Congressional Fellow

STATISTICAL ECOLOGY SECTION NEWSLETTER ~~E.C. Pielou Award

The E.C. Pielou Award is a competitive award made annually to a graduate student or recent Ph.D, and is based on overall quality of the student’s scientific contribution to statistical ecology. The 2003 recipient of the E.C. Pielou Award is Katia Koelle for her talk at the 2003 ESA Annual Meeting in Savannah, “Disentangling the roles of extrinsic and intrinsic factors in nonlinear disease dynamics.” Koelle is a third-year graduate student at the University of Michigan and is advised by Mercedes Pascual. Her dissertation focuses on analyzing the spatiotemporal dynamics of diseases, particularly diseases that may experience significant climatic forcing. Her current research uses cholera case data from Bangladesh to reconstruct the duration of temporary immunity conferred by the disease and to examine the role of the El Niño–Southern Oscillation as an environmental driver of the disease. She received her undergraduate degree in Biology from Stanford University in 1997.

Statistical Section Officers: Sam Scheiner, Randy Balice, Kathy Cottingham

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Meetings

Calendar

Species Exchanges between Eastern Asia and North America:
Threats to Environment and Economy

The Beijing International Symposium on biological invasions will be held 8–11 June 2004 in Beijing, China. The symposium will be co-organized by The Institute of Botany, Chinese Academy of Sciences (CAS), The Sino-Ecologists Club Overseas, Asian Ecology Section of ESA, and the U.S. Geological Survey. Please visit ‹http://bisobi.sino-eco.org› or e-mail ‹smiao@sfwmd.gov› for details.

The Symposium banner (shown here) uses two typical invasive species, one from Eastern Asia and one from North America, to represent the challenges of biological invasions; the Great Wall symbolizes the efforts to prevent and mitigate the threats to environment and economy from such species exchanges.
Shown on the left of the photograph is kudzu (Pueraria montana var. lobata), first introduced to the United States in 1876 at the Centennial Exposition in Philadelphia by a Japanese government sponsored garden, and now covering more than 7 million acres in the southeastern United States. On the right is goldenrod (Solidago canadensis), native to the United States and Canada, but exotic in China. The disproportionate scale of kudzu and density of Solidago canadensis signifies the seriousness of their threats.

American Society of Mammalogists 84th Annual Meeting

The 84th Annual Meeting of the Ameican Society of Mammalogists will be held 11–16 June 2004 at Humboldt State University, Arcata, California. In addition to contributed oral and poster presentations covering all aspects of mammalian biology, this year’s program will feature two symposia. Dr. Penny S. Reynolds will convene “Problems of Sampling and Statistical Assessment in Mammalogy Research.” “Habitat Loss and Fragmentation: Impacts of Urban Sprawl on Mammals” will be convened by Dr. John A. Yunger. Special addresses will be offered by the recipients of the Joseph Grinnell (Dr. David Schmidley) and C. Hart Merriam (Dr. Terry Bowyer) awards, as well as by student honorees. Dr. James Estes, best known for his work on the behavioral ecology of marine mammals (especially sea otters) and on the community ecology of kelp forests, will present the capstone presentation. Also included are the usual ASM socials, ideal for professional interaction.

Nonmembers who are interested in attending the meetings and/or presenting papers should request materials from the Chairman of the Local Program Committee, Dr. Brian Arbogast, Department of Biological Sciences, Humboldt State University, Arcata, CA 95521; (707) 826-4180; Fax: 826-3201; E-mail: bsa2.humboldt.edu. For additional information, please visit the meeting web site at ‹http://www.humboldt.edu/~asm/› For more information about the ASM, please visit our web site at ‹http://www.mammalsociety.org›

Wild Trout VIII Symposium: Working Together to Ensure the Future of Wild Trout: Call for Papers and Posters

The meeting will be held 20–22 September 2004 at the Old Faithful Inn in Yellowstone National Park. The goals of the Symposium are to bring together a broad and diverse audience representing governmental entities, nongovernmental conservation organizations, media representatives, educators, anglers, guides, and business interests associated with trout fisheries to share their viewpoints on wild trout management and related public policy, to exchange technical information, and to seek ways of cooperatively advancing the conservation of wild trout resources.

Authors are encouraged to conslt the Wild Trout VIII web page ‹www.wildtrout8.com› for guidelines for abstract, poster, and manuscript preparation. Abstracts must be submitted by 1 February 2004. For additional information, contact Program Chair Robert Carline at ‹rcarline@psu.edu›.

