Book Reviews: Systematics

Guide to Colorado Wildflowers, volume 2, Mountains. G. K. Guennel. 1995. ISBN 1-56579-119-3. (paper US$24.95) 352 pp. Westcliffe Publishers, Inc., P.O.Box 1261, Englewood, CO 80150.
At first glance the watercolor sketches of each plant catch your eye. You think they are pleasant but not good enough for identification. Then you notice the accompanying excellent color photograph close-ups of flower and other plant details and you realize the two kinds of illustration complement each other well. The paintings give a good sense of the plant as a whole and the color photographs give the necessary details. This combination makes the guide unusual. It also makes you realize the immense amount of time and field work that went into the book since Dr. Guennel was author, photographer and artist. Most field guides divide these tasks among two or three people. It should be a useful book in the field.

There are some 3000 flowering plants in the Colorado flora. From these the author selects, for his two volumes, about 600 species which "the average person might find". Volume 1 covers plants of the Colorado plains and foothills (from 3500 to 8000 feet elevation), and this volume under review includes plants found above 8000 feet in the montane, subalpine, and alpine zones. As the author admits in his preface he interprets "wildflowers" broadly, and includes a few grasses and sedges and some trees and shrubs as well as wildflowers per se.

The plant descriptions are brief but include the common name, scientific name and some synonyms, the plant family, plant kind, size, flowers, fruits, leaves, habitat, flowering time and life zones where found. In the introduction the author explains the book arrangement. "I grouped the plants by color and then I split off trees/shrubs from herbs. Lastly I listed the plants alphabetically by family, and within each family alphabetically by common name." He expands upon this in the text along with a short discussion of Colorado life zones. The book includes a glossary, indexes of common and scientific names, and a bibliography.

It is difficult to produce a really comprehensive field guide to plants of a region. In the Northeast we are fortunate in having two: the Peterson and the Newcomb wildflower guides. Both eliminate grasses and sedges and therefore cover a flora of about 3000 taxa. Each illustrates over 1200 plants, a ratio of about one plant in three. There are several wildflower guides for the Rocky Mountain region, some old, some new. With a flora of probably 4000 taxa even after eliminating grasses, sedges, and trees, these books are all highly selective, and most achieve a coverage ratio of only one to six or seven. For example in the venerable Peterson series: A Field Guide to Rocky Mountain Wildflowers by the Craigheads covers 600 plants and only half are illustrated. The classic Handbook of Rocky Mountain Plants by Ruth Ashton Nelson, 2nd. ed., keys out 970 plants with one third illustrated. Later editions may include more. The recent handsomely illustrated, but slim, three volume set by Dr. Dee Strickler, Prairie Wildflowers, Forest Wildflowers and Alpine Wildflowers together only cover 400 species, as does the equally well illustrated Alpine Wildflowers of the Rocky Mountains by Joseph Duft. In his introduction Guennel says he used more than fifty books to identify Coloradoís plants. He hoped to reduce this need with his new volumes. Their coverage ratio of perhaps one in three is an improvement but I do not think they will eliminate the need for several companion field guides to identify all the plants you are likely to meet in an excursion from the Colorado foothills to Trail Ridge Road. Time will tell. Mary M. Walker, New England Wild Flower Society Library, Framingham, MA.

Niebla and Vermilacinia (Ramalinaceae) from California and Baja California. Spjut, R.W., 1996. ISSN 0833-1475 (paper, no price given) 208 pp. Sida, Botanica Miscellany: 14. Botanical Research Institute of Texas, Inc. The Californian lichen flora comprises over 1000 taxa, of which many, such as species of Niebla sensu lato occur mainly in coastal regions. These fruticose species colonize soil, rocks and trees and often form a conspicuous component of the vegetation. Within the Ramalinaceae, Niebla sensu lato is distinguished by its cortical anatomy. Prior to 1970, Niebla was assumed to be a rather small genus comprising only four species in North America and Mexico. By 1996, 14 new taxa had been added, of which three are endemic to Mexico. Seven of the new species were described in 1994, nearly doubling the number of species for the genus in North America and Baja California. In 1995, Spjut established the genus Vermilacinia to accommodate taxa that lack a chondroid strand, and whose chemical profiles include mainly terpenes. In this publication, he has now presented a revised taxonomic treatment of the Niebla complex in California and Baja California.

