Missouri Botanical Garden Research Articles

Annals of the Missouri Botanical Garden, Vol. 71, No. 3

Annals of the Missouri Botanical Garden, Vol. 71, No. 3

Serotaxonomy of Solanum, Capsicum, Dunalia, and Other Selected Solanaceae

Serotaxonomic comparisons were made on about 50 species of Solanaceae. Immunoabsorption data were obtained from reactions of seed protein antigen systems with antibody systems absorbed by several antigen systems in numerous permutations. Relationships of the taxa were computed by newly developed methods. Twenty species of Solanum, mostly of subg. Leptostemonum, were compared using six antisera. Solanum nigrum (subg. Solanum) was serologically distinct from the others. Several groups of species belonging to different sections of Solanum were recognized. Phenetic and phyletic relationships within sect. Androceras were explored. Re-analysis of a complete cubic matrix of immunoabsorption data for eight species of Solanum rearranged relationships slightly but emphasized the divergence between different sections of this genus. A broad survey was made using an antibody system raised to Capsicum annuum. This revealed the distinctiveness of Nicandra from other Solanaceae, several possible inter-generic relationships, but also some unlikely ones. Further studies revealed closer relationships of Dunalia species to Capsicum than to Iochroma. Protein characters are valuable in plant taxonomy for assessing inter-generic and even interfamily relationships (Jensen & Fairbrothers, 1983). Within the Solanaceae several serological studies have been made by workers such as Chester, Hammond, Tucker, Gray, and Lester (see review by Lester, 1979). Because of the complex physico-chemical properties of proteins, and the even more complex phenomena of immunological reactions, many different serotaxonomic techniques have been developed (Jensen & Fairbrothers, 1983). Immunoabsorption resolves subsets of antigenic determinant sites as recognized by corresponding subsets of antibodies after the removal of common antibodies from the antibody system by absorption by other antigen systems. Recent developments in the theoretical and practical aspects of immunoabsorption techniques and the analysis of the resultant data by appropriate numerical taxonomic procedures have been discussed in detail (Lester, 1979; Lester et al., 1983). For this paper these procedures were applied to selected species of Solanaceae. MATERIALS AND METHODS Seed samples from the Birmingham University Solanaceae Collection (Tables 1-3) were used to prepare protein extracts, to induce antibody production in rabbits, and for subsequent immunological experiments using absorbed antibody systems. Procedures, especially the scoring and analysis of results, followed those described by Lester (1979) and Lester et al. (1983). Three sets of experiments were conducted. 1. Twenty accessions of Solanum, mostly of subg. Leptostemonum (Table 1) were compared using six antibody systems (data published in Lester et al., 1983). 2. Eight species from diverse sections of the genus Solanum (Table 2) were compared using eight antibody systems (data published in Lester, 1979). 3. About 25 species of Capsicum, Solanum, and other genera of Solanaceae, mostly of tribe Solaneae (Table 3) were compared using one antibody system (Lester, unpubl. data). Most of these data have now been analyzed by most of the procedures described by Lester et al. (1983), especially by using Jaccard's and similar coefficients to estimate phenetic relationships. Some of the data sets also were subjected to cladistic analysis. Many dendrograms were produced by different analyses of the various sets and subsets of data, some of which are presented herein (Figs. 1-6). I We are grateful to Sarah Marsh for growing these plants, to the Science and Engineering Research Council (U.K.) for financial support, and to Julie Brean, Barbara Klauza, and Sarah Sutton for technical assistance. 2 This paper was part of the Second International Symposium of the Biology and Systematics of the Solanaceae presented at the Missouri Botanical Garden on 3-6 August 1982. 3 Department of Plant Biology, University of Birmingham, Edgbaston, Birmingham B1 5 2TT, England. 4 Department of Nematology, University of California, Riverside, California 92521. ANN. MISSOURI BOT. GARD. 73: 128-133. 1986. This content downloaded from 207.46.13.60 on Thu, 21 Apr 2016 07:28:58 UTC All use subject to http://about.jstor.org/terms 1986] LESTER & ROBERTS-SOLANACEAE SEROTAXONOMY 129 TABLE 1. Listing and classification of the 20 accessions of Solanum species that were compared immunologically using antibody systems to CAP, ROS, SIS, QUT, TOR, and PRN.