Carnivore Conservation Conference

Defenders of Wildlife is pleased to announce Carnivores 2004: Expanding Partnerships in Carnivore Conservation. The conference will be held 14–17 November 2004 in Santa Fe, New Mexico. This will be our fifth biennial carnivore conservation conference. Defenders is accepting oral (15-minute) and poster presentation submissions for Carnivores 2004. Abstract submissions must be received by 1 June 2004.
While the conference focus is on partnerships in conservation, we will consider any papers covering topics such as biology, behavior, taxonomy, general ecology, recovery, management, and economic impacts of terrestrial, avian, and marine carnivore species (canids, bears, cats, mesocarnivores, raptors, whales, dolphins, sharks, etc). Please visit our official conference website, ‹www.carnivoreconference.org› or leave a message at (202) 789-2844 x315 for complete Carnivores 2004 and abstract submission details.

Aimee Delach
Senior Species Program Associate
Defenders of Wildlife
E-mail: adelach@defenders.org

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Meeting Reviews

Advances in Coastal Habitat Restoration in the Northern Gulf States

“Advances in Coastal Habitat Restoration in the Northern Gulf States,” a symposium cosponsored by the Coastal Restoration and Enhancement through Science and Technology (CREST) Program, with the National Oceanic and Atmospheric Administration (NOAA) and the U.S. Geological Survey (USGS), was held in Thibodaux, Louisiana (USA) on 1-2 July 2003.

In order to integrate available scientific knowledge in support of the restoration of coastal habitats in the northern Gulf of Mexico, the symposium extended the definition of coastal habitat to include adjacent freshwater wetlands, along with estuaries and nearshore marine areas. This symposium is the first major contribution from the new CREST program, which coordinates support from NOAA to university-based research into the restoration ecology of coastal ecosystems in the northern Gulf of Mexico.

Fifteen invited speakers created the atmosphere of a particularly well-attended national meeting of coastal ecologists. These presentations drew on experience from at least nine regional wetland and coastal restoration programs accrued over the past decade. Contributed posters and panel discussions rounded out the program. The largely local audience was composed of state and federal resource managers and academic scientists—those who will shoulder the burden of implementing restoration in the northern Gulf of Mexico. The meeting was timely for its immediate audience, but it also provided an impetus and opportunity to review progress achieved in coastal restoration elsewhere.

The CREST symposium captured a snapshot of coastal restoration in the United States and the ecosystem science behind it. The low-lying, sedimentary coastlines found through much of the United States are defined by a constantly changing geomorphology; it would be quixotic to attempt to reconstruct and maintain coastal ecosystems in a particular former state. Instead, coastal restoration means reestablishing natural patterns in the processes that shape and sustain these ecosystems. In particular, coastal restoration attempts to mitigate the negative impacts of human activities and thus recover natural resources and ecological services that have been lost.

Ecosystem science principally contributes to restoration in three areas: by building an understanding of the mechanics of coastal ecosystems, by identifying the effects of human activities, and by predicting how these ecosystems will respond to attempts to restore them. The symposium organized presentations on these topics around restoration goals related to fish and wildlife (fauna), water quality, and wetlands vegetation (habitat). In addition, several presentations addressed the challenges of integrating science with management of coastal ecosystems. Success here depends on negotiating with managers and stakeholders a common understanding of the goals for restoration and a corresponding set of ecosystem attributes in which changes can be predicted and measured.

The goal of maintaining fisheries and wildlife resources serves as one of the main motivations for public investment in coastal restoration. This presents managers and scientists with the challenging task of establishing a clear link between characteristics of habitat and the supported populations of fish and wildlife. Restoration in northern San Francisco Bay depends on maintaining freshwater discharge in the Sacramento River, which means that this freshwater cannot be provided to dry areas outside the basin. Wim Kimmerer (San Francisco State University) described the principal mechanisms linking higher freshwater discharge to increased fish production in the Sacramento River delta. Counter to the conventional wisdom, this link most likely consists of the direct response of fish to physical conditions related to the amount of freshwater in the estuary, rather than from any general enhanced productivity throughout the food web.

Thomas Minello (NOAA Fisheries) described similar work aimed at documenting the effect of wetland restoration and creation on adult populations of crustacea in Galveston Bay, which rely on these habitats for nursery areas. Brown and white shrimp, as well as blue crabs, all show enhanced populations at the edges of wetland areas, and this has a direct implication for the design of wetland restoration projects. Charles Peterson (University of North Carolina) argued persuasively that restoring oyster reefs provides measurable benefits as habitat for other species beyond the economic value of a sustained oyster fishery. The habitat value of oyster reefs can be measured by increased production of other estuarine fisheries. Restoring oyster reefs also contributes to maintaining water quality, especially clarity, but these benefits are as yet difficult to evaluate.

