Santa Ana Botanic Garden Research Articles

Flavonoid Chemistry and the Phylogenetic Relationships of the Julianiaceae

The phylogenetic affinities of the Julianiaceae were studied by a survey of the leaf and heartwood flavonoids of Amphipterygium adstrigens as compared to those of 40 representative species from 29 genera and ten families, particularly the Anacardiaceae, considered to be closely related to the Julianiaceae. Sixteen flavonoids, including seven 5-deoxyflavonoids, were detected in A. adstrigens. A ll the 5-deoxyf lavonoids were also present in all anacardiaceous species surveyed, and some of the compounds (e.g., sulphuretin, fisetin, and fustin) have been considered characteristic of the Anacardiaceae as a whole. Rengasin and a suphuretin glucoside (aurones) and a butein glucoside (chalcone) are known only from A. adstrigens and from the Anacardiaceae. These unusual 5-deoxyflavonoids were not detected in any of the other families surveyed. The flavonoid constituents of A. adstrigens are most similar to those of anacardiaceous tribe Rhoeae. Evidence from the wood anatomy-anomalous reticulate perforation plates found only in the julianiaceous genera Amphipterygium and Orthopterygium and in the Rhoeae-also shows the similarity of these groups. It is proposed that the Julianiaceae be treated as a subtribe (Julianiinae) of the Rhoeae. The Julianiaceae are a small family of flowering plants consisting of ti monotypic genus Orthopterygium Hemsley (0. huaucui (A. Gray) Hemsley) of Peru and Amphipterygium Schiede (= Juliania Schlechtendal) with four species (Standley, 1923) of southern Mexico and Guatemala. Amphipterygium and Orthopterygium are trees characteristic of the tropical deciduous forests of these regions. The phylogenetic relationships of the Julianiaceae have been the subject of controversy for the past century. The Julianiaceae are wind pollinated and exhibit all the features characteristic of this pollination system (e.g., apetaly, male flowers in a loose catkin), and historically the family has been placed in the Amentiferae. Recently the controversy was revived in a series of papers on the Amentiferae (Mears, 1973; Moseley, 1973; Stern, 1973; Stone, 1973; Thorne, 1973; Wolfe, 1973). The purpose of this study was to investigate the 1 Most of this study was performed while I was a graduate student at Claremont Graduate School-Rancho Santa Ana Botanic Garden, California. I thank Sherwin Carlquist, C. E. Jones, R. Ornduff, Ron Scogin, and Robert Thorne for their help. Field studies were supported by NSF grant GB-38327. Thanks are also due the Consejo Nacional de Ciencia y Technologia, Mexico, for a collecting permit and the Fairchild Tropical Garden for leaf material of A. adstrigens. 2 Botany, University of Illinois at Champaign-UTrbana, Urbana, IL 61801. This content downloaded from 207.46.13.43 on Thu, 14 Apr 2016 06:18:22 UTC All use subject to http://about.jstor.org/terms 150 SYSTEMATIC BOTANY [Volume I

Adaptive Aspects of Insular Evolution, A Symposium

ADAPTIVE ASPECTS OF INSULAR EVOLUTION was the theme chosen by the Association for Tropical Biology for its third international symposium, June 15-19, 1969. Appropriately enough, the symposium was held on a tropical island: at the University of Puerto Rico, Mayagiiez, where over 100 persons convened to hear presentations by an outstanding group of 17 participants assembled by Professor Ernst Mayr, Director of Harvard University's Museum of Comparative Zoology. The tone of the conference was set most effectively when Sherwin Carlquist of the Rancho Santa Ana Botanic Garden and Terrell H. Hamilton of the University of Texas broadly outlined the more important factors responsible for the characteristics of island floras and faunas, respectively. Their introductions re-emphasized how very suitable insular situations are for detailed studies of evolutionary processes, including dispersal, colonization, adaptation, competition, and extinction. With a refreshing balance of botanical and zoological viewpoints, the speakers who followed discussed various aspects of these topics and provoked considerable thoughtful and spirited debate. One of the central topics of the conference was the delimitation of some of the more important genetic and ecological characteristics of successful colonizing species. A recurrent theme was that an insular situation, be it an island or a mountain, may be remote for some species and at the same time very near for others, depending upon their ecological requirements. For certain plants, G. Ledyard Stebbins of the University of California, Davis, found that polyploidy apparently facilitates island colonization. Tremendous quantities of highly dispersible spore propagules were most responsible for the colonizing success of ferns, according to Rolla M. Tryon of Harvard University; he felt this high vagility largely accounted for the relatively low levels of endemicity characteristic of insular fern floras. Based on chromosome banding patterns, the report by Hampton L. Carson of Washington University centered upon certain genetic properties of successful species founders and subspecies colonizers in the genus Drosophila on the Hawaiian Islands. Ernest E. Williams of Harvard IJniversity reviewed his data on the invasion patterns of West Indian Anolis lizards and stressed the relation between the number of ecomorphs occurring on a given island and the structural habitats present. Carabid beetle species most successful in colonizing the West Indies were characterized by Philip J. Darlington, Jr., of Harvard University as being small, winged, and frequently associated with fresh water. Many of the talks focused directly or indirectly on the types of adaptive adjustments made to insular conditions by various floral and faunal elements following successful colonization; effects of impoverishment upon adaptive radiation, niche dimensions, and ecological and behavioral shifts were recurrent topics. Among those presenting new experimental data bearing on these phenomena were Allen Keast of Queens University, speaking on Tasmanian birds, Allen J. Kohn of the University of Washington on the carnivorous marine snail Conus, and Thomas Schoener of Harvard University on West Indian Anolis lizards. The composition, adaptations, and relationships of three very different insular floral communities were the principal subjects of Olov Hedberg of the University of Uppsala, Richard A. Howard of Harvard University, and Bassett Maguire of the New York Botanical Garden. Hedberg characterized the alpine flora of African mountains with particular reference to the rather striking adaptations to the extreme daily temperature fluctuations. Howard focused on the dwarf elfin forest of the higher elevations of Puerto Rico and the possible factors responsible for this peculiar floristic

