (e.g., in T. tau). The same types of idioblasts are often found in the androecious flowers as in the gynoecious ones, most commonly oil and tanniferous cells. Stone cells, however, tend to be less abundant or lacking in the androecious flowers, e.g., pud iie (Endres, 1980b; as T. religiosa). Their LORENCE-— MONIMIACEAE 47 hiscent with a lignified endocarp, an abundant, oily endosperm, and a small embryo. While much larger in fruit, the shape of the receptacle and the carpels' position largely correspond with their shape and position in flower; i.e., significant ge- neric differences exist. Also, the thick, pitted, bony white endocarp of Monimia (Monimioi- deae sensu Thorne, 1974) differs conspicuously from the thinner, finely sculptured, brownish, horny endocarps of the genera in Mollinedioi- deae (e.g., Ephippiandra and Tambourissa; un- known in Decarydendron Fruiting receptacles of all three species of Monimia are similarly subglobose to ovoid, en- closing from one to 12 free, seed-like drupaceous carpels. On herbarium labels these are often mis- taken for multiple-seeded fruits. Fruiting recep- tacles of Monimia are strikingly similar to T of Palmeria (Comer, 1576; Endress, EE BIC fruits i dii they may play a role in protecting the developing ovules from predation. Floral color is relatively constant in many which often have two color m is always constant on a given ad dal and fre- quently constant within a local population. In T. quadrifida, however, both color morphs were ob- served in a single population at Yemen Valley, Mauritius. Incidentally, floral odor was found to be quite constant for each species and provides a diagnostic field character. Floral morphology, color, and odor all appear to be related to pol- lination syndromes (see discussion under Floral Biology). In summary, floral size and morphology, in- cluding number, structure, and color of stamens in the androecious flowers and number, struc- ture, and color of the styles in the gynoecious flowers are distinctive for most species of Tam- bourissa. These features provide some of the most taxonomically useful characters in the genus and should be recorded in the field notes. As flowers lose their shape upon drying, supplementary col- lections of flowers in alcohol should also be made whenever possible, as well as a photographic rec- ord. For a more detailed study of the floral mor- phology and anatomy of Tambourissa, see En- dress and Lorence (1983). FRUITING RECEPTACLE AND CARPELS The seed-like fruiting carpels of all the genera for which information is available are funda- mentally similar in being drupaceous and inde- maturity tł the apex into three to six : irregular, valvate (oles that open to reveal the bright orange fruiting car- pels set against the pinkish red receptacular sur- face (Figs. 14C, D, 19C). Birds appear to be the effective seed vectors for M. ovalifolia in Mau- ritius (Lorence, pers. observ.), and also for the species of Monimia in Réunion (A. Rolland, pers. e ° m.). The indehiscent carpels of Monimia consist of a lignified, bony endocarp enveloped by a soft, orange, aril-like outer coating apparently com- posed of two distinct elements. The outermost is an enation extending downward from the sty- lar base over the apical portion of the carpel and forming a cap-like structure covering the lateral portions of the carpel, while leaving the dorsal and especially the ventral portion exposed (Fig. structure. The inner layer consists of the meso- carp proper which completely pu the en- docarp, but is thickest basally (Fig. 19A, B, D). It is also orange but lacks the oil cells . the former. An ccurs in the neo- tropical Siparuna (Commer, 1976; Endress, ok and has been termed a “‘carpellary aril” or **stylar aril” respectively by these authors. Further stud- ies on the development of Monimia carpels are in progress (Endress & Lorence, in prep.). he endocarp of Monimia is whitish, irregu- larly ovoid-compressed, longitudinally crested and apically mucronate with the micropyle pass- ing through the base of the beak. The wall is hard and bony, ca. 0.5-2 mm thick (probably ade- quate to survive passage through a bird’s gut), 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FicunE 17. Flowers and fruits of some Tambourissa pisa p T. alaticarpa, Esp an. showing ebrios Bynoecious flowers and lateral ied iria flowers in bud, inset shows androecious flower at anthesis and | , Perrier de n Báthie 1 01 02 (P). —B. T. comorensis, inflorescences with androecious and gynoecious flowers, and dehisced fruiting receptacle exposing ripe carpels (c), Lorence 1985] LORENCE- MONIMIACEAE 49 TABLE 7. Selected floral anatomical features of Tambourissa species according to Group. Data given for both androecious and gynoecious flowers, except where noted. + indicates presence of a small amount. p Sasam. Tanniferous Stone Cells mpitum Species Group Idioblasts (Sclerids) Oil Cells LE T. thouvenotii 1A + — T. trichophylla 1A — + + - T. pde inm (ó) lA — — + ? T. dec 1B — + + + T. hildebrandtii 1B + — + — T. humbertii 1B — — + - T. nitida 1B + — + ? T. nosybensis IB = + + + T. floricostata IC + = + ? T. parvifolia 1C = + + ? T. beanjadensis 2A = + + + T. comorensis 2A = + + + T. crassa 2A = — + + T. elliptica 2A + -= + T. leptophylla 2A = + + — T. madagascariensis 2A + + + + T. kirkii 2B ? ? ? - (9) T. paradoxa (8) 2B B + + T. capuronii 3 + + _ - T. moheliensis 3 + + + — T. gracilis (é) 4 _ = + 7 T. purpurea E + + _ + T. religiosa 4 + + — — (?) T. amplifolia 5 + + + — T. cocottensis 5 + + — — T. cordifolia 5 + + — — T. pedicellata 5 = + + — T. tau 5 + F + — T. tetragona 5 2s s= F = T. bathiei 6 + + + — T. sieberi 6 = + + — T. perrieri 7 =: + + — T. quadrifida fi + + — — T. alaticarpa 8 + + — - T. longicarpa 8 + + + + T. castri-delphinii 9 + + + — T. 9 + + + = T. peltata 9 + + = + and consists of many layers of contiguous scle- — a thin single-layered tegmen composed of elon- rids. The endocarp surface is externally deeply gated, rectangular cells encloses the abundant, pitted as in Peumus and Siparuna. Surface sculp- oily endosperm. The small embryo is situated turing ranges from foveate to reticulate or sul- near the micropyle (Fig. 19A) and has slightly cate, varying even within a single species. Inside, divaricate cotyledons. Germination of both M. — 2870 (MO).—C. T. params gynoecious (left) and PUER (right) flowers at anthesis, Lorence 2384 (MO).— D. T. peltata, andr us flower at prr and two androecious buds, Perrier Nature Reserve, Mau- ritius (R. E. Vaughan weisst photo). — E. "dentis emis opened gynoecious flower, note copious mucilaginous exudate (m) on gynoecial disc, Lorence 2555 (MO).—F. T. abies submature fruiting receptacle on branch, Lorence 2197 (MO). Bars equal 10 mm in A, C-E, and 50 mm i 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURE 18. Androecious flowers and stamens of some Tambourissa species, drawn to approximate scale.— A. T. peltata. —B. T. amplifolia.—C. T. capuronii.—D. T. moheliensis. —E. T. purpurea.—F. T. pedicellata. — G. T. ficus. —H. T. tau.—I. T. quadrifida. Bar equals ca. 10 mm for flowers and ca. 1 mm for stamens. 1985] LORENCE-— MONIMIACEAE 51 TABLE 8. Floral color morphs of individuals of some species of Tambourissa from Mauritius and Réunion (inner receptacle surface, stamens or styles Species Androecious Flower Color Gynoecious Flower Color T. amplifolia white T. cordifolia T. crassa T. elliptica subsp. elliptica white; pink us white; purple-red orange; red T. sieberi T. tau white pale yellow; dark purple-red purple-red; salmon pink pale yellow; purple-red greenish white; purple-red orange; pale yellow pink; purple-red pale yellow-green; dark purple-red pale yellow-green; pink yellow-pink lobes purple-red, disc orange greenish white; reddish pink pale yellow-green; purple-red rp d ovalifolia and M. rotundifolia seeds took over four months and viability was quite low, ca. 10- 15%. Fruiting carpels of Decarydendron are un- known (Cavaco, 1959). Because its flowering car- pels are free and subsessile on the receptacular surface, the fruiting carpels may resemble those of Ephippiandra in being free and exposed on the receptacle, but probably lack a basal cupule and are sessile or shortly stalked as in Hedycarya or Mollinedia. Mature fruiting receptacles of Ephippiandra (including Hedycaryopsis) are discoid, flat to convex, fleshy and red when fresh, and bear ca. 10—75 free, ovoid to ellipsoid carpels ca. 7-12 mm long. Each is partly surrounded basally for about one-third to one-half its length by a cupule (Fig. 14H) derived from the receptacle. The ma- ture carpels are composed of a thin, fleshy green exocarp which turns black on ripening and sur- rounds the thin, pale brown, horny endocarp. The endocarp surface is smooth and finely scro- biculate as in Tambourissa and encloses a co- pious, fleshy endosperm s in the Tambourissa (including Phanerogonocarpus) are many times larger than the flowers but otherwise similar in shape, ranging from flat and discoid (e.g., some T. peltata fruits), or somewhat open and cupuliform in the majority (e.g., Figs. 17B, F, 42J), to ellipsoid and closed with a small ori- fice in T. quadrifida, or cylindrical with four to five prominent external ridges in T. alaticarpa and T. /ongicarpa. The majority are corky brown externally, but may be smooth and dark (reddish when fresh) in T. purpurea and T. religiosa. Most have an internal cavity, but some fruits of T. hildebrandtii are almost solid. The styles persist in fruit and line the cavity, providing important other genera, fruiting receptacles of diagnostic features as to their size, shape, a ner. Other taxonpmicaliy useful features bt ep (on the trunk, branches, Or ` terminal), whether solitary or in clusters, its size, shape, and the proportional size of the orifice. At maturity the receptacle splits open irregu- larly, exposing the numerous, bright red-orange 19. Monimia fruiting receptacles and car- FIGURE l gitu tudinal section, Lorence 2350 (M am plexicaulis, ripe receptacle split open m reveal single carpel, Lorence 2503 (MO).—D. M. amplexicaulis, ripe Mg came abaxial view, Lorence 2503 (MO). Bars qual 2 m ap genes 0 mm in C. pins — endocarp; em — acy core en = endospe pa si r= doctiss s — stylar aril. 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN carpels which separate from the pale orange re- ceptacular ground tissue (Fig. 17C). In Tam- DORTESSG cordifolia (Lorence 2630, MO), the re- d tissue intermixed with sheets and columns of tannifer- ous cells and brachysclerids comprising nearly half the total tissue mass. In carpels of 7. cor- difolia the mesocarp is composed of parenchy- matous cells containing numerous orange plas- tids interspersed with abundant oil cells and has a sweet taste when fresh (D. Lorence, pers. ob- serv.). Beneath the mesocarp lies a horny, brown endocarp with a smooth, finely sculptured sur- face composed of numerous subspheroidal to cu- boidal, lignified sclerids united into longitudinal bands and overlain by a single outer layer of flat, discoid sclerids. Beneath this lies an endotesta of elongated, annular tracheids three to four cell layers thick, and finally a unicellular tegmen of cuboidal cells surrounding the abundant endo- sperm whose cells are full of oil droplets. The embryo is 3-4 mm long, situated apically near the micropyle, and has two flat, erect cotyledons. Carpels of various species differ somewhat in size and also color and sculpturing of the en- docarp. In three Mauritian species belonging to Group 5 (i.e., Tambourissa amplifolia, T. pedi- cellata, and T. tau), it has sparsely branched, longitudinal ridges. Otherwise the carpels pro- vide few taxonomically useful characters. Birds appear to be the effective seed vectors in Tam- bourissa (D. Lorence, pers. observ.; A. Rolland, pers. comm.; Benson 142, BM) In summary, morphological and anatomical features of the wood, stems and phyllotaxy, leaves, inflorescence, flowers, and fruits of the Malagasy Monimiaceae often display marked trends and provide important diagnostic char- acters at the generic level. Furthermore, features of trichomes, inflorescence, and in particular flo- ral morphology and anatomy also provide valu- able diagnostic features at the specific level. POLLEN As a detailed palynological study of the Mal- agasy Monimiaceae has been published else- where (Lorence et al., 1984), only a brief sum- mary of the results will be given here. The four genera studied here all possess inap- erturate spheroid, subspheroid, or ovoid pollen characteristic of the majority of the Monimi- aceae sensu stricto excluding the Atherosper- moideae, whose pollen is disulcate (Money et al., [Vor. 72 1950; Walker, 1976). Although Walker (1976) probably meant to exclude the spinose pollen of Monimia (Monimioideae), his statement that pollen of“. . . most of the genera on Madagascar and the Mascarenes is distinctive in possessing spiral bands on the surface" in the manner of Hortonia (Hortonioideae; Ceylon) does not ac- curately describe the pollen of the other three genera (all Mollinedioideae). Their sexines ex- hibit variable pide Ay us from spinose in Monimia, corrugate in Decarydendron, either granulate, reticulate, striato-rugulate, striato-re- ticulate, or tectate in Tambourissa (including Phanerogonocarpus), to granulate with verru- cate, gemmate, or even vesiculate processes in Ephippiandra (including Hedycaryopsis) (Lo- rence et al., 1984 It was found that pollen size, exine thickness, species examined in this study. Although paly- nological evidence does not support the segre- neric differences in pollen are otherwise distinc- tive. CHROMOSOME NUMBERS As chromosome numbers provide some de- gree of support for the segregation of at least some of the subfamilies of Monimiaceae (e.g., Ehrendorfer et al., 1968; Goldblatt, 1974), and because all of the genera in the Malagasy region were previously unknown cytologically, it was decided to accumulate as much chromosomal information as possible during this study. All the reported counts were obtained from androecious flower buds fixed in 3: 1 absolute ethanol : glacial acetic acid and later examined by the acetocar- mine squash method. Chromosomal study of all three species of Monimia has proven extremely difficult because of their small size and high number, a difficulty compounded by their tendency to stick together and by the presence of cytoplasmic inclusions. As a result, only approximate counts were ob- tained, both for Monimia rotundifolia: Réunion, pis cloud forest, Lorence 2429 (MO), n = . 44; Réunion, Col de Bellevue, cloud forest, D 2473 (MO), n = ca. 48. The gametic chromosome number for this species therefore lies somewhere between 40 and 50, and it is ob- viously paleopolyploid. 1985] These chromosomal findings for Monimia do not appear to suggest any close numerical affinity with Peumus (2n = 78; Morawetz, 1981). Both genera appear to be e however, as op- posed to the majority of dide E sensu stricto (e dus M pers. com All nine species of T: SE eats examined cy- tologically in this investigation had a gametic chromosome number of n = 19 (Table 9). Although six of the species were from Mau- ritius, a few from Grande Comore, Réunion, and Madagascar were also examined with the same results. This suggests that the extensive adaptive radiation of Tambourissa in the Malagasy region has occurred at the same chromosomal level. The paleohexaploid base number x = 19 (Goldblatt, 1980) is shared with a number of carya, Levieria, Palmeria, and Wilkiea (Ehren- dorfer et al., 1968)—as well as by Hortonia in the Hortonioideae (Goldblatt, 1974). As in Tam- bourissa, these genera are characterized b droecious flowers bearing numerous, irregularly spaced subsessile stamens with lateral dehiscence (Ehrendorfer et al., 1968), although gynoecious floral construction is more variable (Endress, 1980b). Similarities in cytology and androecious floral morphology among these genera suggest they are relatively closely allied within the Mol- linedioideae. The remaining endemic Madagascan genera belonging to the Mollinedioideae—i.e., Decary- dendron and Ephippiandra (including Hedycary- opsis) as well as both Tambourissa species for- merly included in Phanerogonocarpus — are unknown cytologically. FLORAL BIOLOGY Although the vegetative and floral anatomy (Money et al., 1950), palynology (Walker, 1976), and cytology (Ehrendorfer et al., 1968; Gold- blatt, 1974) of various extant primitive angio- sperm families have been relatively well studied, comparatively little is known of their floral bi- ology and breeding systems. I have therefore at- tempted to accumulate as much data as possible on these aspects of the Monimiaceae while in the Malagasy area. The only confirmed report on pollination in Monimiaceae sensu stricto is by Gottsberger (1977), who observed female thrips making holes and ovipositing in androecious and gynoecious LORENCE—MONIMIACEAE 53 flower buds of Mollinedia i in Brazil. The larvae Je the buds and the adults emerge as the buds open, flying out and carrying pollen to other flowers as they seek mates. En- dress (1979, 1980b) also reported finding holes and eggs in flowers of Wilkiea and Stegnanthera (Mollinedioideae), which may have been from pollinating insects. More recently, investigations by Thien (1980) on the pollination of species of primitive angiosperms belonging to the Dege- neriaceae (Degeneria) and Winteraceae (Drimys, Belliolum, and Zygogynum) in the South Pacific islands show that members of the orders Co- leoptera, Diptera, Lepidoptera, and Thysanop- tera are important pollinators of these plants. The present study was conducted in Mauritius during 1978-1979 and includes observations on seven species of Tambourissa and one species of Monimia. The species studied, study sites, and observation periods are given in Tables 10 and 11. Insect visitors were captured, killed, and mounted or stored in 7096 alcohol for identifi- cation. Flowers of all species studied were ob- served and photographed during various stages. Voucher specimens of the plants are deposited at MAU and MO Tambourissa is a genus of monoecious or dioe- cious shrubs, treelets, or trees. All species have highly specialized, unisexual flowers with mi- nute, scale-like tepals, a well-developed recep- tacle which assumes the functions of a perianth and ovary wall, and inferior carpels. As an un- derstanding of the floral morphology is essential to understand the pollination biology, this chap- ter should be read in conjunction with the section on floral morphology and anatomy. Female receptacles of most species are some- what globose and closed in bud, and split open at maturity into (3—)4-5(-10) lobes, depending on the species. Once open, they do not close again. The receptacle is lined with numerous short to long styles, each corresponding to a single car- pel. These become receptive gradually and cen- tripetally during floral anthesis, which typically lasts 10-15 days. Receptive styles of most species produce a slight mucilaginous exudate in which the pollen tubes germinate and grow (e.g., Tam- bourissa tau, T. cordifolia, Figs. 13E, 16C). In others, the styles are reduced and covered by a massive mucilaginous secretion (compitum) (e.g., T. peltata, Fig. 17E). In yet others (e.g., 7. pur- purea), the flower remains closed except for a mucilaginous plug which occludes the floral en- trance and has a stigmatic function (Endress, 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 TABLE 9. Collections of Tambourissa in which the haploid chromosome a was determined as n = 19. All collections are mine; vouchers are deposited at MO and most at MAU a Lorence Species Number Locality T. comorensis 2878 Grande Comore. La Grille, wet forest. T. cordifolia 2571 Mauritius. Pétrin Nature Reserve, Philippia heath formation. T. elliptica subsp. elliptica 2526, 2530 Réunion. Takamaka, wet forest. subsp. micrant 2483 Réunion. Mare Longue, wet forest. T. pedicellata 2384 Mauritius. Pieter Both Mt., degraded wet forest. ltat 2553 Mauritius. Cascade Aenda stunted Me forest. T. purpurea 1726 Madagasca Mes: locality unknown (cult. MO). T. quadrifida 2539 Mauritius. n a Valley (Yemen), aa dry forest. T. sieberi 2250 Mauritius. MEI ocotte Nature Reserve, cloud forest T. tau 1964 Mauritius. Piton de la Riviére Noire, wet forest T. tau 2951 Mauritius. Perrier Nature Reserve, wet forest. 1979, 1980b; as T. religiosa). This **hyperstig- ma" occurs in some Madagascan and Réunion species, but is absent from Mauritian and Com- orean plants. The sweet mucilage of 7. purpurea tested positive for reducing sugars with Bene- dict's solution and for glucose with Lilly TES- tape (paper for analysis of urine sugar), respec- tively (Endress & Lorence, 1983). It therefore probably has a nectar function in a number of species that produce an abundant secretion (e.g., T. crassa, T. elliptica, T. peltata, T. purpurea). Flowers of most Mauritian species also produced strong floral odors which presumably attract pol- linators. Androecious flowers are similarly lined with unos siis that an mature Jracuay into (3-M- 5(-7) subequal vanes ec aiente: “AL th species open by a small oe orifice, lobes of most species open flat or even recurve and remain permanently open during their 7-12 day life span. Androecious flowers produce large amounts of pollen, which TRPIESEDH a substantial ps re- ward, although s t.I ition, floral odors like those in the aeons flowers are emitted. In the Mauritian Tambourissa species, from one to more than 100 flowers were found to be open on a given day, depending on the species and individual, and flowering generally occurred over a three to five month period. These aspects will be discussed below in more detail by species. A synopsis of the results of this study for Tam- bourissa in Mauritius is as follows. ough a few Comorean 1. Major pollinators are insects belonging to the orders Coleoptera and Diptera. Gynoecious floral morphology, notably size of the floral entrance, plays a key role in reg- ulating insect entry. Floral odor, and possibly also color, plays an important role in attracting pollinators. 4. Floral reward in the androecious flower is pol- i a tract by deceit, while in others a mucilaginous exudate seems to have a nectar function. 5. Demography, phenology, and breeding sys- tems are related to pollinator syndromes. TAMBOURISSA SIEBERI Tambourissa sieberi, a large, monoecious can- opy tree 8-10 m tall, is known from only a few restricted areas of mature wet and cloud forest in Mauritius. It is scarce and occurs as scattered individuals in low population densities, e.g., at Macabé forest (Table 4), and at Mt. Cocotte where it was studied (Table 10). Hundreds of flowers (16-90 per inflorescence) are produced in uni- sexual pleiochasia and thyrses arising in large numbers from the trunk and major branches from December to February. Gynoecious flowers are globose with a small apical pore only 2-3 mm diam. (Fig. 20A). Sev- eral hundred long styles which are coal tinto groups fill the receptacle, leaving only a small central cavity (Fig. 15E). The styles are moist at anthesis due to a slight surface exudate. An- droecious flowers are relatively large and split into five spreading, reflexed lobes bearing ca. 100— 1985] LORENCE— MONIMIACEAE 55 TABLE 10. Localities for pollination biology studies in Mauritius. Ele- Annual vation Pptn. No. Locality Vegetation Type (m (mm) Date Season 1. Mt. Cocotte cloud forest 760 5,000 Jan. 1979 wet 2. Matala valley (Yemen) semideciduous dry forest 200 1,600 June-July 1979 dry 3. Perrier Sideroxylon 600 3,400 Sept.-Oct. 1978 dry lower montane wet forest Sept. 1979 dry Apr. 1979 end wet 4. Pétrin Philippia heath formation 630 4,000 May 1979 start dry 5. Bel Ombre Lowland wet (high) forest 300 2,400 Dec. 1978-Jan. 1979 wet 6. Mare Longue Plateau Lower montane wet forest 660 3,400 Apr. 1979 end wet 7. Piton Brise Fer Lower montane wet forest 700 2,400 Dec. 1978 et 140 large stamens that release abundant, coales- cent pollen. Flowers may be either pale yellow or dark reddish purple, depending on the indi- vidual. Flowers of both sexes produce the same strong, cloying odor of overripe or fermenting fruit. Observations made near midday, totalling 3.4 hours, revealed that introduced honeybees (Apis mellifera) were the most frequent visitors of an- roecious flowers, from which they gathered pol- len. They showed no interest in the gynoecious flowers, however, nor could they enter them. Other less frequent visitors to the androecious and buctler unidentified dipteran (Table 1 1). No visitors were observed on the gynoecious flowers during this time, and no observations were made at night. Examination of the contents of ten gynoecious flowers from the tree and 15 fallen flowers re- vealed the presence of various small arthropods, notably a variety of small Coleoptera only 2— mm long, in over half the flowers (Table 12). These Coleoptera, belonging to the families Hy- drophilidae, Nitidulidae, Rhizophagidae, and Staphylinidae, are presumably active at night. They probably visit androecious flowers to feed on the pollen, attracted by the strong floral odors. They also enter the gynoecious flowers to feed on the styles, which were damaged in certain flowers, and probably oviposit there, as eggs and larvae were found in some flowers (Table 12). Because the stylar canal is situated near the style's base, damage to its terminal portion probably has little if any effect on pollen tube germination and growth which may occur anywhere on the stylar surface. Reduction of the floral entrance can be viewed as a strategy to protect the ovules o from damage by larger, more destructive beetles (cf. Gottsberger, 1974, 1977; Endress, 1980b), at the same time allowing access to the less destruc- tive small beetles. Elongation and coalescence of the styles may provide food for the coleopterans and also protect the ovules. Carpel group for- mation of this type, by connivence of the styles and cohesion by the mucilaginous secretion, functionally divides the gynoecium into several subunits, each being composed of a number of carpels (Endress & Lorence, 1983). This phe- nomenon is unknown elsewhere among the an- giosperms. Breeding experiments suggest that Tambouris- sa sieberi is the only self-compatible monoecious species of the four tested in Mauritius (Table 13). I suspect self-compatibility in 7. sieberi may be a strategy to offset its low population densities and probable inefficiency of the small pollina- tors. Production of a large number of flowers and a prolonged flowering period may also compen- sate for these factors. TAMBOURISSA QUADRIFIDA Tambourissa quadrifida is the only species of Monimiaceae to occur in the dry forest zone of Mauritius. A mid stratum or subcanopy tree, it occurs in small, scattered populations and readi- ly forms multiple coppice stems. Individuals are subdioecious, producing flowers of predomi- nantly one sex, and flower from June to August. The numerous flowers are produced in pleiocha- sia and thyrses along the trunk, and hundreds of flowers may be open on a given day, each per- sisting for ca. 10-15 days. During the 1979 flow- ering season, a predominantly androecious tree (Lorence 2656, MO) was estimated to produce nearly 24,000 flowers, and a predominantly gyn- 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 1985] oecious tree (Lorence 2655, MO) os 15,000 IC out on three individuals showed them all to be self-incom- patible (Table 13). The gynoecious flowers of Tambourissa quad- rifida are ellipsoid to obovoid and open by a small apical pore only 1-2 mm diam. (Fig. 20B). Inside, 200-300 short, glistening pink or yellow styles line the hollow receptacle; only a small amount of stylar exudate is produced. The or- ange or red androecious flowers split into four ultimately recurved segments bearing 300—500 stamens with slightly coalescent pollen (Fig. 20B). A strong, pleasant odor of ripe fruit is produced by flowers of both sexes. Studies were carried out on a population of four individuals at Yemen (Matala Valley) on the edge of a dry forest degraded by numerous exotics. Observations totalling 13.4 hours re- vealed that large numbers of small Diptera (Dro- sophila spp., Homoneura, and Zaprionus) con- gregated on the androecious flowers in the early morning, but did not enter the gynoecious flow- ers (Table 11). For this reason, the drosophilids presumably have little, if any, effect as pollina- tors. Three species of syrphids also visited the androecious flowers occasionally during the day to feed on the pollen, and pollen foraging activity by honeybees was also intense throughout the day. As these insects are all too large to enter the gynoecious flowers in which the styles are con- cealed, they cannot function as pollinators. n examination of 438 gynoecious flowers from the tree and 132 fallen flowers revealed that 93.6% and 75.8% respectively were devoid of any contents (Table 14). A number contained mealybugs which were presumably brought in by the small ants also found in some flowers. Ants are probably ineffective as pollinators, as they did not visit androecious flowers. mall per- centage of flowers contained silk webbing, often with small larvae or pupae (probably Lepidop- tera). These larvae appeared to graze on the styles and probably represent predators, although pol- linating activity by the ovipositing adults cannot LORENCE— MONIMIACEAE 57 be discounted. The only likely pollinator ob- served was a small Nitidulidae beetle, Haptoncus luteolus (Tables 11, 14) seen emerging from a gynoecious flower. It subsequently visited an an- droecious flower to forage for pollen, which ad- hered to its body. The beetle is probably able to travel the short distances of several meters sep- arating trees in this population. Significantly, another species of Haptoncus was seen in the structurally similar gynoecious flowers of Tam- bourissa sieberi (Table 12), and Thien (1980) ound the same genus in Degeneria flowers. One factor responsible for the scarcity of pol- linators on Tambourissa quadrifida at Yemen may have been the disturbed, partly secondary nature of the habitat. However, the predomi- nantly gynoecious tree of the four usually pro- duces some fruit clusters each year (J. A. Lal- ouette, pers. comm.), indicating that effective cross pollination does occur to a certain degree. As in Tambourissa sieberi, an extreme reduc- tion of the gynoecious floral entrance excludes larger, more destructive insects, permitting only smaller pollinators like the nitidulid to enter. The tremendous floral production in 7. quadrifida presumably offsets the lower efficiency of these insects as pollinators as in 7. sieberi, whereas self-incompatibility and subdioecy would en- force outcrossing. TAMBOURISSA TAU Tambourissa tau is a slender, monoecious understory treelet rarely exceeding 2- ig Flowers are produced from meristematic odi. ings along the main stem, or in the leaf axils. Depending on its size, each plant may produce over 100 solitary or fasciculate flowers over a three to five month period, but only several open at a time, each lasting for 10-15 days. Crossing experiments revealed it to be self-sterile (Table 13 Gynoecious flowers are relatively small, ex- ternally corky, and split about half way open into 6-10 irregular, suberect lobes (Fig. 39E, F). Each E20. Flowers of some Tambourissa species.—A. T. ais gynoecious flowers in bud (left) and at it anthesis (right, note insect egg (e) in orifice, Lorence 2250 (MO).— . T. quadri androecious plant with flowers in bud and at anthesis, note visiting E (d), Lorence 2632 (MO).— T. crassa, gynoecious flower at anthesis, note visiting dipteran (d) dri with partly closed white androecious flowers in bud, at anthesis and post a purple androecious flower at anthesis, note visiting syrphid (s), Lorence T. ficus, x (MO).—F. T. ficus, form n open 2115 M Bars equal 10 m nking mucilage, Lorence 2468 MC nthesis, Lorence 1485 [Vor. 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN 89 I pinspu pjnjdosoaq Apuim Əepilrtudosoiq 7? Auuns ‘WY STETIT 6L61 Bioidiq Sv 97-61 0} UMEp —W''Y Șp:9 KeJN o£ r 99 ‘ds pinauowopy oepliuexney 29 cT [e101 42801114 snuo1dvz ÁpUtA Əepirrudosoiq % Auuns '"W'v ç0:6 6L61 Bisidiq 0't 97-81 0} UMEp —W'v $0:9 ABW OE v çL I pinspu pu dosoq xsnp 0} Əepilrtudosoidq 9I Apuim "Wd çt:9 6261 "odi 8I -$'6I % Auuns —W'd 00:6 &EN6I bt 69 ‘ds pu11ə5(1/2Dq oepriruexneT 89 cT [2101 pinspu vj1ydosoiq Apuim Əepirrtudosoiq $'9c 7? 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Cocotte, Mauritius in January TABLE 12. Contents of Tambourissa sieb 979. o. No. Flowers Flowers ro From % Contents Tree Total Ground Total Empty 10 45.5 15 41.7 Small Coleoptera: 6 27.3 13 36.1 (Hydrophilidae, Dactylosternum vinsoni B.B.; Nitidulidae, Carpo- philus mutilatus Er., Epuraea terminata Rtt., Haptoncus minutus Rtt., Lasiodactylus pictus MacL.; Rhizophagidae, Europs cf. brevis Gronv.; Staphylinidae, Atheta dilutipennis Mots., Philonthus flav- ocinctus Mots. Lepidoptera (Opogona sacchari Bojer, larva, presumably a predator) l 4.5 5 13.9 Unidentified larvae (possibly Diptera) 2 9.1 0 0 Ants (Hymenoptera, Formicidae) 0 0 l 2.8 Isop 2 9.1 0 0 Unidentified insect eggs l 4.5 2 5.5 Total 22 100.0 36 100.0 a Three individuals of Epuraea terminata also found on bagged androecious flowers of T. sieberi at this site. flower contains 80-100 long, dark purple-red se- flowers split into 5-7 irregular, flat lobes and contain 100-150 T-shaped stamens with loculi held on lateral arms (Figs. 18H, 39B, C). The pollen is only slightly cohesive. Androecious flower color may be either dark purple-red or greenish white internally. I was unable to detect any floral odor from 7. tau flowers during either day or night. A population of Tambourissa tau at Perrier Nature Reserve was observed for pollinators for a total of 10.8 hours. The only insect to visit the flowers with any regularity was Scaevinus trun- catus, a cryptic, nocturnal folivorous weevil found to occur on most species of Monimia and Tam- bourissa in Mauritius and Réunion. Scaevinus normally feeds on the leaves of T. tau, but durin the flowering season weevils were found feeding on buds and flowers, grazing on the stamens, styles, and receptacle lobes. Large numbers o pollen grains were found trapped among the in- sects’ body hairs and scales. To test the weevil’s vagility, marking experiments were carried out on Scaevinus at Perrier. Among 52 individuals of T. tau studied, 8-2896 were found to harbor Scaevinus on a given day. Out of 27 Scaevinus marked at the onset of the experiment, 18 (67%) had moved to other plants in the same popula- tion after three days. As T. tau occurs at com- paratively high population densities at Perrier (57 plants/512 m2, or an estimated 1,113/ha), with nearest neighbor distances averaging only 160 cm (Fig. 3B), and fruit set in the wild is relatively high, it appears that Scaevinus is ef- fective in cross pollinating 7. tau. Although Scaevinus was also seen feeding on flower buds of 7. peltata at Perrier, it was never observed to visit its flowers, nor those of any other Tam- bourissa or Monimia species. Finally, no other insects were observed to visit the flowers of T. tau. TAMBOURISSA CORDIFOLIA A study of this species ied out at Pétrin Nature Reserve where Tambourissa cordifolia is common and forms dense, local populations in a low, scrubby Philippia heath formation. It is a small, dioecious shrub 1-2 m high with erect, multiple stems bearing medium-sized solitary or fasciculate flowers on leafless nodes. The an- droecious flower splits into four flat, spreading- reflexed lobes with 300—400 white stamens (Figs. 20C, 42B), which release white, coalescent pol- len. The hollow, obconic gynoecious receptacle is shallow and splits apically into four erect or outcurved deltoid lobes that are ribbed and whit- ish within (Fig. 42G, H). The floral entrance is 10-12 mm wide and allows access to 100—400 short, purple-red conical styles that produce a sparse surface secretion (Fig. 13E). Flowers of both sexes emit a strong, pleasantly fruity floral odor likened by various observers to that of ripe 1985] LORENCE-— MONIMIACEAE 63 13. Summary of results of Tambourissa crosses carried out in Mauritius (1978-1979) and at the TABL Missouri Botanical Garden (1980) Species & Locality No. 9 Flowers Outcrossed T. amplifolia, Bel Ombre T. cordifolia, Pétrin T. E x T. ROMS T. peltata, P T. peltata x T Caa T. purpurea, cultivated, St. Louis T. quadrifida, Yemen Valley T. sieberi, Mt. Cocotte T. sieberi x T. tau T. tau, Perrier 8 (3 individuals) 4 (2 individuals) 8 (2 individuals) 7 (2 individuals) 2 (1 individual) 15 (3 individuals) 2 (2 individuals) 2 (1 individual) 12 (8 individuals) n No. ruits 96 Fruits 96 Devel- Fruit No.9? Flowers Devel- Fruit oped Set Selfed oped et 1 12.5 8 (3 individuals) 0 0 3 75.0 (Dioeci "n — — l 12.5 — — 7 100.0 (Dioecious) — — 2 100.0 — — — — — 15 (1 individual) 0 0 14 93.3 16 (3 individuals) O0 0 2 100.0 11 (2 individuals) 4 36.4 2 100.0 — — -— 2 100.0 16 (8 individuals) 0 e apricots, ripe stewed bananas, or ripe jackfruit, which is perceptible for several meters. The species is dioecious and therefore obligately out- crossing A mature flowering individual of each sex was selected for study totalling 23 hours of obser- vation time (Table 11). Pollinator activity began at dawn (ca. 5:45 A.M.) as the dense mists dis- persed. Numerous small dipterans then began congregating on both androecious and gynoe- cious flowers (up to 25 insects per flower; Fig. 20C) drawn by the strong, fruity odor. Drosoph- ila nasuta and Zaprionus vittiger were the most frequent visitors, with smalle f Ly- ciella and H. à (both Lauxaniidae) (Table 11). The insects fed on the pollen of the androe- cious flowers and drank dew and possibly stylar exudate from the gynoecious flowers. Consid- erable interfloral and interplant movement oc- curred at this time. Insect activity began to abate after 7:00-8:00 A.M. with an increase in temper- ature and wind velocity and generally ceased completely by 9:00 or 10:00 A.M. Pollen grains adhered to the bodies of all the insects examined, suggesting they are effective as pollen vectors, although no actual pollen counts were made. In addition, the population ned. of Tambourissa cordifolia was extremely at Pétrin (69 individuals/410 m?, or an ence 1,684/ha), with a mean nearest neighbor distance of 78.5 cm (Fig. 3C). Over 5096 ofthe individuals in this population were nd situated within 50 cm of each other (Fig. Judging by the relatively eer wild fruit set and good regeneration of Tambourissa cordifolia at Pétrin, ample cross pollination takes place. As Drosophilidae breed in decaying fruit and fungi, and Lauxaniidae breed in decaying vegetation (Borror & Delong, 1971), it is likely that signif- icant pools of these insects are available during the plants’ three to five month flowering period. Significantly, Thien (1980) also recorded two of the same dipteran genera (Drosophila and Homoneura) as pollinators of Drimys in New Guinea TAMBOURISSA AMPLIFOLIA A slender treelet 6-8 m tall, Tambourissa am- plifolia is occasional to locally common in the understory of some lowland and lower montane wet forest areas in Mauritius. It is monoecious m long-pedicellate flowers are open at a given time, each lasting for 10-15 days The white androecious flowers split into four flat lobes and resemble those of Tambourissa cordifolia, but are only half as large with 150- 200 stamens (Fig. 18B). The shallowly napiform gynoecious flowers also resemble those of T. cor- difolia, but are half as large with four everted lobes, a broad orifice, and ca. 300 short, yellow or purple styles situated in the shallow, discoid receptacle. Only slight amounts of mucilage are secreted by the styles. Flowers of both sexes pro- duce a faint, fruity odor. One individual of Tambourissa amplifolia was observed at Bel Ombre for a total of 4.5 hours during the morning (Table 1 1). Several drosophi- lids (Drosophila cf. nasuta and Homoneura sp.) repeatedly visited flowers of both sexes. Al- 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 E 14. Contents of Tambourissa quadrifida gynoecious flowers sampled at Matala Valley (Yemen), 979. TABL Mari in March, June, and July 1 No. Flowers No. Flowers Fallen From Contents From Tree 96 Total Ground % Total Empty 410 93.6 100 75.8 Mealybugs (Homoptera, probably Pseudoccidae) l 0.2 15 11.4 Ants (Hymenoptera, Formicidae) 14 3.2 4 3.0 Silk webbing & pupa (probably oe 6 1.4 3 23 Silk webbing & larv (probably ass hal 2 0.5 2 1.5 Coleoptera, Nitidulidae, aptoncus luteolus l 0.2 — — Detritus (feces) 3 0.7 5 3.8 E y egg cases 1 0.2 2 1.5 Springtail (Collembola) — 1 0.7 Total 438 100.0 132 100.0 though visits were less extensive than in T. cor- difolia, the same pollinator syndrome appears to e operative. Individuals of T. amplifolia are generally separated by distances less than 10 m, which the insects should be able to cross. Out- crossed flowers of T. amplifolia yielded a low fruit set (12.596) although it is unclear why (Table 13) TAMBOURISSA PELTATA A medium-sized dioecious canopy tree, Tam- bourissa peltata is common and widespread in wet and cloud forests throughout the island. Flowering lasts from March to June and each individual generally has three to ten flowers open at a given time, although a large androecious plant may produce up to 58 open flowers during peak flowering. The exposed flowers are pro- duced either singly or in groups of two to three on the gi leafless branches in the canopy and the upper trunk, thus making pollination Studies difficult. It is dioecious and obligately outcrossing. The broad gynoecious receptacle splits open by four or five ornate, dark purple-red verrucose sterile lobes that spread flat, resembling a Sta- pelia flower (Figs. 15A, 17E). The flat gynoecial disc is orange and bears ca. 400—500 flat, densely crowded columnar styles. At anthesis, the entire disc is covered by a layer of thick, sweet mucilage that functions both as a compitum and as nectar. Androecious flowers split gradually into four or five ultimately reflexed lobes bearing 200—400 stamens that release abundant, coalescent pollen (Fig. 17D). Both salmon pink and dark purple- red androecious color morphs occur, according to the individual. Flowers of both sexes produce a strong, sour odor of rotting fruit or tomatoes and persist on the tree for 10-15 days The species was studied at Mare Longue Pla- teau and at Perrier and Pétrin Nature Reserves (Table 11) for a total of 8.2 hours and supple- mented by additional insect collections. Most frequent visitors were Diptera, notably Syrphi- dae and Drosophilidae (Table 11). In the early morning, large numbers of Drosophila and Za- prionus congregated on both androecious and gynoecious flowers as in Tambourissa cordifolia, in addition to a few small Coleoptera (Enicmo- soma, Epuraea). As the sun rose higher and the temperature increased, the drosophilids gradu- ally left and were replaced by syrphids. The Syr- phidae Askarina and Ornida were active throughout the day, periodically visiting differ- ent flowers on the same and on different trees. They often remained on flowers for extended pe- riods to feed on the pollen (up to 43 minutes on a single flower for Ornida obesa), and also drank mucilage from the gynoecious flowers. Captured insects were found to have abundant pollen on their bodies. Other insect visitors included hon- eybees, which visited androecious flowers to gather pollen, but displayed no interest in the gynoecious flowers and their muciliage. The fo- livorous weevil, Scaevinus truncatus, while liv- 1985] LORENCE—MONIMI ACEAF 65 ing on T. peltata and feeding on its leaves and young flower buds, did not visit mature flowers and actually displayed an aversion to them, pos- sibly repelled by the strong odor. At Perrier, Tambourissa peltata occurred at comparatively low population densities of 26 in- dividuals/12,700 m" (or an estimated 21/ha), with a mean nearest neighbor distance of 975 cm (Fig. 3A). Because of the relatively large distances sep- arating individuals of this dioecious species, the strong-flying syrphids probably represent the most effective pollinators, although the role of drosophilids and small coleopterans as pollina- tors cannot be dismissed. TAMBOURISSA FICUS Tambourissa ficus is related to T. peltata but differs in being a small understory tree. It is usu- ally subdioecious, producing one or two open flowers generally of a single sex at the swollen base of the trunk (Fig. 20E, E) Although only gthe flow- ering season (Sept. -Dec. ), they have the greatest dimensions of any in the famil The urceolate-cupuliform gynoecious flower has a broad orifice and is lined with ca. 1,000- 2,000 long, yellowish pink setose styles which secrete small amounts of mucilage (Fig. 15C). wo androecious floral forms occur, the first being cupuliform-urceolate like the gynoecious flower, with erect lobes and white stamens (Fig. 20E). The other form splits open flat into five to seven ultimately reflexed lobes with pinkish purple sta- mens (Fig. 20F). The thick, apiculate stamens number 900-1,800 in both forms. Flowers of both sexes emit a fetid-aminoid, but slightly fruity, odor. A tree with flat purple androecious flowers at Brise Fer was observed for 0.8 hours at midday (Table 11). Male and female syrphids of the ge- nus Eumerus were attracted to the flowers (Fig. 20F), and both captured insects had pollen ad- hering to their bodies. The cupuliform white an- droecious flowers on a tree at Riviére des Galets (Fig. 20E) were visited by a different dipteran (unidentified). At Mt. Cocotte, however, large numbers of small Hydrophilidae beetles of the genus Cercyon were found in gynoecious flowers and on open purple androecious flowers on a single tree. Significantly, members of the same family also visited gynoecious flowers of Tam- bourissa sieberi at the same locality, although their role in the pollination of 7. ficus is uncer- n. Although no crosses were carried out for Tam- bourissa ficus, it is probably at least partly out- crossing because of its tendency towards dioecy. Scattered individuals as well as denser local pop- ulations occur, and syrphids would seem to rep- resent the most efficient pollinators, although the Coleoptera may also play a role. More extensive observations on the pollination biology of T. fi- cus are obviously required. Flowers of Tambourissa crassa (Réunion) emit a strong, rancid-fruity odor and appear to display the same pollination syndrome as 7. ficus. A gynoecious individual of this dioecious species at Col de Bellevue (Lorence 2468, MO) was re- peatedly visited by an unidentified dipteran which drank the mucilaginous secretion partly filling the mature gynoecious flower (Fig. 20D). MONIMIA OVALIFOLIA A low, dioecious tree restricted to small patches of cloud forest on the island's highest mountains, Monimia ovalifolia is the only member of the genus to occur in Mauritius. Its numerous small flowers are produced in ramiflorous inflores- cences from August to November. The globose gynoecious receptacle takes on a pink to orange color at anthesis, but remains closed except for a small pore through which the six to 12 glisten- ing white styles are exserted (Fig. 14B). These produce a slight surface secretion. The externally yellowish androecious flowers split into four or five recurved valvate segments (Fig. 14A) much like those of Tambourissa. The numerous white stamens possess paired basal appendages (as in Fig. 13A) and release coalescent pollen. Flowers of both sexes produce a strong, sweet odor. Androecious and gynoecious trees of Monimia ovalifolia were observed for 2.2 hours at Mt. Co- cotte (Table 11). The flowers appear to be my- ophilous and were repeatedly visited by small, unidentified Syrphidae that licked the styles and fed on the pollen. They were found to carry Mo- nimia pollen and could probably traverse the relatively short distances separating individuals of Monimia at Mt. Cocotte. It is therefore likely that syrphids are effective pollinators of Monim- ia ovalifolia in Mauritius. DISCUSSION Tambourissa species in Mauritius exhibit two different modes of pollination involving Coleop- 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN tera and Diptera, modes that occur in a number of other extant primitive angiosperms (e.g., Fae- gri & van der Pijl, 1971; Gottsberger, 1974; Thien, 1980). Thrips, which are known to visit flowers of other primitive angiosperms including Mo- nimiaceae (Gottsberger, 1977), were not found in this study, although it is possible that they may visit some of the small flowered Comorean and Madagascan species of Tambourissa. Flowers of Tambourissa quadrifida and T. sie- beri seem to be primarily cantharophilous with the fruity odors, dull colors, and numerous floral parts (i.e., stamens and styles) characteristic of this syndrome (e.g., Faegri & van der Pijl, 1971). he inferior ovaries and abundance of receptacu- lar idioblasts (see Morphology and Anatomy sec- tion) are probably also protective in this context. Gynoecious flowers of both species are further adapted for pollination by a variety of small, probably allotropic, Coleoptera and differ from the classic Magnoliaceous cantharophilous flow- er in the following ways: the flowers are long lived, apetalous, and remain permanently open. ermore, the gynoecious floral entrance is small, thus excluding many destructive preda- tors, although larvae of the indigenous moth, Opogona sacchari, successfully feed on flowers and developing fruits of at least three species, including 7. sieberi (Table 11). Both species pro- duce only traces of mucilage on their styles and ips deceive the insects by their strong, pe fruity odors, although the long styles of 7. pedes may also be utilized as food bodies. An- droecious flowers of all Tambourissa species of- fer only pollen as a reward, a resource taken ad- vantage of by a variety of Diptera and the introduced Apis, as well as by the effective pol- linators. Mass flowering of these species may compensate for pollinator inefficiency. The drab, apparently odorless flowers of Tam- bourissa tau seem to represent a specialized can- tharophilous syndrome adapted to pollination by a single insect, the folivorous curculionid, Scae- vinus truncatus, which feeds specifically on Mo- nimiaceae leaves. The flowers of this species rep- resent a supplementary food source for Scaevinus, nd T. tau has taken advantage of its presence for use as a readily available pollinator. The long publ pum a food source for the insect, while the basal sty bly trap pollen grains Homi its body and legs. The - peculiar T- o stamens with widely separated loculi may maximize pollen contact with Scaevinus, while e rendering the pollen load less liable to [Vor. 72 be eaten at once. In any case, flowers of T. i differ from the usual appear to be highly specialized. The energetic aspects of this relationship (cf. Heinrich & Ra- ven, 1972) should prove to be most interesting. The predominantly myophilous flowers of Tambourissa amplifolia, T. cordifolia, T. ficus, and T. peltata share a number of cantharophi- lous features with those of T. quadrifida and sieberi, i.e., strong fruity or aminoid odors and drab colors (in 7. ficus and T. peltata). In fact, the latter two species also attracted some Co- leoptera (Table 11), which is perhaps suggestive of a shift from cantharophily to myophily in Tambourissa. A number of other trends in these species suggest adaptations to allotropic fly pol- linators (cf. Faegri & van der Pijl, 1971), includ- ing: paler floral colors (white androecious and pale yellow or pink gynoecious flowers in T. cor- difolia and T. amplifolia, with white and salmon pink morphs occurring in androecious flowers of T. ficus and T. peltata, respectively), apically confluent anther loculi presenting the pollen more effectively (7T. amplifolia, T. cordifolia, T. pel- tata); shallower and broader gynoecious recep- tacles with a larger entrance (extreme in the flat, discoid flowers of 7. peltata), surrounded by at- tractive, spreading sterile lobes that act as land- ing platforms; more copious, sweet mucilaginous exudate with a nectar function in the gynoecious flower (T. peltata, Figs. 15A, 17E); a shortening of the stamens and styles to accommodate the insects' short probosces. Although certain common features occur in these species, there appear to be two distinct trends. The first is a “fruit fly" syndrome in- volving small drosophilids that occurs in Tam- bourissa amplifolia and T. cordifolia. These plants are small in stature, occur in comparatively dense populations, and have smaller and paler flowers with a sweet, fruity odor produced near the ground. Significantly, Thien (1980) recorded two of the same fly genera as pollinators of Drimys in New Guinea, suggesting that these insects may pollinate a greater number of extant primitive angiosperms than previously suspected. The second trend involves larger and more vigorous flies, primarily syrphids (in Tambouris- sa ficus and T. peltata). These are larger, more widely dispersed forest trees with more massive, pinkish to purplish flowers that emit sour, rancid, or otherwise fetid-aminoid odors with fruity overtones. Flowers are produced on the leafless upper branches (T. peltata) or near the ground [n 1985] LORENCE-— MONIMIACEAE 67 (T. ficus) and are exposed for easy access by the insects. Although androecious flowers of T. ficus were visited by syrphids and another dipteran in two cases (Table 11), flowers of both sexes of another individual attracted numerous hydro- philid beetles (Cercyon sp.). Hydrophilids were also found in gynoecious flowers of 7. sieberi at this same site. A more thorough study of T. ficus is needed to detail its significant pollinators. Monimia ovalifolia displays a number of my- ophilous features, namely, a sweet floral odor, yellow to pink or orange receptacles, shallow, permanently open white androecious flowers, well exposed stamens and styles. Its flowers are well adapted to the small, probably allotropic syr- phids that visit them In summary, species of Tambourissa in Mau- ritius exhibit two different modes of pollination based on Coleoptera and Diptera, and each mode displays two distinct trends involving modifi- cations in size, shape, color, and odor ofthe flow- er. Pollinator modes also appear to be related to demography and life forms. Pollinator special- ization presumably contributes to reproductive isolation in simultaneously flowering sympatric species (e.g., Tambourissa amplifolia and T. tau). Another factor contributing to reproductive iso- lation between sympatric species is different flowering times, clearly operative in Monimia species in Réunion (see section on Habitat and Ecology), and also for Mauritian Tambourissa species with common pollinators (e.g., T. ficus flowers from September to December; 7. peltata flowers from March to June). Reproductive iso- lation appears to be effective, as hybridization is extremely rare in the wild in Mauritius, although artificial crosses resulted in fruit set in three dif- ferent interspecific combinations (Table 13). Prolonged flowering periods and high floral output may compensate for low pollinator effi- ciency. Finally, dioecism and self-sterility in monoecious species can be viewed as represent- ing mechanisms to enforce outcrossing (e.g., Bawa, 1974; Bawa & Opler, 1975). Self-com- patibility in Tambourissa sieberi most likely off- sets low population densities and low efficiency of its small pollinators. SYSTEMATIC TREATMENT MONIMIACEAE Jussieu, Ann. Mus. Natl. Hist. Nat. 14: 133. 1809 (sub Monimiées). Evergreen trees or shrubs, rarely lianas, usually aromatic, frequently dioecious or monoecious, sometimes hermaphroditic. Leaves simple, usu- ally opposite and decussate, rarely ternate, al- ternate or anisophyllous, exstipulate, with lam- inar oil glands, the trichomes simple, fasciculate and/or stellate, the venation festooned brochi- dodromous, rarely craspedodromous, the mar- gin entire or serrate to dentate with glandular teeth. Inflorescence cymose or thyrsic, or the flowers fasciculate or solitary, either terminal, axillary, ramiflorous or cauliflorous, the brac- teoles small and caducous; flowers small to large, actinomorphic, rarely oblique, bisexual, polyg- amous or unisexual, the receptacle + well de- veloped, globose, ovoid to obovoid, urceolate, napiform, campanulate or discoid, the tepals subequal, spiral, whorled or decussate, large and petaloid to small or + lacking; androecious flow- ers with few to numerous stamens in 1-many series or irregularly disposed, the filaments usu- ally short and flattened, occasionally with a pair of basal appendages, the anthers erect, (2-)4- sporangiate, the loculi separate or confluent api- cally or rarely basally, lateral or + unifacial, val- vate or longitudinally dehiscent; gynoecious flowers with or without staminodes, the carpels several to numerous, rarely solitary, unilocular, shortly stalked or sessile, free or enclosed or im- mersed in the receptacle wall, the style short or elongated, the stigma terminal, dry or secreting mucilage, the ovule solitary, erect, horizontal or pendulous. Fruiting carpels exposed, or enclosed or immersed in the accrescent receptacle wall, indehiscent, often drupaceous with a membra- nous or fleshy mesocarp and/or aril-like appen- dage, or plumose (Atherospermoideae); endo- ca pendulous, the testa membranous, the endo- sperm abundant, oily and fleshy, the embryo straight, the cotyledons erect or divaricate. The Monimiaceae sensu lato include the subfamilies Atherospermoideae, Hortonioideae, Siparunoideae, Monimioideae, and Molline- dioideae, and pri 30g d near- ly 350 species in the tropics and subtropics, with the greatest diversity occurring in the southern hemisphere. KEY TO GENERA OF ueste IN THE MALAGASY REGIO la. Trichomes of leaf and stem in part or all stel- to x Blei or in fascicles of 2-4; abaxial laminar surface obscured by a dense, whitish indu- 68 ANNALS OF THE MISSOURI BOTANICAL GARDEN nt of trichomes; stamens bearing a pair of VAR DE on filament (Monimioideae) ...... 1. Monimia lb. Trichomes of leaf and stem either simple or rarely in fascicles of 2-4, never stellate; abax- ial laminar surface visible, not obscured by trichomes; stamens lacking appendages on filament (Mollinedioideae ). 2a. wt the e carpels superior, subses- aie Pin ane free on the disc, clavate to columnar; fruiting receptacle + discoid, concave to convex, bearing the exposed, + free drupes, not splitting at maturity. 3a. Buds and flowers with 7-15 large, thick obtuse tepals imbricate in 1-2 radial series; male floral receptacle Eid gradually at anthesis, not splitting „a 2. Decarydendron . Buds and flowers with 4—8(-16) small to minute decussate tepals; male flo- ral receptacle To a 4 valvate segments at a Ae 2b. Gynoecious aL an urceolate, c puliform to napiform, ellipsoid or b dric (discoid in T. peltata), the carpels w ° ] ing receptacle + closed, d in the wall, split- ting open irregularly at ma is Monimia Thouars, Hist. Vég. Iles France 35, tab. 9. 1804; Thouars, Hist. Vég. Iles Austral. Afriq. ed. 1: 35, tab. 9. 1805; Willd., Sp. PI. 4(2): 647. 1805; Juss., Ann. Mus. Natl. Hist. Nat. 14: 116. 1809; Poir., Encycl. Suppl. 3: 726. 1813; Spreng., Syst. 3: 906. 1826; Boj., Hortus Maurit. 289. 1837; Tul., Monogr. Monum. P 1855; A.DC., Prodr. 16(2): 661. . Adansonia, Ser. 1, 9: 133. 1868- , Hist. Pl. ed. 2, 1: 299. 1871; & Hook. Gen. Pl. 3: 139. 1883; Bak- er, Fl. Mauritius 289. 1877; Hobein, Bot. Jahrb. Syst. 10: 56. 1889; Pax, Pflanzenfam. 3(2): 101. 1891; Cordem., Fl. Réunion 299. 1895; Perk. & Gilg, Pflanzenr. 4, 101: 65. 1901; Perk, Pflanzenr. 4, 101 (Nachtr.): 39. 1911; Vaughan, Mauritius Inst. Bull. 1: 76. 1: : u Bullock, Kew Bull. 14: 44. 1960. D ti green trees, rarely shrubby; bud scales present. Leaves op- posite, decussate, entire, strongly discolorous, the abaxial surface obscured by a dense indument of simple, fasciculate and/or stellate-peltate tri- chomes, the venation festooned brochidodro- mous. Inflorescence pubescent, a thyrse or cyme, [Vor. 72 rarely leafy, either ramiflorous, axillary or subfoliar on leafless nodes, rarely terminal, the bracts and bracteoles caducous, the flowers borne in umbelloid units of 2-7. Male receptacle ped- icellate, obovoid or globose in bud, the apex with several pairs of minute, decussate tepals, splitting at anthesis into 4—6 + regular valvate segments, tomentose within, the short stamens numerous, multiseriate, the filament thin, subtended by a pair of basal appendages, the connective and fil- ament reddish brown, full of oil cells, blunt or slightly prolonged and rounded, the anther ovoid, bilocular, the loculi separate, parallel, longitu- dinally dehiscent, introrse; pollen n inaperturate, globose-urceolate to ovoid-urceolate, the apex with several pairs of minute decussate tepals, the carpels 4-14 per flower, unilocular, subsessile, free, enclosed within the receptacle which is lined with dense, simple hairs, the styles simple, + papillose, exserted through the narrow, 3-5-lobed orifice at anthesis, the ovule solitary, pendulous, anatropous. Ripe fruiting receptacle splitting into 3-5 irregular valvate lobes, pinkish red and part- ly velutinous within, exposing 4-12 ovoid-com- pressed fruiting carpels (drupes) surrounded by a fleshy orange mesocarp, the apical half De by a fleshy orange stylar aril, the endocarp w ish, thick, bony, mucronate, and dish crested, the surface foveate, foveate-reticulate to foveate-sulcate. Distribution. Monimia consists of three species endemic to the Mascarene Islands of Mauritius and Réunion. The presence of Monim- ia in Rodrigues is doubtful and based on a single collection of M. ovalifolia (Balfour s.n., K). As Balfour failed to mention the genus in his com- prehensive work on the botany of Rodrigues (1879), I suspect the specimen is wrongly labelled and may actually have come from Réunion where Balfour also collected. The Balfour s.n. collection most closely resembles plants of M. ovalifolia rom oe ponen by orbiculate leaves. Furthermo rigues does not even attain the minimum sae at which the species occurs in Mauritius and Réunion. TAXONOMIC HISTORY OF MONIMIA Monimia was described by A. Du Petit- Thouars in 1804 based on two species, M. ovali- folia and M. rotundifolia, which he collected in Mauritius and Réunion respectively. Du Petit- Thouars described the genus accurately and not- 1985] LORENCE—MONIMIACEAE 69 ed the close similarity between the androecious flowers of Monimia and those of Tambourissa (as Ambora Juss., and Mithridatea Comm. ex Schreb.). As Monimia differed from the latter most conspicuously in the structure of its gynoe- cious flowers, he named it after Monima, wife of Mithridates. In 1809, Jussieu created a new family (as Mo- nimieae) for Monimia, Tambourissa (as Am- ceae by various authors. Willdenow (1805) and Sprengel (1826) placed Monimia in subgroups of their *Dioecia," along with other anomalous genera belonging to unrelated families, but fol- lowed Du Petit-Thouars in noting its affinities to Tambourissa. In his monograph of the family, Tulasne (1855b) placed Tambourissa and Monimia in separate tribes. Monimia was included in his *Drupaceae" (Monimieae) together with Sipa- runa Aubl. (as Citrosma Ruiz & Pav.), Molli- nedia Ruiz & Pav., Kibara Endl., Hedycarya Forst., and Peumus Mol. (as Boldea Juss.) on the basis of their free, drupaceous carpels. Tulasne (1855b) also described an additional species of Monimia, M. citrina, which I consider to be syn- onymous with M. rotundifolia. Baillon (1868- 1870, 1871) generally followed Tulasne, includ- ing Monimia under his * Hortonia series” (Hor- tonieae) with many of the same genera as Tu- lasne, although he included a number of others also having free, drupaceous carpels. Bentham and Hooker (1883) recognized two large tribes of Monimiaceae on the basis of sta- minal dehiscence and position ofthe ovule. They placed Monimia with the bulk of genera now included in the Monimiaceae sensu stricto, a treatment largely followed by Hutchinson (1964). Monimia has been placed in the tribe Monim- ieae within the subfamily Monimioideae, along with Tambourissa, Palmeria F. Muell., and Hen- necartia Poisson by Pax (1891), Perkins and Gilg (1901), and Perkins (1911). Schodde (1970) cre- ated a separate subfamily for Peumus based on features of wood, pollen, and chromosome num- ber but ignored Monimia, leaving it in the Mo- nimioideae with the bulk of Monimiaceae sensu stricto. More recently, Thorne (1974) joined Monimia with Peumus as the sole members of the subfamily Monimioideae and created a new subfamily, the Mollinedioideae, to house the re- maining genera formerly included in the Monim- ioideae. SYSTEMATIC POSITION OF MONIMIA Baillon (1871), Money et al. (1950), and Ca- vaco (1965) all noted floral morphological sim- ilarities between Monimia and Palmeria. In ad- trends (Garratt, 1934; Money et al., 1950). Other shared characters in Monimia and Palmeria in- c ind tofsimple iculate an stellate trichomes, an urceolate gynoecious re- ceptacle enclosing a small number of subsessile or sessile carpels, and a fruiting receptacle which splits open to reveal the ripe carpels set against a pink or red inner surface (Corner, 1976; En- dress, 1980b). In spite of superficial similarities in the fruiting receptacles of the two genera, how- ever, which may represent parallel lines of evo- lution, important differences also exist. The car- pels of Palmeria are anatomically different from those of Monimia and possess a well-developed, hard and shiny black exocarp and thin mesocarp of starch-filled cells absent in the latter (Corner, 76) — o Monimia appears to have closer affinities with Peumus, a monotypic genus endemic to Chile. Both genera have horizontal axillary buds, sta- mens with paired appendages on the filaments, large, spinose pollen grains, the spines and sur- face composed of cable-like strands (smaller and spinulose in Palmeria), receptacles covered by a dense indument of simple, fasciculate and stel- late trichomes, carpels with a fleshy mesocarp surrounding the bony white, sculptured endo- carp (Fig. 19A, B), and also share the dioecious habit. In Peumus, however, the stamens are tetraspo- rangiate and dehisce laterally, whereas in Mo- nimia they are bisporangiate and dehiscence is nearly introrse along the inner margin (Fig. 13A). In these respects Monimia and Peumus appear to provide a link between the Atherospermoide- ae, with their bisporangiate anthers having val- vate dehiscence and similar staminal append- ages, and the Mollinedioideae, which have tetrasporangiate anthers with lateral dehiscence and lack staminal appendages (Sampson, 1969a, 1969b). As a result, the two genera do not fit satisfactorily into either subfamily. Specialization in Monimiaceae is generally considered to be expressed in terms of floral sim- plification and reduction, both in terms of size and number of parts (Money et al., 1950; En- dress, 1980b). In these respects, Monimia has attained a greater degree of floral morphological 70 ANNALS OF THE MISSOURI BOTANICAL GARDEN specialization than Peumus, in spite of the lat- ter's more specialized wood anatomy (Garratt, 1934; Money et al., 1950; Lemesle & Prichard, 1954). Although also dioecious, Peumus has re- tained a greater constellation of unspecialized ta shallow floral receptacle bearing exposed carpels in the manner of the putatively primitive Hor- tonia (Lemesle & Prichard, 1954; Endress, 1980a). Flowers of Monimia have reduced, scale- like decussate tepals, are closed in bud, and the female Peri encloses the carpels—all ad- vanced featur Similarities eh Monimia shares with Peu- mus, and to a lesser degree with Pa/meria, sug- gest possible origin from a common ancestral stock. A Monimia precursor may have originally existed in Africa or Madagascar where it sub- sequently became extinct, but not before reach- ing the Mascarene Islands via long distance dis- persal. The Mascarenes thus provided a refuge for Monimia. Current work with sieve tube plastid analysis by Behnke (1977, and pers. comm.) has shown that Palmeria and Peumus possess fundamen- tally different types of sieve tube plastids (S-type and P-type, respectively). As Monimia ovalifolia possesses S-type plastids (Behnke, pers. comm.), this feature at least would seem to deny a close relationship to Peumus. Sieve element plastids unquestionably provide an additional taxonom- ically useful character when used in conjunction with other available data, but do not provide an adequate basis for fragmenting the Monimiaceae into some five families as proposed by Behnke (1981) In summary, Monimia displays a number of advanced features, certain of which (i.e., its sta- mens) are transitional between the Mollinedioi- deae and the Atherospermoideae. As Monimia shares the greatest constellation of features with Peumus, however, the two genera are best placed together as constituting the Monimioideae as proposed by Thorne (1974). KEY TO THE SPECIES OF MONIMIA la. Leaves amplexicaul with a ined cordate base, sessile to subsessile; Réunion .......... "M. amplexicaulis i Leaves A petiolate, the Loi neithe er union and = Mauri 28. Stems. petioles and inflorescence hir- sute-velutinous; trichomes on adaxial [Vor. 72 surface of lamina mixed simple, ene e ov a with free, + ascen 2. M. rotundifolia 2b. site scabrous or idis not velutinous; tri- h s on adaxial surface of lamina uni- with horizontal united basally = xw their length; Réuni ion and Mauriti 3. M. ovalifolia 4: 1. Monimia amplexicaulis Lorence, Bull. Mus. Hist. Nat. (Paris), Ser. 4, Sect. B, Adansonia 3(3): 294, pl. 1, fig. 1a. 1982. TYPE: Réunion. Hauts du Tévelave: forestry road along the Domanial line, cloud forest transitional to heath formation, ca. 1,700 m, 6 Mar. 1979 (fl, fr), Lorence 2504 (holotype, MO; iso- types, K, MAU, P, REU, Z). Shrub or small tree attaining 10 m tall and 30 cm diam., the bark grayish brown, longitudinally fissured, the new growth pale grayish to fulvous, velutinous-tomentose with predominantly short- armed stellate trichomes, mixed with long simple and fasciculate trichomes 0.5-1 mm long, the mature leafy stems 3-5 mm diam., velutinous- tomentose. Leaves opposite, sessile tó subsessile; petiole stout, 3—5 mm long by 3—5 mm diam., velutinous-tomentose; lamina subcoriaceous to coriaceous, ovate to broadly ovate, broadly el- liptic to suborbiculate, (40—)60—130 mm by (32-) 40-110 mm, the apex shortly acute, shortly acu- minate, mucronate or rounded, the base deeply cordate, amplexicaul, the secondar y veins (4—)5— 7 pairs, making a 40-60? angle with the costa, the venation depressed adaxially, visible to 3°, prominent and raised abaxially, visible to 4°, the adaxial laminar surface pubescent with predom- inantly stellate trichomes with 4—12 short hori- zontal arms, mixed with varying proportions of long simple and fasciculate trichomes with 2-4 ascendant arms, eventually glabrescent and + pustular, glandular, the abaxial surface obscured by a dense whitish to pale yellowish indument of the same trichomes, the margin entire, pu- bescent, plane to slightly revolute. Inflorescence a congested, fulvous velutinous-tomentose thyrse or rarely short cyme of (3-)7-37 flowers (gynoe- cious inflorescence more congested than the an- droecious), branching to 2*(-3?) leafy or not, ramiflorous or subfoliar on leafless nodes (the primary axis often continuing to grow as a leafy stem), axillary or rarely terminal, the floral axis (5-)10—-60(-75) mm by 2-3 mm, often subtended by l-several pairs of caducous, brown ciliate- pubescent naviculate bracts to 7 mm by 3 mm, LORENCE— MONIMIACEAE 7 1 1985] the peduncle 4-12 mm long, bearing an umbel- loid unit of (2—)3(—6) flowers, occasionally sub- tended by a caducous, velutinous bracteole to 2 mm long or by a small leaf. Androecious flower in bud ellipsoid to obovoid, 2-7 mm long by 3- 5.5 mm diam., velutinous-tomentose, the pedi- cel 3-7 mm by 1-1.5 mm, often subtended by 1-2 minute subulate bracteoles; at anthesis deep- ly 3-6-fid, 10-13 mm diam., the lobes ultimately reflexing; stamens (40—)60-150, 1.5-2.5 mm long, the anthers ellipsoid, ca. 1 mm wide, occupying ca. '4—'2 total length of the stamen, the connec- tive slightly prolonged and rounded, the internal receptacle surface tomentose, the simple hairs 0.3-0.8 mm long. Gynoecious flower in bud glo- bose-urceolate to ovoid-urceolate, 4-5 mm long by 3-4 mm diam., at anthesis the orifice 0.8-1.2 mm diam., the styles 5-10, exserted for 1-2 mm, the internal receptacle surface velutinous with simple hairs 0.5-1.5 mm long, the pedicel 1.5- 2.5 mm by 1-1.5 mm. Fruiting receptacles in clusters of 1-10 on the leafless branches, ovoid to subglobose, 7-15 mm diam., externally pu- bescent with stellate, fasciculate and simple tri- chomes, the carpels 1-10 per receptacle, ovoid to ellipsoid, 5-7 mm long by 3-5 mm thick, mucronate, with a longitudinal crest, the endo- carp deeply foveate-reticulate. Figure 21. Distribution. Endemic to Réunion (Fig. 22). abitat. Monimia amplexicaulis is occa- sional to abundant locally as a component of mid to upper level cloud forest and lower limits of the Philippia heath formations from ca. 1,400 to 2,100 m. It is most frequent on the sheltered, leeward escarpments of the “cirques,” e.g., at Petit Matarum, Coteau Kerveguen above Cilaos, Hauts du Tévelave, etc. ÉUNION. CIRQUE DE CILAOS: ascent to Col de Taibit, 22 Mar. 1974 (fr), Bosser 21677 (P); Plaine des Fraises, low Col de Taibit, 1,900 m, 2 Feb. 1967 (fl), Cadet 663 bis (REU); Cilaos to Col de Taibit, 1,950 m, 22 Mar. 1974 (fr), Coode 2558B (K); Montée Cilaos, Co- teau Kerveguen, Feb. 1971 (fl, fr), Bosser 20752 (P); path to Coteau Kerveguen, 1,500 m, 16 Nov. 1973 (fr), Cadet 4512 (REU); Coteau Kerveguen, 1,500-1,700 m, 24 Feb. 1979 (fl), Lorence & Rolland 2443 (MAU, MO); (f), Lorence & Rolland 2444 (MAU, MO); forest of Mare à Joseph, 1,400 m, 10 Jan. 1967 (fl), Cadet 663 (REU); 1,200-2,000 m, Aug. 1943 (fl), Rivals s.n. (TL-R); 2,000 m, 11 June 1943 (fr), Rivals s.n. (TL-R); wet forest at Petit Matarum, 2,000 m, 22 Feb. 1975 (fl, P Cadet 5049 (REU); (fl), pe 5050 (REU); 4 Feb. 1968 (fl), Capuron 28210-SF (P); 6 Mar. 1971 (fl), I 1118 (P); (fr, Friedmann 1119 (P); path to Piton des Neiges, 15 Apr. 1978 (fl, fr), Bosser 22498 (P); 28 Mar. 1973 (fr), Lorence sub MAU 15693 (MAU), (fl, fr), Rivals s.n. (TL-R). CIRQUE DE MAFATE: Col de Fourche, 1,800 m, 29 May 1976 (fr), Cadet 5435 (REU); valley of Bras Bémale, 1,500 m, 13 May 1976 (st), Cadet 5404 (REU); above Belier, on Sentier des Scouts to Aurére, 1,500 m, 14 May 1976 (st), Rich- 132 (K). PLAINE DES CAFRES: base of Piton Bleu, 1,500 m, 11 July 1943 (fl), Rivals s.n. (TL-R); (fr), Rivals s.n. (TL-R); 24 Feb. 1945 (fr), Rivals s.n. (TL- x i 2,100 m, Feb. 1971 vi R). P Cirque de Salazie near La Nouvelle (sic), 1,300 m, 17 L (TL-R). PLAINE DES TAMARINS: 20 Aug. 1944 (st), Rivals .n. (TL-R). HAUTS DU TÉVELAVE: 1,700 m, 21 Sept. 1974 (fr), Cadet 4797 (REU); (fl), Cadet 47 97 bis (REV); along the yes line, 1,700 m, 6 Mar. 1979 (st), Lorence 2501 (MO); (fl), Lorence 2502 (MAU, MO); (fl, fr), Lorence py (MAU, MO). Monimia amplexicaulis shares morphological features with M. rotundifolia, including the pres- ence of three types of foliar trichomes (stellate, fasciculate, and simple; Fig. 9A). In M. amplex- icaulis, the trichomes are predominantly stellate with short arms and not predominantly long, fasciculate and simple as in M. rotundifolia. Monimia amplexicaulis further differs from the latter by its sessile, deeply cordate, and amplex- icaul leaves with 5-7 secondary veins and gen- erally occurs at somewhat higher elevations in more sheltered habitats than M. rotundifolia, often at the cloud forest/heath formation inter- face. Vernacular names. Mapou, Mapou des hauts (Réunion). 2. Monimia rotundifolia Thouars, Hist. Vég. Iles France 35, tab. 9, fig. 2. 1804; Thouars, Hist. Vég. Iles Austral. Afriq. 21, 34, tab. 7, fig. 2. 1805; Willd., Sp. Pl. 4(2): 647. 1805; Spreng., Syst. 3: 906. 1826; Poir., Encycl., Suppl. 3: 727. 1843; Tul., Ann. Sci. Nat. d 4(3): 32. 1855; Tul., Monogr. Mon- . 310, tab. 29a-j. 1855; A.DC., Prodr. 160): 661. 1868; Cordem., Fl. Réunion 299. 1895; Perk. & va ogee: 4, 101: 66, fig. 18A-D. 1901; , Pflanzenr. 4, 101 (Nachtr.): 39. 1911 dise iniu Balfour s.n. collection from Rodrigues); Ll Gatt. Monim. fig. 34A-D. 1925. TYPE: Ile Bour- bon (Réunion). Without precise E o (ca. 1800-1801), Petit- Thouars s.n. (holotype, P). Ambora tomentosa Bory, Voy. Iles Afrique 1: 317, tab. 13. 1804. Myrtus villosa Spreng., Syst. 2: 487. 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 FicunE 21. Monimia amplexicaulis Lorence. — A. Habit, male, one leaf removed. — B. Detail of adaxial leaf surface. — C. Androecious flower at anthesis. — D. Stamen, adaxial view.—E. S i Habi : : i ower i tion. — I. Gynoecious flower, apical view to show pore (styles removed).—J. Ripe fruiting receptacle, dehisced. — K. Fruiting carpel, lateral view. A-E. Lorence 2444 (MO). F, J, K, Lorence 2404 (MO). G-I. Lorence 2503 (MO). Bars equal 10 mm in A-C, F, J, and 1 mm in D, E, G-I, K. 1985] 21°20'S 55920'E LORENCE-— MONIMIACEAE 10 km A M. amplexicaulis O M. ovalifolia .O M. rotundifolia FiGureE 22. Distribution map of Monimia in Réunion. 1825, nom. superfl., based on type of Ambor tomentosa Bory. Eugenia bier be Poir. Encycl., Suppl. 3: 124. 184 3,n PE: ably R Réunion), Bory de St. Vincent s. n. in 1821 (holotype: G-DC,n microfiche, MO). Monimia citrina Tul., Ann. Sci. Nat. ues AGE 32. without precise locality, ichaud s.n. in 183 1837 (lectotype, P, here j patas M. ovalifolia sensu Cordem., Fl. Réunion 300. 1895 non Thouars. Shrub or low tree attaining 10 m tall and 60 cm diam., the new growth velutinous-tomentose, pale gray to fulvous, the simple and fasciculate trichomes 0.5-1 mm long, mixed with short stel- late trichomes, the mature leafy stems 3-6 mm diam., velutinous-tomentose. Leaves petiolate; petioles 5-20 mm by 2-4 mm, velutinous-to- mentose; lamina coriaceous, ovate to broadly ovate, elliptic to broadly elliptic or orbiculate, (35-)45-150 mm by (23-)35-160 mm (attaining 320 mm by 300 mm on vigorous sucker shoots), the apex shortly acuminate, acute, mucronate, obtuse, rounded or rarely retuse, the base die ded decurrent, acutely cuneate, obtuse, rounded o shallowly cordate, the secondary veins 3-7 pairs, making a 45-60? angle with the costa, the ve- nation depressed adaxially, visible to 3?, abaxi- ally raised and visible to 4°, the adaxial laminar surface strigose-hirsute, generally with predom- inantly long, simple hairs and fasciculate tri- chomes with 2-4 long, ascendant arms, mixed with varying proportions of stellate trichomes with 4-12 shorter horizontal arms, the surface glandular, eventually + pustular, the abaxial laminar surface obscured by a dense, whitish to pale yellowish indument of the same trichomes, the margin entire, pubescent, plane to slightly 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN revolute. Inflorescence a congested to lax thyrse (rarely a short cyme) of (3—6-76 flowers (the gynoecious infl congested than the androecious), branching to 2—3°, leafy or not, ve- lutinous-tomentose, either ramiflorous, subfoli- ar on leafless nodes (the primary axis sometimes continuing to grow as a leafy shoot), or axillary, the floral axis (7—)15—90(-130) mm by 1.5-2.5 mm, often subtended by 1-4 pairs of brown, scale-like, pubescent or ciliate bracts to 10 mm by 4 mm, the secondary axes 2-25(-50) mm long, occasionally subtended by a single caducous brown bracteole 2-3 mm long, the peduncle 2- 12 mm long, bearing an umbelloid unit of 2-6 flowers. Androecious flower in bud ellipsoid, obovoid or subglobose, 4.5-8 mm by 3-5 mm, velutinous-tomentose, the pedicel 3-13 mm by 0.5-1 mm, often subtended by 1—2 minute brac- teoles; at anthesis deeply 3—5-fid, 4-14 mm diam., the lobes ultimately reflexing; stamens (25-)50- 105, 2-3.5 mm long, the anther 1-1.4 mm wide, ellipsoid, occupying !⁄4—1⁄ total length of the sta- men, the connective + thickened, blunt or slight- ly apiculate, the internal receptacle surface to- mentose, the hairs simple. Gynoecious flower in bud ovoid-urceolate to globose-urceolate, 3.5-5 mm by 2.5-4 mm; at anthesis the orifice 1-1.2 mm diam., the styles 5-10, exserted for 1-2 mm, the internal receptacle surface velutinous, the short hairs simple, the pedicel 1.5-5 mm by 1- 1.5 mm. Fruiting receptacles borne in clusters of 1-20 on leafy or leafless branches, ovoid to sub- globose, 6-14 mm diam., externally pubescent with simple, stellate and fasciculate trichomes, the carpels 1-8 per receptacle, ovoid, 6-8 mm long, 4-6 mm wide, 4-5 mm thick, mucronate with a longitudinal crest, the endocarp thick, the surface deeply foveate, foveate-reticulate or fo- veate-sulcate Distribution. Endemic to Réunion (Fig. 22). Habitat. Monimia rotundifolia is an occa- sional to locally abundant component ofthe cloud forest zone, rarely ranging into the Philippia heath zone, from ca. 1,000 to 2,000 m. RÉUNION. BELOUVE: Apr. 1956 (fr), Bosser 7586 (P); 1,500 m, Apr. 1956 d "poi 9354 (P). BOIS DE NEFLES: 1,500 m, 17 Sept. 1974 (fl), Cadet 4787 (REU); (fl), Cadet 4788 ds (fl), Cadet 4789 (REU). UCAN-LAUNAY: woods above Boucan-Launay (fr), Boivin 1120 (P). BRETAGNE: above Srs 1,000 m, fe 1945 (fr), goce s.n. (TL-R). BRÜLÉ DE ST. DENIS: 1,300 m, May 1957 (fl), Bosser 11577 (P); 1,000 m, 22 Oct. 1973 Pa Bosser 21332 (P, 2 sheets), near Mamode Camp, 1,200 m, 13 Apr. 1968 (fr), Cadet [VoL. 72 1333 (REU); vicinity of Mamode Camp, at Plaine des Chicots, 31 Jan. 1968 (fr), Capuron 28161-SF (P, 2 sheets); St. Francois, St. Denis (fr), Cord BÉBOUR: Bébour Forest, 4,500 ft. (ca. 1,350 m), 1967 (fl), Barclay 554 (K, MAU); B lébour Pass and Riviére des Marsouins, 1,350 m, 21 Oct. 1973 (fl), 1974 (fl, fr), Cadet 5197A (REU); 29 May 1975 (fl), Cadet 5197B (REU); Bébour Peak, 1,700 m, 27 1973 (fl), Coode et al. 4231 (K, 2 sheets; REU); (fl), Coode et al. 4235 (K); (fr), Coode et al. 4237 (K, REU); (fl), Coode et al ann 2 (P); 1,300-1,600 m, 23 Feb. 1979 (fl), Lorence 2M O); 1,300 m, 6 May 1976 (fl), Richardson et al. 4086 (K, MO); (fl), Richardson et al. 4087 (K, MO); (fl), Richardson et al. 4088 (K, 2 sheets; MO). CIRQUE DE CILAOS: Petit Matarum, 6 Mar. 1971 (fr), Dun 1119 (P); Cilaos at Matarum, 1,300 m, June 1943 (fl), Rivals s.n. (TL-R). CIRQUE DE MAFATE: Sentier des Scouts, crest above Ilet à Malheur, 1,200 m, May 1976 (fr), Richardson et al. 4138 (K, 2 sheets). CIRQUE DE SALAZIE: path from Grand Ilet to Roche Ecrite, 1,900 m, 15 Oct. 1974 (fl, fr), Cadet 4836 (REU); below Grand Ilet at Salazie, 1,600 m, 26 Sept. 1977 (fl), Cadet 5254 (REU); ae Cadet 5255 (REU); Terre Plate, 1,600 m, 7 Apr. 1976 (st), Cadet 5334 (REU). MAiDo: road to Maido ea Petit France, 1,200 m, 22 Feb. 1979 (fr), Lorence & Cadet 2413 (MAU, MO); Maido to Le Guillaum road, 1,550 m, 12 May 1976 (fr), Richardson et al. 4121 (K, MO); (fl), Rich- ardson et al. 4122 (K). MASSIF DE LA FOURNAISE: below the shelter at Pas de Bellecombe, 2,000 m, 25 July 1962 (fl), Cadet x. (REU). MORNE DE PATATES À DURAND: 1,200 m, Hauts du Bois de Néfles à St. Denis, n Sept. 1976 (fl), es 5242 (REU); (fl), Cadet 5243 REU). PLAINE D'AFFOUCHES: June 1945 (fl), Rivals s.n. (TL-R). PLAINE DES CAFRES: open heathland, 5,200 ft. (ca. 1,700 m), 2 Aug. 1968 (8) Barclay 47 1 (K, MAU); Sentier Coteau Maigre, Feb. 1971 (fl, fr), Bosser 20476 (P); Piton Mare a Bouc, 1,600 m, 7 Nov. 1966 (fl), Cadet 503 (P, REU); foot of Piton Lepervanche, 1,700 m, 26 Oct. 1974 (fl), Cadet 4859 (REU); Col de Belle- vue, 30 Nov. 1974 (fl), Friedmann 781 (P); (fl), Fried- mann 782 (P); Col de Bellevue, 1,600 m, 27 Feb. 1979 (fl, fr), Lorence 2471 (MAU, MO); (fl), Lorence 2472 (MAU, MO); (fl), Lorence 2473 (MAU, MO); (st), Lo- rence 2474 (MO); 17 July 1979 (fl, fr), Lorence 2773 (MO, Z); (f), Lorence 2774 (K, MO, Z); (fl), Lorence 2775 (K, MAU, MO, Z); base of Piton Bleu, 11 June 1943 (st), Rivals s.n. (TL-R); Grande Montée, mid ides upper parts, 12 June 1943 (fl), Rivals s.n. (TL-R); low p art 2 ce at La Grand Montée, 1,200 m (st), Rival R); scrub and cloud forest, 1,700 m Tos X Schlieben y (B, MAU); Grand Mone 1,600 m, 28 964 (fl), Staub sub MAU 11105 RB : Roche Ec 975 (fl), Bernardi 14936 (P); (fl), Bernardi 14943 (Py; Roche Ecrite, moist forest, 1,300 m, 23 July 1961 (st), St. John 26527 (G, 2 sheets; K). LORENCE— MONIMIACEAE 75 1985] PLAINE DES MAKES: 1,100 m, June 1957 (fl), Bosser 12216 (P); 1,400 m, A e 1975 (st), Cadet 5154 (REU). PLAINE DES PALMISTES: mountainous regions (fl), Cor- demoy s.n. (TL-R); Petite Plaine above Plaine des Pal- mistes, 1,200 m, 25 Feb. 1979 (fr), Lorence & Rolland 2453 (MO). PLAINE DES SALAZES: high altitude ericoid Me duis Apr. ^d (fr), Alphonsine s.n. (P). RIVIERE E L'EST: far u e Riviére de l'Est, 1,750 m, Nov. 1976 (fl), Cadet 5710 (REU); (fl), Cadet 5711 (REU). RIVIÈRE DES REMPARTS: Roche Plate path, 6,500-6,700 ft. (ca. 2, a m), 16 Nov. 1967 (fl), Barclay 498 (K, AU); ca. 5,400 ft. (ca. 1,700 m), 20 Nov. 1968 (fl, fr), Barclay d (K, MAU); Vallée de la Riviére des us rts, 1,700 m, 20 Nov. 1968 (fl, fr), Cadet 1727 REU); 7 Mar. 1975 (fi), Cadet 5103 (REU); (fr), biis 5104 (REU); (fr), Cadet 5105 (REU). SAINTE AGATH 19 July 1875 (fl), De L'Isle 383 (K; P, 2 sheets). SAINTE KA: : 979 (st), Lorence & Rolland 2532 (MAU, MO). TÉVE- AVE: Hauts évelave, 1,500 m, 21 May 1975 (fr), push 22189 (P. 2 ii 1,300 m, 5 Sept. 1975 (fl), Cadet 5255 (REU); ca. 1,500 m, 25 Nov. 1968 (fl), Edgerley E4 Sea MAU 13372 S May 1976 (fl), Friedmann 281 1,300-1,400 m, s Feb. 197 MO); 1,700 m, 6 Mar. 1979 nr ’ Lorence 2505 (MAU, MO), (fr), Lorenee 2506 (MAU, MO). WITHOUT PRE- Richard 27 (P, 2 sheets). LOCALITY (ISLAND) UN NOWN: (ñ), Anon. sub Herb. ORSTOM 4999 (MAD); (A), Anon. sub MAU 2993 (MAU). Monimia rotundifolia is an extremely variable species as to size and shape of the lamina, al- though its leaves are never sessile to subsessile and deeply cordate-amplexicaul as in M. am- plexicaulis. Composition of laminar trichomes is also a variable feature. Long simple hairs pre- dominate on the adaxial leaf surface of many individuals (e.g., Fig. 9C), but in others large mbers of long fasciculate and shorter stellate trichomes are present (e.g., Fig. 9B), approaching the condition found in M. amplexicaulis. Bojer (1837), followed by Baker (1877), erroneously stated that M. rotundifolia occurred in Mauri- tius, but cited no collections, and no authentic specimens are known from Mauritius. Its wood is occasionally used for construction in Réunion and is said to be virtually incombustible. Vernacular names. Mapou, Mapou a grandes feuilles, Mapou blanc, Mapou des hauts (Ré- union). 3. Monimia ovalifolia Thouars, Hist. Vég. Iles France 35, tab. 9, fig. 1. 1804; Thouars, Hist. Syst. 3: 906. rit. 289. 1837; Poir., Encycl., Suppl. 3: 727. 1843; Tul., Monogr. Monim. 309. 1855; A.DC., Prodr. 16(2): 661. 1868; Baker, Fl. Mauri- tius 289. 1877; Cordem., Fl. Réunion 300. 1895; Perk. & Gilg, Pflanzenr. 4, 101: 65, 1: 76. 1937. TYPE: Ile de France (Mauritius). Without precise locality (ca. 1800-1801), Petit-Thouars s.n. (lectotype, P, here des- ignated; isolectotypes, P, 2 sheets M. did var. - boryana A.DC., Prodr. 16(2): 661. PE: without precise locality (probably Tua us or Réunion) (ca. 1821), B Vincent s.n. (holotype, G- DC, n.v.; microfiche, MO). M. M sensu Cordem., Fl. Réunion 300. 1895 non M. "ow Ve sensu Perk., Pflanzenr. 4, 101 (Nachtr.): on Thouars, as to Balfour s.n. collec- tion on Redndd es (K). Shrub or small tree 7-15 m tall and 40 cm diam., the bark pale brown, smooth, flaking, the new growth scurfy-pubescent, the mature T stems scurfy to scabrous, glabrescent, 2.5-5 m diam., the stems, petioles, costa, and inficles- cence axes scabrous to scurfy with mostly small, appressed stellate-peltate trichomes with 8-16 short, horizontal arms, mixed with fewer simple hairs and fasciculate trichomes with 2-8 long, ascendant arms. Leaves opposite, petiolate; pet- ioles EF pe mm a 2-4 mm; lamina subcoria- ceous to us, ovate-elliptic, elliptic, broadly iie suborbiculate or orbiculate, oc- casio Fame obovate, (30-)50-130(-170) mm by (20-)30- Phin mm (attaining 200 mm by 200 mm with petioles 35 mm long on vigorous sucker shoots), the apex shortly acuminate, shortly acute, obtuse, rounded or retuse, the base cutely d t, acute, obtuse, rounded or shal- lowly cordate, the secondary veins 4-7 pairs, making a 50—65? angle with the costa, the ve- nation P dide adaxially and visible to 2(—3?), raised abaxially and visible to 3(—4°), the adaxial laminar surface scabrous with scattered, + ap- £e 76 ANNALS OF THE MISSOURI BOTANICAL GARDEN — 20%0's T p P srsoE / ` ) 7 i / ç Ç x € M. ovalifola — _ SSA / i N PR S P 3 \ < aN “ a E ad N ( | a / y ( S N I T 20 9 - 4 ES > ` bes] > ~ d 1 . Í "n " . P. ^. J^ ud e px ( t N Uf \ 10 km FIGURE23. Distribution map of Monimia in Mau- ritius pressed stellate-peltate trichomes with 6—16 short, horizontal arms, the costa and secondary veins si scattered simple and fasciculate trichomes, eventually glabrescent and + pustular, glandular, the abaxial laminar surface obscured by a dense whitish indument of the same trichomes, the margin entire, plane to slightly revolute. Inflo- rescence a scurfy to scabrous thyrse (rarely a short e or pleiochasium) of (3—)8-87 flowers, y or not, ramiflorous, either subfoliar on laei nodes (the primary axis sometimes continuing to grow as a leafy stem), axillary, or rarely terminal, the floral axis (5-)10-120(-165) mm by 1.5-3 mm, often sub- tended by I—several pairs of caducous brown, ciliate to pubescent naviculate bracts to 7 mm by 3 mm, the peduncle 3-20 mm by 0.5-1 mm, often subtended by a single caducous brown bracteole 2-5 mm long, or by a small leaf, bearing an umbelloid unit of (2—)3(—8) flowers. oe cious flower in bud ellipsoid to obovoid, 3-5 m by 2.5-3 mm, the ied 1-7 mm by 0.3.0.6 mm, racteolate; at anthesis deeply 4-6-fid, 5-7 mm Ga the lobes reflex- ing; stamens ca. 45-60, 1.5-2 mm long, the an- ther ellipsoid, ca. 1 mm wide, comprising ca. '^- % total length of the stamen, the connective rounded to apiculate, the internal receptacle sur- face pilose with short simple hairs. Gynoecious flower in bud ovoid-urceolate to subglobose-ur- [Vor. 72 ceolate, 2.5-4 mm by 2.5-3.5 mm; at anthesis the orifice 0.6-0.8 mm diam., the styles 6-12, exserted for 0.5-1.5 mm, the internal receptacle tacles borne on the leafy or leafless stems, in clusters of 1—20, ovoid to subglobose, 5-12 mm diam., the apex + acuminate, externally sca- brous with scattered stellate trichomes, the car- pels 1-5 per receptacle, ovoid-compressed, 5-7 mm long, 4.5-5.5 mm wide, 3—4.5 mm thick, mucronate, with a longitudinal crest, the endo- carp surface deeply foveate-sulcate. Distribution. Endemic to the Mascarene Is- lands of Mauritius and Réunion (Figs. 22, 23). Habitat. The species is occasional to locally common in lower montane wet and cloud forest in Réunion from ca. 500 to 1,700 m. In Mau- ritius, Monimia ovalifolia is known only from patches of cloud forest on the summits of Mts. Le Pouce, Cocotte, Lagrave, and Piton Grand Bassin (ca. 550—770 m), and also occurs in wet forest at Plaine Champagne (ca. 450 m). URITIUS. BASSIN BLANC: path near Bassin Blanc, Te PS sub MAU 13081 — GRAND BASSIN: summit area, Piton Grand Bassi . 700 m, 31 Jan. 1976 (fr), Coode 4696 (K, 2 Hen. 18 Oct. 1978 (fl), Lorence 1857 (MAU, MO); Chemin Cheval near Grand Bassin, 23 Jan. 1939 (fr), Vaughan sub MAU 18546 (MAU). LE POUCE MT.: around summit, June 1860 (fl, fr), Ayres s.n. (K); at the upper limit of woods at the base of Mt. Pouce, 5 Sept. 1849 (fl), Boivin s.n. (P); (fl, fr), Bojer s.n. (K); base of the shoulder (fl), Bouton s.n. in 1864—1865 (K); (fr), ous s.n. in 1831 (P, summit of the e. highes Bouton s.n. (G-D shoulder (fl), Bouton pu MAU 1 307 (MAU); SE slope near the summit, 19 Nov. 1973 (fl), Coode 4125 (K); im > below summit on windward slope, 800 m, 14 978 2 ve poiius (M MA 18 Aug. 1979 (fl), je 291 OCOTTE: mossy forest, sum- mit, 10 June o» (B. Cans uu MAU -- — (MAU, 2 sheets); indigenous vegetation along c Nov 1969 (fr), ed sub MAU 14130 (M MAU. s sheets); summit, ca. 770 m, 1 Dec. 1978 (fl), Lorence 1982 (K MO); (fr), quid 1983 (B, G, K, MAU, MO, P, REU, Z); 19 Dec. 1978 (st), Lorence & Lecordier 2168 (MAU, O, P); 17 Co 1979 (fr), Lorence 2285 (MAU, MO); 29 Jan. 1979 (fr), Lorence 2350 (MAU, MO); 10 Aug. 1979 (fl), Lorence 2895 (MO); (fl), Lorence 2896 (B, AU, MO, P, REU, Z); 29 Jan. 1979 (st), Lor- ence 2964 (MO). Mr. LAGRAVE: dense thicket along crest, 27 Sept. 1969 (fl), Guého sub MAU 14097 (MAU, 2 sheets) dense thicket on N flank, 27 Sept. 1969 (fl), Guého sub MAU 14105 (MAU, 2 sheets); S slope facing Eau Bleue Reservoir, 620 m, 24 May 1979 (st), Lorence & Lecordier 2650 (K, MAU, MO). PLAINE CHAMPAGNE: track from Plaine Champagne to Black River Gorges, 1,400 ft. (ca. 450 m), 4 Feb. 1975 (st), 1985] Coode s.n. (K). WITHOUT PRECISE LOCALITY: (fl), Ayres s.n. (MO, 2 sheets); e Feri sub MAU 1308 (MAU: MAU); M aud sub eets); (fl), a . iJ > 1, 12215 (P); Piae = on a La Coudée, 22 Nov. ; Basse Vallée near 979 e. Lorence & Cadet O); (fl), Lore e a 2727 (MAU, MO); 17 May 1976 (st), Richardson 4152 (K); 17 Feb. 1956 (st), Severin 50 m ca d BLANC: above Bois Blan 2 (fl), Cadet 3721 (REU); 800 m, region of Bébour near Pl. (fr), Bernardi 15171 GRAND ÉTANG: m, 1979 (fl), Lorence & Rolland 2754 (MO); (f), L & Rolland 2755 (MAU : O y May 1976 (fl), Cadet 5400 (REU); valley to Langevin below Plaine des Sables, 1,400 m, 10 May 1976 (fl, fr), (TL-R); Mont the e p d pi 1,300- 1,500 m, 12 July 1943 (m. NE y B of Riviére des Remparts, Vallon ise 20 Oct. 1948 (fl), Rivals s.n. (TL- LAINE DES PA ES: high pla teaux, (fl, fr), Cordemoy s.n. (TL-R); de Montée, A 2 (fl, fr), Rivals s.n. Nov. 19 Feb. 1968 (fr), ipod 1240 (K, MAU), rain forest, Tacamaca, u Dec. 1971 (fr), Bosser 20926 (P); Tak- amaka fores et à cR ca. 800-900 m, 10 Mar. 1979 (fr), Ld Rolland 2531 (MO); ca. 1,000 m, 8 May 1976 (st), epee et al. 4100 (K, MO). WITHOUT PRECISE LOCA pis 2 s.n. (G); (st), m. s.n. (probably Cordemoy) ; (f), Frappier 269 (P); (fr), Frappier 270 (P); Anon. ex Herb. Richard 66 (P); (fl), ; Richard s.n. (P); (st), Anon. sub Herb, Willd. 18080 (B, LORENCE— MONIMIACEAE 77 n.v.; microfiche, MO); *Madagascar" (probab ly Mau- ritius) (fl), Colebrooke s.n. (G); “Rodrigues” (probably Réunion) (fr), Balfour s.n. in Aug.—Dec. 1874 (K). Monimia ovalifolia differs from M. amplexi- caulis and M. rotundifolia in having much spars- er and uniquely short stellate-peltate laminar tri- chomes with the arms united basally for ca. !^— 12 their length (Fig. 9D, E), flowers with much smaller receptacles, and in having tanniferous idioblasts in the petiolar cortex. Although char- acterized by a generally lower altitudinal range, M. ovalifolia occurs sympatrically with M. ro- tundifolia i in many localities in Réunion (e.g., at B. Populations of Monimia ovalifolia from Ré- union tend to have fewer simple and fasciculate trichomes on the stem, petiole, costa, and inflo- rescence. The leaves are larger than in Mauritius plants and more variable in shape, often obtuse or even orbiculate. Furthermore, crushed leaves of the Réunion plants usually produce a faint, citron-like odor not apparent in Mauritian plants, even though ethereal oil cells are present in the mesophyll of both. Intergradation of plants from the two islands is total, however, and formal rec- ognition of infraspecific variants is impossible. Vernacular name. Mapou (Réunion). EXCLUDED SPECIES Monimia lastelliana Baill., Bull. Mens. Soc. Linn. (Paris) 1: 342. 1882; Perk. & Gilg, Pflanzenr. 4, 101: 65. 1901. [= Tambourissa lastelliana (Baill) Drake, see discussion under this species]. Decarydendron Danguy, Bull. Mus. Hist. Nat. (Paris), Ser. 1, 34: 279. 1928; Cavaco, Bull. Soc. Bot. France 105: 38. 1958; Cavaco, Blu- mea Suppl. 4: 28, fig. 1. 1958; Cavaco in Humbert, Fl. Madagascar 80: 8, fig. II:1-8. 1959. TYPE: D. helenae Danguy. Monoecious treelets or small trees 4—8 m tall, the trunk sometimes swollen basally, the new growth velutinous to canescent, the hairs simple or in fascicles of 2. Leaves opposite, petiolate, adaxially glabrescent, abaxially + pubescent es- pecially along costa and major veins, the vena- tion festooned brochidodromous, the margin serrate to dentate; juvenile and sucker leaves un- known. Inflorescence cauliflorous, sometimes produced at the swollen base of the trunk, or 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN D. helenae @ var. helenae O var. stenophyllum A D. lamii A D. perrieri B T. alaticarpa DNE E O T. longicarpa FiGURE 24. Distribution map of Decarydendron and two species of Tambourissa in Madagascar ramiflorous, a short, pubescent unisexual or sex- ually mixed pleiochasium of 5-15 flowers, often terminated by a gynoecious flower, the pedicels short, bracteolate. Androecious flower in bud subglobose to napiform, pubescent, the tepals imbricate, opening gradually, not splitting, at an- thesis shallowly cupuliform to hemispherical, the rim bearing 10-15 large, subequal, broadly ob- tuse, radially arranged tepals in 1—2 series; sta- mens eglandular, 16—60, situated on the + gla- brous inner receptacle surface, filament gitudinally dehiscent, extrorse, the connective small, scarcely or not prolonged; pollen inaper- turate, spheroid to subovoid, the sexine corru- gate to rugulate, the sexinous elements short, fin- ger-like. Gynoecious floral ptacle subglobose [Vor. 72 in bud, expanding gradually, not splitting, at an- thesis broadly obconical to turbinate, + angular, the walls fleshy, the rim bearing 9-15 broadly obtuse, radially arranged tepals, often eventually caducous, the carpels numerous, ca. —1,000, e lining the receptacle, subsessile, clavate, ca. —2 mm long, + polyhedral, the small stylar vum situated laterally in the basal half, the sol- itary ovary unilocular, the small loculus confined to the base of the carpel, containing a single pen- dulous, anatropous ovule, the carpels inter- spersed with + dense, short simple hairs. Fruit- ing receptacle, mature carpels, and chromosome number unknown. The genus was named after the French botan- ical explorer, R. Decary, who collected in Mad- agascar during the early part of the twentieth century Distribution. The three described species of Decarydendron are endemic to the wet evergreen forest zones of central and eastern Madagascar Fig. 24 TAXONOMIC HISTORY OF DECARYDENDRON The genus Decarydendron was described by Danguy (1928) based on the species D. helenae. Thirty years later Cavaco (1958a, 1958b), de- scribed two additional species, D. perrieri and D. lamii, as well as another variety of D. helenae, var. stenophyllum, in preparation for his treat- ment for the “Flore de Madagascar" (1959). Her- barium material is scarce, and many additional flowering and fruiting collections and field ob- servations are required for a meaningful revision of the genus. SYSTEMATIC POSITION OF DECARYDENDRON Decarydendron possesses the greatest constel- lation of primitive characters of any of the Mal- agasy Monimiaceae genera, notably the broad, orm e ally without splitting into segments as in the oth- er genera (Fig. 14E). These features readily distinguish Decarydendron from the other two Madagascan genera which form an increasingly specialized series. Decarydendron resembles He- dycarya from Oceania in a number of respects, including the gynoecious floral morphology. Its mature fruiting receptacle presumably resembles 1985] that of the latter genus in having numerous ses- sile carpels situated on a flat, discoid receptacle. Money et al. (1950) placed Decarydendron in the subfamily Monimioideae with the other Madagascan genera. Hutchinson (1964) placed it in the tribe Hedycaryeae within the same subfamily on the basis of its free, sessile carpels. In redefining the Monimioideae to include only Monimia and Peumus, Thorne (1974) created a new subfamily, the Mollinedioideae, to house the remaining genera formerly included in the Mo- nimioideae. Decarydendron 1s therefore now in- cluded in the Mollinedioideae. As its pollen and vegetative and floral anatomy correspond most closely with the other members of this subfamily, I agree with its placement there KEY TO THE SPECIES OF DECARYDENDRON la. Leaves abaxially velutinous; petioles 3-5 m long; tepals of androecious flowers du ads l. D. perrieri lb. Leaves abaxially glabrate or at most puber- ulent; petioles 10-20 mm long; tepals of an- droecious flowers entire, not crenulate. 2a. Laminar teeth 2-4 mm long; androe- cious flowers with 40-60 stamens ........ 2.D 2b. Laminar teeth 0.5 mm long; androecious flowers with 16-26 stamens ........... 3. D. lamii 1. Decarydendron perrieri Cavaco, Bull. Soc. Bot. France 105: 38. 1958; Cavaco in Humbert, Fl. Madagascar 80: 9, fig. I1:6—8. 1959. TYPE: Madagascar. Tamatave: Analamazaotra forest (near Perinet) (fl), Perrier de la Bathie 10089 (holotype, P). Monoecious treelet or small tree 5-8 m tall, the base of the trunk swollen, the new growth densely velutinous, the mature leafy stems to- entose. Leaves opposite to subopposite, peti- olate; petioles sparsely tomentose, 3-5 mm b 1.5-2 mm; lamina sparsely tomentose adaxially, velutinous abaxially, chartaceous, narrowly el- liptic to narrowly ovate, 80-165 mm by 35-65 mm, the apex acuminate, the base broadly cu- neate to obtuse, the secondary veins 5-7 pairs, making a 55—65? angle with the costa, depressed adaxially, the venation visible to 3? adaxially, to 4° abaxially, finely reticulate, prominent, the margin serrate-dentate with 30—35 pairs of sub- equal, acuminate teeth 1-3 mm long. Inflores- cence cauliflorous at the swollen base of the trunk, sometimes extending up the trunk for some dis- tance, a velutinous pleiochasium of 5-7 flowers, either unisexual of androecious flowers or sex- LORENCE-—MONIMIACEAE 79 ually mixed and terminated by a gynoecious flower, the floral axis 120-220 mm by 3-4 mm, the pedicels 5-17 mm by 1.5-3 mm, basally bracteolate. Androecious flowers smaller than the gynoecious, the receptacle thick, externally ve- lutinous, cupuliform, 7-14 mm diam., the mar- gin bearing 10-18 subequal thick, deltoid-ovate or suborbicular tepals 5-6 mm long by 4-5 mm wide in two series, the external ones wre s the internal ones sinuate; stamens ca. 25, brous, 2-3 mm long, ellipsoid, the lament UI not prolonged, the anther ovoid, occupying 7 total length of the stamen. Gynoecious flower at anthesis externally velutinous, cupuliform, thick, 14-17 mm diam., with two longitudinal external ridges, each terminated by an ovate-acuminate, denticulate bracteole, the tepals subequal, del- toid to ovate, 3-6 mm by 5-7 mm, numbering 12-15, in two whorled series, the outer ones ex- ternally velutinous, the inner ones glabrate; car- pels numerous, ca. 800-1,000, lining the recep- tacle, free, clavate, ca. 1.5 mm long. Fruiting receptacle unknown. Distribution. Endemic to Madagascar (Fig. Habitat. The species is known only from the type locality, where it occurs in Tambourissa/ Weinmannia wet forest at ca. 90 MADAGASCAR. TAMATAVE: Perinet forest, 900 m, 30 Mar. 1974 (fl), Cremers 3017 (MAD). In addition to the characters cited in the key, Decarydendron perrieri diff its congeners y the presence of a series of smaller tepals al- ternate with the larger outer series in the female flowers. Further collections, including fruiting material, and field observations are required to further assess its affinities within the genus. 2. Decarydendron helenae Danguy, Bull. Mus. Nat. Hist. (Paris), Ser. 1, 34: 279. 1928; Ca- vaco in Humbert, Fl. Madagascar 80: 10. 1959. TYPE: Madagascar. Fianarantsoa: for- est at N base of Ivohibé Peak, 20 Sept. 1926 (fl), Decary 5391 (holotype, P). Small monoecious tree 7 m tall and 25 cm diam., the new growth golden tomentose, the mature leafy stems glabrescent, the trichomes simple or in fascicles of 2. Leaves opposite, pet- iolate; petioles 10-18 mm by 1.5-2.5 mm, gla- rate; lamina ovate-elliptic to narrowly obo- vate-elliptic, 150-200 mm -130 mm, chartaceous, adaxially hirsutulous when young, 80 ANNALS OF THE MISSOURI BOTANICAL GARDEN glabrescent, abaxially hirsutulous, velutinous along the costa and veins, the apex acute to short- ly and abruptly acuminate, the base narrowly cuneate to obtuse, the secondary veins 5-8 pairs, making a 50-70? angle with the costa, the ve- nation visible to 3? adaxially, to 4? abaxially, the margin prominently serrate-dentate, the teeth 15— 25 pairs, unequal, acute to acuminate, 2-4 mm long, the glandular tips indurated. Inflorescence probably cauli- or ramiflorous, a finely veluti- nous pleiochasium of 5-7 flowers, unisexual of androecious flowers, or sexually mixed and ter- minated by a gynoecious flower, the floral axis 130-230 mm by 3-4 mm, the pedicels 10-15 mm by 1-1.5 mm, flattened near the junction with the receptacle, basally bracteolate. Androe- a single series of 7-15 unequal, obtuse tepals 3— 8 mm long by 6-10 mm wide, pilose on both surfaces; stamens ca. 40—60, glabrous, ellipsoid, 3-4.5 mm long, the filament ca. 0.5 mm long, the connective thick, not prolonged, the anther - PR mm long. oo flower larger than at broad- ly obconic to cupuliform, 15-35 mm alone by 20- m wide, externally pilose-velutinous, the margin bearing a single series of 8-15 thick, ob- tuse tepals 2-5 mm long by 2-4 mm wide; carpels numerous, up to 300 or more, lining the recep- tacle, clavate, 1.5-2.5 mm long, glabrous, inter- pera with POTONE short, simple trichomes. 1 unkno YV 11. oO r r Distribution. Endemic to Madagascar (Fig. 4) Decarydendron helenae differs from its con- geners in having a combination of adaxially gla- brate or only sparsely puberulent leaves with large marginal teeth, and more numerous stamens (up to 60 per flower). The species, particularly the variety stenophyllum, appears to be most closely allied to D. perrieri. More numerous collections and field studies are required to critically assess its status. Y TO THE VARIETIES OF DECARYDENDRON HELEN. la. Leaves 90-130 mm wide; tepals ca. 15 ...... 2a. var. helenae lb. Leaves 50-75 mm wide; tepals 7-1 . var. stenophyllum ° [VoL. 72 2a. Decarydendron helenae var. helenae. Variety helenae differs primarily in having larger leaves, a more robust inflorescence, and larger flowers than var. stenophyllum. Also, its receptacle lacks the two lateral ridges present in the latter variety. It was named after Helen De- cary, who assisted her husband in his botanical research. Habitat. Known only from the type, which was collected at the base of Ivohibé Peak in the Andringitra Massif in southern central Mada- gascar, it probably occurs in montane evergreen wet forest, most likely in the upper limits of the Tambourissa/ Weinmannia zone (Fig. 24). 2b. Decarydendron helenae var. stenophyllum Cavaco, Bull. Soc. Bot. France 105: 39. 1958; Cavaco in Humbert, Fl. Madagascar 80: 10. 1959. TYPE: Madagascar. Tulear: upper ba- sin of the Mandrare River (SE), pass and summit of Marosoui, 1,000-1,400 m, forest on gneissic laterite, 14-15 Nov. 1928 (fl), Humbert 6613 (holotype, P; isotypes, K, MAD, US) n addition to its narrower leaves, variety stenophyllum has a less robust inflorescence, smaller flowers with fewer tepals, and floral re- ceptacles with two prominent, lateral ridges end- ing in small bracteoles. Whether these differences are constant or merely represent stages within a range of variat n (type [o however, geographically RUM bs about 250 km Distribution. Endemic to Madagascar (Fig. Habitat. Variety stenophyllum is known only from the southeastern part of Madagascar, prob- ably in the Tambourissa/ Weinmannia wet forest zone. 3. Decarydendron lamii Cavaco, Blumea Suppl. 4: 28, fi 1938 (fl), Lam & Meeuse 5835 (holotype, presumably BO, n.v., photo, P; isotype, P) Small monoecious understory tree 5 m tall, the new growth fulvous-velutinous, the mature 1985] LORENCE-— MONIMIACEAE 81 leafy stems villous, 3-5 mm diam. Leaves op- posite, e petioles 10-20 mm by 1.5-2 mm, velutinous; lamina chartaceous, narrowly elliptic to dede obovate-elliptic, 100-190 mm by 40-72 mm, the apex abruptly short acumi- nate, the base cuneate, the adaxial surface gla- brate, the abaxial surface appressed pilose, es- pecially on the costa and veins, the secondary veins 10-12 pairs, making a 65-75? angle with the costa, raised on both surfaces, the venation visible to 3? adaxially and to 4? abaxially, the margin plane, finely dentate, the teeth 40—50 pairs, 0.5 mm long, glandular tipped. Inflorescence cauliflorous from meristematic swellings on the trunk or ramiflorous, a pleiochasium of 8-15 flowers, unisexual of androecious flowers or sex- ually mixed and terminated by a single gynoe- cious flower, the floral axis 100-220 mm by 2 mm, densely fulvous-velutinous. Androecious flowers smaller than the gynoecious, at anthesis shallowly cupuliform to disciform, open, 12-16 mm diam. by 5-8 mm long, externally finely strigose-sericeous, the margin bearing 10-12 en- tire, obtuse, ovate to ligulate tepals 2-3 mm wide and long, in two series, the pedicel 6-25 mm by 1 mm, basally bracteolate, the bracteole to 4 mm long; stamens 16-26, ellipsoid to obovoid, 1.8- 3 mm long by 1.5-1.8 mm wide, the filament distinct, short, the anther occupying 74—% total length of the stamen, the connective thick, not prolonged, the internal receptacle surface gla- brous, purple. Gynoecious flower at anthesis ob- conical, 30 mm diam. by 24 mm long, externally finely velutinous, the carpels numerous (proba- bly ca. 500 or more), lining the receptacle. Fruit- ing receptacle and mature carpels unknown. Distribution. Endemic to Madagascar (Fig. 4). Habitat. The species is known from the low- er montane wet forest zone of Myristicaceae/An- thostema in the eastern region, below NÁ e c ga, Bassin of Androranga River, 700 m, 18 Nov. 1950 (f), Capuron 827-SF (P). TAMATAVE: Andapa District, basin, E of Ambalamansy II, 450-80 on gneissic laterite, 28 Nov. 1948 (st), Humbert & Ca- puron 22154 (MO In addition to the characters cited in the key, Si oiii lamii differs from its congeners urpl e within. Like the other species in the genus, it is poorly known Ephippiandra Decne., Ann. Sci. Nat. (Paris), Ser. 4, 9: 278, pl. 7. 1858; A.DC., Prodr. 16(2): 662. 1868; Baill, Hist. Pl. ed. 1, 1: 304. 1869; Perk., Engl. Bot. Jahrb. 25(4—5): 551, t. 5: 1-5. 1898; Perk. & Gilg, Pflanzenr. 4, 101: 50, fig. 9. 1901; Perk., Pflanzenr. 4, 101 (Nachtr.): 15. 1911; Perk., Gatt. Monim. 32, fig. 19. 1925; Cavaco, Bull. Soc. Bot. France 104: 610. 1957; Cavaco in Humbert, Fl. Madagascar 80: 4. 1959; Hutch., Gen. Fl. Pl. 1: 117. 1964. TYPE: E. myrtoidea Decne. F pear Bull. Mus. Hist. die Pe d “apos synon. nov. TY | Dangu Monoecious shrubs, treelets or trees 10-25 m glabrous or + pubescent abaxially, apeti along the costa and veins, the venation festooned brochidodromous, semicraspedodromous or craspedodromous, the margin entire or sparsely dentate apically with acute, glandular teeth, or sinuate-dentate with rounded, apically depressed glandular teeth; juvenile and sucker leaves un- known. Inflorescence axillary, terminal, or ram- iflorous on leafless stems, a short, 5-flowered uni- sexual pleiochasium or a 3-5-flowered sexually mixed dichasium, the gynoecious flower solitary and terminal, maturing first, the androecious flowers lateral, or the flowers solitary and axil- lary, the pedicels + long and slender, basally bracteolate. Androecious flower in bud globose to ellipsoid, bearing 2—4(—5) small tepals apical- ly, at anthesis splitting deeply into 4(-5) + reg- m reflexing, the apex often like tepals internally; stamens eglandular, ca. 50, situated irregularly or in several series on the glabrous to puberulent internal receptacle sur- face, ligulate to shortly deltoid, the filament broad, short or sessile, the anther tetrasporangiate, the 2 loculi lateral and separate or connivent or con- fluent apically, longitudinally dehiscent, ex- trorse; pollen inaperturate, spheroid to ovoid, the exine thick, the sexine granulate with few to many, small to large scattered verrucae or gemmae. e mar u small, decussate deli tepals; at anthesis ex- panding gradually, not splitting, broadly obconic to discoid, the torus flat to concave, bearing ca. 82 ANNALS OF THE MISSOURI BOTANICAL GARDEN 15-120 free, short sessile, 4—6-sided columnar carpels ca. 1-1.5 mm long, the apex flat, the small stylar canal lateral, medial, the ovary unilocular, the solitary ovule anatropous, situated in a small locule near the base of the carpel, the carpel interspersed with short, simple hairs. Fruiting receptacle discoid, enlarged, fleshy and red when fresh, the flat to convex surface bearing ca. 10- 75 free carpels, each surrounded basally by a cupule for '4—'4 its length; mature carpels ovoid to ellipsoid, 7-12 mm long, the thin fleshy me- socarp ripening black, the endocarp thin, horny, pale brown, the surface scrobiculate, the endo- sperm fleshy. Chromosome number unkno ue Distribution. All six species are endemic to The genus ranges from lower mon- tane wet and cloud forest to high altitude mon- tane sclerophyllous forest and ericoid forma- tions TAXONOMIC HISTORY OF EPHIPPIANDRA The genus Ephippiandra, described by De- caisne in 1858 based on E. myrtoidea, was named in reference to the low, saddle-shaped anthers pne pc of the species. A second species, described by Perkins (1911) as Tambourissa mi- cro E lla, was subsequently transferred to Ephippiandra by Cavaco (1957a). In the same year Cavaco (1957c) described a third species and established two sections in the genus based on the presence (section Staminodia) or absence (section Ephippiandra) of fleshy tepals on the inner tips of the receptacle lobes, which he re- ferred to as “‘staminodes.” Danguy described Hedycaryopsis in 1928 based on H. madagascariensis, and Cavaco (1958c) subsequently described three additional species preceding his treatment for “Flore de Madagas- 1959) 39 car GENERIC DELIMITATIONS AND SYSTEMATIC POSITION OF EPHIPPIANDRA Danguy (1928) established Hedycaryopsis as a segregate genus primarily on the basis of its gy- noecious flowers, which he felt approached those of Hedycarya Forst. (syn. Carnegieodoxa Perk.) from Oceania. The discoid to broadly obconic gynoecious flowers of Hedycaryopsis, like those of Ephippiandra, are usually bordered by several pairs of minute, decussate deltoid tepals and the torus bears few to numerous free, sessile poly- [VoL. 72 hedral-columnar carpels on its surface (15-38 in Ephippiandra, 22-120 in Hedycaryopsis, Fig. 14F, G). Furthermore, carpels of both possess oblique lateral stylar canals (Fig. 13C, D). Dan- guy apparently never examined gynoecious flow- ering material of Ephippiandra, because he makes no mention of it m his dispussion (the type of E. d fruit) Clearly, AR floral morphology does not support segregation of Hedycaryopsis from Ephippiandra. Furthermore, fruiting receptacles of both gen- era are morphologically identical. Mature fruit- ing receptacles in both consist of a fleshy, discoid torus, red when ripe, bearing few to many ovoid to ellipsoid carpels surrounded basally by cu- pules (Fig. 14H). Carpels of both have a thin mesocarp, green when fresh, overlying the thin, horny endocarp. Understandably, fruiting char- acters have never been used to distinguish the genera. Although Danguy (1928) noted that androe- cious flowers of Hedycaryopsis scarcely differed from those of Tambourissa, Cavaco (1957c, 1959) felt that stamens were a stable key character sep- arating Ephippiandra from Hedycaryopsis and Tambourissa. He stated that in Ephippiandra the stamens were sessile and broader than long (‘‘dé- c esed aryopsis which had separate, lateral loculi and an apically prolonged connective. He ig- ee or at least failed to mention, however, that in species such as E. myrtoidea the loculi may be both connivent as well as confluent in the same flower. Tambourissa displays the entire range of conditions and the loculi may be sep- arate, connivent, or confluent not only apically but basally as well in various species. Two or more of these conditions may occur in a single androecious flower of T. cordifolia (Fig. 42C—E). ther species such as 7. purpurea possess broad, sessile stamens with a single loculus similar to those of some Ephippiandra species. Further- more, the number of stamens is relatively small in both and overlaps (10-36 in Ephippiandra; 9-26 in Hedycaryopsis), as does the presence or absence of tepals on the Eu of the bs ceptacle lobes. And characters are us j insufficient for generic segregatio Examination s the pollen of two species each of Ephippiandra and Hedycaryopsis (Lorence, 1980; Lorence et al., 1984) has revealed that their 1985] LORENCE-— MONIMIACEAE 83 sexinous patterns correspond much more closely to each other than to any of the other Malagasy genera. Pollen of E. microphylla and E. myrtoi- dea both have thick, granulate to verrucate sex- ines with larger, scattered gemmae and folds oc- curring in the latter. In H. perrieri and H. tsaratanensis the sexine is also thick and gran- ulate with large, dome-shaped verrucae and gem- mae. Sexinous patterns in these two genera differ primarily in the size of the processes, but cor- respond much more closely to each other than to the remaining Malagasy genera. I do not feel that pollen supports separation ofthe two genera. Cavaco (1959) further distinguished these gen- era on the basis of inflorescence structure: di- chasial with a central gynoecious flower and lat- eral (rarely solitary) androecious flowers in Hedycaryopsis (Fig. 14F, G); solitary or pleiocha- sial in Ephippiandra. The only species suppos- edly having pleiochasia, Ephippiandra capuron- ii, actually belongs in Tambourissa (see below). Collections of E. myrtoidea (Anon. sub Herb. ORSTOM 3366, MAD; Anon. sub Herb. Alaotra 300, MAD) possess sexually mixed dichasia, however, thus undermining the validity of this character in separating the genera. Furthermore, an even greater range of inflorescence types oc- curs in the otherwise very cohesive genus Tam- bourissa. Leaves of the various Hedycaryopsis species are extremely diverse and display a striking array of venation patterns (see section on Leaf Archi- tecture). For example, leaves of the type species, H. madagascariensis, are highly specialized and generally characterized by open craspedodro- mous venation ending in sunken teeth at the sin- uate-dentate margin (Fig. 6F), a pattern appar- ently derived from the brochidodromous condition (R. Keating, pers. comm.). In this re- spect, they differ strikingly from all other known Monimiaceae genera and may have influenced Danguy (1928) to describe the genus as new. Ve- nation in two other species, H. capuronii (= Ephippiandra domatiata; Fig. 6C, D) and H. tsaratanensis (Fig. 6E), as well as some collec- tions of H. madagascariensis, is wansitional be- a nimia cluding Ephippiandra myrtoidea (Fig. 6A) and E. microphylla. This intergrading range of ve- nation patterns provides another criterion for uniting the two genera. Because there seems to be no basis for contin- ued segregation of Hedycaryopsis, I propose that it be reduced to synonymy under Ephippiandra, which would thus encompass six species, neces- sitating one new name and three new combina- tions given below Among the Ma la gasy Monimiaceae belonging to the Mollinedioideae, Ephippiandra occupies a position intermediate between Decarydendron and Tambourissa in terms of floral morpholog- ical specialization. The strong androecious/gy- noecious floral dimorphism, highly reduced te- pals, sessile carpels enveloped em by cupules fruit, l in bud and splitting open at anthesis, and tendency towards short, sessile stamens with confluent loculi are all specializations over Decarydendron. Female floral construction in Ephippiandra is not so ad- vanced as in Tambourissa, however, where the closed gynoecious receptacle splits partly open at anthesis and the carpels are inferior and syn- carpous. 8 KEY TO THE SPECIES OF EPHIPPIANDRA la. Leaf margin sinuate-dentate with rounded, apically + depressed teeth. 2a. Abaxial laminar surface densely ferru- gineous tomentose; floral pedicel 25-50 mm long, ferrugineous tomentose 1. E. tsaratanensis . Abaxial laminar surface at most grayis or yellowish puberulent along costa and veins; floral pedicel 8-15 mm long, gray- ish canescent or villous N c ee Ib. Leaf margin entire or at most A with mel. of acute, deltoid teeth in apical port 3a. Basal pair of secondary veins ee up to or beyond middle portion of lam- ina before joining next pair above; bar- bate domatia present PEE in sec- E. domatiata dary veins pcd next pair above before attaining middle of lamina; domatia absent abaxially in sec- ondary vein axils. 4a. Apex of lamina retuse, folded down- ward into a thickened flap w c = 9 Š a E ^a ° Sa $ e o 2 . E. perrieri 4b. Apex of lamina shortly acuminate to acute or rounded, not retuse or a f each androe- bearing 1(—2) scale- y tepals . microphylla 5b. Inner portion of ape oecious floral receptacle lobes lacking scale-like tepals ......... 5. E. myrtoidea 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN @ E. domatiata "65 \ E E. madagascariensis o Y 4 E. tsaratanensis N FiGuRE 25. Dis a map of some Ephippian- dra species in Madagas p . Ephippiandra tsaratanensis (Cavaco) Lo- rence, comb. nov. Hedycaryopsis tsarata- nensis Cavaco, Bull. Soc. Bot. France 105: 41. 1958; Cavaco in Humbert, Fl. Mada- gascar 80: 14, fig. III:1—5. 1959. TYPE: Mad- agascar. Diego Suarez: Tsaratanana Massif, forest with herbaceous undergrowth, 1,800 m (fl), Perrier de la Báthie 15302 (holotype, P) Monecious tree 10-25 m tall, the new growth densely ferrugineous tomentose, the mature leafy stems sparsely tomentose. Leaves opposite to subopposite, petiolate; petioles 8-15 mm by 2.5- 3.5 mm, ferrugineous tomentose; lamina sub- coriaceous, the apex rounded, obtuse, the tip de- pressed, the base broadly cuneate, rounded, trun- cate or rarely subcordate, adaxially pubescent, glabrescent, abaxially densely ferrugineous to- [Vor. 72 mentose, the secondary veins 5-8 pairs, making a 50-65? angle with the costa, the venation cras- pedodromous or partly brochidodromous, visi- ble to 3° adaxially, to 4° abaxially, the margin slightly revolute, sinuate with 5—8 pairs of broad- ly rounded, apically depressed teeth. Inflores- cence axillary, ferrugineous velutinous, a 3-5- er terminal and maturing first, the peduncle 15— 30 mm by 2-3 mm, apically diatated, the ie cels each subtended by a linear bracteole 4—6 m long. Androecious flower in bud ellipsoid to die. bose, 4-6 mm diam., ferrugineous tomentose, . near the apex 4 linear-ovate tepals 3-4 mm g, the pedicel 25-50 mm by 1 mm; at ea deep. 4-fid, 12215 mm diam., the lobes spreading flat, each bearing a broad tepal apically within; stamens 18—20, ovate to ligulate, 2-2.5 mm long by 1-1.5 mm wide, slightly incurved, the filament thick, short or sessile, the loculi lat- eral, separate, occupying almost entire length of the stamen, the connective slightly prolonged, apiculate. Gynoecious flower at anthesis broadly obconical, concave, externally ferrugineous to- m the pedicel 32-37 mm long; carpels 40-70, lining the receptacle, columnar, 4—5-sided, 1-1.5 mm long. Mature fruiting receptacle red when fresh, drying dark brown, irregularly discoid, convex, 60-85 mm diam., the ground tissue containing numerous stone cells, bearing (4—)40—70 free car- pels surrounded basally by cupules for '4—'/ their length, the pedicel and peduncle ca. 45 mm long by 3-4 mm diam.; mature carpels ovoid-ellip- soid, 10-12 mm long by 6-7 mm diam., green when fresh, drying brown, the mesocarp thin, fleshy, the endocarp whitish, 0.2 mm thick, the surface scrobiculate. Distribution. Endemic to Madagascar (Fig. ). Habitat. The species is known only from the type locality on the Tsaratanana massif in north central Madagascar where it occurs from ca. 1,800 to 2,500 m, presumably in montane sclerophyl- lous cloud forest. jagen DIEGO SUAREZ: Tsaratanana massif, trail up S ridge 2,000-2,500 m; iiim cloud forest, 9 May 1974 (fi, fr), Gentry 11605 (MO). The species is most closely allied to Ephip- piandra madagascariensis, from which it differs 1985] by its suborbicular, abaxially ferrugineous to- mentose leaves with mixed craspedodromous/ brochidodromous venation, its more densely fer- rugineous tomentose parts, more robust inflo- rescence with longer pedicels subtended by long, linear bracteoles, and by its androecious buds bearing 4 long, linear-ovate tepals. N . Ephippiandra madagascariensis (Danguy) Lorence, comb. nov. Hedycaryopsis mada- gascariensis Danguy, Bull. Mus. Hist. Nat. (Paris), 1, 34: 278. 1928; Cavaco in Humbert, Fl. Madagascar 80: 14, fig. IV:1- 6. 1959. TYPE: Madagascar. Diego Suarez: Antsalaky forest, 700-1,000 m, 17 Oct. 1927 (fl, fr), Ursch 66 (lectotype, P, here desig- nated). Monoecious tree 10-25 m tall and 50 cm diam., the new growth grayish to yellowish pubescent or canescent, the mature leafy stems 3—5 mm nt. Leaves opposite to subop- ed, the tip sparsely puberulent, glabrescent, the abaxial sur- face puberulent, especially along the costa and veins, the secondary veins 3-5 pairs, making a 40-50? angle with the costa, depressed adaxially, prominent abaxially, dichotomizing and running into the marginal teeth, the venation craspedod- romous, rarely semicraspedodromous, visible to 2? adaxially, to 3? abaxially, the margin revolute, sinuate with 2-7 pairs of broadly obtuse, apically depressed teeth. Inflorescence axillary, a sexually mixed, 3-5-flowered dichasium, the 2-4 an- droecious flowers lateral, subtending the termi- nal gynoecious flower, or the gynoecious flower rarely solitary, the peduncle grayish pubescent, 15-30 mm by 1-1.5 mm, apically dilated. An- droecious flower in bud globose, 3-4 mm diam., the canescent pedicel 8-20 mm by 0.6-0.8 mm, each subtended by a naviculate bracteole 2 mm long, the apex bearing 3-4 obtuse, puberulent tepals 1-2 mm long; at anthesis deeply 4(—5)-fid, ca. 10 mm diam., the lobes spreading flat; sta- mens 35-50, ligulate-subulate, 1.5-2 mm long by 0.7 mm wide, the filament thick, subsessile, the loculi lateral, separate, occupying almost en- tire length of the stamen, the connective thick, prolonged, obtuse, the internal surface glabrous. Gynoecious flower at anthesis discoid, LORENCE— MONIMIACEAE 85 flat, 10-15 mm diam. by 1.5 mm thick, exter- nally canescent, the margin bearing 8 or 16 mi- nute, deltoid tepals 0.5 mm long, the pedicel canescent, 6-15 mm by 1.5 mm; carpels ca. 140- 150, lining the receptacle, sessile, columnar, 5— 6-sided, 0.8-1 mm long by 0.6-0.8 mm wide, the apex flat, interspersed with numerous short, simple hairs, the internal receptacle surface with mucilaginous secretion. Mature fruiting recep- tacle swollen, everted, 35-60 mm diam., bearing 30-50 carpels, the pedicel and peduncle ca. 25 mm by 3 mm; carpels ovoid-ellipsoid, 8-10 mm long by 6-7 mm diam., the basal 4—!^ enveloped by a cupule, the mesocarp reddish, fleshy, ca. 0.5 mm thick, the endocarp brown, foveolate-retic- ulate. Distribution. Endemic to Madagascar (Fig. ). Habitat. Ephippiandra madagascariensis appears to be primarily restricted to the Tam- bourissa/Weinmannia wet forest zone (ca. 700- 1,000 m) in northern and eastern Madagascar. MADAGASCAR. DIEGO SUAREZ: Montagne d’Ambre, S of sae Forestry Station by Petit Lac and above on the summit, 1,100-1,200 m, 22 Dec. 1967 (fl), Hernani 1 1977 (K, MO); Montagne d'Ambre Na- wet forest, ca. 1,200 m, 23 May MO); quon d'Ambre, y; 1,100 m, 7 Dec. 1952 (st), Schedl 170 a srg (st), ’ Schedl 177 (MAD). TAMATAVE: Bean rinet, 12 Sept. 1948 (fl), Anon. sub Service Forestier 15 96-SF (MAD); Analamazaotra forest sisi 1919 (fr), Thouvenot s.n. (P, syntype of He ukan psis madagascariensis); road from Ano- sibé to Mor manga, 9 May 1969 (st), Anon. sub Service Forestier 26852. SF (MO, P). bagni Ahimia- ana , Andramasina, forest SE of Lake Tsia- oa ee god 1961 (fl), Capuron po SF (MO). The species is most closely allied to Ephip- piandra tsaratanensis, differing by its obovate, abaxially glabrous or sparsely puberulent leaves with nen dichot 4. í ly partly brochidodromous) venation, its yellow- ish to grayish canescent pubescence, and less ro- bust inflorescence with shorter pedicels, bracteoles, and androecious tepals. It yields a yellowish wood said to be good for construction. Vernacular names. Ambora, Amborabé, Tambonaika (Madagascar). 3. Ephippiandra domatiata Lorence, nom. nov. Hedycaryopsis capuronii Cavaco, Bull. Soc. Bot. France 105: 40. 1958; Cavaco in Hum- 86 ANNALS OF THE MISSOURI BOTANICAL GARDEN bert, Fl. Madagascar 80: 13. 1959 non Ephippiandra capuronii Cavaco, Bull. Soc. Bot. France 104: 610. 1957. TYPE: Mada- gascar. Diego Suarez: Beanjada Massif (N of the Masoala Peninsula), ridge at ca. 1,100 m, 30 Dec. 1953 (fl), Capuron 8828-SF (ho- lotype, P). Monoecious shrub, the new growth with sparse, scattered hairs, glabrescent, the mature leafy stems 1-2 mm diam., becoming lenticellate. Leaves opposite, petiolate; petioles glabrous, 6— 10 mm by 0.7-1 mm; lamina glabrous, charta- ceous, obovate-elliptic to rhombic, 25-40 mm by 14-25 mm, the apex obtuse to retuse, often apiculate, the base cuneate, the secondary veins 2-3 pairs, the strong basal pair making a 40—45? angle with the costa, attaining top half of the lamina before uniting brochidodromously with the next pair, adaxially depressed, abaxially prominent, the axils of the basal veins domatiate and barbate abaxially, the venation visible to 2? adaxially and to 3-4? abaxially, the margin revo- lute, with 1-2(—3) pairs of minute teeth in the apical '^-!^. Inflorescence axillary, a 3-flowered dichasium, the terminal gynoecious flower sub- tended by 2 androecious flowers, or the male flowers solitary or in cymes of 3, the peduncle glabrous, (4-)12-15 mm by 0.5 mm, with 2 basal bracteoles. Androecious flower in bud globose, 2.5-3 mm diam., glabrous, the apex with 2 pairs of broadly obtuse tepals 1.5-2 mm wide, the pedicel 7-8 mm by 0.5 mm, subtended by a brac- teole 1-1.5 mm long; at anthesis deeply 4-fid, 5- 6 mm diam., the lobes spreading flat, each bear- ing apically 2 thin, broadly obtuse tepals inter- nally; stamens 9-10, ovoid-subulate, 1-1.5 mm long by 0.6-0.8 mm wide, the filament short, distinct, the loculi lateral, separate, occupying ca. 3⁄4 total length of the stamen, the connective slightly prolonged, apiculate. Gynoecious flower at anthesis discoid, flat, 4.5-5 mm diam. by 1.5 mm thick, externally with a few scattered hairs, the margin entire or with a few minute, deltoid teeth, the pedicel glabrous, 9-19 mm by 0.5 mm, occasionally basally bracteolate; carpels 24-25, sessile, columnar, 0.8 mm long by 0.6-0.7 mm wide, 5—6-angled, interspersed with tufts of short, simple hairs. Fruiting receptacle unknown. Distribution. Endemic to Madagascar (Fig. Habitat. The species is known only from the type locality, presumably in Tambourissa/Wein- mannia wet forest at ca. 1,100 m. [Vor. 72 Ephippiandra domatiata is readily distin- guishable from its congeners by its glabrous in- florescence and stems, and particularly by its leaves with abaxially domatiate, barbate second- ary vein axils. Further collections, especially fruiting material, and field observations are greatly desired. 4. Ephippiandra perrieri (Cavaco) Lorence, comb. nov. Hedycaryopsis perrieri Cavaco, Bull. Soc. Bot. France 105: 40. 1958; Cavaco in Humbert, Fl. Madagascar 80: 16. 1959. TYPE: Madagascar. Diego Suarez: Tsaratan- ana massif, forest with herbaceous under- story, 2,000 m, Apr. 1929 (fl, fr), Perrier de la Báthie 16249 (holotype, P). Monoecious shrub, treelet or tree 12-15 m tall, berulent, glabrescent, often with large, elliptic lenticels. Leaves opposite to subopposite, peti- olate; petioles 5-11 mm by 1 mm, finely puber- ulent, glabrescent; lamina chartaceous to sub- coriaceous, glabrous, elliptic to obovate or obcordate, 25-55 mm by 15-35 mm, the apex obtuse, rounded, the tip retuse or emarginate, folded under into a thickened flap, the base nar- rowly cuneate, the secondary veins 3-4 pairs, making a 45-55? angle with the costa, obscure adaxially, prominent abaxially, the venation vis- ible to 3? adaxially and to 4? abaxially, the margin entire, + revolute. Inflorescence axillary, a 3-flowered, sexually mixed dichasium, the gy- noecious flower terminal, subtended by a pair of androecious flowers, or the androecious flowers solitary, axillary, the peduncle 10-15 mm by 1 mm, finely puberulent, subtended by several small, deltoid bracteoles. Androecious flower in bud globose, 3 mm diam., with scattered ap- pressed hairs, the apex bearing 2 obtuse to subu- late, ciliate tepals 1-1.5 mm long, the pedicel 8— 20 mm by 0.7 mm, often with 1-2 minute brac- teoles; at anthesis deeply 4—5-fid, 9-10 mm diam., the lobes spreading flat or recurving slightly, in- ternally glabrous, each terminated by a broad, retuse or bifid tepal internally; stamens 15-20, recurved, 2-2.5 mm long, the filament slender, 1-1.5 mm long, the anther ovoid-ellipsoid, 1 mm long, the loculi lateral, separate, the connective prolonged, apiculate. Gynoecious flower on a pedicel 12-20 mm by 1 mm, bracteolate basally or medially, the receptacle open in bud; at an- thesis flat, discoid, 4-6 mm diam. by 2 mm thick, 1985] externally finely gray sericeous, the margin bear- ing 4-8(-16) minute, deltoid tepals ca. 0.5 mm long; carpels 20-25, bluntly columnar, 1 mm long by 0.6-0.8 mm wide, 5—6-angled, covering the torus, interspersed with short, simple hairs. Mature fruiting receptacle swollen, 15-20 mm diam., usually with 2—4 carpels, the pedicel and peduncle 25-45 mm by 1.5 mm; carpels ovoid, 8-9 mm by 6-7 mm, the basal '4—'^ enclosed by a raised cupule, the mesocarp thin, fleshy, the endocarp light brown, striate-reticulate. Distribution. Endemic to Madagascar (Fig. ). Habitat. The species is restricted to sclero- phyllous montane cloud forest and ericoid for- mations on Madagascar’s highest mountains, the Marojejy and Tsaratanana massifs, from ca. 2,000 to 2,650 m. MADAGASCAR. DIEGO SUAREZ: Marojezy (Marojejy) assif, summit, 29 Mar. 1949 (fl, fr), Anon. sub Herb Alaotra 3479 (MAD, 3 sheets); Marojejy Natural Re- serve, sclerophyllous forest, 2,000 m, 27 Nov. 1972 (f), Guillaumet the sum- mit of Marojejy, 2 .000—2, 137 m, Dec. 1972 (fl), Morat 4035 (MAD, P); Tsaratanana massif, 2,200-2,650 m, Nov. 1966 (fl), Capuron 25000-SF (MO, P); > 100 m, Nov. 1966 (fr), Morat 2268 ( anisam- birano, 2,500 m, Nov. 1966 (fl), Morat 2332 (MAD); (fr), Morat 2333 (MAD). Ephippiandra perrieri most closely resembles E. domatiata, but its leaves have entire margins, a more deeply retuse or emarginate apex which is thickened and folded under into a flap, lack domatia, and the inflorescence is sparsely pu- bescent. In sheltered forest E. perrieri becomes a tree, but takes on a shrubby habit on exposed slopes. 5. Ephippiandra myrtoidea Decne., Ann. Sci. Nat. (Paris), Ser. 4, 9: 278, pl. 7. 1858; A.DC., Prodr. 16(2): 662. 1868; Baill., Hist. Pl. ed. 1, 1: 304. 1869; Perk., Engl. Bot. Jahrb. 25(4— 5): 551, t. 5:1-5. 1898; Perk. & Gilg, Pflan- zenr. 4, 101: 51, fig. 9. 1901; Perk., Pflan- zenr. 4, 101 (Nachtr.): 15. 1911; Perk., Gatt. Monim. 32, fig. 19. 1925; Cavaco in Hum- bert, Fl. Madagascar 80: 6, fig. 1:7-9. 1959. Mollinedia myrtoidea (Decne.) Drake in Grandidier, Hist. Phys. Madagascar 1(1): 21. 1902. TYPE: Madagascar. Tananarive: vicin- ity of Tananarive (fl), Goudot s.n. in 1840 (holotype, P) Monoecious treelet 3—4 m tall, densely branch- LORENCE—MONIMIACEAE 87 Ú Bi E. microphylla @ E. myrtoidea A E. perrieri E 26. = map of some Ephippian- d ils in Madaga ing, the new growth sparsely appressed pilose, the mature leafy stems 1.5—2 mm diam., shortly velutinous-tomentose, the hairs pale yellowish brown. Leaves opposite, shortly petiolate; peti- oles 1-5 mm by 1 mm, velutinous-tomentose; lamina chartaceous to subcoriaceous, ovate to elliptic, 12230 mm by 8-22 mm, the apex acute to abruptly short acuminate, the tip thickened, the base cuneate to obtuse or rounded, adaxially with a few appressed hairs when young, abaxially with hairs along the costa and margin, the sec- ondary veins 3-6 pairs, making a 65-85? angle with the costa, the venation obscure adaxially, visible to 2? abaxially, the margin entire, pilose, slightly revolute. Inflorescence axillary or ter- minal, a Aun 5- flowered dichasium with a single subtended by 2-4 an- droecious flowers, or the flowers axillary, solitary 88 ANNALS OF THE MISSOURI BOTANICAL GARDEN or in cymes of 2-3, the peduncle 3-4 mm by 1 mm, pubescent, basally bracteolate. Androe- cious flower in bud globose, 2-4 mm diam., with scattered hairs, the apex apiculate with 1—2 pairs of small, obtuse tepals to 1 mm wide, the pedicel 2.5-20 mm by 0.6 mm, bracteolate; at anthesis deeply 4-fid, 6-8 mm diam., the lobes spreading flat, each usually terminated by a small, obtuse tepal externally; stamens 10-13, 2-3 per lobe, sessile, obtuse, 1 mm long by 1.5-2.5 mm wide, the loculi connivent or usually confluent apically into a single crescentiform loculus, the filament broad, short or sessile, the inner receptacle sur- face glabrous. Gynoecious flower on a pedicel 6— 25 mm by 0.8 mm; at anthesis the receptacle discoid, flat or + concave, 6-8 mm diam. by 1 mm thick, the surface crowded with 25-30 ses- sile, 4—6-sided columnar carpels 0.8-1 mm long nt 9 mm "ade, interspersed with short, 1 1 ter minal, the torus swollen, convex, 15- 301 mm diam., red when fresh, drying black, the surface sparsely pubescent, bearing (1—)3-14 carpels, the pedicel and peduncle 25-50 mm by 1.5-2.5 mm; carpels ovoid to ellipsoid, 8-9 mm by 5-6 mm, the basal 15—1⁄2 surrounded by a raised cupule, the meso- carp black when ripe, drying blackish, thin, fleshy, the endocarp thin, brown, the surface finely scrobiculate. Distribution. Endemic to Madagascar (Fig. 6). pau The species appears to be restricted to the Tambourissa/ Weinmannia montane sclerophyllous forest and high altitude ericoid zones of eastern and central Madagascar, from ca. 1,300 to 2,000 m. — ae a s high valley of the Rienana E 1,400 m PROVINCE UNCERTAIN: Manijarivol m, 30 Oct. 1970 (fl), Anon. sub Herb. (MAD); Central Madagascar (fl), dpi 1263 (K); (fr), Baron 1355 (K); Andrangaloatra 1881 (K); Andohakelo Massif (R.N gu 1 800-1 ,850 m, Jan. 1974 (fl, fr), Morat 4430 (P); Morat 4434 (P); Morat 4480 (P). WITHOUT PRECISE LOCALITY: (fl), Anon. sub Herb. Alaotra 300 (MAD). The species is virtually indistinguishable from Ephippiandra microphylla except for its androe- cious flowers which have fewer stamens and lack tepals on the inner apical portion of the recep- tacle lobes. Pedicel length and inflorescence type intergrade, and further collections and field stud- les are necessary to ascertain whether the taxa [Vor. 72 are distinct or merely represent variations of a single, polymorphic species. 6. Ephippiandra microphylla (Perk.) Cavaco, Kew Bull. 12: 228. 1957; Cavaco in Hum- bert, Fl. Madagascar 80: 7, fig. III:6—9. 1959. Tambourissa microphylla Perk., Pflanzenr. 4, 101 (Nachtr.): 41. 1911. TYPE: Madagas- car. Province uncertain: Ambohimitombo forest (Tanala), T p? m, 22 Dec. 1894 (fr), Forsyth- Major 326 (holotype, K). Monoecious treelet or small tree 2-3 m tall, the new growth sparsely pilose to yellowish ve- lutinous-tomentose, the mature leafy stems to- mentose or glabrate, 2-3 mm diam. Leaves op- posite to subopposite, dines petioles pilose to tomentose, 2-5 m mm; lamina chartaceous, ovate, in or oblong, 20-50 mm by 10-25 mm, the apex acute to abruptly acu- minate, the base cuneate to broadly cuneate or obtuse, the secondary veins 4-6 pairs, making a 60-75? angle with the costa, the venation scarcely visible to 2? adaxially, visible to 3? abaxially, the surfaces glabrous, the margin slightly revolute, entire or frequently bearing a pair of small, acute glandular teeth in the apical portion. Inflores- cence axillary or terminal, the flowers solitary. Androecious flowers axillary, in bud globose to obovoid, 3-4 mm diam., subglabrous, the s ga bearing 4 obtuse tepals, the pedicel 12—27 m by 0.5 mm, basally O ase at anthesis deeply 4-fid, ca. am., the lobes spreading flat, each dee a rh truncate te- al ca. 2 mm wide internally near the apex; sta- ens 16-18, broadly deltoid to ovoid-obtuse, 1— 2 mm long by 1.5-2 mm wide, the filament very short or sessile, the loculi confluent apically into a single crescentiform loculus, the connective thick, not prolonged, the internal receptacle sur- face glabrous. Gynoecious flower subglabrous externally, on a pedicel 20-2 m by 0.8 mm; at anthesis discoid, flat to slightly concave, 5-6 mm diam., the surface bearing ca. 38 densely crowded, free sessile, 4—6-sided, columnar car- pels 0.5 mm diam., interspersed with short, sim- ple hairs. Mature fruiting receptacle enlarged, the torus 15-35 mm diam., bearing 15-25 carpels; carpels ovoid to ellipsoid, 7-8 mm by 4-5 mm, immersed basally in cupules for ca. ' their length, the mesocarp thin, fleshy, the endocarp thin, brown. Distribution. Endemic to Madagascar (Fig. ) 1985] Habitat. The species is known primarily from south central Madagascar where it occurs in the montane Tambourissa/ Weinmannia wet forest zone from ca. 900 to 1,500 m, but also extends up into the sclerophyllous forest zone from ca. 1,300 to 2,000 m DAGASCAR. TULEAR: Mt. Itrafanaomby (Ank B vine and its SW ca: (high Mandrare be: ib rain forest and ericoid scrub on gneissic crests, 0—1,963 m, end Dec. 1933 (fl), Humbert 13518 The species is most closely allied to EpAip- piandra myrtoidea, from which it differs by its androecious flowers, which have more numerous stamens (16-18), and a single broad, truncate tepal on the inner extremity of each receptacle lobe EXCLUDED SPECIES Ephippiandra capuronii Cavaco, Bull. Soc. Bot. France 104: 610. 1957 (= Tambourissa a Lorence, see discussion under this speci Tambourissa Sonn., Voy. Ind. Orient. ed. 1, 3: 267, tab. 134. 1782; ed. 2, 4: a t 134. 1806; Gmel. Syst. Nat. 2(1): 16. 1791, *Tamburissa' (sphalm.); A.DC., Ad 16(2): 658. 1868; Baill., Hist. Pl. ed. 1, 1: 341. 1869; ed. 2, 1: 332. 1871; oil FI. Mau- ritius 287. 1877; Benth. & Hook., Gen. Pl. 139. 1883; Pax, Pflanzenfam. 3Q): 101. 1899; Cordem., Fl. Réunion 300. 1895; Perk. & Gilg, Pflanzenr. 4, 101: 66. 1901; Drake in Grandidier, Hist. Phys. Madagascar 1(1): 21. 1902; Perk., Pflanzenr. 4, 101 (Nachtr.): 39. 1911; Perk., Gatt. Monim. 44. 1925; Cavaco, Bull. Soc. Bot. France 104: 284. 1957; Cavaco in Humbert, Fl. Madagascar 80: 18. 1959; Hutch., Gen. Fl. Pl. 1: 118. 1964. rvPE: T. quadrifida Sonn. Ht cissa Flacourt, Hist. Madagascar 133, pl. 1661 (pre- -Linnaean polynomia PP sie Comm. ex Schreb., Gen. PI. 2: 783. 1791; Willd., Sp. Pl. I1: 27. 1797; es Syst. 5(3): a Son 1855, nom. nov. for Tambour- issa Schrameckia Doai: Bull. Mus. Hist. Nat. (Paris) 28: 249. 1922; Perk., Gatt. Monim. 45. 1925; Metay, LORENCE—MDONIMIACEAE 89 Arch. Bot. Bull. Mens. 3(11): 177. 1929; — Bull. Soc. Bot. France 105: 39. 1958. TYPE: S. madagascariensis Danguy (= Tambourissa. flori- costata Cavaco). Phanerogonocarpus €— Bull. Soc. Bot. France 104: 612, figs. 1- mbert, Fl. — qp 80: Z fig. I:1-6. 1957, synon. nov E: P. capuronii Cavaco (= Tambourissa lon- pis Lorence). Small to moderate sized monoecious or dioe- cious trees, treelets or shrubs, + aromatic. Leaves simple, usually opposite and decussate, rarely clustered into pseudoverticels, or ternate to al- ternate, glabrous or pubescent with simple, rarely fasciculate, unicellular trichomes, the margin en- tire to serrate-dentate with glandular teeth, the venation festooned brochidodromous, the ju- venile and sucker leaves usually heterophyllous, apically serrate-dentate. Inflorescence terminal, axillary, ramiflorous or cauliflorous, sometimes leafy, the axes bracteolate, the flowers solitary, fasciculate or disposed in dichasia, pleiochasia or thyrses; flower in bud fleshy, globose, globose- depressed, E to obovoid, urceolate or e gynoecious bud usually flattened several pairs of minute decussate tepals extend- ing into the orifice; male receptacle at anthesis usually splitting inte (3-2)4—6(-7) + regular, val- vate segments, these remaining partially closed or usually spreading flat and + reflexing; stamens eglandular, few to very numerous, multiseriate, situated on the glabrous or rarely hairy internal receptacle surface, the anthers tetrasporangiate, bilocular, the 2 loculi free and lateral, connivent, or confluent apically or rarely basally, longitu- dinally dehiscent, extrorse, sessile or with a dis- tinct filament, the connective sometimes pro- longed apically; pollen inaperturate, spheroid to ovoid, the sexine striato-reticulate to rugulate, granulate or granular-tectate; female receptacle at anthesis cylindrical, napiform, cupuliform or rarely discoid, the orifice generally 4(-10)-lobed or entire, the lobes reflexing or not; carpels nu- merous, immersed in and fused with the recep- tacle wall, the styles free and protruding, setose or conical to columnar, the internal receptacle surface frequently bearing 7 hairs, in some ucilaginous exu- pendulous, anatropous to amphitropous. Fruit- ing receptacle cupuliform, subglobose, urceolate or cylindrical, corky brown externally, the fruit- ing carpels (drupes) enclosed within the accres- cent receptacle wall which splits open irregularly 90 ANNALS OF THE MISSOURI BOTANICAL GARDEN at maturity; carpels ovoid-compressed, sur- rounded by a fleshy, oily red-orange mesocarp, the endocarp horny, whitish to black, the surface smooth or variously sculptured, the embryo straight, the cotyledons flat, oval to elliptic. Ga- metic chromosome number: n = 19. Distribution. Tambourissa is a genus of about ritius (10 spp.), and Réunion (2 spp.). The species display a high degree of endemism, each being restricted to a given island and frequently to a specific habitat within the island. Habitat. Species of Tambourissa are occa- sional to locally common components of ever- green moist, wet, and cloud forest formations, often conspicuous because of their peculiar flow- ering and fruiting receptacles. TAXONOMIC HISTORY OF TAMBOURISSA The name Tambourissa, adopted from the pre- amb was first validly published by Sonnerat in 1782. Sonnerat based his genus Tambourissa on a sin- gle species, 7. quadrifida, which he undoubtedly described and illustrated while in Mauritius (Ly- Tio-Fane, 1976). A genus characterized by pe- culiar, often large, cupuliform or urceolate to cylindrical fruiting receptacles containing nu- merous immersed carpels, Tambourissa goes by the colorful local appellations of “Bois tambour" (drumwood) or **Pot de chambre jacot" (mon- key's chamber pot) in Mauritius. In neighboring Réunion it is called “Bois de bombarde” (bom- bardment wood) because the ripe fruiting recep- tacle splits open irregularly to reveal an array of numerous, red-orange carpels set against the pale orange receptacular pulp (Fig. 17B). In Anjouan and Grand Comore (Comore islands) Tambou- rissa is known as “‘M’bweza,”” meaning to calm or sooth the pain, because its leaves are used medicinally in tea. Many species in Madagascar and Mayotte (Comores) go by the name of *Am- bora." The species in Mohéli is called ‘“‘Diarou.”’ Sonnerat felt the genus occupied a position somewhere between Ficus and Dorstenia be- cause of its closed, fig-like buds and gynoecious floral receptacle with an inferior, syncarpous gy- noecium. In 1789, the genus was redescribed by A. L. de Jussieu as Ambora from the Malagasy [Vor. 72 vernacular name, and again as Mithridatea by Commerson, Sonnerat's mentor, in Schreber (1791). It is unclear why these authors rede- scribed Tambourissa under other names, be- cause both genera were based on Sonnerat's orig- inal species, 7. quadrifida. It was not until 1855 that Tulasne correctly placed the genus in Mo- nimiaceae, publishing 11 new species of Ambora (18552) for his monograph of the family (1855b). In 1868, Alphonse de Candolle transferred all of Tulasne's species to Tambourissa, describing no new taxa but providing one new name. Baker (1877) described two new species of Tambouris- sa from Mauritius in his treatment of the family for “Flora of Mauritius and the Seychelles" and later described three more from Madagascar. At the turn of the century, Perkins and Gilg (1901) monographed the entire family for Engler's "Pflanzenreich,' "pnovidug s a synoptic treatment for Ti new species. In her revision of the family ten: years later, again for the *“Pflanzenreich,” Perkins (1911) pub- lished four additional species from the Comores and Madagascar. During this same period, Drake del Castillo (1902) published two new species, and Danguy (1922) published another (as Schra- meckia because of its few, large stamens), all from Madagascar. After a hiatus of some 35 years, Cavaco revised the Malagasy Monimiaceae for “Flore de Madagascar et des Comores" (1959), publishing nine new Malagasy species of Tam- bourissa (1957a, 1957c, 1957d, 1957e, 1958c). Most recently, Lorence (1982) published seven new species and one subspecies of Tambourissa from the Malagasy region. Phanerogonocarpus, described by Cavaco (1957c), was named for its narrowly winged, tubular gynoecious flowers and fruits and based on two Madagascan species. In the present revision of Tambourissa, three new names are proposed, and at the same time nine other taxa are reduced to synonymy. GENERIC DELIMITATIONS AND SYSTEMATIC ITION OF TAMBOURISSA The genus Phanerogonocarpus was described by Cavaco (1957b) on the basis of its winged or ribbed, narrowly opera to cylindrical gynoe- les (Fig. 17A), which he compared to the ribbed flowers of Schrameckia. Cavaco (1959) later undermined this character when he reduced Schrameckia to synonymy under 7ambourissa. Furthermore, two species of Tambourissa, T. perrieri, and T. quad- 1985] LORENCE-— MONIMIACEAE 9] rifida, both possess ellipsoid to obovoid gynoe- cious floral receptacles approaching those of Phanerogonocarpus, although not as narrow. Carpels of both genera are identically immersed in and united with the receptacle wall (Figs. 13E, 16A-D). Furthermore, styles of Phanerogono- carpus are conical with an acute apex, possess an oblique stylar canal, and are interspersed with short simple hairs as in Tambourissa (Fig. 16D). Fruiting receptacles of both genera are exter- nally corky brown and split open irregularly at maturity to reveal the numerous ripe carpels im- mersed in a pale orange matrix. That the fruit of Phanerogonocarpus splits lengthwise leaving the central tube more or less intact is related to its elongated shape; the ellipsoid to obovoid fruits of Tambourissa quadrifida behave similarly. Mature fruiting carpels of both genera consist of mesocarp enclosing the of degree and fail to support generic segregation. In his original description of Phanerogono- carpus, Cavaco (1957c) noted the resemblance of its androecious flowers and stamens to those of Tambourissa, except for the presence of tepals (^staminodes") on the inner tips of the receptacle lobes in the former. This feature is, however, shared with a number of Tambourissa species such as T. comorensis, T. floricostata, and T. paradoxa to name a few. Cavaco correctly in- terpreted the staminal morphology of Phanero- gonocarpus as falling within the range of vari- ability displayed by Tambourissa, however. Pollen of Phanerogonocarpus capuronii (— Tambourissa longicarpa) is subspheroid to ovoid with a striato-rugulate sexine composed of short, compactly spaced finger-like elements. That of P. perrieri (— T. alaticarpa) is spheroid to subo- nous patterns of both species are strikingly sim- ilar to those found in many Tambourissa spe- cies, e.g., 7. comorensis, T. Heg T. ficus, T. floricostata, and T. a. A survey of pollen from a range of Haer prins species shows the genus to have sexinous sculpturing ranging from striato-reticulate or rugulate to granular- tectate, thus readily accommodating the patterns displayed by Phanerogonocarpus (Lorence et al., 984 Regarding inflorescence structure, the sexually mixed pleiochasium terminated by one or two gynoecious flowers in Phanerogonocarpus (Fig. 17A) also occurs in a number of Tambourissa species, e.g., 7. hildebrandtii and T. perrieri. Fi- nally, leaf architecture of the two genera over- laps. Adult leaves of a number of Tambourissa species, e.g., 7. thouvenotii and T. trichophylla, as well as juvenile and sucker leaves of many species are dentate, as are adult leaves of Phane- rogonocarpus. I can find no obvious basis for maintaining Phanerogonocarpus as a distinct ge- nus and therefore propose that it be reduced to synonymy under Tambourissa. Tambourissa has attained the highest level of floral morphological specialization of all the alagasy Monimiaceae in the subfamily Mol- linedioideae. Most species are characterized by a strong androecious/gynoecious floral dimor- phism, have flowers of both sexes closed in bud, have highly reduced tepals, show a range of stam- inal specialization via reduction and, most sig- nificantly, the gynoecious flowers have their car- pels immersed in and fused with the massively developed ovary wall, with the fruiting carpels likewise enclosed in the accrescent receptacle which splits open irregularly at maturity. INTRAGENERIC RELATIONSHIPS Tambourissa, with about 43 species, is the largest genus of Monimiaceae in the Old World. Its members often display extensive popes variation in flora olor in y and number of p arts), EM cr pe r- phology, and bu. characters. Even species ien are obviously closely related with der to IN tures such as foliar and gynoecious floral m phology may differ radically and Inexplicably in in other features suc inal morphology. This happens frequently pie i to render the establishment of meaningful subge- neric categories such as subgenera or sections impossible at present. I have, therefore, chosen to place the species into informal groups of mor- phologically similar taxa to facilitate their com- arison, but no phylogenetic implications are in- tended by this sequence. Nine species groups are listed below, each followed by a discussion (refer also to Tables 5-7): 1. Group A. ..... Tambourissa thouvenotii, T. trichophylla, T. uapaci- folia Group B. ..... Tambourissa decaryana, T. hildebrandtii, T. hum- 92 ANNALS OF THE MISSOURI BOTANICAL GARDEN bertii, T. nitida, T. nosy- ensis Group C. ..... Tambourissa floricostata, . parvifolia Tambourissa beanjaden- sis, T. comorensis, T. crassa, T. elliptica, T. lep- tophylla, T. madagascar- iensis d kirkii, T. paradoxa inr s capuronii, T. moheliensis Tambourissa gracilis, T. purpurea, T. religiosa Tambourissa amplifolia, T. cocottensis, T. cordifo- lia, T. pedicellata, T. tau, T. tetragona Tambourissa bathiei, T. sieberi Tambourissa perrieri, T. quadri Tambourissa alaticarpa, T. longicarpa Tambourissa castri-delphi- nii, T. ficus, T. peltata 2. Group A. Group 1. This is a group of Madagascan species with canescent to velutinous or tomen- tose stems, leaves, and inflorescences (Table 5). The inflorescence is usually a ramiflorous or ax- illary (rarely cauliflorous) pleiochasium, often sexually mixed and frequently terminated by a gynoecious flower, or the flowers may be solitary. Long acute, subulate or deltoid bracteoles sub- tend the pedicels and floral axis. The androecious flower splits open flat into 4(—5) segments, bears scattered hairs internally in many species, and has fairly numerous stamens, frequently with prolonged connectives. The urceolate, napiform or opens by a small four- lobed, X- -shaped orifice in most, and the densely velutinous internal surface is lined with many shortly conical styles. In most species, an abundance of oil cells occurs in the floral ground tissue of both sexes. Group 1A. This group is comprised of three species with densely pubescent, usually api- cally toothed, adult leaves (Tambourissa thou- venotii, T. trichophylla) and densely velutinous- tomentose mature stems (Table 5). Tambourissa [Vor. 72 uapacifolia fits here on the basis of its dense pu- escence, although its leaves are entire and the inflorescence is apparently unisexual. oup 1B. Five species with entire leaves and sparsely pubescent to canescent adult leaves and stems comprise this group (Table 5): Tambouris- sa decaryana, T. hildebrandtii, T. humbertii, T. nitida, and T. nosybensis. Of these, T. decary- ana, T. humbertii, and T. nosybensis show trends rss cauliflory and unisexual inflorescences. IC. The last group contains two species differing in having subglabrous mature growth and internally glabrous female flowers (Tables 5, 6). Tambourissa floricostata has few, large sta- mens and floral receptacles with longitudinal lat- eral ridges; it was formerly placed in Schram- eckia etry 1928). In T. pens the leaves are small and sclerophyllous and the stamens slender with distinct filaments lacking prolonged connectives. roup 2. This diverse group includes species from the Comores, Madagascar, and Réunion, some of which appear to be much more closely related than others (Tables 5, 6). Most have gla- brous to sparsely pubescent mature growth and inflorescences. Varying degrees of reduction oc- ly mixed or unisexual pleiochasium to a fascicle or solitary flowers. Small deltoid brac- teoles subtend the often long, slender floral axis and pedicels. Two subgroups can be distin- guished. Group 2A. This category consists of species with medium-sized to large, 4-fid androecious flowers that open Ent and contain numerous long stamens confl uent loc uli and scarcely or non- SiGe connectives (Ta- les 6, The gynoecious flowers are globose- depressed, closed in bud, internally + glabrous with crowded, shortly columnar blunt styles, open by an X-shaped orifice, and produce a copious mucilaginous secretion in the two Réunion species (Tambourissa crassa, T. elliptica) and T. bean- jadensis (Madagascar). In 7T. madagascariensis (Madagascar) and the Comorean species (7. comorensis, T. leptophylla) the orifice of the gy- noecious receptacle is entire, internally pubes- Styles ofthe two latter species are bluntly colum- nar and shortly conical, respectively. Group 2B. This group consists of two Como- rean species in which the androecious flowers are 1985] small and globular, splitting only slightly and remaining nearly closed (Tambourissa kirkii), or merely opening by a small apical pore lined with scale-like tepals (7T. paradoxa) (Table 6). In both species stamens are fewer, smaller, and more acute than in Subgroup A and have separate, nsis corre- spond well with this group, but because its gynoe- cious flowers differ markedly from the other members, it will be discussed under Group 3. Gynoecious flowers of T. kirkii have an entire, circular orifice like T. comorensis and T. lepto- phylla, thus linking it to Group 2A; those of T. paradoxa are unknown Group 3. Members af this group are charac- terized by comparatively large, brown napiform gynoecious flowers that are closed and apically depressed in bud, later opening by a four-lobed, X-shaped orifice (Table 6). Unlike Group 2, the floral ground tissue contains numerous dark brown, presumably tanniferous, idioblasts. An- droecious flowers of Tambourissa capuronii (Madagascar) are large, split open flat into four lobes, and have numerous large stamens with elongated lateral loculi. Those of T. moheliensis (Comores) are much smaller than in the forme species, only split slightly open at anthesis (as in Group 2B), and differ in having fewer stamens with loculi that unite basally into a U-shape. Its androecious flowers therefore link Group 2B with 3 Group 4. This group is composed of three Madagascan species which are glabrous to sub- acteristic terminal dichasia, condensed pleio- chasia or solitary flowers (Tables 5, 6). The gy- noecious flowers and fruit are terminal, whereas the androecious flowers are either lateral, axil- lary, or ramiflorous. Floral receptacles are small ceptacles are smooth and dark reddish external- ly, not pale and corky brown as in most other species. Tambourissa gracilis and T. purpurea both have similar androecious flowers with short, sessile stamens having apically confluent, cres- centiform loculi; gynoecious flowers of the for- mer are unknown. Flowers of T. religiosa are somewhat larger with longer, shortly conical styles. Also, its stamens have distinct filaments and separate loculi. Gynoecious flowers of T. LORENCE-— MONIMIACEAE 93 purpurea produce a mucilaginous plug or **hy- perstigma" which occludes the pore; it is not known if this phenomenon occurs in the other two species. Group 5. This group of subglabrous to gla- brous trees, treelets and shrubs includes the bulk the Mauritian species (Tables 5, 6). Tam- de pedicellata seems to be the least E cialized member in view of its monoecy, vari- ability of floral disposition and s j. structure, and particularly its monomorphic an- droecious and gynoecious flowers with shallowly subglobose to cupuliform receptacles, relatively few parts, shortly conical styles, and stamens with separate loculi and distinct filaments. ther members of the group (Tambourissa amplifolia, T. cocottensis, T. — T. tau, and T. tetragon is theme with varying pile of i scary in one or more features. Leaves of most species are mod- erately small, opposite and decussate, and have distinct petioles. Trends of specialization are to- wards macrophylly (7. amplifolia, T. tau), clus- tering of the leaves into pseudoverticils of two to four pairs (7. tau), winged stems (7. tetra- gona), and reduction in petiole length (7. cor- difolia, T. tau). Modifications of the inflores- cence tend towards reduction from pleiochasia to dichasia, and finally to fascicles and mono- chasia, towards a shortening of the pedicels and floral axis, and from ramiflory to (often basal) cauliflory. Other apparent trends are towards greater flo- ral dimorphism, towards larger flowers with more numerous parts fu of the loculi apically (T. anpa basally (T. cordifolia), or their extreme separation on lateral arms (T. tau). Gynoecious mira display a greater deepening and closure (7. cocottensis, T. tetragona) or opening (T. peltata), modifications of the styles via elongation (7. tau) or shortening (T. peltata), and elaborate development of the receptacular lobes with bright coloration to en- hance their attractive potential for pollinators (T. cordifolia, T. peltata). The latter two species are rther specialized in being dioecious. Gynoe- cious flowers of 7. ficus correspond to this group in spite of their gigantism. Because of the ternate leaves and androecious floral morphology, how- ever, I have included T. ficus under Group Group 6. This group consists of two species of large, monoecious trees, i.e., Tambourissa sie- beri (Mauritius) and 7. bathiei (Madagascar). 94 ANNALS OF THE MISSOURI BOTANICAL GARDEN Both have relatively small leaves, moderately to densely pubescent new growth and densely yel- lowish velutinous, cauliflorous inflorescences which are generally unisexual pleiochasia or thyrses (Tables 5, 6). The globose female recep- tacle opens by a small pore in 7. sieberi (mature gynoecious flowers unknown in T. bathiei), and is lined with numerous long, narrowly conical, generally coalescent styles. Androecious flowers of both species are large and 4—5-fid with lobes that spread open flat and have numerous large stamens. Group 7. Apparently related to Group 6, this category also contains two species not closely allied to other members ofthe genus: Tambouris- sa quadrifida (Mauritius) and 7. perrieri (Mad- agascar). Both are glabrous cauliflorous trees with relatively small, opposite leaves, grow in rela- tively drier habitats, and produce numerous uni- sexual or sexually mixed pleiochasia or thyrses along most of the trunk (Tables 5, 6). Androe- cious flowers of both species split deeply into four flat or strongly recurved segments and bea hundreds of short stamens. The ellipsoid to ob- ovoid or obpyriform gynoecious flowers open by a small, three- to four-lobed apical pore, are in- ternally velutinous and contain hundreds of very short, basally ventricose styles. The mature fruit- -t ° d = Q P — [4] P4 oO 3 = ge ER = ° 72 oO ° me = This isolated pair of Madagascan species, Tambourissa alaticarpa and T. longi- carpa, formerly comprised the genus Phanero- gonocarpus. Both are pubescent, monoecious, cauli- to ramiflorous treelets of the eastern wet forest zone with large, dentate leaves (Tables 5, ly three) terminal gynoecious flowers with lateral clusters of relatively small androecious flowers grouped along the floral axis, a feature shared with certain members of Groups 1 and 7. The stamens have distinct filaments, free loculi, and the connective may or may not be prolonged. Most distinctive of Group 8 are the slender, ob- ovoid to cylindrical, narrowly ribbed or winged gynoecious flowering and fruiting receptacles with a small orifice. Gynoecious receptacles of T. flo- ricostata (Group 1C) are also ribbed, however, and those of Group 7 are ellipsoid to obovoid, although not as narrow, with comparably small orifices. The conical to ventricose styles are in- terspersed with dense short hairs in both species, [Vor. 72 and a mucilaginous exudate is apparently pro- duced in T. longicarpa. Group 9. Characterized by ternate to subal- ternate adult leaves with prominent, highly or- ganized and finely reticulate venation, this group consists of three species (Tables 5, 6), i.e., Tam- bourissa castri-delphinii (Madagascar), T. ficus, and 7. peltata (both Mauritius). Flowers are pro- duced either at the base of the trunk (T. ficus), or on the upper trunk and older branches (7. peltata and T. castri-delphinii). The highly re- duced inflorescence ranges from a short, con- other two species. The group also displays a trend towards dioecy. Androecious flowers of Tambourissa ficus are the largest in the entire genus, with five to seven incurved or spreading lobes bearing from 900 to 1,800 ligulate stamens with separate, lateral to unifacial loculi and a thick, prolonged connec- tive. Androecious flowers of T. peltata and T. castri-delphinii are considerably smaller, deeply split into four or five recurved lobes, and bear numerous thin, broadly deltoid subsessile sta- mens with connivent or confluent lateral loculi. Gynoecious flowers of Tambourissa peltata are highly specialized, flat and discoid with four to seven elaborate, spreading verrucose sterile lobes ly r inous exudate. Those of T. castri-delphinii have an X-shaped orifice and conical styles, similar to some species in Group 5. In T. ficus the gynoe- cious flowers are huge and urceolate, splitting into numerous straight or inflexed lobes with a broad orifice, and contain ca. 1,000-2,000 setose styles. Apart from their great size, gynoe flowers of T. ficus are morphologically similar to those of species belonging to Group 5 and link the two groups. In the following key I have attempted to utilize vegetative characters as far as possible. Although certain species with outstanding vegetative char- acters are identifiable even from sterile speci- mens (e.g., Tambourissa tetragona is the only species with winged stems), for the majority, complete collections consisting of leafy stems, androecious and gynoecious flowers, and fruits are essential for reliable identification. Unfor- known only from androecious or gynoecious 1985] flowers or fruits, and this has imposed certain limitations on the construction of this key. Al- though I was unable to examine all the Mada- gascan material cited by Cavaco (1959), I have seen additional material not cited by him. All collections in the Exsiccata and virtually all those used to plot the distribution maps have been seen y me. The last five species in the taxonomic treatment (nos. 39—44) are imperfectly known and consequently have not been included in the key. See key cited on following pages. 1. painless trichophylla Baker, J. Linn Soc., 0: 240. 1884; Perk. & Gilg, Pflanzenr. 4, er 68. 1901; Drake in Grandidier, Hist. Phys. Madagascar 1(1): 24. 1902; Perk., Pflanzenr. 4, 101 (Nachtr.): 41. 1911; Ca- vaco in Humbert, Fl. Madagascar 80: 24, fig. VII:8-9. 1959. rvPE: Madagascar. (“Central Madagascar," probably Tanana- rive or Fianarantsoa): without precise lo- cality, 1882 (fl), Baron 1953 (lectotype, K, here designated). Monoecious treelet or tree attaining 15 m tall and 15-20 cm D.B.H., the new growth densely pilose-tomentose, the hairs simple, pale yellow- ish to grayish, the mature leafy stems 2-4 mm diam., tomentose, the hairs + matted. Leaves opposite to subopposite, petiolate; petioles to- mentose, 4-10 mm by 1.5-2 mm; lamina char- taceous to cn oblong, narrowly ob- long, owly obovate, lanceolate or rarely cibus pur s 95 mm by 20-52 mm, the apex shortly acuminate, shortly acute or rarely obtuse, id base obtuse to rounded, rarely acute, abax- ially tomentose-hirsute, especially along costa and secondary veins, the trichomes curved, adaxially pilose when young, glabrescent, the secondary veins 3-5 pairs, making a 50-60? angle with the costa, the arches low and broad, the venation disorganized, obscure adaxially and visible to 2°, prominent abaxially and visible to 3?, the margin slightly thickened and revolute, ciliate, entire or usually serrate-dentate in the apical 45-4 of the lamina with 1—2 pairs of short, broadly deltoid teeth. Inflorescence ramiflorous on leafless nodes or axillary, a usually unisexual, sometimes leafy pleiochasium of 3-7 flowers, or the flowers sol- itary, the floral axis fulvous to grayish, veluti- nous-tomentose, 14-55 mm by 1.5-2 mm, sub- tended by several deltoid-naviculate bracteoles 1-2 mm long, the pedicels 3-5 mm by 1.5 mm, LORENCE-— MONIMIACEAE 95 each subtended by 1(-2) — velutinous bracteoles 4-5 mm long. Androecious flower in bud globose, 5-7 mm diam., sema apiculate with several pairs of minute hirsute tepals; an- droecious flower at anthesis unknown (presum- ably deeply 4-fid with spreading lobes as in 7. thouvenotii); stamens in bud ca. 100, ovoid- E toid, 1.5-2 mm long by 0.8-1 mm wide, loculi lateral, separate, occupying ca. 3⁄4 Apu length of stamen, the filament short or sessile, the connective thick, slightly prolonged apically, the internal receptacle surface with clusters of hairs between the stamens. Gynoecious flower in bud solitary or occasionally in pleiochasia, 7-8 mm diam pressed, hirsute: apiculat tepals; at anthesis ibid 4(—5)-fid, the vun + incurved, the small X-shaped orifice 2-3 m diam., the duci ca. 100, conical, 0.6—0.8 mm long by 0.3-0.4 mm wide basally, + ventricose, acuminate, the internal receptacle surface velu- tinous between the styles. Fruiting receptacle sol- itary on the smaller branches, shallowly cupuli- form, 30-48 mm diam. by 30 mm long, the walls Ee i33 mm hisk, both surfaces pale corky brown, 5 total diam. of the fruit, the scattered. styles conical, 0.3—0.5 mm long and wide basally, the internal receptacle surface hirsutulous; pedicel and peduncle stout, 6-8 mm long by 7-12 mm diam., hirsutulous. Submature carpels 6-8 mm long, 4-5 mm wide, 3—4 mm thick, ovoid-compressed, the endocarp brown. Distribution. Endemic to Madagascar (Fig. Habitat. The species is locally common and fairly widespread in the Tambourissa/ Wein- mannia wet forest zone in the eastern and central regions ranging from 800 to 1,500 m. MADAGASCAR. TAMATAVE: Ankaraoka, ca. 1,500 m, (MAD); National Route no. 2 at 69 k rive, Mandraka forest reserve, 1,400 m, 28 Oct. 1978 (fl), Lorence 1879 (MAD, MAU, MO); 30 Oct. 1978 (fl), Lorence 2034 (MAD, MO), (fl), Lorence 2035 MO); La Mandraka, Sept. 1963 (st), Morat 7 (MAD); National Route no. 2 at ca. 20 km W of Peri- net, 950 m, 29 Oct. 1978 (st), Lorence 2000 (MAD, MO). TANANARIVE: Andringitra Massif, Ambodigraiss Forest near Antsirabe (sic), 1,200 m, 12 Jan. 1945 (fr), Anon. sub Herb. Alaotra 2242 (MAD). 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(K, syntype); Dec. 1883 (st), Baron 2243 (K); (f), Baron 2994 ( (BU, K); 1885 (fl), Baron 3433 (K); Feb. 1882 (st), Parker s K). Tambourissa trichophylla and its close ally, T. thouvenotii, are both characterized by apically rower oblong leaves with fewer and broader teeth (only 1-3 pairs), its more widely spaced second- ary veins with lower, stronger arches, its sparser and more curved trichomes, and its smaller flow- ers with much shorter pedicels and longer brac- teoles. Both species remain distinct where they occur sympatrically and both belong to species Group 1A ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Vernacular name. Ambora saha (Madagas- car). 2. Tambourissa thouvenotii Danguy, Bull. Mus. Hist. Nat. (Paris), Ser. 1, 28: 250. 1922. TYPE: Madagascar. Tamatave: Analamazaotra (Perinet), ca. 1919 (fl), Thouvenot 109 (lec- totype, K, here designated; isolectotypes, P, 2 sheets) T. trichophylla Baker var. thouvenotii Cavaco in Hum- ert, Fl. Madagascar 80: 25, fig. VII:1-4. 1959. Monoecious or subdioecious treelet or tree 8— 22 m tall and 15 cm or more D.B.H., the new growth densely velutinous, the trichomes simple, yellowish, the mature leafy stems ipiis 3- mm diam. (sucker stems to 12 mm diam.). Leaves opposite, petiolate; petioles RA 10-20 mm by 2-3 mm; lamina subcoriaceous, broadly elliptic to elliptic, rarely broadly oblong, ovate or obovate, (70-)85-185 mm by (37-)50- mm, the apex shortly acuminate, rarely ob- tuse, the base obtuse to rounded, the young lam- ina adaxially pilose, glabrescent except on costa and major veins, abaxially densely velutinous with soft, straight yellowish hairs, the secondary veins 4-7 pairs, making a 35-50? angle with the costa, the arches high and narrow, the venation obscure adaxially and visible to 2?, prominent of the lamina usually serrate-dentate with 1—8 pairs of short teeth. Inflorescence cauliflorous, ramiflorous or axillary, a sexually mixed or uni- sexual pleiochasium of 3-7 flowers, or the flow- ers rarely solitary, the floral axis velutinous, 25- 57 mm by 2-3 mm, subtended by several ve- lutinous, deltoid-subulate bracteoles 2-3 mm l a several minute, velutinous tepals, the pedicel 12- 17 mm by 1.5-2 mm; at anthesis deeply 4-fid, 12-25 mm diam., the lobes spreading flat; sta- mens numerous, ca. 50-100, deltoid-lanceolate to subulate, 3-7 mm long by 1-1.5 mm wide, slightly incurved, the filament short or sessile, the loculi lateral to slightly unilateral, separate, occupying OM total length of the stamen, the d apically, acute, the internal receptacle surface pubes scent. Gynoecious flower in bud a at anthesis shallowly and irregularly 5-6-lobed, LORENCE-— MONIMIACEAE 103 1985] the orifice 3-4 mm diam., the lobes incurved, not reflexed; styles numerous, ca. 100-150, very shortly conical, 0.3 mm long by 0.2 mm wide basally, the apex acuminate to acute, the internal receptacle surface velutinous between the styles. Fruiting receptacle solitary on old branches, ir- regularly cupuliform, 40-100 mm diam. by 32- 55 mm long, the walls 12-18 mm thick, both surfaces corky, reddish brown, externally hir- sute-velutinous, the orifice subentire, comprising V4—V» total width of the fruit; styles numerous, shortly conical, 0.3-1.6 mm long by 0.3-0.5 mm wide basally; pedicel and peduncle 15-50 mm long by 9-15 mm wide. Mature carpels 15-16 mm long, 6-7 mm wide, 4-5 mm thick, the en- docarp light brown, the surface scrobiculate. Distribution. Endemic to Madagascar (Fig. Habitat. Tambourissa thouvenotii is fairly widespread and often locally common in the Tambourissa/ Weinmannia montane wet forest zone of the eastern and central regions from ca. 800 to 1,400 m. In addition to the obvious mor- phological differences, both T. thouvenotii and T. trichophylla appear to occur sympatrically in at least some areas, e.g., Massif d'Andringitra (Anon. sub Herb. Alaotra 2242, MAD, Anon. sub Herb. Alaotra 2311, MAD), where they remain distinct. MADAGASCAR. MAJUNGA: Circagri Bealanana (near Tsaratanana massif), June 1953 (st), Anon. su ae herana, Fianarantsoa, 24 Nov. 1949 (fl), Anon. ns SF 1248 (MAD); Andringitra Massif, Ambatapaiso near Antsirabe (sic), 12 Jan. 1945 (st), Anon. sub Herb. Ala- otra 2311 (MAD); in rain forest, Tsararano, Vondrozo district (E coast, near Farfangana), Dec. 1963 (fl), Bos- rence 2014 (MAD, MAU, M! y (MAD, MAU, MO); Perine 912 (st), Viguier & rs and 1 106 ps Ambatondrazaka (fl), Anon. sub SF 2548 (MAD). wiTHOUT PRECISE LOCALITY: North Madagascar, 1892 (fr), js 6721 (K). Tambourissa thouvenotii seems most closely related to T. trichophylla, which also possesses dentate-serrate adult leaves. It is distinguishable from the latter species by its much larger, broader and usually elliptic leaves with more numerous teeth (up to 8 pairs) and more closely spaced secondary veins with higher and weaker arches showing a euo to become almost craspe- 7 A). Tambourissa thouvenotii straight hairs, much longer floral pedicels, short- er bracteoles, and larger flowers than does T. trichophylla. Both species are closely allied to T. uapacifolia which also belongs to species Group lA Vernacular names. Ambora, Ambora bera- vina (Madagascar). 3. Tambourissa uapacifolia Cavaco, Bull. Mus. Hist. Nat. (Paris), Ser. 2, 29: 351. 1957; Ca- vaco in Humbert, Fl. Madagascar 80: 24, fig. V:6-7. 1959. TYPE: Madagascar. Tulear: upper basin of the Mandrare River (SE), pass and summit of Marosoui, 1,000-1,400 m, 14-15 Nov. 1928 (fl, fr), Humbert 6604 (ho- lotype, P; isotypes, P, 2 sheets) Monoecious tree, the new growth densely ve- lutinous, the mature leafy stems 4-5 mm diam., velutinous-hirsute. Leaves opposite to subop- posite, petiolate; petioles 5-16 mm by 1.5-2.5 mm, velutinous; lamina subcoriaceous, obovate to elliptic, 65-155 mm by 20-85 mm, hirsutu- lous when young, later glabrescent except abax- ially on the major veins and margin, the apex obtuse or rarely shortly acute, the base acutely cuneate, rarely obtuse, the secondary veins 4-6 pairs, making a 40—50? angle with the costa, the venation depressed and visible to 3? adaxially, raised and visible to 4? abaxially, the margin thickened, slightly revolute, ciliate. Inflorescence cauliflorous or ramiflorous on leafless nodes, a velutinous, pendulous unisexual (always?) pleio- chasium of 5-9 flowers, the floral axis 20-55 mm apiculate, the apex with several pairs of small, hirtellous-velutinous tepals, the pedicel veluti- nous, 10-17 mm by 1-1.5 mm; at anthesis 4-fid; 17-20 mm diam., stamens ca. 100-300, clavate, 1-2 mm long by 0.6-1 mm wide, the filament comprising ca. 3—2 the total length, the loculi lateral, separate or confluent apically, + parallel, comprising ca. !5—/ total length of the stamen, the connective not prolonged, the external sur- face of loculi bearing medially several scattered simple hairs, the internal receptacle surface bear- 104 ing scattered simple hairs between the stamens. Gynoecious flower unknown. Fruiting receptacle solitary on older leafless stems, the submature fruit cupuliform, 35-41 mm diam. by 20-27 mm long, the orifice ca. 10-15 mm diam., subentire, the walls 14-16 mm thick, externally pale corky brown with scattered hairs, the internal surface hirsute, the styles scattered, broadly conical, 0.5- 0.6 mm long by 0.6-0.8 mm wide basally; pedicel and peduncle hirsute-velutinous, 25-50 mm long by 3-4 mm diam.; submature carpels 9-12 mm long, 7-8 mm wide, 5-6 mm thick, the endocarp light brown, the surface foveolate. Distribution. Endemic to Madagascar (Fig. Habitat. Tambourissa uapacifolia is current- ly known from three collections. The type is from southeastern Madagascar at 1,000 to 1,400 m, presumably in Tambourissa/ Weinmannia wet forest where Humbert also collected 7. capuronii and T. hildebrandtii. The other two collections are from near Moromanga, probably in the same vegetation type. Additional collec ink and field data are obviously neede MADAGASCAR. TAMATAVE: S of Moromanga, 9 Feb. 1930 (fl), Decary 7042 (Pj; Moromanga to Andasibe (Perinet), pk 51, 26 Oct. 1953 (fr), Anon. in SF-7805 (P). Tambourissa uapacifolia belongs to species Group 1A, which also includes 7. trichophylla and 7. thouvenotii (for which Humbert bracteoles, stamens with scattered hairs on the loculi, unprolonged connectives, and by its ob- tuse leaves with entire margins resembling those of certain species of Uapaca (Euphorbiaceae). 4. Tambourissa nitida Danguy, Bull. Mus. Hist. Nat. (Paris), Ser. 1, 34: 280. in Humbert, Fl TYPE: Madagascar. Dieg f Bir amanja (Ambilobe), ca. 1927 (fl, fr), Ursch 267 (holotype, P; isotype, P). Small tree attaining 6 m tall, the new growth pilose-hirsute, the mature leafy stems 1.5—4.5 mm diam., blackish brown, hirsutulous. Leaves glabrous, opposite, petiolate; petioles 10-16 mm by 1-1.5 mm; lamina chartaceous to subcoria- ceous, ovate to narrowly ovate, elliptic, rarely oblong, 60-110 mm by 26-40 mm, the apex long ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 acuminate, + recurved, the base acutely decur- rent, the secondary veins 6-9 pairs, making a 60—70? angle with the costa, the venation prom- inent on both surfaces, visible to 3? adaxially and to 4? abaxially, the adaxial surface highly lus- trous, the abaxial surface dull, the margin slightly revolute. Inflorescence axillary, a sexually mixe pleiochasium of 3-5 flowers, or the flowers sol- itary, the axes L canescent with pale, sim- ple hairs; floral axis 5-12 mm by 1 mm, sub- tended by several subulate-naviculate, hirsute bracteoles 1.5-2 mm long. Androecious flower in bud globose, 6-7 mm diam., sparsely pubes- tended by a bracteole; androecious flower at an- thesis unknown; stamens in bud ca. 60, ellipsoid to clavate, 1-1.5 mm long by 0.5 mm wide, the loculi separate, lateral, occupying ca. !^ total length of the stamen, connivent at the obtuse apex, the connective not prolonged, the internal receptacle surface with scattered groups of simple hairs between the stamens. Gynoecious flower at anthesis unknown, in bud turbinate or napiform, .5-8 mm diam. by 4-5 mm long, usually with two low, basal longitudinal ridges, externally hir- sutulous-canescent, apiculate with 3—4 acute te- pals; styles numerous, ca. several hundred, con- ical or clavate, 0.7—0.9 mm long by 0.3 mm diam., medially ventricose, the apex abruptly ormed fruit axillary on the leafy stem, + napi- form, ca. 15 mm diam,, externally blackish and corky brown, pubescent, the internal surface hir- sute, the scattered styles broadly conical, taper- ing to an acuminate apex, 0.5—0.6 mm wide by 0.8-1 mm long; pedicel and peduncle stout, ob- conical, 10-12 mm long by ca. 5 mm diam. me- dially. Carpels unknown. Distribution. Endemic to Madagascar (Fig. ). Habitat. Tambourissa nitida is only known from two collections, both from the Sambirano region of Diego Suarez near the northern tip of Madagascar. The area where both collections originated appears to represent a transition zone between wet and deciduous forest. MADAGASCAR. DIEGO SUAREZ: Ifazy Valley, Aug. 1909, Perrier P la Báthie 10126 (P). Tambourissa nitida belongs to species Group 1985] LORENCE- MONIMIACEAE 105 1B. Although the slight ridges on the female re- of T. floricostata, T. adaxially lustrous leaves with long acuminate, recurved apices. Also, its gynoecious flowers are more densely velutinous internally and its sta- mens lack the prolonged connectives of most al- lied species. Mature flowers, fruiting receptacles, as well as the habitat of this remarkable species are unknown. 5. Tambourissa hildebrandtii Perk. in Perk. & Gilg, Pflanzenr. 4, 101: 69, fig. 18:N-P. 1901; Perk., Gatt. Monim. fig. 34:N-P. 1925; Ca- vaco in Humbert, Fl. Madagascar 80: 29. 1959. Type: Madagascar. Tananarive: North Betsileo (Ant-) Sirabe, Aug. 1880 (fl), Hil- debrandt 3563 (lectotype, K, here designat- ed; isolectotypes, BM; G, 2 sheets; P, 2 sheets) T. baroni Drake in Grandidier, Hist. Phys. Madagascar 1(1): 24. 1902; Perk., Pflanzenr. 4, 101 (Nachtr.): 41. 1911; Cavaco, Bull. Soc. Bot. France 104: 284, figs. 2, 5. 1957; Cavaco in Humbert, Fl. Mad PAP 80: 26, fig. VII:5-7. 1959. TYPE: s out precise locality (ca. 1890), Baron 2954 dectotpe K, here designated; iso- lecto T. id sen su Drake in Grandidier, Hist. Phys. Madagascar 1(1): 21. 1902, pro parte, non Sonn. Monoecious tree attaining 10 m tall and 30- 40 cm D.B.H., the bark thin, gray, flaking, the new growth villous to velutinous with straight or matted simple hairs, the mature leafy stems 1.5- mm diam., dark brown, villous, glabrescent. Leaves opposite to subopposite, petiolate; peti- oles villous-hirsute, 4-12 mm by 1-1.2 mm; lamina chartaceous to subcoriaceous, sparsely pilose abaxially, more densely so along the costa and major veins, adaxially glabrescent, ovate, elliptic to narrowly elliptic, oblong or narrowly oblong, lorate or rarely obovate, 40-115(-135) mm by 17-44 mm, the apex shortly acuminate, shortly acute, obtuse or emarginate, the base acute to obtuse, sometimes slightly decurrent, the sec- ondary veins 4-9 pairs, making a 55-80? angle with the costa, the venation visible to 2(-3?) adaxially, to 3(—4°) abaxially, the margin thick- ened, slightly revolute. Inflorescence terminal, axillary or ramiflorous, cinereous, velutinous-to- mentose, a unisexual or sexually mixed pleio- chasium of 3-7(—9) flowers, usually i peni by a single gynoecious flower, the floral axis 5- 30 mm by 1.2-2 mm, the axis and pedicels sub- tended by naviculate-subulate bracteoles 1—3.5 mm long. Androecious flower in bud finely ci- nereous-tomentose, globose to globose-de- pressed, 6-7 mm diam. by 5-6 mm long, apic- ulate with 4 narrowly acute tepals ca. | mm long, the pedicel 8-10 mm by 0.6-1 mm; at anthesis deeply 4-fid, 15-18 mm diam., the lobes spread- ing flat; stamens ca. 75-90, deltoid to ovoid- ellipsoid, slightly recurved, 1.2-2 mm long by 0.8-1.2 mm wide, the loculi + unilateral, in- trorse, separate, occupying 75—À total length of the stamen, the filament subsessile, the internal receptacle surface with scattered to dense simple hairs between the stamens. Gynoecious flower in bud subglobose-napiform, more densely tomen- tose than androecious flower, apiculate with 4 acutely deltoid tepals, the pedicel 10-15 mm by 2 mm; at anthesis 7-9 mm diam. by 5-6 mm long, the orifice X-shaped, 3-3.5 mm diam., splitting into 4 deltoid, suberect to outcurved lobes; styles numerous, ca. 150 , narrowly conical, 1-1.5 mm long by 0.4-0.6 mm wide basally, the surface strongly papillose, the inter- nal receptacle surface d lutinous-tomen- tose between the styles. Fruiting receptacle usu- ally solitary on leafless stems, + shallowly cupuliform, 30-50 mm diam. by 20-35 mm long, the cavity shallow to almost absent, the external surface pale, mottled corky brown, the orifice comprising 1—12 total width of the fruit, suben- tire, the walls 15-225 mm thick, the internal sur- face with scattered, narrowly conical papillose styles 0.5—0.7 mm long by 0.5 mm wide basally, interspersed with scattered to dense hairs; ped- icel and peduncle tapered, to 55 mm long by 4— 5 mm diam. medially. Mature carpels 7-8 mm long, 4-5 mm wide, 3-4 mm thick, ovoid-com- pressed, the endocarp light brown, the surface scrobiculate. l Distribution. Endemic to Madagascar (Fig. Habitat. Tambourissa hildebrandtii displays a wide ecological amplitude comparable to that of T. purpurea. It ranges from the Tambourin wet forest zone abo m in the eastern region, to montane forests of the central plateau (ca. 1,000-1,400 m) and ex- tends to the dry semideciduous forests of the western domain where it finally disappears. MADAGASCAR. FIANARANTSOA: Fianarantsoa (fr), Anon. sub SF 2145 (MAD); peg - W of Amba- lavao, at Pk 475, 970 m, 31 Jan. 5 (fl, fr), oT 30194 (MAD, MO); National rd no. 7 at 11 k 106 W of Ambalavao, 950 m, 5 Nov. 1978 (fl, fr), Lorence 2043 (MAD, MAU, MO). MAJUNGA: Natural reserv no. 8, Namoraka (Saolala), 7 Feb. 1951 (fr), Anon. sub RN 3298 (MAD). TAMATAVE: Ambatondrazaka (near Lake Alaotra) (fl, fr), 4non. sub. SF 1723 (MAD); Am- bohijanahary, 1,000 m (by Lake Alaotra), 5 Jan. 1945 (fl), Anon. sub SF 2188 (MAD); small forest at Ma- rovato, East Manaka (near Lake Alaotra), 13 Nov. 1946 (fl), Anon. sub Herb. Alaotra 3102 (MAD, 2 sheets); Didy Forest, Ambatondrazaka (near Lake Alaotra), ca. 1,200 m, 5 July 1952 (st), Anon. sub SF 5498 (MAD); 2 June 1952 (fl), Botoalma sub SF 4145 (MAD, 2 : 3 (K, 2 sheets); Bongolave, 800—900 m, l sj. July 1974 (fr), PROVINCE UNCERTAIN: to Nickelville, 29 Dec. 1944 (fl, fr), Anon. sub Herb. Alaotra 2091 (MAD, 2 sheets); Menavavakely, Mev- atanana, 16 Apr. 1951 (fl), Anon. sub SF 3588 (MAD). WITHOUT PRECISE LOCALITY: Central Madagascar, Jan. 1882 (fr), Baron 1144 (K, P, syntypes of T. baroni); (fl), Baron 2417 (K, P); 1883 (fl), Baron 2477 (K). Tambourissa hildebrandtii belongs to species Group 1B and is distinguishable from other members of the group, notably its closest ally, T. nosybensis, by its strongly papillose styles, fruiting receptacle with a very small to almost nonexistent central cavity, and its villous stems. The type of T. baroni has fewer flowers and smaller, more obtuse leaves than that of T. hil- debrandtii, but easily falls within the range of variation displayed by the species. A comparable degree of variation in leaf size and shape fre- quently occurs on a single individual (e.g., Lo- rence 2043, MAD, MAU, M Fresh flowers of Tambourissa hildebrandtii are externally grayish green to peach colored. The gynoecious flowers have purple styles, whereas the androecious flowers are beefy red within and produce a sweet, fruity odor. In mixed inflores- cences, anthesis occurred in gynoecious flowers before it occurred in androecious flowers (Lor- ence 2043 Vernacular names. Ambora, Ambora saha (Madagascar). 6. Tambourissa nosybensis Lorence, Bull. Mus. Hist. Nat. (Paris), Ser. 4, Sect. B, Adansonia 3(3): 302, pl. 3, figs. 12, 13. 1982. TYPE: Madagascar. Diego Suarez: Nosy Be Island (Nossi-Bé); Lokobe, Natural Reserve no. 6, 350 m, 12 Dec. 1967 (fl), Bernardi 11847 (holotype, G; isotypes, K, P, Z). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Monoecious to subdioecious tree 6 m tall, the bark pale gray, licheniferous, longitudinally fis- sured, the new growth pilose with short, fulvous hairs, the mature leafy stems terete, 2-3.5 mm diam., brown, pilosulous. Leaves opposite, pet- iolate; petioles 6-11 mm by 1-1.2 mm, pilose- pubescent; mature lamina adaxially glabrescent, abaxially sparsely hirsutulous, especially alon , discolorous, o long, occasionally + falcate, 60-125 mm by 20- 6 mm, the apex acuminate, the base acutely cuneate, the secondary veins 4-6 pairs, making a 35-45? angle with the costa, the intersecondary and tertiary branches straight with strong ad- medial orientation, + perpendicular to the costa, the venation visible to 2(-3?) adaxially, and to 3(-4°) abaxially, the margin slightly revolute, thickened. Inflorescence cauliflorous, moderate- ly to densely fulvous velutinous, a unisexual or rarely sexually mixed pleiochasium of 9-12 flow- ers, the floral axis 50-87 mm by 2 mm, the ped- iceland peduncle 7-20 mm by 1-2 mm, the joint swollen, subtended by a caducous, subulate-na- viculate bracteole 1-2 mm long. Androecious flower in bud unknown; at anthesis deeply 5-fid, 17 mm diam., the lobes thick, spreading flat, the receptacle externally hirsute; stamens ca. 70, in- curved, ovoid-acuminate, subsessile, 1.5-2 mm long by 0.7-1 mm wide, the loculi lateral, sep- arate, the connective acute, prolonged for ca. 0.5 mm, the internal receptacle surface with rare, scattered hairs among the stamens; pedicel ca. 2 mm diam. Gynoecious flower in bud discoid-pa- telliform, 9-10 mm diam. by 3-4 mm long, lon- gitudinally multi-ribbed, velutinous, the apex depressed, with 4 subulate tepals ca. 1 mm long, the pedicel 9-20 mm by 1-1.2 mm; at anthesis splitting apically into 4(—6) short, straight or in- curved deltoid lobes, the orifice small, X-shaped; styles numerous, ca. 400, broadly conical to co- lumnar, apiculate, 0.4—0.6 mm long by 0.4-0.5 mm diam. basally, not papillose, the internal re- ceptacle surface velutinous between the styles. Fruiting receptacle unknown. Figure 28 Distribution. Known only from Nosy Be Is- land off the northwestern coast of Madagascar (Fig. 29). Habitat. Tambourissa nosybensis was col- lected in depauperate lower montane wet forest (presumably of Chlaenaceae, Myristicaceae, and Anthostema) at 350 m 1985] LORENCE—MONIMIACEAE 107 o CE b CE RM ERUNT O RE: G: OL O O OO . FicuRE 28. Tambourissa E dan Lorence.— A. Pai — B. Androecious bd i at E ed AE abaxial view.— D. Gynoecious inflorescence at anthesis.—E. Gyn oecious flower at anthes Gynoecious flower at AE longitudinal section. ir Styles, ong todinal Lb. All Mende 11 1847 7 d Bars equal 10 mm in A, B, D, E, and ] mm in C, F, G. 108 200 km 4 (C ` Í y J i ^ { e / ae pur + 14°s wA | 21 ( (^S À \ "4 rt N a = ( V 4 \ f À < \ ) Q C l \ í I i ° A AA x M ) | i r A / E ~~ FP | V^ ] O j F [| O T. decaryana ^ T. floricostata B T. humbertii J @ T. nosybensis Vi A T. parvifolia . ç FiGuRE 29. Dist Vii. map of some Tambou- rissa kind in Madagasc GASCAR. WITHOUT PRECISE LOCALITY: ca. 1883 m Humbolt 642 (K, P). Tambourissa nosybensis belongs to species G 1B. Although most closely allied to T. hildebrandtii (for which Bernardi mistook it), T. These include a number of specialized features, notably the production of a mucilaginous exu- date in the mature gynoecious flowers (conspic- uous in rehydrated flowers), and also an apparent trend towards dioecy. I first thought the species was dioecious but then found a single androe- cious flower among some 34 gynoecious flowers, suggesting it is monoecious. Further collections, particularly of androecious flowers and fruit, and observations on the biology of this interesting species are required. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 7. Tambourissa decaryana Cavaco, Bull. Mus. Hist. Nat. (Paris), Ser. 2, 29: 287. 1957; Ca- vaco in Humbert, Fl. Madagascar 80: 22, fig. VI:1-4. ae TYPE: Madagascar. Ta- matave: Mt. Vatovavy (Mananjary), 22 Nov. 1938 (fl, fr), aoe 13697 (holotype, P). Cauliflorous tree 10-16 m tall, possibly dioe- cious (?), the new growth pilose-strigillose, the adult leafy stems terete, pilose, 4-5 mm diam Leaves opposite, petiolate; petioles 15-20 mm 150-325 mm by 85-115 mm, adaxially glabrous, abaxially strigillose with white, ap- pressed hairs, the apex acuminate, the base ob- tuse to rounded, the secondary veins 6-14 pairs, making a 60-70? angle with the costa, the ve- nation prominent, visible to 4? on both surfaces, the margin slightly revolute. Androecious flow- ers unknown. Gynoecious inflorescence cauliflo- rous, a fulvous, tomentose or velutinous pleio- chasium of 6-11 flowers, the floral axis (10-)30- 135 mm by 1.5-3 mm, subtended basally by several deltoid-naviculate bracteoles ca. 1 mm long, the pedicel 6-23 mm by 1-1.5 mm, sub- tended by a velutinous subulate bracteole (1—)4— 5 mm long. Gynoecious flowers in bud napi- form-depressed, 7-8 mm diam. by 4-5 mm long, xternally tomentose to velutinous, the apex apiculate with 4-5 minute velutinous tepals; at anthesis 8-11 mm diam. by 7-8 mm long, the orifice shallowly 4-5-fid, comprising '2—% total width of the receptacle, the lobes suberect, the margins thickened; styles numerous, ca. 200-300, shortly conical, 0.4-0.5 mm long by 0.2-0.3 mm wide, basally + ventricose, the apex acute, slight- ly papillose, the ovule amphitropous; receptacle lobes bearing internally several series of larger, sterile, hirsutulous conical appendages (ca. 25- 40), these 1-1.5 mm long by 0.5-0.8 mm diam., the internal receptacle surface densely velutinous between the styles and on the lobes. Fruiting re- ceptacle produced on the trunk and branches, cupuliform, 40-50 mm long by 50-70 mm diam., externally corky, the orifice comprising '4—"4 to- tal width of fruit, the styles conical, 0.5 mm long, interspersed with hairs; pedicel and peduncle 37— 110 mm by 6-9 mm Distribution. Endemic to Madagascar (Fig. 9). Habitat. Tambourissa decaryana occurs in the lower altitude Myristicaceae/Anthostema wet forest zone in the eastern domain, from ca. 250 to 1985] TABLE 15. LORENCE—MONIMIACEAE 109 Some distinguishing morphological features of Tambourissa hildebrandtii and T. nosybensis. Character T. hildebrandtii T. nosybensis Angle of secondary veins costa Intersecondary veins and ertiary branches Inflorescence position 55-80° zig-zag, weak admedial orientation ramiflorous or axillary 35-45? straight, strong admedial orientation cauliflorous Inflorescence pubescence Sta mens Loculi unilateral Connective short, obtuse, or absent Gynoecious receptacle nally ri Orifice lobes 4, suberect to everted Styles strongly papillose Ovule anatropous, pendulous Mucilaginous exudate absent Tanniferous cells present Stone cells absent cinereous, hirsute, or tomentose subglobose- -napiform, not longitudi- narrowly conical, 1-1.5 mm long, fulvous, velutinous bilateral acute, 0.5 mm long discoid-patelliform, longitudinally ribbed 4(—6), incurved or strai broadly conical to columnar, apiculate, 0. mm long, not papillose ¿mnhityoppus, obliqu present, covering is present ig etes TAMATAVE: Mt. Vat n, 250 m, tovavy, Manan- Aug. 1911 (fr), Perrier de la Báthie 1957 (fr), Capuron 18111-SF (P). WITHOUT PRECISE LOCALITY: Feb. 1881 (fl), Parker s.n. (K). Parker s.n. differs from the type of Tambouris- sa decaryana in having a longer, less densely pu- bescent floral axis, and furthermore has mature gynoecious flowers (Decary 13697 is mostly in bud). Although Parker s.n. lacks leaves, the pres- ence of large sterile appendages within the flow- ers, an abundance of oil cells in the receptacular ground tissue, amphitropous ovules, and pro- duction of mucilage by the styles all correspond well with the type of 7. decaryana. The species is characterized by a moderately to densely pubescent inflorescence, long brac- teoles, 4—5-fid gynoecious flowers with an X- shaped orifice and suberect lobes, and sparsely pubescent leaves, characters which place it in species Group 1B. Its huge, macrophyllous leaves (assuming they are not from a vigorous sucker shoot) and presence of several rows of large, style- like sterile appendages surrounding the styles set it apart from all other Madagascan species, al- though Tambourissa parvifolia has similar ap- pendages. The sterile appendages in 7. decary- ana resemble the styles but are 2-4 times larger, hirsutulous, and lack an ovule at the base. Nu- merous oil cells occur in the receptacular ground tissue. The species also displays certain special- ized features shared with T. nosybensis, i.e., pro- duction of a mucilaginous exudate and a ten- dency towards dioecy. 8. Tambourissa humbertii Cavaco, Bull. Soc. Bot. France 104: 283. 1957; Cavaco in Humbert, Fl. Madagascar 80: 28, fig. IX:1—-4 (non fig. VIII). 1959. Type: Madagascar. Fianarant- soa: high valley of the Rienana River, Ma- titanana Basin, 18-24 Nov. 1924 (fl), Hum- bert 3593 (holotype, P; isotype P). Large tree, the new growth velutinous, the ma- ture leafy stems flattened, stramineate, 3.5-6 mm diam., sparsely hirsutulous, glabrescent. Leaves opposite, petiolate; petioles hirsutulous, glabres- cent, 10-13 mm by 1.5-2 mm; lamina glabrous, subcoriaceous, elliptic to oblong, 82-126 mm by 35-60 mm, the apex shortly acute, the base acute, slightly decurrent, the secondary veins 6—9 pairs, making a 60-70° angle with the costa, the ve- nation visible to 3° on both surfaces, the margin revolute, the adaxial laminar surface moderately cauliflorous, a fascicle of 2-3 flowers, or a short, 5-17-flowered pleiochasium, the floral axis stout, 30-50 mm long, lenticellar, canescent, the ped- icel velutinous-canescent, jointed below the middle, the pedicel and peduncle 12-20 mm by -3 mm, with 1 to several velutinous, del- toid-naviculate basal bracteoles 1-2 mm long. Gynoecious flower in bud canescent, broadly 110 napiform-depressed, 13-15 mm diam. by 5 mm long, apiculate with several minute, coalescent velutinous tepals; at anthesis broadly napiform- discoid to patelliform, 18-21 mm diam. by 6-8 mm long, the orifice X-shaped, ca. 10 mm diam., splitting for ca. '^ the length ofthe receptacle into 4(—5) suberect, fleshy deltoid lobes, the margins thickened; styles numerous, ca. 500 or more, densely crowded, ovoid-ventricose, 1-1.2 mm long by 0.5-0.7 mm wide basally, the apex short- ly acuminate or apiculate, papillose, the surface bearing scattered simple hairs ca. 1-1.5 mm long, the internal receptacle surface velutinous be- tween the styles and on the lobes the numerous short hairs mixed with scattered long hairs. Fruit unknown. Distribution. Endemic to Madagascar (Fig. ). Habitat. Tambourissa humbertii is known only from the type collection, presumably from the Tambourissa/ Weinmannia montane wet for- est zone from ca. 1,000 to 1,400 m. Further col- lections of this striking species, particularly an- droecious flowers and fruit, and observations on its life history are needed. In contrast to Cavaco’s (1957b) statement that Tambourissa humbertii approaches T. ficus from Mauritius, I find that it differs from the latter species in several important characters. Most no- tably it lacks the ternate-whorled to subalternate leaves and tanniferous idioblasts in the floral ground tissue characteristic of 7. ficus. Tam- bourissa humbertii appears rather to belong to species Group 1B and is distinguishable from other members of the group by its much larger gynoecious flowers and presence of long hairs on the styles. Styles of 7. kirkii (Group 2B; Co- mores), incidentally, bear similar but shorter scattered hairs. In addition, the internal surface of the gynoecious receptacle of T. humbertii bears mixed long and short hairs, a feature found in no other species. In spite of their relatively large size, gynoe- cious flowers of Tambourissa humbertii struc- turally most closely resemble those of 7. decary- ana. In both species the orifice lobe margins are conspicuously thickened and numerous oil cells occur in the ground tissue. Flowers of 7. hum- bertii are much larger, however, and lack the peculiar sterile appendages characteristic of the latter species. 9. Tambourissa floricostata Cavaco, Bull. Soc. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Bot. France 105: 39. 1958; Cavaco in Hum- ve: lamazaotra forest, Feb. 1919 (fl), Raman- antoavolana sub T. Thouvenot 134 (holo- type, P; isotypes, P, 2 sheets). A madagascariensis Danguy, Bull. Mus. Nat. (Paris), Ser. 1, 28: 249. 1922 non Tam- ea madagascariensis Cavaco, 1957. Monoecious tree 18-20 m tall, the new growth unknown, the mature growth subglabrous, the mature leafy stems terete, 1.5-2 mm diam. Leaves opposite to subopposite, petiolate; petioles 7-13 mm by 0.8-1 mm; lamina subcoriaceous to co- riaceous, glabrous, elliptic to obovate, rarely ob- long, d slightly falcate basally, 35-85 mm b m, the apex obtuse to retuse, its extremity Meus thickened or apiculate, the base acutely cuneate to acutely decurrent, the second- ary veins 4—7 pairs, making a 55—60? angle with the costa, the venation visible to 3? adaxially and to 4? abaxially, the margin thickened, slightly revolute. Inflorescence ramiflorous, a puberu- lent, sexually mixed pleiochasium of (3-)5-9 flowers attaining 120-160 mm by 60-70 mm, axillary or on leafless nodes, or the flowers sol- itary and axillary or terminal, the floral axis (17-)23-140 mm by 1-3 mm, + flattened, often leafy. Androecious flower in bud quadrangular- globose, 9—12 mm diam., with 2-4 prominent longitudinal ridges, apiculate with 4 deltoid- rounded hirsute tepals; pedicels 16-30 mm by 1-1.5 mm, flattened, puberulent, each subtended by a naviculate-deltoid bracteole; at anthesis 4-fid, 25 mm diam., splitting to ca. !^—/ total length of the receptacle, the lobes incurved, not spread- ing, each lobe with ca. 2 fleshy, obtuse scale-like tepals internally near the apex; stamens 16-22, large, ovoid to deltoid, acute, 3-4.5 mm long by 1.5-2.5 mm wide, the loculi lateral, free, the con- nective broad, + sessile, the apex apiculate, acute, iam. by 5-7 mm long in bud, with 2 lateral ridges, the orifice small, 1-2 mm diam., flanked by 4 obtuse, decussate tepals; pedicels to 55 mm by 2 mm. Immature fruiting receptacle ca. 47 mm diam., broadly na- piform-depressed, the orifice ca. 5 mm diam., subentire, the external surface corky brown, the internal surface with numerous scattered, shortly conical styles ca. 0.5-1 mm long by 0.5 mm wide LORENCE-— MONIMIACEAE 111 1985] basally, the surface between the styles hirsute. Submature fruiting receptacle solitary on leafless 57 mm diam., externally venose, corky, the ped- icel and peduncle tapered, 120-150 mm by 4-5 mm; mature carpels unknown. Distribution. Endemic to Madagascar (Fig. Habitat. Known only from three collections, all from the Perinet area (Analamazaotra), in the Tambourissa/ Weinmannia wet forest zone from ca. 900 to 950 m. Thouvenot 134 was collected in February with flowers and young fruit. As the Analamazaotra forest is included in a nature re- serve, it is probable that 7. floricostata still oc- curs there; much intact forest still remained in 1978. However, additional flowering and fruiting collections and data on its life history are needed. MADAGASCAR. TAMATAVE: Analamazaotra, Feb. 1912 (seed), Perrier de la Báthie 10128 (P); Zedsiadra (or Djedsiatsa?) near Perinet (st), Sched! 40 (MAD) In spite of the large size and small number of stamens in androecious flowers of Tambourissa dictum the species possesses a number o attributes which suggest inclusion in species dm pol along with 7. parvifolia (Table 5). It differs from the latter species by its larger leaves, flowers, and receptacles with two to four prom- inent external longitudinal ridges. Vernacular names. Ambora, Ambora lahy, Ambora vavy (Madagascar). 10. Tambourissa parvifolia Baker, J. Bot. n.s. 11: 267.1882; Baillon, Bull. Mens. Soc. Linn. Paris 1: 342. 1882; Perk. & Gilg, Pflanzenr. 4, 101: 70. 1901; Perk., Pflanzenr. 4, 101 (Nachtr.): 42. 1911; Cavaco in Humbert, FI. Madagascar 80: 36, fig. X:1—4. 1959. TYPE: Madagascar. Without precise locality (“Central Madagascar," probably Tanana- rive): 1880 (fl), Parker s.n. (holotype, K). T. quadrifida sensu Drake in Grandidier, Hist. Phys. Madagascar 1(1): 21. 1902, pro parte, non Sonn. Monoecious treelet or tree 5—15 m tall, the new growth Tr gi gps. glabrescent, the ma- ture leafy stems 1.5-3. m diam. Leaves op- posite, eleme petioles 3 5 mm by 0.7-1 mm; lamina glabrous, coriaceous, elliptic to oblong or obovate, 15-47 mm by 9-22 mm, the apex ob- tuse to retuse, the base acutely cuneate to obtuse, slightly decurrent, the secondary veins 4-5 pairs, making a 55-60? angle with the costa, the ve- nation obscure adaxially or the 2? veins scarcely visible, abaxially visible to 3°, the margin revo- lute. Inflorescence ramiflorous on leafless stems, an (always?) unisexual, hirtellous-puberulent pleiochasium of 2-11 flowers, the floral axis 12- 70 mm by 1-2 mm, basally bracteolate. An- droecious flower in bud globose, 5-6 mm diam., apiculate with several minute, coalescent tepals, the pedicel 6-10 mm by 1-2 mm, subtended by a deltoid, hirsute bracteole 2 mm long by 1 mm wide; at anthesis deeply 4-fid, 17-30 mm diam., the lobes spreading flat, the internal receptacle surface bearing several sterile, columnar ap- pendages near the apex of the lobes; stamens ca. 50-60, + ligulate, 1-1.5 mm long by 0.6-0.8 mm wide, the loculi lateral, free, parallel, comprising ca. 3⁄4 total length of the stamen, the connective not or only slightly prolonged, the internal re- ceptacle surface glabrous. Gynoecious receptacle at anthesis subglobose to napiform, ca. 9-10 mm diam., externally hirsutulous-pubescent, the ori- fice small, 4-lobed, X-shaped, comprising ca. “4 total width of the receptacle, the pedicel 7-14 mm long; styles numerous, ca. 200 or more, con- ical, ca. 0.5 mm long by 0.5 mm wide basally. Fruiting receptacle solitary or in pleiochasia of 2 or more, > the axis to 60 mm by 5mm, hirtellous, th llowly cupuliform to napiform- doses 30-95 mm wide by 25-40 mm long, the orifice Para. ca. %—' total width of the receptacle, 8-12 mm wide, subentire with several pale corky patches, glabrous or sparsely hirsu- tulous, the internal surface with numerous con- ical styles 0.3-0.5 mm long and wide basally with scattered simple hairs in between, the pedicel 10— 22 mm by 3.5-6 mm. Carpels 5-13 mm long by 5-9 mm wide by 3-4 mm thick, ovoid-com- pressed, the endocarp tan, the surface scrobicu- late Distribution. Endemic to Madagascar (Fig. 29). Habitat. Tambourissa parvifolia is restricted to the sclerophyllous montane forest zone of the central plateau at elevations from ca. 1,400 to 2,000 m. i AP beca E of Antsirabe, 2,000 d ov. 1928 (fl), Humbert 6351bis (P); E slope s eee Aun Mar. 1958 (fl), Capuron 18507- 112 SF (P). FIANARANTSOA: Andringitra, Imaisoa, 1,500- 2,000 m, 28 Oct. 1960 (fl), Leandri et al. 3393 (P). WITHOUT PRECISE LOCALITY: Central Madagascar, 1855 (fr), Baron 3864 (BM, P); (st), Baron 4204 (P); (fr), Baron 4205 (K); (fr), Baron 6048 (P); (fl), Parker s.n. (P, 5 sheets The sclerophyllous leaves of Tambourissa parvifolia are smallest of any known species of Tambourissa and probably represent an adap- tation to the high altitude montane habitat. In adaga dense sclerophylle de montagne" (Koechlin et al., 1974) In spite of its extremely reduced leaves, Tam- bourissa parvifolia is best placed in species Group 1C along with T. floricostata. It displays a num- ber of specialized trends, including reduced leaves, a tendency towards unisexual inflores- cences (and possibly dioecy) and the presence of sterile, b dn like dta on me internal sur- es with T. d euni DRE RANA parvifolia is easily distinguished from 7. floricostata by its small, obutse or emarginate leaves and slender anther filaments. rnacular names. Ambora, Amboravato Price scar 11. Tambourissa beanjadensis Lorence, nom. nov. Ephippiandra capuronii Cavaco, Bull. Soc. Bot. France 104: 610. 1957; Cavaco in Humbert, Fl. Madagascar 80: 8, fig. I:10- 12. 1959 non Tambourissa capuronii Ca- Suarez: Beanjada Massif (N of the Masoala Peninsula), E forest at its upper limits (“forêt des cimes"), ca. 1,100 m, 30 Dec. 1953 (fl), Capuron 8826 bis-SF (holotype, P). Medium-sized monoecious tree, the new growth with rare, scattered hairs, the mature lea- fy stems glabrous, brown, 1.5-2 mm diam. Leaves glabrous, opposite to subopposite, petiolate; pet- ioles 6-9 mm by 1 mm; lamina ch ous to subcoriaceous, adaxially lustrous, minutely punctate, elliptic, (23-)34-47 mm by (12-)18- mm, the apex abruptly and shortly acuminate, the tip thickened, the base cuneate to broadly cuneate, rarely obtuse, the secondary veins 3-5 pairs, making a 75-85? angle with the costa, adaxially scarcely visible, abaxially prominent, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 the venation visible to 3? abaxially, the margin entire, slightly revolute. Inflorescence ramiflo- rous or cauliflorous, solitary or clustered, an ap- parently unisexual pleiochasium of 3-7 flowers, or the flowers rarely solitary, the floral axis 20- 40 mm by 1 mm, pilose when young, glabrescent, subtended by several minute, naviculate brac- teoles 1 mm long, the pedicel 3-7 mm by 1 mm, subtended by a minute, obtuse naviculate brac- teole ca. 1 mm long. Androecious flower in bud globose, 4 mm diam., glabrous, the apex with 4 sparsely pilose obtuse tepals; at anthesis deeply (3-)4(-5)-fid, 8-10 mm diam., the lobes spread- ing flat, each bearing internally near the apex a small, scale-like tepal 1 mm wide; stamens ca. 35-36, obtuse to deltoid, 0.6-1 mm long and wide, the filament short or sessile, the loculi con- fluent apically into a single, crescentiform loculus occupying 73 or more of the total length of the stamen, the connective thick, not prolonged, the internal receptacle surface glabrous. Gynoecious flower in bud globose-napiform, ca. 6 mm diam. by 4-5 mm long, glabrous, the apex with 4 mi- nutely puberulent obtuse tepals; at anthesis split- ting into an X-shaped orifice ca. 1-2 mm wide, the 4 lobes deltoid, the internal receptacle surface with a pubescent ring near the orifice; styles nu- merous, ca. 200, shortly and broadly conical, 0.3-0.4 mm long and wide, papillose, producing a mucilaginous exudate, the internal receptacle surface with scattered hairs between the styles, the receptacle wall 1 mm thick. Fruiting recep- tacle unknown. Distribution. Endemic to Madagascar (Fig. Habitat. The species is known only from the type collection from the upper limits of what is presumably mid altitude Tambourissa/ Wein- mannia wet forest on mountainous summits. Further collections and field observations are ob- viously needed. Features which immediately place this species in the genus Tambourissa are the closed, globose gynoecious floral receptacle with an X-shaped orifice and the conical carpels completely im- mersed in and fused with the receptacle wall. The broadly deltoid, sessile stamens with a single, crescentiform loculus and presence of tepals on the inner surface of the androecious receptacle lobes occur in a number of other Tambourissa species, as previously noted. Pollen of the species, figured by Walker (1976) and presumably taken LORENCE— MONIMIACEAE 113 1985] from the type, also conforms to that of Tam- bourissa rather than that of Ephippiandra. The small leaves and ramiflorous to cauliflo- rous, pleiochasial inflorescence of Tambourissa beanjadensis resemble those of 7. parvifolia, which differs by its more coriaceous, obtuse leaves, much larger androecious flowers, and longer stamens with distinct filaments and sep- arate loculi. Tambourissa beanjadensis is prob- ably best placed in species Group 2A, however, a diverse series of species from Madagascar, Ré- union, and the Comores characterized by their deeply 4-fid androecious flowers which spread (T. crassa and T. elliptica) are globose-depressed, open by an X-shaped orifice, are nearly glabrous within, and produce a Both latter species are easily distinguished by their much larger leaves, obtusely columnar styles, and stamens with separate thecae. 12. Tambourissa madagascariensis Cavaco, Bull. Soc ural Reserve no. 4 (Tsaratanana, Ambanja), Mangindrano Canton, Bealanana District, 7 July 1953 (fl), Rababoto sub R.N. 6028 (ho- lotype, P). Dioecious (?) tree 5-15 m tall, or a lianescent shrub, the new growth with scattered hairs, gla- brescent, the mature leafy stems glabrous, 2-3 mm diam., with longitudinal striations. Leaves glabrous, opposite to subopposite, petiolate; pet- ioles 4-7 mm by 1-1.5 mm; lamina chartaceous, broadly elliptic, 42-80 mm by 24-34 mm, the apex acuminate, the tip + indurated, the base acutely or rarely obtusely decurrent, the second- ary veins 5-7 dn making a 60—70? angle with ion obscure, visible to 2° y, the margin plane to slightly revolute. ons ramiflorous, axillary or on leafless nodes, a subglabrous uni- sexual pleiochasium of 3-9 flowers, sometimes leafy in the terminal portion, the floral axis 20— 85 mm by 1-2 mm, longitudinally striate, sparse- ly puberulent around the pedicels, subtended by several broadly deltoid, ciliate bracteoles. An- droecious flower in bud globose, 6-7 mm diam., + glabrous, apiculate, the tepals fused, the ped- icel 8-23 mm by 0.6-1 mm, glabrous or sparsely 200 km : PN < V S wy / ` 4dE RS cv 14 S k T m ° Ay, l4 \ Pd x rha Vi ài = 7 f ) TAS ( ) Y, wm Š ) b ) ) ( Y / ) > ^ a I O i p MM y | P o ne ) í 1 i i M b / V ^ | | / f ) j Ça w sa ai < de / / f a E j / N / / f / _ f N O T. bathiei 1 A T. beanjadensis k dri A T. castri-delphinii @ T. madagascariensis ` š B T. perrieri FiGuRE 30. Di = map of some Tambou- rissa c in Madagasca puberulent basally, subtended by a deltoid-subu- late, ciliate bracteole 0.5-2.5 m ceolate to subulate, 2.5-4 mm long by 0.7-1 wide, sessile or with a short filament, the loculi lateral, separate, occupying almost the entire length of the stamen, the connective slightly pro- longed, obtusely apiculate, the internal recepta- cle surface glabrous. Gynoecious flower in bud subglobose, slightly depressed apically, 4-5 mm diam., externally glabrous to sparsely hirsutu- lous; at anthesis (?) T mm diam., the orifice circular, entire, 2-3 m m., comprising ca. 12 total width of the hs dara styles numerous, several hundred, shortly columnar, 0.2-0.3 mm long, the apex rounded, the ‘eternal. receptacle surface puberulent around the orifice and be- 114 tween the styles, the pedicel 6-8 mm long, short- erthan in the androecious flower. Mature fruiting receptacle cupuliform, attaining 150 mm diam. (fide notes on Anon. sub SF 2126, MAD); carpels unknown. Distribution. Endemic to Madagascar (Fig. ). Habitat. The type of Tambourissa mada- gascariensis was collected on the Tsaratanana lections of this species are from mo bourissa/ Weinmannia wet forest (800-1,000 m), and Myristicaceae/Anthostema wet forest (ca. 400-800 m) in the eastern region. This seemingly disjunct distribution pattern may be the result of undercollecting. MADAGASCAR. TAMATAVE: Mt. Ambatosoratra, 400 . 1949 (fl), Anon. sub Herb. Alaotra 3247 Kon EE 29 Dec. 1944 (fr), Anon. sub Herb. Alaotra 2142 (MAD Tambourissa madagascariensis is anomalous in a number of respects and does not readily fit into any of the more or less natural groups of Madagascan species, but is best placed in species Group 2A. Its small, thin glabrous leaves are reminiscent of T. purpurea and T. gracilis, but the flowers are borne in long, slender subglabrous and long-pedicelled pleiochasia that are more suggestive of 7. gracilis, T. beanjadensis, and T. parvifolia. Androecious flowers of T. madagas- cariensis are relatively large, however, and split open into four lobes which spread flat and bear numerous (ca. 200) long stamens with separate, parallel loculi, thus differing strikingly from the small, globular, nearly closed androecious flow- ers of T. gracilis and T. purpurea, which have short, sessile stamens with apically confluent loc- uli Gynoecious flowers of Tambourissa mada- gascariensis are small and globose-depressed with an entire, circular orifice that is internally pu- berulent, and numerous short, columnar-round- ed styles suggestive of those species which pro- duce copious mucilage at anthesis. The orifice apparently remains open throughout much ofthe floral development, thus differing markedly from the gynoecious flowers of 7T. beanjadensis and T. parvifolia (unknown in T. gracilis), which open ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 by an X-shaped orifice at anthesis and, further- more have conical styles. In terms of both androecious and gynoecious floral structure, Tambourissa madagascariensis appears to be most closely related to two of the Comorean species in Group 2A, T. comorensis and 7. leptophylla, although their leaves and flowers are much larger. Presence of abundant oil cells in the gynoecious floral ground tissue of all three species and large fruits are common features. This group of Comorean species could have been derived from a common ancestor sim- ilar to T. madagascariensis. Tambourissa madagascariensis also appears to share characters of androecious and gynoe- cious floral morphology with 7. elliptica subsp. micrantha from Réunion, notably the pleiocha- sial inflorescence, similar androecious flowers and stamens, and gynoecious flowers with short, co- lumnar styles and a mucilaginous exudate. Gy- noecious flowers of the latter species lack the open, circular orifice, however. Vernacular names. Ambora, Mahimasina, Valiramarnia (Madagascar). 13. Tambourissa elliptica (Tul.) A.DC., Prodr. 16(2): 660. 1868; Cordem., Fl. Réunion 301. 1895; Perk. & Gilg, Pflanzenr. 4, 101: 71. 1901. Ambora elliptica Tul., Ann. Sci. Nat. (Paris) 4(3): 31. 1855; Tul., Monogr. Mo- nim. 304. 1855. TYPE: Réunion. Woods of the Hauts de la Riviére des Galets, July 1851 (fr), Boivin s.n. (holotype, Tambourissa amplifolia sensu Cordem., Fl. Réunion 301. 1895 non (Boj. ex Tul.) A.DC. Treelet or tree of variable habit, attaining 12 m tall, the new growth glabrous or with sparse, scattered hairs, the mature leafy stems terete, 2— 6(-8) mm diam. Leaves glabrous, opposite to subopposite, petiolate; Roni (810-25 mm by 1.5-5 mm; lamina chartaceous to coriaceous, narrowly to broadly PR SIE obovate-el- liptic, obovate, broadl ovate or orbiculate, (50-)80-180(-250) mm s (15-)25-140(-190) mm, the apex shortly acuminate, acute, obtuse or rarely rounded, the base acutely decurrent, acutely cuneate to obtuse, truncate or subcor- date, the secondary veins (5-)6-12 pairs, making a 55-75? angle with the costa, the venation prom- inent, raised, visible to 3—4(—5°) adaxially, to 4— 5(—6?) abaxially, the margin plane to slightly rev- olute. Leaves of seedlings and suckers hetero- 1985] LORENCE— MONIMIACEAE 115 phyllous, small and apically dentate in subsp. elliptica, long, narrow and entire in subsp. mi- crantha. ,ashort rami- florous pleiochasium (in subsp. micrantha), or the flowers solitary, sometimes axillary (in both subspecies), or cauliflorous and the flowers sol- itary or in fascicles of 2-3 (in subsp. sey aes Androecious flower in bud globose, 7-15 m diam., apiculate with 4 minute, obtuse ae the d 10-40(-60) mm by 1-1.5 mm; at anthesis (3-4(-5)-fid, 16-45 mm diam., the lobes spreading flat; stamens numerous, 65-300, lig- ulate to linear-ellipsoid, 1.5-6.5 mm long by 0.5- 1 mm wide, the filament distinct, the loculi lat- eral, comprising 75—À total length of the stamen, separate or confluent at the obtuse apex, the con- nective not prolonged, the internal receptacle surface glabrous. Gynoecious flower in bud glo- bose-depressed to napiform-depressed, apiculate with several pairs of minute tepals, the pedicel (5—)10-20(-40) mm by 0.6-2 mm; at anthesis 6-15 mm diam., the orifice X-shaped, 1-5 mm wide, else into 4 shallowly deltoid lobes; . 150-200 or more, bluntly brous or with rare scattered hairs, p n mucilaginous exudate. Fruiting receptacle soli- tary or in clusters of 2-3 on the trunk (subsp. elliptica), or on the leafless branches (subsp. mi- crantha), urceolate-globose or cupuliform, 25- 90 mm diam. by 15-50 mm long, the orifice subentire, comprising ca. %—'4 total width of the m polyhedral styles 0.5 mm wide. Mature d ovoid, 9-12 mm long, 5-7 mm wide, thick, the endocarp whitish (subsp. pse or pale brown (subsp. micrantha). Distribution. Endemic to the wet and cloud forest zones of Réunion Island (Fig. 31). E. J. de Cordemoy recognized four species of Tambourissa in his “Flore del'Ile dela Réunion” (1895), but used names of a Madagascan (7. re- ligiosa) and a Mauritian (T. quadrifida) species for two endemic Réunion taxa that I have named T. crassa and T. elliptica subsp. micrantha re- spectively (Lorence, 1982). Tambourissa elliptica, the only previously de- scribed species from Réunion, is quite polymor- phic, but two modally distinct subspecies are dis- tinguishable both by morphology, and, to a lesser degree, altitudinal distribution. Both subspecies with pink or pale yellowish white flower color morphs (never dark purple as is 7. crassa), caulinary flowers, conspicuous higher or- der venation, contain stone cells in the floral ground tissue, and have a less well-developed hypodermis than in 7. crassa, the second Ré- union species. Sı ibspeci es ists of a series of most- ly mid to upper elevation forms which Corde- moy considered to represent two distinct species (T. elliptica and T. amplifolia, respectively), based on leaf characters which prove to be variable. Subspecies micrantha usually occurs at lower el- evations, particularly in the southeastern part of the island where the most lowland wet forest still remains. Each subspecies tends to occupy its own alti- tudinal zone throughout much of Réunion (Fig. 31), at least in higher altitude wet and cloud for- est, significant portions of which are still pre- served due to the island's accentuated relief and steep, inaccessible slopes largely unsuitable for cultivation. A thorough distributional analysis is not possible for the lowlands, however, because little natural wet forest habitat remains there. KEY TO THE SUBSPECIES OF TAMBOURISSA ELLIPTICA la. Flowers large, usually solitary or fasciculate, LA : & | 1 š 1. A 13-14 mm diam.; expanded ue flowers 30-45 mm diam., the stamens ca. (100-)200-300 in number, ivan mm long; gynoecious flowers 10-12 diam.; es; high elevations (1,000-2,000 m, rarely low er) ubsp. eliptica lb. Flowers smaller, usually in pleiochas illary; androecious buds 8-1 0 mm diam.; ex “tsa a diam.; large trees with Wein laws (200-600 m, d higher 13b. subsp. micrantha 13a. Tambourissa elliptica subsp. elliptica. Monoecious or subdioecious treelet or small tree attaining 6 m tall and 25 cm D.B.H., bark soft, corky, pale brown, the new growth glabrous, rarely with a few scattered hairs, the mature leafy stems terete, 3-6(-8) mm diam. av abrous, opposite to subopposite, peti- eco pes 10-25 mm by 2-5 mm; lamina subcoriaceous to coriaceous, elliptic to broadly 116 -F 21920's 55920'E FIGURE 31. elliptic, obovate to broadly obovate or orbicu- late, 100-180(-250) mm by 60-140(-190) mm, the apex acute, rarely acuminate, obtuse or re- tuse, the base acute to obtuse, cuneate to decur- rent, truncate or subcordate, the secondary veins 6-8 pairs, making a 55-75? angle with the costa, the venation prominent, visible to 3(—4?) adax- ially and to 4(—5?) abaxially, the margin slightly revolute; juvenile and sucker leaves heterophyl- lous, with 1—2 pairs of glandular teeth in the apical portion ofthe lamina, the petiole red when living. Inflorescence glabrous or rarely with scat- tered hairs, the flowers solitary or in fascicles of —3 on meristematic swellings along the trunk, or the flowers ramiflorous and usually solitary, either axillary or on leafless nodes. Androecious flower in bud globose, (812-15 mm diam., apiculate with several + coalescent pairs of mi- nute, glabrous tepals, the pedicel 20-40(-60) mm by 1-1.5 mm, jointed near the base with the short peduncle, subtended by several pairs of small, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 10 NERO E, @ T. crassa T. elliptica Ñ A subsp. elliptica ^ À subsp. micrantha (x intermediate colls. Distribution map of Tambourissa in Réunion. deltoid, decussate ciliate bracteoles; at anthesis 4(-5)-fid, 30-45 mm diam., the lobes spreading flat; stamens numerous, (100—)200-300, ellip- soid to linear, 3-5 mm long by 0.6-1 mm wide, the filament broad, distinct, comprising ca. '4— ^ total length of the stamen, the loculi subpar- allel, lateral, connivent or confluent at the obtuse apex, comprising 75—/ total length of the stamen, the connective not prolonged, the internal re- ceptacle surface glabrous, glaucous. Gynoecious flower in bud globose-depressed to napiform-de- pressed, apiculate with several pairs of minute, + coalescent tepals, the pedicel 5-10(-40) mm by 1.2-2 mm, jointed near the base with the short peduncle, subtended by several pairs of decussate bracteoles; at anthesis 8-15 mm diam. by 5-10 mm long, the orifice 4-fid, 2-5 mm diam., X-shaped with 4 short, straight to incurved del- toid lobes; styles numerous, ca. 200 or more, bluntly columnar, ca. 0.5 mm long by 0.2-0.5 mm wide, + polyhedral, the internal receptacle 1985] LORENCE—MONIMIACEAE 117 surface glabrous or with rare, scattered hairs. Fruiting receptacle solitary on the trunk or major branches, urceolate-globose, 45-90 mm diam. by 30-50 mm long, the orifice 10-30 mm diam., comprising '4-'4 total width of the receptacle, subentire with remnants ofthe lobes, the external surface light corky brown, the internal surface glabrous, corky, with numerous short, + poly- hedral styles ca. 0.5-1 mm wide by 0.2-0.5 mm long, the apex flat or depressed, the pedicel and peduncle 10-60 mm long by 8-10 mm diam. Mature carpels ovoid-compressed, 10-12 mm long, 5-6 mm wide, 3—4 mm thick, the endocarp whitish, the surface smooth, unsculptured. Ga- metic chromosome number 7 — Distribution. Endemic to Réunion (Fig. 31). Habitat. It ranges from lower montane wet forest at altitudes from ca. 200 to 1,100 m, to cloud forest at altitudes from ca. 1,100 to 2,000 m. RÉUNION. BÉBOUR: from Bébour to Casse de Taca- ,220 m, 18 Jan. 1975 (fl), Bernardi 15151 é 972 (fr), m, 23 Feb. 1979 SWF] Lorence 2431 (MAU, MO); (fl), : 600 m, July 1972 (fr), Friedmann 1891 (P). BRAS wen aa Oct. 1944 (st), n. (TL-R); along canal of Bras Jeanne, Oct. 1944 (st), Rivals s.n. (TL-R). BROLE DE ST. DENIS: Hauts du Brülé de St. Denis, Mar. 1943 (st), Rivals s.n. (TL- R); Ilet à Quinquina, near St. Denis, July 1945 (st), Rivals s.n. Io kE pr DE CILAOS: Mare à Joseph forest, 1,400 m, 25 Mar. 1967 (st), Cadet 664 bis (REU); 19 July 1975 (fl), Cadet va (REU); 22 Feb. 1975 (fl), Cadet 5062 (REU); Petit Matarum, 1,700 m, 22 Feb. 1975 (fl), hp iie (REU); R bank of Bras de Ben- join, road to s, 950 m, 3 Feb. 1968 (fl), Capuron 28002-SF (P, 2 sheets) Mare a Joseph above Cilaos, 975 iae Coode 4928 (K); (fl), Coode n cloud forest, 1,500-1,700 0 (MAU, MO); (fl), Lorence & Rolland 2441 (MAU, Mie (fi, fr), Lorence & Rolland 2442 (MAU, y; rum and Mare à Joseph, Aug. 1943 d Rivals : : SE Ilet des Salazes, Cilaos, 1,300 4 (fl), Rivals s.n. (TL-R); slopes of the d zt pied 1,900-2,000 m (fr), Rivals s.n. (TL- R); Mare à Joseph, 1,200 m (fr), Rivals s.n. (TL-R m. = E DE MAFAT E: Mafate, Aug. 1883 (st), Cordemoy n. (TL-R); Sentier des Scouts, from Le Belier to top of Galets kao ca. 1,800 m (fl), pude 4136 (K). MOKA: Moka (st), Rivals s. n. (TL-R); 20 Feb. 1945 (fl), pro de s.n. » (TL. R); Hauts de Moka at the ee 600 m, Apr. 1946 (fr), Rivals s.n. (TL-R). LA MONTA rande Montagne, 1,200 m, 22 Mar. 1968 (fr), Cadet 1291 (REU). NOUVELLE: La Nouvelle, near forestry sta- = tion, May 1949 (fl), Rivals s.n. (TL-R). PITON RAVINE MALHEUR: heights of Piton de la Ravine à Malheur, 900 m, May 1943 (st), Rivals s.n. (TL-R). PLAINE D’AF- FOUCHES: forest along path to Plaine d’Affouches, 900 m, May 1943 (st), Rivals s.n. (TL-R). ROCHE ECRITE: path to Roche Ecrite, 28 Feb. 1975 (fl), Coode 5000 c 2 sheets; REU); 5 Oct. 1938 (st), Rivals s.n. (TL- PLAINE DES CAFRES: Plaine des Cafres (st), Cordemoy s.n. p Piton Bleu, 26 pi: 1882 (st), Cordemoy LAINE ips CHICOTS: near Mamode Camp, 966 (st), Cadet 595 (REU). PLAINE om mainial line above Tévelave, along edge of] Plaine des Makes, Oct. 1943 (st), Rivals s.n. (TL- R); ravine of the Col a sae Rouge, 28 June 1967 (st), Cadet 1039 (REU). P E 1,200 m, 19 Feb. dig (fl), "Rivals s.n. (TL-R). PLAINE laine a pov 25 Feb. 1979 (fl), Lorence 2454 (MO): 13 Jan. 1973 (fl, fr), Cadet 4021 (REU). ssp ANN ve Sainte Anne, 1,200 m, 30 May 1974 (fr), Cadet 4714 a near Sainte Anne bridge (st), Cordemoy s.n. (TL-R). TAKA Taka- Ilet 4 Bananes, 600-1, 000 fr), Lorence & Rolland 2530 ( Cabot, slopes of the Ilet de Patience, 1,200-1,5 ; Grand Fond forest, Takamaka, 900 m, 23 J . WITHOUT PRECISE LOCALI 7 (f), Cae dichaud s.n. (P); (st), Rivals s.n. c E Both the upland and lowland forms of Tam- bourissa elliptica subsp. elliptica include indi- viduals with either pink or pale greenish to yel- lowish white floral color morphs (as does subsp. micrantha), but never dark purple as in T. crassa. oral odors of both 7. elliptica subsp. elliptica and T. crassa are strong and rancid-fruity with overtones of butyric acid, never pleasantly sweet- fruity as in subsp. micrantha. In addition, gy- noecious flowers of bot T. elliptica and also of T. crassa are morphologically similar and produce a mucilaginous exudate that partly fills the floral cup. Strong similarities in the floral construction of all three Réunion taxa suggest a fairly recent derivation from a single, common ancestor. Both species belong to Group 2A. Although upland and lowland forms of Tam- bourissa elliptica subsp. elliptica are quite dis- tinct in their extremes, they intergrade gradually and clinally. For exa ample, collections from Ta- kamaka (eastern Réunion) form a continuous se- ries ranging from lowland wet forest at 600—700 m elevation, to the lower limits of cloud forest at 900-1,000 m (Lorence & Rolland 2525-2530, all MO). Both forms are diploids with n = 19. Because they are not separable by any single con- heneriec 118 stant character, nor even by any constant con- stellation of characters, it is not possible to rec- ognize them formally as taxa. The type of subsp. elliptica from the Hauts de la Riviére des Galets (Boivin s.n., P) presents an intermediate form, y leh al though most closely PP Vernacular names. Bois de bombarde, Bois de bombarde à grandes feuilles (Réunion). 13b. Tambourissa elliptica subsp. micrantha Lorence, Bull. Mus. Hist. Nat. (Paris), Ser. 4, Sect. B, Adansonia 3(3): 298, pl. 2, fig. 8. 1982. TYPE: Réunion. Brülé de Baril: lower montane wet forest, ca. 300 m, 2 Mar. 1979 (fl), Lorence, Lorence & Cadet 2489 (holo- type, MO; isotypes, K, MAU, P, REU, Z). T. quadrifida sensu Cordem., Fl. Réunion 301. 1895 non Sonn. Monoecious canopy tree 10-12 m tall and 30 cm D.B.H., the bark soft, pale yellow-brown, corky and peeling, the new growth with sparse, short simple hairs, soon glabrous, the mature leafy stems 2-5 mm diam. (to 8 mm diam. on suckers), longitudinally striate. Leaves glabrous, opposite to subopposite, petiolate; petioles (8—) 10-20 mm by 1.5-2 mm; lamina chartaceous to subcoriaceous, elliptic, narrowly elliptic, oblong or obovate-elliptic, (50-)80-180(-220) mm by (15-)25-50(-65) mm, the apex shortly acumi- nate, shortly acute, obtuse or rarely rounded, the base acutely cuneate to acutely decurrent, the secondary veins (5-)7-12 pairs, making a dip: 5? angle with the costa, the venation prominent finely reticulate, visible to 4(—5?) adaxially and to 5(—6°) abaxially, the margin plane to slightly revolute; juvenile and sucker leaves heterophyl- lous, long and narrowly elliptic, the lamina at- taining 180 mm by 40 mm, the apex acute to acuminate, the secondary veins 10-14 pairs, the petiole and costa bright red when living. Inflo- rescence ramiflorous on old or new stems, or axillary, a unisexual or sexually mixed pleiocha- sium of 3-7 flowers, with sparse scattered hairs, the floral axis 8-20 mm by 1-1.5 mm, subtended by several minute, ciliate naviculate bracteoles 0.6-0.8 mm long, or the flowers rarely solitary. Androecious flower in bud globose to globose- depressed, 7-9 mm diam., the apex with 4 mi- nute, obtuse tepals, the pedicel 10-28 mm by 1 mm, subtended by a minute bracteole; at anthe- sis (3-)4-fid, 16-26 mm diam., the lobes spread- ing flat; stamens 65—95, ligulate, 1.5-2.5 mm long, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 0.5-0.8 mm wide, the filament short, the loculi lateral, comprising ca. 25—⁄4 total length of the stamen, apically connivent or confluent, the con- nective not prolonged. Gynoecious flower in bud napiform-depressed, apiculate with 4 minute te- pals, the pedicel (5-)10—-20 mm by 0.6-0.8 mm, subtended by a single bracteole; at anthesis 6-7 mm diam. by 4-6 mm long, the orifice X-shaped, 1-2 mm diam., splitting into 4 suberect deltoid lobes; styles numerous, ca. 150- , Short, bluntly columnar, 0.5 mm long by 0.3-0.4 mm wide, interspersed with rare, scattered hairs. Fruiting receptacle solitary or in clusters of 2-3 on leafless branches, cupuliform-urceolate, 25- 45 mm diam. by 15-30 mm long, the orifice 6- 14 mm diam., comprising ca. !^—'^ total width ofthe fruit, subentire, the walls 10-15 mm thick, the external surface dark reddish brown mottled with pale corky patches, the internal surface with scattered, short, bluntly columnar-polyhedral styles ca. 0.3-0.5 mm long by 0.5 mm wide, interspersed with rare, scattered hairs, the ped- icel and peduncle 12-16 mm by 3-6 mm. Mature carpels 9-11 mm long, 5-7 mm wide, 3-4 mm thick, the endocarp pale brown, the surface with sas striations. Gametic chromosome num- = 19. Figure 32. Distribution. Endemic to Réunion (Fig. 31). Habitat. Tambourissa elliptica subsp. mi- crantha is occasional to locally common from ca. 300 to 700 m in the southern sector of Ré- union on the lower slopes of the active volcano, the Piton de la Fournaise (le Volcan) massif (at Basse Vallée, Brülé de Baril, Forét de la Mare Longue, Mare d’Arzul, and Bois Blanc). Collec- tions are also known from l'Echo below Plaine des Palmistes (ca. 700 m), and the subspecies reaches ca. 1,000 m at Brülé de St. Denis and Plaine d’Affouches. REUNION. BOIS BLANC: 300 m, Apr. 1949 «, the s.n. (TL-R). BROLE DE ST. DENIS: rainy seaso Feb.) 1945 (fr), Rivals s.n. (TL-R). BAssE epis é- loni, 21 Oct. 1971 (fr), Friedmann 3265 (P). BRÜLÉ DE BARIL: = dia 1975 (fl), Coode et al. 4963 (K, 2 sheets; REU); 300 m, 2 Mar. 1979 (st), Lorence, Lorence & Cadet 1 (MAU, MO); m Lorence, Lorence & Ca- det 2488 (B, K, MAU, MO, P, REU, Z); 13 July 1979 (fr), Lorence & Cadet 2741 (MO); (fr), mpi a Cadet 2742 (MAU, MO); (fr), Lorence & Cadet 3 (B, K, MAU, MO, P, REU, Z). Dos D'ANE: K m, Mar. 1974 (fl), Friedmann 2299 (P, intermediate form). GRAND ÉTANG: (E Réunion) ca. 500 m, 15 July 1979 om mils & Rolland 2758 (MO, intermediate form). o: St. Benóit to Plaine des Palmistes route, 700 m, wi July 1967 a prs 1038 bis (REU); (st), Cadet 1985] LORENCE—MONIMIACEAE 119 FIGURE 32. hese ion elliptica (Tul.) A.DC. subsp. micrantha Lorence.—A. Habit.—B. Androecious flower at anthesis, apical v w.—C. Androecious flower at anthesis, = view. 2 s men, abaxial view.— E. Stamen, adaxial view. M Gynoecious flower at anthesis, apical view.—G. G s flower at anthesis, longitudinal section. — H. Submature ae receptacles. A-G. Pane 2489 (MO). H. pani 2741 (MO). Bars equal 10 mm in A-C, F-H, and | mm in D, E 1039 bis (REU, intermediate ae 27 Feb. 1979 (st), (fr), Rivals s.n. (TL-R). RIVIERE DES REMPARTS: high sah iin 2462 (MO, intermediate form). PLAINE D'AF- bes of the Riviére = bom tele 1 ,000 m, 7 Mar. OUCHES: La Mon e road to Plaine d'Affouches, 975 (fl), Cadet 5123 (R form); 1,100 1 ,000 m, pk 4.5, 27 Dec. 1966 (st), Cadet 612 (REU), m, OR Cadet 5124 ion 2d form). sT. PHIL- 120 IPPE: Mare d'Arzul forest, 9 Dec. 1943 (st), Rivals s.n. (TL-R); St. Philippe (st), Rivals s.n. (TL-R); Mare Lon- m, 14-15 Jan. 1975 (fr), Bernardi 44 (G, 3 sheets; P, REU); ca. 500 m, 25 Feb. 1975 (fl), Coode et al. 4956 (K, 2 sheets; REU); 250-500 m, 2 Mar. 1979 (fl), Lorence, Lorence & Cadet 2483 (MAU, MO); Jan. 1949 (st), Rivals s.n. (TL-R); May 1943 (st), Rivals s.n. (TL-R). TAKAMAKA: Takamaka forest, Ilet à Bananes, ca. 800—900 m, 10 Mar. e (fl), Lorence & Rolland 2524 (MO, intermediate form). WITHOUT PRECISE LOCALITY: 1853-1863 (fl, fr), Ln 4 (P, 5 sheets). At Brülé de Baril, Tambourissa elliptica subsp. micrantha becomes a subcanopy or canopy tree attaining 10-12 m with a straight trunk attaining 25-30 cm D.B.H. and a narrow, compact crown with numerous, erect branches. It differs most conspicuously from subsp. elliptica in having el- liptic to narrowly elliptic leaves with a laminar length to width ratio exceeding 2:1 and often approaching 3:1. Juvenile and sucker leaves of subsp. micrantha are longer and narrower than its adult leaves, have more numerous secondary veins, bright red petioles and midribs when liv- ing, but lack dentate margins. Subspecies e/lip- tica has instead apically dentate-serrate juvenile leaves, as do most other congeners for which data is available. Flowers of both sexes are conspicuously small- er than in subsp. elliptica, hence the epithet mi- crantha. Large individuals of subsp. micrantha are monoecious, often with sexually mixed plei- ochasia, whereas sex expression appears to be size and/or age related in subsp. elliptica. Flowers of subsp. micrantha are generally pleiochasial and ramiflorous or axillary, whereas those o xillary. of s generally smaller, iun produced in clusters of two or three on the branches, and are more con- spicuously mottled in contrast to the larger, sol- itary and often caulinary fruits of subsp. elliptica which are more uniformly pale brown. Although the two subspecies of Tambourissa elliptica occur together in certain areas of wet forest at low to medium altitudes (e.g., Hauts du Bois Blanc; Hauts du Brülé de St. Denis) and remain distinct there, they intergrade in other localities, e.g., Takam Riviére des Remparts (Cadet 5123, 5124, REU), and Grand Etang (Lorence & Rolland 2758, MO). Both subspecies have n = 19. Despite occasional intermediate collections, the two taxa are sepa- rable by relatively constant, modally distinct ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 characters of floral size and odor, staminal length and number, ratio of laminar length to width, and fruit size. I have therefore chosen to recog- nize them as subspecies. Tambourissa elliptica subsp. micrantha be- longs to species Group 2A. In view of its monoe- Cy, less reduced pleiochasial inflorescences, and flowers intermediate in size and number of parts, it seems to be the least specialized of the Réunion taxa and therefore probably most like the an- cestral species from which the others were de- rived. Vernacular names. Bois de bombarde, Bois de bombarde à petites feuilles (Réunion). 14. Tambourissa crassa Lorence, Bull. Mus. Hist. Nat. (Paris), Ser. 4, Sect. B, Adansonia 3(3): 298, pl. 2, fig. 7. 1982. TYPE: Réunion. Road to Maido above Petite France, on forestry road C.F. 81; low cloud forest, ca. 1,200- 1,400 m, 19 July 1979 (fl), Lorence 2780 (holotype, MO; isotypes, K, MAU, P, REU, Z) T. religiosa sensu Cordem., Fl. Réunion 301. 1895 non (Tul.) A.DC. Dioecious tree, rarely shrubby, 7—8 m tall, the trunk stout, to 30 cm D.B.H., the bark pale gray- ish brown, smooth to irregularly fissured-flaking, the crown compact, the new growth glabrous, rarely with a few scattered hairs, the mature leafy stems 4-9 mm diam., terete, glabrous, longitu- dinally wrinkled. Leaves glabrous, opposite to subopposite, petiolate to subsessile; petiole stout, 4-15(-20) mm by (2-)3-6 mm; lamina coria- ceous, obovate, broadly obovate, broadly elliptic or suborbiculate, (40-)60-175 mm by (20-)40- 140 mm, the apex obtuse, rounded, or retuse- emarginate, the base acutely cuneate to acutely decurrent, the secondary veins 5—6 pairs, making a 45-55? angle with the costa, the venation ob- scure, visible to 2(-3?) adaxially and to 3(-4°) abaxially, the margin moderately revolute, the color pale brown to bright yellow-green when ry. Inflorescence a solitary monochasium ter- minal on the leafy stems (the subtending leaves + caducous), or rarely cauliflorous or ramiflo- rous and terminal on short, leafy or leafless brac- teate shoots. Androecious flower in bud globose, 13-14 mm diam., glabrous, apiculate with 4 small, obtuse-deltoid, + glabrous tepals, the ped- icel and peduncle stout, erect or recurved, 15- 32 mm by 4-5 mm, bearing several caducous, LORENCE— MONIMIACEAE 121 1985] ciliate deltoid bracteoles basally; at anthesis 4— 5-fid, 32-45(-50) mm diam., the lobes thick, spreading almost flat or somewhat incurved, the internal receptacle surface glabrous; stamens nu- merous, ca. 125-250, ligulate-ellipsoid, 4-5 mm long by 1-1.5 mm wide, the filament short and thickened basally, the loculi lateral, comprising ca. 25—⁄4 total length of the stamen, usually con- fluent apically, occasionally free, the connective not prolonged. Gynoecious flower in bud napi- form to globose-depressed, the apex with 4 small, obtuse or deltoid, glabrous or sparsely ciliate te- pals, the pedicel and peduncle suberect, 8-2 mm by 2-4 mm, bearing several small, + ciliate bracteoles basally; at anthesis urceolate-globose to urceolate-napiform, 122-14 mm diam. by 8- 10 mm long, the walls thick, the apex splitting into 4(-8) thick, s sees deltoid lobes, the orifice 5-6 m comprising ca. !⁄4—!A total width of the ee styles numerous, ca. 200 or more, crowded, bluntly columnar, ca. 0.5 mm long by 0.3-0.5 mm wide, the internal receptacle surface glabrous, producing a muci- udate. Submature fruiting receptacle globose-urceolate, ca. 60 mm diam. by 40 mm long, corky brown externally, internally glabrous with numerous scattered, bluntly columnar poly- hedral styles ca. 0.3 mm long by m wide, the pedicel and peduncle stout, ca. 50 mm by 10 mm. Mature carpels unknown. Figure 33. Distribution. Endemic to Réunion (Fig. 31). Habitat. Tambourissa crassa is local and oc- casional in cloud forest formations from ca. 1,200 o 2,000 m, rarely extending as low as 700 m (e.g., at Basse Vallée, Bosser 21313). RÉUNION. BASSE VALLÉE: humid forest, 600-700 m, 20 Oct. 1972 (fr), Bosser 21313 (P). BOIS DE NEFLES: Hauts du Bois de Néfles, St. Paul, 1,400 m, 17 Sept. 1979 (fl, fr), Cadet 4790 (REU); (fl), Cadet 4791 (REU). Les Mares, 1,500 m (fl), Friedmann 3001 m, 22 Feb. 1979 (st), Lorence & Cadet 2412 (MAU, MO); ca. 1,200 m, 22 Feb. 1979 en, z Lorence & Cadet 2414 (MAU, MO). PLAINE DES n s.n. (TL-R); Col de Bellevue at Grande Mo Plaine des Cafres, low be Miei 1,600 m, 2 1979 xr Lorence 2468 (M. MO); (fl), Lorence 2469 MO); (fl), Lorence pr O); 17 July 1979 s.n. (st), Friedmann 1 031 P) PLAINE DES PALMISTES: ‘banks of the Ravine Petite 4 Bras des Calumets, 1,300 m, 17 Jan. 1972 (fl), Cadet 3477 (REU). PLAINE DES SALAZES: 2,000 m, 19 Feb. 1944 (st), Rivals s.n. (TL-R). PLAINE DES TAMARINS: 1,800 m, below the Col de Fourche, Mafate, 28 Nov. 1974 (fl), Cadet 4604 (REU). RIVIERE DE L’EST: low thicket along the Riviére de l'Est, 1,550 m, Nov. 1967 (fl), Cadet 5717 (REU). TEVELAVE: for- estry road above Tévelave, cloud forest, 1,300-1,400 m, 28 Feb. 1979 (fl), Lorence 2478 (MO); along the domainial line, 1,700 m, cloud forest transitional to heath formation, 6 Mar. 1979 (fl), Lorence 2500 (MAU, MO); Hauts du Tévelave, 1,500-1,900 m (st), Rivals s.n. (TL-R). WITHOUT PRECISE LOCALITY: (st), Rivals s.n. (TL-R). Tambourissa crassa belongs to species Group 2A and is most closely allied to 7. elliptica subsp. elliptica, also from Réunion, from which it can be distinguished by its truly dioecious habit, thicker leaves, and solitary, terminal flowers and fruits borne at the ends ofthe top branches (rarely caulinary on short, leafy or leafless bracteate shoots, but even then solitary and terminal). The succulent, strongly discolorous leaves of T. cras- sa have a well-developed hypodermis 4-6 cell layers thick, which is only 2-3 cells thick in both subspecies of T. elliptica. Other distinguishing features are its somewhat pachycaulous habit, sparse and compact crown, the almost cam- phor-like fragrance of the dried specimens, and absence of stone cells and c us, idioblasts from the floral ground tiss Unlike the more cbe Tanais ellip- tica subsp. elliptica, T. crassais quite stable mor- phologically. As in most species, a certain amount of intraspecific variation occurs in leaf size and shape, flower color, and number of stamens, but both species are quite distinct morphologically. oth T. crassa and T. elliptica subsp. elliptica frequently occur sympatrically in the cloud forest zone (1,200-2,000 m), e.g., at Hauts du Téve- lave, and along the road to Maido, where neither intermediates nor putative hybrids were found. For these reasons I regard 7. crassa as a discrete species rather than a subspecies of T. elliptica in spite of certain morphological similarities. Both Tambourissa crassa and T. elliptica subsp. elliptica have large, structurally similar leaves and large flowers with numerous stamens and styles, which are presumably adaptations to a shared habitat and similar pollinators. Appar- ently both taxa have evolved from a common ancestor with less specialized features, which 7. elliptica subsp. micrantha appears to approach most closely. Tambourissa crassa has attained the highest degree of specialization of the three Réunion taxa, however, in being dioecious and having inflorescences reduced to a single, ter- minal flower. 122 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 FIGURE 33. Tambourissa crassa Lorence. — A. Habit, with androecious flower at anthesis. — B. Stamen, adax- ial view. —C. Habit, with gynoecious flower at anthesis. — D. Gynoecious flower at anthesis, mes view.— Gynoecious flower at anthesis, longitudinal section. — F. Gynoecious flower at anthesis, apical v G. Fruiting receptacle, longitudinal section. A, B. Lorence 2767 (MO). C-F. Lu 2780 (MO). G. Cade 4790 (REU). Bars equal 10 mm in A, C-G, and 1 mm in B. 1985] Flowers of both sexes produce strong, rancid- fruity floral odors of butyric acid, as do those of Tambourissa elliptica subsp. elliptica. Both pale yellow-green and dark purple floral color morphs occur, although flower color appears to be con- stant within a given population (e.g., yellow at Col de Bellevue, purple at Hauts du Tévelave). Vernacular names. Bois de bombarde, Bois de bombarde à grandes feuilles (Réunion). 15. Tambourissa comorensis Lorence, Bull. Mus. Hist. Nat. (Paris), Ser. 4, Sect. B, Adansonia 3(3): 295, pl. 1, figs. 3, 4. 1982. TYPE: Co- more Archipelago. Grande Comore: La Grille forest above Maoueni, ca. 800 m, 29 July 1979 (fl, fr), Lorence & Banfi 2878 (ho- lotype, MO; isotypes, B, G, K, MAU, P, REU, Z) Large monoecious canopy tree 15-20 m tall and 50 cm D.B.H., the bark pale brown, smooth, flaking, the new growth glabrous or with rare, scattered hairs, the mature leafy stems pale greenish, terete, glabrous, 3-4 mm diam. Leaves opposite to subopposite, glabrous, petiolate; pet- Sw. 10-28 mm by 1-1.5 mm; lamina charta- s, elliptic, narrowly elliptic, ovate, oblong or lanceolate (60-)80-140 mm by (20-)30-75 mm, e apex shortly acuminate, rarely acute or ob- e the tip rounded, the base acutely cuneate to obtuse, generally slightly decurrent, the sec- ondary veins 4-7 pairs, making a 50-60? angle with the costa, the venation prominent, visible to 4(-5?) on both surfaces, the margin plane to slightly revolute; juvenile leaves heterophyllous, the apical '—* of the lamina serrate-dentate with 1-5 pairs of large teeth, the petiole red when living. Inflorescence cauliflorous or ramiflorous, n meristematic swellings on the trunk or on leafless nodes of the branches, a short, unisexual or sexually mixed pleiochasium of 4-9 flowers, the floral axis 5-20(-60) mm by 2-3 mm, when young finely velutinous-pubescent with short, simple or fasciculate hairs, each pedicel sub- tended by a minute naviculate Sipaspa, An- droecious flower in bud globose, 9-14 mm diam., externally puberulent to glabrous, + eee dai apiculate with a small pore flanked by 2-4 pairs of dark, hirsutulous, broadly deltoid tepals, the pedicel 8-30 mm by 1-1.5 mm; at anthesis deep- ly 4(-6)-fid, 23-35 mm diam., the lobes spread- ing flat and reflexing slightly, the apex of each lobe with several fleshy tepals internally, the in- ternal receptacle surface glabrous; stamens nu- LORENCE— MONIMIACEAE 123 merous, ca. 125-150, linear-ligulate, 2.5-4 mm long by 0.7-1 mm wide, the loculi separate, par- allel, lateral or slightly abaxial, occupying almost the entire length of the stamen, the filament short, thick, the connective not prolonged. Gynoecious flowers 1-6 per inflorescence, terminal or lateral, the pedicel 12-20 mm by 2 mm; at anthesis broadly patelliform-urceolate, 12-15 mm diam. by 5-7 mm long, the orifice circular, entire, 7— 10 mm diam., comprising ?^ total width of the receptacle, the rim velutinous, open even in bud, the external surface puberulent, + pustular; styles merous, several hundred, coalescent, short, bluntly columnar-polyhedral, 0.3-0.5 mm long and wide, the apex papillose, the internal recep- tacle surface with scattered hairs. Fruiting recep- tacle solitary or in clusters of 2-3 on the trunk and major branches, irregularly cupuliform, often + distorted, 90-175 mm diam. by 50-75 mm long, the orifice comprising ca. 3-2 total width of the receptacle, entire, subcircular, 45-90 mm diam., the walls 16-25 mm thick, externally smooth, mottled reddish brown, internally gla- brous with numerous scattered columnar-poly- hedral styles 0.3-0.4 mm long by 0.5-0.8 mm wide. Mature carpels ovoid to ellipsoid, com- pressed, 11-13 mm long, 5-8 mm wide, 4-6 mm thick, the endocarp dark brown to black, the sur- face scrobiculate. Gametic chromosome num- ber, n = 19. Figure 34 Distribution. Endemic to Grande Comore (Fig. 35) Habitat. Tambourissa comorensis occurs in lower montane wet forest formations. All known collections are from La Grille forest above Ma- oueni (northern part of the central ridge, 800- 900 m), and from forest above Ntsorale Dimane along the eastern part of the same massif (250- 900 m, fide Bernardi). At La Grille, I found it to be locally abundant and codominant as a sub- canopy or canopy tree, striking when in fruit with numerous, large brown receptacles hanging from the trunk and branches, some split open to reveal an array of red-orange carpels set against a pale orange background. One of the largest known members of the genus, both in habit and in fruit, it flowers from August to December concurrently with the ripening ofthe previous year's fruit crop. GRANDE COMORE. NTSORALE DIMANE: 250-900 m, uxia and Weinmannia formation, 9 Dec. 1967 (fl), scias formation, 1-3 Dec. 1 (G, 2 sheets; K, P, Z); May 1963 (fl), Bosser 18096 124 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 (HII Mh b. pis jr P, yy SEN SSS = Se = A IN 7 ANYS WV UN y (| Ú N FIGURE 34. Tambourissa comorensis Lorence.— A. o — B. Infl ith and “asa —C. Androecious flower in bud, apical view.—D. Androecious flower at anthesis, Du view.—E. Sta adaxial view. — F. Gynoecious flower, longitudinal section. — G. Submature fruiting receptacle, longi- ier sectio n.—H. Fruiting carpel, lateral view. All Lorence 2878 (MO). Bars equal 10 mm in A-D, G, H, and 1 mm in E, F 1985] O T. paradoxa A T. kirkii @ T. comorensis —+ 12940o's | 4320'E | 30 km FIGURE 35. Anjouan, Mayotte MAD, P); ca. 800 m, 29 July 1979 (fr), Lorence & Banfi 2877 (MO, K, P); 9 Feb. 1957 (fl), Anon. in SF- 16529 (P). Tambourissa comorensis belongs to species Group 24A, also including 7. /eptophylla which differs by its smaller androecious buds, fewer sta- mens, and by its gynoecious receptacle with a much smaller orifice and acute, conical styles. Also, fruits of the latter are uniformly dull, corky brown externally with a smaller orifice. Dried fruits of T. comorensis are strongly fragrant in the herbarium, like those of T. moheliensis (Group 3). Tambourissa kirkii (Group 2B) differs by its small, globular androecious flowers, which open only partially and contain fewer stamens with acute, prolonged connectives. In living ma- terial, androecious flowers of 7. comorensis are white and produce a pleasant, sweet odor of fer- LORENCE—MONIMIACEAE 44°30'E 11°50'S d | 43°50'E 125 1390's : 45°20'E 12°30'E B T. leptophylla A T. moheliensis Distribution maps of Tambourissa in the Comore Islands (left to right): Grande Comore, Mohéli, te. menting fruit. Buds and gynoecious flowers are pale green externally. At anthesis, gynoecious flowers have glistening, pale green styles, al- thou parent. In mixed inflorescences of Lorence & Banfi 2788, the gynoecious flowers matured be- fore the androecious flowers. was not ap- Vernacular name and uses. M’Bweza (Grande Comore), meaning “to calm the pain” or “to heal." Natives of Grande Comore infuse the leaves to make a medicinal tea. 16. Tambourissa leptophylla (Tul.) A.DC., Prodr. 16(2): 658. 1868; Baill., Hist. Pl. ed. 2, 1: 310, 344, 345. 1871; Cavaco in Hum- bert, Fl. Madagascar 80: 34, fig. VIII:1-4. 1959. Ambora leptophylla Tul., Ann. Sci. 126 Nat. (Paris) 4(3):29. 1855; Tul., Monogr. Monim. 298, tab. 25, fig. 1. 1855. TYPE: Co- more Archipelago. Mayotte: Zuaby (ca. 1851) (fl), Boivin 3133 (lectotype, P, here designated; isolectotypes, P, 4 sheets) Monoecious or subdioecious tree attaining 10 m tall and 40 cm D.B.H., the bark pale brown, flaking, the new growth finely pubescent, the ma- ture leafy stems glabrescent, terete, 3-5 mm diam. Leaves opposite to subopposite, glabrous, peti- olate; petiole 15-20 mm by 1.5-2.5 mm; lamina membranaceous to chartaceous, ovate to elliptic or oblong, 120-250 mm by 60-110 mm, the apex shortly acuminate to acute, the base acute to ob- tuse, often slightly decurrent, the secondary veins 5-8 pairs, making a 50-65? angle with the costa, the venation visible to 4? adaxially and to 4(—5?) abaxially, the margin plane to slightly revolute; juvenile and sucker leaves heterophyllous, the margin serrate with 2-5 pairs of teeth in the api- cal half, the petioles red when living. Inflores- cence a pleiochasium or thyrse of 7-25 flowers, sometimes leafy, cauliflorous from meristematic swellings on the trunk, or a ramiflorous plei- ochasium of 5-11 flowers on the leafless nodes, or the flowers solitary and axillary, when young finely velutinous, later puberulent, the floral axis 20-150 mm by 1-3 mm, subtended by several minute, caducous bracteoles. Androecious flow- er in bud globose-depressed, 6-8 mm diam., smooth, the apex with 4 minute puberulent te- pals, the pedicel 10-40 mm by 1-1.5 mm, sub- tended by a small, velutinous deltoid bracteole; at anthesis deeply 4-fid, 20-25 mm diam., the lobes spreading flat, the internal receptacle sur- face glabrous; stamens 80-90, linear-ligulate, re- curved, 2-3 mm long by 0.8-1 mm wide, the loculi separate, parallel, occupying almost the entire length of the stamen, apically connivent or the connective slightly prolonged and apicu- late. Gynoecious floral receptacle globose-patel- liform, apically depressed; at anthesis 10-15 mm diam. by 5-7 mm long, the orifice entire, circular, comprising ca. '4—' total width of the receptacle, open even in bud, the rim hirsute; styles very numerous, ca. 300—600, broadly conical, 0.6—0.8 mm long by 0.3-0.5 mm wide basally, the in- ternal receptacle surface hirsute between the styles. Fruiting receptacle solitary or in clusters of 2-4 on the trunk and major branches, globose- patelliform, apically depressed, 90-130 mm diam. by 75-90 mm long, the orifice circular, entire, 15-40 mm diam., comprising ca. !⁄¿—!1⁄ ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 total width of the receptacle, the walls 16-25 mm tered, sharply conical styles 0.7-1 mm long by 0.5 m wide basally, interspersed with stud cdanf simple hairs, the pedicel and peduncle tapered, 70-80 mm long by 9-10 mm diam. me- dially. Mature carpels ovoid-compressed, 13-17 mm long, 8-10 mm wide, 3-6 mm thick, the endocarp light to dark brown, the surface shal- lowly foveolate-reticulate. Distribution. Endemic to Mayotte, Comore Archipelago (Fig. 35). abitat. It is local and occasional in lower montaine evergreen wet forest formations often dominated by Aphloia and Nuxia. Recent col- lections are from the Benara massif (400—653 m) and also the Hachiroungou massif (400—600 m), in remnants of indigenous forest. The species also occurs in secondary forest with native rem- nants, the recent collections being from Barakani and Sada, both at ca. 100 m. Flowering occurs from August to September concurrently with the ripening of last year's fruits. MAYOTTE. o secondary forest and along n ca. 100 m, 24 July 1979 (fr), Lorence & Saida 2824 (MO); Suec rakani, Aug. 1977 (fl, fr), Tattersall a EA wet forest on sum- an Lorence & Saida 2829 (MAU, MO, P). DZOMNOGNE: (fl), 86 (P). HACHIROUNGOU MASSIF: (NW Mayotte) a dose with Nuxia and Aphloia, ca. 400-500 m, 26 July 1979 (fr), usq s Saida 2854 (K, MAU, MO , P, REU, Z). sa . 100 m, Aug. 1977 (fl), Tattersall s.n. (MO). d leptophylla belongs to species Group 2A. Its androecious flowers most closely approach e of T. comorensis, which also split into four flat segments and bear numerous sta- mens with long anthers, although flowers and buds of the latter are larger. Gynoecious flowers of T. leptophylla have acutely conical styles as in T. kirkii (Group 2B), but androecious flowers of the latter are much smaller, open only par- tially, and have fewer stamens (ca. 50) with short, medial loculi and acutely prolonged connectives. All three species are presumably derived from a common ancestor, perhaps one similar to 7. madagascariensis. Vernacular name. Ambora (Mayotte). 17. Tambourissa kirkii Cavaco, Kew Bull. 12(2): 228. 1957; Cavaco, Bull. Soc. Bot. France 1985] LORENCE—MONIMIACEAE 127 104: 284, fig. 3. 1957. TvPE: Comore Ar- chipelago. Johanna (Anjouan): without pre- cise locality, Zambesi Expedition, 1,000 ft. (ca. 300 m), small tree, Aug. 1862 (fl, fr), Kirk s.n. (holotype, K) (see also Kirk’s fig. 355, with type). Monoecious shrub or small tree, the new growth hirsutulous, the mature leafy stems glabrous, stramineate, 2.5-4 mm diam. Leaves opposite to subopposite, glabrous, petiolate; petioles 10- 15 mm by 1-1.5 mm; lamina ovate to elliptic, 100-135 mm by 55-70 mm, the apex shortly acuminate, the base obtuse to rounded, the sec- ondary veins 6—7 pairs, making a 50-55? angle with the costa, the venation visible to 3-4? on both surfaces, the margin slightly revolute. An- droecious inflorescence unknown. Androecious flower (from Kirk's fig. 355) globose, ca. 9-10 mm diam.; at anthesis deeply 3-fid, opening slightly, the lobes incurved; stamens probably ca. 50, subulate, ca. 3-4 mm long by 0.5 mm wide, the loculi separate, lateral, occupying medial '^ of stamen, the filament basally broad, compris- ing ca. 3 total length of the stamen. Gynoecious inflorescence a 4(—5)-flowered, finely velutinous pleiochasium, the position unknown, the floral axis ca. 20 mm by 1.5 mm, bearing rare, scattered bracteoles; female floral receptacle cupuliform- patelliform, 8-12 mm diam. by 5-6 mm long, the orifice broad, circular, entire, comprising ca. 34 total width of the receptacle, the rim veluti- nous, the external surface finely velutinous, + basally, bearing short, scattered hairs, the inter- nal receptacle surface velutinous between the styles. Fruiting receptacle (from Kirk's fig. 355) broadly cupuliform-patelliform, ca. 250-300 mm diam., the broad, subcircular orifice comprising ca. !^—/ total width of the receptacle, the conical styles persistent in fruit. Mature (?) carpels ca. 10-12 mm long by 7-8 mm wide, ovoid-com- pressed, the endocarp “black and hard, sur- rounded with a red fleshy body" (fide Kirk). Distribution. Endemic to Anjouan (Fig. 35). Habitat. According to the information given on the label, Tambourissa kirkii occurs at 1,000 ft. (ca. 300 m), but on his figure 355 (an unpub- lished pencil drawing), Kirk notes “shrub grow- ing at height of 2,000 feet (ca. 600 m) . . . seeds said to be cathartic. Monoecious." It probably occurs in montane wet forest. Tambourissa kirkii belongs to species Group 2B. As noted by Cavaco (19572), it is closely allied to T. leptophylla (Group 2A) from neigh- boring Mayotte. The latter differs by its more sparsely pubescent inflorescence, smaller gy- . a el dám] 46 ola- ifr. land brous styles, and by its androecious flower which is about twice as large with more numerous sta- mens having much longer loculi extending for almost the entire length of the stamen, and a scarcely prolonged connective. Tambourissa paradoxa (Group 2B), also from Anjouan, has smaller, globular androecious flow- ers opening by a small pore and structurally sim- ilar stamens: it may even be conspecific with T. kirkii. As gynoecious flowers are unknown in the former, however, and androecious flowers of the latter are known only from a single pencil sketch, I have decided to recognize both species until further data and collections become available. Tambourissa comorensis and T. leptophylla dif- fer by the characters mentioned above. 18. Tambourissa paradoxa Perk., Pflanzenr. 4, 101 (Nachtr.): 43. 1911. TYPE: Comore Ar- chipelago. Johanna (Anjouan): without pre- cise locality (fl), Anon. sub Herb. Blackburn s.n. (possibly Bojer) (holotype, K). T. pa Perk., Pflanzenr. 4, 101 (Nachtr.): 43. 1911. omore Archipelago. Johanna (Anjovany Zylangi i Lake, 27 Oct. 1862 (fl), Anon. sub Zam $m Expedition s.n. (possibly Kirk) (holotype, K, efthand specimen with flowers). Monoecious shrub or small tree attaining 10 m tall, the new growth unknown, the adult leafy stems glabrous, terete, 3-4 mm diam., yellowish brown. Leaves opposite to subopposite, gla- brous, petiolate; petioles 10-20 mm by 1- mm: lamina chartaceous, elliptic to ovate ellip- tic, 60-140 mm by 25-50 mm, the apex acute to acuminate, dosi base ri cuneate to acutely decurrent, t ins 5-8 pairs, making a 40-45? iid with the pim the venation prom- inent and visible to 3? adaxially and to 4? abax- ially, the margin moderately revolute. Androe- cious inflorescence ramiflorous, a sparse, puberulent pleiochasium of 3-11 flowers, or the flowers solitary and axillary, the floral axis 6-12 mm by 1 mm, bearing scattered, simple hairs, subtended by several pairs of minute, ciliate del- toid bracteoles. Androecious flower in bud glo- bose, the pedicel 5-12 mm by 0.6-1 mm, bearing a few sparse hairs, subtended by 1(-2) minute, 128 caducous deltoid bracteoles; at anthesis globose, mm diam., not splitting but opening by a shallowly 4- lobed apical orifice ca. 2 mm diam. flanked by 4 small, decussate, obtusely deltoid ciliate tepals ca. 0.6-1 mm wide by 0.3—0.5 mm long, the internal surface of each lobe bearing 2- 10 fleshy, obtuse to deltoid tepals intergrading with the stamens; stamens ca. 40—50, subulate- lanceolate, 2-3 mm long by 0.5-0.8 mm wide, the loculi lateral, separate, occupying basal 7^ of stamen, the connective prolonged, acuminate, slightly exserted from the orifice at anthesis; in- ternal receptacle surface glabrous. Gynoecious flowers unknown. Fruiting receptacle solitary on leafless SERE Md shallowly cupuliform, 53-150 mm diam. by 25-50 mm long, externally corky, the orifice FR ida ca. !^ total width of re- ceptacle, the internal surface with numerous broadly conical styles 0.7—0.8 mm long, more or less glabrous in between; pedicel and peduncle 15-17 mm by 5-7 mm. Mature carpels ovoid- compressed, 11 mm long by 6 mm wide. Distribution. Endemic to Anjouan (Fig. 35). Habitat. The species is apparently restricted to evergreen montane wet forest from ca. 650 to 1,400 m, according to available information. UAN. WITHOUT PRECISE LOCALITY: Traprock (?) 18 NJO soils of hills, 2,000-3,000 ft., 29 Oct. 1862 (st), Anon. u mbesi Expedition s.n. (possibly Kirk) (K, sterile wa > N t. (st), gia s.n. (P). N'T m), in evergreen forest, food of pigeons, bulbuls and thrushes, tree : + 20 ft. tall with huge fruits weg on t. 1958 Stell MUCU (fr), Benson 142 (BM). The types of Tambourissa johannae and T. ee are 2er from Anjouan. Both collec- similar leaves, small, Mure androecious flowers which open by a narrow apical orifice flanked internally by de- cussate, scale-like tepals, and have virtually iden- tical stamens containing oil cells and a few scle- rids, but lacking tanniferous idioblasts. It seems clear that these two concurrently published taxa are conspecific. I have chosen to retain the epi- thet paradoxa, as its type possesses more nu- merous and mature flowers than the type of T. o tions and collections, E of female flow- ers and fruit, are desir As is the case for NE gracilis and T. purpurea (both Madagascar), the androecious ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 flowers of 7. paradoxa open by small apical pores. Unlike the stamens of the former two, which are nearly sessile with apically confluent loculi, those of T. paradoxa are long and lanceolate with sep- arate, lateral loculi and acute, prolonged con- nectives A close affinity appears to exist between Tam- bourissa paradoxa and T. kirkii (also Anjouan), which has strikingly sinar floral ang staminal logy , although of the e latter species split into 3—4 segments and lack the exserted connectives of the former. Both be- long to species Group 2B. Similar androecious floral and staminal morphology occurs in 7. moheliensis (Group 3) from neighboring Mohéli, although here the loculi are united basally into a U-shape, and the gynoecious flower differs in being closed in bud. The similarities in androe- cious floral and staminal morphology of T. par- adoxa, T. kirkii, and T. moheliensis may be a result of convergence to accommodate similar pollinators, as the very different gynoecious flow- er of the latter suggests a closer affinity to the Madagascan 7. capuronii (also Group 3) Vernacular name. M'Bweza (Anjouan, fide ) Benson 142, B 19. Tamb i heliensis Lorence, Bull. Mus. Hist. Nat. (Paris), Ser. 4, Sect. B, Adansonia 3(3): 300, pl. 2, figs. 9-11. 1982. TYPE: Co- more Archipelago. Mohéli: mountains be- tween Fomboni and Drodoni, low montane forest, 500-700 m, 6 Dec. 1967 (fl, fr), Ber- nardi 11781 (holotype, Z; isotypes, G, K, P) Monoecious or subdioecious tree 4-15 m tall by 70 cm diam., the new growth glabrous or with sparse, scattered hairs, the mature leafy stems pale green, terete, 2-5 mm diam. Leaves oppo- site to subopposite, glabrous, petiolate; petioles 7-22 mm by 1-1.5 mm; lamina subcoriaceous, ovate, broadly to narrowly elliptic, oblong or rarely obovate, 45-220 mm by 30-70 mm, the apex acute to acuminate, rarely rounded, the base acute to obtuse, cuneate or decurrent, rarely ob- tuse or rounded, the secondary veins 4-8 pairs, making a 45-65? angle with the costa, the ve- nation visible to 2(-3?) adaxially and to 3(-4?) abaxially, the margin slightly revolute. Gynoe- cious inflorescence a finely pubescent, cauliflo- rous pleiochasium or thyrse of up to 20 flowers; androecious inflorescence a ramiflorous plei- 1985] LORENCE— MONIMIACEAE 129 FiGuRE 36. Tambourissa moheliensis Lorence. — A. Habit. — B. Stamen, adaxial view. — C. Submature gyn- oecious inflorescence. — D. Gynoecious flower at anthesis, lateral view. — E. Gynoecious flower, longitudinal section. — F. Styles, lateral view.—G. Submature fruiting receptacle, longitudinal section. — H. Fruiting carpel, lateral view. A-E. Bernardi 11781 (G). F. Bernardi 11759 (G). G, H. Bernardi 11781 (Z). Bars equal 10 mm in A, C-E, G, H, and 1 mm in B, F. 130 ochasium of ca. 5 flowers on the leafless nodes, or the flowers solitary and axillary or rarely sub- terminal on short leafy shoots; floral axis 15-70 mm by 1-5 mm, finely pubescent or with scat- tered simple and fasciculate hairs, subtended by several ciliate-velutinous deltoid bracteoles. An- m apiculate with 4 minute, decussate velutinous tepals, the pedicel 10-12 mm by 0.8 mm, bearing scattered hairs, subtended by a minute, deltoid caducous bracteole; at (Complete?) anthesis glo- bose, 5-6 mm diam., splitting into 4 shallow, inflexed lobes, each with 3-4 obtuse, scale-like tepals apically on the internal surface, these in- tergrading with the stamens, the internal recep- tacle surface glabrous; stamens ca. 50, narrowly acute to subulate, 2-2.5 mm long by 0.6—0.8 mm wide basally, the loculi subparallel, occupying ca. ?^ total length of stamen, usually united basally into a single, U-shaped loculus, rarely free, the filament short, broad, the connective acute, pro- longed. Gynoecious flower in bud napiform-de- pressed, pubescent, apiculate with 4 velutinous deltoid tepals 0.5 mm wide, the pedicels 20-40 mm by 1.5-2 mm, pubescent; at anthesis napi- form-depressed, 12-16 mm diam. by 7-10 mm long, the orifice X-shaped, 1-2 mm wide, open- ing by 4 straight to suberect deltoid lobes, these finely pubescent-velutinous on both surfaces; styles numerous, ca. 100—200, conical, slightly ventricose medially, 1-1.3 mm long by 0.3-0.5 mm wide basally, glabrous, the internal recep- tacle surface velutinous with dense, simple and fasciculate hairs between the styles. Fruiting re- ceptacle borne on the trunk, patelliform to na- piform or subglobose, depressed, ca. 120 mm m long, the internal surface ifi c = j = 4e] E. & ° ° len | =Ñ < 9 < -4 Q o aring remnants of the black, deltoid lobes, the walls ca. 20—26 mm thick, the internal surface bearing numerous scattered, narrowly conical, angular styles 1-1.5 mm long by 0.8-1 mm wide basally, interspersed with scattered to dense, simple and fasciculate hairs. Mature car- pels ellipsoid to ovoid, compressed, 12-15 mm long, 6-7 mm wide, 4-5 mm thick, the endocarp dark brown, the surface scrobiculate. Figure 36. c m Endemic to Mohéli, Comores (Fig. 35). Habi tat. The species occurs in montane ev- ergreen forest from ca. 500 to 700 m. Both flow- ers and fruits are produced in December. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 MoHÉLI. FOMBONI: Bamboa, forest, 8 Mar. 1957 (fl), Anon. in SF-16731 (P). Mr. sr. ANTONIO: 600-700 m, in montane and submontane forest, 5 Dec. 1967 (fl, fr), Bernardi "1759 (G, K, P, Z). Mohéli, smallest of the four Comores, is prob- ably also the most poorly explored lieu with only three collections of Tambourissa known from the island. After much deliberation I have decided. to treat al as a single mera although d observations are required to ascertain ern or not they are truly conspecific. The first collection, Bernardi 11781, possesses two gynoecious inflorescences with numerous flowers and a single submature fruit. Because it has numerous flowers in both thyrses and pleio- chasia in addition to the fruit, I selected it as the type. Anon. in SF- 16731, from the same locality, corresponds well with the type oe but the buds are unfortunately owth, smaller and more shortly acute or api- cally rounded leaves with fewer secondary veins oriented at a greater angle to the costa, and stronger, more distinct secondary vein arches. The more sparsely pubescent androecious flow- ers are in short, ramiflorous pleiochasia of up to five flowers, or rarely solitary and axillary or sub- terminal. Apart from the fact that the anther loc- uli are basally confluent in Bernardi 11759 (Fig. 36B), the narrowly acute to subulate stamens with long connectives and closed, globular androe- cious flowers most closely resemble those of 7. kirkii and T. paradoxa from nearby Anjouan, except that they contain dark tanniferous idio- blasts. In terms of androecious floral and sta- minal morphology, Bernardi 11759 could rep- resent an undescribed species. The specimen at Geneva, however, has a single corky brown im- mature fruit with the orifice bearing six deltoid lobes that corresponds well with Bernardi 11781, and for this reason I tentatively consider them conspecific. On the basis of androecious floral and staminal morphology, Tambourissa moheliensis, T. kir- kii, and T. paradoxa would appear to be derived from a common ancestor. However, unlike the female receptacles of T. kirkii, T. comorensis, and T. leptophylla (all Group 2), which all have a circular, entire orifice open even in bud, that of T. moheliensis is closed in bud and splits open by four, deltoid segments as in 7. capuronii (Madagascar). Moreover, the floral ground tissue 1985] dark of both the latter sp tal brown tanniferous idioblasts not found in the other Comorean species. These characters sug- gest that 7. moheliensis was derived from a dif- ferent ancestor than the other Comorean species and appears most closely allied to 7. capuronii with which I have placed it in species Group 3 Vernacular name. | Diarou. 20. Tambourissa capuronii Cavaco, Bull. Mus. Hist. Nat. (Paris), Ser. 2, 29: 287. 1957; Ca- vaco in Humbert, Fl. Madagascar 80: 20, fig. VI:5-8. 1959. TYPE: Madagascar. Diego Suarez: Betsomanga Massif, valley on the S flank, ca. 550 m, 17 Nov. 1950 (fl), Capuron 820-SF (holotype, P; isotype, P). Large monoecious tree 25-30 m tall and 40- 60 cm D.B.H., the new growth hirtellous, the mature leafy stems glabrous, petiolate; petioles 12-30 mm by 1.2-1.5 mm; lamina chartaceous to subcoriaceous, abaxially hirtellous to glabrate, elliptic to oblong or obovate, (68-)90-165 mm by 30-74 mm, the apex shortly acuminate, acute or rarely obtuse, the base cuneate to acute, rarely obtuse, the secondary veins 7-10 pairs, making a 55-65? angle with the costa, the venation prom- inent, visible to 4? on both surfaces, the margin moderately revolute. Inflorescence a cauliflo- rous, unisexual or sexually mixed pleiochasium of 4-15 flowers, or rarely a fascicle of 2-3 flowers, all parts corky brown externally, the floral axis 10-170 mm by 1-5 mm. Androecious flower in bud globose, 15-19 mm diam., corky brown, slightly apiculate with coalescent tepals, the ped- icel 28-55 mm by 2-2.5 mm; at anthesis deeply 4(-5)-fid, 40-55 mm diam., the lobes spreading flat; stamens numerous, ca. 300, lanceolate, 6-8 mm long by 1-1.5 mm wide basally, subsessile, the loculi narrow, separate, lateral, occupying the basal ⁄⁄—⁄ of stamen, the filament short or lack- ing, the connective acuminate, slightly pro- longed, comprising apical 4-4 of stamen, the apex sometimes bearing up to 5 simple hairs, the internal receptacle surface glabrous or with scat- tered simple hairs. Gynoecious flower in bud na- piform-depressed, corky brown, 15-20 mm diam.; at anthesis to 35 mm long by 40 mm wide, the apex splitting into ca. 6 thick, irregularly del- toid lobes; styles numerous, ca. 200—300, nar- rowly conical, slender, 4-5 mm long by 0.5 mm wide basally, solitary or coalescent into groups of up to 12, the internal receptacle surface dense- ly pilose between the styles and around the ori- LORENCE— MONIMIACEAE 131 200 km : aas \ * # D Aux A - if e k eee a | ; C | a 4 J i ( Pa uot. ate 7 | A ( "i pee. J J l pee / | vA 3 Jf | ( & f if D 4 2 à v a / L "i A B / í — / O T. capuronii M UM E T. gracilis A / A T. purpurea r d EN à @ T. religiosa FicunE 37. Distribution map of some Tambou- rissa species in Madagascar. fice; pedicel to 75 mm by 5 mm. Submature fruiting receptacle cupuliform, externally corky, 35 mm long by 55 mm diam., the carpels ovoid- compressed, to 13 mm by 8 mm Distribution. Endemic to Madagascar (Fig. Habitat. Contrary to Cavaco's original (1957d) statement that the species was a highly restricted endemic, his subsequent list of exsic- cata (1959) shows Tambourissa capuronii to be fairly widely distributed both in lowland Myris- ticaceae/Anthostema wet forest from ca. 300 to 800 m, and also in Tambourissa/ Weinmannia montane wet forest from ca. 800 to 1,400 m, ranging from Mt. D'Ambre (Diego Suarez) in the north, south along the eastern escarpment, to the Mandrare River near Fort Dauphin in the south- east. The species was said by Capuron to be very abundant in 1950 at Betsomanga between 350 132 and 600 m. A canopy tree reaching ca. 30 m, it flowers in November-December and is one of the largest members of the genus. MADAGASCAR. DIEGO SUAREZ: forét d’Ambre, 1,200 Perrier de la Báthie 10087 (P); Perrier de la Báthie 10095 (P); Antasibe, Soanierana, E coast, 350 m, 10 Dec. 1938 (fl), Lam & Meeuse 5835A (P). TULEAR: upper basin of the Mandrare River (SE), pass and sum- mit of Marosoui, forest on gneissic laterite, 1,000-1,400 m, 14-15 Nov. 1928 (fl), Humbert 6606 (B, G, 2 sheets; K, MAD, P). Tambourissa capuronii appears to be most closely related to the Comorean 7. moheliensis with which it comprises species Group 3. The latter is distinguishable by its much smaller An- droecious flowers and stamens, and styles which are never coalescent into groups as in 7. capu- ronii. None of the other Madagascan species ap- pears to be closely allied to 7. capuronii. 21. Tambourissa religiosa (Tul.) A.DC., Prodr. 16(2): 659. ; Perk. & Gilg, Pflanzenr. 4, 101: 71. 1901; Drake: in Grandidier, Hist. Phys. Madagascar 1(1) 23. 1902; Perk., Pflanzenr. 4, 101 (Nachtr.): 44. 1911; Ca- vaco in Humbert, Fl. Madagascar 80: 36. 1959, excluding fig. X:5 (which is T. elliptica Mo : *Bourbon" (Réunion). (Fide Richard, but probably Madagascar), ca. 1851 (fl), Boivin s.n. (lectotype, P, here designated). Ambora rond Boiv. ex Tul., Monogr. Monim. n Lam. ex Steud. E en boivinii A. DC., Prodr. Piu 659. 1868; k & , Pflanzenr. 4, 101: 1902; Perk., Pflanzenr. 4, 101 Rue 42. 1911. TYPE: Madagascar. Tamatave: Sai land, on forest, Mar. 1847 (5), Boivin 1 728 (holot Tambourissa “uddrifida sensu Drake in Grandidier, t. Phys. Madagascar 1(1): 21. 1902, pro parte e to Baron 2504), non Sonnerat. Monoecious shrub, treelet, or small tree 4-5 m tall, the new growth glabrous, the mature leafy stems ascendant, dark brown to stramineate, 2.5— 6 mm diam. Leaves opposite to subopposite, gla- Ride o petioles (10-)15-30 mm (1—)1. 