Eutrophication and contaminants pose chronic threats in all coastal systems. Coastal restoration in the northern Gulf of Mexico involves redistributing flow out of the main channel of the Mississippi River. What risks does this pose for the estuarine areas that will receive augmented flow and to the nearshore marine areas normally affected by the river’s plume? Based on results of monitoring studies presented by Charles Demas (USGS), water quality in the Mississippi River falls within existing quality standards for contaminants. However, these standards generally do not control for the risk of chronic stress, e.g., from long-lived herbicides acting on wetland vegetation. Several other speakers addressed the risk of eutrophication resulting from increased nutrient loading.

Increased nutrient loading can benefit restoration efforts if it stimulates vegetative growth without the adverse effects of eutrophication in the water column. In coastal Louisiana, increasing uptake of nitrogen in wetlands may decrease the intensity of hypoxic conditions in shelf waters fed by nutrients in the plume of the Mississippi River. Some scientists believe that stimulating increased production of organic matter is the key to converting increased sediment delivery from diversions into net emergent marsh area. John Day (Louisiana State University) reported on studies into the fate of nutrients introduced in Mississippi River water diverted into estuarine basins that previously received little or no input of riverine water and sediment. Dubravko Justic (Louisiana State University) described the role of climatic factors on magnitude and extent of the hypoxic zone of the Mississippi River, and the possible confounding effects of long-term climate change on restoration efforts.

The role of hydrologic variability emerged as a minor theme in discussions of threats to water quality and restoration strategies. Hans Paerl (University of North Carolina) delivered a nuanced discussion of the role of climate-driven variability in estuarine response to nutrient loading and management control measures, based on experience in the Nuese River and Pamlico Sound. Runoff is the primary factor determining the annual load of nutrients in this system, but runoff also influences residence time, which in turn modulates estuarine response to nutrient loading. Although the influences of climate-driven pulsing on coastal ecosystems are readily understood, it appears that such pulsing will make it difficult to predict future trends in estuarine water quality.

Some encouraging results have been achieved in restoration efforts in the United States, and the symposium provided an opportunity to decipher strategic elements contributing to these successes. Chief among these is knowing how to measure success. Chris Craft (Indiana University) provided information from salt marsh creation projects implemented in North Carolina over the past 10 years. Craft and others advocated tracking success by using a variety of indices descriptive of vegetation and soil properties, with target values keyed to representative “reference systems.” Beth Middleton (USGS) argued, based on life history requirements of freshwater wetland vegetation species, that recreation of wetland systems dominated by riverine influence requires attention to the natural variability in hydrologic forcing. Other speakers offered perspectives based on experience in large-scale restoration in the Chesapeake Bay, the Columbia River, and the Everglades of South Florida.

The underlying causes of wetland loss in the northern Gulf differ from those of other coastal systems in the United States, and consequently so will the mechanisms for reversing this loss. Missing from the symposium was any information on the dynamic geomorphologic processes active along this deltaic coast. Knowledge of these processes is keenly needed in planning for coastal restoration. Beginning in the late 1800s, major construction of various structures for flood control and navigation has interrupted the natural geomorphic processes, although construction of levees began with the settlement of New Orleans more than a century earlier. Similar stories can be told for other large river deltas. To a large degree, restoration of the ecosystems of a deltaic coast will depend on successfully recreating, through river engineering, these lost geomorphic processes. This will require gathering information about comparable coastal systems, i.e., large river deltas, worldwide. Perhaps this will be pursued in a future CREST symposium.

Overall, the CREST program’s inaugural symposium succeeded in assembling an integrated view of current ecological science relevant to coastal restoration, albeit limited to restoration efforts in the United States. A set of papers based on the invited presentations, to be published in a special issue of Ecological Engineering, should be of interest to anyone working on restoration of coastal ecosystems. More information on coastal restoration in the northern Gulf of Mexico can be found at the CREST web site, ‹http://www.gulfcrest.org/›

William Nuttle
11 Craig Street
Ottawa, ON, Canada K1S 4B6
E-mail: wnuttle@eco-hydrology.com

Piers Chapman --Director, CREST Office
1143 Energy, Coast and Environment Bldg
Louisiana State University
Baton Rouge, LA 70803
E-mail: pchapman@lsu.edu

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Contributions

Commentary

Two Bodies Cannot Occupy the Same Place at the Same Time, or The Importance of Space in the Ecological Niche

Studies of resource partitioning and niche had their golden era in the 1970s to the 1980s, but fell in disgrace after being convicted of circular reasoning and not considering alternative hypotheses. Nevertheless, niche theory persists and has a central role in contemporary ecological theory (Cheesson 1991, Leibold 1995).

The initial definitions of ecological niche stressed the idea of niche as an organism’s position or place in the community, or in the biotic environment (Grinnell 1917 and Elton 1927, respectively). Hutchinson in 1957 described niche as a hypervolume in the n-dimensional ecological space, also distinguishing the fundamental and realized niche. After extensive literature surveys on community ecology, Schoener (1974, 1983) ranked the five most important factors of resource partitioning as macrohabitat, microhabitat, food type, time of day, and seasonality of activity. Pianka (1999) summarized niche dimensions in a subset of the three most important—food type, space and time—once more suggesting the importance of space and time in the organism’s ecological niche.