Spjut started this systematic study in the mid 1980s and collected more than 1000 specimens of the Niebla complex in the following ten years. The Niebla complex in California and Baja California is here considered to consist of seventy-two taxa (including one undescribed species), organized into two genera Niebla and Vermilacinia with the latter genus divided into two subgenera. Fifty-three of these taxa (74%) are new to science: 36 species and one variety in Niebla and 16 species in Vermilacinia. An overview of the taxonomy of Niebla, Vermilacinia and Ramalina opens the systematic portion of this publication. For each genus, Spjut discusses the species concepts applied prior to presenting the character-state concepts he used. Species descriptions are preceded by a discussion on the variation in morphological and chemical characters in both genera and on the species concepts applied within each genus. Two keys to all 72 taxa are provided. The first key emphasizes morphological characters and uses secondary chemistry mainly to distinguish species. The alternative key relies primarily on chemical attributes derived from thin layer chromatography to define species groups which are then distinguished based on morphological features. The actual species descriptions occupy the next 136 pages. Species are arranged alphabetically within each genus or subgenus. For each species, a description of morphological and chemical characters is presented, and the variation and diagnostic characters are briefly discussed. Type material was examined for all previously published species except for V. procera, V. tuberculata, and V. tigrina. The treatment is richly illustrated and includes a) 11 introductory plates of color photographs of general habitats and species habit, b) illustrations accompanying the keys and 3) black and white photographs of individual species that complement the species descriptions. The photographs are of good quality and appear to be correctly labeled (except for plate 11 C & D). By contrast the drawings, prepared by four different authors, are of rather poor quality, ranging from mere pencil sketches to ink drawings with discontinuous outlines and excessive or inadequate shading. The distribution of the species is briefly described and represented by a list of selected collections; unfortunately maps for individual distributions are not provided. A glossary to the terminology mainly used in the species description and discussion completes this publication.

The taxonomy of Niebla relies on both chemical and morphological characters. Secondary metabolites are primary characters, in the sense that they define species groups. The 42 species of Niebla are arranged into 11 chemical groups of which the three main ones are: the divaricatic acid group (22 species), the sekikaic acid group (10) and salazinic acid group (6). Within these groups, species are differentiated by morphological characters. These morphospecies thus represent morphotypes within a chemical group. Although this approach is compared by the author to a phylogenetic species concept within which chemical characters serve as diagnostic features, a formal discussion of relationships among the species is not given, nor is a discussion on the metabolic relationships among these chemicals provided in support of their taxonomic or phylogenetic use. It is thus not clear whether species within a chemical group are indeed more closely related to each other than are similar morphotypes with distinct chemical profiles. Monophyly of chemical groups is tentatively supported by Spjut by such characters as erect free branches, strong yellowish-pigmented holdfast (in the depsides group), versus interwoven branches and no colored pigmentation of the cortex (depsidone-group). Within each of these two groups, however, morphological differentiation between chemically related sub-groups is however less clear. Morphological characters are not only very variable and thus overlapping between groups, but also very plastic within the chemical groups as morphological differences supporting the taxonomic distinction of some species disappear when these species are growing together. Chemically related species also tend to have similar preferences for substrate-type; how the various substrate types actually differ is not made very clear, however. In summary, morphologically similar but chemically and thus ecologically distinct populations of Niebla are recognized as distinct species. The alternative hypothesis, that species vary morphologically and chemically along an ecological "gradient" is rejected mainly because of its impracticality: morphological character variation could not be broken down into "discrete" classes without the integration of chemical characters. Nevertheless, morphological characters may be indicative of phylogenetic relationships across chemical groupings: "Many species of Niebla within a chemical subgroup appear to have siblings in other chemical subgroups, but morphological differences within species or related chemo-complexes are not parallel." "Not parallel" does not, however, exclude the possibility that morphological variation in species may be overlapping, and may thus represent tendencies correlated with chemical variation. Excessive splitting of morphosyndromes into morphospecies certainly leads to species that are so cryptic (defined by narrow classes of morphological character variation) that addressing relationships based on morphology becomes nearly impossible. Whether these morphotypes should have been recognized at an infraspecific rather than a specific level is uncertain, and subjective in the absence of experimental data defining the species limits in a biological, ecological or phylogenetic scenario. What is needed, and is lacking in this treatment, is a synopsis of the species, with a clear proposal of species relationships within or between chemical or morphological groups.