Augustus Fendler (1813-1883), Professional Plant Collector: Selected Correspondence with George Engelmann

Augustus Fendler (1813-1883), Professional Plant Collector: Selected Correspondence with George Engelmann

Convergent Evolution of the 'Homeria' Flower Type in Six New Species of Moraea (Iridaceae-Irideae) in Southern Africa

Convergent Evolution of the 'Homeria' Flower Type in Six New Species of Moraea (Iridaceae-Irideae) in Southern Africa

Cytology and Systematics of the Moraea fugax Complex (Iridaceae)

Cytology and Systematics of the Moraea fugax Complex (Iridaceae)

Notes on the Systematics of Hesperantha (Iridaceae) in Tropical Africa

Notes on the Systematics of Hesperantha (Iridaceae) in Tropical Africa

New Neotropical Species of Meliosma (Sabiaceae)

New Neotropical Species of Meliosma (Sabiaceae)

Studies in the Araliaceae of Nicaragua, and a New Widespread Species of Oreopanax

Studies in the Araliaceae of Nicaragua, and a New Widespread Species of Oreopanax

Sudanian Elements in the Flora of Israel

In the Dead Sea Rift Valley of Israel, a northern tongue of penetration of Sudanian elements exists that has traditionally been regarded as composed of Miocene relicts. We postulate that most of these elements have penetrated the area since the end of the Pleistocene. The principal habitats harboring Sudanian elements in Israel are described, including the pseudo-savanna in wadi beds, cliffs and rock formation, oases and in the Dead Sea Rift as well as a variety of secondary habitats outside the Rift. In addition, a statistical analysis of the 116 species in the flora of Israel considered to be Sudanian elements is presented, covering growth form, phytogeographic distribution, dispersal mechanisms, and degree of endemism. Special attention is given to the arboreal elements, including 49 species of trees, shrubs, and perennial vines. A comparison is made between the habitats of these species in East Africa and in Israel: significant habitat shifts were found in some cases. Although paleo-macrofossils are almost non-existent in the area, other evidence, such as low degree of endemism, disjunctiveness, and adaptations to aridity and long-distance dispersal indicate that the great majority of the Sudanian elements in the Dead Sea Rift are of recent origin. The relationships between the typical evergreen Mediterranean elements with paleotropical origins and the extant Sudanian elements in Israel are discussed. Within the portion of the Syrian-African Rift lying between the Red Sea to the south and the watershed of the Jordan River in the Golan Heights, a large number of Sudanian elements, plants as well as animals, are found considerably farther north than anywhere else in their distribution range and in many cases growing under ecological conditions quite different from their norm. In addition, a handful of Sudanian species also occur in the low-lying coastal plain of Israel, almost entirely cut off from the Dead Sea Rift Valley and the bulk of the Sudanian region proper (Fig. 1). Climatic and geological conditions in the Dead Sea Rift Valley explain in part the presI A. Shmida wishes to express his considerable debt of gratitude to the late Prof. M. Zohary who introduced him (and dozens of other students) to the intricacies and unique attributes of the Israeli flora. We thank G. E. Wickens who reviewed the manuscript with meticulous care, helped gather information for some of the distribution maps, and made valuable suggestions for improvement of the text. J. B. Gillett and P. Quezel also kindly reviewed the manuscript and provided stimulating comments and suggestions. Our thanks also go to Sandy Greenberg, Seemah Shemesh, and Marion Milner who provided computer and typing assistance, to Nancy R. Morin and Dr. Marjorie Tiefert for patient and professional editorial guidance, and to Miri Shmida who drew the figures. The assistance of J. Dransfield in preparing Figure 5 is gratefully acknowledged. Any errors or inaccuracies therein are entirely the responsibility of the authors. Finally we thank the numerous guides of the Israeli Society for the Protection of Nature who helped us locate some of the rare and little-known populations of Sudanian plants in Israel. We gratefully acknowledge financial assistance received during the course of this research from the Bath-sheva de Rothschild Fund to A. Shmida and the Clifford M. Hardin Fund of the Fund for Higher Education to J. A. Aronson. 2 Department of Botany, The Hebrew University of Jerusalem, Jerusalem, Israel. 3Rudolf and Rhoda Boyko Institute for Agriculture and Applied Biology, Institutes for Applied Research, Ben-Gurion University of the Negev, P.O. Box 1025, Beer-Sheva 84110, Israel. ANN. MissouRi BOT. GARD. 73: 1-28. 1986. This content downloaded from 157.55.39.253 on Sat, 11 Jun 2016 07:14:14 UTC All use subject to http://about.jstor.org/terms 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 73

