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ANATOMY OF A ROSE -- EXPLORING THE SECRET LIFE OF FLOWERS

SELECTED BIBLIOGRAPHY AND NOTES

Flower in the crannied wall,
I pluck you out of the crannies;
Hold you here, root and all, in my hand,
Little flower -- but if I could understand
What you are, root and all, and all in all,
I should know what God is and man is.
-- LORD ALFRED TENNYSON

The Physics of Beauty

The information on Neanderthal burials was mainly taken from Arlette Leroi-Gourhan, ''The Flowers Found in Shanidar IV, a Neanderthal Burial in Iraq," Science 190 (November 1975).

The quote by Annie Dillard is from her collection of essays Teaching a Stone to Talk: Encounters and Expeditions (New York: Harper Collins Books, 1982).

The quote from Aldo Leopold is from his Sand County Almanac (New York: Oxford University Press, 1949).

Frederick Turner, Rebirth of Value: Meditations on Beauty, Ecology, Religion, and Education (State University of New York Press, 1991), has a longer list of the themes or tendencies of the universe. These are also explored in Brian Swimme and Thomas Berry, The Universe Story: From the Primordial Flaring Forth to the Ecozoic Era (San Francisco: Harper, 1992), as well as in other books and articles.

More about the giant arum can be found in textbooks and in Susan Milius, "The Science of Big, Weird Flowers," Science News 156 (September 11, 1999).

A college textbook I referred to frequently and recommend is Randy Moore et al., Botany (Wm. C. Brown, 1995). Their selection "Leonardo the Blockhead" summarizes the math behind a sunflower's seed spirals. This material is also known as the Fibonacci series.

Andy Coghlan, "Sensitive Flower," New Scientist, September 26, 1998, nicely summarizes recent work on how flowers "see," "smell," "touch," and "taste." Numerous others, such as Stephen Day, "The Sweet Smell of Death," New Scientist, September 7, 1996; Garry C. Whitelan and Paul E. Devlan, "Light Signaling in Arabidopsis," Plant Physiology Biochemistry 36 (1998), issue 1-2; and Paul Simons, "The Secret Feelings of Plants," New Scientist, October 17, 1992, discuss these subjects in depth.

Dagmar von Helversen and Otto von Helversen, "Acoustic Guide in a Bat-Pollinated Flower," Nature, April 29, 1999, give more information on how bats and flowers use sonar signals.

The Blind Voyeur

The man who lost color sight as a result of brain damage is described in Oliver Sacks, An Anthropologist on Mars (New York: Alfred Knopf, 1995).

Moore et al., Botany, provided me with a good description of the visible spectrum and the function of pigments in a flower petal.

Deni Brown, Alba: The Book of White Flowers (Portland, OR: Timber Press,1989), is a comprehensive discussion of white flowers and includes passages on how and why white flowers look white.

Rob Nicholson, "The Blackest Flower in the World," Natural History 108 (May 1999), is my source for the information on the Oaxacan flower.

Moore et al., Botany, includes a passage called "Why Plants Are Not Black."

The information on bees comes from many sources. Naturally I consulted the seminal works of Karl von Frisch, including his Bees: Their Vision, Chemical Senses, and Language (Ithaca, N.Y.: Cornell University Press, 1971); and The Dance Language and Orientation of Bees (Belknap Press, Cambridge, MA 1967).

A translated version of Georgii A. Mazokhin-Porshnyakov, Insect Vision (Plenum Press, New York, NY 1969), was helpful for background and a sense of history.

An excellent book that I often referred to is Friedrich G. Barth, Insects and Flowers: The Biology of a Partnership (Princeton, N.J.: Princeton University Press, 1991).

Another important source for the behavior and physiology of pollinators is Michael Proctor, Peter Yeo, and Andrew Lack, The Natural History of Pollination (Portland, OR: Timber Press, 1996).

Lars Chittka was another primary Source and a great help. Lars has pioneered much of the most recent research in insect vision, particularly in terms of what colors bees actually see. His most pertinent articles, related to this chapter, are Lars Chittka and Randolf Menzel, "The Evolutionary Adaptation of Flower Colours and the Insect Pollinators' Colour Vision," Journal of Comparative Physiology A 171 (1992 ); Lars Chittka, Avi Shmida, Nikolaus Troje, and Randolf Menzel, "Ultraviolet as a Component of Flower Reflections and the Colour Perception of Hymenoptera," Vision Resolution 34, no. II, p. 1489-1508 (1994); Lars Chittka and Nickolas Waser, "Why Red Flowers Are Not Invisible to Bees," Israel Journal of Plant Sciences 45 (1997); Peter Kevan, Martin Giurfa, and Lars Chittka, "Why Are There So Many and So Few White Flowers?" Trends in Plant Sciences I (August 1996); Lars Chittka, "Bee Color Vision Is Optimal for Coding Flower Color, but Flower Colors Are Not Optimal for Being Coded: Why?" Israel Journal of Plant Sciences 45 (1997); and Lars Chittka and Nickolas Waser, "Bedazzled by Flowers," Nature, August 27, 1998.