5 mm; lamina subcoriaceous to cori- aceous, pd elliptic, elliptic, oblong, narrowly oblong or obovate, 70—150(-175) mm by (16-)30- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 60 mm, the apex shortly acuminate, bluntly acute, obtuse or retuse, the base acute, cuneate or ob- tuse, often + decurrent, the secondary veins 6- 10 pairs, making a 60-70? angle with the costa, the venation obscure or visible to 2-3? adaxially, visible to 3-4? abaxially, the intramarginal loops uniting to form a conspicuous, partial marginal fimbrial vein, the margin thickened, moderately to strongly revolute. Inflorescence glabrous, either a terminal, axillary, or ramiflorous dichasium of 3 flowers or condensed pleiochasium of 5-7 flow- ers (the gynoecious flower terminal, if present), or the flowers solitary and terminal or axillary, the floral axis 1-7 mm by 1.5-2 mm, the axis and pedicels subtended by several glabrous to sparsely ciliate, broadly deltoid bracteoles with pale margins. Androecious flower in bud black- ish to yellowish, globose to napiform, 7-10 mm diam. 4-6 mm long, + depressed, apiculate with 2-4 minute, deltoid glabrous tepals, the pedicel 3.5-5 mm by 1-1.5 mm; at anthesis un- known, presumably splitting into 4 shallow lobes (as in 7. purpurea); stamens in bud ca. 60-82, linear-ellipsoid to slightly ovoid, 0.8-1 mm long by 0.5-0.7 mm wide, the filament short, distinct, slender, the loculi lateral, separate (rarely con- fluent apically), the connective rarely prolonged into an apicule, the internal surface with sparse hairs. Gynoecious flower in bud napiform-de- pressed, the pedicel 6-7 mm by 1-1.5 mm; at anthesis 8-11 mm diam. by 5-7 mm long, the orifice shallowly 4-lobed, X-shaped, 0.5-1.5 mm wide, the walls thick; styles numerous, ca. 300, conical, acuminate, slightly ventricose, 0.5—1.7 mm long by 0.3-1.5 mm wide basally, the in- ternal receptacle surface with clusters of simple hairs between the styles. Fruiting receptacle ter- minal, solitary, or rarely paired, napiform-de- pressed to subglobose, to 52 mm diam. by 40 mm long, th th, dark brown with corky patches, the orifice ca. 12 mm diam., comprising '4-'4 total width of receptacle, sub- circular, subentire, the walls 10-17 mm thick, the internal surface with scattered conical-co- lumnar styles 0.3—0.5 mm long and wide basally, interspersed with scattered hairs. Mature (?) car- pels ovoid- epe 7-9 mm long, 5-6 mm wide, 4-5 mm thick, the endocarp light brown to beige, the Du shallowly scrobiculate. Distribution. The species is endemic to Mad- agascar. Cordemoy (1895) erroneously cites the species as occurring in Réunion, but was actually referring to Tambourissa crassa (q.v.). The lec- LORENCE— MONIMIACFEAF 133 1985] totype of 7. religiosa was said by Richard to have come from Bourbon (Réunion), but this is un- doubtedly due to a mixup in herbarium labels, because his other collections of the species are all from Madagascar and no other authentic spec- imens are known from Réunion (Fig. 37). It was selected because it represents the best of the four syntypes. The type of 7. boivinii, although in poor condition, is clearly referable to T. religiosa. Habitat. In Madagascar Tambourissa reli- giosa appears to be commonest in the eastern littoral antl where it occurs sympatrically with T. purpurea. The species ranges inland to the mountainous central plateau from altitudes of 1,500 to 1,700 m, and across the northwestern region to the offshore island of Nosy Be. DAGASCAR. DIEGO SUAREZ: Nossi Be, 23 July 1840 (st), Pervillé 295 (P, syntype); 10 Oct. 1840 (fr), Pervillé 328 (P, syntype). TAMATAVE: Ambila, 1 May 1928 (fl), Decary 6288 (MAD, P); 10 ee 1928 (fl, fr), Decary 6554 (K, MAD, P); Rantabe, N of Tamatave, littoral forest, 1 "June 1969 (st), Guillaumet 3002 bis (MAD); Maroa, forests within Antongil Bay, 1867 (fl), Mo- querys 66 (G, Z); Reserve no. 1, Tamatave, 21 June 1950 (fl), Rakotoniama sub RN 3956 (MAD, 2 sheets); Sainte Marie Island (East Coast), Sept. 1851 (fl), Boivin Lemaitsoa, littoral forest, Aug. 1975 (fl, fr), Rakotozafy 1293 (MAD); Androndramanika, Rahobevava, 830 m (near Lac Alaoatra), 10 May pd (fr), Cours sub Herb. is 4270 Seager’ WITHOUT PRECISE LOCALITY: Cen- a scar, 1883 (fl), Pon 2499 (K); (fl), Baron 25 04 (K); p 883 (fl), Baron 2766 (K, 2 sheets); East Coast, 91 (fr), Baron 6043 (K); 1865 (fl), Ger- rard 35 (K); us (fl), Humbolt 190 (K). Ti } ] lj t Group species 4 and appears to be most closely allied to T. purpurea, another Madagascan species with structurally similar leaves and flowers. Obser- vations on living material of 7. religiosa, how- ever, are required to ascertain whether or not its gynoecious flowers produce a mucilaginous **hy- perstigma" as do those of 7. purpurea (studied by Endress, 1979 and 1980b, as T. religiosa). Ripe fruiting receptacles of both species are smooth and reddish as opposed to those of most congeners which are pale brown and corky. Tambourissa religiosa is distinguishable from T. purpurea by its floral receptacles which are more than twice as large, its more numerous sta- mens with separate loculi and distinct filaments, its more numerous, acute styles, and by its larger, more coriaceous leaves with a thicker hypoder- mis, prominent partial marginal fimbrial vein, and revolute margins Vernacular names. Ambora, Laingafora (Madagascar). According to Commerson, certain Malagasy tribes used Tambourissa religiosa in a religious context, burying their dead with its branches 22. Tambourissa purpurea (Tul.) A.DC., Prodr. 6(2): 659. 1868; Perk. & Gilg, Pflanzenr. 4, 101: 69, fig. 18:5. 1901; Perk.. Pflanzenr. 4, 101 (Nachtr.): 42. 1911; Cavaco in Hum- bert, FI. Madagascar 80: 39, fig. V:1. 1959. Ambora purpurea Tul., Ann. Sci. Nat. (Paris) 4(3): 30. 1855; Tul., Monogr. Monim. 301, pl. XXVI. 1855. TYPE: Madagascar. Tama- tave: Sainte Marie Island (ca. 1850) (fl, fr), Bernier 262 (lectotype, P, here designated; isolectotype, P). um rota Baker, J. Linn. Soc., Bot. OPER 240. 3; Perk. & Gilg, Pflanzenr. 4, 101: 1901; [oa Pflanzenr. 4, P (Nachtr.): 42. T Perk., Gatt. Monim. fig. 1925; Cavaco in Humbert, Fl. pa cred ^a 35. — X:6. 1959. TYPE: Ma r. Central M. without —— locality po 1881) (fi, fr). on 764 (lec- T. rota var. PR Perk., Pflanzen (Nachtr.): 4 11. TYPE: Madagascar. Central adagascar: ee precise locality (fr), Baron 1642 (holotype, K). Monoecious shrub, peg or small tree at- taining 10 m tall and 30-4 .B.H., the bark gray-black, longitudinally “wisi the Dbranelibs often decumbent and trailing, the new growt glabrous or with rare, scattered hairs, the mature leafy stems brownish, 2- diam., terete. Leaves opposite to subopposite, rarely ternate on vigorous sucker shoots, glabrous, petiolate; pet- ioles 3—10(—17) mm by 0.8-1.3 mm, red to pur- ple when living; lamina chartaceous to subco- riaceous, ovate-elliptic, elliptic, oblong, narrowly oblong or obovate, (25-)40-85(-100) mm by (10-)15-—40(—44) mm, the apex acuminate, acute, obtuse or retuse, the base cuneate, acute, obtuse, truncate or even subcordate, often + decurrent, the secondary veins 4-11 pairs, making a 60—70? angle with the costa, the venation visible to 2— 4° or obscure adaxially, visible to 2—3? abaxially, the margin plane to slightly revolute. Inflores- cence ramiflorous, axillary, or terminal on leafy shoots, a 3-flowered dichasium or a contracted pleiochasium of 5 flowers (the gynoecious flower terminal, if present), or the flowers solitary, or in sparse fascicles on old stems, the floral axis 134 1-4 mm by 1 mm, glabrous to puberulent, the axis and pedicels subtended by 1 to several ciliate to hirsute, naviculate bracteoles. Androecious flower in bud sparsely pubescent to glabrous, glo- bose-depressed to napiform-depressed, 2-3(-4) mm diam. by 2.5-3.5(-4) mm long, the apex with 1—3 pairs of minute, acute to rounded puis tepals, the Uu 2.5-6(-10) mm by 0.6-1 m at anthesis 2.5-3.5 mm diam., shallowly aoe fid for up to '^ its length, the lobes incurved or spreading slightly, the orifice 1-3 mm diam.; sta- mens 20-32, acutely deltoid to obtusely deltoid, 0.7-1.2 mm long by 0.6-1 mm wide, the loculi confluent apically into a single, crescentiform lo- culus, the filament very short or sessile, the in- ternal receptacle surface glabrous or with rare, scattered hairs. Gynoecious flower in bud glo- bose-depressed to napiform-depressed, glabrous to sparsely pubescent, apiculate with 2-3 pairs of minute tepals, the pedicel 5-6 mm by 1 mm; at anthesis urceolate-globose to urceolate-napi- form, 4(-7) mm diam. by 4(-7) mm long, the orifice 3-4-lobed, 0.5-1 mm diam., when living occluded by a mucilaginous plug, the inner tepals stigmatic; styles 35-70(—90), shortly columnar, apiculate, 0.3-0.4 mm long by 0.2-0.3 mm diam., the internal receptacle surface glabrous or with rare, scattered hairs between the styles. Fruiting receptacle terminal, solitary, globose-urceolate, + depressed, 28-50 mm diam. by 25-40 mm long, the external surface dark reddish brown to blackish, occasionally mottled with pale gray- brown, corky patches, the orifice subentire, + circular, 2-7 mm diam., comprising '49—' total width of the receptacle, the walls 8—1 1 mm thick, the internal surface with scattered, subcircular styles ca. 0.5 mm diam., glabrous or with rare, scattered hairs. Mature carpels ovoid-com- pressed, 6-9 mm long, 4-7 mm wide, 3.5-5 mm thick, the endocarp brown, the surface scrobic- ulate to nes le ovens, Gametic chromo- some number, 9. Distribution. Endemic to Madagascar (Fig. Habitat. Tambourissa purpurea is one of the most widely distributed members of the genus in Madagascar, along with 7. hildebrandtii and T. religiosa. Quite variable vegetatively, it dis- plays a wide ecological amplitude ranging from littoral forest along the east coast, through the lowland and submontane wet forest zones of the eastern escarpment (up to ca. 1,400 m), and into montane forest of the central plateau (ca. 1,400— ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 1,600 m). It also extends westward into the semi- deciduous dry forest zone of the western sector as far as the Isalo massif, where it is apparently able to survive annual burning, taking on a shrubby habit with numerous erect coppice shoots. In living plants, floral receptacles of Tam- bourissa purpurea range from green to dark red- dish purple. The gynoecious flowers produce a mucilaginous plug or “‘hyperstigma” (see En- dress, 1979, 1980b) that is probably functional in pollination. Androecious flowers of Lorence 1725 (MO) produced a faint sweet, almost musky, odor. MADAGASCAR. FIANARANTSOA: Isalo, 13 Apr. 1961 (fl), Anon. sub Herb. Peltier 3044 (MAD); Ihosy, Mont Lalandro, 1,100-1,200 m, 5 Nov. 1967 (fr), Bernardi 11230 (K); Route no. 7, 39 km N of Ambositra, 1,250- 1,350 m, 24 Jan. 1975 (fr), Croat 29458 (MAD, MO); W of Ranohira, Route 7, 810 m, 3 Feb. 1975 (fl), Croat 30568 (MAD, MO); Ambohimitombo forest (Fanala), 1,350-1,440 m, 4 Jan. 1895 (fl), Forsyth Major 328 (K); (fl), Forsyth Major 340 (K); 21 Jan. 1895 (fr), of Ranohira (Isalo), 800 m, 6 Nov. 1978 (fr), Lorence 2058 (MAD, MO); (fr), Lorence 2059 (MAD, MO); (st), "dp 2060 (MAD, MO). MAJUNGA: Natural Re- serve no. 4, Marovato, Ambanja, 10 July 1952 (fr), Safy sub RN 4 180 (MAD, 2 Sheets). E Soan- jerana, 1) c. 1949 (fr), An on. sub SF 2346 (MAD); joue ien: “Foule Pointe, 12 June 1950 (fr), Anon. sub SF 2585 (MAD); Natural Reserve no. 3, Sahatavy, Fenoarive, 17 June 1952 (fl), Anon. sub RN 4137 (MAD); Vohimaranitra, Tamatave, 26 Jan. 1951 (fr), Anon. sub RN 1989 (MAD); Sainte Marie Island, Tafondron Forest, Mar. 1847 (fl, fr), Boivin 1729 (P, syntype, 2 sheets); Mora- rano, pseudosteppes on dry slopes (W of Lac Alaotra), Dec. 1954 (fl), Bosser 7502 (MAD); on sand between Mahanoro and Vatomandry, 2 July 1972 (fr), Cremers 2310 (MAD, 2 sheets); Ambila, littoral forest, 8 May km E arive d Oct. i (s), Lorence 2036 (MAD, MO); Perinet, 70 t. Gar den) on 1976 (fl), Rauh M 138 (Z). TANANARIVE: An- abe, Anjozorobe, Mar. 1953 (fr), Bosser 5219 (MAD, 2 sheets) € etsa, Ankazobe, Pk 1 Junga; Apr. 5 (fr), Bosser 7855 (MAD sin DE py (fl, fr), end 12466 (MAD, 2 sheets): "forest at Pk 142 along the road from Tanana- rive to Majunga, 8 July 1971 (fr), CINES 1641 (MAD); Tananarive, 1 dh Park, 1,200 m, 17 Jan. 1975 1985] LORENCE-— MONIMIACEAE 135 (fl), Croat 28636 (MAD, MO); near Tananarive, 28 July 1928 (fr), Decary 6610 (MAD); Tananarive, Tsim- bazaza Park, cultivated in back of Philippe Morat's house (originally from desi 28 May 1974, Gentry & Morat 11964 (MAD, MO); forests of Tampoketsa, May 1956 (fr), Rialinansy 1027 (MAD), bottom of the valley Ambohimanga south, 9 Nov tozafy 204 (MAD); “tapias” (Uapaca for fl 8 km fr 97 5 (O, oe » 778 (MAD, D, ); dunes, Fort Dau- phin (fr) b. Inst. Sci. Madagascar 29 (MAD); Fort — AE Rauh 1440 (MAD). PRO- VINCE UNCERTAIN: À ofa, Natural Reserve, 8 July 1950 "5 pit eg SF 43 (MAD), forest N of the road to Nickaville, 29 Dec. 1944 (st), Anon. sub Herb. Alaotra 2081 (MAD); Antsahabe to Antsakalal- ina, 20-40 m, 5 Apr. 1949 (fr), Anon. sub Herb. Alaotra 3607 (MAD); Andrangoalaka, Central Madagascar, 1881 (fr), Baron s.n. (K); 1882 (fl), Baron 1239 (K, syntype of T. p Nei Parker s.n. (K), Central Ma- 955 (fr), Bosser 8709 (MAD); An- (fr), Rakotozafy 228 (MAD); Tsaravinany, Apr. 1966 (fr), Rakotozafy 588 rota); (fr), Baron 2996 (K); 1883 (fr), Baron 3024 (K, P); 1885 (fr), Baron 3515 (K, P); (st), Baron 357 5 (BM); (fr), Baron 4241 (K, P); (fr), Baron 4325 (K); 1892 (st), Baron pU (BM); (st), Parker s.n. (K); North Mada- 892 (fr), Baron 6439 (K). CULTIVATED: Mis- gascar, (MO); (fl), Lorence 1726 (MO); 13 Feb. 1980 (fl), Lo- rence T 4 (MO). Although Tambourissa purpurea is extremely variable in terms of foliar morphology, even on a single individual (e.g., Croat 29458, MO), its flowers are quite constant and the smallest in the genus. Along with 7. gracilis and T. religiosa, it belongs to species Group 4. Flowers of T. reli- giosa, however, are about twice as large as those of T. purpurea, have more numerous parts, and the leaves are thicker with a well-developed hy- podermis of (3—)5—6 cell layers comprising ca. 4—V, of the total laminar thickness. That of T. purpurea is only (12-3 cell layers thick and comprises only !⁄—'!⁄ of the total laminar thick- ness. Also, T. purpurea lacks the prominent par- tial marginal fimbrial vein of the former species. Both occur sympatrically in eastern and central Madagascar and no hybrids or intermediates are known to occur. Tambourissa purpurea is perhaps even more closely allied to 7. gracilis in terms of foliar and ers are unknown in the latter). The pleiochasia are much laxer in T. gracilis, however, with a much longer floral axis and pedicels. Vernacular names. Ambora, Ambora vavy, Rota, Villainque-possa (Madagascar). 23. Tambourissa gracilis Perk., Pflanzenr. 4, 101 (Nachtr.): 43. 1911; Cavaco in Humbert, Fl. Madagascar 80: 35. 1959. TYPE: “Cen- tral Madagascar." Without precise locality (probably Tananarive or Fianarantsoa): ca. 1888 (fl), Baron 2885 (holotype, K; isotype, P). T. leptophylla sensu Drake in Grandidier, Hist. ET Madagascar 1(1): 23. 1902 non (Tul.) A Dioecious (?) treelet or small tree 8-10 m, the mature leafy stems glabrous, 1.5-2.5 mm diam. Leaves opposite, glabrous, petiolate; petioles 1 4— 7 mm by | mm; lamina chartaceous, elliptic or rarely oblong, 75-120 mm by 26-45 mm, the apex long acuminate, the base acutely cuneate, decurrent, the secondary veins 6—9 pairs, making a 65-85? angle with the costa, the venation raised and visible to 4? on both surfaces, the margin plane to slightly revolute. Androecious inflores- cence ramiflorous on leafless nodes or axillary, a unisexual pleiochasium of 3-7 flowers, sparsely pubescent Ne pale, appressed hairs, the floral axis 7-25 m ] mm, subtended by several basal ee Androecious flower in bud glo- ose, 6-8 mm diam., apiculate with a pair of scilioircuter: obtuse tepals, the pedicel 5-13 mm by 1 mm, subtended by 1(-2) hirsute, naviculate bracteoles 1-2 mm long; at (full?) anthesis glo- bose, 6-8 mm diam., the subcircular orifice 1-2 mm diam., lightly, flanked internally by several pairs of obtuse, scale-like tepals; stamens ca. 50, broadly deltoid, 1.2-1.7 mm lon: 1 mm wide, the loculi occupying almost entire length of the stamen, confluent api- cally into a single, crescentiform loculus, the fil- ament very short or sessile, the internal recep- tacle surface with rare, scattered hairs. Gynoecious inflorescence unknown. Submature fruiting receptacles borne in clusters of 2-3, shal- lowly cupuliform, 30-55 mm wide by 20-30 mm long, the orifice comprising 2⁄—⁄4 total width of T 136 fruit, the styles conical, 0.6—0.8 mm long; fruiting carpels ovoid-compressed, 9-10 mm long by 7- 8 mm wide, tan. Distribution. Endemic to Madagascar (Fig. 37). Habitat. According to Humbert (1965) and Cavaco (1959), Tambourissa gracilis occurs in high altitude ericoid heath formation (ca. 1,800- 2,700 m) on mountains of the central plateau and vicinity. However, a fruiting specimen (Humbert 3688, P) named by him and cited by Cavaco (1959) is closer to 7. purpurea, from which it differs in having disorganized venation characteristic of neither species. As the specimen lacks flowers, a more complete analysis is im- possible. MADAGASCAR. DIEGO SUAREZ: Tsaratanana Massif, 1,400 m, Nov. 1912 (fr), Perrier de la Bathie 10105 (P). Tambourissa gracilis belongs to species Group 4. With respect to its androecious flowers, which have short, subsessile stamens and apically con- fluent loculi, it appears to be most closely related to T. purpurea which differs in having a shorter floral axis and pedicels. Foliar morphology of the two species is virtually identical, also resembling that of T. madagascariensis (Group 2A). Further collections of this imperfectly PNA species, es- pecially g uits, are neede to better understand its ios 24. Tambourissa pedicellata Baker, Fl. Mauri- tius 289. 1877; Perk., Pflanzenr. 4, 101 (Nachtr.): 42. 1911; Vaughan, Mauritius Inst. Bull. 1(1): 76. 1937. TYPE: Mauritius. With- out precise locality; in the shade of humid forests, on the high mountains, 1864 (fl, fr), Bouton s.n. (lectotype, K). Ambora amplifolia sensu Tul., id Monim. 299. 1855, pro p odr. 16(2): 659, 1901, pro parte, non (Boj ex Tul.) A. Monoecious tree attaining 15 m tall and 30 oth, fla puberulent, glabrescent, the mature leafy stems iam., terete, the nodes dialated. Leaves opposite, rarely subopposite, glabrous, petiolate; petioles (2-2)6-13 mm by 1.5-3 mm; lamina chartaceous to coriaceous, elliptic to nar- rowly elliptic or oblong, rarely broadly elliptic to ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 suborbiculate, (30-)50-160(-200) mm by (15-) 22-80(-120) mm, the apex acuminate, acute, ob- tuse, rounded or retuse, the base acutely decur- rent to acutely cuneate, the secondary veins 5- 12 pairs, making a 50-65? angle with the costa, the venation obscure adaxially and visible to 2(-3°), dark brown abaxially and visible to 3(-45), the margin slightly thickened, moderately revolute. Inflorescence a glabrous, ramiflorous (rarely terminal), leafless or leafy, unisexual or sexually mixed pleiochasium of 5-14 flowers, or a cauliflorous, leafy or leafless pleiochasium or rarely dichasium, the floral axis (15-2)35-200 mm by 1.5-4 mm, often + flattened, frequently sub- tended by several deltoid-naviculate puberulent bracteoles; flowers monomorphic. Androecious flower in bud napiform-depressed, 7-11 mm diam. by 4-7 mm long, externally smooth with scattered hairs, the apex with 4 minute, puber- ulent tepals, the pedicel long, slender, 15-66 m by 0.7-1.5 mm, subtended by a single puberu- lent, subulate-deltoid bracteole 1-2 mm long or by a small leaf; at anthesis shallowly 4-6-fid, subglobose, 10-14 mm diam., the lobes thick, short, acuminate, incurved to erect, the orifice comprising !^—/ total width of the receptacle; stamens 50-70, minute, deltoid to ellipsoid, 0.6- 1 mm long by 0.5—0.8 mm wide, the loculi short, broad, subparallel, separate or occasionally con- fluent apically, sometimes + unilateral, occu- pying 75—A total length of the stamen, the fila- ment short, basally ventricose, the connective not prolonged, the internal receptacle surface gla- brous. Gynoecious flower in bud subglobose to napiform-depressed, 8—10 mm diam. by 5-7 mm long, externally smooth with scattered hairs, the apex with 3-4 minute, puberulent tepals, the pedicel as in the androecious flower; at anthesis shallowly 4—7-fid, subglobose, 6-12 mm diam. by 5-7 mm long, the lobes thick, acute to acu- minate, erect, the orifice subentire, comprising ca. !5—/ total width of the receptacle, the walls thick; styles ca. 60-1 80, conical, 0.8—1.2 mm long by 0.3-0.4 mm wide basally, the apex slightly papillose, the base ventricose, the internal re- pleiochasium of up to 5 fruits, + subglobose, 18- mm diam. by 12-25 mm long, the orifice small, shallow, comprising ca. %4—'2 total width of the receptacle, the rim subentire, thickened, with persistent deltoid lobes, the external surface mottled, pale corky brown with darker patches, LORENCE-— MONIMIACEAE 137 1985] 2090's + 63930'E A T. amplifolia A T. O T. B T. . tau ficus pedicellata quadrifida FiGuRE 38. Distribution map of some Tambourissa species in Mauritius. the cavity small, the numerous styles conical, 0.6-1 mm long by 0.4—0.7 mm diam. basally, interspersed with rare, scattered hairs, the ped- icel slender, 20-55 mm by 1.5-2.5 mm. Mature carpels ovoid-compressed, 8-11 mm long, 6-8 mm wide, 4—6 mm thick, the endocarp dark brown with 1 to several longitudinal, + branch- ing ridges, the surface ms scrobiculate. Ga- metic chromosome number, 7 — 19. Distribution. The species is endemic to Mau- ritius (Fig. 38). Habitat. Tambourissa pedicellata is present- known only from four localities in wet and cloud forest: the southern flank ofthe Pieter Both mountain range, where I estimate the total pop- ulation to consist of 50 to 100 individuals; at Mt. Lagrave, where a small population occurs in cloud forest near the summit; in Bassin Blanc crater, where a single individual was located. Vaughan and Wiehe (1941) recorded 7. pedicellata from Macabé forest (Vaughan 1327 sub MAU 2358, MAU), but I was unable to relocate it there dur- ing extensive searches made in 1974-1977 and again in 1978-1979. In 1983, W. Strahm (pers. comm.) located an “individual” of 7. pedicellata at Macabé whose trunk was fused with that of T. sieberi in what apparently represents a perfect natural graft. My observations on the Pieter Both popula- tion during the course ofa year showed the plants to be evergreen and not deciduous as stated by Vaughan and Wiehe (1941). Most individuals 138 here flowered from February through April 1979, although one tree was flowering in November 1978. Fruits generally ripened in February. Liv- ing flowers of T. pedicellata produce little ap- parent odor, although a faint, fruity smell was detected in some. Buds and receptacles are mot- tled green externally, and the stamens may be cream, yellow, or yellowish pink, whereas the styles range from bright orange to reddish orange. MAURITIUS. BASSIN BLANC CRATER: Wet forest near lake, ca. 500 m, 6 Fea 1979 (st), Lorence 2967 (MO). ACABE FORES acabé transect (symbol T,) 16 June 1937 (st), pugil. 1327 sub MAU 2358 diei MT. LAGRAVE: S slope facing Eau Bleue reservoir m- mit, ca. 600—628 m, 24 May 1979 idt Mrs 2966 e) : S flank, 500- 1978 (fl), Lorene 1834 (K, MAU, 979 (fl), Lorence 2384 (K, MAU, MO, 2 sheets; Z); (f, fr), Lorence 2385 (B, G, K, MAU, MO), (fl, fr), Lorence 2386 AU, MO, P, REU); 19 Apr. 1979 (fl), Lorence 2598 (MAU, MO); (fl), Lorence 2599 (MAU, MO). QUARTIERE MILITAIRE: P (MAU); 1814 (fl, fr), Carmichael s.n. (K); ca. 1755 (fl), Commerson s.n. (P); (st), Anon. sub Herb. Ventenat s.n. (G, 2 sheets). In terms of floral morphology, Tambourissa pedicellata appears to be one of the least spe- cialized species in Mauritius and possibly in the entire genus. It appears to fit best in species Group 5, however. Among Mauritian species, 7. pedi- cellatais most similar to T. amplifolia which also has long, slender floral pedicels, small subglobose buds and receptacles, and small stamens and styles; the two species have been confused in the past (De Candolle, 1868; Perkins & Gilg, 1901). Tambourissa amplifolia is readily distinguish- able, however, by its smaller, more slender habit, basal cauliflory, much larger leaves, discoid gy- noecious flowers with everted lobes, and an- droecious flowers deeply split into four flat lobes car) also has monomorphic a noecious flowers, it appears to represent a highly specialized condition because of the mucilagi- nous "hyperstigma" of the gynoecious flowers (see Endress, 1979, 1980b; as T. religiosa). The majority of Mauritian species with opposite leaves, more or less open, cupuliform to napi- form or globose gynoecious flowers and conical to setose styles also belonging to Group 5 may have been derived from a common ancestor ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 which 7. pedicellata most closely approaches to- day. 25. Tambourissa amplifolia (Bojer ex Tul.) A.DC., Prodr. 16(2): 659. 1868; Baker, FI. : o parte; Perk., Pflanzenr. 4, 101 (Nachtr.): 43. 1911. Mith- ridatea amplifolia Bojer, Hortus Maurit. 290. 1837 nom. nud. Ambora amplifolia Bojer ex Tul., Monogr. Monim. 299. 1855. TYPE: Mauritius. Woods of Grand Port: Pouce Mountain, Aug. 1851, Boivin s.n. (holotype, P) Monoecious understory treelet or small tree attaining 8 m tall and 9 cm D.B.H., sometimes shrubby, the branches few, erect, the new growth sparsely hirsutulous, the leafy stems terete, 5-14 mm diam., glabrous, the nodes dialated. Leaves often clustered near the ends of the branches, opposite to subopposite, sometimes congested into pseudoverticels of 4, glabrous, petiolate; pet- ioles 14-30 mm by 4-8 mm; lamina chartaceous to subcoriaceous, elliptic to ovate or narrowly ovate, sometimes + falcate, 150—450(-500) mm by 80-230(-240) mm, the apex acute to acu- minate, the base obtusely decurrent to obtusely cuneate or acutely cuneate to acutely decurrent, the secondary veins 7—9 pairs, making a 60-70? margin plane to slightly revolute; seedling and sucker leaves heterophyllous, with 1—3 pairs of serrate teeth in the apical portion, the petiole red when living. Inflorescence basally cauliflorous (exceptionally ramiflorous), on meristematic swellings up to ca. 1 m above ground level, the flowers solitary, fasciculate or in a unisexual or sexually mixed pleiochasium of 3-7, the gy- noecious flower often terminal, the floral axis 5- 20(-45) mm by 1.2-2 mm, finely pubescent, ba- sally bracteolate. Androecious flower in bud glo- bose, puberulent, 7-8 mm diam., apiculate with 4 minute tepals, the pedicel 25-60 mm by 0.8- l mm, pubescent, subtended by a Toinute, ca- 4-fid, 14-18 mm diam., the lobes thick, spreading al- most flat or slightly incurved, the internal recep- tacle surface glabrous; stamens numerous, ca. 100-250, obovoid to clavate, 0.8-1.5 mm long by 0.5-0.6 mm wide, the loculi confluent at the obtuse apex (very rarely separate), occupying the apical !^—^ of the stamen, the connective not LORENCE-— MONIMI ACEAE 139 1985] p appui: the filament distinct, slender. Gy- w. several pairs of minute tepals, the pubescent ped- icel 38-80 mm by 1.5 mm; at anthesis shallowly napiform- discoid, "P 13 wide by 4-6 mm long, pli 11 4( 6) everted A de eltoid lo bes the orifice + square or cube comprising ca. !^ total width of the receptacle; styles nu- merous, ca. 300, shortly conical, 0.6-1 mm long by 0.2-0.3 mm wide basally, the internal recep- tacle surface velutinous with dense, simple hairs between the styles. Fruiting receptacle solitary at the base of the trunk (exceptionally on the branches), napiform-urceolate or subglobose, 30- 56 mm pen 22-45 mm long, the orifice ca. 25 mm rising ca. !^ total width of the i. EN bearing remnants of the indurated lobes, the walls ca. 12 mm thick, the external surface corky brown, bumpy, with sev- eral prominent veins, the internal surface with numerous shortly conical styles 0.6-1 mm long by 0.2-0.3 mm wide basally, interspersed with scattered simple hairs, the pedicel and peduncle (15-)35-80 mm by 4-5 mm. Mature carpels ovoid-compressed, 12-13 mm long, 6-8 mm wide, 4-5 mm thick, the endocarp light brown, with an irregular network of low ridges, the sur- face slightly foveate-scrobiculate. the apex r Distribution. Endemic to Mauritius (Fig. 38). Habitat. Taml amplifolia is local and occasional to common in the lawet montane wet forest zone at Bel Ombre (200-400 m), at Plateau Colophane below Macabé Forest (ca. 500 m), on the summit of Mt. Corps de Garde (ca. 750 m), in cloud forest on Mt. Lagrave (500—628 m), and in moist forest on the crest of Piton du Fouge (650 m). MAURITIUS. BEL OMBRE FOREST: Bel Ombre Il Reserve, 7 Dec. 1971 (fl), Guého jd ro 15195 (K, m,27D 1978 (st), Lo- O REU); (fl), . (MO, P); 10 Jan. 1979 (fl), Lae 2259 (MAU, MO); 16 Feb. 1979 (fl), sl 2408 (MO); 3 July 1979 (st), Lorence 2691 (MO, 2 sheets); (fr), Lorence 2692 (MO). CORPS DE GARDE MT.: summit ridge, + stunted forest, 15 Nov. 1973 (fl), Coode et N. VUA (K, MAU); 8 Nov. 4928 (MAU, 2 sheets). e, low, dryish , 24 Jan. 1976 (fl), PUER 1629 AU). ers FOREST: Plateau Colophane below Macabé, ca. 600 m, 15 Jan. 1979 (st), Lorence 2280 (MAU, MO); ek Lorence 2281 (MO); near Macabé, uplands, ca. 600 (MAU); Macabé high forest, 6 Vaughan sub MAU 1603 (MAU). MT LA š 600 m, 19 Feb. 1979 (st), Lorence & Lecordier 2362 (MO). WITHOUT PRECISE LOCALITY: (fl), Anon. s.n. (BM); 1863 (fl), Anon. sub Herb. Blackburn s.n. (K); (fr), Anon. : .a NUN 1814 (st), Carmichael s.n. (K); ca. 1755 (fr), Commerson s.n. (P); (fl), Commerson 67 (P, 2 sheets); (fl), s.n. sub Herb. Juss. 16708 (P-JUSS; MO, photo); (st), Ilardwicke s.n. (G). Tambourissa amplifolia belongs to species Group 5 and appears to be most closely related 1 4 to T. tau, which differs inl small- er leaves consistently aggregated into pseudo- verticels of two to four pairs separated by long “internodes,” by its T-shaped stamens with sep- arate loculi, and by its long, setose styles, in internally gl ing carpels of 7. amplifolia, T. pedicellata, ae T. tau all have endocarps with a series of low, X branching ridges, a feature unknown in other species and perhaps suggestive of a close affinity between these species. Bel Ombre forest harbors the largest known population of Tambourissa amplifolia, where it occurs sympatrically wit tau. No interme- diates or suspected hybrids were found here de- spite concurrent flowering times from December to Febru ake as that approach Tambourissa tau, however, occur in the eastern mountain ranges of Mt. des Créoles (Lorence & Julien 2337, MAU, MO) and also Piton Bambou (Staub sub MAU 13493, MAU). Although these plants resemble T. amplifolia in having large, acuminate petio- late leaves, their flowers most closely approach T. tau in being axillary (in Staub sub MAU 13493) and externally corky brown with multiple, in- curved orifice lobes and long, setose styles (in Lorence & Julien 2337). Stamens are critical in distinguishing the two species; unfortunately, an- droecious flowers are absent in both collections. Another intermediate collection, without precise locality (Anon. sub Herb. Brown s.n., K), ap- proaches T. tau in having ramiflorous, shortly pedicellate fruits, but the shortly conical styles are characteristic = T. amplifolia, androecious flowers are absent a The holotype of bar E amplifolia, either 140 from Grand Port or the Pouce Mt., possesses abnormally short floral pedicels (8-10 mm long) and almost glabrous buds. Although lowland forest i in the Grand Port region no longer exists, Boivin's type is similar to recent collec- tions from the nearby Mts. Bambous and Des Créoles in some respects. The type's obovate, obtuse stamens with confluent loculi are char- acteristic of the species as a whole, as are the broad, petiolate leaves, so there can be no doubt distinct over much of their range, particularly at Bel Ombre where I studied them, I have decided to regard them as distinct species. Living flowers of Tambourissa amplifolia have externally yellow-green to purple-red recepta- cles, cream to pale orange-yellow stamens, and pale yellow-pink to dark purple styles. A vague, faintly fruity odor is produced by the flowers. Vernacular name. Bois tambour (Mauriti- us). 26. Tambourissa tau Lorence, Bull. Mus. Hist. Nature Reserve: wet forest with Epis dominant, 600 m, 8 Oct. 1978 (fl), Lorence 1835 (holotype, MO; isotypes, B, K, MAU, P, Z) Little-branched monoecious understory tree- let, rarely shrubby, 1—4 m tall, the stems to 75 mm diam., the new growth sparsely puberulent or with rare, scattered hairs, the mature leafy stems glabrous, terete, 4-7 mm diam. Leaves opposite, decussate, clustered into pseudoverti- cels of 2-4 pairs, sessile to subsessile, rarely shortly petiolate, glabrous; petioles stout, 1-8 (-15) mm by 3-6 mm; lamina subcoriaceous to coriaceous, broadly elliptic to narrowly elliptic, obovate or oblanceolate, rarely ovate, (70—)1 50- 300 mm by (30-)45-130 mm, the apex shortly acuminate, shortly acute, obtuse, rounded or re- tuse, the base subcordate, rounded, obtuse, cu- neate or acutely cuneate, the secondary veins (4—)6-9 pairs, making a 50—65? angle with the costa, the venation raised, visible to 3(—4?) adax- ially, visible to 4(—5°) abaxially, the margin slightly to moderately revolute; seedling leaves hetero- phyllous with 2-3 pairs of short, serrate teeth in the apical portion, the petioles red when living. Inflorescence cauliflorous on meristematic swell- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 es along entire length of the stem, occasionally miflorous or axillary, the flowers solitary, in a de or dichasium of 2-3, or rarely in a short pleiochasium of 3-6 flowers, sexually mixed or not, the floral axis 2-3(-5) mm by 1-1.5 mm, + corky, glabrous or sparsely puberulent basally, subtended by several small, ciliate, naviculate- deltoid bracteoles. Androecious flower in bud rown, mottled, the pedicel 6-20 mm by 1-1. mm, basally bracteolate; at anthesis 4—6-fid, 20— 5 mm diam., the lobes spreading flat, the in- ternal icc Drache surface glabrous; stamens nu- merous, ca. 100-150, T-shaped, 1-1.5 mm long by 1.5-2 mm wide, the loculi separate on lateral arms (rarely + sessile, but never confluent), the filament distinct, slender, the connective slightly prolonged apically or not. Gynoecious flower in bud turbinate-depressed to napiform-depressed, 7-10 mm diam. by 4-5 mm long, externally corky brown, apiculate with several minute tepals, the pedicel 6-20(-40) mm by 1-1.5 mm, subtended by several pairs of deltoid pipe at anthesis napiform-urceolate, 10—12 mm diam., the orifice splitting into 5-10 thick, ENS deltoid, straight or incurved lobes, the orifice 3-6 mm diam., comprising ca. 3—'2 total width of recep- tacle; styles numerous, ca. 80-100, slender, se- tose, 2-3 mm long by 0.3-0.5 mm wide at the ventricose base, the internal receptacle surface glabrous. Fruiting receptacle solitary on the main stem or major branches, rarely axillary or sub- terminal, shallowly cupuliform-patelliform to subglobose, 25-50 mm diam. by 10-27 mm long, the orifice 10-26 mm diam., comprising ca. !^ total width of the receptacle, subentire with 5—6 indurated deltoid lobes, the walls 10-14 mm thick, the external surface corky brown, bumpy, the internal surface with numerous scattered, se- tose styles 1.5-3.5 mm long by 0.3-0.4 wide ba- sally, otherwise glabrous, the pedicel stout, 8-20 y 2-7 mm. Mature carpels ovoid-com- pressed, 10-14 mm long, 6-8 mm wide, 4-6 mm thick, with occasional low, irregular ridges, the surface scrobiculate to foveate. Gametic chro- mosome number, 7 — 19. Figure 39. Distribution. Endemic to Mauritius (Fig. 38). Habitat. This is one of the commonest and most widespread members of the genus in Mau- ritius, occurring in most medium to upper alti- tude wet and cloud forest communities from ca. 300 to 800 m. 1985] LORENCE—MONIMIACEAE 141 Ficure 39. Tambourissa tau Lorence.—A. Habit. —B. Androecious bud and newly opened androecious flower, lateral view . Stamen, lateral view. — D. Gynoecious flower in bud, lateral view. —E. Gynoecious flow ad — sera view. — F. Gynoecious flower, longitudinal section. — G. Submature fruiting receptacle, lateral v —H. As in G, apical view.—I. Fruiting carpel, lateral view. A. Perrier Nature Reserve, Mauritius rence spesa unvouchered). B-F. Lorence 1835 (MO). G, H. Lorence 2950 (MO). I. Lorence 2629 (MO). Bars equal 10 mm in A, B, D-I, and 1 mm in C. 142 MAURITIUS. BASSIN BLANC: side of track to E side of lake, 550 m, 25 May 1976 (fr), Richardson et al. 4163 (K). BEL OMBRE FOREST: ca. 200—400 m, 27 Dec. 1978 (fl), Lorence 2215 (MO); 10 Jan. 1979 (fl), Lorence 2257 (MO); (fl), Lorence 2258 (MO). CUREPIPE BOTAN- ICAL GARDEN: near lake, 30 June 1966 (fl), Vaughan sub MAU 12203 (MAU, 3 sheets). GAULETTES SERREES: low mixed wet forest over lava, 400-450 m, 19 May 1979 (fl), Lorence et al. 2641 (MO). GRAND BASSIN: upland thicket, 22 Sept. 1932 (st), Orian sub MAU 47 (MAU). MACABE FOREST: high forest, 660 m, 18 Oct. 1978 (fl), dig d al. 1863 (MO); Macabé to Brise Me road m, 8 Dec. 1978 e» Lorence 2102 (MAU, MO); 758 Ds 2103 (MO); Mare Longue to Ma- low indigenous Tun E Nov. 1962 (fl), 0472 (MA DLANDS: vestige ; S 1975 (MO); (f), Lorence 1976 (MAU, MO); (fl), Lo- rence 1977 (MAU, MO); 19 Dec. 1978 (fl), Lo Lecordier 2161 (MAU, MO); (st), Lorence & Lecordier ev- ergreen forest, 250—300 m, 25 Jan. 1979 (fl), Lorence & je 2337 (MAU, MO); (st), Lorence & Julien s.n. (M U). MONTAGNE LAGRAVE: flank facing Eau Bleue reservoir, 5 Apr. 1974 (fr), Coode et al. sub MAU 17615 rence & Lecordier 2363 (MO); S slope facing Eau Bleue reservoir, cloud forest invaded by exotics, 600 m May 1979 (fr), Lorence & Lecordier 2649 (MO). PERRIER NATURE RESERVE: dense indigenous thicket, 14 May 1970 (fr), Guého sub MAU 14297 (MAU); lower mon- tane wet forest, ca. 600 m, 8 Oct. 1978 (fl), Lorence 1836 (MO); (f), Lorence 1837 (MO); 13 Oct. 1978 (st), Lorence 1841 (MO); 12 May 1979 (fr), Lorence 2628 K, MAU, MO, P); (fr), Lorence 2629 (MO); Perrier Sideroxylon wet forest, 630 m, 12 Dec. 1978 (fl), Lo- rence 2122 (MAU, MO). PITON BAMBOU: near crest of hill on the leeward side, 14 Aug. 1968 (fl), Staub sub MAU 13493 (MAU; approaches T. amplifolia). PITON DE LA RIVIÈRE NOIRE: lower montane wet forest, 700— 827 m, 25 Nov. 1978 (fl), Lorence 1964 (MAU, MO); 26 Dec. 1978 (fl), den 2204 (MO). PLAINE pelas by side of road to e Fer, indigenous thicket, 1969 (fr), Guého ee MA U 13750 (MAU); stunted i est around clearing, 600 m, 27 Aug. 197 e et al. 2 , MO). vA OF CASCADE 5 E Calophyllum/Sapotaceae wet forest, 550 m, ay e UT PRECISE . WITHO :( sub Herb. Richard s.n Jan. 1930 (fl, Te i 382 (K, 3 sheets). Tambourissa tau is most closely allied to 7. amplifolia, with. which it has been confused in the past (e.g., Vaughan & Wiehe, 1937, 1941, in their vegetational sampling analyses at Mt. Co- cotte, Macabé Forest and Perrier). Both belong to species Group 5 and have large leaves, making them difficult to separate vegetatively, although ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 they are easily separable by floral characters. Tambourissa amplifolia has pubescent inflores- cences, internally velutinous gynoecious flowers with shorter, non setose styles, and apically con- fluent anther loculi never found in 7. tau (see T. amplifolia for further discussion). The specific epithet tau is Greek, Latin, and Celtic for “T,” referring to the characteristically T-shaped sta- mens Mim are unique in the entire genus. A plant growing in the Curepipe Botanical nesis (Vaughan sub MAU 12203, MAU) is typical of Tambourissa tau in all characters ex- cept the anomalous staminal loculi, which are sessile and not borne on extended lateral arms, although neither are they apically confluent as in T. amplifolia. Furthermore, gynoecious flowers of Vaughan sub U 12203 are typical of T. tau. Flynn and Rowley (1972) studied the ultra- structural development of the pollen wall of this same plant as 7. amplifolia. Flower color morphs with either dark reddish purple or pale pink to greenish white stamens and styles occur; these are generally constant within a given population (e.g., dark red-purple as Perrier, pale pink or white at Mt. Cocotte). The flowers produce no apparent odor. Vernacular name. Bois tambour (Mauriti- us). 27. Tambourissa tetragona (Boiv. ex Tul.) A.DC., Prodr. 16(2): 659. 1868; Baker, Fl. Mauri- tius 289. 1877; Perk. & Gilg, Pflanzenr. 4, 101: 70. 1901. Ambora tetragona Boiv. ex Tul., Ann. Sci. Nat. (Paris) 4(3): 30. 1855; Tul., Monogr. Monim. 302. 1855. TYPE: Ile de France (Mauritius). Without precise lo- cality (ca. 1755) (st), Commerson s.n. (lec- totype, P, here designated). Dioecious (?), sparsely branching slender tree- let or small tree attaining 6 m tall and 10 cm D.B.H., the bark pale brown, smooth with lon- gitudinal fissures, the new growth glabrous, the mature leafy stems quadrangular, 4—6 mm diam., with 4 low wings ca. 1-1.5 mm wide. Leaves opposite, rarely subopposite petiolate, glabrous; petioles stout, 5-15 m m, winged, the wings decurrent onto pe stem; lamina subco- riaceous, narrowly obovate, oblanceolate, nar- rowly oblong or rarely narrowly elliptic, 52-180 mm by 17-48 mm, the apex shortly acuminate, rarely shortly acute, the tip indurated, the base acute, strongly decurrent along the petiole, the 1985] LORENCE-— MONIMIACEAE 143 20°0'S O T. e T. À T, D T. cocottensis cordifolia peltata sieberi . tetragona 10 km FIGURE 40. Distribution map of some Tambourissa species in Mauritius. secondary veins 8—13 pairs, making a 65-80? an- gle with the costa, the venation obscure, visible to 2? on both surfaces, the margin revolute. Cau- liflorous, the androecious inflorescence (fide Tu- lasne) a sparse pleiochasium, the buds globose, inaperturate, shortly pedicellate, the stamens and mature androecious flowers unknown; gynoe cious inflorescence a short, glabrous, ica du pleiochasium of 5-11 flowers, the floral axis quadrangular, 7-13 mm by 2-2.5 mm, subtend- ed by several pairs of glabrous, decussate navic- ulate bracteoles 1 mm by 0.5 mm, the pedicel 3-4 mm by 1.5-2 mm, subtended by a glabrous, deltoid bracteole 1-2.5 mm long. p erase flower in bud napiform-depressed, 8-10 m diam. by 6-8 mm long, apiculate with 4 E glabrous obtuse tepals, the external surface drying wrinkled, glabrous, the walls thick; at anthesis the apex splitting into 4(—5) deltoid lobes ca. 3 mm long and wide, the orifice small, ca. 1 mm wide, X-shaped, the lobes hirsute internally; styles numerous, ca. 200, shortly conical, 0.6—0.8 mm long by 0.5—0.7 mm wide at the strongly venii- cose base, the internal between the styles. Fruiting receptacle and car- pels unknown Distribution. Endemic to Mauritius (Fig. 40). Habitat. Tambourissa tetragona is presently known only from low cloud forest on the summit of Mt. Cocotte at the southern extremity of the central plateau. 144 MAURITIUS. GRAND PORT: woods of the hes Port iei Aug. 1851 (st), Boivin s.n. (P, type). NE COCOTTE: Aug. 1977 (st), Piu 3201 (P): su desit 11 Dec. 1979 (st), Guého & Lecordier sub MAU 19501 (MAU), low, Bowen cloud forest on it, 770 m, 19 Dec. 1978 (st), Lorence & Lecordier 2164 (MO); (st), Lorence & Lecordier 216 , MAU, O); 17 Jan. 1979 (st), Lorence 2286 y 30 Aug 1979 (fl), sd a a 28 y Dec. 1937 (fl), Vaughan Py 1508 (MAU), (fl), Vaughan sub MAU 1 Kos WITHOUT PRECISE LOCALITY: (St), Anon. sub see Richard s.n. (P). The leaves, female receptacle, and styles of Tambourissa tetragona are morphologically similar to those of 7. cocottensis, a species also confined to a small patch of cloud forest on the summit of Mt. Cocotte, one of the island’s high- est and wettest spots. Although androecious flowers and fruiting receptacles of the former are required to further assess its affinity, both belong to species Group 5. The distinctive quadrangu- lar, winged stems of 7. tetragona, however, set it apart from all other members of the genus. In the living state, leaves of Tambourissa te- tragona have a lustrous, dark green lamina and red petiole, the stems being pale green with trans- luscent wings. Vegetatively, it is one of the most attractive members of the genus. Gynoecious uds are shiny green flushed with dark red-pur- ple. Buds are initiated in July and August and appear to mature in December. Known only from a single, small population in a highly degraded habitat, the species should be considered endan- gered. My attempts to propagate it vegetatively by cuttings failed. Vernacular name. Bois tambour (Mauriti- 28. Tambourissa cocottensis Lorence, Bull. Mus. Hist. Nat. (Paris), Ser. 4, Sect. B, Adansonia 3(3): 295, pl. 1, fig. 2. 1982. TYPE: Mauritius. Low cloud forest, summit of Mt. Cocotte, 760 m, 19 Dec. 1978 (fl, fr), Lorence & Le- cordier 2167 (holotype, MO; isotype, MAU). Monoecious treelet or small tree 4-5 m tall and 10-35 cm D.B.H., often multistemmed as coppices, the new growth glabrous, the mature leafy stems terete, 2-4 mm diam., longitudi MEN striate. Leaves opposite to subopposte, petiolate, glabrous; petioles 7-20 mm by 1-2 mm; lamina subcoriaceous, elliptic, 45-105 mm by 15-40 mm, the apex shortly acute to shortly acuminate, the tip indurated, the base acutely cuneate, slight- ly decurrent, the secondary veins 6-9 pairs, mak- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 ing a 55-65? angle with the costa, the venation visible to 3? on both surfaces, slightly raised, the margin moderately revolute. Androecious flow- ers solitary or in fascicles of 2, borne along the trunk. Androecious flower in bud globose, apic- ulate, 6 mm diam., externally corky brown, the pedicel and peduncle 4-5 mm by 1.5 mm, quad- rangular, pilose, bearing 4 decussate rows of bracteoles m long; at anthesis 4-7-fid, the lobes spreading flat, 20-21 mm diam., reddish pink; stamens ca. 150-200, subsessile, linear- subulate, 3 mm by 0.6-0.8 mm, the loculi lateral, free, occupying ca. ?5—4 total length of stamen, the connective prolonged, slender, 1 mm long, the internal receptacle surface glabrous. Gy- noecious inflorescence cauliflorous from meri- stematic swellings along the basal 1 m of the trunk, the flowers solitary or in fascicles of 2. Gynoecious flower in bud napiform-depressed, 9-11 mm diam. by 5-7 mm long, the apex bear- ing 3-4 minute, pubescent deltoid tepals, the ex- ternal receptacle surface pale corky brown with scattered simple hairs, the pedicel and peduncle 4—7 mm by 1-1.5 mm, jointed near the base, quadrangular, pilose, bearing 4 decussate rows of minute, ciliate naviculate bracteoles 1-1.5 mm long; at anthesis the apex splitting into 5-7 ir- regular, + inflexed deltoid lobes, the orifice com- prising ca. '4 total width of the receptacle; styles numerous, ca. 150-200, narrowly conical, setose, 1.5-2 mm long by 0.2-0.3 mm diam slightly ventricose base, the apex long acuminate, the internal receptacle surface with scattered clusters of simple hairs between the styles. Young, submature fruiting receptacle globose-urceolate, 15 mm diam. by 12 mm long, the apex de- pressed, the orifice subcircular, 3-4 mm diam., .5 mm wide basally, pilose between the styles and on the lobes, the peduncle and pedicel 5 mm by 3 mm. Mature fruiting receptacle and carpels unknown. Figure 41. Distribution. Endemic to Mauritius (Fig. 40). abitat. Tambourissa cocottensis is known only from a single population on the summit of Mt. Cocotte in low cloud forest (annual precip- itation 5,000 mm). Extensive searching revealed only four individuals growing within an area ca. 100 m across. 1985] LORENCE- MONIMIACEAE 145 FIGURE 41. Tambourissa cocottensis Lorenc W . a 1 M 1 + Pes fruiting receptacle, lateral view.—F. As for E,a e.—A. Habit. — B. Gynoecious flower in bud, lateral view. —C. D. Gynoecious flower in bud, longitudinal section. — E. Submature pical view. A. Lorence 2288 (MO). B-D. Lorence 2287 (MO). E, F. Lorence 2167 (MO). Bars equal 10 mm in A, E, F, and 1 mm in B-D 146 MAURITIUS. MONTAGNE COCOTTE: summit region, low degraded cloud forest, 760 m, 17 Jan. 1979 (fr), Lo- rence 2287 (K, MO, P); (st), Lorence 2288 (MO); crest, 14 July 1982 (fl), Lecordier sub MAU 20412 (MAU); 23 Sept. 1983 (fl), Lecordier & Strahm sub MAU 20563 Tambourissa cocottensis closely resembles 7. tetragona in a number of respects, including leaf size, shape, texture, margin, and venation, but lacks the winged stems and petioles of the latter. Both are cauliflorous with structurally similar flowers, but those of 7. cocottensis are solitary or in fascicles of two and externally corky, dif- fering from the smooth, glabrous pleiochasial flowers of T. tetragona. They more closely re- semble those of T. tau, except for the distinctive quadrangular floral pedicels bearing four decus- sate rows of ciliate bracteoles. Both T. cocottensis and T. tetragona Rune n to aperies Group 5, al- though tł tne p l fthe latter are suggestive of certain Madagascan species. The four known individuals of Tambourissa cocottensis are of different size classes, suggesting some regeneration has taken place in recent years. The two largest individuals (both gynoecious) were in bud in 79, and young fruits were present in December 1978 and January 1979; androecious flowers were not found. AI- though officially a nature reserve, the forest on Mt. Cocotte is highly degraded and has under- gone extensive invasion by vigorous, weedy ex- otic plant species. As a result, there is a grave danger that the population is too small to main- tain itself, as is also the case for T. tetragona. Both appear to be relictual species confined to the summit of Mt. Cocotte. 29. Tambourissa cordifolia Lorence, Bull. Mus. Hist. Nat. (Paris), Ser. 4, Sect. B, Adansonia 3(3): 296, pl. 1, figs. 5, 6. 1982. TYPE: Mau- ritius. Pétrin Nature Reserve: low Philippia heath formation, 630 m, 15 May 1979 (fl, fr), Lorence 2631 (holotype, MO; isotype, K) Glabrous dioecious shrub, rarely a small tree- let, the multiple erect stems 50-200 cm tall and 6-10 mm diam., the mature leafy stems 2-4 mm diam., longitudinally striate. Leaves opposite, decussate, subsessile or sessile, glabrous; petiole 1-1.5 mm long by 2-3 mm wide; lamina sub- coriaceous, ovate to elliptic, 30-80 by 25-55 mm, the apex shortly acute, obtuse or retuse, often ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 rounded or mucronulate, the base cordate, rarely subtruncate, the dary veins 5-6 pairs, mak- ing a 50-60? angle with the costa, the venation raised and visible to 3(—4?) adaxially and to 4(-5?) abaxially, the margin slightly revolute; seedling leaves heterophyllous, the petioles short, red when living, the lamina narrowly elliptic, entire or with 1-2 pairs of small teeth apically. Inflorescence glabrous, either ramiflorous on leafless nodes or cauliflorous, the flowers solitary or in fascicles of 2(—3), very rarely in a short di- or pleiochasium of 3(—5) flowers, or the flowers rarely solitary and axillary or terminal on leafy shoots. Androecious flower in bud globose, 10-16 mm diam., the apex with 4 minute, obtusely deltoid tepals, the ped- iceland peduncle jointed basally, 9-30(-50) mm by 1-1.5 mm, bearing basally 1-3 pairs of api- cally ciliate, deltoid bag dee at anthesis 4(—5)- fid, 28-36 mm diam., the lobes spreading flat, then reflexing; 6 CuneHs numerous, ca. 300-400 narrowly ellipsoid or clavate, 1-2 mm by 0.5-1 mm, the loculi occupying !^—/ total length of the stamen, usually separate, sometimes confluent basally into a U-shape, or confluent apically at the obcordate apex, the con ecious flower in bud napi- form-depressed, apiculate with 4 minute tepals, the pedicel and peduncle jointed basally, 10-15 mm by 1-1.5 mm, basally bracteolate as in male flower; at anthesis shallowly napiform-urceolate, 8-12 mm diam. by 4-6 mm long, the orifice splitting into 4(—6) nie deltoid lobes, 5-8 mm diam., comprising !^—/ total width of the receptacle, the internal surface of the lobes shal- lowly rugose, glabrous; styles numerous, ca. 100- 400, shortly conical, 0.8-1 mm long by 0.4-0.5 mm wide basally, the apex papillose, the internal receptacle surface with clusters of simple hairs between the styles. Fruiting receptacle solitary on the stems, patelliform to shallowly cupuli- form, 25-60 mm diam. by 13-40 mm long, the orifice 16-40 mm diam., comprising /2—% total width of the receptacle, the rim subentire with 4—6 indurated deltoid lobes, the walls 8-12 mm thick, the external surface corky brown, the in- ternal surface with numerous short, conical styles 0.5-1 mm long by 0.5 mm wide basally, inter- spersed with clusters of hairs, the pedicel and peduncle 16-25 mm by 2-3 mm. Mature carpels 12-13 mm long, 6-8 mm wide, 5-6 mm thick, ovoid-compressed, the endocarp dark brown, the surface scrobiculate. Gametic chromosome number, n = 19. Figure 42. 1985] LORENCE— MONIMIACEAE 147 FIGURE 42 pir aca cordifolia pni — A. Habit, male. — B. Androecious flower at anthesis, apical view. —C. Stamen with free espa oe w.—D. Stamen with free loculi, adaxial view. —E ith basally conflu eu loculi, des ] view.—F. nor e flower in bud, lateral view.—G. ious flower at apie — ak ious flower at a apical view. —I. Gynoecious flower at anthesis, lon- secti a Subma ture fruiting receptacle. — K. Fruiting carpel, lateral view. A—E. y pides 2632 (MO). EL Lorence 2630 (MO). J, K. Lorence 2631 (MO). Bars equal mE mm in À, B, F-K, and 1 mm in C-E 148 Distribution. Endemic to Mauritius (Fig. 40). Habitat. Tambourissa cordifolia is restricted to upland regions of hard, unweathered groun water laterite or “cuirasse” supporting low scrub formations, often of Philippia heath. It is partic- ularly abundant at Pétrin (630 m, annual pre- cipitation 4,000 mm), and also occurs in com- parable Philippia heath on Mt. Laselle (550 m, annual precipitation 4,500 mm), in low, marshy scrub at Les Mares near the base of Mt. Cocotte (650 m, annual precipitation 4,800 mm), at Crown Land Declerc, and in low thicket near the summit of Piton de la Riviére Noire (827 m, annual precipitation 4,200 mm) MAURITIUS. CROWN LAND DECLERC: 1 5 Aug. 1968 (fl), Vaughan sub MAU 13343 (MAU). LES MARES: track from Plaine Paule to Mt. Cocotte, 650 m, 13 May 1979 (fl), Lorence 2626 (MO); 10 Aug. 1979 (fl), Lorence 2894 (MO). MARE LONGUE PLATEAU: June 1977 (fl), Friedmann 3132 (P). MONTAGNE LASELLE: N flank, 4 Oct. 1973 (fl), Lorence sub MAU 16282 (MAU); Mid- lands, exposed situations, Mar. 1931 (fr), Vaughan sub MAU 729 (MAU). PÉTRIN NATURE RESERVE: low in- digenous thicket, 2 June 1971 (fr), Guého sub MAU 14762 (MAU), Phylica/Philippia heath on lateritic soil, Oct. 1975 (fl), Guého sub MAU 17585 (MAU, 2 sheets); indigenous thicket near Pétrin, 2,150 ft. (630 m), 23 Mar. 1963 (fr), — MAU 10337 (MAU); low MO); 12 Dec. 1978 (fr), Lorence 2119 (MAU, MO); 10 Feb. 1979 (fr), Lorence 2378 (MAU, MO, P); 4 Apr. 1979 (fl), Lorence 2571 (MO); 15 Apr. 1979 (fr), Lo- rence 2595 (MAU, MO); 9 May 1979 (fl), Lorence 2618 (MO); 12 May 1979 (fl), Epid 2623 (MO); 15 May 1979 (fr), Lorence 2630 (MO); (fl), Lorence 2632 (MAU, MO); 22 May 1979 (fl), jp 2645 (MO, P, Z); (fr), Lorence 2646 (MO); 19 June 1979 (st), Lorence 2660 (MO); 1 May 1976 (fl), Richardson et al. 4074 (K); (fl), Richardson et al. 4075 (K); = Richardson et al. 4076 (K); (fr), Richardson et al. 4077 (K); C ç Vaughan sub MAU 13776 (MAU). PITON DE LA RIVIËRE NOIRE: (fl), Randabel sub MAU 679 (MAU). WITHOUT PRECISE LOCALITY: (st), right hand leafy collection only, Anon. sub MAU 1314 (MAU). Its sessile, cordate leaves and shrubby habit are presumably adaptations to the unique eco- logical niche which Tambourissa cordifolia oc- cupies. Among other Mascarene species of the genus, it appears to be most closely allied to 7. amplifolia in terms of floral morphology. Both are cauliflorous (rarely ramiflorous) species with long pedicellate, often solitary flowers, the an- droecious flowers being 4-fid, internally white, and having numerous clavate stamens, the gy- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 noecious receptacles being napiform, opening by 4(-6) everted deltoid lobes, and having numer- ous short, conical styles interspersed with hairs. Tambourissa amplifolia is readily distinguish- able by its larger habit, basal cauliflory, monoecy, pubescent inflorescences with longer pedicels and smaller floral receptacles, and very much larger, non-cordate petiolate leaves. Both belong to species Group p he only other dioecious Mauritian species besides pe cordifolia is T. peltata, which has ternate to subalternate leaves and be- longs to species Group 9. Although the two species are known to hybridize naturally, at least one such hybrid (Lorence 2624, MO) is highly sterile with ca. 95% aborted pollen grains. In living material, the flower buds and outer surface of the receptacle are cream or yellowish flushed with purple-red. The inner, staminal sur- face of the androecious flowers and lobes of the white and the styles purple- red. Flowers of both sexes produce strong, fruity odors akin to that of ripe apricots or stewed ripe bananas. Vernacular name. Bois tambour (Mauriti- us). 30. Tambourissa sieberi (Tul.) A.DC., Prodr. 16(2): 6 1868; Baker, Fl. Mauritius 287. 1877; Perk. & Gilg, Pflanzenr. 4, 101: 68. 1901; Vaughan, Mauritius Inst. Bull. 1(1): 76. 1937. Ambora vindi Tul., Ann. Sci. Nat. (Paris) 4(3): 31. . Monogr. Mo- nim. 304. 1855. precise locality (ca. Maurit. II, 316 (holotype, P; isotypes, G, K). A. vestita Tul., Ann. Sci. Nat. (Paris) 4(3): 31. 1855; Tul., Monogr. Monim. 306. 1855. Tambourissa vestita (Tul.) A. DC., Prodr. 16(2): 660. 1868; Cor- d : 69. 1901. TYPE: ““Bourb ably Mauritius), without precise id (ca. 1755), Commerson s.n. (holotype Monoecious tree 10-15 m tall and 30-40 cm D.B.H., the bark light brown, flaking, the new growth velutinous-tomentose with simple, yel- lowish wavy hairs, the mature leafy stems ve- lutinous, terete, 2-5 mm diam. Leaves opposite to subopposite, petiolate; petioles velutinous, 5— 15 mm by 1-1.5 mm; lamina chartaceous to sub- coriaceous, both surfaces mined a when young, adaxially glabrescent, elliptic to 1985] LORENCE—MONIMIACEAE 149 broadly elliptic, rarely obovate or suborbiculate, 40-76 mm by 18-40 mm, the apex shortly acu- minate, acute, rarely obtuse, the base acutely cu- g a 60-70? angle with the costa, the ve- nation apis to 2(-3?) adaxially, and to 3(—4?) abaxially, the margin revolute. Inflorescence cauliflorous on mid to upper part of the trunk and on the major branches, an erect, generally unisexual, rarely leafy thyrse of 16-90 flowers, irregularly branching to 2(—3°), all parts pale yel- owish velutinous-tomentose, the floral axis stout, 50-310 mm by 3-8 mm, flattened and dialated at the nodes, the secondary axes opposite to sub- opposite, + ascendant, 6-90 mm by 2-4 mm, the ultimate branches bearing cymose units of 1—4 flowers, the buds of both sexes globose, 12- 14 mm diam., velutinous, apiculate, the apex crowned with 2-3 pairs of persistent, decussate velutinous naviculate tepals 1.5-2.5 mm long by 1 mm wide, the pedicels and peduncles often subtended by a subulate, velutinous caducous bracteole 2-3 mm long, or rarely by a leaf, the pedicels 10-25 mm by 1.5-2 mm. Androecious flower at anthesis deeply (4—)5(—6)-fid, 25-35 mm diam., the lobes spreading flat; stamens numer- ous, ca. 100-140, ovoid to ellipsoid or oblong, 2-5 mm long by 1—2 mm wide, subsessile, the loculi lateral, separate, occupying almost the en- tire length of the stamen, the filament broad, short or sessile, the connective broad, slightly prolonged or not, the apex obtuse to emarginate the internal receptacle surface glabrous. Gy- noecious flower at anthesis globose, 15-18 mm diam., the orifice subcircular, 2-4 mm diam., with 4—6 irregular, shallow lobes; styles numer- ous, ca. 850-875, narrowly conical to setose, 3— 4 mm long by 0.4-0.5 mm wide basally, coales- cent into groups of 1 2—30, the internal receptacle surface densely velutinous between the styles. Fruiting receptacles produced on the trunk and major branches, in clusters of 3-15, the major axis 15-20 mm diam., woody, the secondary axis ca. 12 mm diam., the pedicel and peduncle 10- 35 mm long by 6-15 mm diam., the receptacle irregularly globose-urceolate, depressed, 55-160 mm diam. -75 mm long, the orifice small, comprising ca. 4—'4 total width of the receptacle, the walls 18-20 mm thick, externally pale corky brown, internally with numerous setose styles 2.5-3.5 mm long by 0.5-0.6 mm wide basally, the sides channelled, the surface sparsely tomen- tose between the styles. Mature carpels ovoid- compressed, 10-13 mm long, 6-7 mm wide, 4— 5 mm thick, the smooth endocarp pale brown. Gametic chromosome number, 7 — 19 Distribution. Endemic to Mauritius (Fig. 40). Habitat. Isolated individuals and small pop- ulations of Tambourissa sieberi are known from mature wet forest at Macabé (660 m) and in the Black River Gorges (400—500 m) where the an- nual precipitation is ca. 3,200-3,400 mm, and also from low cloud forest on Mt. Cocotte (650- 760 m, annual precipitation ca. 5,000 mm). AI- though the species appears to set fruit readily and several seedlings presumably of 7. sieberi were found at Mt. Cocotte, no recent regeneration was Observed at any of the other sites, undoubtedly due to severe competitional pressures exserted by vigorously invasive exotic species. Although Tulasne thought that the sterile type of Ambora vestita may have been collected in Réunion (formerly Bourbon) by Commerson, he in fact questioned the origin of the specimen. Commerson actually did collect in Réunion (Ly- Tio-Fane, 1976), but a number of his other col- lections are wrongly labelled. I therefore suspect that the type of Tambourissa vestita is actually from Mauritius where he collected more exten- sively. That the type of 7. sieberi lacks leaves also seems to have prompted Tulasne to describe cas ome ih subsequently collected or noted from Ré- union (Cordemoy, 1895; Rivals, in herb.; Cadet, pers. comm.). ae the type of 7. vestita is indistinguishable from material of 7. sieberi, I consider the species synonymous and have cho- sen to retain the latter name because the type has flowers and is more widely distributed in herbaria. MAURITIUS. BLACK RIVER GORGES: | Dec. 1933 ria Carcenac sub MAU 794 pia: Crown Land Le B ton in Black River Gorg mary wet forest, ca. 400 m, 10 Jan. 1976 (fl), maskay 1585 (K, 2 sheets; MAU, 2 sheets; MO). MACABÉ FOREST: (fl) 12 Mar. 1933 (fl), Carcenac 48 sub MAU 1313 (MAU): Crown Land Macabé, 20 Jan. 1950 (fr), Dulgeet sub MAU s.n. (MAU); left hand side of path to Macabé plot, 20 Dec. 1965 (fr), Forest Department sub MAU 11935 (carpol. coll. 161) (MAU); — forest of Ma- cabé a F . Dept. sub MAU s.n 6 Jan. 12499 (M 1967 (fl), POUR sub MAU 13094 (MAU); corner of plot near ] July 1968 (fr), Vaughan sub MAU 13229 ies a) 187) (MAU, 2 sheets); upland cli- 150 max dp of (end 24 Nov. 1938 (st), iie sub MAU s.n. (MAU). MONTAGNE COCOTTE: degraded for- est, 2, 200 ft. (760 ey " Nov. 1969 (fr), Barclay ys (K); on crest, 9 Nov. 1969 (fr), Guého sub M. , | Dec. 1978 (f), Lorence 1978 (M 1979 (Hl), Lorence 2250 (K, MAU, MO, P); Lo 2251 (K, MAU, MO, P, REU, Z). WITHOUT PRECISE LOCALITY: 1865 (fl), De Franqueville s.n. sub Herb. DeCandolle 16, 2: 660, no. 12 (G-DC; microfiche, MO); (fl), Anon. sub Herb. Richard s.n. (P); (fr), Vaughan V 35 sub MAU spirit coll. 52 (MAU). Tambourissa sieberi seems to be most closely related to 7. bathiei from Madagascar, particu- larly in terms of floral morphology, which is dis- cussed under the latter species. Both belong to species Grou Mauritian species, 7. sieberi shares several features with 7. quadrifida, including cauliflory, a thyrsic inflorescence struc- ture, closed gynoecious floral receptacles which open by small apical pores, and similar strong, fermenting-fruity floral odors. Tambourissa quadrifida is totally glabrous, however, and ap- pears to be more closely allied to the Madagascan northern Madagascar (Baron 6720, K) with sim- ilarly tomentose leaves, accompanied by a pencil sketch of a solitary, terminal fruit, apparently represents an undescribed species possibly allied to T. sieberi. Besides having solitary, terminal fruits, the leaves of Baron 67 20 differ from those of T. sieberi in containing numerous oil cells in ultimate veins. It is described as T. *sp. D" (no. 43) at the end of this treatment. Flowers of Tambourissa sieberi are either bright yellow or dark purple, often within the same pop- ulation. Floral odor in both sexes is strong and cloying, reminiscent of overripe or fermenting fruit Vernacular name. Bois tambour (Mauriti- 31. Tambourissa bathiei Cavaco, Bull. Soc. Bot. rance 104: l. galava, right hand tributary of the Ikopa River, W of Antsiafabositra (probably July 1899) (fl), Perrier de la Báthie 495-A (ho- lotype, P). Monoecious tree, the new growth hirsute. Leaves (known only from the inflorescence) pre- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 sumably opposite, petiolate; petioles 5-6 mm by 8 mm, hirsutulous; lamina chartaceous, ellip- tic, 40-58 mm by 20-25 mm, the apex shortly acuminate, the base acutely cuneate, the second- ary veins 5-7 pairs, making a 65-70? angle with the costa, the venation prominent, visible to 4? on both surfaces, the margin plane. Inflorescence cauliflorous, unisexual, a condensed pleiocha- sium, or thyrse of (8—)20—34 flowers, sometimes leafy, the floral axis (20-)55-95 mm long by 3- 4 mm wide medially, hirsutulous-pubescent. An- droecious flower in bud globose, 10-13 m diam., sparsely pubescent, apiculate with E minute, obtuse velutinous tepals, the pedicel 25— 45 mm by 1.5-2 mm, subtended by a small, caducous ciliate deltoid bracteole 2 mm long; at anthesis deeply 3-4-fid, 25-35 mm diam., the lobes spreading + flat; stamens numerous, ca. 350—450, clavate, 2-3 mm long by 1-1.5 mm wide, the filament well developed, comprising ca. 2—7} total length of stamen, the loculi apical, connivent or confluent at the obcordate-emar- ginate apex, occupying ca. !^-'^ total length of stamen, the internal receptacle surface with clus- ters of short, dense hairs between the stamens. Submature gynoecious inflorescence a fulvou velutinous-pubescent condensed ipt niit of 8 flowers, the floral axis 24 mm mm. Submature gynoecious flower in bud abos. 7- 8 mm diam., apiculate with 3-4 obtuse, veluti- nous tepals, the pedicel 12-15 mm by 1.5 mm; styles numerous, several hundred, completely lining the receptacle, narrowly conical, 1 mm long .4 mm wide at the slightly ventri- cose base, the acuminate apex slightly papillose, the interna surface densely velutinous between the styles. Mature gynoecious flowers, fruiting receptacle, and carpels unknown. r Distribution. Endemic to Madagascar (Fig. Habitat. Tambourissa bathiei is known only from the type, which was collected along the Iko- pa River, probably in riverine forest, in the Ma- forest of Dalbergia and Commiphora, but is now mainly savannah due to human disturbance. On the basis of floral morphology, Tambou- rissa bathiei appears to be most closely related to T. sieberi, a species endemic to Mauritius, which also belongs to Group 6. Both species have small leaves, densely pubescent new growth, are cauliflorous with yellowish velutinous, generally 1985] LORENCE- MONIMIACEAE 151 unisexual thyrses or pleiochasia, and globose gy- noecious receptacles lined with numerous nar- rowly conical styles. Although mature gynoecious flowers of Tam- bourissa bathiei are unknown, the buds are sim- ilar to those of 7. sieberi, which open by a small, subentire apical pore only 1-2 mm diam. An- droecious flowers of T. bathiei have numerous clavate stamens with long, distinct filaments and loculi connivent or confluent at the obcordate apex. In contrast, stamens of T. sieberi are fe braaa: ly deltoid-ovate, sessile, and h loculi along most of their length. Further collec- tions, particularly mature gynoecious flowers and fruit, of this poorly known species are required. See discussion under species number 42 32. Tambourissa perrieri Drake in Grandidier, Hist. Phys. Madagascar 1(1): 24. 1902; Perk., Pflanzenr. 4, 101 (Nachtr.): 42. 1911; Ca- vaco in Humbert, Fl. Madagascar 80: 30, Be VIII:5-7. 1959. TYPE: Madagascar. Ma- unga: vicinity of Maevatanana, Ampassiri, jr July 1899 (fl), Perrier de la Báthie 495 (holotype, P; isotypes, P, 2 sheets). Monoecious shrub or tree 2-8 m tall, the bark smooth, dark, the branches spreading, the new owth sparsely pilose, glabrescent, the mature leafy stems pale green, smooth, glabrous, 2-2.5 mm diam. Leaves opposite, qiiis glabrous; petioles 5—10(—20) mm by 1-2 mm; lamina char- taceous to Aa id both D sopa slightly lustrous, narrowly elliptic, narrowly ovate to nar- rowly oblong, 75-250(-280) mm by 27—-50(-75) mm, the apex acuminate, the base acute to slight- ly obtuse, the secondary veins 7-13 pairs, mak- ing a 75-85? angle with the costa, the venation very prominent, finely reticulate, visible to 4(—5?) on both surfaces, the margin plane. Inflorescence cauliflorous on meristematic swellings on the trunk, ra be tiis or rarely terminal, a short, unisexual or sexually mixed pleiochasium of 7- 9 flowers, the floral axis glabrous or with scat- tered hairs, 55-125 mm by 2-4 mm, each pedicel subtended by a broadly deltoid, ciliate bracteole. Androecious flower in bud globose, 9-11 mm diam., smooth, glabrous or with rare, scattered hairs, apiculate with 4 small, thick hirsutulous deltoid tepals, the pedicel 10-32 mm by 1-1.5 mm, with scattered hairs; at anthesis deeply 4-fid, 18-25 mm diam., the lobes spreading flat; sta- mens numerous, ca. 150-200, ovoid to obovoid or clavate, 1.8-3 mm long by 1.2-1.5 mm wide, more or less recurved, subsessile or the filament comprising up to !^ total length of the stamen, the loculi confluent at the obtuse apex or sepa- rate, the connective retuse, not prolonged, the internal receptacle surface with scattered to clus- tered short, simple hairs between the stamens. Gynoecious flower in bud obovoid to obpyri- form, 9-11 mm long by 6-7 mm wide, + gla- brous or with a few hairs basally, apiculate with 4—5 small, thick hirsutulous deltoid tepals, the pedicel stout, tapered, 3-15 mm by 1.5-2 mm wide medially; at anthesis obovoid to obpyri- rm, 12-14 mm long by 7-9 mm wide, the apex shallowly 4—5-fid, the lobes thick, erect, deltoid, the orifice 3-6 mm diam., comprising ca. '^ total width ofthe receptacle; styles numerous, ca. 200, crowded, shortly conical, 5-angled, 0.5—0.7 mm long by 0.5 mm diam. at the slightly ventricose base, the apex shortly acuminate, the internal receptacle surface densely velutinous with short, white hairs between the styles and on inner sur- face of the lobes. Fruiting receptacle solitary, borne on the trunk or branches, rarely terminal, obovoid, subglobose or rarely cupuliform, 50- 150 mm long by 35-130 mm wide, externally grayish, corky, smooth or venose, the orifice comprising '45—'4 total width of fruit, the styles conical; pedicel and peduncle 28-45 mm by 6- 10 mm. Mature carpels ovoid-compressed, 8-10 mm long by 7-8 mm wide, the endocarp brown. S Distribution. Endemic to Madagascar (Fig. ). Habitat. Tambourissa perrieri appears to be commonest in the northwestern sector of Mad- agascar (the Diego Suarez, Sambirano and Ma- junga regions) where it apparently occurs In vege- tation types ranging from sclerophyllous moist forest o naceae and dry forest of Dalbergia/Commi- phora/Hildegardia, frequently along rivers, to wet forest of Tambourissa/ Weinmannia. As most collections lack precise data, it is often difficult or impossible to determine the exact habitat. In addition, two collections of what appear to be T. perrieri (but differ in having cupuliform fruits) are known from the lower limits of the Tam- bourissa/ Weinmannia zone (ca. 900 m) of the eastern domain at Analamazaotra near Perinet, a seemingly disjunct distribution pattern possi- i iesidhen to undercollecting. MADAGASCAR. DIEGO SUAREZ: Manongarvio Massif W of Sambirano River, above 510 m, Apr. 1909 (fr), Perrier de la Bathie 10116 (P); Sambirano, Feb. 1923 152 (fr), Perrier de la Báthie 15465 (P). MAJUNGA: Bema- 300 ae Aug. 1907 (fl), Perrier i osy, near Maormand- la, Mar. 1909 (fr), Perrier de " ped 10117 (P). TA- MATAVE: near Analamazaotra, ed 1912 (st), Perrier de la Bathie 10128 (P); phas saba, near Analama- zaotra, 1,000 m, Feb. 1921 (f. "Perrier de la Báthie 15958 (P). Morphologically, Tambourissa perrieri most closely approaches 7. quadrifida from Mauritius, the only other member of Group 7. Both species are cauliflorous, rarely ramiflorous or terminal, with unisexual or sexually mixed pleiochasia or d produced along most of the length of the runk. Androecious flowers of both split deeply is four flat or recurved segments bearing nu- merous short stamens. The gynoecious flowers of both are obovoid to obpyriform or ellipsoid with a small orifice; it is much smaller (1-2 mm diam.) in 7. quadrifida which also has thicker, obtuse or retuse leaves with more obscure ve- nation. Both species occur in relatively dry hab- itats and form a group apparently not closely allied to other members of the genus, with the possible exception of Group 6. Vernacular name. Ambora-lavaravina (Mad- agascar). 33. Tambourissa quadrifida Sonn., Voy. Ind. ra ental ed. 1, 2: 237, tab. 134. 1782; ed. 2, 4: 405, tab. 134. 1806; Gmel., Syst. Nat. 2(1): 16. 1791; A.DC., Prodr. 16(2): 658. 1868; Baker, Fl. Mauritius 288. 1877; Perk. & Gilg, Pflanzenr. 4, 101: 71. 1901, excl. synon. T. obovata; Drake in Grandidier, Hist Phys. Madagascar 1(1): 21. 1902, pro parte, 1911. Mithridatea quadrifida (Sonn.) Comm. x Schreb., Gen. Pl. 2: 783. 1791; Willd., Ss Pl. 1(1): 27. 1797; Spreng., Syst. 5(3): 866. 1826. Ambora quadrifida (Sonn.) Poir., Encycl. 7: 565. 1806; Tabl. Encycl. 784. 1806; Tul, Monogr. Monim. 297. 1855. TYPE: Ile de France (Mauritius). Without precise locality (ca. 1775) (fl), Commerson s.n. sub Herb. A. L. de Jussieu 16708 [lec- totype, P-JUSS, here designated (photo, MO); isolectotypes, P-JUSS (photos, MO), both mixed with 7. elliptica subsp. micran- tha] Ambora sempervirens F. G. Dietr., Lexic. Gart. 1: 342. 2, nom. superfl., based on T. quadrifida. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 A. godes Lam. ex Steud., Nom. Bot. 37. 1821, superfl., non A. tamburissa Boiv. ex Tul. Mithradatea Jamburissa Bojer, Hortus iri 290. . superfl., based on T. quadri Ambora ape Tul., Monogr. Monim. 432. “1855. Tambourissa neglecta (Tul.) A.DC., Prodr. x Y. 659. 1868; Perk. & Gilg, Pflanzenr. 4, 101: 1901. rYPE: Mauritius. July 1855, add s.n. (lectotype, P, here designated). Monoecious to subdioecious tree attaining 15 m tall, the stems often multiple, to 15-30 cm D.B.H., the bark light brownish gray, flaking or checking, the new growth totally glabrous, the mature leafy stems glaucous, smooth to longi- tudinally striate, 2-4 mm diam. (suckers to 7-8 mm diam.) Leaves opposite to subopposite (sometimes alternate on suckers), petiolate, gla- brous; petioles 5-12 mm by 1-1.5 mm; lamina subcoriaceous, broadly to narrowly elliptic, ob- long or obovate, (30-)45-100 mm by (13-)17- 52 mm, the apex shortly acute to mucronate, obtuse, rounded or retuse, the base acutely de- current to acutely cuneate, the secondary veins 4—8 pairs, making a 55-65? angle with the costa, the venation + obscure adaxially, visible to 2- 3? on both surfaces, the margin plane to slightly revolute. Inflorescence a glabrous, generally + unisexual, sometimes leafy, condensed pleio- chasium or sparsely branching thyrse of (3-) 5-35(-110) flowers, cauliflorous on meristematic swellings along the entire length of the trunk, or the flowers rarely solitary on leafless nodes or axillary, the floral axis (30-)40-180 mm by 2-4 mm, the flowers borne singly or in cymose units of 2-3. Androecious flower in bud smooth, obo- void to ellipsoid, 10-20 mm long by 7-10 mm wide, apiculate with 3—4 minute, glabrous, ob- tuse-deltoid tepals, the pedicel 10-30 mm by 1- mm, occasionally jointed to a distinct pe- duncle up to 12 mm long, often subtended by an obtuse, ciliate bracteole 0.5 mm long by 0.5-1 mm wide; at anthesis deeply (3-)4-fid, splitting deeply to the base, 20-32 mm diam., the lobes ultimately completely reflexed; stamens numer- ous, ca. 300—500, ellipsoid to oblong, 1.5-2.5 mm long by 0.8-1.4 mm wide, the filament dis- tinct, comprising ca. '4—!^ total length of the sta- men, n ones eee to slightly apie ulate, t clusters of short hairs between the stamens. Gy- noecious flower in bud obovoid to ellipsoid, 10- 16 mm long by 7-10 mm wide, smooth, gla- brous, apiculate with 3-4 minute deltoid tepals, 1985] LORENCE— MONIMIACEAE 153 the pedicel 7-10 mm by 1.5-2 mm, sometimes jointed to a peduncle attaining 10 mm long, often subtended by a minute! le; at anthesis with a subentire, shallowly 3-5-fid orifice 1-2 mm diam., the thick deltoid lobes straight or scarcely outcurved; styles numerous, ca. 200—300, lining the receptacle except near the apex, i. con- ical, 0.4 . at the abruptly swollen base, the internal paso surface velutinous between the styles and on the lobes, the dense hairs short, the external surface soon becoming corky. Fruiting receptacles borne on the trunk and major branches, solitary or in clusters of 2-3, irregularly obovoid-pyriform to globose-urceolate and depressed, 50-170 mm long by 50-100 mm diam., the walls 16-25 mm thick, externally corky brown, the orifice small, 5-10 mm diam., comprising ca. o total width of fruit, the internal surface with numerous, scat- tered shortly conical styles 0.7—0.8 mm long by 0.3-0.4 mm wide basally, the pedicel and pe- duncle stout, 10-20 mm long by 7-18 mm diam. medially. Mature carpels ovoid- compressed, 13- m nutely scrobiculate-rugose. Gametic chromo- some number, n = Distribution. Endemic to Mauritius (Fig. 38). Habitat. Tambourissa quadrifida is restrict- ed to the western escarpments of the central pla- teau, ranging from the Trois Mamelles mountain range in the north to the mountains opposite Le Morne Brabant in the south. It is local and oc- casional to common in evergreen or semidecid- uous moist forest of Elaeodendron orientale Jacq., Diospyros spp., Mimusops petiolaris Du- bard, and other species characteristic of the rel- atively dry rain shadow area. Although older trees of T. quadrifida are still relatively common, I observed no recent regeneration of the species, possibly due to grazing pressures from the high populations of stag which are maintained in the region for hunting. MAURITIUS. CHAMAREL: Chamarel Hill, near first hairpin ben 600 ft. (ca. 200 m), 19 Apr. 1939 e d V 1512 (MAU, 5 sheets); Cham (st), Vaughan Vaughan 544 (K). YEMEN MATALA VALLEY: under Brise Fer Mountain, dry forest (st), Bernardi 14846 (G); 4 Dec. 1973 (fl), Coode et al. 4270 (K, 2 sheets; MAU); W foothills of Brise Fer Mt., Yemen, Matala Valley, 17 Jan. 1976 (fr), Lalouette sub MAU 17744 (MAU); 17 Mar. 1979 (fl, fr), Lorence & Lalouette 2538 (K, MO, P, REU, Z); (fl), Lorence & Lalouette 2539 (MO); 31 May 1979 (fl), Lorence 2654 (K, MO, P, Z); (fl), Lorence 2655 (K, MO, P); (fl), Lorence 2656 (MO, REU); Yemen estate, 4 July l 178) (MAU, 4 sheets). WITHOUT PRECISE LOCALITY: (St), Anon. 4663 (G, K); (fl), Anon. sub Herb. Delessert s.n. Sa (fl), Anon. sub Herb. DeCandolle 16(2): 658 (G-DC, ; (st), Anon. sub Herb. Rich- 181 s.n. (K); (fl), Ilardwicke 6 (G); (st), Martin s.n. (G); (fl), Martin 607 (G, 2 sheets). Tambourissa quadrifida appears to be most closely allied to 7. perrieri from Madagascar, also of species Group 7. The latter is distinguishable by its globose androecious flower buds, gynoe- cious flowers with larger orifices (3-4 mm diam.), longer, acuminate leaves with more prominent venation, and floral ground tissue with abundant oil cells lacking in 7. quadrifida. Living flowers produce a strong odor of ripe fruit similar to that of Tambourissa sieberi. Two floral color morphs occur in the population at Yemen, the androecious flowers being either to- mato red or salmon orange, whereas the gynoe- cious flowers have either red or greenish white to pink styles. Buds and outer receptacle surfaces are marbled green, often flushed with red Vernacular names. Bois tambour, Pomme de singe (Mauritius). 34. Tambourissa alaticarpa Lorence, nom. nov hanerogonocarpus perrieri Cavaco, Bull. Soc. Bot. France 104: 613. 1957; Cavaco in Humbert, Fl. Madagascar 80: 4. 1959 non Tambourissa perrieri Drake in Grandidier, Hist. Phys. Madagascar 1(1): 24. 1902. TvPE: Madagascar. Tamatave: E coast, bank ofthe Fandrarazana River, 200 m, 1930 (fl), Per- rier de la Báthie 10102 (holotype, P). Monoecious treelet, the new growth pale yel- ibn: villous, the hairs simple, the submature s 3 mm diam., stramineate, villous. Leaves s petiolate; petioles 15-16 mm by 2 mm, villous; lamina chartaceous, elliptic to oblong, 210-220 mm by 75-80 mm, the apex abruptly acuminate, the base cuneate, adaxially sparsely ay 154 pilose, abaxially densely pilose, the costa velu- tinous, depressed adaxially, prominent abaxial- ly, the secondary veins ca. 7 pairs, making a 50- 60° angle with the costa, festooned brochidod- romous, the venation raised and visible to 3—4? on both surfaces, the margin serrate-dentate with 9—10 pairs of narrowly acute, antrorse teeth ca. ] mm long, the tips glandular, indurated. Ap- parently cauliflorous (also ramiflorous?), the in- florescence either a sexually mixed pleiochasium 150-250 mm long with 7—10 lateral androecious flowers and a single, terminal gynoecious flower, the floral axis 100-240 mm by 1.5-3 mm, pilose to velutinous, or a short, unisexual pleiochasium 20-30 mm long of 7-10 androecious flowers, the floral axis 6-20 mm by 0.5-1 mm, pilose to vil- lous. Androecious flower in bud glabrous or with rare, scattered hairs, obovoid to ellipsoid, 4—5 mm —3.5 mm, apiculate, the apex with 3-4 acute, scarious deltoid tepals 0.5 mm long and wide, the pedicel 13-19 mm by 0.5 mm, sparsely pilose, subtended by a subulate-linear bracteole 2-3 mm long; at anthesis deeply 4-fid, 7-9 mm diam., the lobes spreading flat; stamens ca. 38, clavate, 1.2-1.6 mm long by 0.7—0.8 mm wide, the filament distinct, stout, the loculi separate, lateral or oblique, occupying !⁄4—!⁄ total length of the stamen, the connective obtuse or apiculate, scarcely or not prolonged, the internal receptacle surface glabrous. Gynoecious flower terminal, sparsely pilose or glabrate, in bud narrowly ob- ovoid, tubular, 22-28 mm by 4-5 mm, angular with 4—6 longitudinal wings 1-1.5 mm wide run- ning from apex to pedicel, the apex crowned by 6-8 subulate tepals 1-1.5 mm long, the wall ca. 0.5 mm thick, the pedicel scarcely differentiated, 20-25 mm by 1.5 mm diam. medially; at an- thesis apparently opening by a small (ca. 1 mm diam.) apical orifice flanked by several deltoid lobes; styles numerous, ca. 250-300, sharply conical, | mm long by 0.5 mm diam. basally, acuminate, curved antrorsely, the internal apical portion of the receptacle velutinous with short, simple white hairs, the styles interspersed with short, dense hairs. Fruiting receptacle probably resembling that of T. /ongicarpa (i.e., cylindrical, narrowly winged), the fruiting carpels (fide Per- rier de la Bathie) embedded in the orange recep- tacular tissue, the mesocarp red, the endocarp black Distribution. Endemic to Madagascar (Fig. ). Habitat. Presumably from lowland wet for- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 est of Myristicaceae/Anthostema, the species is known only from the type. Along with Tambourissa longicarpa, T. ala- ticarpa comprises the very natural species Group 8, characterized by elongate, narrowly winged gynoecious flowers and fruits. It differs from the former species in having larger, more sparsely pubescent leaves with smaller, more numerous teeth (only 2-3 pairs in T. /ongicarpa), and ex- ternally sparsely pilose to glabrate buds and flow- 35. Tambourissa longicarpa Lorence, nom. nov. Phanerogonocarpus capuronii Cavaco, Bull. Soc. Bot. France 104: 612, figs. 1-4. 1957; Cavaco in Humbert, Fl. Madagascar 80: 3, fig. I:1—6. 1959 non Tambourissa capuronii Cavaco, Bull. Mus. Hist. Nat. (Paris), Ser. 2, 29: 287. 1957. TvPE: Madagascar. Diego Suarez: Atondradama Pass, N of the Ma- soala Peninsula, E forest, ca. 500 m, 22 Dec. 53 (fl, fr), Capuron 8775-SF (holotype, P). Large monoecious treelet, the new growth pale yellowish villous, the hairs simple, the mature leafy stems fulvous velutinous, 2-3 mm diam. Leaves opposite, petiolate; petiole 10-13 mm by 1.5-2 mm, fulvous velutinous; lamina broadly elliptic, 75-105 mm by 43-65 mm, chartaceous, the apex deltoid, acute, the tip thickened, the base cuneate, the adaxial surface sparsely pilose, especially along the costa and veins, glabrescent, the abaxial surface densely pilose with pale yel- lowish white hairs, especially along the costa and veins, the costa depressed adaxially, prominent and rounded abaxially, the secondary veins 5—6 pairs, making a 45-55? angle with the costa, the basal pairs festooned brochidodromous, the api- abaxially, the margin slightly revolute, strongly dentate in the apical s with (1-)2-3 pairs of broadly deltoid teeth 3-5 mm long, the tips thickened, glandular. Inflorescence basally cau- liflorous, produced on meristematic swellings from the trunk, either a sexually mixed, 15-22 flowered pleiochasium with 1-3 terminal gy- noecious flowers, the androecious flowers lateral in opposite to subopposite pairs or cymose groups of 2-3, the floral axis velutinous, 70-95 mm by 1-1.8 mm, subtended by several deltoid brac- teoles, or a smaller unisexual pleiochasium of 7— LORENCE—MONIMIACEAE 155 1985] 9 androecious flowers, the floral axis 18-35 mm by 1 mm. Androecious flower in bud hirsute, globose, 4-5 mm diam., apiculate with 1(-2) pairs of minute, subulate- “deltoid tepals 0.3-0.5 mm long, the pedicel 10-22 mm by 0.5 mm, pilose, subtended by a subulate-naviculate, pilose brac- teole 1 mm long; at anthesis deeply 4-fid, 9-11 mm diam., the lobes spreading flat, ultimately reflexing, each bearing 2-3 minute, fleshy tepals apically within; stamens 40-50, subulate, 1.5- 2.5 mm long by 0.8-1 mm wide, the filament distinct, thick, the loculi separate, lateral or abax- ially + unifacial, 1-1.5 mm long, occupying ca. V; total length of stamen, the connective pro- longed, acute to apiculate, the inner receptacle surface with scattered hairs between the stamens. Gynoecious flower in bud narrowly obconical, tubular, 30-35 mm by 3.5-4 mm, + quadran- gular with 4(—5) lateral longitudinal wings ca. 1 mm wide, fulvous hirsute-velutinous, the apex bearing 4—6 deltoid-subulate hirsute tepals 1—2 mm long, the receptacle wall ca. 1 mm thick, the pedicel tapered, scarcely differentiated from the receptacle, 20-25 mm by 1 mm, hirsute, sub- tended by 2 subulate bracteoles 2-3 mm long; at anthesis opening by 4 deltoid lobes, the orifice basally, the base slightly ventricose, the internal receptacle surface apically velutinous, the cavity ca. 2-2.5 mm diam., the styles interspersed with numerous short hairs, a mucilaginous exudate present. Fruiting receptacle pendulous, cylindri- cal-ellipsoid, 160 mm by 70 mm, externally pale corky brown, bearing 4—5 longitudinal wings, 4— 5 mm wide, the orifice narrow, at maturity split- ting lengthwise and exposing the central tube bearing part of the carpels, the pedicel and pe- duncle ca. 120 mm by 6 mm. Mature carpels ovoid-compressed, 12 mm by 5-6 mm, the me- socarp thin, red-orange, fleshy, the endocarp brown, minutely foveolate. Distribution. Endemic to Madagascar (Fig. Habitat. The type and only known collection is from the eastern lowland Myristicaceae/A7z- thostema wet forest zone. Most closely allied to Tambourissa alaticarpa, the only other member of species Group 8, T. longicarpa differs by its smaller leaves with few- er, much larger teeth, its more densely villous inflorescence and flowers, and by its stamens with the connective prolonged beyond the anther. Further collections and field studies are greatly desired. 36. Tambourissa castri-delphinii Cavaco, Bull. Soc. Bot. France 104: 283. 1957; Cavaco in Humbert, Fl. Madagascar 80: 32, fig. IX:9- 10 (non fig. VIII). 1959. rype: Madagascar. Tulear: Ebakika, Fort Dauphin district, in secondary forest (savoka), small tree, 17 Nov. 1932 (fl), Decary 11052 (holotype, P). Small monoecious or subdioecious trees 5-7 m tall, the mature leafy stems terete to subtri- gonous, 3-7 mm diam., glabrous, stramineate. Leaves ternate, rarely i ani patil ate, gla- brous; petioles 7-13 m m; in: lamin na subcoriaceous, both ae diris lus- trous, ovate to oblong, 88-170 mm by 38-68 mm, the apex abruptly and shortly acuminate, the base acutely cuneate, the secondary veins 5- 7 pairs, making a 45-50? angle with the costa, the venation raised and prominent, visible to 4? on both surfaces, the margin thickened, shghuy revolute. ; axillary on the leafless nodes, a contracted pleio- chasium or fascicle of 4-9 flowers, the floral axis 1-9 mm by 1.5-2 mm, sparsely puberulent ba- sally, subtended by several pale, naviculate-del- toid, ciliate bracteoles 0.5-1.5 mm long. An- droecious flower in bud glabrous, globose, 9-18 mm diam., apiculate with 4—6 short, obtuse te- pals, the pedicel 10-18 mm by 0.8-1 mm, sub- tended by a bracteole; at anthesis deeply 4—5-fid, 25-32 mm diam., the lobes spreading flat, re- flexing, purple within; stamens numerous, ca. 200-300, ovoid to deltoid, thin, 2.5-4 mm long by 1.5-2 mm wide, the filament short, subsessile, the loculi lateral, confluent at the obtuse apex, rarely free, the connective not prolonged, the in- ternal receptacle surface with scattered or clus- tered one hairs between the stamens. Gy- bly ramiflorous like the male, at anthos napiform-depressed, 4-6 mm long by 10-14 mm diam., externally glabrous, internally densely hirtellous on the lobes, the ori- fice small, X-shaped, the styles numerous, ca. 800-900, conical acuminate, 0.7-0.9 mm long and wide basally, interspersed with numerous hairs; pedicel and peduncle 8 or more mm long. Mature fruiting receptacle oo eade -cupuli- form, 40-45 mm long by 40-60 mm wide, ex- ternally corky, the walls 10-12 mm bu the orifice comprising ca. !⁄4—1⁄ total width of fruit, 156 the styles conical 0.8-1.5 mm by 0.8 mm, in- terspersed with rare hairs; pedicel and peduncle 17 mm by 5 mm. Mature carpels ovoid-com- pressed, 10-11 mm by 5-8 mm, pale tan. Distribution. Endemic to Madagascar (Fig. 0). Habitat. Tambourissa castri-delphinii is known only from the SE coastal region between Mananjary and Fort Dauphin. Capuron notes that the species occurs in coastal “savoka” (sec- ondary or disturbed primary forest) over old sand dunes. njary, Mar.-Apr. 1909 cm rid 8287 (P). TULEAR: ae. Fort Dau- phin and Manantenina, near Manambato (fl), Capuron 28667-SF (P); " Ebakika, 18 Nov. 1932 (fl), Decary s.n. (P); Mendena, 16 Feb. 1949 (fr), Capuron 378-SF (P); Dauphin, Manambato, 12 Dec. 1972 (fl), Jac- quenim 1197 (P). Cavaco (1957b) states that Tambourissa cas- tri-delphinii is related to T. religiosa, also Mad- agascar, because leaves of both species are sim- ilarly subcoriaceous or coriaceous with prominent venation. This similarity is most likely because both speçies occupy a similar habitata in ponsi forest. N tly, adult pł ferent, being opposte in T. religiosa and usually ternate in T. castri-delphinii, the latter character otherwise found only in T. ficus and T. peltata, both Mauritian endemics. In addition, venation of all three ternate-leaved species is prominent and finely reticulate, showing a high degree of organization. Furthermore, androecious flowers of T. religiosa and T. castri-delphinii are totally different. Both T. peltata and T. castri-delphinii and globular with fewer, much smaller stamens. For these reasons T. peltata and T. castri-del- phinii appear to be more closely related and I have placed them together in species Group 9. Tambourissa peltata differs from T. castri-del- phinii by its larger, solitary or paired androecious flowers with thicker pedicels, and discoid female receptacle with broad, sterile lobes and columnar styles. Gynoecious flowers of T. castri-delphinii differ from both T. ficus and T. peltata in having an X-shaped orifice. 37. Tambourissa ficus (Tul.) A.DC., Prodr. 16(2): ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 659. 1868; Perk. & Gilg, Pflanzenr. 4, 101: 69, fig. 18Q. 1901; Perk., Gatt. Monim. fig. 34Q. 1925. Ambora ficus ps Ann. Sci. Nat. (Paris) 4(3): 30. 1855; Tul., Monogr. Monim. 300. 1855. TYPE: “Java” ‘(orobably Mauritius). Without precise locality (ca. 1800) (fl), Leschenault de la Tour s.n. (ho- lotype, P). Ambora v iron Tul., Ann. Sci. Nat. (Paris) 4(3): 31. 1855; Tul, Monogr. Monim. 305. 1855. Tambourissa alternifolia (Tul) A.DC., Prodr. 16(2): 660. 1868; Baill., Adansonia, Ser. 1, 9: 131. 1868-1870; Perk. & Gilg, Pflanzenr. 4, 101: 72. 1901; Cavaco, Bull. Soc. Bot. France 104: 284. 1957. TYPE: Ile Maurice (Mauritius). At the foot of the Pouce, above the plain at Moka, Oct. 1849, Boivin s.n. [lectotype, P, here designated, as to androecious floral buds only, excluding the stem and leaves which are Homalium integrifolium (Lam.) Baill. ap waq sua: Ambora obovata Tul., Ann. Sci. Nat. (Paris) 4(3): 31. 55; Tul., Monast Moat: 306. Tam- bourissa obovata (Tul.) A.DC., Prodr. 16(2): 660. 1868; Perk. & Gilg, Pflanzenr. 4, 101: 70. 1901. TYPE: Ile Maurice (Mauritius). Woods of the Pouce, Sept. 1849, Boivin s.n. (holotype, P). T. elliptica sensu Baker, Fl. Mauritius 288. 1877; sensu , Pflanzenr. 4, 101 (Nachtr.): 44. 1911 non .D Ficus sapotoides Baker, Fl. Mauritius 284. 1877. TYPE: s. Pouce above ep Sept. 1862, Ay- ers s.n. Gia K; photo, NÁ = LA: treelet or tree 10— 15 m tall and 10- 15 cm D.B.H., the base of the trunk frequently swollen, the new growth finely pilose to fulvous tomentose, the mature leafy stems glabrous, stramineate, subterete, 2-5 mm diam. Leaves ternate to subalternate, rarely op- posite to subopposite, petiolate, glabrous; peti- oles 10-25 mm by 1.5-2 mm (to 45 mm long on vigorous suckers); lamina chartaceous to sub- coriaceous, broadly to narrowly obovate, oblan- ceolate, narrowly to broadly elliptic, 60-150 (2180) mm by (20-)30-75 mm (to 170 mm by 100 mm on vigorous suckers), the apex acumi- nate, acute, obtuse or retuse, the base acutely de- current to acutely cuneate, the secondary veins 5-8(-11) pairs, making a 50-70? angle with the costa, the venation finely reticulate, prominent, visible to 3-4? adaxially and to 4? abaxially, the margin plane to slightly revolute; juvenile and sucker leaves often heterophyllous, the petioles red when living, the lamina apically serrate-den- tate with 1-3 pairs of teeth. Inflorescence cau- liflorous on the swollen base of the trunk, some- times extending up the lower portion ofthe trunk for several meters or rarely on the leafless parts 1985] LORENCE— MONIMIACEAE 157 of the lower branches, the flowers solitary or in fascicles of 2(—3) on the meristematic swellings. Androecious flower in bud globose-depressed to napiform-depressed, 20-30 mm diam. by 15-25 mm long, pubescent, smooth to corky, longitu- dinally ribbed basally, the walls thick and fleshy, apiculate with 3—5 minute, hirsute tepals, the pedicel stout, tapered, Jointed near the base, the pedicel and peduncle 15-25 mm by 3-4.5 mm diam. medially, pubescent to velutinous, bearing several hirsute, subulate-naviculate bracteoles 1— 2 mm long; at anthesis shallowly to deeply 4— 6(—7)-fid, either partially closed and urceolate, 20-40 mm diam. by 17-30 mm long, orthe lobes id iin flat open, 35-75 mm diam.; stamens very numerous, ca. 900-1,800, curvate, linear to ellipsoid or ligulate, 3-6 mm long by 0.5-1 mm e, the filament basally thickened, comprising ut total length of the stamen, the loculi sep- arate, parallel, lateral, or abaxially + unilateral and extrorse, comprising ca. '4 total length of the stamen, the connective thick, apically prolonged, obtuse to apiculate or acute, comprising ca. !⁄4— / total length of the stamen, the internal recep- tacle surface bros cu of simple hairs be- tween the stam oecious flower in bud napiform- Pee oe fleshy, the size and pubescence as in the androecious bud; at anthesis urceolate-napiform, 30-40 mm diam. by 15-30 mm long, externally corky brown, sparsely pu- berulent, the orifice ca. 15-25 mm diam., com- prising ca. 7^ total width of the receptacle, the 6— 8 thick, deltoid lobes incurved; styles very nu- merous, ca. 1,000-2,000, narrowly conical or se- tose, 1.5-5 mm long by 0.3—0.5 mm wide basally, the sides longitudinally furrowed, the internal receptacle surface densely velutinous between the styles. Fruiting receptacle solitary at the base of the trunk, irregularly cupuliform, often + com- pressed dorsally, 40-85 mm long, 135-175 mm wide, the orifice subentire, 90-130 mm wide, comprising ca. 7^—/A total width of the receptacle, the walls 10-16 mm thick, both surfaces corky brown, internally with numerous, setose styles 1.5-4.5 mm long by 0.5-0.8 mm wide basally, the pedicel and peduncle 20-65 mm long by 9- l m diam. medially. Mature carpels ovoid- compressed, 10-14 mm long, 5-7 mm wide, 4- 6 mm thick, the endocarp dark brown, the sur- face scrobiculate. Distribution. Endemic to Mauritius (Fig. 38). Habitat. Tambourissa ficus is fairly wide- spread in mid to upper elevation wet and cloud forest fi ti (ca. 600—800 m), often forming localized populati th its of moun- tains or along rivers and lakes. MAURITIUS. BAMBOU MOUNTAIN: top of Bambou Mt., 7 Oct. 1962 (fl), Mulnier sub MAU s.n. (MAU). BASSIN BLANC: forest inside Bassin Blanc Crater, 4 Nov. 1967 (fl), Guého sub MAU 13228 (MAU); lower montane rain forest inside crater, 550 m, 21 Dec. 1978 (fr), Lorence 2188 (K, MAU); be Lorence 2189 (MO); (fl, fr), Lorence 2190 (MO); ca. 2 m from outside of lake, S side, 500 m, 25 May 1976 (fr), Richardson et al. 4164 (K, MAU, MO); indigenous forest, 13 Mar. 1971 (fr), Staub sub MAU 19283 (MAU). BRISE FER: 200- 300 yards before summit of Piton Brise Fer, 29 Dec 1966 (fl), Guého sub MAU 12505 (MAU, 2 sheets); (A), Guého sub MAU 12509 (MAU); Brise Fer Mt., 609 m J s 1960 (fr), Lalouette sub MAU 10020 (carpol. coll. 11) (MAU, 2 sheets); 20 Aug. 1966 (fr), Lalouette sub po 12194 (carpol. coll. 158) (MAU); mature ever- Pen forest transitional to dry forest, 600 m, 8 Dec. 78 (fl), Lorence 2114 (MO); (fl), Lorence 2115 (MAU, es (st), Lorence 2116 (MO); (st), Lorence 2117 (MO); 11430 (MAU); (f), Vaughan sub MAU 12505 (spirit coll. 167) (MAU). CROWN LAND DANGUET: near Port Louis, 4 Sept. 1947 (st), etd V 3177 (MAU). MACABÉ FOREST: ravines of Black River Gorges near Macabé, 28 Nov. 1950 (fl), Duljeet FD 105 (spirit coll. 41) (MAU, 2 sheets); Macabé forest, near kiosk, tran- sitional wet forest, ca. 660 m, 16 Dec. 1978 (st), Lo- rence 2155 (MAU); Ses Lorence 2156 (MO); Plateau Colophane below Mac O m, 15 Jan. 1979 (st), Lorence 2952 (MO i C near screen, 10 Dec. 1938 (st), Vaughan sub MAU 1336A (MAU); Macabé forest transect, 11 Aug. 1937 (st), Vaughan 27 sub MAU 2359 (MAU); Macabé climax forest near large “natte” tree in plot, 16 Nov. 1967 (fl), Vaughan sub MAU 13055 (MAU). wr. COCOTTE: degraded forest, 2,200 ft. (ca. 760 m), 8 Nov. 1969 (fl), Barclay 1692 (K); dense indigenous thicket along crest, 8 Nov. 1969 (fl), Guého sub MAU 141 a D a 7: forest with Nuxia, invaded by exotics, N slop 0 m, 19 Dec. 1978 e Lorence & Lecordier 2162 K, MO); near summit, 0 m, 19 Dec. 1978 (fl), Lorence & Lecordier 2163 (MO, P); forest i St. Marie near Mt. Cocotte, high forest, Nov. 1937 (fl), Wiehe sub MAU 1511 (spirit coll. 45) wet forest invaded by exotics, 630 m, 22 Jan. Lorence 2303 (MO). MT LAGRAVE: low forest on slopes, much invaded, 5 Apr. dd (fr), ii gan MAU 17598 (K, MAU). PIETER BOTH S flank below “La Fenétre," pr de udi de PES, ca. 550 m, 29 Sept. 1978 (st), Lorence 1833 (MO). PITON DU MILIEU: dense indigenous thicket, rocky ridge at foot of the mountain, 25 Oct. 1969 (fl), Guého sub MAU 14181 (MAU); low native thicket halfway up steep summit, 24 Nov. 1971 (fl), Guého sub MAU 14983 (MAU); degraded lower montane wet forest, 650-750 m, 29 Dec. 1978 (st), Me ud O); (st), Lorence 2236 (MO). PITON DU FOUG . 500 m, 27 Apr. 1976 (st), Richardson i 404 2(K ). POUCE MT.: shoulder of Le Pouce, Dec. 1860 (fl), Ayres s.n. (K); 158 shoulder of Le Pouce, Nov. 1862 (fl), Ayres s.n. (K); , Calophyllum wet te Lorence 2193 (MO). ), Vaughan sub MAU VUILLEMAN FOREST: near Midlands (fl), mon in forests of medium altitude (st), Vaughan iu MAU 1513 (MAU). Although one of the most polymorphic Mau- ritian species, Tambourissa ficus is nevertheless distinct from all other Mascarene species, and is the only one with flowers and fruit produced on the swollen base of the trunk. It is potentially monoecious [e.g., Wiehe 1511 (MAU), (t MAU 12505 (MAU), Lorence 2162 (M MO but frequently produces flowers of a in sex. Although only four to eight flowers are pro- duced during a single season per plant, this pau- city has been compensated for by a floral gigan- tism unrivalled in the genus and indeed the entire covered by ca. 900-1,800 stamens, whereas gy- noecious flowers are large, urceolate cups 30—40 mm diam. lined with a comparable number of styles. The cupuliform fruits, which attain 175 mm diam., are also some of the largest in the genus Both foliar and floral morphology are variable in Tambourissa ficus, but are relatively constant within a given population. For example, popu- lations from the central (Le Pouce Mt., Piton du Milieu) and western (Piton Brise Fer) mountain ranges have broadly elliptic to obovate, acute to obtuse leaves, hence the specific epithet obovata given by Tulasne. Those from the southern edge ofthe central plateau (Bassin Blanc, Mt. Cocotte, Riviére des Galets) tend to have narrower, ellip- tic to oblanceolate, acute to acuminate leaves. Such variation may occur within a single pop- ulation, however, e.g., at Macabé forest. Variation in floral morphology is expressed in length of stamens and styles, and form and color of androecious flowers. Populations from Bassin Blanc and Riviére des Galets, which grow along river and lake banks, have shorter styles and creamy white, urceolate androecious flowers that tend to split only about halfway open. Popula- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 tions from mountain summits (e.g., Le Pouce and Piton Brise Fer) have pinkish to reddish pur- ple androecious flowers that split completely open into a star shape and the gynoecious flowers have longer styles. Flower color and size and shape of stamens and styles are similarly variable in other species, e.g., 7. tau and T. quadrifida. These flo- ral and foliar differences intergrade gradually in T. ficus, making it impossible to formally rec- ognize infraspecific taxa. For example, the Bassin Blanc population has androecious flowers and styles that are transitional between the Riviére des Galets and Brise Fer forms. As such, they are best recognized as variants of a single, poly- morphic species. In view of this arano, a n nd re that Tami as three cna species of Ambora oe Tulasne (1855a). He based A. obovata on a single sterile, leafy twig from P Pouce and based A. alternifolia on a mixed collection consisting of androecious buds of T. ficus and leaves of Homalium (Flacourti- aceae). Furthermore, because the type of A. ficus consists of leaves and gynoecious flowers of the narrow-leaved form, which moreover was sup- osed to have been collected in Java, it is un- derstandable that Tulasne described it as another species. He was apparently unaware that J. Les- chenault de la Tour first visited Mauritius, where he apparently collected the type of A. ficus, before continuing on to Java (Steenis-Kruseman, 1950). Tambourissa is unknown from Java, and the lo- cality on the label is evidently erroneous (Per- kins, 1911) I have lectotypified Tambourissa alternifolia on the basis of the large, solitary ficiform an- droecious flower buds with characteristic ligu- late, apiculate stamens obviously belonging to T. ficus. I recognize the leaves as belonging to a Mascarene species of Flacourtiaceae [Homalium integrifolium (Lam.) Baill.] also occurring on Le P (1877) wrongly referred material of Tambourissa ficus from Le Pouce to T. elliptica. More surprisingly, he described comparable ma- terial of the same species, also from Le Pouce, as Ficus sapotoides, suggesting he failed to dissect flower buds and also overlooked the absence of stipule scars on the stem. Of the three available concurrently published names, I have retained LORENCE—MONIMIACEAE 159 1985] T. ficus because the type is the most complete, with a androecious and gynoecious flowers and lea e ficus belongs to the small species Group 9, also including T. peltata (Mauritius) and T. castri-delphinii (Madagascar). The ternate to subalternate leaf condition of these species appears to be derived, as seedlings and suckers of T. ficus and T. peltata have opposite leaves. Tambourissa ficus is most closely allied to T. peltata in leaf morphology but its leaves are thin- ner and generally lack the bright yellow-green tinge characteristic of 7. peltata, and the stems are more slender. In addition, 7. ficus is mon- oecious, usually basally cauliflorous, and has much larger flowers with more numerous parts. Furthermore, its gynoecious flowers are urceo- late with setose styles as opposed to those of 7. peltata, which are flat and discoid with columnar styles and produce a thick, mucilaginous exu- date. With respect to gynoecious floral mor- phology, T. ficus more closely approaches Group 5, thus linking the two groups. Vernacular names. Bois tambour, Pot de chambre jacot (Mauritius). 38. Tambourissa peltata R. Br. ex Baker, Fl. Mauritius 288. 1877; Perk. & Gilg, Pflan- zenr. 4, 101: 71. 1901. TYPE: Mauritius. Without precise locality, 1814, Carmichael s.n. (lectotype, K, here designated, as to gy- noecious collection only). Dioecious treelet or tree 12-15 m tall and 30- 40 cm D.B.H., the outer bark smooth, pale brown, longitudinally fissured, the new growth glabrous, the mature leafy stems smooth, stramineate, 3— 7 mm diam., angled or subterete. Adult leaves ternate to subalternate, petiolate, glabrous; pet- iole dpi mm by 1-2 mm; lamina subcoria- ceous to coriaceous, broadly to narrowly ob- ovate, nives bm elliptic, broadly elliptic or rarely ovate, 60-150(-180) mm by 30-70(-85) mm (reaching 225 mm long on vigorous suckers), the apex acuminate to acute, frequently obtuse or retuse, the base acutely cuneate to acutely de- current, the secondary veins 6-9(-12) pairs, making a 55-65? angle with the costa, the ve- nation very prominent and finely reticulate, vis- ible to 4? on both surfaces, the margin slightly revolute; seedling and sucker leaves opposite, heterophyllous, the petiole red when living, the lamina oblanceolate to narrowly oblanceolate, with 1—3 pairs of short teeth apically. Inflores- cence ramiflorous on the leafless branches or up- per portions of the trunk, rarely axillary or on short, leafy shoots, the flowers solitary or in fas- cicles of 2-3. Androecious flower in bud gla- brous, globose to ellipsoid, 10-14 mm diam. by 12-18 mm long, apiculate with 5—6 small, acute glabrous tepals, the pedicel and peduncle 6-20 mm by 1.5-3 mm, bearing few to many deltoid- naviculate, ciliate bracteoles to 1.5 mm long; at anthesis deeply 4—6(-7)-fid, 30-50 mm diam., the thick lobes ultimately reflexing; stamens nu- merous, ca. 200-400, ovoid-deltoid to linear- lanceolate, 4-7 mm long by 1.5-3 mm wide, the filament short or subsessile, the loculi separate or confluent apically, occupying almost entire length of the stamen, the connective broad, slightly prolonged or not, the internal receptacle surface glabrous, rarely with scattered hairs. Gynoecious flower in bud glabrous, broadly na- piform-depressed, 14-17 mm diam. by 7-9 mm long, apiculate with 4 minute tepals, the pedicel and peduncle 6-12 mm by 2 mm, bracteolate as in androecious flower; at anthesis deeply 4—7-fid, 23-30 mm diam., the deltoid lobes ultimately reflexing, their internal surface + verrucose- wrinkled, the receptacle discoid, flat to + con- cave, the peltate central portion 13-16 mm diam., pubescent around the margin and between the styles; styles numerous, ca. 400-500, crowded and coalescent, virtually indistinguishable, co- lumnar, 4—-6-sided, 0.3-0.5 mm long by 0.3-0.4 mm diam., the apex flat to depressed, the gy- noecium covered with a mucilaginous exudate. Fruiting receptacle solitary on the old stems or rarely on the upper part of the trunk, sometimes produced in large numbers, cupuliform, patelli- form or discoid-peltate, 40-130 mm diam. by 15-60 mm long, externally corky brown or mot- tled, the walls 8-20 mm thick, the orifice broad, subentire, comprising 3⁄4 or more ofthe total width of the receptacle, the internal receptacle surface corky brown, with numerous short, bluntly co- lumnar 4-6-sided styles 0.5-1 mm diam. by 0.5 mm long, often subtended by clusters of hairs, the pedicel and peduncle stout, 10-25 mm long by 6-20 mm diam. Mature carpels ovoid-com- pressed, 9-13 mm long, 4-8 mm wide, 3-6 mm thick, the endocarp dark brown, lustrous, the sur- face scrobiculate. Gametic chromosome num- ber, n — Endemic to Mauritius (Fig. 40). e commonest and most widely Distribution. Habitat. T distributed species of Tambourissa in Mauritius, 160 T. peltata displays a wide ecological amplitude, ranging from lowland wet forest (e.g., at Bel Ombre, ca. 300 m, annual precipitation 2,400 mm), to upland evergreen moist forest (e.g., Mt. Corps de Garde, 750 m, annual precipitation 1,600 mm), or cloud forest (e.g., Mt. Cocotte, 760 m, annual precipitation 5,000 mm). Locally abundant in some areas, it may become a sub- canopy or canopy tree (e.g., at Macabé high for- est), or remain a stunted treelet (e.g., at Pétrin in Philippia heath formation). MAURITIUS. BAMBOU MOUNTAIN RANGE: crest be- 3151 (P); wet forest with Sapotaceae, ca. m, 27 Dec. 1978 (st), PE (st), Lorence 221 4 (MO). ves We MT.: 960 (fr), t 003 qu P 101) (fl), Fi riedmann 200-4 top of hill, 1,100 ft. y g pipe to water tower, 30 July 1964 (fr), Vaughan "S MAU 11009 (MAU). cuREPIPE: Forestry Depart- ment nursery, along stream bank with remnants of indigenous trees, ca. 700 m, 15 Dec. 1978 (fr), Lorence 2151 (MAU, MO); 14 Mar. 1971 (fr), Remy sub MAU eg (a U). GAULETTES SERREES: north-central Mau- us, low mixed wet forest over lava, ca. 400-450 m 19 May 1979 (fl), Ted et al. 2642 (K, MO); (fl, fr), Lorence et al. 2643 P). LION MT.: by side of path leading to ae along crest, 22 Mar. 1976 (fr), Lalouette sub MAU 17 (MAU). MACABE FOREST: Macabé to Brise Fer al low mixed evergreen forest over laterite, ca. 660 m, 8 Dec. 1978 (fr), Lorence 2101 (MAU, MO); rea forest, near kiosk, transitional wet forest, ca. 660 m, 16 Dec. 1978 (st), Lorence 2154 (MO); Macabé forest, ca. 700 m, 7 Feb. 1975 (fr), Lo- rence sub Coode 4791 (K, MAU); near path leading to Macabé via the marshes, 14 Feb. 1963 (fr), Vaughan (fl), Guého sub MAU 12543 (MAU, 2 sheets); low, mixed wet forest along stream, ca. 700 m, (fl), Lorence 2586 (K, MAU, MO AGNE DES CREOLES: Mauritius, EL e evergreen forest, ca. Tae m, 25 Jan. 1979 (st), Lorence & Julien 2338 (MA MO): (fr), ids & Julien 2339 (MAU, MO); o Lorence & Julien 2340 (MO); (fr), Lorence & Julien 2341 wae P). PERRIER NATURE RESERVE: near Mare aux Vac ow mixed wet forest, ca. 600 m, 27 Mar. 1979 (f). Lorence 2555 (MO); 9 Apr. 1979 (fl), Lorence ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 2585 (K, MO); 23 Apr. 1976 (fr), Richardson et al. 4009 (K, MAU), (fr), Richardson et al. 40 14 May 1970 (fl), Vaughan sub MAU 14184 (MAU): (fl), Vaughan sub MAU 14185 (MAU, 2 sheets); 22 Apr. 1970 (fl), Vaughan sub MAU 14193 (MAU, 2 Ue by stream, 9 Oct. 1975 (fr), Vaughan sub MAU 17543 (MAU). PÉTRIN NATURE RESERVE: Philippia heath formation over laterite, 630 m, 4 Apr. 1979 (fl), Lo- rence 2572 (MO, P). PIETER BOTH MT.: S flank below La Fenétre, very degraded wet forest invaded by ex- otics, ca. 400 m, 23 Jan. die (st), dar d icis MO). PLAINE CHAMPAGNE: road to Plaine Cha gne, dense indigenous thicket, 21 July 1966 p Guého sub eau, invaded by exotics, grading mS cloud fori mit, 600-700 m, 14 D 8 (fr), ice 2146 (K, MO); 14 Feb. 1979 o, Loren 2402 (B, K, MAU, MO, P, REU, Z) DES: = Hardy s.n. (MAU). TAMARIN RIVER: ri ear Tamarin River Bridge, at Mare aux Pu 2 p 1966 (fl), Vaughan sub MAU 12168 (MAU, 2 sheets); 5 May 1966 (fl), Vaughan sub MAU 12240 (MAU, 2 sheets). VACOAS IDGES: 21 Jan. 1968 (fr), Guého sub MAU 13077 (car- pol. coll. 175) (MAU). WITHOUT PRECISE LOCALITY; (fr), A AU); 16 Aug. 1962 (fr), Tilbrook sub MAU 10548 (MAU); 3 Sept. 1962 (fr), d'Unienville sub MAU 10624 (MAU); 30 Mar. 1964 (fl), d'Unienville sub MAU 11206 (MAU, 2 sheets). Leaf shape in Tambourissa peltata is relatively variable, a character which tends to remain con- stant within populations. Populations from the eastern mountain ranges (e.g., Montagne des Créoles, Mt. de Lion, and Mt. Bambous) are characterized by narrowly elliptic to oblanceo- late leaves, whereas most populations from the upland plateau have the usual obovate leaves. Fruit morphology ranges from typically cupuli- form to flat and discoid, both extremes occa- sionally on the same tree (e.g., Lorence 2151). Androecious flower size and color are variable within populations but remain constant on a giv- en tree. Dark purple-red androecious flowers are most common, whereas the salmon pink ones are less frequent. Gynoecious flower color is less variable, the central disc being reddish orange and the sterile lobes dull purple-red and verru- cose. At anthesis, the disc secretes a copious thick, clear mucilage which covers the styles and is probably functional in pollination (see Endress, 161 1985] 1979, 1980b). Flowers of both sexes produce strong, sour odors of fermenting fruit or toma- s. Tambourissa peltata belongs to the small species Group 9, whose members are character- ized by ternate to subalternate leaves, also in- cluding 7. ficus (Mauritius) and 7. castri-del- phinii (Madagascar). Tambourissa peltata strongly resembles and is apparently closely re- lated to the latter species; both are ramiflorous with similar androecious flowers and stamens, and have leaves with prominent, finely reticulate venation. Tambourissa peltata differs by its larg- £ er, solitar y OI is the only species known to have flat, discoid gynoecious flowers with star-like sterile lobes. Flowers of 7. ficus are much larger than those of the preceding species and are produced at the swollen base of the trunk. The only other dioe- cious species in Mauritius, 7. cordifolia, does not seem to be closely related to 7. peltata, but rather belongs to species Group 5, although the two are known to hybridize naturally at Pétrin. Vernacular name. us). Bois tambour (Mauriti- IMPERFECTLY KNOWN SPECIES 39. Tambourissa lastelliana (Baill Drake in Grandidier, Hist. 1902; Perk., Pflanzenr. 4, 101 (N 1911. Monimia lastelliana Baill., Ball Mens. Soc. Linn. Paris 1: 342. 1882; Perk. & Gilg, Pflanzenr. 4, 101: 65. 1901. TYPE: Mada- gascar. Without precise locality, 1841, de Lastelle s.n. (holotype, P). Dioecious (?) shrub or tree (?), the mature leafy stem glabrous, 4 mm diam., longitudinally striate- pustular, the rays relatively narrow, the nodes dilated, with multiple axillary buds, the inter- node 80 mm long. Leaves opposite, petiolate; petiole recurved, 15 mm by 2 mm, glabrescent; lamina thickly chartaceous, brown, adaxially gla- brescent, abaxially with scattered simple, straight to slightly curved simple hairs 0.5 mm long, denser and longer (to 1-1.5 mm) along the major veins and costa, the lamina broadly obovate- cuneate, + involute, the apex shortly acuminate, the base cuneate, the apical portion of the lamina with 4—5 pairs of broad, deltoid teeth 1.5-4 mm long, the tips indurated, the venation mixed, ba- sally festooned brochidodromous, apically cras- pedodromous, the secondary excurrent into LORENCE-— MONIMIACEAE the marginal teeth, ca. 5 pairs, making a 35-40? angle with the costa, the venation strongly de- pressed adaxially and visible to 3°, raised abaxially and visible to 4?, the margin revolute. Androecious inflorescence ramiflorous (?), pre- sumably a dichasium or the flowers solitary, the floral axis pilose-tomentose basally, fulvous, ca. 10 mm by 1.5 mm or longer, subtended by sev- eral obtuse-deltoid tomentose bracteoles. Sub- mature androecious flower in bud 6—7 mm diam., globose, densely golden pilose externally, the apex slightly depressed, with several minute tepals, the pedicel ca. 5 mm by 1 mm, golden pilose; stamens ca. 60—70 per flower, in bud ca. 3-4 mm long by 1 mm wide, the loculi lateral, separate, ccupying the medial ? of the stamen, laterally dehiscent, the filament short, the connective pro- longed, acute, the internal receptacle surface with scattered simple hairs between the stamens; tan- niferous idioblasts and oil cells present in the receptacle wall. Distribution. Presumably endemic to Mad- agascar. The type is said to be from eastern Mad- agascar (Baillon, 1882), presumably from low or mid altitude evergreen wet forest. The type of Monimia lastelliana consists of a single, detached leaf, a leafless twig, and several detached androecious flower buds. Unlike Mo- n or uniquely stellate trichomes, the leaf of de Las- telle s.n. has dentate margins in the apical por- tion, mixed craspedodromous and brochidod- romous venation, and bears only simple trichomes as do the flower buds. Furthermore, stamens of de Lastelle s.n. lack the paired basal appendages characteristic of Monimia and have instead long, laterally dehiscent loculi and an acutely prolonged connective, all lacking in Monimia as noted by Perkins (1911). The pollen is too immature to permit diagnosis. Neverthe- less, it is clear that de Lastelle s.n. does not rep- resent a species of Monimia, nor even belong in the subfamily Monimioideae (sensu Thorne, 1974), and was correctly removed from the genus by Drake del Castillo (1902). Its eglandular stamens with laterally dehiscent loculi, and long, uniquely simple trichomes in- stead suggest placement in the subfamily Mol- linedioideae (sensu Thorne, 1974) with the other Madagascan genera. The globose buds with mi- nute, reduced tepals most closely recall those of 162 Tambourissa. Although venation is festooned brochidodromous in most Tarmbourissa species, that of T. longicarpa and T. thouvenotii is basally brochidodromous and tends to become craspe- dodromous in the toothed apical portion of the lamina, much as in de Lastelle s.n. On the basis of androecious floral morphology and leaf struc- ture, it is most probable that de Lastelle s.n. ac- tually does represent a species of Tambourissa as proposed by Drake del Castillo (1902). 40. Tambourissa sp. A. Habit unknown. Leaves opposite, petiolate; petiole 17-24 mm long; lamina chartaceous, el- liptic, 80-100 mm by 34-43 mm, the apex acu- minate, the base cuneate, the venation visible to ongate floral m long; the androecious flower in bud cylindrical, obovoid-ellipsoid, ca. 12215 mm long, apiculate, the pedicel 7 mm by 1 mm; at anthesis splitting deeply into 3 segments; sta- mens numerous, ligulate, ca. 2 mm long, the lo- culi confluent apically, occupying nearly the en- tire length of the stamen; flowers red when living. Gynoecious flowers, gynoecious inflorescence, and fruit un MADAGASCAR. TAMATAVE: Iaroka, Maroantsetra, 1,000 m, 20 June 1968 (fl), Rakotozafy 847 bis(MAD). The elongate androecious buds most closely resemble those of Tambourissa quadrifida from Mauritius. It probably represents an undescribed species from the mid altitude Yambourissa/ Weinmannia wet forest zone. 41. Tambourissa sp. B. These collections have opposite leaves, the petiole to 23 mm long; lamina coriaceous, ob- ovate, to 85 mm by 48 mm, the apex obtuse, emarginate, the base cuneate, the margin revo- lute. Immature androecious flower buds rami- florous on leafless nodes, solitary, the pedicel ca. 20 mm long, the bud 7-8 mm diam., the stamens with lateral loculi and a prolonged connective. MADAGASCAR. FIANARANTSOA: Ampamaherana, 14 Dec. 1949 (fl), Anon. sub SF-1252 (MAD, P). The species resembles Tambourissa crassa from Réunion in leaf, but in the latter species the flowers are larger and always terminal, and the connectives are not prolonged. This collec- tion probably represents an undescribed species for which I could not find the precise locality on any map. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 42. Tambourissa sp. C. Tambourissa amplifolia sensu Cavaco in Humbert, Fl. Madagascar 80: 34. 1959 non (Bojerex Tul.) A.DC Tree to 5 m, the leaves large, the petiole 10- 16 mm long, the lamina ovate-elliptic, 140-180 mm by 60-80 mm, shortly acuminate apically, rounded basally, the secondary veins 4-7 pairs. Unlike Tambourissa amplifolia (Mauritius), the petioles, costa, and veins are densely hirtellous xiall ADAGASCAR. FIANARANTSOA: Farafangana, Befo- taka, 17 Aug. 1926 (st), Decary 5040 (MAD, P); NE Madagascar, without precise locality (st), Thiny s.n. in 1904 (P). These collections either represent an unde- T. decaryana, for which only inflorescence leaves are known. 43. Tambourissa sp. D. Habit unknown. Leafy stems pale brown, ve- lutinous, 1.5-2 mm diam., the new growth ve- lutinous. Leaves opposite, petiolate; petioles 5— 10 mm by 1 mm, velutinous; lamina subcoria- ceous, adaxially glabrescent, abaxially pubes- cent, especially along the costa and major veins, ovate-elliptic, the apex acute to shortly acu nate, the base acutely cuneate, slightly Sinis the secondary veins pairs, making a 6 angle with the costa, the venation visible to 2(—3°) adaxially, to 4? abaxially, the margin entire, revolute. Flowers unknown. Fruiting receptacle (fide Baron's pencil sketch of 6720, K) terminal, bglobose, ca. 45-55 mm diam., the orifice sub- circular, subentire, comprising ca. '4—'4 total width of the fruit. Mature carpels unknown. MADAGASCAR. N Madagascar (probably piEGO SUAREZ): without precise pii ca. 1882 (st), Baron 67 - abe see also Baron’s sketch of a fruit with the specimen). Distribution. Presumably endemic to Mad- agascar The collection consists of two sterile leafy branches plus Baron’s pencil sketch of the fruit. The leaves are similar to those of Tambourissa sieberi (Mauritius), but differ in having numer- ous oil cells dispersed throughout the laminar tissue and in having less highly branched vena- tion (only to 4°), nor does 7. sieberi ever have solitary, terminal fruits. It probably represents an undescribed species. 1985] LORENCE— MONIMIACEAE 163 LITERATURE CITED AXELROD, D. I. & P. H. RAVEN. 1978. Late Creta- ceous and Tertiary vegetation history in Africa. Pp. 77-130 in M. J. A. Werger (editor), Biogeog- — and Ecology of Southern Africa. W. Junk, BALLON, E 1868-1870. — sur les Mo- ées. Adansonia 9: 111- 71. The Natural pegs a Plants, Volume 1. L. Reeve & Co., London. [Translated by M. M. Hartog.] — 1882. Liste ps _— de Madagascar. Bull. Mens. Soc. Linn. BAKER, J. G. 1877. Flora al Mauritius and the Sey- chelles. L. Reeve & Co., 1882. TOT to the flora of central Madagascar. J. Bot. n 266-2 BALFOUR, I. B. 79. An o of the petrological botanical and zoological collections made in guelen's Land and Rodriguez during the etin of Venus expedition carried out by orders of Her Majesty’s Government in the years 1874-75. Phil- os. Trans. 168: 289-387 Bawa, K. S. 1974. Breeding systems of tree species of a lowland tropical community. Evolution 28: 92. P. A. OPLER. 1975. Dioecism in tropical forest xin Evolution 29: 167-179. BEHNKE, H.-D. 1977. Transmission electron micros- copy and systematics of flowering plants. /n K. Kubitzki (editor), Flowering Plants: Evolution and Classification of iis Categories. Pl. Syst. Evol., Suppl. 1 . 1981. Sieve- pM Nord. J. Bot. 1: 381-400. BENTHAM, G. & J. D. HOOKER. 1883. Genera Plan- tarum, Volume 3. L. Reeve & Co., London. Hortus Mauritianus. Port Louis, Borror, D. J. & D. M. DELONG. 71. An Intro- duction to the Study of Insects, 3rd edition. Holt, Rinehart & Wilson, New York. BRENON, P. 1972 ogy of Madagascar. Pp. 27-86 i Richard-Vinard (ed- itors), Biogeography and Ecology in Madagascar. W. Junk, The Hague. Brown, R. 1814. General remarks, geographical and systematical, on the botany of Terra Australis. Pp. 553—554 in M. Flinders, A Voyage to Terra Aus- tralis. London. BULLOCK, A. A. 1960. Nomenclatural notes: XII. The types of some generic names. Kew Bull. 14: 40- 45. CADET, L. J. T. 1977. La végétation de l'Ile de la Réunion: étude K... et phytosocio- logique. 2 volumes. Ph.D. thesis. L'Université d em Marseille I CANDOLLE, A. 1868. Prodromus Systematis Naturalis ee Vegetabilis 16(2): 658-661. Mig oot S. Island Life. Natural History New York. . 1974. Island Biology. Columbia Univ. Press, New York. Carr, S. G. M. & D. J. CARR. 1961. The functional significance of syncarpy. Phytomorphology 11 249-256. 1957a. A new combination in Ephip- , and a new species of Tambourissa (Mo- nimiaceae). Kew Bull. 12: 228. 7b. Sur les Tambourissa (Monimiacées) e Madagascar et des Comores. Bull. Soc. Bot. France 104: 283-285. . 1957c. Sur deux genres de Monimiacées. Bull. Soc. Bot. France 104: 610-613. 1957 Deux Monimiacées nouvelles de Madagascar: Tambourissa decaryana Cavaco et T. capuronii Cavaco. Bull. Mus. Hist. Nat (Paris), Ser. 2, 29: 287-288. 1957e. Quelques espéces nouvelles de Mad- agascar: Monimiaceae et Annonaceae. Bull. Mus. Hist. Nat. (Paris), Ser. 2, 29: 351-352. 58a. Sur le genre Decarydendron (Moni- miacées) Bull. Soc. Bot. France 105: 38-39. 1958b. Une espéce nouvelle de Decaryden- CAVACO, A. piandra m nouveau pour une espèce de Tambourissa. Bull. Soc. Bot. France 105: 39-41. Monimiacées. /n H. Humbert Sai a Flore de Madagascar et un Comores. 80* Famill Firmin- "a 19 ' Quelques considérations à propos de su ey Webbia 29: 587-592. CORDEMOY, E. J. DE. 1895. de de l'Ile de la Ré- n. Paul Klincksieck, 76. The Seeds of Dicotyledons, bridge Univ. Press, Cambridge . Contribution à l'étude des espèces gascar. Bull, Mus. Hist. Nat. ap 28: 247-251. 928. Contribution à l'étude des Monimi- acées de Madagascar. Bull. Mus. Hist. Nat. (Paris) 34: 278-280. DEcAISNE, M.J. 1858. Description d'un nouveau gen- re de plantes de la famille des Monimiacées. Ann. Sci. Nat. Bot., Ser 4, 9: 278-279. DIETRICH, F. G. 1802. T n der Gärtnerei und Botanik, Volum DircHER, D. 1974. Approaches to de identicaiihn of angiosperm leaf remains. Bot. Rev. (Lancaster) 40: 1-157. 1972. The climate of Madagascar. Pp. n R. Battisti . Richard-Vindard (ed- 2 Biogeography, wae Ecology in Madagascar. The Ha: DRAKE DEL red El 1902. Histoire des plantes de Madagascar. Volume 1 in A. Grandidier (edi- tor), Histoire Physique, Naturelle et oe de Madagascar. à Wr rimerie Nationale, Pari Du PrrIT-THOUA . A. 1804. unc des végétaux = sur r les d de France, la Ré- Mad DONQUE, G. Paris Pikpnpoxayk, F., F. KRENDL, E. HABELER & W. SAU- ER. 1968. Chromosome numbers and evolution in primitive angiosperms. Taxon 17: 337-353. ENpRzss, P. K. 1972. 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La Grande Comore: climat et végé- tation. ae Sect. Sci. Tech. Inst. Fr. Pondichéry 3(5): 1 LEMESLE, R. po . PRICHARD. 1954. Les charactéres histologiques du bois des Monimiacées. Rev. Gen. Bot. 61: 69-96. Leroy, J.-F. 1978. Composition, origin and TE E the Madagascan vascular flora. Ann. Missou Bot. Gard. 65: 535-589. LINDLEY, J. 1853. The Vegetable nn 3rd edi- 1976. The end dius: of Ro- uin Island. Bot. J. Linn. Soc. 72: 269-282. . The Wade E of pr (Indian t. J. Linn. Soc ue 207-247. . 1980. A Systematic and Eco-evolutionary Study of the Monimiaceae in the M : 82. region (SW Indian Ocean). B (Paris), Ser. 4, apt B, sd 293- 304. [1981.] . ZENGER & P. ViN 1984. Pollen mor- phological Sud on the Mod e ofthe Mal- agasy region. Grana 23: Lv-Tio-FANE, M. 1976. piene sonnei 1748- Imprimerie & Pateterie Commerciale, ue Mauritius. McDouaarL, I. & F. H. CHAMALAUN. 1969. 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Out-of-print. many now considered synonyms, giving family placement for each. Monogr. Syst. Bot. Missouri | sat rd. 3. Mosses (1)— FLo F SOUTHERN AFRICA. R. us deeds xv * 291 pp. 1982. 930. 00. The first of ES ded fascicles. One-h phag throu Gri laceae are treate OSSES OF EASTERN NORTH AMERICA. H. A. Crum & L. E. Anderson. 2 vols. 1328 pp. Illustrated. 1981. $7 This highly acclaimed flora is the product of two lifetimes' SUT and will be the sadi eferenc decades to come. We include two copies o eric index, which can be glued conveniently over a LE papers of each volume. C MossES OF GUATE E. B. Bartram. 442 pp. Illustrated. 1949, reprinted 1972. $8.00. Still the only moss ia ora for any Central American country and is a fundamental reference for those studying the mosses of the Neotropics. Fieldiana: Bot. 2 S MOSSES OF THE INTERIOR HIGHLANDS E Wed AMERICA. P. L. Redfearn, Jr. 104 pp. 1972, reprinted with changes and additions 1983. $15 Treats about 300 species with Mae E key, generic descriptions, keys to species. Notes 0n habitats, occurrence, distribution. Ann. Missouri Bot. Gard. 59 (1). Musci AUSTRO-AMERICANI. G. Mitten. 652 + vi pp. 1982. $15.00. 21 A reprint of Mitten's 1869 classic, long out-of-print, which treats about 1700 mosses from the West Indies and Central and South America. Monogr. Syst. Bot. Missouri Bot. Gard. 7. Md ORDER FORM 1. Please type or print mailing address. 2. No shipments until — eceived: 3. Make check or NOM order He to Missouri Botanical Garde funds, and payable Drang U. - bank. Send order to: ven Missouri Botanical Garden P.O. Box St. Louis, MO 63166-0299 U.S.A. Title Adda $1 nn; ANNALS JF THE ISSOURI BOTANICAL GARDEN ^LUME 72 1985 NUMBER 2 CONTENTS tt, Familial Position of the Cape Genus Empleuridium Peter Goldbla Hiroshi Tobe, Sherwin Carlquist & Varsha C. Patel n- A. Why do Some Compositae Have an Inconsistently Deciduous Pappus’ ae BERN — — — — — -.— — —— Z: Z E Loran- Fine Structure of Mistletoe Pollen VI. Small-Flowered Neotropical Lc 167 187 thaceae Sylvia M. Feuer & Job Kuijt .————7—— ca, Fuc a nuda PE The Systematics of the Apetalous Fuchsias of South America, Fuc pie Mu Hemsleyella (Onagraceae) Paul E. Berry . d "ioi 252 Collecting ang Preparing Specimens of Araceae “Thomas Pu Dimorphic Incompatibility in Turnera hermannioides Camb. nA 259 Spencer C. H. Barrett & Joel S. Shore n- TE ” nee G1 D'Arcy The Plants of ‘Ocoquili’ Island, San Blas Coast, Panama ras 264 & Barry Hammel . ye ; Antirheg aromatica UNE Güetitrdeae), a New oui from Veracruz, = Mexico Gonzalo Castillo-Campos & David H. Lorence — J ard W W. > New Species of Rhipidocladum from Mesoamerica Rich 272 NH — . — Systematics of the South African Genus _ (ridaceae- kaod FFF PA Contents continued on back cover VOLUME 72 SUMMER 1985 NUMBER 2 ANNAL OF THE MISSOURI BOTANICAL GARDEN The ANNALS, published quarterly, contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden, | t. Louis. Papers originating outside the Garden will also be ac- cepted. 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 first issue of this volume. EDITORIAL COMMITTEE | NANCY Morin, Editor | Missouri Botanical Garden CHERYL R. BAUER, Editorial Assistant Missouri Botanical Garden MARSHALL R. CRosBv j | Missouri Botanical Garden GERRIT DAVIDSE Missouri Botanical Garden JOHN : Missouri Botanical Garden & St. Louis University PETER GOLDBLATT Missouri Botanical Garden For subscription information contact the Business Office of the Annals, P.O. Box 299, St. Louis, MO 63166. Subscription price is $65 per volume U.S., $70 Canada, and Mexico, dvds ee | $75 all other countries. Personal subscriptions are available at $30 and $35, respec 3 Airmail delivery charge, $30 per volume. Four issues per volume. x n b- The ANNALS OF THE MISSOURI BOTANICAL GARDEN (ISSN 0026-6493) et p postage paid at St. Louis, changes to the ANNALS OF THE MISSOURI BOTANICAL GARDEN, P.O. Box 29. St. Louis, MO 63166. à: € Missouri Botanical Garden 1985 ANNALS OF THE MISSOURI BOTANICAL GARDEN VOLUME 72 1985 NUMBER 2 FAMILIAL POSITION OF THE CAPE GENUS EMPLEURIDIUM! PETER GOLDBLATT,? HIROSHI TOBE,? SHERWIN CARLQUIST,* AND VARSHA C. PATEL? ABSTRACT The monotypic southwestern Cape genus Empleuridium was assigned with owe ies Rutaceae binis first described in 1860. Po orly known until recently, it has been rediscovered an made possible detailed studies of its embryology and reproductive anatomy; en pur leaf suci and pollen. Results confirm that Empleuridium is misplaced in pear and it seems best Te ferr ed to Celastraceae, in which it is mue in its ericoid habit, small s t. The very s small flowers have a Hosa disc in hich the inferior 1S flow into a Wes like single-seeded fru e incorrect ;the ow ovary is embedded. Previous pedal that are ZR The monotypic genus Empleuridium was first described in 1860 and was based on a collection made by the early Cape collectors Ecklon an Zeyher in the 1820s. Publication was nearly si- multaneous in “Flora Capensis” (Sonder, 1860) and in a series of illustrations, the “Thesaurus Capensis,” intended to supplement the Flora (Harvey, 1859-1860). Empleuridium is usually attributed to Sonder, and the species E. juniper- inum to Sonder & Harvey. It was assigned with doubt to Rutaceae by Sonder and has since re- mained in this family in generic floras of South- ern Africa (Phillips, 1926, 1951; Dyer, 1975). Empleuridium is a small shrub with needle- like leaves and small tetramerous flowers, the A Esterhuysen, Univers ity our study and for C in the collection of more material, together wit ry. We also thank Peter Raven for encouraging this study ae for mts the authors Department of Fores four stamens located at the edge of a floral disc and alternating with the petals. It is rare and apparently had not been recollected for over 120 years when Elsie Esterhuysen, the well known Cape botanist who has made a study of the Cape mountain flora, found plants growing on rocky mountain slopes at two localities in the south- western Cape. She realized that her find was un- usual and collected ample material, including a supply of specimens fixed in FAA. Subsequently, - the first author suggested to her that the plants in question might be the poorly known Empleu- ridium. An examination of the type material at Stockholm (S), made by the Director, Dr. B. Nor- denstam, confirmed this. together, and Rolf Dansa. Botanical Museum, Copenhagen, for reviewing the manuscri A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, S Louis, Missouri 63166. ? Department of Biology, Jes of Science, Chiba University, E * Yayoi-Cho, Chiba 260, Japan. 917 * Rancho Santa Ana Bota c Garden, Claremont, California 5 Department of Botany, Tin ersity of Oklahoma, Norman, iud 73019. ANN. Missouni Bor. GARD. 72: 167-183. 1985. 168 Empleuridium apparently does not belong in Rutaceae according to Dr. I. Williams (pers. comm.), who has revised several southern Af- rican genera of this family, including Empleu- rum, to which E L i lly sim- ilar. Because the affinities of E mpleuridium are not known, this study was undertaken in the hope of establishing more satisfactorily the relation- ships of the genus. Embryological and floral anat- omy studies were made by Hiroshi Tobe; stem and leaf anatomy by Sherwin Carlquist; and pol- len by Varsha Patel. MATERIALS AND METHODS Plants used in this study were collected in the wild and preserved in FAA. Voucher data is as news Empleuridium juniperinum Sonder & Harvey South Africa: Cape, Caledon district, above De Rust, 3,000 ft., Esterhuysen 34318 (BOL, MO, RSA). South Africa: Cape, Caledon district, upper slopes of Mt. Lebanon, 3,800 ft., Goldblatt 6918 (MO) Wood studies were supplemented with her- barium material ofa portion of root (Esterhuysen 34318-RSA). EMBRYOLOGY For light-microscopic observations, FAA pre- served flowers and fruits at various stages of de- ` velopment were dehydrated through a t-butyl al- cohol series and embedded in Paraplast with a melting point of 56—58°C. Serial sections cut 5— 10 um thick were stained with Heidenhain's he- matoxylin, safranin and fastgreen FCF, and were mounted in Canada Balsam. Electron-micro- scopic observations on seeds were also made fol- lowing a standard technique and using JSM-25S VEGETATIVE ANATOMY Dried root material was boiled preparatory to further treatment. Prior to sectioning, both the ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FAA and dry material were treated in ethylene diamine for softening in preparation for paraffin sectioning, a technique (Carlquist, 1982) in which small twigs or objects with both hard and soft portions can be sectioned successfully. A safra- nin-fast green combination was used for staining sections. Macerations were prepared with the aid of Jeffrey's flui POLLEN Buds containing pollen were treated by ace- tolysis (Erdtman, 1960), air dried from 9596 eth- anol, sputter coated with gold, and examined with n International Scientific Instruments Super II scanning electron microscope. Slides of glycerine jelly mounts were also examined with transmit- ted light using a Leitz Ortholux microscope. OBSERVATIONS FLORAL MORPHOLOGY The flowers are solitary and borne in the axils of the needle-like leaves. The pedicels are 1.5-2 mm long at flowering, elongating up to 3 mm in fruit (Fig. 4). A pair of tiny opposed scaly bracts (Fig. 4) are present at the base of the pedicel. The flowers are small, 1.5-2 mm long and ca. 2 mm in diam.; perfect and tetramerous, comprising four sepals and four petals apparently arranged in whorls, but decussate in young buds (Fig. 2). There are four stamens and an ovary (Figs. 2, 3; see also Fig. 6) of probably four carpels (see next section). The stamens are alternate with the pet- als (Fig. 2), i.e., haplostemonous. Aestivation of the sepals and petals is imbricate (Fig. 2). The filaments are short, ca. | mm long. A style is also very short with a capitate stigma (Fig. 4). A non- nectariferous disc is slightly developed in the in- trastaminal area. Both the sepals and the style are persistent. The ovary is hemi-inferior and embedded in the receptacle (Fig. 1; see also Fig. 6A, B). The ovary is unilocular with four Nocrasionally a erect ovules, an 1, 3) that would represent the fused lateral walls — FiGures 1-5. Morphology and anatomy of flower and fruit of Empleuridium. se, sepal; pe, petal; st, stamen; ov, ss sc, scaly bract; pd, pedicel; fr, fruit coat; sd, seed.— 1. Longi 200 um.— 2. Tr dead section (TS) of a flower bud at the level of ake Note i 100 u and ie Scale = contains only three ovules. i = 200 um.—4. is nearly mature. cavities. Scale = 200 um tudinal section of a flower bud. Scale = mbricate aestivation of sepals m.—3. TS of a flower bud at ovary level. Note the ovary is unilocular and occasionally Fruit in various sages i development. The one on the ri — 5. TS of a young fruit: tl * VU VIUI y 1985] GOLDBLATT ET AL.—EMPLEURIDIUM 169 170 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 VIDEA YII 5555153224 NOSE IIT cate š FiGURE 6. Diagrams illustrating the floral anatomy and vasculature in Empleuridium. Symbols are the same as those in Figures 1-5.— A, B. Median longitudinal sections of a flower through the line marked a and b in — C-H. Transverse sections of a flower at levels marked c-h in A and B. 1985] of the constituent carpels. The ovules are alter- nate to the stamens (see Fig. 6F). Placentation is basal (Figs. 1, 6B). Although the ovary is initially inferior, it elon- gates beyond the level of the stamens after fer- tilization (Fig. 4). At maturity all but a tiny part of the fruit extends beyond the receptacle. The fruit is 7-9 mm long, fusiform and slightly curved with a tapering persistent style. It is always one- seeded (Fig. 5) and is follicle-like, although strict- ly a capsule since it is derived from a compound ovary. The fruit wall is thin and consists of four to five cell layers and lacks secretory cavities expected in Rutaceae. The large, black seed has a distinctive white aril or elaiosome at the base which has several thread-like coiled extensions. This is described in more detail in a later section. FLORAL ANATOMY Vasculature of the flower is very simple, and does not form a network by mutual fusions of Vascular bundies in dou toceptecle: At the basal part o lakral) bundles (Fig. 6C), sac supplying the: vas- cular traces to either a sepal/stamen or a peta particular ovary-wall part/ovule, depending on the dispositional relationship between the vas- cular bundle and appendages (Fig. 6D—H). The sepals, petals, and stamens each have a single bundle (Fig. 6H). In the wall of the inferior ovary there are eight small vascular bundles that extend upward into a short style (Fig. 6E, G). Vascular strands entering the four ovules are separately derived from the four vascular bundles, which are opposite the four petals respectively, and each of which emits traces not only to a petal but also to the dorsal part of the ovary wall. The vascular anatomy suggests that the four ovules in the single ovary are originally those of four carpels. It seems that the unilocular hemi- inferior ovary of Empleuridium, containing four ovules evolved from a four-carpellate, septate compound ovary by loss of septa. VEGETATIVE ANATOMY Wood (Figs. 7-12). Wood of stems (Figs. 7, 8) is described first here; features by which root wood differs follow. Stem wood with occasional vague growth rings demarcated by wider vessels in earlywood. Vessels are angular to rounded in transectional view; mean vessel diam., 15 um; mean vessel wall thickness, 1.4 um. Mean num- GOLDBLATT ET AL.—EMPLEURIDIUM 171 ber of vessels per sq. mm is 844. Mean vessel element length, 136 um. Perforation plates sim- ple, a few with a transverse bar. Lateral walls of vessels with alternate or scalariform pitting. Im- perforate tracheary elements all tracheids with fully perdened pits. Mean tracheid wall thickness, 3 sel abundance, contact of axial parenchyma is virtually inevitable), axial parenchyma in strands of two cells. Rays few, uniseriate, all composed of markedly erect cells; the wood appears nearly rayless (Fig. 8). Wood non-storied. Wood of root like that of stem except for the following features. Root wood with little growth ring activity (Fig. 9). Mean vessel diam., 25 um; mean vessel wall thickness, 2.5 um. Mean num- ber of vessels per sq. mm is 494. Mean vessel element length, 316 um. Perforation plates sim- ple (Fig. 11) or with one to three bars (Fig. 12). Lateral walls of vessels with alternate pits or pits which may be provisionally termed scalariform (Fig. 11). Mean wall thickness of tracheids, 4.6 m; mean tracheid length, 520 um. Wood nearly rayless (Fig. 10), an ^ bisce uniseriate ray composed of erect cells presen Node, cortex, and phloem. UR of Em- pleuridium juniperinum are all unilacunar with a single trace. The cortex is composed wholly of parenchyma cells except for a few scattered pro- tophloem fibers in stems; fibers absent in roots (Fig. 15). Scattered tannin-bearing idioblasts and cells are subdivided into strands of crystal-con- taining cells; the crystals are single and rhom- boidal, or few per cell, but are not aggregated into druses as in cortical cells. Leaf. Although linear, the leaf is slightly flat- tened as seen in transection (Fig. 13). Stomata are ranunculaceous and present on all portions of the leaf surface. A thick cuticle is present and is raised into an overarching hood external to stomata (Fig.16). Chlorenchyma is two or three layers thick; cells are only a little longer than nly main vein is present; a small veinlet may branch from this on either side. The main vein consists 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 " . ^ ` ` ` ` a , ` a e ` L] Eu "Sea! er w Or 7 oS VETE "y - “<Ñ t) » sloe SA) See duca 7-12. Wood PES of Empleuridium. 7, 8. Stem, from Goldblatt 3418 (MO).—7. Transection, t left, pith at right. — 8. Tangential section. A few elongate ray cells may be seen. 9-12 ot, fi pieds 34318 (RSA).—9. Abi iua tracheids are thicker walled than in stem.— 10. Tangential section. A single ray is present, u upper left. 11, 12. Vessels from radial section. — 11. Simple perforation plate e left); aha. lateral wall pitting (right). — 12. Perforation plate traversed by a single bar. Figures 7-10, scale above Figure 7 = 10 wm; Figures 11, 12, scale above Figure 11 = 10 um 1985] GOLDBLATT ET AL.—EMPLEURIDIUM 173 i P Ficures 13-16. Sections of sisi rig idi 13, 14, 16, sections i = from Goldblatt 3418 (MO).— 13. Leaf transection, adaxial surface above.— 14. Longitudinal section of vein in leaf; tracheids with scalariform and alternate circular pits at ape fiber at left. — 15. Transection of nias of. root, from Esterhuysen 34318 (RSA). ses visible in some cells, tannin in others; periderm at right.— 16. Stoma from leaf, showing overarching cuticular ridges. Figures ti 15, scale above Figure 7; Figures 14, 16, scale above Figure 11. 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 1985] ofa laterally wide bundle sheathed adaxially and abaxially by fibers. The xylem ofthe vein consists of annular, helical, and scalariformly pitted tra- cheids, but also tracheids with circular bordered pits (Fig. 14, right). No secretory cavities were observed. EMBRYOLOGY Anther and microspores. The anther is tetra- Les The wall structure prior to matu- ration comprises four layers; epidermis, endo- thecium, one middle layer, and tapetum (Fig. 18). Cells of the middle layer and endothecium are derived from the same cells (Fig. 17), and the wall formation is regarded as conforming to the dicotyledonous type (Davis, 1966). During mat- uration, the epidermis mostly collapses and the middle layer degenerates while the endothecium develops fibrous thickenings. The tapetum is glandular and its cells become two-nucleate (Fig. 19) and finally degenerate. Thus the mature an- ther wall at dehiscence consists principally of the fibrous endothecium (Fig. 20). Dehiscence is lon- itudinal. Meiosis in microspore mother cells is accom- panied by simultaneous cytokinesis (Fig. 19). The shape of the microspore tetrads, on the basis of the examination of 50 selected tetrads, is usually (9490) tetrahedral and very occasionally (696) de- t Pollen grains are released singly and are two-celled (Fig. 2 Megagametophyte and nucellus. The ovule is anatropous and tenuinucellate. À single ar- chesporial cell differentiates beneath the apical dermal layer of the small nucellus (Fig. 22), and directly develops into a megaspore mother cell (Fig. 23). No case was observed in which two or more archesporial cells were borne in a single nucellus. The megaspore mother cell normally undergoes meiosis, resulting in a linear or T-shaped tetrad of megaspores (Fig. 24). The chalazal megaspore of the tetrad always func- tions and successively develops into a two- (Fig. GOLDBLATT ET AL. —EMPLEURIDIUM 175 25), four- (Figs. 26, 27), and eight-nucleate Polygonum-type embryo sac. The synergids ex- hibit no particular specialization. Two polar nu- clei are united into a central nucleus before fer- tilization (Fig. 28). The three antipodal cells are ephemeral and always absent in the organized mature embryo sac. Thus prior to fertilization, the embryo sac consists only ofan egg, two syner- gids, and the central nucleus (Fig. 28). uring megasporogenesis and megagameto- genesis, a remarkapie breakdown of the nucellar the embryo sac occurs (Fig. 24). Asa result, the greater part of the embryo sac after the two- or four-nucleate ici is no longer enclosed by the nucellar tissue 1 the inner Figs. PRA The bread ma of the remaining nu- cellar tissue continues until it almost disappears. In a young seed, the enlarged embryo sac is whol- ly surrounded by the inner integument alone. No hypostase is ey during the development of the ovule and s Integuments an endothelium. The micro- pyle is formed by the inner integument alone (Fig. 29). Both the inner and the outer integu- ments are initiated by involving an oblique-peri- clinal or periclinal division of dermal cells in an ovule primordium (see arrows in Fig. 30). The subsequent growth of the integument takes place y the division of cells derived from the dermal initials, while the subdermal cells do not con- tribute to the formation of the integuments. The integuments are thus regarded as being of “‘der- mal origin" (Bouman & Calis, 1977). As early as the megaspore mother cell stage both the in- teguments are usually three cell-layers thick. In older stages, the constituent cells further divide periclinally, so that the outer integument be- comes four cell-layers thick while the inner in- tegument increases the thickness into more than six cell-layers. Neither the outer nor the inner integument is vascularized. A vascular bundle in the funiculus ends at the chalaza and does not exhibit post-chalazal branching. — FIGURES 17-21. thecium; ml, middle layer; t, tapetum Development of anther walls and microspores in Empleuridium. ep, epidermis; en, endo- ; mc, microspore mother cell; ms, microspores in telophase of meiosis II; v, nucleus of the vegetative cell; g, nucleus of the generative cell. — 17. Transverse section (TS) of a young anther. Scale — 10 um.— 18. TS of an older anther than that shown in Figure 17. Scale — — 19. TS of an older anther than that shown in Figure 18. Scale = 10 um.— 20. TS of a mature anther. Note that the wall comprises the fibrous andothecium alone. Scale = 100 um. — 21. Section of a mature pollen showing the two-celled state at the time of shedding. Scale = 10 um. 176 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 1985] As the nucellar tissue breaks down and the embryo sac comes to border directly on the inner integument, cells of the inner layer of the inner integument tł di 11 AGNX1011 bj elonga ted and enlarged (Figs. 24, 28) to form a weakly de- veloped endothelium (i.e., integumentary tape- um Fertilization, endosperm, and embryo. Fertili- zation is porogamous and endosperm formation is nuclear. Wall formation in free endosperm nu- clei occurs in later stages, and the young seed is filled with a cellular endosperm. The mature seed is albuminous with copious endosperm (Fig. 31). The endosperm is oi Our fruit samples were not adequate to pursue the embryogeny, but based on microtome sec- tions of several proembryos, the embryogeny of Empleuridium appears to proceed normally. A suspensor appears to be small and short. The mature embryo is narrow, straight and dicoty- ledonous (Fig. 31). Both cotyledons develop equally. The mature seed has a large white aril (elaiosome) (Fig. 33) formed by cells derived from the outer integument and probably from the fu- niculus. It consists of a cushion-like base con- nected to the micropyle and is drawn distally into coiled filaments. The surface is covered with mi- nute trichomes (Fig. 34). e seed coat is formed principally by cells of the exotesta and the exotegmen (Figs. 31, 32) that were previously those of the outer layer of the outer and the inner integument, respectively. In the testa, only the cells of the exotesta are persistent and become tanniferous and cuboidal in shape while cells of the remaining part are completely crushed. The cells of the exotegmen are elongate longitudinally and radially, though the longitudinal elongation is more conspicuous, GOLDBLATT ET AL.—EMPLEURIDIUM 177 and become fibrous with pits at places; in ad- dition, cells of the endotegmen that previously formed the endothelium, become slightly elon- gate longitudinally and tanniferous; cells of all the remaining parts are completely crushed. POLLEN The pollen grains are Hicolporate, radially isopolar, subp te in lateral view Fin. 35, 37), and triangular-pleurotreme in po- lar view (Fig. 36). The surface is reticulate with the widest lumina at the poles (Figs. 35-37). Lu- mina size decreases toward the equator (Fig. 35) with the smallest lumina around the colpi (Fig. 37). Columellae supporting the muri (i.e., the lumina ridges) are spindle-shaped, unbranched, erect, and with lengths directly proportional to o elliptic er rounded polar axis is ca. 26 um and the eiua Pied is ca. 21 um. DISCUSSION RELATIONSHIPS Empleuridium does not belong in Rutaceae nor to any families in this alliance. It was as- signed here when the nature of the gynoecium was unknown and the genus was believed to be dioecious. Empleuridium is discordant with Ru- taceae in having a nonglandular floral disc; the five stamens alternate with the petals rather than opposite to them; an inferior, non-septate uni- locular ovary probably composed of four fused carpels; a single-seeded fruit; and in entirely lack- ing glands in the leaves and other organs (Cron- quist, 1981). At the embryological level Em- — FiGuREs 22-30. Development of ovules and digi qni gie preis in Empleuridium. arc, archesporium; mc, megaspore mother cell; fm, functional megaspore; nl and n wo nuclei in the two-nucleate embryo sac; nl- n4, four nuclei in the four-nucleate embryo sac; eg, egg in comprising an egg cell and two synergids; cn, central nucleus formed by fusion of two polar nuclei; ii, inner dnt, degenerating nucellar tissue; mic, micropyle. —22. Longitudinal section (LS) of an ovule primordium at the archesporial cell stage. The archespo Antipodal cells are not seen because showing the micropyle formed by th showing the mode of initiation of integu orium is one-celled. Scale — 20 mise: LS megaspore mother cell stage showing the tenuinucellate condition. Scale = 20 um.— the megaspore tetrad stage. Note that the tissue of the um.—25. LS of a young ovule at the two-nucleate embryo sa m.— 26, 27. Two of an im M at the oung ovule at aped p art of jm nucellus is jpn Scale = tage. Note that the embryo sac directly borders successive LSs of a young ovule at the four eh. cai with an ep Polygonum- -type e le = 20 u S ofa ca = 50 scone P. of an ovule wheads indicate that both the Sie aig are “initiated by oblique-periclinal or periclinal divisions of deb cells of the ovule primordium. Scale = 178 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 1985] pleuridium contrasts with Rutaceae in having two-nucleate anther tapetum cells in contrast to two multinucleate cells forming polyploid masses by fusion (Johri & Ahuja, 1957; Desai, 1962); a nucellus that degenerates early (persistent in Ru- taceae); and the absence of a hypostase (Mau- ritzon, 1935, 1936; Davis, 1966). Vegetative anatomy also supports the exclusion of Empleu- ridium from Rutaceae. The wood has tracheids, rather than libriform fibers, and lacks the secre- tory cavities typical of Rutaceae (Metcalfe & Chalk, 1950). Morphologically and embryologically, Em- pleuridium accords well with Celastrales in the broad sense (e.g., Cronquist, 1981), and once its characters are correctly interpreted it keys out to this order in standard keys to the flowering plants. It agrees with Celastraceae in particular, having small bisexual haplostemonous flowers, with a non-secretory floral disc, short filaments and a short persistent style. Embryological characters also accord with Celastraceae, notably the weakly qevelopeo and early — nucellus, dad oily endosperm be 1953; Copeland, 1966: Corner, 1976). The anatomy of Empleuridium is relatively unspecialized. Most of its features are distributed widely in the dicotyledons (ranunculaceous sto- mata, druses, rhomboidal crystals, tannin cells). The relatively primitive wood with tracheids rather than libriform fibers suggests affinities with several orders with similar wood, e.g., Saxifra- gales, Rosales, as well as Celastrales. In fact, Em- pleuridium accords particularly well with Stack- housiaceae, an Australasian group related to Celastraceae, but morphologically very different from Empleuridium and clearly not directly re- lated to it. The strikingly reticulate surface, subprolate shape, tricolporate apertures, and size of Em- pleuridium pollen all agree with morphological descriptions of Celastraceae (Erdtman, 1971; Lobreau-Callen, 1977, 1978; Lobreau-Callen & Lugardon, 1972-1973). In personal correspon- GOLDBLATT ET AL.— EMPLEURIDIUM 179 dence with D. Lobreau-Callen, the pollen au- thority for the Celastrales, Empleuridium is fa- vorably compared with several members of the Celastraceae (viz., Pterocelastrus rostratus, Mys- troxylon aethiopicum, Plenckia bahiensis, South African species of Maytenus, and some species of Cassine). A difference with Celastraceae pollen is apparent in the poor development or absence of an endexine fold in the aperture, a character heretofore noted in all members of the family (Lobreau-Callen & Lugardon, 1972-1973). Em- pleuridium pollen is distinguished within the family (and order) by the reticulum lumina which are larger at the poles than at the equator (see Fig. 35). Empleuridium is discordant with Celastraceae in its being a low ericoid shrublet, and in flora structure in the inferior unilocular ovary and the micropyle formed from the inner integument alone. It is also unusual in its one-seeded fruit, basal placentation with erect ovules, and the re- markable aril with coiled filamentous extensions. However, in a few Celastraceae the ovary is metimes nearly inferior (e.g., Paxistima) al- (e.g., Bhesa, Plenckia, Zinowiewia, Microtropia) although usually many-seeded (Corner, 1976); and the ovule is sometimes basal and erect (e.g., Euonymus, Behsa, Paxistima) (Berkeley, 1953; Corner, 1976). There is even a comparable seed in the Malaysian Sarawakodendron, in which the aril has branched filaments on the cushion-like base (see Corner, 1976, vol. 2: 81). Thus, as regards the reproductive character- istics, the difference between Empleuridium and Celastraceae seems to be limited to the non-sep- tate and unilocular ovary and the micropyle formed by the inner integument alone. Even in these features there are examples that reduce the isolation of Empleuridium. In Celastrus orbic- ulatus Lam. the septa do not fuse in the center of the ovary (Berkeley, 1953), a condition that can be regarded as transitional from (septate) plurilocular to (non-septate) unilocular. The di- — FiGuREs 31-34. Seed and seed coat of Empleuridium. em, embryo; end, endosperm; entg, endotegmen; extg, exotegmen; exts, exotesta; ar, aril.— 31. a straight embryo and the endosperm. Scale = 500 u Longitudinal section of a micropylar half of the mature seed containing m. — 32. Transverse section of a nearly mature seed. Note the seed coat is formed principally by the exotesta pa the fibrous exotesta. Scale = 200 um.— 33. Scanning electron micrograph of the mature seed showing an aril with elaborate structure. Scale = 1 mm.— 34. Scanning electron micrograph of a part of the aril. Scale = 100 um 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 FIGURES 35-37. with lumina at the equator. Scale Pollen of E AK dii dade — 35. Lateral view. The widest lumina are on nh AL apt ey =5 Polar view emphasizing the wide lumina. Scale = 5 um.— 37. y um. Lateral view. Around the margin of the copus the lumina are greatly reduced. Scale = versity of ovary structure in the Celastraceae is far from thoroughly studied, and similar po ET carpel fusion and septa reduction may occ other members of the family, not necessarily r tia & Gavde, 1962). The number and the kind of the components of the micropyle is a very important taxonomic character at higher levels, but the significance of the condition in Empleu- ridium should perhaps not be emphasized. ANATOMY The very narrow vessels of Empleuridium stems, ca. 15 mm in diam., are noteworthy. Few — Morphology of xi ene — A. Branch with ripe fruits as well as flowers. — B. Leaf with E 38. E sasa stipules. — C. Flower buds. — D. tage. — F. Sam S e, longitudinally sesiones —G. Fruit in stage of dehiscenc to cares the ER arillate seed; the aril is partly remo removed and laid open. All from [eim 34318 (LD, C). Del. B. Johnsen (orig.). posed. — J. Aril, Flower after anthesis, the anthers shed. —E. Flower in even later e. — H. Fruit, half of the wall removed ved in the right figure, so that the conical funicle is GOLDBLATT ET AL.—EMPLEURIDIUM 1985] 182 dicotyledons characteristically have vessels this narrow, although Metcalfe and Chalk (1950) re- ported vessels of about 15 um diam. in such species as Bergenia delavayi (Franchet) Engl. (Saxifragaceae). Vessel elements so narrow, so short, and so numerous per sq. mm oftransection as those of Empleuridium are clear indicators of xeromorphy (Carlquist, 1977). The wood of the root of Empleuridium is somewhat less xero- morphic than that of the stem according to these measures. Wood of roots is generally more me- somorphic than that of stems (e.g., Carlquist, 1978; Carlquist et al., 1983) The simple perforation plates in the xylem of Empleuridium are also an indicator of xeromor- y. Perforation plates with one to three bars, which are occasional in the species, seem relic- tual of a more primitive condition. They are suf- ficiently regular in appearance that they cannot be considered a malformation. The scalariform lateral wall pitting on vessels might be a remnant of a primitive condition, although a pseudosca- lariform expression (lateral widening of pits based on an ancestry of alternate circular pits) cannot be ruled out. The occurrence of tracheids in Empleuridium wood rather than fiber-tracheids or libriform fi- bers is a primitive feature in the wood, despite the specialization of perforation plates in vessels. The combination of simple perforation plates in vessels with occurrence of tracheids as the im- perforate tracheary element type has been noted as unusually abundant in woods of dry areas, notably chaparral and desert (Carlquist, 1980; Carlquist et al., 1983). The adaptive value of this formulation is the ability to conduct water rap- idly during the brief periods when it is available, permitted by simplification of the perforation plates, combined with the safety of tracheids (adapted for conduction, but air embolisms formed under water stress do not spread from one tracheid to another as they do with vessel elements). The wood of Empleuridium is nearly rayless. In dicotyledons at large, raylessness characterizes phylesis to woodier stature (Carlquist, Empleuridium is not very woody: it can be de- scribed as a diminutive woody herb. The ray- lessness (or presence of uniseriate rays, com- posed of erect cells only) in Empleuridium can be considered related to its habit. There is so little wood accumulation that one cannot say that this genus is in the process of becoming woodier. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 The leaf of Empleuridium shows several ad- aptations to xeromorphy, such as condensed form, thick cuticle, cuticular domes over sto- mata, and isolaterally distributed chlorenchyma with rather small intercellular air spaces. The presence of unilacunar nodes in Empleuridium may also be related to the condensed linear leaves, because many species with linear leaves have unilacunar nodes, even in groups in which broad- er-leaved species have trilacunar nodes. SUMMARY It seems reasonable to assume that Empleu- ridium is a highly specialized member of Celas- traceae, and that the unusual features of its flower and fruit are reductions from the basic type for Celastraceae. Clearly its small size, reduced woodiness, and needle-like leaves are specializa- crophyllous habit is found in numerous taxa that in many cases have broad-leafed relatives. The specializations in wood anatomy reflect the ad- aptation to a Mediterranean climate by a small ericoid subshrub, based on a plan compatible with celastraceous woods. SYSTEMATIC TREATMENT Empleuridium Sonder in Harvey, Thesaurus Ca- pensis 1: 49. 1859; Flora Capensis 1: 442. 1860; Phillips, Genera S. African Fl. Plants, ed. 1. 353. 1926; ed. 2. 442. 1951; Dyer, Genera Southern African Fl. Plants 1: 292. 1975. TYPE: E. juniperinum Sonder & Har- vey. Figure 38. E. juniperinum Sonder & Harvey, Thesaurus Ca- pensis 1: 49-50. 1859; Flora Capensis 1: 442. 1860. TvPE: South Africa. Cape Province: *Bergrucken zwischen Caledon und Baby- lonschetoorn, 1,000-2,000 ft., Ecklon & Zeyher s.n. (52.8) (S) Ericoid, erect, glabrous, eglandular, under- shrub. Leaves simple, needle-like, scattered, 9- 11 mm long, with small scale-like stipules. Flow- ers axillary, solitary, small, tetramerous, acti- nomorphic, with a non-secretory receptacular disc, aestivation imbricate; pedicels 1.5-2 mm long, elongating to 3 mm in fruit, with small, paired opposed, scale-like bracts at the base; se- pals 4, persistent, ovate, acute; petals alternate to the sepals, whitish, ovate, acute, free, spread- ing. Stamens 4, opposite the sepals, erect, fila- 1985] ments short, ca. 1 mm long, anthers with 2 the- cae, introrse. Ovary hemi-inferior, initially embedded in the receptacular disc, elongating well beyond the level of petal and stamen inser- tion after anthesis, unilocular, with 4 erect, basal ovules; style short, persistent, stigma capitate. Fruit large, ca. 7 mm long, fusiform, follicle- likes 1-seeded black, large, ca. x mm long, with a large white basal aril (elaiosome). Distribution. SW Cape, mountains in the Ca- ledon district, mainly above 600 m, on rocky sandstone slopes. Material examined. 33.19 (Worcester) Blokkop (Aasvogelberg) above Villiersdorp, sun and shade at the rocky summit, ca. 5,200 ft. ew Esterhuysen 35233 a Ye NBG, S), 35575 (BOL, S). .19 (Caledon) Groenland Mts., Mt. Lebanon, 3, i ,800 ft. (AA), Goldblatt 6918 (MO); Mt. Leb- , local, Esterhuysen ,Sy “bergriicken zwischen Ca- ledon und Babylonschetoorn, 1,000-2,000 ft.” (?AC), Ecklon & Zeyher s.n. (52.8) (S) LITERATURE CITED ADATIA, R. D. & S. G. GAvpr. 1962. Embryology of the Celastraceae. In Plant Embryology — A Sym posium 1-11. November 11-14, 1960. CSIR, New Delhi. BERKELEY, E. 1953. Morphological studies in the Cel- astraceae. J. Elisha Mitchell Sci. Soc. 69: 185-206. Bouman, F. & J. J. M. Cauis. 1977. Integumentary shifting—a m way to unitegmy. Ber. Deutsch. Bot. Ges. 20: 28. CARLQUIST, S. on Wood anatomy of insular species of Plantago and the problem of raylessness. Bull. Torrey Bot. Club 97: 353-361. 1977. Wood anatomy of Onagraceae: addi- tional species and concepts. Ann. Missouri Bot. Gard. 64: 627-637. 1978. Wood anatomy of Bruniaceae: corre- lations with ecology, phylogeny, and organogra- phy. Aliso 9: 323-364. 1980. Further concepts in ecological wood a y, with comments on recent work in wood anatomy and evolution. Aliso 9: 499-553 . 19 The use of ethylenediamine in soft- ening hard plant structures for paraffin sectioning. Stain Technol. 57: 311-317. khart & D. C. Michener. 1983. Wood 5 GOLDBLATT ET AL.— EMPLEURIDIUM 183 anatomy of Hydrophyllaceae. I. Eriodictyon. Aliso 10: 397-412. COPELAND, H. F. . Morphology and embryology of Euonymus japonica. Phytomorphology 1 334. 6: 326- Corner, E. J. H. 1976. The Seeds of Dicotyledons. Cambridge Univ. Press, Cam CRONQUIST, A. 1981. An Integrated System of Clas- tion of or Plants. Columbia Univ. Y 1966. Systematic Embryology of the perms. John Wiley Sons, New Yor ^1962. Cytology d ner be aÀ of the Ru- ae. Phytomorphology 12: 178-184. Drar. R A. 75. The Genera of Southern African Flowering Plants, Volume 1. 3rd edition. Depart- ment of Agricultural Technical Services, Pretoria. ERDTMAN, G. 1960. The acetolysis method, a revised description. Svensk Bot. Tidskr. 54: 561—564. I E: 71. Pollen Morphology and Plant Tax- onomy — Angiosperms. Hafner, New York. Haney, W. 1859-1860. Thesaurus Capensis. Vol- me 1. erus Smith & Co., Dublin M. & M. HUJA. 1957. A contribution — S. Les pollens des Celas- trales. (Illustrations, commentaires.) Mem. Trav. Inst. Montpellier 3: 1-116. New interpretation of the variations of the exine structure of simple apertured pollen grains in the Celastrales. IVth E Palynol. Conf., Lucknow (1976-1977) 1: 185- ——— LUGARDON. e pA L'aperture a ae du pollen des Celastraceae. Naturalia Mon- , Ser. Bot. 23, 24: 2 ! MAURITZON J. ; ber die Embryologie der milie Rutaceae. Svensk Bot. Tidskr. 29: 319- Zur Embryologie und systematischen ung der Reihen Terebinthales und Celas- ea Bot, gi rond us 2. LFE, 1950. Anatomy of the Dicoty hy Ciarendon Press, Ox = E. P. 1926 The Gen era of South African wering Plants. v edition. Cape Times, Cape own : 1951. The Genera of South African Flow- ame oo 2nd edition. Government Printer, oon R. ^1982. Descriptors used to indicate abun- dance and frequency in ecology and systematics. Taxon 31: 89-94. SONDER, O. W. Rutaceae. /n W. H. Harvey & O. W. Sonder (editors), Flora nne 1: 360- 447. Hodges, Smith and Co., Dublin WHY DO SOME COMPOSITAE HAVE AN INCONSISTENTLY DECIDUOUS PAPPUS?! A. SHMIDA? Many species of Compositae produce seeds equipped with a pappus that acts as a parachute, allowing the seeds to be dispersed by wind (Zo- hary, 1937, 1950; Pijl, 1972; Burtt, 1961, 1977). Surprisingly, in some other species the pappus is formed but deciduous, i.e., separates very easily from the body of the seeds, usually before dis- persal. This trait seems paradoxical — why man- ufacture a dispersal apparatus and then drop it before dispersal? To our knowledge, no adaptive value for deciduousness has been put forward in the literature. Nonetheless, brun Is a common phenomenon recorded in Compositae from virtually all tribes, regions, ind habitats. As an initial stage in investigating possible adaptive values of a deciduous pappus, I have analyzed the Compositae in the flora of Israel for correlations between deciduousness and other ecological parameters. The results of this survey, which I present here, suggest that inconsistent deciduousness is adaptive primarily as a mech- anism for achieving a mixed dispersal strategy, in which seeds that lose their pappus prior to dispersal remain in the vicinity of the mother plant (“‘atelechory”’ sensu Ellner & Shmida, 1981). The analysis is based on Feinbrun-Dothan (1978) supplemented by extensive observations of species distributions and seed characters in Israel. Several species which have been found only sporadically in Israel, and appear not to be established in the flora, were excluded from the analysis. Seeds bearing only a few pales or awns, or a corolla, were regarded as epappose. Copies of the species-list with scorings for all attributes used in the analysis are available on request. À naspas ani en attributes were examined by constructing 2 x 2 contingency tables in which the rows were de- ed by deciduousness versus persistence of the pappus, and the columns by presence versus ab- sence of the other attribute. The tables were tested for significance at the level a = 0.05 by either the chi-square test or (if any of the expected cell counts were <5) the exact multinormal test for a I Of the 230 Compositae found in Israel, 42 have a deciduous pappus (DP), 112 have a per- sistent pappus, and the remainder are epappose. About half (22) of the species with a deciduous pappus are annuals. However, no significant as- sociations were found between DP and growth- orm (annual, biennial, perennial) or life-form (annual, hemicriptophyte, chamaephyte) in the Mediterranean region, the desert region, or both regions combined. I also investigated possible associations between DP and the following spe- cial habitats: Mediterranean region, desert re- gion, rock outcrops, vado and ruderal. A significant positive association was found be- tween DP and ruderal eA (Table 1a) (be- cause all but two ruderals were classed as Med- iterranean, only Mediterranean species were considered). However, this association appears to be an artificial consegnence or two other as- sociations: a strong ation between DP and pappus dimorphism (Table 2), and a negative association between pappus dimor- phism and ruderal habitats (Table 1b). If I re- move this confusing trend by excluding species with a pappus dimorphism, there remains a pos- itive but highly non-significant association be- tween DP and ruderal habitats (Table 1c). Thus, the only significant trend is that DP gen- erally does not occur in species with pappus di- morphism. The term dimorphic pappus is used here in any case in nen the achene of the same capitulum l l type of pappus. The transition between. type of pappus Shmida & Ellner, in prep). In the Se of ferae pappu 1s di variably PS | » | *4 r invah ination of the pappus on a portion of the achenes ! [ want to express my gratitude to Professors D. Cohen and S. Ellner p Missae discussions and to of Isra Professors M. Zohary and N. Feinbrun who att racted me to the flora l 2 Department of Botany, Hebrew University, Jerusalem, Israel. ANN. Missouni Bor. GARD. 72: 184—186. 1985. 1985] TABLE Deciduous pappus and pappus dimor- phism in relation to ruderal habitats in the Mediter- ranean region Compositae of Israel (cell entries are numbers of species). R on-ru- deral deral (a) Deciduous pappus* 16 15 Persistent pappus 20 52 (b) Pappus dimorphism present? 3 27 Pappus dimorphism absent 23 40 (c) Species without pappus dimor- phism? Deciduous pappus 16 14 Persistent pappus 17 26 a The 2 x 2 tables in (a) and (b) (involving all Med- iterranean region Compositae) depart significantly from independence (x? = 4.42 an .84, P « 0.05 and P « 0.025 respectively). The table in (c) (Mediterra- nean Compositae without a pappus dimorphism) does not depart significantly from independence (x? — 0.86, P > 0.25). in each capitulum. DP and pappus dimorphism can therefore be viewed as alternative mecha- nisms which reduce the efficacy of wind-dispersal in some (but not all) seeds. The finding is that species tend not to utilize both alternatives. This is illustrated nicely in the genus Crepis: of the 12 species found in Israel, six have a deciduous monomorphic pappus, and six have a persistent pappus with a marked pappus dimorphism be- tween marginal and central seeds in a capitulum (Feinbrun-Dothan, 1978). Similarly, in the Fi- lago group (sensu Wagenitz, 1969; Feinbrun-Do- than, 1970), Cymbolaena and Lasiopogon spp. in Israel have a deciduous, monomorphic pap- pus, while in Filago and Ifloga, the central fertile achene in each capitulum has a persistent pappus and the marginal seeds are epappose (Feinbrun- Dothan, 1978). These results suggest that DP and pappus di- morphism are alternative means to some end, but give no clue as to what that end might be. The most obvious conjecture has to do with seed dispersal distance. A pappus classed as “‘decid- uous" may separate from the seed either before or after dispersal. My observations of Lactuca spp., Carduus spp., Onopordon spp., Silybum marianum, Urospermum picroidis, and Noto- basis syraica indicate that some of the seeds pro- duced by an individual lose their pappus before dispersal, while others lose their pappus only upon impact after dispersal. SHMIDA—INCONSISTENT DECIDUOUS PAPPI 185 TABLE 2. Deciduous pappus in relation to pappus dimorphism in the Compositae of Israel (cell entries are numbers of species). Medi- terra- nean Desert Entire Region* Region* Flora Pappus dimorphism present? Deciduous pappus 2 2 Persistent pappus 14 24 38 Pappus dimorphism absent Deciduous pappus 14 25 38 Persistent pappus 32 46 74 a Five species found in both Mediterranean and des- ert regions are recorded in eac > The paucity 2 dispersal- heterocarpic species with deciduous pappu is significant at a in each region and i si mm ntire flora (exact test for a 2 x table iod. to An column individually). DP and pappus dimorphism thus both result in a partial loss of “long-distance” dispersal by wind. Mixed dispersal strategies, in which some seeds disperse while others remain near the par- ent plant, have been found to be optimal in var- ious models of species in spatiotemporally vari- able environments (Hamilton & May, 1977; Comins et al., 1980; Motro, 1982a, 1982b; Levin et al., 1983) and also in spatially structured but temporally constant environments (Hamilton & May, 1977; Motro, 1982a, 1982b). In patchy, variable habitats, dispersal allows a species to colonize suitable patches. Exploitation of suit- able patches by subsequent generation requires retention of some seeds in the patch. Partial dis- persal achieves both these ends, and a partial dispersal strategy is therefore generally optimal (Shmida & Ellner, 1984). Other adaptive values for the lack of a pappus can also be conjectured. For example, epappose seeds may enter more readily into small cracks or openings in the soil, and buried seeds may avoid predation and thermal extremes faced by seeds lying at the soil surface (Ellner & Shmida, 1981). Conversely, epappose seeds are smaller and presumably easier for insect predators to handle. The question remains, why produce a pappus if it is advantageous to lose it? Why not reduce or eliminate the pappus on some seeds? I hy- pothesize that the reason is evolutionary flexi- bility. A pappus “‘lost” by deciduousness can be regained by strengthening the attachment of pap- 186 pus to seed, a minor modification compared to redeveloping the entire pappus. However, the contemporary suggestion that a deciduous pap- pus can easily be reconverted to a functional con- dition (= high evolutionary flexibility) should be demonstrated experimentally in the future. The fraction of wind-dispersed seeds can therefore vary over ecological time. This feature would be most valuable in patchy, variable habitats where newly-created patches remain suitable for exten- sive periods, during which time non-dispersal would be favored, a hypothesis which should be tested in the future. When a patch starts to be- come unsuitable, dispersal would be favored. Preliminary observations in the field with the Israeli Compositae flora indicate much variation within the dispersal dynamic of species with an inconsistently deciduous pappus: a e pappus can break before or after dis- persal and the percentage of pre-dispersal break- age versus post- -dispersal breakage changes with- n known from the systematic literature, at least in some species, there is prominent variation in the field in the percentage of the capitulum’s achenes which dispersed with or without a pappus. We hypothesize that such variation can serve has a ynamic mechanism of evolutionary flexibility toward selection (and reversed selection) of an adaptive phenomenon of a deciduous pappus on the whole, as well as the adaptability of an in- consistently deciduous pappus. The large, spiny ruderals with deciduous pappi found on anthropogenic mounds ofnitrogen-rich soil (species of Carduus, Silybum, Notobasis, Cynara, and Onopordon) may be an example of this syndrome. New mounds are constantly being created, and persist for several years. Popula- tions of Silybum marianum and Onopordon cyn- arocephalum in different localities vary mark- edly in the fraction of seeds losing the pappus prior to dispersal (70—10096 in ten populations of Silybum, 60-100% in three populations of On- opordon), possibly representing different phases ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 in the cyclic selection for dispersal and non-dis- persal, a hypothesis that should be tested. LITERATURE CITED Burtt, B. L. 1961. functional evolution. Trans Edinburgh 39: 216 1977. azas of diversification i in the capit- ulum. Pp. 41-59 in V. H. Heywood, J. B. Har- borne & B. L. Turner (editors), The Biology and Chemistry of the Compositae. Academic Press, ndon and New York. Comins, N. H., W. D. HAMILTON & R. M. May. 1980. eibi M stable dispersal strategies. J. Theor. Biol. 82: 205-230. ELLNER, S. & A. SHMIDA. 1981. Why are adaptations for long-range seed dispersal rare in desert plants? Oecologia 51: 133-144. FEINBRUN-DOTHAN, N. 1970. A key to the species of Filago re sensu lato oe in Palestine. Israel J. Bot. 19: 260 Compositae and the study of roc. Bot. Soc. Flor a Pa mam Part 3: Ericaceae to mpositae. Israel Academy of Science and Hu- lem HAMILTON, R. M. Mav. 1977. Dispersal in stable habitats. bor 269: 578-581. LEVIN, , D. CoHEN & A. HasriNGs. 1983. Dis- persal sirategis ii in P cud environments. Theor. Pop. Biol. Morno, U. 19822. rs mn rates of dispersal. 1. Hap- loid populations or. Pop. Biol. 21: 394-411. . 1982b. Optimal rates of dispersal. 2. Diploid populations. The . 21: 4 Pur, L. VAN DER. 1 2. Principles of Dispersal in Higher Plants. Springer Verlag, New Y SuMIDA, A. & S. ELLNER. 1984. Coexistence of plant species with similar es Vegetatio 58: 29-55. VENABLE, D. L. & L. LAwLoR. 1980. Delayed ger- mination and diced m in desert annuals — escape ace and time. Oecologia 46: 272-28 969. A WHITTAKER, F S. A. LEVIN. 1977. The role of ities. Theo Pop. Biol. 12: 117-139. ZoHARY, M. 1937. Die Melodien ded sn Verhaeltmisse der Pflanzen Palaestinas. Beih. Bot Centralbl. 61A: 1-155. Evolutionary trends in the fruiting head of Compositae. Evolution 4: 103-109. FINE STRUCTURE OF MISTLETOE POLLEN VI. SMALL-FLOWERED NEOTROPICAL LORANTHACEAE! SYLVIA M. FEUER? AND JOB KUIJT? ABSTRACT Pollen of small-flowered neotropical Loranthaceae (8 genera; ca. 135 species) was examined in the light, scanning, and transmission electron microscopes. Pollen is typically medium-sized and oblate. Pollen amb is more variable than equatorial shape, ranging from trilobate deeply concave to circular. Both isopolar and heteropolar pollen grains occur within the complex, the latter restricted to species of Phthirusa and Struthanthus in which the apertures differ at each of the polar faces. Simple apertures arranged i in a diploaperturate configuration predominate. Such apertures ne from diploporate to to C und apertures are rare, restricted to Oryctanthus and particular species of Cladocolea and. | Sasha s Pun The (3-)4-5 colpate apertures of Ixocactus are unique in the famil ily. Sculpturing is — uniform with sculpturing ridges and/or striat e. Ultrastructurally, the basic ektexine structure IS composed of a thin, perforate tectum, prac ated nie interstitium, and a thick continuous foot layer usually twice as thick as the tectum plus interstitium. Pollen data suggest 2 Group I genera, dropemon is closely linked to PAthirusa through the species P. pyrifolia and P. platyclada. Oryctanthus is a highly derived genus with only remote ties to other Group I taxa. Among Group II genera, pollen data indicate a close relationship between Cladocolea and the Mexican species of Struthanthus. Pollen characters of /xocactus indicate no relationship with any small-flowered neotropical genus. Rather its pollen features are closer to those of the Eremolepidaceae and the African species of Viscum also shows a marked tendency towards unisexual flowers and dioecism. Only three of the eight genera exhibit strictly bisexual flowers (Oryctan- thus, m Ixocactus); two genera hirusa, Cladocole In a previous paper we detailed the pollen of the large-flowered neotropical Loranthaceae (Feuer & Kuijt, 1980). The present paper focuses on pollen characters of the small-flowered gen- with era. Data from these two complexes will be used to analyze pollen evolution among neotropical Loranthaceae. The small-flowered neotropical complex com- prises eight genera: Struthanthus (ca. 50 spp.), Cladocolea (ca. 25 spp.), Phthirusa (ca. 30 spp.), Dendropemon (ca. 12 spp.), Cladocolea (ca. 25 spp.), PAthirusa (ca. 30 spp.) Dendropemon (ca. 12 spp.), Oryctanthus (10 spp.), Maracanthus (2 spp.), Oryctina (2 spp.), and Ixocactus (1 sp.). The small flower size and generally inconspic- uous flower color (often pale green, buff) wawas an insect pollination mechanism, sharpl trasting with the large, showy bird- pi flowers of most Old and other New World mis- tletoes (Kuijt, 1969, 198 1a). In addition to having small flowers, the group ! Acknowledgements — We tha We are grateful to M. Forster for his critical comment i a to ue ? Department 3 t strictly bisexual Foca and others with exclu- sively unisexual flowers; three genera (Struthan- thus excluding S. panamensis, Maracanthus, Oryctina) demonstrate only unisexual flowers. Small flower size, the tendency towards unisex- ual flowers and dioecism, and an x = 8 chro- mosome number (Barlow & Wiens, 1971) to- gether suggest an independent evolution of this group unlike Old World and other New World Loranthaceae. MATERIALS AND METHODS Dried flowers were collected from sheets in the following herbaria: A, CAS, F, GH, K, LEA, MO, NY, RB, S, SMU, U, UBC, UC, US, and UT. Pollen was processed according to methods out- nk the curators and staff of the herbaria who allowed removal of pollen material. ^ o J. Kuijt are gratefully acknowledged. istory, Chicago, "Illinois 6 of ield ? Department of Biology, pps of Lethbridge, Don Alberta TIK 3M4, Canada. ANN. Missouni Bor. GARD. 72: 187-212. 1985. 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 TABLE l. Summary of pollen data for small-flowered neotropical Loranthaceae. Size (um) Shape Taxon Polar Equatorial Polarity Amb P/E Cladocolea Tieghem 13-31 23-45 isopolar triangular to 0.64—0.70 (ca. 25 spp. rounded oblate nvex rarely slight- l ncave Dendropemon Blume 23-46 38-56 isopolar triangular 0.52-0.98 (12 spp.) slightly to oblate, subob- deeply con- late, rarely ob- cave late spheroidal Ixocactus Rizz. (1 sp.) I. hutchisonii Kuijt Cuatrecasas 23970 29.7 + 0.13 36.4 + 0.14 isopolar circular 0.82 (0.74—0.87) (F), Colombia (27.8-32y (34.2-38.5) suboblate Hutchison & Idrobo 28.4 + 0.09 32.8 + 0.10 isopolar circular 0.86 (0.80-0.93) 3008 (F), Colom- (27.3-29.8) (31.1-34) suboblate bia Steyermark et al. 28.1] + 0.17 37.9 - 0.11 isopolar circular 0.74 (0.69—0.81) 111592 (F), Vene- (26.9-30.1) | (36.6-38.6) oblate zuela Tamayo 2526 (ILL), — — isopolar circular — Venezuela Maracanthus Kuijt (2 spp.) M. chlamydatus (Rizz.) Kuijt Steyermark 99522 17.5 + 0.28 28.9 + 0.29 isopolar triangular to 0.60 (0.46—0.71) (UC), Venezuela (14.4-19.9) (25.8-29.5) slightly con- oblate vex Oryctanthus Eichler 27-35 45-56 isopolar triangular 0.56-0.70 (10 spp.) rounde oblate convex to circular Oryctina Tieghem (2 spp.) O. subaphylla Rizz. Anderson et al. 15.8 + 0.11 24.9 + 0.09 isopolar triangular 0.63 (0.55—0.65) 36949 (RB), Bra- (14.3-17.8) (23.2-26.1) rounded oblate zil convex Phthirusa Martius 17-39 27-51 isopolar; triangular to 0.60-0.77 (ca. 30 spp.) heteropo- slightly con- oblate, subob- lar; isopo- cave; trilo- late, oblate lar-hetero- bate (2 spp.) spheroidal polar? Struthanthus Martius 14-38 28-50 isopolar; triangular to 0.53-0.91 ) heteropo- rounded oblate to sub- lar; isopo- convex, oblate, rarely rarely con- cave oblate sphe- roidal 1985] FEUER & KUIJT— MISTLETOE POLLEN TABLE l. Continued. 189 Apertures Sculpturing Exine Structure simple, compound; diplo- and synapertur- °... foveolate, low profile granular/columellate, gran- ate (diploparasyndemicolpate, diplopara- riato-rugulate syndemicolporate, diplosyndemicolpate, parasyncolpate) simple; diploaperturate (diploporate to dip- ^ psilate-imperforate polar exine; lobrevicolpate) psilate-perforate equatorial exine 4(—5)-colpate irregular bifurcating processes and spines 4-colpate irregular bifurcating processes nd spine (32), 5-colpate irregular bifurcating processes and spines 4-colpate irregular bifurcating processes and spines syncolpate short ridges, low profile stria- to-rugulate, perforate compound(?); diploaperturate (three sagit- psilate-imperforate tate slits at each face form the endoaper- tures) diplosyndemicolpate psilate, foveolate(?), perfo- rate(?) simple; diplo-, rarely synaperturate (1 sp.) psilate-perforate; short furrows, (diplosyndemicolpate, diplodemibrevicol- ridges; low profile striato-ru- pate, diploporate, diplo-2-demicolpate, gulate syncolpate) simple, compound; diplo- and synapertur- pronounced striato-rugulate, ate (syncolpate, 3-demicolpate, diplosyn- tectate-perforate with low demicolpate, diploparasyndemicolpate, profile rugulae; aud perfo- syncolporoidate, dele mayas tata rate to imperfor: iplosyndemicolporoida iplosyndemi- e colporate, diplopa aep ae. rate) columellate/baculate granular columellate/granular ektexinous strands travers- nh narrow osmiophilic zon columellate/granular, colu- llate/baculate, colu- mellate granular, rarely with spo- radic columellae ? Populational mean followed b. VoU deviation; range in parentheses. b Occurring within same popu * Listed in order of dominant nene element. 190 lined in Feuer and Kuijt (1978) with the follow- ing addition. For light microscopy, acetolyzed pollen was mounted in glycerine jelly and pho- Ee with a Nikon-UFX system at either at 68°F in Rodinal diluted 1 :25. Specimens ex- amined with the transmission electron micro- scope (TEM) are marked with an asterisk. Aperture terms used in the present paper are briefly defined followed by a listing of all aperture types occurring among the small-flowered gen- era. Tue term: 1) demi- is used bs describe a pair ofap ly to the equa- tor but not confluent, thus appearing as if each set of apertures were derived by equatorial con- striction and subsequent splitting; 2) diplo- is used to denote two sets of apertures (in these cases usually three to a set), each set restricted to a polar face and not continuous across the equator; 3) syn- is used to denote apertures fusing at the pole; and 4) parasyn- refers to apertures bifurcating at the pole, the bifurcations isolating a portion of exine at the genter of the polar face. The foll in the small-flowered complex: syncolpate (e.g., Figs. 83, 84), parasyncolpate (e.g., Figs. 68-71), diplosyndemicolpate (e.g., Fig. 74), diplopara- syndemicolpate (e.g., Figs. 74-77), 2- and 3- de 'micolpate (e. 8-» Fig. Th ane compound vari- ( porate) of these same types, as well as diploporate (e.g., Figs. 25-29), diplobrevidemicolpate (e.g., Figs. 1, 52) and 3- 4—5 colpate (e.g., Figs. 99, 100) Pollen characters of the small-flowered genera are summarized in Table 1. with- GENERIC POLLEN DESCRIPTIONS Cladocolea Tieghem (Figs. 5-11, 68, 69, 72, 73, 7-89) ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 TYPE I Exomorphology: radial symmetry, isopolar. Shape: amb triangular concave; oblate. Aper- tures: diplosyndemicolpi; colpi nu of uni- form width, with granular membranes. Sculp- turing: short ridges to low profile striato-rugulae. Exine equatorially thickened near apertures. Species and specimens examined: C/adocolea inconspicua (Benth.) Kuijt, Gentry 7056 (F), Mexico (Sinaloa). TYPE II Exomorphology: radial symmetry, isopolar. Shape: amb triangular to rounded convex; ob- te. Apertures: diploparasyndemicolpori; ec- toapertures shallow colpi deirmiüng a prominent round or triangular apocolpium oapertures ranging from U-shaped to elliptical slits to large circular subsurface thinnings lying perpendicular to and in midregion of colpi on polar faces. Sculpturing: psilate-foveolate to shallow striato- rugulate. Equatorial exine thickened near aper- tures. Species and specimens examined: Cladocolea andrieuxii Tieghem, Pringle 10244 (SMU), Mex- — P (H.B.K.) Kuijt, *Pringle 4369 (UC), Mexico (Ja- lisco). TYPE III Exomorphology: radial symmetry, isopolar. late-foveolate to shallow striato-rugulate. Exine evenly thickened at equator. Species and specimens examined: Cladocolea FIGURES 1- Polar views of Phthirusa spp. (1-3 — s chlamydatus Rizz. (4), Cladocolea spp. m Panama. Pur 1 1), Oryctina STE E ido p> Struthanthus spp. es ç 16, 18-24) All light rp x 700.— troflexa (Ru lepidobotrys Eichler, Jam ace with protrudin: gcolpal margins. ders P. retr Opposite p face o uxii Tieghem Kuijt. Grain with elliptical endoapertures. S. densifl andley.— ee Pavón) Kuijt, Pd (Ruíz & Pavón) Kuijt, Nee E PA 14038 (F), Panama. ame g Focal plane slightly below apocolpial exin dial Kuijt Pringle 46 97 (UC), Mexico. — 7. C. andrieuxii Teighem. Focal plane at the level of De —8. C. andrie . Focal plane at the equator. Note prominently thickened exine microphylla (H.B.K.) Kuijt. Grain with U-shaped endoapertures. — 10. — 11. C. yc dares (Tieghem) P Hinton 10500 (GH), = leptostachyus (H — 14. S. — a Standle ey.— Nee & Hansen 14038 (F), —5. C. inconspicua (Benth.) C. microphylla G. Don, Skutch 2618 (A), Costa Rica. . S. palmeri Kuijt.—16. S. belizensis Lundell.— 17. Focal plane at ACIU, — 18. S. oerstedii Standley, Barlow 1443 (UC), Costa Rica. — 191 FEUER & KUIJT—MISTLETOE POLLEN 1985] —21. S. 19. S. panamensis Voy ) Barlow & Wiens, Davidson 431 (MO), Panama. — 20. S. concinnus Martius. rnell) G. Don, Hatschbach 17641 (UC), Brazil. — 22. S. uraguensis var. brevipedunculata “23. S. vulgaris Martius. Focal plane at the apocolpium.—24. S. vulgaris Martius. Focal h plane at the ae 192 glauca Kuijt, Borys et al. 6 (LEA), Mexico (Pueb- la); C. grahamii (Benth.) Tieghem, Hinton 10149 (GH), Mexico (Oaxaca), Pringle 6987 (F), Mex- ico (Mexico); C. harlingii Kuijt, *Harling 6094 g Breckon & Breckon 803 (GH), E (Oaa, Conzatti 1899 (F), Mexico (Oaxaca), Pringle 4697 (UC), Mexico (Oaxaca). TYPE IV Exomorphology: radial symmetry, isopolar. Shape: amb triangular; oblate. Apertures: diplo- syndemicolpi. Sculpturing: short ridges to low profile rugulae. Polar faces exhibiting central cir- cular thickening; equatorial exine evenly thick- ened Species and specimens examined: Cladocolea oligantha (Standley & Steyerm.) Kuijt, Rze- dowski 22674 (CAS), Mexico (Guerrero). I^ J L 1 Lee d endexine pres- ent. Ektexine organized into tectum, intersti- tium, and foot layer. Tectum evenly thickened, occasionally foveolate, rarely perforate. Intersti- tium narrow, typically sparse and finely granu- late with short, irregularly shaped columellae rom tectum or projecting upwards from foot layer, these intercalated between horizontally coalesc- ing ektexinous strands. Foot layer with typically smooth upper surface and shallowly scalloped lower edge. Endexine absent in interapertural equatorial regions, present in central regions of polar faces and regions adjacent to colpi; polar endexine bistratified with upper zone adjacent to foot layer characterized by numerous osmio- philic channels, basal zone slightly thicker, ho- mogeneous; endexine near apertures irregularly thickened, disrupted by numerous small gaps; apertural endexine thin, disrupted, loosely or- ganized into small lamellae or more granular strands. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 Tectum/Equatorial interstitium//Foot layer: l: Species and specimens examined: all species listed under exomorphology that are marked by an asterisk. Dendropemon Blume (Figs. 25, 26, 29, 32, 36- 40) Exomorphology: radial symmetry, isopolar. Shape: amb triangular slightly concave to trilo- bate deeply concave; oblate to suboblate or ob- late spheroidal. Grains diploaperturate. Aper- tures: ranging from elliptical pores to broad brevicolpi, typically restricted to tips of pollen lobes and encircled by subsurface thickened ex- ine continuous with subsurface triradiate polar thickening, membranes disrupted. Sculpturing: psilate imperforate at the polar faces and tips of pollen lobes, exine minutely perforate in inter- apertural equatorial areas. Exine with triradiate thickening at polar faces and equatorially thick- ene lobes. ktex1 endexine pres- ent. Differences i In n interapertural equatorial, lo- tectum, columellate/baculate interstitium, and foot layer. Tectum with narrow perforations. In- terstitium exhibiting short, basally thickened columellae interspersed among rounded baculae pendent from tectum or extending up from foot layer. Foot layer thick with smooth upper and Polar ektexine thicker than equatorial ektexine, bistratified; upper stratum imperforate with smooth upper surface; lower stratum signifi- cantly thicker, upper surface slightly granular; strata separated by a narrow, poorly defined zone often filled with granules and occasionally bridged by extremely narrow short columellae. Endexine ranging from thick, continuous and coarsely granular to thin and locally absent beneath the FIGURES 25-35. Comparisons between Dendropemon Blume and d Martius (LM, SEM). 25, 26, 29. 32. Dendropemon spp. 27, 28, 30, 31, 33-35. Phthirusa spp.— 25. s Ste D. unifloru ud., Leonard et al. 15026 (UC), Haiti. Focal plane just below the apocolpial exine, x 500.— 26. D. one Steud., Leonard et al. 1 5026 (UC), Haiti. Light micrograph with focal plane at equator, x 500. ith —28. P. pyrifolia (H.B.K.) Eichler, ile enced collected Kuijt, focal plane just below the apocolpial exine, x 500. —27. P. platyclada Ule. Light m micrograph w Costa Rica. Light micrograph with focal plane just below the apocolpial exine, x 500.— D. purpureus g & Urban, Liogier 15253 (F), Hispaniola. Polar view, x 1,330. P. pyrifolia (H.B.K.) Eichler, Greenhouse-collected Kuijt, Costa Rica. Polar view, x 2,080.— — 30. P. platyclada Ule. aq de view, x 2,000.— 32. D. constantiae 1985] FEUER & KUIJT— MISTLETOE POLLEN 193 Krug & Urban. Equatorial view, x1,400.—33. P. platyclada Ule. Equatorial view, x 2,000.— 34. P. pyrifolia (H.B.K.) Eichler, Greenhouse-collected Kuijt, Costa Rica. Equatorial view, x2,100.— 35. P. pyrifolia (H.B.K.) Eichler, Greenhouse-collected Kuijt, Costa Rica. Detail of aperture. Note the symmetrical break in the aperture membrane, x 6,000. 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FicunES 36-41. Ultrastructural comparisons between Dendropemon Blume (36-40) and Phthirusa Martius (41) (TEM).—36 ensis Britton. Detail of lobar exine. The ektexine (ek) is organized into irregularly ped pe t structures connected by thin strands of endexine (en), x9, bahamensis Britton Detail of polar exine, ; D. pycnophyllu g an. Detail la e exhibiting well- dev d endexine, x9,75 mensis Britton. Section through polar thickening (pt djacent lobe ne x2 D. bahamensis Britton. Detail o eee) al exine revealing columellate/baculate inter- S tratu . P. platyclada Ule. Ektexine ed of sporadic, itium and n m o comp o irregularly shaped ee and irregularly thickened roe layer, x 11,000. All lines in micrographs equal 1 um 1985] polar faces, thinning in lobar regions present here as thin strands connecting lower surfaces of ir- regularly shaped ektexinous segments, typically absent in interapertural equatorial areas, but in some species (e.g., D. bahamensis) present as a thin continuous stratum. Tectum/Equatorial interstitium//Foot layer: 1:3 to 2:1 Species and specimens examined: Dendrope- mon bahamensis Britton, *Popenoe 5 (A), Ba- hama Is.; D. caribaeus Krug & Urban, Britton et al. 45 (MO), West Indies (St. Thomas), Duss 2419 (GH), West Indies (Guadeloupe); D. con- stantiae Krug & Urban, Cicero et al. 6040 MOL Hispaniola; D. emarginatus Steud., Correll 44095 (MO), Bahama Is., Correll & Correll 47922 (F), Bahama Is. (Great Exuma), Smith et al. 3345 (F), Cuba; D. pauciflora Tieghem, Maxon 8761 (GH), Jamaica; D. picardiae Krug & Meis Ek- man H12947 (GH), Hispaniola; D. purpureus Krug & Urban, Liogier 15253 (F), Hoe Wilson 7647 (F), Bahama Is. (Caicos Is.); D. pyc- nophyllus Krug & Urban, *Liogier 17966 (F), Hispaniola; D. rostratus Urban, Ekman H 10199 (GH), Hispaniola; D. uniflorus Steud., Leonard & Leonard 12470 (MO), Haiti (Tortue Is.), Leon- ard et al. 15026 (UC), Haiti. Ixocactus Rizz. (Figs. 98-104). Exomorphology: radial symmetry, isopolar. Shape: amb circular; oblate, suboblate rarely ob- late spheroidal. Apertures: colpi, varying in number within and among populations from (3 to) 4 to 5 colpate. Sculpturing: densely clustered blunt-tipped spines and more irregularly shaped often bifurcating segments. Exine thickest in in- terapertural equatorial regions thinning slightly near apertures End 1 g texi d endexine pres- ent. Non-apertural ektexine e two types of organization on same grain: Type I with bi- stratified ektexine; upper stratum composed of variably thick, solid, broad, frequently irregu- larly branching segments, the branches some- times elaborated into blunt-tipped spines; basal stratum solid, variably thick with granular upper and slightly scalloped lower surfaces; both strata separated by extremely narrow, sparsely granular interstitium; and Type II, composed of a basi- cally single stratum of ektexine similar in struc- ture to Type I resting on small, highly channeled, loosely aggregated ektexinous segments random- ly attached to lower surface of upper ektexine. FEUER & KUIJT—MISTLETOE POLLEN 195 Endexine, beneath Type I ektexine, solid, con- tinuous irregularly thickened but, beneath Type II, organized into loosely paii small isolated segments attached to e ERR e E layer: 4:1 (Type I ektexine). Species and specimens examined: /xocactus hutchisonii Kuijt, Cuatrecasas 23970 (F), Co- lombia, *Hutchison & Idrobo 3008 (F), Colom- bia, Steyermark et al. 111592 (F), Venezuela, Tamayo 2526 (ILL), Venezuela. Maracanthus Kuijt (Figs. 4, 50, 53, 67). Exomorphology: radial symmetry, isopolar. Shape: amb triangular to slightly convex; oblate. Apertures: syncolpi; colpi narrow, shallow, slightly broader on the polar faces, narrowing at the equator. Sculpturing: short ridges, low profile striato-rugulae, exine foveolate and perforate, these most common in interapertural equatorial areas; polar faces psilate, imperforate. Exine thickened in a circular configuration directly at the: center of the polar faces. and endexine pres- ent. Slight structural differences between equa- torialand polar ektexine. Equatorial ektexine or- d into m oot thickened, foveolate, perforate, and channeled, the latter thin and not continuous through tec- tum. Interstitium clearly defined, composed of irregularly thickened columellae interspersed among baculae and baculae-like structures, the latter pendent from the tectum and surrounded by a finely granular matrix. Foot layer discon- tinuous in small localized areas, extremely thick, with a smooth upper surface and sporadically scalloped lower edge. Polar ektexine thinner than equatorial ektexine, organized into a thick, rarely perforate or foveolate tectum and poorly delim- ited, predominantly granular interstitium exhib- iting a few short, irregularly t ex columel- areas, irregularly thickened, bun lower edge. Endexine present at the polar faces and beneath apertures but typically absent in equatorial areas though sometimes filling the small gaps in lower edge of foot layer. Tectum/Equatorial interstitium//Foot layer: :2 Species and specimens examined: Maracan- hus chlamydatus (Rizz.) Kuijt, *Steyermark 99522 (UC), Venezuela. 196 Oryctanthus Eichler (Figs. 42-49). Exomorphology: radial symmetry, isopolar. Shape: amb circular; polar face characterized by three circular depressions each bounded by a nar- row, raised rounded ridge, the ridges confluent at the center of the polar face forming a triradiate configuration; equatorial ridge also present; ob- late. Grains compound diploaperturate. Aper- tures: ectoaperture a short, elliptical colpus; en- doaperture a sagittate opening; apertures placed 120? apart at the tips ofthe triradiate central ridge and between the ridges which encircle each of the polar depressions. Sculpturing: psilate, im- perforate with loosely granular exine occasion- ally present in depressed circular polar areas. Ex- ine thickened in equatorial areas aligned with apertures and in the center of each of the three DIT pour wd DES d endexine pres- ent. Ektexine typically bistratified, organized into upper and lower solid strata separated by a nar- row, highly undulate, osmiophilic layer; strata connected by numerous, sporadic, extremely short strands of ektexine; equatorial ridges formed by the outward looping and subsequent appres- sion or fusion of the bistratified ektexine; ektex- inous strata at the base of the polar depressions separated by a broad, coarsely granular zone, the upper stratum giving rise to a thin network of fine, randomly anastomosing granules. Endexine not continuous around grain, absent beneath po- lar depressions but prominent and irregularly thickened directly beneath equatorial ridges, thin and locally discontinuous in nondepressed polar areas; apertural endexine thin, loosely granular. Tectum/Equatorial interstitium//Foot layer: :1 Species and specimens examined: Oryctanthus - Broadway 681 (GH), French Guiana, Wiens 3741 (UT), Ecuador; O. occidentalis (L.) Eichler, Kuijt ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 2440 (UBC), Costa Rica; O. phanerolomus (Standley) Kuijt, Gentle 1405 (GH), British Hon- duras; O. spicatus (Jacq.) Eichler, *Killip & Smith 16807 (GH), Colombia. Oryctina Tieghem (Fig. 17). Exomorphology: radial symmetry, isopolar. Shape: amb triangular slightly convex; oblate. Apertures: diplosyndemicolpi; colpal mem- branes slightly disrupted on polar faces ap- exine at equator. Sculpturing: psilate, foveolate and perforate(?). Exine evenly thickened at equator, slightly thick- er along colpal margins at centers of polar faces. Endomorphology: not examined. Species and specimens examined: Oryctina subaphylla Rizz., Anderson, Stieber & Kirkbride 36949 (RB), Brazil Phthirusa Martius (Figs. 1—3, 27, 28, 30, 31, 33- 35, 41, 51, 52, 54-56). TYPE I Exomorphology: similar to Dendropemon spp. in Phthirusa species differing by smaller grain E DUIS similar to that of Dendro- pemon spp., particularly D. bahamensis. Tectum/Equatorial interstitium//Foot layer: 1:1.66 Species and specimens examined: Phthirusa pyrifolia (H.B.K.) Eichler, Croat 18038 (LEA), Peru, Revilla 2173 (MO), Peru, *Greenhouse (collected Kuijt), Costa Rica; P. platyclada Ule, * Asplund 12376 (S), Peru. TYPE II Exomorphology: radial symmetry, iso- and/or heteropolar. Shape: amb triangular to slightly concave; oblate to suboblate. Differences in po- larity, aperture types and sculpturing distinguish the following subtypes: Subtype A. Grains isopolar. Apertures: dip- — FiGunEs 42-49. Oryctanthus Eichler. 42, 43. O. spicatus (Jacq.) Eichler. 44—49. O. cordifolius (Presl) Urban, Kuijt 2571 (UBC), Costa Rica.— 42. thickening revealing three sagittate apertures, x 500.—43 Light micrograph (LM) with focal plane at the level of the triradiate polar LM with focal plane at the equator clearly revealing central polar thickenings in each of the polar depressions (arrowhead), x 500. —44. Polar view. The sagittate apertures, though prominent in the LM, are barely discernible in the SEM, x1,340.— 45. Oblique equatorial view revealing equatorial ridge and raised polar thickening, x 1,535.— 46. Thin section perpendicular to polar 1985] FEUER & KUIJT—MISTLETOE POLLEN 197 face through a polar ip gone including its geme d (pt), an adjacent aperture (ap), and equatorial nee (ear), x p 000. — 47. Detail of polar exine. Narrow bands of ektexine ect the ektexinous strata, 14,000. . Detail of sya thickening which lies at the center of each of the polar depressions. In some species a ü : ular matrix is attached to the tectum, x8,700.— 49. Detail of equatorial ridge adjacent to e nu polar Fuad, revealing looped appressed ektexine and thickened endexine (en), x 8,700. All lines in micro- graphs equal 1 um. 198 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Ficures 50-58. Comparisons between Maracanthus Kuijt and Phthirusa Martius (SEM). 50, 53. Maracan- Je chlamydatus (Rizz.) Kuijt. 51, 52, 54-58 Phthirusa spp.— 50. ith. Polar view of diplosyndemicolpate grain (Type II-A), x1,330.— . P. coarctata A. C. Sm Polar view of syncolpate grain, x1,250.— 52. P. o Eu Polar view of 3- diplodemibrevicolpate grain (Type II-B3), x1,250.—53. Oblique prs s view 500.— 55. x1,250.—54. P. micrantha Eichler. magdalenae (Cham. & Schldl.) Eichler. Polar view show — 56. P. ovata (Pohl) Eichler. Polar view of diplosyndemicolpate grain Se II- W i pe II-C1), x 1,250. x1,2 Polar view of neis) ip grain (Type II-A), x1 ing protruding colpal margin of di plosyndemicolpate P. theloneura Eichler. Diplosyndemicolpate grain (Type II-C2). Polar view, x1,2 200. vun B Equatorial view, x1, losyndemicolpi; colpi terminating subequatori- ally, broadening near equator with membranes frequently disrupted. Sculpturing: psilate, exine perforate and foveolate, these grandom- ly, tending to be more numerous in equatorial than polar areas. Species and specimens examined: Phthirusa coarctata A. C. Smith, Smith 2204 (U), British uiana; P. micrantha Eichler, Carreira 31 (RB), ail CS 1782 (K), Brazil; P. monetaria Sandw., Forest Dept. 82942 (K), British Guiana; P. sandwithii Maguire, Sandwith 1404 (K), Brit- ish Guiana Subtype B. Grains isopolar. Sculpturing: nu- merous randomly arranged short furrows and perforations frequently surrounded by low coarse ridges. Exine bordering apertures and apocol- pium (when present), psilate, imperforate. Dif- ferences in apertures distinguish the following species: 1985] Bl. Apertures: syncolpi Species and specimens examined: Phthirusa squamulosa Klotzsch ex Eichler, *Maguire & Stahl 24972 (D, Surinam. A pertu rec A 1 ] pi; colpi nar- TOW becoming lightly broader near equator. Species and sp ens examined: Phthirusa ovata (Pohi) Ei rep *Irwin et al. 7974 (UC), Brazil; P. rufa var. tentaculifera Rizz., Ducke 26 (RB), Brazil, *Ducke 1919 (K), Brazil. . Apertures: 3-diplobrevidemicolpi; colpi displaced entirely onto polar faces, forming large psilate, e apocolpia. pecies and specimens examined: Phthirusa enidobor Eichter. *Nichols s.n. (K), Jamaica. Subty, rains iso- and/or heteropolar wii respect to apertures, Isopolar grains diplo- gr ains with one po- lar face syndemicolpate, opposite face 3-demi- colpate, rarely 2-demicolpate, the latter through extrusion of a colpal margin and subsequent fu- sion with opposite colpal border. Sculpturing: ranging from ridges to striato-rugulae on same grain; ridges short, undulating, largely confined to midequatorial regions; striato-rugulae in low relief, largely confined to areas bordering psilate margins of colpi. Differences in polarity distin- guish the following species: Cl. Grains isopolar and heteropolar within populations. Species and specimens examined: Phthirusa lobaterae Ferrari, Steyermark et al. 120118 (MO), Venezuela; P. magdalenae (Cham. & Schldl.) Eichler, *Langenheim 3101 (UC), Colombia; P. retroflexa (Ruíz & Pavón) Kuijt, Gentry & Berry 14667 (MO), Venezuela, *Nee & Hansen 14038 (F), Panama. C2. Grains isopolar. Species and specimens examined: Phthirusa krukovii A. C. Smith, Krukoff 5938 (A), Brazil; P. retroflexa (Ruíz & Pavón) Kuijt, *A/len 80 (MO), Colombia; P. theloneura Eichler, *Spruce s.n. (K), Brazil. 3. Grains heteropolar. Species and specimens examined: Phthirusa angulata Krause, Maguire & Fanshawe s.n. (K), British Guiana; P. jamaicensis Krug & Urban, Purdie (K), Jamaica; P. stenophylla Eichler, Spruce 3307 (GH), Brazil. Endomorphology: exine organization similar to that of Maracanthus. Phthirusa species differ in the following characters: 1) tectum ranges from highly to rarely perforate; 2) a strictly columellate FEUER & KUIJT— MISTLETOE POLLEN 199 interstitium exists in several species where the columellae, though small and sporadic, are not associated with granular elements; and 3) con- tinuous foot layer. Tectum/Equatorial interstitium//Foot layer: 1:1.33. Species and specimens examined: all species listed under exomorphology that are marked by an asteris Struthanthus Martius (Figs. 12-16, 18-24, 70, 71, 74-86, 90-97). TYPE I Exomorphology: radial symmetry, grains iso- b triangular to u ranging from a small well-developed rectangular to elliptical slit to an obscurely margined area lying perpendicular to and in midregion of ec- tocolpus on the polar faces. Sculpturing: psilate- imperforate to perforate or rarely low profile striato-rugulae. Differences in T and po- larity distinguish the following subtype pe A. Endoapertures well e Em ped. Al. Grains isopolar; qiplosyndemicolporete, Species and specimens examine truthan- thus belizensis Lundell, *Lundell 6273 (SMU), British Honduras; S. densiflorus Standley, * Moore & Wood 3666 (UC), Mexico (Hidalgo); S. /ep- tostachyus (H.B.K.) G. Don, Mori & Kallunki 5308 (LEA), Panama; S. quercicola (Cham. & Schldl.) Blume, *Barlow 1429 (UT), Costa Rica, Chrysler 5031 (F), Costa Rica, Smith A519, 1479(F), Costa Rica; S. rotundatus Rizz., Lewis et al. 600 Santa Panama, Nee 10651, 14137 (MO), Panam A2 uw sss ‘(amulet with respect to ap- ertures; diploaperturate: one polar face parasyn- micolporate, opposite face either syndemicol- porate or 3-demicolporate. Species and specimens examined: Struthan- thus dichotrianthus Eichler, *Smith 1279 (MO), Colombia, Woytkowski 8269 (UC), Peru. A3. Grains isopolar; syncolporate. Species and specimens examined: Struthan- thus orbicularis (H.B.K.) Blume, Barlow 1507 (UT), Costa Rica, *Burger & Baker 10095 (F), Costa Rica; Williams 16049 (F), Costa Rica; Or- tiz 798 (UC), Guatemala. 200 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 A FiGures 59-67. Ultrastructural comparisons between PAthirusa Martius M). [p opem moms (Rizz.) Kuijt. Fic 59- 62. Phthirusa ovata (Pohl) Eichler. 63-66. Phthirusa spp. 67. Section through en x 4,500. — 60. Section through polar thickening (pt) at the apocolpium and oppo equatorial exine (eq), x 3,200. — 61. Detail of equatorial exine revealing short, sparsely distributed E x11,250.— 62. Detail of apocolpial exine (ac), x6,000.— 63. P. theloneura Eichler. Detail of equatorial exine, 1985] Subtype B. Endoapertures poorly developed with often obscure margin Bl. Grains isopolar; diplosvudemiecinetan date. Species and specimens examined: Struthan- thus escuintlensis Lundell, Matuda 4185 (A), Mexico (Chiapas); S. /eptostachyus (H.B.K.) G. sS Mexico (Chiapas), Janzen s.n. acc. #303522 (UC), exico (Oaxaca). Grains heteropolar; diploaperturate; one polar face parasyndemicolporoidate, opposite face either syndemicolporoidate or demicolpo- roidate. Species and specimens examined: Struthan- thus marginatus (Desr.) Blume, *Barlow 1420 ), Costa Rica, Thorne & Lathrop 40173 (UT), Mexico (Chiapas). B Grains heteropolar; synaperturate, one polar face parasyncolporoidate, opposite face syncolporoidate. Species and specimens examined: Struthan thus deppeanus (Cham. & Schldl.) Blume, *Balls 4340 (A), Mexico (Veracruz). TYPE II Exomorphology: radial symmetry, grains iso- polar or heteropolar, rarely subisopolar. Shape: oblate, suboblate, rarely oblate spheroidal. Ap- ertures simple, diplo- or synaperturate, the colpi sometimes broader in midregions of polar faces narrowing at the equator. Sculpturing: psilate- pepon ate and/or foveolate, rarely low profile riato-rugulae. Differences in amb, aperture SE and patterns of exine thickening dis- tinguish the following species: Subtype A. Grains isopolar; triangular to rounded convex; syncolpate; exine evenly thick- ened around grain. Species and specimens examined: Struthan- thus costaricensis Standley, *Barlow 1402 (UC, UT), Costa Rica; S. interruptus (H.B.K.) Blume, Nagel 8026 (GH), Mexico (Morelos). Subtype B. Grains isopolar rarely subisopo- lar (S. cassythoides) the latter with one polar face FEUER & KUIJT— MISTLETOE POLLEN 201 less rounded convex than opposite face; trian- gular rounded convex; diploparasyndemicol- pate; exine evenly thickened around equator but thinning beneath colpi at periphery of polar faces. Species and rr dad examined: Struthan- th tandley, Gentle 1660 (UC), British Houde ; S. palmeri Kuijt, *LeSueur 1071 (F, GH) | Mexico (Chihuahua). Subtype C. Grains isopolar; triangular to concave; diplosyndemicolpate; exine thinning in midequatorial regions and beneath colpi in pe- ripheral regions of polar faces, thickening near apertures in equatorial areas and at centers of polar faces. cies and specimens examined: iig an- thus oerstedii ipee Barlow 1443 (UC, UT), Costa Rica, Howell 10243 (F), Costa E Utley et al. 2701 (F), Costa - S. panamensis (Rizz.) Barlow & Wiens, *Davidson 431 (MO, US), Pan- ama, Luteyn 3786 (F), Panama. L J. ] le d endexine pres- ent. Ektexine organized into tectum, intersti- tium, and foot layer. Tectum regularly thickened, rarely perforate. Interstitium narrow, poorly de- fined, coarsely panur Occetionally exhibiting irregularly mellae-like structures. Foot layer continuous, with an un- even upper surface and typically smooth lower surface showing an occasional small gap. Endex- ine present at polar faces and beneath apertures, rarely present in interapertural areas and then only filling gaps in lower edge of foot layer; polar endexine slightly thicker than apertural endex- ine, homogeneous. Tectum/Equatorial interstitium//Foot layer: zl Species and specimens examined: all species marked by an asterisk and exhibiting Types I and II exomorphologies, excluding S. panamen- sis. I J L ] Le d endexine pres- ent. Ektexine organized into tectum, intersti- tium, and foot layer. Tectum irregularly thick- — x11,000.— with apos Sa (long arr P. rufa var. tentaculifera Rizz., Ducke 1919 (K), Brazil. Detail of . exine, x 8,700.— . & Schldl.) euni Detail of equatorial exine, x 6,900. , Panama. Detail of equatorial exine, x ,200.— 67. Deta ow) and localized endexine is lower surface uallu ex ed. of Eu E ii arrow). Perforation (p), x 8,000. All lines in micrographs eq 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 quat : : usa ap ge grain. The prominent endoapertures are not visible in the SEM, x1,120.— 73. C. microphylla (H.B.K.) Kuijt (Type II). Inerapertura iini view, xl, 120.— 74. S. dichotrianthus Eichler, su dA 8269 (UC), Peru (Type I-A2). Polar grain. Opposite face of this grain is diplosyn S x] “120. L75. S. belizensis Lundell (Type I- AD. Polar face with ed discernible endoapertures, x1,400.— 76. s. marginatus (Desr.) Blume, Barlow 1420 (UC), Costa Rica (Type I-B2). Polar face s. J s s uku us heteropolar dus Opposite face 3- or rarely 2-demi- Ran pu. x], 120.— 7. S. margi inatus (Desr.) Blume, Barlow 1420 (UC), Costa Rica (Type I-B2). Equatorial at the equator, x1,120.— 78. S. dichotrianthus Eichler, Woytkowski 8269 UO, dr (Type I- KS gn face of 3-demicolporate heteropolar grain. Opposite face syndemicolporate, Rizz.) Barlow & Wiens, Davidson 431 eA Panama (Type II-C). Note the concave d. and M defined gs near the center of the polar face, x 1,12 1985] FEUER & KUIJT— MISTLETOE POLLEN 203 FiGuREs 80-86. Brazilian Struthanthus spp. (Type III).—80. S. uraguensis (Hook. f. & Arnell) G. Don, Hishbsch 17641 (UC), Brazil. Polar view, x 1,500.— 81. S. uraguensis (Hook. f. & Arnell) G. Don, Hatschbach 17641 (UC), Brazil. Equatorial view, x1,500.— 82. S. concinnus Martius. Polar view, x1,500.—83. S. vulgaris Martius. Polar view, x1,500.— 84. S. vulgaris Martius. Equatorial view, x1,500.—85. S. concinnus Martius. Equatorial view, x 1,500. —86. S. concinnus Martius. Detail of equatorial sculpturing revealing striato-rugulae, , ened, highly perforate. Interstitium well defined, ine continuous piss grain. Polar endexine ex- granular rarely exhibiting columellae or colu- tremely thick, up to eight times thicker than mellae-like structures. Footlayercontinuous, with equatorial Rod a dense, filled with osmio- smooth upper and scalloped lower edge. Endex- philic granules; equatorial endexine irregularly 204 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 FAR Do— Rar *AARGN Tu ws XS ` FIGURES 87-97. Ultrastructural comparisons between Cladocolea Tieghem and Struthanthus Martius (TEM). pinus C. harlingii Kuijt, Harling 6094 (S), Ecuador. 90—92. S. panamensis (Rizz.) Barlow & Wiens, Davidson 31 (MO), Panama. 93, 94. S. costaricensis Standley. 95-97. S. uraguensis (Hook. f. & Arnell) G. Don, Hatsch- bach 1 764 1 (UO, Brazil. —87. Thin section through equator, x 1,200. — 88. Section perpendicular to polar face x 2,800. — 89. Detail of equatorial exine revealing rarely perforate tectum, “granular | interstitium, and ‘thick foot layer, x 14,000.—90. Section perpendicular to pole 1985] thickened with small internal gaps often filled with osmiophilic granules thickest in midequa- torial regions, thinning near ektexine bordering apertures; apertural endexine loosely granular. Tectum/Equatorial interstitium//Foot layer: :2 Species and specimens examined: Struthan- thus panamensis (Rizz.) Barlow & Wiens TYPE III Exomorphology: radial symmetry, isopolar. Shape: amb variable ranging from triangular to triangular slightly concave and convex; oblate rarely suboblate. Apertures: syncolpi, rarely parasyncolpi, colpi typically broadening and dis- rupted near the equator; colpal margins often thickened at the center of the polar face. Sculp- turing: ranging from uniformly to nonuniformly prominent striato-rugulate sculpturing elements sometimes randomly coalescing in equatorial areas to form mounds or more elongate ridges; colpal margins psilate or more rarely exhibiting striato-rugulae. Exine usually evenly thickened at equator, but several species (e.g., S. flexicaulis, S. vulgaris) exhibiting thickened midequatorial exine hol kt d endexine pres- ent. Ektexine organized into tectum, granular in- terstitium, and foot layer. Tectum irregularly thickened, outer surface elaborated into small excrescences representing striae in cross-section, rarely perforate. Interstitium narrow, granular, surrounded by a finely granular matrix. Foot lay- er thick, continuous, upper surface ranging from smooth to granular, lower edge randomly scal- loped. Endexine homogeneous, absent in inter- apertural equatorial areas, present beneath ap- ertures and polar faces. Tectum/Equatorial interstitium//Foot layer: 1:2 to 2:1 Species and specimens examined: Struthan- thus concinnus Martius, Irwin 2250 (UC), Brazil; FEUER & KUIJT—MISTLETOE POLLEN 205 S. flexicaulis Martius, Irwin et al. 10771 (MO), 736 < £f g = eo 2 Q & M ° o Dv Ç, A joa Qo < 9 Ej S [65] "S ~ ç 7 ygopus Martius, Glaziou 2599, 4821 (NY), Bra- zil; S. salicifolius Martius, Pereira 897 (MO), Brazil; S. syringifolius Martius, Krukoff 562AA ab Brazil; S. uraguensis (Hook. f. & Arnell) G. Don, *Hatschbach 17641 (UC), Brazil, Smith & Reitz 12449 (MO), Brazil; S. uraguensis var. brevipedunculata Chodat & Hassler, Hassler 7531 (A), Paraguay; S. vulgaris Martius, *Hatschbach 16483 (UC), Brazil. DISCUSSION The following sections discuss pollen charac- ters among small-flowered neotropical Loran- thaceae and their bearing on a the interpretation ofintra- and i within the complex. e + SIZE Most small-flowered genera possess medium- sized grains which, because of their predomi- nantly oblate shape, exhibit a significantly longer equatorial than polar axis. The majority of largest size pollen (P: 25-30 um; E: 32-50 um) characterizes Dendropemon, Oryctanthus, and Ixocactus. At the opposite end of the range, the smallest size pollen occurs in Maracanthus and Oryctina (P: 14-20 um; E: 23- 30 um). Pollen size is variable among species of Phthi- rusa, Cladocolea, and Struthanthus. Both Cla- docolea and Struthanthus exhibit clusters of species whose pollen sizes significantly differ. In Cladocolea, pollen of C. harlingii (15 um by 25 um), for example, is significantly smaller than that of C. grahamii (28 um by 41 um). Similarly, pollen of Struthanthus leptostachyus (15 um by 30 um) is strikingly smaller than pollen of most Brazilian Struthanthus, particularly that of S. syringifolius (33 um by 47 um) (cf. Figs. 13, 22). — granular &llad d torial exine, x 2,500. —91. Detail of Au en revealing perforate tectum, well-defined interstitium, thick foot layer, and well-developed 1,500. endexine (en), x11, —92. T and ei equatorial endexine, x 2,000. x 1,500.—94. tail of stitium, = foot layer, x 11 ,500. —95. Section through equator, x] ,250.— equatorial exine organized into rarely perforate tectum, in section of portion of grain through equator showing granular filled apertures —93. Section through equator including colpal membranes (c), poorly defined, granular inter- . Section perpendicular to equator along po x1,250.—97. Detail of equatorial exine Missis ized into irregularly thickened tectum representing striae in cross-section (arrows), narrow, poorly defined interstitium and irreg ularly thickened undulate foot layer, x8,500. All lines in ea unless otherwise indicated, equal 1 um. Micrographs with um marking on photograph equal 5 & 206 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 EA v. XS: Ficures. 98-104. Ixocactus hutchisonii Kuijt. —98. Polar view (SEM), Tamayo 2526 (ILL), Venezuela, x 1,950.—99. Polar view of 4-aperturate grain (LM), Steyermark et al. 111592 (F), ates x 500.— 100. Polar view of 5-aperturate grain (LM), Steyermark et al. 111592 (F), Venezuela, x — 101. Detail of surface perture (lower right) (TEM). The point of demarcation between Types I and II exine organization is marked by the arrow, x 3,200. — 103. Detail of Type I exine organization showing solid sculptural-structural elements and a solid, variably thickened foot layer separated by a narrow, granular ; interstitium (in) (TEM), x8,000.— 104. Detail of Type II exine organization. Th oosely organized, discontinuous endexine (en) (TEM), x 8,000. All bud in micrographs equal | um. 1985] POLARITY Though all small-flowered genera are typified by isopolar pollen, several species of PAthirusa and Struthanthus exhibit heteropolar pollen which may be uniformly present within a pop- ulation or mixed with isopolar grains. Heteropolar pollen features are restricted pri- marily to differences in aperture types between the two polar faces. The most common hetero- polar aperture combination is associated with species of Struthanthus in which, among diplo- aperturate grains, one polar face i is parasyndem- icolpate while the ite f: either syndem- icolpate or, less frequen e: 3-demicolpate (cf. Figs. 74, 78). Rarely, one set of apertures is longer than those of the opposite face (e.g., Phthirusa angulata, Struthanthus cassythoides). SHAPE Oblate pollen characterizes the small-flowered complex. Though all genera, excluding the sub- oblate /xocactus, exhibit a mean oblate pollen shape, species of Cladocolea, Oryctina, and Oryctanthus are strictly oblate in contrast to those of Dendropemon, Phthirusa, and Struthanthus which also possess suboblate, and more rarely, oblate spheroidal grains. Pollen amb is more variable than equatorial shape ranging from trilobate extremely concave among Dendropemon and Phthirusa pro parte to circular in Oryctanthus and Ixocactus. Despite this wide variation, the amb of most small-flow- ered genera is triangular to triangular slightly convex or, less frequently, concave. APERTURES Small-flowered genera are predominantly sim- ple aperturate though particular species of C/a- docolea and Struthanthus do, however, exhibit small, rectangular to elliptical to U-shaped slits lying perpendicular to the colpus at the periphery of the polar thickening (Figs. 7-11). The simple apertures of small-flowered genera are predominantly diploaperturate, i.e., a set of apertures each restricted to a polar face and dis- continuous at the equator usually separated by a narrow bridge of exine. Three diploaperturate types prevail: 1) diploporate-diplobrevidemicol- pate; 2) diplosyndemicolpate; and 3) diplopara- syndemicolpate. The first type characterizes all Dendropemon and three Phthirusa species. Apertures in these taxa are connected by a subsurface triradiate po- FEUER & KUIJT— MISTLETOE POLLEN 207 lar thickening at each face (Figs. 25, 27, 28). The species of Cladocolea and Struthanthus. The third type is restricted to a few species of Cladocolea and Struthanthus. ough a diploaperturate condition predom- inates among the small-flowered genera, Phthi- rusa squamulosa, Maracanthus, and several species of Struthanthus and Cladocolea possess synaperturate grains. The compound apertures among species of Cladocolea and Struthanthus range from poorly defined colporoidate to well-defined prominent colporate and sagittate types pipi associated with pee idus arrangem The compound apertures ae ts inane are unique wun the family. When viewed in the scanning electron microscope they appear as nar- row, short, slightly opened slits to broad, short, open, elliptical colpi located at the tips of the raised triradiate polar thickening. Optical sec- tions just below the pollen surface reveal, how- ever, a Sagittate endoaperture (Figs. 42-45). The (3-)4—5 colpate apertures of /xocactus, which vary within and among populations (see Table 1), are unique within the family (Figs. 99, 100) SCULPTURING Small-flowered genera are basically uniformly sculptured though some species do show slight sculptural differences between polar and/or col- pal margins and equatorial areas. Most taxa exhibit only slightly sculptured sur- faces ranging from psilate-perforate and/or fo- veolate to shallowly ridged and/or striato-rugu- Cladocolea, Phthirus. Brazilian species of Phthirusa and Struthanthus, the exine is elaborated into pronounced striato- rugulae and ridges. In Struthanthus uraguensis, S. flexicaulis, S. vulgaris, and S. polyrhizus these striato-rugulae coalesce to form small rounde mounds with striate bases and psilate apices (Figs. 80, 81, 83, 84). The sculpturing of /xocactus, composed of blunt-tipped spines and rectangular to more ir- regularly shaped ramifying elements, is unique in the complex. EXINE STRUCTURE The following section details exine characters within the complex. The unique exine features 208 of Ixocactus are reviewed separately at the end of this section. The exine of all small-flowered genera (in- cluding /xocactus) is divisible into ektexine and endexine. Ektexine. The ektexine is basically of uni- form organization throughout the grain. Only Oryctanthus and to a lesser degree Dendropemon exhibit disparate polar and equatorial ektexine structures. The equatorial ektexine is composed of tectum, interstitium, and foot layer. The tectum is typically perforate ranging from rarely to highly perforate. Only Oryctanthus and a few species of Struthanthus exhibit an imper- forate tectum throughout the grain. Usually evenly thickened, the tectum is variably thick- ened in Struthanthus panamensis (Fig. 91) and those Species. of Phthirusa: and Struthanth us with (Figs. 95— 97). A granular or granular-columellate intersti- tium characterizes most genera. These columel- lae, ranging from narrow to broad, are typically irregularly shaped, often sporadic and clustered, and usually associated with a granular matrix (Figs. 63-65, 67, 89, 94). Strictly collumellate or columellate/baculate interstitia are rare, restrict- ed to a few species each of Dendropemon (Fig. 40) and Phthirusa (Fig. 61). These columellae and/or baculae, though the dominant structural tial organization is difficult to interpret. Tum sec- tions reveal a narrow traversed by extremely thin ektexinous strands (modified columellae?) (Fig. 47). All genera, excluding Maracanthus, exhibit a continuous foot layer. Varying at the species level, the foot layer ranges from evenly thickened to undulate and variably thickened, the latter most pronounced in Phthirusa platyclada and the Bra- zilian species of Struthanthus (Figs. 41, 97). The upper surface is either smooth and clearly de- fined or granular and poorly defined. The lower surface can be smooth, coarsely granular, and/ or broken by small gaps (Figs. 67, 2 In most genera, the foot layer ranges from as thick as or up to twice as thick as the tectum plus equatorial interstitium (1:1 to 1:2) with the re- verse ratio (2: 1) occurring among Brazilian Stru- thanthus where the tectum is prominently sculp- tured In most genera the polar ektexine reflects an ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 organization similar to that in equatorial areas. The former differs in the usually thicker imper- forate tectum and narrower less well-defined interstitium. Oryctanthus, with its strikingly dif- ferent polar and equatorial ektexine organiza- nee is unique within the complex. Here ick polar interstitium, filled with irregularly i ektexinous segments and numerous os- miophilic granules, is in sharp contrast to the equatorial interstitium represented by a narrow osmiophilic stratum traversed by thin ektexi- nous strands (cf. Figs. 47, 49). Ektexine organization in lobar and peripheral equatorial areas near apertures is represented by isolated, irregularly shaped segments pendent from a solid, imperforate tectum (Figs. 36, 39, In /xocactus, the organization ofthe upper ekt- exine into large, solid, irregularly shaped seg- ments and bifurcating spines as well as the ab- sence of an interstitium and basal ektexine in large non-apertural regions of the grain, isolate this genus from other members ofthe small-flow- ered complex. Endexine. Endexine is typically present in apertural, lobar, and polar regions but absent in interapertural equatorial areas. In most Stru- thanthus, Cladocolea, Phthirusa species and Maracanthus, polar endexine is thickest at the center of the polar faces, this reaching its greatest development in Struthanthus panamensis (Fig. 90 Endexine patterns in Oryctanthus, Dendrope- mon, and Phthirusa species with trilobate pollen differ from the majority of small-flowered gen- era. Polar endexine is typically absent but equa- torial endexine is present though thin, irregularly thickened, and locally discontinuous. In Oryc- tanthus, endexine is also present and promi- nently thickened beneath the equatorial ridge (Fig. 49). The absence of polar endexine in these taxa suggests that the triradiate polar thickening vis- ible in the light microscope is largely due then to development of ektexine rather than endexine, in contrast to those polar thickenings among C/a- docolea and Struthanthus species. The highly discontinuous granular endexine restricted to certain portions of the pollen wall in 7xocactus is unique within the complex. POLLEN TRENDS The following sections analyze pollen trends among the small-flowered genera. Pollen char- 1985] acters of the more primitive large-flowered neo- tropical genera are used for out-group compar- isons. Pollen features of /xocactus are discussed separately at the end of these sections. Polarity. Isopolarity is basic for the complex while heteropolarity represents the derived state. Heteropolarity, characterizing several species in Stru- thanthus and Phthirusa, has developed indepen- dently among small-flowered loranths. The re- cent evolutionary occurrence of heteropolarity is suggested by its frequent occurrence alongside isopolar grains within the same population, its characterization of species rather than genera, and a consistent association with aperture anom- alies. Shape. A trilobate amb is primitive for the omplex. Restricted to Dendropemon and two species of Phthirusa, this shape most closely ap- proaches that typical ofthe large-flowered genera and Loranthaceae as a whole. The rounded con- vex amb in Oryctina and particular species of Cladocolea and Struthanthus is derived. The tru- ly circular amb, found among a few species of Oryctanthus, is also a highly derived condition having arisen through modification of a rounded convex shape typical of most Oryctanthus species. Oblate pollen shape is basic for the complex. The presence of suboblate and, more rarely, ob- late spheroidal shapes among ves advanced 1 Vil dos Ulva EV lq do large-flowered loranths suggest their derived na- ture. Apertures. A diploaperturate arrangement of simple apertures is basic for the complex. The basic type consists of colpi typically fused at the poles (syn-) but narrowly discontinuous at the equator (diplodemi-) with smooth, narrow mem- branes sometimes disrupted and/or broader near the equator. Both the parasyn- colpal form and the synaperturate arrangement, largely peculiar to species of ~ = Struthanthus, rep- resent derived conditi The status of the dico dub MM colpate apertures is obscure. Their occurrence in Dendropemon and three Phthirusa species and their closer similarity to apertures of the large- flowered Psittacanthus [see for example P. di- latatus, P. peronopetalus (Feuer & Kuijt, 1979)] than to any small-flowered taxa suggest primi- tive, probably vestigial, character states. Though primitive, the pori-brevicolpi of the small-flow- ered taxa do show unique modifications: encir- FEUER & KUIJT — MISTLETOE POLLEN 209 clement of each aperture by thickened, often slightly raised exine and interconnection of all apertures by a narrow, clearly defined triradiate subsurface polar thickening. e compound apertures of Oryctanthus, Cla- docolea, and Struthanthus are judged to be “‘in- cipient" and recently derived both because of their often ill-defined shape and the absence of a compound aperture type among Loranthaceae to date (Feuer & Kuijt, 1978, 1979, 1980, work in progress). ME close 'palynological similamties between their elliptical to U-shaped Beni iE dii may have developed independently from those of Oryctanthus Sculpturing. Though the psilate-perforate sculpturing of Dendropemon-Phthirusa pro parte is primitive for the complex, the basic sculptur- ing Rn adores by: most small- flowered taxa nda/ar etria OI SU Id- arc low p to- rugulae. Two sculpturing trends are evident within the complex. The first is the development of p forate psilate exine. This has occurred once the morphologically remote Oryctanthus se again among particular Mexican and Central American Struthanthus and Cladocolea species. A second and opposite trend, restricted to Bra- zilian populations of Struthanthus, involves the elaboration of the exine into pronounced striato- rugulae which sometimes coalesce into short mounds in equatorial areas. Exine structure. Similar ektexine organiza- small-flowered genera. The equatorial ligni organization in n Oryctanthus may be an ind in the small-flowered complex but is common among large-flowered neotropical genera. The basic ektexine of most small-flowered taxa is organized into a thin, perforate tectum, gran- ular/columellate or columellate/granular inter- stitium and a thick, continuous foot layer typi- cally twice as thick as the tectum plus interstitium. The imperforate tectum in Oryctanthus and par- ticular species of Struthanthus and Cladocolea with- overall loss of columellae. The extremely narrow 210 interstitial strands in Oryctanthus would then represent highly modified, though vestigial col- umellae. Endexine patterns of most small-flowered gen- era are strikingly similar to those of large-flow- ered genera. Both groups exhibit triradiately con- figured, thickened, often stratified polar endexine but no interapertural equatorial endexine. The thickened polar endexine arranged in a circular configuration in particular species of Cladocolea and Struthanthus is unique to the flowered com- plex and represents a derived condition. INTRAGENERIC RELATIONSHIPS Only PAthirusa, Cladocolea, and Struthanthus exhibit significant specific pollen variation to permit a discussion of intrageneric relationships; of these, only Cladocolea has been monographed (Kuijt, 1975). Phthirusa. Pollen characters divide the genus into two major groups: Group I composed of P. platyclada and P. pyrifolia and Group II con- taining all remaining species. Group I species are characterized by trilobate pollen with diplopor- ate apertures, midequatorial and triradiate polar thickenings, and a columellate/baculate exine structure. In contrast, pollen of Group II taxa, excluding P. squamulosa, are diplosyndemicol- pate with evenly thickened midequatorial and columellate/granular exine. Phthirusa lepidobo- trys, with its unique diplobrevidemicolpate pol- len, midequatorial and triradiately thickened po- lar exine ar shape a Ss e€ e e o = z rn e e 3 -= - a ° = ge = = u a mediate position between the two groups. Cladocolea. Pollen characters indicate a par- ticularly close relationship among C. andrieuxii, C. loniceroides, and C. microphylla, the latter two closer to each other than either is to C. an- drieuxii. These species exhibit the only endoap- ertures in the genus; the latter two additionally share a rounded convex amb and a triangular polar thickening along colpal margins. The nat- uralness of this group, as suggested by the pollen membranes in C. inconspicua suggest an isolated position within the genus. However, pollen of the closely related C. inorna and C. clandestina was unavailable for study. Cladocolea oligantha also occupies an isolated ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 position with its triangular pollen shape, thick exine, and sharply defined circular central polar thickening. Morphologically, this species is dis- tinguished by dimorphic inflorescences—a fea- ture present in only one other Cladocolea species. Struthanthus. Two major groups are delim- ited by pollen characters: Group I containing Brazilian species populations and Group II com- posed of remaining non-Brazilian species pop- ulations which range from Mexico to South America. This geographic distinction refers largely to populations of species since the Stru- thanthus taxa which occur in several regions (e.g., S. marginatus, S. orbicularis, S. rotundatus) were not examined throughout their total range. Pollen data suggest that the Brazilian species populations are particularly closely interrelated. Pollen is uniformly isopolar, syncolpate with pronounced striato-rugulate sculp t verse. Both iso- and heteropolar grains occur, which can exhibit either simple or compound apertures arranged in either syn- or diploaper- turate arrangements. Sculpturing ranges from typically tectate-perforate to low profile striato- rugulate to rarely psilate-imperforate. The unique occurrence of highly sculptured exines among Group I species and the restriction of heteropolar pollen, compound apertures, and psilate-imper- forate features to Group II species suggest two independently evolving complexes. Among Group II species, S. panamensis is dis- tinguished by its trilobate, slightly concave pol- len, thinning midequatorial exine and the pres- ence of a well-defined interstitium and midequatorial endexine, features closer to those distinguished from all other S inuthanihus species y its bisexual flowers arranged in monads as opposed to unisexual triads in all other species. Such characters have led to the proposal that S. L i y represent an inde[ branch leading into the Struthanthus pool (Kuijt, 198 1a). INTERGENERIC RELATIONSHIPS Excluding the disparate pollen characters of Ixocactus, the pollen data suggest two basic groupings of small-flowered genera: a first group composed of Dendropemon, Phthirusa pro parte, and Oryctanthus and a second generic cluster 1985] containing Phthirusa pro parte, Cladocolea, Struthanthus, Maracanthus, and Oryctina. Such an arrangement does not, oe preclude weaker intergroup relationship Within Group I, Daye is closely linked to Phthirusa through the species P. pyrifolia and P. platyclada. All exhibit trilobate, extremely concave pollen amb, diploporate apertures, and a midequatorially thickened columellate/bacu- late exine structure. PAthirusa lepidobotrys, with its diplobrevicolpate midequatorially thickened but triangular pollen links the Dendropemon-like Phthirusa species to the remaining 28 PAthirusa species, the pollen of which is strikingly similar to particular Struthanthus species. Pollen characters support a relationship be- tween Oryctanthus and Dendropemon-Phthirusa pro parte as suggested by floral and inflorescence structures (Kuijt, 1976a) but offer no evidence for an affinity with Maracanthus (Kuijt, 1976b). Oryctanthus pollen, though quite distinct, does possess several features common to Dendrope- mon-Phthirusa pro parte pollen: a narrow, well- defined triradiate polar thickening, small diplo- apertures restricted to the tips of this triradiate thickening, and an equatorial ektexine structure similar to the polar e t f Den- dropemon. Orvedtihiüs however, exhibits sev- eral uniquely derived pollen features: circular amb, raised polar ridges encircling polar depres- sions, compound apertures composed of sagit- tate endoapertures, and an equatorial intersti- tium traversed by numerous thin strands. Such unique characters suggest that Oryctanthus is a highly derived genus with only remote ties to the Dendropemon-Phthirusa pro parte complex. ic cluster, pollen char- acters indicate a relationship between Struthan- thus and Phthirusa on the one hand and Stru- thanthus and Cladocolea on the other Struthanthus is linked to Phthirusa through pollen characters of heteropolarity, oblate-sphe- roidal shape, apertures with protruding colpal margins, and pronounced striato-rugulate sculp- turing. Phthirusa species remain palynologically distinct from Struthanthus through consistent triangular amb and a well-defined columellate- granular interstitium —the latter closely resem- bling a Dendropemon feature. Pollen suggests an even closer relationship be- tween the Mexican-Central American Struthan- thus species and Cladocolea, particularly species with Types II and III pollen. The Struthanthus- Cladocolea species constellation similarly ex- ong the second g FEUER & KUIJT—MISTLETOE POLLEN 211 hibits a triangular rounded convex amb, simple and compound apertures, syn- and parasyncolpi arranged in either diplo- or synaperturate ar- rangements, and a granular poorly defined in- terstitium. The parasyncolpi, the elliptical to U-shaped endoapertures, and granular intersti- tium are features unique to this group. The strong ties suggested by the pollen data are corroborated by gross morphology. The unique occurrence of geniculate styles among particular Cladocolea and Mexican Struthanthus evinces close interspecific relationships. Striking morphological similari- ties between certain intergeneric species pairs (C. harlingii, S. orbicularis; C. lenticellata, S. poly- stachyus; C. pedicellata, S. deppeanus) have been used to purport a polyphyletic origin for Stru- thanthus (Kuijt, 1981a). Since pollen of several key Cladocolea species was not examined and as Struthanthus pollen in each of the species pairs is not strikingly different from other Struthan- thus nor closer to particular Cladocolea species, the data do IL to support or deny polyphylet- ism in the The prop dinde affinity of Maracanthus and Oryctanthus (Kuijt, 1976a) is unsupported by the pollen data. Rather, the following pollen char- acters indicate an intermediate position between Struthanthus and Phthirusa: triangular to round- ed convex amb, syncolpate apertures, perforate tectum with low profile striato-rugulae, circular polar thickening, and a well-developed colu- mellate/granular interstitium Little pollen evidence exists ios the suggested relationship between Oryctina and Maracanthus (Kuijt, 1981b) though these two genera are closer psilate pollen with thickened apocolpial colp " margins resembles pollen of Struthanthus species, particularly S. leptostachyus and S. palmeri. The pollen characters of /xocactus indicate no relationship with any small-flowered genus. The intra- and interpopulational aperture variation, the irregularly spinulose sculpturing, and ektex- ine structure characteristic of 7xocactus are clos- African species, than to any Loranthaceae sensu stricto. LITERATURE CITED BARLow, B. A. & D. Wiens. 1971. Thecytogeography of loranthaceous mistletoes. Taxon 20: 291-312. FEUER, S. & J. Kuyt. 1978. Fine structure of mis- 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 tletoe pollen I. ws eir Lepidoceras, and Plants. bd of California Press, Berkeley and Tupeia. Canad. J. Bot. 56: 2853-2864. Los Angele & 1979. Pollen evolution in the genus . 1975. The genus Cladocolea (Loranthaceae). Psittacanthus Mart. Fine structure of mistletoe J. Arnold Arbor. 56: 265-335. pollen II. Bot. Not. 132: 295-309. —— ———-. 1976a. Revision of the genus Oryctanthus . Fine structure of mistletoe (Loranthaceae). Bot. Jahrb. Syst. 95: 478—534. pollen. III. Large-flow ered neotropical Lorantha- | ———. 1976b. Maracanthus, a new genus of Lo- ceae and their Australian relatives. Amer. J. Bot. ranthaceae. Brittonia 28: 231—238. : 34-50. — ——-. 1981a. Inflorescence morphology of Loran- ar J. 1967. The genus /xocactus (Loranthaceae e em an evolutionary synthesis. Blumea 27: .S.): description of its first species. Brittonia 19: -7 š A rejoinder on Oryctina (Lorantha- ——, 1969. The Biology of Parasitic Flowering ceae). Pl. Syst. Evol. 137: 215-219. THE SYSTEMATICS OF THE APETALOUS FUCHSIAS OF SOUTH AMERICA, FUCHSIA SECT. HEMSLEYELLA (ONAGRACEAE)! PAUL E. BERRY? ABSTRACT 4: . a 4 A systematic revision of Fuchsia sect. Hemsleyella, a to the tropical Andes, is presented. Fourteen species are decidi die xb ares three new species. Members of this group are protogynous and are hummingbird pollinated; raid are generally rare and present little opportunity for sympatry or interspecific hybridization. The firs with 1 s that are opposite-ternate, persistent and elliptic-ovate to cordate and long-petioled pls om terrestrial to epiphytic or rock-inhabiting shrubs; - tuberous to tuberous plants; a - or medium-length floral tubes to ] tubes over 10 cm m a ra long. In its quia of oed sect. Hemsleyella is most similar to sect. Ellobium, from Mexico and Central America Within the highly distinctive Onagraceae, Fuchsia contains over 100 of the ca. 675 species in the family. It shows no obvious relationships to any of the other 16 genera and is unique in the family in its fleshly fruits and basically 2-aperturate pollen, being placed accordingly in its own tribe, Fuchsieae. The last generic mono- graph of the group was published by Munz in 1943. In recent years, an effort has been made towards a systematic reassessment of the entire genus, under the direction of Peter H. Raven; this has resulted so far in the revision of all the Mexican and Central American species of Fuch- sia (Breedlove, 1969; Breedlove et al., 1982), as well as the large South American sect. Fuchsia (Berry, 1982). Over three-quarters of the species of Fuchsia occur in South America. Among them, Hemsley (1876) first recognized the existence of a partic- ular group of species characterized by the ab- sence of petals, generally alternate leaves, and a mostly deciduous, epiphytic or saxicolous habit. ! This study is based upon research financed partially d. the U.S. National Science Foundation under b ne 821487 9t Pet av uar ére under their care: ; © MO, MPU VEN, W, ey 2 This group was not formally designated, how- ever, until Munz (1943) conferred upon it a sec- tional status, under the name Hemsleyella. Munz included 12 species and 13 different taxa in this section; one of these, F. decidua, is now placed in the recently described sect. E/lobium (Breed- love et al., Members of sec Hemsleyella are a difficult group to study because the plants are generally scarce, strongly seasonal, and occupy relatively inaccessible habitats. This section occurs sym- patrically with sect. Fuchsia over most of its range in the tropical Andes. While conducting field studies on me tatter group in pene 1980, the fsect posuer qn in i the field and to collect series of erbarium specimens and cytological samples. In addition, loans of sect. Hemsleyella from nu- merous institutions were examined. As a result, it was possible to make a substantial revision of Munz's (1943) treatment, including new ecolog- ical, cytological, palynological, and taxonomic elp preciated. I wish to thank the curators of the following herbaria for allowing me A, MER, , > s > > > , , MY, NA, NY, OXF, P, PH, POM, Q, RSA, S, TCD, TEX, U, UC, US, USM, partamento de Biología de Organismos, Universidad Simón Bolívar, Apartado 80659, Caracas 1080, Venezuela. ANN. MissounRni Bor. GARD. 72: 213-251. 1985. 214 information. The following revision now rec- ognizes 14 species in sect. Hemsleyella, the sec- ond largest in the genus; only six of these species are the same as ones recognized by Munz (1943), and three new species are described. GEOGRAPHICAL DISTRIBUTION AND ECOLOGY Distribution. Fuchsia sect. Hemsleyella is re- stricted to the moist slopes of the tropical Andes, from ca. 10°N to 17°S latitude. Its range coincides very closely with that of the larger sect. Fuchsia (see Berry, 1982, fig. 1), and therefore it is useful to discuss the distribution of the different species in light of the major structural units of the trop- ical Andes recognized by Berry (1982, fig. 12). Colombia, and Ecuador; two of these occupy the northern extreme of the Cordillera Oriental in Venezuela, whereas the other two occur in Ec- uador. Curiously, there is no sure record of sect. Hemsleyella from Colombia. In the Central An- es, which include Peru and Bolivia, ten different species occur. As in sect. Fuchsia, a single species is found along the relatively dry Pacific slopes of the Cordillera Occidental in northern Peru; one species is restricted to the Cordillera Central in Peru, two occur in both the Cordillera Central and Cordillera Oriental, while six different species are restricted to the Cordillera Oriental in south- . ern Peru and Bolivia The greatest local concentrations of species in sect. Hemsleyella occur in the moist valleys of Dept. Cuzco, Peru, and in the yungas of Dept. Cochabamba, Bolivia, with five species present in each area The most specialized species in the section oc- cur in the Central Andes, where the most prom- inent tuber development, longest floral tubes, and highly seasonal species occur; unspecialized species are found in both the Northern and Cen- tral Andes. Habitat and phenology. Fuchsia cestroides inhabits montane scrub vegetation in nort western Peru and occupies the lowest altitudinal range in the section (1,100-1,700 m). All other species in the section occur in Andean cloud for- est above 1,600 m, and some extend into the lower reaches of the open puna or subpáramo. Fuchsia apetala is most common at the upper cloud forest-puna limit and is known to reach 4,200 m in Peru, where it is subject to frequent frosts. In Venezuela, F. membranacea extends ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 from 2,600 to 3,400 m at the upper limit of cloud forest and the lower reaches of open páramo. Broad altitudinal ranges of more than 1,000 m are common in species of sect. Hemsleyella, by virtue of their versatile habit; plants are gen- erally terrestrial towards their upper altitudinal limit and are often epiphytic towards the lower limit of their range. The species of sect. Hemsleyella are among the most strongly seasonal in the genus. All species flower primarily in the dry season, when many are totally deciduous. This results in a limited flowering season, unlike the members of sect. Fuchsia, which are far less seasonal and flower over a more extended period, even though they inhabit the same areas as species of sect. Hem- sleyella. Strong vegetative growth occurs during the rainy season, and the presence of tubers or thickened stems in species of sect. Hemsleyella enables food and water to be stored for the dry season flowering. Due in part to the infrequent occurrence of plants belonging to sect. Hernsleyella in their na- tive habitat, their often epiphytic or scandent habit, and their adaptation to a short dry season and extended moist season, members of this group are particularly intolerant of habitat dis- turbance, and the survival of many species with limited geographical ranges is being endangered by the rapid advance of forest clearing along the steep Andean slopes. FLORAL BIOLOGY AND POLLINATION In details of floral biology, sect. Hemsleyella is similar to the situation described by Berry (1982) for sect. Fuchsia: the flowers are strongly protogynous (Fig. 1), yet they last several days, and self-pollination can occur before stigma re- ceptivity ends. The stigmas are generally exserted beyond the anthers, but since the flowers are pen- dant and the pollen contains viscin threads, pol- len grains can in some cases reach the stigma of the same flower by wind or gravity. The species sect. Hemsleyella are hermaphroditic, and self- incompatibility is not known for any of the pecies. All species of sect. Hemsleyella are character- ized by the presence of reddish (pink, violet, or orange), odorless, hanging flowers, all with floral tubes from 18 to 160 mm long and copious nectar production from a large nectary situated at the base of the floral tube and followed by a con- striction just beyond it. With clusters of flowers [^4] 1985] flowering simultaneously on different branches, those species offer a striking floral display, and all species are visited by hummingbirds. On le- gitimate visits, the hummingbirds approach the tions of pollinators were possible, but an emerald hummingbird (Amazilia sp.) was seen pollinat- ing a population of F. tillettiana (i ding Berry 4289) sporadically from daybreak to dusk, at 1,600 m in Edo. Mérida, Venezuela, in April 1984. Three species of sect. Hemsleyella, F. apetala, F. garleppiana, and F. inflata, often have floral tubes over 10 cm long. The only hummingbird with a bill long enough to reach the nectar res- ervoir without piercing the tube is the swordbill, Ensifera ensifera. This remarkable humming- bird has a bill almost as long as its body, often m, Dept. Cuzco, Peru; since the flowers of this species hang down off leafless and erect or divergent shoots and often from rock overhangs, it is easy for a swordbill to approach from below and insert its bill in the tube. Because of the correspondence in bill and tube length, the bird makes contact first with its ead to the stigma, then with the anthers. Indi- viduals of Ensifera were also seen visiting short- er-tubed flowers of F. chloroloba, but their bod- ies did not contact the anthers In populations of long-tubed plants of F. in- flata examined in August 1979, on the road from the Abra de Málaga towards Quillabamba (Prov. La Convención, Dept. Cuzco, Peru), the majority of the flowers had been pierced at the base of the floral tube by nectar robbers, as evidenced in specimens such as Berry & Aronson 2567 and CYTOLOGY Fuchsia has retained the original basic chro- of the other tribes in the family, aneuploidy and translocation systems are unknown in the Fuch- sieae. Kurabayashi et al. (1962) considered the chromosomes of Fuchsia to be large and unspe- cialized for the family, with poor differentiation into heterochromatic and euchromatic portions. BERRY — FUCHSIA SECT. HEMSLEYELLA FIGURE 1. Portion of a flower. of Fuchsia inflata Dies & Aronson 301 ) fanthesis, ing the sticky pti stigma and idoneo an- ber (protogyny). Polyploidy does occur in Fuchsia and has proven to be somewhat useful in characterizing the dif- ferent sections of the genus. All species of sects. Ellobium, Schufia, Jimenezia, Skinnera, and Encliandra have been counted and were found to be diploid, except for one individual of sect. Encliandra (Breedlove, 1969; Breedlove et al., 1982; Raven, unpubl. data). Berry (1982) found that sect. Fuchsia is predominantly diploid; 37 species examined were diploid, five were tetra- ploid, and one species had both diploid and tet- raploid populations. The monotypic sect. Kier- schlegeria is tetraploid, and sect. Quelusia is entirely polyploid, with three known tetraploid and one octoploid species (Berry & Ramamoor- thy, unpubl. data Prior to this study, no chromosome counts had been published for sect. Hemsleyella. Twenty- two new counts are reported here for the first time in the section, as listed in Table 1, including nine of the 14 species and two possible hybrids. The methods used were the same as those de- scribed in Berry (1982). Diploid chromosome 216 TABLE 1. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Chromosome numbers in Fuchsia sect. Hemsleyella. Taxon Collection Data F. apetala Ruiz & Pa- vón F. aff. apetala Ruiz & Pavón F. cestroides Schulze- F. chloroloba I. M. Johnston F. garleppiana Kuntze & Wittmack F. inflata Schulze-Menz F. x inflata (probable hybrid) F. juntasensis O. Kuntze F. membranacea Hem- sley F. tillettiana Munz F. tunariensis O. Kuntze ca. 44 44 PERU. cuzco: Tres Cruces, Prov. Paucartambo, Berry et al. 2593 (MO). PERU. cuzco: below Abra de Málaga, Cuzco-Quillabamba Road, Prov. Convención, Berry & Aronson 3030 (MO). PERU. JUNÍN: 32 km E of Comas, on road to Satipo, Berry & Aronson 3069 (MO). PERU. cuzco: 91 km from Ocongate, above Marcapata, Ber- ry & Aronson 3014 (MO) PERU. CAJAMARCA: 6 km above San Benito towards Guz- mango, Berry et al. 3600 (liquid specimen, author’s col- lection) PERU. cuzco: above Buenos Aires, Acanaco to Pilcopata, Prov. Paucartambo, Berry et al. 2599 (MO). PERU. CUZCO: 92 km from Ocongate, above Marcapata, Ber- ry & Aronson 3015 (MO). CULTIVATED: Berkeley, California, Univ. California Botani- cal Garden accession N° 49.817. Originally from Peru, no voucher. BOLIVIA. COCHABAMBA: 41 m from Cochabamba on road to Chapare, Berry 3640 (live tubers sent for cultivation to MO; same site where Berry & Berry 2582 was collected). PERU. cuzco: Km 145-146 from Cuzco to Quillabamba, Berry & Aronson 3036 (MO). PERU. cuzco: Km 150 from Cuzco to Quillabamba, Berry & Aronson 3040 (MO). PERU. cuzco: 88 km from Ocongate above Marcapata, Ber- ry & Aronson 3010 (MO). PERU. cuzco: 88 km from Ocongate, above Marcapata, Ber- ry & Aronson 3011 (MO) BOLIVIA. COCHABAMBA: above ENDE pumping station at Corani, Berry 3638 (MO). UT MÉRIDA: 13 km above Timotes, Berry 3278 (MO, VEN). VENEZUELA. MÉRIDA: 12.5 km below Apartaderos, Berry 3465 (MO, VEN) oe LARA: Laguna La Blanquita, Parque Nacional acambú, Berry 3259 (MO, VEN). VENEZUELA. LARA: Parque Nacional Yacambü, above San- are, Berry 3469 (MO, VEN). VENEZUELA. MÉRIDA: 1 km above San José towards Mucu- tuy, Berry 3463 (MO, VEN). VENEZUELA. TACHIRA: between Zumbador and Queniquea, Berry 3033 (MO, VEN) . AYACUCHO: 45 km E of Tambo, towards Calicanto, posa 3051 (MO). a Counted by Peter H. Raven. * Counted by Peter Goldblatt. * With bridges and fragments present at Anaphase I. 1985] numbers were obtained for seven species and tetraploid counts for two species and two puta- tive hybrids apparently with a somewhat higher proportion of tetraploid species. The two tetraploid species are not closely related to each other, but each is closely related to a different diploid species. In sect. Fuchsia, on the other hand, most of the tetraploid species were either geographically iso- lated or morphologically very distinct from the rest of the species in the section. SYMPATRY AND HYBRIDIZATION Because of the scarcity and relatively few col- lections of members of sect. Hemsleyella, little is known about the occurrence of interspecific id pene in this group. For some species, uch as F. cestroides, F. insignis, and F. pila- dod eis there is virtually no chance for sympatry, as they occur in areas well isolated from other species in the section. Other species, such as F. tillettiana and F. membranacea, or F. garlep- piana and F. juntasensis, grow in the same areas, but in different altitudinal ranges. Fuchsia nana and F. apetala generally occur close to timber- line, well above cloud forest occupied by other members of sect. Hemsleyella The greatest chance for sympatry and possible interspecific hybridization is found in two areas where many members of the section have been collected: several valleys of Dept. Cuzco, Peru, and the yungas of Dept. Cochabamba, Bolivia. Nonetheless, the only observed case of close plants in this area shówed a series of individuals with unusual morphological variability (Berry & Aronson 3010, 3011, 3012, 3013, and 3016), as well as tetraploid individuals with somewhat aberrant meiosis, providing some evidence that natural interspecific hybridization may be oc- curring between different members of the section there. Members of sect. Hemsleyella commonly oc- cur in the same locations as species belonging to sect. Fuchsia. With the marked morphological differences between those two sections, one would not expect to find intersectional hybrids. How- ever, one possible case of intersectional hybrid- ization has been detected in a series of Bolivian collections made by W. M. S. Brooke in August BERRY — FUCHSIA SECT. HEMSLEYELLA 217 FiGuRE2. Habit of Fuchsia inflata (Berry & Aron- son 3036) growing from tubers out of rocks and flow- ering on leafless stems, Prov. La Convención, Dept. Cuzco, Peru (Aug. 1978) 1950, at Incachaca, Dept. Cochabamba. The sets of Brooke 6666, 66 ferent conditions of sun or shade. Sheets of Brooke 6 , however, at BM, F, and NY, show some very different features. The specimen of Brooke 6666bat F (Herb. N° 1548145) was found “‘grow- ing in the open but in high grass,” according to the label; it has well-developed, opposite to ter- nate leaves present, whereas plants of F. junta- sensis found growing in the sun generally flower when leafless. The shape of the flowers on this sheet correspond roughly to that expected for F. juntasensis, but some flowers have petals pres- ent; some of the petals are deformed and others are more or less oblanceolate and up to 15 mm long. Other flowers from the same lack petals altogether, and all have malformed, non- functionalanthers. A band type nectary and con- nate sepal bases attest to the Hemsleyella par- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FiGURE 3. Epiphytic habit of Fuchsia inflata (Berry & Aronson 3038), several km below Berry & Aronson 3036 (Fig. 2). entage of this plant. The sheet of Brooke 6666b at NY is similar to that described above, only the locality is not the same: “under trees in a light forest, flowers, varying shades of mauve.” The third sheet at BM bears the same label in- formation as the NY sheet, but appears to be F. denticulata, which had been collected previously in the area. Most likely, Brooke 6666b 1s a series of naturally occurring hybrids between F. jun- tasensis and F. denticulata (sect. Fuchsia). MORPHOLOGICAL CHARACTERS All species of sect. Hemsleyella are lettiana, usually occur as upright to scandent shrubs 1-3 m tall, but F. tillettiana has also been found as a liana with long, flexuous branches clambering through woody vegetation on steep slopes. The closely related F. membranacea is generally terrestrial, but requires the aid of neigh- boring shrubs and trees to support its branches; it is also occasionally found as an epiphyte. Fuch- sia apetala, F. inflata, and F. juntasensis all oc- cur terrestrially near their upper altitudinal limits and epiphytically at lower elevations (Figs. 2, 3). Fuchsia pilaloensis grows either terrestrially or epiphytically at the same elevation in western cuador. more specialized species in the section inflata, and F. chloroloba, grow primarily among rocks and secondarily as epiphytes. Fuchsia gar- leppiana and F. inflata produce upright, slender stems when terrestrial and divergent to hanging branches when epiphytic (see Figs. 2, 3). Fuchsia apetala, on the other hand, is unique in having a prostrate, almost creeping habit, with highly congested, knotty stems when terrestrial; as an epiphyte at lower elevations, the stems are more elongate and produce slender stolon Leaves. Fundamental ee in leaf ar- rangement, shape, and persistence occur in dif- ferent groups of species in sect. Hemsleyella. One 1985] group, presumably the primitive condition in the genus and in this section, has leaves that are opposite to ternate, medium-sized (3-10 cm long by 2-5 cm wide), with short petioles (mostly 5- 25 mm long), elliptic to elliptic-ovate = shape, and slightly to moderately decidu econd, more advanced group in a) em- sibvelia is strongly seasonal, and the species tend to flower in the dry season when all the leaves have fallen. In some species, such as F. insignis and F. nana, the leaves are practically unknown. In cases where the leaves are known, however, such as F. apetala, F. chloroloba, and F. garlep- piana, they are alternate, large (5-14 cm long by 2-7 cm wide), long-petioled (15-270 mm long) and narrowly to broadly ovate in shape, with cordate bases and typically acuminate tips. Re- maining species, such as F. pilaloensis, F. sali- cifolia, and F. tunariensis, are intermediate be- tween the two groups describe Tuber development. Outside of sect. Hem- sleyella in Fuchsia, the presence of tubers is known only in two species of sect. Ellobium, which are principally rock-inhabiting or epi- BERRY — FUCHSIA SECT. HEMSLEYELLA à , a7 R P = ME = > É at POI SF i 1 i" e E xa ON / A4 "k ANS an^ FiGURE 4. Extensive tuber development in Fuchsia aff. inflata (Berry & Aronson 3010), a shrub growing among rocks in Prov. Quispicanchis, Dept. Cuzco, Peru. phytic. Likewise in sect. Hemsleyella, strong tu- ber development occurs in the species that are most markedly epi Such species as F. apetala, F. garleppiana, and F. inflata bear cylindrical tubers that often grow between rocks or even may be exposed above the soil (see Fig. 4). The tubers of F. chloroloba Fig. 12) are smaller and more spherical than those ofthe above-mentioned species. In general, the most seasonal species of this section, with marked leaf drop during the dry season, exhibit the strongest tuber development. On the other hand, most species with opposite, basically el- liptic leaves tend to grow terrestrially and pre- serve their leaves for most of the year. These species, such as F. membranacea, F. tillettiana, and F. huanucoensis, show very little evidence of tuber development; instead, swollen, flexuous stems sometimes occur (see Fig. 21), and appear to provide a moderate food and water storage capacity to the plant. Floral characters. Hemsleyella is the only section of Fuchsia that is characterized by the absence of petals in all species. The only other phyt tiv Vi 220 species in the genus lacking petals is F. procum- bens (sect. Skinnera), although other species of sects. Skinnera and Ellobium show strong ten- dencies toward petal reduction. The length and shape of floral tubes serve as reliable specific characters in sect. Hemsleyella. Fuchsia cestroides has tubes just 18-20 mm long, and F. nana has tubes 21-31 mm long, whereas tubes exceeding 140 mm are known in F. apetala and F. garleppiana. With the exception of F. apetala, relatively little variation in floral tube length occurs within species; in some ofthe long- tubed species, the lower range of tube lengths can be attributed to measurements of young flowers whose sepals open upon drying. In F. apetala, two distinct floral lengths have been found, those with tubes 10-16 cm long and those with tubes 4—8 cm long. Although only two collections with intermediate tube lengths were seen, the paucity of collections and the close similarity of stem, leaf, and pubescence characters in plants of dif- ferent tube length preclude ioa. F. ape- tala into separate taxa at this ti In sect. Hemsleyella, a uui band-type nec- tary lines the lowermost portion of the floral tube (see Berry, 1982, fig. 48). The nectary is similar to those found in sects. Skinnera and Ellobium. It usually causes the tube to appear bulbous at the base, a condition that also results in part from the occurrence of a constriction of the floral tube Just beyond the nectary. The size of the basal, bulbous part of the tube varies between species, but in all of them it serves as a kind of nectar reservoir. Hairs on the inside of the tube are generally absent in the basal part of the tube and usually appear at the constriction and further up the tube; these hairs serve to prevent the nectar from flowing down and out of the tube. Partic- ularly large nectar reservoirs and strong floral tube constrictions are characteristic of F. inflata (Fig. 14) and F. chloroloba (Fig. 12). The tion at the base of the sepals is char- acteristic of all species of sect. Hemsleyella and is found elsewhere in the genus only in sect. Que- lusia. The degree of connation, as well as the shape of the sepals, largely determine how far the sepals spread or become reflexed at anthesis. Strongly reflexed sepals occur in F. tillettiana, F. insignis, F. huanucoensis, and F. juntasensis (see Fig. 21), where they are characteristically long and narrow. The sepals of most other species are moderately spreading at anthesis. Floral color is an important character in the section, but it is usually lost upon drying. The ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 sepals of most species differ in color from the tube, generally becoming greenish towards the apex. In F. chloroloba, a particularly striking contrast occurs between the glaucous, orange flo- ral tube and the parrot green sepals. Fuchsia in- flata also has mostly green sepals; however, their color intergrades less abruptly with that of the floral tube. Pale pink floral tubes occur in both F. garleppiana and F. pilaloensis. Other char- acteristic colors of floral tubes are red-purple (F. nana), cerise (F. tillettiana), and mauve or lav- ender (F. juntasensis and F. cestroides). Pubescence. Vestiture characteristics, espe- cially on the floral tubes and young d parts, can be useful in distinguishing certain species of sect. Hemsleyella. Fuchsia chloroloba is remarkably glabrous throughout and is the only species with glaucous floral tubes. Fuchsia jun- tasensis and F. membranacea are generally subglabrous on both leaves and flowers. The flowers of F. tillettiana have gland-tipped hairs when fresh, a characteristic that helps distinguish -P have densely pilose or pilulose branchlets and young leaves; their floral tubes are likewise densely pilose. Fuchsia tunariensis is character- ized by a fine, velutinous pubescence throughout, whereas F. apetala has a very shaggy, hirsute vestiture on young growth, ovaries, and the basal part of “m ubes. Pollen. Pollen grains of Fuchsia share the distinctive features of other onagraceous pollen, such as viscin threads, protruding apertures, par- acrystalline-spongy ektexine, and solid endexine. The genus is unusual, however, in its basically 2-aperturate condition, with bilaterally symmet- rical grains, although a number of species have grains that are 3-aperturate and radially sym- metrical. During the past ten years, Fuchsia has been the subject of intensive palynological stud- ies, using light, scanning electron, and transmis- sion electron microscopy (Nowicke et al., 1984; Praglowski et al., 1983; Skvarla et al., 1976, 1978). As a result of these studies, the most useful taxo- nomic characters within the genus have proven to be aperture d viscin thread morphol- ogy, and pollen grain Pollen grains of sect. Honsievella are basically 2-aperturate, but 3-aperturate grains have been found in several collections. Pollen of Berry & Aronson 3010, 3011, and 3012 (F. aff. inflata, 1985] see Nowicke et al., 1984, fig. 76) is almost en- tirely 3-aperturate; chromosome counts of the first two collections indicate that they are tetra- ploid, but with meiotic abnormalities (see Cy- tology). In addition, pollen stainability of these collections varies widely, for example, 47%, 61%, and 9996 in counts of 200 grains from different flowers of 3010 and 3796, 77%, and 8896 in dif- ferent flowers of 3012. Collections of F. inflata from other localities have entirely 2-aperturate pollen, and two chromosome counts from these plants were both diploid, so it seems possible that the series of Berry & Aronson 3010, 3011, and 3012 are allotetraploid hybrids of F. inflata and some other member of sect. Hemsleyella. In two collections of diploid species, F. tillet- tiana (Berry 3267-B) and F. membranacea (Ber- ry 3278), 5—10% of the grains were 3-aperturate. A probable tetraploid collection of F. juntasen- sis, Berry 3638, has entirely 2-aperturate grains. On the other hand, grains of Berry & Aronson 3071, a long-tubed collection of F. apetala, are mostly 3-aperturate; F. apetala is presumably a tetraploid species. In this manner, the correlation noted by Nowicke et al. (1984) for 3-aperturate grains in Fuchsia to be associated with poly- ploidy is not clearly evident in sect. Hernsleyella. The viscin threads of sect. Hemsleyella are all segmented-beaded, which is considered the primitive condition for the genus and is shared wic 1 sparsely segmented threads characterize the re- maining sections. The unusually large size of the pollen grains in sect. Hemsleyella is useful in distinguishing its members from other species in the genus. In seven species of sect. Hemsleyella examined by Praglowski et al. (1983), the longest axis mea- sured is between 97 and 130 pm, far longer than Leh all but two of g THE EVOLUTION OF FUCHSIA SECT. HEMSLEYELLA A number of unique characters or ones found only in one or two other sections of Fuchsia occur in sect. Hemsleyella and make it a very distinc- tive Sand specialized group within the genus. These the following, with other sections sharing them indicated in paren- theses: 1) Loss of petals (one species of sect. Skinnera, BERRY — FUCHSIA SECT. HEMSLEYELLA 221 although strong petal reduction occurs in oth- er species of sects. Skinnera and Ellobium). 2) Smooth band nectaries (sects. Skinnera and Ellobium). 3) Presence of tubers in most species (sect. E/- lobium). 4) Alternate leaves in some species (sects. Skin- nera and Kierschlegeria). 5) Sepal connation beyond the floral tube (sect. Quelusia 6) Floral e more than 10 cm long in some specie T) EE ee epiphytism, and dry sea- son flowering with accompanying leaf drop (one species in sect. Ellobium). All species in sect. Hernsleyella lack petals, have band nectaries, and have partially connate sepals. However, not all the species possess the remaining specialized features; those species with all or most of the above traits can be assumed to be most advanced within the section, whereas the species with characters common to the ma- jority of the other sections in the genus, such as opposite-whorled leaves, little or no tuber de- velopment, medium-length floral tubes, and lit- tle seasonality, are interpreted as indication of lower levels of specialization and the more an- cestral condition of the section. Based upon these criteria, the ancestors of sect. Hemsleyella would have been diploid, terrestrial, non-tuberous, moderately seasonal shrubs, with opposite-ter- nate leaves, moderate-length floral tubes (30-50 mm long), band nectaries, and reduced petals. The present-day group corresponding most closely to this assemblage of characters is sect. Ellobium, which includes three species ranging from e Rica to Central Mexico. Although occupying a more northerly range than sect. Hemsleyella, sect. Ellobium shows specializa- tion trends very similar to those found in sect. Hemsleyella, including i increasing Spon din epiphytism, reduction in petal size, and a m marked presence of tubers iue et ii 1982). On the other hand, these same trends in sect. Ellobium are accompanied by a progression to much more congested and defined inflores- cences than those found in sect. Hemsleyella. Berry (1982) considers sect. Ellobium to be a relatively recent arrival to Central America, based on the similarities of its pollen, leaves, and flow- ers to the more widespread and more numerous species belonging to the South American sects. Fuchsia and Hemsleyella. Whether sect. Ellob- 222 Nge Ta RoLoBA INFLATA La conto r PLE, sien! SALICIFOLIA vue — — — — UNTASENSIS e res EMBRANACEA HYPOTHETICAL ANCESTOR (SEE TEXT) FiGuRE 5. Specialization trends and interspecific eerie within Fuchsia sect. Hemsleyella. ium is closely related to sect. Hemsleyella or whether their similarities are the result of con- vergent eyolufon i is met i died clear. On the other hand, t nd g lrang of sect. Ra is very similar to that of sect. Fuchsia, a much larger and more general- ized section of the genus; sect. Hemsleyella likely evolved from an early offshoot of sect. Fuchsia and then radiated in similar areas, occupying lo- cally different habitats. Appropriate habitats for the epiphytic and saxicolous species of sect. Hemsleyella may have been more limited than the terrestrial forest edge habitat of most mem- bers of sect. Fuchsia, which may in part explain the greater success of the latter group, in terms of number of species and greater population sizes. Within sect. Hemsleyella, individual species can be arranged schematically from least to most highly specialized, based on trends towards greater tuber development, increased epiphy- tism, longer floral tube length, higher altitudinal ranges, and stronger seasonality accompanied by a more deciduous habit, with large, alternate cor- interspecific relationships within sect. Hemsley- ella is shown in Figure HISTORY OF CULTIVATION Because of their scarcity, non-shrubby habit, and adaptations to dry-season flowering, mem- bers of sect. Hemsleyella have rarely been cul- tivated. Fuchsia apetala (as F. macrantha) was ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 introduced into Great Britain in 1846 (Hooker, 1846); problems associated with its cultivation were treated by van Houtte (1848). Within the past 50 years, three different apet- o uchsia chloro 36.1901) flowered and was collected there in 1939. Likewise, F. inflata (N° 38.1052) was col- lected in flower at Berkeley in 1940. Finally, F. pilaloensis (N° 58.782-1) was collected in flower rom plants in cultivation in 1961 and 1965. None of these plants or their progeny have sur- vived until the present, however. In a popular article on Bolivian fuchsias, Brooke (1953) stated that two tuberous species of sect. Hemsleyella were in cultivation in Great Britain then, but she failed to mention where, or what species were involved. Fuchsia garleppiana, F. apetala, and F. tu- nariensis were briefly in cultivation at the Mis- souri Botanical Garden during 1979-1980, but all subsequently died. According to Wright (1979) and Ewart (1982), there are currently no species of sect. Hemsley- ella known to be in cultivation. SYSTEMATIC TREATMENT Fuchsia sect. Hemsleyella Munz, Proc. Calif. Acad. Sci., Ser. 4, 25: 74. 1943. TYPE: F. apetala Ruiz & Pavón. Erect to scandent shrubs, prostrate subshrubs, lianas, or epiphytes. Cylindrical to subspherical tubers or thickened stems often present. Leaves dant, usually axillary or clustered at the branch tips, or racemose in F. cestroides. Sepals shorter than the floral tube and connate at the base. Pet- als lacking. Stamens erect, in two whorls. Pollen 2-(3-)aperturate, mu or radially symmet- ric, with segmented-beaded viscin threads. Berry with ca. 50-2 50 seeds, these d laterally and irregularly triangular-oblong in outline. Ga- metic chromosome number, n = Distribution. Cloud forest or less commonly scrub forest, puna or subpáramo in the tropical Andes from Venezuela 5 “apa (10°N to 17°S), at altitudes of 1,100—4, Measurements used in hh i and species de- scriptions are based on dried herbarium speci- mens. Leaves are often lacking when flowering specimens are collected, but leaf position can be 1985] determined by leaf scars on the ste m. Termi- nology for leaf venation follows Hickey (1973). The use of the term “floral tube" refers to the position of the flower from the top of the ovary to the point of insertion of the filaments or the base of the sepals. = w e CA P Q e 7a. 7b. ternate .... KEY TO THE SPECIES OF FUCHSIA CT. HEMSLEYELLA Leaves or leaf scars all or mostly opposite or Leaves or leaf scars all or mostly alternate ....... 