Relationships among species of Vermilacinia appear much clearer than in Niebla, as they are formally arranged into two subgenera that are best defined by cortical and medullary characters. Eighteen saxicolous species compose the type subgenus in the studied area, whereas subgenus Cylindricaria includes 20 primarily corticolous species. Most species are defined by a particular morphotype rather than a chemotype, and thus a species of Vermilacinia can be represented by a chemosyndrome (a suite of chemotypes). However, morphological intergradation still remains a problem (e.g., among V. polymorpha, V. procera, V. paleoderma and V. reptilioderma) and some species can be identified reliably only by their chemical profile (V. reptilioderma and V. paleoderma are here considered sibling species differing mainly in the nature of their terpenoids).

With this publication, the Niebla complex has moved from obscurity to the forefront of lichenology in western North America. Indeed, with 42 species to its name, Niebla has become one of the largest macrolichen genera in North America including Baja California. Certainly the author deserves credit for his detailed observations, and extensive chemical studies. Whether the taxonomic conclusions drawn from these are biologically, ecologically or phylogenetically meaningful remains to be tested; one earnestly hopes the required studies will be undertaken prior to the description of more taxa in this group. Bernard Goffinet, Department of Botany, Duke University, Durham, NC, 27708

The Biology of Grasses. Chapman, G.P., 1996. ISBN 0-85199-111-4 (cloth US$85) 273 pp. CAB International, 198 Madison Avenue, New York NY 10016 As a botanist who simply wanted to know more about grasses, I enjoyed this book immensely. In fact, I devoured it. Chapman has a lively, delightful writing style that incorporates metaphor as well as current issues in science and the news. The book surveys grass utility and importance, morphology and development, ecology, reproduction, mating systems, taxonomy, biogeography, and evolution.

The two major themes of the book are 1) the immense morphologic, physiological and ecological diversity found at every taxonomic level in the grass family, and 2) the implications of this diversity to grass taxonomy and phylogeny. Chapman favors the view that the subfamily Bambusoideae, which includes bamboo and rice, is phylogenetically most basal and illustrates nearly every chapter with some example from the bamboos. Chapman also seems to have a fondness for the unusual and exceptional. While this is one of the characteristics that makes the book so delightful, students may need to be reminded of the rarity of such exceptions.

However, some material that one would expect to find in a book titled "The Biology of Grasses" is conspicuously absent. Chapman makes no apologies for this. The back cover calls this book a bridge between introductory texts and technical papers. Chapman explicitly states in the Preface that his intent was not to duplicate material in any of his other well-known grass texts. He calls "The Biology of Grasses" an "individual perspective," a reaction to the abundant grass literature he is expert in, a chance to make connections between narrow fields within grass biology and draw attention to unexplored territories. As such, its coverage of some areas seems shallow while other topics are not included at all. His description of embryo and leaf development is accurate but brief. Inflorescence and floret development is never mentioned. Photosynthesis is discussed only in the context of its mechanistic diversity and the insights it provides into evolutionary relationships. Except for a mention of secondary metabolites, these is no further discussion of grass physiology or biochemistry what so ever. The most glaring omission is the huge amount of research into carbohydrate storage and partitioning in grasses. In some of these areas, Chapman quickly refers the reader to the appropriate reviews and then moves on.