An earlier version of this chapter contained a longer version of why white flowers look bee-green and why green leaves look gray: "White flowers that reflect UV are actually rare. Most human-white flowers, like this daisy, absorb UV. They do not look bee-white, since they are not reflecting back the bee's entire spectrum. They are reflecting back blue and green and, to a bee, look blue-green. To a bee, the green, serrated leaves of a daisy probably look gray. In bee vision, green foliage has a weak and uniform reflectance, which makes it dull or uncolored. For humans, leaves absorb relatively more light in the red range."

The idea that bees had color vision before the appearance of flowers also comes from Lars Chittka's research, described in Kathleen Spiessbach, "The Eyes of Bees," Discover, September 1996. An example of Chittka's style and humor can be found in his article about flower color coding in Chittka, "Bee Color Vision": "But how can we determine in what colors insects saw the world 200 myr (million years) ago? Since time machine projects habitually run into complications (e.g., Wells, 1885), it is now difficult to obtain funding for them and so evolutionary biologists resort to an alternative strategy called comparative phylogenetic analysis."

Nickolas Waser was also an important source for this and other chapters. A related article is Nickolas Waser, Elvia Melendrez-Ackerman, and Diane Campbell, "Hummingbird Behavior and Mechanism of Selection on Flower Color in Ipomopsis," Ecology 78, no. 8 (1998).

I should also mention Beverly J. Glover and Cathie Martin, "The Role of Petal Shape and Pigmentation in Pollination Success in Antirrhinum majus," Heredity 80: 778-784 No.6, June 1998; Adrian Horridge, "Bees See Red," Trends in Ecology and Evolution 13 (March 1998); and A. C. Dyer, "The Color of Flowers in Spectrally Variable Illumination and Insect Pollinator Vision," Journal of Comparative Physiology A 183 (1998): 203-212 No. 2 August 1998.

Stephen L. Buchman and Cary Paul Nabhan, The Forgotten Pollinators (Washington, D.C.: Island Press, 1996), have a good discussion on the history and use of pollination syndromes.

Martha Weiss has done considerable work on the pollination behavior of butterflies. I used her research on swallowtail butterflies, taken from Martha Weiss, "Innate Color Preferences and Flexible Color Learning in the Pipevine Swallowtail," Animal Behavior 53 (1997): 1043-1052 no. 5.

Weiss also gave this quote in Susan Milius, "How Bright Is a Butterfly?" Science News 153 (April 11, 1998): "Honeybees are considered to be the intellectuals of the insect world." Earlier in the article, Milius writes that butterflies are too often "dismissed as too dumb to find their way out of a wet petunia."

Additionally, Weiss was my major source for information on how flowers change their color: Martha Weiss, "Floral Color Changes as Cues for Pollinators," Nature 354, November 1991; and Martha Weiss, "Floral Color Change: A Widespread Functional Convergence," American Journal of Botany 83, no. 2 (1995).

I should also mention Lynda F. Delph, "The Evolution of Floral Color Change Pollinator Attraction Versus Physiological Constraints in Fuchsia Excorticata," Evolution 43, no. 6 (1989).

Smelling Like a Rose

For this chapter, I am particularly indebted to two books: D. Michael Stoddart, The Scented Ape: The Biology and Culture of Human Odour (New York: Cambridge University Press, 1990); and Diane Ackerman, A Natural History of the Senses (New York: Random House, 1990). Stoddart discuss the desensitization of humans to their own smell, as well as the use of odor in human culture and perfumery. Ackerman continues and broadens that discussion.

Roman Kaiser, The Scent of Orchids: Olfactory and Chemical Investigations (Basel, Switzerland: Elsevier, 1993), was helpful to my understanding of how flowers produce scent, as were other books and articles. Rob Ragusa, "Floral Scent Production in Clarkia breweria," Plant Physiology 116 (1998): 599-604 no. 2, gave a specific example of floral emission.

My understanding of how animals pick up scent came primarily from the Konrad Colbow, ed., R. H. Wright Lectures on Insect Olfaction (Burnaby, B.C. Canada, Simon Fraser University, 1989); and T. L. Payne, M. C. Birch, and C. E. J. Kennedy, eds., Mechanisms in Insect Olfaction (Oxford, England: Clarendon Press, 1986).

B. S. Hansson, "Olfaction in Lepidoptera," Experientia 51 (1995), was also helpful.

The idea of flower constancy is discussed in many books. Nickolas Waser persisted in reminding me that flower constancy is an idea still being explored. I began with his "Flower Constancy: Definition, Cause and Measurement," American Naturalist 127 (May 1986), as well as his "The Adaptive Nature of Floral Traits, Ideas and Evidence," in Pollination Biology edited by Leslie Real (Orlando, Fl.: Academic Press, 1983).

Articles on how bees forage and smell include M. Giurfa, J. Nunez, and W. Backhaus, "Odour and Colour Information in the Foraging Choice Behavior of the Honeybee," Journal of Comparative Physiology A 175 (1994): 773-779; Martin Hammer and Randolf Menzel, "Learning and Memory in the Honeybee," Journal of Neuroscience 15 (March 1995); and B. Gerber et al., "Honey Bees Transfer Olfactory Memories Established During Flower Visits to a Proboscis Extension Paradigm in the Laboratory," Animal Behavior 52, 1079-1085 no. 6, (1996).