7 2a. asia — 18-20 mm long; pedicels 3- 4 mm long 2) F. cestroides 2b. Floral ae 25-56 mm long; pedicels 15- 5 mm long . Floral tubes and ovaries pubescent, with pi- lose, velutinous or gland-tippe d hairs u... Floral tubes and ovaries glabrous or subglab- rous 4a. Floral tubes finely puberulent; sepals spreading; leaves velutinous on under- side. Southern Peru and Bolivia ............ 4) F. tunariensis 4b. Floral tubes usually with gland: -tippe ed leaves subglabrous. Venezuela (13) F. “eS qasi . Floral tubes narrowly funnelform, ca. wide at the mouth, 31-34 mm long; sepals narrowly i 4-5 mm wide. Centr (5) F. Foi. Floral tubes funnelform, 7-12 mm wide h, 25-51 mm long; sepals lance- ovate, 4-11 mm wide. Venezuela, Bolivia... 6 6a. Petioles 4-10 mm long; sepals divergent - to recurved at anthesis, lavender-violet, same color as floral tube. Bolivia ......... (8) F. juntasensis . Petioles 10-35 mm long; sepals an ing at anthesis, greenish; floral tube p 9) Venezuela F. membranacea (10) F Floral tubes 21—31 mm long Floral tubes 32-160 mm long in the lower '4, then abruptly widened to 8— 14 mm in the mi iddle (6) ) F. — Ova the tube generally ons oa slightly bul- bind at the base and 3-7 mm wide at broadest 10a. Foral tubes hirsute in lower '^, rose to dull orange ually borne on short, knotty-tort tems .. . (1) F. apetala 10b. Floral tubes pilose to subglabrous, light F. garleppiana lla. Floral tubes and ovaries completely brous; sepals bright green throughout (3) F. chloroloba BERRY — FUCHSIA SECT. HEMSLEYELLA 11b. 13b. (oz Fuchsia unduavens 223 Floral tubes and ovaries lightly pubescent to densely hirsute; sepals not entirely bright green 12a. Sepals 25-37 mm long 13 12b. Sepals 10-25 mm long „u . 14 . The longer stamens 30—40 mm long; se pals strongly recurved at anthesis ....... (7) F. insignis The longer stamens 15-22 mm long; sepals spreading to divergent at anthesis (12) F. salicifolia 14a. Floral tubes and ovaries densely pi- ose or hirsute 14b. I bead cag ovaries puberulent or on short, knotty a l tubes rose to orange-red. Peru and Bolivia (1) F. di m ot on sho knot si stems; floral tubes many pisk, W (11) F. ee to sparsely pubescent; ae -— ly lanceolate, with long ( 12) F F Mdb 1. Fuchsia apetala Ruiz & Pavón, Fl. Peru v. 3: 89, pl. 322, fig. b. 1802. Macbride, Field Mus. Nat. Hist., Bot. Ser. 13: 546. 1 TYPE: Peru. Dept. Junín: Huasahuasi (7 od " 37'W, 11?15'S), Apr. 1779, Ruiz & Pavón (lectotype, MA, here designated; photo- MO). There are four Ruiz and Pavón collections at MA which are labelled as Fuchsia apetala; the lectotype is one of two with “N° 11/90" written on a separate label at the base of the sheet, which also contains a lengthy, handwritten description of the plant, presumably by the collectors (see Fig. 6). The label cites at the end, “Habitat Huasa- huasi in elevatis humidis et frigidis. Floret abri." Figures 6—9. Fuchsia Tu TUM Hooker, Bot. Mag. 72: t. 4233. 1846 unz, Proc. Calif. Acad. Sci., Ser. 4, 25: n 1837, Mathews 1197 (holotype, K-Hooker Her- barium; isotypes, K-Bentham Herbarium, OXF, P, TCD, mm hirsuta Hemsley, J. Bot. 14: 69. 1876. Munz, Proc. Calif. Acad. Sci., Ser. 4, 25: 81, pl. 13, fig. 70. 1943. TYPE: Peru. Dept. unknown, June 1854, Lechler 1989 (holotype, K). is Munz, Proc. Calif. Acad. Sci., Ser. 4, 25: 81, pl. 13, E 71. 1943. TYPE: Bolivia. Dept. o gas, Unduavi, 3,600 m, 13 Feb. 1907, peti 2925 (holotype, US-1177335, photographs, NY, UC; isotypes, NY, Z). 224 ANNALS OF THE MISSOURI BOTANICAL GARDEN Herbarium Peruvian Ruiz et Pavon Tuta ¿tala R.P me d HERBARIUM HORT! BOTANIC! MATRITENSIS FiGURE 6. Photograph of the lectotype of Fuchsia apetala, Ruiz & Pavón, 1779 (MA). [Vor. 72 1985] FiGurE 7. Flowering branch of Fuchsia apetala, long tubed form, from Berry & Berry 3070, Dept. Ju- nín, 1 1 1 = 2 1 P rostrate t or on the edge of 1 banks rich in Sphagnum or humus, or vine-like epiphytes on moist tree trunks. Tubers ellipsoid to cylindrical, 2-8 cm long and 1-5 cm thick, covered by copper-tan, flaking bark, often interconnected by slender stems. Stoloniferous shoots 2-12 dm long when epiphytic (lacking when terrestrial), 2-4 mm thick with freely exfoliating red-copper bark; leaves BERRY — FUCHSIA SECT. HEMSLEYELLA 225 ^ AA ASKA. M 1 | À m ç FicureE 8. Flowering branch of — form of Fuchsia — d from E erry & Berry 2565, Prov. La Convención, Cuzco, Peru. Note the hirsute pubescence at the pid of the floral tubes. and flowers borne on short, lateral, knotty-tor- tuous stems with tightly packed leaf and pedicel scars, these stems rarely more than 4 cm long, 3-8 mm thick, dull gray, pilose to hirsute. Leaves alternate and usually tightly clustered at the branch tips, mostly deciduous at anthesis or pres- ent on plants flowering in shady habitats, soft membranous, subcordate to ovate-elliptic, 60— 120(-150) mm long, 35-65 mm wide, apex acute to acuminate, base rounded to obtuse or sub- cordate, densely pilose-hirsute when young, green and subglabrous to strigose above at maturity, slightly paler and pilose below, with appressed, densely villous hairs along veins; margin sub- 226 O 1! 2 3cm. FIGURE 9. Leaves of Fuchsia apetala, from ial E cm 2564, Prov. La Convención, Dept. Cuzco, entire to denticulate, secondary veins 6-8 on either side of the midvein. Petiole 25-50(-70) mm long, 2-3.5 mm thick, reddish, pilose; stip- ules filiform, 2-3 mm long, deciduous. Flowers crowded near the tip of lateral, knotty twigs, usu- ally 3-12 per shoot; pedicels pilose to hirsute, (1018-35 mm long; ovary cylindrical, subte- tragonous, 6-12 mm long, 2-4 mm thick, short pilose to densely hirsute, the hairs often shaggy and to 2 mm long. Floral tube subcylindrical, 4— 16 cm long, 3-6 mm wide and somewhat nodose at the base around the nectary, then narrowed to 2-4 mm wide followed by a gradual widening until 7-10 mm wide at the rim, sometimes di- lated to 8-13 mm wide just below rim, pilose to densely hirsute in the basal 1—2 cm, loosely pilose for the rest of length, inside subglabrous to vil- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 lous 10-35 mm above the base. Sepals ovate, broadly acute, 16-25(-30) mm long, 7-12(-16) mm wide, usually broader than the tube, connate at the base for '4-' the total length, free segments spreading to slightly recurved at anthesis. Tube rose to light orange-red, at times with a dark purple, nitid ring at the rim, sepals rose red and usually dull yellow-green distally. Nectary lus- trous green-purple, lining the basal 5-6 mm of the tube. Filaments green to dull pink with age, the antesepalous ones 10-18 mm long, the an- tepetalous ones 8-15 mm long; anthers narrowly oblong, 3.5-4.5 mm long, 1.5-2 mm wide, yel- low. Style pale green, subglabrous to densely (re- trorse) villous between 10—40 mm above the base, the basal 10-15 mm usually glabrous; Pues green, globose- -o slightly 4-cleft, 2-4 m long, 1.5-2. e, exserted 1-15 mm be- vond the anthers. Bor oblong, red, subtetrag- onous, 17-25 mm long, 8-12 mm thick, subgla- brous to densely hirsute, more or less verrucose, many-seeded. Gametic chromosome number, Distribution. Eastern slopes of the Peruvian Andes from Dept. San Martín south to Dept. Puno and in Bolivia, Depts. La Paz and Coch- abamba; in upper cloud forest and the lower reaches of the puna, on rocks, tree trunks and Sphagnum banks, (2,800—)3,200-3,850(-4,200) m, but most common around 3,600 m. Flow- ering principally in the dry season, from June to September. Figure 10. Specime ined. PERU. SAN MARTIN: valley of Rio Apisoncho, 30 km above Jucusbamba, 7°55’S, 77*10"W, Hamilton & Holligan 1003 (K), 1073 (K, UC). HUÁNUCO: Carpish summit, Ferreyra 2395 (USM), 10030 (USM); about Muña and Pillao, Lobb s.n. (W); UNÍ oncepción, 22 km E of Comas, Con- cepción- Pond Berry & Aronson 3068 (MO, USM); m E of Comas, road to Satipo, Berry & Aronson ( AC Cordillera et between Tambo, Hda. Luisiana, ca ta, entrance to (USM); Prov. Caneallo, desce (CUZ); Punaccuanca Pass, West 3669 (GH, MO, UC). CUZCO: Prov. Convención, Km 140-144 sip Cuzco to Quillabamba, below Abra de Málaga, Ber. erry 2562 (MO), 2563 (MO), 2564 (MO), 2565 (MO), 2566 (MO), Berry & Aronson 3030 (MO, USM), 3033 (MO, 1985] BERRY — FUCHSIA SECT. HEMSLEYELLA 227 = ` X ) p A! r / T / : LA @Fuchsia apetata ® Fuchsia tunariensis * Fuchsia salicifolia e P d oFuchsia garleppiana X *Fuchsia juntasensis =“ * Fuchsia nana e £. -10° [ i -—7 b n qu ER e | e La: y= i A ‘ e +e, é Lise , L - 20? r T l O 300 600 Km. 80 T5 L L FiGu Peru and Bo USM); Prov. Paucartambo, 1 km from Tres Cruces, road to Acanaco, Berry et al. 2593 (MO—2 sheets); 4 wer Panticalla Pass, Cook & Gilbert 1857 (US); v. La vención, ca. 25 km NE of Hda. Luisiana a y Rio Ap urimac, Dudley 11215B (NA—2 sheets); Tres Cruces, Gentry et al. 23435 (MO), Pennell 13897a (PH), Vargas 12207 (CUZ), Weberbauer 6975 (F—2 sheets, GH); Salcantay, Heim, 1848 (Z), Rauh P1473 (RSA); Colinas de Sacsayhuamán, Herrera 2189 (BH, 209 (CUZ); Yucay, Soukup 715 (BH, F, Picchu, 3 km from ba, Chincheros, Vargas 1641 (CAS, CUZ, UC), 9602 H, G-DEL, GH, K, MO, NA, UC); Combapata, Can- chis, Vargas 1900 (CUZ, GH); Prov. Quispicanchis, Pucuta, Chutacuchu, Marcapata, Vargas 3750 (CUZ); Panticalla-Yanamanche, Vargas 4432 (CUZ); Prov. RE 10. een of the species of Fuchsia sect. Hemsleyella restricted to Bolivia or occurring in both liv Quispicanchis, Layampampa, Vargas 6649 (CUZ); Prov. Anta, Mollepata, toward Ramicruz, Vargas 8343 (CUZ); Prov. Calca, near Lares, Vargas 1 0962 (CUZ); Prov. Paucartambo, Dtto. Marcachea, forest of Pucara, viis 11170 (F, G-DEL, K, NA, UC); Prov. bamba, Peñas, Vargas 15730 dur without locality, Wedel 4771 (P); Beck 4168 (MO); Nor Yungas, 2 km from Unduavi to 228 Coroico, Berry & id 2576 e. d Braun km low ' Ta- O Paz 6904 (BM); Unduavi, Buchtien, 1910 (F); Ingenio, be- n Sorata and ics pas 1142 (NY), Wil- ( ve Pongo, D'Arcy & Be- B , W); Pelechuco, Pearce, 1864 (BM); Murillo, 22.5 km below Zongo ese : 1 E o Solomon1 1277040; Pongo, Tate 188 (NY —2 sheets); Cordillera , Ri , Tate 718 (NY); Prov. pr Valley of i say Weddell, 1851 (P). cocua- : Rodeo-Mizque road, Badcock 830 (K): prima Aparcita, Brooke 6098 (BM, F, G-DEL, NY NW of Cochabamba, above cacha ; 6920 (U); Luruni, hot springs Fun Quillacollo, ‘rail to Cochabamba, Brooke 6786 (BM); Ayopaya, — Cardenas 3369 (US); To- gas da Vandiola, Cardenas 3754 enka-Carrasco, Cárdenas 6135 (US); Prov. Ayopaya, Weddell 4129 (P, US). WITHOUT DEPARTMENT: Bang 146 (BM), 2834 (F, GH, K, LE, MICH, MO, NY), Bridges s.n. (BM), Pearce 771 (K). Fuchsia apetala was the first apetalous species described in the genus, but the notable differ- ences between Ruiz and Pavón's type description and the accompanying illustration, as well as the apparent wide variation in floral tube lengths in the species, have led to differing interpretations as to what plants should be included under this name. It is clear from the original description and from most of Ruiz and Pavón's collections labelled by them as F. apetala, that they applied this name to plants with pilose floral tubes 10— 14 cm long, with relatively small sepals and with flowers borne on short, knotty stems (see Figs. 6, 7). The plate that accompanies the type de- scription, however, shows a plant with much shorter floral tubes, long-spreading sepals and smooth flowering and leaf-bearing stems; these discrepancies appear to be the result of a con- fusion of different apetalous specimens that they collected in central Peru, and not to different forms of the same species. The herbarium at MA contains four different Ruiz and Pavón collections labelled as F. apet- ala. One of these bears a slip of paper with the name “‘Cavanilles” above the label and includes a plant with ampliate, short-tubed flowers 55— 60 mm long and a single, large, ovate leaf 15 cm long by 9 cm wide, but with the typical knotty ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 stem of F. apetala. The remaining EA all have long, narrow-tubed, subcylindrical flowers on verrucose stems, agreeing closely un the type description. The sheet chosen as the lectotype (Fig. 6) also contains a lengthy handwritten de- scription very similar to the one used to describe the species and cites as the collecting locality one of the two sites mentioned in the type descrip- tion, which states, “Habitat copiose in Huassa- huassi et Muña nemoribus" (Ruiz & Pavón, 1802: 89). Unless there is a Huasahuasi in Dept. Huá- nuco, these localities correspond to two different areas, the first in Dept. Junín and the second on the Dept. Huánuco-Pasco border. Additional Ruiz and Pavón collections labelled as F. apetala are found at BM (three sheets, two with long floral tubes and one with a mixture of a long- tubed and a short-tubed plant), F (two fragments taken from MA, one from a long and narrow- tubed plant, the other from a short, broad-tubed individual), G (three sheets, - long-tubed, in G- , G-BOIS, and G-“‘Pavén 213"), MO (“ex BM," long-tubed), and P fois Pavón 36," long-tubed). Subsequent authors treating the apetalous fuchsias based their concept of F. apet- ala either on the type description (Macbride, 1941) or else on the illustration (Hemsley, 1876; Munz, 1943, 1974). As defined here, F. apetala is best character- ized by the presence of short, knotty or verrucose stems on which the leaves or flowers are tightly grouped, as well as by a typically pilose to hirsute pubescence on the leaves and flowers. The leaves, although generally absent when the plant flowers, are larger and more ovate-cordate than any other apetalous species (except possibly F. insignis), and the tubers are particularly well developed. Populations of F. apetala may differ consider- ably in floral tube length, the only species in the section to do so; nonetheless, almost all of the plants share the aforementioned traits, as well as similarly short, ovate and dull-colored sepals with rose scarlet to dull orange floral tubes In general, the specimens of F. apetala can be divided into long-tubed (10-16 cm) or short- tubed (4-8 cm) groups. The long-tubed plants, including the type of F. macrantha, seem to grow mostly as epiphytes in cloud forest and are re- stricted to Depts. Huánuco, Pasco, and Junín, in Peru. Short-tubed plants, including the types of F. hirsuta and F. shinier pies pow at higher elevati tions, either terrestrially along banks or among rocks; these 1985] plants occupy a wider range, from Dept. San Martín in Central Peru to Dept. Cochabamba in Bolivia. yen few specimens with intermediate ed; Macbride 4901, with tubes 85 mm long, and Macbride 4970, with tubes 70-120 mm long, both from Dept. Huán- uco in Peru, were the only such sheets seen. On the other hand, both short- and long-tubed plants have been found to occur in the same general area; in Dept. Junín, between Concepción and Satipo, individuals with floral tubes 6 cm long were found growing out of road banks at 3,600 m, 123-133 km west of Satipo (Berry & Aronson 3068, 3069). Further east along the same road, plants kaya floral tubes 1 2-1 3 em pe were found 350-3,400 m, 72-73 km west of Satipo eds & Aronson 3070, 307 D. Except for the difference in the tube length, the two populations are very similar in characters such as coloration, pubescence, tube shape, knotty stems, and leaf type; interestingly, the sepals are approximately the same size and shape in both populations, unlike the large, flar- ing sepals shown in the short-tubed plant in Ruiz and Pavón's illustration. Before deciding wheth- er two distinct taxonomic entities should be rec- ognized for the short-tubed and the long-tubed plants, it will be necessary to undertake more intensive collections in central Peru, where both forms and possibly some with intermediate tube lengths occur. Three different short-tubed populations of F. apetala were examined cytologically, and all were tetraploid (n = 22). The only other tetraploids known in sect. Hemsleyella are the distantly re- lated F. juntasensis and in a population of pos- sible hybrids (x F. inflata) in eastern Peru. Al- though no counts were obtained from long-tubed plants of F. apetala, Berry & Aronson 3071 has a large percentage of triporate pollen grains, a character generally associated with polyploidy in the genus (Praglowski et al., 1983; Nowicke et al., 1984). A number of collections are doubtfully re- ferred to F. apetala because they differ from the description and general characters of the species in one or more traits. Two ofthese are long-tubed plants from Ampay, Dept. Apurimac, Peru, that cannot be unequivocally placed either in F. in- flata or F. apetala. Vargas 1000 (BH, CUZ, GH, UC, 10 June 1938, 3,200-3,500 m) has floral tubes 11-12 cm long, which are pilose, yet am- pliate as in F. inflata (10-16 mm diam. in the BERRY — FUCHSIA SECT. HEMSLEYELLA 229 middle ofthe tube); the leaves, however, are large and ovate-cordate, as in F. apetala. The second collection from Ampay, Marín 2253 (F, July 1950, 3,600 m), has subcylindric, pilose floral tubes 90-105 mm long and ovate leaves, but lacks the knotty stems typical of F. apetala. Dud- ley 11215B (NA, 19 July 1968, Peru, Dept. Cuz- co, Prov. La Convención, ca. 25 km NE of Hda. Luisiana and Río Apurimac) is a fairly typical short-tubed example of F. apetala, except for its very large and broad sepals 36 mm long by 17 mm wide. Berry & Aronson 3014 (MO, 21 July 1978, Peru, Dept. Cuzco, Prov. Quispicanchis, 3,120 m, 91 km from Ocongate above Marca- pata) has short, pilose floral tubes and knotty stems, but the tubes are more constricted than is usual in F. apetala, and the collection was diploid, 2n = 22. Chávez 3489 (MO, 12 Nov. 1976, Peru, Dept. Cuzco, Quebrada Las Peñas, 3,150 m, 10 dm tall in rock fissures) is a plant with somewhat knotty stems and moderately ampliate floral tubes, yet the flowers have fine, whitish pubescence and pink sepals, and it is seemingly intermediate between F. apetala and F. tunariensis. Another group of old collections, presumably all from central Peru, recall the illustration that accompanies Ruiz and Pavón's type description of F. apetala. Lobb s.n. (K) has rather ampliate tubes 46-48 mm long, large and broad ovate sepals (18 mm long, 9 mm wide), smooth stems, and large leaves 13 cm long and 8 cm wide. Lobb 112 (K, W, from Muña, Dept. Huánuco) has similar flowers, with tubes 58—60 mm long, se- pals 21 mm long and 10 mm wide, and filaments 17-20 mm long. Similar collections are Gay, 1839-1840 (P), but with very pilose leaves and knotty stems, and Dombey s.n. (F), from **Chu- rupallana," with petioles to 6 cm long. It is pos- sible that these collections correspond to Ruiz and Pavón's illustration accompanying the type description of F. apetala and that they represent a yet undescribed apetalous species from the Huánuco-Pasco region, most closely resembling F. insignis from Ecuador. Further field work will be necessary to determine if this is so. 2. Fuchsia cestroides Schulze-Menz, Notizbl. Bot. G 13: 550. 1941. Munz, Proc. Calif. Acad. Sci., Ser. 4, 25: 83, pl. 14, fig. 74. 1943. TYPE: Peru. Dept. Piura: Prov. Ayavaca, W slopes 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Gi - 10° - 15° r ü s 1 7 Jo mum A! ¿ / c». d Xe (@Fuchsiq cestroides iGoFuchsia huqnucoensis e Fuchsia inflata O Fuchsia chloroloba FiGURE 11. of the Andes below Frias, 1,000-1,200 m, 21 May 1912, Weberbauer 6423 (holotype, B, destroyed in World War II; isotypes, F— 2 sheets, GH, US; photographs of US iso- type, BH, NY, UC). Erect to scandent shrubs 0.5-3 m tall. Branch- lets terete, 1.5-3 mm thick, puberulent to pilose; older stems 5-40 mm thick, with exfoliating bark Leaves opposite or ternate, membranous, ellip- tic, 26-100 mm long, 14-15 mm wide, apex and base acute, subglabrous to puberulent above, pu- berulent below, especially on the veins; margin subentire to serrulate, secondary veins 6-14 on either side of the midvein. Petioles strigose, (2—)4— 8 mm long. Stipules lanceolate, ca. 1 mm long, ca. 0.5 mm wide. Flowers generally numerous on terminal and axillary racemes with small, de- ciduous, lance-filiform bracts; rachis (1 5-)25-35 mm long. Pedicels puberulent, 3-4 mm long. Ovary cylindrical, 3-4 mm long, 1-1.5 mm wide, puberulent to pilose. Floral tube narrowly fun- nelform, 18-20 mm long, 2-3 mm wide at the Distribution of the species of Fuchsia sect. Hemsleyella restricted to Peru. base, narrowed to ca. 1.5 mm wide in the lower !^, then gradually widened to ca. 4 mm wide at the rim, lightly puberulent to pilose outside, re- trorse villous inside in lower !^. Sepals oblong- lanceolate, 5-7 mm long, 2.5-3 mm wide, obtuse to acute at the apex, connate for ca. 1.5 mm at the base, spreading at anthesis. Tube rose pink to lavender, sepals greenish (in bud) to dull pur- ple. Nectary ca. 3 mm high. Filaments green, the antesepalous ones 3.5—6(—7) mm long, the an- tepetalous ones 2-5 mm long; anthers broadly elliptic, yellow, ca. 1.5 mm long, ca. 1 mm wide. Style glabrous, exserted 5-8 mm beyond the an- thers. Stigma clavate, 1.5-2 mm long, ca. 1 mm wide, slightly 4-cleft at the apex. Immature fruit oblong, 8 mm long, 5 mm wide, with ca. 50 seeds. Gametic chromosome number, n = 11 Distribution. Northwestern Peru. Rare and endemic to scrub vegetation between 1,100 and 1,700 m on the Pacific-facing slopes ofthe Andes in Depts. Cajamarca, Lambayeque, and Piura. Figure 11. 1985] Specimens examined. PERU. CAJAMARCA: Andaloy, between San Benito and Guzmango, 1,600-1,800 m, Berry et al. 3600 (liquid-preserved specimen, author's collection), López & Sagdstegui 6251 (US) LAMBAYEQUE: Prov. Lam yea ! E of OI- mos, between Olmos and Jaen on Mesones-Muro hi way, 1,150 m, Hutchison & Wright 3408 (RSA, UC). This is an extremely rare species that is some- what unusual for the section. Indeed, Munz (1943) remarked that in its shrubby habit, opposite-ter- nate, elliptic leaves and dense inflorescences of small flowers, the type specimen of F. cestroides resembled fairly closely certain species in sect. Fuchsia, such as F. lehmannii. Nonetheless, the lack of petals and the band-type nectary clearly place it in sect. Hemsleyella. Geographically, F. cestroides is strongly isolated from other mem- rs of the section, by inhabiting the rather dry, open scrub vegetation of the Pacific slopes of the northern Andes of Peru. e small number of specimens collected makes it difficult to assess the degree of vari- ability in vegetative characters in F. cestroides. Both collections from Cajamarca were leafless when collected in flower, and the leaves present on Hutchison & Wright 3408, from Lamba- yeque, are quite distinct from those of the type specimen: they are opposite, semicoriaceous, smaller (30 mm long, 15 mm wide), with obtuse tips and petioles just 2-3 mm long. Further col- lections from the northwestern departments of Peru may eventually show that these specimens belong to a different taxon from F. cestroides, but one that is related to it. 3. Fuchsia chloroloba Johnston, J. Arnold ar bor. 20: 243. 1939. T O £ Zz E < y: Š S ww ^E: 5 N s Pennell 13973 (holotype, GH, photograph at NY; isotypes, F, NY, PH). Figure 12. Fuchsia god sensu Macbride, Publ. Field Mus. Nat. Hist., Bot. Ser. 13: 564. 1941, pro parte. Fuchsia tuberosa Krause var. typica sensu Munz, Proc. . Acad. Sci., Ser. 4, 25: 76, pl. 12, fig. 66. 1943, pro parte. Shrubs 1-2 m high, usually growing in rocks or epiphytic on trees, with globose to ellipsoid tubers 1-4 cm thick. Branchlets terete, smooth, purplish, 1.5-5 mm thick; older stems 0.5-2 m long, 5-18 mm thick, with freely exfoliating bark. Leaves alternate, usually lacking at anthesis, membranous, narrowly lance-ovate, 60-120 mm long, 20—55(-60) mm wide, apex acuminate, base BERRY — FUCHSIA SECT. HEMSLEYELLA 231 cordate to subcordate, glabrous on both sides; margin subentire to remotely denticulate, sec- ondary veins 5—9 on either side of the midvein. Petioles 14—45 mm long. Stipules filiform, ca. 2 mm long, 0.2-0.3 mm wide, deciduous. Flowers few to numerous, axillary or clustered at the tips of young shoots. Pedicels dull red or green, 26- 60(-70) mm long. Ovary (narrowly) cylindric, 10-24 mm long, 2-4 mm wide, lustrous light green. Floral tube (38-)41—-51 mm long, 4-5 mm wide and bulbous at the base, thereupon strongly constricted to 1.5-2 mm wide in the lower !^—!4 of the tube, then abruptly widened until 10-13.5 mm wide in the upper half before narrowing slightly to 9-11(-12) mm wide at the rim, gla- brous outside, pilose inside in the lower '4. Sepals lance-ovate, thin membranous, 16-24 mm long, 5-10(-12) mm wide, apex acute to acuminate, connate for 5-7 mm at the base, tetragonous in bud, spreading-divergent at anthesis. Tube glau- cous, bright scarlet to light orange-pink, sepals parrot green. Nectary 6-8 mm high. Filaments green, the antesepalous ones 10-16 mm long, the antepetalous ones 8-13 mm long; anthers ob- =i yellow, 4-5 mm long, 2-3 mm wide. Style pale green, pilose inside in the lower !^; dins N clavate, green, 4-4.5 mm long, ca. 2 mm wide, exserted 5-15 mm beyond the anthers. pos berry not seen. Gametic chromosome number, n = 11. Distribution. Southern Peru, endemic to iit g forest in several valleys in Prov. Paucartam and Prov. Quispicanchis, Dept. Cuzco, (1, 600) 2,200-2,800 m. Flowers in the dry season, prin- cipally from May to October. Figure 11. Specimens examined. PERU. cuzco: Prov. Paucar- v T Prov. qi esp ry & y eine 3015 (MO, EOM 3024 a > km below Opispata arcapa ks DE Aronson 3017 (MO, USM), "Metcalf 30719 (A, MO, UC, US), Rauh P1282 (RSA), Vargas 6214 (CUZ), 16567 (CUZ), Wasshausen & Encarnación 783 (MO); Prov. Paucartambo, Tambomayo, Vargas 60 (CUZ— 2 sheets, F, MO); Pillahuata, Paso del Aguila, Vargas 16764 (CUZ), 22997 (MO); Prov. Paucartambo, Achirani, Marcachea, Vargas 1625 (CUZ, MO); be- tween Achirani and Medias-Mayu, Vargas 11116 (A, F, G-DEL, K, NA, UC); Cosuepata, Prov. Paucartam- bo, Velarde 1197 (RSA) Prov. ío Tambomayo, from Pillahuata Pridge to head of Tam- bomayo Grande; West 7094 (GH, MO—2 sheets). CULTIVATED: Berkeley, California, m California Bo- 232 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 AY FiGureE 12. Fuchsia chloroloba, leafy and flowering branch from Wasshausen & Encarnación i Prov. Quispicanchis, Dept. Cuzco, Peru; tubers from Berry et al. 2599, Prov. Paucartambo, Dept. Cuzco, Peru. 1985] tanical Garden Accession N° 36.1901, from seed of West 7094, in 1939 (UC). This is a very localized and distinctive species, characterized by its short, tubular floral tubes with a very marked constriction in the lower third and a very striking color pattern, with glau- cous, orange floral tubes and parrot green sepals. Fuchsia chloroloba is highly seasonal, forming well-developed tubers and flowering in the dry season, when most of the leaves are absent. This species has been confused by Munz (1943) with F. inflata, with which it grows sympatrically above Marcapata in Prov. Quispicanchis. The latter species has a similar floral tube shape, but the flowers are much longer (without apparent intergradation) and are reddish pink instead of orange, with the sepals less entirely green, as in F. chloroloba. Nonetheless, it is possible that nat- ural hybrids occur between the two taxa, as de- scribed in the section on Sympatry and Hybrid- ization. 4. Fuchsia garleppiana Kuntze & Wittmack, Gartenflora 42: 461, fig. 96. 1893. Munz, Proc. Calif. Acad. Sci., Ser. 4, 25: 82, pl. 14, fig. 72. 1943. TYPE: Bolivia. Dept. Cocha- bamba: cloud forest at Cocapata, Tunari Range, 3,000 m, Apr. 1982, Kuntze (holo- type, NY; isotypes, F-297181, F-297182, NY, photographs of F-297182 at NY, UC). Figure 13 Shrubs 0.5—4 m tall, growing epiphytically or more commonly out of rock crevices, with strongly developed cylindrical tubers 10-80 cm long and 2-4 cm thick and more or less erect branches 1-2.5 m long. Young growth densely white-pilose; branchlets terete, 2-4 mm thick, red-green; older stems 4-10 mm thick, with free- ly exfoliating bark. Leaves alternate, usually ab- sent at anthesis, membranous, ovate to elliptic- ovate, mature blades 90-130 mm long, 55-75 mm wide, apex subacuminate, base rounded to cordate, light green and subglabrous above, pi- lose along veins and margin below; margin sub- entire to remotely denticulate, secondary veins 9-10 on either side of the midvein. Petioles pi- lose, 40-73 mm long on mature leaves. Stipules filiform, ca. 2 mm long, deciduous. Flowers few to many, axillary or grouped at branch tips. Ped- icels densely pilose, greenish pink, 7-30(-60) mm ong. Ovary narrowly cylindric, 7-8 mm long, 2—2.5 mm thick, densely pilose. Floral tube nar- rowly tubular-funnelform, (45-)70-140 mm long, BERRY — FUCHSIA SECT. HEMSLEYELLA 233 3.5—6(—7) mm wide and bulbous at the base, then constricted to 2-2.5 mm wide and gradually wid- ened above until 7210 mm wide below the slight- ly constricted mouth, pilose outside, villous in- side for most of length. Sepals lance-ovate, 10— 18 mm long, 5.5-7(-8) mm wide, apex acute, connate for ca. 3 of length at the base, spreading- divergent at anthesis. Tube and sepals light to very pale pink, with a dark purple ring on the throat where the stamens are inserted. Nectary lustrous green-purple, 4—6 mm high. Filaments green, the antesepalous ones 10-12 mm long, the antepetalous ones 8—9 mm long; anthers yellow, 3-4 mm long, 1.5-2 mm wide. Style light pink, villous for most of length; stigma green, globose, 2-2.5 mm long, 2.5-3 mm wide. Young berries narrowly cylindric, ca. 22 mm long, ca. 4 mm wide. Gametic chromosome number, n = 11. Distribution. Endemic to cloud forest in Dept. Cochabamba, Bolivia, at 2,500-2,950 m. Flow- ers principally in the dry season, when leafless, from (March to) April to August. Figure 10. Specimens examined. BOLIVIA. COCHABAMBA Chapare, 54 km towards Villa Turari, Beck 14 Lr 2 sheets); 41—42 mi. from Cochabamba to Chapare, Berry & Berry 2582 (MO—2 sheets); 2583 (MO—2 sheets), 2590 (MO); 5 km from Incachaca on old road to Aguirre, Berry 3635 (MO). Incachaca, road to Aguirre, Brooke 6741 (BM, F, G-DEL, NY, S, U); Incachaca, Cárdenas 688 (US); near Chulumani, way to Yungas de Tablas, Cárdenas 6279 (NY, US); way Cárdenas s.n. (POM); Incacorral-Aduana , J. Steinbach 9538 (BM, F, G-DEL, GH, K, MO, NY, PH, S, U, Z). Of all the native Bolivian fuchsias, F. garlep- piana is the only one with floral tubes over 7 cm long. It is also characterized by the very pale pink coloration of the flowers, the dense white-pilose pubescence on the young growth, and its pri- marily shrubby habit. Although this species has well-developed, cylindrical tubers and flowers in the dry season when it is leafless, it is rarely truly epiphytic; rather, it prefers exposed rocks, from which erect stems to 1 m tall emerge. Only two other species of Fuchsia, both from Peru, can have similarly long flowers; Fuchsia apetala is basically epiphytic or terrestrial at higher ele- vations, with creeping, tortuous stems and red- dish orange flowers. Fuchsia inflata, the other species with long floral tubes, sometimes has a habit similar to that of F. garleppiana, but its floral tubes are pink-orange, much more con- stricted at the base and broader at the top, and it has yellow-green sepals. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 s Et Aj BS c ^ a E = AF t L Fi ” $ Z À IR £ yes i ad E. 3 iet ? m tf E F > i H = ,. PE E s i E i E. v" i iu À 1 ro T H K : pan Tw Et 13. Fuchsia garleppiana, flowering branch from Berry & Berry 2583, Dept. Cochabamba, Bolivia; Ten om Beck 1414, Dept. Cochabamba, Bolivia 1985] 5. Fuchsia huanucoensis P. Berry, sp. nov. TYPE: eru. Dept. Huánuco: Carpish Divide, Chinchao Valley, 8,500 ft., Oct. 1945, Sandeman 5166 (holotype, K; isotypes, K, OXF) Frutex scandens, ramis oppositis. Folia coaetanea, Si me acea, anguste elliptico-ovata, apice subacuminata, basi acuta vel rotundata, 40-80 mm longa 0 mm lata, petiolis 6-10 mm longis. m ta 2-3 mm limbis sub anthesi valde patentibus. scandent shrubs with oppositely mm thick, purplish, pilose with w branches pilose, with lustrous purple bark split- ting longitudinally. Leaves present at anthesis, opposite, firmly membranous, narrowly elliptic- ovate, 40-80 mm long, 18-30 mm wide, apex subacuminate, base acute to rounded, apparently dark green and subglabrous above, paler below with pilose midvein and margin; margin sub- entire, secondary veins 6-10 on either side of the midvein. Petioles strigose, purplish, 6-10 mm long. Stipules lance-triangular, 1-1.5 mm long, deciduous. Flowers few and axillary, close to the branch tips. Pedicels slender, drooping, sparsely pilose, 24-35 mm long. Ovary oblong, 6-7 long, ca. 3 mm wide. Floral tube narrowly fun- nelform, 31-34 mm long, ca. 3.5 mm wide at the base, then gradually widened above until ca. 7 mm wide at the rim, subglabrous outside, pi- lose inside in lower 2. Sepals narrowly lanceo- late, apex narrowly acute to acuminate, 22-23 mm long, 4-5 mm wide, connate for 2-3 mm at the base, tips strongly spreading or recurved at anthesis. Tube and sepals magenta or bright red- dish pink. Nectary ca. 3 mm high. Filaments reddish, the antesepalous ones 13-20 mm long, the antepetalous ones 9-14 mm long; anthers narrowly oblong, 3-4 mm long, ca. 2 mm wide. Style reddish, pilose for most of length; stigma subclavate, barely 4-cleft at the apex, ca. 2 mm long, ca. 1.5 mm wide. Berry not seen. Distribution. Central Peru, known only from the type locality in cloud forest at the Carpish Divide, Dept. Huánuco, 2,750-2,950 m. Is known to flower from October to November. Figure 11. BERRY — FUCHSIA SECT. HEMSLEYELLA 235 Specimen examined. PERU. HUÁNUCO: Carpish, 9,000 ft., Sandeman 3480 (K). This species is allied to F. juntasensis, F. membranacea, and F. tillettiana, by virtue of its fruticose habit, opposite and narrowly ovate leaves, apparent lack of tubers, medium length flowers, and reflexed sepals. It is readily distin- guished from these species, however, by its firm- er, narrower leaves, with sparse pilose pubes- cence, narrowly lanceolate sepals, and disjunct distribution. Fuchsia huanucoensis is apparently very rare, as the Carpish area has been relatively well collected by botanists. 6. Fuchsia inflata Schulze-Menz, Notizbl. Bot. Gart. Berlin-Dahlem 15: 136. 1940. Fuchsia tuberosa Krause var. inflata (Schulze-Menz) Munz, Proc. Calif. Acad. Sci., Ser. 4, 25: 77. 1943. TYPE: Peru. Dept. Cuzco: Prov. Pau- cartambo, above Cosñipata, between Tam- bo Tres Cruces and Tambomayo, 3,100 m, 25 Apr. 1914, Weberbauer 6935 (holotype, B, destroyed in World War II; isotypes, F— 2 sheets, GH, POM, US—2 sheets). Figures 1-4, 14. Fuchsia macrantha sensu Munz, Proc. Calif. Acad. Sci., Ser. 4, 25: 83, pl. 14, fig. 73. 1943, pro parte. Shrubs 1-3 m high, usually growing among rocks, or epiphytic on trees, with well-developed, cylindrical, tan tubers 2-50 cm long and 2-4 cm thick. Branchlets terete, 2-5 mm thick, tan to ish, glabrous to densely cinereous or velu- tinous; older stems erect to divergent and 1-3 m long, 4-12 mm thick, with freely exfoliating bark. Leaves alternate or rarely opposite, membra- nous, narrowly elliptic to ovate, 55-135 mm long, 20-52 mm wide, apex acuminate, base acute to rounded or subcordate, glabrous to sparsely pu- bescent above, glabrous to pubescent along the veins — margin subentire to remotely den- pies condary veins 6—9 on either side of miedos Petioles glabrous to canescent-ve- ee 12-22(-35) mm long. Stipules filiform, .5-2 mm long, deciduous. Flowers axillary, one or few to numerous, usually densely clustered at branch tips. Pedicels pale green to reddish, 20- 55(—70) mm long, glabrous to densely pubescent. Ovary ellipsoid to narrowly cylindrical, 8-18 mm long, 2-3 mm wide, subglabrous to densely ci- nereous, sometimes 8-sulcate. Floral tube 85- 120 mm long, 5-9 mm wide and strongly bul- 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURE 14. Fuchsia inflata, flowering branch from Berry & Aronson 3036, Prov. La Convención, Dept. Cuzco, Peru; leaves from Vargas 23044, Prov. Calca, Dept. Cuzco, Pe bous at the base, strongly constricted to 2-4 mm wide above the nectary, then abruptly widened to (810-14 mm wide in the upper part of the tube, slightly constricted to 7-12 mm wide at the mouth, glabrous to densely canescent or pilulose outside, pilose inside on lower 3. Sepals narrow- ly triangular to lanceolate, 15-26 mm long, 6- 10 mm wide, apex acute, connate for 4-10 mm at the base, somewhat recurved at anthesis or divergent in lower '2 and suberect in upper !^. Tube (light) pink to orange, the base greenish or dull purple around the nectary, sepals light green to dull yellow-green. Nectary 3-5 mm high. Fil- aments green to pale yellow-green, the antesepal- ous ones 15-20 mm long, the antepetalous ones 8-16 mm long; anthers (narrowly) oblong, yel- low, 4-5 mm long, 2-3.5 mm wide. Style green to white-yellow, villous in lower 2 or for most of length; stigma green, clavate to globose, some- times tetragonous, 4-cleft at the apex, ca. 4 mm long, 2-3 mm wide. Young berries 20-40 mm long, 5-6 mm wide. Gametic chromosome num- ber, n = 11. Distribution. Dept. Cuzco, Peru, endemic to several valleys on the eastern slopes of the Andes, (2,350-)2,850-3,600 m. Flowers principally in the dry season, from June to August. Figure 11. Specimens examined. PERU. cuzco: Prov. Con- 2 Km 145-146 from Cuzco to Quillabamba, Berry & Berry 2567 (MO), 2568 (MO), 2569 (MO), 2570 (MO), Berry & Aronson 3036 (MO, USM), 3037 (MO), 3038 (MO, USM), Gentry 19830 (MO); Km 150 on Cuzco-Quillabamba road, Berry & Aronson 3040 (MO); Km 156 on Cuzco-Quillabamba road, Berry & 1985] & Berry 2572 (MO); Prov. rcd y ti 88-92 from Ocongate to Marcapata, Ber n 3010 (MO), 3011 (MO), 3012 (MO), jos JMO). 3016 ; (MO); across river from Marcapata, & Aronson 3023 (MO- 2 sheets); Quillabamba, Sia, Teresa, 0.5 km W of La Playa at Mandornilloc, Peyton & Tilney-Peyton ag ha Hirsch P1308 rgas 3595 (BH— 2 sheets); Prov. Calca, Amparaes, e 3273 doe Vargas 23044 (MO); Prov. Convención, Lucumayo Valley, Cook & Gilbert 1312 (US); Prov. Calca, near Lares, Vargas 3584 (BH, CUZ, MO), 3595 (BH, CUZ); onvención, Yanamache-Lucumayo, Vargas 4451 (CUZ —2 sheets); near town of Ma ta, Var- e , owering in summer of 1940, Univ. Calif. Botanical Garden Acces- sion N° 38.1052, from Dept. Cuzco, Peru (UC). A complex pattern of variation occurs in the long-tubed plants of sect. Hernsleyella that in- habit a series of adjacent valleys to the north, east, and west of the city of Cuzco, in southern Peru. An adequate understanding of the intra- and interpopulational variability of this group, here treated as belonging to one species, F. in- flata, has not been possible, because of the scar- city of these plants, their strong seasonality and an insufficient sampling of the populations in each of the valleys In general, the specimens treated here under Fuchsia inflata can be distinguished by their flo- ral tubes 8-12 cm long, with a bulbous base around the nectary, followed by a narrow con- striction before forming a tube that tapers slightly towards the rim. Unlike F. apetala and F. gar- leppiana, the only other species in the genus that can have such long-tubed flowers, the sepals of F. inflata are mostly green, and the leaves are much narrower than either of the other species. Furthermore, only F. apetala has knotty-tortu- ous stems, and F. garleppiana alone has pale pink flowers. Macbride (1941) and Munz (1943) treated specimens of F. inflata as long-tubed variants of F. tuberosa, for which both of them refer to plants presently recognized as F. chlo- roloba. Although F. chloroloba and F. inflata can occur sympatrically, and both have marked floral tube constrictions, the floral tubes of the former are quadrangular in cross section, and the tubes never exceed 55 mm in length, while the sepals are entirely parrot green and the tubers are small- er and more spherical than the elongate ones of F. inflata. The type collection of F. inflata is the only BERRY — FUCHSIA SECT. HEMSLEYELLA 237 specimen of this group collected in the Paucar- tambo Valley, which is in the central part of the range of the species; that specimen is glabrous throughout and bears both opposite and alter- nate leaves. Other collections agreeing with the made on July 21, 1978 along the Río Marcapata, 88—92 km from Ocongate, at elevations between 3,000 and 3,280 m, considerable differences in flower color, floral tube shape, and leaf size and shape were found among long-tubed plants most closely agreeing with F. inflata. Berry & Aronson 3010 and 3011 were found just meters apart at 3,280 m, in full sun among rocks and boulders above the river (see Fig. 4); 3010 had green sepals and light pink floral tubes sharply constricted at the base but broad above and was leafless, where- as 3011 had numerous large leaves, yellowish sepals and much narrower, less constricted floral tubes. Fifty meters downriver, Berry & Aronson 3013 had flowers similar in shape to 3010, but with salmon orange floral tubes. At 3,120 m, 1 km below 3013, an odd plant with floral tubes 50 mm long and with knotty stems similar to F. apetala was found, Berry & Aronson 3014. Just 20 m below, F. chloroloba appeared. Chromo- some counts were obtained from several of these plants, yielding n — ca. 22 for 3010, 2n — 44 and n = ca. 22 for 3011, 2n = 22 for 3014, and n = 11 for 3015. Anaphase I of pollen mother cells in 3011 showed meiotic irregularities such as bridges and chromosome fragments. It is unclear what is occurring in this area, but some hybrid- ization may have resulted among entities assign- able to F. inflata, F. chloroloba, and possibly F. apetala. The remaining specimens cited here under F. inflata come from valleys further to the north- west, in the Provinces Calca and La Convención (with a single collection from Prov. Urubamba). Although they all have leaves and floral tubes similar in size and shape to the plants described as “typical” for this species, they differ noticeably in their much longer, slender ovaries (to 4 cm long in fruit) and in the presence of a fine, ashy on leaves, stems, and flow- ers, including the interior of the sepals in some cases. In addition, both the ovaries and the bulb around the nectary are sometimes 8-ridged and purplish. As in the populations from Prov. Quis- picanchis, however, flower color varies from pink tubes with yellowish sepals (Berry & Berry 2567) canescent 238 to bright orange tubes with mostly light green sepals (Berry & Berry 2570, 2572, and 3036). Two collections examined from this area were both diploid (Berry & Aronson 3036 and 3040). Clearly, further cytological information from dif- ferent populations will be very useful in clari- fying the systematics of the F. inflata complex. 7. Fuchsia insignis Hemsley, J. Bot. 14: 69. 1876. TYPE: Ecuador. Provinces Cañar and Chim- borazo: in mossy forest on W slopes of Mt. Azuay, 23-26 Aug. 1859, Spruce 5976 (lec- totype, K — Bentham Herbarium, here des- ignated; isolectotypes, BM, CGE, F, G-BOIS, G-DC, K—Hooker Herbarium, LE, NY, OXF, P—2 sheets, W—3 sheets). Fuchsia apetala sensu Munz, Proc. Calif. Acad. Sci., Ser. 4, 25: 80, pl. 13, fig. 69. 1943, pro parte. Scandent or epiphytic vines or shrubs climbing high into trees, or terrestrial shrubs ca. 1 m high. Branchlets smooth, purplish, pilose, 2-5 mm thick, with prominent, alternate ie ae older stems freely exfoliating in strips, with stolonif- erous runners. Leaves alternate, absent when flowering, membranous, ovate to broadly ellip- tic-ovate, just two young leaves seen, these 45— 90 mm long, 30-40 mm wide, apex acute, base rounded, green and subglabrous above, pale green to purplish and villous below, especially along the veins; margin subentire to remotely dentic- ulate, secondary veins 6-7 on either side of the midvein. Petiole 1-1.5 mm thick, 9-18 mm long. Flowers few to numerous and clustered at the tips of new shoots. Pedicels pilose, 10-23 mm long. Ovary narrowly cylindrical, pilose, 8-10 mm long, 2-3 mm wide. Floral tube funnelform, (38-)40-54(-62) mm long, 4-6 mm wide and bulbous at the base, constricted to 2-2.5 mm in lower !^, then gradually widened until 7211 mm wide at the rim, pilose outsid d inside in lower ^. Sepals oblong-lanceolate, 24-37 mm long, 7- 12 mm wide, usually widest in the middle, apex acute to obtuse, connate for 3—4 mm at the base, strongly recurved at anthesis. Tube and sepals bright scarlet or orange-red. Nectary 4-6 mm high. Filaments green (fide Rose & Rose 22230), the antesepalous ones 30-40 mm wide, the an- tepetalous ones 25-35 mm long; anthers oblong, 4-5 mm long, 2-2.5 mm wide. Style densely re- trorse villous in the lower '4, exserted 10-15 mm yond the anthers. Stigma globose, ca. 2 mm long and wide. Immature berry narrowly cylin- drical, verrucose, 18-20 mm long, 5-7 mm wide. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 Distribution. Southern Ecuador; scarce epi- phytes in cloud forest or shrubs on hillsides in Provinces Bolívar, Chimborazo, and Cañar, 2,000-3,240 m. Flowers in the dry season when leafless, mostly from July to October. Figure 15. Specimens examined. ECUADOR. BOLÍVAR: Chapar- ro de Gualicón Loma, Cordillera Occidental, Acosta Solis 6267 (F). CAÑAR: between Cuenca and Huigra, Hitchcock 21701 (GH, NY, US). cHIMBORAZO: Si- bambe, Hda. La Carmela, Sección Era- Pata, Cord Occidental, Acosta Solis 5454 a ; n s.n. (K); Andes of Ecuador, 9,000 ft., Pearce s.n. (K), Lobb 112 (K). This species is best distinguished by its un- usually large, reflexed sepals and long filaments. It is strongly deciduous, but the presence of tu- bers in this species has not been noted. No col- lections of F. insignis have been made since 1943, probably a reflection of the destruction of large areas of natural habitat for this species, as well as a possibly brief flowering period and the in- accessibility of the remaining habitats. 8. Fuchsia juntasensis O. Kuntze, Revis. Gen. Pl. 3(2): 97. 1898. Munz, Proc. Calif. Acad. Sci., Ser. 4, 25: 77, pl. 12, fig. 67. 1943. TYPE: Bolivia. Dept. Cochabamba: between Co- chabamba and Río Juntas, E side of Cor- dillera, 3,000 m, Kuntze (lectotype, NY, here designated). Figure 16. Fuchsia o ie Contr. Gray Herb. 75: 38. 1925. TYP gn a. Dept. Cochabamba: Prov. Chapare, open w at Incachaca, 2,500 m, 22 Oct. 1920, Getic 5038 hoes NY, pho- tograph, UC; isotypes, A, G-DEL). Terrestrial or commonly epiphytic shrubs 0.5— 1.5 m tall, occasionally with small tubers. Young branches purplish, terete, 1-2 mm thick; older branches 4-20 mm thick with bark turning tan and exfoliating in strips or flakes. Leaves op- osite or less often ternate, membranous, (nar- rowly) ovate, 35-110 mm long, 15-50 mm wide, apex acuminate, base acute to rounded, glabrous to sparsely puberulent when young, often pur- plish below; margin subentire to serrulate, sec- ondary veins 5-7 on either side of the midvein. Petioles 5-9 mm long. Stipules triangular to lan- ceolate, 2-3 mm long, 1.5-2.5 mm wide, occa- sionally connate. Flowers few, axillary or grouped 1985] BERRY — FUCHSIA SECT. HEMSLEYELLA 239 300 Km. NN s OFuchsia membranacea e Fuchsia tillettiana Fuchsia pilaloensis e)Fuchsia insignis FIGURE 15. at branch tips. Pedicels slender, 20-75 mm long. Ovary oblong, greenish, 5-10 mm long, 2-3 mm wide. Floral tube funnelform, 25-50 mm long, 2-4 mm wide at the base, slightly constricted in the lower '4, then widened until 8-10 mm wide at the rim, glabrous outside, villous inside in the lower !^ above the nectary. Sepals ovate-lanceo- late, 14-25 mm long, 4-8 mm wide, apex acute to acuminate, connate for 3—4 mm at the base, divergent to recurred at anthesis. Tube and sepals deep lavender to rose violet. Nectary lustrous green to purplish, 4-5 mm high. Filaments pur- ple, the antesepalous ones 9-21 mm long, the antepetalous ones 6-16 mm long; anthers ob- long, yellow, 3-4.5 mm long, 1.5-2.5 mm wide. Style light purple, villous for most of its length; stigma green, 2.5-4 mm long, 2-3 mm wide, 4-cleft at apex, exserted 4-8 mm beyond the an- Distribution of the Ecuadorian and Venezuelan species of Fuchsia sect. Hemsleyella. thers. Berry oblong, purplish, 16-18 mm long, 8-10 mm wide (immature). Gametic chromo- some number, n = 22. Distribution. Bolivia, restricted to cloud for- est on the northeast slopes of the Andes in Dept. Cochabamba, 1,900-2,800 m. Flowers princi- pally in the dry season, from June to October. Figure 10. Specimens examined. BOLIVIA. COCHABAMBA: Valle de Montepunc nco, 1,900 m, Adolfo 380 (US), Valle de . from Cocha- Berry 3634 (MO); above ENDE pumping station at Corani, Berry 3638 (MO—2 sheets); Incachaca, Brooke 6666 (BM, F, NY, U), 6666a (BM, F, NY, U), 6666c Tablas, Cárdenas 4089 (F, GH, RSA): c cerros de In- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FiGunE 16. Fuchsia juntasensis, floral section and flowering branch from Berry & Berry 2587, Dept. Cocha- ne. "Bolivia: leaves from Berry 3638, Dept. Cochabamba, Bolivia cachaca, Steinbach 5738 (A, G-DEL, K); sys sy de Corani, Steinback 9862 (BM, F, G-BOIS, K, MO, NY , S, U, UC, Z). WITHOUT LOCALITY: Bridges $. n. (BM). This is one of the opposite-leaved species of sect. Hemsleyella with poorly developed tubers; it can be distinguished by its glabrous, mauve or violet-colored flowers, which generally have a lance-ovate sepals. Fuchsia juntasensis is known only from the yungas of Dept. Cochabamba in Bolivia, where it occurs as a terrestrial shrub or as an epiphyte. As Brooke (1953) noted, plants of this species found growing in the shade usually flower and produce leaves simultaneously. A single chromosome count of n — ca. 22 was obtained from Berry 3638; curiously, pollen of the same collection is entirely biporate (Prag- lowski et al., 1983), whereas most other tetra- ploid plants of Fuchsia examined show at least some triporate pollen grains. A possible inter- sectional hybrid between F. juntasensis and F. denticulata (sect. Fuchsia) is discussed in the sec- tion on Sympatry and Hybridization. 9. Fuchsia membranacea Hemsley, J. Bot. 14: 70. 1876. Munz, Proc. Calif. Acad. Sci., Ser. 1985] 4, 25: 78. 1943. TYPE: Venezuela. Edo. Méri- da: Venta de Mucuchíes, June 1842, Linden 372 (holotype, K; isotypes, BM, CGE G-BOIS, G-DC, MICH — fragment from G, MPU —2 sheets, OXF, P—2 sheets). Some labels incorrectly list Caracas, Venezuela as the type locality. Figure 17. » Terrestrial shrubs 1-2 m tall, sometimes climbing to 6 m above ground, growing in rocks, or epiphytic in trees to 10 m above ground; thick- ened roots to 20 mm thick or small tubers to 2 cm diam. sometimes present. Branchlets red-pink to dark purple, 2-5 mm thick, older stems 5-25 mm thick, with freely peeling, flaky, pale tan bark. Leaves opposite or ternate, sometimes sub- opposite, membranous, ovate-elliptic, 50-100 mm long, 18-50 mm wide, apex (sub-)acumi- nate, base rounded to acute, subglossy green and mostly glabrous above, pale green and subgla- brous below; margin subentire to denticulate, secondary veins 6—8 on either side of the mid- vein. Petiole 10-35 mm long. Stipules lance-tri- angular, 2-2.5 mm long, 1-2 mm wide. Flowers 2-12 per shoot, axillary or grouped toward the branch tips. Pedicels slender, usually green, 30- m ored. Floral tube funnelform, (30-)36-51 mm long, 2.5-4 mm wide at base, narrowed slightly in lower !^, then widened until 7-12 mm wide at the rim, mostly glabrous outside, pilose inside in lower !^. Sepals ovate-lanceolate 17-25 mm long, 5-11 mm wide, apex acute, connate for 5- 8 mm at base, lobes often split open before an- thesis with stigma exserted, spreading at anthe- sis. Tube red-pink, sepals usually pale green in upper !^. Nectary green, lustrous, 4-5 mm hi Filaments usually green, antesepalous ones 9- 15(-20) mm long, the antepetalous ones 7-11 (716) mm long; anthers oblong, yellow, 3.5—4.5 mm long, 2—3 wide. Style retrorse villous in low- er 14, exserted 10-15 mm beyond the anthers; stigma clavate, 2.5-3.5 mm long, ca. 2 mm wide, green. Berry cylindrical, 16-18 mm long, 11-12 mm wide, shiny light red when ripe, seeds mostly 60—70 per fruit, triangular-ovate in outline, ca. 2 mm long, 1.3-1.7 mm wide, light brown. Ga- metic chromosome number, n = 11. Distribution. Venezuelan Andes, in subpá- ramo and cloud forest in Estados Mérida and Trujillo, (2,600—)2,750—3,400 m. Flowers mostly in the dry season, from January to April. Figure 15. BERRY — FUCHSIA SECT. HEMSLEYELLA 241 Specimens examined. VENEZUELA. TRUJILLO: Gui- riguay, towards Pefias Blancas, Aristiguieta & Medina 3599 (NY, VEN); Páramo de la Cristalina, Jahn 12 pez-Figuieras 906 (MERF, MO). MERIDA: Páramo de Timotes, Alston 6622 (BM); Sierra Nevada, near La- guna Coromoto, Barclay & Juajibioy 9985 (RSA); Pá- ramo de Santo Domingo, Casca Victoria, Benítez de Rojas 1296 (MY); 5 km above El Rincón de La Venta, Timotes-El Aguila, Berry 3136 (MO, VEN); 36- 38 km from El Aguila to Piñango, Berry 3137 (MO, VEN), Luteyn et al. 6193 (NY); 24 km above Timotes on road to El Aguila, Berry 3278 (MO — 2 sheets, M. a Mesa de Los Pinos, above La Mucuy, Berry 3443 (MO, VEN), 3444 (MO), 3446-B(MO— 2 sheets), 3447 (MO); (MO); entrance to Hotel Los Frailes, Berry 3652 (MO); 73 km d Mérida, road to Barinas, Breteler 3354 US); between Mucubají and Sto. Domingo, de 22 939 (MER, MO, VEN), Dennis 2120 (K); 4 m from Laguna Mucubají towards Sto. Domingo near artificial lake, Foldats 2469 (VEN —2 sheets); Chacho- po, Funck & Schlim 798 (BM, CGE, F, G-DC, MPU, OXF, P); Páramo de Timotes, Jahn 508 (GH, US); Timotes, Karsten s.n. (Wy; Páramo de los Conejos, Lasser 511 (US, VEN); San Martín, 10 km S of San Rafael de Mucuchíes, Ruiz- Terán 992 (MER — 2 sheets); near Minigü, 10-15 km S of San Rafael de Mucuchíes, Ruiz-Terán 997 (MER, MO), 7208 (MO); between La- guna de Coromoto and Qda. El Oso, Ruiz-Terán & López-Palacios 1663 (MERF, MO); Laguna Brava, be- tween Río Los Granates and Loma La Paja, Páramo Los Granates, 15 km E of Sierra Nevada de Sto. Do- Le Ruiz-Terán 6281 (MERF, MO); Cafiada Cer- , between Chachopo and El Aguila, Ruiz- Terán & mui 12947 (MERF, MO); Las Tapias, from Apar- taderos to Barinas, Ruiz-Terán 13438 (MERF); near Laguna de Coromoto, Schulz et al. 326 (MER, U, VEN); above Chachopo, Steyermark 56751 (F, NY, VEN); La Aguada, Vareschi s.n. (MER) Fuchsia membranacea is an opposite-leaved member of sect. Hemsleyella: that inhabits upper cloud forest and p vege- tation. In both areas, it can be found as a liana, epiphyte or terrestrial shrub on rocks or in hu- mus. In habit and in many vegetative characters, it strongly resembles F. tillettiana, which has a forest, including the subpáramo, between 2,750 ,400 m, whereas F. tillettiana grows in cloud forest between 1,600 and 2,650 m. In F. mem- branacea, the filaments are i shorter sind greenish, nd dull pink to green, and the flowers are glabrous and axillary or grouped at the branch tips, but not subrace- mosely, as in F. tillettiana. Because of the ab- sence of morphologically or ecologically inter- 242 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 7. Fuchsia membranacea, flowering branch Em Berry 3465, Edo. Mérida, Venezuela; leaves and Teide a from Berry 3136, Edo. Mérida, Venezuel 1985] BERRY — FUCHSIA SECT. HEMSLEYELLA FIGURE 18. Fuchsia nana, flowering branch and tubers, from Berry & Berry 2581, Dept. Cochabamba, Bolivia. mediate plants, both taxa are recognized as distinct species. 10. rigen nana P. Berry, sp. nov. TYPE: Boliv- a. Dept. Cochabamba: Incachaca Pass, on Brooke 6920 (holotype, BM; isotypes, F, G-DEL, NY, U). Figure 18 Frutex nanus epiphyticus saxatilisve; caules erecti vel repentes 3-10 dm longi, stolonibus 2-8 dm longis et tuberibus numerosis lateralibus ellipsoidalibus cu- preis. Folia alterna saepe hysterantha, m ranacea, ovato-lanceolata, apice acuminata, basi rotunda Flores pauci axillaresque ad apicem ram juni ru infundibuliformes, 21-29(-31) mm longi, basi ca. mm lati bulbosique, superne dilatati summo 7-: lati. Sepala lanceolato-ovata, 12-14(-16) mm loni 5-8 mm lata, basi connata 2-3 m Erect to creeping subshrubs 3-10 dm tall in rocks, or epiphytic in trees, with stoloniferous stems 2-8 dm long and numerous ellipsoid, cop- per-colored tubers lateral to the stem. Branchlets glabrous to loosely pilose, terete, 1.5-3 mm thick, reddish to copper; older stems 2.5-4 mm thick, with freely exfoliating bark. Leaves alternate, usually absent when flowering, membranous, (narrowly) lance-ovate, 20-35 mm long, 10-20 mm wide, apex acuminate, base rounded, gla- brous above, usually pilose along the nerves be- f gm side ofthe midsein. Petioles 7211 mm long. Stip- ules filiform, 1.5-2 mm long, deciduous. Flowers few and axillary near the tips of young shoots. Pedicels slender, glabrous, 10-14(-19) mm long. Ovary cylindrical to ellipsoid, 7-7.5 mm long, 2-3(-4) mm wide, lustrous cherry red, glabrous to pilose. Floral tube funnelform, 21-29(-31) mm long, ca. 3 mm wide and more or less bulbous 244 at the base, constricted above the base to ca. 2 mm wide, then widened above until 7-8 mm wide at the mouth, glabrous to sparsely pilose outside, pilose inside in lower '2. Sepals lance- ovate, 12-214(-16) mm long, 5-7(-8) mm wide, apex acute to obtuse, connate for 2-3 mm at the base, divergent at anthesis. Tube and sepals bright red-pink, with a dark purple ring usually present around the mouth of the tube. Nectary high. Filaments red, the antesepalous ones 8-10 mm long, the antepetalous ones 6-8 mm long; anthers ellipsoid, cream, 2-2.5 mm long, 1-2 mm wide. Style red-pink, densely retrorse villous in lower !^; stigma green, globose, ca. 1.5 mm long and wide, exserted 5-10 mm beyond the anthers. Young berry ellipsoid, 11 mm long, 6 mm wide Distribution. Endemic to upper cloud forest in Dept. Cochabamba, Bolivia, (2,000-)3,000- 3,700 m. Flowering principally in the dry season, from (April to) July to November. Figure 10. Specimens examined. BOLIVIA. COCHAB 100 of Cochabamba-Villa Tunari road, Bade ock PT 3 (K); € and N of La apar — & Berry 2581 trict, Brooke 3 738 ( M grande, Cárdenas 4091 1. sr TRSA): Km between Colomi and truck terminal on way to xm Cárdenas s.n. (RSA). Fuchsia nana is readily recognized by the small size of the plant and its lustrous red-pink flowers with tubes just 2-3 cm long. It isa highly seasonal and almost exclusively epiphytic species, flow- the cloud forest. Both the high altitudinal range and the characters mentioned above would ap- pear to relate F. nana most closely to F. apetala. 11. Fuchsia pilaloensis P. Berry, sp. nov. TYPE: Ecuador. Prov. Cotopaxi: ] Holm-Nielsen & Jeppesen 1547 nee AAU; isotypes, DS, S). Figure 1 Frutex terrestris erectus usque 1 m altus vel epiphy- me ae anguste ovata, apice acuminata, basi ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 acuta vel rotundata, 40-122 mm longa, 25-54 mm lata, petiolis 15-38 mm longis. Flores pauci vel nu- merosi in ramulis axillaribus vel ad apicem ramorum aggregati, pedicellis 15-22 mm longis. Tubi florales i ene infundibuli formes, 44-53 mm longi, basi 3- lati ti bul bosique inde 2-3 mm LM constricti, m lati. la anguste ovata, 13-20 mm longa, Ss m lata, apice acuta, basi 4-7 mm connata, limbis sub anthes: valde patentibus Terrestrial shrubs 0.5-1 m tall or lianas or epiphytes with long, hanging branches to 8 m high in trees; tubers sometimes present or else thickened stems 10-20 mm diam. Branchlets te- rete, dull purple-green, densely pilulose, 2-4 mm thick; older branches reddish purple, freely ex- foliating. Leaves usually alternate, occasionally i bra i l 5-54 m wide, often unequal on either side of the margin, apex acu- minate, base acute to rounded, subglabrous to scattered pilose above, pilose below along nerves and margins; margin subentire to denticulate, secondary veins 6-9 on either side of the mid- vein. Petioles pubescent, 15-38 mm long. Stip- ules filiform, ca. 2 mm long, 0.2-0.4 mm wide, deciduous. Flowers few to many, grouped to- wards the tip of branches or axillary branchlets, sometimes subtended by reduced leaves. Pedi- cels pilose, usually pink, 15-22 mm long. Ovary cylindrical, 5-7 mm long, ca. 2 mm wide, dull purple, pubescent. Floral tube narrowly funnel- form, 44-53 mm long, 3-4.5 mm wide and bul- bous at the base, then narrowed to 2-3 mm wide in lower !^ and gradually widened above until 6-10 mm wide at the rim, densely pilose outside, pilose inside in lower '4. Sepals narrowly ovate, 13-20 mm long, 7-8 mm wide, apex acute, con- nate at base for 4—7 mm, lobes strongly spreading at anthesis. Tube pale pink; sepals whitish to pale pink, sometimes greenish in bud. Nectary green- purple, lustrous, 3-4 mm high. Filaments dull green, the antesepalous ones 12-15 mm long, the antepetalous ones 7-9 mm long; anthers oblong, yellow, 3-4 mm long, 2.5-3 mm wide. Style pale pink, densely retrorse villous in lower 1⁄2, ea , green, ca. 2 mm long, 2-2.5 mm wide, slightly 4- cleft at the apex. Berry 4-angled 1 maturity, oblong, verrucose, purplish; mature berry not seen Distribution. Known only from cloud forest above Pilaló, Prov. Cotopaxi, Ecuador, 2,400- 3,150 m. Known to flower in June and July. Figure 15. 1985] BERRY — FUCHSIA SECT. HEMSLEYELLA 245 FiGuRE 19. Fuchsia pilaloensis, flowering branch from Berry & Berry 2551, Prov. Cotopaxi, Ecuador. Specimens examined. ECUADOR. COTOPAXI: 10 km VATED: Berkeley, California, Univ. Calif. Botanical above Pilaló on Quevedo-Latacunga road, 3,150 m, Garden Accession N° 58.782- I, originally collected by Berry & Berry 2548 (MO, Q); 5 km above Pilaló, 2,760 C. K. Horich in Prov. Cotopaxi, Ecuador, between m, Berry & Berry 2551 (MO); 7 km above Pilaló, Albert ^ Pilalóand Macuchi, Hutchison, 1961 (RSA, UC), 1965 de Escobar & Escobar- Uribe 1479 (MO, TEX). curri- A). ` 246 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FiGunE 20. Fuchsia salicifolia, flowering branch from Vargas 3486, Prov. La Convención, Dept. Cuzco, This species is restricted to an area of western Ecuador where no other species of sect. Hemsley- ella is known, but where several species of sect. Fuchsia occur (Berry, 1982). Fuchsia pilaloensis is characterized by its light pink, narrow-tubed flowers, with pale sepals and narrowly ovate, mostly alternate leaves. In the same local area, this species has been found both as a small ter- restrial shrub with leaves as well as a profusely flowering climber or epiphyte; no tubers were detected in either case. 12. Fuchsia salicifolia Hemsley, J. Bot. 14: 70. 1876. Munz, Proc. Calif. Acad. Sci., Ser. 4, 25: 78. 1943. TYPE: Bolivia. Dept. La Paz: Prov. Nor Yungas, Sandillani, 7,000-8,000 ft., Apr. 1866, Pearce (holotype, K). Figure 20 Mir tuberosa Krause, Repert. Spec. Nov. Regni g. 1(11): 170. 1905. Fuchsia tuberosa Krause 1 if. Aca Puno: Prov. Sandia, between Sandia and Cuyo- cuyo, in rocks, 2,700-2,800 m, Weberbauer 875 (holotype, B, destroyed in World War II, photo- graph at F; isotype, G-DC). Epiphytic shrubs on tree trunks, occasionally growing in rocks. Stems sometimes stolonifer- ous, with globose-ellipsoid tubers 2-3 cm diam., the stems often growing down tree trunks for thic lightly pubesc te with freely tide bark, often 1-3 m long and hanging when epiphytic. Leaves alternate, firmly membranous, (narrowly) lanceolate, 60— 135(-160) mm long, 18-37(—40) mm wide, apex generally long acuminate, base acute to rounded and often unequal, pale green and subglabrous above, light green to purplish and subglabrous below; margin remotely paa: secondary veins 7-9 on either side of the midvein. Petioles 5-20(-30) mm long. Stipules um d m id m long. Ovary (narrowly) cylindrical, 8-11 mm long, ca. 1985] 2.5 mm wide, dull green, puberulent. Floral tube narrowly funnelform, 40-54(—60) mm long, 3- 4(-5) mm wide and more or less bulbous at the base, then narrowed to 1.5-2(-2.5) mm in the lower '^ and gradually widened above until 7- 10 mm wide at the rim, glabrous to puberulent outside, villous inside in lower !^. Sepals nar- rowly ovate, 23-30 mm long, 6-10 mm wide, apex acute to subacuminate, base connate for 2- 4(—5) mm, n, spreading- divergent : anthesis, the distal p ack. Tube red- pink, sepals similar or paler, a Pun purple ring usually present at the throat of the tube. Nectary 3-4.5 mm high. F ilaments red to dark purple, the antesepalous ones 15-26 mm long, the an- tepetalous ones 12-20 mm long; anthers oblong, yellow, 2.5-3.5 mm long, 1-1.5 mm wide. Style pink, retrorse villous in lower 1; stigma globose to subclavate, dull green, ca. 2.5 mm long, 1.5- 2 mm wide, exserted 6-12 mm beyond the an- thers. Mature berry not seen. Distribution. Southern Peru (Depts. Cuzco and Puno) and Dept. La Paz, Bolivia, in cloud forest, (2,000—)2,500-2,900 m. Flowers in the dry season, principally from July to October. Fig- ure 10. r air pua PERU. cuzco: La Conven- , 10 km SW of Incatambo, Peyton & do, La Con- omon 9683 (MO). WITHOUT LOCALITY: Pearce 799 (K). This species is best distinguished by its nar- rowly lanceolate, unequal leaves with long, acu- minate tips and by its pink, narrowly funnelform floral tubes. Most specimens have leaves present together with flowers, but Berry & Berry 2578 was almost leafless when collected. Close examination of the isotype and the pho- tograph of the destroyed holotype of F. tuberosa reveals that it is not specifically different from F. salicifolia, whereas Munz (1943), who did not have the opportunity to examine the type ma- terial, mistook F. tuberosa for what we now rec- ognize as F. chloroloba, which also has tubers, but has orange-red and green flowers, a sharply constricted floral tube and entirely glabrous leaves and flowers BERRY — FUCHSIA SECT. HEMSLEYELLA 247 13. Fuchsia tillettiana Munz, Aliso 7: 410. 1972. TYPE: Venezuela 1,700 m, 24 Mar. 1970, Tillett 703- 35 (ho- lotype, RSA-226553; isotypes, GH, MO, NY, US). Figure 21. Fuchsia pars sensu Munz, Proc. Calif. Acad. Sci., Ser. 5: 80. 1943, pro parte. Terrestrial shrubs 0.5—2 m tall, or growing in rocks, climbing up trees or epiphytic in trees to 10 m above ground; small, lateral tubers occa- sionally present in the soil on large plants. Branchlets puberulent, terete, 2-5 mm thick, red or copper-colored; older branches ascending on shrubs, flexuous and erect to drooping and to 10 m long on hanging bushes, the rooted portions often to 25 mm thick, with freely peeling, flaky bark. Leaves opposite or ternate, soft membra- nous, elliptic to narrowly ovate, (30-)55-100 mm long, (7-2)16—-50 mm wide, apex (sub-)acuminate, base rounded to acute, subglabrous to pruinous or puberulent above, subglabrous to puberulent and pale green below; margin subentire to ser- rulate, secondary veins 7—9 on either side of the midvein. Petioles puberulent, green or reddish, 6—12(—30) mm long. Stipules lance-triangular, 1— 2 mm long, 1-1.5 mm wide. Flowers generally numerous at branch tips or subracemose; leaves often absent, or small bracts 15-30 mm long and 6-20 mm wide present when flowering. Pedicels red-purple, 25-50 mm long. Ovary cylindrical, more or less sulcate, 8-10 mm long, 2.5-3.5 mm wide, green to purplish, usually with glandular hairs. Floral tube funnelform, lower !^, then gradually widened until 7-9(-13) mm wide at the rim, glandular-pubescent out- side, pilose inside in lower '4. Sepals lanceolate, 19-33 mm long, 4—6(-10) mm wide, apex acute, base connate for 2-3 mm, blunt-tipped in bud, strongly recurved at anthesis. Tube and sepals cerise or rose pink. Nectary glossy green, 3-4 mm high. Filaments pink to cerise or dark purple, the antesepalous ones 20-35 mm long, the antepetal- ous ones 15-28 mm long; anthers oblong, yel- low, 4-5 mm long, 2.5-3 mm wide. Style pink, retrorse villous in lower !5-!^, exserted 15-20 mm beyond the anthers; stigma green, clavate, ca. 3 mm long, 2-2.5 mm wide. Berry cylindrical, usually more or less narrowed at the apex, 18- 21 mm long, 6-10 mm wide, puberulent, dull purple-green to lustrous bright red when ripe; seeds ca. 50 per fruit, light tan, 1.8-2 mm long, 248 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURE 21. Fuchsia tillettiana, flowering branch from Berry 3463, Edo. Mérida, Venezuela; leaves and stem portion from Berry 3265, Edo. Lara, Venezuela. 1985] 1.3-1.6 mm wide. Gametic chromosome num- ber, n= 11. Distribution. Venezuelan Andes, cloud forest in Estados Lara, Trujillo, Mérida, and Tachira, 1,600-2,650 m; one doubtful collection from Co- lombia. Flowers mainly in the dry season when leafless, from January to April. Figure 15. Specimens examined. | VENEZUELA. LARA: Parque Nacional Yacambi, 13-17 km S of Sanare, Berry 3265 (MO, VEN), 3267A (MO), 3267 B (MO), 3467 (MO), 3469 (MO), Dunsterville & Dunsterville, 1969 (VEN), Garófolo, 1979 (MO), Steyermark & Carrefio 108848 , VEN) Tillett 711-26 (MER), 714-41 (MER). TRUJILLO: La Mesa de Esnujaque, Box & Alayón 3710 (BM, VEN). MÉRIDA: 1.5 km above San José, road to Mucutuy, Berry 3463 (MO, VEN), 4280 (MO, VEN); forest above La Veguilla, above Mucutuy, d 3464 (MO, VEN), 4289 (MO, VEN); banks of Qda. La Cam- San José, Jahn 969 (US, VEN); Páramo de Mucu- chachí, Jahn 981 (VEN); Páramo de Aricagua, Jahn 030 (US); Mucutaray, between San Isidro and Qda Mucubají, more or less 15 km ESE of El Morro, Ruiz- Terán & Lopez-Figuieras 8603 (MERF, MO); Páramo de Pozo Negro, between San José and Beguilla, Stey- ermark 56293 (F, NY, VEN). TACHIRA: Dtto. Carden- as, 15 km E of Zumbador on road to Queniquea, Berry 3303 (MO—2 sheets, VEN). One of six opposite-leaved species in sect. Hemsleyella, Fuchsia tillettiana is most readily recognized by its glandular-pubescent floral tubes (best seen when fresh), strongly reflexed sepals, long filaments (20-35 mm long in the antesepal- ous whorl) and the cerise color in both the tube and the sepals. The flowers of F. tillettiana are grouped subracemosely at the branch tips, and anthesis usually occurs when the plant is leafless, although shade plants often bear leaves and flow- ers together. Tubers seldom occur on plants of F. tillettiana, but thick semisucculent stems seem to supplant the storage function of tubers in climbing and epiphytic plants. Woody, erect stems predominate when the species occurs as a terrestrial shrub, but when growing as a climber, plants produce long, flexuous stems that climb up to 10 m high in trees and then clamber or droop down over dne adjacent VEBetaHon. Fuchsia tilletti occupies tely same lid range as F. membranacea, to which it is closely related; their main differences are discussed under F. membranacea A specimen of somewhat doubtful origin, Dawe 896 (K, Colombia, in the páramos, tuber- ous), lacks leaves, but has pubescent flowers with the BERRY — FUCHSIA SECT. HEMSLEYELLA 249 narrow, though not evidently reflexed, sepals that agree most closely with F. tillettiana. The no tation of the presence of tubers and of inhabiting páramos casts doubt, however, on it belonging to this species. More complete specimens with detailed locality information would be necessary to evaluate the proper placement of this collec- tion. 14. Fuchsia tunariensis O. Kuntze, Revis. Gen. Pl. 3(2): 98. 1898. Munz, Proc. Calif. Acad. Sci., Ser. 4, 25: 79, pl. 13, fig. 68. 1943. TYPE: Bolivia. Dept. Cochabamba: Tunari Range, 3,000 m, Apr.—May 1892, Kuntze (holotype, NY, photograph at UC). The type collection is a very poor specimen with no flowers left, but the leaves agree well with the type de- scription. Figure 22. P mattoana Krause, Bot. Jahrb. Syst. 37: 599. 906 PE: Peru. De x road from Cuz June 1906, Weberbauer 4976 (holotype, B, de- stroyed in World War II, photograph at F). Shrub 0.8-1.5 m high, growing terrestrially in humus, on rocks, or epiphytically in trees, with cylindrical-ellipsoid tubers 1—3 cm thick and 8- 15 cm long usually present. Young growth ve- lutinous, pilulose or cinereous; branchlets terete, 2—4 mm thick, pilulose; older branches with free- ly exfoliating bark. Leaves opposite or less often alternate, membranous, broadly ovate to elliptic- ovate, 60-110(-115) mm long, 35-65 mm wide, apex (sub- Jacuminate, base acute to cordate, , scattered to densely pubescent below, especially along the nerves; margin remotely denticulate, secondary veins 6— 9 on either side of midvein. Petioles pubescent, (10-)15-33 mm long. Stipules narrowly trian- gular, ca. 1.5 mm long, deciduous. Flowers gen- erally few and axillary towards young branch tips. Pedicels finely pubescent, 15-30(—50) mm long. Ovary cylindrical, velutinous to pilulose, 9-11 mm long, wide. Floral tube funnel- form, (36-)44—56 mm long, 3.5-4.5 mm wide and more or less bulbous at the base, then slightly constricted to ca. 3 mm wide and widened grad- ually above until 6-10 mm wide at the rim, finely pubescent outside, villous inside for most of length. Sepals lanceolate to (narrowly) ovate, 17— 25 mm long, 7-8(-9) mm wide, apex acute to acuminate, connate for 3-4 mm at the base, di- vergent at anthesis. Tube and sepals bright red- pink to light orange. Nectary 2.5-4 mm high. 250 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FiGuRE 22. Fuchsia tunariensis, flowering branch and leaves from Berry 3051, Dept. Ayacucho, Peru. 1985] Filaments greenish, the antesepalous ones 14-20 mm long, the antepetalous ones 8-13 mm long; anthers oblong-ellipsoid, yellow, 3-3.5 mm long, ca. 2 mm wide. Style orange to pink, villous for most of length; stigma globose, green, 2-3 mm long, ca. 2 mm wide, exserted 6-20 mm beyond the stamens. Mature berry not seen. Gametic chromosome number, n = 11 Distribution. Southern Peru and Bolivia. Rare in cloud forest from Dept. Ayacucho, Peru, to Dept. Cochabamba, Bolivia, 2,400-3,400 m. Flowers mostly at the end ofthe dry season, from September to November. Figure 10. Specimens examined. PERU. AYACUCHO: 45 km E of Tambo on road to Ayna, Berry 3051 (MO). cuzco: F); Prov. Convención, “El Villa Tunari road, Badcock 31 2 (K); Choro, above Río Cocopata, 100 mi. NW of Cochabamba, Brooke 6151 (BM, 3 6247 (BM) road to Incachaca, Brooke 6809 M, F of Cochabamba-Chapare road, Cárdenas 5 969 a UC, US); Km 102 of Cochabam- ba-Aguirre-Todos Santos road, between Comercocha and n Ugent & Ugent 5060 (DS, GH, K) This species is best distinguished by its mostly opposite leaves and the soft, velutinous-cinere- ous pubescence on the leaves, flowers, and you ng shoots. The fl tly ored, typically rose salmon to light orange, al- though the sepals may be slightly greenish at the apex. Many specimens bear leaves and flowers at the same time, and although the plants are characteristically tuberous, they can grow in a variety of situations— as epiphytes, or as terres- trial shrubs in rocks or in humus soil. Since this variety of habitats can give rise to considerable vegetative variation, and there are so few col- lections made of this species, with a considerable gap between the Bolivian and the Peruvian col- lection localities, more material is needed to de- termine if the plants from these two areas are really conspecific. A single collection from Dept. Huánuco, Peru, is possibly a new species, most closely allied to F. tunariensis: Meza 360 (MO, Peru, Dept. & Prov. Huánuco, Quebrada before Utao, 2,430 m, 4 Oct. 1965, shrub with pink flowers and greenish sepal tips) has opposite to ternate, ovate leaves, canescent-pilose pubescence, erect stems, pilose floral tubes 40 mm long (in bud) and sepals 12 mm long by 4 mm wide. The sepals of this unifor mly col- BERRY — FUCHSIA SECT. HEMSLEYELLA 251 plant, though still immature, are too small and contrast too strongly in color with the floral tube to be placed in F. tunariensis, but mature flow- ering specimens would be desirable before de- scribing it as a new species of sect. Hemsleyella. LITERATURE CITED Berry, P. E. 1982. The systematics and evolution of ia sect. Fuchsia (Onagraceae). Ann. Mis- d. 69: 1-198. BREEDLOVE, D. E. 69. The systematics of Fuchsia sect ly (Onagraceae). Univ. Calif. Publ. Bot. zh . E. RR H. RAvEN. 1982. The Mex- ican and Central American — of Fuchsia (On- agraceae) except sect. Encliandra. Ann. Missouri Bot. Gard. 69: 209-234. Brooke, W. M. A. 1953. Bolivian fuchsias. Gard. Chron. 133: 220-221. 1982. Fuchsia Lexicon. Van Nostrand HEMsLEY, W. B. 1876. The apetalous fuchsias of South America, with descriptions of four species. J. Bot. 14: 67-70. Hickey, L. J. Classification of the architecture of "oie dodo leaves. Amer. J. Bot. 60: 33. Hooker, W. 1846. Fuchsia macrantha. Large-flow- ered apetalous Fuchsia. Bot. Mag. 72: t. 4233. Hourte. L. vAN. 1848. Culture du Fuchsia macran- a. Fl. KURABAYASHI, M., H. Lewis & P. H. RAVEN. 1962. spei study of mitosis in the Onagraceae. Bot. 2 1003-1026. MACNEIDE, J. F. 1941. Onagraceae. In Flora of Peru. i us. Nat. nd Bot. Ser. 13: 521-566. ME P. A. 1943. A revision of the genus Fuchsia (Onagraceae). Proc. Calif. Acad. Sci., Ser. 4, 25: 974. Onagraceae. In Flora of Ecuador. Op- era ) Bot., Ser. , 3: 3-46. NowICKE, J. W J. SKVARLA, P. H. RAVEN & P. E. BERRY. 1984. A palynological study oft the genus : erue A aiaia Ann. Missouri. Bot. Gard. ml ja J. J. SKVARLA, P. H. RAVEN & J. W NOWICKE. 1983. Onagraceae Juss. Fuchsieae L./ : ussiaeeae L. World Pollen and Spore Flora 12: —4]. ms P. H. 1975. The bases of angiosperm phy- en CENE: Ann. Missouri Bot. Gard. 62: as Ruiz, H. & A PAVÓN. 1802. Flora Peruviana et Chi- lensis 3: 1-95. Madrid. SKVARLA, J. J., P. H. RAVEN & J. PRAGLOWSKI. 1976. The Evolutionary Significance of the Exine. Aca- demic Press, London . F. CHIssoE & M. SHARP. 1978. An ultrastructural study of viscin threads in On- agraceae n n. Pollen & Spo vnd tan 143. WRIGHT, J. O. Fuchsia “ es," a summary of those in A c Fuchsia oui 41: 64-67. COLLECTING AND PREPARING SPECIMENS OF ARACEAE THOMAS B. CROAT! The family Araceae is a conspicuous compo- nent of moist to very wet life zones in the tropics. Because of the frequently succulent and often large parts, the preparation of specimens is often difficult. A detailed discussion of all aspects to be considered in preparing descriptions of An- thurium (applicable as well to most other genera) was published for use by aroid workers (Croat & Bunting, 1979). This paper will deal only with those aspects which should be recorded when specimens are being prepared in the field. Since the family is well suited to cultivation, and since an increasingly higher percentage of the collec- tions are also being collected and sent back alive, this paper will be divided into two parts: 1. her- barium specimens; 2. live collections. HERBARIUM SPECIMENS AND FIELD NOTES ECOLOGICAL ASPECTS OF ARACEAE The diversity of habit types in Araceae is much greater than in most families (Simmonds, 1950; Madison, 1978) and special attention should be given to aspects of habit. Some members of the family, such as Caladium, Chlorospatha, Dief- fenbachia, Spathiphyllum, and Xanthosoma are strictly terrestrial. Most temperate species such as Arisaema, and of course all tuberous members of subfamilies Lasioideae (e.g., Dracontium, Amorphophallus, etc.) and Aroideae, are also ter- restrial. Quite a number of aroids are rooted aquatics in standing water, including Orontium, Montrichardia, Urospatha, and other genera. saeco’ ds Spathiphyllum, and others occur in marshy habits or sometimes in standing water. Some Spathiphyllum and Anthurium species oc- cur only on rocks along streams. Riparian species are particularly common in southwest Asia with genera such as Holochlamys, Piptospatha, and Aridarum frequenting such habitats. Pistia is a free-floating aquatic and Jasarum a rooted but wholly immersed aquatic. The largest percent- ages of aroid species are, however, either epi- petric or epiphytic in one form or another. The types of epiphytism are diverse with some genera such as Anthurium and Stenospermation usually being true epiphytes while other genera like Monstera, Syngonium, and most Philodendron begin their life in the soil then eventually climb trees while becoming hemiepiphytes and at least in part often losing their connection with the ground to become true epiphytes. Some Philo- dendron, such as P. wendlandii Schott, have seeds which germinate only on tree branches and are thus true epiphytes. Still others such as P. ra- diatum Schott grow m as tiig ie S until they eventually at which time they must be classified as hemiepiphytes. Many genera pass most of their adult lives slowly creeping up tree trunks. These are referred plants in certain genera, notably Monstera and Syngonium, can revert again and again to juve- nile forms owing to being dislodged from their host plant or merely because they have run out of space to climb. These reversions have been studied for Monstera (Madison, 1977) and es- pecially for Syngonium (Ray, 1981; Croat, 1982). Leaf forms for juvenile plants, pre-adult plants (usually rouna down low on tree trunks), and 11 4 4 tal ly distinct and ma y even seem to be distinct species. If possible, one should collect juvenile and pre-adult forms. The habit of Philodendron is particularly di- verse with, for example, some species occurring only over the crowns of understory shrubs while others occur only on the lower portions of larger tree trunks and still others grow only high in the canopy where more light is available. To merely refer to all of these as epiphytes (the usual case) or epiphytic vines (in fact most are hemiepiphy- tic vines) does little justice to a complex array oflife forms and habits. Itis thus important when making aroid collections to make note of the habitat and habit of the collection. The habit of the collection is particularly important. IMPORTANT MORPHOLOGICAL FEATURES Since it is often not possible to collect the whole plant, certain morphological features are impor- ! Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. ANN. Missouni Bor. GARD. 72: 252-258. 1985. 1985] TABLE 1. Synopsis of features to note for certain genera of Araceae Anthurium Habit; int ngth; petiole cross-sectional shape; qiapou Dan of inflorescence; color and disposition d shape of fruits. Doui Height of plant; diameter, color and glossiness yasqa or glossy) "E stem; cross-sectional shape, color a fleaves; shape of unen midrib. Homalomena -f 1 2 ] var iegation ofleaves. Monstera Habit; length, coloration, and markings of inter- nodes; collect juvenile and pre-adult leaves if pos- sible; color of spathe and spadix. Philodendron Habit; internode length and color, Sony decid- s or persistent, } sectional shape of petiole; shape of midrib c on up- per surface; prominence of minor veins on lower surface; color of sap; number of inflorescences per axil; color of spathe (inside and outside). Rhodospatha Habit; internode length; color of spathe and spadix. Spathiphyllum Habitat; degree of clustering; spathe and spadix col- or; color of tepals; color and degree of protrudence of pistils. apom abit; int de length; collect j ile and pre-adult leaves if possible: number ‘of inflorescences per Xanthosoma Spathe tube color (inside and outside); color of fe- male spadix tant to note. These are summarized for certain important genera in Table Ste. Numerous usan characteristics tant. If it is not possible to collect the stem either because of its size or because the stem is being used as a live specimen, one should make note ofthe length and width ofthe internodes. In most climbing genera even one or two internodes in- cluded with the specimen yields considerable in- CROAT- COLLECTING ARACEAE FIGURE 1. Stem with several nodes, cataphyll and both Autos of blade. formation because many species have unique colors on drying or are fissured in characteristic manners. The stems of Philodendron and other scandent genera may be variously flattened on one side or variously sulcate providing useful taxonomic characteristics. Though most of the figures in this paper lack stems because the stems were taken for cultivation, it is always best to include the stem if it is not to be used. On large stems it is best to cut off a small portion of the apex of the stem, then split the stem longitudinally (or even remove all but a thin portion of the outer part of the stem so that the uppermost cataphyll is still intact). If possible and the size of the petiole does not prohibit it, the uppermost petiole should be left with the Philodendron, a large and taxo- nomically complex genus, has cataphylls that provide several important taxonomic character- istics. Cataphylls are bract-like structures at least partially encircling the stem, at or near the apex, and subtending each new leaf for its protection as it emerges. They may be present or absent (in 254 FIGURE 2. One-half of large end with midrib and beak “folded to show base and a the case of some entire sections of the genus, e.g., sect. Pteromischum, where protection of the new leaf is accomplished by a sheathing petiole of an earlier leaf) and may be variously colored. Cata- phylls may also be unribbed, 1 -ribbed, or 2-ribbed and at the same time they may be deciduous or variously persistent (either remaining intact or more usually weathering to a mass of persistent fibers which envelop the stem). At least the apical portion of ih stem is thus essential for an ade- quate specim Petioles. Fm ole cross-sectional shape on many genera including both Philodendron and Anthurium is one ofthe most valuable taxonom- ic characteristics and should be noted. Because this feature is so variable and because a typical shape has been described so differently by dif- ferent collectors, an illustration of the various petiole shapes along with descriptive terminol- ogy is included (Fig. 9). Blades. When blades are small, it is best not to fold the blade but to prepare the specimen with the lower surface up (Fig. 1). The lower surface, because of the generally more pro- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 K Ac WH MANA FIGURE 3. Large blade with apex folded back yllateral veins and (slightly to the sinus). nounced venation, provides more characters. On a larger blade it is best to slice off one-half of the blade leaving the midrib and petiole with the half which is to be saved (blades of Araceae are gen- erally bilaterally symmetrical but where not, this technique should be replaced by making one or more sections of the blade perpendicular to the midrib). The half leaf can then be folded accor- dian style to fit a standard sized newspaper for specimen preparation (Fig. 2). It is important to start at the base of the blade with the lower sur- face facing upward, then fold the apicallobe over so that it is directed slightly to one side so the midrib and the sinus (space between the poste- rior lobes) is still visible (Fig. 3). When the blade is particularly long, two or more folds can be made, but particularly large blades should be cut in half perpendicular to the midrib and mounted on two sheets (Figs. 4, 5). When blades are wide and the basal portion will not fit on a standard sized herbarium sheet, the lower portion can be placed on the sheet sideways (Fig. 6) with the remainder folded on a second sheet. If a speci- 1985] FIGURES 4, 5. men is of moderate size, the petiole, stem, and inflorescence can be accommodated on the same sheet. After folding the blade (usually only one can be accommodated) in the manner described above, the petiole, section of stem apex, and the inflorescence (usually in one of uppermost axils) can be folded over the leaf. This method can at least be used for initial field drying, then if the value of the collection warrants it, the collection can later be cut along the fold lines and mounted on separate sheets. Inflorescence. Because of the unusual nature of flower morphology, some discussion is war- ranted (see Croat & Bunting, 1979, for a more detailed treatment). Araceae flowers are invariably quite small and are borne in congested spikes (or spadices) which are, at least initially, always enveloped by a con- spicuous spathe. Flowers may be unisexual (in which case plants are usually monoecious) or bisexual. When they are unisexual, the female (pistillate) flowers are always borne on the lower part of the spadix and mature first (aroids are all believed to be protogynous). The male (stami- nate) flowers are borne on the usually much long- CROAT—COLLECTING ARACEAE Blade cut perpendicular to midrib and placed on two sheets. e. er upper part of the spadix. The lower part of the staminate spadix often bears a small section of sterile male flowers. A few genera in the old world, e.g., Alocasia and Amorphophallus, bear an elongated, often conspicuous, appendage at the apex above the male spadix. In the case of unisexual flowers, the respective sections of the spadix may be of different colors and these should be noted. Bisexual flowers are borne in uniform spadices and are usually of uniform color though the tepals may be a different color than the pistils. Color changes may also occur after anthesis and sometimes it is possible to note these by observ- ing several e on a plant or several plants in a population. The spathe dem is taxonomically important and should be noted. Like the spadix, color changes often occur after anthesis ad if so, these should be noted. Species with convolute spathes (such as Philodendron) may have the spathe col- ored differently on the inner surface than on the outer surface. These important differences should be noted. Since many genera may have massive inflorescences or infructescences, it is often wise to dissect the spadix longitudinally for better ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURE 6. Lower portion of blade placed sideways on sheet to accommodate broad blade (upper half of blade dried in a separate sheet). drying. Genera with convolute, closed spathes pose special problems since the spadix enclosed in them exhibits important characters which must another to expose the spadix. One method is to remove the spathe completely by cutting around the spathe just above its attachment with the peduncle (Fig. 7). Another method is to cut a large window in the spathe at the base to expose the pistillate portion and at least the base of the staminate pattern (Fig. 8). It is important to make note of any differences in coloration of the male and female portions, because in many cases they are not the same. Fruit. Color and shape of fruits are taxonom- ically important and should is ial accu- rately since even where live material exists, it is often not possible to — pollinate the plant and thus study the fruits. Fruits usually are solid colored but are often bicolorous with the lower portions usually paler. Recording the shape of the berries is important because drying greatly distorts them. FIGURE 7. Spathe removed by cutting the base near peduncle attachment and by pulling the two sections apa PREPARATION AND SHIPMENT OF IVING MATERIAL Most Araceae are either epiphytic or have short internodes and are easily shipped live. Generally the most difficult to ship live are slender stemmed vines especially those with long internodes such as Heteropsis or Philodendron (sect. Ptero- mischum). Generally a live specimen should con- sist of at least three nodes (preferably more). The most -a psp aa e are usu- ally about 15-20 c ng . Old cata- phylls, as cell as alii leaves and den should be removed as they usually perish during shipment. Generally dirt and debris can be removed by wiping the stems with your hands or a rag and a knife can be used to scrape off more difficult to remove dirt. Washing with a brush in clean moving water is good but all excess water must be moned on or stems must be allowed to air g to prevent mold. Individual cuttings should be tagged first then wrapped in small pieces of newspaper. The cuttings should be gathered into plastic bags, tightly sealed, and 1985] Spathe cut to expose portion of spadix rile staminate E 8. Spat eh lower pistillate portion, steri flowers and the lower half of the staminate spadix). kept in a shady, preferably cool, place until shipped Aluminum tags that can be written on with a ballpoint pen are excellent and are designed to wrap around the stem. Masking tape wrapped around the stem also persists well but should be marked with an indelible marker. Paper tags often fall off before they arrive. Never attempt to ship stems with the tag attached to a petiole or around the tip of the stem as petioles are soon deciduous and tags often fall off of the generally pointed tip of the plant. For additional information con- cerning the availability of different types of tags, see Croat (1 Generally, the simplest and cheapest way to send back live material is to collect fruits and prepare the seeds for shipment. Aroid fruits are invariably fleshy and the seeds must be removed from the berry to meet importation require- ments. This is easily done by careful maceration in a bowl. If the resulting material is flushed with water, the pulpy mesocarp usually floats to the surface and can be removed by decantation. The seeds are then wrapped in a small piece of dry CROAT—COLLECTING ARACEAE CROSS-SECTIONAL PETIOLE SHAPES IN ANTHURIUM A. Basically terete 1 2 3 4 5 6 7 8 9 10 11 B. D-shaped or broader than thick 1 2 3 4 5 6 7 C. U-shaped or thicker than broad 1 2 3 4 5 6 D. Markedly angular 1 2 3 4 5 6 7 8 E. Markedly ribbed abaxially vd A Cross-sectional petiole shapes in An- y terete: ranging from esulcate and inr su ws = (4), ton cate (5), to narrowly and a arbi sulca and acutely secon (7), shallowly sulcate (8), flat adax- ially (9), flat adaxially with marginal ribs (10), to flat adaxially with marginal and medial ribs (11).—B. D- shaped or broader than thick: ranging from flat adax- ially € obtuse margins (1), to n sulcate with rgins (2), broadly sulcate wi acutely sulcate wi margins* (3), shallowly and P sulcate* (4), sulcate with acute margins (5), go with sharp margins and a medial rib* (6).— wi th acute eds (D, to ) quadrangular with obtuse angles (2), trap- ezoidal (3), obtusely triangular (4), Bde triangular (5), acutely triangular with two marginal ribs (6), ba- sically triangular with two abaxial ribs (7), obtusely triangular, narrowly and sharply sulcate with convex sides (8). — E. Markedly ribbed abaxially: ranging from trapezoidal or quadrangular, sharply and broadly sul- cate adaxially, 3-ribbed abaxially (1), to obtusely and broadly sulcate adaxially, 3-ribbed abaxially (2), broadly and obtusely sulcate adaxially, 5-ribbed abaxially (3), broadly and sharply | sulcate adaxially, narrowly and 1 Or wi with one or more ribs on the sulcus (5). *Not yet observed but to be expected. 258 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 newspaper or toweling, marked, and placed in a sealed plastic bag before being mailed. If there is danger they will be smashed, extra toweling or newspaper could be placed on the outside of the plastic bag or a padded envelope could be used LITERATURE CITED ico T. B. 1982. A revision of eura ~ ae). Ann. espe Bot. Gard. 68: 565-65 1 mportance of labeling living eus Aroideana 7: 27- 30. & G. S. BUNTING. 1979. Standardization of Anthurium descriptions. Aroideana 2: 15-2 MADISON, M. 1977. A revision of Monstera (Ara- ceae). Contr. Gray Herb. 207: 3-1 The genera of Araceae in the northern Andes. Aroideana 1: 31-53. Ray, T. S., JR. 1981. Growth and heterophylly in an herbaceous tropical vine, Syngonium (Araceae). Thesis. Harvard Univ., Cambridge. SIMMONDS, N. W. 1950. Notes on the biology of Ara- ceae of Trinidad. J. Ecol. 38: 277-291. DIMORPHIC INCOMPATIBILITY IN TURNERA HERMANNIOIDES CAMB. (TURNERACEAE)! SPENCER C. H. BARRETT AND JOEL S. SHORE? ABSTRACT Turnera udi d (Turneraceae) is a me diploid, perennial restricted. to 0 open, May sites in northeastern Brazil. Four p flon and short-styled plants. The style and stamen elpa in floral — is associated with differ- ences in pollen size and pollen production. In contrast, flower size, ovule number, and seed fertility of the morphs are not significantly different. A controlled pollination Mis ome demonstrated leri individuals of T. hermannioides possess a self- and intra-morph incompatibility system. The pollinations resulting in significant seed production were between the floral morphs. Populations of T. hermannioides grow intermingled with 7. ulmifolia, a wid was that resemble T. hermannioides. A crossing program between T. hermannioides and diploid and tetraploid varieties of T. — revealed barriers to dw akore in crosses at the diploid level. It is concluded that isolated from T. ulmifolia. Distyly is reliably reported from 23 flowering plant families (Ganders, 1979). In taxa on which experimental work has been undertaken, it has usually been shown that the stamen-style di- morphism is closely associated with a diallelic self-incompatibility system as well as several morphological polymorphisms involving pollen size, pollen production, and stigmatic papillae number and length. Although there is general agreement that distyly is an outbreeding mech- anism (Darwin, 1877; Crowe, 1964; Vuilleu- mier, 1967), the functions of the various floral polymorphisms that accompany self-incompat- ibility are still unclear (Yeo, 1975; Ganders, 1979). The large neotropical genus Turnera (Turner- aceae) contains some 54 species of which 37 are reported to be heterostylous (Urban, 1883). Species within the genus are poorly defined, and there is need for a modern taxonomic revision of the group. With the exception of studies of distyly in the widespread polymorphic weed complex T. ulmifolia L. (including the synonyms T. subulata Smith and T. trioniflora Sims) by Lock (1904), Barrett (1978), and Bentley (1979), there are no published reports of experimental work on the reproductive biology of species of Turnera. To broaden understanding of the na- ture of distyly in the genus, we initiated a study of the breeding system of T. hermannioides, a herbaceous to woody perennial native to north- eastern Brazil. In addition to documenting gen- eral features of distyly, incompatibility relation- ships, and population structure in the species, we also investigated its crossing relationships with T. ulmifolia, with which it is superficially similar in habit (Urban, 1883) and frequently sympatric. MATERIALS AND METHODS All populations of Turnera hermannioides studied occur in the state of Sergipé in north- eastern Brazil. Detailed localities are presented in Table 1. At each population, the number of long- and short-styled plants was recorded. In populations 2, 3, and 4, the entire flowering pop- ulation was scored whereas in population 1, a complete sample of the accessible part of the population (ca. one-third of the total size) was made (Table 5). Deviations from the expected 1:1 morph frequency were analyzed using the G-statistic which was calculated for each popu- lation. In addition, pooled and heterogeneity G-statistics were calculated (Sokal & Rohlf, 1981) to examine whether population structures were similar. ! We thank the staff of the herbarium at the Instituto Pesquisas Agronomicas, Recife, Pernambuco E assis- Ca orthea de bo tance, ER nada for funding field studies in n securing seed collections of Turnera ulmifolia Dárdano of us to Turn a Mercedes Ar aid in late Professor stern "den and Mari ted to the memory of the de — Lima, authority on the flora and vegetation of abbr Brazil, who introduced o iy cease of Botany, University of Toronto, Toronto, Ontario MSS 1A1, Canada. ANN. Missouni Bor. GARD. 72: 259-263. 1985. 260 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 E]. Localities of Turnera hermannioides and Turnera ulmifolia populations used in this study. All populations of T. hermannioides were collected along roadsides on open, waste groun A. Turnera hermannioides Varieties of T. ulmifolia Population Locality Present at Site ls Hwy. 235, ca. 30 km W of Itabaiana, Sergipé elegans 2 Hwy. 235, ca. 15 km W of Itabaiana, Sergipé elegans 3 Hwy. 235, ca. 1 km W of Itabaiana, Sergipé elegans 4° Hwy. 235, 21 km S of BR101, near Aracaju, Sergipé elegans and surina- mensis B. Turnera ulmifolia Chromosome Number Variety Locality and Ploidy Levels surinamensis Santarém, Pará, Brazil 2n 2x = 10 grandiflora Corrientes, Argentina 2n = 2x = 10 intermedia Barreirinhas, Maranhá l 2n = 2x = 10 intermedia St. Cristobal, Dominican K oubli 2n = 4x = 20 elegans Crato, Ceara, Brazil 2n = 4x = 20 * Vouchers for chromosome counts, Shore and Barrett 1 TRT. * Vouchers for chromosome counts, Shore and Barrett 2 TRT * Vouchers for chromosome counts, Shore and Barrett (unpubl. data). At population 1, ten flowers of each style form and corolla diameter were measure pers to the nearest 0.1 mm (Table 2). Style and stamen lengths were measured from the base of the ovary to the top of the male or female re- productive organ. Flower buds, 2-4 mm in length, were collected and fixed in 3:1 ethanol:acetic acid for chromosome number determination. Later, anther squashes were prepared in 296 ace- to-carmine, and meiosis was observed in pollen mother cells. Eighteen larger buds were collected separately from 18 individuals of each style morph to determine pollen production. Pollen counts were made using the hemacytometer method of Lloyd (1965). Anthers from three buds per morph were pooled for each value obtained. Six replicate samples were then determined for each morph. Data were analyzed by nested AN- OVA. A bulk seed sample was collected at population T of 3 parts sterilized potting compost (1 peat:1 clay-loam: 1 sand): 1 sand: 1 peat, in 7.5 cm plas- tic pots. Seeds were covered with about 1 cm soil and allowed to imbibe water for one week. As the soil was allowed to dry, germination pro- ceeded. Nine short-styled and 13 long-styled plants were obtained. When they had reached the 5-leaved stage, these were then transplanted into 12 cm clay pots and grown under uniform glasshouse conditions in air temperatures rang- ing from 25°-35°C. The plants were the material used for experimental work described below. The polar and equatorial axes of five pollen grains were measured for each of seven individ- uals per morph. Pollen grains were measured to the nearest 1.5 um using a calibrated ocular mi- crometer on a compound microscope. Nested ANOVA was used to examine variation among individuals and between style morphs. A controlled crossing program was initiated in May 1983, to determine the compatibility rela- tionships of the floral morphs (Table 4). Each individual was self-pollinated at least twice, crossed to at least two different individuals of the same floral morph, and to a minimum of two individuals of the opposite Toa morph. Polli- on llinator-free glass- l Pollen was transferred from the male to the female parent by removing all stamens with a pair of clean, fine forceps and rubbing the de- hiscent anther sacs on recipient stigmas. The number of seeds set per pollination was recorded. For inter-morph crosses, capsules were harvest- ed prior to dehiscence, which normally occurred approximately 24 days after pollination, and dis- at ina 1985] BARRETT & SHORE— TURNERA HERMANNIOIDES 261 TaBLE 2. Floral characters in the long- and short-styled morphs of Turnera hermannioides. Values are the mean and standard m Measurements of floral organs and pollen production were made from field gan plants i in population g p g in population 4. = = = not significant. Floral Morph Character Long-styled Short-styled F P Flower diameter (mm) 33.9 + 2.8 34+ 5.1 0.007 NS Style length (mm) 12+ 1.1 7.3 + 0.9 114.0 «0.001 Stamen length (mm) 7.3 + 0.6 12 + 0.8 221.0 «0.001 Pollen size (um) (polar axis) 63.6 + 2 72.9 + 2.6 61.7 «0.001 Pollen size (um) (equatorial axis) 3+0. 39.6 + 1. 117.0 <0.001 Pollen d 15,200 + 2,700 11,800 + 2,200 6.14 «0.05 Ovule number -8ower 32.3 + 9.9 39 + 8.8 2.4 NS quis woa 20 + 10.1 18.9 + 10.1 0.14 NS a Following inter-morph pollination. sected open. This procedure enables determi- nation of the number of seeds set and the total number of ovules produced in a flower. Data on seed set and ovule number were analyzed by nested ANOVA using a sample of nine individ- uals of each morph and two replicate measures for each individual. A second crossing program was initiated in June 1983, to determine the crossing relation- ships of T. hermannioides with taxa in the related T. ulmifolia complex (Table 6). Five population samples of four varieties were used. Five indi- viduals from each beue regis T. her- mannioides were used. The varietal status, lo- cality, and ploidal level of Suse of the T. ulmifolia complex are provided in Table 1. Fur- ther details of the reproductive biology of pop- ulations are presented in Barrett (1978). Polli- nations were performed as described above. Capsules were harvested after dehiscence. Seeds were retained in the capsules by placing parafilm over the swollen ovary about ten days after pol- lination. For each cross the number of brown plump seeds produced per pollination was re- orded RESULTS Floral morphology. Flowers of Turnera her- mannioides exhibit a precise reciprocality in the lengths of stamens and styles between the long- and short-styled morphs (Table 2). There was no significant difference between the floral morphs in flower diameter, number of ovules per flower, and seed set following legitimate (intermorph) pollination. In common with many distylous plants, 7. hermannioides exhibited dimorphism in the size of pollen grains. Both axes of pollen grains are longer in the short-styled morph. Pol- len production per flower in the long-styled morph was significantly higher than in the short-styled morph (Table 2). Microscopic observation of stigmas ofthe floral morphs revealed the absence of stigmatic papillae, a feature shared with T. ulmifolia, and in contrast to most heterostylous species. Where appropriate data were available, nested ANOVAs were undertaken on reproductive traits in e to panum the observed variation into thet ng individuals and error). In our samples, pesos cant variation occurred in ovule number per flower among individuals, but not between the floral morphs (Table 3). Large samples clearly would be required to determine if morph-specific differences in ovule number occur. The data for ovule number and pollen production suggest that TABLE 3. Variance components of reproductive characters in d eiii ndn qi Values are the percentage of the variation which is attrib- utable to debeas m: floral morph, and error following nested analysis of variance. Individ- Character ual Morph Residual Ovule number 70* 13 17 Pollen production 40> 38 22 Pollen size (polar axis) 9v 85^ 6 a P < 0.05. * P « 0.001. 262 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 TABLE 4. Fruit and seed set following controlled self, intra-morph and inter-morph cross-pollination of the long- and short-styled morphs of Turnera hermannioides. Plants were grown from seed collected in population 4; all pollinations were performed under glasshouse conditions. Pollination No. of 9 Flowers 96 Fruit No. of Seeds Mean Seed Set per Treatment Plants Pollinated Set Produced Capsule + s.d. L self 13 87 10 13 1.4 + 0.7 S self 9 62 13 10 1.3 + 0.7 LxL 13 28 25 7 2.3 + 0.3 SxS 9 18 6 l 1 LxS 13 27 100 540 20 + 10.1 S x L 9 18 100 341 18.9 + 10.1 considerable individual variation in fertility may exist within populations of 7. hermannioides. The controlled pollination program nstrat of a physiological self-incompatibility system in T. hermannioides. Self- and intra-morph polli- nations resulted in relatively small amounts of seed being produced. In contrast, legitimate pol- linations were highly fertile and all pollinations resulted in capsules and seed (Table 4). Population structure. Morph frequencies within populations of 7. hermannioides are iso- plethic (equal proportions) and the pooled and heterogeneity G-statistics indicate the absence of heterogeneity among populations (Table 5). Pre- sumably, this equilibrium is maintained by dis- assortative mating Deren the floral morphs as aresult of th y SyS- tem. Crossability with Turnera ulmifolia. Ob- servations of meiosis in plants of T. herman- nioides from populations 1 and 4 indicated that TABLE 5. Representation of floral morphs in four populations of Turnera hermannioides in Sergipé, northeastern Brazil. NS — not significant No. of No. of Long- Short- Popula- styled styled tion Plants Plants G3 df P l 60 57 0.08 l NS 2 54 56 0.04 l NS 3 60 53 0.43 l NS 4 47 61 1.82 l NS Total 221 227 2.37 4 NS * G (pooled) 0.08, df 1, NS; G (heterogeneity) 2.29, df 3, they were both diploid (25 = 2x = 10) with reg- ular bivalent formation. Results of the crossing bers of the 7. ulmifolia complex are presented in Table 6. Crosses between the two taxa usually resulted in the initiation of capsules, however, fruits and developing seed commonly aborted after initial swelling. Barriers to c bilit particularly evident between diploid aret es of T. ulmifolia and T. hermannioides. Althou proximately one-third of the pollinations that were conducted resulted in fruit, little seed was formed. Interestingly, significant numbers of seed were produced when 7. hermannioides was used as the male parent in crosses with the two tet- BLE 6. Seed fertility in reciprocal crosses be- tween Turnera hermannioides (population 4) and va- rieties of Turnera ulmifolia. All pollinations were per- formed under glasshouse conditions. No. Pol- 9 Mean Seed Set Varieties and lina- Fruit per Pollination Ploidy Level tions Set + s.d. A. T. hermannioides as 9 parent surinamensis (2x) 9 56 1.8 + 2.6 grandiflora (2x) 12 25 0.4 + 0.8 intermedia (2x) 8 25 0.8 + 1.8 intermedia (4x) 6 83 4.3 + 2.7 elegans (4x) 6 33 13 +1 B. T. hermannioides as à parent surinamensis (2x) 10 30 1 + 1.7 grandiflora (2x) 11 36 1.3 + 2.1 intermedia (2x) 9 22 0.3 + 0.7 intermedia (4x) 7 100 8.3 + 3.6 elegans (4x) 71 13 + 10.1 1985] raploid varieties of T. ulmifolia. The reciprocal crosses were less productive. The crossing pro- gram demonstrates that 7. hermannioides is, in large part, reproductively isolated from T. ul- mifolia at the diploid level. Although F, plants may be produced from crosses with tetraploid varieties, they would be triploid and probably sterile. DISCUSSION Apart from the absence of stigmatic polymor- phisms, the distylous syndrome of T. herman- nioides resembles those described in several oth- er heterostylous families (reviewed in Ganders, 1979). The corolla morphology of distylous Tur- nera species is unusual for heteromorphic taxa because of the absence of a well-developed co- rolla tube. Flowers of 7. hermannioides and T. ulmifolia are bowl-shaped and relatively unspe- cialized. Observations of visitors to flowers of both species indicated that they were visited by a wide range of insects, including Apis mellifera, Xylocopa sp., and various groups of solitary bees (see Barrett, 1978). It seems likely that with rel- atively generalized insect visitors and unspe- cialized flowers, considerable illegitimate polli- nation would occur, particularly in plants with large floral displays. The strongly developed self- incompatibility system in 7. hermannioides and T. ulmifolia (Barrett, 1978) may thus be main- tained to prevent self-fertilization. There is little information on the geographic distribution of 7. hermannioides. Based on our own field observations, as well as surveys of her- barium collections from Brazil, it would appear that the species occurs in scattered localities in the states of Sergipé, Bahia, and Minas Gerais. Diploid varieties of 7. u/mifolia in Brazil also exhibit restricted and widely scattered distribu- tions, frequently in association with local soil conditions. In striking contrast, polyploid vari- BARRETT & SHORE— TURNERA HERMANNIOIDES 263 eties of the 7. ulmifolia complex are ruderal weeds with broad ecological tolerance and widespread distributions (Barrett, 1978; Bentley, 1979). Throughout northeastern Brazil, the tetraploid T. ulmifolia var. elegans is conspicuous as a wee of waste ground and roadsides, including those in which 7. hermannioides occurs. Although T. ulmifolia var. elegans occurred intermingled with T. hermannioides in the four populations sam- pled, no putative hybrids were observed. While small numbers of seed may be produced from crosses between the two taxa all attempts by us to obtain viable seedlings have failed. The taxa are reproductively isolated from one another. LITERATURE CITED BARRETT, S. C. H. 1978. Heterostyly in a tropical weed: the reproductive biology of the Turnera ul- mifolia complex. Canad. J. Bot. 56: 1713-1725. BENTLEY, B. L. 1979. Heterostyly in Turnera trioni- flora, a m weed of the Amazon Basin. Bio- tropica 11: Crowe, L. K. Da The evolution of outbreeding in on Plants of the Same Species. John Murray, Lon- don. GANDERS, F. R. 1979. The biology of heterostyly. New Zealand J. Bot. 17: 607—635. LrLovp, D. 1965. Evolution of een prd and racial differentiation in Leavenworthia (Crucifer ae). Contr. Gray Herb. 195: 3-134 Lock, R. H. 1904. Ecological notes on Turnera ul- mifolia L. var. elegans Urban. Ann. Roy. Bot. Gard. (Peradeniya) 2: 107-119. SoKAL, R. R. & F. J. RoHLF. 1981. edition. wr San Francisco. STEBBINS, G. . Chromosomal Evolution in Higher nd Addison-Wesley, Reading, Mas- Biometry. 2nd 83. Monographie der gres der Tur- eraceen. Gerbruder Borntraeger, LIE B.S. 1967. The origin Bx evolution- a evelopment of heterostyly in the angio- sperms. Evolution 21: 210-226. Yeo, P. F. 75. Some aspects of heterostyly. New Phytol. 75: 147-153. E THE PLANTS OF 'OCOQUILT ISLAND, SAN BLAS COAST, PANAMA! W. G. D'ARCY AND BARRY HAMMEL? ABSTRACT Study of one of the few San Blas Islands (Panama) with intact vegetation indicates that the flora has a greater affinity with distant circum-Caribbean islands than with the nearby mainland. The San Blas coast of northeastern Panama is fringed by a series of small flat islands, few of them as long as 1 km, and few more than 1 km offshore. Nothing is known about the vegetation of the islands before they were settled by the Kuna Indians in the middle of the last century (Jaén, 1978). Today most of the islands are densely populated without a single trace of orig- inal plant life remaining. On 9 October 1978, we hired an outboard motor boat and visited a small island in the group that was uninhabited and appeared to have a large measure of its natural vegetation intact. The 34 species noted during this morning included two new to Panama and an assemblage of species more likely to be found on a flat limestone island in the northern Carib- bean hundreds of miles away than on the main- land of Panama less than 1 km away. The island’s location, 9?14'N, 48?01'45"W, is midway between Isla Nustupo and Ailigandí, and it is approximately halfway along the San Blas coast between Porvenir, the district capital, and Puerto Obaldía at the Colombian frontier. It is only a few hundred meters from the mainland. This island is not marked on any maps we con- sulted, and we refer to it by the name given by our Indian guides, ‘Ocoquili,’ which refers to the coconut trees on it. Upon leaving the area later in the day, we persuaded our pilot to circle ‘Oco- quili' and the photo shown here (Fig. 1) was tak- en. Ocoquili Island is about 1 km long and 300- 400 m wide, and it is formed of coquina-like limestone. Coconuts grow around most of the edges, and on the western end there is an accu- mulation of sand. Much of the interior is low and flooded, covered by mangroves. Across the island toward the eastern end is a shallow chan- nel and holes in the partly emergent ida at the edges of this, harbored small plan Although no one lived there at the Ee our guides reported that boatmen sometimes visited the island for coconuts, fishing, parties, and other reasons but did not stay long at a time. We found no signs of construction and no sign of feral an- imals. A total of 37 collections were made, some of them sterile. Determinations of Gramineae and by the senior author. Thirty-three species were represented, and the sight record of Cocos nu- cifera yielded a total of 34 species (Table 1). Al- though more species may be present, none were apparent at the time of this visit even in sterile condition. Our coverage was nearly complete. All collections are deposited with the Missouri Botanical Garden (MO). Of the trees, three species are mangroves. Red, white, and black mangroves were all present, but we found only one species of red mangrove, Rhi- zophora mangle. Of the other tree species, Hip- pomane mancinella is neotropical, Cocos nuci- fera is introduced, and Cordia sebestena, commonest in the northern Caribbean, has been collected few times in Panama, and then only along this same San Blas coast. All tree species are e mainly known hom neotropical seacoasts, y forests, and all are to be found i in proximity in many Carib- bean sites (Table 2 The shrubs have pan-Caribbean distribution, r elsewhere. Conocarpus erecta and piane 5 hile are found near many Caribbean coastlines, and they also occur in West Africa. Suriana maritima, which nae some 1 Supported by National Science Foundation Grant DEB 79-22192. ? Missouri Botanical Garden, P.O. Box ANN. MISSOURI Bor. GARD. 72: 264—267. 1985. 299, St. Louis, Missouri 63166. TABLE 1. List of the 34 species found on ‘Ocoquili’ Island, San Blas Coast, Panama. Sources: Rubiaceae from Dwyer (1980), Boraginaceae from Miller (in prep.), Amaranthaceae from Mears (1982), and other distri- butions from Adams (1972). Species Family Habit Range Habitat-Ecology Andropogon bicornis L. Gramineae herb neotropical weed species Avicennia germinans (L.) L. Verbenaceae tree neotropical, limestone coasts W Africa Blutaparon vermiculare (L.) Mears Amaranthaceae herb neotropical coastlines Canavalia maritima (Aubl.) Thou. Leguminosae herb pantropical coastal strands Cassine xylocarpa Vent.* Celastraceae shrub N Caribbean dry woodlands Chamaesyce mesembrianthemifolia Euphorbiaceae herb neotropical calcareous m coasts Chiococca alba (L.) Hitchc. Rubiaceae vine Texas to Ar- limestone gentina Cissus sicyoides L. Vitaceae vine neotropical weed species Coccoloba uvifera L. Polygonaceae tree Caribbean seacoasts Cocos nucifera L. Palmae tree introduced seacoasts Conocarpus erecta L. Combretaceae shrub neotropical, coastal W Africa Cordia sebestena L." Boraginaceae tree N a W Carib- seacoasts Crinum erubescens Ait.° Amaryllidaceae herb peer shorelines Cuervea kappleriana (Miq.) A.C. Sm. Hippocrateaceae vine neotropical coastal wood- ands Dalbergia brownei (Jacq.) Urb. Leguminosae shrub Caribbean to coastal wood- Brazil lands Dalbergia ecastophyllum (L.) Taub. Leguminosae shrub neotropical, seacoast thickets Eustachys petrea (Sw.) Desv.” Gramineae herb neotropical limestone & beaches Fimbristylis cymosa R. B Cyperaceae herb pantropical coastal areas Hippomane mancinella "à Euphorbiaceae tree neotropical coastlines Laguncularia racemosa (L.) Gaertn. f. Combretaceae tree neotropical, mangrove rica Lantana involucrata L.^ Verbenaceae shrub Caribbean limestone & coasts Melanthera aspera (Jacq.) Benth. Compositae herb Caribbean, seacoasts Ecuador Pavonia rhizophorae L.** Malvaceae herb Panama to seacoasts Colombia Pilea microphylla (L.) Liebm. Urticaceae herb pantropical weed species Randia aculeata L. Rubiaceae shrub exico to dry limestone Venezuela Rhizophora mangle L. Rhizophoraceae tree neotropical, mangrove Sesuvium portulacastrum L. Aizoaceae herb pantropical seashores Sophora tomentosa L.” Leguminosae shrub pantropical coastal weed Spartina spartinae (Tru.) Merr.* Gramineae herb exas to Ar- coasts & marsh- gentina Sporobolus virginicus L Gramineae herb pantropical shores Stenotaphrum psoas: spp (Walt.) Gramineae herb pantropical limestone coasts Gaertn. f.> Suriana maritima L.* Simaroubaceae shrub Caribbean, In- seacoasts i ans Vigna luteola (Jacq.) Benth. idee dn herb pantropical limestone coasts Wedelia trilobata (L.) Hitchc. Compos herb neotropical, weed species W Africa a New report for Panam: ° [n Panama, found only nearby on this coast. * Not on Jamaica. 266 Aerial view of ‘Ocoquili’ Island. FIGURE 1. is a new record for Panama, is found on seacoasts in the area and also in the Indian Ocean and South Pacific. Dalbergia brownei ranges around it also occurs inland in some places. Randia acu- leata is used as a Christmas tree in some Carib- bean islands: it occurs inland in Panama as well as along the coasts. There are only three species of climbing vines; two of them, Chiococca alba and Cissus si- cyoides, are pantropical and also plentiful in the Antilles, and Cuerva kappleriana is restricted to TABLE 2. Growth forms on ‘Ocoquili’ Island. Trees 7 Shrubs 8 Herbs 16 Climbing vines 3 Total species 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 the Caribbean area. Several of the herb or shrub species might also be called vines: Dalbergia brownei, which has tendrils, Vigna luteola, and Canavalia maritima, but these do not usually new reports for Panama, and eight species are known in Panama only from sites along this coast. All but three species, Crinum erubescens, Pa- vonia rhizophorae, and Spartina spartinae, are known from Jamaica, more than 900 km to the north, and the last of these is widespread on cal- careous coasts in the Caribbean area and on al- kaline flats in inland continental areas. While the florula of the tiny Ocoquili Island is but a token representation ofthe flora that once covered this string of numerous islands, it is suf- an assemblage of species distinct from the as- sociations now known on the mainland. D'ARCY & HAMMEL—PLANTS OF 'OCOQUILT ISLAND 267 1985] LITERATURE CITED del siglo XVI al siglo XX. Omar Jaén Suarez, Pan- ADAMS, C. D. . Flowering Plants of Jamaica. Univ. of the West Indies, Mona, Jamaica. kas J.A : se patel aspis La Dwyer, J. D. Rubiaceae. /n R. E. Woodson low siti tege 8 aep A et laccack 2 I et al., Flora of Panama. Ann. Missouri Bot. Gard. 11 vä 117 7: 1-256. Jaén S., O. 1978. La población del Istmo de Panamá ANTIRHEA AROMATICA (RUBIACEAE, GUETTARDEAE), A NEW SPECIES FROM VERACRUZ, MEXICO! GONZALO CASTILLO-CAMPOS? AND DAVID H. LoRENCE? ABSTRACT new species of Rubiaceae, Antirhea aromatica Castillo-Campos & Lorence, is described from central Veracruz, Mexico. A tree of the tropical semideciduous forest, it appears to be very localized in distribution and is the first record of the genus for Veracruz. Its distribution, habitat, and affinities are discussed, and specimen citations are given RESUMEN ribe una nueva especie de Rubiaceae, Antirhea Maie See Campos & Lorence, del Se árbo a media Ns desc entro de Veracruz, México. des 2 un á área de distribución muy restringida y Se ] que crece en la más se registra e "aen jii primera vez para V dos ana quide elect con discute su distribución, hábitat y afinidades, y se citan los ejemplares examina During the course of studies on Miei Ru- biaceae and on Jal in central siad several collections were en- countered which represent an undescribed species of Antirhea of the tribe Guettardeae. Members of the Guettardeae are characterized by having a two- to many-locular ovary in which each loc- ule has a single anatropous ovule, a corolla with the stamens inserted in the throat, and a usually cymose inflorescence. Antirhea Comm. ex A. L. Juss. is distinguishable from the other neotrop- ical genera in the tribe by the following combi- nation of characters: an axillary, usually dichot- omous, cymose inflorescence, a corolla with strongly imbricate lobes, one or two of them being exterior, i with two to nine cells and a deciduous calyx (Standley, 1934). comulco Ou dated into both the Guettardeae and por itself. Its Il duced to a single flower. Such biflorous cymes, as well as those reduced to solitary flowers, also occur in a number of Antillean Antirhea species (Standley, 1934). Antirhea is a genus of about 50 species dis- tributed in the Paleotropics of Madagascar, the SW Indian Ocean islands (Comores, Mascar- enes), W India, Asia, Indonesia and Australia, and also in the American tropics where it is best represented i in the Antilles. Standley (1934) rec- ognized 31 American species, most of which oc- cur in the larger islands of Cuba, Hispaniola, Jamaica, and Puerto Rico. One Antillean species, A. lucida (Sw.) Benth. & Hook., reaches Me- soamerica where it has been reported from Belize (Standley & Williams, 1975), Guatemala, and more recently from Quintana Roo in Mexico (Téllez V. & Sousa S., 1982). A second species, represents the first report of the genus from Ve- racruz. The majority of Antirhea species appear to have extremely restricted distribution patterns, i.e., most are endemic to a single island or are known only from the type locality. As Veracruz and the adjacent states have been relatively well collect- ed, the new species may be endemic to the state. Antirhea aromatica Paca Campos & Lorence, sp. no . Veracruz: Munici- pality of iowa: ca. 10 km S of Apa- zapan, Barranca de Monterrey, S of Cuetza- lan, between Cuetzalan and Apazapan; ! We are grateful to M. C. Johnston for assistance with the Latin diagnosis, to E. Saavedra for preparing the illustration, to L. Robles for assistance in the field, and to E. J. Lott and F. Chiang for comments on the scri 2 Instituto Nacional de Investigaciones sobre Recursos Bióticos, Apdo. Postal 63 (INIREB), 91000 Xalapa, éxi Veracruz, ? Instituto de "Biología, ooo Nacional Autónoma de México, Apdo. 70-233, Deleg. Coyoacán, C.U., 04510 México, D.F., Mex ANN. Missouni Bor. GARD. 72: 268-271. 1985. 1985] semideciduous forest, 350 m, 17 Sept. 1983 (fl, fr), Castillo-Campos 2957 (holotype, XAL; isotypes, MEXU, XAL). Figure 1. r 6-15 metralis lenticellata. Folia petiolata; lamina elliptica vel ovato-ellipti ta 9 mm longa 40—90 mm lata. Inflorescentiae axillares biflores; pedunculus 20-45 mm longus. Flor ubes centes; cupula calycis 5-6 mm 5 triangularibus subulatis 4-8 mm longis; corolla alba PA gui tubo 90-117 mm longo, lobis 4-5, 10-16 m longis; antherae 4-5 ca. 8 m m lon ngae. Fructus -37 11 longus mm longus 10-17 mm latus, costulis longitudinalibus 6—8 prominentibus; semina 7-10 cylindrica 25-3 longa. Trees 6-15 m tall, 10-30 cm dbh, the bark whitish, flaking near the base, the blaze pinkish h white rays, the wood yellow. Stems brown- ish, lenticellate, 3-4 mm diam., the internodes crowded apically, the new growth hirtellous, res- inous; stipules caducous, brown, resinous, ovate- deltoid, naviculate, 11-14 mm by 5-7 mm, the apex acute to acuminate, the broad margins scar- ious, externally hirtellous, internally puce m sally, lined with a d each 0.5—0.6 mm long. Leaves i in subequal | pairs, petiolate; petioles 15-60 mm by 0.8-1 mm, hir- tellous, slightly canaliculate adaxially; lamina el- liptic to ovate-elliptic, 90-200 mm by 35-90 mm, the apex shortly acuminate or rarely acute, the acumen 5-20 mm long, usually falcate, the base narrowly cuneate to cuneate, often more or less attenuate, the sides subequal, the secondary veins 6-8 pairs, festooned brochidodromous, the ul- timate venation prominently reticulate, visible iscolo- , drying brown apbressed hirtellous-hirsutulous expecially when young, glabrescent, the costa and veins hirsutu- lous, the secondary vein axils barbate abaxially, the margin ciliolate, callose. Inflorescence axil- lary, several generally produced near the branch apex, each with 2 flowers; peduncle 20-45 mm by 0.8-1.2 mm, appressed hirtellous. Flowers sessile, each subtended by 1(—2) persistent often bilobed ovate-naviculate involucral bracts 5-8 mm by 4-6 mm, acute, carinate, the margins scarious, each lined with a deltoid patch of dig- itate glands; hypanthium obconical, 2-2.5 mm long, tomentose; calyx cup 5-6 mm long, tubular, externally spersely hirsutulous, internally with š deltoid p h the lobes; calyx lobes (4-)5, deltoid- saaal: 4- 8 mm by 1.5-2.5 mm basally, externally sparsely CASTILLO-CAMPOS & LORENCE—ANTIRHEA 269 hirsutulous. Corolla white, aromatic when fresh, salverform, the tube cylindrical, 90-117 mm long y 4-5 mm wide medially, externally hirsutu- lous, more or less resinous basally, the lobes (495, imbricate in bud, ovate-elliptic, 10-16 mm by 5-7 mm, obtuse, externally hirsutulous, in- ternally glabrous, recurved at anthesis, the mar- gin undulate; stamens (4-)5, interioblilar: sessile, affixed ca. 5 mm below the faux; anthers linear- subulate, 8-9 mm by 1 mm, the base cordate, the apex acute, exserted for 2-3 mm; style 40- 70 mm long, sparsely spreading-hirsutulous; ovarian disc 1.5 mm long, doughnut shaped; ovary 7-10-locular, the ovules pendulous. Fruit b eid obovoid-ellipsoid, 25-37 mm by 10- mm, 6-8-costate, sparsely hirtellous, resi- nous, the pericarp thin, fleshy, yellowish w ripe, the calyx ultimately deciduous; seeds 7-1 e tightly adherent, narrowly ellipsoid, 25-30 mm by 3-5 mm medially, more or less curved, the testa tough, spongy; embryo cylindrical. Distribution. The species is known only from the type locality in central Veracruz, near the towns of Jalcomulco and Apazapan in the mu- nicipality of Jalcomulco at altitudes of 350- imens examined. MEXICO. VERACRUZ: Munic- ipality of Jalcomulco, ca. 10 km S of Apazapan, Bar- ranca a Monterrey, S of Cuetzalan, between Cuetza- azapan, 350 m, Castillo C. 2987 (XAL), Robles 248 (XAL); vicinity of Jalcomulco, ca. 500 m, 25 July 1973, Gándara & Dorantes 90 (F, MEXU, MO). In addition, ca. 50 more isotypes will be dis- tributed shortly to a wide range of herbaria. Phenology. Flowering occurs during July and August; the fruits are produced concurrently and ripen in October. The somewhat fleshy, yellow pericarp suggests bird dispersal, although no re- cent regeneration was observed locally. cific epithet refers to the strong, gar- denia-like fragrance produced by the flowers in the evening. This, together with the long, tubular white corollas, is characteristic of plants with a hawkmoth pollination syndrome. The patches of digitate glands within the stipules, floral bracts, and calyx tube apparently produce the brown resin that bathes the young developing organs and is presumably protective in nature. The resin is highly aromatic when dissolved in alcohol, to which fact the epithet also alludes. This resin also occurs in a number of Antillean species. 270 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 1985] CASTILLO-CAMPOS & LORENCE— ANTIRHEA 271 TABLE l. Some diagnostic morphological characters of the two species of Antirhea known from Mexico. Character A. aromatica A. lucida Vegetative pubescence hirtellous-hirsutulous glabrous Resin present absent Stipule size (mm) 11-14 by 5-7 4—7 by 1.5-2 Petiole length (mm) Lamina size (mm) 15-60 90-200 by 35-90 6-8 3-10 45-65 by 20-35 Number of secondary vein pairs 10-11 erus ind e per 2 22-30 inflor Calyx nla p (mm) 10-14 2-3 Corolla tube length (mm) 90-117 4-5 Fruit size (nmm 25-37 by 10-16 5-7 by 3-4 Common name. “Chicahuastle. Uses. The wood is said to be used locally for construction of houses. Habitat. The area around Jalcomulco and Apazapan is characterized by series of limestone hills and cany ith altitudes ranging from 350 to 1,000 m dissected by a river system. The lime- stope outcrops are o vered with a thin, black clay The area was originally covered by low, seasonally decid- uous tropical forest intergrading with areas of taller semideciduous tropical forest in canyons and on slopes. Much of this forest has been con- verted for sugar cane and mango plantations. Characteristic arboreal components of the low deciduous forest are: Bauhinia divaricata L., Bursera simaruba (L.) Sarg., Calliandra spp., Comocladia engleriana Loes., Diospyros verae- crucis Standley, Diph d robinioides Benth., Pis- tacia mexicana ., Plumeria rubra L., and Spondias sp., s spp., and columnar a as Lu as Hechtia spp., abound in the lower stra- tum and understory. Dominant tree species of the taller semideciduous forest include: Brosi- mum 'H <ç R. 133a annm i) hin (T ) Sarg Hyperbaena mexicana Miers, Manilkara zapota (L.) Van Royen, and Protium copal (Schldl. & Cham.) Engl Diversity of woody species is high in the area, which may be partly due to the series of ecotones between the two forest types. Antirhea aromatica occurs only in the most humid canyons where it is a fairly common mid-stratum component of the tropical semideciduous forest. Antirhea aromatica is easily distinguished from listed in Table 1. It differs from the other Antil- lean members of the genus by its larger, thinner leaves and much larger flowers and fruits. Its closest relative appears to be A. involucrata Ur- ban & Ekman from Hispaniola, which also grows on limestone, is resinous, and has a two-flowered involucrate inflorescence with a relatively large corolla and fruit. The latter species differs by its thicker glabrous leaves, a smaller subsessile in- florescence, and smaller flowers and fruits with flowered panicle and much smaller flowers and fruits LITERATURE CITED p J. D. 1980. Rubiaceae. In Flora of Panama. n. Missouri Bot. Gar FEN eun P. C. 1934. Rubiaceae. In N. Amer. Fl. 32: 262-275. . O. WILLIAMS. 1975. . In Flora of Guatemala. Fieldiana, Bot. 24(11 TELLEZ V., O. & M. So 1982. on de la Flora Quin ntanarooense. Centro de [OAM de Quintana Roo. Litoarte S. de R.L., Méxic — FIGURE 1. ruits.—b. De peduncle. —d. Stipule, internal s mature fruits. — g. ture fruits, tone longitudinally and transversally.—h. fruit. —i. Seeds. Illustration by E. Saavedra (INIREB), based on Castillo- Campos 2957. Habit and details of Antio ar — Castillo- Campos & Lorence. —a. Habit, with flowers and of opened flower. — Transverse section of mature THREE NEW SPECIES OF RHIPIDOCLADUM FROM MESOAMERICA! RICHARD W. POHL? ABSTRACT Rhipidocladum clarkiae Pohl and R. pacuarense Pohl are described from dade m flowering material and R. panamense Pohl is described from flowering specimens from Pan The genus Rhipidocladum was erected by McClure (1973) to include species of bamboos previously assigned to Arthrostylidium, Arundi- naria, and Ludolphia. The species of Rhipido- cladum are slender and graceful. At each mid- culm node, they bear an adnate triangular flat (apsidate) array. The genus is differentiated veg- etatively from Merostachys, which has a similar branchlet array, by the nature of the main culm leaves. In Rhipidocladum, the base of the erect culm blade is as wide as the apex of the sheath and adnate to it. In Merostachys, the reflexed pex. The species of Rhipidocladum are slender bamboos, ranging from 2 to 10 m or more in length and 1 to 5 cm diam. They inhabit mesic sites in moist forests and are found on canyon walls and river banks at low and middle altitudes. The genus ranges from southern Mexico to northern South America, with one species re- ported from Brazil. McClure indicated that his genus included 11 species and offered a key to distinguish some of these. One species, R. ver- ticillatum, has since been transferred to a new genus, Actinocladum, by Soderstrom (1981). With the addition of these three new species, the cur- rent number is 13. Mesoamerican species of Rhipidocladum are not well known because of their erratic blooming cycles. Recent gregarious flowering of two Costa Rican species and one from Panama has allowed differentiation of species previously known only in the vegetative condition. Two of these species, R. clarkiae and R. panamense, are distinct from the remaining species of the genus because o their extremely large number of filiform branch- es. The third, R. pacuarense, is most closely re- et lated to R. bartlettii of the Petén of Guatemala and Chiapas, and their differences are stated in Table 1. In the absence of adequate type material of many of the species, detailed comparisons with the remaining species cannot be made. The ke given by McClure (1973) offers the best separa- tions to date. Rhipidocladum clarkiae Pohl, sp. nov. TYPE: tion Booth, S of Bajo de Hondura, 1,350 m, 26 July 1982, Pohl & Clark 14103 (holo- type, ISC; isotypes, CR, K, MO, US). Figure la-e Gramen lignosum caespitosum; culmi 10-20, plus 130 in complemento, delicati, 45-65 cm longi. Inflo- rescentia racemosa biseriata delicata. Spiculi 7-8, 15— 30 mm longi; gluma primera acicularis, 0.5-2.5 mm, uninervia; gluma secunda ovata, 4—5.5 mm, arista ca. mm; lemma sterile 4-5 mm longum anguste-ovata, 5-nervata; spiculis supra lemma sterile secedens; lem- mata fertilia 2-4, 7-9 mm longa, scabrida, arista 4—6 mm; palea lemma aequans, apice scabrida; lemmata rudimentaria 2, convoluta; lodiculae 3, crystallis oxa- latis obtectis. Caespitose ligneous bamboo; culms in clumps of 10-20, more than 10 m long; internodes thin- walled, cylindrical, glabrous, 1-2 cm thick. Culm aths pering gradually pressed-hispid above near the base and margins. Flowering branches borne at many nodes, 75- 130/node, unbranched, extremely thin and del- ! Journal paper #J-11484 of the Iowa Agriculture and Home Economics Experiment Station, Ames, Project 1833 ? Department of Botany, Iowa State University, Ames, Iowa 50011. ANN. Missouni Bor. GARD. 72: 272-276. 1985. 1985] icate, 45-65 cm by 0.5 mm, including the inflo- rescences (plants not seen in vegetative phase). Branch leaves 2-3 per branch; sheaths glabrous or puberulent, the apex prolonged into an erect auricle bearing weak erect bristles to 6 mm; ligule membranous, lacerate, ciliolate; blades flat, dark green above, glaucescent beneath, 3-9 cm by 4- 9 mm, the midrib very inconspicuous. Inflores- cence a solitary slender unilateral raceme, ter- minal on the branch; peduncle scarcely exserted, puberulent; rachis 9-13 cm long, very slender, puberulent; pedicels less than 1 mm long, ap- pressed. Spikelets 7-8 per raceme, 10-14 mm apart, erect, appressed, very slender, scarcely compressed, 15-30 mm long, disarticulating above the sterile lemma and between the fertile florets; first glume acicular, 0.5-2.5 mm long, 1- nerved, clasping the rachilla, the awn ca. 2 mm long; second glume narrowly ovate, 4—4.5 mm long, 3-nerved, the awn ca. 3 mm long; sterile lemma 4-5 mm long, ia: ovate, 5-nerved; hs florets 2-4, with 2 rudimentary, short- wned, convolute dn shorter than the fertile "n and borne above, fertile lemmas 7-9 mm long, narrowly ovate, 7—9-nerved, scabrid, the awn 4—6 mm long, scabrous; paleas of fertile flo- rets shorter than or equalling the lemmas, 2- keeled, the keels and tips scabrous; rachilla seg- ments 4.5-6 mm long, glabrous, wrinkled, the apex coroniform; lodicules 3, thin, flat, ovate, the surface beset with numerous ruses; the apex with 2—4-celled microhairs, the vascular bundles 1-2, weak, short, unbranched; anthers 3, ca. 4.5 mm long; style 1; stigmas 2; ovary beset with druse crystals. The only known specimens ofthis species were collected from the same colony along the access road to the Parque Nacional Braulio Carrillo, in both sides of the road. All appeared to be in bloom in 1982 and early 1983. TopoTyPEs: Pohl 14152, 13 Sept. 1982 (CR, F, ISC, K, MEXU, MO, US); Davidse et al. 23228, 23 Jan. 1983 (BM, CR, ISC, MEXU, MO, US). This species is named for Lynn Clark, student of the genus Chusquea, who participated in the original collection of the species. No other col- lections from other colonies are known, but the species should be sought on the wet northern slopes of the Cordillera Central of Costa Rica. The plants have a delicate aspect and have or- namental possibilities. POHL— RHIPIDOCLADUM 273 TABLE 1. Differences between R. bartlettii and R. pacuarense. R. pacuarense bartlettii Branch leaf L/W 5.6-9: 1 15-17: 1 ratio Abaxial blade base, both base, one pubescence sides side Rachis internodes 2-7 ca. 10 mm Spikelets/branch 22-25 6-18 Sterile lemma 7-8 mm 5—6 mm ma awn length «4 mm 0-1 mm Anther length mm 5 mm Rachilla s NE glabrous ultimate segmen scabrous Rhipidocladum pacuarense Pohl, sp. nov. TYPE: Costa Rica. Prov. Cartago: 1.8 km E of Río Pacuare crossing of Hwy. 232, top of hill, N side of road, 750 m, 1 Oct. 1982, Pohl 14161 (holotype, ISC; isotypes, CR, F, K, MO, US). Figure 1f-i Gramen lignosum caespitosum rhizomis pachymor- phis; culmi plus quam 10 m x 2-3 cm, cylindrati, cavi. Ramuli 18—30 in complemento, 20—60 cm longi. Folia estere ses 7-12 cm x 10-22 mm, subter vil- ried pi i i biseriata, arcuata 9-17 c ma ae x 3 mm. ro. pasan 18-19 mm; glu- 2.5-3.5 m s; gluma secunda mm, 3-5 iid og dcos minus quam 4 mm; spiculae supra lemma sterile secedens; lemma sterile 7-8 mm longa; lemmata fertilia 3, 7-8 mm, glabra, 7- 9-nervata, arista 3-4 mm; palea lemma aequans vel longior, apice e re lodiculae 3, vasculatae; antherae 3, 2-3 mm; stigmata 2. A R. bartlettii differt inflores- centia densior, pepe longiores rachilla glabra, fo- liorum forma (vide Table Caespitose, ligneous bamboo; rhizomes pachymorphous; clumps small; culms more than 10 m by 2-3 cm, cylindrical, green, thin-walled, glabrous, arching and the upper portions trailing; internodes —12 cm long; ligule a stiff membrane, 0.5-1 mm long, minutely ciliolate; blades erect, as wide as the sheath apex, the body ca. 6 cm long, with a narrow caudate apex ca. 3 cm long; margins stiffly hispid-ciliate. Branches usually 18— 30 per node, 20-60 cm long, sometimes re- branched from a middle node. Branch leaves usually 3—6 per branch; sheaths glabrous, the apex truncate, bearing fl l mm long; pseudopetiole smooth, purple, ca. 3 br is tles to 1 0 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 274 —]. Leaf blade, abaxial —c. Inflorescence. — d. Spikelet. —e. Culm leaf (all based on Pohl & Clark 14103). f-i. Rhipido- cladum pacuarense. — f. Branch complement. —g. Branch leaf blade, abaxial. —h. Inflorescence. — i. Spikelet (all based on Pohl 14161). j-1. Rhipidocladum panamense. —j. Inflorescence. — k. Spikelet. Rhipidocladum species. a-e. Rhipidocladum clarkiae. —a. Branch complement. — b. Branch leaf (all based on Gentry & Dwyer 3418). Scale lines = 1 cm. FiGURE 1. blade, abaxial. 1985] mm long; ligule less than 0.5 mm long; blades 7-12 mm by 10-22 mm, the length: width ratio 5.6—9:1, flat, without a conspicuous midrib, sca- brid above, rounded to the asymmetric base, the abaxial surface glaucescent, glabrous except for patches of woolly pubescence on both sides near the base. Inflorescence a slender arcuate raceme, terminal on a primary branch or a short second- ary branch from a middle node, 9-17 cm long by 3 mm wide; rachis glabrous; pedicels erect, ca. | mm. Spikelets 22-25 per raceme, borne 2- 7 mm apart, 18-19 mm long, appressed, disar- ticulating above the sterile lemma and between the florets, first glume 2.5-3.5 mm long, awl- shaped, 1-nerved, tapering to an awn 3-5 mm long; second glume narrowly ovate, 4.5-6 mm long, 3-5-nerved, tapering to an awn less than 4 mm long; sterile lemma 7-8 mm long, 5-7- naved, i above, ies awn [4,933 mm ) long; above; UE lemmas 7-8 mm long, rounded on the back, 7-9-nerved, usually purplish, glabrous, stiff, awn 3-4 mm long; palea shorter or longer than the lemma, ciliate above; lodicules 3, flat, hispid at the tip; two lodicules equal, ovate, sev- eral-nerved; third lodicule shorter, lanceolate, | -nerved; anthers 3, 2-3 mm long; style 1; stig- mas 2 Hillsides and canyons; central and southern Costa Rica; 650-1,400 m . c flowering specimens. UA RICA. valley of Rio Tuis, along H 2 gw SE of Jicotea, 1 Oct. 1982, Pohl 14165 (C (CR, F, ISC, K S). Representative vegetative specimens. COSTA PROV. CARTAGO: Pohl & Pinette 13182 (topotype) asc, Pohl & pie 13149 cco Pavas de Turri Pohl A e 13207 (ISC, PROV. PUNTARENAS: Cañas Gor Pohl & cao 13274 (ISC, MO). PRov. SAN JOSE: E onejo, Pohl & Davidse 11056 (ISC, MO), Pohl & cu beni (ISC, MON Rosario, Pohl & Gabel 13579 (ISC). PRov. ALAJUELA: San Ramó » Pohl & Clark MS (ISC). This widespread Costa Rican bamboo differs etén of Guatemala and in Chiapas, in a number of respects. The most conspicuous of these are listed in Table 1. Unfortunately R. bartlettii is incompletely known and the type is in the flow- ering state only. The original description con- tains no information concerning the number of flowering branches per node, the culm height and diameter, or the nature of the culm leaves. The differences between the Costa Rican plants and POHL— RHIPIDOCLADUM 275 the type of R. bartlettii that can be determined from available material are mostly spikelet char- acters, but they appear to indicate that these two populations represent distinct species, a conclu- sion supported by their apparent geographical disjunction. This species occurs in scattered small popu- lations at lower and middle elevations in moist areas of central and southern Costa Rica. I have Observed various of these populations in the veg- etative state at different times between 1968 and 1982. One of these colonies, Pohl & Pinette 13207, collected in 1976, at that time lacked new culms or culm she aths, indicating of blooming. Specimens 13182, 13149, and 14161 are all from the same population at the type locality east of the Río Pacuare. Blooming was first observed at this locality in 1982. The Río Tuís population, which was in bloom at the same time, was about 5 km distant from the type locality. From this collection site, we could ob- serve massive gregarious blooming at a distance, along the Río Tuís. Canes ofthis species arch and droop, the upper young internodes contained liquid. The plants are graceful and have potential as an ornamental. Rhipidocladum panamense Font, Sp. NOV. TYPE: ma n TOV Goofy La (holotype, ISC; isotype, MO). Figure j-l. Gramen lignosum; culmi scandentes, ad 5 m longa, cylindrati, cavi, 5-7 mm diam uli floriferi supra 200 in quoque nodum, ad 35 cm longi; folia ramulo- rum 2-4, plana, 3-6 c -3 , linearia. Inflores- centia racemosa Esp tenuis. "Spiculae 5-7, 12- 16 mm longae; gluma pri -3.5 mm longa; se- cunda 4-7 mm, 5-nervata, anda 3-4. 5 mm m onga; lem- mata fertilia 2, 5.5-6 mm, 7—9-nervata, arista 3-6 mm longa; lemma dorsalis bala: flosculi radical 2-3. A R. clarkiae recedit foliis tenuiores, spiculis breviores, et rami supra 200 in complemento. Ligneous bamboo; culms clambering, to 5 m long, the basal diameter unknown; flowering in- ternodes hollow, cylindrical, glabrous, 5-7 mm thick; branches 200-230 per node, very slender, to 35 cm long, including inflorescences; nodes swollen. Branch leaves 2-4 per branch; sheaths glabrous or sparsely puberulent; auricular bris- tles sparse or absent, ca. 2 mm lo to the pseudopetiole, glabrous or sparsely his- 276 pidulous above, glabrous or puberulent beneath; 1 strong nerve near one margin. Inflorescence a very slender unilateral raceme, 4-12 cm long, terminal on a leafy primary branch or a short secondary branch arising from a middle node. Spikelets 5—7 per raceme, appressed, borne 5-20 mm apart, 12-16 mm long, very slender, scarcely compressed, disarticulating above the sterile lemma and between the fertile florets; first glume 2.5-3.5 mm long, narrowly ovate, awnless; sec- ond glume 1-nerved, 4-7 mm long, clasping, 5- nerved, the awn 3.5—4.5 mm long; sterile lemma narrowly ovate, 5.5 mm long, 7-nerved, the awn 3-4 mm long; fertile florets 2, with 2-3 rudi- mentary, convolute florets above; fertile lemmas narrowly ovate, 5.5-6 mm long, 7—9-nerved, the awn 3-6 mm long, the back hispidulous along the margins and near the apex; paleas of fertile lemmas longer or shorter than the lemma, ciliate at the apex; anthers not seen; style 1; stigmas 2. Forests, provinces of Darién and Panamá, Panama. This species is similar to R. clarkiae, both having large numbers of branches at the flowering nodes, which separates them from all other known species of Rhipidocladum. Rhipi- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 docladum panamense differs from R. clarkiae in the much narrower and shorter leaf blades, smaller spikelets, and the much larger number of flowering branchlets. Representative flowering specimens. PANAMA. PROV. PANAMÁ: Goo e, 16 Aug. 1967, Dwyer & Hayden 8037 (MO); 2 mi. above Goofy Lake, 16 Aug. 1967, Stimson 5377 (MO); 7 mi. N of Cerro Azul, road to Cerro Jefe, 2,600 ft., 13 Nov. 1965, Blum, Godfrey & Tyson 1810 (MO); Cerro Azul, 2,000 ft., 3 Nov. 1965, Tyson 2116 (MO). Nee 10711 (ISC, MO) from the Río San Felix, Chiriqui, is similar to the preceding, but has much onger, awnless lemmas. Until specimens with intact spikelets can be obtained, I cannot place . — LITERATURE CITED McCLuRnE, F. A. 1973. Genera of bamboos native to the New World. (Gramineae: Bambusoideae). Smithsonian Contr. Bot. 9: i-x, 1-148. SODERSTROM, T. R. Observations on a fire- adapted bamboo of the Brazilian Cerrado, Acti- nocladum verticillatum V or Bambusoideae). Amer. J. Bot. 68: 1200- SYSTEMATICS OF THE SOUTHERN AFRICAN GENUS GEISSORHIZA (IRIDACEAE —IXIOIDEAE)! PETER GOLDBLATT? ABSTRACT Geissorhiza is a large genus of small, corm bearing perennials endemic in the Cape Province of i of some 25 over the 52 species of true Geissorhiza admitted by Foster (1941) in his revision of the os and the eight species placed by Lewis ( 1 941)i in E ngysiphon, a p here reduced to synonymy. ssorhiza (34 spp.) and Weihea y Spp.) are recognized, each including several sections. Pici ns is seen as a late Tertiary genus EM pog differentiated in upland southeastern Africa when much of southern Africa including pe Region was still heavily forested. The continuing deterioration of the climate in the Pliocene fa ite i Cape probably gave the impetus to the extensive radia nt apii agoe belt in this area, where most species, including all of the more speciali taxa, e taxa are concentrated along the relatively well-watered southern Cape coast, wher little isis jai ccurred and the primitive taxa a appear to have differentiated slowly by a process of phyletic a ea The geography of Geissorhiza is analyzed in detail and several aaa centers of endemis rre- sponding closely with Wei k’s phytogeographical ppp eain are luni, uius The major trends i in the evolution of peat include: the development of imbricate corm tunics from the basic con- centric type; the reduction or extreme elongatio N the perianth tul ; the development of floral zygomorphy; and the elaboration of the leaf lamina, whic ome T ri id grooved (section Geissorhiza), or may have raised and winged ciliate margins winge b (section Ciliata) or thickened margins and midrib (especially sections nece o d Props. Cytolosica : studies have brought the number of species in which the chromosome number half the genus. Basic chromosome number is x — 13, and while most species are e diploid, kie d has been recorded i ina few species including the triploid vegetative apomict, G. bolusii. . The conser- vation status almost all those c to the coastal plain between Cape Town and Piketberg, an intensively farmed area rich in speci Geissorhiza is a large genus of medium to small, sorhiza is a member of Ixioideae, a predomi- corm bearing plants, restricted entirely to the nantly African subfamily, strongly developed in winter rainfall region ofthe south and westcoasts southern Africa, and is distinguished by its woody of southern Africa. It is centered in the moun- to papery (rarely fibrous) corm tunics, asym- tains and western coastal belt ofthe southwestern metric corms, herbaceous floral bracts, and long, Cape, with species extending as far east as Gra- exserted style (included in two species) with short hamstown in the eastern Cape and as far north recurved style branches. It is most closely related as Steinkopf in northern Namaqualand. Geis- to Hesperantha (ca. 55 spp.—Goldblatt, 1982a, ! Support for this research from the U. S. National hri — grant DEB-78-10655 and 81-19292 is gratefully acknowledged. I also wish to f Forestry and the Cape Department of Nature Environmental Conservation for their sistas pe this project and for collecting permits. For their assistance in the field or in providing me with material for study I want to thank Dee Snijman, qo dei Herbarium, Cape Town; Ion Williams, Vogelklip, Hermanus; J. H. J. Vlok, Department of Forestry, Georg Neil MacGregor, Glenlyon, Nieuwoudtville; Mike Viviers, Department of Forestry, Cedarberg; and Pio d Elsie Esterhuysen, Bolus Herbarium, University of Cape Town, who helped me extensively with the rare high mountain species and after whom two species have been named. The support and hospitality provided by John Rourke and his staff, Compton Herbarium, Kirstenbosch, Cape Town in the course of field work is acknowledged here with gratitude. I also wish to thank Margo Branch for the many excellent illustrations made for this study. Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. ANN. Missouni Bor. GARD. 72: 277—447. 1985. 278 1984) from which it differs mainly in its style and style branches. In addition, Hesperantha is unusual in the family in having introrse anthers at anthesis. Since its treatment in “Flora Capensis” (Baker, 1896) in which 29 species were recognized, Geis- sorhiza has been completely revised only by Foster (1941). Foster admitted 55 species to the genus, three subsequently transferred to Gladi- olus, including the only species he recognized from Madagascar, G. bojeri (Goldblatt, 1982b). A second Madagascan species Geissorhiza am- bongensis, which he regarded as “doubtful or un- known," is here also excluded from Geissorhiza but its correct generic position is uncertain. De- Geissorhiza has, for reasons outline poorly understood. Foster’s revision was based on abe sassa material only but he did not ex- ine th in South African din and was thus unaware of several un- described species in these collections. The revi- sion was also in part unnatural, combining un- related taxa in some subsections, and the key was difficult to use. In addition, extensive collecting in South Africa, especially in recent years, has resulted in the discovery of many species that were clearly undescribed. A few of these are local lowland endemics but most are montane species of middle and upper elevations, many discov- ered only in the last 40 years, as the Cape moun- tain flora was systematically explored, largely by the Cape botanist, Elsie Esterhuysen. Eighty-one species in two subgenera and 12 sections are recognized in this treatment (Table 1), in which Engysiphon Lewis (8 spp.) is reduced to sectional rank. Some 12 species admitted by Foster are reduced to synonymy, two to subspe- cies status, while two of Foster's varieties are raised to specific rank, and 29 species are de- scribed here for the first time. The subgeneric classification followed here is novel, being based primarily on the characteristics of the corm rath- er than the leaf which were considered of more fundamental importance by Foster. Both corm and leaf structure vary to an unusual extent : Geissorhiza and these aspects are examined 1 detail in this study. Chromosome cytology, also investigated here, is relatively uniform with x — ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 13 the basic chromosome number. Most of the 40 species known cytologically are diploid, but G. bolusii is an apomictic triploid, and three species are tetraploid, or have tetraploid races. RELATIONSHIPS AND SYSTEMATIC POSITION OF GEISSORHIZA Geissorhiza is a member ofthe largely African subfamily Ixioideae, an alliance defined by a spi- cate inflorescence, the sessile flowers of which are subtended by a pair of opposed bracts. The tepals are always united in a tube which is often well developed, and the rootstock is a basally rooting corm that typically has a stele (De Vos, 1977 It is reasonably well established on ron logical and cytological grounds (Lewis, 1954; Goldblatt, 1971, 1982a, 1984) that Pau i is most closely related to Hesperantha, a wide- spread African genus of some 55 species, also centered in the southern African winter rainfall region. The apparently primitive species of both general with light brown, firm to woody, concentric tunics, her- sl. floral bracts, small actinomorphic flow- ers, and membranous capsules with numerous turbinate to angular seeds. Unspecialized species of both genera can in fact only be distinguished by their different style and style branches and anther dehiscence in the actinomorphic flowered species. In Geissorhiza, the long style is well- exserted from the perianth tube and divides to- wards the apex of the anthers into three short recurved branches (near the base of the anthers periant branches. The anthers of Hesperantha are in- trorse at anthesis, contrary to the normal con- dition for species of Iridaceae with actinomor- phic flowers. The two genera share a common basic chro- mosome number, x = 13, a number unusual in Iridaceae, but also found in the related Schizo- stylis (mono- or di-typic) and in some species of the apparently unrelated Romulea (De Vos, 1972). Lewis (1954) placed Geissorhiza in a sub- — FIGURES 1-6. From left to right and top to bottom.— 1. Geissorhiza elsiae.—2. G. corrugata.—3. G. kar- ooica. —4. G. cedarmontana. — 5. G. nubigena. —6. G. longifolia. N - x < ° [o < t; 1 = = < - m Q - Q Q 280 tribe Ixiinae close to Hesperantha in a lineage that also includes G/adiolus and the small, highly specialized genera such as Homoglossum and Anomalesia, closely allied to G/adiolus. In a more refined classification (Goldblatt, 1971), Geisso- rhiza is placed together with Hesperantha, Schizostylis, and the monotypic Melasphaerula in Hesperanthinae, a classification that still seems to best reflect the relationships of these genera. Melasphaerula (x = 11) is probably isolated in the alliance, if it correctly belongs here, but it has the woody corm tunics that define the sub- tribe. The difference between Geissorhiza and Hes- perantha appears to be based on relatively minor features but nevertheless, they appear on phe- netic grounds to comprise monophyletic assem- blages. The patterns of variation are coherent in both genera and are consistent with the conten- tion that they are indeed natural groups, even though Geissorhiza can at present only be defined negatively, with respect to Hesperantha, by a combination of unspecialized characteristics. The unspecialized species of Hesperantha have pale, white or pink flowers and are usually eve- ning blooming and strongly sweet scented. The genus is believed to be a basically moth polli- nated group but day blooming species with col- ored flowers are probably bee pollinated (Vogel, 1954; Goldblatt, 1984). In contrast, the species of Geissorhiza are always day blooming and typ- ically scentless, and the genus is believed to be largely bee pollinated (Vogel, 1954). MORPHOLOGY OF GEISSORHIZA Species of Geissorhiza are small, geophytic corm bearing perennials. Actual age is difficult to determine but plants with up to 12 annually produced corm tunic layers are known. With the exception of G. outeniquensis, species grow ac- tively in the wet seasons of the year, usually win- ter and spring (or summer at higher altitudes), flower in spring or summer, and become dor- mant during the dry, later spring or summer and autumn. Geissorhiza outeniquensis is apparently evergreen and has a very reduced underground storage organ. It grows in permanently moist montane habitats and appears to have no dor- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 mant period during the year. Geissorhiza four- cadei flowers in the autumn but it is restricted to sites that remain moist throughout the sum- mer or all year. All parts of the plant body have some spe- cialized features and these are discussed in the following pages, under the main headings of corm, leaves, stem, inflorescence, flower, stamens, gy- noecium, and fruit CORM The corm of Geissorhiza consists of a swollen internode with an apical bud, typically covered with hard, woody to firm and papery tunics. The live portion of the corm is a white (Fig. 13A—E) asymmetric body, being somewhat flattened on one side or towards the base, and with a peculiar downward oriented projection at the lower end of the flattened side, from which the roots are produced. The corms vary considerably in the genus but fall into two main groups, those with concentric or with imbricate tunics (the termi- nology of Foster, 1941, 1948). The basic tunic type is assumed to be one with concentric layers, a type shared with the related genus Hesperantha, in which corm and tunic morphology have recently been described in de- tail (Goldblatt, 1982b). Here the overall shape is often asymmetric (Fig. 13), reflecting the in- ternal ri but the pene of Ie lateral projection is no y. Corms with concentric fanice are usually small of con- stant size, and the older tunic layers split irreg- ularly from base and apex into somewhat elliptic sections which completely enclose the new tunic layers produced in successive years. The tunics are usually light to middle brown in color. In the second major corm type in Geissorhiza, the outer tunics cover the inner above only, and are thus imbricate. The corm shape is usually nearly symmetric and more or less ovoid with a small flat basal part. The tunics are usually dark brown to blackish, but in G. similis and G. scil- laris they are a light brown color as in species with concentric corm tunics. In either case, the layers are notched below in a regular pattern that gives the corms their very characteristic appear- ance (Fig. 13F-K). The upward displacement of FIGURE 7-12. G. .— Hom —10. G — From left to right and top to bottom.— 7. Geissorhiza malmesburiensis. —8. G. barkerae.—9. G. mathewsii.—11. G. radians.—12. G. splendidissima. GOLDBLATT — GEISSORHIZA 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 TABLE l. The species of Geissorhiza arranged in systematic order, in their sections. Brief distribution and habitat data are also listed for convenience. Subgenus Weihea (Baker) Goldbl. . G. odio ep v G. elsiae G G. zc (Levis) Goldbl. G. outeniquensis Goldbl G. fourcadei (L. Bolus) oe noU T2 — G. foliosa Klatt . G. nigromontana Goldbl. . G. delicatula Goldbl. . G. bracteata Klatt . G. nana Klatt . G. setacea (Thunb.) Ker . G. ornithogaloides Klatt — — N — subsp. marlothii (Foster) Goldbl. subsp. ornithogaloides 13. G. malmesburiensis Foster 14. G. geminata E. Meyer ex Baker 15. G. ovalifolia Foster 18. G. ovata (Burm.f.) Asch. & Graeb. 19. G. corrugata Klatt 20. G. spiralis (Burch) De Vos ex Goldbl. 21. G. karooica Goldbl. 22. G. pusilla (Andr.) Klatt Section Weihea—18 spp. S Cape between enum and Uitenhage Kammanassie Little Karoo, pee to Uitenhage wet shady sites, Outeniqua Mts., George district Swartberg, Outeniqua and Tsitsikamma Mts., Robinsons ss to Humansdorp slopes flats at the foot of the Langeberg Great Swartberg, along streams mts. of the interior S Cape S Cape near Albertinia to E Cape at Grahamstown Caledon district to Riversdale, clay slopes and flats SW Cape, wet sandy widespread in the interior Cape region, Cedarberg E to the Long Kloof Paarl district to Caledon and Bredasdorp Malmesbury, sandy slopes wet sites, interior SW Cape damp sites, SW Cape mts. damp shady sites, SW and W Cape mts SW and W Cape mts., sandy soils and rocks SW and S Cape, sandy soils, mainly montane Section Tortuosa Foster—3 spp. Calvinia district, shale flats Sutherland district, W Karoo flats Matjesfontein Karoo, S-facing shale slopes Section Pusilla Goldbl.—1 sp. damp shady places, Peninsula and surrounding hills Section /ncludanthera Goldbl.—2 spp. 23. G. esterhuyseniae Goldbl. 24. G. cedarmontana Goldbl. Great Winterhoek Mts., rocky S slopes at 2,000 m Central Cedarberg above 1,200 m, steep S slopes Section Angustifolia Goldbl.— 11 spp. 25. G. lithicola Goldbl. 26. G. purpurascens Goldbl. 27. G. humilis (Thunb. 1 ~ 28. G. darlingensis Gol 29. G. hispidula o DUE 0 pappei Baker intermedia Goldbl. unifolia Goldbl juncea (Link) A. Dietr. furva Ker ex Baker stenosiphon Goldbl. UJ ` w . >» Mx ala a: Ww e Q . umbrosa Lewis alticola Gol oT Schltr. cataractarum Goldbl. nubigena Goldbl. w = 2 ° - QQ oo lower slopes of the Kogelberg Mts., in rocky ground SW Cape, medi flats at isolated sites between Siilenbovel and Pike sandy flat dA Cape Peninsula and Flats local in damp flats around Darling sandy flats and slopes, “otoq to Albertinia rare, sandy mt. soils, SW rare, mts. between “us and Porterville high elevations in the Cedarberg sandy flats and slopes, Caledon to Pakhuis Pass SW Cape, stony lowlands between Paarl and Gouda Hexberg, Cold Bokkeveld Mts. Section /xiopsis Goldbl.—5 spp. mid to upper elevations, Peninsula and Hottentots Holland to the Cedarberg higher peaks, Bains Kloof to Wemmershoek mp and marshy slopes, Bains Kloof to Bredasdorp waterfalls and damp cliffs, Bettys Bay to Hermanus higher mts., Gt. Winterhoek to Kogelberg Mts. 1985] GOLDBLATT — GEISSORHIZA 283 TABLE l. Continued. Section Engysiphon (Lewis) Goldbl.— 7 spp. 41. G. brevituba (Lewis) Goldbl. Piketberg Mts. 42. G. schinzii (Baker) Goldbl. mts. of the Caledon district 43. G. jd dil a) Goldbl. stony often shale soils, Michells Pass to Gifberg 44. G. confusa Goldbl. stony mountain soils, French Hoek to Gifberg 45. G. bonae- ipsi Gla S Cape Peninsula 46. G. tenella dbl. sandy flats and dunes, Darling to Bredasdorp 47. G. exscapa ae Goldbl. sandy flats and plateaus, Blouberg to Hondeklipbaai Subgenus Geissorhiza Section /ntermedia Goldbl.—2 spp. 48. G. similis Goldbl. sandy slopes and flats, Cape Peninsula to Bains Kloof 49. G. scillaris A. Dietr. sandy slopes and plateaus, Caledon to the Cedarberg Section Geissorhiza—10 spp. 50. G. imbricata ls la Roche) Ker subsp. im Cape Peninsula to Bredasdorp tae oe abi Goldbl. W Cape, wet sandy flats 51. G. purpureolutea B W Cape coastal belt, wet flats 52. G. barkerae Goldbl. wet marshy sites at foot of the Piketberg 53. G. louisabolusiae Foster Olifants River Valley around Citrusdal 54. G. brehmii Eckl. ex Klatt seasonal pools in the SW Cape 55. G. sulphurascens Foster wet sandy soils, Nieuwoudtville Escarpment 56. G. minuta Goldbl. rare in wet sites in the Pakhuis Mts 57. G. eurystigma L. Bolus rare, Darling to Kalabaskraal 58. G. mathewsii L. Bolus rare, around Darling, vlei margins 59. G. radians (Thunb.) Goldbl. damp sites, SW Cape from Gordons Bay to Saldanha Section Monticola Goldbl.—8 spp. 60. G. burchellii Foster middle elevations, Bains, Kloof to the Langeberg 61. G. grandiflora Goldbl. mts. among rocks, Gt. Winterhoek to the Langeberg 62. G. tabularis Goldbl. semi marshy places, Table Mt., Cape Peninsula 63. G. ramosa Ker ex Klatt mt. slopes, Hermanus to Tulbagh 64. G. bryicola Goldbl. waterfalls and stream edges, ue Mts. 65. G. scopulosa Goldbl. high altitudes, Hex River 66. G. ciliatula Goldbl high altitudes, Cedarber 67. G. pseudinaequalis Goldbl. damp mt. slopes and cliffs Ceres to Stellenbosch Section Planifolia Goldbl.—4 spp. 68. G. aspera Goldbl. sandy flats and slopes, W Cape 69. G. inaequalis L. Bolus W Cape mts., Bains Kloof to Nieuwoudtville 70. G. monanthos Eckl. W Cape coast, often on granite outcrops 71. G. tulbaghensis F. Bolus clay flats, Tulbagh Section Ciliata Goldbl.— 10 spp. 72. G. namaquensis Barker Namaqualand, Kamiesberg to S Richtersveld 73. G. kamiesmontana Goldbl. local in the Kamiesberg above 4 74. G. divaricata Goldbl. Nieuwoudtville ibm and ‘Gifbers 75. G. subrigida L. Bolus Nieuwoudtville Escarpm 76. G. heterostyla L. Bolus widespread, S Cape to Manada and N through the Rog- geveld to S Namaqualand 77. G. arenicola Goldbl Nieuwoudtville Escarpment and Gifberg 78. G. splendidissima Diels near Nieuwoudtville 79. G. inflexa (de la Roche) Ker SW Cape. gcn to iie a shale slopes and flats 80. G. erubescens Goldbl. Pakhuis Pass, shale 81. G. leipoldtii Foster Clanwilliam to ern valley, shale soils FIGURE 13. furva. — B. G. ovata. — C. G. juncea. —D F, G. G. inflexa with tunic layers progressively removed a onanthos with numerous cormlets ea the Bc G. aspera (all more or less life grandiflora.—J. G. m size except A, x0.5). the older tunics is usually easy to recognize, but occasionally the displacement is minimal and the corm type can be difficult to determine. The difference in the corm types in Geissorhiza is so marked and is correlated with other im- portant features in that it is considered a fun- damental character in the genus and the basis for the recognition of subgenera. Species with con- centric tunics are assigned to subgenus Weihea while those with imbricate tunics fall in subgenus Geissorhiza. In both subgenera, species growing in peren- nially moist to wet habitats tend to have much softer textured tunics, and this is believed to be a secondary development. This occurs in other genera with typically hard corm coverings and cannot be considered significant above the species or sectional rank. This condition is most extreme in Geissorhiza outeniquensis, and G. cataracta- rum, species that grow close to permanent water. The corm coverings of G. hesperanthoides, G. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Corm morphology in Geissorhiza. A-E. Subgenus Weihea. F-K. Subgenus Geissorhiza.—A. G. G. ornithogaloides de i Hole pu —E. G. he speranthoides. — naked corm.—H. G. scillaris.—I. G. nubigena, and some forms of G. umbrosa are most unusual in being composed of reticulate fibers. The relatives of these closely allied species have soft textured, concentric tunics, and they are clearly members of subgenus Weihea. In subgenus Weihea, the corms of section En- gysiphon are unusually large and the tunics par- ticularly hard and persistent so that species of this group can often be recognized from their corms alone. In this subgenus, the corms of Geis- sorhiza ornithogaloides subsp. ornithogaloides and G. malmesburiensis are also distinctive (Fig. 13) in having a broad, oblique to horizontal base, and the tunic layers are serrated below In subgenus Geissorhiza, the corms are uni- form in sections Geissorhiza, Planifolia, and Cil- iata, bei and less persistent, possibly reflecting their mon- tane and less extreme habitat. The pale tunics of G. similis and G. scillaris, assigned to section 1985] Intermedia, have already been mentioned. It is not clear whether the corms of these two species represent a convergent development or are sim- ply a variation or modification of the basic im- bricate type of subgenus Geissorhiza. In the ab- sence of further information, it is regarded as the latter and accordingly the section is included in this subgenus. LEAVES Cataphyll. There is usually a single trans- parent and membranous cataphyll sheathing the stem base below ground. It is often lacking or fragmentary in dry material and seldom distinc- tive. Occasionally, the cataphyll is firm textured, persistent and brown, and forms a characteristic feature of species like Geissorhiza erubescens, G. divaricata, G. parva, and G. ovata. Foliage leaves. These are defined as having a free, monofacial apex as opposed to stem bracts which are entirely sheathing. There may be from several to only three, less often two, and one only in Geissorhiza unifolia. The leaf number in most species is constant. The primitive state 1s pre- sumed to be several and indeterminate as in G. foliosa, G. bracteata, and G. inconspicua (all sec- tion Weihea). Leaf number has stabilized at three basal or nearly basal and long leaves and a third cauline and shorter, partly sheathing leaf. This uppermost leaf occasionally has a somewhat in- flated and spathe-like sheath, or it may be re- duced and entirely sheathing in which case it is regarded as a bract rather than a leaf. In G. scil- laris only two leaves are produced, the lower with a long free lamina and the upper also long but sheathing for almost its entire length. The basic leaf type in the genus, as for Irida- ceae in general, is ensiform and equitant with a plane lamina in which the margins and the mid- rib can be readily distinguished but are not sub- stantially thickened or raised. This basic leaf ground plan has been more extensively modified in Geissorhiza than in any other genus of the "x o) with the possible exception of G/adiolus (Fig. 1 The uS modification is a narrowing from ensiform to linear, a common feature of species in the most primitive section Weihea. Among the species with plane leaves, G. pusilla stands out in having partly pubescent leaves. The species is unusual in several ways and has been assigned to its own section, Pusilla. Unusual broad, often GOLDBLATT — GEISSORHIZA wings and edges of raised midrib ciliate (whole plant life size; leaf sections much enlarged). ovate and prostrate or nearly prostrate leaves are found in G. ovata and sometimes G. parva. A more complex development is a thickening of the margins and the midrib so that the leaves become distinctly 2-channelled on each surface (Fig. 14A, C), and the edges of the thickened mar- gins are sometimes minutely ciliate. Leaves of this type o some species of subgenus Weihea, in sections Angustifolia, Ixiopsis, and Engysiphon as well as in subgenus Geissorhiza. In G. exscapus, G. tenella, and G. bonae-spei of section Engysiphon, the margins are raised, forming wings held at right angles to the blade while the midrib is hardly raised at all. These leaves are thus H-shaped in cross section (Fig. -— + A narrowing of the blade is evident in Geis- sorhiza humilis and G. hispidula, and in both these species the leaves are sticky so that soil adheres to the lamina even when dry. With the contraction of the blade width, the longitudinal channels become narrowed to thin grooves. Such leaves are found in section /xiopsis of subgenus Weihea and occasionally in species of other sec- tions, notably G. radians and G. sulphurascens of section Geissorhiza. In a few species, e.g., G. juncea, G. furva, G. brehmii, G. louisabolusiae, G. stenosiphon, and G. fourcadei, the blade is 286 terete (Fig. 14D), but the narrow grooves, indi- cating their ancestry, are preserved and can be seen in cross section or even in intact leaves un- der the dissecting microscope. In subgenus Geissorhiza, the leaves of species in section Ciliata are distinctive in having the margins extended to form wings held at right angles to the blade (Fig. 14F), the midrib also raised and winged, and their edges are ciliate to pubescent. In G. subrigida, and robust forms of G. inflexa, G. divaricata, and sometimes other species of the section, secondary veins may also be raised and ciliate. The leaves of most species of section Geisso- rhiza are unusual in having several veins as well as the margins heavily thickened so that the blade is several ribbed and grooved (Fig. 14E). The sheathing part of the third and uppermost leaf in this alliance is inflated and also heavily ribbed and grooved, and this character is preserved even when the lamina has become narrow and only 2-grooved (G. radians, G. sulphurascens, forms of G. imbricata) or terete (G. brehmii, forms of G. louisabolusiae). Bract leaves. These appear to be modified upper leaves in which the free part of the lamina has been suppressed. They range from green and leaf-like, but entirely sheathing, to membranous and even scale-like. The presence of a bract leaf is typically constant for a species and is present even when the stem is unbranched. Such bracts are a diagnostic feature of G. scillaris, G. heter- ostyla, and all species of section Monticola. STEM The stem is typically erect, smooth, terete, and produced well above the ground. It is usually sheathed below by leaf bases, and in many species the third leaf is specialized as a sheath which envelopes the lower part of the stem. The upper half of the stem may be naked, or bear a spe- cialized bract leaf (see above). The stem is dense- ly (or occasionally sparsely) ciliate in all the species of section Planifolia, in several of section Ciliata, and in G. ciliatula and G. scopulosa (sec- d in two isolated species, G. intermedia and G. unifolia, of subgenus Weihea, section Angusti- olia. Branching is variable, but certain species typ- ically do not produce any branches, while exten- sive branching is characteristic only of several species of section Monticola, such as G. ramosa, G. burchellii, and G. tabularis. Branching is sup- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 pressed in all species of sections Angustifolia and Ixiopsis. INFLORESCENCE The inflorescence in Geissorhiza is, with a few exceptions, a spike of several flowers and may be more or less straight to flexuose. The number of blooms is variable depending on age and health of the plants and seasonal and soil conditions. A few scattered species in subgenus Weihea have solitary flowered stems, thus technically not true spikes. These include all three species of section Tortuosa (G. corrugata, G. karooica, and G. spi- ralis) and G. delicatula and G. fourcadei (section Weihea). The spike is typically drooping in bud, and straightens gradually from the base as the flowers open progressively. This drooping is seldom ev- ident in dried material and although not deter- mined for all the species, appears to be a feature of the genus. The flowers are subtended by paired, opposed bracts, a larger outer enclosing the smaller inner. The inner bract (sometimes called a bracteole), used very little as a taxonomic character, is a double structure with two main veins and a forked apex. While the inner bract may be as long or slightly longer than the outer bract, it is always smaller and narrower. Only in section Engysi- phon are the inner bracts substantially shorter than the outer, and this is one of the diagnostic features of the section. The outer bracts are usually herbaceous with a hyaline upper margin, and firm in texture, but blooming. In subgenus Geissorhiza, sections Planifolia and Ciliata, the bracts become dry and ferrugineous in the upper half. This is particu- larly well developed in G. aspera, G. monanthos, G. inflexa, and G. subrigida. There is little to distinguish the bracts of other species with the exception of the members of section Engysiphon in which they are very large, up to 50 or 60 mm in some species, but only 15-20 mm long in G. brevituba, and much longer than the quite mem- branous inner bracts. Geissorhiza humilis and G. hispidula are distinctive in having sticky bracts, to which sand adheres even when the plants are dried. FLOWER Most species of Geissorhiza have a simple, ac- tinomorphic flower with a short, straight, cylin- dric perianth tube and horizontally extended te- 1985] MLE GOLDBLATT — GEISSORHIZA ve | mm i z 2.5 FiGurE 15. Floral features of Geissorhiza. —A.G mens and style. —D 9 I th deflexed —C. G. grandiflora, zygomorphic flower with declinate nches.—E. G. eurystigma, the broad ns. ciliate style branches found in this species (A-C life size; D-E x pals (Fig. 15A, B). The tube generally reaches to about the apex ofthe bracts surrounding the tube, but after fertilization it may become exserted a few millimeters due to the enlargement of the ovary. The style is slender and exserted well be- yond the mouth of the tube and ultimately di- vides into three relatively short, recurved branches. The three stamens are usually sym- metrically disposed, with the filaments inserted ow the mouth of the perianth tube and well- exserted from the tube. The stamens vary con- siderably and are dealt withi in more bien below. Longer perianth t rmore than 1 or 2 mm from the bracts are jeune and are scattered throughout the genus. In subgenus Weihea, G. ovata (section Weihea), G. nubigena (section Ixiopsis), G. stenosiphon (section An- gustifolia), and both G. esterhuyseniae and G. cedarmontana (section Includanthera) have un- usually long perianth tubes. In section Engysi- phon, the long perianth tube is accompanied by long bracts, and the tube may reach only to the apex of the bracts (G. schinzii, G. bonae-spei) or be exserted for some distance (G. exscapa, G. confusa, G. tenella Short perianth Gites that reach only to about the midline of the bracts are regarded as derived since they occur only in specialized species or sections, while moderately long tubes are found in the less specialized species and in related gen- era. These short tubes are characteristic of a few species in subgenus Weihea, notably G. alticola (section [xiopsis), in sections Planifolia and Cil- iata, and in several species of section Geisso- rhiza, all subgenus Geissorhiza. In these species, the bracts become dry at flowering time and are folded back in the midline by the outspread te- pals. Zygomorphic flowers appear to have evolved repeatedly in both subgenera (Figs. 5, 6, 11, 12, 15). In the zygomorphic flowers in Geissorhiza, the stamens are unilateral and decumbent, and lie above the anterior tepal (Fig. 15C). The style is also decumbent, and it lies under the stamens. This situation is unusual in Iridaceae where zy- gomorphic flowers typically have erect and uni- lateral stamens, arcuate under an erect and often hooded anterior tepal. In subgenus Weihea, zy- gomorphy is developed in G. roseoalba and G. outeniquensis (section Weihea), in G. karooica (section Tortuosa), and in all species of section Engysiphon, while in subgenus Geissorhiza, zy- 288 gomorphic flowers are found in G. tulbaghensis and G. monanthos (section Planifolia), in G. ra- dians and G. barkerae (section Geissorhiza), and in G. grandiflora (section Monticola). Flower colors of white to cream or shades of violet to purple are most common in Geissorhiza (Figs. 1-12), but all colors except shades of or- ange occur. Except for occasional mutants, pop- ulations and even species are uniform in their flower color. Geissorhiza inflexa stands out in having populations of white, purple, pink, or deep red flowers. Unusual color combinations occur in several species. In G. barkerae, most forms of G. purpureolutea and some of G. brehmii and G. imbricata (all section Geissorhiza), and in some populations of G. inflexa the cream to yellow flowers have a dark purple to brown center. Geissorhiza eurystigma, G. radians, and G. ma- thewsii (section Geissorhiza) and forms of G. monanthos have dark violet flowers marked with contrasting concentric bands of white and dark red (Figs. 10-11) STAMENS The three stamens typically have well-devel- oped and equal filaments inserted in the perianth tube shortly below the mouth. The two species of section 7ncludanthera, G. cedarmontana and G. esterhuyseniae, are remarkable in having very short filaments inserted near the base of the peri- anth tube, so that the stamens are entirely in- cluded in the tube. An unusual feature of subgenus Geissorhiza is the presence in many species, of unequal fila- ments, one being consistently shorter, by at least 0.5 but up to 3 mm, than the other two (Fig. 15B). While this development appears to have arisen independently in each section (excepting section Intermedia where the character is not present), it is probably fundamental in the sub- genus, although only expressed in certain species. In section Monticola, unequal filaments occur in all species except the very dwarf G. ciliatula, while in section P/anifolia, unequal filaments oc- cur consistently only in G. inaequalis and oc- casionally in G. aspera and G. monanthos. In section Geissorhiza, only G. barkerae and in sec- tion Ciliata, G. heterostyla consistently have a shorter filament, while this occurs occasionally in G. louisabolusiae (section Geissorhiza) and leipoldtii (section Ciliata). The anthers are usually somewhat longer than the filaments, and erect to ascending. In those ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 species with declinate stamens, the anthers are erect and thus held at right angles to the decum- bent filaments. In G. esterhuyseniae and G. ce- darmontana (section Includanthera), the anthers are included in the perianth tube. Anther size is a generally useful taxonomic character as it is fairly constant in a species, easy to measure, and a good indication of overall flower size. GYNOECIUM Style. The style is filiform, slender, and well exserted from the perianth tube. Ultimately it divides into three comparatively short and slen- der recurved branches (Fig. 15A-D). This char- acter distinguishes the genus from the allied Hes- perantha and varies very little. In Geissorhiza cedarmontana and G. esterhuyseniae (section In- cludanthera), however, the style is short and en- trely included in the perianth tube while in the striking G. stenosiphon, the style may divide any- where between the mouth of the perianth tube and the base of the anthers. The style typically diverges to some degree from an upright position in all species, but is decumbent and placed under POR 2 : os gen 15C). In some populations of G. heterostyla, in- dividuals may have either long (i.e., normal styles), medium, or short styles that reach only to the apex of the perianth tube (Bolus, 1930), hence the name given the species by Bolus. The anthers do not vary at all in length, so that the condition is not true heterostyly. This is presum- ably an adaptation to promote outcrossing. Two species, Geissorhiza eurystigma and G. mathewsii (section Geissorhiza), have distinc- tive, broad flat style branches (Fig. 15E), de- scribed by Lewis (1954) as flat, thick, and cren- nate edged. The style branches ofthese two species are so unusual that they can immediately be rec- ognized from this character alone. Ovary. The oblong to ovoid ovary has little taxonomic value, and, apart from minor size dif- ferences, there is little to distinguish species on this character. FRUIT The fruit is a thin walled loculicidal capsule, typically more or less ovoid-oblong in shape. The ovary develops rapidly after fertilization, and generally grows to reach the apex of the bracts before maturing. The bracts enclose, to some ex- tent, the young fruits but in the later stages of development the bracts become dry and broken. 1985] The species of section Engysiphon stand out in having exceptionally long capsules that are more or less fusiform in shape. The capsules are en- tirely included in the long bracts of these species. eds are basically turbinate to globose in shape with a persistent funicle and a testa of uncon- torted epidermal cells with smooth surfaces. De- viations from this basic type include increasing compression, resulting in a more or less angular to irregular shape and the wrinkling and crum- pling ofthe epidermal cells (Wagner & Goldblatt, 1984). Although the seeds of only a few species have been examined, seed structure appears rel- atively uniform and of limited taxonomic value, differences being mainly of size. CYTOLOGY Fo » ier. uniform cytologically. number is x = 13 (Gwynne, 1958: ea gem 1), this coming from counts for 11 species. Additional studies here, including original counts for 32 species (Table 2), of which 30 are first reports for species, confirm the base number established in 1971. The cytological technique employed involved root tip squashes in contrast to the paraffin section method which I used in earlier studies of the genus. Details of the technique are described elsewhere (Gold- blatt, 1979b, 1980) and consist of a pretreatment hydrolysis in 10% HCl prior to squashing in or- cein based stains. A total of 41 species, over half the genus, is now known cytologically. Counts for most species, including as many as five or six popu- lations of some, are diploid, 2n = 26. Exceptions are G. bracteata (one count) and G. inaequalis (two counts), both tetraploid; G. aspera which has both diploid and tetraploid populations; and G. bolusii, where the five populations examined are triploid, 2n = 39. B chromosomes have been recorded in G. heterostyla, 2n = 26 and 2n = 26 + 2B, G. inflexa, G. bryicola, and G. tulbagh- ensis, 2n = 26 + 1B The basic number of x = 13 is somewhat un- usual in Iridaceae, but it is shared by the closely related Hesperantha (Goldblatt, 1971, 1984) and the mono- or di-typic Schizostylis (Goldblatt, 1971). The karyotype is similar in all three genera and consists of fairly small meta- to submeta- centric chromosomes that with the methods em- ployed, range in size from 2.5-4 um. GOLDBLATT— GEISSORHIZA 289 INFRAGENERIC CLASSIFICATION AND CLADISTICS The classification adopted here rests on the primary assumption that the peculiar imbricate corm tunics found in many species of Geisso- rhiza arose once in the evolution of the genus. The recognition of the two subgenera proposed here is thus based on the nature of the corm tunics, species of subgenus Geissorhiza having imbricate tunics and species of subgenus Wei- hea, the presumably ancestral concentric tunics. Several other characters are correlated with the presence of imbricate corm tunics, all apparently specialized and restricted, or nearly so, to sub- genus Geissorhiza, though not found in all species. These include leaf margin and midrib pubes- cence, corrugate leaves, ferrugineous bracts, un- equal stamens, and stem puberulence (or cilia- tion). Other major morphological specializations in Geissorhiza such as leaf margin and midrib thickening and development of floral zygomor- phy seem to have arisen independently, in both the basal subgenus Weihea and the derived sub- genus Geissorhiza. Both these types of special- ization occur in other genera of Iridaceae and in this light seem secondary to the evolution of im- bricate tunics, and in fact, both appear to have evolved more than twice in each subgenus. SECTIONAL CLASSIFICATION IN SUBGENUS WEIHEA Seven sections are here recognized in subgenus W. corm tunics and plane, usually ensiform to linear leaves without well-developed thickening of the margins or midrib. The flowers of the majority of species are also unspecialized, with tubes of moderate length, exserted and equal stamens. Geissorhiza ovata stands out in the section in havingan unusually long perianth tube and near- ly prostate, ovate leaves, while G. roseoaiba and ni ar fourcadei stands out here in its terete leaves an large flowers that are solitary on the branches. 290 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 TABLE 2. Chromosome numbers in Geissorhiza. Original counts are marked with an asterisk (*). Previous counts were reported by Goldblatt (1971) except as indicated in the table, by Gwynne (1958). Diploid Species Number Collection Data G. arenicola 26* W of Nieuwoudtville, Goldblatt 7056 (MO) G. aspera 26 ien locality or voucher (Gwynn, 1958—as G. secunda) 26* ondebosch Common, Goldblatt 5350 (MO); Burghers Pos, Darling, oes s.n. (no voucher); Olifants River Valley, near Rondegat, Goldblatt 5647 (MO) 52 Cape Peninsula, Signal Hill, Goldblatt 213 (BOL); Simonstown, Gold- blatt 520 (BOL) 52* Caledon, Albertyn road near Leeu River, Goldblatt 48394 (MO); St. James, Cape Peninsula, slopes below Boyes Drive, Goldblatt s.n. (no voucher G. barkerae 26* N of Piketberg, Goldblatt 6391 (MO) G. bolusii 39 Bains Kloof, Goldblatt 458 (BOL); Cape Peninsula, Grootkop, Gold- blatt 515 (BOL) 39* Villiersdorp, High Noon, Goldblatt s.n. (no voucher); Paarl, Simons- berg, Esterhuysen 35281A (MO); Jonaskop, Riviersonderend Mts., Goldblatt 6444 (MO) G. bonae-spei 26* Cape Peninsula, opposite Cape Point Reserve, Goldblatt 5267 (MO) G. bracteata ca. 52* Hankey, Klein River Valley, Goldblatt 4933 (MO) G. brevituba 26 Piketberg, top of Versveld Pass, Goldblatt 204 (BOL) (as Engyvsiphon) G. bryicola 26* Vogelgat, Hermanus, Goldblatt 6741 (MO) 26 + B* Vogelgat, Hermanus, Goldblatt 6742 (MO) G. burchellii 26* Swellendam, Langeberg, Esterhuysen 35363 (MO) G. cataractarum 26* Caledon, cliffs above Bettys Bay, Goldblatt 5355 (MO) G. cedarmontana 26* Cedarberg, Middelberg plateau, Goldblatt 5145 (MO) G. ciliatula 26* Cedarberg, Middelberg, Goldblatt 5144 (MO) G. confusa 26 Bains Kloof, Goldblatt 483 (BOL) (as Engysiphon exscapus) G. corrugata 26* Calvinia, Goldblatt 3895 (MO) G. erubescens 26* Pakhuis Pass, Goldblatt 6396 (MO) G. exscapa 26* top of Vanrhyns Pass, Goldblatt 7061 (MO) G. grandiflora 26* Limietberg, Bains Kloof, Goldblatt 6828 (MO) G. heterostyla 26 Voelvlei, Sutherland, Hall s.n. (NBG); Calvinia, Goldblatt 135 (J) (as G. leipoldtit) 26* Blomfontein road, W of Middelpos, Goldblatt 5815 (MO); Grasberg NW of Nieuwoudtville, Goldblatt 6265 (MO) 26 + 2B* Humansdorp, E of town, Goldblatt 5211 (MO) G. cf. hispidula 26* near Baardscheerdersbos, sandy flats, Goldblatt 5377 (MO) G. humilis 26* Cape Point, near Reserve, Goldblatt 5265 (MO) G. imbricata subsp. 26 without locality, Goldblatt 165 (J) imbricata G. inaequalis 52 Vanrhyns Pass, Goldblatt 129 (J); near Nieuwoudtville, Goldblatt 275 (BOL) G. inconspicua 26* near George, Vlok s.n. (no voucher) G. inflexa 26* Caledon, Queen Anne-Eseljagt, Goldblatt 2497 (MO); Caledon, W end of the town, Goldblatt 6173 (MO) 26 + 1B* Tulbagh cemetery, Goldblatt 5228 (MO) G. juncea 26* Cape Peninsula, Lions Head, Goldblatt 4700 (MO) G. kamiesmontana 26* Kamiesberg, Rooiberg slopes, Goldblatt 5770 (MO) G. karooica 26* Ghaap kop, Matjesfontein, Goldblatt 6371 (MO) 1985] TABLE 2. Continued. GOLDBLATT — GEISSORHIZA Pakhuis Pass, Goldblatt 122 (J) (as Engysiphon) N of Citrusdal, Goldblatt 6267 (MO) Citrusdal, Goldblatt 247 (BOL) Malmesbury Commonage, Goldblatt 6282 (MO) near Swellendam, Goldblatt s.n. (no voucher) Caledon, Goldblatt 214 (BOL) (as G. nana) Outeniqua Pass, George, Vlok 525 (MO) Bot River, Goldblatt 217 (BOL) Cape Peninsula, Signal Hill, Goldblatt 196 (BOL) Paardevlei Mts., Attaquaskloof, Viok 706 (MO) Elands Kloof, Goldblatt 5245 (MO) Bains Kloof, Goldblatt s.n. (no voucher); mts. above Clovelly, Cape ) Peninsula, Goldblatt 7052 (MO G. longifolia 26 26* G. louisabolusiae 26 G. malmesburiensis 26* G. nana 26* G. ornithogaloides 26 subsp. ornithoga- oides G. outeniquensis 26* G. ovata 26 G. pusilla 26 G. roseoalba 26* G. scillaris 26* G. similis 26* G. splendidissima 26 G. tabularis 26* G. tenella 26* G. tulbaghensis 26 + 1B* Nieuwoudtville, Goldblatt 347 (BOL) above Skeleton Gorge, Table Mt., Goldblatt 6726 (MO) S of Koeberg, Goldblatt 7111 (MO) Tulbagh cemetery, Goldblatt 5227 (MO) Section Tortuosa is a small segregate of section Weihea, restricted to the western Karoo. Two species have been added to the one assigned here by Foster. All three species are of small size, have large flowers, and are distinguished by having solitary flowered spikes. The monotypic section Pusilla includes only the distinctive and taxonomically isolated Cape Peninsula species, Geissorhiza pusilla. It has a combination of features that suggest strongly a ybrid origin. Its concentric corm tunics are typ- ical of subgenus Weihea, while the plane ensi- form leaves are unusual.for section Weihea in being pubescent. The very short-tubed flower and dry and ferrugineous floral bracts match exactly those found in sections Geissorhiza and Ciliata. This combination of the specialized features of subgenus Geissorhiza and the primitive concen- tric corm tunics of subgenus Weihea seems so discordant that an intersubgeneric hybrid origin is likely, although the possibility of an indepen- dent origin of the specialized short perianth tube and ferrugineous bracts cannot be discarded. Section Jncludanthera comprises only Geis- sorhiza cedarmontana and the poorly known G. esterhuyseniae, corms of which have not been seen. The two species are united in having the stamens and the style included in the lower part of the long perianth tube. The affinity of these two species is uncertain but they are assumed to be a monophyletic offshoot of section Weihea. Geissorhiza esterhuyseniae has the plane ensi- form leaves typical of section Weihea while the leaves of G. cedarmontana have strongly thick- ened margins and midribs. Section Includan- thera is restricted to the higher mountains of the western Cape. The specialized and derived sections Angus- tifolia and Ixiopsis are closely allied. Both have three or fewer, narrow leaves with heavily thick- ened margins and midribs and stems in which branching is suppressed. Section Angustifolia, corresponding largely to Foster's subsection Fili- formes, comprises some 11 species, sharing un- specialized, usually small, actinomorphic flowers and small corms with hard woody tunics. Species usually have two basal leaves and a third, which is much smaller, inserted in the mid part of the stem, and partly sheathing. Geissorhiza steno- siphon, a very local endemic of the Cold Bok- keveld Mountains is remarkable in the section for its slender perianth tube some 4-5 cm long, and it stands out in the genus in having a style that divides between the mouth of the tube and the base of the anthers. This species, as well as G. juncea and G. furva, stands out in having terete, 4-grooved leaves. The poorly understood G. intermedia and G. unifolia are unique in sub- 292 genus Weihea in having lightly ciliate stems. The section is centered in the southwestern Cape, G. unifolia and G. juncea occurring as far north as the Cedarberg and G. hispidula ranging from the Cape Peninsula to the southern Cape near Al- inia. The related section [xiopsis comprises a close knit alliance of five species that are centered in the mountains of the southwestern Cape. Only one species, Geissorhiza hesperanthoides was known to Foster, who assigned it to his subgenus Ixiopsis together with three species subsequently removed to Gladiolus. All the species of the sec- umbrosa extends much outside this area, occur- ring as well on the Cape Peninsula and in the Cedarberg. In these species, the third leaf is typ- ically reduced and entirely sheathing, and the corm tunics are soft in texture and either mem- branous or fibrous, most unusual for the genus. Section Engysiphon, another close knit alli- ance, comprises seven species, united in having arge, conspicuously veined zygomorphic and declinate flowers, usually with a very long peri- anth tube and long herbaceous bracts, the inner much smaller than the outer. The leaves have thickened to winged margins and the corms are typically large and have very hard woody tunics. The section, treated by Lewis (1941) as a separate genus and including one more species, here as- signed to section Weihea, comprises two distinct assemblages, one in which the leaf has the thick- ened margins and midrib, also found in section Angustifolia, and the other in which the leaf has raised and winged margins that curve over the lightly thickened midrib. Despite the clear cut leaf differences, the two groups appear closely related and there seems no doubt that the section Caledon district north to Namaqualand. SECTIONAL CLASSIFICATION IN SUBGENUS GEISSORHIZA The species of subgenus Geissorhiza are united primarily by the possession of their characteristic imbricate corm tunics. These are remarkably uniform in the majority of species, especially in sections Planifolia, Ciliata, and Geissorhiza, being hard, dark in color, and regularly notched below. In section Monticola, there is a trend for the softening of the texture of the tunics, while in the ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 two species of section Intermedia the corm tunics stand out as being pale in color. Section /nter- media is the only one in subgenus Geissorhiza in which none of the species have unequal sta- mens, a character that is probably a part of the genotype in the other sections although it is not always expressed. Thus it is, for basically nega- tive reasons, that section Intermedia has been recognized, and it stands out as an isolated and residual group ofthe subgenus, the species united by their pale imbricate corm tunics and the fre- quent specialized feature of narrow leaves with strongly thickened margins and midribs Section Monticola comprises a range of species in which branching is normally well-developed, the stamens are distinctly unequal and the corm tunics are often soft in texture with an accom- panying tendency for the tunics to become dis- placed easily. It then often becomes difficult to be certain of the type of tunics in some species. However, well-preserved corms in a few speci- mens or the associated characters of the species is usually sufficient to determine the sectional position. Only two species of the section were own to Foster (1941) who assigned them to his otherwise monophyletic subsection Fili- formes. Section Monticola is centered in the moun- tains of the southwestern Cape, but it has a wide range, G. ciliatula occurring in the Cedarberg in the north and G. burchellii extending as far east as the Langeberg at Heidelberg. Sections Ciliata and Planifolia appear closely allied, and were united by Foster in his subsec- tion Pubescentes together with G. pusilla. They have in common, flowers with very short peri- anth tubes and bracts that become dry and fer- rugineous above, during or after flowering. Some species of both sections have flowers with un- equal stamens, and puberulous or ciliate stems, the sections is that the leaves of section Planifolia are flat, and usually smooth, while those of sec- tion Ciliata have the margins and midribs (and sometimes other veins also) raised and winged, and the edges of the wings ciliate. Two species of section Planifolia have zygomorphic and dec- linate flowers. Section Geissorhiza, which corresponds with Foster’s nomenclaturally illegitimate subsection Ventricosae, comprises a group of closely related species all having the leaves with strongly thick- ened margins and veins. In a few species, such 1985] as G. brehmii and G. louisabolusiae, the leaves are very narrow or terete and 4-grooved, and in these the characteristic ribbing is developed on the typically inflated sheath of the Boyer est leaf. Section G en in the flat plains of the western Cape, betwen Cape Town and Piketberg, but species p as far north as the Nieuwoudtville Escarpment. The formal taxonomic treatment, Ss descriptions and citation of types for the infra- outlined here, are presented in 1e systematic treatment. CLADISTICS A cladogram has been developed (Fig. 16) in which the subgenera and sections are grouped according to their shared derived characteristics (synapomorphies). The cladogram was con- structed manually according to concepts estab- lished by Hennig (1966) and adapted by several bo "we Bree: (Bremer, 1976; Humphries, 1981; , 1982; to mention a few examples). AS D in the preceding pages, the subgen- era and sections are believed to be natural (monophyletic) assemblages. They can usually be distinguished on the basis of one or more spe- cialized features, the polarity of which was de- termined by outgroup comparison and by gen- erally accepted trends in the family. The primitive section Weihea, as well as sec- gustifolia (subgenus W. distically. Section Weihea is a residual grouping of presumably old and unspecialized species while sections Angustifolia and Monticola are defined phenetically. Section Intermedia (subgenus Geis- sorhiza) is a weakly defined group of two possibly residual species that may be misplaced in the subgenus. The major taxonomic characters are listed in Table 3 with their primitive (plesio- morphic) states assigned 0 and derived (apo- morphic) states assigned 1 (or 2 or 3 if the char- acter concerned has more than one derived state). The reasons for regarding the states as primitive or advanced are, unless obvious, discussed at length in the section dealing with the morphology of Geissorhiza. It is perhaps unnecessary to point out that in making a cladogram of a group above the species level, only the presumed primitive state for the taxon concerned should be considered and not other states of the character that occur in that group. Thus section Engysiphon is the only one in which a zygomorphic flower can be used as a GOLDBLATT — GEISSORHIZA 293 Subgenus Geissorhiza Subgenus Weihea FicunE 16. Cladogram of the sections of Geisso- rhiza. Section Pusilla is believed to be of hybrid origin between species of section Weihea and section Ciliata. sectional character. In other sections, both zy- omorphic and actinomorphic flowers occur but the basic state for each of these sections is pre- sumed to be the actinomorphic flower. One arbitrary assumption has been made in the case of the derived state for character 10, filaments unequal. This is, that whether the char- acter occurs in all the species of the section or not, the derived state is regarded as a sectional character. I have done this because the character is an unusual one and it seems reasonable to regard it, in the absence of contradictory evi- dence, as part of the genotype of those sections in which it occurs although not always expressed. The alternative, to treat the character as newly evolved in each section in which it occurs, seems absurd since it would then be necessary to as- sume the independent origin of this very unusual character at least six times in Pii dim Geisso- rhiza and nowhere else in Iridace My treatment of one main ich iof the clado- gram as a subgenus and several equivalent branches from the same node as sections of the second subgenus Weihea violates rigid cladistic theory. However, this seems justified in a situ- ation where a basal and unspecialized group, sec- tion Weihea (species of which are probably little different from the basal stock of the genus) has given rise apparently independently to several lines, one of which has diverged and radiated to 294 TABLE 3. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Important characters in Geissorhiza and their primitive and advanced states. The numbers 0 for primitive state, and 1, 2, or 3 for the derived state are used in the cladogram (Fig. 16) to indicate which characters have changed at branch points and their states. Primitive State Character Number 0 Advanced State 1. Corm tunics concentric 2. Tunic texture woody, brittle 3. Stem ooth 4. Leaf blade flat, ensiform 5. Inflorescence spike of several flowers 6. Bracts green and herbaceous 7. Flower actinomorphic 8. Perianth tube as long as bracts 9. Stamens and style well-exserted 10. Filaments equal 11. Stem branched imbricate membranous, soft/fibrous puberulent to pubescent . linear with thickened margins and midrib . linear with winged and ciliate margins and N — — — — midrib 3. several ribbed and grooved 1. solitary flowere . very long and outer more or less two times the inner . ferrugineous above when plants dried zygomorphic and declinate about half as along as bracts or . very long and well-exserted (unless the bracts also very long . included in lower part of the tube . one shorter than other two 1. branching suppressed Ne — N such a degree that it is best to assign it higher rank than its sister groups rather than follow strict cladistic theory in classifying taxa. The possible intersubgeneric hybrid origin of section Pusilla discussed above is not reflected in the cladogram. Instead, section Pusilla is placed in subgenus Weihea (concentric corm tunics), and the fea- tures shared with sections Geissorhiza and Cil- iata emerge as parallelisms. I have not attempted to develop cladograms for the species in any infrageneric group as there seems little value in such effort in a large genus such as Geissorhiza in which there are few sig- nificant characters that define species and rela- tionships appear reticulate in many sections due to convergence in many separate characters (ho- moplasy). In addition, some species that seem distinct on overall morphology appear to have no distinguishing features that can readily be po- larized and used in a cladistic analysis. GEOGRAPHY OF GEISSORHIZA GENERAL TRENDS IN DISTRIBUTION Geissorhiza is restricted to the southern Afri- can winter rainfall region (as defined by Gold- blatt, 1976b, 1979a, 1981) which extends along the west coast from southern Namibia to the Cape Peninsula, and along the south coast to Port Elizabeth (Fig. 17). Only one species, G. bract- eata occurs significantly outside this area. It ranges from the George district in the southern Cape into the southeastern Karoo and Eastern ape, where some winter precipitation is usual but the area is usually regarded having a summer rainfall pattern. The genus is largely restricted to the Cape Floristic Province, which lies entirely within the winter rainfall region (Goldblatt, 1978). Only a handful of species occur outside the limits of the Cape Flora, two in the north in Nama- qualand and four in the western and winter rain- fall part of the Karoo, as well as G. bracteata, mentioned above, Thed atterns of the two subgenera, Weihea and c differ in several ways. The more primitive subgenus Weihea is well rep- resented in the southern Cape (Fig. 18), the east- ern half of which receives substantial summer precipitation, and is often regarded as a region of year round rainfall. It is here that the majority of the least specialized species occur, most of which have comparatively wide ranges but there are amongst them several local endemics. Three species, comprising section Tortuosa, are unusu- al in this subgenus in being restricted to the west- ern Karoo, where Geissorhiza is otherwise rep- resented only by G. heterostyla (subgenus Geissorhiza). Subgenus Weihea has radiated ex- 1985] T LA ENS SEN me SNS AA =X Z| =\= ISSN == — NA h AA WSS “Iç A Mea. SN "It FiGURE 17. Concentration of species of Geissorhiza in southern Africa. Numbers indicate: the total number of The entire range of the genus is within the | heavy li The area usually regarded as receiving winter rainfall is to the west and south of the broken line. tensively in the southwestern Cape, where two sections, Angustifolia and Ixiopsis have almost all their species. The relatively few species of subgenus Weihea that extend further north are amongst the most specialized in the alliance. These include the two species of the peculiar section Zncludanthera, and representatives of sections Engysiphon (G. longifolia, G. confusa, G. exscapa) and Angustifolia (G. unifolia, G. jun- ea). Subgenus Geissorhiza is, in contrast, poorly represented in the southern Cape. It has its center in the southwestern Cape and is well represented northwards through the Cedarberg and Nieu- woudtville Plateau, while two species are endem- ic in Namaqualand. Despite the different overall distribution patterns in the two subgenera, both exhibit similar modes of extensive radiation in the southwestern Cape, both in the mountains and along the coastal belt from Cape Town to Piketberg. Subgenus Geissorhiza has radiated to a greater degree in the mountains of the north- west Cape and has two important centers of endemism here, one in the Cedarberg and one on the Nieuwoudtville Plateau and the Gifberg. In all, 76 species of Geissorhiza occur in the Cape Floristic Region in a strict sense, and of these 73 occur exclusively here. Those occurring in the Cape Region as well as in the Karoo and Namaqualand are: G. bracteata (George in the southern Cape to Grahamstown and Cradock); GOLDBLATT— GEISSORHIZA 4 A 2057 p ^ fi n Ale r e ca 0% Ad ES >> ze MAS Stitt medi? E PE VIN q S3 IZN N DIE DA dia P7 S: audi * iret whereas subgenus Geissorhiza has radiated extensively along the west coast, with species extending into north- ern Namaqualand. i heterostyla (southern Cape, the western Ka- oo, Nieuwoudtville escarpment and southern Namaqualand); and G. exscapa (Malmesbury to o ipbaai in Namaqualand). Geissorhiza namaquensis and G. kamiesmontana are re- stricted to Namaqualand and G. karooica, G. corrugata, and G. spiralis are local endemics in the western Karoo. The high concentration of species in the west (Fig. 17) suggests that the substantial radiation in Geissorhiza in this semi-arid, and summer area of rugged and varied landscape has been the result of strong selective pressure through ex- treme climatic variation during the glacial pe- riods of the later Pleistocene (see chapter on Evo- lution). Contractions of ranges of species and the reduction and fragmentation of populations dur- ing periods of extremely unfavorable conditions presumably enhanced the tendency for diver- gence of species by the random effects of genetic drift in conjunction with locally differing selec- tive forces. It is not surprising that concentration falls sharply in Namaqualand and in the interi where the climate is most arid. The PRAE equable climate and higher, more evenly spread rainfall in the eastern southern Cape has evi- dently not provided the same stimulus to spe- ciation that is so developed in the west, and this 296 TABLE 4. Centers of endemism in Geissorhiza. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 SOUTHWEST CAPE CENTER Hottentots Holland-French Hoek Subcenter Malmesbury Flats Subcenter cens, Cape Peninsula Subcenter Bredasdorp Subcenter Species occurring in more than one Subcenter of the Southwestern Center NORTHWEST CAPE CENTER Cedarberg Subcenter (here includ- ing the Piketberg) cens, G. leip G. unifolia Great Winterhoek Subcenter Gifberg-Bokkeveld Mts. Subcenter G. barkerae, G. brevit G. alticola, G. lithicola, G. nubigena G. darlingensis, G. eurystigma, G. furva, G. mathewsii, G. purpuras- G. radians G. bonae-spei, G. tabularis G. bryicola, G. schinzii G. cataractarum, G. hesperanthoides, G. hispidula, G. humilis, G. pusilla, G. setacea, G. tenuis uba, G. cedarmontana, G. ciliatula, G. erubes- oldtii, G. louisabolusiae, G. minuta, G. stenosiphon, G. esterhuyseniae, G. scopulosa G. arenicola, G. divaricata, G. inaequalis, G. splendidissima, G. sub- rigida, G. sulphurascens Species occurring in more than one G. longifolia Subcenter of the Northwestern Center LANGEBERG CENTER G. foliosa KAROO MT. CENTER KNYSNA REGION SOUTHEAST CAPE CENTER none G. nigromontana G. elsiae, G. outeniquensis area may be seen as a refugium for the primitive species of the genus. CENTERS OF ENDEMISM Geissorhiza is clearly a genus of the Cape Flo- ristic Region and a predominantly montane one. It exhibits the patterns of endemism that have come to be seen as characteristic of the Cape Region (Weimarck, 1941) and the following dis- cussion is framed in the context of Weimarck's phytogeographical subdivision of the Cape Flo- ristic Region. Weimarck recognized six main centers of endemism and these are dealt with in sequence. The Southwestern Center. The area of high- plain north of the Cape Peninsula (Weimarck's Malmesbury Flats Subcenter) and on the moun- tain complex that includes the Kogelberg, Hot- tentots Holland, French Hoek, Du Toits Kloof, and Bains Kloof Mountain Ranges (the French Hoek and Hottentots Holland Subcenters). Six species are restricted to the Malmesbury Flats (Table 4) and two to the Hottentots Holland- French Hoek Subcenters. The Cape Peninsula Subcenter is small but has two endemic montane to submontane species (Table 4). These presum- ably evolved as a result of their separation from the main populations of their relatives that grow inland across the inhospitable Cape Flats barrier that isolates this subcenter. The Bredasdorp Sub- center, so frequently an important center of ra- diation for taxa of the Cape Flora, has only two endemics (G. schinzii, G. bryicola), also both montane to submontane. Most of the mountain species of this subcenter also occur in the Hot- tentots Holland Subcenter, the mountain ranges being continuous or almost so. The lowland species are either shared with the Langeberg Cen- ter (G. nana) (there is virtually no barrier to mi- gration between the lowlands of the two regions) or the Malmesbury Flats (G. "aN (isolated by substantial mountain barrier The Northwestern Center. is includes two important subcenters, the Great Winterhoek 1985] Subcenter, and the Cedarberg Subcenter (Table 4), while Weimarck admits two noncontiguous areas, the Kamiesberg and Hantam-Roggeveld t Winterhoek Subcenter and ten to the Cedarberg Subcenter. This area includes several of the most specialized species in the genus including G. stenosiphon, the two species of section /nclu- danthera, G. esterhuyseniae and G. cedarmon- tana. Evidently the fairly rich development of the Cape flora on Cape Sandstone soils north of the Cedarberg, on the Gifberg and Bokkeveld Mountain complex was unknown to Weimarck, and I feel compelled to extend the Northwestern Center to include this area, which I recognize as the Gifberg-Bokkeveld Subcenter. The signifi- cance of this subcenter is particularly evident in Iridaceae, not only in Geissorhiza but in several other genera, notably Hesperantha, Gladiolus, and Sparaxis as well as in other monocots. The Geissorhiza endemics in this subcenter include G. subrigida, G. divaricata, G. arenicola, and G splendidissima, all closely allied species of sec- tion Ciliata and G. sulphurascens, section Geis- sorhiza. Neither the Kamiesberg nor the Hantam-Rog- geveld Subcenter is regarded today as part of the Cape Flora, but there is a development of vege- tation that has several elements clearly of Cape origin. Two species of Geissorhiza, G. tortilis and G. corrugata are endemic in the Hantam-Rog- geveld area while G. heterostyla is common but it extends through the interior southwestern Cape and the southern Cape to Port Elizabeth. On the Kamiesberg, there are only two species, the en- demic G. kamiesmontana and G. namaquensis, that extend northward to Namaqualand to the southern Richtersveld. Neither species is com- mon, and they seem to be restricted to areas of higher moisture availability in this arid country. The Southern Cape Centers. Few species of Geissorhiza in the southern Cape fall convenient- ly into the four phytogeographic divisions rec- a Karoo Mountain, berg Center, G. foliosa, and two to the Knysna Region, G. outeniquensis and G. elsiae. No species is endemic to the Eastern Center and its sepa- ration from the Knysna Region has no signifi- cance for Geissorhiza or for Iridaceae in general. In the combined southern Cape Centers, the en- GOLDBLATT — GEISSORHIZA 297 demics include G. fourcadei, G. roseoalba, and G. inconspicua, in addition to the more restricted G. foliosa, G. outeniquensis, G. nigromontana, and G. elsiae. This is not surprising since the Langeberg and Outeniqua Mountains extend vir- tually weer from Montagu in the west to Humansdorp in the east, and the few minor gaps seem not to have affected the distribution of Geissorhiza species to any degree. The drier in- terior southern Cape mountains that include the small and somewhat isolated Little Karoo ranges (Rooiberg, Warmwaterberg, Kammanassie Mountains) and the Swartberg Mountains have a poor Geissorhiza flora in contrast to the ap- parently climatically and geologically similar western Cape mountains. Reasons for this are not clear but presumably they had very different climatic histories. In summary, the southwestern part of the Cape Floristic Region is the center for the evolution and radiation of Geissorhiza. Within the region are several important centers of endemism. The most important of these are the coastal plain between Cape Town and Piketberg (Weimarck’s Hangklip and Tulbagh (W and Hottentots Holland Subcenters). The moun- tains of the west coast, the Northwestern Center, constitute another important center for specia- tion especially for subgenus Geissorhiza; and a Gifberg-Bokkeveld Mountain Subcenter, not recognized by Weimarck, emerges as a major center of endemism in Geissorhiza as well as many other geophytic monocots. ECOLOGY Species of Geissorhiza have radiated exten- sively in the southwestern Cape and have come to occupy a greater variety of niches than most other genera of Iridaceae, here or elsewhere. Per- haps the most important ecological distinction in the genus is an edaphic one. Species are gen- erally found on either a shale-derived clay soil or a sandstone or granite-derived sandy soil. The importance of soil type in the distribution of Cape taxa (Goldblatt, 1978) has been discussed by several authors for various genera, notably Dahlgren (1968) for Aspalathus (Leguminosae) and Rourke (1972) for Leucadendron (Protea- ceae) and it seems particularly significant in geo- phytic taxa (Goldblatt, 1976b, 1979a, 1981). In Geissorhiza, which is almost entirely re- stricted to ake area of the Cape Flora, relatively 298 few species grow on the heavy clay soils that are covered by renosterbosveld, a characteristic mi- This habitat is with numerous geophytic taxa of several families. These soils are found mainly on flats or lower mountain slopes. They are relatively well watered, especially along the coast, in the wet winter months, but dry out rapidly in the later spring when much of the geophyte flora of these areas blooms in a four to six week period in the spring. Common species of this habitat are G. inflexa, G. leipoldtii, the rare and rather local G. tulbaghensis (section Planifolia), G. nana and G. bracteata (section Weihea), and G. ornithoga- loides, to mention some of the more significant examples. Geissorhiza erubescens (section Cil- iata) and G. longifolia (section Engysiphon) are montane to submontane species, but they occur on the upper shale band of the Table Mountain Series. This is a broad layer of shale located in the upper part of an otherwise sandstone geo- logical system, of which the Cape mountains are largely composed e very characteristic nutrient poor quartz- itic sandy soils of the Cape region have a quite different range of Geissorhiza species, and it is on these soils that most of the radiation in the genus has occurred. The lowland alluvial sands have a very rich Geissorhiza flora. These soils also dry out fairly rapidly at the end of the wet months and flowering is condensed into an eight week period, but lasting longer than in the clay- loving species. Most common on flats and lower mountain slopes in sandy soil is G. aspera, of the southwestern and western Cape, while G. het- erostyla is found in similar habitats in the south- ern Cape. Curiously, outside the Cape Region, G. heterostyla occurs on heavy clay soils in the Roggeveld, which seems a strange contradiction. Species of section Angustifolia are also well suit- ed to lowland alluvial sands, and G. humilis and G. hispidula are common here in the western Cape, while the related, very local endemic G. darlingensis occurs near Darling in similar sites. In seasonally moist sites in the southwestern Cape, section Geissorhiza has speciated to an unusual extent, and all the species of section Geis- sorhiza occur in such habitats. Geissorhiza pur- pureolutea, G. imbricata, and G. barkerae occur in moist sandy flats between Cape Town and Piketberg, while G. brehmii favors even wetter situations in this area and is semi-aquatic, grow- ing in seasonal pools. In the Darling district where the parent rock is granite, G. radians, G. ma- thewsii, and G. eurystigma, also of section Geis- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 sorhiza, fill this seasonally wet lowland habitat. Further north this wet sandy habitat is occupied by G. louisabolusiae in the Olifants River Valley and by the closely allied G. sulphurascens on the Nieuwoudtville escarpment. Only a few species in other sections grow in this habitat, notably G. setacea wakes Weihea) which occurs between Cape Town and Gouda. The d Ld montane species all grow in well-drained, nutrient poor soil derived from rocks of the Cape Sandstone series. Although these soils are very low in nutrients and have poor water retaining properties, the mountains themselves receive a higher rainfall than the sur- rounding valleys and flats, and this appears to have allowed geophytic groups such as Geisso- rhiza to have filled some unusual niches. In mon- tane marshy sites, G. umbrosa is often common in the early summer when the surrounding vege- tation has been burned. Damp southerly facing slopes often harbor G. hesperanthoides, G. nu- bigena, and open rocky situations are a habitat for G. ramosa, G. grandiflora, and G. burchellii. The flowering season is later and longer in the higher mountains, presumably because of cooler temperatures an ecause moisture may be available for a longer time, or may even be per- manent, as for the January flowering G. cata- ractarum and G. outeniquensis, both species of stream and waterfall edges. Species of section Engysiphon often occur on dry mountain slopes and flower in the early sum- stricted to the isolated Piketberg Mountains and G. bonae-spei to the southern Cape Peninsula. POLLINATION The pollination biology of Geissorhiza is large- ly unknown, but the genus is believed to be main- ly bee pollinated (Vogel, 1954: 101, 115). Vogel suggested this on the basis of the predominant flower form in the genus: the flowers are radially symmetric, typically short-tubed, open during the support his hypothesis, including those of Elliot (1891), who made numerous records of insect visitors on southern African plants. Elliot found that Geissorhiza aspera (as G. secunda) was vis- ited by several species of bees. His record of a dipteran visiting ‘Ixia excisa Thunb.’ is also probably for Geissorhiza, I. excisa being an early synonym for the distinctive G. ovata. Vogel re- 1985] cords the observations of Hans Herre of bee pol- lination for G. radians (as G. rochensis) and pre- sumably his own for a few more species: G. splendidissima, G. furva, and a species identified as G. ramosa, all from ‘Little Namaqualand,’ probably the northwestern Cape. Vogel suggests that the pale, small flowers of G. juncea are moth pollinated, on the assump- tion that they open in the evening. The species is, however, day blooming and very short-tubed therefore, moth pollination is unlikely. If direct bservation was actually involved, then a mis- identification must have been made. Vogel also suggests that the few long-tubed species of Geis- sorhiza may be butterfly pollinated, mentioning by name G. fourcadei. On the basis of their flower form, he predicts that G. ovata and ‘G. nama quensis, perhaps G. kamiesmontana, also have this type of pollination. Elliot's observation of a dipteran visiting G. ovata would seem to con- tradict butterfly pollination for this species. Cl more data is required before the some- what confusing and uncritical suggestions made by Vogel can be accepted. It seems likely that pollination studies in Geissorhiza will be of con- siderable value in understanding the evolution and speciation in this large genus in which flower form is often the major or only distinction be- tween related species. CONSERVATION There are a large number of very localized species in Geissorhiza and by certain definitions all of these may be regarded as potentially en- dangered. However, all of those that occur in mountain habitats appear reasonably secure be- cause this habitat is not likely to be modified to any appreciable degree in the foreseeable future except by too frequent fires and the spread of alien woody vegetation that poses a severe threat to some of the Cape mountain vegetation (Gold- blatt, 1978; Stirton, 1978). As far as I am aware, ment conducted by the South African Depart- ment of Forestry, which controls a large part of the Cape mountains, is in fact likely to improve the conservation status of what remains of this vegetation. The situation is, however, very different in the lowlands, particularly on the coastal plain north of Cape Town as far as Piketberg. Here the spread of agriculture and the growing population are placing considerable pressure on what little re- GOLDBLATT — GEISSORHIZA 299 mains of the rich and highly endemic flora of the area. This coastal plain is one of the two major centers of radiation of Geissorhiza and of the approximately 35 species that occur here, 15 have the larger part of their range in this relatively small area, and six species, G. purpurascens, G. darlingensis, G. eurystigma, G. mathewsii, G. urva G. radians are endemic. A further two nearly restricted to this area and are sim threatened. Of the species mentioned above name, only G. radians (better known as G. ro- chensis) has a wide distribution throughout the coastal plain. Although now reduced to fewer and smaller populations, it is still relatively common. The exact status of the other species mentioned is uncertain but all must be regarded as severely threatened in some degree. Geissorhiza darling- ensis and G. mathewsii, are both restricted to few sights and require protection. Both occur in the Darling area, northwest of Cape Town in a rich agricultural region, and the changes in agricul- tural practice in recent years including aerial spraying of weed killers and insecticides, as well as the enrichment of the soil through annual ap- plication of fertilizers, have changed the habitat of what is left of the native flora so that it seems probable that some of the rarer coastal plain en- demics may become extinct within the next few decades th h habitat loss or impoverishment. As demonstrated by Parker (1982), there is very little undisturbed habitat along the west coast and the need for preserves to save what remains of this once rich and interesting flora should be given the highest priority. To the north, the situation is similar in lowland areas and Geissorhiza louisabolusiae, endemic to the mid Olifants River Valley and adjacent val- leys, is near extinction as a result of the spread of agriculture in this area and the draining of the semi-marshy situations to which it is restricted. The several local endemics of the Nieuwoudt- ville escarpment have always been very local in range and should be considered under pressure. Flat sandy areas here are increasingly being put to agricultural use and the striking G. splendi- dissima, to name one species, is becoming par- ticularly rare and should be regarded as seriously endangered. EVOLUTION It seems likely that the ancestral stock that gave rise to Geissorhiza and the closely allied Hesperantha differentiated in upland eastern 300 southern Africa in the Miocene (26-5 Ma) when the climate of this area was becoming increas- ingly seasonal and the predominant forest vege- tation was giving place to larger areas of open grassland and heathland (Axelrod & Raven, 1978; Goldblatt, 1983). A seasonal climate presumably provided the selective force for the deciduous and geophytic habit. During the Miocene, Africa moved rapidly some 10? northwards and at this time the southern and western Cape must have been well forested perhaps with local areas of drier climate suitable for geophytes. Conditions along the wetter parts of the south- ern Cape coast today are probably similar with forest clothing the valleys and the southern mountain slopes but locally either rock outcrops, shallow soils or rainshadow effects provide suit- able habitats for species of Geissorhiza and many other paries taxa. It was not until the late Pliocene or the Pleistocene, however, that a Med- Vend winter wet and summer dry, climate developed along the Cape south western and west coasts and truly arid conditions prevailed to the interior and further north along the west coast. Thus the large numbers of species of Geissorhiza that occur today in the southwestern Cape and seem so specialized in comparison to most of the species of the southern Cape, probably did not exist until relatively recent times. Many probably evolved only during the Pleistocene as the dis- tinctive Cape climate became established. The climatic fluctuations of the Pleistocene, reason- ably well documented along the Cape west coast at Rietvlei and Langebaan, are believed to have provided the stimulus for the burst of radiation that produced the wide array of species of Geis- sorhiza in the southwestern Cape The unspecialized southern Cape species such as Geissorhiza inconspicua, G. bracteata, and G. foliosa have probably diverged very little from their ancestors that were present in the late Mio- cene and the Pliocene. They probably evolved slowly by a process of phyletic evolution (G. G. Simpson, 1953; B. B. Simpson, 1973) rather than true speciation. These species or their ancestors probably migrated from the forests of south- eastern Africa into the eastern and southern Cape when habitats opened up for them as aridity spread south ward in the Pliocene and Pleistocene in Pleistocene and the recorded climatic fluctua- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 tions of the time are known to have led to major shifts in the vegetation belts here, with the forest alternately contracting to sheltered sites and then expanding into the lowlands where they do not occur today. These shifts must have affected the ranges and population sizes of species of Geis- sorhiza very strongly, with the resulting local ex- tinctions and fragmentation of populations that are believed to precede geographic speciation. Increasing aridity also accentuated habitat dif- ferences and increased the diversity of ecological niches available to species. The large numbers of species that occur along the west coast and in the adjacent mountains are thus seen as having evolved under circumstances of increasing but fluctuating aridity, more or less in place in a relatively short time, following the classical model for geographic and eats speciation or a combination of these. The ma trends of phenotypic specialization are xero- morphic. They include a reduction in the number of leaves, decrease in leaf surface area, the de- velopment of heavy marginal and costal thick- enings alternating with grooves, and leaf edge ciliation. Stem modifications include the devel- opment of pubescence, the reduction of branch- ing and of flower number in the spike. Similarly, floral modifications such as the decrease or in- crease in the tube length are associated with spe- cialization for different pollinators, a feature not evident among the primitive, less drought adapt- ed species. Such floral specializations seem con- sistent with adaptation to drier climates, where bees and other insects with specialized pollina- tion behavior are more numerous. Subsequent migration and the spread of newly evolved forms must certainly have occurred but time and physical barriers have limited their spread throughout suitable areas of the south- western Cape, and these factors have combined to produce the patterns of distribution that pre- vail today. The centers of endemism and geog- raphy of the genus are discussed more fully in the section on Geography. Section J/ncludanthera affords a striking ex- ample of quantum evolution or speciation (G. G. Simpson, 1953; Grant, 1977), usually regard- ed as distinct from geographic speciation. In quantum speciation, novel phenotypes appear to have been produced abruptly, without a series of intermediates among their allies. The two species of section /ncludanthera are remarkable in their long-tubed flower with the stamens in- 1985] cluded in the middle part of the tube and the style and style | and no more than a few millimeters long. The species with this remarkable flower might seem better treated as a separate genus but examples of similar specialization occur in a few isolated species of Hesperantha (Goldblatt, 1984) and in some forms of Ixia paniculata (Lewis, 1962). The examples of convergence for included sta- mens and styles makes it seem best to maintain species with this character combination in the genera to which they are allied. Speciation through gross chromosome muta- tion is apparently rare in Geissorhiza. Examples of polyploid speciation include G. inaequalis and possibly G. bracteata, both tetraploid, but the single count for the latter is not sufficient for this hypothesis to be accepted without reservation. In G. aspera where tetraploid populations were initially found, further examination has revealed the presence of diploid populations as well, and the possibility of its polyploid origin is no longer tenable. Geissorhiza bolusti, however, seems to have arisen throu gh JF arn 11 + triploid 2n = 39. The ancestors of this widespread, sexually sterile species have not been identified, but the morphologically similar and sexually fertile G. ovalifolia (chromosome number unknown) is very likely one of the progenitors of G. bolusii. With about half the species of Geissorhiza known cytologically, it is not possible to fully assess the role of chromosomal mutation in spe- ciation, but it seems likely to have been small as in most other genera of Ixioideae in Africa (Goldblatt, 1971), with the notable exception of Romulea (De Vos, 1972). TAXONOMIC HISTORY The first species of Geissorhiza known to sci- ence were described during the later eighteenth century, and assigned to the broadly circum- scribed genus Ixia. When Geissorhiza was first regarded as distinct from Ixia (Ker, 1803, 1804) some 15 different species had been described, three by de la Roche (1766), one each by Bergius (1767), Burman f. (1768),and Linnaeus f. (1782), eight by Thunberg (1783, 1800, 1803) and two by Ker. Ker actually admitted only 11 species to Geissorhiza (one of which is Romulea sp. and another a nomen nudum but it is probably the species currently known as G. furva Ker ex Bak- er). He was apparently unaware of Burman's G. GOLDBLATT— GEISSORHIZA 301 ovata, of which G. excisa (L. f.) Ker is a synonym, and of three Thunberg species, G. exscapus pub- lished in 1800 (as Gladiolus) and G. monanthos and G. radians published in early 1803. Ofthese early botanists, the contribution made by Thunberg is by far the most important (Gold- blatt, 1983). Thunberg actually collected speci- mens of at least 13 species during the three years he spent exploring the Cape (1772-1775). He regarded these as belonging to eight different species, seven assigned to Ixia and one, G. exsca- pus to Gladiolus. Thunberg treated all but two, I. secunda sensu Berg. and I. hirta Thunb. (nom. superfl. pro J. inflexa de la Roche) as new. The identity and typification of the species described by Thunberg is complex and although the Iri- daceae of Thunberg's Herbarium were examined and reviewed by Brown in 1928, several prob- lems were only resolved after a recent reexami- nation of the collection (Goldblatt, 1983). The typification of the three species of Ixia described by Daniel de la Roche and now known to belong in Geissorhiza is also problematic (Goldblatt & Barnard, 1970). Authentic material was discov- ered in 1969 and as a result several de la Roche species could for the first time be identified and typified, including the long misinterpreted G. (Ixia) inflexa. No sme material je G. (Ixia) imbricata was discovered and a neotype now identifies this sii (Goldblatt & Barnard, 1970). The name of a third species, 7. secunda (syn. G. eurystigma) cannot be transferred to Geissorhiza as the epithet is occupied in the ge- nus. The important cycle of collecting in the period 1820-1830, by Ecklon and Zeyher, by Drége, and to a lesser extent by Pappe, resulted in the dis- covery of many more new species of Geissorhiza, collected in large sets and widely distributed among European, North American, and South African herbaria. Ecklon (1827) enumerated his and Zeyher’s collections, several of these being new species, but mostly as nomina nuda. Ecklon also introduced the new generic name Weihea ) for the unusual G. ova- ta (as W. excisa) as s well as G. bracteata (as W. elatior). Weihea was validated as a subgenus by Foster (1941) and is recognized here with a much broader circumscription. Amongst Ecklon's 18 listed species of Geissorhiza and Weihea, 13 are apparently new, but only two, G. rosea and G. candida can be regarded as legitimate. Most of the Drége, and Ecklon and Zeyher collections were (also without 302 studied later in the nineteenth century by Klatt (1866, 1882) and by Baker (1876, 1892, 1896), both of whom described several new species from their collections. aker’s treatments of Iridaceae in his **Hand- book of the Irideae" (1892) and in “Flora Ca- pensis" (1896), essentially identical, are a wa- tershed for the taxonomy of the family in southern rica. Baker recognized 29 species in Geisso- rhiza (six of which have since been transferred to Hesperantha, Ixia, and Gladiolus) and three more were assigned either to Acidanthera (G. exscapa as A. tubulosa; and G. tenella as A. ro- sea) or Romulea (G. spiralis). Klatt's (1895) treatment in the “Conspectus Florae Africae" is seldom given much attention by R uen but it is noteworthy that he recognized 40 speci in Geissorhiza. Ofthese some five belong to se genera, leaving the genus still appreciably larger than Baker treated it a year later. Botantical activity in southern Africa acceler- ated in the late nineteenth century, when im- portant d of Geissorhiza were made b Peter MacOwan, Harry Bolus, Arnold Penther, and notably Rudolph Schlechter. Baker de- scribed three species based on Penther collec- tions that he saw after the “Flora Capensis” had been published. Schlechter's collections were de- scribed partly by Schlechter himself (1899), but only one of his species, G. hesperanthoides, is currently recognized. Later, Foster (1941) de- scribed several new species based on the Schlech- ter collections, of which G. burchellii, G. mal- mesburiensis, and G. leipoldtii are currently recognized. The modern cycle of collecting, initiated by H. M. L. (Louisa) Bolus in the 1920s continued es- sentially without a break until today. Louisa Bo- lus and her students, W. F. Barker and G. J. Lewis,as wellas R. H. Compton made important collections of Iridaceae generally. Their activity resulted in the discovery of several new species of Geissorhiza (Bolus, 1930, 1931; Bolus & Lew- is, 1933)and extended the known ranges of many more. Foster's studies, published at this time, however, benefited little from this work, and he saw only the few duplicates that then reached European herbaria. Nevertheless, Foster (1936, 1941) clarified the nomenclature of several pre- viously problematic species, and he described as new some 11 species: G. burchellii, G. louisa- bolusiae, G. sulphurascens, G. leipoldtii, G. ova- lifolia, G. malmesburiensis, currently recognized and G. marlothii, G. rubicunda, G. ixioides, G. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 lewisiae, G. montana, here reduced to synony- my. During this time the artificially constituted ge- nus Acidanthera was largely disassembled by Lewis (1941). Species were shown to belong to Gladiolus, Hesperantha, Tritonia, and Geisso- rhiza while a few, regarded by Lewis as allied to Geissorhiza, were assigned to a new genus En- gysiphon. The criteria she used for distinguishing this genus from Geissorhiza are: unilateral and declinate stamens and style; filaments exceeding anthers; bracts firm throughout; and a long flow- er, usually with a well-developed perianth tube. Critical analysis shows that Engysiphon has no significant characters or character combinations not found in Geissorhiza. The genus is reduced to sectional subgeneric rank here. The extensive botanical exploration of the higher Cape mountains, undertaken by Elsie Es- terhuysen, began in the 1940s. Esterhuysen’s Cape mountain collections, supplemented by those of Thomas Stokoe, Peter Jackson, and Ernest Gal- pin have brought to light a whole range of mon- tane species of Geissorhiza and expanded the ranges of several more. As a result, G. alticola, G. nubigena, G. esterhuyseniae, G. stenosiphon, G. scopulosa, G. unifolia, G. elsiae, and G. gran- diflora, previously unknown or poorly collected, were discoverd and are described here, while species such as G. burchellii and G. ramosa pre- viously known only from one or a few gatherings from isolated mountain localities are now known to have extensive ranges. The high mountain species are still relatively poorly known, and it seems likely that novelties will continue to be discovered in the mountains of the western and southern Cape. SYSTEMATIC TREATMENT Geissorhiza Ker, Ann. Bot. (Kónig & Sims) 1: 223. 1804; Baker, Handb. Irid. 152-159. 1802, Fl. Cap. 6: 65-76. 1896; Klatt, Abh. Naturf. Ges. Halle 15: 390-393 (Erg. 56- 61). 1882; Durand & Schinz, Consp. Fl. Af- ric. 5: 176-181. 1895; Foster, Contr. Gray Herb. 135: 3-78. 1941; Lewis, Fl. Cape Pen- ins. 252-256. 1950. TYPE: G. obtusata Ker [^ G. imbricata subsp. bicolor (Thunb.) Goldbl.] Ixia L. sensu L. f., Suppl. Pl. 91-94. 1782; sensu Thunb., Diss. de Ixia 1783, pro parte. Weihea Ecklon, Topogr. Verz. Pflanzensamml. Ecklon ud. Sphaerospora Klatt, Linnaea 32: 725. 1863, hom. illeg. 1985] non Sweet ex Loudon (1841). TYPE: S. exscapa (Thunb.) Klatt (lectotype here designated). Acidanthera Bm sensu Baker, Fl. Cap. 6: 130-134. 1896, pro part Buber! pien 1 S. African Bot. 7: 19-24. 1941. TYPE: E. schinzii (Baker) Lewis (lectotype here designated, type not indicated by Lewis). Plants small to medium in size. Rootstock a tunicate corm, with hard and woody to mem- branous, or rarely fibrous coverings, either asym- metric with one side flattened below and some- times extending downwards shortly or evidently symmetric, but usually with the roots developing from a small ridge on one side; tunics either con- centric (subgenus Weihea) with the outer layers usually light brown and completely enclosing the inner, fragmenting into irregular to elliptic seg- ments, often drawn into points above and below; or imbricate (subgenus Geissorhiza) with the out- er layers dark brown to blackish and covering the inner above only, regularly V-notched below into nearly rectangular segments. Leaves several to only 2 (rarely solitary), at least the lowermost basal and usually largest, decreasing in size above and the uppermost often becoming bract-like and entirely sheathing, erect to falcate or sometimes prostrate, linear, ensiform or linear; surface plane and smooth (rarely pubescent, G. pusilla), or with the margins and midrib raised and thus 2-grooved on each surface, sometimes (section Ciliata) the margins held at right angles to the blade and ciliate along the edges and the midrib (and oc- casionally other veins) raised and winged, with cilia along the edges, or lamina 3 to several ribbed and grooved (section Geissorhiza); margins smooth, ciliate to pubescent, or sticky and with adhering soil. Stem erect, terete, simple or branched and then either from the base or from cauline nodes. Spike usually several to many flowered, or sometimes only 1- or 2-flowered; always drooping in bud and becoming erect as the flowers open; bracts herbaceous to membra- nous, sometimes reddish on the upper margins, dry and brownish in the upper half (especially GOLDBLATT— GEISSORHIZA 303 sections Planifolia and Ciliata), the outer larger and usually slightly longer than the inner, the inner bifid at the apex. Flower actinomorphic and stellate 0 hypoctateriorm or Zygomorplue SRS See upri unilateral anthers and a declinate style, variously colored with shades of purple and pink, or deep violet to blue most common, but also red, white, or yellow, the tepals often darker colored on the reverse and frequently reddish when flowers a pale color; perianth tube straight, shorter or about as long as the bracts, less often much longer than the bracts, usually cylindric; tepals oblong to ob- ovate, the outer usually somewhat larger than the inner, spreading equally or with the posterior tepals upright and the anterior tepals horizontal and lying below the decumbent stamens and style. Filaments inserted near the top of the perianth tube, occasionally in lower part of the tube, erect and symmetrically disposed to unilateral and de- clinate, all equal or one consistently shorter than the other two (especially subgenus Geissorhiza); anthers oblong to linear, shortly tailed, subbas- ifixed, extrorse when uh pr ipn disposed, well- n section Incl ), pol- len yellow, blue, or violet. Ovary ovoid, style slender, initially erect, usually becoming dis- placed laterally or declinate and lying under the filaments, usually well-exserted and branching nearthe apex ofthe anthers, occasionally entirely included (section Includanthera), sometimes di- viding at the mouth of the tube or at the base of the anthers, style branches usually short, ascend- ing and ultimately recurved, occasionally flat and broadly lobed. Fruit a globose to obovoid cap- sule, usually about as long as the bracts; seeds globose to angular, i to dark brown. Basic ee number, x = 13. ty-one NRI Qm to the winter Noo areas of southern Africa, and concen- trated in the southwestern Cape; one species ex- tending into northern Namaqualand and another into the eastern Karoo and Eastern Cape as far east as Cradock and Grahamstown. 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(K, PRE); Buffelsrivier, SW of oe 2,900 ft. (DA), Oliver 3645 (K, P E). 3324 (Steytlerville) ee Mts., between 1985] GOLDBLATT — GEISSORHIZA 313 HD eA eA CQN d UY2 A URE 20. Morphology and distribution of Geissorhiza roseoalba. Habit x0.5; flower life size (Vlok 706, FIG Attaquaskloof). Smitskraal and Wilgehof, 3,500 ft. (CB), Oliver 4586 (K, STE). 3325 (Port Elizabeth) Deysels Plaat farm, Groendal Wilderness Reserve (CA), Scharf 1930 (PRE); Groen- dal Wilderness Reserve, Enon Conglomerate, on a burnt S-facing slope, Scharf 1517 (PRE). 4. Geissorhiza outeniquensis Goldbl., sp. nov. TYPE: South Africa. Cape: George, Jonkers- berg, Outeniqua Mts., wet rocky stream banks and waterfalls, Esterhuysen 19379 (holotype, BOL; isotypes, K, PRE). FIGURE 21. theris 6-9 mm longis, stylo diviso propre apicem an- therarum, ramis recurvatis. Plants 20—50 cm long, usually decumbent or FIGURE 21. 525, Outeniqua Pass). trailing, forming clumps. Corm depressed-glo- bose, symmetric, 6-8 mm diam., tunics appar- ently concentric, light brown, papery and soft, decaying readily. Cataphyll evidently lacking. d sometimes bract-like, 3—6 mm wide, linear, usually reaching to the base of the spike, sometimes longer. Stem decumbent, unbranched. Spike 2—4-flowered; bracts 16-20 (-24) mm long, herbaceous, becoming submem- branous above, the inner slightly shorter than the outer. Flower zygomorphic, declinate, pink ANNALS OF THE MISSOURI BOTANICAL GARDEN gs [Vor. 72 x 7 = V SJ al iy Up ten ES pis NATED VV 7 ALLY 2. aa AE we SA ES RT rz Lo a ¢ ç = m m ° 0 Om n n n ——— | i = EP Morphology and distribution of Geissorhiza outeniquensis. Habit x0.25; flower life size (Vlok to purple with darker markings near the base of the lower tepals, purple on the reverse; perianth tube 10-13 mm long, cylindric, about as long or slightly longer than the bracts; tepals 26-32 mm long, oblong, 10-14 mm wide. Filaments equal, declinate, 15-20 mm long; anthers 6-8 mm long, pollen yellow. Ovary ca. 5 mm long, style divid- ing at the apex of the anthers, branches 3-4 mm long, recurved. Capsule unknown. Chromosome number, 2n = 26 (Vlok 525). Flowering time. December to mid-January. Distribution. Local in the Outeniqua Moun- tains, in shady situations near water. Figure 21. 1985] Geissorhiza outeniquensis has until now been considered a form of the more widespread G. roseoalba, which until recently was referred to the genus Engysiphon. It clearly belongs in sec- tion Weihea and seems closely related to G. ro- seoalba. These are the only two species of the section which have a zygomorphic flower with declinate stamens and style. They also have the largest flowers in the section and are so distinc- tive that they are unlikely to be confused with other species of the section. The flowers of G. outeniquensis are very similar to those of G. ro- seoalba but are normally slightly larger and have a longer perianth tube, some 10-13 mm long, compared to 8-10 mm in G. roseoalba. The very soft textured corm tunics of G. outeniquensis, owever, are so different from those of G. ro- seoalba that the two species are unlikely to be confuse bpeissatliizá outeniquensis is apparently re- stricted to the Outeniqua Mountains immedi- ately around George, where it occurs in moist shady kloofs, along streams and waterfalls. It has an unusual caespitose habit, with the main plants producing short runners which sprout to form new stems and leaf clusters. The species seems to have lost the deciduous habit of the genus, and does not die back and become dormant dur- ing the later summer and autumn. Specimens examined. ed i524 CAPE: 3322 (Oudtshoorn) Jonkersberg, Out ua Mts., stream banks and waterfalls (CC), DURO 19379 (BOL, K, PRE); Outeniqua Pass, streambanks and seepages (CD), Acocks 21738 (B, K, NBG, PRE), Vlok 525 (MO, NBG, PRE); *Geelhoutbosch," George, damp places, -— 1628 (BOL); bri Pass, McNeil & Ising n. (PRE), Zinn s.n. (SAM 54823). 5. Geissorhiza fourcadei (L. Bolus) Lewis, J. S. African Bot. 7: 32. 1941. Acidanthera four- cadei L. Bolus, Ann. Bolus Herb. 4: 118. 1928. TYPE: South Africa. Cape: S slopes of Witte Els Berg, Humansdorp, Fourcade 2968 (lectotype, BOL; isolectotypes, BOL “Herb. Fourcade," MO, PRE, S, STE). FiGure 22. Plants 15-30 cm high. Corm nearly spindle- shaped, 6-10 mm diam., corms of previous sea- sons persistent and accumulating below current corm, tunics brown, woody to soft-papery, layers concentric but sometimes apparently imbricate by displacement especially when soft, fragment- ing irregularly from above and below, outer lay- ers sometimes becoming fibrous, extending up- ward in a neck together with decaying leaves. Cataphyll membranous, light to dark brown. GOLDBLATT — GEISSORHIZA 315 Leaves 3-5, terete or nearly terete, the margins and midrib actually much enlarged, and thus nar- rowly 4-grooved, half as long to as long as the stem, the lower 2-4 leaves basal, the uppermost cauline, partly to entirely sheathing, shortest. tem erect, smooth, simple or 1-2-branched, bearing one partly sheathing foliage leaf and 2- 3 dry sheathing stem bracts. Spike single flow- ered (both main and lateral axes); bracts herba- ceous, drying near the apex, (12-)16-30 mm long, S inner as long, or slightly shorter than the Flower hypocrateriform, pink to mauve; perdis tube (8—)17—20 mm long, cylindric, wid- ening gradually towards the apex, exserted, or just short of the apex of the bracts; tepals 25-30 mm long, oblanceolate, to 11 mm wide. Fila- ments 13-20 mm long, equal; anthers 10-12 mm long. Ovary ca. 4 mm long, style declinate, di- viding beyond the anthers, branches recurved Capsule unknown. Chromosome number, un- Flowering time. Mid-January to early May. Distribution. Mountains of the southern Cape, in seasonally moist sites, such as seeps, streamsides, and marshes. Figure 22 Geissorhiza fourcadei is distinctive in Geis- sorhiza in the persistent bases of leaves and cata- phylls which accumulate in a neck around the stem base and in the spike and its branches al- ways being single flowered. It is also peculiar in that the corms of past seasons often persist in- stead of decaying, so that a series of old corms accumulate behind the current year's corm. It is also unusual in its late summer to autumn blooming habit. The affinities of G. fourcadei are uncertain but the corm tunics correspond fairly well to the concentric type characteristic of sub- genus Weihea. It seems isolated in the subgenus but is best placed in section Weihea in which there is a marked tendency to the reduction of the number of flowers in the spike. It stands out in the section in its terete 4-grooved leaf. Diffi- culties in placing the species to subgenus arise because the corm tunics are often soft in texture, sometimes even fibrous, and are then easily dis- placed from their concentric position when up- rooted. The tunics then match neither the typical concentric tunics of subgenus Weihea nor cor- respond well with the imbricate tunics of sub- genus Geissorhiza. The tunic layers of some col- lections, notably Vlok 812, are hard and perfectly concentric, and it seems clear that the tunics are basically concentric even though they are often easily displaced when disturbed. 316 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 TN ` | SR > SN << >i NE e ox = RE23. Distribution a aes foliosa, G. On and G. delica and differences are discussed under G. nigro- montana. y UN LI ollected, and there are few recent collections of the species, despite the fact that it is recorded from a well-collected and easily accessible area. Specimens examined. SOUTH AFRICA. CAPE: 3321 (Ladismith) Kampsheberg, 1,000 ft. (CD), Muir 3336 (BOL). 3420 (Bredasdorp) Swellendam (AB), Ross Frames s.n. (BOL 31888); below 10 O'Clock Mt., Swellendam, in pastureland, Wurts 438 (NBG), 480 (NBG): Kolo- niesbos, fynbos, van Wyk 351 (STE); roadside near Zuurbraak (BA), Hurling & Neil s.n. (BOL 31889); Buffeljagsrivier bei Rietkuil, Ecklon & Zeyher 3961 (124.10) (BM, BOL, FI, G, K, S, SAM, W, Z); Ruggens near Zuurbraak, 800 ft., Galpin 4711 (BOL), 4715 (K, PRE); *Zwellendam, berge bei Puspas Valei, Duivels- bos etc." (BA-BB), Ecklon & Zeyher Irid. 218 (70.10) (B, BM, E, FI, K, L, MO, P, PRE, S, SAM, W, Z pug: ee and Heidelberg (BB), Lewis 5655 (NBG 3421 as near Riversdale (AA), Muir 4718 (BOL); between s. and Albertinia (AB-BA), Stayner s.n. (NBG ). WITHOUT ss LOCALITY: Caledon district, Pappe s.n. (SAM 20879). 7. Geissorhiza nigromontana Goldbl., sp. nov. TYPE: South Africa. Cape: Groot Swartberg, Blesberg at Tierkloof, along stream, Vlok 93 (holotype, MO; isotypes, NBG, SAAS). Planta 10-16 cm alta, tunicis cormi manifeste con- u : ein prostrato supra erecto, infra bulbilifero, spica 2 rum, floribus verosimiliter stellatis, caeruleo-purpur- eis, tubo ca m longo, tepalis 14-16 mm longis, antheris ca. 5 mm longis, idi diviso ad edd an- therarum, ramis recurvatis ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 Plants 10-16 cm high. Corm globose, asym- metric with an obliquely flattened side extending downward shortly, ca. 6 mm diam., tunics ap- parently concentric, light brown, outer layers ap- parently brittle-papery. Cataphyll pale, mem- branous. Leaves 3—6, falcate, at least the lower two basal, upper cauline and becoming smaller, ometimes 3-4 on a stem, the uppermost often bract-like, 4-10 mm wide, reaching to the base of the spike or only half to one-third as long as the stem. Stem apparently prostrate below, erect above, simple or sometimes branched, both from the base or from upper stem nodes, bearing cormlets from the lower nodes. Spike 2-3-flow- ered; bracts 8-10 mm long, green, becoming membranous above, the inner slightly shorter than the outer. F/owers apparently stellate, blue- purple; perianth tube 2-3 mm long, cylindric, included in the bracts; tepals 14-16 mm long, obovate, ca. 6 mm wide. Filaments ca. 6 mm long; anthers about 5 mm long, pollen yellow. vary ca. 3 mm long; style dividing at about the midline of the anthers, branches long, 4-5 mm long, recurved. Capsule unknown. Chromosome number, unknown. Flowering time. January to February. Distribution. Great Swartberg Mountains, local along streams on Blesberg at ca. 1,800 m Figure 23. Geissorhiza nigromontana is apparently a nar- row endemic of the Swartberg Mountains, where it has been collected only twice by J. J. Vlok in 1981 and 1985. It blooms towards the end of summer, in January and probably into February, in this area of low rainfall and depends for mois- ture on the permanent streams along which it OWS. It apparently has fairly soft corm tunics, but few of the specimens available have corms, and their exact nature is difficult to assess with so little material to examine. Geissorhiza nigro- montana is most probably closely related to G. foliosa, a species of the Langeberg foothills, which blooms mainly in the months of October and November, and it has the broad flat leaves typical of that species. It is distinctive in its short tubed flower, with the tube ca. 2 mm long, few flowered spike and relatively long style branches 4-5 mm long. In contrast, G. foliosa has a tube 5-6 mm long, a spike of (1—)3-7 but usually several flow- ers and style branches to 3 mm long. Geissorhiza nigromontana is also distinctive in bearin cormlets in the lower stem axils, a characteristic rare in the genus. 1985] Specimens examined. SOUTH AFRICA. CAPE: 3322 (Oudtshoorn) lower S slopes of the Swartberg next to waterfall, near Rus en Vrede (AD), Vlok 893 (MO, NBG, SAAS); Groot Swartberg, Blesberg at Tierkloof, along stream (BC), Vlok 93 (MO, NBG, SAAS) 8. Geissorhiza delicatula Goldbl., sp. nov. TYPE: South Africa. Cape: shady mountainside, Seven Weeks Poort, Compton 4016 (holo- type, NBG; isotype, BOL) Planta 3-9(-1 2) cm alta, foliis 3-6, inferioribus 2-4 rum, floribus stellatis, pallide caeruleis, tubo 2-3 mm longo, tepalis 7-12 mm longis, antheris 2.5-3.5 mm longis, ne diviso prope apicem antherarum, ramis recurvatis Plants 3-9(-12) cm high. Corm globose, asym- metric with an obliquely flattened side extending 3-8 mm diam., tunics con- oft membranous, frag- menting irregularly into sections. Cataphyll brown to transparent, membranous, sometimes conspicuous. Leaves 3—6, erect to falcate, 1.5- 4.5 mm wide, linear to ensiform, often prostrate, half as long as the stems, at least the lower 2-3 basal, the upper cauline and becoming smaller, uppermost often bract-like. Stem erect, simple 3-flowered; bracts 4-8 mm long, herbaceous, be- coming membranous and dry above and some- times reddish along the margins, the inner about as long as the outer. Flowers stellate, pale lilac purple, cream in the throat and tube; perianth tube 2-3 mm long, cylindric, included in the ima dn Mes 7-12 mm long, obovate, 2.5-4.5 mm wide. Filaments 3-5 mm long; anthers 2.5— 3.5 mm long, pollen yellow. Ovary 1.5-3 mm long, pis dividing towards the apex of the an- thers, branches to 1.5 mm long, recurved. Cap- sule 4-7 mm long, narrowly obovoid. Chromo- some number, unknown. Flowering time. August to September in drier sites, November to December at higher altitudes. Distribution. Sheltered places in the interior mountains ofthe western and southern Cape from Jonas Kop in the west to the Swartberg Pass in the east. Figure 23. Geissorhiza delicatula is apparently related to the more we own G. inconspicua and G. d liosa, both p fthe wetter coastal ranges, extending from the Lange- berg at Swellendam to near Port Elizabeth. It is GOLDBLATT— GEISSORHIZA 319 always a much smaller and more delicate plant and it can be distinguished from apa species by its normally s n the most robust individuals of G. delicatula seem to have this habit. It also differs from G. inconspicua and G. foliosa in its slightly smaller flowers, with te- pals 7-12 mm long and short perianth tube, 3- 4 mm long, which is always included in the bracts. Geissorhiza inconspicua and G. foliosa normally have flowers with tepals 8-17 mm long and a tube 4—6 mm long, and typically have spikes of 3-6 flowers. A collection from the Swartberg, Oliver 3056, seems to differ in having leaves of a somewhat firmer texture than other collections. It also has larger flowers than most other gatherings of the species, but at least in the flower size, a collection from nearby, Stokoe s.n. from the Swartberg Pass, is comparable. From the material available, it seems best to regard these specimens merely as a form of G. delicatula. All the collections of the species except Oliver 5359 from the Rooiberg, per branch. It is possible that this collection represents a very depauperate form of G. inconspicua. Specimens examined. SOUTH AFRICA. CAPE: 3319 (Worcester) Karoopoort % Hafstrom & Acocks 288 (PRE, S); Jonas Kop, ca. 3,000 ft., wet shaded sites on S side of ae rocks (DC), Goldblatt 4723 (MO), 5880 (M 3321 cm Buffelskloof, Swartberg near Ladi- smith (AC), Esterhuysen 14007 (BOL); shade on S slopes below Towerkop, Esterhuysen 13967 (BOL); Seven Weeks Poort, mountain slopes, SW aspect (AD), Es- terhuysen 24838 (BOL); shady mountainside, Seven Weeks Poort, Compton 4016 (BOL, NBG); above Sev- en boo Poort, Phillips 1543 (SAM); Swartberg, be- tween Waboomsberg and Kangoberg, 6,000 ft. (BD), Oliver 3056 (PRE, STE); Rooiberg, summit ridge above Pass (AC), Stokoe s.n. (SAM 68346); La to Daskop near turnoff to Matjesrivier (DC-DD), 301 (MO). Vlok 9. Geissorhiza bracteata Klatt, Abh. Naturf. Ges. Halle 15: 391 (Erg. 57) 1882; Baker, Handb. Irid. 158. 1892, Fl. Cap. 6: 74. 1896; Foster, Contr. Gray Herb. 135: 62. 1941. TYPE: South Africa. Cape: Olifantshoek, Uitenhage, T lon & Zeyher s.n. (holotype, B “Herb. Lu- beck”), Ecklon & Zeyher Irid. 221 (10. a (B, BM, E, FI, G, LD, MO, PRE, S, W, Z, from the same locality are probably isotypes). FIGURE 24. 320 ANNALS OF THE MISSOURI BOTANICAL GARDEN M [Vor. 72 FIGURE 24. Morphology of Geissorhiza bracteata and distribution of G. bracteata and G. nana. Habit x0.5; flower and separated outer and inner bracts life size; leaf section much enlarged (no voucher, Uitenhage). Weihea elatior Ecklon, Topogr. Verz. Pflanzensamml. E 27, nom. nud. Geissorhiza tabularis var. elatior Ecklon ms. [Ecklon & Zeyher Irid. 221 (10.9)]. Geissorhiza recurvifolia (Poir.) Klatt sensu Klatt, Lin- naea 34: 655. 1866, pro parte excl. basion. Plants (4-)6-18 cm high. Corm ovoid, asym- metric with an oblique flat side extending shortly downward, 4—6 mm diam., tunics pale to dark rown, concentric, the outer layers fragmenting irregularly into sections and drawn into points above. Cataphyll pale, membranous. Leaves (3—4 to several, the lower basal, upper cauline and becoming smaller, sometimes bract-like, 2— 9 mm wide, ensiform, half to one-third as long as the stems. Stems erect, single or 2-5 produced from base and occasionally higher on the stem. Spike usually 1-flowered, rarely 2-3; bracts green, often reddish along the margins, 6-10 mm long, the inner usually slightly shorter than the outer. Flowers stellate, white, the outer tepals some- times red-flushed on the reverse; perianth tube 3-5 mm long, infundibuliform, widening from the base; tepals (S—)8—12 mm long, obovate, 4.5— 6 mm wide. Filaments 5-7 mm long; anthers 2— 3.5 mm long, pollen yellow. Ovary 2-3 mm long; style dividing at the apex ofthe anthers, branches recurved. Capsule 7-9 mm long, narrowly ovoid, seeds angular. Chromosome number, 2n — ca. 52 (Goldblatt 4933). Flowering time. September to mid-October. Distribution. Southern and eastern Cape, from Albertinia east to Grahamstown and in the southern Karoo, often in clearings or on edge of forest. Figure 24. Geissorhiza bracteata is one of the least spe- cialized species in the genus, having several flat, ensiform leaves, a branched stem, and concentric corm tunics, all of which are considered primi- tive in Geissorhiza. It is closely allied to G. in- conspicua and sometimes difficult to separate from that species. It can be recognized by its 1(-3)-flowered spikes and branches produced from the base of the plant. Geissorhiza incon- spicua is a variable species, and depauperate, few flowered specimens of the white-flowered form approach G. bracteata. It is also possible to con- fuse G. bracteata (e.g., Bayliss 6035, Duthie 1132) with G. setacea although G. setacea generally has much shorter and narrower leaves, typically less than 2 mm wide. Geissorhiza setacea occurs on damp sandy flats in the southwestern Cape. This is well to the west of the range of G. bracteata, which is centered in the eastern Cape between Grahamstown and Humansdorp, although pop- ulations have been recorded as far to the west as Albertinia. Geissorhiza bracteata appears to be a poly- ploid species, 2n = ca. 52, at least on the basis 1985] of counts for one population. This population, from near Hankey, has proved in the greenhouse to be autogamous, which is unusual in Geisso- rhiza. Specimens examined. SOUTH AFRICA. CAPE: 3224 (Graaf -— near Jansenville (DC), Bayliss 5925 (BR, K, MO, W 3225 Astoria East) Cradock Common (BA), Bay- liss 4967 (GH, MO, UC, US). 3321 (Ladismith) Welgevonden near Plattebosch, Albertinia (DC), t 1811 (BOL); Cloetes Pass (DD), Goldblatt 4158 (M 3322 asa ie Cradock Berg, George (CD), Galpin 4673 (K, PRE); pepe Lewis 5603 (NBG). 3323 (Willowmore) Avontuur-George road, farm Arbor Nook, yu apa ane Nee. Snijman 365 (NBG); station above Uplands, 11.3 mi. from Kruis River Nek (CC-CD), oe u 12 (STE); Lottering (DC), Fourcade s.n. (BO 3324 (Steytlerville) Ds River, 4 mi. W of Com pany's Drift (CC), Fourcade 2317 (BOL); Bo Plaas, near Humansdorp (CD), Bayliss 6035 (MO); 5.5 km N of Hankey (DD), Goldblatt 4933 (K, MO, PRE, US). 3325 (Port Elizabeth) hills at Addo (BC-DB), Ecklon & Zeyher Irid. 220 (SAM), Ecklon & Zeyher Irid. 221 (S); Zuurberg Pass, cool SE-facing slopes (BD), Snij- man 489 (MO, NBG); Groendal Wilderness Reserve (CA), Scharf 1493 (PRE); Despatch, OMNE (CD), Holland 2209 (BOL), Holland s.n L . Gard. 2687/25 in BOL); Port Elizabeth Td Fries, p eo & Weimarck 216 (LD, SAM), Paterson 152 (BOL). 3326 (Grahamstown) Grahamstown (BC), Mac- 9 (SAM); along the the rivulet at Grahams- town, Burchell 3546 (K); Mountain Drive, Grahams- town, Bayliss 7231 (MO); Brickfields, Grahamstown, Daly 861 (B, BOL, Z); Hopewell, S of Southwell (BD), Acocks 16127 (PRE). 3422 (Mossel Bay) between Malgaten and Great Brak Rivers (AA), Fourcade 6481 (BOL); turf near Church, Belvedere, Knysna (BB), Duthie 1132 (BOL, MO, STE). 3423 DIM flats 14 mi. E of Knysna (AA), Gillett 1 = Agr L). iB roue hillside, Humansdorp (BB), Galpin uda E dud LiTy: Riversdale €: Sehlechter s s.n. 0) Uiteinde Ecklon & Zeyher s.n. (10.9) (B, L); Z aii near podium Loubser sos (NBG); Cape, Krebs (B), Verrea .( ay E o foci & Bud "irid. 221 (10. m (B, doe MO, P W, Z); Uitenhage, ae Wm n. (K); M sel EI Fries, Nor- lindh & poen 1287 (K, LD, PRE). 10. Geissorhiza nana Klatt, Abh. Naturf. Ges. Halle 15: 391 (Erg. 57). 1882; Baker, Handb. Irid. 157. 1892, Fl. Cap. 6: 73. 1896; Foster, Contr. Gray Herb. 135: 60-61. 1941. TYPE: South Africa. Cape: "Rivierzondereinde," Swellendam, Ecklon & Zeyher 3967 (90.9) (as Drége 3969) (holotype, B “Herb. Lu- beck”; isotypes, B, FI, G, K, S, SAM, W, GOLDBLATT — GEISSORHIZA 321 Plants tiny, 2.5—4(—8) cm high. Corm narrowly ovoid, asymmetric, with an oblique flat side, 3- 5 mm diam., tunics pale, concentric, drawn into bristles above. Catap pale, membranous. Leaves (3—)4—6, usually all basal, or uppermost inserted on the lower part of the stem, flat, half as long, to slightly longer than the stem, 1-2 mm wide, linear-ensiform. Stems erect, single, or to 4, produced from near the base, rarely branched above. Spike 1-2(-3)-flowered; bracts 4-6 mm long, green, sometimes red along the margins. Flowers tiny, stellate, white, red-brown on the reverse of the outer tepals; perianth tube 1—2.5 mm long, nearly cylindric; tepals obovate, 3-6 (-8) mm long. Filaments ca. 2 mm long; anthers 1.2-1.8 mm long. Ovary ca. 2 mm long; style dividing at mid-anther level, branches less than 1 mm long, recurved. Capsule 4-6 mm long, obovoid, usually slightly exceeding bracts. Chro- mosome number, 6 (Goldblatt s.n. no voucher, near Swellendam). M T: Flowering time. September to early October. Distribution. Clay flats and slopes in renos- terbosveld, in the Caledon district and eastwards to Heidelberg. Figure 24. Geissorhiza nana is a very dwarf species of Geissorhiza clearly related to the southern and eastern Cape G. bracteata. It is in fact difficult to distinguish Ln forms of the latter rom G. nana. The flower is critical in separating the two: : G. nana has tiny flowers with tepals 3— 6 mm long, with very pronounced darker mark- ings on the reverse of the outer tepals, while G. bracteata usually has quite large flowers with te- pals in the 8-12 mm long range, and coloring on the reverse of the outer tepals is rare. Difficulties are, however, experienced in distinguishing G. nana from the depauperate forms of G. bracteata found in the southern Cape, which may have tepals as short as 6-7 mm, - even these plants are usually larger than G. n here is also the acini of confusing Geis- sorhiza nana with G. setacea, another dwarf species of section uu Geissorhiza setacea is very similar in general appearance, but it is larger in all features, having flowers with a tube ca. 6 mm long and tepals in the 8-12 mm range. Geis- sorhiza setacea grows on damp sandy flats along the western Cape coastal belt, whereas G. nana grows in drier clay soils in the southern Cape. My impression is that Geissorhiza nana is not uncommon in renosterbosveld from the Caledon district east to Riversdale. The few collections 322 — Q "d SE = J FiGunE 25. Morphology and distribution of Geiss- orhiza setacea. Habit x0.5; flower life size (Goldblatt 6193, flats near Gordons Bay). of the species probably reflect the fact that it is very inconspicuous and thus easily overlooked. T divin examined. SOUTH AFRICA. CAPE: 3419 of Caledon (AB), Goldblatt B 1 (BOL); Caledon, commonage S of the town, in cla gravel, pred 6198 (MO); turnoff to Speelmansri- vier, E o on near Caledon (BA), Oliver 6020 (STE); Ae sic ai ard (BB), Ecklon & a 5 ed (90.9) (B, FI, G, K, S, SAM, W, Z); c ar Mier farm, SW of Bredasdorp (DB), Goulblat 61 85 (K, MO, NBG). 3420 (Bredasdorp) clay road verge between Heidel- berg and Strawberry Hill (BB), Goldblatt 4961 (MO); es Heidelberg, Loubser 2126 (NBG). 1 (Riversdale) Korente River Dam, clay hill (AA), Bohnen 6663 (STE). 11. Gei hiza setacea (Thunb.) Ker, Ann. Bot. (Kónig & Sims) 1: 224. 1804, Bot. Mag. tab. 13: 146. 1783. TYPE: South Africa. Cape: without precise locality, Thunberg s.n. [lec- totype, Herb. Thunb. 995, effectively des- ignated by Foster (1941); isolectotypes, S ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 *Herb. Osbeck,” “Herb. Alstroemer,” “Herb. Montin"]. FiGURE 25. Plants small, 4-8 cm high. Corm globose, asymmetric, one side flattened below and ex- tending downward in a fan-like ridge, 3-4 iam., tunics concentric, outer layers irregularly areca Cataphyll membranous. Leaves usually 4, the lower three basal, about half as long as stem, erect to falcate, linear-ensiform, 1— 2.5 mm wide, lamina flat, the uppermost leaf cauline, usually smaller than the basal leaves, the lower half sheathing the stem. Stem erect, oc- casionally 1 or rarely 2-branched. Spike 1(-rarely 2)-flowered; bracts 5-7 mm long, herbaceous, the upper margin sometimes reddish, the inner slightly longer than the outer. Flower hypocra- teriform, cream white or pale yellow with a dark center, flushed red to purple on the reverse of the outer tepals especially with age; perianth tube ca. 6 mm long, infundibuliform; tepals (8—)10— 12 mm long, obovate, 6-8 mm wide. Filaments 3-4 mm long; anthers 2.5—4 mm long. Ovary ca. 3 mm long; style dividing at the apex of the an- thers, branches recurved. Capsule unknown. Chromosome number, unknown. Flowering time. July to August (to Septem- ber). Distribution. Wet flats in the southwestern Cape from the Peninsula east to the foot of the Hottentots Holland Mountains and north to Gouda. Figure 25. This small species can easily be recognized by its fairly large, and relatively long-tubed flowers borne singly on short stems. Its corm tunics are of the concentric type, and it belongs in subgenus Weihea, near Geissorhiza bracteata and G. nana. Geissorhiza setacea can readily be distinguished from these species by its large flower, with tepals 8-12 mm long, on a very small plant seldom exceeding 8 cm in height. The very dwarf G. nana is similar but smaller, usually only 4-5 cm high and has tiny flowers with tepals 3-6 mm long. It is typical of clay soils in contrast to G. setacea which grows on damp and often water- logged sandy flats. It was one of the early species of Geissorhiza to be named, having been described by Thunberg in 1783, as Ixia setacea, from material he col- lected during his travels at the Cape in the years 1772-1775. It has generally been recognized as a distinct species of Geissorhiza, although partly confused with G. bracteata by J. G. Baker (1896) 1985] and Foster (1941), both of whom included spec- imens of G. bracteata under. G. setacea. . This confusion i 1 of G. bracteata do resemble G. setacea, The plant illustrated by Ker (1810) as G. setacea in the Botanical Magazine tab. 1255 is G. hispidula Goldbl. (G. humilis var. bicolor Baker), a differ- ent and not very closely related species. Geissorhiza setacea usually has white flowers, but in the north of its range, near Gouda, at Elandsberg farm there is a creamy yellow form, which has a blackish center. This form grows sympatrically with and blooms at the same time as G. purpureolutea which has almost identical flowers, but imbricate corm tunics and ribbed leaves and belongs in subgenus Geissorhiza. This is one of several examples of floral convergence found in Geissorhiza, where similar flower color and marking occur repeatedly in unrelated species or in other genera. Specimens examined. SOUTH AFRICA. CAPE: 3318 (Cape Town) near pd Town (CD) H. Bolus 4804 (BOL), 4803 (E, K, Z); Table Mt. near ped Kloof, Schlechter 412 (Z); flats near Hoorn Kloof, P. Tg, , K); near Stellenbosch tsi E); clay slope 300 452 orcester) Elands i farm, S of Gouda, wet fats TEN Goldblat tt 6210 (K, M O, NBG, PRE, S). 418 (Simonstown) Van Der E Stellenbosch (BB), S 2. 3204 (PRE); low lying, wet area SW of Somerset West, Boucher & Mauve 4946 (PRE); sandy flats be- tween Strand and Gordon’s p Parker 4227 (BOL, NBG); Faure, flats Sir pu Pass, Stokoe s.n. (SAM 49615), Schlechter 1 T As. , Z). PRE ISE LOCALITY: Caledon, Pappe s. (SAM $5323 C ape, Thunberg s.n. (Herb. Thunb. 995 UPS, S), Oldenberg 489 (BM). 12. Gelssorhiza ornithogaloides Klatt, Linnaea 56. 1866. TYPE: South Africa. Cape: Caledon Zwartberg and about the Baths, Ecklon & Zeyher Irid. 225 (51.8) (lectotype, B, here designated; isolectotypes, B, BM, BOL, E, FI, G, GH, L, LD, MO, W, Z); Worcester, Tulbaghskloof, Winterhoeks- berg, Tulbaghsthal etc., Ecklon & Zeyher Irid. 224 (77.9) (syntypes, B, LD, Koue Bokkeveld, Lichtenstein 15 (syntype, B) [chosen arbitrarily by Foster as lectotype for G. ornithogaloides, being the first of the three cited type collections; it lacks a corm — ee GOLDBLATT— GEISSORHIZA 323 and cannot be certainly identified and dis- tinguished from G. ornithogaloides subsp. marlothii (syn. G. marlothii)]. FIGURE 26. Plants small, 4—10 cm high. Corm either more or less ovoid or campanulate in outline with a broad flat base, 4—6 mm at widest diam., tunics medium brown, concentric, either fragmenting into elliptical sections or, in two parts, a cam- panulate cap and basal plate, then lower margins of cap toothed. Cataphyll membranous, often evidently lacking. Leaves usually 4, the lower two basal, linear to lanceolate, erect to falcate, 1-2 (-3) mm wide, flat, the upper two inserted on the stem at or above ground level, and sheathing below, the sheath often inflated. Stem erect, usu- ally branching from both cauline leaves. Spike a 2)-flowered; bracts pale green, becom- ing dry above, 7-10 mm long. Flower clear yel- low, coat "Dern tube 2-3 mm long, not exserted e bracts; — obovate (6—)7— 12(-18) mm ise 3.5-6 mm wide. Filaments 2- 4mm long, erect; anthers 3-4.5 mm long, yellow. Ovary ca. 2 mm long, style dividing at the apex of the anthers, branches 2-3 mm long and re- curved. Capsule obovate, 7-9 mm long, usually exceeding bracts. Chromosome number, 2n — 26 (Goldblatt 214 as G. nana). Flowering time. Late August to October. Distribution. Sandy and clay flats and slopes from the Paarl district eastward through the Cold Bokkeveld, Worcester district and southern Cape to the Long Kloof. Figure 26. Geissorhiza ornithogaloides is typically a dwarf and rather aac species with unusual, , bright yellow flowers. It some i RUN ina population may be robust and spikes with either one or two flowers, two nar- row, flat, basal leaves, and two cauline and partly sheathing leaves. This pattern is found in Geissorhiza marlothii, a species which is identical to G. ornithogaloides in all features except the corm, the reason that Foster (1941) recognized the former as distinct. Geissorhiza marlothii has a small corm with con- centric tunics of the type characteristic of all species of subgenus Weihea. The corm of G. or- nithogaloides is unusual in the genus, being cam- panulate in outline, with an oblique flat base and ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 G. ornithogaloides un subsp. marlothii Q subsp. ornithogaloides GTE A] : AÑ ged Eu PM < < 20 21 22 3 FIGURE 26. Morphology and distribution of Geissorhiza ornithogaloides. Habit with flower life size; opened flower and gynoecium x2 (Goldblatt 6202, Villiersdorp-Brandvlei). serrate lower margins. The morphological sim- ilarity between the two species, excepting the corms, seems to me so striking, however, that I prefer to treat G. marlothii as a subspecies of G. ornithogaloides. e only close ally of Geissorhiza ornithoga- loides is G. malmesburiensis, described by Foster in 1941 on the basis of a single Schlechter col- lection, which lacked corms. A second sheet of the Schlechter collection, and collections made by Lewis and by myself, have corms which are exactly like those of the typical subspecies of G. ornithogaloides, which substantiates Foster’s be- lief that they were in fact closely related. Geis- sorhiza malmesburiensis, in fact, differs from all forms of G. ornithogaloides, mainly in its large flower size which contrasts with its very dwarf vegetative morphology. I have found it necessary to alter the lectotyp- ification of Geissorhiza ornithogaloides, the rea- sons being as follows. Of the three collections cited by Klatt in the protologue, two are Ecklon and Zeyher collections from the Caledon district, both widely well-distributed in major herbaria, and often complete with corms. The first speci- men cited, however, is a Lichtenstein collection supposedly from the Cold Bokkeveld which to- day lacks corms, though they were present at one time according to the description. Both the dis- tribution and description of the corms suggests 1985] that this last collection is G. marlothii, yet was chosen as lectotype by Foster, for G. ornitho- galoides, a species only distinguishable from G. marlothii by its corms. Foster was clearly arbi- trary in choosing this first cited collection as lec- totype, and I propose in place Ecklon & Zeyher Irid. 225 in the Berlin Herbarium as the lecto- type. The present interpretation of G. ornitho- galoides is in this way preserved. The two subspecies of Geissorhiza ornitho- galoides are treated below, with subsp. marlothii first, as it is clearly the less specialized, having corms of the basic type for the subgenus, whereas corms of subsp. ornithogaloides are unusual and must be regarded as derived. Y TO THE SUBSPECIES OF GEISSORHIZA ORNITHOGALOIDES la. Corm nearly ovate, tapering at iie end: not campanulate in outline; and low argins of tunics not or rarely slightly ng MM subsp. marlothii b. with an oblique flat base; the lower margins of the tunics serrate subsp. ornithogaloides 12A. Subsp. marlothii (Foster) Goldbl., comb. et stat. nov., Geissorhiza marlothii Foster, Contr. Gray Herb. 135:66. 