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For Further Reading

Darwin, C. R. 1859. On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life.

Reference for: Chapter 12, The Foundations of Evolutionary Theory

Haldane, J. B. S. 1932. The Causes of Evolution. Longman: UK.

Reference for: Chapter 12, The Foundations of Evolutionary Theory


*Long, John A. 1995. The Rise of Fishes: 500 Million Years of Evolution. John Hopkins University Press: MD

This lavish overview of the evolution of fishes is not the most detailed but its illustrations and photographs give a rich sense of the evidence on which our understanding of fish evolution is based. It makes a highly readable reference for students and a terrific desk reference for instructors called upon to teach aspects of fish evolution.

Reference for: Chapter 12, Spotlight 12.1

*Raup, David. 1991. Extinction: Bad Genes or Bad Luck? W.W. Norton: NY

This “little” book summarizes the evidence for five major extinctions in the geologic records and their causes. It’s a highly readable and engaging account that will quickly bring the reader up to date on this fascinating topic.

Reference for: Chapter 12, The Foundations of Evolutionary Theory


*Stott, Rebecca. 2003. Darwin and the Barnacle: The Story of One Tiny Creature and History’s Most Spectacular Scientific Breakthrough. Norton: NY

This book brings to the forefront Darwin’s painstaking and highly important work on barnacles. It might be argued that Darwin formulated his ideas about evolution and natural selection from studying barnacles. Although this is a “storybook”, in the sense that it weaves a narrative about Darwin’s barnacle work, it does illuminate this important and little known work in an engaging and instructive manner.

Reference for: Chapter 12, The Foundations of Evolutionary Theory

*Carroll, Sean B. 2006. The Making of the Fittest: DNA and the Ultimate Forensic Record of Evolution. W. W. Norton: NY

The evolutionary record is contained in the DNA of organisms. It is a history that we can finally begin to read.



*Coyne, Jerry A., and H. Allen Orr. 2004. Speciation. Sinauer Associates: MA.

Coyne and Orr have written a textbook covering all aspects of speciation, emphasizing modern research on this topic.


*Ellis, Richard. 2001. Aquagenesis: The Origin and Evolution of Life in the Sea. Viking Penguin Books: NY

Ellis is a masterful storyteller and illustrator. There are better books on this subject but if you like Ellis way of weaving facts, this book should please you.


*Fortey, Richard. 1997. Life: A Natural History of the First Four Billion Years of Life on Earth. Vintage Books: NY

Fortey narrates the history of life on Earth, citing his own work and the research of other scientists to piece together the puzzles of how life evolved.


*Fortey, Richard. 2000. Trilobite! Eyewitness to Evolution. Alfred A. Knopf: NY

All you ever wanted to know about trilobites in an engaging, delightful prose.

*Gould, Stephen Jay. 1989. Wonderful Life: The Burgess Shale and the Nature of History. W. W. Norton: NY

Stephen Jay Gould delights some and irritates others but he always manages to inspire thoughtful reflection on a topic. In this book, he discusses in great detail the Burgess Shale and how it paints a picture of the “progression” of evolution unlike what is commonly perceived. Gould sees evolution not only as “survival of the fittest” but also as “survival of the lucky.”

*Gould, Stephen Jay. 2001. The Book of Life: An Illustrated History of the Evolution of Life on Earth. W.W. Norton: IA

*Gould, Stephen Jay. 2002. The Structure of Evolutionary Theory. Belknap Press of Harvard University Press: MA

This immense volume details Gould’s provocative and often controversial views on the evolution of life on Earth. To his credit, Gould is typically entertaining, and this book reads like a good novel. Unfortunately, you have to read a lot of it if you are generally unfamiliar with his ideas or the nuances of evolution. Nonetheless, it’s an essential reference for a biologist’s library.

*Hull, David L. 2001. Science and Selection: Essays on Biological Evolution and the Philosophy of Science. Cambridge University Press: UK

Hull’s essays educate and entertain and get the reader to thinking more deeply about science and its effects on humanity. His essays on evolution are a big help to those who need a refresher or those who require greater ammunition in the verbal wars with antievolutionists.

*Johnson, Kirk R., and Richard K. Stucky. 1995. Prehistoric Journey: A History of Life on Earth. Roberts Rinehart Publishers: CO.

Based on dioramas at the Denver Museum of Natural History, this delightfully illustrated book traces the history of life from microbes to mammals, with an emphasis on dinosaurs. Its brevity notwithstanding, this book does a great job of providing the fossil evidence on which the scientific interpretation of the history of life is based.