For information on worldwide agriculture, I referred to Collier's Encyclopedia, s.v. "Agriculture." Vol. 21 (out of 24), New York: P.F. Collier, 1984.

Concerning the interplay of sex and food, as well as other passages, I was much helped by Elizabeth A. Bernays, ed., Insect-Plant Interactions, vol. 5 (Boca Raton, Fl.: CRC Press, 1994). Specifically I consulted the long and comprehensive chapter by H. Dobson, "Floral Volatiles in Insect Biology."

The tidbit that moth pheromones and elephant pheromones have similarities is from Stephen Day, ''The Sweet Scent of Death," New Scientist, September 7, 1996, and comes from the research done by Bers Rasmussen at the Oregon Graduate Institute of Science and Technology.

Stoddart, The Scented Ape, discusses the experiment with women and musk and includes more information on those flower-based compounds that resemble human steroids.

Information on the giant arum and dead horse arum can be found in many books, including David Attenborough, The Private Life of Plants: A Natural History of Plant Behavior ( Boston: Compass Press, 1995). Another source for general information is Bastiaan Meeuse and Sean Morris, The Sex Life of Plants (New York: Faber Publishers, NY, 1984). These books are also good sources for the variety of flowers that smell like fungi, female wasps, and the like. The story of the drag-queen flowers is also told in these books and in many others. In addition, I consulted articles such as Marlies Sazima et al., ''The Perfume Flowers of Cyphomandra (Solanaceae): Pollination by Euglossine Bees, Bellows Mechanism, Osmophores, and Volatiles," Plant Systematics and Evolution 187,(1993): 51-88

Florian P. Schiestl et al., "Variation of Floral Scent Emission and Post-Pollination Changes in Individual Flowers," Journal of Chemical Ecology 23, no. 12 (1997), is one of many articles on this subject.

M. Gierfa, "The Repellent Scent Mark of the Honeybee Apis mellifera ligustica and Its Role as Communication Cue During Foraging," Insect Society 40 (1993), discusses the memos left by some bees.

Ackerman, Natural History of the Senses, describes Joy as the most expensive perfume in the world.

The Shape of Things to Come

I am grateful to my neighbors for growing passionflowers (Passifiora incarnata).

Peter Bernhardt, The Rose's Kiss: A Natural History of Flowers (Washington, D.C.: Island Press, 1999), gives thorough and wonderful passages about the shapes of flowers and the vocabulary that botanists use when talking about flowers. The quote I use from this book is in his chapter "The Pig in the Pizza."

Moore et al., Botany, also gives a good description of the parts of a flower.

The paragraphs on evolution were particularly difficult to write, given the complexity of the subject. I consulted a number of books, including Niles Eldredge, Life in the Balance: Humanity and the Biodiversity Crisis (Princeton, N.J.: Princeton University Press, 1998); Niles Eldredge, Fossils: The Evolution and Extinction of Species (New York: H. N. Abrams, 1991); and E. O. Wilson, The Diversity of Life (New York: W. W. Norton and Company, 1992). Another very readable book on the subject is David Quamman, The Song of the Dodo (New York: Scribner, 1996).

The latest information on monkeyflowers comes from Susan Milius, "Monkeyflowers Hint at Evolutionary Leaps," Science News 156 (October 16, 1999). The information on how the bills of hummingbirds evolve to fit the shape of corollas is in Ethan Temeles and Paul Ewald, "Fitting the Bill?" Natural History 108 (May 1999).

R. Dawkins and J. R. Krebbs, "Arms Races Between and Within Species," Proceedings R. Society of London B 205, 489-511 (1979), was helpful, as were Candace Galen, "Why Do Flowers Vary?" Bioscience 49 (August 1999), and Graham Pyke, "Optimal Foraging in Bumblebees and Co-evolution with Their Plants," Oecologia (Berl.) 36, 281-293, (1978).

The quote by Charles Darwin is from his Origin of Species (1859) and was quoted by Friedrich Barth in Insects and Flowers.

Sex, Sex, Sex

Lack, Natural History of Pollination, gives a good description and explanation of flower sex, as does Barth, Insects and Flowers; Bernhardt, The Rose's Kiss; and Moore et al., Botany. Karl Niklas, "What's So Special about Flowers?" Natural History 108 (May 1999), is helpful. I also recommend Karl Niklas, The Evolutionary Biology of Plants (Chicago: University of Chicago Press, 1997). In addition, I used Bob Gibbons, The Secret Life of Plants (Blandford, London, England, 1990).