The importance of the spatial dimension of niche is stressed in these definitions, but it was not followed by recognition of its a priori importance over food type and time. Space is a limited resource by definition. Theoretically, two bodies cannot occupy the same point in space and time. In practice, species such as parasites and symbionts differing greatly in body size could be in almost the same place at the same time, where the body of one species is the immediate environment of another. However, for organisms of similar size the theory holds: they may overlap in habitat or microhabitat, but it is not likely that they would overlap at the same point of the space–time bi-dimensional axis. Through deduction, partitioning of space–time axes is expected to be intense. When in close proximity, one will eventually be expelled, be it an organism, a social group, or a colony. In close proximity, diet segregation will not avoid exclusion of one of the organisms from the site. Segregation in time could avoid this spatial exclusion, but it is only possible if food resources are not limiting, or have high renewal rates. Therefore, complete overlap in microhabitat and time cannot be counteracted by segregation in food type. On the other hand, differentiation (not necessarily segregation) in space use might counteract complete overlap in diet.

Since MacArthur (1958) and Connell (1961), the use of space is an important niche dimension for understanding species coexistence and competition. Space use seems to be the main factor explaining community structure and species richness at the local (examples below) and continental scales (e.g., Kelt et al. 1996), along elevational gradients (Patterson et al. 1998), and along the evolutionary process of vertebrate radiation (Streelman and Danley 2003). Differentiation in space use is a key factor in community structure (see Cunha and Vieira [2002] for small mammals; Peres [1997] for primates; Arlettaz [1999] for bats; Howard and Hailey [1999] for reptiles, Brodman and Jaskula [2002] for amphibians; MacArthur [1958] for birds, Krijger and Sevenster [2003] for insects, Bouchon-Navaro [1986] for reef fishes, and Kohda and Yanagisawa [1992] for lake cichlids). In most cases, space use patterns clearly distinguish species with extensive overlap in diet (see references above and the respective citations). Segregation in space use may be more important in three-dimensional or complex environments (sensu August 1983), such as tropical forests, coral reefs, and other aquatic communities.

With space as the most important niche dimension, analysis of space use patterns becomes a central requisite to study species coexistence, especially in complex environments. In addition, use and availability of space are frequently easier to measure than other dimensions of the niche, particularly use and availability of food resources. Other dimensions usually considered important, such as food resources and time use, should always be considered as complementary to space use.

Literature Cited

Arlettaz, R. 1999. Habitat selection as a major resource partitioning mechanism between the two sympatric sibling bat species Myotis myotis and Myotis blythii. Journal of Animal Ecology 68:460–471.
August, P. V. 1983. Heterogeneity in structuring tropical mammal communities. Ecology 64:1495–1507.
Bouchon-Navaro, Y. 1986. Partitioning of food and space resources by chaetodontid fishes on coral reefs. Journal of Experimental Marine Biology and Ecology 103:21–40.
Brodman, R., and J. Jaskula. 2002. Activity and microhabitat use during interactions among five species of pond-breeding salamander larvae. Herpetologica 58:346–354.
Chesson, P. 1991. A need for niches? Trends in Ecology and Evolution 6:26–28.
Cunha, A. C., and M. V. Vieira. 2002. Support diameter, incline, and vertical movements of four didelphid marsupials in the Atlantic forest of Brazil. Journal of Zoology 258:419–426.
Elton, C. 1927. Animal ecology. Sidgwick and Jackson, London, UK.
Howard, K. E., and A. Hailey. 1999. Microhabitat separation among diurnal saxicolous lizards in Zimbabwe. Journal of Tropical Ecology 15:367–378.
Hutchinson, G. E. 1957. Concluding remarks. Population Studies: Animal Ecology and Demography. Cold Spring Harbor Symposia on Quantitative Biology 22:415–427.
Grinnell, J. 1917. The niche-relationships of the California Thrasher. Auk 34:427–433.
Kelt, D. A., J. H. Brown, E. D. Heske, P. A. Marquet, S. R Morton, J. R. W. Reid, K. A Rogovin, and G. Shenbrot. 1996. Ecology 77:746–761.
Kohda, M., and Y. Yanagisawa. 1992. Vertical distribution of two herbivorous cichlid fishes of the genus Tropheus in Lake Tanganyika, Africa. Ecology of Freshwater Fishes 1:99–103.
Krijger, C. L., and J. G. Sevenster. 2001. Higher species diversity explained by stronger spatial aggregation across six neotropical Drosophila communities. Ecology Letters 4:106–115.
Leibold, M. A. 1995. The niche concept revisited: mechanistic models and community context. Ecology 76:1371–1382.
MacArthur, R. H. 1958. Population ecology of some warblers of Northeastern coniferous forests. Ecology 39:599–619.
Patterson, B. D., D. F. Stoz, S. Solari, J. W. Fitzpatrick, and V. Pacheco. 1998. Contrasting patterns of elevation zonation for birds and mammals in the Andes of southeastern Peru. Journal of Biogeography 25:593–607.
Peres, C. A. 1997. Primate community structure at twenty western Amazonian flooded and unflooded forests. Journal of Tropical Ecology 13:381–405.
Pianka, E. R. 1999. Evolutionary ecology. Sixth edition. Addison-Wesley, Boston, Massachusetts, USA.
Schoener, T. W. 1974. Resource partitioning in ecological communities. Science 185:27–39.
Schoener, T. W. 1983. Field experiments on interspecific competition. American Naturalist 122:240–285.
Streelman, J. T., and P. D. Danley. 2003. The stages of vertebrate evolutionary radiation. Trends in Ecology and Evolution 18:126–131.