*Kirschner, Marc W. and John C. Gerhart. 2005. The Plausibility of Life: Resolving Darwin’s Dilemma. Yale University Press: CT

Kirscner and Gerhart tackle the origins of new species and evolutionary complexity.

*Knoll, Andrew. 2003. Life on a Young Planet: The First Three Billion Years of Evolution on Earth. Princeton University Press: NJ

This is an outstanding book on the evolution of Earth and its biota. Knoll is one of the pioneers in the field of geobiology and his up-to-date scientific account of the field makes this an excellent reference and an entertaining read. Knoll exposes the controversies and examines the evidence that surrounding them. Most narratives don’t make good reference books but Knoll’s is an exception. If you are trying to choose between “histories of life on Earth”, pick this one.

*Larson, Edward J. 2004. Evolution: The Remarkable History of a Scientific Theory. Modern Library: NY

This book sketches the development of evolutionary theory. It’s primarily written for general audiences and so loses some of the detail required for students and instructors.

*Margulis, Lynn, and Dorion Sagan. 1986. Microcosmos: Four Billion Years of Microbial Evolution. Simon and Schuster: NY.

A provocative hypothesis about the interdependency of higher organisms and bacteria.

*Margulis, Lynn. 1998. The Symbiotic Planet: A New Look at Evolution. Weidenfeld & Nicolson: UK

Margulis is not one to shy away from controversy. Her endosymbiotic hypothesis was met with great skepticism originally but is now widely accepted. In this book, she applies her principles of symbiosis to the full range of life and its communities, including Earth.

*Margulis, Lynn, and Michael F. Dolan. 2002. Early Life: Evolution of the PreCambrian Earth, 2nd Edition. Jones and Bartlett: MA

*Mayr, Ernst. 1982. The Growth of Biological Thought: Diversity, Evolution, and Inheritance. Belknap Press of Harvard University Press: MA

Professor Sean thinks this is one of the most important books ever written. It defends the place of biology in science and retells the history of evolutionary thinking from pre- to neo-Darwinism. At more than 900 pages, it’s an intimidating volume, but Mayr’s prose and his way of explaining concepts makes this book a delight to read. You will only want to read several pages of it at a time as Mayr provokes deep reverie with every page. But you will have a more comprehensive and deeper understanding of evolution upon reading this book than is possible with just about any other book.

*Mayr, Ernst. 2001. What Evolution Is. Basic Books: NY

Any book by Ernst Mayr is worth reading, according to Professor Sean. This book provides a solid foundation for different aspects of evolution and evolutionary processes.

*Weiner, Jonathan. 1994. The Beak of the Finch. Vintage Books: NY

This Pulitzer Prize-winning book has become a textbook for learning about evolution.

*Zimmer, Carl. 1998. At the Water’s Edge: Fish With Fingers, Whales With Legs, and How Life Came Ashore but Then Went Back to Sea. Simon and Schuster: NY

An excellent narrative on macroevolution.

*Zimmer, Carl. 2001. Evolution: The Triumph of an Idea. HarperCollins: NY

This is the companion book to the Evolution video series by PBS.

*Moorehead, Alan. 1969. Darwin and the Beagle. Harper & Row: NY

This “old” book is notable for its abundant photos, illustrations and drawings, many of which are full page and stunning, and for its highly readable and intimate account of Charles Darwin’s voyage aboard HMS Beagle. It’s not as dense with information as other books on Darwin but it captures the spirit of his curiosity and scientific reasoning.

Reference for: Chapter 12, The Foundations of Evolutionary Theory

The Endless Voyage: Building Blocks, Water World and Survivors (written by W. S. Chamberlin) (Episodes 18, 19 and 21). 2002 (VHS and DVD). Intelecom.

Professor Sean appeared in several of the episodes of this series and helped develop learning activities to support it. While some episodes are better than others, The Endless Voyage provides one of the most complete and up-to-date series on oceanography available

: : Encyclopedia of the Sea : :
Chapter Two Image

The Benthic Annelids: Polychaetes, Beard Worms, Girdle Worms, and Sea Leeches by Sean Chamberlin

Bristle worms (polychaetes), girdle worms (oligochaetes) and beard worms (pogonophora) belong to the same phylum as earthworms but most hardly resemble earthworms. All are members of the taxon Annelida, the segmented worms, though the pogonophorans were previously thought to represent their own phylum. Segmentation in these animals, including arthropods, is thought to confer several advantages but agreement concerning the evolutionary significance of this body plan is lacking. Be that as it may, the diversity of these animals and their importance to biogeochemical cycles is evidence of their success, regardless of the reasons.