Nickolas Waser reminded me that any discussion of natural selection has to avoid certain land mines and that any discussion on the function and usefulness of sex is still theoretical. On his urging, I consulted such articles as F. F. Green and D. L. G. Noakes, "Is a Little Bit of Sex as Good as a Lot?" Journal of Theoretical Biology 174, 87-96 (1995); Harris Bernstein, Gregory S. Byers, and Richard Michod, "Evolution of Sexual Reproduction: Importance of DNA Repair, Complementation, and Variation," American Naturalist 117 (April 1981); D. G. Lloyd, "Benefits and Handicaps of Sexual Reproduction," Evolutionary Biology 13, 69-111 (1980); L. Nunney, "The Maintenance of Sex by Group Selection," Evolution 43 (1989) 245-257; and Nickolas Waser and Mary Price, "Population Structure, Frequency-Dependent Selection, and the Maintenance of Sexual Reproduction," Evolution 36 (1982).

I also looked at more popular articles, such as Bryant Furlow, "Flower Power," New Scientist, January 9, 1999.

In the Heat of the Night

A primary source of information for this chapter was Roger Seymour, a professor in the Department of Environmental Biology at the University of Adelaide, Australia, whose works include "Plants That Warm Themselves," Scientific American, March 1997; and "Analysis of Heat Production in a Thermogenic Arum Lily, Philodendron selloum, by Three Calorimetric Methods," Thermochimica Acta 193 (1991), 91-97. I also read Roger Seymour, George Bartholomew, and Christopher Barnhart, "Respiration and Heat Production by the Inflorescence of Philodendron selloum Koch," Planta 157 (1988); Roger Seymour and Paul Schultz-Motel, "Thermoregulating Lotus Flowers," Nature, September 26, 1996; Roger Seymour and Paul Schultz-Motel, "Temperature Regulation Is Not Associated with Odor Production in the Dragon Lily (Dracunculus vulgaris)" (poster presented at Sixteenth International Botanical Congress, St. Louis, Mo., August 1999); and Roger Seymour and Amy J. Blaylock, "Switching of the Thermostat: Thermoregulation by Eastern Skunk Cabbage (Symplocarpus foetidus)" (poster presented at Sixteenth International Botanical Congress, St. Louis, Mo., August 1999).

Several other articles were helpful for this chapter as well. They included Bastiaan Meeuse and Ilya Raskin, "Sexual Reproduction in the Arum Lily Family, with Emphasis on Thermogenicity," Sexual Plant Reproduction (1988) I: 3-15; Gerhard Gottsberger and Ilse Silberbauer-Gottsberger, "Olfactory and Visual Attraction of Eriscelis emarginata (Cyclocephalini, Dynastinae) to the Inflorescences of Philodendron selloum (Araceae)," Biotropica 23, no. I (1993); Hanna Skubatz, William Tang, and Bastiaan Meeuse, "Oscillatory Heat Production in the Male Cones of Cycads," Journal of Experimental Botany 44 (February 1993); and Bastiaan Meeuse, "The Voodoo Lily," Scientific American, July 1966.

Dirty Tricks

Judith Bronstein, a professor of evolutionary biology, was a good source for this chapter. Her publications included Judith Bronstein, "Our Current Understanding of Mutualism," Quarterly Review of Biology 69 (March 1994); Judith Bronstein, John F. Addicott, and Finn Kjellberg, "Evolution of Mutualistic Life-Cycles: Yucca Moths and Fig Wasps," in Insect Life Cycles: Genetics, Evolution, and Coordination, edited by Francis Gilbert (New York: Springer-Verlag, 1990); and Judith Bronstein and Yaron Ziv, "Costs of Two Non-Mutualistic Species in a Yucca/Yucca Moth Mutualism," Oecologia (1997) 112: 379-385.

Other sources were Olle Pellmyr and Chad Hurth, "Evolutionary Stability of Mutualism Between Yuccas and Yucca Moth," Nature, November 17, 1994; M. C. Ansteet, Judith Bronstein, and M. Hossart-McKay, "Resource Allocation: A Conflict in the Fig/Fig Wasp Mutualism," Journal of Evolutionary Biology 9, 417-428 (1996); Judith Bronstein, Didier Vernet, and Martine Hossart-McKey, "Do Wasp Figs Interfere with Each Other During Oviposition?" Entomologia Experimentalis et Applicata 87: 321-324 (1998); Susan Milius, "How Moths Tell if a Yucca's a Virgin," Science News Vol. 156 July 3, 1999); Jerry Powell, "Interrelationships of Yuccas and Yucca Moth," Trends in Evolution and Ecology 7 (January 1992); and A. J. Tyre and J. F. Addicott, "Facultative Non-Mutualistic Behavior by an 'Obligate' Mutualist: 'Cheating' by Yucca Moths," Oecologia (1993) 94:173-175.

Stephen Buchman and Gary Paul Nabhan, The Forgotten Pollinators, wonderfully describes the partnership of yuccas and yucca moths, as do other books on flowers.

The quote from Darwin is taken from his Origin of Species. The quote about how cheating pollinators are more rare than cheating plants comes from Jorge Soberon Mainero and Carlos Martinez del Rio, "Cheating and Taking Advantage in Mutualistic Associations," in The Biology of Mutualism, edited by Douglas Boucher (New York: Oxford University Press, 1985). These authors also discuss the idea of the aprochevado.