1941. TvPE: South Africa. Cape: Cold Bokkeveld, Houden- beck, 850 m, Marloth 10612 [holotype, B; isotypes, PRE, STE (without corms)]. ? Ixia e, ye ret! Licht. ex Roem. & Schult., hn Veg. 1: 376. 1 frica. Ca Bokkeveld, Lichtenstein 15 (holotype, B). (Spec. ns lac d to distinguish the species from n G. ornithogaloides anh but from the stated locality, must be su arlothii. This is not the basionym of G. pars diruit Klatt.) Corm nearly ovoid, tapering at both ends, is Aid flattened below on one side, sometimes with the flat a extending apis conspic- bs. 3-5 m iam., tunics concentric, frag- menting peer into elliptical segments Flow- ers sometimes very small, with tepals (6—)7-12 (-15)mm long. Filaments 2-4 mm e anthers 3-4 mm long. Chromosome number, unknown. Flowering time. Late August to October. Distribution. Sandy and shale slopes and flats, Cold Bokkeveld, and east through the Warm Bokkeveld and the Witteberg and Voetpadsberg, with outlying disjunct populations in the Little Karoo, the Albertinia district and Long Kloof in the southern Cape. Figure 26 GOLDBLATT — GEISSORHIZA 325 Subspecies marlothii has an ovate-elliptic corm with concentric tunics. This corm is typical of subgenus Weihea and must be viewed as the an- cestral type in G. ornithogaloides, while the cam- panulate corm of subsp. ornithogaloides is spe- cialized and derived. In all other features, the two subspecies are essentially identical, although there are several populations of subsp. marlothii on the edges of the range of this subspecies that have smaller flowers than any found in subsp. ornithogaloides. The smaller flowered collections are those to the north and east of the main range for subsp. marlothii, which is centered in the Cold and Warm Bokkeveld. The northern extension is rep- resented by a collection from the Koudeberg near Compton 3806) and populations in the es Karoo, Al trict (southern Cape) and in the Long Kloof. The eastern populations are few and scattered and subsp. marlothii is apparently rare outside its main range. There is nothing as yet known to suggest that these eastern dier iam represent a distinct taxon as they are similar to several flowered plants from other Nue notably to the Schlechter and Compton collections men- tioned above. Specimens examined. SOUTH AFRICA. CAPE: 3219 (Wuppertal) pei Wuppertal (AC), Schlechter dede BM, BOL, BR, E, G, H, K, L, MO, PH, PRE, S, Z), Houdenbeck, Cold Bokkeveld (AD), Marloth 10612 (B, PRE, STE), Cold Bokkeveld, 3 mi. S of Leeurivier (CC), Lewis 2637b (SAM); Bokkeveld Ta- felberg (CD), Leighton s.n. (BOL); Cold Bokkeveld, flats at De Keur, Esterhuysen 13006 (BOL, PRE); Schoongesig, Ceres, Hanekom 1222 (K, PRE, STE). 3319 (Worcester) 7 mi. W of Gydo Pass, Cold Bok- keveld (AA), Hutchinson 1055 (K, NBG); Gydo, Ceres (AB), Leipoldt 3811 (BOL); 6 mi. N ofthe top of Gydo 5120 (K, MO, PRE, S, STE); "— (BC), Olivier s.n. (NBG 60692); Theronsberg Pass, De Vos 1673 (STE); Orchard Siding, Worcester, poda 16570 (K), 16666 (K). 3320 (Montagu) poort N of Pienaarskloof, Cere (NBG); vov aig x l uur-George road, farm Arbor Nook, M (CA), Snijman 361 BG). 3324 (Steytlerville) Company's Drift, Long Kloof (CO), Fourcade 2313 (BOL, K, STE). 326 3421 (Riversdale) Albertinia (BA), Erasmus 1578 (STE), 1624 (STE); Plattebosch, near Albertinia, Muir 1810 (BOL) WITHOUT PRECISE LOCALITY: Ceres Wild Flower Show, Lewis s.n. (BOL 21246). 12B. Subsp. ornithogaloides. FIGURE 26. Geissorhiza flava Reichb. ex Klatt, Abh. Naturf. Ges. Halle 12: 392 (Erg. 58). 1882. TYPE: South Africa. Cape: without precise locality, Breutel s.n. (ho- lotype, B). Waitzia flava Reichb. ms., Klatt, Abh. Naturf. Ges. Halle 12: 392. 1882, in syn. Geissorhiza ornithogaloides var. n (Klatt) Foster, Contr. Gray Herb. 135: 68. 1941. Geissorhiza romuleoides Ecklon, o: s Verz. Pflan- zensamml. Ecklon 27. 1827, nom. nud. et ms. [Ecklon & Zeyher Irid. 225 (51.8)]. Corm campanulate with an obliquely flattened base, 4—6 mm at widest diam., tunics medium brown, concentric, in two parts, with a cap, and separate basal disc, lower margins of cap serrate. Flowers with tepals 8—12(-18) mm long. Fila- ments 2.5-4 mm long; anthers 3-4.5 mm long. Chromosome number, 2n — 26 (Goldblatt 214 as G. nana). Flowering time. September to October. Distribution. Clay and sandy flats and slopes from the Paarl district east to Worcester and the Hex River Valley and south to the Caledon dis- trict. Figure 26. As already discussed, subsp. ornithogaloides differs from subsp. marlothii only in its corms. The corms of subsp. ornithogaloides are cam- panulate with a flat base, and have a serrate lower margin. Subspecies marlothii has ovate-elliptic corms, tapering at both ends, a type found in most species of subgenus Weihea. In other fea- tures, the two entially identical, except that some populations bf subsp. marlothii have smaller flowers than any found in subsp. ornithogaloides. Geissorhiza flava Klatt is a name applicable to larger flowered, robust forms of G. ornithoga- loides, and although corms are lacking from the what larger and usually 2-branched. As he com- ments, it tends to intergrade with the typical va- riety. With the considerably greater number of collections available to me, it seems clear that even this treatment is unacceptable. Variation within populations of G. ornithogaloides can be ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 considerable and robust plants corresponding to G. flava may grow next to small slender individ- uals matching the lectotype of G. ornithogaloides (e.g., Lewis 6052). Specimens examined. SOUTH AFRICA. CAPE: 3318 (Cape Town) near Hercules Pillar (DD), Salter 4721 (BM, BOL, K), Mathews s.n. (Nat. Bot. Gard. 1419/ 30 in BOL); Agter Paarl, Barker 404 (NBG 3319 (Worcester) Romansrivier (AC), Lewis 2636 (SAM); Romans River Nature Reserve, Goldblatt 6290 (MO, WAG); Buffelsrug, Osplaats Station, NE of De Doorns (BC), Mauve & Oliver 235 (STE); esa sandy flats near the water (CB), Lewis 6052 (N STE); Du M Aqu idi ina pets Barker ies (NBG); E s Kloof, sandy a Lewis 6043 (NBG, p ae 2412 ac NBG, S); be- n its the Breede River on stony roadside, Goldblatt 4703 (MO, NBG); r from Vil- liersdorp to Worcester (CD), PE wad yom Barker 75 BG); S of Quagga & Oliver 264 (STE); between Brandvlei and Jit 6 be dorp, Goldblatt 6202 (MO, PRE, S, WAG). 18 (Simonstown) sandy flat inier Gordons Bay and Strand (BB), Parker 4241 (GH , K). 3419 (Caledon) between French aa and Villiers- dorp (AA), Lewis 2898 (SAM); Sondereinde River bridge on Villiersdorp-Caledon road (AB), Salter 4792 Saas K); Caledon, Leipoldt s.n. (BOL); Caledon, wartberg and about the Baths, Ecklon & Zeyher Irid. ~ (51.8) (B, BM, BOL, E S, W, Z); Zwartberg, Caledon, Zeyher 3966 (B, FI, G, K, ; commonage below Zwartberg, Caledon, “a la n. (BOL 16917); hillside above vlei, Caledon oad 1 mi. from Villiersdorp turnoff, Barker 13 (BOL, Kx stony flats S of Villiersdorp at Theewater farms turnoff, Goldblatt 4016 (MO); road from Shaws Pass to riety ey Goldblatt 214 (BOL); Onverwacht, 19 m f Riviersonderend (BA), Acocks 22665 (PR E). 3420 (Bredasdorp) Potberg flats, De Hoop Nature Reserve asa Burgers 2248 (STE WITHOUT PRECISE LOCALITY: Cape, Breutel s.n. (B “Herb. Lube J. 13. Geissorhiza malmesburiensis Foster, Contr. Gray Herb. 135. 1941. TYPE: South Africa. Cape: hill near Malmesbury, 300 m, Schlechter 1654 (holotype, B; isotypes, B, BOL, BR, L, MO, PRE, Z). FIGURE 27. Plants small, 5-8 cm high. Corm campanulate with an oblique flat base, 4—6 cm at widest diam., tunics medium brown, concentric, in margins of the cap toothed. Cataphyll membra- nous, evidently often lacking. Leaves usually 4, the lower two or three basal, linear-filiform, more or less erect, to 1 mm wide, blade plane, the upper two or uppermost inserted on the stem at or above ground level, sheathing and inflated 1985] below, narrow above. Stem erect, simple or branching from the upper leaf or leaves. Spike 1 -flowered; bracts pale green, with a hyaline mar- gin, becoming dry above, 7-11 mm long. Flower clear yellow, cupped; perianth tube ca. 4 mm long, included in the bracts; tepals obovate, 13— mm long, to 8 mm wide. Filaments ca. 6 mm long, erect; anthers 4-5 mm long, yellow. Ovary ca. 3 mm long, style dividing at the apex of the anthers, branches ca. 4 mm long, recurved. Cap- sule unknown. Chromosome number, 2n — 26 (Goldblatt 6282). Flowering time. September to early October. Distribution. Restricted to the Malmesbury district. Figure 27. When Foster (1941) described this species from a single sheet of a Schlechter collection, he was unaware ofthe nature of its corm. A second sheet of the type collection also at Berlin Herbarium has several corm fragments, while the only other collections of Geissorhiza malmesburiensis, made by Lewis in 1952 and by myself in 1981, have intact corms, and it is now clear the species has the striking corm previously thought restricted to Geissorhiza ornithogaloides (here subsp. or- nithogaloides). The corm is campanulate in out- line with an oblique flat base, and toothed lower margins. The corm structure bears out Foster's suggestion that G. malmesburiensis is closely re- lated to G. ornithogaloides. The two in fact differ mainly in the size of the floral parts. Geissorhiza malmesburiensis has flowers much larger than any known in G. ornithogaloides, although all plants so far collected are rather small, slender, and few branched. Flowers of G. malmesburien- sis have a perianth tube ca. 4 mm long and tepals 13-25 mm long but usually in the 20-25 mm range, while in G. ornithogaloides the perianth tube is 2-3 mm long and the tepals (6—)7-15 mm long, rarely to 18 mm, and robust plants with larger flowers are always branched and have spikes of more than one flower each. This species is evidently extremely rare. The only known habitat, on the slopes above Mal- mesbury, is being developed for housing, and this population will soon be extinct. Geissorhiza mal- mesburiensis may occur elsewhere in the Mal- mesbury district, but collecting in this heavily agricultural area has so far failed to produce other records of the species. It must be regarded as seriously endangered and if not extinct, then on the verge of extinction. GOLDBLATT — GEISSORHIZA FIGURE 27. Morphology and distribution of Geis- sorhiza malmesburiensis. Habit life size; separate flow- er x2 (Goldblatt 6282, Malmesbury Commonage). Specimens examined. SOUTH AFRICA. CAPE: 3318 (Cape Town) Malmesbury commonage (BC), Lewis 3633 (SAM), Goldblatt 6282 (MO); hill near Malmes- bury, 300 m, Schlechter 1654 (B, BOL, BR, L, MO, PRE, 2). 14. Geissorhiza geminata E. Meyer ex Baker, Handb. Irid. 159. 1892, Fl. Cap. 6: 76. 1896; Foster, Contr. Gray Herb. 135: 63. 1941. TYPE: South Africa. Cape: between Slangen- K, designated here; isolectotypes, B, BM, E, G, K, L, MO, P, S) [Foster (1941) failed to indicate which of the sheets at K was the *type" hence his apparent lectotypification cannot be accepted]. FIGURE 28. Geissorhiza geminata Drége ms.; E. Meyer, Flora 26 (Beigabe): 119-187. 1844, nom. nud. Plants (12-)16-30 cm high. Corm globose, more or less symmetric, obliquely flattened be- low, tunics concentric, light brown, fragmenting irregularly into segments, drawn into short bris- tles above. Cataphyll membranous. Leaves 3-5, 328 FIGURE 28. Morphology and distribution of Geiss- orhiza geminata. Habit and corm x0.5; intact and opened flower and floral bracts life size; leaf section much enlarged (Goldblatt 6981, flats near Worcester). the lower two or three basal, shorter than the stem, 1-2 mm wide, blade plane when dry, pos- sibly inflated and terete when live in some pop- ulations, the upper leaves cauline and partly sheathing, uppermost bract-like, often vestigial. Stem erect, bearing leaves below and leafy bracts above, often branched and then dichotomously. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Spike 1—3-flowered; bracts herbaceous, hyaline or reddish along the margins, 6-8 mm long, the inner about as long, or longer than the outer. Flower white (to cream), stellate, sometimes pink on the reverse of the tepals; perianth tube to 8 mm long, reaching the apex of the bracts or long- er; tepals to 14 mm long, ca. 6 mm wide. Fila- ments to 4 mm long, erect; anthers 5 mm long. Ovary 1.5-3 mm long, style dividing at the apex ofthe anthers, branches ca. 4 mm long, recurved. Capsule unknown. Chromosome number, un- known Flowering time. September to November. Distribution. Marshes and vleis, in sandy areas from Brandvlei north to the Cold Bokke- veld. Figure 28. Geissorhiza geminata is a semi-aquatic species, typically found in seasonally marshy areas, sometimes in standing water. It occurs in the Worcester and Ceres districts, north into the Cold Bokkeveld. It is often confused with the super- ficially similar G. brehmii (syn. G. teretifolia) which also grows in aquatic habitats and has fair- ly large white flowers. The two species are, how- ever, quite unrelated. Geissorhiza brehmii has imbricate corm tunics typical of subgenus Geis- sorhiza, while G. geminata has the concentric tunics of subgenus Weihea. Geissorhiza gemi- nata is most probably allied to G. setacea and G. ornithogaloides (section Weihea), and it has the flat leaf lamina characteristic of the section. It can be distinguished by its large white flower, long perianth tube usually exserted from the bracts and also by its characteristic pattern of nearly dichotomous branching in the upper stem. This branching pattern is so distinctive that most specimens of G. geminata can be recognized at once from this character alone. The name Geissorhiza geminata was first used by Drége on his collections of the species and the name was subsequently published by Meyer (1843) in this treatise on Drége's travels and col- lections. A valid description was not published until 1892 by J. G. Baker. Earlier, Vahl (1805) described an Ixia geminata, which is according to Lewis (1962), synonymous with G. geminata. It is not, however, the basionym for G. geminata which was treated as a new name in Geissorhiza. The identity of Vahl's Ixia geminata is unknown to me. The type specimen in the Vahl Herbarium has been lost, although a sheet, now bearing only the faint impression of a plant, still exists. 1985] Specimens examined. SOUTH AFRICA. CAPE: 3219 (Wuppertal) Cold Bokkeveld, vlei at Leeurivier bridge Elands Kloof, Leipoldt 3036 (BOL), 3038 (BOL). 3319 (Worcester) near Lakenvlei (BC), Phillips 2077 (SAM); Brandvlei (CB), Lewis s.n. (BOL 20414, K); near Worcester, in water, L. Bolus s.n. (BOL 19180), Goldblatt 6981 (E, K, MO, NBG, PRE, S, US, WAG); Moordkuil, Worcester (CD), Lewis 2896 (SAM); Scherpenheuwel vlei (DA), Barker 7507 (NBG). WITHOUT PRECISE LOCALITY: between Slangenheuwel, Franschehoek and Donkerhoek, Drége s.n. “G. gemi- nata EM. aœ” (B, BM, E, G, K, L, MO, P, S); Cape of Good Hope, Heatley 6394 (BM). 15. Geissorhiza ovalifolia Foster, Contr. Gray Herb. 135: 58. 1941. TYPE: South Africa. Cape: ‘‘Drakensteinsbergen, 4,000-5,000 ft., Oct.," Drége s.n. (holotype, B; isotypes, G, S) Plants small, 3-7(—9) cm high. Corm ovoid, 2 mm diam., tunics concentric, papery and soft. Cataphyll evidently dark, membranous or be- coming fibrous. Leaves 4-5, the lower three to four basal and prostrate, the uppermost cauline and bract-like, lanceolate-oblong, occasionally ovate, acute, 2-5 mm wide, 1—4 cm long, bearing a cormlet in the axil of one or more basal or rarely cauline leaf. Stern often prostrate near base, then erect, with a leaf bract in lower third to mid- line, rarely 1-branched. Spike 1—4(—6)-flowered, flexuose; bracts herbaceous, becoming membra- nous above, 4-7 mm long, the inner usually slightly shorter than the outer. Flowers stellate, white, yellow in the throat; perianth tube 2-3 mm long, widening from the base; tepals ovate, 6-8 mm long, to 3.5 mm wide. Filaments 2-3 mm long; anthers 1.5-3 mm. Ovary ca. 2 mm long, style dividing at the apex ofthe anthers, branches 1.2-2 mm long, recurved. Capsules ovoid, 3-5 mm long. Chromosome number, unknown. Flowering time. Late September to mid-No- vember. Distribution. Mountains of the western Cape from French Hoek north to Visgat and the Twen- ty Four Rivers Mountains, usually in shaded sites in damp moss. Figure 29. Geissorhiza ovalifolia is a poorly known mountain species of damp shady sites in mossy banks, cracks in cliffs and rocky ledges, or under boulders. It is often confused with the mor widespread and common G. bolusii which occurs in similar habitats in the Cape mountains. Geis- oO GOLDBLATT — GEISSORHIZA = NS c KS SSS METERS [] mm TEM Z... ULL 18 19 20 n 1 n 1 FIGURE 29. Distribution of Geissorhiza ovalifolia. enrhi7n hifnlin1 3751 41 J y sexually fertile, and capsules develop rapidly after blooming. Unfor- tunately, few mature plants are known. It also produces a cormlet in one or two basal or rarely cauline leaf axils which gives it a characteristic appearance. In contrast, G. bolusii is a triploid, of cauline leaf bracts, one being characteristic of G. ovalifolia while there are usually several in G. bolusii. This distinction does not always hold, and some specimens of G. bolusii may also have a single leaf bract on the stem. In some specimens, the slightly darker yellow color in the perianth tube and throat gradually intensifies with age, and in the early nineteenth century collections of Drége, and Ecklon and Zey- her, it has turned dark blue. This has been the cause of some taxonomic confusion and Foster (1941) believed the dark center to be a significant taxonomic feature of G. ovalifolia. Specimens examined. SOUTH AFRICA. CAPE: 3319 (Worcester) Twenty Four Rivers Mts., damp shady spots (AA), Esterhuysen 16164 (BOL, PRE); Visgat, Stokoe s.n. (SAM 63131); Groot Winterhoek farm, mts. behind Porterville, Boucher 1902 (PRE Porterville, Edwards s.n. y; Wit i Ridge above De Eiker, Esterhuysen 336 OL, K, MO, PRE, S, US); mts. W of Gydo Pass, Hutchinson 330 ANNALS OF THE MISSOURI BOTANICAL GARDEN “> os NS nm | x O i AC 17 a i EZ foy S s SNS NN e" 72777 as: 277 FiGuRE 30. Morphology and distribution of Geissorhiza bolusii with the range of the closely allied, sexually fertile G. ovalifolia outlined. Flowering plant x0.5; sterile plant and flower life size (Esterhuysen 352814, ). Simonsberg 1013 (K); Hartebeeskloof, S slopes of Vaalboskloofberg (AB), Oliver 5162 (STE); Worcester, "beim waterfall" (AC), Ecklon & Zeyher Irid. 223 (B, FI, G, LD, MO, S); Hex River Mts., Milner Ridge Peak (AD), Ester- huysen 9341 (BOL); Michells Pass, Esterhuysen 6202 (BOL), Compton 11934 (NBG); Ceres side of Michells Pass, Thorne s.n. (MO, SAM 51236); rugged slopes above Merweda, E of Eselfontein, Esterhuysen 32678 (BOL, K, MO, PRE); French Hoek Forest Reserve near Bushmans Castle (CC), Salter 6869 (BOL, SAM); Langeberg, above Berg-en-dal farm, ca. 2,000 ft., steep S facing slopes and cliffs (DB), Esterhuysen 35683 (K, MO, NBG, PRE, WAG). 3419( berg, Somerset West (BB), Dummer 590 (E). WITHOUT PRECISE LOCALITY: Witsenberg, Tygerkloof, Zeyher s.n. (SAM 20889), Drége s.n. (B). 16. Geissorhiza bolusii Baker, Handb. Irid. 158. 1892, Fl. Cap 6: 75. 1896; Foster, Contr. Gray Herb. 135: 57-58. 1941; Lewis, FI. Cape Penins. 254. 1950. TYPE: South Africa. Cape: Du Toits Kloof, 2,300 ft., H. Bolus s.n. [lectotype, BOL 5247 in K, effectively designated by Foster (1941); isolectotype, BOL]. FIGURE 30. Geissorhiza rupestris Schltr., Bot. Jahrb. Syst. 27: 98. 99; Foster, Contr. Gray Herb. 135: 57. 1941. TYPE: South Africa. Cape: mts. above Bains Kloof, 2,400 ft., Schlechter 9167 [lectotype, B, designat- ed by Foster (1941); isolectotypes, B, BM, BOL , G, H, K, L, MO, PH, PRE, Z]. Geissorhiza dregei Baker, Handb. Irid. 158. 1892, FI. Cap. 6: 74. 1896; Foster, Contr. Gray Herb. 135: 56. 1941. TYPE: South Africa. Cape: Paarl Mts., Drége s.n. “G. secunda 8 Ker a" [lectotype, K, effectively designated by Foster (1941); isolecto- types, BM, E, G, K, L, MO]; without locality, Thunberg s.n. (syntype, Herb. Burman G = G. parva); Olifantshoek, Ecklon & Zeyher Irid. 221 ; — G. parva). Geissorhiza tabularis Ecklon ms. (Ecklon 311). Plants 3-8(-11) cm high. Corm more or less ellipsoid, 2-4 mm diam. at widest, tunics pale, concentric, brittle-papery, notched below into sections, and drawn into points above. Cataphyll transparent, membranous. Leaves 2—5, basal, en- siform to lanceolate, acute, usually prostrate, blade flat, the stem also bearing leafy bracts. Stern usually prostrate at the base, erect above, occa- sionally branched, with a few to several leafy bracts, bearing a small cormlet in the axil of each leaf and bract. Spike 2-8-flowered, or flowers all bracts green, 3-6(-10) mm long, the inner about as long or longer than the outer in those sub- tending flowers, much smaller where flowers 1985] aborted, a cormlet produced in the axil of each sterile bract and, after flowering, in the axils of the floral bracts. Flowers stellate, white (or not produced on some plants); perianth tube infun- dibuliform, 3-4 cm long, reaching apex, or slight- ly exserted from bracts; tepals 7-10 mm long, spreading, ovate, 2-3 mm wide. Filaments 3-4 mm long; anthers 2-3 mm long. Ovary 3-4 mm long, style dividing near the apex of the anthers, branches ca. 1.5 mm long, recurved. Capsules and seeds never developed. Chromosome num- ber, 2n = 39 (Goldblatt 458, 515, 6444, Gold- blatt s.n. no voucher, High Noon Mts.; Ester- huysen 352814). Flowering time. October to December (to anuary). Distribution. Mountains along the western Cape coast and interior from the Cape Peninsula to the Cedarberg, shady south-facing slopes, rock cracks, often in damp sites with moss. Figure 30 Geissorhiza bolusii is a poorly collected, yet widespread species, common in suitable habitats in the mountains of the western Cape. It favors cool damp sites in shallow soil under rocks or in cracks of south-facing cliffs or along watercours- es, and it is often associated with moss. The species is unusual, being triploid, 27 — 39 (counts from five separate populations from different parts of its range), and it never reproduces by seed. Instead a small cormlet develops in the axil ofeach leaf and stem bract. Some flowering stems produce no flowers, but a cormlet in the axil of each flower bract. Other stems produce small white flowers, but no capsules, and a cormlet later develops in place of each aborted ovary. The presence of these two kinds of stems, flow- ering and vegetative, is one characteristic Mature of G. bolusii. of both kinds, including the type collections of G. bolusii and G. rupestris, the latter reduced to synonymy here. One of the three type collections of G. dregei, Drége s.n. from Paarl Mountain, chosen here as lectotype for this species, also exhibits this characteristic. The two other type Ld consist of rather poor specimens of G. pa hus G. dregei, recognized as distinct by Foster n is likewise reduced to synon- ymy in G. bolus Geissorhiza n is probably most closely related to G. ovalifolia which grows in similar habitats and also has small, white flowers. Geis- sorhiza ovalifolia usually has none or only one cauline leaf or bract, and produces one or few GOLDBLATT— GEISSORHIZA 331 cormlets in its leaf axils. In G. ovalifolia, fruit development is evidently fairly rapid and im- mature capsules can be seen in some collections in which fading blooms are present at the ends of spikes. pecimens examined. SOUTH AFRICA. CAPE: 3218 (Clanwilliam) Piketberg (D-), Barker 7856 (NBG); Gryskop, Piketberg, 3,500 ft. (DC), Pillans 7400 (BOL); Zebrakop, Piketberg, 3,000-4,000 ft., Esterhuysen 35805 (MO). 3219 (Wuppertal) N Cedarberg at Koupoort (AA), Esterhuysen 12188 (BOL); Cedarberg, Middelberg vlakte, under boulders (AC), Goldblatt 5135 (K, MO, PRE, S); Elandskloof, rock ledges (CA), Bond 677 (NBG); Hexberg, Cold Bokkeveld Mts., Goldblatt 7 114 (K, MO); Warm Baths, Olifants River Valley, Edwards S. n. (BOL 14430); Bo-Boskloof, Cold Bokkeveld (CC- CD), Rourke 651 (NBG). 3318 (Cape Town) Table Mt., S aspect (CD), H. Bolus 4617 (BOL); Table Mt., Ecklon 311 (G, K, M, PRE, S), Marloth 8041 (PRE); iras DUE Dum- mer 858 (E); Devils Peak, Wolley 8 (BOL); Cecilia, Barker 20181 (NBG); Lions Ea iak places 2, 900 ft. (DD), Kerfoot 6097 (NBG); Simonsberg, kloof r Helshoogte, Esterhuysen 35281A (MO). 319 (Worcester) Groot Winterhoek, moist rocks T Marloth 2329 (PRE); Witsenberg (AC), Zeyher 3958 (SAM); shady damp rocks, upper Tulbagh Wa- terfall, Burgers 2586 (STE); Milner Peak, Hex River Mts. (AD), Esterhuysen 14240 (BOL); Hex River val- ley (BC), Davidson 116 (SAM), 53 (SAM); Bains Kloof (CA), Schlechter 9167 (B, BM, BOL, E, G, H, K, L, MO, PH, PRE, Z), Esterhuysen 32723 (BOL), Gold- blatt 458 E dg 380 (PRE), L. Bolus s.n. ie Barker 4250 G, W); Cossacks, Slanghoek Mts., 4,000 ft., Eoin 35823 (C, K, WAG); a Slanghoek Mts., Esterhuysen 9454 ( Peak, damp bank (CB), Esterhuysen 18190 (BOL); French ipiis (CC), Barker 4910 (BOL); Jonaskop, Riviersonderend Mts., under rock overhang (DC), Goldblatt 6444 JA PRE). 3418 d slopes of Constantiaberg (AB), M); scs Ait. , Compton 14171 (NBG), us dr 1933 (K); near M corner, Lewis 1488 (NBG). 17. Geissorhiza parva Baker, Kew Bull. 1906: 26. 1906; Foster, Contr. Gray Herb. 135: 60. 1941. TYPE: South Africa. Cape: Houw Hoek, Penther 723 (holotype, K). FIGURE 31 Ixia [iiie f., Suppl. Pl. 92. 1782; Thunb., Diss. de Ixia no. 24. 1783, pro parte excl. Thunber, rg s.n. “Tia excisa o” [lectotype of I. excisa L. f. (Gold- blatt, 1983) — G. ovata (Burm. f.) Asch. & Graeb.]; Thunberg s.n. “Ixia excisa 8" (syntype, UPS = G. parva). 332 METERS LJ ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 21 7 1811381521 fy Z 4 M) >) c SZ SASS NN SS c. Sy - Hinr A g "P "s , i u MSN TITTY FIGURE 31. life size; opened flower x2 (Goldblatt 5897, Caledon Zwartberg). Geissorhiza dregei Baker, Handb. Irid. 158. 1892, Fl. Cap. 6: 74. 1896, pro parte excl. Drége s.n. “G. secunda B Ker a” (lectotype of G. dregei = G. bo- lusii Baker). Geissorhiza tabularis Ecklon ms. [Ecklon & Zeyher Irid. 222 (90.9)]. Plants small, 4—7(-12) cm high. Corm ovoid- conic, 3-5 mm diam., asymmetric, obliquely flattened towards base, the flat part extending downwards shortly, tunics dark brown, concen- tric, the layers firm textured, fragmenting irreg- ularly. Cataphyll membranous brown, usually persistent and accumulating around the base. Leaves 3-4, all basal or the uppermost cauline, 1.5-2(-4.5) cm long, prostrate or inclined, much shorter than the stem, ensiform to oblong, rarely linear, occasionally ovate, usually minutely red- dotted or streaked, acute or obtuse, margins oc- casionally ciliate to papillose towards the apex. Stem smooth, erect above, often prostrate for a he base, simple or rarely branched, flexed above the cauline leaf. Spike (1-)2-6-flowered; bracts herbaceous, usually red on the margins, 4—6(-8) mm long, the inner slightly shorter than the outer. Flower stellate, creamy white to pale yellow; perianth tube 3-4 31. Morphology and distribution of Geissorhiza parva. Habit x0.5; corm, flower, and gynoecium mm long, included or just 1 from the bracts, gradually widening from the base; tepals extend- ed horizontally, more or less ovate, 6-8 mm long, .5-3.5 mm wide. Filaments ca. 2 mm long, erect; anthers 2-2.5 mm long, pollen yellow. Ovary ca. 2 mm long, style dividing at the apex of the an- thers, branches 1.5-2 mm long, recurved. Cap- sule globose to oblong, 3-7 mm long. Chromo- some number, unknown. Flowering time. Mid-August to October (to early December at high altitudes). Distribution. Sandy and often very thin mountain soils in the southwestern Cape, ex- tending from the Cedarberg south to the Lang- eberg near Swellendam; mainly montane at mid- dle to upper elevations but occasionally at low altitudes near the coast in the Caledon district. Figure 31. Geissorhiza parva is clearly closely allied to G. ovata, a species similar in habit with short pros- semi-prostrate | d a long perianth tube, but larger in all features and having par- ticularly long-tubed white flowers, marked red at the base of the tepals and on the reverse of the tepals and tube. The difference in size, flower 1985] color, and length of the perianth tube in the two species makes confusion unlikely. Geissorhiza parva sometimes grows in open soil, occasionally together with G. ovata, but more often on thin soil on wet rocky pavement or on rocks in damp moss. It has a wide range, extending from the Cedarberg southwards to the Caledon district and east to the Langeberg near Swellendam. Popu- lations occurring north of the Hex River Moun- tains can usually be distinguished by having four leaves, in contrast to the southern populations which normally have only three. The difference is not without exception, e.g., Stokoe s.n. from the Riviersonderend Mountains in the south, and Esterhuysen 35719 from the Hex River Moun- tains to the north, consist of plants with either three or four leaves. Some northern populations of Geissorhiza parva can be difficult to distinguish from the re- lated G. ovalifolia. This species has white flow- ers, soft-textured leaves, and usually an axillary cormlet near ground level. The two species can usually be separated by a combination of these characters. Specimens from the Commonage below the Zwartberg at Caledon, Guthrie s.n., are probably hybrid with Geissorhiza ovata. Some individuals of this collection have very large flowers with the reverse of the outer tepals flushed red. The flow- ers have tepals to 13 mm long and a tube as long as 6 mm, a size intermediate between G. parva and G. ovata. Specimens examined. SOUTH AFRICA. CAPE: 3219 (Wuppertal) Cedarberg, Middelberg (AC), Goldblatt 5137 (MO, NBG); Elands Kloof (CA), Lewis s.n. (BOL 31893), Barker 3087 (NBG); Cold Bokkeveld, 33.5 km E of Citrusdal, Goldblatt 4203 (K, MO); Olifants River Valley, Grootfontein above the Toorgat (CC), Oliver 3973 (K, PRE, STE); Hexberg, Cold Bokkeveld Mts., 4,000 ft., Goldblatt 7115 (MO, PRE); Cold Bokkeveld, Wabooms River-Ysterfontein (CD), Lewis 2644 (SAM); E side of Bokkeveld Sneeukop, after fire, in wet sand, rcester) Visgat ( 63130); Michells Pass (AD), ep ied 6145 (BOL); (BOL 17233); Skurf- ); M OS- , shallow muddy soil (BC), Pn. 35719 (BOL, K, MO, PRE,. S). 3320 (Montagu) near base of Crown Mt., Swellen- dam (CD), Wurts 282 (NBG). ottentots Holland, Pus a 131 (BOL n 550 (NBG); Eg Fo D), Bouche 1942 (PRE, STE); Platteberg, Stokoe s.n. (SAM 61723). GOLDBLATT— GEISSORHIZA 333 3419 (Caledon) flats E of Viljoens Pass (AA), Davis s.n. (SAM 61726), Stokoe s.n. (SAM 61724); Viljoens Pass, Stokoe s.n. (SAM 49600); Lebanon State Forest, Jakkalsrivier Catchment II, moist path in peaty soil, Haynes 1243 (STE); Houw Hoek Pass, near Bot River, van Niekerk 785 (BOL); Caledon Zwartberg (AB), Goldblatt 5897 (K, MO, NBG); Zwartberg near Cale- don, Galpin 4675 (K, PRE); Caledon, commonage be- low Zwartberg, Guthrie 16916 (BOL); Highlands For- est Reserve, wet flats (AC), Goldblatt 4779A (MO); Hawston, sand hills near lagoon, Walgate 1087 (BOL); Vogelgat, cleared path at 1,500 ft. (AD), Goldblatt 5348 (MO); Vogelgat, roads end, 410 m, Williams 2859 (NBG); 8 mi. from Stanford on road to Elim, Gillett 4499 (BOL, BR, MO); Riviersonderend Mts. (BA-BB), Stokoe s.n. (SAM 63482); Swellendam, “am Ufer des Riviersondereinde bei Appelskraal" (BB), Ecklon & Zeyher Irid. 222 (90.9) (B, FI, G, LD, MO); Olifants- hoek, Ecklon & Zeyher Irid. 221 (K, SAM); 2 mi. N x Papiesvlei (BC), Acocks 22868 (K, PRE); Salmons- m Nature Reserve, Goldblatt 437 (BOL); Sandys Glen (BD). Goldblatt 4849 (K, MO, NBG); flats near Baard- scheerdersbos (DA), Lewis 6007 (NBG). 420 (Bredasdorp) Potteberg, De Hoop (BC), Bur- gers 2247 d STE). T PRECISE LOCALITY: Cape of Good Hope, Herb. Burman s.n. 18. Geissorhiza ovata (Burm. f.) Asch. & Graeb., i . 1906; Foster, man s.n. (holotype, Herb. Burman G). FIGURE 32. Ixia excisa L. f., Suppl. Pl. 92. 1782, Thunb., Diss. de ia no. 24: 151. 1783. Geissorhiza excisa (L. f.) Ker, Ann. Bot. (Kónig & Sims) 1: 224. 1804; Bak- er, Handb. Irid. 159. 1892, Fl. Cap. 6: 76. 1896. TYPE: South Africa. Cape: exact locality unknown, Thunberg s. " M excisa a’ ` [lectotype, uid Thunb. 951 y Brown n slectonnes, S* *Herb. Montin, d “Herb, " “Herb. Swartz’); “Ixia excisa B" (syn- Herb. Thunb. 952 UPS). Weihea excisa (Burm. f.) Ecklon, Topogr. Verz. Pflan- zensamml. Ecklon 22. 1827, comb. illeg. gen. non. val. pev Plants 6-15 cm high. Corm asymmetric, ovoid, obliquely flattened below, the flat part extending shortly downwards, tunics light to dark brown, concentric, drawn into points above. Cataphyll ry, brown, papyraceous. Leaves 2-3, the upper often bract-like and inserted on upper part of the stem, the lower two basal, prostrate or inclined, much shorter than stem (1—)1.5-3(—6) cm long, 5-12(-14) mm wide, ovate-oblong, acute or ob- tuse, the margins usually ciliate, sometimes the blade also ciliate-pubescent on the veins (rarely 334 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 X E c VAM S Wes “| MI US on S Z< 2 Tana — a = SS ASE S 2 X> ae, Se TOY 6 SLIEP: =>») (C= re d — L FIGURE 57 Morphology and distribution of Geissorhiza scillaris. Habit x 0.5; intact flower life size; partial ). flower and gynoecium x2; leaf section much enlarged (Esterhuysen 35495, Zebrakop, Piketberg sheets, it differs from G. juncea in having only one free leaf instead of two, in having a second basal leaf sheathing most of the stem instead of a short third leaf only partly sheathing, and in having a different type of corm. Lewis could have added that G. scillaris is also distinctive in con- sistently having a short, membranous and sheathing bract on the upper part of the stem, a feature found rarely in G. juncea. The fact that G. scillaris was described, albeit from mixed ma- terial, as early as 1783 by Thunberg, as Ixia scil- laris (an illegitimate homonym for 7. scillaris L.) has largely been overlooked. Typification of Thunberg's illegitimate homonym, J. scillaris, is complex owing to the fact that the type material consists of three different taxa. The arguments leading to the selection of a lectotype have been dealt with in a separate paper (Goldblatt, 1983). The subsequent history of Ixia scillaris is com- plex. The species was renamed T. ramosa by Ker (1802c) (who must have chosen the epithet from the protologue and not from the type material which he did not see). The name did not get taken into general use, and Ker (1827) later placed 7. 386 ramosa in the synonymy of Geissorhiza imbri- cata. Roemer and Schultes (1817) also renamed I. scillaris, calling it I. phalangioides, but the epithet is illegitimate and superfluous for /. ra- mosa Ker, cited in synonymy. Schultes (1822) distinguished /. scillaris from I. juncea as did Dietrich (1833), who transferred J. scillaris to Geissorhiza, but few other authors have. Baker (1892, 1896), in his important treatments of Iri- ceae, ignored the species, and Foster (1941) placed /. scillaris Thunb. in the synonymy of G. Juncea. Foster and probably Baker did not see Thunberg's collections. Brown, who examined the Thunberg Herbarium, also failed to distin- guish G. juncea from G. scillaris. Foster actually recognized the species here called Geissorhiza scillaris as G. juncea var. pal- laine pei ) Foster without realizing that the type erial of G. scillaris, at least in part, une ne variety. Geissorhiza juncea var. pal- lidiflora, based on G. pallidiflora Schltr., is typ- ified by Schlechter 9088, a collection that ap- pears to differ in no significant way from G Juncea. The type collection of G. pallidiflora is, however, interesting in one respect, in that of the many plants of the gathering, two at least have a membranous stem bract, but this is probably coincidence and not indicative of any relation- ship with G. scillaris. It is difficult to place G. scillaris with certainty to subgenus, but the limited number of known corms have tunics of the imbricate type, though unlike the typical tunics of subgenus Geissorhiza, species of which have heavy black tunics. The brown corm tunics of subgenus G. scillaris tend to accumulate more than is usual in subgenus Geissorhiza and over the years become rather distorted and broken. The species is here as- signed to the ditypic section Intermedia of sub- genus Geissorhiza, together with G. similis, which has corms with similar tunics. The two species are otherwise rather different and although prob- ably not closely allied, they appear to belong to a single, perhaps very old line. Geissorhiza scillaris is a fairly common species in the mountains of the western Cape, and it extends north from Shaws Pass and the Groen- land Mountains at Elgin to Pakhuis Pass in the northern Cedarberg. It usually grows in open rocky places, in sandstone-derived soil, and usu- ally at altitudes above 800 m. It may occasionally be found in damp places, but is clearly well adapted to dry stony situations. Most popula- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 tions consist of plants with white flowers, but in the Cold Bokkeveld and Cedarberg there is an attractive blue-flowered form in which the flower center is white (Goldblatt 5072, Esterhuysen 3013, 20507, 13111) cimens examined. SOUTH AFRICA. CAPE: 3218 2 ek Mts. above The Rest, Pikeniers Kloof (DB), Gillett 3740 (BOL); c eritis Ester- huysen 35495 (K, MO, NBG, PRE, S Meri Versveld Pass, Piketberg (DC), Compton 10991 (NBG 219 (W , STE); Top of AA Pass, Persson road, Goldblatt 2752 (MO, WAG); Middelberg de ndskloof, Hafstrom Acocks 299 (PR i S); Crystal Pool hut, Pocock 6 (STE); Sneeuberg, summit, " Taylor 6 152 (PR S rocky i ed 2050 7 (BOL. River farm y flats, Goldblatt 567 0 Mos top of (N " om (K, r (CD), Lewis 2504 (BOL, K); Porterville Mts., NW end of Zuurvlakte, Thompson 1476 (STE). 18 (Cape a Klein Drakenstein (DD), Du- Plessis s.n. (STE 19 (W orcester) Visgat (AA), Stokoe s.n. (SAM 63132); Winterhoe 11 senberg, near Steenda Peak, 4,000 ft., Michells Pas (PRE, STE); flats NW of Prince Alfreds Hamlet, Oliver 5016 (K, MO, PRE, STE); Mts. in Hex River Valley (BC), Tyson 722 (BOL, SAM); Wellington (CA), Grant 2391 (BOL, BR, MO); Wolwehoek Forest Reserve, Bains Kloof, Barker 4336 (NBG); Bains Kloof, Comp- ton 18266 (NBG); Mias Poort, above Du Toits Kloof, souls ee 35517 (BOL, MO, US, WAG); slopes at Du Toits Kloof, Esterhuysen 18912 (BOL, PRE); Moins Peak, Du Toits Kloof, Stokoe s.n. (SAM 59845); Breede River flats near Worcester (CB); Goldblatt 6980 (MO, P 3419 (Caledon) Vijoens Pass (AA), Werdemann & Oberdieck 395 (B, PRE); stony flats S of Villiersdorp at Theewater farms turnoff (AB), Goldblatt 4014 (MO, PRE); hillside N of Theewater Kloof, above dam, Gold- blatt 4734 (MO); top of Shaws Pass, recently burned (AD), Marsh 1432 (PRE, STE). 1985] THOUT PRECISE LOCALITY: Thungberg s.n. (“Ixia scillaris a”) n Thunb. 992 in part, S, UPS; Herb. Burman s.n., SECTION GEISSORHIZA 9. Section Geissorhiza. TYPE: G. obtusata Ker. Geissorhiza section Rochea wasa. J. Linn. Soc., Bot. 1878, nom. illeg. superfl. — subsection V. entricosae Foster, Contr. Gray erb. 135: 27. 1941. TYPE: G. imbricata (de la Roche) Ker, nom. illeg. s superfl. Corm tunics hard and woody, stems smooth, leaves with the margins and midrib and often other veins strongly thickened, ensiform to lin- ear, or terete, bracts herbaceous below, becoming dry above, prowess stellate to hypecratentorm, eclinate, the tube short or reaching to the apex of the bracts, stamens equal or unequal. Species. 10. 50. Geissorhiza imbricata (de la Roche) Ker, Ann. Bot. (Kónig & Sims) 1: 224. 1804; Bak- er, Handb. Irid. 156. 1892, Fl. Cap. 6: 72. 1896; Foster, Contr. Gray Herb. 135: 37. 1941, pro parte excl. var. brehmii; Lewis, Fl. Cape Penins. 255. 1950. Ixia imbricata de la Roche, Descr. Pl. Aliq. Nov. 17. 1766. TYPE: South Africa. Cape: Cape Peninsula, Lakeside, Salter 7034 [neotype, BOL, des- ignated by Goldblatt & Barnard (1970)]. FIGURE 58. (For additional synonyms see under the subspecies). Plants 6-25(-30) cm high. Corm 5-10 mm diam., globose, tunics blackish, imbricate, the outer layers accumulating above, and regularly notched below into segments. Cataphyll single, membranous, often lacking in dry material Leaves 3, reaching from about middle to apex of spike, sometimes longer, ribbed with (2-)4 to several longitudinal grooves running the length of the leaf, the lower two basal, (linear to) ensi- form, (1-)2-6(-10) mm wide, the third leaf larg- est, inserted on The SEHE near the base or in the low for half its length, the ‘sheath strongly ribbed and inflated. Stem erect, simple or 1-branched, flexed above the leaf sheath. Spike (2-)6—8(—14)-flowered, flexed at the base and flexuose; bracts herbaceous below, often becoming dry and membranous above at flow- GOLDBLATT — GEISSORHIZA 387 ering time, an beset mm long, sometimes red above, the r narrower than the outer. Flower stellate a pe cupped, white to cream or pale yellow, usually red on the reverse of the outer tepals, sometimes dark in the center; perianth tube 2-8 mm long, enclosed to slightly exserted from the bracts; tepals (9—)1 1—25(—30) mm long, obovate, (4.5-)5.5-12 mm wide, ob- tuse. Filaments equal, (354-8(-10) mm long. anthers (3-)4-7 mm long, yellow. Ovary 2-3.5 mm long, style dividing near the apex of the anthers, branches - mm long, recurved. Cap- sule oblong-globose, to 10 mm long. Chromo- some number, 2n — = (Goldblatt 165). Flowering time. (August to) September to October. Distribution. Southwestern Cape, from Bre- dasdorp north to the Olifants River Valley; mainly on damp poorly drained flats. Figure 58. As treated here, Geissorhiza imbricata is one of the more variable species in the genus. It in- cludes no less than three species recognized by Foster (1941) in his monograph of the genus: G. wrightii, G. rubicunda, and G. bicolor (including var. macowanii), as well as G. sulphurea var. arenicola. The main forms or races of Geissorhiza im- bricata are: 1. A short-tubed (2-4 mm) form, usually fairly low growing, with small, white to cream flowers that was once common on the Cape Peninsula and adjacent Cape Flats (G. sabulosa, G. imbri- cata sensu Baker and sensu Lewis). 2. A very robust form with a longer tube (3.5— 6 mm) and with medium-sized flowers and sometimes very broad leaves (G. wrightii; G. im- bricata, at least as to type). 3. A medium- to long-tubed form (5-8 mm) with large cream to pale yellow flowers and nar- row to moderately broad leaves (G. rubicunda, G. obtusata). 4. A form similar to the previous but with a dark throat and often dark tube to the flower (G. bicolor). It would seem convenient to give taxonomic recognition to all four forms or races as did Foster, but the existence of many intermediates, col- lected since Foster's monograph was publishe makes a revised treatment necessary. Lewis, in her treatment of Geissorhiza in the ‘‘Flora of the Cape Peninsula" (Lewis, 1950) maintained both the short-tubed form (as G. imbricata) and the 388 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 72 Í aV 2 NUE y SS DR CAA NS Y J N NN Y G. imbricata [i Q subsp. imbricata 2 DN. ANSA A Bb subsp. bicolor | 9} x = `< "nm apis m e die s . Big € e Tuas == < [|]. e | sn e 9 | Ze ° m" oe III MMC CE SA 18 19 20 21 SH ALE FIGURE 58. Morphology of Geissorhiza imbricata subspecies imbricata and the distribution of the two subspecies of G. imbricata. Habit Kirstenbosch). rons and iesu G. wrightii, the only two I nd most specimens on the Peninsula can readily be assigned to one of these. However, to the east in the Caledon and Bre- dasdorp districts a range of intermediates occur which I find impossible to assign with confidence to either of these taxa. The only solution seems to be to treat these as one taxon, which has in common small- to medium-sized flowers with a short to moderately long perianth tube. This tax- on has a distinct southern distribution from the Cape Peninsula east to Bredasdorp It has also become impossible to continue to regard the cream- to pale yellow-flowered longer x 0.5; flower life size; leaf section much enlarged (Goldblatt s.n. no voucher, tubed forms with self-colored flowers as distinct from those with dark markings in the throat and tube. The difference here seems solely to be one of coloration and there are populations of plants with and without the dark color in the throat and ube. However, at present it does seem useful to formally recognize this larger, cream- to yellow- flowered form with a generally longer perianth tube as a distinct taxon. The presence of inter- mediates between this and the shorter tubed, white-flowered form in the Worcester and Tul- bagh districts that cannot be assigned with con- fidence to either form suggests that subspecific e. 1985] status may be most appropriate. The cream- to yellow-flowered form is thus treated as Geisso- rhiza imbricata subsp. bicolor (Thunb.) Goldbl. KEY TO THE SUBSPECIES OF GEISSORHIZA IMBRICATA la. d white to cream (often red on the re- rse of the outer tepals), without a dark cen- hen dpi dar boe 2-4(—6) mm long; tepals 9 15(-17) mm lo subsp qm Flowers cream Ps pale yellow (sometimes re on the reverse of the outer — occasion- ally with a dark center; perianth tube (4—)5- 8 mm long; tepals (16-)20-30 mm ise T subsp. bicolor = 50A. Subsp. imbricata. FIGURE 58. Ixia scillaris Baar Diss. de /xia no. 14. 1783, pro parte, hom. illeg. non L., excl. lectotype (Gold- blatt, 1982). Geissorhiza sabulosa Klatt, Trans. S. African Philos. Soc. 3:2 1 Africa. Cape: near totype Geissorhiza wrightii Baker 2 Fl. Cap. 9. 1896; Foster, Contr. Gray Herb. 135: 35- 36. 1941; Lewis, Fl. Cape inm 255. 1950. TYPE: is Africa. Cape: Simons Bay, Wright 243 [lectotype, K, designated us Foster (1941: 36); Dorn iei Geissorhiza — var. arenicola Foster, — Gray He 1941. TYPE: South a. Cape: Ceres Road peers Schlechter 3992 (holo- type, B; isotypes, BM, BOL, BR, E, G, H, K, L, MO, PRE, S, Z). Geissorhiza arenaria Ecklon ms. (Ecklon 1597, 313). Plants (6—)10—20(-30) cm high. Leaves (1-)2- 5(-10) mm wide. Spike (2-)6-8(-14)-flowered, flexed at the base and flexuose; bracts (6—)8-11 mm long. Flower stellate, white to cream, usually red on the reverse of the outer tepals; perianth tube 2-4(-6) mm long, enclosed in the bracts; tepals (9-)11-15(-17) mm long, obovate, (4.5-) 5.5-7 mm wide, obtuse. Filaments (3—-)4—5 mm long; anthers (3—)4-5 mm long, yellow. Ovary 2- 3 mm long. Capsule oblong-globose, to 10 mm long. Chromosome number, 2n = 26 (Goldblatt 165) Flowering time. (August to) September to October. Distribution. Cape Peninsula eastwards to the Caledon and Bredasdorp districts, and local in the Worcester district north to near Wolseley. Figure 58. Geissorhiza wrightii was described by Baker in 1876 and although it has remained poorly under- stood, it has been consistently recognized ever GOLDBLATT — GEISSORHIZA 389 since (Baker, 1896; Foster, 1941; Lewis, 1950). Foster actually saw little material of the species and was in some doubt about its status (Foster, 1941: 36) but he realized that it belonged to the “imbricata group." Lewis, who had a consider- able field knowledge of Iridaceae, stated in “Flora of the Cape Peninsula" that it was very like G. imbricata but larger and more robust and with a longer perianth tube (Lewis, 1950). The key distinction she made was that the leaves of G. imbricata were 1-5 mm wide with the blade of the sheathing leaf about as long as the sheath, while the tepals were 9-20 mm long and the tube 1—2 mm long. In contrast, the leaves of G. wright- ii were 5-8 mm wide with the blade ofthe sheath- ing leaf usually much longer than the sheath, while the bie were 16-20 mm long and the tube 3-5 mm long. The leaf blade versus the sheath distinction is than the sheath (Goldblatt 6189, Page s.n.) while several narrow-leaved specimens have a blade much longer than the sheath (Zeyher 1597). Moreover, a few of the specimens determined by Lewis as G. wrightii have leaves less than 5 mm wide. It seems that like many species of the ge- nus, Geissorhiza imbricata can be very variable as regards height; leaf width; and length, number, and size of the flowers. Accordingly, G. wrightii seems better regarded as merely robust G. im- bricata subsp. imbricata, possibly genetically dis- tinct, but equally likely, simply large due to good growing conditions. It is here regarded as taxo- nomically identical with G. imbricata and the less robust G. sabulosa. Geissorhiza imbricata subsp. imbricata has a southern distribution from the Cape Peninsula and Cape Flats to Bredasdorp. To the north, it is replaced by the larger and cream- to yellow- flowered subsp. bicolor which grows in similar wet to semi-marshy places. In the Worcester dis- trict and around Stellenbosch where the two sub- species meet, it is often difficult to assign spec- imens with confidence to either and some populations seem completely intermediate be- tween the two in some or all their characteristics. The differences between the two subspecies are dealt with in more detail under subsp. bicolor. Specimens examined. SOUTH AFRICA. CAPE: 3318 (Cape Town) foot of Table Mt. near Kirstenbosch (CD), H. Bolus 4710 (BM, BOL, K); foot of Table Mt., MacOwan s.n. Herb. Norm. Austr. Afr. 258 (B, BM, G, GH, K); slopes near Blinkwater, Levyns s.n. (CT); 390 nu Bay, MacOwan 2273 (K, S, SAM), Cassidy 287 (CT, SAM), Lavranos 10993 (MO); above Camps Bay, Wolley Dod 2767 (BOL, K), Lewis 1213 (SAM), Bark- er 3850 (NBG); Raapenberg, Mowbray, Guthrie s.n. (CT); Camp Ground, Salter 9640 (BM); Kenilworth, L. Bolus s.n. (BOL 13944); Kenilworth Race Course, Barker 4793 (NBG), 4114 (NBG), Lewis 4793 (SAM), Levyns s.n. (CT 5889); Claremont, Page s.n. (BOL); Wynberg, damp fields, H. Bolus s.n. (BOL 31876); bog- gy peng oe Pillans 10640 (G, MO); Mait- land flat metery, Wolley Dod 2888 (BM, BOL, K); Aneren flats (DD), Strey s.n. (M), Garside 87 (K), 1049 (K). 31 ; (Worcester) us Road (Wolseley) jr Schlechter 8992 (B, BM, BOL, E, G, RE, S, Z); Lael s eg Worcester (CB), joe ia 8199 (BOL); Klein Drakenstein (CC), Du Plessis s.n. (STE 19731) 3418 (Simonstown) Schusterskraal vlei (AB), Barker 3888 (BOL, NBG); marsh at Lakeside Station, Salter 7034 (BOL); Modderdam, Salter 8689 (BOL, CT, K, SAM); Constantiaberg, Compton 15142 (NBG): lower , Hout Bay, Acocks 692 (PRE); Orange Kloof, in swamp, Wolley Dod 3454 (BM, BOL, K); Three Beacons, Bergvliet flats, Purcell s.n. (MO, SAM 93340); Hout Bay, Bond 1227 (NBG), Compton 11295 (NBG); Constantia Nek, under pines, Goldblatt 423 (BOL); Fish Hoek flats, Salter 7397 (SAM 20871), Ecklon 313 (E, K, M, MO, P, S, Z), Moss 4307 (Z), 4309 (BM); Cape Flats near the salt pans, Bowie 1 " kerk 276 (BOL); Faure, Barker 1824 (NBG), 2987 : s Bay, wet soil (BD), Ebersohn 358 (NBG). 3419 (Caledon) flats E of Viljoens Pass (AA), Davis & Stokoe s.n. (S eed at the foot of the lisi Guthrie s.n. (BOL 16919); wet area on Kleinmond road just S of Her- manus turnoff (AC), Goldblatt 2989 (MO, PRE, WAG); Karwyders l, between Bot River and Hawston, Lewis 5289 (NBG); F ernkloof Nature Reserve, Her- a (AD), icula 307 (BR, E, K, MO, WAG); ermanus-Stanford road, wet recently burned ground, Gillen 4166 (BOL), 4177 (BOL, K); kloof SW of Napier — n 6189 (MO, NBG, PRE, S, US, WAG); wet fl skraal (CB), Goldblatt 7098 (MO, NBG, PRE), boggy ground near Elim (DA), Muir 5019 (NBG); near The Poort, Bredasdorp (DB), Barker 4594 (NBG), Acocks 1526 (S). WITHOUT PRECISE LOCALITY: eei ie n. (S "Herb. Montin," Herb. Thunb. 993 UPS), Herb. Burman s.n. (G), Oldenburg 45 (BM), 629 “Ixia virginea” (BM). 50B. Subsp. bicolor (Thunb.) Goldbl., comb. et stat. nov. Ixia bicolor Thunb., Phytogr. Blatt. 1: 3. 1803. Geissorhiza bicolor (Thunb.) N. E. Brown, J. Linn. Soc., Bot. 48: 44. 1928; Foster, Contr. Gray Herb. 135: 32-33. 1941. TYPE: South Africa. Cape: without precise locality, Thunberg s.n. “Ixia bicolor æ” [lec- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 totype, Herb. Thunb. 921 UPS, effectively designated by Brown (1928)]; “Ixia bicolor B8" (syntype, Herb. Thunb. 922 UPS) Geissorhiza ol Sol ex Ker, Bot. Mag. 18: tab. 672. 1803. Geissorhiza imbricata var. obtusata (Ker) Baker, Fl. Cap. 6: 73. 1896. TYPE: South Africa. Cape: cultivated in England (originally collected by Niven) (illustration in Bot. Mag. tab 803). Geissorhiza imbricata var. concolor Baker, Fl. Cap. 72. 6; Foster, Contr. Eid Herb. 135: 38. 1941. TYPE: South Africa. Cape: near Somerset West, H. Bolus 5567 [lectotype, K, designated by Foster (1941); jeep ar BOL]; Somerset West, Eck- lon & Zeyher Irid. 111 (syntypes, B, G). Geissorhiza rubicunda Foster, Contr. Gray Herb. 135: 1. TYPE: South Africa. Cape: Michells Pass, Schlechter 8931 (holotype, B; isotypes, BM, BOL, ,G,H,K, : Geissorhiza bicolor var. macowanii Foster, Contr. Gray 135: 33. 1941. TYPE: South Africa. Cape: near Malme sbury, MacOwan s.n. Herb. Norm. Austr. Afr. 1568 (holotype, G; isotypes, B, G, K, Z). eine aurea Ecklon, Topogr. Verz., Pflan ml. Ecklon 21. 1827 et ms. (Ecklon & 25 Trid. 211). Plants 6-25 cm high. Leaves (1-)3—6(-9) mm wide. Spike 2—6(—9)-flowered, flexuose; bracts 10— 12(-15) mm long. Flower somewhat cupped, cream to pale yellow, often red to purple-flushed -)5-8 mm long, infundibuliform, usually slightly shorter than, or reaching the apex of the bracts, rarely exserted from the bracts; tepals (16-)20-30 mm long, obovate, 10-12 mm wide, obtuse. Filaments 7-10 mm long; anthers 5-7 mm long. Ovary 2.5-3.5 mm long. Capsule ob- long to s. 6-8 mm long. Chromosome number, unknown. Flowering time. September to early October. Distribution. Wet, usually sandy flats from False Bay north to Porterville, and in the Olifants River Valley. Figure 58. Subspecies bicolor includes plants with pure yellow flowers as well as those with a dark center. Populations which have flowers with a dark cen- ter correspond closely to the type collection of Geissorhiza bicolor and are relatively rare, hav- ing been recorded from the Stellenbosch, Paarl, and Malmesbury districts, 1.e., the south central part of the range of the subspecies. As far as I can tell all of the collections differ in no other character but the dark central color of the flower, 1985] from the species known as G. rubicunda Foster. This was poorly known when first described but plants matching the type have now been col- lected at many sites in the Tulbagh and Porter- ville ed as Pas as in the Olifants River Valley to the Collections pom Geissorhiza bicolor and G. rubicunda share the following significant fea- tures: a large pale yellow to cream, actinomor- phic flower with tepals (16—)20-30 mm long; a relatively long perianth tube (4—)5-8 mm long, reaching nearly to the apex of or slightly exceed- bricata typically has smaller white to cream flow- ers, with tepals 12-217 mm long, a short perianth tube 2.5-4(-6) mm long, not reaching the apex of the bracts, and fairly dense spikes of up to 12, but usually 6—8 flowers. The differences in flower size, color and, in particular, length of the peri- anth tube make confusion between most collec- tions of these two subspecies unlikely. Geissorhiza obtusata, Ker's “yellow flowered tile root," is included here in synonymy of subsp. bicolor as it seems to match almost perfectly G. rubicunda. The only type material known is the painting (tab. 672) in the “Botanical Magazine," the flowers of which are today either white (per- haps faded) or yellow in different copies of the volume. Baker (1896) treated this species as a variety of the related G. imbricata. Also included here in subsp. bicolor is G. imbricata var. con- color Baker (syn. G. aurea Ecklon nom. nud.) a distinct form from the southern Cape Flats. It has relatively broad and unusually obtuse leaves and self-colored flowers even lacking the usual red flush on the reverse of the tepals. Geissorhiza bicolor var. macowanii is apparently a name that Foster applied to a form with 3-4 flowers per spike and longer leaves than the two other col- lections which he e to G. bicolor, and it too is placed in synonym Subspecies bicolor is S adis related to the southwestern Cape species G. brehmii (syn. G. ule. and the two can be distinguished only in their vegetative characteristics. Geissorhiza brehmii is a semi-aquatic species with terete, somewhat succulent or inflated leaves which if sectioned or viewed carefully with a lens can be seen to have four narrow longitudinal grooves running the length of the blade. Often its low- ermost leaf is much reduced, vestigial, or even absent while the uppermost leaf is largest and GOLDBLATT— GEISSORHIZA 391 ribbed on the inflated sheath and also occasion- ally on the free upper part. Such specimens are especially difficult to separate from narrow-leaved individuals of subsp. bicolor. The white to creamy yellow flowers of G. brehmii also occasionally have a dark center. The differences between subsp. bicolor and G. brehmii are consistent and it seems useful to om to recognize the latter as a distinct speci Gei. Mic TE subsp. bicolor is also sometimes confused with G. purpureolutea. This species has narrow two-grooved leaves, usually ca. 15 mm wide and a smaller flower with tepals 10-17(-22) mm long and a short perianth tube, 2-4 mm long. The plants are also usually short, 6-12 cm high, although they may reach 17 mm. e two species grow in the same habitat and occasionally close to one another. Specimens examined. SOUTH AFRICA. CAPE: 3218 (Clanwilliam) aca Mt., Clanwilliam (BB), MacOwan s.n. in Herb. Norm. Austr. Afr. 1969 (B, BM,G,GH,K, s SAM, Z), MacOwan 3328 (S, SAM). 3219 Piga Platkloof, Citrusdal (CA), Hane- kom 1241 (K, P 3318 (Cape iie near Mamre (AD-BC), MacOwan 2488 (K, SAM); Porterville (BB), Loubser 485 (NBG), 401 (BOL); Twenty Four Rivers, —— slope near road bridge, Goldblatt 6409 (K, MO, NBG, P Twenty Four Streams, near Saron, Lewis s.n. (BOL 31877); damp places near Malmesbury, 120 m (BC), Schlechter 1625 > BOL, BR, L, P ° Davis (SAM); near Stellenbosch, L. Bolus s.n. (BOL); slopes of the Bottelary hills, Acocks 3703A (S); Stellenbosch Golf Course, Bos 659 (STE, WAG); Stellenbosch Flats, Garside 87 (K), 1050 (K). 3319 (Worcester) Saron, sandy flats (AA), Lewis 5736 (NBG), Martin s.n. (NBG 60409); De Hoek Estates, near Saron, Lewis s.n. (BOL 31878); foot of Elands- ., Elandsrivier Farm (AC), Goldblatt 5856 ( ommonage at Gouda, damp onse in Goldblatt 4214 (MO); Steendal, Tulbagh, Pappe (BOL); Wolseley, Loubser 475 (NBG); Michells Pass. 1,500 ft. (AD), Schlechter 8931 (B, BM, BOL, K, L, MO, PH, S, Z); Breede River flats near Worcester eD Goldblatt 6982 (MO). 418 (Simonstown) near Somerset West (BB), Eck- *. & Zeyher Irid. 211 (83) (B, BM, FL, K AM); low lying fields, Somerset West, H. Bolus 5567 (BOL, K); Vergelegen, diei Lewis 5672 (NBG, STE); Faure, van Niekerk 2 6 (B sad WITHOUT PRECISE TY: Harvey 873 a 880 (E), Thunberg s.n. (Herb. b. Thunb. 991, 992, 961, UPS); between Piketberg and Piketberg Road Station, H. Bo- 392 lus s.n. (BOL), Masson s.n. “Ixia obtusata B" (BM), s.n. *Ixia graminea J. B." (BM), Roxburgh s.n. (BM “Ixia obtusata B," G). . Gei hi lutea Baker, J. Bot. n.s. 5: 238. 1876, Fl. Cap. 6: 68. 1896; Foster, Contr. Gray Herb. 135: 33-34. 1941. TYPE: South Africa. Cape: between Paarl and Pont, Drége 8476 [lectotype, K, effectively desig- nated by Foster (1941); isolectotypes, G, L]. FIGURES 9, 59. Geissorhiza sulphurea Schltr., Bot. Jahrb. Syst. 27: 99. dam near Cer Sc 8981 [lectotype, B, effectively desig- nated by Foster (1941); isolectotypes, BM, BOL, BR, H, K, MO, P, PH, PRE, S, Z]. Plants (4—)6-12(-17) cm high. Corm globose, more or less symmetric, 5-7 mm diam., tunics blackish, imbricate, regularly notched below into segments. Cataphyll not evident. Leaves 3, linear to narrowly ensiform, 1-2.5 mm wide, half as long to rarely as long as the stem, the margins and midrib strongly thickened, 2-grooved on each surface, more rarely 4 to several ribbed and grooved, the lower two leaves basal, the upper- most leaf largest, inserted near the base or on the lower part of the stem, sheathing the stem below for half its length, sheath inflated and strongly ribbed. Stem erect, simple. Spike 1—3(-5)-flow- ered, flexuose; bracts herbaceous, usually red- flushed near the apex, sometimes dry in the up- per half at flowering time, 7-8 mm long. Flowers stellate, cream to pale yellow, usually intense brown (to purple) in the basal third of the tepals and tube, rarely self-colored, often reddish on mm long, infundibuliform, usually about half as long as the bracts; tepals 10-17(-22) mm long, obovate 7-8 mm wide. Filaments equal, 5—6 mm long, often dark in the lower half; an- thers 4-6 mm long, yellow. Ovary 2-3 mm long, style dividing near the apex of the anthers, ranches ca. 2.5 mm long, recurved. Capsule un- known. Chromosome number, unknown. Flowering time. Mid-August to mid-Sep- tember. Distribution. Wet, sandy gravel flats, on the coastal plain between Paarl and Piketberg and in the Tulbagh Valley. Figure 59. Geissorhiza purpureolutea is a rare species of damp, poorly drained flats in the southwestern ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Cape. Its main range is the coastal lowlands be- tween Paarl and Piketberg but it also occurs in the upper Tulbagh Valley, to the interior. Typical G. purpureolutea has moderate-sized cream to yellow flowers with a dark brown to purple cen- ter. In a collection from west of Malmesbury (Lewis & Davis 2233) several plants have either a typical or much reduced marking or it may be lacking. A second collection from this same gen- eral area in the west of the range of the species, Salter 4756, consists mostly of plants lacking markings entirely (one plant has a dark throat), but in all other respects match the Lewis an Davis collection, as well as some other collec- tions (e.g., Loubser 2034 from Piketberg) which have flowers slightly on the small side for the species. These specimens that lack the central contrast- ing blotch match very closely collections in the Wolseley-Romans River area in the lower Tul- bagh Valley, amongst which is Schlechter 8961, the type collection of Geissorhiza sulphurea. The 11 Schlech ter frather small- flowered plants (tepals ca. 10 mm long), but the other collections from here (Salter 6836, Gold- blatt 2432), have flowers of the same size (tepals 12-15 mm long) as typical G. purpureolutea. It seems likely that these collections represent nothing more than self-colored G. purpureolutea, and accordingly G. sulphurea is reduced to syn- onymy here. Geissorhiza purpureolutea is closely allied to the G. imbricata complex from which it can usu- ally be distinguished by its shorter perianth tube, 2—4 mm long, comparatively small flowers with tepals 10-17(-22) mm long, usually with a large central blotch of dark color, spikes of 1—3(-5) flowers and narrow basal leaves to 2.5 mm wide and usually 2 (rarely 4 to several) grooved. A collection from the Darling area (Lotter s.n.) represents an unusual form of Geissorhiza pur- pureolutea. Plants have broad, several grooved leaves, and large flowers (tepals 20-22 mm long) with the typically short perianth tube of the species. The spikes have three or four flowers each. These plants are taken to be an extremely robust form of G. purpureolutea, rather than a separate taxon. ecimens examined. SOUTH AFRICA. CAPE: 3218 š rg ( , Loubser kop, S of Eendekuil turnoff in gravelly soil, M 6129 (MO, N 8 (Cape from Malmesbury on Hopefield ae pus ground (BA), Lewis & Davis s.n. 1985] GOLDBLATT — GEISSORHIZA FIGURE 59. Morphology and distribution of Geissorhiza purpureolutea. Habit x0.5; flower life size; leaf section much enlarged (Goldblatt 6129, foot of Zebrakop). ); ) Thompson 2613 (PRE, STE); near Wellington (DB), L. Bolus s.n. (BOL 20327, K, PRE), Compton 11731 (NBG), Barker 408 (NBG); near Paarl, Barker 5948 (NBG); Paarl flats, N end, Esterhuysen 6125 (BOL); between Paarl and Pont, Drége 8475 (G, L), 8476 (G, K, L); flats N of Paarl, Leighton 1960 (PRE); Huguenot, Compton 18257 (NBG); ? near Stellenbosch (DD), Ed- wards s.n. (BOL). 3319 (Worcester) Gouda commonage, heavy clay soil (AC), Goldblatt 2432A (K, MO, PRE); Ceres Road (Wolseley), 700 ft., Schlechter 8981 (B, BM, BOL, E, , H, K, MO, P, PH, PRE, S, Z); between Darling Bridge and Romans River Siding, Salter 6836 (BOL); flats between Bains Kloof and Wolseley, Goldblatt 2432 (MO, PRE, US); Elandsberg farm, S of Gouda, Gold- blatt 6209 (MO). 52. Geissorhiza barkerae Goldbl., sp. nov. TYPE: South Africa. Cape: N of Piketberg, wet sites at the foot of Zebrakop, Goldblatt 6391 (ho- lotype, MO; isotypes, K, NBG, PRE, S, W AG). FIGURES 8, 60. Planta 10-20 cm alta, foliis 3 inferioribus duobus basalibus lineari-ensiformibus marginibus costisque incrassatis, superiore infra inflato et vaginante, caule erecto saepe simplici, spica 2-4(-10) florum, floribus zygomorphis flavis atropurpureis insignis, tubo peri- anthii 7-8 mm longo, tepalis 22-30 mm longis, fila- mentis inaequalibus declinatis 15-20 mm longis, un —6 mm breviore quam aliis, antheris 5-6 mm longis. Plants 10—20 cm high. Corm globose, more or less symmetric, 7-11 mm diam., tunics blackish, imbricate, regularly notched below into seg- ments. Cataphyll not evident. Leaves 3, linear (to ensiform), 1-4 mm wide, (2 to) 4 to several ribbed on each surface, about as long as the stem, the lower two basal, the uppermost largest, in- serted near the base or in the lower part of the stem, sheathing the stem for half its length, the sheath inflated and strongly ribbed. Stem erect, usually 1-branched, flexed above the sheath of the upper leaf. Spike (2—)4-10-flowered, flex- uose; bracts herbaceous, the upper margins hya- line, becoming dry and usually red-flushed near ti ry above at flowering time, 10-13 mm long. Flowers zygomorphic, with the style and stamens declinate and held above the nearly horizontal upper tepal, pale yellow marked with purple in the lower half of the tepals and yellow in the center and tube, often reddish on the reverse of the outer tepals; perianth tube 7— 8 mm long, infundibuliform; tepa/s 22-30 mm long, lanceolate, 8-9 mm wide. Filaments un- equal, two longer, 15-20 mm, the shorter, 10- 14 mm long, yellow; anthers 5-6 mm long, yel- the apex r > 394 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 i? | EP SS ZR ; ET N S UN "S Haz ANIN WAA 1 y Z i. ee b A e METERS S 0 >> @) Bu | p ZZ M t = u^ A LB 18 b 2 21 E 60. Morphology and distribution of Geissorhiza barkerae. Habit x 0.5; flower life size; leaf section p). IG much piri (Goldblatt 6391, foot of Zebrako low. Ovary ca. 2-3 mm long, style dividing near the apex of the anthers, branches ca. 5 mm long, recurved. Capsule unknown. C} ber, 2n — 26 (Goldblatt 6391). Mid-September to mid-Oc- [41,2442 Flowering time. tober. Distribution. Seasonal marshes and stream banks in sand or light clay north and northwest of Piketberg, and locally in the Olifants River Valley. Figure 60. Geissorhiza barkerae is a very distinctive species, with a local distribution, north and west of Piketberg, where it grows in seasonally flooded or waterlogged sandy or light clay soils. It is allied to G. imbricata especially subsp. bicolor which Occurs in similar habitats to the south and to G. louisabolusiae which occurs in the Olifants River Valley to the north. It can readily be distin- guished from these two species by its large, zy- gomorphic flowers borne on drooping spikes of up to ten flowers. The flowers are pale yellow with dark purple in the lower part of the tepals while the throat and tube are also yellow. The stamens are declinate and very unequal, with the two longer filaments 15-20 mm long and the 1985] shorter one about two-thirds this length. In con- trast, G. imbricata subsp. bicolor has somewhat smaller and actinomorphic flowers sometimes with a dark center, and then usually a dark peri- anth tube (not so in the type). It also has fewer flowers in a spike (2-5, rarely to 10), and equal filaments. Geissorhiza louisabolusiae has self- colored, and smaller, deeper yellow flowers with a shorter perianth tube, and subequal to slightly unequal filaments, and typically much narrower, linear, 2-grooved leaves. Ihave pleasure in naming this species in honor of W. F. Barker, the South African botanist, and retired Curator of the Compton Herbarium, Kir- stenbosch. Her extensive collecting over many years has been of great value, particularly as she has taken great care in preparation of her spec- imens, thus making many characters easy to see which would otherwise be lost, and thus require recollecting. ens examined. SOUTH AFRICA. CAPE: 3218 (Clanwillam), chi reg Mas uh Leighton 121 (BOL, RE), Com G, W), Barker 2639 (BOL, NBG} dU. near Sepa (DB), Marloth 9247 (PRE); flats at foot of Zebrakop, S of Concordia farm, LIS 6391 (K, MO, NBG, PRE, S, WAG); N of Piketberg, wet ee along road to Het Kruis, Gold- Mah 542 C 3219 (W uh Warm Baths, Clanwilliam (CA), EX s.n. (BOL 31881 Ky Pass and ridus Leipoldt 3585 T Mene Town Wild Flower Show, Oct. 1939, Anon. (BOL 31880); “Tulbagh,” Thom s.n. (K). 53. Geissorhiza louisabolusiae Foster, Contr. Gray Herb. 135: 34. 1941, pro parte excl. var. longifolia. TYPE: South Africa. Cape: 6 mi. E of Graafwater, Salter 2790 (holotype, K; isotypes, BM, BOL). FIGURE 61. Plants (10—)15-20 cm high. Corm globose, 7— 9 mm diam., symmetric, tunics blackish, imbri- cate, notched into regular segments below. Cata- phyll not evident. Leaves 3, linear to nearly ter- ete, 1-2 mm wide, reaching to about the base of the spike, narrowly 4—6(—8)-grooved on each sur- th ribbed and inflated. Stern erect, sheathed for e to two-thirds its length, flexed above the sheath, occasionally with 1 branch. Spike flexuose, in- clined (2—)3-7-flowered; bracts herbaceous, oc- casionally the margin lined red above, often be- coming dry and membranous from the apex, 8- GOLDBLATT — GEISSORHIZA 395 10 mm long, the inner shorter than the outer. Flower actinomorphic, the tepals somewhat cupped, bright yellow; perianth tube 4—6 mm long, nearly reaching the apex of the bracts (exserted in old blooms), infundibuliform; tepals 18-28 mm long, 7-9 mm wide, oval. Filaments nearly equal to unequal, longer two 7-9 mm long, the third if shorter, 5-7 mm long; anthers 4.5-6 mm long, yellow. Ovary ca. 2-3 mm long, style di- viding near the apex of the anthers, branches ca. mm long, recurved. Capsule unknown. Chro- mosome number, 2n — 26 (Goldblatt 427). Flowering time. Late August to September. Distribution. Damp often waterlogged sand or shallow vleis, in the Olifants River Valley and valleys of the Olifants River Mountains to the west. Figure 61. Geissorhiza louisabolusiae has a limited dis- tribution, occurring in the Olifants River Valley and the adjacent valleys of the Olifants River Mountains to the west. It is restricted to wet sandy flats and vleis that are waterlogged in the late winter and spring. This habitat is now be- coming much reduced owing to the expanding ritmic and d » | fruit I dc that have been developed in this area, and G. /ouisabolusiae is, as a result, rapidly becoming endangered. It appears to be a member of the G. imbricata group and although it does not have the corru- gated leaves typical of the alliance, the sheathing part of the upper leaf is characteristically ribbed and grooved. It appears to be most closely related to the southwestern Cape species G. brehmii (syn. G. teretifolia) but it can readily be distinguished from this species. Geissorhiza louisabolusiae has deep yellow flowers with a relatively short peri- anth tube, 4-6 mm long, a flower with ascending somewhat cupped tepals, and usually one stamen slightly to appreciably shorter than the others. In contrast, G. brehmii usually has a longer perianth tube, 5-9 mm long, either white, cream, or pale yellow flowers, sometimes with a dark center and equal stamens. The leaves of G. /ouisabolusiae are more or less terete and narrowly 4—6(-8)- grooved, with the two basal equally well-devel- oped, while those of G. brehmii are fully terete and inflated, always 4-grooved and the lower- most of the basal leaves is much smaller than the other, sometimes being vestigial or even ab- sent. The variety Geissorhiza louisabolusiae var. longifolia recognized by Foster (1941), based on a Bachmann collection from near Moorreesburg ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 EE UO? ` = Ae — = o. OL are E x X022, GFP CA Morphology and distribution of Geissor FIGURE 61. section much enlarged (Goldblatt 6268, flats N of Citrusdal) is here referred to Geissorhiza brehmii. It appears to be no more than a robust form of this species but quite characteristic in its long perianth tube, inflated terete leaves, and much reduced or lack- ing lowermost leaf. Specimens examined. SOUTH AFRICA. CAPE: 3218 (Clanwilliam) 6 mi. E of Graafwater (BA), Salter 2790 (BM, BOL, K). 3318 (Wuppertal) Citrusdal (CA), Barker 7343 (NBG), 3078 (BOL, NBG), Leighton 1090 (BOL); flats N of Citrusdal, Salter 3659 (BM, BOL, K), Goldblatt 247 (BOL), Goldblatt 6268 (MO); 5 mi. N of Citrusdal, Lewis 1416 (SAM); Citrusdal vlei, Barker 3604 (NBG); Warm Baths, Olifants River Valley, Leipoldt s.n. (BOL 31879); Leeu River bridge, growing in vlei (CC), Hor- rocks 32 (NBG) 54. Geissorhiza brehmii Ecklon ex Klatt, Lin- naea 34: 653-654. 1866. Geissorhiza im- hiza louisabolusiae. Habit x0.5; flower life size; leaf 1). bricata var. brehmii (Klatt) Foster, Contr. Gray Herb. 135: 38. 1941. TYPE: South Af- rica. Cape: Hottentots Holland, about Som- u G. bicolor (Thunb.) N. E. Brown sensu lato]. Geissorhiza louisabolusii var. longifolia Foster, Contr. Gray Herb. 135: 34. 1941. TYPE: South Africa. Cape: road to Mooreesburg, Malmesbury, Bach- mann 1091 (holotype, B; isotype, Z). Geissorhiza teretifolia Lewis, J. S. African Bot. 7: 48. 1941. TYPE: South Africa. Cape: Kenilworth Race Course, Lewis 97 (holotype, SAM 53064). Plants 20-30 cm high. Corm 8-12 mm diam., globose, tunics blackish, imbricate, the outer lay- 1985] ers accumulating above, regularly notched below into triangular segments. Cataphyll single, ex- tending shortly above ground, firm memb nous. Leaves 2 or 3, usually terete and inflated or somewhat succulent with 4 very narrow lon- gitudinal grooves running the length of the leaf, the lower one or two basal, the lowermost small, often broken, or vestigial (occasionally lacking), the second about as long as the stem, the upper- and inflated, the free upper part often as long as the basal leaf (or leaves), terete or occasionally ribbed. Stem erect, often branched. Spike 2- 6-flowered, flexuose; bracts herbaceous, becom- ing dry and membranous from the apex, 7-10 (-15) mm long, the inner smaller than the outer. m tepals, occasionally dark colored in the throat and tube; perianth tube 5—7(—9) mm long, slightly shorter than, or reaching the apex of the bracts; tepals 15-20(-26) mm long to 9 mm wide, ob- ovate, obtuse. Filaments 5-9 mm long, equal; anthers ca. 5 mm long. Ovary ca. 2 mm long, style dividing near the apex of the anthers, branches 3-4 mm long, recurved. Capsule un- nown. Chromosome number, unknown. Flowering time. (August to) September to ctober. Distribution. Cape Peninsula, north to the Malmesbury district and east to Caledon and Bredasdorp in sandy ground, either swampy, or seasonally flooded, or at edges of pools. Figure Geissorhiza brehmii is closely related to G. im- bricata especially to subsp. bicolor (here includ- ing G. rubicunda) and the two taxa have a similar general appearance and flower. Geissorhiza breh- mii grows in wetter habitats, usually in standing water, or in sites usually flooded in the winter, but sometimes nearly dry by flowering time. It can be distinguished by its habitat and otherwise only in its vegetative characteristics, by its slen- der terete and slightly succulent, narrowly 4-grooved leaf, the lowermost of which is often much smaller than the others, or even vestigial or lacking. Geissorhiza imbricata has in contrast linear to ensiform, several ribbed, flat leaves, the lower of which is always well developed. Occa- sional narrow-leaved individuals of G. imbricata subsp. bicolor are encountered (e.g., Goldblatt 6409) but leaves of these plants are not terete or GOLDBLATT — GEISSORHIZA 397 | VAKA ⁄ Y TÑ LAE 4 DV YZ JUD Š ⁄ Z < Z < EE CIA 7251 4 FIGURE 62. Distribution of Geissorhiza brehmii. inflated and can be seen under a lens to be 1- 2-ribbed on each surface. In some collections of G. brehmii one or a few plants may have ribbed leaves (e.g., Compton 20095), and it becomes very difficult to distinguish such specimens from subsp. bicolor. As published, Geissorhiza brehmii appears to be a superfluous name for Klatt (1866) who cited two apparent synonyms in the protologue, “G. setacea Gawl.” and “G. aurea Ecklon." How- ever, Klatt was almost certainly citing G. setacea in reference to the “Botanical Magazine" fig. 1255 which Ker (Ker-Gawler) identified as G. setacea although in fact it is not this species, but probably G. hispidula (syn. G. humilis var. hispidula from the true G. setacea based on the Thunberg type. The other name cited, G. aurea Ecklon, is a nomen nudum. Klatt cited two collections in his description of Geissorhiza brehmii, both from near Somerset West, Ecklon & Zeyher Irid. 295 (G. brehmii Ecklon ms.) and Ecklon & Zeyher Irid. 211 (G. aurea Ecklon ms.). These are different species, the latter being a cream- to yellow-flowered form of G. bicolor with self-colored flowers and rela- tively broad effectively designated Ecklon & Zeyher Irid. 295 as the lectotype for G. brehmii when he reduced distinguished by its narrow leaves. Lewis (1941) quite independently described G. teretifolia based on collections from the Cape Peninsula, before Foster’s (1941) revision was published. Geisso- rhiza teretifolia differs in no significant way from G. brehmii, although the material cited by Lewis 398 is in general more robust than the lectotype of G. brehmii and has white flowers. Most populations of Geissorhiza brehmii have white- to cream-colored flowers, with a faint pink flush on the reverse of the outer tepals. Plants Cape Flats, however, apparently have cream flowers (yellow on drying) with a distinct red streak on the reverse of the outer tepals and a dark color in the throat and tube of the peri- anth. The Kraaifontein population (Compton 20095) comprises plants with either a dark, or self-colored throat and tube, while in the Muld- ers Vlei population (Lewis 4447) all plants ped the dark color in the center of the flower. Som plants of this collection also have unusually b. flowers, with tepals to 26 mm long and perianth tube to 9 mm. There seems no reason, however, to treat this form as a separate taxon. Other large-flowered populations have been recorded, e.g., Bachmann 1091 and H. Bolus 4339. Plants of these collections have apparently cream-colored flowers without any marking on the reverse of the tepals. These have either only one basal leaf, or at most a vestigial second basal leaf. The Bachmann collection is the type col- lection of Foster's Geissorhiza louisabolusiae var. longifolia. Geissorhiza louisabolusiae is in fact, superficially similar to G. brehmii, but always has two well-developed basal leaves, usually a shorter perianth tube, and subequal to unequal stamens. Differences between these two species, in leaf morphology and in the length of the peri- anth tube, make it easy to distinguish them. Geis- sorhiza louisabolusiae var. d is here re- duced to synonymy in G. brehm Geissorhiza brehmii has been heed at times with G. hispidula (Klatt, 1866) and with G. gem- inata (Foster 1941: 78). Several collections of G. hispidula made by Ecklon and Zeyher bear the manuscript name G. brehmii. Both G. hispidula and G. geminata (subgenus Weihea) have con- centric corm tunics and are unrelated to G. breh- mii, which has imbricate tunics typical of sub- genus Geissorhiza. Specimens examined. SOUTH AFRICA. CAPE: 3318 (Cape Town) Hopefield district QAB), Bachmann 1090 (Z); road to Moorreesburg (?BA), Bachmann 1091 (B, Z); Kenilworth Race Course, in pools (CD), Lewis 4796 (PRE, SAM), 97 (SAM), 480 (SAM), Barker 4795 (NBG), 4115 (NBG), Salter 7738 (BM, BOL, CT, K, 95 (BOL, NBG, STE); Bergrivier at Paar “feuchten Wiesen und in Dümpfel" (DD), Drége y ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 P, S); Stellenbosch flats, Hol- di ó E de 1063 (K); Old Golf Course, Stellenbosch, Du Preez s.n. (STE); Mulders Vlei, Lewis 4447 (PRE, 3319 (Worcester) Hex River Valley (BC-CB), Da- vidson 75 (SAM); between Worcester and Villiersdorp weve vis 2899 (SAM), 2900 (SAM). 8 (Simonstown) Ottery Road Station (AB), Lewis i SAM; Wynberg, Wilms 3733 (B), Schlechter 1557 (B, G, M, W, Z); Cape flats (BA), Ecklon & Zeyher 427 G). T PRECISE LOCALITY: Cape of Good Hope, Siebold. 526 (B), z s.n. (PRE); Zonderend Mts., Stokoe s.n. (SAM 57388). 55. Geissorhiza sulphurascens Schltr. ex Foster, Contr. Gray Herb. 35: 39. 1941. TYPE: South Africa. Cape: Bokkeveld W of Nieuwoudt- ville, Marloth 7658 (holotype, B; isotype, PRE). FIGURE 63. Plants 12—20 cm high. Corm globose, 6-8 mm diam., symmetric, tunics blackish, imbricate, notched into regular segments below. Cataphyll not evident. Leaves 3, narrow, linear, ca. 1 mm wide, reaching the midline of the stem or the base of the spike, the margins and midrib en- larged and 2-grooved on each surface, the lower two leaves basal, the uppermost basal or inserted just above the ground, largest, sheathing the stem in its lower two-thirds, the sheath ribbed and times becoming dry and brownish from the apex, 7—9 mm long, the inner shorter than the outer. Flower stellate, or the tepals slightly cupped, cream; perianth tube 3-4 mm long, enclosed in the bracts, infundibuliform; tepals 13-18 mm long, 5-7 mm wide, oval. Filaments equal, 5-8 mm long; anthers ca. 6 mm long, pale yellow. Ovary ca. 2.5 mm long, style dividing at the apex of the anthers, branches ca. 2 mm long, recurved. Capsule more or less globose about 5 mm long. Chromosome number, unknown. Flowering time. Late August to September. Distribution. Nieuwoudtville Escarpment, in damp sand. Figure 63 1985] GOLDBLATT — GEISSORHIZA @ G.sulphurascens $ G. minuta FIGURE 63. Morphology of Geissorhiza sulphurascens and distribution of G. sulphurascens and G. minuta Habit x0.5; flower life size; partia Grasberg road, Nieuwoudtville). Geissorhiza sulphurascens is endemic to the poorly drained, wet, sandy sites. It is related to the G. imbricata group, the other species of which occur to the south from the Olifants River Valley to the Cape Peninsula and Bredasdorp. While it seems somewhat isolated taxonomically, it is perhaps most closely related to G. /ouisabolusiae a species with fewer and larger, bright yellow flowers on a lax inflorescence and typically un- equal stamens. The pale creamy yellow flowers and congested inflorescence of G. sulphurascens are distinctive, as are the linear and 4-grooved basal leaves and well-developed uppermost leaf which sheaths at least two-thirds of the stem. Specimens examined. SOUTH AFRICA. CAPE: 3119 (Calvinia) Nieuwoudtville Escarpment NW of Van ower and gynoecium x 1.5; leaf section much enlarged (Goldblatt 5841, Rhyns Pass (AC), Oliver 3849 (K, PRE, STE); Nieu- woudtville, L. Bolus s.n. (PRE 36875), Loubser 953 (NBG); Willemsrivier, Leipoldt 720 (BOL, PRE); mar- shy soil near Nieuwoudtville, Leipoldt 792 (BOL); Bokkeveld W of Nieuwoudtville, Marloth 7658 (B, PRE); Grasberg road, NW of Nieuwoudtville, da Nieuwoudtville, Lewis 587 ; 4 km NW of Nieuwoudtville on Grasberg road, Goldblatt 6218 (MO); 2 mi. W of Nieuwoudtville, Hall 3874 (NBG); Uit- komst farm SW of Nieuwoudtville, in sand, Barker 10740 (K, NBG). 56. Geissorhiza minuta Goldbl., sp. nov. TYPE: South Africa. Cape: Pakhuis Pass, Ester- huysen 3193 (holotype, BOL; isotypes, K, MO). Planta 3-8(-12) cm alta, tunicis cormi imbricatis, foliis 3, inferioribus duobus basalibus linearibus, 400 superiore caulem vaginante, caule pru spica 2-6 florum, floribus stellatis, albis, tubo ca. 1 mm longo, tenali 7-8 mm longis, antheris 2— 3n mm lae stylo diviso prope apicem antherarum. Plants 3-8(-12) cm high. Corm globose, 5—6 m diam., symmetric, tunics blackish, imbri- cate, notched into regular segments below. Ca- taphyll not evident. Leaves 3, linear, ca. 1 mm wide, about as long as the spike, the margins and midrib enlarged and 2-grooved on each surface, the lower two leaves basal, the uppermost basal or inserted just above the ground, longer or short- er than the lower leaves, sheathing the stem in its lower two-thirds, the sheath ribbed and some- what inflated. Stem erect, sheathed for about two- thirds its length, flexed above the sheath of the upper leaf, occasionally with 1 branch from the Spike flexuose, inclined, 2—6-flowered; ape m lo ner about as long as the outer. Flower evidently white, stellate; peri- anth tube ca. 1 mm long, enclosed in the bracts, infundibuliform; tepa/s 7-8 mm long, ca. 2.5 mm wide, oval. Filaments apparently equal, 2-3 mm long; anthers 2-3 mm long, pale yellow. Ovary ca. | mm long, style dividing just beyond the apex of the anthers, branches 1-1.5 mm long, recurved. Capsule unknown. Chromosome num- ber, unknown. Flowering time. September to early October. Distribution. Moist, rocky places in the Pak- huis Mountains. Figure 63. Geissorhiza minuta is a dwarf species of sub- genus and section Geissorhiza, apparently most closely related to G. sulphurascens, an endemic of the Nieuwoudtville Plateau. Geissorhiza mi- nuta has white flowers, which are amongst the smallest in the genus. Its short perianth tube, ca. 1 mm long, tepals 7-8 mm long, and short an- thers, 2-3 mm long, as well as slender habit and dwarf stature make confusion with any other species unlikely. It has only been collected in the vicinity of Pakhuis Pass and is probably endemic to this limited area to the north of the main Cedarberg range. Specimens examined. SOUTH AFRICA. CAPE: 3219 (Wuppertal) Pakhuis Pass, w Lewis 1871 (SAM); Pakhuis Pass, moist spot among rocks, Esterhuysen 3193 (BOL, MO). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 57. Geissorhiza eurystigma L. Bolus, S. African Gard. 21: 282. 1931. Geissorhiza mathewsii var. eurystigma (L. Bolus) Foster, Contr. Gray Herb. 135: 27-28. 1941. TYPE: South Africa. Cape: near Mamre, L. Bolus s.n. (lec- totype, BOL 19876, here designated; isolec- totype, K). FIGURE 64. Ixia igs de la Roche, Diss. 17-18. 1766. Geisso- rhiza monantha Sweet, Hort. Brit. 2nd edition, 5 3. 1830. nom.nov. pro Ixia secunda de la Roche [G. Secunda (Ker) Ker, 1804 bars transfer to Geis- of G. mon- anthos Ecklon, 1827 after Articles 64. 2 and 75 of (holotype, L). Ixia rochensis Ker non sensu Ker, Bot. Mag. 17: tab. £P p. nom. illeg. superfl. pro T. secunda de l he (misapplied to G. radians). G i s. (Ker) Ker, Ann. Bot. (Kónig & Sims) 1 . 1804, nom. illeg. basion. illeg. su- perfl. ae to G. radians). Geissorhiza rochensis var. monantha Steals pen non sensu Baker, Handb. Irid. 156. 1892, Fl 71. 1896 (misapplied to G. monanthos Ecklon). Ixia rochensis var. spithamaea Ker, Bot. Mag. 17: tab. 802. Geissorhiza rochensis var. spithamaea (Ker) Baker, Handb. Irid. 156. 1892, FI. Cap. 6: 71. 1896, pro parte; Foster, Contr. Gray Herb. 135: 29-30. 1941, pro parte. TYPE: South Africa. Cape: precise locality unknown, Masson s.n. “Ixia violacea” (lectotype, BM, here designated). Plants 8-15(-20) cm high. Corm globose, more or less symmetric, 8-11 mm diam., truncate at the base, tunics imbricate, blackish, layers notched below into regular segments. Cataphyll membranous or evidently lacking. Leaves 3, en- siform, conspicuously several ribbed, reaching to the base or middle of the spike, 3-6 mm wide, the lower two leaves basal, the uppermost in- serted at the base or in the lower part of the stem, sheathing and inflated below, often as long and larger than the lower leaves. Stem flexed above sheath of the upper leaf and at the base of the spike, rarely bearing a bract-like leaf in the upper half, simple or 1—2-branched. Spike flexuose (1— 2-5(-7)-flowered; bracts herbaceous, green, be- coming dry and membranous from the apex and often reddish on the upper margin, 9-13 mm long, the inner about as long as the outer but narrower. Flowers evidently actinomorphic, the tepals widely cupped, dark blue with a red center; perianth tube fairly long, and widening gradually from the base, 6—9 mm long, exserted from the bracts; tepals 16-24 mm long, 9-13 mm wide, obovate, blue above, red in the lower half. Fil- 1985] GOLDBLATT— GEISSORHIZA 401 — on ae, = Dorsa —= 2 D Le. Zm nit 18 19 l L n L FiGuRE 64. Morphology and distribution of Geissorhiza eurystigma. Habit x 0.5; flower and separated bracts o life size; gynoecium and leaf section much enlarged (Burns s.n., farm Dassenberg, Kan aments to 9 mm long, erect, equal; anthers 4.5— 5.5 mm long, yellow. Ovary 2-3 mm long, style dividing near the apex of the anthers, branches 4 mm long, broad and heavily ciliate-pubescent. Capsule ovate-oblong, about as long as the bracts. Chromosome number, unknown. Flowering time. September to early October. Distribution. estern Cape coastal belt, cen- tered around Darling and Mamre, and extending southeast to Kalabaskraal. Figure 64. Geissorhiza eurystigma was first described in 1766 by Daniel de la Roche, who named it Ixia secunda. The type, in the Leiden Herbarium, is today still well enough preserved to make this identification certain (Goldblatt & Barnard, 1970). A year after the publication of 7. secunda, Bergius (1767) described a second species (now G. aspera Goldbl.) also calling it 7. secunda, and in the appendix to this work he cited de la Roche's I. secunda. While Bergius’ species is today not nkop). considered a validly published species, but mere- ly a misapplied name, the eighteenth century bo- tanical community either ignored the priority of de la Roche's species, or considered it and the better known Bergius plant conspecific (e.g., Thunberg, 1783), and the name was used widely for the small blue-flowered G. aspera. Ker (18022, 1802b) realized that Ixia secunda de la Roche was different from the species then bearing this epithet, but instead of applying the name 7. secunda to what he believed was de la Roche's species, he renamed it 7. rochensis and form leaved plant figured in the “Botanical Mag- azine," now G. radians (long known as G. ro- chensis) and a variety (B) spithamaea, which in fact corresponds with de la Roche's 7. secunda, now G. eurystigma. The two species are similar in flower form and color but G. eurystigma has broader, multi-ribbed leaves, broad flat ciliate 402 style branches and tepals which lack the white and and curious pits in the tepals characteristic of G. radians Robert Sweet realized that Ker's Geissorhiza rochensis, as figured in the “Botanical Maga- zine," was distinct from Ixia secunda de la Roche and in 1830 he renamed the latter G. monantha, citing a figure published by Houttuyn (1780) which is correctly named J. secunda de la Roche. In his catalogue of plants cultivated in Britain, Sweet (1830) also recognized Ker's G. rochensis (i.e., G. radians) as G. rocheana, indicating that he regarded the two as separate species. The name G. monantha is here considered a homonym o G. monanthos Ecklon. The two names have slightly different endings, one Greek and the oth- er Latin, but they are certain to be confused and should not both be used in the same genus. When Geissorhiza eurystigma was collected in the 1820s by Zeyher and then in the later nine- teenth century by Templeman, it was again con- fused with the more common G. radians (G. ro- chensis sensu auct.) and robust plants of both species were referred to G. rochensis var. spitha- maea Ker. Then in 1931 Louisa Bolus recol- lected the species and, believing that it was new to science, described it as G. eurystigma, a name referring to its unusual and conspicuous style branches. Foster (1941) regarded Geissorhiza eu- rystigma as a variety of the related G. mathewsii, a treatment not followed here. The differences between the two seem to me sufficiently great to merit separation at the species level, even though they share so unusual a character as their broad flat style branches. The two species are almost certainly ed allied. Both have relatively broad, ri leaves, deep blue flowers with a red center, Send conspicuous broad flat style branches. The differences between the two species are discussed under G. mathewsii, the following species. Wa edi exami dup EAD AFRICA. CAPE: 3318 (Cape Town), Mamre AD), Barker 1810 (BOL, NBG), 3834 (NBG), Waser 436 (NBG), van Nie- kerk 272 (NBG), Compton 11765 (NBG); near e (AD-CB), L. Bolus Hs ‘BOL 19876), Sidey 2283 (S); AM mi. No Mamre on Darling road, Lewis 1 037 (SAM): abi amre and Darling, Lewis 3634 (SAM), Lam a eluate 2074 (K); between Mamre and Suid a rnoff to Mud River, Goldblatt 7266 (MO); first pes n Ganzekraal (CB), Barker 723 (NBG); Kalabas- kraal (DA), Werdermann & Oberdieck 323 (B, GH, K, PRE), Strey s.n. (M); farm Dassenberg, N slopes of Kanonkop Range, Burns s.n. (MO). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 WITHOUT PRECISE LOCALITY: Cape, Anon. s.n. "Ixia secunda de la Roche" (L), Herb. Burman s.n. '*Gladi- olus junceus" (G), Masson s.n. (BM). 58. Geissorhiza mathewsii L. Bolus, Ann. Bolus Herb. 4: 42. 1926; Foster, Contr. Gray Herb. 135: 27. 1941, excluding var. eurystigma. TYPE: South Africa. Cape: damp marshy places near Darling, Mathews s.n. flor. 1923 (lectotype, BOL 18502, here designated); 1926 ( BOL 18502). FIGURES 10, 65. syntype, Plants 8-18 cm high. Corm globose, more or less symmetric, 7-10 mm diam., truncate at base, tunics imbricate, blackish, layers notched below into regular segments. Cataphyll membranous or evidently lacking. Leaves 3, the lower two basal, reaching to base or middle of spike, 2-4 mm wide, ensiform, conspicuously several ribbed, lowermost smallest, the third and uppermost leaf inserted at base or in lower part of stem, sheath- ing and inflated below, also several ribbed, longer and larger than lower leaves. Stem flexed above sheath of upper leaf, and at base of inflorescence, smooth, rarely bearing a bract-like leaf in upper half, simple or 1—3-branched. Spike secund, flex- uose, 2-6-flowered; bracts herbaceous, green, be- coming dry and membranous from apex, and often reddish on upper margin, 6-12(-15) mm long, inner about as long but narrow. Flowers actinomorphic, nearly stellate when fully open, mm long, included in the bracts; tepals 14-16 mm long, 9-10 mm wide, obovate, blue above, reddish in lower half. Filaments 6.5-8 mm long, erect; anthers ca. 4.5 mm long, articulated and horizontal, lying over the style branches, brown- purple. Ovary ca. 2 mm long; style dividing near the base of the anthers, branches 2-3.5 mm long, broad and flat, evidently smooth, concealed by the anthers. Capsule globose, to 16 mm long. hromosome number, unknown Flowering time. Mid- to late September. Distribution. The Darling area of the west coast in wet low lying situations. Figure Geissorhiza mathewsii is a rare local endemic of the Darling area of the southwestern Cape coast where it grows in wet, low lying areas. It is related to and often confused with G. eurystig- ma. The two have a similar appearance and broad, ribbed leaves, blue flowers with a red cen- ter, and the peculiar broad flat style branches, 1985] GOLDBLATT — GEISSORHIZA 403 Ficure 65. Morphology and distribution of Geissorhiza mathewsii. Habit x0.5; flower and separate tepals life size; gynoecium and anther x 2; leaf section much enlarged (Goldblatt 7268, Waylands, Darling). unique to these two species. Foster (1941) treated G. eurystigma as a variety of G. mathewsii, re- marking that it differed primarily in its longer perianth tube, shorter filaments, and longer style. In this distinction Foster is only partly correct, G. mathewsii has a short tube, to 2.5 mm long and a shorter style but its filaments are shorter than those of G. eurystigma. However, there are other differences between the two and their treat- ment as separate is desirable. In addition to the short perianth tube, G. mathewsii has an alto- gether smaller flower with tepals to 16 mm long, whereas the tepals of G. eurystigma are up to 24 mm long. More significantly, the stamens and style are quite different. In G. eurystigma, the stamens are erect and held below the broad and heavily ciliate style branches, but in G. mathew- sii the style divides near the base of the anthers which are articulated and lie horizontally when the flower is fully open, covering the nearly smooth style branches. The floral bracts also tend to be shorter than those of C. eurystigma and the lowermost leaf smaller than the others. The species is known from relatively few collections and is apparently very restricted in its distribu- tion. The distinctive flower coloring of Geissorhiza mathewsii and G. eurystigma is also found in G. radians and in some forms of the unrelated G. monanthos (section Planifolia), and they may be sometimes be confused. Geissorhiza radians can be recognized by the pitted markings on the te- pals, narrow leaves, and longer perianth tube and G. monanthos by its puberulous stem and smooth, flat leaves. Similarity of floral coloration in G. monanthos is most likely due to color mim- icry, common in the southwestern Cape flora. Geissorhiza mathewsii grows together with G. radians and G. monanthos at some sites and the differences between their habitat preferences is then evident. Geissorhiza radians grows in the wettest places, often in standing water; G. ma- thewsii prefers slightly higher ground, but still in waterlogged soil; and G. monanthos occurs only in well-drained areas surrounding wet depres- sions. Specimens examined. SOUTH AFRICA. CAPE: 3319 (Cape Town) “in humidis" near Darling (AD), Ma- thews s.n. (BOL 18502); near Darling, Compton & Lamb Lamb s.n. (K); Waylands farm, edge of vlei N of the railway line, Goldblatt 7268 (K, MO, NBG, PRE), Jar- vis s.n. ( . WITHOUT PRECISE LOCALITY: Malmesbury (? distr.), Kassner s.n. (B). 404 59. Geissorhiza radians (Thunb.) Goldbl., . 1983. Ixia Ja is 03. TYPE Thunberg s.n. [lectotype, Herb. Thunb. 982 UPS, designated by Goldblatt (1982); syn- type, Herb. Thunb. 983 UPS = Romulea sp.]. FiGunES 11, 66. Ixia rochensis Ker, Bot. Mag. 17: tab. 598. 1802, nom. illeg. superfl. pro Z. secunda de la Roche (1766) = excl. var. on Foster, Contr. Gray Herb. 135: 29. Ixia Vd ad Roemer & iip Syst. Veg. 1: 379. 1817 . superfl. a dela Roche = G. eu md ma L. Bolus. Hog did larochei Paper & Schult.) Lo 1830, nom. illeg. basion. ies. a T rocheana Sweet, Hort Ist re 399. 7, Hort. Brit. 2nd o 503. TYPE: South Africa. Cape: without locality e tration in Bot. Mag. 71: tab. xd 180 — iac Klatt, Abh. Naturf. Ges. Halle 90 (Erg. 56) 1882. TYPE: als Africa. Cape: near oa Drége 8486 KAAN B, “Herb. Lu- beck”; isotypes, i Geissorhiza cyanea Ecklon, Topo gr. erz. Pflanzen- ml. Ecklon 20. 1827, nom. i et ms. (Eck- ná Zeyher Irid. 219). Gaso ha rochensis var. Ve Logo en Baker, Handb. Irid. 156. 1892, Fl. Cap. 6: 1896 as to most specimens Eo but excl. MN Foster, Contr. Gray Herb. 135: 29-30. 1941, pro parte (the type of var. es Is G. eurystigma L. olus). Rochea venusta Salisb., Trans. Hort. Soc. London 1: : 2. 1812, nom. nov. pro Ixia rochensis Ker sen- u Ker, nom. illeg. gen. illeg. non v " Geissorhiza radians Gawl.," Nom. Bot. 2nd edition, 1: 668. 1840, in syn. Plants 8-16 cm high. Corm globose, symmet- ric, truncate below, 6-9 mm diam., tunics dark, imbricate, notched below into regular rectan- gular segments. Cataphyll membranous pale. Leaves 3, linear, 0.5-1.5 mm wide, with thick- ened margins and enlarged midrib and 2-grooved on each surface, about as long, somewhat longer, or shorter than the stem, the lower two leaves basal, the upper inserted on lower part of stem, the sheath inflated and ribbed. Stem erect, flexed above the sheath of the upper leaf and at the base of the spike, simple or 1-branched. Spike 1—4 (-6)-flowered; bracts 8-13(-16) mm long, her- baceous, the inner shorter than the outer. Flowers zygomorphic with the stamens declinate, the te- pals widely cupped, deep blue with a red center ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 and narrow white band between, with a dark, excavated pit in the midline of each tepal; peri- anth tube 6-8 mm long, infundibuliform, exsert- ed from the tube; tepals 15-22 mm ovate, 12-15 mm wide. Fil long, dm unilateral and declinate, held above the dorsal tepal, curving upward near the an- thers; anthers unilateral, erect, 5-6 mm long, pol- len red-brown. Ovary 2-3 mm long, style lying under the stamens, ca. 2 mm long, branches 4 mm, recurved. Capsule unknown. Chromosome number, unknown. Flowering time. September to early October. Distribution. Flats in the western Cape coast- al belt, from the Darling area south to Paarl and possibly to False Bay, damp sites in sandy soil, often of granitic origin. Figure 66. This species is one of the most striking in the genus, and one of few, fairly widely known. It is occasionally found in cultivation and its large, deep blue flowers with contrasting white band around the red center make it worth the effort to maintain the plant in garden situations. As is the case with many species that have received botanical attention over a long period, it has a complex nomenclatural history. It is also unfor- tunate that the name it has been known by for over 150 years, Geissorhiza rochensis, is illegit- imate (Goldblatt, 1983), so that this well-known species now has to carry the new name G. radians (Thunb.) Goldbl. The type collection of pies foci pega was made by Thunberg in the 1770s, although h only described the species years see in 1803, as as Ixia radians. A year before, Ker (1802a) pub- lished Ixia rochensis as a new name for Ixia secunda de la Roche, but based on plants grown in England, and figured in the “Botanical Mag- Ker mistakenly believed the figured species to be the same as the Ixia secunda, de- scribed in 1766 by Daniel de la Roche, but this is actually conspecific with the superficially sim- llar G. eurystigma L. Bolus. Ker used the name Ixia secunda for a third species, the small, blue- flowered G. aspera, and so cited I. secunda de la Roche as a synonym of his T. rochensis. Ker's nomenclatural treatment is unacceptable and to- day I. rochensis is regarded as a superfluous and illegitimate name (Goldblatt, 1983). The next available synonym for the species figured by Ker is Ixia radians Thunberg, and the combination in Geissorhiza was made in 1983 (Goldblatt, 1983) 1985] GOLDBLATT — GEISSORHIZA i KA v CUZ a A Zl d ay FiGURE 66. Morphology and distribution of Geissorhiza radians. Habit life size; opened flower x 1.5; leaf section much enlarged (cultivated plant, hort. Kirstenbosch, no voucher). Sweet (1830) seems to have been aware of this confusion and he renamed /. secunda de la Roche, monantha, and G. rochensis sensu Ker, G. rocheana. Other synonyms of G. radians include G. tulipifera Klatt, merely a later synonym, and G. cyanea Ecklon, a nomen nudum. Robust specimens of G. radians have often been referred to G. rochensis var. spithamaea by both Baker (1896) and Foster (1941). There is no character other than size to distinguish the plants to which they applied the name and no infraspecific taxon of G. radians is recognized here. The type of var. spithamaea Ker, Masson s.n. “Ixia violacea," in the British Museum is actually the closely related G. eurystigma (syn. G. monantha Sweet), which Ker regarded as a variety of his G. rochensis. Geissorhiza radians was once fairly common along the Cape west coast, originally extending from False Bay in the south to the Malmesbury district in the north. Its range is now much re- duced due to expanding farming activity. This is unfortunately inevitable, as G. radians grows along streams and in wet flats in a rich agricul- tural area. It is protected to some degree in small reserves in the Darling area by local farmers, at least from direct ploughing, but alien plant species, animal grazing, and effects of fertilizer and weed killer runoff are gradually changing the environment, and this and many other lowland west coast species are becoming increasingly threatened. Geissorhiza radians is distinctive in its blue and red flower coloration and peculiar pitted markings in the midline of its tepals as well as in its linear ( ti ] )! itt enlarged margins and midrib. Similarly colored flowers are found in G. mathewsii and G. eury- stigma, both of which lack the pitted tepal mark- 406 ings and have broad, ribbed leaves. Some forms of the unrelated G. monanthos also have simi- larly colored flowers but this species has flat leaves and a puberulent stem. Specimens examined. SOUTH AFRICA. CAPE: 3318 (Cape ida a sie (AC), Barker 3844 (NBG); pv AD), n & Zeyher 219 (66) (B, FI, G, MO), Ze he 2 (SAM), MacOwan s.n. in Herb. . Afr. 5 Pappe s. n. (BM, K); near Darling, H. Bolus 12843 (BM, BOL, BR, Z), Levyns 3248 (K); Darling, Guthrie 2054 (NBG), Stokoe s.n. (SAM 69599), Grant 2527 (BOL, BR, M, MO, WAG), Marloth 9562 (STE); Darling Flora Reserve, uibs 8644 (NBG), Lewis 5068 (NBG), 5537 (N ; 8 mi. W of Darling, Strey s.n. (M); Mamre Hills, Wasserful 437 (NBG), Barker 1806 (NBG), 3838 (NBG); between Mamre and Darling, Lewis 3634 (SAM); 3 mi. S of Darling, Lewis 1306 (SAM); Cas near Malmesbury (BC), Schlechter 1610 (B, G, W Spenglersdrift, near 20645 (NBG, PRE); Agterpaarl, damp sites (DB), Loubser 468 (NBG); near Paarl, Drége 8486 (B, BM, G, K, L, MO, P, S); Muldersvlei (DD), Lewis 4446 (SAM); damp place S of Bottelary road, Acocks 2162 (S 3418 Span e Hottentots Holland (BB), Eck- lon & Zeyher Irid. 292 (MO). WITHOUT PRECISE LOCALITY: Herb. Burman s.n. “Ixia secunda Berg. Cap. 6" (G), "OldenPurs 59 (BM). SECTION MONTICOLA 10. Section Monticola Goldbl., sect. nov. TYPE: G. burchellii Foster. Tunicae cormi durae et lignosae vel molles, foliis plerumque 3, ensiformibus ad linearibus vel teretibus natis, tubo brevi vel longo, staminibus aequalibus vel inaequalibus Corm tunics hard and woody to fairly soft in texture, /eaves ensiform to linear or terete, more or less plane or with the margins and midrib variously thickened, bracts herbaceous below, becoming dry above, flowers stellate to hypocra- teriform, actinomorphic or declinate, the tube short or reaching to the apex of the bracts, sta- mens equal or more often strongly unequal. Species. 9. 60. Geissorhiza burchellii Foster, Contr. Gray Herb. 135: 25-26. 1941. TYPE: South Africa. Cape: Langeberg near Zuurbraak, Schlechter ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 2160 (holotype, B; es B, BM, BOL, BR, PRE, Z). FIGURE 67. Plants 12-20 cm high. Corm globose, tapering above, 7 mm diam., tunics light brown, more or less imbricate, papery in texture, cut below into narrow segments, drawn into points above. Cataphyll membranous, apparently solitary. Leaves 2-4, about half as long as the stem, oc- casionally about as long, 1.5-3 mm wide, the margins and midrib thickened and 2-grooved on each surface, often sticky and with soil adhering below, the lowermost leaf basal, the others in- serted towards base and sheathing the stem for some distance above the ground, the uppermost sheathing for half its length. Stem erect, smooth, bearing in addition to a cauline leaf, 1-3 short, sheathing bracts, simple or 1—3-branched, flexed at the base of the inflorescence when in bud. Spike (1-)3-9-flowered, flexuose; bracts (6—)8— 10 mm long, herbaceous, flushed red above. Flower stellate when fully open, the tepals some- times becoming reflexed with age, pale to deep purple or violet, pale yellow in the throat and tube; perianth tube 4—6 mm long, cylindric, about as long or slightly longer than the bracts; tepals 12-18(—22) mm long, 5-6 mm wide. Filaments unequal, the two longer 10-14 mm long, the shorter 6-10 mm long; anthers yellow, 4—5.5 mm long. Ovary 1.5-2 mm long, style dividing near the apex of the anthers, branches ca. 4 mm long, recurved. Capsule more or less globose, 5-7 mm long. Chromosome number, 2n = 26 (Esterhuy- sen 35363 Flowering time. December to January. Distribution. Mid to upper elevations in the a Kloof and Heidelberg. blooming well only after fires. Figure 67 Geissorhiza burchelliiis an attractive but poor- ly known species of the western Cape mountains. It extends from the Slanghoek Mountains in the north to the Langeberg at Swellendam in the east and appears to grow at middle elevations on moist, often south-facing slopes. It flowers in the wild only after veld fires and in this character seems typical not only of the group of species to which it is allied but to most Cape mountain species of Geissorhiza. It is related to Geissorhiza grandiflora, a species with a similar montane distribution and to the Cape Peninsula endemic G. tabularis, and is eas- 1985] GOLDBLATT — GEISSORHIZA FIGURE 67. Morphology and distribution of Geissorhiza burchellii. Habit x0.5; flower and corms life size; berg). leaf section enlarged (Esterhuysen 35159, Goedehoop Peak, Langeber ily confused with these species as well as with the more widespread G. ramosa. All are montane species with characteristic branched stems; pap- ery to woody, more or less imbricate corm tunics; leaves with slightly to heavily raised margins and midrib; and distinctly unequal stamens. Geis- sorhiza ramosa stands out in having very small flowers with a perianth tube 2-3 mm long, tepals 5-9 mm long and anthers 2-3.5 mm long. The flowers of G. tabularis have a similar short tube 1.5-2 mm long, but are larger, with tepals 15— 18 mm long and anthers 3.5-5 mm long. The flowers of G. grandiflora are even larger with a tube 10-22 mm long, tepals 22-30 mm long and anthers 5-6 mm long. Geissorhiza burchellii has flowers of intermediate size with a tube about 4— 6 mm long, tepals 12-22 mm long and anthers 4—5.5 mm long. Its purple flowers and moder- ately long perianth tube give G. burchellii a very different appearance from the pale-flowered G. tabularis but it is easily confused with G. gran- diflora and care must be taken in separating these two species. A collection from Grassy Dome, Du Toits Kloof, Jackson s.n., is tentatively assigned here. The single specimen lacks leaves and corm, but the flowers match those of G. burchellii well. Notes with sheet indicate that the flowers had white and dark red markings at the base of the tepals. Collections assigned here to G. burchellii 408 from near Caledon, H. Bolus 9891, and Land- drost Kop, Stokoe s.n. and Thorne s.n., consist of plants within which the thickenings on the margins and midrib are poorly developed. In leaf morphology, these plants thus approach G. gran- diflora but perianth tube and tepal length are nearly typical of G. burchellii and this is where they probably belong. Specimens examined. SOUTH AFRICA. CAPE: 3318 (Cape Town) Landdrost Kop (BB), Stokoe s.n. (SAM 55712); E slopes of Landdrost Kop, Thorne s.n. (SAM 51559). 3319 (Worcester) Cossacks, Slanghoek Mts., 4,000 ft. (CA), Esterhuysen 24006 (BOL, K, PRE); Grassy 3,000 ft., after burning, Esterhuysen 35604 (K, MO, NBG, PRE, S, US, WAG); Langeberg near Zuurbraak, : 300 ft. (DC), Schlechter 2160 (B, BM, BOL, BR, RE, hes Goedehoop Peak, Heidelberg, steep moist S ise n burned area, 3,500-4,000 ft. (DD), Ester- nicae 35159 (BOL, K, MO, PRE, S, US); Langeberg above Grootvadersbos, 3,500—4,000 ft., Esterhuysen 18273 (BOL, K, 2 vue WITHOUT PRECIS Y: hills near Caledon, H. Bolus 9891 (BOL, K). "Burchell 7322 (K). 61. Geissorhiza grandiflora Goldbl., sp. nov. TYPE: South Africa. Cape: N side of Limiet- berg, Bains Kloof, ca. 4,000 ft., Esterhuysen 35832 (holotype, BOL; isotypes, C, E, K MO, NBG, PRE, S, US, WAG). FIGURE 68. » Planta 16-35 cm alta, foliis (2-)3 inferioribus duo- bus basalibus linearibus, marginibus peii incras- satis, caule erecto saepe ramoso, spica (1—)3-8 fl floribus hypocrateriformibus roseis, tubo aaah (10-)15-22 mm longo, tepalis 22-30 mm longis, fila- mentis inaequalibus declinatis 11-20 mm longis, uno mm breviore quam aliis, antheris 5-6 mm longis. Plants 16-35 cm high. Corm globose, 10-12 mm diam., tunics brown, more or less imbricate, woody, cut below into segments and drawn into short points above. Cataphyll membranous, ap- parently solitary. Leaves (2-)3, linear, 1.5-4 mm wide, with thickened margins and midrib, 2-grooved on each surface, sometimes sticky be- low and with soil adhering, the lower one to two leaves basal, reaching to about the base of the spike, the uppermost inserted on the lower part ofthe stem and sheathing for some dist Stem erect, smooth, bearing in addition to the cauline leaf, 1 or 2 short sheathing bracts, simple or 1— 3-branched, flexed at the base of the spike in bud. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Spike (1-)3-8-flowered, flexuose; bracts 13-16 mm long, herbaceous, flushed red above. Flower zygomorphic, the stamens and style declinate, deep pink with dark markings at the base of the tepals, pale in the tube; perianth tube (10-)15- 22 mm long, une slightly longer to about twice as | e bracts; tepals 22-30 mm long, 6.5-9 mm wid Filaments unequal, unilateral and declinate, two 11-20 mm long, one 4-5 mm shorter than the others; anthers yellow, 5-6 mm long. Ovary 2-3 mm long, style dividing near the apex of the anthers, the branches 4-5 mm long, recurved. Capsule obovoid to globose, 5-8 mm long, enclosed in the bracts. Chromosome num- ber, 2n = 26 (Goldblatt 6828). Flowering time. Late November to early Jan- ry. Distribution. Mid to upper elevations in the Slanghoek, Hottentots Holland, Riviersonde- rend and Langeberg Mountains, in rocky habi- tats. Figure 68. Geissorhiza grandiflora is endemic to the mountains of the southwestern Cape. It has been collected on isolated high areas from Great Win- terhoek Mountain and Bains Kloof in the north to the Langeberg at Swellendam in the east. The relatively few collections suggest that it flowers irregularly, perhaps only after mountain fires. It has one of the largest flowers in the genus, com- parable in size with those of some species of section Engysiphon (subgenus Weihea). Despite its large and zygomorphic flowers with declinate stamens and style, unusual in section Monticola, G. grandiflora has the imbricate corm tunics, un- equal stamens and branched stems characteristic of this section and is assigned here rather than to section Engysiphon. It resembles most closely the smaller flowered G. burchellii, a generally smaller species that can be distinguished by its leaves with heavily thickened margins and mid- rib and actinomorphic flowers with tepals in the 12-18(-22) mm range ane a penent tube 4-6 mm long. Geissorhiza b t common in the Langeberg, but has been recorded to the west in the Hottentots Holland and Slanghoek Mountains as well. The two species are occa- sionally sympatric or nearly so and both have been collected on the Cossacks, Slanghoek Mountains, by Elsie Esterhuysen at the same time some years ago. A collection from Great Winterhoek, Phillips 1886, is tentatively assigned to Geissorhiza gran- diflora. It appears to have somewhat smaller 1985] GOLDBLATT — GEISSORHIZA 409 FIGURE 68. Pesci and distribution of Geissorhiza podium Fruiting plant and corm x0.5; spike with flower g Bains Kloof). flowers than are typical, but it is uncertain wheth- er the poorly pressed blooms were incompletely open or shrunk on drying, or whether this represents a form that eis has flowers with shorter tepals. Also included are collections from the Riviersonderend Mountains, Esterhuy- sen 35131, and the Langeberg, Wurts 543. These comprise plants with somewhat atypical flowers, seemingly larger and with a wider perianth tube. flowering Esterhuysen 35832, both Limietberg, The Esterhuysen collection clearly has yellow coloring in the throat of the flower. More collec- tions from here are needed to determine whether this form represents a separate species. Specimens examined. H AFRICA. CAPE: 3319 (CA), Esterhuysen 23997 (BOL, G); Baviaans Kloof off Bains Kloof, 3,000 ft., Lewis 770 410 or s.n. (BOL, NBG, PRE, SAM); N side of rag asta Wemmershoek Tafelberg, rugged slope, 5,000 ft., Es- terhuysen 35863A (MO). 0 (Montagu) Crown Mt., above Swellendam, he new power 984); p E side nadendal valley, low wet cliffs, ca. 3,000 ft. (BA), a 35131 (B 62. Geissorhiza tabularis Goldbl., sp. nov. TYPE: South Africa. Cape: Table Mt., between Skeleton and Window Gorges, Goldblatt 6726 (holotype, MO; isotypes, K, NBG, PRE, S, US, WAG). FIGURE 69. Geissorhiza ramosa Es Klatt sensu G. J. Lewis, FI. Cape Penins. 255. 1950. Planta 25-35 cm alta, foliis (2-)3 pisi s duo- bus basalibus linearibus marginibus 18 mm longis, filamentis inaequalibus, longioribus duobus 5-6 mm longis, breviore 3-4 mm longo, an- theris 3.5-5 mm longis. Plants 25-35 cm high. Corm globose, tapering above, 7 mm diam., tunics light brown, more or less imbricate, papery in texture, cut below into narrow segments and drawn into points above. Cataphyll membranous, apparently solitary. Leaves (2-)3, linear, 2-3 mm wide, with slightly thickened to winged margins and midrib, and broadly 2-grooved on each surface, somewhat sticky below and with soil adhering, half to oc- casionally about as long as the stem, the lower two (or one) basal, the uppermost cauline, sheathing for half its length. Stem erect, smooth, bearing in addition to a cauline leaf, 1(—2) short, sheathing bracts, simple or 1—3-branched, flexed at the base of the inflorescence when in bud, later becoming straight. Spike 3-8-flowered, flexuose; bracts 7-8 mm long, herbaceous, flushed red above, recurved above at flowering time. Flower stellate when fully open, pale mauve to nearly white, usually becoming white after drying; peri- anth tube 1.5-2 mm long, cylindric, much short- er than the bracts; tepa/s 15-18 mm long, 5-7 mm wide. Filaments unequal, the two longer 5— 6 mm long, the shorter 3-4 mm long; anthers 3.5-5 mm long. Ovary 1.5-2.5 mm long, style dividing at the apex of the anthers, branches ca. 2.5 mm, recurved. Capsule unknown. Chro- mosome number, 2n — 26 (Goldblatt 67 26). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Flowering time. Late October to mid-De- mber. Distribution. Restricted to the lower plateau able Mountain, in very wet or marshy sit- uations in peaty sand, blooming rarely (if at all) except after fire. Figure 69. This species, collected at least as early as 1825 so treated in “Flora of the Cape Peninsula" by Lewis (1950). Neither Foster (1941) nor Baker (1896) dealt with Geissorhiza tabularis and prob- ably had no material of the species. It is endemic to Table Mountain on the Cape Peninsula and grows in wet or marshy sites. It flowers in the late spring or early summer only in the years following a veld fire Geissorhiza tabularis is related to the more common and widespread G. ramosa but is quite distinct from this species which typically has a more branched stem and many, much smaller flowers, the tepals of which are 5-9 mm long in contrast to the few branched G. tabularis in which the tepals range from 15-17 mm sere Geisso- esprea d Cape mountain species, G. burchellii, which has large, purple flowers with a longer perianth tube to 5 mm and longer filaments, the long pair being 10-13 mm long and the short one, 5-8 mm. Geissorhiza tabularis has an unusually short perianth tube, 1.5-2 mm long and characteris- tically short filaments 3-6 mm long. Specimens examined. SOUTH AFRICA. CAPE: 3318 (Cape Town above Kirstenbosch, Lewis 7 73 3 (PRE De 56034); summit of Table Mt., Zeyher s.n. (SAM Kop, shady S face, Lewis 4792 (SAM (BOL, NBG); Waai Vlei, Wolley Dod 2146 (BM, BOL, K in part) 63. Geissorhiza ramosa Ker ex Klatt, Linnaea 34: 657. 1866 (perhaps intended as a com- bination but no basionym cited and thus South Africa. Cape: Tulbaghskloof, Tul- baghsthal, am Fuss des Winterhoeksberg, am Witsenberg und bei Vogelvalei, Ecklon & 1985] GOLDBLATT — GEISSORHIZA ZZ 74 A MAE M and distribution of Geissorhiza tabularis. Habit and corm x0.5; flower life size FIG 9. (Goldblatt PUR Table Mt. Zeyher Irid. 229 (77.11) [lectotype, B, des- ignated by Foster Ar py dae B, BM, E, FI, G, GH, LD, MO, P, PRE, SAM, W, Z], Perm p Zeyher me 234 (syntype, not seen). un ar montana Foster, Contr. Gray Herb. 135: 55. 1941. TvPE: South Africa. Cape: Genadendal m : 3,000—4,000 ft., Drége s.n. “G. imbricata a” (holotype, B; isotypes, BM, E, G, K, L, MO, S). Plants 20-45 cm high. Corm globose, tapering above, 6-8 mm diam., tunics brown, woody or more often more or less papery in texture, incised below into regular sections. Cataphyll membra- nous, solitary. Leaves 2-3(-4), linear, 1-2.5 mm wide, the margins and midrib thickened and nar- rowly 2-grooved on each surface, half as long, to about as long as the stem, the lower one or two leaves basal, the upper one Do two) cauline, and sheathing the stem bel Stem erect, smooth, with 2-4 branches an simple), usu- ally bearing 1 cauline leaf and short bracts in the axils of the branches. Spike few to several (to 1 1)-flowered, inclined, flexuose; bracts (3-)4—5.5 412 mm long, herbaceous, or becoming dry and membranous from the apex. Flower evidently stellate and actinomorphic (not seen live), blue, purple, or violet (fading to pink then white when dry); perianth tube short, 2-3 mm long; tepals (5-)7-9 mm long, 3-4.5 mm wide, obovate. Fil- aments unequal, the two longer 3-4 mm long, the other shorter by 0.5-1.5 mm; anthers 2-3.5 mm long, yellow. Ovary 1-2 mm long, style di- viding towards the apex of the anthers, branches ca. 1.5 mm long, recurved. Capsule globose to oblong, 3.5-5 mm long. Chromosome number, unknown Flowering time. October to December (to January at high elevations). Distribution. Mountains of the southwestern Cape from Groot Winterhoek in the north to the Langeberg near Riversdale in the east, often in marshy or wet sites; frequent after fires. Figure 70. Geissorhiza ramosa is relatively common in the mountains of the southwestern Cape but it is not often collected, probably because it grows mainly at high altitudes and also due to its habit of flowering well only after fires. It is a most stem; small, short-tubed flowers, and unequal make confusion with other species un- likely. It is most closely related to the Cape Pen- insula endemic, G. tabularis a species with much larger flowers, but a shorter perianth tube. The relationships of G. ramosa are discussed more fully under this species. Specimens collected at Vogelgat near Her- manus, Schlechter 9577 and in Bains Kloof, Grant s.n., are unusually short and have mostly un- branched stems and seem at first rather different from other collections of G. ramosa. The spec- imens match exactly the smaller individuals of collections such as Esterhuysen 33703, and there seems no reason to regard this material as any- thing but depauperate G. ramosa. hen Klatt described Geissorhiza ramosa he attributed the name to Ker, although as far as is known, Ker never knew this species. Klatt per- haps cited Ker in error or possibly intended to make a new combination based on Ixia ramosa Ker, but if so he neglected to cite a basionym, as was his normal practice. Given this situation, Foster (1941) suggested that G. ramosa be re- garded as a new species described by Klatt, al- though he was aware that some 30 years later ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Klatt (1895) cited Ixia ramosa Ker in the syn- onymy of G. ramosa when he subsequently dealt with this species. Foster suggested that after this period of time this action could not serve to in- dicate that G. ramosa was originally a new com- bination. I propose to follow Foster's treatment of G. ramosa. Foster also recognized Geissorhiza montana, here included in G. ramosa. He knew this only from the type collection, a population of small plants, which he considered distinct from G. ra- mosa, on account of its crowded basal leaves "with a different type of venation" and soft, im- n. “G. imbricata a,” have only two basal leaves, as in all the specimens he placed in G. ramosa, with margin and midvein only slightly less thickened than in those of other spec- imens. The corm tunics differ, in my estimation, hardly at all from many other collections of typ- ical G. ramosa. Geissorhiza montana is therefore reduced to synonymy in G. ramosa. Specimens examined. SOUTH AFRICA. CAPE: 3318 (Cape Town) Jonkershoek (DD), Compton 15310 NBG). 3319 (Worcester) Winterhoeksberg (AA), Ecklon & Zeyher Irid. 227 (72.11) (B, LD, P); Witsenberg, Tul- ,PRE,S,S bagh (* Worcester, Tulbaghskloof), Ecklon & Zeyher ae 229 (77.11) (B, E, FI, LD, MO, PRE, S, SAM, W, 2 edel ea Esterhuysen mo? (BOL, NBG): mts. near Michells (BOL); Steenboksberg, Bains Kloof, marshy stream bank near **Veepos," Esterhuysen 33703 (BOL, K, MO, S, US, WAG); Bains Kloof, Grant s.n. (LMU, MO, WAG); Witte River ih (BC E Thorne. s.n. (SAM 46520), Wasserfall61 BG nme farm Eendracht, A (CB), Pica Survey ; mt. slopes S of Wemmershoek, 2,000 ft. (CC) uL. 1309 (SAM); Adolph's Kop, French Hoek Esterhuysen 11198 (BOL, PRE); ridge from French Hoek Pass towards Paarde Kop, Esterhuysen 35862 (MO); Zachariashoek, Kalsteelkloof, Smith 46 (STE); Villiersdorp (CD), Esterhuysen s.n. (BOL 31883). 2 8495), ip di 4679 (PRE). 3321 (Ladismith) dala) e River farm, Riversdale (CO), Muir 5364 (PRE); ‘Valley River, on the Lange Bergen near K , Bur chell 7044 (K). 3418 (Simonstown) Somerset Sneeukop (BB), Sto- koe s.n. (SAM 49610); E slopes of Landrost Kop, Hot- GOLDBLATT — GEISSORHIZA + ramosa . bryicola . scopulosa . ciliatula MLE Ficure 70. Morphology of Geissorhiza bryicola and distribution of G. ramosa, G. bryicola, G. scopulosa, and G. ciliata. Habit x0.5; flower and gynoecium x1.5 (Goldblatt 7032, Vogelgat, Hermanus). tentots Holland, Thorne s.n. (SAM 51545); slopes above Steenbras Siding, Thorne s.n. (SAM 50416); Leopard Gorge, Betty's Bay, forest edge (BD), Boucher 1704 STE). 3419 (Caledon) Lebanon, burned area, 3,000 ft., Kruger 227 (PRE, STE); Vogelgat (AD), Schlechter 9577 (B, BM, BR, E, G, K, L, , P, PH, PRE, Z), Goldblatt 6741 (MO); mts. above Genadendal (BA), Drége s.n. (B, BM, E, G, K, MO, S); Rivier Zonder End Mts. (BA- BB), Stokoe s.n. (SAM 57329); Pilaarkop above Lin- deshof, path below summit, 4,500 ft. (BB), Esterhuysen 70 , MO, NBG, PRE, S, US, WAG). 3420 (Bredasdorp) limestone hills, Potteberg, De Hoop (AD), Burgers 1197b (PRE). WITHOUT PRECISE LOCALITY: Caledon, Pappe s.n. (K); C. B. S., Roxburgh s.n. (BM). 64. Geissorhiza bryicola Goldbl., sp. nov. TYPE: South Africa. Cape: Vogelgat Nature Re- serve, Hermanus, along the path at Psoralea Drip, Goldblatt 7032 (holotype, MO; iso- types, K, NBG, PRE, S, WAG). FIGURE 70. Planta 15-30 cm alta, cormo subgloboso, 5-8 mm diam., tunicis duro-membranaceis infra incisis, foliis 3(-4), inferioribus duobus basalibus linearibus, l reclinato interdum ramoso, spica 2-6 florum, bracteis herbaceis, 4—6(-8) mm longis, floribus stellatis albis, tubo ca. 2.5 mm longo, tepalis 8-10 mm longis, fila- mentis inaequalibus, longioribus 5-6 mm longis, bre- longis. viore 3.5-5 mm longo, antheris ca. 3 mm lo 414 Plants 15-30 cm high. Corm subglobose, sym- metric, 5-8 mm diam., tunics brown, brittle- membranous, imbricate, incised below into reg- ular segments. Cataphyll membranous. Leaves usually 3(-4), linear, weak and trailing, usually slightly longer than the stem, 1.5— m wide, nearly flat, the midrib and margins lightly raised, the lower two leaves basal, the third inserted in the lower to mid part of the stem, partly sheath- ing, occasionally bract-like and with a short free apex, the fourth leaf if present, inserted in the upper part of the stem and short, more often reduced to a scale. Stem weak, inclined to trail- ing, simple or with 1 (occasionally 2) branches, bearing 1 short leaf and sometimes a second, often bract-like shorter leaf. Spike (2-)3-6(-10)- flowered, flexuose; bracts 4-6(-8) mm long, her- baceous, the upper margins transparent. Flower stellate, white; perianth tube short, ca. mm long, usually just exserted from the bracts; tepals 8-10 mm long. Filaments unequal, the two long- er 4-6 mm long, the shorter 3.5-5 mm long; anthers ca. 3 mm long, yellow. Ovary 2-2.5 mm long, style dividing at the apex of the anthers, branches ca. 1.5 mm long, saus x iia asa bose, 3—5 mm long. Chromosome ber 26 (Goldblatt 6741), 26 + B (Goldblatt b Flowering time. September to mid-Novem- ber. Distribution. Klein River Mountains, in the vicinity of Hermanus, wet situations, in shade or on damp cliffs near seeps or waterfalls. Figure 70 Geissorhiza bryicola grows in much the same habitat as G. cataractarum, which occurs along the slopes of the mountains between Hermanus and Bettys Bay some 30 km to the west. The two are, however, unrelated, and are treated here as belonging to different subgenera. Geissorhiza bryicola has smaller and white flowers (tepals 8— 10 mm long) on spikes of 3-6 (occasionally to 10) flowers; unequal stamens; persistent, brown, imbricate corm tunic layers; and leaves with a nearly flat lamina. In contrast, G. cataractarum has large bluish flowers with tepals 14-20 mm long; spikes of 1—5 flowers; equal stamens; eva- nescent, pale evidently concentric corm tunics; and leaves with much thickened margins and midrib. Geissorhiza bryicola is closely related to the widespread western Cape mountain species G. ramosa and the two have similar flowers with short unequal stamens. Geissorhiza ramosa is a more robust species, usually with erect 2- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 4-branched stems, spikes of up to 11 flowers and leaves with thickened margins and midrib. This combined with a purple to violet flower make it unlikely to be confused with the white-flowered G. bryicola Specimens examined. SOUTH AFRICA. CAPE: 3419 (Caledon) Fernkloof Nature Reserve, Hermanus, roc face in spray of waterfall, 200 m (AD), Robertson 132 (K, MO, NBG, PRE); Vogelgat Nature Reserve, along the path to Main Falls, Goldblatt 6742 (MO, NBG), 7037 (MO, PRE, S); Vogelgat, wet S-facing rocks at Psoralea Drip, Goldblatt 7032 (K, MO, NBG, PRE, S, AG). WITHOUT PRECISE LOCALITY: Hermanus Mts., Stokoe s.n. (SAM 55711); Klein River Mts., Stokoe s.n. (SAM 65. Geissorhiza scopulosa Goldbl., sp. nov. TYPE: South Africa. Cape: Hex River Mts., Milner Ridge Peak, 4,000-5,000 ft. (AD), Ester- huysen 9328 (holotype, BOL: isotypes, NBG, PRE) Planta 6-15(-20) cm alta, foliis 3 goin dues duo bus ELE linearibus, marginibus costoque incras- i e erecto i fl fl mm longo, tepalis ntis qualibus, longioribus duobus ca. 3 mm longis, breviore 2 mm longo, antheris 2-2.5 mm longis Plants (6—)8-15(-20) cm high. Corm globose, 8-10 mm diam., tunics brown, firm to soft-mem- branous, imbricate, regularly notched below. Cataphyll membranous, brownish above. Leaves 3, linear, 1-2 mm wide, the margins raised at right angles to the lamina, the midrib thickened, the lower two leaves basal, the lowermost long- est, reaching at least to the base of the spike, sheath sticky and with sand adhering, second leaf sheathing the lower part of the stem for a short distance, the uppermost inserted in the upper part of the stem, short, sheathing for at least half its length, sometimes almost entirely. Stern up- right or inclined, simple, minutely ciliate-papil- lose, bearing in addition to a cauline leaf, a sheathing bract in the upper third. Spike secund, flexuose, 2-5(-7)-flowered; bracts 3.5-5 mm long, green to membranous in bud, sometimes reddish above, becoming dry and pale in the upper part at flowering time. Flower stellate, pale violet blue, darkening in color with age, cream in the throat and tube; perianth tube 2-3 mm long, in- fundibuliform; tepals 8-9 mm long, ca. 3 mm wide, elliptic. Filaments unequal, the two longer ca. 3 mm long, the shorter 2 mm long; anthers 2-2.5 mm long, yellow. Ovary ca. 1 mm long, 1985] style dividing near the apex of the anthers, branches ca. 1 mm long, recurved. Capsule glo- bose, to 4 mm long, not exceeding bracts. Chro- mosome number, unknown. Flowering time. November Distribution. amp rocky sites, in cliffs and under boulders, usually south-facing, at high al- titudes in the Hex River Mountains. Figure 70. Geissorhiza scopulosa is very like G. ramosa in floral morphology and general appearance, al- though much reduced in size, and the two are almost certainly closely related. Geissorhiza sco- pulosa is restricted to the Hex River Mountains, an area where G. ramosa does not occur. It is apparently confined to rocky habitats at altitudes above 1,500 m. In addition to its smaller size, Geissorhiza scopulosa can be recognized by its unbranched stems, which are minutely ciliate, an unusual character in this section. Its reduced size and absence of branching make it seem likely that G. scopulosa is a local segregate of the very widespread G. ramosa. Specimens p soi AFRICA. CAPE: 3319 (Worcester) Hex River ner Ridge peak, 4,000- 5,000 ft. (AD), ieiuna 9328 (BOL, NBG, PRE); Milner Ridge Peak, bg ge cliffs, 5,000—6,000 ft., Es- n. 9333 (BO ); Baboon opa Hex River Mts., 4,800 ft., trs 33337 (BO 66. Geissorhiza ciliatula Goldbl., sp. nov. TYPE: South Africa. Cape: Cedarberg, Middelberg, 1,500 m, Goldblatt 5144 (holotype, MO; isotypes, K, NBG, S) anta 6-12 cm alta, tunicis cormi imbricatis mol- libus, foliis (2—)3, inferiore basali falcato, marginibus ndo basali caulem v vaginanti florum, floribus stellatis "age ed pu rpurascenti- bus, tubo ca. 1.5 mm longo, tepalis 6-7 m m ngls, filamentis aequalibus, hera 5^ 2 mm lon Plants 6-12 cm high. Corm globose, 3.5-5 mm diam., tunic layers brown, more or less imbri- cate, soft-textured, incised above and below. Cataphyll membranous, often fragmented. Leaves 3 or 2, linear to falcate, to 1 mm wide, oblon in section with margins and midrib strongly thickened, thus narrowly 2-grooved on each sur- face, the lowermost leaf basal, longest, the second also basal, but sheathing the lower half of the stem and free only above, the uppermost inserted on the upper third of the stem and sheathing for at least half its length (occasionally entirely GOLDBLATT — GEISSORHIZA 415 sheathing and bract-like). Stem erect, minutely ciliate, unbranched, flexed above the sheathing part of the upper leaf. Spike 1—4-flowered, flex- stellate, white, turning purplish on fading; peri- anth tube ca. 1.5 mm long, not exserted from the bracts, widening from the base; tepa/s 6-7 mm long, obovate, to 3 mm wide. Filaments equal, ca. 2.5 mm long; anthers ca. 2 mm long, yellow. Ovary ca. 1.5 mm long, style branching at the apex of the anthers, branches ca. 1.3 mm long, recurved. Capsule globose, 3-4 mm long, shorter than bracts. Chromosome number, 2n — 26 (Goldblatt 5144). Flowering time. November Distribution. Cedarberg, above 1,400 m, in sheltered wet mossy places on cliffs and cracks in rocks, mainly south-facing. Figure 70. Geissorhiza ciliatula is a dwarf, white-flowered species from the Cedarberg and Cold Bokkeveld Mountains. In the Cedarberg it grows on the rocky peaks that rise above Middelberg Plateau. It grows in sheltered cracks in south-facing cliffs or under boulders, where shade and water are available late in the growing season. It occurs together with the very different and distinctive large pink-flow- ered G. cedarmontana in a habitat also shared with moss and Drosera sp., and occasionally Hesperantha cedarmontana, which flowers ear- lier in the season The exact nature of the corm tunics of Geis- sorhiza ciliatula is difficult to determine, as the tunics are soft-textured and not firmly fixed in position. However, they appear to be more or less imbricate and resemble those found in sub- genus Geissorhiza section Monticola. Tentative- ly, I suggest that G. ciliatula may be related to the G. ramosa alliance, perhaps closest to the fairly small and also ciliate leaved G. scopulosa which occurs in similar habitats in the Hex River Mountains, some distance to the south. Specimens examined. SOUTH AFRICA. CAPE: 3219 (Wuppertal) Cedarberg, S slopes of pct Central (AC), Goldblatt 5144 (K, MO, N S); Bokkeveld Sneeuberg, 4, 3005, 000 2 seaso damp narrow ledges below hig Finca en 36127 (BOL, K, MO, PRE, ST 67. Geissorhiza pseudinaequalis Goldbl., sp. nov. TYPE: South Africa. Cape: Witteberg, Slanghoek Mts., wet overhung cliffs on S side, Esterhuysen 22293 (holotype, BOL; isotypes, K, MO). FIGURE 71. 416 FIGURE 71. terhuysen 35281, Simonsberg). ginantibus, caule erecto, spica (22)3-7 florum, flo- ribus stellatis caeruleo-violaceis, tubo 3-5 mm longo, tepalis 12-17 mm longis, filamentis inaequalibus lon- gioribus duobus 8-10(-12) mm longis, breviore 4—6 mm longo, antheris 3.5-5 mm longis. Plants 9-18(-30) cm high. Corm globose, sym- metric, 5-9 mm diam., usually heavily bulbifer- ous at base, tunics membranous-papery, more or less imbricate, incised below, light brown, the outer layers becoming irregularly broken. Leaves usually 3 (or 4), with a sheathing stem bract as well, linear, plane, the margins and midrib slight- ly raised, occasionally sparsely ciliate on the edges, 2-4 mm wide, usually about as long as the stem, the lower two (or three) leaves basal, the upper- most inserted in the lower to mid part of the stem, and sheathing below. Stem erect, smooth, bearing 1 or more sheathing, herbaceous bracts ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 vA R Ç X M Z D Li aes — Morphology and distribution of Geissorhiza pseudinaequalis. Habit x0.5; flower life size (Es- in the upper part, and often branching at this point, and sometimes also from the axil of the upper leaf. Spike (2—)3-7-flowered, flexed at the base and flexuose; bracts 7-10 mm long, her- mauve) and usually pale in the tube, the throat sometimes ringed with darker color; perianth tube 3-5 mm long, not reaching the apex of the bracts; tepals 12-17(-20) mm long, obovate, 5-6.5 mm wide. Filaments unequal, the two longer 8- 10(-12) mm long, the shorter 4-6 mm long; an- thers 3.5-5 mm long. Ovary 2-3 mm long, style dividing near or beyond the apex of the anthers, branches recurved. Capsule unknown. Chro- mosome number, unknown. Flowering time. Mid-October to January. Distribution. Mid to upper elevations in the western Cape mountains between Ceres and Si- 1985] monsberg, on damp, sheltered, south- or west- facing rock ledges and cliffs. Figure 71 Geissorhiza pseudinaequalis is one of several species of Geissorhiza that occupy specialized niches in the south western Cape mountains. This species has a limited range from the Hex River Mountains and Bains Kloof to Simonsberg, and apparently grows in rocky places, on ledges, or cracks in cliffs where shade and moisture are available in mid summer, and it may bloom as late as January at higher elevations. It has often been confused with Geissorhiza inaequalis (section Planifolia) on account of its markedly unequal stamens and flat leaves. The corms, however, are quite different, G. inae- qualis having woody, imbricate tunics, smaller flowers and in addition, distinctively puberulent stems. Geissorhiza pseudinaequalis has papyra- ceous corm tunics, relatively large flowers and smooth stems. These two species are probably not closely allied and are here assigned to dif- ferent sections of subgenus Geissorhiza. Geiss- orhiza pseudinaequalis is probably related to other montane species with similar soft-textured imbricate corm tunics such as G. burchellii and G. tabularis. These two species also have unequal stamens but differ in their larger size and leaves with strongly raised margins and midribs. The single collection from the Hex River Mountains, Esterhuysen 15882 at BOL, is mixed, and perhaps partl brid. It consists of a spec- imen of ie bee heterostyla with its charac- teristic small flowers and brown stem bract, sev- eral specimens of P pseudinaequalis, and some putative hybrids with flowers of intermediate size and partly brown stem bracts. The leaf margins and midrib of plants of this collection are mi- nutely ciliate, also a feature of G. heterostyla. Specimens examined. SOUTH AFRICA. CAPE: 3318 (Cape Town) Simonsberg, kloof on W side near Hels- hoogte (DD), Esterhuysen 35281 (BOL, K, MO, PRE). 3319 (Worcester) Prospect Peak, Hex River Mts., side of waterfall on E n (BC), Esterhuysen 1 5882 (BOL, NBG, PRE); Bains Kloof, along road 2-3 m from hotel (CA), remi 32722 (BOL, PRE, S). Bains Kloof, L. Bolus s.n. (BOL 31895, 31909), John- son 1 (MO), Barker 4279 (NBG), Goldblatt 457 (BOL), 679 (BOL); Upper Wellington Sneeukop, ledges on W side, Esterhuysen 26546 (BOL, MO); Witteberg, Slanghoek Mts., wet overhung cliffs on S side, Ester- huysen 22293 (BOL, K, MO); Witteberg, in Sarean and vertical cracks in S-facing ledges, Esterhuys 22293A (BOL, K, PRE); kloof between Molen ip sean and Witteberg, Du Toits Kloof, Esterhuysen 24150 (BOL); mts. S of Wemmershoek (CC), Andreae 821 (PRE). GOLDBLATT — GEISSORHIZA SECTION PLANIFOLIA 11. Section Planifolia Goldbl., G. aspera Goldbl. sect. nov. TYPE: Tunicae cormi durae sich foliis plerumque 3, ensiformibus ad linearibus plan is, caule puberulo-cil- iato, baies infra herbaceis im siccis ferrugineis, ib omorphis stellatis vel declinatis, tubo brevi, saison aequalibus vel inaequalibus. Corm tunics, blackish, hard and woody, /eaves usually 3, blade plane and smooth, lanceolate to ensiform, rarely linear, stem puberulous-ciliate, bracts herbaceous below, dry and ferrugineous in the upper half, flowers stellate and actino- morphic or declinate, the tube short, stamens equal to unequal. Species. 4. 68. Geissorhiza aspera Goldbl., J. S. African Bot. 36: 303. 1970. TYPE: South Africa. Cape: Rondebosch common, Goldblatt 501 (ho- lotype, BOL; isotypes, K, NBG, PRE). FIGURE Ixia secunda de la Roche sensu Berg., Pl. Cap. 6, 360. : 7.1783 non Ixia secunda Ker sensu Ker, 1802 teer hom. illeg. (type of 1 secunda dela Roche specifically excluded), et nom. superfl. ( Gladiolus junceus Burm. f., Fl. Cap. Prod. 2. 1768, nud. Plants 8-35 cm high. Corm ovoid-globose, 3— 6 mm diam., tunics dark, often blackish, imbri- cate. Leaves usually 3, flat, linear to ensiform, erect to falcate, the margins and midrib slightly raised, smooth or sparsely puberulent-ciliate, about as long as the stem or somewhat longer, 2—5(-7) mm wide, the lower two basal, the upper basal or inserted shortly above the ground and sheathing the lower part of the stem. Stem in- clined or erect, simple, more often 1—3-branched, first flower and secund; bracts herbaceous below, dry and brown in the upper half, 6-12 mm long, spreading above, the inner smaller, 6-8 mm long. Flower stellate, usually deep blue to purple, oc- casionally white or pale bluish; perianth tube 1— YK e Yee <> M < SD FIGURE 72. size (Goldblatt 6274, Commonage at Malmes 2 mm long, infundibuliform; tepa/s 11-15 mm long, 4-6 mm wide. Filaments 3-5 mm long, one sometimes shorter by up to 1 mm; anthers erect, 3-5.5 mm long, white to pale yellow. Ovary 1.5-2 mm long, style 7-8 mm long, branches to 2 mm long. Capsule 5-6 mm long, ovoid to ro- tund. Chromosome number, 2n — 52 (Goldblatt 213, 520, 48394, s.n. no voucher, St. James, Cape Peninsula), 2n = 26 (Goldblatt 5350, 5647, s.n. no voucher, Burghers Pos, Darling). Flowering time. August to September. Distribution. Widespread, Cape Peninsula east to the Langeberg at Tradouw Pass and north into the Olifants River Valley as far as the Doorn River and Gifberg. Figure 72. Geissorhiza aspera is acommon southwestern Cape species. It is most often found on sandy soils, but also light clay or granitic ground, both on mountains and flats, and it typically grows in large numbers. It is persistent and is often one ANNALS OF THE MISSOURI BOTANICAL GARDEN 77 ⁄ A ¿Z LAT 2 /ZZzzzZz2zz=> Ca [VoL. 72 Morphology and distribution of Geissorhiza aspera. Habit x 0.5; corm and flowering spike life esbury). of the last survivors of the native flora after se- vere disturbance. Plants vary greatly in size, de- pending on soil fertility and the amount of mois- ture available, and may grow well over 30 cm high, or be stunted and less than 10 cm. The flowers are deep glistening blue to mauve in the southwestern Cape, but occasional white mu- tants occur, sometimes even in pure colonies. It is more frequent in the south, in the Caledon district, on the Cape Peninsula and locally to the north, around Darling and Malmesbury, but it extends north through the Olifants River Valley and Cedarberg Mountains to the Gifberg Moun- tains near Van Rhynsdorp. Here G. aspera is more variable, and plants with white to pale bluish flowers rather than the typical dark blue-purple are frequent. In the north of its range plants are often found in sheltered sites, sometimes in the lee of rocks or on moister south trending slopes. The stamens are often described as being equal (Foster, 1941), but careful examination of live 1985] plants shows that one filament is usually slightly shorter, and in populations in the north of its range the filaments are distinctly unequal. Geis- sorhiza aspera is unusual in the genus in being heteroploid with both diploid, 2” = 26, and polyploid, 2n = 52, populations. From the sam- ple of seven populations so far counted, there is a pattern of diploids in the north of the range and tetraploids in the south, but both cytotypes have been recorded on the Cape Peninsula. Geissorhiza aspera is closely allied to G. in- aequalis, a more robust species, which has large per Dius-n mauve flawers. M shares with G. aspera stem and broad flat jedes. In G. EIE the short filament is about half as long as the others and the bracts are green to membranous during flowering and unlike the typical dry, brown-tipped bracts of G. aspera. The nomenclature of this common species is unfortunately complex and confusing. Its history begins when Bergius published a description of a plant he called Ixia secunda in his “Plantae Capensis" in 1767, based upon specimens in his collection in Stockholm. However, in the adden- dum to this work he cites 7. secunda de la Roche (1766), in effect, but perhaps not by intention as a synonym. Bergius' 7. secunda is regarded here as having no nomenclatural validity, but is mere- ly a misapplied name. Ixia secunda de la Roche is the species currently known as G. eurystigma (Goldblatt, 1983). De la Roche's study of Cape Iridaceae, published in 1766, was largely mis- understood by contemporary and succeeding botanists and the name 7. secunda was usually applied to plants matching Bergius' description and specimens (e.g., Thunberg, 1783; Ker, 1802a, 1802b), and this situation led to considerable nomenclatural confusion when the two species named Ixia secunda were transferred to Geis- sorhiza. Ker, who actually understood that Ixia secun- da de la Roche and /. secunda sensu Bergius were different species, published descriptions and il- seii of both in Curtis' “Botanical Maga- zine.” He renamed what he thought was Z. se- posh de la Roche (i.e., the earlier epithet) 7. rochensis, citing of course 7. secunda de la Roche as a synonym. This name is therefore, nomen- claturally superfluous. It has, unfortunately, been applied in Ker’s sense ever since to the well known Cape species, now to be called G. radians (Thunb.) Goldbl. Ker (1802b) named the small, blue-flowered GOLDBLATT — GEISSORHIZA 419 and pubescent stemmed plant Ixia secunda cit- ing, of course, I. secunda sensu Bergius, but ex- cluding de la Roche's species. According to cur- rent rules of nomenclature, Ker's J. secunda is a new name and a homonym for I. secunda de la Roche. It is also superfluous, since T. pusilla Andr. was cited as a synonym. Upon transfer to Geis- sorhiza, G. secunda cannot be treated as a legit- imate new name in terms of article 72.2 as it is still superfluous, Z. pusilla not having been ex- cluded when the transfer was made. The new name G. aspera was provided by myself (Gold- blatt & Barnard, 1970) for G. secunda Ker 2 as the only legitimate synonym, the name mu be used for this well known Cape plant icd blatt, 1983). Specimens examined. SOUTH AFRICA. CAPE: 3118 hynsdorp) near Doorn River bridge, old road 7520 (N pes of plateau, paleta d arker 3613 (NBG); 20 mi. N of Citrusdal, Olifants ena Valley, Lewis 1033 (SAM); irr S Berg, Barker 5338 (N BG); pass over Kransvlei Mts., Lewis 1870 (SAM); 8 mi. S of Clanwilliam, ‘Goldblatt 266 (BOL); 14 km S of Clanwilliam, Goldblatt 2472 (MO, US); between Clanwilliam and Algeria, Goldblatt 2569 (MO, PRE, S); Olifants River Valley, N of Ron degat, Goldblatt 5647 (MO, NBG, S, US, WAG); Kip- teins Kloof (DA), van Niekerk 636 (BOL), Modder- n or River (DB), Diels 209 (B); Greys Pass, Lewi K), Hafstrom & Acocks 298 (PRE, S); near Piketberg A Barnes s.n. (BOL 31899), Barker 7564 NBG); Levant Peak, Marloth 11528 (BOL), Es- eae s.n. (MO). 3219 (Wuppertal) Pakhuis Pass (AA), Bond 590 (NBG); Middelberg, Cedarberg, shady rocky crevices (AC), Compton 6613 (NBG); Cedarberg, Uitkyk Pass, Gillett 4113 (MO); Warm Baths (CA), Leipoldt s.n. (BOL 31901); Waterfall River, Citrusdal, Barker 3775 (NBG); Platkloof, Hanekom 1236 (STE). 3318 (Cape Town) near Hopefield (AB), Bachmann 217 (Z); Darling rere B), ne r- ling, Stokoe s.n. (SA Oy 4 Z Iba ad near Porterville (BB), Wilman 715 (BOL): Porterville, Steyn 583 (NBG); Malmesbury commonage, on S slope of hill (BC), Gold- 420 blatt 6274 (MO, WAG); flats N of Hermon (BD), Gold- blatt 6411 (MO); = du (CB), Barker 806 (NBG); Hof Str, Cape Town (CD), Page s.n. (BOL); Signal Hill, Penfold s.n. (SAM 52898). Barker 406 (NBG), 703 (NBG), diri 3739 (B), Dümmer 886 (E), Salter 531 (K); Lion , MacOwan s.n. (SAM), Marloth 7394 (PRE), Schlechter 1579 (B, FI, G, W, Z); Signal Hill, W slopes, Goldblatt 213 (BOL), Camps Bay, lower slopes, Cassidy 250 (BOL); lower Blinkwater, Cassidy 12 (BO : Rondebosch common, Goldblatt 501 (BOL, K, NBG, PRE); Camp Ground, H. Bolus 4609 (BM, BOL, GH, K, Z); Bergplatze bei Capstadt, Ecklon & Zeyher Irid. 228 (64.9) (B, FI, G, GH, L, LD, MO, W, Z); slopes of Table Mt., MacOwan in Herb. Norm. Aust. Afr. 807 (B, BM, BOL, G, SAM, Z); Table Mt., A id 5034 (SAM), Ecklon 310 (B, BM, E, G, K, M, S, Z), Rogers 16973 (PRE, Z); Salt River, Zeyher 70 or SAM); near Wellington (DB), L. Bolus s.n. (BOL 31900), Moss 307 (J); Paarl Mt., Henderson 1184 (NBG), Drége P); Langverwacht, above Kuils River (DC), Oliver 47 10 (PRE, STE); Jonkershoek Valley (DD), Lewis 1611 (SAM); Swartbos Kloof, Jonkershoek, Lewis s. 58954); Klapmuts, Lewis 4451 (NBG); Stellenbosch, Prior s.n. (K), Garside 1067 (K). 3319 (Worcester) lower slopes, Great Winterhoek Mts. (AA), Thorne s.n. (SAM, PRE); Tulbagh (AC), Kassner 1306 (BR, E), Garside 1544 (K); Lakenvlei (BC), Barker 1474 (NBG); Bains Kloof (CA), Lotsy & Goddijn 879 (L); French Hoek Pass (CC), Werdemann & Oberdieck 381 (B, K, PRE), Boucher 2347 (K, PRE). 3320 (Montagu) foot of Tradouw Pass (DC), Marsh 878 (STE). 3418 (Simonstown) Simonstown (AB), Wolley Dod 582 (BOL); between Witsands and Redhill, Lewis 1074 (SAM); Chapmans Peak, van Niekerk 465 (NBG); Red- hill, W slopes, Barker 5912 (NBG); Simons Bay, Wright s.n. (K, L); Grasslands Kloof, Cape Point Reserve (AD), Taylor 8620 (STE); Cape Flats (BA), MacOwan s.n. (SAM 48596), Krauss 1394 (MO), Rehmann 1837 (BM, BR, Z), Marloth 7261 (PRE); Helderberg (BB), Parker 4257 (GH, K, N PH E) Sir Lowrys Pass, Schlechter 1579 (B); near Somerset West, Ecklon & K, rin L Bay, Hangklip (BD), ‘Gillett d (BOL): Palmiet River a Gillett 4241 (BR, LD, MO); Kogel Bay, Wer- dermann & Oberdieck 288 (B, BR, K, PRE, WAG). 3419 (Caledon) 8 mi. N of Bot River (AA), Goldblatt 293 (BOL); quarry above Onrus (AC), Horocks 19 (MO Albertyn-Caledon road, Avontuur farm, Goldblatt 4981 (MO); Hemel-en-Aarde (AD), Gillett 1403 (BOL, STE); nd s.n. (BOL 31899); roadside pia Hermanus and Stanford, Gillett 4512 (BOL, P 3420 CA ie dii near a (CA), Sidey 1831 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 (MO, S); eee eet Buffeljagsrivier, farm Midden- plaas (BA), Vlok MO). WITHOUT PRECISE LOCALITY: Cape of Good s (CBS), Lalande s.n. (B, P), iic tA (B), B. gius s.n. (B), M. & Maire (B), Harvey 867 (E) 76 ds. Commerson s.n. (G), Niven s.n. (BR), 4 rea n. (G), Bowie 403 (G), Alexander s.n. (BR, Z Men (S, UPS), Grondal s.n. (S); “Olifants ^. bei villa Brakfontein,” Ecklon & Zeyher Irid. 213 (76) B, SAM); banks of Olifants River, Schlechter 5029 (B, sun BR, E, G, GH, K, P, S, STE, Z); Herb. Burman "Gladiolus junceus" (G); Olifantsrivier, Penther 666 (Z), 679 (Z). 69. Geissorhiza inaequalis L. Bolus, S. African ard. 20: 346. 1930; Foster, Contr. Gray Herb. 135: 45-46. 1941. TYPE: South Africa. Cape: Klipkoppies 6 mi. from Nieuwoudt- ville, L. Bolus s.n. (lectotype, BOL 19256, here designated; isolectotypes, BM, BOL, K). FiGURE 73. Plants 12-25 cm high. Corm ovoid-globose, 6-8 mm diam., tunics dark, often blackish, im- bricate, bearing several cormlets around the base. Leaves usually 3, flat, ensiform, erect to falcate, e, the lower two leaves basal, the third er or ed above the ground and sheathing for some distance. Stem more or less erect, sim- ple or 1-3-branched, lightly papillose-puberu- lent, usually bearing a short leafy bract from the upper node. Spike 2-7-flowered, usually flexed below the first flower and secund; bracts herba- ceous below, drying and becoming pale an membranous above, rarely brownish at apex, 10- le: pale blue-mauve, whitish in the center; perianth tube 2-3 mm long, infundibuliform; te- pals 20-23 mm long, 4-6 mm wide. Filaments unequal, the two longer 8—10 mm long, the short- er 5-6 mm long, nearly horizontal and initially more or less unilateral; anthers inclined to hor- izontal, 5-6 mm long, one sometimes slightly shorter, white. Ovary 2-3 mm long, style divid- ing near the apex of the anthers, branches ca. 3 mm long, recurved. Capsule oblong, 5-8 mm long, ca. 3.5 mm wide, shorter than the bracts. Chromosome number, 2n — 52 (Goldblatt 129, 2 Flowering time. Late August to early Octo- er. Distribution. Heavy soils in the Nieuwoudt- ville district, often in rocky situations. Figure 73. 1985] GOLDBLATT — GEISSORHIZA 1 r: IGURE 73. Morphology and distribution of Geissorhiza inaequalis. Habit x 0.5; flower life size [Nieuwoudt- ville Reserve (ex hort.), no voucher]. Geissorhiza inaequalis is most closely related to the very common and widespread G. aspera (syn. G. secunda) and it is sometimes difficult to distinguish them when dry. Both have a similar habit, flat ensiform leaves and puberulent stems. Geissorhiza aspera typically has deep blue-vi- olet, less often whitish, flowers and at flowering time the upper parts of the floral bracts are often dry and rusty brown. It usually has equal or sub- equal stamens, but populations with one shorter stamen are not uncommon. In contrast, G. in- aequalis has larger, pale blue-mauve flowers, typ- ically green or pale, membranous bracts at flow- ering time, the brown coloration developing only after the flowers fade. It also has one filament about half as long as the others and even one anther may be slightly shorter then the other two. In G. inaequalis, the stamens are inclined to hor- izontal and nearly unilateral while they are erect and symmetrically disposed in G. aspera. Geis- sorhiza inaequalis may also be confused with G. heterostyla L. Bolus [syn. G. rosea (Klatt) Foster], which is similar in habit, and has one short fil- ament. The latter, however, has leaves ciliate on raised margins and midrib, usually a glabrous stem and it typically has a short dry bract leaf in the upper part of the stem. Dry material must be examined carefully to avoid misidentification. The species is unusual in the genus in being tetraploid, 2n = 52. It is restricted to heavier soils 422 in the Nieuwoudtville district where it occurs at the edge of the Karoo, often in rocky sites. Specimens examined. SOUTH AFRICA. CAPE: 3119 us s.n. (BM, BOL 19256, K); a farm, Noui dE Oliver & Mauve 30 (PRE, STE); near the top of Van Rhyns Pass, Goldblatt 269 (BOL); Grasberg road, 4 mi. from Nieuwoudtville, Goldblatt 275 (BOL); 5 mi. from Nieuwoudtville, Barker 9560 (NBG); Nieuwoudtville Reserve, among dolerite boul- ders in red clay, Perry & Snijman 2360 (NBG); Uit- komst farm, SW of Nieuwoudtville (CA), Barker 10733 ). 70. Geissorhiza monanthos Ecklon, Topogr. Verz. Pflanzensamml. Ecklon 21. 1827; Baker, Fl. Cap. 6: 71. 1896, in syn. sub G. rochensis, Foster, Contr. Gray Herb. 135: 39. 1941 [as G. monantha (Thunb.) Ecklon]. Ixia monanthos Thunb., Fl. Cap. Ist edition hom. illeg. non de la Roche (1766) (= Spar- axis sp.). TYPE: South Africa. Cape: exact locality unknown, Thunberg s.n. [lectotype, Herb. Thunb. 975 UPS, designated by Gold- blatt (1982)]. FIGURE 74. Geissorhiza ere MacOwan, J. Linn. Soc., Bot. 25: 393. 0; Baker, Handb. Irid. 156. 1892, Fl. 810 Apte SAM; isolectotypes, B, BM, BOL, M). Geissorhiza rochensis var. monantha (Sweet) Baker u Baker, genns hs 156. 1892, Fl. Cap. 6: '1896, excl. basi Geissorhiza lewisiae Foster, Contr. Gray Herb. 135: 941. TYPE: South Africa. Cape: Langebaan, i s.n. esas, p Bolus 20303 in K; isotype, BOL), syn N.B. si eps dederim Sweet (1830) is a different species, base a different type. It is regarded here asahomonym of. Gm onanthos and is currently known as G. eurystigma. Plants (6—)10-20 cm high. Corm globose, sym- metric, 5-10 mm diam., l cormlets around th ensiform, plane with the midrib distinct, (1—)2.5- 4 mm wide, usually slightly shorter than the stem, the lower two leaves basal, the upper cauline, decreasing in size and sheathing below, the up- permost sometimes bract-like, smooth entirely or ciliate to pubescent along the margins and sometimes other main veins. Stem erect, flexed ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 at the base of the spike, minutely puberulous, n ke 1-4(-5)-flow- baceous in bud, becoming dry and brown above as the flowers open, folded outwards in the upper part, the inner slightly smaller than the outer. Flower either dark blue-violet and pale yellow in the center and the tube or sometimes red in the lower part of the tepals, stellate or slightly cupped, zygomorphic with upper tepal tilted downwards, and stamens declinate, held above upper tepal, i style lying under the stamens; perianth tube ca. 2 mm long, d tepals obovate, 14-18 mm long, 9-12 mm wide. Fil- aments 6-8 mm long, usually equal, aed one slightly shorter, declinate, held above the upper tepal; anthers unilateral, red-brown or yellow, 4.5-5.5 mm long. Ovary ca. 2 mm long, style decumbent, lying below the stamens, branching at mid anther level, branches recurved. Capsule ovate, 7-8 mm long, enclosed in the bracts. Chromosome number, unknown. Flowering time. Late August to mid-Octo- ber. Distribution. Southwestern coastal belt from the foot of the Hottentots Holland Mountains to the Vredenburg-Saldanha area, with outlying populations in the Olifants River Valley near Citrusdal: on sandy or rocky soils, mostly of granitic origin, sometimes growing in cracks in granite rocks. Figure 74 This relatively common and conspicuous west coast Geissorhiza has been known at least since it was collected by Thunberg in the 1770s. It remained undescribed until 1811 when the first edition of Thunberg’s “Flora Capensis” was pub- lished. Unfortunately the name Thunberg se- lected, Ixia monanthos, was a homonym for a species described in 1766 by Daniel de la Roche, y well known name monanthos can be preserved in its current sense. Much later, in 1890, MacOwan described G. bel- lendenii quite independently, based on plants he collected near Mamre, at Groene Kloof. At the time MacOwan was evidently unaware of the existence of the name G. monanthos for this species, and was concerned in drawing attention to the differences between his plant and G. ra- dians (then known as G. rochensis Ker). 1985] GOLDBLATT — GEISSORHIZA 423 ZZ ; VE “ral CAD Gey FIGURE 74. Morphology and distribution of Geissorhiza monanthos. Habit x 0.5; flower life size; leaf section enlarged (Goldblatt 6286, Mud River road, near Darling). Geissorhiza monanthos is closely allied to G. aspera (G. secunda sensu auct.), but also resem- bles fairly closely the unrelated G. radians. Geis- sorhiza aspera has a similar puberulous stem and flat leaf lamina, but smaller and actinomorphic flowers, while G. radians has the zygomorphic flower found in G. monanthos, but different floral markings, a more cupped flower, smooth stem, and narrow 4-grooved leaves, the upper of which has a strongly ribbed sheath. Plant size and leaf width vary considerably in Geissorhiza monanthos. Plants receiving little rainfall or in dry situations are short, with soli- tary flowered spikes and filiform leaves while plants growing in seasons of generous rainfall or in favored sites can be to 25 cm high with broad leaves and several flowered spikes. A single pop- ulation can yield plants covering the whole range of variation for these characters (e.g., Barker 3597, Goldblatt 2705). Populations in the Darling-Mamre area have flowers that are blue and red or blue with a yellow center ringed in dark purple (Margin 1168, Barker 724, Loubser 954, Goldblatt 6286, 6272), the type of Geissorhiza bellendenii matching the blue- and red-colored form. These plants seem to differ in no other way from pure blue-flowered plants and are treated here as the same species. This bicol- ored flower is like that found in the related G. radians with which G. monanthos occasionally grows (Barnard s.n., SAM 68439). The similarity is probably an example of color mimicry, a com- mon phenomenon in the Cape flora, or less likely to hybridization and introgression. Foster (1941) recognized as a distinct species Geissorhiza lewisiae, here reduced to synonymy. He distinguished the species on the basis of the type having slightly unequal filaments, and he grouped it with species such as G. inaequalis which also has unequal filaments. I have ex- amined the type, collected by G. J. Lewis at Langebaan, and the only collection of G. /ewisiae 424 known to Foster and it is clear that one of the filaments is shorter than the other two, but the difference is no more than 0.5 mm, and this small difference can also be detected in several collec- tions of G. monanthos. What seems more re- markable about this collection is that the leaves have a conspicuous, fine pubescence on the mar- gins and major veins. This condition is unusual in G. monanthos but can be seen in some col- lections of the species from the north of its range (Goldblatt 2705, 4084) where it is developed to a lesser degree. There seems little in fact to dis- tinguish G. /ewisiae and the species is here re- duced to synonymy. In “Flora Capensis,” Baker (1896) treated G. monanthos as a variety of G. radians (as G. ro- chensis), confusing it with G. eurystigma L. Bo- lus, while recognizing as distinct G. bellendenii. There is no doubt that Baker was mistaken in considering this a different species. The confu- sion is probably based on his inadequate knowl- edge of the type material of G. monanthos rather than any fundamental difference between it and the type of G. bellendenii. Specimens examined. SOUTH AFRICA. CAPE: 3217 (Vredenburg) Witteklip, near B cuu rg (DD), Lewis 1039 (SAM), Compton 1 NBG), Leishion 597 (BOL), Goldblatt 5846 un TR US); near Vreden- burg, Lewis 1041 3218 (Clanwilliam) s a; Cove e pi 1038 (BOL, PRE, SAM), Leighton 601 (BOL, K, PRE). 3219 (Wuppertal) 2 mi. N of Citrusdal, odds Riv- er Valley (CA), Lewis 1415 (SAM); 5 mi. N of Citrus- dal, Lewis 5201 (NBG), nemine 20761 (NBG); Cit- rusdal, Barker 3597 (NBG). 3318 (Cape Town) Olifants Kop, E of ene (AA), Goldblatt 2705 (BR, MO, PRE, S, US, WAG); Postberg, Lewis 5244 (NBG), Goldblatt 4084 (MO); Langeban, Lewis s.n. (BOL 20303, PRE); Waylands, Darling, Barker 274 (NBG), Darling commonage, Barker 8045 (NBG); Darling, Loubser 954 G), Stokoe s.n. (SAM 56869), Salter 1350 (BM); Darling Hill, Goldblatt 6272 (MO); granite outcrop- ping, Mud River road, Goldblatt 6286 ( (K, MO, PRE); eed from Darling on Hopefield road, Barnard s.n. , K); Oudepost, between Malmesbury and Hopefield (BA), Salter 3873 (BM, BOL, K); Ab- botsdale Station (BC), Martin 1168 (NBG); 33 mi. N of Cape Town on Malmesbury road (DA), Gillett 4146 (MO); 5.3 mi. NW of Klipheuwel, Acocks 20644 (PRE): t sandy flats, Lewis 5902 (NBG); Hercules Pillar, Leighton 556 (BOL, K). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 3418 Ry. Hottentots Holland (BB), Zey- her s.n. o 67). HOUT PRECISE LOCALITY: Malmesbury road, L. Bolus s.n. (BOL 31902); Cis Siebold 577 (B), Masson 71. Geissorhiza tulbaghensis F. Bolus, Ann. Bolus Herb. 2: 160. 1918; Foster, Contr. Gray Herb. 135: 40-41. 1941. TYPE: South Africa. Cape: Tulbagh, L. Bolus s.n. [lecto- type, BOL 14851, designated by Foster (1941); isolectotypes, K, PRE]. FIGURE 75. iens 6-15 cm high. Corm globose, symmet- , truncate at the base, 5-8 mm diam., tunics imbricate, blackish, the outer layers regularly ed below into truncate sections. Cataphyll UDINE often broken. Leaves 3-4, ensi- form to lanceolate, rarely oblong, plane, (1.5—)2- 6(-10) mm wide, shorter to slightly longer than stem, often sticky below and with soil adhering, the lower two or three leaves basal, the two low- erect, minutely puberulent, flexed above the leaf sheaths, simple or with 1 or 2 branches from the axils of the cauline or basal sheathing leaf. Spike 1, rarely 2-flowered, flexed at the base; bracts 10-18 mm long, green below, brown and dry above, the inner shorter than the outer. Flower zygomorphic, the stamens unilateral and decli- nate, held over the horizontal dorsal tepal, the style lying under the stamens, tepals white with maroon-red to brownish markings towards the base; perianth tube 4-5 mm long, infundibuli- form; tepals 18-22 mm long, the outer 10-13 mm wide, the inner 8-11 mm wide. Filaments 10-12 mm long, equal, unilateral and declinate; anthers 3—4.5 mm long, erect, contiguous, pollen dark red-brown. Ovary ca. 4 mm long, style dec- linate, lying under the stamens, branches ca. 5 mm long, recurved. Capsule 6-8 mm long, short- er than the bracts, ovate-oblong. Chromosome number, 2n — 26 1B (Goldblatt 5227). Flowering time. August to mid-September. Distribution. Heavy, often stony clay soil, in renosterbosveld on the flats at the foot of the mountains between Wellington and Porterville, and in the Tulbagh Valley. Figure 75. Geissorhiza tulbaghensis is one of the most beautiful species of the genus, having large white 1985] ZÀ w MdB GOLDBLATT — GEISSORHIZA EN FicunE 75. Morphology and distribution of Geissorhiza tulbaghensis. Habit and corm x 0.5; flower life size; leaf section enlarged (Goldblatt 4755, Tulbagh Cemetery). flowers marked in the center with maroon-red to brown. It is fairly rare today, and its former range is becoming restricted to fewer sites as the fertile clay flats where it grows are planted to vines or cereals. It appears similar to the well known **wine cup," G. radians, and shares with it very similar floral morphology, having large zygomorphic flowers with declinate stamens and style. They are, however, unrelated, G. tulba- ghensis having plane leaves and a puberulous stem typical of section Planifolia while G. ra- dians has the linear, 2-grooved leaves and ribbed leaf sheaths characteristic of section Geissorhiza. in its flower coloration, but in its flat leaves which have a peculiar, slender petiole-like base, and puberulous stem. It is worth horticultural atten- tion, and is easy to grow. Loubser has recorded an interesting case of sympatry and hybridization involving Geisso- rhiza tulbaghensis on a farm NW of Porterville where this speci together with G. inflexa and G. aspera. Hybrids between both the latter and G. tulbaghensis are present and exhibit var- ious degrees of intermediacy between the parents [Loubser 2185, 2187, 2188 (BOL)]. Specimens examined. SOUTH AFRICA. CAPE: 3218 (Clanwilliam) Nurust farm, 8 mi. NW of Porterville (DD), Loubser 2103 (NBG), 2190 (BOL). 3318 (Cape Town) Wellington (CB), Dawson s.n (BOL 14011), Grant s.n. (MO). 3319 (Worcester) Tulbagh (AC), F. Bolus & L. Bolus s.n. (BOL 14851, K, PRE), Grant 2423 (MO, PRE), Zeyher s.n. (SAM 20866), Stanford s.n. (BOL); 2 mi. S of Tulbagh, Lewis 5793 (NBG); Tulbagh Cemetery, Goldblatt 4755 (K, MO, PRE, US), j ; PRE); 3 mi. W of Tulbagh, Marloth 7113 (PRE); Ar- tois, S of Tulbagh, H. Bolus s.n. (BOL 7585); foot of Vogelvlei Mts., near Gouda, Esterhuysen 18798 (BOL); 426 ed oe Road Station (Gouda), Diels 1164 (B); bagh turnoff, from Hermon, Mauve 4645A (PRE, TE) WITHOUT PRECISE LOCALITY: Cape, Boucher s.n. (B “Herb. Lubeck”). SECTION CILIATA 12. Section Ciliata Goldbl., sect. nov. TYPE: G. inflexa (de la Roche) Ker. Geissorhiza subsection Pubescentes Foster, Contr. Gra Herb. 135: 39. 1941. TYPE: G. eros a (Salisb.) Foster [= G. inflexa (de la Roche) Ker]. Tunicae cormi durae lignosae, foliis plerumque 3, ciliato- o iiu caule glabro vel puberulo-cilia- tis, bracteis = herbaceis supra siccis ferrugineis, flo- us actinomorphis stellatis, tubo brevi, aa ice aequalibus vel inaequalibus Corm tunics hard and woody, /eaves with the margins and midrib moderately to strongly raised and winged, the midrib and sometimes other veins also winged, the edges ciliate-pubescent, ensiform to linear, bracts herbaceous below, d and ferrugineous in the upper half, flowers stel- late or hypocrateriform, actinomorphic, the tube short or reaching to the apex of the bracts, sta- mens equal or unequal . Species. 10. 72. Geissorhiza namaquensis Barker, Fl. Pl. Af- rica 18: tab. 688. 1938; Foster, Contr. Gray Herb. 135: 51. 1941. TYPE: South Africa. Cape: Namaqualand, Klipfontein, Phillips s.n. (holotype, BOL 22163) Plants (8-)12-30 cm high. Corm globose, 8— 10 mm diam., somewhat asymmetric with one side obliquely flattened below and extended downward shortly, tunics dark brown, imbricate, notched below into more or less regular seg- ments, and drawn into points above. Leaves usu- ally 3, occasionally 4, about as long as the stem, 2-4 mm wide, evidently flat, but the margins raised and extending for a short distance at right angles to the blade and with ciliate edges, midrib also raised and winged and ciliate on the edges, sometimes a second pair of veins also ciliate, the lower two leaves basal, the third leaf inserted above the ground and sheathing below, similar but shorter than the basal leaves, fourth leaf, if present, shortest and becoming bract-like. Stem more or less erect, flexed at the base of the spike, minutely and sparsely puberulous below, some- times with 1 branch. Spike inclined, flexed at the ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 base, flexuose, 2-8-flowered; bracts green, her- baceous, dry and membranous, pale becoming brownish only near the apex, (92)12-16(-20) mm long, the outer completely enclosing the smaller inner bract. Flower hypocrateriform, blue-pur- ple; perianth tube (6—)8-11 mm long, reaching the apex of, or just exceeding the bracts, cylin- dric, widening near the throat; tepals 14-18 mm long, elliptic, to 4 mm wide. Filaments equal, 5— 6 mm long; anthers (2-)4—6 mm long, yellow. Ovary 2-4 mm long, style dividing near the mid- dle or towards the apex of the anthers, becoming unilateral, branches 1-3 mm long, recurved. Capsule globose, to 8 mm long. Chromosome number, unknown. Flowering time. September to October. Distribution. Northern Namaqualand, in Renosterbos type vegetation on shale and quartz- ite slopes. Figure 7 Geissorhiza namaquensis and its more spe- cialized, long-tubed relative, G. kamiesmontana are a closely related pair of species and the only members ofthe genus endemic in Namaqualand. Although at first glance Geissorhiza namaquen- sis resembles the Western Cape G. aspera (syn. . secunda sensu auct.), its winged and ciliate leaf margins and midrib and its well-developed perianth tube suggest a very fundamental differ- ence between them. Geissorhiza namaquensis and C. kamiesmontana are most likely related to species of section Ciliata where similar leaves are found. However, they seem taxonomically isolated in the section and probably represent a fairly old line, perhaps close to the ancestral type of the section, while the remaining species have evolved more rapidly and become specialized in their very short perianth tubes and dry bracts that are brown by the time the flowers open rath- er than drying after the flowers fade as in G. namaquensis. Geissorhiza namaquensis occurs locally on the Kamiesberg on granitic soil, but elsewhere in Na- maqualand is restricted to the few outcrops of shales and quartzites of the ancient Nama sys- tem, which once covered Namaqualand, but is now largely eroded away. All collections away from the Kamiesberg have been made in the north-south trending belt BE Nama Shale that occurs west of S somewhat beyond Steinkopf. This soil, very dif- ferent from the granitic sands typical of Nama- qualand, supports an interesting flora which shows many links with the Cape Flora to the 1985] south (Goldblatt, 1979a: 388). It supports several Iridaceae either occurring elsewhere only in the Cape Region, or endemic, but having their clos- est affinities with Cape taxa. Specimens examined. SOUTH AFRICA. CAPE: 2917 (Springbok) Klipfontein (B-), Phillips s.n. (Nat. Bot. Gard. 1471/30 in BOL, K); Steinkopf (BA), Marloth 6782 (B, BOL, PRE, STE), Herre s.n. (STE 11844); stony hillside, Steinkopf, Lewis 5495 (NBG); Kosies PRE, WAG); 15.25 mi. W by So terveld on shale and quartzite hills, Acocks 79580 (K, M, NBG, PRE). 3018 (Kamiesberg) Leliefontein, foot of Kamiesberg Mt. (CC), Le Roux & Ramsey 672 (PRE). 73. Geissorhiza 1 i t TYPE: South Africa. Cape: Kamiesberg, E slopes of Rooiberg near Welkom, Goldblatt 5770 (holotype, MO; isotypes, K, NBG, PRE, WAQ). Goldbl., sp. nov. Planta (10-)1 5-30 cm alta, foliis 3—4, inferioribus duobus basalibus, marginibus elevatis, pubescentibus, costis centralibus elevatis alatis pubescentibus, caule pubescente, spicis florum 1-4, floribus hypocraterifor- mibus, atrocaeruleis, tubo 18-25 mm longo, tepalis 16-20 mm longis, stylo diviso prope basem anthera- m. Plants (10-)15—30 cm high. Corm globose, ca. mm diam., somewhat asymmetric with one side flattened obliquely below and sometimes ex- tending downward, tunics dark brown, imbri- cate, notched below into nearly regular segments and drawn into points above. Cataphyll 1 or 2, membranous, brownish. Leaves 3 or 4, ensiform, 2-3 mm wide, reaching to about the base of the spike, the margins lightly raised and extending at right angles to the blade, the midrib also raised and winged, the edges of the margins, midrib and other veins pubescent, the lower two leaves bas- al, the upper of these sheathing the stem shortly, the upper one or two cauline, the fourth leaf (when present) smallest and becoming bract-like. Stem erect, puberulent-pubescent throughout, including the spike axis, 1-2-branched. Spike 1- 4-flowered; bracts green, becoming dry from the apex, eventually turning brown above, 12-18 (-21) mm long, the inner completely enclosed by the much longer outer bracts. Flower hypocra- teriform, deep blue, with pale throat and tube; perianth tube 18-25 mm long, cylindric below, widening gradually above the bracts; tepals 16— 20 mm long, elliptic, ca. 3 mm wide. Filaments 8-10 mm long, equal; anthers 3-4 mm long, yel- GOLDBLATT — GEISSORHIZA Bia ee he ae oe 4 IT T E h A = y VY G. kamiesmontana f. € G. namaquensis x {| Hig J 2 Z FiGugRE 76. Distribution of Geissorhiza nama- quensis and G. kamiesmontana. low. Ovary ca. 3 mm long, style dividing near the base ofthe anthers, branches ca. 2.5 mm long, recurved, and reaching the middle of the anthers only. Capsule oblong to globose, 6-9 mm long. Chromosome number, 2n = 26 (Goldblatt 5770). Flowering time. September. Distribution. Middle altitudes in the Kam- iesberg, in central Namaqualand. Figure 76. Geissorhiza kamiesmontana is known from only two gatherings from middle altitudes in the Kamiesberg Mountains of central Namaqua- land, where it is apparently endemic and re- stricted to a few sheltered sites. I made the type gathering on the eastern slopes of the Rooiberg where shade from rocks and tall shrubs gave shel- ter to the plants which grow in shallow soil near seasonal pools and streamlets. e species is clearly related to the other en- demic Namaqualand species of Geissorhiza, G. namaquensis which is similar in general appear- ance. The two have identical corms, puberulent stems and similar leaves with raised and winged margins and pubescence on the margin and mid- rib edges. Geissorhiza kamiesmontana is easily distinguished from G. namaquensis by its well- developed perianth tube, 18-25 mm long which much exceeds the long bracts; its deep blue flower with a relatively short style, the branches of which do not exceed the anthers; and the heavy, almost 428 velvety, white pubescence on the leaves, stem, and inflorescence. The collection made by Lei- poldt “in a kloof near Kamieskroon" has par- ida long bracts (to 21 mm) compared with the type material, in which the bracts are to 18 mm long. The flowers of Geissorhiza nama- quana are pale blue and have a much shorter tube, 6-11 mm long, which is usually about as long as the bracts, while the stem and leaf pu- bescence is less pronounced. Specimens examined. SOUTH AFRICA. CAPE: 3017 (Hondeklipbaai) Kloof near Kamieskroon (BB), Lei- S 3815 (BOL). 018 (Kamiesberg) E slopes of Rooiberg near Wel- Ns (AC), Goldblatt 5770 (K, MO, NBG, PRE, WAG), 6703 (MO). 74. Geissorhiza divaricata Goldbl., sp. nov. TYPE: South Africa. Cape: W of Nieuwoudtville, among rocks in fynbos, Goldblatt 5838 (ho- lotype, MO; isotypes, K, NBG). FIGURE 77. Eras | (15-)20-45 em ala, toris 3, kasa ae NC r= SIS, piens (23-5, sellis ius vel Lang bus, tubo 1-2 mm longo, tepalis ca. 10 mm longis, saan aqa aequalibus, ca. 3 mm longis, antheris 3-4 i tł rami ea i Plants (15—-)20-45 cm high. Corm globose, ta- pering above, and obliquely flattened below, 10- 12 mm diam., tunics blackish, imbricate notched below into segments. Cataphyll firm, brownish, persistent, and accumulating with leaf bases in a fibrous neck around the stem base. Leaves 3, linear to ensiform, 2-3 mm wide, the margins thickened and extending at right angles to the veins ciliate-pubescent, the lower two leaves bas- al, half to about as long as stem, the uppermost leaf inserted in the middle part of the stem, EE. the uid half of the stem for half to ngth. Stem erect, smooth, usually bearing in ind to the sheathing cauline leaf, one or more short, dry, membranous bracts, and branched from these nodes and also sometimes from the cauline leaf axil, sharply flexed above the nodes and at the base of the spike. Spike flexed at the base, inclined, (2-)3-5-flowered, flexuose; bracts dry, membranous and brown at flowering, flexed outwards in the upper half, 5.5— 7 mm long, the inner shorter than the outer. Flowers stellate, white to pale mauve, reddish purple on the reverse of the outer tepals; perianth ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 tube 1-2 mm long, much shorter than the bracts; tepals ca. 10 mm long, ovate-elliptic, ca. 4 mm wide. Filaments equal, ca. 3 mm long; anthers 3-4 mm long. Ovary 1-2 mm long, style dividing near the upper third of the anthers, branches recurved. Capsule globose, 4-5 mm long. Chro- mosome number, unknown. Flowering time. September Distribution. Niguwoudtville Escarpment and Gifberg, in rocky situations in fynbos. Figure 77. (ipicenrhi7n Hi t 1 y very closely allied to G. subrigida, a poorly known species of the Nieuwoudtville area. The two are distinctive in having a sheath of fibers around the stem base, smooth stems, ciliate to pubescent and ribbed have tended to obscure their differences and G. divaricata has until now been identified with G. subrigida. The differences between the two species, however, seem significant. Geissorhiza divaricata has small white to pale mauve flowers with a perianth tube 1-2 mm long and tepals ca. 10 mm, while G. subrigida has large blue to violet flowers with perianth tube 3-4 mm long and tepals 12-17 mm long. Also G. divaricata tends to be taller and more branched and it has smaller corms than the shorter and few branched G. sub- rigida. Both grow along the Nieuwoudtville Es- carpment in the rocky sandstone soils of the area, but have not been recorded growing together. The Nieuwoudtville area is a well known center of endemism for geophytic plants, Iridaceae in rticular, so the occurrence of two distinct, but closely allied species here is not particularly sur- ri sing. Included in Geissorhiza divaricata are two col- lections from the Gifberg, a short distance to the southwest, Barker 9571 and Bayliss 6187, spec- imens of which have either simple or single branched stems and lack the distinctive dry stem bracts found in the Nieuwoudtville populations. The specimens approach Geissorhiza erubescens in outward appearance, but do not have the pu- bescent stems of that species. , Specimens examined. SOUTH rites CAPE: 3118 ), Barker 9571 MN Bayliss 6187 (MO 3119 (Calvinia) near Nieuwoudtville (AC), Leipoldt 821 (BOL, SAM), L. Bolus s n. (BOL 21060); fynbos W of Nieuwoudtville in rock crevices, Goldblatt 5838 (K, MO, NBG); Uitkomst farm, SW of Nieuwoudt- ville, Barker 10745 (K, NBG). 1985] GOLDBLATT — GEISSORHIZA J Y S WY WZ SSS P SSX D SN > : ENS EX SS AS HS @ G. divaricata j $ G. subrigida P33 m d e METERS N AA S E De us iliii FIGURE 77. Morphology and distribution of Geissorhiza divaricata (left) and G. subrigida (right). Habits x 0.5; flowers life size; leaf sections much enlarged (G. divaricata, Goldblatt 5838, Nieuwoudtville Escarpment; G. subrigida, Goldblatt 7220, W of Nieuwoudtville). 430 75. Geissorhiza subrigida L. Bolus, S. African Gard. 22: 275. 1932. TYPE: South Africa. Cape: near Nieuwoudtville, cult. in hort. L. Bolus, Cape Town, Buhr s.n. [holotype, BOL 19995 T as 19943); isotypes, K, PRE]. FIGURE 7 Plants 12-20(-30) cm high. Corm globose, ta- pering above, and obliquely flattened below, 1- 1.5 cm diam., tunics blackish, imbricate, notched below into segments. Cataphyll firm, brownish, persistent and accumulating with leaf bases in a fibrous neck around the stem base. Leaves 3, half to about as long as the stem, 3-6 mm wide, the margins thickened and extending at right angles to the blade, the midrib also raised and winged and at least two other pairs of veins enlarged, ciliate to pubescent on the veins and margin edges, the lower two leaves basal, the uppermost in- serted in the middle part of the stem and sheath- ing 7 at cai two- praia its length, several ribbed. Stem erect, h, bearing in addition to the masia ind ur a short, dry, membranous ract and usually branched from this node and also sometimes from the axil of the cauline leaf, flexed above the nodes and at the base of the spike. Spike flexed at base, (2-)3-5-flowered, flexuose; bracts 6-10 mm long, dry, membra- nous and brown at flowering time, flexed out- ward in the upper half, the inner shorter than the outer. Flowers eee violet blue, pale in the tube; perianth tube 2-3 mm long, shorter than the bracts; tepals 12-17 mm long, ovate-elliptic, to 7 mm wide. Filaments equal, ca. 4 mm long; anthers 4-5 mm long. Ovary 1.3-2 mm long, style dividing at or near the apex of the anthers, becoming declinate, branches recurved. Capsule globose, ca. 4 mm long. Chromosome number, unknown. Flowering time. August to mid-September. Distribution. Sandy soils around Nieuwoudt- ville. Figure 77. Geissorhiza subrigida is an unusual species with comparatively large blue to violet flowers, strongly ciliate-pubescent leaves and a divaricate branching pattern. It is very local in distribution, ern Cape. Closely allied is G. divaricata which is similar in vegetative morphology, but has small- er, white to pale mauve flowers. The similarities and differences between these two species are ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 pee more fully following the description of G. divar The diei uf Geissorhiza subrigida and G. di- varicata, with their raised and flattened margins extending at right angles to the lamina and the raised and winged midrib, both ciliate on their edges are similar to those of G. inflexa and G. heterostyla in particular, although the latter two have a lighter pubescence. They seem well placed in section Ciliata, in which their branched habit, the presence of stem bracts, and their equal sta- mens suggest that they are fairly unspecialized. Specimens examined. SOUTH AFRICA. cAPE: 3119 (Calvinia) Nieuwoudtville (AC), Buhr s.n. (cult.) (BOL 19995, K, NBG 60617, 60618, PRE, STE 17319); Glenridge farm, Nieuwoudtville, Lewis 5886 (NBG); shallow sandy soil in recently burned veld W of Nieu- woudtville, S of the old road to the pass, Goldblatt 7220 (K, MO, PRE). 76. Geissorhiza heterostyla L. Bolus, S. African 148, here designated; isolec- totypes, BM, BOL, K). FIGURE 78. Den rogersii N. E. Br., Kew Bull. 452. 1931. outh Africa. Cape: "Worcester, between Os- ira and Tunnel Sidings, 600-900 m, Rogers 16725 (lectotype, K, here designated; isolecto- ypes, J, K). Hesperantha rosea Klatt, Abh. Naturf. Ges. Halle 15: 395 (Erg. 61). 1882. TYPE: South Africa. Cape: Knysna, Newdegate s.n. (holotype, B *Herb. Lu- beck” Geissorhiza rosea (Klatt) Foster, Contr. Gray Herb. a : 48. 1941, hom. illeg. non Ecklon, Topogr. U nzensamml. Ecklon 20. 1827 [= G. in- xa (de la Roche) Ker]. Hesperantha ciliata E. Mey. ex Klatt, Abh. Naturf. Halle 12: 394 (Erg. 60). 1882. TYPE: South Africa. Cape: Hexrivierskloof, Drège s.n. (holo- type, B “Herb. Lubeck”), nom. superfl. pro Geis- sorhiza rosea Ecklon. Plants 12-45 cm high. Corm symmetric, 4- 12(-15) mm diam., globose, tunics blackish, woody, outer accumulating above, and notched regularly into segments below. Cataphyll mem- branous, often dark above. Leaves 3, linear to ensiform, 2-5 mm wide, half to about as long as the stem, the margins raised and extended at right angles to angles to the blade and ciliate on the edges, the midrib also raised and ciliate, the lower two leaves basal, the uppermost basal or cauline, sheathing for most of its length, several 1985] ribbed on the sheath. Stem erect, puberulous to ciliate or smooth, simple or more often branched, sheathed for up to three-fourths its length, bear- ing a membranous, often dry, bract leaf in the upper part, and usually branching from this axil, or also from the axil of the sheathing leaf. Spike (1-)2-7-flowered, flexed at base and flexuose; bracts 7-10(-15) mm long, initially pale green with hyaline margins, but drying from apex an becoming transparent, or flushed red or brown, the inner shorter than the outer. Flower stellate, either white, flushed pink to purple on reverse of the tepals, or pale blue to mauve, white to yellow or occasionally dark purple in the throat; perianth tube 1-2 mm long; tepals 10-18(-22) mm long, obovate, 6-9 mm wide. Filaments un- equal, the two longer 4-6 mm long, the shorter 3-4 mm long; anthers (2.5—)4—6 mm long, yellow or blue-black. Ovary 1.5-3(-4) mm long, style short to long, either dividing near the base of filaments, with long branches, or reaching to the base or to the apex of the anthers and branches short or long, ultimately recurved. Capsule glo- bose, 5-8 mm long. Chromosome number, 2n — 26 (Hall 193—as Hall s.n., Voelvlei, Suiberland: Goldblatt 135—_as G. leipoldtii, 5815, 6265), 26 + 2B (Goldblatt 5211). Flowering time. August to early October. Distribution. Widespread from the Kubis- kouw Mountains in south Namaqualand through the Roggeveld and Witteberg to the southern Cape as far east as Port Elizabeth; on various soils but usually on sand in the Cape Floristic Region. Figure 78. Geissorhiza heterostyla was described by H M. L. Bolus in 1930 based on a collection of plants from Whitehill in the western Karoo. T outstanding characteristic of the type collection is the variable length of the style. Plants have either long styles, dividing beyond the apex of the anthers; medium styles dividing at anther level; or short styles, dividing below anther level. Other distinguishing features of the collection are a puberulous stem; a submembranous stem bract; leaves with winged, ciliate margins; and unequal stamens, the latter feature not noticed by Bolus, but quite evident in the ample type material. Other collections matching the type have since been made in the same area, e.g., Lewis 4790— Eendracht; Barker 7492— Tweedside; Compton 3772— Cobita —all heterostylous, as well as a few like Compton 15172 and Leighton 201, both from = © GOLDBLATT— GEISSORHIZA 431 the nearby Witteberg that cannot be distin- guished from G. heterostyla except that the styles are invariably long. Phillips 1549, from Seven Weeks Poort, consists ofl ts with either smooth or sparsely papillose stems. The nonheterostylous collections from the Witteberg match closely plants from the Ceres, Koo, and Hex River Valleys, e.g., Esterhuysen 20344— Eselsfontein; Barker 1473— Theronsberg; Loub- ser 2061—Koo Valley; Oliver 5015— Prince Alfreds Hamlet; and Rogers 16725— Osplaats— Tunnel Sidings, the latter the type collection of G. rogersii (Brown, Heterostylous exitii of Geissorhiza het- erostyla have also been made in the Roggeveld, in the Calvinia district, and in the Kubiskouw Mountains in southern Namaqualand. Again, some collections that match the heterostylous material appear to consist solely of long-styled plants (Goldblatt 549— Verlaten Kloof; Gold- blatt 5815—near Middelpos; Goldblatt 5824— Hantamsberg). It is clear that the homostylous populations are the same taxon, and G. hetero- styla must now be regarded as consisting of either heterostylous or homostylous populations. Geissorhiza rogersii accordingly falls into syn- onymy of G. heterostyla. The Witteberg, Ceres, and Hex River Valley populations of G. heterostyla are generally small- er, and less robust than plants from the Rogge- veld and the Calvinia district (though not in- variably), and in turn are easily confused with plants named G. rosea (Klatt) Foster (actually a homonym for G. rosea Ecklon), which can be separated by their having leaves with less strong- ly elevated margins and usually smooth stems. Populations of plants matching G. rosea sensu Foster extend from Worcester south and east along the Langeberg and southern coast to Port Elizabeth, usually occurring on sandy soils. Plants are not always slender or invariably with smooth stems. Some notable exceptions are Goldblatt 2544 — Op de Tradouw, robust and with puber- ulous stems; Goldblatt 4929— Humansdorp, also very robust; Beacom s.n. —near Joubertina, slen- der but with a puberulous stem; and Esterhuysen 24654 — foot of Leeuriviersberg, slender to fairly robust plants with a puberulous stem. Plants that would be considered to be Geis- sorhiza rosea grade into the homostylous form of G. heterostyla to the extent that it seems un- reasonable to maintain them as a distinct species. Nor is there any good character except perhaps ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 «ff —Ç | Up Ç Yh, | ig UT d i | Y JAN. d UAE e Z ¿6 FiGURE 78. Morphology and distribution of Geissorhiza heterostyla showing both long and short styled flowers. Habit and corm x 0.5; flowers life size; leaf section much enlarged (Goldblatt 6265, Nieuwoudtville). the degree of leaf margin elevation that can be consistently used to separate them. Thus G. ro- sea and its synonyms Hesperantha ciliata E. Mey. ex Klatt nom. superfl. and G. ciliaris Salisb. are here regarded as conspecific with G. heterostyla. While there may be some merit in establishing a subspecific taxonomy for G. heterostyla it seems quite arbitrary to which subspecific taxa some of the intermediate forms should be assigned. I have decided therefore, not to formally recognize sub- specific taxa. Those who need to refer to one of the forms of G. heterostyla should use an infor- mal trinomial terminology as suggested by Burtt (1970). Thus slender and smooth (rarely puber- ulous) stemmed and long-styled plants with leaf margins raised relatively little compared to leaf width can be referred to as G. heterostyla (rosea). Populations of heterostylous plants, typically with puberulous stems (rarely smooth) and leaves with margins raised well above the leaf surface can 1985] similarly be designated G. heterostyla (hetero- "a Pes: homostylous m can be des ted = heterostyla (rogersi Two more forms of Pads uit heterostyla need to be mend The first consists of dis- tinctive plants with medium to large white flow- ers (Thomas s.n. — Eilandia Road; Olivier 283— Karoo Garden) which initially appears different from other populations. Little more than a color difference however seems to be involved, as sim- ilarly large-flowered plants occur elsewhere. The second population, from Stettynskloof, south of Worcester (Barker 9455), incidentally the most westerly station, seems fairly typical at first hav- ing very unequal stamens and pale flowers. All but one of the several specimens on the sheet lack the membranous stem bract, and except for the unequal stamens these plants are very like the related G. inflexa which occurs in this area and to the south. It seems that this population, on the edge of the range of G. heterostyla, is intermediate in some respects with G. inflexa, possibly as a result of introgressive hybridiza- tion. Some collections from the Sutherland district, notably Hanekom 1562—Houthoek and Acocks 15963 — Klipbanksrivier, are remarkably robust and the large flowers are purple with darker pig- mentation in the center. These specimens appear to represent a distinct local race or form of G. heterostyla. Specimens examined. SOUTH AFRICA. CAPE: 3019 (Lo D. Kubiskouw, near summit (CD), Mar- x 12870 (PR 119 (Calvinia near Nieuwoudtville on the Loer- di road (AC), Lewis 5850 (NBG); 1 mi. N of Nieuwoudtville, Barker 9563 (NBG, S, STE), Goldblatt (BC), Goldblatt 5824 (K, MO, NBG); upper slopes be- low cliffs at the W end of the Hantamsberg, Goldblatt 5805 (K, MO, NBG, PRE); wet red clay, top of Han- tamsberg (BD), Goldblatt 426 (BOL); Driefontein voor Hantam (DA), Marloth 12811 (PRE) 3120 (Williston) Blomfontein farm, E of Middelpos (CC), Barker 10774 (K, NBG); 66 km S of Calvinia, on road to Blomfontein, Goldblatt 5815 (MO, PRE); 3220 (Sutherland) D a ine Men bs 6 (K e, Golda 6322 BI berg > £ o foot of the Roggeveld Mts. (CB), Acocks PRE); 13.5 mi. SSW ofthe foot of Komsberg Pass (DC), GOLDBLATT — GEISSORHIZA 433 od 18447 (BOL, K, PRE); 20 mi. aed of the foot of Komsberg Pass, Leistner 269 (PR 3319 (Worcester) Gydo Pass, we slopes (AD), as rom & Acocks 284 (K, PRE); flats NW of Prince p og 5015 (PRE); Eselsfontein, Ceres, ( B pares 15941 (BOL); Onse Rug farm ker 9443 (NBG); lower slopes of Apes pem Onse Rug farm, posea 4089 (E, MO, PRE, S, US, WAG); Worcester, Nature Reserve, Olivier 283 (K); Stettyns- kloof (CD). Pih 9455 (NBG); Eilandia S mi. from Worcester-Robertson road (DA), Thomas s.n. (NBG 93061, PRE 58032); NW — to pi valley (DB), Loubser 2061 (NBG); Langeberg, above Berg- en-Dal, cliff face, 2, I ke a fsterhsen 35684 (BOL, MO); Berg-en-Dal, a s, slopes near the base of the pci L 5 35 797 (MO, PRE G). 3320 (Montagu) Voetpadsberg, Pages Ae Barker 7492 (NBG); Tweedside, Lewis s.n. (Na Gard. 2694/32 in BOL, K, SAM); Wiltabert ae burg (BA), Leighton 201 (BOL); Witteberg at Bantams 5,000 ft., Compton 12244 (NBG); Witteberg, White- hill, Compton 13997 (NBG), 15172 (NBG), 16277 (NBG); Whitehill, L. Bolus s.n. (BM, BOL 19148); Donkerkloof, Montagu (CC), Lewis 4789 (SAM); slopes at foot of the Langeberg near Leeuriviersberg (CD), Esterhuysen 24564 (BOL, PRE); Op-de-Tradouw, W of Barrydale (CD), Goldblatt 2844 (MO, PRE). 3321 (Ladismith) Elandsberg (Torenberg), Klein dismith, Loubser 2086 (N serve, Calitzdorp (BC), Vlo " Oaks, Gysmanshoek Pass, Riversdale (CC), 722 (MO, STE); damp slope, Garcias Pass, Lewis 5646 BG). 3322 (Oudtshoorn) Swartberg, hills above the Cango Caves (AD), Vlok 292 (MO, NBG, SAAS), Kamma- nassie Mts., 2,800 ft. (DB), Viviers & Vlok s.n. (MO, SAAS). 3323 (Willowmore) Bo Koega, near Joubertina, Bay- liss 5954 (MO); hills N of Kruisfontein (CC), Fourcade 4430 (STE); Rede beside National Road (DD), Beacom s.n. (NBG 87524, 92829). 3324 This cia 4 p N of Humansdorp, Hankey road (DD), Fourcade 616 3420 (Bredasdorp) bani (AB), Fry sub Galpin s.n. A Limestone hills, De Hoop (AD), Lewis 6019 (M, NBG); Ponebers. ledges on S. slopes (BC), Ester- huysen 23210 (BOL, K), Hamerkop, S side of Potte- berg, Acocks 22745 PE valley behind Oyster Beds Hotel, Fort Beaufort (BD), Marsh 824 (K, STE); Cape Infanta, Blum 18 (Ey; Duiwenhokrivier Kloof, between Port Beaufort and Riversdale, Lewis 6013 (MO, NBG, STE ). 3421 (Riversdale) Still Bay, ridge below the reservoir (AD), Bohnen 4251 (STE); Roodehoogte, near Her- bertsdale (BB), Muir 1271 (BOL); E foothills N of Herbertsdale, Goldblatt 4156 (MO, P 3423 (Knysna) Victoria Bay, George (AA), ` Barker 8181 (NBG, STE); slopes above Victoria Bay, Lewis 3631 (SAM); near George, Burchell 6088 (K); Noetzie 434 RE 79. Morphology and distribution of Geiss- orhiza arenicola. Habit x0.5; intact flower life size; opened flower x 1.5; leaf section much enlarged (Go/d- blatt 7056, heathland W of Nieuwoudtville). (BA), Middlemost s.n. (NBG 60637, SAM 61725); na- tional road outside Plettenberg Bay (AB), Ene 2912 (MO, WAG); Keurbooms River, Galpin 4 (K, PRE); Plettenberg Bay, Fourcade 4806 (BOL, Mo ere Salter 6941 (BOL). 424 (Humansdorp) Oudebos flats (AA), Fourcade s (BOL), 3364 (BOL, K, PRE, STE); Krom River, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 W of Humansdorp (BA), Acocks 21476 (PRE); duine veld, E of Kromme River mouth (BB), Fourcade 2781 (STE); 4 km E of Humansdorp, Goldblatt 4929 (MO), 5211 (MO, PRE); 8.5 mi mansdorp, , Barker 9068 (NBG), d yr (RE, os 467 1 (K, PRE). 3 n akens River (DO), Long 47 : E) 762 (BOL, PRE), Fries, Norlindh & Weimarck 217 (K, e € ns River, Long 441 (K, PRE). Í » 30) 55 (Bx Cape ofc Good Hope (C.B.S.), Sparrman n. (S), Bowie 18 (BM). 77. Geissorhiza arenicola Goldbl., sp. nov. TYPE: South Africa. Cape: sandy wet heathland west of Nieuwoudtville, Goldblatt 7056 (holo- type, MO; isotypes, K, NBG, PRE, S). FiGURE 79. 12-26 cm alta, tunicis cormi imbricatis duris, duobu pe- , tubo m longo, tepalis 10-15 mm longis, fil lamentis inaequali- bus, antheris 3-5 mm longis, WAR diviso prope apicem antherarum. Plants 12-26 cm high. Corm globose, 6-10 mm diam., symmetric, tunics blackish, imbri- cate, woody, notched into regular segments be- low. Cataphyll solitary, membranous. Leaves 3, narrow, linear, 1-2 mm wide, reaching the mid- line of the stem or the base of the spike, the margins and midrib enlarged and thus 2-grooved on TR SHE. Ane eat oF the margins and ently smooth, often sticky below and with aog sand, the lower two leaves basal, the uppermost inserted shortly above ground, often largest, sheathing the stem for two-thirds of its length, the sheath several ribbed and somewhat inflated. Stem erect, pu- berulent above the third leaf, smooth below, oc- casionally with 1 branch, without a sheathing bract. Spike flexuose, inclined, (1—)3—8-flowered; bracts herbaceous, the margins often lined with red above, sometimes becoming dry and mem- branous from the apex, 7-9 mm long, the inner shorter and much narrower than the outer. Flow- er stellate, white or deep blue-mauve and then pale in the throat; perianth tube 1.5-2 mm long, enclosed in the bracts, infundibuliform; tepals 0-15 mm long, 4-8 mm wide, oval. Filaments slightly unequal, 3-4 mm long, one 1-1. shorter than the other two; anthers 3-5 mm long, pale yellow. Ovary 1.5-2 mm long, style dividing at the apex of the anthers, branches ca. 2 mm Ó 1985] long, recurved. Capsule unknown. Chromosome number, 2n = 26 (Goldblatt 7056). Flowering time. September to mid-October. Distribution. Nieuwoudtville Escarpment, between Grasberg and Lokenberg, and on the Gifberg, in stony Cape mountain soil. Figure 79. Geissorhiza arenicola is endemic to the Gif- berg and Nieuwoudtville Escarpments where it grows in the rocky, poor sandstone-derived soil typical of the mountains of the Cape System. It is nowhere common, but has been collected at four places along the Nieuwoudtville Escarp- ment as well as on the Gifberg a short distance to the southwest. Geissorhiza arenicola 1s dis- tinctive in its puberulent stem; heavily ribbed and grooved upper leaf sheath; sticky basal leaves with strongly raised margins; and slightly to dis- tinctly unequal stamens. While blue is the usual flower color, the collection from Lokenberg, Acocks 17251, is described as having white flow- ers. It is apparently related to the better known and striking, brilliant blue-flowered Geissorhiza splendidissima with which it shares several im- portant features, including a puberulent stem; a strongly ribbed and grooved upper leaf sheath; short perianth tube; and basal leaves with strong- ly raised margins and midrib, the edges of which are usually lightly ciliate. These two species have been assigned to section Ciliata, but as discussed under G. splendidissima, they are exceptional in this alliance in their strongly ribbed upper leaf sheath, a characteristic of section Geissorhiza. Geissorhiza arenicola shows some resemblance to the Nieuwoudtville endemic, G. sulphuras- cens, a species which has creamy yellow flowers and a smooth stem. The similarity is probably coincidental, and the two species can always be distinguished by several different characters. Specimens examined. SOUTH AFRICA. CAPE: 3118 (Vanrhynsdorp) Gifberg (DC), Phillips 7518 (NBG, 3119 (Calvinia) Nieuwoudtville Escarpment in fyn- bos of town, 3 km S of the Calvinia road (AC), Goldblatt 5837 (MO); 5 mi. NW of Nieuwoudtville on Grasberg road, Barker 9560 (M, NBG); wet sandy heathland W of Nieuwoudtville, Goldblatt 7056 (K, MO, NBG, PRE, S); wet sandy meadows, Bokkeveld, Marloth 7641 (PRE); Lokenberg, arid fynbos (CA), Acocks 17251 (PRE). 1909; Foster, Contr. Gray 78. Geissorhiza splendidissima Diels, Bot. Jahrb. Syst. : ; Herb. 135: 31. 1941. TYPE: South Africa. GOLDBLATT — GEISSORHIZA YW, 1200 iili ML " 7 FiGurE 80. Morphology and distribution of Geiss- orhiza splendidissima. Habit x 0.5; flower life size; leaf section much enlarged (Goldblatt s.n., Glenlyon farm, Nieuwoudtville). Cape: Oorlogskloof, Diels 627 (holotype, B). FIGURES 12, 80. Plants 8-20 cm high. Corm ovoid, truncate at base, 7-10 mm diam., tunics dark, imbricate, the outer layers incised below into regular trun- cate segments. Cataphyll not evident. Leaves 3, linear, shorter to slightly longer than the stem, (1-)1.5—2 mm wide, the margins raised and ex- tending at right angles to the blade, the midrib thickened, the blade thus narrowly 2-grooved on each surface, the edges of the margins and the midrib minutely ciliate, slightly sticky and with sand adhering in places, the lower two leaves basal, the lowermost longest, the uppermost in- with a short bract leaf above, simple or with a single, or rarely 2 branches. Spike (1—)2-5(—6)- 436 flowered, flexed at the base of the spike; bracts green, becoming dry and brown from the apex, the outer 16-20(-25) mm long, the inner 8-16 mm long. Flower deep blue-violet, darker to- wards the base of the tepals, and yellowish in the throat, tepals cupped, zygomorphic, the stamens unilateral and declinate, held above the dorsal tepal; perianth tube 2-4 mm long, widening from the base; tepals broadly ovate, 15-22 mm long, outer 9-14 mm wide, inner 8-12 mm. Filaments ca. 5 mm long, equal, unilateral and declinate; anthers unilateral, horizontal, lying over the dor- sal tepal, 7-8 mm long, pollen red-brown. Ovary ca. 3 mm long, style 10-11 mm long, lying below the stamens, dividing at the apex of the anthers, branches 2 mm long, recurved. Capsule globose, 6-7 mm long, to 5 mm diam., enclosed in the bracts. Chromosome number, 2n = 26 (Gola- blatt 347) Flowering time. Late August to September. Distribution. Sandy to stony flats to the north and south of Nieuwoudtville, east of the escarp- ment above Van Rhyns Pass. Figure 80. Geissorhiza splendidissima is endemic to the area immediately around Nieuwoudtville in the northwestern Cape, where it grows in light clay soil among low-growing shrubs. It flowers well then it may make a striking display. The large, glossy deep blue-vi- olet flowers are amongst the most beautiful in the genus and the species is well worth horticul- tural attention. The affinities of Geissorhiza splendidissima seem to be with section Ciliata, the species of which h y, with strong- ly raised margins and winged margins that are ciliate on the edges. Its puberulent stem is also consistent with this disposition for several species of the section also have this feature. The zygo- morphic flowers of G. splendidissima are unique in the section and must be regarded as having evolved independently in this species and not indicative of a relationship to other BECHER in which th leaf with similar flowers. It is somewhat unusual in section Ciliata also in the somewhat ribbed sheath of the up- permost leaf. This character is shared with the apparently allied G. arenicola and this seems to set these two species somewhat apart in the sec- tion. cimens examined. SOUTH AFRICA. CAPE: 311 on Nieuwoudtville- isis road (AC), a ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 blatt 347 (BOL); ca. 1 mi. N of Nieuwoudtville, L. Bolus s.n. (BM, BOL 19439, K); Nieuwoudtville, Salt- er 1645 (BM), Buhr s.n. (Nat. Bot. Gard. 636/29 in BOL, Nat. Bot. Gard. 300/37, a i in NBG, 822/ i Martley, B Grasberg, 15 mi. o (NBG, UO); Charlies Hoek, Barker 10759 (NB apkuilsfontein, Leipoldt 3035 alvinia, "say 817 (SAM); Bokkeveld, Oorlogskloof, Diels 627 (B); Glen- lyon farm, Nieuwoudtville, Hardy 67 (G, K, PRE, Z). 79. Geissorhiza inflexa (de la Roche) Ker, Ann. Bot. (Kónig & Sims) 1: 224. 1804; Baker, Handb. Irid. 157. 1892, Fl. Cap. 6: 73. 1896, non sensu Baker (intended for G. imbricata sensu lato); Goldblatt & Barnard, J. S. Af- rican Bot. 36: 296-298. 1970. Ixia inflexa de la Roche. Descr. Pl. Aliq. Nov. 15. 1766. Herb. 135: 77. 1941, excl. ba- sion. name misapplied to ieee va- ginata (Sweet) Goldbl.]. TYPE: South Africa. Cape: Cape Peninsula, Barker 3869 [neo- type, BOL, designated by Goldblatt & Bar- nard (1970); isoneotype, NBG]. FIGURE 81. Ixia ye V eue. Diss. de Ixia no. 6. 1783, hom. illeg. o I. infl d sup isb., od. 36. sorhiza erosa Salisb.) Foster, Contr. Gray Herb. 135: 52. 1941. : South Africa. Origin un n: cultivated in England [location of type unknown, possible at G (Herb. Ixia clar: d ES eid Bot. Mag. 18: sub. tab. 672. "n ci us are D tly ses on a specimen collected Es Auge s.n. (BM), annotated “Ixia cil- iaris Salisb. Prod." ydo. in L Salisbury ir It is the large, red- or purple-flowered form o inflexa.) Gei. iliaris Salisb., Trans. Hort. Soc. London 1: 321. 1812, nom. nud. (apparently intended as a combination in Geissorhiza of “Ixia ciliaris Salisb., Prod. 36. > a name never published — possibly Salisbury changed the epi- - in print: Ixia erosa Salisb., which is conspe- appears on the page cited in his Prodromus). Trim ciliaris (Ker) n Index Kew. 1: 1006. 1895, nom. nud. basion. Geissorhiza rosea Ecklon, Topogr. iccorhi za) ry Verz. Pflanzen- s.n. [lectotype, S, designated by Nordenstam 1970 ( ee candida Ecklon, Topogr. Verz. Pflanzen- - Mars n 21. 1827. TYPE: South Africa. Ca Hot tots Holland, Zeyher s.n. (lectotype, S). RUE quinquangularis Ecklon ex Klatt, Linnaea 1985] GOLDBLATT — GEISSORHIZA 437 E > = > s RU YA, 1200 T Mili K 439 g 18 Yu æ 21 r Y ! L N) Ficure 81. Morphology and distribution of Geissorhiza inflexa. Habit life size; opened fl d gy x 2: leaf section much enlarged (Goldblatt 5643, N of Bot River). 34: 654. 1865; Foster, Contr. Gray Herb. 135: sheet designated by Foster (1941: 54) as the lec- 53-54. 1941, nom. superfl. pro G. rosea Ecklon totype; Ecklon & Zeyher Irid. 215; Ecklon & Zey- if this species is accepted as validly described, as her Irid. 296; and Lalande s.n.] [Speci ited in the d ipti Eck- | Hesperantha kermesina Klatt, Abh. Naturf. Ges. Halle 50 ere lon & Zeyher Irid. 214 (Caledon Zwarteberg), the 15: 395 (Erg. 61). 1882. Geissorhiza erosa var. 438 kermesina (Klatt) Foster, Contr. Gray Herb. 135: 53. 1941. TYPE: South Africa. Cape: between Paarl and Pont, Drége 8480 [lectotype, B, effectively designated by Foster (1941: 53); isolectotypes, B, M K, L, MO, S] Hesperantha quinquangularis Ecklon, Topogr. Verz. Pflanzensamml. Ecklon 23. 1827, nom. nud. diri eos graminifolia Baker, Handb. rid. 155. 1892, Fl. C : 6. TYPE: South Africa. Cape: near Swellendam, Drége 3496 (lectotype, K, here designated; isolectotypes, B, E, G, L, MO, PA near R Burchell 6088 (syntype, K = G. hetero- sty Tei d var. bicolor Baker, Handb. Irid. 155. 1892, Fl. Cap. 6: 70. 1896. TYPE: South Africa. Cape: Hout Bay, MacOwan in Herb. Norm. Austr. Afr. 261 (lectotype, K, here designated; iso- lectotypes, B, BOL, G, SAM; syntypes, see Flora Cap. 6: 70. 1896). Geissorhiza quinquangularis var. atrofaux Foster, r. Gray Herb. 135: 55. 1941. Type: South de Cape: Piketberg, Schlechter 5210 (holo- pe, B; isotypes, B, BM, G, K, SAM, W, Z a pilosa Pappe ms. [Cape Flats, Pappe s.n. (SAM)]. Plants T 30(-40) cm high. Corm sym- metric, 7-12 mm diam., tunics blackish, imbri- cate, the outer : e notched regularly below into segments. Cataphyll membranous, dark above. Leaves 3, usually half to two-thirds as long as the stem, sometimes exceeding the spike linear-ensiform, (1.5-)3-4(-10) mm wide, the margins extended outward at right angles to the blade, the midrib also raised and winged, the edges of the margins and midrib and sometimes other larger veins ciliate-pubescent, the lower two leaves basal, the uppermost basal or more often cauline, sheathing for most of its length, the sheath ribbed and ciliate-pubescent, slightly inflated. Stem erect, smooth, sheathed below by the up- permost leaf, rarely branched. Spike (1-)2- 6-flowered, flexed at the base; bracts membra- nous, dry, reddish brown, occasionally greenish below, 7-15(-22) mm long, the inner somewhat smaller than the outer. Flowers stellate, either purple, pink, red, or white, cream or pale yellow and then usually red-flushed on the reverse of the outer tepals and occasionally dark in the cen- ter; perianth tube short 1-2.5 mm long; tepals (8-)10-18(-24) mm long, obovate, 8-10 mm a widest. Filaments 4—7 mm long, equal; anthers (2.5—)3—6(-8) mm long. Ovary 2-3 mm long, style dividing near the apex of the anthers, branches ca. 4-5 mm long, recurved above. Capsule glo- bose to oblong, 3-6 mm long. Chromosome number, 2n = 26 (Goldblatt 2497, 6173), 2n = 26 + 1B (Goldblatt 5228). _ ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Flowering time. August to September Distribution. Southwestern Cape, from Bre- dasdorp and Swellendam in the east to the Cape Peninsula and north to Piketberg, extending in- land in the Tulbagh and Worcester valleys. Fig- ure 81. Geissorhiza inflexa was described in 1766 by Daniel de la Roche in an important publication, actually his doctoral dissertation, in which 20 species of Cape Iridaceae were described. Placed in the genus Ixia, it was misinterpreted by Baker (1892, 1896) who cited specimens of G. imbri- cata under this name and by Foster (1948) who confused it with the large yellow-flowered Hes- perantha vaginata from the Calvinia district in the northern Cape. Authentic material at the Lei- den Herbarium (Goldblatt & Barnard, 1970) makes it quite clear that 7. inflexa is a species of Geissorhiza, as realized by Ker in 1804. It is conspecific with what was long known as G. quinquangularis, or G. graminifolia. A neotype, Barker 3869 from the Cape Peninsula, a goo match for the material at Leiden, was designated by Goldblatt and Barnard. The species is common in the southwestern Cape on clay and shale slopes and sometimes is found on very stony ground. It extends from Bre- dasdorp and Swellendam in the east to the Cape Peninsula and Mamre and northward to the Pi- ketberg district with inland extensions into the Tulbagh and Worcester valleys. It belongs to sec- tion Ciliata of subgenus Geissorhiza, the species of which have leaves with a raised margin which extends at right angles to the blade and itself has ciliate edges. The midrib is also slightly raised and winged with ciliate edges. Occasionally one or more secondary pairs of veins may also be ciliate. Of the group, G. inflexa seems one of the more specialized, having few flowered spikes and only occasionally branched stems. It always has three leaves and lacks the stem bract found in some of the other species of the group. The flow- ers are moderate to large in size, and range in color from cream to pale yellow with pink or red on the reverse of the outer tepals to shades of pink, red, or purple. Geissorhiza heterostyla, a southern Cape species that extends north into the western Karoo is closely allied, but can readi- ly be distinguished by its one filament much shorter than the others, a more branched inflo- rescence, and conspicuous dry stem bracts below the spike. There is a tendency for reduction in flower size 1985] and narrowing of the leaves in populations of Geissorhiza inflexa towards the east of its range, south and east of Caledon. At Bredasdorp spec- imens seem very different from the robust plants that occur at Tulbagh, and it is easy to see why earlier authors should have recognized these as separate species. However, the narrowest leaved form which is found at Bredasdorp, with leaves 1-2 mm wide and flowers with tepals ca. 9 mm long (Wasserfall 376, Barker 2527, 2475, Comp- ton 9094) is connected through a series of inter- mediates to the type specimens of G. quinquan- gularis (leaves 3 mm wide, tepals 15 mm long) and G. inflexa (leaves 4-6 mm wide, tepals to 12 mm long). The latter specimen in turn match- es, except in flower color, several collections from Stellenbosch, Mamre, and Tulbagh, with purple or red flowers [e.g., Leighton 1336, Zeyher s.n. (SAM 20861), Mathews 10, Grant 2459 (leaves 5-8 mm wide, tepals to 14 mm long)] which are clearly conspecific with such strikingly robust specimens as Lewis 5335, 5740, MacOwan s.n. (Herb. Norm. Austr. Afr. 590) (leaves 5-10 mm wide, tepals 18-20 mm long) Traditionally the red, pink, or purple and ro- bust form of Geissorhiza inflexa has been re- garded as a species distinct from plants with white to cream flowers marked red on the reverse of the outer tepals. Baker (1892, 1896) recognized pink- to purple-flowered plants as G. hirta (an ue name) and the paler flowered and often slender plants as G. graminifolia. The latter is actually based on two collections, one, Burchell 6088, is G. heterostyla and the other, a Drége collection, here designated the lectotype, match- es several other specimens of G. inflexa cited in *Flora Capensis." The Drége collection corre- sponds closely to the type material of G. quin- quangularis Ecklon ex Klatt, a much earlier name, and the one used by Foster in his monograph for this form. Foster used the name G. erosa for the robust, and red- to purple-flowered plants as he realized G. hirta was superfluous and illegiti- mate The recognition of the two forms as separate species seems unjustified, as it is based largely on color differences. Specimens with cream flow- ers from the Cape Peninsula, and north of Cape Town, as well as the type collection of G. quin- quangularis match some of the less robust, red- flowered specimens of G. erosa-hirta in every respect except flower color, and separation of species on color differences alone is unaccepta- ble, and especially since cream-flowered speci- GOLDBLATT — GEISSORHIZA 439 mens occur occasionally in populations of the red-flowered form from Tulbagh (Lewis 5738, Garside 1541). These two forms are consequent- ly united here in one species, the earliest name for which is G. inflexa. There is obviously in- tense selection for red flower color in the Tulbagh valley, and several species of Iridaceae, else- where white- or blue-flowered, have flowers in red to purple colors here (e.g., Sparaxis gran- diflora, Babiana villosa, and Galaxia versicolor). Similar selection for a cream (to pale yellow) flower with a dark center and throat is evident along the coastal belt north of Cape Town es- pecially in the Piketberg-Porterville area and populations of Geissorhiza inflexa here also have pale flowers with a dark center. This form was recognized by Foster (1941) as G. inflexa var. atrofaux. It seems to me no more than a minor color variant, and not worth taxonomic recog- nition. Loubser has recorded probable hybrid- ization between this form of G. inflexa and G. tulbaghensis where they occur together on a farm northwest of Porterville, also in association with G. secunda. Loubser 2188 and 2104 have large flowers with a red throat, a character of G. tul- baghensis, while Loubser 2102 has self-colored flowers Specimens examined. SOUTH AFRICA. CAPE: 3218 (Clanwilliam) Piketberg Mt. ~ Schlechter 5210 P BM, G, K, SAM, W, Z); Nurust farm 8 mi. NW Porterville (DD), Loubser 2102 (BO, 2188 (NBG). 3318 (Cape Town) Contreberg slopes (AD), Pillans i 3 6945 (BOL); Mamre hills, Barker 1813 (NBG); Groenekloof, Zeyher s.n php 1), MacOwan in ib Norm. Austr. cee 590 (B, BOL, G, GH, K, SAM); ar Cape Town (CD), H. porum 4805 (BOL, K); Signal Hill Barker 405 (NBG), Wolley Dod 521 (BM, BOL, K); Signal Hill, above Tamboers Kloof, Michell s.n. Dep ) Saler 7629 v SAM); Lions Head, Ecklon 312 (B, 2840 (E); Stellenbosch (DD), Ecklon & Zeyher Irid. 215 (FI, G, MO); near Stellenbosch, Lightfoot s.n. (BOL 3194); between Paarl and Pont, Drége 8480 (B, MO, S); Onderpapagaisberg, lower slopes, Taylor 6882 (STE); Jonkershoek, Garside 1531 (K); Jonkershoek valley, Lewis 1995 (SAM); Stellenbosch, Duthie 585 (BOL), Worsdell s.n. (K); Paarl, Loubser 3319 (Worcester) Tulbagh (AC), Hall 1426 (NBG), Grant 2459 (MO), Marloth 7801 (PRE), $909 (PRE), Garside 1541 (K), Rogers 11097 (Z), MacOwan 2678 (E, K); 2 mi. S of Tulbagh, stony clay vis Lewis 5740 (NBG); S-facing hill slopes on clay at Tulbagh, Lewis 5738 (NBG, STE); i Leighton 1338 (BOL, NBG); foot near Michells Pass, Goldblatt 4077 (MO) Tulbagh cemetery, Goldblatt 5228 (MO); 1 mi. S of Tulbagh 440 Road Station, Goldblatt 304 (MO); near Artois, H. Bolus s.n. (BOL); Ceres, Reyneke 12 (BOL); Ceres Road (Wolseley), Schlechter 8975 (B); 3 mi. from Hermon on Tulbagh road, Marsh 738 (PRE, STE); Slanghoek valley (CA), Barker 9470 (NBG); shale hills between Villiersdorp and Brandvlei (CD), Goldblatt 6203 (MO). 3418 (Simonstown) near Hout Bay neck (AB), MacOwan in Herb. Norm. Aust. Afr. 261 (B, BM, BOL, G, K, SAM, W) = MacOwan 2632 (SAM); Cape Flats (BA), Pappe s.n. (K, SAM); Hottentots Holland (BB), Ecklon & Zeyher Irid. 214 (91.8) (K, SAM); wet flats at Gordons Bay, H. Bolus 9939 (BOL, K, PRE, Z); near Faure at Macassar Beach turnoff, Jackson s.n. (NBG 88444); near Oudebos farm, Palmiet River Val- ley, shale band (BD), Boucher 1894 (PRE, STE). 3419 (Caledon) N of Bot River (AA), Goldblatt 5643 (BR, MO, US, WAG), Goldblatt 49824 (MO); Bot Riv- er-Caledon road via Goedvertrou, Goldblatt 3996 (MO); clay slope W of Eseljag Pass on road to Queen Anne, Goldblatt 2497 (K, MO, PRE, S, US, WAG); stony clay flats S of Villiersdorp at Theewater farms turnoff (AB), Goldblatt 4015 (MO, NBG); between Caledon and Villiersdorp i d d bridge, Salter 4793 (BM, BOL, K); Caledon (AB), Barker 2495 (NBG); commonage below Zwartberg, Caledon, Guthrie s.n. (BOL); lower S slopes of Swartberg, Caledon, Ester- huysen 32670 (BOL, MO), Ecklon & Zeyher Irid. 214 (51.8) (B, BR, E, FI, G, LD, W, Z), Schlechter 5561 B, PRE, Z); Caledon Zwartberg lop t Gardens, Goldblatt 5896 (MO, WAG); clay slope on W outskirts of Caledon, Goldblatt 6173 (MO); 18.4 km SW of Greyton, Acocks 24340 (K, MO, PRE); Albertyn-Ca- «Di Glen, near Fairfield, Goldblatt 4848 (MO); The Poort, SW of Bredasdorp (DB), Wasserfall 378 (NBG), Gold- blatt 4862A (MO), Barker 2475 (NBG), 2527 (NBG), Acocks 24238 (EA, STE); Poort hills, Bredasdorp, Compton 9094 (NBG). WITHOUT OCALITY: Breede River, near Swel- lendam, Drége 3496 (B, BM, E, G, K, L, MO); Cape of Good Hope (C.B.S.), Herb. Burman s.n. (G), Thun- berg s.n. (Ixia hirta) (S "Herb. Montin," UPS), Rox- burgh s.n. (BM), Pappe s.n. (BM), Prior s.n. (K). 80. Geissorhiza erubescens Goldbl., sp. nov. TYPE: South Africa. Cape: Pakhuis Pass, on shale slopes near top of pass, Goldblatt 6396 (holotype, MO; isotypes, K, NBG, PRE, S, WAG). FIGURE 82. Planta (5—)8—15 cm alta, foliis 3, duobus inferioribus basalibus erectis linearibus, floribus (1—)3-7, albis, ru- i S rum exteriorum, tubo 1.5-2 mm longo, tepalis 8-12 mm longis, filamentis aequalibus, antheris 2.5-3 mm longis, stylo diviso prope apicem antherarum, ramis recurvatis. Plants (S—)8—15 cm high. Corm more or less globose, obliquely flattened below, 5-12 mm diam., tunics blackish, imbricate, notched reg- ularly below into segments. Cataphyll firm, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 v | FIGURE 82. Morphology and distribution of Geis- sorhiza erubescens. Habit 0.5; flower life size; leaf section much enlarged (Goldblatt 6396, Pakhuis Pass). brownish, persistent, and accumulating with leaf bases in a fibrous neck around the stem base in older individuals. Leaves 3, linear and erect, 2- 3mm wide, halfto two-thirds as long as the stem, the margins raised with wings extending at right angles to the blade, the midrib also raised and winged, minutely ciliate to visibly pubescent on the edges of the margins, midrib, and sometimes other veins, the lower two leaves basal, the up- permost leaf largest and inserted in the middle part of the stem, sheathing below. Stem erect, ciliate to pubescent below the third leaf, ciliate or smooth above, simple or 1-branched. Spike flexed at the base, (1—)3-7-flowered, flexuose; bracts membranous and greenish to transparent below, rust brown above, 6-8 mm long, flexed outward in the midline, the inner shorter than the outer. F/owers stellate, creamy white, bright red on the reverse of the outer tepals; perianth 1985] tube 1.5-2 mm long, much shorter than the bracts; tepals 8-12 mm long, ovate-elliptic, 3-4 mm wide. Filaments equal, ca. 3 mm long; anthers 2.5-3 mm long, cream. Ovary 1-2 mm long, style dividing at the apex of the anthers, branches re- curved. Capsule unknown. Chromosome num- ber, 2n = 26 (Goldblatt 6396). Flowering time. September. Distribution. Known only from Pakhuis Pass, common on the shale band, especially after fires. Figure 82. Geissorhiza erubescens is a distinctive small- flowered species of section Ciliata, probably most closely allied to the southwestern Cape G. inflexa and to G. leipoldtii which occurs in the same general area as G. erubescens. It can be distin- guished by the unusual sheath of fibers around the stem base, at least in older individuals; rel- atively small flowers with tepals 8-12 mm long; strongly winged leaf margins; and its lightly cil- iate-pubescent stem, at least below the upper- most leaf. It is perhaps closest to G. /eipoldtii which sometimes has unequal stamens and typ- ically larger flowers, the tepals (13-18-28 mm long, without the deep red markings on the re- verse of the outer tepals characteristic of G. eru- bescens. It also bears some resemblance to the Nieuwoudtville and Gifberg species, G. divari- cata, but this typically has distinctive, dry stem bracts and always has a smooth stem. Geissorhiza erubescens has been collected only in the upper part of Pakhuis Pass in the shale band, growing in a light sandy clay gravel. When I found it in 1981 it was flowering in profusion in the season following a veld fire. It seems likely that it flowers only in the spring after a burn. Specimens examined. SOUTH AFRICA. CAPE: 3219 (Wuppertal) E slopes of Pakhuis Pass (AA), Compton 19964 (NBG); Pakhuis Pass, near top of pass on shale slopes, Goldblatt 6396 (K, MO, NBG, PRE, S, US, WAG), Goldblatt 6397 (MO). 81. Geissorhiza leipoldtii Foster, Contr. Gray Her 47. 1941. TYPE: South Africa. Cape: Pakhuis-Wuppertal road, Leipoldt s.n. (holotype, BOL 20778 in K; isotype, BOL). FIGURE 83. Geissorhiza ixioides Schltr. ex Foster, Contr. Gray Herb. 135: 43-44. 1941. Type: South Africa. Cape: “Agtertuin, in collibus," Schlechter 10866 (ho- lotype, B; isotypes, BM, BOL, BR, E, G, K, L, LD, MO, P, PH, PRE, S, Z). GOLDBLATT — GEISSORHIZA 441 € NI N > as D 2 ASS ASS y NS: AN TA G ANS (Q SS METERS ° E. p Ze Mill " 19 20 1 AB 83. Morphology and distribution of Geis- sorhiza leipoldtii. Habit x 0.5; flower life size; leaf sec- tion much enlarged (Goldblatt 6165, S of Clanwilliam). Plants (12-)16-30 cm high. Corm symmetric, globose, 5-8 mm diam., tunics dark brown to blackish, imbricate, the outer layers regularly notched below into segments. Cataphyll solitary, membranous, brownish above. Leaves 3, linear to ensiform, 3-5 mm wide, usually shorter than the stem, apparently plane but the margins raised at right angles to the blade and winged, ciliate along the edges, the midrib also slightly raised and winged, sparsely ciliate along the edges, the lower two leaves basal, the uppermost inserted above the ground and sheathing for at least half its length. Stem erect, sparsely puberulent to cil- iate, simple or 1-branched. Spike flexed at the base of the spike, 2—6-flowered, flexuose; bracts membranous, dry and pale brown above, green- ish below, 10-16(—22) mm long, the inner small- er than the outer. Flowers somewhat cupped, 442 white or pale pink to purple (often becoming pink with age), apparently dark in the throat in the type population of G. ixioides; perianth tube 1.5— 2 mm long; tepals narrowly obovate, (13-)18- 28 mm long, 7-10 mm wide. Filaments either equal and 5-6 mm long or unequal with one shorter and 2.5 mm long; anthers 6-8 mm long, yellow. Ovary ca. 2 mm long, style dividing to- wards the apex of the anthers, branches 3-6 mm, recurved. Capsules unknown. Chromosome number, unknown Flowering time. August to September. Distribution. Shale soils especially on south- facing slopes in the Olifants River Valley and the mountains east of Pakhuis Pass. Figure 83. Geissorhiza leipoldtii is native to the arid hills of the Olifants River and northern Cedarberg Mountains and its foothills east of Pakhuis Pass. It is evidently allied to the southwestern Cape species G. inflexa and it resembles especially the larger flowered forms of this species. It can, how- ever, readily be recognized by its puberulous or ciliate stems, only sparsely ciliate leaves and often unequal stamens. Geissorhiza erubescens is un- doubtedly closely allied to G. /eipoldtii. This en- demic ofthe Pakhuis Mountains has small, white to cream flowers with deep red markings on the reverse of the outer tepals, a distinctive fibrous sheath around the stem base and consistently ual filaments. Geissorhiza leipoldtii is poorly collected, probably because plants bloom well only in the years when more favorable rains have fallen. Included in the species are two collections made in the wet spring of 1974 on the upper slopes of Biedouw Pass (Mauve & Oliver 82, Goldblatt 2540) which are unusually robust and have very large flowers, perhaps owing to par- ticularly good rainfall that year. Geissorhiza ixioides Foster is here reduced to synonymy. This species was recognized on the basis of its smaller flowers (tepals ca. 13 mm long) with a dark center (possibly an artifact of drying), supposed short style and equal stamens, in contrast to the supposed unequal stamens of G. leipoldtii. A tendency can be detected in some recent collections of G. /eipoldtii to develop a dark color in the center of the flower but this character has no taxonomic significance. Foster described the filaments of G. leipoldtii as un- equal, one being shorter but examination of the type collection at my disposal including the am- ple material at BOL, not seen by Foster, has shown the filaments of the type of G. leipoldtii ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 to be essentially equal, while type material of G. ixioides has a style of normal length. A collection from the Olifants River Valley near Clanwilliam, Goldblatt 6165, is unusual in having distinctly unequal filaments, one being about half the length of the other two. In the absence of any other differences, this form is in- cluded in Geissorhiza leipoldtii. Specimens examined. SOUTH AFRICA. CAPE: 32 (Clanwilliam) 10 km S of Clanwilliam, clay bank m Goldblatt 6165. 3219 (Wuppertal) Pakhuis-Wuppertal road (AA), Leipoldt s.n. (BOL 20778, K, SAM); slopes above Wel- near S cca Ac BM, BOL, BR, E RE, S, Z), 313a (PRE); jim id pr (B, BM, BOL, BR, G, K, L, MO, P, PH, PRE, S, Z); top of Biedouw Pass, SE slopes, Mauve & Oliver 82 (K, MO, PRE, S); Goldblatt 2540 (K, MO, NBG, S, US, WAG); Cedarberg Forest Reserve, Lang- rug, 1,000 ft. (AC), Viviers 584 (MO); Matjesrivier, Cedarberg (AD), Wagener 214 (NBG). SPECIES INCOMPLETELY UNDERSTOOD The following represent apparently unde- seribed species, but the material available is too letely known to form the basis for the pe of a new species. 1. Geissorhiza sp. 1 (Rogers 28746, Z). A collection of only two plants made by F. A. Rogers at Ceres clearly represents a species not included in this revision. The corm appears to have imbricate tunics which places the species in subgenus Geissorhiza. Other significant char- acteristics include two basal, and a third cauline and partly sheathing, narrow and linear, foliage leaves that have thickened margins and midrib and thus are narrowly 2-grooved on each surface; a puberulent, unbranched stem; an apparently membranous bract in the upper part of the stem; and spikes of 2 long-tubed flowers. The plants are 15-20 mm high and very slender, the leaves about 1 mm wide and half as long as the stem. The flowers appear to be salver shaped, with a ube 9-12 mm long and horizontally extended tepals ca. 16 mm long. The stamens are unequal, the two longer with filaments 5 mm long and a shorter one with a filament 2.5 mm long. The style is of the Geissorhiza type and reaches to about the apex of the anth d has 3 spreading branches. The morphology of the leaves and stem, as well as the flower with unequal stamens all sug- 1985] gest that this species is correctly assigned to sub- genus Geissorhiza, in which unequal stamens are common and pubescent stems not unusual. The affinities of the species are most likely with sec- tion Monticola, most species of which have un- equal stamens, but usually branched stems. However, stem pubescence is unknown in the section. Further material is needed before the species can be described and its relationships more adequately assessed. Locality information is unfortunately limited to Ceres and the altitude 1,400 ft. Ceres may represent the town itself, the immediate surroundings or the district that ex- tends well to the north. The altitude suggests that the species grows on lower to middle mountain slopes. 2. Geissorhiza sp. 2 (Stokoe s.n., SAM 65812). A collection of a small species, made in De- cember 1952 by T. P. Stokoe at Oudebosch in the Palmiet River Mountains, appears to rep- resent either an undescribed species or a very aberrant form of a known one. The plants are 10-15 cm high, have linear, plane leaves and smooth, simple or 1-branched stems with soli- tary flowered spikes. The corms are small and apparently have soft-textured tunics that do not accumulate to any degree, and it is not possible to determine whether the tunics are concentric or imbricate. Accordingly, the species cannot be assigned to subgenus with certainty. The overall appearance of the plants and the soft plane leaves, however, suggests that the species belongs in sec- tion and subgenus Weihea. The branching pat- tern is unusual. The branches are almost equal in length and so appear dichotomous. This type of branching is known in Geissorhiza only in G. geminata, also section Weihea and it is to this species that G. sp. 2 is assumed to be allied if not simply a depauperate form of G. geminata. More material of the species is needed before a final decision can be taken on the status of this collection. EXCLUDED SPECIES Geissorhiza abyssinica R. Br. ex A. Richard, Tent. ss. 308. 1850 — Lapeirousia abyssinica (R. Br. ex A. Rich- ard) Baker (Baker, 1898; Cufodontis, 1972). Geissorhiza albens E. Mey. in Drége, Zwei Pflan- ngeogr. Dokum. 187. 1843, nom. nud. — Gladiolus debilis Ker (Lewis et al., 1972). GOLDBLATT— GEISSORHIZA 443 Geissorhiza alpina Hook. f., J. Linn. Soc., Bot. 7: 233. 186 = Hesperantha petitiana (A. Richard) Baker. Geissorhiza ambongensis H. Perrier, Notul. Syst. (Paris) 8: 130. 1939. = ? Tritonia sp. (Goldblatt, 1982b). Geissorhiza anemonaeflora (Jacq.) Klatt, Lin- naea 34: 657. 1866. Ixia anemonaeflora Jacq. Icon. Pl. Rar. 2: tab. 273. 1793. = Ixia campanulata signs er Lewis (1962). Geissorhiza bojeri Bake . 14: 239. 1876. = Gladiolus bojeri rae een (Goldblatt, 1982b). Geissorhiza briartii Wildem. & Durand, Bull. Soc. Roy. Bot. Belgique 39: 105. 1900 = Lapeirousia erythrantha (Klotzsch ex Klatt) Baker var. briartii (Wildem. & Durand) Geerinck, Lisowski, Malaisse & Symoens eerinck et al., 1972 Geissorhiza erecta Baker, J. Bot. n.s. 5: 238. 1876. = Hesperantha erecta (Baker) Benth. ex Baker (Goldblatt, 1984). Geissorhiza gracilis Baker, Handb. Irid. 155. = Gladiolus parvulus Schltr. (Lewis et al., 1972). Geissorhiza grandis Hook. f., Bot. Mag. 96: tab. 5877. 18 A floribundus Jacq. subsp. milleri (Ker) Oberm. (Lewis et al., 1972). A “a run rg la Roche) Baker, J. Linn. i. 1878. = Ixia pus p la Roche (Lewis, 1962; Goldblatt & Barnard, 1970) Geissorhiza longituba Klatt, Linnaea 35: 383. 1867-1868. — Hesperantha longituba (Klatt) Baker (Gold- blatt, 1984 s lutea Ecklon, Topogr. bag Pflan- zensamml. Ecklon 21. 1827, n = usaspa, E falcata (L. f.) a (Goldblatt, 1984) (syn. Hesperantha lutea Ecklon e Baker). Geissorhiza macra Baker, Bull. Herb. Boissier, r. 2, 4: 1004. 1904. TYPE: South Africa. Transvaal: Mt. vra Jacottet s.n. (G, not seen; GH, pho = Hesperantha sp., n fairly large flowers are borne singly on the stems. Geissorhiza minima Baker, J. Bot. n.s. 5: 239. 1876 — Hesperantha minima (Baker) Foster (Foster, 1948; Goldblatt, 1984). Geissorhiza patersoniae L. Bolus, Ann. Bolus Herb. 1: 132. 1915. 444 — Gladiolus stellatus Lewis (Lewis, 1966; Lew- is et al., 1972) Geissorhiza pauciflora Baker, Bull. Herb. Bois- sier, Ser. 2, 10: 1004. 1904. — Hesperantha falcata (L. f.) Ker (Goldblatt, 1984). Geissorhiza quadrangula (de la Roche) Ker, Irid. Gen. 88. 1827. Ixia quadrangula de la Roche; Descr. Pl. Aliq. Nov. 16. 1766. — Gladiolus quadrangulus (de la Roche) Bar- nard (Goldblatt & Barnard, 1970). vifolia (Poiret) Klatt sensu Klatt, Linnaea 34: 655. 1866. Ixia bees es [sic] iie in — e Suppl 3(1): 2 813, om. superfl. pro x. recurva Red. ise, pee tab. 251, f. 2. 1809. = Romulea cf. flava (Lam.) De Vos (De Vos, 1972 Geissorhiza ). Geissorhiza schlechteri Baker, Bull. Herb. Bois- sier, Ser. 2, 9: 43. 1 — Hesperantha schlechteri (Baker) Foster (Foster, 1936). Geissorhiza sublutea (Lam.) Ker, Ann. Bot. (Kónig & Sims) 1: 224. 1804. Ixia sublutea Lam., Encycl. 3: 335. 1789. = Romulea triflora (Burm. f.) N. E. Brown (De Vos, 1972). Geissorhiza vaginata Sweet, Brit. Flow. Gard., Ser. 1, tab. 138. 2 = Hesperantha vaginata (Sweet) Goldbl. PSN latt & EE 1970; Goldblatt, 984). LITERATURE CITED AXELROD, D. I. & P. H. Raven. 1978. Late Creta- ceous and Tertiary vegetation history of Africa. Pp. 77-130 in M. J. A. Werger (editor), Biogeog- e and Ecology of Southern Africa. Junk, The Pie J. S 1876. New species of Ixieae. J. Bot. 14: 236-239. . 1878. Systema Iridacearum. J. Linn. Soc., Bot. 16: 61-180. ——l 1892. Handbook of the Irideae. George Bell & E London 1896. Irideae. In W. T. Thiselton-Dyer ed: itor), Flora Capensis 6: 7-171. Reeve & Co, Ash ford, 1898. Irideae. In W. T. Thiselton-Dyer (ed- itor), Flora of oe Africa 7: 337-376. Lovell e & Co, "1904. Beitrag z zur Kenntnis der Afrika- e. Bull. Herb. Boissier, Sér. BERGIUS, P.J. 1767. Descriptiones Plantarum Capite Bonae Spei. Salvius, Stockholm ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 Borus, H. M. L. 1930. Plants new and noteworthy: Geissorhiza heterostyla and G. inaequalis, new species. S. African Gard. 20: 345-346. . 1931. Plants new and noteworthy: Geisso- rhiza bellendenii MacOwan and G. eurystigma L. Bol. r African Gard. 21: 281-282. G. J. Lewis. 1933. Plants new and note- worthy: Acidanthera roseoalba Lewis. S. African Gard. 23: 255 BREMER, K. 1976. ' The genus Relhania (Compositae). 0: r5 The Iridaceae of Thunberg's Herbarium. J. Linn. Soc., Bot. 48: 34-42. 29 | .T d ^ a pensis Prodromus. Kew Bull. 1929: 129-139. des Kewenses CXXVII. Kew Bull. Travels in the Interior of gma ce J.L. (f. M 1768. Florae Capensis Prodromus. 1 n. Bone] B. L. 1970. Infraspecific categories in flow- ering plants. Biol. J. Linn. Soc. 2: 233-238. ud i tarum Aethi- piae Spe phyta. Bull. Jard. Bot. Natl. Bel- save. si Suppl. i 1579-1657. DAHLG . M. 1968. Distribution and sub- adi in ere ies African genus Aspalathus (Le- guminosae). Bot. Not. 121: 505-534 1972. The genus dee in South E: African Bot. Suppl. 9: 1-307. — —, "19 ol ontwikkeling by so sommige genera van die ru en die systematiese posisie. Tyds. Natuurwetensk. 1833 DIETRICH, A. I s. Plantarum. 6th edition. li n C. F. Ecklon. Reise EISTNER. 1971. A degree ref- erence system for citing biological records in southern Africa. Mitt. Bot. Staatssamml. Mün : 501-509. ELLIOT, "d "i Sc COTT 1891. i on the fertilisation f Ann. pn 5: 333- e II. Ar revision of Geissorhiza Ker-Gawl. Contr. oan Herb. 135: 1 78 1948. Studies in Iridaceae V. Some new o noteworthy species of Hesperantha. Contr. Gray Herb. 166: 3-27. FUNK, V. A. 2. The systematics of Montanoa (Asteraceae, Heliantheae). Mem. New York Bot. Gard. 36: 1-133. GEERINCK, D., S. Lisowski, F. MALAISSE & J. J. SYMOENS. 1972. Le genre Lapeirousia Pourr. (Iridaceae) au Ziare. Bull. Soc. Roy. Bot. Bel- gique 105: 333-351 GOoLDBLATT, P. 1971. Cytological and morphological studies in the Southern African Iridaceae. J. S African Bot. 37: 317-460. ————. 1976a. Evolution. cytology and subgeneric 1985] — in Moraea (Iridaceae). Ann. Mis- . Gard. 1-23. 3: The genus Moraea in the winter rain- fall a area of Southern Africa. Ann. Missouri Bot. Gard. 63: 657-7 — 78. An analysis of the flora of Southern frira: ite Ann. Missouri. Bot. Gard. 65: 369— 436 and kary change in Galaxia (Iridaceae). PL gent Evol. 3 161-169. 1980. Redefinition of Homeria and Moraea (Iridaceae) in the light of biosystematic data, with Rheome gen. nov. Bot. Not 1981. prag and biology of Homeria is MED . Missouri Bot. Gard. 68: 413- ` 1982a. Corm morphology in Hesperantha (Iridaceae — Ixioideae) and a proposed infragener- ic taxonomy. Ann. Missouri Bot. Gard. 69: 370- 378. . 1982b. Notes on Geissorhiza: the species of Madagascar. Ann. Missouri Bot. Gard. 69: 379- 381. . 1983. of Thunberg's Herbarium Nordic J. Bot. 3: 437—442. 1984. A revision of Hesperantha (Iridaceae) in the winter rainfall area of Southern Africa. J. S. African Bot. 50: 15-141. . BARNARD. 1970. The Sg c Daniel de la Roche. J. S. African Bot. 36: The species of Geissorhiza (Iridaceae) (Uppsala, Sweden). 318. GRANT, V. sie Organismic Evolution. Freeman, GwYNNE, E. 1958. Cytological studies in the Irida- ceae. Cytologia 23: 68-71. HENNIG, 1 Phylogenetic Systematics. Univ. of Illinois Press, Urbana (translated by D. D. Da- Za ` Natuurlyke Historie part 2, 12. š C ocn and cladistic tudies i in Anacyclus (Compositae: Anthemideae). Nordic J. Bot. 1: 83-96. KER (GAWLER), J. B. 1802a. Ixia rochensis (a). Plaid pe t. Mag. 17: tab. 598. 02b. Ixia secunda. One-ranked Ixia. Bot. TA 17: tab. 597. . 1802c. Ixia maculata, var. a viridis (v). Green- stained Ixia. Bot. Mag. 16: tab. 549. 1803. Geissorhiza obtusata. Yellow-flowered Geissorhiza. Bot. Mag. 18: tab. 672. Ensatorum Ordo. Ann. Bot. (Kónig & Sims) 1: 219-247. — Np 1810. Geissorhiza setacea. Narrow leaved tile- root. Bot. Mag. 31: — 1827. Iridearum Gen a. De Mat, Brussels. Katt, F. W. 1866. Revisio Iridea (Conclusio). Linnaea 34: 537—689. . Erganzungen und Berichtigungen zu Baker's Systema Iridacearum. Abh. Naturf. Ges. Halle 15: 337-404. GOLDBLATT— GEISSORHIZA 1895. Irideae. Jn T. Durand & H. Schinz (editors), Conspectus Florae Africae 5: 143-230. Lewis, G. J. 1933. Plants new and noteworthy — Aci- danthera roseoalba. S. African Gard. 23: 255, 266. . Iridaceae. New genera and species and miscellaneous notes. J. S. African Bot. 7: 19-59. 50. Iridaceae. Pp. 217—264 in R. S. Ad- amson & T. M. Salter S Flora of the Cape Peninsula. Juta, Ca 1954. Some aspects ofi the Yi Vii phy- logeny and taxonomy of the daceae. Ann. S. African Mus. 40: S113. . 1959. The genus Babiana. J. S. African Bot. Suppl. 3: 1-149. . 1962. South African Iridaceae. The genus Ixia. J. S. African Bot. 28: 45-195. . 1966. Thunberg’s South African species of uai dris name changes. Bot. Not. 119: 295. A. A. OBERMEYER & T. T. BARNARD. 1972. A revision of the South African species of Gladi- . S. African Bot. Suppl. 10: 1—316. LINNAEUS, C. (f). 1782. Supplementum Plantarum. Orphanotropheus, Braunschweig. MEYER, E. 1843. Zwei A air mud ographische Doc- umente von J. F. Drége. Flora 26 (Beigabe): 1- Types of Ecklon’s ‘Topogra NORDENSTAM, B. phis edish Museum of 2 1972. ches Mesa d in the Sw History. J. S. African Natural Bot. 38: 277—298. PARKER, D. 82. The western Cape lowland fyn- bos — what is there left to cuc Veld & Flora 64: 98-101. RocHE, D. DELA 1766. Descriptiones Plantarum Ali- uot Novarum. Verbeek, Leiden ROEMER, J. J. & J. A. SCHULTES. 1817. Systema Ve- getabilium Secundum. Volume 1. Cottae, Stutt- 1972. Taxonomic studies on Leuco- S. African Bot. Suppl. 8: 1- gart. ROURKE, J. P. spermum R. Br. J. 194. SCHLECHTER, R. 1899. Plantae Schlechterianae novae vel minus cognitae describuntur II. Bot. Jahrb. Syst. 27: 86-220. SCHULTES, J. A. 1 Mantissa in Volumen Primum Systematis Vegetabilium. Cottae, Stuttgart. Simpson, B. B. 19 Contrastin ting modes of evolution of two groups of Perezia (Mutisieae; Compositae) of southern South America. Taxon 25: 525-536. SiMPSON, G. G. 1953. The Lam — of Evo- lution. Columbia Univ STAFLEU, F. A. 1978. Soothe Code of Botanica menclature. Bohn, Holkema & Schelte 978. €— Invaders — Beautiful but s. Department of Nature and Environ- ment enisi tein "a the Cape EE Cape Utr ; STIRTON, C. H. 1830. Sweet's Hortus Britannicus. 2nd edi- es acie London. _ Dissertatio de Ixia. O. J. SWEET, R l am THUNAE, C. Am 300. Prodromus Plantarum Capensium. Part 2. Edman, Uppsala. 1803. Novae species plantarum em examine et descriptae. Phytogr. Blatt. 1: 1—5. 446 ANNALS OF THE MISSOURI BOTANICAL GARDEN . 1807. Flora Capensis. 1st edition. Volume 1. Edman, Uppsala. ——. 1810. Beskrivelse over 19 Arter af Gladiolus fra Afrikas sondre Odde. Skr. Naturhist.-Selsk. 6: 1-15. . 1818 (or 1820). Flora Capensis. 2nd edition. gen Enumeratio Plantarum. Volume 2. Moller & fil., Copenhag VOGEL, S. 1954. Blutenbiologische Typen als Ele- mente der Sippengli pend Bot. Stud. 1: 1-338. WAGNER, W. ed FA OL phology in Hesperantha (Iridaceae) Ann. s- Gard. 71: 176. souri Bot. WEIMARCK, H. 1 Bu ccm groups, centres and intervals within Hii Cape flora. Acta Univ. Lund. n.s. 37(5): 1-14 INDEX (See p. 344 for excluded species.) Acidanthera Hochst. 303, 367 exscapa i ) Baker ex Bouche & Wittmack 378 i 31 section nanoia Goldbl. 343 section Cia Goldbl. 426 sectio gysiphon (Lewis) Goldbl. 367 section cA pa section 7ncludanthera Goldbl. 340 section Intermedia Goldbl. 382 section c section Planifolia Goldbl. 417 tion Pusilla G section Rochea 8 section Tortuosa Foster 336 section Weihea Ecklon ex Baker 309 subgenus Eugeissorhiza Foster 382 subgenus Geissorhiza 382 subgenus /xiopis Foster 359 subgenus Weihea (Ecklon ex Baker) Goldbl. 309 en Foliosae Foster 309 section Ventricosae Foster 387 atticola Goldbl. 361 arenaria Ecklon ms. 389 s Goldbl. 434 aspera Goldbl. 417 aurea Ecklon 390 barkerae Goldbl. 393 bellendenii MacOwan 422 — (Thunb.) N. E. Brown 390 . macowani Foster 390 brehmii Ecklon ex Klatt 396 brevituba (Lewis) Goldbl. 368 bryicola Goldbl. 413 burchellii Foster 406 candida Ecklon 436 cataractarum Goldbl. 364 cedarmontana Goldbl. 341 ciliaris (Ker) Jackson 436 ciliaris Salisb. 430 ciliatula Goldbl. 415 confusa Goldbl. 373 corrugata Klatt 336 cyanea Ecklon darlingensis age 348 Idbl. 3 erosa (Salisb.) Foster 436 var. kermesiana (Klatt) Foster 437 erubescens Goldbl. 440 esterhuyseniae Goldbl. 341 Ker exscapa (Thunb. ) Go 378 filifolia Baker 354 flava Reichb. ex Klatt 326 foliosa Klatt 317 fourcadei (L. Bolus) Lewis 315 fi geminata E. Meyer ex Baker 327 graminifolia 438 var. bicolor Baker 438 grandiflora Goldbl. 408 hesperanthoides Schltr. 362 heterostyla L. Bolus 430 hirta (Thunb.) Ker 436 hispidula (Foster) Goldbl. 349 humilis (Thunb.) Ker 346 var. bicolor Baker 349 var. grandiflora Baker 346 var. hispidula Foster 349 . humilis Baker 346 indicar: (de la Roche) Ker subsp . bicolor (Thunb.) Sogni 390 var. usata (Ker nn 390 n L. B inconspicua B in inflexa (de la L. Ns 436 intermedia Goldbl. 352 ixioides Schltr. ex dei 441 juncea (Link) A. D var. pallidiflora (Schltr. E aid 354, 384 kamiesmontana Goldbl. 4 1985] karooica Goldbl. 338 larochei (Roem. & Schult.) Loudon 404 leipoldtii Foster 441 lewisiae Foster 422 lithicola Goldbl. 343 longifolia (Lewis) Goldbl. 371 louisabolusiae Foster 395 var. longifolia Foster 396 malmesburiensis Foster 326 obtusata Solander ex E 390 ornithogaloides Klatt subsp. marlothii (Foster) Goldbl. 325 outeniquensis Goldbl. 313 ovalifolia Foster 329 ovata (Burm. f.) Aschers. & Graeb. 333 pallidiflora Schltr. 354 pilosa Pappe ms. AU plicata Pappe m pseudinaequalis Goldbl. 415 pusilla (Andr.) Klatt 339 quinquangularis Ecklon ex Klatt 436 var. atrofaux Foster 438 radians (Thunb.) Goldbl. 404 “radi E ramosa (Ker) Klatt 410 ramosa Ker ex Klatt 410 recurvifolia (Poir.) Klatt 320 rocheana Sweet 404 u— (Ker) Ker 400, 404 var. monantha (Sweet) Baker 400, 422 var. Wee psp (Ker) Baker 400, 404 rogersii N. E. Brown 430 romuleoides Ecklon 326 rosea Ecklon 436 rosea (Klatt) Foster 430 To a ee 312 cunda F. rupestris Sen p sabulos 89 china (Baker) Goldbl. 368 scillaris r. 384 eti similis Goldbl. 382 GOLDBLATT — GEISSORHIZA spiralis (Burchell) De Vos 337 spiralis (Burchell) De Vos ex Goldbl. 337 splendidissima Diels 435 stenosiphon Goldbl. 358 striata Ecklon ms. 349 subrigida L. Bolus 430 sulphurascens Schltr. ex Foster 398 2 ri cca Goldbl. 410 elatior Ecklon ms. 320 Pc Goldbl. 377 teretifolia Lewis 396 tulbaghensis F. Bolus 424 tulipifera Klatt 404 umbrosa Lewis 359 unifolia Goldbl. 353 violacea Baker 309 spe ch 378 zt ex Klatt 430 inflexa (de la E Foster 436 kermesina Klatt 4 quinquangularis Ecklon 438 rosea KI 0 uias Baker 354 Ixia L. 302 mia Thunb. 390 ciliaris Salisb. ex Ker 436 erosa Salisb. 436 excisa L. f. 331, 333 imbricata (de la Roche) Ker 387 inflexa de la Roche 436 juncea Link 353 larochei Roem. & Schult. 404 monanthos Thunb. 422 ornithogaloides Licht. ex Roem. & Schult. 325 ovata Burm. f. 333 phalangioides Roem. & Schult. 384 pusilla Andr. 339 radians Thunb. 404 ramosa Ker 384 rochensis Ker 400, 404 scillaris Thunb. 384, 389 secunda de la Roche 417 secunda Ker d Thunb. | a Burm. f. 3 M o secunda Pappe ms. 373 Rochea venusta Sa Romulea spiralis (BurchelD Baker 337 —— Klat pa (Th unb) Klatt 373, 378 excisa (Burm. f.) Ecklon 333 NOTES A NEW SYNGONANTHUS (ERIOCAULACEAE) FROM SOUTHERN MEXICO While preparing a treatment of the Eriocau- laceae for “Flora Mesoamericana,” I discovered two recent collections of a distinctive species of Syngonanthus from the State of Chiapas in southern Mexico here described as S. davidsei Huft. This is only the second species of the genus to be found in Mexico. Syngonanthus caulescens has been collected in bogs near Minatitlán, Ve- racruz (2 Feb. 1892, Smith 354, MO), and is also known from Costa Rica and northern South America. The only other species of this largely South American genus that are known from con- tinental North America are S. flavidulus (Michx.) Ruhl. of the southeastern United States, S. pit- tieri from Panama, and four poorly-known species endemic to Belize. Syngonanthus davidsei Huft, sp. nov. TYPE: Mex- ico. Chiapas: Municipio of Ixtapa, grassy flats with Quercus, Acacia, and Byrsonima near Ixtapa, 915 m, 1 Nov. 1981, Breedlove & Davidse 54339 (holotype, CAS!; F neg. no. 59185; isotypes, to be distributed to ENCB, MEXU, MO, not seen). -lancenlata ras parvae perennes, acaulescentes. Folia li- Pe- dupcali aliquot, terminales, glanduloso- pilosi. Capi- tula solitaria, a ovatae, Apna aureo-brunneae, glabrae, apice ob- tusae vel parum acutae; bracteolae receptaculares nul- lae ceterum glabri; sepala à ô late obovato- falcata, aureo-brunnea; sepala 9 elliptico-oblonga, basi alba, apice aureo-brun- nea; petala ? anguste oblonga, sepalis parum breviora alba, basi libera, versus apicem conniventia vel leviter connata. ` Small perennial herb, monoecious, acaules- cent. Leaves tufted, spreading or recurved, often flat on the ground, linear-lanceolate, 1-1.5 cm long, 0.8-1.2 mm wide, moderately strigose, the apex acuminate. Peduncles several, terminal, 6— 17 cm long, slender, glandular-pilose, the hairs spreading; sheath 12-20 mm long, appressed, densely glandular-pilose, the apex long-acumin- S, broadly ovate, navicular, 1.8-2 mm long, 0.8-1 mm wide, golden brown, glabrous, the apex ob- ANN. Missouni Bor. GARD. 72: 448—449. 1985. tuse to acute: receptacle flat, glabrous; recepta- cular bracts lacking. Staminate florets stipitate, the stipe 0.3-0.5 mm long, long-ciliate at base; sepals 3, free, broadly obovate-falcate, navicular, ca. 1 mm long, ca. 0.5 mm wide, glabrous, golden brown, the apex obtuse to acute; petals connate into a narrow, glabrous, stramineous tube, the x with 3 short lobes; stamens 3, the anthers white. Pistillate florets stipitate, the stipe ca. 0.5 mm long, long-ciliate at base; sepals 3, free, el- liptic-oblong, 1.3-1.5 mm long, ca. 0.4 mm wide, dai ipn D toward base, golden brown to- ard a e apex acute; petals 3, narrowly oblong, slightly shorter than the sepals, glabrous, long, united ca. two-thirds their length; seeds not seen. Additional specimen examined. MEXICO. CHIAPAS: Municipio of La Trinitaria, 10 km E of La Trinitaria on road to Lagos de Montebello National Park, sandy ats with Pinus and Quercus, 1,555 m, 7 Nov. 1981, Breedlove & Davidse 54973 (CAS). This species is named in honor of Dr. Gerrit Davidse, who was instrumental in establishing the “Flora Mesoamericana” project. Syngonanthus davidsei is very similar to the four species endemic to Belize. Indeed, with the exception of S. caulescens, the Mesoamerican species are nearly identical in aspect and differ chiefly in the pubescence of the peduncle and in minor characters of the involucral bracts and sepals. Because of the great rarity of these six species (they are known from a } total of only 12 collections), a proper ic value of their characters is Senata at this time. In fact, since three of the Belize endemics (S. hondurensis, S. lundellianus, and S. oniellii) are known only from collections made by Hugh O’Niell in August and September 1936 within a few miles of Boomtown, it is tempting to con- sider these as belonging to a single, somewhat variable, species. It is therefore with some hes- itation that I venture to describe a new species in this group. However, S. davidsei seems dis- tinctive on the basis of its broadly ovate, blunt involucral bracts and its broadly ovate-falcate A 71117111 1985] NOTES 449 staminate sepals, which are unlike those of any city of information available, recognition of these of the other Mesoamerican species. six species seems unavoidable. Further collections of Syngonanthus from Me- Syngonanthus davidsei may be separated from soamerica are much to be desired in order to its Mesoamerican congeners by means of the fol- arrive at a proper taxonomic understanding of lowing key: these species. For the time being, given the pau- la. Stems elongate; leaves arranged along stem (Veracruz, Costa Rica) _ S. caulescens (Poir.) Ruhl. lb. Stems very short, the plants essentially acaulescent; leaves tufted. 2a. Peduncles glabrous, or nearly so. 3a. Pistillate sepals glabrous; staminate corolla tube with petals free only at apex (Panama) .. S. pittieri Mold. 3b. Pistillate sepals long-ciliate on the margins; staminate petals connate at middle, free at base and at apex (Belize) S. oniellii Mold. 2b. Peduncles conspicuously pubesc 4a. Involucral kn hyaline, olores the — white (Belize) S. bartlettii Mold. 4b. teni bracts golden brown 5a. Involucral bun broadly iei bt t taminate sepals t dl E obtuse to acute at apex, ria brown (Chiapa s) Cip ai Huft 5b. Involucral bracts lanceolate to narrowly veg or E acute to Dn at apex; staminate — elliptic, acuminate at apex (Be 6a. Pubesc sis peduncles strongly Lima ak a few pow glanduliferous hairs prese S. hondurensis Mold. 6b. Pd of peduncles densely spreading glandular-pilose ........ S. /undellianus Mold. I am grateful to Dr. Frank Almeda (CAS) for —Michael J. Huft, Missouri Botanical Garden. the loan of specimens and for information re- Mailing Address: Department of Botany, Field garding the disposition of the isotypes. Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605. BOOK REVIEW Snijman, D. 1984. A revision of the genus Haemanthus L. (Amaryllidaceae). J. S. African Bot. Suppl. 12: 1-139. Since the recognition of Scadaxus by Friis and Nordal in 1976, for tropical African species until then treated as Haemanthus, the genus has had a restricted application to species with true bulbs and somewhat fleshy distichous leaves. The ge- nus as thus circumscribed is restricted to south- ern Africa where it has a wide range. This ex- cellent revision by Dierd nbosch in Cape Town, is the only comprehensive treatment of the genus since “Flora Capensis” (1896) and ad- mits 21 species of which four are new The study is the result of patient research over the several years needed to complete fieldwork and grow and flower the plants to observe the entire life cycle. Haemanthus presents the work- er with an unusual problem: leaves of most species are produced after flowering. Thus populations found in flower must be revisited or grown to obtain leaf material, which in several cases is essential for species identification as well as for the preparation of type material. Conversely, plants collected in leaf had to be grown, some- times for several years, before flowers could be obtained. The result of this labor is presented in a beautifully illustrated monograph that will be the envy of many a systematist. Twenty-four species and subspecies are illustrated with wa- tercolors by Elaphie Ward-Hilhorst (and one is by Fay Anderson). The artwork and reproduc- tion are of such quality that the plants come to life on the pages. Haemanthus is concentrated in the winter rainfall area of southern Africa; which includes extreme southwestern Namibia, the south and west coasts, and interior of the Cape Province of South Africa. The pattern of speciation is one that is typical of many geophytic genera in south- ern Africa. There are a limited number of wide- spread and often in the summer rainfall part of southern Africa. Then in contrast, there has been marked specia- tion in the south and west and many ofthe winter rainfall species are very restricted in distribution although well-known species such as H. cocci- neus and H. pubescens have wide ranges in this region.— Peter Goldblatt, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. Volume 71, No. 2, pp. 344-630 of the ANNALs or THE MISSOURI BOTANICAL GARDEN, was published on 19 March 1985. Volume 71, No. 3, pp. 631-986 of the ANNALs OF THE MISSOURI BOTANICAL GARDEN, was published on 19 April 1985. Volume 71, No. 4, pp. 987-1187 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN, was published on 14 May 1985. Volume 72, No. 1, pp. 1-166 of the ANNALS or THE MIssoURI BOTANICAL GARDEN, was published on 10 June 985. ANN. Missouni Bor. GARD. 72: 450. 1985. Studies in Fuchsia This special issue of the ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 69, no. 1, 1982) is devoted to several papers on the systematics of the genus Fuchsia (Onagraceae). The large section Fuchsia, which contains such horticul- turally important species as F. corymbiflora, F. triphylla, and F. fulgens, contains 61 species, about 6096 of the entire genus. Paul Berry's monograph treats the section in detail; extensive descriptions of flowers, blooming periods, habitats, and distribution are given. These are supplemented with beautiful color plates of several species. In addition to traditional keys to all the species, principal mor- phological differences between similar species and hybrids are treated in table form. CONTENTS The Systematics and Evolution of Fuchsia Sect. Fuchsia (On- agraceae) | Poul E Demy... V ous a n o 5 Pollinator Maintenance vs. Fruit Production: Partitioned Re- productive Effort in Subdioecious Fuchsia lycioides. oe Peter R. Atsatt & Phillip Ra de |... o a The Mexican and Central American Species of Fuchsia (On- agraceae) except for Sect. Encliandra. X "ne Dennis E. Breedlove, Paul E. Berry & Peter H. Raven 2 236 Index 245 ORDER FORM Please send me copy/ies of Studies in Fuchsia at $7.50. 3. Make check or money order payable to Missouri Botanical Garden, in U.S. funds, and payable through U.S. bank. Date: d Payment enclosed Ship to: d Send invoice ($1.00 fee will be added to total) COCOS UIN DNI ERE EE Send order to: M MN E o gueuiianueue EE Department Eleven Missouri Botanical Garden P.O. Box 299 St. Louis, MO 63166-0299 U.S.A. ee EET To place an order, use this form or a photocopy of it. Contents continued from front cover NOTES A New Syngonanthus (Eriocaulaceae) from Southern Mexico Michael 448 Book Review 450 ANNALS F THE SSOURI BOTANICAL GARDEN JUME 72 1985 NUMBER 3 New Memecyleae CONTENTS The Histogenesis and Evolution of Integuments in Onagraceae Hiroshi ERE Peter AH. Raven . L — ———— oen Chromosome Numbers in Compositae, XV: Liabeae Harold Kaon: A. Michael Powell, Robert M. King & James F. Weedin Chromosome Studies on Hydromystria laevigata (Hydrocharitaceae) Eduardo A. Moscone & Luis M. Bernardello 480 Studies in Neotropical Paleobotany. III. The Tertiary € 'ommunities of Pan- ama— Geology of the Pollen-bearing Deposits Alan Graham, R. H. : x Stewart & J. L. Stewart T Studies in Neotropical Paleobotany. IV. The Eocene Communities of Pan- ama Alan Graham i = ^ New Cymbopetalum (Annonaceae) from Costa Rica and Panama with edid Observations on Natural Hybridization George E. Schatz 533 Notes on and Descriptions of Seven New Species of Mesoamerican Cleth- : Taceae Clement W. Hamilton 539 B umelia reclinata var. austrofloridensis (Sapotaceae), a New Variety from South Florida, U.S.A. R. David Whetstone Fi UN Taxa of New World Memecyleae (Melastomataceae) forley Thomas VOLUME 72 AUTUMN 1985 NUMBER 3 ANNAL OF THE MISSOURI BOTANICAL GARDEN The ANNALS, published quarterly, contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the Garden will also be ac- cepted. 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 first issue of this volume. EDITORIAL COMMITTEE NANCY Morin, Edito. Missouri Botanical osa CHERYL R. BAUER, Editorial Assistant Missouri Botanical Garden MARS ER CRO Nie pesi sasa ss ERRIT DAVIDSE Missouri Botanical Garden JOHN D. DWYER Missouri Botanical Garden & St. Louis University PETER GOLDBLATT Missouri Botanical Garden For subscription information contact the Business Office of the Annals, P.O. Box 299, St. Louis, MO 63166. Subscription price is $65 per volume U.S., $70 Canada, and Mex $75 all other countries. Personal subscriptions are available at $30 and oF fhe irmail delivery charge, $30 per volume. Four issues per volume. The ANNALS OF THE MISSOURI BOTANICAL GARDEN (ISSN 0026-6493) is pu lished quarterly by the Missouri Botanical Garden, 2345 Tower Grove Ave., St. Louis, M 63110. Subscription price is $65 per volume U.S., $70 Canada and Mexico, $75 all esis countries. Personal subscriptions are available at $30 and $35, respectively. Second C postage paid at St. Louis, MO and additional mailing offices. POSTMASTER: Send m changes to the ANNALS OF THE MISSOURI BOTANICAL GARDEN, P.O. Box ^'^ St. Louis, MO 63166 tively. € Missouri Botanical Garden 1985 OF THE ANNALS MISSOURI BOTANICAL GARDEN VOLUME 72 1985 NUMBER 3 THE HISTOGENESIS AND EVOLUTION OF INTEGUMENTS IN ONAGRACEAE! HIROSHI TOBE? AND PETER H. RAVEN? ABSTRACT The mode of initiation and subsequent growth of the inner (i.1.) and outer (o.i.) integuments were a and compared i in 40 species, representing : all seven tribes and all 17 genera of Onagraceae. The our different types of developmental iude of the o.i. are characterized: D o.i. of subdermal origin— — Lopezia (Lopezieae), Fuchsia (Fuch- sieae), and Circaea (Circaeeae); 2) 0.1. subdermal cells dividing more a (Onagreae); 3) o.i. of both derm less actively than those of the ame l cells— Hi 4) o.i. of dermal o sonia, Clarkia, Gayophytum, Gongy both dermal and subdermal origin, actively than ee 2 the dermal cells— Oenothera and Stenosiphon subderm with derivatives of the ay, RRE and Calylophus and Gaura (Onagreae); rigin— Ludwigia (J Sere. on aac and Epilobium (Epilobieae); and Camis- locarpus, Heterogaura, and unexpected close similarity of Hauya (Hauyeae) to peor dbs iin and Gaura (Onagreae), among which is shared an apparently derived state of this character ra e patterns of relationships suggested on the basis of the ae Sea mode of. the o.i. closely accord with th viden derived from other lines of evidence. The results from Onagraceae confirm that the is bot characters of integuments are valuable indicators of relationship at a generic level. The family Onagraceae, composed of seven tribes, 17 genera, and about 674 species (Raven, 1979), is one of the most intensively investigated families of angiosperms. In recent years, in ad- dition to systematic studies (e.g., Raven, 1963, 1964, 1969; Plitmann et al., 1973), there have also been studies on chromosome number and morphology (e.g., Kurabayashi et al., 1962; Ra- ven & Gregory, 1972; Raven & Tai, 1979), wood anatomy (Carlquist, 1975, 1977, 1982), pollen morphology (e.g., Skvarla et al., 1975, 1978), floral anatomy (Eyde & Morgan, 1973; Eyde, 1977, 1978, 1981, 1982), and leaf anatomy (Keating, 1982). As a result of these detailed studies, our knowledge of phylogenetic relation- ships within the family at a generic level has become quite detailed. he major outlines of the pattern of relation- ships that have been revealed are as follows. Lud- wigia represents a line distinct from all other genera of the family (Eyde, 1977, 1978, 1981; Raven & Tai, 1979). It shares with two of the more primitive genera in the other phylogenetic branch of the family, namely Fuchsia (Fuchsieae) and Hauya (Hauyeae), the generalized charac- teristic of lacking interxylary phloem (Carlquist, 1975, 1977). In addition to Fuchsia and Hauya, Lopezia (Lopezieae) and Circaea (Circaeeae), in ! We are grateful to the U.S. National Science Foundation for support of studies of Onagraceae through grants F. to Peter Raven, most recently grant BSR82-14879, and to Drs. criteal reading of and comments on the manuscript. Bouman and F. D. Boesewinkel for their epartment of Biology, Chiba University, 1-33 Yayoi-cho, Chiba 260, Japan. 3 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. ANN. Missouni Bor. GARD. 72: 451—468. 1985. 452 | WS 2 X TOBE & RAVEN — INTEGUMENTS IN ONAGRACEAE 453 Ludwigia peploides. — A-E. Median longisections of the o.i. in es ead successive stages o the FIGURE 2. development showing t development of a distinctive three-lay of ered part o rmal initials of the o.i.; e', daughter cell ed the dermal initial e, and functioning as an initial A p ‘third (middle) layer of the Ow o.i. E. scales — 10 which at least the most primitive species have a basic chromosome number of n = 11, present an array of relatively primitive characteristics, so that Circaea, Fuchsia, Hauya, and Lopezia may be regarded as distantly related, generalized lines within this major branch of the family—the branch that does not include Ludwigia. Epilo- bieae, consisting of Boisduvalia and Epilobium only, is advanced in many of its characteristics but occupies a rather isolated position within the family (Raven, 1976). Within the group of ten genera recognized as Onagreae (Raven, 1964, 1969), Clarkia and Heterogaura constitute one distinctive group; Oenothera and Stenosiphon another; Calylophus and Gaura a third; and Ca- missonia, Gongylocarpus, Gayophytum, and Xy- lonagra a fourth (Tobe & Raven, 1985). Hauya (Hauyeae) has been separated as a distinct tribe by virtue of its generalized characteristics in- cluding the presence of stipules, generalized leaf anatomy distinct from that of all Onagreae (Keating, 1982), and lack of interxylary phloem. On the other hand, its possession of obviously divided sporogenous tissue generally similar to — FIGURE 1. stages of development, showing that both the inner (1.1. ) and outer (o.i.) integuments are of dermal o Ludwigia arcuata. — A-D. Median longisections of ovule primordia in progressively ug n n. a, b, c, dermal initials of the i.i.; d, e, dermal initials of the o.i. — E. 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Q20H (A9q za [euliop [4 [4 (OW) L961 “dupuç “oD uuv :eruisojreO "v'S( -Ieg) 11405704 `dsqns jew wnjoy1s wuniqojidq = [euliop [4 [4 (OW) ZI¿6 uoldhumpiO “oO o[oA :erusojt[eO "vs `dleAA CUNN) v112q013 ^g uueul = Teuuop [4 [4 ‘OD S0rF uosmm P asaayD *SqqnN MYO “Tey (uoAeq % "zm n;pmqns vippanpsiog eearqord AUL (OW) = ewp z T LL6I ‘92111109 “OD əuwnjon :erulojyie) `V `S`n ƏIIIAoO (LOL) Dipunsajay DANDZO4LIJI H (OW) 6991 = Iguup z z ppbzəAO P Du24021140JA. *OeoeIn;) ONEN MYD p]jəuə1 `dsqns SIMITȚ (Ae) DJ]2uəa] `J = [eulisp [4 [4 (OW) 09927 uv4ojy “eturojtIeO efeg "oorxojJq "IQW 79 `SIƏN (SureIQV) DJDO1J2P DDJAU]-) [euuopqns (TAO) Fr pue [guuəp v-t T 8£0I UDAIING “OD ute[O3NW WOLNO VSA "QuAoH (NNN) snyofiuy uoudisouaIs Teuuopqns (OW) 6I SF 2401 ++ pue [guuəp s [4 P urn?) 42u3044 “oO SMOT 1S :unossryw "v'S'( psojjia "dsqns -qunu L psojjt4 ‘O Iguuəpqns ++ pue ewp S-Z z (OW) IZ£F uowojosg 3 42u32,44 *oSuein(q ‘OILIN pap]/ 'dsqns Walley ('S[9N `V) vanif 242110u2() Teuuopqns (OW) 225+ 2901 + pue [euuop £ [4 P II 4203044 “oO uosisgef MOSSIN VSN qoeds n40jfi8uo] `D [euuopqns + pue Iguuəp LaK [4 (OW) 68ç£ i204 “OD soded :sexə[ `v Sin YSINd D2u12202 VANDO Teuopqns '('aNVD T pue Iguuəp t-c T ££ççI Syoodg “OD ueuususS :sesuey "v'S'() UALY (NNN) 57DIRA498. `D rewpqns (OW) 27259 WoAey + pue [euuap t [4 sw o» wwugar “OD upoourT :epeA9N "V'S'(1 (&e10) "y 79 "110 L) snijofipmpuvap] snydojAjoD juouinzoju yuəum3əluIr J9jn() 1ouu UONVULIOJU] uonoəlloO pue Aeon uoxe J2NO Jo 19jnQ JO BUO uonguro,] ashes ur spenru] quid A jo so»4n "EAUX Jo uonedriorneg Jo 991821 "penunguo) “| sigV TOBE & RAVEN— INTEGUMENTS IN ONAGRACEAE 457 1985] that found in Calylophus, Clarkia, Gaura, and 1979), as have its vespertine flowers, long floral tubes, white petals, and strong floral odor, this syndrome of characteristics closely resembling that found in white-flowered species of Oeno- thera. The histogenesis of integuments which is the subject of this paper is used here as another ap- proach to clarify the relationships between the genera of Onagraceae. Although such character- istics have traditionally been included within the scope of investigations of embryologists, they have largely been confined to considerations of the number and thickness of the integuments, the development of integuments into seed coats, and the participation of integuments in forma- tion of the micropyle. In contrast, little attention has been devoted to the developmental mode o the outer and inner integuments (see reviews in Bouman, 1971, 1974, 1984). Twenty-five years ago, the studies of Roth (1957) on Capsella began to interest embryologists again in the develop- mental mode of integuments, a field that had Mi largely neglected since the studies of Warm- ng (1878) over a century ago. The subsequent n of Bouman and his collaborators on Ju- glans and Pterocarya (Boesewinkel & Bouman, 1967), on Polycarpicae (Bouman, 1971, 1977, 1978; Boer & Bouman, 1972, 1974), and on Ru- taceae (Boesewinkel, 1977, 1978; Boesewinkel & Bouman, 1978) have increasingly called our at- tention to the histogenetic characters of integu- ments as a probable indicator of affinity among taxa. Bouman (1971) emphasized the impor- tance of observations of the histogenesis of in- teguments in their evaluation, citing the fact that seemingly identical structure of integuments often developed as a result of different histogenetic processes. He also warned against the taxonomic value of data based on the structure of the mature seed coat for similar reasons. For example, Dri- B 1974), whereas Magnolia ae (Sieb. & Zucc.) Maxim., M. virginiana L., and Liriodendron tu- lipifera L. (Magnoliaceae) have an outer integ- ument of subdermal origin (Boer & Bouman, 1972; Bouman, 1977). This difference in the his- togenetic origin of the outer integument has been used as additional evidence for separating Win- teraceae from Magnoliaceae, with which they were combined earlier (Boer & Bouman, 1974). We have made observations of the histogen- esis of the integuments as part of a study of the embryology of Onagraceae. In this paper we shall deal only with the mode of initiation and sub- sequent growth of oe 3nieiueenis not consid- ering the more divers of the seed coat or testa. We shall utilize the mode of ini- tiation and subsequent growth of integuments solely as an indicator of affinity between taxa and shall attempt neither morphological interpreta- tions of these structures nor detailed studies of other aspects of embryology in these plants. Only one earlier paper deals with the histo- genesis of integuments of Onagraceae (Geerts, 1908). Geerts provides excellent illustrations of the young ovules of Oenothera glazioviana Mich. (= “O. lamarckiana”), stating “Das innere In- tegument entwickelt sich nur aus Dermatogen- zellen, zur Bildung des äusseren Integumentes finden im Periblem Teilungen statt.” Several other papers dealing with the embryology of On- alpina subsp. pacifica = a holm, 1914, Lopezia racemosa subsp. racemo- sa = “L. coronata”) have illustrations of sections of young ovules including views of developing integuments; all of these papers, however, lack an explanation of the histogenesis of the struc- tures illustrated and are not detailed enough to be used in connection with this present compar- ative study. MATERIALS AND METHODS Forty species representing all seven tribes and all 17 genera of Onagraceae were included in our study. They are listed in Table 1, along with the results obtained from each. All the samples were fixed in FAA (90 parts 50 or 70% ethanol, 5 parts acetic acid and 5 parts formalin) and then de- hydrated through tertiary-butyl alcohol series and embedded in paraplast. Serial microtome sec- tions of ovaries 5-7 um thick were stained with Heidenhain's hematoxylin, safranin, and fast green FCF. In all Onagraceae, the ovule has two integu- ments, inner (i.i.) and outer (0.i.); it becomes anatropous or hemitropous (Lopezia) at matu- rity. The terminology, abbreviation of terms, and the manner of drawing microtome sections ba- sically follow Bouman (1974). In every diagram illustrating the histogenesis of integuments, Fig- ures 1, 2, 4-8 for example, bold lines are used 458 the o.i.— udwigia arcuata. Note tha is initiated subdermally. Scales — 20 um to represent the boundary between dermal and subdermal tissues. OBSERVATIONS TRIBE JUSSIAEEAE Ludwigia arcuata, L. bonariensis, L. latifolia, L. leptocarpa, L. linearis, L. peploides, L. peru- viana, and L. virgata—In every species the ini- tiation of the i.i. commences by periclinal divi- sions of dermal cells located at the flank of an ovule primordium (Fig. 1A, see cells b). In me- Fig. 3A), or occasionally only two, a and b, were ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 _ Ra! FicuRE 3. Median longisections of the ovule primordia, showing two contrasting developmental modes of t the 0.1. is initiated dermally.—B. Lopezia racemosa. Note that the o.i. recognized. The first periclinal divisions of these three initials form a swelling of the i.i., first dis- cernible externally. The subsequent ertt ofthe 1.1. primordium takes place chiefly by divisions of daughter cells of two, a and b, initials (Fig. IC, D), so that the i.i. grows as a two-layered thin organ. In a developed ovule the i.i. usually becomes three-layered because of periclinal di- gps, of cells of the inner layer of the i.i. (Fig. E, F). The tip of the i.i. ultimately becomes ies forming an endostome similar to rms growth of the i.i. takes place by divisions of der- mal initials and their derivatives, the i.i. of Lud- E4. Lopezia semeiandra.—A-C. Median Pauqar of ovule primordia i In progressively successive igi tt f subdermal origin. — D. Median uc n development, showing that the i.i. is of derm longisections of a growing ovule. — E. Longisection of a lateral part of a developed ov the o.1 section of a ule. — F. Longise micropylar part of a MEE aa ovule. Note that the derivatives of the subdermal initials reach up to the tip of the o.i. Bracket scales = 100 uv TOBE & RAVEN — INTEGUMENTS IN ONAGRACEAE 459 1985] 460 Fuchsia radicans. Note that the o.i. is of FIGURE 5. subdermal origin and the derivatives of the subdermal initials reach up to the tip of the o.i. Bracket scale — 0 um. wigia is of dermal origin, a term used consistently by Bouman and his collaborators to express this pattern of development. The o.i. is also of dermal origin in all species examined. At the initiation stage, two dermal initials (d, e in Fig. 1B; see also Fig. 3A) undergo periclinal divisions. Subsequent growth takes place by repeated divisions of derivatives of the dermal initials (Fig. 1C, D). The mature o.i. is two-layered in all the species except L. peploides (Fig. 1E, F), and only its tip becomes multilay- ered to form an exostome. The o.i. hardly de- velops at the funicular side, and, if it does, it is made up on this side of derivatives of dermal initials only (Fig. 1E). In the sole exceptional species, L. peploides, a developed o.i. is two-layered in the upper part and three-layered in the lower part (Fig. 2E). This three-layered part of the o.i. is formed in a dif- ferent mode from that of the i.i. The first peri- clinal division of the dermal initial e gives rise to two daughter cells, the lower cell e' functioning as a distinct initial of the third (middle) layer in ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 the subsequent growth (Fig. 2A, B). Consequent- ly, the upper part of the o.i. grows by divisions of the upper daughter cell e as well as of another initial d, and the lower part by divisions of de- rivatives of an additional initial f as well as of e' and d (Fig. 2C, D). TRIBE LOPEZIEAE Lopezia langmaniae, L. racemosa and L. se- meiandra — The i.i. is of dermal origin and two- layered in all species examined (Fig. 4A—-F). The mode of initiation and the subsequent growth of the o.i. is consistent in all species ex- amined, differing completely from the pattern seen in Ludwigia. On the antiraphial and lateral sides, the initiation of the o.i. commences by periclinal divisions of subdermal cells, and not by those of dermal cells (Fig. 4A). Repeated peri- clinal divisions of subdermal cells give rise to a primordium of the o.i. discernible externally as a swelling (Fig. 4B, C; see also Fig. 3B). Subse- quent growth proceeds by divisions of both sub- dermal and dermal cells (Fig. 4D). Thus, even if dermal cells will have divided periclinally to make an exostome at the tip of the o.i., derivatives of the subdermal initials reach the tip on all anti- raphial and lateral sides (Fig. 4F). The mature 0.1. is usually three-layered in the upper part and three- to four-layered in the lower part (Fig. 4E, F). In Lopezia, the mature ovule is hemianatro- pous, and therefore the o.i. is well developed on the funicular side also (Fig. 4D). Thus, since subdermal initials and their deriv- atives contribute to the formation of the integ- ument, the o.i. of Lopezia is of subdermal origin, a term used to express a mode contrasting with that in Ludwigia. TRIBE FUCHSIEAE Fuchsia jimenezii, F. microphylla subsp. quer- cetorum, and F. radicans— The i.i. is of dermal origin and two-layered in all species examined. The o.i. is of subdermal origin in all species examined. Derivatives of subdermal initials par- ticipate in the formation of a major part of the o.1., and reach its apex as in Lopezia (Fig. 5). The thickness of the o.i. differs from species to species: three- to four-layered in F. jimenezii, and three- to six-layered in F. radicans (Fig. 5) and F. microphylla. 1985] FicuRE6. | Hauya elegans subsp. elegans. — A. Median | TOBE & RAVEN—INTEGUMENTS IN ONAGRACEAE 4: £ 1 — B. Longisection of a developed ovule. Note that the o.i. is of both dermal and subdermal origin, and the derivatives of the dermal initials divide more actively than those of the subdermal ones in the formation of the o.i. Bracket scales = 100 um. TRIBE CIRCAEEAE Circaea alpina subsp. pacifica and C. corda- ta— The i.i. is of dermal origin and two-layered in both species. The o.i. is of subdermal origin and three- to five-layered in both species. Derivatives of sub- dermal initials reach the tip of the o.i. as in Lo- pezia and Fuchsia (Fig. 9A). TRIBE HAUYEAE Hauya elegans subsp. elegans and H. hey- deana — The i.i. is of dermal origin and two-lay- ered in both species (Fig. 6A, B). The o.i. is of both dermal and subdermal origin in both species. On the antiraphial and lateral surfaces, the initiation of the o.i. commences with periclinal divisions of dermal as well as of sub- dermal initials (Fig. 6A). Subsequently, however, derivatives of the dermal initials divide more actively than those of the subdermal initials, so that the derivatives of the subdermal initials re- main in the basal part of the o.i. on the antira- phial and lateral sides (Fig. 6B). Consequently, the mature o.i. is mostly two-layered, but three- to five-layered in its basal portion in both species (Fig. 6B). TRIBE ONAGREAE Gongylocarpus fruticulosus and G. rubricau- lis— Both the i.i. and the o.i. are of dermal origin and two-layered in both species. Gayophytum heterozygum and G. humile— Both the i.i. and the o.i. are of dermal origin and two-layered in both species. Xylonagra arborea — Both the i.i. and the o.i. are of dermal origin and two-layered. Camissonia californica and C. ovata— Both the ii. and the o.i. are of dermal origin and two- layered in both species. Calylophus lavandulifolius and C. serrulatus— The i.i. is of dermal origin and two-layered in both species. The mode of the initiation and the subsequent growth of the o.i. is similar to that in Hauya. In both species the o.i. is initiated by periclinal divisions of dermal as well as of sub- dermal initials (Fig. 7A). Derivatives of the der- mal initials divide more often than those of the subdermal initials subsequently (Fig. 7B). Ulti- mately, the derivatives of subdermal initials par- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 __] L... Ficure 7. Calylophus lavandulifolius.