Three New Species of Solanum Section Geminata (G. Don) Walp. (Solanaceae) from Panama and Western Colombia

Three New Species of Solanum Section Geminata (G. Don) Walp. (Solanaceae) from Panama and Western Colombia

Morphology and Development of Pistillate Inflorescences in Extant and Fossil Cercidiphyllaceae

Morphology and Development of Pistillate Inflorescences in Extant and Fossil Cercidiphyllaceae

Temperate-Zone Biogeography: Biogeogaphical Relationships between Temperate Eastern Asia and Temperate Eastern North America . Missouri Botanical Garden, St. Louis, 1984. 329 pp., illus. Paper, $15. Reprinted from Annals of the Missouri Botanical Garden , vol. 70, nos. 3 and 4 (pp. 421- 749). From a symposium, St. Louis, Sept. 1982.

Temperate-Zone Biogeography: <i>Biogeogaphical Relationships between Temperate Eastern Asia and Temperate Eastern North America</i> . Missouri Botanical Garden, St. Louis, 1984. 329 pp., illus. Paper, $15. Reprinted from <i>Annals of the Missouri Botanical Garden</i> , vol. 70, nos. 3 and 4 (pp. 421- 749). From a symposium, St. Louis, Sept. 1982.

Temperate-Zone Biogeography: Biogeogaphical Relationships between Temperate Eastern Asia and Temperate Eastern North America . Missouri Botanical Garden, St. Louis, 1984. 329 pp., illus. Paper, $15. Reprinted from Annals of the Missouri Botanical Garden , vol. 70, nos. 3 and 4 (pp. 421- 749). From a symposium, St. Louis, Sept. 1982.

Temperate-Zone Biogeography: <i>Biogeogaphical Relationships between Temperate Eastern Asia and Temperate Eastern North America</i> . Missouri Botanical Garden, St. Louis, 1984. 329 pp., illus. Paper, $15. Reprinted from <i>Annals of the Missouri Botanical Garden</i> , vol. 70, nos. 3 and 4 (pp. 421- 749). From a symposium, St. Louis, Sept. 1982.

Costa Rican Natural History

volume is a synthesis of existing knowledge about the flora and fauna of Costa Rica. The major portion of the book consists of detailed accounts of agricultural species, vegetation, amphibians, reptiles, mammals, birds, and insects. This is an extraordinary, virtually unique work...The tremendous amount of original, previously unpublished, firsthand information is remarkable.--Peter H. Raven, Director, Missouri Botanical Garden An essential resource for anyone interested in tropical biology...It can be used both as an encyclopedia--a source of facts on specific organisms--and as a source of ideas and generalizations about tropical ecology.--Alan P. Smith, Ecology

Annals of the Missouri Botanical Garden

Annals of the Missouri Botanical Garden

Studies in Neotropical Paleobotany. III. The Tertiary Communities of Panama-Geology of the Pollen-Bearing Deposits