Alison Brody directed me to various articles on nectar theft and robbery, including Alison Brody and Rebecca Irwin, "Nectar-Robbing Bumblebees Reduce the Fitness of Ipomopsis aggregata (Polemonicea)," Ecology, in press; Alison Brody and Rebecca Irwin, ''Nectar Robbing in Ipomopsis aggregata: Effects on Pollinator Behavior and Plant Fitness," Oecologia (1998) 116: 519-527; and Alison Brody, "Effects of Pollinators, Herbivores, and Seed Predators on Flowering Phrenology," Ecology 78 (6) 1997 pp. 1624-1631 no. 6; as well as others.

Meeuse and Morris, The Sex Life of Plants, discusses forms of flower traps, deception, and mimicry. The book also describes various "murderous" arum plants and tells the story of the water lily Nymphaea capensis, which drowns the hapless hoverfly. This material is also covered in other books. Ethan Temeles and Paul Ewald, "Fitting the Bill," Natural History 108 (May 1999), has a good sidebar on the cruelty of flowers.

The material on daisies and armyworms comes from Dennis Bueckert, "Plant Warfare," Canadian Geographic, July 1994.

The role of ants in pollination is discussed in Proctor, Yeo, and Lack, The Natural History of Pollination.

The association of politics and the scientific idea of mutualism is brought up in Douglas Boucher, "The Idea of Mutualism, Past and Future," in The Biology of Mutualism, edited by Douglas Boucher (New York Oxford University Press, 1985).

The quote by the unnamed biologist is from Nickolas Waser. His ideas can be found in many articles, some already noted. Others include Nickolas Waser and Mary Price, "What Plant Ecologists Can Learn from Zoology," Perspectives in Plant Ecology, Evolution, and Systematics Vol. 1/2 pp. 137-150, 1998; Nickolas Waser et al., "Generalization in Pollination Systems and Why It Matters," Ecology 77 (June 1996); and Nickolas Waser, "Pollen Shortcomings," Natural History 7, no. 93 (1984).

Time

The basic material on physics and the examples of the clocks at the bottom of the tower and the twins at different places on the earth comes from Stephen W. Hawking, A Brief History of Time: From the Big Bang to Black Holes (New York: Bantam Books, 1988).

Material on the cereus cactus can be found in many books. I used Gary Paul Nabhan, Desert Legends: Re-storying the Sonoran Borderlands, with photography by Mark Klett (New York Henry Holt and Company, 1994). Nabhan is quoted regarding the "ugly duckling" and cuddly appeal of the cereus cactus. He also mentions that he briefly thought the flowers were left-behind flashlights. I also consulted and used material from Susan Tweit, Seasons in the Desert: A Naturalist's Notebook (San Francisco: Chronicle Books, 1998).

More information on the history of Silver City and its parties can be found at the Silver City Museum, run by its esteemed director, Susan Berry.

Bernhardt, The Roses Kiss, has many good passages on the how and why of a flower's life span.

Tweit, Seasons in the Desert, and Nabhan, Desert Legends, are both good sources for information on the century plant.

Travelin' Man

Bernhardt, The Rose's Kiss, has a number of good sections on pollen, as does Barth, Insects and Flowers; and Proctor, Yeo, and Lack, The Natural History of Pollination.

The quote about the "thin and particulate sheet" comes from Douglas Boucher, ed., The Biology of Mutualism (New York: Oxford University Press, 1985).

I also consulted S. Blackmore and I. K. Ferguson, eds., Pollen and Spores: Form and Function (Orlando, Fl.: Academic Press, 1985), particularly the chapter by W. Punt, "Functional Factors Influencing Pollen Form"; and Irene Till-Bottraud et al., "Selection of Pollen Morphology: A Game Theory Model," American Naturalist 144 (September 1994).

The information on Neanderthal burials was mainly taken from Arlette Leroi-Courhan, "The Flowers Found in Shanidar IV, a Neanderthal Burial in Iraq," Science 190 (November 1975).

For the material on the murders in Germany, I consulted R. Szibor et al., "Pollen Analysis Reveals Murder Season," Nature 395 (October 1998). A general article is Meredith Lane et al., "Forensic Botany," BioScience 40 (January 1990).

The information on the Shroud of Turin was discussed in Avinoam Danin, "Traces of Ancient Flower Pollen on the Shroud of Turin: New Botanical Evidence to Date and Place the Burial Cloth of Jesus of Nazareth" (media presentation at the Sixteenth International Botanical Congress, St. Louis, Mo., August 1999); and in many newspaper articles, including Jack Katzenell, "Plant Cues Place Shroud in Holy Land," Albuquerque Journal, June 16, 1999.

For the discussion on buzz pollination, I read Stephen Buchman, "Buzz Pollination in Angiosperms," in Handbook of Experimental Pollination Biology edited by Eugene Jones and John Little (Princeton, N.J.: Princeton University Press, 1983); and Susan Milius, "Color Code Tells Bumblebees Where to Buzz," Science News 155 (April 3, 1999), as well as other articles.