Polychaetes are among the most beautiful and diverse worms in the ocean; some are even named after Greek goddesses. Familiar ones, like the suspension-feeding Christmas tree worm or feather duster, live in tubes. Others, like the predatory sand worm, Nereis, are free-roaming. Polychaetes may be found inhabiting a wide range of habitats from the shallow intertidal to deep-sea muds.

The body plan of a motile polychaeate resembles an earthworm except for the numerous bristle-covered flaps, called parapdodia, that extend from each side of its segments. A prominent proboscis, an eversible mouth, acts like a vacuum cleaner to feed on sediments and may, in some species, contain powerful chitinous jaws for subduing prey. In the polychaete, Glycera convolute, the proboscis wields four fangs that inject a neurotoxin called glycerotoxin. This compound, like many naturally occurring neurotoxins, has been especially useful in studies of synaptic transmission (e.g., Meunier et al., 2002). Tube-dwelling polychaetes differ largely by modifications to their heads, namely, the presence of colorful or elongate tentacles used in suspension or surface deposit feeding. Eudistylia, a common species of feather duster worm along the Pacific coast, bears tentacles that resemble something you would see at a show in Vegas. The Christmas tree worm, Spirobranchus, exhibits a wide range of exotic colors in a pair of tentacles shaped like a well-trimmed blue spruce.

Feeding in polychaetes presents interesting challenges for studies of the flow of energy and materials in benthic food webs. Fauchald and Jumars (1979) categorized several feeding modes in polychaetes as a means of distinguishing their potential ecological roles. These include the raptorial feeders, the motile polychaetes who actively seek food as carnivores, herbivore, detritivores, or omnivores; the filter feeders, usually tube-dwelling polychaetes who remove and/or select suspended particles; selective deposit feeders, tube-dwelling or burrowing polychaetes who collect and sort food particles at the surface using tentacles or other specialized body parts; and non-selective deposit feeders, the “earthworm” types, many motile but some tube-dwelling, who ingest sediments non-discriminately at the surface or within the sediments. Because the feeding behavior of many soft-bottom organisms is so difficult to observe, discriminating between these types of feeders and their role in digenesis remains difficult. Surface deposit feeders, like the cirratulid poychaetes, have been shown in laboratory and field studies with glass beads ingest sediment particles at the surface and egest them at depth, a process known as downward sediment transport (e.g., Shull and Yasuda, 2001). Some researchers have suggested that such processes may be important in the burial of carbon (e.g. Emerson et al., 1985).

Given their diverse lifestyles and forms of feeding, it should be no surprise that polychaetes often dominate soft-bottom communities. In estuarine, coastal and deep-sea sediments, they may comprise between 24-73% of all the species present (Hutchings, 1998). Nonetheless, the factors that influence species diversity in sediments are hotly debated, particularly in the deep sea where species diversity appears higher than coastal sediments (e.g., Levin et al, 2001). Also uncertain is the role that polychaetes (and other dominant benthic organisms) play in the global carbon cycle and the biogeochemical cycles of nitrogen, sulfur, iron and other elements. Research using benthic landers, which enclose a portion of the sea bottom for measurements of exchanges of gases and chemical compounds, indicate that bio-irrigation is an important component in the flux of biologically important nutrients from sediments on the Monterey Bay shelf (e.g. Berelson, 2003).

Polychaetes also play an important role on hydrothermal vents where they are renowned for their thermal tolerance. Lee (2003) reports that one species, Alvinella pompejana, can tolerate temperatures from 50 - 56° C (122-132° F), making it the “hottest” animal on Earth (sorry, Britney). Alvinella also boasts one of the most highly developed gills among polychaeates and a high-temperature hemoglobin with a very high oxygen binding constant that allow it to survive in poorly oxygenated waters (such as those leaving a vent). The dorsal surface of this polychaete apparently serves as a microhabitat for many different species of bacteria, some visible to the eye. This epibiotic association (epi = living on; biotic = life) has been called “the most highly evolved…among all invertebrates” (Van Dover, 2000; following Desbruyères et al. 1998). Numerous other polychaetes make their home on hydrothermal vents and likely serve as a link between chemosynthetic primary production and higher trophic levels.