The California Islands

The California Islands off the Pacific coast of southern and Baja California display almost diagrammatically the insular phenomena that characterize fringing archipelagos. To a lesser extent they also illustrate some of the insular features associated with oceanic islands. Several of these islands have been much studied and are relatively accessible. The location, size, elevation, and geological history of the islands are described. To determine the degree of disharmony of the island floras, they are analyzed in comparison with selected California mainland floras. The degree of richness of the island floras is discussed in relation to size of the islands, proximity to the mainland, and other variables. Such phenomena of island life as the immigrant pattern of vertebrate distribution, relict versus autochthonous endemism of plants and animals, insular gigantism, continuous flowering, high degree of hybridization, vulnerability, and loss of dispersibility in island species are considered in reference to the California Islands. Some suggestions as to available facilities on the islands and persons or institutions to contact are given in an epilogue. Off the Pacific coast of southern California and Baja California, Mexico, lie the islands variously called the Channel Islands or California Offshore Islands and the Baja California Islands or, collectively, the California Islands. Although they are in no sense tropical, they are worthy of mention in this symposium because they are a good example of a fringing archipelago and because they illustrate many of the insular phenomena that one might otherwise have to travel far to tropical areas to study. Some of them are quite accessible from the heavily populated areas of southern California, and some of them have been rather thoroughly studied by biologists. Although they are, thus, rather well known to California biologists, they are surprisingly little known to non-Californians. It is hoped that this short discussion of them and of some of their insular characteristics will create more interest in them among those biologists concerned with evolutionary and environmental phenomena. The Southern California and Baja California Islands consist of 16 major islands or island groups spread over some 500 miles between Point Conception, California, and Punta Eugenia, Baja California, Mexico (Fig. 1). From east to west the Northern Channel Islands are Anacapa, Santa Cruz, Santa Rosa, and San Miguel. The Southern Channel Islands are San Nicolas, Santa Barbara, Santa Catalina, and San Clemente Islands. The Baja California Islands include Los Coronados just south of San Diego, Todos Santos off Ensenada, San Martin off San Quintin, San Geronimo south of El Rosario, the San Benitos, Cedros, and Natividad off Punta Eugenia, and Guadalupe 157 miles off the Baja California peninsula. In order of size, with area in square miles, the islands are: Cedros (134), Guadalupe (98), Santa Cruz (96), Santa Rosa (84), Santa Catalina (75), San 11 am grateful to Dr. Reid Moran for critically reviewing the manuscript, for making many valuable suggestions, and for supplying useful unpublished information. 2Rancho Santa Ana Botanic Garden, Claremont, California 91711. ANN. MISSOURI BOT. GARD. 56: 391-408. 1969. This content downloaded from 157.55.39.247 on Wed, 27 Apr 2016 05:48:03 UTC All use subject to http://about.jstor.org/terms [VOL. 5 6 392 ANNALS OF THE MISSOURI BOTANICAL GARDEN Clemente (56), San Nicolas (22), San Miguel (14), Natividad (2.8), San Benito (2.5), Anacapa (1.1), Santa Barbara (1.0), Los Coronados (1.0), San Martin (0.9), Todos Santos (0.5), and San Geronimo (0.2). Their distances from the mainland, in statute miles, are: San Martin (3), Todos Santos (4), Natividad (5), San Geronimo (6), Los Coronados (8), Anacapa (13), Cedros (14), Santa Cruz (19), Santa Catalina (20), San Miguel (26), Santa Rosa (27), Santa Barbara (38), San Benito (41), San Clemente (49), San Nicolas (61), and Guadalupe (157). All these figures are from a summary by Philbrick (1967). The highest elevations on the largest islands, in feet, are: Guadalupe (4,257), Cedros (3,950), Santa Cruz (2,165), Santa Catalina (2,069), San Clemente (1,965), Santa Rosa (1,574), and San Nicolas (907).