A number of previously mentioned books and articles talk about the life of bees. I enjoyed Susan Brind Morrow, "The Hum of Bees," Harper's Magazine, September 1998.

The quote from the Navajo chant is from Margaret Link, ed., The Pollen Path: A Collection of Navajo Myths (Stanford, Calif.: Stanford University Press, 1956).

Living Together

The article "Sensitive Flower" by Andy Coghlan in New Scientist, September 26, 1998 summarizes recent work on how flowers "see," "smell," "touch," and "taste." Numerous other articles, such as Stephen Day, "The Sweet Smell of Death," New Scientist, September 7, 1996; Garry C. Whitelan and Paul E. Devlan, "Light Signaling in Arabidopsis," Plant Physiology Biochemistry 1998 36 (1-2) 125-133; and Paul Simons, "The Secret Feelings of Plants," New Scientist, October 17, 1992, discuss these subjects.

For speculation on how plants react to thunderstorms, see Stephen Young, "Growing in Electric Fields," New Scientist, August 32, 1997.

Autar K. Matoo and Jeffrey C. Suttle, eds., The Plant Hormone Ethylene (Boca Raton, Fl.: CRC Press, 1988), gives important information about this hormone. Bernhardt, The Roses Kiss, also discusses the signals that start the development of a flower.

Specifically for plant intercommunication, I looked at Jan Bruin, Maurice W. Sabelis, and Marcel Dicke, "Do Plants Tap SOS Signals from Their Infested Neighbors?" Trends in Evolution and Ecology 10 (April 1995); Irene Sconle and Joy Bergelson, "Interplant Communication Revisited," Ecology 76 (December 1995); and Marcel Dicke et al., "Jasmonic Acid and Herbivory Differentially Induce Carnivore-Attracting Plant Volatiles in Lima Bean Plants," Journal of Chemical Ecology 25, no. 8 (1999), as well as other articles.

Proctor, Yeo, and Lack, The Natural History of Pollination, provides a good base for understanding communities of flowers. More about allelopathy can be found in numerous textbooks like Moore et al., Botany, and in articles like Gail Dutton, "Yo Buddy -- Outa My Space," American Horticulturist, Vol. 72 March 1993; and Chang-hung Chou, "Roles of Allelopathy in Plant Biodiversity and Sustainable Agriculture," Critical Reviews in Plant Sciences 18, no. 5 (1999).

Dutton, ''Yo Buddy 00 Outa My Space," discusses how plants use their roots, as does A. Tayler, J. Martin, and W E. Seel, "Physiology of the Parasitic Association Between Maize and Witchweed (Striga hermonthica)," Journal of Experimental Botany 47, no. 301 (1996); and Charles Mann "Saving Sorghum by Foiling the Wicked Witchweed," Science, August 22, 1997.

James Tumlinson, W. Joe Lewis, and Louside E. M. Vet, "How Parasitic Wasps Find Their Hosts," Scientific American, March 1993, and other articles discuss the relationship of some plants to wasps.

For the mimicry of Puccinia rust fungi, I consulted Robert Raguso and Barbara Roy, "Floral Scent Production by Puccinia Rust Fungi That Mimic Flowers," Molecular Ecology (1998) 7: 1127-1136; and Barbara Roy, "Floral Mimicry by a Plant Pathogen," Nature 362, March 1993.

Batesian mimicry and Mullerian mimicry are described in many textbooks. I also used articles such as Barbara Roy and Alex Widmer, "Floral Mimicry: A Fascinating yet Poorly Understood Phenomenon," Trends in Plant Sciences 4 (August 1999); James Marden, "Newton's Second Law of Butterflies," Natural History Vol. 1 (January 1992); Lori Oliwens, "Royal Flush," Discover, January 1992; and James Brown and Astrid Kodric-Brown, "Convergence, Competition, and Mimicry in a Temperate Community of Hummingbird-Pollinated Flowers," Ecology 60, no. 5 (1979).

Quamman, The Song of the Dodo, has a good discussion on some of the controversies concerning Darwin and Wallace, as well as accounts of Henry Bates's and Wallace's collecting adventures. I also read Henry Bates, The Naturalist on the River Amazon: A Record of Adventures, Habits of Animals, and Sketches of Brazilian and Indian Life (Dover Publications, 1975, and Mea Allan, Darwin and His Flowers: The Key to Natural Selection (Taplinger Press, 1977).

The Tower of Babel and the Tree of Life

A very helpful book for this chapter was William Stearn, Botanical Latin (Hafner Publishers, 1966), as well as portions of Moore et al., Botany; and Tod F. Stuessy, Plant Taxonomy: The Systematic Evaluation of Comparative Data (New York: Columbia University Press, 1982).

Information about Carl Linnaeus came from many sources, including Tore Frangsmyr, ed., Linnaeus: The Man and His Work (Science History Publications, 1994) and Bil Gilbert, "The Obscure Fame of Carl Linnaeus," Audubon Vol. 86 (September 1984).