Samples containing fossil pollen and spores were collected from six sites representing five Tertiary formations in Panama. The Gatuncillo Formation is middle(?) to late Eocene, based on foraminifera and mollusk fossils. The formation is exposed around Madden Basin and samples were collected in the vicinity of Alcalde Diaz (Pefioncito). Material from the early Miocene Culebra Formation was obtained from a well drilled in the Contractor's Hill locality at the base of Gold Hill. The Cucaracha Formation overlies the Culebra and is also early Miocene in age. The Cucaracha material was collected from exposures along the Gaillard Cut section of the Canal. The early Miocene La Boca Formation outcrops in the same area. Although these three formations are similar in age (early Miocene), field evidence demonstrates that they are sequential from the Culebra (oldest), through the Cucaracha, to the La Boca (youngest). The Gatun Formation is late Miocene and early Pliocene in age and core samples were taken from a series of wells drilled in Gatun Lake. The location, stratigraphy, lithology, and depositional environments are summarized for these five formations. In subsequent publications the paleocommunities, paleoenvironments, and biogeographic implications will be considered for each of the microfloras. The present studies on the Cenozoic history of neotropical vegetation began in 1964 with a collection of pollen-bearing samples from Tertiary 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 Museum, and various institutions in Central America was rapidly providing new information on the composition of the flora, and on the range, ecology, and biogeographic relationships of its components. By comparison, the vegetation of more restricted areas to the north, such as Puerto Rico and Veracruz, was somewhat better known (Howard, 1973; Gomez-Pompa, 1973), and it seemed logical to begin paleofloristic studies on geologically younger deposits in these areas where modem analogs were more clearly defined. Consequently, our first studies were on the middle Oligocene San Sebastian palynoflora from Puerto Rico (Graham & Jarzen, 1969), followed by the late Miocene Paraje Solo palynoflora from Veracruz, Mexico (Graham, 1976). The decision to defer consideration of the material was fortunate for several reasons. The older Gatuncillo assemblage from proved the most difficult of our neotropical studies, and recently accumulated information on the modem vegetation was most helpful in making paleoenvironmental reconstructions. The completion 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 available to our studies on vegetational history. Recent biogeographic summaries are also valuable for interpreting the paleofloristic data (e.g., Gentry, 1982). Also, the pollen and spore reference 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 Canal and Vicinity, Republic of Panama (Stewart et al., 1 980) and modem plate tectonic summaries (e.g., Coney, 1982; Raven & Axelrod, 1974) are important for placing the paleobotanical data in a sound geologic context. Pollen-bearing samples from five Tertiary formations were collected in within or adjacent to the former Canal Zone (Figs. 1-3). These I Supported by NSF grants DEB-8205926, DEB-8007312, and GB-1 1862. 2 Department of Biological Sciences, Kent State University, Kent, Ohio 44242. 3Chief Geologists (retired), Canal Commission. Present address: 204 Tampa Downs Blvd., Lutz, Florida 33549. ANNm. MISSOURI BOT. GARD. 72: 485-503. 1985. This content downloaded from 157.55.39.104 on Mon, 20 Jun 2016 06:29:04 UTC All use subject to http://about.jstor.org/terms 486 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 72

Pollen Morphology of the Genus Erythrina (Leguminosae: Papilionoideae) in Relation to Floral Structure and Pollinators

Pollen Morphology of the Genus Erythrina (Leguminosae: Papilionoideae) in Relation to Floral Structure and Pollinators

IUCN—WWF Plant Advisory Group, First Meeting, held in the Missouri Botanical Garden, St Louis, Missouri, USA, during 17–18 December 1984

IUCN—WWF Plant Advisory Group, First Meeting, held in the Missouri Botanical Garden, St Louis, Missouri, USA, during 17–18 December 1984

Miocene Fossil Grasses: Possible Adaptation in Reproductive Bracts (Lemma and Palea)