THE BIOTA OF LONG-DISTANCE DISPERSAL. II. LOSS OF DISPERSIBILITY IN PACIFIC COMPOSITAE.

EvolutionVolume 20, Issue 1 p. 30-48 ArticleFree Access THE BIOTA OF LONG-DISTANCE DISPERSAL. II. LOSS OF DISPERSIBILITY IN PACIFIC COMPOSITAE Sherwin Carlquist, Sherwin Carlquist Claremont Graduate School, Rancho Santa Ana Botanic Garden, Claremont, CaliforniaSearch for more papers by this author Sherwin Carlquist, Sherwin Carlquist Claremont Graduate School, Rancho Santa Ana Botanic Garden, Claremont, CaliforniaSearch for more papers by this author First published: March 1966 https://doi.org/10.1111/j.1558-5646.1966.tb03341.xCitations: 58AboutPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinked InRedditWechat Citing Literature Volume20, Issue1March 1966Pages 30-48 ReferencesRelatedInformation

CHROMOSOME NUMBERS IN COMPOSITAE. II. HELENIEAE

Raven, Peter H. (Rancho Santa Ana Botanic Garden, Claremont, Calif.), and Donald W. Kyhos. Chromosome numbers in Compositae. II. Heleniae. Amer. Jour. Bot. 48(9): 842–850. Illus. 1961.—Chromosome counts are now available for 42 of the approximately 55 genera of Compositae, tribe Helenieae, which is predominantly a group of western North America. These chromosome numbers are summarized here at the generic level, and 100 original counts for the tribe are added, including what seem to be the first published reports for the genera Amblyopappus, Baeriopsis, Hulsea, Jaumea, Pericome, Rigiopappus, Trichoptilium, and Venegasia, as well as for many species. The phylogeny of Chaenactis is discussed in the light of published records and 46 original counts, and C. douglasii is shown to include plants in which n = 6, 12, and 18, which differ somewhat morphologically. Helenium has species which have a complete series of aneuploid numbers from n = 13 to n = 17. Chromosome numbers coincide with morphological variability in indicating that Helenieae are a diverse group. More detailed studies of various kinds will be necessary before the genera of Helenieae can be re‐aligned effectively, but it is evident that different genera show affinities with various other tribes of the family. Nevertheless, it is thought to be convenient to continue to recognize Helenieae at the tribal level for the present.

A California Flora . Philip A. Munz and David D. Keck. Published for the Rancho Santa Ana Botanic Garden by the University of California Press, Berkeley, 1959. 1681 pp. Illus. $11.50.

<i>A California Flora</i> . Philip A. Munz and David D. Keck. Published for the Rancho Santa Ana Botanic Garden by the University of California Press, Berkeley, 1959. 1681 pp. Illus. $11.50.

A California Flora. Philip A. Munz and David D. Keck. Published for the Rancho Santa Ana Botanic Garden by the University of California Press, Berkeley, 1959. 1681 pp. Illus. $11.50

A California Flora. Philip A. Munz and David D. Keck. Published for the Rancho Santa Ana Botanic Garden by the University of California Press, Berkeley, 1959. 1681 pp. Illus. $11.50

Glandular Structures of Holocarpha and Their Ontogeny

Carlquist, Sherwin. (Rancho Santa Ana Botanic Garden, Claremont, California.) Glandular structures of Holocarpha and their ontogeny. Amer. Jour. Bot. 46(4): 300–308. Illus. 1959.—Two types of advanced glandular structures occur in the 4 species of the genus Holocarpha. Sessile disk-shaped glands occur at the tips of upper leaves and of involucral and receptacular bracts. Unlike all other glandular structures of Madinae which have been investigated, these originate from several protodermal initials rather than a single one. These glands, however, represent modifications of a glandular trichome. The other type of glandular structure, termed hollow-stalked trichome here, occurs on the outer surface of involucral and receptacular bracts. These trichomes originate from a single cell but differ from others in the formation of a hollow stalk, the wall of which is one cell in thickness. Mesophyll of the bract, often with an included vascular bundle, is present as an intrusion into the base of the hollow stalk. Corresponding to the advanced nature of the glandular structures, the leaves show specializations in the “inrolling” of margins. Upper leaves have a cylindrical organization of vascular tissue, whereas basal leaves are “normal” and leaves of the main stem are intermediate. The species of Holocarpha differ in certain details of leaf anatomy and structure of hollow-stalked trichomes. The systematic distribution of these is given. The essential unity of the various glandular structures of Madinae is discussed both in terms of mature structure and ontogeny, and the steps in the evolution of these are suggested.