— A, B. Median longisections of ovule primordia in progressively ive stages of develop , showing that the o.i. is of both dermal and subdermal origin. — C. Longisection of a developed ovule. Note that the derivatives of the dermal initials divide more often than those of the subdermal ones in the formation of the o.i. Bracket scales = 100 um. ticipate only in the formation of the basal part of the o.i. (Fig. 7C). The mature o.i. is mostly two-layered but three- to four-layered in its bas- al portion (Fig. 7C). aura coccinea and G. longiflora— The i.i. is of dermal origin and two-layered, and the o.i. of The degree of participation of derivatives of sub- dermal initials in the formation of the o.i. closely resembles that seen in Hauya and Calylophus. Therefore, the mature o.i. is mostly two-layered but three- to four-layered in its basal portion, which is made up of derivatives of dermal and subdermal initials. Oenothera flava subsp. flava and O. villosa subsp. villosa— The 1.1. is of dermal origin, as Geerts (1908) observed, and two-layered in both species. On the antiraphial and lateral sides, the o.i. is of both dermal and subdermal origin in both species. Geerts (1908) mentioned that in O. glazioviana (= “O. lamarckiana”), the o.i. is pro- uced by divisions of subdermal cells. But his figure (Geerts, 1908, table 19, fig. 3) illustrating a median longisection of the very young ovule primordium shows the o.i. swelling by divisions of derivatives of both dermal and subdermal ini- tials. In both species examined in this paper, the dermal and the subdermal initials appear to di- vide almost simultaneously (Fig. 8A). In any event, at least it is obvious that the derivatives of both the dermal and subdermal initials con- tribute to the formation of the o.i. Of more im- portance to the final outcome is the fact that the derivatives of the dermal initials divide less ac- tively than those of the subdermal ones, a feature in which Oenothera differs from Hauya, Caly- lophus, and Gaura (Fig. 8B). The mature o.i. is two-layered in the upper portion and three-lay- ered in the lower portion (Fig. 8C). Stenosiphon linifolius— The i.i. is of dermal origin and two-layered. The mode of initiation and the subsequent growth of the o.i. is almost identical to that in Oenothera. On the antiraphial and lateral sides, the o.i. is of both dermal and 1985] TOBE & RAVEN — INTEGUMENTS IN ONAGRACEAE 463 Y AN FiGURE 8. Oenothera flava subsp. flava. — A-C. Median longisections of ovule primordia in progressively successive stages of development, showing that the o.i. is of both dermal and subdermal origin. Note that the derivatives of the dermal initials divide less actively than those of the subdermal ones in the formation of the o.i. Bracket scale = 100 um. subdermal origin with the derivatives of the der- Clarkia delicata and C. tenella— Both the 1.1. mal initials dividing less often than those of the and the o.i. are of dermal origin and two-layered subdermal initials subsequently. In some ovules, in both species (Fig. 9C). the derivatives of the subdermal initials extend Heterogaura heterandra — Both the 1.1. and the close to the tip of the o.i. (Fig. 9B). o.i. are of dermal origin and two-layered. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 464 1985] TRIBE EPILOBIEAE Boisduvalia glabella and B. subulata — Both the 1.1. and the o.i. are of dermal origin and two- layered in both species. Epilobium canum and E. ciliatum— Both the i.i. and the o.i. are of dermal origin and two- layered in both species (Fig. 9D). DISCUSSION In Table 1 the results of this study on the his- togenesis of integuments are presented compar- atively. Although all Onagraceae have a two-lay- ered i.i. of dermal origin, four different types of o.i. are recognized with respect to mode of ini- tiation and subsequent growth. These are found y, (Fuchsieae), and Cir the o.i. is of subdermal origin, and the derivatives of the subdermal initials reach the tip of the o.i. (Fig. 10A); 2) Oenothera and Stenosiphon (On- agreae), in which the o.i. is of both dermal and subdermal origin, and the derivatives of the sub- dermal T extend beyond the middle of the 0B); 3) Hauya (Hauyeae) and Caly- is both dermal and subderma f o derivatives of the to the basal part of the o.i. (Fig. 10C), and 4) Ludwigia (Jussiaeeae), Boisduvalia and Epilo- bium (Epilobieae), and Camissonia, Gayophy- tum, Gongylocarpus, and Xylonagra (Onagreae), in which the o.i. is of dermal origin and mostly two-layered (Fig. 10D). l rms, the i.i. is almost always of ceae have a generalized i.i. with respect to his- togenesis and thickness. The amplification in thickness of the i.i. because of periclinal cell di- visions, which often is characteristic of Ludwig- TOBE & RAVEN —INTEGUMENTS IN ONAGRACEAE 465 „is of is of subdermal origin. — B. The o.i a ermal origin with the derivatives of the subdermal initials dividing more actively than those of s dermal ger pea The o.i. of both dermal and subdermal origin with the derivatives of the der- mal initials dividing more often than those of the sub- dermal o — D. The o.i. is of dermal origin ia, would in that case be sea as a sec- ondary characteristic (Bouman, pers. comm.). Bouman (1974, 1984) has me that the most primitive type of angiospermous ovule has — FiGURE 9. A. Circaea cordata. Median longisection of a growing ovule, bcp that the o.i. is of subderm the origin and the derivatives of the subdermal initials reach up to the tip al o.1. — B. Stenosiphon linifolius. Longisection of apical part of integuments. Note that, since the o.i. is of pee dermal and subdermal origin, a certain part of the o.i. is made u of a growing ovule showing two-layered i.i. and o.i. o subdermal initials. Scales = 20 p p of derivatives of both the dermal and subdermal initials. In this ovule, the derivatives of the subdermal initials extend close to the tip of the 0.1.—C. C larkia ten dermal origin. — a basal part of a developed Vind SURE two-layered i.i. and o.i. of dermal origin. . Longisecti sub, derivatives of the 466 a subdermally initiated o.i. and that thick o.1.'s are more primitive than thinner ones. In light of Bouman's principles, the histogenesis of the o.1.'s of Lopezia, Fuchsia, and Circaea would be con- sidered the most primitive found in Onagraceae, because in these genera the o.i. is entirely of sub- dermal origin. At least some species of all three genera have the original basic chromosome num- ber for the family, n = 11, and generalized chro- mosome structure (Kurabayashi et al., 1962; Ra- ven, 1979). As indicated by Eyde and Morgan (1973), Lopezia, Fuchsia, and Circaea have no evident direct relationship to one another. For example, although both have reduced numbers of flower parts, Lopezia differs from Circaea in its anther disposition and nectary morphology. Fuchsia, despite its mostly red, bird-pollinated flowers and unique fleshy fruits, may be the modern genus closest to the ancestors of most Onagraceae. The pattern of histogenesis of its integuments is in accordance with such a con- clusion. The tribe Onagreae, with ten genera, exhibits three patterns with respect to the histogenetic characters of integuments (see Table 1). Based on Bouman's (1974, 1984) postulates, Oenothera and Stenosiphon are more primitive than other members of the tribe in that their o.i. retains a stronger tendency to entirely subdermal initia- tion (see Fig. 10B). Calylophus and Gaura appear to be more specialized than Oenothera and Stenosiphon, and less specialized than other On- apice] in _ Mem o.i. still retains a weak ten- i ermal i enc sub 10C). The remaining six genera, Camissonia, Gayophytum, Gongylocarpus, earn Clark- ia, and Heterogaura, in whic e o.1. has no tendency to a subdermal . (see Fig. 10D), constitute a third group, which on the basis of this feature would be considered the most ad- vanced in the tribe. It is almost certain that the patterns of integ- umental histogenesis found in these three groups were derived from one in which the o.1. was en- tirely of subdermal origin, like that which per- sists in Fuchsia, Lopezia, and Circaea. It is not, however, possible to determine whether they were derived sequentially or independently on the ba- sis of this characteristic alone. Gongylocarpus is the only genus of Onagreae that retains the orig- inal basic chromosome number for the family, n = 11, but it has a highly specialized pattern of ovary development (Carlquist & Raven, 1966). If it is an offshoot from the line leading to Ca- missonia, Gayophytum, and Xylonagra (Raven, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 1964, 1969), then, since all of these genera have identical specialized patterns of integumentary histogenesis in agreement with their distinctive stigma morphology, it would be implied that there is no direct relationship between this line and other Onagreae. Certainly, these relationships make it appear that the reduced basic chromo- some number, n = 7, must have been derived o ' separately in this line and in the other parts of he trib the tribe The large genus Oenothera (about 125 species) is identical to the monotypic Stenosiphon in the pattern of histogenesis of its integuments, as it is in stigma morphology and other features. Thus relationship between these genera and others within the tribe remains obscure. The basic chro- mosome number, n = 7, in these genera certainly seems to have evolved independently from the same number in other genera of Onagreae. Clarkia and Heterogaura share the specialized feature of the o.i. with the first group discussed above. If they are directly related to Calylophus and Gaura, as claimed by Raven (1964, 1969) on the basis of the thick partitions that divide the sporogenous tissue of their anthers into pack- ets, it would have to be assumed that the pattern of development of their o.i.'s was independently derived from that in the "Gongylocarpus— group"; if not, then their specialized anthers and stigmas must have evolved within that group. Hauya (x = 10), now placed in a separate tribe Hauyeae (Raven, 1979) on the basis of the many features, all apparently retained primitive ones, in which it differs from all genera currently as- signed to Onagreae, was formerly grouped with Calylophus, Clarkia, Gaura, and Heterogaura on the basis of its divided sporogenous tissue. The histogenesis of its integuments is identical to that in Calylophus and Gaura, differing sharply from the more advanced condition found in Clarkia and Heterogaura and the more primitive one in Oenothera and Stenosiphon. This may indicate that Calylophus and Gaura (both x = 7) were derived from a phylogenetic line separate from that leading to other Onagreae, and that Hauya may be a surviving early evolutionary offshoot of that line. The relationships of the different genera of Onagreae and their affinity to Hauya will be discussed in more detail in a separate paper focusing on the exact nature of the divided sporogenous tissue in the Onagraceae (Tobe & Raven, 1985). The remaining two tribes of the family, Jus- 1985] siaeeae (Ludwigia only) and Epilobieae (Epilo- bium and Boisduvalia), share with six genera of Onagreae the most advanced type of integumen- tary histogenesis in the family, with a two-lay- ered o.i. of dermal origin. Fundamental floral differences discussed by Eyde (1977, 1978, 1981) make it clear that Ludwigia represents a phylo- genetic line different from all other Onagraceae, so that it must have evolved its specialized 0.1. independently. There appears to be no evidence to suggest a close grouping of Epilobieae with any genera of Onagreae, so that we may assume that a two-layered o.i. of dermal origin evolved independently in it, in the Gongylocarpus-line, and perhaps also in Clarkia-Heterogaura —thus three or even four times within Onagraceae. A similar condition has evolved independently in various angiosperms, as for example, Lactori- daceae among “‘Polycarpicae”’ ud 1971). Within the family, Ludwigia peploides (sect. Oligospermum) is distinct from all other species in having a three-layered o.i. instead of a two- layered one. Eyde (1977), pointing out that sect. Oligospermum stands apart from other ludwigias because of its characteristic aerenchymatous tis- sue surrounding the endocarp, inferred that this section seems to have diverged early from the other ludwigias and developed its own aquatic adaptations. The characteristic developmental mode of its integuments, unique in Onagraceae, constitutes additional evidence supporting his ypothesis. his comparative study covering the whole family enables us to evaluate the histogenetic characters of integuments. As has been described and seen in Table 1, this character is nearly ge- nus-specific. Even in cases in which the seed coat or testa histology differs from species to species (e.g., in Gongylocarpus, Carlquist & Raven, 1966), the mode of initiation and the subsequent growth of the integuments is consistent within a genus. In view ofthis, detailed comparisons ofthis char- acter have made it possible to evaluate the affin- ities ofgenera and to consider their relative states of advancement. LITERATURE CITED Boer, R. DE & F. Bouman. 1972. Integumentary studies in the Polycarpicae. II. Magnolia stellata and Magnolia virginiana. Acta Bot. Neerl. 21: 617- 629. & ————. 1974. Integumentary studies in the Polycarpicae. III. "reca winteri (Winteraceae). Acta Bot. Neerl. 23: = F. = l 2 Development of ovule and esta in Rutaceae. I. Ruta, on and reium pen B Neerl. 26: 193- TOBE & RAVEN—INTEGUMENTS IN ONAGRACEAE 467 1978. Development of ovule and testa in Ruta taceae. II. Some representatives of the Auran- tioideae. Acta eh Neerl. 27: 342-354. & F. Boum 1967. Integument initiation in gm and predi Acta Bot. Neerl. 16: —— q 1978. Development of ovule and testa in Rutaceae. II. The unitegmic and pachy- chalazal seed of Glycosmis cf. arborea (Roxb.) D.C. Acta Bot. Neerl. 27: 69-78. Bon, J. & F. BouMAN. 1974. Development P ovule and integuments in Euphorbia milii an o carpicae. I. Lactoridaceae. Acta Bot. Neerl. 20: 565-569. 74. V. “pies studies of the ovule, integuments and ps =a e angiosperms. The- sis. Univ. of Amste , Ne ; riage E studies in the I IV. Liriodendron tulipifera L. Acta Bot or 26: 213-223 978. Integumentary studies in the Polycar- picae. V. Nigella damascena L. Acta Bot. Neerl. 27: 2. ror The ovule. Pp. 123-157 in B. M. Johri (edi tor), Embryology of Angiosperms. Springer- Verlag, Heidelberg CARLQUIST, S. 1975. Wood anatomy of Onagraceae, with notes on alternative modes of "pig ime movement in dicotyledon woods. Ann. Misso Bot. Gard. 62: 386-424. 1977. Wood anatomy of Onagraceae: addi- tional species and concepts. Ann. Missouri Bot. Gard. 64: 627-637. . 1982 [1983]. Wood anatomy of Onagraceae: further species; root anatomy; significance of ves- tured pits and allied structures in S EO a Ann. Missouri Bot. Gard. 69: 755-7 P. H. Raven. 1966. The systematics and anatomy of Gongylocarpus (Onagraceae). Amer. J. Bot. 53: 378-390. EvpE, R. H. 1977 [1978]. and evolution in Ludwigia (Onagrac droecium, placentation, merism. A Bot. Gard. 64: 644-655. sap Spi structures ae). I. An- nn. Missouri oem structures and evolution a (Onagrae ae). III. Vasculature, nec- TI Missouri Bot. Gard. 68: in Ludwi taries, Mice Evolution pu systematics of n. Missouri Bot 1982 [1983]. Gard. 69: 735-747 & J. T. MORGAN. 1973. Floral structure I evolution in Lopezieae (Onagraceae). Amer. J. B 0: 771-787. GEERTS, J. M. 1908. Beitrage zur Kenntnis der cy- tologischen Entwicklung von Oenothera La- marckiana. Ber. Deutsch. Bot. Ges. 26: 608-614. JOHANSEN, D. A. 1934. Studies on the morphology of the Onagraceae. VIII. Circaea pacifica. Amer. J. Bot. 21: 508-510 KEATING, R. C. 1982 [1983]. The evolution and sys- 468 tematics of Onagraceae: leaf anatomy. Ann. Mis- —-803. S & P. H. RAVE comparative study of mitosis in the eee Amer. J. Bot. 49: 1003-1026. O'NEAL, C. F. 1923. A study of the ste sac de- phenomena in Oe- nothera irre Bull. Torrey Bot. Club 50: 133- 146. PLITMANN, U., P. H. RAVEN & D. E. BREEDLOVE. 1973. The systematics of Lopezieae (Onagraceae). Ann. Missouri Bot. Gard. 62: 621-646. RAVEN, P. H. 1963. The Old World species of Lud- Mio (including Jussiaea), with a synopsis of the yore de ceae). Reinwa rdtia 6: 327-427. "The generic subdivision of Onagraceae, nu Onagreae Britania 16: 276-288. A revision of the genus Camissonia (Onagraceae) Contr. U.S. Natl. Herb. 37: 161- 1976. Generic and sectional delimitation in Onagraceae, tribe Epilobieae. Ann. Missouri Bot. Gard. z 326-340. A survey of reproductive biology in E New Zealand J. Bot. 17: 575-593. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 . GREGORY. 1972. Observations of meiotic I in Gaura (Onagraceae). Brit- tonia 24: 71-86. 1979 [1980]. Sisi of à oe). Ann ard. . Die Histogenese der Integumente von Capsella bursa-pastoris und ihre morphologische Bedeutung. Flora 145: 212- PR: RAGLOWSKI. 1975. The evolution of pollen tetrads in Onagraceae. Amer. J. Bot. 62: 6- “= W. F SOE & M. SHARP. 1978. An ultrastructural m of viscin threads in On- agraceae pollen. Pollen & Spores 20: 5-143 TACKHOLM, G. . Zur Kenntnis der entwicklung von Lopezia coronata Andr. waseq Bot. Tidskr. ^ 223-234. r L11101 Tose, H. & P. H. Raven. 1985. The evolution of ae anthers in Onagraceae. Amer. J. Bot. (in pre WARMING, M . 1878. De l'ovule. Ann. Sci. Nat. Bot., Ser. 6, 5: 177—266. CHROMOSOME NUMBERS IN COMPOSITAE, XV: LIABEAE HAROLD RoBInson,! A. MICHAEL PowELL,? ROBERT M. KING,’ AND JAMES F. WEEDIN?? ABSTRACT Lim ted I lad £ AL bers in the small, mostly Andean tribe Liabeae acne records of 12 of the 16 genera: Cacosmia (x = 7); Chionopappus (n = ca. 9); Ferreyranthus, Oligactis, and Liabum (x = 18); S 12?); Erato (x = 9); Philoglossa (n= — (x = 16); Paranephelius (x 18); Chrysactinium (x= 12); Munnozia (n = 10, = 9, 14); een aad SR = The tribal base number is apparen i x= tern Hemisphere. Apparent aneuploid series of the type seen in he highest > Mi osito known elements i). the tribe with extensive number. No correlation is seen between chro- nalyunini ua adori ser almost war qaa generic gine aa in the tribe, and no evidence is noted of hybridization ossa extant gener. The present paper continues a series dealing with chromosome numbers of Compositae (Ra- ven et al., et al., 1963, 1967; Payne et al., 1964; Solbrig et al., 1964, 1969, 1972; Anderson et al., 1974; Powell et al., 1974, 1975; Kingetal., 1976; Tomb et al., 1978; Robinson et al., 1981) and is the first dealing with the elements of Liabeae as a united group recognized at tribal level. Because data for the tribe are limited, reports from the literature are included in the table with altered identifications where necessary. New reports are provided for 31 populations of 15 species in- cluding new reports for nine species and one ge- nus (marked respectively in Table 1 The new reports in this paper are based on material collected by R. M. King and counted by A. M. Powell and J. F. Weedin. The chro- mosome counts have been made from aceto-car- mine or aceto-orceine squashes of microsporo- cytes in meiosis. Voucher specimens of the King collections are in US, a second set is in MO. Robinson (1983a) recognized about 157 species of Liabeae in 15 genera; his paper is the basis for comparison in this paper. The subsequently de- scribed monotypic Bishopanthus of Peru (Rob- inson, 1983b) is unknown cytologically and will not be considered in the present study. In spite of the comparatively small size of the tribe, Lia- beae show considerable diversity in many struc- tural details (Robinson & Brettell, 1974) and pol- 1960; Raven & Kyhos, 1961; Ornduff len (Skvarla et al., 1977) which are used as the basis for three subtribes, Liabinae, Paranephe- liinae, and Munnoziinae, in the recent revision considerable extent with the revised generic con- cepts and show distinctive trends in different subtribes. Previous records of chromosome numbers of Liabeae are scattered and often confusing. Orn- duff et al. (1963) offer the only cytological eval- uation of Liabeae as a group, but only those parts treated under the traditional concept as subtribe Liabinae in Senecioneae. The traditional dispo- sition, derived from Bentham (1873) and Hoff- mann (1894), was totally flawed by the inclusion of foreign elements such as Neurolaena and Schistocarpha, which are now placed in Helian- theae, by the exclusion of true members of Lia- beae such as Chionopappus, Cacosmia, and Phil- oglossa, and by the placement of all remaining members of the tribe in a single genus, Liabum. Ornduff et al. (1963) were able to mention only one chromosome count of a true member of the tribe, a Liabum sp. (L. ovatum vel aff.) reported by Diers (1961) as n= 14. The chromosome counts available in the unnatural group were said by Ornduffet al. to “attest to the isolated position of the Liabinae." A number of additional chromosome counts ! Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 ? Department of Biology, Sul Ross State University, Alpine, Texas 79830. 011. ? Community College of Aurora, Aurora, Colorado 80 ANN. Missouni Bor. GARD. 72: 469—479. 1985. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 470 6£I6 doysig y Suty *eo1eurefe;) Jo FN Wy ss orwel ‘neg (ajnvop `J se 'zZ861 “Sur L X Loa) Nog £176 doysig P Sury ‘OZU jo S WJ £z :seuozeury ad SZO doysig X 3ury *equieqouiroT JO MS WY op :seuozeury ‘nog 0026 doysig P Sury ‘seXodeyoeyD Jo JS peol Zuoye :seuozeury Md 8716 doysig 3 suty ‘eoreweled jo IN WJ [r :eo1eurefe;) `nioq £964 vpaw]y y 3ury 'seqourq;) Jo N wy z :efo'T Jopenog (z861 '1oum 79 uona) wd (w4nqvrT s? :(96[ “Te 19 UML) Md (1961 's191qq) Nog 6C64 PP -ətu]V 3 3ury *efo jo 4 wy LI :edrgourg;)-eoure7 -Jopenoq £169 Suly ‘elo jo d WA c eo elo] '1openoq 6884 ppatu]y P Bury ‘eloy jo M uni L :efoT 1openoq £049 Bury ‘equind jo 3S WJ 9 “ed :Aenzy -1openoq e(nZ 1 = u UuƏAƏ Á[Qqissod) 121d.131u1 0} 3[noggrp I oxeui sjug[eArq orgdaouiro1213q ‘(ABU -UWI99S) SjU9[UAIQq Joj[Teuis J9q10 “sluə[eAIq pozis-o1e1opour ç *juo[eAtq J98Ie] [] pI 10 £1 cl »«(1uopeAnpnur e SI A[QIssod ,jU9[PAIQ,, 19312] os sI[95 AJ ut uaos puq :3up[oo[-1uourge1j uəAə səuo 1ə[[gtus IWOS PLus 0} UMOP Sozis SUIAIVA pue JJ ZIR, | 33 noggrp uon ejoud -191ur 3xeui sluə[pAIq orgdaouio1919g ‘pI 10 ‘çq “ZI ÁAjqissod 3uouSeg [ + ZI = u 10 ZI = t 2x SYOO] Sj[29 əwos uD | + £T *[SII95 9UIOS UI IVAP AJOA JUSTRAIG IZIN :(;1uo[ -EAn mW JO) ju9[eAIq BILI Á19A [ “H£ I Jo jusufddeg | + ZI = u Jogo ZI =u Je ÁA[8utuuəəs s[[29 ətuos ut] çI 10 ZI «(183/9 10U) [ + €I AQT ‘SI “I UƏA9 10) ç] UdAd A[qrssod ZI “eo eb I-I cI cI (07-81 = uc) 6 “BO a(ƏZtS oures [Je sjuo[eA1Q) / qjuoaurdej [Tews [ + L q(9ZIS oues fe sjyuo[eAtq) L «(1uourdejy pews 1 + A[qtssod) / I[23324g ?? UosuIqoY 'H CuosərH) wumpgnsoa wuniuovsiau) manag ?? uosuiqos `H (M g`H) sapiomopaang wuninovsiau) ALIA `A `S !upt/juəq snddodouon/O uos -uiqow ^H Usury JeA DS08nT4 pttusoopO WaH ?»so3n4 ptuusoopƏ IMOS 2JnjeJojrT JO Á LIO (u) 1SqdumyN Əəutosouroruo soar»odg `əeəqer1 107 sooJnos pue sJaquinu a2uiosouioIq^) ‘| a18vL 471 ROBINSON ET AL.—LIABEAE CHROMOSOME NUMBERS 1985] (z861 '1oum X uma) nəd (wunqvrT se ‘p861 “Te 19 uosue() OULIEN “eIquIO[OD (w4nqvrT se ‘0861 'Ássonig ?p uosue() lopenoq 6959 3u1y *oAng JO N WY g[ `V :ezejsed -Jopenog £969 Sury ‘oon OFA JO qd uni z "eo :enqemáun |, 1openoq (C861 “Sum % uoma) wd (0161 “Borg 79 sox10 [) orqndos ueorururoq Z£68¿Z so4pun;) P spsp2ə41pnO *uiotA op BONA (LEMA -opueJge[e3ng) easy A QUN vT ouo ojuouropord '[ejuop -1990 JUMIA ENU BII[IPIOD VML) [op AVLA `#Iquio[oO SSS9 Bury 'or8əN ONJ JO 4 WA c “eo :enyem3un L -Jopenod $644 Upaw]y 3 Surly ‘TeueD jo 4 WA L9 :JeuUeD -Jopenog 1969 3ury ‘Keong Jo AN WY 6£ “Bd :oze1oquimq;) -Jopenog $489 Sury “eouənO Jo AN WA 9-6 “ed :Aenzy '1openoq 9899 3ury “UID jo AS WJ pI “ed :Aenzy "10penog IF$9 sury 'soueg Jo q Wy / :enyemsuny -1openoq (T861 '1oum X UMA) Md OIOL Sury ‘SIULA JO 3S WA ZI “ed :seAenr) -Iopenog (0861 ‘UƏS1O) LF99 Bury ‘BIS3IS JO N WJ 6 "eo :AeNzy "Jopenog (tunqorT se :L961 “Te 19 1oum [) məd (Jo wnqvrT se *6,61 “Te 19 1oum L) eIquio|oO (DIZOUUNPY se ‘0861 “uəs|O) equo (wu nqprT se ‘pg6I “Te 19 uəsuer) oAeumjng *erquioj[o;) (Pizouunyy se ‘0861 'uss[O) Jopenoq pc “BO ZI gI 95 A e (red og1ep | ue) II cl 8I *6T :0c-LI el] Stiseude1əuu Aq Sur -je1edos ÁA[snormo002ud pue (u6] = u Jt SII -Se1j puno: ç pu? „61 A[šurusəəs] 6] "eo »SjuouldeJj / *9 + 6I »SjuauldeJg Z (| + Qc "vo -— 99S) JOI UI pəA9I[9q 6 JO Woda e(S}UZTBAIG STYdIOWIOIINI9Y) | F 6I 8T Anaa Š mng 2 uos -uiqo» `H (AVID `V) vpad] nizouunjy I[2324g 79 uosuIqoYy ^H ('sseO) inaissnf pizouunjw mng 2 uosut -qow 'H (37$ `d) Pijofuspu vizouungy uosuIgoYy `H 7/14(2442f piIzouunpy 'qpAy əjnpəopqns wunqorT] SSX] (N g H) səptotuolsp]ətu wnqprT uosutqo `H usury wnqgvrT `ssə'1 wnpunquoJf uunqprT `UOJƏIH 11542832 uunqpr] `UOJƏIH 7712284n0q tunqprT mng ?? wosuIqoy `H (Tg H) Snyofisspqiaa snuun44244241 meunag X uosutiqos H (e1K21194) SNSOSNA snu1unaa4424 uosuiqo» ^H (NEA) p3tup3]na 01045] 'Od sapiomuudjod 0143 991JnoS ƏIn18191r1 JO Á VIOT (u) i9qummN JWIOSOWOIYD sar oodg 'penunuo) “| slgv L ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 472 ‘snuod 10j 110d21 1SIJL] q *SLIOddI MƏN e 8674 Suly *oSueuoj[ezanQ) jo S WY gI “ed :oguguəl[ezənO "epeurojenr) I LIL 8uty *ugjneury jo MS um zz "eo :epurosg eewo Sp Bury *o8ueuojezejJA JO AA WY oe “Bd :zənbədəluons eeuwen Gunqpr'T se ‘pL 6] “Te 19 [[JaMog) oorxoj (7861 “Sum l 7 UOTTIG) Neg (v4d4720421d vss0J80jtd Se ‘1961 's1o1q) Neg (ye [9^ UNDA uunqt17 se :[96] 's1o«q) Nag OFS L doysig É 3u1y NOYO 1e :equreqeuooO enog (runmımq winqprT se ‘1961 “Te 19 UMI) məd (tungo! se ‘p861 “Te 19 uosue() eoreureurpun;) ‘equo (w4riqprT se ‘0861 ‘Assonig 2 uosue() 1openoq 6064 vpaw]y y Suty “elo jo d wy c :efo'] `1openoq £84 vpawu]y P Sury ‘oMBesag JO S WJ 01 :efo j10penog ELLL Dpaujpy P 3ury ‘aquind jo S WJ 01-8 :Aenzy -1openog 8169 3ury “elo jo 3 Wy g “ed :efo'q '1openoq (winjnj)180s UNQUIT se *p86] “Te 19 uosue[() eozeureurpunO *erquioj[o?) (wnjpudaov32w wnqvrT se ‘6961 ‘BUA Y p[240g) equo 1894 doysig P Suty “ue -UNL 0} peor UO TUIO[O/) woy wy $7 :equreqguooO euog e(ƏZIS JUSIOYIP s}USTeA -1q ‘sodeys onsuojoereuo Sururoj jou SIS -Superp 1? sired 9p “çI = u 3q 0} sreodde -o[[eurs ç—[ *ore| | *o1yepoo[onunjnur) 91 «LT = u UƏAƏ K[qrssod 10 *9[ = u JO ‘ZIET ezixo jU9[EAIQ IUO YIM CT = U) 9] ‘CI »([&uniou -qX pue orudiouuo1ə919q sjuopeAtq :9[ = u JO jU9[EAIQ 281e[ DUO WIM cT = u) 9] JO çI 81 “¿I (1x21 ut pəuonsənb) z1 6t “BO (PERL SB oues) or «(1x21 90g "ord -1ouro1j91əq Ápy3ys siuə[eAIq Ioj[euis g Jo Əuuos jet po1ou 1dəoxə '£/// s? 2uies) ¿ 01 s(sluə[eAIq exi 3urAeqoaq 1nq sjus[peA nur A[qissod S]U9[EAIQ Ə8Ig[ BY} ‘ZIS [eULIOU aq) IM} SjUu9[EAIQ 7 pue? sjugp[eAIq pozis-[eunriou g) oT eC “89 Ol OI el + ZI ‘QAPAN (uosutiqo 7T `@) zi;vgojqns puijouig ‘PAY (»xeIg “A `S) Dsopysoddy pumpurs "ry % "400H 40JO2SIp plulp]ou1sç Mng ?? UosuIqoY "H ('uoJotH) PENON ERS Sldasouopnasg '"PP9A SNIDAO snyaydaupivg mng ?? uosut -QOY `H (uodorg) 118721 snijoudaubapq meunag ?? uosutqoq 'H CuosərH) sisuau2ur2id siop3i]o "qiueg sipiuoiauas vizouunp uos -uiqow `H (aypuy) muoa4pui pizouunpy 99JnoS 3Jnj?JojrT 10 Áji[e90] (u) 19qurnN ourosouio1q.) sateds 'Penunuo) ‘| sigv L 1985] have become available for the group since the earlier summary, some as a result of recognition of the previously reported Philoglossa and Chionopappus (Diers, 1961) as members of the tribe, others as a result of new reports. Norden- stam (1977), in his short summary of the tribe, cited chromosome numbers for seven genera. Dillon and Turner (1982) gave reports for four genera. The few reports in recent years by other au- thors, and the reports in the present study are still limited in number, but the total is now suf- ficient to see some patterns in the tribe that have not been evident before. The rather complete understanding of the tribe on the basis of other provides a basis for important phyletic conclu- sions. The reports in the present paper represent a small return on many attempts. The members of the Liabeae are much less easily counted than most members of such tribes as the Eupatorieae (King et al., 1976) or Heliantheae (Robinson et al., 1981). However, the various ways in which chromosomes are difficult to count are them- selves characters worthy of study, and are often characteristic of genera or tribes. One example in the subfamily Asteroideae (sensu Robinson, 1977) is Mikania in the normally rather easily counted Eupatorieae. Mikania seems to have some bg in the actual number of chro- osomes, but the primary difficulty lies in the slight differential in the stainability of the chro- mosomes and the cytoplasm. The chromosomes stain poorly and the cytoplasm takes enough stain to make the chromosomes difficult to count. Most of the examples of less easily counted Asteraceae, however, are in Cichorioideae (sensu Robinson, 1977) in which Liabeae is included. Collections of Mutisieae, Vernonieae, and Liabeae all seem to yield characteristically poor results. The comparison of this feature that is partic- ularly important is that between Liabeae and Vernonieae, because the former have often been included in the latter in spite of their opposite leaves, rays, yellow corollas, and frequent milky sap (Cassini, 1823, 1825, 1830; Nash, 1976; Jan- sen & Stuessy, 1980). That both tribes have chro- mosomes that are difficult to count might be tak- en as an indication of relationship, but according to the observations by Powell, the nature of the problem in the two tribes is not the same. In ROBINSON ET AL.—LIABEAE CHROMOSOME NUMBERS 473 Liabeae there are uncertainties in the interpre- tation of heteromorphic bivalents that appear to meiosis I configurations occur, one or t them appear to be much larger inia (or mul- tivalents?), and then the other bivalents occur in varying sizes down to small, with some of the smaller configurations often resembling frag- ments in size and shape. When bud material is adequate it is often possible to resolve uncertain counts by observing meiotic chromosome be- havior through several stages. In Liabeae, how- ever, aceto-carmine staining is often poor at pro- phase I stages before and through diakinesis. Metaphase I and anaphase I chromosomes usu- ally stain deeply with aceto-carmine. In Verno- nieae, aceto-carmine preparations often reveal poorly stained bivalents that remain clumped or “sticky” in meiosis I stages. According to Sterling Keeley (pers. comm.), Vernonieae yield better results when salcie t midday, a trait not tested for in Liabeae. CvTOLOGICAL EVIDENCE REGARDING ORIGINAL CHROMOSOME NUMBER AND RELATIONSHIPS In many tribes in the Asteraceae, original base numbers have been determined with reasonable confidence (Senecioneae — Ornduff et al., 1963; Eupatorieae — King et al., 1976; Heliantheae — Robinson et al., 1981). The various chromosome numbers known for Liabeae are adequate for a similar determination for that tribe. There have been various proposals of a base number for the Asteraceae, but only two are re- garded here as credible. The number x — 9 with its multiples is most common in the family (Sol- brig, 1977) and has been suggested as the base number for the family by Raven (1975). More recently, Robinson et al. (1981) have suggested x = 10 as a base number for at least the entire subfamily Asteroideae. Evidence of predomi- nant decreasing aneuploidy in Asteraceae would or x — 10 as ancestral in most Cichorioideae as well. Still, internal evidence from Liabeae does not iid agree with the external evidence. The the tribe seem e n — 9andn = 180rca. 18. The counts of n — 9 seem centrally located in the tribe and have been cited from elements of all three sub- tribes. The number n = 10 is reported, perhaps BS < (11131 m (0m 474 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Munnozia | Liabum Austroliabum Sinclairia | Paranephelius x = 12 x= 18 x= 16 | x = 9, 14 | | Chrysactinium lo igactis Microliabum Liabellum | | | Pseudonoseris x = 12 x = 18 | | x = 12? | | | | | Philoglossa | Ferreyranthus | x = 18 | xw 78 | Ë | Chionopappus | | Erato x= 9 | | Cacosmia x= 9 | | X= 7 | I | | | x=9 | Munnoziinae Liabinae Paranepheliinae FiGURE 1. Schematic representation of possible relationships of genera and subtribes of the Liabeae (from Robinson, 1983a) with postulated basic chromosome numbers inserted erroneously, in Munnozia but in a compara- tively specialized element of that genus (Rob- inson, 1983a) as explained below. A base of x — 9 would explain adequately all the elements of the Liabeae, and there is no need to assume a higher base of x — 10 within the reconstructible history of the tribe. No evidence of an ancestral x = 10 survives in extant Liabeae. The basic chromosome number in Liabeae and the pattern of variation in chromosome numbers both furnish significant contrasts from other ele- ments of the family which have been considered related. The base of x = 9 and the lack of stability in the chromosome number in the tribe contrast with the nearly consistent n = 10 of Senecioneae in which members of the Liabeae were tradi- tionally placed (Bentham, 1873; Hoffmann, 1894). Thi id dditional fo 4 _ the removal of Liabeae from the Asterioidean Senecioneae, which has been proposed by nu- merous recent workers (Robinson & Brettell, 1974; Wagenitz, 1976; Nordenstam, 1977; Nash, 1976; Jansen & Stuessy, 1980; Robinson, 1983a). Most of these recent workers place Liabeae near or in Vernonieae in the series of tribes pres- ently included in the expanded Cichorioideae (or a segregate group Vernonioideae Turner ex Jan- sen & Stuessy, 1980). It is Vernonieae to which Liabeae have closest resemblance in the general aspect of their styles, anthers, and pappus. The base number of either x = 9 or 10 suggested by Jones (1977) for Vernonieae, refined to a prob- ble x = obinson et al. (1981), is not significantly different from that in Liabeae Nevertheless, actual relationships between the two tribes may not be as direct as many authors suppose, since there are significant differences in morphological, anatomical, and palynological characters. A more careful analysis of the cytol- ogy reveals an important discrepancy between the tribes in chromosomes as well. In the Ver- nonieae, the x = 10 is characteristic of the Pa- neces pene and various specialized ele- persed from that area (Jones, 1977). aue onini the Neotropical elements, that overlap geographically with Lia- beae, h ] I fx = 19 (Funk in Turner, 1981) and x = 17 derived from poly- ploids of x = 10. On the basis of present evi- dence, n = 10 is not represented in the basically American elements of the Vernonieae. This rais- es the possibility that the Vernonieae were orig- inally restricted to the Paleotropical Region and were initially introduced into the Western Hemi- sphere in a polyploid condition. Liabeae are strictly Andean in origin, with a 1985] series of chromosome numbers that seem to be based on an original x = 9. These numerical and al di iscrepancies, correlated with mor- o crepancies (Robinson, 1983a; Robinson & Mar- ticorena, unpubl. data), suggest that the two tribes had separate origins in separate hemispheres and that their common ancestor is quite remote. REVIEW OF THE CHROMOSOME NUMBERS The following discussion of chromosome numbers in Liabeae follows the three subtribal groupings of Robinson (1983a). For purposes of orientation, it should be noted that according to Robinson, Liabinae and Munnoziinae appear to represent a basal divergence in the tribe, and Paranepheliinae seem to have arisen from slight- ly more advanced members of Liabinae. As such, all three subtribes contain elements showing comparatively primitive characters LIABINAE As shown below and in Table 1, and as sum- marized in Figure 1, genera of the subtribe fall into two groups. The first has chromosome num- bers apparently consistently at n = the second has numbers apparently consistently near n = 16 or 18. The latter group includes all of the more widely distributed and more richly speciated genera of the subtribe. A series of new reports clearly establishes x — 7 as the chromosome number for Cacosmia, and it is the only known occurrence of the number in the tribe. It is not difficult to interpret x = 7 as a reduction from x = 9 that seems basic in the tribe, although Cacosmia is a shrub, and such reduction usually involves reduction in the habit to less woody or shorter-lived plants (Bennett, 1972; Robinson et al., 1981). The primary tra- ditional distinction of Cacosmia, the lack of the pappus, along with such features as the ranked involucral bracts and the modified arrangement of thickenings in the endothecial cells (Robinson, 1983a) do not justify placement in a separate tribe such as the Helenieae (Bentham, 1873) but do indicate some phyletic distance. The genus dence from other tribes and from other Liabeae against a base of x = 7 and against frequent aneu- ploid increase (Robinson et al., 1981) prohibits the acceptance of x = 7 as the forerunner of x = ROBINSON ET AL.—LIABEAE CHROMOSOME NUMBERS 475 9 in Liabeae. Furthermore, some specialized as- pects of Cacosmia discourage any thought of a direct connection between the x = 7 of Cacosmia and the x = 12-14 of various Munnoziinae and Paranepheliinae. Chionopappus of coastal Peru has been re- ported to have n = ca. 9 by Diers (1961), and this is the only report of the supposed tribal base number in Liabinae. The genus is distinguished by its uniseriate plumose pappus and was orig- inally placed in Mutisieae (Bentham, 1873). Oth- er features of Chionopappus, however, are not unusual for Liabinae. The n= 9 in the genus indicates a point of origin in the subtribe before the incidence of polyploidy that characterizes the advanced genera, a point similar to the ones at which Cacosmia and the Paranepheliinae may have diverged. The remaining known chromosome numbers in Liabinae are apparently at the polyploid level. In Ferreyranthus of Peru and southern Ecuador, there are reports of n — 18 or 19 plus or minus 1 on the basis of two closely related species. In this genus, polyploidy is associated with the most subarborescent members ofthe tribe, but the habit difference from genera such as Cacosmia is not sufficiently marked to indicate any particular correlation with polyploidy. There is no reason to assume that polyploidy in Ferreyranthus is anything but a part of the general polyploid trend the subtribe are Bon Eom and Oligactis. The y in the An- des of Colombia and Ecuador, but Liabum has attained the widest range of any genus in the tribe, from central Mexico and the Greater An- tilles to Bolivia. The various reports indicate a chromosome number of n = 18 or 19. Because of the difficulties in counting chromosomes in the tribe, it is uncertain whether the variety in reports represents a true variety in numbers. The presence of a probable base of x = 9 in the tribe causes the present authors to favor x = 18 as basic for Liabum, Oligactis, and Ferreyranthus. The 7n — ca. 39 reported from Oligactis pichin- chensis indicates a further level of polyploidy in the group. The report of n = 9 for Liabum bourgeaui by Olsen (1980) is anomalous in view of all other evidence for the generic group. Also, the species occurs in Mexico and Central America, but not in Colombia from where the collection is cited. The voucher from SWMT is actually Verbesina two genera 476 barragana Cuatrecasas or a closely related species, which casts total doubt on the report because n = 9 is not characteristic of either genus. Un- fortunately there are no other reports for L. bour- geaui or of its two closest relatives, L. asclepi- adeum Schultz-Bipontinus of the northern Andes or L. ferreyrii H. Robinson of Peru. The Sinclairia group of Mexico and Central America has been reported as n = 17,18 by Pow- ell et al. (1974) and Nordenstam (1977) on the basis of a single count of SS. discolor. It seems notable that this report was uncertain toward a lower number from n = 18 because the present study shows n — 15,16 for the genus on the basis of a number of species with one apparently clear count of n = 16. In the absence of better evi- dence, the x = 16 is regarded here as basic for the group. Sinclairia, Ferreyranthus, and the genus-pair Liabum and Oligactis all have higher numbers of chromosomes presumably derived from poly- ploidy, but these comparatively advanced mem- bers of the subtribe may not share the same poly- ploid ancestry. Certainly, Sinclairia has structural features and latex that suggest closer relationship to Chionopappus with n — ca. 9 than to the other polyploid genera. PARANEPHELIINAE The subtribe contains Paranephelius and Pseudonoseris. As can be seen in the table, there are only three chromosome number reports for Paranephelius showing two basically different numbers. The new report of n= 14 or 15 for Paranephelius ovatus supports the report of n = 14 (2n = 28) by Diers (1961: 465), although Diers's report for P. ovatus vel aff. from coastal Peru probably represents the closely related P. uniflorus Poepp. & Endl. At the same time, the n — 9 reported by Turner et al. (1967) for P. jelskii (as P. bullatus) fits well into the concept of x — 9 as a base for the tribe and is reported from the most divergent element in the genus. The element for which this ancestral number is reported is restricted in distribution in Amazo- nas in northern Peru while the probable derived group with n = 14 is widely distributed from more coastal northern Peru south to Bolivia and north- ernmost Argentina. These counts do not provide a basis for much speculation on the actual origin of the n = 14, but some numbers in the n = 11, 12 range occur in the Munnozia element of the Munnoziinae where possible patterns in the or- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 igin of such aneuploid increases are discussed more fully. The recent report of n = 12 from Pseudono- seris szyszylowiczii (Dillon & Turner, 1982) seems somewhat anomalous and needs confirmation. As reported, the number would represent an ad- ditional aneuploid increase in the subtribe sep- arate from that cited above within Paranephe- lius. We hesitate to accept the present report because the voucher from F, collected 5 Jan. 1979, and a duplicate at US, while correctly deter- mined, show only mature heads, a condition, which according to R. M. King is typical of the species at that time of year. It seems possible that the buds counted might have come by error from some intermixed Munnozia or Chrysactinium in which the number 7 = 12 is common. MUNNOZIINAE The subtribe contains four genera that fall eas- i roups, Erato—Philoglossa and Chrysactinium-Munnozia, on the basis of pu- bescence, endothecial cell structure, and pollen structure (Robinson, 1983a; Robinson & Mar- ticorena, unpubl. data). Chromosome reports in Table 1 and the summary in Figure 1 show that the two groups are distinct cytologically also. Erato and Philoglossa have ranges that broad- ly overlap in the central Andes south to Bolivia, but Erato is concentrated in the main Andean chains that reach north to Venezuela and Colom- bia with an extension to Costa Rica, whereas four of the five species of Philoglossa are restricted to the coastal ranges of Peru. The two genera have been placed in separate tribes in the traditional systems of classification of Bentham (1873) and Hoffmann (1894), although they prove inti- mately related on the basis of all characters ex- cept their pappus. The two genera share most notably a stiff type of vegetative hair, short, transversely polarized endothecial cells, a re- duced number of ribs on the achene, a somewhat irregularly spinulose pollen, and a chromosome number based on x = 9. Philoglossa is generally more advanced in its more reduced pappus and its two rather than four ribs on the achene. Re- ports by Turner et al. (1979), Olsen (1980), and Jansen et al. (1984) firmly establish n = 9 as the number for E. vulcanica. Unfortunately, a num- ber of attempts to count the closely related E. polymnioides DC. during the present study have failed, but one count of n = 9 has been provided by Olsen (1980). The report of n = 18 for Phil- 1985] oglossa stenocarpa by Diers (1961) fits well with the n = 9 reported from the closely related Erato. The limited examples give little indication of the full distribution of polyploidy in the two genera; nevertheless, the polyploidy in the smaller though not necessarily more herbaceous Philoglossa is still another example of the lack of correlation between habit and chromosome number in the tribe. Chrysactinium and Munnozia also have geo- graphic ranges that overlap in the Andes of Ec- uador and northern Peru, but the latter, larger genus extends much more widely to Costa Rica, Venezuela, and Bolivia. The two genera share a tomentose type of vegetative pubescence, verti- cally or obliquely polarized endothecial cells, usually ten-ribbed achenes, pollen with regularly distributed — and chromosome numbers of n = 10 to ca. 13. Chrysactinium hieracioides now has n. reports by Turner et al. (1967), Dillon and Turner (1982), and the new reports (see Table 1) indicate a chromosome number of n = 12, the same number reported from the Pe- ruvian species C. rosulatum (as C. acaule, Dillon & Turner, 1982). The new report of n = 13, 14 in C. rosulatum (see Table 1) and reports of pos- sible n = 13 in C. hieracioides may or may not represent true variation in the genus, but the re- ports of n = 12 seem more reliable, and that number also seems basic in the closely related Munnozia. In the examination of the King col- lections of Chrysactinium from Peru (9128, 9139, 9200, 9245, 9273), Powell noted a uniformity of karyotype that contrasted with that of a Fer- reyranthus studied with uncertain results at the same time. In Munnozia, most reports are based on M. of n = 10 or 11. There is one report of n = ca. 12 (see Table 1). However, on the basis of the drawings provided (Figs. 2, 3), Strother (pers. comm.) suggests the report of n = 10 is in error. He suggests that the cell illustrated in Figure 2 exhibited 8,, + 2,, for 2n = 24 or 10, + liy for 2n — 24. This presumes that the larger configu- rations noted by Powell are actually multivalents even though they exhibited meiotic behavior that is characteristic of bivalents. Other species rep- resenting the great diversity of the genus, in- cluding M. jussieui (Jansen & Stuessy, 1980), M. lyrata and M. ferreyrii (Dillon & Turner, 1982), and M. hastifolia and M. maronii reported here, show mostly n = 12 or ca. 130r n = ca. 24, which ROBINSON ET AL.—LIABEAE CHROMOSOME NUMBERS C? Q 99, "i SN (0o Q SS A o S 0c T T 0225 e 3 Ficures 2, 3. Munnozia senecionidis Benth. King and Almeda 7834. —2. Metaphase I. —3. Anaphase II. Line = agrees more with Chrysactinium. On this basis we regard x — 12 as basic for the generic pair. The x = 12 of the Chrysactinium-Munnozia group and the x = 14 of typical Paranephelius appear to be two clear examples of aneuploid gain in Liabeae in spite of the evidence from other groups such as Heliantheae (Robinson et al., 1981) regarding the comparative rarity of such gains in chromosome number. We have no inclination to regard either number as the result relationship to the one genus in the tribe, cosmia, having a chromosome number less than n = 9. As already indicated, there is also no close relationship between the Munnozia group and Paranephelius, and the two examples of aneu- ploid gain are thus totally independent. There is, however, no need to accept more than one in- stance of aneuploid gain in Munnozia. The series of numbers in Munnozia is at best a downward series that occurs in a specialized element of the genus. The 7 = 12 is basic to both genera and is obviously ancestral in the group. The occasional reports of still higher numbers in the Munnozia group might indicate that the ancestral number of the generic pair was actually higher than n = 12. Although no direct evidence is available, the . . "1.5 | -nht A š fr. some extinct polyploid ancestor having n = 18. It is notable that series of numbers above the ancestral base number similar to those in Mun- nozia can be found in other tribes of Asteraceae. One such example is in the Microspermum-Ste- via-Piqueria relationship of Eupatorieae. At present we favor an interpretation of that series like that of Munnozia with the highest number in the series being the first, followed by reduction. We suspect that many other seeming series of aneuploid increases should be interpreted in this GENERAL CONCLUSIONS Given the overall pattern of chromosome numbers in the Liabeae, there remain two gen- eral points of interest to be discussed. The foregoing discussion of the genera and subtribes includes mention of many examples of polyploidy in the tribe, mostly concentrated in two series of advanced members of Liabinae. There seems to be an unusual lack of correlation of chromosome number with habit in the tribe, with little tendency for lower numbers in shorter- lived or more herbaceous plants as commonly occurs in other tribes (King et al., 1976; Solbrig, 1977) and in many other groups of plants. How- ever, the polyploids in Liabeae do include all the elements that have their primary ranges north of the Ecuadorian-Peruvian center of the tribe. It is as though the diploid members of the tribe were restricted to their ancestral area, and only polyploids were able to invade new territories. This provides the possibility of an interesting test. Among the genera not yet counted is Aus- troliabum, a genus that has exceeded the ances- tral range of the tribe to the south in Bolivia and Argentina. If the hypothesis is correct, then this genus should also be polyploid. It has been observed (Robinson, 1983a) that intergeneric hybridization has left its mark on many of the large tribes of Asteraceae and may be an important factor in their success. However, there is no evidence of such hybridization in the Liabeae, and it has been suggested that this is correlated with the comparatively restricted size and distribution of the tribe. The present genera of Liabeae that are not geographically isolated tend to have different chromosome numbers, which would inhibit hybridization. The number of Gent basic numbers represented. is re- l number of extant genera (Fig. 1). From a cytologically uniform oup of ancestors, Cacosmia can be seen as an early drop-out with reduction of its chromosome number to n = 7, whereas other genera became isolated by various higher chromosome numbers or by geography. LITERATURE CITED ANDERSON, L. C., D. W. KyHos, T. Mosquin, A. M. P. H. RAvEN. 1974. Chromosome numbers in Compositae, IX: Haplopappus and other Astereae. Amer. J. Bot. 61: 665-671 BENNETT, M. D. 1972. Nuclear DNA content and minimum generation time in herbaceous plants. Proc. Royal Soc. London, Ser. B, 181: 109-135. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 BENTHAM, G. 1873. Compositae. Pp. 163-537 in G. Bentham & J. D. Hooker, Genera Plantarum. Vol- ume 2. Lovell Reeve & Co., Williams & Norgate, London ari H. 1823. iabon. Volume 26, pp. 203-211 nG. Cuvier (editor wil deu des Sciences Na- i Paris. [Reprinted in R. M. & Dawson (editors). irt Cassini on Compositae. 3 Volumes. Oriole Editions, New York.] 1825. Oligacte. Volume 36, pp. 16-18 in G. Cuvier (editor), Dictionaire des Sciences Natu- relles. Paris. [Reprinted in R. M. King & H. W Dawson (editors). 1975. Cassini on Compositae. 3 Volumes. Oriole Editions, New York. 1830. Zyégée. Volume 60, pp. 560-619 in G. Cuvier (editor), Dictionaire des Sciences Na- turelles. Paris. [Reprinted in R. M. King & H. W Dawson (editors). 1975. Cassini on Compositae. 3 Volumes. Oriole Editions, New York.] Diers, L. 1961. Der Anteil an Polyploiden in den Vegetationsgürteln der Westkordillere Perus. Z. Bot. 49: 437—488. DiLLON, M. & B. L. TURNER. 1982. Chromosome numbers of Peruvian Compositae. Rhodora 84: 131-137. HOFFMANN, uin 1894 [1890]. Compositae. Jn H. G. A. En ngler K. A. E. Prantl (editors), Nat. Pflan zenfam. Mm vit 391. Wilhelm Engelmann, Leine JANSEN, R. K. & T. F. Srusssv. 1980. Chromosome counts of Compositae from Latin America. Amer n Bot. 67: 585-594. IAZ-PIEDRAHITA & V. A. FUN 1984. Recuentos cromosomicos en apos de “ran oq Caldasia 14: 7—20. JONES, S. B., 1977. Vernonieae—systematic re- view. Cham 17, pp. 503-521 in V. H. Heywoo J. B. Harborne & B. L. Turner (editors), The Bi- ology and Chemistry of the Compositae. Academ- ic Press, p and New York. KiNG, R. M., D. W. Kvuos, A. M. PowELL, R. H. RAVEN & H. hung 1976 [1977]. Chromo- some numbers in Compositae, XIII: Eupatorieae. Ann. Missouri Bot. Gard. 63: 862-888. Nasu, D. L. 1976. Tribe I: Vernonieae. Jn D. L. Nash L. O. Williams, Compositae. Flora of Guate- mala 24(12): 4-32. NORDENSTAM, B. 1977. Senecioneae and Liabeae— glia ien Chapter 29, pp. 799-830 in V. B. Harborne & B. L. Turner (ed- ion, The Biolo ogy and Chemistry of the Com- posita S x J. . Lóve, Chr ome Number Reports. LXVIL Taxon 29: 366- , D. W. KvHos & P. m some numbers in Hi positae, VI: Secs II. Amer. J. Bot. 54: 205- 213. w sess : N, D. W. KvHos & A. R. KRUCKE- BERG. 1963. ip eet ree numbers in Compos- itae, III: s, deed J. Bot. 50: 131-139. PAYNE, W. W., N & D. W. KvHos. 1964. Chro omosome nune in Compositae, IV: Am- brosieae. Amer. J. Bot. 51: 419—424. 1985] PowELL, A. M. & R. M. KiNG. 1969. ee numbers in the Compositae: Colombian specie Amer. J. Bot. 56: 116-121. , D. W. Kyuos & P. H. Raven. 1974. Chro- mosome numbers in Compositae, X. Amer. J. Bot. 61: 909—913. & 1975. Chromosome num bers i in Compositae, XI: Helenieae. Amer. J. Bot. 62: 1100-1103. Raven, P. H. 1975. The bases of angiosperm phy- logeny: cytology. Ann. Missouri Bot. Gard. 62: 724—764. ———— & D. W. Kyuos. 1961. Chromosome num- bers in Compositae, II: Helenieae. Amer. J. Bot. 48: 842—850. , O. T. Soranic, D. W. KvHos & R. Snow. 1960. Chromosome numbers in Compositae, I: 32. An analysis of the characters and relationships of the tribes Eupatorieae and Vernonieae (Asteraceae). Syst. Bot. 2: 199-208. 1983a. A generic review of the tribe Liabeae (Asteraceae). Smithsonian Contr. Bot. 54: 1—69. 3b. Studies in the Liabeae (Asteraceae). XVI. New taxa from Peru. Phytologia 54: 62-65. . D. BRETTELL. 1974. Studies in the Lia- beae (Asteraceae). II. Preliminary survey of the genera. Phytologia 28: 43-63. : . PowELL, R. M. KiNG & J. F. WEEDIN. 1981. Chromosome numbers in Compositae, XII: Heliantheae. Smithsonian Contr. Bot. 52: 1—28. SKVARLA, J. J., . TURNER, V. C. PATEL & A. S. Toms. 1977. Pollen morphology in the Com- positae and in ved etis related families. sis quai pp. 141-265 in V. H. Heywood, J. B Harborn . L. Turner (editors), The Biology and oe of the Compositae. Academic Press, London and New York ROBINSON ET AL.—LIABEAE CHROMOSOME NUMBERS 479 SorBRIG, O. T. 1977. kin saan cytology and evolution in the family Compositae. C B e istry of the Compositae. Academic Press, London and New Yor L.C. ANDERSON, zd W. KvHos & P. H. RAVE 1969. Chromosom mbers in Compositae, VIE Astereae III. p pw n ; ‘Bot 56: 348-353. , D. W. KvHOS, eae Saatine P. H. RAVE 1972. Chromosome n in Compositae, VIIL Heliantheae. Amer. J. p 59: 869-878. , L. C. ANDERSON, D. W. Kyuos, P. H. RAVEN Ruben, 1964. omosome numbers positae, V: Astereae IL Amer. J. Bot. 51: TowB, A. S., K. L. "Ice D. W. Kyuos, A. M. PowELL & P. H. RAvEN. 1978. Fossil di numbers in us Compositae, XIV: Lactuceae. Amer. J. Bot. 65: 717-721. Torres, A. M. & A. H. Liocier. 1970. Chromosome numbers of Dominican Compositae. Brittonia 22: 0-245. TR B. L. 1981. New species and sonibinatipns teraceae). with a revisional conspectus in D oups. Brittonia 33: 401-412. A. M. PowrziL & J. CUATRECASAS. 1967. Chromosome numbers in Compositae, XI: Pe- ruvian species. Ann. Missouri Bot. Gard. 54: 172- 177. , J. BACON, L. URBATSCH & B. SIMPSON. Chromosome numbers in po American Com- positae. Amer. J. po 66: 173-1 WAGENITZ, G. 1976. shots and phylogeny of the Compositae P Pl. Syst. Evol. 125: 9-46. CHROMOSOME STUDIES ON HYDROMYSTRIA LAEVIGATA (HYDROCHARITACEAE)! EDUARDO A. MOSCONE AND Luis M. BERNARDELLO? ABSTRACT Chromosomes of individuals from two Argentine populations of Byeromysias pian, ilar 8. A. T. Hunz. were analyzed for n umbers, meiotic behavior , and karyotype. It has n Both populations have regular meiosis and show no significant aan differences Mira them. The k 6 sm pairs + 4 st pairs. No satellites were foun Hydromystria is a monotypic genus of Hydro- charitaceae. The only species, H. laevigata, is an aquatic plant that grows in Mexico, West Indies, and South America (Hunziker, 1981, 1982). Some authors include Hydromystria within Limnobi- m (Dandy, 1959; Cook et al., 1974; Cook & Urmi-Kónig, 1983), while others recognize them as distinct although closely re- lated (cf. Hunziker, 1981, 1982). No comprehensive cytological study has been carried out on H. laevigata, although some chro- 2n = 26-28, and the same author (1940) pue 4n = 56 in counts on colchicine-induced tetra- ploid root cells. Recently, Urmi-Kónig (in Cook Urmi-Kónig, 1983) reported five different chromosome numbers for this plant [sub Lim- nobium laevigatum (Willd.) Heine]: 2n — 26, 27, 28, 29, 30. In the present contribution, the so- matic and gametic numbers of two Argentine populations of H. /aevigata are reported, as well as their meiotic behavior and karyotype MATERIALS AND METHODS The materials came from two morphologically identical populations (hereafter named P, and P.) collected in the following localities: Argen- tina, Córdoba: P, = Punilla: Cabalango, Bernar- dello, Weigel & Moscone 359 bis, 29 Aug. 1982; P, = Calamuchita: Emblase Río III, camino a la segunda Usina, Moscone, Carrizo & Moscone 1, 24 Oct. 1982; we analyzed 35 plants of P, and 16 of P,. Voucher specimens have been depos- aryotype is similar in both populations: asymmetrical and bimodal; it is formed by: 4 m pairs + ited in the herbarium of Museo Botánico de Cór- doba, Argentina (CORD). Mitotic chromosomes were examined in root tip squashes; the root tips were fixed in 1 : 3 acetic acid: ethanol mixture after pretreatment in a sat- urated solution of p-dichlorobenzene in water for 5-7 hours at room temperature, and then stained with alcoholic hydrochloric acid carmine (Snow, 1963) for at least 12 hours. Karyograms were prepared from microphotographs, using the ter- minology introduced by Levan et al. (1964). Two parameters were used: the arm ratio and the cen- tromeric index. The chromosomes were first ar- ranged according to their increasing arm ratio and then according to the decreasing order with- in each group. The idiogram is based on ten metaphase plates (five from each population). Meiotic behavior was observed in pollen mother cells obtained by squashing young an- thers fixed in 1:3 acetic acid : ethanol and stained with acetic carmine. All the permanent mounts were made with Bradley's method (1948). The statistical methods followed Sokal and Rohlf (1979); as regards the :-test, the levels of signif- icance considered were 0.05 and RESULTS Meiotic observations. Table 1 summarizes the data obtained from the analysis of meiotic be- havior. Both populations have regular meiosis, forming usually 14 bivalents (Fig. 1A, B). Some- times univalents were observed: a 13 II + 2 I configuration was detected in 2.296 of P, cells and in up to 3.396 of P, cells. ! The authors thank Prof. A. T. Hunziker for suggesting the iiis and for his e support, Dr. J. H. Hunziker for the critical nds of fund manuscript, Dr E. Cocucci and M Münch for technical assistance. This work was supported i rt by a grant from the Consejo de rendo fiuc amm Científicas y i inia ue d d Pub de Córdoba, Pastas This paper was presented on 22 Sept. in the XIX as Argen organized by the Sociedad Argentina de Botanica at Santa E esie dudes de Biología Vegetal (IMBIV), Casilla de Correo 495, 5000 Córdoba, Argentina. ANN. Missouni Bor. GARD. 72: 480-484. 1985. 1985] Meiotic behavior of Hydromystria laevigata. TABLE 1. MOSCONE & BERNARDELLO — CHROMOSOMES OF HYDROMYSTRIA 481 23 Bd 25 ar S. EE gogu LE edr SORS S s" > ë as ad E š EG Chiasmata Total Chromosomal Associations in Diplotene- Diakinesis per Cell: Range and Mean + Standard Error 1.48 +0.026 10-19 16-25 0-2 13-14 0.044 +0.044 +0.365 +0.319 +0.295 +0.022 1.49 +0.016 P, 0.065 +0.037 fis = 0.34 +0.260 t, = 0.99 +0.226 tias = 0.12 +0.168 tiss = 0.63 P = +0.019 t s = 0.32 t-test tias = 0.34 P 0.7 P = Pe B e ç w A ° aQ O 0 RO V. ` n! i o ut š % A,A p? 10 pm 4 s=. * _ FIGURE 1. Hydromystria Laid ll Diakine- s, 14 II (Moscone et al. 1).—B. Diakinesis, 14 II (Ber- nardello et al. 359 bis). — C. First mitotic metaphase of pollen grain, n = 14 (Moscone et al. D. Two bivalents much bigger than the others have from two to six chiasmata; they correspond to pairs 1 and 11 from the karyotype, although it is not always possible to distinguish one pair from the other. The remaining bivalents are shorter, similar to each other in length, and gen- erally have only one terminal chiasma. Some- times they sustain two terminal chiasmata, thus forming ring bivalents. oieri p the asso- ciation of some bivalents to the nucleolus and the first mitotic division of the microspore were observed (Fig. 1C). Mitotic observations. The somatic chromo- their arm ratio, the chromosomes can be clas- sified as follows (Fig. 3A, B): 4 m pairs (1 to 4), 6 sm pairs (5 to 10) and 4 st pairs (11 to 14). It should be emphasized that no chromosome is satellited, at least according to our observations. aryotype is asymmetrical and bimodal. E: E 2. Mitotic metaphases of Hydromystria Mese 2n = 28.—A. Moscone et al. 1. —B. Bernar- dello et al. 359 bis. 482 A Go S ) | L ) i FIGURE 3. ANNALS OF THE MISSOURI BOTANICAL GARDEN SEBBE Karyograms and idiogram of Hydromystria laevigata.—A. Karyogram of Figure 2A. Scale, 5 [VoL. 72 œ © = o E ert = N = w = A sm st sm 5 pm i st _—— a | e òè à €* (à 8 9 10 11 12 13 14 1 pm IDEE m. — B. Karyogram of Figure 2B. Scale, 5 um.— C. Idiogram. Scale, 1 um Pairs 1 and 11 are considerably larger than the others, which are in general homogeneous i in size for which are presented in Table 2. Pair 1 is the longest, and pair 10 is the shortest. The lowest arm ratio corresponds to pair 4 and the highest one to pair 11. DISCUSSION From the meiotic behavior and the f-test re- sults it is evident that both populations have regular meiosis and show no statistical differ- ences in their meiotic systems. Furthermore, the analysis of mitotic chromosomes demonstrates a striking similarity between P, and P, karyo- grams although Chaudhuri and Sharma (1978) have mentioned, - ed on their investigations on Vallisneria spiralis L., Ottelia alismoides Pers., Hydrilla See ihe (L. f.) Royle, and oth- ers, the frequent presence of cytotypes in Helo- biales. These authors found several cytotypes of those species varying in both chromosome num- ber and morphology. This circumstance has been reported in other members of the Hydrochari- taceae by other botanists (Bhattacharya & Gosh, 1976; Harada, 1956; Misra, 1974; Sharma & Bhattacharyya, 1956) In contrast to our results, Urmi-König (in Cook & Urmi-Kónig, 1983), studying plants from Guadeloupe, Buenos Aires, and of unknown or- igin, indicated 2n = 26, 27, 28, 29, 30 for H. laevigata with several chromosome numbers in an individual specimen. Although some differ- ences can be drawn from her karyogram data, it is difficult to compare them because she did not state the terminology used and did not show any figure or photograph. She found one pair of long submetacentric and one medium pair designed as acrocentric. After the widely accepted termi- nology of Levan et al. (1964), both long pairs are m and sm according to our results. Urmi-Kónig did not report sm chromosomes within the short pairs but did report a number of acrocentric ones Geitler (1938) could find neither satellites nor 1985] MOSCONE & BERNARDELLO— CHROMOSOMES OF HYDROMYSTRIA 483 TABLE 2. Measurements and indexes of somatic chromosomes of Hydromystria laevigata. Measurements 6 given in um. Abbreviations after Levan et al. (1964). Chromosome Lengths: Range and Mean + Standard Error omen- Pair S l c r i clature 1 1.72-2.55 2.55—4.00 4.33-6.55 1.53 39.53 m 2.00 + 0.092 3.06 + 0.152 5.06 + 0.243 2 0.80-1.05 1.05-1.45 1.90-2.50 1.32 43.19 m 0.92 + 0.028 1.21 + 0.046 2.13 + 0.070 3 0.73-1.07 0.90-1.38 1.70-2.45 1.27 44.10 m 0.86 + 0.036 1.09 + 0.052 1.95 + 0.083 4 0.60—0.80 0.70-0.95 1.30-1.75 1.15 46.53 m 0.67 + 0.019 0.77 + 0.024 1.44 + 0.042 5 0.68—0.90 1.35-2.05 2.08-2.90 2.09 32.35 sm 0.77 + 0.024 1.61 + 0.065 2.38 + 0.081 6 0.60-0.92 1.28-1.83 1.90-2.75 2.17 31.51 sm 0.69 + 0.030 1.50 + 0.056 2.19 + 0.078 7 0.60-0.80 1.15-1.65 1.80-2.45 1.97 33.66 sm 0.69 + 0.020 1.36 + 0.051 2.05 + 0.066 8 0.57-0.85 1.02-1.50 1.60—2.35 1.77 36.13 sm 0.69 + 0.027 1.22 + 0.046 1.91 + 0.073 9 0.45-0.65 0.83-1.15 1.30-1.80 1.90 34.44 sm 0.52 + 0.019 0.99 + 0.027 1.51 + 0.043 10 0.40—0.58 0.80-1.10 1.20-1.68 2.02 33.09 sm 0.45 + 0.018 0.91 + 0.029 1.36 + 0.044 11 0.60—0.90 2.65—4.20 3.30-5.10 4.43 18.42 st 0.70 + 0.030 3.10 + 0.144 3.80 + 0.172 12 0.35-0.50 1.35-2.10 1.70-2.60 3.79 20.87 st 0.43 + 0.019 1.63 + 0.067 2.06 + 0.082 13 0.37-0.60 1.23-1.90 1.60-2.50 3.31 23.20 st 0.45 + 0.021 1.49 + 0.062 1.94 + 0.082 14 0.37-0.52 1.20-1.70 1.60-2.20 3.23 23.63 st 0.43 + 0.016 1.39 + 0.053 1.82 + 0.068 secondary constrictions in this species. We agree with his observations, although Urmi-König (in Cook & Urmi-König, 1983) reported that one pa bears satellites but without specifying which Em other Hydrocharitaceae examined (Mis- ra, 1974; Chaudhuri & Sharma, 1978; Bhatta- charya & Gosh, 1976; Sharma & Bhattacharyya, 1956) Hydromystria has an asymmetrical karyo- type. The basic number of x = 7 for H. laevigata is one of several previously recorded for the family. The basic numbers, in decreasing order of fre- quency, are: x — 8, 11, 10, 7, and 6 (Fedorov, 1969; Goldblatt, 1981, 1984; Moore, 1973, 1977; Packer & Witkus, 1982). Some of the genera of Hydrocharitaceae have not been studied cytologically: Apalanthe, Ap- pertiella, Limnobium sensu stricto, and Thalas- sia, and the genera Nechamandra and Stratiotes are insufficiently known. A cytological study of Limnobium may help elucidate its relation to Hydromystria, on which we are currently work- ing. LITERATURE CITED BHATTACHARYA, G. N. & D. K. Gos 1976. Cyto- logical analysis of different cyiotypes of Ottelia alismoides Pers. Caryologia 195-202. BRADLEY, M. V. 1948 de niet for making aceto- carmine squa uashes anent without removal of cover slip. Stain Technol, 23: 41-44. 484 CHAUDHURI, J. B. & A. SHARMA. 1978. Cytological studies on three aquatic members of Hydrochar- itaceae in relation to their morphological and eco- m EDS Cytologia 43: 1-19. Cook, C. D . URMi-KóNiG.. 1983. A revision of the 5 Limnobium including H MM (Hydrocharitaceae). ders Bot. 17: ,B.J. GuT x, J. EM “SEITZ. 1974. Water P of the World. Dr. W. Junk , The Hague. DANDY, J. E. 1959. 5. (keys for the tribes and genera). Pp. 540-541 in J. Hutchinson a Mis ia ies of Flowering Plants. Vol- on Press rd. PR. P eat) 1969. Chromosome diei of dis ring Plant .K ov Bot. Leni d {Reprinted by [o Koeltz Sci. PH 74. Über das Wachstum von Chro- ozentrenkernen und zweierlei Heterochromatin bei Blütenpflanzen. Z. Zellf. Mikroskop. Anat., Abt. B, Chromosoma 28: 133-1 . 1940. Die Polyploidie der Dauergewebe hó- herer Pflanzen. Ber. Deutsch. Bot. Ges. 58: 131- 142 Mire P. (editor). 1981. Index to plant vd numbers 1975-1978. Monogr. Syst. Nissan Bot Gard. 5: 279. 984. Index to plant chromosome numbers 1979-1981. Monogr. Syst. Bot. Missouri Bot. Gard. HARADA, L 1956. Cytological studies in Helobiae, I. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Ch * "m ologia 21: 306-328. Hunziker, À. T. 1981 Todd. laevigata (Hy- dm) en el centro de Argentina. Lorent- zia 4: 5- —. 1982. Observaciones biológicas y taxonómi- cas sobre RAM dp ud (Hydrocharita- ceae). pu -477 LEvAN, A., K. FREDGA MS A. SANDBERG. 1964. No- menclature for centromeric Tue on chromo- somes. Hereditas 52: 201- MisRA, M. P. 1974. iretur studies in Ottelia alismoides Pers. Cytologia 39: 419—427. MOORE s J. (editor). 1973. Index to plant chro- moso e numbers for 1967-71. Regnum Veg. 90: e = 1 £ . Index to plant chromosome numbers 124. . 1982. Hydrocharitaceae. In ^. Lóve (editor), IOPB Chromosome Number Reports LXXV. Taxon 31: 363 SHARMA, À. K. & B. BHATTACHARYYA. 1956. A study of the cytology of four members of the Hydro- Phyton (Buenos Aires) 6: 121-132 Snow, R. 1963. Alcoholic hydrochloric acid-carmine as stain for chromosomes in squash preparations. Ee uus 38: 9-13. , R. R. & F. J. ROHLF. 1979. Biometría. H. “Blume a Madrid. [Translated from the 1969 English 1st edition.] STUDIES IN NEOTROPICAL PALEOBOTANY. III. THE TERTIARY COMMUNITIES OF PANAMA — GEOLOGY OF THE POLLEN-BEARING DEPOSITS! ALAN GRAHAM, R. H. STEWART,? AND J. L. STEWART? ABSTRACT amples pis ciae] fossil "Hh and spores were collected from six sites representing five Tertiary he the La Boca (youngest). The Gatun samples were ta and depositional environments are marized fe along the Gaillard Cut section of the Canal. The early Miocene La Boca Fo ae gage dies ncillo Formation is middle(?) to late Eocene, based on foraminifera ken from a series of wells drilled in Gatun Lake. The location, stratigraphy, lithology, the paar soar paleoenvironments, and biogeographic implications will be considered for each of the microflora The present studies on the Cenozoic history of neotropical vegetation began in 1964 with a collection of pollen-bearing samples from Ter- tiary deposits in central Panama. Subsequently, other samples were obtained from the Oligocene of Puerto Rico and the Miocene of Veracruz, Mexico. At that time the vegetation of Central America was poorly known, but research by staff of the Missouri Botanical Garden, the Field Mu- seum, and various institutions in Central Amer- ica was rapidly providing new information on the composition of the flora, and on the range, i aw: and biogeographic relationships of its . By comparison, the vegetation of ico and Veracruz, was somewhat better known (Howard, 1973; Gomez-Pompa, 1973), and it seemed logical to begin paleofloristic studies on sibi Sopa younger deposits in these areas where ern analogs were more clearly defined. Con- LE. our first studies were on the middle Oligocene San Sebastian palynoflora from Puer- to Rico (Graham & Jarzen, 1969), followed by the late Miocene Paraje Solo palynoflora from Veracruz, Mexico (Graham, 1976 The decision to defer consideration of the Pan- ama material was fortunate for several reasons. The older Gatuncillo assemblage from Panama proved the most difficult of or our neotropical stud- les, and recently on the modern vegetation was most helpful in making paleoenvironmental reconstructions. The com- pletion of the “Flora of Panama" (1980) and Croat's (1978) “Flora of Barro Colorado Island," initiation of Burger's “Flora Costaricensis," and the “Flora Mesoamerica,” coordinated through the Missouri Botanical Garden and Kew, are making both specimens and information avail- able to our studies on vegetational history. Re- try, 1 collection, used for identification of the fossil palynomorphs, has increased from 10,000 slides in 1964 to nearly 22,000. From the standpoint of geology, the recently revised “Geologic Map of the Panama Canal and Vicinity, Republic of Panama” (Stewart et al., 1980) and modern plate tectonic summaries (e.g., Coney, 1982; Raven & Axelrod, 1974) are important for placing the pa- leobotanical data in a sound geologic context. Pollen-bearing samples from five Tertiary for- mations were collected in Panama within or ad- jacent to the former Canal Zone (Figs. 1—3). These ! Supported by NSF grants DEB-8205926, DEB-8007312, and GB-11862. ? Department of Biological Sciences, Kent Sta tate University, Kent, Ohio 44242. ? Chief Geologists (retired), Panama Canal Commission. Present address: 204 Tampa Downs Blvd., Lutz, Florida 33549. ANN. Missouni Bor. GARD. 72: 485—503. 1985. 486 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 80°00 45 30 N Ç 9 PORTOBELO —9°30' "a < ? a ° i M S: v? coron £ We s n £ Š e: / È | 5 rd - q: Yeo QU f 7 ie 9 diii t . —~— he N GS > P ion Lo | = w. —9°15 ñ i d N ^ BOYD NUEVO CHAGRES f _ `Ë ROOSEVELT D Buenos Aires N L b < N Ç Cumbres CoU N ` š ‘a = ce — 9900 SE [coN PANAMA `. BALBOA N i d \ \ A tsa SS sem i i x LU QO » : Š, » SY LA CHORRERA BAHIA í MJ l TABOGA Q DE = 3 @CAPIRA 9 PANAMA e a, / Scale ( 1:500,000 d (0) 5 IO miles SEE Å (0) 10 kilometers © CHAME A | | | FiGURE 1. Locality map for Tertiary pollen and spore- -bearing samples, Republic of Panama.— A, B. ard , Panam nal.—C. Cucaracha Formation DE Gatuncillo Formation [Middle(?) to late Eocene], n near Alcalde Díaz (Pefioncito). — 1. Drilling iu through the Culebra Formation (early Miocene), Contractor's Hill locality in front of Gold Hill (core GH-9). — 2. Drilling n through the Gatun Formation (late Miocene to Pliocene), Gatun Lake (cores SL-48, 49, 60, 103 La Boca 1985] GRAHAM ET AL.—NEOTROPICAL PALEOBOTANY III 9°0500" 79°37'30" =, I \ "d TN ) ` Scale J s SN 25,000 ` TIc "S X / o | i 2 kilometers 3 — Tica x A Tico Q A < a o I miles / 2 X H li ¢ \ J ) > A \ ? o i Tic Š! Pu d ' 4 ^. Ñs j i ; EE tm e I x \ "-— — TI Ens I Ó Tic BOR e S CASCADAS cec B | — w di N 79°40'00" zd p and B (La Boca Formation) are along ENS DADA `. < PEAR S — ie we SWCE i a. 123755: "s Linear DA. Z eli LS -2 u SV ^S Tca BS, 9°02 50" RE 2. Generalized surface geology map, Gaillard Cut section of Canal, Republic of Panama. Collecting a Canal-facing slope overlain by the Las Cascadas Formation (Tlc), and C (Cucaracha Formation) is unc in part by the Pedro Miguel Misi d (Tpa). Note extensive faulting. Modified from Stewart et al. (1 Limestone member (coralliferous limestone) of La Boca Cucaracha; Tlca basalts (middle and late M. a are the middle(?) to late Eocene Gatuncillo For- mation, the early Miocene Culebra, Cucaracha, and La Boca Formations, and the late Miocene to Pliocene atum Formation. Th gthe Gail- lard Cut, as reflected by the faults shown i in Fig- ure 2, presents problems in recognizing the age and stratigraphic : relations between the numer- l, are frequent- ous, sma ly concealed by dense vegetation cover. In ad- dition, special problems arise from maintenance procedures used along the Canal, which often involve removal of vast amounts of slope sedi- ments (Figs. 9-11). This significantly alters the dscape, even between collecting seasons. These procedures are required by the unusual geologic configuration through which the Canal was con- structed. During the Tertiary, lava filled many of the valleys and river basins. Subsequent ero- sion of the softer surrounding sediments below the basalt-filled basins produced a present-day massive, dense basalts resting on top of softer mbols: Tlc— Las Cascadas Formation; Tl—La Boca; Tle— Emperador E] ca Formation; Tpa— Pedro Miguel; Tcb—Culebra; Tca— o Las Cascadas Formation (early Miocene); Tb— intrusive and extrusive underlying sediments. When the Canal was dug through this mosaic, the weight of the overlying basalts began forcing enormous volumes of sed- iment into the Canal. The scars from these slip- pages are a common and characteristic feature of the banks along the Canal. To reduce land- slides, the slope face is cut back, and phic re- lationships La neo many of the collecting sites. TI positional environments, and ancrently mabe i ages for five Tertiary formations in Panama from which pollen-bearing samples have been col- lected. Subsequent studies will trace the history of the vegetation in central Panama as preserved in the palynofloras of the Gatuncillo, Culebra, Cucaracha, La Boca, and Gatun Formations. THE GATUNCILLO FORMATION The Gatuncillo Formation was named by Thompson in Panama Canal reports for 1944 488 79°35 | canal ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 79°30) ND | —9°10' ( ¿ pT N l Re | ç Tpm Ny Xo, Tas ra Es Las Cumbres CARRETERA (- BOYD ROOSEVELT T O Bess 5 o 5 miles kilometers Generalized surface geology map, vicinity of Alcalde Díaz [Peñoncito; locality D—Gatuncillo FIGURE 3. Formation, middle(?) to late Eocene]. Symbols: Tp—Pana i i i i and extrusive andesite (Oligocene and earl ocene); T porphyry (Miocene); Ter Caraba (late Oligocene); Qa-Holocene oea or m an For other symbols see puso to Figure 2. and first published in Woodring and Thompson (1949). Maximum thickness is about 1,000 m and the formation *unconformably overlies a basement complex of unknown, possibly Cre- taceous age consisting of strongly deformed al- tered volcanics-flows, pyroclastics, and indurat- ed fine-grained sediments that probably were ama Formation (early to late Oligocene); ien —Pa mn originally fine-grained tuffs" (Woodring & Thompson, 1949: 227). In the vicinity of our collection sites the for- mation outcrops around the margins of Madden Basin and Lake Madden north of Panama City (Fig. 3). The Rio Gatun fault terminates the ex- posure to the north, and the Azota fault defines 1985] GRAHAM ET AL.— NEOTROPICAL PALEOBOTANY III 489 portions of its western limits. A few isolated out- crops occur to the west along Gatun Lake an at the north end of Culebra Reach. Elsewhere the formation has been identified in cores to the south as far as Gold Hill and north of Gamboa. In addition, it has been recognized eastward into the Bayano River basin and westward as far as the Costa Rica-Panama border. There are also massive outcrops in the southern part of the Azuero Peninsula. Local extrusive basalts of Miocene age overlie small areas of the Gatuncillo Formation, and just north of Alcalde Díaz (Penoncito) there are very local deposits of the Panama Formation forming the tops of hills. The latter is early to late Oli- gocene in age and composed mainly of agglom- erates (andesitic in fine-grained tuff) and stream- deposited conglomerates (an agglomerate is composed of angular volcanic ejecta, in contrast to conglomerate particles rounded by stream transport). The Panama Formation outcrops are rapidly being altered, especially near the former Canal Zone, because these local hilltop deposits are extensively removed for valley fill and road construction. The Gatuncillo sediments consist of mud- stones, siltstones, quartz sandstone, impure ben- tonite, and coralline and foraminiferal lime- stone. Interbedded are lenses of lignite ranging from a few centimeters to 1-2 m in thickness. Locally there are overlying deposits of marine limestone. The sequence is typical of nearshore deposition in tropical environments, since lig- nites presently form under such conditions (Co- hen & Spackman, 1972; Scholl, 1964a, 1964b), and those in the Gatuncillo sediments contain Rhizophora pollen. The corals fringing the vol- canic islands that occupied the region of pres- ent-day central Panama during the Eocene, and contributing to the coralline component of the Gatuncillo Formation, further reflect tropical de- positional environments. Surrounding most of the Gatuncillo Formation to the east, north, and west are extensive areas mapped as pre-Tertiary. TI š 14331 14 A lt; A 1 11160] and tuff, and other dioritic and dacitic intrusives. Woodring (1957-1982) has made extensive studies on the Tertiary mollusks from the Canal Zone. From the Gatuncillo Formation he reports a fauna most similar to those elsewhere of middle to late Eocene age. Cole (1952) studied the larger foraminifera of the Gatuncillo Formation. He reported 21 species, 18 of which are recorded elsewhere in the late Eocene. The remaining three occur in the middle Eocene, and two of these d Uni: inis and Fabiania cubensis) from the middle Eocene. Con- c uud. us (1952) refers the Gatuncillo For- mation to the middle(?) and late Eocene. Samples for pollen and spore analysis were obtained from a roadcut section near Alcalde Díaz just off the Boyd-Roosevelt highway (our locality D, Figs. 1, 3-5; samples 1-15; 9?7'N, 79°32'W). The concurrence of the mollusk and foraminifera data, in the absence of any conflict- ing stratigraphic information, indicates the paly- nomorphs recovered from the Gatuncillo For- mation represent remnants of a middle to late Eocene plant community growing in the region approximately 40 Ma. THE CULEBRA FORMATION The Culebra Formation (Hill, 1898) takes its name from a town located in the central Gaillard Cut during the construction period. Or- mation outcrops in very limited areas on either side of the Canal along the Gaillard Cut (Fig. 2). Stratigraphic relations are exceedingly complex because of extensive faulting in the area. In a distance of ca. 3 km, more than 113 faults have been observed along Las Cascadas Reach to the north. In one section adjacent to the Canal, along Las Cascadas, Empire, and Culebra Reaches, left- handed fault motions represented in a distance of 4 km total 4 km of displacement. The Culebra Formation lies unconformably on the Gatuncillo Formation, and has a thickness ofabout 165 m. Itisoverlain by the Las Cascadas Formation as revealed by a core (SC-108) drilled at the Las Cascadas Reach (9?04'N, 79°41'W; Canal Station 1615). The Culebra is composed mainly of calcareous sandstones and siltstones, with interbedded lenses of coalified lignite and carbonaceous shales. Like the Gatuncillo For- mation, the Culebra also represents a nearshore, transgressive depositional environment and specifically an estuary deposit as evidenced by the very thinly bedded, dark, fine-grained sedi- ments, some of which contain fragments of land plants. In the older literature reference is made to the Empire or Emperador limestone member of the Culebra Formation. This **massive, cream- colored coralliferous limestone, formally ex- posed in shallow quarries at Empire, was named the Empire limestone by Hill (1898: 195—196). Empire, now abandoned, was a village on the original line of the Panama Railroad near Cu- lebra and about half a mile west of Gaillard Cut" 490 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURES 4, 5. Roadside exposure of the middle(?) to Late Eocene Gatuncillo Formation near Alcalde Díaz (9°7'N, 79*32'W; locality D), Republic of Panama (January 25, 1983). —4. General view showing basal lignite and overlying siltstone. — 5. Close-up of lignite area shown in lower- left corner of Figure 4. 1985] MU S£ FIGURE Core material from the lower Miocene Culebra Formation was obta GRAHAM ET AL.—NEOTROPICAL PALEOBOTANY III 6. The southwest face of Gold Hill on the Gaillard Cut section of the Canal, Republic of Panama. ained from a well drilled in front of Gold Hill on the west side of the Canal (9*02'N, 79*38'W; drill site 1 on Fig. 1). The subsurface stratigraphy for the Culebra Formation at this site is summarized in Tables 1 and 2 from the well log data. e Miocene or early Pliocene age (above) from upper lower Miocene agglomerates of the unnamed basalt of lat Pedro Miguel Formation (below (Woodring & Thompson, 1949: 237). The Em- perador is now included as a member within the lower La Boca Formation. Woodring (1957-1982) assigns the Culebra to the lower Miocene based on mollusk data. Strati- graphic relations with overlying fossiliferous de- posits, and one tentative potassium-argon date from the Cucaracha Formation (see below), fur- ther suggest a lower Miocene age. In previous stratigraphic columns (e.g., Woodring & Thomp- son, 1949), the Culebra Formation was assigned to the upper Oligocene. A well was drilled through the Culebra For- mation (locality 1 on Fig. 1) that provided fos- siliferous samples and log data detailing the sub- Subsequently the cores were discarded, but cop- ies ofthe logs, unprocessed samples, vials of pro- cessed material, and slides are preserved in the palynology collections at Kent State University. The line separates an The samples were obtained from Hole No. GH-9; latitude 9°02’N + 2912.3, longitude 79°38'W + 5561.5; ground elevation 29.6 m (+96.1 ft.); core recovery 124 m (402.6 ft.; 80.5%) from a total section of 154 m (500.2 ft.); project — Gold Hill Investigation; location—in front of Gold Hill on the west side of the Canal and in front of the Model Slope which is just north of Contractor's Hill (Fig. 6). Fifty-seven samples were taken along the 124 m section (Table 1), primarily from carbonaceous layers within the predominantly sandstone section (labeled in our collections as PAN Core, Culebra Formation). From these, 21 samples were processed and all contained fossil pollen and spores. The stratig- raphy and lithology for portions of the core sam- pled are given in Table THE CUCARACHA FORMATION The Cucaracha Formation outcrops on both sides of the Canal from Hodges Hill southward to the area of Pedro Miguel Locks. The most 492 TABLE 1. Samples from Panama Canal Company's Hole No. GH-9, Contractor's Hill locality, Culebra Formation (lower Miocene), Republic of Panama. Fig- ures represent depth in feet along the core. Arrange- t of the samples is subdivided according to core box sequence used by the PCC. Although these boxes and samples have been discarded, the sequence is still referred to in the summa n of the Geological Field Log of the Contractor's Hill locality, Gold Hill Investigation. Box 12 Box 13 Box 14 Box 15 Box 16 of 17 of 17 of 17 of 17 f 17 370.2* 392 424 446.7 474 376.58 394.3 424.6 4482 476 3778 407^ 425* 449.4 479 378.4* 409 427.5 450 481 3822 411 428.6 451 482.5 383 412.6 433.3 4528 483.6 385.7 415.58 435 454.6 486 388.7 437.98 456° 488* 438.6 459.6 490.6* 440.7 461? 4914 441.9 463.48 491.68 442.98 466 494 444.6 469.8 494.9 70.67 495.8 496.8 ? Samples from which fossil pollen and spores have been recovered. prominent and accessible outcrops are on the southwestern side of the Cucaracha Reach sector of the Gaillard Cut between Canal Stationing 1983 and 2010. The formation is named after a railroad station and a small town that existed prior to Canal construction. The Cucaracha Formation is composed mostly of bentonitic clay shales, tuffaceous siltstones, and sandstones with discontinuous lenses of lig- nite and freshwater conglomerates. The total thickness of the Cucaracha Formation is about 183 m. The formation is divided into an upper and a lower zone. A layer of ash flow or ignim- brite divides the two zones. r part consists of the following: clay 5 shale, and the ash flow at the base. The dip of the beds is 15°, and the dip of the slope is about 20°. The lignites range from a black, friable, fissile, lignite shale to a coalified lignite. All of these beds in the upper part are in fault contact with the basalt in the upper part of the hill. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 The ash flow or ignimbrite is a layer of volcanic debris 8 m thick with tentative potassium-argon dates of 22.2 + 1.7 and 18.9 + 2.2 Ma (Wood- ring, 1982: 551). The lower 0.5 m of the ash flow contains nu- merous carbonized logs, twigs, and roots. In places, the old stumps were found still upright with roots extending into the strata below. This eruption. Immediately to the north of the area just north of Contractor's Hill at the Model Slope, 7596 of the log bases exposed during Canal wid- ening excavations pointed north and northeast- erly indicating the direction from which the erup- tion came The 100 m of strata below the ash flow consists primarily of alternating, recurrent layers of clay shale, tuffaceous siltstone, tuffaceous sandstone, and conglomerate with scattered layers of lignitic shale. This sequence is repeated many times throughout the lower part of the section. At the base of the section is a basal conglomerate con- taining abundant oysters, indicating a nearshore depositional environment. Occasional well-worn specimens of Miogypsina cushmani can be found in some of the upper layers of sandstone. These were likely derived from nearby outcrops of the Culebra Formation that had been uplifted by tec- tonic activity prior to the deposition of this part of the Cucaracha Formation. The dominant plant is a coarse grass, Saccha- rum spontaneum L., introduced from the Old World tropics and now widespread throughout the Canal area. On fresh exposures this grass initially aligns itself along the lignite outcrops (Fig. 8) and is a convenient marker for these organic-rich sediments. Later, as the grass spreads and slumping of the sediments occurs, this re- lationship is obscured. Along road K-2 about 0.8 km northwest of the intersection with K-15 in the Gaillard Cut sec- tion of the Canal is a roadside exposure of the Cucaracha Formation (our locality C, samples 57a—66, Figs. 1, 2, 7, 8). Across the road on the Canal side the ash flow continues and the lower 1.5 m includes highly decomposed logs repre- senting the forest layer at this site. Below the tuff is another clay shale, then a 0.7 m lignite near water level that is not evident on the roadside section. This lignite rests on a clay shale with 25 cm diameter siderite (FeCO,) concretions. At this locality there is a small utility building adjacent to a power pole (Fig. 7). About 4 m 1985] GRAHAM ET AL.—NEOTROPICAL PALEOBOTANY III 493 TABLE 2. Stratigraphy and lithology of samples from the Culebra (lower Miocene) Formation, Republic of Panama. Representative data selected from Panama Contractor's Hill locality, Gold Hill Investigation. Canal Company's Geological Field Log, Hole No. GH-9, Depth (ft.) Elevation (ft.) Descripton of Material Core Recovery —270.2 366.3 SANDSTONE, RH 1-2, moderately soft to medium hard, moderate strength, moderate mn very thin bedding, medium to coarse 7299.9 396.0 dark, grays. NOTE: E highly carbonaceous layers make innumerable dark gray lin SANDSTONE, RH 1-2, moderately soft to medium hard, moderate and carbonaceous, fossiliferous; color: 26.9 (90.5%) strength, moderate eae very thin bedding, medium to coarse an doi variably calcare ottled medium and par “wasa make innumerable dark gray lines SANDSTONE, RH 1-2, soft to medium hard, moderate strength, ith —353.3 449.4 grays. NOTE: thin, highly carbonaceous d carbonaceous, fossiliferous; color: 49.4 (92.5%) moderate jointing, very thin bedding; fine to medium grained wi very high silt content; variably calca careous, carbonaceous, and fos- siliferous; color: mottled medium and dark gray; thin carbonaceous layers m: —404.1 500.2 e innumerable dark gray lines. SANDSTONE, RH 1-2, soft to medium hard, moderate strength, with moderate jointing, very thin bedding; fine to medium graine very high silt siliferous; color: mottled medium and dark gra layers make innumerable dark gray lines. Bottom of hole. content; variably calcareous, carbonaceous, and fos- 50.0 (98.4%) ; thin carbonaceous above and to the left of the pole is a conspicuous conglomerate layer 1 m thick. The conglomerate terminates abruptly and continues on the other side of the pole about 4 m down in the section. This displacement is 9 m of left lateral motion, and the fault is one of the hundreds that char- acterize the Gaillard Cut section of the Canal. All samples were collected to the right of the fault line (facing the slope), and no faults were ob- served through the section sampled. out 0.7 km further to the right (NW) is part of the locality from which Whitmore and Stewart (1965) reported fossil mammal bones, all be- extended from Ca to Canal Station 2011 in the south. This site also contains large numbers of presumably crocodil- ian coprolites. Several of these were collected and processed, but the palynomorphs were poorly preserved and in low concentrations. In this dis- tance of ca. 0.7 km, two faults intervene, and, although the mammal beds are altitudinally equivalent to the layers sampled for fossil paly- nomorphs, they are stratigraphically lower. Clearly, sampling in this complex region must take into account the unusually large number of faults, frequently obscured by slumping and dense vegetation, that occur within very short dis- nces. A well was drilled near this site on October 4, 1958 (Hole No. PA-33; latitude 9°01'N + 5914.9, longitude 79°38’W + 2300.0; ground elevation 56 m (+ 182.7 ft.); core recovery 40.8 m (132.5 ft.; 71.4%); project—Cucaracha Reach Widening Studies; location—Cucaracha Reach). The drill- ing logs are filed with the Panama Canal Com- mission, and copies are in the palynology col- lections at Kent State University. Relevant portions of the log are summarized in Table 3. It is noted that only 71.4% of the core was re- covered, and among the missing strata are me and the lower described earlier. The initial lignite referred to in the bid is the uppermost layer in the roadside sectio ‘ae general sedimentology of the section, and the sequence of lit form under coastal, brackis where slow-flowing rivers empty onto a relative- ly broad coastal plain and where a dense vege- 494 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 FIGURES 7, 8. Exposures of the Cucaracha Formation (locality C) along the banks of the Canal, Republic of Panama. — 7. Alternating layers of dark lignites and lighter ash and conglomerates. —8. Lignite layer with 15— 20 cm thick overburden of id sh and lignite. Note the ee alignment of the grass Saccharum spontaneum L. along the lignite zon 1985] GRAHAM ET AL.—NEOTROPICAL PALEOBOTANY III 495 TABLE 3. Stratigraphy and lithology of samples from the Cucaracha (lower Miocene) Formation, Republic of Panama. Representative data selected from Panama Canal Company's Geological Field Log, Hole No. PA- 33, Cucaracha Reach locality Elevation (ft.) Depth (ft.) Description of Material Recovery +111.2 71.5 +109.5 73.2 +102.7 80.0 +85.0 97. (Canal water level) +78.7 104.0 +35.0 147. (Bottom of canal) +30.4 152.3 +28.0 154.7 2.9 185.6 LIGNITIC SHALE, RH- 1, soft, weak, closely jointed, thin bed- ; contains an abundance of carbonaceous plant debris; color: dark gray to black. (This is the uppermost lignite layer described from the roadside exposure from which fossil pol- len-bearing samples were collected.) LAY SHALE, RH- 1, soft, weak, very closely jointed with join highly slickensided; consists of a fine- water-laid is canic tuff highly altered to itontmorilinie and bentonitic clay minerals; color: medium gray-green. (Note that only 5396 of the core was recovered; missing is the conglomerate and basal lignite layer described from the roadside section.) ASH FLOW, RH-3, hard, strong, moderate jointing, massive bedding, fine-grained, porphyritic, agglomeratic; consists of a dacitic ignimbrite; color: purple, greenish gray, and brown. (This is the layer from which the K-Ar date was obtained; the lower 2 feet represent the old forest layer.) CLAY SHALE, RH- 1, soft, weak, very closely jointed and slick- ensided, thinly bedded; consists of a fine-grained volcanic tuff highly altered to montmorillonite and bentonitic clay min- erals; silt content increases with depth; color: gray-green. LIGNITIC SHALE, aed 1, Deiat, weak, closely jointed, thinly bedded, contains a f carbonaceous plant E color: dark gray to E (This is the layer on the Canal si of the road below the lowermost layer on the roadcut expo- sure.) CLAY SHALE, RH- 1, soft, weak, very closely jointed and slick- ensided, thinly bedded; consists of a fine-grained volcanic tuff highly altered to montmorillonite and bentonitic clay min- erals; variably silty and sandy; contains some calcareous con- cretions in upper portion; color: 157.4 to 165.6, gray-green, 165.6 to 172.9, gray-green mottled with red-brown, 172.9 to 184.4, red-brown, 184.4 to 185.6, gray-green mottled with red-brown. NOTE: Some scattered calcareous concretions in red-brown layer from 180.6 to 181.4 ft. depth (these are the siderite concretions discussed previously). Bottom of hole. 1.7 (100%) 6.8 (53%) 19.2 (80%) 48.3 (37%) 2.0 (83%) 28.6 (93%) Y and tation (frequently dominated by ma moderate off-shore currents allow Gai detri- tus to accumulate. Climatic conditions under which modern lignites are presently forming are warm-temperate, subtropical to tropical (Cohen & Spackman, 1972; Scholl, 1964a, 1964b). The shoreline was periodically inundated, probably from local subsidence of the land rather than changes in sea level, as evidenced by the folding and faulting of the strata. The clays and Hs contain both marine invertebrates and debri from land vegetation. Extensive volcanism is documented by the tuffs (water-lain volcanic ash) and the basalt that caps the section. THE LA BocA FORMATION The La Boca Formation outcrops along both sides of the Canal from the Pacific entrance to Las Cascadas Reach. It is composed of mud- stones, siltstones, sandstones, lignited shales, tuffs, and limestones. The Emperador limestone is a ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FiGuURE 9. Aerial photograph of pa Cascadas Reach of the Canal, Republic of Panama, 1967, when this 5m section was being widened from 90 m A in the early Miocene Boca Formation at canal marker 1600 (1766 new system). Double arrow designates drilling site LBW-88 through the La Boca Formation at canal marker 1619 (1785 new system; 9*04'N, 79°40'W; log data summarized in Table 5). Triple arrow designates collecting locality B in the La Boca Formation at canal marker 1622 (1788 new system). Other samples were collected at markers 1625 (1791) and 1627 (1793) further to the right. member of the lower part of the La Boca com- posed of coralliferous limestone reflecting warm, subtropical to tropical marine environments. These conditions are consistent with the layers of lignite that occur throughout the lower part of the section (see previous discussion, Cucaracha Formation). The sequence of sediments indicates a period ofsubsidence with bathyl siltstones (Bla- cut & Kleinpell, 1969) capping the section. The lignitic layers contain shells of marine inverte- brates and substantial amounts of pyrite result- ing from the oxidation of soluble ferrous com- pounds by iron bacteria. Woodring (1982) recognized 130 species of mollusks from the La Boca and their collective ranges suggest an early Miocene age. In September 1967, samples were collected from two Canal-side exposures of t Boca Formation (Fig. 2). Our locality A qus 1- 6) was on the Las Cascadas Reach at station marker 1600 (1766 new stationing), and locality B (samples 27-54) was at markers 1622 (1788), 1625 (1791), and 1627 (1793). At that time the Canal was being widened from 90 m to 155 m (Figs. 9-11), and by the January 1983 field season this construction was complete. Thus, compared to the present physiography, the 1967 samples were collected from strata 25 m up and 60 m out over the Canal. The beds dipped back into the slope 20-25? so that now the same lignite layers are nearly at water level and, in this area, near the La Boca-Las Cascadas contact (Fig. 12). In addition to these challenging alterations in the landscape, the Canal marking system has changed. Marker number 1625 (in our 1967 col- lections) is now 1791. This complicates any study dealing with the former Canal Zone and vicinity where canal markers were used as reference points. The Panama Canal Commission has a list correlating the new and old numbers, and 1985] GRAHAM ET AL.—NEOTROPICAL PALEOBOTANY III 497 Ficures 10, 11. Exposures of the La Boca Formation at locality A, Las Cascadas Reach of the Canal, Republic of Panama, in 1967 during widening of the Canal [near canal marker 1600 (1766 new w system); see Figure 9]. — 10. General view of the locality. —11. La Boca Formation. Note the fault abruptly terminating the white tuff at the lower left of the exposure. Over 113 of these faults have been mapped in a distance of 3 km in the Gaillard Cut section of the Canal. 498 TABLE 4. List correlating the old iin site numbers to the new collecting site numbers. F Panama Canal Commission. 1967 Stationing Present Stationing 1600 1766 1619 1785 1620 1786 1622 1788 1625 1791 1627 1793 those pertaining to our collecting sites are sum- marized in Table 4 It should also be noted that reference points may be cited on North American datum or Canal Zone datum. To change North American to Ca- nal Zone datum, add 941 ft. to longitude and 942 longitudes cited in this study are on North Amer- ican datum ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 On April 6, 1960, a well was completed at canal marker 1619 (old system; 1785 new sta- tioning) near our collecting sites. The log pro- vides general information on the stratigraphy and lithology of the La Boca Formation [Hole No. LBW-88; latitude 9?04'N + 4586.4, longitude 79°40'W + 4557.1; ground elevation 52.7 m (+ 171.4 ft.); core recovery 38 m (123.5 ft.; 72.0%); project — Canal Improvement; location — Las Cascadas Reach; old station 1619, 1785 new sta- tioning]. The logs are filed with the Engineering and Construction Bureau of the Panama Canal Commission, and copies are in the palynology collections at Kent State University. The rele- vant portions of this log are summarized in Ta- ble 5. It is evident that the Culebra, Cucaracha, and La Boca Formations are similar in age (lower Miocene), and that dienen relations are complicated by the highly faulted and altered nature of the sediments along ‘the Gai lard Min nothe om an aaa coring error in the 1950s s led to incorrect correlations in some major paleon- TABLE 5. Stratigraphy and lithology of samples from the La Boca (lower Miocene) Formation, Republic of Panama. Representative data selected from Panama Canal Company's Geological Field Log, Hole No. LBW- 88, Las Cascadas Reach locality (station 1619). Elevation Depth Co (ft.) (ft.) Description of Material Recovery +48.0 123.4 LIGNITIC SHALE, soft rock, closely jointed, joints slickensided, thin bed- s contains abundant carbonaceous plant debris and some marine bor- 1.5 ings filled with fossiliferous material; color: dark gray to black. (5796) +45.9 126.0 SILTSTONE, soft rock, thin bedding, moderate jointing with joints slick- 3.5 ensided, fossiliferous at base; color: medium gray-gree (77%) +40.9 130.5 LIGNITIC SHALE, soft rock, closely jointed, joints slickensided, thin bed- ding; contains abundant carbonaceous plant debris and some marine bor- 0.0 ings filled with fossiliferous material; color: dark gray to black. (0%) +38.4 133.0 SILTSTONE, soft rock, thin bedding, moderate jointing with joints slick- 0.9 ensided, fossiliferous at base; color: medium gray-green. (45%) +36.4 135.0 | LIGNITIC SHALE, soft rock, closely jointed, joints slickensided, thin bed- ding; contains abundant carbonaceous plant debris and some marine bor- 0.5 ings filled with fossiliferous material; color: dark gray to black. (16%) +33.4 138.0 s tuffaceous, SOR rock, fugere jointing with joints open; assive bedding; consists of andesitic and basaltic pebbles and cobbles in a fine-grained, tuffaceous matrix of similar composition; some fragments are pumaceous; somewhat altered to various clay minerals and chlorite; material checks rapidly on exposure to air and falls apart; color: mottled grays and light greens. NOTE: 25.9 Very tuffaceous in lower 11.0 ft., almost a tuff. (7796) —0.1 171.5 Bottom of hole. 1985] GRAHAM ET AL.—NEOTROPICAL PALEOBOTANY III 499 o3 wes tt. C 12. New exposures of lower Miocene La Boca lignites (January 1983), Le of Panama 967 was completed (see Figs. 9-11). This is locality B, and when FIGURES 12, 13. m out over the Canal. Since the beds dip after 61.5 m widening project started in 1 topie 2 Pis right (nort hough beds are “watami through uplift and left-lateral faulting. Slumping has obscured lithological 9). on top, e difftrences io the formations at the contact (canal marker 19 500 TABLE 6. Location of pollen-bearing samples from the upper Miocene and Pliocene Gatun Formation and other Cenozoic deposits, Republic of Panama. Cores are designated by the Panama Canal Commission's system, and samples from each core are listed accord- ing to depth in feet (except HU's SL-60 sample at 40 ft. which is at a depth of ca. 127 ft.). Samples were collected in December 1962, by Elso Barghoorn and Alexandra Bartlett (HU), or in December 1963, by A. KSU). All slides are presently in the palynology collections at KSU. Core SL-48 (Latitude 9°11’ + 1331; Longitude 79°55’ + 4501) KSU HU 157 Gatuncillo 158 162 162 Core SL-49 (Latitude 9°08' + 4113; Longitude 79*57' + 1987) Gatun Core SL-60 (Latitude 9?11' + 6042; Longitude 79°52' + 411) KSU HU 40 Caimito Formation Core SL-103 (Latitude 9°16’ + 5945; Longitude 79°52' + 2963) KSU HU 178 Quaternary 250 Gatun 253 255.5 257 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 tological publications dealing with the Culebra and La Boca Formations. Van den Bold (1972, 1973) correlated the La Boca with the Culebra bee on _ data and similarities in li- Ogy. correlation was made by egi Mos que mollusk evidence. R. H. Stewart (pers. comm.) found field evidence that was inconsistent with this correlation (see below) and re-examined the core from which the inver- tebrate fossils had been obtained. Stewart dis- and worked loose in the next drilling which was through the La Boca. Woodring (1982) subse- quently corrected the error for the mollusk stud- ies. Field stratigraphy reveals that the three for- mations are sequential within the early Miocene from the Culebra (oldest) through the Cucaracha, to the La Boca (youngest). This stratigraphy, used on the most recent version of the geologic map of Panama (Stewart et al., 1980), is based on the observation that in local areas the Cucaracha Formation sits on top of the Culebra. Above the Cucaracha is another formation, the Pedro Miguel. The terrestrial phase of the La Boca has not been observed directly on the Cucaracha, but it does interfinger with the Pedro Miguel, and therefore must be younger than the Cucaracha. In addition, the marine phase of the La Boca does rest directly on the Las Cascadas Formation (Fig. 13) which is the same radiometric age as the Cucaracha Formation. Thus, the Culebra, Cucaracha, and La Boca Formations, as pres- ently understood, represent a time-stratigraphic sequence for the early Miocene in central Pan- Summary of log data for four cores through Tertiary and Quaternary formations in Panama from which fossil pollen and spores were recovered. Depth and elevation are in feet. See Table 6, Figure 1 (locality 2), and text for location of drilling sites. Core Elevation Depth Description of Material SL-48 +86.5 0.0 Water +6.5 80.0 Overburden — 64.9 151.4 Top of weathered rock (samples SL-48, 157, 158, 162) SILTSTONE, medium hard to hard (OH 3-4); slightly to medium plastic; medium tough to tough; medium to high dry strength; clayey with streaks weathered to clay; slightly sandy, with very fine sand; color: mottled, gray and brown. (Gatuncillo Formation.) Recovery 10.4 ft —76.6 163.1 Top of sound rock SILTSTONE, medium hard (RH-2); massively jointed, no joints apparent; core in 0.1 ft. and smaller pieces broken during drilling by dry blocking; 1985] GRAHAM ET AL.—NEOTROPICAL PALEOBOTANY III 501 TABLE 7. Continued. Core Elevation Depth Description of Material —77.8 SL-49 +86.9 +14.9 — 133.2 SL-60 +87 —40.3 —60.0 —99.3 SL-103 +85.7 +47.7 +17.4 —163.3 —168.4 —178.4 225.8 0.0 81.0 88.0 91.7 127.3 147.0 186.3 0.0 38.0 68.3 249.0 254.1 264.1 slightly sandy with fine sand; fairly clayey and marly; massively bedded, no bedding apparent; color: dark gray to gray. (Gatuncillo Formation.) Recovery 1.2 ft. Final depth Water Overburden Top of sound rock (samples SL-49, 222.5, 223, 223.5 SILTSTONE, soft to medium hard (RH 1-2); massively jointed, no joints apparent; core in one piece 4.6 ft. long and in 0.1 ft. and smaller pieces, broken during drilling by dry blocking; slightly sandy with fine sand; fairly clayey; quite marly with scattered white streaks of shell fragments; mas- sively bedded, no bedding apparent; friable; color: dark gray. (Gatun For- mation.) Recovery 4.7 ft. Final depth Water Overburden SILT, CLAY, AND ORGANIC MATERIAL, laid down since formation of Gatun ; GRAVEL AND SILT. SILT AND CLAY, soft to medium soft (OH 1-2); loose; medium plastic; slightly spongy at plastic limit, nis belay plastic limit; very: weak; low dry strength; sticky, | wood fragments; color: dark gray and black. (Atlantic Muck.) Recovery 32.2 ft. Top of weathered rock (sample SL-60, 40) SANDSTONE, medium hard to very hard (OH 3-5); jointing apparent at base only; silty; bedding not apparent; color: gray-green and brown (Weathered Caimito Formation.) Recovery 9.8 ft. Top of sound rock SANDSTONE, medium hard to hard (RH 2-3); closely jointed; core re- covered i in poor condition in 0.01-0.4 ft. lengths with 0.1 ft. lengths and silty, with occas sional calcareous lenses near -the base; bedding d about 5°; color: light gray-green with calcareous lenses almost white. (Caimito Formation.) Re- covery 6.18 ft. Final depth Water Fill Overburden (sample SL-103, 178) Top of weathered rock (samples SL-103, 250, 253) SANDSTONE, hard (OH-5 to RH-1), silty; locally clayey; scattered streaks of light blue-gray clay; slightly waxy locally; checks upon drying; color: dark gray; grades into underlying sound rock. (Weathered Gatun For- mation.) Top of sound rock (samples SL-103, 255.5, 257) SANDSTONE, soft to medium hard (RH 1-2); massively jointed; medium grained; silty; carbonaceous; fossiliferous; contains scattered small pyrite grains; dark gray and greenish gray; core recovered in 0.5-2 ft. lengths. (Gatun Formation.) Recovery 10 ft. Final depth 502 ama. These strata provide a rare opportunity in tropical paleobotany to study the vegetational history of a single region through a defined seg- ment of Tertiary time. THE GATUN FORMATION The Gatun Formation was named by Howe (1907: 113-114) from the type region extending from the bluff overlooking Gatun Lake near Ga- tun, northward to Mount Hope, formerly known as Monkey Hill, near Colon (Woodring & Thompson, 1949). Surface outcrops are exten- sive on the north (Caribbean) side of the Canal bordering the northern shore of Lake Gatun. The rocks are sandstone, siltstone, tuff, and conglom- erates typical of nearshore deposition. The Ga- tun Formation is relatively unfaulted, whereas the nearby Eocene, Oligocene, and lower Mio- cene formations are heavily faulted. Thus, the stratigraphic position and geomorphology indi- cate a post lower Miocene age, although the Ga- mation does rest with a strong, angular uncon- formity on the pre-Tertiary, Cretaceous volca- nics and sediments. There are ash flows or pumice beds within the Gatun sediments that may even- tually provide radiometric dates. The upper part of the Gatun Formation is dis- tinctly marine. Foraminifera from offshore, pre- sumed Gatun sediments (DSDP, R. H. Stewart, pers. comm.), contain some Pliocene species, and Vokes (1983) believes, on the basis of other in- vertebrates, that the Gatun could be as young as basal Pliocene. Near the Refineria Panama, Co- lon, a small knoll is composed almost exclusively of shells of marine invertebrates, especially Tur- ) of 339 species have been described, and collec- tively they were interpreted to suggest a middle Miocene age. This assignment is used on the most recent version of the geologic map of Panama (Stewart et al., 1980). More recently, however, R. H. Stewart (pers. comm.) believes that a reas- sessment of the paleontological evidence, com- bined with observations on the stratigraphy, sug- gests that the Gatun Formation may be upper Miocene and Pliocene in age. A preliminary sur- vey of the pollen and spores reveals an assem- blage quite modern in aspect, and, for purposes ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 of our studies, we are now tentatively accepting the younger age assignment. In 1962 the Panama Canal Commission drilled a series of wells in Gatun Lake to test the foun- dation for new water stora t as part of the Trinidad Dam studies, The upper portions th gla cial sediments and were e used by Bartlett and Barghoorn (1973) for their study of Quaternary vegetational history and sea-level changes in ama. The lower portions of the cores that penetrated surface Gatun Formation sediments also contained a rich pollen and spore flora. It is fortunate that samples from the cores were ob- tained because all surface exposures are marine and none are known to contain plant microfos- sils. Samples containing fossil pollen and spores are listed in Table 6, and the location of the drilling site is indicated as locality 2 on Figure l. Relevant log data from the cores are sum- marized in Table 7 In the preceding sections the age, stratigraphic relationships, lithology, and depositional envi- ronments are summarized for five pollen/ spore-bearing Tertiary formations in Panama. Attention is devoted to the original location of collecting sites; alteration in the landscape that m and datum reference points that can confuse comparison with previ- ously published studies; noting errors in earlier correlations; and recording the permanent lo- cation of unpublished well logs, samples, pro- cessed material, and slides in the palynology col- lections at Kent State University. The rationale for providing these details in an introductory paper is three-fold. First, it is con- venient to integrate geological information on the several formations in a single publication. Second, it is possible that these lignite beds will be further altered or destroyed through construc- tion activities along the Canal, or rendered no longer accessible because of political changes af- fecting Central America. For example, the core material used by Bartlett and Barghoorn (1973) for their study of Quaternary vegetational history in Panama, previously housed in storage facili- ties of the Panama Canal Commission, has been destroyed. The Tertiary portions of these cores, including all core material mentioned in this pa- per, measuring hundreds of thousands of feet and containing vast amounts of irretrievable infor- mation on the biology and geology of Panama, 1985] were also discarded. Fortunately, samples were obtained in 1964 and 1967, and these constitute a major part of the present study. Finally, indi- viduals now in residence in Panama (R. H. Ste art and J. L. Stewart) and other life-long iden of Central American paleontology (e.g., P. Woodring) will soon be altering these direct as- sociations with Panama. Their cooperation in facilitating almost every recent paleontological investigation within the Canal Zone is widely recognized, together with their impressive knowledge of local field geology and the exten- sive unpublished literature. Consequently, it seems timely to initiate these studies on the Ter- tiary history of vegetation in Panama with a re- vised summary of the geology of the formations. LITERATURE CITED BARTLETT, A. S. & E. S. BARGHOORN. 1973. Phyto- geographic history of the Isthmus of Panama dur- nd V History of Northern Latin America. Elsevier Publ. Co., Amsterdam BLAcuT, G. & R. M. KLEINPELL. 1969. Ast uad il sequence of benthonic smaller foraminifera fro Canal the La Boca Formation, Panama ne. Contr. Cushman Found. Foram. Res. 20: 1-22. COHEN, A W. SPACKMAN. 1972. Methods in peat petrology and their application to reconstruc- tion of oo s. Bull. Geol. Soc. Amer 83: 129-142. Corr, W. S. 1952. Eocene and Oligocene larger fo- raminifera from the Panama Canal Zone and vi- cinity. Profess. Pa . Geol. Surv. 244: 1-41. Coney, P. J. 1982 [1983]. Plate tectonic constraints on the biogeography of Middle America and the be . Ann. Missouri Bot. Gard. 69: 432-443. Croat, T. B. 1978. Flora of Barro Colorado Island. Stanfo pap Press, Stanford. 1982 [1983]. Neotropical floristic di- GoMEz-POMPA, A. 1973. Ecology of the vegetation of Veracruz. Pp. 73-148 in A. Graham (editor), Vegetation and Vegetational History of Northern Latin America. Elsevier Publ. Co., Amsterdam. GRAHAM ET AL.—NEOTROPICAL PALEOBOTANY III 503 GRAHAM, ALAN. 1976. Studies in neotropical paleo- botany. II. The Miocene communities of Vera- cruz, Mexico Ann. Missouri Bot. Gard. 63: 787- 842. D. M. JARZEN. 1969. Studiesin ipi Wer D I. The Oligocene communities of Puerto Rico. Ann. Missouri Bot. Gard. 56: 308- 357. Hit, R. T. 1898. The pnr history of the Isth- of Costa Ri Mus. Comp. Zool., H ll . 1973. The vegetation of the Antilles. n A. Graham Ste li Vegetation and rn Latin America. HowE, E. 7. Report on im E of the Canal Zone. Annual Rep. Isthmian Canal Comm. 1907: 108-138. Raven, P. H. & D. I. AxELROD. 1974. Angiosperm biogeography and past continental movements. Ann. Missouri Bot. Gard. 61: 539-673. 964a. Recent vun record in e ` swamps s and rise in sea level over the southwestern coast of Florida, Part IL Marine Geol. 1: eos 1964b. aa M record in man- grove swamps a sea level over the south- western coast of f Florida, Part II. Marine Geol. 2: 343-3 iine R. . & J. L. STEWART (with the collabo- n of S 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 (map also included in Wood- VAN DEN Botp, W. A. 1972. Ostracoda of the La rmation, doc Canal Zone. Micropa- ( deonology 18: 410-44 1973. La d estratigráfica de la for- ación oca, Panama, Zona del Canal. Publ. Geol. ICAITI 4: 167-170. Mrs E. H. 1983. Additions to the Typhinae (Gas- poda: Muricidae) ofthe Gatun Formation, Pan- ama. Tulane Stud. Geol. Paleontol. 17: 123-134. WHITMORE, F. C. & R. H. Stewart. 1965. Miocene d Central A i S 148: 180-185. WoopRING, W. P. 1957-1982. Geology and paleon- tology of Canal Zone and — parts of Pan- ama. Profess. Pap. U. eol. Surv. 306A-F. & T. F. THOMPSON. 1949. Tertiary forma- tions of Panama Canal Zone and adjoining parts of Panama. Bull. Amer. Assoc. Petrol. Geol. 33 223-247 STUDIES IN NEOTROPICAL PALEOBOTANY. IV. THE EOCENE COMMUNITIES OF PANAMA! ALAN GRAHAM? ABSTRACT The Pa to upper Eocene Gatuncillo Formation outcrops near pense Díaz, Panama. Fro n the formation an assemblage of fossil pollen and spores ype cf. f. ragastris, Combretum/Terminalia, Casearia, Lisianthius, cf. Tontalea, Alfar Engelhardia (rare), Crudia, Malpighiaceae, cf. Ficus, Eugenia/Myrcia, Coccoloba, Rhi ra, F. e ubiaceae, Cardiospermum, Serjania, Paullinia, cf. Chrysophyllum, Pelliceria, and phi tal conditions in the immediate iced of the Gatuncillo ment of re re Edi paleoclimates and paleophysiography must await dis- covery of other fossil oras, bee the Gatuncillo assemblage is the only one of Eocene age known for northern Latin Am The vegetation characterizing the Eocene ep- kn published on an extensive flora of this age for Central America, Mexico, or the Antilles. Con- sequently, the Gatuncillo palynoflora of Panama is of interest in providing a first view of this vegetation. The nearest Eocene fossil floras to the south are from the upper Los Cuervos and Mirador Formations of Colombia (Gonzalez Guzman, 1967) and the Rio Turbio Formation of Argentina (Romero, 1977; for a more com- plete bibliography see Graham, 1973a, 1979a, 1982). The former are of early and middle Eocene age, compared to the late Eocene Gatuncillo pal- ynoflora, and the latter presently treats only the gymnosperms and the Fagaceae. Both belong to a completely different paleophysiographic prov- ince because at this time South America was iso- lated between Africa and North America (e.g., Raven & Axelrod, 1974), while the present re- gion of Central America was occupied by a series of volcanic islands trailing south from the North American continent (Coney, 1982; Dengo, 1973). To the north the closest Eocene floras are from the Mississippi Embayment and Gulf coast re- gion studied by Dilcher and collaborators (e.g., Dilcher, 1973), Elsik (1974), Frederiksen (1980), and others. These are some 2,500 km from the Panama localities. Additional information is provided by Germeraad et al. (1968), Muller (1981), and Graham (1977), but these are either preliminary reports or deal with the fossil record of specific palynomorphs rather than entire paly- ofloras. The results presented here are consid- ered an initial effort to characterize the Eocene vegetation of central Panama and must be aug- mented considerably before a detailed concept of the regional paleocommunities, paleoenviron- ments, and vegetational history can be devel- oped. THE COLLECTING LOCALITY Samples were collected from a roadside ex- posure of the Gatuncillo Formation near Alcalde Diaz (Pefioncito). The area is shown on the Al- calde Diaz quadrangle map (sheet 4243 II NE, ' The author gratefully acknowledges information on the modern vegetation provided by Thomas B. Croat and Alwyn H. Gentry. Research supported by NSF grants DEB-8205926, DEB-8007312, GB-11862. ? Department of Biological Sciences, Kent State University, Kent, Ohio 442 ANN. Missounmi Bor. GARD. 72: 504—534. 1985. 1985] | SILTSTONE E LIGNITE s. aes NOE MERC ute : Dij GRAHAM -NEOTROPICAL PALEOBOTANY IV u DIRT ROAD e Tomus Ba em . 505 jer foa 3 FT. CA 2 FT. THICK €" 9 o: ha Yet ua E t AS eO - —À — as. — Diagrammatic section through roadcut exposure of the Gatuncillo Formation near Alcalde Díaz, IGU i Panama (see Graham et al., Series E866, scale 1:25,000) prepared by the Army Map Service, Corps of Engineers at coordinates 9°7'N, 79°32'W (North American datum). The site is reached from Panama City by the Boyd- Roosevelt Highway. About 1.8 km from Las Cumbres a dirt road departs to the right. The locality is along this road 2.2 km from the high- way intersection. This site is designated as our locality D and in January 1967, 15 samples were collected (Fig. 1). The beds are tilted at an angle of about 30°. Samples 1-9 are from a basal lignite ca. 0.6 m (2 ft.) thick. This layer is overlain successively by a 0.9 m (3 ft.) siltstone, a second lignite 1.2 m (4 ft.) in thickness (samples 10-12), a 0.6 m (2 ft.) siltstone, a 150 cm (6 in.) lignite (sample 13), a 1.2 m (4 ft.) siltstone with a thin lignite lens (sample 14), and a 1.2 m (4 ft.) siltstone (sample 15) separated from the underlying silt- stone by a thin erosion surface. The section is capped by early to late Oligocene conglomerates of the Panama Formation. A visit to the locality in January 1983 revealed that considerable slumping had taken place and the exposure was covered by a nearly uniform veneer of silt 300- 460 cm (12-18 in.) thick with a dense over- growth of herbaceous vegetation (mainly grasses) beginning to develop. 1985 for discussion of the locality). From among the samples, numbers 2, 4, and 9 (all from the lower lignite) yielded the most well-preserved and abundant palynomorphs, and the present study is based primarily on these samples. Slides from the other lignites and the uppermost siltstone were also examined, how- ever, and contained essentially the same micro- flora. Thus the data base for Hus study i is some- what limited t from the lower lignite. Woodring (1957-1982) has studied the mol- lusks of the Gatuncillo Formation, and Cole (1952) studied the larger foraminifera. On the basis of the stratigraphic distribution of these fossils in other Caribbean deposits, both authors conclude the Gatuncillo is middle(?) to late Eocene in age. The possibility ofa middle Eocene age is considered because three of the larger fo- raminifera are known elsewhere in strata of this age, and two (Yaberinella jamaicensis and Fa- biania cubensis) are known only from the middle Eocene. In addition to the lignites and mudstones/silt- stones, the Gatuncillo in surrounding areas in- cludes quartz sandstones and coralline and fo- raminiferal limestones. This lithology reflects deposition in a nearshore coastal environment, and the lignites specifically indicate warm-tem- 506 TABLE 1. ANNALS OF THE MISSOURI BOTANICAL GARDEN Formation, Panama. Figures are percentages based on counts of [Vor. 72 Identification and numerical representation of fossil palynomorphs from the Eocene Gatuncillo 00. Sample Sample Sample 2 4 9 Sample Sample Sample 2 4 9 Selaginellaceae Selaginella Parkiaceae Ceratopteris Polypodiaceae Pteris Trilete Fern Spores Bromeliaceae cf. Tillandsia Type 1 cf. Tillandsia Type 2 Palmae Anacardiaceae cf. Campnosperma Aquifoliaceae Ilex cf. Araliaceae Bignoniaceae cf. Paragonia/Arrabidaea Burseraceae cf. : 1m cf. P. a. Combretum/Terminalia Flacourtiaceae Casearia Gentianaceae Lisianthius Hippocrateaceae cf. Tontalea Juglanda yee E DU 2.5 - 0.5 — 2 = 2.5 0.5 4 1.5 5 1.5 1 = 0.5 a 0.5 l = 2 05 0.5 4 2 3 1.5 — l = 1 ú 6 - l — 3 0.5 2.5 Leguminosae/Caesalpinioideae Crudia Malpighiaceae Type 1 Type 2 Moraceae cf. Ficus Myrtaceae Eugenia/ Myrcia Polygonaceae Coccoloba Rhizophoraceae Rhizophora Rubiaceae Sapindaceae C. ardiospermum cf. Chrysophyllum Theaceae Pelliceria Tiliaceae Mortoniodendron Unknown Echinate Types Unknown Intectate Type Type 6 Unknown Triangular Type Type 7 = -— Unknown Oblate/Oblate-Spheroidal Types 0.5 0.5 3.5 NEOTROPICAL PALEOBOTANY IV 507 1985] GRAHAM- TABLE l. Continued. Sample Sample Sample 2 4 9 Type 17 1 1.5 1 Type 18 2.5 — Type 19 1 — — Type 20 — — — Type 21 — Unknown Prolate, Tricol(por)ate Types Type 22 3.5 Type 23 1.5 — Type 24 3.5 1.5 — Type 25 — 2 — Type 26 5.5 3 — Type 27 3 3 2 Type 28 4 2.5 0.5 Type 29 — — — Type 30 6.5 4 2 Type 31 5.5 0.5 1 Type 32 — 0.5 — Type 33 0.5 — Type 34 0.5 0.5 ype 35 — 2 — Type 36 2.5 2 — Type 37 1.5 0.5 Type 38 — — 1 Type 39 — — — Type 40 — Other Unknowns 14 11.5 17.5 perate to tropical conditions (Cohen & Spack- man, 1972; Scholl, 1964a, 1964b). Other details on the geology of the Gatuncillo Formation are summarized by Graham et al. (1985). MATERIALS AND METHODS Samples were prepared by cleaning the sur- faces to remove contaminants and palyno- morphs damaged by oxidation. Pieces of lignite between two and three grams were broken in a mortar and pestle and covered with 10% HCl to remove carbonates. The samples were then passed through HF (to remove silicates), nitric acid (to oxidize lignins and other organic debris), and 10% KOH. Residues were mounted unstained in glycerine jelly and sealed with CoverBond. Pho- tographs were taken with a Zeiss photomicro- scope using Panatomic X film. Location of paly- nomorphs on the slides is indicated by England Slide Finder coordinates. All materials from the study are deposited in the palynology collections at Kent State University. SYSTEMATICS A total of 47 palynomorphs were identified from the Gatuncillo Formation, and 40 other morphological types were recovered whose bi- ological affinities are unknown (Table 1). The prefix cf. designates the identification is proba- ble, but not as certain as those referred to specific taxa. This is usually due to minor quantitative differences between the modern and fossil pollen or spore that could be the consequence of pres- ervation, or reflect the age difference between the Eocene specimens and their presumed modern analogs. In other instances the specimens may be similar both to a common neotropical genus, and to a taxon from some remote geographic region, characteristic of a completely different biozone (e.g., northern boreal), a rare species, or a plant with exceptionally low pollen production and a highly specific entomophilous pollinating mechanism. Although it may be likely the spec- imens represent the pollen-prolific, common neotropical genus, it is theoretically possible they represent the other form. The cf. designation is also useful in providing examples of morpho- logically similar pollen to augment the discrip- tion and illustration of the fossil specimen (for a more specific example of identification pro- cedures see cf. Tillandsia Type Identification of the microfossils is based on comparisons with taxa in a modern pollen and spore reference collection numbering about 22,000 slides (ca. 20,000 species). To minimize bias in the identifications towards present-day Panamanian or Central American vegetation, our preparations include species from throughout Latin America (Mexico, the Antilles, Central and orld en a and published sod of sensi paleopalynologcl studies on Cenozoic deposits in various regions, are also consulted. Consequently, the possibility of bias in the identification of the microfossils towards a given geographic region or a particular community is recognized and is not regarded as a major problem considering the broad coverage of the reference collection and the available lit- below, lpt In the pollen d ipti terns are described a: as seen under light micro- scope (LM) magnifications of about 430-970 x. In some instances a sculpture pattern appearing as faintly scabrate under LM may be revealed as [Vor. 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1985] punctate or microreticulate under electron mi- croscope (EM) magnifications. In such instances the EM pattern can be given in parentheses; e.g., faintly scabrate (EM — microreticulate). RESULTS SELAGINELLACEAE Selaginella (Fig. 2a, b). Amb ca. circular; tri- lete, laesurae frequently obscured by wall thick- ness and sculpture elements, straight, narrow, inner margin entire, ca. 12-15 um long, extend- ing nearly to spore margin, tapering to acute apex; echinate, echinae short (ca. 2-3 um), occasionally curved, base broad, distal face appearing irreg- ularly reticulate due to arrangement of spine bas- es, proximal face less sculptured; wall 2-3 um thick (excluding echinae); 25-30 um and 40-50 um. Two size groups of Selaginella microspores are present, 25-30 um (Fig. 2b) and 40-50 um (Fig. 2a). Both are moderately frequent in samples 2 cene of Puerto Rico (Graham & Jarzen, 1969) and in the Miocene of Veracruz, Mexico (Gra- ham, 1976). Selaginella is presently distributed widely in tropical regions and is particularly abundant in shaded humid habitats. PARKIACEAE Ceratopteris (Fig. 3. Amb oval-triangular, (2 um) ridged lip, lip surface psilate, margin en- tire; wall coarsely and conspicuously striate, striae psilate, margin slightly undulahog, S 2-3 um wide, ss dis- tinct approaching laesurae: wall 2-3 un E 55-80 u The genus Ceratopteris is a floating aquatic fern distributed from Florida through the An- GRAHAM —NEOTROPICAL PALEOBOTANY IV 509 tilles and Central America into northern South America. The spores are common in the fossil record under the form-genus name Cicatricosis- porites. Germeraad et al. (1968) described sim- ilar spores as Magnastriatites, but Dueñas (1980) considered these congeneric with Cicatricosis- porites. In the Gatuncillo samples the spores are common in sample POLYPODIACEAE Pteris (Fig. 4). Amb triangular, margin en- tire; trilete, laesurae straight, narrow, inner mar- gin entire, ca. 20 um long, extending nearly to spore margin; wall with coarse, irregular verrucae on distal surface, proximal surface more laevi- gate, marginal flange ca. hyaline, 4-5 um wide; 45-50 um The genus Pteris includes about 250 species widely distributed in Latin America. Although some species can occupy dry habitats as primary invaders, most grow in moist areas (in Panama, tropical moist forest; Croat, 1978). TRILETE FERN SPORES (FiGS. 9, 10, 13-16) In addition to Ceratopteris and Pteris, five oth- er trilete spore types are represented in the Ga- tuncillo Formation. Although affinities may be suggested for some of these spores, their mor- phology is too general for definite generic iden- tification. Type 1 (Fig. 10). Amb triangular; trilete, lae- surae slightly undulating, narrow, inner margin entire, 20-25 um long, extending nearly to spore margin; wall 2 um thick, laevigate; 35-40 um This spore is similar to several species of Adiantum, but lacks diagnostic features separat- ing it consistently from others (e.g., variants in spores of some species of Dennstaedtia). It is most common in sample 2 (2%). Type 2 (Fig. 9). This spore differs from Type 1 in that the wall thickens at the apices and the ends of the laesurae are branched. 45 um. Type 3 (Fig. 13). The distinctive feature of this spore is its large size (120 um), nearly double — FIGURES 2-15. 2a. qucd. 2-1, V-27,1-3.—2b. Selaginella, 2-1, Q-31,2.—3. Ceratopteris, 9-1, S-36.— -39. Monolete fern Le type 1, 4-1, O-22,2. —6. Monolete fern spore type 2, 2-1 . G-39, 2.— š —8. Monolete fern Leg: type 4, 2-1, T-32,1-3.—9. Trilete fern spore —10. Trilete fern iion type 1, 4-1, X-8.— 13. Trilete E spore type 3, 4-1, N-26,3-4.-14, 15. Trilete Heins spore type 4, 4-1, M-38,1-2. (Numbers fo llo , 12. Monolete fern spore type 5, 4-1, W-12,4.— owing plant names indicate the sample and slide number. and England Slide Finder coordinates. All material is labeled locality Pan D, = Gatuncillo Formation from the site described in the text.) 510 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 1985] that of the similar Type 5 (55-65 um; Fig. 16). It is most common in sample 2 (2.596). Type 4 (Figs. 14, 15). Amb circular; trilete, laesurae straight, narrow, 27-30 um long, ex- tending !2—/ distance to spore margin, bordered by narrow, ridged lip; wall hyaline, 3-4 um thick, laevigate, minute pits surrounding laesurae, con- centrated in central contact region; 90-100 um. Type 5 (Fig. 16). This spore is similar to Type 3 except it is about half the size (55-65 um). Similar spores are known in older sediments as Deltoidospora and have been reported recently rom the Eocene of the Burgos Basin (north- eastern Mexico; Martinez-Hernandez et al., 1980). Among spores in our reference collection it is similar to Antrophytum (Polypodiaceae). It is common in samples 2 (4%) and 4 (1.5%). MONOLETE FERN SPORES (FiGS. 5-8, 11, 12) Monolete fern spores usually cannot be iden- tified to genus within the Polypodiaceae/Blech- naceae complexes. This is especially true of fossil spores that lack the enclosing perine. In the Ga- tuncillo material, two distinct monolete fern spores are common, and three others are present that may represent only variants or preservation forms. Type 1 (Fig. 5). Ambreniform, outer margin entire; monolete, laesura along concave side, straight, narrow, 20-25 um long, extending ca. 5 spore length; wall 2 um thick, laevigate; 70— 75 um by 45-50 um. This spore belongs to the form-genus Laviga- tosporites, a isa ka highly artificial taxon and, (Paleozoic to Recent an Type 2 (Fig. 6). Amb reniform, outer margin lobate reflecting sculpture elements; laesura straight, narrow, inner margin entire, 30-35 um long, extending 2⁄4 spore length; wall 3—4 um thick, verrucate, verrucae prominent on distal surface, diminishing about laesura; 55-65 um by 35-45 um. This spore type belongs to the form-genus Ver- rucatosporites which ranges from the Paleozoic to Recent. GRAHAM —NEOTROPICAL PALEOBOTANY IV 511 Type 3 (Fig. 7). This spore is similar to Type 1 but has minute punctations over the entire surface. 45 um by 35 um. Type 4 (Fig. 8). This spore is coarsely and irregularly punctate (corrosion artifact?). 50 um by 35 um. TypeS(Figs.11,12). This spore has echinae- like structures with the bases arranged to give a reticulate appearance. It is possible that this ap- parent sculpture represents remnants of a perine apressed to the spore surface. 50 um by 35 um. BROMELIACEAE (FIGS. 17-21) cf. Tillandsia Type. 1 (Figs. 17-19). Prolate; ht , narrow, inner mar- gin entire, 45 um long, extending nearly entire length of grain; tectate-perforate, wall thin (ca. 1.5-2 um), reticulate, reticulum fine, lumen slightly irregular in outline, ca. 1-1.5 um on dis- tal surface, diminishing toward colpus and es- pecially toward poles, muri ca. 1-1.5 um wide, smooth, margins entire; 50-55 um by 25-30 um. cf. Tillandsia Type 2 (Figs. 20, 21). Pollen of Tillandsia Type 2 differs from the preceding in having a much coarser reticulum. The di- ameter of the largest lumen is 3-4 um (versus 1.5—2 um). In other features the grains are sim- llar. The assignment of these grains to the Bro- meliaceae is based on similarity to several species of Tillandsia. The specimen illustrated in Figure 19, for example, is especially similar to some grains of T. excelsa Griesb. (Ocampo 001159, Costa Rica, CR). Other fossilspecimens are com- parable in morphology but differ slightly in mi- nor, quantitative features (e.g., almost immea- surable, minute differences in diameter of the nomic rank (family in then com- pared (cf.) to the genus Tillands The fossil record suggests an) the middle and FIGURES 16-31. 16. Trilete fern spore type 5, 2-1, J-24,4.— 17, 18. cf. Tillandsia type 1, 4-1, W-30,2-4.— — 20, 21. cf. Tillandsia type 2, 4-1, M-30,2-4. — 22. Palmae type 3, ^d D- 16,2. — 27. Palmae type 5, 2-1, 'L-22,1- 3. — 28, 29. cf. Campnosperma, 4-1, Q-25,3.—30, 31. cf. Aral apos 4-1, S-14. 512 late Eocene includes the time interval during which many modern genera. In slightly older deposits (e. g., Paleocene, early Eocene) many, perhaps most, angiosperm pollen types cannot be assigned to modern genera. In slightly younger deposits (e.g., Oligocene) most do resemble modern genera. The upper Eocene is intermediate and the Gatuncillo Formation is part of this transition period. The concepts and procedures illustrated by the identification of the Bromeliaceae cf. Tillandsia specimens are considered a conservative ap- proach that minimizes misidentifications while from this difficult tropical Eocene material. PALMAE (FIGS. 22-27) Five morphological categories of palm pollen were recovered from the Gatuncillo Formation. The moderate diversity and sustained numerical representation suggest that palms were a signif- icant, but not a dominant, element in the upper Eocene landscape of Panama (7.596 in sample 2; ; Table 1). Oblate, amb triangular to slightly oval triangular with rounded apices; tri- chotomocolpate, arms straight to slightly sin- uous, 18 um long, narrow, tapering to acute apex, inner margin entire to faintly dentate, extending within ca. 2-4 um of outer margin; tectate, wall 1.5-2 um thick, slightly thicker at apices; mi- croechinate, echinae minute, blunt to pointed, ensely aggregated, spaces between echinae giv- ing microreticulate aspect to some areas of grain surface; u Pollen ofthe extant genus to the microfossils in being trichotomocolpate but differs in details of sculpture pattern and wall thickness. Type 2 (Figs. 24, 25). Reniform; monosul- cate, sulcus straight to slightly sinuous, 24 um long, inner margin faintly dentate, extending en- tire length of grain; tectate to tectate-perforate, wall ca. 1.5 um thick; microreticulate (diam. of lumen less than 1 um); 28-30 um by 22-24 um. A . . . :1 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 Species within a number of extant genera have pollen similar to the microfossils, including Ai- phanes, Chamaedorea (e.g., C. caspariana Klotzsch), Coccothrinax [e.g., C. argentea (Lodd.) Sang.], and Copernica. Type 3 (Fig. 22). Amb ca. wedge-shaped (widest part of grain above equator); monosul- cate, sulcus sinuous, 52 um long, inner margin entire, extending entire length of grain; tectate to tectate-perforate, wall ca. 1.5 um thick; finely reticulate, muri smooth, inner margin entire, diam. of lumen ca. equal to width of muri (ca. 1 um or less); 56-60 um by 24-28 um. ype 4 (Fig. 26). Amb oval (elongated); monosulcate, sulcus straight, 36 um long, inner margin minutely dentate, Mim. entire oe of grain; tectate, wall ca. 1.5 um thick; micro- reticulate (diam. of Duns less than 1 im: 39- 43 um by 25-29 um. This grain is similar in aperture and sculpture pattern to Type 3, but the widest part of the grain is around the equator and it is considerably smaller ig. 27. Amb ca. wedge-shaped (widest part above equator); monosulcate, sulcus straight, 32 um long, inner margin dentate, ex- tending entire length of grain; tectate-perforate, wall 2 um thick; finely reticulate, muri smooth, inner margin entire, lumen ca. 1 um; 37-41 um by 20-24 um. ANACARDIACEAE cf. Campnosperma (Figs. 28, 29). Prolate, ranged, meridionally elongated, equidistant, cos- tae colpi, pores equatorially elongated, 1 um by 3 um, situated at midpoint of colpus, inner mar- gin entire; tectate-perforate, wall 1.5 um thick; finely striato-reticulate; 24—26 um by 19-21 um. The grains are similar to pollen of Campno- sperma and, to some extent, Comocladia. Campnosperma consists of two species with one (C. panamensis Standley) native to Panama. They are most abundant in the West Indies and com- FIGURES 32-53. ond usus 4-1, L-7 br erminalia, polar view, 4-1, U-38,1-2.— D-22, 1.—45, 46. Casearia, 4-1, K-35,2; 4-1, D- 21, — 32, 33. Ilex, equatorial view, 4-1, M-31.— 34. Ilex, polar view, 9-1, G-23,3.—35. cf. Par- ,2.—36, 37. Combretum/Terminalia, equatorial view, 4-1, Q-25,4. —38, 39. Com- l. ^s Tetragastris, 4-1, X-8, 2. —42-44. cf. Protium, 9-1, 48, 52, 53. Lisianthius, 4-1, X-23; 4-1, V-34,1.— 49, 50. cf. Tontalea, 9-1, M-38.—51. MEE 4-1, C-35,1 1985] GRAHAM —NEOTROPICAL PALEOBOTANY IV 513 514 monly grow in lowland, wet habitats such as swamps. AQUIFOLIACEAE Ilex (Figs. 32-34). Oblate-spheroidal, amb circular; tricolporoidate, colpi straight, 10 um long, broad (ca. 9 um at midpoint), inner margin diffuse, tapering to acute apex, equatorially ar- ranged, meridionally elongated, equidistant, ex- tending within 6 um of pole (P.I. 0.27), pores obscure, diam. ca. 2-3 um, ca. circular, situated at midpoint of colpus; intectate, wall thickness (viz., height of columellae) 3 um; clavate; 25-32 m. Ilex is a widespread genus of about 400 species. In Latin America it commonly occurs in mesic to slightly drier habitats. For example, in the Antilles it grows in the high-altitude pine forests (Howard, 1973); in Veracruz it is a member of the low evergreen selva, the high semi-evergreen selva, and the bosque caducifolio (Gomez-Pom- pa, 1973); and in Panama it is found at mid- altitudes in drier forest habitats and late second- ary forests (Porter, 1973) Fossil pollen of Tlex appears in the early Late Cretaceous of Australia, late Late Cretaceous of North America, and Paleocene of South America (Muller, 1981). By Tertiary times it is virtually cosmopolitan. In northern Latin America //ex is known from the Eocene of Panama (this report), the Oligocene of Puerto Rico (Graham & Jarzen, 1969), the Miocene of Veracruz, Mexico (Gra- ham, 1976), the Miocene of Panama (Graham, vei and the Quaternary of Panama (Bartlett & Barghoorn, 1973). CF. ARALIACEAE (FIGS. 30, 31) Prolate to prolate-spheroidal, amb circular; tricolporate, colpi straight, 10 wm long, inner margin entire, tapering to acute apex, equato- rially arranged, meridionally elongated, equidis- tant, pore obscure, slightly elongated equato- rially, ca. 1-1.5 um by 2.5 um, situated at midpoint of colpus; tectate-perforate, wall 1.5- 2 um thick; finely reticulate; 21-24 um. The primary purpose of including this micro- fossil is to document the occurrence of Aralia- ceae-type pollen in the assemblage, even though the specimens cannot be identified to genus. BIGNONIACEAE cf. Paragonia/Arrabidaea (Fig. 35). Oblate, amb circular; tricolporate, colpi straight, 18 um ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 long, broad (ca. 10 um at equator), equatorially arranged, meridionally elongated, equidistant, tapering to acute apex, margin diffuse; tectate to tectate-perforate, wall ca. 1.5 um thick; finely reticulate; 45 um. Modern pollen of the Bignoniaceae has been studied by Gentry and Tomb (1979). Based on their survey, and material in our reference col- lection, the microfossil is most similar to Par- agonia [P. pyramidata (Rich.) Bur.] and to Ar- rabidaea [e.g., A. corallina (Jacq.) Sandw.]. One collection of Paragonia (Croat 7895, BCI, Pan- ama, MO) is especially similar in being relatively large, thin-walled, and having a diffuse colpus margin. Pollen from two other collections (Har- vard exchange) differ in being ia thick- walled, and with distinct costae co Arrabidaea and Paragonia are both lania dis- tributed from Mexico to Brazil, Bolivia, and the former to Argentina. Paragonia is widespread in the tropical moist and wet forests of Panama. Both Arrabidaea and Paragonia produce nu- merous flowers in terminal panicles and are a Bu lasting about a month, dry season and erratically throughout the year” (Croat, 1978: 768), and the various Arrabidaea species have similar flowering peaks spaced throughout the year (Gentry, 1974). These fea- tures are consistent with small percentages in the fossil record (1% or less, Table 1) even though the genera are insect-pollinated lianas of the for- est canopy BURSERACEAE The two principal neotropical genera of the Burseraceae are distinguished palynologically by the striato-reticulate pollen of Bursera (ca. 80 species) and the psilate pollen of Protium (ca. 90 species). Within the latter, however, there are other smaller genera with similar pollen (e.g., Crepidospermum, 2 species; Hemicrepidosper- mum, | species; Tetragastris, 8—12 species). Two subtypes of this prolate, psilate pollen are rec- ognized in the Gatuncillo assemblage. One is comparatively thick-walled and is most similar o pollen of Protium. The other is thin-walled with a more hyaline appearance and is compa- rable to Tetragastris. Because there is some over- lap in pollen morphology among these modern genera, however, the fossils are referred to cf. Protium and cf. Tetragastris. cf. Tetragastris (Figs. 40, 41). Prolate; tri- 1985] colporate, colpi narrow, straight, ca. 25 um, inner margin entire, sides parallel, apex acute, equa- torially arranged, meridionally elongated, equi- distant, distinct costae colpi and costae pori 4 situated at midpoint of colpus, inner margin en- tire; tectate, exine relatively thin (1.5 um); psi- late, hyaline; 32 um by 24 um Tetragastris is a tree or shrub distributed from Brazil northward through the Antilles (to His- paniola) and Central America to British Hon- duras. The most common species is 7. pana- mensis (Engl.) O. Kuntze found in British onduras, Honduras, Nicaragua, Costa Rica, Panama, Venezuela, the Guianas, Brazil, and Peru. It is widespread in Panama and Porter (1970), citing Johnston (1949), noted that it is *an important forest tree on San Jose Island, where it may make up more than half of the " Accordi are no other reports of Tetragastris in the fossil record cf. Protium (Figs. 42-44). Prolate; tricolpo- rate, colpi narrow, faint, straight, ca. 26 um, in- ner margin entire, sides parallel, apices acute, equatorially arranged, meridionally elongated, equidistant, costae colpi and costae pori thick in equatorial region surrounding pore, pore equatorially elongated (colpi transversalis), 2.5 ituated at inne margin entire; tectate, wall 2. 5 um thick; psilate; 36 um by 24 um. Protium is a tree or shrub of the tropical forests of both the New and Old Worlds. In the neo- tropics it is most common in South America, with seven species listed for Panama (Porter, 1970). Throughout Central America it occurs in the tropical moist, premontane wet, and tropical wet forests. The fossil record of the Burseraceae is poorly known, with tentative reports that may extend the range of the family back to the lower Eocene (Muller, 1981). In the Caribbean area, Protium-type pollen is recorded from the upper Miocene Paraje Solo Formation of Veracruz, Mexico (Graham, 1976). | COMBRETACEAE Combretum/Terminalia (Figs. 36-39). Pro- late to prolate-spheroidal, amb circular (lobate reflecting arrangement of apertures); tricolporate with 3 pseudocolpi, colpi straight, ca. 22-25 um, GRAHAM —NEOTROPICAL PALEOBOTANY IV 515 inner margin entire, narrow, sides parallel for most of length, then tapering near terminus to acute apex, equatorially arranged, meridionally Wines equidistant, extending within 5 um of pole (P.I. 0.2), pore (endoaperture) frequently obscure, circula. diam psilate a faintly scabrate; 25-30 um by 18-22 The microfossils oriented in polar view (Figs. 38, 39) are most similar to Combretum/Termi- nalia pollen because the pore is more evident. Some microfossils oriented in equa tiple apertures (colpi), but the pore is obscure. In the pollen of most modern species of Combre- tum/Terminalia the pore is visible in equatorial view, but occasionally it is less obvious. The grain illustrated in Figures 38 and 39 is typical of Com- bretum/Terminalia, while those represented by Figures 36 and 37 are regarded as likely belong- ing to the same complex. It is not possible to consistently distinguish Combretum and Terminalia pollen un t designation Combretum/Terminalia. Similar pollen occurs in the Melastomataceae, but it is either all (9-15 um) and/or has distinct cos- tae co Bur jo and Terminalia are wide- spread in Latin America and most commonly occur in wet to moist forests, although C. fruti- bretum, Bartlett & Barghoorn, (1981) cited other occurrences that extend the complex into the upper Eocene of Cameroon, and it is now known also from the upper Eocene of Panama (this report). Ifthis is the approximate complete stratigraphic range of Combretum/Ter- minalia, it may explain the difficulty in identi- fying certain types from the Gatuncillo assem- blage, because the group would be in its early stage of differentiation. FLACOURTIACEAE Casearia (Figs. 45, 46). Prolate; tricolporate, colpi narrow, straight, 22-24 um, inner margin faintly undulating, apices acute, equatorially ar- ranged, meridionally elongated, equidistant, nar- 516 row costae colpi, pore equatorially elongated (colpi transversalis), 2 um by 5 um, situated at midpoint of colpus, inner margin entire; tectate, wall 1.5 um thick; microreticulate; 30-35 um by 22-25 um Casearia includes about 250 species of trees and shrubs widely distributed in tropical to sub- tropical regions of both hemispheres. Eight species are listed for Panama (Robyns, 1968), and these generally occur in moist forest types, although some (e.g., C. arguta H.B.K., C. commersoniana Camb.) range into the drier premontane or trop- ical dry forests (Croat, 1978). Pollen morphology of the family has been studied by Keating (1973). The fossil pollen is most similar to C. sylvestris Sw. In the Caribbean region, Casearia has been reported from the Oligocene of Puerto Rico (Graham & Jarzen, 1969) and the upper Miocene ent report from tends the fossil record to the upper Eocene. GENTIANACEAE Lisianthius (Figs. 47, 48, 52, 53). Oblate to oblate-spheroidal, amb circular; tricolporate, colpi broad at equator, tapering to acute apex, ca. 18 um long (equator to apex), extending with- in 6-7 um of pole (P.I. 0.02), equatorially ar- ranged, meridionally elongated, equidistant, in- ner margin entire to slightly diffuse, faint margo formed by gradual diminution of reticulum near colpus margin, pore circular, 4-5 um, situated at midpoint of colpus, margin diffuse; tectate-per- forate, height of columellae 2. 5 um in ‘equatorial mesocolpal region, di margins of colpi; reticulate, reticulum somewhat irregular, diam. of lumen ca. 2-3 um in equa- torial mesocolpal region, diminishing toward poles and margins of colpi, muri tall (ca. 2.5 um) in equatorial mesocolpal area, giving boxwork effect to o muri surface psilate, margins entire; 35—45 u he presence of Lisianthius pollen in the Ga- America, into northwest Colombia (Weaver, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 1972; Elias & Robyns, 1975). They are found in a variety of habitats including secondary scrub, pine forests, cloud forests, and savannahs rang- ing from sea level to about 1,800 m. Of particular interest from the standpoint of the fossil pollen record is that, although primarily insect-polli- nated, many species are highly floriferous and occur in dense stands at moderately low eleva- tions. Fossils of the Gentianaceae are known from the lower Eocene of Texas, and other records extend its range into the Paleocene (Crepet Daghlian, 1981). The Gatuncillo material rep- resents the first report of Lisianthius in the fossil record. HIPPOCRATIACEAE cf. Tontalea (Figs. 49, 50). Oblate, amb ca. circular, undulating; tricolporate, colpi narrow, straight, 12 um long, inner margin minutely den- tate, apices acute, extending within 3-4 um of pole (P.I. 0.015), equatorially arranged, meridio- nally elongated, equidistant, narrow margo, pore circular, 3 um, situated at midpoint of colpus; tectate to tectate-perforate, ektexine thicker (col- umellae longer) in equatorial mesocolpal area (ca. um), thinning toward apertures (1.5 um); finely reticulate; 26 um Tontalea is a neotropical genus consisting of about 31 species of lianas, slender shrubs, or small trees. Only one [T. richardii (Pery.) A. C. Smith] occurs in Panama (Dodson & Robyns, 1965). This species is a liana common in the upper can- opy of the tropical moist forest, although it is occasionally found at lower elevations and along the coast (Croat, 1978). Two other genera in the family contain species with pollen generally sim- ilar to Tontalea (Cueruea, Pristimera), but in the fineness of the reticulum and in wall structure (columellae) as seen in optical section, the fossil pollen is most similar to Tontalea. The genus has not previously been reported in the fossil record. JUGLANDACEAE Alfaroa/Engelhardia (Fig. 51). Oblate, amb oval-triangular; triporate, pore circular, ca. 2 um, inner margin entire, equatorially arranged, equi- FIGURES 54-71. 59. Eugenia/Myrcia, 2-1, E-30, — 54. Crudia, 2-1, T-21,1-3.— 55, 56. Malpighiaceae type 1, 4-1, C-22.— 57. Malpighiaceae type 2, 9-1, H-45,3.—58. cf. Ficus, 4-1, W-12,1-2.— tetra-aperturate form, 4-1, V-24.—61. Rhizophora, 4-1, K-24,1.—62, 63. Coccoloba, 4-1, 3.—60. Eugenia/Myrcia X-7,2-4.— 64, 65. Faramea, 2-1, G-14,2; 9-1, K-39,4.—66, 67. Rubiaceae type 1, 4-1, G-17,1-3.— 68, 69. Rubiaceae type 2, 4-1, S-14.— 70, 71. Paullinia, 4-1, V-11,1-2 1985] GRAHAM —NEOTROPICAL PALEOBOTANY IV 518 distant; tectate, wall 1.5 um thick; psilate to faint- ly scabrate; 26 um. Whitehead (1965) and Nichols (1973) con- clude that Engelhardia and Alfaroa cannot con- sistently be distinguished on the basis of pollen morphology. Crepet et al. (1980) noted that one exception may be the distinctly smaller and tri- angular pollen of section Psilocarpeae of Engel- hardia. Since our Gatuncillo material is not of this type, it is referred to A/faroa/Engelhardia. Similar pollen is stratigraphically and geograph- ically widespread throughout the Caribbean Ter- tiary and is a common microfossil in temperate regions of the northern hemisphere (Muller, 1981; Crepet et al., 1980). The fossil pollen is frequent- ly referred to the form-genus Momipites. It is known from the Oligocene San Sebastian For- mation of Puerto Rico (Graham & Jarzen, 1969), the Oligo-Miocene of Chiapas, Mexico (Langen- heim et al., 1967), and the upper Miocene Paraje Solo Formation of Veracruz, Mexico (Graham, 1976). In the Gatuncillo material the pollen is rare (three grains encountered among the three samples studied). Both Alfaroa and Engelhardia occur in Central America (Manning, 1960) and are typically, but not exclusively, associated with temperate forests. LEGUMINOSAE/CAESALPINIOIDEAE Crudia (Fig. 54). Prolate; tricolporoidate; colpi narrow, straight, 22-38 um, frequently ob- scured by sculpture elements, extending nearly entire length of grain, equatorially arranged, me- ridionally elongated, equidistant, pore area faint, ca. circular, situated at midpoint of colpus; tec- tate, but with occasional separations betw sculpture RE wall 1.5 um thick; distinctly and coarsely stria allel to long axis ie grain, surface psilate, margin entire, occasionally appearing beade m un- derlying pores in foot layer/endexine; 42. um by 7 um. In the Caesalpinioideae 12 genera have striate pollen, but only two of these (Crudia, Macro- lobium) are closely similar to the microfossils and occur in the New World. The genus Macro- lobium has about 60 species that grow in tropical America. The pollen of these differ from the fos- sil in having more coarse, thick striations and/ or with a much more conspicuous beaded ap- pearance to the striations imparted by the un- derlying pores. The genus Crudia has about 55 species with ten en growing in "e bua qos These are mainly Amazonian an p ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 (Cowan & Polhill, 1981: 131; pollen morphol- ogy, Graham & Barker, 1981). Crudia-type pol- len (under the form-genus name Striatocolpites catatumbus Gonzalez) is known back to the Eocene in northern South America and to the Paleocene in Nigeria (Muller, 1981). MALPIGHIACEAE Pollen of the Malpighiaceae is distinctive, but within the family generic identifications are dif- ficult. Fossil pollen is common, but at low fre- quencies, in the Caribbean Tertiary. In the Ga- tuncillo Formation two pollen types belonging to the Malpighiaceae are present, but these can- not be referred to any one genus in the family. Type 1 (Figs. 55, 56). Spherical, amb circu- lar; pericolporate, colpi ca. straight to slightly sinuous, narrow um, apices acute, inner margin Hingd donde pores oval to slit-like, long axis at right angle to colpus, 1 um by 3.5 um, situated at midpoint of colpus, inner margin entire to minutely dentate; tectate, wall conspic- uously thick (5 um); minutely punctate-micro- reticulate; 44 um. Type 2 (Fig. 57). Spherical, amb circular; .5 um, situated at midpoint of colpus, inner margin entire, distinct annulus ca. 2.5 um wide; tectate, wall 3 um thick; scabrate; 27 um. MORACEAE cf. Ficus (Fig. 58). Prolate, amb oval; dipo- rate, pores ca. circular, ca. 1.5 um, situated at apices of grain; tectate, wall ca. 1.5 um thick; scabrate; 10 um In most diporate Moraceae pollen the pores somewhat thicker wall. The fossil pollen is most similar to Ficus, but generic distinctions are so subtle that the microfossils are referred to cf. Ficus. The genus is large (ca. 1,000 species world- wide) and poorly understood taxonomically. In Panama, Ficus i ly associated with the moist forest type but can range into drier habitats. As expected from the pollination mech- anism and floral structure, pollen of cf. Ficus is rare in the Gatuncillo palynoflora. In the strati- graphic literature the pollen type is known as Psiladiporites minimus van der Hammen & mstra and ranges from lower Miocene to Re- cent (Germeraad et al., 1968). The Gatuncillo 1985] Occurrence extends the range back to the upper Eocene MYRTACEAE M hi (Figs. 59,60). Oblate to per- oblate, amb triangular; tricolporate, colpi nar- row, straight, 10:12 um, inner margin slightly dentate, equatorially arranged, meridionally elongated, equidistant, syncolpate, pores ca. 1 um, situated on equator at midpoint of colpus; tectate, wall thin (ca. 1.5 um), faintly scabrate; —19 um. Pollen of Eugenia and Myrcia is similar, and it is not possible to consistently refer ced microfossils to either genus (Graham, 1980). This pollen type is common in Gulf/Caribbean Ter- tiary deposits, ranging from the middle Eocene (Elsik & Dilcher, 1974) to Recent. An occasional tetracolporate form is encountered (Fig. 60), r portant components of tended vegetation but den ads through a wide variety of habitats which reduces their usefulness in paleoecological reconstructions. POLYGONACEAE Coccoloba (Figs. 62, 63). Prolate; tricolpo- rate, colpi narrow, straight, extending nearly en- tire length of grain, 30-38 um long, apices acute, inner margin entire, equatorially arranged, me- ridionally elongated, equidistant, costae colpi, pore (endoaperture) frequently obscure, ca. cir- cular, 2-3 um, situated at midpoint of colpus; tectate-perforate, wall thick (3 um), columellae me and eicany evident ü in 1 optical section, sex- t from nexine (foot ayez) pore diam. of lumen ca. 1 um; 39- 50 um by 23-36 um. Coccoloba is common in moist forest types of tropical America. Howard (1960) recognized 12 species for Panama, and several of these range into lowland coastal habitats where pollen can readily be incorporated into accumulating sedi- ments. Microfossils are common in the Gatun- cillo Formation (3.596 in sample 4; 296 in sample 9. Table 1), and in the upper Miocene Paraje Solo Formation of Veracruz, Mexico (Graham, 1976). The Eocene sp from Panama can- not be related to a single modern species but are quite similar to C. belizensis Standley. GRAHAM —NEOTROPICAL PALEOBOTANY IV 519 RHIZOPHORACEAE Rhizophora (Fig. 61). Prolate to prolate- pube tricolporate, colpi narrow, straight, , apices acute, equatorially arranged, point of colpus, inner margin entire forate, wall thick n um); (um reticulate; 19— 21 um by 22-24 u Fossil pollen of bins also listed in the literature under the form-genus name Zonoco- Stites, is almost by definition an important com- ponent of Tertiary lignites in warm-temperate to subtropical regions that form under coastal, brackish-water conditions (Cohen & Spackman, 1972; Scholl, 1964a, 1964b). Pollen of the six species (and one hybrid) are similar, but suffi- cient differences exist to tempt species distinc- tions in the fossil record. Modern and fossil pol- len of Rhizophora was studied by Langenheim et al. (1967), Leopold (1969), and Muller and Caratini (1977). The last authors concluded that three groups of species can be recognized on the basis of pollen morphology: R. mucronata/sty- losa; R. apiculata/lamarckii/mangle, and R racemosa. The microfossils in the upper Eocene Gatun- cillo Formation are difficult to identify because they are near the apparent time of early diver- sification and radiation of the genus. The oldest known records are from the lower to middle Eocene of southeast Asia (Muller & Caratini, 1977), and the genus first appears in tropical America in the late Eocene (viz., Gatuncillo time). As a result, there are a number of microfossils in our Panama material that appear similar to Rhizophora but are quite small (14-16 um by 18-20 um) and sufficiently close to spherical as to orient in various views, precluding determin- ing exact shape and/or aperture morphology. y ‘unknowns’ in this category represent Rhi- zophora-like pollen and may be early forms of the pollen in these tropical American sediments. The specimen in Figure 61 is a more typical ex- ample of Rhizophora in the Gatuncillo Forma- tion. RUBIACEAE Faramea (Figs. 64, 65). Oblate, amb oval; diporate, pores (short colpi) 6-7 um (apex to equator), inner margin entire, apices ca. rounded, situated at poles (apices of long axes), broad dif- fuse costae pori (colpi) 4-5 um wide, giving 520 stained effect to region surrounding aperture; tec- tate-perforate, wall ca. 1 um thick; finely retic- ulate; 27-30 um by 24-30 um. Faramea is a genus of about 125 species of trees and shrubs widely distributed in the An- tilles, Central America, and South America (Dwyer, 1980). The pollen 1 is typically triporate and reticulate, common to the family Rubiaceae, and these pollen types have been recorded from the Oligocene San Sebastian Formation of Puer- to Rico (Graham & Jarzen, 1969) and the upper Miocene Paraje Solo Formation of Veracruz, Mexico (Graham, 1976). Among the species with normal triporate pollen, however, are some in our reference collection (e.g., F. coarinensis Muell. Arg.) in which many of the grains are diporate. The Gatuncillo specimens are all of the diporate type. Consequently, the known stratigraphic range of the genus is upper Eocene to Recent, but the diporate form is known only from the upper Eocene, and the triporate type ranges from the middle Oligocene to Recent. The present report extends the range back into the Eocene from that reported in Muller (1981). Pollen of the Rubiaceae is abundant in the Gatuncillo Formation, but it frequently belongs to a generalized type common to many genera in the family. Two of the more common forms are described below. Type | (Figs. 66, 67). Oblate to oblate-sphe- roidal, tricolporate, colpi short (6-8 um), broad (ca. 4 um at equator), tapering to ca. acute apex, inner margin entire, equatorially arranged, me- ridionally elongated, equidistant, pore elongated equatorially (colpi transversalis), | um by 4 um, situated at midpoint of colpus; tectate-perforate, wall ca. 1 um, slightly thicker in mesocolpal re- gion; finely reticulate; 18-23 um This pollen type is common to several genera of the Rubiaceae, including especially Antir- rhoea, but also Calycophyllum, Erithalis, and others. Type 2 (Figs. 68, 69). This rubiaceous grain is similar to Type 1, except it is slightly larger (26 um versus 18-23 um) and more distinctly reticulate. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 SAPINDACEAE Cardiospermum (Fig. 72). Oblate, amb tri- angular; triporate, pores protruding ca. 3-4 um, —2 um diam., equatorially arranged, equidis- tant, aspidate, vestibulate; tectate-perforate, wall relatively thin (ca. 1.5 um); reticulate, muri equal to or slightly broader than diameter of lumen, giving punctate effect to portions of the exine; 45 um. Cardiospermum is a genus of about 14 species of herbaceous, annual or perennial vines widely distributed in tropical America and in Africa. Two species are currently known from Panama (C. grandiflorum Swartz, C. microcarpum H.B.K.), and another (C. halicacabum L.) likely occurs there (Croat, 1976). As noted by Muller (1981), Cardiospermum pollen has been men- tioned in the Tertiary fossil record (e.g., Leopold & MacGinitie, 1972, Eocene, Rocky Mountain region) but without documentation (description or illustration). Bartlett and Barghoorn (1973) record the genus from the Quaternary of Pana- ma. Paullinia (Figs. 70, 71). Oblate, amb trian- gular; triporate, pores 3—4 um, equatorially ar- ranged, equidistant; tectate-perforate, wall 2-3 um thick; finely reticulate; 45 um. Paullinia is a genus of scandent shrubs or lia- nas widely distributed in tropical America, with one species in Africa (P. pinnata L.). It is most characteristic of moist forest types. About 194 species are listed for the neotropics, and Croat (1976) recorded 37 for Panama. Pollen of the genus is known from the upper Miocene of Ve- racruz, Mexico (Graham, 1976) and from the Quaternary of Panama (Bartlett & Barghoorn, 1973). The Gatuncillo occurrence extends the stratigraphic record of Paullinia back to the up- per Eocene. Serjania (Figs. 73, 74). Oblate, amb trian- gular; tricolporate, colpi narrow, straight, 4-6 um long, inner margin entire to minutely dentate, equatorially arranged, meridionally elongated, equidistant, pores slit-like, situated at midpoint p FIGURES 72-89. 4-1, F-36,3-4; 2-1, M-39,2. — 72. Cardiospermum, 4-1, D-38,1.— 73, 74. Serjania, 4-1, S-22,1.— 75, 76. Mortoniodendron, —77, 78, 80. Pelliceria, 4-1, T-37,1; 4-1, S-27,1.—79. cf. C hrysophyllum, 4-1, -4.— S-19,2. —81. Unknown 3, 4-1, K-25,2. — 82, 83. Unkno wn 5, 4-1, P-22,4.— 84. Unknown 1, 4-1, X-9,2 Unknown 2, 4-1, C-19,3. — 86, 87. Unknown 7, 9-1, X-35.—88. Unknown 4, 2-1, M-28,4.— 89. Unknown 6, 9-1, K-43,1 1985] GRAHAM —NEOTROPICAL PALEOBOTANY IV "YS 522 of colpus; tectate-perforate, wall ca. 2 um thick; finely reticulate; 45 um is genus consists of about 215 species dis- tributed from southern United States to tropical South America. The plants are scandent shrubs or lianas and occur in a wide variety of habitats. Fifteen species are listed for Panama (Croat, 1976). Pollen of Serjania is known from the up- per Miocene of Veracruz, Mexico (Graham, 1976) and from the Quaternary of Panama (Bartlett & Barghoorn, 1973). The Gatuncillo oc- currence extends the range of the genus back to the upper Eocene. Among modern species, the microfossils are similar to S. conigera Turcz. of the tropical moist forest. SAPOTACEAE cf. Chrysophyllum (Fig. 79). Prolate, tricol- porate, colpi long (25-27 um), narrow, straight, inner margin minutely dentate, equatorially ar- ranged, meridionally elongated, equidistant, cos- tae colpi ca. 1-2 um wide, pore elongated equa- torially (colpi transversalis), ca. 1 um by 8-10 um, inner margin entire, situated at midpoint of colpus; tectate, wall 2.5 um thick; psilate to faint- ly scabrate; 52-54 um by 28-32 um. Chrysophyllum is a large genus of tropical trees (150 species), especially well-represented in the American tropics. The microfossils are similar to several genera of the Sapotaceae (e.g., Micro- 2 Pouteria) but more commonly resemble ; hence, the PEDIR cf. anna At There i is consid- erable range in size among the species (18-46 um in length), and a smaller form (13.5 um by 18.9 um) is reported from the middle Oligocene San Sebastian Formation of Puerto Rico (Graham & Jarzen, 9). The Gatuncillo specimens are larger (52-54 um) and extend the range of this pollen type from the middle Oligocene to the upper Eocene THEACEAE Pelliceria (Figs. 77, 78, 80). Oblate, amb oval- triangular to nearly circular; tricolporate, colpi narrow, straight, 15 um (equator to apex) (P.I. 0.4), inner margin entire, equatorially arranged, meridionally elongated, equidistant, colpi trans- versalis 1 um by 12-14 um, situated at midpoint of colpus; tectate to tectate-perforate, wall thick (7-10 um), separation of ektexine from endexine clearly evident on most specimens; sculpture variable from mound-like clusters (verrucae) with ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 subreticulum, to distinctly reticulate; size vari- able, ca. 50-70 um The pollen morphology and fossil record of Pelliceria has been summarized by Graham (1977). Although presently confined to lowland, coastal areas from Costa Rica to northwestern Colombia, the genus is widespread in Caribbean Tertiary deposits. In the stratigraphic literature the specimens are referred to the form-genus Psi latricolporites crassus van der Hammen & Wym- stra. It is known from the lower middle Eocene of Jamaica, upper Eocene of Panama, middle Oligocene of Puerto Rico (Graham & Jarzen, 1969), Oligo-Miocene of Chiapas, Mexico (Lan- genheim et al., 1967), and the Tertiary of the Guiana Basin (van der Hammen & Wymstra, 1964). A notable feature of modern Pelliceria pollen is the extensive range in size and sculpture patterns, which is also matched by the micro- fossils. Pollen of Pelliceria is common in the Ga- tuncillo Formation and was probably an impor- tant component of the fringing mangrove vegetation. TILIACEAE Mortoniodendron (Figs. 75, 76). Peroblate, amb oval-triangular to circular; tricolpate, colpi short (3 um equator to apex), apices ca. rounded, inner margin entire, equatorially arranged, me- ridionally elongated, equidistant, conspicuous costae colpi ca. 1.5—2 um wide; tectate-perforate; finely reticulate, reticulum closed, regular, muri ca. 0.5 um wide, m surface psilate, margins entire; 23-32 u Pollen of o PUES and its fossil rec- ord has been summarized by Graham (1979b). In that report microfossils had not been re- covered from the Gatuncillo Formation, so these specimens represent new occurrences in the Ca- ribbean Tertiary. Other records include the mid- dle Miocene Gatun Formation of Panama (Gra- ham, 1979b), the upper Miocene Paraje Solo Formation of Veracruz, Mexico (Graham, 1976), and the Quaternary of Panama (Bartlett & Barg- hoorn, 1973). Similar fossils have been reported by Tschudy and van Loenen (1970) under the name Tiliaepollenites from the upper Eocene of Mississippi. The complete stratigraphic range, as presently known, is upper Eocene to Recent. There are about five species of these small shrubs to tall trees growing from southern Mexico through Central America. They are most char- acteristic of tall moist forests. The diversity in 1985] GRAHAM fossil pollen illustrated by Graham (1979b) sug- gests that other species or related taxa may yet be described from modern tropical forests. UNKNOWNS In addition to these specimens, there are others in the Gatuncillo Formation for which biological affinities are unknown. The number of such spec- imens increases with age of the strata, and the point at which most fossil pollen cannot be re- ferred to a modern genus or family is frequently in a broad transition zone between the upper and middle Eocene. Consequently, the upper Eocene microfloras include a mixture of specimens that can be referred to modern genera or families, depending on the amount of variation in the modern analogs (the knowns y; others for which modern y have not yet been encountered in the reference collection or liter- ature (the unknowns); and a substantial number that are not known to match the pollen of any extant taxa and probably represent extinct forms (the unknowables). The last decrease upward to- ward the Oligocene and younger strata, and in- crease below the upper Eocene to the point that they begin to significantly hamper the recon- struction of paleocommunities and paleoenvi- ronments is based on fossil palynomorphs. Re- semblage although nese can a b recog- nized ł by their er color, and/or distinct morphology jme if rede- posited from Paleocene or older sediments). The most common unknowns from the Ga- tuncillo Formation are illustrated in Figures 81- 144 and briefly described below. These undoubt- edly include both extinct forms and those for which biological affinities will ultimately be es- tablished in future studies. UNKNOWN ECHINATE TYPES Type | (Fig. 84). Spherical, amb circular; non- aperturate; tectate, wall 1 um thick; echinate, spines ca. 1 um, evenly spaced, are te dense (1.5-2 um between spines); small (10 um). Type 2 (Fig. 85). Oblate, amb oval-triangu- lar; tricol(poroid?)ate, colpi ca. 3 um wide at equator, tapering to acute apex, 6-7 um long (P.I. 0.2), inner margin entire, equatorially arranged, meridionally elongated, equidistant; tectate, wall 1.5 um thick; echinate, echinae short (2 um), acute, some slightly curved, ca. evenly spaced, —NEOTROPICAL PALEOBOTANY IV 523 moderately dense (ca. 3 um between spines); 18 um. Type 3 (Fig. 81). Spherical, amb circular; non- aperturate (possibly monocolpate?); tectate, wall 1.5—2 um thick; echinate, echinae short (2 um), blunt, ca. evenly spaced, moderately dense (ca. 3 um between spines); 23 um. Type 4 (Fig. 88). Spherical, amb circular; non- aperturate; p ae wall 1.5 um; echinate, echi- nae short 4 um), longer ones curved, acute, widely od (6-7 um); 39 um Type (Figs. 82, 83) Prolate tricol- (poroid?)ate; colpi 18-20 um long, inner mar- gin entire, equatorially arranged, meridionally elongated, equidistant, possibly slight costae col- pi; tectate, wall thick (3-4 um), separation of ektexine and endexine clearly evident; minutely echinate; 27 um. UNKNOWN INTECTATE TYPE Type 6 (Fig. 89). Spherical, amb circular; non- aperturate; intectate, columellae short, mound- like, slightly elongated baculae, some constricted at base (clavae/gemmae), diam. of sculpturing elements 2-4 um, smooth, some ca. hyaline, moderately dense (2-4 um between sculpturing elements); 40 um UNKNOWN TRIANGULAR TYPE Type 7 (Figs. 86,87). Oblate, amb triangular, outer margin undulating; triporate, pores circu- lar, diam. 4-5 um, inner margin entire, situated at apices of triangular grain, equidistant; tectate, wall thick (ca. 3 um), separation of ektexine and endexine evident; psilate to faintly scabini, some coarser, irregular equatorial region near r pores; 31 um. UNKNOWN OBLATE/OBLATE-SPHEROIDAL TYPES Type 8 (Figs. 90, 91). Oblate-spheroidal, amb by diminution of reticulum along colpus margin equatorially arranged, meridionally elongated, quidistant; tectate-perforate, wall 2 ick; reticulate, reticulum closed, muri thin, smooth, d sinuous, diam. of lumen ca. 4 um; 36 m 9 (Figs. 92-95). Oblate, amb oval-tri- angular; tricolpate, colpi short diam. 3-4 um, equatorially arranged, equidis- tant; tectate-perforate, wall 1.5 um thick; retic- 524 ulate, reticulum closed, regular, muri thin, smooth, diam. of larger lumen ca. 3 um; 20-22 m The specimen in Figures 94 and 95 may be a slightly flattened, corroded form of the specimen in Figures 92 and 93. They are representative of rubiaceous is of pollen common in the Ga- tuncillo Formation. Type 10 "sei 96, 97). Oblate, amb circular; tricolporate, colpi faint, 14-16 um, equatorially arranged, meridionally elongated, equidistant, pore large (diam. 8-10 um), situated at midpoint of colpus; tectate-perforate, wall 2.5 um thick; reticulate, reticulum closed, ca. regular, muri thin, smooth, diam. of larger lumen ca. 2 um; 33 um. Type 11 (Figs. 98, 99). Oblate, amb circular; triporate, pores circular, 2.5 um, narrow annulus, inner margin entire, equatorially arranged, equi- distant; tectate, wall 1.5 m thick; scabrate (pos- sibly microreticulate); 2 um. T (Fi ced amb circular; tri- colpate, colpi 10 um, extending within 4 um o pole (P.I. 0.2), tapering to acute apex, inner mar- gin minutely dentate, equatorially arranged, meridionally elongated, equidistant; tectate-per- forate, wall 1-1.5 um thick; finely reticulate; 20 um. Type 13 (Fig. 101). Oblate, amb circular; tri- colporate, colpi 5-6 um, extending within 5 um of pole (P.I. 0.2), apex acute, inner margin entire to minutely dentate, equatorially arranged, me- ridionally elongated, pore conspicuous, diam. 3 um, narrow annulus ca. 1 um; tectate-perforate, wall 1.5 um thick; finely and regularly reticulate; 23 um. Type 14 (Fig. 102). Oblate, amb triangular; papse. colpi short (3-4 um), narrow (P.I. 0.2), tapering to acute apex, inner margin entire, um y 3 um, situated at midpoint of colpus; eae. wall 1.5 um thick; psilate (punctations, possibly artifacts); 13 wm Type 15 (Fig. 103). This grain is similar to ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Unknown 14 except it is more oval-triangular, the wall is slightly thinner, and the surface ap- pears more sculptured (faintly rugulate, possibly some punctations). Type 16 (Figs. 104, 105). Oblate-spheroidal, amb circular; tricolporate, colpi long (12 um), tae colpi, pore elongated equatorially, sit- uated at midpoint of colpus; tectate-perforate, wall 1.5—2 um thick; reticulate, reticulum closed, regular, diminishing along colpus (faint margo), muri smooth, equal in diam. to lumen (ca. 1 um) giving punctate effect, appearing ca. linearly aligned; 18 um Type 17 (Fig. 106). Oblate, amb circular; tri- colpate, colpi 12 um, tapering to acute apex, straight, inner margin ca. entire, extending with- in 6 um of pole (P.I. 0.2), equatorially arranged, meridionally elongated, equidistant; intectate, columellae thin needle-like baculae, some slight- ly clavate; ca. 2 um long; Type 18 (Figs. 107, 108). Oblate, amb cir- cular; tricolpate, colpi 10 um long, extending within 6 um of pole (P.I. 0.2), tapering to acute margin minutely dentate, tant; tectate-perforate, wall thin am. of grain (1-1.5 um); reticulate, reticulum EM regular, dde of muri and diam. oflumen ca. 1 um; 30 u Type 19 (Fig. 109). Oblate, amb circular to oval-triangular; triporate (short colpi) pores slightly elongated meridionally, 1 um by ca. 3 um, inner margin entire, conspicuous costae pori (colpi) 2.5 um wide, pores equatorially arranged, equidistant; tectate-perforate, wall 1.5 um thick; Sarai reticulum regular, closed, eee Es diam. of lumen 1-2 um); pe 20 (Fig. 111). Oblate, Ab. oval: trian- "E tricolporate, colpi short, faint, equatorially arranged, meridionally elongated, equidistant, pores elongated equatorially (1 um by 3 um), situated at midpoint of colpus, costae pori, slightly — FIGURES 90-123. 90, 91. Unknown 8, 4-1, W-9,1.—92-95. Unknown 9, 4-1, N-15; 4-1, W-23,1-3.—96, 97. known 21, 4-1, déco mire 115. Unknown 23, 4-1, 1, 13.24 —116. Unknown 24, d s rA 1985] GRAHAM —NEOTROPICAL PALEOBOTANY IV 525 526 aspidate; Pes wall 1.5 um thick; rugulate- striate; 32 u ype 21 (ees 112, 113). Oblate, amb ca. circular; stephanoporate (short colpi; 5 in num- r), pores slit-like, equatorially arranged, equi- distant; tectate-perforate, wall thin (1.5 um); finely reticulate; 40 um. UNKNOWN PROLATE, TRICOL(POR)ATE TYPES Type 22 (Fig. 110). Prolate; tricolpate, colpi long (24 um), narrow, straight, apices acute, inner margin entire to minutely dentate, equatorially arranged, meridionally elongated, equidistant; tectate, wall 1.5 um thick; psilate to faintly sca- brate; 27 um by 15 um Type 23 (Figs. 114, 115). Prolate; tricolpo- rate, colpi long (12 um), narrow, ene apices acute, inner tire, ranged, meridionally elongated, eanidistant pores faint, situated at midpoint of colpus; tectate, wall 1 um thick; psilate to faintly scabrate; 15 um by 12 um. Type 24 (Fig. 116). Prolate; tricolpate, colpi long (16 um), narrow, straight, apices acute, inner margin entire, equatorially arranged, meridio- nally elongated, equidistant; tectate, I 1 um thick; faintly scabrate; 20 um by 14 uv Type 25 (Fig. 117). Prolate; ricolporoidate colpi long (16 um), narrow, straight in entire, equatorially arranged, meridionally Mon gated, equidistant; tectate-perforate, reticulate, diam. of lumen ca. 1 um; 20 um Type 26 (Figs. 118, 119). Prolate; tricolpate, colpi long (24 um), narrow, straight, inner margin entire to minutely dentate, equatorially arranged, meridionally elongated, equidistant; tectate-per- forate, wall 2 um thick; finely reticulate, diam. of lumen and width of muri ca. 1 um; 27 um by 23 um. Type 27 (Figs. 120, 121). Prolate; tricolpo- rate, colpi long (18 um), narrow, straight, inner um thick; finely reticulate, diam. of lumen and width of muri ca. 1 um; 27 um by 23 um. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Type 28 (Figs. 122, 123). This type is similar to Unknown 27, except costae colpi are not ev- Es e 29 (Figs. 124, 125). Prolate; tricolpo- Na s long (18 um), narrow, straight, inner margin entire to minutely dentate, costae colpi 2.5 um wide, equatorially arranged, meridionally elongated, equidistant, pore circular, 2 um, inner margin entire, situated at midpoint of colpus; tectate-perforate, wall 1.5 um thick; finely retic- ulate, reticulum closed, regular, diam. of lumen k i of muri ca. 1 um; 21 um by 18 um e 30 (Figs. 126, 127). Prolate: tricolpo- NU pue long (22 um), narrow, straight, inner margin entire to minutely dentate, costae colpi 3 um wide, equatorially arranged, meridionally elongated, equidistant, pore slightly elongated equatorially (1.5 um by 3 um), situated at mid- point of colpus; tectate-perforate, wall 1.5 um thick; finely reticulate, reticulum closed, regular, iam. of lumen and width of muri ca. 1 um; 27 um by 19 um. Type 31 (Fig. 128). This specimen is similar to Unknown 30, but slightly smaller (22 um by 14 um). Type 32 (Fig. 129). Prolate; tricolpate, e long (28 um), narrow, straight, inner margin m thick; psilate to faintly striate; 33 um by 23 um. Type 33 (Figs. 130, 131). Prolate; tricolpate, colpi long (32 um), narrow, straight to slightly sinuous, inner margin minutely dentate, equa- torially arranged, meridionally elongated, equi- distant; tectate to minutely tectate-perforate, wall 1.5 um (thin in relation to size of grain); very finely irri, 36 um m. Type 34 132). Prolate to prolate-sphe- roidal; tri(to firii tetra-)colporate, colpi long (38—40 um), narrow, straight, apices acute, costae gated, equidistant, pores conspicuous, oval (13 um by 5 um), inner margin entire, situated at midpoint of colpus; tectate, wall 1.5—2 um thick; minutely reticulate; 45 um by 33 um. Ficures 124-141. Unknown 31, 4-1, W-15,3-4.— — 124, 125. Unknown 29, 4-1, G-8,1-3.—126, 127. Unknown 30, 4-1, V-43,1.—128. 129. Unknown 32, 4-1, E-35,2-4.— 130, 131. Unknown 33, 4-1, W-23.—132. Unknown 34, 4-1, K-37,1.—133-135. Unknown 35, 4-1, N-24,1.—136, 137. Unknown 37, 2-1, R-30,1-3.— 138, 139. Unknown 36, 2-1, M-35,1.—140, 141. Unknown 38, 4-1, T-22,3 GRAHAM — NEOTROPICAL PALEOBOTANY IV 527 1985] 528 Type 35 (Figs. 133-135). Prolate to prolate- spheroidal; tricolporate, colpi long (30 um), nar- TOW, straight, inner margin entire, apices acute, costae colpi 4 um wide, pores (endoapertures) elongated equatorially (2-2.5 um by 4-4.5 situated at midpoint of colpus; tectate-perforate, wall 2.5-3 um thick, ektexine clearly distinct from endexine; finely reticulate, diam. of lumen ca. 1 um; 36 um by 29 um. Type 36 (Figs. 138, 139). This species is sim- ilar to Unknown 35, but is slightly larger (41 um by 24 um) with a thinner exine (1.5-2 um). Both are similar to several Vip dii of the Anacar- gy with some Type 37 (Figs. 136, 137). This specimen (40 um by 21 um) is typical of the prolate, tricol- porate, costate, reticulate type previously de- scribed, but is distinctive in that the sharp edges of the muri give a c ee ‘scooped out’ appearance to the e Type 38 (Figs. ps a This large speci- men (68 um by 40 um) is similar to the smaller specimens described as Unknowns 35 and 36, and resembles some Euphorbiaceae, as well as the larger Anacardiaceae. In many specimens of this type, the bases ofthe columellae are arranged into a subreticulum, and the wall is compara- tively thick (3—3.5 um). Type 39 (Fig. 142). This specimen (58 um by 40 um) is also anacardiaceous/euphorbia- ceous in aspect but has finer, more delicate col- umellae and a relatively thin wall (2 um). Type 40 (Figs. 143, 144). This specimen (68 m by 45 um) is of the same general type de- BENE for the Anacardiaceae/Euphorbiaceae forms but has a wall intermediate in thickness (2-3 um) between Unknowns 38 and 39. AI- though these minute differences in wall thickness and columellae coarseness are difficult to quan- tify with exact measurements under light mi- croscopy, such differences do impart recogniz- able differences between the specimens. PALEOCOMMUNITIES Arrangement of taxa from the Eocene Gatun- cillo Formation into paleocommunities is diffi- cult because of the age of the deposits. The mi- crofossils represent remnants of a vegetation existing ca. 40 Ma, which was a transitional pe- riod when many lineages were differentiating into modern taxa (see previous discussion under 77/- landsia). Consequently, the morphology of nu- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 merous specimens do not match, exactly, any modern analog, even though they may clearly be recognized as belonging to a given family. Such specimens often combine the pollen characters of two or more genera that are now palynologi- cally distinct. This is no different in kind from similar problems encountered in -younger Ter- 1 tiary [ numerous with increasingly older deposits. Further, it can not always be assumed that the ecological requirements of the fossil represen- tative and its presumed modern analog are iden- tical. This proven is lessened to some degree when pal ona large number of genera, rather dian O ona few genera regarded as key or indicator co ponents. The many unknowns in the es palynoflora also complicate interpretations. These limitations, however, do not automat- ically preclude detecting the general kinds of pa- eocommunities present, even in late Eocene de- posits. When fossil taxa cluster into community types that include many genera, based on the habitat of presumed modern analogs, it is likely that assemblage was present, even though indi- vidual components may be difficult to identify or may have changed associations through time. Another problem involved in recognizing pa- leocommunities based on fossil palynomorphs is a bias toward lowland vegetation and wind-pol- linated components. In tropical environments these limitations are minimized, although not eliminated, by the considerable outwashing of pollen into lowland basins of deposition by high rainfall (Germeraad et al., 1968). Muller (1959) demonstrated that pollen from upland commu- nities is readily transported into coastal envi- (Graham, 1976) include lowland communities of Rhizophora (manglar), the mid-altitude bosque caducifolio, the higher altitude bosque de pino y encino, and the high altitude bosque de oyamel (Abies). Although there is some evidence that cooler temperatures shifted the vegetation zones downward (viz., the presence of Picea pollen in the sediments), many of these communities were still altitudinally removed some distance from the coastal basin of deposition. Similarly, the presence of such large, ento- mophilous pollen types as Hauya (Onagraceae, 85 um), Ludwigia (Onagraceae, 104 um), and Hampea/Hibiscus (Malvaceae, 135 um) in trop- 1985] GRAHAM —NEOTROPICAL PALEOBOTANY IV 529 TABLE 2. Distribution of taxa identified ied the middle(?) to upper Eocene Gatuncillo Formation among comparable modern community types in Panam Tropical Moist Forest (21 genera + 1 widespread) T bisous inii Pteris, Tillandsia, Campnosperma, Casearia, Chrysophyllum, Coccoloba," Combre- um, a, Faramea,? Ficus, Myrcia, Paragonia, Paullinia, Pelliceria, Protium, Rhizophora, Serjania,* pena labaa Tontalea Tropical Wet Forest (19 genera) Selaginella, Pteris, Tillandsia, Casearia, Chrysophyllum, Coccoloba, Combretum, Crudia,* Eugenia, Faramea, Ficus, Lisianthius,! Mortoniodendron, Myrcia, Paragonia, Paullinia, Protium, Terminalia, Tetragastris Premontane Wet Forest (14 genera + 1 widespread) Selaginella, Pteris, Tillandsia, Casearia, Coccoloba, Combretum, Eugenia, Faramea, Ficus, Paragonia, Paul- linia, Protium, Serjania, Terminalia, Tetragastris Premontane Moist Forest (3 genera + 6 widespread) Combretum, Eugenia,* Faramea,* Ficus,* Ilex, Paullinia,* Serjania, Terminalia,’ Tetragastris* Premontane Rain Forest (4 genera) Casearia, Paullinia, Serjania, Tillandsia Tropical Dry Forest (4 genera + 2 widespread) Cardiospermum, Casearia, Coccoloba,* Combretum, Serjania, Tetragastris* Premontane Dry Forest (4 genera + 1 widespread) Cardiospermum, Casearia, Combretum, Eugenia, Ficus Lower Montane Moist Forest (3 genera) Pteris, Alfaroa, Engelhardia (Oreomunnea) Lower Montane Wet Forest (2 genera) Pteris, Tillandsia Lower Montane Rain Forest (none) Montane Wet Forest (none) Montane Rain Forest (none) a The genus occurs in this community but is more common or typical in other habitats > Most common in more or less stressed envirionments (swamps, beaches, limestone outcrops) but occurs in all lowland life zones. * Myrtales tend to be more common in Panama in more or less stressed envrionments (e.g., Cerro Jefe) in somewhat drier forest types. 4 The genus is most common in the tropical wet forest, but the most common species (F. occidentalis) is very abundant in the tropical moist forest. * Most common in South America in inundated fores f Tropical wet forest, but the most common species 7 skinneri) is more or less weedy and occurs in drier forest types. ni sediments (Graham, 1976; Graham & Jar- data would automatically preclude the presence zen, 1969) is nian to the effectiveness of out- wallin ng and water transport in depositing a variety of pollen = in lowland habitats. There paren are obvious exceptions, such as low pollen-pro- ducing communities removed great distances from the basins of deposition (e.g., paramo), or distinctly desert communities where pollen has little chance of long-distance water transport or types, are commonly incorporated into lowland basins of deposition in tropical regions. Con- surviving oxidation under the arid conditions. Evidence for both vegetation types is meager in the Tertiary plant microfossil record for Latin America, but no rational interpretation of the versely, when the common and defining elements of a particular community or ecotype are con- sistently rare or absent in a large tropical Tertiary palynoflora, with such exceptions as noted ear- 530 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURES 142-144. lier, it is likely that assemblage was absent, poor- ly defined, or restricted in distribution (i.e., not a prominent component of the vegetation). d : ern counterparts (Table 2). The life-zone classi- fication is from Holdridge (1967; Holdridge et al., 1971) and was used by Croat (1978) for de- scribing the vegetation of Barro Colorado Island. It is noted that the purpose of Table 2 is not to record the complete range of the genera among present-day communities in Central America. Rather, the genera are placed according to their principal or common occurrence in recognition that some are more widely distributed or may contain a species that extends beyond the range ham, 1980), both are listed to maximize the number of communities potentially represented in the Gatuncillo palynoflora. When these genera are assigned to life-zones, using distributions cited by Croat (1978), family treatments in the “Flora of Panama," and com- ments provided by Croat and Gentry (pers. comm.), an obvious pattern emerges (Table 2). Elements from lowland, moist communities con- stitute the overwhelming majority in the Gatun- 142. Unknown 39, 4-1, X-8,2-4.—143, 144. Unknown 40, 4-1, X-32,2. cillo assemblage. Equally evident is the abrupt decrease in genera characteristic of 1) montane, and 2) dry forest types. Even within the con- straints on interpretation previously discussed, and acknowledging that some genera range be- yond the communities listed in Table 2, it is difficult to avoid the conclusion that higher al- titude and drier vegetation types were poorly rep- resented. A conservative interpretation ofthe ex- isting data is that the listings in Table 2 probably reflect, in a general way, the relative importance of the paleocommunities characterizing the late Eocene landscape in this part of central Panama. The principal paleocommunities present were the tropical moist forest, tropical wet forest, and pre- montane wet forest. Less prominent were the premontane moist forest, premontane rain for- est, tropical dry forest, and premontane dry for- est, although representation in these forest types is so low, and involves primarily genera that are also found elsewhere, that some may have been because the pollen cannot consistently be distin- guished under the magnifications of light mi- croscopy, and the specimens are rare in the assemblage (Table 1; ca. three specimens re- covered). This forest type, together with the low- er montane wet forest, lower montane rain forest, montane wet forest, and montane rain forest were probably poorly represented or absent in the re- 1985] gion. This general reconstruction of the paleo- communities is consistent with other indepen- dent geological and paleontological evidence on paleoenvironments and paleophysiography. PALEOENVIRONMENTS Reconstruction of paleoenvironments for the Gatuncillo flora is facilitated by information available on the geology (particularly sediment ypes), marine invertebrate and terrestrial mam- mal paleontology, and plate tectonic history o i leobotanical results can be interpreted. The microfossils were recovered from lignite deposits that are common in Tertiary formations throughout the Gulf/Caribbean region. The northernmost limit for any significant Present- ern pe Scholl, 19642. 19640). Thus the presence of the lignite itself suggests tropical to subtropical en- vironments. Since these lignites commonly con- tain pollen of Rhizophora, Conocarpus, Avicen- nia, Laguncularia, and Pelliceria, this is further confirmation that the sediments reflect tropical SE es Finally, the Gatuncillo lignites are associated with coralline limestones character- istic of deposition in near-shore, tropical habi- tats. Plate tectonic reconstructions show that dur- ing most of the Tertiary the present region of Panama was occupied by a series of volcanic islands located above the subduction zone be- tween the east Pacific (Cocos) and developing Caribbean plates (Coney, 1982; Dengo, 1973). Intense tectonic activity is reflected by more than 113 faults observed within a 3 km section of the Gaillard Cut portion of the Canal (R. H. Stewart, rs. comm.). Volcanic c activity is evidenced by wealth of biogeographic information is consis- tent with a model of discontinuous land surface between North and South America during most of the Tertiary, culminating with uplift of the Panama land bridge in latest Pliocene and Pleis- tocene times (Graham, 1973b; Marshall et al., 1976,1981,1982; Raven & Axelrod, 1974; Webb, 1976; Whitmore & Stewart, 1965; Woodring, 1966). This documentation of the insular nature of the landscape is important in interpreting cli- GRAHAM —NEOTROPICAL PALEOBOTANY IV 531 matic histories because biotas of small- to mod- erate-sized islands are often more insulated from climatic changes than comparable biotas of in- land continental areas. If these islands are of relatively low relief, climatic changes that are well-documented in paleofloras derived from continental areas of more diverse physiography (viz., like those from Veracruz, Mexico or the Andean region) may not be as evident in paleo- floras derived from low-lying insular vegetation. The palynological evidence previously dis- cussed suggests that the Eocene landscape, in the vicinity of the Gatuncillo depositional basin, was of moderately low relief. There is no evidence for either locally deposited or long-distance wind or water transported pollen from montane com- munities. Equally rare is pollen from dry vege- tation types. Thus, the local paleoenvironmental setting for these late Eocene biotas appears to include a se- ries of volcanic islands of moderate to low relief, bordered seaward by shallow limestone-depos- iting coralline communities, fringed by man- grove vegetation of Rhizophora and Pelliceria, supporting inland communities of tropical moist, tropical wet, and premontane wet forests, grow- ing under the general tropical conditions of high and evenly oo rainfall and high uniform temperature These d paleoclimatic conditions under which the Gatuncillo flora existed can be refined, to some extent, by a comparison with modern communities of similar composition. Normally, communities as old as the late Eocene resemble modern assemblages of comparable habitats only in a very general way. In the case of the Gatun- cillo flora, however, there is a modern analog in which virtually all the components of the paleo- swam nd b Atlantic side of Gatun fC where the vegetation shows considerable simi- larity to the paleocommunities, is nearly double that at Balboa, on the Pacific side where the vege- tation shows distinctly less similarity to the pa- leocommunities (107.3" versus 68”; Croat, 1978: 3). A very general estimate of temperature re- gimes is provided by data from the forest floor (versus open clearings) on Barro Colorado Island in Gatun Lake (Rubinoff, 1974, cited in Croat, 532 1978). There the mean maximum forest floor temperature is 28.0°C and the mean minimum is 21.1°C. Although these figures provide only the most general of estimates for upper Eocene conditions in the vicinity of the Gatuncillo de- positional basin, they are of interest as the first approximation of terrestrial paleoenvironments available for all of northern Latin America based on paleobotanical data. As noted earlier, the nearest Eocene floras are in South America, which during this time was isolated between Africa and orth America and the Mississippi Embaymen floras to the north. More precise reconstructions of regional paleoenvironments must await dis- covery of other fossil floras from the upper Eocene elsewhere in Central America. Of biogeographic interest is the list of families provided by Raven and Axelrod (1974) consid- ered to be 1) of South American origin and mi- grating to North America in Eocene or later times, and 2) those probably already established in North America by the late Eocene. An exact cor- relation would not be expected because the pa- leoflora is of an age and location that could pre- serve the earliest migrants from both continents. Nonetheless it is worthwhile to record the anal- yses because eventually similar information from other paleofloras, combined with biogeographic studies ofthe modern vegetation, will allow more precise estimates as to the region of origin of certain angiosperm families. Among the families listed by Raven and Axelrod (1974) as primarily of South American origin and migrating into North America during Eocene or later times, only four are recorded in the Gatuncillo palynoflora: ia e rti Malpighiaceae. Among those listed as already established in North America by the Eocene, 12 were represented in the Gatuncillo palynoflora: Anacardiaceae, Aquifoliaceae, Araliaceae, Big- noniaceae, Leguminosae (Caesalpinioideae), Moraceae, Myrtaceae, Rubiaceae, Sapindaceae, Sapotaceae, Theaceae, and Tiliaceae. At the fam- ily level the palynoflora seems to have somewhat greater affinities with North American-derived elements than with South American ones. None ofthe families listed by Raven and Axelrod (1974) as primarily characteristic of arid habitats in tropical America were represented in the Gatun- cillo assemblage. Genera identified from the Gatuncillo For- mation that have not previously been reported in the fossil record include Lisianthius (Gentia- naceae) and Cardiospermum (Sapindaceae). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Three other tentative reports will have to await additional specimens, and possibly SEM/TE studies, for confirmation. These include cf. Par- agonia/Arrabidaea (Bignoniaceae), cf. Tetragas- tris (Burseraceae), and cf. Tontalea (Hippocra- tiaceae). All are rare in the assemblage (Table 1). Range extensions are recorded for six pollen types in the Caribbean Tertiary (based primarily on previous stratigraphic ranges summarized by Germeraad et al., 1968; Muller, 1981): Casearia (from middle Oligocene), cf. Ficus (7 Psiladispo- rites minimus; on Moraceae), Faramea (di- porate form, first report; triporate known from middle Oligocene), Paullinia (from upper Mio- cene), Serjania (from upper Miocene), and cf. Chrysophyllum (from middle Oligocene). The next palynofloras to be treated in this study are from the Culebra, Cucaracha, and La Boca Formations exposed along the Gaillard Cut sec- tion of the Panama Canal (Graham et al., 1985). They will reflect the changes in climate, physi- ography, and vegetation during a 16 Ma period between the late Eocene (ca. 40 Ma) and the early Miocene (ca. 24 Ma; Harland et al., 1982). LITERATURE CITED BARTLETT, A. S. & E. S. BARGHOORN. 1973. Phyto- geo ographic history of the Isthmus of Panama dur- el d past rs 00 "years (a history of Meat e). Pp. 203-299 in Al Graham (edion) Vegetation and d History of Northern Latin America. Elsevier Publ. Co., Amst ens COHEN, A. D. & W. SPACKMAN. 1972. Methods in peat petrology and their application to reconstruc- tion of paleoenvironments. Bull. Geol. Soc. Amer. : 129-142. anal Zone and vi- cinity deer Pap. U. eol. Surv. 244: 1-41. MOREM P.J. 1982 [1983]. Plate tectonic constraints e America and the n the biogeography of ribbean region. Ann. Missouri Bot. Gard. 69: 432- Cowan, R. S. &R M. PoLHILL. 1981. Detarieae. Pp. 117-134 inR. M. Polhill & P. H. Raven (editors), Advances in Legume Systematics. Royal Botanic .P. DAGHLIAN. 1981. Lower Eocene and Paleocene Gentianaceae: floral and palyno- logical evidence. Science 214: 75-77. & M. ZAVADA. 1980. Investigations of angiosperms from the Eocene of North Amer- ica: a new juglandaceous catkin. Rev. Palaeobot. Palynol. 30: 361-370. Croat, T. B. 1976. Flora of Panama —Sapindaceae. Ann. Missouri Bot. Gard. 63: 419-540. Flora of Barro Colorado Island. Stan- ford Univ. Press. Stanford. 1985] DENGO, G. 1973. Estructura geológica, historia tec- tónica y morfológia de América Central. 2nd edi- E US Regional de Ayuda Técnica, A.I.D., éx sae => "D. L. 1973. A paleoclimatic interpretation e Eocene ve E southeastern North Amer- 39-5 n Graham (editor), Vegeta- tion and Vege cal dou) of Nort — Latin merica. Elsevier Pu , Amster: cia ih C. OBYNS. ^1965. ora Rs Pana- — Hippocrateaceae. Ann. Missouri Bot. Gard. - 81 -98. DueENas, H. 1980. Some remarks about the genus Magnastriatites Germeraad, Hopping et Muller, 1968. Rev. Palaeobot. Palynol. 30: 315-317 Dwyer, J. D. Flora of Panama— Rubiaceae. Ann. Missouri Bot. Gard. 67: 1-522 ELiAs, T. S. & A. RoByns. 1975. Flora of Panama— Tad Ann. Missouri Bot. Gard. 62: 61— E s ^ C. 1974. Characteristic Eocene palyno- morphs in the Gulf Coast, U.S.A. Palaeontograph- ica, Abt. B, Paláophytol. 149: 90-111. — & D. L. DitcHER. 1974. Palynology and age P clays exposed in Lawrence Clay Pit, Henry unty, Tennessee. Palaeontographica, Abt. B, Paliophytol. 146: 65-87. FREDERIKSEN, N. O 1980. Sporomorphs from the of Mississippi and western Alabama. Profess. Pap. U.S. Geol Surv. 1084: 1- 974. Coevolutionay patterns in Central American Bignoniaceae. Ann. Missouri Bot. Gard. 61: 728-759. . S. Toms. 1979. Taxonomic implica- tions of Bignoniaceae palynology. Ann. Missouri Bot. Gard. 66: 756-777. GERMERAAD, J. H., d HOPPING & J. MULLER. 1968. Palynology of Tertiary sediments from tropical areas. Rev. Palaeobot. Palynol. 6: 189-348. GOMEZ-PoMPA, A. 1973. Ecology of the vegetation of Veracruz. Pp. 73-148 in Alan Graham (editor), GONZALEZ GUZM Study on n Uppe r Los Cuervos and Mirador Formations E et rem Eocene; Tibü Area, Colombia). E. J. Brill, Lei N. 1973a. Literature on vegetational merica. Pp. S 282 in Alan etational rdam. 73b. iced of the arborescent temperate element in the merican biota. Pp. 223-236 in Alan erdum (edito Vegetation and Vege- rn Latin America. El- tational History of Nort sevier Publ. Co., Amste eiim 1976. Studies i in neotropical agii ei II. ruz, Mexico. New records of Pelliceria (Theaceae/ Pelliceriaceae) in the Tertiary of the Caribbean. Biotropica 9: 48-52. . 1979a. Literature on vegetational history in Latin America. Supplement I. Rev. Palaeobot. Palynol. 27: 29-52. GRAHAM —NEOTROPICAL PALEOBOTANY IV 533 1979b. Mortoniodendron — and Sphaeropteri/T richipteris (Cyatheaceae) in Ce- no ozoic depo f -Caribbean ed n. Missou ri Bot. ‘Gard 66: 572-576. 1980. Morfologia del polen de Eugenia/Myr- cla , (Myrta ceae) y Combretum/Terminalia (Com- bretaceae) en relación a su alcance estratigráfico en el Terciario del Caribe. Biotica 82. Literature on vegetational history in Latin America. Supplement II. Rev. Palaeobot. Palynol. 37: 185-223 1984 [1985]. Lisianthius pollen from the Eocene of Panama. Ann. Missouri Bot. Gard. 71: 987-993 . 1985. Vegetational ipid studies in Panama and adjacent Central America. In W. G. D'Arcy D.C aA (editors The Botany and Nat- ural History of ES a u Historia & . BARKER. 981. _ Palynology and tribal classification in the inioideae. Pp. —834 in R. M. Polhill & P. H. E (edidi) Advances n Legume Systematics. Royal Botanic Gardens, &D D. M. JARZEN. 1969. — sa bici s paleobotany. I. The Oligocene munities of Puerto Rico. Ann. Missouri Bot. "Ga rd. $6: 308- 57. STEWART & J. L. STEWART. 1985. Stud- bearing deposits. Ann. Missouri Bot. Gard. 72: 485-503. HARLAND, W. B., A. V poni P. G. LLEWELLYN, C. A G. PICKTON, A. G. SMITH & R. WALTERS. 82. A Geologic Time Scale. Cambridge Univ. Press, Cambridge HOLDRIDGE, L. R. 1967. Life Zone m Tropical Science Center, San Jose, Cos 5 . GRENKE, W . H. To T Tost, JR. 1971. Se Environments in Tropical Life Zones. Pergamon Press, New York. HowaARnp, R. 1960. P am In J. A. Duke, onaceae. Ann. Missouri 73. The vegetation of the Antilles. Pp. 1- 38 in Alan Graham (editor), Vegetation and Vege- Latin America. El- . 1949. The botany of San Jose Island (Gulf of Panama). Sargentia 8: 1-306. KEATING, R. C. 73. Pollen morphology and rela- tionships of = Flacourtiaceae. Ann. Missouri Bot. Gard. 60: 273-305. LANGENHEIM, ee . L. HACKNER & A. H. BARTLETT. 67. Mangrove pollen at the depositional site of sce Miocene amber from Chiapas, Mexico. Bot. s. Leafl. 21: 289-324. ie E. B. 1969. Miocene pollen and spore flora of Eniwetok Atoll, Marshall Islands. Profess. Pap. U.S. Geol. Surv. 260-II: 1133-1185. MacGiNiTIE. 1972. Development and affinities of Tertiary floras in the Rocky Mo tains. Pp. 147-200 in Alan Graham (editor), Flo- ristics and Paleofloristics of Asia and Eastern North America. Elsevier Publ. Co., Amsterdam. 534 MANNING, W. E. 1960. Flora of Panama— gan: daceae. Ann. Missouri Bot. Gard. 47: MARSHALL, L. G., R. F. BUTLAR, R. E. DRAKE ri G. H. Curtis. 1981. Calibration of ree beginning of the Age of Mammals in Patagonia. Science 212: 3-45. , S. D. WEBB, J. J. SEPKOSKI, JR. & D. M. RAUP. 1982. Mammalian evolution and the great Amer- ican interchange. Science 215: 1351-1357. , R. F. BUTLAR, R. E. DRAKE, G. H. Curtis & 1976. Calibration of the great merican interchange. Science 204: 272 MARTINEZ-HERNANDEZ, E., B. LUDLOW- Wicie RS & . SANCHEZ-LOPEZ. 1980. Catálogo palinológico de la Cuenca Fuentes-Río Escondido. Volume 1 Esporas Monoletes, esporas Tri len sulcados. Comisión Fed. Electricidad (Mexico, , Ser. Tec. 6: 1-234. MULLER, J. 1959. Palynology of Recent Orinoco delta and shelf sediments: reports of the Orinoco Shelf Expedition, 5. Micropaleontology 5: 1-32. 1981. Fossil pollen records m extant angio- sperms. Bot. Rev. (Lancaster) 47: 1-142. & C. CARATINI. 1977. Pollen a Rhizophora (Rhizophoraceae) as a guide fossil. Pollen & Spores 19: 361-389. NicHors, D. J. 1973. North American and European species of Momipites ("Engelhardtia") and related genera. Geosci. & Man 7. PoRTER, D. M. 1970. Flora of Panama — Burseraceae. Ann. Missouri Bot. Gard. 57: 5-27. 973. The vegetation of Panama: a review. Pp. 167-201 in Alan Graham (editor), bianca and Vegetational History of Northern Latin Amer- ica. Elsevier Publ. Co., Amsterdam. RAVEN, P. H. & D. I. AXELROD. 1974. Angiosperm biogeography and past continental movements. Ann. Missouri Bot. Gard. 61: 539-673. Flora of Panama — Flacourtiaceae. 4. ilet etes eranos wou noO- ROMERO, E. J. 7. Polen de gimnosperm ig de la formación Río Turbio (Eoceno). Santa , Argentina. Centro Invest. Recursos Geol. (CIRGEO), Buenos Aires. ANNALS OF THE MISSOURI BOTANICAL GARDEN TscHupy, R. H. & S. D. [Vor. 72 RuBiNorr, R. W. 1974. Environmental Monitoring and apii Data. Smithsonian Inst. Press, Wash- ingto SCHOLL, D. W. 1964. duel nep uat record in mangrove swamps and ri a level over the icu erue Bae of Florida, Part I. Marine Geol. 1: 344-366. MES Recent sedimentary record in man- grov mps and rise in sea level over the south- en facies of Florida, Part II. Marine Geol. 2: 343-364. 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 [map also included in Woodr- ing, 1982]. AN LoEÉNEN. 1970. Illus- trations of plant cn from the Yazoo Clay (Jackson pee upper Eocene) oo Pro- fess. Pap . Geol. Surv. 643-E: 1-5. VAN DER dl N, T. & T. A. “cansa 1964. A Sóc paw E study on the Tertiary and upper Cre- taceous of British Guiana. Leidse Geol. Meded. 30: 183-241. WEAVER, R. E., JR. 1972. A revision of the neotrop- ical genus Lisianthus (Gentianaceae). J. Arnold Arbor. 53: 76-100, 243-311. Wess, S. D. 1976. Mammalian faunal dynamics of the great American interchange. Paleobiology 2: 220-234. WHITEHEAD, D. R. 1965. Pollen morphology in the Juglandaceae, II: a of the family. J. Arnold Arbor. 46: 369-4 WHITMORE, F. C. & 2 H. STEWART. Central A 1965. Miocene y Science 148: 180-185. WooprING, W. P. . The Panama land bridge as barrier. Trans. Amer. Philos. Soc. 110: 425- 433. . 1957-1982. Geology and paleontology of Ca- nal Zone and adjoining parts of Panama. Profess. Pap. U.S. Geol. Surv. 306A-F A NEW CYMBOPETALUM (ANNONACEAE) FROM COSTA RICA AND PANAMA WITH OBSERVATIONS ON NATURAL HYBRIDIZATION! GEORGE E. SCHATZ? ABSTRACT Cymbopetalum torulosum G. E. Schatz, sp. nov., known from northeastern Costa Rica and a single o n b dyn Cy nd are dispersed by ochre- bellied flycatchers a ata redes. The species hybridizes with C. costaricense. describe y se illustre una especie nueva de Ken iila ep nordeste de Costa Rica y de una ta Rica al lado del Pacífico. Es muy fácil de io de los monocarpos toru- losos. Populaciones en la Estací encuentran en mido recientes alluviales Ellos florecen de Febrer por el coleóptero y, y hybride con C. costaricense. o, estan po Ome el pajaro ro Mioneiget oleaginea. La especi Cymbopetalum Bentham is one of the best cir- cumscribed and most easily recognized genera of neotropical Annonaceae. It is characterized by the inner whorl of thick, fleshy, saccate petals. When last revised (Fries, 1931), the genus con- 1974a, 1974b). Another ees species has now been discovered at La a Bi- ological Station of the Organization for Tropical Studies. Cymbopetalum torulosum G. E. Schatz, sp. nov. TYPE: Costa Rica. Heredia: La Selva Biolog- ical Station, at the confluence of the Rio Puerto Viejo and Rio Sarapiqui, 3 km S of the town of Puerto Viejo de Sarapiqui, 10°26’N, 84*00'W, 35 m, 19 July 1984 (fl, fr), Schatz & Grayum 1029 (holotype, MO; isotypes, CR, DUKE, NY, WIS. Arbor usque ad 5 m alta, ramulis novellis sockets ha sassa nie Folia brevipetio ata, pe- ! [ thank Hugh H. Iltis and Kamal S. assistance with the Latin pine i iren i € Ratcliffe for identification of the en Herbarium fund and Ries Apia Studies Program o eed Company, Rochester, New York, and the Jesse Smith Noyes Foundation tiolo applanato tomentello, lamina membranacea bul- lata, elliptica-oblonga ad obovata, 15-22 cm longa, 5- , apice acuminato ad caudato, basi inaequi- fee rotundata ad cuneata, nervis lateralibus utrinque . Flores solitarii, penduli, supra-axillares; pedi- Ben tomentelli, 2-4 cm longi; sepala flavescentia, late ova m longa, 5-8 mm lata, apiculata, tomen- tella; ni. exteriora flavescentia, plana, tenuia, late ovata, mm longa, 16-19 mm lata, apiculata, to- mentella, intus reticulata; odia interiora flavoviren- tia, crasse carnosa, cymbiformia, late ovata ad rotun- data "a rs mm ito. 17-24 mm lata, extus sulcata. Stamina numerosa, 3.5-3.8 mm longa, filamentis 0.4 mm longis, connectivis minute papillatis. Carpella 8— 24, 3m mm longa, stigmatibus connatis; ovula 8-12 d 24, torulosis, 9cm longis, apicibus acutis, ‘stipitibus 10-15 mm lon- gis. Small trees to 5 m tall and 4 cm diam. at breast height. Young branches minutely gray-tomen- tose. Leaves alternate, short-petiolate, the peti- oles 2-3 mm long, flattened, tomentulose; leaf blade membranaceous, conspicuously bullate when fresh, grayish green with a slight sheen, elliptic-oblong to iip - 22 cm long, 5-9 cm wide, the apex long acuminate to caudate, the base slightly [E din. rounded to cu- Bawa for critical review of the manuscript, Duane A. Kolterman for ucy d ylor for assistance with the illustrations, Brett 1, J ames s. and Helen Young for stimulating rt for field work during 1982 a of the tudies. ? Herbarium, Department of Botany, Caivenifs of Wisconsin, Madison, Wisconsin 53706. ANN. MIssourRI Bor. GARD. 72: 535—538. 1985. 536 wers of Costa Rican Cymbopet- —]. . C. torulosum at the end of female receptivity FiGuREs 1-3. Flow iei Pil) —2. C. tor ulosum during early uen receptiv- x cence (note reflexed outer petals). Scale bar = 1 cm neate, the midrib sericeous on both surfaces, es- pecially toward the base, elevated beneath, the 17—22 pairs of major lateral veins slender and Flowers protogynous, acti- the pedicels rigid, minutely tomentose, especially at the base, 2-4 cm long; sepals 3, rarely 4, broad- ly ovate, 4-6 mm long, 5-8 mm wide, apiculate, minutely tomentose; petals 6, rarely 8, in two ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURES 4-6. dns of Pd t Cymbopetal- um.—4. C. torulos —5. icense. —6. Puta- tive C. pese x "C costaricense "hybrid (note the acute apex). Scale bar = 2c strongly differentiated whorls: outer petals closed down at anther dehiscence, appressed to inner petals, green initially, turning yellow at the end of female receptivity, thin, broadly ovate, 19 mm long, 16-19 mm wide, minutely tomentose, the apex acute, apiculate, the inner surface slightly reticulate; inner petals yellowish green, thick, fleshy, cymbiform, broadly ovate to rounded, 26- 36 mm long, 17-24 mm wide, minutely tomen- tose, the margins strongly involute, 2 mm wide, the apex acute, the outer surface strongly sulcate, the inner surface only slightly striated. Stamens numerous, 3.5-3.8 mm long, the filaments 0.4 mm long, the anthers 3.1—3.4 mm long, the con- nectives minutely papillate. Carpels (8—)15—20 (724), 3 mm long, the stigmata fused into a head which abscises as a unit just prior to anther de- hiscence, the ovules 8-12 in a single row. Fruit 1985] TABLE 1. SCHATZ— CYMBOPETALUM 537 Distinguishing characteristics of Costa Rican Cymbopetalum. C. torulosum C. costaricense Young branches tomentulose Leaves elliptic-oblong to obovate, grayish green, with a slight s Leaves with 17—22 pairs of major lateral veins, these strongly impressed, the lamina bullate Flowers solitary, borne midway between nodes bearing full-sized leaves Calyx and corolla tomentulose Outer petels closed at anther dehiscence, appressed to inner Outer petals slightly reticulated on the inner sur- face Inner petals strongly sulcate on the outer surface, only slightly striated on the inner surface Monocarps torulose at maturity, the apex acute, with a single row of seeds Young branches glabrous Leaves elliptic, olive green, very glossy Leaves with 9-12 pairs of major lateral veins, these only slightly impressed, the lamina smooth Flowers borne on new “‘short-shoots,” subtended by small bract-like leaves, thus clustered, and either leaf-opposed or pseudo-termina Calyx slightly tomentulose, corolla glabrate Outer petals usually strongly reflexed at anther dehis- Outer petals strongly striated on the inner surface Inner petals glabrous on the outer surface, striated on the inner surface Monocarps not torulose at maturity, the apex round- ed, with two rows of seeds apocarpous, a cluster of up to 24 monocarps, the appears at maturity along the abaxial surface op- posite the indehiscent carpellary suture; seeds black, ellipsoid, 12 mm long, 7 mm diam., with a thin orange aril (Figs. 1, 2, 4). Distribution. Cymbopetalum torulosum is known from the Río San Juan drainage in north- eastern Costa Rica, both from the Río Sarapiquí region and the Llanuras de San Carlos, and from a single specimen from lowland Chiriquí Prov- ince, Panamá, adjacent to the Costa Rican border along the Pacific coast, an area comparable in rainfall. It is also to be expected in southeastern Nicaragua, as yet a poorly collected area. Ecology. At the La Selva Biological Station of the Organization for Tropical Studies, Cym- bopetalum torulosum is restricted to the most recent alluvial soils bordering the Río Puerto Viejo, Río Sarapiquí, and Río Peje, and slightly FIGURES 7, 8. center, Monocarps of Costa Rican Cymbopetalum. In each figure: left, C. costaricense with rounded pex; putative C. torulosum x C. costaricense hybrid with acute apex = variable morphology of putative hybrid monocarps); right, C. torulosum with acute apex. Scale bar = 538 inland from the rivers along their tributaries and in low-lying swamps. It flowers over a long pe- riod from February to August, during the dry season and early part of I: ied is die with the peak of flowering occurring durin ril and May, and is pollinated by goin ded sparsa Arrow (Scarabaeidae: Dynastinae), attracted to female phase flowers during the early evening hours by a subtle odor reminiscent of linseed oil (unpubl. data). Fruit takes six to seven months to mature before the carpels split open to reveal the orange-arillate seeds. Seeds of C. torulosum are dispersed by birds that feed upon the aril and either pass or regurgitate the seeds. Regurgitated Cymbopetalum seeds have been recovered from an ochre-bellied flycatcher (Mionectes oleagi- nea), an apparent specialist on arillate seeds (D. Levey, pers. comm.). Seeds germinate in ap- sal Sena. four weeks (unpubl. data). Etymology. From the Latin torulosus, mean- ing cylindrical with constrictions at intervals, and ro patric C. costaricense (Donn. Sm.) R. E. Fries (Figs. 3 Specimens examined. CosTA RICA. HEREDIA: La Selva Biological Station, confluence of the Río Puerto Viejo and Río Sarapiquí, 10 May 1982 (fl), Schatz 589 (WIS); 7 Apr. 1982 (fl), Hammel 11607 (CR, DUKE, WIS); 16 July 1981 (fr), Hammel 10990 (DUKE); 14- 17 June 1968 (fl), Burger & Stolze 5747 (F, NY); 26 Apr. 1973 (fl), Burger & Gentry 9248 (F); 27 Apr. 1981 (fl), Folsom 9879 (DUKE, WIS); 25 May 1982 (fl), Hammel 12499 (DUKE, WIS); 23 Apr. 1982 (fl), mmel & Trainer 12753 (DUKE, WIS). ALAJUELA: near Caño Negro, 18-22 km N of Agua Zarcas in Llan- uras de San Carlos (10°31'N, 84?24'W), 60 m, 21 May 1968 (fl), Burger & Stolze 5200 (F). PANAMA. CHIRIQUi: Comarca del Baru, area W of Puerto Armuelles, 100 ft., 15 June 1957 (fl), Stern & Chambers 127 (MAD). There appears to be a natural subdivision within Cymbopetalum between torulose and non- torulose fruited species. Cymbopetalum toru- losum is more closely related to the other tor- ulose-fruited species than to the sympatric C. costaricense, from which it is easily distinguished (Table 1). Within the torulose-fruited group, C. torulosum seems to be most closely related to C. tessmannii R. E. Fries from Peru. It differs from C. tessmannii by having larger leaves with more major lateral veins, larger flowers, and more ovules per carpe ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Determination of Costa Rican Cymbopetalum becomes problematic in light of the discovery of hybridization between C. torulosum and C. cos- taricense at La Selva Biological Station. Cym- bopetalum torulosum is restricted to the most recent alluvial soils; C. costaricense occurs on older alluvium and residual soils. Putative hy- brid individuals have been located in a zone of overlap, where the most recent alluvium grades into older alluvium. Flowering phenology of the two species also differs at La Selva. Peak flow- ering of C. torulosum occurs during April and May, whereas that of C. costaricense occurs in July and August. Nevertheless, overall flowering periods of each species do overlap. Cymbopet- alum costaricense is also pollinated by Cyclo- cephala sparsa, and marked individuals of C. sparsa have been observed to move from one species to another within a 24 hour period. Putative hybrids are intermediate in both veg- etative and reproductive characters, such as the amount of pubescence on young branches and S complete and the other incomplete, and always having an acute apex (Figs. 6-8). The morphol- ogy of hybrid fruits is variable, the monocarps often resembling more closely one or the other of the parental types (Figs. 7, 8). Artificial interspecific crosses have yielded mature fruit and seed (unpubl. data). Seeds from the spontaneous putative hybrid fruit pictured in Figures 6 and 7 germinated successfully. LITERATURE CITED Fries, R. E. 1931. Revision der Arten einiger An- nonaceen Gattungen II. Acta Horti Berg. 10(2): 180-194. . 1952. Contributions to the knowledge of the Annonaceae in veda South America. Arkiv Bot. n.s. 1(6): 329-34 Some new pM to the knowl- edge of the Annonaceae i 1 Vr ES and Mexico. Arkiv Bot. n.s. 3(12): 4 LuNpELL, C. L. 1974a. SEA of American plants VI. nd 5(2): 23-44 74b. Studies of American plants VII. im 5(3): an SCHERY, R. W. Annonaceae. In R. E. Woodson W. dut Contributions Toward a Flora of Panama, V. Ann. Missouri Bot. Gard. 28: 427- 429. NOTES ON AND DESCRIPTIONS OF SEVEN NEW SPECIES OF MESOAMERICAN CLETHRACEAE! CLEMENT W. HAMILTON? ABSTRACT Taxonomic confusion in Mesoamerican Clethraceae, caused by important differences between Sleu- mer's 1967 monograph and the 1966 “Flora of Guatemala" treatment by Standley and Williams, is resolved. Seven new species of Clethra are —— also: three from Mexico, Clethra breedlovei, C. oaxacana, and C. michoacana; one from Nica agua, C. nicaraguensis, one from Costa Rica, C. tala- mancana; and two from Panama, C. tutensis di C. coloradensis. The genus Clethra Gronov. ex L. comprises approximately 70 species, of which about 40 are neotropical (Sleumer, 1967). Sleumer (1967) wrote his monograph of the genus at the same time that Standley and Williams (1966) prepared their treatment of Clethraceae for the “Flora of Guatemala." Standley and Williams (1966: 74) stated that “many of Dr. Sleumer's annotations are accepted here but others are not." The dis- agreements and resulting taxonomic confusion involve several widespread species that are treat- ed under different names in the two treatments. In an earlier work on the family, Britton (1914) recognized many more distinct species than sub- sequent authors have thought warranted (Stand- ley & Williams, 1966; Sleumer, 1967; Robertson, 1968) In preparing Clethraceae for “Flora de Nica- ragua" (Hamilton, in prep.), I have attempted to reconcile these differences in order to identify the many specimens collected since 1965. The major differences between Sleumer (1967) and Standley and Williams (1966) are considered be- low, and recommendations and m are e taxo- nomic problems a are especially Eu be- cause the species in question account for ap- proximately 7096 of Central American material of Clethra. Complete keys to the Mesoamerican species will be included in the “Flora Mesoamer- icana" treatment of the family (Sutton, in prep.). I examined 75 sheets at MO, the duplicates of which were cited by Sleumer, especially closely. Additional material from F, HNMN, and MO was studied. LEGITIMACY OF CLETHRA OCCIDENTALIS Standley and Williams (1966) doubted the le- gitimacy of the combination Clethra occidentalis (L.) O. Ktze., noting that Kuntze (1898) did not cite the basionym. As Sleumer (1967) correctly pointed out, however, the transfer was legiti- mately accomplished earlier by Kuntze (1891), who clearly cited the basionym, Tinus occiden- talis L. Steudel (1821) was actually the first to use the name Clethra occidentalis, but he did so without citing basionym or authority. THE CLETHRA MACROPHYLLA GROUP Standley and Williams (1966) treated Clethra macrophylla Mart. & Gal., C. vicentina Standl., and C. occidentalis (L.) O. Ktze. as one species, C. macrophylla, whereas Sleumer (1967) recog- nized them as separate entities. The three species appear to be closely inter- related: they share grey-brown inflorescences, usually entire leaf margins, and usually striking color differences between the dark upper surfaces ever, by its glabrous leaf blades which often dry grey-green instead of dark brown above. Fur- thermore, material collected of both C. vicentina and C. occidentalis in Honduras and Nicaragua shows no evidence of intergradation relative to the characters mentioned above. The distinction between C. macrophylla and C. occidentalis is more problematic. The former has longer leaf blades (15—20 cm versus 7-12 cm) and thicker, more reddish pubescence on the leaf ! I thank the staff of the Missouri Botanical Garden, particularly Alwyn H. Gentry, John D. Dwyer, and Amy Pool, for their assistance; and Peter F. Stevens, John W. Thieret, and Nancy R. Morin for helpful comments on the manu uscript. ? Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. ANN. Missouni Bor. GARD. 72: 539-543. 1985. 540 undersides. The two species occur only allopat- rically, however, and careful field study may re- veal that they represent one variable species. occurs in Ede Honduras, and Nica aragua. If they are to be considered one species, the correct name is C. occidentalis (L.) O. Ktze. (the specific epithet published by Linnaeus in 1759) rather than C. macrophylla Martens & Galeotti (1842), the name used by Standley and Williams (1966). Standley and Williams (1966) did not mention C. hartwegii Britt. as growing in Guatemala. It is a chiefly Mexican species that also reaches Guatemala and is related to the species consid- ered above. It may be distinguished from them by its short (3-4 mm), robust pedicels and the deep rust-colored tomentum on the inflores- cences. Another Mexican species, C. pringlei Watson, also belongs in this species group. THE CLETHRA LANATA GROUP Standley and Williams (1966) treated C. mex- icana A. DC. and C. lanata Mart. & Gal. as conspecific and ee the penne name. They are, however, distinct species. Clethra mexicana, which appears to be solely Mexican, has inflo- rescence rachises that are extremely stout, fur- rowed, and deep red-brown, and the fruit is large (5-6 mm diam.). Clethra lanata, a variable species found from Mexico to Panama, has more slen- der, pale brown rachises and smaller fruit (3 mm iam.). I differ with Sleumer (1967), however, on the status of one species in this group. Clethra rosei Britt. appears quite distinct and striking in Mex- ico, with its upper leaf surfaces scabrous to pu- bescent and its styles usually longer than those of C. lanata sensu stricto. The apparent distinc- tion between these two species breaks down, however, because the degree of upper leaf pu- bescence and style length forms a continuum. Therefore C. lanata should include C. rosei. CLETHRA OLEOIDES AND CLETHRA GELIDA Clethra oleoides L. Wms. and C. gelida Standl. appear to be closely related. They are unique in having small leaves and peduncles that curve downward and are collected usually above 2,800 m elevation, in elfin forest. I can find only one good character to differentiate the two: the leaf underside texture of C. gelida is ferrugineous, whereas that of C. oleoides is rough and glandular ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 but not pubescent. Clethra oleoides occurs in Mexico and Guatemala, whereas C. gelida oc- curs in Costa Rica; presumably the present-day paucity of suitable habitat between their ranges has resulted in the isolation of populations and consequent speciation. THREE NEW SPECIES FROM MEXICO Clethra breedlovei C. Hamilton, sp. nov. TYPE: Mexico. Guerrero: ENE of Puerto El Gallo, 2,700-2,850 m, 11 Nov. 1973 (fl), Breedlove 36098 (holotype, MO-2243945; isotype, CAS) . 10 m alta, d t f ] rbor c lato- we ss anu Folia petiolis 1.3-1.8 cm rper lami- nis 7-12 cm longis, 2.5-5.5 cm latis, supra brunneo- stellato- pubescentibus sulcis nerviorum mediorum infra dense pal- lido- domentotis atque ferrugineo-stellato- -pubescenti- bus, venis secundariis 12-15 infra porphyreo-stellato- pubescentibus. Inflorescentiae terminales paniculatae contractae, ramis 6-10, ramis maturis (5-)6-8 cm lon- gis, bracteolis usque ad 6 mm longis. Flores pedicellis mm longis; sepala linearia, 4-5 mm longa; nga; stamina filamentis 2.5 mm longis, antheris 1.5 mm longis; stylus usque ad 3 mm longus. Fructus non visi. tellato O-STeHAato Tree ca. 10 m; young branchlets densely deep brown setose and stellate-pubescent. Leaves with petioles 1.3-1.8 cm long, densely deep brown setose and stellate-pubescent, terete; blade rigid, elliptic, the apex acute or mucronulate, the base cuneate to obtuse, the margin entire or obscurely denticulate, not revolute at base, 7-12 cm long, 2; wide, green-yellow and regularly brown stellate-pubescent above with midvein furrow densely brown stellate-pubescent, densely pale tomentose and red-brown stellate-pubes- cent below; secondary veins 12-15 pairs, not furrowed above, prominulous below, rusty stel- late-pubescent below. Inflorescences terminal contracted panicles with 6-10 branches, the ma- ture branches (5-)6-8 cm long; rachises densely deep brown setose and stellate-pubescent; brac- teoles to 6 mm long, linear, caducous. Flowers with pedicels 3-4(-5) mm long; sepals 5, linear, 4-5 mm long, tomentose without; petals 5, entire or short-fringed, 5.5-6.5 mm long; stamens 10, the filaments 2.5 mm long, the anthers 1.5 mm long; ovary pale tomentose, the style to 3 mm ong. Fruit not seen. This species is probably related to C. mexi- cana, with which it shares similar stout dark brown inflorescence rachises. Clethra mexicana 1985] leaf blades are obovate to oblong and essentially glabrous above, whereas C. breedlovei leaf blades are elliptic and pubescent above. Clethra oaxacana C. Hamilton, sp. nov. TY Mexico. Oaxaca: road between Teotitlán He Camino and Huatla de Jiménez, 35 mi. E of Teotitlán, 1,650 m, 22 Feb. 1979 (imm. fl), Croat 48252 (holotype, MO-2981877). or ca. 10 m alta, dense rufo-stellato- tomentosa. nales, racemis 6—10 cm longis, bracteolis 2 mm longis. Flores pedicellis 3-4 mm longis; sepala linearia, 4 mm longa; pe onga; stamina filamentis ca. 2 mm longis, antheris 1.2 mm longis. Fructus non visi. Tree ca. 10 m; young branchlets densely red- brown stellate-tomentose. Leaves with petioles 1.5-2 cm long, densely brown stellate- pubescent, terrete; blad , elliptic the apex acute to cuspidate, ‘the tee obtuse to truncate, the margin denticulate, not revolute at base, 10-14 cm long, 4.5-7.5 cm wide, indu- mentum stellate, deep brown regularly setose above with vein furrows densely brown setose, red-brown setose below; secondary veins 9-12 pairs, shallow furrowed above, prominent below, densely red-brown tomentose below. Jnflores- cences terminal clusters of 5-8 racemes, the ra- cemes 6—10 cm long; rachises densely red-brown stellate-tomentose; bracteoles 2 mm long, to- mentose, linear, caducous. Flowers with pedicels 3-4 mm long; sepals 5, linear, 4 mm long, dense- ly tomentose without; petals 5, fringed, ca. 4 mm long; stamens 10, the filaments ca. 2 mm long, the anthers 1.2 mm long; ovary velutinous. Fruit not seen. V UV YG LV, This species appears related to C. alexandri Griseb., a Jamaican species with long (ca. 10 mm) bracteoles and larger flowers. Clethra oaxa- cana is related also to C. michoacana C. Ham- ilton (see following species description). Clethra michoacana C. Hamilton, sp. nov. TYPE: ,000 fl), Oliver et äl 941 (holotype, MO- 1099851: isotype, BM) Arbor usque ad 15 m alta, dense ferrugineo-stellato- tomentosa. Folia petiolis 2.5—4 cm longis, laminis 15- 18 cm longis, 8-10 cm latis, supra glabris sulcis vena- HAMILTON -—CLETHRACEAE 541 rum rufo-stella to-tomentosis infra pallido -incanis atq ue ferrugineo- stellato-tomentosis, venis secundariis 13- gis, robustis; sepala ovato-linearia, minimum 4 mm longa. Fructus non visi Tree to 15 m; young branchlets densely red- brown stellate-tomentose. Leaves with petioles 2.5-4 cm long, densely red-brown stellate-to- mentose, terete; blade rigid, obovate, the apex obtuse, the base obtuse, the margin entire, not revolute at base, 15-18 cm long, 8-10 cm wide, green-brown and glabrous above with vein fur- rows red-brown stellate-tomentose, pale pannose and red-brown stellate-tomentose below; sec- ondary veins 13-16 pairs, furrowed above, prominent below, densely red-brown stellate-to- mentose below. Inflorescences terminal clusters of 5-7 racemes, the racemes 7-12 cm long; ra- chises densely red-brown stellate- tomentose; bracteoles 9-13 mm flow sepals 5, ovate-linear, at least 4 mm long, red- brown tomentose without; petals 5; stamens 10; ovary velutinous. Fruit not seen. This species differs from the related C. oaxa- cana C. Hamilton in having more secondary veins, lacking setae on the upper leaf surface, and having longer, more persistent bracteoles. Cleth- ra michoacana differs from the Jamaican C. al- exandri Griseb. in having longer leaf blades (1 5- 18 cm versus 8-12 cm more secondary veins (13-16 versus 8-12). A NEW SPECIES FROM NICARAGUA Sleumer (1967) cited only one Nicaraguan specimen in his entire treatment; intensive col- Jecting since then has provided much more ma- terial, including a new species related to C. /ana- ta. Clethra nicaraguensis C. Hamilton, sp. nov. TYPE: arag Jinotega, _ Feb. 1979 (fl), Grijalva & Araquistain 166 (holotype, MO-3103450; isotypes, BM, CR, HNMN, MBM, MEXU, PMA, TEX, U, US, VEN, XAL) Arbor 3-10 m alta, atrorufo-velutina. Folia petiolis (1-)1.5—2 cm longis, laminis 8-13 cm longis, 3.5—6 cm latis, supra glabris sulcis venarum sparse stellato-pi- losis, infra rufo-lanatis atque stellato-pilosis, venis se- cundariis (9—)10—12(-14) infra atroferrugineo-villosis. 542 Racemi 4-12 fasciculati fermen. et saepe racemis solitariis in 13-22 cm longis, bracteolis Cs mm longis. Flores n 2-4(-5) mm longis; sepala ovato-linearia, 4 onga; petala 4 mm longa; stamina filamentis 2 mm s. antheris 1 mm longis; stylus usque ad 2.5 mm longus. Fructus 4 mm longi, 4 mm diam Tree 3-10 m; young branchlets densely deep red-brown velutinous. Leaves with petioles (1-)1.5-2 cm long, densely deep rusty velutinous, often flattened above; blade coriaceous, obovate, the apex obtuse to subacute, the base cuneate, the margin entire or rarely denticulate near apex, revolute at base, 8-13 cm long, 3.5-6 cm wide, dull brown and glabrous above with vein furrows sparsely stellate-pubescent, pale red-brown woolly and stellate-pubescent below; secondary veins (9—)10-12(-14) pairs, furrowed above, prominent below, deep rusty villous below. Zn- florescences terminal clusters of 4—12 racemes, often also with axillary racemes in distal 3-4 nodes, the mature racemes 13-22 cm long; rach- ises densely deep rusty velutinous; bracteoles 3—5(—6) mm long, linear, caducous. Flowers with pedicels 2-4(-5) mm long; sepals 5, ovate-linear, 4 mm long, tomentose within and without; petals 5, fringed, 4 mm long; stamens 10, the filaments 2 mm long, the anthers 1 mm long; ovary ve- lutinous, the style to 2.5 mm long. Fruit a 3- lobed loculicidal capsule, 4 mm long, 4 mm diam., 7 mm diam. after dehiscence. Distribution. Endemic to Nicaragua, 1,200- 1,600 m. Flowers mostly from December to March, and fruits mostly from April to June. Additional specimens examined. NICARAGUA. JINOTEGA: rH de Matagalpa-Jinotega, 1,400-1,500 m, 2 July 1980 (fl), Moreno 1108 (MO); 20 Aug. 1980 (fr), Moreno 1874 (MO); Cerro Zamaria, 1,450 m, 5 July 1975 (fl), Atwood & Neill 97 (BM, MO); Volcán Yalí, (i 500—1,542 m, 9 Apr. 1981 (fr), Moreno 7924 ; Montaña Cuspire, 1,500-1,539 m, 10 Apr. 1981 (fr), Moreno 8029 (BM, HNMN, MO). MATAGALPA: Cerro El Picacho, 1,500 m, 11 Feb. 1965 (fl), Williams et al. 29181 (MO); finca Sta. María de Ostuma, 10 km N of Matagalpa, 12 Aug. 1977 (fr), Neill 2312 (MO); El Porvenir, 1,700 m, 11 Mar. 1967 (fl), Molina 20514 (MO). RIvAs: Ometepe, NE of Vol- Concepción, 13 Mar. 1981 (fl), Sandino d (BM, O). ZELAYA: Cerro El Hormiguerro, 1,100-1,183 m, 15 Apr. 1979 (fr), Pipoly 5173 (BM, CR, HNMN, MEM MO). Clethra nicaraguensis is distinguished from C. lanata, probably its closest relative, by its deep rusty pubescence on the leaf blade undersides and rachises and by its smaller flowers and fruits. Clethra nicaraguensis superficially resembles C. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 mexicana in having a dark-pubescent, sturdy in- florescence but differs in having smaller fruit (4 mm versus 5-6 mm diam.), much less robust rachises, and shorter leaf blades (8-13 cm versus 10-16 cm long) A NEW SPECIES FROM COSTA RICA Clethra talamancana C. Hamilton, sp. nov. TYPE: Costa Rica. Cartago: SE slope of Cerro de la Muerta, Cordillera de Talamanca, along Interamerican Highway, 2,700 m, 23 May 1976 (fl), Croat 35400 (holotype, MO- 3099952; isotype, BM). Arbor ca. 3 m alta, rufo- nen Folia petiolis 1. 33 cm longis, laminis (5-)6— m longis, 2-4 cm latis, indumento stellato, esa een O-pu- venarum e pu ra florescentiae terminales dp pies cont is ca. i s ca. 1.5 mm longus. Fructus non visi. Tree ca. 3 m; young branchlets red-brown stel- late-pubescent. Leaves with petioles 1.3-2 cm long, densely red-brown stellate-pubescent, te- rete; blade subcoriaceous, elliptic-obovate, the apex mucronate to acuminate, the base cuneate, the margin mucronate-denticulate throughout, not revolute at base, (5-)6-10 cm long, 2-4 cm wide, indumentum stellate, dull brown and reg- own t ana shallowly furr below, red-brown ea 1. nflores- cences terminal contracted pande of ca branches, the mature branches 8-12 cm long; rachises red-brown pubescent; bracteoles 2-3 mm long, linear, caducous. Flowers with pedicels 3—4 mm long; sepals 5, linear, 3-3.5 mm long, pu- berulent without; petals 5, fringed, 4 mm long; stamens 10, the filaments 1.2 mm long, the an- thers 0.8 mm long; ovary pubescent, the style ca. 1.5 mm long. Fruit not seen. The dense red-brown pubescence, small leaves, and short sepals distinguish C. talamancana from other Clethra species. As more material is col- lected from high elevations in Costa Rica and Panama, perhaps additional new species will be found to shed light on the presently unknown 1985] affinities of this and the following two species from Panama. Two NEW SPECIES FROM PANAMA Clethra tutensis C. Hamilton, sp. nov. TYPE: Panama. Veraguas: Cerro Tute, ca. 10 km NW of Santa Fe, on ridgetop in cloud forest, over 1,000 m, 3 Aug. 1975 (fr), Mori et al. 7574 (holotype, MO-3099953; isotypes, BM, MEXU, PMA). Arbor ca. 7 m alta, brunneo-adpresso-setosa. Folia petiolis 0.5-1 cm longis, laminis 2.5—3.5(—4) cm longis, 1.4-2.2 cm latis, supra glabris sulcis venarum glabris, infra glabris, venis secundariis 7—9 infra rufo-adpresso- setosis. Racemi 5-10 fasciculati terminales, racemis maturis (6-)7-10(-1 1) cm longis, bracteolis non visis. Flores non visi. Fructus 4 mm longi, 4-5 mm diam ca. 7 m; young branchlets brown ap- d blade ja NUR elliptic to slightly basate: the apex mucronate-acute, the base cuneate, the margin mucronate-denticulate throughout, not revolute at base, 2.5-3.5(-4) cm long, 1.4-2.2 cm wide, dark brown and glabrous above, pale brown and glabrous below; secondary veins 7— 9, furrowed and glabrous above, prominent and red-brown appressed-setose below. [nflores- cences terminal clusters of 5-10 racemes, the ma- ture racemes (6—)7—10(—1 1) cm long; rachises red- brown puberulent; bracteoles not seen. Flowers not seen. Fruit a 3-lobed loculicidal capsule, 4 mm long, 4-5 mm diam., 6-7 mm diam. after dehiscence. The small leaves, large fruit, and clustered in- florescences distinguish C. tutensis from all other species of Clethra; its affinities are unknown. Clethra coloradensis C. Hamilton, sp. nov. TYPE: Panama. Chiri 3099954; isotypes, BM, MEXU, PMA) bor 8-12 m alta, dense rufo-stellato- oe Folia petiolis 0. 8-1. 5 em longis, laminis 4-6. cml ongis, brescentibus sulcis venarum dense puberulis, infra pal- lido-incanis atque ferrugineo-puberulis, venis secun- dariis 8-10 infra ferrugineo-puberulis. Racemi 6-8 fasciculati terminales, racemis maturis 5-8 cm longis, bracteolis 2.5-3 mm longis. ipn pesi > 5-1 mm longis; sepala ovata, 3 mm . 3 mm longa. Fructus 2-2.5 cm longi, 3 di HAMILTON —CLETHRACEAE 543 Tree 8-12 m; young branchlets densely red- brown stellate-tomentose. Leaves with petioles 0.8-1.5 cm long, densely red-brown stellate-to- mentose, terete; blade rigid, obovate, the apex mucronulate-obtuse, the base cuneate, the mar- gin entire, not revolute at base, 4—6 cm long, 2.5- 4 cm wide, indumentum stellate, brown puber- ulent becoming glabrous above with vein fur- rows densely puberulent, pale pannose and red- brown puberulent below; secondary veins 8-10 pairs, shallow furrowed above, prominent below, red-brown puberulent below. Inflorescences ter- minal clusters of 6-8 racemes, the mature ra- cemes 5-8 cm long; rachises red-brown tomen- tose; bracteoles 2.5—3 mm long, linear, caducous. Flowers with pedicels 0.5-1 mm long; sepals 5, ovate, 3 mm long, puberulent within and with- out; petals 5, fringed, ca. 3 mm long; stamens 10; ovary tomentose. Fruit a 3-lobed m capsule, 2-2.5 cm long, 3 mm diam., 4 mm after dehiscence. Additional specimens examined. |. PANAMA. CHIRIQUÍ: N of San Felix at Chiriquí-Bocas del Toro border, on Cerro Colorado copper mine road along continental divide 5,000—5,500 ft., 3 May 1975 (imm. fl), Mori & Kallunki 5812 (MO, PMA). The small, stellate-pubescent, and puberulent leaves, relatively short racemes, minute pedicels, and small fruit distinguish C. coloradensis from all other neotropical species of Clethra; its affin- ities are unknown LITERATURE CITED BRITTON, N. L. 1914. Clethraceae. In N. L. Britton, W. A. Murrill & J. H. — (editors), North merican Flora 29(1): 3-9. KuNrzE, O. 1891. Revisio Generum Plantarum 2. Arthur Felix, Leipzi 1898. E Generum Plantarum 3(2). Ar- thur Felix, Leipz LINNAEUS, C. 1759. j O Naturae. 10th edition. Stockholm. MARTENS, G. M. von & H. G. GArEorri. 1842. No- tice sur les plantes des familles des Vacciniées et des Ericacées. Bull. Acad. Roy. Sci. Bruxelles 9(1): 5 KI R. 1968. Clethraceae. In R. E. Wood- n & R. W. Schery, Flora of Panama. Ann. Mis- ouri Bot. Gard. 54(3): 389-392. E H. 1967. Monographia Clethracearum. Bot. Jahrb. Syst. 87: 36-116. STANDLEY, P. C. & L. O. WILLIA 1966. Clethra- ceae. In Flora f Guatemala. Fieldiana, Bot. 24(8): 74-81. STEUDEL, E. G. von. 1821. Nomenclator Botanicus 1: 207. Stuttgart. BUMELIA RECLINATA VAR. AUSTROFLORIDENSIS (SAPOTACEAE), A NEW VARIETY FROM SOUTH FLORIDA, U.S.A.! R. DAVID WHETSTONE? ABSTRACT Taxonomic studies in the Saptotaceae of the Southeastern United States support the recognition of three infraspecific taxa of Bumelia reclinata (Michaux) Ventenat: var. reclinata, var. rufotomentosa (Small) Cronquist, and var. austrofloridensis Whetstone, var. nov. A key to the infraspecific taxa and distribution map are included. Bumelia reclinata is endemic to the outer revised by Asa Gray (1886), J. K. Small (1900), and R. B. Clark (1942) who primarily studied the North American taxa and by Arthur Cron- quist (1945) who studied the New World rep- resentatives. Bumelia reclinata (Michaux) Ventenat com- prises three varieities, i.e., var. reclinata, var. rufotomentosa (Small) Cronquist, and var. aus- trofloridensis Whetstone. Of the more recent treatments, Clark (1942) recognized B. rufoto- mentosa and B. reclinata whereas Cronquist (1945) considered the two taxa as conspecific and relegated B. rufotomentosa to varietal status un- der the latter taxon. My conclusions are most similar to those of Cronquist but recognize the existence of a third variety that was overlooked by previous ride - E Cronquist; Clark, and Small wh eral specimens of var. aus- irofloridensis); Sd js to the paucity of ma- terial available at that time. Those specimens studied by Small, Cronquist, and Clark were con- sidered to belong to var. reclinata. The primary bases for my conclusions are close examination of almost 3,000 exsiccatae of Bumelia and field observations on all U.S. species. Individuals of B. reclinata from the vicinity of Long Pine Key in the Florida Everglades are con- sistently and clearly separable from other taxa of the species by the following suite of character tates: a. leaves persistently tawny-pubescent across the abaxial surface of the lamina; . twigs essentially SPSS or glabrate; pedicels pubescent inner and outer sepals pubescent; ovary pubescen fruits basally a NN and fruits not exceeding 9 mm in length. m m o n. ° = (See Table 1 for summary of key characteristics of all varieties.) Variety austrofloridensis is com- mon in hammocks and slash pine flatwoods in the Florida Everglades. The known range exten ds localities are within the Everglades National Park. As far as I have determined, no other members of B. reclinata are sympatric although B. celas- trina H.B.K. and B. salicifolia (L.) Swartz occur on the Key (Fig. 1). Due to the limited geographic distribution and rarity of this taxon, I offer the name “Everglades Buckthorn” for those individ- uals and governmental agencies dealing with deed and endangered species. KEY TO THE INFRAsPECIFIC TAXA OF B. RECLINATA la. Abaxial surfaces of fully expanded leaves per- sistently pubescent across the lamina; mp essentially glabrous; NR inner and o sepals strigose seer eae lb. Abaxial geris of eat nied leaves gla- along the midrib; twigs ae strigose ses to densely pubescent; pedicels, inner and outer sepals glabrous to densely strigose. ! The dui i d way esp dE the loan of specimens from the herbaria cited herein (acronyms are from l., 1 xcep U,” vide I outgrowt a treatment of the Sapotaceae fo n addition, support for this study was provided S i i ith field work o Radford, elu cds editor, in prep. Univ. of North Carolina Press, Chapel Hill). Descriptions generally follow the preferred format set forth by the editorial committee. ? Herbarium, Department of Biology, Jacksonville State University, Jacksonville, Alabama 36265. ANN. MISSOURI Bor. GARD. 72: 544—547. 1985. 545 1985] WHETSTONE—BUMELIA RECLINATA VAR. AUSTROFLORIDENSIS TABLE 1. Comparison of key characteristics within varieties of Bumelia reclinata. rufotomentosa reclinata austrofloridensis Twigs densely strigose Twigs glabrous Twigs essentially glabrous Leaf trichomes coppery Leaf trichomes tawn trichomes tawny f pubescence along abaxial Leaf pubescence along abaxial Leaf pubescence across the abaxial surface of midvein surface of midvein surface Outer sepals and pedicels dense- Outer sepals and pedicels gla- Outer sepals and pedicels pubes- cent ly pubescen brous Ovaries densely strigose at an- thesis Mature fruits ca. 11 mm long Mature fruits ca. 9 (or more less) Ovaries sparsely strigose at an- thesis Ovaries densely strigose at anthe- Mature fruits ca. 9 mm long (or less) mm long (or 2a. Leaves coppery pubescent on the peti- oles and along the abaxial surface of the midvein; twigs densely ig pedicels, inner and outer sepals densely pubescent var. rufotomentosa 2b. Leaves rufous to tawny-pubescent (whit- ish) on the petioles and abaxial surface of the midvein (glabrate); twigs sparsely strigose to glabrous; pedicels and outer sepals glabrous, inner sepals glabrous to pubescent var. reclinata Bumelia reclinata (Michaux) Ventenat var. aus- trofloridensis Whetstone, var. nov. TYPE: United States. Florida: Dade County, Long TE H- Fr q k= ECES T LIA E ™ Ud A YY L - TNE n LE TIO AIEO EY pe Minus En. COH ede sor ARPES Eh | i CES DPI A IEEE ^ LETRAS Aw Hed x MIL qu A f DAS i Le Li I - sub / 7 H E ZINS AN, HN EN 2 eia ANTA os? REDE T Pre ER TRE T HTEH-UL "E ref PIES U TAHHA AIN CARN 2277 2: gs Í FHT L: zi < STOREY AERA x° TITI CUT DLL Eae ee LO Saa MUERTE AEO + Ae a] jon N! HAUS < ES LUE ELLECI EH DO T i HERA (UNE THERE WEM " C hen "m XXE BW) + MEET WA T > UON MILES WA H4 | w^ o S. 200 300 juri NS, 959 FIGURE 1. rufotomentosa; and star — var. austrofloridensis. Map of the documented distribution of Bumelia reclinata. Triangle = var. reclinata; cross = var. 546 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 FIGURE 2. Bumelia reclinata var. austrofloridensis— A. Branch. — B. Fruit. — C. Adaxial view of androecium and corolla. — D. Gynoecium at anthesis. — E. Leaf details (blade). 1985] Pine Key in the Everglades National Park, recently burned glade; over oolite. Abun- dant shrub, to ca. 1.5 m (isotypes collected from the same individual as holotype). 7 July 1984, Whetstone 14459 (holotype, JSU 23555; isotypes, to be distributed to A, FLAS, MO, NCU, USF, VDB). Ev- erglades Buckthorn. FIGURE 2. arietas nova B. reclinata optime distinguitur lobis calycis strigosis, ramunculo sparse strigoso ad glabres- cens, ovario strigoso, et foliaris pubens persistens. Small thorny shrubs. Twigs strigose, armed tially glabrous. Leaves evergreen, alternate to fascicled, oblanceolate, 1-4 cm long, fully ex- panded leaves with tawny pubescence on the abaxial surface of the lamina, apices rounded to emarginate, margins entire and somewhat revo- lute, bases acuminate with attenuate lamina forming an adaxial channel; petioles 2.5-3 mm long, densely pubescent basally. Flowers borne in loose, axillary fascicles at leafy and defoliated nodes. Flowers ca. 2.8 mm long, calyx of 5 im- bricate sepals, sepals ovate, ca. 2 mm long , densely with lateral appendages, margins erose; stamens epipetalous, anther slightly exerted, staminodes petaloid; gynoecium ca. 1.5 mm long, ovary WHETSTONE—BUMELIA RECLINATA VAR. AUSTROFLORIDENSIS 547 densely vias style and stigma glabrous; ped- icels 4-5(-9) mm long, pubescent apically with rufous DIOE Fruits obovoid, ca. 9 mm long (excluding the apiculate stylar remnant), slightly pubescent basally and occasionally apically. Seeds ovoid, light brown, ca. 7 mm long, scar basilat- eral with a smaller adjacent scar. Specimens examined. UNITED STATES. FLORIDA: Dade Co., Bessey 2 (A), 75 (A), Craighead, s.n. (FLAS 146613, 146614, 146617), Godfrey et al. 63394 (FSU 86781, NCU 253411), 63460 (FSU 86487), 77025 (FSU 154085), Hill & Harvey, III, 3215 (NCU 462019), Korsakoff, s.n. (FLAS 42452), Rehder 898 (A), Small & ri 2912 (NY), 2986 (NY), 2988 (NY), 2988 (NY), 2988a (NY), Whetstone 14427 (JSU), 14470 (JSU), p (JSU). LITERATURE CITED CLARK, R. B. A revision of the genus Bumelia in the UA States. Ann. Missouri Bot. Gard. 29: 2 155-182. CRONQUIST, A. 45. Studies in the Sapotaceae, III. Dipholis and Bumelia. J. Arnold Arbor. 26: 435- 471 GRAY, A. 1886. Synoptical Flora of North America. Revised edition. Ivison, Blakeman, Taylor, and Company, New Ri. k. HOLMGREN, P. K., W. KEUKEN fn E. K. ScHoriELD. 1981. Index MM ais rt I. The Herbaria of the World. 7th edition. shi Sheltema and 1900. The genus Bumelia in North America. Bull. New York Bot. Gard. 1: 437-447. WHETSTONE, R. D. 1983. JSU. Herb. News 3: 49. FIVE NEW TAXA OF NEW WORLD MEMECYLEAE (MELASTOMATACEAE)! THOMAS MORLEY? ABSTRACT new species and one subspecies of Mouriri are described, with one new species of Votomita. rir a an elevated regions of southeast Pará and north Mato Grosso, Brazil, and M. viridicosta is from French uiana. Relations of the firs t three are clear but M. viridicosta has no close relatives and does not fit well in um of the EN sections of the genus. Mouriri cearensis subsp. carajasica is a distinct inland form of the specie New species of the Memecyleae (Melastoma- taceae) from the New World tropics are still dis- covered occasionally, usually where little-col- — regions are explored more intensively but metimes as with M. viridicosta through con- aaa collecting in a relatively well-known country like French Guiana. The tribe is known well enough by now so that new discoveries can usually be placed to genus, subgenus, and section with certainty, but sometimes an exception ap- pears, and M. viridicosta is one of these. This species has no known close relatives and its sec- tional affinities are in doubt. e other three species present no real problems. Mouriri peru- viana is a western relative of M. nervosa, a wide- spread species of Amazonia. Mouriri pranceana is a close relative of M. huberi recently discov- ered through explorations of the Serra do Ca- chimbo and nearby regions; the new species is most readily distinguished by its fruit. Votomita pubescens is a seventh species of that genus, from farther west than the other species; it is the only known member of the tribe in the New World with a pubescent lamina. Mouriri peruviana Morley, sp. nov. TYPE: Peru. pto. Huá lo largo del Río Pachitea cerca del campa- mento Miel de Abeja (1 km arriba del pue- blo de Tournavista o unos 20 km arriba de la confluencia con el Río Ucayali). En bos- que alto en el caserio Miel de Abeja a 2 km del campamento de Iparía. Altura sobre el mar 300—400 m. Arbusto 4—6 m, flores blan- cas con sépalos de color amarillo palido. *Sachavaca Shahuinto." 23 June 1967, Schunke V. 2077 (holotype, F; isotype, US). FIGURE 1. Frutex vel arbor usque 15 m alta; fola 11-25 cm | mm lon- find : In} gus calyx 1.5 “es pen lo * 1.5-2.5 mm en ab staminibus, 3.5- 5mm ost anthesin; eundi 4.3-4.5 mm distan € 351 basi styli ad stamina 3.6—4.8 m ovarii n l ate e separati ps eres ii basim hypanthii; plac parietales in quoqu e loc n fructus 9.5- 14 mm B e semina 9-12 mm Shrub or tree to 15 m high; young twigs round- ed-quadrangular, not winged. Petioles 2-4 mm long; blades coriaceous, intensely green or yel- lowish green, 11-25 cm long, 5.7-8.5 cm wide, oblong, oblong-elliptic, or oblong-ovate, abrupt- ly acuminate at the apex, rounded to cordate at base with a notch 2-8 mm deep; midrib plane above, prominent below with a flattened lower surface and short wings; lateral nerves obscure to moderately prominent above and below when dry. Midrib xylem tubular; stomatal crypts Type III (see Morley, 1976), averaging in a leaf 42-52 um diam., 23-28 um high, 95-108 per mm? (ex- tremes 31-71 um diam., 20-32 um high, 89-141 mm?2); upper epidermis one cell thick, most of the cells with mucilagi ypodermis short-branching, with several to many short usu- ally sharp arms. Inflorescences at leafless nodes of twigs 3-7 mm diam., 1—4 per side, each usually branching near the base, forming a dense cluster, each 1—9-flowered, 4-12 mm long to base of far- ul costs have been supported in part by the Hayden Fund of the Botany Department, University ur iwan ANN. Missouni Bor. GARD. 72: 548-557. 1985. es Botany PYP 220 Bioscience Center, University of Minnesota, St. Paul, Minnesota 55108. 1985] thest pedicel measured along the axes and with 2-3 internodes in that length; bracts esl to ovate-triangular, 1.6-2 mm long, 1.8 mm wide, mostly dada by anthesis or sione after. True pedicels 5-10 mm long, to 13 mm in fruit; calyx including inferior ovary 6-8.5 mm long, cup-shaped, the 5 locules widely separated and appearing as if in the base of the hypanthium and bulging somewhat externally; apparent free hypanthium as measured from style base to sta- men attachments 3.6—4.8 mm; sepals yellowish green to pale yellow, 1 mm long, 2 mm wide, triangular, 1.5-2.5 mm long when measures from stamen attachment, t a further distance of 1.5-1. 8n mm at anthesis, the sepals then 3.5-5 mm wide. Petals white. Fila- ments 7-9 mm long; anthers 4-4.5 mm long; sporangia 4—4.4 mm long, dehiscing by apical pores; gland 0.7—0.9 mm long, 2.5-2.8 mm from apex of anther when measured from center of gland; cauda none. Placentae parietal-basal in each of the 5 locules, the ovules 5, borne on all sides of a short parietal-basal column, ca. 25 in all; style ca. 16-18 mm long. Fruit consisting of 3-5 subglobose lobes independently attached to the old hypanthium and calyx, the lobes some- times contacting each other but not laterally fused, reddish orange, 9.5-14 mm diam., 1-seeded o rarely 2-seeded, each lobe developed from 1 loc- ule of the ovary. Seeds brown, polished except for the large hilum, irregularly spheroid, 8.5-10.2 mm high, 9.8-12 mm wide, 6-8 mm thick, with a small rounded hump on the polished side and a larger one on the hilum side, the hilum occu- pying all of one side dien ed a broad extension of the polished surface w curves over and down on the hilum side in e of the height of that side. Paratypes. PERU.LORETO: O woods, Masisea, 275 m, (fr) 25 July 1929, Killip & E Smith 26845 (F, NY, US); Coronel Portillo, Bosque Nacional de von Hum- MORLEY —MEMECYLEAE 549 boldt, Km sn, Pucallpa- -Tingo Maria road, 270 m, Ar- 75 boretum an 0' 8?40'S, (fr) : Aug. 1980, Gentry & Salazar 29438 (US). N: high N MARTÍN: hi d in for E of house, Don Diogenes del Aguila, E of Aguaytia between Pucallpa d and Río Aguaytia, (defl, fr) 28 June 1960, Ma- 7879 ;12k fT , near Río Tocache, mature flatland forest on lateritic soil, 500 , 76°3 , 8*10'S ds) 13 Mar. 1979, Gentry, Schunke V. & Aronson 25651 (US Distribution. High forest to open woods in central Peru in the departments of Loreto, San Martin, and Huanuco in the drainages of the Huallaga and Ucayali Rivers at elevations of 270 o 500 m e Mouriri peruviana is very simlar to M. ner- vosa, differing from the latter mostly in the great- er size of its parts. The leaves of M. peruviana are somewhat larger than those of M. nervosa, the pedicels are somewhat longer, the calyx in- cluding the inferior ovary is larger, the calyx lobes are longer as measured from the stamen attach- ment, the anthers are longer and with a different shape, the lobes of the fruit and the contained seeds are larger, and the seeds have a different shape. Although most of these are differences of size, the new plants are considered to represent a species rather than a subspecific taxon of M. ner- vosa because of the gaps between the size-ranges of the calyx and ovary, anthers, fruit lobes, and seeds in the two taxa, and because differences of shape are also seen in the anthers and seeds. The geographic range is also very distinct. Previously I mistook the collection Killip & Smith 26845 to be M. nervosa. When this plant is properly rec- ognized as M. peruviana the nearest M. nervosa is about 1,000 km to the east. RE 1. Mouriri peruviana. —A. Leave veins (Mathias & Taylor 5006). oe Cross section of leaf blade showing upper epi ction crypts (Killip & sii 26845).— (Schunke 2077).—F. Long section flower, for the sporangia (Schnuke 2077). Mouriri pranceana. — A. Leave 5 Me —F. uin section 263). except for the sporangia (Prance et al. 25263). (Pires 16105). before anthesis (Schunke ).— —H. Fruit (Schunke 7879).—1I, J. Seed (Schunke 7879). 63). — D. Cross section of leaf midrib (Prance et al. mut. n of flower, before anthesis (Prance et al. 25263).— —H. Petal (Prance et al. 25263).—I. P (Pires 76105). — s. — B. Cleared portion of leaf blade showing terminal sclereids and ermis, sclereids, and stomatal midrib (Killip & Smith 26845).—E. Inflorescence 20 G. Anther, shown cleared except . — B. Cleared portion of leaf blade showing terminal sclereids and — C. Cross section of leaf blade showing upper epidermis, sclereids, and stomatal —E. v pee (Prance shown cleared —J. Seed MORLEY — MEMECYLEAE 551 1985] 552 Mouriri peruviana is also somewhat similar to M. angustifolia, a close relative of M. nervosa from the east-central border of Colombia. Mou- riri angustifolia has still smaller leaves than M. nervosa, Type II crypts instead of Type III, short- er pedicels, smaller flowers and flower parts, and only three ovules per locule in the ova The new species has sometimes been mistaken for M. cauliflora because the flowers of the latter are also rather large. However, M. cauliflora has a smooth lamina when dry, a prominently rounded midrib above with a groove on each side, and columnar foliar sclereids; the ovary loc- ules of the ovary do not bulge outwardly in the flower, and the fruits and seeds are smaller. Mou- riri cauliflora does occur in Peru, in Maynas and Requena Provinces of Loreto, but has not yet been found in the range of M. peruviana. Mouriri pranceana Morley, sp. nov. TYPE: Brazil. Mato Grosso: Serra do Cachimbo, Cuiabá- Santarém highway, BR 163, Km 764, 15 km S of Mato Grosso/Pará border, 520 m. Dis- turbed campina forest. Tree 3 m; petals b I. ad 1977, Prance, Silva, Berg, enderson, Nelson, Balick, Bahia & dos Santos P 25263 (holotype, MG; isotype, US). E 2. Frutex vel arbor usque 5 m alta; laminae apice acuto .5 mm depressa, vix manifesta; hilum seminis 2.2-2. longum, 1.8-2 mm latum. Shrub, or tree to 5 m high, glabrous except for the inflorescence; young twigs rounded, with 2 shallow channels on opposite sides. Petioles 3- 5 mm long; blades coriaceous, with a dull shine when dry, 8.5-12 cm long, 3.3-5.2 cm wide, ovate-elliptic to elliptic or narrowly so or these slightly oblong, broadly to medium acute or abruptly very short acute at the apex, rounded to broadly or medium acute at base, the blade abruptly short-attenuate on the petiole; midrib ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vo. 72 flat above or occasionally with a slight narrow central groove, prominent below, low-rounded near the petiole, becoming 2-angled !⁄—!⁄4 of the way to the apex or very narrowly 2-winged; lat- eral nerves invisible to slightly visible above and below when dry. Midrib xylem tubular; stomatal crypts a highly modified Type II, averaging in a leaf 40-50 um diam., 115-131 um high, 69-120 per mm? (extremes 30-65 um diam., 105-141 um high, 54-135 per mm?); upper epidermis varying with the thickness ofthe leaf, in the thin- ner leaves being mostly 2 cells thick, rarely one, in thicker leaves 2 or 3 cells thick; conspicuous mucilaginous walls usually present in the cells contacting the palisade and also in the middle cells when the epidermis is 3 cells thick; hypo- dermis none; terminal sclereids columnar, nar- leafless nodes of twigs to 7 mm thick or some- times axillary, 1 per side, each 3—9-flowered, 14- 3 mm long to base of farthest pedicel measured along the axes and with 2-3 internodes in that length; bracts early deciduous, unknown. Pedi- cels and at least the upper internodes of the in- florescence glabrous or very minutely puberu- lent. True pedicels 2.5-7 mm long; calyx including inferior ovary 6-8 mm long dry, to ca. 8.5 mm long fresh; calyx lobes fused to the apex of the flower in bud, splitting apart regularly at anthe- sis, each then ovate-triangular and acute, 3-3.8 mm long, 2.5-3 mm wide; free hypanthium 2.3- 2.6 mm long measured to middle of the stamen attachment; calyx circumscissile a bit irregularly in the free hypanthium ca. 0.5 mm above the base after anthesis. Petals white, sometimes with a red line on the center of the exterior, broadly elliptic and acute, i sessile, ca. 5.4 mm long, : e; matu laments unknown; an- thers yellow, 3.1-4 mm long; sporangia 2.4—3.3 mm long, dehiscing by apical slits; gland 0.8-1 mm long, 2.1-2.6 mm from apex of anther when measured from center of gland; cauda 1-1.5 mm long. Ovary 5-locular; placentae basal in each locule, the ovules borne on all sides of a short basal column, (3-)4—5 per placenta, 20-25 per ovary; mature style unknown. Fruit yellow and edible when ripe, tart when green, subglobose, FiGURE3. Mouriri oe (Lescure 834).—A. Leaves.—B. ign portion of leaf blade showing terminal deeds and veins. — of leaf midrib. — E. Flow LE. es J. Anther, shown ded. except ir ihe sporan section of leaf blade showing upper section of is before ikea —G. Inflor —D. Cross section rescence. — H. Fruit. — I. Seed. — idermis and sclereids. 1985] PH 30 MORLEY —MEMECYLEAE 553 | <` zz — rey 554 estimated ca. 28 mm diam., with a smooth round calyx scar 5.4—6.5 mm diam. that is scarcely vis- ible. Seeds 5 in the fruit examined, dark brown and polished, narrowly and irregularly ellipsoid, 12.8-12.9 mm long, 6.8-7.6 mm wide, 6.7-7.4 mm thick; hilum basal, angled slightly upward away from the center of the fruit, irregularly roundish in shape, 2.2-2.8 mm long, 1.8-2 mm wide. Paratypes. BRAZIL. PARÁ: Serra do Cachimbo, BR m Cuiabá: Santarém highway, Cachoeira de Curuá, ocky terrain, 300 m, (bud, fr) 4 Nov. 1977, Prance et al P 24831 (US); Serra do Cachimbo, BR 163, Cuiabá- tarém highway, Bis 879, 2 km from Rio Curuá, ow forest on terra e, (fl) 11 Nov. 1977, Prance et d P 25189 (RB, US): a i Xingu, Fazenda Rio Doura- o, R. Dou — aflue . Fresco, ca. 52°W, 85S, ancha de na pedrogosa no alto de morro, (fl, fr) 28 J June 1978. "Pires 16105 (NY). Distribution. Low forest to savanna to dis- turbed campina forest, often rocky, in southern Pará and adjacent Mato Grosso from the Serra do Cachimbo to the Xingu drainage at elevations from 300 to 520 m. Mouriri pranceana clearly falls in the small section Huberophytum, owing largely to char- acters of the calyx and ovary. The calyx lobes are fused for their length in the bud and separate regularly at anthesis, the hypanthium is circum- scissile post anthesis, and the placentation is typ- ical for the section. Within Huberophytum, M. pranceana is distinct from M. elliptica and M. cearensis on the basis of habitat, habit, and leaf form and structure, but it closely resembles the remaining species, M. huberi. From the latter it differs chiefly in its fruit, which is roundish with a small smooth calyx scar. The smooth scar ap- parently results from the fact that the hypan- thium is circumscissile at its base, ms little tissue on top of the enlarging ov n M. huberi the hypanthium is pcan er VEN of the way from the base to the stamen attachment, leaving on the developing fruit a ring of tissue which becomes a broad circular inward-pointing rim 0.5-1 mm high. Apparently there is less expan- sion at the end of the developing fruit in M. pranceana than in M. huberi, and thus a smaller scar is produced in the former. Other distinguishing features of the new species in comparison to M. huberi are the former's more acute blade apex and less grooved midrib above, its thicker leaves with higher stomatal crypts and more strictly columnar sclereids, its longer ped- icels and larger calyx including the inferior ovary, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 its larger petals and anthers, its greater number of ovules, and its smaller seeds and hilum at least in the few examples checked. It ] tfor Dr. Prance, wide- -ranging collector in Amazonia. Mouriri cearensis Huber subsp. carajasica Mor- ley, subsp. nov. TYPE: Brazil. Pará: Serra dos ee Serra do paid ca. 20 km N of AM mp (ca. 6°S, 50?15'W). In oe forest. Tree er m by 25 cm; corolla white. 18 Oct. 1977, Berg, Henderson & Ba- hia BG 615 (holotype, MG; isotypes, MICH, NY, US). Petioli alati 4.5-9 mm longi; laminae petiolis 11-17 E longiores; lobi calycis 2.4-2.9 mm longi in sicco, m longi in vivo; cicatrix calycis 4—6.5 mm diam. in rants Tapering winged petiole 4.5-9 mm long, the lamina 11-17 times as long as the petiole; calyx lobes 2.4-2.9 mm long when dry, 2.6-3.2 m long boiled es probably when fresh; calyx scar on dried fru .5 mm diam.; seeds 9.5-11 mm high with a em constriction about '4 of the way above the base, mostly dark brown above and below the constriction and medium brown at the constriction, the hilum 4-5 mm long, 2-3 mm wide. a en IL. PARA: nats dos Carajás: Mara- bá, caminho para eis o do aeroporto, (im Msg eft) 3 jade 1977, Silva 4 ps 3034 (NY "US); orte near AMZA Exp pied Camp (ca. 6°S, e15) W), low Tor est on edge fet pe, ca. dm 11 Oct. 1977, Berg & l tas BG 469 (NY, US); ZA Exploration Camp, in fina duda (fr) 19 Oct. 1977, Berg & Henderson BG 638 (MICH, NY). Distribution. Low forest or liana forest on terra firme in the Serra dos Carajás in southeast Pará, Brazil, at elevations of ca. 300 to 500 m. Subspecies cearensis has the following con- trasting characters: winged petiole 2-5.5 mm long, lamina (14—)18—30 times as long as the petiole, calyx lobes 3-3.8 mm long when dry, 3.2-4 mm long when boiled, calyx scar on dry fruit 7.5-10 mm diam., seeds 10.7-12.5 mm high, without a constriction, medium brown, the hilum 4-5 mm long, 3-3.4 mm wide. Subspecies cearensis occurs near the coast in Ceara, Piaui, and Maranhão, disjunct nearly 600 km from the new subspecies, as the distributions are now known. Although subsp. carajasica is 1985] distinct in morphology as well as being isolated geographically, in my estimation it is best re- garded as a subspecies because there is little or no gap between the differing character-ranges of the two taxa and because of the great overall similarity in leaf anatomy, flower, and fruit in e two Mouriri viridicosta Morley, sp. nov. TvPE: French Guiana. St. Georges de l'Oyapock: Foret prés de la Crique Gabaret. Arbuste de 6 m Feuilles opposées à nervation peu visible. Fleurs fanées à calice jaune. Petits fruits isotype, CAY; note on US label says Lescure 834 — Oldeman B2731). FIGURE 3. Te 4 r4 14 £42 in sicco viridis; xylema costae mediae canaliculatum, non matophorae nullae; pedicelli 0.5-1.5 mm longi; calyx ovarium inferum includens 2.5 mm longus; ovarium 1-loculare, ovula rs fructus 6.2-7.9 mm diam., stylo persistenti; semen 1, ca. 5 mm longum, 6.4 mm latum. Shrub 6 m high; young twigs terete, gray. Pet- ioles 1.5-2 mm long; blades 11—-17.7 cm long, 3.3-5.6 cm wide, elliptic-oblong to ovate-oblong or elliptic-ovate-oblong, abruptly acuminate at the apex, acute at base; midrib grooved above, rounded below; lateral nerves faintly to moder- ately visible above when dry, faintly visible be- low; blade including the underside of the midrib a medium green when dry. Margins of the leaf midrib xylem extended and turned i in and down but not fused to each upper epidermis one cell thick; hypodermis none; mucilage walls none; free stone cells present only at base of midrib; terminal sclereids filiform, oc- casionally branching; tanniniferous compounds lacking. Flowers 1—several in crowded clusters at leafless nodes oftwigs to 3 mm diam., the flowers per peduncle, the peduncle ca. 0.5 mm long with a pair of short triangular bracts at top and bottom, the bracts 0.7-0.8 mm long. True ped- icels 0.5-1.5 mm long; calyx including inferior ovary 2.5 mm long, campanulate, glabrous; calyx yellow; free hypanthium 1.2 mm long; calyx lobes low-triangular, 0.3-0.5 mm high, 1.2-1.4 mm wide, not separating further at anthesis. Mature petals unknown. Mature filaments unknown; an- thers 1.2-1.4 mm long; sporangia 1.1-1.3 mm long, their mode of dehiscence unknown; gland 0.2-0.3 mm long, placed ca. 0.4 mm below the MORLEY —MEMECYLEAE 555 apex of the anther; cauda none. Ovary 1-locular with 4 ovules arranged around a short central and basal placenta; style 2.8-3 mm long. Pr doni subglobose, 5.7—6.2 mm high, 5.9—6 iam. when dry, estimated 7-8.5 mm ido Men fresh, crowned with the remains of the hy- panthium and calyx and with the persistent style. Seed 1, ca. 5 mm high, 6.4 mm wide, 2 mm thick, the polished center part (the enlarged outer face of the ovule) ca. 2.3 mm wide, the rest of that face of the seed wrinkled irregularly with subparallel lines radiating away and upward from the polished center. Distribution. Known only from the type lo- cality in tropical forest near the Oyapock River in east French Guiana at about 40 m elevation. The seed of M. viridicosta unmistakably places this species in the subgenus Taphroxylon, where its non-tubular midrib xylem also fits best. How- ever, within this subgenus the new plant has no close relatives. Even its section is uncertain. The evergreen habit, short pedicel, single tiny flower, and seed structure exclude it from section Abun- diflos. The lack of tannins and the form and structure of the anther are agreeable with section Taphroxylon, but this relation is disputed by the limited distribution of the free stone cells and by he filiform sclereids, very short pedicel, tiny flower, and separate calyx lobes. Of the present sections the new species is most at home in Brevipedillus. The limited distribu- tion of the free stone cells, very short pedicel, tiny flower, and separate calyx lobes are all matched in this section, and even the filiform sclereids are approached in M. duckeana and M. duckeanoides. However, the lack of tannins and the anther form and structure are discordant. a section of its own, since the fully filiform sclere- ids, small number of ovules, persistent style, and small fruit and seed are unique in the subgenus. However, because the majority of the known dis- tinguishing characters for sections fall into an existing section, I am inclined to view M. viri- dicosta as an aberrant member of section Brev- ipedillus until further evidence indicates other- wise. The persistence of the style on the fruit was known previously only in Votomita monantha; I had thought this might be a generic character for Votomita, but was mistaken. 556 A p d pm CA Oo m FIGURE 4. and veins. — B. Cross section of leaf midri Votomita pubescens (Lao M. 22). Votomita pubescens Morley, sp. nov. TYPE: Peru. Loreto: Requena, Sapuena, Jenaro Herrera, 120 m. Collection from the Dep't. of Forest Management, Natl. Agric. Univ. at La Mo- lina, Lima. Ties number R-D12 JH. “Lanza caspi." Arbol; bosque humedo tropical. Abd Rafael ao M. 22 (holotype, F). FIGURE 4. Arbor; folia supra glabra, subtus a scle- rides foliorum ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 Gss ee ss 300 pm wo === 3 S —A. Cleared portion of leaf blade showing terminal sclereids Cross section of leaf = showing upper e ib. — D. Leaf. — pidermis, sclereids, dm crypts, and hairs.— E. Inflorescence. M^ Long section of flow rium 10.3-13.3 mm longum, tantum parvum contrac- tum in sicco, in sicco laeve, induratum, olivaceum, duplo crassius ad placentas quam ad apicem, 2-locu- lare, placentis 2 in quoque loculo. Tree. Petiole 5-6.5 mm long; blades 12-14 cm above but with a slight groove down the center, 1985] prominent below, the latter part rounded at base but soon 2-angled with 2 very narrow wings for its length, pubescent on the sides; lateral nerves not visible above or below when dry. Stomatal crypts multiple, each branched above the mouth into 2-7 small cavities, the multiples 90—175 um total diam., 65-87 um deep, averaging 35 per mm?; upper epidermis mostly 1 cell thick, oc- casionally 2, of large square cells as seen in sec- usually with an irregular short-branched central body and a dendroid branching system against each epidermis; hairs 80-190 um long, appar- ently 1-celled but with 1—6 septae and an ex- panded base set in the epidermis. Peduncles ax- illary, 1 per side, each 2-3-flowered, 19-27 mm long to base of farthest pedicel measured along thes: axes and with 1 or 2 pakistan in that length; idu True pedicels 8-10 mm pee pui 4-merous; calyx including inferior ovary 10.3-13.3 mm long, 7.8- 9 mm thick, glabrous, shrinking only slightly on drying, when dry with a hard smooth somewhat polished olive green outer layer, the ovary about twice as thick at the midpoint as at the apex; calyx lobes separate, 0.9-2 mm long, 2.4-2.7 mm wide, triangular and slightly apiculate, not sep- arating further at anthesis. Petals and stamens unknown. Ovary 2-locular, each locule with 2 axile placentas and an irregular very incomplete ridge running down the outer wall representing a missing partition; ovules 5—6 per placenta; style unknown, apparently deciduous at anthesis. Fruit and seed unknown MORLEY — MEMECYLEAE 557 Distribution. Known only from the type lo- cality in a humid tropical forest in northeast Peru in the drainage of the Ucayali River at an ele- vation of 120 m The previously known species of Votomita form two groups, three species (V. plerocarpa, V. monadelpha, V. orbinaxia) with (3—)4 locules and 36-48 ovules and three species (V. guianensis, V. orinocensis, and V. monantha) with one locule and 5-10 ovules. The new species falls in between these groups, because it has two locules (with two placentas in each) and 20-24 ovules. It is not similar to any of the other six. However, it is doubtless closest to the first group because of the four placentas and the fact that two of the first group (V. plero- carpa and V. monadelpha) have pubescent mid- set it apart from all the other six species: the pubescent lamina, the ovary that is twice as thick at the midpoint as at the apex and that is smooth and hard when dry, and the two locules with a total of 20-24 ovules. LITERATURE CITED Mor ey, T. 1976. Memecyleae (Melastomataceae). Fl. Neotrop. 15: 1-295. Tosi, J. A., Jk. 1960. Zonas de vida natural en el TU: memoria explicativa sobre el Mapa Ecoló- gico del Perú. Bol. Técn. Inst. Interamer. Ci. Agríc. 5: 39. NEW SPECIES OF SOLANUM SECTION GEMINATA (G. DON) WALP. (SOLANACEAE) FROM SOUTH AND CENTRAL AMERICA! SANDRA KNAPP? ABSTRACT Four new species of Solanum section Geminata are described: S. tillum, S. heleonastes, S. pas dasyneuron, and S. aphyodendron. Two of these are Central American, one from Guatemala and one from Costa Rica. The third is only found along the Río Paraná along the border of Argentina and Paraguay. The fourth is a widespread species that has long been confused with S. nudum H.B.K. ex Dunal. Nomenclature and details of natural history are discussed. Solanum section Geminata (G. Don) Walp. is one of the largest sections in the genus, with ap- proximately 70 species, all but one of which is exclusively neotropical. The section has previ- ously own as section Leiodendra Dunal (D'Arcy, 1972), but for reasons of priority (Knapp, 1983), Geminata (G. Don) Walp. is the correct sectional name for this species group. Members of section Geminata are generally small trees or shrubs, often growing in the pri- mary forest understory, an unusual habitat in the genus Solanum. Species in the section share di- foliate nodes with many (but not all, see Danert, 1967; Child, 1979) other Solanums, which in section Geminata, as the name implies, are often geminate. In this situation the two leaves of the node arise from the same height on the stem due to failure of mesopodial elongation (Danert, 1958): therefore the subtending leaf of the pre- vious shoot generation is at the same level as its daughter shoot. The leaves in a geminate cluster of equal size and shape or can differ in one or both of those characters. The most com- mon situation in section Geminata is for the geminate cluster to be anisophyllous. The larger further characterized by the leaf-opposed nature of the inflorescence, which also arises from the same level on the stem as the geminate leaves. _' I thank Ww G. . D'Arcy, R. L. Dressler, W. Haber, B. Hammel, W. J. Kress, J. Mallet, M. an This “‘leaf-opposedness” is due to meso- and epi- podial suppression and adnation of the stem and the peduncle (Danert, 1967; Child, 1979) and is not perfect in all species. Pubescence in the sec- tion is absent or consists of uniseriate trichomes "finger hairs" sensu Seithe, 1962, 1979), but branched trichomes do occur. The flowers are small, white or greenish white, and the degree to which the petals reflex at anthesis is often a good specific character. Fruits of most species are green and remain hard until the day of maturity, when they become soft and yellowish (pers. observ.), but several species have brightly colored, bud sumably bird-dispersed fruits. The pedice of flowers that fall off without producing fruit become corky and are prominent in later stages of inflorescence growth. The spacing and ar- rangement of these scars is generally an over- looked character (but see D’Arcy, 1973) and is useful in distinguishing the species of section Geminata. While preparing a evan of Solanum section Geminata, I have di S both in the herbarium and in the field. Four of these are described here. Further details of the natural history of S. pastillum and S. aphyo- dendron can be found in Knapp (1985). Lam pecies, Solanum pastillum S. Knapp, sp. nov. TYPE: Cos- ta Rica. Puntarenas: Monteverde, Camp- nd M Sy i following herbaria for access to their specimens: BH, WIS. Field work in Costa is gratefully acknowledge 2 L. H. Baile rol Kalafatic fo and Jack Putz for field assistance, and W. J. Dress for x with the Latin diagnoses. I thank ne A of the POM Rica was funded by an Organiza on pu Tropical Studies Inc. Jessie Smi th Noyes n Jos C . G. D'Arcy of the Missouri Botanical Garden. The support of these institutions ailey Hortorium, Cornell University, Ithaca, New York 14853. ANN. Missouni Bor. GARD. 72: 558—569. 1985. 