Fossil reproductive bracts of Berriochloa, Nassella, and Paleoeriocoma of the tribe Stipeae, Archaeoleersia of the tribe Oryzeae, and Panicum of the tribe Paniceae (Gramineae) were collected from late Oligocene-Miocene strata in central North America, examined, and compared with modem taxa. One objective of these studies was to elucidate the evolution and possible adaptive significance of bract features. Berriochloa, the oldest known grass, appears in late Oligocene-early Miocene strata, where it has an indurate, cylindrical anthoecium (lemma and palea collectively) and an elongated, pointed callus. From early Miocene forms of Berriochloa two mid-late Miocene lineages evolved, one retaining a cylindrical anthoecium and the other developing a prominently inflated anthoecium. Both lineages retained the elongated, pointed callus. Paleoeriocoma, Nassella, Archaeoleersia, and Panicum first appear in the late Miocene with anthoecia similar to those of living taxa. The indurate anthoecium of the fossil and related living grasses probably evolved primarily as an adaptation to mammals and insects that use grasses as food. Related living taxa show adaptive mechanisms similar to those of the fossils, although some features such as the strongly interlocking bracts and blunt callus of some species of Piptochaetium are a post Miocene development. The Tertiary deposits of central North American Plains have produced rich and varied floras of considerable paleobotanical interest (Stansbury, 1852; Engelmann, 1876; Cockerell, 1914; Berry, 1928; Elias, 1931, 1932, 1934, 1935, 1942; Frye et al., 1956, 1978; Leonard, 1958; Leonard F Frye L Segal, 1965, 1966a, 1966b, 1966c; Skinner et al., 1968; Galbreath, 1974; Diffendal et al., 1982; Voorhies & Thomasson, 1979). Grasses in these floras are found as silicified reproductive bracts or as remains of leaves, stems, and roots and frequently exhibit detailed epidermal features that aid in determining phylogenetic relationships of fossils and their modem counterparts (Thomasson, 1976, 1977, 1978, 1979, 1980a, 1980b, 1984). Fossils described include reproductive bracts assigned to Archaeoleersia of the tribe 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 (lemma and palea) or bract (lemma) enclosing the mature grain or caryopsis of fossil grasses and their related living taxa. It reviews the geologic and paleoecologic background, summarizes the morphologic features of the fossil and living grasses, examines evolutionary trends among the fossil grasses, and speculates 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 widespread 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 diatomites, and volcanic ashes and vary in thickness in individual sections from more than 200 m to 1 m. Although some early studies suggested rather simple models of deposition and biostratigraphy, more recent investigations have demonstrated their depositional and biostratigraphic complexity (Bart, 1975; Breyer, 1976, 1981; Skinner et al., 1968, 1977; Swinehart, 1979; Thomasson, 1979; Diffendal, 1980, 1982). Among the most richly fossiliferous Tertiary strata in the world, they contain assemblages of 1 This paper is one result of some studies that have been supported by the National Geographic Society (188379, 2197-80) and the National Science Foundation (DEB-7809150, DEB-800228 1, DEB-820468 1, CDP-8000918, and R1 1-8213915). I thank numerous colleagues and assistants who contributed their time and expertise in the field during these studies and N. Morin, P. Palmer, and anonymous reviewers for helpful suggestions that improved this paper. 2 Department of Biological Sciences, Fort Hays State University, Hays, Kansas 67601-4099. ANN. MISSOURI BOT. GARD. 72: 843-851. 1985. This content downloaded from 157.55.39.215 on Sun, 24 Jul 2016 04:34:41 UTC All use subject to http://about.jstor.org/terms 844 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 72 TABLE 1. Stratigraphic relationships and ages of late Tertiary strata and study sites in the plains of central

Phylogenetic Patterns and Hybridization

Hybridization is an important part of the evolutionary history of flowering plants. If hybridization has occurred among 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 hybridization 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 polyploidy in evolution because they are outstanding features of many plant groups. According to some authorities 30-80% of 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 do not include diploid hybrids. Despite its importance, hybridization has been virtually ignored by those who have dominated the discussion 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 hybridization have a strong relationship with unisexuality and are not considered important in evolution. There are different types of hybridization. Figure 1 summarizes some of the possibilities (a more detailed explanation is found in Funk, 1981) but does not include introgression. For the purpose of this paper it is important to note that many hybrids are sexually reproducing individuals and are morphologically distinct and in some manner reproductively isolated from both parents. 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 pattern or hierarchy. The method of cladistics (phylogenetic systematics) seeks to discover these patterns by grouping together taxa that share apomorphies (evolutionarily novel, unique, or derived characters). Hybridization, or reticulate evolution, is inconsistent with a method designed to depict hierarchies. Hybridization is, therefore, a cause of incongruent, intersecting data that obscure phylogenetic information. Cladists have been concerned with this problem for several 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 hybridization and phylogenetic systematics include Bremer (1983), Bremer and Wanntorp (1977), Clark (1982), Funk (1981), Humphries (1983), Humphries and Funk (1984), Nelson (1983), Nelson and Platnick (1980), Rosen (1979), Wagner (1969, 1983), Wanntorp (1983), and Wiley and Brooks (1982). One favorite method 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): reproductive compatibility is a primitive attribute for the members of a lineage and has, therefore, no power to specify relationships within a genealogical framework. We cannot use the ability of two or more species to hybridize as an indication of close relationship because the ability is relaI A number of people, who do not necessarily agree with everything that I have said, have kindly provided me with data and comments on various drafts of the manuscript and this paper would not have been possible without their assistance. They include: R. Jansen, J. Semple, C. Clark, R. Sanders, C. Humphries, H.-E. Wanntorp, K. Bremer, G. Nelson, N. Platnick, D. Rosen, and P. Weston. I appreciate the assistance of B. Kahn in helping prepare the illustrations. 2 Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. ANN. MISSOURI BOT. GARD. 72: 681-715. 1985. This content downloaded from 207.46.13.148 on Sun, 11 Sep 2016 04:15:12 UTC All use subject to http://about.jstor.org/terms 682 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 72