For some of the newest ideas on taxonomy, I attended a number of sessions on the subject at the Sixteenth International Botanical Congress in St. Louis, Missouri, in August 1999. I also read Brent Mishler, "Getting Rid of Species," in Species: New Interdisciplinary Essays (Cambridge, Mass.: MIT Press, 1999); Rick Weiss, "Plant Kingdoms' New Family Tree," Washington Post, August 5, 1999; Susan Milius, "Should We Junk Linnaeus?" Science News 156 (October 23, 1999); William Stevens, "Rearranging the Branches on a New Tree of Life," New York Times, September 23, 1999; and Glennda Chui, "Tree of Life Proposal Divides Scientists," Mercury News, September 23, 1999, as quoted on the Deep Green Web page, http://ucjeps.herb.berkeley.edu, with additional keywords "bryolab" and "greenplantpage"; "Team of Two Hundred Scientists Presents New Research That Reveals Full Tree of Life for Plants" (press release prepared by International Botanical Congress, August 4, 1999); and Jeff Doyle, "DNA, Phylogeny, and the Flowering of Plant Systematics," BioScience 43 June 1993).

Flowers and Dinosaurs

For some of the newest information on the evolution of green plants, including the idea that they came out of fresh and not salt water, I attended key sessions at the Sixteenth International Botanical Congress, St. Louis, Mo., August 1999. See also, for example, Kathryn Brown, "Deep Green Rewrites Evolutionary History of Plants," Science Magazine 285 (September 1999), listed on the Deep Green Web page.

Loren Eiseley's essay "How Flowers Changed the World" was republished as a book by the same title, with photographs (San Francisco: Sierra Club Books, 1996).

Bernhardt, The Roses Kiss, recounts the story of the evolution of flowers, as does the Moore et al., Botany.

Else Marie Friis, William G. Chaloner, and Peter R. Crane, eds., The Origins of Angiosperms and Their Biological Consequences (New York: Cambridge University Press, 1987), was an important source, particularly the following chapters in the book: Else Marie Friis, William G. Chaloner, and Peter R. Crane, "Introduction to Angiosperms"; Peter R. Crane, "Vegetational Consequences"; Else Marie Friis and William Crepet, "Time and Appearance of Floral Features"; William Crepet and Else Marie Friis, "The Evolution of Insect Pollination"; and M. J. Cow et al., "Dinosaurs and Land Plants."

See also Conrad C. Labandeira, "How Old Is the Flower and the Fly?" Science 280 (April 3, 1998); Ge Sun et al., "In Search of the First Flower," Science 282 (November 27, 1998); Peter R. Crane, Else Marie Friis, and Raj Pedersen, "The Origin and Early Diversification of Angiosperms," Nature, March 2, 1995; Ollie Pellmyr, "Evolution of Insect Pollination and Angiosperm Diversification," Trends in Evolution and Ecology 7 (February 1992); and David Winship Taylor and Leo Hickey, "An Aptian Plant with Attached Leaves and Flowers," Science 247 (February 9, 1990).

Information on the fossil flowers in New Jersey came from William Crepet, "Early Bloomers," Natural History 108 (May 1999); and Carol Yoon, "In Tiny Fossils, Botanists See a Flowery World," New York Times, December 21, 1999.

Information about the floral landscape of dinosaurs came from sessions at the Sixteenth International Botanical Congress, St. Louis, Mo., August 1999, such as Peter R. Crane, "Plants and Flowers from the Age of Dinosaurs: New Discoveries and Ancient Flowers" (talk given at the congress).

Many books and articles discuss the extinction of dinosaurs and the theories and controversies surrounding that event. Among others are Carl Zimmer, "When North America Burned," Discover, February 1997. I should also include Frank DeCourten, The Dinosaurs of Utah (Salt Lake City: University of Utah Press, 1998); and Tim Haines, Walking with Dinosaurs (BBC Worldwide, 1999).

Kirk Johnson, "Leaf Fossil Evidence for Extensive Floral Extinction at the Cretaceous-Tertiary Boundary, North Dakota, USA," Cretaceous Research (1992) 13, 91-117, provided specific information about the KT Boundary and the floral extinction of that time.

Regarding mass extinction and empty niches, the theory of punctuated equilibrium is well described in Eldredge, Fossils.

The news about Amborella was presented at the Sixteenth International Botanical Congress, St. Louis, Mo., August 1999, and was reported in articles like Susan Milius, "Botanists Uproot Their Old Tree of Life," Science News 156 (August 7, 1999).

The Seventh Extinction

Local and regional newspapers provided statistics on the effects of the heat wave in the summer of 1999. See, for example, Bob Herbert, "When Summer Turns Deadly," New York Times, August 8, 1999.