Reproduction in many polychaetes involves the bizarre transformation of a benthic, non-reproductive atoke to a pelagic, gonad-bearing epitoke with enlarged eyes and parapodia modified for swimming. Like the mass spawning of corals, polychaetes spawn synchronously under specific conditions and at particular times of the year. In a phenomenon known as swarming, thousands of epitokes rise from the sea floor, usually in response to light cues (most being negatively phototaxic). At the surface, they release their eggs and sperms, oftentimes conducting a kind of nuptial dance, like the swarming palolo worms who twirl and twist as they spawn. American Samoans celebrate the night of the palolo by gathering along the shore with nets to catch and eat the annelid caviar (raw, fried with eggs or spread on toast). At least one species, the Bermuda fireworm (Odontosyllis enopla), employs bioluminescence in a mating ritual between “steady-glow” females and “blinker” males, ending, as one author puts it, “in a stunning luminescent final” of spewed eggs and sperm (Ruppert et al, 2004).

Beard worms (Giant tube worms, vent worms or lipstick worms)

Beard worms are named for the beard-like “tentacles” (palps) that line the anterior part of their body (prostomium). They gained their fame in 1977 when the deep-sea submersible Alvin discovered colonies of “giant tubeworms” on hydrothermal vents of the Galapagos Rift. These vent worms (more aptly called lipstick worms for the vivid red prostomium that extends and contracts from a tube like lipstick) were initially placed in the Phylum Pogonophora but since have been classified as polychaetes (Black et al., 1997). Nonetheless, their legendary status in the annals of oceanographic history give them special attention here.

When first discovered, Riftia pachyptila (as it is known) was a biological oddity for its extreme size (animals up to nine feet have been found, but typically they range from 2-4 feet); its lack of a mouth, digestive system or anus; and its specialized organ, the trophosome, packed with bacteria. The first question scientists asked: how does an animal get so large without any obvious means of nutrition? The answer was found in anatomical studies of the worm, revealing four distinct body regions that support their symbiotic relationship with chemoautotrophic bacteria (see Figure ##). The plume of the worm (the lipstick “beard”) extends into vent water containing hydrogen sulfide and other nutrients which can be absorbed and transferred via blood vessels to the internal compartments of the animal, namely the trophosome. Here reside trillions of bacteria (several trillion bacteria per gram body weight of trophosome have been observed) that produce ATP by oxidizing hydrogen sulfide (chemoautotrophy) and use that energy to supply sugars (presumably) to their worm host. Rates of chemoautotrophy in Riftia rank highest among chemosymbiotic species and are more than sufficient to meet the metabolic demands of the organism by at least twofold (Van Dover, 2000; following Childress et al. 1991).

Girdle Worms

Within the class Oligocheata (Phylum Annelida), are around 200 species of girdle worms that live interstitially or in tubes in beach sediments although some deep-sea species are known. Girdle worms, which include the familiar earthworm (and are sometimes called aquatic earthworms), display a girdle-like “gland”, called a clitellum, that serves various reproductive functions and give these animals their name. Otherwise, their body plan closely resemble burrowing polychaetes.

Among the most conspicuous girdle worms are the tube-dwelling ones (tubificids), notably Tubifex. These hardy animals can be found in the mud flats of estuaries, especially where human impacts are high, like San Francisco Bay and New York/New Jersey harbor. Able to withstand large fluctuations in salinity, temperature and water conditions, these animals are an important indicator species of “pollution” in shallow water, soft-bottom habitats. They also serve as an important source of food for bottom-feeding fishes. Unfortunately, the accumulation of toxic chemicals in their tissues leads to biomagnification of those toxins by fish who feed on them. Tubifex has also been shown to feed selectively on silts and clays (which are the most susceptible to metal and toxin binding) and egest them on the surface, which has important implications for ecotoxicological studies and possibly, for sediment transport in the sediments.

See Hydrobiologia, November 2001, Vol. 463, 1-3, for an entire series of articles on aquatic oligochaetes. Google-search keywords: tubificids, marine oligochaetes, girdle worms

Sea leeches

Made infamous by Humphrey Bogart in The African Queen, the leeches—a taxa within Annelida—often evoke sounds of disgust at their mention. Nonetheless, leeches continue to serve an important medical role for humans, being used to reattach severed appendages (like fingers and toes).

In the ocean, only a few marine leeches are known and these act as parasites on sharks and rays. Should you ever lose a finger to a shark and can retrieve it, the shark’s companion leech could come in handy.