1985] KNAPP—SOLANUM SECT. GEMINATA 559 FIGURE |. bell’s Woods, 1,450 m, 20 Aug. 1982, Knapp, Holbrook & Putz 6063 (holotype, BH; iso- types, CR, F, MO, US). Figure 1. Frutex gracilis; caules virides petiolorum basibus snp alati; folia ovata geminata utrinque glabra sub- us pallide viridia, apice acuminato, basi attenuata de- urrente; inflorescentiae foliis opposita simplices gla- brae pendulae; see sub anthesi deflexi; calycis lobi orbiculati succulenti 1 mm diam.; corolla. rotata m viridi-alba, lobi vo- viridis, pedicello lignoso infra calycis tubum pedis semina ovoidea reniformia fulva, testa foveolata. Slender arching shrubs, 1—3 m tall; stems gla- brous, green, horizontally spreading, winged be- tween the nodes with the decurrent leaf bases; bark of the older stems pale green, white-lenti- cellate. Leaves geminate, elliptic-ovate, widest at or just below the middle, dark green and slightly bullate above, pale sea green beneath, glabrous; major leaves 11.2-22 cm long (occasionally to 25 cm on the lower branches), 3.4—6.6 cm wide (to 11.3 cm on the lower branches), with pairs of primary veins, raised above, paler and prominent beneath, the apex long acuminate, the base decurrent on the petiole, attenuate to cu- neate; petioles 0.5-1 cm long; minor leaves dif- Solanum pastillum (from Knapp, Holbrook & Putz 6063). fering from the major ones only in size, 2.1-6 cm long, 1.1-3.5 cm wide, the apex long acu- minate, the base attenuate; petioles 3-5 mm long. Inflorescences opposite the leaves, simple, gla- 5-50-flowered; pedicel scars irregularly nM 1-3 mm apart, beginning ca. 1 cm from the bas ofthe Paani pedicels at anthesis cea 1.4-1.5 cm long, tapering from the calyx tube to a ino base ca. 0.5 m diam.; buds conical, the corolla soon exserted from the globose calyx lobes, glabrous; calyx tube ca. 1.5 mm long, broadly cup-shaped, translucent green, lobes on fresh specimens consisting of 5 orbiculate projections ca. 0.5 mm diam. on the rim of the calyx tube, in dry specimens of irregularly shaped projec- tions 0.5-1 mm long from the rim of the calyx tube, glabrous and translucent green; corolla greenish white, 1.1—1.2 cm across, lobed ca. three- fourths of the way to the base, the lobes reflexed at anthesis, the tips ofthe lobes carinate-hooded, minutely papillose; anthers 2.5-3.25 mm long, 1.25-1.5 mm wide, poricidal at the tips, the pores becoming slit-like upon drying; free portion of the filaments ca. 0.5 mm long, the filament tube 4s dasyneuron € S. pastillum CENTRAL AMERICA FIGURE 2. veni of Schon pastillum and Sian dasyneur ca. 0.5 mm long; ovary glabrous; style 5-6 mm long, straight or curved in dry specimens; stigma capitate, minutely papillose. Berry globose, greenish yellow at maturity, ca. 50-seeded, 1-1.5 cm diam., usually only 1 or 2 from an inflores- cence; fruiting pedicels 1.9-2.6 cm long, woody, deflexed, greatly ded at the distal end, there ca. 4—7 mm diam., the base slender, ca. 0.5 mm diam.; seeds pale brown or tan, ovoid reniform, 3.5-4 mm long, 2-2.5 mm wide, the surfaces minutely pitted, the pits very shallow, the seeds appearing smooth. Chromosome number, n = 12 (Knapp, unpubl. data; voucher, Knapp: et al. 6063). Distribution. Found only in the cloud forests of montane central Costa Rica from 1,000 to 1,700 m. Figure 2. Solanum pastillum is an isolated species in section Geminata. The most probable close rel- ative of S. pastillum is S. tuerckheimii Greenm., also of montane central Costa Rica (but extend- inflorescences of greenish white flowers with re- flexed petals at anthesis, and hard green fruits which become yellow and soft upon maturity. Solanum pastillum blooms at the beginning of the wet season at Monteverde de Puntarenas, Costa Rica, the type locality, and is pollinated primarily by bumblebees, Bombus ephippiatus (Knapp, 1985). The local distribution pattern of S. pastillum is typical of many of the forest understory species of section Geminata. Small clumps of three to five individuals grow at widely ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 spaced intervals in the understory, often at the edges of old gaps caused by fallen trees or branch- es. The inflorescences of S. pastillum are many- flowered, but only a few fruits per inflorescence develop. Whether this is due to poor pollination success or to fruit abortion is not known. The presence of a single fruit on a long inflorescence causes the effective pedicel length of the fruit to be extremely long. This may make the fruit more visible to small frugivorous bats, known to take the fruits of this and many other species of sec- tion Geminata in Monteverde (E. Dinerstein, pers. comm.). The specific epithet, “‘pastillum” meaning small round loaf of bread, refers to the orbiculate calyx lobes, which are quite distinctive in this species. Additional specimens examined. COSTA RICA. ALAJUELA: Llanura de San Carlos, 100 m, 21 Feb. 1966, Molina R. et al. 17659 (F); Penas Blancas Trail, 3.5- 5 km ESE of ndis ca. 1,450 m, 17 Aug. 1976, Kennedy & Guindon F); Mont Sven © on road to Peñas Blancas (over ci Divide below the Divide on Atlantic slope, 1,450 m, 10 84*50'W, 13 Apr. 1981, Knapp & Mallet 864 qn. Alfaro Ruíz, Guade Zarcero, 1,400 m, 11 July 1938, Smith NY904 (F, NY); Alfaro Ruíz, Zapate, 1,425 m, 10 May 1940, Smith P2663 (F); Alfaro Ruíz, 5 m o 1,500 m, 17 June 1941, Smith 2789 (P. CARTAGO: El Mufieco on Río Navarro, 1,400-1,500 m Mar. Standley & Torres R. 51016, 51040 (US). PUNTAR- ENAS: Sendero El Brillante, Monteverde Cloud Forest 1,580 m, 16 June 1976, Dryer 208 (F); Monteverde, near the Ventana (Continental Divide), 1,500-1, m, 12 July 1976, Dryer 426 E of road, Monteverde Cloud Forest Reserve, 1,5 1,580 m, Aug. 1976, on, 1,150 m, 29 July 1937, Brenes 22615 (F); 5 mi. S of Santa María, 6,800 ft., 5 Feb. 1928, Stork 1757 (F) Solanum heleonastes S. Knapp, sp. nov. TYPE: Argentina. Corrientes: Capital, Puerto Italia en montes costeros del Río Paraná, 6 Nov. 1972, Schinini 5671 (holotype, MO; iso- types, MO, WIS). Figure 3. Frutex vel arbor parva; caules juniores incani tri- homatibus minutis uniseriatis dense pubescentes; caules veteres succulenti nitidi rubiginosi; folia ovata geminata supra glabra subtus in nervis minute papil- lata, nervis stramineus, apice acuto vel acuminato, basi 1985] KNAPP—SOLANUM SECT. GEMINATA FIGURE 3. Solanum heleonastes (from Schinini 5671, fruit from Schwindt 30). cuneata valde obliqua; inflorescentiae = oun simplices modo caulium juniorum pubescentes; pedi- celli sub anthesi erecti; d lobi ne DM e; corolla rotata alba, lobis planis leviter carinatis; bac globosa lignosa, in specimenibus exsiccatus ochracea, formia, testa foveolata; habitat in paludibus fluvii Pa- rana inter Argentina et Paraguay. Shrubs or small trees, 1—5 m tall; young stems and leaves densely covered with appressed uni- seriate papillate trichomes, 0.05-0.1 mm long, these irregular and floccose in appearance, grey- ish brown, the young growth with a grainy tex- ture; bark of the older stems red-brown, shining, y white-lenticellate. Leaves glabrous above, minutely papillate on the veins beneath, the veins yellowish, epidermis large- celled and appearing crystalline; major leaves 10.2-18.5 cm long, 5.8-7.9 cm wide, with 8-10 strong, evenly spaced primary veins, these yellow beneath, the secondary venation obscure, the apex acute or acuminate, the base broadly cuneate, strongly oblique; petioles 1.7—3.2 cm long, slight- ly winged from the decurrent leaf bases; minor leaves differing from the major ones only in size, 7-11 cm long, 3.4—5.5 cm wide, the apex acute or acuminate, the base broadly cuneate, atten- uate, oblique; petioles 0.8-1.2 cm long. Inflores- cences opposite the leaves, simple, 1-2 cm long, 10—30-flowered, densely covered with papillate overlapping; pedicels at anthesis erect, 1.1—1.4 cm long, tapering from the base of the calyx tube to a slender base ca. 0.5 mm diam., minutely white-speckled, sparsely papillate with uniseriate 562 T — C Ken aie / À 1 cm ANNALS OF THE MISSOURI BOTANICAL GARDEN N | D [VoL. 72 d 3». 3 4, ; Y, <, iw Ficure 4. Solanum dasyneuron (from Matuda 2849). trichomes like those of the young stems and leaves; buds at first globose, enclosed in the ur- ceolate calyx tube, later ovoid and exserted from the clayx tube; calyx tube 1.5-2 mm long, ur- ceolate in bud, later cup-shaped, the lobes long triangular, 2.5-5 mm long, white-speckled, mi- nutely papillose at the tips; corolla white (blue in Schwarz 8182), 1.2-1.5 cm across, lobed near- ly to the base, the lobes planar at anthesis, tips of the lobes slightly carinate, minutely papillose at the tips and the margins; anthers 3.5-4 mm long, tightly connate in young flowers, 1-1.5 mm wide, poricidal at the tips, the pores becoming slit-like upon drying; free portion ofthe filaments ca. 1 mm long, the filament tube ca. 0.5 mm long; ovary glabrous; style filiform, 5-7 mm long, straight; stigma minutely capitate, white-papil- lose. Berry globose, dry and woody with very little pulp, 1-1.2 cm diam., mustard yellow in dried specimens, the septum between the carpels dry and papery in dry specimens; fruiting pedi- cels erect and woody, 1.1-1.4 cm long, ca. 1 mm 1985] ARGENTINA € S. heleonastes e—— 200 km Ficure 5. Distribution of Solanum heleonastes. diam. at the base; calyx lobes becoming woody and reflexed in fruit; seeds dark brown, flattened reniform, ca. 2 mm long, 1.5 mm wide, the sur- faces minutely pitted. Chromosome number, not known Distribution. Found only in the swampy for- ests on the margins and islands in the upper Rio Paraná, on the border between Argentina and Paraguay, from 100 to 200 m. Figure 5. Solanum heleonastes is closely related to S. robustifrons Bitter of eastern Peru and adjacent Brazil, differing from that species in its longer calyx lobes, smaller leaves, and generally more pubescent young stems and leaves. Solanum he- leonastes is usually a large tree, and S. robusti- frons is a small shrub or treelet that grows in the understory of dense forests at middle elevations. Solanum heleonastes is one of the few species of section Geminata growing in swamp forest and e p meaning in itant, refers to the unusual habitat of this species. Additional specimens examined. ARGENTINA. CHACO: 1? de Mayo, Col. Benítez, Bermejo, Isla Bra- siliera, 4 Oct. 1964, Schulz 9422 (F, WIS). — Capital, on the Río Paraná 5 km from Corri , 10 Oct. 1944, Hunziker 5700 (US); Arroyo desee) 15 Dec. 1944, Sesmero 201 (NY); Capital, Puerto Italia, 27 July 1974, Schinini & Gonzalez 9502 (MO, WIS); Ituzaingó, Isla San Martín, 9 Oct. 1949, Schwarz dn (BH). MISIONES: Posadas, Río Alto Paraná, 18 1907, Ekman 820 (MO); Posadas, 17 Apr. 1930, Ro: KNAPP—SOLANUM SECT. GEMINATA 563 dríguez 188 (F); Candelaria, San Juan, 216 m, 4 Jan. 1947, Schwindt 30 (RSA/POM). PARAGUAY: Sin. loc., 1853-1856, Palmer s.n. (US). Solanum dasyneuron S. Knapp, sp. nov. TYPE: exico. Chiapas: Volcán Tacaná, Chiqui- huite, 2.800 m, 27 Mar. 1939, Matuda 2849 (holotype, F; isotypes, F, NY). Figure 4. (tex; caules gla b Fru vecta albicantes; : folia obovata vel anguste obovata flexo; habitat in sylvis nubosis Volcan Tacana insi- dentibus inter Mexico et Guatemala. Shrubs 2-3 m tall; young stems and leaves glabrous or occasionally minutely rusty-papil- late, drying black and shiny; bark of older stems white, exfoliating in small transverse pieces. Leaves obovate to narrowly obovate, geminate, glabrous above, the midrib depressed, the pri- mary veins raised, densely pubescent on the pri- mary and secondary veins beneath with unise- riate trichomes 0.25-0.5 mm long, trichomes restricted to on the veins, none on the lamina; major leaves 12.5—17.3 cm long, 5—6.5 cm wide, with 11-15 pairs of primary veins, the apex acute, the base attenuate, not decurrent on the petiole; petioles 1.5-1.9 cm long; minor leaves differing from the majors only in size, 2.2-7 cm long, 1.6— 2.9 cm wide, apex acute, the base attenuate; pet- ioles 0.6-1.1 cm long. Infl leaves, simple or occasionally once furcate, 1-5 cm long, 10-20-flowered, glabrous and shining; pedicel scars closely spaced and overlapping, be- ginning 4—6 mm from the base of the inflores- cence; pedicels at anthesis deflexed or somewhat m n to the calyx lobes; na ovoid, shy, p nly par- tially exserted from the calyx; calyx m l. 5-2. 5 mm long, conical, the lobes deltoid, apiculate, 1-2 mm long, papillose within, glabrous without, the margins of the lobes white and thickened; corolla white, thick and fleshy, ca. 1.5 cm across, lobed nearly to the base, planar at anthesis, the lobes minutely papillose on the margins and tips, tips cucullate; anthers 3.5-4 mm long, 1-1.5 mm 564 € KaAsd obe ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 1 cm FiGURE 6. Solanum aphyodendron (from Knapp, Kress & Hammel 4136). wide, poricidal at the tips, the pores becoming slit-like upon drying; free portion ofthe filaments ca. 0.25 mm long, the filament tube ca. 0.5 mm long, drying black; ovary glabrous; style ca. 7 mm long, straight; stigma large and capitate, ca. 1 mm diam., minutely papillose. Berry globose, 8 mm diam. (immature), green; fruiting pedicels de- flexed, woody, 1—1.5 mm diam. at the base; calyx lobes enlarging and becoming woody in fruit, ca. 4 mm long; seeds unknown. Chromosome num- ber, not known. Distribution. Endemic to the upper slopes of Volcan Tacana, on the border of Mexico and Guatemala, in wet forest from 2,500 to 3,000 m. Figure 2. Solanum dasyneuron is closely related to S. America. Solanum dasyneuron differs from both of those species in its large fleshy flowers, thick pedicels, its tendency to dry blackish, and by the dense pubescence on the veins on the undersides of the leaves. The species epithet, *dasy" meaning hairy and "neuron" meaning nerve, refers to the distinctive pubescence on the veins of the leaf undersides. Additional specimen examined. GUATEMALA. SAN MARCOS: between La Vega ridge along Río Vega and NE slopes of Volcán Tacaná, to 3 mi. from Guatemala- Mexico border, in vicinity of San Rafael, 2,500-3,000 m, 20 Feb. 1940, Steyermark 36198 (F, 2 sheets). 1985] KNAPP— SOLANUM SECT. GEMINATA 565 ° ° ' 2J ° g a ° s E ub s Pai - dts a e j X re °@° y \ { f , AN — 8 Po) po < | (4 rem *. ` | e. d. 3 . ^ š » % xa a p se $° f ° $ * v e "e * S. aphyodendron S 2? Wa s l 200 km $ ev CENTRAL AMERICA < } FiGure 7. Distribution of Solanum aphyodendron in Central America and Mexico. Solanum aphyodendron S. Knapp, sp. nov. TYPE: Panama. Chiriqui: along Quebrada Aleman, 8 mi. N of Los Planes de Hornito, IRHE Fortuna Hydroelectric Project, 1,200 m, 8°45'N, 82°12'W, 13 Mar. 1982, Knapp, Kress & Hammell 4136 (holotype, BH; iso- we MO, others to be distributed). Fig- e6. ai ns Brandegee, Univ. Calif. Publ. Bot. 6: 917n non Solanum "umala Link in Buch. YPE: e acru uapan, Dec. 1915 Purpus 7565 (CAL, isotype, MOD. Solanum nudum auct. .B.K. ex Dunal, Solan. Syn. 20. 1816. Solanum parcebarbatum Morton & Standley, Publ. Field Mus. Nat. ms Bot. Ser. 18: 1088. 1938 pro parte, non Bitt Frutex vel arbor parva m apertos habitantes; caules bis, ap- simplices glabrae; gemmae glo thesi deflexi; calycis lobi deltoidei trichomatibus unise- riatibus pubescentes 1 mm longi; corolla rotata alba lobis sub anthesi reflexis; bacca globosa flavo-viridis, P s" gd suo Md testa foveolata, marginibus incrassata. Shrubs or small trees, 3-15 m tall; young stems and leaves glabrous or with a few scattered uni- seriate trichomes, drying blackish; bark of older stems greenish white and rough; stems rl winged from the Epiri petiole bases. Leave narrowly ovate, geminate, widest at or just nat e the middle, drying pale greyish green, glabrous above and below, except for tufts of uniseriate trichomes in the axils of the primary veins be- neath, the trichomes ca. 1 mm long, white; major leaves 8.5-15 cm long, 3-6.1 cm wide, with 8— 10 pairs of primary veins, these depressed above, raised and pale greenish white beneath, the apex acute or acuminate, the base cuneate or slightly attenuate, with a small wing on the petiole; pet- ioles 0.6—1.7 cm long; minor leaves differing from the major ones only in size, 3.2-8.6 cm long, 1— 4 cm wide, the apex acute or acuminate, the base cuneate; petioles 4-8 mm long. Inflorescences opposite the leaves, simple, 1-2.6 cm long, 20- 30-flowered, glabrous or occasionally with a few scattered uniseriate trichomes distally; pedicel scars closely spaced, occasionally overlapping, ca. 1 cm fro uniseriate trichomes 0.1-1 mm long; calyx tube ca. 1 mm long, crateriform, the lobes broadly triangular, 0.5—1 mm long, densely to sparsely 566 pubescent with uniseriate trichomes 0.1-1 mm long, the margins ofthe lobes white and scarious; corolla white, often drying with a lavender tinge, 1-1.2 cm across, lobed nearly to the base, the lobes slightly reflexed at anthesis, tips and mar- gins of the lobes minutely papillose; anthers 2.5- 3 mm long, 1-1.5 mm wide, poricidal at the tips, the pores becoming slit-like upon drying; free portion of the filaments ca. 0.25 mm long, the filament tube ca. 0.5 mm long; ovary glabrous; style 5-7 mm long, straight; stigma not distinct from the body of the style, minutely papillose. Berry globose, bright green, ripening yellow-green, 1-1.2 cm diam., remaining hard until maturity, then softening overnight; fruiting pedicels de- flexed, woody, 1.5-2.3 cm long, ca. 1 mm diam. at the base; seeds tan, flattened reniform, 2.5-3 mm long, 1.5-2 mm wide, the surfaces minutely pitted, margins incrassate, pitting of margins larger than that of the body. Chromosome num- ber, n = 12 (Knapp, unpubl. data; voucher, Knapp et al. 4136). Vernacular names. “*Capulin de pajaro,” "yerba de zopilote," Mexico; "zorrillo," Las Cruces, Puntarenas, Costa Rica; “kaqi sakyol," Quecchi language, Alta Verapaz, Guatemala; “huele de noche," Chiquimula, Guatemala; **he- dionilla," Quezaltenango, Guatemala; “hiede hiede," Cerro Punta, Chiriqui, Panama. Distribution. Widely distributed in second growth at middle to high elevations (800-2,500 m) from Mexico to Bolivia. Figures 7 and 8. Solanum aphyodendron is a common species of roadsides throughout Central America and northern South America, and it often forms large monospecific stands in open areas. Material from Central America, Colombia, and Venezuela is quite uniform. Variability in pubescence in- creases in Peruvian and Bolivian specimens, with some men calyx lobes and denser trichomes i in the vein ax- ils. In Monteverde de Puntarenas, Costa Rica, Solanum aphyodendron blooms at irregular in- tervals throughout the year (Knapp, 1985). Data from herbarium specimens and observations of this species in Panama indicate that its flowering behavior is very much the same throughout its range. At Monteverde, S. aphyodendron is pol- linated by Meliponine bees, primarily by Meli- pona fasciata (Knapp, 1985). The occurrence of large monospecific stands of S. aphyodendron trichomes on the ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 f } § \ > ^ BOLIVIA | à ° e| Qu N i ° @ S. ophyodendron -— | 200 km URE8. Distribution of Solanum aphyodendron in Sou America combined with the foraging behavior of Meli- pona produces high fruit set in this species. The ripe fruits are eaten by small frugivorous bats and disappear soon after they become ripe (E. Dinerstein, pers. comm.). Solanum aphyodendron has long been con- fused with S. nudum K. ex Dunal, to which it is extremely closely related, but differs in hab- itat, flower size, pubescence of flower parts, and in the color and texture ofthe bark of older stems. Solanum nudum occurs in low elevation thick- ets, sometimes on beach strands, and has dark brown bark and glabrous, globose buds. The type specimen of S. nudum in the H.B.K. herbarium at Paris clearly belonged to the lowland, glabrous budded taxon, which left the pubescent, middle to high elevation taxon without a name. Bran- degee's (1917) epithet "foliosa" unfortunately An unpublished name, S. chamaecerasus Bitter, has been applied to this species in a few herbaria, 1985] but I have chosen not to use it, as Sauce to the characteristics of the spec This species is named for its i NN whit- ish bark, *aphyo" meaning becoming white and *dendron" meaning tree. 1 specimens examined. MEXICO. CHIAPAS: Tenejapa, paraje of Koltol Te', 4,750 ft., 10 July 1964, Breedlove 6121 (US); Unión Tuárez, Bosque de Pino, 1,360 m, 18 Nov. 1977, Calzada et al. 3711 (F); along Mexican Hwy. 195, 2 mi. N of Colonía San José, 8 mi. S of Layón, 1,500 m, 3-4 June 1973, Han- sen et al. 1689 (BH, US, WI S); Mt. Tacaná, 1,000- 2,000 m, Aug. 1938, Matuda 2466 (F, US); Mt. Pastal, 12 Apr. 1948, Matuda 17683 (F); near Finca El Sus- piro, 11 Jan. 1953, Miranda 7654 (US); near Tumbala, 4,000—5,500 ft., 20 Oct. 1895, Nelson 3332 (US); Finca Mexiquito, July 1913, Purpus 6980 (F, US); Cerro del Es e dog 1914, Purpus 7318 y (mixed collec- tion w coticosmum Bitter paraje iss K anal 8,400 ft - Z + Q E m (US). HIDALGO: Chapulhuacan, 1,300 m, 12 July 1937, Lundell & Lundell 7192 (LL, NY). JALIsco: SW foot- hills of the “utasa de Colima, ca. 10 mi. SW of Aten- quique on the Tonila road, 1,600 m, 5 Apr. 1951, McVaugh 1 1301 (NY, US); above Ahuacapán, road to Corralitos 10-12 mi. SSE of Autlán, 1,500-1,800 m, 29 Sept. 1960, McVaugh 19586 (US); 12 mi. SW of Chante along road to Sierra Manantlan, 5,500 ft., 4 Feb. 1975, Gentry & Gentry 23510 (US). MExiCO: Te- mascaltepec, Tejupilco, 1,340 m, 9 Apr. 1934, Hinton et al. 5753 (LL, NY, US). MORELOS: Sierra de Ocuila, 18 Sept. 1941, Lyonnet 3329 (US). NAYARIT: along Hwy. 28, between Tepic & Jalcatlán (Rt. 66 on some maps) at Km 14-16, 950-1,050 m, 8 Jan. 1979, Croat 45256 (MO). oaxaca: 10 mi. S of Sola de Vega along Puerto Escondido, 7,000 ft., 8 May 1965, Breedlove 9855 (US). vERACRUz: Huatusco, road to Coscomatepec, 1,580 m, 14°1 O'N, 97°00'W, 10 Aug. 1979, Avendaño & Calzada 411 (F); environs of Ori- zaba, Bottieri & Sumichrast 1857 (P); Jalapa, 6 Sept. 1936, MacDaniels 934 (BH, F); Xico, 6 km NE of Xico, 1,500 m, 19?27'N, 97°01'W, 12 May 1973, Marquez & Gandara 101 (F); near La Calavera, 10 km N of m by road) on road to Tlapacoyan, 1,350 m, 19551'N, 97?13'W, 28 June 1980, Nee & Han- sen 18643 (BH, F); 2.5 km (by road) E of Ayahualulco & 1.6 km (by road) W » Ixhuacán de los Reyes, 1,900 m, 19?2'N, 97*08'W, 7 Nov. 1981, Nee 22959 (P); Jilotepec “El Esquilón, ? 1,390 m, 7 Jan. 1976, Ortega cuatla, Lomas de Santa Rita, 1 m, 3 June 1971, Ventura A. 326 (F); Atzalán, La Florida, 1,600 m, 20 Mar. 1978, Ven- tura A. 151 GUATEMALA ALTA VERAPAZ: near V Coban, 1,260-1,440 m, 26 Mar.-15 Apr. 1939, Stan- dley 69240 (F, US); 3 km W of San Julián, Tactic, 1,600 m, 15?20'N, 90°15'W, 1 Feb. 1969, Williams et al. 40415 (F). BAJA VERAPAZ: below Patal, 1,550 m, 4 KNAPP—SOLANUM SECT. GEMINATA 567 Apr. 1941, Standley 90958 (F). CHIMALTENANGO: road to Iximche Ruins, Tecpán, 2,500 m, 12-23 Jan. 1966, Molina R. et al. 16128 (F lage of Brujo, 1,500-2,000 m, 1 1939, Steyermark 30939 (F, US). EL PROGRESO: Sierra de las Minas, between El Jute op Cobara & Finca Pia- montes, L 400-2,400 m, 3 Feb. 1942, Steyermark 43381 F, NY). ESCUINTLA: between ard Maria de Jesüs & Palin, 1,800 m, 29 Dec. 1938, Standley 61307 (F, US). GUATEMALA: Km 28 F.D.R. between San Lucas & Gua- temala City, 2,000 m, Molina R. et al. 16657 (F, TEX). HUEHUETENANGO: 3 mi. NW of Santa Cruz Barillas along road to San Mateo Ixtatlan, 6,000 ft., 7 Aug. 1965, Breedlove 11651 (F); yry a ft., 15 Dec. 1934, Skutch 1949 (F, NY, US), 13 km W of Hue- huetenango near Puente de Xinaxó, I i m, 30 Dec. 1940, Standley 81575 (F, US); spring of Río San Juan near Aguacatán, 1,800 m, 6 Dec. 1962, Williams et al. 22501 (F, NY, US, WIS). QUEZALTENANGO: Los Positos of San Martín Chile Verde, 1,500 m, 8 Mar. 1939, Standley 67901 (F, US); vicinity of Fuentes Georginas, slopes of Volcán de Zunil, 2,300-2,500 m, 3 Feb. 1941, Standley 85865 (F, US); w slopes of Volcán de Zunil, opposite Santa M 2 ú 500 m, 21 Jan. 1940, Fi Mee 3512 portion o Chiquimulilla, 250-2,000 m, 20 Dec. 1939, Steyer- mark 33149 (F). SUCHITEPEQUEZ: SW lower slopes of Volcán de Zunil between Finca des Asturias & Finca Alto Mira, NE of Pueblo Nuevo, 1,000-2,000 m, 1 Feb. 1940, Steyermark 35349 (F). HONDURAS. COMAYAGUA: Montaña Le Choca, Cordillera Comay- FE near Coyocutena, 1,200 m, 22 May 1956, Molina R. 7113 (F); Montana El Cedral, Cordillera Montecil- ‘ortés, (MO). MORAZAN: Mt. San Juancito, 7, T 1948, Glassman 1666 (F, NY, TEX, Uyuca, 1,600-2,000 m, 10-20 Mar. id y; 7167 (US). OCOTEPEQUE: around Belen Gualcho, 40 km E of Nueva Ocotepeque, 1,500-2,000 m, 29 June-3 July 1976, Nelson et al. 3817 (MO). vono: El Portillo Grande, 4,000 ft., July 1937, van Hagen & van Hagen 1006 (F, NY); Quebrada Olotillo, 15 km from Yoro, 1,100 m, 8 May 1956, Molina R. 6829 (US). Er SAL- VADOR. CHALATENANGO: E slope of Los Esemiles, 2,100- 2,300 m, 14?21'N, 89?09'W, 1 Apr. 1942, Tucker 1186 (F, LL). SANTA ANA: along road to Cerro Monte Cristo at Los Planes at Km 22, 1,800 m, 31 July 1977, Croat 42332 (MO). SONSONATE: Cerro Verde, 1,800 m, 25 Feb. 1968, x R. & Montalvo 21726 (F, NY). Ni- CARAGUA Mombacho Volcano, 960 m, 5 July 1923, yrs et al. 7781 (F, US). JINOTEGA: along 568 Hwy. 3 from Jinotega to Metagalpa, 5-8 mi. SW of Jinotega, 1,500 m, 7 Aug. 1976, Croat 43055 (MO). mi. NW of Zarcero, 850 m, 15 Aug. 1977, Croat 43541 (MO); along Río Alajuela, Alajuela-Carrizal road 5 km S uen 1,200 m, 24 Mar. 1974, Hartshorn 1423 (F, MO); Monteverde, road to Peñas Blancas (over Continental Divide) ca. 1 km below the Divide, At- lantic slope, 1,400 m, 10?25'N, 84*50'W, 13 Apr. 1981, Knapp & Mallet 865 (BH, CR, to be distributed); Santa María National Park, Caribbean slope, 600 m, 10?27'N, 85°17'W, 7 Feb. 1978, Leisner 5139 (MO); Zarcero, Palmiras, 6,000 ft., 16 Aug. 1937, Smith A152 (F, MO, 332 (F, 84°01' w, Lent 3160 (F, MO); El Muñeco on the R Navarro, 1,400-1,500 m, 6-7 Mar. 1926, Standley 4 Torres R. 50939 (US); near La Sierra ca. 25 of Cartago, Cordillera de Talamanca, 2, 000 m, 23 Jan. 1965, Williams et al. 28048 (F). GUANACASTE: along road between Santa Elena & Monteverde, ca. 2 mi from Santa Elena-Monteverde junction, 1,350 m, 7 Feb. 1979, Croat 47100 (BH, MO). HEREDIA: vicinity of Alto la Palma between Paracito & Bajo La Hondura, 1,500 m, 14 Jan. 1978, Croat 44484 (MO); Vara Blanca de Sarapiquí, N slope of Central Cordillera, 1,500- 1,750 m, July-Sept. 1937, Skutch 3286 (MO, NY, US). PUNTARENAS: Monteverde, pasture edges in village, 1,400-1,500 m, 15 Aug. 1976, Dryer 566 (CR, F); Fin- ca Las Cruces, 3 km S of San Vito de Java, 4,000 ft., 11 Feb. 1971, Gillis & Plowman 10129 (F, TEX). SAN Sept. Grande, 3 km NW of Cascajal, 1.750 m, 12 Dec. 1971, Lent 2295 (F, MO); hills above Aserri, 14 June 1955, Schubert & Lt 701 (US); vicinity of El General, 1936, Skutch 2987 (MO, NY Ltn o = = - N — D'Arcy 6492 (F, MO, NY, P, R Hornito, 4,500 ft., 8 May 1978, Hammel 3028 (MO). VERAGUAS: Cerro Tute, E slopes, 1 km beyond Escuela Agricola Alto Piedra above Santa Fé, 900-1,200 m, 14 May 1981, Sytsma & Andersson 4644 (MO). VENEZUELA. MERIDA: Quebrada El Oso, Palmira, Justo Briceno, 1,600 m, 3-11 Oct. 1973, López-Palacios & Bautista- Bautista 3481 (MO). COLOMBIA. ANTIOQUIA: between Río Guapa & Río León, 1,000 m, 18 Mar. 1948, Ruíz Landa et al. 118 (US); 4 km from Palmitas, 1,700 m, 5 Mar. 1949, Skolnik et al. 19An168 (F, US); road between Yarumal & Valdívia, region of Ventanas, (Gutierrez et al.) Valdivia 5 (US); road between Cura- monta & Valparaíso, (Travecedo & Gouzy) Valparaiso ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 3 (US). cauca: Popayán, Timbio in Hato-Viejo, 1,800 m, 14 July 1939, Cuatrecasas & Pérez-Arbeláez 6070 (F, US); Cordillera Occidental, E slope, Cuchilla de ipi 1949, Idrobo & Fernandez a ` Micay valley, near Río Joa- quim, Cordillera Beal 1,400-1,500 m, 29-30 June 1922, Killip 7846 (NY); Popayán, 1,500-2,000 m, Jan. ma “apauni CUNDINAM : David 4852 (US); Santana station, above Sasaima, 16 1,700 m, 25-29 July 1945, Dugand & Jaramillo 3877 dg Cordillera Oriental, Finca, “Alto Oscar" 11 km La Palma, 5,600 ft., 10 Mar. 1944, Litle 7388 Y, US). HUILA: Finca Cedral, Quebrada Urraca, Río E above Vega Larga, 30 km E of Neiva, 1,300 m, 2°58'N, 74°58' W, E 1943, Fosberg 19765 (NY, US). SANTANDER: Río Suratá valley 2,000-2,500 m, 5-6 Jan. 1927, "Killip & Smith 16531 (NY, US); vicinity of Charta, 2,000-2,600 m, 1-11 Feb. 1927, Killip & Smith 19089 (NY, US); vicinity of Tona, 1,900-2,100 m, 17 Feb. 1927, Killip & Smith 19505 (NY, US). Tota: El Líbano above San José, 1,580-1,620 m, 19 July 1947, García-Barriga 12239 (US). VALLE DE CAUCA: Cordillera Occidental, E slope between Bitaco mbo, 1,700 m, 5 Apr. 1979, Cua- trecasas & Cuadros 2883 7 (US); Río Nima above Ten- jo, 1,850 m, 2 Oct. 1974, Maas & Plowman 1827 (MO). EcuADOR. Loja: Km 25, Loja to San Lucas, 2,200 m 15 Sept. 1961, Dodson & Thien 635 (MO, US). PERU. sin. loc. 1840, Mathews 3249 (NY). AMAZONAS: Bon- gará, Jalca zone 3 km S of Pomacocha, E of Shipas- bamba trail, 2,400 m, 20 June 1962, Wurdack 997 6 5 N e° 2 E massif of Cordillera Central, 1 : 73°50'W, 20 Aug. 1968, Dudley 11879 (F); Ccarrapa between Huanta & Río Apurímac, 2,200 m, 5, 6, 17 May 1929, Killip & Smith 2306 (NY); La Mar, just below Huanhuachayo on the Capprichio-Puncu trail, W slope of Río Apurímac valley, 1,590 m, 12?43'S, 73°47'W, 15 July 1970, Madison 10265—70 (F). cuzco: *Pillahuata" Cerro de Cusilluyoc, 2,400-2,500 m, 3- 20 May-1 June 1923, Macbride 4155 (F); ‘road from +e uco to Tingo María N of Carpish Pass, 54 km E of Huanuco, 2, 310 m m, 6 Dec. 1981, Plowman & a 11164A (F). JUNÍN: Tarma, at little bridge on Tarma-San Ramon road, | km above Matichacra, just below Carpupata, E side of Rio Tarma valley opposite mouth of Rio Huasihuasi, 11 km NNE of Palca, 2,000 m, 4 Dec. 1962, Iltis et al. 322 (US, WIS); Pichís trail, between Yapas & Enefias, 1,800 m, 28 June-8 July 1929, Killip & Smith 25618 E Schunke hacienda above San Ramón, 1,300-1, Schunke A101 :S Y of Río Res pta (near w Calisaya) 750-900 m, 1-22 July 1939. rm Mff 104 F, MO, NY, US). SANTA CRUZ: Rí Piraymiri, a 5 nde, 2, o0 m, Feb. 1956, Carde- nas 5118 (US). 1985] LITERATURE CITED BRANDEGEE, T. S. 1917. Plantae Mexicanae Purpu- sianae VIII. Univ. Calif. Publ. Bot. 6: 373. BucH, C. L. von. 1825. Physicalische Beschreibung der Canarischen Inseln. Berlin. CHILD, A. 79. A review of branching patterns in the Solanaceae. Pp. 345-356 in J. G. Hawkes, R. N. Lester & A. D. Skelding (editors), The Biology and Taxonomy of the Solanaceae. Linnaean So- ciety, London. 1958. Der Verzweigung der Solanaceen in reproduktiven Bereich. Abh. Deutsch. Akad. Wiss. Berlin (K1. 1957) 6: 1-18 67. Die Verzweigung als infragenerisches ruppenmarkel i in der Gattung Solanum L. Kul- turpflanze 15: 275-292. D’Arcy, W. G. 1972. iji ee studies II: typifi- cation of the wow: i Solanum. Ann. Missouri Bot. Gard. 59: 262- KNAPP—SOLANUM SECT. GEMINATA 569 1973. Solanaceae. Jn R. E. Woodson & R. S. Schery (editors), Flora of Panama. Ann. Missouri Bot. Gard. 60: 573-780. Knapp, S. 1983. Sectional nomenclature in Solanum (Solanaceae). Taxon 32: 635 Reproductive biology of Solanum sec- a Costa Rican G. D’Arcy (editor), Solanaceae: Systematics and Biology. Columbia Univ. Press York. SEITHE, A. 1962. Die Haararten der Gattung Solanum L. und ihre taxonomische Verwertung. Bot. Jahrb Syst. 81: 261-336. . 1979. Hair types as taxonomic — in Solanum. Pp. 307-319 in S, Lester & A. D. Skelding (editors), The B Biology and Taxonomy of the Solanaceae. Linnaean Society, London. POLLEN MORPHOLOGY OF THE GENUS ERYTHRINA (LEGUMINOSAE: PAPILIONOIDEAE) IN RELATION TO FLORAL STRUCTURE AND POLLINATORS! A. J. HEMSLEY AND I. K. FERGUSON? ABSTRACT The unacetolysed pollen of 99 species of Erythrina was studied with scanning electron microscopy. The exine sculpturing is basically reticulate but displays considerable variation in several features. Pollen types may be divided into those species with tubular, hummingbird-pollinated flowers and E with gaping flowers, pollinated by a variety of passerine birds. eaii type” pollen may iin as medium-sized, with medium-sized lumina and sparse or no sexinous granules. Pollen a passerine-pollinated sections is more heterogeneous but may be distinguished generally by mod- erate to ruin granularity or small i sparse or lacking in hummingbird groups. These observations are discussed and possible explanations suggested Palynological contributions to taxonomy, in disclosing and suggesting relationships between taxa, have emerged hitherto principally from comparative study of structure and ornamenta- tion among pollen. Recently, however, attention has become focused increasingly on functional aspects of the structural details observed. The environmental pressures many features of pollen structure and physiology may lead to failure, or conversely promote success, at any stage. Thus, 1t seems reasonable to expect that pollen, subject to natural selection Just as much as the whole plant, v; evolved to meet the demands imposed upon Bue the situation is unlikely to be simple. A plant is constrained within certain limitations m phological features: for example, an inefficient pollination system may be compensated for by vironmental conditions are both continuous chance but not all features are nec- essarily of any immediately obvious adaptive significance. In a relatively close-knit genus, such as Ery- thrina, it seems reasonable to assume that the species have radiated relatively recently from an ancestral stock and have preserved a basic sim- orlds. Although all species are pri- marily pollinated by birds of one sort or another (Toledo, 1974; Raven, 1979), there seems to be distinct syndromes in floral biology within the genus, apparently associated with pollination p o modifications of the pollen itself might be ex- ' We are greatly indebted to C. H. Stirton (K) for suggesting the topic and for much initial guidance and to Botanic d the and we especially thank the Bentham-Moxon Trust for support to AJH to allow com- pletion of the su ? The Herbarium. " Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, England. ANN. MissouRi Bor. GARD. 72: 570—590. 1985. 1985] pected. However, climatic, ecological, or other factors may also influence evolution, and ran- dom variation, whether genetically or environ- mentally induced, may also occur. Ferguson and Skvarla (1982) have shown that in the subfamily Papilionoideae there is a close association between pollen ornamentation and stratification with flower structure and pollina- tion system, notably in some species with flowers adapted for pollination by birds or bats. This association between pollen morphology and flo- ral structure occurs in species and genera from widely separated tribes, and it is suggested that it is the result of convergent evolution and a secondary adaptation to pollination. The pollen 1977) made a detailed study of the pollen of Erythrina in relation to taxonomy but without consideration of the approach discussed above. These workers gave an extensive bibliography of the pollen morphology of the genus, which can be supplemented by reference to Thanikaimoni (1976, 1980) The aim of the present study, using herbarium material, is to examine the links between pollen morphology, flower structure, and pollination ecology. MATERIALS AND METHODS Pollen was obtained primarily from herbarium specimens at the Royal Botanic Gardens, Kew (K). Material was also obtained from the her- barium of the New York Botanical Garden (NY) Table 1. The data viuis) in Table 1 are based on scanning electron microscope (SEM) micro- graphs of untreated pollen removed directly from herbarium sheets. This procedure was followed because, although acetolysis (Erdtman, 1969) in most cases somewhat improved the clarity of the ornamentation and increased the size of the pol- len grains, it removed the residual pollenkitt found on many unacetolysed pollen. Differences in the occurrence and volume of pollenkitt pres- ent in pollen samples were found to be of some significance. This will be discussed later. The pollen was prepared by removing mature anthers Hom herbarum a sheets and shaking or uu ersten oti thle-cided sticky tape on SEM: specimen stubs. The samples were sput- ter coated with platinum and examined with a Jeol T20 SEM. HEMSLEY & FERGUSON — ERYTHRINA POLLEN MORPHOLOGY POLLEN MORPHOLOGY From the 99 species of Erythrina examined in this study, and from the results of Graham and Tomb (1974, 1977), the pollen may be described as follows: oblate; rounded-triangular in polar view; ranging in size from 22 to 55 um diam.; triporate, occasionally tetraporate, pores more or less equally spaced, equatorially arranged at the corners; surface ornamentation basically reticu- late, but with considerable variation in the size and shape of the lumina, the height and width of the muri, and the occurrence and prominence of sexinous granules in the lumina. In occasional samples the muri are discontinuous on either the polar or equatorial faces (e.g., dominguezii, E. lysistemon). Rarely is there little or no orga- nized reticulum, and the ornamentation consists of discontinuous muri or sexinous granules. Each of the main features of variation in the pollen morphology is discussed briefly in turn and the data for each species are summarized in Table 1 SIZE AND SHAPE The pollen diameter of the majority of species falls in the range 23—30 um, as measured on elec- tron micrographs of unacetolysed material [these sizes are proportionally smaller than those given by Graham and Tomb (1977) for acetolysed pol- len A few species have pollen that is smaller than average, 20-23 um diam. These include E. cris- ta-galli, E. arborescens, E. breviflora, E. edulis, E. suberosa, E. stricta, and E. resupinata. Ery- thrina eggersii has atypically small pollen among the six sections characterized by tubular, hum- mingbird pollinated flowers in subgenus Ery- thrina, which otherwise have medium, rarely medium- to large-sized pollen. Large pollen, 30—35 um diam., occurs predom- inantly in subgenus Erythraster, in a number of species in subgenus Chirocalyx, and in the mono- typic section Acanthocarpae in subgenus Ery- thrina. The pollen of E. fusca is particularly variable in size. Large, medium, and small pollen are all represented by different samples (see Table 1). The shape of the pollen of the vast majority of species is rounded-triangular in polar view. However, that of a few sections, i.e., Suberosae, Arborescentes, Breviflorae, Leptorhizae, and Oli- viae—all subgenus Erythrina, is markedly tri- angular, while that of other species (e.g., in sub- genus Erythraster) is more rounded. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 572 - u IS s d e s (D eIpu] APESOZ 244012 'QXOWw $422$240q4D `q Jo»xnry $27422$240q4y. 1298 F w s s d e s (SD eipu £94 (2uoojw + w/u stp s/w d ə s OD eIpul ¿£ ¿F£ 234D]O `qxoq pjpuidnso4 ^q + w s s/w d e E OD epu $46 X201Mg "qxosp DIAS 7] uo} + ul S S d e ul (SD (1m9) eAef "us LOYD -IJL A %@ SIOPIOOY Dd4D72042114 ^q ++ u sIp+ s d e s GD erpu LL 42400H `qxow rso42qns ^q POJMY apso42qns 1598s Dui: `3qns + ur 1 ul d q u OD €Iquio[o/) 00IS "JP 12 səllnt/9ç yoo) + ul 1 uil d q ul (SD p[ƏnzZƏuə A 80£ 42]pua4 ‘J 'O (s1od[e A) Dunisiddaod `: + q 1 w d 2 u OD Izeg Z-818 lyong OSO[[SA Du424 `T + u sIp I d 3 tu OD nod WILEC AUDId suueH 12]n "T Ta ul sip ul d 9 ul OD eunuoary OFOS 14n1u2A4 Jo|sseg :1zan8unuop `q x(421d0421JA 1298 + ul 1 ul d 5 s/w OD eunuosiy €046 14n1u2A4 "pug 77Dojpf d = ul s s d e s (D punuəsiy 0c 8 uas4apad "UI 1/108 -01S142 ^q Jjo»xnay 17/03 -201814 1298 + u sip I d 3 I OD [zesg C848p umar + ul Sip S/UI d 9 S/UI OD e[eurojenr) ‘u's XnT X apu + ul 1 SA d e S OD `S[ OIOWIOZ F6€I Ipuraqappng + tu 1 SA d e I D BouInt) MƏN 8I8ET PIW ++ q SIp ul d P u OD eÁe[e]o 8r£ funy OII9InO'] p2sn/í `F yoynry (s19d[eAA) visuipsspyong 1298 Joxeg `O `f (sji9d[e M) x«a22d0421Jy "qns asom PPIM sd pÞƏZIS Ol OA ,2ZIS pə Áy[eoo'T uono səroəds -ug Uny -33A -usodəaq snou -Ixag eurum uod uM emu UlOJJ POALIOp UIQ Ə9Aeu pu? 2ATj[21 21e səzt8 "9S[q9[IeA* ÁA[rpea1 JOU SPA [ELI9]EUI 9[Q?]Ins əsngoəq pouluiexo UIQ 1ou seu K8o[ouduoui II. u L. `səroəds jo Iep IUL '(p/61) Áqoureg pue? pgo»xnry JO Japso orurouoxej eui ur — ose sorxods eq L `S1O199A pue ‘sodA} siojpereqo uojod ureu oy} jo Kreurums e pue uuo JO s[rejop YIM poeururexo suauroedgs '[ ATAV IE HEMSLEY & FERGUSON—ERYTHRINA POLLEN MORPHOLOGY 573 1985] * u IF u H q tu (AN) O9IX9JA 64-0461 ffoxniy Aqouseg ?g goxnry voipnd `q * u 1 u H q u (AN) OO9IX9JA[ OET -0461 Hony ‘OC S2p10j]402 "ge `J + w JS I H q u on Oo9IX9JA[ FCI-0461 fJosnay = u IS I H q w OD O9IX9JA 8cI Joynay ‘OC səplo]]p4io2 `q - ul 1 ul H q ul OD OOIXOJA[ C0cc «au21) ÁouJe2N siu40ft]jaqp]f ` = w 1 u H q tu on O9IX9JA[ £L-0L61 ffoxniy Joxnry pubojpuvis `q &qoureg 7 HOJ — ul IS ul H q s/w (D OOIX9JA 969 Dso41404 -NITY D2$040481u “dss n22Dq424 `J Dua `199S = u 1 SA H e w OD OSIXƏJA L89F ABU "Od 9Pud0w 7 9¿9rg — u 1 s/w H q I (SD OOSIXƏ]JA] ADIANOD X spg ‘OC Pz14401d3] `q — u IS s H q ul (SD OOIXƏJA] OS 42w]pd ÁƏ9[pue1S ?p 3soy punjuou `q Jjo»xnry avz1ysojdaT 1298 + u IS s H q/e ul OD Jopenoq £F9£ `F duvo spo nffawrnjos `q — — — — — — — — — — suneH Djapyodjod `q Áqouueg W goxn.ry səjnpə-opnəsq 7199s + ul S s H q/e ul OD [rzeg "ww$ uiapy P 211124 SAAƏ1puv Dsomods `q Joynrs ( 3sseH) sidoilouə1ç 1298 — u IS S 4 e s (SD Jopenod F69 apgovyw euer] SIjnpa ^ Jioxnry sampa “199s HOA x u IS E H e u (ON) O9IX9JA[ 0ç9ç JN -nry 9 Aqouseg wniqojoing q — ul JS SA d e s (D OOIXƏJA] C8SET t/SnpA2JWw _ w Is SA d e s GD OOIX9JA OIC£I u3npAoIN ‘Od ?40[fià24q 7 Jjoxnry ar4opfia24g 1298s T u IS S d 3 u OD səuiddrirtudq OS6T 1142W ++ ui JS s d 5 u (D puejreu L @0¿ZI ‘JP 12 uasua4eg IHIN (3SSeH) supaquanqns `q Jjoxnry (»sseH) sraoudod«g 71298 sS9o[n PPIM — dA] »9ZIS 0} gQ9dÁ] ,.ƏSZIS pə ÁA1tIeoo T uono sar oodg -ue MWN -99A -iÀodaq snou -1x35 purum uə[[oq 'pəonunuoO “| z18v[ Áqou N P — ul IS ul H q ul OD e[eurojenr) €-C461 ffoyniy -Ieg ?? Ponary vuvosanbivg ^ $ = u 1 u H q u OD Bory BISOD FPIIII ?2242N HIYAW 5284221401802 `F B _ w/u Is w H q ul OD seJipuoH 149 Spaompy Áo[puelS 27DJO22UuU] `q Aqouieg = uu s u H q u OD ejeurojenre) 8-£ 461 foynsy 7? pony sisuasupuajanyany `J — ui 1 ul H q ul (AN) OOTX9JAI LES Aoduog-zawuoy Aqouleg ?p yoxnaw $2214242q `F * uu IS ur H q u OD erwan Z-1 L61 ffoonaxy Aqəureg 7 POJATA 2r12u240]f 1 Aqou — ul 1 ul H q ul (AN) ENZeIVSIN] £Z09 suaaaig -Jeg 8 POJMY :D/401442421$ ^41 + ul IS u H q u OD gry €1SO,) IF 1-6961 foynsy `PojnO 79 uos1og x4jp2oqoj2 `q Z = uu IS u H q ur OD e[eurojenze) 1£-6961 foynsy POTY sisuəjpuolpn2 77] Q F w Is w H q ur OD eewo £-6961 Joxnay OG vjjiudo4orui ‘q < = uu IS ur H q u (AN) eueued L6F0I 10049 POY sisuanbinpo ^q - Aqou S _ uu 1S u H q u CD OoTX9JA £9-0461 ffoyniy -1eg 79 goxnay sisuaopuunfp] `q Z + wu JS u H q u (AN) e[eurojenzc) 09c0F SuD M Aqəureg V POTY zstupijlu T < Áqou ° E u IS w H q w OD e[eurojenr) I I-0461 foynsy -Ieg P POJMY sz$u2unrqoo `; = Aqou = F q s u H q ur OD e[eurojenr) €4I-6961 ffoonaxy -Ieg ?g PONY s1$42u7])D `I 2 Aqou = _ uu 1 u H q pu OD ewm çIZ-6961 JOJY -1eg 9 goxnry vuvsvdviys `q I = q s w H q ju c = q stp u H 3 wu (AN) sejpuoH CIOL s42punpg Kajpueis sisuasnpuoy `q Š + u IS u H q yu (AN) Bory B1SOD COTS 12N KajpuelS D772/1/202 `T lai = u JS u H q w Op Jopensy 96 2pagovjy Jjoxnry nuntius “7 = = ul Is ul H q ul (AN) OOIX9JA [IZZ 0 zajpzuoy Aqouseg ?g jyoxnry pupxn] “7 2 = u JS u H q ur OD O9IX9JA 16-0461 Joynsy exuopIoJA 7? YOY 11942»]0/ `T + u 1 I H q tu OD OO9IX9JA €6-0461 foynsy Aqouseg 7 joxnry vanquys `T = u sIp+ w H q u OD OoTXOW LL-0L61 foynsy Ag[puelg uunupjos `q KaJ = ul 1 u H q ur (D OOIX9JA 99ISC `]? 12 5401S -PURIS s1[D1u2p1220 ‘dss pipup) `q + u IS u H q u OD OOTXOTA 0£00I uomiH soy vun) ‘dss DIDUD] `q səm — ,QPIA «= OAL. PSZIS — JO) QdÁ[ ,ƏZIS pə Kipeoo'T uono səroəds uw MW -99A -isodaq inde €urumnT ulod -IXƏS : 574 'penunuo;) ‘| dav HEMSLEY & FERGUSON — ERYTHRINA POLLEN MORPHOLOGY 575 1985] ¥ w IS ul d 3 ul (SD ƏAqequurz 9ç[ uaniiajyy + ul S ul d q uu (D ƏAqequurz 088 25742) “Yon uO142]S1s4] ` + u IS tu d q u OD BOLYY 'S 8864 PP?) 'qunu[ 24ffpo 3 Joynry X Aqoweg apaffpD 1298 - uu S s/w d e ul (AN) OOIX9]JA Z zapuvruuaH Jjoxnry 2r110 `q Áqouueg Y gyoxnry əpupiua1O 1298 x ur 1 w ó q Uu OD eqn) $886 112m0 P uonug WBUM `O sisuaqno `q Joxnry sasuaqnyD 1228 = — — — — — — — — — s33ug 77 pIeMoH 22u9J2 “J a wu S uu H q w OD neH ‘U's UDUY uewyg 2 ueqJ() Dpodojdaj `q = uu 1 tu H q w (AN) eH Lt6 23pup]oH ueqJ[) nyong ` I0ç6 F w/u IS w H q S (AN) oory oLang uonug y uonag DUSP[OW #@ Joxnry 1542332 `J + wu IS uu H o ul (D 1uƏOSUIA 1S [ZZ uyoH "1 "unapuapojr]aoo ^q T uiu IS ui H o ul OD pepruu FI9€ (pappoig + uu IS ui H 2 u OD pepruu L ws SPADYIIY asoy X Uong vpıpd `q = uil S ul H q uil OD e[9nZ9US3A POF 42]pua urnboe[ sina ^q _ wu s w H q w (AN) lopenoq FCS$6I punjdsp Jgjoxnry puprinaod ^q = uiu S w H q wu (AN) enog u's azjuny YOFNTY $1718 `F = uu I ul H q w (SD Jopensq SIS əp(ugopiw Yoynry p3tuoznpuap 3 Jo»xnry p4puapojjp40;) 1298 — u 1 ul H q Ww OD eureued POE 142q21S ‘POJNO vsoqqis ` Áqouaeg #@ goxnry 22s50qq1£) 1298 Aqou = ur/u s u H q uu OD e[eurojenr) £££-6961 JJoyniy -Ieg ?9 pony 240]f1/2]]802 1 * u s u H q ui OD e[euro]enr) LS-6961 f#foyniy Aqougeg V goxnay p4ojfita]Us `F = uu S ul H q yw OD OOIX9JA 9/9 2ssp]auv'] Joxnry pupomxəta cu + uu J u H q wu OD E[ƏnZƏU9 A £8C I f#foynay AQ H VIULUA ` s uu 1 u H q wu OD Bory EISOD 101-6961 Joynsy ueqI() pupo421432q `T — ur/u IS ur H q ul OD OOTX9]AL "u's anala IN 2uno142040 `J sM PPIM OAT, p2ZIS 30] q«ƏdÁL —.2zig pə Áureo Uuoloə[|loO sərpəds -URIO uny -33A -isodoqg snou -Ixag eurn ulod 'Penunuo) 'I 318v] ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 576 pueun(q + ur Js s d 2 yw OD UOOJIQSUIE;) 040€ xnə.T W WPM pupisuputuSooip `A F q is SA d e w D eueyo £8I I 2u81A "W9InH avuosippo I = T = E e — a — x = enH pupluo]]ol/] `T + ul IS u d q I OD omqgequilZ £8c Sdjayd Joyeg vupiuosssuial “J + w IS u d 3 ul OD ROLY "ALS 69£ Japtag SULIeH{ D4029p `T = - — — — — — — — — SWJeH :z1unpq `q $Cc9 uos + q IS ul d 9 ul (D eIquiez -YOO) Y puouuunaq dO], 11Səpuətu "T = — — — — — — — — — ono vanwsdd `q XAJDIOAIY 1298 + u s s/u d 3 ul OD 9UOvT LIIS F88£ uonjdiaq suureH 71pav4qpina `T suueH uopadspa20[nf21q 1298 + q s wu d ə yw OD epues[() 848I Uapmous’ Joxeg Dsj2oxo "T suueH snjnj420qopiq 71598 + q Is ul d o ul (D QUOI] E.LIƏIS C8 x40X "Od sisuajpaauoas `q ++ q sip I d P yw OD ELIƏSIN "ws (uojojy ‘J OOH 1172804 3 Joxnry X Áqouaeg (S12d[e A) tuniquí4A20420JA. 129 Ter ul IS u d ə I OD eidorqi £489 (2u00jy JULSMYIG 120n4q "T Joxnry ?? Áqoureg apunaonag 1298 Á9AJeH (1ousto]JA) x/(020414:) “3qns = — — — — — — — — — 'Op19A 1(DMu2248 `T snqojo421d14 J, "1298s Poyry wz Kqougeg 57:g0j0421d14 L, "dqns + ul IS ul d o I OD ROLY `S ‘US USAT DÁIN 4 pdupooulupop “yz ypoynry wz Aqouieg apdupd0yjuvop 1228 + q/ui 1 tu d q/e ul OD ROLY `S ‘U's qnjoH F w/u s SA d € ul (D vuy `S FISEI S422804 KAIGH 1421/42Z `J + tu IS ul d q u OD anbiquiezoJA LIZZ UDIJOUM + u IS ul d q ul OD eoXulv `S L6C6 PPOD "Suse1ds pupauunt `q Yoynry ?? Aqəureg əpupəutuungi 1228 INM PPIM AAL p9ZIS 0} «ƏdÁL ƏZIŞ pə Áji[e90/T uonoo[[0) sar oodg -uvo UNN -33A -yusodəaq snou m = -1x9§ eurumn'T uojod 'Penunuo) '[zs18v[] HEMSLEY & FERGUSON— ERYTHRINA POLLEN MORPHOLOGY 577 1985] MO] ut jnq əsuəp ə)mb əq souirrouios Ageu) juosaJd = + 'osreds K19A = F *jussqe = — '(unur 3y} ut sxyeajq 0} onp 1941980) ə8;əur gurung 319UA) snonuruoosrp POI y3 ur juoururoud pue BIE] JO osuap AIDA = ++ ‘Goro ‘so[nuess snourxos Jo Ajrsuop/oououruiroJd Iane pue 9ouaLmooQ) s "P£oq = q *umrpour = ut mone = u pm UN, , = SIP ‘punou pue snonuis = Js ‘snonuls = s ‘punoi = 1 :adÁa purum, ‘odse] = | 'umnrpour = ui ews = s ews ÁA = SA :ƏZIS purun p 'spiiqSururumq = H “Sspitq ouuossed = d :10193A uƏ||Od ; `1X91 398 :ƏdÁ} uo[[og q 'o&1e| = | 'umrpour = ui ‘ews = s :9zIS uo[ogd e T q E uis d p I Op eAuoy 99ç£ pjnoqman "QNEL pt/zupopunpjəauu ‘q = = — = = = — = = z suureH 142qoiuos "qr — — — — — — — — — — IMIA 1401442d ` + q IS pwu d p I OD eidomg ‘U's JOoOuptuuIDH = q SIp s d p I OD eidomiq SEP] 4284ng “AOTY) pupunq `T + q SIp w d p I OD ekusy 6256 (DAuə221D J 19388 Ming `q + q IS w/s d P I OD eqn) LFEC 84M Ueq1() MYIDgGasUs `T PF q IS I d P I OD gIquIO|oO L8Z A2041 &inqupg “PIMA Punja `T + q I rs d J I OD gumy MON CHIT 2ppouos Joxnry VUDI jU `] (w03 + q IS u d ¿P u OD eensny $668 puvj4H pues?) ‘yug orm42dsas 3 + q IS p/u d p I OD eiensny 6£ç uosuuof YMDIŞ "muog oi42ds24 7 + q 1 s d p I OD earl 988p Sqoovr 3sseH 2jjudipona `: + q I bs d J I OD nemĪmeH VCO6F] uvuaug DESpEN $7542710] "T Q)oouis/1e[nuej8 Ajauy svens d w OD gumy MON Et LZ əppouoç ++ q IS Ul d p I (D eruezue r "ws ISDT ++ q IS I d P I (D exueT us 960€ Ppny TT q IS I d P I OD pue[reu p, EECKI 4424 ++ q IS I d p I OD eneuNns EES uosulqoy "T pJD8214DA “J AIISDAY Ig `199S Jjoxnry Y Aqousreg 4275741147] "Sqns + w IS s d 2 ul OD eruezue[ 926 Aquio + ui 1 S d 9 ul (D anez 6£ 1208utq4 "ule T DOIu1SSÁQD ^q ++ q s I d P I OD 9^qequirZ PITT S2^0]d Taha ^S DiutSSHD] `T + ul s w d 5 s GD UOOJ9UIP.) 0£29 p4ofupg enH Doaptiotustis `q F ul IS ul d q Ul Op vruezue [, u's YMMI 9pI9A !!p42Du `q ++ ul S I d p I GD pruezue yl 6F9F (DAU2324D eng lxnapovs ` 7 + ul s ul d 3 wu (D anez 0£0$ asginbsayH 91ombsouo) nj1udo4o ‘q SS9[ PPIM —dÁ[ — ;9zIS O} qƏdÁL ,Ə2ZIS pe Kyie207T uono sətoəds uy MW -33A -1IsodəGq he euruinT uo[[od -IX9S : "Donunuo^) `I AIAVT 578 ORNAMENTATION SHAPE AND SIZE OF LUMINA The average width of the lumina varies from extremely narrow, ca. 1 um, to much broader, ca. 6-8 um, but with typical values in the range 1-3 um. The shape of the lumina varies from distinctly sinuous to more rounded in outline but wi thermore, the reticulum may closed, or the muri may have frequent free ends, making the reticulum discontinuous and irreg- ular. On the whole, these characteristics do not cor- relate readily with taxonomic groupings in the (e.g., compare Figs. 25-27, all sect. Chirocalyx, or Figs. 28-30, all sect. Erythraster). Isolated in- stances of pollen showing a discontinuous retic- ulum occur eed throughout the subgen- era and sectio The largest don in the genus, Erythrina, is also the most consistent with regard to lumina size, nearly all species in this section having pol- len with medium-sized lumina. This is true also of the closely related sections Corallodendra and Gibbosae. However, within this group of species characterized by tubular, “hummingbird flow- ` the small sections PER Pseudoe- dules, and Stenotropis are somewh t anomalous having pollen with markedly smaller lumina (Figs. 19, 20). Among the passerine-pollinated species, several small sections contain species that also have pollen with predominantly medium- sized lumina (e.g., Micropteryx, Caffrae, Hu- meanae). Other sections are marked by pollen with much smaller lumina [Suberosae, Arbores- centes, Hypaphorus, Breviflorae, Edules, and Oliviae (see Figs. 7-12 A particularly distinct itteni, in which very small round lumina occur on polar faces, opening out to much broader lumina and more discon- tinuous muri on equatorial faces, is found in only two species, E. tahitensis and E. merrilliana, both in sect. Erythraster. Erythrina fusca was found once again to be very variable with respect to lumina size (Figs. l, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 APPEARANCE AND SIZE OF THE MURI Differences in the size and general appearance of the muri occur. The muri are usually of mod- erate relief and width (ca. 1 um high and 1 um wide), somewhat rough and ridged in appearance on the sides and fairly smooth and domed on top. Also the sides of the muri may sometimes be more or less smooth (see Figs. 14, 16, 18). This is found in the majority of species. Particularly broad, flat-topped muri occur in some species of subgenus Erythraster [e.g., E. variegata (Fig. 36) and E. velutina]. However, in other species of the subgenus the muri are of high relief and are irregularly ridged (e.g., E. grise- bachii). A number of species in subgenus Chirocalyx, and also E. zeyheri, have very smooth muri sup- ported by long columellae that are higher than the sexinous granules in the lumina and give the ornamentation a characteristic rather raised ap- pearance (e.g., see Figs. 23, 25). In E. poeppigiana and E. cubensis, the muri are fairly broad and in low relief. Species of section Suberosae (E. microcarpa, E. stricta, and E. resupinata) have flattened, square-topped muri that give an impression of regular segmentation along their length (Fig. 4). SEXINOUS GRANULES The sexinous granules in the lumina of the reticulum show a complete range from virtually absent or of very low relief to extremely prom- inent and numerous. Four of the five specimens of E. variegata ex- amined showed the most prominent sexinous granules, these occupying most ofthe area within as the broad muri (Fig. 36). Ano respect was one sample of E. variegata (Schodde 2743) from New Guinea, which had no reticulate patterning on any of its pollen and an almost completely smooth appearance (Ferguson & Skvarla, 1981); this specimen needs further in- vestigation. A number of other species, mainly in the sub- genera Chirocalyx and Erythraster, but also in- cluding E. fusca (Haniff 348; Fig. 2) and E. do- minguezii, have particularly dense and prominent granules. By contrast, sexinous granules are ab- sent, or sparse and low, in virtually all species within the six sections characterized by tubular, 1985] HEMSLEY & FERGUSON —ERYTHRINA POLLEN MORPHOLOGY 579 FıGures 1-6. Whole pollen grains and exine surfaces of pollen grains from species pollinated by poem birds. 1-3. Whole pollen grains, x1,500; E. fusca showing variation in the pollen of the species. — showing small lumina (Millar 13818). —2. Type (d) showing dense sexinous granules (Haniff 348). d. Type ra showing more or less large lumina with discontinuous muri ce in beide 4—6. Exine surfaces, x 3,600. — Wt s showing pollen of Type (a) with sinuous lumina and segmented muri.— 5. E. arborescens showing Es of Type (a) with small, sinuous/round lumina.— 6. E. pa ie (Merrill 1950) showing pollen of Type (c) shine sinuous/round lumina and dense sexinous granules. 580 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FiGuREs 7-12. Whole pollen grains from species p by passerine birds, x 1,500. All Type (a). 7- pe l. Showing small-sized pollen.— 7. E. crista-galli —8. E. pinata (Mooney 762 E —9. E. arborescens. — 10. breviflora (McVaugh 13210). — 11. E. edulis. —12. E. ae medium-sized pollen 1985] HEMSLEY & FERGUSON — ERYTHRINA POLLEN MORPHOLOGY 581 FicunEs 13-18. Whole pollen grains and exine surfaces of eus grains from species “eran by hum- mingbirds. edi E 13,15,17, x1,500; 14,16, 18, x 3,600. 13, mina, sexinous granules absent. — 13. E . barqueroana. — 14. E. tajumulcensis. 15, lo Sho owing more or less iet lumina with sparse very low s exinous granules. — 15. E. pudica. — 16. E. aff. coralloides. 17, 18. Showing sinuous lumina with muri sometimes discontinuous. — 17. E. leptopoda. —18. E. mexicana. 582 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 . 22-24. From species pollinated by sunbirds.— humeana (Codd 9297) yu spe pollen of Type (b) similar to that found in m (see Figs. 13-18). 23, 24. E. zeyheri showing variation in pollen of species. nico, Showing pollen with rather smaller lumina and dde muri than typical Type (b) and sexinous granules (Holub s.n.). — 24. Showing pollen of Type (a) with small lumina (Rogers 14814). 1985] HEMSLEY & FERGUSON — ERYTHRINA POLLEN MORPHOLOGY 583 FIGURES 25-30. Whole pollen grains from species pollinated by passerine birds, x 1,500. 25, 26. Pollen of Type (c). — 25. broad muri. — 26. E. abyssinica (Hornb 926) showing atypically small lumina and sexinous granules. 27- 30. Pollen of Type (d) showing large-sized pollen with bro ad muri.— 27. E. latissima mowing large lumina and numerous sexinous granule es.— 28 discontinuous muri. —30. E. pee showing a 4- eae pollen grain with very discontinuous muri. 584 “hummingbird flowers" in subgenus Erythrina. Notable exceptions are E. pallida, E. corallo- dendron, E. speciosa, and a second sample of E. coralloides ( Krukoff 1970-124), which have moderate granularity. Outside these *humming- buit sections, granules are absent or low on the pollen of a number of passerine- id EC species, although their absence is fre- quently associated with smaller than average lu- mina size (e.g., see Figs. 7-12) or broad muri (e.g., see Fig. 29). However, species in sections Cubenses, Caffrae, and Humeanae (subgenus Erythrina), and E. haerdii and E. livingstoniana in subgenus Chirocalyx, have absent or low gran- ules and medium lumina size and muri width. All ne eae — ae or EXINE STRATIFICATION Thin sections for transmission electron mi- croscopy of acetolysed exines were cut offa num- ber of representatives from the range of variation found in the pollen to see if there were also dif- ferences in the exine stratification. No significant differences that were not already visible in SE studies were apparent. Some details of the exine stratification are described in Larson (1964) and Ferguson and Skvarla (1981) and further infor- mation will be illustrated elsewhere. INTRASPECIFIC VARIATION If comparisons of pollen types between species are to be valid, it is important to have some idea of the degree of variability within species. From the above results, this is evidently considerable specimens of E. variegata from different geo- graphical regions were found to be consistent. Erythrina variegata from New Guinea, with its lack of reticulation and small size, appears to be anomalous. One sample of E. resupinata showed incom- plete development ofthe reticulum with granules on the polar faces; and one sample of E. /ysis- temon had unusually low relief of muri and lack of granules, not unlike the ornamentation on the pollen from an anther of deformed pollen of E. goldmanii. Perhaps these features are abnor- malities of development. One sample of E. zey- heri had a much finer reticulum than another, and one sample of E. burana had broader lumina with flatter muri than the other. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 The most variable species found was E. fusca, which had both broad and fine reticula, large and small size, and one without any apparent orga- nized reticulum. All seemed to show evidence of moderate to high granularity. POLLEN TvPES The pollen of Erythrina can be divided into four main types described below. The pollen type to which each species is assigned is shown in Table 1. Two additional types (e) and (f) have been established for convenience. Type (a). Pollen usually small in size (rarely medium or large), often markedly triangular in shape, with small or very small lumina, usually without visible sexinous granules (Figs. 1, 4, 5, 7-12, 19, 24). This type is found largely in species pollinated by passerine birds with the exception of E. hor- rida. The species included are mainly in subge- Suberosae nus Bren and Edules. Erythrina speciosa and E. schimpfii are intermediate between Type (a) and Type (b Type (b). Pollen medium-sized (rarely large), rounded-triangular in shape, with medium-sized lumina, muri of medium width, very sparse or no sexinous granules present (Figs. 13-18, 22, 23 This occurs in the New World species polli- nated by hummingbirds that comprise sections Leptorhizae, Erythrina, Gibbosae, and Corallo- dendron together with S. African E. humeana and s zeyheri, which are pollinated by sunbirds. Micropteryx) and E. haerdii (section C hirocalyx) are pollinated by pou bir T c). L in medium, sometimes large, In tid with medium-sized lumina, medium to broad muri, pres Ee numerous and lower than height o This type occurs Fist in subgenus Chi- rocalyx, among African species pollinated by passerine birds (Figs. 3, 6, 21, 25, 26, 31-34). Type (d). Pollen large, lumina variable in size, muri broad, usually large, dense sexinous gran- ules present (Figs. 2, 27-30, 35, 36). This occurs in Old World species of subgenus Erythraster pollinated by passerine birds. Type (e). Pollen usually medium-sized with very discontinuous muri on equator or poles or on both, muri merging into sexinous granules. This occurs in five or more unrelated species 1985] HEMSLEY & FERGUSON—ERYTHRINA POLLEN MORPHOLOGY 585 Type i es. — 31. E. . E. droogmansiana show rather aati lumina. — 33. E. orophila. — 34. E. brucei. 35, 36. e (d) xim wide muri and very large sexinous granules. — 35. E. melanacantha. —36. E. variegata pete 533). that have distinctive but not necessarily exactly lumina that are sometimes sinuous, the muri dis- similar pollen. continuous on the equa torial area, low relief sex- Type (f. Pollen usually medium-sized with inous granules present small, round lumina on the polar area and wide This occurs only in E. merrilliana and E. tahi- 586 tensis (subgenus Erythraster) that occur in the Pacific OCCURRENCE OF POLLENKITT During the course of dissection of the flowers to remove pollen, it was noticed that pollen from the tubular, “hummingbird flowers" was consis- tently dust-like and easily spread, while pollen rom other, gaping flowers often had a tendency to be more sticky and difficult to separate. In order to see whether this might have any basis, whole anther preparations and groups of pollen grains of several species were examined under the electron microscope. Of the "hummingbird flower" anthers and pollen g in some other species such as E. humeana and E. lysistemon (visited by sunbirds), only very slight quantities of amorphous material or none at all could be seen between the pollen grains. e *pollenkitt." Variable quantities of pollenkitt were present, from little or none to quite large amounts, in groups with “passerine flowers.” Er- ythrina fusca, E. euodiphylla, E. velutina, and E. variegata were all found to have moderate to high quantities, while species such as E. mendesii, E. IT Occ n Ferguson (1984) illustrated these differences in the occurrence of pollenkitt in Erythrina and briefly discussed the phenomenon in some other papilionoid legumes. Changes occur in the composition and nature of the lipidic substances on the pollen wall prior to and at dehiscence of the anther (Hesse, 1981 and pers. comm.). These changes may occur to different degrees as a result of the methods used in the preparation of herbarium specimens. In view this the differences found in stickiness may no rue representation of the pollen under d conditions. Investigations of fresh material would be interesting. DISCUSSION The reticulate ornamentation of the exine of Erythrina pollen evidently shows considerable morphological variation between, and some- times within, species. How far may such varia- tion be explained in terms of an adaptive re- sponse to the pollen vector? In order to tackle such a question, a number of aspects of the pollination biology of the genus ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 must be considered. Especially in such a large and complex pantropical genus such as Erythri- na, our knowledge of the detailed pollination bi- ology of each species is necessarily incomplete. Nonetheless, a few generalizations may be made. All species of Erythrina are pollinated primarily by birds that do not appear to discriminate be- tween species (Raven, 1974); any one plant is likely to be visited, and pollinated, by a wide variety of local bird species (e.g., see Feinsinger et al., 1979; Guillarmod et al., 1979; Hernandez & Toledo, 1979; Morton, 1979; Raven, 1979; nation syndromes exist, the tubular “humming- bird flowers" in six adjacent sections of subgenus Erythrina (Stenotropis, Pseudoedules, Leptorhi- zae, Erythrina, Gibbosae, and Corallodendra) and the more or less gaping flowers, pollinated predominantly by passerines (Raven, 1974). In the Old World, and therefore outside the range of hummingbirds, pollination is effected by a va- riety of passerine species in diverse families (Ar- royo, 1981). A number of other features of floral biology further di h in “hummingbird flowers" e inflorescence axis is vertical, with the individual flowers directed outwards, horizontally or somewhat deflexed, providing easy access to the nectar for a long- billed, hovering bird (Fig. 37A, B). By contrast, a typical passerine-pollinated species has a hor- izontal inflorescence axis, and the gaping flowers are directed inwards (Fig. 37E, F), permitting a bird perched on the horizontal axis to reach the nectar (Cruden & Toledo, 1977). Quantitative and qualitative differences between the nectars of passerine- and hummingbird-pollinated flow- ers have also been found. The former tend to ndromes. Thus, fined generally as medium-sized, with medium- sized lumina in the reticulum and none to sparse low sexinous granules [Type (b), Figs. 13-18]. By contrast, the pollen of passerine sections, in both the Old and New Worlds, is more heterogeneous in character with respect to both size and or- namentation. However, it may be distinguished 1985] HEMSLEY & FERGUSON—ERYTHRINA POLLEN MORPHOLOGY 587 FiGURE 37. A, B. E. chiapasana, a typically hummingbird-pollinated species. — A. Showing a vertical inflo- rescence with flowers horizontal, pointing outwards, x25. — B. Tubular flower, x1. C, D. E. falcata, a passerine- bird-pollinated species with floral characters rather intermediate between those of the typical hummingbird and typical passerine bird type. — C. Showing a pendulous i ith flowe ess pointin X24. — D. Showing a somewhat shorter tubular flower, x 1. E, F. E. variegata, a typical passerine-bird-pollinated species. — E. Showing a horizontally orientated inflorescence with flowers pointing inwards, x25. —F. Showing a gaping flower with a broad reflexed standard, x1. 588 generally from the "hummingbird Mud either arkedly rin both small lumina size. S t groups as discussed later. The relative uniformity of pollen in the hum- mingbird-pollinated sections may indicate a rel- atively recent radiation of the group, and this is supported by Barneby (pers. comm.), who noted that these species are separated by much nar- rower morphological discontinuities than else- where in the genus. Conversely, the greater het- erogeneity observed in passerine-pollinated sections may reflect the greater spread of taxo- nomic relationships consequent on a longer pe- riod of Overall eyolukonary divergenpe, But per- underlies the observed differences between hummingbird and passerine pollen types, related to structural dif- ferences in the surfaces on which the pollen is carried. Neill (pers. comm.) indicated that the feathers of hummingbirds (and African sunbirds) may be finer at a microscopic level than those of larger passerine birds. Greenwalt (1960) stated that the hummingbirds may well be more closely feathered than any other bird family. The most striking feature of the plumage of hummingbirds is the iridescent patch of scaly *gorget" feathers found on the throat of the male and, in some species, extending over the head. These feathers have a quite different structure from the typical contour feathers of birds (Greenwalt, 1960). Fur- thermore, according to the observations of Neill (pers. comm.), the pollen of hummingbird-pol- linated species is generally placed on the bill or very fine throat feathers of the bird, whereas that of passerine-pollinated species is placed on the coarser chest feathers of the bird (see also Cruden & Toledo, 1977; Toledo, 1974; Toledo & Her- nandez, 1979). As previously indicated, some overlap of pol- len types does occur between the species of these two main pollination classes, but this is perhaps to be expected in a closely knit genus that is apparently still in a state of evolutionary flux with respect to pollination syndromes, and in which pollen morphology varies along a number of continua. For instance, pollen rather similar to the “hummingbird type" is found in a few Old World species. The southern African small sec- tion Humeanae in subgenus Erythrina is dis- tinctive in having deflexed, tubular rather than gaping corolla and is apparently adapted for pol- lination by sunbirds (Cruden & Toledo, 1977; Guillarmod et al., 1979). The African sunbirds, though quite unrelated, show certain parallels haps enama ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 with the New World hummingbirds in their de- gree of specialization as flower visitors (Stiles, 1981) and in the possession of patches of irides- cent plumage, for example. However, the pollen of E. zeyheri in this section was found to show some variability (Figs. 23, 24). The neighboring HE section | Caffrae, found i in ih me region, g fk Wer sunbirds are frequent visitors to pn flowers of E. caffra so are larger, less specialized passerines. Again, the pollen of E. /ysistemon in this section was somewhat variable. Two other Old World species, E. haerdii and E. livingstoniana in sub- genus Chirocalyx, have pollen not dissimilar to the “hummingbird type." Visits from both hummingbirds and passer- ines are reported for several of the New World species in subgenus Micropteryx with more or less gaping flowers, including E. poeppigiana and E. crista-galli (Toledo, 1974). In the latter, the flowers are outwardly directed by virtue of their almost resupinate position, and it has been sug- gested that the resupinate position may be viewed as an alternative strategy to a straightened keel, 4 th + 99 VU MIV llvVtdai, the low $ sucrose: hexose ratio of the nectar, how- ever, is consistent with perching bird pollination Baker & Baker, 1982). The racemes of E. do- minguezii and E. falcata are pendulous rather than horizontal, and the flowers less widely gap- ing than the typical passerine type, with the keel petal forming a short tube around the base of the stamens (Fig. 37C, D; see also Lucas & Theobald, 1982). Of these species, the pollen of E. poep- pigiana most closely approaches the *'*humming- bird type," although that of E. falcata is only a little more granular; E. crista-galli differs in the small size of the pollen and the narrow lumina. Erythrina cubensis, placed in a monotypic sec- tion in subgenus Erythrina, has more or less tu- bular flowers, and although pollination data were not available, there seems no reason why it should not be hummingbird pollinated. Among the hummingbird- pollinated sections, mm finities of E. leptorhiza (section Leptorhizae) with species of sections Breviflorae and Edules, and this is supported by the pollen morphology (Gra- ham & Tomb, 1977): pollen in these sections is characterized by small lumina size and distinctly triangular amb (also found in sections Suberosae, Arborescentes, and Oliviae). Section Breviflorae is unusual in its pollination biology, containing 1985] both the passerine- -pollinated E. breviflora and are pollinated by medium-sized hummingbirds (Hernandez & Toledo, 1982). The pollen of both species is in general rather similar and not of the "hummingbird type." Perhaps these sections represent a cluster of species around the **evo- lutionary borderline" between pollination syn- romes (Hernandez & Toledo, 1982). The other exceptions to the “hummingbird type" pollen are E. speciosa (section Stenotropis) and E. schimpf- fii (section Pseudoedules) with smaller than av- erage lumina size (Fig. 20), E. pallida (Fig. 21) and E. corallodendron showing moderate gran- ularity, and E. hondurensis with discontinuous m uri. A further feature of the pollen morphology was the sometimes quite considerable intraspecific variation. The most during this study was E. fusca (Figs. 1-3), al- though other species also showed some vari- ability. Erythrina fusca is the most widespread species in the genus, occurring on all four con- tinents, and was also considered by Krukoff and Barneby (1974) to be the most primitive. Per- haps this variability in pollen morphology is an indication of the greater plasticity of expression of a relatively unspecialized genotype. Or per- haps it is a function of the wide distribution and long evolutionary history, permitting local dif- ferentiation of pollen types. From the foregoing, it may be seen that al- though a broad correlation can be made between pollen morphology and pollen vector, this is sub- ject to several constraints. However, as previ- ously mentioned, many gaps still remain in our knowledge of the plant-pollinator relationships in the genus, and such relationships are not al- ways straightforward. A thorough comparative study of the structural differences between the field, outside the scope of this investigation. In this context, it should be noted that the actual surface of interaction between pollen and pollen vector may well not be the exine itself but an adhering layer of tapetal-borne materials (pol- lenkitt). Hesse (1981) has suggested that the exine sculpturing may influence the distribution and adhesion of pollenkitt on the surface of the exine, thereby modifying the stickiness of the pollen. During this study, several of the passerine-pol- linated species were found to have an abundance, or irregular and variable distribution of pollen- HEMSLEY & FERGUSON—ERYTHRINA POLLEN MORPHOLOGY 589 kitt over groups of pollen grains, whereas this was not apparent in hummingbird flowers. May- be the presence of sexinous granules perd p'a the adherence of pollenkitt by providing a * prominent granules or closely packed muri es- haps may support the layer of pollenkitt at the outermost surface of the pollen grain, where it is most effective. Q + +h sculpturing at an evolutionary, laal or de- velopmental level. How strong a selective pres- sure must operate to disrupt established patterns of exine ornamentation influenced more by evo- lutionary relationships than present pollination ecology? How long must the relationship be- tween plant and pollen vector have been estab- lished before any modification of the pollen sur- face becomes apparent? What is the effect of the possibly quite different surface structures? Can adaptation of pollen ornamentation occur at a populational level in response to locally oper- 1979), influence the form of the exine sculpture? How sensitive is the form of the reticulum to minor fluctuations in, for example, temperature or humidity (Hebda & Lott, 1973) during de- velopment? Any or all of such factors, or many others, may interact to determine the nature and extent of the variation observed on the pollen surface. CONCLUSIONS Although considerable variation occurs in the exine ornamentation of the pollen of Erythrina, some general conclusions can be drawn from this study. New World species of Erythrina with long, slender, miS presented, tubular flowers and vertica by ukasa snakes tend to have similar medium- sized pollen with a simple, regular, reticulate or- namentation and no sexinous granules in the lu- mina. Species with more or less gaping flowers held reflexed with the standard petal enlarged, with horizontal inflorescences that occur in both the Old and New World, and that are pollinated c v with almost always either small lumina or sex- inous granules present. The variation in the pol- len morphology of passerine-bird-pollinated species may reflect the large range of types of 590 birds involved as much as taxonomic and mor- phological differences. Pollen from hummingbird-pollinated species appears to be dry and powdery when dissected from anthers taken from herbarium specimens while that of many passerine-bird-pollinated species appears to be rather sticky. The observations suggest some modification of pollen morphology in response to pollen vec- tor LITERATURE CITED ARROYO, M. T. K. 1981. Breeding systems and pol- ee biology in Leguminosae. Pp. 723-769 in . M. Polhill & P. H. Raven (editors), Advances in Legume Systematics. Royal Botanic Gardens, BAKER, I. &H. G. BAKER. 1982. Some chemical con- stituents of floral nectars of Erythrina in relation to pollinators and systematics. Allertonia 3: 25- CRUDEN, R. W. & V. M. ToLEpo. 1977. Oriole pol- lination of Erythrina breviflora (Leguminosae): evidence for a polytypic view of ornithophily. PI. Syst. Evol. 126: 393-403. ERDTMAN, G. 1969. Handbook of Palynology. unksgaard, Copenhagen. FEINSINGER, P., Y. B. LiNHART, L. A. SWARM & J. A. OLFE. 1979. Aspects of the pollination biology of three Erythrina species on Trinidad and To- bago. Ann. Missouri Bot. Gard. 66: 451-471. FERGUSON, I. K. 1984. Pollen morphology and bio- systematics of the subfamily Papilionoideae (Le- guminosae). Pp. 377-394 in W. F. Grant (editor), Plant Biosystematics: Forty Years Later. Academ- ic Press, Canada. & SKVARLA. 1981. The pollen mor- phology of the subfamily died (Legu- minosae). Pp. 859-896 in R. M. Polhill & P. H. jer (editors), Advances in phu Systemat- . Royal Botanic Gardens, Kew . Pollen morphology in re- lation to pollinators in Papilionoideae (Legum nosae). Bot. J. S 3. GRAHAM, A. A. S. Tome. 1974. Palynology of Er- ythrina (Leguminosae: Papilionoideae): prelimi- nary survey of the subgenera. Lloydia 37: 465- 481. 1977. Palynology of Erythrina (Le- guminosae: Papilionoi ideae): the subgenera, sec- tions, and generic relationships. Lloydia 40: 413- 435 GREENWALT, C. H. 1960. Hummingbirds. Doubleday & Co. Inc., New Yor GUILLARMOD, A. J., R. A. Jus & C. J. SKEAD. 1979. Field studies of six sou cen African species of Erythrina. Ann. Missouri Bot. Gard. 66: 521-527. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 HEBDA, R. J. & J. N. A. Lorr. 1973. Effects of dif- ferent temperatures and humidities during growth on pollen morphology: an SEM study. Pollen & Spores 15: 563-571. HERNANDEZ, H. M. & V. M. TOLEDO. 1979. The role of nectar robbers and pollinators in the reproduc- tion of Erythrina leptorhiza. Ann. Missouri Bot. Mig 66: 512-520. — n 1982. Floral biology of Erythrina patolobium and the evolution of pollinatio etin n American species of the genus. Allertoni $ 71-84. T n J. systems. Ann. Mi HESSE, M. "1981. relation to the stickiness of angio ad Palaeobot. Palynol. 35: 81-92. 1979. Pollen walls as adaptive issouri Bot. Gard. 66: 813-829. The fine structure of exine in sperm pollen KnRuxorr, B. A. & R. C. B Conspectus of: species ofthe genus E rythrina. Lloydia 37: 332- 459. LARSON, D. A. 1964. Further electron microscopic eturiac nt 4 + + A ct s. 2i Palynol. 5: 265-276, pls. 1-8. Lucas, S. A. & W. L. THEOBALD. 1982. Observations of flowering behavior in selected species of Ery- thrina in cultivation in Hawaii. Allertonia 3: 85- E. S. 9. MORTON, 1979. Effective pollination of Ery- coevolved behavioral manipulation. Ann. Mis- souri Bot. Gard. 66: 482-489. RAvEN, P. H. 1974. Erythrina (Fabaceae): achieve- ments and opportunities. Lloydia 37: 321-331. 1979. Erythrina ipepe Faboideae): in- troduction to symposi II. Ann. Missouri Bot. Gard. 66: 417- Romeo, J. T. 1973. A Chemotaxonomic Study of the Genus Erythrina oe osae). Ph.D. Thesis, Univ. of Texas, Aus STEINER, K. E. 1979. Passerine pollination of Ery- thrina megistophylla Diels oen Ann. Mis- souri Bot. Gar -5 SriLEs, F. G. 1981. Geographical aspects of bird- flower coevolution, with particular reference to Central America. Ann. Missouri Bot. Gard. 6 323-351. THANIKAIMONI, G. 1976. Index i sur la morphologie des pollens d'angios up- plement-2. Trav. Sect. Sci. Tech. Inst. gone Pon- dichéry 13: aor atriéme index E sur an rme ct. Sci h. Inst. Franc. Pondichéry 17 71-336. OLEDO, V. 974. Observations on the relation- ships between hummingbirds and Erythrina species. Lloydia 37: 482-487. & HERNANDEZ. 1979. Erythrina oli- viae: a new case of oriole oe " Mexico. Ann. Missouri Bot. Gard. 66: 503-51 Volume 72, No. 2, pp. 167-450 of the ANNALS OF 19 July 1985. THE MISSOURI BOTANICAL GARDEN, was published on MONOGRAPHS in Systematic Botany from the Missouri Botanical Garden MSB-1 MSB-2 MSB-3 MSB-4 MSB-5 MSB-6 MSB-7 MSB-8 MSB-9 MSB-10 MSB-11 MSB-12 A Provisional Checklist of Species for Flora North America. Revised. 1978. Out-of- Print. Missouri Wildflowers of the St. Louis Area. E. R. Eisendrath. 390 pp. Illustrated. 1978. $7.95. A Dictionary of Mosses. Third Printing. 1981. Out-of-Print. Orchids of Panama. L. O. Williams & P. H. Allen. viii + 484 + xxvi pp. Illus- trated. 1980. $20.00. Index to Plant Chromosome Numbers, 1975-1978. P. Goldblatt, Editor. vii + 552 pp. 1981. $15.00. Les Ombelliferes. A.-M. Cauwet-Marc & J. 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Mail form 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 the MONOGRAPHS checked above to: O Payment enclosed. Name O Send invoice ($1.00 fee will be Address added to total). Payer T Pe ee nee Tib. eS, ey a POE P Clauntryv ANNALS JF THE HISSOUR] BOTANICAL GARDEN OLUME 72 1985 NUMBER 4 Lagenostoma and Tenlangium CONTENTS The Implications of Phylogenetic Analysis for Comparative Biology: The Thirtieth Annual Systematics sedis Vicki A. Funk & Joel Ch ae — —— ——— — o 2 7 LU 5 591 Species and Speciation in Phylogenctic Ë Sy stematics, with b Examples om the North American Fish Fauna E. O. Wiley & Richard L. Mayden ... 596 Geological Hierarchies and Biogeographic Congruence in the Caribbean Donn E. Rosen ... s 636 Historical Ecology: A New Aupmach b to Studying t the Evolution of Ecological Associations Daniel R. Brooks . — BELASH 660 Phylogenetic Patterns and Hybridization A. jus = 681 Phylogenetic Analysis of Seed Plants and the s: of Angiosperms Peter KC 95 7 o om š ecu) RED Red Diversification and Its Causes dod Ca. ee A lological Basis for Adaptation in Grasses: An Introduction Khidir W. h. Hilu & Thomas R Soderstrom 1 — 823 olyploidy, Hybridization, and the Invasion of New Habitats G Led: MSN. — —— —— — 824 Adaptation of G1 to Water Stress— Leaf Rolling a and j Siomate 1 Distri- bution R. E. Redmann . SSE SLOTS > E 833 Contents continued on back cover VOLUME 72 WINTER 1985 NUMBER 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN The ANNALS, published quarterly, contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden, t. Louis. Papers originating outside the Garden will also be ac- cepted. 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 first issue of this volume. EDITORIAL COMMITTEE Nancy Morin, Edit ties Botanical Garden CHERYL R. BAuER, Editorial Assistant 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 MES phos Garden For subscription information contact the Business Office of the Annals, P.O. Box 299, St. Louis, MO 63166. Subscription price is $65 per volume U.S., $70 Canada, and Mexic NES $75 all other countries. Personal subscriptions are available at $30 and $35, respec Airmail delivery charge, $30 per volume. Four issues per volume. The ANNALS OF THE MISSOURI BOTANICAL GARDEN (ISSN 0026-6493) is je lished quarterly by the Missouri Botanical Garden, 2345 Tower Grove Ave., St. Louis, der 63110. Subscription price is $65 per volume U.S., $70 Canada and Mexico, $75 all o countries. Personal subscriptions are available at $30 and $35, respectively. ci postage paid at St. Louis, MO and additional mailing offices. POSTMASTER: FC x 299, changes to the ANNALS OF THE MISSOURI BOTANICAL GARDEN, P.O St. Louis, MO 63166. € Missouri Botanical Garden 1985 — a NE OF THE ANNALS MISSOURI BOTANICAL GARDEN VOLUME 72 1985 NUMBER 4 THE IMPLICATIONS OF PHYLOGENETIC ANALYSIS FOR COMPARATIVE BIOLOGY: THE THIRTIETH ANNUAL SYSTEMATICS SYMPOSIUM This symposium brings together seven sys- tematic biologists to discuss the relationship be- tween phylogenetic systematics (cladistics) and various branches of comparative biology. Cla- distic theory and method were not the focus of this symposium, but rather it addressed how the results of cladistics—that is, hypotheses about phylogenetic pattern—are an essential compo- nent of historical analysis. Thus, the papers pre- sented here illustrate six specific problems that, directly or indirectly, rely on cladistic analysis for their solution Cladistics icine an important force within systematic biology following the English trans- lation of Willi Hennig's ““Grundzuge einer Theo- rie der phylogenetischen Systematik" (1950), which was updated in 1966 as “Phylogenetic Sys- tematics.” From that time, cladistics has steadily increased in influence, first within zoology, and somewhat later in botany (for an introduction to cladistics, see Eldredge & Cracraft, 1980; Wiley, 1980, 1981; Patterson, 1980; Nelson & Platnick, 1981; Cracraft, 1983; Humphries & Funk, 1984). During this period many workers have discussed the pros and cons of cladistics, and frequent mis- understandings about the theoretical and meth- dological content of cladistics have arisen. Per- haps contributing to this situation is the fact that the views of cladistic theoreticians have also evolved, often along divergent pathways so that the field of cladistics is now broader, and con- sequently more internally contentious, than it was 15 years ago. Despite this diversity, the ques- tion can be asked whether cladistics is united by any underlying principles so as to form a coher- ANN. Missouni Bor. GARD. 72: 591—595. 1985. ent research program. An answer to this question is of interest beyond being able to differentiate cladists or their work, because it also helps to place the various transformations of cladistics into a common framework and to gain a per- spective on some of the arguments that are cur- rently being expressed over the meaning and con- tent of cladistic thought. THE PRINCIPLES OF CLADISTICS We do not wish to engage in an extended dis- cussion of the historical changes that have taken place in the thinking of individual cladists. We want, instead, to call attention to a common in- tellectual thread that has passed through the writ- ings of cladists from Hennig to the present. We perceive this thread to consist of two principles: (1) taxa are united into natural groups on the basis of shared derived characters, or synapo- morphies (the Principle of Synapomorphy), and (2) classifications must express those taxic pat- terns of synapomorphy explicitly (the Principle of Strict Monophyly). To be sure, each of these principles may rely on certain unexpressed as- sumptions, but none we think that are not also shared with For instance, Platnick's (1979: 538) rud princi- ple of cladistics, that nature has a single historica pattern with a hierarchical structure, would ue be acceptable to the majority of sys- temati dn oe to these two principles, the Prin- ciple of Synapomorphy is clearly primary in that the Principle of Strict Monophyly depends on it. vstematics 592 Pheneticists and their followers reject the first principle, and therefore, logically also reject the Principle of Strict Monophyly. Some evolution- ary (or eclectic) systematists might well accept the Principle of Synapomorphy but advocate paraphyletic taxa, thereby rejecting the Principle of Strict Monophyly; in fact, however, we believe that most evolutionary systematists, like phe- neticists, reject both principles. We also suggest that a third principle has been central to cladistic theory and methodology, namely, the Principle of Strict Parsimony. AI- though an acceptance of parsimony is basic to all scientific reasoning, cladists have certainly been much more concerned with elucidating the relationship of parsimony to systematics and ap- plying parsimony analysis to their work than have advocates of phenetics or evolutionary system- atics. Some of the critical literature discussing the application of parsimony to systematic hy- potheses includes Kluge and Farris (1969); Gaff- ney (1979); Farris (1982, 1983), Sober (1983a, 1983b), and Maddison et al. (1984) Within a cladistic framework taxa are grouped on the basis of shared character transformations. All such characters are termed synapomorphies. Some of these transformations will be unique, others may have arisen more than once within the group being studied and may be termed ho- moplasious. Obtaining a final interpretation of the pattern of the character transformation for any specific character depends on the tree to- pology (phylogenetic hypothesis) upon which all character transformations are optimized. Thus, we choose that tree by maximizing the congru- ence of character transformations across the en- fewer! ad hoc hypotheses jore to explain ley, 1975; Farris 1982, 1983). Within this iss, 1cal frame- work, it is clear that homoplasies (parallelisms, convergences, reversals) are also synapomor- phies: they represent derived character transfor- mations defining two or more unrelated groups of taxa (“unique” synapomorphies define only a single group). The Principle of Strict Parsimony has been a core methodological component of mainstream cladistics for nearly two decades. In recent years, a small number of workers, also claiming to prac- tice cladistics, have advocated abandoning (or at least relaxing) this principle in favor of a method that constructs trees using only a subset of the ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 original character data. This method, called com- patibility or clique analysis, unites taxa by uti- lizing those characters that exhibit congruence and eliminating those showing homoplasy (Es- tabrook et al., 1976; Meacham, 1984). Compat- ibility methods have been strongly criticized from several standpoints (Farris & Kluge, 1979; Mick- evich & Parenti, 1980; Churchill et al., 1984). We note here the primary philosophical objec- tion: a given scientific hypothesis, including those within systematics, has veracity compared to competing hypotheses when it best explains all of the relevant data, thus it is unclear to what extent hypotheses can be objectively compared when we exclude from consideration data that might be incongruent with one or more of the alternatives. THE CHANGING FACE OF CLADISTICS Cladistics emerged as a solution to a long- standing problem within systematic biology: how o we come to have knowledge about the phy- logenetic relationships of organisms? Most con- temporary systematists employing cladistic methods still see this as the central focus of their research. For all these workers, branching dia- grams (cladograms) are interpretable a history. Within this framework, synapomorphies are taken to be evolutionary transformations or derivations from more primitive conditions. The preceding describes a view of cladistics sometimes termed "phylogenetic" or “evolu- tionary" cladistics. To most of its practitioners, an evolutionary process underlies the rationale for the method. A claim for a specific evolution- ary process is generally not made, but without the assumption of evolution, we are sometimes told, cladistics is set adrift in a sea of “conceptual confusion" (Beatty, 1982: 33). Thus, according to Wiley (1981: 22), “the formalism of taxonomy must be subservient to the demands of evolu- ion." An alternative viewpoint about the relation- ship between cladistic analysis and evolutionary theory has surfaced within the last five years. Unfortunately, it has generated substantial mis- understanding, particularly outside the cladistic e neously, a number of workers proposed that there could be a separation between reconstructing his- torical pattern on the one hand and assumptions 1985] about the process underlying that pattern on the other. These systematists suggested that the anal- ysis of systematic pattern was (or certainly could be) independent of any preconceived notion of process (Gaffney, 1979; Eldredge, 1979; Plat- nick, 1979; Eldredge & Cracraft, 1980; Patterson, 1980, 1982a; Nelson & Platnick, 1981). The reaction to transformed or pattern cladis- tics, as the above view has been called, has run from quiet acceptance "à buys en hostility. Most (Beatty, 1982; Ridley, 1983) and has been found- ed more on its misunderstandings of cladistics than on any cogent criticism of pattern cladism see responses of Platnick, 1982; Patterson, 1982b; Brady, 1982). Nevertheless, some potentially in- teresting issues are being raised by cladists them- selves. For example, if a specific notion of evo- lutionary process is unnecessary, where does that leave our interpretation of the concept of syn- apomorphy? Phylogenetic cladists continue to view character transformation in terms of **prim- itive to derived sequences" and to argue that in constructing pleas of tas transformation must have some prior cor ofhistorical relationships in ae ph assasi outgroup analysis. Pattern cladists might respond that syn- apomorphy is strictly a problem of deciding the level of generality of defining characters and that this can be arrived at by inspection and com- parison of y histories, thus eliminating the need for prior (see Nelson & Platnick. 1981: Rosen, 1984). The is- sue of which method of comparison (outgroup or ontogenetic) à js primary, is this is being debated at this time (Kluge, Brooks & Wiley, 1985; Nelson, 1985; Platnick, 1985). Matters of principle aside, in practice cladists from either side of the debate will use both ontogenetic and outgroup data to resolve systematic relationships, so perhaps in that re- gard the debate will not concern those system- atists interested solely in cladistics as a meth- odological tool. lthough cladists might differ in their percep- tions about the role of evolutionary assumptions within cladistics, as we noted earlier a shared set of common principles can be identified. To our knowledge, all pattern cladists believe in natu- ralism, whereby the hierarchical pattern of na- ture is assumed to be the result of naturalistic processes. Thus, to claim that pattern cladists do not believe in some form of “evolutionary” (his- FUNK & CRACRAFT — INTRODUCTION 593 torical) process behind phylogenetic pattern or that pattern cladistics is **antievolutionary" is simply mistaken. Pattern cladists merely claim that a prior commitment to a specific process is unnecessary in order to generate hypotheses about that pattern. And given that hypotheses about pattern have been proposed for hundreds of years, under different theoretical paradigms about the causes of that pattern (Patterson, 1977; Nelson & Platnick, 1981), their point seems to be well taken. THE SYMPOSIUM Speciation analysis. In the first paper, E. O. Wiley and R. L. Mayden show how the results of a cladistic study can be used to examine pat- terns and processes of speciation. Using species- level taxa within the eastern North American fish fauna, they begin by briefly discussing attitudes towards species concepts as they have been used by phylogenetic systematists. Wiley and Mayden then investigate patterns of relationships for nu- merous clades of fishes having common species borders of endemism. They show that these his- torical hypotheses inhibit intercladal congru- ence, which they then use as components of an analysis of speciation modes. Historical biogeography. Donn Rosen begins his paper by addressing two widely held, but in- correct, assumptions: that fossils can tell us how old a taxon is, and that the ages of geologic events have been correctly assigned. He emphasizes the need for precision in specifying how historical biology i IS telated to historical geology and shows OW between these two systems. He seeks to discover patterns of congruence between historical biogeography and geological events so explicit that the congruence discovered cannot be dismissed as being due to chance or coincidence. Rosen stresses that it is the “independence of biological from geological data that makes the comparison of the two so interesting... ." He reviews Caribbean geologic history and presents a cladistic hypothesis for the historical interrelationships of the areas of that region. He points out that complex histories should lead us to expect complex patterns and that all potential hypotheses of area relationships may be corroborated by one or more cladistic patterns exhibited by the endemic taxa. Rosen e raises a Wasp for biogeographic analysis: the pas me workers have assumed that dispersal iS ad by failure to discover - B zy 594 congruence in area relationships, yet even though dispersal may be widespread “theories of dis- persal to explain biotic complexity are no more informative than theories of relationship based on symplesiomorphy Historical ecology ud coevolution. Daniel Brooks discusses analytic methods of historical ecology and shows how they provide a missing component in studies analyzing the evolution of ecological associations. Direct estimates of eco- logical history are obtained by constructing cla- distic hypotheses for as many interacting groups of organisms as possible. This method is con- trasted with evolutionary ecology, which often uses indirect estimates such as the assumption that the age of ecological associations is propor- tional to its diversity. Using host-parasite data, Brooks addresses three questions within histor- ical ecology: (1) How did species occurring in a given area come to be assembled? (2) How did two or more species having a close and evident ecological relationship come to be that way? And (3) Under what conditions did the ecological life history traits that we observe today emerge? Brooks shows how historical analysis through cladistics can . insights into all of these ecological ques S. I Vicki Funk addresses one of the major concerns of plant systematists, namely how to analyze hybridization within the context of a phylogenetic hypothesis. She summarizes the problems that arise during a cladistic analysis of a group whose taxa hybridize with one another, and she presents dmi for NAE. hy- brids into a cladogra rder to exemplify these methods, she dapi cladistic ee of seven different genera exhibiting various de- grees of hybridization among their component taxa. Funk stresses that all hypotheses regarding hybrid identification must be corroborated by chromosomal, distributional, and ecological data. A major conclusion is that cladistic analysis is indispensable when analyzing hybridization, but in cases of taxa exhibiting extensive hybridiza- tion, all systematic methods, including cladistics, may fail to give a clear indication of the history of the group. Origin of the angiosperms. In the next paper, Peter Crane applies cladistic on to the question of the origin of the iosperms. His central E is fo ) delineate then major groups of seed plan and dun to establish to panic group of gym- nosperms the flowering plants are most closely ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 related. Crane accomplishes this task by provid- ing an exhaustive analysis of extant and fossil groups of seed plants and establishes a frame- work within which competing theories of rela- tionships are compared. By formulating the rel- evant phylogenetic questions more explicitly, he also provides an alternative to the established tradition of searching for ancestral groups. Biological diversification theory. In the final paper, Joel Cracraft briefly describes the patterns of diversification that fall within the umbrella of a general theory of diversification. He proposes a hypothesis in which speciation rates are a func- tion of the rate of change in large-scale geomor- phological complexity, whereas extinction rates are a function of temporal and spatial changes in environmental harshness. Together, these two rate-controls describe a m process of diversification. He then summarizes some of the evidence supporting i B Wawap pointing out that the data themselves are often dependent upon knowledge of the phylogenetic relationships of many groups of organisms. LITERATURE CITED BEATTY, x 1982. Classes and cladists. Syst. Zool. 31: 25-3 BRADY, 4 H. 1982. Theoretical issues and pattern cladistics. Syst. Zool. 31: 286-291. Brooks, D. R. & E. O. Wirev. 1985. Theories and methods in different approaches to phylogenetic systematics. puces 1: 1-11. CHURCHILL, S. P., E. O. WILEY & L. A. HAUSER. 1984. A critique of Wagner groundplan-divergence stud- ies and a comparison with other methods of phy- logenetic analysis. Taxon 33: 212-232. CRACRAFT, J. Cladistic ei and vicariance biogeography. Amer. Sci. 7 81. ELDREDGE, N. 1979. Cladism u a sense. Pp. 165-197 in J. Cracraft & N. Eldredge (editors), Phylogenetic Analysis and Paleontology. Colum- bia Univ. Press, New Yor J. CRACRAFT. 1980. Phylogenetic Patterns and the Evolutionary Process. Columbia Univ. Press, New York. ESTABROOK, G. F., C. S. Jou Mo A mathematical foundation js the analysis of veri character compatibil- y. Math. Biosci. 29: -187. Nd J.S. 1 i eeu and parsimony. Syst. Zool. 31: s id 83. e logical basis of phylogenetic anal- ysis. Adv. Cladistics ar —36. & A. G. KLU NSON, JR. & F < O 79. A botanical clique. Syst. Zool. 28: 400-4 GAFFNEY, E. S. 1979. An introduction to kis logic of phylogeny reconstruction. Pp. 79- n J. Cra- craft & N. Eldredge (editors), Phviosenctc Anal- ysis ~ Paleontology. Columbia Univ. Press, New Yor 1985] HENNIG, W. 1950. Grundzuge einer Theorie der phy- um Systematik. Deutscher Zentralver- bad Ber 1966. pe Systematics. Univ. Illi- nois Press, Urba HUMPHRIES, .& "V. FuNK. 1984. . me thodology. Pp. 323-362 in V. H. He D. ore oe mn Concepts in | Plant Taxonomy. Academic Pres eii d phylogenetic sys- tematics. e 1: 13-27. . FARRIS. 1969. Quantitative phyletics and the evolution of anurans. Syst. Zool. 18: 1- 34. MADDISON, W. P., M. J. DONOGHUE & D. R. MAD- DISON. 1984. Outgroup analysis and — dud Syst. Zool. 33: 83-103. MEACHAM, C. 1984. The role of hypothesized direc- history. Taxon 33: 26-38 F. & L. PARENTI. 1980. Review of ch ee analysis” by J. Strauch. Syst. Zool. 29: 8-113. 1985. Outgroups and ontogeny. Cla- distics 1: 29-45. N. I. PLATNICK. 1981. Systematics and Bio- geography: Cladistics and Vicariance. Columbia Univ. Press, New York. pu. C. 1977. The contribution of paleontol- ogy to teleostean phylogeny. Pp. 579-643 in M. K. Hecht, P. C. Goody & B. M. Hecht (editors), Major cialis in Vertebrate Evolution. Plenum, New Yor 1980. ` Cladistics. Biologist 27: 234—240. FUNK & CRACRAFT — INTRODUCTION 595 1982a. Morphological d and ho- mology. Pp. 21-74 in K. A. Joy: . E. Friday (editors), Problems of PLE Reconstruc- tion. Academic Press, London 1982b. Classes and cladists or individuals and evolution. Syst. Zool. 31: 86. PLATNICK, N. I. 1979. Eara in and the transfor- mation of cladistics. Syst. Zool. 28: 537- 1 Defining characters and evolutionary x groups. Syst. Zool. 31: 282-284. 1985. Philosophy and the transformation of 7 eadistics revisited. Cladistics 1: 87-94. 1983. Can classification do without evo- 47 l. n D. E. 4. Hi erarchies and history. Pp. 77- in J. W. Pollard iig eria Sagat Theory into the Future. John Wiley, New York. Soper, E. 1983a. ME in ini dba pom sophical issues. Annual Rev. Ecol. Syst. 14: 335- 357 3b. Parsimony methods in systematics. Adv. Cladistics 2: 37-47. WiLEY, E. O. . Karl R. Popper, systematics, and classification: a reply to Walter Bock and other iii taxonomists. Syst. Zool. 24: 233-243. 1980. Phylogenetic systematics and vicari- ance biogeography. Syst. Bot. 5: 194-220. — PN 1981. Phylogenetics: The Theory and Prac- tice of Phylogenetic Systematics. John Wiley, New York. — Vicki A. Funk, Department of Botany, Smith- sonian Institution, Washington, D.C. 20560; and Joel Cracraft, Department of Anatomy, Univer- sity of Illinois, Chicago, Illinois 60680. SPECIES AND SPECIATION IN PHYLOGENETIC SYSTEMATICS, WITH EXAMPLES FROM THE NORTH AMERICAN FISH FAUNA! E. O. WILEY AND RICHARD L. MAYDEN? Phylogenetic systematics as developed by Hennig (1950, 1966) is a system of methods and a view of the world designed to integrate system- atics with the evolutionary "paradigm." By evo- lutionary paradigm" we mean the world view that organisms always have parents and that or- ganismic diversity is a byproduct of descent with modification. If our view of phylogenetic system- atics is correct, or even partly so, the results de- rived from Hennig's methods should be of gen- eral utility to the evolutionary community. We shall attempt to show this utility by considering how the results of phylogenetic analysis can be used to infer the mode of speciation involved in the origin of species. Before doing so, however, we need to discuss what constitutes a species because our concepts will influence our choice of units to be analyzed. SPECIES IN PHYLOGENETIC SYSTEMATICS There is no consensus concerning either the nature of species-as-taxa or the importance of species among those who align themselves with Hennig's (1966) basic philosophy. This should come as no surprise because no consensus on either question has been reached by the rest of the biological community. At the risk of over- simplifying, we will define two basic attitudes. The first is what might be called the heuristic attitude. Rosen (1979: 277) defined species as “ʻa : 11) stated "species are simply the smallest detected samples of self- erpetuating organisms that have a unique set of characters." Rosen's statement is somewhat more restrictive, both in its inclusion of a geographic criterion and in its criterion that the species must have at least one apomorphous character. Both, of course, are not “process free" concepts since one (Rosen's) contains a statement based on character evolution while the other (Nelson and Platnick's) contains a statement based on repro- duction. Nevertheless, both concepts are as heuristic as the authors can make them he second attitude might be termed ibé **pro- cess" attitude. The concept applied is not re- stricted to purely heuristic considerations but also the evolutionary nature of the entities. Of all phylogeneticists, Hennig (1966) himself was the most radical proponent of this attitude, a fact that is not always appreciated. Hennig (1966) spent a substantial part of his book arguing against heuristic concepts because he rejected ideal mor- phology and the “morphological system." To Hennig (1966: 79-80), it was phylogeny that was important, morphology being a way of getting at the phylogeny: The categories [= taxa] of phylogenetic sys- tematics are not constructed by abstraction. They are not defined as bearers of a complex of characters that remains if, starting with the individuals, we subtract more and more char- acters that are specific to the individuals and then to progressively more inclusive groups of individuals. In the phylogenetic system the categories [= taxa] at all levels are determined by genetic [= genealogical] relations that exist among their subcategories [= taxa included within them]. Knowledge of these relations is a prerequisite for constructing these categories — taxa], but the relations exist whether they are recognized or not. Consequently here the morphological characters have a completely different significance than in the logical and morphological systems. They are not them- selves ingredients of the definition of the higher categories [= taxa] but aids used to apprehend the genetic [= genealogical] criteria that lie be- hind them. [Brackets and emphases ours.] ! This study was funded by a grant from the National Science Foundation (DEB-8103532) and two grants from ad University of Kansas General Research Fund (Nos. 3338 and 3591). um of Natural History and Department of Systematics and Ecology, The University of Kansas, Law- 045. rence, Visa 66 ANN. Missouni Bor. GARD. 72: 596-635. 1985. 1985] Thus Hennig (1) rejected taxa as being defined by characters a 2) accepted ibat the only char- were those that indicated genealogical connections. Hennig wished to work with entities having a “real” ex- istence in nature. Of course one consequence of holding to this realist view of the world con- cerned species — to Hennig species are important only to the extent that they function in the evolutionary process. Thus a concept of a species as a unit functioning in the evolutionary process was an important tenet in his system. For all practical purposes, Hennig (1966) adopted the concept that species are (or were at some stage) reproductive units, but differed from such work- ers as Mayr (1963) and Simpson (1961) in re- jecting chronospecies. However, he did more than this. Hennig understood very clearly that species have many of the characteristics of individuals, especially singular places of origin in time and space (1966: 81) If we now attempt to evaluate the categories [= taxa] of the phylogenetic system from the viewpoint thus gained, there can be no doubt that all the supra-individual categories [= taxa], from the species to the highest category rank, have individuality and reality. They are all (. ..) segments of the temporal stream of suc- cessive “interbreeding populations." As such they have a beginning and an end in time (N. Hartmann), and there is a constant causal con- nection between the phases in which they are found at different times (Ziehen). All this is missing in the categories [= taxa] of the mor- phological or typological system, which con- sequently are timeless adai (Woodger) and therefore have neither individuality nor reality. It would be a mistake to assume that his con- demnation was directed only at paraphyletic and polyphyletic taxa. Hennig was perfectly aware that species and natural higher taxa (i.e., monophyletic taxa) were not the same sorts of individuals as individual organisms. Michael Ghiselin isn’t the same sort d apiens, who isn’t the r, he recognized that the concept of individuality provided a link in our understanding of the tran- sition between species, higher taxa, and natural hierarchies (see Hennig, 1966: 82-83). Specifi- cally WILEY & MAYDEN— PHYLOGENETIC SYSTEMATICS 597 Thus all categories [= taxa] of the phylogenetic system are characterized by individuality an reality, in contrast to the abstract and timeless categories 2 taxa] of the morphological i s not mean that they are all “ and which we ordinarily call individuals. De- pending on the purpose we have in mind, we may emphasize the common traits in the mode of existence of the individuals (in the custom- ary, strict sense) and the supra-individual sys- all the criteria of individuality that character- ize one category rank [= taxa at one rank] to Mrs: category ranks [= taxa at higher or lower nk] if we are to avoid faulty conclusions P xu 1966: 83). Hennig went on to that his thoughts, apparently developed from considering Hart- mann (1942, not seen, cited in Hennig, 1966), applied m to taxa that were natural in the phy- logenetic sen Michael Chisin (1969, 1974, 1980, 1981) in- dependently came to the same conclusion as Hennig (1966): a species are individuals and most emphatically, they are not mere clus- ters, nor are they natural kinds. This distinction is discussed at length by Ghiselin (1969, 1974, 1980, 1981) and by Hull (1976, 1983). Natural kinds are held by many philosophers to be eter- nal, immutable, and discrete. The problem, pointed out by both Ghiselin and Hull, is that philosophers have traditionally thought of par- ticular species as natural kinds, even after ac- cepting that evolution renders particular species and changeable entities. We stress an additional confusing aspect to the problem — “species” refers to both particular species such as Homo sapiens and Pinus ponderosa and to a natural kind that defines a class of entities on which process operates: although Pinus ponder- osa is changeable and temporary, the natural kinds “biological species" and “evolutionary species” are eternal and immutable. There may be no individual species that are the natural kind “biological species," but that doesn’t mean there has never been such a species or that such a species will never appear in the future. The pos- sibility of biological or evolutionary species is a permanent possibility, entities which fill the class 598 may evolve many times and in many places. An individual species, such as Pinus ponderosa, owever, can evolve but once and once extinct is extinct forever. PROCESS AND PATTERN IN SPECIATION Speciation is one of the major processes un- derlying the patterns of descent that is the phy- logeny of life on earth. As summarized by Wiley (1981) from many sources, speciation is not a single process but several more-or-less distinct processes that result in increases (and theoretical decreases) in the number of species in a clade. Patterns of genealogy at the level of species are the result of speciation. In some cases patterns e and space. In other cases they are different. purus patterns may be the result of mistakes in reconstructing phylogenetic his- tories, the result of undetected extinctions or spe- ciations, or the result of dispersals that alter the original pattern. These are all the result of our inabilities to detect what has actually happened. It is also possible that we have correctly recon- structed history and that the similarities and dif- ferences we observe in patterns of descent have a causal reason. Specifically, different modes of speciation might result in different patterns of genealogy in time and space. If so, and if we can couple differences of pattern with differences in mode, then we have additional tools with which we can gain a better understanding of the evo- lutionary process. ASSUMPTIONS Following the analysis of Wiley (1981) we shall explore the phylogenetic and biogeographic pre- dictions derived from various proposed specia- determine which of the various modes might be operating. Obviously we do not expect all, or even most, groups to fulfill these conditions. At best we can hope that a few groups will fulfill the conditions and act as exemplars for studying the evolutionary process. Assumption 1. Character evolution provides a reliable basis for inferring the history of spe- ciation. There are two conditions under which this assumption would hold: (1) character evo- lution is coupled with speciation; or (2) character evolution keeps pace with lineage splitting, that ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 is, there is enough anagenesis between lineage splits so that the sequence of splits leaves a “‘his- torical trail" that can be followed. Assumption 1 is violated if vicariance proceeds at a faster rate than character evolution, producing isolated populations that are similar. Subsequent evolu- tion would not document the history of vicari- ance and a polytomy would result. (We shall see in a later section that a polytomy may exist even if character evolution does keep pace with vi- cariance.) Two examples illustrate this. Fundulus sciadi- cus 1s composed of at least three disjunct pop- ulations (Fig. 1), presumably separated during the Kansan glaciation. If future divergent char- acter change occurs before sympatry is achieved or before further splitting, a future phylogenetic reconstruction will not reflect the vicariance event involved. Fundulus catenatus is composed of at wo of nessee highlands vicariance event that we will discuss in a later section. Subsequent character evolution would not document the sequence of geographic isolation that this species has under- n one. Assumption 2. There are no extinctions of species in the clade. If we examine the results of particular speciation events, we must have some confidence that the species compared are the product of: a single speciation event and D sep- arated on the and unknown, species. If iie are, then the dif- ferences observed may be the result of differences Desert (Uyeno & Miller, 1963). Ob- viously ascribing the differences between Recent western Fundulus and their eastern counterparts to a single speciation event would be a dubious hypothesis. We simply don't know how many events were involved. The case of the Fundulus nottii species group (Wiley, 1977) is different. In this. case (see Fig. 5 and discussion in a later patterns because the group is distributed contin- uously over space. ud od 3. _ Dispersal has not obscured the or clades that show large- scale dispersal may be identified WILEY & MAYDEN 599 d ue T m. e PESCA ON "s ge s C aa t S S. | i I i FIGURE 1. by widespread sympatry. For example, the pat- tern of speciation in the Fundulus olivaceus group (Fig. 3) is obscured by the widespread sympatry of F. olivaceus and F. notatus (Thomerson, 1966) and by the sympatry of F. euryzonus with both F. notatus and F. olivaceus (Suttkus & Cashner, 1981 iously we cannot insure that any clade ful- fills all three assumptions. However, we will sug- gest that some clades do because their histories and distributions show concordance with pre- dictions based on particular modes of speciation and, in some cases, partial congruence in time and space with other groups. One final caveat— the modes we discuss are not our inventions and we have no interest in defending any of them as dominant or, in some cases, even realistic. MODES AND PREDICTIONS Following Mayr (1963), we may distinguish among three basic models of speciation. Reduc- Distribution of Fundulus sciadicus (from Lee et al., NS on — 3 À \ kn i i = $ wo i i. SN 7 r X » p XA ro NS, < 2 (ox f N Y | E a TA b US vli \ Soo i| ex up 2S7 : 7 i / 7# i. . n js 1 SA Ç j CN TA y h i pao Ne nA v AS 1980 and Ellis, 1914). tive speciation involves lineage fusion; two species fuse to produce a third new species. While theoretically possible, there have been no doc- umented cases of such events (Wiley, 1981). The nearest example would be a “‘compilospecies” (Harlan & deWet, 1963), in which one preexist- ing species simply absorbs the other. Additive speciation involves lineage splitting and reticu- late speciation. The result is an increase in species diversity. There are several models for nonretic- ulate additive speciation that can » distin- guished on the basis of the importance of geo- graphic subdivision from essential (allo b ic speciation) to nonessential (sympatric specia- tion). Additive speciation models are rich in pre- dictions and we shall discuss several of them in sections below. The final model is “‘phyletic spe- ciation," the proposition that there is species turnover within a single lineage without adding species in any one time frame. This “mode” has been rejected as a true mode of speciation by Hennig (1966) and by Wiley (1981) who pointed 600 ANNALS OF THE MISSOURI BOTANICAL GARDEN EM # <— 6 r [VoL. 72 FL e NACE p “= I \ s FIGURE 2. Distribution of Fundulus catenatus (from Lee et al., 1980). out that subdividing a continuous lineage is an inherently arbitrary procedure. We shall not dis- cuss it further. ALLOPATRIC SPECIATION Allopatric speciation is an umbrella term for a spectrum of modes that involve complete geo- graphic separation of two or more subpopula- tions of a species during its evolution into two or more daughter species (one of which may be the ancestor itself). Allopatric speciation is thought to the dominant mode by many in- vestigators (cf. Mayr, 1963; Grant, 1971). Two basic models can be discerned based on popu- lation structure, a UER that gums gene flow ates hn and nne that assumes gene flow is absent or rare. The "gene flow model” can be further subdivided on the basis of the relative percent of the ancestral geographic range occupied by the subdivided populations. At one extreme we can envision equal partitioning of the ancestral range. At the other extreme we can envision the budding off of a very small part of the ancestral range, per- haps a single deme or even a single individual. Thus three models may be distinguished, two of which are extremes of a continuum. ALLOPATRIC SPECIATION I— VICARIANCE Vicariance speciation (Wiley, 1981) is a gene Differentiation may be manifested before geo- graphic separation in the form of geographic variation, or it may not. Speciation may be rapid or slow depending on the amount of geographic variation existing before subdivision and the oc- currence of evolutionary novelties after subdi- vision. Wiley (1981) suggested that the following phylogenetic and biogeographic predictions could be made based on this mode: PHYLOGENETIC 601 WILEY & MAYDEN FIGURE 3. 981) 1. Dichotomous phylogenies should predom- inate because of ancestral “extinctions,” i.e., be- cause there is no particular reason to think that either geographic subunit will be identical with the ancestor as a whole (Fig. 4). 2. The range of an ancestral species may be estimated by adding the ranges of daughter species. 3. Different clades should show common (congruent) patterns of speciation over time and pace. Predictions | and 3, dichotomy and some level of biogeographic congruence, are the important predictions. Of course, the congruence observed will be limited to those groups who (1) inhabit Distribution of the Fundulus olivaceus species group (from Lee et al., 1980 and Suttkus & Cashner, the same region, (2) are actually affected by the geographic event in question, and (3) have a his- tory of pev i queue that has kept pace with the subdivi Vicariance — Cont tral Gulf Coast specia- tion. A number of clades of fishes contain sister species whose boundaries can be defined by the Mobile Bay drainage of the Central Gulf Coast of the U.S.A. Within the freshwater Fundulus nottii species group (Wiley, 1977; Fig. 5), F. not- tii is found from streams draining into Lake Pontchartrain, Louisiana eastward to and in- cluding the lower reaches of the Mobile Bay drainage, Alabama. Its sister species, F. escam- biae, is found from the Perdido River Drainage, 602 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 NO P Alabama to the lower Suwannee River, Florida. > Phylogenetic analysis (Wiley, 1977) indicates that x this species pair is most closely related to F. lin- eolatus, an Atlanti tal i tric witl F. escambiae (C. Gilbert, pers. comm.), and that P4 VV RULE the three together have affinities with the western sister pair F. blairae—F. dispar (see Fig. 5). There ; u an 6), coastal and estuarine killifishes (Wiley, in Ss press). 2. Etheostoma chlorosomum and E. davisoni (Fig. 7), freshwater darters (Howell, 1968; Page, 1983) FiGURE4. Speciation in the hypothetical clade NOP illustrating phylogenetic patterns expected in model 1 speciation. Md l L ; ) el - ty hein FE, SB 4 K AA | == ba^ ze LA a fd) s et ANNA es | Z ⁄7 UR pg Ufa e pl CHR AOAN, PEALE NP INOA oe \ TA T Sud Ç ° h i + ET UN ^s il RTI NS HY il visto, es SOS t if HT SKA MP isl NIE, — ^ Nr hi D A H \ 1 M & yas MANTA NQ ih Bl, NU ATT Y ASAT TD VF NOU) 0 SARA HN NG KK , N Ni y | LLAP + V Ka I l: > N F d "aa. Mi an j HA EB" ada" oR. >) Ad n | DEDERIT ay SRE A d) HHEES 2 EN | ET ARE N FiGURE 5. Distribution of the Fundulus nottii species group (from Wiley, 1977, with additional data from C. R. Gilbert, Univ. Florida). Tree: [4+5]+[3+(1+2)]. Characters supporting tree: see Wiley (1977, in press). 1985] WILEY & MAYDEN PHYT / : V — `> — M f SENS. Z Al I AX WA Sep = w: pA € < Eo A 17 PNA { ! PA AH CENE. Á 5 \ à \ i (M SN A A — MT A \ z f S ES TM 3 S uu. E CY i N (Cx -— ue ) f ¢ oc d Po ed | "« EA. a 4 = | JE | Oe ed N. T Cran Pai Š> Pt 1 Pi — a WE. M ; M À 1 je uc] NE fi LA E | AU: T XS X M. j ER Ji ENS R) I F. pulvereus, fe x URE6. Distribution of Fundulus confluentus sd x. pulvereus. (Reylea, M.S. Thesis, Florida State Univ., lon FiG 1965, recognizes most Atlantic populations north o ida as belonging to F. pulvereus and h e recognizes a zone of intermediates between the Apalachicola River sies! Mobile Bay. We do er follow this, but we recognize the need for further study.) 3. Ammocrypta beani and A. bifasciata, freshwater sand darters (Fig. 8; from Williams, 1975) 4. Hybopsis winchelli and an undescribed species (Fig. 9), freshwater chubs (Clemmer, 1971; Lee et al., 1980 In addition, several groups of snakes show vi- cariant patterns along the Mobile Bay basin, some on the level recognized as species, some as sub- species. These include: Nerodia rhombifera and N. taxispilota, eter snakes (Fig. 10; Mount, 1975) . Two danapecies of Nerodia cyclopion (Fig. 11; Mount, 1975). 3. Two subspecies of Farencia abacura, the mudsnake (Fig. 12; Smith, 1938; McDaniel & Karges, Additional possible examples of this pattern also exist, but are complicated by the presence of a Mobile Bay drainage endemic, borders which do not exactly correspond to the event, dispersal, or the lack of an adequate phylogenetic hypoth- esis. These include: 1. Notropis longirostris and an undescribed species, two shiners, with the undescribed species endemic to the Mobile Bay drainage and N. /on- girostris found both east and west (Fig. 13; Swift, 1970; Lee et al., 1980; R. D. Suttkus, pers. comm. ). 2. The Notropis roseipinnis species group, FIGURE 7. three shiners, with the Mobile endemic being the sister of the species east of the Mobile rather than west (Fig. 14). Snelson (1972) presented dif- ferent possible phylogenies of these species and recently Stein et al. (1985) considered the above relationships to be most parsimonious. . Ammocrypta meridiana, a M a ee en- denne is the sister species of th X—A. pellucida species group (Fig. 15; Sia. 1975. Page, 1981). This pattern is similar to that of the ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 » Fr [i I as "i qanuna LISTA Se ase Distribution of Etheostoma chlorosomum and E. davisoni (from Page, 1983). Notropis roseipinnis group and has a Mobile en- demic, but differs from it and the F. nottii group in lacking a species east of the Mobile. 4. The Notropis hypselopterus species group in which there is an apparent peripheral isolate, N. euryzonus, closely related to the eastern species N. hypselopterus. The western species, N. sig- nipinnis, shows a broad area of sympatry east of the Mobile (Fig. 16; Bailey & Suttkus, 1952; Sutt- kus, 1955; Gilbert & Platania, 1980). 1985] 5. Percina sciera and P. nigrofasciata (Fig. 17) show a pattern opposite of the Notropis hypse- lopterus group with the eastern species being widely sympatric with its sister west of the Mo- bile. This is further complicated by having two additional species (P. aurolineata and P. lenticu- la) of this clade in the middle of the distribu- tional pattern of the clade (Lee et al., 1980; Page, 1981). 6. Two species of the turtle genus Sternothe- rus, S. carinatus and S. minor, show a pattern that might have resulted from a vicariance event involving the Mobile Bay drainage, but the pat- tern is complicated by the presence of S. minor in the Pearl River drainage of Mississippi and Louisiana (Fig. 18; Tinkle, 1958). 7. Two subspecies of the cottonmouth, Ag- kistrodon piscivorus (Fig. 19; Conant, 1975), two races of kingsnakes, Lampropeltis getulus, and two races of the pinesnake, Pituophis melano- leucas, have borders that correspond to a Mobile Bay vicariance (Mount, 1975) but in each case there are additional races or subspecies and there are no phylogenetic hypotheses demonstrating that the races with borders at the Mobile Bay drainage are closest relatives. nother group of species show evidence of a more western vicariance event loosely associated with the Mississippi River. Within the Fundulus nottii species group, F. blairae and F. dispar are largely confined to the western part of the range (Fig. 5) and are the sister group of the eastern members of the group. A similar pattern is seen in Ammocrypta, with A. clara being the sister species of the beani—bifasciata species pair (Fig. 8). Additional examples might include: 1. Notropis sabinae, the sister species of the N. oo complex (Swift, 1970; Coburn, 1982; Fig d. edu. umbratilis-N. ardens species pair, sister group of the N. dapi group (Snelson, 1972; Mayden, unpubl. data; . 14). Other groups may show a Cale pattern, but the phylogenetic relationships among the mem- bers have not been investigated. These include: 1. Two subspecies of the turtle Deirochelys re- ticularia (Fig. 20; Zug & Schwartz, 1971), but there is a third subspecies. 2. *Notropis" amnis and the Hybopsis am- blops complex (Fig. 9), but there are at least five species inhabiting other regions whose relation- ships are not known prenne 1971; Coburn, 1982; Mayden, unpubl. dat In summary, there are igna two large-scale WILEY & MAYDEN—PHYLOGENETIC SYSTEMATICS | | o| | | € tag L. 1 IN s) Distribution of io Ammocrypta beani 975). Tree: AY À 2). Characters supporting tree: see Williams (1975) M a rer o P ^ GURE 8. species group (from Williams, 1 vicariance events on the Northern Gulf Coast, one splitting the areas east and west ofthe Mobile Bay drainage and one involving the areas east and west of the Mississippi River. For those groups that show both patterns, it appears, based on the phylogenies, that the vicariance centering around the Mississippi is older than that center- ing around the Mobile Bay. Vicariance—the Central Highlands. The Central Highlands of the U.S. may be broadly conceived of having two major components. The Interior Highlands comprises the Ozark Plateau and Ouachita Highlands. The “Tennessee High- lands" comprises the Interior Low Plateau, the Ridge and Valley Province, and the Blue Ridge Province—essentially that part of the Central Highlands drained by the Ohio, Tennessee, and rl Rivers between the Appalachian doubt floral) endemics and the closest relative(s) of an endemic in one area is likely to be found in the other area. In Figure 21, we have plotted mx ss xe 9X wXxo002 59€ ZEBOXONS A Xo BARUK XO x ots 2 5 er GURE 9. Distribution of bsc sae: and Apa n ipe E A al. ophyly of the k of the urohyal, (3) the total ranges of 44 species of freshwater fishes whose ranges are largely confined to one or both regions of the Central Highlands. (The many areas plotted north of these regions define various dis- junct populations of species found in the region.) Not shown on this map, but plotted on subse- quent maps are the ranges of close relatives out- lands. The southern boundaries of the majority of the species shown in Figure 21 are largely congruent and form a border with the margins ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 Zi - - = . US P=. mir. , 1980). Tree: (1+2)+3+ p: (1) oval (2) prominent lateral s spines on fi s ad T (4) long ges on first eo. (5) enlarged first pair of ypobranchials, and (6) elongate interhyal (Mayden, ie data). of the Cretaceous sea s: with the Coastal Plain. The n rn boundaries of the Central Highlands s are not distinct owing to numerous disjunct populations of species oth- erwise confined to the two regions. Geologic data indicate that prior to the Pleis- tocene many of the streams of Ohio, Indiana, and Illinois were high gradient and thus suitable for highland fishes (Thornbury, 1967; see May- den, 1985, for summary). Further, drainage pat- terns were different, and what is now loosely re- ferred to as the “Central Lowlands” was a high- land region prior to the Pleistocene. Much of the 1985] WILEY & MAYDEN —PHYI 607 ] ` x c ` S 2 I D ^ * E: — UY t //) f H ^ ^n < ww. S - AS ATA 2S So SN: aw; A x SSO D E: (Oc C 9*09 909,9 4 4.9 Serene ees SOO Sa SO ve SN * > PR > F. esos Ch SN Mw". M 39:95 95.920069 a gre! 15, Me thes soi e. SPS SOW OSCAR e OSS RS EP CORO FRO SSG SAIS KER ORC: Noe PEAS $ — OD `< 9.5 s 4 > A 9j - z FiouRE 11. Distribution of two subspecies of Nerodia cyclopion (from Mount, 1975 and Conant, 1975). 608 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 t CN Ay Ed — -— “sis s x ~ s S Ü -A ie 25 CoS un A wa f h `. ` A k y d. (S ; ` ? rd Y SRS ^ e L =o à Ur F T NR 1 pP UT LN rh i == AUR A eee U, S UM * |j ANILARA IT EDEN (i "WEE We v^ F. abacura abacura ZL EE Vox v IOAN XT af os f N 1 Y 5 ; i AD } H l l $ ' d k t y 2 ii ^x | lil L^ 1 B = ^ ur š Tm yi N oii M | pe » É H ÜN | S LIP D 5 i: I ili hac ~ | LTEM à n | X i in TNI ! LY 2 [es 2X y D [4 i °: " N mma. "ET ease - Ko / 2 Y \ l m — F. a. reinwardti: k: AY i LCLC d iS — a 2 Lei A sp — L4 u SS TR CAEN RUN AER e FT t; IE Bull n id ee — LE X ——— -— 2-02 h ; r 424 k” SA ZI am par m "a. w = “ram am = Zuma — Ld £. K— ry W on ED SS LOT m ru ze un a tA eed Y s Y. [-cn -— ) SS NL m. tra — a J É "ED PARE OPED AEG) aN > Dar - = a siR.` ~j-—--"* "n 1 — ÁÀ mÓ FicunE 12. Distribution of two subspecies of Farencia abacura (from Smith, 1938 and McDaniel & Karges, 1983). Tennessee Highlands was drained by the Teays ( i 7 } 5i A, AN N a ) `S : š é x "E . y te EU boc DNO Dux . River (labeled 9 in Fig. 22) that joined the Mis- SEAT CY vo EVITE c sissippi north of St. Louis. To the south the old < SV Vi teo 1777. 7. Ohio (labeled 10 in Fig. 22) joined with the Cum- — hes, “orangetin shine. — berland to empty directly into the Mississippi Z ka s: WoS M and did not join with the Tennessee (see Hocutt ^ UNA N : : dol NORMA = et al., 1978, for a review). The present drainages 1 hs are a consequence of Pleistocene phenomena. Thus, prior to the Pleistocene there is evidence that (1) the highland fauna covered a more ex- tensive geographic area than today and (2) the Aia drainage patterns were much different. longirostris | | NN | | | b pem E icm d de ie exe | — pers. comm.). Tree: 3+(1+2). Characters supporting x | | | | | 1 | l monophyly of the group: (1) maxillary process of pal- t ongate an eriorly directed, and (2) enlarged atine el te and a FIGURE 13. Distribution of the Notropis longiros- plate-like second infraorbi tris group (from Lee et al., 1980 and R. D. Suttkus, Characters uniting (1 +2): see Swift (1970). ar. S Y ox Ly LE———— -— m e taba’ L5 — a REV ne a. WILEY & MAYDEN L | p.t : > V “Et IE —— ee SS ee ED — ’ N AZ SAN | Sere EE - 485770 A i / ` | een — eae ~ Sa 1-4 2————Z ' - 4 if Ww ——— ————- PZN f p =< ` / E == - ; 4 d fF es A Á = Or SES ) Pi l : — = == : - an 4; = — ^ - Z= A À ~ Fe ——À—— oe 7 === LI TE E. Y Y, —— ——= x LARL CEDE or x WY um —— === LATA Eu PU [X x 5 L; C—O e "uy =S Z — CAM e, aCA, Mis S ; H =>: Ne — = 1 EL NE k ; WU I^ ) u ee oe Lm — — 2Ó ES——1 22. F MN eS eee e h eaa í SS Lf -—g —— BË / i/o +N ! x a T, —À T /&rF V = See eee Mest — D Y Uy 7^ : — W— — EL ERU T LE A CE / (7 /, f ——Á — i. m e, Ep ARE * / 9⁄2 f/ l m es P S= sa ==. xx M, 7 y SS ee ee >= — J UY T. ; — — u; i r= Oé i a Z ó U SS SSS SS uk 14 S < LN Á ` FS NT a AE: cere. Y . ( N Si LOU ESS ES SE Sen” \ y LLL —rr RA A Re! SS NEEN NAA ME. — "a ag a" Se a it t < — geen v — et i. SS SS | "Riz = j SS UN N. umbratilis — Z— —— tL; NS a E= E RY — Ee WSS Ç EC ES ES Ss / { ji al n | SS FE | | k.;-2Z=— = 'F M C SSS === lilii IN — P RUNG Mba \ ee ee we —-- Sr eae Vs cir MM aY Waku. KN "Z d x A. . < S > Je N. roseipinn [X - \ Z X ¿À š 5.) SW \ I a b ERN x 1 ) Y — d <4 o F M x w y | 14. Distribution of the Notropis roseipinnis species group [37-4] [2- (1 - 2]. Character uniting the group: breeding colors of males : dark vertical (Mayden, unpubl. data). Character uniting (3+ 3) Tare ON ASS E N. atrapiculus S. 1980). Tree: from Lee et al., ting of bright blues and reds f consis bars on dorsum of breeding males (Mayden, unpubl. data). Relationships of [2+(1+2')] are discussed in Stein et al. (1985). The distinctiveness of the Central Highlands is partly reflected by the presence of species large- ly endemic to both areas. Among these are the minnows Notropis galacturus (Fig. 23) and N. boops (Fig. 24), the extinct sucker Lagochila la- cera (Fig. 25), and cavefish Typhlichthys subter- raneus (Fig. 26), the darter Percina evides (Fig. 27), and the salamander Eurycea lucifuga (Fig. 28). The distinctiveness of each region is also reflected by the presence of numerous endemic species that are not considered in this analysis. When we presented data at the symposium concerning the Central Highlands fauna we thought that speciation within the clades inhab- iting these regions might be due to Pleistocene glaciation. Glacial activity caused the regions to the north to become disjunct by depositing till and/or loess and silting of the streams in areas now referred to as the Central Lowlands. This might be termed the “Pleistocene hypothesis." The question is not whether the Illinoian (and probably the earlier Kansan) Glacier caused a 610 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FiG 15. Distribution of the Ammocrypta pellucida species group (from Williams, 1975). Tree: 1+(2+3). reru discussed by Williams (1975). disjunction. Rather, was some of the speciation within the Central Highlands fish fauna corre- lated with the event(s) or, were the species we observe today extant at the beginning of the Pleistocene and perhaps speciated, in a drainage system pattern that was subsequently replaced by the pattern we see today? This latter hypoth- esis might be termed the “‘pre-Pleistocene hy- pothesis.” Clades that conform to the Pleistocene hy- pothesis would be expected to show the following attributes: (1) they have one speciation event that WILEY & MAYDEN Tas: xA : PX N. guryon / ” `= - AA D " I c É N. signipinpis AS wA l à > S ` R d z L) rey a^ um as = Lr === oes rs Fre — FiGuRE 16. Distribution of the Notropis hypselopterus species group (from Gilbert & Platania, 1980). Tre hyly of : e Per p: (1) very shallow preethmoid socket on the mb scent colors (Mayden, unpubl. data). Characters (3) iride supporting (2+3): breeding tubercles on mandi i. directed laterally and in “comb” rows (Mayden, unpubl. data). is congruent with the Interior-Tennessee dis- junction, and (2) no speciation event within either region can be correlated with an event that was pre-Pleistocene or pre-Illinoian in origin. Three species groups conform to these attributes: . The Notropis spectrunculus species group (Fig. 29). A clade of two described and one un- described species (Ramsey, 1965; Mayden, un- publ. data). Only the described species ranges are shown, the undescribed species is found in the Tennessee and Cumberland River systems and is the sister species of N. spectrunculus (Branson, 1983). 2. Etheostoma niangue and E. saggita (Fig. 30). Most workers (e.g., Bailey, 1948; Page & Whitt, 1973) have considered this pair sister species. om to other members of the genus are not c 3. Percina s ona" ia and P. sp. cf. cy- matotaenia (Fig. 31) (Page, 1974, 1983). is is not an impressive list. Further all three clades show wide geographic disjunctions be- tween species that indicates much extinction and, in no case do we have the phylogenetic infor- mation necessary to relate any of these clades to other species within their genera. Finally, while they conform to our expectations regarding the Pleistocene hypothesis, they are also perfectly consistent with the pre-Pleistocene hypothesis. Clades that conform to the pre-Pleistocene hy- pothesis may be expected to show the following attributes. (1) They contain one or more species in both the Interior and Tennessee Highlands and one or more additional species endemic to one of these regions, indicating that speciation in the clade and the achievement of sympatry occurred before the: Pleistocene. (2) 1 contain a species (or populations of eu Pleistocene ‘origin, dating that species as existing before Illinoian glacia- tion, the last known event to split the Central Highlands. (3) They contain species whose origin can be dated as early Pleistocene or pre-Pleis- tocene and who are of more recent origin than the taxa conforming to the disjunction between the Tennessee and Interior Highlands. at has relict a X — a Ya Í — f n í i i Y E : — hp NÀ `í — ZA Eo y A es ciat ERN oe EMEN rr ET NE —+ — +4 -— Z—=— 71 ST a (í a EN R21 —— ae [— — rJ F ~ a a s ee § — iP. p MEL amm v pend ph — 24d f — 7 $ , RN ——— ea WO REA U == I A aD e m x. MEA ' Um ATEEERERZUM Y. — - Í — Z -— | 9 qam amn a: A FA, w— M = T. >— P amarum cwm ~ ° - = pe asd eres — SI 2 LT í URE 17. Distribution of the subgenus Hadropterus, genus Percina (from Page, 1974, ANNALS OF THE MISSOURI BOTANICAL GARDEN TT [Vor. 72 f e — € ro | an fe Fy Ze Y y ^ esI Ç V LI LT S i ` i SLAI x - r AS NS bs PA ur 7 eee SS ee 55 5 W pt = \ c3 — v XOU --- š re 7 e" Xe A = a % EOM X Y £—— h Ly ^ w Co Cod" vH Lx 3 Nd KY XX ` Le DO LE . REAL A "s t ORZ SNAN PUO Q5 N x FN ANOS s E — = È \ AAA NOS _ Š N eQ ——— 5 v = W f ` % ` > ` D am WM I \ i ” = — — ». B OD. r. reticularia ^ —r t- N wl : Y yy I VP Za Xe d " pa FM eX xS A ded as i Y D ey | I AW _ 1 A FA, v 1 y ^ Bl I» Z T Yai ete =s == OO ————— TT p ee WW 1 7 a i= —— C o 225. oe oS ep E ee ——— 4 NS ST E TA I E S ee ZT Esa m EO aE — ri 2Á———= — a jg — a TNS mC —— o — SL ru q ==. — =, =——— XU == W 5 s ALL A J NS ESI A. v. CIE by he \J Ficure 20. Distribution of three subspecies of Deirochelys reticularia (from Zug & Schwartz, 1971). Tree: unresolved. This group contains five species of chubs char- acterized by having a unique barbel morphology (Mayden, unpubl. data). The species relation- ships are unanalyzed, but H. insignis and H. dis- similis share one derived character (presence of lateral pigment blotches on the flanks), which indicates they are sister species. Hybopsis dissim- ilis is found in both highland regions and is broadly sympatric with H. insignis. Further, H. insignis has a number of relict populations, some of which correspond to regions along the south- ern edge of the Illinoian Glacier. Hybopsis x-punctata is found in the Ozark Highlands and has several disjunct populations including one in the Ouachita Highlands, several north of the Ozarks and several congruent with the eastern disjunct populations of H. dissimilis. 3. The Notropis telescopus group (Fig. 35). This four species, which includes two species (N. telescopus an burn, 1982). Notropis telescopus is found in both highland regions. Notropis ariommus is more 1 1 lat+ 4 «+h tt LC + LI;:—-hl » | : VANOMVA SWIG LV MII L D F and it has a number of disjunct populations in the Central Lowlands. 4. The Notropis zonatus—coccogenis group (Fig. 36). In this case we have five species and two clades (Buth, 1979; Buth & Mayden, 1981; Mayden, unpubl. data). The clades show the In- terior- Tennessee split. However, only N. cocco- genis is a Tennessee form and it is most closely related to N. zonistius from the Apalachicola River drainage. Further, of three disjunct populations, one of which is confined to the Ouachita, dating it as pre-Saga- monian. Thus the origin of N. zonatus and N. pilsbryi must have occurred prior to this (unless N. pilsbryi is the ancestor of N. zonatus). 5. The Notropis nubilus group (Fig. 37). Three species are members of this clade (Swift, 1970). Two are found in the highlands and one in the Mobile Bay drainage. Relationships among the three are unresolved. If two highland species, N. nubilus and N. leuciodus, are closest relatives, _— 615 WILEY & MAYDEN "E $ Y E t lor HI E de ad. MP UAA XR y AS VEO d q No Sp. AN Bd T Ta FF Pe N aei Uy - B v M t a y FicunE 21. Composite ranges of 44 species of fishes largely confined to the Central SE of the United slc nete disjunct populations found outside the region (mostly from Lee et al. then the group would seem to be a candidate for the Pleistocene hypothesis. However, N. nubilus has a northern disjunct population that occurs, in part, in the “Driftless Area" of Wisconsin, Minnesota, and Iowa (Flint, 1957) and is prob- ably a pre-Pleistocene relict (Pflieger, 1971). In addition, N. nubilus has a coiled gut found only inone othe 1970; Snelson, 1971) indicating that i it cannot be considered an ancestor that dispersed eastward uring the —— to give rise to the eastern members of the c . The Fiheostoma variatum group (Fig. 38). This group of darters comprises six species “Lory g d 1940; Richards, 1966; Mayden, unpubl. data). It is interesting because the Inte- FiGURE 22. Pre-Pleistocene drainage patterns of the Central Highlands and adjacent regions. 1 = Plains Stream, 2 = Red River, 3 = Ouachita River, 4 = Ar- kansas River, 5 = White River, 6 = Grand River, 7 = F< 3or Iowa River, 8 — upper Mississippi River, 9 — Teays River, 10 = Cumberland and Tennessee Rivers. (Mod- ified from Mayden, 1985.) É š F Y . galacturus j i ve E “ [S ea n VE. ` " E 2 al Ç FIGURE 23. (from Lee et al., Distribution of Notropis galacturus 1980). rior- Tennessee split does not correspond to two monophyletic groups. Rather, E. blennius is the oldest member ofthe group and there is evidence that there was a separation of the southern part ofthe Tennessee Highlands from the presumably continuous highland fauna to the north and west (essentially Tennessee River versus Teays-Mis- sissippi). The next speciation event involved E. sellare and the ancestor of the remaining species and was a highlands versus East Coast vicariance we cannot i pd (stream ula oe ty isolation?). The remaining i m a clade and have an ior Tennessee fuic w esa Whether the speciation event that separated members of this clade was the result of Pleistocene glaciation is problemat- ical. However, we could resolve this by showing that the origin of the E. osburni-E. kanawhae species pair was early Pleistocene or pre-Pleis- tocene. Unfortunately there are no data con- cerning the age of the Kanawha Falls (Hocutt et al., 1978), the geological feature that marks the boundary between these species and their sister, E. variatum. Alternately we could test the hy- pothesis by dating the time of origin of E. tetra- zonum and E. euzonum. Thus this clade may show “attribute 3" patterns, but this hypothesis awaits more data. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Other taxa are of interest. Five additional groups, one species of amphibian and four fish groups, deserve mention as possible examples pertinent to our analysi ryptobranchus allesunienem has an over- all distribution similar to the Etheostoma var- iatum group. One subspecies alleganiensis is found in both highland regions and there is a disjunct population on the East Coast (Fig. 39). The four fish groups comprise either a single species for which subspecies are recognized or several closely related species. Lack of phyloge- netic hypotheses concerning their relationships precludes their use in the analysis. 2. The subgenus Swainia, genus Percina (Fig. 40). The group contains four highland darters (one undescribed) and a widespread species (P. phoxocephala) not confined to highland habitats (Page, 1974). 3. The Etheostoma maculatum species group (Fig. 41). This group is interesting in having high- land species, disjunct populations of one species (E. maculatum), and a southern disjunct species found in the drainage adjacent to that containing the southern disjunct population of Fundulus ca- tenatus (Williams & Etnier, 1982). Thus it con- tains species relevent to testing a Gulf Coastal Plain connection as well as a highlands connec- z o n. 4. Etheostoma blennioides. Several subspe- cies are recognized (Miller, 1968), and the dis- tribution of these is similar to others included in the analysis. 5. Cottus carolinae. This species includes three highland subspecies and a subspecies endemic to the eastern Mobile Bay drainage (Lee et al., 980 — Vicariance—the Central Highlands—discus- sion. The evidence we have presented indicates that speciation in highland fishes cannot be cor- related with the vicariance of the Central High- lands by the Illinoian Glacier. Rather, the evi- dence indicates that most or all of the species analyzed were present and living in the region before the Pleistocene and thus speciated over a landscape including drainage patterns not found today. Our data are not consistent with the idea that most of the groups analyzed lived north of several clades have relatives in the Coastal n (Mobile and Apalachicola drainages) and some have relatives on the Atlantic slope. This implies a history of contact between the high- WILEY & MAYDEN PHYTI K US », “Qe A qr v, A Lng 2 a By A FicurE 24. Distribution of Notropis boops and N. xaenocephalus (from Burr & Dimmick, 1983). lands and the Gulf Coastal Plain. We do not wish to imply that the present disjunction is not cor- related with the Illinoian Glacier. It probably did cause the disjunction we now observe by mod- ifying the area north ofthe present highlands into the Central Lowlands. Further, we do not wish to imply that some speciation within either the Interior Highlands or the Tennessee Highlands had no correlation with Pleistocene events. It probably did. What we do wish to imply is that the simplest explanation for speciation observed in three species pairs we first discussed is not corroborated when we analyze the fauna as a ole. While large-scale vicariance speciation (Model I) might explain much of the evolution in the clades we have analyzed, glaciation does not provide a simple causal mechanism of vi- cariance correlated with speciation. ALLOPATRIC SPECIATION II — PERIPHERAL ISOLATION Peripheral isolation models postulate the or- igin of a new species in a very small portion of the range of an ancestor or colonization and sub- sequent differentiation of a part of the earth not occupied by the ancestor. One problem with de- ucing phylogenetic and biogeographic predic- tions from this mode is that there are at least the case of what might be called “classic ripheral isolation, the model invoked has a dis- tinct ecological component as well as a historical component. New species arise in marginal hab- itats, usually but not always at the periphery of the ancestral range. This is the model favored by Mayr (1963), Hennig (1966), and Brundin (1966). W "4 Ig 1 — Pi - n im Sk > Y 4 `X FIGURE 25. Known collection localities of Lago- chila pons (now extinct, from Lee et al., 1980). I+; LEE | +h ti lat: — AK < S kN FiGure 26. Distribution of Typhlichthys subter- raneus ion Lee et al., 1980). ancestral species always go “extinct” at branch points. We do not believe it is necessary to pos- tulate this. Without arguing the merits of the metaphysical case (we have no stake in the out- come of such an argument), the hypothesis is clear. seed s ( l Pe) ee rule applies. One specie isolate, diverges more from the em condition ls the other. A variety of predictions can be generated under this model or models). These fall into two general classes, predictions from single speciation events and predictions from multiple speciation events. Peripheral — isolation—single speciation events. Peripheral isolation involving one cen- tral population and one peripheral population results in a dichotomous tree at the level of species. Thus the tree pattern is identical to that — r the North American killifish genus Fundulus, F. diaphanus is a wide-ranging species and its sister species, F. waccamensis, is restricted to Lake Waccamaw in North Carolina, and one artifi- cially introduced population (that can be ig- nored) found in Phelps Lake, North Carolina. In describing F. waccamensis, Hubbs and Raney WILEY & MAYDEN FiGure 27. Distribution of Percina evides and P. palmaris (from Page, 1983). (1946) suggested that it was a peripheral isolate derived from F. diaphanus. To what extent is this hypothesis justified? One problem in considering this hypothesis is that the ranges now occupied by F. diaphanus and F. waccamensis could have been achieved in several different ways. In Figure 42a we show the geographic relationships between two hy- pothetical species, A and B. In Figure 42b we show the origin of B as a peripheral isolate de- rived from a “small-scale” vicariance event. (The same effect would be achieved by dispersal of one or more populations ancestral to B into the area followed by differentiation.) This hypothet- ical scenario, similar to that favored by Hubbs and Raney (1946), requires little or no alterations in the range of A nor does it require doengrios The same result, however, could be achieved by very different scenarios, three of which are shown in Figure 42c, d, and e. These include (1) peripheral isolation of A with subsequent displacement of B into its present range (Fig. 42c), (2) a large- scale vicariance event resulting in the origins of both A and B, with subsequent displacement of B (Fig. 42d), and (3) widespread extinction of the 620 v ( `N ——N f A = / Zo MU AL VAM T / K 1. L- "" +, pr : I N y ; Z A AAS RK in E NTC nA: "4 Eurycea lucifuga ui y^ AY ANNALS OF THE MISSOURI BOTANICAL GARDEN FicunE 28. Distribution of Eurycea lucifuga (from Conant, 1975). original ancestral populations with both A and B originating in the resulting refugia, followed by a re-invasion of the original range of the ancestor by A (Fig. 42e). The problem is—how can we sort out these four very different scena- rios? One rather direct approach might be to adopt a parsimony argument and claim an 1t is sim- plerto adopt a model that xtinctions or dispersal (i.e., Fig. 42b) because uh a mode contains the fewest ad hoc assumptions. We would reject this purely logical approach for two reasons. First, parsimony is a logical device and cannot be used to test a proposition. It proceeds by weighing conflicting data and there are none. Second, the importance (or lack thereof) ascribed to this model of speciation by some make it im- portant enough to be approached with some cau- tion. We suggest that certain lines of evidence might be developed. Some are “extrinsic” in that they deal with data concerning the histories of the respective geographic areas inhabited by the species involved. Some are “intrinsic” in that they deal with data derived from the species themselves. Extrinsic evidence might take several forms. We might demonstrate that there is no evidence for climatic change or large-scale vicariance events occurring in the region presently inhab- ited by species A that would have effected it or its ancestor. This would reinforce the idea that neither Figure 42d nor e are applicable. We might find that the region occupied by species B either did not exist or was uninhabitable in the recent past, requiring a hypothesis of dispersal into the area. We might be able to identify a “small-scale” vicariance event with the isolation of the ances- tral populations of B. Intrinsic evidence is more difficult to speculate about, because the kinds of data derived from character analysis may depend on levels of gene flow between populations, the time that has passed since vicariance or dispersal, the history (or lack thereof) of gene flow between the isolate and the central population, and the effect (or lack 1985] WILEY & MAYDEN-—PHYI IC 621 NA M à A S M. ozarcanus M ys RE 29. Distribution of the Notropis spectrunculus species group (from Ramsey, 1965; Mayden, e n » ul An undescribed species reported by Branson (1983) is found west of the range of N. spectrunculus an artly e with this species. We have not plotted the range of the undescribed species for hen of dilkributional data VS J. AY fret ; ANIS MON f AN > NEM, "S xv E 1 ux jM T 7/ < E 4 ; [oS NC f E i ç] Mee š u^. E. nianguae : > (Of ACA TM URS d doe sta a f 4 " P “` | G NY Ct Lote MN & CN a b; — NY M I Bo pr s i | = - Z FicuRE 30. Distribution of Etheostoma niangue and E. sagitta (from Page, 1983). 622 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 NM N g NS A ies s < r =. AN et c N " rd GAS 9 E \ | «€ AR See PR AVI 5 ` P. cymatotaenia ao FIGURE 31. Distribution of Percina cymatotaenia and an undescribed sister species (from Page, 1983). F E 32. Distribution of the Fundulus catenatus species group (from Lee et al., 1980 and Williams & Etnier, 1982). Tree: 3 [4 5 * (14 2)]. Characters supporting tree: see Wiley (in press). WILEY & MAYDEN PHYLOGENETIC 1 A m X: X : uh i Í Ç IS 1 a j 2 ob FiGURE 33. saree tase AA E HH eS HEH RLL A ar CUTE LINT] ri Miu [1 LT [| Distributions of some of the members of the Hybopsis dissimilis species group (from Lee et al., 1980). See Figure 34 for tree and characters supporting the tree. thereof) of local selection on each population. Controversies exist concerning the population genetical aspects of founder effects and even the importance of peripheral isolation as a mecha- nism of speciation (for recent contrasting reviews see Carson & Templeton, 1984, and Barton & Charlesworth, 1984). In spite of these problems, we believe that certain lines of inquiry might be fruitful. One line of inquiry might be developed around the putative peripheral isolate relative to the re- constructed ancestral phenotype or genotype. For this to count, however, we would have to assume that the characters examined represent a random sample of all characters that have diverged. Fur- ther, lack of a sharp deviation (that is, the pres- ence of equal divergence in both lineages) might be compatible with peripheral isolation if the two species are of sufficiently ancient origin (thus pro- viding time for anagenesis in the wide-ranging species) and the wide-ranging species is charac- terized by high levels of gene flow (thus providing the opportunity for the spread of evolutionary novelties). Another line of evidence might be developed by considering the geographic variation of cer- tain kinds of characters. If species B is a periph- eral isolate, it was probably derived from pop- ulations of A near its range or, alternately, was in genetic contact with such populations before vicariance. This is probable if the scenario pre- sented in Figure 4b is true. Rapidly evolving sys- tems, especially if they were immune from se- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 iN a * V. Si D e E lh y d us & ? NS > — t "bs = J “monacha (5) `` M f \ IGURE 34. Distribution of the remaining species of the Hybopsis dissimilis species group (from Lee et al., 1980). Tree: 3+4+5+(1 +2). Characters supporting monophyly of the group: (1) an elongate basihyal, (2) stellate barbel morphology, (3) high vertebral count. Character supporting (1-2): distinct lateral blotches on the side of the body. (Characters from Mayden, unpubl. data.) lection and recombination, might show this relationship if a cladistic analysis was per- ormed on geographic samples of populations of A and B. Mitochondrial DNA has the kinds of evolutionary characteristics suitable for such an analysis (Avise et al., 1979). Another line of evidence might be developed by considering “ancestral clines," i.e., clines ap- parently present in ancestors that are retained in descendants (Wiley, 1981). At this level of anal- ysis, does the putative peripheral isolate fit into the pattern of geographic variation exhibited by populations of the supposed ancestor? If so, can we use this as evidence to prefer the scenario in Figure 42b relative to the alternatives? Perhaps if we can show that the characters are derived and heritable or that the pattern indicates a past history of genetic contact, then the species pair is a good candidate for the peripheral hypothesis. Returning to the F. diaphanus-E. waccamen- exist in the Middle Pleistocene. Thus its present freshwater fish fauna is the result of a colonizing event. (2) The lake is unique ecologically, being a clear, spring-fed lake surrounded by a tannic acid aquatic environment typical of the area. In terms of intrinsic evidence we note that while no detailed analysis of geographic variation has been published, F. diaphanus is comprised of two rec- . waccamensis than is the northwestern form. Thus what we know about the relationships of 1985] WILEY & MAYDEN—PHYLOGENET WIA I LLN Ic 625 , [^ _ i id i I es usd f ' N. ariommus + .- / JESN e : n> ee vd eo: í N. semperasper Y 1 = T SPs ci We À x > te WAS x AME ie — — = m === . Y. A rea \ N. telescopus , £ A, I uu p t LS LE: oo URS pon P F a~ ^ E n - SH RU AES POR FIGURE 35. Distribution of the Notropis telescopus species group (from Lee et al., 1980). Tree: 1+[4+(2+3)]. circus supporting tree: see Coburn (1982). - NL YZJ ` š WT 4N va WW 2 do LA — — CS ^. A AX > ao f T A MR Z1/A fo oy ae | | ms, NN I ha 4 Ip i} } ~ N \ šu ` aN BE 1 \ 9 H iy Í deut " MCA s [x » Rar i £ > N b ` d 4 n foa T 3 “Z 4 N. zonatus / ;— ` n" V £ ON | 2: N Ea us M px Pra In {~ Se NPP < PENN ER en E 36. Distribution of the Notropis zonatus—coccogenis species group (from Lee et al., 1980). Tree: uk 5]+D +(1+2)]. Characters supporting tree: see Buth (1979). — ae N — oo m — 3 —— — h =. — m em | < K — —— —— pod ir Distribution of the Notropis nubilus species group (fr ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 rom Lee et al., 1980). Tree: 3+(1 +2). FIGURE 37. Characters supporting monophyly of the group: (1) iridescent breeding colors, (2) dorsolateral pigment pattern cales heavily scalloped. Character supporting (1--2): presence of a dashed iridescent predorsal stripe. ta.) vios from Mayden, unpubl. data the two species does not preclude the hypothesis that F. waccamensis is a peripheral isolate de- rived from F. diaphanus. The problem is that such evidence is relatively ] idering tl di rt fthe phenomenon. In many cases the tentative nature of such speculations are compounded by the fact that the species pair involved is a singular and unique example. In the next section we will sug- gest that hypotheses of peripheral isolation ma be strengthened if we can find additional ex- amples under certain circumstances either in the same clade, or in different clades inhabiting the same region Peripheral isolation — multiple speciation events. Single and unique speciation events are 1985] WILEY & MAYDEN—PHYI 627 ys AB n ry Z ta S es AAA. dot š g Me s v . | ý Jte m j ad L p. 3 a së li s 274 5 ~ : j I ee pe yor > £ "s ED ATION Y X LEIA eke ee oar f p A À a ai poly NI "i^ vw m 4 P2 > w se » E f Í “A Pelee I (AA A or, [N C l «Nim dA t M I ú hv yas ) d . E. V y ; A 1 4 — 7YÀ y pm x PENG Z H anie Dr EM V ` ~~ E. euzonum ` i Mo ç t D x Jj Cu . AE I Qt ME. Ses M uf Y ays M i Á Z Z > ` ¿Z .sellare 77 E. se e / 20 Y y ^r 7? 228 LM " 41 TIR y | ue E. kanawhae = Vw >, — a xry - M. AUS > NN E KA TES “aN V T Sei T us K E. blennius. ~ X | | FIGURE 38. Distribution of the Etheostoma variatum species group (from Page, 1983). Tree: 7+([6]+[(3+(1 +2))+(4+5)]]. Characters supporting monophyly of the group: (1) fifth infraorbital pore and canal di rge dark dorsal saddles on the body. Characters uniting 1 to 6: (1) anal rays 9, (2) pr body, (2) yellow-gold bar on face. (Characters from Mayden, unpubl. data.) not easy systems to work with because we lack degrees of freedom to speculate about the pos- sible effects of dispersal, selection, and other phe- nomena that could lead us to the wrong hypoth- esis. A clade containing several species, more than one of which had some characteristics of peripheral isolates, or a biota containing an area of endemism inhabited by several suspected pe- . ripheral isolates would give us more room to speculate. Wiley (1981) considered patterns of descent involving peripheral isolation. One case involves an ancestor that “buds off" more than one de- scendant. Since the two descendants do not share a unique history, they have nothing in common other than sharing an ancestor (i.e., they lack synapomorphies uniting them). If the ancestor remained conservative and if it were treated as a single unit, a trichotomy would be obtained. If the peripheral isolation events were far enough separated in time and the ancestor was evolving, a dichotomy would be obtained. If the ancestor was very widespread and geographically vari- able, and there were numerous peripheral iso- a bones the ancestor appearing on the phylogeny next to or close to peripheral i tes in geographic p imity. We have no such examples in fishes, but if Shaffer's (1984) cladistic analysis of Mexican byst tid sal ders is fairly accurate, the phylogenetic pattern he obtained is what we would expect to see. Populations of Ambystoma tigrinum (the suspected ancestor, widespread in North America and found in each of the endemic basins of the Mesa Central) appears in the phy- logeny interdigitated among no less than 12 species (including both terrestrial and aquatic forms), of very limited range. The second approach is to search for different ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FiGuRE 39. Distribution of Cryptobranchus alleganiensis (from Conant, 1975). groups that have endemics in the same restricted area. We can then use the methods of vicariance biogeography to examine the phylogenetic and biogeographic relationships of the endemics to their sister species. For example, Lake Wacca- maw has three endemic species of fishes, each with a widespread sister species. One is F. wac- camensis. The second is Menidia extensa (a sil- verside). It is related to the wide-ranging coastal species M. beryllina. The third is Etheostoma perlongum, a darter related to E. omstedi. The distributions of the three species pairs are shown in Figure 43. Note that the three widespread species have little in common, their ranges do not coincide, they have different ecologies, and they are very distantly related. Given these ob- servations, coupled with the relatively late origin of Lake Waccamaw, it is improbable that the scenarios in Figure 42c, d, and e can explain a distributional pattern in which there is geograph- ic congruence of three unrelated endemics but little congruence between the closest relatives of these endemics other than the fact that each is found around the lake. The best explanation seems to be Figure 42b. Finally, we note that Lake Waccamaw has a diverse fish fauna (Hubbs & Raney, 1946), and the vast majority of species have not speciated in the lake. This indicates that while unique ecological circumstances might promote differentiation; differentiation is not an automatic consequence of changing the ecolog- ical parameters a population finds itself in. The North American fish fauna contains many species whose distributions suggest peripheral ies especially among darters and min- SW For example, wanqa, procne Is a wide- s, N. mekisto- cholas (Snelson, 1971) i a risi and peripheral distribution (Fig. 44). We might en- tertain the hypothesis that the origin of N. me- kistocholas involved peripheral isolation and that N. procne is the extant ancestor. However, our 1985] WILEY & MAYDEN—PHYLOGENETIC 629 mi (Gera R ` in p : EN i di | qe TOR x 5 Un i uM `Y y Wi qu l Numi M | O. x Wie 5 ri y rten ia Z3 (f | T x — x EN J, Nene l i E € À—E ====-. š ===. € = SS S= — SSS ZL———————À E————————À Z¿ZZ==x a S E R), TIL E ——— = === | = == ==: C — > — — —ÀE——t-1 AS (tt à A SES Q im i 1 | - V AU | n o M pS BU A A | | ha | ail LI d | i » A nh f i 7 wy PY s jude M 1j E E 40. Distribution of subgenus Swainia genus Percina (from Page, 1983). Tree: 1+[(2+3)+(4+5)]. P. debba supporting tree: see Page (1983) and Mayden (1985). lack of knowledge of the cladistic relationships tor, Fundulus catenatus (Fig. 32). However, what among populations of the hypothesized ancestor little we know of the relationships of this group coupled with our lack of knowledge of the closest indicates that F. catenatus is more closely related relative(s) of the species pair limits our ability to to F. stellifer, and thus these species do not con- ollow up our analysis. We can neither test the form to the expected pattern of peripheral iso- proposition by showing that some population or lates and their ancestors (Wiley, in press; data populations of N. procne are cladistically nearer from Williams & Etnier, 198 the descendant for some characters nor can we The continuum of allopatric speciation from show that the putative descendant conforms to large-scale vicariance and peripheral isolation i IS the deviation rule of Hennig (1966). apparent when we consider “‘mini-vicariance.” Perhaps as important as finding criteria from Mayden (1985) has examined several species which one can use phylogenetic analysis to cor- found in the Ouachita Highlands whose closest roborate hypotheses of peripheral isolation is to relatives inhabit the surrounding lowlands. These find cases in which phylogenetic analysis causes include eight species of fishes: two minnows (No- rejection of the hypothesis or indicates a need tropis perpallidus and an undescribed species), for collection of additional data. For example, two madtom catfishes (Noturus lachneri and N. Fundulus albolineatus and F. julisia are two ob- taylori), four darters (Etheostoma pallididorsum, vious candidates for the peripheral isolates hy- — E. radiosum, Percina pantherina, and an unde- pothesis and are surrounded by a possible ances- scribed species). In addition there are six species r ce maculatum | Ji. 7 cy YS RO AR , Distribution of the Etheostoma mac- FIGURE 41. latum species group (from Williams & Etnier, 1982 and Page, 1983). Tree: el ih s: unresolved. of crayfishes: Orconectes leptogonopodus, O. menae, Procambarus ouachitae, P. parasimu- lans, P. reimeri, and P. tenuis (for references see Mayden, 1985). The congruence observed points ent involving the dissection of the Interior nsas River, and a common event with species surrounding or south of the highlands. The fact that a relatively small geographic region was in- volved makes this pattern a candidate for “*vi- cariant peripheral isolation." There are many North American fishes that have restricted distributions, making them can- do the necessary kinds of analyses to test the model ALLO-PARAPATRIC AND PARAPATRIC SPECIATION Allo-parapatric speciation occurs when two populations of an ancestral species become dis- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 C Et s t Pl EL Ae | | ch Ce Cb EL FIGURE 42. Scenarios concerning the evolution of species A and junct, differentiate to some degree during the time they are allopatric, become sympatric in a lim- ited area, and complete their divergence because of interactions in the zone of sympatry. Para- patric speciation occurs when two populations attain lineage independance while maintaining a narrow zone of contact (see Wiley, 1981, for summaries of both models). Both models are hard to test, at least in fishes. An investigator postu- lating the allo-parapatric model must demon- strate that (1) the contact zone is a secondary zone achieved after a disjunction, and (2) that the differences observed between the species have either been evolved or “perfected” because of the interactions within the contact zone. An in- vestigator postulating the parapatric model must demonstrate that (1) the contact zone is a pri- mary zone, and (2) the differences we observe have been achieved in spite of the contact be- tween the two species. The simple observation of the presence of the contact zone is not suffi- cient. Testability is confounded because except for the presence of the contact zone itself, the phylogenetic and biogeographic predictions we can derive from both models are similar to those we can derive from the various modes of allo- patric speciation (Wiley, 1981). not have examples of either of these modes of speciation in the North American fish fauna. There are several reasons for this. First, it has been a tradition among North American ichthyologists to favor a rather strict definition of the biological species concept. Thus nominal taxa showing even limited interbreeding within a contact zone are usually considered as a single species and the possibility of either mode oper- ating is never considered. Second, freshwater fishes are confined to the streams they inhabit 1985] WILEY & MAYDEN— PHYLOGENETIC SYSTEMATICS Se oN \ \ | , F. waccamensis | i & ui? | Ir Ni ji ! M. extensa \ i Hi | \ Iib HINH lj" ij J * 2 E. perlongum [W H IND \ \ \ \ t l \ + \ J Wl; \ 1 \ E43. Distributions of three species endemic to Lake Waccamaw, North Carolina, and their widespread ) R closest relatives (from Lee et al., 1980 and this limits the possibilities for establishing contact zones and enhances the possibilities for allopatry between drainages. Thus, parapatric and allo-parapatric speciation would be expected to be largely confined to within-drainage systems situations. Third, we of the ichthyological com- munity simply have not been looking for evi- dence of either mode because we tend to think “‘allopatrically.” STASIPATRIC SPECIATION Alt though stasipatric speciation has frequently ynon u some mechanism such as drift or meiotic drive to fix the mutation in a small population, fol- lowing the model presented by White (1978). We have no examples of this mode of speciation in N findin methods for banding fish chromosomes are gen- erally made available. SYMPATRIC SPECIATION As discussed by Wiley (1981), there are several modes of speciation that do not depend on any degree of geographic subdivision and thus can be termed “sympatric” modes. There are no ex- amples of North American fishes that show an ecological mode of sympatric speciation. There are examples involving hybridization and sexual shifts that have produced unisexual species. These include two Nai ngage examples Poecilia formosa is a gynogenetic species (Hubbs, 1964) d in northern Mexico and 632 Seo Xcaa < MI 2 NN LI = —— m E A aY me h oe e G l ` | ( € ` À A Pa i AR N , \ y ` ) M o A y ayy zi J ` me J H. - { Y ^ rd A. ^ ^^. we 4 ki ` WA < p IC ` Y. v^ d wer 4 — \ EN | A, PLAT = (S uM > s Lam ta mam... ` UMS c M uk 16) < k. Ere qs Fa d 1 A 2.7 SÉ P - An ONT uu = 23 ~ \ E Cem reeves me ae £ E e ———34 Faw oaan mmen. en 1 T L è j X 4 = à w as we — W———! 44. Distribution of . procne and N. ; (from Lee et al., 1980). southern Texas. Hubbs and Hubbs (1932) sug- gested that this species originated by hybridiza- tion between the bisexual species P. /atipinna 1971) and the species involved in the hybridiza- tion event could have been P. mexicana or P. sphenops.] Although P. /atipinna and members of the P. sphenops complex are in the same sub- genus, they are not considered very close rela- tives. Menidia clarkhubbsi is a unisexual species of silverside found at three different locations in Texas (Echelle & Mosier, 1982). Echelle and Mo- sier (1982) hypothesized that the species arose M. ber- closely related to M. extensa than to M. penin- sulae (Johnson, 1975 In addition, there are at least five “forms” of the topminnow genus Poeciliopsis that are com- posed of parthenogenic females (Schultz, 1967; Moore et al., 1970). However, the nomenclature and relationships of these forms are not clear (R. R. Miller, pers. comm.). They may be examples of species derived from hybridization and sexual ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 shifts, sexual shifts alone, or morphs within their respective species. DISCUSSION AND CONCLUSIONS Analysis of phylogenetic and biogeographic patterns of descent and attempts to correlate these patterns with modes of speciation has hardly be- gun. Our attempts with selected North American fishes is a preliminary effort, but they suggest that such correlations can be made. Our first example, the Northern Gulf Coast, suggests a pattern of large-scale allopatry (model I) involving several clades. We note that the number of species in- volved is small relative to the number of species inhabiting the region and that the groups them- selves do not comprise a “biota” in the sense discussed by vicariance biogeographers (Fig. 45). Further, not all speciation followed a common pattern. Our second example, the Central High- lands, can only be considered a counter example. We observe vicariance without consequent spe- ciation. Apparently the species inhabiting the re- gions were present before the vicariance event that caused the disjunction of the fauna. How- ever, the effects of vicariance can be seen within the Interior Highlands in a pattern of “‘mini- vicariance" in the Ouachita Highlands. Our attempt to corroborate the peripheral iso- lation hypothesis proposed by previous workers graphic patterns involving putative ancestors and descendants are suggestive of this phenomenon, we lack the phylogenetic and biogeographic in- formation necessary to pursue such hypotheses. Two examples of multiple peripheral isolation were examined. In one (Fundulus catenatus, F. julisia, and F. albolineatus) hypotheses that the two species of restricted distribution are periph- eral isolates of the wide-ranging species are not corroborated because the wide-ranging species, . catenatus, is more closely related to F. stellifer. of the two peripheral isolates if it is, indeed the ancestor. (But if this is true, would the species that gave rise to F. albolineatus and F. julisia be F. catenatus?) The other example, species of am- bystomatid salamanders of the Mesa Central seems to offer a clearer picture, but we need ex- amples among fishes. WILEY & MAYDEN PHYLOGENETIC \ Kd E q-7- y ¢ Roa I I ! n I I + t 1 Si Y ÑW AU ent ÀA Ab SU RAIDS WA. EM NX x JU A te VEZ yor FIGURE 45. Summary map of distributions of taxa from the Central Gulf Coast and adjacent regions con- sidered in this paper. We were not able to provide evidence for either allo-parapatri parapatri lati tis pos- sible that, because freshwater fishes are confined to their streams, the opportunity for either mode to operate is restricted. Perhaps studies of coastal species where this restriction is not present will provide an opportunity to demonstrate these modes of speciation. The only sympatric mode of speciation well documented in the North American fish fauna is speciation via hybridization associated with asexual or gynogenetic reproduction of all-fe- male descendants. In neither case were the species of hybrid origin the result of hybridization be- tween closest relatives. 4 LITERATURE CITED AvisE, J. C., C. GIBLIN-DAviDSON, J. LAERM, J. C. PATTON & R. A. LANSMAN. 1979. Mitochondrial DNA clones and matriarchal phylogeny within and among geographic populations of the pocket go- pher, Geomys pinetis. Proc. Natl. Acad. U.S.A. 76: 6694—6698. BAILEY, R. M. 1948. 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MILLER. 63. Summary of late Cenozoic freshwater fish records for North Amer- ica. Occas. Pap. Mus. Zool. Univ. Michigan 631: 1-34. WnurirE, M. J. D. 1978. Chain processes in chromo- somal speciation. Syst. Zool. 27: 285-2 WILEY, E. O. 19 The phylo ogeny and systematics of the Fundulus nottii species group (Teleostei: uade. Univ. Kansas Occas. Pap. Mus. Nat. Hist. 66: 1-31. 81. P AN ics. The Theory and Prac- tice of Phylogenetic Systematics. John Wiley and E ssa York. ress. The evolutionary relationships of Finds topminnows (Teleostei: Fundulidae). WILLIAMS, x D. 75. inp ex the percid fishes ofthe subgenus Ammocrypta us Ammocrypta, with Prep cu i a asd. cma Bull. Ala- bama Mus. Nat. Hist. 1: 1- & D. A. ETNIER. "1978. theostoma aquali, a new percid fish (subgenus Nothonotus) from the ig and Buffalo ENS Tennessee. Proc. Biol. Wash. 91: 463-4 9 e odis of a new species, undulus julisia, with a redescription of Fundulus Are de and a diagnosis of the subgenus Xe- nisma (Teleostei: Cyprinodontidae). Univ. Kan- sas Occas. Pap. Mus. Nat. Hist. 102: 1-20. Zuc, G. R. & A. SCHWARTZ. 1971. Deirochelys, D. reticularia. Cat. Amer. Amphib. Reptil. 107.1- 107.3. GEOLOGICAL HIERARCHIES AND BIOGEOGRAPHIC CONGRUENCE IN THE CARIBBEAN! DoNN E. ROSEN? ABSTRACT If it is agreed that an understanding of biohistory in some ways is tied to an understanding of geohistory, the biohistory is explicitly tied to a particular geohisto geology. A propo different historical geologies of the Cari used by previous bioge ADDE in which g n as an example that contrasts with the con n one might also agree that what is needed is a precise means of specifying how a given we s type of analysis is the ographic area c cladograms ry. The constraint in ori notion of process, e.g., dispersal or extinction, was used to direct the outcome of TOORE a analysis. Whether espousing dispersalism TE 1957) or vicariism (Croizat, 1958, 1962), biol- ogists have always assumed that the EAER of organisms in some way reflect the nature of the world’s geologic history. It was, thus, implied that an understanding of biohistory is tied to an understanding of geohistory. In the Darwinian- arlingtonian tradition the relevant geohisto was assumed to be one of stabilism necessitating an interpretation of biohistory as one of active or passive dispersal. In other words, so long as the continents stood still something had to move to account for the occurrence of closely related organisms spanning large water gaps. Thus were invoked temporary land bridges suitable for crossing ocean barriers, birds with feet and feath- ers to which seeds and small animals would ad- floating debris that was spewed into the ocean's currents from river mouths and deltas. The difficulties inherent in these scenarios of a haphazard biohistory are apparent in Darling- ton's (1957) hypothesis that ostariophysans arose in central Asia and made their risky way along ephemeral freshwater routes to the southern con- tinents without leaving a trace of these great mi- grations. But the hypothesis of a steadfast ge- ography and a dancing biota was deemphasized after the theory of plate tectonics was elaborated. Since most biologists had a penchant for be- lieving that geologists had a special hold on the truth, the acceptance of comment drift panen £4 some of these L g nuring egy of proposing that it was the geography that moved while the organisms got carried about to their new longitudes and latitudes. Other biol- ogists (Darlington, 1965; McDowall, 1971; Briggs, 1984) tried and still try to rescue the past by agreeing that the geography did in fact move but that the timing of these great events was wrong in relation to the ages of the biotas. Such attitudes might invoke the ages of fossils to show that all the taxa are too young to have been influenced by the geographic cataclysms. This view involves two assumptions, both wrong at some level: 1) that fossils can tell us how old a taxon is and 2) that the ages of the geologic events have been correctly assigned. The first assumption is wrong because fossils give a minimum rather than max- imum age ofa taxon, and the second assumption is put into question by recent age reassignments. For example, parts of the Caribbean which were originally supposed to have been moving along a transform fault at the moderate rate of 2 cm per year (Kellogg & Bonini, 1982) are now be- lieved to be moving at the brisker pace of 4 cm per year (Sykes et al., 1982; Wadge & Burke, 1983), thus doubling the rate of motion and halv- ing the ages of the events associated with trans- location. But these relations leave us still at the mercy of general supposition when what is need- terms of time spans involved in human ue tual history, might be said to have d more g the work and review of the typescript I thank Drs. Kevin Burke, Arnold ordi uns ! For Parenti, Norman Plat nick, Edward Robinson, Peter Tolson, Lynn Sykes, and, especially, Gareth Nelso ? Department of Ichthyology, American Museum of Natural History, New York, New York 10024. ANN. Missouni Bor. GARD. 72: 636—659. 1985. 1985] or less simultaneously in systematics and bio- geography. This revolution has been called cladistics, the science of character analysis and the use of branching diagrams. Its premises permit precise comparisons between biological and geological systems (e.g., Rosen, 1978). The general objec- tive of cladistics is to discover congruence be- tween the two that constrains historical expla- nation. In the Darwinian tradition in biogeography it is dispersal that was the constraining concept, requiring an interpretation of the history of life in space contrary to what the data of life sug- gested. Leon Croizat (1958, 1962) was one of the pioneers who recognized that the biological data tell their own story, which can be at odds with a stabilist geology. Now that stabilist geology has been rejected in favor of a concept of mobilism, some biogeographers (Nelson & Platnick, 1981; Wiley, 1981) accept that biology has an inde- geological data that makes the comparison of the two so interesting because it is hard to imagine how congruence between the two could be the result of anything but a causal history in which geology acts as the independent variable provid- ing opportunities for change in the dependent biological world. The comparison becomes es- pecially interesting if there is a congruence among geohistories based on different approaches to the geographic problem, and if there is a congruence among cladistic relations of different taxa with respect to the same geographic areas. The specific questions are: 1) do the members of different monophyletic groups of organisms have the same relations to each other with respect to geographic regions in which they are endemic and is their congruence with respect to these areas non-ran- dom; and 2) does this non-random congruence of different groups of organisms correspond to a branching diagram that represents part of the history ar some geographic regia on? The ed straint in these gram rather than a A process saunas to be of causal importance. In 1976 and 1978 I suggested a history for the Caribbean land and water that, at a rather general level, is consistent with the present distributions of plants and animals in the region including the Antilles, Central America, northern South America, and the southeastern and southwestern United States. The 1976 proposal, which was ROSEN—GEOLOGICAL HIERARCHIES 637 based largely on a descriptive history by Malfait and Dinkleman (1972) and Tedford (1974), was questioned by Prepi uen) whose claim was that there is no ge a proposal. This ‘anni: was S priticaliy evaluated recently by Hedges (1982) who disagreed with Pregill on the grounds that ample geologic data had been available for some time in support of the Malfait and Dinkleman-Tedford theory (he cited 13 literature sources). Since the time of Hedges' reply to Pregill, I became aware of sev- eral other accounts of Caribbean history based 1977: on a variety of ge movement along major fault zones; Pindell & Dewey, 1982: plate contour matching and pa- leomagmatic data; Kellogg & Bonini, 1982: grav- earthquake data; Sykes et al., of shallow earthquakes and other seismic data; Wadge & Burke, 1983: reconstructing plate mo- tions by closing the Cayman Trough along its bounding transform faults). It is apparent, there- fore, that Pregill was operating with a different set of constraints, namely, that the biota is re- cently distributed by means of dispersal. Hence, no number of geologic accounts or amount of data would be expected fundamentally to alter his position. My constraint is the cladogram and how it describes relationships of taxa and areas. What I propose is to divide the historical geology of the Caribbean into the minimum number of time periods in which different geologic theories agree on the geographic contacts between differ- ent areas and the severing of those contacts. In this way I have covered four main periods span- ning 165 million years. CARIBBEAN HISTORY As I interpret the geological literature the fol- lowing events took place in Caribbean geohis- tory; some prefatory remarks are in order. There are two basic kinds of theories about the history of the Caribbean: 1) those of Salvador and Green (1980) (Figs. 1-9) and Anderson and Schmidt (1983) (Figs. 10-16). These describe a north-south expansion ofthe Gulf-Caribbean re- gion by the latitudinal displacement of North and South America. Both of these are primarily pre- Cenozoic theories that focus principally on the opening of the Gulf of Mexico so that the detailed placement of the Greater Antilles is secondary in importance. 2) All remaining theories, as ex- emplified by (Figs. 17-29) Malfait and Dinkle- 638 ANNALS OF THE MISSOURI BOTANICAL GARDEN panar | — [Vor. 72 «i y q 1 FIGURE 1. The Gulf of Mexico and Caribbean region in the Kimmeridgian (143 Ma) according to Salvador and Green (1980). This, and Figures 2-9, is one of two pre-Cenozoic theories focusing primarily on the opening of the Gulf, and is based on the notion that the Caribbean, like the Gulf, was created by north-south spreading or separation of North and South America. It is distinguished by proposing an origin of Cuba from volcanism along a northern South American transform fault and the gradual shift northward toward the Bahama Bank of Cuba. Other Greater Antillean, and the Lesser Antillean components are also viewed as having in situ volcanic origins, although the Beata Ridge is depicted as a volcanic chain of which Hispaniola is the most northerly element. Black dots represent primary sites of volcanism. Vertical hatching is land; cross-hatching is shallow- water shelves with sedimentary deposits; heavy lines with half arrows are transform faults; and heavy lines with solid arrows are subduction zones. Only the main areas of volcanism are shown by the solid dots. man, and Tedford (see Rosen, 1976), Pindell and Dewey (1982), Sykes et al. (1982) (Figs. 30-33), Wadge and Burke (1983), and Dickinson and Coney (1980) (Fig. 34), describe longitudinal dis- placement events of land and sea floor that are primarily Cenozoic and focus principally on the Caribbean heartland, particularly the origins and movements of the Antillean islands. Readers of these two kinds of historical geologies will notice immediately that the latitudinal expansion the- ories place the origin of the Greater Antilles along the northern edge of South America, not unlike the older theory of Vandel (1972), whereas the longitudinal displacement theories identify the main Antillean islands as part of an ancient is- land arc that connected northern Central Amer- ica (Yucatan) with northwestern South America. Although seemingly contradictory theories of Caribbean history, the consequences of the two theories in terms of the sequence of land con- nections and disconnections are similar. Their common elements include: 1) contact between North and South America via a proto-Greater Antilles; 2) Cenozoic longitudinal displacement of the Antilles; 3) hybridity of Hispaniola and contact of central Hispaniola with eastern Cuba; 4) approximation of eastern Cuba with Florida and western Cuba with northern Central Amer- ica (Yucatan) at some stage; 5) secondary closing of the Central American, South Am and Nicaragua) along a transform fault (= Mo- tagua-Polochic fault and Cayman Trench) with comparable motions ofthe Greater Antillean ele- ts. Although both theories accept a later origin for lower Central America connecting Honduras and Nicaragua with Colombia, in Pindell and Dew- 1985] ROSEN — GEOLOGICAL HIERARCHIES 639 LI $ | LIIIII LI j | TH i Hr jit [LI : l K : H E2. The Gulf of Mexico and the Caribbean region In the Berriasian rin Ma) according to Salvador an ‘Green (1980). At this stage some Sai translocation i stern margin of the Caribbean gion (note the position of Honduras and Dicis E H-N, and compare wit: Buiscauent figures) although a no dioc ecu expansion is still considered primary. Pe IGURE 3. The Gulf of Mexico and the Caribbean region in the Berremian (117 Ma) according to Salvador and Green (1980). 640 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 E The Gulf of Mexico and Caribbean region in the Cenomanian (93 Ma) according to Salvador and e (1980). Jamaica (J) is depicted as a consequence of submarine volcanism and Hispaniola is shown just to the east of Jamaica with a small subaerial component. UN NN al | ú X | v i | FiGURE 5. The Gulf of Mexico and the Caribbean region in the Maastrichtian (70 Ma) according to Salvador and Green (1980). The outlines of all of the Greater Antillean islands are now shown, with the western, subaerial part of Cuba (C) in contact with the Yucatan (Y) platform 1985] ROSEN — GEOLOGICAL HIERARCHIES oo sa . x, 4 L1 ses sss sas as as FIGURE 6. The Gulf of Mexico and the Caribbean region in the Paleocene (55 Ma) according to Salvador and Green (1980). The Beata and Aves Ridges (BR and AR respectively) are now emplaced and the Caribbean has undergone its maximum north-south expansion. A volcanic chain now extends between northwestern South America and the ultimate site of Honduras and Nicaragua. ey's (1982) longitudinal displacement theory Honduras and Nicaragua (and by implication of polar position data, also Jamaica) had, along with Panama and Costa Rica, an older origin in the eastern Pacific and were emplaced as the proto- Caribbean/eastern Pacific sea floor was sucked into the Caribbean heartland. Thus, both kinds of theories include the potential for Antillean juxtaposition with northern Central America (Yucatan) at some time in their histories. The largest difference between them is how far east one should situate an island arc that joins north- ern Central America (Yucatan) with Colombia and in the ages and rates of motion of the An- tillean components. But there seems little reason to argue that a single ancestral biota might have occupied large parts of what now are a series of isolated islands, thus providing major opportu- nities for biotic differentiation in a pattern of land connections and disconnections very much alike for both latitudinal and longitudinal dis- placement theories. From the standpoint of bio- geography, then, the different theories disagree in some details and in focus, but agree in ways that concern biohistory. HISTORICAL SUMMARY 150-165 MA At this early stage North and South America were closely approximated, separated in what is now the Gulf region by the northern Central America (Yucatan) block, which later became the principal component of northern Central America (Yucatan). The Bahama platform lying due ESE of southeastern North America existed at this time. No elements of the Antilles would be recognizable at this period. Northern Central America (Yucatan), including o d Guatemala, Honduras, and Nicaragua, would, at this stage, be Pacific land (the Chortis SER ee to the southwest of what was to become Mexic Africa, North America, and South America were all still in contact with a small piece of Africa lying athwart the southeastern segment of the already formed Appalachian chain and including Florida and the northern Bahamas. Major fault zones were beginning to form 1) to the north and south of northern Central America (Yucatan) in the Gulf region, 2) to the east, west, and south of the Bahama platform, the eastern part forming 642 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 | | '| vl | | | | BRL ORE, lll il FIGURE 7. The Gulf of Mexico and the Caribbean region in the Middle Eocene (42 Ma) according to Salvador and Green (1980). A spreading center has now appeared in the Cayman Trough (CT), accentuating east-west translocation along the faults to the north and south of the Caribbean basin and the latter is being compressed by the motion of North and South America toward one another. In this model only Hispaniola (HI), Puerto Rico (PR), two sections of Cuba and the Isle of Pines are considered to be subaerial. The proto-Lesser Antilles se ni have now appeared on the eastern border of the Aves Ridge. x i Ë FEN | mur a ~ pee 4", D p s — í gesaanlB E CTT 5 Q a< ert i Elm TI ` Te ii í DT II CENTS a. esa | x fr s Í li 1985] i I! II t ia | c ROSEN —GEOLOGICAL HIERARCHIES E ezass HM 643 in uz Ae = sia HE oo 2 Com Ë" £a Rš "aea. ansar a| FILES A Ud il m Bi LI oe FIGURE 9. The Gulf of Mexico and ie Caribbean region in the late Miocene (7 Ma) according to Salvador and Green (1980). The Caribbean bas n has undergone nearly its maximum compression in relation to the Recent configuration, Cuba has uei: into three sections, Hispaniola into two, and Jamaica andi a major al. part of the Yucatan is subaeri a northern segment of the proto-mid-Atlantic idge, 3) a southern segment of the proto-mid- Atlantic ridge between northeastern South America and the proto-Gulf of Guinea region of Africa, and 4) a major part ofthe western Central American Trench off the west coast of North America and South America. 95 MA North and South America have now separated substantially, opening up the Gulf of Mexico and an area to the south that was later occupied by Caribbean sea floor. Northern Central America (Yucatan) has now become joined to Mexico, and the Chortis block has moved east to a po- sition south of Mexico and west of northern Cen- tral America (Yucatan), the area of contact rep- resenting pul Nur of the Motagua-Polochic transform fault (= Cayman Trough). A proto- Antilles island arc, of volcanic origin, has formed Central America (Yucatan) and ican trench in this region has jumped to the east of the arc to form the leading edge of a subduc- tion zone that will enable the proto-Antilles to override the proto-Caribbean sea floor. The western end ofthis island arc moved north, where; colliding with northern Central America (Yu- catan); Jamaica, the Caymans, and southwestern Hispaniola were sheared off from western Cuba. Thus, Jamaica, the Caymans, southwestern His- paniola, and western Cuba all accreted respec- tively to the southern and eastern parts of north- URE 8. The Gulf of Mexico and the Caribbean region in the early Miocene (21 Ma) according to Salvador S migrated viene under the Yucatan (Nuclear Central Aun pen mim rinde HERI ‘of Ts pepe Trou 644 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 (| t ` S HA Ky N \ ‘see q Th = | ry ^ s: A yl Ma: š: UN : l d Š V 3 || EEE i min according to uec. and Schmidt (1983). Vertical hatching is land, cross- hatching i is shallow-water shelf, and oblique hatching is a reconstructed curvilinear belt o Paleozoic rocks that show lithologic and i ions of Mexico and Caribbean by the north-south db ane of North and South America (see Figs. | 1-17). ern Central America (Yucatan). The attenuated tacting the Bahama platform along its north- Bahama platform made a southeastward ap- eastern margin and nuclear Hispaniola to the proach to the northern South America trench, south. Southwestern Hispaniola, the Caymans, and the mid-Atlantic ridge is now well developed and Jamaica have begun an eastward migration between Africa and North America, the former away from the southern edge of northern Central having left behind Florida and the small Appa- America (Yucatan), moving along the proto- lachian segment mentioned above. Cayman trough, which has a complex structure including pieces to the north, west, and south of Cuba and a north-south arm southeast of the 65 MA North and South America have separated fur- eastern edge of the Caribbean plate. The Aves ther along a modern north-south axis and the Ridge or proto-Lesser Antilles has formed just proto-Antillean arc had begun to stream east- behind this leading plate margin. The Chortis ward into the Caribbean heartland opened up by block has also begun a southeastward migration the separation of North and South America. The along the Motagua-Polochic fault, carrying it Oriente of Cuba is now situated to the east, con- closer to northern Central America (Yucatan), FiGURE 12. The Gulf of Mexico and Caribbean region in the Middle - urassic (150 Ma) according to ndn on and Schmidi (1983). As a result of latitudinal displacement of North and South America the proto-Greater Antilles are depicted as moving north away from South America. 1985] ROSEN —GEOLOGICAL HIERARCHIES 645 P M Ell. TheGulfof Mexico and Caribbean region in the Middle : urassic ee Ma) peel Anderson I Da ? t ah Schmidt (1983). The proto-mid-Atlantic ridge can be seen between south-eas ort and north- western Africa as a three-step inferred fault of broken lines. Vertical hatching 1S Eu cross- ate is shallow- water shelf, unbroken heavy lines is the original position of the Mojave-Sonora megashear that bounds the Yaqui microplate to the north (see Fig. 10), broken heavy lines are w la faults ~} antl H I LANE a TT inl 7 l| 646 Ji FiGURE 13. The Gulf of Mexico and Caribbean re- gion in the Kimmeridgian (ca. 140 Ma) according to Anderson and Schmidt (1983). Compare with Figure 1. ANNALS OF THE MISSOURI BOTANICAL GARDEN | INIM | m Il Ill Ill TINI Y and Bahama platforms. Compare with Figure 4. [Vor. 72 | J | V W r The Gulf of eae oe Caribbean re- t Cretaceous (ca. 70 Ma) according to (1983). dier dat displace- RE 15. on in pot lates dnd and Schmidt ment is still continuing. and lower Central America has moved in from the eastern Pacific to suture with the Chortis block. PRESENT TO 9 MA During this pre-modern epoch in Caribbean history nuclear Hispaniola continued its east- ward movement, separating from the Oriente of Cuba and being joined by southwestern Hispan- iola (about 9 Ma) and picking up a small southern piece of the Bahama platform along its northern edge (about 3 Ma). Jamaica and the Caymans have trailed behind maintaining a steady dis- tance from southwestern Hispaniola, so that, al- though at one time connected by contiguous land to Cuba and Hispaniola through northern Cen- tral America (Yucatan), the former islands have been isolated fragments for most of Caribbean geohistory (during which, the White Limestone Group of central Jamaica was continuously sub- aerial, Robinson, 1976; Robinson et al., 1977). Central America now occupies the site, formerly the province of the proto-Greater Antilles, and is joined to the precursor lands of western Co- lombia (the area ofthe Rio Cauca and Magdalena basins). The Cayman Trough has a complex con- figuration including a spreading center between its northern and southern slip-fault components. 1985] ROSEN—GEOLOGICAL HIERARCHIES 647 Í FiGure 18. The Gulf of Mexico and the Caribbean region prior to the separation of Africa and South America (ca. 150 Ma) according to Pindell and Dewey sii The heavy east-west lines across Mexico are as of thrusting that separate Mexico from south- P ML A (| we aan North America at roughly the level of the Rio L^ II = j f, Grande and that subdivide Mexico into four discrete ul zones E 16. _ The Gulf pn Mexico and the Caribbean connects with the subduction zone along the regior in t and Schmidt . Eod with Figures ri dh Q leading eastern edge of the Caribbean plate be- Latitudinal expansion has ceased and northern Central hind which the modern Lesser Antilles have America (Honduras and Nicaragua) remain isolated emerged as chains of volcanic stepping-stones from northwestern South America. from the precursor Aves Ridge. With the ulti- mate disappearance of the Cauca-Magdalena, The southern component is now continuous with transgressing seaway in northwestern Colombia, the active Motagua-Polochic left-lateral trans- a new continuous land connection was made be- form fault, which connects with the again un- tween North and South America. broken western Central Americas Trench. northern component of the major Antillean HISTORICAL SUMMARY trench system continues eastward between the Bahama platform and the formerly Bahamian section of northern Hispaniola and ultimately a | I alll { To the extent that the foregoing account is cor- rect, one can say that the Caribbean land and FiGuURE 19. The Gulf of Mexico and Caribbean re- Figure 17. The Gulf of Mexico and the Caribbean gion in the Kimmeridgian (140 Ma) according to Pindell region in late Pangaean time (ca. 165 Ma) as and Dewey (1982). The Yucatan block has rotated into to Pindell and Dewey (1982). Yucatan (Northern Cen position and sutured to An Mexico. North- tral America, Y) is depicted here as a separate micro- ern Central rica (Honduras and Nicaragua, H-N) plate between North and South America is depicted as already sutured i uod Mexico. 648 FiGurE 20. The Gulf of Mexico and Caribbean re- gion at the time of formation of a proto-Antillean ar- chipelago (PA) at 125 Ma, according to Pindell and Dewey (1982). The Bahama platform (B) can be seen extending in an ESE direction toward the proto-mid- Atlantic ridge. water had a complex history of connections be- tween different geographic features being made and then broken and then reformed in another configuration. The history includes eight events of fragmentation and seven of hybridization. This is summarized below in the series of included maps that show bn vicariant opportunities for the biota are abo events that e opportunities for mixing of biotas. The geology of Hispaniola provides per- haps the most dramatic example of these two aspects of geohistory since it is alleged to have had a history of prior connection with northern Central America (Yucatan), Cuba, Jamaica, the FiGURE21. The Gulf of Mexico and the Caribbean region at the onset of separation of Africa and South America at 110 Ma, according to Pindell and Dewey (1982). The proto-Antillean archipelago has already moved slightly to the east of its position in the pre- ceding figure ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 gs into the Caribbean region at about 95 Ma, ETS to Pindell and Dewey (1982). Bahamas and an Pay. procesi of inamentanon from other proto-An p jor segments that behaved ind r most of their reconstructed history. But SIE the most interesting part of Caribbean geohistory is that involving the Chortis block and lower Cen- tral America, which appear to have been Pacific land accreted into the eastern margin of the Pa- cific borderlands, thus recalling some elements of the Pacifica concept of Nur and Ben-Avraham (1977). Another recent study has further docu- mented the incorporation of allochthonous ter- ranes into western North America (Minckley et al., in press). If part of the same phenomenon, FiGURE23. The Gulf of Mexico and the Caribbean region as the proto-Antillean archipelago at 80 Ma, according to Pindell and Dewey (1982), moves ENE toward the Bahama platform and a contact with the southeastern margin of the Yucatan. Note also the re- construction of lower Central America as an eastern Pacific a. oy near the western boundary of the Caribbean pla 1985] ROSEN—GEOLOGICAL HIERARCHIES la FiGURE 24. The Gulf of Mexico and the Caribbean region: as s the proto- -Antillean archipelago begins to dif- fthe Greater Antilles at 65 Ma : according to Pindell and Dewey T. Jamaica yid igi Hispaniola (SWHI), iris o (PR), and Cuba (C) that lies just north of th in is of the yos Trough (jagged line and CHI). the history of Central America might well be shared with that of the exotic terranes along the western margins of North and South America, including perhaps the Carnegie Ridge system (the Galapagos and Cocos Islands) (see Rosen, 1976). DERIVING GEOLOGICAL AREA CLADOGRAMS With a history this complex it must be obvious that no simple branching diagram can exactly express all the implied relationships nor, because of hybridizations, is a completely resolved FiGURE25. The Gulf of Mexico and the €— eat rthern A cm to es nd Dewey (1982 FIGURE 26. The Gulf of Mexico and the Caribbean many of the Antillean been differentiated, the Cayman Trough ha as be ecome active and northern Central America has taona UlUV Tl Motag ua o fault and central Hi t Cuba according to Pindell and cien (1982). branching diagram possible. It is, therefore, my conclusion that most profit is to be gained from isolating parts of Caribbean history that repre- sent stages in the evolution of land and water and therefore stages during which today's com- plex biota can be understood. There is evidence land iguanas, decapod crustaceans, and probably also cichlid fishes (see Rosen, 1976). Regarding the West Indian butterfly fauna, Brown (1978) has written: “It is possible that the original fauna The Gulf of Mexico and the Caribbean when, according to Pindell and Dewe ey (1982), a spreading center has appeared in the FIGURE 27. region at about 21 M ayman of Jamaica, southwestern Hispaniola, and the disjunc- tion of central Hispaniola and Cuba. ANNALS OF THE MISSOURI BOTANICAL GARDEN URE28. The Gulf of Mexico and the Caribbean n, according to Pindell and made its closest Cu approach to the southwestern tip of the Bahama plat- of the region had its roots in Africa at the close of the Mesozoic Era. Very little of this fauna is left but there are some indications of it." Other taxa seem to reflect events in the less remote [Vor. 72 FiGunE 29. The Gulf of Mexico and present plate boundaries and land areas in the Caribbean region from Pindell and Dewey (1982). periods of North and Middle American biohis- tory (Rosen, 1978). It would, therefore, be most helpful to be able to compare biological area cladograms with specific periods in geohistory rather than, as was done before (Rosen, 1976), to simply identify components the modern biota that are consilient with some features of today's complex geography. Thus one must introduce a Y 1 e AM 24 Gulf of Mexico PLATE Mesco ` uie f 3 — ae O Atlontic TONS aM. Q : o yor™ 3 : D Y(09Q" — ed be 2», ceon i, m e AES 10 o Ta (as Kae, ALT rrp. Se! oni LF EER Mig E hn. a) Renal an" TA Uc es i € S. Hoiti 9. gem. bana Q # A pducred T a e ea v PLATE d A Sens 38 Mo. ` Central ` 9 y» ee” € © i ae 24 IT YW Mh: al Venezuela mplex incor- t Am with the Chortis block by the early Miocene prior to an isthmian connection between North and South America. 1985] ROSEN—GEOLOGICAL HIERARCHIES NORTH AMERICAN Son, PLATE GD = ^o / 6, Ong Ationtk Oceon o9? e o SC uiu EK: Pacii "i EOCENE 48 MA Q " SOUTH i AMERICAN PLATE DA MUR A nont Oc eo^ sayy SOUTH AMERICAN ais PLATE d. Cocos CARIBBEAN FLATE PLATE _ r LATE MIOCENE SOUTH AMERICAN 7 MA PLATE FIGURES 31-33.—31. The eastern Pacific and Carib- bean in mid-Eocene times (ca. 48 Ma) according to Sykes et al. (1982). An intimate association between Jamaica, Cuba, and Hispaniola is depicted.— 32. The Caribbean plate in n Miocene times (ca. 20 Ma), Sykes et al. (1982), showing the formation ofa Cayman Trench spreading center on the site of the older Cayman Ridge, the continued association of cen- tral Hispaniola with eastern Cuba, and a now com- pleted Chortis block/Isthmian Bridge. —33. The Ca- ration in late (1982), showing the still — ids LM Bridge with northwestern South America a e movement of Cuba eastward away from Pa The Eocene age of this depiction is consistent with the age determi- nations of Wadge and Burke (1983) who also a a rate of plate motion of about 4 cm per 1 , cladistic constraint so that £g dia- grams represent present areas iof endemism that can be shown to be congruent with a branching diagram representing a specific part of Caribbean geohistory. The branching sequence must also agree with the time constraints (relative times) of accretion and fragmentation. da-Bahama bank where arc activity is extin- guished. CLADISTIC ANALYSIS From the standpoint of Antillean history, the simplest place to begin is at that point 65 Ma when a proto-Antillean archipelago connected northern Central America (Yucatan) with north- western South America (Figs. 33-44). This his- tory can be divided into five components that will take us up to the modern period. In the earliest configuration (Fig. 33), the archipelago consists of 1) western Cuba and its sheared off fragments (Jamaica and southwestern Hispan- iola) that are closely associated with northern gua) is either Pacific land unconnected to north- ern Central America (Yucatan) or is sutured to western Mexico considerably to the north of the proto-Antillean archipelago. Branching diagrams representing the complex history of the Caribbean must take into account both accretionary events, of which I identify sev- en since the main continental separation began about 165 Ma ago, and fragmentation events, of which I identify eight, including the initial con- tinental separation. Fragmentation events (roughly in order of de- creasing age 1. North America-Africa 2. North America-South America 3. Southern Mexico-North America 652 CA NCA JAM SWHISP WCUB ECUB CHISP A FIGURE 35. A diagrammatic consensus dip ai 2 mid-Cenozoic relations among s nents of the Caribbean heartland ud. a ba aa dia- gram that summarizes these relations. South America-Africa Southern Mexico (Yucatan)—western Cuba, Jamaica, southwestern Hispaniola Eastern Cuba-Bahamas Eastern Cuba-central Hispaniola Hispaniolan Bahamas (fragment)-southern ahamas > mW E Accretionary events (roughly in order of decreas- ing age) l. s Cuba, Jamaica, southwest- ern Hispa nduras, s (northern Costa Rica)- western to southwestern Mexico, and even- tually to northern Central America (Yucatan) Western Cuba-eastern Cuba Eastern Cuba-central Hispaniola Central Hispaniola-southwestern Hispaniola Southern Bahamas (fragment)—central His- paniola m pode ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 7. Southeastern Costa Rica, Panama-northern Central America (Yucatan), northwestern South America For the accretionary events there is an alter- native model, differing in details of island se- quence, and that is the one proposed by Carey (1958), recently discussed and illustrated by Na- gel (1971, fig. 15) in which, in sequence from north to south, Jamaica, Puerto Rico, Hispan- iola, and Cuba were a part of southern Mexico prior to their translation and rotation to a more easterly position. In this model the Bahamas lay between the tip of Florida and northern South America with an unspecified amount of inter- vening land, and Honduras-Nicaragua (the Chortis block), and the Yucatan (labelled Gua- temala in Nagel's figure) were narrowly joined and both a part of the North American gulf coast. Also, in this model, it was the Lesser Antilles that formed an island arc South America in the early Mesozoic. From the standpoint of its relation to biogeography even this seemingly inverted model of land-area re- lationships yields the same general expectation of the biological relationships of the Caribbean subregions. The general features of the relation between n tectonics and biogeography were discusse ost recently by Durham (1985), but primarily in connection with the model pro- posed by Sykes et al. (1982). If we adopt the position that land areas formed of mutually ex- otic terranes through accretion, might exhibit taxic area relationships when later fragmented, a consequential inference is that accretionary events might be followed by biotic transfer across comparison with biological area cladograms must incorporate both accreted and fragmented areas. Since a suture zone might serve to isolate orig- inal and transferred biotic elements because of increasing geographic disruption during the later phases of suturing, my geologic area cladograms adopt the potentially most informative interpre- tation by incorporating sutured areas (that have not secondarily fragmented) on an equal basis e based on both n and accretion, land ybridizations would be rep ted either as re- ticulations (lines joining two or more branches) 1985] or, as in the case of biological hybridizations, as unresolved b gthe parent taxa and the hybrid. This is s the conservative proce- dure adopted here. One might suppose that land hybridization (e.g., the ms block with south- with we a sharing of biotas by the joined fragments, but that supposition requires the subsidiary idea, probably correct in some instances but not oth- ers, that the suture zone presents no obstacle to biotic mixing. At least some suture zones seem perfectly hospitable to organisms, as in the case of the Chortis block suture zone (referred to as the Polochic-Motagua transform fault) where persistent, small-scale seismic activity, and the formation of a major down-dropped block of land (e.g., Lago Isabal) have not been inimical to the survival of faunal elements within the re- Motagua-Polochic synclinal depression]. Other organisms, such as the swamp eels of the genera Ophisternon and Synbranchus, seem unusually sensitive to this fault line (see Rosen & Green- wood, 1975), although species of Ophisternon occur in the Yucatan and in Cuba, indicating that distributional boundaries cannot be assumed from eta ata. Even though one can identify at least 16 geo- Won areas involving Caribbean history (if the continental regions are included), I have chosen only eight of these to illustrate a method whereby geological data might be presented for compar- ison with biological data. The chief objective 1s to be able to relate a given biological distribution í. to a specified period in Caribbean geo- hist ene area cladogram representing these relationships can be written as shown in Figures . Th Hispaniola, western Cuba]. convincing case we will take the most conser- vative stance and derive from this general clado- gram all possible completely resolved four-taxon statements, of which there are six shown in Fig- ure 36. Many more three-taxon statements are derivable, but three-taxon statements have the disadvantage of representing relatively trivial statements of relationship and therefore requir- ing much more comparative data to demonstrate general congruence. For example, for three-area ROSEN — GEOLOGICAL HIERARCHIES 653 Resolved four-t tat ts derivable from A: X ` oe A WU X NCA WCUB ECUB CHISP SWHISP WCUB ECUB — CHIsP C Cuba; ECUB, eastern Cuba; CHISP, central or nuclear Hispaniola. d qi there are only three possible solu- s: A-B-C, A-C-B, B-A-C and two replications dts one of these patterns would yield a probabilit y possible solutions to a four-taxon system (Nelson & Platnick, 1981; Rosen, 1978) and two repli- cations of a given pattern would yield a proba- bility of once in every 225 trials (15 x 15 x 15) that the congruence was random. In the next geographic configuration in which Central America). The general cladogram (Fig. 37) representing these relationships includes one land hybridization and three vicariant events and 654 W CUB NCA ECUB o c S HISP JAM SW HISP CA NCA JAM SWHISP WCUB ECUB CHISP B FiGURE 37. A diagrammatic consensus map of the late ende relations among som mponents of the Caribbean heartland and the branching diagram that summarizes these relations. includes a pentachotomy. From this general statement only a single resolved four-taxon state- ment is derivable (Fig. 38). As eastward movement of the Antilles contin- ues, central Hispaniola separates from eastern Cuba but western Cuba rived but a single resolved four-taxon statement (Fig. 40). Continued movement of the Caribbean plate brings southwestern Hispaniola and central Hispaniola into contact and, although no new mainland relationships are being established, the Chortis block continues to slide eastward against northern Central America (Yucatan) on the Mo- tagua-Polochic fault so that at least new contig- uous land-area-relationships are being estab- lished (Fig. 41). Here, too, the general cladogram includes a pentachotomy but gives rise to three resolved four-area cladograms (Fig. 42). In the dern configuration a new land contact is es- tablished between southern Bahamas and His- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 CA NCA JAM SWHISP WCUB ECUB CHISP Resolved four-taxon statements derivable from B: CA NCA ECUB C HISP FiGURE 38. The branching diagram as in Figure 37 and the single, completely resolved four-taxon, four- i dcr aes derived from it: abbreviations as in WCUB DO ° HISP SW HISP WCUB ECUB JAM SWHISP CHISP CA c FiGURE 39. A diagrammatic consensus map of late Cenozoic relations among some main components of the Caribbean heartland and a branching diagram that summarizes these relations. 1985] WCUB ECUB JAM SWHISP | CHISP Resolved four-taxon statements derivable from C : ECUB FiGURE 40. The branching diagram as in Figure 39 and the simple, completely resolved, four-taxon, four- area statement derived from it: abbreviations as in Figure 36 paniola, and the Chortis block continues to move eastward along the Motagua-Polochic fault. The general cladogram incorporates two dichoto- mies, one trichotomy, and one tetrachotomy (Fig. 43), but allows derivation of seven resolved four- area statements. The sum of all of the proposed historical geo- logies leads to a number of predictions: —_ . Hispaniola will have a mixture of elements, some related to the Bahamas, some to eastern Cuba, some to Jamaica, others to north- ern Central America (Yucatan). Some of the Antillean S p be parts of natural groups s in North and South America and Afric . Honduras, iE and perhaps western Costa Rica, will share related taxa in southern and southwestern Mexico, and some of these should be parts of more inclusive taxa that include the Antilles and South America. The most inclusive taxa should have repre- sentatives in all the foregoing areas and in Africa. Panama and eastern Costa Rica, by and large, should share species or species groups not unique to themselves but shared with South tə WwW » m ROSEN — GEOLOGICAL HIERARCHIES WCUB E ECUB oe NCA SS HISP um SW HISP CA NCA JAM | S8AH WCUB ECUB SWHISP CHISP D FiGURE 41. A diagrammatic consensus map of late iacu relations among some main components of e Caribbean heartland and the branching diagram di summarizes these relations. JAM S3AH WCUB ECUB SWHISP CHISP Resolved four-taxon statements derivable from D : CA WCUB ECUB CA SWHISP CHISP i MA WCUB ECUB SWHISP CHISP FiGURE 42. The branching diagram as in Figure 41 and the three completely resolved, four-taxon, four- area statements derived from it: abbreviations as in Figure 36. 656 ECUB SWHISP CHISP SBAH E FiGuRE 43. A d map of the modern relations among s mponents of the Carib- bean heartland a a (esqon s ng diagram summarizing these A (SBAH, southern Bahamas). America and southern Mexico and northern Central America (Yucatan). Endemic sister taxa will be found in eastern and western Cuba on each side of the uplifted cap marine deposits at about 79? west longitu EN Whether the details of this geohistory are true or false is not at issue here. What is at stake is the possibility that a geohistory of this complex- ity might well have occurred, meaning that there might be 19 possible four-area cladograms with which biological area cladograms can be con- gruent. In other deg: 19 different cladistic sos lutionsto reflect the dramatic events in Caribbean geohis- to ry. In the recent past some authors (Rosen, 1976; Tolson, 1982) have assumed that failure to dis- dispersal may account for the lack of congruence. Tolson (1982), for example, assumed that the occurrence on Hispaniola of three boid snakes each with a different relationship to those on Jamaica, Puerto Rico, Cuba, and the Bahamas meant that dispersal might have occurred in or- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 ECUB SWHISP CHISP SBAH E Resolved four-taxon statements derivable from E: SWHISP = CHISP W CUB ECUB wCUB ECUB SWHISP SBAH . Sg CA NCA SWHISP SBAH XZ CA NCA C HISP S BAH FIGURE 44. The branching diagram as in Figure 43 and ie seven completely resolved four-taxon, four- area statements derived from it: abbreviations as in Figures 36 and 43. der to permit the coexistence on one island of each of these endemics. But the complex history of Hispaniola should lead one to expect complex relationships of endemic Hispaniolan taxa. e fact that several different historical geo- logies of the Caribbean specify similar kinds of complexities and support an origin ofthe Antilles from an ancient island arc between South Amer- ica and northern Central America (Yucatan) is reason enough to suppose that land hybridization as well as fragmentation will yield a picture of unique endemism and mixed biotas in which endemic taxa from formerly isolated land frag- ments have been brought together in a way that conflicts with simple sister-group analysis. e is a message here for all interested in biogeography and it is not that dispersal never occurs, but that theories of dispersal to explain m w edd in SED (Nelson & Platnick, 1981) 1985] Although geology has yet to fill in the details associated with the above five general clado- grams, biology has a constructive role here be- cause, by discovering congruent four-taxon area cladograms like the 19 derived above, it can in- dependently support any one of the five general area cladograms and therefore details of geohis- tory that are still out of reach of geologists' data. If, in fact, there is a causal relation between an independent variable of geohistory and a depen- dent one of biohistory it should work just that way and biology should be at least as infi iv to the geologists as geology has been assumed to be for biologists in the past. Clearly biologists have always been ready to make that judgment since the last 150 years of biogeography has been based on the assumption that modern biotic dis- tributions have been achieved through dispersal, signifying to geologists their belief that the rel- evant part of geohistory has been one of stabil- ism—a view that could have been challenged only by a theory of plate tectonics or a biological the- ory of general area congruence as proposed by Leon Croizat (1958, 1962). In such detailed historical summaries of the geology of a region there is an unfortunate air of real knowledge, but Wey] (1980) has warned that, not only does the problem of reconstruction be- come increasingly difficult as we go back in time, but that the only criteria we have to go by are the data of prehistoric magmatism and the struc- ture of the uppermost part of the crust, while the seismic and gravimetric data that are so impor- tant either are lacking or are poorly known rel- ative to what is needed for establishing a detailed historical picture. Even though one of the most detailed recent historical geologies (Sykes et al., 1982) is based on the paleoseismic data required by Weyl, bi- ological data from the living and fossil biotas might have still more to say about Caribbean history. For example, if each of the 19 resolved four-taxon area cladograms were corroborated O and to abandon the search for cladistic area-con- gruence in favor of some a priori notion that all distributions might be explained by guesswork liberally laced with dispersalist intuition is to ensure that future generations of biogeographers will regard such proposals lightly. The fact that ROSEN—GEOLOGICAL HIERARCHIES 657 there are 19 resolved four-area cladograms that need to be corroborated might seem somewhat daunting to someone venturing for the first time into cladistic biogeography because it will, in- deed, require a stupendous multidisciplinary ef- fort to resolve decisive patterns for the region — and more still if one adds Mexico, North and South America, and Africa. But I presume that if the problem were less difficult there would have been fewer attempts t scenarios about which pe no two scenario-writers can exactly agree, or provide that decisive constraint that shows one or another scenario to be flawed and scientifically unacceptable. But the history of today's biogeography has been characterized by the casual, almost offhand, manner of the scenario-writers that populate the field. Never- theless, the temptation to propose imprecise, non- cladistic, solutions to the problem of Caribbean history is great because of the abundance of en- demic taxa that link certain areas. Patterson (1981: 458) complained that these seemingly re- assuring biogeographic data add up to a phenetic concept that cannot lift biogeography out of its present narrative phase But all problems in comparative biology begin with a data gathering phase that un- known amounts of noise and signal. Cladistics identifies the signal. The need for cladistic vi- cariance analysis is underscored by questions such as those following. How significant is the co-occurrence on His- paniola of boid snakes in relation to a geological theory of a threefold origin for Hispaniola? How significant is the occurrence of swamp eels of the genus Ophisternon restricted to Trin- idad, Cuba, and northern Central America north of the Chortis block? How significant are the occurrences of Carl- hubbsia and Quintana plus Girardinus in north- ern Central America (Yucatan) and Cuba re- spectively (Rosen & Bailey, 1959) in relation to eories of ancient oe between western Cuba and the Yuc How si E cuit IS ae occurrence of related taxa of the poeciliid genus Gambusia in Cuba, the Bahamas, Hispaniola, and northern Central America (Yucatan) in relation to mid-Cenozoic interconnections of these areas? These distributions, linking disjunct or hybrid areas, appeal to our sense of discovery. A cla- distic analysis of areas will help us decide wheth- er the distributional components that drew our 658 attention specify a general problem. I predict that such analysis, on a grand scale involving many different kinds of organisms, will corroborate some, if not all, of the five general geological area- cladograms that describe the longitudinal dis- placement theories of Caribbean history of Pin- dell and Dewey (1982), Sykes et al. (1982), and Wadge and Burke (1983). But, if such corrobo- ration is not forthcoming, as biologists we are bound by the message of biological data in de- scribing a biotic history of those geographic areas regardless of any possible conflicts with geologic theory. LITERATURE CITED eee T. H. & V. A. ScHMIDT. 1983. The evo- lution of Middle A d the Gulf of Mexico- Caribbean Sea region during Mesozoic time. Bull. Geol. Soc. Amer. 94: 941-966. BiRNIE, R. W. 1977. The Guatemalan earthquake — Fe 4, 1976. Explorers J. 55: 97-118 BRiGGs, J. 1984. Freshwater fishes and biogeography of Central America and the Antilles. Syst. Zool. 33: 428-435. Brown, F. M. 1978. The origins of the West Indian butterfly fauna. Acad. Nat. Sci. Philadelphia Spe- = Publ. 13: 5-31. CAREY, S. W. 1958. A tectonic approach to conti- nental drift. Pp. ipsis in Lm Drift, A posium. Hobart 1958. Panbiogeography. “Published by the author, Carac 1962 1964]. Spac ce, Time, Form: The Bio- EN Synthesis. Published by the author, Cara- Destmcros P. J. 1957. Zoogeography, The Geo- ical Distribution of Animals. John Wiley and eim Inc., New York. 1965. Biogeography of the Southern End of Cam Ea North Atlantic Ocean. Louisiana State Univ., Bat- on Rouge. DURHAM, J. W. 1985. Movement of the Caribbean plate and its importance for biogeography. Geol- ogy HEDGES, B. bean biogeography: impli- . 19 Carib cations of recent plate tectonic studies. Syst. Zool. 31: 518-522. KELLOGG, J. N. & W. E. Bonini. 1982. Subduction of the Caribbean plate and basement uplifts in the overriding South American plate. Tectonics 1: 251— 276. MALFAIT, B. T. & M. G. DINKLEMAN. 1972. Circum- Caribbean tectonic and igneous activity and the evolution of the Caribbean plate. Bull. Geol. Soc. Amer. 83: 251-272. McDowarir, R. M. 1971. : Fishes of the e Aplo- chitonidae. J. Roy. Soc. New Zealand 1 : 31-52. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 MINCKLEY, W. L., D. HENDRICKSON & C. E. Bond. In press. Geography of Western North American Freshwater Fishes: Description and Relations to Intracontinental Tectonism. John Wiley and Sons, l. Systematics = Biogeography: peus and Vicariance. Colu a Univ. Press, o NUR, a & F. BEN a hua sassa 1977. “Lost Pacifica Continent.” Nature 270(5632): 41-43 PATTERSON, C. 1981. Methods of rp 446-489 in G. Nelson & D. E. Rosen (editors), Vicariance Biogeography: A Critique. Columbia Univ. Press, New York. PINDELL, J. & J. F. Dewey. 1982. Permo-Triassic reconstruction of western Pangea and the evolu- tion of the Gulf t Mexico/Caribbean region. Tec- . K. e An appraisal of the vicariance hypothesis of Caribbean biogeography and its n plication to West I ol. 30: 147-155. ROBINSON, E. 1976. Minerals and ans tectonics in A vicariance model o

w my Oo nN a wo > nN w + FiGuRE15. Finding congruent historical aspects of ecological pe (a) and (e) are cladograms for asso ocia ated with another group [(b) and group in (b), and (h phylogenetic relationships of the group in (e) to the group in (f) (redrawn from Brooks, 1981b). tative approach, at least implicitly, available for studies of historical ecology. III. Under what conditions did the ecological life history traits that we observe today emerge? This category provides historical ecological contributions to questions concerning the evo- lutionary effects of competition and the occur- rence of adaptive radiations. It allows possible distinction between factors responsible for the maintenance of traits seen today and factors re- sponsible for the emergence of those traits in the 670 Elipesurus Potamotrygon r= o. 1 Š F H R MO C Y MA FiGuRE 16. Cladogram . phylogenetic re- een among eight spe of neotropical fresh- water stingrays based on the pi Hylogeneric relationships of their helminth ae (redrawn from Brooks, 1981b) first place (see also Ross, 1972b). Evolutionary ecology, by virtue of its reliance on indirect es- timates of history, must rely on a form of uni- formitarianism — if competition maintains traits today, it was responsible for their origin in the past. Historical analysis also allows one to compare the relative degree of parallel (sometimes called adaptive) evolution exhibited by different groups. ra rasites, id rampie i are supposed to be par- (Price, 1977, 1980). And yet, recent studies of various groups of para- sitic worms (Brooks & Glen, 1982; Brooks et al., 1985a, 1985b) have documented consistency in- dices of 0.74 to 0.95, suggesting very low levels of parallel evolution. Lastly, by preparing a cladogram for a given group of organisms based on non-ecological traits (i.e., structural rather than functional traits), then mapping ecological diversification onto the tree (Farris optimization: Farris, 1970), one can de- termine whether ecological diversification pre- cedes or lags behind morphological diversifica- tion (see e.g., Ross, 1972a, 1972b; Resh et al., 1976; Morse & White, 1979). Such analyses can have startling effects. Brooks et al. (1985b) re- cently finished a family-level analysis of dige- netic trematodes, based on 211 morphologica characters. When all data for six different cate- gories of ecological diversification were mapped onto the cladogram, only 26% of the tree was resolved. Ths historical picture is one of ecolog- ical div agging behind diversification. Ross (1972b) estimated that ı one ecological shift occurs for approximately 30 spe- ciation events in phytophagous insects. Boucot (1983b) has suggested that this ecological con- servatism is a general picture in the fossil record. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Association Origin. of Coevolution Colonization I- < plesiomor phic H-H H-R I— zd < Q (5 O M | E autapomorphic R-H R-R Ll FIGURE 17. Two by two contingency table showing Pse evolutionary explanations for observed eco- logical associates and individual ecological life history traits. H = his solace or inherited; R = recent, or mod- ified in the given ca The applications of historical ecological anal- ysis in this third area are just beginning. I wil now present a case study to illustrate the kinds of applications I envision for historical ecology. We may establish the topology of interactions for a given ecological association by direct ob- servation. I have tried to show that it is possible to establish robust hypotheses of the relative re- cency of association among the members. By us- ing those two types of data, it is possible to es- timate the extent to which contemporary interactions are responsible for ecological life history traits. Species may occur together be- cause they evolved together, vicariantly, or be- cause at least one dispersed from its area of or- igin. Ecological life history traits exhibited by two or more interacting species may be traits inherited unchanged from ancestors or they may be traits that evolved concomitantly with the contemporary association. Figure 17 summariz- es the four possible combinations of circum- stances this provides. For members of a vicariant association whose ecological interactions result from plesiomorphic (ancestral) ecological life history traits, their in- teractions may be completely explained by ref- erence to historically determined properties of the organisms, namely, their inherited traits and their geographical history. There may also be associations in which at least one member has dispersed from its area of origin but for which interactions still result from expression of ple- 1985] siomorphic ecological life history traits. For such associations only the contemporary ability of one (or more) ofthe species involved to disperse need be added to the historically determined traits to explain the observed interactions. For neither of these two classes of observations may contem- porary ecological interactions be logically in- volved to explain the traits exhibited by the species involved. The other two classes of observations are char- temporary association. Those trai arisen independently of ancestral interactions or may have arisen as a result of ancestral inter- actions. Only for the latter case can it be said that influences extrinsic to the organism, in this case the ecological interactions of other species, were responsible in any way for a particular set of traits that we observe today. In many cases, it is possible for different equal- ly-parsimonious interpretations of the ancestral condition to be postulated (Farris, 1970; Mick- evich, 1982; Swofford & Maddison, unpubl. data). The examples that I will present next have been chosen so that they are relatively unambiguous, but research into the properties and assumptions of various optimization schemes for predicting ancestral traits is of utmost importance for ap- plying this technique generally. Holmes (1973) reported that the digeneans Psettarium sebastodorum and Aporocotyle mac- arlani, inhabiting the circulatory system of rockfishes, exhibited little niche overlap. Pset- tarium sebastodorum occurs almost exclusively in the heart whereas A. macfarlani is almost en- tirely restricted to the blood vessels of the gill arches. Holmes assumed that parasite commu- nity structure is affected to a large degree by in- terspecific competition. From this, he concluded that the non-interactive site selection by the two species of blood flukes was de facto evidence of past competition. Rohde (1979) examined the same data as Holmes. He assumed that natural selection could explain the observed differences in site selection easily. All one need do is invoke the notion that selection would favor flukes that clumped to- gether in the host because they would have a higher probability of mating. His conclusions: one need not invoke competition to explain the observations. The data are evidence of the effec- tiveness of natural selection. Price (1977, 1980) also examined Holmes' data. BROOKS—HISTORICAL ECOLOGY Aporocotyle Psettarium heart ^^ and gills gills 1 E 18. Predicted sets of cladograms for rock- blood flukes, including the h of the FIGUR recent (autapomorphic) traits (redrawn from Brooks, 19802). His assumption was that parasite communities, because they are generally found to be below MacArthur-Wilson (MacArthur & Wilson, 1967) equilibrium, are never established and persistent for long periods but are always young colonizing communities. He further asserted that general- ized parasites are young colonizers and special- ized parasites are not. Given the two specialized parasites in a young colonizing community, Price concluded that the parasites were pre-adapted to their specialized niches and therefore colonized em. Competition and selection played mini- mal roles in their site selection in rockfish. Finally, Brooks (1980a, 1980b) suggested that the parasites could be part of a coevolving fauna, that is, a vicariant ecological association, exhib- iting plesiomorphic ecological life history traits e 672 for site specificity (habitat loyalty). The four dif- ferent opinions each correspond to one of the pum classes listed in Figure 17 and are shown n Figure 18. Brooks (1980a, 1980b) suggested m each of the four explanations predicted. an explicit set of phyl S in characters and/or hosts. For Brooks' contention to be correct, the two different parasites must exhibit a vicariant (7 coevolving) relationship with the same host group and must exhibit plesiomorphic life his- tory traits. Figure 18a summarizes those attri- butes. Price's hypothesis (Figure 18b) would not differ too greatly, except that either Psettarium or Aporocotyle, or both, would be colonizers in rockfish, so the host-parasite relationships in- dicated by each parasite group would differ Rohde's hypothesis involves hosts and parasites that are together for a considerable period oftime uring which natural selection promotes pro- gressively narrower site selection. This is shown in Figure 18c, where the parasites exhibit vicar- iant host relationships as well as autapomorphic ecological life history traits. Finally, we consider Holmes' hypothesis of competitive interactions mediating site selection (Fig. 18d). Corrobora- tion of this class of hypotheses requires both ele- ments of dispersal and the expression of auta- pomorphic traits relevant to the dispersal. Autapomorphic traits are features of the evolu- tionary context of the ecological interactions. Holmes and Price (1980) attempted to analyze the Psettarium/ Aporocotyle/rockfish system ac- and were “ecological specialists" in other hosts. Note that this corresponds to the situation found in Figure 18b, in which the ecological life-history traits exhibited are plesiomorphic, that is, his- torically constrained. Thus, Holmes and Price were incorrect in asserting that their conclusions supported strict ecological determinism. They also asserted that . the genealogies of para- sites and their hosts are clearly not congruent Dapur. all four cases of Brooks' Figure 4)." Brooks' “Figure 4" refers to Figure 18. Note that for only Figure 18a and 18c are host and parasite genealogies congruent. Note also that Figure 18b ponds to th lusi Hol d Price rew. Thus, they were also incorrect in saying that they had eliminated all four cases Brooks (1980a) proposed. These inconsistencies in their ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 conclusions led me to re-examine Holmes and Price’s phylogenetic hypotheses. Regarding Pset- tarium, they wrote: “The genealogy of three species of Psettarium is not clear (see differen- tiating characteristics in Holmes, 1971). How- ever, it does not appear that P. japonicum and P. tropicum are more closely related to each oth- er than to P. sebastodorum. The male terminal genitalia of P. sebastodorum are more similar to those of P. japonicum than to those of P. tropi- cum, suggesting ` genealogy of figure 2B.” Upon examining the primary literature, I found that the similarity in ale terminal genitalia between P. sebastodorum and P. japonicum is based on their possession of a cirrus sac (a bag containing the male intromittent organ)— P. tropicum lacks a cirrus sac. However, the presence of a cirrus sac is plesiomorphic not just for fish blood flukes, but for all flukes. Therefore, its absence in P. tropicum is not indicative that P. sebastodorum and P. japonicum are each other's closest rela- tives. However, P. tropicum and P. japonicum are both wide-bodied, whereas P. sebastodorum is slender and elongate. In order to determine which trait might be plesiomorphic, we needed to examine close relatives (out-groups) of Pset- tarium spp. Manter (1954) had stated that the differences between Psettarium and the genus Cardicola were “‘not clear" and “seemed rather inadequate." Holmes (1971) stated: “These two genera [Psettarium and Cardicola] are very sim- ilar, and certainly do not belong to different subfamilies." Of the nine species of Cardicola, some are elongate and some have wide bodies. Finding no clearcut support within Cardicola, I then examined two other related species, namely, Paracardicola hawaiiensis and Neoparacardico- la nasonis. Both of these species have two testes, the plesiomorphic condition for flukes. Members of Cardicola and Psettarium have only one tes- tis; thus, they would seem to form a monophy- letic group based on that trait. Both P. hawaiien- sis and N. nasonis are elongate worms, suggesting that elongate bodies are plesiomorphic for the group. Thus, Psettarium tropicum and P. japoni- virtue of possessing a bend in the posterior mar- gin of the body at the point of the male genital pore. However, Neoparacardicola nasonis ex- hibits that trait, making it either a convergent trait or a plesiomorphic trait among these flukes. Thus, Psettarium has no real support as a mono- FicuRE 19. Cladogram depicting phylogenetic re- lationships of species of Aporocotyle, a genus of blood flukes inhabiting fish. phyletic group distinct from Cardicola. Cardi- cola, however, has been distinguished only be- cause it lacks the body bend of Psettarium; thus, there is no character support for Cardicola as a real group either. I will return to this point later. The genealogy for Aporocotyle presented by Holmes and Price (1980) suffers from similar analytical flaws. Much of their cladogram is based on presence or absence and arrangement of body spines and number of testes. All members of Aporocotyle have multiple testes, as do members of their closest relative, Sanguinicola. However, as mentioned before, the plesiomorphic number of testes in flukes is two. Holmes and Price's genealogy of Aporocotyle arranges species begin- ning with those having more than 130 testes and terminating with those having 25-32 testes. Out- group comparisons would support a reversal of that trend. In addition, enlarged lateral body spines are used as indicators of relationships among species of Aporocotyle, even though San- guinicola spp. have them as well. Figure 19 de- picts the phylogenetic hypothesis best supported by those poi re-interpreted in strictly phy- logenetic term In Figure 2o the cladogram for Aporocotyle and one for the three Psettarium species are dis- played, with their host groups superimposed in place of species names. Although there is not much resolution, there is no evidence supporting host-switching. The possible vicariant associa- tion between the two groups of blood flukes and BROOKS— HISTORICAL ECOLOGY 673 IGURE 20. Host relationships indicated by phy- eri relationships of Psettarium spp. (top) and Aporocotyle spp. (bottom). the various host groups could be better examined if species of Psettarium occurred in cottoids or scombroids. Interestingly, four species of Car- dicola occur in cottoids and three in scombroids. At least one of those occurring in cottoids, C. laruei, has a wide body like P. japonicum and P. tropicum. [The remaining two species of Car- dicola inhabit mugiloid and labroid fishes, re- spectively.] Figure 21 shows Figure 20a redrawn to include the species of Cardicola; Figure 22 shows the relative relationships ofthe fish groups involved and their associated blood flukes. The new analysis demonstrates no incongru- ence among the host and parasite cladograms (Fig. 23). However, the vast majority of species of fish in each of the host groups have no blood flukes reported from them. This suggests that (sampling error aside) although both groups of blood flukes have coevolved with their hosts, that coevolution has been marked by indepen- dent failure of the two groups of flukes to persist in all evolving host lineages. The absence of par- asites from many hosts does not indicate the de- gree of relationship among those that survive. It is not inconsistent with coevolution that in only one known case do members of Aporocotyle and ANO T ` o ar ce Q d oc o C € o gor" we _Phylogenetic Mog S of Psetta- FIGURE 21. polyphyletic and Cardicola is paraphyletic Psettarium- Cardicola occur in the same host. This is especially true if the hosts in which the co-occurrence ar pn are relatively primitive, as they are in this c Having EE. ds temporal context of the association between Aporocotyle macfarlani, Psettarium sebastodorum, and Sebastes spp., we may now examine the possibility of interactive or non-interactive factors determining the site- selection exhibited by the parasites. For this we take the two parasite cladograms and superim- pose their habitat site rather than the species names at the ends of the branches. We then es- Farris, 1970) (Figs. 24, 25). Having those data, we may see that A. macfarlani occurs in the gills because its ancestor was a gill-inhabiting parasite but P. sebastodorum is most likely descended from ies that inhabited tl teric blood vessels. Thus, P. sebastodorum exhibits an aut- apomorphic trait for site selection. This would appear to support Rohde's hypothesis, for that species at least. However, no other species in the group shows a similar tendency. The shift in traits ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 FiGURE 22. Host relationships indicated by phy- ped. relationships of Psettarium/Cardicola spp. d IGURE 23. Summary set of host relationships sup- di, both Aporocotyle spp. SR and Psettarium/ Cardicola spp. (PC). Note congruence 1985] gills mesenteric blood vessels, gills BROOKS— HISTORICAL ECOLOGY 675 bulbous and $ auricles, ventricle bulbous arteriosus heart and gills FIGURE 24. Summary of site selection traits for species of Aporocotyle. Sorig indicate required evolutionary abe question mark indicates unknown data; would not seem to be explicable in terms of com- petitive exclusion, because A. macfarlani does not occur in the mesenteric blood vessels. The change in site by P. sebastodorum does not seem to be correlated with an extrinsic variable, so we conclude that the structure of this ecological as- sociation is historically determined. This con- trasts with the South American freshwater sting- ray parasite associations that include some colonizers. It is important to emphasize that historical evidence for adaptive changes of the type sought by evolutionary ecologists appear as exceptions to the historical pattern of structural and func- tional traits. Adaptive shifts are manifested by the same, or highly similar, traits arising in species that are not each other's closest relatives, but that share common ecological regime, in which the convergent traits are correlated with that partic- ular ecological regime. In asking the question, *How often do such shifts occur?," Ross (1972b) concluded that for a survey of various insect groups the answer was 1-2096, with an average of about 396 (one in every 30 speciation events). This accords well with my studies of parasitic helminths. If this is generally true, there does not star indicates A. macfarla mesenteric blood vessels mesenteric blood vessels FIGURE 25. Summary of site selection traits for species of Psettarium/Cardicola. Arrows indicate re- T evolutionary changes; star indicates P. sebas- todoru 676 p w o © (A) (B) (C) (D) 1 2 3 4 | ` (Y) (X) (Z) (A) (B) (C) (D) 2 3 4 Cc Y) (A) (x) (C) (Z) d FiGuRE 26. Demonstration that degree of co-ac- commodation (host specificity) need not be great in order for the method proposed in this study to detect historical patterns.—a. Depicts a host taxon’s phylo- genetic relationships. — b. Shows that the ded taxon 1-2-3-4 has co-speciated with the host taxo c. De- ote that co-speciation of d 1-2-3-4 can still be discerned. — d. Depicts the host-parasite relationships of parasite taxon 1-2- 3-4 if B and D are no longer host members of the parasite taxon. In this case, even though a number of unrelated hosts are used, the historical pattern is still present (redrawn from Brooks, 1981b) seem to have been enough ecological diversifi- cation to explain evolutionary diversification. If historical constraints play a major role in determining ecological structure, we would ex- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 pect to find evidence of them at higher systematic levels than the species. As mentioned before, Brooks et al. (1985b) recently presented a phy- logenetic analysis of the 62 families of digeneans (to which Psettarium, Cardicola and Aporocotyle belong). Only 26% of the groupings and 3596 of all branches on the phylogenetic tree were char- acterized by ecological changes. Classes of eco- logical traits considered were: (1) changes in de- velopmental programs producing increased numbers of infective larvae, (2) changes in en- cystment behavior of ee oae iA ud in first host, (4) ond LB I hosts, (5) shifts in final host, and (6) d in site of infection in the final host. Ecological changes definitely lagged behind morphological changes. Trends seen in species-level analyses are found in family-level analyses as well, attesting to the influence of historical constraints on eco- logical diversification. SUMMARY Historical ecology provides a missing com- ponent in studies of the evolution of ecological associations — history. This may seem like a strange assertion, since it is certainly true that evolutionary ecologists are vitally interested in historical phenomena. I think two perceptions have led to the exclusion of historical compo- nents in evolutionary ecology. The first is the perception that there exists no protocol for pro- ducing direct estimates of history that has any degree of empirical rigor. I hope that my few examples and the rest of the contributions in this volume provide reason for optimism that such a protocol is now firmly entrenched. The second perception goes something like this: even if we could provide direct estimates of history, the in- direct measures we use now give us the same answers. Historical ecology allows us to inves- tigate this perception. In the past, I have examined three components of the evolutionary ecology of host-parasite sys- uili numbers of species, (2) host specificity, and (3) ‘‘resource-tracking”’ models (see Brooks, 1979b, 1980a, 1980b, 1981b). Consider the following possibilities de- rived from historical analysis. Suppose we dis- cover an association that is very old and histor- ically structured, such as the Parana River system stingray parasite fauna, and yet that association exists below predicted “equilibrium” numbers. Price (1980) noted that parasite communities 1985] generally exist below equilibrium numbers and used that observation as evidence that parasite community structure reflects young, colonizing phenomena. From the historical perspective, it seems that the notion of “equilibrium” numbers of species may be an artifact of one theory of community evolution rather than empirical ne- cessity. Now let us consider the question of host spec- ificity from the historical perspective. I have sug- gested previously that degree of host specificity is decoupled from historical association of lin- eages. Figure 26 provides an example of this. Notice that no matter what differences exist in degree of host specificity, the history of the par- asite lineage is still coordinate with that of the original host lineage. Conversely, it is not nec- essarily true that very pronounced (host) speci- ficity indicates a long-standing association. Fi- nally, a related concept is that of "resource tracking" models of coevolution (see e.g., Keth- ley & Johnston, 1975). Resource-tracking models imply a set of specific historical predictions that are seldom, if ever, acknowledged. Chief among these is the assertion that not only do ancestors always persist, ancestors are always among the known species at any given time (see Brooks, 1981b for a fuller discussion). Thus, it seems that itis not true that the indirect estimates of history will always provide the same answers as expla- nations derived from direct estimates of history. Hence my assertion, at the beginning of this re- view, that historical ecology provides a comple- mentary approach with evolutionary ecology to attaining insights into causal mechanisms of the evolution of ecological associations. Certainly the most interesting durstions ling involve cases in which the two app fferent answers Finally, I would like to comment on the view, from historical ecology, of the current enthu- ner hansos g ecologists concerning the relative merits of *competition" or “random as- sociation" null hypotheses in explaining the structure, or organization, of communities (see Lewin, 1983a, 1983b for a review). To do so, we must first represent the question in historical terms, then produce maximum randomness and minimum randomness configurations. I tried to show how to represent such questions in histor- ical terms using the fish blood fluke example. A convenient measure of the relative randomness of an association of species and their ecological life history traits is the statistical entropy of the community ensemble. Using the method pi- BROOKS—HISTORICAL ECOLOGY 677 oneered by Karreman (1955) and modified for use in examining phylogenetic questions (Wiley & Brooks, 1983), it is possible to show that the minimum randomness configuration is the one that is wholly determined historically. It is also possible to show that maximum randomness is not achieved by a random assemblage of species with fixed traits, but by the assemblage (random or not) of species with unlimited polymorphism for ecological traits; in other words the maxi- mum randomness null hypothesis is indistin- guishable from a maximum competition model when viewed historically. Somewhere in between lie all systems experiencing some competitive interactions. All of these are relatively more ran- m than historically determined ecological as- sociations. Thus, from the perspective of histor- ical ecology, it is difficult to understand the hypothesis that competition is an organizing force in evolution. Accounting for such differences in perspective should provide exciting research projects in the future. LITERATURE CITED ANDREWS, J. M. 1927. Host-parasite re tru in the coccidia E mammals. J. Parasitol. 13: 194. ASHLOcK, P. D. 1974. The uses of cladistics. Annual . 5: 81- 48. Les helminthes, parasites des ver- e leur evo- lution et celle de leurs hotes. Ann. Sci. Franche- Comté 2: 99-113. Boucor, A. J. 1983a. , Area- -dependent- -richness hy- Naturalist 121: 294—300. 1983b. Does evolution take place in an evo- lutionary vacuum? J. Paleontol. 57: 1-30. modi us 5 1983. The Eclipse of Darwinism. Johns s Univ. Press, Baltimore m 1979a. Testing hypotheses of evolu- Testing the context and extent of host- parasite coevolution. Syst. 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Studies on hah al trema- todes of Hawaiian fishes: wee Opecoelidae Ozaki, 1925. Zool. Jahrb. 93: 173-202. Resu, V. H., J. C. Morse & I. D. WALLACE. 1976. The evolution of the sponge-feeding habit in the tion in parasites. Amer. Naturalist 114: 648-671. Rosen, D. E. 1975. A vicariance yov of Caribbean biogeography. Syst. Zool. 24: 431-464. 1985. Ge ological Ed and biogeo- graphic c ongruence in the Caribbean. Ann. Mis- souri Bot. Gard. 72: "636-659. Ross, H. H. 1972a. An uncertainty principle in eco- pion iw cdi In R. T. . All en & F. C. Jam (edito Pap. ium Arkansas Mus. 4: 133-161. 1972b. The origin of species diversity in eco- logical communities. Taxon 21: 253-259. STAMMER, H. J. 1957. Gedanken zu den parasito- phyletischen Regeln und zur Evolution der Para- siten. Zool. Anz. 159: 255-267. SzibpAT, L. 1956 marine charakter der Parasi- Sc des Rio de la P 1 utung als Re- liktfauna des Tertiasen Tethys-Meeres. Proc. XIV Int. Congr. Zool. 1953: 128-138. 1960. La parasitologia como ciencia auxiliar paras develar problemas hidrobiologicos, zoogeo- graficos y geofisicos del Atlantico Sud. Libro Ho- menaje Eduardo Cabellero VERSCHAFFELT, E. 1910. The cause determining the 680 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 72 selection of food in some herbivorous insects. Proc. Plant Biogeography. Chronica Botanica Co., Hid cad. Sci. Amsterdam 13: 536-542. tham dd sachusetts. [Translation of 1932 R WILEY, E. O. & D. R. BRooks. 1983. Nonequilibrium sian n tex thermodynamics and evolution: a response to — ZSCHOKKE, F 1933. Die puis als Zeugen fur die Levtrup. Syst. Zool. 32: 209-219. geologische V Trager. Forsch. & Wu.rr, E. V. 1950. An Introduction to Historical Fortschr. 9: 466—467. PHYLOGENETIC PATTERNS AND HYBRIDIZATION! V. A. FUNK? ABSTRACT Hybridization is an important part of the evolutionary history of flowering plants. If hybridization has occurred amon the species of a taxon under cladistic analysis the results are varied but always present additional difficulties. Hybridization results in incongruent intersecting data that obscure the underlying hierarchy. Guidelines and methods are examined for their usefulness in identifying possible hybrids in a cladistic study. Seven genera are analyzed cladistically and the resulting cladograms examined for possible hybrids. These hypotheses of hybridization are then compared to other data, such as distribution and cytology, to see if the hypotheses of hybridiza tion are supported or rejected. The more hybrids an analysis contains and the more complex the interactions, the more difficult it becomes to identify possible hybrids and their parents. It is difficult to overemphasize the importance of hybridization and yanasa h in evolution be- cause they turesof many plan "pu According to some od 30-80% the species of angiosperms are polyploids (Goldblatt 1979; Lewis, 1979; Stebbins, 1974), which allows for the possibility of a tremendous amount of hybridization. Of course, these figures h e nored by those who have dominated the discus- sion of evolutionary theory. This is a result, no doubt, of evolutionary theory being largely in the hands of scientists who work on groups in which such phenomena as polyploidy and hybridiza- tion have a strong relationship with unisexuality and are not considered important in evolution. There are different types of hybridization. Fi ure ] Summarizes some of the possibilities (a nis found in Funk, 1 but does not include introgression. For the pur- pose of this paper it is important to note that many hybrids are sexually reproducing individ- uals and are morphologically distinct and in some manner reproductively isolated from both par- ents. Thus, they behave as species no matter whose definition you chose to adopt. - — THE STUDY OF HYBRIDIZATION The basic concept of phylogenetic systematics (sensu Hennig, 1966) is an ever branching pat- tern or hierarchy. The method of cladistics (phy- logenetic systematics) seeks to discover these patterns es grouping together taxa Wees share m les (evolutionarily novel, unique, or derived edd rs). idization, or PLAN evolution, is inconsistent with a method de- signed to depict hierarchies. Hybridization is, therefore, a cause of incongruent, intersecting data that obscure phylogenetic information. Cladists have been concerned with this problem for sev- eral years. Most realize that any method that seeks to identify patterns of relationship must be able to accommodate hybridization because of its frequency. Workers in the problems of hy- bridization and phylogenetic systematics include Bremer (1983), Bremer and Wanntorp (1977), Clark (1982), Funk (1981), Humphries (1983), Humphries and Funk (1984), Nelson (1983), Nelson and a (1980), Rosen (1979), Wag- c Wanntorp (1983), and Wiley One favorite aware of botanists in estimating the closeness of relationships among taxa is the percentage of hybridization in crossing studies. An important point about such hybridization studies was made by Rosen (1979: 277): “‘repro- ductive compatibility is a primitive attribute for the members of a lineage and has, therefore, no power to specify relationships within a genea- logical framework.” We cannot use the ability of two or more species to hybridize as an indication of close relationship because the ability is rela- ! A number of people, who do not necessarily agree with everything that I have said, have kindly provided with data and comments on various drafts of the manuscript and this paper would not have been possible me ^4 S papar They include: R. Jansen, J. Semple, C. Clark, R. Sanders K. B r, G. Nelso C. Humphries, H.-E. Wanntorp, n, N. Platnick, D. Rosen, and P. Weston. I appreciate the assistance of B. Kahn in helping prepare po feta ns Depart ment of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. ANN. MissouRi Bor. GARD. 72: 681-715. 1985. 682 | ASEXUAL | SEXUAL | I I | [ | | | I | | l I Possible pangs and polyploid relation- o species (A and B) and their reproductive nad (Funk, 1981). I nen do tively ancestral, possessed at one time by all members of the group. In fact, it is the loss of the ability to hybridize that is apomorphic. Be- cause it is never certain that any two taxa are unable to reproduce, whether or not species hy- bridize is uninformative in determining the pat- tern of relationship. Among cladists, three different approaches st parsimonious cladogram(s) and leaving the homoplasies (char- acters appearing more than once on the clado- gram) resulting from the presence of hybrids as the true reflection of the character pattern. Others have advocated removing the hybrids that have been identified by their ‘intermediacy’ at the be- ginning of the analysis. The third group advises leaving all of the taxa - the sir ia and then closely examining the cladograms for polytomies (nodes with more than Two branches) and Chat t may acter There are problems with all three approaches. The first does not accurately reflect the character pattern; as we shall see, hybrids may appear on the cladogram in a polytomy or with character conflicts or even as ancestors when they are none of these. The patterns do not reflect the accurate sister-group relationship (nor are they true rep- resentations of phylogeny). The second approach assumes one can identify the hybrids prior to the analysis and this is not possible in many cases. The third position relies to a great extent on hy- brids causing polytomies. Further analysis has shown this usually does not happen (Humphries, 1983) ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Wagner (1983) has suggested a method for dealing with hybrids that he calls reticulistics. This method works with groups in which hybrids are intermediate in character and progressively less well with those that are not intermediate. Often the hybrids to which Wagner refers are F,'s that are being formed continually and are sex- ually inviable. Certainly for well-defined plant groups in which the hybrids are intermediate and obvious and are characterized by being rare and either sterile or polyploids (definition, Wagner, 1983: 71), Wagner's method should be consid- ered. These individuals are not units ofevolution and thus are not species. In this paper I am con- cerned with hybrids that are regarded as evolu- tionary units and that are usually designated as species, subspecies, or varieties. Theoretically, in cladograms, hybrids show up by causing character conflicts; so also do homo- plasies (Fig. 2, character 4; Appendix A). One must be diligent in trying to distinguish between character conflicts caused by hybridization and those that are the result of parallel or convergent evolution. It is advisable to use the cladogram (developed with the concept of parsimony using all taxa) to examine the apomorphies that appear more than once on the cladogram (homoplasies). Perhaps a closer examination will reveal that m homologies) or are combinations of characters. For instance, not all black anthers in the Com- positae genus Montanoa Cerv. are homologous. Although originally treated as one apomorphy (Funk, 1982) a close examination showed that some black anthers were black only around the edges of the thecae, some were black only on the op of the connective, while others were com- pletely black. This additional information indi- cated that “black anthers” was not a single apo- morphy but three apomorphies. Character conflicts can also be the result of a designated apomorphy actually containing several charac- ters. Characters such as habit, chromosome number, and flower regularity, all can be divided into several characters. New apomorphies can be added to the cladistic study to reflect the addi- tional information because the “groups of char- acters" should not be viewed as character con- flicts but rather as separate apomorphies. Such changes should be made only when available evi- mai either undetected homoplasy or hybridization. FIGURE 2. flict that can a the result t of either ipa eria OT parallel evolution A lack of apomorphies can also be caused by hybridity. The hybrid does not necessarily in- herit all of the apomorphies of both parents. This observation is important: there is no reason to FUNK—HYBRIDIZATION 683 assume that all apomorphies are dominant over the more plesiomorphic characters of a trans- formation series. The data presented later in this paper indicate that in the heterozygous condition of the hybrid there might well be a higher per- centage of plesiomorphic characters showing in the phenotype. Therefore, it is possible for the hybrid to show only preemorpiies PME ancestral, or on the cladogram in an ancestral ee Ta Figure 2, taxon D could be a hybrid, between taxon C and any other taxon, that inherited the plesiomorphies of both parents. Indeed, it is in- teresting to speculate on whether or not one could use such cladistic studies to identify interesting genetic problems. Often the hybrids in a cladistic analysis will be grouped with one or the other of the parents depending on with which parent they share the most apomorphies (Humphries & Funk, 1984). When the putative parents are sister species (two species that are more closely related to one another than they are to any other species), hy- FIGURES 3-6. Cladograms illustrating the pattern of species A and B and their hybrid, H. FIGURE 7. Cladogram illustrating the pattern of species A and B and their hybrid, H. brids are quite often apparent regardless of whether or not they form polytomies. Hybrids that form dichotomous branching patterns can be identified as such so long as they have at least one apomorphy of both parents or if they lack an autapomorphy of the parent with which they are grouped. For instance in Figures 3-5 (for characters see Appendix B), species A and B hy- bridize to give species H. If there were an equal number of apomorphies in A and B (characters 4 and 5) and if both were found in the hybrid, the result could be expressed as two equally par- simonious cladograms (Figs. 3, 4) or as a tricho- tomy (Fig. 5), although the latter involves one more character change and is therefore one step longer. If the incongruent character set is inferred to be the result of hybridization, the hybrid could be placed above the diagram connecting with both parents (Fig. 6). However, if one parent taxon had one more autapomorphy than the oth- er, or if the hybrid showed unequal character inheritance, then a single cladogram results. For instance, if taxon A (Fig. 7; Appendix C) had two autapomorphies (5 and 6), and the hybrid in- herited all apomorphies of both parents, the most parsimonious cladogram would give the result shown in Figure 7. Nonetheless, because they are sister taxa, the possibility of hybridization is ap- parent, so long as the hybrid has at least one apomorphy from each parent. However, as men- tioned earlier it is possible for the hybrid not to display all the apomorphies. If the hybrid in Fig- ure 7 did not have character 4 there would be no indication that it was a hybrid (except for the ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 more indirect evidence of lack of apomorphies in taxa A and H). If one parent has very few autapomorphies there is less chance that the hy- brid will have any indication of its history. Sometimes there is more than one hybrid from the same two parents (Figs. 8-12; Appendix D). The two hybrids are most parsimoniously grouped with either parent (Figs. 8, 9) but the two equally parsimonious cladograms indicate the hybridity of H1 and H2. The most parsi- monious cladogram has two reticulations (Fig. 12). It 1s, of course, possible that taxa H1 an H2 are the result of a single hybridization event followed by segregation. One way to evaluate this possibility is to examine the distribution of the taxa in question. The possibility of hybridization followed by segregation lessens as the distance between the hybrid taxa increases. The only time it is “most parsimonious" to have a polytomy is when the hybrid does not have any of the autapomorphies of either parent (Fig. 13; Appendix E). However, parent taxa are not always sister taxa. For instance, Wagner (1954) has shown that at least three diploids are involved in producing the hybrids in Asplenium, and Grant (1953, 1964) has shown that species from different species groups are hybridizing in Gi/ia (Funk, 1981). In cases such as this the task of identifying hybrids becomes more difficult. For instance, given the cladogram in Figure 14 (Figs. 14-17, 19; Ap- pendix F) the most parsimonious cladogram that includes the hybrid, places the hybrid (H, Fig. 15) as the sister taxon of the parent that involved the least number of homoplasious events (the number of character changes or length of this cladogram is L — 11). The length would increase if the hybrid were grouped with the other parent (Fig. 16, L = 12) because there is one more ho- moplasy. It is much longer to form a polytomy (Fig. 17, L = 14). The only time a polytomy would be formed in the most parsimonious cladogram is if the hybrid had none of the apo- morphies of either parent, at least those above the first node shared by both parents (Fig. 18; Appendix G). If the hybrid is identified as such it can be removed from the cladogram and placed above it giving an even shorter cladogram (Fig. 19, L = 9). The identification of possible hybrids is only the beginning of an analysis. Those cladograms indicating hybrids (e.g., Fig. 19) are merely hy- potheses of hybridization and should be tested by using other data, such as distribution, chro- 1985] FUNK—HYBRIDIZATION 685 Ficures 8-12. Cladograms illustrating some of the results when the same two parent species (A and B) produce more than one hybrid, H1 and H2. mosome number, karyotyping, and pollen fer- gram without a reticulation and continues by tility before they can be referred to as hybrids. adding reticulations one at a time so as to min- Nelson (1983) has suggested a method forana- imize character conflict. It is based on the idea lyzing cladograms for possible hybrids. His pro- that when there are two equally parsimonious cedure begins with the most parsimonious clado- ways of representing a homology on a cladogram FIGURE 13. Cladogram illustrating that it is more not show any of the autapomorphies of either parent. one should investigate the possibility of inserting a reticulation. If the reticulation results in a de- crease of apparent homoplasy and if the taxon exhibits character conflict of the “intermediate” type, the reticulation can be maintained. For a certain set of data (Appendix H) there are two equally parsimonious cladograms, with the same branching pattern. One cladogram (Fig. 20) has a parallel acquisition of character 4 (Figs. 20-22, Appendix H) and the other has one acquisition of character 4 and a subsequent loss (Fig. 21). However, one can introduce one reticulation and eliminate the need for homoplasy and/or loss (Fig. 22). All this diagram indicates is that if hybridization is involved it is most likely that taxon B is of hybrid origin. A more complicated example involves nine taxa and 12 characters. There are two equally parsimonious cladograms with the same branch- ing pattern with different amounts of homoplasy and loss (Figs. 23, 24; for characters for Figs. 23- 25 see Appendix I). By progressively adding re- ticulations, all need of reversal and/or homoplasy can be eliminated. Taxa H and I may be hybrids Fig. 25). Nelson's method of examination of to the one in Figure 27 by adding one oo and thereby shortening the clado Some groups have Aap that cause difficulties when using Nelson's method. For in- stance, some have their origin in hybridization ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 but have since speciated (developed autapomor- phies), some have numerous hybrids and even hybridization among hybrids, and some hybrids do not have all of the apomorphies of the parents (as in Fig. 2). The workability of Nelson's meth- od is dependent on the hybrid inheriting the apo- morphic characters from the parent taxa without too many character losses; otherwise the clado- gram with the reticulations (Fig. 28; for char- acters for Figs. 28, 29 see Appendix K) will be longer than the most parsimonious cladogram without reticulations (Fig. 29). Also, this method is only feasible when the percentage of hybrids in the data set is low because the possibilities become endless, especially when the hybrids are hybridizing. There are additional guidelines and methods that can be used when examining gr for possible hybrids. Use of these on seven data sets indicates that insights into the identification of possible hybrids and their parents can be gained by studying. e dioe patterns : oe the clado- grams as oidy levels of the taxa eer Some of mad guidelines and methods are elaborations and evaluations of previously published ideas and others are new. GUIDELINES AND METHODS FOR IDENTIFYING PossiBLE HYBRIDS AND THEIR PARENTS 1. When there are two cladograms of similar length and one taxon position changes, the taxon that is moving may be a hybrid and the two taxa between which it is moving may be the parents. In Figures 3 and 4, taxon H (the hybrid) shifts etween taxa A and B in the two most parsi- monious cladograms. Taxon H may be a hybrid and A and B may be its parents. In Figures 15 and 16 taxon H shifts between taxa C and D and may be a hybrid. 2. As an extension of number 1, it is possible to follow a path of character conflicts. Figure 15 has characters 6 and 7 appearing twice and this identifies taxon D, the parent with which H (the have to have all of the characters. For instance, in Figure 16, H might not have character 3 (Fig. 30; for characters for Figs. 30, 31 see Appendix L) but as long as it had 1 and 5 taxon C would emerge as the other parent (Fig. 31). 3. Taxa that are defined solely by character conflicts may be hybrids or parents. In Figure 1985] FUNK—HYBRIDIZATION 687 Ficures 14-19.— 14-17, 19. Cladograms illustrating the pattern of speci ies C and D and their hybrid, H.— 18. Cladogram illustrating when a polytomy is formed in the most parsimonious cladogram 15, taxa H (the hybrid) and D are defined only by characters that appear elsewhere on the clado- gram and in Figure 16, taxa C and H also have only homoplasies. The same is true for taxa C and B in Figure 20, taxa A and I in Figure 23, and taxa B and C in Figure 26. 4. Taxa with reversals may be hybrids. In Fig- ure 32 (for characters for Figs. 32, 33 see Ap- pendix M) taxon H (the hybrid) has not inherited all of the apomorphies of both parents and there- fore is defined not only by characters that appear more than once on the cladogram but also by 688 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 FicunEs 20-22. Cladograms illustrating a simple example of the Nelson (1983) method for analyzing clado- grams for possible hybrids. 23 à FIGURES 23-25. Cladograms illustrating an example of the Nelson (1983) method for analyzing cladograms for possible hybrids. 1985] FUNK—HYBRIDIZATION 689 FIGURES 26, 27. Cladograms illustrating the use of Nelson’s method for analyzing cladograms for possible hybrids in the genus Microloma character loss. Taxon H does not have characters and 8 from taxon C and is also missing char- acter 10 from taxon D. In fact, there has been enough inheritance of plesiomorphies to make the cladogram. With reticulations (Fig. 2H the (Fig. 32). , Another possibility is when there i one parent species rich in apomorphies and another lacking them altogether—hybrids might be in- termediate or they might evidence multiple loss. 5. Taxa without autapomorphies may be par- ents. If the hybrid inherits all ofthe apomorphies of the parent with which it 1s grouped, then the parent will have no autapomorphies (Fig. 2, tax- n A; Fig. 7, taxon A; Fig. 15, taxon C; Fig. 16, taxon D). If the hybrid fails to inherit any of the apomorphies of the parent with which it is not grouped, the hybrid will have no autapomor- phies (Fig. 13, taxon H)— normally in evidence as homoplasies. 6. Consensus Trees— Consensus analysis is developing rapidly as an aid in evaluating a num- gram formation P red by two or more cladograms. The consensus tree is a compromise classifica- FIGURES 28, 29. Cladograms illustrating that the cladograms with reticulations are not necessarily the most parsimonious. 690 FIGURES 30, 31. both parents are not present in the hy tion. Consensus trees were first developed by d (1972) and have been used in the context comparing “ao s Mas phenetic methods sed 1978; h & Polhemus, 1980; Sokal & Rohlf, 0s or to compare a clado- gram constructed from a chemical data set and an intuitive tree (Seaman & Funk, 1983). There are many different types of consensus analysis, Adams consensus (Adams, 1972), strict consen- sus (Sokal & Rohlf, 1981), majority consensus ush & McMorris, 1981), and durchschnitt consensus (Neuman, 1983). Only Adams con- sensus trees and Nelson component analysis will be discussed in this paper because of ease of use and explanation. For further investigation, other references include Mickevich (1978), McMorris et al. (1983), and a series of papers in Felsenstein (1983). ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 Cladograms illustrating that possible parents can be identified even if all apomorphies of rid. An Adams consensus tree (ACT) may have a topology different from any of the cladograms used to construct them. Figures 15 and 16 have a different topology, but if we concentrate on agreements we get Figure 14. Then we can add the taxon left out to the first node common to both of its positions (Fig. 17). A detailed dis- cussion of how to construct an ACT is found in Adams (1972). 