GLANDULAR STRUCTURES OF HOLOCARPHA AND THEIR ONTOGENY

Carlquist, Sherwin. (Rancho Santa Ana Botanic Garden, Claremont, California.) Glandular structures of Holocarpha and their ontogeny. Amer. Jour. Bot. 46(4): 300–308. Illus. 1959.—Two types of advanced glandular structures occur in the 4 species of the genus Holocarpha. Sessile disk‐shaped glands occur at the tips of upper leaves and of involucral and receptacular bracts. Unlike all other glandular structures of Madinae which have been investigated, these originate from several protodermal initials rather than a single one. These glands, however, represent modifications of a glandular trichome. The other type of glandular structure, termed hollow‐stalked trichome here, occurs on the outer surface of involucral and receptacular bracts. These trichomes originate from a single cell but differ from others in the formation of a hollow stalk, the wall of which is one cell in thickness. Mesophyll of the bract, often with an included vascular bundle, is present as an intrusion into the base of the hollow stalk. Corresponding to the advanced nature of the glandular structures, the leaves show specializations in the “inrolling” of margins. Upper leaves have a cylindrical organization of vascular tissue, whereas basal leaves are “normal” and leaves of the main stem are intermediate. The species of Holocarpha differ in certain details of leaf anatomy and structure of hollow‐stalked trichomes. The systematic distribution of these is given. The essential unity of the various glandular structures of Madinae is discussed both in terms of mature structure and ontogeny, and the steps in the evolution of these are suggested.

THE LEAF OF CALYCADENIA AND ITS GLANDULAR APPENDAGES

Carlquist, Sherwin. (Rancho Santa Ana Botanic Garden, Claremont, Calif.) The leaf of Calycadenia and its glandular appendages. Amer. Jour. Bot. 46(2) : 70‐80. Illus. 1959.—Large tack‐shaped glands are characteristic of the leaves of Calycadenia which are associated with the inflorescence. These glands may be divided into those which are terminal on leaves and those which occur laterally on the surface of the leaf. Lateral glands show stages early in their development which are identical with those of simpler trichomes of Madinae. Terminal glands, which possess more vascularization of the stalk, show a more modified form of development. Vascularization is not derived from protoderm, but from more deeply‐seated cells. These cells are included in a zone of elongation which forms the stalk. Vascular bundles may extend to the base of glands which lack vascularization in their stalks. Tack‐shaped glands are considered an advanced form of trichome in which internal tissues of the leaf are involved. Within the genus Calycadenia, ontogenetic and comparative studies suggest that the following characters are advanced: reduction to a single terminal gland, “inrolling” of margins to form a cylinder of bundles, concomitant with a central core of fibers or a pectic channel. Systematic distribution of gland occurrence and of types of foliar structure are given.

The Leaf of Calycadenia and its Glandular Appendages

Carlquist, Sherwin. (Rancho Santa Ana Botanic Garden, Claremont, Calif.) The leaf of Calycadenia and its glandular appendages. Amer. Jour. Bot. 46(2) : 70-80. Illus. 1959.—Large tack-shaped glands are characteristic of the leaves of Calycadenia which are associated with the inflorescence. These glands may be divided into those which are terminal on leaves and those which occur laterally on the surface of the leaf. Lateral glands show stages early in their development which are identical with those of simpler trichomes of Madinae. Terminal glands, which possess more vascularization of the stalk, show a more modified form of development. Vascularization is not derived from protoderm, but from more deeply-seated cells. These cells are included in a zone of elongation which forms the stalk. Vascular bundles may extend to the base of glands which lack vascularization in their stalks. Tack-shaped glands are considered an advanced form of trichome in which internal tissues of the leaf are involved. Within the genus Calycadenia, ontogenetic and comparative studies suggest that the following characters are advanced: reduction to a single terminal gland, “inrolling” of margins to form a cylinder of bundles, concomitant with a central core of fibers or a pectic channel. Systematic distribution of gland occurrence and of types of foliar structure are given.