André Almeida Cunha and Marcus Vinicius Vieira
Laboratório de Vertebrados
Departamento de Ecologia
Universidade Federal do Rio de Janeiro
CP 68020
Rio de Janeiro, RJ
CEP 21941-590 Brazil
(55) 21-25626314/92362024
Fax: (55) 21-25626313
E-mail: cunha@biologia.ufrj.br

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A History of the Ecological Sciences, Part 12:
Invertebrate Zoology and Parasitology during the 1500s

The terms invertebrate zoology and parasitology were coined later, but since our concern here is the history of ecology, not classification, these terms are convenient indicators of subjects discussed. A number of physicians advanced the knowledge of these subjects; the most prominent are discussed here.

Girolamo Fracastoro (1478–1553) was from a patrician family of Verona, and he studied medicine at the University of Padua, which had one of the best medical schools in Europe. He was a true Renaissance humanist who had broad interests and many friends in high places (Thorndike 1941:488–493, Zanobio 1972). His most famous work, a 1346-line poem in three books, Syphilis sive morbus Gallicus (1530), was more famous for its literary than its scientific merits; it is the most famous Renaissance poem in Latin (Eatough 1984:1). Perhaps he chose a verse format because his own ideas were uncertain, and he wanted to convey them ambiguously. Our name for this disease comes from the title of his poem.

Fig. 1. Girolamo Fracastoro [Wright 1930]

He tells us that this disease came to Italy with the invasion of Charles VIII’s French army, hence the name morbus Gallicus (Fracastoro 1984, I:5), but that earlier it had come to Europe with Columbus’ sailors (Fracastoro 1984, I:69–70). He thought it was contagious (Fracastoro 1984, I:130), but did not mention sexual transmission. What was its source? Perhaps the air, corrupted by the planets and modified by climate (Fracastoro 1984, I:182–185). He was more explicit in describing its symptoms (Fracastoro 1984, I:320–367). Book I was devoted to such speculations, and Books II and III were devoted to its cure. He warned that south winds and the filth and sweat of unclean marshes are dangerous (Fracastoro 1984, II:81–86), and he thought that fish was a dangerous food (Fracastoro 1984, II:118–120). A concoction of thyme, hops, fennel, parsley, and other herbs could help (Fracastoro 1984, II:174–180), but if not effective, then try a mixture of styrax, mercuric sulphide, lead, antimony, and grains of incense (Fracastoro 1984, II:260–263). However, another cure comes from the land where the disease originated: lignum-sanctum, or guaiacum (Fracastoro 1984, III). None of his advice seems original (Munger 1949, Eatough 1984:20). Neither remedy was effective, and of course mercury and lead are dangerous. Mercury might cause some sores to shrink, but at the expense of poisoning the body.

After 16 years of further medical practice, reading, and pubishing, Fracastoro’s ideas on contagious disease were more definite, and he published the prose treatise that is the basis of his place in the history of medicine and his relevance for the history of ecology. This book contained two distinct works: the first, on sympathy and antipathy, was quite traditional and vacuous in its speculations (Wright 1930:xxxiv–xxxv, Thorndike 1941:493–496); the second part, on contagion, contagious diseases, and cures, broke new theoretical ground. One historian of medicine calls it “a truly marvellous triumph of close observation and clear reasoning,” and the most important statement of contagion theory before Pasteur (Winslow 1943:143). A later evaluation of Fracastoro’s concepts emphasizes their speculative nature (Howard-Jones 1977). Both evaluations seem valid; he studied the problem for many years, but could only reason about it before development of the experimental method.