Several press releases from the Sixteenth International Botanical Congress, St. Louis, Mo., August 1999, gave information on extinction, for example, "World's Biodiversity Becoming Extinct at Levels Rivaling Earth's Past Mass Extinctions"; "Nearly Half of Earth's Land Has Been Transformed by Humans: Fifty Dead Zones Found in Oceans"; and "World Conservation Union (IUCN) Mobilizes International Team of Experts to Save Plant Species." Many others also gave presentations concerning extinction and human influence on that process (e.g., Peter Raven [president of the congress], "Mass Extinction of the Earth's Plant Species: Can We Prevent It?" Jane Lubchenco, "The Human Footprint on Earth: New Research"; Mike Wingfield, "Alien Invasions: Combating Aggressive Takeovers"; David Brackett, "Survival of Plant Species: A Plan of Action for the New Millennium"; and Gregory Anderson, "Threatened Islands: Storehouses of Biological Treasures"). Many other presentations at the congress dealt also with these subjects.

See also David Quamman, "Planet of Weeds," Harper's Magazine, October 1988. A good book on extinctions of island life, of course, is Quamman, Song of the Dodo.

More information on the extinction of plant species can be found in Sally Deneen, "Uprooted," E: The Environmental Magazine 10 (July 1999); Carol Kearns, David Inouye, and Nickolas Waser, "Endangered Mutualisms: The Conservation of Plant Pollinator Interactions," Annual Review of Ecology Systematics 29, 1998, 83-112; Fred Powledge, "Biodiversity at the Crossroads," Bioscience 48 (May 1998); and Carol Kearns and David Inouye, "Pollinators, Flowering Plants and Conservation Biology," BioScience 47 (May 1997), as well as other articles and sources.

What We Don't Know

In addition to interviews and discussions with Rob Raguso, I also read Robert Raguso and Mark Willis, "The Importance of Olfactory and Visual Cues in Nectar Foraging by Nocturnal Hawkmoths," Proceedings of Third International Congress of Butterfly Ecology and Evolution (Chicago: University of Chicago Press, 2000); Robert Raguso and Eran Pichersky, "A Day in the Life of a Linalool Molecule: Chemical Communications in a Plant Pollinator System," Plant Species Biology (in press); Natalia Dudareva et al., "Floral Scent Production in Clarkia breweri," Plant Physiology 116 (1998); and Robert Raguso and Barbara Roy, "Floral Scent Production by Puccinia Rust Fungi That Mimic Flowers," Molecular Ecology 7 (1998).

Alchemy of a Blue Rose

Bernhardt's The Rose's Kiss has good passages on double roses and the evolution of the roses. For more information on crossbreeding, see Steve Kemper, "Ron Parker Puts the Petals on Their Mettle," Smithsonian 25 (August 1994). Ron Parker is the source for the quote on flower colors and nonwhite siding.

"Programs Are Launched to Analyze Impact of Bt Corn on Monarch Butterflies," Chemical Market Report 256 (November 1999); and "Of Corn and Butterflies," Time 153 (May 1999), both concern the controversy about genetically engineered corn and its potential harm to butterflies.

For more about the blue rose, see the following articles in New Scientist, October 31, 1998: David Concar, "Brave New Rose"; Phil Cohen, "Running Wild"; Martin Brookes and Andy Coghlan, "Live and Let Live"; and Debbie Mack, "Food for All." I also consulted Andy Coghlan, "Blooming Unnatural," New Scientist, May 22, 1999; Rozanne Nelson, "Not Making Scents," Scientific American, September 1999; Ruth Pruyne, "Green Genes," Penn State Agriculture Magazine (winter 1997); and Ruth Pruyne, "Shedding Light," Breakthroughs (magazine for alumni of the College of Natural Resources at University of California Berkeley) (summer 2000).

Jeremy Rifkin's quote is from his Biotech Century (Penguin Putnam, 1998).

Phytoremediation

For the use of sunflowers to clean up radiation, I read a number of articles, including Andy Coghlan, "Flower Power," New Scientist, December 6, 1997. Another good article on phytoremediation was Amy Adams, "Let a Thousand Flowers Bloom," New Scientist, December 1997. The Sixteenth International Botanical Congress, St. Louis, Mo., had numerous lectures and presentations on phytoremediation, including the media presentation by Ilya Ruskin, "Plants That Are Decontaminating the Environment."

Information on the bitter nut kola came from Maurice Iwu, "Ethnobotany: A New Plant Discovery to Cure Disease" (media presentation at Sixteenth International Botanical Congress, St. Louis, Mo., August 1999); and from "Edible Plant Stops Ebola Virus in Lab Tests" (press release of Sixteenth International Botanical Congress, St. Louis, Mo., August 1999). Several newspapers and magazines had follow-up articles on this discovery.

Information on the rosy periwinkle comes from Systematics Agenda, 2000, Charting the Biosphere (distributed at Sixteenth International Botanical Congress, St. Louis, Mo., August 1999).

For information on the medicinal uses of plants, I used Michael Moore, Medicinal Plants of the Mountain West (Museum of New Mexico Press, 1979), as well as other sources.

For material on flower essences, I consulted Clare Harvey and Amanda Cochrane, The Encyclopedia of Flower Remedies: The Healing Power of Flower Essences Around the World (Thorsons, 1996); and Anne McIntyre, Flower Power (New York: Henry Holt, 1996).

Material on sunflowers can be found in many books and articles, including Rita Pelczar, ''The Prodigal Sunflower," American Horticulturist, August 1993.

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