Though there are superb line drawings of grasses to illustrate subfamilies and some of the specifically discussedgrass species, illustrations are missing at other points where they would be helpful, such as the discussion of embryo and leaf development, and the involvement of Tripsacum and teosinte in the evolution of maize. His written descriptions are fabulous and conjured up accurate pictures of plants and their organs in my mind, but I am a plant morphologist. A student may need the extra help of a drawing. His final chapter on maize domestication is quite evocative and very nicely summarizes the competing hypotheses about maize evolution without reference to any of the difficult molecular data on the subject. Botanical and grass-specific terms are used without definition throughout the text. This facilitates the flow of reading and is appropriate to an intermediate-level text, as students can refer to the excellent glossary.

Because of the absence of discussion of carbohydrate storage and partitioning, I would not recommend this book for use in classes with an agronomic focus. However, "The Biology of Grasses" would make a good companion text to a taxonomy or grass systematics manual in an agrostology or grass systematics class. It introduces students to the diversity, value and wonder of grasses, and provides an introduction into how traits other than conventional morphologic and molecular data, such as ecological function, geographic distribution and physiology, can be useful in phylogenetic reconstruction. Rebecca A. Sherry, Division of Biological Sciences, University of Missouri, Columbia

Rare Lilies of California. Fiedler, P. L. 1996. ISBN 0-943460-30-1 (paper) California Native Plant Society, Sacramento, California As a graduate student back at San Francisco State University, I was a teaching assistant for Peggy Fiedler in a non-majors economic botany course. Some of the students concocted a vignette to compare me, their enthusiastic but gritty lab instructor, to Peggy, a polished lecturer and accomplished botanist. It went something like this: In a garden of little people (the students), there lived a gentle fairy princess (the lecturer), and an evil gnome (guess who). Impressions like this, though colorful, are not entirely accurate, at least not always predictive. One of us may remain gnomish, but the other has gone on to put together a solid, quite unfairy-like piece of work in conservation biology.

Rare Lilies is a book by a botanist with a mission. Fiedler sets out to "document and articulate" rarity in the diverse and beautiful California Liliaceae, a large plant family with 34 native genera in California and dozens of species found only in the state. She documents her topic with an appropriate mix of prose, tables, and graphics. She articulates her work to readers through a series of gorgeous full color plates. Fiedler and her illustrator, Catherine M. Watters, combine visual poetry with hard science in examples of species that demonstrate the beauty and the evolutionary complexity of the California lilies.

This is a most attractive book, accessible to non-specialists and useful as well for scientists. Fiedler presents hard questions in soft sentences like, "What makes a lily a lily?" and "What does it mean to be rare?" By asking these questions, she challenges the reader to puzzle out answers for difficult topics such as endemism and relictualism, and their role in an evolutionary framework. More questions go unanswered than resolved as Fiedler outlines the causes and consequences of rarity but significantly, we are invited to construct a model that goes beyond the organisms she studies. We learn about more than lily biogeography and conservation here.

The book has broad appeal for students in many areas of plant biology. Its contents will surely find their way into new textbooks that touch on the subject of floral variation and evolution, patterns of endemism and rarity, and monocot biology and taxonomy. The excellent work of Randy Zebell, which Fiedler highlights throughout the book, was done at SF State while he was struggling to support himself as a graduate student, driving a cab at night. Rare Lilies of California is a testament to the continued ability of the California State University system to produce first rate scientific research as well as accomplished students. It builds on the strong tradition of collaboration between the California Native Plant Society and the academic community. It also illustrates the great and ongoing potential of California as a hotbed of plant evolution. Samuel Hammer, College of General Studies, Boston University

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