He identified three means of contagion: direct contact with a sick person, contact with objects contaminated by a sick person, and contagion at a distance (Fracastoro 1930: 7). Although reasoning by analogy often led astray early naturalists, including Fracastoro, his reasoning about contagion through contact was perceptive; he compared it to spoiled grapes or apples spoiling others adjacent to them. Like Lucretius in ancient Rome, he believed invisible germs could travel through air, though neither of them argued that such germs were alive. Fracastoro’s concept of germs was chemical (Winslow 1943:133).

Although unable to penetrate the fundamental mystery of contagious diseases, it was a valuable step to survey the different kinds. He realized that some diseases mainly attack children, and that some of them cause fevers and pustules on the skin (Fracastoro 1930:73). He interpreted these “variolae” as diseases of blood, and the pustules as putrefactions escaping the body. Once that happens, the person seldom suffers the same disease again. Rabies was a disease primarily of dogs, and it was known since antiquity that they transmit it by their bites. Then it incubates for 20 or 30 days, but sometimes for 4–6 months (Fracastoro 1930:125). The symptoms he describes were well known, but his further speculations on hydrophobia beyond that were fruitless. His ideas on syphilis were now more definite than when he wrote his poem on it. He again tells us it came to Naples from France with King Charles’ army in the 1490s, and that it had come from the New World with Spanish seamen. But now he also tells us that it is transmitted mainly by sexual intercourse, though infants could get it from an infected mother or wet nurse by suckling their milk (Fracastoro 1930:135–137). Syphilis can cause pustules, and he wanted to separate it from other maladies such as elephantiasis, leprosy, and scabies, but that was too difficult a challenge, since one might evolve into another: “Psora, which is nowadays called scabies, is a still milder affection than leprosy, but in a severer form it may pass into leprosy, just as leprosy in a severe form may pass into elephantia” (Fracastoro 1930:171). To go further would require the laboratory techniques of the 1800s.

Girolamo Gabuccini was an Italian physician who published the first separate treatise on parasitic worms (Gabuccini 1547), which attracted enough attention to appear in a second, possibly pirated edition at Venice in 1547, and a third edition at Lyon in 1549. Parts of his account of tapeworms (which were known since antiquity) are quoted in Latin by Hoppeli (1959:104, 152, note 9); Gabuccini believed that the lesser heat in the intestine leads to tapeworm formation (Hoppeli 1959:134). He also described the liver fluke of sheep and goats (quoted in Latin by Reinhard 1957:209), first identified by a French sheep farmer, Jean de Brie, who, at the encouragement of Charles V, wrote a treatise on managing sheep in 1379. De Brie believed that sheep got flukes (Fasciola hepatica) by eating a herb, la dauve, in marshy places. He was close to the truth, since larval stages encyst on vegetation in marshy places to await sheep, but de Brie thought the herb’s leaves actually turned into flukes. Anthony (or his brother John) Fitzherbert went further in his Boke of Husbandrye (1523), describing two dangerous plants in marshy places, but also describing the fluke itself (de Brie’s and Fitzherbert’s accounts are quoted by Kean, Mott, and Russell [1978:561–562], [Fitzherbert 1882:50–51]). Gabuccini came no closer to solving the mystery of the liver fluke’s life cycle than his two predecessors, and it would be three centuries before breakthrough discoveries were made (Reinhard 1957:216–220). His book is digested in a long chapter by Thomas Mouffet entitled “Of the Signs and Cure of Worms out of Gabucinus” (1967:1111–1122). It is so heavily documented with references to ancient and medieval authors that it is difficult to detect any innovations beyond the synthesis itself.

Not only did physicians in the 1500s write medical books, but some also wrote extensively on plants and animals (Egerton 2003a, b). Edward Wotton (1492–1555) received his undergraduate education at Oxford and his medical doctorate at Padua, then returned to practice in London, where he served a term as president of the Royal College of Physicians (Wheeler 1976). His De differentiis animalium libri decem (Paris, 1552) was a survey of the animal kingdom, compiled largely from classical sources. His original contributions were inserted in Book 9 on insects, where he struggled to reconcile his observations of sexual reproduction with traditional belief in spontaneous generation. The weight of tradition was too strong for him, and he concluded that both methods of reproduction occur (Raven 1947:42).

During the second half of the 1500s there was a race to publish the first book entirely on insects, although the participants probably did not know of each other’s work. The Italian Ulisse Aldrovandi became known in Europe first for his three volumes on birds (1599–1603); the Englishman Thomas Penny had devoted much of his research to plants. Aldrovandi and his assistants easily won the race with publication of De animalibus insectis Libri VII (Bologna, 1602; later editions in 1618, 1623, 1638), because Penny died before he had progressed beyond collecting his information.