7. Component Analysis (NCA) was devel- oped by Nelson (1979) as a consensus method. A component is any monophyletic group on a cladogram or phylogenetic tree. Every cladogram can be divided into a certain number of com- ponents that have more than one terminal taxon. In order to construct a Nelson Consensus Tree it is possible to add together components that are common to two or more cladograms. The Ficures 32, 33. Cladograms illustrating that character loss may be an indication of hybridization. FUNK— HYBRIDIZATION FIGURES 34, 35. Cladograms illustrating that by comparing the components (Table 1) one can identify the source of incompatibility. movement of one taxon from one side of the cladogram to the other can have the effect of changing all of the components (Figs. 34, 35, Table 1) and no consensus tree can be construct- ed. However, instead of searching for complete agreement among the components, we can ex- amine in what way they are different and identify which taxa are responsible for the lack of con- gruence. Examining the list of components for the two figures (Figs. 34, 35, Table 1) itis obvious that taxon H is causing the incompatibility. These various suggestions are not to be used individually but collectively to generate hypoth- eses of hybridization. We can then turn to other forms of data to corroborate or falsify our hy- potheses. Two of the most readily available in pow ner a Peli many group»5a I yP y others such as karyotyping and pollen fertility can be employed. We will see how such infor- mation can be used in the examples below. All cladograms were constructed manually. EXAMPLE 1. MICROLOMA (ASCLEPIADACEAE) BREMMER AND WANNTORP (1979) Microloma is an African genus of 19 species, nine of which are represented in the most par- simonious cladogram (Fig. 36, Appendix J) and an alternative cladogram that is three steps long- er (Fig. 37). Figure 37 shows taxon C sharing an apomorphy with taxon B and Figure 36 shows taxon C sharing four apomorphies with taxon D indicating that B and D may be the parents if C is a hybrid. Also, there is a path of parallel char- acters to follow to the parents especially in Figure 37. Furthermore, taxon D (Fig. 36) and taxon B (Fig. 37) haven "em hice, indicatinsthes might be the parents with which the hybrid is grouped. In addition, an ACT (Fig. 38) can be constructed that identifies the hybrid by placing it at the first node shared by the two parents. A NCA (Table 2) shows that C is responsible for the incompatibility between the two sets of com- ponents. EXAMPLE 2. ANACYCLUS (ASTERACEAE) HUMPHRIES (1979, 1981) Anacyclus (Appendix N) is a genus of Medi- terranean distribution with 14 species (Figs. 39— 42). The monograph and subsequent paper by Humphries (1979, 1981) included a cladistic analysis and speculations on the hybrid origin for three of the species. Figure 39 is the most parsimonious cladogram (L = 45) for the data furnished by Humphries (1983); the number in parentheses next to the lower case letter indicates the number of apomorphies that have that pat- tern of distribution. If we concentrate on the character types that conflict with the most par- simonious cladogram (p, b, f, q, 1) we can draw alternative cladograms that are slightly longer (Fig. 40, L — 50; Fig. 41, L — 52). Figure 40 has four character types in conflict (k, p, q, o) and a TABLE l. Components for Figures 34, 35. Figure 34 Figure 35 AH AB AHB ABC AHBC HF EF EHF DEF DEHF ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURES 36-38. Two cladograms of Microloma and their Adams consensus tree. gain and subsequent loss of two character types (c, d). Figure 41 also has four types of conflict (o, b, c, q) and two types of loss (c, d). The con- sensus tree for these three cladograms is illus- trated in Figure 42 and shows that four taxa are potential hybrids (N, C, I, J) because they have dropped in resolution and because they are de- fined either completely or primarily by charac- ters that are found elsewhere on the cladogram. In addition by listing the components ofthe three figures (Table 3) it is evident that all of the com- TABLE 2. Components for Figures 36, 37. Figure 36 Figure 37 CD BC ECD ED FGECD FGED HIFGECD HIFGED BHIFGECD BCHIFGED ABHIFGECD ABCHIFGED ponent incompatibility is caused by taxa I, J, C, and N. These four taxa are potential hybrids (Fig. 43) and their possible parents are as follows: possible possible hybrids parents N = AB x EF-LM I = J = GxH C = AB x D-LM Figure 43 is 42 steps long. Based on his analyses Humphries (1979, 1981) suggested three puta- tive hybrids (N, L, J)and indicated corroboration hypothesis of hybridization for these three species. According to Humphries in addition to sharing apomorphies with AB and EF-LM, A. officinarum (N) is known only as an extinct cul- tivar grown in the nineteenth century for phar- maceutical reasons. Anacyclus valentinus (I) and A. inconstans (J) have florets of intermediate length between A. clavatus (G) and A. homoga- mos (H). Also, A. valentinus (I) is a weedy taxon 1985] FUNK —HYBRIDIZATION 693 FIGURES 39-42. Cladograms of Anacyclus. occurring only on disturbed land in the southwest Mediterranean region, and A. inconstans (J) oc- curs sympatrically with the Algerian population of A. clavatus (G). Cytogenetic studies carried out by Humphries (1981) corroborated the hy- potheses of hybridity for A. valentinus (I) and A. inconstans (J). There was no material for cyto- genetic studies available for either A. officinarum or A. monanthos (C). There is, then, good TABLE 3. Components for Figures 39-41. reason to list both A. valentinus (I) and A. in- constans (J) as possible hybrids because the hy- potheses of hybridity have been supported b independent data. Anacyclus officinarum (N) has been supported as a hybrid by its cultivated na- ture but is more difficult to analyze because it is not known to be extant. Anacyclus monanthos (C) was not indicated by Humphries to be of hybrid origin. In his monograph, Humphries Figure 39 Figure 40 Figure 41 Conflicts IJ IJ IJ IJH GIJ GIJ IJ GIJH GIJH GIJH IJ GIJHK GIJHK GIJHK GIJHKLM GIJHKLM EFGIJHKLM EFGIJHKLM EFGIJHKLM DEFGIJHKLM NEFGIJHKLM NEFGIJHKLM N CDEFGIJHKLM DNEFGIJHKLM DNEFGIJHKLM C, N BN KLM AB C, N CDNEFGIJ C ABNCDEFGIJHKLM ABCDNEFGIJHKLM ABCDNEFGIJHKLM 43 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 H K LM m(3) 9g(3) e(3) a(5) FIGURE 43. Cladogram of Anacyclus with reticulations. (1979) listed it as a pioneer of sandy soil and in some areas a dominant weed; these are charac- teristics of some hybrids. He also mentioned that it resembles one of the two subspecies of A. ho- mogamos (H). However, Humphries thinks that A. monanthos (C) is a taxon exhibiting inter- mediate characters and not a hybrid because it has many autapomorphies (Humphries, pers. comm.). Because of the lack of additional infor- mation on this taxon I have left it as a dotted line on the cladogram (Fig. 43). The cladogram in Aron ao differs from that of Humphries hecau g number of parallel n as and was not as concerned if this created additional character losses. Therefore, taxon N is connected as an internode further along the diagram. This is im- portant because it is, in my opinion, unlikely that a hybrid will inherit all of the apomorphies of both parents. Thus, some plesiomorphies will be inherited resulting i in the loss of some characters. ited are the guide to the identification of possible parents. Anacyclus is a good example of the use of cla- distics to identify possible hybrids and then em- ploying other techniques including uia Lai that O of hybridization (Humphries, 1979, 1981) EXAMPLE 3. AGASTACHE (LAMIACEAE) SANDERS (1982, AND IN PREP.) The 14 species of Agastache Clayt. sect. Brit- tonastrum (Appendix O) are eat diploid and are confined to th t United 1985] micrantha comp. mexicana comp. r 1 FUNK—HYBRIDIZATION 695 pallidiflora comp. cana comp. qu———l T&F 1 mic prn wrt mex epl pim coc pd-c pd-p mrn brv pf-r pf-i pf-pne-hne-naur can rup ' 21 10 1 11(2) 28(1')X i? 27 à 11 107 11? 24/34 4 11(2) 5(2") 28X 7,3929 h on 21(2') 5(2” 6 30 8 4 2117) 20 at 9 12 17 33 10 24 32 23 21(2) 24 aur can rup 10 $ 5(2'') 21(2’) 5(2') 13 2 1 5(2) 25 1 3 1*2) 45 1 26 *25, 26, 27 $ 11(2) 44 FIGURES 44, 45. Cladograms of Agastache. States and Northwest Mexico. Looking at the neomexicana ne-n, A. pallidiflora var. greene pf-r, A. mexicana mex), taxa with only character conflicts (A. breviflora brv, A. mearnsii mrn, A. coccinea coc, A. pringlei prn), and taxa with character losses (A. pallida var. coriacea pd-c, A. pallidiflora var. gilensis pf-i). Three of these taxa (prn, brv, pd-c) are immediately identifiable as possible hybrids, A. pringlei and A. breviflora pallidiflora var. gilensis (because of the absence of character 24 and some connection with A. coccinea) and A. mearnsii [because of character 21(1)]. Examining each one individually, breviflora has characters 25, 26, and 27 (an sometimes 11). If A. breviflora is a hybrid then one parent (A. mearnsii) could be its sister species. In order to identify a candidate for the second parent, emphasize the characters A. breviflora has and not the characters it does not have. Then locate where else on the cladogram these char- acters are found (25, 26, and 27; Fig. 44). The presence of these three characters indicates A. Mexico, and northern Sonora an Agastache mearnsii is found a short distance from A. wrightii in southern Sonora and Chihuahua. The distribution pattern would suggest a hybrid- ization event is possible. A second possible hybrid, A. pringlei (prn) is sister to A. micrantha (one subspecies of which has no autapomorphies). Agastache micrantha has four character conflicts (13, 16, Ap times sympatric with A. micrantha (southern 696 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 72 . A brv TE micrantha comp. mexicana comp. ~~ pallidiflora comp. cana comp. LI 3 up ¥ 1 1f 1 m coc N, -— ` 1 P T / n2 pt-i \ - / 13 ^ 41307) / / ` ` / N PÀ / ` ES | i rty mex epl pim ; Pdp pdo Nm ie pf-r / pfp ne-h ne-n aur can 28(1°) ME ^ y 22 f 4 N ! T! 14 11(2) 11(2) / 3 / 10$ + n? V |! 5(2'') "d V X207 ?! / mnm r as 28 v É E: E '210) ma 35 I 21(2') 5(2') / N / ^ ` 6 727 \ 17/ N \ I 8 4 X ; \ | N 18 N I 9 A 3 \ l N \ l N \ Ys I 23 E: cc V ! 24 22 SP is, 21(2) \ Zt 10 11 25 5(2) 26 3 1 21 16 46 FIGURE 46. Cladogram of Agastache with reticulations. Chihuahua and southern New Mexico) and para- patric with A. mearnsii (southern Chihuahua and the Chihuahua/Sonora border). The A. pallidi- flora subspecific taxa and A. breviflora are dis- tributed in the southwest United States. If A. pringlei is a hybrid the most likely putative par- ents are A. micrantha and A. mearnsii. A third potential hybrid, 4. pallida var. co- riacea (pd-c), is sister to A. pallida var. pallida lli sibly something in the **Pallidiflora complex” be- cause the putative hybrid has characters 16 and 21. Based on geographical distribution, the most likely parent in the ‘“‘Pallidiflora complex” is A. mearnsii that has a parapatric distribution with A. pallida var. coriacea. The last two possible hybrids, A. coccinea (coc) and A. pallidiflora var. gilensis (pf-i), are more difficult to place because they have less infor- mation (fewer characters). We can estimate that A. coccinea is a hybrid and that one parent may be found in the group defined by character 32; because of the presence of character 21(1') the other parent would probably be A. mearnsii. The distribution of the species does not help in this in the **Pallidiflora complex” that have a distri- bution that would allow for the formation of this hybrid have character 24 except A. mearnsii. However, A. pallidiflora var. gilensis has an aut- apomorphy, and this makes it less likely that it is of hybrid origin (but does not exclude it). Based on this analysis the following hybrids are pos- 1985] FUNK -—HYBRIDIZATION 697 x-9 x=3 x=4 x= 9 x=9 flo gdg gdv lan lat gs-gd sca BR pil mar gs-gg gs-c gs-h sub gs-gt lil li-d 22 3 4 2B 13 TA 21 14 19 22 14 12 15 16 16 21 Am Š 5. u 14 14,19 4 18 17 8 ° 3h gs-c gs-h sub 14 4 19 16 11 2A 18 12 21 2B 14 49 6 gs-gg gs-c gs-h sub gs-gt li-l li-d 8 47 (x2 5) FicunEs 47-49. Cladograms of Chrysopsis and Bradburia. sible: bry = wrt x mrn, prn = mic x *'Pallidiflora complex," pd-c = pd-d x mrn, coc = mex-plm, and pf-i = pf-r x some taxon without 24 (possibly mrn). Figure 46 shows the hypothesized hybrids and their possible parents. According to Sanders (1981), the hybrids and their parents are as follows: brv = wrt x mrn, prn = mic x mrn, -c = pd-p x mrn, coc = pfr x mrn (mex-epl + rn), and pf-i = pf-r x mrn (pf-f — pf-r + mrn). Although Sanders' estimates are in some cases more specific, there are no conflicts. EXAMPLE 4. CHRYSOPSIS AND BRADBURIA (ASTERACEAE) SEMPLE (1981, AND PERS. COMM.), SEMPLE AND CHINNAPPA (1984) Chrysopsis (Nutt) Elliot (Appendix P; 10 species) and Bradburia Semple & Chinnappa (Appendix P; 1 species) are yellow-rayed golden- asters distributed in the southeast United States (especially Florida) except for one species of Chrysopsis that occurs in the eastern United States. Using the characters: furnished by Semple ted (Figs. 47-49). There is a high level of homoplasy in the clado- am because only nine apomorphies lacked character conflicts and only four of these 11 are synapomorphies (10, 20, 11, 2A). Such a high level of character conflict and character loss is an indication of possible hybridization. e chromosome numbers were not used as char- acters in the analysis and are indicated on the cladogram to facilitate the discussion. Also, there is one report of x = 4 for Bradburia that is not indicated on the cladogram. Taxa C. lanuginosa (lan), C. gossypina subsp. gossypina f. gossypina (gs-gg), C. gossypina subsp. hyssopifolia (gs-h), and C. linearifolia subsp. li- nearifolia (li-I) are defined only by character con- flicts and/or character losses and may be hybrids themselves or the parent with which another hy- brid is not grouped. Two taxa C. god/reyi f. viri- dis (gd-v) and C. gossypina subsp. cruiseana (gs-c) are possible parents with which the hybrids are grouped because they have no autapomorphies 698 flo gd-v gd-g lat gs-gd sca 8 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Ficure 50. Cladogram of Chrysopsis and Bradburia with reticulations. and have a single taxon as a sister taxa. If we construct hypotheses based on these data we ob- tain the following results: lan = gd-g x li-l, gs-c & h = sub x pil or mar, and gs-gg = sub x some taxon without 8. Because so many of the taxa are sympatric or parapatric, the distributions are +1 1 not of refining The exceptions are C. gossypina subspp. cru- iseana and hyssopifolia (gs-c & h) that can be attributed to a cross between C. subulata and C. mariana sa x mar). There are four taxa with no autapomorphies [gd-g (C. godjreyi f. godfreyi), gs-gd (C. gossypina subsp. gossypina f. decumbens), gs-gt, li-d)] that are potential hybrids or parents. Their chro- mosome numbers show that all of the gs taxa are x = 9, and based on outgroup comparison the base number for the genera is probably x = (this agrees with Semple, 1981). Therefore, the subspecific taxa of C. gossypina are most likely CA hybrids. This supports the hypotheses of hybrid- ization for three of the subspecific taxa of C. gossypina (gs-gg, gs-c & h), however C. gossypina f. trichophylla and f. decumbens (gs-gt, gs-gd) were not identified, except for noting that they lacked autapomorphies. The cladogram clearly indicates why C. gossypina f. decumbens was overlooked. it h autapo- morphies. If it is a hybrid it is an excellent ex- ample of a hybrid inheriting all plesiomorphies of both parents and appearing in an ancestral position on the cladogram. Likewise, C. gossy- pina f. trichophylla was overlooked because it inherited most, but not all, of the plesiomor- phies. The chromosome number of x = 5 does not support, and in fact falsifies, the hypothesis of hybridization for C. lanuginosa because one of the hypothesized parents has x — 5 and the other x — Figures 47-49 show that the x — 9 taxa do not 699 frutescens group 1985] FUNK - HYBRIDIZATION californica group ve lac cal can al phe rad 10(2) 100074 1400 14(2) 12 16 13 9 3 7(2) FAN 14(2) i2 13 7(2) 16 8 1 3 2 12 5 51 FIGURE 51. Cladogram of Encelia. form a monophyletic group. Semple (1981) pro- posed that the x — 9 group is from one hybrid- ization event between an individual of C. su- bulata and one of C. mariana with subsequent selections to give different combinations of pa- rental genes. The cladogram does not support that statement because of the different combi- nations of characters in the five different sub- specific taxa. However, it also does not reject Semple's suggestion. It simply suggests that one should also consider the possibility of several different hybridization events involving the same species as parents with different characters being inherited each time (Fig. 50). Using cladistics alone has not given us a clear answer to the question of hybridization in Chry- sopsis and Bradburia, however when the hy- potheses of hybridization are tested with addi- tional information, such as distribution and ploidy level, we have been able to make five putative hybrids and gain some insight into pos- sible parents. EXAMPLE 5. ENCELIA (ASTERACEAE) (CLARK, PERS. COMM.) Encelia Adanson (Appendix Q) comprises 18 taxa distributed in the western United States. All taxa are diploids. The data matrix and the taxon distributions were furnished by Curtis Clark. There are at least two cladograms that are equally parsimonious (Figs. 51, 52) and one that is one step longer (Fig. 53); all have different branching patterns. There are several additional cladograms that have slightly different character distributions but do not have different branching patterns, and these are not illustrated. On the three cladograms there are several taxa that change positions. Encelia farinosa can be placed as the sister taxa of E. farinosa var. phenicodonta (Figs. 52, 53) or near the base of the cladogram (Fig. 51). In either case it is only defined by char- acter loss and/or homoplasy. Likewise E. asperi- folia is either the sister taxon of E. ventorum/E. laciniata (Fig. 51) or it is near the basal node (Figs. 52, 53). In all three positions E. asperifolia 700 californica group ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 frutescens group LI ven lac rad far phe can pal 1 10(1) 2 1 10(2) 12 4( 14(2) 9 7(2) 13 8 12 7(1) 16 3 13 8 1 12 2 5 52 FIGURE 52. Cladogram of Encelia. is identified by character loss and/or homoplasy. Encelia canescens is either the sister taxon of E. palmeri (Fig. 51) or it shares the node defined by character 7(1) with several other taxa (Figs. 52, 53). Two other taxa have some indication that they m may be hybrids, E. /aciniata with in- a cated as possibilities but cannot be placed as hybrids ancestors, or parents because they have few apomorphies. Two taxa (E. virginensis and E. actoni) have only one apo- morphy each and they appear on the cladogram at the basal node, and E. californica has a slight change in position depending on whether char- acter 3 is treated as a character loss or not (Figs. 51-53). Encelia farinosa and E. asperifolia stand out because of character losses and homoplasy. The best estimate for E. farinosa is that it is a hybrid between E. farinosa var. phenicodonta (its sister asp cal SC act vir rav res GC SF fru taxon in Figs. 52, 53) and something without characters 1, 2, and 3. Encelia asperifolia may be a hybrid between something without char- acters 3 and 12 (perhaps something in the **Fru- tescens group" *) and something in the **Califor- nica group" with character 8 (perhaps E. californica with which it is closely grouped and whose lack of apomorphies may account for E. asperifolia's similar situation). Encelia canes- cens might be a hybrid, between E. palmeri its sister taxon in Figure 51 and E. farinosa var. phenicodonta, because of the intermediate na- ture of character 14. Encelia laciniata may be a hybrid between E. ventorum and some other tax- on that has not left a trace. If so, E. laciniata is an example of a hybrid inheriting all of the apo- morphies of one parent. If the taxon from Santa Clara (SC) is a hybrid, one parent might be E. ventorum because of characters 13 and 16 and the other parent something from the ‘‘Californica group" that has character 12. A summation of possible hybrids is as follows: far = phe x some- frutescens group 1 r 1 GC SF fru 1985] FUNK- HYBRIDIZATION californica group v lac cal rad far phe can pal 10 3 GN oG) 2X Aa 1 12 1 14(2) 13 7 (2) 9 16 8 7(1 13 12 12 3 8 1 2 5 53 FicunE 53. Cladogram of Encelia. thing without 1, 2, and 3, asp = cal x something in the **Frutescens group" with 8, can — pal x phe, lac = ven x ?, and SC = ven x “Frutescens group" with 12. Listed above are five hypotheses of hybridiza- tion and some possible parents. Examination of the distribution patterns and other data do not support two of the hypotheses (E. farinosa and SC) and an additional one is added. An F, that has been identified as E. virginensis has been found growing with E. actoni SO that E. virgi- With E. actoni as one of the parents. Using the distributions we can narrow down the choice of possible parents to the following: asp = cal x SF, can = phe x pal, lac = ven x ?, and vir = act x ?. The reticulate cladogram is illustrated in Fig- ure 54. The hybrids indicated with solid lines are those that were Supported a as hybrids by ide the ;those with dotted lines were supported by only one of e two EXAMPLE 6. MONTANOA (ASTERACEAE) (FUNK, 1982) Montanoa Cerv. (Appendix R) has 20 species in Mexico and Central America and five in northern South America. Examining one of the equally parsimonious cladograms (Fig. 55), only one taxon (M. hexagona) shows any strong in- dication of hybridization. This taxon has the only two character losses on the cladogram. There are however, three known high level polyploids in the genus (Fig. 55). Two of these (M. revealii and M. guatemalensis) show no evidence of being hybrids. The third is M. hexagona and, it could be a hybrid between its sister taxon, M. hibis- cifolia, and something outside of the group de- fined by character 34. None of the polyploids are sympatric with any other species and they all have at least 9596 pollen viability, and during meiosis there is at least one stage where a com- plete bivalent can be observed. So, we are left with these three species being either very old polyploids, two of which have developed autapo- 702 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 4 \ F lac E Pa \ gz ven, ? rad ` phe - pal cal | act \ 2- N = I N ` l I \ \ I N I Ñ I \ I : I N [ A I \ i : l N [ N I ` I ) I / ! / I / I / I / [ / [ / / I [ [ [ I I I [ l l I I I [ [ I I l lj lj FiGuRE 54. Cladogram of Encelia with reticulations of Spilanthes Jacq. but were removed by Jansen (1981). There are 39 taxa (30 species), 16 of which orphies, or the parents are extinct so the rela- are diploids (23 polyploids; ploidy level is esti- m tionships do not show up on the diagram. Or less likely, they are autopolyploids with the dip- loids no longer extant. The cladogram cannot genus is pantropical with one species in the help us in resolving this matter. EXAMPLE 7. ACMELLA (ASTERACEAE) (JANSEN, IN PRESS) ually This last example presents the most difficult There are at least ten equally parsimonious case: one where there are more hybrids than non- cladograms of Acmella and a large number (over hybrids in a genus, where the hybrids are hy- — 100) that are only a few steps longer. Many of bridizing, where there are few characters in the these cladograms have very different structures. analysis and some of the hybrids have inherited I have selected one to discuss as a representative mostly plesiomorphies, and where the hybrids (Fig. 56), but in no way am I indicating that this are weeds that disperse readily and tend to hy- particular cladogram is to be preferred over any bridize wherever they are. Although not the rule, other. In Figure 56 there are only three apo- such situations are not that unusual in the As- morphies (excluding autapomorphies) that are teraceae family. One such genus is Acmella (Ap- — not either subsequently lost or found elsewhere pendix S). The species of Acmella used to be part on the cladogram [7, 1, 2(2)]. The three major FUNK -HYBRIDIZATION 703 55 FIGURE 55. groups of Jansen (in press; indicated in Fig. 56 by the large numbers 1, 2, and 3) are obvious on the cladogram and only one, number 2, is non- monophyletic (this group was non-monophyletic on all ofthe cladograms that I constructed). Some taxa may be hybrids between the three major groups, some of the more obvious ones are as follows: 1. A. decumbens var. decumbens (23a) may be a hybrid between some taxon in group 1 and one in group 2 because it has apomorphies 1 and 2. 2. A. poliolepidica (1) may be a hybrid be- tween a taxon in group 1 that has apomorphy 20 and a taxon in group 3 with apomorphy 16. 3. The ancestor of A. darwinii (7) and A. so- diroi (8) may have been a hybrid between a taxon in group 2 and one in group 3 that has apomor- phies 18(2) and 17(2). 4. A. paniculata (19) may be the hybrid of a taxon in group 3 and one outside of it because of its lack of apomorphy 22 (only two taxa in group 3 lack apomorphy 22). X O F H HEAT P K LE L GA B ! S 37 43 2019x434 |31 2 42 N35 46 38 39 45 $ 42 44 20 43 43 22,29,41 35,43 41 = 44 34 24 24 23 40 7 6 5 4 * 23,24,25,28 $ 28 Cladogram of Montanoa, numbers above taxa indicate ploidy level of known polyploids. Other taxa show some indication of hybrid- ization within the groups. However, there is no strong indication of hybrids and their parents within the groups because there are so many al- ternative groupings and so few characters. Some taxa are obviously hybrids or of hybrid ancestry because of their ploidy level (Fig. 56, Appendix S) but there is little indication of what their his- tory might be. The different parsimony clado- grams give us different possible hybrids and par- ents. With the exception of two or three small groups of species that appear repeatedly on many, if not all, of the cladograms there are a few ad- ditional questions, such as biogeography, char- acter evolution, or ecology that can be investi- gated using these cladograms. To a large extent we are dealing with straight character patterns. We have apparently reached the limits of cla- distics with genera such as Acmella. I say this because cladistics is merely an organized way of looking at the relevant data that have been gath- ered. If no consistent pattern develops using cla- distics then the data are responsible, not the 704 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 ~. ws r @ í nar 3 — d u 29 [3 © q* q^ o9 PD a??? a A A9 9 4 ° ° A 2 ae 42? 3° 6 av ^ T $Š A69 OP Ah A 9 q^ 90% 70°42 49946? N `x, = "m y v T UN Po A X den 24 ux w RS X«I od tes 71 M. MI I I o s x 4 =. Y j ; ç H E : ' + Tom ' I H n | um woe 38 \ I H ; q =, | f 4 + I[ E. ! |! I v l I = I tod I rpm [ I I I I I = hexaploid tetraploid 56 IGURE 56. One of the many equally parsimonious cladograms for Acmella, many of which have different F topologies. method. So the lack of resolution in genera such as Acmella is simply a reflection of the data. In the future we may be able to gain more infor- and that a cladogram can be reconstituted from a classification. In such a classification only monophyletic groups are recognized. An exam- ple using the genus Anacyclus from Figure 39 is listed below: Classification of Anacyclus without hybrids mation from genetic level research to increase the data base and obtain further resolution from cladistic analyses. CLASSIFICATION OF HYBRIDS A number of papers have been published that discuss the possibilities of classifying hybrids (Wiley, 1979; Wagner, 1980; Humphries, 1983; Humphries & Funk, 1984; Nelson, 1973) so the alternatives need not be discussed in this paper. As discussed in Humphries and Funk (1984), I prefer the method called phyletic sequencing or the annotated Linnean Hierarchy. This method works on the basic principle that all information from a cladogram is available in a classification Anacyclus A. pyrethrum A. monanthos A. maroccanus A. radiatus Clavatus species group A. linearilobus A. nigellifolius The hybrids can be added in several ways; one FUNK- HYBRIDIZATION 705 1985] is to make them the sister taxon of either one of the parents. Using the first parent in the phyletic sequence we arrive at the following classification: Classification of Anacyclus with hybrids Anacyclus Sect. Pyrethraria A. pyrethrum +A. officinarum sedis mutabilis (A. pyrethrum x radiatus Sect. Anacyclus Clavatus species group A. linearilobus A. homogamos +A. inconstans sedis mutabilis (A. omogamos x clavatus) +A. valentinus sedis mutabilis (A. monogamos x clavatus A. clavatus A. latealatus A. nigellifolius The special notations include a plus sign (+) for hybrids, the parental species listed are the hy- brids in parentheses, and the latin phrase sedis mutabilis (“changeable position") that is used to mean a polytomy in the cladogram. The clado- gram is recovered in the following manner. Sec- tion Pyrethraria is the sister group of sect. An- acyclus, and A. pyrethrum and A. officinarum are sister taxa within sect. Pyrethraria (Fig. 39). In succession; A. monanthos is the sister taxon of the remaining species; A. marocannus is the sister taxon to the remaining species; A. radiatus is the sister taxon to the remaining species; the “clavatus group" is the sister group to A. /ateala- tus and A. nigellifolius (Fig. 39); within the *'cla- vatus group," A. /inearilobus is the sister taxa of A. clavatus and A. homogamos and their two hybrids; and A. homogamos is the sister taxon of A. m but forms a polytomy with the two hybri I add to "m the provision that should the two parents occur in different subgeneric groups (or different genera) then the hybrid should be listed in both groups. CONCLUSION In general, phylogenetic systematics can be used to help identify possible hybrids and their par- ents for further study. However, several condi- tions exist in some groups that make a cladistic analysis more difficult. Such conditions include the following: 1) having an increased percentage of species within a group that are hybrids, 2) 3) having hybrids that hybridize with one another, 4) having species that can reproduce asexually, and/or 5) having introgression occur. LITERATURE CITED ADAMS, E. N., III. 1972. Consensus techniques and the comparison of taxonomic trees. Syst. Zool. 21: 397. BREMER, K. 1983. Angiosperms and Lip ie systematics: some problems and examples. Ver Naturwiss. Vereins Hamburg 26: 343-354 . WANNTORP. 1977. Phylogenetic sys- tematics in botany. Taxon 27: 317-329. 79. Hierarchy and reticulations in systematics. Syst. Zool. 28: 624-627. CLARK, C. 1982. Relationships between experimental ~ phylogenetic 1 an overview. Bot. mer. Misc 8. Ex m J. 983. Numerical Taxonomy. Spring- dw Berlin, Heidelberg, New York, Tokyo. Fu . A. 1981. Special concerns in iar eg cae phylogenies. Pp. 73-86 in V. A. Funk & D. R. Brooks (editors), Advances in Cladistics: Pro- The systematics of Montanoa (Aster- aceae, Heliantheae). Mem. New York Bot. Gard. 36: Goines P. 1979. Fed in angiosperms: monocotyledons. Jn Lewis (editor), Poly- ploidy: Biological Relevance. Plenum Press, New York. GRANT, V. 1953. The role of hybridization in the evolution of the leafy-stemmed Gilias. Evolution 51-64. ] aid Genetic and taxonomic studies i in Gil- ia.X pw E iiw Gilias, Aliso 5: 479— 507 HENNIG, W. Phylogenet Systematics. Univ. of Illinois Press, Urban HUMPHRIES, i J. 1979. re revision of the genus An- acyclus L. (Compositae: a Bull. Brit. Mus. (Nat. Hist.), Bot. 7 1981. Cytogenetic py cladistic studies in Anacyclus L. (Compositae: Anthemideae). Nordic J. Bot. Di Pri rimary data in hybrid analysis. Pp. 89-103 in N. I. Platnick & V. A. Funk (editors), Advances in Cladistics: Proceedings of the Second Meeting of the Willi E Society. Columbia Univ. Press, New Y V. A. FUNK. 1984. Cladistic methodology. Pp. d in V. H. Heywood & D. M. Moore (editors), Current Topics in Plant Taxonomy. Ac- ademic X Es JANSEN, R. K. Systematics of Ud LU positae: Tetas Syst. Bot. 6: 231- 706 In press. Systematics of € (Asteraceae, Heliantheae) Syst. Bot. Mon ; Lewis, W al 9. Polyploidy i in angiosperms: di- cotyledon n W. H. Lewis (editor), Polyploidy: Biological "Mein Plenum Press, MARGUSH, . R. McMonnis. 1981 n-trees. Bull. Math. Biol. 43: 239-24 McMonnis, F. R., D. B. MERONK & D. ^i NE EUMANN. 19 A view of some consensus methods for trees. Pp. 122-126 in J. Felsenstein (editor), Nu- merical Taxonomy. Springer-Verlag, Berlin, Hei- delberg, New York, Tokyo MiCKEVICH, M.R. 1978. Taxonomic congruence. Syst. Zool. 27: 143-158. NELSON, G. 1973. Classification as an expression of phylogenetic relationships. Syst. Zool. 22: 344- 3 or : , Consensus à 1979. Cladistic analysis and synthesis: prin- ciples and definitions, with a historical note on Adanson's Familles des Plantes (1763-1764). Syst. Zool. 28: 1-21. 3. Reticulation i in reir Pp. 105- 111 in N. I. Platnick & V. A. Funk (editors), Ad- vances in Cladistics: Proceedings of the Second Meeting of the Willi Hennig Society. Columbia Univ. Press, ew Yor TNICK. 1980. Multiple branching in cladograms: two interpretations. Syst. Zool. 29: 86-91. NEUMANN, D. A. 1983. Faithful consensus methods for n-trees. Math. Biosci. 63: 271-287. Rosen, D. E. 1979. Fishes from the uplands and intermontane basins of Guate mala: dC el y ]. Amer. Mus. Nat. Hist. 162: 267-376. SANDERS, R. W. 1981. cpu» analysis of Agastache (Lamiaceae). Pp. 95- in V. A. Funk & D. R. Brooks (editors), he eens in Cladistics: Proceed- ings of the First Meeting of the Willi Hennig So- ciety. New York Botanical Garden, New York. ScHuH, R. T. & J. T. PoLHEMUS. 1980. Analysis of taxonomic congruence among morphological, ecological, and FR er woe an sets for the Lep p t. Zool. 29: 1- 26. SEAMAN, F. C. & V. A. FUNK. 1983. Cladistic analysis of complex natural products: developing transfor- mation series from sesquiterpene lactone data. Taxon 32: 1-27. SEMPLE, J. C. A Men, of the goldenaster genus Chrysopsis (Nutt.) Ell. nom. cons. (Com- positae — Astereae). rae 83: 323-384. bs is ao 1984. Observations on rphology and ecology of Brad- puria P eee uas Astereae). Syst. Bot. 9: 95-101. SOKAL, R. e eh 1981. Taxonomic con- gruen n the re eer re-examined. Syst. Zool. 30: 304- STEBBINS, G. 74. wering Plants: Evolution Abov athe Species pas Belknap-Harvard Press, . J. ROHLF. am WAGNER, W. 54. Reticulate — in the Appalachian Aspleniums. a n 8: 103-108. 1969. The role and taxonomic treatment of hybrids. BioScience 19: 785-7 89. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 . 1980. Origin and philosophy of the ground- plan divergence method of cladistics. Syst. Bot. 5: l 3. 1983. Reticulistics: the recognition of hah and their role in cladistics oy ——— 79 in N. I. P e Willi He ennig Society. Columbia w York. E. 1983. men cladograms and the identification of hybrid taxa. Pp. 81-88 in N. I. Platnick & V. A. Funk (editore. Advances in Cladistics: Proceedings of the Second Meeting of the Willi Hennig Society. Columbia Univ. Press, York WiLEv, E. O. 1979. An annotated Linnaean hier- archy, with comments on natural taxa and com- peting systems. Syst. Zool. 28: 308-337. — & D. R. Brooks. 1982. Victims geri libri t. Zool rr 31: 1-24. APPENDICES A-S. Only the apomorphies are indi- cated in the data matrices because in phylogenetic sys- tematics only the apomorphies are used to group taxa. APPENDIX A. Data matrix for Figure 2. Apomorphies Taxa l 2 3 4 5 6 7 l l l l l l l l l l l m g O = > APPENDIX B. Data matrix for Figures 3—6. Apomorphies APPENDIX C. Data matrix for Figure 7. Apomorphies 1985] FUNK — HYBRIDIZATION 707 APPENDIX D. Data matrix for Figures 8-12. APPENDIX I. Data matrix for Figures 23-25. Apomorphies Apomorphies Taxa 1 2 3 5 Taxa 1 2 3 4 5 6 7 8 9 1011 12 A 1 1 1 A 1 1 1 B 1 1 l B l 1 C l C l 1 1 H1 1 1 l l D l 1 1 1 1 H2 1 1 l l E l 1 1 1 1 F l 1 1 1 1 1 1 G l 1 1 1 1 1 1 1 APPENDIX E. Data matrix for Figure 13. H ll 11 1 1 11 ! I l 1 1 1 l 1 1 Apomorphies Taxa l 2 3 5 1 1 | 1 1 — — — — APPENDIX F. Data matrix for Figures 14-17, 19. Apomorphies Taxa 1 2 3 4 5 6 8 9 A 1 1 1 B 1 1 1 l C 1 1 l l D l l E l l l H l l 1 1 l APPENDIX G. Data matrix for Figure 18. Apomorphies Taxa 1 2 3 4 5 6 8 9 A 1 1 1 B l 1 1 1 C 1 1 1 l D l l E 1 l l H l APPENDIX J. ay R. Br. p apri = 4 Abbreviations. — A. M. incanum Decn gitubum janan —C. P burchellii N. E. ok i — D. M. um.—E. M. sei heli —F. M. pilis oe M. spinosum N. E. Brown.—H. M. M ard N. E. Brown.—I. M. la s biis Charac- ters. — Data published in Bremer and Wanntorp (1979) and Humphries (1983) but no character list was fur- nished in either publication. Data matrix. — For Figures 26, 27, 36-38 Apomorphies Taxa 1 2 3 4 5 6 N A l B 1 1 1 C 1 1 1 1 1 1 1 D 1 1 1 1 1 1 E 1 l 1 1 1 F 1 1 1 1 G l l 1 1 H 1 l l I 1 1 l APPENDIX K. Data matrix for Figures 28, 29. APPENDIX H. Data matrix for Figures 20-22. Apomorphies Taxa 1 2 3 4 UA — p — — Apomorphies Taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 A 1 1 1 1 1 B 1 1 1 1 1 € 1 1 1 1 D 1 1 1 E 1 1 1 F 1 1 1 1 G 1 1 1 1 H 1 1 1 1 708 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 APPENDIX L. Data matrix for Figures 30, 31. APPENDIX M. Data matrix for Figures 32, 33. Apomorphies Apomorphies Taxa 1 2 3 4 S 6 7 8 9 Taxa 1 2 3 4 5 6 7 8 9 10 11 I2 13 A l l l A l 1 1 1 1 1 l B l l 1 1 B l l 1 1 1 I 1 C l l l l C ll 1 1 | 1 l D l l l D l 1 l E l l l E 1 1 1 H l l l l l H | 1 l l l l APPENDIX N. í L. (Asteraceae, Anthemideae). Abbreviations.—A. A. pyrethrum (L.) Link var. pyrethrum.—B. A. pyrethrum (L.) Link var. ie (Ball) Maire. — C. A. monanthos (L.) Thell.—D. A. maroc- canus (Ball) Ball.—E. A. radiatus Loisel.—F. A. coronatus (Murb.) Humphries.—G. A. clavatus (Desf.) Pers.— H. A. homogamos (Maire) Humphries. — I. P valentinus L.—J. A. inconstans Pomel.—K. A. linearilobus Boiss. & Reuter.—L. A. /atealatus Hub.-Mor.—M. A. nigellifolius Boiss.—N. A. officinarum Hayne. Characters.— Published in Humphries (1979). Data matrix.— Lower case letters represent groups of characters that display that pattern so that there are five apomorphies that have the distribution patterns of a, etc. For Figures 39-43. Apomorphies b c d e f g k l m n o p q (3) (3) G (0 G ( D G) 6) (4 (Q0 (1 l l l l l l — um — M — — ~~ ~ -— — ~ — p ~~ ~ — xw — — — — — — — — — — — — — — — — — — — — — m — m - — — tr Z m abu an — — — — — — — — — — — — — — — — — — — — — — — — — — — — — FUNK—HYBRIDIZATION 709 1985] ‘Aydiowotsajd pue Aydiourode ay} u99A19q SILILA e c I 4C I 1 1 du — — — — — — — — — — — — — Cl _ I I I 1 I I 202 I I I el I I I iC 9€ SE PE EE Z€ TE OE 6% 8C LC 9C SC vC ET CC IZ Oc BI LI OI Hl €I CI I OL 6 8 9 S v t C sorydiowody S > qç E '9p—pp samy J04—^ on geq 'sisA[eue sty} ur posn jou uoxej oyroodsqns e 10j AÁqd1ourodejne ue sem I 9sneooq L jn vd jo [£^ 0u121 əy} 10} 1d90x9 (186 D: V oe APUAN 29 uoloo ^ DAE "muss Y u4— es (9u2211)) s14js2dn4 710 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 72 APPENDIX P. o e Eu Ell. and Bradburia Semple & Chinnappa (Asteraceae). Abbreviations. — BR. B. hirtella T. & G.— C. floridana Small.—gd-g. C. godfreyi Semple f. godfreyi.—gd-v. C. aaa Semple f. viridis Semple.—gs-c. C. gossypina (Michx.) Elliot subsp. cruiseana (Dress) ioe —gs-gd. C sypina (Michx.) Elliot se gossypina f. decumbens (Chapm.) Godfrey.—gs-gg. C. gossypina (Michx.) Elliot subsp. gossypina f. gossypina. —gs-gt. C. gossypina (Michx.) Elliot subsp. gossypina f. trichophylla (Nutt.) Semple. — gs-h. C. gossypina (Michx.) Elliot subsp. Ayssopifolia (Nutt). Semple.—lan. C. lanuginosa Small.—lat. C. /a- tisquamea Pollard.—li-d. C. linearifolia Semple subsp. dressii id —li-l. C. eh ien Semple subsp. /i- nearifolia. — mar. C. mariana (L.) Elliot. —pil. C. pilosa Nutt.—sca. C. scabrella T. & G.—sub. C. subulata Small. Characters — Original characters furnished by Semple and db slightly by Funk. Transformation Series Plesiomorphic Character Apomorphic Character 2A. Growth form biennial perennial 2B. Growth form biennial annual 4. Pubescence of basal rosettes wooly pilose 6. Achene body no translucent ribs translucent ribs 7A. Outer pappus bristles narrow bristles broad 7B. Outer pappus present absent 8. Upper cauline leaves wooly not wooly 9. Glandulation of upper cauline sparsely densely leaves 10. Upper leaf base sessile, nonclasping clasping 11. Upper leaf size not greatly reduced P tue 12. Glandulation of peduncles glandular eglan 13. Upper leaf shape elliptic ag 14. Upper leaf apex obtuse acute 15. . Glandulation of phyllaries densely eglandular 16. dau ld phyllaries glabrate pubescent 17. Phyllary si narrow roa 18. Phylla mi acute-attenuate long, subulate 19. Outer w w pas size much shorter than similar in size inner phyllaries 20. Head orientation in bud erect nodding 21. Inflorescence t open cymose subumbellate 22. OMM Ur ERE branch few many 23. Disc flor hemaphroditic staminate Data matrix. — For Figures 47—50. Apomorphies Taxa 2A 2B 4 5 6 7A 7B 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 l 1 l 1 l l 1 ! l £e las — — _ — — — — — — — _ 1985] FUNK—HYBRIDIZATION 71 1 APPENDIX Q. Encelia Adanson (Asteraceae). Abbreviations.—act. E. actoni Elmer.—asp. E. asperifolia (S. F. eqe Clark & Kyhas.—cal. E. californica Nutt. —can . E. canescens Cav.—far. E. farinosa Gray.— j wass Gray.— GC. Und it Canyon (C. Clark, pers. comm.).—lac. E. waspa dán se.—pal. E. palmeri Vasey & Rose.—phe. E. farinosa Gray var. phenicodonta I. M. Johnsto . E. Main: Brandegee.—rav. E. ravenii Wiggins.—res. E. resinosa Brandegee. — SC. Undescribed taxon pius Pi- cachos de Santa Clara, Baja (C. Clark, pers. comm.). — SF. Undescribed taxon from San Felipe (C. Clark, pers. comm.).— ven. E. ventorum Brandegee.— vir. E. virginensis Nelson. Characters. — To be published by Clark (pers. comm.). Data matrix.— For Figures 51-54. Apomorphies Taxa l 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 E: — M — — — — — — — — — — — 2a a Varies between apomorphies 2 and 1. 712 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 72 APPENDIX R. Montanoa Cerv. (Asteraceae). s —AN. M. andersonii McVaugh.—AG. M. an- gulata Badillo. — AT. M. ud uie ) Sch —B. M. bipinnatifida (Kunth) K. Koch.—E. M. echinacea S. F. Blake.—F. M. fragrans Badillo. . M. und —GA. M. grandiflora DC.—G. M. guatemalen- sis Robins. & Greenm.—HE. M. hexagon Robins. & Greenm. —H. M. hibiscifolia Benth. —lI. M. imbricata V. A. Funk. —J. M. josei V. A. Funk.— pi pisi —L. M. laskowskii McVaugh. —LE. M. leucantha (Lag.) S. F. Blake. —LI. M. liebmannii m Bip.) S. F. Blake. — M. M. mollissima shai —O. M. ovalifolia C.—P. M. pteropoda S. F. Blake.—Q. M. quadrangularis Sch. Bip.—R. M. revealii H. Robinson.—S. speciosa DC.—ST. M. standleyi V. A. Funk. — T. M. tomentosa Cerv. Erud bt Published in Fank (1982). Data matrix.— For Figure 55. Apomorphies l 2 3 4 5 6 7 8 9 10 1» 12 13 14 15 16 17 AN 1 1 1 1 1 9 20 21 22 so = CRTI Q Q m > p — — — — — — — — — — — — — uu d4ávzxovozct 1985] APPENDIX R. Continued. FUNK—HYBRIDIZATION 713 Apomorphies 23 24 25 26 27 28 29 30 31 32 34 35 36 37 38 39 40 41 42 43 44 45 46 47 1 1 l 1 l 1 1 I l l 1 1 l 1 1 | l l 1 l l l l l l l l I l l l l l l l l l 1 |] 1 1 l l 1 l l I 1 l I l l l 1 |! l 1 1 l l l 1 l l l 1 I l 1 l l 1 l l 1 |] 1 1 l l l l l 1 l l l 1 l l 1 1 1 l l 714 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 APPENDIX S. Acmella Rich. ex Pers. (Asteraceae). Abbreviations (* indicates chromosome number is an estimate from pollen diameter). — 1. A. poliolepidica (A. H. Moore) R. K. Jansen (4x). —2. A. pilosa R. K. Jansen (2x).—3a. A. aaa sm i uk var. repens (Walt.) R. K. Jansen (4x).—3b. A. oppositifolia Rich. var. opposi- Lips (2x, 3x, 4x, 6x).—4. A. ciliata (H.B.K.) Cass. (6x).—5. A. caulirhiza Delile (2x).—6. A. calva (DC.) R. K. Jansen P^ 24. A. pes: (D. M. Porter) R. K. Jansen (6x*).—8. A. sodiroi (Hieron) R. K. Jansen 6x).—9. A. ramosa (Hensl.) R. K. Jansen. (2x).—10. A. pusilla (Hook. & Arn.) R. K. Jansen (2x, 4x).—11. A. lundellii R. K. Jansen (2x*).—12a. A. papposa (Hemsl.) R. K. Jansen var. papposa (4x).—12b. A. papposa (Hensl.) R. K. Jansen var. macrophylla scr K. Jansen (4x). — 13. A. iodiscaea (A. H. Moore) R. K. Jansen (4x).—14. A. uliginosa (Sw.) Cass. (4x).—15a. A. filipes (nens R. K. Jansen var. filipes (2x*),— 15b. A. filipes (Greenm.) R. K. Jansen var. cayensis R. K. Jansen (2x*).—15c. A. filipes (Greenm.) R. K. Jansen var. parvifolia (Benth.) R. K. Jansen (2x). —16a. A. alba (L'Herit.) R. K. Jansen var. alba (6x*).— 16b. A. alba (L'Herit.) R. K. Jansen var. ecuadorensis R. K. Jansen (6x*).—18. A. leucantha (H.B.K.) R. K. Jansen (6x*).— 19. A. paniculata (DC.) R. K. Jansen (4x*).—20a. A. radicans (Jacq.) R. K. Jansen var. radicans (6x). —20b. A. radicans (Jacq.) R. K. Jansen var. devilis (H.B.K.) R. K. Jansen (6x). — 21. A. brachyglossa Cass. (6x).—22a. A. grandiflora (Turcz.) R. K. Jansen var. grandiflora (4x*).—22b. A. grandiflora (Turcz.) R. K. Jansen var. brachy- glossa (Benth.) R. K. Jansen (4x).—22c. A. grandiflora (Turcz.) R. K. Jansen var. discoidea R. K. Jansen (6x*).— 23a. A. decumbens (Smith) R. K. Jansen var. decumbens (Ax). — 23b. A. decumbens (Smith) R. K. Jansen v affinis (Hook. & Arn.) R. K. Jansen (2x).—24. A. leptophylla (DC.) R. K. Jansen (2x).—25. A. pelidioides (Smith) R. K. Jansen (2x).—26. A. grisea (Chodat) R. K. Jansen (2x).—27. A. serratifolia R. K. Janse 28. A. alpestris (Griseb.) R. K. Jansen (2x).—29. A. psilocarpa R. K. Jansen (2x*).—30. A. glaberrima RR R. K. Jansen (2x). Characters. — Taken from Jansen (in press) with the following changes: transformation series 13-15 were eliminated because I felt they were the same as 20-22; transformation series 1 1 and 12 were combined into one transformation series with two independent apomorphies. Data matrix. — The apomorphies of trans- formation series 8, 11, and 21 were all Hes ss independent of one another while in transformation series 2; 17, and 18, apomorphy li t p p p phy 2. For Figure 56. Apomorphies - = > b -— N w A cA a N oo Ne) — o - e - an — - _ oo — " t2 ° N — N N gom — — — — — — — — Www Ww — — — — o0 —1 O. Q + Q Q h2 = — NNN — N N —_ — — — — — — — — — tet k. os e — 0 M a E E og — N N N — — — — — — — — N - — N — — NNNNN —= NON = — NNN NY — 1985] APPENDIX S. Continued. FUNK—HYBRIDIZATION 715 Apomorphies 9 10 21 22 pi — — — — — — — pb pb w w w w w w w o2 PHYLOGENETIC ANALYSIS OF SEED PLANTS AND THE ORIGIN OF ANGIOSPERMS! PETER R. CRANE? ABSTRACT Principles of phylogenetic analysis (cladistics) are introduced with an examination of relationships among extant genera of Gnetales netales can be supported as a monophyletic group, wit Gnetum and Welwitschia more closely ne to each other than either is to Ephedra. Characters of the progymnosperm Archaeopteris and 19 extinct and extant seed plant taxa are then reviewed as a basis for a cladistic analysis of their interrelationships. The seed plant taxa included are: medullosan cycads, Lyginopteris, Cordaixylon, Mesoxylon, Lebachia, extant conifers, Ginkgo, Callistophyton, peltasperms, glossopterids, Caytonia, corystosperms, Bennettitales, Pentoxylon, Gnetum, Welwitschia, w tales, and corystosperms will be of maximum interest in further elucidating the phylogenetic Sues: of flowering plants. The origin of flowering plants is one of the of the century and reflected a natural extension major unsolved problems of plant phylogeny and an enigma in which most major groups of vas- cular plants have been implicated. Together with the related question of the origin of the angio- sperm flower it has generated a body of literature unrivalled in phylogenetic botany for its size and diversity of opinion. The suggestion that Mag- nolia and related taxa are the most primitive living angiosperms (Arber & Parkin, 1907; Bes- sey, 1897, 1915) originated at around the turn tion of the flower as a reduced bisexual axis bear- ing ovules and pollen organs on modified leaves (Anthostrobilus or Euanthial theory, Arber & Parkin, 1907) was similarly an extension of the classical idealistic morphology of the nineteenth century. Although the magnoliid theory has nev- er been without its competitors (e.g., Croizat, ! This paper is dedicated to the late Dr. P. D. W. Barnard, who first encouraged my interest in paleobotany. The work was su useful risiede with the late P. D. W. Ba upra W. L. Cr the reconstuctions of fossil plants, and W. Kovac 2 g oO g Illinois 60605. ANN. Missouni Bor. GARD. 72: 716-793. 1985. pported in part by National Science Aper grant BSR-8314592, and has benefitted from rnard, and w k, M. N. B W. E. artment of Geology, Field Muséum of Natural History, di Road at Lake Shore Drive, Chicago, 1985] 1960; Engler, 1897; Meeuse, 1966; Melville, 1962, 1963; Rendle, 1925, 1930; Sporne, 1971b; Strasburger et al., 1898; Wettstein, 1935), com- parative studies of extant flowering plants par- ticularly over the last 20 years (Bailey, 1944; Bailey & Swamy, 1951; Cronquist, 1968, 1981; Dickison, 1975; Eames, 1961; Hickey & Wolfe, 1975; Raven, 1975; Stebbins, 1974; Takhtajan, 1969, 1980; Walker, 1974a, 1974b, 1976; Walk- er & Doyle, 1975; Wodehouse, 1935, 1936) have reinforced the view that the Magnoliidae sensu lato (Cronquist, 1968, 1981; Takhtajan, 1969, 1980) and perhaps certain monocotyledons (Burger, 1977) are a heterogeneous assemblage of plants that retain a relatively large number of primitive characters for flowering plants as a whole. This conclusion now has been corrobo- rated broadly by: the Qocumenoe of the predicted adaiospetim fossil record (Doyle, 1969, 1978; Doyle & Hickey, 1976; Hickey & Doyle, 1977; Muller, 1970, 1981, 1984; Sporne, 1974; Upchurch, 1984; Walker & Walker, 1984; Wolfe et al., 1975). The sequential stratigraphic ap- pearance of progressively more complex leaves and pollen (Brenner, 1963; Doyle, 1969; Doyle & Hickey, 1976; Hickey & Doyle, 1977; Hughes, 1976; Muller, 1970, 1984) has been interpreted as documenting a major mid-Cretaceous flow- ering plant radiation beginning in the Barremian or earlier in the Lower Cretaceous (Brenner, 1984; Trevisan, pers. comm.). Palynological evidence suggests that this radiation began at low, tropical, paleolatitudes and was followed by predomi- nantly poleward migration (Brenner, 1976; Hickey & Doyle, 1977). Reports of pre-Creta- ceous angiosperms either have been rejected or must be regarded as of uncertain relevance due to lack of critical information (Doyle, 1978; Hill & Crane, 1982; Scott et al., 1960). Reconstruc- tions of some early angiosperms from several dispersed organs (Dilcher & Crane, 1984) and studies of flowers and other reproductive struc- tures (Basinger & Dilcher, 1984; Crane & Dil- cher, 1984; Dilcher, 1979; Krassilov et al., 1983; Vakhrameev & Krassilov, 1979) also have pro- vided important insights into the systematic re- lationships, floral morphology, and rep biology of some early flowering plants. Despite this success in clarifying the timing and pattern ofthe early flowering plant radiation, oO the fundamental question of the we po- of sition of angiosperms in the overall schem seed plant evolution remains unanswered. a CRANE-SEED PLANT PHYLOGENETICS 717 have claimed that we are no closer to a consensus as to which group of gymnosperms might be most relevant than we were at around the turn of the century (Beck, 1976; Harris, 1960; Hughes, 1976). Discussions of this problem have generally cen- tered on two D issues: the origin of the fflowering enerm carne! Bisnis. There have been many hypotheses as to the derivation of the angiosperm carpel from the ovulate reproductive structures of living and fos- sil gymnosperms. The most common proposi- tion derives the conduplicate carpel from the ovulate structures of pteridosperms (seed ferns) sensu lato: either Caytonia (Andrews, 1963; Doyle, 1978), corystosperms (Stebbins, 1974), Glossopteris (Retallack & Dilcher, 1981), or even Carboniferous seed ferns (Long, 1966, 1977). Al- though none of these hypotheses has received universal acceptance, they have contributed to a widespread view that the outer integument ofthe normally bitegmic angiosperm ovule is homol- ogous to the cupule of seed ferns (Smith, 1964), and that the pteridosperms are the ancestors of flowering plants (Cronquist, 1968, 1981; Knoll & Rothwell, 1981; Stewart, 1983) Much of the literature reviewed above implies that solution of the angiosperm problem must necessarily involve the recognition of angio- sperm ancestors (e.g., Mabberley, 1984). This is a position confronted by theoretical and practical difficulties. First, if our current understanding of evolutionary processes is accurate, then it is species, not higher taxa, that are the units of evolution, and thus only species and not higher taxa can be truly ancestral (Eldredge & Cracraft, 1980: 114); and second, ancestral groups cannot be defined by features that they alone possess, because by definition these features also char- acterize members of the descendant group (Wi- ley, 1979: 212-214). Under such circumstances it is inevitable that many ancestral groups have been construed loosely and used broadly to in- dicate inferred proximity of descent. À system in which groups are not clearly defined, and sim- ilarity is hazily interpreted as phylogenetic re- lationship, may have heuristic value, but inev- itably it is only capable of nis broad, relatively unspecific ifficult to compare. It does not fücilitate 2. iei and thus critical, evaluation of alternative sugges- tions of relationship (Hill & Crane, 19 Phylogenetic systematics (Hennig, 1966; often treated as synonymous with cladistics) offers an 718 alternative approach to the question of angio- sperm origins. It can provide a relatively explicit assessment of relationships from which straight- made and a a methodological framework i in which com- peting theories of relationship may be compared usefully. It also provides an alternative to the search for ancestral groups, allowing phyloge- netic problems to be formulated in a different and more tractable fashion. In cladistic terms the problem of angiosperm origin is to recognize and define the major groups of seed plants, to deter- mine their phylogenetic interrelationships, and thus to establish with which group of gymno- sperms the flowering plants share a most recent common ancestor (Crane, 1984). The large-scale phylogenetic relationships of seed plants have been the subject of very little critical discussion, and there is little agreement on how the major taxa should be grouped to- gether. Many recent classifications have either adopted or modified the views of Coulter and Chamberlain (Chamberlain, 1935; Coulter & Chamberlain, 1917) formulated over half a cen- tury ago (Bierhorst, 1971; Cronquist et al., 1966; Foster & Gifford, 1974; Sporne, 1971a). Other authors have merely treated all the major groups as of equivalent rank (Bold, 1973; Bremer & Wanntorp, 1981; Taylor, 1981a). The late Pro- fessor Tom Harris succinctly summarized the situation when he commented (1976: 133) that the “‘classes” of gymnosperms were no longer the “branches of a single phylogenetic tree," but “a frr ward relationships, and establish a coherent frame- work within which the position of flowering plants may be assessed. There have been two previous attempts to evaluate relationships within seed plants using cladistic techniques. Both were preliminary and intended in part to stimulate future work. Parenti ized by Richardson (1982) (see also reply by Parenti, 82). The principal difficulties with her analysis d i heterogeneity, and hence mono- hyly, of some taxa, and the level of detail of character PPAR Hill and Crane (1982) pre- sented an analysis based solely on extant seed plants using living Equisetum, lycopods, and ferns as out-groups. Our analysis also encountered dif- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 ficulties with character definition and recogni- tion. Fifty characters were surveyed but yielded several alternative solutions that, although dra- matically different, were almost equally parsi- monious. This paper extends and attempts to improve on the analysis by Hill and myself. It core and redefines many of the characters mployed, includes additional characters, and i E for the first time some of the better understood fossil gymnosperms. In the absence of detailed cladistic analyses of the larger gym- advanced and primitive characters in those taxa. The paper is divided into four major parts. First, the principles of cladistics are introduced briefly and illustrated by a simple analysis of relationships between the three extant genera of Gnetales. Second, the characters of extant, and sufficiently well known fossil, seed plants are re- viewed and phylogenetic relationships are as- sessed. Third, an overview of seed plant phylog- eny is presented and compared with previous classifications and phylogenetic hypotheses. Fi- nally, the possible phylogenetic relationships of other more poorly understood plant fossils are reviewed and problems for future research are discussed. Throughout, it should be noted that higher taxa are referred to as far as possible by informal names that have no implication oftaxo- nomic rank. The primary concern is to clarify relationships, and I make no suggestions as to how these should be summarized as a formal nomenclatural scheme. Fossil taxa are treated as “whole plants.” Justification for individual cases is given in the text. In the descriptive sections the megasporangiate reproductive structures are out for simplicity of comparison, but it is by no logues. Individual cases are discussed in the text. The numbers assigned to characters are prefixed by the number of the table in which they are listed. PHYLOGENETIC RELATIONSHIPS IN THE GNETALES—A THREE-TAXON PROBLEM PRINCIPLES In a cladistic analysis each of the taxa under consideration should be a monophyletic group. 1985] As far as I know no one has seriously questioned the naturalness and, by implication, the mono- phyly of any of the three extant gnetalean genera. Welwitschia is in any case monotypic. It is a bizarre xerophyte occurring only in restricted desert areas of southwest Africa (Chamberlain, 1935). Gnetum comprises about 40 species of predominantly tropical lianes but also includes a few trees and shrubs (Sporne, 1971a). Ephedra comprises about 40 species of “switch” shrubs with less common tree-like and vine-like forms (Sporne, 1971a). The genus occurs in xeric hab- itats at low, but generally extra-tropical, lati- es. Not all of the characters of these genera can contribute to resolving relationships within the Gnetales. Features restricted to a single genus may account for its distinctiveness but are clearly of no value for interpreting relationships with other genera. Similarly, features that occur in all u to defining inter-generic relationships. ters that occur in two of the three extant genera may provide an indication of relationship but are of two kinds: those that could define an ex- clusive group of two gnetalean genera in which no other taxon would be included, and less re- stricted characters that could only define a group in which non-gnetalean taxa would also have to be included. Characters of the first kind suggest that two gnetalean genera are more closely re- lated to each other than they are to any other taxon, while characters of the second kind merely indicate that the two genera are no more closely related to each other than either is to some non- gnetalean plant. Characters of the second kind are of no value for assessing phylogenetic rela- tionships among the three gnetalean genera, and cladistics attempts to recognize and utilize only characters of the first kind. This is accomplished Crane, 1982: 280; Stevens, 1980, 1981; Watrous & Wheeler, 1981). Theoretically the out-group for the Gnetales could be all other organisms, but for practical purposes a more manageable out-group could be all other seed plants or all other vascular plants. In this example all seed plants, with the exception of flowering plants, are used as the out-group. The effect of incor- porating flowering plants is considered later in this paper. To assess whether the tetrasporic megagame- CRANE-SEED PLANT PHYLOGENETICS 719 tophyte of Welwitschia and Gnetum defines an exclusive group, and therefore suggests close re- lationship, the distribution of the tetrasporic and monosporic condition is examined in all seed plants. All other seed plants (excluding some angiosperms) are monosporic. The tetrasporic condition is relatively restricted and defines an exclusive group (Welwitschia plus Gnetum). Conversely, the cellular female gametophyte seen at fertilization in Welwitschia and Ephedra is merely an expression of the that occurs in all other seed plants except Gnetum and an- giosperms. In Gnetum the female gametophyte is partially free nuclear at the time offertilization. The character is therefore of no value for as- sessing relationships between the gnetalean gen- era. A E +h 1:14 í 1 Ly anes f Al "1 by out-group comparison inevitably involves a logical regress in that decisions can be made only by reference to groups already circumscribed (perhaps wrongly) by the distribution of yet other characters (Hill & Crane, 1982: 280). However, the process can be used to develop a hypothesis about which of a pair of contrasted characters is more, or less, restricted. This is open to test by broadening the analysis still further. When ap- plied to a variety of characters these assessments of generality result in a character hierarchy that efines groups within groups, for example, a mul- ticellular embryo developing within an arche- re) cheids and/or vessels (tra plants, nd or a megapro four to 16 nuclei (angiosperms) (Fig. n Ti this way relationships can be defined in relative terms (Hennig, 1965: 98). Ginkgo and Pinus both have seeds (spermatophytes) and are therefore more closely related to each other than either is to Lycopodium (a pteridophyte), which is free spor- ing and lacks seeds. All characters may be useful for defining groups at some level in the hierarchy (Wiley, 1981: 126), but at any given level of inclusiveness (univer- sality) some characters will be useful for group definition, whereas others will not. The imple- mentation of this distinction is the fundamental difference between cladistics and phenetics. At any level in the hierarchy the phenetic concept of “overall similarity” is separated into a gen- eralized, cladistically non-definitional compo- nent (symplesiomorphy), and a less generalized, definitional component (synapomorphy). In practice recognizing apparent synapomor- 720 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 EMBRYOPHYTES [ 1] TRACHEOPHYTES | 1 UNNAMED GROUP | | Bryophytes Pteridophytes Progymnosperms FIGURE 1. die Groups definable by presence of a single c in this diagram are give they ar and Churchill (1984). phies and identifying their distribution seldom produces a single unambiguous character hier- archy. Different characters often indicate con- flicting groups and hierarchical patterns. Apply- ing the principle of parsimony, it is the simplest of competing patterns that is preferred. In ef- fect, parsimony minimizes the conflict between apparent synapomorphies and therefore the con- flict between apparent groups (Farris, 1982; Wi- ley, 1981). In cladistic terms the “true synapo- morphies" recognized in this way are homologies, and the problems of determining polarity, syn- apomorphy, and homology can therefore be viewed as synonymous (Eldredge & Cracraft, 1980: 35-40; Hill & Crane, 1982: 282-287; Pat- terson, 1982a; Stevens, 1984). The result of a cladistic analysis is therefore merely a pattern of nested synapomorphies. It defines relationships in a relative way and like any other hierarchical pattern may be summarized in a tree-like dia- gram. There is no necessary relation to the theory of evolution and no necessary time axis. SPERMATOPHYTES ANGIOSPERMS | L Gymnosperms nuclei in megaprothallus secondary xylem and phloem tracheids with differentially thickened walls zygote producing a multicellular — with early development onium or embryo-sac A simplified summary of character distributions in embryophytes and the subclasses that they character given in upper case. Such grou P may be regarded monophyletic. Widely = he irt only definable E i presence and absenc n low Unless these gro n be shown to hav e regarded as eae: For further details p nia ou s uius the bryophyte aa see Mishler = p < [72 e un jen defining characters, l Although such diagrams can thus be largely independent of evolutionary preconceptions (Nelson, 1979; Patterson, 1982b; Platnick, 1979, 1982), most systematists wish to interpret them in phylogenetic terms (Crane & Hill, in press; Hill & Crane, 1982; Wiley, 1981). Indeed, for many systematists, the compatibility between cladistic reasoning and current concepts of char- acter change during phylogeny is an important reason for preferring a cladistic rather than a phenetic approach. Interpreted in terms of phy- logeny the nested synapomorphies can be viewed as a nested sequence of homologous evolutionary novelties, and the problem of determining char- which characters are relatively de- a Or nc Groups defined by synapo- morphies (clades) can be interpreted as including a common ancestor and all its descendants (monophyletic). Groups such as the gymno- sperms and pteridophytes (Fig. 1) that contain 1985] GNETUM, WELWITSCHIA Sterling (1963) Tube nucleus Microspore nucleus < Sterile nucleus Generative nucleus CRANE-SEED PLANT PHYLOGENETICS Male gamete Spermatogenous nucleus Martens (1971) Prothallial nucleus Microspore nucleus <Ç Antheridial EPHEDRA Martens (1971) Prothallial nucleus Microspore nucleus < Prothallial Prothallial nucleus nucleus Male gamete Tube nucleus mitia Male gamete Generative nucleus Male gamete Tube nucleus Antheridial initiale Sterile nucleus Generative nucleus Male gamete Spermatogenous nucleus Male gamete E2. Comparison of male gametophyte development in Gnetum, Welwitschia, and Ephedra. See text Fic for E explanation. some, but not all, of the descendants of a com- mon ancestor (paraphyletic), may be interpreted as definable only by a combination of shared primitive and advanced features. The vertical axis of the cladogram can be interpreted as rel- ative time and the nodes as hypothetical ances- tors. Relationships may be viewed in terms o relative recency of common ancestry (Bigelow, 1956; Hennig, 1966). Ginkgo shares a more re- cent common ancestor with Pinus than either does with Lycopodium. Finally, the principle of parsimony can be nes as a criterion for se- lecting between competing cladograms in which homoplasy PAN a mn: ñawi ` ce and re- versal) is minimized. = ANALYSIS OF CHARACTERS 1.1 Male gametophyte composed of four nu- clei. In Gnetum and Welwitschia three haploid mitoses produce a male gametophyte of four nu- clei (Martens, 1971; Sterling, 1963), though the pollen may be dispersed at the bi-nucleate or tri- nucleate stage. According to Martens (1971: 146, 260), the gametophyte consists of a prothallial nucleus, a tube nucleus, and a generative nucleus, which produces two male gametes directly with- out division into sterile and spermatogenous nu- clei. Sterling (1963: 195) interpreted the pro- thallial nucleus of Martens as a “tube nucleus" and the tube nucleus as a "sterile nucleus" (Fig. 2; see Sterling, 1963, for a more detailed consid- eration of terminology). In Ephedra, the micro- spore undergoes five haploid mitoses to produce a six-celled microgametophyte consisting of two prothallial nuclei, a tube nucleus, a sterile nu- cleus (stalk nucleus), and a spermatogenous nu- cleus (body nucleus) (Martens, 1971: 48, 260). The pollen is dispersed with five free nuclei. The development of the male gametophyte in Ephedra is similar to that in Araucariaceae, Pi- naceae, and Podocarpaceae (Sterling, 1963). In Ginkgo there is also a six-celled microgameto- phyte, but the details of development (particu- larly of the motile gametes) differ from those of Ephedra and conifers (Lee, 1955). Within the conifers, Cephalotaxaceae, Taxaceae, Taxodi- aceae, and most Cupressaceae apparently have only three haploid mitoses and lack prothallial cells (Sterling, 1963), as in Gnetum and Wel- wits: vid In most cycads, there are four mitotic divisi and only one prothallial cell is pro- duced pecora 1963). By out-group comparison with pte ipse las I interpret: the more extensive ve- and six-c generalized Bis and regard the fous: cellsd condition as relatively specialized. Because the conifers are accepted as a monophyletic group in this paper (p. 727), reduction of the male ga- metophyte must have occurred at least twice in seed plants, once within the conifers, and once within the Gnetales. The character can never- 722 theless be regarded as a potential synapomorphy in the Gnetales that unites Gnetum and Wel- witschia. 1.2 Female gametophyte tetrasporic. In Gnetum and Welwitschia, the female gameto- phyte is tetrasporic, being the meiotic product of a single megaspore mother-cell without the formation of cell walls (Martens, 1971: 261). In Ephedra, which is monosporic, the megaspore mother-cell produces a linear tetrad of mega- spores, the lowermost of which enlarges and di- vides to form the female gametophyte. Occa- sionally wall formation between the megaspores may be slightly delayed (Lehmann-Baerts, 1967). The monosporic condition occurs in conifers, Ginkgo, cycads, and most angiosperms, and the tetrasporic condition therefore is interpreted on out-group comparison as a derived character. 1.3 Archegonia absent. In Gnetumand Wel- witschia, no archegonia differentiate in the fe- male gametophyte (Martens, 1971; Sporne, 1971a). In Gnetum this reflects the free nuclear condition of the micropylar end of the gameto- phyte at fertilization, whereas the female ga- metophyte of Welwitschia is cellular at fertiliza- tion, and although archegonial initials can be detected, they do not develop (Singh, 1978). In phedra, two or three archegonia differentiate from the superficial cells of the gametophyte and consist of an egg cell, a ventral canal cell, and a distinct neck (Martens, 1971, figs. 30, 31). In conifers, Ginkgo, and cycads, the archegonia are well differentiated (Sporne, 1971a: 115, 137,169). From out-group comparison I interpret the ab- sence of archegonia in Gnetum and Welwitschia as a derived character. 1.4 Embryo with “feeder.” The mature em- bryo in Gnetum and Welwitschia has a lateral, finger-like extension of the hypocotyl, termed a "feeder" (Sporne, 1971a: 179). It remains embedded in the seed after germination and may have an absorptive function. In Gnetum the Pollen of Gnetum is spheroidal, generally 20 um or less in diameter, with small spinules. The grains are typically in- aperturate, but some have a single, thin and poorly defined pore-like area (leptoma) (Erdtman, 1957, 1965: 42-44). Pollen of Welwitschia is ellipsoi- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 dal, approximately 50 um long, and the exine has about 20 longitudinal ribs separated by shal- low grooves. One of the grooves is larger and forms a poorly defined, elongated aperture (Erdt- man, 1957, fig. 30, 1965: 80-81; Wodehouse, 1935, 1936). Pollen of Ephedra is similar to that of Welwitschia. Grains are ellipsoidal, approxi- mately 20-40 um long, with 15-18 straight or zig-zag longitudinal ribs separated by grooves that are V-shaped in section. There is no aper- tural region like that in Welwitschia (Erdtman, 1957, fig. 30, 1965: 37-41; Steeves & Barghoorn, 1959; Wodehouse, 1935, 1936). The pollen of living conifers, Ginkgo, and cycads shows no close similarities with that of any gnetalean genus; however, the ribbed pollen in Welwitschia and Ephedra is unique and interpreted here as spe- cialized within seed plants. 1.6 Vessels. Vessel elements occur in all three genera of Gnetales. The early protoxylem tra- cheids are unusual and have circular bordered pits associated with annular or helical wall thick- enings (Bierhorst, 1960: 281). These are modified to form pores only in the elements differentiated later in ontogeny (Bierhorst, 1960). In Gnetum the later protoxylem tracheids have pores to- wards their ends. These appear to develop from large pits by loss of pit membranes and frequent- ly by a reduction in the width of the pit border (Bierhorst, 1960: 280—283; Bliss, 1921). Gener- ally there are fewer than six widely spaced pores arranged in a uniseriate row (Bierhorst, 1971: 473). The metaxylem tracheids generally contain fewer pores, and they are arranged in contact with each other. In the secondary xylem, the per- foration plates are often transversely oriented and may consist of groups of pores or a single pore (Bierhorst, 1971: 473; Thompson, 1918). These simple plates represent at least two different sit- uations: true simple pores, and compound struc- tures formed by loss of the secondary wall be- tween several adjacent pores (Bierhorst, 1971: 73). Muhammad and Sattler (1982) document the extreme variability in the perforation plates of Gnetum and discuss their mode of formation. In Welwitschia the pores in the protoxylem are generally single, anda a row of two or three is rare. In y xylem only single pores occur, r, and there is no evidence that they arise by fusion of several smaller ones as in Gnetum (Bierhorst, 1960: 287). In Ephedra the late protoxylem and early metaxylem vessel members typically have more pores than those of Gnetum. They are arranged 1985] in a uniseriate row, but in the late metaxylem and secondary xylem the pores tend to be ar- ranged in compact groups. Simple perforation plates are very uncommon (Bierhorst, 1960: 281). Vessels do not occur in any other gymnosperm and are clearly relatively specialized within seed us in They are, however, present in some, pre- mably only distantly related, pteridophytes vn wg 1958, 1960, 1971). On this basis they are accepted here as a potential synapomorphy of the Gnetales (see p. 767 for discussion of ves- sels in flowering plants). 1.7 Microsporangiate and ovulate "flowers" with opposite pairs of bracteoles. The micro- sporangiate and ovulate “flowers” of the Gne- tales are arranged on a fundamentally similar plan that provides a unifying character for the group. Each “flower” is comprised of a system of opposite and decussate bracteoles axillary to a primary bract (Martens, 1971: 256). Interpre- tations of the homologies of the bracts, brac- teoles, and various envelopes that they form have varied, and an enormous literature has accu- mulated (Martens, 1971; Pearson, 1929). The explanation of homologies given here is based on a straightforward interpretation of the cur- rently available data In Gnetum the inflorescences may be “‘unisex- ual" or “bisexual,” but both are elongated and have whorls of reproductive structures. Micro- sporangiate z biras “flowers” are aggregated into one or more whorls subtended by a fleshy ring-like collar paa of Pearson, 1929). The base of the inflorescence is subtended by a pair of opposite bracts (Martens, 1971, figs. 102.1, 105.1, 107; Pearson, 1929, fig. 35). The lower- most collar frequently has two teeth oriented at 90? to the two basal bracts, and, if several collars with teeth are present, their arrangement is op- posite and decussate (Martens, 1971; Pearson, 1929: 56). Although most collars do not have teeth and show no evidence of bipartite structure in their ontogeny (Martens, 1971: 208-209), they generally are interpreted to have arisen by the p fusion of two opposite bracts (Pear- son, ied 56). In ovulate spikes there are usually two rings of fertile ovulate descr. Dia d iid above each other. In mi- frequently several whoris of “flowers,” the upper of which in some species (e.g., G. gnemon, Pearson, 1929: 57) are ovulate but “incomplete” (see below) and non- functional. “Floral” primordia develop adaxially from an annular meristem at the base of the su- «€ CRANE-SEED PLANT PHYLOGENETICS 723 praadjacent developing collar (Martens, 1971, fig. 102.4), but the axillary position of the “flowers” is clearly demonstrated by their vasculature. The arrangement of **floral" vascular bundles may be complex, but the traces are predominantly de- rived from the bundles that supply the collars (Pearson, 1929: 62-68; Thoday, 1911). The microsporangiate structures of Gnetum are surrounded at the base by a tubular **perianth" that has two median lobes at the apex (Martens, 1971: 257; but see Pearson, 1929). There is some indication of bipartite structure during early on- togeny (Martens, 1971: 213, fig. 110.5, 110.6), and the perianth is generally interpreted as hav- ing been phylogenetically derived from two pos- terior-anterior bracteoles (Fig. 3A; Bierhorst, 1971: 475; Sporne, 19712). The nucellus in Gnetum is surrounded by three envelopes (Fig. 3B) that have been variously in- terpreted (Martens, 1971; Pearson, 1929). All three layers arise ontogenetically as circular mer- istems, the outer differentiating first (Martens, 1971, fig. 104). The inner layer, interpreted here as the only integument, is extended at the apex beyond the two outer envelopes. The m outer layers give no indication of an origin from pairs of opposite bracteoles (Martens, 1971), and that interpretation (Fig. 3C) hinges solely on rec- onciling the structure of Gnetum with that of Ephedra and Welwitschia (see below). In “in- complete" flowers of Gnetum the middle layer begins 5 develop but then aborts (Chamberlain, 1935: 416). In Welwitschia the microsporangiate and ovu- late inflorescences are dichasially branched (Sporne, 1971a: 180), and the ultimate branches terminate in cones with opposite and decussate bracts (*cone-scales," Sporne, 1971a: 180). The microsporangiate “flowers” (Fig. 3D) are the most complex “floral” structures in the Gnetales. Each consists of a central sterile ovule with a single integument and elongated micropylar tube ex- panded at the apex into a papillose funnel-shaped opening (Martens, 1971: 134-135; Sporne, 1971a). At the base of the ovule there are two small lateral flaps (Martens, 1971, fig. 71.11). The ovule is surrounded by a cup-like structure bearing two lateral, two dorsal, and two ventral microsporangial stalks. Each stalk has a terminal trilocular sporangium that dehisces by three ra- dial slits at the apex. In addition to the two lateral sporangia there are also two lateral “‘pseudo- staminal" primordia (Martens, 1971: 137-142). 724 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURE 3. Comparison of microsporangiate and ovulate “floral” diagrams in the Gnetales. — A. Gnetum microsporangiate “flower,” after Martens (1971, figs. 105.4, 105.7, 110.5, 110.6). — B. Gnetum ovulate “flower,” after Martens (1971, figs. 103, 104). C. Gnetum ovulate “flower,” interpretation, see text (character 1.7) for explanation.— D. Welwitschia microsporangiate “flower,” redrawn from Martens (1971, fig. 70.6). — E. Wel- witschia ovulate “flower,” redrawn from Martens (1971, fig. 63.6). —F. Ephedra microsporangiate “flower, .. after Eames (1952, fig. 1A). —G. Ephedra ovulate “flower,” based on Martens (1971).—H. Ephedra abnorma microsporangiate “flower,” after Mehra (1950: 176, pl. V.2). N = nucellus. Outside the microsporangiate cup there is an in- er anterior-posterior pair of bracteoles oriented parallel to the primary bract and an outer pair of lateral bracteoles. The ovulate “flowers” of Welwitschia (Fig. 3E) are organized on a plan very similar to those with microsporangia. The single ovule has an elon- gated micropylar tube but it is not expanded at the apex. There are no microsporangiate struc- tures, or any indication of them, but there is an inner pair of anterior-posterior bracteoles (that the wing in “fruit”? and an outer pair of lateral bracteoles. e microsporangiate and ovulate inflores- cences of Ephedra are borne in axillary position and vary considerably in their degree of branch- 1985] ing (Pearson, 1929). The “flowers” are clustered into cones" with an opposite and decussate ar- rangement of primary bracts. The microsporan- iate **flowers" (Fig. 3F) consist of a two-lipped **perianth" that arises from two anterior-poste- rior primordia (Pearson, 1929: 76) and sur- rounds the base of the microsporangiate struc- tures. The morphology of the microsporangiate structures varies among species. In E. distachya, E. intermedia, and E. trifurca there are two or more separate sporangiophores, sometimes with a protruding, vascularized axial remnant be- tween them (Bierhorst, 1971: 467; Chamberlain, 1935: 369; Eames, 1952). Eames (1952) inter- prets this condition as primitive. In other species, such as E. antisyphilitica (Hiechorst, us 467), "1m rahimnar tube. Eames (1952) regards this condition as rel- atively advanced. The number of sporangia var- ies from two to eight, forming a trilocular, quad- rilocular, or more typically bilocular synangium at the apex of the columnar tube (Pearson, 1929: 75). The ovulate “flowers” of Ephedra (Fig. 3G) are organized similarly to those with microspo- rangia. Generally only the uppermost pair of pri- mary bracts in a “cone” is fertile (Pearson, 1929: 54-55), and the lower bracts frequently become red and fleshy at maturity: presumably they aid in dispersal. Each bract subtends one “flower.” The ovule has a single integument with an elon- gated micropylar tube that exhibits a range of morphologies in different species (Pearson, 1929: 79). The “‘perianth” has been variously inter- preted (Martens, 1971: 41), but consists of two anterior-posterior bracteoles that may be free or connate at their margins (Eames, 1952: 88). These orm the “husk” of the “fruit” at maturity. There is a direct correspondence between the structure of microsporangiate and ovulate “‘flow- ers” in Ephedra, and the “‘perianth” of both ap- pears homologous (Eames, 1952). This is sup- ported further by the similarity of poorly developed *perianths" in ovulate “flowers” to the normal *perianth" in microsporangiate “flowers” (Mehra, 1950: 177-178). There is also a close similarity between the microsporangiate “flowers” of Ephedra and Gnetum (Pearson, 1929: 75), and the microsporangia in both are supplied by two vascular strands (Mehra, 1950). The arrangement of bracts and bracteoles in these three “flowers” differs from the organization in Welwitschia only in the absence of the outer lat- eral bracteoles from the microsporangiate and CRANE- SEED PLANT PHYLOGENETICS 725 ovulate “flowers.” I suggest that the “‘perianth” in Ephedra and Gnetum is homologous with the inner bracteoles of Welwitschia, and that merely the lateral outer bracteoles are missing. This sug- gestion is supported by the fact that the micro- sporangia in Ephedra are in fact arranged in a whorl like those of Welwitschia, and that where bisporangiate “flowers” of Ephedra have been reported (Mehra, 1950: 168-169) the microspo- rangia occur where expected (Fig. 3H): surround- ing the ovule but inside the “‘perianth” (Mehra, 1950: 177). Only the two outer envelopes around the ovules in Gnetum remain unaccounted for. These show no sign of having fused, either phy- y (but see Martens, 1971: 203), - Boni pairs of bracteoles. However, their position, and the occurrence of microspo- rangia between the integument and the middle layer (inner bracteoles) in abnormal flowers of G. scandens (Thompson, 1916), suggest a direct equivalence with the inner and outer bracteoles of Welwitschia (compare Fig. 3C, E). Certainly they cannot be reconciled more easily with the reproductive structures of any other gymno- sperm. These views on the homology of gneta- lean flowers are summarized in Figure 3; their organization is unique within seed plants. On this basis they are interpreted here as a potential syn- apomorphy of the Gnetales. DISCUSSION AND INTERPRETATION The data matrix for the seven characters of the Gnetales is given in Table 1 and the resulting cladogram in Figure 4. The results suggest that the Gnetales are a monophyletic group (see also Arber & Parkin, 1908; Hill & Crane, 1982) with- in which Gnetum and Welwitschia are more closely related to each other than either is to Ephedra. The Gnetales are separated from all other gymnosperms and united as a group by their fundamentally similar floral organization (character 1.7) and the presence of vessels (char- acter 1.6). Their monophyly is further supported f y pp in all genera (but not all species, Bierhorst, 1971; Foster & Gifford, 1974; Gifford & Corson, 1971; Sporne, 1971a) and the presence of a stem apex with a discrete tunica (Foster & Gifford, 1974; Martens & Waterkeyn, 1963, Voronin etal., 1973) unlike the typical condition in gymnosperms (Gifford & Corson, 1971; Johnson, 1951; Sporne, 1971a: 110, 133, 167, 184). This agrees with the view expressed by Bierhorst (1971) and Coulter 726 EPHEDRA WELWITSCHIA GNETUM the data in Table 1. See text fo Viewed methodologically character l. 5 cou ld Eun ra eer Welwitschia (a, j or as having been secondarily n Gnetum (1.5*). Fro iens ribbed pollen is more "likely to be bomine within the Gnetales and to have been subsequently modified in Gnetum. and Chamberlain (1917: 402) that, “whatever may be the connections of Ephedra with other gymnosperms ... it cannot be separated from Welwitschia and Gnetum Additional evidence for a close phylogenetic relationship of Gnetum and Welwitschia is pro- vided by: the presence of asterosclereids (Bier- horst, 1971: 475; "spicularcells," Martens, 1971: 255), the occurrence of vestured pits in the early protoxylem (Bierhorst, 1960), the occurrence o ut not exclusively syndetocheilic) mata (Florin, 1951, fig. 7e; Maheshwari & ai 1961; Martens, 1971, fig. 961), totally cel- lular embryogenesis (Martens. 1971: 765), and the irregular cellularization that occurs during female gametophyte development resulting in multinucleate cells and the formation of fusion TABLE 1. ANNALS OF THE MISSOURI BOTANICAL GARDEN Data matrix for characters of the Gnetales. [Vor. 72 nuclei (Martens, 1971: 262). Ephedra is cladisti- cally more primitive. Eames (1952) has empha- sized how *'distantly related" Ephedra is from elwitschia and Gnetum, and Coulter and Chamberlain (1917: 403) Ul that, “ Wel- witschia and Gnetum are farther removed from other gymnosperms than is Ephedra.” Some of the similarities that Ephedra shares with conifers have been mentioned above, and in cladistic terms these are interpreted as similarity owing to the retention of plesiomorphic (primitive) characters. PHYLOGENETIC RELATIONSHIPS IN THE CYCADS Extant cycads comprise ten genera (Bowenia, Ceratozamia, Cycas, Dioon, Encephalartos, Lepidozamia, Macrozamia, Microcycas, Stan- geria, Zamia) with approximately 100 species in tropical and sub-tropical regions (Greguss, 1968; Sporne, 1971a). They are typically pachycaul, unbranched, with persistent leaf bases, and are generally treated as a natural group (Bierhorst, 1971; Chamberlain, 1935; Sporne, 19712). A de- tailed analysis of relationships within cycads is not available, but in this paper I am concerned only with characters that appear to define cycads as a monophyletic group. ANALYSIS OF CHARACTERS 2.1 Girdling leaf-traces. The leaf-traces in cycads arise on the opposite side of the stem to the leaf they supply. They then girdle the stem at a slightly oblique angle before entering the leaf (Chamberlain, 1935: 88-89; Sporne, 1971a: 109). Girdling leaf-traces are not known to occur in any other gymnosperm and are interpreted as a Other Seed Plants (exclud- Character ing angiosperms) ` Ephedra Welwitschia Gnetum 1.1 Male gametophyte of four nuclei +/— — + + 1.2 Female gametophyte tetrasporic _ _ + + 1.3 Archegonia absent — — + + 1.4 Embryo with “feeder” — — * * 1.5 Ribbed polle _ + + _ 1.6 Vessel — + + + 1.7 Microsporangiate and ovulate "flowers" with opposite pairs of bracteoles _ + + + 1985] OTHER EXTANT CYCADS SEED PLANTS r— | 2.3 2.2 2.1 seeds Ficure 5. Probable synapomorphies of extant cy- cads. See text for further explanation. € emg Sauer The traces to the cotyle- cycad pP are apparently ra- 53). All extant genera of cycads have megasporophylls aggregated into a simple cone. In all extant genera except Cycas each cone terminates the activity of the apical meristem, and further growth continues sym- podially by the activation of another meristem at the base of the cone stalk (Bierhorst, 1971: 373; Sporne, 1971a: 109). This mode of growth is reflected anatomically by domes of old cone vascular tissue (“cone domes") that extend across the pith. In Cycas only the microsporangiate plants grow in this fashion, and the ovulate plant is truly unbranched. Ovulate cones of Cycas only differ from those of other cycads in being inde- rminate, and after the production of megaspo- rophylls, further growth of the same apical mer- istem produces vegetative leaves. The simple ovulate cones of all cycads are interpreted here asa aped feature quite different from the “compound” ovulate cones of conifers, or the ““cone”’-like Basora of “flowers”? in Gne- les 2.3 Presence of cycasin. The methylazoxy- methanol glycoside cycasin is present in all ten cycad genera and is not known to occur in any other gymnosperm (DeLuca et al., 1980; Moretti et al., 1983). A similar compound macrozamin is also ubiquitous in cycads and is a potential synapomorphy of the group (Moretti et al., 1983). DISCUSSION AND INTERPRETATION Girdling leaf-traces (character 2.1), simple ovulate cones (character 2.2), and the presence of cycasin (character 2.3) are provisionally in- CRANE—SEED PLANT PHYLOGENETICS 727 TABLE 2. Data matrix for characters of cycads. Other Seed Character Plants Cycads 2.1 Girdling leaf-traces = + 2.2 Simple ovulate cone = + 2.3 Presence of cycasin = + terpreted as uniting cycads as a monophyletic group (see Fig. 5 and Table 2). PHYLOGENETIC RELATIONSHIPS IN THE CONIFERS The living conifers are generally treated as a natural group comprising six families: Araucari- aceae, Cephalotaxaceae, Cupressaceae, Pinaceae, Podocarpaceae, and Taxodiaceae, with approx- imately 50 genera and 550 species (Bierhorst, 1971; Chamberlain, 1935; Sporne, 1971a). The Cupressaceae and Taxodiaceae are treated as a single family by some authors (Eckenwalder, 1976a, 1976b). A further family, the Taxaceae with five genera, sometimes is included with oth- er conifers (Bierhorst, 1971; Chamberlain, 1935; Foster & Gifford, 1974) or sometimes separated from them (Sporne, 1971a). When included the Taxaceae is often placed with the Cephalotaxa- ceae in the Taxales (Greguss, 1972). Separation of the Taxaceae is based on Florin’s view (1951) that the ovuliferous structures of conifers and taxads are fundamentally different (see below). A preliminary cladistic analysis of conifers was presented by Miller (1982). In this paper I deal only with the relationship between the extinct Pennsylvanian conifer Lebachia, extant conifers and all other seed plants. I attempt to justify the conifers sensu lato (conifers plus taxads) as a monophyletic group. Although the group is in- tuitively natural, a single defining character can- not be specified without invoking a series of sub- sidiary hypotheses. These are discussed below. ANALYSIS OF CHARACTERS 3. E Narrowly triangular leaves. The leaf gy of fi narrowly tri- ted dles to b d, short scales angular t In Agathis and some species of Araucaria and Podocarpus, the leaves are broad, leathery, and laminar. Despite this variability, all families, in- cluding the Araucariaceae (Stockey, 1982) and 728 Podocarpaceae (Buchholz & Gray, 1948), have at least some species with narrowly triangular leaves (Bierhorst, 1971, table 25-1; Dallimore & Jackson, 1948). In many conifers ontogenetic evidence is generally interpreted as indicating that helically arranged, narrowly triangular leaves are primitive within the group (Chamberlain, 1935: 358). This is particularly clear in the Cupressa- ceae, in which the adult foliage frequently con- sists of opposite and decussate, or whorled, scale- like leaves, but is also seen in other families. The ontogenetic sequence in conifers with well-de- veloped broad, laminar leaves has never been surveyed in detail, but in Araucaria angustifolia (A. brasiliensis) the early leaves of the seedling si broadly triangular (Hill & DeFraine, 1909). n Lebachia the leaves are generally narrowly de occasionally with bifid tips (Florin, 1951; Mapes & Rothwell, 1984; Rothwell, 198 2a; Scott & Chaloner, 1983). I interpret narrowly triangular leaves as the primitive condition in all conifers and regard all other leaf morphologies as secondary modifica- tions of this basic type (but see Discussion and Interpretation). If this view is accepted then leaf phology distinguishes living me Les bachia from other seed plants, which character- istically have large megaphyllous leaves. The closest similarity to the leaves of conifers is found in Ephedra, but these are typically arranged in pairs or whorls of three that are joined by a sheathing base around the stem (Foster & Gif- ford, 1974: 523-524). 3.2 Resin canals. Resin canals occur in all conifers, although their distribution in different organs of the plant varies. In some Pinaceae they occur in the cortex of roots and stems, the wood, the leaves, and the ovulate cones. In the Arau- cariaceae, Cephalotaxaceae, Cupressaceae, Po- docarpaceae, and Taxaceae there are typically no resin canals in the secondary xylem (Greguss, 1972) although they usually occur in other parts of the plant, and resin cells are generally present in the xylem parenchyma. Resin canals are par- ticularly sparse in the Taxaceae, especially in Taxus (Chamberlain, 1935), but they do occur in the leaves of Torreya and the roots of Austro- taxus and Taxus cuspidata (Chamberlain, 1935: 252). Resin canals occur in Middle Pennsylva- nian Lebachia-like conifers (Rothwell, 1982a), but lack the epithelial lining seen in many extant conifers. It is not clear whether the “mucilage canals" of cycads, certain seed ferns, and Ginkgo (Bier- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 horst, 1971: 375; Sporne, 1971a: 168; Stewart, 1983: 253) are comparable to those of conifers. According to Bierhorst (1971: 375) the contents of these canals in cycads are partly miscible with water and do not appear to have a high terpene content. In this paper I interpret the possession of resin canals as a potential synapomorphy of conifers but closer comparisons, including de- velopmental studies, are needed with secretory structures in other gymnosperms. I interpret the relatively sparse resin canals of Taxus as due to secondary loss. There is some limited ontoge- netic evidence to support this conclusion. Jeffrey (1903) has reported that in Sequoia gigantea res- in canals are present only in the secondary xylem of the first year. Resin canals also can be induced by injury in wood that otherwise lacks them (Jef- frey, 1 3.3 Fertile axillary shoot lacking an apical meristem. It is now widely accepted that the ovulate cones of most conifers are compound structures, the ovuliferous scale being a modified shoot bearing megasporophylls with ovules. Each ovuliferous scale is borne in the axil of a bract. This hypothesis (the polyaxial or *Brachyblast" theory, Florin, 1954) had been proposed by a succession of plant morphologists since the first half of the nineteenth century (see Coulter & Chamberlain, 1917: 245), but was advocated most strongly as a unifying concept by Florin (1951) supported by his observations on Recent and fossil conifers. On this interpretation the most primitive cones would be like those of Cryp- tomeria in which there is little fusion between the ovuliferous scale and the bract and in which the ovuliferous scale has leaf-like lobes. Cones such as those of the Cupressaceae, that show con- siderable fusion between the bract and ovulifer- ous scales, are considered relatively advanced. Ontogenetic evidence supports these ideas. In some Taxodiaceae the bract and ovuliferous scales are free when young, but not during sub- sequent growth (Sporne, 1971a: 141). According to the theory, each ovulate cone is a reduced "inflorescence," and hence conifers with very poorly developed cones, or with a single bract by reduction (Florin, 1951: 363-367; Wilde, 1944, fig. 40) According to Florin (1951), only the Taxaceae cannot be accounted for in terms of the polyaxial theory, since the single terminal ovule shows no evidence of having evolved by reduction from a 1985] EXTANT CONIFERS LEBACHIA | Wiss l 3.3 3.2 3.1 FiGUREÓ. Preliminary cladogram summarizing re- latonships between Lebachia and other conifers based n the data in Table 3. See text for further explanation. bract and ovuliferous shoot system, or from a reduced “inflorescence” (Florin, 1951: 372). Flo- rin (1948, 1951, 1955) regarded this difference from other conifers as fundamental and indica- tive of a totally independent origin for the two groups. Harris (1976) argued for a closer rela- tionship between conifers and taxads based on a plausible scenario for deriving a terminal taxad ovule from a single fertile axillary shoot. Evi- dence from stelar morphology is consistent with this view, and “the stelar system of Taxaceae... differs in no significant way from the stelar sys- tem of conifers with helical phyllotaxy” (Beck et al., 1982: 753). I treat the Taxaceae as another family within the conifers based on their leaf morpholgy and the presence of resin canals. Like Harris (1976), I regard the terminal ovule as a secondary sim- plification from forms with ovuliferous scales (shoots) and bracts. Similar but less extensive simplification is interpreted as having occurred in the Podocarpaceae and Cephalotaxaceae (Florin, 1951; Wilde, 1944). This view is also implicit in those classifications of conifers that regard the Cephalotaxaceae, Podocarpaceae, and Taxaceae as closely related (Buchholz, 1934; Keng, 1975). Modified fertile shoots in the axils of bracts or leaves also occur in cordaites, Ginkgo, and the Gnetales. In cordaites and Lebachia the apical spiral (Florin, 1951). nifers the shoot apex is not differentiated (Harris, 1976: 124), and there is no regular phyllotactic spiral. I interpret this as an advance over the condition in Lebachia and cordaites, and there- fore as a feature that unites all extant conifers. CRANE-SEED PLANT PHYLOGENETICS 729 TABLE 3. Data matrix for characters of conifers. Other Extant e - coni- Character Plants bachia fers 3.1 Narrowly triangular leaves - + +/— 3.2 Resin canals +/— + + 3.3 Fertile wawapa shoot la s. pical meristem +/— = + DISCUSSION AND INTERPRETATION The «+h harartara £ My . | A here are listed in Table 3, and the resulting cladogram in Figure 6. The presence of narrowly triangular leaves and resin canals unites Lebachia and ex- tant conifers as a monophyletic group. Rothwell (1982a: 18) has also pointed out that the single plane of symmetry of Lebachia seeds is a feature that separates them from those of other platy- spermic Paleozoic gymnosperms. This could perhaps be used as a defining character of the group after further study. Within the extant co- nifer plus Lebachia group, the lack of the ovu- liferous shoot apex provides a character that de- fines extant conifers. These conclusions may not b understood fossil, conifers. e (1981: 216) raises the intriguing alternative pos- sibility of conifer polyphyly. Based on a consid- eration of Wilde’s work (1944) on Podocarpa- ceae, he suggests that whereas most conifers may be most closely related to Lebachia, the Podo- carpaceae may in fact be more closely related to cordaites. PHYLOGENETIC RELATIONSHIPS N THE GLOSSOPTERIDS The glossopterids are a loosely defined group centered around the characteristic leaves, Glos- sopteris sensu lato, that are a dominant element in Permian and Triassic “Gondwana floras.” Pu- levoryas, 1969, but see Schopf, 1976: the Permian of Turkey (Archangelsky & Wagner, 1983). Glossopterid leaves are simple, narrow- ly elliptical, with a midrib and well-developed reticulate venation (Fig. 7E). The group was di- verse, and a variety of different reproductive 730 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 D> / AMA: Nd RD S NS s mm ! = ? j Z ER | PRX = Ta’ AT JE IVAN — M / > y ` — ANNI WA 2 |! d) S IGURE 7. Morphology of glossopterids. —A. peyi Ii e Ae and associated leaf, redrawn from Pant (1977a, fig. 10E-G), orient n Pant and Nautiyal (1984); x 1.— B. Lidgettonia africana megasporophyll, based on Thomas (1958), debis d Chandra (1975, text-fig. 12), Schopf (1976, fig. 8D); x1.5.—C. Eretmonia microsporophyll, redrawn from Surange and Chandra M text-fig. ID); x2.— D. “Glossopteris” (?Dictyopteridium) megasporophyll in axil of a vegetative leaf, redrawn fr Delevoryas (1977, fig. 1d), note that details of sporophyll attachment and sporophyll eae are uncertain, see text for discussion; xl. —E. Glossopteris sastrit leaves borne ona shoot, based on Pant and Singh (1974, text-fig. 2B-D); x0 , based on Pant and Nautiyal (1960, text-fig. 3A); x25. ar Pollen grain from micropyle of P.) raniganjense, redrawn from Pant and Nautiyal (1960, text-fig. 4G); x 550. 1985] structures have been described (Pant, 1977a; Schopf, 1976; Surange & Chandra, 1975; White, 1978). Unfortunately, many are known only as impressions in which cellular details have not been observed, and their interpretation is there- fore equivocal. In this initial analysis I consider only three of the more completely understood glossopterid reproductive structures, “‘G/lossop- teris" (?Dictyopteridium, Gould & Delevoryas, 1977), Lidgettonia (Thomas, 1958), and Otto- karia (Pant, 1977a; Pant & Nautiyal, 1965). The ovule bearing structures of most glossop- terids were borne adnate to the surface of a leaf. Although they have often been reconstructed adjacent to the adaxial leaf surface (Fig. 7D), this has never been convincingly demonstrated. Glossopterid reproductive structures could be interpreted morphologically in several different ways (e.g., Retallack & Dilcher, 1981, fig. 3), but there are two main possibilities. The ovule bear- ing structure is either a leaf homologue (mega- sporophyll) or it is a modified axis (i.e., a cladode). Despite its axillary position, Gould and Dele- voryas (1977) emphasized its leaf-like nature, and in this paper I treat it as a megasporophyll. This interpretation also brings the ovulate re- productive structures of glossopterids into line with those of other Mesozoic seed ferns that ap- pear to have megasporophylls. The anatomically preserved megasporophylls described by Gould and Delevoryas (1977) from the Late Permian of Queensland, Australia were simple, narrowly elliptical with an inrolled lam- ina, bearing ovules on its inner surface (Fig. 7D). Although Gould and Delevoryas provided some evidence to suggest that the ovule bearing surface faced the subtending leaf, the interpretation re- mains equivocal and at variance with the recon- struction of Ottokaria by Pant and Nautiyal (1984, see below). The gross form of the Queens- land material is similar to Dictyopteridium de- yas, 1977: 397; Surange & Chandra, 1975). The interior of the structure was filled by a delicate network of filaments, or plates of cells, between the ovules. Based on clusters of S gli pollen sacs in the Queensland chert, Gould and Dele- voryas (1977) suggested that the microsporan- giate organs of their plant were of the Eretmonia or Glossotheca type (Fig. 7C). From other local- ities Eretmonia and Glossotheca are known to consist of a fertile portion (interpreted here as a microsporophyll) adnate to a leaf. The micro- sporophyll consisted of a bifurcated axis, each CRANE-SEED PLANT PHYLOGENETICS 731 branch of which bore numerous Arberiella microsporangia. Pollen was bisaccate (Protohap- loxypinus), with numerous distinct striations on the body of the grain oriented at 90° to the distal suture (Fig. 7 Lidgettonia africana (Fig. 7B) was described by Thomas (1958) from the Upper Permian of South Africa. It consists of a sterile leaf with an adnate megasporophyll. The megasporophyll dif- fers from that in Gould and Delevoryas' ma- terial in bearing about eight fertile “cupules” on slender stalks adnate to the lamina of the sub- tending leaf. Each **cupule" consisted of an ex- panded disc, which is thought to have borne ovules on its lower surface. Thomas (1958) in- terpreted these discs as peltate, but Schopf (1976: 45-48) has shown that the p was attached laterally, as in the Indian specie cronata (Surange & Chandra, 1975). yrs aris sacs and Pa dames. of the Eretmonia type are associated with the Lidgettonia fructifications (Schopf, 1976: 50; Thomas, 1958), but no in- formation is available on the pollen. In this study I assume that it was of the Protohaploxypinus type. Ottokaria (Pant, 1977a; Pant & Nautiyal, 1965, 1984) from the Permian of India is preserved as compressions from which some cellular details are known. The megasporophyll was adnate to the upper surface of a leaf and expanded distally to form a *spoon-shaped" head with sterile mar- ginal lobes, bearing ovules on the concave sur- face (Fig. 7A). Contrary to Gould and Delevor- yas' interpretation of the Queensland material, the ovule bearing surface faced away from the subtending leaf (Pant & Nautiyal, 1984). The ovules are similar to the dispersed ovule genera pinus pollen (*'Striatites, " the micropyles of these dispersed ovules. Apart from the inference that they contained Proto- haploxypinus pollen, the microsporophylls or microsporangia of Ottokaria are unknown. In this paper they are assumed to have been similar to Eretmonia. ANALYSIS OF CHARACTERS 4.1 Glossopterid leaves. The leaves of all three genera are narrowly elliptical, with retic- ulate venation. The leaves are distinctive, not known to occur in any other group of gymno- sperms, and are interpreted as a specialized char- 732 "GLOSSOPTERIS" (?DICTYOPTERIDIUM) LIDGETTONIA OTTOKARIA D. IM ZIMA RE 6. y cladog izing lationships between Lidgettonia, ‘“‘Glossopteris” (?Dic- tyopteridium), and Ottokaria based on the data in Ta- e 4. See text for further explanation. acter. The closest similarity in venation occurs in leaflets of Caytonia (Sagenopteris), but this general pattern of reticulate venation is also widespread in ferns and other groups (Alvin & Chaloner, 1970). 4.2 Megasporophylls adnate to a subtending leaf. The megasporophylls in all three genera are adnate or closely adjacent to the subtending leaf. This arrangement is unknown in any other group of seed plants and is interpreted as a spe- cialized character. 4.3 Microsporophylls adnate to a subtending leaf. Glossopterid microsporophylls from sev- eral localities (Pant, 1977a) are borne adnate to the subtending leaf (Gould & Delevoryas, 1977). This basic arrangement is apparently general in glossopterid plants and is assumed (although not known) to be present in the three glossopterids considered here. It is not known to occur in any other group of seed plants and is interpreted as a potential glossopterid synapomorphy ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 4.4 Striate bisaccate pollen. Bisaccate pol- n with a transverse distal sulcus, and longitu- dinal striations on the body of the grain, is thought to be general in glossopterids. It occurs rarely in other groups of gymnosperms and some conifers (see also Nipurdia, Pant & Basu, 1979). It is in- tepreted as a specialized character. Simple megasporophyll. In **Glossopter- s” QDictyopteridium) and d Ottokaria, the ovules are borne on the surface of a simple laminar structure, d here as a simple mega- sporophyll. Megasporophylls of Cycas are sim- ilar, but are not adnate to a subtending leaf, and bear ovules only on the margin rather than over the lamina surface. In Lidgettonia the presence of several ovule bearing structures suggests that the adnate megasporophyll was pinnate rather than simple as in “Glossopteris” (?Dictyopterid- ium) and Ottokaria. Pinnate megasporophylls occur in the following seed ferns: Callistophyton, peltasperms, corystosperms, and Caytonia; and are therefore interpreted here as primitive within the glossopterids. The simple lamina is inter- preted, conversely, as a specialized feature that unites **G/ossopteris" (?Dictyopteridium) and Ot- okaria ~ DISCUSSION AND INTERPRETATION The data matrix for the five characters of glos- sopterids is given in Table 4 and the resulting cladogram in Figure 8. Glossopterids are defined on their simple axillary megasporophylls. These conclusions, and the assumptions made, need to be tested by further work. As pointed out by Pant (1977a), there is no evidence to support the sug- gestion of Surange and Chandra (1975) that the TABLE 4. Data matrix for characters of glossopterids. Glossop- teris/ Other Seed Dictyop- Character Plants Lidgettonia teridium Ottokaria 4.1 Glossopterid leaves + + + 4.2 ssi n ate adnate subtending leaf T + + 4.3 Mierosporophylis adnate subtending leaf + + + 4.4 s. bisaccate pollen +/— + + + 4.5 Simple megasporophyll +/— — + + 1985] glossopterids sensu lato should be divided into *pteridospermous" and “‘glossopterid”’ forms. PHYLOGENETIC RELATIONSHIPS THE BENNETTITALES The Bennettitales are diverse and abundant plants in Upper Triassic to Upper Cretaceous floras. The leaves may be simple (Nilssoniopte- ris) or, more typically, pinnate, with parallel, oc- casionally dichotomous veins in the pinnae. In the te. Theleave are superficially like those of cycads but can usu- ally be distinguished by characters of gross form and always by their characteristically cutinized syndetocheilic (paracytic) stomata (Fig. 11C) that contrast with the haplocheilic (anomocytic) sto- mata of true cycads (Harris, 1932a; Thomas Bancroft, 1913). Bennettitalean microsporo- phylls or ovules were aggregated into flower-like i wa usually surrouodad by a “perianth” of acts. The “flowers” were unisexual (monospo- be or bisexual (bisporangiate). In all Ben- nettitales the ovules were orthotropous, with two integuments joined to the nucellus only at the chalaza (see interpretation of Vardekloeftia and Bennetticarpus crossospermus below; also char- acters 9.19, 9.22). Distally, the inner integument generally formed an elongated micropylar tube. The ovules were crowded together into heads and individually were not subtended by any othe organ. In most compressions of bennettitalean ovules cuticles are obtained from the outer, and bai acp the inner, surface of the inner integu- nt, the micropylar tube, and the surface of the n rne but there is no maceration resistant megaspore membrane (Harris, 1932b, 1954, 1969). The ovules were separated by sterile in- terseminal scales, which had a single vascular undle. morphological nature of these in- terseminal scales is uncertain, but the structure of B. crossospermus described by Harris (1932b) provides evidence to support Seward's view (1913: 118, 1917: 403) that they are PENAN with ovules (see also Delevoryas, 1968: , in- terpreted here as bitegmic, p. 764). In 5 cros- sospermus (Fig. 9E, F) the elongated micropylar tube passed through the center of a thickly cu- tinized **micropylar plate" very similar to the flattened cutinized apices of the interseminal scales in the same gynoecium (Fig. 9E). This sug- gests that the ovules and interseminal scales may be homologous and that the scales “are formed by the diverted development of seed initials" o = CRANE-SEED PLANT PHYLOGENETICS 733 (Harris, 1932b: 116). Studies of ovule develop- ment in Williamsonia (Sharma, 1974) also sup- port this idea. There is some evidence that B. crossospermus is associated with Prerophyllum known to assess the relationships of the plant in more detail. In this paper relationships between 11 of the better understood bennettitalean taxa are considered (Table 5). ardekloeftia is a genus erected by Harris ‘oa for two species of spherical, bennetti- talean, ovulate heads from the Rhaetic of Green- land. Each head was composed of flattened, bi- laterally symmetrical ovules separated by numerous interseminal scales. The ovules were relatively few (two to 20), and relatively large (5— mm long), compared to those of most Ben- nettitales. They also were unusual in having an outer cutinized layer (the **cupule," Harris, 1932b, interpreted here as the outer integument) sur- rounding the inner integument in an identical position to the **micropylar plate" of B. crosso- spermus. Two very similar species have been de- scribed, V. conica, and the more completely known V. sulcata. In V. sulcata (Fig. 9B, C), each head contained two to six seeds. In young ovules the cutinized layer outside the inner integument formed a flattened plate through the center of which the micropyle projected. At this stage the outer layer (outer integument) was very similar to the micropylar plate of B. crossospermus but was less substantial at maturity. Vardekloeftia sulcata is associated with Pterophyllum kochi leaves (Fig. 9A; Harris, 1932b: 111), but the mi- crosporangiate structures are unknown. With the exception of Vardekloeftia all other bennettitalean taxa considered in this paper have more ovules in their megasporangiate “flowers” and a well-differentiated **perianth." The basic structure is illustrated by two very similar species, Williamsonia harrisiana (Bose, 1968) and W. sewardiana (Sahni, 1932b), both based on silic- ified material from the U 10A). Williamsonia sewardiana is thought to have been attached to Bucklandia indica stems that bore Prilophyllum cf. cutchense leaves (Sahni, 1932b: 10). Both species were apparently uni- € (monosporangiate). Although microspo- angiate “flowers” have been described from the scq Hills (Bose, 1967; Sharma, 1969; Sith- 734 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 [A 299^ » uh Sve S Sola ECT AS RSQ * ` 1 NI Ñ NN N NN N ( N ~ N M NI No ` a uto uet f : À Z í Cs INE | t Ex = 2 (` L^ Ay = — d } / f | \ Y k y] Y \ NY X N Sp een WT - 2 i Hii E. e — $ F i š E : FiGuRE 9. Morphology of the Vardekloeftia and Bennetticarpus plants.—A. Pterophyllum kochii, based on Harris (1932b, fig. 29); x0.5.— B. V. sulcata, spherical head composed of ovules and interseminal scales, based on Harris (1932b, pl. 15, fig. 1, pl. 17, figs. 1, 2, pl. 18); x2.—C. Vardekloeftia, longitudinal section through ovule, redrawn from Harris (1932b, fig. SOE); x 5.—D. Pterophyllum ptilum, based on Harris (1932b, fig. 30D); X0.5.—E. B. crossospermus, outer surface of interseminal scales showing position of ovule, based on Harri (1926, pl. 11; 1932b, fig. 48J); x 10.—F. B. crossospermus, longitudinal section through ovule, redrawn from Harris (1932b, fig. 50C); x10. CRANE- SEED PLANT PHYLOGENETICS 735 1985] (je (snoave, pL6I VAI) səltupz¿ (enxosiq) ‘ds vapioappro;) -nxasiq) ‘ds rvapioappo«;) ‘ds papi02ppo(? -310 "] B1OFeq ymos (renxestq) 6961 'sured — 4O[Dtu S1421d0140SS]INI (renxesiq) 1421421] p]jə1uosuup1]]1⁄44 1421081] DjJ21408WDI]]UM ó (jenxosiq) (Lvl :6961 6961 LH ppa Stuajdotuossj1IN — ([enxosIq) pJDuO402 D]JaUOsSWDIIA, || DIDu0402 p]jəluostun1]]1⁄4 *sujeH 99s) 1opuo[s ps 6961 'suuJeH uəpəd tun])KXudoln1q uaj2ad n11/2141]2 M 1(quə2322] Diuoswvi]]ug -omisnd pnipup])yo5ng jo sapiou 696] 'su1eH -119Əd wnjjadopiQ SISUAIQUYM D11/2141]Ə M anpinu ptuosutun1]]144 psojnjsnd vipuppjong (S31ssem f 6961 ‘SULH Sp818 sə1lttupz JOS »n(2uilaM SD318 piluostun1]]1⁄44 s0813 pDipuppyong ‘W) ayso “WN (dIssel 40u piJof -nf "T-omsseu[ `Q) Qct61I (SuJeHn -1U S21upzowuouy Ssapioiisapua wunisuniajdospapyi, — vijofusnauv vjjaipuvjary, -1]SN8UD vjJaipur]as44( pueru FW uəpəAs 6S6I (snoooei *seÁ1oA9[9(] é é poyfiudvw DISayJUDUOP p>ox/0uSpu DISIYIUDUOW, -310 (]) OOIxƏJA] MON 8961 “soq ¿ ¿ DUDISIAIDI| DluOStuD1]]1⁄4 ¿ Əsuət/2]no (orsseiní) qc€61 ‘Tues 3o uun]]((t(doln11q ¿ DUDIPADMAS pluosuub1]]1⁄4 popu: vipuvjyong SWH jeyewfey *erpug (orsseun( '—31S qc£6l1 ‘SeH 11490) wnjjdo4a1d ¿ DIDIINS Di]f20pJ2p40A ¿ -SEUL `()) puepuaa1n) 4 95u919JƏ%[ JE97T IMO 21erdug1odsoJorJq J9^O0[J NEMAO UI91S ÁujevoT Kreurnq `səre1nəuuəg UMOUY 19119q ƏY) JO sues1o parejos] `ç ATV 736 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Limp D 1 S A š ti LALA asss (o AA a ; "A S WSS NIS QOO 2) Jam 77 S DN NOV UP JHOY O e RC uM E FIGURE 10. Morphology of Bennettitales. — A. Williamsonia harrisiana, longitudinal half section of “flower,” based on Bose (1968, pl. 1, figs. 8, 9); x 1.5.— B. Bennettites albianus, cutaway section through an ovule, based Williamsonia gigas, longitudinal half section of young "flower," 1985] oley & Bose, 1953, 1971), the microsporangiate structures of both species, = = vegetative parts of W. harrisiana, are unkno Monanthesia described “tas the Upper Cre- taceous Mesaverde Formation of New Mexico (Delevoryas, 1959) is similar to Williamsonia sewardiana and W. harrisiana. The “flowers” were monosporangiate and borne on long, slen- der peduncles in the axil of almost every leaf. The receptacle was conical and covered with ovules and interseminal scales. The microspo- rangiate “flowers” and pollen are unknown. Four species of Williamsonia have been de- scribed based on compressions from the Middle Jurassic of Yorkshire, W. gigas, W. hildae, W. leckenbyi, and W. himas (Table 5; Harris, 1969). The first three are the best understood as “whole” plants, and W. himas is not considered further in this review. Williamsonia gigas (Fig. 10C) has been linked to the leaves Zamites gigas and the microsporangiate “flowers” Weltrichia sol (Fig. 10D, E; Harris, 1969: 163). Williamsonia hildae has been linked with the leaves Ptilophyllum pec tinoides, the microsporangiate "flowers" Wel- trichia whitbiensis, the **perianth" bracts Cyca- dolepis hypene, and the stem Bucklandia pustulosa (Harris, 1969: 172). Williamsonia leckenbyi has been linked to the leaves Ptilo- phyllum pecten and the microsporangiate “‘flow- ers" Weltrichia pecten (Harris, 1969: 170) (Table 5). The structure of the ovulate “flowers” in all three species is basically similar to that in Wil- liamsonia harrisiana and Williamsonia seward- iana. The Yorkshire species differ however in the presence of a distinct sterile “corona” above the level at which ovules are borne. The corona is formed from the receptacle and interseminal scales (Fig. 10C; C. R. Hill, pers. comm.; Harris, 1969: 129). The microsporangiate “flowers” of the three plants also are basically similar. Wel- trichia sol consisted of a whorl of up to 30 “rays” (interpreted here as microsporophylls) more or less fused proximally into a cup-like structure. The inner surface of the cup bore large numbers of “resinous sacs" and the free parts of the mi- crosporophylls bore two-valved synangia (Fig. monosulcate (Fig. 10F). In Weltrichia ue (Fig. 10D) there were about 30 microsporophylls, and the synangia were borne alternately on short fer- tile appendages that projected from the inner sur- face of the microsporophyll. Comparison with the microsporophylls of Cycadeoidea suggests that CRANE-SEED PLANT PHYLOGENETICS 737 these appendages are fertile pinnae. Each valve of the synangium contained about 12-15 micro- sporangia. In Weltrichia pecten and Weltrichia whitbiensis there were between ten and 20 mi- crosporophylls, each ond Hina synangia in two rows on their inner su Wielandiella ple rond from the Rhaetic of Scania, southern Sweden (Nathorst, 1909), and Greenland (Harris, 1932b) had monosporangiate “flowers” (Harris, 1932b: 90-91). The ovulate “flowers” are similar to those of Williamsonia gigas, Williamsonia leckenbyi, and Williamso- nia hildae and have a distal sterile corona. The microsporangiate flowers are poorly understood and require restudy (Lundblad, 1950: 73), but the “palisade rings" described at the base of the “flower” by Nathorst (1909) are similar to Hy- Md anii marsilioides (Lundblad, 1950). The leaves of Wielandiella angustifolia are thought to te Anomozamites minor. Two similar species of Williamsoniella are known as compressions from the Middle Jurassic of Yorkshire (Harris, 1944, 1969, 1974). Both were bisporangiate (bisexual) “flowers” (Fig. 11A, D). A whorl of bracts formed a “‘perianth” around a whorl of about 12 laterally compressed, wedge- shaped EHAA Each i muerosporophpil bore two on the awl suffice (interpreted as fertile bin- nae), each of which bore a single two-valved syn- angium containing pollen sacs with psilate monosulcate pollen. At the center of the “flower” the receptacle bore ovules and interseminal scales and was expanded distally into a terminal sterile corona. The two species, sos lignieri and Williamsoniella coronata (Fig. are linked with Nilssoniopteris major and a vittate (Fig. 11B, C), respectively (Harris, 1969: 146, 149). Williamsoniella lignieri was larger than W. coronata and generally is less well understood. “Perianth” bracts are associated with the “‘flow- er" but have not been seen attache Many species of Cycadeoidea have been de- scribed from a wide range of Upper Jurassic and Lower Cretaceous localities in the Northern Hemisphere (Crepet, 1974; Stewart, 1983; Wie- land, 1906, 1916), but all show a basically similar structure. Cycadeoidea was pachycaul, with short, stout trunks and helically arranged leaf bases that probably bore Zamites-like leaves (Seward, 1917; Wieland, 1906: 87). The “flowers” were bispo- rangiate (bisexual) (Fig. 1 1E) and borne on short pedicels in leaf axils. They did not project beyond the persistent leaf bases. There was a perianth of short ft 738 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 Ti NS" Wy = jj ZZ FicuRE 11. Morphology of Bennettitales.—A. Williamsoniella coronata, longitudinal half section through “flower,” based on Harris (1944, fig. 1); x 3.— B. Nilssoniopteris vittata, redrawn from Harris (1969, fig. 32A), an unusual specimen showing two lateral pinnae at the base; x1.— C. N. vittata, detail of stoma showing paracytic guard cells and sinuous anticlinal flanges, based on Harris (1969, fig. 32D); x1,000.— D. W. coronata, surface of immature “gynoecium” showing micropyles and interseminal scales, based on Harris (1969, fig. 62F); x 50. — E. Cycadeoidea sp., longitudinal half section through “flower,” based on Wieland (1906, fig. 88) and Crepet (1974, pl. 61, fig. 21); x1.5. 1985] helically arranged outer bracts surrounding ap- proximately 20 microsporophylls. These were fused proximally into a shallow cup but were pinnate distally and compressed laterally. Two rows of appendages (pinnae) were borne adaxi- ally, each with a row of about six kidney-shaped synangia. The synangia contained eight to 20 tu- bular sporangia and produced psilate, monosul- cate, dps iig pollen (Crepet & Zavada, pers. comm. wards their apices the micro- sporophylls were Pi with their abaxial surface against a conical or domed receptacle covered with ovules and interseminal scales, and showing no sign of a corona. Delevoryas (1963, 1968) and Crepet (1972, 1974) both support the view that the microsporophylls never opened to become fully expanded as Wieland (1906) sup- posed. ANALYSIS OF CHARACTERS 6.1 Interseminal scales. Interseminal scales occur in all Bennettitales considered in this anal- ysis. They are not known to occur in any other seed plant, and following Harris (1932b) I inter- pret them as developing from ovule primordia (see characters 9.19, 9.22). They are regarded here as a specialized character. 6.2 Guard-cells, paracytic and heavily cutin- ized. The stomata of all Bennettitales are para- cytic and interpreted as ontogenetically synde- tocheilic (Harris, 1932a; Thomas & Bancroft, 1913). This is sufficient to distinguish them from the stomata of most other gymnosperms except Gnetum, Welwitschia, and some conifers (Scott & Chaloner, Mes However, ui guard- cells of the B ti cutinized, outer periclinal walls and dorsal an- ticlinal flanges. This feature occurs in leaves with both sunken and superficial stomata but is not thought to occur in other gymnosperms (Harris, 1932a: 94-108) 6.3 Ovulate heads with numerous ovules. heavily All though only two to six seeds develop there are a large number of interseminal scales that are interpreted as sterile ovule primordia. No other gymnosperms (except the Vojnovsky- aceae and Ruflorinaceae, Meyen, 1984) have such a large number of ovules clustered together, and I interpret this as a potential synapomorphy of the Bennettitales. CRANE-SEED PLANT PHYLOGENETICS 739 6.4 Bivalved synangia. Most bennettitalean microsporophylls have p e with- in bivalved synan ough the microspo- o iana, W. harrisiana, an a unknown, the bivalved condition is interpreted here as general in the Bennettitales. In many seed plants (e.g., conifers, cycads) the microsporangia are not in synangia; where they do occur (e.g., Caytonanthus, medullosans, and Gnetales), they are not arranged to form a bivalved capsule. Here I interpret the bivalved synangia of Bennettitales as a specialized condition, although it should b noted that they do not occur in all bennettitalean pollen organs (e.g, Leguminanthus siliquosus Kráusel & Schaarschmidt, 1966). 6.5 “Perianth” of helically arranged bracts. All of the Bennettitales considered in this study, with the exception of Vardekloeftia (Harris, 1932b), have ovulate heads that are sur- rounded by a “perianth” of numerous helically arranged bracts. This flower-like arrangement does not occur in any other group of gymno- sperms and is thus interpreted as a relatively specialized Sisi within the Bennetütales. 6.6 “Coron *corona" occurs in Wil- liamsonia gigas, W. hildae, W. leckenbyi, Wie- landiella angustifolia, and the two species of Wil- liamsoniella, W. coronata and W. lignieri. No comparable feature occurs in other gymno- sperms, and I regard this as a specialization over the condition in other Bennettitales. Bisporangiate “flowers the two species of Williamsoniella. They are not known in other gymnosperms except the Gne- tales (see character 1.7). They are interpreted here asa specialized feature within the Bennettitales, dtl "m "flowers" of the pg is considered later in this paper (p. 7 6.8 n od laterally flattened. In those Bennettitales for which the microsporo- phylls are known, only in Cycadeoidea, William- soniella coronata, and W. lignieri are they lat- erally flattened. Dorsiventrally flattened leaf-like microsporophylls are generalized in seed plants, and the laterally compressed forms found in some Bennettitales are therefore interpreted as spe- cialized. 6.9 Microsporophylls with three to four pairs of bivalved synangia. Both species of William- soniella have very similar microsporophylls, with three to four pairs of bivalved synangia. Other 740 VARDEKLOEFTIA SULCATA WILLIAMSONIA SEWARDIANA WILLIAMSONIA HARRISIANA MONATHESIA MAGNIFICA WIELANDIELLA ANGUSTIFOLIA WILLIAMSONIA GIGAS WILLIAMSONIA HILDAE WILLIAMSONIA LECKENBYI WILLIAMSONIELLA CORONATA WILLIAMSONIELLA LIGNIERI CYCADEOIDEA SP. FIGURE | 12. _Preliminary cladogram summarizing relationshi the data in Table 6. “An asterisk indicates reversal of character 6.6. See text for further explanatio gymnosperms generally have larger numbers of sporangia on each microsporophyll, and most other known bennettitalean microsporophylls have numerous synangia. The condition with few synangia is SEE as a synapomorphy of Williamsoniella species. Ni rE leaves. Both species of Walliamseniella have been linked with Nilsson- iopteris leaves, and although recognized as two distinct pied N. vittata and N. major are very similar in morphology, venation, and cuticle (Harris, 1969: 2. Megaphyllous leaves in most seed plants, including most Bennettitales, are pinnate. In the context of the Bennettitales as a whole I therefore interpret Ni/ssoniopteris leaves as a synapomorphy of Williamsoniella species. 6.11 Synangia borne directly on the "rays" (microsporophylls). The microsporangiate "flowers" of Williamsonia hildae (Weltrichia in having the synangia borne directly on the sur- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 face of the “rays” Movil qa ic i rather than on fertile lateral p Weltrichia whit- biensis, W. pecten, poris and Cycad- eoidea. The presence of these distinct fertile lat- eral pinnae is interpreted here as a primitive feature within the Bennettitales based on com- parison with the depend divided microsporo- phylls of most “seed fern DISCUSSION AND INTERPRETATION The data matrix for the Bennettitales is given in Table 6 and the resulting cladogram in Figure 12. The Bennettitales have long been widely ac- cepted as a natural group and at this level of analysis may be defined as monophyletic by the presence of interseminal scales, the characteristic cutinization of the guard-cells, the numerous ovules, and the bivalved synangia. The prelim- inary analysis presented here is the first attempt to clarify the interrelationships of the various taxa within the group. Although it provides some resolution, it is handicapped by lack of knowledge of microsporangiate organs in Var- dekloeftia, Williamsonia sewardiana, W. har- risiana, Wielandiella angustifolia, a nå ona thesia. In addition there is considerable diveret among Upper Triassic bennettitalean reproduc- tive structures that needs to be incorporated into the analysis (Kräusel, 1948, 1949; Kräusel & Schaarschmidt, 1966). Several of these fossils (e.g., Haitingeria krasseri, Leguminathus sili- are currently being reinvestigated (Crane, work in progress). It has been traditional to recognize two loosely defined families, the Cycadeoidaceae and Wil- liamsoniaceae, within the Bennettitales (Stewart, 1983; Taylor, 1981a), but these do not form monophyletic groups in the cladogram. The Cy- lly regarded as unified by having the flowers borne within an armor of per- sistent leaf bases and would include Cycadeoidea and Monanthesia considered in this analysis. The habit of these plants is similar but other char- acters do not place them together, and the flowers e (Sahni, nless other characters can be found to unite the Cycadeoida- ceae, the two traditional families of Bennettitales have dubious phylogenetic utility. It should, however, be noted that the pattern of characters CRANE-SEED PLANT PHYLOGENETICS 741 1985] 4SÁBI,, 94} uo — E = * * = é= ¿— ¿— ¿— ¿— = Apoastp uzoq visueu&g Iro = $ d as = = = i- ¿= z — =a SOABI $1421d01uOSS]1N. OI`9 - + + - = - i- i- i- ¿— i- = erdueuás Poalealq JO sired p—¢ yum sjAydosodso niw 6'9 + + + = = = ¿— ¿— ó ¿— ¿— = pousney ÁA[[e1o1v] sj[Aydor1odso1ip 8'9 + + + _ = = E = E se = = «$190],, (jenxasiq) Ə)erdgueiodsiq /'`9 _ + + + + + + _ _ — = = «BU0IOD,, 9'9 + + + + + + + + + + _ _ sjoe1q posuee Áe Jo ,,qiueusd,, ¢°9 + + + + + + + ¿+ ¿+ ¿+ ¿+ = ergueuds poA[pAlg t`9 + $ + + + + + + + + + — S9[nAO SsnoJoumnu YUM speou IMAO €'9 ¿+ + + + + + + ¿+ ¿+ ¿+ + _ pezrunno ÁqrAeog pue onÁoeBIed s[[2 preny 79 + + + + + + + + + + + = so[pos [pPururssIo]u] ['9 ‘ds papi 1420481] DjDuO402 4q apnppi — sp3i8 — DOfijsns 721 Dui puni pivojns (suuəds IIOLIVYJ -02ppoK?) vjjauos pjjətuos -uə2Ņ42ə] DIUOS puos -up vj -fugo -Suivy — -papwas loo] -ordue -upi "Mura puos wur] -uom -punim pisay DIuOS Duos -api0A — Surpn[oxo) "uiua -upuojW -uvm -luD1]]144 Sjue|d pees 19g10 'So[peiniouuog Jo SIIVLILYI 10] XLW QUA `9 14V L FIGUR (1981, fig. 6.31) and Phillips et al. (1972, pl. 37, fig. 8); c rilet on Phillips et al. (1972, pl. 45, fig. 49); x 500.—C. s s ea es sp. trilete MD based on Phillips et al. (1972, pl. 43, fig. 28); x200. accepted here suggests that the absence of the “corona” (character 6.6) from Cycadeoidea may be interpreted as secondary loss. REPRESENTATIVE EXTANT AND FossıL GYMNOSPERMS Having attempted to provide an outline of re- lationships within some groups of gymnosperms, the relationships between major seed plant taxa can be assessed. The principal constraint in such an analysis is the information available for many fossil seed plants. In this analysis I have at- tempted to keep assumptions concerning un- known characters to a minimum by using only the better understood fossil seed plants. This has excluded many potentially important fossil plants that need to be incorporated in future, more in- clusive analyses. In this section I briefly review the characters of extant and fossil seed plants, emphasizing in particular the basis and limita- tions of our knowledge of fossil taxa. ARCHAEOPTERIS Archaeopteris (Fig. 13A) ranges from the mid- Devonian (Givetian) to Lower Mississippian (Tournaisian). It is an important Late Devonian ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 RE 13. Morphology of Archaeopteris. — A. A. g fertile lateral Bis ind based on Beck —B. microspore, based plant and is the most completely understood of all progymnosperms (Beck, 1981; see Fig. 1). The plants were arborescent and eustelic with a large parenchymatous pith. There was a bifacial cam- bium that produced both secondary xylem and secondary phloem (Beck et al., 1982). The sec- ondary xylem (Callixylon) was pycnoxylic and similar to that of extant conifers. The tracheids had uniseriate to multiseriate, circular, bordered pits on the radial walls (Beck, 1: 202), and there was no axial parenchyma. The rays were uniseriate to multiseriate. Callixylon wood is own to have been attached to Archaeopteris branch systems (Beck, 1960, 1970). Although stem morphology and the pseudomonopodial branching of Archaeopteris is well understood, regular axillary branching does not occur (Beck, 81: 212; Scheckler, 1978). Leaves were heli- cally or decussately arranged and decurrent at the base (Fig. 13A), typically wedge-shaped, and varied from entire to deeply divided. The sto- mata of Archaeopteris have not been described. sporous (Phillips et al. 1972), but whether other 1985] species were homosporous or heterosporous is unknown. The microsporangia were long and slender, while megasporangia were shorter and broader. Both dehisced by a single longitudinal suture. TI about 40 um in diameter, with a proximal trilete suture and a finely granular or smooth surface (Fig. 13B). The gr i Pet- titt, 1966). Ther g (Bihari about 200 um in diameter, with a trilete RI and a reticulate to rugulate surface (Fig. 13C; Phillips et al, 1972). Both microspores and megaspores presumably germinated proximally to produce free-living male gametophytes with motile spermatozoids and female gametophytes with well-developed archegonia. However, noth- ing is known of gametophyte development, ga- metes, or embryogenesis. Y J Wu MEDULLOSANS The medullosans are among the most exten- sively studied of all seed ferns (Stewart, 1983; Stidd, 1981a, 1981b). Typically, they have two or more vascular segments in the stem but this apparent *polystely" appears to be only a mod- ification of the fundamentally eustelic arrange- ment in all other seed plants (Basinger et al., 1974; Beck et al., 1982). However, Quaestora amplecta (Mapes & Rothwell, 1980) from the Upper Mississippian of Arkansas, the earliest medullosan stem known, has an exarch proto- stele. If Quaestora is correctly interpreted as a medullosan it suggests that *polystely" may have arisen within the medullosan clade. The bifacial cambium produces secondary xylem and sec- ondary phloem. The wood has multiseriate rays and is comprised of tracheids with multiseriate, oval, bordered pits on their radial walls. Despite extensive studies of medullosan anatomy (Stew- art, 1983) and ontogeny (Delevoryas, 1955), ax- illary branching in medullosans has been dem- onstrated only once (Hamer, 1984 and pers. comm.). Medullosan leaves (e.g., Alethopteris, Neuropteris, Odontopteris) were large and pin- nately organized, often with a bifurcated primary rachis. The petioles (Myeloxylon) contained a large number of scattered vascular bundles. Sto- mata have been described as paracytic (Oestry- Stidd & Stidd, 1976), but reinterpretation and better preserved material shows that they are an- omocytic (Mickle & Rothwell, 1982; Reihman & Schabilion, 1978; Stidd, 1981a: 67). Although a few specimens are known that show CRANE-SEED PLANT PHYLOGENETICS 743 seeds apparently attached to medullosan foliage (Stewart, 1983: 260-261), most medullosan seeds, particularly those that are well understood ana- tomically (e.g., Pachytesta illinoensis), are linked with the vegetative parts on the basis of associ- ation evidence (Stewart, 1983: 260). The ovules (Pachytesta, Trigonocarpus, etc., see Hoskins & Cross, 1946; Taylor, 1965) are large (up to 11 cm long) and radiospermic. Each has a single integument with a sarcotesta, sclerotesta, and en- otesta. The integument and nucellus are free except at the chalaza. Frequently the sclerotesta has three primary ribs, and often secondary an tertiary ribs also are developed. The ovules have a radially arranged double vascular system. The outer system the sarcotesta between the ribs, while the inner system within the nucellus is formed by bundles or a network originating from a pad of vascular eem at the chalazal end. The megaspore mem- brane in medullosan seeds is thick (5-25 um) (Taylor, 1965). Some medullosan pollen organs are large, complex synangiate structures with tubular spo- rangia that dehisce longitudinally (e.g., Dolero- theca), while others such as Aulacotheca and Halletheca are simpler, with fewer sporangia cupida 1981b). Only a few of these pollen or- gans are known to have been attached to medul- losan foliage (Dennis & Eggert, 1978), but con- vincing association and structural evidence has been presented for others (Ramanujam et al., 1974; Stewart, 1983). The majority of the pollen organs are known to have produced Monoletes (Schopfipollenites) pollen. These grains are large (100—500 um long) and ellipsoidal. Distally there are often two longitudinal grooves and proxi- mally a monolete suture with a marked deflec- tion in the center. Occasionally (Parasporites, Dennis & Eggert, 1978) the grains have a pair of small sacci. The pollen wall is alveolar, with nu- merous fine cavities (Taylor, 1978). Pollen wall ultrastructure is similar to that of cycads, but the ontogeny is apparently different (Taylor, 1982; Taylor & Rothwell, 1982), and the wall cavities are much less regular. Germination is thought to have been proximal (prepollen sensu Chaloner, 1970a); male gametes were probably motile. Monoletes pollen has been described in the pol- len chamber of Pachytesta hexangulata (Stewart, 1951). Little is known of male gametophyte de- velopment, but the pollen contents figured by Stewart (1951) are suggestive of the spermato- zoids of extant cycads and hence zooidogamy. trun wit thin 744 The details of female gametophyte develop- ment are unknown in medullosans, but the ga- metophyte is cellular, with archegonia at fertil- ization (Stewart, 1951). Nothing is known of embryogenesis, and an embryo has never been described from a medullosan seed (Stewart, 1983: 282-283). CYCADS Possible defining characters of cycads have al- ready been discussed (Fig. 5). The primary vas- cular architecture of extant cycads is not under- stood clearly (Beck et al., 1982), but evidence from microsporangiate cones suggests that the lature is fund tally telic (Beck et al., 1982: 755). Typically the stem has a large pith, and the bifacial cambium (or cambia in Enceph- alartos and Macrozamia, Sporne, 1971a: 108) produces relatively small amounts of secondary xylem and phloem. The secondary xylem con- sists of tracheids with multiseriate bordered pits and has many rays. In Zamia and Stangeria the wood has tracheids with scalariform pitting (Sporne, 1971a: 108). The growth of most cycad stems is sympodial (see character 2.2), and some genera (e.g., Stangeria and Bowenia) are irregu- larly branched. According to Bierhorst (1971: 373), Stopes (1910), and Stevenson (1980) branching is adventitious, and Foster and Gif- ford (1974: 419) observed that some lateral buds form from the living tissues of persistent leaf bases while others arise due to injury. There are no detailed developmental studies of branching in cycads. Cycad leaves are generally large and pinnate with dichotomous venation, although some anastomoses do occur (Foster & Gifford, 1974: 422-423). The stomata are anomocytic. The ovules are borne on simple megasporo- phylls that are generally arranged in cones (see character 2.2). There are generally two ovules per megasporophyll, although some species of Cycas may have up to eight. The ovules are large (up to 6 cm long, Chamberlain, 1935: 104) and ra- dially symmetrical. Each has a single massive integument with three layers: a fleshy sarcotesta, a hard sclerotesta, and an endotesta. The integ- ument and nucellus are “fused” except close to the micropyle. The ovule is supplied by a double, radially arranged, vascular system. There are about ten vascular bundles that run up the out- side of the sclerotesta, and an inner anastomos- ing system of vascular bundles that run in the endotesta to the level at which the integument ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 and nucellus are free. The megaspore membrane is well developed and may be up to 10 um thick in mature ovules (Chamberlain, 1935: 126; Erdt- man, 1965), but the nucellus and inner lining of the integument is only weakly cutinized (Harris, 1954) The microsporophylls of all cycads are borne in cones and bear microsporangia on their abax- ial surface. In Cycas there may be more than 1,000 microsporangia, but in Zamia there are 25 or less (Chamberlain, 1935: 115). The micro- sporangia are arranged in sorus-like clusters of two to six, each supplied by a single vein (Ste- venson, 1982). Each microsporangium dehisces by a single slit towards the center of the **sorus." The pollen is boat-shaped, non-saccate, and more or less smooth-walled. The pollen wall is alveo- late with numerous fine cavities (Audran & Ma- sure, 1977). The male gametophyte is generally five-celled (see character 1.1), and the final di- vision of the spermatogenous cell produces two large spermatozoids up to 400 um in length (Nor- stog, 1977) with a spiral band of flagella (Sporne, 1971a: 116). The pollen germinates distally, pro- ducing a haustorial pollen tube, and the sper- matozoids are released by a rupture in the pollen tube close to the distal pollen wall. The female gametophyte develops from a sin- gle functional megaspore (see character 1.2) that undergoes free nuclear divisions until over 1,000 nuclei have been produced, before centripetal cellularization (Sporne, 1971a: 114) and the for- mation of archegonia. Following fertilization, free nuclear division produces 64-1,000 nuclei be- fore cellularization occurs (Sporne, 1971a: 116). LYGINOPTERIS Lyginopteris oldhamia is known from the Up- per Carboniferous (Westphalian A) of Europe (Oliver & Scott, 1904; Stewart, 1983). The name is applied to small stems, up to 4 cm in diameter (Taylor & Millay, 1981: 38), of possibly vine- like habit (Stewart, 1983; Taylor & Millay, 1981). The primary vasculature was eustelic (Beck et al., 1982; Scott, 1923), and a bifacial cambium produced relatively little secondary xylem and phloem. The wood contained large tracheids with multiseriate, oval bordered pits on the radial walls. The rays were wide and multiseriate. Reg- ular axillary branching occurred. Leaves were widely separated and borne helically on the stem. They were small, pinnately organized, and highly dissected. The primary rachis bifurcated shortly 1985] CRANE-SEED PLANT PHYLOGENETICS 745 Morphology of € and Telangium. — A. L. lomaxii, cupule and ovule, based ee pe Fic 14. and eon (1904, text-fig. 2); x after leaving the stem. On the basis of association and the occurrence of large, distinctive, stalked, capitate glands, Sphenopteris hoeninghausii (Ra- chiopteris aspera when anatomically preserved) is thought to be the foliage of L. oldhamia. The ovules of Lyginopteris (Lagenostoma lo- maxii, Fig. 14A) are known attached to Sphenop- teris hoeninghausii foliage. They were borne ter- minally on the distal part of We res in pedicellate cupules. Each cupule containe sin- gle ovule. The upper half of the cupule bore pee to ten distal lobes, each supplied by a single vas- cular strand. The ovules were small (about 5.5 mm long) and radiospermic. The single integu- ment was "fused" with the nucellus except near The integument was supplied by (Oliver & Scott, 1904) and may be up to 5 um in some lyginopterid pteridosperms (Taylor & Millay, 1981). It is unknown whether the mega- spore of L. lomaxi had a trilete mark, but this feature does occur in other lyginopterid mega- spores (Pettitt, 1969; Schabilion & Brotzman, 1979), and in Stamnostoma huttonense there is a cluster of three aborted spores at the apex of the megaspore (Long, 1975). This suggests that lyginopterid megaspores were derived from a tet- rahedral rather than linear tetrad. The microsporangiate organs of Lyginopteris are frequently considered to be Crossotheca (Kidston, 1906; Sporne, 1971a: 55), but there is no evidence linking these two organs as part of the same plant (Jongmans, 1930). Lyginopteris oldhamia and Lagenostoma lomaxii come from — B. T. scottii, pollen organ, based on Benson (1904, figs. 1, 9, 10); x strata older (Westphalian A) than those in which Crossotheca is typically abundant. Benson (1904) made a good case for Telangium scottii (Fig. 14B) being the Lyginopteris microsporangiate organ, b ally symmetrical synangia with six to eight ellip- soidal sporangia that dehisced along their inner wall. The pollen (prepollen) was small wit proximal trilete mark. The pollen wall tee ie ture is unknown, but in other supposed lyginop- terids (e.g., Crossotheca, Millay et al., 1978) it is structurally homogeneous. Male gametophyte development is unknown, but presumably ger- mination was proximal and produced motile sperm. The large flagellate spermatozoids de- scribed in the micropyle of Lagenostoma ovoides (Benson, 1908) would be worth reexamining. The female gametophyte of Lagenostoma lo- maxii was cellular, and the development prob- ably involved an initial free nuclear phase fol- lowed by centripetal cellularization (McLean, 1912; Oliver & Scott, 1904). The female game- tophytes of L. ovoides and Hydrasperma tenuis were also cellular, and archegonia have been de- scribed (Long, 1944; Matten et al., 1984). Long (1975) has described a possible lyginopterid em- bryo from the Lower Carboniferous of Scotland, but embryos of Lagenostoma and all lyginop- terid pteridosperms are otherwise unknown CORDAIXYLON Several species of microsporangiate and ovu- late Cordaianthus inflorescences have been de- 746 scribed in the literature (Rothwell, 1977; Dagh- lian & Taylor, 1979), but only one is known as part of a "whole plant" (Rothwell & Warner, 1982, 1984). The description given here is based on the Cordaianthus inflorescences attached to Cordaixylon dumusum stems and leaves from the Late Pennsylvanian of eastern Ohio (Roth- well & Warner, 1982, 1984) The mature stems were up to about 5 cm in diameter and “haapa abundant ien roots. The plan primary a system was an aah cic and a bifacial cambium produced sec- ondary xylem and secondary phloem. The sec- ondary xylem was composed of tracheids with crowded multiseriate oval bordered pits on the dec walls. The rays were predominantly uni- riate. Leaves were helically arranged and ranged vin needle-like with a single vein to spatulate forms up to 3 cm wide with numerous veins. Some branches bore needle-like leaves at the base and spatulate leaves distally. Others bore needle-like leaves throughout. Anatomical de- tails of the leaves are described by Rothwell and Warner (1984). The stomata consisted of two guard cells with two lateral and two subsidiary cells. Inflorescences were borne irregularly as epi- cormic branches on the stems. Frequently the xylem of the cone axis was not continuous with the xylem of the stem. Ovulate inflorescences (Cordaianthus duquesnensis Rothwell, 1982b) consisted of a bilaterally symmetrical primary axis bearing four rows of fertile secondary shoots, each in the axil of a bract. The secondary shoots had 60-70 helically arranged scales of which the distal 20-30 were fertile and bore a single ter- minal ovule. Immature ovules are known at- tached to the inflorescence and are linked by a continuous morphological series with mature ovules of Cardiocarpus oviformis Leisman (1961) that occur in the same deposit. These mature ovules were up to about 9 mm long and distinctly flattened. There was a single integument that was free from the nucellus except at the chalaza. Each ovule was supplied by a single vascular bundle that expanded to form a flattened plate at the base of the nucellus. Below this plate the main vascular bundle produced two lateral traces that ran through the sarcotesta in the primary plane, one on either side ofthe nucellus. The megaspore membrane was about 5 um thick. The structure of the microsporangiate inflo- rescence was similar to that of the ovulate inflo- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 rescences, consisting of a bilaterally symmetrical primary axis with four rows of fertile secondary shoots, each in the axil of a bract. The secondary shoots bore helically arranged scales the distal- most of which were fertile. rus fertile scale bore ged in a ring at the apex. The microsporangia were fused proximally and free distally. Pollen was saccate and of the F/or- inites type. Florinites pollen has a distal thinning of the wall and an alveolate wall structure with large cavities. Nothing is known of cordaite male gameto- phytes from North American material, but Flor- in (1936) described male gametophytes from Florinites in situ within Cordaianthus from Eu- rope (Millay & Eggert, 1974). Florin's material shows axial rows of four or five cells in each grain, and the male gametophyte was therefore basi- cally similar to that seen in many conifers and Ephedra (Millay & Eggert, 1974; Fig. 2, see char- acter 1.1). Germination was probably distal (Mil- lay & Taylor, 1974). The female gametophyte of Cardiocarpus oviformis was cellular and pos- sessed two archegonia as described in C. spinatus by Andrews and Felix (1952). Nothing is known of the embryo of the '*Cordaixylon plant." MESOXYLON The cordaite inflorescence Gothania was de- scribed originally from European material (Scott, 1919), but knowledge of the Gothania plant has been increased considerably by description of G. lesliana from the Middle Pennsylvanian of Ken- tucky (Daghlian & Taylor, 1979) and Mesoxylon priapi from the Upper Pennsylvanian of eastern Ohio (Trivett, 1983; Trivett & Rothwell, 1985). On the basis of association in both European and North American coal balls, Daghlian and Taylor (1979) suggest that G. /esliana is the microspo- rangiate fructification of a M. sutcliffi stem with Cordaites felicis foliage and Mitrospermum compressum ovules. Trivett and Rothwell (1985) describe Mesoxylon priapi based on stems with attached Cordaites felicis foliage and microspo- rangiate inflorescences of the Gothania type. This material forms the basis for the description given here Mesoxylon stems (Beck et al., 1982; Scott & Maslen, 1910) had a primary vascular system consisting of a modified mesarch eustele. A bi- facial cambium produced secondary xylem and secondary phloem. The secondary xylem was composed of tracheids with crowded multise- 1985] riate oval bordered pits on the radial walls. The rays were two to four cells wide. Axillary branch- ing is known to occur and axillary buds were covered with needle-like bud scales. The leaves (Cordaites felicis) were helically arranged, prob- ably spatulate, and several wide with numerous veins. Anatomical details of the leaves are described by Benson (1912), Good and Tay- lor (1970), and Trivett and Rothwell (1985). The stomata consisted of two guard cells with two lateral and two subsidiary cells. Inflorescences were borne in the axils of leaves. Occasionally both an inflorescence and a vege- tative bud occur in the axil of a single leaf. Ovu- late inflorescences are unknown, but were pre- sumably similar to those of Cordaixylon dumusum, consisting ofa bilaterally symmetrical primary axis bearing four rows of fertile second- ary shoots, each in the axil of a bract. Mature ovules of Mitrospermum vinculum (Grove & Rothwell, 1980) occur in the same deposit as M. priapi and contain abundant pollen from the Mesoxylon plant in the micropyle. Mature ovules were up to about 4 mm long and distinctly flat- tened. There was a single integument that was free from the nucellus except at the chalaza. The ovule was supplied by a single vascular bundle that expanded to form a flattened plate at the base of the nucellus. Below this the main vascular bundle produced two lateral traces that passed through the sclerotesta and extended to the mi- cropyle in the sarcotesta. The bundles were po- sitioned one on either side of the nucellus in the primary plane of the ovule. The megaspore membrane was 6-13 um thic The structure of mmictospoennpiats inflores- cence was similar to that of the ovulate inflores- cences, consisting of a bilaterally symmetrical primary axis with four rows of fertile secondary shoots. The secondary shoots had up to 28 he- lically arranged scales. There were about five fer- tile scales confined to the distal part of the shoot and each bore four microsporangia arranged in a row at the apex. The microsporangia were free throughout their length. Pollen was saccate and of the Sullisaccites type. Sullisaccites has a distal monolete or trilete suture and an alveolate wall structure with large cavities (Millay & Taylor, 1974). Gothania lesliana produced monosaccate Felixipollenites pollen with a monolete or trilete proximal suture. The pollen wall was alveolate, with irregular, medium-sized cavities (Taylor & Daghlian, 1980). Nothing is known of male ga- metophyte development, but both Felixipolle- CRANE-SEED PLANT PHYLOGENETICS 747 nites and Sullisaccites (prepollen) are thought to have germinated proximally. Nothing is known of the female gametophyte or embryo of Mesoxylon. Male and female ga- metophytes of other cordaites are known and discussed under Cordaianthus. CONIFERS Possible defining characters of conifers have been discussed already (Fig. 6). The primary vas- culature of the stem is eustelic (Beck et al., 1982) with a single bifacial cambium producing sec- ondary xylem and phloem. The secondary xylem is pycnoxylic, characteristically with uniseriate rays, and the tracheids have uniseriate to mul- tiseriate circular bordered pits, generally con- fined to the radial walls. Some genera have long- and short-shoot systems, and axillary branching occurs. Although there is considerable variation in leaf morphology (see character 3.1), no conifer has pinnate leaves. Stomata are anomocytic (but see Scott & Chaloner, 1983). Ovulate cones are diverse in form, but all are regarded as fundamentally compound, with cone scales interpreted as fertile shoots in the axils of bracts (see character 3.3). The ovules vary con- siderably in size and are flattened. Each has a single Paeria which frequently has two “horns” on either side of the micropyle in the primary pis (Bierhorst, 1971, figs. 25.3B, 25.3C, 25.5E). These “horns” reflect the onto- genetic development from two opposite primor- dia that also are positioned in the primary plane (Bierhorst, 1971, fig. 25.5C, 25.5D; Florin, 1951: 364). The integument and nucellus may be “fused” for most of their length or free except at the chalaza. The ovule lacks a well-developed vascular system, and frequently there are no bun- dles at all (Chamberlain, 1935: 298). The mega- Podocarpaceae it may be 4-6 um thick, whereas in the Araucariaceae, Cupressaceae, Cephalotax- aceae, Taxaceae, and Taxodiaceae it is generally less well developed and may be extremely thin (Erdtman, 1965). It is also thick in Lebachia (Mapes & Rothwell, 1984, pl. 15, fig. 2). Out- group comparison with cycads, Ginkgo, and pteridophytes suggests that the thick megaspore membrane is the primitive condition in the co- nifers, thinner membranes being more special- ized. The nucellus and inner lining of the integ- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 FIGURE 15. well (1975, text-fig. 1); x0 e pes ofthe Callistophyton plant. —A. Callistophyton, branch system, redrawn mom Roth: allospermarion pusillu iles, based on Rothwell (1981, re 3), note that details of seed attachment i in Callistophyton are not well cid hw see text; x 5. —C. re sil ss pinnule (1981, fig. 3A); x 10.— . Vesicaspora pol on Rothwell (1981, pl. 3, fig. 7); x 11 1 L1 e MAJ Wany 1954). The microsporangiate cones of conifers are ex- gia. There are several kinds of pollen (Erdtman, 1965; Wodehouse, 1935), but saccate grains with a distal aperture are interpreted here as primitive for extant conifers based on out-group compar- ison with Lebachia (see Fig. 6). Saccate pollen is accepted here as primitive for the group. The pollen wall structure in most saccate grains is alveolate with large cavities, but in the non-sac- cate conifers (e.g., Araucariaceae, Cupressaceae) the pollen wall is granular (Doyle et al., 1975) Male gametophyte development varies (Sporne, 197 1a: 142), but the developmental pat- tern that occurs in cycads, Ginkgo, and Ephedra bearing three adaxial microsporangiate sy . boyssetii detail of proximal pinnules, base len d from an Idanothekion glandulosum pollen sac showing microgametophyte, based 0. angia, based on Rothwell d on Rothwell (1981, fig. 2B); x 2.5.— is accepted here as primitive for the group (see character 1.1). The pollen germinates distally and fertilization is typically siphonogamous, al- though there are rare reports of zooidogamy in conifers (Christiansen, 1969). The female gametophyte develops from a sin- gle functional megaspore (see character 1.2) that 3 do [cH g N e Em ° 5 e = a o a < short free nuclear phase in the development of the embryo GINKGO The genus Ginkgo contains a single extant species, G. biloba, although there are several fos- sil species, mostly known from foliage only (Tra- lau, 1968). The primary vasculature of the stem 1985] is eustelic (Beck et al., 1982) with a single bifacial cambium producing secondary xylem and phloem. In long shoots the secondary xylem is pycnoxylic and conifer-like with narrow rays, and uniseriate or biseriate circular bordered pits on the radial walls of the tracheids. Short shoots have manoxylic wood. Axillary branching oc- curs. The distinctive fan-shaped leaves have two vascular bundles in the petiole. The venation is dichotomous with a few anastomoses (Foster & Gifford, 1974: 449). Stomata are anomocytic. The ovules are borne in pairs at the end of stalks in leaf axils. The stalks are slightly dorsi- ventrally flattened and contain two pairs of vas- cular bundles. Each ovule bearing stalk is inter- preted as a short shoot bearing two uniovulate fused megasporophylls each represented by a pair of vascular bundles (Rothwell, pers. comm.). The ovules are large (up to 4 cm long) and either flattened or trigonous. Each has a single, massive integument that frequently has two apical “horns,” one on either side of the micropyle in the primary plane (Bierhorst, 1971: 421, fig. 24- 4B). There is a sarcotesta, a sclerotesta, and en- dotesta. The integument and nucellus are “fused” except close to the micropyle. The ovule is sup- plied by two (or three) poorly developed vascular strands that run inside the sclerotesta to the level at which the integument and nucellus are free. The vascular bundles run in the primary plane, and the sclerotesta has two (or three) distinct ribs. Occasional ovules may have three or four vas- cular bundles and a corresponding number of ribs. The megaspore membrane is well devel- oped and up to 4—6 um thick at maturity (Cham- berlain, 1935: 208; Erdtman, 1965: 10, but see Karkenia, Archangelsky, 1965); the nucellus and inner lining of the integument are only weakly M Soap (Harris, 1954). The m PAPE DEE organs also arise in the axils of pais and consist of a lax inflorescence bearing numerous lateral stalks interpreted here as microsporophylls. Each microsporophyll bears a pair of pendant WD dies and an apical mucilage-containing “knob” (Sporne, 1971a: 169). At the base of each Es stalk there is a pair of minute projections. The sporangia de- hisce by a single longitudinal slit. The pollen is boat-shaped, non-saccate, and more or less smooth-walled. The pollen wall is alveolate, with large cavities similar to those in many conifers a & Crepet, pers. comm.). Male gameto- phyte development is similar to that in Ephedra and some conifers (character 1.1). As in cycads, CRANE-SEED PLANT PHYLOGENETICS 749 the final division of the spermatogenous cell pro- duces two large spermatozoids with a spiral band of flagellae (Sporne, 1971a: 170). The pollen ger- minates distally, produces a haustorial pollen tube, and the motile spermatozoids are released by a rupture in the pollen tube close to the distal pollen wall. The female gametophyte develops from a sin- gle functional megaspore (see character k éh which 256 pulei an are formed, before cellularization oc- curs centripetally and archegonia are formed (Sporne, 1971a: 169). Following fertilization, free nuclear divisions produce 256 nuclei before cel- lularization of the embryo occurs (Chamberlain, 1935: 211; Sporne, 1971a: 171). CALL Callistophyton (Fig. 15) is currently the best understood of all seed ferns based on Middle and Upper Pennsylvanian material from North America. Its morphology and reproductive bi- ology are reviewed by Rothwell (1975, 1980, 1981). Stems of Callistophyton are up to 3 cm in diameter, and the plant is thought to have been small and shrubby with a scrambling habit (Fig. 15A). The primary vasculature of the stem is eustelic with a pith and a ring of five sympodia, as is typical of Lyginopteris and many fossil co- nifers. The secondary xylem is well developed, typically with biseriate rays but with some up to five cells thick. Tracheids have alternate multi- seriate oval bordered pits on the radial walls. The secondary phloem produced by the bifacial cam- bium consists of radial files of sieve cells and phloem parenchyma with scattered rays. The stem and all other parts of the plant contained spherical secretory cavities, each lined with an epithelium. Adventitious roots were formed at many of the nodes, and axillary branching oc- curred. The axillary buds were covered by small linear scales. The leaves (Fig. 15D) were very variable in size, and bi-, tri-, or quadripinnate. They were helically arranged on the stem. The stomata have not been described. he ovules (Callospermarion) were borne scattered on the abaxial surface of the leaves (Fig. 15B). Each was flattened with two planes of sym- metry. There was a single integument that was free from the nucellus except at the chalaza. It was vascularized by two vascular bundles in the primary plane of the sarcotesta. At the base of 750 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 FIGURE 16. Morphology of peltasperms. — A. Lepidopteris stormbergensis, bipinnate frond, based on Thomas (1933, fig. 53); x0.25.— B. A Harris (1932a, fig. 28E); x 4.—F. P. rotula, transverse section of ovule, redrawn utunia thomasii, megasporophyll based on Townrow (1960, t x 3.—C. Antevsia zeilleri, based on Harris (19322, pl. 7 from Harris (1932a, fig. 28E); x 4.—G. P. rotula, peltate organ bearing a circle of ovules on the lower surface, most ovules removed for clarity, based on Harris (1932a, fig. 281); x 1.5.—H. P. rotula, peltate organ bearing ovules on the lower surface, based on Harris (1932a, fig. 28I, pl. 8, fig. 1); x3. the nucellus there was a distinct disc of vascular tissue. The megaspore membrane was up to 10 um thick (Rothwell, 1971). The microsporangiate organs (Jdanothekion) were radially symmetrical synangia borne abax- ially on pinnules (Fig. 15C). Each consisted of six to eight sporangia united for half their length. Distally they formed a tube and dehisced towards the center along a single median suture. The spo- rangia produced monosaccate pollen of the Ves- icaspora-type with a distal sulcus (Fig. 15E). The pollen wall is alveolate, similar to that of coni- fers Pollen occurs frequently in the micropyles of Callospermarion ovules, and a slender, branched pollen tube similar to that of extant conifers has been described (Rothwell, 1972). Germination was distal, and the tube is unlike the larger in- flated haustorial structures of extant cycads and Ginkgo (Rothwell, 1980: 92). It seems likely that the Callistophyton pollen tube indicates siphono- gamy. Vesicaspora pollen has been described with cellular contents preserved, and the four-celled axial row has been interpreted as three prothallial cells with a larger antheridial cell (Rothwell, 1981: 114). Microgametophyte development was 1985] CRANE-SEED PLANT PHYLOGENETICS 751 TABLE 7. Isolated organs of the three better understood peltasperms (after Townrow, 1960; Kerp, 1982). Locality Leaf Megasporophyll Microsporophyll South Africa Lepidopteris Autunia thomasii Antevsia (M. Triassic) stormbergensis Sweden & E eal Lepidopteris Peltaspermum Antevsia (U. Trias ottonis rotula zeilleri Nahe, Went un Callipteris Autunia miller- (L. Permian) conferta J therefore basically similar to that in Ephedra, some conifers, and cordaites (Florinites, Millay & Eggert, 1974; see character 1.1). Megagametophyte development is thought to have involved free nuclear divisions, followed by centripetal cellularization and differentiation of two archegonia. Cellular embryos have not been recognized. PELTASPERMS The clipe A range from the Lower rmi riassic and are known from leaves ioe conferta, Lepidopteris), mi- crosporophylls (Antevsia), and megasporophylls (Autunia, Peltaspermum) (Fig. 16) (Antevs, 1914; Harris, 1932a, 1937; Kerp, 1982; Lundblad, 1950; Mamay, 1975, 1978; Salmenova, 1979; Thomas, 1933; Townrow, 1960). Two species are known from all three organs (Table 7). The individual organs never have been found con- nected, but they all show highly characteristic blister-like swellings of the cuticle. This an er cuticular similarities, combined with associ- ation, provide evidence to link them together. Almost nothing is known of the stem of the peltasperms (Townrow, 1960: 340), but the leaves apparently were deciduous, and this suggests that the plants were woody. The leaves were bipin- nate (Fig. 16A) with bulbous frond bases. Pin- nules were borne both on the pinnae and directly on the leaf rachis. Stomata were anomocytic. Four species of megasporophylls have been de- scribed (Kerp, 1982): Autunia dzungaricum (Pel- taspermum dzungaricum Salmenova, 1979); A. milleryensis; A. thomasii (P. thomasii, Harris, 1937; Thomas, 1933; Townrow, 1960); and P. rotula (Harris, 1932a). They were dorsiventrally organized and branched in a single plane (Town- row, 1960). It is not known how they were at- tached to the plant. Each lateral “branch” (in- terpreted here as a pinna of the megasporophyll) terminated in a swollen lamina that bore seeds on the lower surface. In A. dzungaricum, A. mil- leryensis, and A. thomasii the lamina bore two seeds lateral to the insertion ofthe “branch” (Fig. 16B). In P. rotula the seed bearing lamina was peltate with the “branch” inserted centrally to the undersurface: ten to 12 seeds were arranged around the insertion of the “branch” (Fig. 16G, H). The lobed margin of the lamina in P. rotula was reflexed and attached to the “branch.” Only the ovules of P. rotula are known in detail (Fig. 16E, F). They were flattened and unitegmic with a long, bilobed, micropylar tube. The cuticle of the outer integument separated into two halves on maceration (Harris, 1932a: 69), and I inter- pret this as reflecting a platyspermic organi- zation of the seed. Nothing is known of seed vasculature. The nucellus was free from the in- tegument except. P s base. The megasnpre membrane w Microsporophylls (Antevsia) were bipinnate, with alternate primary branching in one plane (Fig. 16O). It is not known how they were at- tached to the plant. Secondary branching was irregular and frequently at 90° to that of the pri- mary branching. Secondary branches were short and bore four to 12 unilocular, pendulous pollen sacs that dehisced along a single longitudinal su- ture. Pollen was ellipsoidal, non-saccate, with a single distal sulcus (Fig. 16D). Fine structure of the pollen wall is unknown. Nothing is known of the gametophytes or em- bryo of peltasperms. GLOSSOPTERIDS An outline cladogram of glossopterids has been presented already (Fig. 8); only additional fea- tures thought to be general for the group are men- tioned here. The Glossopteris plant is thought to have been a large tree (Gould & Delevoryas, 1977; Pant, 1977a). The primary vasculature was eustelic, and a bifacial cambium produced secondary phloem and large amounts of secondary wood (Araucarioxylon). The wood was pycnoxylic, with 752 tracheids with multiseriate opposite or alternate circular bordered pits. The rays were uniseriate. Secondary xylem of identical structure is known to have been produced in the roots (Vertebraria, Gould, 1975). Axillary branching is known to have occurred. The leaves were borne in whorls or spirals, probably on long and short shoots (Pant, 19772). The leaves were simple and nar- rowly elliptical with a midrib and well-developed reticulate venation (character 4.1). Stomata were anomocytic. The ovules were borne on different kinds of megasporophylls (see characters 4.2, 4.5) in the axil of a leaf. The ovules of the silicified Austra- lian material (Gould & Delevoryas, 1977) were not described in detail, but they appear to have been pyriform, and some appear flattened (Gould & Delevoryas, 1977, fig. 7c). Those associated with permineralized Antarctic G/ossopteris ma- 6: 57). The Glossopteris ovules described from compressions (e.g., Pterygospermum, Pant & Nautiyal, 1960) also were flattened, with a single integument and a long micropylar tube. The in- tegument and nucellus were apparently free ex- cept at the chalaza. The nucellar cuticle was ro- bust, but a megaspore membrane up to 6 um thick also was present (Pant & Nautiyal, 1960). The microsporophylls were borne adnate to a Glossopteris leaf (character 4.3). The sporangia (Arberiella) had a finely striated wall and de- hisced by a single longitudinal slit. They pro- duced saccate pollen of the Protohaploxypinus type with a distal sulcus. The wall structure of glossopterid pollen has not been studied in detail but is interpreted here as probably alveolar, sim- es lular and contained either one (Gould & Dele- voryas, 1977: 392) or two archegonia (Schopf, 1976: 56). The embryos of Glossopteris are un- known. CAYTONIA Sagenopteris leaves range from the Upper based on a limited number of localities in east ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Greenland (Harris, 1933, 1937), eastern U.S.S.R. (Krassilov, 1977a), Poland (Reymanówna, 1973), Sardinia (Edwards, 1929), Sweden (Lundblad, 1948), and most importantly the Middle Jurassic of England (Thomas, 1925; Harris, 1940a, 1940b, 1941, 1945, 1951b, 1958, 1964). Three similar species are known from leaves, microsporophylls, and megasporophylls (Fig. 17, Table 8). The or- gans have never been found organically con- nected but are linked by association evidence, similarity of cuticular structure, and the consis- tent occurrence of Caytonanthus pollen in the micropyles of Caytonia ovules (Thomas, 1925; Harris, 1933, 19402, 1941, 1951b, 1964). Other genera of reproductive structures that may be related to Caytonia (e.g., Pramelreuthia Kráusel, 1949. Ktalenia Archangelsky, 1963) are not in- cluded in this discussion of characters. The stem of the Caytonia plant is known from several small twigs (Harris, 1940b, 1971), but they are sufficient to show that it was a woody shrub or tree with alternate leaves. The twigs were thicker than in most extant temperate trees and shrubs. They bore alternating zones of bud- scales and leaves. Buds occur in the axils of the leaves. Branching was sympodial, the apical bud ceasing to grow and being succeeded by two lat- eral buds to give forked branching as occurred for example in the bennettitalean Wielandiella angustifolia (Nathorst, 1909). The leaves (Sa- genopteris, Fig. 17A, B) were borne on raised “leaf cushions" and had a long petiole bearing four narrowly elliptical leaflets at the apex. Al- though superficially palmate, the leaflets were ar- ranged in two pairs (Fig. 17B). They each have a midrib and reticulate venation. Small leaves with broad stipule-like petiolar flanges and bud- scales also are known (Harris, 1940b). The leaves apparently were deciduous, with both leaflets and petioles shed. Stomata were anomocytic. The megasporophylls (Caytonia, Fig. 17E) were pinnate, bearing lateral “cupules” on short stalks. It is not known how they were attached to the stem. The “cupules” were reflexed, with a small lip on one side close to the stalk. They were all borne on one surface of the rachis, and all were reflexed 1 in the same direction. Differentiation of were on the upper (presumed adaxial) surface (Harris, 1940a). The fleshy and “‘berry-like”’ (Harris, 1951b) and shed individually. The “flesh” of the *cupule" con- sisted of a network of vascular strands, large 1985] CRANE-SEED PLANT PHYLOGENETICS 753 Tolua nna Wa dot FiGure 17. Morphology of the Caytonia plant.—A. Vi siad colpodes, based on Thomas (1925, pl. 15, fig. 50); x0.75.—B. S. aae baie of leaflet attachment — tion, based on Harris (1964, fig. 2H); x 4.— 2, 3 "D I C. Caytonanthus arberi, base arris (1941, pl. g. 3); x7. . Pollen from C. arberi, m on Townrow (1962b, fig. 3d, e); x 1,200. —E. Catonia nathorstii megasporophyll, based on Harris (1964, fig. 10A-C); x 5.— F. Caytonia “cupu le” containin sed on Reymanówna (1973, particularly text fig. 12E, F), x12.5.— eeds, ba E Caytonia “‘cupule,”’ longi yarn section, based on Reymanówna (1973, m text-fig. 12E, F); x 12.5.— H. C. nathorstii ovule longitudinal section, redrawn from Harris (1958, fig. 7); x1 754 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 TABLE 8. Isolated organs of the three best known species of the Caytonia plant (after Harris, 1941, 1951b). Locality Leaf Megasporophyll Microsporophyll U.K., Yorkshire Sagenopteris Caytonia Caytonanthus (M. Jurassic) colpodes sewardi oncodes Sagenopteris Caytonia Caytonanthus phillipsii nathorstii arberi E Greenland Sagenopteris Caytonia Caytonanthus (U. Triassic-L. Jurassic) nilssoniana thomasii kochii rounded cells, and branched sclereids (Reyma- nówna, 1973). Each *cupule" contained a “‘pack- et" of several small ovules surrounded by a com- mon cuticular membrane that was continuous with the outer cuticle of the cupule (Fig. 17F; Krassilov, 1977a; Reymanówna, 1973). The ovules were arranged with their micropyles ori- mc towards the mouth, below the lip of the pule." Narrow channels ran between the cu- ticles of the mouth connecting the micropyles with the outside (Fig. The ovules were flattened and unitegmic, with a narrow micropylar slit (Harris, 1940a, 1958; Thomas, 1925). The integument was free from the nucellus except at the chalaza and was sup- plied by a pair of vascular bundles (Reyma- nówna, 1973). Each bundle consisted of scalari- form tracheids that ran along the edge of the seed in the primary plane, almost to the micropyle. The integument and nucellus were strongly cu- tinized, but no acid-resistant megaspore mem- brane was detected (Fig. 17H; Harris, 1958). The microsporophylls (Caytonanthus, Fig. 17C) were pinnate, with irregular, short, lateral “branches.” It is not known how they were at- tached to the plant, but they were dorsiventrally flattened, with different cuticles on the upper and lower surfaces. On their lower surface the lateral synangia, w The locules dehisced longitudinally towards the inside, separating in the middle but remaining attached at the apex and base. The pollen (Vitrei- sporites, Fig. 17D) was bisaccate with a distal sulcus. Krassilov (1977a) has described mono- saccate grains. The pollen wall was alveolate with large cavities (Crepet & Zavada, pers. comm.). Although Caytonia was claimed originally to be an angiosperm (Thomas, 1925), Harris (1933, 1940a) demonstrated Vitreisporites pollen in the micropyles of Caytonia seeds. Pollination prob- ably involved a pollination drop mechanism that drew pollen along the channels of the mouth and onto the micropyles. Nothing is known of the male or female ga- metophyte of Caytonia, and the embryo is only known from “aleurone cells" (Harris, 1958). CORYSTOSPERMS The Corystospermaceae was established by Thomas (1933) for leaves, microsporophylls, and megasporophylls from the Middle Triassic Mol- teno Beds ofthe Upper Umkomaas Valley, Natal (see Anderson & Anderson, 1983). Some of Thomas' specimens have been subsequently re- interpreted (Townrow, 1962a), and additional material has been described from the Molteno flora of Zimbabwe (Lacey, 1976), the Upper Triassic/Lower Jurassic of Tasmania (Townrow 1965), the Triassic of New South Wales (Holmes & Ash, 1979), the Triassic of India (Pant & Basu, 1973, 1979; Srivastava, 1974), the Triassic of Argentina (Archangelsky, 1968; Petriella, 1979, 1980, 1981, 1983), and the Middle Jurassic of Yorkshire (Harris, 1964). The individual organs have never been found in organic connection but are treated together on the basis of consistent association, similarity of cuticular structure, and the occurrence of pollen from the microsporan- gia in micropyles of the ovules (Archangelsky, 1968; Harris, 1964; Thomas, 1933; Townrow, s 1965). woody plants, but their stems are ; poorly kon: Harris (1983a) de- scribed the stem of Pachypteris papillosa based on compressions, and Archangelsky (1968) has suggested that RAexoxylon (Fig. 18G; Archan- gelsky, 1968; Archangelsky & Brett, 1961) may be the stem of some Triassic corystosperms. Ex- ternally Rhexoxylon showed leaf bases and branch scars. It is not known whether axillary branching occurred. Internally there was a large pith sur- rounded by a ring of vascular segments with sec- ondary xylem developed both centripetally and centrifugally in relation to the ly a modification of a basic eustele (Beck et al., 1985] CRANE- SEED PLANT PHYLOGENETICS 755 FicunRE 18. Morphology of corystosperms.— A. Pachypteris papillosa stem with leaves, based on Harris (1983a, fig. 2); x0.25.—B. Pteroma thomasii synangium, abaxial view, redrawn from Harris (1964, fig. 66B); x25 —C.P é f Io 3 africanus, redrawn from Townrow (1962a, fig. 10F); x 600.—I. P. africanus, based on Townrow (1962a, fig. 1A- 5. 756 1982; Stewart, 1983). The primary xylem con- tains tracheids with spiral thickenings or sca- lariform pitting. The secondary xylem segments were pycnoxylic, although some axial paren- chyma occurred. The tracheids had uniseriate to triseriate, circular bordered pits on the radial walls. The rays were uniseriate (Archangelsky & Brett, 1961). Corystosperm leaves have been referred to a variety of genera, including Dicroidium (Fig. 18D), Xylopteris, and Pachypteris (Townrow, 1965). The leaf rachis was generally forked, with a simple or bipinnate arrangement of pinnae (Re- tallack, 1977: 255; Townrow, 1965). Stomata were anomocytic. Megasporophylls were dorsiventrally orga- nized and branched in a single plane (Fig. 18E). They consisted of a main axis with irregularly positioned lateral **branches" bearing several re- curved *'cupules." Thomas (1933) described “bracts” and pairs of "bracteoles" on these structures, and he therefore interpreted them as fertile branches. However, a similar interpreta- tion ofthe microsporangiate organs was not sup- 62a), who showed the bearing structures never have been reinvestigat- ed but are interpreted here as megasporophylls. It is not known how they were attached to the stem. The megasporophylls were highly variable, and lobes by clefts in the plane of branching, and lacked hairs or hair bases on the inner surface. Pilophorosperma had “cupules” with a single cleft and their inner surface covered by hairs. Sper- matocodon had “cupules” with no conspicuous clefts, which lacked hairs on the inner surface. In all three genera each “‘cupule” contained a single ovule. Other subsequently described co- rystosperm *'cupules" (e.g., Karibacarpon, La- cey, 1976; Holmes & Ash, 1979) also contain a single ovule. The ovules were flat (Fig. 18F), and although their vasculature is unknown, they had files of small rectangular cells arranged in longitudinal rows along their two margins (Thomas, 1933: 229). At their apex the ovules had two elongated micropylar lobes that were curved to one side in the primary plane. The curved micropylar tube was oriented away from the cupule stalk. De- tailed observations on the seed membranes of ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 corystosperms still are needed, but Thomas (1933: 228) noted “a well cutinized testa" and an "exceedingly delicate inner membrane." The ovules are assumed to have been unitegmic, and the megaspore membrane probably was not well developed. Microsporophylls (Pteruchus, Fig. 181) were branched irregularly and bore groups of pendu- lous pollen sacs on expanded distal laminae. The "branches" (rachises, Townrow, 1962a) were at- tached either in a single plane (Townrow, 1962a) or spirally (Pant & Basu, 1973, 1979; Srivastava, 1974; Thomas, 1933). It is not known how the main axis was attached to the plant. Townrow (1962a) emphasized the leaf-like nature of Pte- ruchus, and it is interpreted here as a micros- porophyll. Pollen sacs were ellipsoidal, uniloc- ular, and dehisced along a single suture. Pollen was bisaccate (Fig. 18H), with a single distal sul- cus, and very similar to that produced by Cayto- nanthus (Townrow, 1962b). The pollen wall was granular (Crepet & Zavada, pers. comm.; see also Taylor et al., 1984). Pollen has been described in the micropyle of one of the South African ovules (Thomas, 1933). The microsporophyll Pteroma thomasii Har- ris (Fig. 18B, C) from the Middle Jurassic of Yorkshire also is included in the Corystosper- maceae by Townrow (1965). On the basis of as- sociation and some similarity in cuticle, it is thought to be part of the Pachypteris papillosa plant (Harris, 1964). Pteroma is a pinnate struc- ture with oval or round fertile laminae with a ring of microsporangia embedded in their un- dersurface. The microsporangia were unilocular, dehisced by a single longitudinal slit and pro- duced bisaccate pollen. Nothing is known of the male or female ga- metophyte, or embryo of corystosperms. PENTOXYLON Knowledge of the Pentoxylon plant is based mainly on silicified material from the Jurassic of the Rajmahal Hills, India. Sahni (1948) gave the most detailed account and linked the ovulate heads (Carnoconites) with the stem (Pentoxylon, Fig. 19D) and the leaves (Nipaniophyllum, Fig. 19C) on the basis of association evidence and structural similarity. Microsporangiate organs (Sahnia, Fig. 19E) were described subsequently by Vishnu-Mittre (1953), and knowledge of the Pentoxylon plant was reviewed by Rao (1976, 1981). The other published reports of Pentoxy- 1985] CRANE-SEED PLANT PHYLOGENETICS 757 E Jc | E Ficure 19. Morphology of Pentoxylon plants. — A. Carnoconites cranwelliae, ovulate heads, based on Harris (1962, text-fig. 2B, fig. 1); x 2.5.— B. Carnoconites, longitudinal section through ovule, based on Sahni (1948, fig. 21); x10.—C. Nipaniophyllum raoi, redrawn from Sahni (1948, fig. 34a, b); x 1.—D. Pentoxylon sahnii, transverse section of stem showing vascular strands, based on Sahni (1948, fig. 9); x 8. — E. Sahnia microspo- rangiate “flower,” based on Vishnu-Mittre (1953, fig. 11) and Bose et al. (in press); x2.5. lon are based on ovulate heads (Fig. 19A) and toria (Drinnan & Chambers, 1985), New South leaves preserved as compressions from the up- Wales (White, 1981), and Queensland, Australia permost Jurassic or lowermost Cretaceous of New (Turner, pers. comm.). The description given be- Zealand (Harris, 1962, 1983b) and a variety of low and used in the phylogenetic analysis is based different organs in the early Cretaceous of Vic- on both the silicified and compression material. 758 The stem (Pentoxylon) was differentiated into long and short shoots, both with helically ar- ranged leaf-cushions showing leaf scars. In trans- verse section the stem typically had five vascular segments, but as in medullosans this is inter- preted as a modification of a fundamentally eu- stelic arrangement (Beck et al., 1982: 749; Stew- art, 1976). The secondary xylem was pycnoxylic with uniseriate rays and no axial parenchyma. Secondary xylem tracheids had uniseriate or bi- seriate circular bordered pits on their radial walls, although scalariform pitting has been reported in the microsporangiate structures and short shoots of Sahnia (Vishnu-Mittre, 1953). Leaf- traces arose in pairs, one from each of two ad- jacent vascular segments (Stewart, 1976). The leaves (Nipaniophyllum) were simple, strap- d of the taeniopterid-type, with a mid- originally described as “fundamentally Bennet- titalean" (Sahni, 1948: 56), but many are ap- parently anomocytic, and the morphology and ontogeny of the stomata require detailed reex- amination (Rao, 1976). The ovules were ue narhotropaus, nen cally arrange (Carnoconites compactum, C. un phi i or elongated heads (C. laxum). Each head of C. compactum d of about 20 i dee but there were more in C. cranwelliae and C. laxum No bracts or other structures were associated with these heads, nor were there interseminal scales as in Bennettitales. The ovulate heads were borne terminally on short shoots. In the Indian mate- rial a single pedicel produced several branches, each of which bore a single ovulate head. In the New Zealand material (C. cranwelliae) 12 sep- arate pedicels arose from the same point in an umbel-like arrangement, and each bore a single ovulate head. vules of Carnoconites compactum were flat- tened, with two integuments that were free from the nucellus except at the chalaza. The sclerotesta was bicarinate and distinctly flattened in trans- verse section. Immediately surrounding the sclerotesta was a distinct layer of dark cells. The remaining outer part ofthe “integument” formed a thick, fleshy *sarcotesta" interpreted here as an outer integument and probable *'cupule" homologue (see character 9.19). Each ovule was supplied by a single vascular bundle that pene- trated th al end; no ot vascular tissue has been reported in the ovules. oO = ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 The ovule cuticles of C. compactum and C. lax- um are unknown, but in C. cranwelliae the sar- cotesta had a thick outer cuticle. The inner cuticle of the integument was delicate but the nucellar cuticle was robust. There was no megaspore membrane preserved. The microsporophylls ae were borne in a whorl, or low helix, form surrounded themselves by deciduous bracts. Ac- cording to Vishnu-Mittre (1953), the microspo- rophylls were fused proximally to form a shallow cup, but according to Rao (1981), they were free at the base. The microsporophylls were numer- ous, and each bore ten to 20 spirally arranged, stalked unilocular sporangia. The sporangia were borne singly or in groups of two or four. Eac ulcate non-saccate pollen. Pollen wa stratification was probably granular (Taylor & Crane, in progress). Monosulcate pollen has been described in the micropyles of C arnoconites ovules. Germination may be the remains of megagametophyte tissue. No well-preserved embryos have been described, although Harris (1962) mentions a flat, cutinized plate in C. cranwelliae seeds, which he tenta- tively interpreted as the remains ofthe *aleurone layer" of the “endosperm.” BENNETTITALES An outline cladogram of the Benettitales al- ready has been presented (Fig. 12). Knowledge of other characters is very uneven, and in this section a general description of the Bennettitales is synthesized from features preserved in the dif- ferent taxa. The Bennettitales exhibited considerable di- versity in habit ranging from unbranched pachy- caul forms (Cycadeoidea, À orms that were more slender and highly branched (De- levoryas, 1975; Ischnophyton, Delevoryas & Hope, 1976; Williamsonia leckenbyi plant, Har- 4 mary vasculature was eustelic, and a bifacial cambium produced both secondary xy- lem and secondary phloem. The wood was dis- 1985] sected into numerous segments by broad rays, but individual segments were pycnoxylic with only uniseriate or biseriate rays. Tracheidal pit- ting was highly variable (Bose, 1953) ranging from multiseriate circular bordered pits to scalariform pitting (Sahni, 1932a). Leaf-traces pass directly into the leaf and do not girdle the stem as in cycads (Crepet, 1974: 178). Axillary branching is known to have occurred. Leaves generally were pinnate (see character 6.10) although there was some variation (Ash, 1975), and in Eoginkgoites there were several pairs of pinnae at the apex of the petiole giving a pseudopalmate arrangement (Ash, 1976, 1977). Stomata were paracytic, with distinctive cuticular thickenings (see character Ovules were grouped into **flower-like" heads, generally with surrounding bracts forming a *perianth" (but see Vardekloeftia, character 6.5). The ovules were small, generally less than 7 mm long, and frequently only 1-2 mm. Ovules were radially symmetrical (although flattened in Var- dekloeftia and Bennetticarpus wettsteinii), with two integuments that were free from the nucellus except at the chalaza (see p. 764). Frequently the nucellus was borne on a long stalk. Each ovule was supplied by a single vascular trace that ran to the base of the nucellus, but no other vascu- lature is known in the integument. Distally the integument was elongated into a long micropylar tube, and at maturity the integument was differ- entiated into four layers (Crepet, 1974: 157). The megaspore membrane was thin or not acid-re- sistant, but the cuticle of the nucellus and inner surface of the integument was well cutinized (Harris, 1954, 1969). The ovules were not in- dividually subtended by any other organs but are surrounded by numerous sterile interseminal scales that develop from primordia of bitegmic ovules (see character 6.1). The interseminal scales generally have a single vascular bundle. Microsporophylls were aggregated into “flow- er-like" heads surrounded by a "perianth" of bracts. The microsporangia are borne in various ways, but most of those known are aggregated into bivalved synangia (character 6.4). Pollen is monosulcate, boat-shaped, and more or less smooth-walled. The pollen wall is granular (Cy- cadeoidea, Crepet, pers. comm.; Taylor, 1973). The multicellul leg tophytes reported by Wieland (1906) are now interpreted as inter- nal folds in the pollen wall (Taylor, 1973). The female gametophyte apparently was monosporic, developing from the innermost of a linear tetrad CRANE-SEED PLANT PHYLOGENETICS 759 of megaspores (Cycadeoidea, Crepet & Delevor- yas, 1972). Little is known of the subsequent development in Cycadeoidea, but according to Sharma (1974), in Williamsonia free nuclear di- visions were followed by cellularization begin- ning at the micropylar end and proceeding to- ward the base. Cellular dicotyledonous embryos are known in Cycadeoidea (Crepet, 1974; Wie- land, 1906, 1916). GNETALES A cladogram for the Gnetales has been pre- sented already (Fig. 4). Only a general description of the Gnetales is given here as a basis for as- sessing their relationship with other seed plants. telar morphology of Gnetum and Wel- witschia is poorly known, but in Ephedra the primary vasculature is a eustele (Beck et al., 1982: 757-759). In Ephedra there is a single ring of vascular bundles with secondary xylem and phloem, but in Gnetum and Welwitschia there may be several concentric rings. The secondary xylem tissue consists of tracheids with circular bordered pits, vessels, and axial parenchyma. Axillary branching occurs in all genera, but leaf morphology is variable, ranging from the retic- ulate veined leaves of Gnetum to the strap-shaped parallel veined leaves of Welwitschia, and the scale-like or needle-like leaves of Ephedra. The morphology and arrangement of the ovu- late and microsporangiate reproductive struc- tures of the Gnetales have been discussed al- ready. The ovules generally are small, and the innermost layer of the various ovular coverings generally is regarded as a single integument. It is “fused” with the nucellus for about half its length. Apically the integument forms an elongated mi- cropylar tube. The ovules of Gnetum and Wel- witschia are not obviously flattened, although in Welwitschia there are a pair of minute lateral flaps at the base of the non-functional ovule in the microsporangiate flower. In some species of Ephedra (e.g., E. distachya) the integument orig- inates from a pair of dorsiventral primordia (Martens, 1971, fig. 24.4, 24.5), and at maturity there are two integumentary vascular bundles (Eames, 1952: 87). In some species of Ephedra, therefore, the ovules appear to be platyspermic, but the orientation of the primary plane is at 90° to the bract unlike the situation occurring in co- nifers. The acid-resistant megaspore membrane of mature seeds in the Gnetales is either ex- 760 tremely thin (about 1 um) as in Wel hia and Ephedra or lacking altogether in Gnetum (Erdt- man, 1965; Thomson, 1905) The microsporangiate structures of Welwitsch- ia and some species of Ephedra are arranged in a whorl, although in Gnetum the microsporangia are borne on a solid structure (character 1.7). Gnetalean pollen has already been described (character 1.5), and the evidence currently avail- able (Van Campo & Lugardon, 1973) suggests that the exine is comprised of a tectum, a gran- ular interstitium, and a thick, laminated inner layer. Microgametophyte development (charac- ter 1.1) and megagametophyte development (characters 1.2, 1.3) in the Gnetales have already been described. Embryogenesis varies in the three genera. In Ephedra there is a short free nuclear phase, but in Welwitschia embryogenesis is com- pletely cellular. In Gnetum there are reports of oth a free nuclear phase and completely cellular embryogenesis (Martens, 1971: 265 ANGIOSPERMS Most estimates place the total number of an- "A il species at around 250,000-300,000. us nu quura diversity and inadequate com- parative knowledge, makes the angiosperms ex- tremely dificult to deal with as a group in a broad s this (Hill & Crane, 1982). Although it i is widely pa that the Magno- liidae and some monocotyledons have retained a relatively large number of primitive characters, they are still an extremely heterogeneous assem- blage within which it is difficult to come to def- inite conclusions concerning likely plesiomor- phic features. This difficulty is well illustrated by the argument over whether the angiosperms are primitively vesselless (Young, 1981). In the case of the few higher dicotyledons that lack vessels, this can be reasonably hypothesized as secondary loss, but in the case of vesselless magnoliid plants, this is much more problematic. The issue can only be resolved by a well corroborated phylo- genetic analysis of relationships within the prim- itive angiosperms (Riggins & Farris, 1983). Until such a phylogenetic hypothesis is available for primitive flowering plants, the analysis provided here can only be regarded as preliminary. Char- acters of flowering plants are considered indi- vidually as part of the character discussion given below ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 PHYLOGENETIC RELATIONSHIPS OF EXTANT AND FOSSIL GYMNOSPERMS ANALYSIS OF CHARACTERS 9.1 Eustele. Beck et al. (1982: 701) define a eustele as “a stele with a hollow cylinder or tu- bular mass of tissue (i.e., with or without defin- able pith) and with discrete sympodia usually either as a discontinuous cylinder or in a scat- tered and dispersed arrangement." This kind of stele occurs in Archaeopteris and most seed plants and contrasts with the protosteles or siphono- steles typical of pteridophytes, including the aneurophytalean progymnosperms (Beck et al., 1982: 726). Fossil taxa with unknown stem anat- omy (peltasperms, Caytonia) are assumed for present purposes to have had a eustele. The an- giosperms too are interpreted here as fundamen- tally eustelic although this has been disputed by Tomlinson (1984) particularly for the monocot- yledons. A few seeds plants, however, are pro- tostelic, particularly among the “‘lyginopterid seed ferns” (e.g, Heterangium, Microspermopteris, Tetrastichia, Taylor, 1981a). Out-group com- parison with all pteridophytes, except Archaeop- teris, suggests that this may be the primitive con- dition for seed plants as a whole. On this basis Quaestora probably exhibits the primitive med- ullosan condition, the **polystelic" eustele hav- ing evolved within the group. The medullosans are therefore treated as protostelic in this anal- ysis. 9.2 Embryogenesis with a free nuclear phase. In extant bryophytes and pteridophytes all of the divisions of the zygote following fer- tilization involve the formation of cell walls (Sporne, 1970). Embryogenesis is cellular throughout. In embryos of extant cycads and Ginkgo a coenocytic phase with 250 or more nuclei may be produced before cellularization occurs. In Ephedra and most conifers there is a short free nuclear phase of approximately three zygotic divisions. The presence of a free nuclear phase in embryogenesis is interpreted (Hill & Crane, 1982) as a specialized feature of seed plants by out-group comparison with pteridophytes. In Welwitschia and angiospe Gnetum whether early embryogenesis is cellular or has a free nuclear phase is uncertain (Martens, 1971: 265). The primitive condition is assumed in this analysis. A few conifers have wholly cel- lular embryogenesis (e.g., Sequoia, Martens, 1971: 265), but these taxa are specialized within the 1985] conifer clade re 1982). I interpret this fea- ture as a secon advance within the group. As Chamberlain vna 342) suggested, there may be a correlation in seed plants between egg-cell size and the duration of the free nuclear phase. Embryogenesis for all fossil seed plants consid- ered is unknown but is treated here as having involved a free nuclear phase. In Archaeopteris, however, in line with other presumed pterido- phytic aspects of its reproductive biology (Pettitt, 1970), I have treated embryogenesis as being cel- lular throughout. 9.3 Single functional megaspore mother-cell per megasporangium. ith the exception of Archaeopteris the megasporangium in all of the taxa considered here produces one functional megaspore mother-cell. In all seed plants except Gnetum, Welwitschia, and a few angiosperms (character 9.28) one functional megaspore is sub- sequently formed, and the remaining three mega- spores abort. Under this interpretation the fe- male gametophyte develops entirely within the megaspore, and the tissues of the megasporan- gium are regarded as homologous with the nu- cellus. In heterosporous species of Archaeopteris and most heterosporous pteridophytes (e.g., Se- laginella) more than one megaspore is produced in each megasporangium. A few pteridophytes such as the extant fern Marsilea, the fossil lep- idodendroid genera Achlamydocarpon and Lep- idocarpon, the probable herbaceous Pennsylva- nian lycopod Miadesmia, and the fossil sphenopsid Calamocarpon also have a single functional megaspore per megasporangium, but the distribution of other characters (not analyzed here) indicates that these are not homologous with the seed plant situation. This is the widely accepted view (Stewart, 1983). 9.4 Integument. All of the taxa considered in this analysis except Archaeopteris have the yan in- tegument. This feature has been given various interpretations in different groups of seed plants (see characters 9.7, 9.19), but no comparable structure occurs in extant pteridophytes. Enclo- sure of the megasporangium in the fossil lyco- pods Lepidocarpon and Miadesmia is achieved by lateral extensions of the megasporophyll that extend around the equator of the megaspore. This is not homologous with the situation in seed plants (Stewart, 1983) where enclosure has apparently occurred along the polar axis of the megaspore. Micropyle. All seed plants considered in this study have a clearly defined micropyle, and CRANE-SEED PLANT PHYLOGENETICS 761 although the integument may have a pair of lat- eral “horns” (e.g., conifers, Ginkgo) or be elon- gated to form a long micropylar tube (e.g., corys- tosperms, Bennettitales, Gnetales), it is never deeply lobed around the micropyle as in early seed plants such as Archaeosperma (Pettitt & Beck, 1968; Pettitt, 1970). I interpret the unlobed micropyle as a derived feature that unites all the seed plants considered in this study (see also Stewart, 1983). Archaeosperma, the early seeds described by Gillespie et al. (1981), Genomosper- ma kidstonii, and G. latens (Long, 1959) all have a poorly differentiated micropyle that I interpret as the primitive condition. The lobed micropyle of some angiosperms (e.g., Hernandia) is clearly not homologous with the condition in early seed plants. 9.6 Linear tetrad of megaspores. Meiosis of the megaspore mother-cell in all extant seed plants produces a linear tetrad of megaspores of which only the inner develops, or female gametophyte development is tetrasporic (Gnetum, Welwitsch- ia, and a few angiosperms, character 9.28). Most angiosperms produce a linear tetrad (Sporne, 1974: 162). In the fossil taxa considered in this study probable linear tetrads of megaspores have been described only in Cycadeoidea (Crepet, 1974; Crepet & Delevoryas, 1972), but the ab- sence of haptotypic markings on the megaspores of all other taxa where this has been examined suggests that they also may have been produced in linear tetrads. Comparison of Lyginopteris with other "lyginopterid seed ferns” suggests that the megaspores may have developed from a tetra- hedral tetrad. By out-group comparison with ~ E Aa Cs > Q Q 2 . iu] M 3 R Q = £e [2 ° a © m Ro as) Q e 21 — e ^ — o ON SB Pettitt, 1970) I interpret Lagenostoma (Lyginop- teris) as primitive in this character. 9.7 Double vascular supply to ovules. Cycads and medullosans both have large ovules, which are supplied by a radially arranged double vas- cular system. The outer ring of bundles supplies the sarcotesta. In medullosans, the nucellus and integument are free except at the chalaza, and the inner ring of bundles supplies the nucellus; but in cycads where the nucellus is “fused” to the integument for most of its length, it is rarely clear whether the inner bundles are in nucellar or integumentary tissue. This double vascular supply to the ovules is not known to occur in any of the other taxa considered in this study. Rodin and Kapil (1969) have compared the ovule vasculature of Gnetum with that of Pachy- 762 testa, but under the interpretation given here (character 9.33) only the inner envelope ofa Gne- tum ovule is a true integument, and therefore the two outer layers do not have homologues in cy- cads or medullosans. In Lagenostoma (Lyginop- teris) there is a unitegmic ovule with a radial vascular supply inside a vascularized cupule. It has been suggested (Sporne, 1971a: 61; Wordsell, 1906) that this cupule may be homologous with the integument of cycads and medullosans and that their inner vascular system represents the remains of an "inner" integument. However, the "inner" integument of Lagenostoma is differ- entiated into a sarcotesta, a sclerotesta, and en- dotesta as in most gymnosperm seeds, including those of cycads and medullosans. There is no evidence in cycads and medullosans, other than the vasculature, for an additional tissue layer in the integument, and the "double integument” interpretation is not accepted in this paper (see also Stewart, 1983). Similarly the pad of vascular tissue at the base of Cardiocarpus and Mitro- spermum ovules is not taken to indicate the for- mer presence of an "inner integument" (Gra- ham, 1935). In peltasperms and corystosperms ovule vasculature is unknown but assumed no to have been double as in cycads and medullo- sans. Only the inner integument of angiosperms is interpreted in this study as homologous to the integument of other seed plants (character 9.19). Very few angiosperms have a vascularized inner integument (Sporne, 1974: 154), and I know of none in which a double vascular system has been described. 9.8 Axillary branching. Despite intensive study of anatomically preserved material, axil- lary branching has not been reported in progym- nosperms. Axillary branching has also never been demonstrated in extant cycads (p. 744). In all other plants considered in this study axillary branching is either known or is assumed (pel- tasperms, corystosperms) to have been present. 9.9 Saccate pollen. Saccate pollen or pre- pollen is generally regarded as the primitive con- dition in conifers and cordaites, but also occurs in ales lossoptrids Caytonia, and Orysto ate pollen is interpreted as a relatively speciale feature within seed plants, although it should be noted that saccate spores occur in some aneurophytalean progymno- sperms (e.g., Rellimia, Tetraxylopteris; Bonamo, 1977; Bonamo & Banks, 1967; Leclerq & Bona- mo, 1971; Scheckler & Banks, 1971) and Penn- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 sylvanian lycopods (Kosanke, 1969; Pigg & Rothwell, 1983). Saccate pollen reported in med- ullosans (Parasporites in Parasporotheca Dennis & Eggert, 1978) is treated here as specialized within that group. 9.10 Flattened ovules. one or two planes of symmetry occur in conifers, cordaites, uni iid and all of the “Me- sozoic seed ferns" (peltasperms, glossopterids, zie cory , Pentoxylon). Flat- tened ovules also occur in some lyginopterid seed ferns (not Lagenostoma) that are too poorly own to evaluate their (e.g.. Lyrasperma, Long, 1960). In cordaites, conifers, Callistophyton, Caytonia, and Ephedra there are two vascular bundles in the primary plane that run in the sarcotesta for most of their length, but in Ginkgo the bundles run within the endotesta. In the Bennettitales the ovules are small, with no integumentary vascular tissue, and are gen- erally regarded as radiospermic. However, in the primitive bennettitalean genus Vardekloeftia (Figs. 9B, C, 12) and in the Upper Triassic Ben- netticarpus wettsteinii (Krausel, 1949), the ovules are relatively large (7210 mm long) and flattened. ore detailed examinations of the ovules in primitive Bennettitales are required, but here I interpret this platyspermy as primitive for the Bennettitales on the basis that the ovules resem- ble those of Mesozoic seed ferns, Callistophyton, cordaites, and other groups mentioned above. The integument of Gnetum (“endotesta” of Ro- din & Kapil, 1969) has a well-developed radial vascular system; and Welwitschia ovules are un- vascularized and radiospermic. The bitegmic ovules of many angiosperms are bilaterally symmetrical and anatropous. How- ever, if the interpretations accepted in this paper are correct (character 9.19), then this is homol- ogous to a flattened recurved “cupule” contain- ing a single unitegmic ovule. A comparable sit- uation occurs in the corystosperms, and in this group the primary plane of the ovule is aligned in the same plane as the *cupule" stalk. Although many bitegmic ovules of angiosperms could be flattened “cupules,” each containing a flattened seed, there is no convincing evidence that the nucellus and inner integument of angiosperm ovules are fundamentally platyspermic. 9.11 Megaphyllous leaves organized on a pin- nate plan. Flattened ovules with witschia, and Ephedra. In these taxa leaf vena- 1985] tion is either univeined, dichotomous and or dichotomous and basically parallel. chotomously veined leaves of Kingdonia, Cir- caeaster, and many monocotyledons (Foster & Gifford, 1974) are interpreted as secondary ad- vances within the angiosperm clade. Megaphyl- lous leaves are not known to occur in progym- nosperms. 9.12 Narrowly triangular awl-shaped leaves. This feature already has been discussed in rela- tion to conifers (character 3.1). The triangular leaves in Ephedra and a few, presumably de- rived, angiosperms are only superficially like those of conifers, but in Cordaixylon, Mesoxy- lon, Lyginopteris, and Callistophyton scale-like leaves similar to those of conifers are occasion- ally produced on proximal parts of branches and associated with axillary buds (Rothwell, 1982a). These heteroblastic leaf series may be important in the origin of conifer foliage (Rothwell, 1982a), but in this analysis the consistent production of narrowly triangular awl-shaped leaves only oc- curs in Lebachia and extant conifers. 9.13 Megasporophylls borne on short fertile shoots in the axil of a bract or leaf. This feature already has been discussed in relation to conifers (character 3.3). The megasporophylls of conifers, cordaites, and Ginkgo are all borne on fertile shoots in the axil of a bract or leaf. In the Gne- tales the ovules are terminal. In peltasperms, Caytonia, and corystosperms it is not known how the megasporophylls were attached to the plant, but from their morphology it is unlikely that they were borne in a manner homologous with that of cordaites and conifers, and the unspecialized condition is assumed. In glossopterids (character 4.2) the megasporophylls were adnate to a sub- tending leaf or bract, but the morphological sit- uation and homologies are unclear. The interpretation of this character in flow- ering plants is dependent on whether the angio- sperm carpel is regarded as a modified simple megasporophyll (e.g., Doyle, 1978) or a com- pound structure formed from a fertile branch in the axil of a subtending leaf or bract (e.g., Steb- bins, 1974). This paper adopts the former inter- pretation (character 9.34). In some angiosperms (e.g., *Amentiferae," Chloranthaceae) the car- , pels are borne in the axil of a bract or leaf, but this is generally regarded as a secondary advance within the angiosperm clade (Walker & Walker, 1984). CRANE-SEED PLANT PHYLOGENETICS 763 9.14 Fertile axillary ovulate shoots aggregat- ed into an “inflorescence.” The fertile axillary ovulate shoots of cordaites and conifers are ag- gregated into an “inflorescence,” but in Ginkgo the fertile axillary shoot is single in the axil of a leaf. The situation in cordaites and conifers is not known to occur in other seed plants consid- ered in this study. The relationships of the enig- matic Southern Hemisphere conifers Buriadia and Walkomiella, and the possibility that they represent *the coneless ancestors of Lebachia- like conifers" (Pant, 1977b: 31) is not examined in this paper, although it is of interest to note that they may post-date Lebachia in the fossil record (Rothwell, 1982a; Pant, 1977b) 9.15 Pollen with distal germinal aper- ture. Pollen with a distal aperture (and there- fore, presumably, distal germination) occurs in all of the seed plants considered here, with the exception of medullosans, Lyginopteris, and that distal germination is a derived feature. The pollen of Lebachia (Potonieisporites) has a prox- imal monolete suture but also frequently has two curved folds in the exine of the distal surface (Scott & Chaloner, 1983). However, material de- scribed by Mapes and Rothwell (1984) suggests that germination in Potonieisporites was proxi- mal. In the Gnetales the distal aperture of Wel- witschia is accepted as the primitive condition in the group relative to the inaperturate grains of Ephedra and Gnetum (character 1. Although a few angiosperms have a proximal germinal aperture (Walker, 1974b), distal ger- mination is widely considered to be primitive in the group (Chaloner, 1970a; Doyle, 1978; Walk- er, 1974b), equatorial germination being a sub- sequent development within the clade (Doyle, 78) 9.16 Cordaites foliage. The characteristical- ly strap-shaped foliage of Cordaites is known at- tached to Cordaixylon dumusum (Rothwell & arner, 1984) and Mesoxylon priapi (Trivett & Rothwell, 1985). The leaves have dichotomous, more or less parallel veins and a characteristic internal anatomy with well-developed longitu- dinal fibrous ribs (Harms & Leisman, 1961). The leaves of Welwitschia are superficially similar but grow continuously from a basal meristem, an have much less well-developed sclerotic Paci (Martens, 1971: 94). 9.17 Primary axis of ovulate and microspo- 764 rangiate “inflorescences” dorsiventrally piu with bracts and fertile shoots four-ranked. Thi character occurs only in Cordaixylon and Me- soxylon among the taxa studied. 9.18 Siphonogamy. Zooidogamy occurs in extant cycads and Ginkgo. All spores of ipn plants with proximal germination produce m tile gametes (bryophytes, pteridophytes). This is the basis for inferring zooidogamy in those fossil taxa with proximal germinal apertures in their pollen (medullosans, Lyginopteris, Mesoxylon, Lebachia). Callistophyton is known to have pro- duced a distal pollen tube, which was probably involved in gamete transfer. In this analysis all other fossil plants with distal germinal apertures in the pollen are scored uniformly as plesio- morphic in this feature. 9.19 Ovules borne in a "cupule." The ovules of glossopterids, Caytonia, and corystosperms are enclosed or partly enclosed by the mega- sporophyll lamina to form a “cupule.” These Permian and Mesozoic “cupules” have frequent- ly been compared, or considered homologous, to the cupules of Upper Devonian, Mississipppian and Pennsylvanian seed ferns, such as Archaeo- sperma and Lyginopteris (Thomas, 1933, 1955; Townrow, 1960). Morphologically, however, there are at least two different kinds of structure. The cupules of Lyginopteris and other early pte- ridosperms are axially organized with respect to the long axis ofthe ovules they enclose. The stalk is axially attached and the construction of the upule may reflect an origin either from sterile deeply divided leaf (e.g., Archaeopteris). In glos- sopterids, Caytonia, and corystosperms, the stalk of the cupule is lateral with respect to the long axis of the enclosed ovules. This lateral arrange- ment is most easily interpreted as reflecting an origin from modified pinnae or pinnules, such as those of Autunia, which have become rolled or expanded in various ways to form “cupular” pouches. Doyle (1978: 383) expressed a similar view that the “cupules” of Permian and Meso- zoic seed ferns are more easily explained as “‘leaf- lets modified by various degrees of circinnate enrollment than as dir of the bell- shaped cupules of eee terids.” According to the interpretation of “cupules” G may be interpreted as a uniovulate “cupule.” The idea that the outer integument in a bitegmic an- giosperm ovule is homologous to the “‘cupule” ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 of “Mesozoic seed ferns” has been advocated by numerous authors (Doyle, 1978; Stebbins, 1974; Stewart, 1983). This view is supported by the recurved “‘anatropous” arrangement of the “cu- pule" in Caytonia and corystosperms, the zig- zag micropyle of many bitegmic angiosperm ovules, and evidence that the funicle, chalaza, and outer integument are part of a single devel- opmental unit (Bor, 1978), while the inner in- tegument and nucellus are part of another. The complex net-like vasculature of the outer integ- ument of many angiosperm ovules is a further similarity with the “cupules” of “Mesozoic seed ferns." The homology of the “cupules” of Cay- tonia and corystosperms with angiosperms is ac- cepted in this paper as providing the most straightforward explanation of the second integ- ument that does not require a de novo origin (cf. Long, 1966). Unitegmic ovules occur in rela- tively specialized flowering plants and are inter- preted as a secondary development within the angiosperm clade. In Pentoxylon and Bennettitales the homolo- gies of the outer integument are more difficult to determine. In Vardekloeftia and Bennetticarpus crossospermus (Bennettitales) there is a distinct outer layer around the inner integument, which has been variously termed the “‘cupule”’ or “‘mi- cropylar plate" (Harris, 1932b) and is interpreted here as the outer integument of tained more than one ovule (Harris, 1932b: 108). In Cycadeoidea Wieland (1906: 120) has de- scribed a “cup-shaped supporting basal husk,” and in Bennettites albianus Stopes (1918) has described a “cupule or aril of elongated tubular cells." I suggest that both of these layers are ho- mologous with the *cupule" of Vardekloeftia and Bennetticarpus crossospermus and the outer layer of Bennetticarpus wettsteinii seeds (Krau- sel, 1949). In Pentoxylon I interpret the bennet- titalean ‘“‘cupule” as represented by the well-de- veloped outer fleshy ‘“‘sarcotesta” around each ovule, and the cuticle obtained from this layer in Pentoxylon cranwellii has been compared with that of the "cupule" in Vardekloeftia (Harris, 1962: 24) On the interpretations of “cupules” given above the outer integument in the ovules of Ben- nettitales and Pentoxylon may be interpreted in three different ways. Either it is homologous with the cupule of Lagenostoma (Lyginopteris), or ho- 1985] mologous with the ''cupule" of glossopterids, Caytonia, corystosperms, and angiosperms, or it is independently derived and not homologous to either. In this paper I exclude the first possibility. Although the ovules of Bennettitales, Pentoxy- lon, and Lagenostoma are all orthotropous, they differ considerably in other features. The cupule of Lagenostoma is lobed, heavily vascularized, does not enclose the ovule, and is thought to have been borne on well-developed megaphylls. ““Cu- pules” of primitive Bennettitales and Pentoxylon are unlobed, unvascularized, enclose the ovule, and are not borne on megaphylls. > by ES different methods of scoring in the data matrix for this paper. In one analysis seis l, p. 770), the bennettitalean and Pentoxylon ovule is interpreted as non- Fidis denis ien the “cupules” of glossopterids, Caytonia, and corys- tosperms. In the second analysis (cladogram 2, p. 770) the homology of these different “cupules” (but not the cupule of the Lyginopteris plant) is accepted. The details of adjusting the data matrix for these two different interpretations are dis- cussed later in the paper (p. 774). In the Gnetales the only morphological fea- tures that may be interpreted as evidence for the presence of a “cupule” are the regular develop- ment of two nucelli within a single integument in some species of Ephedra (Thoday & Berridge, 1912), the two lateral flaps of tissue at the base of the non-functional ovule in microsporangiate ments" (Pearson, 1915). In the absence of better evidence the Gnetales are treated as lacking a *cupule" in this analysis. 9.20 Megaspore membrane thin. Com- parative information on the megaspore mem- branes of gymnosperms is inadequate. Currently available data suggest that a thick, well-devel- oped acid-resistant megaspore membrane, like that typical of pteridophytes, is primitive within seed plants (Hill & Crane, 1982) and that the weakly developed membrane in Caytonia, co- rystosperms, Pentoxylon, Bennettitales, and an- giosperms is a specialized condition (Erdtman, 1957, 1965; Harris, 1954; Th .T megaspore membrane in these plants is only weakly resistant to maceration, and frequently this seems to be correlated with the presence of a well-developed nucellar cuticle (Harris, 1954). In conifers the megaspore membrane varies in thickness between genera. A thin megaspore CRANE-SEED PLANT PHYLOGENETICS 765 membrane, however, is most common in rela- tively advanced taxa (Erdtman, 1965; Miller, 1982). In angiosperms the megaspore membrane can be detected with transmission electron mi- croscopy (Mogensen, 1978) but is not acid-re- sistant. 9.21 Granular pollen wall. The pollen of all the seed plants considered here has two well- defined wall layers, the outer of which is either alveolar or granular in construction (Doyle et al., 1975). Within the alveolar forms there are two basic types; one in which the alveolae are com- prised of small, typically densely crowded cavi- ties (““cycad type," Doyle et al., 1975), the other in which the alveolae are larger and more highly organized (“‘pinaceous type," Doyle et al., 1975). The microspores of Archaeopteris (Pettitt, 1966) and the pollen of corystosperms, Bennettitales, and Gnetales all have a granular wall structure. Pollen of cycads and medullosans is of the al- veolate type, with small cavities, and the pollen of conifers, cordaites, Ginkgo, Callistophyton, peltasperms, glossopterids, and Caytonia is of the more highly differentiated alveolate type (Crepet & Zavada, pers. comm.; Doyle et al., 1975; Millay & Taylor, 1976). Pollen of Lygi- nopteris has not been studied with transmission electron microscopy but is assumed to have had homogeneous wall as in the pollen of Crosso- theca. Pollen of Pentoxylon is currently under investigation (Taylor & Crane, in progress). Pre- liminary results from transmission electron microscopy suggest that the pollen wall is granu- ar. Granular pollen wall € occurs in orniine Tustandacess. Beni it is erence as a secondary modification within flowering plants. However, granular exines also occur in man magnoliid families (e.g., Amborellaceae, Anno- naceae, Canellaceae, Magnoliaceae) (Walker, 1976). Walker (1976: 278) suggests that these granular exines represent a stage in the origin of tectate wall structure from a homogeneous, walled, atectate primitive condition. However, in the gymnosperms studied in this analysis a homogeneous exine occurs only in Lyginopteris, and the basic pollen wall type appears to be the alveolar condition with granular exines a rela- tively specialized feature seen in corystosperms, Bennettitales, Gnetales, and, presumably as a secondary advance, in some conifers (e.g., Ar- aucariaceae, Taxodiaceae). In the context of seed plants as a whole, I accept the hypothesis that 766 granular pollen wall stratification is primitive within angiosperms (see also Doyle, 1978: 375- 376) 9.22 Uniovulate "cupule." The "cupules" of Caytonia and the glossopterids considered here contain several ovules. In the glossopterid Den- kania (Pant, 1977a; Surange & Chandra, 1975) the *cupules" apparently contain a single ovule, but this is interpreted as a specialized feature within the glossopterid clade. According to the interpretation of character 9.19, the corysto- sperms, Bennettitales, Pentoxylon, and angio- sperms also have a single ovule per “‘cupule.” The homologies of “cupules” in all of these taxa are treated in two different ways (p. 770) but are not considered homologous with the cupules of Lyginopteris and other early seed ferns, for rea- sons given earlier (character 9.19). 9.23 Microsporophylls forming OWers. ' The microsporophylls of the Bennettitales, Pen- toxylon, Ephedra, and Welwitschia are aggregat- ed together in a whorl or pseudowhorl to form cup-like **flowers" quite distinct from the pollen cones of conifers. In Bennettitales, Welwitschia, Ephedra, and angiosperms they are clearly ar- ranged in a whorl or tight helix (see also character 9.33). The microsporophylls are similarly aggre- gated in Pentoxylon but details of their arrange- ment are uncertain. It is not known how the mi- crosporophylls of coryst Caytonia, glossopterids, and Peltaspermum were arranged on the plant, but in this analysis I have assumed that they were not aggregated into “flowers” (ple- : "cupule" erect. The uni- ovulate “cupules” of corystosperms like the mul- tiovulate **cupules" of Caytonia were recurved, and I interpret this as the primitive condition (see character 9.19). In Bennettitales and Pent- oxylon the uniovulate **cupules" are borne erect. In flowering plants the most widespread, and ap- parently primitive, configuration for the bitegm- ic ovule is anatropous. This is consistent with cur, these are considered as specializations with- in the flowering plant clade. 9.25 Megasporophylls "unicupulate." In the Bennettitales and Pentoxylon the presumed homologues of the megasporophylls consist of a single *cupule" (character 9.19). In other Perm- ian and Mesozoic plants with “cupules” (Cay- tonia and corystosperms) there are several “cu- ye ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 pules" per megasporophyll, and in Glossopteris this is also interpreted as the primitive condition (character 4.5). In angiosperms the primitive ian ird ie to b ds (Bai- ley & Swamy, 1951). Under the iE adopted in this paper, each carpel would there- fore consist of several cupules, perhaps originat- ing in the manner suggested by Doyle (1978). Although some of the carpels in the Magnoliidae (e.g., Chloranthaceae) contain only a single ovule, they are interpreted here as a secondary modi- fication within flowering plants (Walker & Walk- er, 4 9.26 "Unicupulate" megasporophylls aggre- gated to form heads. In Pentoxylon and Ben- nettitales the “unicupulate’’ megasporophylls ge ue 9.19, 9.25) are aggregated together to m heads. No comparable arrangement occurs in any other seed plant considered in this anal- sis. 9.27 Male gametophyte of three or four nu- clei. In gymnosperms, a male gametophyte of four nuclei occurs only in Gnetum, Welwitschia, and some conifers (character 1.1). In angi sperms the male gametophyte is even less exten- sive and undergoes two mitotic divisions to pro- duce three cells. 9.28 Female gametophyte tetrasporic. In Gnetum and Welwitschia (character 1.2), the fe- male gametophyte is tetrasporic. Tetrasporic Bie ae are uncommon in angio- rms (Sporne, 1974: 162) and are interpreted asa dir development within the flowering plant clade. 9.29 Archegonia absent. Asin Gnetum and Welwitschia (character 1.3), no archegonia are differentiated in the female gametophyte of an- giosperms. However, the megagametophytes in these two groups differ markedly in both devel- opment and mature structure. For these reasons they are not accepted a priori as homologous in this analysis. 9.30 Embryo with "feeder." There is no structure comparable to the “feeder” of Gnetum and Welwitschia (character 1.4) in the embryo of angiosperms or any other seed plant. 9.31 Ribbed pollen. Ribbed pollen superfi- cially similar to that of Ephedra and Welwitschia (character 1.5) occurs in some angiosperms (e.g., tectate columellate wall rather than by granular 1985] exine as in Ephedra and Welwitschia (Trevisan, 1980). Ribbed pollen known to occur in Acan- thaceae is also interpreted as a secondary devel- e. ration plates. Vesels occur in t cter 1.6) and most angiosperms. Although they have also been reported in the Bennettitales (Krassilov, 1984) they are not known to be general within that group. Typically the vessels of Gnetales and ngiosperms have been regarded as indepen- dently derived from tracheids, with circular bor- dered pits and scalariform pitting, respectively (Cronquist, 1968). It also has been generally ac- cepted that vessels arose within the angiosperms and that magnoliid angiosperms that lack vessels are primitively vesselless. However, Young (1981) has recently questioned whether the an- primitively vesselless. His ana alysis of relationships within the Magnoliidae suggests that it is more parsimonious to invoke loss of vessels rather than the multiple vessel origins, but this should not be taken to imply that an- giosperm and gnetalean vessels are necessarily homologous. Muhammad and Sattler (1982), however, have described *'scalariform" and “sclaroid” perforation plates in Gnetum, which further blur the distinctions between angiosperm and gnetalean vessels. Despite these difficulties the traditional view, that vessels of Gnetales and angiosperms are non- mologous, is adopted in this paper. À more detailed evaluation of relationships within the Magnoliidae is required to evaluate the system- atic position of vesselless taxa. Also, the simi- larities in perforation plates, of Gnetum and an- giosperms described by Muhammad and Sattler (1982), involve taxa that are regarded as rela- tively derived within flowering plants (e.g., var- ious *Amentiferae"). 9.33 Microsporangiate and ovulate "flowers" with opposite pairs of bracteoles. The repro- ductive structures of the Gnetales and several angiosperm groups are arranged on an opposite and decussate plan (character 1.7). In many cases, such as the "*Amentiferae," this arrangement is thought to be a secondary modification within the flowering plant clade. This probably also is the case in more primitive angiosperms such as Chloranthaceae (Walker & Walker, 1984, fig. 9.34 Bitegmic ovule enclosed or partially en- closed by a structure with a pollen-receptive stig- matic surface external to the second integu- CRANE-SEED PLANT PHYLOGENETICS 767 ment. Several living and extinct gymnosperms show total or partial enclosure of a unitegmic ovule (e.g., Caytonia, corystosperms, Bennetti- tales, glossopterids, and some conifers), and some (e.g., Tsuga and perhaps Leptostrobus) have a pollen-receptive surface external to the micro- le. However, no known gymnosperm has an enclosed, or partially enclosed, bitegmic ovule and a pollen-receptive stigmatic surface external to the outer integument. It is assumed here that those angiosperms with unitegmic ovules rep- resent a secondary modification within the flow- ering plant clade. Numerous mechanisms have been suggested for the enclosure of angiosperm ovules and the formation ofa simple conduplicate carpel. In this paper the hypothesis is adopted that the carpel represents a *cupulate" megasporophyll such as occurs in Caytonia and corystosperms, perhaps modified by progenesis (Doyle, 1978). 9.35 Female gametophyte of 4-16 nu- clei. In all seed plants where it is known, the female gametophyte is much more extensive than in angiosperms. 9.36 Double fertilization. The process of double fertilization in angiosperms that results in the formation of a zygote and endosperm nu- cleus does not occur in other seed plants. The closest approach to the angiosperm situation is in Ephedra, in which one of the male gametes fuses with the ventral canal nucleus of the ar- chegonium, but this second zygote undergoes no further divisions (Moussel, 1978; Sporne, 1971a: — oo 9.37 Laminate endexine lacking. All gym- nosperm pollen that has been studied in sufficient detail has a laminate endexine. In angiosperms to the apertures (Doyle et al., 1975), and some angiosperms appear to lack any endexinous layer (Walker, 1974a). Lugardon and LeThomas (1974) have suggested that the laminated endexine of nosperms is in fact homologous to the foot layer (innermost ektexine) of angiosperm pollen and that the endexine of angiosperms arose de novo (Zavada, 1984). If this interpretation is ac- cepted, then the apparent endexinous lamina- tions of some Annonaceae (Walker, 1976) and Trimeniaceae (Sampson & Endress, 1984) are not homologous to those of gymnosperms. In- terpreting this complex character is difficult (Hill & Crane, 1982: 344—345) and requires further investigation aimed particularly at clarifying general patterns of pollen wall ontogeny. gq 768 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 TABLE 9. Data matrix for characters of seed plants: apomorphic characters marked +, plesiomorphic char- acters marked —, not applicable marked NA. Cladogram 1 (Fig. 20) was constructed from version one of the data in which Bennettitales and Pentoxylon were scored as plesiomorphic for characters 9.19 and 9.22; and character 9.25 was excluded as redundant. Cladogram 2 (Fig. 22) was constructed from version two of the data in which Bennettitales and Pentoxylon were scored apomorphic for characters 9.19, 9.22, and 9.25. See text for further discussion of these two alternatives. Ex- Lygin- Cor- Le- tant piles Medul- opte- daixy- Mes- bach- Coni- Character eris losans Cycads ris lon oxylon ia fers 1. Eustele + = + + + + + + 2. Embryogenesis with a free nuclear phase =? +? + +? +? +? +? + 3. Single functional megaspore mother- cell per megasporangium = + + + + + + t 4. Integumen — + + + + + + + 5. Micropyle NA + + + + + + + 6. Linear tetrad of megaspores - +? + _ +? +? +? + 7. Double vascular supply to ovules NA + + _ — — -— _ 8. Axillary branching = + — + + + + + 9. Saccate > NA — _ — + + + + 10. Flattened ovules NA _ - + + + + 11. sasaqa salka leaves organized on a pinnate plan = + + + = ~ = = 12. Narrowly triangular awl-shaped leaves _ = = — — _ + + 13. Megasporophylls borne on short fertile shoots in the axil of a bract or leaf = = _ -— + + + + 14. Fertile axillary ovulate shoots aggregat- ed into an “inflorescence” NA NA NA NA + + + + 15. Pollen with distal germinal aperture = = + = + _ _ + 16. Cordaites foliage = — — _ + + — — 17 mary axis of ovulate and microspo- rangiate “inflorescences” dorsiventrally flattened with bracts and fertile shoots four-ranked NA NA NA NA + + = _ 18. Siphonogamy NA =? B =? zT =? =? + 19. Ovules borne in a “cupule” NA — _ — = _ = _ 20. Megaspore membrane thin = _ _ — _ — _ _ 21. Granular pollen wall + - — +? — — — -= 22. Uniovulate “cupule” NA NA NA NA NA NA NA NA 23. Microsporophylls E "flowers" — — = = _ _ _ — 24 iovulate **cupule" NA NA NA NA NA NA NA NA 2 egasporophylls deam NA NA NA — NA NA NA NA 5. M 26. "Unicupulate" megasporophylls aggre- gated to form heads NA NA NA NA NA NA NA NA 27. Male gametophyte of three or four nu- clei =? —? = =? — _ —? = 28. Female gametophyte tetrasporic = =? _ =? =? =7 —? _ 29. Archegonia absent =? Es — = = —? =? = 30 ith “feeder” =? =} = =? =? zy =? = 31. Ribbed pollen — — = — = — _ _ 32. Vessels with porose perforation plates = — = -= i _ _ _ 33. and ovulate “‘flow- with opposite pairs of bracteoles NA NA NA NA NA NA NA NA : Mies ovule enclosed or partly en- closed by a structure with a pollen-re- ceptive stigmatic surface external to the second integument NA NA NA NA NA NA NA NA W A 1985] CRANE- SEED PLANT PHYLOGENETICS 769 TABLE 9. Continued. Cal- Glos- Co- listo- Pelta- sopter- Cay- rysto- Bennet- Pent- - Ephed- Angio- Ginkgo phyton sperms ids tonia sperms titales oxylon Gnetum witschia ra sperms + + +? + +? + + + + + + + + +? +? +? +? +? +? +? + _ + - + + + + + + + + + + + - + + + + + + + + + + + + + + + + + + + + + + + + + 4? +? +? +? + +? NA NA + + € = P P ass —? = = = e. CR - + + +? + + +? - + + + + + - + - + + + - - - - _ _ + + + + + + + + - - + _ = + + + + + + + + - - + + — —? —9 —? _ = > ss = = = NA NA NA? NA? NA? NA? NA NA NA NA NA NA + + + + + + + + + + + + NA NA NA NA NA NA NA NA - - - NA = + = = =? =? =? =a + + + + - - _ + + + —/+ —/+ - - - + _ _ - E + + + + + + + + - = - _ _ + + + + E + + NA NA NA - - * —/+ —+ NA NA NA + _ _ -? -? -? -? + + = + + + NA NA - - - - + + NA NA NA - NA NA - - - - -* —+ NA NA NA >- NA NA NA NA NA NA + + NA NA NA NA =: = =f m: =? =? =? —? + + — + — —? -? —? -? —? — —? + + — — — — —? — —? =? =? =? + + — - = = 9 xx —? -" = —? + + I = _ _ -? - -? - _ - + + + - NA NA NA NA? NA? NA - - + + + = NA NA NA NA NA = - > NA NA NA + 770 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 TABLE 9. Continued. Ex- Lygin- or- L tant plier Medul- t daixy- es- bach- Coni- Character eris losans Cycads ris o oxylon ia fers 35. Female gametophyte of four to 16 nu- clei 36. Double fertilization 37. Laminate endexine lacking 38. Axially aligned companion cells de- rived ontogentically from the same mother-cells as the sieve elements 9.38 Axially aligned companion cells derived ontogenetically from the same mother-cells as the sieve elements his sner apparently oc- curs only in angiosperms and is not known in any other group of seed a (Hill & Crane, 1982 CLADOGRAMS Cladograms were generated from the data ma- trix using the PAUP Program (Phylogenetic Analysis Using Parsimony) of D. L. Swofford with the global branch swapping and MULPARS option. Cladograms were rooted by incorpora- tion of a hypothetical ancestor scored as plesio- morphic for all characters. Two different forms of the matrix (Table 9) were a to generate two different nea passa as follow Cladogram 1 (Fig. 20). dens 1 was ence from version one of the data matrix in which the “‘cupules” of Bennettitales and Pent- oxylon were not coded as homologous to the "cupules" of glossopterids, Caytonia, corysto- sperms, and angiosperms. That is, Bennettitales and Pentoxylon were coded as plesiomorphic for characters 9.19 and 9.22. Character 9.25 was ex- cluded as redundant (coded by character 9.24) giving in a total of 37 characters. The analysis resulted in five equall (61 character state transitions) all with en sim- ilar topologies. A strict consensus tree (Rohlf, 1982) derived from those five cladograms is giv- en in Figure 20. This kind of consensus tree pre- serves only those aspects of tree topology that occur in the five original trees. In this case the consensus tree only resulted in one polychotomy. Cladogram 2 (Fig. 22). Cladogram 2 was generated from version two of the data matrix in which the “cupules” of Bennettitales were cod- ed as homologous to those of glossopterids, Cay- tonia, corystosperms, and angiosperms. That is, Bennettitales and Pentoxylon were coded as apo- morphic for characters 9.19, 9.22, 9.24, 9.25, giving a total of 38 characters. The analysis re- sulted in ten equally parsimonious cladograms (62 character state transitions) all with very sim- ilar topologies. A strict consensus tree derived from these ten cladograms is given in Figure 22. DISCUSSION AND INTERPRETATION (GYMNOSPERMS) The pattern of relationships in cladograms 1 and 2 is very similar. In both, seed plants are interpreted as a monophyletic a well-differentiated micropyle (character 9.5) surrounding a megasporangium with a single functional megaspore mother-cell (character 9.3) and having a free nuclear phase in embryogenesis (character 9.2, secondarily cellular in Welwitsch- Lyginopteris is resolved by both cladograms as the sister taxon to all other seed plants con- sidered. However, the characters on which this division is based are not strong. Distal pollen aperture (character 9.15) is particularly problem- atic (see below), and although Lyginopteris prob- ably had a tetrahedral tetrad of megaspores, the distributrion of tetrahedral and linear tetrads in other fossil plants is not well known. A close relationship between the Carbonifer- ous pteridosperms and cycads has been recog- nized by many authors. Arnold (1953) and Scott (1923) suggested that cycads were closely related 1985] TABLE 9. Continued. CRANE-SEED PLANT PHYLOGENETICS 771 a Glos- Co- listo- Pelta- sopter- Cay- rysto- Bennet- Pent- Wel- Ephed- Angio- Ginkgo phyton spe ids tonia sperms titales oxylon Gnetum witschia ra sperms — — —? — —? —? —? —? — — — + a] —9 —9 29 —? —9 —9 —? DE = = $ = _9 —9 —? —? —9 —9 —? = = = + m —? —? —? —? —? —? —? x _ = En to lyginopterid pteridosperms, while Delevoryas (1955), Stewart (1983), and Wordsell (1906) have favored a closer, even ancestor-descendant, re- lationship from medullosans. Cladogram 1 sup- ports the view that medullosans and cycads are sister taxa based on the double vascular system in the ovule (character 9.7). This assumes that the Lagenostoma cupule has no homology in the cycad ovule, and that the chalazal pad of vascular tissue in the ovules of conifers, cordaites (Cor- daixylon, Mesoxylon), and Callistophyton is not the remains of a previous inner integument (see character 9.7). In cladogram 2 the position of cycads and medullosans relative to each other, and relative to all other seed plants considered (except ipis nopteris), is not resolved. Both cladograms suggest that the apparent xe of axillary branching in cycads ( .8) is the result of secondary loss, and that s. branch- ing is a potential seed plant synapomorphy. Loss of axillary branching may be one result of the specialized pachycaul habit of extant cycads. The relationships of Spermopteris, Phasmatocycas, and Archaeocycas to cycads (Mamay, 1976) are discussed later in this paper. Similarly, both cladograms suggest that the protostele of Quaes- tora is reduced from the eustele that characterizes all other seed plants considered. This conclusion may have to be modified if » could be demon- strated that cycads and were closely related to aneurophytalean progymnosperms (protostelic) as suggested by some authors (e.g., Stewart, 1983 The evidence for a close relationship between conifers and cordaites was presented convinc- ingly by Florin (1951) and has been accepted widely by subsequent authors (Mapes & Roth- well, 1984; Rothwell, 1982b). Recently, how- ever, Beck (1981: 193) has suggested “that the Cordaitales and Lebachiaceae represent inde- pendent lines of evolution originating from Ar- chaeopteris or some closely related, similar pro- gymnosperm." This hypothesis would require at least a biphyletic origin of seed plants that is not supported by the analyses in this paper. The cladograms reflect the views of Florin and other authors. They suggest that conifers and cordaites are more closely related to each other than either is to Archaeopteris on the basis of several derived characters: single functional megaspore mother- cell per megasporangium (character 9.3), integ- ument (character 9.4), micropyle (character 9.5), axillary branching (character 9.8), saccate pollen (character 9.9), flattened ovules (character 9.10), megasporophylls borne on short fertile shoots in the axils of bracts (character 9.13), and ovulate short shoots aggregated into an “inflorescence” (character 9.14). Rothwell (1982b) has similarly emphasized the importance of these characters in his analysis of Beck's hypothesis. The mi- “inflorescence” of Cordaites is a 8 ge p 3 z ecludin sean (Beck, Tn ena 1977b). In cladistic terms this character would be a synapomorphy of cor- daites but would not obviate a close relationship with conifers. The difficulty only arises when the relationship between the two groups is viewed as that of an ancestor and descendant. Similar arguments could be applied to other, possibly unique, specializations of cordaites listed by Beck (1981: 215) (leaf morphology, bilaterally sym- metrical distichous inflorescences). The closest relatives of Ginkgo have long been thought to be conifers and cordaites (Chamber- lain, 1935; Emberger, 1954; Pant, 1977b), and this is supported by both cladograms in which Ginkgo is placed as the sister taxon to the conifer 772 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 = z o JP" £ RP. zŠ u ® z Š = > O & ú Ú z < < = C < = z I Z u a < o I = n. ul [ 7) > o ° o. a = o = E Oo u S F o w wu s € O wù a < o F ` oO < Y uc d 9 = > I = o - a O Z F L x - C€ Ó < O 3 QO < x O z o L 7? Qo G uo 5 F a o O G AO KE o m F z -J 3 O > t Z z u J r O cC X uU > O u y x = < u 3 < O u u 2 u a Z < l Z O O Z j u OG O a G O O m a G 3 u < | | | | Í © © © @ © © (is) ORORO G) © I "a @ @ L @ - © @ T Ll T 4 a © (2) C @ W (s © € a ges [ | 33 OR e (8) qQ ( o O © | J | ë @ T L ® EN Qi € d URE20. Relationships beween major groups of seed plants. Strict ived from five equally Md cladograms constructed from version one of the data matrix. In this lanier the “cupules” of Bennettitales and Pentoxylon weri coded as homologous x the "cupules" of glossopterids, Caytonia, corystosperms, and ea ad pure ips and Pentoxylon were coded as plesiomorphic for characters 9.19 and 9.22, and character 9.25 was excluded as redundant. Total “waqsa 37, character state transitions 61. A dot next to the number sl homoplas B 1985] plus the cordaite clade. The synapomorphy of the Ginkgo plus conifer plus cordaite clade is the presence of megasporophylls on an axillary fertile shoot (character 9.13). The morphological interpretation presented earlier (p. 749) of the Ginkgo ovulate reproductive structure is there- fore critical. There is no support in the analysis of characters presented here for Meyen's (1984) view that Ginkgo and related plants constitute a very distinct major lineage in seed plant evolu- tion. Ginkgo is resolved as more closely related to conifers and cordaites than to any of the taxa included by Meyen in his Ginkgoopsida. Simi- larly, although Ginkgo shares some features such as zooidogamy and the long free nuclear phase of embryogenesis with the living cycads (Stewart, 1983: 319), in cladistic terms these are primitive characters that do not indicate close phylogenetic relationship (Chamberlain, 1935: 216). Rothwell (1981) discussed two possibilities for the phylogenetic relationships of Callistophyton: either it is more closely related to conifers and cordaites than it is to other pteridosperms, or the similarities with conifers and cordaites are be- cause of parallel evolution. He rejected both and suggested a position intermediate between pte- ridosperms and cordaites or conifers. Both clado- grams reflect the difficulties in resolving the pre- cise debe relationship of Callistophyton and hav lycł jm os CHER peltasperms, and. the glossopterid plus Caytonia plus corystosperm clade. These four groups are themselves united by flattened seeds (character 9.10) and saccate pollen (lost secondarily in peltasperms, character 9.9 One advanced character that may ultimately be important in assessing the relationships of Callistophyton is the presence of pollen with a distal aperture. In both cladograms it is meth- odologically more parsimonious to interpret the proximal germinal apertures in medullosans, Mesoxylon, and Lebachia as reversals from dis- tal germination primitive for all seed plants ex- cept Lyginopteris (a total of three reversals). However, from a botanical perspective such re- versals seem unlikely and most paleobotanists would probably view the prepollen of medullo- sans, Mesoxylon, and Lebachia as retention of a rimitive feature. If this view is adopted it ne- cessitates five (cladogram 2) or six (cladogram 1) independent origins of distal germination. Cha- loner (19702) recognized that the switch from proximal to distal germination had occurred sev- CRANE-SEED PLANT PHYLOGENETICS 773 eral times in seed plant evolution, but the extent of this parallelism is of considerable interest. It suggests that detailed studies of pollen ontogeny might detect different mechanisms of aperture formation, and it clearly indicates that pollen with a distal aperture is not a good synapomor- phy of seed plants as might be concluded from a cladistic uen restricted solely to extant gymnosper At the same level as Callistophyton in the cladogram the systematic relationships of pel- tasperms are unresolved; but the other group in- corporated at this polychotomy is the glossop- terid plus Caytonia plus corystosperm clade de- fined by the presence of a *'cupule" (character 9.19). Within this group Caytonia and corysto- sperms are more closely related to each other than either is to glossopterids based on the poorly developed acid-resistant megaspore membrane (character 9.20). These relationships among “Mesozoic seed ferns” are maintained in both cladograms. The relationships of Pentoxylon previously have been considered enigmatic. Frequently the genus has been viewed as a peculiar and isolated gymnosperm of uncertain phylogenetic relation- ship (Andrews, 1961; Stewart, 1983) though var- ious similarities with other groups of seed plants have been pointed out. For example, monosul- cate pollen occurs in cycads, Ginkgo, and.the Bennettitales, while pycnoxylic conifer-like wood is a generalized feature within seed plants as a whole. Other authors (e.g., Ehrendorfer, 1971) have suggested a close relationship between Pentoxylon and the Bennettitales. Tn the analysis given here t useful in establishing the ual We relation- ships of Pentoxylon: the “‘flower-like” arrange- ment of the sban (character 9.23), which also occurs in Bennettitales, Gnetales, and angiosperms; the aggregation of the ovules into a head (character 9.26), which otherwise occurs only in the Bennettitales, and the structural sim- ilarity of bitegmic bennettitalean and Pentoxylon ovules (characters 9.19, 9.25, 9.26). If the inter- seminal scales of Bennettitales are accepted as being derived ontogenetically and phylogenet- ically from “cupule” primordia (characters 6.1, 9.19) then the existence of fully fertile ovulate heads like those of Pentoxylon, which lack in- terseminal scales, is a straightforward and rea- sonable prediction. This general view (though before Pentoxylon had been studied) was ex- pressed by Wieland (1906: 120). The stomata of 774 AT: ° L 1). D ] L pn) W enioxyrion) Bl y de scribed by Sahni (1948: 56) as syndetocheilic, and they deserve careful reexamination to clarify the range of variation "nich mey exon and to establish their preci fferences with those of the Bennettitales. It is also inter- esting that the anticlinal flanges of Nipaniophyl- lum leaf cuticles are finely sinuous, like those of most Bennettitales. A close relationship between Bennettitales and cycads is not supported by the cladogram, al- though the two groups frequently have been treated together on the basis of their leaf mor- phology and the similar habit of cycads and Cy- cadeoidea Sip ide 1916). Both clado- grams milarities are the result of convergence. Many Aras (Arber & Parkin, 1907: 51; Arnold, 1953: 58; Stewart, 1983) have recognized that the cycad plus Bennettitales group is artificial and probably does not reflect close phylogenetic relationship. The two groups are treated together primarily for reasons of con- venience and tradition, but according to the anal- ysis in this paper even the general designation of the Bennettitales as “‘cycadophytes” obscures their probable phylogenetic relationships with other taxa. On the basis of the characters given here, the two groups are very far apart In both cladograms the Bennettitales plus Pentoxylon clade is resolved as a sister group to a clade comprising the angiosperms plus Gnetales exactly as proposed by Arber and Par- kin (1908). The probability of a close relation- ship between the Bennettitales and Gnetales fre- quently has been suggested (Arber & Parkin, 1907, 1908; Martens, 1971: 269; Takhtajan, 1969: 17; Thoday, 1911), but at this level of analysis the cladogram does not support the view that the gnetalean “flower” is a highly reduced bisexual bennettitalean “flower” consisting of a single ovule or that the Gnetales and Bennettitales are sister taxa. It is nevertheless possible that a more detailed character analysis might place the Gne- tales as a relatively advanced monophyletic group within the bennettitalean clade. If this could be supported then a bennettitalean origin for the gnetalean “flower” would be more plausible. In this case the bennettitalean '*perianth" (character 9.33) may be interpreted as homologous to the opposite bracteoles of the Gnetales, and the "slight protruberance slightly below the epicot- yl" in Cycadeoidea embryos (Crepet, 1974: 158) could be compared to a poorly developed **feed- er" in Welwitschia or Gnetum (character 9.30). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Other similarities between the Bennettitales and Gnetales include the details of the micropyle and seeds (Berridge, 1911; Thoday, 1911), paracytic stomata, bisporangiate flowers, the possible pres- ence of vessels (Krassilov, 1984), and the pres- ence of a long micropylar tube. Cladogram 2 also supports the view that the apparent absence of a true outer integument in Gnetales is most easily explained as a secondary loss. The main alternative to a bennettitalean or angiosperm relationship of the Gnetales is the possibility of a close relationship between Ephedra and cordaites (Eames, 1952). Eames emphasized the difference between Ephedra ver- sus Gnetum and Welwitschia. The cladogram supports the view that Gnetum and Welwitschia are more closely related to each other than either is to Ephedra based on the reduced male ga- metophyte (character 9.27), the tetrasporic fe- male gametophyte (character 9.28), the lack of archegonia (character 9.29), and the presence of a "feeder" in the embryo (character 9.30; see also p. 766). However, contrary to Eames (1952) the cladogram suggests that the Gnetales are mono- phyletic and united by fundamentally similar re- productive structures (character 9.33), vessels (character 9.32), and possibly by their distinc- tive, ribbed pollen (character 9.31). Eames based his discussion mainly on the fertile axillary shoot that occurs in both Gnetales and cordaites, and his views hinge in particular on an interpretation of the ovules in Ephedra as appendicular rather than cauline. The other Ephedra-cordaite simi- larities cited by Eames (1952) are interpreted here as primitive features within seed plants (pyc- noxylic wood, anomocytic stomata) or indepen- dently derived in the spermatophyte clade (non- megaphyllous leaves). In most features already discussed cladograms l and 2 scarcely differ, but the most prominent and almost the sole effect of interpreting the other integument of Bennettitales and Pentoxylon as homologous with the “‘cupule” in glossopterids, Caytonia, corystosperms, and angiosperms is to i gana the Bennettiin pis PERONON plus group to the corystosperms. Under this inter are Mon the orthotropous bitegmic ovules of Bennetti- tales and Pentoxylon may be considered the ul- timate product of a reduction series involving the Caytonia megasporophyll with several **cu- pules" and several seeds in each; and the cor- ystosperm megasporophyll with fewer **cupules" each containing a single seed. The bennettitalean Ww megasporophylls con taining a i. vule, sterile “unicupulate,” seminal scales. or Pentoxylon ovule would be interpreted as a highly reduced megasporophyll of one “cupule” containing a single seed (Fig. 21, see also Thom- as, 1955: 655). In my view the only major ar- gument against this interpretation is the ortho- tropous orientation of bennettitalean and Pentoxylon “cupules” versus the recurved pules” of Caytonia and corystosperms. Clearly it would be of considerable interest if the pre- dicted (but unknown) recurved bitegmic ovules were detected in primitive Bennettitales. In favor of the relationship suggested in clado- am 2 is the similarity of ovule structure and cuticles in Caytonia, corystosperms, Bennetti- tales, and Pentoxylon. Additional evidence is provided by the similarity between Rhexoxylon (probable corystosperm stem) and Pentoxylon. Both have a “polystelic” arrangement of vascular tissue and also show an identical pattern of leaf- trace origin (Stewart, 1983: 312). On the basis of this slim evidence I currently favor cladogram 2 over cladogram l a most likely Hs primi. thet eed plants, and this view is also more PAS consistent with the stratigraphic evidence. Cladogram 1 places the origin of four fundamentally Mesozoic groups (Bennettitales, Pentoxylon, Gnetales, and CRANE-SEED PLANT PHYLOGENETICS ophyll containing one ovule.— aggregated into a head as in Pentoxylo aie Ser cael as in Bennettitales. Sterile megasporophylls forming inter- —F. An ngio sperm carpel opened and viewed adaxially. See text for further explanation. angiosperms) at a level equivalent to the origin of major Paleozoic groups such as medullosans and cordaites. Further work is clearly necessary to supplement our knowledge of characters and plants relevant to the competing hypotheses of (Stopes, Kráusel, 1949), Pentoxylon, and corystosperms is a pri- mary requirement. In addition to the particular questions of re- lationship already considered, t raise two general issues: are the pt phylogenetically useful group, and how well do existing classifications of seed plants reflect phy- logenetic relationships? The extreme heteroge- neity of the “pteridosperms” is clearly demon- strated in both cladograms. The relationships of medullosans, Lyginopteris, Callistophyton, pel- tasperms, glossopterids, Caytonia, and corysto- sperms are extremely diverse, and there is no pair of pteridosperm taxa that together constitute a monophyletic group in both cladograms. Gen- erally the concept of the pteridosperms has been used intuitively with little attempt at rigorous 776 1 . “ae e TABLE 10. I l gy Extinct taxa indicated with an asterisk. Chamberlain, 1935 Bierhorst, 1971 Cycadophytes Cycadopsida *Cycadofilicales *Pteridospermales *Bennettitales Cycadales Cycadales *Cycadeoidales Coniferophytes Caytoniales * Cordaitales Coniferopsida Ginkgoales *Cordaitales Coniferales Coniferales Gnetales axales Ginkgoales Gnetopsida Ephedrales netales Welwitschiales ngiospermopsida definition. However, Townrow (1962a: 316) of- fered a more precise circumscription defining the pteridosperms as *gymnospermous plants with leaves, pollen- and seed-bearing organs pinnate. t aggregated into cones or flowers.” This defines the pteridosperms en- urely in terms of pinnately organized leaves, mi- crosporophylls, and megasporophylls (character 9.11), which are all generalized features within most groups of seed plants. On this definition the pteridosperms are merely the residue of seed plants after taxa with “cones” (cycads, conifers, cordaites, Pentoxylon) and taxa with “flowers” (presumably Bennettitales, Gnetales, and angio- sperms) have been removed. The group is de- fined by exclusion and primitive characters. For phylogenetic purposes the current concept of the pteridosperms is valueless and has been a major source of confusion in attempts to analyze the phylogenetic relationships of seed plants. t , however, be recognized that classifications ins be viewed as having many urposes, and not all were intended to reflect Vui relationships. Considering only ex- tant gymnosperms the cladogram compares fa- vorably with the classification of Chamberlain (1935) (Table 10). Chamberlain divided the ex- tant gymnosperms into two groups, cycado- phytes, in which he placed the cycads, and conif- erophytes, in which he included the conifers, Ginkgo, and Gnetales. He clearly recognized the distinctiveness of the cycads, and that the co- ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 nifers, Ginkgo and Gnetales were more closely related. Bierhorst (1971) (Table 10) separated the Gnetopsida from the Coniferopsida, although he acknowledged that they are probably more closely related than either is to cycads. However, when the comparisons are extended to include fossil plants, the cladogram conflicts with the classifi- cations of Chamberlain (1935) and Bierhorst (1971). Both these authors recognized the Pte- ridospermales (Cycadofilicales) and the Cycade- oidales (Bennettitales) as two groups within the Cycadopsida (Cycadophytes). The heterogene- ity of the pteridosperms already has been dis- cussed, and according to cladogram 1, conifers, cordaites, and Ginkgo are more closely related to cycads than to Bennettitales. In cladogram 2 Gnetales. Bier- horst (1971) also included Caytonia in the Cy- cadopsida. In both cladograms Caytonia is more closely related to conifers, cordaites, and Ginkgo. The results presented here suggest that in ei for the cycadophytes to be considered a mono- phyletic group, it should oo include jera the medullosans and cyca If the cycadophytes of Casa baqana S system phytes" according to both cladograms expand to include Callistophyton, glossopterids, Caytonia, and corystosperms. In cladogram 2 the Bennet- titales, Pentoxylon, Gnetales, and angiosperms are also included in the coniferophyte clade. In this context many of the traditional distinctions between the cycadophytes and coniferophytes break down, and the *'coniferophytes" no longer include only forms with profuse branching, small simple leaves, small pith and cortex, and pyc- noxylic secondary xylem (Foster & Gifford, 1974: 85). If the relationships in cladogram 2 are ac- cepted, it may be more useful to recognize a broad group (*platysperms") defined by the presence of flattened ovules. Radiospermic ovules has been one of the characters traditionally used to link the cycads with the Bennettitales and even has been used to suggest a relationship between the Gnetales and medullosans (Rodin & Kapil, 1969: 429). Under the interpretation in cladogram 2, the Bennettitales and Gnetales are only second- arily radiospermic, and some, such as Varde- kloeftia and Ephedra, are platyspermic. There is, in any case, very little resemblance between the small, poorly vascularized ovules of the Bennet- titales and the large, doubly vascularized ovules of cycads. According to the analysis of characters 1985] in this paper, the radiospermy: of cycads, thed- ullosans, and L) the sec condary ‘ ‘pseudoradiospermy” of Bennet- titales and Gnetales DISCUSSION AND INTERPRETATION (ANGIOSPERMS) The angiosperms along with the Gnetales and the Bennettitales plus Pentoxylon clade form a monophyletic group. In cladogram 1 this clade cladogram 2 the corystosperm group to the Bennettitales plus Pentoxylon plus Gnetales plus angiosperm clade. is second pattern of relationships is broadly consistent with the view that the “Mesozoic seed ferns” were important in angiosperm origins (Andrews, 1961; Doyle, 1978; Gaussen, 1946; Retallack & Dilch- er, 1981; Stebbins, 1974; Stewart, 1983) and rec- onciles this idea with the suggestions of Arber and Parkin (1907, 1908) concerning the rela- tionships of angiosperms to the Gnetales and Bennettitales. The analysis of characters presented here does not address the possibility that the angiosperms are polyphyletic (Hughes, 1976, 1977; Krassilov, 1973, 1975; Meeuse, 1966) and accepts the tra- ditional view that the angiosperms are mono- phyletic (Cronquist, 1968; Takhtajan, 1969). In cladistic terms polyphyly could only be sup- ported if a group of flowering plants could be shown to be more closely related to some group of gymnosperms than to other flowering plants. This has never been explicitly demonstrated, but is implicit in Krassilov’s (1977b) suggestion that there are three major mid-Cretaceous angio- sperm “lineages,” the Hamamelidales, Laurales, and monocotyledons, that are “rooted” in the Dirhopalostachyaceae, Caytoniales, and Czeka- nowskiales, respectively. In both cladograms the angiosperms could be justified as monophyletic y five characters (9.34—9.38; see also Hill & Crane, 1982). It should, however, be recognized that most of these features are difficult to assess eyen in x living plants, miat most species in the y poorly understood, and that there are real probleme of sampling in such a large group. An interesting aspect of the cladogram is that the flowering plants are more closely related to Gnetales than any other group of gymnosperms. This view was advocated in the earlier part of CRANE-SEED PLANT PHYLOGENETICS 777 this century (Arber & Parkin, 1908) but has been out of favor for over 50 years, apparently for two principal reasons. The first is the apparent dif- ficulty of “transforming” the ovulate structures of the Gnetales into an angiosperm carpel. Al- though this has been attempted by several au- thors (Meeuse, 1966; Thompson, 1916), the re- sults have been unconvincing and usually have involved comparison with the ovary in some * Amentiferae" that are more likely to represent secondary modifications within the flowering plant clade. These hypotheses also have been unable to account for the origin of the condu- plicate carpel. he second reason stems from the widely ac- cepted view that vessels arose independently in the Gnetales and angiosperms (see earlier dis- cussion of characters 1.6, 9.32). This view has been so influential that, even when the many similarities between Gnetales ine Hi ig ia have been acknowledged, the as been that *the evidence of vascular anatomy pues controverts any suggestion of relationships" uae 1968: 43). Both of these problems seem to arise out of i. preoccupation with identifying angiosperm ancestors and the apparent impossibility that the Gnetales could have fulfilled this role. From a cladistic standpoint, such an ancestor-descen- dant relationship is untenable; some of the sim- ilarities between the two groups indicate a close phylogenetic relationship, but each has its own specializations with neither being ancestral to the other (see also Arber & Parkin, 1908; Just, 1948). Interestingly too, several of the gnetalean fea- tures that appear to be present in flowering plants (e.g., the dicotyledonous leaves of Gnetum) may well be secondary advances within the gnetalean a, ade. In both cladograms, the Gnetales and angio- sperms are resolved as sister groups (cf. Arber & (character 9.27), embryogenesis with no free nu- clear phase (character 9.2), precocious differen- tiation of integuments at the time of megaspo- rogenesis (Crepet, 1974: 163), presence of astrosclereids (Bierhorst, 1971: 475), and the oc- currence of paracytic stomata that, according to Baranova (1972), are primitive within angio- sperms. In addition, angiosperms and Gnetum are similar in leaf morphology and in having a 778 tal pal ti ially ( giosperms b female me hu at fertilization: Further similarities between the Gnetales and angiosperms, that require more detailed exami- nation, are similarities in chemistry (Gottlieb & Kubitzki, 1984) particularly of the wood (Gibbs, 1958; McLean & Evans, 1934; Melvin & Stew- art, 1969; Nishio, 1959), the occurrence of latic- ifers (Muhammad & Sattler, 1982), the small chromosomes, and the occurrence of polyploidy in Ephedra n 8 qi. 1976). Ehrendorfer es- timates the base chromosome number of Gne- tales as n = 7, the same value as that estimated for the base number of angiosperms (Raven, 1975). Arber and Parkin (1908: 513) explicitly rec- ognized a sister-group relationship between the angiosperms and Gnetales, with the Bennetti- tales as a sister group to both. The Bennettitales were an important factor in the development of the Anthostrobilus theory of the angiosperm flower (Arber & Parkin, 1907) and were part of the justification for considering the **Ranalian plexus" as the most primitive living flowerin plants. The cladogram provides an explicit base from which to examine further the idea that the Bennettitales, like Pentoxylon and Gnetales, are closely related to flowering plants and from which to pursue the functional parallelisms in terms of pollination and dispersal biology that can be traced in these clades. Other hypotheses for the origin of the angio- sperms are not supported by the distribution of characters in the cladogram. The cupules of ly- ginopterid seed ferns are not treated as homol- ogous with those of **Mesozoic seed ferns” (char- acter 9.19) and the cladogram lends no a pos- teriori support to this idea. Lyginopterid cupules therefore seem unlikely to be helpful in any par- simonious explanation ofthe origin of the second integument or carpel in angiosperms (Long, 1966, 1977). Indeed, results from a PAUP analysis of Table 9 in which the cupule of Lyginopteris is scored as homologous with the “cupules” of glos- sopterids, Caytonia, and corystosperms and the second integument of Bennettitales, Pentoxylon and angiosperms, place Lyginopteris in various highly derived positions inconsistent with other aspects of its morphology (and stratigraphic po- sition). The suggestion that the carpel evolved by inrolling ofa cycad-like megasporophyll (Ma- may, DU is ase URSUDportec by bee dein gram. Inrolling of a simple secret ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 would, in any case, only produce a carpel con- taining unitegmic ovules. Additionally, some of the plants (Spermopteris, Phasmatocycas) on which this hypothesis is based are interpreted here as platyspermic, and possibly more closely related to Callistophyton, the Mesozoic seed ferns, Bennettitales, Gnetales, and angiosperms, rather than to cycads (see below) Similarly, the notion that the angiosperms are cycadopsid seed plants (Doyle, 1978; Stewart, 1983) is not supported by the cladogram. This idea can be traced to the view that the seed ferns are uniformly a cycadopsid group. In fact, the jd apa historical development of the seed- concept, and their original designation as “ ir t has obscured their phyloge- netic heterogeneity. In my view the seed ferns include both cycadopsid and coniferopsid forms (p. 776), and it is the latter taxa that are more closely related to angiosperms. PHYLOGENETIC RELATIONSHIPS OF OTHER FossiL GYMNOSPERMS EREMOPTERIS, NYSTROEMIA, AND SIMILAR PLANTS Leaves, seeds, and probable microsporangia of Eremopteris were described by Delevoryas and Taylor (1969) from the Upper Pennsylvanian of Pennsylvania. The bipinnate leaves were closely associated with stalked flattened ovules. Each sporangia of Nystroemia pectiniformis from the Permian of China (Halle, 1929) are similar to those of Eremopteris. Although important de- tails of both Eremopteris and Nystroemia are unknown, the flattened ovules, combined with the lack of a *cupule" or conifer-cordaite-like inflorescence, suggests a combination of rela- tively advanced and primitive features similar to that seen in Callistophyton and peltasperms. Permian pinnately organized leaves bearing ovules on the lamina or at the margin have been described as *'callipteroid" (Mamay & W 1971). Tinsleya (Mamay, tenuis (Halle, 1929), a of interest that microsporophylls thought to be associated with these callipterid plants produced bisaccate pollen (see Mamay, 1966: 1) 1985] SPERMOPTERIS, PHASMATOCYCAS, SOBERHEIMIA Spermopteris coriacea (Goeppert) Cridland & Morris (1960) from the Upper Pennsylvanian of Kansas consists of entire-margined taeniopterid leaves bearing ovules on the abaxial surface. The ovules are confined to the distal portions of the leaf and form a row on either side of the midvein. They were oriented at right angles to the midvein with their micropyles arranged approximately along the margin. They were probably flattened with a distinct micropylar notch Phasmatocycas kansana Mamay (1976) is a megasporophyll with sessile ovules borne in two lateral rows. Distally the megasporophyll may have expanded into a lamina with taeniopterid venation (Mamay, 1976, but see Kerp, 1983). he ovules were flattened (Stewart, 1983: 283), had a shallowly bifid, notched apex similar to those of Spermopteris, a blunt, funnel-shaped micropyle, and a thick megaspore membrane. Recently collected material from Kansas (Pfef- erkorn, pers. comm.) also has flattened ovules. Soberheimia jonkeri Kerp (1983) from the Permian of Germany is similar to Spermopteris and Phasmatocycas. It is a leaf-like, bilaterally symmetrical structure with two rows of lateral organs interpreted as seeds. The lamina is thought to have been lobed with expansions between the seeds. Mamay (1976) interpreted Spermopteris and Phasmatocycas as important plants in the evo- lution of cycads. According to the analysis of characters given here, a close relationship to cy- cads is unlikely. Cycads and medullosans do not have platyspermic seeds and, as in Eremopteris and Nystroemia, the possession of this feature, combined with the lack of a conifer-cordaite-like inflorescence or “‘cupules,” is a combination of advanced and primitive features similar to that in Callistophyton and peltasperms. VOJNOVSKYALES The Vojnovskyales are a group of Permian plants described from Asia, North and South America, and Africa (Krassilov & Burago, 1981). Leaves of Vojnovskya (Nephrolepis) are fan- shaped and resemble those of the enigmatic Triassic plant Sanmiguelia (Ash, 1982; Becker, 1972; Tidwell et al., 1977) in the plications of the lamina and the lack of a clearly differentiated petiole. The reproductive structures are poorly understood but apparently consisted of “heads” CRANE-SEED PLANT PHYLOGENETICS 779 with several platyspermic ovules and numerous elongated scales. These scales have been inter- preted as microsporophylls but were probably sterile (Maheshwari & Meyen, 1975). Compar- ison of these few characters ofthe Vojnovskyales with the cladogram suggests that the combina- tion of platyspermic ovules (character 9.10) clus- tered together into heads (character 9.26) could indicate a possible relationship with Bennetti- tales and Pentoxylon. In view of this possibility, a detailed study of the stomata of Vojnovskya would be useful, and it would be of considerable interest to compare the scales on Vojnovskya “heads” with the interseminal scales of Bennet- titales (Harris in Mamay, 1978). PUTATIVE GNETALES The Gnetales currently have no well-estab- lished macrofossil record (Taylor, 1981a), al- though this may reflect difficulties of recognition rather than real absence (Arber & Parkin, 1908; Hill & Crane, 1982; Crane & Upchurch, work in progress). Upchurch and Crane (1985) describe a probable gnetalean with attached leaves and seeds from the Lower Cretaceous Potomac Group of Virginia, and pollen resembling that of extant Ephedra and Welwitschia is widely distributed during the Triassic and early Cretaceous. Recent transmission electron microscope studies of this *gnetalean" pollen have shown that the similar- ities also extend to the granular exine structure (Trevisan, 1980). Little is known of the plants that produced these grains. The Triassic dis- persed pollen Equisetosporites chinleana was referred to Ephedra by Scott (1960) and has niei. been demonstrated within micro- rangiate cones (Masculostrobus clathratus pde 1972b; see Zavada, 1984, for details of pol- len). The cones consisted of a main axis bearing spirally arranged mero with mp sporangia on the tal lamina. Bosea indica is a further 1 microspor- angiate structure from the Middle Triassic of Nidpur, India (Srivastava, 1973). It consisted of a stout axis bearing opposite or sub-opposite mi- crosporophylls with unilocular sporangia on the lower surface. The sporangia produced non-sac- cate, monosulcate pollen with pronounced lon- gitudinal ribs. Neither M. clathratus nor B. in- dica is similar to the microsporangiate structures of Ephedra, but both could be interpreted as sim- ilar to Pteruchus in which both dorsiventral and 780 helical arrangement of microsporophylls occurs (Pant & Basu, 1973). On the basis of cuticular structure Srivastava (1973) suggested that B. in- dica might be part of the plant that produced Lepidopteris indica leaves. These similarities with Gnetales and corystosperms require more de tailed examination but are of considerable in- terest in view of the relative positions of these two groups in the cladograms. A further possible macrofossil record of gne- talean-like plants is Hexagonocaulon minutum from the Upper to Middle Triassic of the South Shetland Islands. Hexagonocaulon minutum La- cey and Lucas (1981) was established for a large number of small in situ axes, each 0.5-4 mm in diameter. The axes bore alternating whorls of three scale-like bilobed appendages at the nodes. Nothing is known of the reproductive structures, oul y uley Witil Equisetites and Ephedra (Lacey & Lucas, 1981). Another fossil for which a gnetalean relation- ship has been suggested is the dispersed pollen Eucommiidites. Eucommiidites troedsonii was described first from the early Jurassic of southern Sweden (Erdtman, 1948) and interpreted as a tricolpate angiosperm grain with unequally de- veloped colpi. Based on analyses of the sym- metry, Couper (1956, 1958) suggested that a gymnospermous relationship was more likely, and Hughes (1961) interpreted the grain as dis- tally monosulcate with a proximal zonosulcus. Pollen wall structure of Eucommiidites is gran- ular (Doyle et al., 1975), but in some forms there e “‘pillar-like elements" in the pollen wall (Trevisan, 1980). The gymnospermous affinity of Eucommiidites was confirmed by its pollen wall structure (Doyle et al., 1973) and By D oc- currence in the Lower Cretaceous (Wealden) of southern En- gland (Hughes, 1961), the Lower Cretaceous Pa- tuxent Formation of Virginia (Brenner, 1967), and the Upper Liassic of Poland (Reymanówna, 1968). van MR Cittert (1971) has described con like structure from the Ju urassic of Yorkshire that produced Eu- commiidites pollen in numerous microsporangia on the adaxial surface of a microsporophyll. Knowledge of Eucommiidites is therefore too fragmentary to permit a detailed evaluation of relationships. However, the granular pollen wall (character 9.21) and the occurrence in ovules with an elongated micropyle suggests the possibility of a relationship with corystosperms, Gnetales, Bennettitales, or Pentoxylon. Similar views have ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 been expressed by Hughes (1961), Trevisan (1980), and Doyle et al. (1975). Against this sug- gestion is the occurrence of a well-developed megaspore membrane in the ovules (Reyma- nówna, 1968). Further information on the plants producing Eucommiidites is clearly required. LEPTOSTROBUS Leptostrobus is an ovulate reproductive struc- ture of Jurassic and Lower Cretaceous age. It consists of an axis bearing small scale leaves at the base and widely separated "capsules" con- taining ovules in the upper region (Harris, 195 1a; Harris et al., 1974; Krassilov, 1977b). The “‘cap- sules” are sessile and two-valved, with each valve containing a single row of three to eight small ovules. The ovules are poorly known, but were arranged with their micropyles facing the cone axis, and had a resistant megaspore membrane. Based on field association and cuticular similar- ity, Leptostrobus has been attributed to the same plant as Czekanowskia and Solenites. The leaves are borne on short shoots and may be either simple (Solenites) or finely and dichotomously branched (Czekanowskia). The microsporan- giate organ Jxostrobus has been linked with Czekanowskia at some localities (Krassilov, 1970), but in the Middle Jurassic flora of York- shire the evidence for linking Leptostrobus can- cer with Ixostrobus whitbiensis is weak, and Ixo- strobus may be attributed equally well to Desmiophyllum gramineus (Harris et al., 1974). Leptostrobus is not well understood, but some inferences concerning its systematic relation- ships are possible with reference to the clado- gram. The presence of scale leaves at the base of the **capsule"'-bearing axis suggests that it is mor- phologically an axial structure. The “capsules” therefore may be homologous to "cupulate" megasporophylls. The valves of the "capsule could be interpreted either as a single bivalved "cupule" or as a pair of closely associated “‘cu- pules" (character 9.19). However, Leptostrobus has a resistant megaspore membrane and has several seeds in each cupule (i.e., lacks characters 9.20, 9.22). Its position in the cladogram there- fore could be hypothesized as above the level of Callistophyton and peltasperms but below that of Gnetales, Bennettitales, and Pentoxylon. This hypothesis, however, may conflict with evidence from leaf morphology (character 9.11) and fur- ther characters are needed to pursue the issue. 1985] PROBLEMS AND POSSIBILITIES FOR FUTURE RESEARCH The analysis presented in this paper is open to criticisms of various kinds. However, criticisms of the lack of direct paleobotanical evidence of morphological transitions and the supposed in- validation ofthe results by i E homoplasy are straightforwardly reject The view that “direct” do evi- dence is the final arbiter in elucidating phylog- eny, and therefore in determining phylogenetic relationships, is entrenched in much botanical literature, but the kind of decisive paleontolog- ical evidence that might be required or expected is rarely elaborated. Finely graded series of in- termediates that can be “read directly from the rocks" are not a common feature of the paleon- tological record (Crane, 1984), and it is naive to expect the simple description of further fossils to miraculously reveal plant phylogeny. As in other areas of paleontology, paleobotanists will always be faced with a problem of determining phylogenetic relationships between superficially dissimilar organisms without an unequivocal se- ries of intermediates. The problem of determin- ing plant phylogeny is therefore to produce a classification that reflects phylogenetic relation- ships. To accomplish this effectively requires an explicit method of classification that has a straightforward evolutionary interpretation. The second common criticism, that homopla- sy (parallelism, convergence, and reversal) is so pervasive that the character patterns revealed by phylogenetic analysis are meaningless, is fre- quently linked to functional arguments that in- voke the influence of strong selective pressures. In plants there is certainly considerable homo- plasy (Fig. 22). Perhaps this is to be expected in dominantly modular organisms with highly vari- able genetic mechanisms and the potential for extreme phenotypic plasticity. However, either character Palle any attempt to clarify plant phylogeny i is aban- doned; or the problem becomes how may ho- moplasy be recognized particularly in cases where the normal criteria of structural, positional, and developmental similarity are fulfilled? Homol- other features indicate. The fact that homoplasy CRANE-SEED PLANT PHYLOGENETICS 781 can be assessed only in this way leads logically to a search for the simplest nested pattern for explaining the distribution of all features. ore serious, in my view, are problems that involve the analysis of characters on which the cladogram is based. A brief attempt has been made to justify the monophyly of Bennettitales, cycads, conifers, glossopterids, Gnetales, and an- giosperms in order to simplify the analysis, but the assumptions involved need to be tested as a more detailed understanding is developed of re- lationships within these groups. Similarly, gen- eralizations concerning groups of fossil plants such as the medullosans may be shown to be unjustified if future work demonstrates that they are heterogeneous fossil taxa (Beck, 1981: 193). Assumptions about unknown characters in cer- tain fossil plants are also problematic. These were kept to a minimum and are explicitly identified in the character tables. The basis for many is explained in the text, but others were merely scored as plesiomorphic to minimize their influ- ence on patterns of relationships. The necessity for these assumptions highlights the relatively poor understanding of many fossil taxa as whole plants. Current knowledge is highly uneven, and it is clear that there is as much need to under- stand our present fossils better as to discover new fossil taxa (Harris et al., 1974: 85). Recognition of these problems has highlighted specific questions that need to be addressed by future neontological and paleobotanical re- search. Those particularly important to resolving the relationships between ‘‘Mesozoic seed ferns,” Bennettitales, Gnetales, and angiosperms are: a) Clarification of integument ontogeny, anat- omy, and vasculature in peltasperms, Caytonia, corystosperms, and particularly in glossopterids. b) Clarification of the manner in which mi- crosporophylls and megasporophylls were borne in peltasperms, glossopterids, Caytonia, and co- rystosperms. Reexamination of the corysto- sperms would be particularly useful to reveal de- tails of megnaeroinyy) © nde ide and the Further study of silicified glossopterid material should also clarify the relationship between the megasporo- phyll and ‘“‘subtending bract.”’ c) Comparison of the outer seed covering (“‘cu- ule" of Vardekloeftia with the outer integu- mentary layer of anatomically preserved Ben- nettitales and Pentoxylon. d) Comparison of the anatomy and biology : af gnetalean, o r [Vor. 