Aldrovandi, as he explained in a letter to Cardinal Maffeo Barberini (later Pope Urban VIII, who condemned Galileo), collected his insects on little expeditions (translated in Ley 1968:158):

   I was in the habit of going into the country for months during the summer and autumn, not for relaxation, like others; for at these times I employed all my influence, as well as money, to induce the country-people to bring me such insects, whether winged or creeping, as they could procure, in the fields or under ground, and in the rivers and ponds. When any were brought to me, I made inquiries about its name, habit, locality, &c. I often, too, wandered over the marshes and mountains, accompanied by my draughtsman and amanuenses, he carrying his pencil, and they their notebooks. The former took a drawing if expedient, the latter noted down to my dictation what occurred to me, and in this way we collected a vast variety of specimens.

Since Gessner had not published on insects, Aldrovandi was much more on his own for De Animalibus insectis than in other volumes that depended heavily on Gessner’s work. Some of the original pages of illustrations made by his draughtsman survive, and two of them are published, along with extensive translations of the Latin text into German, and modern identifications by Bodenheimer (1928–1929, I:250–276, II:336–345), including the first dichotomic key ever written, to determine the higher groups. Aldrovandi’s three volumes on birds contained 1009 pages of text; he devoted 300 pages to insects. Section I, of 95 pages, describes bees, wasps, and hornets; Section II, “De papilionibus,” includes 81 species of Lepidoptera and also dragonflies; Section III describes two-winged insects, including flies and mosquitoes; Section IV, those with more than two wings: grasshoppers, crickets, beetles, praying mantis; Section V, insects without wings: true insects—bugs and ants—but also spiders, scorpions, centipedes, and millipedes; Section VI, “De vermibus,” includes earthworms and shell-less snails; Section VII is on water insects, but also includes marine worms and starfish (Ley 1968:158–159, Beier 1973:85–86).

Fig. 2. Aldrovandi 1602. Reproduced from and identifications by Bodenheimer 1928–1929, I:268.

Thomas Penny (c.1530–1588/1589) followed in the footsteps of William Turner, and was a friend of Turner’s son, Peter (Raven 1947:153). Like Turner, he studied theology and medicine at Cambridge, and in 1565 he traveled to Zurich to study medicine with Conrad Gessner. That lasted only a few months before Gessner died suddenly of plague. Before going there, Penny had already begun studying both plants and insects, and after arriving he and Gessner shared their knowledge of these subjects with each other. Gessner had planned a volume on insects (Théodoridès 1966), but his only “insect” account ever published was on scorpions (Topsell 1967:750–757). After Gessner’s death, Penny assisted in preparing some of the manuscripts for publication, and he acquired some of the manuscripts on insects (Raven 1947:157). Penny then traveled and studied in Europe, 1566–1569, before returning with his M.D. degree to England, where he practiced medicine in London. When he returned, his botanical interest was dominant, but he spent the last 15 years of his life preparing to write a book on insects, using Wooton’s, Gessner’s, and his own materials. However, he never wrote the book, and gave his materials to his young friend and colleague, Thomas Mouffet (1553–1604).

Mouffet had also studied at Cambridge, and had then received his M.D. at Basel before practicing in London (Simpkins 1974). He reported that Penny’s materials were ill-arranged and ill-written (Raven 1947:172, Mouffet 1967, Preface). As he organized them he added his own comments, which were less useful than Penny’s (Raven 1947:180, 189). Mouffet was distracted by his busy medical practice and never published his Latin manuscript. His widow sold it to Theodore Mayerne, who finally did publish it at London in 1634. It has about 500 wood engravings of varying quality, some of which are quite good (Beier 1973:86). An English translation was published in 1658 as Volume 3 to Edward Topsell’s History of Four-Footed Beasts and Serpents (Mouffet 1967). There was a strong emphasis on insect pests and ways to combat them (Ordish 1976:54–64). Although Raven, an Englishman, found much to criticize in Mouffet’s book, he nevertheless concluded that it was superior to Aldrovandi’s (1947:191). It seems doubtful, however, that Italian scholars would agree; Aldrovandi was quite interested in naming the different kinds of insect, but Mouffet, when faced with a variety of different but similar kinds (such as butterflies), was content merely to number them, though he did provide illustrations and descriptions of each, which are often adequate for modern identifications (Bodenheimer 1928–1929, II:345–352). Although not denying the possibility of spontaneous generation, he expressed skepticism about it because no one claimed to have seen it happen (Mouffet 1967:984). On the other hand, he was not always skeptical of folklore, such as the report of India having locusts three feet long, whose hind legs were used as saws (Mouffet 1967:985).

In at least one instance Mouffet’s (or Penny’s?) medical and entomological interests converged to produce original observations—on the itch mite, Sarcoptes scabiei (1967:1094–1096), which had been known since medieval times. We have a summary of his discoveries (Friedman 1934:627).

Fig. 3. Mouffet 1934. Reproduced from and identifications by Bodenheimer 1928–1929, I:280

(1) Moffet’s description of the acarus, as well as its itch-provoking and burrow-forming activities, was not only the first account of the itch-mite published by an English author but was also the best and most accurate that had anywhere been given up to his time. Moffet, of course, was also familiar with the method of removing the parasite by means of a needle. (2) He was the first to differentiate the acarus from the pediculi. (3) His observation that the acarus was to be found in the “mines” or “furrowes” hard by the vesicle of scabies and not in the vesicle, antedated Renucci’s epochal statement to the same effect by exactly two hundred years.

This last comment on anticipating Renucci points to the disadvantage of the vast encyclopedic works of the 1500s: pearls of wisdom were easily lost in compendiums of ancient and medieval learning.
The honey bee had been domesticated in antiquity and was common in Europe and elsewhere (Chauvin 1968). Mouffet’s longest discussion is on bees (1967:889–921), but their social structure was not understood. Among other things, the queen was considered a king. Although Mouffet made the connection between caterpillars and butterflies and moths, he nevertheless discussed them in different chapters for the good reason that he seldom could connect particular adults with particular caterpillars (1967:957–974, 1029–1041). The same was true about the connection between grubs in the ground and beetles, which he also discussed in different chapters (1967:1005–1016, 1042–1044), though he knew that the whurlworm “The next year after they are bred, they are alwaies transformed into May Beetles” (1967:1043).

Fig. 4. Mouffet 1967:971: “The middle sort of Day-Butterflies” 1 and 2.

Invertebrate animals and diseases were more difficult subjects to study during the 1500s than either vertebrate animals or vascular plants, because there were so many invertebrates, and they, and the causal organisms of parasitic diseases, were smaller beings. Although eyeglasses were invented about 1285 (James and Thorpe 1994), stronger magnifying lenses and microscopes were not developed until the 1600s, stimulated by development of the telescope in 1608 (Bud and Warner 1998:387). Despite the difficulties, physicians and naturalists made important progress in describing invertebrates and diseases, as well as interactions with other organisms.

Literature Cited

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Bodenheimer, F. S. 1928–1929. Materialien zur Geschichte der Entomologie bis Linné.
Two volumes. W. Junk, Berlin, Germany.
Bud, R., and D. J. Warner, editors. 1998. Instruments of science: an historical encyclopedia. Garland, New York, New York, USA.
Chauvin, R., editor. 1968. Traité de biologie de l’abeille. Volume 5: Histoire, ethnographie et folklore. Masson, Paris, France.
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Egerton, F. N. 2003a. A history of the ecological sciences, Part 10: Botany during the Italian Renaissance and beginnings of the scientific revolution. ESA Bulletin 84:130–137.
Egerton, F. N. 2003b. A history of the ecological sciences, Part 11: Emergence of vertebrate zoology during the 1500s. ESA Bulletin 84:206–212.
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Friedman, R. 1934. Thomas Moffet. The tercentenary of his contribution to scabies. Medical Life 41:620–621.
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Acknowledgments

I thank Anne-Marie Drouin-Hans, Université de Bourgogne, Jean-Marc Drouin, Musée National d’Histoire Naturelle, Paris, Benedicte Bilodeau, Centre Alexandre Koyré, Paris, and Sidney A. Ewing, College of Veterinary Medicine, Oklahoma State University, Stillwater, Oklahoma, for their assistance.

Frank N. Egerton
Department of History
University of Wisconsin-Parkside
Kenosha WI 53141
E-mail: frank.egerton@uwp.edu

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A Mussel’s Midday Migration:
Reflections on the Intertidal Zone

On a summer day two mussels met,
One a traveler the other set
On the rock where he was born,
Feeling quite bored and a tad forlorn.
Edulis told his cousin tales
Of acorn barnacles and humpbacked whales,
And the one who was always held down by his byssus
Suddenly realized just what he misses.
So he told his brother and the one convinced another,
And then there were three who went down to the sea.

But before they’d gone far, they saw a sea star,
The reason they never did roam so far from home.
They called out for help as they raced by some kelp.
A Thais replied, contemplating low tide,
But the snail had grown fat and only wanted to chat.
Said Californianus, please don’t detain us.
We’re in quite a hustle, so said the mussel,
Who harbored the fine wish of avoiding the starfish.

The three tried to go faster, in the face of disaster
But right behind was Pisaster: the Intertidal Master.
All three longed for the life before
When they’d stayed away from their predator,
And just when they thought it would be their last day,
A wave came and swept the Pisaster away.
But their byssus threads held our three heroes on tight
And they came through the onslaught feeling all right.
They returned to their rock and from that day swore
They would stay in their niches for evermore.

Meghan MacLean Weir
Green College, Oxford OX2 6HJ
E-mail: meghan.weir@green.ox.ac.uk

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