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN 782 SWH3dSOI9NV VUuQOaHda VIHOSLIIM13AM WNLAND NOTAXOLNAd S31IVLILLANNIG SWH3dSOL1SAHOO VINOLAVO Saiduaridosso19 SWH3dSVlla3d NOLAHdOLISITl1VO OS NIS SH33INOO LNVLX3 VIHOV831 NOTAXOSAW NOTAXIVQHYODS SQGVOASD SNVSOTINGIN Siu31dONI9A1 Siu31dO3VHOuV @ @ WW. ASP NO & (8) -6-6 WOOO e. REE "S. — L-eee QO ® © = © 6 © © | ACN fe n @ Fee @ © (X24 -OOOOH = @ (m ten equally Af; parsimonious cladograms constructed from version two of the data matrix. In this analysis the “cupules” of A 264 FiGuRE 22. Relationships between major groups of seed plants. St 1985] and detailed evaluation of the evidence for platy- spermy in these taxa, especially angiosperms. e) Comparative study of “seed cuticles,” in- cluding the megaspore membrane in living and fossil plants using modern techniques (Hill & Crane, 1982). f) Co TETE study of pollen wall stratifi- cation in “Mesozoic seed ferns,” Pentoxylon, and Bennettitales. g) Attempts to identify the plants producing dispersed Mesozoic “Ephedra-like” pollen Further difficulties with the analysis are the use of “reduction” or “loss” characters, e.g., lack of archegonia (character 9.29) and the relatively small number of characters available. It is a ten- uous position to regard two taxa as identical with respect to a condition that cannot actually be observed, but such characters may provide use- ful synapomorphies if they can be correlated with other features. Similarly, examination of additional features clearly is desirable but is constrained in part by lack of comparative information for seed plants as a whole, including living representatives. In many cases characters that have been surveyed in detail in one group of seed plants have been examined only cursorily, or not at all, in others. ost botanical comparative studies have con- centrated on taxa within seed plants and, as a result, have tended to reinforce their apparent differences. CONCLUSIONS got approach. outlined i in this paper does not differ fi used in paleobotany to interpret early land plant evolution (Banks, 1975; Chaloner, 1970b), early angiosperm evolution (Doyle, 1978; Doyle & Hickey, 1976), and studies, of Hie maor pi & ` 1975). It differs only in being relatively dee and in attempting to define a highly resolved hierarchy of relationships. In the context of seed plants it has attempted, for the first time, to pro- vide a detailed scheme of relationships account- ing for both neontological and paleontological data CRANE- SEED PLANT PHYLOGENETICS 783 The characters considered support a mono- phyletic origin of seed plants. Lyginopteris is placed as the sister taxon to all other seed plants. Flattened seeds and saccate pollen together de- fine a major group of seed plants (**platysperms"") including conifers, cordaites, Ginkgo, Callisto- phyton, peltasperms, glossopterids, Caytonia, and corystosperms. If the outer integument of ben- nettitalean, Pentoxylon, and angiosperm biteg- mic ovules is homologous to the “‘cupule” of glossopterids, Caytonia, and corystosperms then these groups plus the Gnetales form a clade of “higher” ‘‘platysperms” within the “‘platys- perm" group. Such an interpretation would rec- oncile the widely accepted view that “Mesozoic seed ferns" are highly relevant to the problem of angiosperm origins with the idea proposed at the beginning of this century that Bennettitales and Gnetales are closely related to flowering plants. If this is accepted the sister group to the “higher platysperms.” The Bennet- titales and Pentoxylon are resolved as sister taxa and together form the sister group of angio- sperms plus the Gnetales. The Gnetales are monophyletic and the sister group of flowering Ü Inte ) 9 1 3 phylogenetic connection between cycads and Bennettitales and emphasizes the heterogeneity of the seed ferns as currently recognized. seed ferns include taxa of very diverse relation- ships. The current concept of the p MUSAE a ' is therefore phylogenetically meaningless an confuses discussions of inis elati ships. The interpretation of seed plant relationships presented in this paper (particularly cladogram 2) is in broad agreement with the first strati- graphic appearance of the various taxa. It sug- gests that increased understanding of Triassic Bennettitales, Gnetales, and corystosperms wi major interest in further elucidating the phylogenetic relationships of flowering plants. It may also suggest a substantially earlier time of origin for the flowering plant clade than is cur- rently envisaged. A Triassic or Jurassic origin would not conflict with the hypotheses of angiosperm diversifica- 2 m— and Pentoxylon were coded as d to the “cupules” of glossopterids, Caytonia, corysto- perms, and angiosperms. Bennettitales and Pen xylon were coded as a pomorphic for characters 9.19, 9.22, and 9.25. Total characters 38, character state hii 2 a- 62. A dot next i the number indicates homoplasy. 784 tion proposed by Doyle and Hickey (1976) and would suggest a pattern of evolutionary history for flowering plants similar to that of mammals with an early Mesozoic origin followed by low diversity and a delayed radiation. There are still, owever, no unequivocal pre-Cretaceous angio- sperm records, although there is an abundance of enigmatic Mesozoic plants about which we know almost nothing. Among these problemat- ica that show angiosperm-like characters are the schmidt, 1968), and Phyllites (Seward, 1904). In palynology, the reticulate, columellate, mono- sulcate grains reported by Cornet from the Upper Triassic-Lower Jurassic Newark Supergroup of the eastern United States are very similar to those of angiosperms (Doyle, 1978). The principal difficulty with the phylogenetic analysis presented in this paper is the problem of dealing with missing characters in many po- tentially critical fossil plants. 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O. dá polos species and clado- grams —rema n the symposium. Pp. 211-244 am in J. Cracra LAN. Eldredge (editors), Phylogenetic c ah and Paleontology. Columbia Univ. Press, New 1 81. ` Phylogenetics: The Theory and Prac- tice of Phylogenetic Systematics. Wiley and Sons, New WODEHOUSE, R. P. Hill, New York. 1936. ers oy of pollen grains. Bot. Rev. T eee 2: 67- Wo ret, J. A., J. A. PES M. PAGE. 1975 [1976]. The bases of angiosperm phylogeny: paleobotany. n. Missouri Bot. Gard. 62: 801-824. E icm W. C. 1906. The structure and origin of the Cycadaceae. Ann. Bot. (London) 20: 129-1 59. Young, D. A. 1981. Are the angiosperms primitively vesselless? Syst. Bot. 6: 313-330. & . RICHARDSON. 1982. A ara = < analysis of extant seed plants: the need to utiliz homologous characters. Taxon 31: 250-254. 1935. Pollen Grains. McGraw- structure. Ann. Missouri Bot. Gard. 71: 444—463 BIOLOGICAL DIVERSIFICATION AND ITS CAUSES! JOEL CRACRAFT? ABSTRACT Three major patterns of diversity are defined and a general hypothesis proposed to explain them. munities or biotas vary spatially and temporally, and (o) Phanerozoic posila which describes large- rrent explanatory hypotheses for these patterns are generally formulated for systems that are pecan to be in a state of ecological equilibrium, with speciation and extinction rates being diversity-dependent. This paper describes an diversity. It is postulated that shayana ne is CORB ded primarily by large-scale aspis in lithospheric (geomorphological) complexity. T is hypoth cal data showing that allopatric speciation is the "ms. BER of differentiation, and of ea ie showing that tectonic changes within the lithosphere are responsible for the formation of pisei a and d barriers. This hypothesis makes a number of predictions about patterns of en m, orical biogeography, and spatial gradients of diversity, and data consistent with these predictions are pre- sented. Other p riabilit behavioral- ecological variability within species, intensity of sexual selection) : are discussed and their yp may occasionali ly nportant 4 + 1 Aeal ++ 4A. = š +L : m - 1 1 for ae patterns Or long-term changes of diversity within biotas. Extinction rate is postulated to be controlled primarily by spatial and temporal changes in envi- ronmental harshness, particularly as the latter is manifested by gradients of temperature and moisture. ian e of extinction, particularly the tectonic elimination of habitats, may be ofi impo ortance fo or specific groups of organisms (e.g., marine shelf at aici at aie and times (e.g., near a volcanic eruption) but are not likely to play : as ; important a role as does change in harshness. Together, the two main controls on speciation and iine Au define a diversity- independent process of diversification. The biosphere can be odynamic system t be expected to grow in complexity (including diversity) through time as a result of a el Misa The temporal history of the biosphere therefore has been nonequilibrial rather than equilibrial. Clades of organisms exhibit variation in di- versity through space and time, and closely re- lated clades often are characterized by significant differences in relative diversity. Species diversity in large- -scale com dances of species are apportioned nonrandomly within and among local habitats. These and other patterns of biotic div: ave been known for a long time and biologists have fervently sought their explanation. This paper addresses this question: Can a major component of the observed variance in these patterns be explained by a single causal hypothesis? The analysis of biological diversification has recently shifted its emphasis. For many years, particularly in the 1960s, the field was dominated by the study of large-scale and local patterns of ! [ am exceedingly grateful to many colleagues who have discussed the ideas of this paper with me or who [o . Gentry, Jurgen Haffer, W an uscript, including Daniel Axelrod, Teresa Bone, Daniel Brooks, G. Brent Warren Hamilton, James R. Karr, Peter en, Thomas J. M. SA and Larry Wolf. I am especially indebted to Jennifer Kitchell and John Wiens for their critical comments on a later version of the paper even thou by the National Science Foundation. 2 Department of Anatomy, Univer Museum of Natural History, Chicago, Illinois 6060 ANN. Missouni Bor. GARD. 72: 794—822. 1985. manuscript. All of these colleagues have improved this gh they do a necessarily agree with its conclusions. Portions of this research were supported sity of Illinois, Chicago, Illinois 60680; and Department of Zoology, Field 5. 1985] species diversity, particularly from an ecological point of view. Since the early 1970s, the problem of diversification has been addressed increasing- ly from a taxonomic viewpoint, especially by paleontologists. This change has arisen partly as a result of a reformulation of macroevolutionary analysis, from one focusing on the adaptive transformation of organismal form to an ana- lytical assessment of changes in diversity within and among clades (Eldredge & Gould, 1972; Stanley, 1979; Eldredge & Cracraft, 1980; and many papers in Paleobiology). As a consequence, the literature is now dominated by the study of speciation and extinction rates using primarily fossil taxa. Significantly, while attempting to ac- count for patterns of diversification within clades and for temporal changes in the entire biosphere, this work has often neglected the spatial analysis of diversity. Many explanations have been proposed for patterns of diversification, yet there has been little recognition that macroevolutionary diver- sity and spatial diversity might constitute a single problem. Biologists realize that complex biolog- ical patterns such as those of diversity only rarely result from single causal processes. Yet, science is an endeavor that searches for generality, and thus scientific explanations, to the extent that they are in fact general, should attempt to ac- count for as much of the variance within a data- set as possible. In so doing, patterns not ex- plained by that hypothesis may be identified and perhaps explained by second order causal factors. Patterns of diversification are first-order func- tions of speciation and extinction. À theory of biological diversification, therefore, must begin by delineating the rate-controls of speciation and extinction. This paper advances the hypothesis that spatiotemporal changes in speciation and extinction rates describe patterns of diversifica- tion both within clades and across large-scale species assemblages. In particular, diversifica- tion is postulated to be diversity-independent, with speciation rate being a function of the rate- change in lithospheric (geomorphological) com- plexity and extinction rate being a function of the rate-change in environmental harshness. This hypothesis makes the claim of identifying the major causal factors shaping these diversity pat- terns. Other factors certainly will be of impor- tance in particular instances, and some of these will also be discussed below. Intra- and inter- cladal changes in species abundances and large- scale spatial gradients in diversity constitute the CRACRAFT — BIOLOGICAL DIVERSIFICATION 795 domain of the hypothesis, and it makes no at- tempt to account for local, among- or within- habitat diversity. Contemporary diversification theory is based largely on the assumption that the controls of speciation and extinction are di- n el elsewhere (Cracraft, in prep.) and will not be con- sidered in detail here. Bioric DIVERSIFICATION THE PATTERNS Patterns of diversity (species richness) can be expressed in terms of the diversifying lineages (clades) themselves, in which case these can be called macroevolutionary patterns (Eldredge & Cracraft, 1980; Cracraft, 1982a), or they can be characterized spatially as a statistical summation of all the cladal patterns present at one or more locations. These latter patterns of diversity will be termed global when the scale is large, and local (or ecological) when it is small. Brief mention should be made of several as- sumptions underlying the recognition of diver- sity patterns (discussed in detail in Cracraft, 1981a, 1982a). In defining diversity, we seek to count all those taxonomic entities thought to be evo- lutionary units. Thus, concepts of species have important theoretical and empirical conse- quences. To the extent that prevailing species concepts vary from group to group, measures of diversity will not necessarily be comparable. Within much of vertebrate zoology (especially in ornithology and mammalogy), the biological species concept is widely applied (e.g., Mayr, 1963, 1969). That this species concept cannot be used to describe diversity accurately is not gen- erally understood. Many differentiated, evolu- tionary taxonomic units, for example, can be in- cluded within a single “biological species," and this often occurs in practice (Rosen, 1978; Cra- craft, 1983a). Use of the biological species con- cept significantly underestimates the numbers of taxa that have diversified. It is more appropriate, therefore, to adopt a species concept that is de- fined in uns of the evolutionary units them- selves uses both phylogenetically within clades and ecologically within and among biotas (for one such concept, see Nelson & Platnick, 1981, and Cracraft, 1983a). Patterns of diversity can be described accu- rately only to the extent that highly corroborated relative 796 radiation reduction reduction radiation radiation A B ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 radiation mass extinction radiation C D FIGURE 1. Four common patterns of diversity observed within lineages. Patterns A-C involve combinations of i and reduction diversity; pattern D illustrates steady-state diversity (after Cracraft, 1982a). phylogenetic relationships are available. This is especially true with respect to macroevolution- ary patterns, which are defined in terms of those E, uibus hypotheses of relation- ships change patterns of diversity signifi- cantly Gan 1981a, 1982a; Novacek & No- rell, 2 nowledge about phylogenetic relationships may be less critical when diversity of biotas is being measured, unless that measure is being partitioned taxonomically or examined historically. Three major ee E INED) patterns will be addressed in this l. MEC ene diversity. This in- cludes patterns of species richness within and among clades. Intra-cladal patterns are expressed in terms of radiation diversity, in which species numbers increase with time; reduction diversity, in which numbers decrease with time; and steady- state, in which diversity is maintained relatively constant throughout time. These patterns may occur in combination within any one clade (Fig. A second macroevolutionary p describes sister-clades that vary greatly in relative diversity (Table 1). Because they are hypothesized to be sister-groups, the diversity of the two clades can be compared in terms of the same age of origin. 2. Global diversity. Large-scale gradients of species richness among communities constitute a well-known pattern, particularly as it is ex- pressed in terms of latitude, but longitudinal gra- dients also exist. Many such patterns have been described in the literature (e.g., Fischer, 1960; Stehli, 1968), and some will be discussed below. 3. Phanerozoic diversity. Temporal varia- tion in species richness within the biosphere has been extensively studied, particularly for marine organisms (Raup, 1972, 1976a, 1976b; Sepkoski, 1976, 1978, 1979). EQUILIBRIUM OR NONEQUILIBRIUM? Current models of diversification generally have borrowed their conceptual framework from population ecology. Speciation rate is analogized with birth rate, extinction rate with death rate. approach, even in those studies examining pat- terns over vast stretches of time, appears to have been the widespread acceptance of island bio- geographic theory (MacArthur & Wilson, 1963, 1967). The use of diversity-dependent, equilib- rial models is pervasive, and they have been ap- plied whether the system under discussion is di- versity within the entire biosphere through the Phanerozoic (Stehli et al., 1969; Sepkoski, 1976, 1985] n and h ict taxa CRACRAFT — BIOLOGICAL DIVERSIFICATION 797 TABLE 1. to tachytelic and bradytelic rates of speciation. y probably related Tachytelic Taxon Bradytelic Taxon Aves? pens rmes er pelecaniforms (59) Phaethontidae (3) Pea sisi Ardeiidae (62) Balaenicipitidae (1) eriform Anatidae (147) Anhimidae (3) Falconiformes Accipitridae + Falconidae (279) Pandionidae (1) Galliformes usasapa + Phasianidae (256) Megapodiidae (12) Gruiform Rallidae 3 Heliornithidae (3) Columbiformes Columbidae (306) Pteroclididae (16) Cuculiformes Cuculidae (129) Musophagidae (18) St orme Strigidae (135) Tytonidae (11) Piciformes Picidae ban, Indicatoridae (16) Passerifor other DIE (1,097) Acanthisittidae (4) Furnariidae (218) Dendrocolaptidae (52) Laniidae (74) Vangidae (13 Corvidae (106) Grallinidae (4) Estrildidae (136) Bubalornithidae (2) Estrildini (113) Viduini (10) Mammalia’ Chiroptera (981) Dermoptera (2) Rodentia (1,728) Rodentia + Lagomorpha (1,794) gomorpha (66) Macroscelidea (21) a For a discussion of these interrelationships see Cracraft (1981b). * For a discussion of these interrelationships see Novacek (1982). 1978, 1979, 1984; Raup, 1976a, 1976b; Raup et al., 1973; Simberloff, 1974; Webb, 1976; Gould, 1976; Stanley, 1979; Valentine, 1980; Carr & Kitchell, 1980) or diversity within Recent com- munities (Hutchinson, 1959; MacArthur, 1965, 1972; MacArthur & Wilson, 1963, 1967; Wilson, 1969; Terborgh, 1973; Saunders & Bazin, 1975; Rosenzweig, 1975; Levinton, 1979; Fowler & MacMahon, 1982). The use of diversity-dependent, equilibrial models has influenced our understanding of di- versification in two ways: 1. By an expectation of pattern. Obser- vations are theory-laden and our expectation of patterns in nature is dependent on the predic- tions of our theoretical worldview. By building an equilibrial viewpoint into diversification models, however, expected and observed pat- terns are more likely than not to confirm (and conform to) that viewpoint. Thus, many pale- ontologists have interpreted the Phanerozoic di- versity curve to be equilibrial (e.g., Sepkoski, 1978, 1979; see Discussion), with mass extinc- tions perturbing diversity away from the equi- librium, but only for relatively short periods of time. Likewise, the expectation of pattern is also prevalent when diversity-dependent models are applied to patterns of species diversity at the local level. Some of these latter observational expectations include: saturation of habitats, competitively structured community assem- blages, predominance of density-dependent mortality, and the presence of limiting similar- ities among co-existing species, to name only a few (see Wiens, 1984, for further examples and discussion). 798 h th : En dd í . u of diversity-dependent models is that there is an upper limit to diversity (MacArthur, 1969; Mar- galef, 1972; Van Valen, 1976; Levinton, 1979). The upper-limit hypothesis has its strongest rationale within current ecological theory when attempts are made to explain within-habitat di- versity, but it also has been extrapolated to the biosphere as a whole. I will return to this hy- pothesis in the Discussion. 2. By an expectation of process. Underlying assumptions also carry with them explicit ex- pectations "us boni causal structure of nature. ted, e of an equilibrial worldview has resulted in | the dud use of diversity- dependent models of speciation and extinction rate-control. Metaphorically speaking, the bio- sphere is viewed as a large Tribolium population bottle, in which space and resources are assumed to be limited; then, using the same logistic growth equations of population biology, paleontologists have extended the analogy to expect a decrease in speciation rate and rise in extinction rate with increasing diversity (MacArthur, 1969; Valen- tine, 1973; Rosenzweig, 1975; Sepkoski, 1978; Carr & Kitchell, 1980). Diversity-dependent " of these rates is postulated to result directly from intensified ecological interactions as diversity increases, in particular increased competition and species packing, along with re- duced dispersal ability (see above cited authors for a more detailed explanation and Cracraft, in prep. for a critical evaluation of this model). The hypothesis presented here states that the patterns and processes of biological diversifica- tion are more properly characterized within diversity-independent, nonequilibrial frame- work. There are several reasons for applying this conceptual approach to questions within evo- lutionary biology. One is that historical systems, whether physical or biological, behave as if they are nonequilibrial rather than equilibrial, and therefore it seems appropriate that their descrip- tion be cast i (M , 1981; Peacocke, 1983). Biological systems, including individual organisms, species, communities, and the biosphere, appear to exhibit a transforma- tional history of increasingly more organized steady-states; with rare exceptions, they increase in complexity tbrough time. Thus, an increasing number of biologists ideas to questions about the origin of macro- molecular complexity (Prigogine et al., 1972; Saunders & Ho, 1976, 1981; Wicken, 1978, 1979, e ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 1980; Eigen, 1971; Eigen & Schuster, 1977, 1978a, 1978b), general evolutionary theory (Wi- ley & Brooks, 1982; Brooks & Wiley, 1984), de- velopmental biology (Zotin, 1972; Lurie & Wag- ensberg, 1979), and community structure and evolution (Gallaucci, 1973; Ulanowicz, 1980; Phipps, 1981; Johnson, 1981). A second reason for considering a nonequilib- equilibrial systems are inherently transforma- tional and nongradualistic when seen macro- scopically (Mercer, 1981; Wiley & Brooks, 1982). In contrast, equilibrial systems, by their nature, are not persistently transformational, and within this conceptual framework biologists frequently h t $24 ala P +l lal the “forces” maintaining the system at or near equilibrium, or returning that system to equilib- rium following a perturbation by some outside "force." A nonequilibrial viewpoint places the description of biological systems behavior in a new li THE DIVERSITY-INDEPENDENT RATE-CONTROL OF SPECIATION The primary aim of this section is to describe a hypothesis of speciation rate-control that is di- versity-independent. Other factors have also been hypothesized to modulate the rate of speciation, and these will be discussed separately. THE HYPOTHESIS OF LITHOSPHERIC COMPLEXITY That speciation rate is controlled primarily by nonequilibrial changes in large-scale lithospheric geomorphological) complexity can be deduced from biological and geological premises. The hy- pothesis itself can then be used to generate test- able predictions about patterns of biological di- versification. Consider two biological and two geological premises: Premise 1. Speciation is a geographical phe- nomenon and is allopatric in most groups of or- ganisms (Mayr, 1963; Grant, 1971; Bush, 1975; White, 1978). Speciation via genetic alterations such as poly- ploidy is frequent in some groups, particularly angiosperms (Grant, 1971; White, 1978), but there is little evidence to suggest that allopatric within tephritid flies of the genus Rhagoletis, 1985] which has been studied extensively because it is said to provide a possible example of sympatric speciation, most of the species are allopatric and almost certainly exhibited geographic disjunc- tion during their diversification [Bush (1966); see also the arguments of Futuyma & Mayer (1980) against the commonness of non-allopatric dif- ferentiation]. Premise 2. Allopatric speciation is depen- dent upon the origin and maintenance of geo- graphic or ecological barriers. This assumption is straight-forward and em- pirically well-corroborated (Mayr, 1963; Bush, 1975; White, 1978). Geographic barriers are im- portant in two ways, by promoting vicariance and isolation of widespread populations and by providing the possibility of isolation given long- distance dispersal across those bayeiers. ne though not dependent up the relative frequencies. of vicariance and long- distance dispersal, the hypothesis can be used to predict spatial characteristics of differentiation (see below). Premise 3. The lithosphere is an irreversible thermodynamic system exhibiting a nonequilib- rial increase in complexity. Decay of radioactive isotopes of uranium, thorium, and potassium produces heat that ul- timately drives mantle convection currents and lithospheric plate motion (Davies & Runcorn, 1980; Bott, 1982). Being a historical system, the lithosphere is irreversibly transformational. Geological degradation of complexity may occur locally (for example, from erosional processes), but the overall historical trend has been for com- plexity to increase globally, as evidenced by our ability to reconstruct past lithospheric (geomor- phological) configurations. Premise ^ The Tate of formation of geo- function of rate- changes in lithospheric (geomorphological) evo- lution Relative plate motions create large-scale geo- morphological complexity in some areas and contribute to processes of geological degradation in others. Regions of high complexity (e.g., areas of high relief) are generally associated with high- ly active, converging plate margins or with active deformation within plates. Changes in the litho- sphere also have first-order effects on climatic change (Hamilton, 1968; Cox, 1968; Crowell & Frakes, 1970; Frakes & Kemp, 1972, 1973) and thus on the rate of climatic barrier formation. These premises lead to the hypothesis that the graphi ca CRACRAFT — BIOLOGICAL DIVERSIFICATION 799 rate of speciation is a first-order function of litho- spheric (geomorphological) complexity. A causal connection between speciation and large-scale plate motions has been suggested before (see es- pecially Valentine & Moores, 1970, 1972; Val- entine, 1971, 1973; Bakker, 1977). Simpson (1964) and Nel (1975) also noted a correlation between geologic (topographic) complexity and diversity, thereby inferring a relationship to spe- ciation. Bakker (1977) postulated cycles of orog- eny and marine transgressions producing cycles of speciation in the higalands of continents, with subsequenti 1) of species into lowland habitats. Ross (1972), in a highly stim- ulating and underappreciated paper, viewed di- versification as a rate-function of geographic dis- junction within a nonequilibrium biosphere; his short contribution is one of the more explicit statements to date regarding the rate-control of speciation by earth history. he hypothesis of lithospheric complexity de- scribed here pois significantly from this pre- vious work. It sta plicitly that the rate-change of geological oe is the primary control of speciation rate. The hypothesis also makes a larger claim: because the lithosphere exhibits a nonequilibrial increase in complexity through time, Phanerozoic patterns of diversification should also be nonequilibrial. Furthermore, the hypothesis is more general: although it acknowl- edges the correlation between high topographic relief and high diversity, it posits a causal rela- tionship between lithospheric complexity and speciation rate across both terrestrial and marine environments. Thus, topographic relief per se is not the central causal agent of speciation control; rather, it is the rate-change in geomorphological complexity that is causally related to the rate at which barriers arise and taxa therefore speciate. Areas of high relief may be geologically stable and produce few species, whereas areas of lower relief may be tectonically active and produce species at a higher rate. e now turn to some predictions and impli- cations of the hypothesis of lithospheric com- plexity gini examine gbservational data relevant to them. Critical p exist with any attempt to decipher dc causal mechanics of diversifica- tion. If we are examining the expected influence of a particular causal hypothesis (as with litho- spheric complexity, for example), not only are there the confounding contributions of other fac- tors controlling the rate of speciation, but also 800 the concomitant effects of extinction. Except un- der very special circumstances, we cannot expect to measure speciation rates directly and accu- rately. Consequently, diversity patterns them- selves constitute the data with which we must test alternative causal processes. However, di- versity patterns are the end result of the spatio- temporal history of all the separate causes of spe- ciation and extinction. Therefore, predictions derived from any one hypothesis of speciation or extinction rate-control must of necessity be proposed within the context of ceteris paribus. This requirement naturally raises the specter of ad hoc escapes from potentially conflicting ob- servational data: if the data do not fit the hy- pothesis being tested, are we to claim that the hypothesis itself is not rejected because devia- tions from expectation can be ascribed to other causal factors? This onerous problem can rarely be avoided when many causal factors are in- volved; one can only attempt to discuss its ram- ifications as openly as possible (e.g., see Hillborn & Stearns, 1982). This situation does not mean that causal hypotheses about speciation and ex- tinction rate-controls are untestable; as long as the relative influence of other potential causal factors can be minimized, then the importance of a specific causal mechanism can be investi- ated A series of predictions can be derived from the hypothesis that geomorphological change con- trols the rate of speciation, including: Prediction 1. Species will be distributed non- randomly and will be clustered in areas of ende- ism. If geomorphological change mediates the spa- tial pattern and tempo of speciation, then geo- logically induced barriers (physiographic or eco- logical-climatic) will partition biotas through time, and thus produce common areas of ende- mism. This is a strong prediction, but the ex- pected patterns can be obscured (1) by popula- tion dispersion in which case species ranges will exhibit varying degrees of congruence, or (2) by long-distance dispersal across barriers. If taxo- nomic differentiation took place only by long- distance dispersal and not by the vicariance of species that were already widespread, then the frequency of well-defined areas of endemism would be low. Long-distance dispersal is a sto- chastic property of individuals, and thus major centers of endemism are not likely to arise by this process. In summary, if areas of endemism did not exist, or were very uncommon, then the ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 hypothesis of lithospheric complexity would not corroborated. Observational data strongly support this pre- diction. It has been known for several centuries that organisms are nonrandomly distributed and cluster in areas of endemism. The literature per- taining to areas of endemism (also termed “areas of provinciality," or “centers of dispersal”) is vast. For example, well-marked areas of ende- mism are apparent in the avifaunas of Australia (Keast, 1961); Africa (Hall & Moreau, 1970; Snow, 1978), and South America (Muller, 1973; Haffer, 1978; Cracraft, 1985). Likewise, Gent (1982a) and Prance (1982a, 1982b) have pre- sented data documenting the existence of areas of endemism within the South American flora. Many more examples could be cited. Prediction 2. Species will exhibit congruent patterns of historical biogeography. There is no stronger test of the hypothesis of lithospheric complexity than the relative fre- 1979; Platnick P: Nelson, 1978; Nelson & Platnick, 1981; Cracraft, 1982b, 1983b). The vicariance of biotas is the only known mechanism producing congruent histories in- volving three or more areas of endemism; the other mechanism producing allopatric disjunc- tion, long-distance dispersal, can yield two-area congruence (as from a mainland to an island), but invoking it to explain patterns involving three o tat + 41 A more areas ne hoc assumptions about dispersal, the relative timing of differentiation, and extinction (see above citations). Thus, the degree to which phylogenetic rela- tionships of species confirm hes existence of con- gruent | of the rate- control of : speciation by geologic (in- cluding climatic) fragmentation. Unfortunately, the field of vicariance biogeography is still in its infancy, and few studies bearing on this predic- tral American freshwater fishes; Cracraft (1982b, 1983a, 1983b) does likewise for some Australian birds; Wiley and Mayden (1985) also present strong evidence that vicariance was extremely important in the diversification of the eastern North American fish fauna. Finally, the Ama- zonian forest avifauna also presents examples of PRUM (1982) 1985] CRACRAFT- BIOLOGICAL DIVERSIFICATION 801 š : 2 i | | `—— > By. = T3 I | eee 77" ` PTEROGLOSSUS | hand p oum t| PIONOPSITTA | : A ka e haematotis S - v as (pyrilia L" U” — Prey EN coccineicollaris p) ES N Í | | A/S | pulchra / / ` FIGURE 2. Four lineages of tropical South American birds exhibiting congruence for four areas of endemis "m iet. Other taxa in Central pom erica, n within these clades indicate the following pattern of historical bio- a, and southeast B components in these comparisons. Data after Prum (1982) and Tanh au observ taxa whose biogeographic patterns are congruent with one another (Prum, 1982; Cracraft, unpubl. observ.; see Fig. 2). Congruent t are compelling support for the hypothesis of lithospheric complexity. That congruence im- plies that the biotas themselves have been frag- mented by common geographic barriers, and if so, we can predict that other taxa endemic in these same areas eventually will be shown to have histories congruent with those already stud- ied. If geological change has not set the tempo hasthese of diversification, or has played only a minor role, then oe congruence should be rare or nonexist Prediction 3. "ien graphic gradients in species diversity will parallel gradients in lithospheric complexity. All things being equal, species diversity should be higher in regions of higher complexity. But things are rarely, if ever, equal. Regions of lower complexity might have higher diversity if ex- tinction has been significant in those regions of higher complexity. Thus, in attempting to ex- 802 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 P200 ul 3000 + 150 o O < D m = o F 2000| +100 = < < 1000 4 + 50 120 115 110 105 100 95 90 85 80 75 LONGITUDE FIGURE3. A soo } th n Un ited States at 35°N latitude showing the relationship tween top l d species diversity gradients of birds and mammals (data ie MacArt thur, 1969: Simpson, arn Bui in both groups rises in those areas having the greatest topographic complexity (see text). amine this prediction, it is important to mini- mize the potential effects of variation in extinc- tion along the geographic transect being investigated. As will be discussed below, this is probably one of the reasons why latitudinal tran- sects of diversity do not show close correlation with complexity. Longitudinal transects, on the other hand, can be chosen so as to reveal the relationship between diversity and complexity while at the same time minimizing the influence of extinction in obscuring that pattern Consider first data from a continental setting. Figure 3 shows a longitudinal transect across the southern United States at 35?N. In this figure, relative topographic relief is taken to be an ap- proximation of lithospheric (geomorphological) complexity. Data on species richness for mam- mals (Simpson, 1964; Wilson, 1974) and land birds (MacArthur, 1969; see also Cook, 1969) are also displayed. The gradient in diversity of both birds and mammals parallels the "pesi in topographic complexity, as predicted by t 64: 70) ciation in the West (see also Cook, 1969).] Comparable data for similarly chosen gradi- ents are rare indeed, in part because the taxa being analyzed must not be sensitive to any marked change in environmental harshness along the gradient. Thus, it is not surprising that am- phibians exhibit a strong westward decrease in diversity along the same transect (Kiester, 1971), and the major change in diversity corresponds to a steep decline in moisture availability in the central United States. Reptiles also exhibit a westward decrease in diversity (Kiester, 1971) but the gradient is, not unexpectedly, much less steep than that for amphibians: reptiles are, in general, less affected by arid conditions than are but to Incr eases in harshness (as determined by apas than are birds or mammals. The South American vertebrate fauna offers considerable potential for evaluating this pre- iction, but distributional data are lacking or ru- dimentary for most taxa. Available information, owever, seems to support the third prediction. Diversity of amphibians (Lynch, 1979), reptiles (Dixon, 1979), and birds (Slud, 1976; Donahue et al., in prep.; Terborgh et al., 1984) is highest along the lower slopes of the Andes and the im- mediately adjacent lowlands as compared to the lowlands of the Amazonian basin itself. More- over, as expected, diversity decreases with in- creasing altitude and a rise in the harshness of montane environments. Species diversity patterns of plants, to the ex- tent that they have been resolved, parallel the data for vertebrates. Based on current knowl- edge, and on projections of the numbers of species estimated yet to be discovered and described, 1985] CRACRAFT BIOLOGICAL DIVERSIFICATION 803 Diversity (ES 59) 5 10 15 coast 1000} gastropods invertebrate = 2000F megafauna [9] a 3000F 4000F 5000 FIGURE 4. A topographic profile of the continental margin in the western Atlantic (at 35°N latitude) nos with P e diversity of representative invertebrate groups at the given depths (data from Rex, 1981a). The shape of th Vertical exaggeration about x130. plant diversity appears to be highest along the lower, tropical and subtropical slopes of the An- des when compared to the Amazonian basin (Gentry, 1982b, 1982c; pers. comm.). This must remain a tentative conclusion, of course, but the large number of species with very restricted ranges within the Andes is enormous and even many small mountain valleys le large numbers of narrowly endemic specie Geographic transects of conidia species di- versity that are appropriate for testing this pre- diction are scarce and potentially susceptible to influences other than that of lithospheric com- plexity. Alternative ways of examining the influ- ence of complexity on diversity will be pursued in subsequent predictions. A further evaluation of the third prediction can be made using data from the marine realm. In this case, longitudinal transects probably ex- hibit less geographic variation in harshness and thus presumably show fewer effects of extinction than do transects across continents. Three pat- terns of diversity/complexity are relevant for the third prediction. a. North Atlantic diversity gradients. From the coast, species diversity of benthic inverte- e two species diversity curves is exemplary of that found in other benthic marine sasa kese brates increases on the continental slope and rise but decreases again as abyssal depths are ap- proached (Hessler & Sanders, 1967; Sanders, 1968; Rex, 1981a). Indeed, diversity is highest for virtually all groups at depths of 2,000-3,500 meters (Rex, 1981a). Much attention has been paid to the causes of high diversity in the deep- sea (see Discussion), but one explanation has not been seriously considered, namely, increased lithospheric complexity. The slope and upper rise represent the region where continental and oceanic crustal regimes meet. Relative to shelfs, rises, and the abyssal plain, continental slopes are more highly com- plex geomorphologically, with many deep sub- marine canyons and gullies generally traversing their flanks (e.g., Uchupi, 1968). The relation- ship between the continental margin profile of e North American Atlantic coast and two rep- resentative diversity profiles for marine inver- tebrates is pictured in Figure 4. In this figure, the profile itself does not adequately convey the complexity ofthe slope and upper rise sufficiently because the scale is too small. It can be hypothesized, therefore, that in- creased diversity with increasing depth is related En = 804 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 West coast - 0 - - 1000 - - 2000 - - 3000 - - 4000 - - 5000 - - 6000 - - 7000 - East coast E 5. Topographic profiles of the Pacific and Atlantic continental margins of South America at 25°S A Bütude. The profiles begin at the coast and end active and much more complex g diversity along th e Pacific coast is postu complex region rie text). Vertical exaggeration about + id 1 1 : 1 1 + A 4 to the opportunities that complexity creates for hic isolation and differentiation. This h geograp y- pothesis differs markedly from the prevailing ecological explanations for these diversity pat- terns such as the time-stability hypothesis, which will be reviewed and analyzed in the Discussion. b. Active versus passive plate margins. Active plate margins such as the western coast of South America are much more complex geo- logically than passive margins as exemplified by the Atlantic (Fig. 5). Active margins gain their complexity from the extreme structural relief re- sulting from plate-plate interactions; passive continental margins, in contrast, may even de- grade in complexity through time due to the de- position of continental sediments. Consequently, we would predict that active margins, which are continually changing their configuration, should promote higher diversity. The available data seem to support the pre- diction. Thus, species diversity of bivalve mol- luscs is nearly twice as high along the western coast of South America than along the eastern coast (Stehli et al., 1967; Valentine, 1971). This at the abyssal plain ( eomorphologically than the passive margin of the Atlantic. Higher species tulated to result from increased rates of isolation in that spatially more x 70. ). The Pacific margin is tectonically pattern of diversity extends northward to North America but there the pattern is confounded by Late Cenozoic extinction event in the western Atlantic (Stanley & Campbell, 1981). arge-scale diversity patterns of the Pacif- ic. The Pacific basin shows considerable vari- ation i from being the least complex on the abyssal plain of the central Pacific, to very highly complex in the Indo-Australian region where microplates, is- land arcs, and areas of subduction abound. Using this gradient in complexity as a predictor of di- versity, greater diversity would be expected along the western margin of the Pacific. This is pre- cisely what is seen The highest diversity of bivalve molluscs, for example, is found in Australasian waters (Stehli etal., 1967), with over 1,000 species as compare to the 150-250 species found in the central and eastern Pacific. Zooxanthellate (Rosen, 1981) and hermatypic corals (Stehli & Wells, 1971) also show greatest diversity in the western Pacific. Stehli (1968) presented data for other groups (e.g., gastropods) having a similar pattern of diversity. Prediction 4. Large clades, widely distribut- 1985] TABLE 2. Patterns of diversity in neotropical fly- db. (Tyrannidae) and tanagers (Thraupidae). en- tral Ama- Ameri- ZO- Andean can nian Region Region Region Relative area 0.27 0.30 1.0 Rank order in geomor- phological complexity l 2 3 Tyrannidae’ Differentiated taxa 310 177 184 Rank order 1 3 2 Differentiated taxa/ genus 6.89 4.12 3.76 Thraupidae’ Differentiated taxa 297 123 105 Rank order 1 2 3 Differentiated taxa/ genus 9.00 6.15 5.53 a Data for the tyrannids from Traylor (1979). * Data for the thraupids from Paynter and Storer ). (1970 ed over the same areas, will exhibit covarying gradients in diversity that parallel the gradient in lithospheric complexity. The third prediction was primarily a statement about gradients of diversity averaged over many clades, that is, a statement about biotic patterns. The fourth prediction partitions that biotic pat- tern and makes a statement about the causes of diversity within and among the clades them- selves. If a clade is distributed widely over an area that has significant variation in geomorpholog- ical complexity, the hypothesis of lithospheric complexity predicts diversity will be higher in regions of higher complexity. But again other causal factors must be held constant, thus we assume that taxa within the clade have been dis- tributed in the different areas for about the same amount of time, and that extinction rates have been roughly equivalent across the entire distri- bution. If these assumptions are satisfied, then complexity should ied as the primary pre- dictor of diversity pattern In order to examine this m" two highly diverse and widespread avian families of tropical America were examined, the tanagers (Thrau- pidae) and the flycatchers (Tyrannidae). Three biogeographic regions, the Andes (upper part of tropical zone and above), Central America, and CRACRAFT — BIOLOGICAL DIVERSIFICATION 805 tropical lowland Amazonia, were chosen for comparison, with each representing an area of very high, intermediate, and low geomorpholog- ical complexity, respectively. Diversity was cal- culated by tabulating the numbers of endemic subspecies (in the case of polytypic species) or monotypic species in each region, following the recent systematic treatments of the Peters' “Check-list of Birds of the World" (Paynter & Storer, 1970; Traylor, 1979); subspecific, rather than species-level, taxa were ennumerated in or- der to estimate more accurately the number of differentiated evolutionary units (Cracraft, 1983a). The results are shown in Table 2. Diversity in both families parallels the gradient in litho- spheric complexity. It is also noteworthy that this is especially true even when area is taken into account; diversity is not related to a simple area- effect. Amazonia covers a much greater area, yet contains only about the same number of differ- entiated taxa within these two families as does Central America. Also revealing is the number of taxa per genus that have differentiated in the three areas. For both families, the Andean region shows the greatest average amount of differen- tiation per genus, followed by Central America, and then Amazonia. OTHER RATE-CONTROLS OF SPECIATION Spatial and temporal variation in lithospheric complexity has been hypothesized above to effect a major control on speciation rates, and thus to contribute significantly to explaining patterns of diversification. Other factors are also widely ac- knowledged to contribute to variation in specia- tion rates, and the purpose of this section is to assess their contribution. Not all the evidence tion. It will be argued that, although més mw in some cases perhaps significantid so), they ap- pear to have had a limited impact on defining major spatial patterns of diversity. INTERNAL MORPHOGENETIC COMPLEXITY Differentiation following isolation is a func- tion of the amount of morphogenetic complexity 806 within the individuals of a species. If those in- dividuals exhibit little or no variation, then the rate-change in character divergence following isolation should be low, and thus too the prob- ability of differentiation (Stanley, 1975, Wiley & B ). Morphogenetic com- plexity means the amount of variation as ex- pressed within the developmental pathways of individuals of a population, a notion equivalent to the concept of canalized information as used by Wiley and Brooks (1982). Any changes within the genome that have effects on developmental athways, and thus on phenotypic expression, are potentially important in establishing the probability that a population will differentiate once it becomes isolated. Thus, variation in mor- phogenetic complexity within the species of a clade might influence their speciation rate (Cra- craft, 1982a): clades comprised of species having high internal complexity should speciate at a higher average rate than clades with less complex species (all things being equal). Present evidence suggests, for example, that genomic modifications such as gene duplication, polyploidy, and in addition to point mutations, can alter developmental pathways and thus adult phenotypes (Wilson et al., 1977). Asa consequence, the prediction might be made that if individuals within a clade are characterized by these types of genomic modi- fications, then the probability of speciation will be relatively high. Wilson et al. (1977), in fact, proposed that speciation rates in mammals have been higher than those for anurans partially be- cause of a higher rate of chromosomal change in the former. Levin and Wilson (1976) made a comparable claim to explain patterns of diversity within angiosperms. Properly designed comparative studies should give us much more insight into the causal rela- tionship between large-scale genomic change and ciation rates within and among clades. If this relationship proves significant, the question will arise as to how that might be relevant at the level of diversity gradients within biotas. At first glance, it might appear difficult to conceive of numerous clades exhibiting congruent patterns of genomic modification among their species, yet such geo- graphic gradients have been suspected for some time. The incidence of polyploidy, for instance, increases with increasing latitude (Grant, 1971); what effect that has had on geographic variation in speciation rates is a separate, as yet unan- swered, question ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 BEHAVIORAL-ECOLOGICAL ATTRIBUTES OF SPECIES Biologists have long hypothesized a causal connection between certain behavioral-ecologi- cal attributes of species and the likelihood that they might speciate. Unfortunately, conflicting predictions or expectations are common. Sub- stantial literature suggests that ecological gen- eralists (or eurytopes) would be more likely to topes (see Parsons, 1983 for a summ Kohn, 1971; MacArthur, 1972; Diamond, 1975, among many others). Yet, stenotopy, rather than eurytopy, has been hypothesized to lead to spe- ciation through small, ecol 1980; Vrba, 1980, 1983). The best Spi ds ta supporting this latter hypothesis are those of Ja- blonski (1982; see also Valentine & Jablonski, 1983),in which d nonplanktotrophic larvae (stenotopes) presum- ably speciate at higher rates than those with planktotrophic larvae (eurytopes). While it seems clear that possession of certain ecological or behavioral characteristics may help determine the probability of speciation for species within a clade, this can translate into geographic patterns of diversity only if the characteristics themselves vary geographically, if they vary in the right direction, and if the different clades of organisms comprising the biota being investi- Wi th +L ge £e displayed geographically. Such a conjunction of events suggests this woul occur Hd very rarely, f speciation rate- control probably has little iniuence on geograph- ic diversity gradients. SEXUAL SELECTION Fisher (1930) proposed that a genetic corre- lation between assortative mating created by fe- male choice, on the one hand, and secondary sexual characteristics of males (the basis for the choice), on the other, can lead to rapid evolu- tionary change within populations. Lande (1981, 1982) and West-Eberhard (1983) have also ar- h be responsible for high rates of speciation in some ups. In theory, this mechanism for rapid divergence might be potentially important in those groups of organisms possessing the appropriate mating 1985] systems (West-Eberhard, 1983). Yet, an upper- level constraint on this process of divergence would still exist and include those factors con- trolling the rate of geographic isolation; in most cases, sexual selection itself could influence the rate of divergence only after populations became isolated. The influence of sexual selection on speciation rate thus remains highly speculative. Unless the incidence of sexual selection varies geographi- cally —e.g., if sexual selection were more frequent in highly diverse ecosystems (as perhaps is the case)—then its contribution to establishing geo- graphic gradients in diversity would be minimal. Even if this geographic variation existed, how- ever, speciation would not be expected to occur faster than the rate of geographic isolation. THE RATE-CONTROL OF EXTINCTION Despite a large literature, biology still lacks a widely accepted deterministic theory of extinc- tion. Emphasis has been placed on the occur- rence and explanation of short-term pulses (so called “mass” extinctions) rather than on the pat- terns and processes of normal “background” ex- tinction. Another factor mitigating against the development of a theory of extinction has been the difficulty of testing any proposed hypothesis. The analysis of causation within extinction the- ory is particularly susceptible to scale effects: at one level climate may change, thereby initiating numerous environmental consequences leading to extinction, but at th time that extinction also can be examined causally at the more “‘prox- imate" level of processes within populations. At issue here are patterns of extinction calling for explanations at hierarchical levels above that of populations. Such patterns are taxonomic, at the level of clades, and biotic, at the level of assemblages of species, or some combination of both. Thus, taxonomic patterns of extinction within or among clades can be expressed either geographically or temporally at a single location. Patterns of extinction within biotas are statistical mmations of extinction within and among clades. Within a biota, changes in extinction rate are expressed temporally, whereas patterns be- tween biotas can be both temporal and geograph- ic. These large-scale patterns, above the level of populations, can be expected to have first-order causes which themselves are expressed at a hi- erarchical level above that of populations. It is CRACRAFT — BIOLOGICAL DIVERSIFICATION 807 proposed here that change in the temporal and geographic gradient of environmental harshness (favorableness) is the primary control of extinc- tion rates, and thus of the large-scale extinction patterns just mentioned. This hypothesis will developed in more detail now and later will be compared with other potential rate-controls of extinction. THE HYPOTHESIS OF ENVIRONMENTAL HARSHNESS The twin concepts of environmental harshness and favorableness have been discussed exten- sively in the ecological literature (especially An- drewartha & Birch, 1954). Most of that literature relates harshness with density-independent pop- ulation regulation (see Enright, 1976 for a dis- cussion of how harshness might function within a density-dependent context) The hypothesis presented here expands that thinking to account for large-scale gradients in extinction. Premise Individual organisms (and thus populations "e species) possess a definable physiological capacity permitting them to exist within a specific environmental regime; no or- ganism or species can live in all environments. Premise 2. As the physical characteristics of an environment shift away from the “optimal” conditions for a species, the intrinsic rate of in- crease declines and eventually becomes negative, resulting in extinction of the population. In this case, we define “optimal” in terms of the conditions (narrow or broad) at which the rate of increase is at or near maximum, but other measures could also be chosen. This premise has considerable empirical support, and physical at- tributes of the environment are known to influ- ence physiological functions, which in turn have effects on demographic parameters such as fe- cundity, survivorship, age of reproduction, among others (Andrewartha & Birch, 1954; Ricklefs, 1973; Krebs, 1978). Premise 3. The environments of the earth can be arranged | along a scale defined i in ters of the Ule Te, I olosical tolerances of a random sample of the world’ nisms. = ajah physical parameters have the potential to influence the survivorship and reproductive eepapiities d organisms, Moisture and tengper- B to ex- erta greater overall effect than do other param- eters, and that assumption is adopted here. Therefore, the premise states that, in principle, 808 700 (000 60001 5000 4 4000 RAINFALL (mm) 3000 4 2000 4 1000 ° ° 0 5 10 15 20 25 TEMPERATURE (oC) A theoretical diagram postulating a re- lationship between the probability of extinction and the position of a population in a space of environ- mental base q sp DE as measured i in terms of mean annual rainfall and temperatu da ter Birch, 1953 and Krebs, 1978). the joint influences of temperature and moisture on the intrinsic rate of natural increase can be experimentally determined for each species (e.g., the classic study of Birch, 1953), and a measure of that influence (ahus RET E can be mapped geographically. Premise 4. The frequency of species occur- rences along the harshness/favorableness scale is decidedly nonrandom. Also having much empirical support, this premise merely states that environmental ex- tremes in temperature and/or moisture are tol- erated by relatively few organisms. In areas of low harshness (generally speaking, in environ- ments that are warm and moist; see below), organisms require less energy to maintain phys- iological homeostasis relative to their surround- ings than do organisms in harsher environments. These premises yield the following hypothesis: the probability of extinction is a function of the change in the valųe of environmental harshness to which a species is exposed (Fig. more explicit what bi- also makes a larger claim not often recognized: that taxonomic and biotic patterns of extinction are primarily a function of spatio- temporal changes in environmental harshness. Ecologists have long proposed a relationship be- tween environmental harshness/favorableness ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 and diversity (e.g., Richerson & Lum, 1980), but most of these discussions have not pinpointed the causal connection of this relationship: that is, harshness influences only one component of diversification, namely extinction rate (see Con- nell, 1975 for a discussion of the ways in which harshness gradients influence community struc- ture). Extinction rates are exceedingly difficult to study because their analysis is more dependent on the quality of the fossil record than is the analysis of speciation rates. For example, if we compare Recent sister-clades differing greatly in diversity, the minimal number of speciation events for each clade can be specified (N-1, where N is the number of species) even in the absence of a fossil record; relative rates can thus be es- timated directly. On the other hand, a compa- rable statement regarding the number of extinc- tions in each lineage is not possible in the absence of a fossil record. A method must be found to document the loss of a species, not its mere pres- ence, and a historical record is our only way of doing this. Measurements of extinction rates based upon an analysis of the fossil record entail additional difficulties in that paleontologists rarely use species-level taxa but often are compelled in- stead to study taxa at the generic or family levels, precisely because of the lack of detail in the fossil record. Unless the supraspecific taxa contain the same average number of species (and with little variance), however, comparisons among them will not yield accurate statements about the rel- ative amounts of extinction at the relevant evo- lutionary level, namely that of species. Evaluation ofthe hypothesis of environmental harshness not only relies upon the fossil record for an estimation of relative rates of extinction, e a moisture, are retrievable from both paleontolog- ical data and from physical and chemical char- acteristics of the rock record itself. Unfortu- nately, detailed study of the relationship between extinction rate and harshness calls for a well- preserved historical record, which is usually lacking for terrestrial environments. Despite its inadequacies, paleontology can provide, never- theless, abundant data at a coarse-grained level with which to evaluate the hypothesis of envi- ronmental harshness. 1985] A major prediction of this hypothesis is that as a specific num gei harsher, the average extinction rate o of the biota will rise. As defined earlier, harshness camnonen t speci es extreme values. Under these conditions, more organisms become physiologically stressed, pop- ulation sizes decline, and the probability of ex- tinction increases. Paleontologists have docu- mented extinction d harsh- Pies 1 er dhvironments (Axelrod, 1967). For example, the marked increase in elevation of the Andes in the middle to late Cenozoic resulted in a change toward extreme aridification along the west coast of South America, thereby causing a once diverse biota to become extinct and be replaced by one much less so (Berry, 1938). A widespread, rela- tively diverse biota disappeared in North Afric as the climate became more arid in the late Ceno- zoic (Moreau, 1232, 1300) IUD changes in lso took place in Australia (Kemp, 1981; Galloway & Kemp, Increases in harshness as measured by tem- perature are also known to have increased ex- tinction rates. Hickey (1981, 1984) reported a gradual increase in the extinction rates of floras as global temperatures became cooler across the Cretaceous-Tertiary boundary. Furthermore, ex- tinction rates were higher in higher latitudes where the temperature gradient was steeper. That de- viations in temperature away from physiologi- cally more favorable regimes increase extinction rates of floras has been noted by many workers, ndp psthe p ith the Pleistocene climate fluctuations (e.g., Axelrod, 1967; Kershaw, 1981). Finally, in the marine realm, temperature change is also strong- ly implicated in large-scale extinction events (Stanley, 1984; Kauffman, 1984). That large-scale, geographic increases in en- vironmental harshness result in concomitant re- sponses in the steepness of extinction gradients can scarcely be denied, yet this does not inform us about the relative role of increased harshness, when compared to other postulated causes of extinction, in effecting a control over patterns of diversification. At the level of biotas, changes in harshness seem to play an important part in the evolution of geographic patterns of diversity, but their relative contribution to controlling patterns of diversity, within and among clades is some- what more difficult to establish because a rela- CRACRAFT —BIOLOGICAL DIVERSIFICATION 809 tively extensive fossil record is qua. Mos changes in harshness should alter extinction: rates of clades nonrandomly (e.g., as in latitudinal gra- dients). Changes in harshness, however, often re- sult from relatively local tectonic evolution, and thus patterns of extinction within clades might exhibit a “‘stochastic” appearance over a large geographic area (i.e., clades would go extinct, in this case, purely as a result of being in the wrong place at the wrong time). Many clades show a pattern of reduction di- versity during their history, but the causes of this common pattern have not yet been analyzed comparatively. Unfortunately, it is not possible at this time to evaluate the generality of the en- vironmental harshness hypothesis by reference to a large suite of phylogenetically controlled data. As an alternative, we can undertake a partial evaluation by examining other postulated causes of extinction and asse their potential for ex- plaining bot of biotic diversity. a. Tectonic elimination of habitats. Tec- tonic activity brought about by lithospheric plate motion directly obliterates habitats (e.g., shallow and desp-water ana environments during continent changes in their extent through mountain building, alterations in sea-level, or volcanism (Valentine & Moores, 1970, 1972; Valentine, 1971, 1973; Schopf, 1974; Bakker, 1977; Hallam, 1981; Axelrod, 1981). Tectonic events can have major effects on the extinction patterns of local biotas, and generally are recognized to contribute to mass extinctions. In particular, if these events have a large geo- graphic scale, then they can alter patterns of di- versity within higher taxa (examples can be found among the major Phanerozoic mass extinction events). Whether tectonic events are a major, direct control on long-term patterns of reduction diversity within clades is questionable, although marine regressions and transgressions can span many millions of years and might alter diversity gradually. Tectonic activity perm se is thus not likely to be e thep of most dients in extinction o or of long-term patterns of reduction diversity (particularly within terrestri- al organisms). But tectonic alterations of envi- ronments play a significant role in geologically “short-term” mass extinctions, and thereby coul leave their signature on “long-term” patterns of diversity within clades. The extent to which these 810 geological events could help explain congruent changes in diversity over many clades would de- pend upon the magnitude of the events them- selves, their location, and the clades of organisms that happen to be exposed to them. b. Competition. Ecologists have long iden- tified competition as a major cause of extinction within populations, and the tradition goes back to Darwin himself. Paleontologists and others interested in large-scale patterns of diversifica- tion have adopted many ecological ideas regard- ing competition and its role "m extinction and have reified tl higher taxa. A common application of this think- ing occurs when one group of organisms is said to “replace” another, trophically similar group over a period of time (perhaps many millions of years). Well-known examples ascribed to com- petition include the replacement of e by Vane (Simpson, 1953), South American arsupials by North icon placentals (Simpson, 1950), multituberculates by primitive placentals (Simpson, 1953; Van Valen & Sloan, 1966), dinosaurs by placental mammals (Van Valen & Sloan, 1977), and even invertebrates by primitive gnathostome fishes (Thomson, 1977), to name only a few. If the causal assessment of these replacements is correct, then by implica- None of these putative examples is well doc- umented, however, at least by the standards cur- rently acceptable within modern ecology (Wiens, 1977; Connell, 1975, 1980, 1983). Many diffi- culties arise, first in recognizing the mere pres- ence of competition within Recent communities, and then, second, in determinin the problems become intractable. Showing that one species competitively replaces another re- quires extensive ecological analysis; arguing that many species of one clade (or one biota) can competitively replace many species of another clade or biota requires detailed data that a fossil record is incapable of supplying. The “replace- ment" can be documented, but a causal hypoth- esis of competition cannot. The proposition that competition is a major reach this conclusion, moreover, even accepting ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 the hypothesis that competition is ubiquitous and important in the regulation of species abun- dances within local communities. But given that this latter hypothesis is still being debated within ecology, it seems premature, and perhaps pre- sumptive, to conclude that competition is a ma- or causative agent of extinction either within clades or between biotas (see Gould & Calloway, 1980, for an analysis ofthe “classic” brachiopod- to-clam replacement, which they conclude re- sulted Ll the Permian extinction event not d, . Pr DM Predation is postulated to be an Gaga factor in regulating relative abun- dances within populations. Depending upon the ecological system being studied, the presence and effect of predation may or may not be easily de- termined. Within paleontological assemblages, predation has been investigated most Mb aed in marine invertebrates in which evidence is often obtained directly from the fossils ace. These studies suggest the possibility that pre- dation pressure has increased the rates in some clades or organisms. Stanley (1977), for example, postulated that the post-Paleozoic in- crease in the frequency of shelled infaunal bi- valves as compared to exposed epifaunal forms was the result of more intense predation on the latter. This “species selection," Stanley (1977: 238) proposed, led to higher extinction rates of epifaunal species and major changes in diversity within and between clades. Likewise, Stanley (1977) attributed the post-Late Mesozoic decline of articulate brachiopods to increased predation from teleost fish, snails, and crabs e significance of predation within ecosys- tems cannot be denied, yet acceptance of its im- portance as a causal agent controlling rates of extinction is less warranted. Under most natural conditions, predators do not extirpate their prey, although at times they may affect their numbers substantially. If predation does not cause ex- tinction of a prey species directly, population sizes might be so reduced that extinction could occur stochastically or via other agents. Preda- tion might be expected to play a role in species extinction in communities with simple structure in which prey species cannot (or have failed to) develop any effective means of predator avoid- ance. In contrast, if prey species are widely dis- tributed or have high dispersal abilities (or are unavailable to predators) during some part of the life cycle, then the probability of extinction via predation would be expected to be relatively low. T. 1985] € 0 SD C PR J K T FIGURE 7. Phanerozoic scape pees curve for marine invertebrates showing a rise in diver- sity in the Cretaceous and Tertiary (after Raup, 1976a and Sepkoski et al., 1981) Asa consequence of these factors, predation will probably not be an effective mediator of extinc- tion rates in most groups of organisms. Any hy- pothesis that predation is important needs to be critically evaluated and thoroughly documented. d. Behavioral characteristics. The degree of behavioral-ecological plasticity of species has been postulated to influence the probability of extinction. Jackson (1974), Eldredge (1979), El- dredge and prat (1980), and Vrba (1980) have suggested tha ytopic species will be less likely to go extinct uu apa Eurytopic species tolerate a relatively wid and are thus assumed to n more widely distrib- uted and have larger overall population sizes than stenotopes. Extinction rates are postulated to be higher in stenotopes because of their smaller, more narrowly distributed populations. The correlations (e.g., Vrba, 1980) between eurytopy and clades of low diversity, and be- tween stenotopy and pi of higher diversity, are not easily interpre sumably have higher ende of both ex- tinction and speciation; in such cases, therefore, high speciation rates would be postulated to override high extinction rates. Eurytopic species, in contrast, are assumed to be characterized by both low extinction and low speciation rates. If eurytopic clades are generally characterized by low diversity, then that would probably result from the low speciation rates and not low ex- tinction rates, which should instead promote high diversity. Consequently, this suggests that sten- otopy and eurytopy, to the extent that they in- fluence intracladal diversity, do so through the mediation of speciation rates and not through effects on extinction. The preceding discussion makes clear the com- plex causation underlying extinction rate-con- trols. Any temporal change in diversity that is CRACRAFT — BIOLOGICAL DIVERSIFICATION 811 attributable to extinction rather than speciation almost certainly will be a manifestation of dif- f£ 4 1 PF. ° 14 1 Accepting this, we should seek to identify the relative contributions of each cause to the extent possible. In most cases, the available data prob- ably will not permit us to discriminate muc more than one or two main factors. The evidence reviewed here suggests that in- tracladal patterns of reduction diversity and geo- graphic gradients of diversity, insofar as they re- flect variation in extinction, are generally the result of gradients in environmental harshness. This claim is merely one of frequency. Many kinds of tectonic activity can alter extinction rates of biotas on a local scale or sometimes even over areas of considerable extent. Behavioral attri- butes such as eurytopy and stenotopy have the potential to influence extinction rates, although because they can also affect speciation rates, their analysis is complicated and their role in extinc- tion is questionable. In principle, competition and Pie through their effects on species s, might contribute to regulating the Moreover, even though subject to these presumed density-dependent in- teractions, density-independent factors might actually be the primary controls on mortality (Enright, 1976). The conclusion is that only en- vironmental harshness is likely to constitute a significant influence on the types of extinction atterns considered here. Because of its perva- sive, and well-documented, influence on extinc- tion, the effects of environmental harshness should be eliminated prior to considering other causal hypotheses. Such a procedure will ensure that those hypotheses are evaluated under the best possible circumstances. DISCUSSION WHAT IS THE SHAPE OF THE PHANEROZOIC DIVERSITY CURVE? The Phanerozoic diversity curve has been the subject of much discussion over the past decade (Valentine, 1972; Raup, 1972, 1976a, 1976b; Bambach, 1977; Sepkoski, 1978, 1979; Sepkoski et al., 1981; Signor, 1982). Arguments have cen- tered on the empirical shape of the curve itself, in particular whether diversity has been at equi- librium since the end ofthe Paleozoic or whether 812 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 200 MARINE METAZOAN ORDERS 100 4 v |[€jejsjD;c|PJ|* J | kit 600 400 200 0 FiGURE 8. A Phanerozoic diversity curve for marine “orders” of Metazoans (from Sepkoski, 1978). Curves n as ps have been used to support the hypothesis that diversity has been in equilibrium since the late Paleo there was a late Mesozoic-Cenozoic rise. Various authorities now generally agree that present data show a curve for marine invertebrates that rises abruptly from a Cambrian low to one of apparent equilibrium through the Permian, followed by a dip in the Triassic, and a subsequent increase in the late Mesozoic and Cenozoic to levels that were four to eight times those of the Paleozoic and early Mesozoic (Sepkoski et al., 1981; Sig nor, 1982; Fig. 7) Our ee here is not with the actual em- pirical shape of the curve but with its expected shape given specific models of diversification. With respect to the former, one might argue that available data, which are primarily of marine organisms, are inadequate to test any of the curves that might be predicted. For one thing, a mean- ingful evaluation of the pattern of diversification cannot be restricted solely to marine taxa (e.g., Sepkoski et al., 1981), because many of them also have radiated into nonmarine environ- ments. Consequently, although the marine pat- tern is of interest in its own right, most workers would probably agree that it cannot be the bench- k of the Phanerozoic curve itself. Restriction sample of the results of diversification; none of these ecological subdivisions is a closed system as far as diversification is concerned. Furthermore, estimations of the shape of the diversity curve based on counting supraspecific taxa are inherently biased and inaccurate (Cra- craft, 1981b, 1982a; Novacek & Norell, 1982; and earlier discussion). Higher taxa are not lutionary units within those taxa. Finally, the Phanerozoic record for freshwater and terrestrial environments is much too poor to provide an accurate estimate of the numbers of species from one time period to the next. As a simple example, the estimated number of living arthropods increased tenfold (to perhaps 30-40 million species) in the last few years after it was realized just how diverse the tropical dad can- opy is (T. L. Erwin, 1982; pers. comm.). Tropical invertebrate faunas, in general, are still virtually nknown. Given record, accurate estimates of diversity based on that record should not be expected. o summarize, we can only praise the efforts of those who have attempted the arduous task 1985] DIVERSITY EQUILIBRIUM MODEL TIME NONEQUILIBRIUM MODEL .— A. Curve illustrating the equilibrium model of biotic diversification. Perturbati M . À nonequilibrium model of biotic diversi- fication. An intrinsic exponential i increase in diversity is T with the rate of increase constrained by wor harshness/fa- voribien tions are pin to “reset” the curve (see text). of compiling patterns of diversity from the fossil record, and their emphasis on the marine fossil record is readily understandable. The message here is merely that some skepticism is necessary regarding what these curves actually tell us. They may reveal an accurate picture of diversification patterns within the marine realm itself, yet clear- ly that record does not depict actual patterns of diversification of all those clades within the bio- sphere as a w Consequently, the question can be asked: what might be the expected shape of the Phanerozoic diversity curve? Investigators of marine diversity have often assumed the marine environment to be a closed (or semi-closed) system, and the di- versity curve therefore to be logistic, or equilib- rial (Sepkoski, 1978, 1979; Figs. 8, 9A). Within such a system, perturbations such as mass ex- tinctions depress the curve, but diversity even- tually rebounds to the equilibrial value. Only CRACRAFT — BIOLOGICAL DIVERSIFICATION 813 minor variation above equilibrial values would be expected. If diversification is controlled significantly by lithospheric evolution, isolate formation would be some exponential function of the rate of bar- rier formation (Cracraft, in prep.). Diversity, therefore, would increase through time. But the path of the diversity curve is constrained by ex- tinction, including both mass and the back- ground rate. In this model, mass extinctions “‘re- set" the curve of diversification, not perturb it from an equilibrial value (Fig. 9B), and changes in the background rate of extinction— P woul occur, for example, ing or warming— operate to calibrate the expo- nential rise function. Ecologically, a diversity-independent diversi- fication model would not entail a saturation of the resource-space of the biosphere. Indeed, in the absence of external constraints (extinction), that space should grow through time. When dif- ferentiation produces new species, p also pro- duces taxa having new (or modified) ways of uti- lizing the physical and biotic space of the biosphere. Because different trophic levels of a biota have different relative abundances, re- peated vicariance of that biota might be expected to yield different diversification rates within each trophic level: we would predict, for example, that the primary consumer (herbivore) level would increase at a higher rate than the secondary con- sumer (carnivore) level. This may explain the inordinately high numbers of herbivores in sys- tems that have experienced relatively few ex- tinction events over time (e.g., terrestrial envi- ronments in the tropics as compared to temperate regions; see below). Empirically, the Phanerozoic diversity curve seems to fit the diversity-independent model more closely than an equilibrial model. Sepkoski et al. (1981) and Signor (1982) document a pre- cipitous rise in marine diversity since the Trias- sic, exactly what might be expected given the absence of a major mass extinction during that time (only two “intermediate” events are re- corded in the marine record; Sepkoski, 1982). Because this rise is so far above the Pulcozbió “equilibrium” level, an equilibrial model cannot easily explain it, whereas the diversity-indepen- dent model can. The hypothesis that Phanerozoic diversity was at some equilibrial value from the Ordovician to the end of the Permian (almost 250 Ma) is also open to challenge and to the claim that di- 814 aqp en could have been nonequilibrial (Ross, 1972). First, two significant extinction events, H Ashgillian in the late Ordovician and the Frasnian in the middle to late Devonian (Raup & Sepkoski, 1982), can be hypothesized to have "reset" the diversification curve of the marine a ea arboniferous until the end of the Permian, many other groups were apparently radiating into ter- restrial and freshwater environments at that time: e.g., amphibians (Carroll, 1977), conifers (Florin, 1951). These groups are underrepresented in cal- culations of the Phanerozoic species diversity rve. Finally, the Permo-Carboniferous was a pe- riod of harsh global climates (Frakes et al., 1975; Olson & Vaughn, 1970), which can be postulated to have constrained diversification. If true, then d expected exponential rise might have been mped," thus M EREMO. to the appearance Eo a diversity equilibri Insuffici lent evidence. thus eR 1 eee over that I all pattern of equilibrium. Moreover, there is little reason to expect that speciation and ex- tinction are diversity-dependent (Cracraft, in prep.). The expectation of equilibrium, which has been so widely accepted within the paleon- tological community, has arisen from an accep- tance of only one view within contemporary ecology regarding population regulation and from the assumption that processes at the hierarchical level of clades and biotas are simple extrapola- tions of ecological processes within and between populations. As these latter processes do not ap- pear to constitute first-order determinants of spe- ciation or extinction, that extrapolation is prob- ably not valid. IS THERE AN UPPER LIMIT TO DIVERSITY? Acceptance of the idea that biomass has at- tained an equilibrial value within the biosphere has been the basis for concluding there exists an upper limit to diversity (MacArthur, 1965, 1969; Margalef, 1972; Levinton, 1979). The previous discussion, however, has questioned the under- lying assumption of a limit to the evolution of biomass, and that viewpoint will be amplified here (see also Benton, 1979; Cracraft, in prep.). As long as the biosphere is an open thermo- dynamic system (Ulanowicz, 1980), the influx of energy and matter will continue to be captured ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 400 € S Rain forest z & 30°! (250-400) E < E a 200 | S Deciduous forest LU (70-250) c a < 100| Savanna Coniferous forest ul (5-35) x (25-70) s Prairie Tundra Desert (1-18) (0-1) (5) 30 20 10 0 -10 MEAN TEMPERATURE ( C) FIGURE 10. Biomass production (tons/hectare) of differ in environmental harshness/favorableness, as mea- sured by mean annual temperature and precipitation (after Smith, 1980). Potential biomass production is constrained by worldwide patterns of environmental harshness (see text). by organisms, ultimately through the processes of reproduction and ontogeny, with biomass thus increasing. Degradation of the earth by chemical and physical processes, and also by organisms, releases matter, some of which is incorporated into living organisms. Incoming solar energy is transduced and used to manufacture organic ma- terial, yet only a very small portion (196 or less; ebs, 1978: 522-524) of the available energy is utilized. Given these observations, then, an in- crease in biomass would appear to be inevitable, unless some extrinsic factor can be shown to con- strain the flow of energy and matter. Ecologists have often viewed the evolution of to the evolution of biomass in yet another way. e flux of energy and matter through individuals and populations is the mo- tive force for ontogenetic and evolutionary dif- ferentiation (Wiley & Brooks, 1982; Brooks & Wiley, 1984). Thus, diversification takes place as a consequence of biomass expansion. More- over, as long as energy flow through the litho- sphere continues to drive its evolution, taxo- nomic differentiation will persist unabated ut are there no limits to biomass evolution? Biologically more meaningful, perhaps, is to speak of constraints on biomass evolution within the 1985] context of a nonequilibrium increase (Ulano- wicz, 1980). Through the history of life the most important constraint has probably been a steep gradient in environmental harshness. On the earth today, for example, climates of harsh tempera- ture and moisture regimes restrict net primary productivity (Fig. 10). This observation also im- plies that space itself has not been a limiting factor of global biomass production. At any one location, however, biomass production could be limited by a variety of factors, abiotic or biotic. ECOLOGICAL EXPLANATIONS AND THE ANALYSIS OF LARGE-SCALE DIVERSITY GRADIENTS Ecologists have long played a leading role in the analysis of diversity, particularly since Hutchinson's (1959) seminal paper. These efforts have been directed primarily toward the solution of two questions: (a) How can latitudinal gra- dients in diversity be explained? and (b) How can variation in diversity among different hab- itats be explained? By and large ecologists have considered the answer to the first question to be an extension of the answer to the second. The discussion that follows takes the approach that the answers to the two questions are largely in- dependent —that is, mes is a logical or bio- logical corollary of the o Ecological dubiis to divers have been extensively reviewed by others (Connell & Orias, 1964; MacArthur, 1965; Pianka, 1966; Whitta- ker, 1972, 1977; Ricklefs, 1973; Rosen, 1981; Thiery, 1982), and although each categorizes po- tential explanations somewhat differently, a common list can be generated. Explanation 1. The time-stability hypothesis states that the longer a community has main- tained stability and not been subject to major disturbance (extinction), the more species will accumulate. ` Explanation 2. The spatial heterogeneity hy- pothesis predicts more species will co-exist in environments that are more complex spatially. As Pianka (1966: 36—37) noted, the concept of spatial heterogeneity is also a problem of scale, and ecologists, more often than not, refer to local within- or between-habitat complexity when comparing differences in diversity (MacArthur et al., 1962, 1966; MacArthur, 1964; Ricklefs, 1977). Explanation 3. The productivity hypothesis states that as environments increase in produc- CRACRAFT — BIOLOGICAL DIVERSIFICATION 815 tivity, diversity will increase (Connell & Orias, 964) Explanation 4. The competition hypothesis argues that as competition increases, organisms become more specialized and niche sizes de- crease, and hence diversity can increase (MacArthur, 1965, 1972). A variant of this hy- pothesis states that the evolutionary impetus to reduce competition results in more niche overlap and thus higher species diversity (Klopfer & MacArthur, 1961 Explanation 5. The predation hypothesis states that as predators increase in numbers, they reduce population sizes of their prey, thus allow- ing more of those species to coexist (Paine, 1966; Janzen, 1970). Ecologists have "em ue | to pont out UM no single hypothesis al gra in diversity, and co RR some combina- tion of the above five explanations is generally invoked (Pianka, 1966; Menge & Sutherland, 1976; Huston, 1979). Yet even this misses the point because supporters of these hypotheses rarely address the question of the rate-control of speciation and extinction (Ricklefs, 1973 is an exception), which, it has been argued, is the first- order cause of large-scale diversity gradients. Ecological explanations have generally con- founded two issues: the generation of diversity versus its maintenance. By and large, ecologists have spoken only to the latter, and thus a causal relationship between their explanations and the ios determinants of diversity — specia- on and extinction— usually is not addressed. exte (1973: 714) classification of diversity hypotheses is one attempt to do this, but that effort primarily functions to expose the incon- sistencies among some of these hypotheses. For example, competition is said to be more intense in tropical situations, thus increasing natural se- lection and thereby leading to faster evolution (Dobzhansky, 1950). Yet at the same time, com- petition is claimed to be /ess stringent in the tropics, thus leading to lower extinction rates (Ricklefs, 1973: 714, table 43-5). Likewise, great- r “productivity” is hypothesized to lead to greater turnover of populations, thus increased natural selection, and faster evolutionary (spe- ciation) rates; in contrast, more “resources” also are said to result in less competition, thus damp- ing natural selection and thereby lowering ex- tinction rates. The hypothesis advanced here is that these presumed ecological correlates of high diversity, 816 rather than being causes, are simply effects of having more species together in the same place. None of them is sufficient to account, in any regular manner, for the spatial gradients of spe- ciation and extinction rates that are necessary to produce observed patterns of diversity. In con- trast, these ecological correlates can be explained as consequences of those same gradients. Thus, higher productivity, narrower niches, more niche overlap, more competition, more predation, and more habitat heterogeneity might be expected in communities having relatively more species. DEEP-SEA DIVERSITY AND TIME-STABILITY HYPOTHESIS One of the most intriguing spatial gradients of diversity is the increase of the deep-sea benthos relative to the continental shelf and abyssal plain (Hessler & Sanders, 1967; Sanders, 1968; Rex, 1981a, 1981b; see Fig. 4). Although this increase has been explained as an effect of area and thus not requiring a separate deterministic explana- tion (Abele & Walters, 1979a, 1979b), Rex (198 1a) reanalyzed available data and concluded that the additional diversity on the continental slope and rise is not area dependent. Explanations for the parabolic shape of the deep-sea diversity gradient (Fig. 4) have been ecological, and the time-stability hypothesis has been particularly influential (Grassle & Sanders, 1973; Hessler & Sanders, 1967; Sanders, 1968, 1969). desit to this model, the deep-sea en- much more predictable than that ted" fauna. Shelf faunas, on r hand, are in more stressful environ- ments and are therefore predominately “‘physi- cally controlled" (Sanders, 1968). Given time, it is hypothesized, these stable environments will accumulate many stenotopic (narrow niched) species whereas shelf environments will be char- acterized by fewer, more eurytopic species. Orig- inal formulations of the hypothesis did not con- sider the generation of diversity, only its maintenance, but Slobodkin and Sanders (1969) attempted to argue that stenotopic species in the predictable environments would have a higher probability of speciating than would eurytopes in unpredictable situations e time-stability hypothesis has not been ac- cepted by some ecologists, particularly because it is difficult to test (Peters, 1976; Abele & Wal- ters, 1979a). Yet alternative explanations have ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 also been ecological and equally difficult to test. Rex (1981a, 1981b) has adopted the “‘competi- tive equilibrium" model of Huston (1979) to ex- plain the increased diversity of deep-sea com- munities. At equilibrium, competition is assumed to eliminate some species, thus lowering diver- versities l. other ecologists have argued that communities in “competitive equilibrium” result in narrower niches and thus more species; e.g., MacArthur, 1965, 1969, 1972). Rex (1981a: 336—338) postulated that the higher diversity of predators on the slope and rise prevents com- petitive equilibrium, thus raising diversity (see also Rex, 6). As Rex (1981a: 338) noted, this hypothesis could account only for the maintenance of di- versity, so the problem still remains why there are more species at increased depth, including more predators. Not only does the hypothesis fail to address the aaa of speciation and extinction rates in a ictive manner, but whether the hypothesis is duisi to empirical evaluation also needs to be considered. How, for example, might one determine whether these deep-sea communities are at equilibrium or not? And, how might the predicted historical roles of competition and predation be evaluated in com- munities located at depths of 1,500-3,000 me- et 8 rs? The critical weakness with these ecological hy- potheses is that they do not explain gradients in speciation and extinction rates that must exist from the continental shelf to the deep-sea. As discussed earlier, a simpler hypothesis exists: namely, that continental slopes are regions of high geomorphological complexity and that this has promoted a higher rate of speciation. Wheth- er these deep-sea environments have had lower extinction rates than those of the continental shelf or abyssal plain is an open question. Predict- ability and stability do not imply that an envi- ronment will necessarily be more favorable in a physiological sense than are less stable environ- ments, although we might expect greater stability to contribute to lowering the probability of ex- tinction. The time-stability hypothesis, if it has validity or usefulness, would seem to speak only to the issue of a possible difference in extinction rates between environments of the shelf as com- pared to those of the slope. Present consider- ations suggest that the deep-sea diversity gradi- ent is better explained by a nonecological 1985] hypothesis that emphasizes variation in specia- tion and extinction rates. CONCLUSIONS Conventional interpretations of biological di- versification generally are characterized by as- sumptions that the biosphere is at or near some global equilibrium, that gradients in diversity can be explained by ecological theories of commu- nity dynamics, and therefore that speciation and extinction are diversity-dependent. This paper presents an alternative conceptualization of bi- ological diversification and its causes. Some of the main arguments underlying that conceptual- ization include: 1. The biosphere is a nonequilibrial thermo- dynamic system, open to a flux of energy and matter from outside the system. The energy-flow from the sun drives biosphere evolution through the transductive capabilities of organisms, name- ly reproduction and ontogeny. New matter enters the biosphere primarily as a result of the chem- ical breakdown of the outer surface of the litho- sphere. 2. Because the biosphere is an open system, biomass would be expected to increase through time, subject to constraints originating from out- side the system. Such constraints would include the physical and chemical properties of the earth and its history. 3. Patterns of diversity are a first-order func- tion of rates of speciation and extinction. 4. The rates of speciation and extinction are diversity-independent. The rate of speciation is hypothesized to be mediated by the rate-change of lithospheric evolution operating through geo- morphological complexity. Increased complexi- : hi — US SS SS a typ Pp long-distance dispersal to produce isolation and differentiation. A deterministic theory of extinction is for- mulated and relates an increased probability of extinction to an increase in the gradient of en- vironmental harshness. The latter is a measure of the physiological stress to which populations are exposed. . The theory of diversification proposed here is testable and has considerable empirical sup- port. Latitudinal and longitudinal gradients of diversity, in both Heo ind and marine envi- ronments, can be explained most simply by vari- ation in lithospheric seu and environ- mental harshness. The theory, as does any CRACRAFT — BIOLOGICAL DIVERSIFICATION 817 conjecture dealing with complex historical pat- terns, does not claim to explain all observations; rather, it attempts to account for the greatest amount of variance seen in large-scale diversity gradients. 6. Because the biosphere is an open thermo- dynamic system, we would not expect the Pha- nerozoic species diversity curve to show stabil- ity, except perhaps over short time periods, and then only because outside constraints (extinc- tion) are damping diversification. Present data support the hypothesis that diversity has not yet reached an upper limit. The exponential rise of the curve itself can be “reset” by mass extinc- tions or it can be “‘calibrated”’ by constraints im- posed from outside the system, e.g., by major shifts in global harshness, which might increase the gradient in background extinction. . Ecological theories proposed to explain large- scale gradients in diversity have been un- extinction. Assumed ecological correlates of high diversity (greater niche specialization and niche overlap, changes in the intensity of competition, increased predation), even if they do exist, are postulated here to be effects of that diversity, not its first-order cause. LITERATURE CITED iar L. G. & K. WALTERS. 1979a. The dg enia me hypothesis: reevaluation of the data. Amer ima sua 114: T 568. — & ——. 79b. Marine “esse diversity: a critique and alt logeogr 6: 115-126. ANDREWARTHA, H. G. & L. C. Birch. 1954. The Distribution and Abundance of Animals. Univ. Chicago ru Mer. ue — D. I. . Quaternary extinctions of large mals. DI Calif. Publ. 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BIOLOGICAL BASIS FOR ADAPTATION IN GRASSES: AN INTRODUCTION The idea of this symposium, which was held at the 34th Annual AIBS meeting in 1983, began with the question “Why have grasses become one of the most successful plant groups?" The grass family, although only the fourth largest family among flowering plants, is the most dominant and widely distributed. Members of the Poaceae, which account for an estimated 25—4596 of the world vegetation, are found in almost every hab- itat and constitute the dominant species in such biomes as grasslands, steppes, and savannas. Morphologically, grasses appear to be simple and uniform; their vegetative parts are similar and their flowers consist of little more than sta- mens and pistil. In the words of Edgar Anderson, they are “‘stream-lined.” The humble appearance of grasses is, however, deceptive, for these plants have evolved numer- ous sophisticated modifications in their wawa tive and reproductive structures at both m roscopic and microscopic ies bon ipi ttheleve — of embryo and seed and proceed l the seedling stage to maturity. Several of these spe- cializations are discussed by the contributors. Living grass genera and tribes have acquired different combinations of morphological and an- atomical characteristics. These grasses could be grouped into several adaptive lines that have ra- diated from an ancestral complex to exploit spe- cific habitats. The pooid (festucoid) line invaded r equatorial zone; andropogonoid, the tropical, seasonal-rainfall zone; chloridoid-eragrostoid, the xeric regions, mostly the dry belts of the tropics of Cancer and Capricorn; bambusoids, temper- ate and tropical forests; and oryzoids, aquatic ANN. Missouni Bor. GARD. 72: 823. 1985. habitats. There are numerous other adaptive lines such as the aristidoid, and stipoid, to name a few. Grasses have also displayed an enormous adaptability to man-made habitats for our most productive crops are the cereals such as wheat, barley, corn, and rice. It is not a coincidence that these cereals became the first staples more than 10,000 years ago. Adaptation to man-made hab- itats and high fitness in those habitats require special traits such as tolerance to oo competition in monocultural population This enormous and systematic a in grasses ought to be based on a rather flexible genetic system that can accommodate changes of different magnitudes. Chromosome number and size are quite variable in grasses and poly- ploidy is estimated by Professor Stebbins to oc- cur in 80% of the species, with hybridization and apomixis quite common. Some of the odd tribes of grasses such as Lygeeae, Nardeae, Brachye- lytreae, and Diarrheneae may have evolved from early, wide hybridization and polyploidy. This genetic system of grasses has been able to accom- modate diversification and high efficiency in traits related to fruit dispersal, seedling establishment, wind pollination, interaction and coevolution with grazing regimes, water balance, carbon dioxide and energy exchanges, and others. The next four papers will elaborate on and discuss some of these features and relate them to the success of grasses. —Khidir W. Hilu, Department of Biology, Vir- ginia Polytechnic Institute and State University, Blacksburg, Virginia 24061; and Thomas Soderstrom, Department of Botany, Smithsoni- an Institution, Washington, D.C. 20560 POLYPLOIDY, HYBRIDIZATION, AND THE INVASION OF NEW HABITATS G. LEDYARD STEBBINS! ABSTRACT Experiments are described showing that most artificial autopolyploids derived from native or in- troduced perennial grass iade Hie California are far inferior in behavior eir field conditi ions than their diploid ancestors. In a single species, Ehrharta erecta in two o autotetraploid maintained itself for, respectively, 19 and 39 years, but rem locality of planting e ny. These results, along evidence derived from several literature sources, strengthen the hypothesis that successful polyploids among natural populations are usually or almost always the result of increased videos ta accom- panying either interracial or interspecific hybridization. Th of their greater tolerance ate severe ecological or climatic conditions i is again rejected, and i which postulates secondary co changed in extent and position repeatedly during the geological periods since the initial evolution of ily. the fam The family Gramineae contains higher per- centages of species and races or cytotypes of poly- ploid origin than any other large family of an- giosperms. More than 8096 of its species have undergone polyploidy some time during their evolutionary history. Polyploidy is expressed by four different kinds of numerical series, as fol- lows: (1) Multiples of an original low basic number. Examples: Triticum, Bromus, and Festuca that include species having somatic numbers of 2n = 14, 28, 42, ... , basic number x = 7. (2) Multiples of a secondary basic number, that was itself derived from the original number by an earlier cycle of polyploidy. ere Poa scabrella complex, 2n = 42, 84, = 21 (Har- tung, 1946); B 'ochl omplex, 2n = 60, 120, 180, x = 30 (Gould, 1966); and Australian species of Danthonia, 2n = 24, 48, 72, x = 12 (Brock & Brown, 1961). (3) Multiples of a basic number that is the lowest in its genus, but was probably derived from that of preexisting genera by a cycle of poly- ploidy in the remote past. Examples: Oryza, 2n = 24, 48, x = 12 and Tripsacum, 2n = 36, 72, x = 18 (Fedorov, 1969). (4) Aneuploid series that most probably rep- resent successions of alloploids based upon dif- ferent basic numbers. Example: Stipa, 2n = 22, 24, 28, 32, 34, 36, ... 82. All of the numbers listed in Fedorov (1969) can be derived from various combinations of the basic gametic num- bers x = 5, 6, and 7. The e of these four kinds of situations indicates that polyploids in this family include, in addition to those that have originated recently during the past few thousand years, other poly- ploids that are intermediate in age from one to several million years and still other genera that acquired a secondary polyploid number during the early evolution of the family 50-70 million years ago. The original basic number for the fam- ily has been the subject of much speculation, but no convincing evidence has been presented for any of the various hypotheses. In this writer's opinion, the basic numbers x — 5, 6, and 7 are almost equally probable. They could, in fact, have all been acquired by the species complex from which the family first arose. Moreover, the pres- ence of x — 11 in Streptochaeta, a genus that is very much isolated with respect to morpholog- ical characters, and at the same time is a mosaic of very primitive characteristics along with oth- ers that are highly specialized, suggests that poly- ploidy, along with trends toward aneuploidy and a high degree of specialization with respect to some morphological characteristics, evolved quickly and soon after the family first became differentiated. Accordingly, the early stages of grass evolution must have produced many species ! Department of Genetics, University of California, Davis, California 95616. ANN. Missouni Bor. GARD. 72: 824—832. 1985. 1985] and genera that are now completely extinct. Hence in the absence of diagnostic fossils, any attempt to reconstruct the early evolution of grasses, in- cluding the first occurrences of polyploidy must be regarded as futile and self-defeating. Another distinctive feature of the Gramineae is that the high frequency of polyploidy saree throughout the family. Nearly all of the o large families of angiosperms: Liliaceae, pants daceae, Ranunculaceae, Rosaceae, Legumino- sae, Onagraceae, Umbelliferae, Scrophulariace- ae, Rubiaceae, and Compositae, include a few large genera in which polyploids are a minority of species or are totally absent. Not so in the Gramineae: its largest genus in which the ma- jority of p dsis Melica containing about 60 species; a modest number compared to such giants as Festuca (150 spp.), Panicum (500 spp.), and Paspalum (200 spp.). Because of this situation, Gramineae are not well fitted for mak- ing comparisons between related genera that dif- fer greatly from each other with respect to the frequency of polyploidy. Hence in discussing rea- sons for high frequencies of polyploidy, I shall make comparisons between genera belonging to other families. CURRENT HYPOTHESES TO EXPLAIN HIGH FREQUENCIES OF POLYPLOIDY During the past 60 years, several hypotheses have been advanced to explain different fre- quencies of polyploidy in a given flora or group of plants. The principal ones are: (1) Many poly- ploids are more resistant to extreme tempera- tures, either cold or heat, than their diploid rel- y i (2) Polyploids include a higher proportion ought resistance genotypes than do related dibloids (3) Polyploids are better adapted than their diploid relatives to invasion of new habi- tats. None of these is entirely satisfactory. The hypothesis of greater cold resistance, first proposed by Tischler (1935) and more recently supported by Lóve and Lóve (1949) and others, has been severely criticized by Favarger (1957), Gustafsson (1948), and the present writer (Steb- bins, 1950). It is contradicted by most experi- mental autopolyploids, which are less resistant to frost than are their diploid progenitors. If it were the major reason for high frequencies of polyploidy in Gramineae, we would expect to find higher frequencies of polyploidy in temper- ate and arctic genera than in tropical genera. As is evident from the high degree of polyploidy in STEBBINS—POLYPLOIDY AND HYBRIDIZATION 825 such tropical genera as Paspalum, Saccharum, and various genera of Bambuseae, this is not the case. The same criticisms can be raised against the hypothesis that polyploidy has become gen- erally and significantly more prevalent because of present or past exposure to severe drought. With respect to drought, in fact, some examples of increasing polyploidy have been associated with adaptation to more mesic habitats on the part ofthe higher polyploids. This is true of hexa- ploid as compared to tetraploid varieties of wheat, tetraploid as compared to ancestral diploid species of perennial Triticeae such as E/ymus (Psathyrostachys) junceus (Dewey, 1970), and polyploid as compared to diploid species of Bou- eh (Gould, 1966; Stebbins, 1975). Hence one can safely conclude that greater re- sistance to cold or drought is not produced by polyploidy or chron doubling per se. In IlOIC resistant than o one sofi its diploid colativer. this is probably due to the acquisition of greater re- sistance from another source, either mutation or, more probably, hybridization. With respect to both of these hypotheses, a relevant set of data is the geographical and eco- family. These are listed in Table 1. They encom- pass a wide range of taxonomic diversity, in- cluding eight tribes (Agrosteae, Andropogoneae, Aveneae, Chlorideae, Danthonieae, Eragrosteae, Festuceae, and Paniceae) and four subfamilies (Arundinoideae, Eragrostoideae, Panicoideae, and Pooideae), thus emphasizing the wide dis- tribution of high polyploidy in the family. Their geographic distribution is equally diverse. They grow in a variety of climatic zones and habitats, from tropical to polar and desert margins, al- though the largest single category of habitats is cool or warm temperate, and insular, with rela- tively little seasonal fluctuation in temperature. These data support many others in pointing to- ward the lack of correlation between distribution of polyploids and any particular kind of habitat or geographic region. The principal reasons for the high frequency of polyploids in grasses must be sought among factors other than edaphic- physiological or geographic. yp p ther fac- tors is that which postulates greater ability of polyploids to invade and colonize new or dis- turbed habitats than that of their diploid pro- genitors. This hypothesis assumes that poly- 826 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 TABLE 1. Twelve species of Gramineae having the highest levels of polyploidy. Somatic Basic Ploidy Species Number Number Level Geographic Distribution Alopecurus alpinus 130 7 18-20x Circumpolar Andropogon (Bothriochloa) barbinodis 180 10 18x SW United m n Calamagrostis crassiglumis 140 7 20x Pacific North nium j 160 10 16x South Afri Danthonia induta 120 6 20x Southwestern Australia Dupontia Fisheri 132 11 12x i chinochloa stagnina 126 9 14x Assam, India Helictotrichon (Avena) pratense 126 7 18x Britain, Scandinavia Poa litorosa 263-265 7 38x Auckland Islands, New Zealand Poa rigens 127 7 18x Iceland Saccharum robustum 194 10 18-20x New Guinea Sporobolus airoides 126 9 14x SW United States ploids can arise and maintain themselves in a variety of habitats, depending upon their genetic constitution, but only those having an guise strong capacity for colonization will becom Nissan It = advanced by Reese (1958) as an altern ion for the high frequency of s. in the flora of northern Europe and has been supported by the present author (Steb- bins, 1972), Darlington (1973), and Ehrendorfer (1980). It is, however, incomplete. It simply rais- es another question: How do successful, wide- spread polyploids acquire their greater capacity for colonization? The following sections of this contribution will be devoted to a discussion of this problem. ARTIFICIAL TETRAPLOID EHRHARTA ERECTA: LoNG TERM FIELD EXPERIMENT The most direct evidence that can be obtained on the possible effect of polyploidy in increasing evolutionary success consists of field experi- ments in which polyploids and their known dip- loid ancestors are planted simultaneously in the same area and their relative success is compared. With respect to strict autopolyploids of non-hy- brid origin, the one that I have conducted during 39 years on a South African a Ehrharta erecta, which became establis n adventive in northern California about 1930, is the most extensive and long lasting that is known to me. As stated in the account published with the rec- ord of the first few years of this experiment (Steb- bins, 1949), it was begun after screening colchi- cine induced polyploids belonging to 20 species and nine genera of grasses, the diploid progeni- tors of which all are well adapted to persistence in the natural vegetation of northern California. Of these species, only E. erecta survived the pre- liminary screening and was judged worthy of comparing the artificially induced autopolyploid with its diploid progenitor under a variety of conditions. Autopolyploids derived from all of the other species failed completely. Consequent- ly, the results of this experiment can inter- preted as indicating the maximum success that the non-hybrid autopolyploid of a grass species can achieve in a semi-natural habitat in com- parison with its diploid progenitor. The fact should be emphasized that the ex- periment to be reviewed below was not a single, isolated effort. It was, rather, the most successful of a series of attempts to establish in nature ar- tificially induced tetraploids belonging to several different and diverse genera. In all, 22 different plantings were made of dip- loid and autotetraploid Ehrharta, some of which consisted of seed sown in plots 5 x 5 m", the diploid plot being sown adjacent to that of the tetraploid, while others were started by planting well-rooted clonal divisions, in which diploids and tetraploids alternated with each other. As shown previously (Stebbins, 1949), distinctive morphological characters enabled me to follow the spread or extinction of diploids and tetra- ploids, but from time to time these results were checked by actual chromosomal counts. Sixteen of the plots were on the campus of the University of California, Berkeley, and two each in the inner Coast Ranges of Napa County, the campus of 1985] STEBBINS—POLYPLOIDY AND HYBRIDIZATION FIGURE 1. neighborhood of Plot 7, Strawberry Canyon, Berkeley 1973. At left a plant from the original diploid planting; center, a plant from about 30 m up the hill, in beaten ground under a picnic table; at right, a plant from about 100 m away, in deep shade of a planted redwood grove Three plants of Ehrharta erecta, descendants of the planting made in 1943, dug in 1972 in the and growing under u: niform conditions in a green nhouse, All of these plants are diploid; variation of this kind was not observed among the tetraploids. the University of California at Santa Cruz, and the town of Carmel, Monterey County. Although diploid E. erecta has survived and is reseeding itself in most of the plots, and from some of them has spread extensively into sur- rounding areas, the autotetraploid has failed completely except in two of them, both of them in Strawberry Canyon near Berkeley's Botanical Garden. The one described as Plot 7 in the pre- vious account (Stebbins, 1949) has been followed for 39 years. Up until 1948, the tetraploid was more successful there than was the diploid. A few years later a small building was erected just below this plot, and apparently dirt from the excavation that contained seeds of Ehrharta was thrown into a grove of oaks where no planting had been made. This condition was first noticed in 1957, and the frequency of diploids and tet- raploids was determined in the accidentally seed- a lower area in 1965. In the area as a whole, e number of established plants was: diploids b. tetraploids 230. The distribution patterns of the two cytotypes was highly distinctive: as in the originally seeded upper plot, the tetraploid plants were concentrated in that portion of the area that occupied the steepest, best drained slope, and was most heavily shaded under Quercus agrifolia. For several years after these observa- tions were made, little change was noticed, but about 1965 Ehrharta spread extensively west- ward. The plants colonized relatively diverse areas, some of them in hard packed soil Ee others in well-drained under ds (S. quoia sempervirens). In 1972 these areas were plotted and the most extreme types, including some that resembled the original tetraploids, were taken to the greenhouse for chromosome count- ing. All of them proved to be diploids. Never- theless, some of them maintained their charac- teristic growth habits under uniform conditions, as shown in Figure 1. The first plot planted in 1943 and the second pair of plots established in 1964 show similar results. In both areas, the diploid has spread ex- 828 tensively and the autotetraploid, although it still maintains itself, has remained in the type of hab- itat that exists where it was first established, and has decreased in numbers The relative lack of success of tetraploid E. erecta during the long period of the principal experiment cannot be ascribed to low seed fer- tility. Although precise records of seed fertility have not been kept, it has consistently been higher than 8096, and during the first years of the ex- periment, large numbers of seedlings were found every year near the mother plants. These experiments should not be interpreted to mean that autopolyploids are always inferior to allopolyploids as colonizers of new habitats. Those that are derived omi hybridizakon be- tween he same species, or between genetically different diploid genotypes that exhibit good combining ability may often be superior to their parents. This con- clusion is supported by the experience of agron- omists who have attempted to breed new auto- tetraploids of commercial value. The most extensive efforts in this direction have been made by Müntzing (1954) with cereal rye. He has achieved success only by intercrossing cultivars either at the original diploid level or their in- duced tetraploids, in order to increase the base of genetic diversity. Dealing chiefly with alfalfa e importance of hybridization and heterozygosity for the production of successful autopolyploids. How does this relate to the sit- uation in nature? POLYPLOIDY AMONG SPECIES THAT HAVE RECENTLY COLONIZED NEW AREAS Except for intentional attempts at introduc- tion, like that just described for Ehrharta, the most direct evidence available that bears on the problem can be obtained by comparing chro- mosome numbers of the same species or closely related species pairs, some populations of which are in their original native habitat, while others have been introduced in historical times into a new region. Recent spontaneous allopolyploidy is known for two such examples: Spartina town- sendii (S. maritima x alterniflora) (Marchant, 1966) and Tragopogon mirus plus T. miscellus (Ownbey, 1950). These examples show that new allopolyploids can often be highly successful. On the other hand, a large number of diploid species have become adventive and established in large ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Js » 42 +c AR: fF + from their original homes. In many of them, dip- loid chromosome numbers have been counted both in their original home and on the continent where they are adventive. Fingering through the chromosomal encyclopedia edited by Fedorov (1969), I ind 14 such examples in the Gramineae (Poaceae) alone. Admittedly, this evidence is weak. If autopolyploids of adventive species, originating recently, had still a much restricted geographic distribution they could easily be over- looked. Nevertheless, in some genera, such as Tragopogon, in which even the diploids have very large Qu) the increase in size that would certainly acco likely onably safe statement is that in Tragopogon species estab- lished as adventives in the New World, allo- polyploidy has been a more successful evolu- tionary change than autopolyploidy. Another kind of indirect evidence is derived from comparing the frequency of polyploidy in neighboring genera of the same family, or in dif- ferent subgenera of the same genus. If the source of polyploid series is autopolyploidy in which the change in chromosome number is the chief reason for success, no difference would be ex- pected between these closely related groups, par- ticularly if they have similar geographic and eco- logical distributions. On the other hand, if a major reason for success depends upon generating and arated populations in areas of secondary contact, then the frequency and success of polyploids in a genus or subgenus should be positively corre- lated with the number of secondary contacts that have been made during its evolutionary history. This number would obviously be greater for groups that, because of their ecological prefer- ences, tend to have a **patchy" distribution than for those that contain species that are distributed chiefly in more continuous habitats such as cli- max forests and riverine habitats. Two families that exhibit, in a dramatic fash- lon, the pattern predicted on the basis of the secondary contact hypothesis are the Salicaceae and Betulaceae. In the Salicaceae, species of Pop- ulus are mostly confined to the margins of streams and rivers, and so exhibit continuous distribu- tion patterns. All of the established species are diploid, although individual polyploid trees are not uncommon. On the other hand, the species of Salix include both riverine species and those 1985] that have patchy distributions in early stages of forest succession or on mountain slopes. Among them are many polyploids, having chromosome numbers as high as 2n = 190 (Fedorov, 1969). In the Betulaceae, the genera Carpinus, Corylus, and Ostrya grow in well-developed continuous forests, often on stream margins. All but one are iploid. Species of Betula grow in a variety of habitats, but many of them are found principally on mountain slopes and in bogs, and therefore patchy distribution patterns were likely to have been found also in the past. Of the 37 species of Betula that have been counted, 19 are polyploid or contain polyploid cytotypes, their polyploidy being well correlated with ecological preferences that would favor patchy distribution patterns. Another family that provides evidence of this kind is the Liliaceae. Among forest loving genera like Lilium, Disporum, and Trillium, polyploid series are rare. Related genera, having species adapted to more open, patchy habitats like Tu- lipa, Gagea, and Lloydia contain high percent- ages of polyploid species and cytotypes. Among the Gramineae, differences of this kind are rare, since nearly all of the larger genera con- tain high percentages of polyploids. Neverthe- less, suggestive patterns exist in the subfamily Bambusoideae, that have been carefully ana- lyzed by Soderstrom and his associates (Soder- strom, 1981; Hunziker et al., 1982). Bamboos are of particular importance in this connection, since they are confined to mesic forests or forest margins, almost entirely in the tropics, and never in climates or situations characterized by ex- tremes of drought or cold. Soderstrom s the subfamily into two groups: the woody boos, containing chiefly the tribe Bambuseae; dod the herbaceous bamboos, that resemble typical bamboos in most of their reproductive charac- teristics but are low growing, herbaceous, and confined to the understory of tropical rain for- ests. They include several tribes. Assuming that basic gametic numbers are x = 12, 11, and 10 (the probable polyploid origin of these numbers is discussed later in this article), the 108 woody species that have been counted include 86 tet- raploids and 18 hexaploids making 9696 poly- ploidy, while the 35 herbaceous species include six tetraploids, or 1796. A further point of great importance is that within the woody group the same level of polyploidy exists within entire gen- era or groups of genera, while two of the her- baceous genera include both diploids and tetra- id the Gramineae, STEBBINS—POLYPLOIDY AND HYBRIDIZATION 829 These differences are best explained by assum- ing that the polyploidy among the woody group in agreement with conclusions about woody an- giosperms in general, as compared to herbaceous groups (Stebbins, 1980; Ehrendorfer, 1980; Ra- ven, 1975). The hypothesis is that arboreal bam- boos first became differentiated in forest mar- gins, perhaps in montane regions, where patchy distributions and frequent secondary contact promoted the success a hybrid polyploids. These an ndt More recently, successful species of woody bam- boos have become incorporated into forest flo- ras, have reduced tendencies for colonization and secondary contact, and so do not exhibit poly- ploid series. Herbaceous bamboos, on the other hand, have always been adapted principally to the understory of rain forests, but from time to time have become colonizers in connection with which they have given rise to a few polyploid species within genera that still contain diploids. The implications of this hypothesis are far- reaching. First, the success of polyploid cyto- types, species, and genera is connected only in- directly to increase in chromosome number per se and is due primarily to favorable gene com- binations that they contain. As I have repeatedly emphasized elsewhere (Stebbins, 1950, 1980), polyploidy throughout the continuous spectrum that extends from auto- to allopolyploidy buffers and tends to preserve favorable gene combina- tions, particularly those in which adaptive fitness depends largely upon heterozygosity and epistat- ic gene interactions. Buffering is achieved in two different ways: tetrasomic inheritance ratios and preferential pairing of completely homologous hromosomes, i.e., those having identical or very similar pui sequences in their DNA. These mechanisms have been described in numerous publications, incha those that have been cit- ed, and need not be repeated here. Th dary tact hypothesis predicts that isolated polyploid races as well as localized au- topolyploid cytotypes should be found in many diploid or homoploid species: this prediction is in accord with presently available data. On the other hand, the difference between failure, lo- calized establishment, and widespread success depends not upon changes in chromosome num- er but upon acquisition of favorable gene com- binations. Although these may sometimes arise by mutation alone, they are much more likely to . sha d: IPE PE f e o "Oo 830 hybridization between previously isolated and adaptively differentiated populations. These populations may be interfertile races of the same species so that polyploid segregants from their hybrid would be autopolyploids. Other diploid ancestors may belong to distinct species that form sterile hybrids at the diploid level so that poly- ploid derivatives would incorporate various de- grees of allopolyploidy. Again, what counts is the adaptiveness of population environment inter- actions, not the taxonomic status of parental populations. CYCLES OF POLYPLOIDY IN THE GRAMINEAE As stated in the introduction, one reason for unusually high levels of polyploidy in Gramineae is the frequent establishment of polyploid basic numbers after an initial cycle of polyploidy fol- lowed by diploidization. This condition is best interpreted by assuming that within the same genus occur successive cycles of polyploidy, be- tween which evolution takes place at homoploid levels. The secondary contact hypothesis would predict that these cycles would coincide with pe- populations. Does the evolution of the family as e know it agree with such a prediction? Unfortunately, the fossil record is too incom- plete to provide a definite answer to this ques- tion. Nevertheless, shifts in paleoclimates, par- of the Cre d ern discussions of the phylogeny of the family, I assume that the earliest grasses inhabited mar- gins of tropical forests, then invaded open trop- ical savannas, entered still later into expanding temperate grasslands during the middle and end of the Tertiary Period, and finally occupied weedy habitats created by human m after the retreat of the Pleistocene glacier The first cycle of polyploidy s s. probably soon after the family originated in the Upper Cretaceous Period. From the widespread initial ase numbers of x = 6 and 5 arose in different groups the secondary base numbers x = 12 and 11. The following facts support retention of the hypothesis that the base numbers x = 10, 11, nd 12 are not primitive as suggested by Raven (1975), but derived by polyploidy from x = 5 and 6. First, they appear in a few distantly related ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 groups that are among the most primitive genera of their tribes, such as Danthonia and Penta- schistis in the Danthonieae, as well as Elionurus (Kammacher et al., 1973) and Sorghum in the Andropogoneae. Second, the alternative hypoth- esis proposed by Raven that x = 7, 6, and 5 are derived by aneuploid reduction from x = 12 is made improbable by the extreme rarity of x — 8 in the family as a whole, and the almost equal rarity of x = 9 except in highly specialized mem- bers of the Paniceae and Erogrosteae. The early cycles of polyploidy may well have been asso- ciated with mountain building “revolutions” that took place at the end of the Cretaceous Period and brought about many shifts in the location of forests and forest margins. The second cycle may have coincided with the opening up of warm savanna habitats during the Eocene epoch, par- ticularly in South America, where the first graz- ing mammals evolved (Andreis, 1972; Patterson & Pascual, 1968). Fragments of siliceous bodies e = pue loess deposits of South America robably remnants of Eocene grasses E 1930). Entrance into north temper- ate habitats is documented partly by the numer- ous fossil grass seeds that have been found on the plains of North America (Stebbins, 1950; Thomasson, 1978) and may well have coincided with shifts in position, expansion, and contrac- tion of temperate grasslands. These events were followed by extensive mountain building throughout the world during the Pliocene epoch creating, in the Northern Hemisphere, the con- tinental type of climate characterized by great seasonal differences in temperature to which the majority of genera belonging to the tribe Festu- ceae are adapted. New polyploids probably arose y the advances and retreats of the Pleistocene P arsa and the shifts in vegetation that accompanied them. A final cycle of poly- ploidy, affecting chiefly weedy genera like Bro- mus, Triticum (Aegilops), Hordeum, Eragrostis, Chloris, and Digitaria, took place in association with human disturbance and cultivation since the retreat ofthe glaciers and the onset of warmer climates through much of the world. In my opin- ion, Gramineae contain higher percentages of polyploids, both primary and secondary, than do other families of angiosperms because their fa- vored habitats and methods of reproduction pre- adapted them particularly well for taking advan- tage of the climatic cycles just outlined. 1985] SUMMARY AND CONCLUSIONS The percentage of species having had poly- ploid events during their evolutionary history is in the Gramineae between 80 and 9096, the high- est of any large angiosperm family. This includes both polyploid series within genera and ancient polyploidy that gave rise to high basic diploid- ized numbers for many genera. The evidence presented indicates that this high percentage ex- ists not because the grass genome is more prone than others to the occurrence of polyploid in- dividuals within populations or that raw autopolyploids are in any way superior to their tion between differentiated diploid populations have generated highly adapted, aggressive gene combinations that have been buffered and main- tained largely by the effects of polyploidy in fa- voring tetrasomic inheritance and preferential ogous, as compared to partly mparison between artificially produced antlioclvplaids and their diploid ancestors, par- ticularly an experiment of 39 years' duration with Ehrharta erecta, has shown that without hybrid- ization, autopolyploids, even when highly fertile, are greatly inferior to their diploid progenitors under field conditions or at best can maintain themselves but lack OM and the abil- ity to invade new habita 2. The hypotheses id eee by early workers that polyploids owe their success to their greater ability to withstand severe ecological conditions, such as cold and drought, have been refuted by recent data showing that polyploids are neatly or quite as conditions as in arid and cold regions. 3. Comparing genera in various families, some having high and others low frequencies of poly- ploidy, a correlation is found between high fre- quencies of polyploidy and patchy geographical or ecological distributions, which have permitted many secondary contacts between differentiated populations. he secondary contact hypothesis just pro- posed is supported also by the fact that through- out the geological periods during which the fam- ily Gramineae has evolved, habitats favored by grasses such as forest margins, savannas, tem- perate grasslands, and arctic-alpine tundra have become alternately extended and localized and have changed their geographic positions. These ° ° uvplva STEBBINS— POLYPLOIDY AND HYBRIDIZATION 831 successive habitat changes have promoted the successive cycles of polyploidy that are needed to explain both the high frequency and the high levels of polyploidy found in the family. LITERATURE CITED ANDREIS, R. R. 1972. Paleosuelos de la formacion Musters (Eocene medio), Laguna del Mate, Prov. de Chubut., Rep. Argentina. Rev. Asoc. Arg. Min- eral. Petrol. Sedimentol. 3: 91-97. BINGHAM, E. T. 80. Maximizing mp in autopolyploids. Pp. 471-490 in W. H. Lewis (ed- itor), Polyploidy, Se pid Relevance. Plenum , New York and London. Cytotax- onomy of Australian Danthonia. Austral. J. Bot. Dz ma C.D. 1 Chromosome Botany and the Origins of (una Plants, 3rd edition. Allen and Unvin, Dewey, D. R. 70. Genome relationships among diploid Elym n tetraploid and octoploid EDI species. i J. Bot. 57: 633- EHRENDORFER, F. 1980. isa and distribution. Pp. 45-60 in W. H. Lewis (editor), Polyploidy, Biological Relevanch Plenum Press, New Yor and Lo nn C. 19 Sur le pourcentage des poly- ploides dans la flore de l'étage nival des Alpes Suisses. Compt. Rend. 8th Congr. Int. Bot. (Paris) FEDOROV, A. A. (editor). 1969. Chromosome Num- s of Flowering Plants. Izdatelstvo *Nauk," Leni sa ct rn J. 1930. Particulas de silice organizada n el loess y en los limos pampeanos. Selulas si- vae de Gramineas. Anales Soc. Ci. Santa Fé 2: 66. Chromosome numbers of some d. J. Bot. 44: 1683-1696. š 1948. Polyploidy, life form and veg- etative reproduction. Hereditas 34: 1-22. HAS M. F. . Chromosome numbers in Poa, Agropyron and Elymus. Amer. J. Bot. 33: 516-532. HUNZIKER, J., T. SODERSTROM & A. F. WULFF. 1982. Chromosome numbers in the Bambusoideae (Gramineae). Brittonia a 30-35. KAMMACHER, P., G. ANOM Tes ADJANOHOUN & L. AxE-AssIL. 1973. Nom chromosomiques de Gramines de Cote-d’ Pat. Candollea 28: 191- 217. Love, A. & D. Lóvr. 1949. The geobotanical signif- icance of polyploidy. I. Polyploidy and latitude. Portugaliae Acta Biol., Sér. A, Special Vol. R. B. Goldschmidt: 273-352. MARCHANT, C. 1966. The ice M pcd and the origin of S. Townsendii. P n C. D. Darlington & W. L. Lewis eund CUL omes Today. Volume 1. Oliver and Boyd, Edinburgh Lon MüNTzING, A. 1954. An analysis of co vigour in tetraploid rye. Hereditas 40: 265- 832 OwNBEY, M. 50. Natural hybridization and am- Siue in the genus Tragopogon. Amer. J. B 37: 487-499. PATTERSON, B. & R. PASCUAL. 1968. The fossil mam- mal fauna of South America. Quart. Rev. Biol. 43: 409-451. Raven, P. H. 1975 [1976]. The bases of angiosperm phylogeny: cytology. Ann. Missouri Bot. Gard. 62: 4—764. — = 2s Polyploidie und Verbreitung. Z. Bot. 9-354 Pic T. R. 81. Some evolutionary trends in the Bambusoideae (Poaceae). Ann. Missouri Bot. Gard. 68: 15-47. STEBBINS, G. L. 49. The evolutionary significance of natural and artificial polyploids in the family Gramineae. Proc. 8th Int. Congr. Genetics (He- reditas, Suppl. Vol.), pp. 461-485. 1950. Variation and Evolution in Plants. Co- lumbia Univ. Press, New York. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 . ps The evolution of the grass family. Pp. 1-17 in V. B. Younger & C. B. McKell Poi die The Biology and Utilization of Grasses Press Inc., New York and London 75. The raid of polyploid complexes in the evolution of North American grasslands. Taxon 1980. Polyploidy in plants: unsolved prob- ms and prospects. Pp. 495-520 in W. H. Lewis (editor), Polyploidy, Biological Relevance. Ple- num Press, New York and London. rsen J. R. 1978. Epidermal patterns of the ma in some fossil and living grasses and their oray upa S Science 199: 975-977. TiscHLER, G. 1935. Die Bedeutung der Polyploidie für die Verbreitung der Angiospermen, erlautert an den Arten Schleswig-Holstein, mit Ausblicken auf andere Florengebiete. Bot. Jahrb. Syst. 67: 1- 36. ADAPTATION OF GRASSES TO WATER STRESS—LEAF ROLLING AND STOMATE DISTRIBUTION! R. E. REDMANN? ABSTRACT Leaf dimension, degree of leaf rolling Or dip and stomatal densities on adaxial and abaxial leaf ium specim imens. Representative surfaces also were e from dry habitats had narrow rolled Or folded. leaves (4 mm or 1 ns of 39 grass species from a range of dry to wet habitats mined using sagen electron microscopy. All species ess). The proportion of stomata on the abaxial surface of species from dry habitats ranged from 0 to 65%, but 56% of the species were cancentiial strongly amphistomatous. The amphistomatous leaves would pe -schéeted for habitats in which water supply and demand fluctuate widely on seasonal or diurnal time scales The evolutionary adaptation of grasses in- volves a syndrome of physiological, anatomical, and morphological characteristics. Grass leaf structure is closely coupled with major physio- logical processes such as photosynthesis, water relations, and energy balance. For example, vas- cular bundle and mesophyll cell arrangement are related to the type of photosynthetic pathway (Black et al., 1973; Hattersley & Watson, 1975), which, in turn, can influence niche separation in grasses (Monson et al., 1983). Specialized leaf tissue structure in some grasses results in leaf rolling, which strongly influences water and en- ergy balances by changing the characteristic leaf dimension and the conductance of heat, water vapor, and carbon dioxide (Ripley & Redmann, 1976). This paper concentrates on the relation- ship between leaf rolling and stomatal distribu- tion and conductance. The anatomy and mechanism of leaf rolling n grasses have been studied for over a century CEsbbirch. 1882; Shields, 1951). Loss of turgor in the bulliform cells on the adaxial (upper) sur- face generally is considered to induce rolling. Shrinkage of the adaxial subepidermal scleren- chyma and mesophyll, due to water loss, also contributes to involution; rollin n occur in leaves that lack bulliform cells (Shields, 1951). Some grasses have permanently rolled or folded leaves. Leaves of native grasses from semi-arid grass- land roll in response to increased plant water stress during dry periods (Ripley & Redmann, 976). More mesic grasses such as cereal crops also exhibit leaf rolling when exposed to water stress (Hurd, 1976; O’Toole et al., 1979). Leaves of Sorghum bicolor roll and unroll in response to diurnal changes in plant water status, provided stress is not too severe (Begg, 1 The early textbooks on plant em generally explained rolling as a xeromorphic adaptation for reducing transpiration by "protecting" the stomata, which were considered to be concen- trated on the upper surface (Warming, 1909; Weaver & Clements, 1929; McDougall, 1949). Unfortunately, references to data supporting this explanation were not given, but the idea pros ably originated with early work on ecol anatomy. Tschirch (1882) classified grasses as meadow type, with flat leaves, or steppe type, with rolling leaves. Lewton-Brain (1904) grouped British grasses into four categories representing progressively more drought resistant types: (1) leaves with flat upper surfaces, amphistomatous, (2) leaves with ribbed upper surfaces, amphisto- matous, (3) leaves with ridged upper surfaces, rolled with drying, epistomatous, and (4) leaves permanently rolled or folded, epistomatous. I was unable to find published comparative data on relative stomatal distributions on grass leaves in relation to rolling or drought resistance that would verify the generalizations describe above. Parkhurst (1978) pointed out that the lim- ited data in the literature show no clear trends l plant ! This research was supported by a Natural Science and Engineering Research Council of Canada grant. Thanks go to G. of Saskatchewan. V. L. Harms tributed valuab Shaw who did the d microscopy work using facilities in the Department of le discussion regarding drought resistance en iology, University ? Department of Crop ae Eis Plant Ecology, University of Saskatchewan, Saskatoon S7N 0W0, Canada. ANN. Missouni Bor. GARD. 72: 833-842. 1985. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 834 (SuruuoX M) usniqə8eS [4 t ea S£/S9 U UWS %# "uquog (ysind) unjooids ^y pue[sse18 ÁoÁv[.) 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INAN 19910J snonproqq L L 94 01/06 U 7] snomuidam `J 1$310jJ [£910g 9 9 94 07/08 U ‘Teog Snjpaouul `T siojd paean mo 9 SI eu Sp/SS ry “HOW Y "uquog sna42u12 "T Xueq I3AU Apues 9 8 ey O£/OL -p 7] sisuappuvo snu] q 1s310} [eə10q 9 L ed ST/SL U “Oe (UIT) wnnpotuopai `V Suruodo 1s2310J [£310g 9 L LAS | 0t/09 U 'ouyodrg (Jur) wunpunoesqns y yeqIqeH xopu] (ww) pta, dh] % eodeys səroəds souRISISoOy Ev] pərnseəjw Jeri qe/pe ?jeulojs je] 1gsnodq 'penunuo) `I #slgsv L 836 regarding the relationship of drought resistance to adaxial-abaxial stomatal distributions. Park- hurst concluded from a survey of four families that amphistomatous leaves occur most often in xeric and hydric habitats; oe he did not include grasses in his ana Absolute stomatal wee. varies widely among species and with environmental preconditioning (Muenscher, 1915; Salisbury, 1927). The relative numbers of stomata on upper and lower surfaces are not as strongly influenced by precondition- ing. Salisbury (1927) found that stomatal density on upper and lower surfaces tends to “augment or diminish in a parallel manner” depending on growth conditions. Tan and Dunn (1975) showed highly significant positive correlations between adaxial and abaxial surfaces of Bromus inermis for both stomatal frequency and length. seo and molarova 1980) ex luded that hayvial vida Was ae ‘crudely” related to the ratio of stomatal densities on the two surfaces. However, the pro- portion of carbon dioxide exchange through the adaxial and abaxial surfaces of Zea mays is re- lated to the relative number of stomata on each surface (Bertsch & Domes, 1969). For purposes of the conductance model described later in this paper it is sufficient that the ratio of stomata on each leaf surface indicates at least the trend in the lason of total conductance contributed by each surfac The first ane of the work reported here was to determine the relationships between de- gree of leaf rolling, leaf dimension, and stomatal distribution for grass species collected from na- tive habitats with different water stress regimes. The second objective was to develop a concep- tual model to help explain the advantages of dif- ferent stomatal distribution patterns and leaf rolling in wet or dry habitats METHODS The grass flora of the Prairie Provinces of west- ern Canada was surveyed in order to select species from a range of habitat types along a gradient of water availability. The species were assigned a crongat remsiançe Indes (DRI) value based on Best et al. (1971) and Hitchcock Ege in n addition to discussions with V. L. Harms, a taxonomist with extensive experience collecting in western Canada. The DRI ranged from 1 (species present in the driest hab- itat) to 10 (species present in the wettest habitat). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Specimens from the W. P. Fraser Herbarium, University of Saskatchewan, were selected for measurements of leaf width, degree of rolling or folding, and stomatal density on adaxial and abaxial surfaces. In a few cases the DRI was mod- ified slightly depending on annotation data for the particular specimen selected. Stomatal density was determined on fully-de- veloped leaves from mid-culm position, or on basal leaves, where culm-leaves were small or absent. In all cases impressions were made at the mid-blade position of the adaxial and abaxial surfaces in order to standardize the sampling. Impressions were made using m thick transparent polyvinyl chloride film. Details of the method are described in Redmann (1985). This technique was successful even with leaves having trichomes and corrugated adaxial sur- aces. The number of stomata in a minimum of ten fields randomly located on the impression was determined at 500 x magnification. In a few cases where stomatal density was small, counts were made at 125 x. The mean stomatal density per mm? for each surface was used to calculate per- centages of total number of stomata on adaxial and abaxial surfaces. Variation in percentages of the total number of stomata on the two surfaces ofleaves from five plants in a population of PAal- aris arundinacea was smaller than variation in absolute stomatal density. This was because den- sities on the surfaces increased or decreased to- gether, as was found by Salisbury (1927) and Tan and Dunn (1975). Only percentage ratios are pre- sented here in order to avoid some of the diffi- culty in comparing species with widely-differing absolute stomatal densities. Because only one or two populations of each species were sampled, variability among different populations of each species was not assessed. Even relatively large variation in percentage ratios (e.g., 96 probably would have little effect on the conclu- sions reached. Maximum flattened leaf width was determined on each herbarium specimen. The degree of roll- ing or folding on the dried specimens was noted and used to supplement information on folding, involution, or convolution reported in Best et al. (1971) and Hitchcock (1950). Leaf surfaces of eight species with widely different DRI values were examined using a Philips 505 scanning elec- tron microscope. Micrographs for three species with distinctive surface features were selected for inclusion here. 1985] 25] e 15F ° e E F E 10F e = r e o [L ° 0 = - 0 o " o u L ° = e u sr e 8 e - o ° |° O re O O O O [9] [e] [9] P o |° O - o o. OQ 0 9 r1 A ri L L | — — — T 1 2 3 4 5 6 7 8 9 10 dry wet DROUGHT RESISTANCE INDEX FicureE 1. Relationship between measured leaf dth and drought resistance index for 39 grass species from Saskatchewan and adjacent areas. Closed circles a represent flat leaves, open circles are rolling or folding leaves, and half-closed circles are permanently rolled or folded leaves. RESULTS The 39 grass species included in Table 1 occur in a range of habitats in western Canada and adjacent geographic regions. The four major tribes (Gould, 1968), Festuceae, Aveneae, Triticeae, and Stipeae each have representatives from several habitat types. The minor tribes, Meliceae and Arundineae, include mainly wetland types. Tribes with the C4 photosynthetic pathway (Waller & Lewis, 1979) were excluded because their special physiology and higher water use efficiency might have complicated the analysis. In the populations sampled, 19 species have flat leaves that either roll or fold on drying. tuca saximontana, F. scabrella, Oryzopsis hy- menoides, and Stipa richardsonii have leaves that are more or less permanently folded or rolled. The remaining 16 species have flat, non-rolling leaves. Leaf widths ranged from 1.5 mm in F. saxi- montana and Poa secunda to 25 mm in Phrag- mites communis. There was a clear relationship between drought resistance index (DRI) and leaf width (Fig. 1). No species with a DRI of 5 or lower had a maximum leaf width greater than 4 m. All four species with more or less perma- nently rolled or folded leaves had diameters or REDMANN-— WATER STRESS ON GRASSES 25} e ° ° £ El e I = e a ° e e = 8 ° 8 ° = e M s ° e ° = o o ° 9 o oo ° ° 8 = = =o s s. oe s oe oO-—-——————-—---— o n 1 L 1 1 L 10 30 40 50 60 ABAXIAL STOMATA o URE 2. Relationship between leaf width and abaxial surface for 39 di Closed circles represent flat leaves, open circles rolling or folding leaves, and half-closed circles are dashed line in n permanently rolled or folded leaves and other t LE. widths of 1 mm or less. None of the species with a DRI of 6-10 had a leaf width less than 5 mm. Neither leaf width nor stomatal distribution showed a clear relationship to tribe (Table 1). In the Stipeae, five of the eight species had stomata mainly on the adaxial surface; the exceptions were O. hymenoides, Stipa spartea, and S. viridula. The average adaxial/abaxial stomatal ratio for each tribe was: Stipeae 80/20, Festuceae 74/26, Triticeae 68/32, and Aveneae 64/36 There was no correlation of stomatal density on leaf surfaces with either DRI or leaf width (Fig. 2). The results do show that all species with leaves less than about 4.5 mm wide either had flat leaves that rolled or folded, or had per- manently rolled or folded leaves. However, the proportion of stomata on the abaxial surfaces of these leaves varied from 0 to 6596. Ten of 18 species (56%) classified as DRI 1-5 had at least 2596 of their stomates on the abaxial leaf surface. Six of 21 species with a DRI of 6-10 had 2596 or fewer stomata on the abaxial surface. The mean percentage of abaxial stomata (and coefficient of variation, c.v.) for the 18 species with DRI 1-5 was 23.996 (c.v. — 8396), and for the 21 species with DRI 6-10 was 3596 (c.v. = 5196). These 1Bum3G@BkU 26: 3 215793 AGTRA 180uym3880KkU far FIGURE 3. ANNALS OF THE MISSOURI BOTANICAL GARDEN fam d t i tL cal fate’ 1 [Vor. 72 caulum, ys —b. A. raccaium. ° ab. —c. . Á. 4 rachrauum, ab, stomate.—d. eq spartea, a ab.—f. S. s, e.—g. Poa arida, ad.—h. P surfaces. — a. aspen trachy- . S. spartea, a, ab.—1. P. arida, ad, stomate. Alisrunüng black art and white bars NE ic pu (0.1 mm in a, b, d, e, g, ra d lOuminc,f,i means were not significantly different (t-test, Steel & Torrie, 1960). Variation among species in numbers of abaxial stomata was greater in the more drought resistant group (DRI 1-5) The adaxial surfaces of species exhibiting leaf rolling were strongly ridged or corrugated. An extreme example of this is Stipa spartea (Fig. 3d). Stomata were in rows at the bottoms of the furrows between ridges. Stipa spartea had 4096 of the total stomata located on the abaxial sur- face, which was relatively smooth and encrusted with cuticular wax (Fig. 3e). Poa arida had a less strongly corrugated adaxial surface; however, the stomata on both surfaces were sunken into dis- tinctive pits, which, in the case of the adaxia surface, were d by peculiar w wax ios di — * 31). Flat- leaved species, as s represented by do pyron trachycaulum, ha surfaces similar to al bud. (Fig. 3a, b). adaxial Only one species with non-rolling epistoma- tous leaves, Calamagrostis canadensis, appeared among the 39 grasses studied (Table 1). This grass is common in moist to wet sites, sometimes shaded, in the boreal forest. Two other flat-leaved forest grasses, Bromus ciliatus and Elymus vir- ginicus, and one leaf-roller, Oryzopsis asperifo- lia, also tended strongly toward the epistomatous condition and could be grouped with C. cana- densis. All other strongly epistomatous species, including Festuca saximontana, F. scabrella, Oryzopsis pungens, Stipa curiseta, and S. rich- ardsonii have rolling or folding leaves and are found on relatively dry sites in western Canada RI 1-5 9g Flat-leaved amphistomatous species were me- sic forest- or Minn di (DRI 6-10): Bro- mus pumpellianus, Cinna VET Catabrosa aquatica, and ten other species her am- phistomatous species had collage or folding leaves. 1985] uj : T @ < ° 5 NS ap a TS z ° ` š Š ` Q9 — —, u. u. AB N ^ x e 4 4 NA A A LEAF WATER STRESS LEAF WATER STRESS ul LPN.. @ < x` 5 Ë `-. AMPHI- al Sues Él => Š EPI-T < <. u NA. eB < c u 4 — _ LEAF WATER STRESS E 4. Generalized graphs s showing relative p = abaxial surface, AD = adaxial surface.—2. istomatous | leaf. — 3. Mean conductances of. adaxial and II-)a `> = n p (EPI-) leaves. Dotted line abun poss ssible anced conductance at a particular stress ud due io leaf rolling. See text for further explanatio All those that occurred in habitats with DRI of 1-5 had rolling or folding leaves. A few species from relatively wetter habitats (DRI 6-10) also had leaves that tended to roll: Scolochloa fes- tucacea and Hierochloe odorata are found in wet- lands that can dry out in late season; Elymus canadensis and E. cinereus occur on sites with wide variation in water availability The relationship of leaf type to m may be summarized as follows: (1) Flat amphistomatous leaves—all 12 species with oe a type occurred only in relatively moist s (2) Flat, Hee MR leaves — four species with this type were hes moist, often shaded, for- est edge habita (3) Rolling or ea amphistomatous leaves— nine of 13 species in this category were from relatively dry sites. (4) Rolling or folding epistomatous leaves — five of six species came from relatively dry hab- 1tats. Permanently rolled amphistomatous leaves— the only species in this category, Oryzopsis ~ CA — REDMANN —WATER STRESS ON GRASSES 839 Rolled leaf of Elymus cinereus photo- graphed several minutes after excision from shoot. Di- ameter is 3.5 mm FIGURE 5. hymenoides, was collected in its typical sand- dune habitat. (6) Permanently rolled epistomatous leaves — all three species were from dry habitats. DISCUSSION The strong trend toward reduction in leaf di- mension in grass species from dry habitats (Fig. 1) confirms a well-known trend toward leaf re- duction in plants from dry habitats (Dauben- mire, 1974). Narrow leaves are more efficient heat exchangers and are less likely to overheat when exposed to drought and high vise ud (Parkhurst & Loucks, 1972; Gates, 1980). Le rolling reduces leaf dimension even further, re- 976). Rolling influences latent heat transfer through its effects on (1) leaf tempera- ture, and therefore on water vapor density in the leaf, and (2) boundary layer conductance. Stomatal distribution is an important factor in determining the degree to which rolling reduces leaf water vapor conductance. À simple, concep- 840 tual model was devised to help explain how leaf nductance is related to stomatal distribution, leaf rolling, and water stress regime (Fig. 4). The model presented here is meant to aid in inter- pretation of the observations of leaf character- istics. Hypotheses derived from this analysis ould be examined using a quantitative simu- lation model, and verified by field measurements of leaf conductance. The model assumes that at low water stress, conductances of the adaxial and abaxial surfaces fth flat and rolling leaves are the same. With greater water stress, rolling in- creases until complete closure is reached and adaxial conductance drops to zero (Fig. 4-1, point A). Whether rolling completely closes off the adaxial surface is uncertain; however, this seems to be the case for species that I have observed (Fig. 5). The conductance of the abaxial surface of the rolled leaf is considered to be somewhat higher than for the flat leaf, owing to the smaller dimension and thinner boundary layer ofthe for- mer. At point B, stomatal conductance is zero, and only cuticular water loss occurs from the two surfaces of the flat leaf. Point C represents the average cuticular conductance of the adaxial and abaxial surfaces of the rolled leaf. A complete average curve for the rolling amphistomatous leaf is given in Figure The adaxial surfaces of the flat and rolled ep- istomatous leaves are assumed to respond in the same way as the amphistomatous types (Fig. 4-2). The abaxial surface has only cuticular conduc- tance, which increases slightly with rolling. The average whole leaf conductance of the ep- istomatous leaf is lower than the amphistoma- tous type, except at high water stress, where they are equal (Fig. 4-3). The average conductance for the upper and lower surfaces (point C) is the same for both the epistomatous and amphistomatous rolled leaves and is lower than the average of the surfaces of flat leaves (point B). The major reason for decreasing conductance with water stress is stomatal closure. Boundary layer resistance is a relatively small component of total leaf resistance to water vapor transfer, at least in the open canopies typical of dry grass- land. Only at maximum water stresses, assuming there is a complete “seal” of the rolled leaf edges, does boundary layer resistance become large (and conductance very small). Natural selection of leaf form and function must optimize all of the following for any environ- mental regime: (1) maximum carbon dioxide ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 conductance, (2) minimum water vapor conduc- tance (when water supply is limiting), and (3) leaf temperature (Parkhurst & Loucks, 1972). Cost of constructing a particular leaf form also is a factor in the Parkhurst and Loucks analysis. A rolling-type leaf, with its extensive support tissue and other non-photosynthetic cells, thick cuticle and wax layers, trichomes, etc., must be more "costly" to produce than a simple, flat leaf. The model (Fig. 4) suggests a hypothesis that rolled epistomatous leaves with their lower total conductance are disadvantageous except in hab- itats with sustained water stress, where minimi- zation of water vapor conductance and avoid- ance of leaf overheating are important factors. Some recent data (Cohen et al., 1982) support this idea: The lowest-yielding of several slections of Festuca arundinacea had 4496 lower abaxial stomatal frequency, and 1096 higher adaxial sto- matal frequency compared with other selections in field plot studies. Flat amphistomatous leaves would be favored in habitats with lower water stress, where high carbon dioxide conductance is a benefit and water loss is less a problem. Am- phistomatous rolling leaves are a “compromise” design selected for in habitats where water avail- ability fluctuates widely on a seasonal or diurnal basis. In this way the contradictory goals of max- imizing carbon dioxide conductance and mini- mizing water conductance can be resolved. A number of studies suggest that partial rolling might actually be a mechanism to sustain rather than reduce transpiration when evaporative de- mands are high. For example, Stocker (1972) described Aristida pungens as a dune grass with rolled leaves “of highest perfection," which maintains positive photosynthesis and low tran- spiration even during extreme dry periods. This implies that leaf conductance might be higher than for an equivalent unrolled leaf. From data in Willis and Jefferies (1963) it appears that with water stress the transpiration of Ammophila ar- enaria is sustained, even though “‘inrolling” of the leaves occurs. More recently, O'Toole and Cruz (1980) speculated that leaf rolling shelters the upper surface in such a way that adaxial con- ductance is higher than abaxial in rolled leaves. Johns (1978) found that conductance of forcibly unrolled leaves of Festuca arundinacea was sim- ilar to the normally flat leaves of other species. Bennett-Clark (1935, cited in Grieve, 1955) ar- gued that xeromorphic modifications, such as leaf rolling, maintain a moister atmosphere around stomata, thus inhibiting closure and 1985] maintaining conductance. Recently much atten- tion has been paid to the direct response of sto- mata to humidity (e.g., Losch & Tenhunen, 1981). Rolling might be a mechanism to reduce this sensitivity, permitting higher stomatal conduc- tance under conditions of high evaporative de- mand. This could result in lowering of tissue water content in rolled leaves, as reported by Rychnovska and Kvet (1963). Higher stress would still result in complete stomatal closure and low conductance. This modified type of re- sponse is included in the conceptual model (Fig. 4-3, dotted lines). Of the 18 species from dry habitats that I ex- amined, only seven (3996) had predominantly epistomatous, rolled or folded leaves. The rela- tive number of species with epistomatous leaves may become larger in consistently drier habitats. When low total leaf conductance is a strong se- lective advantage, then species with epistoma- tous leaves may be selected for. This could be tested by measuring stomatal distribution in grass species from more arid habitats than those ex- amined here. LITERATURE CITED BEGG, J. E ee adaptations of leaves to water stress. Pp. 33-42 in N. C. Turner Kramer (editors), Adaptation of Plants to Water and High Temperature Stresses. John Wiley and Sons, Toronto. Bertscu, A. & W. Domes. 1969. CO,-gaswechsel am- phistomatischer Blatter. I. Der Einfluss unter- schiedlicher Stomaverteilung der beiden Blatte- mem auf den CO,-Transport. Planta 85: 183- nb F., J. LooMAN & J. B. CAMPBELL. 1971. Prai- rie grasses identified and described by vegetative characters. Agric. Canada. Publ. 1413. Informa- tion Canada, Ottawa. Black, C. C., W. H. CAMPBELL, T. M. N & DITTRICH. 1973. The v curd dens: their e evolution and comparative biology. III. Pathways of carbon metabolism related to net carbon diox- ide assimilation by monocotyledons. Quart. Rev. Biol. 48: 299-313. COHEN, C. J., D. O. CHILCOTE & R. V. FRAKES. 1982. Leaf anato 4 tall fescue ieiections differing in forage yield. Crop Sci. (Madison) 22: 704-708. . 1974. Plants and HS Gates, D. M 0. Biophysical Ecology. A ned Verlag, New Yor ys E J. 1955. The physiology of sclerophyll s. J. & Proc. Roy. Soc. Western Australia 39: n HATTERSLEY, P. W. & L. WATSON. 1975. Anatomical par rameters for predicting photosynthetic path- REDMANN —WATER STRESS ON GRASSES 841 ways of grass leaves: the ‘maximum lateral cell count’ and the ‘maximum cells distant count.’ Phytomorphology 25: 325-333. Hitcucock, A. S. 1950. Manual of the Grasses of the United States, 2nd Yes edi by Agnes Chase.) Misc. Pu c. 200: 1-1051. Hurp, E. A. 1976. Plant irte id drought resis- tance. 3 n T. T. Kozlowski (editor), 3i Water Deficits and Plant Growth. Volume 4. Ac- ademic Press, New York. Jouws, G. G. Transpirational leaf area, sto- matal and photosynthetic responses to gradually mperate herbage -125. . D. TENHUNEN. 1981. pid quiin re- echa Mans- 1949. Plant Ecology. Lea and Febiger, Philadelphia. Monson, R. K., R. O. LITTLEJOHN & G. J. WILLIAMS. 1983. Photosynthetic adaptation in 4 species from the Colorado Shortgrass steppe — a physiological model for coexistence. Oecologia 58: 43-51 MUENSCHER, W. L. C. 1915. A study of ee relation- ship of transpiration to the size and number of . Amer. J. Bot. 2: 487-504. O’TOOLE, J. C. & R. T. CRuz. 1980. Response of leaf water potential, stomatal resistance and leaf roll- ing to water stress. Pl. Physiol. (Lancaster) 65: 428-432. N. SimcH. 1979. Leaf rolling and transpiration. T Sci. Lett. 16: 111-114. PARKHURST, 1978. The adaptive significance of stomatal occurrence on one or both surfaces of leaves. J. Ecol. 66: 367—384. . Loucks. 1972. Optimal € Lv. in relation to environment. J. Ecol. 60: PosristtovA, J. &J. SOLABQUA. 1980. Environmental and b ru and abaxial pe epidermis. Photosynthet- ica 14: 80-127. REDMANN, R. E. 1985. A simple technique for mak- ing epidermal imprints using clear vinyl film. Ca- nad. J. Bot. (in press). uir E. A. & R. E. REDMANN. 1976. Grassland. p. 349-398 in J. L. Monteith (editor), Vegetation 2 the Atmosphere. Volume 2. Case Studies. Ac- ademic Press, don. RYCHNOVSKA, J. Kvet. 1963. Water relations of some psammophytes with respect to their dis- Whitehead (editors), The Water Relations of Plants. n Wiley and Sons, New York. SALISBURY, E. J. 1927. On the causes and ecological significance of stomatal frequency, with special reference to the woodland flora. Philos. Trans., 1-6 : 1951. The involution mechanism in leaves of certain xeric grasses. Phytomorphology 1: 225-241. 842 STEEL, R. G. D. & J. H. TOME. „1960. Ries and Procedures of Stati w-Hill, York. STOCKER, O. 1972. Water- jer, photosynthesis- rela- tions of desert-plants in the Mauritanian Sahara III. Small shrubs, hemi- Pe a grasses. Flora 161: 46-110. [In German.] TAN, G.-Y. & G. M. Dunn. 1975. Stomatal length, iege ncy and distribution in bromegrass. Crop i. (Madison) 15: 283-286. 1882. Beitrage zu der Anatomie und ie A. A ‘yes Wiasvialtel, Jahrb. Wiss. Bot. 13: 544-568. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 WALLER, S. S. & J. K. Lewis. 1979. Occurrence of orth and C4 photosynthetic pathways in i Man C3 American rae . Range Managem. 32: 12-28. WARMING, E. 1909. Oecology of Plants. Oxford Univ. Press, London. WEAVER, J. E. & F. E. CLEMENTS. 1929. Plant Ecol- ogy. McGraw-Hill, New York. kuy. A. J. & R. L . 1963. Investigations n the water relations of sand-dune plants under canal peera p. 168-189 in A. J. Rutter & F. H. tehead din The Water Relations of Plants. John Wiley and Sons, New York. MIOCENE FOSSIL GRASSES: POSSIBLE ADAPTATION IN REPRODUCTIVE BRACTS (LEMMA AND PALEA)! JosEPH R. THOMASSON? ABSTRACT Fossilreproductive bracts of Ee Nassella, and Pal. I ibe Sti rchaeo- d Pani leersia of the tribe Oryzeae, an rica, exam Oligocene-Miocene strata in centra | North Am with of = tribe raene (Gramineae) were collected pa late ned, a ern taxa. One objective of these studies was to elucidate ey ue esas and possible adaptive significance of bract features. Berriochloa, the oldest kno d the fossil and related living grasses probably ev sects that use grasses as food. Related living taxa appears in late Oligocene-early Miocene strata, where volved primarily as an adaptation to mammals and w adaptive mechanisms similar to those of the sho fossils, although some features such as the strongly Musso bracts and blunt callus of some species of Piptochaetium are a post Miocene developmen The Tertiary deposits of central North Amer- ican Plains have produced rich and varied floras of considerable paleobotanical interest (Stans- bury, 1852; Engelmann, 1876; Cockerell, 1914; Berry, 1928; Elias, 1931, 1932, 1934, 1935, 1942; Frye et al., 1956, 1978; Leonard, 1958; Leonard & Frye, 1978; Frye & Leonard, 1959; Segal, 1965, 1966a, 1966b, 1966c; Skinner et al., 1968; Gal- breath, 1974; Diffendal et al., 1982; Voorhies & Thomasson, 1979). Grasses in these floras are found as silicified reproductive bracts or as re- mains of leaves, stems, and roots and frequently exhibit detailed epidermal features that aid in of fossils and their modérn counterparts (Thomasson, 1976, 1977, 1978, 1979, 1980a, 1980b, 1984). Fossils described include reproductive bracts as- signed to Archaeoleersia of the t Oryzeae, Berriochloa, Nassella, and Paleoeriocoma of the tribe Stipeae, and Panicum of the tribe Paniceae and leaf fragments assigned to the subfamilies Festucoideae and Arundinoideae. This paper presents results of my studies of the reproductive bracts ou and palea) or bract (1 or cary- opsis of fossil grasses and their related living taxa. It reviews the geologic and paleoecologic back- ground, summarizes the morphologic features of the fossil and living grasses, examines evolu- tionary trends among the fossil grasses, and spec- ulates on the adaptive significance of features seen in fossil and living grasses. GEOLOGIC AND PALEOECOLOGIC BACKGROUND The late Tertiary (Oligocene-Pliocene) strata of the central part of North America are a wide- spread group of continental sediments extending from North Dakota and Wyoming to Texas and New Mexico that were deposited principally in fluvial and aeolian environments. They consist of a large variety of sediments including clays, silts, sands, conglomerates, freshwater diato- mites, and volcanic ashes and vary in thickness in individual sections from more than 200 m to 1 m. Although some early studies suggested rath- er simple models of deposition and PE , more recent investigations have dem Paid their depositional and wi hip yawa pol complexity (Bart, 1975; Breyer, 1976, 1981; Skinner et al., i Thomasson, 1979; Diffendal, Among the most richly fossiliferous Tertiary strata in the world, they contain assemblages of B ! This paper is one result of some studies that have been supported by the National Geographic Society (1883- 79, 2197-80) and the National Science and R11-8213915). I thank numerous colleagues and assistants who con Foundation (DEB-7809 150, ain adt DEB-8204681, CDP- Magd n their time and expertise in the field during ipod mos and N. Morin, P. Palmer, and anonymous pecu , for helpful boss that x sina this , iaio of I Biological Sciences, Fort Hays State University, Hays, Kansas 67601-4099. ANN. Missouni Bor. GARD. 72: 843-851. 1985. 844 TABLE 1. North America ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 Stratigraphic relationships and ages of late Tertiary strata and study sites in the plains of central Epoch North American Land (Ma) Mammal Age (Ma) Study Sites? Formation Blancan Pliocene 4 PE 5 OMEN ES Hemphillian 85 9 9. 50, 56 Ogallala Clarendonian Barstovian 56 Miocene 64, 65 Hemingfordian Sheep Creek 79, 81 Harrison-Monroe Creek — 24 — Arikareean Oligocene * Geographic localities: Kansas, west-central (9); Nebraska, west-central (50, 79), northeastern (56), north- (85). western (64, 65); Texas, northern fossil plants and animals that have been the sub- ject of numerous taxonomic and biostratigraphic studies for nearly 150 years. Some of the most paleontologically productive of these deposits are, Hollow, and Kimball) Formations. The geographic locations of several major study sites, the general stratigraphic relationships, and ages of the geologic formations and the strati- graphic levels from which fossil grasses have been studied are shown in Table 1. The stratigraphy, biostratigraphy, and lithologic characteristics of each of these formations have been studied ex- tensively, and the reader is referred to these stud- ies for further information (Harrison — Yatkola, 1978; Swinehart, 1979; Sheep Creek — Skinner et al., 1977; Ogallala — Breyer, 1976, 1981; Thom- asson, 1979; Swinehart, 1979; Diffendal, 1982). An excellent historical review of the various in- terpretations of the stratigraphic relationships of these formations is that by Galusha (1975). e vertebrate faunas found in each of the formations include abundant grazing and semi- grazing cursorial animals such as camels, horses, and rhinoceroses. This suggests savanna or sa- vanna parkland was present in the central plains of North America throughout the late Oligocene and Miocene. Based on the abundance of aeolian sediments in the late Oligocene and early Mio- cene strata, the environment during this period appears to have been arid to semi-arid (Swine- hart, 1979). However, by the middle and late Miocene the environment was more humid and probably 21 as indicated by the pre- ponderance of sediments deposited by fluvial processes and by the presence of vertebrates such as large land tortoises and crocodiles in the strata (Holman, 1971; Voorhies, 1971; Thomasson, 1979). The abundance of volcanic ash in the sed- iments indicates widespread and massive vol- canic activity during eigen of all of the for- 1., 1977; Yatkola, 1981), but the Gimatological and perhaps evo- lutionary effects of this activity have not been explored. Fossil wood with annual rings suggests that wet and dry seasons were probably present throughout the period from the late Oligocene to late Miocene, but the extremes of winter and summer present on the plains today are probably a post Miocene development. FEATURES OF GRASS ANTHOECIA In order to reasonably speculate about the adaptive significance of features seen on the bracts of fossil and living grasses it is necessary to re- 1985] THOMASSON —MIOCENE FOSSIL GRASSES 845 TaBLE 2. Husk features of Late Tertiary fossil grasses and related living taxa. Explanation of symbols: ob T e c = obconic, d = cylindrical, f = fus =o Ae en ji = “pointless” hooks, p = pric iform, k = biconvex, | = lenticular, n = ; o = absent, + = present; b = susu s = sharp; a = papillae, h = hooks, i = microhairs, m = ng, s = s Ei Interlock- Surface Silica Bodies ing Lemma Callus Structures on the Genus Shape and Palea Type of the Lemma Lemma Berriochloa* c, d, n, S, V o s h, m, p o Piptochaetium c,d,n,s, v + b,s h,m,pt o ipa d, n o S h, i, m, p o, + Nassella* f, v o b h, m o assella f,1 o b h o Paleoeriocoma* f o b h, m o ZOpsis f,n o b,s h, i, m, p o, + Panicum* k o b a o Panicum k o b a o Archaeoleersia* l + b a,1, p o Leersia l + b a,1, p + view the nature and occurrence of the features. Only those taxa represented by both living and related fossil forms will be considered, although discussions of the features are undoubtedly rel- evant to many other taxa. Summaries of the fea- tures of the fossil grasses and their related living counterparts are given in Tables 2 and 3. SHAPES OF THE ANTHOECIUM In living grasses the shape of the mature an- thoecium is extremely variable, being narrowly to broadly cylindrical or oblong in Stipa and some Piptochaetium; spindle-shaped, obovate, oblong, or nearly spheroid in Oryzopsis, Pipto- chaetium, and Nassella; biconvex in Panicum, and lenticular in Leersia and some Nassella. In ap studies the anthoecium shape was thought o be generically distinct, but as Barkworth bo 1983b) and I (Thomasson, 1976, 1979, 1980b) have shown, this is not true. For example, in the genus Piptochaetium the shape of the an- thoecium varies from elongate, cylindrical in P. avenaceum (formerly Stipa avenaceum), to ob- conic in P. uruguense, to spheroid in P. stipoides. Further, I have suggested that taxa such as S. viridula and S. robusta, which have been as- signed to the genus Stipa on the basis of their oblong to cylindrical anthoecia with pointed cal- li, probably belong in the genus Oryzopsis on the basis of micromorphological features (Thomas- son, 1981). More studies such as the one con- ducted by Barkworth (1983b) are needed to de- termine accurately generic relationships and limits in the Stipeae. Anthoecium shapes found among the fossil grasses are similar to those found in living forms. Although Stipa and Piptochaetium are not found as fossils, their common ancestor Berriochloa is present and varies in shape from cylindrical to spheroid (Parodi, 1944). Similarly the living grasses Oryzopsis and Leersia are represented in the fossil record by Paleoeriocoma and Archaeo- leersia respectively, and the shapes are nearly identical (Thomasson, 1980a, 1980b) TYPES OF CALLI The callus in grasses is an extension of the anthoecium below the base of the lemma and palea. Generally it is an extension of the lemma, 314 + 1 + 1 tc y de elon ga point- ed or blunt (Figs. 1, » Commonly the sharply pointed type is associated with an elongate, cy- lindrical or oblong anthoecium and the blunt type with a more robust, obovate or spheroid shape. Grasses assigned to Stipa usually have pointed calli whereas those assigned to Oryzopsis and Nassella have short, slightly pointed to blunt cal- li. Piptochaetium is the most variable with re- y to the callus elongate and r P d blun in n P | stipoides and P. montevidense. The various types of callus found in living grasses were also found i m fossil ue but T dietnbhiutio no om that in astaq grasses. Thus, while calli “asss: on species of fossil Paleoeriocoma and Nassella were similar to those on species in related, living Ory- zopsis and Nassella, all species of Berriochloa, regardless of the shape of the anthoecium, have 846 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 TABLE 3. Features of the husks of Miocene fossil grass genera and related living genera of the tribe Stipeae. Explanation of sy mbols: * = fossil; ** = Thomasson, 1978; ; a = well developed and indurate; b = > moderately or weakly developed and not indurate; c = well developed and prominent; d = moderately or weakly developed Raised Lateral Long Cells alls o Several Grooved Long Spheroid Times Longer Sha Genus Palea Palea Cells** Husks than Wide Callus Berriochloa* a o o c c c Piptochaetium a c c c o,c o, c ] a, b o o o o, c c,d Nassella* o o o o o o Nassella o, b o o o o o Oryzopsis a o o o o, c, d o Paleoeriocoma* a o o o d o sharply pointed calli. This suggests that the evo- lution of ihe blunt callus as cons in several species of Piz CIIL. INDURATION OF THE BRACTS In all of the living genera considered during this study, the lemma at maturity was indurate, but the palea was equally indurate in Leersia, Oryzopsis, Panicum, Piptochaetium, and Sti Nassella, and absent in most Nassella. In any case, at maturity the lemma and palea or lemma alone form a rigid container that encloses the ripe A s (Fig. 3). mong the fossils, the lemma and palea were well developed and apparently indurate (as judged y the preservation of the bracts) in all taxa ex- cept Nassella, in which the palea was absent. INTERRELATIONSHIP OF THE LEMMA AND PALEA At maturity the lemma and palea or lemma alone form a container for the caryopsis, but this happens in several different ways depending on the interrelationship of the lemma and palea. In species of Stipa and Nassella with an indurate lemma and a membranaceous palea or no palea at all, the lemma wraps around on itself and forms the rigid anthoecium. In other taxa in which both the lemma and palea are indurate and play a role in the development of the anthoecium, two conditions may occur. First, as in many species of Stipa and Oryzopsis, the two bracts may simply overlap with the lemma externally, thereby forming the anthoecium (Fig. 3). In the second condition the rigid anthoecium is formed by the physical interlocking of the lemma and palea as in Leersia, Panicum, and Piptochaetium (Figs. 4, 5). The lemma and palea may be weakly interlocked as in Panicum or strongly so as in Leersia and Piptochaetium in which the lemma interlocks into prominent grooves of the palea (Thomasson, 1976, 1980a The interrelationship of fossil lemma and pa- lea is the same as in living taxa except the grooved palea of Piptochaetium is absent in Berriochloa so that this feature apparently evolved after the Miocene. The strongly grooved and interlocked palea found in Leersia is present in the related Miocene Archaeoleersia. MICROMORPHOLOGICAL STRUCTURES OF THE LEMMA AND PALEA Types of mi hological st on lemmas and paleas of living taxa investigated include macrohairs, prickles, hooks (= barbs, crochet, or crown cells), papillae, silica bodies, and microhairs (see Metcalfe, 1960; Ellis, 1979; or Palmer and Tucker, 1981, 1983 for detailed descriptions of the different types of structures). One interesting variation of these structures is a type of hook found on Piptochaetium montevi- dense in which the point at the apex of the hook found FIGURES 1-6.— brevicalyx. section o — longate, pointed callus of Piptochaetium avenaceum. —2. Blunt callus of Piptochaetium the mature anthoecium of Stip Grori elianka of the lemma (L) and palea (P). — a comata with enclosed caryopsis (C). Note the 4. Cross section of the anthoecium of Piptochaetium THOMASSON — MIOCENE FOSSIL GRASSES ` / y" i J UC i ; P R; ww i PIO OY In / I0) mii TBA m , AO | ore bicolor. Observe that the lemma (L) interlocks with the palea (P) in a palea groove (G).—5. Palea removed from the anthoecium of Piptochaetium avenaceoid show distinct medial groove (arrows).—6. Lemma surface of Piptochaetium montevidense showing “pointless” hooks (arrows). 848 is greatly reduced or entirely absent (Fig. 6). While I refer to these structures as “pointless” hooks, I speculate later that, in combination with the anthoecium shape, they actually have an impor- tant function in protecting the caryopsis. Except for “pointless” hooks, all surface structures ob- served on living taxa were also observed on fossil taxa. EVOLUTIONARY TRENDS IN FEATURES OF THE GRASS ANTHOECIUM One of the most obvious trends that I and others (Elias, 1942) have observed in fossil grass- es is the change in anthoecium shape in the genus Berriochloa. The oldest known late Oligocene- early Miocene fossil grasses [Harrison- Monroe Creek (undifferentiated) Formation, 23 Ma] have anthoecia that are elongate cylindrical (e.g., B. schereri). These are succeeded in the middle iocene (Sheep Creek Formation, 17 Ma) pri- marily by grasses such as Berriochloa minima, that have elongate cylindrical anthoecia, al- though at least two grasses (B. primaeva and B. dawesense) from this period have more inflated, oblong anthoecia. By the late Miocene (Ogallala Formation, 8-12 Ma) two lines of grasses emerge based on the shape of the anthoecia. In one line (e.g., Berriochloa communis and B. grandis) the anthoecium is elongate and cylindrical whereas in the second (e.g., B. glabra and B. inflata) the anthoecium is inflated and obconic to nearly roid. A second trend among fossil grasses is an in- crease in overall size ofthe anthoecia in the genus Berriochloa. Species found in older strata of the Harrison and Sheep Creek Pec e (e.g., Ber- riochloa schereri and B. minima) have anthoecia that are predominantly — iien to oblong and small (length 1.5-5 mm and width 0.4-1.4 mm), whereas in much younger strata of the Ogallala Formation species (e.g., B. grandis and B. com- munis) with similarly shaped, but larger anthoe- cia (length 3-12 mm and width 0.7-2.5 mm) are common. Likewise, the only species with inflat- ed, obovate to fusiform anthoecia that occur in older strata ( Berriochloa primaeva and B. dawes- ense in the Sheep Creek Formation) have small anthoecia (length 1.1—4.5 mm and width 0.6-1.4 mm), whereas there are several species (e.g., B. maxima, B. glabra, B. tuberculata, and others) with very robust, fusiform to spheroid anthoecia (length 2.5-8 mm and width 2.2-3.5 mm) that ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 are found only in the much younger Ogallala Formation. A final trend among the fossil grasses is an increasing variety of taxa from older to younger strata. This is true even though thousands of specimens from each stratigraphic level are known. In the oldest levels only the genus Ber- riochloa is known and is represented by two species, whereas in the younger strata ofthe Sheep Creek Formation as many as seven species of Berriochloa are known. In the late Miocene Ogal- lala Formation an explosion in the numbers of taxa of fossil grasses occurs, with as many as 20 species of Berriochloa being known, in addition to species of Archaeoleersia, Nassella, Panicum, and Paleoeriocoma. Although exact explanations for the trends among fossil grasses are unknown, it is possible to speculate on their causes and adaptive signif- icance I believe that evolution of the two lines of anthoecium shapes (elongate, cylindrical and in- flated, fusiform to spheroid) appears primarily to represent a response to the coevolution of grasses with animals and insects that ate the caryopsis as food (see section later in this paper on the adaptive significances of anthoecium fea- tures). This suggests that the shape of the an- thoecium has limited taxonomic utility (i.e., un- related taxa may have similarly shaped anthoecia), and this finding has been supported by my studies, at least at the generic level. To illustrate, when the first fossil grasses of the tribe Stipeae were recovered from the High Plains Ter- tiary strata two taxa, Stipidium and Berriochloa, were described primarily on the basis of the an- thoecium shape, the former having cylindrical anthoecia and the latter having inflated anthoe- cia (Elias, 1932). However, I have since shown that anthoecium Shape alone cannot be used in the fossil and modern Stipeae, but rather that a combination of micromorphological and macromorphologi- cal features of the anthoecium must be used (Thomasson, 1979, 1981) (Table 3) I speculate that increasing size of anthoecia is a long term response to foraging insects such as harvester ants. It is well known that these insects collect large numbers ofgrass anthoecia (Wheeler & Wheeler, 1963; Davison, 1982), and that they select seeds according to size (i.e., the smaller the body length of the forager the smaller the seeds collected) (Davison, 1982). An increasingly larg- er anthoecium might have reduced foraging on 1985] the caryopsis of species with large anthoecia and thus favored those species with increased sur- vival of caryopses, each of which could poten- tially germinate and produce a new plant. inally, with respect to the increasing variety of taxa, several explanations are possible includ- ing increased immigration of new taxa, more complex environments with more diverse eco- logical niches, or more favorable conditions for preservation (e.g., more alkaline groundwaters resulting in more silica in solution). In the genus Berriochloa the increasing variety of species might reflect increasingly diverse responses of grasses to their coevolution with foraging animals and insects. ADAPTIVE SIGNIFICANCE OF ANTHOECIUM FEATURES An important goal of my studies of Tertiary grasses and their living counterparts has been to discern the adaptive significance of the anthoe- cium features. Based on my studies and obser- vations and those of others, it is possible to speculate on the adaptive significance of grass anthoecium features: 1. Rigid indurate anthoecium. a) Protection from damage to the caryopsis during ingestion by mammals. Evidence from recent and fossil fecal deposits and from anthoecia associated with skeletal re- mains (Parodi, 1944; Voorhies, 1974; Thomasson, 1976; Voorhies & Thomas- son, 197 b) Protection from insects that feed the car- yopsis to larvae. Evidence from both mod- ern and fossil burrows (Wheeler & Wheel- er, 1963; Davison, 1982; Thomasson, 1982). Pointed callus. a) Aids in dispersal of anthoecium with en- closed caryopsis through attachment to the hide or entanglement in the hair of mam- mals (Parodi, 1944; unpubl. data). b) Burial of the anthoecium and caryopsis in the soil (Barkworth, 1983a; unpubl. data). . Blunt callus. The absence of a sharply pointed callus may facilitate rapid movement of the anthoecium through the digestive tracts of mammals. . Cylindrical anthoecium. The narrow, elon- gate anthoecium facilitates burial in cracks in the soil (Barkworth, 1983a; unpubl. data). p W + THOMASSON —MIOCENE FOSSIL GRASSES 849 5. Inflated anthoecium. a) In combination with the blunt callus, the inflated shape allows for more rapid movement of the anthoecium through the digestive tracks of mammals (Parodi, b) The inflated shape protects against use by many insects that have great difficulty in moving all other similarly shaped dissem- inules (Davison, 1982). 6. Surface structures. a) Pointed surface structures such as hooks may aid in the entanglement of the an- thoecium in the hair of animals. rge pointed feat h especially on the callus, aid in the burial of the anthoecium by making it difficult for the anthoecium to come free once lodged in cracks in the soil (Barkworth, 19832; unpubl. data). Blunt, raised structures such as papillae and “pointless” hooks, especially in com- bination with inflated anthoecia with blunt calli, facilitate rapid movement of the an- thoecium through the digestive tracts of mammals. . Grooved palea. May act as a mechanism to securely lock the lemma and palea together for increased protection of the caryopsis dur- ing passage of the anthoecium through the digestive tracts of mammals. iz <~ O ~ N The degree to which anthoecium features are in- terrelated is particularly interesting. For exam- ple, the anthoecium of the South American grass Piptochaetium montevidense has a combination of features that include an indurate, obovate an- thoecium formed by a strongly interlocking lem- ma and palea, a short, blunt callus, a lemma surface covered with large “pointless” hooks, and a readily deciduous awn. Rather than asking what the function of any single feature is, the question that I think must be answered is: What is the function of the combination of characteristics observed? For example, common sense suggests that the rigid, indurate anthoecium probably functions in protecting the caryopsis from some- thing. But from what? In order to answer that question and others concerning the adaptive sig- nificance of anthoecium features, I have exam- ined evidences from both the past and present, and I have concluded that the principal factor acting on the evolution of many of the features seen on the fossil and living grasses is the co- 850 evolution of the grasses with animals and insects that ingest (purposely or inadvertently) the cary- opsis. In arriving at this conclusion all of the features of the anthoecium have been consid- ered. Thus, in the case of P. montevidense the features observed should protect the caryopsis by aiding the rapid movement ofthe anthoecium through an animal's digestive tract (rounded shape, rough surface, and blunt callus) while pre- venting damage by digestive juices (indurate, in- terlocking bracts). Features seen on a grass such as Stipa comata, however, would allow for easy burial of the anthoecia in cracks in the soil (cy- lindrical anthoecium with a hairy, sharp callus and persistent awn) while providing protection to the caryopsis from foraging insects and mam- mals (indurate, overlapping bracts) Although factors of the environment such as mesic and xeric habitats probably have influenced the evo- lution of certain features of the anthoecium (e.g., the sharp callus) the features have not been in- fluenced to the degree suggested by Barkworth (1983a) CONCLUSION Many features of grasses that have allowed them to be phenomenally successful in compe- tition with other plants, animals, and insects have a long geologic history. The results of my studies of fossil and living grasses have suggested pos- sible explanations of the adaptive significance of many features of grass anthoecia and have pro- vided a better understanding of the evolution of grasses. Whereas most of the fossil history of grasses and the actual origins of many grass fea- tures are unknown, the widespread distribution of Cretaceous and Tertiary deposits containing abundant floras provide unlimited opportunities for future exploratory studies on the intriguing question of grass evolution. LITERATURE CITED BARKWORTH, M. E. 1983a. Patterns of evolution in niv. Wyoming Contr. Geol. 14: 27-39. 1928. Fossil nutlets of the genus Lith- ospermum. Proc. U.S. Natl. Mus. 73(2734): 1-3. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 1976. The d of Ogallala sed- a. Univ. Michigan Pap BREYER, J. iments in western Nebra Paleontol. 12: 1- jn The Kimballi ian land-mammal age: ene, tekel upharsin (Dan. 5:25). J. Pa- eee "55: 1207-1216. COcKERELL, T. D. A. 1914. Two new plants Mes bs Tertiary rocks of the west. Torreya 14(8): DAVISON, E. A. 1982. Seed utilization by harvester ants. Pp. 1-6 in R. F. Buckley (editor), Ant-Plant Interactions in Australia. Dr. W. Junk, The Hague, . 1980. The Rush Creek-Lisco structural basin, Garden and Morrill Cou Nebraska. Trans. Nebraska A ounty, Nebraska ni adjacent areas. Bull. Geol. Soc. Amer. 93: 964-976. . PABIAN & J. R. THOMASSON. 1982. Geo logic history of Ash Hollow Park Nebraska. Circ. Ed. Nebraska Univ. Conservation Surv. Div. 5: 1-31. ELiAs, M. K. 1931. The geology of Wallace County, Kansas. Bull. Kansas Univ. Geol. Surv. 18: 1-254. 1932. Grasses and other plants from the Ter- tiary rocks of Kansas and Colorado. Univ. Kansas Sci. Bull. 20: 333-367 . 1934. Zones of fossil herbs in the later Ter- tiary of the High Plains. Proc. Geol. Soc. Amer. 1934: 332. [Abstract.] . Tertiary grasses and other prairie vege- pde: from the High Plains of North America. mer. J. Sci. 29: 24- 1942. Tertiary prairie grasses and other herbs from the Pigs Plains. Special Pap. Geol. Soc. Amer : 1-17 ELLIS, R. P. A procedure for SR ai comparative leaf anatomy in the Poaceae. II. epidermis as seen in surface view. Bothalia 12: 1-671. ENGELMANN, H. 1876. Appendix I. Report on the geology of the country between Fort Leavenworth, ley. Pp. 243-335 in J. H. Simpson, Report of Ex- plorations Across th i i of Utah for a Direct Wagon-Route from Camp Floyd to Geneva, in Carson Valley, in 1859. Gov- ernment Printing Office, Washington FRYE, J. C. & A. B. LEONARD. 1959. Co rrelation of the Ogallala Formation (Neogene) in western Tex- as with type pns in a . Rep. Invest. Bur. Econ. Geo s 39: p H. E Cae 1978. Late Cenoz oic sediments, molluscan faunas, and clay minerals in northeastern New Mexico. Circ. hc Mexico Bur. Mines Mineral Resources 160: 1-32. & A. SWINEFORD. "uon Stratigra- phy ofthe Ogallala Formation (Neogene) of north- ern Kansas. Bull. Kansas Univ. Geol. Surv. 118: GALBREATH, E. C. 1974. Stipid grass “seeds” from the Oligocene and Miocene deposits of northeast- ern Colorado. Trans. Illinois State Acad. Sci. 76: 366-368. 1985] GALUSHA, T. 1975. Stratigraphy of the Box Butte formation. Bull. Amer. Mus. Nat. Hist. 156: 1- 36 HOoLMAN, J. A. 1971. Climatic significance of giant tortoises from the Wood Mountain Formation pper Miocene) of Saskatchewan. Canad. J. Earth Sci. 8: 1148-1151. LEONARD, A. b the Pliocen Sci. Bull. 38: | 1393-1403. & . 1978. Paleontology of Ogallala Fo rmation, northeastern New Mexico. Circ. New . Mines Mineral Resources 161: 1-21. 960. Anatomy of the Monocoty- ledons. I. Gramineae. Clarendon Press, Oxford. PALMER, P. G. & A. E. TUCKER. A scanning electron microscope — 2 of the epidermis of East African G Contrib. Bot. 49: Two new lege plants from s. Univ. Kansas 1983. A scanning electron micro- epidermis of East African 1944. vision de las gramineas aus- tral americanas del “seso Piptochaetium. Revis- ta Mus. La Plata, Secc. Bot. 6: 213-310. SEGAL, R. H. New fossil fruit (Compositae) from the Pliocene of western Kansas. Amer. Midl. Naturalist 73: 430-432. Taxonomic study of the fossil species of the genus Cryptantha (Boraginaceae). Southw Naturalist 11: 205-210. review of some Tertiary endocarps of Celtis (Ulmaceae). Southw. Naturalist 11: 211- 216. 1966c. Biorbia (Boraginaceae) in the central U.S. Pliocene. Univ. Kansas Sci. Bull. 46: 495- 508 SKINNER, M. F., S. M. SKINNER & R. J. Goonis. 1968. Cenozoic rocks and faunas of Turtle Butte, south- central South Dakota. Bull. Amer. Mus. Nat. Hist. 138: 379-436. & 1977. Stratigraphy and biostratigraphy oflate Cenozoic deposits in central Sioux County, western — a. Bull. Amer Nat. Hist. 158: 265-37 STANSBURY, H. 52. ain and Survey of the Valley of the Great e: aene sd Utah, Including u Lemay e Through the Rocky he ounta ins. coit ais & Co., Philadelph d pese J. B., II. 1979. Cenozoic Geology of the Plat tte River Valley, Morrill and Garden on url Nebraska. M.S. Thesis. Univ. of Ne- braska, Lincoln. THOMASSON, J.R. 1976. Tertiary Grasses and Other Angiosperms from Kansas, Nebraska, and Colo- THOMASSON — MIOCENE FOSSIL GRASSES 851 rado: Relationships to UM Taxa. Ph.D. Thesis. Iowa State Univ., 1977. Late ena grasses, basi and hackberries from southwestern Nebraska. Univ. aps Contr. Geol. 16: 39-43. 978. Epidermal patterns of the lemma in some fossil and living grasses and their phyloge- netic significance. Science 199: 975-977 1979. Late Cenozoic grasses and other an- osperms s from Kansas, Nebraska and Colorado: biostratigraphy and relationships to living taxa. Bull. Ka Univ. Geol. Surv. 218: 1-68. 1980a. Archaeoleersia nebraskensis gen. et sp. nov. (Gramineae — Oryzeae), a new fossil grass from the late Tertiary of Nebraska. Amer. J. Bot. 67: 876— "o 1 b. Nisem. aw “sawna Stipeae) from the Mioce d Nebr phylogenetic significa 9 Mic Pal oTe a Sti- pa robusta and Stipa viridula (Gramineae: Sti- peae): her ae significance. Southw. Naturalist 26: 211-214. n Fossil grass vee and other plant fossils from arthropod burrows in the Miocene of western Nebraska. J. Pale pehar 56: 1011-1017. l ji sp. n., Cyperocarpus elia- j nd C the Miocene of Nebraska. Amer. J. Bot. 70: 435- 449 . 1 . Miocene grass (Gramineae: Arundi- d leaves showing external micromorpho- logical and internal anatomical features. Bot. Gaz. (Crawfordsville) 145: 9. Vooruies, M. R 71; Pied aii significance of crocodilian remains from the Ogallala red (up- per Tertiary) in northeastern Nebraska. J. Paleon- tol. 45: 119-121. . 1974. Fossil pocket mouse burrows in Ne- braska. Amer. Midl. Naturalist 91: 492-497. 1981. Ancient ashfall creates a de cue of prehistoric animals. Natl. Geogr. Mag. 159: 66 75. . R. THOMASSON. 1979. Fossil grass an- thoecia within Miocene rhinoceros skeletons: di- rect evidence of diet in an extinct species. Science 206: 331- m G. J. WHEELER. 1963. The Ants of rth Dakota. Univ. of North Dakota Press, Grand etn YATKOLA, D. A. 1978. Tertiary stratigraphy of the Niobrara River Valley, Marsland quadrangle, western Nebraska. Nebraska Geol. Surv. (Pap.) 19: 1-66 GRAMINOID RESPONSES TO GRAZING BY LARGE HERBIVORES: ADAPTATIONS, EXAPTATIONS, AND INTERACTING PROCESSES! MICHAEL B. COUGHENOUR? ABSTRACT problem of d adaptive significance to traits that enable graminoids to tolerate or evade The ungulate herbivory is exam grazing selection pressures, pm constituting grazing exaptations rather than ned. Some of these traits may have originally evolved in ced non - to true adap s. The fossil record indicates Saal ped habitats, extensive grasslands, and grazers appeared, “interacted, and evolved together. How effects for grazed plants. Other traits, such as developmental vena enhance competitive ability in certain environments, but also increase grazing tolerance or resis oes xpe modeling showed that defoliation responses are embedded including photosynthesis, transpiration, nutrient uptake, a aptations to defoliation must be interprete ents and simulation ina pier non-herbivorous selection pressures, they may subsequently have been selected, combined, or am- ic “awa aa usb , polymorp plastic individuals, or communities of species that evade, resist, or tolerate herbivo Graminoid grazing tolerance and the nearly simultaneous increase of grasses and grazers in the fossil record (Stebbins, 1981) suggest that grasses are adapted to herbivory, perhaps as a result of coevolution. Grasses and herbivores may, consequently, be somewhat mutualistic (Owen & Wiegert, 1981). Graminoid tolerance of herbivory due to continued elongation from the base of the leaf and lateral growth through tillering following defoliation have long been known (Lisle, 1757; Arber, 1934). Interacting physiological and morphological characteristics ofgraminoids that contribute to such grazing tol- erance have been identified (McNaughton, 1979a) and analyzed (Coughenour et al., 1984b, 1985a). However, the origin of present traits in organ- isms cannot always be attributed to selection pressures of their present en t (Gould & Lewontin, 1979). The current beneficial effect of a trait is actually incidental or secondary if the trait frequency increased in response to another previous selection pressure. Thus, we must dis- tinguish between beneficial traits (aptations) that are only incidentally beneficial (exaptations), and traits that have actually resulted from selection pressures to confer their present benefits (adap- tations) (Gould & Vrba, 1982). We should con- sider alternative non-adaptive hypotheses in any investigation of the presumed adaptive signifi- cance of traits (Eriksson et al., 1983). Evolutionary constraints reduce the variety of evolutionary options available to a species. A trait may be non-adaptive in the sense that it is a necessary consequence of another trait (Eriks- son et al., 1983). More fundamentally, individual traits are not in themselves the targets of natural selection. It is the fitness of the whole organism, or interacting system of traits, that determines the outcome of natural selection (Mayr, 1983). This “cohesion of the genotype" results in in- evitable compromises among competing de- mands and a residue of sub-optimal parts that are necessary for the working of the whole (Mayr, 1983). Most plants are in essence metapopula- tions (White, 1979) of repeating modular units (Harper, 1981). The effect of herbivory on mod- ular differs from the effect of predation on non-modular organisms in that modular or- ganisms survive predation through vegetative re- production, whereas non-modular organisms do not. Therefore, it is necessary to consider defense against herbivory in a context of other interacting Factorial experimentation and simulation mod- ' I thank Sam McNaughton for an entree into this field via an extended post-doctoral fellowship on his Serengeti project. He and Linda Wallace were gg Soran for most of the experimental results. Interactions with them and with Roger Ruess were invalua Foundation DEB-77 20350 and DEB-79 ? Biol 291 to . This work was funded by grants from the National Science . J. McNaughton ogical Research Lab, Syracuse University, 130 College Place, Syracuse, New York 13210. Present address: 523. Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, Colorado 80 ANN. Missouni Bor. GARD. 72: 852-863. 1985. 1985] eling are used here to examine interactive pro- cesses, causal networks, and possible competing demands that are relevant to herbivory response in perennial graminoids. EvoLUTIONARY HISTORY OF GRASSES AND GRAZERS The original selection pressures that gave graminoids their current morphological and physiological characteristics can never be abso- lutely known. However, some insight can be gained by examination of the origins of grasses in relation to grazers in evolutionary time and space. These origins must be related as well to pre-grazing ancestors to determine possible pre- adaptiveness of traits, and to non-grazing selec- tion pressures such as climate, which may have given rise to grazing exaptations. Monocotyledons as a broad group are evolu- tionarily more advanced than dicotyledons, ex- hibiting neotony as an evolutionary response to extreme environmental conditions, particularly to an aquatic habitat (Takhtajan, 1969). Linear, parallel-veined leaves that grow from their bases arose from modified bladeless petioles, and the sheathing bases of monocot leaves are a conse- ce of the single cotyledonary structure (Cronquist, 1968). The first monocotyledons probably arose in the Cretaceous period (135- 70 Ma) (Raven & Axelrod, 1974). By the tertiary period (70 Ma) most ancestral grasses had differentiated. Predominantly xero- phytic types developed to the south into Africa (Jacques-Felix, 1962). However, there are basi- cally three contrasting theories (Whyte, 1974) re- garding the origin of primitive grasses: (1) origin in tma humid tropical ee with increasing rrelated with great- er adaptation to aridity nen 1929); (2) origin in open forest followed by migration into xero- phytic zones (Valentine, 1970); (3) origins in me- sophytic and xerophytic zones with subsequent mixing of types (Aubreville, 1962; Thomas, 1966). A typical primitive grass may have been a small, low-growing perennial, somewhat tuft- ed, with short leaves, and probably forest-loving (Stebbins, 1972). Early grasses and hypsodont-toothed mam- mals first appeared in the fossil record in early Eocene (60 Ma) strata of South America. During the Oligocene (40-25 Ma) several new groups of notungulates appeared in the first open savan- nahs in South America. In the Miocene (25-12 COUGHENOUR —GRAMINOID RESPONSES TO GRAZING 853 Ma) horses with more refined hyposodont teeth appeared, and predominated the large herbivore fauna of North America (Stebbins, 1981). Semi- arid conditions appeared in South America 20 million years earlier than in North America, but the greater continentality of the North American climate caused more rapid changes in environ- mental selection pressures (Stebbins, 1981). The Rocky Mountain uplift, and resulting aridity of the Great Plains caused a retreat of forests in late Oligocene and Miocene (30-12 Ma) leaving drought-tolerant grasses and herbs behind (Dix, 1964). Close-grazing bison and sheep arrived in North America from Eurasia in the Pleistocene (3-0.01 Ma) (Mack & Thompson, 1982; Steb- bins, 1981). Equidae and Camelidae migrated from North to South America at the end of the Pliocene (3 Ma) or early Pleistocene (Stebbins, 1981). Existence of grass savannahs in Africa begin- ning in the Eocene (60—40 Ma) is suggested by the sparse fossil record of this period (van der Hammen, 1983). Poaceous grasses may have en- tered South America from Africa during the Pa- leocene (70-60 Ma) or even uppermost Creta- ceous (Stebbins, 1981). In Africa, the Eocene-Miocene (60-12 Ma) was dominated by perissodactyls and proboscideans. Increasing continental elevation was accom- panied by increased aridity in the Oligocene (40- 25 Ma) (Raven & Axelrod, 1974). Grazing, rumi- nating bovids appeared first in fossils of the late Miocene (18-12 Ma) in Eurasia and radiated greatly during the Pliocene (12-3 Ma), but they did not invade Africa until late Pliocene (Sin- clair, 1983). In the late Pliocene-Pleistocene (4— 0.01 Ma) the climate fluctuated in Africa, causing fluctuations in vegetation types and expansion- contraction cycles of savanna habitats. As a re- sult, there was great adaptive radiation in the African large herbivore fauna (Morton, 1972). There may have been significant evolutionary development of grass species at this time (Love, 1972) Thus, grasses evolved at an early date and ex- isted for quite some time prior to abundant graz- ing animals. Increasing specialization and ad- aptations to aridity occurred among grasses subsequently. Extensive grasslands, grazers, and semiarid climates appeared almost simulta- neously. This simultaneity makes it difficult to ascertain the original adaptive values of traits that enhance survival under both aridity and grazing. 854 ADAPTATIONS FOR DROUGHT OR GRAZING? A number of traits that allow graminoids to evade, resist, or tolerate drought provide similar benefits to plants subjected to grazing. These traits are basal meristems, small stature, high shoot density, deciduous shoots, belowground nutrient reserves, ane rapid transpiration and growth inear t romanintercalary mer- istem at the base of the leaf is an ancient trait of monocotyledons. This trait probably made it more feasible for some monocots to maintain intercalary meristems in a protected position at the base of the plant during vegetative growth. Basal meristems, protected by the Dasa sheath in grasses, may be better able to with ght (Barlow et al., 1980). Many perennial grasses have reduced bud height, which is regarded as a means to semiarid habitats “probably had already de- veloped enough of a basal intercalary meristem to enable leaves to recover from grazing” (Steb- bins, 1972). Small stature is a well known characteristic of many graminoids that are adapted to more arid habitats. Water and time both impose an upper limit on individual shoot size. In general, how- ever, there is an inverse relationship between shoot size and shoot number (G sponse to ephemeral water availability, distrib- utes the risk of drought-induced mortality, and increases the rate of recovery from defoliation. A shorter growth form is also less likely to have a large proportion of its leaves removed by large herbivores. Deciduous shoots allow perennial graminoids to survive drought by reduction of both tran- spiring leaf area and shoot maintenance respi- ration. However, senescent leaves are low quality food items that limit herbivore population sizes (Sinclair, 1975) and, therefore, herbivore im- pacts. Drought-deciduous leaves are relatively inexpensive to construct compared with leaves of drought-enduring plants (Orians & Solbrig, 1977; Mooney & Gulmon, 1982), and therefore herbivore damage is less costly to the plant. Ex- portation of nutrients back to perenniating or- gans during senescence (Williams, 1955) further reduces the impact of herbivory (Mattson, 1980). Perennial grasses endure the dry season with carbon and nutrient reserves stored in sub-sur- face crowns and roots. When moisture returns, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 regrowth proceeds at a rate in excess of current photosynthesis, dependent on upward translo- cation from reserves. This adaptation to a non- growing season is critical for regrowth following defoliation (Trlica, 1977). Drought-deciduous perennials that are active only in a wet season should be under strong se- lection pressure to evolve morphological and physiological traits to maximize production when water | ble (Orians & Solbrig, 1977). Whereas drought-tolerant shrubs use water slow- ly when wet and continue to transpire when dry, grasses use water rapidly when wet and cease transpiration when dry. Rapid transpiration im- } d growth, which help plants recover from defoliation. plies rapid [ ADAPTATIONS FOR COMPETITION OR GRAZING? In a competitive environment it is adaptive for a plant to have higher plasticity of root-shoot allocation to enable rapid exploitation of light, or soil water, or nutrient resources, as they be- come available (Grime, 1977). Rapid shoot re- growth and associated diversion of photosyn- thate from roots to shoots enables rapid response to changing resource availability. For example, a sudden reduction ofleafarea causes a shift from water to light limitation, thus requiring rapid di- version of resources to shoots The most primitive grasses were small, and the most general trend in grass evolution has been to increase in size, through increased activ- ity of intercalary meristems (Stebbins, 1972). Through sequential activation of the meristems of stem internodes, the meristems of leaf blades are elevated on culmed shoots. This provides a more effective display of leaf area than does a basal origin of all leaf blades, thereby increasing light competitive ability. Increasing lignification and canopy heights associated wit pothesized to be inherently less resistant to her- bivory than more rhizomatous or stoloniferous growth forms, possibly due to coevolution of the latter with large herbivores (Mack & Thompson, 982). However, the way plant modules are ar- ranged relative to one another may have adap- tive significance distinct from herbivory toler- ance. Vegetative reproduction by tillering is an efficient method of colonizing open sites. Exten- -— 1985] sive asexual reproduction also yasin the probability Vl £ ook, 1979) A plant with short horizontal internodes and til- lers packed in a bunch or tussock grows as an advancing front or “phalanx,” forming an im- penetrable zone of resource preemption. Plants with vigorous horizontal spreading or tillering, on the other hand, have the “guerilla-like” tactic of spreading over an unoccupied area quickly (Harper, 1981). The guerilla growth form is adaptive for plants that compete for space in early successional stages or disturbance sites. Once established, however, protection against invaders is more advantageous. The guerilla- phalanx dichotomy involves a trade-off of re- source investment inherent in lateral versus hor- izontal growth Several other traits may allow rapid coloni- zation of open space and increased competitive ability in early successional environments (Ba- zazz, 1979). These include rapid growth rate, non-light-saturated photosynthetic rate, rapid transpiration and photosynthesis, and reduced sensitivity to decreased water availability. These traits may enable better recovery from defolia- tion as well. DEFENSIVE CHEMISTRY-DOES IT, SHOULD IT, OR CAN IT OCCUR IN GRASSES? Lignin and silica both reduce palatability; however, it is likely that their primary functions in grasses are non-defensive. Silica reduces di- gestibility in some grasses, increases it in others, invariably wears down herbivore teeth, and can cause kidney stones (van Soest, 1982). Silica con- centrations are sometimes higher in more heavi- ly grazed plants, genotypes or species (Mc- Naughton & Tarrants, 1983; Brizuela, 1985), but this might be taken as evidence either for or against defensive utility. Defoliation induces el- evated silica levels in individual plants of some grasses (McNaughton & Tarrants, 1983) but not of others (Cid, 1985). Although invertebrate her- bivory is deterred by silica, deterrence of rumi- nant herbivory has not been demonstrated. Lig- nin is indigestible, and it can reduce the digestibility of associated carbohydrates (van Soest, 1982). However, both silica and lignin im- part rigidity to the cell wall and prevent wilting when cell water contents decrease, as during mild droughts. Xerophytic plants generally have thicker cell walls and are more lignified (Sinnott, COUGHENOUR —GRAMINOID RESPONSES TO GRAZING 855 1960). Lignin (Higuchi, 1981) and silica Pung: 1973) enable terrestrial plants to develop upri forms and be resistant to environmental stress and disease. Thus, grasses that grew in drier, more open habitats evolved firmer and more silica- ceous leaves, which probably prompted the evo- lution of hypsodont dentition among grazers (Stebbins, 1981). Non-lignin phenolic compounds could affect ruminant herbivory of grasses. Tannins and fla- vones have been isolated from East African grasses (Field, 1976); nevertheless, these species are important forage for native ruminants. Ter- penoids occur in another African grass, Cyrn- bopogon excavatus (Field, 1976). Regrowth is grazed but mature plants are avoided. Flavo- noids and coumarins also occur (Wong, 1973). These d have not been shown to deter ruminant grazing. Grasses lack the variety of toxic chemicals that deter herbivory in dicotyledons. Alkaloids have been isolated from only 21 grasses out of 8,00 examined (Culvenor, 1973). The only alkaloid- bearing grasses that have been shown to affect ruminants, however, are Phalaris tuberosa, Loli- um perenne (Culvenor, 1973), and Festuca arun- dinacea (Bush et al., 1970). Cyanogenic com- pounds were detected in only two of 16 grasses (Gibbs, 1974); however, these have not been shown to affect ruminants. Ruminal microflora are able to degrade anum- +1 they ber o uic reach the stomach. It has been | hypothesized that pregastric fermentation originally evolved be- cause it could detoxify secondary compounds in browse plants (van Soest, 1982). Although some tropical grasses contain oxalates that inhibit nu- trient absorption by the non-ruminant digestive system of horses (Blaney et al., 1981), the mi- croflora of ruminants degrade the oxalate before it reaches the stomach (Allison et al., 1977). Ru- men microflora of domestic sheep have been shown to detoxify alkaloids (Dick et al., 1963) and toxic isoflavones (Nilsson et al., 1967). Sev- eral wild ruminants commonly feed on the poi- sonous fruit of the East African herb Solanum incanum L. (Pratt & Gwynne, 1977). Thus, some chemical constituents do, and therefore can, defend grasses from herbivory, at least to some extent. However, the effectiveness of chemical defenses is limited by the ability of rumen microflora to degrade them. Although lig- nin and silica have non-defensive functions, they may reduce palatability enough to decrease se- 856 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 TABLE l. Summary of clipping effects on growth and photosynthesis of a short sedge, a mid-grass, and a tall-grass subjected to factorial variations in clipping height (cm) and frequency (d). Compared to unclipped controls, there were stimulations (+), inhibitions (—), and no differences (0). K. nervosa T. triandra H. filipendula Height of plant 10-25 2, 5 cm/2, 7 d Clipping regimes Aboveground growth Total ES Tillering = a Leaf elongation rate + Residual leaf area — +4 Belowground growth Root growth — 0° Crown growth — pd Photosynthetic rate 4—e Stomatal conductance +—e Internal conductance 0 Water status +e 50-120 100-150 3, 6 cm/3, 7 d 10, 15 cm/7, 14 d 0 0-* + + = 0 = 0 — +f + 0 + — +f + 0 +" a Inhibited in 14 d, no effect in 7 d clipping treatments. » Short-duration measurement mid-way through experiment. * [nhibited in 2 d, 2 cm treatments. 4 Inhibited in 2 cm, stimulated in 5 cm treatments. * Stimulated in 2 cm, inhibited in 5 cm treatments. f Inhibited in S d, stimulated in 7 d treatments. g Soil water c rvation. ^ Plant water yon m lection pressure for other defensive compounds. The benefits of chemical defenses are further re- duced by other, morphological or i aa nate traits that enable deterrence, tolerance, or eva sion of herbivory. INTERACTIVE PROCESSES AND MORPHOMETRIC TRAITS IN THREE SERENGETI GRAMINOIDS The responses of grasses to defoliation are , 1983; Wallace oughenour et al., 1985a, 1985b) elucidated this causal network. The short-, mid-, and tall-statured species represented a range of habitat aridity in the Serengeti ecosystem of northern Tanzania Qualitatively different responses to defoliation were exhibited by the three species (Table 1). Total aboveground growth, residual leaf area, and tillering were stimulated by clipping only in the short-statured species. However, leaf elongation rate was enhanced in all three species. In turn, clipped yield was significantly correlated with leaf elongation rate in all species. Positive photo- synthetic responses to defoliation, mediated by stomatal and internal leaf conductances, com- pensated for reduced leaf area consistently in the tall-statured species and variably in the short- and mid-statured species. This prevented nega- tive belowground growth responses in the tallest species. Belowground growth was l residual leaf area and tiller numberin all species. In general, the tallest species had the most ho- meostatic response, but the underlying adjust- ments were the most complex. Water status of the short and tall graminoids was improved by defoliation. Water additions affected growth in both species and photosyn- thesis of the latter, suggesting that responses to defoliation were partially mediated by water. Variations in habitat aridity and light com- petition exert differential selection pressures upon plant growth form that subsequently influence defoliation responses. The inherent tendencies of shorter-statured species to allocate shoot growth in the horizontal direction via tillering, and for taller-statured species to allocate in the vertical direction via leaf elongation, largely influenced d wi th COUGHENOUR—GRAMINOID RESPONSES TO GRAZING 1985] A. š BELOWGROUND Ë PRODUCTION Š pe REUS ei i air e o N x N = \ o N ( | ABOVEGROUND ` $ | STIMULATION ` ° a N \ B. IO p CARBON RELATIVE EFFECT ON GROWTH GRAZING INTENSITY FIGURE 1.—A. The relationship between above- grazing intensity on the simulated plants. E DE in the model. (Adapted from Coughe- nour et al., a.) which of these mechanisms was the more im- portant defoliation response. Similarly, shorter species had inherently greater root/shoot and trient uptake and crown biomass on nutrient storage and meristem production. Additional experimentation with Ky/linga nervosa has revealed that plant nitrogen relations and grazing responses are highly interactive. Ni- trate and ammonium had differential effects on plant responses to clipping, e clipping stim- ulated uptake rates of both ions (Ruess et al., 1983). Urea nutrition d material flows to grazers and to plant reproduction, while clip- ping and urea together resulted in stimulated leaf and flower production, thus establishing a pos- itive feedback from grazers to plants to grazers Ruess & McNaughton, 1984). Furthermore, grazing affected the balance of several other plant I400r 2 IOOOF * SHORT GRASS o M GRASS L GRASS TOTAL PHOTOSYNTHESIS gC/m o o o d L n L L L n L 20 40 60 80 100 [4 LIVE SHOOT BIOMASS gC/m FiGURE2. Total photosynthesis over a single grow- ing season versus "-— ve shoot biomass of sim- i tall-grasses. Each point xd resents a different pii regime. (Data fro Coughenour et al., 1985a.) nutrients via clipping and nitrogen source (Ruess, 1984) MECHANISTIC SIMULATION ANALYSES OF CLIPPING RESPONSES A simulation model was constructed (Coughe- nour et al., 1984a) that incorporated morpho- logical and physiological aspects of graminoid o physiological processes that controlled carbon, water, and nitrogen uptake rates and allocations within the plant. The model simulated meristem types and numbers, tissues types and masses, and canopy geometry. Simulation experiments were erformed with short-, mid-, and tall-grasses subjected to various clipping heights and fre- quencies (Coughenour et al., 1984b; Coughe- nour, 1984) Response surfaces of yield to grazers (clipped yield) versus grazing height and frequency indi- cated that shorter graminoids were optimally de- foliated at greater frequencies than taller gram- inoids, primarily because a large number of small shoots reach their maximum sizes more rapidly than a small number of large shoots. Maximum aboveground productivities were achieved at in- termediate grazing intensities (Fig. 1A) when aboveground growth was maximally stimulated. Belowground growth was depressed little at low to moderate grazing intensities but declined rap- idly at higher grazing intensities. Greater pho- tosynthetic rates pc Dd ons in be- lowgroun veground growth response to grazing kafa was unimodal pri- oo CA oo ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 N E O @ SHORT o LOOF 5$ W > A TALL C D UNGRAZED > = 150r O = e © a IOOF e = 5 a 50r C z O J a L L L L L _ ey u 200 400 600 800 lOO0O 1200 1400 2 puede ¿ka Tu 9c /m FIGURE 3. Total bel 7 1 f 154m nada season. These are the same simulations as in Figure 2. marily because relative water availability in- creased while relative carbon availability de- creased as grazing intensity increased. Total available photosynthate decreased as more was removed by defoliation, but total transpiration and rainfall interception decreased. Total photosynthate production was maximal at intermediate levels of live shoot biomass (Fig. 2). The increase in photosynthesis was greatest at low levels of shoot biomass, indicating the critical dependence on leaf area at higher grazing intensities. At higher shoot biomass levels an increase in shoots had little or no impact on total photosynthesis because photosynthetic rate per unit leaf area decreased. Total photosynthesis was maximized at lower shoot biomass levels in shorter grasses. Photosynthetic rates of grazed plants were stimulated by grazing. Light and water limitations on photosynthesis were lessened as greater quantities of water and light were avail- able to each unit of leaf area of grazed plants. Shoot morphology affected the relationship between total photosynthesis and BeoNEround 3). The tall-grass exhibited a non-linear map because more photosynthate was allocated to J E [24 e eo shoots at low levels of total photosynthesis. The tall-grass produced the lowest quantity of be- lowground growth with a given quantity of pho- tosynthate because it diverted a greater fraction of photosynthate to fewer, taller, and more lig- nified shoots. Clipped mid- and tall-grasses had much less belowground growth per unit of pho- tosynthate than corresponding unclipped plants, while clipping did not have this effect on the short-grass. The effect of shoot hol as tested with the model by interchanging shoot morphologies of short- and tall-grasses (Table 2). All param- eters regulating shoot growth, including shoot number, type, and size, tissue types, and growth rates, were reversed while the remaining biotic and abiotic parameters were held constant. The shoot system of the short-grass had at least twice as many meristems; however, tall-grass maxi- mum individual shoot growth rates were several times higher, and the shoots could grow roughly three to four times larger. Placing a tall-grass shoot system on the short-grass decreased above- and belowground productivities in all grazed plants. Only above- ground production of ungrazed plants was in- 1985] COUGHENOUR —GRAMINOID RESPONSES TO GRAZING 859 TABLE 2. Effect of reversing shoot morphology between short- and tall-grasses on above- and belowground production (g/m?). Aboveground Production Belowground Production Grazing Experiment Ungrazed Moderate Severe Ungrazed Moderate Severe Control 35 116 97 140 112 62 Short-grass with tall-grass shoots 46 84 24 118 94 7 Control 214 512 90 301 158 8 Tall-grass with short-grass shoots 196 419 509 353 324 82 creased. The interchange was especially harmful to severely grazed plants. Aboveground produc- tion of the ungrazed tall-grass was depressed by the short-grass shoot morphology, but be- lowground production was increased because less photosynthate was required by the smaller shoots. (Of course, the experiment did not account for reduced light competitive ability of shorter shoots.) When severely grazed, the tall-grass with short — cana considerably better. Because shorter grasses had Lgs densities of E ster tillers, panei i removed proportionately less ao. and foliage below the grazing height was more concentrated. Grazing stimulated tillering and converted sparse, tall canopies into shorter ones with denser con- centrations of younger, high quality foliage in th the model and in reality (McNaughton, 1976, 1984). Modeling has explicitly shown that grazing re- sponses are the result of interactions between e and that the traits, the processes, and the inter- actions are, conversely, affected by grazing. ADAPTATIONS IN INDIVIDUALS, POPULATIONS, SPECIES AND COMMUNITIES The importance of plant form in grazing re- sponse mas been — both through ex- odeling. Shorter grasses had greater densities s smaller tillers, so defoliation at a given height removed proportionately less foliage, and foliage below the grazing height was more concentrated. However, morphological shifts to grazing-resistant forms can occur within individuals, populations, or communities. Graz- ing stimulates tillering, and convertes a sparse, tall canopy into a shorter “grazing lawn" that has a denser concentration of younger, higher quality forage (McNaughton, 1984). Grazing lawns can result from phenotypic plasticity. This was sim- ulated by the model, and shown in controlled experimentation discussed above. Phenotypic responses to grazing are reversed when grazing ceases (Quinn & Miller, 1967) Another type of grazing lawn can be induced through long-term repeated grazing. Prostrate or dwarfed growth forms, often with a high ratio of basal to culmed meristems, may exist as infre- quent ecotypes in ungrazed polymorphic popu- lations. Recurrent, localized grazing may then increase the predominance of grazing-resistant ecotypes in the local sub-populations (Stapledon, 1928; McNaughton, 1979b, 1984; Detling & Painter, 1983; Detling et al., 1985), and the pop- ulation as a whole would then be a mosaic of genetic variability. Alternatively, somatic mu- tations during asexual reproduction may facili- tate evolution of genets and mosaics of genetic variability (Whitam & Slobodchikoff, 1981). hus, a third type of grazing lawn could be com- dum of plants that are ° inherently shorter, but 1984). * Finally, by reduci leaf area, opening up the canopy, creating small disturbance sites, and generally reducing the survival of grazing-intol- erant species, grazing may alter competitive re- lationships in plant communities (Watkin & Clements, 1978; Edroma, 1981). In a grazed plant community lateral resource investment is more important, and shorter species may be favored where taller species would otherwise be com- petitively superior. Thus, it is necessary to distinguish the impacts of grazing on individual plants from impacts on localized shifts in the ia ssim of ecotypes in a population, or s n a community. Since phenotypic dies. is “heritable (Bradshaw, 1974) and since polymo m can be induced through disruptive selection, mm could include examples of true grazing adaptiveness. However, 860 one must show that the grazing-resistant forms of plastic individuals are usually or only induced by grazing, that saa with grazing- re- where there iS, Or was periodic grazing, or ae Suus ecotypes com- prise more heavily grazed patches of a mosaic. In the case of non-plastic individuals or non- polymorphic populations, a trait can clearly be attributed to grazing selection pressure only if it is always more predominant, or if it only exists in reproductively isolated populations that are, or were, regularly grazed. m CONCLUSIONS Grasses and large grazing herbivores evolved together. However, it is difficult to show which graminoid traits arose or are solely maintained by grazing. Part of the difficulty, of course, is that we are unable to determine from the fossil record the precise origin of graminoid traits in relation to herbivore evolution. Extant grasses do with- stand, avoid, or deter grazing, yet this does not constitute sufficient evidence for the selection of traits through coevolution with herbivores. I have given a number of examples of possible exap- tations, or traits that could have arisen through non-herbivorous selection pressures, but which nevertheless confer benefits to grazed plants. Either adaptive or exaptive traits may be more frequent among grazed (or ungrazed in the case of deterrence or avoidance) populations or species. If such a trait originally arose through non-herbivorous selection, but is now main- tained in the gene pool of a population by graz- ing, or becomes more apparent in patches of grazed plants through differential survival, the trait should be considered truly adaptive. Simi- larly, phenotypic plasticity that is maintained by grazing is adaptive irrespective of original selec- tion pressure. Grasses sel markedly from dicotyledons i in their lack of chemical d High concentrations of lignin and silica deter herbivory of grasses, but their primary functions are probably structural support and drought re- sistance. Ruminant microorganisms may make chemical defenses ineffective, and the array of other non-chemical means of coping with or avoiding herbivory may make c chemi unnecessary or non-cost-effect It is well worth specula inp in view of the novelty and potential importance of the concept, to admit that moderate grazing could have pos- t herbivory. defenses ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 itive long-term effects on grasses (Owen & Wie- gert, 1983). Perennial grasses persist for hundreds of years by vegetative reproduction, and mod- erate grazing can increase vegetative reproduc- tion (McNaughton, 1983a). Thus, grazing can stimulate fitness. If so, natural selection would tend to encourage moderate herbivory of grasses. The lack of chemical deterrents and the high de- gree of adaptation or exaptation to herbivory among grasses support this possibility. 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CLARK2 ABSTRACT Chusquea pohlii, C. latifolia, and C. serpens are described from Central and South America. Ob- servations on bud morphology and branch development i in Chusquea are included. Bud morphology provides a new set of vegetative characters invaluable i In separating species of Chusquea. The bud a species, including those described pane is defined and contrasted with era usque. pi seep iss branching, typical of the majority of species in the genu Among the woody bamboos, Chusquea Kunth has long been recognized as a large and diverse genus. McClure (1973) and Calderón and So- derstrom (1980) included over 90 species within the genus, but according to my observations the count is nearer 120 species. Early descriptions of Chusquea species (e.g., Nees von Esenbeck, 1835; Munro, Puai Hackel, 1903; Pilger, 1905) fo- cused primarily on spikelet morphology. More rece drm McClure (1973) and Soderstrom and Calderón (19782, 1978b) recognized the impor- tance of vegetative features in distinguishing among species in Chusquea, but id i E variation of bud and branch complem phology in the genus has received little ene study. My work has shown that the bud com- plement provides a new set of vegetative char- acters invaluable in separating species, and that there is greater variability in patterns of branch development than has been previously attributed to Chusquea. The bud complement in Chusquea, according to McClure (1973: 69), consists of “separate pri- mary buds of two size categories in constellate insertion, the smaller ones usually many (rarely only 2)." Although this fundamental bud mor- phology is shared by all members of the genus, the number and arrangement of the buds are highly variable and seem to be species-specific. Two representative bud complements are illus- trated in Figure 1. The sheath scar is the line of attachment of the culm leaf. The nodal region is delimited below by the sheath scar (nodal line), and above by the supranodal ridge, which may or may not be prominent. The largest, or dom- inant bud is the central bud, and the smaller subtending buds are called subsidiary buds (McClure, 1973; Soderstrom & Calderón, 1978a, 1978b). In the chusqueoid bud complement, all the buds are of the same (primary) order. The term "subsidiary" is used here in the sense of McClure (1973) to indicate only that the buds so described are smaller than the central bud. As defined by Foster and Gifford (1974), subsidiary buds also could be called collateral buds. The dimorphism between the central and subsidiary buds in Chusquea is striking and sets this genus apart from all other bamboos. Two types of central buds are found in Chus- quea. The triangular type, exemplified by C. ser- rulata Kunth (Fig. 1A), is very common and like those of other bamboos. The bud is erect and roughly triangular in outline, and the winged pro- phyllum is easy to distinguish. The circular type of central bud, as seen in C. /iebmannii Fournier (Fig. 1B), is more or less circular in outline, pro- jecting outward horizontally from the culm. The prophyllum is atypical, apparently lacking wings. I have observed circular central buds in only eight or nine species of Chusquea and not in any other bamboo genera. Extravaginal branching, in which the emerging branches break through the base of the culm sheath proper (Fig. 2A), is typical of Chusquea ! Field work was supported te NSF Grant DEB-8 102766. I thank Richard W. Pohl for sharing with me his extensive knowledge of the gras s. I am especially grateful to Thomas R. Soderstrom and Cleo of the Smithsonian Institution for Pie me to the bamboos. Alice R. Tangerin é E. Calderón ni provided technical advice regarding the illustrations. I thank Dr. Nels R. Lersten of Iowa State University for critically reviewing the — ? Department of Botany, Iowa State University, Ames, Iowa 50011. ANN. Missouni Bor. GARD. 72: 864-873. 1985. 1985] d, \l Jt) (t ut EN A i mn. RA (o D^ i i W n J We MI fum OY NU M Y ( M X M no CLARK — CHUSQUEA 865 C Representative bud complements of Chusquea. — A. C. serrulata, mid-culm bud complement FIGURE 1. showing triangular paise bud.— B. Scales — Cb— ss— sheath scar, w— wing. (Soderstrom & Calderón, 1978b). The girdle, a horizontal band of elastic, usually persistent tis- at the base of the culm sheath in some bamboos (McClure, 1973), usually is not well developed in species of Chusquea that exhibit extravaginal branching. However, I have ob- served in several species of Chusquea, including the three species described in the present paper, that the girdle is prominent, forming a some- times asymmetrical band of tissue up to 1.5 cm wide at the base of the culm sheath. The girdle usually is persistent, even after the culm leaf sep- arates from it and falls away. In contrast to ex- travaginal branching, here the developing sub- sidiary branches emerge through the girdle without rupturing the culm sheath (Figs. 3F, 4E). Occasionally, as the branches mature, the culm sheath becomes ruptured to a certain extent, par- ticularly if its overlapping edges are fused at the base (Fig. 3F). I am designating this pattern of branch development “‘infravaginal,” a term sug- gested to me by Cleofé Calderón. It is clear that this type of branching is derived from the extra- vaginal pattern by means of the expansion of the girdle, but the distinction between extravaginal C. liebmannii, mid-culm bud comple ntral bud, k— $ed m — margin, p— prophyllum, sb— subsidiary bud, sr — supranodal ridge, ment showing circular central bud. and infravaginal branching is taxonomically use- ful and a parallel, descriptive term is necessary. Infravaginal branching is known in only one oth- bamboo genus, the Asiatic Dinochloa Büse (Dransfield, 1981). In both Chusquea and Dino- chloa, Unies! branching is correlated with a climbing, viny it The first of the is species described here is named in honor of Dr. Richard W. Pohl of Iowa State University, who has devoted much of his career to studying the grasses of Costa Rica. It is fitting that this Costa Rican endemic com- memorate Dr. Pohl's many years of field work in that country and his long-standing interest in the bamboos. Notes of the late Dr. F. A. Mc- Clure, filed in the Hitchcock and Chase Grass Library of the Smithsonian Institution (U.S. Na- tional Herbarium), show that he considered the second of these three species to be distinct, based on the presence of just two subsidiary buds sub- tending the central one. Because of the unusual branch complement, he tentatively proposed that this species be included in a new genus. It is evident that his notes and descriptions were at best preliminary; he used at least three different 866 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 ia FiGURE 2.— A. Extravaginal branching in Chusquea serrulata, scale = 1 cm (based on Calderón et al. 2989). B-F. C. pohlii. — B. Inflorescence, scale = 1 cm.— C. Spikelet, scale = 1 mm.— D. Bud complement, scale = 1 cm.—E. Leafy branch, scale = 1 cm.—F. Culm leaf, scale = 1 cm (B-C based on Croat 36232; D-F based on Clark & Clark 275). 1985] provisional names (of which I chose the most descriptive) for the same entity. Three agrostol- ogists (J. R. Swallen, T. R. Soderstrom, and R. W. Pohl) recognized, apparently independently, that the third species described here was distinct, but the provisional names used would be invalid and no adequate description was ever provided. pi š p pohlii Clark, sp. nov. TYPE: Costa sé: along the Hwy ud ae 44—45, closer to Km 45, 2,190 m, 27 Feb. 1982, Clark & Clark 275 (ho- lotype, ISC; isotype, US). Figure 2B-F. Culmi scandentes, arcuati, usque ad 3 cm diam., m longi. Folia culmorum at panies lon- gior quam nodus s E uens iue 21 -47.6 c uu p con- na gior quam lamina, externae hispidae, internae sparsim hispidae; laminae persistentes, triangulares, apicibus acuminatis, 2.5-8.4 cm longae, scabrae; picem asymmetricum, gerr non nisi prope gem Nodi ad culmum medium gemma centralis cn rotundata subtenta a Q4 91 2)g emmis parvioribus lanceolatae, apicibus acuminatis, basibus attentuatis- rotundatis, 14-25. id m longae 3 cm latae, = dibae inferne, inferme tes- 30 cm longa; rhachis plus minusve complanata, pub- erula; rami Wisi loben puberuli. Spiculae lan- ceolatae, 6.8-9 mm longae, leviter falcatae. Glumae be iquemiotitus 0.5-0.6 mm longa, enervis; glu- ma II squamiformis, 0.8 mm longa, enervis. Lemmata sterilia 2: lemma sterile I triangulatum, apiculatum, e; lemma sterile II lanceo- latum, apiculatum, 4.1-5. à Lemma fertile lanceolatum, apiculatum, 6.6-7.6 m longum, 7-nerve. Palea lan caries bicarinata, ane lata, 6.4—7.7 mm longa, 4-nerv Culms scandent, to 3 cm diam., to 15 m long, arching. Jnternodes terete, often shallowly sul- cate above the central bud on more mature culms, solid, 22-36 cm long, scabrous-hispid below node. Culm leaves persistent, usually extending past the next node, 21—47.6 cm long, chartaceous to cartilaginous; sheaths triangular, fused for 2.3 cm above the base, 18.5-39.2 cm long, 5.4-10.5 cm wide at base, usually 5.5-7.5 times, occa- sionally only 3-4 times, as long as the blade, abaxially hispid but hairs deciduous with age so the sheath often seems scabrous only, adaxially shiny and sparsely hispid toward the apex or sometimes the whole upper half hispid, the mar- gins glabrous, the base often densely pubescent; CLARK— CHUSQUEA 867 blades persistent, triangular, 2.5-8.4 cm long, abaxially scabrous, adaxially glabrous and shiny, the apex acuminate, the margins glabrous; girdle asymmetrically developed, prominent only in the region of the bud complement where the sheath scar dips markedly forming a flap through which the lowermost subsidiary ! hes emerge; inner ligule a short, stiff rim, 1-2 mm long; outer ligule lacking. Nodes only slightly swollen, at mid-culm with one circular central bud subtended by (2-)4- 9(-12) smaller subequal subsidiary buds. Branching infravaginal, central bud rarely de- veloping but sometimes rebranching when de- veloped; lowermost subsidiary buds developing first and emerging through the girdle, the upper ones developing shortly thereafter and emerging through the sheath, forming leafy branches 24- 39 cm long, which do pot pom Foliage leaves 4—6 per ate, hispid but often glabrous just below the pseudopetiole and toward the margins, the overlapping margins cil- iate; blades linear-lanceolate, 14-25.6 cm long, 1-2.3 cm wide, L: W = 9:16, the adaxial surface scabrid and not tessellate, the abaxial surface gla- brous and weakly tessellate, the apex acuminate, the margins serrulate, the base attenuate-round- ed; pseudopetiole more or less distinct, 2-5 mm long; outer ligule a conspicuous, stiff rim 0.5-2 mm long; inner ligule elongate, asymmetrical, (7-)11-30 mm long, chartaceous. Inflorescence a narrow panicle 21-30 cm long; rachis puber- ulent, one side slightly rounded, the other ridged; branches appressed, arising only from the ridged side of the rachis, angular, 5-6 cm long at the base of the panicle, puberulent; pedicels 1-4 mm long, angular, pubescent. Spikelets lanceolate, 6.8—9 mm long, slightly falcate. Glumes 2; glume I scale-like, 0.5—0.6 mm long, glabrous, nerves lacking; glume II scale-like, 0.8 mm long, api- cally obtuse, abaxially pubescent, nerves lacking. Sterile lemmas 2; sterile lemma I triangular, 3.1— 4.5 mm long, apiculate, abaxially pubescent on the upper half, adaxially pubescent just below the apes; marginally ciliate toward apex, 5-ne II lanceolate, 4.1—5.1 mm long. apiculate: abaxially pubescent on the upper half, adaxially pubescent just below apex, mar- ginally ciliate toward apex, 5-nerved. Fertile lemma lanceolate, 6.6-7.6 mm long, apiculate, abaxially scabrous-pubescent on the upper half, adaxially pubescent just below apex, marginall ciliate toward apex, 7-nerved. Palea lanceolate, 2-keeled, sulcate only toward the apex, 6.4-7. mm long, apiculate, abaxially scabrous-pubes- 868 cent between the keels, otherwise scabrid toward apex, 4-nerved. Lodicules 3; 1.5-1.9 mm long, the posterior lodicule narrower than the anterior ones, all ciliolate on the upper margins. Stamens 3; anthers linear, 3.3-3.9 mm long. Ovary gla- brous. Fruit unknown. Representative specimens examined. COSTA RICA. ALAJUELA: Km 15-16, N of San Ramón, Pohl & Clark 14115 (CR, ISC). cARTAGO: Tapantí Hydroelectric Re- serve trail along Río Dos Amigos, 23 June 1976 (fl), Croat 36232 (MO, US); Alto Patillos, NE of Tapantí, along road to Turrialba, Poh! 14139 (CR, ISC); at hy- droelectric dam, canyon of Río Grande de Orosi, S of Tapantí, Pohl & Selva 12886 (ISC, MO). HEREDIA: Vol- cano Barba, wet submontane forest, at end of road, Booth 161 (USy; Route 9, S of the Vara Blanca inter- section, between Km 28 & 29, Clark & Clark 277 (ISC, US); Alto del Roble, Río Las Vueltas, ca. 10 km NNE of Heredia, Pohl & Gabel 13676 (ISC). PUNTARENAS: onteverde, along forest road in forest preserve, Pohl & Pinette 13246 (F, ISC, MO). sAN JosÉ: along road N of Cascajal, N of Río Cascajal, Pohl 14101 (CR, ISO): arque Nacional Braulio Carillo, S boundary, 2 km of Bajo de Hondura, Pohl & Clark 14104 (CR, ISC). This bamboo has solid culms and multiple in- dependent buds at each node, thus placing it in Chusquea, where it is distinct enough to merit specific recognition. Pohl (1980, 1983) has re- ferred to this species as the “Tapantí population" of Chusquea and Chusquea *'hispidissima," re- spectively. Chusquea pohlii is distinguished by its rather thick, coarse, clambering culms, hispid culm and foliage leaf sheaths, subsidiary branch- es that do not rebranch, long ligules, and spikelets with the sterile lemmas extending approximately one-half the spikelet length. This species has been collected only in the cloud forests of Costa Rica at elevations of 1,500 to 2,600 m, where it can form large colonies in disturbed areas. The peculiar bud complement of this species is one of its most distinctive features. The central bud is circular in outline rather than triangular (Fig. 2E), and when it develops, the resulting branch initially diverges nearly 90? from the main culm. Usually four to nine smaller subsidiary buds subtend the central bud. As the subsidiary branches develop, they too diverge strongly, be- coming geniculate and bending outward, away from the central bud. 2. Chusquea latifolia Clark, sp. nov. TYPE: Co- lombia. Tolima: El Libano a Murillo (Km 11 al 22 de la carretera), subpáramo en el Alto de Peñones, 2,200-2,950 m, 20 July 1947 (fl), Garcia-Barriga 11259 (holotype, US; isotype, COL). Figure 3. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 Culmi scandentes, usque ad 1. 3 cm diam., usque ad m longi extensa, 17.8-30 cm longa, pubescentia ad juncturam vaginae et laminae: vaginae persistentes, ipee senis connatae basi, 15.2-26.5 cm longae, usque ad 8.5 c latae basi, multo longior quam E. scabrae vel interdum pubescentes; laminae deciduae, triangulares, mum d gemma centralis singularis b brae inferne, inferme tessellatae; ligula interna plerum- que conspicua, rotundata = truncata, (1-)2-15 mm longa, plerumque pubescen glabra; rami appressi, secundi, angulati, glabri. - -12 mm lo ceolata, — 2.4-2.5 m Lemm latum, E 3.2 lemma ste 6.5 mm longum, 7-nerve. Lemma fe ceolatum, Tid viculare, apiculatum, 8-10.6 mm 9- vel 11-nerve. Palea ovata-lanceolata, bicarinata, sul- cata, non pcc 8.3-9.6 mm longa, 6- vel 8-nervis. Rhizomes leptomorph. Culms viny, scandent, to 1.5 cm diam., to 40 m long, often trailing for part of the length. /nternodes terete to slightly laterally compressed, solid, 25-39 cm long, gla- brous to scabrous or pubescent just below the nodes. Culm leaves extending almost the length of the internode, 17.8—30 cm long, pubescent at the juncture ofthe sheath and blade, chartaceous; sheaths persistent, triangular, fused to 2.5 cm above the base, 15.2-26.5 cm long, to 8.5 cm wide at the base, at least 9 times as long as blade, scabrous or sometimes pubescent; the margins ciliate near the apex; blades deciduous, trian- gular, 1.1-3.5 cm long, Ls scabrous or sometimes glabrous, adaxially apex acuminate, the margins ciliate girdle usu- ally well developed, to 1.5 cm forming a flap through emerge, often disintegrating before the rest of the culm leaf; inner ligule a short, ciliolate rim; outer ligule lacking. Nodes only slightly swollen, at mid- culm with one triangular central bud subtended by two smaller subequal buds. Branching infra- vaginal, central bud sometimes developing into a robust branch more or less equal in size to the main culm and rebranching; subsidiary buds 2, developing into leafy branches 17-66 cm long 1985] CLARK — CHUSQUEA 869 SN ~ LN X > yU DM E SS Z s —— SA c) E. 7 p a hy "P Z I = 37 VA, 77 772 | SSS 22 —— V a MY == < F 7 A U SN A. A P f; w ⁄ /, Ú EN — " ZZ 7 i \ \ — > n ZZ — NS = I! WONT yy E Muy’ Cry 4 Ca Aiete LUANG: e ERG G IOAN VILE - I m Y! (u Dj, FiGURE3. Chusquea latifolia. — A. Leafy branch complement, scale = 2 cm (based on Killip & García 33904).— B. Inflorescence, scale = 2 cm.—C. Spikelet, scale = 1 mm (B, C based on García-Barriga 12259).—D. Culm leaf, scale = 2 cm. —E. Bud complement, scale = 1 cm D, E based on Calderón 2997).—F. Node with branches ing infi inally (b f sheath with fused margins), scale = 1 cm (based on McClure 21748).—G. girdle. oo c 870 that y from their lower nodes. Foliage leaves 5-9 per complement; sheaths car- inate, usually pubescent, especially along the keel, occasionally E the margins usually gla- rous or occasionally sparsely ciliate; blades lan- ceolate to broadly lanceolate, 13.3-32.1 cm long, I cm wide, L: W = 3.5:6, the adaxial sur- ce glabrous, not tessellate, the abaxial surface boobs weakly tessellate, the apex acuminate, the margins cartilaginous, serrulate, the base rounded to rounded-truncate; pseudopetiole dis- tinct, 3-9 mm long; outer ligule a conspicuous, stiff rim 0.5-2 mm long; inner ligule usually con- spicuous, rounded to truncate, (1-)2-15 mm long, usually abaxially pubescent at least on the lower half, chartaceous. Inflorescence a narrow panicle 20.5-42.5 cm long; rachis glabrous, one side flat and slightly ridged, the other rounded; branches appressed, secund, arising only from the flat, slightly ridged side of the rachis, angular, 3-9 cm long at the base of the panicle, becoming pro- gressively shorter toward the apex with the spike- lets eventually arising directly from the rachis, glabrous; pedicels short. Spikelets ovate-lanceo- late, 8.2212 mm long, slightly falcate. G/umes 2; glume I triangular, 0.5-1.3 mm long, apically rounded, glabrous, lacking nerves; glume II ovate- lanceolate, 2.4-2.5 mm long, apiculate, abaxially abrous, sdaxially sparsely pubescent near the apex, 3- . Sterile lemmas 2; sterile lemma I Suche e 3.2-4.2 mm long, apiculate, abaxially glabrous, adaxially pubes- cent below the apex, 7- or 9-nerved; sterile lem- ma II ovate-lanceolate, 4.4—6 ertile lemma ovate- lanceolate, shiny, 8-10.6 mm long, apiculate, abrous, 9- or 11-nerved. Palea ovate-lanceo- late, 2-keeled, sulcate, shiny, 8.3-9.6 mm long, abrous, 6- or 8-nerved, the apex acute but not apiculate. Lodicules 3; anterior pair 2.5-2.7 mm long, the upper margins ciliate; posterior lodicule 1.6 mm long, narrower than the anterior, ciliate on the upper margin. Stamens 3; anthers linear, 4.5—5.2 mm long. Ovary glabrous, with 2 styles bearing plumose stigmas. Fruit unknown. Representative specimens examined. COLOMBIA. selvas densas y hümedas e L). RISARALDA-QUINDI i , McClur 21748 (ISC, US). vALLE: old Cali-Buenaventura road, 17 km NW from Cali, Villa Monica-El Yo, , confluencia del río Pichindecito con el Pichindé, 7 Nov. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 1944 (fl), oo 18767 TU Mig pa den te orien li, An dental, Killip & García 33904 (US); along Cali-Buena- ntura Hwy. on E slope of Cordillera Occidental, McClure 21234 (ISC, US). < © This species is characterized by its large, wide leaves, rebranching subsidiary branches, sca- brous sheath with a relatively Bin blade, and somewhat falcate spikelets with apiculate (but not distinctly awned) lemmas. Known only from Colombia, Chusquea latifolia generally inhabits cloud forests and ranges from 1,600 to 2,700 m elevation, although it may be found as high as 2,950 m in subparamo vegetation. The vine forms an extensive network, which clambers over trees and other vegetation. Frequently portions of the culm will run for several meters over the ground, eventually turning downward and entering the soil to re-emerge as another culm some distance away. The bud complement of C. /atifolia is unusual, consisting of one triangular central bud subtend- ed by two (sometimes three) smaller, subequal subsidiary buds that begin to proliferate early (Fig. 3E). The central bud frequently does not develop, but the subsidiary buds develop and rebranch vigorously, often producing mature branch complements of up to 12 branches. Nodes of the main culm contain one root primordium on each side of the central bud (or branch). The nodal region of the culm is well marked, this band of tissue usually appearing slightly darker in color and being rugose in texture. The rhi- zomatous segments of the culm may be distin- guished by their lack of such differentiation in the nodal region. 3. Chusquea serpens Clark, sp. nov. TYPE: Costa Rica. Alajuela: edge of Valley of Río Cari- blanco, Cariblanco, 830 m, 26 Aug. 1968 (fl), Pohl & Davidse 11023 (holotype, ISC; isotype, US). Figure 4. Culmi scandentes, usque ad 1 cm diam., usque ad rigidis, sparsis usque ad densis; laminae plus minusve cordatae, aapea setosis, 3-8. 4 cm longae, non lon- gior q quam vag minens, ae ad 5 mm longum, confertim pubescens. es ad culmum medium naire centralis singularis riangularis subtenta a 2 gemmis parvioribus subae- ee Ramificatio s Folia 3-7 in 1985] CLARK — CHUSQUEA 871 — ) | | i | i R = ——= j Ficure 4. Chusquea serpens. — A. Flowering branch, scale = 1 cm (based on Pohl & Davidse 11023).—B. Spikelet, scale = 1 mm (based on Pohl & Davidse 11176).—C. Culm leaf, scale = 1 cm (based on Young 101).— D. Bud complement, scale = 1 cm.—E. Node with branches emerging infravaginally, scale = 1 cm (D, E based on Clark 283). 872 complemento: vaginae plerumque glabrae; laminae lineares-lanceolatae vel lanceolatae apicibus setosis, bus rotundatis-truncatis Spiculae ovatae-lanceolatae, 11.3-14.6 mm longae. Glumae 2: glume I triangulata vel lanceolata, apice rotundatis, 1.2-2.5 mm longa, |-nervis; gluma II ova- ta-triangularis, longa-aristata, 3.5-7 mm longa, 1- vel 3-nervis. Lemmata sterilia 2: lemma sterile I ovatum- . Palea ovata-lanceolata, apice 'bicari- m sulcata, RUE 10.1-11.5 mm longa, 2-4(-6- 8)-n Rhizomes leptomorph. Culms viny, scandent, to 1 cm diam., to 20 m long, often trailing for part of the length. /nternodes terete but often shallowly sulcate above the primary bud, solid, to 39 cm long, retrorsely scabrous for most of their length, glabrous just above the node, green to green-mottled with purple. Culm leaves not persistent, extending about halfway up the in- ternode, 10—20.1 cm long, chartaceous; sheaths more or less rectangular but flaring slightly at the base, loosely wrapped around the culm, 6.7-11.3 cm long, 2.5—4 cm wide at the base, usually about 1.5 times as long as the blade, occasionally equalling the blade, or to twice as long as blade, glabrous to verrucose with sometimes numerous rigid hairs,the overlapping margin ciliate; blades usually deciduous before sheaths separate from the culm, moderately to distinctly cordate, 3-8.4 cm long, not longer than the sheath, glabrous, the apex setose, the margins smooth to ciliate in some specimens; girdle more or less prominent, to 5 mm wide, densely pubescent, forming a flap through which the branches emerge, disintegrat- ing before the rest of the culm leaf; inner ligule a short, ciliolate rim; outer ligule lacking. Nodes only slightly swollen, at mid-culm with one tri- al buds. Branching infravaginal, central bud often developing into a robust branch more or less equal in size to the main culm, this branch rebranching extensively, forming a long clam- ering network; subsidiary buds 2, developing into leafy branches 15-35 cm long that rarely rebranch. Foliage leaves 3-7 per complement; sheaths carinate, usually glabrous, occasionally ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 72 scabrous or pubescent near the margins and/or apex, the overlapping m m pes blades lin- ear-lanceolate to lanceolate, 11.7—35.8 cm long, .1 cm wide, L: W = 5: P Athe adaxial sur- face glabrous, not tessellate, the abaxial surface mostly glabrous and pilose only near the apex to entirely pilose, not tessellate to weakly so, the apex tapering into a long bristle-like tip, the mar- gins cartilaginous, serrulate, the base rounded- truncate; pseudopetiole distinct, 3-12 mm long; outer ligule a conspicuous, stiff rim 0.5-1.5(-2.5) mm long; inner ligule truncate, 1-2(-2.5) mm long, chartaceous. Inflorescence. a narrow panicle 33-55(-60) cm long, term side flat and slightly ridged, the other rounded; branches appressed, secund, arising only from the flat, slightly ridged side of the rachis, angular, 5-9 cm long at the base of the panicle, becoming progressively shorter toward the apex with the spikelets eventually arising directly from the rachis, glabrous or pilose; pedicels short. Spike- lets ovate-lanceolate, 11.3-14.6 mm long. Glumes 2; glume I somewhat variable, triangular to lan- ceolate, 1.2-2.5 mm long, apically broadly acute, glabrous, 1-nerved; glume II ovate-triangular, 3.5-7 mm long, long-awned, glabrous to slightly pubescent adaxially just below the apex, 1- or 3-nerved. Sterile lemmas 2; sterile lemma I ovate- lanceolate, 5.4-8.6 mm long, awned, glabrous to densely pubescent adaxially, (5—)7(-9)-nerved; sterile lemma II ovate-lanceolate, keeled, 7.7— 9.9 mm long, awned, glabrous to densely pubes- cent adaxially for the upper one-half to two-thirds of its length, (5-)7—9- or 11-nerved. Fertile lem- ma ovate-lanceolate, boat-shaped, shiny, 10.6— 13.2 mm long, short-awned, glabrous to densely pubescent adaxially for the upper one-half of its ength, usually obscurely 7—18-nerved. Palea ovate-lanceolate, 2-keeled, sulcate, shiny, 10.1— 11.5 mm long, apiculate, marginally ciliate to- ward apex, glabrous, 2-4(-6—8)-nerved. Lodi- cules 3; anterior pair 2-2.5 mm long, the upper margins ciliate; posterior lodicule 2.5-3 mm long, narrower than the anterior, ciliate on the upper margins. Stamens 3; anthers linear, 5.5-7.5 mm long. Ovary glabrous, with 2 styles bearing plu- mose stigmas. Fruit unknown. Representative specimens examined. COLOMBIA. jeg A dillera Oriental, vertiente oriental, Sucre, 4 Apr. 1940 (fl), Cuatrecasas 9068 ( uri oa piis HUI- LA: Río joe near prep San of Colombia, 24 Dec. 2 (fl), pude d Holdridge 19568 (US). META: Las eheu plateau between Río 1985] Papamene and Río Duda, Colombia-Uribe trail, 8-9 km SW of Uribe, 22 Dec. 1942 (fl), Fosberg 19504 ral on road to Dominical, Pohl 14187 (ISC). ECUADOR. of Reventador on the road e finc n Chi riquí Pr cloud forest, 17 Apr. 1968 (fl), Kirk- 2 (MO). nes de Horni little past the hill at 1,640 m, 978 (f), Burandt VO457 (MO); Parque Nacional Yacambi, ca. m from. main entrance, ata picnic area, Clark et g 231 á, above Finca La S 8 km SW of Catanii. pagaspa 56392 (U Chusquea serpens is characterized by its viny, stoloniferous habit, branch complement with only two subsidiary branches, cordate culm leaf blade almost equal to the sheath in length, secund in- florescence, and awned spikelets. Chusquea ser- pens and C. latifolia are similar in many respects, particularly in their clambering habit, bud com- plement of one triangular central bud subtended by two subsidiary buds, and large, secund inflo- rescences (not so marked in C. /atifolia). Both species clearly fall within Chusquea because of their solid culms, multiple independent buds, and typical one-flowered spikelets but may form a natural group within the genus based on the un- usual nature of the bud complement. Chusquea serpens can be distinguished from C. latifolia by its generally less robust appearance, relatively larger, cordate culm leaf blade, and awned spike- lets. Chusquea serpens has appeared previously in the literature as the ‘‘Cariblanco population" of Chusquea and Chusquea “‘cariblanco” (Pohl, 1980, 1983). Chusquea lanceolata Hitchcock from Guate- mala and Mexico, closely related to C. serpens, differs in its relatively smaller culm leaf blades, more open inflorescences, and spikelets with CLARK — CHUSQUEA 873 apiculate, not awned, lemmas. The two species are very similar with respect to their clambering habit, branch complement, and large leaves. Chusquea lanceolata tends to occur at somewhat higher elevations than C. serpens, although still in the montane forest zone. Chusquea serpens is the only species of Chus- quea known to occur in both Central and South America, extending from Costa Rica to Vene- zuela and along the Andes to Ecuador. It is found in cloud forests throughout its distribution, rang- ing in elevation from 800 to 2,100 m. Typically the plant forms large, tangled networks that fes- toon the surrounding vegetation. In contrast to C. latifolia, the central branches in C. serpens rebranch to produce the long, trailing and hang- ing culms. LITERATURE CITED CALDERÓN, C. E. & T. R. SODERSTROM. 1980. The genera of Bambusoideae (Poaceae) of the Ameri- can du ap, keys a and comments. Smithsonian r. Bot. 44: i-iii, 1-27. Daansntt, S. 1981. The genus Dinochloa (Gra- ae — Bambusoideae) in Sabah. Kew Bull. 36: 3. Foster, A. S. & E. M. GIFFORD, JR. 1974. Compar- ative Morphology of Vascular Plants, 2nd edition. . H. Freeman and Company, San Francisco. HACKEL, E. 1903. Neue Gräser. Oesterr. Bot. Z. 5 153-159. McCLuuRE, F. A. 1973. Genera of bamboos native to the New World (Gramineae: Bambusoideae). Smithsonian Contr. Bot. 9: i-xii, 1-148. [Edited by T. R. Soderstrom.] Munro, W. 1868. A monograph of the Bambusa- i 1 2 "| z ve Call 4L - Trans. ceac, g p p Linn. Soc. London 26: 1-157. NEES VON ESENBECK, C. G. D. 1835. Bambuseae Bra- silienses: Recensuit, et alias in India orientali pro- venientes adjecit. Linnaea 9: 461- PILGER, R. 1905. Gramineae Andinae. I. Bambuseae Repert. Spec. Nov. Regni Veg. 1: 145- Pour, R. W. 1980. Family 15. Gramine n W. Burger (editor), Flora Costaricensis. Fi donis. Bot. n.s. 4: 1-608. 1983. Current blooming status ofthe bamboo flora of Costa Rica. Amer. J. Bot. 70: 126. [Ab- SODERSTROM, T. R. be C. E. CALDERON. 1978a. The species of Chusquea (Poaceae: Bambusoideae) with verticillate bade Brittonia 30: 154-164. b. Mies cd and Swallenochloa (Po- aceae: Bambusoideae): generic relationships and new species. eno °, 30: 297-312. NOTES PHARUS PARVIFOLIUS SUBSP. ELONGATUS (POACEAE), A NEW SUBSPECIES FROM TROPICAL AMERICA While working on a revision of the Phareae (Poaceae: Bambusoideae), it became evident that the widespread neotropical species Pharus par- vifolius Nash consists of two entities that merit subspecific recognition. The present note is nec- essary to validate the name of the new taxon, so that it will be available for use in my Flora Me- soamerica treatment. À more detailed study of this species will be presented in a forthcoming revision of Pharus Pharus parvifolius was described from Haiti (Nash, 1908). It is differentiated from other species in i genus by its colonial habit (old culms be bent and root at the nodes), strongly two-ranked foliage, lower (morpholog- ically adaxial, as in all species in the Phareae) leaf blade surfaces with longitudinal sclerenchy- matous bands between the small lateral veins, and linear-oblong pistillate spikelets with purple glumes. Subspecies parvifolius is characterized by the following features: erect stems 25-50 cm long; leaf blades lanceolate, 6.5215 cm long, 1-2.5 cm m m long, the (naked) axis termi- ing the eens 2-5 cm long; and pistil- cc spikelets 8-14 mm long. It is frequent in Cuba, Hispaniola, eastern Venezuela, and Suri- nam, with scattered stations in Central America, the smaller West Indian islands, French Guiana, Ecuador, and Brazil. A more robust series of populations, Mio] distributed in the Neotropics but with a concen- tration of sites in Central America, is d bed Pharus parvifolius Nash subsp. elongatus Jud- ziewicz, subsp. nov. TYPE: Costa Rica. Car- tago: Los Espaveles Nature Trail into for- ested canyon of the Río Reventazón, Centro Agronómico Tropical de Investigación y Ensenanza (CATIE), 3 km E of Turrialba, 9°54’ N, 83?39'W, 550—600 m, 9 May 1983, in young fruit, Liesner, Judziewicz & Pérez 15281 (holotype, CR; isotypes, BAF, BM, CEPEC, COL, CR, CTES, DUKE, F, IBUG, INPA, ISC, LE, LPB, MEXU, MO, MVFA, NY, P, PMA, RB, SP, US, VEN, WIS, XAL). ANN. Missouni Bor. GARD. 72: 874—875. 1985. ffert a P. vel (ir subsp. parvifolius statura al- us (60-125 cm), laminae lineare-lanceolatae long- lore " um 32 cm) latioreque (2-3.5 cm), nervi sub angulo 4-8°, inflorescentiae longiore (20-35 cm), axis termin- alae joins (3.5-9 cm), et spiculae feminae longiore ). (10-16 mm Similar to subspecies parvifolius, but larger: culms stout, 3-6 mm diam.; erect culms 60-125 cm long; leaf blades linear to linear-lanceolate, 18-32 cm long, 2-3.5 cm wide, tapering to both base and apex, the lateral veins diverging from the midrib by 4-8°, panicle 20-35 cm long, the naked, terminal portion of the axis 3.5-7(-9) cm fone pistillate spikelets slightly larger than in the typical variety; glumes 5.5-8 mm long, lemma 10-16(-17) mm long. Representative specimens examined (only one given percountry). MEXICO. VERACRUZ: Estación Biológica Los Tuxtlas, w epey 1186 (ENCB, F, MEXU). BELIZE. TO Sandino 112 . C of Volcán Rincón i la Vieja, 1,150 m, Pohl 12667 (F, ISC, MO, US). PANAMA. CANAL AREA: Barro Col- orado pee Foster 1766 (DU PMA 50 Ekman H-3825 (S, US). SURINAM. Be tween Kabalebo and _Coppename Rivers, 20 km Bakhuis, Flor- schütz & Maas 2523 (U). ECUADOR. “MANABI: El Ro- sario, o, Bages, 15188 (F, P, fragm. US ex K). PERU. LORETO: Pue A Williams 5285 (F, GH, LE, US). BR AZIL. BAHIA: gogi, Curran 110 (US). BOLIVIA. LA PAZ: Geen 2. 000 ft., White 1043 (US). Both subspecies grow in the same habitat, moist to wet forests from near sea level to 1,000 m or more. The tendency toward geographical sepa- ration in these two entities, and their nearly ab- solute vegetative distinctness from each other (there are a few intermediate populations in northern Central America and Peru) suggest the rank of subspecies rather than variety. I do not believe mat specific recognition is warranted. In addition to t men- tioned, there is an apparent lack of ‘temporal separation in flowering time between the two 1985] subspecies, both blooming in the Northern Hemisphere dry season (about January to May). Also, although vegetatively most specimens can be assigned without much difficulty to one taxon or the other, there is a much greater degree of overlap in reproductive characteristics, espe- cially in the pistillate spikelet. Pohl and Davidse (1971) reported a chromo- some number of 2n = 24 (i.e., diploid, as in most other species in the genus) for the (now type) population of subsp. elongatus at Turrialba, Cos- ta Rica. There are no chromosome counts of subsp. parvifolius. Assistance was received from the E. K. and O. N. Allen and J. J. Davis Funds of the University NOTES 875 of Wisconsin, Wisconsin Natural History Coun- cil, Museo Nacional de Costa Rica, and Missouri Botanical Garden. Thanks are due to H. H. Iltis, G. Davidse, R. L. Liesner, and T. H. Cochrane. LITERATURE CITED NasH, G. V. 1908. Two new grasses from the West Indies. e Torrey Bot. Club 35: 301- Pour, R. W. & G. Davipse. 1971. Chromosome cds ‘of Costa Rican grasses. Brittonia 23: 293- 324. —Emmet J. Judziewicz, Department of Botany, University of Wisconsin, Madison, Wisconsin CAMPYLOCENTRUM TENELLUM (ORCHIDACEAE), A NEW SPECIES FROM PANAMA Campylocentrum Bentham is a genus of ap- ama came to my attention. After careful exam- proximately 45 species of small, epiphytic, neo- ination of the South American taxa, I feel con- tropical orchids. In a revision ofthe Costa Rican fident in proposing this new species. species (Todzia, 1980), a new taxon from Pan- FIGURE 1. ym tenellum. — A. Whole plant.—B. Raceme.—C. Flower.—D. Lip and spur.—E. ea petals, and lip ANN. Missouni Bor. GARD. 72: 876—877. 1985. 1985] Campylocentrum tenellum Todzia, sp. nov. TYPE: Panama. Panamá: La Eneida, region of Cer- ro Jefe, 26 Oct. 1969, Dressler 3758 (holo- type, CR). Figure 1. anta . iia 1-5.5 cm alta; folia li- longa, be mm lata, va- 7-10 mm longa; flor albus, sepala ovata, petala Due labellum tri- lobatum, calcar rectum acutum hispidulum Small, epiphytic plant with pendant or as- cending stems, 1-5.5 cm tall. Roots basal or al- Ip with the lower 1 or 2 leaves, fibrous, flexuous, glabrous, to 15 cm long. Stem simple, Ed. ‘slightly fractiflex, 1-2 mm wide, invested by leaf sheaths. Leaves small, thin, distichous, linear-lanceolate, margin smooth to lightly den- ticulate and crenulate, apex retuse, mucronulate to caudate, articulated to the persistent sheathing bases, 0.8-1 cm long, 1-2 mm broad; leaf sheath tubular, compressed, lacerate on the apical mar- gin. Inflorescences produced singly opposite the leaves, not exceeding the leaves, ascendant to spreading, 7-10-flowered; rachis 7-10 mm long, minutely puberulent; floral bracts concave, tri- angular, acuminate, 1 mm long, ciliate along the margin. Flowers small, distichous, white to cream-colored; sepals ovate, acute, 1 mm long, ] -nerved, outer surface lightly hispidulous; pet- als linear, acute, 1.25 mm long, longer than the sepals, 1-nerved; lip ovate, 1.5 mm long, slightly 3-lobed at the base, lateral lobes obtuse, abbre- NOTES 877 viated, mid-lobe ovate, acute; spur straight, cy- lindric, acute, 1.5-2 mm long, outer surface light- ly hispidulous; column short, 0.5 mm long; ovary 0.75-1 mm long, hispidulous with short, stiff hairs 0.1 mm long. Capsule not seen. PANAMA. PANAMÁ: El Llano- N of El Llano, 30 Sept. 1973, Ee cd viridem rti Hwy., 15- pee s.n. (TEX). Campylocentrum tenellum is most similar to C. parvulum Schltr. of Costa Rica, from which it is differentiated by its lacerate leaf sheaths, shorter inflorescences, and flowers with a longer spur. It is easily distinguished from all other leafy erate leaf sheaths, and the small size of the plant. Campylocentrum tenellum belongs to section Campylocentrum as does C. parvulum. I thank R. L. Dressler for making these spec- imens available to me, S. Thurston and L. Vo- robik for graciously providing the drawing, and B. Simpson an . C. Johnston for helpfully critiquing the manuscript. LITERATURE CITED Topzia, C. A. 1980. A revision of the Costa Rican species of Campylocentrum (Orchidaceae). Bre- nesia 18: 117-136. — Carol A. Todzia, Department of Botany, Uni- versity of Texas, Austin, Texas 78713. SAURAUIA MOLINAE, A NEW SPECIES OF ACTINIDIACEAE FROM CENTRAL AMERICA In the course of preparing the taxonomic treat- ment of Saurauia (Actinidiaceae) for Flora Me- soamericana, I was not able to place a group of specimens recently collected from Honduras in the classification scheme of Hunter (1966) or Soejarto (1980). Nor do these specimens fit the recently described species of this genus from Mexico (Keller & Breedlove, 1981). Further study showed that these specimens belong to a new taxon described below. Saurauia molinae Soejarto, sp. nov. TYPE: Hon- uras. Departmento de Comayagua: 3 km S of Siguatepeque, 1,300 m, thickets along creeks in pine forest; weak tree, flowers white, 9 Jan. 1969, Molina 23296 (holotype, MO!; isotype, F!). Figure 1 Saurauia molinae Soejarto; inter species seriei Gy- notrichae foliis valde coriaceis indumento sepalorum expositorum in alabastro et inflorescentiarum omnino . differt Trees, to 12 m tall. Branchlets terete, distinctly hollowed, with leaf scars prominent, sparingly to densely appressed pubescent, pubescence denser along Juvenile parts and shoots, trichomes plu- mulose-setose to plumulose-sericeous, yellow to rally arranged, clustered below tips of branchlets; ades obovate to elliptic-obovate, 10-25 cm long, 5-13 cm wide, coriaceous, in dry state dark to gray-brown adaxially, olive brown abaxially, much darker adaxially, rounded-acute to very shortly acuminate to rarely obtuse at apex, cu- neate to broadly acute to obtuse to rounded, but less frequently oblique at base, subentire to finely setaceo-serrulate along margins, teeth or setae less than 1 mm long, 2-8 mm apart, secondary veins 14-25 pairs, forming a wide-angled V (be- coming more acute toward apex) with straight to somewhat curving arms that lightly curve and usually dichotomize shortly before reaching margins and may or may not form a lightly anas- tomosing network, tertiary veins elevated be- neath, more prominent than lesser reticulation; upper surface usually smooth and sometimes opaque or subglossy, scattered pubescent, tri- chomes limited primarily to major veins, yellow to white, hirsute to sericeous, appressed to the epidermal surface, tufted to barbate at their base; ANN. Missouni Bor. GARD. 72: 878—880. 1985. abaxial surface somewhat smooth, unevenly pu- bescent, denser along major veins, glabrous to glabrescent on other parts and between veins, trichomes along veins appressed to epidermal surface, plumulose-hirsute to plumulose-seri- ceous, mixed with and obscured by cottony or arachnoid indument in juvenile leaves, minor reticulation distinctly discernible; petiole sub- terete, lightly Pug adaxially, 2— cm ong, 2-3 mm diam., a antly to densely pu- bescent, trichomes to ae mm long, plumulose- sericeous and mixed with arachnoid indument, the latter more distinct in juvenile leaves. Jnflo- rescence a thyrse, inserted at 30? angle to per- pendicularly to branchlet, straight, becoming somewhat pendulous in fruit, moderately branched, 15-90-flowered, 7-18 cm long, 4-9 cm wide, abundantly to densely pubescent es- pecially on terminal parts, pubescence yellow- brown, cottony, individual trichomes plumu- lose-hirsute mixed with and obscured by the cottony indument, peduncle 3.5-9 cm long, bracts linear to subulate, to 5 mm long, densely cottony pubescent. Flowers actinomorphic, perfect, of two types, one short-styled belonging to functionally staminate plants, the other long-styled belonging to functionally pistillate plants, 13-18 mm broad when open, buds subglobose, to 5 mm across, pedicels 2-7 mm long, to 10 mm in fruiting stage, bracteoles linear to subulate, to 4 mm long, densely cottony pubescent; sepals 5, reflexed at apex early in bud, remaining so in fruit, aesti- vation quincuncial, outer two ovate to elliptic to oblong-elliptic to orbicular-ovate, acute at apex, imbricated one ovate-oblong to suborbicular, rounded at apex, inner two ovate-oblong to ovate- orbicular to suborbicular, rounded at apex, all 7-13 mm long, 6-13 mm wide, the outer two the smallest, all densely cottony pubescent on exposed parts in bud, glabrous on imbricated parts and inner surface, outer two entire to ob- scurely ciliate, inner three distinctly ciliate; petals 5, white, connate basally and falling off as a unit with stamens after anthesis, oblong to oblong- obovate to oblong-elliptic, 6-10 mm > wn pubescent at base, trichomes filiform, anther lin- ear-oblong, 2 mm long, extrorse; ovary globose, 5-loculed, 5-sulculate, densely pubescent, tri- 1985] NOTES oe Sm E IT Tr, ü i» i. m 4-2 pate e re EL os AE t Winer = E s a E i ee eed Z x di R = =e E. Mare Spa ee SASS ESP > Tameme m ra ell. 31 < A -— = — Y is ue - er FIGURE 1. Saurauia molinae Soejarto. — A. Flowering branch from a functionally staminate plant. —B. A functionally staminate flower, top view; note the obsolete-styled, densely pubescent ovary surrounded by sta- mens.—C. A fi i wi unctionally staminate flower, b ng the pattern of pubescence on the outer surface ottom view, sho of the sepals. —D. A fruit from a functionally pistillate flower. —E. Enlarged portion of the abaxial leaf surface to show details of venation and pubescence along the major veins. A-C, E are based on Molina & Molina 25585 and Molina 23296 (holotype); D is based on Hazlett 14 chomes yellow-brown, styles obsolete in func- tionally staminate plants, to 4.5 mm long in func- tionally pistillate plants, stigmas simple to subcapitate. Berries globose, 5-loculed, 5-sulcate, sparingly to abundantly pubescent (epidermal 71. surface of ovary usually visible), trichomes yel- low-brown, of fimbriate-stellate type, with arms straight or wavy, hyaline and filiform, to 1 mm long and upwardly appressed, styles 5, persistent, to 5 mm long, sepals persistent. 880 E examined. HONDURAS. DEPT. COMAY :10k of Siguatepeque, between El Portillo. mis El nir 1,300 m, cut-over forest, weak tree, flowers white, 10 Mar. 1970, fr., Molina & Molina , common, flowers white, * Montaña,” E Apr. 1971, fr., D 25985 (F); S of i , tree beside stream that Doi through 974, fr., Hazlett | 47 1 (MO). DEPT. Lejarsia, between Kms 9 and 11 2 . DEPT. LA PAZ: Cordillera Guajiquiro, be- tween Las Marias and El Cerron, 1,700 m, cut-over mixed forest, tree 12 m, flowers white, 20 Mar. 1969, fr., Molina & Molina 24232 (F) This new taxon belongs to the series Gynotri- chae (Hunter, 1966). Like all other members of = series (S. veraguasensis Seem., S. sq ucta Hunter, and S. waldheimia Busc.), it is deins by the possession of pubescent ovary. S. molinae Soejarto may be distinguished from the other Central American members of the se- ries by the following key: sth ctallata teo 1 1 la. Leaves abaxial surfac 2a. cur sia ER usually more than 10- flowered veraguasensis 2b. aio M usually less than 7-flow- . squamifructa . Leaves pucr stellate trichomes on t abaxial surface or, if aci. limited only to the axils of secondary vein eo = ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 72 3a. Blades chartaceous, normally less than 5 cm long, axils of secondary veins vil- lous-barbate abaxially, sepals with ap- pressed-setulose or plumulose-setulose trichomes on the exposed surface in bud w ° . Blades coriaceous, normally mor 15 cm long, axils of secondary veins not villous-barbate abaxially, sepals cottony pubescent on the exposed surface in bud molinae The new taxon is dedicated to Dr. Antonio Molina, the Honduran botanist, whose extensive explorations of the flora of Honduras resulted in the discovery of this species. I would like to thank Ms. Marlene H. Werner, a senior scientific illustrator ofthe Field Museum of Natural History, who kindly prepared the il- lustration. LITERATURE CITED Hunter, G. E. 1966. Revision of Mexican and Cen- tral American Saurauia (Dilleniaceae). Ann. Mis- souri Bot. Gard. 53: s 89. KELLER, B. T. & D. Two new species of Saurauia TE a Mexico. yst. Bot. 6: 65-73. SOEJARTO, D. 980. Revision of South American oe (Actinidiaceae). Fieldiana, Bot. n.s -141. —D. D. Soejarto, Botany Department, Field Museum of Natural History, Chicago, Illinois 60605 and PCRPS, University of Illinois at Chi- cago, 833 South Wood Street, Chicago, Illinois 60612 ERRATA The following corrections should be ecd in e bio titled “The order Myrtales: circumscription, variation, and relationships," ' by Rolf Dahlgren and Robe F. Thorne [71(3): 633-699. 1984 (1985)]. A and B should be switched in the legend to Figure 1. The legend d Figure 14 should be switched with the legend to Figure 16. ANNALS F THE MISSOURI BOTANICAL GARDEN The Missouri Botanical Garden is pleased to offer individual subscription rates to the ANNALS OF THE MISSOURI BOTANICAL GARDEN. The ANNALS, published quarterly, contains papers in systematic botany, floris- tics, plant biogeography, reproductive biology and other topics of interest to systematic botanists. Proceedings from symposia on a wide variety of subjects, held both at the Missouri Botanical Garden and elsewhere, are also published regularly in the ANNALS. Annual rates: $30.00 (North America), $35.00 (elsewhere). Enclosed please find $__—__— to cover my individual sub- Name scription to the ANNALS at $30.00__ or $35.00__ (check ER one). Please begin my subscrip- tion with the 1985 ( ) or 1986 ke ( ) volume (check one). ee ee City State Zip Code Contents continued from front cover Miocene Fossil Grasses: Possible Adaptation in Reproductive Bracts (Lem- ma and Palea) Joseph R. Thomasson 843 Graminoid Responses to Grazing by Large Herbivores: Adaptations, Exaptations, and Interacting Processes Michael B. Coughenour ...... 852 Three New Species of Chusquea (Gramineae: Bambusoideae) Lynn G. AKR I 864 NOTES Pharus parvifolius subsp. elongatus (Poaceae), a New Subspecies from Trop- ical America Emmet J. Judziewicz 74 Campylocentrum tenellum (Orchidaceae), a New Species from Panama CIE TEE. t uude ec S S S 5 uM 876 Saurauia molinae, a New Species of Actinidiaceae from Central America J) D Soy — -ea 878 FRA TA 50 o eem M i M UNE 880